# Dhole

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Species of mammal

"Mountain wolf" redirects here. For other uses, see [Mountain wolf (disambiguation)](/source/Mountain_wolf_(disambiguation)).

Not to be confused with [Red wolf](/source/Red_wolf), [Northern Rocky Mountain wolf](/source/Northern_Rocky_Mountain_wolf), or [African wild dog](/source/African_wild_dog).

Dhole Temporal range: Middle Pleistocene – Recent 0.78–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Conservation status Endangered (IUCN 3.1)[1] CITES Appendix II [1] Scientific classification Kingdom: Animalia Phylum: Chordata Class: Mammalia Infraclass: Placentalia Order: Carnivora Family: Canidae Subfamily: Caninae Tribe: Canini Genus: Cuon Hodgson, 1838 Species: C. alpinus Binomial name Cuon alpinus (Pallas, 1811) Subspecies C. a. adjustus C. a. alpinus C. a. fumosus † C. a. hesperius C. a. laniger C. a. lepturus C. a. sumatrensis † C. a. antiquus † C. a. caucasicus † C. a. europaeus dhole range Synonyms Canis alpinus

The **dhole** ([/doʊl/](https://en.wikipedia.org/wiki/Help:IPA/English) [*dohl*](https://en.wikipedia.org/wiki/Help:Pronunciation_respelling_key);[2][3] ***Cuon alpinus***) is a [canid](/source/Canidae) native to [South](/source/South_Asia), [East](/source/East_Asia) and [Southeast Asia](/source/Southeast_Asia). The sole member of the genus ***Cuon***, it is anatomically distinguished from members of the genus *[Canis](/source/Canis)* in several aspects: its [skull](/source/Skull) is convex rather than concave in profile, it lacks a third lower [molar](/source/Molar_(tooth)), and the upper molars possess only a single [cusp](/source/Cusp_(dentistry)) as opposed to between two and four. During the [Pleistocene](/source/Pleistocene), the dhole ranged throughout [Asia](/source/Asia), with its range also extending into [Europe](/source/Europe) (with a single putative, controversial record also reported from [North America](/source/North_America)) but became restricted to its historical range 12,000–18,000 years ago. It is now extinct in [Central Asia](/source/Central_Asia), parts of [Southeast Asia](/source/Southeast_Asia), and possibly the [Korean peninsula](/source/Korean_peninsula) and [Russia](/source/Russia).

The dhole is a highly social animal, living in large clans without rigid [dominance hierarchies](/source/Dominance_hierarchies) and containing multiple breeding females. Such clans usually consist of about 12 individuals, but groups of over 40 are known. It is a diurnal pack hunter which preferentially targets large and medium-sized [ungulates](/source/Ungulate). In tropical forests, the dhole competes with the [tiger](/source/Tiger) (*Panthera tigris*) and the [leopard](/source/Leopard) (*Panthera pardus*), targeting somewhat different prey species, but still with substantial dietary overlap.

It is listed as [Endangered](/source/Endangered) on the [IUCN Red List](/source/IUCN_Red_List), as populations are decreasing and estimated to comprise fewer than 2,500 mature individuals. Factors contributing to this decline include habitat loss, loss of prey, competition with other species, persecution due to livestock predation, and disease transfer from [domestic dogs](/source/Domestic_dog).

## Etymology and naming

The [etymology](/source/Etymology) of "dhole" is unclear. The possible earliest written use of the word in English occurred in 1808 by soldier Thomas Williamson, who encountered the animal in [Ramghur district](/source/Ramgarh_district), India. He stated that *dhole* was a common local name for the species.[4] In 1827, [Charles Hamilton Smith](/source/Charles_Hamilton_Smith) claimed that it was derived from a language spoken in 'various parts of the East'.[5]

Two years later, Smith connected this word with [Turkish](/source/Turkish_language): *[deli](https://en.wiktionary.org/wiki/deli#Turkish)* 'mad, crazy', and erroneously compared the Turkish word with [Old Saxon](/source/Old_Saxon_language): *[dol](https://en.wiktionary.org/wiki/dol#Old_English)* and [Dutch](/source/Dutch_language): *[dol](https://en.wiktionary.org/wiki/dol#Dutch)* (cfr. also English: [dull](https://en.wiktionary.org/wiki/dull#English); [German](/source/German_language): *[toll](https://en.wiktionary.org/wiki/toll#German)*),[6] which are in fact from the [Proto-Germanic](/source/Proto-Germanic_language) **dwalaz* 'foolish, stupid'.[7] [Richard Lydekker](/source/Richard_Lydekker) wrote nearly 80 years later that the word was not used by the natives living within the species' range.[8] The [Merriam-Webster](/source/Merriam-Webster) *Dictionary* theorises that it may have come from the [Kannada](/source/Kannada_language): ತೋಳ, romanized: *tōḷa*, [lit.](/source/Literal_translation) 'wolf'.[9]

Other English names for the species include **Asian wild dog**, **Asiatic wild dog**,[10] **Indian wild dog**,[8] **whistling dog**, **red dog**,[11] and **red wolf**.[12]

## Taxonomy and evolution

Skeletal remains of a [European dhole](/source/European_dhole) dating back to the upper [Würm](/source/Last_Glacial_Period#Würm_glaciation_(Alps)) period from Cova Negra de [Xàtiva](/source/X%C3%A0tiva), [Valencia](/source/Valencia), Spain

Illustration (1859) by [Leopold von Schrenck](/source/Leopold_von_Schrenck), one of the first accurate depictions of the species, based on a single skin purchased in the village of Dshare on the [Amur](/source/Amur)[13]

*Canis alpinus* was the [binomial name](/source/Binomial_name) proposed by [Peter Simon Pallas](/source/Peter_Simon_Pallas) in 1811, who described its range as encompassing the upper levels of Udskoi Ostrog in [Amurland](/source/Amurland), towards the eastern side and in the region of the upper [Lena River](/source/Lena_River), around the [Yenisei](/source/Yenisei) River and occasionally crossing into [China](/source/China).[14][15] This northern Russian range reported by Pallas during the 18th and 19th centuries is "considerably north" of where this species occurs today.[15]

*Canis primaevus* was a name proposed by [Brian Houghton Hodgson](/source/Brian_Houghton_Hodgson) in 1833 who thought that the dhole was a primitive *Canis* form and the [progenitor](/source/Progenitor) of the [domestic dog](/source/Domestic_dog).[16] Hodgson later took note of the dhole's physical distinctiveness from the genus *Canis* and proposed the genus *Cuon*.[17]

The first study on the origins of the species was conducted by paleontologist Erich Thenius, who concluded in 1955 that the dhole was a post-Pleistocene descendant of a golden jackal-like ancestor.[18] The paleontologist [Bjorn Kurten](/source/Bjorn_Kurten) wrote in his 1968 book *Pleistocene Mammals of Europe* that the primitive dhole *Canis majori* Del Campana 1913 —the remains of which have been found in [Villafranchian](/source/Villafranchian) era [Valdarno](/source/Valdarno), Italy and in China—was almost indistinguishable from the genus *Canis*. In comparison, the modern species has greatly reduced [molars](/source/Molar_(tooth)) and the [cusps](/source/Cusp_(anatomy)) have developed into sharply trenchant points. During the Early [Middle Pleistocene](/source/Middle_Pleistocene) there arose both *Canis majori stehlini* that was the size of a large [wolf](/source/Wolf), and the early dhole *Canis alpinus* Pallas 1811 which first appeared at [Hundsheim](/source/Hundsheim) and [Mosbach](/source/Mosbach) in Germany. In the [Late Pleistocene](/source/Late_Pleistocene) era the [European dhole](/source/European_dhole) (*C. a. europaeus*) was modern-looking and the transformation of the lower molar into a single cusped, slicing tooth had been completed; however, its size was comparable with that of a wolf. This subspecies became extinct in Europe at the end of the late [Würm](/source/Last_Glacial_Period#Würm_glaciation_(Alps)) period, but the species as a whole still inhabits a large area of Asia.[19] The European dhole may have survived up until the early [Holocene](/source/Holocene) in the [Iberian Peninsula](/source/Iberian_Peninsula),[20] and what is believed to be dhole remains have been found at Riparo Fredian in northern [Italy](/source/Italy) dated 10,800 years old.[21][22]

The vast Pleistocene range of this species also included numerous islands in Asia that this species no longer inhabits, such as [Sri Lanka](/source/Sri_Lanka), [Borneo](/source/Borneo) and possibly [Palawan](/source/Palawan) in the [Philippines](/source/Philippines).[23][24][25][26][27][28] Middle Pleistocene dhole fossils have also been found in the Matsukae Cave in northern [Kyushu](/source/Kyushu) Island in western [Japan](/source/Japan) and in the Lower Kuzuu fauna in [Tochigi Prefecture](/source/Tochigi_Prefecture) in [Honshu](/source/Honshu) Island, east Japan.[29] Dhole fossils from the Late Pleistocene dated to about 10,700 years before present are known from the [Luobi Cave](/source/Luobi_Cave) or Luobi-Dong cave in [Hainan Island](/source/Hainan_Island) in [south China](/source/South_China) where they no longer exist.[30] Additionally, fossils of canidae possibly belonging to dhole have been excavated from [Dajia River](/source/Dajia_River) in [Taichung County](/source/Taichung_County), Taiwan.[31]

A single record of the dhole is known from North America. This consists of a jaw fragment and teeth of [Late Pleistocene](/source/Late_Pleistocene) age found in San Josecito Cave in northeast Mexico, dating to around 27,000–11,000 years ago.[32] Other researchers have either considered this record as "insufficient" or suggested that further corroboration is required for the definitive taxonomic attribution of these specimens.[33][34]

Phylogenetic tree of the wolf-like canids with timing in millions of years[a] Caninae 3.5 Ma 3.0 2.5 2.0 0.96 0.6 0.38 Domestic dog Gray wolf Coyote African wolf Golden jackal Ethiopian wolf Dhole African wild dog 2.6 Side-striped jackal Black-backed jackal

Dholes are also known from the Middle and Late Pleistocene fossil record of Europe.[35] In 2021, the analyses of the [mitochondrial](/source/Mitochondrial_DNA) [genomes](/source/Genomes) extracted from the fossil remains of two extinct European dhole specimens from the Jáchymka cave, [Czech Republic](/source/Czech_Republic) dated 35,000–45,000 years old indicate that these were genetically [basal](/source/Basal_(phylogenetics)) to modern dholes and possessed much greater genetic diversity.[22]

The dhole's distinctive morphology has been a source of much confusion in determining the species' systematic position among the Canidae. [George Simpson](/source/George_Gaylord_Simpson) placed the dhole in the [subfamily](/source/Subfamily) Simocyoninae alongside the [African wild dog](/source/African_wild_dog) and the [bush dog](/source/Bush_dog), on account of all three species' similar dentition.[36] Subsequent authors, including [Juliet Clutton-Brock](/source/Juliet_Clutton-Brock), noted greater morphological similarities to canids of the genera *Canis*, *[Dusicyon](/source/Dusicyon)* and *[Alopex](/source/Arctic_fox)* than to either *[Speothos](/source/Speothos)* or *[Lycaon](/source/Lycaon_(genus))*, with any resemblance to the latter two being due to [convergent evolution](/source/Convergent_evolution).[37]

Some authors consider the extinct *Canis* [subgenus](/source/Subgenus) *[Xenocyon](/source/Xenocyon)* as ancestral to both the genus *Lycaon* and the genus *Cuon*.[38][39][40][41]: p149 Subsequent studies on the canid [genome](/source/Genome) revealed that the dhole and African wild dog are closely related to members of the genus *Canis*.[42] This closeness to *Canis* may have been confirmed in a menagerie in [Madras](/source/Madras), where according to zoologist [Reginald Innes Pocock](/source/Reginald_Innes_Pocock) there is a record of a dhole that interbred with a golden jackal.[43] DNA sequencing of the [Sardinian dhole](/source/Sardinian_dhole) (*Cynotherium sardous*) an extinct small canine species formerly native to the island of Sardinia in the Mediterranean, and which has often been suggested to have descended from *Xenocyon*, has found that it is most closely related to the living dhole among canines.[44]

### Admixture with the African wild dog

In 2018, [whole genome sequencing](/source/Whole_genome_sequencing) was used to compare all members (apart from the black-backed and side-striped jackals) of the genus *Canis*, along with the dhole and the [African wild dog](/source/African_wild_dog) (*Lycaon pictus*). There was strong evidence of ancient [genetic admixture](/source/Genetic_admixture) between the dhole and the African wild dog. Today, their ranges are remote from each other; however, during the [Pleistocene](/source/Pleistocene) era the dhole could be found as far west as Europe. The study proposes that the dhole's distribution may have once included the [Middle East](/source/Middle_East), from where it may have admixed with the African wild dog in [North Africa](/source/North_Africa). However, there is no evidence of the dhole having existed in the Middle East nor North Africa,[45] though the *Lycaon* was present in Europe during the Early Pleistocene, with its last record in the region dating to 830,000 years ago. Genetic evidence from the Sardinan dhole suggests that both Sardinian and modern dholes (which are estimated to have split from each other around 900,000 years ago) share ancestry from the *Lycaon* lineage, but this ancestry is significantly higher in modern dholes than in the Sardinian dhole.[44]

### Subspecies

Historically, up to ten subspecies of dholes have been recognised.[46] As of 2005[\[update\]](https://en.wikipedia.org/w/index.php?title=Dhole&action=edit), seven subspecies are recognised.[47][48]

However, studies on the dhole's [mtDNA](/source/MtDNA) and [microsatellite](/source/Microsatellite) genotype showed no clear subspecific distinctions. Nevertheless, two major phylogeographic groupings were discovered in dholes of the Asian mainland, which likely diverged during a [glaciation event](/source/Glacial_period). One population extends from South, Central and North India (south of the Ganges) into Myanmar, and the other extends from India north of the Ganges into northeastern India, Myanmar, Thailand and the Malaysian Peninsula. The origin of dholes in Sumatra and Java is, as of 2005[\[update\]](https://en.wikipedia.org/w/index.php?title=Dhole&action=edit), unclear, as they show greater relatedness to dholes in India, Myanmar and China rather than with those in nearby Malaysia. However, the [Canid Specialist Group](https://en.wikipedia.org/w/index.php?title=Canid_Specialist_Group&action=edit&redlink=1) of the [International Union for the Conservation of Nature](/source/International_Union_for_the_Conservation_of_Nature) (IUCN) states that further research is needed because all of the samples were from the southern part of this species' range and the [Tien Shan](/source/Tien_Shan) subspecies has distinct morphology.[49]

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.[50] Fossils of dhole from the early [Middle Pleistocene](/source/Middle_Pleistocene) have been found in [Java](/source/Java).[51]

Subspecies Image Trinomial authority Description Distribution Synonyms C. a. adjustus Burmese dhole,[43] Indian dhole Pocock, 1941[43] Reddish coat, short hair on the paws and black whiskers[52] Northeastern India and south of the Ganges River, northern Myanmar[52] antiquus (Matthew & Granger, 1923), dukhunensis (Sykes, 1831) C. a. alpinus Ussuri dhole[12] (nominate subspecies) Pallas, 1811[14] Thick tawny red coat, greyish neck and ochre muzzle[52] East of the eastern Sayan Mountains, eastern Russia, northeastern Asia[52] – C. a. fumosus[53] Pocock, 1936[53] Luxuriant yellowish-red coat, dark back and grey neck[52] Western Sichuan, China and Mongolia. Southern Myanmar, Thailand, Laos, Cambodia, Vietnam, Malaysia and Java, Indonesia[52] infuscus (Pocock, 1936), javanicus (Desmarest, 1820) †C. a. hesperius Tien Shan dhole[12] Afanasjev and Zolotarev, 1935[54] Long yellow tinted coat, white underside and pale whiskers[52] Smaller than C. a. alpinus, with wider skull and lighter-coloured winter fur.[12] Altai, Tian Shan and Pamir mountain ranges.[1] Currently considered to be extinct since 1946.[55][1] jason (Pocock, 1936) C. a. laniger[53] Pocock, 1936[53] Full, yellowish-grey coat, tail not black but same colour as body[52] Southern Tibet, Himalayan Nepal, Sikkim, Bhutan and Kashmir[52] grayiformis (Hodgson, 1863), primaevus (Hodgson, 1833) C. a. lepturus[53] Heude, 1892[56] Uniform red coat with thick underfur[52] South of the Yangtze River, China[52] clamitans (Heude, 1892), rutilans (Müller, 1839), sumatrensis (Hodgson, 1833) Sumatran dhole and Javan dhole C. a. sumatrensis[57] Hardwicke, 1821[58] Red coat and dark whiskers[52] Sumatra, Indonesia[52] Its range is highly fragmented with multiple protected areas in Sumatra and Java.[1]

## Characteristics

A dhole in [Kanha National Park](/source/Kanha_National_Park), India

Dhole skull and molars illustrated by [St. George Mivart](/source/St._George_Mivart) (1890)

Captive adult dhole

The general tone of the dhole's fur is reddish, with the brightest hues appearing in winter. During the winter coat phase, the back is covered in a saturated rusty-red to reddish color, featuring brownish highlights along the top of the head, neck, and shoulders. The throat, chest, flanks, belly, and upper parts of the limbs are less vividly colored, exhibiting a more yellowish 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 luxurious and fluffy, mainly a reddish-ocher color, with a dark brown tip. The summer coat is shorter, coarser, and darker.[12] The dorsal and lateral [guard hairs](/source/Guard_hair) in adults measure 20–30 mm (0.79–1.18 in) in length. Dholes in the [Moscow Zoo](/source/Moscow_Zoo) moult once a year from March to May.[11] A [melanistic](/source/Melanism) individual was recorded in the northern [Coimbatore](/source/Coimbatore) Forest Division in [Tamil Nadu](/source/Tamil_Nadu).[59]

The dhole has a wide and massive skull with a well-developed [sagittal crest](/source/Sagittal_crest),[12] and its [masseter muscles](/source/Masseter_muscle) are highly developed compared to other canid species, giving the face an almost [hyena](/source/Hyena)-like appearance.[60] The [rostrum](/source/Rostrum_(anatomy)) is shorter than that of domestic dogs and most other canids.[11] It has six rather than seven lower [molars](/source/Molar_(tooth)).[61] The upper molars are weak, being one third to one half the size of those of wolves and have only one [cusp](/source/Cusp_(dentistry)) as opposed to between two and four, as is usual in canids,[12] an adaptation thought to improve shearing ability and thus speed of prey consumption. This may allow dholes to compete more successfully with [kleptoparasites](/source/Kleptoparasite).[52] In terms of size, dholes average about 88–113 cm (35–44 in) in length (excluding a 41–50 cm (16–20 in) long tail), and stand around 42–50 cm (17–20 in) at the shoulders.[11] Adult females can weigh 10–17 kg (22–37 lb), while the slightly larger male may weigh 15–21 kg (33–46 lb). The mean weight of adults from three small samples was 15.1 kg (33 lb).[52][62]

In appearance, the dhole has been variously described as combining the physical characteristics of the [gray wolf](/source/Gray_wolf) and the [red fox](/source/Red_fox),[12] and as being "cat-like" on account of its long backbone and slender limbs.[18]

## Distribution and habitat

Lone dhole strolling through the jungle in [Mudumalai National Park](/source/Mudumalai_National_Park)

Dhole in [Tadoba Andhari Tiger Reserve](/source/Tadoba_Andhari_Tiger_Reserve), Maharashtra, India

Historically, the dhole lived in [Singapore](/source/Singapore) and throughout [Central Asia](/source/Central_Asia) including [Afghanistan](/source/Afghanistan), [Kyrgyzstan](/source/Kyrgyzstan), [Kazakhstan](/source/Kazakhstan), [Mongolia](/source/Mongolia), [Tajikistan](/source/Tajikistan) and [Uzbekistan](/source/Uzbekistan), though it is now considered to be regionally extinct in these regions.[1] Historical record in [South Korea](/source/South_Korea) from the [Veritable Records of the Joseon Dynasty](/source/Veritable_Records_of_the_Joseon_Dynasty) also indicate that the dhole once inhabited [Yangju](/source/Yangju) in [Gyeonggi Province](/source/Gyeonggi_Province),[63] but it is now also extinct in South Korea,[1] with the last known capture reports in 1909 and 1921 from [Yeoncheon](/source/Yeoncheon) of [Gyeonggi Province](/source/Gyeonggi_Province).[64] The current presence of dholes in [North Korea](/source/North_Korea) and [Pakistan](/source/Pakistan) is considered uncertain.[1] The dholes also once inhabited the alpine steppes extending into [Kashmir](/source/Kashmir) to the [Ladakh](/source/Ladakh) area, though they disappeared from 60% of their historic range in India during the past century.[1] In India, Myanmar, Indochina, Indonesia and China, it prefers forested areas in [alpine zones](/source/Alpine_zone) and is occasionally sighted in [plains](/source/Plain) regions.[12]

In the [Bek-Tosot Conservancy](https://en.wikipedia.org/w/index.php?title=Bek-Tosot_Conservancy&action=edit&redlink=1) of southern [Kyrgyzstan](/source/Kyrgyzstan), the possible presence of the dholes was considered likely based on genetic samples collected in 2019.[65] This was the first record of dholes from the country in almost three decades.[65]

The dhole might still be present in the [Tunkinsky National Park](/source/Tunkinsky_National_Park) in extreme southern [Siberia](/source/Siberia) near [Lake Baikal](/source/Lake_Baikal).[66] It possibly still lives in the [Primorsky Krai](/source/Primorsky_Krai) province in far eastern Russia, where it was considered a rare and endangered species in 2004, with unconfirmed reports in the Pikthsa-Tigrovy Dom protected forest area; no sighting was reported in other areas since the late 1970s.[67][*[page needed](https://en.wikipedia.org/wiki/Wikipedia:Citing_sources)*] Currently, no other recent reports are confirmed of dholes being present in [Russia](/source/Russia),[55] so the IUCN considered them to be extinct in Russia.[1] However, the dhole might be present in the eastern [Sayan Mountains](/source/Sayan_Mountains) and in the [Transbaikal](/source/Transbaikal) region; it has been sighted in [Tofalaria](/source/Tofalaria) in the [Irkutsk Oblast](/source/Irkutsk_Oblast), the [Republic of Buryatia](/source/Republic_of_Buryatia) and [Zabaykalsky Krai](/source/Zabaykalsky_Krai).[55]

One pack was sighted in the [Qilian Mountains](/source/Qilian_Mountains) in 2006.[68] In 2011 to 2013, local government officials and herders reported the presence of several dhole packs at elevations of 2,000 to 3,500 m (6,600 to 11,500 ft) near [Taxkorgan Nature Reserve](/source/Taxkorgan_Nature_Reserve) in the [Xinjiang](/source/Xinjiang) Autonomous Region. Several packs and a female adult with pups were also recorded by [camera traps](/source/Camera_trap) at elevations of around 2,500 to 4,000 m (8,200 to 13,100 ft) in Yanchiwan National Nature Reserve in the northern [Gansu Province](/source/Gansu_Province) in 2013–2014.[69] Dholes have been also reported in the [Altyn-Tagh](/source/Altyn-Tagh) Mountains.[70]

In China's [Yunnan](/source/Yunnan) Province, dholes were recorded in [Baima Xueshan Nature Reserve](https://en.wikipedia.org/w/index.php?title=Baima_Xueshan_Nature_Reserve&action=edit&redlink=1) in 2010–2011.[71] Dhole samples were obtained in [Jiangxi](/source/Jiangxi) Province in 2013.[72] Confirmed records by camera-trapping since 2008 have occurred in southern and western [Gansu](/source/Gansu) province, southern [Shaanxi](/source/Shaanxi) province, southern [Qinghai](/source/Qinghai) province, southern and western [Yunnan](/source/Yunnan) province, western [Sichuan](/source/Sichuan) province, the southern [Xinjiang](/source/Xinjiang) Autonomous Region and in the Southeastern [Tibet](/source/Tibet) Autonomous Region.[73] There are also historical records of dhole dating to 1521–1935 in Hainan Island, but the species is no longer present and is estimated to have become extinct around 1942.[30]

The dhole occurs in most of India south of the Ganges, particularly in the Central Indian Highlands and the [Western](/source/Western_Ghats) and [Eastern Ghats](/source/Eastern_Ghats). It is also present in [Arunachal Pradesh](/source/Arunachal_Pradesh), [Assam](/source/Assam), [Meghalaya](/source/Meghalaya) and [West Bengal](/source/West_Bengal) and in the [Indo-Gangetic Plain](/source/Indo-Gangetic_Plain)'s [Terai](/source/Terai) region. Dhole populations in the [Himalayas](/source/Himalaya) and northwest India are fragmented.[1]

In 2011, dhole packs were recorded by camera traps in the [Chitwan National Park](/source/Chitwan_National_Park).[74] Its presence was confirmed in the [Kanchenjunga Conservation Area](/source/Kanchenjunga_Conservation_Area) in 2011 by camera traps.[75] In February 2020, dholes were sighted in the [Vansda National Park](/source/Vansda_National_Park), with camera traps confirming the presence of two individuals in May of the same year. This was the first confirmed sighting of dholes in [Gujarat](/source/Gujarat) since 1970.[76]

In [Bhutan](/source/Bhutan), the dhole is present in [Jigme Dorji National Park](/source/Jigme_Dorji_National_Park).[77][78]

In [Bangladesh](/source/Bangladesh), it inhabits forest reserves in the [Sylhet](/source/Sylhet) area, as well the [Chittagong Hill Tracts](/source/Chittagong_Hill_Tracts) in the southeast. Recent camera trap photos in the Chittagong in 2016 showed the continued presence of the dhole.[79] These regions probably do not harbor a viable population, as mostly small groups or solitary individuals were sighted.[1]

In [Myanmar](/source/Myanmar), the dhole is present in several protected areas.[1] In 2015, dholes and tigers were recorded by camera-traps for the first time in the hill forests of [Karen State](/source/Karen_State).[80]

Its range is highly fragmented in the [Malaysian Peninsula](/source/Malaysian_Peninsula), [Sumatra](/source/Sumatra), [Java](/source/Java), [Vietnam](/source/Vietnam) and [Thailand](/source/Thailand), with the Vietnamese population considered to be possibly extinct.[1] In 2014, camera trap videos in the montane tropical forests at 2,000 m (6,600 ft) in the [Kerinci Seblat National Park](/source/Kerinci_Seblat_National_Park) in Sumatra revealed its continued presence.[81] A camera trapping survey in the [Khao Ang Rue Nai Wildlife Sanctuary](/source/Khao_Ang_Rue_Nai_Wildlife_Sanctuary) in Thailand from January 2008 to February 2010 documented one healthy dhole pack.[82] In northern [Laos](/source/Laos), dholes were studied in [Nam Et-Phou Louey](/source/Nam_Et-Phou_Louey) National Protected Area.[62] Camera trap surveys from 2012 to 2017 recorded dholes in the same Nam Et-Phou Louey National Protected Area.[83]

In [Vietnam](/source/Vietnam), dholes were sighted only in [Pu Mat National Park](/source/Pu_Mat_National_Park) in 1999, in [Yok Don National Park](/source/Yok_Don_National_Park) in 2003 and 2004; and in [Ninh Thuan Province](/source/Ninh_Thuan_Province) in 2014.[84]

A disjunct dhole population was reported in the area of [Trabzon](/source/Trabzon) and [Rize](/source/Rize) in northeastern [Turkey](/source/Turkey) near the border with [Georgia](/source/Georgia_(country)) in the 1990s.[85] This report was not considered to be reliable.[1] One single individual was claimed to have been shot in 2013 in the nearby [Kabardino-Balkaria](/source/Kabardino-Balkaria) Republic of Russia in the central [Caucasus](/source/Caucasus); its remains were analysed in May 2015 by a biologist from the [Kabardino-Balkarian State University](/source/Kabardino-Balkarian_State_University), who concluded that the skull was indeed that of a dhole.[86] In August 2015, researchers from the [National Museum of Natural History](/source/National_Museum_of_Natural_History_(Bulgaria)) and the [Karadeniz Technical University](/source/Karadeniz_Technical_University) started an expedition to track and document possible Turkish population of dhole.[87] In October 2015, they concluded that two skins of alleged dholes in Turkey probably belonged to dogs, pending DNA analysis of samples from the skins, and, having analysed photos of the skull of alleged dhole in Kabardino-Balkaria Republic of Russia, they concluded it was a grey wolf.[88]

## Ecology and behaviour

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.[89] Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries.[90] In contrast to wolves, dholes do not [howl](/source/Howling) or bark.[12]

Dholes have a complex [body language](/source/Body_language). Friendly or submissive greetings are accompanied by horizontal lip retraction and the lowering of the tail, as well as licking. Playful dholes open their mouths with their lips retracted and their tails held in a vertical position whilst assuming a play bow. Aggressive or threatening dholes 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.[91]

### Social and territorial behaviour

Dholes are more social than [gray wolves](/source/Gray_wolves),[12] 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.[92] They live in [clans](/source/Clan) rather than [packs](/source/Pack_(canine)), as the latter term refers to a group of animals that always hunt together. In contrast, dhole clans frequently break into small packs of three to five animals, particularly during the spring season, as this is the optimal number for catching fawns.[93] 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.[94] Intragroup fighting is rarely observed.[95]

Dholes playing, [Pench National Park](/source/Pench_National_Park)

A pack of Asiatic wild dogs

Dholes are far less [territorial](/source/Territory_(animal)) than wolves, with pups from one clan often joining another without trouble once they mature sexually.[96] Clans typically number 5 to 12 individuals in India, though clans of 40 have been reported. In [Thailand](/source/Thailand), clans rarely exceed three individuals.[11] Unlike other canids, there is no evidence of dholes using [urine](/source/Urine) to mark their territories or travel routes. When urinating, dholes, especially males, may [raise one hind leg](/source/Raised-leg_urination) or both to result in a handstand. Handstand urination is also seen in [bush dogs](/source/Bush_dog) (*Speothos venaticus*)[97] and domestic dogs.[98] They may defecate in conspicuous places, though a territorial function is unlikely, as [faeces](/source/Faeces) are mostly deposited within the clan's territory rather than the periphery. Faeces are often deposited in what appear to be communal [latrines](/source/Animal_latrine). They do not scrape the earth with their feet, as other canids do, to mark their territories.[91]

### Denning

Four kinds of [den](/source/Burrow) have been described; simple earth dens with one entrance (usually remodeled [striped hyena](/source/Striped_hyena) or [porcupine](/source/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 30 m (98 ft) of interconnecting tunnels. These "cities" may be developed over many generations of dholes, and are shared by the clan females when raising young together.[99] Like [African wild dogs](/source/African_wild_dog) and [dingoes](/source/Dingo), dholes will avoid killing prey close to their dens.[100]

### Reproduction and development

Dhole pup, [Kolmården Wildlife Park](/source/Kolm%C3%A5rden_Wildlife_Park)

In India, the [mating season](/source/Mating_season) occurs between mid-October and January, while captive dholes in the [Moscow Zoo](/source/Moscow_Zoo) breed mostly in February.[11] Unlike wolf packs, dhole clans may contain more than one breeding female.[94] More than one female dhole may den and rear their litters together in the same den.[95] During [mating](/source/Mating), the female assumes a crouched, cat-like position. There is no [copulatory tie](/source/Copulatory_tie) characteristic of other canids when the male dismounts. Instead, the pair lie on their sides facing each other in a semicircular formation.[101] The [gestation period](/source/Gestation_period) lasts 60–63 days, with litter sizes averaging four to six pups.[11]

The [hormone](/source/Hormone) metabolites of five males and three females kept in Thai zoos was studied. The breeding males showed an increased level of [testosterone](/source/Testosterone) from October to January. The [oestrogen](/source/Oestrogen) level of captive females increases for about two weeks in January, followed by an increase of [progesterone](/source/Progesterone). They displayed sexual behaviours during the oestrogen peak of the females.[102]

Pups are suckled at least 58 days. During this time, the pack feeds the mother at the den site. Dholes do not use [rendezvous](https://en.wiktionary.org/wiki/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](/source/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 for 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](/source/Sambar_deer) by the age of eight months.[100] Maximum longevity in captivity is 15–16 years.[95]

### Hunting behaviour

Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and mounting.[103] Dholes are primarily [diurnal](/source/Diurnality) hunters, hunting in the early hours of the morning. They rarely hunt at night, except on moonlit nights, indicating they greatly rely on [sight](/source/Visual_perception) when hunting.[104] They can chase their prey for many hours.[12] During a pursuit, one or more dholes takes over chasing the 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 (1,600 ft).[105] When chasing fleet-footed prey, they run at a pace of 50 km/h (30 mph).[12] Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.[106]

Dholes attacking a [sambar](/source/Sambar_(deer)) (*Rusa unicolor*), [Bandipur National Park](/source/Bandipur_National_Park)

Once large prey is caught, one dhole grabs 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.[107] They occasionally blind their prey by attacking the eyes.[108] [Serows](/source/Serow) 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.[8] Dholes tear open their prey's flanks and [disembowel](/source/Disembowelment) it, eating the [heart](/source/Heart), [liver](/source/Liver), [lungs](/source/Lungs) and some sections of the [intestines](/source/Intestines). The [stomach](/source/Stomach) and [rumen](/source/Rumen) are usually left untouched.[109] Prey weighing less than 50 kg (110 lb) is usually killed within two minutes, while large stags may take 15 minutes to die. Once prey is secured, dholes tear off pieces of the carcass and eat in seclusion.[110] They give the pups access to a kill.[94] They are generally tolerant of [scavengers](/source/Scavenger) at their kills.[111] Both mother and young are provided with regurgitated food by other pack members.[95]

### Feeding ecology

Prey animals in India include [chital](/source/Chital), [sambar deer](/source/Sambar_deer), [muntjac](/source/Muntjac), [mouse deer](/source/Mouse_deer), [barasingha](/source/Barasingha), [wild boar](/source/Wild_boar), [gaur](/source/Gaur), [water buffaloes](/source/Water_buffalo), [banteng](/source/Banteng), [cattle](/source/Cattle), [nilgai](/source/Nilgai), [goats](/source/Goat), [Indian hares](/source/Indian_hare), [Himalayan field rats](/source/Himalayan_field_rat) and [langurs](/source/Langur).[11][43][112] There is one record of a pack bringing down an [Indian elephant](/source/Indian_elephant) calf in [Assam](/source/Assam), despite desperate defense of the mother, resulting in numerous losses to the pack.[113] In Kashmir, they prey on [markhor](/source/Markhor),[43] and [thamin](/source/Thamin) in Myanmar,[11] [Malayan tapir](/source/Malayan_tapir), [Sumatran serow](/source/Sumatran_serow) in [Sumatra](/source/Sumatra) and the [Malay Peninsula](/source/Malay_Peninsula) and [Javan rusa](/source/Javan_rusa) in Java.[52] In the [Tian Shan](/source/Tian_Shan) and [Tarbagatai Mountains](/source/Tarbagatai_Mountains), dholes prey on [Siberian ibexes](/source/Siberian_ibex), [arkhar](/source/Arkhar), [roe deer](/source/Roe_deer), [Caspian red deer](/source/Caspian_red_deer) and [wild boar](/source/Wild_boar). In the [Altai](/source/Altai_Mountains) and [Sayan Mountains](/source/Sayan_Mountains), they prey on [musk deer](/source/Musk_deer) and [reindeer](/source/Reindeer). In eastern Siberia, they prey on roe deer, [Manchurian wapiti](/source/Manchurian_wapiti), wild pig, musk deer and reindeer, while in [Primorye](/source/Primorye) they feed on [sika deer](/source/Sika_deer) and [goral](/source/Goral). In Mongolia, they prey on [argali](/source/Argali) and rarely Siberian ibex.[12]

Dholes feeding on a [chital](/source/Chital), Bandipur National Park

Dhole feeding on [sambar deer](/source/Sambar_deer) carcass, [Khao Yai National Park](/source/Khao_Yai_National_Park)

Like African wild dogs, but unlike wolves, dholes are not known to actively hunt people.[12][43] They are known to eat [insects](/source/Insect) and [lizards](/source/Lizard).[114] Dholes eat [fruit](/source/Fruit) and [vegetable](/source/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.[115] In summertime in the Tian Shan Mountains, dholes eat large quantities of mountain [rhubarb](/source/Rhubarb).[12] Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves.[116] [Livestock](/source/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](/source/Oxen) are killed more often than [cows](/source/Cow), probably because they are given less protection.[117]

### Enemies and competitors

Dhole killed and cached in a tree by a [leopard](/source/Leopard), India

In some areas, dholes are [sympatric](https://en.wiktionary.org/wiki/sympatric) to [tigers](/source/Tiger) and [leopards](/source/Leopard). Competition between these species is mostly avoided through differences in prey selection, although there is still substantial dietary overlap. Along with leopards, dholes typically target animals in the 30–175 kg (66–386 lb) range (mean weights of 35.3 kg (78 lb) for dhole and 23.4 kg (52 lb) for leopard), while tigers selected for prey animals heavier than 176 kg (388 lb) (but their mean prey weight was 65.5 kg (144 lb)). 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](/source/Langur) 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.[118]

Tigers are dangerous opponents for dholes, as they have sufficient strength to kill a dhole with a single paw strike.[113] Dhole packs are smaller in areas with higher tiger densities due to tigers directly killing dholes and stealing kills they made. The kleptoparasitism causes dholes to prefer hunting smaller animals because they can eat more of a smaller carcass before a tiger arrives to steal it. Direct predation can lead to lower reproductive and recruitment rates, lower hunting success rates and less food for the pups when a helper is killed, and potentially pack destabilization if one member of the breeding pair is killed.[119]

Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs.[43] There are numerous records of leopards being treed by dholes.[95] Dholes were once thought to be a major factor in reducing [Asiatic cheetah](/source/Asiatic_cheetah) populations, though this is doubtful, as cheetahs live in open areas as opposed to forested areas favoured by dholes.[120] Since leopards are smaller than tigers and are more likely to hunt dholes, dhole packs tend to react more aggressively toward them than they do towards tigers.[121]

Dhole packs occasionally attack [Asiatic black bears](/source/Asiatic_black_bear), [snow leopards](/source/Snow_leopard) and [sloth bears](/source/Sloth_bear). When attacking bears, dholes will attempt to prevent them from seeking refuge in caves and lacerate their hindquarters.[43] Although usually antagonistic toward [wolves](/source/Wolves),[12] they may hunt and feed alongside one another.[122]

The dhole is also sympatric with the [Indian wolf](/source/Indian_wolf) (*Canis lupus pallipes*) in parts of its range.[123][124] There is at least one record of a lone wolf associating with a pair of dholes in [Debrigarh Wildlife Sanctuary](/source/Debrigarh_Wildlife_Sanctuary),[125] and two observations in [Satpura Tiger Reserve](/source/Satpura_Tiger_Reserve).[126] They infrequently associate in mixed groups with [golden jackals](/source/Golden_jackal). Domestic dogs may kill dholes, though they will feed alongside them on occasion.[127]

### Diseases and parasites

Dholes are vulnerable to a number of different diseases, particularly in areas where they are [sympatric](https://en.wiktionary.org/wiki/Sympatric) with other canid species. Infectious pathogens such as *[Toxocara canis](/source/Toxocara_canis)* are present in their faeces. They may suffer from [rabies](/source/Rabies), [canine distemper](/source/Canine_distemper), [mange](/source/Mange), [trypanosomiasis](/source/Trypanosomiasis), [canine parvovirus](/source/Canine_parvovirus) and [endoparasites](/source/Endoparasite) such as [cestodes](/source/Cestode) and [roundworms](/source/Roundworm).[52]

## Threats

[Habitat loss](/source/Habitat_loss) is thought to amount to 60% of the dhole's historical range in India. The fragmentation and isolation of dhole populations has resulted in inbreeding and the [Allee effect](/source/Allee_effect), which threaten its long-term viability.[128]

Some ethnic groups like the [Kuruba](/source/Kuruba) and [Mon Khmer](/source/Mon_Khmer)-speaking tribes will appropriate dhole kills; some Indian villagers welcome the dhole because of this appropriation of dhole kills.[95] Dholes were persecuted throughout India for bounties until they were given protection by the [Wildlife Protection Act of 1972](/source/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](/source/British_Raj) did so under the conviction that dholes were responsible for drops in [game](/source/Game_(hunting)) populations. Persecution of dholes still occurs with varying degrees of intensity according to the region.[52] Bounties paid for dholes used to be 25 [rupees](/source/Rupee), though this was reduced to 20 in 1926 after the number of presented dhole carcasses became too numerous to maintain the established reward.[129] The Indochinese dhole population suffers heavily from nonselective hunting techniques such as [snaring](/source/Snaring).[52] The [fur trade](/source/Fur_trade) does not pose a significant threat to the dhole.[52] The people of India do not eat dhole flesh and their fur is not considered overly valuable.[115] Due to their rarity, dholes were never harvested for their skins in large numbers in the [Soviet Union](/source/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](/source/Ussuriysk) during the late 1860s for a few silver [rubles](/source/Ruble). In the early 20th century, dhole pelts reached eight rubles in [Manchuria](/source/Manchuria). In [Semirechye](/source/Jetisu), [fur coats](/source/Fur_coat) made from dhole skin were considered the warmest, but were very costly.[12]

## Conservation

In India, the dhole is protected under Schedule 2 of the [Wildlife Protection Act, 1972](/source/Wildlife_Protection_Act%2C_1972). The creation of reserves under [Project Tiger](/source/Project_Tiger) provided some protection for dhole populations sympatric with tigers. In 2014, the Indian government sanctioned its first dhole conservation breeding centre at the [Indira Gandhi Zoological Park](/source/Indira_Gandhi_Zoological_Park) (IGZP) in [Visakhapatnam](/source/Visakhapatnam).[130] The dhole has been protected in Russia since 1974, though it is vulnerable to poison left out for wolves. In China, the animal is listed as a category II protected species under the Chinese wildlife protection act of 1988. In [Cambodia](/source/Cambodia), the dhole is protected from all hunting, while conservation laws in Vietnam limit extraction and utilisation.[1]

In 2016, the Korean company Sooam Biotech was reported to be attempting to clone the dhole using dogs as [surrogate mothers](/source/Surrogate_mother) to help conserve the species.[131]

## In culture and literature

[Russian ruble](/source/Russian_ruble)

[Kazakhstani tenge](/source/Kazakhstani_tenge)

Drawing of a dhole from the Imperial Chinese Encyclopedia

Three dhole-like animals are featured on the [coping](/source/Coping_(architecture)) stone of the [Bharhut](/source/Bharhut) [stupa](/source/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](/source/Treeing) tigers.[132] In [Tamil Sangam literature](/source/Tamil_Sangam_literature) from 100 BC to 200 AD, the dhole is a recurring character, a device of [metonymy](/source/Metonymy) in song; [Ainkurunuru](/source/Ainkurunuru) 323 narrates a pregnant dhole craving to eat wild pig, so its male mate hunts for pig.[133]

Its fearsome reputation in India is reflected by the number of pejorative names it possesses in [Hindi](/source/Hindi), which variously translate as "red devil", "devil dog", "jungle devil", or "hound of [Kali](/source/Kali)".[113]

Dholes appeared in [The Second Jungle Book](/source/The_Second_Jungle_Book) in the chapter [Red Dog](/source/Red_Dog_(Kipling_short_story)), in which Mowgli and the Seeonee Wolf Pack must battle a pack of dholes (referred to as "red dogs") that have invaded their part of the jungle.[134]

[Leopold von Schrenck](/source/Leopold_von_Schrenck) had trouble obtaining dhole specimens during his exploration of [Amurland](/source/Amurland), as the local [Gilyaks](/source/Gilyaks) greatly feared the species. This fear and superstition was not shared by neighbouring [Tungusic peoples](/source/Tungusic_peoples). It was speculated that this differing attitude towards the dhole was due to the Tungusic peoples' more nomadic, hunter-gatherer lifestyle.[13]

Japanese author Uchida Roan wrote about the declining popularity of indigenous dog breeds in 1901, which he asserted were descended from the dhole.[135]

In China, the dhole was widely known throughout history and mythology. One notable legendary creature is the [Nine sons of the dragon](/source/Nine_sons_of_the_dragon), which was believed to be a creature that was part-dhole, part-dragon. In modern times, the Chinese word for dhole 豺; *Chái* is often confused with 'jackal' or 'wolf', resulting in many confusions and mistranslations of dholes as jackals or wolves.[136]

A dhole may have been presented as a gift to the Akkadian king [Ibbi-Sin](/source/Ibbi-Sin) as tribute referred to in the inscription as the "red dog of Meluhha" or [Indus Valley Civilisation](/source/Indus_Valley_Civilisation).[137]

### Tameability

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.[138] 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.[8]

## See also

- *[Wild Dog Diaries](/source/Wild_Dog_Diaries)*

## Notes

1. **[^](#cite_ref-mtdna_seq_desc_35-0)** For a full set of supporting references refer to the note (a) in the phylotree at [Evolution of the wolf#Wolf-like canids](/source/Evolution_of_the_wolf#Wolf-like_canids)

## References

1. ^ [***a***](#cite_ref-iucn_1-0) [***b***](#cite_ref-iucn_1-1) [***c***](#cite_ref-iucn_1-2) [***d***](#cite_ref-iucn_1-3) [***e***](#cite_ref-iucn_1-4) [***f***](#cite_ref-iucn_1-5) [***g***](#cite_ref-iucn_1-6) [***h***](#cite_ref-iucn_1-7) [***i***](#cite_ref-iucn_1-8) [***j***](#cite_ref-iucn_1-9) [***k***](#cite_ref-iucn_1-10) [***l***](#cite_ref-iucn_1-11) [***m***](#cite_ref-iucn_1-12) [***n***](#cite_ref-iucn_1-13) [***o***](#cite_ref-iucn_1-14) [***p***](#cite_ref-iucn_1-15) Kamler, J.F.; Songsasen, N.; Jenks, K.; Srivathsa, A.; Sheng, L.; Kunkel, K. (2015). ["*Cuon alpinus*"](https://www.iucnredlist.org/species/5953/72477893). *[IUCN Red List of Threatened Species](/source/IUCN_Red_List)*. **2015** e.T5953A72477893. [doi](/source/Doi_(identifier)):[10.2305/IUCN.UK.2015-4.RLTS.T5953A72477893.en](https://doi.org/10.2305%2FIUCN.UK.2015-4.RLTS.T5953A72477893.en). Retrieved 24 January 2022.

1. **[^](#cite_ref-MWeb_2-0)** [""dhole""](https://www.merriam-webster.com/dictionary/dhole). *Merriam-Webster*. [Archived](https://web.archive.org/web/20240107071959/https://www.merriam-webster.com/dictionary/dhole) from the original on 7 January 2024. Retrieved 7 January 2024.

1. **[^](#cite_ref-ColDict_3-0)** [""dhole""](https://www.collinsdictionary.com/dictionary/english/dhole). *Collins Dictionary*. [Archived](https://web.archive.org/web/20210501140052/https://www.collinsdictionary.com/dictionary/english/dhole) from the original on 1 May 2021. Retrieved 7 January 2024.

1. **[^](#cite_ref-4)** Williamson, T. (1808). [*Oriental field sports: being a complete, detailed, and accurate description of the wild sports of the East*](https://books.google.com/books?id=1BlcAAAAcAAJ&pg=PA1). Vol. II. London: Orme.

1. **[^](#cite_ref-smith1827_5-0)** Smith, C. H. (1827). [*The class Mammalia*](https://archive.org/stream/classmammalia21827cuvi#page/326/mode/2up/search/dhole). London: Geo. B. Whittaker.

1. **[^](#cite_ref-smith1839_6-0)** Smith, C. H.; Jardine, W. (1839). [*The natural history of dogs: Canidae or genus canis of authors; including also the genera hyaena and proteles*](https://archive.org/stream/naturalhistoryof139smit#page/166/mode/2up). Vol. I. Edinburgh, UK: W.H. Lizars.

1. **[^](#cite_ref-7)** [Orel, V.](/source/Vladimir_Orel) (2003), [*A Handbook of Germanic Etymology*](https://archive.org/details/handbookofgerman0000orel/page/81), Leiden, DE; Boston, MA: Brill, p. [81](https://archive.org/details/handbookofgerman0000orel/page/81), [ISBN](/source/ISBN_(identifier)) [978-90-04-12875-0](https://en.wikipedia.org/wiki/Special:BookSources/978-90-04-12875-0)

1. ^ [***a***](#cite_ref-lydekker1907_8-0) [***b***](#cite_ref-lydekker1907_8-1) [***c***](#cite_ref-lydekker1907_8-2) [***d***](#cite_ref-lydekker1907_8-3) [Lydekker, R.](/source/Richard_Lydekker) (1907). [*The game animals of India, Burma, Malaya, and Tibet*](https://archive.org/stream/gameanimalsofind00lyde#page/360/mode/2up). London, UK: R. Ward Limited.

1. **[^](#cite_ref-9)** [dhole](http://www.merriam-webster.com/dictionary/dhole) [Archived](https://web.archive.org/web/20161224095139/https://www.merriam-webster.com/dictionary/dhole) 24 December 2016 at the [Wayback Machine](/source/Wayback_Machine). Merriam-Webster Dictionary.

1. **[^](#cite_ref-f_10-0)** [Fox 1984](#CITEREFFox1984)

1. ^ [***a***](#cite_ref-cohen1978_11-0) [***b***](#cite_ref-cohen1978_11-1) [***c***](#cite_ref-cohen1978_11-2) [***d***](#cite_ref-cohen1978_11-3) [***e***](#cite_ref-cohen1978_11-4) [***f***](#cite_ref-cohen1978_11-5) [***g***](#cite_ref-cohen1978_11-6) [***h***](#cite_ref-cohen1978_11-7) [***i***](#cite_ref-cohen1978_11-8) Cohen, James A. (1978). ["Cuon alpinus"](https://doi.org/10.2307%2F3503800). *Mammalian Species* (100): 1–3. [Bibcode](/source/Bibcode_(identifier)):[1978MamSp.100....1C](https://ui.adsabs.harvard.edu/abs/1978MamSp.100....1C). [doi](/source/Doi_(identifier)):[10.2307/3503800](https://doi.org/10.2307%2F3503800). [JSTOR](/source/JSTOR_(identifier)) [3503800](https://www.jstor.org/stable/3503800).

1. ^ [***a***](#cite_ref-heptner1998_12-0) [***b***](#cite_ref-heptner1998_12-1) [***c***](#cite_ref-heptner1998_12-2) [***d***](#cite_ref-heptner1998_12-3) [***e***](#cite_ref-heptner1998_12-4) [***f***](#cite_ref-heptner1998_12-5) [***g***](#cite_ref-heptner1998_12-6) [***h***](#cite_ref-heptner1998_12-7) [***i***](#cite_ref-heptner1998_12-8) [***j***](#cite_ref-heptner1998_12-9) [***k***](#cite_ref-heptner1998_12-10) [***l***](#cite_ref-heptner1998_12-11) [***m***](#cite_ref-heptner1998_12-12) [***n***](#cite_ref-heptner1998_12-13) [***o***](#cite_ref-heptner1998_12-14) [***p***](#cite_ref-heptner1998_12-15) [***q***](#cite_ref-heptner1998_12-16) [***r***](#cite_ref-heptner1998_12-17) Heptner, V. G.; Naumov, N. P., eds. (1998). ["Genus *Cuon* Hodgson, 1838"](https://archive.org/stream/mammalsofsov211998gept#page/566/mode/2up). *Mammals of the Soviet Union*. Vol. (II. Part 1A: Sirenia and Carnivora (Sea Cows, Wolves, and Bears)). Washington, DC: Smithsonian Institution and National science Foundation. pp. 566–586. [ISBN](/source/ISBN_(identifier)) [1-886106-81-9](https://en.wikipedia.org/wiki/Special:BookSources/1-886106-81-9).

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1. ^ [***a***](#cite_ref-Pallas1811_14-0) [***b***](#cite_ref-Pallas1811_14-1) Pallas, P. S. (1811). ["*Canis alpinus*"](https://archive.org/stream/zoographiarossoa11831pall#page/34/mode/2up). *Zoographia Rosso-Asiatica: Sistens omnium animalium in extenso Imperio Rossico, et adjacentibus maribus observatorum recensionem, domicilia, mores et descriptiones, anatomen atque icones plurimorum* (in Latin). Petropoli: In officina Caes. Acadamiae Scientiarum Impress. pp. 34–35.

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1. ^ [***a***](#cite_ref-f80_101-0) [***b***](#cite_ref-f80_101-1) [Fox 1984](#CITEREFFox1984), p. 80

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1. **[^](#cite_ref-109)** Grassman, L. I. Jr.; M. E. Tewes; N. J. Silvy & K. Kreetiyutanont (2005). ["Spatial ecology and diet of the dhole *Cuon alpinus* (Canidae, Carnivora) in north central Thailand"](https://web.archive.org/web/20061123090326/http://www.mnhn.fr/museum/front/medias/publication/6555_m05n1a2). *Mammalia*. **69** (1): 11–20. [doi](/source/Doi_(identifier)):[10.1515/mamm.2005.002](https://doi.org/10.1515%2Fmamm.2005.002). [S2CID](/source/S2CID_(identifier)) [85198149](https://api.semanticscholar.org/CorpusID:85198149). Archived from [the original](http://www.mnhn.fr/museum/front/medias/publication/6555_m05n1a2) on 23 November 2006.

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1. ^ [***a***](#cite_ref-perry1965_114-0) [***b***](#cite_ref-perry1965_114-1) [***c***](#cite_ref-perry1965_114-2) Perry, R. (1964). *The World of the Tiger*. London: Cassell.

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1. ^ [***a***](#cite_ref-mivart1890_116-0) [***b***](#cite_ref-mivart1890_116-1) Mivart, George (1890). ["Genus *Cyon*, Hodgson (1838)"](https://archive.org/stream/dogsjackalswolve00mivauoft#page/n307/mode/2up). *Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ*. London: R.H. Porter. pp. 177–188.

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1. **[^](#cite_ref-118)** Johnsingh, A.J.T.; Yonten, D.; Wangchuck, S. (2007). ["Livestock-Dhole Conflict in Western Bhutan"](https://www.biodiversitylibrary.org/page/48380954). *Journal of the Bombay Natural History Society*. **104** (2): 201–202 – via Biodiversity Heritage Library.

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1. **[^](#cite_ref-venkataraman_122-0)** Venkataraman, A. (1995). ["Do dholes (*Cuon alpinus*) live in packs in response to competition with or predation by large cats?"](http://eprints.iisc.ac.in/25085/1/article_id_069_11_0934_0936_0.pdf) (PDF). *Current Science*. **69** (11): 934–936. [Archived](https://web.archive.org/web/20190713030406/http://eprints.iisc.ac.in/25085/1/article_id_069_11_0934_0936_0.pdf) (PDF) from the original on 13 July 2019.

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1. **[^](#cite_ref-124)** Mukherjee, S.; Zelcer, M. & Kotler, B.P. (2009). "Patch use in time and space for a meso-predator in a risky world". *Oecologia*. **159** (3): 661–668. [Bibcode](/source/Bibcode_(identifier)):[2009Oecol.159..661M](https://ui.adsabs.harvard.edu/abs/2009Oecol.159..661M). [doi](/source/Doi_(identifier)):[10.1007/s00442-008-1243-3](https://doi.org/10.1007%2Fs00442-008-1243-3). [PMID](/source/PMID_(identifier)) [19082629](https://pubmed.ncbi.nlm.nih.gov/19082629). [S2CID](/source/S2CID_(identifier)) [24051254](https://api.semanticscholar.org/CorpusID:24051254).

1. **[^](#cite_ref-125)** Afik, D.; Pinshow, B. (1993). "Temperature regulation and water economy in desert wolves". *Journal of Arid Environments*. **24** (2): 197–209. [Bibcode](/source/Bibcode_(identifier)):[1993JArEn..24..197A](https://ui.adsabs.harvard.edu/abs/1993JArEn..24..197A). [doi](/source/Doi_(identifier)):[10.1006/jare.1993.1017](https://doi.org/10.1006%2Fjare.1993.1017).

1. **[^](#cite_ref-126)** Nair M. V. & Panda S. K. (2013). ["Just Friends"](https://web.archive.org/web/20190724195645/https://www.sanctuaryasia.com/magazines/features/9390-just-friends.html). *Sanctuary Asia*. **XXXIII**: 3. Archived from [the original](http://www.sanctuaryasia.com/magazines/features/9390-just-friends.html) on 24 July 2019. Retrieved 23 December 2014.

1. **[^](#cite_ref-127)** Ghaskadbi, P.; Das, J.; Mahadev, V. & Habib, B. (2021). ["First record of mixed species association between dholes and a wolf from Satpura Tiger Reserve, India"](https://www.canids.org/CBC/23/Dhole_wolf_association.pdf) (PDF). *Canid Biology & Conservation*. **23** (4): 15–17. [Archived](https://web.archive.org/web/20211017142640/https://www.canids.org/CBC/23/Dhole_wolf_association.pdf) (PDF) from the original on 17 October 2021.

1. **[^](#cite_ref-humphrey1990_128-0)** Humphrey, S. R. & Bain, J. R. (1990). *Endangered Animals of Thailand*. Gainesville: Sandhill Crane Press. [ISBN](/source/ISBN_(identifier)) [978-1-877743-07-8](https://en.wikipedia.org/wiki/Special:BookSources/978-1-877743-07-8).

1. **[^](#cite_ref-129)** Modi, S.; Mondol, S.; Nigam, P.; Habib, B. (2021). ["Genetic analyses reveal demographic decline and population differentiation in an endangered social carnivore, Asiatic wild dog"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8361113). *Nature*. **11** (1): 16371. [Bibcode](/source/Bibcode_(identifier)):[2021NatSR..1116371M](https://ui.adsabs.harvard.edu/abs/2021NatSR..1116371M). [doi](/source/Doi_(identifier)):[10.1038/s41598-021-95918-3](https://doi.org/10.1038%2Fs41598-021-95918-3). [PMC](/source/PMC_(identifier)) [8361113](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8361113). [PMID](/source/PMID_(identifier)) [34385570](https://pubmed.ncbi.nlm.nih.gov/34385570).

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1. **[^](#cite_ref-131)** [Zoo to have conservation breeding centre for 'dhole'](http://www.thehindu.com/news/cities/Visakhapatnam/zoo-to-have-conservation-breeding-centre-for-dhole/article6326027.ece) [Archived](https://web.archive.org/web/20240303095610/https://www.thehindu.com/fragment/latestnews?count=4) 3 March 2024 at the [Wayback Machine](/source/Wayback_Machine), *The Hindu* (18 August 2014)

1. **[^](#cite_ref-NewScientist500_132-0)** Zastrow, M. (8 February 2016). ["Inside the cloning factory that creates 500 new animals a day"](https://www.newscientist.com/article/2076681-inside-the-cloning-factory-that-creates-500-new-animals-a-day/). *New Scientist*. [Archived](https://web.archive.org/web/20160222043834/https://www.newscientist.com/article/2076681-inside-the-cloning-factory-that-creates-500-new-animals-a-day/) from the original on 22 February 2016. Retrieved 23 February 2016.

1. **[^](#cite_ref-133)** Van der Geer, A. A. E. (2008). *Animals in stone: Indian mammals sculptured through time*. Leiden: Brill. [ISBN](/source/ISBN_(identifier)) [978-90-04-16819-0](https://en.wikipedia.org/wiki/Special:BookSources/978-90-04-16819-0).

1. **[^](#cite_ref-134)** ["Ainkurunūru 323, Ōthalānthaiyār, Pālai Thinai – What the hero said to his heart"](https://sangamtranslationsbyvaidehi.com/ettuthokai-ainkurunuru/#:~:text=Ainkurun%C5%ABru%20323%2C%20%C5%8Cthal%C4%81nthaiy%C4%81r%2C%20P%C4%81lai%20Thinai%20%E2%80%93%20What%20the%20hero%20said%20to%20his%20heart). *Sangam Translations*.

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1. **[^](#cite_ref-136)** Skabelund, A. H. (2011). *Empire of Dogs: Canines, Japan, and the Making of the Modern Imperial World*. Cornell University Press, p. 85, [ISBN](/source/ISBN_(identifier)) [0801463246](https://en.wikipedia.org/wiki/Special:BookSources/0801463246)

1. **[^](#cite_ref-137)** Schafer, E. H. (1991). "Brief Note: The Chinese Dhole". *Asia Major*. **4** (1): 1–6. [JSTOR](/source/JSTOR_(identifier)) [41645459](https://www.jstor.org/stable/41645459).

1. **[^](#cite_ref-138)** McIntosh, J. (2008). *The ancient Indus Valley: new perspectives*. ABC-CLIO. [ISBN](/source/ISBN_(identifier)) [978-1-57607-907-2](https://en.wikipedia.org/wiki/Special:BookSources/978-1-57607-907-2).

1. **[^](#cite_ref-139)** Hodgson, B.H. (1833). ["Description of the Wild Dog of the Himalaya"](https://www.biodiversitylibrary.org/item/137991#page/584/mode/1up). *Asiatic Researches*. **XVIII**: 226. [Archived](https://web.archive.org/web/20210129081829/https://www.biodiversitylibrary.org/item/137991#page/584/mode/1up) from the original on 29 January 2021. Retrieved 14 January 2021.

### Bibliography

- Fox, M.W. (1984). [*The Whistling Hunters: Field Studies of the Asiatic Wild Dog (*Cuon alpinus*)*](https://books.google.com/books?id=V8qMs6x-PhUC). Albany: State University of New York Press. [ISBN](/source/ISBN_(identifier)) [978-0-9524390-6-6](https://en.wikipedia.org/wiki/Special:BookSources/978-0-9524390-6-6).

- Karanth, U.K. (2006). [*A View From the Machan*](https://books.google.com/books?id=W6ks4b0l7NgC). Karnataka: Permanent Black. [ISBN](/source/ISBN_(identifier)) [978-8-1782413-7-1](https://en.wikipedia.org/wiki/Special:BookSources/978-8-1782413-7-1).

## External links

Look up ***[dhole](https://en.wiktionary.org/wiki/Special:Search/dhole)*** in Wiktionary, the free dictionary.

- [Dhole Home Page](http://www.cuon.net/dholes/) ([Archive](https://web.archive.org/web/20170919112743/http://www.cuon.net:80/dholes))

- ARKive – [images and movies of the dhole](https://web.archive.org/web/20060313170514/http://www.arkive.org/species/GES/mammals/Cuon_alpinus/)

- [Saving the dhole: The forgotten 'badass' Asian dog more endangered than tigers](https://www.theguardian.com/environment/radical-conservation/2015/jun/25/dhole-asia-endangered-tiger-ignored), *The Guardian* (25 June 2015)

- [Photos of dhole in Bandipur](https://web.archive.org/web/20110608190250/http://public.fotki.com/msandeep/wild-dogs---bandipur/)

v t e Extant Carnivora species Kingdom: Animalia Phylum: Chordata Class: Mammalia Infraclass: Eutheria Superorder: Laurasiatheria Feliformia ("cat-like" carnivorans) Feliformia African palm civet (Nandinia binotata) Feloidea Prionodon (Asiatic linsangs) Banded linsang (P. linsang) Spotted linsang (P. pardicolor) Felidae (cats) Pantherinae Neofelis Sunda clouded leopard (N. diardi) Clouded leopard (N. nebulosa) Panthera Snow leopard (P. uncia) Tiger (P. tigris) Jaguar (P. onca) Lion (P. leo) Leopard (P. pardus) Felinae sensu stricto Bay cat lineage Pardofelis Marbled cat (P. marmorata) Catopuma Bay cat (C. badia) Asian golden cat (C. temminckii) Caracal lineage Serval (Leptailurus serval) Caracal African golden cat (C. aurata) Caracal (C. caracal) Leopardus Pampas cat (L. colocola) Geoffroy's cat (L. geoffroyi) Kodkod (L. guigna) Southern tiger cat (L. guttulus) Andean mountain cat (L. jacobita) Ocelot (L. pardalis) Clouded tiger cat (L. pardinoides) Oncilla (L. tigrinus) Margay (L. wiedii) Lynx Canada lynx (L. canadensis) Eurasian lynx (L. lynx) Iberian lynx (L. pardinus) Bobcat (L. rufus) Puma lineage Acinonyx Cheetah (A. jubatus) Jaguarundi (Herpailurus yagouaroundi) Puma Cougar (P. concolor) Leopard cat lineage Pallas's cat (Otocolobus manul) Prionailurus Leopard cat (P. bengalensis) Sunda leopard cat (P. javanensis) Flat-headed cat (P. planiceps) Rusty-spotted cat (P. rubiginosus) Fishing cat (P. viverrinus) Felis Chinese mountain cat (F. bieti) Domestic cat (F. catus) Jungle cat (F. chaus) African wildcat (F. lybica) Sand cat (F. margarita) Black-footed cat (F. nigripes) European wildcat (F. silvestris) Viverroidea see below↓ Viverroidea Viverridae Palm civets Hemigalinae Owston's palm civet (Chrotogale owstoni) Otter civet (Cynogale bennettii) Hose's palm civet (Diplogale hosei) Banded palm civet (Hemigalus derbyanus) Paradoxurinae Binturong (Arctictis binturong) Small-toothed palm civet (Arctogalidia trivirgata) Sulawesi palm civet (Macrogalidia musschenbroekii) Masked palm civet (Paguma larvata) Paradoxurus Asian palm civet (P. hermaphroditus) Brown palm civet (P. jerdoni) Golden palm civet (P. zeylonensis) Viverrinae sensu lato Viverrinae sensu stricto Small Indian civet (Viverricula indica) African civet (Civettictis civetta) Viverra Malabar large-spotted civet (V. civettina) Large-spotted civet (V. megaspila) Malayan civet (V. tangalunga) Large Indian civet (V. zibetha) Genettinae Poiana (African linsangs) Central African oyan (P. richardsonii) West African oyan (P. leightoni) Genetta (genets) Abyssinian genet (G. abyssinica) Angolan genet (G. angolensis) Bourlon's genet (G. bourloni) Crested servaline genet (G. cristata) South African small-spotted genet (G. felina) Common genet (G. genetta) Johnston's genet (G. johnstoni) Letaba genet (G. letabae) Rusty-spotted genet (G. maculata) Pardine genet (G. pardina) Aquatic genet (G. piscivora) King genet (G. poensis) Schouteden's genet (G. schoutedeni) Servaline genet (G. servalina) Hausa genet (G. thierryi) Cape genet (G. tigrina) Giant forest genet (G. victoriae) Herpestoidea see below↓ Herpestoidea Hyaenidae (hyenas) Proteles Aardwolf (P. cristata) Hyaeninae (bone-crushing hyenas) Striped hyena (Hyaena hyaena) Brown hyena (Parahyaena brunnea) Crocuta Spotted hyena (C. crocuta) Herpestidae sensu lato Eupleridae (Malagasy carnivorans) Euplerinae (Malagasy civets) Fossa (Cryptoprocta ferox) Malagasy civet (Fossa fossana) Eupleres (falanoucs) Eastern falanouc (E. goudotii) Western falanouc (E. major) Galidiinae (vontsira) Ring-tailed vontsira (Galidia elegans) Galidictis Broad-striped Malagasy mongoose (G. fasciata) Grandidier's mongoose (G. grandidieri) Narrow-striped mongoose (Mungotictis decemlineata) Salanoia Brown-tailed mongoose (S. concolor) Durrell's vontsira (S. durrelli) Herpestidae sensu stricto (mongooses) Mungotinae Suricata Meerkat (S. suricatta) Liberian mongoose (Liberiictus kuhni) Mungos Gambian mongoose (M. gambianus) Banded mongoose (M. mungo) Pousargues's mongoose (Dologale dybowskii) Helogale Ethiopian dwarf mongoose (H. hirtula) Common dwarf mongoose (H. parvula) Crossarchus (kusimanses) Alexander's kusimanse (C. alexandri) Angolan kusimanse (C. ansorgei) Common kusimanse (C. obscurus) Flat-headed kusimanse (C. platycephalus) Herpestinae Marsh mongoose (Atilax paludinosus) Long-nosed mongoose (Xenogale naso) Urva (Asian mongooses) Small Indian mongoose (U. auropunctata) Short-tailed mongoose (U. brachyura) Indian grey mongoose (U. edwardsii) Indian brown mongoose (U. fusca) Javan mongoose (U. javanica) Collared mongoose (U. semitorquata) Ruddy mongoose (U. smithii) Crab-eating mongoose (U. urva) Stripe-necked mongoose (U. vitticolla) White-tailed mongoose (Ichneumia albicauda) Yellow mongoose (Cynictis penicillata) Selous's mongoose (Paracynictis selousi) Meller's mongoose (Rhynchogale melleri) Bdeogale Bushy-tailed mongoose (B. crassicauda) Jackson's mongoose (B. jacksoni) Black-footed mongoose (B. nigripes) Herpestes (slender mongooses) Angolan slender mongoose (H. flavescens) Egyptian mongoose (H. ichneumon) Somalian slender mongoose (H. ochracea) Cape gray mongoose (H. pulverulenta) Common slender mongoose (H. sanguinea) Caniformia ("dog-like" carnivorans) Canidae (dogs) Urocyon Gray fox (U. cinereoargenteus) Island fox (U. littoralis) Vulpini Bat-eared fox (Otocyon megalotis) Nyctereutes (raccoon dogs) Common raccoon dog (N. procyonoides) Japanese raccoon dog (N. viverrinus) Vulpes (true foxes) Bengal fox (V. bengalensis) Blanford's fox (V. cana) Cape fox (V. chama) Corsac fox (V. corsac) Tibetan fox (V. ferrilata) Arctic fox (V. lagopus) Kit fox (V. macrotis) Pale fox (V. pallida) Rüppell's fox (V. rueppelli) Swift fox (V. velox) Red fox (V. vulpes) Fennec fox (V. zerda) Canini (true dogs) Cerdocyonina (zorro) Maned wolf (Chrysocyon brachyurus) Speothos Bush dog (S. venaticus) Short-eared dog (Atelocynus microtis) Crab-eating fox (Cerdocyon thous) Lycalopex (South American foxes) Culpeo (L. culpaeus) Darwin's fox (L. fulvipes) South American gray fox (L. griseus) Pampas fox (L. gymnocercus) Sechuran fox (L. sechurae) Hoary fox (L. vetulus) Canina (wolf-like canids) Lupulella Side-striped jackal (L. adustus) Black-backed jackal (L. mesomelas) Lycaon African wild dog (L. pictus) Dhole (Cuon alpinus) Canis Golden jackal (C. aureus) Domestic dog (C. familiaris) Coyote (C. latrans) African wolf (C. lupaster) Wolf (C. lupus) Eastern wolf (C. lycaon) Red wolf (C. rufus) Ethiopian wolf (C. simensis) Arctoidea Ursidae (bears) Ailuropoda Giant panda (A. melanoleuca) Tremarctos Spectacled bear (T. ornatus) Ursinae Sloth bear (Melursus ursinus) Sun bear (Helarctos malayanus) Ursus American black bear (U. americanus) Asian black bear (U. thibetanus) Brown bear (U. arctos) Polar bear (U. maritimus) Mustelida Pinnipedia (seals) see below↓ Musteloidea see below↓ Pinnipedia (seals) Odobenidae Walrus (Odobenus rosmarus) Otariidae (eared seals) Callorhinus (northern fur seals) Northern fur seal (C. ursinus) Otariinae (sea lions) Steller sea lion (Eumetopias jubatus) Zalophus California sea lion (Z. californianus) Galápagos sea lion (Z. wollebaeki) South American sea lion (Otaria flavescens) Neophoca Australian sea lion (N. cinerea) New Zealand sea lion (Phocarctos hookeri) Arctocephalus (southern fur seals) South American fur seal (A. australis) Australasian fur seal (A. forsteri) Galápagos fur seal (A. galapagoensis) Antarctic fur seal (A. gazella) Juan Fernández fur seal (A. philippii) Brown fur seal (A. pusillus) Guadalupe fur seal (A. townsendi) Subantarctic fur seal (A. tropicalis) Phocidae (earless seals or true seals) Phocinae ("northern seals") Bearded seal (Erignathus barbatus) Hooded seal (Cystophora cristata) Phocini Harp seal (Pagophilus groenlandicus) Ribbon seal (Histriophoca fasciata) Grey seal (Halichoerus grypus) Phoca Spotted seal (P. largha) Harbor seal (P. vitulina) Pusa Caspian seal (P. caspica) Ringed seal (P. hispida) Baikal seal (P. sibirica) Monachinae ("southern seals") Monachini (monk seals) Mediterranean monk seal (Monachus monachus) Neomonachus Hawaiian monk seal (N. schauinslandi) Mirounga (elephant seals) Northern elephant seal (M. angustirostris) Southern elephant seal (M. leonina) Lobodontini (Antarctic seals) Leopard seal (Hydrurga leptonyx) Weddell seal (Leptonychotes weddellii) Crabeater seal (Lobodon carcinophagus) Ross seal (Ommatophoca rossi) Musteloidea Ailuridae Red panda (Ailurus fulgens) Mephitidae (skunks) Conepatus (hog-nosed skunks) Molina's hog-nosed skunk (C. chinga) Humboldt's hog-nosed skunk (C. humboldtii) American hog-nosed skunk (C. leuconotus) Striped hog-nosed skunk (C. semistriatus) Mephitis Hooded skunk (M. macroura) Striped skunk (M. mephitis) Mydaus (stink badgers) Sunda stink badger (M. javanensis) Palawan stink badger (M. marchei) Spilogale (spotted skunks) Southern spotted skunk (S. angustifrons) Western spotted skunk (S. gracilis) Eastern spotted skunk (S. putorius) Pygmy spotted skunk (S. pygmaea) Procyonidae Kinkajou (Potos flavus) Bassariscus Ringtail (B. astutus) Cacomistle (B. sumichrasti) Procyon (raccoons) Crab-eating raccoon (P. cancrivorus) Raccoon (P. lotor) Cozumel raccoon (P. pygmaeus) Bassaricyon (olingos) Eastern lowland olingo (B. alleni) Northern olingo (B. gabbii) Western lowland olingo (B. medius) Olinguito (B. neblina) Nasuina (coatis) Nasua White-nosed coati (N. narica) South American coati (N. nasua) Nasuella (mountain coatis) Eastern mountain coati (N. meridensis) Western mountain coati (N. olivacea) Mustelidae see below↓ Mustelidae Mustelidae American badger (Taxidea taxus) Mellivora Honey badger (M. capensis) Melinae Arctonyx (hog badgers) Northern hog badger (A. albogularis) Greater hog badger (A. collaris) Sumatran hog badger (A. hoevenii) Meles (Eurasian badgers) Japanese badger (M. anakuma) Caucasian badger (M. canescens) Asian badger (M. leucurus) European badger (M. meles) Melogale (ferret-badgers) Vietnam ferret-badger (M. cucphuongensis) Bornean ferret badger (M. everetti) Chinese ferret-badger (M. moschata) Javan ferret-badger (M. orientalis) Burmese ferret-badger (M. personata) Formosan ferret-badger (M. subaurantiaca) Guloninae Tayra (Eira barbara) Pekania Fisher (P. pennanti) Gulo Wolverine (G. gulo) Martes (martens) American marten (M. americana) Pacific marten (M. caurina) Yellow-throated marten (M. flavigula) Beech marten (M. foina) Nilgiri marten (M. gwatkinsii) European pine marten (M. martes) Japanese marten (M. melampus) Sable (M. zibellina) Ictonychinae Lyncodontini Patagonian weasel (Lyncodon patagonicus) Galictis (grisons) Lesser grison (G. cuja) Greater grison (G. vittata) Ictonychini (African polecats) Vormela Marbled polecat (V. peregusna) African striped weasel (Poecilogale albinucha) Ictonyx Saharan striped polecat (I. libycus) Striped polecat (I. striatus) Lutrinae (otters) Giant otter (Pteronura brasiliensis) Lontra North American river otter (L. canadensis) Marine otter (L. felina) Neotropical otter (L. longicaudis) Southern river otter (L. provocax) Enhydra Sea otter (E. lutris) Spotted-necked otter (Hydrictis maculicollis) Lutra Eurasian otter (L. lutra) Hairy-nosed otter (L. sumatrana) Lutrogale Smooth-coated otter (L. perspicillata) Aonyx African clawless otter (A. capensis) Asian small-clawed otter (A. cinereus) Congo clawless otter (A. congicus) Mustelinae Neogale (New World weasels) Amazon weasel (N. africana) Colombian weasel (N. felipei) Long-tailed weasel (N. frenata) American mink (N. vison) Mustela (weasels) subgenus Mustela (paraphyletic) Sichuan weasel (M. aistoodonnivalis) Mountain weasel (M. altaica) Stoat/Beringian ermine (M. erminea) Haida ermine (M. haidarum) Yellow-bellied weasel (M. kathiah) Least weasel (M. nivalis) American ermine (M. richardsonii) subgenus Lutreola (paraphyletic) Japanese weasel (M. itatsi) European mink (M. lutreola) Indonesian mountain weasel (M. lutreolina) Malayan weasel (M. nudipes) Siberian weasel (M. sibirica) Back-striped weasel (M. strigidorsa) subgenus Putorius Steppe polecat (M. eversmannii) Ferret (M. furo) Black-footed ferret (M. nigripes) European polecat (M. putorius) See also: Mink Polecat

v t e Extinct Canidae Kingdom Animalia Phylum Chordata Class Mammalia Order: Carnivora Suborder: Caniformia Canidae Canidae †Prohesperocyon †Hesperocyoninae Hesperocyon Mesocyon Cynodesmus Sunkahetanka Philotrox Enhydrocyon Paraenhydrocyon Caedocyon Ectopocynus Osbornodon O. brachypus O. fricki O. iamonensis O. renjiei O. sesnoni O. scitulus O. wangi †Borophaginae Archaeocyon Oxetocyon Otarocyon Rhizocyon Phlaocyonini Cynarctoides Phlaocyon P. achoros P. annectens P. latidens P. leucosteus P. mariae P. marslandensis P. minor P. multicuspus P. taylori P. yatkolai Borophagini Cormocyon Desmocyon Metatomarctus Euoplocyon Psalidocyon Microtomarctus Protomarctus Tephrocyon Cynarctina Paracynarctus Cynarctus Aelurodontina Tomarctus Aelurodon Borophagina Paratomarctus Carpocyon Protepicyon Epicyon Borophagus B. diversidens B. dudleyi B. hilli B. littoralis B. orc B. parvus B. pugnator B. secundus Caninae see below↓ Caninae Caninae †Leptocyon †Metalopex Urocyon †U. citrinus †U. galushai †U. minicephalus †U. progressus †U. webbi Vulpini †Ferrucyon †Prototocyon Nyctereutes (raccoon dogs) †N. abdeslami †N. donnezani †N. lockwoodi †N. megamastoides †N. sinensis †N. tingi †N. vinetorum Vulpes (true foxes) †V. hassani †V. odessana †V. praeglacialis †V. qiuzhudingi †V. riffautae †V. rooki †V. skinneri †V. stenognathus Canini (true dogs) †Canis gezi †Nurocyon Cerdocyonina (zorro) †Protocyon †Theriodictis Speothos †Pleistocene bush dog (S. pacivorus) †Dusicyon Falkland Islands wolf (D. australis) D. avus D. cultridens Lycalopex (South American foxes) †L. cultridens (?) †L. ensenadensis Canina (wolf-like canids) see below↓ Canina (wolf-like canids) Canina †Dire wolf (Aenocyon) †Sardinian dhole (Cynotherium) †Trinil dog (Mececyon) †Merriam's dog (Megacyon) †Xenocyon †Eucyon E. davisi E. debonisi E. ferox E. intrepidus E. khoikhoi E. kuta E. marinae E. minor E. monticinensis E. skinneri E. wokari E. zhoui Lycaon †L. magnus †L. sekowei Dhole (Cuon alpinus) †C. a. antiquus †C. a. caucasicus †European dhole (C. a. europaeus) †Tien Shan dhole (C. a. hesperius) Canis †C. adoxus †C. antonii †C. apolloniensis †Armbruster's wolf (C. armbrusteri) †C. arnensis †C. borjgali †C. cedazoensis †C. chihliensis †Edward's wolf (C. edwardii) †Etruscan wolf (C. etruscus) †C. lepophagus †Mosbach wolf (C. mosbachensis) †C. nehringi †C. palmidens †C. variabilis Coyote (C. latrans) †Pleistocene coyote (C. l. orcutti) Red wolf (C. rufus) †Florida black wolf (C. r. floridanus) †Texas red wolf (C. r. rufus) Wolf (C. lupus) †Prehistoric C. l. bohemica C. l. lunellensis C. l. maximus C. l. santenaisiensis Cave wolf (C. l. spelaeus) Beringian wolf †Recently extinct Sicilian wolf (C. l. cristaldii) Hokkaido wolf (C. l. hattai) Japanese wolf (C. l. hodophilax) Austro-Hungarian wolf Synonymized under C. l. arctos: Bernard's wolf (C. l. bernardi) Synonymized under C. l. occidentalis: Kenai Peninsula wolf (C. l. alces) Manitoba wolf (C. l. griseoalbus) Synonymized under C. l. nubilus: Newfoundland wolf (C. l. beothucus) Cascade mountain wolf (C. l. fuscus) Mogollon mountain wolf (C. l. mogollonensis) Texas wolf (C. l. monstrabilis) Southern Rocky Mountain wolf (C. l. youngi) See also: Evolution of the wolf Category:Extinct canids

Taxon identifiers Cuon alpinus Wikidata: Q132585 Wikispecies: Cuon alpinus BioLib: 1882 BOLD: 73506 ECOS: 3075 EoL: 328688 GBIF: 2434317 iNaturalist: 42101 IRMNG: 10856319 ITIS: 183831 IUCN: 5953 MDD: 1005949 MSW: 14000793 NBN: NHMSYS0000376566 NCBI: 68730 Observation.org: 80328 Open Tree of Life: 313163 Paleobiology Database: 45383 Species+: 5289 TSA: 5499 Xeno-canto: Cuon-alpinus Canis alpinus Wikidata: Q109647079 Wikispecies: Canis alpinus CoL: QLX5 EUNIS: 8462 GBIF: 7705729 ITIS: 998399 ZooBank: 92207BB9-A501-4F50-99B3-61F22B885AE1

Authority control databases National United States Israel Other Yale LUX

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Adapted from the Wikipedia article [Dhole](https://en.wikipedia.org/wiki/Dhole) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Dhole?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.