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Description |
Geographic Range [top]
Range Description: This species occurs in Sri Lanka, most of India, northern Pakistan, Nepal, Bhutan, Bangladesh and southern China, including Hainan and southern Tibet, and into southeast Asia (Myanmar, Thailand, Lao PDR, Viet Nam, Cambodia) south to the northern part of the Thai–Malay peninsula (Roberts 1977, Groves and Grubb 1990, Ohtaishi and Gao 1990, Corbet and Hill 1992, Wang Ying-xiang 2002, Groves 2003, Grubb 2005). The precise southern range limit remains unclear on the Thai–Malay peninsula (see Taxonomy). It has been introduced in the Andaman Islands (Corbet and Hill 1992). It is widespread in most range states, except Pakistan (see Population).
Countries occurrence:
Native:
Bangladesh; Bhutan; Cambodia; China; Hong Kong; India; Lao Peoples Democratic Republic; Myanmar; Nepal; Pakistan; Sri Lanka; Thailand; Viet Nam
Additional data:
? Upper elevation limit (metres): 3500
Range Map: Click here to open the map viewer and explore range.
Population [top]
Population: This adaptable and widespread species remains locally common in most of its range (e.g. Whitehead 1993, Le Xuan Canh et al. 1997, Timmins et al. 1998, Duckworth et al. 1999, Lynam et al. 2001, Lynam 2003, Johnsingh et al. 2004, Polet and Ling 2004, Johnson et al. 2006, Steinmetz et al. 2008), even in Hong Kong, an almost totally deforested landscape which has in the past sustained hunting and habitat disruption heavy enough to force extinction even of squirrels (R. Corlett pers. comm. 2008, Shek et al. 2007). About 20–30 individuals were estimated by Roberts (1977, based on a 1972 estimate) for Margalla National Park in northern Pakistan, this would account for most of the population surviving in that country at the time. In Bangladesh the species is the most widespread wild ungulate, more widely distributed even than wild hogs. However, the population is declining and the species is nowhere very common. It is a rare species with only a small population in the Sundarbans which constitutes half of the remaining forests of Bangladesh, but it is uncommon to rare in the remaining hill forest areas of northeast and southeast of the country (Md Anwarul Islam in litt. 2008, Suprio Chakma pers. comm. 2014), this reflects “rampant...forest destruction” during the British colonial era which was, if anything, even worse after independence (Khan 2003). Hunting is the main current threat, rather than habitat loss as it is a well-adapted species. Even so, muntjacs (assumed to be this species) can still be seen readily in the north-east (P.M. Thompson pers. comm. 2008). In the southeast in the Chittagong Hill Tracts muntjacs were detected in all of 38 surveyed grid cells (each 10,000 ha and >50% forest cover) during a camera-trapping survey in 2010-2011, suggesting occupancy of essentially all remaining suitable habitat patches (Suprio Chakma pers. comm. 2014). The species’ status in China is somewhat clouded by the overlap in range with M. reevesi (as well potentially as other muntjac species), both species have mapped ranges in eastern Yunnan, Guizhou, Guangxi, Guangdong, southern Hunan and southern Jiangxi and possibly western Fujian (Zhang et al. 1997, Corbet and Hill 1992). In some areas camera-trap pictures of these two species may be very difficult to tell apart (R. J. Timmins pers. comm. 2014). For instance there was much uncertainty about the identity of animals in Hong Kong with thoughts that perhaps only M. reevesi occured (Shek 2006), however camera-trapping surveys recently concluded that M. vaginalis [as M. muntjak s.l.] was the only species present (Shek et al. 2007), but further investigation is warranted as although some animals do appear to be diagnosable as M. vaginalis, some other captive animals at the Kadoorie Farm and Botanic Garden and a photo of a female in Shek (2006) show character combinations that make firm diagnosis as M. vaginalis or M. reevesi impossible, suggesting perhaps introgression (R. J. Timmins pers. comm. 2014), Shek et al. (2007) note significant differences in general pelage of muntjac in Hong Kong. In Hong Kong muntjac populations would appear to be stable and “very common”, having increased from a low point at some point in the past (Shek et al. 2007, Pei et al. 2010). Population status of muntjacs is presumed to be worst in the southern provinces especially Guangxi, Guangdong, but it is currently unclear whether there are differences in status between M. vaginalis and M. reevesi in these provinces. On the basis of recent surveys using multiple methods including camera-trapping (Lau et al. 2010) muntjacs (species uncertain) remain relatively widespread and appear to persist in most reasonably sized areas of suitable habitat in Guangxi, Guangdong and southern Jiangxi where hunting pressure is extremely high and populations of almost all large mammals are severely depressed or extirpated (Lau et al. 2010, B.P.L. Chan and M.W.N. Lau pers. comm. 2014). However at a site based level muntjac in these provinces are notably scarcer than in Hong Kong, based on encounter rates of calls, faeces and camera-trap photographs, this is attributed by rural hunters to recent very heavy hunting with gin-traps precipitated by a fire-arm ban (B.P.L. Chan and M.W.N. Lau pers. comm. 2008). Status appears to be similar in Hainan, where M. vaginalis is currently the only known species, and hunting pressure is high (Lau et al. 2010, M.W.N. Lau pers. comm. 2014), although in some well protected areas the species appears to be abundant with incredibly high densities reported (e.g. 54 animals/km²: Teng et al. 2005, such high densities are clearly unnatural and suggest either methodological error or peculiar ecological circumstances).
Although Sheng and Ohtaishi (1993) stated that the total Chinese population of M. vaginalis [as M. muntjak s. l.] was about 500,000, the assumptions behind such a number were not explicit and it should be treated with great caution. Ohtaishi and Gao (1990) estimated that a total of 30,000 individuals were harvested each year on Hainan Island alone.
Populations are certainly also declining in Lao PDR, Cambodia and Viet Nam, through heavy trade-driven hunting (see threats). Muntjacs in a number of Thai protected areas are increasing (Steinmetz et al. 2010, Steinmetz et al. in press) so the era of widespread population decline in Thailand might have largely ended, as few are likely to remain outside of protected areas. ‘Red’ muntjacs are common in Kaeng Krachan National Park, Thailand (Ngoprasert et al. 2007), but this is within the main transition zone between Sundaic and northern taxa, so the species there is/are unknown. Populations in Nepal are thought to be relatively stable (Hem Sagar Baral pers. comm. 2008). Muntjac in deciduous forests in western Thailand attain densities of 2.1–3.1 individuals/km² (Srikosamatara, 1993, Steinmetz and Mather, 1996, Sukmasuang, 2001). Densities are 2.2/km² in the deciduous forests of eastern Cambodia (Gray et al. 2012). The following densities have been estimated in India: Bhadra Tiger Reserve, 3.64 +/- 0.63 (SE) per km² (Jathanna et al. 2003), Kanha moist deciduous forest, 0.6 animals per km² (Karanth and Nichols 2000), Nagarahole moist deciduous forest, 6 animals per km² (Karanth and Nichols 2000), Bandipur dry deciduous forest, 0.7 animals per km² (Karanth and Nichols 2000), Tadoba-Andhari dry deciduous forest, 0.9 animals per km² (Karanth and Kumar 2005), Melghat dry deciduous forest, 0.6 animals per km² (Karanth and Kumar 2005), Wilpattu, Sri Lanka, 2.5 animals per km² (Barrette 1977), Karnali-Bardia, Nepal, 1.7 animals per km² (Dinerstein 1987), and Rajaji National Park, 1 animal per km² (A.J.T. Johnsingh pers. comm. to Barrette 2004). Where studied, population densities are always well below those of herding ungulates (previous citations), but this is not so in areas where hunting has perverted ratios, where Northern Red Muntjac may even be the only deer species remaining.
Current Population Trend: Decreasing
Additional data:
? Population severely fragmented: No
Habitat and Ecology [top]
Habitat and Ecology: The Northern Red Muntjac inhabits a wide variety of forests and scrub, including level plains and rugged mountains, dense and open forests, evergreen and deciduous areas, old-growth and secondary forests, and even grass- and cropland near woody habitats, from sea-level commonly up to about 1,500 m asl and occasionally over, perhaps with some variation regionally (Blanford 1891, Prater 1971, Barrette 1977, 2004, Lekagul and McNeely 1977, Roberts 1977, Timmins et al. 1998, Duckworth et al. 1999, Teng et al. 2004). Within this broad habitat use, densities doubtless vary based on conditions. Steinmetz et al. (2008) found that signs, and presumably animals, were significantly more common in lower- than higher-lying areas of the Tenasserim–Dawna mountains, Thailand, noting that this was the opposite of the patterns that would be predicted if hunting (itself concentrated heavily in the lowlands) was the chief determinant of population densities. Population densities in Thailand are substantially higher in deciduous forest than in evergreen forest (Steinmetz and Mather 1996, Steinmetz et al. 2010). Kushwaha et al. (2004) found that in Kumaon Himalaya (India), M. vaginalis frequently exploits grass- and herb-dominated environments, linked to highly logged locations, and with a preference for lower altitudes (although they did not specify what altitude range was covered, nor discuss the possibility that because logging tends to occur in valley bottoms and lower hills rather than on steep hills and ridges, a preference for lower altitudes could simply be a consequential effect of a preference for logged areas. Teng et al. (2004) found that in Hainan, muntjacs preferred shrub grassland and thorny shrubland, used dry savanna in proportion to its availability, and avoided woods, cultivated grass-plot, and deciduous monsoon forests. Sheng and Ohtaishi (1993) attributed the apparent stability in numbers of the Chinese population to logging activities, which were bringing about an increase of secondary forest and scrub dominated areas. A study of coffee estates around Bhadra Wildlife Sanctuary, India, recorded Red Muntjac more widely than any other herbivorous mammal, including in coffee areas far from the sanctuary’s boundary (Bali et al. 2007).
Its diet is based on fallen fruits, buds, small seeds, twigs, seed pods, tender leaves, and young grass (Barrette 1977, 2004, Lekagul and McNeely 1977, Roberts 1977, Chen et al. 2001, Teng et al. 2004). The species has been cited as a major dispersal agent of fruit-producing plants in southern China (Chen et al. 2001) and Thailand (Lekagul and McNeely 1977) and this is presumably so throughout its range.
In much of its range (although rather less than was popularly supposed until recently) this is the only muntjac species. Where congeners occur, it is widely sympatric, evidently even syntopic, with them, there is no solid evidence for competitive exclusion except perhaps at very high altitudes. Where it overlaps with Fea’s Muntjac M. feae in Thailand, Red clearly predominates in more deciduous habitats and lower altitudes, with Fea’s apparently more common in high altitudes and fully evergreen forest (R. Steinmetz pers. comm. 2008). It probably occurs throughout the range of Large-antlered Muntjac M. vuquangensis (e.g. Timmins et al. 1998). There is extensive overlap with muntjacs of the M. rooseveltorum species-complex (M. rooseveltorum, M. truongsonensis, M. puhoatensis and M. putaoensis), but there may be areas above about 1,500 m asl where the latter greatly outnumber M. vaginalis and some where M. vaginalis is absent (R.J. Timmins pers. comm. 2008, based on examination of numerous camera-trap photographs from many camera-trap programmes). Camera-trap results from northern Myanmar suggest that Red Muntjac occurs largely below the altitudinal range of M. gongshanensis, but fully through the range of M. putaoensis (R.J. Timmins pers. comm. 2008 based on WCS unpublished data). Schaller and Rabinowitz (2004), however, reported many M. vaginalis at high altitudes in Arunachal Pradesh, with one direct sighting at 3,000 m asl near Tawang, and there may therefore be significant local variation in the upper altitudinal limit. Two specimens in the Natural History Museum, London from Nepal have recorded altitudes of 11,400 and 8,500 feet (3,450 and 2,590 m asl) (R.J. Timmins pers. comm. 2008). Camera-trapping in Bhutan has recorded the species relatively commonly between 2000 and 3650 m, over essentially the full elevation range surveyed, where this may be the only muntjac species present (K. Vernes, S. Dorji and R. Rajaratnam based on unpublished data pers. comm. 2014, photos seen by R.J. Timmins 2014).
Northern Red Muntjac appears to be able to breed all year around, although the existence of well-defined breeding seasons has been suggested for some populations, particularly in those inhabiting locations with a more seasonal climate (Barrette 1977, Lekagul and McNeely 1977, Roberts 1977, Chen et al. 2001). In any case, "peaks" of mating activity are present, at least, in some areas (Prater 1971, Barrette 1977, Roberts 1977). Despite only a single fawn being born each occasion, overall lifetime output is high (Barrette 2004). Even where births occur throughout the year, there is still a fairly rigid cycle of antler formation and shedding, synchronised at the population level (Barrette 1977, Mishra 1982, Acharjyo and Patnaik 1984). It tends to be solitary (Eisenberg and Lockhart 1972, Seidensticker 1976, Karanth and Sunquist 1992, Barrette 2004), reports of small groups could be attributed to mating pairs or to female/offspring units, or to temporary incidental proximity of individuals at concentrations of high-value food, e.g. fallen fruit under a tree (Barrette 1977, 2004, Lekagul and McNeely 1977). Individuals seem strongly attached to a certain home range, but there is no convincing evidence of territoriality (Barrette 1977, 2004). Northern Red Muntjac is diurnal in some areas, nocturnal in others (Prater 1971, Barrette 1977, Lekagul and McNeely 1977, Roberts 1977) and is probably best regarded as crepuscular. It is an important prey for Leopard Panthera pardus, Tiger P. tigris, and Dhole Cuon alpinus.
Systems: Terrestrial
Use and Trade [top]
Use and Trade: This species is widely traded and hunted in China and Southeast Asian countries. China appears to be a major consumer and producer of Muntiacus vaginalis products. There is an intense commercial activity of this species from bordering Southeast Asian countries into China which involves meat, hides and medicinal products. (Ohtaishi and Gao 1990, Sheng and Ohtaishi 1993, Zhijun et al. 1996, Duckworth et al. 1999, see also other references in General Information section). It is not usually kept as livestock.
Threats [top]
Major Threat(s): Northern Red Muntjac are hunted widely, both legally and illegally, over most of its range (Sheng and Ohtaishi 1993, Madhusudan and Karanth 2002, Tungittiplakorn and Dearden 2002, Jathanna et al. 2003, Johnson et al. 2003, Santiapillai and Wijeyamohan 2003, Hansel 2004, Kaul et al. 2004, Kumara and Singh 2004, Rao et al. 2005, Steinmetz et al. 2008), and it is probably among the most used and most desired wild meats in South and South-east Asia (e.g. Srikosamatara et al. 1992, Duckworth et al. 1999). Patterns of threat vary across the species range. Viet Nam, Lao PDR and to a lesser extent Cambodia and northern Thailand are apparently the areas where mammal hunting has been heaviest. In its mainland China range, rates may be even higher, because a recent fire-arm ban is said to have pushed up levels of gin-trapping (B.P.L. Chan pers. comm. 2008). Muntjacs are hunted within a thriving hunting and wildlife trading culture in this area and in adjacent countries such as China and Thailand, involving many land vertebrate species, along with other forest products such as orchids and Aquilaria resin (e.g. Compton and Le Hai Quang 1998, Compton et al. 1999, Noreen and Claridge 2001). Such hunting reaches all areas, although in large rugged mountain forest blocks trade-driven hunting of species valued only for their meat (rather than high-value, low-weight, generally medicinal products that can be carried out efficiently from even the remotest areas) is still limited by economics of accessibility. The regional wild meat trade has little likelihood of abating as long as there are any pigs, muntjacs, and civets to be hunted. The human population of Viet Nam is more than 90 million, that of China more than 1,000 million. Together, they comprise an enormous market for wildlife products. For example, tens of millions of wild turtles are imported, legally and illegally, into China annually (van Dijk et al. 2000). There has been no comparable study of ungulate trade levels. Within the Northern and Central Annamites, every square kilometre of Viet Namese forest and of Lao forest that is within 5 km of the Viet Namese border probably has snares capable of capturing muntjacs set in it every year (Timmins et al. 2007, R.J. Timmins pers. comm. 2008). Intensity in some areas probably reaches many thousands of snare-nights per square kilometre per year (Timmins et al. 2007). Snaring is less intensive in Lao PDR at least away from the Viet Namese border, but is increasing dramatically (Timmins and Robichaud 2005, W.G. Robichaud pers. comm. to R.J. Timmins 2007, Coudrat et al. 2014, Timmins and Duckworth 2012, R.J. Timmins in press b, R.J. Timmins pers. comm. 2014). Hunting intensity had in recent decades been generally lower in southern Viet Nam than in Lao PDR and northern Viet Nam, leaving many wildlife populations less depleted (e.g. Le Xuan Canh et al. 1997, Timmins and Duckworth 2000, Polet and Ling 2004), but the general increase in regional hunting intensity and economic wealth indicate that hunting intensity in this region must also be increasing (Timmins et al. 2008), as of 2014 there are indications that hunting intensity has reached equivalent levels to the north, at least in some areas (Timmins in press a). In northern Thailand when muntjacs become rare, the animals are hunted in group drives, the meat is often sold (Tungittiplakorn and Dearden 2002).
Closed tropical evergreen forests have inherently low ungulate biomass (Robinson and Bennett 2000), and beyond this populations of all wild animals larger than 20 kg in Indochina have been significantly reduced by human exploitation (Timmins and Ou 2001, Timmins et al. 2007, R.J. Timmins pers. comm. 2008). Field survey encounter rates with Northern Red Muntjacs in Lao PDR, Viet Nam, and Cambodia are remarkably low compared with other areas in its range (Timmins and Ou 2001, Timmins et al. 2007, R.J. Timmins pers. comm. 2008). In a recent reconnaissance survey of highly fragmented habitat in southern Lao PDR (Pathoumphon district, Champasak province), outside the protected area system, muntjac were the only ungulates (assumed to relate to this species on habitat and distribution) for which signs were commonly found, even wild pigs Sus sp(p). being noticeably more scarce (K. Khounboline and J.W. Duckworth pers. comm. 2008). This pattern is general throughout Lao PDR (K. Khounboline pers. comm. 2008). However, areas of heavy hunting have lost muntjacs or had them reduced to extreme rarity, not only forest isolates such as Houay Nhang (a few square kilometres,, just outside Vientiane, Duckworth et al. 1994) but also, where hunting has been extremely heavy, less fragmented landscapes (such as the western lowlands of Xe Bang-Nouan National Protected Area in south Lao PDR, J.W. Duckworth pers. comm. 2008). In such areas, even though forest within them is in general heavily degraded, habitat is unlikely to be the prime cause of decline, because in some other areas of long-standing forest devastation now dominated by a mix of secondary ruderals, shrubs and grasses, muntjacs are still relatively common, e.g. Phongsali (far northern Lao PDR), where they can even still be heard from the provincial capital town (Duckworth and Robichaud 2005). Similarly, Red Muntjac is common and widespread in Hong Kong, an almost deforested landscape (Chek 2006, R. Corlett pers. comm. 2008), and studies in Hainan indicated selection of openings and other edge habitats rather than forest interiors (Teng et al. 2004), this seems generally true throughout South-east Asia (R.J. Timmins and J.W. Duckworth pers. comm. 2008). There is also evidence from elsewhere in its range that this species is better able than most ungulates to withstand a certain degree of continuous harvesting as well as habitat disturbance (Jathanna et al 2003), although this has rarely been based on quantified status assessment. In Nagarahole National Park, India, Madhusudan and Karanth (2002) found no difference in muntjac abundance between a heavily hunted and a lightly hunted site, compared with enormous differences for most other ungulates. In the context of Indochina, even their heavily-hunted site would, however, rank as lightly hunted (J.W. Duckworth pers. comm. 2008, from based on visit and discussion). Madhusudan and Karanth (2002) cautioned against assuming that these results indicated an intrinsic resilience to hunting by this muntjac: it is not so readily hunted as, for example, Chital Axis axis, because of its solitary nature, affinity for dense cover and therefore risks of getting caught are higher when using appropriate techniques to kill or capture it. However, the ubiquitous disparity in status between M. vaginalis and other ungulates (and indeed almost all other mammals of similar or larger body weight) in Indochina indicates that it is truly much more resilient to hunting. Muntjac populations are capable of recovering quickly from overexploitation if hunting is controlled: an overhunted, low density population freed from hunting pressure in Thailand increased 10-fold in 6 years, a rate of increase that is likely near the biological maximum attainable by the species (Steinmetz et al. 2010). An intensive study of hunting on mammals in two Hmong villages of northern Thailand classified quarry species into tiers representing the sequence of loss through overhunting, muntjacs (most or all were presumably M. vaginalis) fell in the fifth tier, persisting much longer than all other ruminants and with only species like squirrels, civets, and linsangs being placed later in the extinction sequence (Tungittiplakorn and Dearden 2002). Even so, villagers reported that muntjacs were greatly reduced in numbers in the area. The strongest demonstration of its remarkable resilience is that it is now common and widespread in Hong Kong, an almost deforested landscape which has in the past sustained hunting and habitat disruption heavy enough to force extinction even of squirrels. Red Muntjac were reduced to very low populations a few decades ago, but did not become extinct, and has increased rapidly in recent years (Shek 2006, R. Corlett pers. comm. 2008).
While wild meat is the major driver of South-east Asian muntjac trade, parts such as antlers are also heavily traded (e.g. Duckworth et al. 1999, Noreen and Claridge 2001).
Rapid economic development and expanding wealth in the species range, particularly South-east Asia and China, are increasing the demand for wildlife meat and ‘medicines’ (Timmins et al. 2007). Timmins et al. (2007) highlighted a common misconception, especially in documentation of development projects in the region, that poverty is a principal cause of biodiversity loss, but as they pointed out for Saola Pseudoryx nghetinhensis, the main driver of threats to wild ungulates, at least for the mid-term, is not rural poverty but increasing urban wealth in Indochina and China. Similarly, in Thailand, trade-driven hunting has been the main cause of muntjac and other wildlife population declines, regardless of protected area status (Steinmetz et al. 2006). In Viet Nam “the free market economy has resulted in feverish periods of trade in wild species nationwide, with negative impacts on biodiversity” (Government of the Socialist Republic of Viet Nam 2004). In Cambodia, the same factor has fostered a thriving bushmeat market and hunting of species for international wildlife trade (Timmins and Ou 2001, Maxwell et al. 2006, Timmins 2006). There is a rapid, ongoing, expansion of wealthy social strata in Lao PDR, directly accelerated by illegal trade in rosewood, wild meat, and other forest resources, and by the economics of large infrastructure projects (e.g. Nan Theun 2 hydro-electric power project), creating a significantly greater in-country demand for luxury meats such as venison (W.G. Robichaud pers. comm. 2007). Wild meat is chosen in preference to farmed meat despite a greater price per unit weight (e.g. Hansel 2004). Markets along major roads such as route 13 (e.g. at Ban Namthon) have expanded, not contracted, in the last 15 years and have a huge (albeit unquantified) turnover of wild meat, at this specific market, multiple stalls are selling dried muntjac meat daily (balance of muntjac species unknown). The effects of hunting in Lao PDR, Viet Nam, Cambodia and Thailand have been exacerbated during the last two decades by habitat loss and various socio-economic factors, of which the most biologically significant are discussed in the Red List account for Large-antlered Muntjac M. vuquangensis.
Thus, even though this is one of the most resilient of large mammals in Indochina persisting in fragmented landscapes and often in peripheral forest areas, populations in Indochina (Viet Nam, Lao PDR and Cambodia) are likely to be declining significantly (perhaps at a level to trigger VU i.e. 30% in 10–15 years, for the regional population) (Timmins et al. 2008). Densities especially in Lao PDR and Viet Nam are generally well below carrying capacity, although in extensive areas of Cambodia are likely to be more natural, as they are across the range west of the Mekong (Timmins et al. 2008). Since 2008 rates of decline in Indochina have almost certainly increased and are now quite likely to be in the region of ‘30% in 10–15 years’, especially in Lao and Cambodia where economic growth has seen widespread landscape level changes in infrastructure and land use and concomitant rise in the value of bushmeat and in hunting pressure, although most areas in Vietnam may have already been so depleted prior to 2008 that hunting of residual animals has been relatively difficult since then, thus tempering decline rates, even though hunting intensity has probably increased (R.J. Timmins pers. comm. 2014).
Conservation Actions [top]
Conservation Actions: Northern Red Muntjac occurs in many protected areas throughout its range and in most countries is covered by laws regulating hunting at some level. Across large parts of its range, the species would benefit if these national laws involving the protection of the species were more efficiently enforced, particularly those which relate to wildlife trading. Also, greater effort should be devoted to engaging local communities in the protection and recovery of wildlife populations in nearby protected areas. The proactive involvement of communities has resulted in increased population trends of muntjac and other hunted species at various sites in the region.
Citation: Timmins, R.J., Steinmetz, R., Samba Kumar, N., Anwarul Islam, Md. & Sagar Baral, H. 2016. Muntiacus vaginalis. The IUCN Red List of Threatened Species 2016: e.T136551A22165292. http://dx.doi.org/10.2305/IUCN.UK.2016-1.RLTS.T136551A22165292.en. Downloaded on 23 August 2016.
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