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Brandon-Jones Langur Taxonomy 5Jul04.Pmd PAPER ZOOS' PRINT JOURNAL 19(8): 1552-1594 A TAXONOMIC REVISION OF THE LANGURS AND LEAF MONKEYS (PRIMATES: COLOBINAE) OF SOUTH ASIA Douglas Brandon-Jones 32a Back Lane, Richmond, Surrey TW10 7LF, United Kingdom Email: [email protected] ABSTRACT evident as the workshop progressed, but discussion with the Over 300 study specimens from Chicago, Kolkata, Leiden, participants, and an impromptu visit to the Annamalai Hills, Tamil London, Mumbai, New York, Paris and Washington DC are Nadu, clarified some of the issues. Seizing the opportunity of combined with a literature survey to review the taxonomy of Semnopithecus johnii, S. entellus and S. priam. Simia johnii the stopover at Mumbai, five days were spent examining the Fischer, 1829 is declared a nomen protectum. The 15 valuable colobine collection accrued by the Bombay Natural subspecies Ellerman and Morrison-Scott (1966) recognized History Society (BNHS), including 56 hanuman langur for Semnopithecus entellus are reduced to seven subspecies specimens (33 skins and skulls, 10 skins only, and 13 skulls for S. entellus and two for S. priam. Type specimens are only). Together with the 146 specimens (101 skins and skulls, 13 documented, pelage colour variation described, and the nomenclature and subspecific geographic distributions skins only, and 32 skulls only) at the Natural History Museum, discussed. Two previously recognized subspecies are London (ZD); the three skulls at the Odontological Museum, reinterpreted as hybrid populations of S. entellus and S. Royal College of Surgeons, London; two skins and skulls, and johnii. The central Indian S. entellus anchises, previously three skulls at the National Museum of Natural History, Leiden; suspected as being only an intermediate between one skin and skull, and three skins at the American Museum of subspecies, is found to occupy a substantial discrete distribution. The Himalayan subspecies S. e. schistaceus Natural History, New York (AMNH); the eighteen skins and proves to have a much wider distribution than previously skulls at the Field Museum of Natural History, Chicago (FMNH); conceived, extending from northwestern Bhutan possibly to and the seven skins and skulls at the Smithsonian Institution, Afghanistan. The corollary is that the distribution of S. e. Washington, DC (USNM); this results in the most ajax is reduced to three known localities, two of them over comprehensive overview of Indian and Nepal langur taxonomy 930km apart, indicating the intervening occurrence of an as to date, totalling 246 specimens (169 skins and skulls, 26 skins yet unconfirmed high altitude population of this subspecies. only, and 51 skulls only). Blyth (1844b; 1848a), Horsfield (1851) KEYWORDS and the catalogues of the Indian Museum, Kolkata (ZSI), Bangladesh, India, langur, Nepal, Pakistan, primates, published by Blyth (1863), Anderson (1881) and Khajuria (1956a) revision, Semnopithecus entellus, S. johnii, S. priam, supplied data on a further 23 specimens. Oboussier and Maydell subspecies key, taxonomy, zoogeography (1960) presented data on an additional 18 specimens, and Kurup (1965) on two more. Material at the Muséum National d’Histoire ABBREVIATIONS AMNH - American Museum of Natural History, New York; Naturelle, Paris (MNHN) was only briefly examined and BNHS - Bombay Natural History Society; CAMP - descriptions of most of the eleven specimens are taken from the Conservation Assessment and Management Plan; FMNH - literature Field Museum of Natural History, Chicago; ICZN - International Code of Zoological Nomenclature; MNHN - Muséum National d’Histoire Naturelle, Paris; USNM - The Bombay Natural History Society collection proved to Smithsonian Institution, Washington, DC; ZD - Natural include six specimens from Khandala, Maharashtra, India, a History Museum, London; ZSI - Indian Museum, Kolkata locality unrepresented at any of the other above institutions. The peculiarities of these specimens made credible a doubtful This paper springs essentially from an invitation to attend a central Indian subspecies Semnopithecus entellus anchises Conservation Assessment and Management Plan (CAMP) Blyth, 1844 whose recognizability inevitably altered the known Workshop for South Asian Primates, held at the State Forest distribution of other central Indian subspecies, and entailed Service College, Coimbatore, Tamil Nadu, India, from 5-9 March modification of some of the guidance offered to the workshop. 2002 (Molur et al., 2003). The organizers recognized the This negated even some of the recent conclusions by Groves importance of assessing the conservation status of all (2001) and increased the urgency for a thorough reassessment. taxonomically discernible populations, but appreciated the Full justice to langur taxonomy must take account of the now difficulty of accomplishing this aim, given our inadequate extensive literature on langurs. This paper does not purport to knowledge, particularly of some Indian langur subspecies be an exhaustive literature review but covers more key references (subfamily Colobinae). Least known is the South-west Indian than any predecessor. Some relevant sources may have been (Malabar) Langur, but langur subspecific distributions in the overlooked but none, it is hoped, that will significantly affect Himalaya and central India are far from satisfactorily the taxonomic results. The leaf monkeys of Sri Lanka will be understood. A manuscript on the Malabar Langur was drafted covered in a subsequent broader review. They are subject to during the brief interval between the invitation and less pressing, mainly nomenclatural and geographic, taxonomic commencement of the workshop. The deficiencies in our issues. knowledge of langur subspecific distributions became more Manuscript 971; Received 29 November 2002; Revised received 13 April 2004; Revised finally accepted 1 June 2004; © Zoo Outreach Organisation 1552 August 2004 A taxonomic revision of the langurs and leaf monkeys (Primates: Colobinae) of South Asia D. Brandon-Jones Overwhelming evidence now segregates about two-thirds of Along the superciliary hairline Trachypithecus adults have the species formerly amassed in Presbytis Eschscholtz, 1821 moderately lengthened black hair and behind it, except in into Semnopithecus Desmarest, 1822 (Brandon-Jones, 1984; Semnopithecus obscurus shanicus, no frontal whorl. Apart 1993; 1999; 2001; Groves, 1989; 2001; Corbet, 1992). A consensus from most S. obscurus subspecies where integumental divides Semnopithecus into at least two species groups, but depigmentation removes the contrasting background colour, their species composition and hierarchical status remain all Trachypithecus females display an area of varying size, shape disputed (see Oates et al., 1994; Brandon-Jones et al., 2004). and colour (usually white or yellow) of pale pubic skin and hair Semnopithecus vetulus and S. johnii have been assigned their between and below the ischial callosities. Pelage colour is own genus Kasi Reichenbach, 1862 by Hill (1936; 1937c; 1939) otherwise so geographically variable that no further subgeneric and Pocock (1939) but cranial morphology, neonatal pelage characters have been identified. Except for females at Ramnagar colour and sexually dichromatic pubic integument integrate them (27044'N 84027'E), Nepal (Chalise, 1995, p. 35), probably referable with Trachypithecus Reichenbach, 1862. The morphological to S. e. schistaceus, sexually dichromatic pubic integument is resemblance between S. j. johnii and the Vietnamese S. undetected in Semnopithecus (sensu stricto). All (non-hybrid) (Trachypithecus) johnii poliocephalus, two residual adults have a frontal hair whorl, straight projecting black populations of a single species fragmented by glacially-induced eyebrows and a pale ventral pelage colour. During the first two deforestation (Brandon-Jones, 1995; 1996), is closer than that months of life the pelage gradually becomes much paler so that between S. johnii and S. vetulus. Vocal analysis (Hohmann, yearlings are readily distinguished from adults (McCann, 1933, 1988) and intermediate pelage colour (see under S. johnii, S. p. 619). Sugiyama (1966, p. 216) specified that the skin turns priam and S. entellus hypoleucos) indicate natural hybridization from pinkish to white-brown about a week after birth and that of S. johnii with both S. priam and S. entellus. The affiliation the pelage begins to turn white shortly after the skin becomes of S. johnii and S. entellus in mitochondrial cytochrome-b gene predominantly black at about three months. Infants between sequences (Zhang & Ryder, 1998) and their evident hybrid 4.5 to five months and 1-2 years are whiter than adults. When viability corroborate the recent (possibly Holocene) divergence describing adult pelage colour, it is therefore necessary to ensure of Semnopithecus from Trachypithecus inferred from it has passed the transitional condition. The age influence on morphological, biogeographic and climatological evidence adult pelage colour suspected by Blyth (1843b, p. 173) and A. (Brandon-Jones, 1995; 1996). Where genetic and morphological Sharma (pers. comm.) is unconfirmed, but there is individual evidence irresolvably conflict, the interests of field and and geographic variation in the intensity and distribution of institutional specimen identification and consistent taxonomic the yellow or sometimes orange suffusion in the white ventral treatment of extant and fossil taxa are best served by granting pelage colour. This pale colour usually extends onto the flank, precedence to morphology.
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