tendrils. Leaves alternate; petioles usually bearing promi- nent, often stalked glands; blades simple, usually 2- or 3-lobed, the margins entire or serrate; venation palmate, palmate at base, or rarely strictly pinnate; stipules present.
Flowers bisexual, actinomorphic, rarely apetalous, soli- tary or paired (rarely in racemes), axillary; peduncles jointed (often mistaken for a pedicel), bearing 3, often setaceous bracts; sepals 5, free, horned below the tip, often showy; petals 5, free, showy (lacking in P. coriacea);
corona (petaloid structure between petals and stamens) of several series; corona and perianth parts attached to hypanthium; operculum (ring of tissue within corona)
632 DICOTYLEDONEAE
membranous, sometimes plicate; stamens 5, attached to androgynophore (the fused staminate and pistillate parts);
anthers 2-celled, versatile, dehiscing longitudinally;
ovary superior, at apex of androgynophore, 1-locular, 3-carpellate; placentation parietal; ovules many, anatro- pous; styles 3 or stigmas sessile; stigmas 3, capitate. Fruits berries; seeds many, with fleshy endosperm, embedded in usually clear, fleshy, sweet mesocarp.
The family is represented on the island only by the genus Passiflora. The genus is recognized by being ten- driled vines with glandular petioles, by the showy, soli- tary or paired flowers with a modified corona and an androgynophore (complex structure containing both staminate and pistillate parts), and by the fleshy, many- seeded, usually tasty fruits.
The flowers are very specialized. Most probably follow the behavior of Passiflora foetida and P. vitifolia as re- ported by Janzen (1968). After the flower opens, the styles become deflected, which promotes outcrossing. During first visits to the flower the pollinating organism receives only pollen, but later visits bring it in contact with the stigmas. P. vitifolia, which is hummingbird pollinated, has strong stigmal deflection 30-180 minutes after open- ing. Janzen (1968) reported that the flowers open asyn- chronously between 5:30 and 7:30 A.M. and are repeatedly visited. Possibly all blue- and white-flowered species are bee pollinated (G. Frankie, pers. comm.), though the possibility of self-pollination (Lewis, 1966) cannot be overlooked. P. foetida is reported by Janzen to open syn- chronously in the early morning hours and is visited almost immediately by bees of the genus Ptiloglossa (Colletidae); stigmal deflections take place within 30 minutes and all visiting activity ends in about 45 minutes.
Flowers of Passiflora last but a single day and most wither well before the day is over.
Fruits are endozoochorous. Most seeds of the small, thin-walled fruits are probably dispersed chiefly by birds, whereas the larger-fruited species, such as P. ambigua, P. menispermifolia, P. nitida, P. seemannii, P. vitifolia, and P. williamsii, are probably dispersed chiefly by arbo- real frugivores. Peccaries and white-tailed deer eat fruits of P. vitifolia (N. Smythe, pers. comm.). Many of these species have rinds so thick and leathery that only the larger birds would attempt to peck them open. Once fruits are opened by monkeys and other animals, birds might be instrumental in the dispersal of any seeds left, but the fleshy, sweet, translucent substance covering the seeds of nearly all species of Passiflora is very tasty, and opened fruits are seldom found with any seeds remaining.
I suspect that animals spit out the seeds after sucking on them for a while because the seeds become very bitter or astringent.
Twelve genera and 600 species; subtropics and tropics.
PASSIFLORA L.
Passiflora ambigua HemsL, Bot. Mag. 128, pi.
7822. 1902
Coarse vine, essentially glabrous, with simple tendrils in leaf axils. Stipules filiform, inconspicuous, not persistent;
petioles 1.5-3 cm long, bearing 2 thick glands 0.5 to 1 cm above base; blades ovate to oblong-elliptic, acuminate, obtuse to subcordate at base, 10-20 cm long, 5-8 (9) cm wide, subcoriaceous. Flowers solitary, adjacent to ten- drils; peduncles ca 3 cm long (to 7 cm long in fruit), surmounted by 2 involucral bracts, the bracts free, trifid at apex, ca 3 cm long or longer, subfoliaceous; sepals 5, narrowly oblong, 4-6 cm long, maroon, with a slender appendage 2-4 mm long at apex; petals 5, narrowly
KEY TO THE SPECIES OF PASSIFLORA
Stipules subreniform, 1.5 cm long and 1 cm wide p menispermifolia H.B.K.
Stipules mostly linear, to 2 mm wide:
Blades peltate P coriaceaAdt_ juss_
Blades basmxed:
Blades 2- or 3-lobed at least to middle:
Blades deeply 3-lobed, cordate at base; flowers and fruits more than 3 cm across:
Palmate leaf veins 5; flowers deep scarlet, at least 10 cm diam; plants common P. vitifolia H.B.K.
Palmate leaf veins 3; flowers white with pink, at most 7 cm diam; plants rare
'•••'• P- williamsii Killip Blades broadly 2-lobed, rounded at base; flowers and fruits less than 3 cm across:
Blades more than 3 cm long along midvein; flowers paired P. biflora Lam.
Blades less than 2.5 cm long along midvein; flowers generally solitary P. punctata L.
Blades at most shallowly lobed (juvenile leaves of P. seemannii lobed):
Blades shallowly 3-lobed or with the lobes reduced to angles:
Petioles bearing 2 large auriculate glands near the base; flowers less than 3 cm diam, greenish-white p aunculata H.B.K.
Petioles eglandular; flowers more than 3 cm diam, white P. foetida L. var. isthmia Killip Blades not lobed:
Blades deeply cordate and palmately veined p_ seemannii Griseb.
Blades at most rounded at base, not palmately veined:
Blades at least serrulate; sepals lacking appendages p. nitida H.B.K.
Blades entire; sepals with slender appendage at apex p. ambigua HemsL
98.
PASSIFLORACEAE/PASSIFLORA633
lanceolate, 3-4 cm long, white outside, purplish inside;
stamens 5; filaments ca half as long as corolla, purplish.
Fruits ovoid to globose, to 15 cm long, greenish-yellow with whitish spots; exocarp 1-2 cm thick; seeds many, ca 8 mm long, with many small pits. Croat 11858, Zetek 3480, s.n.
Apparently rare; collected only once recently. Flowers from December to April. Mature fruits have been col- lected in the early rainy season, from May to July.
Southern Mexico to Colombia (Choco). In Panama, known from tropical moist forest in the Canal Zone (around Gatun Lake) and Bocas del Toro, from premon- tane wet forest in Code and Panama, and from tropical wet forest in Colon and Panama; no doubt in Darien as well since it has been found in adjacent Colombia.
Passiflora auriculata H.B.K., Nov. Gen. & Sp.
2:131. 1817
Vine, with simple axillary tendrils. Stipules bristle-like, to 1 cm long; petioles 1.5-2.5 cm long, biglandular near base, the glands large, ear-shaped, to ca 3 mm long; blades
± ovate, shallowly trilobate (generally with the lateral lobes reduced to angles), 7-15 cm long, 4-10 cm wide, glabrous to puberulent, with numerous brownish glands ca 1 mm diam. Flowers paired in leaf axils; peduncles ca 1 cm long in flower (ca 1.5 cm long in fruit), incon- spicuously jointed above middle, the bracts inconspic- uous; sepals 5, oblong-lanceolate, ca 1 cm long, foliaceous;
petals 5, linear, shorter than sepals, whitish; filaments of corona about as long as sepals; stamens 5, united to ca 4 mm at base. Berries globose, 1-2 cm diam, yellow-green, becoming black when mature, scattered-puberulent; seeds many, ca 3.5 mm long, reticulate, dark. Croat 5617.
Occasional along the shore and to be expected in clear- ings. Flowers from October to March (sometimes from August), with the fruits maturing from March to July.
Nicaragua to Brazil and Bolivia. In Panama, known principally from premontane wet forest in Colon, Code, Panama, and Darien; known also from tropical moist forest in the Canal Zone (Atlantic slope) and Bocas del Toro and from tropical wet forest in Panama and Darien.
Passiflora biflora Lam., Encycl. Meth. Bot. 3:36. 1789 Guate-guate, Camacarlata, Calzoncillo
Vine, woody at base, essentially glabrous; stems striate.
Stipules minutely setose, with 4 or 5 pairs of round glands essentially only in the central part, the first pair at the very base; petioles eglandular, to 2 cm long; blades broadly bilobed (occasionally with a small central lobe as well), V-shaped, obtuse to rounded at base, 3-5 (10) cm long in the center, 4-9 cm wide, distinctly lighter below, glabrescent, with a slender apiculum at apex of midvein and each lobe; palmate veins 3. Flowers paired in axils, 3-4 cm diam; peduncles to 2 cm long, jointed at or below the middle, with 3 slender bracts at articula- tion; hypanthium ca 3 mm deep, depressed at point of attachment to pedicel; sepals 5, lanceolate, 1.5-2 cm long, hooded, creamy white; petals 5, lanceolate, 1—1.5 cm long, creamy white; corona filaments in 2 series, the outer of
oblong filaments ca 7 mm long, yellow, the inner of narrowly linear filaments ca 5 mm long, reddish; oper- culum closely plicate, 3 mm high; gynophore ca 5 mm long; stamens 5, fused into tube around style; anthers dorsifixed, held below stigma, spreading and dehiscing extrorsely; ovary ± tomentulose. Berries globose, 1.5-1.8 cm diam, yellow-green and mottled, becoming purple when mature; seeds many, to 3 mm long, reticulate, becoming dark. Croat 5416, 5707.
Occasional, along the shore over low vegetation in Annona-Acrostichum associations and in clearings. Flow- ers and fruits throughout the year.
Mexico to Colombia and Venezuela. In Panama, wide- spread and ecologically variable; known from tropical moist forest in all provinces, from tropical dry forest in Code and Panama, from premontane moist forest in the Canal Zone and Panama, from premontane wet forest in Chiriqui, Veraguas, and Panama, from tropical wet forest in Colon and Darien, and from premontane rain forest in Chiriqui.
Passiflora coriacea Adr. Juss., Ann. Mus. Natl. Hist.
Nat. 6:109. 1805
Vine, sometimes trailing on the ground, ± glabrous.
Stipules narrowly linear, ca 5 mm long, persistent; peti- oles 1-4 cm long, bearing a pair of ear-shaped glands in apical half; blades peltate (the petiole attached 3-13 mm from base of blade), trilobate, broadly rounded to cordate at base, 3-7 cm long, 6-25 cm wide, the central lobe occasionally suppressed to a mucro, the lateral lobes nearly horizontal, abruptly acuminate; palmate veins 5.
Flowers generally paired, axillary but also racemose, the racemes opposite the leaves; peduncles jointed above the middle, the bracts inconspicuous and deciduous;
sepals 5, lanceolate, ca 1.5 cm long, greenish-white to cream-colored, apetalous; corona in 2 series, the outer of filaments to 10 mm long, the inner of a few glandular filaments at most 5 mm long; operculum not generally plicate, ca 2 mm high; gynophore ca 10 mm long; ovary glabrous. Berries globose, 1-2 cm diam, the exocarp generally fleshy, glaucous, green and perhaps speckled when immature, maturing dark blue; seeds many, ca 5 mm long, with about a dozen contiguous pits on each side, drying dark. Croat 5414a, 6732.
Occasional, on the shore and in the young forest. Flow- ers throughout the year, principally in the dry season, with most fruits maturing from May to July.
Southern Mexico to the Guianas, Bolivia, and Peru.
In Panama, known from tropical moist forest in the central Canal Zone and from premontane wet forest in Code and Panama.
Passiflora foetida L. var. isthmia Killip, Publ. Field Mus. Nat. Hist., Bot. Ser. 19:497. 1938
P. hispida DC.
Vine, often creeping on the ground, yellow-pubescent on all parts, the trichomes ca 2 mm long, sometimes gland- tipped. Stipules about half encircling stem, deeply cleft with linear lobes, bearing glandular trichomes; petioles
Fig. 406. Passiflorafoetida var. isthmia
Fig. 407. Passiflora nitida
98.
PASSIFLORACEAE/PASSIFLORA635
2-8 cm long, eglandular but with glandular trichomes;
blades variable, deltoid in ours, shallowly trilobate (the lobes abruptly acuminate), cordate at base, 5-8 cm long and 5-8 cm wide; palmate veins 5-7. Flowers strongly scented, solitary in leaf axils; peduncles 3-6 cm long, jointed directly beneath the flower, the bracts foliaceous, 2-3.5 cm long, 2- or 3-pinnatifid, the ultimate segments filiform, gland-tipped; sepals 5, ± lanceolate, ca 2 cm long, white; petals 5, similar to sepals but slightly shorter;
filaments of corona in several series, the longer to 1.5 cm long, purple below, the shorter ca 2 mm long; operculum erect, ca 1 mm long, denticulate. Berries inflated, ovoid, yellowish, ca 2.5 cm long, minutely apiculate at apex, subcordate at base, usually obscurely 3-sulcate and with 3 indistinct lines of dehiscence (apparently nonfunc- tional?); exocarp very thin; seeds numerous, ovoid- flattened, ca 5 mm long and 2.5 mm wide, slightly sculp- tured, black, shiny, trilobate at base, bearing at apex a slender funicle ca twice length of seed. Croat 4641, 7400.
Occasional, in clearings. Flowers and fruits throughout the year. In Costa Rica the species begins to flower in June (beginning of the rainy season) and flowers into the following dry season (Janzen, 1968).
The species is a cosmopolitan weed native throughout the American tropics; the variety isthmia extends from Panama along the Pacific coast to Ecuador. In Panama, widespread in weedy areas; known from tropical moist forest in the Canal Zone, all along the Atlantic slope, and in Panama and Darien, from tropical dry forest in Panama, from premontane wet forest in Bocas del Toro and Chiriqui, and from tropical wet forest in Colon and Darien.
See Fig. 406.
Passiflora menispermifolia H.B.K., Nov. Gen. & Sp.
2:137. 1817
Vine; stems densely hispid-hirsute. Stipules conspicuous, subreniform, ca 1.5 cm long and 1 cm wide, cuspidate at one end, usually glandular-denticulate; petioles 1.5-5(9) cm long, bearing 4-8 glands; blades suborbicular, 8-14 cm long, 8-13 cm wide, angulately trilobate, cordate at base, sometimes glandular-denticulate, definitely lighter and pilose below, densely pilose above, the middle lobes much larger, to 8 cm wide; palmate veins 5-7. Flowers solitary; peduncles 2-3 cm long, jointed directly beneath the flower, the bracts narrowly lanceolate, ca 10 mm long, at most 3 mm wide; sepals 5, lanceolate-oblong, 2-3 cm long, greenish-white, awned at apex, the awn little, off- white; petals 5, oblong, 2.5-3.5 cm long, light violet;
filaments of corona in several series, the outermost to 2.5 cm long, glandular; operculum membranous, erect, divided above into numerous filaments ca 5 mm long;
stigmas 3, ca 13 mm long. Berries narrowly ovoid, 6 cm long or more, ca 2 cm diam, glabrous; seeds ca 5 mm long, coarsely reticulate (fruit description from Flora of Panama, Woodson & Schery, 1958). Aviles 21b, Wood- worth & Vestal 634.
Not seen on the island in recent years. Seasonal behav-
ior uncertain. Probably flowering and fruiting throughout the year.
Nicaragua to northwestern South America. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Colon, Veraguas, and Panama and from premontane wet forest in Chiriqui (Boquete).
Passiflora nitida H.B.K., Nov. Gen. & Sp. 2:130. 1817 Coarse vine, glabrous. Stipules linear; petioles 1.5-2 cm long, biglandular above the middle; blades ovate-oblong, acuminate, rounded to subcordate at base, 11-19 cm long, 6-11 cm wide, subcoriaceous, serrulate. Flowers solitary;
peduncles 2-3.5 cm long, jointed directly beneath the flower, bracteate, the bracts 2, conspicuous, ovate-elliptic, ca 3.5 cm long and 2.5 cm wide, rounded at apex and base; sepals 5, oblong-elliptic, to 5 cm long, greenish outside, white inside; petals 6, similar, white; filaments of corona about 4 cm long, white with pink and blue;
stamens 5, purple; pistil pinkish. Berries obovoid, blunt at apex, red-orange to orange-red or magenta, densely punctate with lighter dots (these often run together), 6-8 cm long, 4.5-6 cm wide, glabrous, the withered flower often persisting; pericarp 1-1.5 cm thick, spongy;
seeds obovate, ca 7 mm long, punctate, black. Shattuck 675, Zetek 3620.
Occasional, high in the forest canopy. Flowers in Panama from December to May. Mature fruits have been seen throughout the year, mostly from April to September.
The Panamanian collections of this species have gener- ally greater dimensions than those from South America.
Also, the petiolar glands are slightly below the apex (about one-third of the way down the petiole), instead of at the apex as on South American species.
Panama to the Guianas, northern Brazil, and Peru.
In Panama, known from tropical moist forest on BCI, from premontane wet forest in the Canal Zone, Colon, and Panama, and from premontane rain forest in Darien (summit of Cerro Pirre).
See Fig. 407.
Passiflora punctata L., Sp. PI. 957. 1753 P. misera H.B.K.
Slender vine, essentially glabrous; stems striate. Stipules usually persistent, narrowly subulate, ca 2 mm long; leaf blades of 2, ± oblong lobes directed at nearly 180° angle to each other (rarely with a small central lobe), truncate or subcordate at base, 1-2.5 cm long, 5-13 cm broad, the lobes rounded and mucronulate, the lower surface glau- cescent with 3-5 pairs of round glands usually along midribs especially in axils of lateral veins; palmate veins 3. Flowers generally solitary in leaf axils; peduncles 3-9 cm long, obscurely jointed immediately below to 1 cm below the flowers, the bracts inconspicuous, setaceous, ca 1 mm long; calyx tube campanulate, ca 4 mm deep, the lobes 5, lanceolate, ca 1.5 cm long, light yellowish-green;
petals 5, oblong, ca 1 cm long, greenish-white; filaments of corona in 2 series, the outer to 10 mm long, mostly
636
DICOTYLEDONEAEpale purple, the inner ca 5 mm long, purplish, narrower;
operculum plicate, ca 3 mm high, incurved, purplish;
gynophore slender, to 8 mm long, purple; ovary narrowly ellipsoid, to 4 mm long, purple, capitellate. Berries glo- bose, ca 1.5 cm diam, deep blue; seeds ovate, ca 3 mm long, transversely ridged, the ridges rugulose. Croat 8273, 8278.
Uncommon, along the shore. Flowers and fruits throughout the year.
The distinction is obscure between this species and P. misera H.B.K., which is therefore not recognized here.
The petals, sepals, and outer series of corona filaments reflexed somewhat at anthesis. Anthesis occurs in the early morning, and plants collected at 10:30 A.M. usually have all flowers closed. Nectar is probably enclosed within the operculum, within which many holes are seen.
Panama to Argentina. In Panama, known only from tropical moist forest around Gatun Lake.
Passiflora seemannii Griseb., Bonplandia 6:7. 1858 Guate-guate
Stout vine, glabrous; stems to at least 5 mm diam, striate.
Stipules somewhat persistent, narrowly linear, 10-15 mm long; petioles 3-10 cm long, biglandular near apex; blades broadly ovate-cordate, rounded and mucronulate to apiculate at apex, 5-10 cm long, 6-11 cm wide, the basal sinus ca 3 cm deep, the lower surface glaucous, eglandu- lar, the margins subentire to serrulate; palmate veins 7-9; juvenile leaves trilobate. Flowers solitary in leaf axils, ca 8 cm diam; peduncles 6-9 cm long (to 10 cm long when in fruit), jointed directly beneath flowers, the bracts 3, fused, 2.5-5 cm long, the lobes broadly ovate, white, purple-tinged; hypanthium ca 2 cm deep; sepals 5, ovate-lanceolate, 3.5-4 cm long, appendaged, greenish- white, sometimes tinged with violet; petals 5, oblong- lanceolate, ca 3.5 cm long, purple, the entire perianth strongly reflexed when open; corona filaments in 2 series, the inner erect, incurved at apex, to 2.5 cm long, banded with purple and white, the outer ca 1 cm long; staminal filaments fused except at apex; anthers shedding pollen after beginning of fruit development; operculum 2 mm high, denticulate; gynophore ca 2 cm long in flower;
ovary ovoid; styles 3; stigmas broad, yellow. Berries broadly oblong to ovoid, at least 4-5 cm long and 3.5 cm broad, green, weakly pruinose, densely speckled with light green spots, the exocarp thick, white inside; seeds many, stalked, orbicular-ovate, ca 3 mm long, punctate.
Shattuck 692, Wetmore & Abbe 147.
Uncommon along the shore. Flowers mostly from October to March, sometimes from August. The fruits mature from January to March.
Native to Panama and Colombia; cultivated elsewhere.
In Panama, known principally from tropical moist forest in the Canal Zone, all along the Atlantic slope, and in Panama and Darien; known also from premontane wet forest in Chiriqui and Panama and from tropical wet forest in Colon and Panama.
See Fig. 408.
Passiflora vitifolia H.B.K., Nov. Gen. & Sp. 2:138. 1817 Guate-guate, Pasionaria, Granadilla, Granadillo de monte Liana; stems to at least 1.5 cm diam, ± densely ferruginous-pubescent on all parts. Stipules deciduous, subsetaceous, ca 5 mm long; petioles 2-6 cm long, incon- spicuously glandular toward base; blades deeply trilobate,
± deeply cordate, 7-14 cm long, 9-14 cm wide, the lobes acute to acuminate at apex, the central lobe larger, the upper surface sparsely strigillose except on tomentulose veins, the lower surface softly ferruginous-pubescent,
the glands minute, 2 to several, around base of each sinus; palmate veins 5. Flowers solitary in axils, often borne on young leafless branches, 10-15 cm wide when open; peduncles 3-5.5 cm long (to 6.5 cm long in fruit), the bracts 3, free, lanceolate, 2.5-3 cm long, glabrescent, glandular-serrate, generally with a larger (1 mm across) disk-shaped pair of glands toward the base; sepals 5, narrowly lanceolate, 6-8 cm long, 1-2 cm wide, with a long slender appendage at apex; petals 5, linear-lanceolate, 4-6 cm long; both sepals and petals deep scarlet to ma- genta, reflexed when open; corona filaments in 3 series, the outer longest, to 2 cm long, erect, bright red or bright yellow; anthers 5, green, held perpendicular on gyno- phore; operculum deflexed, to 1 cm long, fimbriate; gyno- phore ca 1 cm long, pale red; ovary densely tomentulose.
Berries ovoid, 5-8 cm diam, puberulent, greenish-yellow with darker stripes and bands of lighter splotches, very fragrant; seeds ± flattened, ca 6 mm long and 5 mm wide, reticulate with numerous small punctations, light-colored.
Croat 4766, 8319.
Frequent in the forest. Flowers from December to May, elsewhere in Panama also from July to November.
Fruit maturity time not determined.
Native to lowland forests from Nicaragua to Venezuela and Peru; cultivated in the West Indies and elsewhere.
In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, Panama, and Darien, from premontane wet forest in Colon, Code, and Pa- nama, and from tropical wet forest in Colon, Code, Panama, and Darien.
Passiflora williamsii Killip, J. Wash. Acad. Sci.
12:262.1922
Coarse vine, ± densely puberulent on all parts. Stipules 6-7 mm long; petioles 4-6 cm long, bearing a pair of prominent glands 1-2 cm from base and sometimes a second pair above them; blades trilobate to about middle, truncate to generally deeply cordate at base, 9-13 cm long and wide, the lobes acuminate, the central lobe stronger and contracted toward the base, the upper surface gla- brescent except on strigillose veins, the lower surface minutely crisp-villous, sparsely so in age, distinctly bi- glandular at base of each sinus, the glands ca 1 mm long, the margins subentire to serrulate; palmate veins at base 3. Flowers solitary in axils, 6-7 cm diam; peduncles to 2 cm long in flower, the bracts 3, united at base, ca 2.5 cm long, tomentulose; sepals 5, oblong, 2.5-3.5 cm long, with a slender appendage at apex; petals 5, oblong-spatulate,