PASSIFLORACEAE

西番莲科 xi fan lian ke

Wang Yinzheng (王印政)1; Shawn E. Krosnick2, Peter M. Jørgensen3, David Hearn4
Climbing herbs or lianas with axillary tendrils, rarely shrubs or trees, vegetative ramification through a supra-axillary bud.
Leaves alternate (rarely subopposite), spirally arranged, simple or rarely compound, stipulate or not, petiolate, petiole or base of
blade often with 1 to many glands; leaf blade entire or lobed, venation pinnate or palmate, often with glands on margin and abaxial
surface. Inflorescence cymose, axillary, sessile or pedunculate, primary axis often a tendril, secondary axis often reduced; bracts
minute to foliaceous, sometimes glandular. Pedicels articulate distal to bracts. Flowers mostly actinomorphic, bisexual, functionally
unisexual, or polygamous; perianth persistent, segments free or partially fused at base. Sepals (3–)5(or 6). Petals (3–)5(or 6), imbri-
cate. Corona inserted on hypanthium, composed of filaments or hairs often in multiple series. Stamens (4 or)5(–60), inserted on an-
drogynophore or at base of hypanthium; filaments free to base or sometimes partially fused; anthers dorsifixed, 2-celled, dehiscing
via longitudinal slits, sometimes with apical appendage. Ovary (1–)3(–5)-carpellate, superior, 1-loculed, sessile or stipitate on elon-
gate androgynophore; placentation parietal; ovules many, anatropous; styles (1–)3(–5), free to partially fused at base; stigmas glo-
bose, capitate, papillate, or divided. Fruit a berry or loculicidally 3(–5)-valved capsule. Seeds numerous, compressed, often beaked,
enclosed by fleshy aril; testa reticulate or pitted; endosperm copious, oily; embryo straight; cotyledons foliaceous.
About 16 genera and 660 species: mainly in tropical and subtropical areas, particularly diverse in the New World tropics; two genera and 23
species (seven endemic, seven introduced) in China.
Bao Shihying. 1999. Passifloraceae. In: Ku Tsuechih, ed., Fl. Reipubl. Popularis Sin. 52(1): 97–120.

1a. Androgynophore long; flowers large, bisexual; corona well developed; fruit a berry ................................................... 1. Passiflora
1b. Androgynophore absent or short; flowers small, unisexual or polygamous; inner corona lacking, outer corona
reduced; fruit a capsule .......................................................................................................................................................... 2. Adenia

1. PASSIFLORA Linnaeus, Sp. Pl. 2: 955. 1753, nom. cons.

西番莲属 xi fan lian shu

Wang Yinzheng (王印政); Shawn E. Krosnick, Peter M. Jørgensen

Granadilla Miller; Tripsilina Rafinesque.

Herbaceous or woody perennial vines, rarely shrubs or trees. Leaves simple or rarely compound, alternate (subopposite in one
species), entire or dissected, petiolate, usually with extra-floral nectaries on petiole and/or blade; stipules linear to leaflike, often
glandular. Inflorescence axillary, cymose; peduncle often highly reduced or absent, central axis developed into a tendril, secondary
axes often highly reduced to 1 or 2 flowers; bracts minute to foliaceous, sometimes glandular. Flowers bisexual (rarely plants
dioecious). Hypanthium broad to campanulate. Sepals 5, often petaloid, sometimes with a subapical projection. Petals 5 (rarely
absent). Corona present at base of perianth in one to several series of showy filaments; innermost series (operculum) membranous,
partially to entirely fused, margin entire or fimbriate, often incurved over nectar chamber; extra-staminal nectariferous disk (limen)
present around base of androgynophore, fused to base of hypanthium. Stamens (4 or)5(–8); filaments free (rarely connate into a tube
around ovary); anthers linear or oblong, dorsifixed, versatile. Ovary on androgynophore, stipitate or sessile, 3(–5)-carpellate; styles
3(–5), free; stigmas capitate. Fruit a berry (rarely a dehiscent capsule). Seeds arillate, compressed, testa pitted; endosperm oily, abun-
dant; embryo straight; cotyledons elliptic or oblong-elliptic; germination epigeal (rarely hypogeal).
About 520 species: mainly in tropical America and tropical Asia; 20 species (seven endemic, seven introduced) in China.

Passiflora “alato-caerulea” (Lindley, Bot. Reg. 10: t. 848. 1824), a hybrid of horticultural origin, is grown in China and elsewhere for its
flowers and robust, vigorous vegetative growth. The epithet “alato-caerulea” is considered to be a hybrid formula (i.e., P. alata Curtis × P. caerulea
Linnaeus) and not a true epithet (Vienna Code, Art. H.10.3).

The reader is referred to the following references: Killip, The American Species of Passifloraceae (Publ. Field Mus. Nat. Hist., Bot. Ser. 19(1–2):
1–331, 333–613. 1938); de Wilde, A monograph of the genus Adenia Forsk. (Passifloraceae) (Meded. Landbouwhogeschool Wageningen 71(18): 1–
281. 1971); and de Wilde, The Indigenous Old World Passifloras (Blumea 20: 227–250. 1973 [“1972”]).

1a. Bracts not entire, pinnatifid or lacerate.

1 Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.
2 UC Davis Center for Plant Diversity, Plant Sciences MS#7, University of California, One Shields Avenue, Davis, California 95616-8780, U.S.A.
3 Missouri Botanical Garden, P.O. Box 299, Saint Louis, Missouri 63166–0299, U.S.A.
4 Plant Sciences, University of Arizona, Forbes Building, Room 303, 1140 East South Campus Drive, Tucson, Arizona 85721-0036, U.S.A.

141
142 PASSIFLORACEAE

2a. Bracts pinnatifid, lobes apically with glandular hairs; leaves shallowly 3-lobed or entire; flowers 2–3 cm in
diam.; petiole without glands .................................................................................................................................... 16. P. foetida
2b. Bracts lacerate, not glandular; leaves 3- or 5-lobed, never entire; flowers 3–5 cm; petiole with 2 stipitate
glands ................................................................................................................................................................. 15. P. adenopoda
1b. Bracts entire, or caducous, never with glandular hairs.
3a. Leaves unlobed, entire.
4a. Inflorescence reduced to only one flower; outer corona in 5 or 6 series.
5a. Stem 4-angled; leaves broadly ovate or suborbicular; stipules large, leaflike, 2–4 cm; petiole
with 4–6 glands; fruit ovoid, 20–25 cm ........................................................................................ 19. P. quadrangularis
5b. Stem terete; leaves ovate-oblong; stipules linear; petiole with 2 glands; fruit globose, 5–6 cm
in diam. .................................................................................................................................................... 20. P. laurifolia
4b. Inflorescence of more than one flower; coronal filaments in 1 or 2 series.
6a. Laminar nectaries absent on abaxial surface; petiolar glands 2, small, at middle of petiole;
inflorescence 1- or 2-flowered ....................................................................................................... 4. P. kwangtungensis
6b. Laminar nectaries present on abaxial surface; petiolar glands various; inflorescence 2- to many
flowered.
7a. Leaves broadly ovate to cordate, with 20–50 tiny nectaries scattered on abaxial surface; sepals
with apical appendage ...................................................................................................................... 5. P. eberhardtii
7b. Leaves ovate-elliptic, broadly lanceolate, linear-lanceolate, or suborbicular, with 2 to many
nectaries on abaxial surface; sepals lacking apical appendage.
8a. Leaves suborbicular to broadly orbicular; inflorescence 2–8-flowered ........................................... 6. P. henryi
8b. Leaves linear-lanceolate, broadly lanceolate, or lanceolate to ovate; inflorescence 2–16-flowered.
9a. Flowers with 5 stamens and 3 carpels; leaves ovate, subopposite to opposite,
abaxially white pubescent .......................................................................................... 2. P. cochinchinensis
9b. Flowers with variable numbers of stamens and carpels; leaves lanceolate to
oblanceolate, glabrous or pubescent.
10a. Stem and leaves pubescent; leaves oblanceolate, alternate ........................................... 1. P. siamica
10b. Stem and leaves glabrous; leaves lanceolate, spirally arranged ............................... 3. P. tonkinensis
3b. Leaves 2- or 3-lobed or 5-palmatifid.
11a. Inflorescences 1-flowered; coronal filaments in 2–6 series.
12a. Leaves (3–)5(–9)-palmatifid, lobes entire; petiole with 2–4(–6) glands; stipules large, reniform,
clasping ................................................................................................................................................ 18. P. caerulea
12b. Leaves deeply 3-lobed, lobes dentate; petiole with 2 glands apically; stipules linear-subulate ............ 17. P. edulis
11b. Inflorescences 2–20-flowered; coronal filaments in 1 or 2 series.
13a. Laminar nectaries absent on abaxial surface; petals absent; corona in 2 series ................................ 14. P. suberosa
13b. Laminar nectaries present on abaxial surface (2–25); petals present; corona in 1 or 2 series.
14a. Coronal filaments ca. 1/2 as long as sepals or shorter; leaves truncate, lobes generally
equal in length.
15a. Petiole with 2 discoid nectaries at base; laminar nectaries in 2 lines near midvein
...................................................................................................................................... 8. P. jianfengensis
15b. Petiole with 2 peg-shaped nectaries; laminar nectaries 2–4 on abaxial surface.
16a. Plants strongly pubescent throughout; leaf blade often with 2 or 3 pairs of glands
near apex of midvein on abaxial surface, apex truncate, usually 3-lobed ................ 9. P. jugorum
16b. Plants glabrous, rarely glabrescent; leaf blade usually with 1 pair of glands near
apex of midvein on abaxial surface, apex strongly truncate, usually 2-lobed ........ 10. P. wilsonii
14b. Coronal filaments more than 1/2 as long as sepals; leaves shallowly to deeply lobed, lobes
unequal in length, midvein often conspicuously shortened.
17a. Lobes of leaves apically obtuse; abaxial surface with 6–25 nectaries; sepals with
apical appendage ............................................................................................................. 7. P. cupiformis
17b. Lobes of leaves apically acute; abaxial surface with 2–8 nectaries; sepals lacking awn.
18a. Inflorescence 5–8-flowered; leaves truncate, lobes ovate-oblong; petiole with
2 conspicuous, auriculate glands ............................................................................... 11. P. papilio
18b. Inflorescence 1–16-flowered; leaves deeply truncate; base of leaves lanceolate.
19a. Stem striate; leaves and stems pubescent; flowers 2–16 per
inflorescence; coronal filaments white throughout ................................. 12. P. altebilobata
19b. Stem subtriangular, smooth; leaves and stems glabrous, veins
white; flowers 1 or 2 per inflorescence; coronal filaments
brown at base, yellow at apex ..................................................... 13. P. xishuangbannaensis
 PASSIFLORACEAE 143

1. Passiflora siamica Craib, Bull. Misc. Inform. Kew 1911: 55. 3. Passiflora tonkinensis W. J. de Wilde, Blumea 20: 241.
1911. 1973 [“1972”].
长叶西番莲 chang ye xi fan lian 长叶蛇王藤 chang ye she wang teng
Passiflora octandra Gagnepain; P. octandra var. attopen- Passiflora cochinchinensis Sprengel subsp. glaberrima
sis Gagnepain; P. octandra var. cochinchinensis Gagnepain. (Gagnepain) Cusset; P. moluccana Reinwardt ex Blume var.
glaberrima (Gagnepain) W. J. de Wilde; P. octandra Gagne-
Herbaceous vines to lianas, 6–8 m long. Stem ± angular,
pain var. glaberrima Gagnepain.
striate; younger stems densely pubescent. Petiole 1–4 cm, often
with 2 ovoid glands at middle; leaf blade ovate-elliptic to lan- Herbaceous vines to lianas, ca. 6 m long. Stem terete,
ceolate, 6–18(–25) × 2.5–7 cm, leathery, usually pubescent, abax- glabrous. Leaves spirally arranged; petiole 1–3 cm, with 2 dis-
ially with (0–)2–8 laminar nectaries, base cuneate to sub- coid glands 2–8 mm from base of blade; leaf blade lanceolate or
cordate, apex acute or acuminate. Inflorescences (sub)sessile, elliptic to oblong, 6–10 × 2.5–4(–6) cm, membranous, both sur-
axillary, (2–)4–15-flowered, sometimes rust-brown pubescent. faces glabrous, laminar nectaries 4–6, base subcordate, apex
Pedicel 3–15 mm, pubescent. Flowers white, 2.5–3 cm in acute. Inflorescences axillary, (sub)sessile, 1- or 2-flowered, gla-
diam.; hypanthium 7–10 mm in diam. Sepals 9–12 × 6–7 mm, brous; bracts linear. Pedicel 5–25 mm, glabrous. Flowers whit-
margin membranous, pubescent. Petals 9–10 × 3–4 mm. Co- ish, 3–4.5 cm in diam.; hypanthium 6–9 mm in diam. Sepals
rona in 2 filamentous series; outer series 7–10 mm, inner series green, 1.5–2 cm × ca. 5 mm, glabrous. Petals white, 1.2–1.6
1.5–4 mm; operculum incurved, plicate, 2–2.5 mm high, mem- cm × 3–4 mm. Corona in 2 filamentous series; outer series 1–
branous; disk ca. 0.5 mm high; androgynophore 3–5 mm tall. 1.6 cm, yellow at base, purple at apices; inner series 1–3 mm,
Stamens 5–8; filaments 1–1.1 cm, basally connate for 1.5–4.5 filaments yellow, apices capitate; operculum incurved, green,
mm. Ovary (sub)sessile, ellipsoid, 2–3 × 1.5–2.5 mm, pubes- plicate, 1.5–2 mm tall, membranous; disk purple-brown, ca.
cent; styles 3–5, 4–9 mm, fused basally up to 2 mm. Fruit 1 or 0.5 mm high; androgynophore 5–10 mm tall. Stamens (5 or)6–
2 per inflorescence, blue at maturity, subglobose, 1.5–2 cm in 8, filaments 6–10 mm, connate and enclosing ovary; anthers 3–
diam., sparsely pubescent. Fl. Mar–Apr, fr. Jun–Jul. 5 × 1.5–2 mm. Ovary (sub)sessile on androgynophore, ellipsoid,
Forests, scrub; 500–1600 m. EC Guangxi (Guiping), S Yunnan glabrous; styles (3 or)4 or 5, 5–8 mm. Fruit blue at maturity,
(Xishuangbanna) [India, Laos, Myanmar, Thailand, Vietnam]. 1.5–2.5 cm in diam., glabrous.
2. Passiflora cochinchinensis Sprengel, Syst. Veg. 4: (Cur. Scrub in mountain valleys; 100–200 m. SE Yunnan (Hekou)
Post): 346. 1827. [Laos, Vietnam].

蛇王藤 she wang teng 4. Passiflora kwangtungensis Merrill, Lingnan Sci. J. 13: 38.
1934.
Anthactinia horsfieldii M. Roemer; Disemma horsfieldii
Miquel; D. horsfieldii var. teysmanniana Miquel; Passiflora 广东西番莲 guang dong xi fan lian
hainanensis Hance; P. horsfieldii Blume; P. horsfieldii var.
elbertiana H. G. Hallier; P. ligulifolia Masters; P. moluccana Herbaceous vines, 5–6 m long. Stem slender, glabrous,
Reinwardt ex Blume var. teysmanniana (Miquel) W. J. de Wilde; thinly striate. Petiole 1–2 cm, with 2 glands in distal 1/3; leaf
P. pallida Loureiro (1790), not Linnaeus (1753); P. philippinen- blade lanceolate to oblong-lanceolate, 6–13 × 2–4(–6) cm,
sis Elmer. membranous, abaxially puberulous, laminar nectaries absent,
base cordate, margin entire, apex long acuminate. Inflores-
Herbaceous vines, ca. 6 m long. Stem striate, sparsely pu- cences sessile, 1- or 2-flowered, glabrescent. Pedicel 1–1.5
bescent. Leaves subopposite, dimorphic; petiole 7–10 mm, with cm. Flowers white, 1.5–2 cm in diam.; hypanthium 4–5 mm
two glands 2–8 mm from base of blade; leaf blade lanceolate to in diam. Sepals 7–9 mm × ca. 2.5 mm, membranous. Petals 6–7
elliptic, 6–10 × 2.5–4(–6) cm, leathery, abaxially canescent, pin- × 2–3 mm. Corona a single filamentous series, white, 2–3 mm,
ninerved, abaxially with 4–6 laminar nectaries, base subcordate, apices clavate; operculum plicate, ca. 1.5 mm high, membra-
apex obtuse or rounded. Inflorescence sessile, 2(–12)-flowered, nous; disk ca. 0.3 mm high; androgynophore ca. 4.5 mm tall.
± glabrous; bracts linear. Pedicel 5–25 mm, glabrous. Flowers Stamens 5; filaments ca. 3.5 mm, basally connate for ca. 1 mm;
pinkish white, 3–4.5 cm in diam.; hypanthium 6–9 mm in diam. anthers oblong, ca. 2.5 × 1 mm. Ovary subsessile on andro-
Sepals greenish, with occasional purple mottling, 1.5–2 cm × gynophore, ellipsoid, ca. 2.5 × 1.5 mm, glabrous; styles 3, free,
ca. 5 mm, glabrous, margin membranous. Petals whitish, 1.2– 2.5–4 mm; stigmas capitate. Fruit blue at maturity, globose, 1–
1.6 cm × ca. 5 mm. Corona in 2 series; outer series 1–1.6 cm, 1.5 cm in diam., glabrous. Fl. Mar–May, fr. Jun–Jul.
brown at base, white at middle, purple at apices; inner series 1–
3 mm, brownish; operculum incurved, brownish, plicate, 1.5–2 ● Scrub at forest margins; 600–700 m. N Guangdong, NE Guang-
xi, SE Jiangxi.
mm tall, membranous; disk dark brown, ca. 0.5 mm high; an-
drogynophore 5–10 mm tall. Stamens 5; filaments (5–)6–10 5. Passiflora eberhardtii Gagnepain in Lecomte, Fl. Indo-
mm, free; anthers 3–5 × 1.5–2 mm. Ovary (sub)sessile, ellip- Chine 2: 1020. 1921.
soid, 2–4 × 1.5–3 mm, glabrescent; styles 3, free, 5–8 mm.
Fruit blue at maturity, 1.5–2.5 cm in diam., glabrescent. 心叶西番莲 xin ye xi fan lian
Scrub in mountain valleys; 100–1000 m. Guangdong, Guangxi, Herbaceous vines, 2–3 m long. Stem terete, pubescent.
Hainan [Laos, Malaysia, Vietnam]. Petiole (2–)5–6 cm, pubescent, with two large discoid glands at
144 PASSIFLORACEAE

base; leaf blade broadly ovate, ca. 15 × 12 cm, membranous to mm tall. Stamens 5; filaments 4.5–6 mm, free; anthers ca. 2.5 ×
leathery, puberulous to densely pubescent, abaxially with 20–50 1.5 mm. Ovary sessile on androgynophore, subglobose, ca. 2 ×
small scattered laminar nectaries, base cordate, margin entire, 1.5–2 mm, pubescent; styles 3, free, ca. 4 mm. Fruit blue at
apex acute or shortly acuminate. Inflorescences (sub)sessile, maturity, globose, 1–1.6 cm in diam., glabrous. Fl. Apr, fr. Sep.
15–20(–30)-flowered. Pedicel 1–2 cm. Flowers white, 1–1.5 cm
Grass and scrub on slopes, valleys, roadsides; 1000–2000 m.
in diam. Sepals ca. 6 × 3 mm, awn 1.5–2 mm, pubescent. Petals
Guangdong, Guangxi, Hubei (Badong), Sichuan, Yunnan [Vietnam].
ca. 5 × 2 mm. Corona in a single series, 1.5–3 mm; operculum
plicate, ca. 1.2 mm tall, membranous; disk absent; androgyno- 8. Passiflora jianfengensis S. M. Hwang & Q. Huang, Acta
phore ca. 2 mm tall. Stamens 5; filaments ca. 2.5 mm, free; an- Phytotax. Sin. 23: 64. 1985.
thers ca. 2.5 × 0.7 mm. Ovary sessile on androgynophore, ob-
void, pubescent; styles 3, free, 2–2.5 mm. Fruit globose, 1.5– 尖峰西番莲 jian feng xi fan lian
1.8 cm in diam., densely pubescent. Fl. Feb–May, fr. Jun. Herbaceous vines, 8–10 m long. Stem terete, angular,
Forests on limestone hills; ca. 1200 m. W Guangxi (Jingxi), SE brownish pubescent. Leaves alternate; petiole 2–3 cm, glabres-
Yunnan (Xichou) [Vietnam]. cent, with 2 discoid glands at base; leaf blade 10–16 × 6–8.5
cm, leathery, abaxially glabrescent, with 2 lines of laminar nec-
6. Passiflora henryi Hemsley, Hooker’s Icon. Pl. 27: sub t.
taries scattered along midvein, base rounded, margin slightly
2623. 1899.
undulate, apex 2- or 3-lobed, lobes equal in length or central
圆叶西番莲 yuan ye xi fan lian lobe longer. Inflorescences sessile, 2-flowered. Pedicel 3–4 cm.
Flowers whitish, 4–5 cm in diam.; hypanthium ca. 1 cm wide.
Herbaceous vines, 2–3 m long. Stem angular, glabrous.
Sepals pinkish, ovate, 2.5–3 × ca. 1 cm, membranous, sparsely
Petiole 1.5–4 cm, with 2 tuberculate glands near base; leaf
pubescent. Petals white, 2.5–3 cm × ca. 5 mm. Coronal fila-
blade suborbicular to broadly orbicular, 3.5–5.5 × 3–6 cm,
ments in 2 series; outer series 1–2 cm, inner series ca. 5 mm;
leathery, abaxially with 4–6 nectaries, base rounded or sub-
operculum incurved, plicate, membranous; androgynophore ca.
cordate, margin entire, apex truncate, sometimes slightly acute.
1 cm tall. Stamens 5; filaments ca. 1 cm; anthers ca. 8 mm.
Inflorescences sessile, 2–8-flowered, glabrescent. Pedicel 1–2.5
Ovary ellipsoid, densely pubescent; styles 3, free, ca. 1 cm; stig-
cm. Flowers greenish yellow, 2–2.5 cm in diam.; hypanthium
mas capitate. Fruit ovoid or ellipsoid, ca. 3 cm in diam.
ca. 5 mm wide. Sepals green, 8–10 × 2–3 mm. Petals 7–8 × 1–2
mm. Coronal filaments in 2 filamentous series; outer series ● Dense forests. Guangxi (Guiping, Longmen), Hainan.
6–8 mm, center brown, apices whitish; inner series ca. 4 mm,
yellow-green, apices brown and capitate; operculum incurved, 9. Passiflora jugorum W. W. Smith, Notes Roy. Bot. Gard.
green, plicate, 1–2 mm tall, membranous; disk greenish brown, Edinburgh 9: 115. 1916.
ca. 0.3 mm high; androgynophore brown mottled, 4.5–5 mm 山峰西番莲 shan feng xi fan lian
tall. Stamens 5; filaments 4–5 mm, free. Ovary sessile on an-
drogynophore, subglobose, 1.5–2 × ca. 1.5 mm, pubescent; Passiflora burmanica Chakravarty.
styles 3, free, 4–5 mm. Fruit blue at maturity, globose, 1.2–1.5 Woody vines, ca. 8 m long. Stem terete, sparsely white pu-
cm in diam., glabrous. Fl. Jun–Jul, fr. Aug–Oct. bescent, striate. Petiole 3–5 cm, pubescent, with 2 peg-shaped
● Scrub on slopes and in valleys; 400–1600 m. Yunnan. glands in lower 1/3 part; leaf blade 3–11 × 3–10 cm, mem-
branous, pubescent, often with 2 or 3 pairs of glands near apex
7. Passiflora cupiformis Masters, Hooker’s Icon. Pl. 18: t.
of midvein on abaxial surface, apex truncate, (2 or)3-lobed;
1768. 1888.
lobes 5–10 mm, base broadly rounded to cordate, apex acute to
杯叶西番莲 bei ye xi fan lian acuminate. Inflorescences (sub)sessile, 5–15-flowered, pubes-
cent. Pedicel 2.5–3 cm, pubescent. Flowers whitish, 3–4 cm in
Passiflora franchetiana Hemsley; P. kwangsiensis H. L.
diam.; hypanthium 9–12 mm wide. Sepals light purple, 1–2 cm
Li; P. seguinii H. Léveillé & Vaniot; P. yunnanensis Franchet.
× ca. 5 mm, glabrous. Petals 1–1.5 cm × ca. 5 mm. Coronal
Climbing vines, ca. 6 m long. Stem terete, glabrescent. filaments in a single series, 6–8 mm; operculum plicate, 2.5–3
Petiole 3–7 cm, sparsely pubescent, with 2 discoid glands on mm tall, membranous; disk ca. 0.5 mm high; androgynophore
basal 1/4; leaf blade 6–12(–15) × 4–10 cm, leathery, abaxially 7–10 mm tall. Stamens 5; filaments 6–10 mm, basally fused for
with 6–25 scattered nectaries, base rounded to cordate, truncate 1.5–3(–4) mm; anthers 3.5–5 × 2–3 mm. Ovary sessile on an-
to deeply 2-lobed, lobes 3–8 cm, apically rounded or obtuse. drogynophore, ellipsoid, glabrous; styles 3, free, 4–5 mm. Fruit
Inflorescences (sub)sessile, (1–)5–20-flowered. Pedicel 2–3 cm. blue at maturity, subglobose to ellipsoid, ca. 2 cm in diam.,
Flowers creamy-white, 1.5–2 cm in diam.; hypanthium 5–7 mm glabrous. Fl. Mar–Apr, fr. May–Jul.
wide. Sepals 8–10 × 3–4 mm, often with apical appendage, gla- Forests on limestone hills; 1000–1800 m. S Yunnan [Myanmar].
brescent. Petals 7–8.5 × 2–3 mm. Coronal filaments in 2 series;
outer series 8–9 mm, filaments purple-brown mottled in proxi- 10. Passiflora wilsonii Hemsley, Bull. Misc. Inform. Kew 1908:
mal half, creamy-white in distal half; inner series 2–3 mm, base 17. 1908.
brown, apex white and capitate; operculum incurved, brownish
镰叶西番莲 lian ye xi fan lian
green, plicate, ca. 1.5 mm tall, membranous; disk purplish
brown, ca. 0.25 mm high; androgynophore brown mottled, 3–5 Passiflora assamica Chakravarty; P. celata G. Cusset; P.
 PASSIFLORACEAE 145

perpera Masters; P. rhombiformis S. Y. Bao; P. spirei G. ca. 5 mm, filaments white; inner series ca. 1 mm, filaments
Cusset. white, capitate at apex; operculum incurved, white, plicate, 1–
1.5 mm tall, membranous; disk green, ca. 0.2 mm high; andro-
Herbaceous vines, 6–10 m long. Stem terete, glabrous,
gynophore 2.5–3 mm tall. Stamens 5; filaments ca. 3 mm, free;
rarely glabrescent. Petiole 2–2.5 cm, with 2 peg-shaped glands
anthers ca. 2 × 1.5 mm. Ovary sessile on androgynophore,
in lower 1/3 part; leaf blade 4–6(–11) × (3.5–)6–13 cm, mem-
ovoid-ellipsoid, ca. 1.5 mm, glabrous; styles 3, free, 2.5–3 mm.
branous, glabrous, rarely glabrescent, with 1(or 2) pairs of glands
Fruit blue at maturity, globose, 1–1.5 cm in diam., glabrous. Fl.
near apex of midvein on abaxial surface, base broadly rounded
Apr–May.
to subcordate, apex strongly truncate, 2(or 3)-lobed. Inflores-
cences (sub)sessile, glabrous or sparsely puberulous, 2–15-flow- ● Forests in valleys; 600–1500 m. S Yunnan (Simao, Xishuang-
ered. Pedicel 5–15 mm. Flowers white, (2–)2.5 cm in diam.; banna).
hypanthium 5–10 mm wide. Sepals purplish brown, 7–11 × 2–3 13. Passiflora xishuangbannaensis Krosnick, Novon 15: 160.
mm, glabrous. Petals whitish, 7–10 × 2–3 mm. Corona in a 2005.
single series; filaments purple at base, white at apex, 3–6 mm;
operculum white, plicate, ca. 2 mm tall, membranous; disk 版纳西番莲 ban na xi fan lian
brown, 2–3 mm high; androgynophore 6–10 mm tall. Stamens
Vines, 1–3 m long. Stem subtriangular, glabrous. Petiole
5; filaments (2.5–)3–6 mm, free; anthers 2–3(–4) × 1–1.5(–2)
1.5–2.5 cm, with 2 nectaries in distal 1/4; leaf blade 4.5–8.5 ×
mm. Ovary sessile on androgynophore, ellipsoid, 2–3 mm, gla-
2.5–5 cm, abaxially white variegated along the three major
brous; styles 3, free, 3–5 mm. Fruit blue at maturity, subglo-
veins, abaxially with 8–12(–15) nectaries arranged in 2 lines
bose, 2.5–3 cm in diam., glabrous. Fr. Jul.
outward from midvein, margin entire, lateral lobes greatly ex-
● Scrub on slopes, along streams in mountain valleys, forest mar- ceeding midvein, creating a sinus ca. 1/2 as long as leaf. Inflo-
gins; 1300–2500 m. SW Guizhou (Wangmo), SE Xizang (Mêdog), C, rescence sessile, 1- or 2-flowered. Pedicel 1.5–2 cm. Flowers
E, and S Yunnan. 3.2–3.8 cm in diam.; hypanthium 8–10 mm wide. Sepals light
11. Passiflora papilio H. L. Li, J. Arnold Arbor. 24: 447. 1943. green, 1.6–1.8 cm × 5–7 mm. Petals white, 1–1.2 cm × 2–3
mm. Coronal filaments in 2 series; outer series ca. 1 cm, purple-
蝴蝶藤 hu die teng brown in basal 1/3, distal portion bright yellow; inner series ca.
5 mm, purple-brown, apex yellow and capitate; operculum in-
Herbaceous vines, 1–2 m long. Stem slender, glabrous or
curved, green, with purple striations, plicate, 3–4 mm high,
glabrescent, striate. Petiole 2–3 cm, sparsely pubescent, with 2
membranous; disk greenish purple, ca. 1 mm tall; androgyno-
conspicuous, auriculate glands in proximal 1/4; leaf blade 2.5–
phore ca. 7 mm tall. Stamens 5; filaments basally connate for
3.5 × 6–10 cm, leathery, glabrous, truncate-bilobed; lobes ovate,
ca. 1 mm; anthers 3–4 × ca. 1 mm. Ovary ellipsoid, 4–6 × 2–3
5–7 × 2.5–3.5 cm, abaxially with 4–8 discoid laminar nectaries,
mm, sessile on androgynophore, glabrous; styles 5–7 mm, free.
lateral veins 2 or 3 per side of midvein, base truncate or sub-
Fruit dark blue at maturity, ovoid, ca. 1 cm in diam.
rounded, apically acute or obtuse. Inflorescences sessile, 5–8-
flowered, sparsely pubescent. Pedicel 2–6 mm. Flowers yellow- ● Shaded areas near streams in forests and fields; ca. 1200 m. S
green, 1.5–2 cm in diam.; hypanthium ca. 4.5 mm wide. Sepals Yunnan (Xishuangbanna).
8–10 × 3–4 mm, villous. Petals 6–8 × 2–3 mm. Corona in 2 14. Passiflora suberosa Linnaeus, Sp. Pl. 2: 958. 1753.
series; outer series 3.5–4 mm, filaments green at base, yellow at
apex, apex thickened; inner series ca. 1.5 mm, filaments green 细柱西番莲 xi zhu xi fan lian
at base, apex yellow and capitate; operculum yellow-green, ca.
Herbaceous vines, 1–4(–10) m long. Stem white or whitish
1.5 mm tall; disk brownish, ca. 0.4 mm high; androgynophore
gray, triangular, slender, striate. Petiole 2–4 cm, white strigose,
4–5 mm tall. Stamens 5; filaments ca. 3 mm, free; anthers ob-
with 2 glands at middle or slightly above; leaf blade 5–5.5(–8)
long, ca. 2.5 × 1 mm. Ovary sessile on androgynophore, ovoid,
× 5–6(–11) cm, abaxially gray-green, sparsely villous, base cor-
ca. 2 × 1.5 mm, glabrous; styles 3, free. Fruit blue at maturity,
date, 3-lobed, lobes ovate, apex acute, mucronate. Flowers axil-
globose, 1–1.2 cm in diam. Fl. Apr–May, fr. Jun–Jul. lary, solitary or in pairs, light green or white, 1.5–2.5 cm in
● Scrub, forests on limestone hills; 400–500 m. SW Guangxi. diam. Sepals 5–8 mm, oblong or lanceolate, outside hispidu-
lous. Petals absent. Corona in 2 series, filamentous, outer series
12. Passiflora altebilobata Hemsley, Bull. Misc. Inform. Kew 3–7 mm, inner series ca. 1 mm; operculum plicate, 1–2 mm
1908: 17. 1908. high; disk 0.5–1 mm; androgynophore 2–4 mm tall. Filaments
月叶西番莲 yue ye xi fan lian flat, 1.5–4 mm, free; anthers oblong, 2–3 mm. Ovary subglo-
bose, glabrous; styles 3, almost threadlike, 7–8 mm; stigma
Herbaceous vines, ca. 2 m long. Stem striate, pubescent. capitate. Berry turning bluish black at maturity, subglobose, 1–
Petiole 1.5–2.5 cm, with 2 glands in basal 1/3 to middle; leaf 1.2 cm in diam. Fl. Aug–Sep, fr. Sep–Nov. 2n = 24.
blade 2.5–9 × 2–7 cm, abaxially with 4 small nectaries near
apex of midvein, deeply 2-lobed, pubescent, base rounded. In- Escaped weed. Taiwan, S Yunnan (Xishuangbanna) [native to the
florescences 2–16-flowered. Pedicel 1–5 mm, pubescent. Flow- West Indies and Central and South America].
ers greenish, 7–10 mm in diam.; hypanthium ca. 4 mm wide. This species was treated as Passiflora gracilis J. Jacquin ex Link
Sepals green, 3–6 × 3–4 mm, pubescent. Petals whitish, 3–6 × by Bao (in FRPS 52(1): 110. 1999). The two species are distinguished
2–3 mm, glabrous. Coronal filaments in two series; outer series by their leaf lobes in that they are obtuse or rounded in P. gracilis vs.
146 PASSIFLORACEAE

acute in P. suberosa; the petiolar glands are placed below the middle in Naturalized weed, grassy slopes, forest margins, roadsides, riv-
P. gracilis (occasionally with one additional gland above middle) vs. at erine beaches, occasionally cultivated; 100–1200 m. Guangdong, Guang-
or above the middle in P. suberosa; the fruit are bluish black and ovoid xi, Hainan, Taiwan, Yunnan [native to the West Indies and N South
or spheroid in P. suberosa vs. orange or red and ovoid and at least twice America].
as large in P. gracilis; the largest leaves of P. gracilis have two small
teeth less than 1 cm from the petiole vs. no such teeth in P. suberosa. The Chinese material is rather uniform, and Killip’s infraspecific
taxonomy (loc. cit. under genus heading) would key to Passiflora foe-
15. Passiflora adenopoda Candolle, Prodr. 3: 330. 1828. tida var. hispida (Candolle ex Triana & Planchon) Killip ex Gleason.

腺柄西番莲 xian bing xi fan lian 17. Passiflora edulis Sims, Bot. Mag. 45: t. 1989. 1818.
Passiflora acerifolia Chamisso & Schlechtendal; P. aspera 鸡蛋果 ji dan guo
Sessé y Lacasta & Mociño; P. scabra Sessé y Lacasta &
Passiflora minima Blanco (1837), not Linnaeus (1753).
Mociño.
Herbaceous vines, woody at base, ca. 6 m long. Stem
Herbaceous vines. Stem angulate, striate, hispid. Stipules
slender-striate, glabrous. Leaves 6–13 × 8–13 cm, membranous,
suborbicular to reniform, 1–1.3 × 1.7–2 cm; petiole 2.5–6 cm,
base cuneate or cordate, deeply 3-lobed, middle lobe ovate,
with 2 stipitate glands; leaf blade 7–12 × 7.5–15 cm, membra-
lateral lobes ovate-oblong, margin glandular-serrate, with 1 or 2
nous, hispidulous, 3- or 5-lobed, lobes ovate, base deeply cor-
small cup-shaped glands near base of sinuses, glabrous. Inflo-
date, margin dentate, apex acute. Inflorescence reduced to 1 or
rescence a reduced cyme, central flower not developed, one lat-
2 flowers; bracts 3, ovate-oblong, lacerate, placed at or slightly
eral branch converted to a tendril, flower opposite tendril; bracts
above middle. Flowers white, with violet purple center, 3–5 cm
green, broadly ovate or rhombic, 1–1.2 cm, margin irregularly
in diam. Sepals oblong-lanceolate, 1.8–2.7 × 1–1.5 cm, obtuse,
serrulate. Pedicel 4–4.5 cm, biglandular at apex. Flowers 4–7
awned. Petals linear-lanceolate, 8–9 × 4–5 mm, apex obtuse.
cm in diam.; hypanthium 0.8–1 × 1–1.2 cm. Sepals green out-
Corona uniseriate, ca. 1.8 cm, base violet-purple, apex white;
side, light green or white inside, 2.5–4 × ca. 1.5 cm, awn 2–4
operculum incurved, white, plicate; limen annular; androgyno-
phore ca. 9 mm tall. Ovary ovoid, densely tomentose. Fruit blu- mm. Petals 2.5–3 cm × ca. 8 mm. Corona in 4 or 5 series; outer
ish black, subglobose, ca. 3.5 × 3 cm; aril bright orange. Seeds 2 series ligulate with filiform distal half, 2–2.5 cm, base light
brown to black, ca. 6 × 4 mm, reticulate. 2n = 12. green, middle purple, apex white; inner 2 or 3 series filiform, 1–
3 mm, green and purple; operculum recurved, 1–1.2 mm, mar-
Naturalized weed, grassy slopes on limestone hills. SW Yunnan
gin entire or irregularly lacerate apically; disk ca. 4 mm high,
(Menglian) [native to Central and South America (Mexico to Peru)].
membranous; androgynophore 1–1.2 cm tall; trochlea (ring-
16. Passiflora foetida Linnaeus, Sp. Pl. 2: 959. 1753. shaped enlargement on androgynophore) just above disk. Fila-
ments 5–6 mm, flat, coherent at base; anthers light yellow-green,
龙珠果 long zhu guo
oblong, 5–6 mm. Ovary obovoid, ca. 8 mm, glabrous to pubes-
Dysosmia foetida (Linnaeus) M. Roemer; Granadilla cent; styles flat; stigma reniform. Fruit purple at maturity, ovoid,
foetida (Linnaeus) Gaertner; Passiflora foetida var. hispida 3–4 cm in diam., glabrous. Seeds many, ovoid, 5–6 mm. Fl.
(Candolle ex Triana & Planchon) Killip; P. hispida Candolle ex Jun, fr. Nov.
Triana & Planchon; Tripsilina foetida (Linnaeus) Rafinesque. Cultivated, escaped in forests in mountain valleys; 100–1900 m.
Herbaceous vines, foul smelling. Stem slender, striate, Fujian, Guangdong, Taiwan, Yunnan [native to South America (probab-
ly originally from S Brazil)].
spreading pubescent. Stipules partly clasping, deeply parted,
glandular pilose; petiole 2–6 cm, without glands, pubescent, in- 18. Passiflora caerulea Linnaeus, Sp. Pl. 2: 959. 1753.
dumentum dense, spreading, glandular pilose; leaf blade broad-
西番莲 xi fan lian
ly ovate to oblong-ovate, (3–)4.5–13 × (2.5–)4–12 cm, mem-
branous, glandular ciliate, abaxially pubescent with small nec- Passiflora caerulea Loureiro ex Candolle (1828), not
taries in distal part, adaxially sericeous, mixed with occasional Linnaeus (1753); P. loureiroi G. Don.
glandular hairs, base cordate, 3-lobed or unlobed, margin irreg- Herbaceous vines. Stem terete, slightly angulate, glabrous.
ularly undulate, apex acute. Inflorescence reduced to a single Stipules reniform, ca. 1.2 cm, clasping, margin undulate; petiole
flower, opposite tendril; bracts 3, 2- or 3(or 4)-pinnatifid; lobes 2–3 cm, with 2–4(–6) small glands; leaf blade 5–7 × 6–8 cm,
threadlike, glandular pilose apically. Flowers white or light pur- papery, base cordate, margin entire, palmately (3–)5(–9)-lobed;
ple with white spots or lines, 2–3 cm in diam. Sepals ca. 1.5 cm, lobes ovate-oblong, middle lobe slightly larger than lateral lobes,
awned. Petals ca. 1.5 cm. Corona 3–5-seriate, filamentous, two glabrous. Inflorescence a reduced cyme, central flower not devel-
outer series ca. 1 cm, inner three series 1–2.5 mm; operculum oped, one lateral branch converted to a tendril, flower opposite
1–1.5 mm, membranous; disk cupular, 1–2 mm high; andro- tendril; bracts broadly ovate, 2–3 cm, margin entire. Pedicel 3–
gynophore 5–7 mm tall. Stamens coherent at base, flat; anthers 4 cm. Flowers 6–8(–10) cm in diam. Sepals light green outside,
oblong, ca. 4 mm. Ovary shortly stipitate, ellipsoid, ca. 6 mm, white inside, 3–4.5 cm, awn 2–3 mm. Petals white to light green,
glabrous; styles 3(or 4), 5–6 mm; stigmas capitate. Berry orange 2.5–4 cm. Corona in 3 or 4 series, filamentous; outer 2 series
or orange-red, ovoid-globose, 2–3 cm in diam., glabrous. Seeds (0.6–)1–1.5 cm, base dark purple, middle white, apex bright
many, light brown to black, elliptic, 3–4 mm. Fl. Jul–Aug, fr. blue; inner 1 or 2 series 1–2 mm, base light green, apex white
Apr–May of next year. 2n = 20. and capitate; operculum fimbricate, lobes dark purple, with an-
 PASSIFLORACEAE 147

nular nectary at base; disk 1–2 mm high; androgynophore 8–10 styles purple; stigma 3-lobed. Berry yellow or orange at ma-
mm tall. Filaments ca. 1 cm, flat, free; anthers oblong, ca. 1.3 turity, ovoid, 20–25 cm, rind thick and fleshy. Seeds many, 7–9
cm. Ovary ovoid-globose; styles free, purple, 6–8 mm; stigma mm, flat. Fl. Feb–Aug.
reniform. Berry orange-yellow or yellow, ovoid-globose or sub-
Cultivated. Guangdong, Guangxi, Hainan [native to tropical Ameri-
globose, ca. 6 cm. Seeds many, obcordate, ca. 5 mm. Fl. May–
ca].
Jul.
20. Passiflora laurifolia Linnaeus, Sp. Pl. 2: 956. 1753.
Cultivated. Guangxi, Jiangxi, Sichuan, Yunnan [native to South
America (N Argentina, S Brazil)]. 樟叶西番莲 zhang ye xi fan lian
19. Passiflora quadrangularis Linnaeus, Syst. Nat., ed. 10, 2: Passiflora acuminata Candolle; P. tinifolia Jussieu.
1248. 1759.
Woody vines. Stem terete, glabrous, striate. Stipules line-
大果西番莲 da guo xi fan lian ar, 3–4 mm; petiole terete, ca. 1.2 cm, sparsely pubescent, with
Granadilla quadrangularis (Linnaeus) Medikus. 2 glands near base of blade; leaf blade ovate-oblong, 7–10 ×
3.5–6 cm, leathery, glabrous, lateral veins 8–10 per side of mid-
Herbaceous vines, woody at base, 10–15 m long, glabrous. vein, base rounded or subcordate, margin entire, apex mucronu-
Young stem quadrangular, narrowly winged. Stipules ovate-lan- late. Inflorescence reduced to a single flower; bracts 3, ovate,
ceolate, 2–4 cm, margin entire, apex acute; petiole 3–5 cm, with 2–3 cm, puberulous, base cuneate, margin glandular serrate, apex
2 or 3 pairs of cupular glands; leaf blade broadly ovate to sub- obtuse. Pedicel 1.5–3 cm, thick. Flower white, with red spots,
orbicular, 7–13 × 5–15 cm, membranous, glabrous, lateral veins 5–7 cm in diam. Sepals 5, ovate-oblong, 2–2.5 cm, awned and
8–12 pairs, tertiary veins somewhat prominent, base rounded puberulous apically. Petals 5, similar to sepals. Outer corona in
to truncate, margin entire, apex acute. Inflorescence a reduced 6 whorls, purple with white bands; outer 2 whorls 2–4 cm, line-
cyme, flower opposite tendril, tendril thick and strong; bracts ar; inner 4 whorls 1.5–2 mm, threadlike; inner corona membra-
leaflike, ovate, 3–5 cm, base cordate, apex acute. Pedicel 1–3 nous, apically dentate; limen annular around base of androgyno-
cm, triangular, articulate at middle. Flowers light red, 6–8(–10) phore. Ovary ovoid, sparsely pubescent. Fruit citreous or red-
cm in diam. Sepals green outside, red inside, ovate to ovate- dish yellow, ovoid, 4–6 cm. Seeds many, obcordate, 5–7 mm. Fl.
oblong, 3–4 cm, sparsely pubescent. Petals light red, oblong to Jun.
oblong-lanceolate, 2.5–3.5 cm. Corona in 5 series, white or pur-
ple, filamentous; outer 2 series 3–5 cm; middle series ca. 2 mm, Cultivated. Guangdong (Guangzhou) [native to Central and South
capitate; inner 2 series 1–2 mm, threadlike, 4–5 mm, membra- America].
nous, margin lacerate; operculum membranous, similar to fifth A white-flowered form was introduced to China; wild populations
outer corona in shape; disk cupular. Stamens free. Ovary ovoid; most often have red or reddish purple flowers.

2. ADENIA Forsskål, Fl. Aegypt.-Arab. 77. 1775.

蒴莲属 shuo lian shu
Wang Yinzheng (王印政); David Hearn
Modecca Lamarck.
Plants herbaceous or woody vines, perennial, tendrillate, glabrous. Roots tuberous or fleshy. Stems terete; tendrils axillary,
(minutely 3-forked or) simple. Leaves simple, alternate, petiolate; stipules triangular, minute; petiole apex or blade base with 2
glands. Plants (monoecious or) dioecious. Inflorescences axillary, cymose-corymbose, pedunculate, few- to many flowered, tendril-
late at center; bracts and bracteoles triangular, minute. Flowers yellowish or green, often red punctate, unisexual, with solid tissue
(stipe) subtending hypanthium articulate at pedicel. Hypanthium narrowly to broadly cup-shaped. Sepals (calyx lobes) 5, partially
connate into calyx tube, imbricate, persistent. Petals 5, free, attached at hypanthium apex, greenish, creamy, or white, mostly smaller
than sepals, included. Septa, when present, adnate with hypanthium and filaments or staminodes, dividing hypanthium into 5 sacs.
Corona absent or a laciniate to membranous ring at apex of hypanthium. Disk glands 5 or absent, ligulate, truncate, inserted at or near
base of hypanthium, alternating with petals. Male flower: stamens 5, hypogynous or perigynous, variably inserted in hypanthium;
filaments free or partially connate into tube, often fused abaxially to septa; vestigial ovary present. Female flower with stamens
reduced to subulate staminodes; ovary superior on gynophore, globose, with 3 parietal placentas; remnant stamen tube, if present, not
adnate with gynophore; ovules numerous, anatropous; styles 3, free or partially fused; stigmas laciniate or plumose to densely
woolly. Fruit a 3-valved capsule, bright red. Seeds with pitted testa, black at maturity, enclosed in fleshy aril.
About 100 species: Old World tropics and subtropics; three species in China.

1a. Plants herbaceous vines, with tuberous rootstock; mature stems at soil-level narrow, less than 1 cm in diam.;
leaf with paired glands on peltate extension of blade base; flowers with calyx lobes 2–10 × as long as depth
of hypanthium, hypanthium soft; corona usually present ......................................................................................... 1. A. penangiana
148 PASSIFLORACEAE

1b. Plants woody lianas, with fleshy rootstock, rarely tuberous; mature stems at soil-level thick, 2–8 cm or more
in diam.; leaf with paired glands on auriculate extension at apex of petiole; flower with calyx lobes 0.2–1 ×
as long as depth of hypanthium; hypanthium fleshy, turgid, thick; corona absent.
2a. Margins of mature leaves with irregularly spaced black-tipped glandular serrations, bases cordate; tertiary
and secondary veins raised abaxially, distinctly trabeculate, veins from arcuate laterals parallel; exocarp
hard-woody, mesocarp spongy, together 5–20 mm thick; elevation (800–)1000–2000 m ............................. 2. A. cardiophylla
2b. Margins of mature leaves entire or rarely serrate toward apex, bases variable, typically rounded; tertiary
veins typically indistinct, veins from arcuate laterals with variable orientations; pericarp flexible-leathery
externally and internally, 1–3 mm thick; elevation near sea level to 300(–1000) m ...................................... 3. A. heterophylla

1. Adenia penangiana (Wallich ex G. Don) de Wilde, Blumea Modecca cardiophylla Masters in J. D. Hooker, Fl. Brit.
15: 266. 1967. India 2: 602. 1879.
滇南蒴莲 dian nan shuo lian Canopy lianas, to 25 m long. Stems terete, to 8 cm in
diam.; internodes 5–15 cm; tendrils to 20 cm. Stipules dark-
Passiflora penangiana Wallich ex G. Don, Gen. Hist. 3: ening with age, narrowly triangular, ca. 1 × 0.5 mm; petiole 3–
55. 1834; Adenia nicobarica King; Disemma penangiana 12 cm, glands at blade-petiole junction 2, on concave ca. 4 mm
(Wallich ex G. Don) Miquel; Modecca nicobarica Kurz. in diam. auriculate extensions at apex of petiole, 2–3 mm in
Vines, to 6 m long, glabrous. Rootstock tuberous. Stems diam.; leaf blade cordate, rarely (2 or)3-lobed, 7–25 × 5–19 cm,
terete, slender; internodes 1–10 cm; tendrils simple or 3-forked, membranous, not punctate, tertiary and secondary veins raised
1–10 cm. Stipules triangular, ca. 0.5 mm; petioles 0.5–2 cm; abaxially, venules distinctly trabeculate, veins from arcuate lat-
glands at blade-petiole junction 2, on slightly bi-lobed peltate erals parallel, base of mature leaves cordate, margins of mature
extension of base of blade, ca. 1 mm in diam.; leaf blade elliptic leaves with irregularly spaced black-tipped glandular serrations,
apex acute to acuminate, with mucro. Plants dioecious. Inflo-
to lanceolate, 2–16 × 0.5–4 cm, membranous to leathery, not
rescences with tendril emerging from center, pedunculate to 18
punctate, glands near leaf margin 0–9, base rounded, margin
cm, to 30-flowered in males, 1–3-flowered in females; bracts
entire, apex acute to acuminate. Plants (monoecious or) dioeci-
and bracteoles triangular, ca. 1 mm, acute. Pedicel 0.5–10 mm.
ous. Inflorescences usually tendrillate, occasionally of short
Flowers tubiform-campanulate, 9–25 × 3–8 mm, including stipe
racemose shoots, pedunculate for 1–10 cm, to 30-flowered in 6–12 mm. Hypanthium cupuliform, wall fleshy-leathery, ca.
males, 1–3-flowered in females; bracts and bracteoles triangu- 0.75 mm thick, 6–10 mm deep. Calyx tube 0.5–3 mm; lobes
lar, ca. 1 mm, apex acute. Pedicel 0.5–4 mm. Flowers narrowly narrowly triangular, 4–6 mm, rounded to acute, entire. Petals
campanulate, 6–17 × 1–4 mm; stipe cream-colored, 1–7 mm. deltoid to lanceolate, 3–5 × ca. 2.5 mm, red punctate, rounded
Hypanthium narrowly cup-shaped, 1–2.5 mm deep. Calyx tube to acute, entire. Septa to 2 mm high. Corona absent. Disk glands
to 2 mm; lobes reflexed, oblong to linear, 4–15 mm, apex entire linear, to 1 mm high, truncate. Male flowers: filaments 0.5–
to irregular, rounded to acute. Petals inserted at level of corona, 5.5 mm, connate up to 4 mm, inserted at base of hypanthium;
elliptic to linear, 4–10 mm, apex serrulate, rounded to acute. anthers ca. 5 × 1.5 mm, acute, up to 0.5 mm apiculate; vestigial
Septa 0.25–2 mm high. Corona laciniate, membranous, occa- ovary to 3 mm, gynophore ca. 1 mm. Female flowers: stami-
sionally of woolly filaments at apex of hypanthium. Disk nodes 2–3 mm, connate for ca. 1 mm, inserted at base of hy-
glands linear, ca. 1 mm, truncate. Male flowers: filaments 2–3 panthium; ovary ellipsoid, ca. 4 × 4 mm; gynophore to 4 mm;
mm, connate for ca. 2 mm, inserted at base of hypanthium, style arms ca. 1 mm, connate for ca. 1 mm; stigmas laciniate.
septa occasionally spanning entire length of hypanthium; an- Capsules 1 or 2 per inflorescence, outside deep red at dehis-
thers ca. 3 × 0.5 mm, to 1 mm apiculate; vestigial ovary includ- cence, pear-shaped, fruit to 11 cm with gynophore to 7.5 cm,
ing gynophore ca. 1 mm. Female flowers: staminodes ca. 1 mm, exocarp woody, mesocarp and endocarp spongy. Seeds to 40
connate for ca. 0.5 mm, inserted at base of hypanthium; ovary per capsule, reniform; funicles of seeds 1–2 cm. Fl. Apr–Sep,
3–6 mm; gynophore ca. 0.5 mm; styles 1–2 mm, connate for up fr. Jun–Nov.
to 0.5 mm; stigmas globose, woolly. Capsules 1 per inflores- Mixed forests, scrub; (800–)1000–2000 m. Yunnan [?Bhutan,
cence, scarlet at dehiscence, oblong, ca. 4 × 2 cm, with gyno- Cambodia, India, Laos, Malaysia, Myanmar, Thailand, Vietnam].
phore to 4 cm, exocarp and mesocarp leathery. Seeds ca. 10 per 3. Adenia heterophylla (Blume) Koorders, Exkurs.-Fl. Java 2:
capsule, subglobose or flattened; funicles of seeds 5–10 mm. Fl. 637. 1912.
and fr. throughout year, mostly around Oct.
异叶蒴莲 yi ye shuo lian
Dense forests and open fields, occasionally in seasonally marshy
areas; ca. 500 m. Yunnan (Mengla) [India (Nicobar Islands), Indonesia Modecca heterophylla Blume, Bijdr. 940. 1826; Adenia
(Sumatra), Laos, Malaysia, Thailand, Vietnam]. chevalieri Gagnepain; A. maclurei Merrill; A. parviflora
(Blanco) Cusset; A. populifolia K. Schumann & Lauterbach
The above description is based on extra-regional specimens as no
material from China was available for study.
(1900), not Engler (1891); Modecca formosana Hayata; Passi-
flora parviflora Blanco.
2. Adenia cardiophylla (Masters) Engler, Bot. Jahrb. Syst. 14:
Canopy lianas, to 30 m long. Roots fleshy, thickened.
376. 1891.
Stems terete, to 5 cm in diam.; internodes 2–20 cm; tendrils (3-
三开瓢 san kai piao forked or) simple, to 25 cm. Stipules flattened-triangular, ca. 1
 PASSIFLORACEAE 149

mm; petiole 1–10 cm, glands at blade-petiole junction 2, on flowers: filaments 1–4 mm, connate up to 2 mm, inserted at
concave 2–4 mm in diam. auriculate extensions at apex of peti- base of hypanthium; anthers ca. 3 × 1 mm, acute, up to 0.5 mm
ole in mature leaves, 1–3 mm in diam.; leaf blade ovate to lan- apiculate; vestigial ovary ca. 1 mm, gynophore to 1 mm. Fe-
ceolate, membranous to leathery, glands often present, dotlike, male flowers: staminodes 1–3 mm, connate to 1 mm, inserted at
base of mature leaf oblong, unlobed or (2 or)3-lobed with distal base of hypanthium; ovary subglobose to oblong, ca. 4 × 2.5
1/3 of leaf often slightly constricted from slight lobation, mar- mm; gynophore ca. 3 mm; style arms ca. 1 mm, free or connate
gin entire to serrate, apex acute to acuminate. Plants (monoeci- for ca. 0.5 mm; stigmas globose, papillate. Capsules 1–3 per in-
ous or) dioecious. Inflorescences often with tendril emerging florescence, outside deep red at dehiscence, ellipsoid, fruit body
from center, peduncle to 20 cm, to 40-flowered in males, 2–4- to 13 cm, with gynophore to 3 cm, fruit wall leathery. Seeds to
flowered in females; bracts and bracteoles narrowly triangular, 60 per capsule, orbicular; funicles of seeds 5–15 mm. Fl. and fr.
ca. 1 mm, apex acute. Pedicel 0.5–13 mm. Flowers tubiform- throughout year, primarily during wet periods.
urceolate, 15–25 × 2–5 mm, including stipe 1–15 mm. Hypan- Forests, scrub; near sea level to 300(–1000) m. Guangdong,
thium urceolate to cupuliform, wall fleshy-leathery, ca. 0.5 mm Guangxi, Hainan, Taiwan [Cambodia, Indonesia (Java, Sumatra), Laos,
thick, 3–6 mm deep. Calyx tube 2–3 mm; lobes reflexed, tri- New Guinea, Philippines, Thailand, Vietnam; Australia, Pacific islands].
angular, 1–3 mm, rounded to acute, entire. Petals oblanceolate This is a widespread and highly variable species. It has been treat-
to triangular, 2–4 × 0.5–1 mm, red punctate, base hastate, apex ed as Adenia heterophylla [subsp. heterophylla] var. heterophylla accord-
entire to undulate, rounded to acute. Septa 0.25–3 mm high. ing to de Wilde (Meded. Landbouwhogeschool Wageningen 71(18):
Corona absent. Disk glands linear, to 3 mm high, truncate. Male 216. 1971).