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 (Mdog) 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
From the Altai Mountains in Manchuria in Central and Eastern Asia, its range spreads southwards through the forest tracts of India, Burma, and the Malayan Archipelago. Three races of the dhole exist in India alone
(Trans-Himalayan, Himalayan, and Peninsular).
Biogeographic Regions: palearctic (Native ); oriental (Native )
C.I.S., Korea, China, India, Southeast Asia
The dhole is an average size canine with head/body length 90cm (35"), tail length 40-45cm (16"-18"), and shoulder height 50cm (20"). The dhole is set apart from other canids in that it has an unusually thick muzzle and one less molar tooth on each side of its lower jaw. Other members of the family Canidae have a total of 42 teeth. The adult dhole is characterized by a rusty red coat with a pale underside; depending on the region, pelage may vary from light brownish gray to a uniform red coat. A dhole is born with a sooty brown color, acquiring an adult color at three months of age. Dholes also have dark, almost always black, bushy tails.
Range mass: 17 to 21 kg.
Other Physical Features: endothermic ; bilateral symmetry
Habitat and Ecology
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 Dholes 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).
Dholes like open spaces and can often be found on jungle roads, river beds, jungle clearings, and paths, where they rest during the day. Their hunting range is about 40sq km (15sq mi). The dhole can also be found in dense forest steppes, and the thick jungles of the plains as well as the hills. They are never found in the open plains and deserts.
Terrestrial Biomes: forest ; rainforest ; scrub forest
The dhole eats wild berries, insects, and lizards. Packs of dholes feast on mammals ranging from rodents to deer. Some of the dhole's favorites include wild pigs, hares, wild goats, sheep, and occasionally a monkey. Unlike many other "dogs," the dhole seldom kills by biting the throat. Larger mammals are attacked from the rear, while smaller ones are caught by any part of the bodies. The smaller mammals are killed by a swift blow to the head; the larger mammals are immediately disembowled. Dholes compete for the food, not by fighting, but by how fast they can eat. An adult dhole can eat up to 4kg (8.8lbs) of meat in one hour. Two to three dholes can kill a 50kg (110 lb) deer in less than two minutes, and they begin to feed on it before it is dead. The larger prey rarely die from the attack itself, but from blood loss and shock as their intestines, heart, liver, and eyes are feasted upon.
Animal Foods: mammals; reptiles; insects
Plant Foods: fruit
Primary Diet: omnivore
Known prey organisms
This list may not be complete but is based on published studies.
Life History and Behavior
Perception Channels: tactile ; chemical
Status: captivity: 16.0 years.
Lifespan, longevity, and ageing
Each pack contains a dominant monogamous pair. Subordinate pack members help care for the young of the dominant pair.
Mating System: monogamous ; cooperative breeder
The dhole's gestation period is 60-62 days. The mother usually gives birth to eight pups at a time. The pups reach sexual maturity at about a year. Pups are born throughout the end of fall, winter, and the first spring months ( November - March ). Female dhole can have up to 16 mammae, suggesting their ability to take care of large litters. Dens are constructed near streambeds or among rocks. After a female dhole has given birth, a few other adults take part in feeding the mother as well as the pups. The pups, as early as the tender age of three weeks, and the mother are fed regurgitated meat.
Range number of offspring: 2 to 6.
Average number of offspring: 3.5.
Range gestation period: 60 to 63 days.
Average weaning age: 58 days.
Key Reproductive Features: gonochoric/gonochoristic/dioecious (sexes separate); sexual
Average birth mass: 275 g.
Average number of offspring: 5.
Average age at sexual or reproductive maturity (male)
Sex: male: 365 days.
Average age at sexual or reproductive maturity (female)
Sex: female: 365 days.
Parental Investment: altricial ; post-independence association with parents; extended period of juvenile learning
Molecular Biology and Genetics
Barcode data: Cuon alpinus
No available public DNA sequences.
Download FASTA File
Statistics of barcoding coverage: Cuon alpinus
Public Records: 2
Specimens with Barcodes: 2
Species With Barcodes: 1
IUCN Red List Assessment
Red List Category
Red List Criteria
- Endangered (EN)
- 2004Endangered (EN)
- 1996Vulnerable (VU)
- 1994Vulnerable (V)
- 1990Vulnerable (V)
- 1988Vulnerable (V)
- 1986Vulnerable (V)
There are 10 subspecies of the dhole ranging in color and size. Two of the subspecies are listed as endangered by the IUCN (east asian dhole and the west indian dhole). Two other subspecies are on the verge of extinction ( C.a. primaerus, and the C.a. laniger).
CITES: appendix ii
IUCN Red List of Threatened Species: endangered
Date Listed: 06/02/1970
Lead Region: Foreign (Region 10)
Where Listed: Entire
Population location: Entire
Listing status: E
For most current information and documents related to the conservation status and management of Cuon alpinus , see its USFWS Species Profile
Because so little is known about Dhole ecology, we used African Wild Dogs (Lycaon pictus) as a surrogate species for estimating the proportion of mature individuals and generation length. Of the canids, African Wild Dogs are arguably most similar to Dholes. Both species have specialized dental morphology for obligate hypercarnivory (Van Valkenburgh 1991), which is rather unique among canids. Phylogenetic research shows both species are in the same monophyletic group (Lindblad-Toh et al. 2005), and otherwise both species are similar in body size, reproduction, social behaviour and feeding behaviour (Johnsingh 1982). Consequently, much of the text below comes directly from the IUCN assessment for the African Wild Dog (Woodroffe and Sillero-Zubiri 2012).
Proportion of mature individuals
Estimating the number of mature individuals is challenging for Dholes, because like African Wild Dogs, they are obligate cooperative breeders (Johnsingh 1982, Venkataraman 1998). Consequently, within a pack the alpha male and female are the parents of the majority of surviving pups (Venkataraman 1998). Although the IUCN Red List Categories and Criteria (IUCN 2012) define mature individuals as individuals known, estimated or inferred to be capable of reproduction, it does not specify the time period within which reproduction is considered possible. The User Guidelines (IUCN 2014) go on to state in many taxa there is a pool of non-reproductive (e.g., suppressed) individuals that will quickly become reproductive if a mature individual dies. These individuals can be considered to be capable of reproduction. As with African Wild Dogs, a high proportion of individual Dholes within a park are indeed reproductively suppressed (Johnsingh 1982, Venkataraman 1998), but these animals do not necessarily become reproductive quickly if an alpha individual dies. In a mature pack, most pack members are offspring of the alpha pair; for these animals, death of an alpha would usually not open up a breeding opportunity because no unrelated mates would be available within the pack. Given these complexities, and in keeping with the spirit of capturing a snapshot of current conditions, we have chosen to define mature individuals as those considered capable of reproduction within the current breeding season. The number of mature individuals thus comprises the number of alpha males and females, and the number of sub-dominant (i.e. non-alpha) animals that breed successfully.
Following that reported for Africa Wild Dogs (Woodroffe and Sillero-Zubiri 2012), we estimated for Dholes the number of mature individuals (Nm) from populations of adults and yearlings (Nc). The number of alpha males (NaM) and alpha females (NaF) would be estimated with the following equations:
NaM = Nc x 0.55 x 0.176NaF = Nc x 0.45 x 0.215
The above equations result in approximately equal estimates of the numbers of alpha males and alpha females. Also following that reported for African Wild Dogs (Woodroffe and Sillero-Zubiri 2012), we estimated for Dholes the number of sub-dominant breeders (Nsub) as:
Nsub = (NaM x 0.10) + (NaF x 0.08)
Therefore, the number of mature individuals (Nm) for Dholes could be calculated using the following equation:
Nm = NaM + NaF + Nsub
Population size and densities
Population estimates of Dholes are not available for any country. Therefore, we made an attempt to estimate the total population of Dholes by classifying countries within their current distribution as having high (1,500-3,000), medium (750-1,500), or low (250-750) numbers of Dholes. These classifications were based on estimates of relative abundances and area covered by Dholes within each country. We classified one country as having high numbers of Dholes (India), two countries with medium numbers (Thailand, Myanmar), six countries with low numbers (Bhutan, Cambodia, China, Lao PDR, Malaysia, Nepal), and four countries with negligible numbers (Bangladesh, Korea DPR, Pakistan, Viet Nam). Consequently, we estimate the total population of Dholes to be 4,50010,500. Although there is relatively high uncertainty with this population estimate, it has a large range and we feel that it adequately represents the possible population size of Dholes.
To calculate total population of mature individuals, we used the equations above to show an estimate of 436-1,016 alpha males (NaM), 435-1016 alpha females (NaF), and 78-183 sub-dominant breeders (Nsub). This resulted in an estimate of 949-2,215 mature individuals (Nm), which is below the 2,500 threshold (i.e., one of the criteria for listing as EN under C).
The only estimates of local Dhole densities come from a few protected areas in India. Only one of those estimates has been obtained through systematic sample-based survey methods (Selvan et al. 2014), whereas the others are presumably based on estimates of the number of packs within the protected areas (derived using known home range areas and knowledge of mean pack sizes). Reported Dhole densities were 0.066 Dholes/km2 in Pakke Tiger Reserve (Selvan et al. 2014), 0.095 Dholes/km2in Mudumalai Wildlife Sanctuary, 0.13 Dholes/km2in Bandipur Tiger Reserve, and 0.3 Dholes/km2in Pench National Park (Durbin et al. 2004).
The size of subpopulations of Dholes has not been reported anywhere. Therefore, the following is our estimate of the size of one of the largest subpopulations of Dholes, which is presumed to be in the Western Ghats of India, based on high prey numbers and extent of protected forests in the region. This region contains the most intensively studied subpopulation of Dholes in the world (Johnsingh 1982, Venkataraman et al. 1995, Karanth and Sunquist 1995, Venkataraman 1998, Karanth and Sunquist 2000, Andheria et al. 2007, Ramesh et al. 2012, Srivathsa et al. 2014). Srivathsa et al. (2014) estimated the proportion of area occupied by Dholes in the Malenad landscape (Western Ghats in the State of Karnataka), by assessing "true occupancy" in 206 grid-cells (188km2each). This area covers 16 wildlife reserves, where the highest numbers of Dholes are likely restricted to four source populations: Anshi-Dandeli, Bhadra, Nagarahole-Bandipur and Biligiri Rangaswamy Temple (BRT) reserves. Abundance was derived from estimates of occupancy, based on the Royle-Nichols (2003) occupancy model [abundance in a grid-cell= Log (1/(1-PSI))] for all the 206 grid-cells. We calculated a range of abundances for each source population, which likely represent one metapopulation. The lower limit of this range is the sum of grid-cell-wise abundances from those cells that intersect with the reserve boundaries; the upper limit of this range is the sum of grid-cell-wise abundances for the reserve+the cells that are first order neighbours for the reserve cells (with contiguous forest habitats). The following are the abundance ranges for the four source populations: Anshi-Dandeli = 15-46; Bhadra = 23-39; Nagarahole-Bandipur = 85-91, BRT = 24-28. We estimate the remaining forested landscape may support 60-100 Dholes, resulting in a total population of 207-304 individuals. To calculate total number of mature individuals in this subpopulation, we used the above equations which resulted in an estimate of 44-64 mature individuals (20-29 alpha males, 20-29 alpha females, four to six sub-dominant breeders). Clearly, even one of the largest subpopulations of Dholes contains well below 250 mature individuals (i.e., the criterion for listing as EN under C2a(i)).
Change in population size
There is almost no quantitative information on Dhole population trends through their distribution. In fact, a common characteristic of Dhole populations is that they often exhibit severe local population fluctuations (Johnsingh 1982, Venkataraman 1998, Karanth and Sunquist 2000, Durbin et al. 2004, J. Kamler pers. obs.), which makes estimating populations and population trends difficult. We estimate that the total population of Dholes is still decreasing, due to the continuous decline in their distribution.
Habitat loss and transformation: Habitat loss and degradation are serious threats to Dholes in southern Asia, particularly because this threat is closely associated with prey depletion and high levels of human disturbance. Although extensive areas of natural or semi-natural vegetation remain in Lao PDR and Cambodia, habitat conversion and fragmentation are proceeding unabated. In Viet Nam, very few natural areas greater than 50 km2 remain. Habitat loss and fragmentation is a major threat to protected areas in Indonesia, particularly those on Sumatra. Habitat loss is driven by several different factors, including logging, palm and rubber plantations, agriculture expansion, rural biomass extraction, livestock grazing and major infrastructure expansion (e.g., hydropower dams, irrigation projects, new highways, etc.).
Persecution: Persecution of Dholes stems mainly from retaliatory killings due to livestock predation, and this factor is driving some Dhole populations towards local extinction (Lyngdoh et al. 2014). Dholes appear to be especially susceptible to poisoning of carcasses using strychnine or other rodenticides, which often are readily available to rural people in southern Asia. Consequently, unsystematic but consistent poisoning of carcasses can easily wipe out Dholes within an area, especially because the entire pack will feed on a carcass. For example, poisoning of livestock carcasses apparently wiped out Dholes from Bhutan in the 1970s and early 1980s, although this species began to re-occupy the country starting in the 1990s (Wangchuk 2004, Thinley et al. 2011). Killing of Dholes by poisoning livestock carcasses also has been reported in China, Nepal, India and Indonesia, and is likely is widespread in southern Asia. Poisoning campaigns also were thought to have contributed to the extirpation of Dholes in the countries of the former Soviet Union (Ginsberg and Macdonald 1990).
Dholes reportedly have been shot by humans, as Dhole carcasses with gun-shot wounds were found near the boundary of protected areas in Thailand (K. Jenks and N. Songsasen unpubl.). Dholes are probably susceptible to non-selective snaring, particularly where this activity is widespread such as in Viet Nam, Cambodia, China and Lao PDR. In India, and possibly elsewhere, Dholes living outside or on the edge of core protected areas are particularly vulnerable to human kleptoparasitism.
Disease and pathogens: Dholes are susceptible to rabies, canine distemper, canine parvovirus and sarcoptic mange among others (Durbin et al. 2004), which are usually contracted from domestic village dogs that act as reservoirs. Dholes appear to be especially susceptible to disease epizootics due their large pack sizes and high levels of amicable behaviour within packs, even among adults (Johnsingh 1982). Such behaviours likely result in relatively high intraspecific contact rates, which are conducive for disease epizootics, at least compared to other wild canids such as jackals (Canis aureus) which are more solitary. Disease epizootics may contribute to the sudden disappearance of Dholes from protected areas, and often cause severe local population fluctuations resulting in relatively small pack sizes (Karanth and Sunquist 2000, J. Kamler pers. obs.). In Cambodia, a recent outbreak of canine distemper in 2011-2012, thought to have originated in local domestic dogs, caused the near-extirpation of Dholes from protected areas in the eastern and northern parts of the country (J. Kamler unpubl.), although populations appear to be recovering; mortalities associated with canine distemper also have been observed in a protected area in Thailand (N. Songsasen and K. Jenks unpubl).The range-wide effects of diseases on Dhole population dynamics is unknown, but it is likely significant across southern Asia, and might result in an increased probability of extirpation for Dhole populations in isolated protected areas.
Competition with other species: Aside from humans, the main competitors of Dholes for limited resources are Tigers (Panthera tigris) and Leopards (P. pardus). Although Dholes are much smaller in body size, packs of Dholes reportedly have killed both Tigers and Leopards, although the reverse also has been reported (Burton 1940). The dominance hierarchy between Dholes and Tigers is not clear, although Dholes likely avoid tigers especially if packs are small. Dholes appear to be behaviourally dominant over Leopards, and packs of Dholes often tree this species when they interact (Venkataraman 1995). Whether large felids can negatively affect Dhole numbers is unknown, although the exploitive and interference competition between them likely becomes more intense as prey populations are reduced by humans, possibly resulting in spatial exclusion where prey numbers are lowest. Free-ranging dogs also may compete with Dholes for limited food resources where prey numbers are low.
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.
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.
Relevance to Humans and Ecosystems
Although this occurs on rare occasions, dholes can attack livestock at the cost of the owner.
Dholes have become an indirect food source for the residents of the jungles. Dholes do not attack human beings, and they usually retreat at the sight of a person. Human residents of the jungle follow dholes when they are hunting. When the dhole ccompletes its kill, the human hunters scare it away and steal its kill.
The dhole (Cuon alpinus) is a canid native to Central and Southeast Asia. Other English names for the species include Indian wild dog, whistling dog, chennai, Asiatic wild dog, red wolf (not to be confused with Canis rufus), red dog and mountain wolf. It is genetically close to species within the genus Canis,(Fig. 10) though its skull is convex rather than concave in profile, it lacks a third lower molar, and the upper molars sport only a single cusp as opposed to 2–4. During the Pleistocene, the dhole ranged throughout Asia, Europe and North America, but became restricted to its historical range 12,000–18,000 years ago.
The dhole is a highly social animal, living in large clans without rigid dominance hierarchies and containing multiple breeding females. Such clans usually consist of 12 individuals, but groups of over 40 are known. It is a diurnal pack hunter which preferentially targets medium and large sized ungulates. In tropical forests, the dhole competes with tigers and leopards, targeting somewhat different prey species, but still with substantial dietary overlap.
It is listed as Endangered by the IUCN, as populations are decreasing and estimated at less than 2,500 adults. Factors contributing to this decline include habitat loss, loss of prey, competition with other species, persecution, and disease transfer from domestic dogs.
- 1 Characteristics
- 2 Distribution and habitat
- 3 Ecology and behaviour
- 4 Threats
- 5 Conservation
- 6 Discovery, taxonomy and evolution
- 7 In culture and literature
- 8 Etymology and naming
- 9 See also
- 10 References
- 11 External links
In appearance, the dhole has been variously described as combining the physical characteristics of the grey wolf and red fox, and as being "cat-like" on account of its long backbone and slender limbs. It has a wide and massive skull with a well-developed sagittal crest, and its masseter muscles are highly developed compared to other canid species, giving the face an almost hyena-like appearance. The rostrum is shorter than that of domestic dogs and most other canids. The species has six rather than seven lower molars. The upper molars are weak, being one-third to one-half the size of those of wolves, and have only one cusp as opposed to 2–4, as is usual in canids, an adaptation thought to improve shearing ability, thus allowing it to compete more successfully with kleptoparasites. Adults may weigh over 18 kg (40 lb), with females usually weighing 4.5 kg (9.9 lb) less than males. It stands 17–22 inches at the shoulder and measures three feet in body length. Like the African wild dog, its ears are rounded rather than pointed. It has 6–7 teats, sometimes eight.
The general tone of the fur is reddish, with the brightest hues occurring in winter. In the winter coat, the back is clothed in a saturated rusty-red to reddish colour with brownish highlights along the top of the head, neck and shoulders. The throat, chest, flanks, belly and the upper parts of the limbs are less brightly coloured, and are more yellowish in tone. The lower parts of the limbs are whitish, with dark brownish bands on the anterior sides of the forelimbs. The muzzle and forehead are greyish-reddish. The tail is very luxuriant and fluffy, and is mainly of a reddish-ocherous colour, with a dark brown tip. The summer coat is shorter, coarser and darker. The dorsal and lateral guard hairs in adults measure 20–30 mm in length. Dholes in the Moscow Zoo moult once a year from March to May.
Dholes produce whistles resembling the calls of red foxes, sometimes rendered as coo-coo. How this sound is produced is unknown, though it is thought to help in coordinating the pack when travelling through thick brush. When attacking prey, they emit screaming KaKaKaKAA sounds. Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries. In contrast to wolves, dholes do not howl or bark. Dholes have a complex body language. Friendly or submissive greetings are accompanied by horizontal lip retraction and the lowering of the tail, as well as licking. Playful dholes will open their mouths with their lips retracted and their tails held in a vertical position whilst assuming a play bow. Aggressive or threatening dholes will pucker their lips forward in a snarl and raise the hairs on their backs, as well as keep their tails horizontal or vertical. When afraid, they pull their lips back horizontally with their tails tucked and their ears flat against the skull.
Distribution and habitat
There are currently no confirmed recent reports of dhole being present in Russia, Mongolia, Kazakhstan, Kyrgyzstan or Tajikistan, though one specimen was caught in southern China's Jiangxi district. It is unknown if dholes continue to inhabit Tien Shan, though they possibly occur in small numbers in Gansu Province, with one pack being sighted in the Qilian Mountains within that province in 2006. Dholes still occur in Tibet, and may still inhabit North Korea. Although they have not been recorded in Pakistan, they once occurred in the alpine steppes extending into Kashmir. They occur in most of India south of the Ganges, particularly in the Central Indian Highlands and the Western and Eastern Ghats of the southern states. In north-east India, they inhabit Arunachal Pradesh, Assam, Meghalaya, and West Bengal. The situation of dholes in the Himalaya and north-west India is precarious, and populations fragmented. They may occur in Kashmir's Ladakh area.
Dholes once occurred in the Indo-Gangetic Plain's Terai region. In 2011, dhole packs were recorded by camera traps in the Chitwan National Park. Their presence was confirmed in the Kanchenjunga Conservation Area in 2011 by camera traps.
It is unknown whether the species still lives in Bangladesh, where it once inhabited the forested areas of the Chittagong and Sylhet District. The presence of dholes in Myanmar was confirmed by camera trapping in 11 areas and, alongside leopards, have apparently replaced tigers as the country's top predators.
Their range is highly fragmented in the Malaysian Peninsula, Sumatra, Java, Vietnam and Thailand. A camera trapping survey in the Khao Ang Rue Nai Wildlife Sanctuary during January 2008 to February 2010 revealed at least one healthy dhole pack.
In Central Asia, dholes primarily inhabit mountainous areas; in the western half of its range, they live mostly in alpine meadows and high-montane steppes high above sea level, while in the east, they mainly ranges in montane taigas, though may appear along coastlines. In India, Myanmar, Indochina, Indonesia and China, they prefer forested areas in alpine zones, and occasionally also in plains regions.
Ecology and behaviour
Social and territorial behaviour
Dholes are more social than grey wolves, and have less of a dominance hierarchy, as seasonal scarcity of food is not a serious concern for them. In this manner, they closely resemble African wild dogs in social structure. They live in clans rather than packs, as the latter term refers to a group of animals that always hunt together. In contrast, dhole clans frequently break into small packs of 3–5 animals, particularly during the spring season, as this is the optimal number for catching fawns. Dominant dholes are hard to identify, as they do not engage in dominance displays as wolves do, though other clan members will show submissive behaviour toward them. Intragroup fighting is rarely observed. Dholes are far less territorial than wolves, with pups from one clan often joining another without trouble once they mature sexually. Clans typically number 5-12 individuals in India, though clans of 40 have been reported. In Thailand, clans rarely exceed three individuals. Unlike other canids, there is no evidence of dholes using urine to mark their territories or travel routes. They may defecate in conspicuous places, though a territorial function is unlikely, as faeces are mostly deposited within the clan's territory rather than the periphery. Faeces are often deposited in what appear to be communal latrines. They do not scrape the earth with their feet as other canids do to mark their territories.
Four kinds of den have been described; simple earth dens with one entrance (usually remodeled striped hyena or porcupine dens); complex cavernous earth dens with more than one entrance; simple cavernous dens excavated under or between rocks; and complex cavernous dens with several other dens in the vicinity, some of which are interconnected. Dens are typically located under dense scrub or on the banks of dry rivers or creeks. The entrance to a dhole den can be almost vertical, with a sharp turn three to four feet down. The tunnel opens into an antechamber, from which extends more than one passage. Some dens may have up to six entrances leading up to 100 feet (30 m) of interconnecting tunnels. These "cities" may be developed over many generations of dholes, and are shared by the clan females when raising young together. Like African wild dogs and dingoes, dholes will avoid killing prey close to their dens.
Reproduction and development
In India, the mating season occurs between mid-October and January, while captive dholes in the Moscow Zoo breed mostly in February. Unlike wolf packs, dhole clans may contain more than one breeding female. More than one female dhole may den and rear their litters together in the same den. During mating, the female assumes a crouched, cat-like position. There is no copulatory tie characteristic of other canids when the male dismounts. Instead, the pair lie on their sides facing each other in a semicircular formation. The gestation period lasts 60–63 days, with litter sizes averaging 4–6 pups. Their growth rate is much faster than that of wolves, being similar in rate to that of coyotes. Pups are suckled at least 58 days. During this time, the pack feeds the mother at the den site. Dholes do not use rendezvous sites to meet their pups as wolves do, though one or more adults will stay with the pups at the den while the rest of the pack hunts. Once weaning begins, the adults of the clan will regurgitate food for the pups until they are old enough to join in hunting. They remain at the den site 70–80 days. By the age of six months, pups accompany the adults on hunts, and will assist in killing large prey such as sambar by the age of eight months. Maximum longevity in captivity is 15–16 years.
Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and homo- and heterosexual mounting. Dholes are primarily diurnal hunters, hunting in the early hours of the morning. They rarely hunt nocturnally, except on moonlit nights, indicating they greatly rely on sight when hunting. Though not as fast as jackals and foxes, they can chase their prey for many hours. During a pursuit, one or more dholes may take over chasing their prey, while the rest of the pack keeps up at a steadier pace behind, taking over once the other group tires. Most chases are short, lasting only 500 m. When chasing fleet-footed prey, they run at a pace of 30 mph. Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.
Once large prey is caught, one dhole will grab the prey's nose, while the rest of the pack pulls the animal down by the flanks and hindquarters. They do not use a killing bite to the throat. They occasionally blind their prey by attacking the eyes. Serows are among the only ungulate species capable of effectively defending themselves against dhole attacks, due to their thick, protective coats and short, sharp horns capable of easily impaling dholes. They will tear open their prey's flanks and disembowel it, eating the heart, liver, lungs and some sections of the intestines. The stomach and rumen are usually left untouched. Prey weighing less than 50 kg is usually killed within two minutes, while large stags may take 15 minutes to die. Once prey is secured, dholes will tear off pieces of the carcass and eat in seclusion. Unlike wolf packs, in which the breeding pair monopolises food, dholes give priority to the pups when feeding at a kill, allowing them to eat first. They are generally tolerant of scavengers at their kills. Both mother and young are provided with regurgitated food by other pack members.
Prey animals in India include chital, sambar, muntjac, mouse deer, swamp deer, wild boar, gaur, water buffalo, banteng, cattle, nilgai, goats, Indian Hares, Himalayan Field Rats and langurs. There is one record of a pack bringing down an Indian elephant calf in Assam, despite desperate defense of the mother resulting in numerous losses to the pack. In Kashmir, they may hunt markhor, and thamin in Burma. Javan Rusas are hunted in Java. In the Tien Shan and Tarbagatai Mountains, dholes prey on Siberian ibexes, arkhar, roe deer, maral and wild boar. In the Altai and Sayan Mountains, they prey on musk deer and reindeer. In eastern Siberia, they prey on roe deer, Manchurian wapiti, wild boar, musk deer, and reindeer, while in Primorye they feed on sika deer and goral, too. In Mongolia, they prey on argali and rarely Siberian ibex. Like African wild dogs, but unlike wolves, dholes are not known to attack people. Dholes eat fruit and vegetable matter more readily than other canids. In captivity, they eat various kinds of grasses, herbs and leaves, seemingly for pleasure rather than just when ill. In summertime in the Tien Shan Mountains, dholes eat large quantities of mountain rhubarb. Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves. Livestock predation by dholes has been a problem in Bhutan since the late 1990s, as domestic animals are often left outside to graze in the forest, sometimes for weeks at a time. Livestock stall-fed at night and grazed near homes are never attacked. Oxen are killed more often than cows are, probably because they are given less protection.
Enemies and competitors
In some areas, dholes are sympatric to tigers and leopards. Competition between these species is mostly avoided through differences in prey selection, although there is still substantial dietary overlap. Along with leopards, dholes typically target animals in the 30–175 kg range (mean weights of 35.3 kg for dhole and 23.4 kg for leopard), while tigers selected for prey animals heavier than 176 kg (but their mean prey weight was 65.5 kg). Also, other characteristics of the prey, such as sex, arboreality, and aggressiveness, may play a role in prey selection. For example, dholes preferentially select male chital, whereas leopards kill both sexes more evenly (and tigers prefer larger prey altogether), dholes and tigers kill langurs rarely compared to leopards due to the leopards' greater arboreality, while leopards kill wild boar infrequently due to the inability of this relatively light predator to tackle aggressive prey of comparable weight.
On some rare occasions, dholes may attack tigers. When confronted by dholes, tigers will seek refuge in trees or stand with their backs to a tree or bush, where they may be mobbed for lengthy periods before finally attempting escape. Escaping tigers are usually killed, while tigers which stand their ground have a greater chance of survival. Tigers are extremely dangerous opponents for dholes, as they have sufficient strength to kill a single dhole with one paw strike. Even a successful tiger kill is usually accompanied by losses to the pack. Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs. Since leopards are smaller than tigers and more likely hunt dholes, dhole packs tend to react more aggressively toward them than they do towards tigers.
There are numerous records of leopards being treed by dholes. Dholes sometimes drive tiger, leopards, and bears (see below) from their kills. Dholes were once thought to be a major factor in reducing Asiatic Cheetah populations, though this is doubtful, as cheetahs live in open areas as opposed to forested areas favoured by dholes.
Though usually antagonistic toward wolves, they may hunt and feed alongside one another. There is at least one record of a lone wolf associating with a pair of dholes in Debrigarh Wildlife Sanctuary. They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion.
Diseases and parasites
Dholes are vulnerable to a number of different diseases, particularly in areas where they are sympatric with other canid species. Infectious pathogens such as Toxocara canis are present in their faeces. They may suffer from rabies, canine distemper, mange, trypanosomiasis, canine parvovirus, and endoparasites such as cestodes and roundworms.
The dhole only rarely takes domestic livestock. Certain people, such as the Kurumbas and some Mon Khmer-speaking tribes will appropriate dhole kills; some Indian villagers welcome the dhole because of this appropriation of dhole kills. Dholes were persecuted throughout India for bounties until they were given protection by the Wildlife Protection Act of 1972. Methods used for dhole hunting included poisoning, snaring, shooting and clubbing at den sites. Native Indian people killed dholes primarily to protect livestock, while British sporthunters during the British Raj did so under the conviction that dholes were responsible for drops in game populations. Persecution of dholes still occurs with varying degrees of intensity according to region. Bounties paid for dholes used to be 25 rupees, though this was reduced to 20 in 1926 after the number of presented dhole carcasses became too numerous to maintain the established reward. In Indochina, dholes suffer heavily from nonselective hunting techniques such as snaring.
The fur trade does not pose a significant threat to dholes. The people of India do not eat dhole flesh, and their fur is not considered overly valuable. Due to their rarity, dholes were never harvested for their skins in large numbers in the Soviet Union, and were sometimes accepted as dog or wolf pelts (being labeled as "half wolf" for the latter). The winter fur was prized by the Chinese, who bought dhole pelts in Ussuriysk during the late 1860s for a few silver rubles. In the early 20th century, dhole pelts reached eight rubles in Manchuria. In Semirechye, fur coats made from dhole skin were considered the warmest, but were very costly.
The dhole is protected under Schedule 2 of the Wildlife Protection Act, 1972. The creation of reserves under Project Tiger provided some protection for dhole populations sympatric with tigers. In 2014, the Indian government sanctioned its first dhole conservation breeding centre at the Indira Gandhi Zoological Park (IGZP) in Visakhapatnam. The dhole has been protected in Russia since 1974, though it is vulnerable to poison let out for wolves. In China, the animal is listed as a category II protected species under the Chinese wildlife protection act of 1988. In Cambodia, the dhole is protected from all hunting, while conservation laws in Vietnam limit extraction and utilization.
Discovery, taxonomy and evolution
The species was first described in literature in 1794 by an explorer named Pesteref, who encountered dholes during his travels in far eastern Russia. He described the animal as being a regular pack hunter of Alpine ibex, and of bearing many similarities with the golden jackal. It was given the binomial name Canis alpinus in 1811 by Peter Pallas, who described its range as encompassing the upper levels of Udskoi Ostrog in Amurland, towards the eastern side of the Lena River, though he wrote that it also occurred around the Yenisei, and that it occasionally crossed into China. The British naturalist Brian Hodgson gave the dhole the binomial name Canis primaevus, assuming that it is the progenitor of the domestic dog. Hodgson later took note of the dhole's physical distinctiveness from the genus Canis and assigned it to a new genus Cuon.
The first study on the origins of the species was conducted by paleontologist Erich Thenius, who concluded that the dhole was a post-Pleistocene descendant of a golden jackal-like ancestor. The earliest known member of the genus Cuon is the Chinese C. majori of the Villafranchian period. It resembled Canis in its physical form more than the modern species, which has greatly reduced molars, whose cusps have developed into sharply trenchant points. By the Middle Pleistocene, C. majori had lost the last lower molar altogether. C. alpinus itself arose during the late Middle Pleistocene, by which point the transformation of the lower molar into a single cusped, slicing tooth had been completed. Late Middle Pleistocene dholes were virtually indistinguishable from their modern descendants, save for their greater size, which closely approached that of the grey wolf. The dhole became extinct in much of Europe during the late Würm period, though it may have survived up until the early Holocene in the Iberian Peninsula and at Riparo Fredian in northern Italy The fossil record indicates that the species also occurred in North America, with remains being found in Beringia and Mexico.
The dhole's distinctive morphology has been a source of much confusion in determining the species' systematic position among the canidae. George Simpson placed the dhole in the subfamily Simocyoninae alongside the African wild dog and the bush dog, on account of all three species' similar dentition. Subsequent authors, including Juliet Clutton-Brock, noted greater morphological similarities to canids of the genera Canis, Dusicyon and Alopex than to either Speothos or Lycaon, with any resemblance to the latter two being due to convergent evolution. Subsequent studies on the canid genome revealed that the dhole and African wild dog are closely related to members of the genus Canis, and that both are more closely related to grey wolves, coyotes, golden jackals and Ethiopian wolves than the more basal black-backed and side-striped jackals are. This closeness to Canis may have been confirmed in a menagerie in Madras where, according to zoologist Reginald Pocock, a dhole interbred with a golden jackal.
|Subspecies||Image||Trinomial authority||Common names||Description||Range||Synonyms|
|C. a. alpinus||Pallas, 1811||Indian wild dog|
|Large subspecies with bright red coat and narrow skull.||Far eastern Russia, Mongolia, China, Nepal, Indian subcontinent, Bhutan, Burma, Indochina and Java.||adustus (Pocock, 1941), antiquus (Matthew & Granger, 1923), clamitans (Heude, 1892), dukhunensis (Sykes, 1831), fumosus (Pocock, 1936), grayiformis (Hodgson, 1863), infuscus (Pocock, 1936), javanicus (Desmarest, 1820), laniger (Pocock, 1936), lepturus (Heude, 1892), primaevus (Hodgson, 1833), rutilans (Müller, 1839)|
|C. a. hesperius||Afanasjev and Zolotarev, 1935||Northern dhole|
Tien Shan dhole
|Smaller than C. a. alpinus, with wider skull and lighter coloured winter fur.||Altai, Tien Shan and possibly Pamir and Kashmir||jason (Pocock, 1936)|
|C. a. sumatrensis||Hardwicke, 1821||Sumatran dhole||Has short, coarse fur with no woolly underfur, and much black on the back.||Sumatra|
However, studies on dhole mtDNA and microsatellite genotype showed that there are no clear subspecific distinctions. Nevertheless, two major phylogeographic groupings were discovered in dholes of the Asian mainland, which likely diverged during a glaciation event. One population extends from South, Central, and North India (south of the Ganges) into Burma, and the other extends from India north of the Ganges into northeastern India, Burma, Thailand and the Malaysian Peninsula. The origin of dholes in Sumatra and Java is, as of 2005, unclear, as they show greater relatedness to dholes in India, Burma and China rather than with those in nearby Malaysia. In the absence of further data, the researchers involved in the study speculated that Javan and Sumatran dholes could have been introduced to the islands by humans.
In culture and literature
Three dhole-like animals are featured on the coping stone of the Bharhut stupa dating from 100 BC. They are shown waiting by a tree, with a woman or spirit trapped up it, a scene reminiscent of dholes treeing tigers. The animal's fearsome reputation in India is reflected by the number of pejorative names it possesses in Hindi, which variously translate as "red devil", "devil dog", "jungle devil", or "hound of Kali". According to zoologist and explorer Leopold von Schrenck, he had trouble obtaining dhole specimens during his exploration of Amurland, as the local Gilyaks greatly feared the species. This fear and superstition was not however shared by neighbouring Tungusic peoples. Von Schrenk speculated that this differing attitude towards dholes was due to the Tungusic people's more nomadic, hunter-gatherer lifestyle. Dhole-like animals are described in numerous old European texts, including the Ostrogoth sagas, where they are portrayed as hell hounds. The demon dogs accompanying Hellequin in Mediaeval French passion plays, as well as the ones inhabiting the legendary forest of Brocéliande, have been attributed to dholes. According to Charles Hamilton Smith, the dangerous wild canids mentioned by Scaliger as having lived in the forests of Montefalcone could have been based on dholes, as they were described as unlike wolves in habits, voice and appearance. The Montefalcone family's coat of arms had a pair of red dogs as supporters.
Dholes appear in Rudyard Kipling's Red Dog, where they are portrayed as aggressive and bloodthirsty animals which descend from the Deccan Plateau into the Seeonee Hills inhabited by Mowgli and his adopted wolf pack to cause carnage among the jungle's denizens. They are described as living in packs numbering hundreds of individuals, and that even Shere Khan and Hathi make way for them when they descend into the jungle. The dholes are despised by the wolves because of their destructiveness, their habit of not living in dens and the hair between their toes. With Mowgli and Kaa's help, the Seeonee wolf pack manages to wipe out the dholes by leading them through bee hives and torrential waters before finishing off the rest in battle. Japanese author Uchida Roan wrote 犬物語 (Inu monogatari; A dog's tale) in 1901 as a nationalistic critique of the declining popularity of indigenous dog breeds, which he asserted were descended from the dhole.
Brian Houghton Hodgson kept captured dholes in captivity, and found, with the exception of one animal, they remained shy and vicious even after 10 months. According to Richard Lydekker, adult dholes are nearly impossible to tame, though pups are docile and can even be allowed to play with domestic dog pups until they reach early adulthood. A dhole may have been presented as a gift to Ibbi-Sin as tribute.
Etymology and naming
The etymology of 'dhole' is unclear. The earliest possible written use of the word in English occurred in 1808 by soldier Thomas Williamson, who encountered the animal in Ramghur district. He stated that 'dhole' was a common local name for the species. In 1827, Charles Hamilton Smith claimed that it was derived from a language spoken in 'various parts of the East'. Two years later, Smith connected this word with Turkish: deli ‘mad, crazy’, and erroneously compared the Turkish word with Old Saxon: dol and Dutch: dol (cfr. also English: dull; German: toll), which are in fact from the Proto-Germanic *dwalaz ‘foolish, stupid’. Richard Lydekker wrote nearly 80 years later that the word was not used by the natives living within the species' range. The Merriam-Webster Dictionary theorises that it may have come from the Kannada: tōḷa (‘wolf’).
Local and indigenous names
|Linguistic group or area||Indigenous name|
豺蜀 (chai shu)
|Khmer||ចឰ បែ (chkai prey)|
|Lao||ໝາໃນ (ma nai)|
|Malayalam/Tamil||செந்நாய் (chen nai)|
|Russian||Красный волк (krasnyi volk)|
Ди́кая собака (dikaya sobaka)
|Telugu||అడవి కుక్క (resu kukka)|
రేసు కుక్క (adavi-kutta)
|Thai||หมาใน (maa nay)|
|Vietnamese||chó sói lửa|
- Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Durbin, L.S., Hedges, S., Duckworth, J.W., Tyson, M., Lyenga, A. & Venkataraman, A. (IUCN SSC Canid Specialist Group – Dhole Working Group) (2008). "Cuon alpinus". IUCN Red List of Threatened Species. Version 2014.3. International Union for Conservation of Nature.
- Lydekker, R. (1907). The game animals of India, Burma, Malaya, and Tibet. London: R. Ward Limited.
- Fox 1984
- Heptner, V. G. & Naumov, N. P. (1998). Mammals of the Soviet Union Vol.II Part 1a, SIRENIA AND CARNIVORA (Sea cows; Wolves and Bears), Science Publishers, Inc. USA., pp. 566–86, ISBN 1-886106-81-9
- Cohen, J. A. (1978). Cuon alpinus. Mammalian Species 100: 1–3.
- Perry, R. (1964). The World of the Tiger. London: Cassell.
- Lindblad-Toh, K.; Wade, C. M.; Mikkelsen, T. S.; Karlsson, E. K.; Jaffe, D. B.; Kamal, M.; Clamp, M.; Chang, J. L.; Kulbokas, E. J.; Zody, M. C.; Mauceli, E.; Xie, X.; Breen, M.; Wayne, R. K.; Ostrander, E. A.; Ponting, C. P.; Galibert, F.; Smith, D. R.; Dejong, P. J.; Kirkness, E.; Alvarez, P.; Biagi, T.; Brockman, W.; Butler, J.; Chin, C. W.; Cook, A.; Cuff, J.; Daly, M. J.; Decaprio, D. et al. (2005). "Genome sequence, comparative analysis and haplotype structure of the domestic dog". Nature 438 (7069): 803–819. Bibcode:2005Natur.438..803L. doi:10.1038/nature04338. PMID 16341006.
- Clutton-Brock, J., Corbet, G. G., and Hills, M. (1976). "A review of the family Canidae, with a classification by numerical methods". Bulletin of the British Museum of Natural History 29: 179–180.
- Zhang, H.; Chen, L. (2010). "The complete mitochondrial genome of dhole Cuon alpinus: Phylogenetic analysis and dating evolutionary divergence within canidae". Molecular Biology Reports 38 (3): 1651. doi:10.1007/s11033-010-0276-y. PMID 20859694.
- Fox 1984, p. 85
- Fox 1984, pp. 86–7
- Durbin, L.S., Venkataraman, A., Hedges, S. & Duckworth, W. (2004). Dhole Cuon alpinus (Pallas 1811), in Sillero-Zubiri, C., Hoffmann, M. & Macdonald, D.W. (eds.) Canids: Foxes, Wolverhampton Wanderers F.C., Jackals and Dogs: Status Survey and Conservation Action Plan. IUCN/SSC Canid Specialist Group. Gland, Switzerland and Cambridge, UK. x + pp. 210–219
- Karanth, K. U. and Sunquist, M. E. (1995). "Prey selection by tiger, leopard and dhole in tropical forests". Journal of Animal Ecology 64 (4): 439–450. doi:10.2307/5647. JSTOR 5647.
- Thenius, E. (1955). "Zur Abstammung der Rotwölfe (Gattung Cuon Hodgson) [On the origins of the dholes (Genus Cuon Hodgson)]". Österreichische Zoologische Zeitschrift (in German) 5: 377–388.
- Fox 1984, pp. 61–2
- Fox 1984, pp. 41
- Fox 1984, p. 93
- Fox 1984, p. 95
- Fox 1984, p. 97
- Harris, R. B. (2006). "Attempted predation on blue sheep Pseudois nayaur by dholes Cuon alpinus". Journal of the Bombay Natural History Society 103: 95–97.
- Thapa, K., Kelly, M. J., Karki, J. B. and Subedi, N. (2013). "First camera trap record of pack hunting dholes in Chitwan National Park, Nepal". Canid Biology & Conservation 16 (2): 4–7.
- Khatiwada, A. P.; Awasthi, K. D.; Gautam, N. P.; Jnawali, S. R.; Subedi, N.; Aryal, A. (2011). "The Pack Hunter (Dhole): Received Little Scientific Attention". The Initiation 4. doi:10.3126/init.v4i0.5531.
- Wangchuk, T. (2004). "Predator-prey dynamics: the role of predators in the control of problem species". J. Bhutan Studies 10: 68–89.
- Thinley, P., Kamler, J. F., Wang, S. W., Lham, K., Stenkewitz, U. (2011). "Seasonal diet of dholes (Cuon alpinus) in northwestern Bhutan". Mammalian Biology 76: 518–520. doi:10.1016/j.mambio.2011.02.003.
- Jenks, K. E., Songsasen, N. and P. Leimgruber (2012). "Camera trap records of dholes in Khao Ang Rue Nai Wildlife Sanctuary, Thailand". Canid News. online: 1–5.
- Fox 1984, pp. 81–2
- Walker, E. P., Nowak, R. M., Warnick, F. (1983). Walker's Mammals of the World. 4th ed. Baltimore: Johns Hopkins University Press.
- Fox 1984, p. 92
- Fox 1984, pp. 43–49
- Fox 1984, p. 80
- Fox 1984, p. 79
- Fox 1984, pp. 100–1
- Fox 1984, p. 50
- Fox 1984, p. 73
- Fox 1984, p. 67
- Grassman, L. I., Jr., M. E. Tewes, N. J. Silvy, and K. Kreetiyutanont (2005). "Spatial ecology and diet of the dhole Cuon alpinus (Canidae, Carnivora) in north central Thailand". Mammalia 69 (1): 11–20. doi:10.1515/mamm.2005.002.
- Fox 1984, p. 63
- Fox 1984, p. 70
- Fox 1984, p. 51
- Pocock, R. I. (1941), Fauna of British India: Mammals volume 2, Taylor & Francis, pp. 146–63
- Fox 1984, pp. 58–60
- Mivart, G. (1890), Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ, London : R.H. Porter : Dulau, pp. 177–88
- Fox 1984, p. 71
- Johnsingh, A.J.T., Yonten, Deki & Wangchuck, Sangay (2007). "Livestock-Dhole Conflict in Western Bhutan". J. Bombay Nat. Hist. Soc. 104 (2): 201.
- Venkataraman, A. (1995). "Do dholes (Cuon alpinus) live in packs in response to competition with or predation by large cats?". Current Science 11: 934–936.
- Finn, F. (1929). Sterndale's Mammalia of India. London: Thacker, Spink & Co.
- Shrestha, T. J. (1997). Mammals of Nepal: (with reference to those of India, Bangladesh, Bhutan and Pakistan). Kathmandu: Bimala Shrestha. ISBN 0-9524390-6-9.
- Nair, M. V. & Panda, S. K. (2013). Just Friends. Sanctuary Asia, Vol. XXXIII, No. 3
- Humphrey, S. R., Bain, J. R. (1990). Endangered Animals of Thailand. Gainesville: Sandhill Crane Press. ISBN 1-877743-07-0.
- Fox 1984, p. 109
- Zoo to have conservation breeding centre for ‘dhole’, The Hindu (August 18, 2014)
- Pallas, P. S. (1811), Zoographia Rosso-Asiatica : sistens omnium animalium in extenso Imperio Rossico, et adjacentibus maribus observatorum recensionem, domicilia, mores et descriptiones, anatomen atque icones plurimorum, Petropoli : In officina Caes. Acadamiae Scientiarum Impress. MDCCCXI, pp. 34–5
- Hodgson, B. H. (1833). "Description and Characters of the Wild Dog of the Himalaya (Canis primævus)". Asiatic Researches. XVIII, Part 2: 221–237.
- Hodgson, B. H. (1842). "European Notices of Indian Canines, with Further Illustrations of the New Genus Cuon vel Chrysæus". Calcutta Journal of Natural History II: 205–209.
- (German) Schrenk, L. v. (1859), Reisen und forschungen im Amur-lande in den jahren 1854–1856, St. Petersburg : K. Akademie der Wissenschaften, pp. 48–50
- Kurtén, Björn (1968), Pleistocene mammals of Europe, Weidenfeld and Nicolson, pp. 111–114
- Ripoll, M. P. R.; Morales Pérez, J. V.; Sanchis Serra, A.; Aura Tortosa, J. E.; Montañana, I. S. N. (2010). "Presence of the genus Cuon in upper Pleistocene and initial Holocene sites of the Iberian Peninsula: New remains identified in archaeological contexts of the Mediterranean region". Journal of Archaeological Science 37 (3): 437. doi:10.1016/j.jas.2009.10.008.
- Petrucci, Mauro; Romiti, Serena; Sardella, Raffaele (2012). "The Middle-Late Pleistocene Cuon Hodgson, 1838 (Carnivora, Canidae) from Italy". Bollettino della Società Paleontologica Italiana 51 (2): 146.
- Kurtén, Björn (1980), Pleistocene mammals of North America, Columbia University Press, ISBN 0231516967, p. 172
- Simpson, G.G. (1945), The principles of classification and a classification of mammals, Bulletin of the American Museum of Natural History, 85:1–350
- Ellerman, J.R. & Morrison-Scott, T.C.S. (1966). Checklist of Palaearctic and Indian mammals, British Museum (Natural History), London, UK.
- Iyengar, A., Babu, V. N., Hedges, S., Venkataraman, A. B., Maclean, N. and P. A. Morin (2005). "Phylogeography, genetic structure, and diversity in the dhole (Cuon alpinus)". Molecular Ecology 14 (8): 2281–2297. doi:10.1111/j.1365-294X.2005.02582.x. PMID 15969714.
- van der Geer, A. A. E. (2008), Animals in stone: Indian mammals sculptured through time, BRILL, p. 188, ISBN 90-04-16819-2
- Smith, C. H. & Jardine, W. (1839). The natural history of dogs : canidae or genus canis of authors ; including also the genera hyaena and proteles, Vol. I. Edinburgh: W.H. Lizars.
- Skabelund, A. H. (2011). Empire of Dogs: Canines, Japan, and the Making of the Modern Imperial World. Cornell University Press, p. 85, ISBN 0801463246
- McIntosh, J. (2008). The ancient Indus Valley: new perspectives, p. 130, ABC-CLIO, ISBN 1-57607-907-4
- Williamson, T. (1808). Oriental field sports: being a complete, detailed, and accurate description of the wild sports of the East. Volume II. London: Orme.
- Smith, C. H. (1827). The class Mammalia. London: Geo. B. Whittaker.
- Orel, V. (2003), A Handbook of Germanic Etymology, Leiden, Boston: Brill, p. 81, ISBN 90-04-12875-1
- dhole. Merriam-Webster Dictionary.
- Andrew T. Smith, Yan Xie, Robert S. Hoffmann, Darrin Lunde, John MacKinnon, Don E. Wilson, W. Chris Wozencraft (2010). A Guide to the Mammals of China. Princeton University Press. pp. 418–19. ISBN 1400834112
- Fox, M. W. (1984). The Whistling Hunters: Field Studies of the Asiatic Wild Dog (Cuon Alpinus). Albany: State University of New York Press. ISBN 0-9524390-6-9.
EOL content is automatically assembled from many different content providers. As a result, from time to time you may find pages on EOL that are confusing.
To request an improvement, please leave a comment on the page. Thank you!