The Dhole is a habitat generalist, and can occur in a wide variety of vegetation types, including: primary, secondary and degraded forms of tropical dry and moist deciduous forests; evergreen and semi-evergreen forests; temperate deciduous forests; boreal forests; dry thorn forests; grassland–scrub–forest mosaics; temperate steppe; and alpine steppe. Consequently, their elevation range is from sea level to as high as 5,300 m asl in Ladakh (R. Simpson pers. comm.). They have not been recorded in desert regions.

The Dhole is one of only three canid species with specialized dental adaptations for an exclusively carnivorous diet, termed hypercarnivory (Van Valkenburgh 1991). Although Dholes consume a wide variety of prey species, ranging from small rodents and hares to Gaur (Bos gaurus; Karanth and Sunquist 1995, Andheria et al. 2007, Ramesh et al. 2012, Selvan et al. 2013a), the preferred prey are ungulates with a body mass of 40-60 kg (Selvan et al. 2013b). However, if this prey size is not available, Dholes will selectively prey upon both smaller and larger ungulate species (Kamler et al. 2012). Seasonal changes occur in the Dhole diets, reflecting seasonal changes in availability and numbers of prey (Thinley et al. 2011). In India, Dholes form relatively large packs (usually five to 10, but up to 25 adults) to efficiently hunt large numbers of prey, as well as to protect litters, which are usually large (usually five to 10, but up to 12 pups; Johnsingh 1982, Venkataraman et al. 1995, Durbin et al. 2004). However, in tropical evergreen forests of Southeast Asia, Dholes appear to persist in smaller packs and presumably have smaller litters, probably due the low prey biomass and small size of ungulate prey in these habitats (Kawanishi and Sunquist 2008).

Due to the demands imposed by hypercarnivory, sufficient numbers of ungulate prey are the Dhole’s major habitat requirements. In India, tropical dry and moist deciduous forest may represent optimal habitats, based on the areas thought to hold the largest Dhole populations. Ungulate biomass, particularly that of cervid species, is highest in these habitat types when compared to other habitats in the same region (A. Venkataraman and V.N. Babu unpubl.). Besides prey numbers, other important factors that may influence habitat use include levels of human disturbance, water availability, tiger presence, and suitability of breeding sites (Steinmetz et al. 2013, Srivathsa et al. 2014, J.F. Kamler unpubl. data).

Home range sizes of Dholes reportedly ranged from 23-199 km² in India (Johnsingh 1982, Venkataraman et al. 1995, Karanth and Sunquist 2000, Acharya 2007) and from 60-80 km² in Thailand (Grassman et al. 2005, K. Jenks unpubl. data).

Historically, Dholes occurred throughout South and East Asia, to as far north as the southern parts of the Russian Federation (including the Amur region and upper Lena River north of Lake Baikal), and as far west as the mountains ranging from eastern Kazakhstan to northern Pakistan (Altai, Tian Shan, Pamir and western Himalayan mountains; Heptner and Naumov 1967). Dholes have disappeared from >75% of their historic range, and most remaining populations are fragmented and still appear to be declining based on scattered surveys and personal observations. Their current distribution, based on recent information we consider reliable, is described below.

Central and East Asia: there have been no confirmed reports of Dholes in more than 30 years from the Russian Federation, Mongolia, Kazakhstan (formerly in the Altai and Tian Shan mountains), Kyrgyzstan (formerly in the Tian Shan and Pamir mountains), Afghanistan (formerly in Pamir Mountains), Tajikistan (formerly in Pamir Mountains) or Uzbekistan (formerly in Tian Shan Mountains), and they are likely extirpated from these regions (A. Poyarkov and N. Ovsyanikov in litt., D. Miquelle pers. comm.).

Bangladesh: Dholes still occur in some forest reserves in the Sylhet area in northeastern Bangladesh, as well the Chittagong Hill Tracts in southeastern region. However, these areas are unlikely to contain viable populations because Dholes are usually sighted only as solitary individuals, or in small groups of two to three adults. Dhole numbers appear to be decreasing in these areas due to a decreasing prey base (S. Chakma pers. comm.).

Bhutan: Although Dholes were nearly extirpated in Bhutan in the 1970s and early 1980s due to poisoning campaigns, this species began to re-occupy the country starting in the 1990s (Wangchuk 2004, Thinley et al. 2011). Based on sign and interviews with park managers, they now occur in most, if not all, protected areas in the country (J. Kamler unpubl. data).

Cambodia: There are recent (under five years old) records of Dholes in the Cardamom Mountains (FFI-Cambodia Programme unpubl. data), the northern plains landscape (WCS Cambodia Program unpubl. data), the eastern plains landscape (J. Kamler unpubl. data), and northeastern Cambodia (Siem Pang Province; D. Willcox unpubl. data). Dholes do not likely exist in viable populations outside of these areas. A recent outbreak of canine distemper in 2011-2012, thought to have originated in local domestic dogs, caused the near-extirpation of Dholes in both the eastern and northern plains landscapes (J. Kamler and A. Suzuki unpubl. data), but it appears these populations are slowly recovering as of 2015.

China: Historically Dholes occurred throughout China, but their current status and distribution is highly uncertain, and they are likely extirpated throughout most of the country. Despite extensive surveys throughout the country, recent (<10 years) records from China come only from six provinces: Gansu, Sichuan, Shaanxi, Yunnan, Tibet Autonomous Region (TAR) and Xinjiang. In northern Gansu Province, Dholes were recently reported from the Qilian Shan Mountains as a pack was observed there in 2003 (Harris 2006) and a mother with pups was photographed in Yanchiwan Nature Reserve in 2013 (Riordan et al. 2015). As the Qilian Shan Mountains extend into northern Qinghai Province, Dholes still might occur there as well. In southeastern Xinjiang Province, camera traps recorded Dholes during 2012-2013 in the Altun (Altyn-Tagh) Mountains (Xue et al. 2014). In western Xinjiang Province, there were sightings by local people during 2011-2013 in Taxkorgan Nature Reserve (Riordan et al. 2015). In central Sichuan, a Dhole was photographed in 2012 in Heishuihe Nature Reserve in the Qionglai Mountains (Y. Zhou pers. comm.). Although one Dhole was observed in 2003 in Tangjiahe Nature Reserve in northern Sichuan (Y. Zhou pers. comm.), there have been no camera trap photos or other confirmed records in the past 10 years (S. Li pers. comm.). Dholes may occur in the Minshan Mountains (northern Sichua) and Daxue Shan Mountains (western Sichuan), as there are former records from there and current prey densities are relatively high (S. Li pers. comm., Li et al. 2010). In Shaanxi Province, a Dhole was recently recorded during a camera trap study in Guanyinshan Nature Reserve in the Qinling Mountains (Liu et al. 2013). However, camera trap studies in several other nature reserves in Shaanxi failed to record Dholes (S. Li pers. comm.), so their numbers must be extremely low in the area. In Yunnan Province, Dholes were photographed in camera traps in both the Nangunhe and Xishuangbanna nature reserves in 2012-13 near the border with Myanmar (Wildlife Institute, Beijing Forestry University, unpubl.), but Dhole numbers are likely low there due to highly fragmented habitat, high poaching, and extremely low prey densities (S. Li pers. comm.). In TAR, camera traps recorded Dholes in 2014 in Motuo (Mêdog) County (southeastern TAR; S. Li pers. comm.). Although there are few recent records, Dholes may still occur in the southern parts of TAR, as this species was recorded in the Mt. Qomolangma (Mt. Everest) Nature Reserve (Hu et al. 2014). In southeastern China, the last two records of Dholes come from Jiangxi Province: the unconfirmed report of one captured in early 2000s (C. Bellamy pers. comm.), and an unconfirmed photo taken by a camera trap in Taohongling Nature Reserve in 2010 (S. Li pers. comm.). However, there have been no confirmed reports or additional records from other areas, and they are now likely extirpated from the region. Interviews with local people suggested that Dholes disappeared from large areas of central and southern China during the 1980s and early 1990s, after locals starting poisoning carcasses in retaliation for livestock losses from Dholes (S. Li pers. comm.).

India: Dholes occur in several regions of India, and this country undoubtedly contains the largest numbers of Dholes. That said, Dholes have disappeared from 60% of their historic range in India during the past 100 years (Karanth et al. 2010). Relatively high populations of Dholes are still found in the Western Ghats and central Indian forests, due to high prey numbers and extent of protected forests, whereas lower numbers of Dholes are found in the Eastern Ghats (Karanth et al. 2009). Dholes are also found in the northeastern states, although numbers are low and decreasing in this region due to a decreasing prey base and retaliatory killings from livestock predation (Gopi et al. 2012, Lyngdoh et al. 2014). Dholes are found in some areas of Terai region in northern India (Karanth et al. 2009), although their exact distribution there is unknown. In the Himalayan region, Dholes were recently reported from Sikkim (Bashir et al. 2014), and in 2008 near Tso Kar in Ladakh (R. Simpson pers. comm.), thus they may occur in other areas of Ladakh as well.

Indonesia (Sumatra and Java): Historically, Dholes occurred throughout both Sumatra and Java; however, their current distribution on both islands is fragmented and greatly reduced. On Sumatra, Dholes have recently been confirmed in several national parks along the Barisan Mountain range, ranging from the northern to southern parts of the island (e.g., Gunung Leuser, Kerinci Seblat and Bukit Barisan Selatan National Parks; FFI, WCS and WWF country programs unpubl. data). Dholes also have been recently confirmed in several protected areas in lowland forests in the east-central part of the island (e.g., Tesso Nilo and Bukit Tigapuluh National Parks, Harapan Rainforest and Batang Hari Protection Forest; FFI and WWF country programs unpubl. data). On Java, Dholes have recently been confirmed in national parks only in the extreme western (e.g., Gunung Gede Pangrango, Ujung Kulon and Gunung Halimum Salak National Parks) and eastern (e.g., Baluran National Park and Alas Purwo National Park) parts of the island (A. Ario pers. comm.). They are likely extirpated in other regions of the island.

Korean peninsula: Historically, Dholes occurred on the Korean peninsula, but were likely extirpated throughout most of their range by the 1970s (Won and Smith 1999). In the 1980s Dholes were confirmed to still occur on Mt. Pakdoo in northeastern Korea DPR, near the Chinese border (Won and Smith 1999). However, there have been no available data since that time, so their status in Korea DPR remains uncertain. They are certainly extirpated from the Republic of Korea.

Lao PDR: There are recent (under five years old) records of Dholes from northern Lao PDR (Nam Et-Phou Louey [NEPL] and Nam Ha protected areas) and in central Lao PDR (Nakai-Nam Theun and Nam Kading protected areas; WCS-Lao PDR program unpubl. data, C. Coudrat pers. comm.). The most studied and perhaps largest population of Dholes in the country is in NEPL (Kamler et al. 2012). There are no recent records of Dholes in southern Lao PDR, and they are probably extirpated from this region.

Malaysia (peninsula): The historical range of Dholes likely included the entire Malaysian peninsula, possibly even what is now Singapore. Dholes are now extirpated from Singapore and the southern forests of the Malaysian peninsula (Endau-Rompin complex; WCS Malaysia Program, unpubl.). Based on recent camera trapping surveys, Dholes still occur in the central (Taman Negara) and northern (Belum-Temengor complex) forested areas of the Malaysian peninsula (K. Kawanishi pers. comm.). 

Myanmar: The current distribution of Dholes in Myanmar is uncertain. In the late 1990s and early 2000s, they were recorded by camera traps at 11 of 15 survey areas scattered across the country (Durbin et al. 2004). However, numbers and distribution may have decreased since that time. For example, Dholes had been recorded in Chatthin Wildlife Sanctuary in western Myanmar, where they reportedly predated on livestock. Apparently, local people persecuted Dholes in retribution, and consequently Dholes were not detected in Chatthin during recent camera-trapping surveys (N. M. Shwe unpubl. data). Since 2005, Dholes have been recorded from several protected areas in Myanmar, including the northern region (Hukaung Valley Wildlife Sanctcuary and Htamanthi Wildlife Sanctuary), west-central region (Mahamyaing Wildlife Sanctuary and Namataung National Park), southwestern region (Rakhine Yoma Elephant Range), and peninsula region (Tanintharyi Nature Reserve; WCS Myanmar Program unpubl. data). From 1999-2002, Dholes were recorded in at least eight additional forest tracts and protected areas the in northern, western, and central parts of the country (WCS Myanmar Program unpubl. data), thus Dholes may still occur throughout these regions. There are few confirmed records of Dholes from eastern Myanmar, although they likely occur near the tri-border area with Lao PDR and Thailand, as there are records in that area from those countries. There are recent records of dholes from camera-trap studies in Karen (Kayin) State (R. McEwing pers. comm.), but their status in other areas of eastern Myanmar is not known.

Nepal: Dhole sightings in Nepal are not common, yet there are recent reports of this species in several areas of country. In the Himalayan region, Dholes reportedly occur in the western (Rara and Khaptad National Parks, Dhorpatan Hunting Reserve; R. D. Choudhary pers. comm., A. Aryal in litt.) and extreme eastern parts (Kanchenjunga Conservation Area; Khatiwada 2011) of the country. In southern Nepal, Dholes are found throughout much of the Terai Arc Landscape, including Chitwan and Bardia national parks (Thapa et al. 2013, A. Khatiwada pers. comm.), and Parsa and Shuklaphanta Wildlife Reserves (A. Khatiwada pers. comm.). Dholes also are reported in several districts of Nepal outside protected areas, but all populations are extremely low and threatened by low prey base, poisoning and habitat loss (A. Khatiwada pers. comm.).

Pakistan: Although there are no confirmed records of Dholes from Pakistan, they historically occurred in the western Himalayan Mountains in the northern part of the country. They might still occur in the Ladakh region of northern Parkistan, as Dholes recently have been recorded in the India-administered region of Ladakh.

Thailand: Dholes have been extirpated from most areas of Thailand, and are now absent from the peninsula and eastern parts of the country. Dholes are still found in fragmented populations in several large protected-area complexes, including the south-central region (Phayayen-Khao Yai and Eastern Forest complexes), north-central region (Phumieng-Phuthong and Phukhieo-Namnow complexes), northwestern region (Srilanna-Khutan and Doi Phucar-Maeyom complexes), and western region (Western Forest Complex; Jenks et al. 2012; Department of National Park, Wildlife and Plant Conservation, Thailand). It is not yet known how stable are these isolated populations, and if the protected-area complexes are large enough to conserve viable Dhole populations in the long term.

Viet Nam: There are very few recent confirmed records on Dholes in Viet Nam. The last confirmed records of Dholes were in Pu Mat National Park in 1998-99 (D. Willcox pers. comm.) and in Yok Don National Park in 2003 (J. Eames pers. comm.) despite extensive camera trapping in >25 protected areas throughout the country. Along with other large carnivores, Dholes are likely extirpated from Viet Nam, although individuals may occasionally enter the country from neighboring Cambodia or Lao PDR

Legal protection
It is included in CITES – Appendix II (2013). Dholes are legally protected in the countries where they occur. However, enforcement of laws is insufficient to provide effective protection of Dholes in many of their range countries. Local governments sometimes may still offer bounties on Dholes to reduce livestock predation, as was recently the case in western Myanmar. In Thailand, Dholes were recently blamed for the decline of wild ungulates in some protected areas, and as a result some government officials have proposed to eliminate Dholes from those areas (K. Jenks and N. Songsasen unpubl.). Dholes also may have been intentionally extirpated from some protected areas by local officials in southeastern China in an attempt to boost ungulate numbers (S. Li pers. comm.). Providing compensation, incentives or insurance for livestock-Dhole conflicts to reduce retaliatory killing of Dholes may be a good conservation strategy and should be tested (Dickman et al. 2011, Gurung et al. 2011). Similarly, incentives for conservation of habitat, Dholes and their prey should be explored.

Presence in protected areas
Although this species occurs in protected areas throughout its range, there are no conservation measures specifically focused on Dholes, except for a few isolated localities like eastern Nepal. For the putative northern Dholes, their occurrence in China was recently confirmed by camera traps in several isolated nature reserves. However, lack of data on numbers of Dholes and their prey in these reserves prevent a valid assessment as to their potential for conserving Dholes in the long-term. For the putative southern Dholes, both Project Tiger and Project Elephant in India have the potential to conserve populations of Dholes and their prey in areas where they coexist with tigers and elephants. However, Dholes require up to five times the land area as tigers to maintain viable long-term populations, and consequently Dholes have disappeared from more reserves than have tigers (Woodroffe and Ginsberg 1998). Thus, relatively large (>750 km²) reserves in India might be the most effective for conserving Dhole populations. Large protected-area complexes for Tiger conservation in Bhutan, Malaysia, Myanmar, Nepal, Indonesia (Sumatra), and Thailand, also are conserving Dhole populations, thus these areas hold the greatest potential for the long-term conservation of Dholes in South and Southeast Asia. Consolidating more forest areas and including them in protected area networks would greatly enhance the conservation of Dholes in these regions. 

Presence in captivity
As of August 2013, there were at least 223 Dholes in 38 zoos worldwide (International Species Information System [ISIS] unpubl.), including zoos in Europe (24 zoos), Asia (nine zoos), North America (four zoos), and Australia (one zoo). There also are captive Dholes in additional zoos and breeding farms which are not members of ISIS. The origin of most captive Dholes is unclear, and their subspecific classification is probably wrong. The most numerous subspecies in captivity is listed as C. a. lepturus, which occurs in at least 20 zoos worldwide and is the most common Dhole in European zoos. Firstly, inbreeding may be an issue because captive Dholes listed as lepturus trace their origin to only three founders: a single Dhole from a game farm in North America with an unknown origin (H. Maisch pers. comm.), and Dholes from the Moscow Zoo, which originated from only two individuals captured in Qinghai Province, China in 1957 (Sosnovskii 1967). Secondly, the Dholes captured in Qinghai Province should represent either C. a. hesperius or C. a. fumosus, from the putative northern Dhole group, rather than lepturus which historically occurred only south of the Yangtze River and is part of the southern Dhole group (Durbin et al. 2004). The putative southern Dholes are represented in several Indian zoos (probably C. alpinus dukhunensis), and in zoos in Phnom Penh, Cambodia, and Sydney, Australia (C. alpinus infuscus). Other zoos do not list subspecies, thus it is likely that putative subspecies from different origins have been interbred (M. Boeer pers. comm.), such as that done in Singapore Zoo. The European Endangered Species Programme (EEP) does not consider subspecies, but it does regard Dholes in European zoos as a Chinese ecotype, and prevents mixing this type with Dholes from other origins (e.g., India, Cambodia). Nevertheless, the value of any of captive Dholes for potential reintroduction efforts is uncertain, at least until genetic studies can confirm their origin and subspecific classification. Until that time, we recommend that captive Dholes from the putative northern and southern groups be managed separately, such as that done by the EEP.

Research needs
More research is needed on Dholes to better understand their ecology and assist conservation efforts. These include: 1) develop cost-effective surveys to determine the abundance of Dholes, as data on Dhole numbers would allow us to better understand their conservation status; 2) investigate the genetic and morphological differences between the putative northern and southern Dholes, and the distinctiveness of other putative subspecies such as the Sumatran and Javan Dholes; 3) determine the area and prey requirements needed to maintain a viable Dhole population; 4) investigate the effects of disease on Dhole population dynamics, and; 5) investigate effects of Dholes on ecosystems, specifically their interactions with other large carnivores, and their impacts on prey and smaller carnivores.