Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The Lin- nean Society of London, 2004? 2004 1401 63102 Original Article E. MEIJAARD and C. P. GROVESA REVISION OF TRAGULUS Zoological Journal of the Linnean Society, 2004, 140, 63–102. With 23 figures A taxonomic revision of the Tragulus mouse-deer (Artiodactyla) E. MEIJAARD* and C. P. GROVES School of Archaeology and Anthropology, Faculty of Arts, A.D. Hope Building, Australian National University, 0200 ACT, Australia Received October 2002; accepted for publication July 2003 The taxonomy of South-East Asian mouse-deer (Tragulus) is complex, and after some 120 years of considerable tax- onomic revisions of the genus a clear key is still lacking for the determination of species and subspecies. Through craniometrical analysis of 338 skulls of Tragulus and some study of coat coloration patterns we have come to a better understanding of mouse-deer taxonomy. Our results show that there are three species groups: the T. javanicus- group, the T. napu-group, and T. versicolor. Within the T. javanicus-group we recognize three species: T. javanicus (from Java), T. williamsoni (from northern Thailand and possibly southern China), and T. kanchil (from the rest of the range), and within these species we provisionally recognize 16 subspecies. Within the T. napu-group we recognize two species: T. nigricans (from Balabac), and T. napu (from the rest of the range); within these species we provi- sionally recognize eight subspecies. T. versicolor from Nhatrang, south-east Vietnam, is distinct from the two pre- vious groups; it is, however, unclear whether this species is still extant. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 63–102. ADDITIONAL KEYWORDS: chevrotain – coat colours – craniometrics – island fauna – South-East Asia – taxonomy – Tragulidae. INTRODUCTION Smit-van Dort (1989) studied the morphological dif- ferences between the four conventionally recognized The chevrotains or mouse-deer (Tragulidae) are an species of tragulids, i.e. the African Hyemoschus ancient group of ungulates, with a shared history dat- aquaticus, Moschiola meminna, Tragulus javanicus, ing back to the Miocene. They are considered the and T. napu. She could separate the two Tragulus spe- sister-group of the remaining living Ruminantia cies only by using metrical characters in the skeleton, (Groves & Grubb, 1982). In southern Asia, they are i.e. the greater mouse-deer T. napu has stronger and found in fossil assemblages dated at 18 million years broader feet than the lesser mouse-deer T. javanicus. before present (Myr), although they reached their Skull dimensions of T. napu and T. javanicus, espe- highest diversity with five named and 52 unnamed cially in specimens from small islands, converged to species at around 11.5 Myr (Barry et al., 1991). such an extent that the two species could not be reli- Between 16 and 14 Myr, they were also common in the ably separated using skull characters (also see van area that is now northern Thailand, although species Dort, 1988). Skin patterns do differ between the two diversity may have been low (Ducrocq et al., 1994). species, with T. napu having mottled fur and light After 9 Myr, the tragulid family declined significantly superciliary lines on the head, whereas T. javanicus in diversity in southern Asia (Barry et al., 1991), normally has three strong white lines on the throat which may have been caused by the evolution of more and has uniformly coloured upperparts (Smit-van open vegetation types. Presently there are three gen- Dort, 1989). There is, however, considerable variation era left, of which two, Moschiola and Tragulus, occur within each species in the coloration and striping pat- in the South and South-East Asian region. terns, especially on the many islands where the spe- cies occur, and in several instances it is nearly impossible to determine whether a taxon is more *Corresponding author. E-mail: [email protected] closely related to T. napu or to T. javanicus. In fact, © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 63–102 63 64 E. MEIJAARD and C. P. GROVES Groves (1989) considered the two to be sibling species. We propose instead to designate Gmelin’s Moschus Based on this geographical variation, Chasen (1940), javanicus as type, which, with Pallas’ Tragulus, would the first major reviser of Tragulus, recorded 27 sub- fix the name of the Javan mouse-deer as Tragulus jav- species for T. napu and 26 for T. javanicus (some of anicus (this will be part of our submission to the which he had reallocated from one to the other) for the Commission). Malayan region, and at least ten more subspecies, pri- Opinions as to the identity of javanicus, whether marily from mainland Asia, have been described in dated from Osbeck or Gmelin, have fluctuated other publications. The determination of this plethora between authors, from the large (napu group) to the of subspecies has been based primarily on variation in small (kanchil group) and back. Raffles (1822), the skin patterns, but when, during preliminary research, first to mention the existence of two (or even three) we investigated skins of Tragulus we realized that the species of South-East Asian mouse-deer, called the characteristics of the subspecies reported by Chasen larger species Moschus javanicus Pallas and named (1940) and others were not always consistent, and the smaller M. kanchil, mentioning in addition a there appeared to be considerable morphological over- potential third species, ‘the Pelandok’, based on anec- lap between subspecies. Also, some taxa that Chasen dotal information. His specimens were from the had assigned to subspecies appeared to be distinct Bengkulu district of Sumatra. His descriptions of the enough to qualify as species. We therefore decided to two named species are accurate, and recognizably focus on a craniometrical analysis, because the taxon- those of the two that occur today sympatrically on omists who worked on the genus had paid relatively Sumatra. Hamilton-Smith (1827) named Raffles’ little attention to skull form. Pelandok Moschus pelandoc, although his rather In this research we investigated skulls of Tragulus confused description suggests that he thought his to obtain a more transparent and consistent taxonomy specimen to be the same as Raffles’ ‘Napu’. Miller for the genus. In addition we studied some skins, but (1903b) finally rejected all descriptions of because many type specimens especially of small M. javanicus, T. javanicus, and T. pelandoc, and island taxa are located in the Smithsonian Museum, instead proposed a new species, T. focalinus, to desig- which we did not visit, we were unable to assess the nate the species from Java. validity of all taxa. We intend to address the variation Gray (1836) recognized Moschus javanicus Gmelin between these island taxa, and also variation within (with M. napu F.Cuvier a synonym), Moschus kanchil M. meminna in future publications. Raffles, and two new species: Moschus fulviventer (which he thought is probably Raffles’ ‘pelandok’) and Moschus stanleyanus. After these initial descriptions TAXONOMIC HISTORY OF THE TRAGULIDAE several intermediate revisions appeared. In a later The earliest scientific name generally considered publication, Gray (1843) dropped the genus Moschus, applicable to a tragulid is Cervus javanicus Osbeck included it in the Bovidae, and referred the South- (1757), described from Nieu Bay, Ujung Kulon, Java. East Asian mouse-deer to Tragulus; he also introduced We are not convinced that this name actually applies the name Tragulus affinis from Malacca. Blyth (1858) to a tragulid at all; he may have been describing a suggested that four (if not five) species of Tragulus juvenile cervid, perhaps Muntiacus muntjak. As this were distinguishable. Milne-Edwards (1864) wrote a will be the subject of a future application to the Inter- long review of the Tragulidae, recognizing Tragulus national Commission on Zoological Nomenclature, we javanicus, T. napu, T. kanchil and T. stanleyanus will not argue the case here. (as well as South Asian T. meminna). Gray (1872) Pallas (1777) gave what we consider to be the first changed his species listing yet again and recognized description of a specimen of a mouse-deer that he had T. stanleyanus, T. javanicus (of which he maintained obtained from Java; he did not name it, but Gmelin that T. napu of Milne-Edwards was the adult, and his (1788) considered it a new species of musk-deer (Mos- T. javanicus the young), T. fulviventer (= Tragulus chus), and on the basis of Pallas’ description named it kanchil Milne-Edwards, 1864), and Tragulus pyg- Moschus javanicus (see Appendix 1). Pallas’ descrip- maeus. One year later, Gray (1873) dropped the last tion not only clearly refers to a mouse-deer, but his species again. mention of a grey nape confirms that the specimen The late 19th and early 20th centuries were a time was from Java, as no other mouse-deer species has of prolixity in the taxonomy of Tragulus, when zoolo- this characteristic (e.g. Miller, 1903a). Pallas also cited gists such as Miller, Lyon, Lydekker, Kloss, Sody, Bon- Brisson’s (1762) generic name Tragulus, which had hote and Chasen described numerous species and been substituted for Linnaeus’ Moschus; as Gentry subspecies. They differed in their opinions on how spe- (1994) showed, the type of this genus is pygmaea, but cies and subspecies should be defined. For instance, she proposed, following Ellerman & Morrison-Scott Miller (1911) referred his new taxa to subspecies when (1951), to designate Cervus javanicus Osbeck as type. there was some overlap in skin characteristics © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 63–102 A REVISION OF TRAGULUS 65 between the specimens from two geographical groups, paraocciptal processes (PPW) (not shown in Fig.