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The mouse-deer or chevrotains are the smallest of the ungulates and ruminants. They are characterized by a number of traits which are considered plesiomorphic for the Artiodactyla order. The objective of this study was to examine sleep in the lesser mouse-deer (Tragulus kanchil), which is the smallest in this group (body mass < 2.2 kg). Electroencephalogram, nuchal electromyogram, electrooculogram, and body acceleration were recorded in four adult mouse-deer females using a telemetry system in Bu Gia Map National Park in Vietnam. The mouse-deer spent on average 49.7 ± 3.0% of 24 h in non-rapid eye movement (NREM) sleep. REM sleep occupied 1.7 ± 0.3% of 24 h or 3.2 ± 0.5% of total sleep time. The average duration of REM sleep episodes was 2.0 ± 0.2 min, the average maximum was 5.1 ± 1.1 min, and the longest episodes lasted 8 min. NREM sleep occurred in sternal recumbency with the head held above the ground while 64.7 ± 6.4% of REM sleep occurred with the head resting on the ground. The eyes were open throughout most of the NREM sleep period. The mouse-deer displayed polyphasic sleep and crepuscular peaks in activity (04:00–06:00 and 18:00–19:00). The largest amounts of NREM occurred in the morning (06:00–09:00) and the smallest before dusk (at 04:00–06:00). REM sleep occurred throughout most of the daylight hours (08:00–16:00) and in the first half of the night (19:00–02:00). We suggest that the pattern and timing of sleep in the lesser mouse-deer is adapted to the survival of a small herbivorous animal, subject to predation, living in high environmental temperatures in the tropical forest undergrowth.

New remains of the small tragulid Dorcatherium songhorensis Whitworth, 1958 from the Early Miocene fossil site of Napak XXI (Uganda) include the first significant sample of postcranial bones from this species ever described. The limb bones of this tragulid are very similar to that described in the Miocene Asian long-legged tragulids of the genus Siamotragulus Thomas et al., 1990, a type previously unknown in the African Miocene. A cladistic analysis links D. songhorensis to a Siamotragulus clade as its basal offshoot, so we propose the name Siamotragulus songhorensis (Whitworth, 1958) for this species. Also, the Siamotragulus clade belongs to a monophyletic group that includes Afrotragulus Sánchez et al., 2010 and the extant Asian genera Moschiola and Tragulus. This inclusive clade is characterized by both a derived selenodont dentition and an advanced postcranial skeleton. Additionally Siamotragulus shows some cursorial refinements reflected in its postcranial skeleton including the pecoran-like metatarsals III–IV. Siamotragulus songhorensis shows that the genus Siamotragulus was not endemic to Asia as previously thought, and that a highly diverse guild of tragulids, including different members of the advanced selenodont clade, inhabited Africa as early as the Early Miocene (19–20 Ma)., This study is part of the research project CGL2011-25754 (Spanish Government, MINECO), and the Research Group CAM-UCM 910607.

4. Silver-backed Chevrotain Tragulus versicolor French: Chevrotain du Vietnam / German: Annam-Kantschil / Spanish: Ciervo ratdn plateado Taxonomy. Tragulus versicolor Thomas, 1910, Nhatrang, Annam, Viet Nam. This species 1 monotypic. Distribution. If still extant, the species would likely occur in Vietnam, and possibly in neighboring Cambodia and Laos. The true distribution of the species is unknown, because there has been very little awareness of the species. Surveys have not sought it, because many have assumed that only one species, Tragulus kanchil, occurs in most of Indochina. Descriptive notes. Head-body 40 cm, tail 5 cm, hindfoot 11 cm, ear 3-5 cm; weight 1-7 kg (all measurements from one specimen). The upperparts of 7. versicolor have a characteristic shabrack or saddlecloth appearance. The fur on the back is uniformly grayish-silver and relatively dense; the hairs are conspicuously tipped with white. The area of the neck and shoulders, although quite uniform in color, is visibly separated from the back in having more brownish and less dense fur. Thesides of the neck possess a somewhat different pattern of pelage structure. The fur of the neck appears to have a coarser, less spiny appearance compared to the sympatric 7. kanchil. This coarse hair was one of the characteristics used to describe the species initially. There are no dark lines from the eye to the nose. Some specimens have a throat pattern with three white and two colored stripes, which are characters of 7. kanchil, whereas there are others with the napwlike five white and four dark stripes. The white area of the underparts in 1. versicolor continues uninterrupted from the chin towards the groin area, and there is no transverse brownish collar below the throat, unlike most other Tragulus species. Also, there are no darker-colored markings along the sides and midlines of the belly, which are characteristic of 71. kanchilin Vietnam. The skull of 7. versicolor is significantly smaller than in all other Tragulus species, but relatively wide, with wide bullae and very long nasals. Compared to T. napu and T. kanchil, the large auditory bullae are much more rounded and inflated. Dental formulais10/3,C1/1,P 3/3, M 3/3 (x2)= 34. Habitat. Because no specimens have been traced to specific habitats, and there are no observations of the species in the wild, nothing is known about its habitat preferences. Food and Feeding. Nothing known. Breeding. Nothing known. Activity patterns. Nothing known. Movements, Home range and Social organization. Nothing known. Status and Conservation. Classified as Data Deficient on The IUCN Red List. There is no information on the current range or population status of 7. versicolor because of a lack of survey work for the species in appropriate areas of Vietnam. Therefore the species cannot be assessed against range size or population trend criteria. No strong ecological correlates are sufficiently known to allow assessment of population trends through habitat trends. The species’ resilience to hunting is not known. Although there is only one record since 1907, there is no reason to consider it a possibly Extinct candidate, because the paucity of records is assumed to reflect the lack of suitable surveys. A more informed judgment when sufficient data are available is likely to indicate that it is threatened. The population is almost certainly declining due to severe hunting pressure for ground-dwelling mammals, resulting in population reductions of even the most resilient species; it may be found to warrant a very high threat category, particularly as there is no evidence of presence in high altitudes. Bibliography. Kuznetsov & Borissenko (2004), Meijaard & Groves (2004a, 2004b), Meijaard et al. (2004), Thomas (1910), Timmins et al. (2008)., Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Tragulidae, pp. 320-334 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on page 330, DOI: 10.5281/zenodo.5721279

5. Lesser Indo-Malayan Chevrotain Tragulus kanchil French: Chevrotain kanchil / German: Kleinkantschil / Spanish: Ciervo ratén pequeno Other common names: Kancil Taxonomy. Moschus kanchil Raffles, 1822, Bengkulu, Sumatra, Indonesia. Lesser Indo-Malayan Chevrotains are highly variable in coloration, especially the taxa from small islands. This has led to the description of a great number of species and subspecies. A taxonomic review in 2004 brought some clarity about the validity of the many described taxa. Still, many type specimens, especially of small island taxa, were not included in that review, and the authors were unable to assess the validity of all taxa. This includes Sfulvicollis from the Malacca Strait islands (Bengkalis, Padang, Rupat, Tebingtinggi, and Rangsam); pallidus from the small island of Laut, north of Bunguran, Natuna islands group; carimatae from Karimata Island, west of Borneo; lampensis from Lanbi Kyun (= Lampi) Island in the Mergui Archipelago; insularis from Phuket (= Junk Seylon), Ko Sirae (= Sireh) and Ko Yao Yai (= Panjang) islands; and lancavensis from Langkawi Island. Sixteen subspecies presently recognized. Subspecies and Distribution. T.k.kanchilRaffles,1822—Sumatra,islandsoffESumatra(Mendol&Berhala). T.k.abruptusChasen,1935—SubiI,oftWBorneo. T.k.affinisGray,1861—Vietnam,Laos,SE&EThailand,Cambodia. T.k.anambensisChasen&Kloss,1928—AnambasArchipelago(MatakI). T.k.angustiaeKloss,1918—SMyanmar,SWThaimainland(probablylimitedtoWoftheChaoPhrayaRiver). T.k.everettiBonhote,1903—NatunaIs(Bunguran),offWBorneo. T.k.fulviventerGray,1836—SMalayPeninsula(Sof7°N). T.k.hosetBonhote,1903—Borneo(Sarawak,West,Central,East&SouthKalimantan). T.k.klossiChasen,1935—NBorneo(NEastKalimantan,E&CSabah,andpossiblyWSabahandBrunei. T.k.luteicollisLyon,1906—BangkaI,offESumatra. T.k.pidonisChasen,1940—KohPipidonI(=PhiPhiDon),offWMalayPeninsula. T.k.ravulusMiller,1903—islandsoffWMalayPeninsula(KohAdang&KohRawi). T.k.ravusMiller,1902—SThailand,NMalayPeninsula. T.k.rubeusMiller,1903—RiauArchipelago(BintanI). T.k.siantanicusChasen&Kloss,1928—AnambasArchipelago(SiantanI). T. k. subrufus Miller, 1903 — Lingga Archipelago (Lingga & Singkep Is). The range on the Asian mainland is poorly known and could occur as far north as China (S Yunnan). As stated in the Taxonomy section, the subspecific status of the populations of some areas of Borneo (W Sabah, Brunei & N Sarawak) and several other islands remains unclear. Descriptive notes. Head—body 37-56 cm, tail 6-9 cm; weight 1.5-2. 5 kg. Thisis a small chevrotain species that was recently split from 7. javanicus. It is separated from the larger 1. napu primarily by its smaller size, the number of throat stripes, the visibility of the nape line, and the lack of mottling of the upperparts. There is some degree of melanism in 7. kanchil specimens from small islands, leading to the development of extra dark, transverse lines, which makes the throat pattern hard to distinguish from that of 1. napu. The degree of erythrism (reddish pigmentation) and melanism is, however, less pronounced than in 7. napu. Dental formulais10/3,C1/1,P 3/3, M 3/3 (x2)= 34. Habitat. Tall forest in lowlands, amidst undergrowth on edges of heavy lowland forest. It has been suggested that 7. kanchil occurs primarily in hilly areas, but other sources stated that the species was absent from areas above 250 m in Sarawak, whereas 7. napu occurred much higher. This species also occurs in cultivated areas up to 600 m in elevation. The habitat of this species could be described as a mosaic ofriverine, seasonal swamp and dry undulating country, vegetated predominantly by legumes and dipterocarps, with stands of dense bamboo or palms for daytime resting. In Sabah, they also inhabit mangrove forest, and they can be quite common in monocultural tree plantations in a matrix of secondary forest stands. Even though the evidence is somewhat ambiguous,it appears that this species prefers disturbed forests to primary ones. A review of encounter rates in various parts of the species’ range suggested that the distribution of 7. kanchil is perhaps highly patchy and correlated to specific habitat features and microhabitats. What these features are remains unresolved, but their water requirements are important, as authors note commonness in riverine areas with surprising regularity. The habitat use of this species with respect to edge—interior areas shows startling heterogeneity, at least in non-Sundaic areas, but this cannot yet be explained; in particular, disentangling the contributions of intrinsic habitat suitability and the effects of hunting is difficult. Food and Feeding. The speciesis largely frugivorous, but also feeds on shoots, young leaves, and fallen fruits. Fruit mass of consumed fruits varied from 1 g to 5 g and seed mass from 0-01 g to 0-5 g. On Borneo, the species feeds on a range offruiting species, including Polyanthia sumatrana, Diospyros macrophylla, Endospermum peltatum, Quercus sp., Garcinia forbesi and G. parviflora, Litsea caulocarpa and L. orocola, Notaphoebe sp., Dialum imdum, Aglaia sp., Chisocheton sp., Dysoxylum sp., Lansium sp., Artocarpus dadah, Ficus spp., Dimocarpus longan, Paranephelium xestophyllum, and Microros antidesmifolia. Breeding. Males mark their territories and their females with an intermandibular scent gland located under the chin. They chase and fight one another for prolonged periods, slashing with the elongated canine teeth. These teeth are razor sharp and local hunters in Borneo report that sometimes males fight to the death. Mating occurs throughout the year in some areas. In Vietnam the species mates in November-December and gives birth in April-May. The species is polygynous. Females are almost continuously pregnant. Gestation has been estimated at 140-177 days, the mother produces 2-3 young per year, and fawns are kept hidden. Activity patterns. Diurnal to cathemeral. Movements, Home range and Social organization. A study of this species in the Malaysian state of Sabah suggested thatit is mostly solitary, with 93-9% of the observations being single animals and the remainder pairs or a female with one or two young. In apparent contrast with this statement, camera trap photos of the species often show two adult animals together. Further studies are needed to determine the social organization of the species and how this varies spatially as well temporally. Population densities on Borneo were estimated at 21-39 ind/km?, with density positively correlated with fruit mass, seed mass, and total fruit resources. The species does not appearto be territorial. The core areas of neighboring animals were completely separate among same-sex individuals, but overlapped widely among opposite-sex individuals. The results suggest that this species is mostly monogamous, although apparently males can also be polygamous. The core area of a paired female overlapped not only with the core area of the paired male, but also with that of another neighboring male. This suggests that males tolerate the presence of females in their core areas and that paired males do not control the movement of paired females into the home ranges of other males. Females establish new home ranges when giving birth. Home range size for females was estimated at 4-4 ha and for males 5-9 ha using the minimum convex polygon method, but the differ ence between males and females was not significant. Mean daily distance travelled for males was 519-1 m (+ 88-8 m), that for females 573-8 m (+ 219-7 m). In a Bornean study area, the density was negatively correlated with pioneer trees, grass, and herbs, suggesting that the species is negatively influenced by the effects of timber harvest, although, as pointed at above, this remains ambiguous. Data from Indochina, suggest that the species is quite tolerant of forest disturbance and may even be considered an edge species that benefits from disturbance. Further studies are needed to elucidate this issue. Status and Conservation. Classified as Least Concern on The IUCN Red List, because it remains widespread and locally common, and at least in non-Sundaic areas persists in environments of very heavy forest degradation, fragmentation, and hunting. The presumed short generation length of the species (under five years) also influences assessment. Thus although there may be or may have been drastic local reductions, these have probably not been synchronous over a large enough area. The IUCN review points out several major uncertainties in the conservation status assessment. Firstly, there are few modern records of Tragulus from both the Sundaic and non-Sundaic portions of the range that have been identified conclusively as to species. Secondly, the conflicting nature of the information available concerning the effects of hunting (harvest levels are locally very high) and habitat destruction makes it difficult to estimate population declines. Thirdly, there are strong indications that in its non-Sundaic range (i.e. Asian mainland) it is localized in occurrence, a pattern for which the reasons remain opaque, but which might be the result of hunting. And fourthly, the apparent restriction to lowland forest, at least in Borneo, suggests that with the rapidly dwindling lowland forests in this part ofits range, the species is losing habitat and its range might decrease and become fragmented. The species apparently has become extinct in Bangladesh due to high hunting and trapping pressure, although in many other areas it seems to survive despite local poaching and trapping. Bibliography. Caldecott (1988), Davison (1980), Duckworth (1997), Duckworth & Timmins (2008), Endo (2004), Heydon (1994), Heydon & Bulloh (1997), Kim et al. (2004), Liat (1973), Matsubayashi & Sukor (2005), Matsubayashi et al. (2003, 2006), Medway (1978), Meijaard (2003), Meijaard & Groves (2004a, 2004b), Meijaard et al. (2005), Miura & Idris (1999), Nolan et al. (1995), O'Brien et al. (2003), Payne et al. (1985)., Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Tragulidae, pp. 320-334 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on pages 330-331, DOI: 10.5281/zenodo.5721279

8. Greater Indo-Malayan Chevrotain Tragulus napu French: Chevrotain napu / German: Grofskantschil / Spanish: Ciervo raton grande Taxonomy. Moschus napu F. Cuvier, 1822, Sumatra, Indonesia. Restricted by Sody in 1931 to southern Sumatra. Greater Indo-Malayan Chevrotains are highly variable in coloration, especially the taxa from small islands. This has led to the description of a great number of species and subspecies. A taxonomic review in 2004 brought some clarity about the validity of the many described taxa. Still, many type specimens, especially of small island taxa, were not included in that review and the authors were unable to assess the validity ofall taxa. In addition to the subspecies below several other taxa have been described for which the taxonomic status remains unresolved. This includes nigrocinctus from Malacca Strait islands (Kundur and Karimun Besar = Great Karimon); bullitonus from Belitung Island (which might be a subspecies of 1. kanchil); amoenus from Musala Island; and several subspecies from island taxa west of Sumatra, including brevipes from Bangkaru Island, russulus and batuanus from Tanahbala and Tanahmasa islands, pinius from Pini Island. All these taxa require further study to determine their taxonomic status. Seven subspecies presently recognized. Subspecies and Distribution. T.n.napuF.Cuvier,1822—SMyanmar,Thai/MalayPeninsula,islandsoffWMalayPeninsula(Langkawi&Pangkor),Borneo,SSumatra,BangkaI,islandsoffBorneo(Laut&Serasan). T.n.bangue:Chasen&Kloss,1931—BanggiIandBalembanganI,offNBorneo. T.n.bunguranensisMiller,1901—NatunaIs(=Bunguran),oftWBorneo. T.n.neubronneriSody,1931—NSumatra. T.n.nmiasisLyon,1916—NiasI,offWSumatra. T.n.rufulusMiller,1900—TiomanI,offEMalayPeninsula,RiauandLinggaArchipelagos. T. n. terutus Thomas & Wroughton, 1909 — Terutau I, off W Malay Peninsula. The species was recently reconfirmed for Singapore. Maps that include Vietnam, Cambodia, and Laos in the distribution range are based on the earlier assumption that 7. versicolor was a subspecies of 1. napu. Subsequent studies have indicated that 7. versicolor is a distinct species, and that the range of 1. napu therefore does not extend into Cambodia, Laos, and Vietnam. The northern limit on the Thai-Malay peninsula is not well defined. Specimens of 1. napu have been collected from as far north as Bankachon in southern Myanmar (10° 08 ” N), but despite fairly intensive camera-trapping in Kui Buri National Park, Thailand (12° N), 7. napu has not been photographed there. At the northern margin ofits range, it is generally rare. It has been reported, for example, that during the flooding of the Chiew Larn Reservoir (Surat Thani Province; about 9° N, 98° 45’ E), only six 7. napu were rescued compared with 172 71. kanchil. This area is the transition zone from wetter evergreen forest to drier deciduous types, and it might be that 7° napu is not well adapted to the drier forest types towards the northern limit ofits range. There are unconfirmed reports of the species on Java, where it may have been confused with one of the two color morphs of 7. javanicus. As explained in the Taxonomy section, the subspecific status of the populations of several islands remains unclear. Descriptive notes. Head-body 42:5-68 cm, tail 8-5 cm, shoulder height 30-35 cm; weight 3-5—4-5 kg. 1. napu is one of the largest Asian chevrotains. The pelage is orangeybrown in color; the hindquarters are lightly mottled with black. The underparts and legs are paler. The underside of the chin is white, and there are a series of white markings on the neck. The head is triangular in shape, and the large black nose is naked. The eyes are very large, and the ears are medium-sized, lightly covered with black hair. The body is rounded, and the rear quarters are higher than the front. The legs are extremely thin and delicate, being about the diameter of a pencil. Compared to the partly sympatric 1. kanchil, skin patterns of 7. napu are more mottled and 7. napu has pale lines on the throat and neck, whereas 7. kanchilnormally has three strong white lines on the throat and has uniformly colored upperparts. There is, however, considerable variation within each species in the coloration and striping patterns (also see other Tragulus species accounts), especially on the many small islands where the species occur. In several instances it is nearly impossible to determine whether a taxon is more closely related to 7. napu or to T. kanchil. In fact, until an in-depth taxonomic review was recently conducted, 7! napu was sometimes considered to be a sibling species of 1. kanchil. The pelage coloration in 7. napu is more variable than in 7 kanchil, with varying degrees of melanism and erythrism. Specimens from the larger islands (Borneo and Sumatra) and the Asian mainland and Malay Peninsula are generally dull yellowish-brown, with a mottled appearance. On several islands, 7. napu typically has a red-brown color. Even the nape-stripe is red-brown, somewhat darker than the orange-red neck. The lines from the eye to nose are deep black, and the red-brown back is mixed with black. From studies of hair and skins and from information in the literature it appears that increased redness is the result of fewer all-black and black-tipped hairs in the coat and possibly also an actual increased redness in the hair pigment. The color pattern of the primitive type of 1. napu on the mainland can be summarized as follows: a mixed brown and black neck, the black concentrating along the nape to form an evident napestripe; and a normal throat pattern with a median white longitudinal stripe, on each side of which is a similar stripe, the three meeting in a broad white mass; the space between median and lateral stripes is brown, as in the side of the neck, or somewhat darker; a brown transverse band or collar separates the stripes from the white of the chest. The two lines of variation are toward predominance of yellowish-brown or toward predominance of black. This species was readily differentiated from members of the 1. javanicus / kanchil group by its larger size, relatively small auditory bullae and, in most specimens, the distinct coat coloration. The analysis of skull measurements suggested that within this species there are three geographical groups, none of which is distinct enough to be assigned to species level. Dental formula: 10/3, C1/1,P 3/3,M 3/3 (x2)= 34. Habitat. Primary and secondary evergreen rain forest. It has been suggested that the foot morphology of 7. napu indicates a preference for more swampy habitats than 7. javanicus / 1. kanchil. Another source, from Borneo, however, claims that 7. napu is a hill species, found at no great elevation but never on swamps, plains and less often on river banks than 7. kanchil. T. napu seems to be scarcer in logged forest than in primary forest, with an 86% decline in density being reported following timber harvest. Daily home range use also appears to be more extensive in secondary forest. The sensitivity to logging is probably a consequence of diet constraints that limit the species’ ability to compensate for the reduced abundance of selected fruits in disturbed forest by utilizing browsing. In twelve-year-old logged forest, fewer trees remained belonging to genera eaten by 7. napu. On Tioman Island, of the coast of the Malay Peninsula, densities increased away from human settlements probably in direct relationship to hunting pressure; also 83% of observations on this island were in primary forests and 17% in secondary forests or rubber estates. Food and Feeding. This species is mostly frugivorous, although few details on feeding are known. In captivity they eat most fruits and readily take to bananas, though their favorite food was mentioned to be the fruits and particularly the flowers of “Buah Simpoh” (presumably a Dillenia species). It has been asserted that 7. napu are prone to kill chickens, but no direct evidence for this was found. Breeding. The species is possibly polygynous; there is no distinct mating season and females give birth throughout the year in Peninsular Malaysia; one or rarely two young are born. The gestation period is 152-155 days. Weaning occurs at 2-3 months. The young reach sexual maturity at 4-5 months, and maximum life span in captivity is reported to be up to 14 years. 1. napu was observed to stamp with one or both hindfeet when slightly alarmed. Other individuals would or would not stamp in response. Both males and females mark objects with the intermandibular gland. Males mark much more frequently than females. Males often lick the urine of females; less frequently, females lick the urine of males. Males court both receptive and unreceptive females. Courting males mark the female on the back or rump with the intermandibular gland and emit a series of squeaks. Females have a postpartum estrus and return to estrus at approximately 14day intervals unless they become pregnant. Females of 7. napu have the potential to be pregnant throughout adult life, often with only 85-155 minutes passing between giving birth and becoming pregnant again. The young are born fully developed, and they are able to stand after 30 minutes. Mothers spend little time with infants. They emit a vocalization that sounds like the squeak of courting males. Both mothers and infants emit a higher-pitched vocalization. Intense agonistic behavior was seen only when strange individuals were introduced into established groups. Males fight by facing each other and biting each other on the ears, neck, and shoulders with their large upper canines. Fighting males usually hold the tail in a vertical position, exposing the white ventral surface, and may emit loud growls. If one male flees, the other pursues and attempts to bite him on the neck and body. Activity patterns. 7. napu is reported to be cathemeral (active day and night), but there are few data to support this. Movements, Home range and Social organization. The density of 7. napuin one Borneo site was positively correlated with large strangler fig trees (Ficus spp.) and negatively correlated with grass and herbs and vine tangle. In primary forest densities vary from 37-72 ind/km?®. On Tioman Island, densities ranged from 27-8 ind/km? to 312-5 ind/ km? The home range in primary forest is about 7 ha. Home range and core area size do not differ according to sex, but are determined by body size of the individuals. The species is mostly solitary, with a mean group size of 1-37 individuals. Recent observations indicate that the species escapes predators by hiding under water in forest streams. Status and Conservation. Classified as Least Concern on The IUCN Red List, because it remains widespread and at least locally common in the Sundaic region and, at least on Borneo, it appears that this species is resilient to hunting, although probably not to habitat degradation. Better data would allow a more confident assessment. Its wide altitudinal tolerance (presumed to be up to 1000 m above sea level) means that it has large populations above the elevation where very rapid forest loss is occurring. The presumed short generation length of the species, which is likely underfive years, also influences assessment, because it means that the species could quickly recover from local declines. Thus, although there may be or may have been drastic local reductions, these have probably not been synchronous over a large enough area to threaten its survival. Bibliography. Azlan & Engkamat (2006), Chasen (1940), Chua et al. (2009), Dobroruka (1972), van Dort (1986, 1988), Duckworth (1994, 1997), Heydon (1994), Heydon & Bulloh (1997), Matsubayashi & Sukor (2005), Meijaard & Groves (2004a, 2004b), Meijaard et al. (2010), Miura & Idris (1999), Nakhasathien (1989), Payne et al. (1985), Ralls et al. (1975), Smit-van Dort (1989), Timmins et al. (2008), Yasuma (1994, 1996), Yasuma & Andau (2000)., Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Tragulidae, pp. 320-334 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on pages 332-333, DOI: 10.5281/zenodo.5721279

9. Balabac Chevrotain Tragulus nigricans French: Chevrotain de Balabac / German: Balabac-Kantschil / Spanish: Ciervo ratén de Balabac Other common names: Pilandok Taxonomy. Tragulus nigricans Thomas, 1892, Balabac, Philippines. This species was resurrected as a distinct species in 2004 based on skull morphology. 1. nigricans was originally allied with 1" napu due to similarities in body size and markings. Skull measurements demonstrate clear differentiation between T. nigricans, 1. napu, and 1. kanchil, the species from nearby Borneo and surrounding small islands. 7. nigricans is a restricted area species endemic to the Philippines, and there are no subspecies. It has been suggested that the chevrotain population of Pulau Banggi (Malaysia), which lies midway between Balabac and the Bornean mainland, and which was almost certainly (and perhaps repeatedly) connected to both by former (late Pleistocene) land bridges, might belong to this species. Recent morphometric study of specimens from Pulau Banggi indicates its distinctiveness from both 7. napu and 1. nigricans, suggesting that morphometrically the Pulau Banggi population is more closely related to 7. kanchil. Throat patterns and coloration of the upperparts, however, suggests a napurelated taxon, and for now the subspecies T. napu banguei is retained for the Banggi population. Monotypic. Distribution. Balabac, Ramos, and Bugsuk Is, Palawan region, Phillipines. In the late 1990s, a small stock of eight Balabac Chevrotains escaped from their enclosures on Calauit Island, a small island off the coast of Busuanga Island, north of Palawan, where the species had been maintained and bred since 1982. These animals were reported to have increased to at least 21 free-living individuals by 2006. Apparently the species was also introduced to the larger island of Palawan. No recent records confirm that it is still extant there, although unconfirmed reports suggest that it may survive in southern Palawan. Descriptive notes. Head-body 40-47. 5 cm, tail 6.5-8. 5 cm, shoulder height 18 cm; weight unknown. The Balabac Chevrotain is very small in stature, and smaller than its apparent close relative 71. napu from neighboring Borneo. T. nigricans is one of the most darkly-colored chevrotains, with an overall dark brown coat. The upperparts are broadly washed with black, and the neck is mixed black and fulvous. Each individual hair has sections of different colors. The base is generally pale, ranging from white to ashy brown, the mid-section is tawny, orange, or brown, and there is a long black tip. The abdominal region is generally orange-brown in color. The lower abdomen, groin, and insides of the thighs are white. The hindlegs have a white stripe on the front that extends from the upper thigh to the tarsal joint; the rest of the leg is fulvous in color. The inner surfaces of the front legs are white, and this coloration extends across the chest in between the front legs as a whitish patch. The tail is dark brown with a white tip. The most striking markings of the Balabac Chevrotain are on the throat, with three narrow white stripes beginning from a white patch under the chin and extending down towards the chest. In contrast to these white stripes, the rest of the throat is jet black; in some specimens the black coloration even overtakes and obscures the stripes. Towards the chest, these black and white markings disappear into a broad brown band that crosses the lower throat. The head itself is generally darker in color than the rest of the body. Broad rufous or fulvous eyebrow stripes extend from the anterior corners of the eyes to the base of ears. The bridge of the nose and forehead are dark brown, becoming increasingly infused with black towards the crown of the head. The sides of the head are more fulvous. A naked glandular patch on underside of the jaw is bordered with white, which runs into the white patch at the top of the throat. Dental formula is 10/3,C1/1,P3/3,M 3/3 (x2)= 34. Habitat. 7. nigricans inhabits both primary and secondary forests as well as shrubland on Balabac Island, using both large blocks of forest and smaller fragments. It may frequent mangroves and more open areas to forage. Dense vegetation within the forest interior provides shelter for this species during the daytime. Food and Feeding. It is reported that 7. nigricans feeds mostly on leaves, which would set it aside from most other chevrotain species, which are primarily frugivorous. One photograph of a captive animal shows it feeding on grass-like leaves, which would confirm the above observation. Breeding. There is apparently a distinct breeding season, but it is unclear when this occurs and whether it coincides with the distinct dry and wet seasons in the species’ range. Litter size is reportedly almost always one, rarely two. Activity patterns. Reported to be nocturnal. These chevrotains leave their shaded resting spots at night, entering into clearings to feed. Most encounters between humans and the Balabac Chevrotain occur along roads at night, when their eye shine is frequently observed from vehicles. Movements, Home range and Social organization. Nothing known. Status and Conservation. Classified as Endangered on The IUCN Red List, and is probably the most range-restricted of all chevrotain species. The species is subject to poaching for food, using guns or snares, and there is minor domestic trade in live animals for local zoos and private collectors. The species is also affected by habitat loss due to conversion of former habitat to coconut plantations and other agriculture. Bibliography. Allen & White (1910), Huffman (2009), Meijaard & Groves (2004a, 2004b), Oliver (1992), Oliver et al. (2008), Rabor (1977), Rico & Oliver (2008), Sanborn (1952), Timm & Birney (1980)., Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Tragulidae, pp. 320-334 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on page 333, DOI: 10.5281/zenodo.5721279

6. Northern Chevrotain Tragulus williamsoni French: Chevrotain de Williamson / German: Thailand Kantschil / Spanish: Ciervo ratén de Tailandia Taxonomy. Tragulus williamsoni Kloss, 1916, Meh Lem, Thailand. The recent reallocation of 1. williamsoni to species level was based on one specimen from northern Thailand, because the skull dimensions exceeded those of any other T. kanchitlike chevrotains from the Asian mainland. Recently, two additional specimens from Mengla, Yunnan, China have been identified that appear to support the species-level designation of this taxon. This, however, requires further careful study, because even though the two Chinese skulls are similar in size to the specimen from northern Thailand, verification of the measurements is still required. The taxonomic uncertainty of 71. williamsoni is compounded by the lack of available specimens of chevrotains from close to the type locality of T. williamson: with which to test for conspecificity with 7. kanchil and, if the taxon is validated, to determine whetherit is sympatric or parapatric with 7. kanchil. More material is needed for both 7. williamsoni and 1. kanchil, if it occurs there, from the northern parts of Thailand, adjacent southern China, and possibly northern Laos as well as northern Myanmar, to make an informed judgment. Monotypic. Distribution. Thailand and southern China, mainly the Sanchahe part of the Mengman region and the Darongshu of the Longmen region, both in S. Yunnan, but possibly more widespread with potential presence in Laos, Vietnam, and Myanmar. Descriptive notes. No measurements available. The type specimen of this species was described as larger than other 7. kanchil specimens from the Asian mainland, but with color patterns similar to 7. k. affinis. Photos of this type specimen show uniformly colored pale brown upperparts, with a barely visible darker nape streak; the species was described as being like 7. k. affinis but larger and with the upperparts deeper ochraceous and only moderately annulated with brown. Throat patterns are as in 7 k. affinis, with clearly defined white chin and neck stripes and a reddish-brown transverse throat band that separates the white neck and chin from the white chest (unlike in 7 versicolor, in which this dark band is discontinuous, and the white neck and chin and white chest thus connected). Dental formulais10/3; C1/1;P 3/3; M 3/3 (x2)= 34. Habitat. A recent study in Yunnan showed that the species prefers valleys along banks of rivers or streams in low altitude, where they tend to use middle and lower slopes with the favorite fruit trees Ficus hirtal, Baccaurea samiflora, Alpinia kwangsiensis, and Phrynium capitatum. These plants occur along banks ofrivers or streams. Chevrotain habitat use was mostly affected by tall tree coverage, shrub coverage, and herb coverage, and distance to water. Food and Feeding. Based on its habitat use and tree preferences in Yunnan, the diet of T. williamsoni appears to contain significant amounts offruit. Breeding. Nothing known. Activity patterns. Nothing known. Movements, Home range and Social organization. Nothing known. Status and Conservation. Classified as Data Deficient on The IUCN Red List, The species’ status cannot be assessed due to taxonomic uncertainty and a paucity of data on extent of occurrence, threats, habitat preferences and altitudinal range. Hunting is heavy in the species’ likely range and habitat perturbation is increasing there. The lack of understanding of the population-level effects on the species rule out inferential application of decline-based criteria. On the IUCN assessmentit is suggested that even modest additional information would allow categorization of the species, and thatit would likely warrant either Near Threatened or one of the threatened categories. Bibliography. Aidong Luo et al. (1999), Cao Mingl et al. (2010), Kloss (1916, 1918), Meijaard & Groves (20044, 2004b), Shi Liming & Chen Yuze (1989), Timmins et al. (2008)., Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Tragulidae, pp. 320-334 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on page 331, DOI: 10.5281/zenodo.5721279

7. Javan Chevrotain Tragulus javanicus French: Chevrotain de Java / German: Java-Kantschil / Spanish: Ciervo ratén de Java Taxonomy. Cervus javanicus Osbeck, 1765, Udjung Kulon Peninsula, Java, Indonesia. Craniometrical analysis indicates that there may be two distinct chevrotain taxa on Java. The data are, however, inconclusive. Further research, particularly on skin patterns ofJavan Chevrotain, is required. None of the specimens originally used to describe two subspecies, the gray-necked form pelandoc (from the north coast of West Java Province) and the red-necked form focalinus (from the western part ofJava and to the southern coast), were included in a recent taxonomic review of the genus. An earlier review found all kinds of forms that were intermediate between the two supposed Javan subspecies and concluded that there was probably only one taxon on Java. Monotypic. Distribution. Java. Several biogeographic reviews failed to list the genus for Bali, an island directly east of Java with strong biogeographic affinities to the latter. A sighting was, however, reported from Bali Barat National Park, in a birdwatching trip report. Given the live-animal trade of this species on Java, further records are needed to confirm whether or not there is a native population of the genus on Bali. If there is, biogeographic considerations suggest it would be most closely related to the Javan population and quite probably conspecific. Descriptive notes. Head-body 50-53 cm,tail 4-6 cm; weight 1.7-2. 1 kg. Compared to 1. kanchil, T: javanicus is distinguished by its narrow auditory bullae, narrow braincase, and longer and higher mandibles. Compared with 7. kanchil from nearby Sumatra, the rostrum is shorter and the auditory bullae are narrower. Two different taxa of chevrotain have been described from Java, a gray-necked form and a red-necked form. The gray-necked form was distinguished from all other 7. javanicuslike chevrotains by the great width and distinctness of the tawny superciliary stripes and by the grizzled gray neck, which is strikingly contrasted with the tawny body and head; in addition, the nape stripe is absent, although the dark crown sometimesslightly extends onto the nape. The red-necked form, described on the basis of five specimens in the Zoological Museum of the Humboldt University in Berlin, has an orange-brown ground color,slightly grizzled with black. All five specimens had a clear, almost black, nape stripe. The cheeks and the sides of the neck are orange-buff, as opposed to the gray in the gray-necked form. This species was described as having a remarkably long tongue, which “it brings with ease to behind its ears,” but presumably this is a generic rather than species-specific characteristic. Dental formulais 10/3; C1/1; P 3/3; M 3/3 (x2)= 34. Habitat. Preferred habitat of the Javan Chevrotain includes evergreen and semi-evergreen rainforest. The species has been reported in the drier semi-deciduous forests of eastern Java, butit seems to be rarer there than in the wetter forests of central and west Java. One habitat-use study suggested that 7. javanicus uses dense bushes and stands of Salacca zalacca palms during the day. It generally remains not far from rivers. This suggests that chevrotains there might be an “edge” species, apparently preferring areas with thick understory vegetation, such as that along riverbanks. This would not be unusual within the genus (see other Tragulus accounts). The species occurs from sea level to about 1600 m above sea level in the mountains. Food and Feeding. In captivity Javan Chevrotains refuse grass as fodder and do not feed on any grasses that grow in their cages. Instead they are kept on banana peels, jackfruit, and the left-overs of rambutan and mangosteen fruits. In captivity, they are partial to leaves of a number of Acalypha (Euphorbiaceae) species, including A. marginata and A. wilkesiana, as well as leaves of Laeucaena glauca, Ipomoea batatas, I. reptans, and Manihot utilissima. In the wild, the species reportedly feeds on fallen fruit, including wild figs (Ficus spp.), and the leaves and buds of a variety of plants. Breeding. Males ofthis species are reported to produce unusual screams during the mating season. In captivity they have been observed to fight to the death. With bared teeth (lips withdrawn) males circled each other, and by approaching from the side, tried to get underneath the opponentto stab or slice with their canines. As a result, the belly of one victim was ripped open, causing its entrails to spill. The female had calmly observed the fight, lying down with the front legs folded underneath her body; mating with the winning male took place soon after. Females in captivity developed bold patches, sometimes chafed and bloody, after mating, apparently caused by the sharp hooves of the male during copulation. Gestation lasts several months, after which one young is born, “the size of a rat”, dark brown, with a very small head and thin legs. Young can run and jump soon after birth, and suckle only occasionally, only at night and very briefly. The young soon start eating the same leaves and fruits as older animals. Neither male nor female parent made any apparent effort to protect their young. A specimen of 7. javanicus successfully bred with a chevrotain from Bangka Island (presumably 7 kanchil luteicollis) and produced a young with intermediate characteristics. Activity patterns. Observations in captivity suggest that the species is nocturnal, and during the day rests in a quiet, cool, shaded spot. It becomes active around 16:00 h or 17:00 h in the afternoon. Movements, Home range and Social organization. Nothing known. Status and Conservation. The account of 7. javanicus on The IUCN Red List states that the speciesis classified as Data Deficient, partly reflecting the lack of clarity over how many species of chevrotain occur onJava and therefore how much of the available information about the genus refers to 1. javanicus. The authors mention that there are fair indications of a decline, perhaps a major one, and a category such as Vulnerable is quite likely to be applicable. The species is hunted for meat and is also often sold as a pet, for example in markets in West Java. In the 1920s it was reported that the species was commonly trapped using snares, but no recent information on this is available. Dedicated field investigations (throughoutJava) ofstatus are urgently warranted, and the Red List status of the species should be reviewed regularly in light of current uncertainty and concerns. Bibliography. Dakkus (1932), Dobroruka (1967), Doctors van Leeuwen (1921), van Dort (1986, 1988), Duckworth et al. (2008), Farida (2003), Groves & Meijaard (2005), Hoogerwerf (1970), Meijaard & Groves (2004a, 2004b), Supraptomo (1953), van der Vegte (1938)., Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Tragulidae, pp. 320-334 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on pages 331-332, DOI: 10.5281/zenodo.5721279

Tragulus Pallas, 1779. Spicil. Zool., 13:27. TYPE SPECIES: Cervus javanicus Osbeck, 1765. COMMENTS: Original citation, Brisson, 1762, Regn. Anim., 2nd ed., 12:65-68, is not available according to Hopwood (1947)., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 382, DOI: 10.5281/zenodo.7359189, {"references":["Brisson, M. J. 1762. Le regnum animale in classes IX distributum, sive synopsis methodica sistens generalem animalium distributionem in classes IX, & duarum primarum classium, quadrupedum scilicet & cetaceorum, particularem dibvisionem in ordines, sectiones, genera & species. T. Haak, Paris, 296 pp.","Hopwood, A. T. 1947. The generic names of the mandrill and baboons, with notes on some of the genera of Brisson, 1762. Proceedings of the Zoological Society of London, 117: 533 - 536."]}

Tragulus napu (F. Cuvier, 1822). In E. Geoffroy and F. Cuvier, Hist. Nat. Mammifères, pt. 2, 4(37):4 pp. "Chevrotain napu". TYPE LOCALITY: Indonesia, S Sumatra. DISTRIBUTION: Indochina, Thailand, Malaysia, Borneo, Balabac Isl (Philippines), Sumatra, many adjacent islands. SYNONYMS: abjectus, abruptus, amoenus, anambensis, annae, bancanus, banguei, batuanus, billitonus, borneanus, bunguranensis, canescens, flavicollis, formosus, hendersoni, jugularis, lutescens, neubronneri, niasis, nigricans, nigricollis, nigrocinctus, parallelus, perflavus, pretiellus, pretiosus, rufulus, sebucus, siantanicus, stanleyanus, terutus, umbrinus, versicolor. COMMENTS: In older literature, mistakenly given the name T. javanicus; see Van Bemmel (1949b)., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 383, DOI: 10.5281/zenodo.7359189, {"references":["Van Bemmel, A. C. V. 1949 b. Notes on Indo-Australian mammals. 1. A note on Lutrogale perspicillata (I. Geoffroy) (Mustelidae). 2. On the meaning of the name Cervus javanicus Osbeck 1765 (Tragulidae). Treubia, 20: 375 - 380."]}

Tragulus javanicus (Osbeck, 1765). Reise nach Ostindien und China, p. 357. TYPE LOCALITY: Indonesia, W Java, Udjon Kulon Peninsula. DISTRIBUTION: Indochina, Thailand, Yunnan (China), Malaysia, Sumatra, Borneo, Java, many adjacent islands. SYNONYMS: affinis, angustiae, brevipes, carimatae, everetti, focalinus, fulvicollis, fulviventer, fuscatus, hosei; indicus Brisson, 1765 (not available); insularis, kanchil, klossi, lampensis, lancavensis, longipes, luteicollis, masae, mergatus, natunae, pallidus, pelandoc, penangensis, pidonis, pierrei, pinius, pumilus, ravulus, ravus, rubeus, russeus, russulus, subrufus, virgicollis, williamsoni. COMMENTS: In older literature, known as T. kanchil; see Van Bemmel (1949b)., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 382, DOI: 10.5281/zenodo.7359189, {"references":["Van Bemmel, A. C. V. 1949 b. Notes on Indo-Australian mammals. 1. A note on Lutrogale perspicillata (I. Geoffroy) (Mustelidae). 2. On the meaning of the name Cervus javanicus Osbeck 1765 (Tragulidae). Treubia, 20: 375 - 380."]}

Tragulus meminna (Erxleben, 1777). Syst. Regn. Anim., 1:322. TYPE LOCALITY: Sri Lanka. DISTRIBUTION: Sri Lanka; peninsular India; Nepal. ISIS NUMBER: 5301419005002002001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 319, DOI: 10.5281/zenodo.7352976

Tragulus Pallas, 1779. Spied. Zool., 13:27. COMMENT: Original citation, Brisson, 1762, Regn. Anim., 2nd ed., 12:65-68, is not available. ISIS NUMBER: 5301419005002000000., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 318, DOI: 10.5281/zenodo.7352976, {"references":["Brisson, M. J. 1762. Le regnum animale in classes IX distributum, sive synopsis methodica sistens generalem animalium distributionem in classes IX, & duarum primarum classium, quadrupedum scilicet & cetaceorum, particularem dibvisionem in ordines, sectiones, genera & species. T. Haak, Paris, 296 pp."]}

Tragulus napu (F. Cuvier, 1822). In E. Geoffroy and F. Cuvier, Hist. Nat. Mamm., 37:2. TYPE LOCALITY: Indonesia, S. Sumatra. DISTRIBUTION: Indochina; Thailand; Malaysia; Borneo; Balabac Isl. (Philippines); Sumatra; many adjacent isls. ISIS NUMBER: 5301419005002003001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 319, DOI: 10.5281/zenodo.7352976