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Overview

Brief Summary

Biology

Dholes are highly social animals and they live and hunt in packs that closely resemble those of the African wild dog (Lycaon pictus) (4). These packs seem to consist of more males than females (6) and usually contain around 5 to 12 members, although groups of up to 40 have been observed on occasion (5). There is a strict hierarchy within the pack and the group will defend a territory that can be as large as 84 square kilometres depending on the availability of food (6); territories are marked by latrine sites at trail intersections (4). Usually only the dominant female will breed (5), giving birth to a litter of three to four young, or occasionally ten, after a two month gestation period (2). The mating season occurs from September to February (2). Pups are born in a den, which is usually the abandoned burrow of another animal, and all members of the pack help to care for the mother and her litter (6). Individuals feed the pups by regurgitating food for them, and will help to guard the den; when the pups are old enough to accompany the adults on hunting trips they are allowed to eat first at the kill (4). Cooperating in a pack to hunt prey, dholes are capable of killing animals over ten times their own body weight in size (5). Their diet is almost wholly carnivorous, predominantly made up of medium-sized ungulates (2) such as spotted deer (Axis axis), sambar deer (Cervus unicolor) and wild sheep (6). Hunting in thick forest, dholes rely on scent to locate prey, occasionally jumping high into the air to get their bearings (4). Pack members either move forward in a line or stand guard on the edge of dense cover whilst other members flush out the prey (4). Dholes are capable swimmers and sometimes drive their prey into water (5). Like the African wild dog, these animals have acquired a vicious reputation due to the speed with which they eat, and their method of disembowelling prey before it is fully dead (4). Attacks on humans are, however, extremely rare (4).
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Description

Dholes or Asian wild dogs are pack-living canids, although they are unique amongst this family in having a thickset muzzle and one less molar tooth on each side of the lower jaw (4). The bushy coat is usually a rusty red colour with white on the belly, chest and paws (5). Different subspecies exist and those in the northern parts of the dhole's range have lighter and longer hair than their southern relatives (6). The bushy tail is black and the pups are also born a sooty black colour before acquiring their adult coat at around three months of age (4). The large rounded ears are filled with white hair (6) and the eyes are amber (5). Males tend to be significantly larger in size than females (6). Dholes have a wide range of vocalisations including an extremely distinctive whistle that is used to reassemble pack members in the thick forest of their habitat (5).
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Comprehensive Description

Summary

"The Dhole or the Indian/asiatic wild dog is the only extant member of the genus Cuon. It is a highly social animal, living and hunting in packs (clans). They are less territorial than wolves ."
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Distribution

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 )

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Range Description

In Central and eastern Asia, there have been no confirmed, recent reports of dholes from Russia, Mongolia, Kazakhstan, Kyrgyzstan (where they were found formerly in the Tian-Shan area) or Tajikistan (where they were found formerly in the eastern Pamir area) (A. Poyarkov and N. Ovsyanikov in litt. D. Miquelle pers. comm.). There is a recent report of a dhole that was captured in Jiangxi district, south China (C. Bellamy pers. comm.). Dholes were once present in parts of western China in the Tian-Shan Range, but the species' current status in this area is unclear; they do at least still persist, perhaps in low numbers, in parts of the Qilian Shan in north-western Gansu Province (Harris 2006). The species is still found in Tibet today, particularly in areas bordering the Ladakh region of India (R. Wangchuk pers. comm.), and the Tibet Forestry Bureau has reported that dholes are still "common" in parts of southeast Tibet (S. Chan, in litt.). Dholes occurred in northern Korea (Won Chang Man and Smith 1999) and a few small populations may still exist. There have been no records from Pakistan, but the species occurred on the alpine steppes of Ladakh, Kashmir, and India (Johnsingh 1985) that extend into the region termed Pakistan-occupied Kashmir by India.

Dholes are still found throughout much of India south of the river Ganges, and especially in the Central Indian Highlands and the Western and Eastern Ghats of the southern states. They are also found throughout north-east India, in the states of Arunachal Pradesh, Assam, Meghalaya, and West Bengal (A. Venkataraman, A.J.T. Johnsingh and L. Durbin pers. comm.). In the Himalaya and north-western India, the status of dholes seems more precarious with a much more fragmented distribution. Dholes reportedly still occur in the Ladakh area of Kashmir, which is contiguous with the Tibetan highlands in China (R. Wangchuk pers. comm.).

The species formerly was recorded in the Terai region of the Indo-gangetic plain, including the Royal Chitawan National Park in Nepal, but there have been few recent reports. There is an unconfirmed report of dholes in Dhorpatan Hunting Reserve in the late 1990s (R.C. Kandel pers. comm.).

In Bhutan, there have been recent press reports that dholes have recovered from a government-initiated mass poisoning campaign in the 1970s and there have apparently been numerous recent incidents of dholes killing livestock in the lower Kheng region. Two recent, independent, eye-witness reports identify dholes in six protected areas in Bhutan (S. Wangchuk pers. comm., T. Wangchuk pers. comm.). In some regions, dhole predation on wild boar (Sus scrofa) may be viewed in a positive light by local people (T. Wangchuk pers. comm.).

In Bangladesh, dholes were thought to occur in the forested tracts of the Chittagong and Sylhet Districts (Johnsingh 1985). It is not certain whether any remain in Bangladesh.

In Myanmar, dholes were recorded by camera trapping at 11 of 15 survey areas scattered across the country, only four of which were protected. Dholes and/or leopards have apparently replaced tigers as the top predator in these areas (Myanmar Forest Department 2003).

In Indochina, dholes probably ranged over all or almost all of Lao PDR, Cambodia, Viet Nam and Thailand, although reliable site-specific information is scarce. Present distribution is highly fragmented and large parts, particularly of Viet Nam and Thailand, are without any regular occurrence of dholes, although they persist in a number of protected areas (Duckworth et al. 1999, Waltson 2001, M. Baltzer and R. Shore in litt., A. Lynam pers. comm.).

The species' historical range probably included all or most of the Malaysian peninsula and the Indonesian islands of Sumatra and Java, but reliable information is scarce. Current distribution is poorly known but is thought to be highly fragmented. On the Malaysian peninsula, dholes are known to occur in four sites in northern and central areas of the peninsula (from recent camera-trap surveys; J.B. Abdul pers. comm.). On Java, dholes appear to be most common in the protected areas at the eastern and western ends of the island. On Sumatra, very little is known, but dholes are known to occur in major protected areas in the southern, central, and northern parts of the island (e.g., from camera trapping; D. Martyr pers. comm.).

There is no reliable evidence of the presence of Dhole in Turkey (Kryštufek and Vohralík 2001; Can 2004).
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Historic Range:
C.I.S., Korea, China, India, Southeast Asia

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Range

Dholes previously ranged throughout the Indian subcontinent, north into Korea, China and eastern Russia and south through Malaysia and Indonesia reaching as far as Java (2). Today, information on dhole numbers is lacking but the range appears to be greatly reduced and remaining populations are isolated in fragments of former habitat (5). There are 11 subspecies of dhole and these vary in range with the most common being Cuon alpinus dukhunensis found in central and southern India (5) (6).
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Physical Description

Morphology

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

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"General colour bright rusty-red or rufous fawn-colour, paler beneath ; ears erect, rather large, somewhat rounded at the tip ; tail moderately brushed, reaching to the heels, usually tipped blackish; limbs strong ; body lengthened."
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Size

"Length, head and body, 32 to 36 inches ; tail about 16 inches ; height 17 to 20 inches."
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Ecology

Habitat

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

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Habitat and Ecology

Habitat and Ecology
The dhole is found in a wide variety of vegetation types, including: primary, secondary and degraded forms of tropical dry and moist deciduous forest; evergreen and semi-evergreen forests; dry thorn forests; grassland–scrub–forest mosaics; and alpine steppe (above 3,000 m). They are not recorded from desert regions.
In India, tropical dry and moist deciduous forest may represent optimal habitats, based on the regions thought to hold the largest dhole populations. Ungulate biomass, particularly that of cervid species, is highest in these vegetation types when compared to others in the same region (A. Venkataraman and V. Narendra Babu, unpubl.). In India, tropical dry and moist deciduous forests are subject to seasonal monsoon climates.

Important factors that may influence habitat selection include the availability of medium to large ungulate prey species, water, the presence of other large carnivore species, human population levels and suitability of breeding sites (proximity to water, presence of suitable boulder structures and sufficient prey).

Systems
  • Terrestrial
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Dholes are found in forested areas throughout their range from dense montane forest in Thailand to alpine areas in Russia, and thick scrub jungle in India (2). In general, factors such as prey and water availability, den sites and relatively open forest areas with grassy meadows (usually having high prey densities) are required to support dholes (2).
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Trophic Strategy

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

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Associations

Known prey organisms

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Life History and Behavior

Behavior

Perception Channels: tactile ; chemical

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Behaviour

"The wild dog preys both by night and day, but chiefly by day. Six, eight, or ten unite to hunt down their victim, maintaining the chase by their powers of smell rather than by the eye. Theyusually overcome their quarry by dint of force and perseverance, thqugh they sometimes effect their object by mixing stratagem with direct violence."
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Life Expectancy

Average lifespan

Status: captivity:
16.0 years.

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Lifespan, longevity, and ageing

Maximum longevity: 16.1 years (captivity)
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Reproduction

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

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"The bitch has twelve to fourteen teats, and has at least six whelps at a birth. They breed from January to March. They breed in holes under rocks, several females apparently breeding together."
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Molecular Biology and Genetics

Molecular Biology

Barcode data: Cuon alpinus

The following is a representative barcode sequence, the centroid of all available sequences for this species.


No available public DNA sequences.

Download FASTA File
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Statistics of barcoding coverage: Cuon alpinus

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 2
Specimens with Barcodes: 2
Species With Barcodes: 1
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Conservation

Conservation Status

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

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IUCN Red List Assessment


Red List Category
EN
Endangered

Red List Criteria
C2a(i)

Version
3.1

Year Assessed
2008

Assessor/s
Durbin, L.S., Hedges, S., Duckworth, J.W., Tyson, M., Lyenga, A. & Venkataraman, A. (IUCN SSC Canid Specialist Group - Dhole Working Group)

Reviewer/s
Sillero-Zubiri, C. & Hoffmann, M. (Canid Red List Authority)

Contributor/s

Justification
It is estimated that fewer than 2,500 mature individuals remain in the wild and the declining population trend is expected to continue. Main threats to the species include ongoing habitat loss, depletion of prey base, interspecific competition, persecution and possibly disease transfer from domestic and feral dogs.

History
  • 2004
    Endangered
  • 1996
    Vulnerable
  • 1994
    Vulnerable
    (Groombridge 1994)
  • 1990
    Vulnerable
    (IUCN 1990)
  • 1988
    Vulnerable
    (IUCN Conservation Monitoring Centre 1988)
  • 1986
    Vulnerable
    (IUCN Conservation Monitoring Centre 1986)
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Current Listing Status Summary

Status: Endangered
Date Listed: 06/02/1970
Lead Region: Foreign (Region 10) 
Where Listed: Entire


Population detail:

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

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Red List Category & Criteria: Endangered C2a(i) ver 3.1 Year Published: 2008
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Status

Classified as Endangered (EN) on the IUCN Red List (1), and listed on Appendix II of CITES (3).
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Population

Population Trend
Decreasing
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Threats

Major Threats
Depletion of prey base: Across almost all of Cambodia, Lao PDR, and Viet Nam, as well as within protected areas, ungulates occur at levels well below natural. All species of ungulate except muntjacs (Muntiacus spp.), pigs (Sus spp.) and in some areas southern serow (Naemorhedus sumatraensis) are ecologically or fully extinct across extensive parts of the region. Only a few of the largest wildernesses support nearly intact species assemblages and even in these, the larger species (Bos spp., Cervus spp., hog deer Axis porcinus) are very rare. This situation will likely hinder any possibility of recovery by the region's dhole populations, even if the other issues could be addressed. While not as depressed as in Indochina, prey levels in Indonesia also exist at levels much below carrying capacity (because of illegal hunting and habitat degradation). In protected areas in southern and central India, where dhole numbers are stable, prey densities are high. In north-east India, prey densities are very low in protected areas with dholes.

Habitat loss and transformation: Currently, extensive areas of natural or semi-natural vegetation remain in Lao PDR and Cambodia, some areas encompassing many hundreds of square kilometres of potential dhole habitat. However, habitat conversion and fragmentation are proceeding apace. In Viet Nam, very few natural areas of over 50 km² remain. Habitat loss and fragmentation is a major threat to protected areas in Indonesia, particularly those on Sumatra. Habitat loss and degradation are also serious threats to dholes in South Asia and the disappearance of dholes from many of the forested tracts in India has been attributed in large part to loss of habitat.

Persecution: This certainly occurs in Indochina, although it is unclear how often. In Indonesia, too, it is a threat but again its significance is unknown. In India, such persecution can play a serious role in limiting local populations. Dholes living outside or on the edge of core protected areas are particularly vulnerable to human kleptoparasitism, snaring (non-selective) and direct persecution. For example, during a radio-tracking study in 2000, in the buffer zone of Kanha Tiger Reserve, central India, at least 16 out of 24 dholes in one pack died from a sudden strychnine poisoning (L. Durbin pers. obs.). In southern India, such persecution is moderate to low and often occurs indirectly when cattle graziers and others inadvertently go close to dhole dens and disturb adults and pups, disrupting breeding and rearing (A. Venkataraman pers. obs.). "By-catch" in snares and other traps is probably a significant threat to dholes across Indochina at least.

Competition with other species: Apparently, free-living dogs have been seen and/or camera trapped in many parts of Indochina, but there is no evidence for existence of large populations. Undoubtedly, the main competitor for prey species in Indochina is people. There is no evidence that feral dogs are significant competitors with dholes in Indonesia. In many parts of their range, dholes are sympatric with tigers and leopards and so the potential for significant interspecific competition for prey exists, especially if the prey populations are reduced as a result of hunting by people.

Disease and pathogens: Particularly those transmitted by feral and/or domestic dogs (e.g., mange, canine distemper, parvovirus and rabies). The significance of disease is unclear in Indochina, but diseases are a significant threat in South Asia and probably in parts of Indonesia.

There is no widespread exploitation for fur or other purposes, though medicinal use should be investigated in China.
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"Loss of habitat, depletion of prey base, persecution , disease from feral or domestic dogs."
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Dhole numbers have been reduced as their habitat is being destroyed throughout much of the Asian continent; the human population explosion has led to the clearance of vast tracts of forest for timber and to make way for agriculture and development (5). Historically, hunters viewed dholes as competition and thus persecuted them; bounties were also offered for their pelts (4). Today, habitat loss and the elimination of prey species pose the greatest threats to the survival of the dhole (4). Diseases such as distemper and rabies, possibly spread by domestic dogs, are important threats to the Indian subspecies C. a. primaevus (2).
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Management

Conservation Actions

Conservation Actions
Included in CITES – Appendix II (2003).

In Cambodia, the current wildlife decrees give the dhole protection from all hunting. A new forestry law is under preparation, and a proposal to list the species as a fully protected species is under discussion. In India, the dhole is protected under Schedule 2 of the Wildlife Act of 1972 (permission is required to kill any individual unless in self defence or if an individual is a man killer). The creation of Project Tiger Reserves in India has provided some protection for populations of the dukhunensis subspecies (A.J.T. Johnsingh pers. comm., L. Durbin, pers. obs.). In the Russian Federation, dholes received the status of "protected animal" in 1974 (A. Poyarkov and N. Ovsyanikov in litt.); however, the poisoning of grey wolves may inadvertently affect any remnant dhole populations (V. Puzanskii pers. comm.). In Viet Nam, the dhole is protected by Decree 18/HDBT (17/01/1992) and the amendment Decree 48/2002/ND-DP (22/04/2002) under category IIB, which limits extraction and utilization. However, the levels of extraction or utilization are not quantified (B. Long in litt. 2003). Dholes are listed as a category II protected species under the Chinese wildlife protection act of 1988.

The species occurs in protected areas throughout its range.

No conservation measures specifically focused on dholes have been reported for most range states. In India, Project Tiger could potentially maintain dhole prey bases in areas where tigers and dholes coexist. There do not appear to be any specific measures for dhole conservation in Indochina, although the declaration of relatively large protected area networks in Cambodia, Lao PDR, and Viet Nam will, when these areas become functional realities on the ground, form a suitable conservation system for the species in at least Cambodia and Lao PDR.

There are at least 110 dholes in captivity, and the sex ratio is approximately even. Except for some captive populations in India heterozygosity appears to be good, but there is little chance of breeding the putative subspecies as animals from diverse geographical origins have been widely interbred (M. Boeer pers. comm.).
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Conservation

Dholes are protected throughout most of their range, in India they are protected under Schedule I of the Wildlife Act of 1972 and hunting has also been prohibited in Russia since 1971 (2). In India and Nepal, dholes are protected within many tiger reserves and this has helped to keep their stronghold in southern India (4). More data is urgently needed on dhole distribution and numbers and the Dhole Conservation Programme is working to achieve this and to develop a Dhole Action Plan to safeguard the future of this remarkable canid (5).
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Relevance to Humans and Ecosystems

Benefits

Although this occurs on rare occasions, dholes can attack livestock at the cost of the owner.

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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.

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Wikipedia

Dhole

This article is about the species of wild dog. For H. P. Lovecraft's fictional monster, see Dhole (Cthulhu Mythos). For the town in France, see Cuon, Maine-et-Loire.

The dhole (Cuon alpinus) is a canid native to Central and Southeast Asia. Although genetically close to dogs of the genus Canis,[3](Fig. 10) it is morphologically distinguished by its distinctive skull and teeth.[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.[5]

The dhole is a highly social animal, living in large clans without rigid dominance hierarchies.[6] It is a diurnal pack hunter which preferentially targets medium and large sized ungulates.[7] Other than humans, its only natural predators are tigers and leopards, with which it avoids competition by targeting different prey animals.[8]

It is listed as Endangered by the IUCN, on account of its decreasing population, which is now 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.[2]

Etymology and naming[edit]

The etymology of 'dhole' is unclear. The earliest possible written use of the word occurred in 1808 by soldier Thomas Williamson, who encountered the animal in West Bengal's Ramghur district. He stated that 'dhole' was a common local name for the species.[9] In 1827, Charles Hamilton Smith claimed that it was derived from a language spoken in 'various parts of the East'.[10] 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),[11] which are in fact from the Proto-Germanic *dwalaz ‘foolish, stupid’.[12] Richard Lydekker wrote nearly 80 years later that the word was not used by the natives living within the species' range.[13] The Merriam-Webster Dictionary theorises that it may have come from the Kannada: tōḷa (‘wolf’).[14]

Other English names for the species include wild dog,[13] whistling dog, chennai,[15] red wolf[16] (not to be confused with Canis [lupus] rufus), red dog[17] and mountain wolf.[18]

Local and indigenous names[edit]

Discovery, taxonomy and evolution[edit]

Illustration (1859) by Leopold von Schrenck, one of the first authentic depictions of the species, based on a single skin purchased in the village of Dshare on Amur.[19]

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, 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.[20]

Skeletal remains of a European dhole dating back to upper Würm period from Cova Negra de Játiva, Valencia, Spain.

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.[21] 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 tubercular teeth, 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 tubercular tooth 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,[22] though it may have survived in the Iberian Peninsula up until the early Holocene.[23] The fossil record indicates that the species also occurred in North America, with remains being found in Beringia and Mexico.[24]

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.[25] 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.[4] 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.[3] 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.[26]





Side-striped jackal



Black-backed jackal








Golden jackal





Dog



Grey wolf




Coyote





Ethiopian wolf




Dhole





African wild dog






Subspecies[edit]

Historically, up to ten subspecies of dhole have been recognised.[27] As of 2005,[28] only three subspecies are recognised by MSW3.

SubspeciesImageTrinomial authorityCommon namesDescriptionRangeSynonyms
C. a. alpinus

(Nominate subspecies)

Keulemans Cuon alpinus javanicus.pngPallas, 1811Southern dhole[29]
Ussuri dhole[16]
Large subspecies with bright red coat and narrow skull.[16]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), sumatrensis (Hodgson, 1863)
C. a. hesperiusKeulmans Cuon alpinus alpinus.pngAfanasjev and Zolotarev, 1935Northern dhole[29]
Tien Shan dhole[16]
Smaller than C. a. alpinus, with wider skull and lighter coloured winter fur.[16]Altai, Tien Shan and possibly Pamir and Kashmirjason (Pocock, 1936)
C. a. sumatrensisKeulmans Cuon alpinus dukhunensis.pngHardwicke, 1821Has short, coarse fur with no woolly underfur, and much black on the back.[29]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.[30]

Physical description[edit]

In appearance, the dhole has been variously described as combining the physical characteristics of the grey wolf and red fox,[16] and as being "cat-like" on account of its long backbone and slender limbs.[21] It has a wide and massive skull with a well-developed sagittal crest,[16] and its masseter muscles are highly developed compared to other canid species, giving the face an almost hyena-like appearance.[31] The rostrum is shorter than that of domestic dogs and most other canids.[17] The species has six rather than seven lower molars.[32] 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 two to four, as is usual in canids,[16] an adaptation thought to improve shearing ability, thus allowing it to compete more successfully with kleptoparasites.[7] Adults may weigh over 40 lbs, with females usually weighing 10 lbs 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.[32] It has 6-7 teats, sometimes eight.[16]

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.[16] 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.[17]

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.[33] Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries.[34] In contrast to wolves, dholes do not howl or bark.[16] 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.[35]

Behaviour[edit]

Social and territorial behaviours[edit]

Dholes playing, Pench Tiger Reserve.

Dholes are more social than grey wolves,[16] 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.[6] 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.[36] 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.[37] Intragroup fighting is rarely observed.[38] Dholes are far less territorial than wolves, with pups from one clan often joining another without trouble once they mature sexually.[39] Clans typically number 5-12 individuals in India, though clans of 40 have been reported. In Thailand, clans rarely exceed three individuals.[17] 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.[35]

Reproduction and development[edit]

In India, the mating season occurs between mid-October and January, while captive dholes in the Moscow Zoo breed mostly in February.[17] Unlike wolf packs, dhole clans may contain more than one breeding female.[37] More than one female dhole may den and rear their litters together in the same den.[38] 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.[40] The gestation period lasts 60–63 days, with litter sizes averaging 4-6 pups.[17] 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.[41] Maximum longevity in captivity is 15–16 years.[38]

Hunting and feeding behaviours[edit]

Dholes attacking a sambar, Bandipur National Park

Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and homo- and heterosexual mounting.[42] 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.[43] Though not as fast as jackals and foxes, they can chase their prey for many hours.[16] 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.[44] When chasing fleet-footed prey, they run at a pace of 30 mph.[16] Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.[45]

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.[31] They occasionally blind their prey by attacking the eyes.[46] 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.[13] 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.[47] 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.[48] 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.[37] They are generally tolerant of scavengers at their kills.[49] Both mother and young are provided with regurgitated food by other pack members.[38]

Ecology[edit]

Habitat[edit]

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.[50] Like African wild dogs and dingoes, dholes will avoid killing prey close to their dens.[41]

Diet[edit]

Dholes feeding on a chital, Bandipur National Park

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.[17][26][51] 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.[18] In Kashmir, they may hunt markhor,[26] and thamin in Burma.[17] Javan rusas are hunted in Java.[7] 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.[16] Like African wild dogs, but unlike wolves, dholes are not known to attack people.[16][26] 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.[29] In summertime in the Tien Shan Mountains, dholes eat large quantities of mountain rhubarb.[16] Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves.[52] 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.[53]

Enemies and competitors[edit]

Dhole killed and cached in a tree by a leopard, India.

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[8] 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).[8] 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.[8] 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.[18][26] 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.[18] Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs.[26] Because leopards are smaller than tigers, and are more likely to hunt dholes, dhole packs tend to react more aggressively toward them than they do with tigers.[54] There are numerous records of leopards being treed by dholes.[38] Dholes sometimes drive tiger, leopards, and bears (see below) from their kills.[38] 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.[55]

Dhole packs occasionally attack Asiatic black bears and sloth bears. When attacking bears, dholes will attempt to prevent them from seeking refuge in caves, and lacerate their hindquarters.[26]

Though usually antagonistic toward wolves,[16] they may hunt and feed alongside one another.[56] They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion.[57]

Diseases and parasites[edit]

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.[7]

Range and conservation[edit]

There are currently no confirmed recent reports on the animal's presence 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. It still occurs in Tibet, and may still inhabit North Korea. Although it has not been recorded in Pakistan, the species did once occur in the alpine steppes extending into Kashmir.[2]

It still occurs 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, it inhabits Arunachal Pradesh, Assam, Meghalaya, and West Bengal. The situation of dholes in Himalaya and north-west India is precarious, with the species occurring in fragmented populations. It may still occur in Kashmir's Ladakh area.[2]

Dholes once occurred in the Indo-Gangetic Plain's Terai region, including in Nepal's Chitawan National Park, though recent reports are scant.[2]

Dholes in Bhutan have apparently recovered from a mass poisoning campaign during the 1970s, and have begun to frequently attack livestock in the lower Kheng region.[2]

It is unknown whether the species still lives in Bangladesh, where it once inhabited the forested areas of the Chittagong and Sylhet District.[2]

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.[2]

Its range is highly fragmented in Indochina, though it still persists in protected areas in Vietnam and Thailand.[2]

Although the species probably once occurred in most of the Malaysian Peninsula, Sumatra and Java, its populations there are highly fragmented.[2]

Relationships with humans[edit]

Hunting and persecution[edit]

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.[38] 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.[7] 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.[58] In Indochina, dholes suffer heavily from nonselective hunting techniques such as snaring.[7]

The fur trade does not pose a significant threat to dholes.[7] The people of India do not eat dhole flesh, and their fur is not considered overly valuable.[29] 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.[16]

In culture and literature[edit]

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.[59] 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.[19] 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. The dangerous wild canids mentioned by Scaliger as having lived in the forests of Montefalcone could have been 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.[11]

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 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.[60]

Tameability[edit]

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.[29] 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.[13] A dhole may have been presented as a gift to Ibbi-Sin as tribute.[61]

See also[edit]

References[edit]

  1. ^ 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. 
  2. ^ a b c d e f g h i j 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. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 22 March 2009. Database entry includes justification for why this species is endangered
  3. ^ a b Lindblad-Toh, K.; Wade, CM; Mikkelsen, TS; Karlsson, EK; Jaffe, DB; Kamal, M; Clamp, M; Chang, JL et al. (2005). "Genome sequence, comparative analysis and haplotype structure of the domestic dog". Nature 438 (7069): 803–819. doi:10.1038/nature04338. PMID 16341006. 
  4. ^ a b Clutton-Brock, J., Corbet, G.G., and Hills, M. (1976). "A review of the family Canidae, with a classification by numerical methods." Bull. Brit. Mus. Nat. Hist. 29, 179–180.
  5. ^ 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), 1651-1660.
  6. ^ a b Fox 1984, p. 85
  7. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw ax ay az ba bb bc bd be 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, Wolves, Jackals and Dogs: Status Survey and Conservation Action Plan, IUCN/SSC Canid Specialist Group. Gland, Switzerland and Cambridge, UK. x + pp. 210-219
  8. ^ a b c d 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. 
  9. ^ Williamson, T. (1808), Oriental field sports: being a complete, detailed, and accurate description of the wild sports of the East, Vol. II, Orme, pp. 1-12
  10. ^ Smith, C. H. (1827), The class Mammalia (1827), London : Printed for Geo. B. Whittaker, p. 326
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  12. ^ Orel, Vladimir (2003), A Handbook of Germanic Etymology, Leiden, Boston: Brill, p. 81, ISBN 90-04-12875-1 
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  14. ^ dhole. Merriam-Webster Dictionary.
  15. ^ Fox 1984
  16. ^ a b c d e f g h i j k l m n o p q r s 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
  17. ^ a b c d e f g h Cohen, J. A. (1978), Cuon alpinus. Mammalian Species 100: 1-3.
  18. ^ a b c d Perry, Richard (1965), The World of the Tiger, Cassell & Company ltd., pp. 149-162, ASIN B0007DU2IU
  19. ^ a b c (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
  20. ^ 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
  21. ^ a b Thenius, E. (1954), On the origins of the dholes, Osterreich Zoologie Zietshcrift, 5, 377–388
  22. ^ Kurtén, Björn (1968), Pleistocene mammals of Europe, Weidenfeld and Nicolson, pp. 111-114
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Bibliography[edit]

  • Fox, M. W. (1984), The Whistling Hunters: Field Studies of the Indian Wild Dog (Cuon Alpinus), Steven Simpson Books, ISBN 0-9524390-6-9 
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