Tragulus Brisson 1762

Wilson, Don E. & Reeder, DeeAnn, 2005, Order Artiodactyla, Mammal Species of the World: a Taxonomic and Geographic Reference (3 rd Edition), Volume 1, Baltimore: The Johns Hopkins University Press, pp. 637-722 : 648

Tragulus Brisson 1762

Tragulus Brisson 1762 , Regn. Anim., 2nd ed.: 12.

Type Species: Cervus javanicus Osbeck 1765

Synonyms: Lagonebrax Gloger 1841 .

Species and subspecies: 6 species with 50 subspecies:

Species Tragulus javanicus (Osbeck 1765)

Species Tragulus kanchil Raffles 1821

Subspecies Tragulus kanchil subsp. kanchil Raffles 1821

Subspecies Tragulus kanchil subsp. abruptus Chasen 1935

Subspecies Tragulus kanchil subsp. affinis Gray 1861

Subspecies Tragulus kanchil subsp. anambensis Chasen and Kloss 1928

Subspecies Tragulus kanchil subsp. angustiae Kloss 1918

Subspecies Tragulus kanchil subsp. brevipes Miller 1903

Subspecies Tragulus kanchil subsp. carimatae Miller 1906

Subspecies Tragulus kanchil subsp. everetti Bonhote 1903

Subspecies Tragulus kanchil subsp. fulvicollis Lyon 1908

Subspecies Tragulus kanchil subsp. fulviventer Gray 1836

Subspecies Tragulus kanchil subsp. hosei Bonhote 1903

Subspecies Tragulus kanchil subsp. insularis Chasen 1940

Subspecies Tragulus kanchil subsp. klossi Chasen 1935

Subspecies Tragulus kanchil subsp. lampensis Miller 1903

Subspecies Tragulus kanchil subsp. lancavensis Miller 1903

Subspecies Tragulus kanchil subsp. longipes Lyon 1908

Subspecies Tragulus kanchil subsp. luteicollis Lyon 1906

Subspecies Tragulus kanchil subsp. masae Lyon 1916

Subspecies Tragulus kanchil subsp. mergatus Thomas 1923

Subspecies Tragulus kanchil subsp. pallidus Miller 1901

Subspecies Tragulus kanchil subsp. penangensis Kloss 1918

Subspecies Tragulus kanchil subsp. pidonis Chasen 1940

Subspecies Tragulus kanchil subsp. pinius Lyon 1916

Subspecies Tragulus kanchil subsp. ravulus Miller 1903

Subspecies Tragulus kanchil subsp. ravus Miller 1902

Subspecies Tragulus kanchil subsp. rubeus Miller 1903

Subspecies Tragulus kanchil subsp. russeus Miller 1903

Subspecies Tragulus kanchil subsp. russulus Miller 1903

Subspecies Tragulus kanchil subsp. siantanicus Chasen and Kloss 1928

Subspecies Tragulus kanchil subsp. subrufus Miller 1903

Species Tragulus napu F. Cuvier 1822

Subspecies Tragulus napu subsp. napu F. Cuvier 1822

Subspecies Tragulus napu subsp. amoenus Miller 1903

Subspecies Tragulus napu subsp. bancanus Lyon 1906

Subspecies Tragulus napu subsp. banguei Chasen and Kloss 1931

Subspecies Tragulus napu subsp. batuanus Miller 1903

Subspecies Tragulus napu subsp. billitonus Lyon 1906

Subspecies Tragulus napu subsp. bunguranensis Miller 1901

Subspecies Tragulus napu subsp. flavicollis Miller 1903

Subspecies Tragulus napu subsp. hendersoni Chasen 1940

Subspecies Tragulus napu subsp. lutescens Miller 1903

Subspecies Tragulus napu subsp. neubronneri Sody 1931

Subspecies Tragulus napu subsp. niasis Lyon 1916

Subspecies Tragulus napu subsp. nigricollis Miller 1902

Subspecies Tragulus napu subsp. nigrocinctus Miller 1906

Subspecies Tragulus napu subsp. parallelus Miller 1911

Subspecies Tragulus napu subsp. pretiellus Miller 1906

Subspecies Tragulus napu subsp. rufulus Miller 1900

Subspecies Tragulus napu subsp. sebucus Lyon 1911

Subspecies Tragulus napu subsp. stanleyanus Gray 1836

Subspecies Tragulus napu subsp. terutus Thomas and Wroughton 1909

Species Tragulus nigricans Thomas 1892

Species Tragulus versicolor Thomas 1910

Species Tragulus williamsoni Kloss 1961

Discussion: Tragulus was attributed to Brisson (1762) by many authors (A. Gentry, 1994) though some (e.g. Chasen, 1940; Lydekker, 1915) assigned it to Pallas (1779). Recent rejection of Brisson (1762) was on the assumption that it was unavailable. Brisson (1762:65-68) listed the species T. indicus , T. guineensis (= Neotragus pygmaeus ), T. surinamensis (= Mazama americana ), T. africanus (= Sylvicapra grimmia ), and [ T.] moschus (= Moschus moschiferus ). Of these, only T. indicus is referred to Tragulus as currently used, according to A. Gentry (1994:141). Brisson's Tragulus was defined by lacking horns (or antlers). However, three species were included on the strength of females or immatures which lack horns or antlers, while adult males possess them, so the character can not help to confirm that T. indicus is a mouse-deer. This nominal species is also based on descriptions of specimens of N. pygmaeus by Linnaeus and Seba (and secondary citations by Klein) and on Kolbe's "Chevre de Congo ". Brisson's own description is mostly undiagnostic but indicates a small animal with upperparts of head, neck and body reddish-yellow mixed with blackish; throat, belly, and inner parts of thighs whitish; and 26 teeth (so not with the whole tragulid complement of 34), including two upper canines, one on each side. The vernacular name is "Le Chevrotain des Indes". The streaked pelage suggests a species of Tragulus rather than the uniformly-coloured N. pygmaeus , and so does the presence of upper canines, as noted by A. W. Gentry (1995). Although infants of N. pygmaeus often have upper milk canines, canines are rarely present in subadults and adults and are then very small. Reference to the Indies is suggestive. This is the evidence for inferring that Brisson had studied a subadult specimen of mouse-deer, one of the syntypes of T. indicus . Tragulus indicus Brisson could be identified as a mouse-deer and the genus could apply to these mammals. Merriam (1895 c:375) designated Tragulus indicus as the type of Tragulus Brisson , but identified it as the same as " Capra pygmea Linn. , which becomes Tragulus pygmeus (Linn.) 1758" (i.e. N. pygmaeus ), probably for reasons suggested by A. W. Gentry (1995). He did not designate a lectotype for T. indicus that was N. pygmaeus (for instance, Linnaeus' or one of Seba's specimens) and ignored the evidence that a tragulid was among the syntypes: the species can still be regarded as a tragulid. Gardner (1995:79, 81) preferred to treat Tragulus Brisson as unavailable and to date Tragulus from Pallas (1767, fasc. 6, p. 6), but this would appear to be a nomen nudum, with Tragulus not distinguished from Moschus . It is also the only place where Pallas cited Tragulus pygmaeus (possibly a misidentification of N. pygmaeus as a tragulid), which contrary to A. Gentry (1994:140) is not the single species included in the genus by which Tragulus Pallas was made available. Elsewhere, Pallas (1779, fasc. 13, p. 28) provided a diagnosis of Tragulus and in a footnote referred to T. ceylonensis , nomen nudum, and T. javensis , a name validated by a reference to Pallas' earlier description of a tragulid from Java (Pallas, 1777, fasc. 12, p. 18). Hopwood (1947:534) considered Tragulus Brisson to be unavailable and by quoting selected text, denied that Pallas defined or formally adopted the genus Tragulus (he did not refer to the diagnosis in fasc. 13, p. 28). He therefore assigned Tragulus to Boddaert (1785:131), who included in the genus T. moschus (= Moschus moschiferus ), T. (= Moschiola ) meminna , and T. pygmaeus . Hopwood (1947:534) selected T. pygmaeus as the type and as this is Neotragus pygmaeus , Tragulus Boddaert would be referred to the Bovidae (A. Gentry, 1994: 141) . But because Tragulus Boddaert is a replacement for Moschus Linnaeus, 1758 and thus an objective synonym, the type of the genus must be T. moschus (= Moschus moschiferus ) according to Ellerman and Morrison-Scott (1951:350). This would have to be set aside if T. meminna = Moschus meminna Erxleben, 1777 were to be treated as the type of Tragulus , should Tragulus Brisson or Tragulus Pallas be regarded as unavailable, and should this genus be therefore referred to Boddaert ( Gardner, 1995:81). Brisson (1762) was rejected for nomenclatural purposes (but only in 1998), so it might seem that Tragulus Pallas, 1779:29 is now the first available designation of the genus; however, Tragulus Brisson, 1762 has been conserved (Opinion 1894, International Commission on Zoological Nomenclature, 1998), with Cervus javanicus Osbeck, 1765 as type species. In the same Opinion, Tragulus Pallas, 1767 (though not Tragulus Pallas, 1779 ) and Tragulus Boddaert, 1785 were placed on the Official Index of Rejected and Invalid Generic Names in Zoology. It has long been assumed that Tragulus includes only two species, T. javanicus ( T. kanchil in the older literature) and T. napu (mistakenly given the name T. javanicus in the older literature); see Van Bemmel (1949 b). A revision by Meijaard and Groves (2004) recognised six species in three species-groups: napu group (including T. napu and T. nigricans ), versicolor group (monotypic) and javanicus group (including T. javanicus , T. kanchil , and T. williamsoni ).