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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

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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Geographic Range

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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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

"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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Physical Description

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.

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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

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 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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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

Food Habits

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

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

Lifespan/Longevity

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

"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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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.

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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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

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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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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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

Economic Importance for Humans: Negative

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

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Economic Importance for Humans: Positive

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 (/dl/), Cuon alpinus, also called the Asiatic wild dog or Indian wild dog, is a species of canid native to South and Southeast Asia. It is the only extant member of the genus Cuon, which differs from Canis by the reduced number of molars and greater number of teats. The dholes are classed as endangered by the IUCN, due to ongoing habitat loss, depletion of its prey base, competition from other predators, persecution and possibly diseases from domestic and feral dogs.[2]

The dhole is a highly social animal, living in large clans which occasionally split up into small packs to hunt.[3] It primarily preys on medium-sized ungulates, which it hunts by tiring them out in long chases, and kills by disemboweling them. Unlike most social canids (but similar to African wild dogs), dholes let their pups eat first at a kill.[4][5][6] Though fearful of humans, dhole packs are bold enough to attack large and dangerous animals such as wild boar, water buffalo, and even tigers.

Naming and etymology[edit]

Since these canids are called dholes only in English,[7] the etymology remains unclear,[8] but it may have come from Kannada: tōḷa (‘wolf’).[9] Some 19th-century authors[10] connected this word with Turkish: deli ‘mad, crazy’, and erroneously compared the Turkish word with Old Saxon: dol and Dutch: dol (cf. also English: dull; German: toll), which are in fact from Proto-Germanic *dwalaz ‘foolish, stupid’.[11]

Other names for the species include wild dogs,[12] whistling dogs, chennai,[13] red wolves[14] (not to be confused with Canis [lupus] rufus), red dogs[15] and mountain wolves.[16]

The generic name Cuon is an alternative spelling of cyōn (Ancient Greek: κύων, cf. also Procyonidae), meaning "dog".

Vernacular names[edit]

Vernacular names include lal rakshasa (red devil) and rakshur kukur (devil dog) in India, and jangli rakshasa (jungle devil) or hounds of Kali in Assam.[17] In the Himalayas, they are variously known as Bhaosa, Bhansa and Buansu.[7]

Evolution and taxonomy[edit]

Dholes are post-Pleistocene in origin, and are more closely related to jackals than they are to wolves.[19] One theory has dholes becoming social animals as an adaptation to living with tigers and Indian leopards.[20]

George Gaylord Simpson placed dholes under the subfamily Symocyoninae along with the African wild dog and bush dog on account of shared anatomical features, namely the reduction of postcarnassial molars. Many have questioned this classification, arguing that these shared features are due to convergent evolution. Juliet Clutton-Brock concluded from comparing the morphological, behavioural and ecological characteristics of 39 different canid species that with the exception of skull and dentition, dholes more closely resembled canids of the genus Canis, Dusicyon and Vulpes/Alopex than to African wild dogs and bush dogs.[19] A comparative study on dhole and other canid mtDNA in 1997 showed dholes diverged from the Lupus lupus lineage before the black-backed jackal and the golden jackal diverged, a few million years before the domestication of the dog.[21]

Subspecies[edit]

SubspeciesTrinomial authorityDescriptionRangeSynonyms
Indian dhole
Cuon a. dukhunensis[22]

Indian wild dog by N. A. Naseer.jpg

Sykes, 1831Peninsular India, south of Ganges River
Himalayan dhole
Cuon a. primaevus[22]
Hodgson, 1833Gentically distinct[23]Himalaya and northern parts of the Indian subcontinent. Probably Khangchendzonga National Park, Sikkim, India[23]
Kashmir dhole
Cuon a. laniger[22]
Pocock, 1936Kashmir, Lhasa
Kiangsi dhole
Cuon a. lepturus[22]

Cuon alpinus.jpg

Heude, 1892Southern Chinagrayiformis (Hodgson, 1863)

clamitans (Heude, 1892) rutilans (Müller, 1839)

Indochinese dhole
Cuon a. infuscus[22]
Pocock, 1936Myanmar, Thailand, Laos, Cambodia, Vietnamadustus (Pocock, 1941)
Sumatran dhole
Cuon a. sumatrensis[22]

Sumatran dhole.jpg

Hardwicke, 1821A small subspecies, it measures only two feet in length, and stands 360 mm (14 in) high at the shoulder. It has a pointed, black, fox-like muzzle with long, dark whiskers. The nose and lips are foxy brown mixed with black. The general colour is foxy ferraginous red, with lighter shades on the belly and inner sides of the legs.[24]Sumatra, Indonesia
Javan dhole
Cuon a. javanicus[22]
Desmarest, 1820Java, Indonesia
Eastern[25] or Ussuri dhole[26]
Cuon a. alpinus

Ussuridhole.JPG

Pallas, 1811This is the largest subspecies, with a long, narrow face and a skull measuring 189 mm long on average. The winter fur's general tone is intense rusty-red. The top of the head and the outer ears are brownish-rusty with black-brown highlights. The shoulders and upper surface of the back is brownish-rusty with black-brown highlights. The outer sides of the legs are rusty brown, while the inner sides of the legs and lower sides of the body are yellowish.[26]Russian Far East, China, Tibet, Mongoliafumosus (Pocock, 1936)
Western[25] or Tien Shan dhole[27]
Cuon a. hesperius

Tien shan dhole.jpg

Afanasjev and Zolotarev, 1935A small subspecies, it has a short, wide face and a skull measuring 180 mm long on average. The general tone of the winter fur is lighter-coloured than C. a. alpinus, with weakly developed rusty-red tints. The top of the head and outer sides of the ears are reddish-straw coloured. The upper surface of the neck is dirty-white, with a narrow, sandy-yellow-coloured band running along the upper surface of the back from the ears to the shoulders. The outer surfaces of the limbs are sandy-yellow, while the flanks and inner sides of the limbs have little to no yellowish tint.[27]Transoxiana, Eastern Russia and Chinajason (Pocock, 1936)
Late Pleistocene dhole
Cuon a. europaeus
Bourguignat, 1875The earliest form to evolve a singlely cusped, sharply trenchant tooth in place of the lower tubercular molar[28]Czech Republic, Hungary, Austria, Switzerland and French Riviera
Late Middle Pleistocene dhole
Cuon a. fossilis
Nehring, 1890An intermediate form between Cuon a. priscus and Cuon a. europaeus[28]Heppenloch, Germany
Early Middle Pleistocene dhole
Cuon a. priscus
Thenius, 1954

Physical description[edit]

Anatomy[edit]

Dhole skull, as illustrated by N. N. Kondakov.

Dholes have relatively short, heavy and massive skulls, with shortened facial regions, widely separated zygomatic arches and well-developed sagittal crests.[29] The frontal bone is inflated, and passes down onto the snout, giving the animals a convex rather than concave profile.[30] The masseter muscles are highly developed compared to other canid species, giving the face an almost hyena-like appearance.[31] The skull is broader, and has a shorter rostrum than that of domestic dogs and most other canids.[15] The dental formula is 3.1.4.23.1.4.2

The species uniquely 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,[29] an adaptation thought to improve shearing ability, thus allowing it to compete more successfully with kleptoparasites.[33] The canine teeth are slightly curved and short.[29]

Their limbs are moderately long, and their thoraces are proportional.[34] Along with African wild dogs, dholes are often referred to as "cat-like" canids, due to their long, fine limbs and backbones.[32] They have great jumping and leaping abilities, being able to jump 3.0–3.5 m (10–12 ft) high, and leap 5– to 6-m (17– to 20-ft) distances in one leap with a running start.[35] Their tails measure 16–17 in long,[32] and are almost half the length of their bodies, nearly touching the ground when in full winter fur.[36] They are smaller than African wild dogs.[37] Weights range from 10 to 25 kg (22 to 55 lb), with males averaging about 4.5 kg (9.9 lb) heavier than females. This dog is 88 to 113 cm (35 to 44 in) long from the snout to the base of the tail, with the tail averaging 45 cm (18 in) in length.[38][39] Shoulder height is 42 to 55 cm (17 to 22 in).[40] Like African wild dogs, their ears are rounded rather than pointed. However, unlike the former species, male dholes do not have a clearly visible prepuce, thus making the sexing of individuals difficult even at close proximity. Unlike members of the Canis genus, females have 12–14 teats rather than 10.[32] They are not as odorous as wolves, jackals and foxes, having a smaller number of anal scent glands.[41] Their stomachs have been estimated to hold 6.5 lb (2.9 kg) of food.[42]

Fur[edit]

The general tone of the fur is reddish, with the brightest hues occurring in winter. When in their winter fur, 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.[43] 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.[15]

Behaviour[edit]

Social and territorial behaviours[edit]

A pair of dholes at Port Lympne Wild Animal Park, Kent, UK

Dholes are more social than wolves,[18] and have less of a dominance hierarchy, as seasonal scarcity of food is not a serious concern for them as it is with wolves. In this sense, they closely resemble African wild dogs in social structure.[3] 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.[6] Intragroup fighting is rarely observed.[44] 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.[45] Dholes are far less territorial than wolves, with pups from one clan often joining another without trouble once they mature sexually.[46] Clans typically number 5-12 individuals in India, though clans of 40 have been reported. In Thailand, clans rarely exceed three individuals.[15] 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.[47]

Reproduction and development[edit]

Among Indian dholes, the mating season occurs between mid-October and January, while captive dholes in the Moscow Zoo breed mostly in February.[15] Unlike wolf packs, dhole clans may contain more than one breeding female.[6] More than one female dhole may den and rear their litters together in the same den.[48] During mating, the female assumes a crouched, cat-like position. There is no "tug of war" characteristic of other canids[further explanation needed] when the male dismounts. Instead, the pair lie on their sides facing each other in a semicircular formation.[49] The gestation period lasts 60–63 days, with litter sizes averaging four to six pups.[15] 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.[50] Maximum longevity in captivity is 15-16 years.[51]

Denning behaviours[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.[52] Like African wild dogs and dingoes, dholes will avoid killing prey close to their dens.[50]

Diet, hunting and feeding behaviours[edit]

Dholes attacking a sambar, Bandipur National Park
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.[4][15][53] 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.[16] In Kashmir, they may hunt markhor,[53] and thamin in Burma.[15] Javan rusas are hunted in Java.[33] 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.[54] Like African wild dogs, but unlike wolves, dholes are not known to attack people.[55][56] 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.[57] In summertime in the Tien Shan Mountains, dholes eat large quantities of mountain rhubarb.[54] Bael fruits are also eaten.[58] Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves.[59] 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.[60]

Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and homo- and heterosexual mounting.[61] 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.[62] Though not as fast as jackals and foxes, they can chase their prey for many hours.[54] 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.[63] When chasing fleet-footed prey, they run at a pace of 30 mph.[58] Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.[64]

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.[65] 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.[66] 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.[5] 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.[67] 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.[6] They are generally tolerant of scavengers at their kills.[68]

Tiger hunted by wild dogs (1807)

Both mother and young are provided with regurgitated food by other pack members.[69]

Relationships with other predators[edit]

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[70] 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).[70] 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.[70] 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.[55][71] 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.[72] Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs.[53] 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.[20] There are numerous records of leopards being treed by dholes.[73] Dholes sometimes drive tiger, leopards, and bears (see below) from their kills.[74] 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.[75]

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.[53]

Though usually antagonistic toward wolves,[76] they may hunt and feed alongside one another.[58] They infrequently associate in mixed groups with golden jackals. Domestic dogs may kill dholes, though they will feed alongside them on occasion.[77]

Communication[edit]

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.[78] Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries.[79] In contrast to wolves, dholes do not howl or bark.[80]

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.[47]

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.[81]

Range[edit]

Fossils of Cuon alpinus dated from Würm glaciation in the Iberian Peninsula.

Historical[edit]

Dholes once ranged throughout most of South, East and Southeast Asia, extending from the Tien Shan and Altai Mountains and the Primorsky Krai southward through Mongolia, Korea, China, Tibet, Nepal, India, and south-eastwards into Myanmar and Indochina, Thailand, Malaysia, Sumatra and Java.[82]

During the last glacial period, they ranged across most of Eurasia, and are known to have once inhabited North America from a single fossil find in the Gulf of Mexico.[83] A canid called the Sardinian dhole (Cynotherium sardous) lived on the Mediterranean island of Sardinia during the Pleistocene, but it is not as closely related to the living species as its name would imply.[84]

Current[edit]

Sleeping dhole at the Toronto Zoo

Dholes have not been reported recently in Russia, Mongolia, Kazakhstan, Kyrgyzstan and Tajikistan. One capture was reported in southern China's Jiangxi province. Dholes still occur in Tibet, particularly in southeast Tibet. They may still be present in North Korea. They still occur in India south of the Ganges River, especially in the central Indian Highlands and the Western and Eastern Ghats. Dholes also occur in northeast India's states of Arunachal Pradesh, Assam, Meghalaya and West Bengal and in Ladakh. They have a precarious, fragmented distribution in Himalaya and northwest India. They are occasionally reported in the Ladakh area of Kashmir, contiguous with the Tibetan highlands and China. In Bhutan, dholes have since recovered from a government-sponsored poisoning campaign started in the 1970s, with reports of livestock predation occurring in the lower Kheng region. It is uncertain if they still occur in Bangladesh. Camera trapping has confirmed dholes still occur in 11 survey areas in Myanmar, where they have replaced tigers as main predators. Dhole populations are highly fragmented in Thailand and Indochina, particularly in Vietnam. They are known to occur in four sites in northern and central Malaysia. In Java, they appear to be most common in the island's protected eastern and western ends. They are also known to occur in Sumatra's protected areas in the southern, central and northern areas.[82]

In Nepal, dholes were recorded in 2011 in the Churia hills of the Chitwan National Park located in the Terai.[85] They were also reported in the Dhorpatan Hunting Reserve in the late 1990s.[82]

Relationships with humans[edit]

Hunting and persecution[edit]

The dhole only rarely takes domestic livestock.[86] 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.[87] 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.[81] 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.[88] In Indochina, dholes suffer heavily from nonselective hunting techniques such as snaring.[81]

The fur trade does not pose a significant threat to dholes.[81] The people of India do not eat dhole flesh, and their fur is not considered overly valuable.[89] 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.[80]

Dholes in folklore, mythology, literature and popular culture[edit]

A dhole pack, as illustrated by Winifred Austen

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.[90] 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.[41] 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.[91]

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. They would reappear in two animated television adaptations: Jungle Book Shōnen Mowgli and Jungle Cubs.

In an episode called "Alpha" in season six of The X-Files, a cryptid dhole from China is blamed for multiple killings. This dhole (Wanshang dhole) is also mentioned in the Angel episode That Vision Thing. Both episodes were written by Jeffrey Bell.

Tameability and possible relation to the dog[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.[57] 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.[92] A dhole may have been presented as a gift to Ibbi-Sin as tribute.[93]

Although genetic distance and mitochondrial DNA studies have proven dogs descended from wolves, the dhole is still put forward by some researchers as a more likely candidate. Points raised in favour of the dhole ancestry cite the general domestic dog-like passiveness of the dhole in having its kills taken by humans, which contrasts with the behaviour of sympatric wolves, which will defend their prey aggressively against humans. It is also claimed that dhole skulls bear more similarities to dog skulls than wolves do, with the glaring exception of the dentition.[94]

References[edit]

Notes[edit]

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  2. ^ a b 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 Fox 1984, p. 85
  4. ^ a b Fox 1984, pp. 58–60
  5. ^ a b Fox 1984, p. 63
  6. ^ a b c d Fox 1984, pp. 86–7
  7. ^ a b c d e f g h i j k l m n o p q r Lydekker 1907, p. 360
  8. ^ dhole. Oxford dictionary on-line.
  9. ^ dhole. Merriam-Webster Dictionary.
  10. ^ Smith & Jardine 1839, p. 179
  11. ^ Orel, Vladimir (2003), A Handbook of Germanic Etymology, Leiden, Boston: Brill, p. 81, ISBN 90-04-12875-1 
  12. ^ Lydekker 1907
  13. ^ Fox 1984
  14. ^ Heptner & Naumov 1998
  15. ^ a b c d e f g h Cohen, James A. (1978). "Cuon alpinus". Mammalian Species 100 (100): 1–3. doi:10.2307/3503800. 
  16. ^ a b Perry 1965, p. 147
  17. ^ Perry 1965, p. 145
  18. ^ a b Shretha 1997, p. 121
  19. ^ a b Sillero-Zubiri, Hoffman & MacDonald 2004, p. 210
  20. ^ a b 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. 
  21. ^ Wayne, Robert K.; Geffen, Eli; Girman, Derek J.; Koepfli, Klaus P.; Lau, Lisa M. and Marshall, Charles R. (1997). "Molecular Systematics of the Canidae". Systematic Biology 46 (4): 622–653. doi:10.1093/sysbio/46.4.622. PMID 11975336. 
  22. ^ a b c d e f g Iyengar, A., et al. Phylogeography, genetic structure, and diversity in the dhole (Cuon alpinus). Molecular Ecology 14.8 (2005): 2281-2297.
  23. ^ a b Bashir, T. A. W. Q. I. R., et al. Precarious status of the Endangered dhole Cuon alpinus in the high elevation Eastern Himalayan habitats of Khangchendzonga Biosphere Reserve, Sikkim, India. Oryx (2013): 1-8.
  24. ^ Smith & Jardine 1839, pp. 186–7
  25. ^ a b Fox 1984, p. 40
  26. ^ a b Heptner & Naumov 1998, p. 578
  27. ^ a b Heptner & Naumov 1998, p. 579
  28. ^ a b Kurtén 1968, pp. 112–14
  29. ^ a b c Heptner & Naumov 1998, p. 567
  30. ^ Pocock 1941, p. 149
  31. ^ a b Fox 1984, pp. 61–2
  32. ^ a b c d Fox 1984, p. 41
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