34;•«fer
20. CYCLANTHACEAE
Epiphytic vines or terrestrial, acaulescent or short- stemmed, palmlike herbs arising from a rhizome. Leaves alternate or forming a rosette, petiolate, the petioles dilated at base forming a clasping sheath; blades simple, entire to cleft, often plicate; venation palmate; stipules lacking. Flowers unisexual (monoecious), actinomorphic, the sexes interspersed and closely congested on a simple, axillary spadix subtended by several spathes; perianth cupular or lobed or rudimentary in staminate flowers, many-lobed or rudimentary in pistillate flowers; stamens many; filaments connate at base, epitepalous if a perianth is present; anthers 2-celled, dehiscing longitudinally;
staminodia 4 in pistillate flowers, short or flexuously filiform; ovary inferior, 1-locular, 2-4-carpellate; placen- tation parietal; ovules numerous, anatropous; style none or 1, with 1-4 stigmas. Fruits fleshy syncarps of separate or connate berries; seeds numerous, with copious endo- sperm.
The Cyclanthaceae may be confused with the Palmae (19), but are distinguished by having conspicuous stam- inodia, several usually deciduous spathes subtending the simple spadix, and a fleshy syncarpous fruit with numer- ous small seeds.
Most Cyclanthaceae are visited by small weevils (Cole- optera) and other insects. The first day after anthesis the staminodia appear to yield some fragrance, which prob- ably aas as an attractant. Marling (1958) reported that
20. CYCLANTHACEAE/CARLUDOVICA 179
KEY TO THE TAXA OF CYCLANTHACEAE
Plants vines; leaf blades entire Ludovia integrifolia (Woods.) Harl.
Plants not vines; leaf blades cleft to divided:
Leaf blades becoming deeply 2-parted, almost divided, the segments usually entire; fruiting spadix like a series of concentric rings Cyclanthus bipartitus Poit.
Leaf blades not deeply 2-parted, either 2-cleft or 4-parted, the segments dentate to deeply split;
fruiting spadix not a series of concentric rings:
Leaf blades broadly obovate, bifid halfway at most (the segments often irregularly split); peti- oles winged, at most half as long as blade; plants short-stemmed Asplundia alata Harl.
Leaf blades rounded in outline, 4-parted; petioles at least half again as long as blade; plants acaulescent Carludovica
during this stage the stigmas are receptive and the anthers are unopened, but most species I have observed have the stigmas completely covered by a mass of anthers until later. Moreover, I have seen anthers open and Trigona bees collecting pollen at this stage on Carludovica pal- mata. Usually by the following day the staminodia have withered, and Harling reported that the pleasant aroma is replaced by a "sweetish, somewhat sickening" smell but that the weevils are still present on the spadices among the staminodia. He said the weevils are often very destructive but are nevertheless probably the legiti- mate pollinators of most Cyclanthaceae. Trigona bees collecting pollen might also effect pollination, and visits by xylocopid bees were also reported by Harling.
Fruits are mostly bird dispersed, perhaps except for Cyclanthus, but they may also be taken by lizards, mam- mals, and perhaps even ants (van der Pijl, 1968). Fruits of Carludovica palmata are eaten by the bat Micronycteris hirsuta (Phyllostomidae) (Wilson, 1971) and by the tam- arin (Hladik & Hladik, 1969).
According to Harling (1958), a family of 11 genera and 178 species ranging from southern Mexico and Lesser Antilles throughout Central America and the Amazon basin; also in eastern Brazil.
ASPLUNDIA Harl.
Asplundia alata Harl, Acta Horti Berg. 18:223- 224. 1958
Large terrestrial herb, ca 2 m tall; stems to 30 cm high and 12 cm thick. Leaves to 1.8 m long, closely spiraled, clustered at apex; petioles about half the length of blade;
blades broadly obovate, to ca 1 m long and 60 cm wide near apex, bifid, tapering to narrow base, decurrent onto petiole and continuous with broadly vaginate wing of petiole, becoming very much dissected in age, the sur-
faces pleated, the ribs 3, the lateral ribs branching from the midrib near base, the segment ribs prominently raised on upper surface, with furfuraceous scales scattered on lower surface; juvenile leaves similar to adults in shape, at first entire, splitting in maturity. Inflorescences from leaf axils, often several; peduncles to ca 20 cm long;
spathes usually 3 or 4, ± oblong, 6-10 cm long, widely spaced below spadix, the outer one keeled; spadix cylin- drical, blunt at apex, 4.5-6 cm long, white in flower, becoming green in fruit; staminate flowers alternating with short, conical pistillate flowers, densely covering spadix, but soon falling free; perianth lobed; pistillate flowers with tepals thick, longer than style at anthesis, light green; staminodia to ca 6 cm long, caducous; stigmas medial, narrow, elongated, dark green. Seeds ca 1.5 mm long and ca 1 mm wide. Croat 10204, 14088.
Uncommon, on creek banks and in ravines in the vi- cinity of the Laboratory Clearing. Flowers from late dry season to the middle of the rainy season, March to August.
Mature fruits have been seen from June to September.
Known only from Panama in tropical moist forest in the Canal Zone, San Bias, and Chiriqui and from tropical wet forest in Colon.
CARLUDOVICA R. & P.
Carludovica drudei Mast., Gard. Chron. n.s.
8:714. 1877
Large, acaulescent herb, 2-3 m tall. Petioles to 2.5 m long, terete, vaginate only near base; blades to 1.8 m wide, palmately and unequally 4-lobed, the lobes to 75 cm long, 35-60 cm wide at apex, with 9-16 teeth per lobe, the teeth 5-13 cm long, glabrous and shiny above, dull below; juvenile leaves entire, the apex V-shaped, becoming 4-lobed, the lateral lobes more deeply divided.
Peduncles 40-50 cm long; spathes 4, congested immedi-
KEY TO THE SPECIES OF CARLUDOVICA
Lobes of blade toothed less than one-fourth the way to base; blades shiny on upper surface, dull on lower surface; stigmas laterally compressed; tepals clearly exceeding stigmas even in flower C. drudei Mast.
Lobes of blade toothed deeply, often past middle; blades shiny on both surfaces; stigmas ovate to suborbicular; tepals scarcely, if ever, exceeding stigmas C. palmata R. & P.
Fig. 72. Carludovica drudei, inflorescence with staminodia exserted from pistillate flowers
Fig. 74. Carludovica drudei, cross section of spad
ately below spadix; spadix narrowly cylindrical, 11-12 cm long, 1.5 cm thick in flower, to 22 cm long and 4.5 cm thick in fruit; staminate and pistillate flowers alternating spirally on spadix; staminate flowers in clusters of 4, lack- ing perianth, the stamens numerous, closely congested, obscuring all of pistillate flower but the staminodium, falling within a few days after anthesis; pistillate flowers sunken into fleshy axis of spadix; tepals 4, 5-6 mm long in flower, distinctly surpassing length of stigmas (to 8 mm long in fruit); staminodium slender, flattened, very long and showy, white, falling soon after anthesis; stigmas 4, laterally compressed; fruiting spadices rupturing at ma- turity, beginning at apex, to expose bright orange matrix with embedded fruits. Fruits oblong to rounded, ca 10 mm long, 6-8 mm broad; seeds numerous, ± ovoid, ca 2 mm long, flattened. Croat 10838, 12305.
Occasional, in the forest, usually along streams, pos- sibly preferring steep banks. Flowers in June. The fruits mature from July to October.
Distinguished by having the leaf lobes toothed much less than halfway to the base.
Lowland forests in Mexico (Chiapas and the Yucatan Peninsula), Costa Rica, Panama, and possibly Colombia.
In Panama, known from tropical moist forest on both slopes in the Canal Zone and in Chiriqui and Darien.
See Figs. 72, 73, and 74.
Carludovica palmata R. & P., Syst. Veg. 291. 1798 Panama hat palm, Palmita, Jipijapa, Portorrico, Atadero, Rampira, Iraca, Guachiban, Canagria
Large, glabrous, acaulescent herb, to over 3 m tall. Peti- oles to 3.5 m long, round; blades palmately lobed, pleated, usually light green in color, to 1 m long and 1.9 m wide, usually with 4 ± irregular lobes to 80 cm long and 85 cm wide at apex, the lobes irregularly toothed to past middle, with 10-16 teeth per lobe. Peduncles 20-45 cm long;
spathes usually 3(4), congested just below spadix, ca 25 cm long, greenish to white, maroon near apex; spadix cylindrical, white in flower, ca 15 cm long and 2.5 cm thick, to 25 cm long and 6 cm thick in fruit; staminate and pistillate flowers alternating spirally as in C. drudei;
tepals scarcely if at all exceeding stigmas; staminodia 4-6 cm long; stigmas ovate to suborbicular. Fruits orange, ca 1 cm long; seeds many, white, 2 mm long. Croat 10385.
Common locally, primarily in clearings and on steep banks of the shoreline, but also in the forest on Orchid Island; common and widespread in the Canal Zone.
Flowers mostly in January and February, possibly March.
The fruits mature from April to June, rarely later in the rainy season.
Except for leaf differences, tepal length, and shape of stigma the species is very similar to C. drudei, which is treated in much greater detail. The leaf segments of C.
palmata are at least in part toothed more than halfway to base.
Native from Guatemala to Bolivia; introduced in the West Indies. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Col6n, Code, Panama, and Darien.
See Figs. 75 and 76.
20.
CYCLANTHACEAE/LUDOVIA I8I CYCLANTHUS Poit.Cyclanthus bipartitus Poit., Mem. Mus. Hist. Nat.
9:36. 1822
Portorrico, Lengua de buey
Acaulescent, monoecious herb, to 3.5 m tall. Petioles 1.5-2 m or more long; blades to 1.6 m long, almost com- pletely bisected, occasionally trifid, each segment 15-24 cm wide, the base of each lobe oblique, the midrib near upper edge at base, becoming centered by apex. Pedun- cles 40-90 cm long; spathes 5-7, broadly ovate, 15-25 cm long, the inner cream-colored or pinkish, the outer
± foliaceous; spikes 5-11 cm long when flowering, 1.5-2.5 cm broad, with staminate and pistillate flowers in 10-15 separate alternating whorls, the flowers of each whorl connate; staminate flowers naked; pistillate flowers en- closed by two rims of fleshy tissue, these becoming much enlarged; fruiting syncarps to 6 cm diam, ± pendent.
Each segment of syncarp falling off individually, with many seeds in a fleshy matrix; seeds to 1.5 mm long and 1 mm wide. Croat 5386, 6016.
Occasional, inhabiting chiefly stream banks and the lakeshore, though numerous juvenile plants are found within the forest away from any streams. Flowers mostly April and May, but also sometimes in June and perhaps later as well. The fruits mature throughout the rainy season, mostly by October.
The soft fleshy segments of the syncarp float and are dispersed in part by water currents and in part by animals or birds.
Guatemala to Peru and Brazil; Lesser Antilles. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, Chiriqui, Panama, and Darien and from tropical wet forest in Panama (beyond Cerro Jefe) and Col6n.
See Fig. 77.
LUDOVIA Brongn.
Ludovia integrifolia (Woods.) Harl., Acta Horti Berg.
18:338. 1958
Carludovica integrifolia Woods.
Nonrosulate epiphytic vine; stems 9-12 mm diam, at first green, becoming brown, forming long, ± tough, brown roots at most nodes; internodes mostly 5-7 cm long.
Leaves drying thin, all but the upper 3-8 withering but often persisting; petioles 10-15 cm long, narrowly cana- liculate; sheaths thin, extending one-third to almost entire length of petiole; blades narrowly elliptic to oblanceolate, acuminate, cuneate and often ± unequal at base (one side slightly shorter), 13-23(33) cm long, 4-7(17) cm wide, weakly bicolorous, the lower surface dull, with 4-7 major veins running almost parallel to the midrib but extending to the blade margin; margins weakly toothed at termination of each veinlet, the midrib raised on both surfaces but most prominent in basal third of blade.
Spathes 3, enveloping spadix, acuminate, greenish to white, the outer 6-8 cm long, the inner to 5 cm long, all caducous, breaking loose from base at anthesis and carried
Fig. 77. Cyclanthus bipartitus
Fig. 79. Ijjihvia itite^rifolia
upward by the enlarging spadix; spadix ca 3 cm long at anthesis; staminodia widely dispersed in clusters of 4 (rarely 5), to ca 3 cm long; staminate flowers ca 3.5 mm long, the perianth 20-30-lobed, glandular; pistillate flowers connate with each other, the tepals much reduced, the long, immersed stigmas sessile, longer than broad;
fruiting spadices 8.5-9.5 cm long, ca 1.5 cm diam, blunt, becoming orange, fleshy and tasty at maturity, the pedun- cles with 3 prominent spathe scars; stipes 5-7 mm long;
stigmas round, ca 0.5 mm diam. Fruits diamond-shaped, 11-17 mm long, 9-12 mm wide, conical; seeds numerous,
± ovoid, ca 1.5 mm long, embedded in a sweet, sticky, gelatinous matrix at maturity. Croat 12560.
Occasional, in the old forest. Reportedly flowers in June. Mature fruits have been recorded in November and December.
Obscurely resembling Stenospermation (21. Araceae), but distinguished by having more than one spathe.
Panama to Ecuador on the Pacific slope. In Panama, known from tropical moist forest in the Canal Zone and Darien.
See Figs. 78 and 79.