Trichosanthes.
Trichosanthes L.
Trichosanthes L. Sp. Pl. 2: 1008. 1753.
Type: Trichosanthes anguina L., Micheli, Nov. Pl. Gen. pl. 9 (lectotype). 1729; accepted name: Trichosanthes cucumerina L.
Anguina Mill., Fig. Pl. 21. 1755.
Cucumeroides Gaertner, Fruct. 2: 485. 1791.
Involucraria Ser., Mém. Soc. Phys. Genève 3: 27. 1825.
Gymnopetalum Arn. in R. Wight, Madras J. Lit. Sci. 12: 52. 1840.
Tripodanthera M. J. Roem., Fam. Nat. Syn. Monogr. 2: 11, 48. 1846.
Scotanthus Naudin, Ann. Sci. Nat. Bot. ser. 4. 16: 172. 1862.
Eopepon Naudin, Ann. Sci. Nat. Bot. ser. 5. 5: 31. 1866.
Platygonia Naudin, Ann. Sci. Nat. Bot. ser. 5. 5: 33. 1866.
Annual or perennial climbers with several meters long herbaceous shoots, often with woody rootstock, and dioecious or rarely monoecious sex system. The leaves are simple, petiolate, often with small probract at the petiole base, with entire or palmately 3-7(-9)-lobed blade, rarely compound, 3-5-foliate and usually denticulate margin. The tendrils are 2-5-fid (rarely simple), sometimes with apical adhesive pads. The flowers are medium-sized to large, mostly fragrant and opening at night. The male flowers are usually bracteate, in racemes, rarely solitary and coaxillary with a raceme, the female flowers are produced solitary (rarely in racemes). The receptacle-tube is elongate, tubular to cylindric, often dilated at the apex with five entire, serrate or laciniate, triangular to lanceolate sepals. The five petals are long-fimbriate less often entire, white, rarely pink or red. The three stamens are inserted halfway up the tube on very short, free filaments. Two anthers are bithecous, one is monothecous with triplicate thecae. The pollen is 3-(4)(col)porate, psilate, perforate, rugulate, verrucate or (micro)reticulate, medium-sized to large (polar axis 32-98 µm, equatorial axis 34-125 µm, (Khunwasi 1998, Pruesapan and van der Ham 2005)). The ovary is ovoid or fusiform, glabrous to villous with three placentae and many, horizontal, pendent ovules. The style is slender to filiform with three entire or bifid stigmata. The fruit is fleshy, pulpy, globose, ovoid to ellipsoid or fusiform, indehiscent, usually glabrous and smooth, sometimes ribbed, ripening orange to red (or rarely metallic blue). The many seeds are oblong or ovate, sagittate, or rounded, 1-loculed, compressed or 3-loculed, turgid, the two lateral locules empty. The testa is smooth, yellowish to white, black or dark brown, with or without distinct margin. The chromosome number ranges from n = 11 or 12 (Beevy and Kuriachan 1996) to 2n = 88 in T. kirilowii (Huang et al. 1994).
The ca. 100 species grow in Asian rainforests from India to Northeastern Australia. Trichosanthes cucumerina var. anguina is cultivated as vegetable in tropical regions of Africa, Asia, Central and South America.
The genus is placed in tribe Sicyoeae, where it groups as sister to the New World clade (Schaefer & Renner 2011, Bellot et al. (in press)), from which it split about 30 million years ago (Schaefer et al. 2009).
Accepted species
Trichosanthes adhaerens W.J. de Wilde & Duyfjes, Sandakania 14: 11. 2004.
Trichosanthes anamalaiensis Bedd., Madras J. Lit. Sci. III, 1: 47. 1864.
Trichosanthes auriculata Rugayah, Reinwardtia 11: 216. 1998.
Trichosanthes baviensis Gagnep., Bull. Mus. Natl. Hist. Nat. 24: 379. 1918.
Trichosanthes beccariana Cogn., Monogr. Phan. 3: 380. 1881.
Trichosanthes borneensis Cogn., Monogr. Phan. 3: 369. 1881.
Trichosanthes bracteata (Lam.) Voigt, Hort. Suburb. Calcutt. 58. 1845.
Trichosanthes celebica Cogn., Monogr. Phan. 3: 385. 1881.
Trichosanthes cordata Roxb., Fl. Ind., ed. 1832 3: 703-704. 1832.
Trichosanthes coriacea Blume, Bijdr. Fl. Ned. Ind. 15: 935. 1826.
Trichosanthes costata Blume, Bijdr. Fl. Ned. Ind. 15: 933. 1826.
Trichosanthes cucumerina L., Sp. Pl. 1008. 1753.
Trichosanthes dafangensis N.G. Ye & S.J. Li, Acta Phytotax. Sin. 26: 465-466. 1988.
Trichosanthes densiflora Rugayah, Reinwardtia 11: 252. 1999.
Trichosanthes dentifera Rugayah, Reinwardtia 11: 253. 1999.
Trichosanthes dieniensis Merr. & L.M. Perry, J. Arnold Arbor. 30: 59. 1949.
Trichosanthes dioica Roxb., Fl. Ind., ed. 1832 3: 701. 1832.
Trichosanthes dolichosperma Duyfjes & Pruesapan, Thai Forest Bull., Bot. 32: 84. 2004.
Trichosanthes dunniana H. Lév., Repert. Spec. Nov. Regni Veg. 10 (243-247): 148. 1911.
Trichosanthes edulis Rugayah, Reinwardtia 11: 254. 1999.
Trichosanthes ellipsoidea Merr., Philipp. J. Sci., C 13: 332. 1918.
Trichosanthes elmeri Merr., Univ. Calif. Publ. Bot. 15: 299. 1929.
Trichosanthes emarginata Rugayah, Reinwardtia 11: 258. 1999.
Trichosanthes erosa Duyfjes & Pruesapan, Thai Forest Bull., Bot. 32: 85. 2004.
Trichosanthes fissibracteata C.Y.Wu ex C.Y.Cheng & Yueh, Acta Phytotax. Sin. 12: 438. 1974.
Trichosanthes floresana Rugayah, Reinwardtia 11: 260. 1999.
Trichosanthes fusca W.J. de Wilde & Duyfjes, Sandakania 14: 17. 2004.
Trichosanthes globosa Blume, Bijdr. Fl. Ned. Ind. 15: 936. 1826.
Trichosanthes hastata Cogn. ex Harms, Bot. Jahrb. Syst. 60: 160. 1925.
Trichosanthes holtzei F. Muell., Australas. J. Pharm. 1: 447. 1886.
Trichosanthes homophylla Hayata, Icon. Pl. Formosan. 10: 8-10. 1921.
Trichosanthes hylonoma Hand.-Mazz., Symb. Sin. 7: 1066. 1936.
Trichosanthes integrifolia Thwaites, Enum. Pl. Zeyl. 127. 1859.
Trichosanthes intermedia W.J. de Wilde & Duyfjes, Sandakania 14: 19. 2004.
Trichosanthes inthanonensis Duyfjes & Pruesapan, Thai Forest Bull., Bot. 32: 86. 2004.
Trichosanthes ishigakiensis E. Walker, J. Jap. Bot. 46: 71. 1971.
Trichosanthes jinggangshanica C.H. Yueh, Acta Phytotax. Sin. 18: 342, pl. 2. 1980.
Trichosanthes kerrii W.G. Craib, Bull. Misc. Inform. Kew 1914: 7. 1914.
Trichosanthes khasiana Kundu, J. Bot. 77: 11. 1939.
Trichosanthes kinabaluensis Rugayah, Reinwardtia 11: 262. 1999.
Trichosanthes kirilowii Maxim., Prim. Fl. Amur. 482. 1859.
Trichosanthes kostermansii Duyfjes & Pruesapan, Thai Forest Bull., Bot. 32: 89. 2004.
Trichosanthes laceribractea Hayata, J. Coll. Sci. Imp. Univ. Tokyo 30: 117-118. 1911.
Trichosanthes laeoica C.Y. Cheng & Lu Q. Huang, Bull. Bot. Res., Harbin 16: 503. 1996.
Trichosanthes lepiniana (Naudin) Cogn., Monogr. Phan. 3: 377. 1881.
Trichosanthes leuserensis Rugayah, Reinwardtia 11: 265. 1999.
Trichosanthes lobata Roxb., Fl. Ind. 3: 703. 1832.
Trichosanthes longispicata Rugayah, Reinwardtia 11: 266. 1999.
Trichosanthes mianyangensis C.H. Yueh & R.G. Liao, Bull. Bot. Res., Harbin 2: 115-117. 1992.
Trichosanthes miyagii Hayata, Icon. Pl. Formosan. 10: 11. 1921.
Trichosanthes montana Rugayah, Reinwardtia 11: 218. 1998.
Trichosanthes morrisii W.E. Cooper, Austrobaileya 8: 381. 2011.
Trichosanthes mucronata Rugayah, Reinwardtia 11: 268. 1999.
Trichosanthes multiloba Miq., Ann. Mus. Bot. Lugd.-Bat. 2: 82. 1865.
Trichosanthes nervifolia L., Sp. Pl. 1008. 1753.
Trichosanthes obscura Rugayah, Reinwardtia 11: 269. 1999.
Trichosanthes obtusiloba C.Y. Wu ex C.Y. Cheng & Yueh, Acta Phytotax. Sin. 12: 431. 1974.
Trichosanthes odontosperma W.E. Cooper & A.J. Ford, Austrobaileya 8: 126. 2010.
Trichosanthes orientalis (W.J. de Wilde & Duyfjes) H.J. de Boer, Phytokeys 12: 30. 2012.
Trichosanthes pallida Duyfjes & Pruesapan, Thai Forest Bull., Bot. 32: 90. 2004.
Trichosanthes papuana F.M. Bailey, Queensland Agric. J. 7: 349. 1900.
Trichosanthes pedata Merr. & Chun, Sunyatsenia 2: 20. 1934.
Trichosanthes pendula Rugayah, Reinwardtia 11: 219. 1998.
Trichosanthes pentaphylla F. Muell. in Benth., Fl. Austral. 3: 314. 1867.
Trichosanthes philippinensis Rugayah, Reinwardtia 11: 271. 1999.
Trichosanthes phonsenae Duyfjes & Pruesapan, Thai Forest Bull., Bot. 32: 91. 2004.
Trichosanthes pilosa Lour., Fl. Cochinch. 588. 1790.
Trichosanthes planiglans Rugayah, Reinwardtia 11: 273. 1999.
Trichosanthes postarii W.J. de Wilde & Duyfjes, Sandakania 14: 26. 2004.
Trichosanthes pubera Blume, Bijdr. Fl. Ned. Ind. 15: 936. 1826
Trichosanthes pulleana Cogn. ex Harms, Bot. Jahrb. Syst. 60: 160. 1925.
Trichosanthes quinquangulata A. Gray, U.S. Expl. Exped., Phan. 1: 645. 1854.
Trichosanthes quinquefolia C.Y. Wu ex C.Y. Cheng & C.H. Yueh, Acta Phytotax. Sin. 18: 351-352. 1980.
Trichosanthes refracta C.H. Yueh, Bull. Bot. Res., Harbin 16: 500. 1996.
Trichosanthes reticulinervis C.Y. Wu ex S.K. Chen, Bull. Bot. Res., Harbin 5: 114-115. 1985.
Trichosanthes rosthornii Harms, Bot. Jahrb. Syst. 29: 603. 1901.
Trichosanthes rotundifolia Rugayah, Reinwardtia 11: 223. 1998.
Trichosanthes rubriflos Thorel ex Cayla, Bull. Mus. Natl. Hist. Nat. 14: 170-172. 1908.
Trichosanthes rugatisemina C.Y. Cheng & C.H. Yueh, Acta Phytotax. Sin. 12: 440-441. 1974.
Trichosanthes scabra Lour., Fl. Cochinch. 589. 1790.
Trichosanthes schlechteri Cogn. ex Harms, Bot. Jahrb. Syst. 60: 159. 1925.
Trichosanthes sepilokensis Rugayah, Reinwardtia 11: 275. 1999.
Trichosanthes sericeifolia C.Y. Cheng & C.H. Yueh, Acta Phytotax. Sin. 18: 346-347. 1980.
Trichosanthes siamensis Duyfjes & Pruesapan, Thai Forest Bull., Bot. 32: 97. 2004.
Trichosanthes smilacifolia C.Y. Wu ex C.H. Yueh & C.Y. Cheng, Acta Phytotax. Sin. 18: 347-348: 1980.
Trichosanthes subrosea C.Y. Cheng & C.H. Yueh, Acta Phytotax. Sin. 18: 349: 1980.
Trichosanthes subvelutina F. Muell. ex Cogn., Monogr. Phan. 3: 366. 1881.
Trichosanthes tetragonosperma C.Y. Cheng & C.H. Yueh, Acta Phytotax. Sin. 12: 425: 1974.
Trichosanthes tricuspidata Lour., Fl. Cochinch. 2: 589. 1790.
Trichosanthes trifolia (L.) Blume, Bijdr. Fl. Ned. Ind. 903. 1826.
Trichosanthes truncata C.B. Clarke, Fl. Brit. India 2: 608 1879.
Trichosanthes tubiflora (Wight & Arn.) H.J. de Boer, PhytoKeys 12: 31. 2012.
Trichosanthes valida Rugayah, Reinwardtia 11: 277. 1999.
Trichosanthes villosa Blume, Bijdr. Fl. Ned. Ind. 15: 934. 1826.
Trichosanthes wallichiana (Ser.) Wight, Madras J. Lit. Sci. 12: 52. 1840.
Trichosanthes wawraei Cogn., Monogr. Phan. 3: 384. 1881.
Trichosanthes yixingense Z.P. Wang & Q.Z. Xie, Acta Phytotax. Sin. 19: 483. 1981.
Literature
Beevy, S. S., and P. Kuriachan. 1996. Pollen morphology and taxonomy of the genus Trichosanthes (Cucurbitaceae). Journal of Palynology 32: 21-27.
Bellot, S., Mitchell, T.C., and H. Schaefer. Phylogenetic informativeness analyses to clarify past diversification processes in Cucurbitaceae. Scientific Reports (in press).
De Boer, H.J. and M. Thulin. 2012. Synopsis of Trichosanthes (Cucurbitaceae) based on recent molecular phylogenetic data. PhytoKeys 12: 23-33.
De Wilde, W.J.J.O. and B.E.E. Duyfjes. 2004. The genus Trichosanthes (Cucurbitaceae) in Sabah. Sandakania 14: 5-32.
De Wilde, W.J.J.O. and B.E.E. Duyfjes. 2006. Review of the genus Gymnopetalum (Cucurbitaceae). Blumea 51: 281-296.
Huang, L. Q., Yue, C.X. and M.X. Li. 1994. Chromosome numbers of 8 species in Trichosanthes. Acta Botanica Yunnanica 16: 95-96.
Khunwasi, C. 1998. Palynology of the Cucurbitaceae. Doctoral Dissertation Naturwiss. Fak., University of Innsbruck.
Pruesapan, K. and R.W.J.M. van der Ham. 2005. Pollen morphology of Trichosanthes (Cucurbitaceae). Grana 44: 75-90.
Rugayah, and W.J.J.O. de Wilde. 1997. Trichosanthes L. (Cucurbitaceae) in Java. Blumea 42: 471-482.
Rugayah, and W.J.J.O. de Wilde. 1999. Conspectus of Trichosanthes (Cucurbitaceae) in Malesia. Reinwardtia 11: 227-280.
Schaefer, H. and S.S. Renner. 2011. Phylogenetic relationships in the order Cucurbitales and a new classification of the gourd family (Cucurbitaceae). Taxon 60: 122-138.
Schaefer, H., Heibl, C., and S.S. Renner. 2009. Gourds afloat: a dated phylogeny reveals an Asian origin of the gourd family (Cucurbitaceae) and numerous oversea dispersal events. Proceedings of the Royal Society B 276: 843-851.
