Trees, shrubs, lianas, or herbs; stems and twigs sometimes quadrangular. Leaves opposite, petiolate; blades simple, entire or serrate, sometimes lobed {Clerodendrum), some- times glandular beneath; venation pinnate; stipules lack- ing. Panicles, racemes, spikes, cymes, or heads terminal or axillary; flowers bisexual, actinomorphic or zygomorphic;
calyx 4- or 5-lobed; corolla tubular, 4- or 5-lobed, of- ten showy; stamens 4(5), didynamous or equal; an- thers 2-celled, dehiscing longitudinally; ovary superior, 2-4-carpellate, the locules as many as carpels or twice as many (by false septation); placentation axile; ovules 1 per apparent locule, anatropous; style 1; stigma simple or bilobed. Fruits of nutlets or dupes of several pyrenes;
seeds lacking endosperm.
Verbenaceae are easily confused with the Labiatae (123), but are distinguished by the undivided ovary, the terminal style, and inflorescences that are not verticillate.
Flowers are probably all insect pollinated (Chapman, 1970). Lantana camara is visited by several species of butterfly, as species of Aegiphila probably are (Chapman, 1970). Stachytarpheta in Costa Rica is heavily visited by skippers (Hesperiidae) and by Eulaema, Euglossa, and large anthophorid bees (Heithaus, 1973).
Lantana camara is chiefly bird dispersed (Leek, 1972).
KEY TO THE SPECIES OF VERBENACEAE
Leaves compound (opposite and each trifoliolate) vitex cooperi Standl.
Leaves simple:
Leaves toothed or lobed:
Leaves broadly cordate at base, semiorbicular with 3-7 short-acuminate lobes; inflorescences large terminal panicles Clerodendrum paniculatum L.
Leaves not cordate, ovate, finely serrate; inflorescences not paniculiform:
Flowers in small, long-pedunculate heads; plants common weeds Lantana camara L.
Flowers scattered along long slender spikes; plants rare, probably no longer on the island Stachytarpheta jamaicensis (L.) Vahl Leaves entire:
Flowers with calyx and corolla violet; calyx lobes exceeding corolla; inflorescences racemes;
leaves stiff, asperous below petreaaspera Turcz.
Flowers white, yellow, or greenish; calyx shorter than corolla; inflorescence not racemose;
leaves not stiff and asperous:
Trichomes of plants (stems, inflorescence branches, and calyces) dense, appressed, white, ca 1 mm long; inflorescences capituliform; fruits in virtually contiguous clusters at maturity Aegiphila cephalophora Standl.
Trichomes lacking or less than 1 mm long; inflorescences ± open panicles; fruits not con- tiguous at maturity:
Calyx lobed to one-third its length, in fruit splitting to the base; inflorescence branches and young stems terete to slightly angulate; fruits pointed at apex; leaves usually less than 2.5 times as long as broad, usually more than 7 cm wide, often softly vil- lous below Aegiphila elata Sw.
Calyx entire to minutely lobed, in fruit ± entire; inflorescence branches and young stems usually 4-sulcate; fruits truncate to depressed at apex; leaves usually more than 2.5 times as long as broad, less than 7 cm broad, strigillose, puberulent below
Aegiphila panamensis Moldenke
Most species of Aegiphila are probably bird dispersed and probably also dispersed to some extent by other animals. Petrea aspera and possibly also Stachytarpheta are wind dispersed.
About 100 genera and some 2,600 species; primarily in the subtropics and tropics in the Southern Hemisphere.
AEGIPHILA Jacq.
Flowers of Aegiphila were reported by Moldenke (1934) as being either male-predominant or female-predominant.
I am considering these to be merely short-styled and long-styled forms, respectively.
Aegiphila cephalophora Standl., Publ. Field Columbian Mus., Bot. Ser. 4:156. 1929
Liana, becoming arborescent, to 10 m long; trunk 6-8 cm diam near ground; stems, petioles, and inflorescence branches bearing dense, long, appressed pubescence.
Petioles 0.5-1.1 cm long; blades elliptic to ovate-elliptic, acuminate, obtuse to rounded at base, 10-16 cm long, 5-6.5 cm wide, sparsely appressed-pubescent above but more densely so on midrib, the trichomes below dense, acropetal, appressed only on midrib. Cymes terminal or axillary, in dense capitate heads; inflorescence branches and calyces with dense long trichomes; flowers sessile or subsessile; calyx ca 3.5 mm long, enlarging to ca 5 mm in fruit, lobed in apical third, glabrous inside, the lobes 4, obtuse to rounded, persisting in fruit; corolla white, lobed, 6-12 mm long, the tube 4-7 mm long, the lobes 4, ovate, spreading; stamens 4, in long-styled forms to 1 mm long, in short-styled forms to 10 mm long; fila- ments fused to basal two-thirds of tube; anthers oblong, ca 0.8 mm long; style with the stigma bifid, in long- styled forms the style and stigma each to 8 mm long, in short-styled forms the style and stigma each ca 1.5 mm long. Fruits globose to ellipsoid, to 9 mm diam, in dense,
± globose clusters, green becoming orange at maturity, the surface granular; seeds (2) 3 or 4, irregular, to 5 mm long. Croat 12543,16511.
Common; normally growing high in the canopy, but occasionally flowering near the ground in tree-fall areas.
Flowers from late June to September (sometimes to December). The fruits mature from September to De- cember.
Apparently endemic to the Canal Zone in tropical moist forest.
Aegiphila elata Sw., Prodr. Veg. Ind. Occ. 31. 1788 Bejuco de peine mico, Guairo santo, Guaro
Liana, becoming shrubby, to 6 m high; stems, petioles, inflorescence branches, and calyces moderately short- pubescent. Petioles 1-2 cm long; blades elliptic to ovate- elliptic, acuminate, obtuse to rounded and often some- what inequilateral at base, 10-20 cm long, 5-10 cm wide, short-villous below especially on veins, puberulent on veins above, with glandular punctations near midrib especially near base. Panicles axillary or terminal, often thyrsoid; pedicels to 5 mm long; calyx bell-shaped, ca
122. VERBENACEAE/AEGIPHILA 733 4 mm long, lobed ca one-third its length, in fruit gla- brate, persistent, enlarging to ca 12 mm long, the lobes 4 (5), sometimes irregular, ovate, with a short acumen;
corolla greenish to cream-colored, 8-9 mm long, the lobes 4 (5), ovate, acute to blunt at apex, glabrous outside, puberulent inside on tube and on basal part of filaments;
stamens 4, long-exserted, 10-12 mm long; anthers oblong, ca 1.3 mm long; pollen white, tacky; style ca 2.5 mm long, slender; stigma bifid, ca 2 mm long. Fruits ± globose, to 1 cm diam, mostly enclosed by the weathering calyx, bright orange at maturity, glabrous; seeds 2-4, to 6 mm long. Croat 14639, Foster 1240.
Infrequent, in the forest. Some flowers have been seen in May and old fruits in September.
A few plants showed the calyx to be merely split and essentially bilobed, with one of the lobes bearing two minute teeth.
Mexico to Colombia, Venezuela, and the Guianas;
West Indies. In Panama, known from tropical moist forest in the Canal Zone, Panama, and Darien and from tropical wet forest in Colon (Santa Rita Ridge).
Aegiphila panamensis Moldenke, Trop. Woods 25:14. 1931
Shrub or tree, to 15 m tall, to 15 cm dbh; young stems and inflorescence branches usually 4-sulcate; stems, leaves, and exposed parts of inflorescences glabrate to densely pubescent, the trichomes very short, stiff, erect or appressed. Petioles to 1.2 cm long; blades elliptic to narrowly ovate-elliptic, acuminate, acute to obtuse at base, 8-19 cm long, 3-7 cm wide. Panicles axillary and terminal, to 13 cm long, the cymes open; flowers numer- ous; calyx and pedicel like an inverted cone; pedicels ca 4 mm long; calyx ± entire, ca 2 mm long, enlarging to ca 4 mm; corolla cream, white, or yellowish, puberu- lent outside, 7-9 mm long, tubular below, soon falling, the lobes 4, ca 3 mm long, globular in bud; stamens 4, in long-styled flowers to 1.5 mm long, in short-styled flowers to 10 mm long; filaments coiled in bud, pubes- cent near point of fusion with tube; long-styled forms with the style long-exserted, to 10 mm long, the stigma bifid, ca 3.5 mm long, the short-styled forms with the style inserted, the style and stigma each to ca 2 mm long.
Fruits drupaceous, ± oblong, truncate and ± depressed at apex, to 12 mm long and 9 mm wide, encircled at base by the accrescent calyx, green turning orange at maturity;
seeds 4, white, only slightly shorter than the fruit. Croat 4094, 6749, 6780.
Occasional, as a shrub in clearings and open areas or as a tree in the younger forest. Flowers from July to December (sometimes from April). The fruits mature from October to January.
This species is similar to A. martinicensis Jacq.
Southern Mexico, Costa Rica, and Panama. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, Panama, and Darien, from premontane moist forest in the Canal Zone and Panama, from premontane wet forest in Chiriqui, and from tropi- cal wet forest in Code.
See Fig. 478.
Fig. 478. Aegiphila panamensis
Fig. 479.
Petrea aspera
CLERODENDRUM L.
Clerodendrum paniculatum L., Mant. PL 90. 1767 Scarlet glorybower, Danger flower, Red pagoda flower Herb or shrub, to 3 m tall; stems angulate, glabrous ex- cept for rings of long trichomes at leaf nodes. Petioles to 30 cm long; blades broadly ovate to suborbicular, acuminate and lobed, cordate and palmiveined at base, 9-25 cm long, 12-30 cm wide, with conspicuous glands 3-8 mm apart on margins, the lobes 5 (sometimes 3 or 7), shallow, acuminate, the lower surface punctate, with many small, whitish, peltate scales. Inflorescences ter- minal panicles ca 15 cm high and ca 20 cm wide, with reduced leaves at branching nodes; inflorescence branches rose-red, minutely puberulent; pedicels to 12 mm long;
flowers 5-parted; calyx campanulate, ca 13 mm long, the tube orange, the lobes obtuse, ca 3 mm long, glandular be- fore bud opens, red; corolla red-orange, the tube slender, 1.5-2 cm long, the lobes paler, spreading, ca 6 mm long;
stamens 4, orange-red, exserted to ca 3.5 cm above throat, curved; style exserted ca two-thirds as far as stamens, minutely bilobed at apex, violet-purple. Drupes small, green, ± enclosed in persistent calyx. Croat 7000.
Cultivated in the Laboratory Clearing. Flowers prin- cipally in the rainy season. No fruits have been seen on BCI.
Native to Asia; cultivated in Europe and in the Ameri- can tropics and subtropics. In Panama, known only from tropical moist forest in the Canal Zone.
LANTANA L.
Lantana camara L., Sp. PL 627. 1753
Pasorin, Bandera espanol, Camara, Cinco negritos, Chi- chiquelite
Herb, usually less than 1.5 m tall, sometimes woody, hir- tellous on most parts, sparsely so on upper leaf surfaces.
Petioles 4-20 mm long; blades ovate, acute to acuminate, rounded to obtuse at base, then abruptly attenuate to half the length of petioles, 4—11 cm long, 2-5 cm wide, mi- nutely crenate-serrate, scabrous or scaberulous above.
Flower heads 1.5-2 cm diam; peduncles to 8 cm long;
flowers sessile, bracteate, puberulent outside, the outer- most of each head orange-red to burnt-orange, the inner- most orange, the bracts acute, ca 3 mm long; calyx ca 2.5 mm long, obscurely 4-lobed, ciliate; corolla tubular, ca 10 mm long, 4-lobed, the tube somewhat expanded and bent above the middle, the limb ca 4 mm wide; sta- mens 4, fused to tube at 2 levels about midway down tube, the free part of filaments short; anthers often shed- ding pollen in bud, their thecae distinct; style held some- what below the lowest pair of anthers; stigma attached laterally, becoming fleshy after anthers have lost their pollen, sometimes secreting a nectarlike substance visited by tiny insects; nectary small, but the nectar often filling base of corolla tube. Drupes round, several to many in a tight cluster, grayish-blue, to 5 mm diam; exocarp thin;
mesocarp sweet, tasty; seeds obovate, to 4 mm long.
Croat 6004.
122. VERBENACEAE/PETREA 735 Abundant in weedier areas of clearings. Flowers and fruits sporadically throughout the year.
Hummingbirds (Damophila julie and Amazilia tzacatl) visit the flowers for nectar (Leek, 1972).
The fruits are taken by a variety of birds, including manakins, flycatchers, honeycreepers, and tanagers (Leek, 1972).
Pantropical. In Panama, known from tropical moist forest in the Canal Zone, San Bias, Herrera, and Darien, from premontane moist forest in Veraguas and Los San- tos, and from premontane wet forest in Code.
PETREA L.
Petrea aspera Turcz., Bull. Soc. Imp. Naturalistes Moscou36(3):211. 1863
Viuda, Flor de mayo, Flor de la cruz
Liana, growing into canopy, appearing to be an arch- ing shrub when juvenile; young parts sparsely short- pubescent, glabrate in age; stems sometimes 4-angled.
Petioles to 1.5 cm long; blades elliptic, often broadly so, rounded to acuminate at apex, acute to rounded at base, 7-21 cm long, 3.5-11 cm wide, usually stiff, asperous below. Racemes long, ± pendent, upper-axillary, to ca 30 cm long; pedicels ca 11 cm long; flowers 5-parted;
calyx light violet, the tube membranous, ridged, ca 6 mm long, the lobes 5, thin, membranous, narrowly obo- vate, 2-2.5 cm long, a crownlike projection occurring at summit of tube and at base of lobes, ca 3 mm high, acutely 5-lobed to near base, often drying brown, per- sisting and acting as dispersal mechanism for fruit; corolla violet, salverform, ca 1.5 cm long, lobed to middle, some- what zygomorphic, puberulent on both surfaces, the tube villous inside near apex, 1 lobe with a white spot near its base; stamens 4, fused to basal two-thirds of tube;
filaments villous, ca 2 mm long; anthers held closely together on one side of the tube at its rim; pollen some- what sticky, adhering to anthers after being shed; style bent slightly at apex in the direction of the white-spotted corolla lobe; stigma held just below anthers. Fruits of
1 or 2 tiny nutlets completely enclosed by the persistent calyx; nutlets ovoid, ca 1.5 mm long, glabrous. Croat 9432.
Abundant in the canopy and at the edge of the forest over the lake; one plant grows as an epiphyte from a large ant nest {Croat 9432). Flowering and fruiting throughout the year, often in synchronous waves throughout the forest.
Petrea volubilis L. was reported by Moldenke (1973) in the Flora of Panama for BCI based on Aviles 14 and Shattuck 412. These specimens do not differ from the typical P. aspera, however, and I believe that only a single species of Petrea occurs on the island.
Widespread in tropical America from northern Mexico to southern Brazil; Cuba, West Indies; widely cultivated.
In Panama, known from tropical moist forest in the Canal Zone, San Bias, Veraguas, Los Santos, Panama, and Darien.
See Fig. 479.
736 DICOTYLEDONEAE STACHYTARPHETA Vahl
Stachytarpheta jamaicensis (L.) Vahl, Enum. PL 1:206. 1804
Berbena, Blue porterweed, Gervao, Jamaica, Verbena, Vervain, Rinchao, Verbena azul, Brazilian tea, Cola de millo Herb, to ca 1 m tall, sparingly pubescent to glabrate, sometimes slightly woody basally. Petioles absent or to
1 cm long; blades oblong to oval, obtuse to acute at apex, cuneate at base, decurrent onto petiole, to ca 11 cm long and to ca 4.5 cm wide, scaberulous above, glabrous below but with sparse jointed trichomes on primary veins, the margins serrate and ciliate-scabrous. Spikes terminal, to 20 (50) cm long, terete, 3-4 (5) mm wide; flowers at first erect, later embedded in the thickened rachis; bracts subtending pits in which flowers are immersed, acute, to 5 mm long and ca 1—1.5 mm wide, the margins scari- ous; calyx tubular, ca 5 mm long, shallowly 4-toothed on the side facing away from stem, notched to 1 mm on side next to stem; corolla salverform, shades of blue, violet, and purple, ca 10 mm long, the tube slightly curved, the limb ca 8 mm wide; perfect stamens 2, inserted above middle of corolla tube, included; filaments very short;
thecae divergent; staminodia 2, posterior, minute; style ca 6 mm long, filiform; stigma terminal, persistent on fruit after corolla falls. Fruits oblong-linear, emerging from rachis pits at maturity, dry, splitting into cocci, the cocci 2, long, hard, narrow; seeds 1 per coccus, linear.
Woodworth & Vestal 739.
Rare. Flowering and fruiting throughout the year.
Alabama and Mexico to Ecuador and Brazil; West Indies; introduced into parts of tropical Africa, Asia, Australia, and Oceania. In Panama, known from tropical moist forest in the Canal Zone, Colon, San Bias, Chiri- qui, Los Santos, Panama, and Darien, from premontane wet forest in Chiriqui and Panama, and from tropical wet forest in Bocas del Toro and Colon.
VITEX L.
Vitex cooperi StandL, Trop. Woods 16:26, 29, 32, nomen nudum 1928; Publ. Field Columbian Mus., Bot. Ser. 4:256. 1929
Tree, to 26 m tall; trunk buttressed and fluted, to 75 cm dbh; outer bark thin, light-colored; branchlets acutely
tetragonal to terete or compressed, densely puberulent when young. Leaves trifoliolate, opposite; petioles 1.3-9.5 cm long, finely appressed-puberulent; petiolules 1-8 mm long, the central one longest, flattened, margined;
leaflets unequal with the lateral pair smaller and nar- rower, the central leaflet rounded and acuminate at apex, acute to acuminate at base, 4-22 cm long, 2.5-10.5 cm wide, entire, shiny and glabrous above, finely puberulent on veins beneath when young, becoming glabrate in age.
Inflorescences axillary, solitary, 3.5-14 cm long, 4-6 cm wide, dichotomously branched, finely puberulent throughout, bearing many flowers; peduncles conspicu- ously flattened, 4.5-7.5 cm long; pedicels 1-3 mm long;
bracts ca 1.5 cm long and 4 mm wide; flowers not seen, reportedly blue and lavender. Fruits not seen.
Rare, known only from the vicinity of the Laboratory Clearing; collected sterile once by R. Foster. Elsewhere in Panama, flowers during June and July, with the fruits maturing during July and August.
Guatemala to Panama; sea level to 600 m. In Panama, known from tropical moist forest in the Canal Zone and Darien and from premontane wet forest in Chiriqui (Progreso); reportedly fairly common on the Atlantic watershed around Gatun Lake {Fisher 1), and no doubt more widespread and common than collections indicate.
123. LABIATAE (LAMIACEAE)
Annual or perennial herbs, erect or vinelike, pubescent, usually aromatic; stems usually quadrangular. Leaves opposite, sessile or petiolate; blades simple, serrate; vena- tion pinnate; stipules lacking. Cymes bracteate, capituli- form or verticillate, contracted, axillary or terminal;
flowers bisexual, zygomorphic; calyx tubular or campan- ulate, 5-lobed; corolla bilabiate, 5-lobed; stamens 2, or 4 and didynamous, epipetalous, often forcefully ejected from flower; anthers 2-celled, the thecae often diver- gent, introrse, dehiscing longitudinally; ovary superior, 4-lobed, 2-locular, 2-carpellate; placentation basal; ovules 4, anatropous; style 1, arising from depression between lobes; stigma bilobed. Fruits of 4 nutlets (in Salvia usu- ally only 1 maturing); seeds 1 per nutlet, with little or no endosperm.
KEY TO THE SPECIES OF LABIATAE
Inflorescences capituliform; flowers white; stamens 4:
Calyx tubular, more than 5 mm long; nutlets ca 1 mm long Hyptis capitata Jacq.
Calyx campanulate, less than 4 mm long; nutlets less than 1 mm long Hyptis brevipes Poit.
Inflorescences verticillate; flowers blue, lavender, or violet; stamens 2 or 4:
Herbs small, cultivated, once planted at the laboratory, probably no longer present on the island;
leaves often variegated Coleus blumei Benth.
Herbs small or large, not cultivated:
Flowers pedicellate; stamens 2; fruiting calyx with short blunt teeth, conspicuously covered with large gland-tipped trichomes Salvia occidentalis Sw.
Flowers sessile; stamens 4; fruiting calyx with long slender teeth, lacking gland-tipped tri- chomes but with sessile glands Hyptis mutabilis (A. Rich.) Briq.
Recognized by the opposite leaves, squarish stems, and pleasant aroma. Members of the family are closely related to the Boraginaceae (121) and Verbenaceae (122), but are generally not confused with either of these fami- lies on BCI.
All BCI species have typical gullet flowers, which are scentless and generally bee pollinated (Faegri & van der Pijl, 1966). The flower is two-lipped, the lower lip serv- ing as a landing platform. The genus Hyptis probably has the same pollination syndrome as that of Eriope studied by Harley (1971) in Brazil, since most Hyptis have the same morphology and triggered release of sta- mens. Eriope flowers open early in the day and are visited by tiny bees. The stamens then spring out, the style later elongates past the stamens, and the stamens become deflexed with the lower lip. The flower usually falls before the day is over.
Nutlets are small and usually numerous. Hyptis nut- lets exude mucilage copiously when wetted (H. Baker, pers. comm.). They are probably in part dispersed by small birds of clearings and in part merely spilled and scattered by passing animals or the wind. The burlike pseudoheads of Hyptis brevipes and H. capitata are bal- lasts set loose by passing animals (van der Pijl, 1968).
Ridley (1930) suggested that H. brevipes might be epizoo- chorous by means of its long-pubescent sepaloid points.
About 180-200 genera and 3,200-3,500 species; cos- mopolitan but concentrated in the Mediterranean region.
COLEUS Lour.
Coleus blumei Benth., Lab. Gen. et Sp. 56. 1832 Coleus, Pompolluda, Chontadua, Jacob's coat
Sprawling herb, to 70 cm long; younger stems and peti- oles densely pubescent. Petioles 1-4 cm long; blades ovate, acute at apex, obtuse to truncate at base, 2-15 cm long, 1.5-10 cm wide, sparsely pubescent above and on veins below, resin-dotted below, crenate, sometimes variegated. Inflorescences terminal, branched, 15-30 cm long; flowers in cymes, purplish; pedicels 1-2 mm long; calyx ca 2 mm long, resin-dotted, unevenly lobed and toothed; corolla ca 1 cm long, the tube recurved, the upper lip bilobed, the lower lip entire, enlarged, boat-shaped; stamens 4; style bifid near apex. Nutlets 4, ca 1 mm diam. Shattuck 158.
Formerly cultivated at the laboratory; not seen in recent years, but so commonly cultivated in Panama it is included here. Seasonality uncertain. Possibly flowers and fruits year-round.
Native to the East Indies; cultivated in many places.
In Panama, known from BCI and Code (El Valle), but no doubt cultivated elsewhere.
HYPTIS Jacq.
Hyptis brevipes Poit., Ann. Mus. Natl. Hist. Nat.
7:465. 1806
Herb, to 60 cm tall, pubescent throughout. Petioles ob- solete or to 1 cm long; blades lanceolate-elliptic or ovate,
123.
LABIATAE/HYPTIS737
acute at apex, attenuate at base, to 5 cm long and 2 cm wide, irregularly or doubly serrate, resin-dotted. Inflo- rescences capitulate, ca 1 cm diam; peduncles to 1 cm long; bracts awl-shaped, ciliate, to 6 mm long; flowers sessile, tufted at base; calyx campanulate, to 3.5 mm long, with 5 spinulose teeth ca 2 mm long; corolla bilabiate, 5-lobed, ca 3 mm long, white; stamens 4, weakly exserted;
style bifid at apex. Nutlets 4, oblong-ovoid, ca 0.6 mm long. Shattuck 479.
Collected once by O. Shattuck on Wheeler Trail; pos- sibly no longer occurring on the island. Seasonal behavior uncertain. Elsewhere flowers and fruits to some extent all year, but possibly with most flowering from Novem- ber to January.
Native to Brazil, but now a weed throughout tropical America, Asia, and Polynesia. In Panama, known from tropical moist forest in the Canal Zone and Panama and from tropical wet forest in Colon.
Hyptis capitata Jacq., Coll. 1:102. 1787 Suspiro de monte
Herb, to 2 m tall, sparsely to moderately pubescent throughout. Petioles slender, 5-20 mm long; blades ovate to rhombic, acute at apex, attenuate at base, 5-12 cm long, 2-6 cm wide, irregularly serrate. Flower heads at first hemispheroid, later globular; flowers numerous, many open at a time in each head; calyx scabrid, hispid at base outside, with a pubescent line inside above nut- lets, the lobes 5, slender, at first equaling tube, the tube much longer in fruit; corolla white, ca 4 mm long, slightly exceeding calyx, bilabiate, the middle lobe of upper lip hooded, loosely enclosing the style and 2 longest stamens, folding backward following anthesis; anthers red, closed at anthesis; filaments curling over the edge of upper lip in time; stigma held midway between long and short pairs of stamens, its 2 lobes open at anthesis. Nutlets usually 4, ovoid, ca 1 mm long, smooth. Croat 4406, 8683.
Common in clearings. Flowers mostly in the early dry season, with the fruits maturing late in the dry season.
This species differs considerably from H. mutabilis in its pollination behavior. Those individuals of H.
mutabilis observed shed the pollen in the bud, and the stamens were thrown forward violently after being opened forcibly (while the stigma was still unopened).
In H. capitata only two stamens are held within the upper lip and then only very loosely. Their flowers are probably not forced open and the stamens, when released, do not spring forward violently. Both protandrous and proto- gynous forms were observed in a single population of H. capitata, though the anthers in some plants were shed in bud, with the style remaining unopened until most of the pollen was shed.
Mexico to Colombia, Venezuela, Ecuador, and Peru;
Asia and Polynesia. In Panama, widespread; known from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, Veraguas, Los Santos, Panama, and Darien, from premontane moist forest in Los Santos and Panama, and from premontane wet forest in Chiriqui, Code, and Panama.