NYCTAGINACEAE

AMARANTHUS L

48. NYCTAGINACEAE

Trees, shrubs, or lianas (if lianas, with axillary spines);

young branchlets often ferruginous-pubescent. Leaves opposite or nearly so, petiolate; blades simple, entire;

venation pinnate; stipules lacking. Flowers unisexual (dioecious), apetalous, in terminal, bracteate, corymbose clusters; calyx 5-lobed, petaloid; stamens 6-10; filaments united near the base; anthers 2-celled, dorsifixed, dehisc- ing longitudinally; ovary superior, 1-locular, 1-carpellate;

placentation basal; ovule solitary, anatropous or campylo- tropous; style 1, slender; stigma divided and appearing brushlike. Fruits achenes, enclosed in the persistent, modified calyx (i.e., the anthocarp); seed 1, with endo- sperm.

Members of the family are distinguished by the simple, often subopposite leaves, which usually dry darkened, by the distinctive anthocarpic fruit, and by the unisexual apetalous flowers, which usually have an undulate peri- anth limb and a brushlike or much-divided style.

Pollination systems are unknown.

Fruits of Neea and Guapira are endozoochorous.

Those of Neea are chiefly bird dispersed but are also taken by monkeys. Oppenheimer (1968) reported that white-faced monkeys usually remove the pulp and discard the seed. Fruits of Guapira are probably dispersed in part by birds, but white-faced monkeys eat them as well.

Pisonia aculeata has sticky fruits that appear suited for

KEY TO THE SPECIES OF NYCTAGINACEAE

Plants lianas with stout axillary spines Pisonia aculeata L.

Plants shrubs or trees, unarmed:

Shrubs usually less than 2 m tall (to 7 m); staminate flowers tubular-urceolate, the stamens in- cluded; pistillate flowers with ovary sessile, the style included; anthocarps reddish to violet- purple at maturity, usually less than 10 mm long, the surface drying smooth Neea amplifolia Donn. Sm.

Trees to 20 m or more tall; staminate flowers campanulate, the stamens ca half-exserted; pistil- late flowers with ovary stipitate, the style exserted; anthocarps dark blue-violet at maturity, mostly more than 10 mm long, drying with prominent longitudinal grooves Guapira standleyanum Woods.

epizoochory by becoming attached to the fur of animals or to the feathers or beaks of birds.

About 30 genera with 300 species; mostly in the Amer- ican subtropics and tropics.

GUAPIRA Aubl.

Guapira standleyanum Woods., Ann. Missouri Bot.

Gard. 48:404. 1961

Dioecious tree, to 20(35) m tall and 70 cm dbh, some- times buttressed, often ribbed near base; outer bark very thin, flaking in minute pieces; inner bark tan; sap moder- ately strong and somewhat foul-smelling; glabrous but with the branchlets and inflorescences densely ferrugi- nous-tomentose and the petioles and midribs puberulent.

Leaves opposite, crowded; petioles 1-3 cm long, variable on same stem; blades variable, ovate to elliptic to obovate, acute to blunt or long-acuminate at apex with tip down- turned, acute to rounded at base, 5.5-20 cm long, 2.5-9 cm wide, often inequilateral, broadly undulate, dry- ing gray-brown. Inflorescences terminal, corymbose- thyrsiform, 4-13 cm long, all exposed parts minutely ferruginous-pubescent; flowering peduncles to 8 cm long;

flowers unisexual, greenish-yellow, 3-7 mm long, densely tomentose; perianth campanulate, flared abruptly above base, the rim undulate, not flared; staminate flowers cam- panulate; stamens 7-10, about half-exserted; filaments united near base. Pistillate flowers with the perianth accrescent and persistent in fruit; ovary stipitate; style exserted, whitish at anthesis; stigma with many divisions.

Anthocarps oblong-ellipsoid, 12-16 mm long, minutely ferruginous-pubescent, becoming dark blue-violet at maturity, longitudinally striate when dry; seed 1, whitish, ribbed, oblong-ellipsoid, to 13 mm long; mesocarp fleshy, purple, to 2 mm thick. Croat 5555, 5704.

Frequent in the forest. Flowers from February to June.

The fruits mature mostly from May to July.

More field work is necessary on this species before the name can be certain. Guapira standleyanum does not differ appreciably from the type of G. costaricana Standl.

from the Nicoya Peninsula of Guanacaste Province in Costa Rica. Nor does it differ greatly from G. itzana Lund, from Guatemala or from the following species of Torrubia, which are also to be considered Guapira: T.

uberrima Standl. (Colombia), T. rusbyana (Heim.) Standl.

(Venezuela), and T. myrtiflora Standl. (Peru). Probably the species is much more widespread, but currently it should be considered to be endemic to Panama.

Differing considerably from Guapira standleyanum, however, are plants going by the name Guapira costari- cana Standl. from premontane wet forest at higher eleva- tions such as El Valle in Code and Loma Prieta in Los Santos (e.g., Lewis et al. 2199). This is possibly a new species.

Known only from Panama, where it is ecologically variable; known from tropical dry forest in Panama, from tropical moist forest in the Canal Zone and Darien, and from probably premontane wet forest in Colon (Santa Rita Ridge).

See Fig. 226.

48.

NYCTAGINACEAE/PISONIA

385

NEE A R. &P.

Neea amplifolia Donn. Sm., Bot. Gaz. (Crawfordsville) 61:386. 1916

N. pittieri Standl.

Dioecious shrub, usually less than 2 m tall (to 7 m);

younger branches densely ferruginous-pubescent, be- coming glabrate in age. Leaves opposite or nearly so;

petioles 0.5-5 cm long, canaliculate, often somewhat reddish; blades ± elliptic, ovate to obovate-elliptic, abruptly to gradually long-acuminate, obtuse to attenuate at base, 7-36 cm long, 3-15 cm wide, entire, glabrous above, glabrate to puberulent below. Inflorescences terminal, obscurely dichasial thyrses, essentially glabrous to densely ferruginous-pubescent, 4-15 cm long, bearing few to many flowers; staminate flowers narrowly tubular- urceolate, 5-10 mm long, about 2.5 mm wide, the peri- anth with 5 short lobes, the limb weakly spreading; sta- mens 8, to 5.5 mm long, included, attached near base of tube; filaments of different lengths, united into a short tube around the sessile pistillode. Pistillate flowers similar to staminate flowers but with a prominent constriction about one-third of the way down the perianth tube, thickened within at point of constriction; style and stam- inodia (usually 9) fitting tightly through the constriction, included, later exposed when the upper third of perianth above the constriction withers and falls; staminodia held just above constriction; ovary narrowly elliptic, sessile, at first loosely enveloped by perianth, by maturity com- pletely filling it; style slender; stigma with few divisions.

Anthocarp elliptic-oblong, ca 1 cm long, at first reddish, becoming violet-purple at maturity, the persistent peri- anth fleshy and sweet; seed solitary, somewhat shorter than fruit. Croat 4213, 5626.

Frequent in the forest. Flowers throughout the year, most commonly from March to September. The fruits are most common from June to December.

Standley (1933) also reported N. psychotrioides Donn.

Sm., which is a Costa Rican species earlier confused with N. laetevirens Standl. of the Atlantic slope of Panama.

N. laetevirens probably does not occur on BCI. Most of the material from BCI assigned the name N. psychotrio- ides is N. amplifolia. A single sterile collection, Shattuck 121, is in doubt; it does not appear to be typical of N.

amplifolia. However, it can be stated with confidence that a second species of Neea does not now occur on the island. Leaves of the species are consistently host to var- ious cryptogamic epiphytes, including mosses and lichens.

Costa Rica and Panama. In Panama, known principally from tropical moist forest in the Canal Zone, all along the Atlantic slope, and in Darien; known also from pre- montane wet forest in Colon, Chiriqui, and Code and from tropical wet forest in Colon.

See Fig. 227.

PISONIA L.

Pisonia aculeata L., Sp. PI. 1026. 1753

Dioecious, climbing shrub or liana, growing into canopy, its climbing aided by stout, recurved, axillary spines on

Fig. 227. Neea amplifolia

smaller stems, nearly glabrous to sparsely villous, espe- cially on younger parts, underside of leaves, and axes of inflorescences; trunk to 12 cm diam, unarmed; branching divaricate. Leaves opposite to subopposite, the pairs often markedly unequal; petioles 0.5-3 cm long; blades variable, mostly obovate to obovate-elliptic, acute to short-acuminate at apex, rounded to acute at base, 2-10 cm long, 1.5-5 cm wide, usually glabrate above except midrib. Inflorescences usually terminal on condensed short shoots; flowers greenish-yellow, numerous, in dense,

± globular, short-pedunculate clusters to 4 cm diam;

pedicels short, bearing few bracts; pedicels and perianth densely and coarsely short-pubescent; staminate flowers campanulate, ca 3 mm long, the limb 5-7 mm diam, the lobes 5, acute; stamens 6-8 (often 7), unequal, widely exserted, to 8 mm long; anthers as broad as or broader than long, longitudinally dehiscent, dorsifixed at base;

pistillode with a slender style and brushlike stigma, usually held to one side and above rim. Pistillate flowers tubular, ca 2.5 mm long; style short-exserted; stigma brushlike; staminodia reduced. Fruiting inflorescences usually much expanded; anthocarps club-shaped and 5-sided, to 1.5 cm long, densely short-pubescent, bearing a longitudinal row of prominent stalked glands on the angles, the glands sticky. Croat 5390, 8313.

Occasional, in the canopy of the forest, sometimes hanging down over the edge of the lake. Plants may lose their leaves just before flowering. Flowering usually occurs from January to April, with the fruits maturing as early as February.

The sticky, stipitate glands on the fruit presumably function in distribution by adhering to the feathers of birds.

Widely distributed in the American tropics. In Pan- ama, known from tropical moist forest in the Canal Zone and Panama, from premontane moist forest in Panama, and from premontane wet forest in Code.

See Fig. 228.