Taxonomy and classification
A revision of the infrageneric classifications suggested by previous authors on the basis of morphological studies (Yueh and Cheng 1974; Jeffrey 1980b; Huang et al. 1997; Rugayah and de Wilde 1999; de Wilde and Duyfjes 2004) is here proposed on the basis of the molecular phylogenetic data (de Boer et al. Submitted). A synopsis is presented in which we attempt to assign all 91 species recognized here to sections using the clades recovered in the phylogenetic analysis as a framework, along with a plenitude of data from macromorphological studies of herbarium vouchers (Chakravarty 1959; Jeffrey 1980a; Jeffrey 1980b; Jeffrey 1982; Telford 1982; Ohba 1984; Rugayah and de Wilde 1997; Rugayah and de Wilde 1999; Duyfjes and Pruesapan 2004; de Wilde and Duyfjes 2004; de Wilde and Duyfjes 2006; Cooper and Ford 2010; de Wilde and Duyfjes 2010; Lu et al. 2011; Cooper and de Boer 2011) and palynological work (Huang et al. 1997; Pruesapan and Van Der Ham 2005; van der Ham et al. 2010). Synonyms are only included if these are new or relevant for this paper. Names that have been placed in synonymy by previous authors can be found in the above-cited morphological studies.
Trichosanthes L. (1753) Sp. Pl. 2: 1008 – Type: Trichosanthes anguina L. [= Trichosanthes cucumerina L.]
Trichosanthes subg. Scotanthus (Kurz) H.J.de Boer, comb. nov. – Gymnopetalum subg. Scotanthus Kurz (1877) J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 46: 99. – Lectotype, designated here: Bryonia cochinchinensis Lour. [ = Trichosanthes costata Blume]
Scotanthus Naud., nom. illeg. (1862) Ann. Sci. Nat., Bot. sér. 4, 16: 172. – Type: Momordica tubiflora Roxb. [ = Trichosanthes costata Blume]
Trichosanthes sect. EdulisRugayah (1999) Reinwardtia 11: 232. – Type: Trichosanthes edulis Ruguyah.
Trichosanthes densiflora Rugayah (1999) Reinwardtia 11: 252
Trichosanthes dentifera Rugayah (1999) Reinwardtia 11: 253
Trichosanthes dieniensis Merr. & L.M.Perry (1949) J. Arnold Arbor. 30: 59
Trichosanthes edulis Rugayah (1999) Reinwardtia 11: 254
Trichosanthes hastata Harms (1925) Bot. Jahrb. Syst. 60: 160
Trichosanthes laeoica C.Y.Cheng & Lu Q.Huang (1996) Bull. Bot. Res., Harbin 16: 503
Trichosanthes odontosperma W.E.Cooper & A.J.Ford (2010) Austrobaileya 8: 126
Trichosanthes pulleana Harms (1925) Bot. Jahrb. Syst. 60: 160
Trichosanthes schlechteri Harms (1925) Bot. Jahrb. Syst. 60: 159
Trichosanthes sect. Involucraria(Ser.) Wight (1840) Madras J. Lit. Sci. 12: 52. – Involucraria Ser. (1825) Mém. Soc. Phys. Genève 3: 27, t. 5. – Type: Involucraria wallichiana Ser. [ = Trichosanthes wallichiana (Ser.) Wight]
Trichosanthes anamalaiensis Bedd. (1864) Madras J. Lit. Sci. III, 1: 47
Trichosanthes borneensis Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 369
Trichosanthes bracteata (Lam.) Voigt (1845) Hort. Suburb. Calcutt. 58
Trichosanthes celebica Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 385
Trichosanthes cordata Roxb. (1832) Fl. Ind. 3: 703
Trichosanthes coriacea Blume (1826) Bijdr. Fl. Ned. Ind. 15: 935
Trichosanthes dolichosperma Duyfjes & Pruesapan (2004) Thai Forest Bull., Bot. 32: 84
Trichosanthes dunnianaH. Lév. (1911) Repert. Spec. Nov. Regni Veg. 10: 148
Trichosanthes ellipsoidea Merr. (1918) Philipp. J. Sci., C 13: 332
Trichosanthes elmeri Merr. (1929) Univ. Calif. Publ. Bot. 15: 299
Trichosanthes emarginata Rugayah (1999) Reinwardtia 11: 258
Trichosanthes erosaDuyfjes & Pruesapan (2004) Thai Forest Bull., Bot. 32: 85
Trichosanthes fissibracteata C.Y.Wu ex C.Y.Cheng & C.H.Yueh (1974) Acta Phytotax. Sin. 12: 438
Trichosanthes floresana Rugayah (1999) Reinwardtia 11: 260
Trichosanthes globosa Blume (1826) Bijdr. Fl. Ned. Ind. 15: 936
Trichosanthes intermedia W.J.de Wilde & Duyfjes (2004) Sandakania 14: 19
Trichosanthes inthanonensis Duyfjes & Pruesapan (2004) Thai Forest Bull., Bot. 32: 86
Trichosanthes khasiana Kundu (1939) J. Bot. 77: 11
Trichosanthes kinabaluensis Rugayah (2000) Reinwardtia 11: 419
Trichosanthes kostermansii Duyfjes & Pruesapan (2004) Thai Forest Bull., Bot. 32: 89
Trichosanthes laceribractea Hayata (1911) J. Coll. Sci. Imp. Univ. Tokyo 30. Art. 1: 117
Trichosanthes lepiniana (Naud.) Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 377
Trichosanthes leuserensis Rugayah (1999) Reinwardtia 11: 265
Trichosanthes longispicata Rugayah (1999) Reinwardtia 11: 266
Trichosanthes montana Rugayah (1998) Reinwardtia 11: 218
Trichosanthes morrisii W.E.Cooper (2011) Austrobaileya 8: 381
Trichosanthes obscura Rugayah (1999) Reinwardtia 11: 269
Trichosanthes pallidaDuyfjes & Pruesapan (2004) Thai Forest Bull., Bot. 32: 90
Trichosanthes papuana F.M.Bailey (1900) Queensland Agric. J. 7: 349
Trichosanthes pedata Merr. & Chun (1934) Sunyatsenia 2: 20
Trichosanthes pentaphylla F.Muell. in Benth. (1867) Fl. Austral. 3: 314
Trichosanthes philippinensis Rugayah (1999) Reinwardtia 11: 271
Trichosanthes planiglans Rugayah (1999) Reinwardtia 11: 273
Trichosanthes pubera Blume (1826) Bijdr. Fl. Ned. Ind. 15: 936
Trichosanthes quinquangulata A. Gray (1854) U.S. Expl. Exped., Phan. 15: 645
Trichosanthes quinquefolia C.Y.Wu ex C.Y.Cheng & C.H.Yueh (1980) Acta Phytotax. Sin. 18: 351
Trichosanthes refracta C.H.Yueh (1996) Bull. Bot. Res., Harbin 10: 500
Trichosanthes rugatisemina C.Y.Cheng & C.H.Yueh (1974) Acta Phytotax. Sin. 12: 440
Trichosanthes sepilokensis Rugayah (1999) Reinwardtia 11: 275
Trichosanthes subrosea C.Y.Cheng & C.H.Yueh (1980) Acta Phytotax. Sin. 18: 349
Trichosanthes tricuspidata Lour. (1790) Fl. Cochinch. 2: 589
Trichosanthes valida Rugayah (1999) Reinwardtia 11: 277
Trichosanthes wallichiana (Ser.) Wight (1840) Madras J. Lit. Sci. 12: 52
Trichosanthes wawrae Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 384
Trichosanthes sect.Tripodanthera (M.Roem.) H.J.de Boer, comb. nov. – Tripodanthera M.Roem. (1846) Fam. Nat. Syn. Monogr. 2: 48. – Gymnopetalum sect. Tripodanthera (M.Roem.) Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 390. – Type: Bryonia cochinchinensis Lour. [ = Trichosanthes costata Blume]
Trichosanthes costata Blume (1826) Bijdr. Fl. Ned. Ind. 15: 933. — Type: Blume s.n. barcode L0589632, (lectotype L, designated by de Wilde and Duyfjes (2006); 2 isotypes L), Java, Indonesia. Heterotypic synonyms: Evonymus chinensis Lour. (1790) Fl. Cochinch. 1: 156. – Gymnopetalum chinense (Lour.) Merr. (1919) Philipp. J. Sci. 15: 256 – Type: Loureiro †. Bryonia cochinchinensis Lour. (1790) Fl. Cochinch. 1: 595. – Gymnopetalum cochinchinense (Lour.) Kurz (1871) J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 40: 57 –Type: Loureiro s.n. (BM), Vietnam. Momordica tubiflora Roxb. (1832) Fl. Ind. 3: 711 – Type: Wallich Cat. 6749 (holotype? K-W), Dacca, Bangladesh. Note: The existence of Trichosanthes chinensis Ser. (1828) Prodr. [A.P. de Candolle] 3: 315 blocks the transfer of Gymnopetalum chinense, based on the basionym Evonymus chinensis, to Trichosanthes. The second name in line of priority would be Gymnopetalum cochinchinensis, based on the basionym Bryonia cochinchinensis. However, the combination Trichosanthes cochinchinensis M.Roem. (1846) Fam. Nat. Syn. Monogr. 2: 96, based on Trichosanthes cucumerina Lour. (1790) 722 (non L.), blocks the transfer. The third name in line of priority is Trichosanthes costata Blume (1826) Bijdr. Fl. Ned. Ind. 15: 933, and this name is available for Gymnopetalum chinense.
Trichosanthes scabra Lour. (1790) Fl. Cochinch. 2: 589. – Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes (2008) Reinwardtia 12: 268. – Type: Poilane 11322 (neotype P; isoneotype L, designated by de Wilde and Duyfjes (2008)), Annam. Heterotypic synonym: Cucumis integrifolius (‘integrifolia’) Roxb. (1832) Fl. Ind. 3: 724. – Gymnopetalum integrifolium (Roxb.) Kurz (1871) J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 40: 58. –Type: Wallich Cat. 6730 (holotype? K-W), Burma.
var. scabra
var. pectinata (W.J.de Wilde & Duyfjes) H.J.de Boer, comb. nov. – Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes var. pectinatum (W.J. de Wilde & Duyfjes) W.J.de Wilde & Duyfjes (2008) Reinwardtia 12: 268 – Gymnopetalum integrifolium (Roxb.) Kurz var. pectinatum W.J.de Wilde & Duyfjes (2006), Blumea 51: 287. – Type: W.J. de Wilde and Duyfjes 21692 (holotype L), Java, Indonesia.
var. penicaudii (Gagnep.) H.J.de Boer, comb. nov. – Gymnopetalum penicaudii Gagnep. (1918) Bull. Mus. Natl. Hist. Nat. 24: 374. – Gymnopetalum scabrum (Lour.) W.J.de Wilde & Duyfjes var. penicaudii (Gagnep.) W.J.de Wilde & Duyfjes (2008) Reinwardtia 12: 268 – Type: Pénicaud 43 (lectotype P), Hainan, China.
Trichosanthes orientalis (W.J.de Wilde & Duyfjes) H.J.de Boer, comb. nov. – Gymnopetalum orientaleW.J.de Wilde & Duyfjes (2006) Blumea 51: 290. – Type: De Wilde and Duyfjes 21937 (holotype L), Lombok, Indonesia.
Trichosanthes subg. Trichosanthes
Unresolved placement within this subgenus:
Trichosanthes reticulinervis C.Y.Wu ex S.K.Chen (1985) Bull. Bot. Res., Harbin 5(2): 114
Trichosanthes smilacifolia C.Y.Wu ex C.H.Yueh & C.Y.Cheng (1980) Acta Phytotax. Sin. 18: 347
Trichosanthes sect. Asterosperma W.J.de Wilde & Duyfjes (2004) Sandakania 14: 6. – Type: Trichosanthes postarii W.J.de Wilde & Duyfjes.
Trichosanthes auriculata Rugayah (1998) Reinwardtia 11: 216
Trichosanthes fusca W.J.de Wilde & Duyfjes (2004) Sandakania 14: 17
Trichosanthes postarii W.J.de Wilde & Duyfjes (2004) Sandakania 14: 26
Trichosanthes rotundifolia Rugayah (1998) Reinwardtia 11: 223
Trichosanthes sect. Cucumeroides (Gaertn.) Kitam. (1943) J. Jap. Bot. 19: 35. – Cucumeroides Gaertn. (1791) Fruct. Sem. Pl. 2: 485, t. 180, t. 4. – Type: Trichosanthes cucumeroides (Ser.) Maxim. [ = Trichosanthes pilosa Lour.].
Trichosanthes adhaerens W.J.de Wilde & Duyfjes (2004) Sandakania 14: 11
Trichosanthes beccariana Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 380
Trichosanthes mucronata Rugayah (1999) Reinwardtia 11: 268
Trichosanthes pendula Rugayah (1998) Reinwardtia 11 (3): 219
Trichosanthes pilosa Lour. (1790) Fl. Cochinch. 2: 588. Heterotypic synonyms according to Cooper and de Boer (2011): Trichosanthes baviensis Gagnep. (1918) Bull. Mus. Natl. Hist. Nat. 24: 379; Trichosanthes trichocarpa C.Y.Wu ex C.Y.Cheng & C.H.Yueh (1980) Acta Phytotax. Sin. 18: 340; Trichosanthes holtzei F.Muell. (1886) Australas. Journ. Pharm. 1: 447.
Trichosanthes siamensis Duyfjes & Pruesapan (2004) Thai Forest Bull., Bot. 32: 97
Trichosanthes tetragonosperma C.Y.Cheng & C.H.Yueh (1974) Acta Phytotax. Sin. 12: 425
Trichosanthes sect. Foliobracteola C.Y.Cheng & C.H.Yueh (1974) Acta Phytotax. Sin. 12: 427. – Type: Trichosanthes kirilowii Maxim.
Trichosanthes homophylla Hayata (1921) Icon. Pl. Formosan. 10: 8
Trichosanthes hylonoma Hand.-Mazz. (1936) Symb. Sin. Pt. 7: 1066
Trichosanthes ishigakiensis E.Walker (1971) J. Jap. Bot. 46: 71
Trichosanthes jinggangshanica C.H.Yueh (1980) Acta Phytotax. Sin. 18: 342
Trichosanthes kirilowii Maxim. (1859) Prim. Fl. Amur. 482
Trichosanthes mianyangensis C.H.Yueh & R.G.Liao (1992) Bull. Bot. Res., Harbin 2: 115
Trichosanthes miyagii Hayata (1921) Icon. Pl. Formosan. 10: 11
Trichosanthes multiloba Miq. (1865) Ann. Mus. Bot. Lugd.-Bat. 2: 82
Trichosanthes rosthornii Harms (1901) Bot. Jahrb. Syst. 29: 603
Trichosanthes sect. Gymnopetalum (Arn.) H.J.de Boer, comb. et stat. nov. – Gymnopetalum Arn. (1840) Madras J. Lit. Sci. 12: 52. –Type: Bryonia tubiflora Wight & Arn. [ = Trichosanthes tubiflora (Wight & Arn.) H.J.de Boer].
Trichosanthes tubiflora (Wight & Arn.) H.J.de Boer, comb. nov. – Bryonia tubiflora Wight & Arn. (1834) Prodr. Fl. Ind. Orient. 1: 347. – Gymnopetalum tubiflorum (Wight & Arn.) Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 388 –Type: Rottler s.n. ex Herb. Klein in Herb. Wight Cat. 1118, February 1796 (holotype K; isotypes E, several duplicates), Trincomalee, Ceylon.
Trichosanthes sect. PseudovariiferaH.J.de Boer, sect. nov. Diagnosis: Similar to sect. Foliobracteola, but male flowers with stamens inserted low in receptacle tube, with pseudo-ovary (a thick-walled basal part of the receptacle tube, without staminodes), and fruit with white pulp. Type: Trichosanthes villosa Blume.
Trichosanthes kerrii Craib (1914) Bull. Misc. Inform. Kew: 7
Trichosanthes phonsenae Duyfjes & Pruesapan (2004) Thai Forest Bull., Bot. 32: 9
Trichosanthes sericeifolia C.Y.Cheng & C.H.Yueh (1980) Acta Phytotax. Sin. 18: 346
Trichosanthes villosa Blume (1826) Bijdr. Fl. Ned. Ind. 15: 934
Trichosanthes sect. Trichosanthes
Trichosanthes cucumerina L. (1753) Sp. Pl.: 1008
Trichosanthes dafangensis N.G.Ye & S.J.Li (1989) Acta Phytotax. Sin. 27: 153
Trichosanthes dioica Roxb. (1832) Fl. Ind. 3: 701
Trichosanthes integrifolia Thwaites (1859) Enum. Pl. Zeyl. [Thwaites]: 127
Trichosanthes lobata Roxb. (1832) Fl. Ind. 3: 703. – Heterotypic synonyms: Trichosanthes perrotetiana Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 362; Trichosanthes villosula Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 342
Trichosanthes nervifolia L. (1753) Sp. Pl.: 1008
Trichosanthes sect. Truncata C.Y.Cheng & C.H.Yueh (1974) Acta Phytotax. Sin. 12: 427. – Type: Trichosanthes truncata C.B.Clarke
Trichosanthes truncata C.B.Clarke (1879) Fl. Brit. India [J.D. Hooker] 2: 608. – Heterotypic synonym: Trichosanthes ovata Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 365
Trichosanthes sect. Villosae (Yueh & L.Q.Huang) H.J.de Boer, stat. nov. – Trichosanthes subsect. Villosae Yueh & L.Q.Huang (1997) Act. Phytotax. Sinica 35: 127. – Type: Trichosanthes subvelutina F.Muell. ex Cogn.
Trichosanthes subvelutina F.Muell. ex Cogn. (1881) Monogr. Phan. [A.DC. & C.DC.] 3: 366