Cheniella gen. nov. (Leguminosae: Cercidoideae) from southern China, Indochina and Malesia

Ruth P. Clark, Barbara A. Mackinder, Hannah Banks


For much of the last thirty years, the caesalpinioid genus Bauhinia has been recognised by numerous authors as a broadly circumscribed, ecologically, morphologically and palynologically diverse pantropical taxon, comprising several subgenera. One of these, Bauhinia subg. Phanera has recently been reinstated at generic rank based on a synthesis of morphological and molecular data. Nevertheless, there remains considerable diversity within Phanera. Following a review of palynological and molecular studies of Phanera in conjunction with a careful re-examination of the morphological heterogeneity within the genus, we have found strong evidence that the species of Phanera subsect. Corymbosae are a natural group that warrant generic status. We describe here the genus Cheniella R.Clark & Mackinder gen. nov. to accommodate them. It comprises 10 species and 3 subspecies, one newly described here. Generic characters include leaves that are simple and emarginate or bilobed; flowers with elongate hypanthia which are as long as or much longer than the sepals; pods that are glabrous, compressed, oblong, indehiscent or tardily dehiscent; and with numerous seeds, the seeds bearing an unusually long funicle extending most of the way around their circumference. A further distinctive floral character was found to be a fleshy disc on which the staminodes are mounted. An analysis carried out for this study reveals Cheniella to be characterised by a pollen type that is unique to the genus and previously unknown in the Leguminosae. Species diversity is richest in southern China, the full distribution extending westward to India and south- and eastward through Indochina into Malesia.


Bauhinia; Caesalpinioideae; Cercideae; Fabaceae; Phanera; pollen; staminodes

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Arroyo M.T.K. 1981. Breeding systems and pollination biology in Leguminosae. In: Polhill R.M. & Raven P.H. (eds) Advances in Legume Systematics 2: 723–769. Royal Botanic Gardens, Kew.

Bandyopadhyay S. 2001. On the distribution range of Bauhinia glauca (Benth.) Benth. (Leguminosae: Caesalpinioideae) in India. Journal of the Bombay Natural History Society 98 (1): 149–150.

Bandyopadhyay S., Ghoshal P.P., & Pathak M.K. 2012. Fifty new combinations in Phanera Lour. (Leguminosae: Caesalpinioideae) from Paleotropical Region. Bangladesh Journal of Plant Taxonomy 19: 55–61.

Bandyopadhyay S. & Ghoshal P.P. 2014. Two new combinations in Lysiphyllum (Leguminosae-Caesalpinioideae). Phytotaxa 178 (4): 298–300.

Bandyopadhyay S. & Sharma B.D. 1992. Lectotypification of Bauhinia glauca ssp. tenuiflora (Leguminosae). Journal of the Bombay Natural History Society 89 (1): 393–395.

Banks H. & Klitgaard B.B. 2000. Palynological contributions to the systematics of detarioid legumes (Leguminosae: Caesalpinioideae). In: Herendeen P.S. & Bruneau A. (eds) Advances in Legume Systematics 9: 79–106. Royal Botanic Gardens, Kew.

Banks H. & Rudall P. 2016. Pollen structure and function in caesalpinioid legumes. American Journal of Botany 103: 423–436.

Banks H., Feist-Burkhart S. & Klitgaard B. 2006. The Unique Pollen Morphology of Duparquetia (Leguminosae: Caesalpinioideae): Developmental Evidence of Aperture Orientation Using Confocal Microscopy. Annals of Botany 98: 107–115.

Banks H., Forest F. & Lewis G. 2013. Palynological contribution to the systematics and taxonomy of Bauhinia s.l. (Leguminosae: Cercideae). South African Journal of Botany 89: 219–226.

Banks H., Forest F. & Lewis G. 2014. Evolution and diversity of pollen morphology in tribe Cercideae (Leguminosae). Taxon 63: 299–314.

Banks H., Klitgaard B.B., Lewis G.P., Crane P.R. & Bruneau A. 2003. Pollen and the systematics of tribes Caesalpinieae and Cassieae (Caesalpinioideae: Leguminosae). In: Klitgaard B. & Bruneau A. Advances in Legume Systematics, Part 10. Higher Level Systematics: 95–122. Royal Botanic Gardens, Kew.

Bentham G. 1852. Leguminosae. In: Miquel F.A.W. Plantae junghuhnianae 2: 205–269.

Bruneau A., Forest F., Herendeen P.S., Klitgaard B.B. & Lewis G.P. 2001. Phylogenetic relationships in the Caesalpinioideae (Leguminosae) as inferred from chloroplast trnL intron sequences. Systematic Botany 26: 487–514.

Bruneau A., Mecure M., Lewis G.P. & Herendeen P.S. 2008. Phylogenetic patterns and diversification in the caesalpinioid legumes. Botany 86: 697–718.

Chen H.Y. 1938. New species of Bauhinia from China. Journal of the Arnold Arboretum 19: 129–133.

Chen T.C. 1988. New taxa of the genus Bauhinia L. from China. Guihaia 8 (1): 43–51.

Clarke C.B. 1889. On the plants of Kohima and Muneypore. Journal of the Linnean Society, Botany 25 (165–107): 1–107.

Dezhao C., Chen T., Zhang D., Larsen K. & Larsen S.S. (eds) 2010. Flora of China 10, Fabaceae: 6–21. Science Press, Beijing and Missouri Botanical Garden Press, St Louis.

Duarte-Almeida J.M., Negri G. & Salatino A. 2004. Volatile oils in leaves of Bauhinia (Fabaceae Caesalpinioideae). Biochemical Systematics and Ecology 32: 747–753.

Erdtman G. 1960. The acetolysis method. Svensk Botanisk Tidskrift 54: 561–564.

Ferguson I.K. & Pearce K.J. 1986. Observations in the pollen morphology of the genus Bauhinia L. (Leguminosae: Caesalpinioideae) in the Neotropics. In: Blackmore S. & Ferguson I.K. (eds) Pollen and Spores, Form and Function: 283–296. Academic Press, London.

Gagnepain F. 1912. Bauhinia nouveaux d’Extrême-Orient. Notulae Systematicae 2 (6): 168–182.

Gamerro J.C. & Fortunato R.H. 2001. Morfología del polen de las especies argentinas de Bauhinia (Cercideae, Caesalpinioideae, Fabaceae). Annals of the Missouri Botanic Garden 88: 144–158.

Gogoi R. & Bandyopadhyay S. 2014. A new variety of Phanera glauca subsp. tenuiflora (Fabaceae: Caesalpinioideae) from India. Journal of the Botanical Research Institute of Texas 8 (1): 71–75.

Graham A. & Barker G. 1981. Palynological and tribal classification in the Caesalpinioideae. In: Polhill R.M. & Raven P.H. (eds) Advances in Legume Systematics 2: 801–834. Royal Botanic Gardens, Kew.

Hao G., Zhang D.X., Zhang M.Y., Guo L.X. & Li S.J. 2003. Phylogenetics of Bauhinia subgenus Phanera (Leguminosae: Caesalpinioideae) based on ITS sequences of nuclear ribosomal DNA. Botanical Bulletin of Academia Sinica 44: 223–228.

Hind N. 2015. Compositae (Asteraceae). In: Utteridge T. & Bramley G. (eds) The Kew Tropical Plant Families Identification Handbook, second edition: 206–207. Kew Publishing, Royal Botanic Gardens Kew, Richmond.

Hokche O. & Ramírez N. 1990. Pollination ecology of seven species of Bauhinia L. (Leguminosae: Caesalpinioideae). Annals of the Missouri Botanical Garden 77 (3): 559–572.

Hou D., Larsen K. & Larsen S.S. 1996. Caesalpiniaceae. In: Hou D., Larsen S.S., Laferriere J.E. & Duyfjes B.E.E. (eds) Flora Malesiana. Series 1, 12 (2): 409–784. Leiden University, Leiden.

Hughes J.B., Round P.D. & Woodruff D.S. 2003. The Indochinese-Sundaic faunal transition at the Isthmus of Kra: an analysis of resident forest bird species distributions. Journal of Biogeography 30: 569–580.

IUCN. 2001. IUCN Red List Categories and Criteria: version 3.1. IUCN, Gland, Switzerland and Cambridge.

JSTOR Global Plants. 2016. Available from: [accessed 16 Feb. 2016].

Kumar R., Bandyopadhyay S. & Sharma S. 2014. A new variety of Phanera glauca subsp. tenuiflora (Leguminosae: Caesalpinioideae) from India. Phytotaxa 166 (2): 155–159.

Lai M., Sceppa J., Ballenger J.A. & Doyle J.J. 1997. Polymorphism for the presence of the rpL2 Intron in chloroplast genomes of Bauhinia (Leguminosae). Systematic Botany 22: 519–528.

Larsen K. & Larsen S.S. 1973. The genus Bauhinia in Thailand. Natural History Bulletin of the Siam Society 25 (1–2): 10.

Larsen K. & Larsen S.S. 1982. Notes on some Asian Bauhinia. Nordic Journal of Botany 2: 329–332.

Larsen K. & Larsen S.S. 1991. Notes on the genus Bauhinia (Leguminosae-Caesalpinioideae) in SE Asia. Nordic Journal of Botany 11: 6: 629–634.

Larsen K. & Larsen S.S. 1993. New taxa & nomenclatural combinations in Malesian Bauhinia (Leguminosae-Caesalpinioideae). Nordic Jornal of Botany 13: 6: 657–665.

Larsen K., Larsen S.S. & Vidal J. 1980. Légumineuses–Césalpinioïdées. In: Aubréville A. & Leroy J.-F. (eds) Flore du Cambodge, du Laos et du Viêtnam: 1–227. Muséum National d’Histoire Naturelle, Paris.

Larsen K., Larsen S.S. & Vidal J. 1984. Leguminosae-Caesalpinioideae. In: Smitinand T. & Larsen K. (eds) Flora of Thailand. The Forest Herbarium, Royal Forest Department, Bangkok.

Larsen S.S. 1975. Pollen morphology of Thai species of Bauhinia (Caesalpiniaceae). Grana 14: 113–131.

Lau C.P., Saunders R.M.K. & Ramsden L. 2009. Floral biology, breeding systems and population genetic structure of three climbing Bauhinia species (Leguminosae: Caesalpinioideae) in Hong Kong, China. Journal of Tropical Ecology 25: 147–159.

Lewis G.P. 1998. Caesalpinia–a Revision of the Poincianella-Erythrostemon Group: 15–17. Royal Botanic Gardens, Kew.

Lewis G. & Forest F. 2005. Cercideae. In: Lewis G., Schrire B., Mackinder B. & Lock M. (eds) Legumes of the World: 57–67. Royal Botanic Gardens, Kew.

Lewis G.P., Simpson B.B. & Neff J.L. 2000. Progress in understanding the reproductive biology of the Caesalpinioideae (Leguminosae). In: Herendeen P.S. & Bruneau A. (eds) Advances in Legume Systematics 9: 65–78. Royal Botanic Gardens, Kew.

LPWG (The Legume Phylogeny Working Group). 2013. Legume phylogeny and classification in the 21st century: progress, prospects and lessons for other species-rich clades. Taxon 62 (2): 217–247.

LPWG (The Legume Phylogeny Working Group). 2017. A new subfamily classification of the Leguminosae based on a taxonomically comprehensive phylogeny. Taxon 66 (1): 44–57.

Mackinder B. & Clark R. 2014. A synopsis of the Asian and Australasian genus Phanera Lour. (Cercideae: Caesalpinioideae: Leguminosae) including 19 new combinations. Phytotaxa 66: 49–68.

Merrill E.D. 1942. Records of Indo-Chinese Plants 3. Journal of the Arnold Arboretum 23: 156–197.

O’Toole C. & Raw A. 1991. Bees of the World. Blandford, London.

Parnell J. 2013. The biogeography of the Isthmus of Kra region: a review. Nordic Journal of Botany 31: 1–15.

Pérez-Barrales R., Arroyo J. & Armbruster W.S. 2007. Differences in pollinator faunas may generate geographic differences in floral morphology and integration in Narcissus papyraceus (Amaryllidaceae). Oikos 116: 1904–1918.

Punt W., Hoen P.P., Blackmore S., Nilsson S. & Thomas A. Le. 2007. Glossary of pollen and spore terminology. Review of Palaeobotany and Palynology 143 (1–2): 1–81.

Queiroz L.P. de 2006. New species and new combinations in Phanera Lour. (Caesalpinioideae: Cercideae) from the Caatinga Biome. Neodiversity 1: 6–10.

Richardson J.E., Costion C.M. & Muellner A.N. 2012. The Malesian floristic interchange: plant migration patterns across Wallace’s Line. In: Gower D.J., Johnson K., Richardson J., Rosen B., Rüber L. & Williams S. (eds) Biotic Evolution and Environmental Change in Southeast Asia: 138–163. Cambridge University Press, Cambridge.

Roxburgh W. 1814. Hortus Bengalensis. Mission Press, Serampore.

Roxburgh W. 1832. Flora indica ed. 1832. W. Thacker & Co., Calcutta.

Salatino A., Blatt C.T.T., Santos W.A.C. dos & Vaz A.M.S. da F. 1999. Foliar flavonoids of nine species of Bauhinia. Revista Brasileira de Botânica 22 (1): 17–20.

Santos F.A.R. dos, Novaes D.M. & de Queiroz L.P. 2012. Pollen of Bauhinia L. and Phanera Lour. (Leguminosae-Caesalpinioideae) from the Brazilian Caatinga. American Journal of Plant Sciences 3: 909–920.

Sinou C., Forest F., Lewis G.P. & Bruneau A. 2009. The genus Bauhinia s.l. (Leguminosae): a phylogeny based on the plastid trnL-trnF region. Botany 87: 947–960.

Trethowan L.A., Clark R.P. & Mackinder B.A. 2015. A synopsis of the neotropical genus Schnella (Cercideae: Caesalpinioideae: Leguminosae) including 12 new combinations. Phytotaxa 204 (4): 237–252.

Vaz A.M.S. da F. 2010. New combinations in Phanera (Leguminosae; Cercideae) from Brazil. Rodriguésia 61 (Suppl.): S33–S40.

Verdcourt B. 1979. A Manual of New Guinea Legumes. Papua New Guinea, Botany Bulletin 11. Office of Forests, Division of Botany, Lae.

Walker-Larsen J. & Harder L.D. 2000. The evolution of staminodes in angiosperms: patterns of stamen reduction, loss, and functional re-invention. American Journal of Botany 87 (10): 1367–1384.

Walter K.S. & Gillett H.J. (eds) 1997. 1997 IUCN Red List of Threatened Plants. IUCN, Gland, Switzerland and Cambridge.

Wang Q., Song Z., Chen Y., Shen S. & Li Z. 2014. Leaves and fruits of Bauhinia (Leguminosae, Caesalpinioideae, Cercideae) from the Oligocene Ningming Formation of Guangxi, South China and their biogeographic implications. BMC Evolutionary Biology 14: 1471–2148.

Wilkin P. & Schuiteman A. 2015. Orchidaceae. In: Utteridge T. & Bramley G. (eds) The Kew Tropical Plant Families Identification Handbook, second edition: 36–37. Kew Publishing, Royal Botanic Gardens Kew, Richmond.

Wit H.C. de 1956. A revision of Malaysian Bauhinieae. Reinwardtia 3: 381–539.

Woodruff D.S. 2010. Biogeography and conservation in Southeast Asia: how 2.7 million years of repeated environmental fluctuations affect today’s patterns and the future of the remaining refugial-phase biodiversity. Biodiversity and Conservation 19: 919–941.

Wunderlin R. 1976. Enumeration and typification of genera in tribe Cercideae. Rhodora 78: 750–760.

Wunderlin R. 1979. Consideration of Barklya and the subtribes of the Cercideae (Caesalpinioideae: Fabaceae). Phytologia 44: 325–327.

Wunderlin R.P. 2010. Reorganization of the Cercideae (Fabaceae: Caesalpinioideae). Phytoneuron 48: 1–5.

Wunderlin R.P. 2011. New combination in Phanera (Fabaceae). Phytoneuron 19: 1–2.

Wunderlin R., Larsen K. & Larsen S.S. 1981. Cercideae. In: Polhill R.M. & Raven P.H. (eds) Advances in Legume Systematics, Part 1: 107–116. Royal Botanic Gardens, Kew.

Wunderlin R., Larsen K. & Larsen S.S. 1987. Reorganization of the Cercideae (Fabaceae: Caesalpinioideae). Biologiske Skrifter 28: 1–40.

Zhang D.X. 1994. Leaf venation of Cercideae (Leguminosae). Journal of Tropical and Subtropical Botany 2: 45–57.

Zhang D.X. & Chen T.C. 1992. Systematics and Biogeography of Bauhinia L. (Leguminosae) II. Cladistic Analysis of Ser. Corymbosae. Acta Botanica Austro Sinica 8: 31–35.


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