AMARANTHUS L
61. CHRYSOBALANACEAE
Trees or shrubs; sap sometimes colored. Leaves alternate, petiolate; blades simple, entire; venation pinnate; stipules present. Flowers bisexual, ± actinomorphic, in terminal or axillary panicles or racemes; receptacles present; calyx 5-lobed, the lobes imbricate; petals 5 (rarely lacking), imbricate, free, inserted on the margin of the disk at the top of the hypanfhium; stamens 3-15, free, sometimes unilateral; anthers 2-locular, longitudinally dehiscent;
ovary superior, with 1 fertile carpel (and 2 aborted car- pels), 1-locular; placentation basal; ovules 2; style filiform, basally attached at one side of ovary; stigma truncate or slightly lobed. Fruits drupaceous; seed exalbuminous.
Distinguished by having their styles attached basally to the ovary. Though no other clearly defined morphological features distinguish them, all the BCI species are distinc- tive and are not easily confused with members of any other family.
The flowers of the family are open. Pollinators are unknown.
The fruits of Hirtella are dark-colored and fleshy.
They are possibly dispersed chiefly by bats, but are also
KEY TO THE GENERA OF CHRYSOBALANACEAE
Stamens more than 8 mm long; inflorescences narrow racemose panicles or racemes; leaf blades pubescent but never arachnoid-tomentose Hirtella Stamens less than 5 mm long; inflorescences ± pyramidal panicles; leaf blades white arachnoid-
tomentose on lower surface or completely glabrous Licania
6i.
CHRYSOBALANACEAE/HIRTELLA417
taken by monkeys. H. triandra is taken by the white-faced monkey, which eats the fleshy part and spits out the seed (Oppenheimer, 1968). Licania platypus is also probably bat dispersed (R. Foster, pers. comm.). A Licania species was among those whose seeds were found in bat detritus of Artibeus jamaicensis in Mexico.
Seventeen genera and about 420 species; lowland tropics of Western and Eastern hemispheres.
HIRTELLA L.
Hirtella americana L., Sp. PL 34. 1753 Pigeon plum
Tree, to 20 m tall, to 30 cm dbh; young branchlets and petioles densely rufous-tomentulose. Petioles stout, very short; stipules subulate, paired, velutinous, 5-9 mm long, subpersistent; blades oblong to oblong-elliptic, short- acuminate, rounded at base, 7-15 cm long, 2.5-6.5 cm wide, glabrous to sparsely pubescent above, velutinous below especially on veins, the trichomes denser on veins, with a few round glands near base below. Inflorescences narrow terminal panicles 10-20 cm long, densely pubes- cent except on petals and inner flower parts; branches with rounded or oval, gland-tipped bracts, the glands sessile or short-stipitate; pedicels 1-2 mm long; calyx lobes 5, oval, rounded at apex, often tinged with purple;
petals 5, oval, ca 4 mm long, white to reddish, spreading at anthesis; stamens 3, ca 1 cm long, mounted on a whit- ish disk (2 aborted stamens sometimes visible as subulate trichomes); filaments white, straight and divergent at anthesis; style erect, ± equaling ovary, violet-purple near apex, villous near base. Drupes ellipsoid, 1.5-2 cm long, sparsely pubescent to glabrous, black, shiny; pericarp thin, fleshy. Croat 4866, 7751.
Occasional, in the forest, especially the younger forest.
Flowers from February to May. The fruits probably mature in the early to middle rainy season.
Southern Mexico to northern Colombia and Venezuela.
In Panama, known from tropical moist forest in the Canal Zone, Veraguas, Herrera, Panama, and Darien and from tropical dry forest in Code (Penonome).
Hirtella racemosa Lam., Encycl. Meth. Bot. 3:133. 1789 H. americana sensu Aubl. non L.
Small tree or arching shrub, 1-5 m tall, usually less than 10 cm dbh; young branches usually moderately pubes- cent. Petioles 1-3 mm long, densely hispid to glabrous;
stipules paired, linear, 2-5 mm long, persistent; blades elliptic to oblong, bluntly acuminate at apex, obtuse to subcordate at base, 4-12(19) cm long, 1.5-5(7) cm wide, sparsely appressed-pubescent on midrib above and below and sometimes on surface below. Racemes axillary or terminal, 5-15(29) cm long; rachis and pedicels puberu- lent; bracts and bracteoles 1-3 mm long, narrowly trian- gular, appressed-pubescent, usually with 1 or more round, sessile glands near base; pedicels slender, (1.5)4-11 mm long, perpendicular to rachis; calyx lobes 5, ovate, 2-3 mm long, sparsely puberulent; petals 5, obovate to ellip- tic, 3-5 mm long, lavender; stamens 5-7; filaments 1-2 cm long, lavender; ovary densely pilose-tomentose; style to 1.5 cm long. Drupes oblong-obovoid, to 1.5 cm long, ca 6 mm diam, purplish-brown turning black, sparsely pubescent. Croat 9518.
Frequent in the forest and along the shore. Flowers from November to June, principally in the dry season.
The species may flower more than once a year. The fruits probably mature in a few months, mostly in the late dry and early rainy seasons.
Panama through Colombia, Venezuela, and the Gui- anas to Peru, northern Bolivia, and Amazonian Brazil.
In Panama, known from tropical moist forest in the Canal Zone, Colon, Los Santos, Panama, and Darien and from tropical dry forest in Code (Penonome) and Panama (Taboga Island). Reported from premontane moist, pre- montane wet, and premontane rain forests in Costa Rica (Holdridgeetal., 1971).
See Fig. 252.
Hirtella triandra Sw., Prodr. Veg. Ind. Occ. 51. 1788 Camaroncillo, Carapoto, Chicharron, Conejo, Wild pigeon plum
Tree or shrub, to 20 m; trunk to 23 cm dbh; outer bark thin, fissured, weakly flaky; inner bark reddish-brown, granular; wood light brown, hard, heavy; youngest stems pubescent; sap inconspicuous. Petioles to 5 mm long, moderately pubescent; stipules paired, subulate, to 5 mm long, subpersistent; blades elliptic-oblong, acuminate at apex, acute to rounded or subcordate at base, 4-12(14.5) cm long, 1.5-5 cm wide, glabrous above except on mid- rib, sparsely pubescent below, the trichomes denser on veins. Flowers in terminal racemose panicles to 8 cm long; rachis, branches, and pedicels tomentulose; pedicels slender, 2-3 mm long (appearing longer since flowers are often solitary at apices of branches); bracts and bracte- oles narrowly lanceolate, 1-3.5 mm long, pubescent, lack-
KEY TO THE SPECIES OF HIRTELLA
Inflorescences strict racemes (pedicels ± perpendicular to axis); stamens 5-7; flowers lavender ...
H. racemosa Lam.
Inflorescences short-branched panicles; stamens 3; flowers white:
Branches of inflorescences and younger stems very densely golden-brown tomentose; panicles usually more than 10 cm long, many times longer than broad; bracteoles of inflorescence bearing conspicuous glands; drupes black, shiny, sparsely pubescent H. americana L.
Branches of inflorescences and younger stems not very densely golden-brown tomentose; pani- cles less than 8 cm long, often about as broad as long; bracteoles of inflorescence lacking conspicuous glands; drupes brownish-purple, densely golden-brown tomentose H. triandra Sw.
Fig. 252. Hirtella racemosa
ing glands; calyx lobes 5, ovate, 3-4 mm long, rounded at apex, reflexed at anthesis, puberulent on both sides;
petals 5, broadly elliptic, ca 5 mm long, rounded at both ends, white; stamens 3, long-exserted, arising from side of flower; filaments purplish above, 1-2 cm long, fused into a low ring arising from rim of hypanthium (remnants of a 4th and a 5th stamen persisting as sharp lobes on this ring); style ± equaling stamens and opposite them at anthesis, pilose and white below middle, purple above;
ovary pilose-tomentose; stigma small, capitate. Drupes ovoid-oblong to rounded, ca 2.3 cm long, densely short- pubescent; exocarp thin, brownish-purple; mesocarp fleshy, sweet, tasty; seed ± ovate, ca 1.7 cm long, with irregular longitudinal grooves, abruptly narrowed at base. Croat 7171, 10969.
Common in the forest, mostly in the old forest. Flowers sporadically throughout the year, principally from No- vember to May. Individuals flower more than once a year, possibly as many as three times. The fruits mature throughout the year; white-faced monkeys eat the fruits in June and July (Oppenheimer, 1968).
Unusual growths were found on Croat 6010, which consisted of long-stipitate, obovate, greenish-white, fruitlike structures covered with short erect trichomes.
The structures were regularly disposed on the stems and were at first believed to be fruits. Dr. Ghillean Prance (New York Botanical Garden), who made sections of the structures, has confirmed that they are not fruits. They are possibly due to gall insects.
Central Mexico to northern and western South Amer- ica, Bolivia, and Brazil; West Indies. In Panama, known from tropical moist forest all along the Atlantic slope and in Chiriqui, Panama, and Darien and from premontane wet forest in Colon.
See Fig. 253.
LICANIA Aubl.
Licania hypoleuca Benth., Bot. Voy. Sulphur 91, t.
32. 1844 Garapata
Tree, to 15 m tall; trunk ca 20 cm dbh; outer bark unfis- sured, minutely roughened, thin; inner bark reddish- brown, thin, hard; branches enlarged at base; young branchlets and petioles puberulent; sap with faint, pun- gent odor. Petioles 0.5-1 cm long; stipules minute, paired, subpersistent; blades ovate, acuminate at apex, obtuse to rounded at base, 5.5-10 cm long, 2.5-5 cm wide, glabrous above, whitish below with dense, minute, arachnoid tri- chomes. Flowers minute, in terminal or upper-axillary panicles 10-20 cm long, the branches and pedicels densely and minutely puberulent; hypanthium campanu- late or turbinate, to 2 mm long; calyx lobes 5, minute,
6i.
CHRYSOBALANACEAE/LICANIA419
ovate, acute; petals lacking; stamens 3, less than 1 mm long; anthers about half as long as filaments; ovary white- hirsute; style to 2 mm long, adjacent to the 2 stamenless calyx lobes. Fruits pyriform to obovate, to 2.5 cm long and 2 cm diam, at first pink to red, becoming white and soft at maturity, usually with a minute depression at apex;
mesocarp thick, fleshy, sweet, somewhat pithy; seed obovate, ca 1 cm diam. Croat 14648.
Infrequent, in both the younger and older forests.
Probably flowers in the early dry season. The fruits are mature mostly from April to June.
Southern Mexico to Colombia, Venezuela, the Gui- anas, and Amazonian Brazil. In Panama, known from tropical moist forest on BCI and in Darien and from pre- montane wet forest in Panama (Cerro Azul).
Licania platypus (Hemsl.) Fritsch, Ann. K.K. Natur- hist. Hofmus. 4:53.1889
Sangre, Wild pear, Zapote
Tree, 10-30(50) m tall; trunk to 75 cm dbh (sometimes buttressed to 1-2.5 m elsewhere); branches and leaves glabrous; branchlets often reddish; sap red in age. Petioles ca 1 cm long; stipules ovate, ca 2.5 mm long, stiff, adnate to petiole at base, persistent; blades mostly narrowly oblong-elliptic, shortly acuminate at apex, acute to rounded at base, 10-20(30) cm long, 3-6(8) cm wide, lustrous above, pale and ± glaucous below, with small round flattened or sunken glands below, especially near margin. Panicles terminal or upper-axillary, 10-25 (35) cm long, the branches gray-tomentose, flattened at base;
flowers minute, white, fragrant, sessile or very short- pedicellate; hypanthium, sepals, and edges of petals gray- tomentose; hypanthium ± turbinate, ca 1.5-2 mm diam;
sepals 5, triangular, ca 1 mm long and wide, spreading;
petals 5, obovate, 2-3 mm long; stamens 15, glabrous; fila- ments ca 3 mm long, attached separately to disk; style 5-6 mm long. Drupes variable in size, reported to 20 cm long and 14 cm diam, green turning brown; mesocarp granular, yellow, juicy, sweet; seed usually 1, ovate-oblong, flat- tened, to 5 cm or more long. Croat 8695, 11851.
Frequent in the forest, especially the old forest. Flow- ers principally in the dry season, especially from Febru- ary to April. Time of fruit maturation is uncertain, since fruit size is so variable, but fruits believed to be mature were falling in June and late August. Some fruits prob- ably persist on the tree much longer.
At maturity, the mesocarp smells much like fresh pumpkin.
Southern Mexico (both coasts) south to Colombia;
reported also from the valley of the Magdalena (Jimenez S., 1970). In Panama, known from tropical moist forest in the Canal Zone, Chiriqui, Panama, and Darien; Allen (unpublished) reported the species to be very common
KEY TO THE SPECIES OF LICANIA
Stamens 15; petals 5; leaves glabrous; sap red; fruit to 20 cm long; branchlets reddish
. L. platypus (Hemsl.) Fritsch Stamens 3; petals lacking; leaves densely whitish-arachnoid below; sap not colored; fruits to 2.5 cm
long; branchlets not reddish L. hypoleuca Benth.
Fig. 255. Licania platypus