Dhole
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The dhole (''Cuon alpinus''; ) is a
canid Canidae (; from Latin, '' canis'', " dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found withi ...
native to
Central Central is an adjective usually referring to being in the center of some place or (mathematical) object. Central may also refer to: Directions and generalised locations * Central Africa, a region in the centre of Africa continent, also known as ...
,
South South is one of the cardinal directions or Points of the compass, compass points. The direction is the opposite of north and is perpendicular to both east and west. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Pro ...
,
East East or Orient is one of the four cardinal directions or points of the compass. It is the opposite direction from west and is the direction from which the Sun rises on the Earth. Etymology As in other languages, the word is formed from the fa ...
and
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
. Other English names for the species include Asian wild dog, Asiatic wild dog, Indian wild dog, whistling dog, red dog, red wolf, and mountain wolf. It is genetically close to species within the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''
Canis ''Canis'' is a genus of the Caninae which includes multiple extant species, such as wolves, dogs, coyotes, and golden jackals. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and de ...
'', but distinct in several anatomical aspects: its
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
is convex rather than concave in profile, it lacks a third lower molar and the upper molars sport only a single
cusp A cusp is the most pointed end of a curve. It often refers to cusp (anatomy), a pointed structure on a tooth. Cusp or CUSP may also refer to: Mathematics * Cusp (singularity), a singular point of a curve * Cusp catastrophe, a branch of bifurc ...
as opposed to between two and four. During the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
, the dhole ranged throughout
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
,
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
but became restricted to its historical range 12,000–18,000 years ago. The dhole is a highly social animal, living in large clans without rigid dominance hierarchies and containing multiple breeding females. Such clans usually consist of about 12 individuals, but groups of over 40 are known. It is a diurnal pack hunter which preferentially targets large and medium-sized
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, cam ...
s. In tropical forests, the dhole competes with the
tiger The tiger (''Panthera tigris'') is the largest living cat species and a member of the genus '' Panthera''. It is most recognisable for its dark vertical stripes on orange fur with a white underside. An apex predator, it primarily preys on u ...
(''Panthera tigris'') and the
leopard The leopard (''Panthera pardus'') is one of the five extant species in the genus '' Panthera'', a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, a ...
(''Panthera pardus''), targeting somewhat different prey species, but still with substantial dietary overlap. It is listed as
Endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inva ...
on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
, 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 dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is Domestication of the dog, derived from the extinct Pleistocene wolf, and the modern wolf is the dog's n ...
s.


Etymology and naming

The
etymology Etymology ()The New Oxford Dictionary of English (1998) – p. 633 "Etymology /ˌɛtɪˈmɒlədʒi/ the study of the class in words and the way their meanings have changed throughout time". is the study of the history of the Phonological chan ...
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,
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
. He stated that ''dhole'' was a common local name for the species. In 1827,
Charles Hamilton Smith Lieutenant-Colonel Charles Hamilton Smith, KH, KW, FRS, FLS, (26 December 1776 in East Flanders, in the United Provinces of the Netherlands – 21 September 1859 in Plymouth) was an English artist, naturalist, antiquary, illustrator, soldie ...
claimed that it was derived from a language spoken in 'various parts of the East'. Two years later, Smith connected this word with tr,
deli Deli may refer to: * Delicatessen, a shop selling specially prepared food, or food prepared by such a shop * Sultanate of Deli, a former sultanate in North Sumatra, Indonesia Places * Deli, Boyer-Ahmad, a village in Kohgiluyeh and Boyer-Ahmad Pro ...
'mad, crazy', and erroneously compared the Turkish word with osx, dol and nl, dol (cfr. also en,
dull Dull may refer to: __NOTOC__ Places * Dull, Perth and Kinross, Scotland, United Kingdom, a village * Dull, Ohio, United States, an unincorporated community * Dull, Texas, United States, a ghost town People * Jack Dull (1930–1995), American prof ...
; german:
toll Toll may refer to: Transportation * Toll (fee) a fee charged for the use of a road or waterway ** Road pricing, the modern practice of charging for road use ** Road toll (historic), the historic practice of charging for road use ** Shadow toll, ...
), which are in fact from the
Proto-Germanic Proto-Germanic (abbreviated PGmc; also called Common Germanic) is the reconstructed proto-language of the Germanic branch of the Indo-European languages. Proto-Germanic eventually developed from pre-Proto-Germanic into three Germanic branc ...
*''dwalaz'' 'foolish, stupid'.
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker, ...
wrote nearly 80 years later that the word was not used by the natives living within the species' range. The
Merriam-Webster Merriam-Webster, Inc. is an American company that publishes reference books and is especially known for its dictionaries. It is the oldest dictionary publisher in the United States. In 1831, George and Charles Merriam founded the company as ...
''Dictionary'' theorises that it may have come from the kn, tōḷa ('wolf').


Taxonomy and evolution

''Canis alpinus'' was the
binomial name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
proposed by
Peter Simon Pallas Peter Simon Pallas Fellow of the Royal Society, FRS FRSE (22 September 1741 – 8 September 1811) was a Prussian zoologist and botanist who worked in Russia between 1767 and 1810. Life and work Peter Simon Pallas was born in Berlin, the son ...
in 1811, who described its range as encompassing the upper levels of Udskoi Ostrog in
Amurland Outer Manchuria (russian: Приаму́рье, translit=Priamurye; zh, s=外满洲, t=外滿洲, p=Wài Mǎnzhōu), or Outer Northeast China ( zh, s=外东北, t=外東北, p=Wài Dōngběi), refers to a territory in Northeast Asia that is now ...
, towards the eastern side and in the region of the upper
Lena River The Lena (russian: Ле́на, ; evn, Елюенэ, ''Eljune''; sah, Өлүөнэ, ''Ölüöne''; bua, Зүлхэ, ''Zülkhe''; mn, Зүлгэ, ''Zülge'') is the easternmost of the three great Siberian rivers that flow into the Arctic Ocean ...
, around the
Yenisei The Yenisey (russian: Енисе́й, ''Yeniséy''; mn, Горлог мөрөн, ''Gorlog mörön''; Buryat language, Buryat: Горлог мүрэн, ''Gorlog müren''; Tuvan language, Tuvan: Улуг-Хем, ''Uluğ-Hem''; Khakas language, K ...
River and occasionally crossing into
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
. This northern Russian range reported by Pallas during the 18th and 19th centuries is "considerably north" of where this species occurs today. ''Canis primaevus'' was a name proposed by
Brian Houghton Hodgson Brian Houghton Hodgson (1 February 1800 or more likely 1801 – 23 May 1894) was a pioneer naturalist and ethnologist working in India and Nepal where he was a British Resident. He described numerous species of birds and mammals from the Himala ...
in 1833 who thought that the dhole was a primitive ''Canis'' form and the
progenitor In genealogy, the progenitor (rarer: primogenitor; german: Stammvater or ''Ahnherr'') is the – sometimes legendary – founder of a family, line of descent, clan or tribe, noble house, or ethnic group.. Ebenda''Ahnherr:''"Stammvater eines G ...
of the
domestic dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is Domestication of the dog, derived from the extinct Pleistocene wolf, and the modern wolf is the dog's n ...
. Hodgson later took note of the dhole's physical distinctiveness from the genus ''Canis'' and proposed the genus ''Cuon''. 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. The paleontologist 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 Villafranchian age ( ) is a period of geologic time (3.5–1.0 Ma) spanning the Late Pliocene and Early Pleistocene used more specifically with European Land Mammal Ages. Named by Italian geologist Lorenzo Pareto for a sequence of terrestrial s ...
era
Valdarno The Valdarno is the valley of the river Arno, although this name does not apply to the entire river basin. Usage of the term generally excludes Casentino and the valleys formed by major tributaries. Some towns in the area: * Rignano sull'Arno * ...
, Italy and in China—was almost indistinguishable from the genus ''Canis''. In comparison, the modernspecies has greatly reduced molars and the
cusp A cusp is the most pointed end of a curve. It often refers to cusp (anatomy), a pointed structure on a tooth. Cusp or CUSP may also refer to: Mathematics * Cusp (singularity), a singular point of a curve * Cusp catastrophe, a branch of bifurc ...
s have developed into sharply trenchant points. During the Early
Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. The ...
there arose both ''Canis majori stehlini'' that was the size of a large
wolf The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly u ...
, and the early dhole ''Canis alpinus'' Pallas 1811 which first appeared at
Hundsheim Hundsheim is a town in the district of Bruck an der Leitha in Lower Austria, in northeast Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Easter ...
and Mosbach in Germany. In the
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...
era the
European dhole The European dhole (''Cuon alpinus europaeus'') was a paleosubspecies of the dhole, which ranged throughout much of Western and Central Europe during the Middle and Late Pleistocene. Like the modern Asiatic populations, it was a more progressi ...
(''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 The Würm is a river in Bavaria, Germany, right tributary of the Amper. The length of the river is , or including the ''Steinbach'', the main feed of Lake Starnberg. It drains the overflow from Lake Starnberg and flows swiftly through the villag ...
period, but the species as a whole still inhabits a large area of Asia. The European dhole may have survived up until the early
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...
in the
Iberian Peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, defi ...
. and what is believed to be dhole remains have been found at Riparo Fredian in northern
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...
dated 10,800 years old. The vast Pleistocene range of this species also included numerous islands in Asia that this species no longer inhabits, such as
Sri Lanka Sri Lanka (, ; si, ශ්‍රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங்கை, Ilaṅkai, translit-std=ISO ()), formerly known as Ceylon and officially the Democratic Socialist Republic of Sri Lanka, is an ...
,
Borneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and eas ...
and possibly
Palawan Palawan (), officially the Province of Palawan ( cyo, Probinsya i'ang Palawan; tl, Lalawigan ng Palawan), is an archipelagic province of the Philippines that is located in the region of Mimaropa. It is the largest province in the country in ...
in the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
. Middle Pleistocene dhole fossils have also been found in the Matsukae Cave in northern
Kyushu is the third-largest island of Japan's five main islands and the most southerly of the four largest islands ( i.e. excluding Okinawa). In the past, it has been known as , and . The historical regional name referred to Kyushu and its surroun ...
Island in western
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
and in the Lower Kuzuu fauna in
Tochigi Prefecture is a prefecture of Japan located in the Kantō region of Honshu. Tochigi Prefecture has a population of 1,943,886 (1 June 2019) and has a geographic area of 6,408 km2 (2,474 sq mi). Tochigi Prefecture borders Fukushima Prefecture to the ...
in
Honshu , historically called , is the largest and most populous island of Japan. It is located south of Hokkaidō across the Tsugaru Strait, north of Shikoku across the Inland Sea, and northeast of Kyūshū across the Kanmon Straits. The island separ ...
Island, east Japan. Dhole fossils from the Late Pleistocene dated to about 10,700 years before present are known from the Luobi Cave or Luobi-Dong cave in
Hainan Island Hainan (, ; ) is the smallest and southernmost province of the People's Republic of China (PRC), consisting of various islands in the South China Sea. , the largest and most populous island in China,The island of Taiwan, which is slightly ...
in south China where they no longer exist. Additionally, fossils of canidae possibly belonging to dhole have been excavated from
Dajia River Dajia River () is the fifth-longest river in Taiwan located in the north-central of the island. It flows through Taichung City for 142 km. The sources of the Dajia are: Hsuehshan and Nanhu Mountain in the Central Mountain Range. The Dajia R ...
in
Taichung County Taichung County was a county in central Taiwan between 1945 and 2010. The county seat was in Yuanlin Township before 1950 and Fongyuan City after 1950. History Taichung County was established on 26 November 1945 on the territory of Taichū P ...
, Taiwan. The
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
indicates that the species also occurred in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, with remains being found in
Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72 degrees north latitude in the Chukchi Sea; and on the south by the tip ...
and
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
. In 2021, the analyses of the mitochondrial
genomes In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding gen ...
extracted from the fossil remains of two extinct European dhole specimens from the Jáchymka cave,
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
dated 35,000–45,000 years old indicate that these were genetically basal to modern dholes and possessed much greater genetic diversity. The dhole's distinctive morphology has been a source of much confusion in determining the species' systematic position among the Canidae. George Simpson placed the dhole in the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Simocyoninae alongside the
African wild dog The African wild dog (''Lycaon pictus''), also called the painted dog or Cape hunting dog, is a wild canine which is a native species to sub-Saharan Africa. It is the largest wild canine in Africa, and the only extant member of the genus '' Ly ...
and the
bush dog The bush dog (''Speothos venaticus'') is a canine found in Central and South America. In spite of its extensive range, it is very rare in most areas except in Suriname, Guyana and Peru; it was first identified by Peter Wilhelm Lund from fossi ...
, on account of all three species' similar dentition. Subsequent authors, including
Juliet Clutton-Brock Juliet Clutton-Brock, FSA, FZS (6 September 1933 – 21 September 2015) was an English zooarchaeologist and curator, specialising in domesticated mammals. From 1969 to 1993, she worked at the Natural History Museum. Between 1999 and 2006, she w ...
, noted greater morphological similarities to canids of the genera ''Canis'', ''
Dusicyon ''Dusicyon'' is an extinct genus of South American canids. The type species is ''Dusicyon australis'', the Falkland Islands wolf. In 1914, Oldfield Thomas established this genus, in which he included the culpeo and other South American foxes. ...
'' and ''
Alopex Alopex may refer to: * ''Alopex lagopus'', a taxonomic synonym for the Arctic fox, ''Vulpes lagopus'' * ALOPEX a correlation-based machine learning algorithm * Alopex (Teenage Mutant Ninja Turtles), a character in the ''Teenage Mutant Ninja Turt ...
'' than to either ''
Speothos ''Speothos'' is a genus of canid found in Central and South America. The genus includes the living bush dog, '' Speothos venaticus'', and an extinct Pleistocene species, '' Speothos pacivorus''. Unusually, the fossil species was identified and ...
'' or '' Lycaon'', with any resemblance to the latter two being due to
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
. Some authors consider the extinct ''Canis''
subgenus In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
''
Xenocyon ''Xenocyon'' ("strange dog") is an extinct subgenus of ''Canis''. The group includes ''Canis'' (''Xenocyon'') ''africanus'', ''Canis'' (''Xenocyon'') ''antonii'' and ''Canis'' (''Xenocyon'') ''falconeri'' that gave rise to ''Canis'' (''Xenocyon'' ...
'' as ancestral to both the genus ''Lycaon'' and the genus ''Cuon''. Subsequent studies on the canid
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
revealed that the dhole and African wild dog are closely related to members of the genus ''Canis''. This closeness to ''Canis'' may have been confirmed in a menagerie in
Madras Chennai (, ), formerly known as Madras ( the official name until 1996), is the capital city of Tamil Nadu, the southernmost Indian state. The largest city of the state in area and population, Chennai is located on the Coromandel Coast of th ...
, where according to zoologist
Reginald Innes Pocock Reginald Innes Pocock F.R.S. (4 March 1863 – 9 August 1947) was a British zoologist. Pocock was born in Clifton, Bristol, the fourth son of Rev. Nicholas Pocock and Edith Prichard. He began showing interest in natural history at St. Edward ...
there is a record of a dhole that interbred with a golden jackal.


Admixture with the African wild dog

In 2018,
whole genome sequencing Whole genome sequencing (WGS), also known as full genome sequencing, complete genome sequencing, or entire genome sequencing, is the process of determining the entirety, or nearly the entirety, of the DNA sequence of an organism's genome at a s ...
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 The African wild dog (''Lycaon pictus''), also called the painted dog or Cape hunting dog, is a wild canine which is a native species to sub-Saharan Africa. It is the largest wild canine in Africa, and the only extant member of the genus '' Ly ...
(''Lycaon pictus''). There was strong evidence of ancient
genetic admixture Genetic admixture occurs when previously diverged or isolated genetic lineages mix.⅝ Admixture results in the introduction of new genetic lineages into a population. Examples Climatic cycles facilitate genetic admixture in cold periods and gene ...
between the dhole and the African wild dog. Today, their ranges are remote from each other; however, during the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
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 The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
, from where it may have admixed with the African wild dog in
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
. However, there is no evidence of the dhole having existed in the Middle East nor North Africa.


Subspecies

Historically, up to ten subspecies of dholes have been recognised. , seven subspecies are recognised. However, studies on the dhole's
mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
and
microsatellite A microsatellite is a tract of repetitive DNA in which certain DNA motifs (ranging in length from one to six or more base pairs) are repeated, typically 5–50 times. Microsatellites occur at thousands of locations within an organism's genome. ...
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. 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, , 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 Canidae (; from Latin, ''canis'', "dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found within ...
of the
International Union for the Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natura ...
(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 The Tian Shan,, , otk, 𐰴𐰣 𐱅𐰭𐰼𐰃, , tr, Tanrı Dağı, mn, Тэнгэр уул, , ug, تەڭرىتاغ, , , kk, Тәңіртауы / Алатау, , , ky, Теңир-Тоо / Ала-Тоо, , , uz, Tyan-Shan / Tangritog‘ ...
subspecies has distinct morphology. 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. Fossils of dhole from the early
Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. The ...
have been found in
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's List ...
.


Characteristics

In appearance, the dhole has been variously described as combining the physical characteristics of the gray wolf and the
red fox The red fox (''Vulpes vulpes'') is the largest of the true foxes and one of the most widely distributed members of the Order (biology), order Carnivora, being present across the entire Northern Hemisphere including most of North America, Europe ...
, and as being "cat-like" on account of its long backbone and slender limbs. It has a wide and massive skull with a well-developed sagittal crest, and its
masseter muscle In human anatomy, the masseter is one of the muscles of mastication. Found only in mammals, it is particularly powerful in herbivores to facilitate chewing of plant matter. The most obvious muscle of mastication is the masseter muscle, since it ...
s are highly developed compared to other canid species, giving the face an almost hyena-like appearance. The
rostrum Rostrum may refer to: * Any kind of a platform for a speaker: **dais **pulpit * Rostrum (anatomy), a beak, or anatomical structure resembling a beak, as in the mouthparts of many sucking insects * Rostrum (ship), a form of bow on naval ships * Ros ...
is shorter than that of domestic dogs and most other canids. The species has six rather than seven lower
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
. The upper molars are weak, being one third to one half the size of those of wolves and have only one
cusp A cusp is the most pointed end of a curve. It often refers to cusp (anatomy), a pointed structure on a tooth. Cusp or CUSP may also refer to: Mathematics * Cusp (singularity), a singular point of a curve * Cusp catastrophe, a branch of bifurc ...
as opposed to between two and four, as is usual in canids, an adaptation thought to improve shearing ability, thus allowing it to compete more successfully with
kleptoparasite Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when fo ...
s. Adult females can weigh from , while the slightly larger male may weigh from . The mean weight of adults from three small samples was . Occasionally, dholes may be sympatric with the
Indian wolf The Indian wolf (''Canis lupus pallipes'') is a subspecies of gray wolf that ranges from Southwest Asia to the Indian Subcontinent. It is intermediate in size between the Himalayan wolf and the Arabian wolf, and lacks the former's luxuriant wint ...
(''Canis lupus pallipes''), which is one of the smallest races of the gray wolf, but is still approximately 25% heavier on average. The general tone of the fur is reddish, with the brightest hues occurring in winter. In the winter coat, the back is clothed in a saturated rusty-red to reddish colour with brownish highlights along the top of the head, neck and shoulders. The throat, chest, flanks, and belly and the upper parts of the limbs are less brightly coloured, and are more yellowish in tone. The lower parts of the limbs are whitish, with dark brownish bands on the anterior sides of the forelimbs. The muzzle and forehead are greyish-reddish. The tail is very luxuriant and fluffy, and is mainly of a reddish-ocherous colour, with a dark brown tip. The summer coat is shorter, coarser and darker. The dorsal and lateral
guard hair Guard hair or overhair is the outer layer of hair of most mammals, which overlay the fur. Guard hairs are long and coarse and protect the rest of the pelage (fur) from abrasion and frequently from moisture. They are visible on the surface of the fu ...
s in adults measure in length. Dholes in the
Moscow Zoo The Moscow Zoo or Moskovsky Zoopark (russian: Московский зоопарк) is a zoo founded in 1864 by professor-biologists, K.F. Rulje, S.A. Usov and A.P. Bogdanov, from the Moscow State University. In 1919, the zoo was nationalized. In ...
moult once a year from March to May. A melanistic individual was recorded in the northern
Coimbatore Coimbatore, also spelt as Koyamputhur (), sometimes shortened as Kovai (), is one of the major metropolitan cities in the Indian state of Tamil Nadu. It is located on the banks of the Noyyal River and surrounded by the Western Ghats. Coimbato ...
Forest Division in
Tamil Nadu Tamil Nadu (; , TN) is a States and union territories of India, state in southern India. It is the List of states and union territories of India by area, tenth largest Indian state by area and the List of states and union territories of India ...
.


Distribution and habitat

The dhole can be found in
Tibet Tibet (; ''Böd''; ) is a region in East Asia, covering much of the Tibetan Plateau and spanning about . It is the traditional homeland of the Tibetan people. Also resident on the plateau are some other ethnic groups such as Monpa people, ...
and possibly also in
North Korea North Korea, officially the Democratic People's Republic of Korea (DPRK), is a country in East Asia. It constitutes the northern half of the Korea, Korean Peninsula and shares borders with China and Russia to the north, at the Yalu River, Y ...
and
Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's List of countries and dependencies by population, fifth-most populous country, with a population of almost 24 ...
. It once inhabited the alpine steppes extending into
Kashmir Kashmir () is the northernmost geographical region of the Indian subcontinent. Until the mid-19th century, the term "Kashmir" denoted only the Kashmir Valley between the Great Himalayas and the Pir Panjal Range. Today, the term encompas ...
to the
Ladakh Ladakh () is a region administered by India as a union territory which constitutes a part of the larger Kashmir region and has been the subject of dispute between India, Pakistan, and China since 1947. (subscription required) Quote: "Jammu and ...
area. In Central Asia, the dhole primarily inhabits mountainous areas; in the western part of its range, it lives mostly in alpine meadows and high-montane steppes, while in the east, it mainly ranges in montane
taiga Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruce ...
s, and is sometimes sighted along
coast The coast, also known as the coastline or seashore, is defined as the area where land meets the ocean, or as a line that forms the boundary between the land and the coastline. The Earth has around of coastline. Coasts are important zones in n ...
lines. In India, Myanmar, Indochina, Indonesia and China, it prefers forested areas in
alpine zone Alpine tundra is a type of natural region or biome that does not contain trees because it is at high elevation, with an associated alpine climate, harsh climate. As the latitude of a location approaches the poles, the threshold elevation for alp ...
s and is occasionally sighted in
plain In geography, a plain is a flat expanse of land that generally does not change much in elevation, and is primarily treeless. Plains occur as lowlands along valleys or at the base of mountains, as coastal plains, and as plateaus or uplands ...
s regions. In the
Pamir Mountains The Pamir Mountains are a mountain range between Central Asia and Pakistan. It is located at a junction with other notable mountains, namely the Tian Shan, Karakoram, Kunlun, Hindu Kush and the Himalaya mountain ranges. They are among the world ...
of southern
Kyrgyzstan Kyrgyzstan,, pronounced or the Kyrgyz Republic, is a landlocked country in Central Asia. Kyrgyzstan is bordered by Kazakhstan to the north, Uzbekistan to the west, Tajikistan to the south, and the People's Republic of China to the east. ...
, the presence of the dhole was confirmed in 2019. The dhole might still be present in the
Tunkinsky National Park The Tunka or Tunkinsky National Park (Russian: Тункинский) is a national park located in south central Siberia. It covers a mountainous region centered on the Irkut River valley (also referred to as the Tunka Valley) that continues from ...
in extreme southern
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part of ...
near
Lake Baikal Lake Baikal (, russian: Oзеро Байкал, Ozero Baykal ); mn, Байгал нуур, Baigal nuur) is a rift lake in Russia. It is situated in southern Siberia, between the federal subjects of Irkutsk Oblast to the northwest and the Repu ...
. It possibly still lives in the
Primorsky Krai Primorsky Krai (russian: Приморский край, r=Primorsky kray, p=prʲɪˈmorskʲɪj kraj), informally known as Primorye (, ), is a federal subject (a krai) of Russia, located in the Far East region of the country and is a part of the ...
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. Currently, no other recent reports are confirmed of dhole being present in
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
. However, the dhole might be present in the eastern
Sayan Mountains The Sayan Mountains (russian: Саяны ''Sajany''; mn, Соёны нуруу, ''Soyonï nurû''; otk, 𐰚𐰇𐰏𐰢𐰤, Kögmen) are a mountain range in southern Siberia, Russia ( Buryatia, Irkutsk Oblast, Krasnoyarsk Krai, Tuva Republic ...
and in the
TransBaikal Transbaikal, Trans-Baikal, Transbaikalia ( rus, Забайка́лье, r=Zabaykalye, p=zəbɐjˈkalʲjɪ), or Dauria (, ''Dauriya'') is a mountainous region to the east of or "beyond" (trans-) Lake Baikal in Far Eastern Russia. The steppe and ...
region; it has been sighted in Tofalaria in the
Irkutsk Oblast Irkutsk Oblast (russian: Ирку́тская о́бласть, Irkutskaya oblast; bua, Эрхүү можо, Erkhüü mojo) is a federal subject of Russia (an oblast), located in southeastern Siberia in the basins of the Angara, Lena, and Nizh ...
, the
Republic of Buryatia Buryatia, officially the Republic of Buryatia (russian: Республика Бурятия, r=Respublika Buryatiya, p=rʲɪsˈpublʲɪkə bʊˈrʲætʲɪjə; bua, Буряад Улас, Buryaad Ulas, , mn, Буриад Улс, Buriad Uls), is ...
and
Zabaykalsky Krai Zabaykalsky Krai ( rus, Забайкальский край, r=Zabaikal'skii krai, p=zəbɐjˈkalʲskʲɪj kraj, lit. "Transbaikal krai"; bua, Yбэр Байгалай хизаар, Uber Baigalai Xizaar) is a federal subject of Russia (a krai) t ...
. One pack was sighted in the
Qilian Mountains The Qilian Mountains (, also romanized as Tsilien; Mongghul: Chileb), together with the Altyn-Tagh (Altun Shan) also known as Nan Shan (, literally "Southern Mountains"), as it is to the south of Hexi Corridor, is a northern outlier of the Kunlu ...
in 2006. In 2011 to 2013, local government officials and herders reported the presence of several dhole packs at elevations of near
Taxkorgan Nature Reserve The Taxkorgan Nature Reserve (officially spelled Taxkorgan Natural Reserve) is a nature reserve in Kashgar Prefecture, Xinjiang, China. It is situated around the Taghdumbash Pamir of Pamir Mountains and Karakorum Mountains. It covers about wa ...
in the
Xinjiang Xinjiang, SASM/GNC: ''Xinjang''; zh, c=, p=Xīnjiāng; formerly romanized as Sinkiang (, ), officially the Xinjiang Uygur Autonomous Region (XUAR), is an autonomous region of the People's Republic of China (PRC), located in the northwest ...
Autonomous Region. Several packs and a female adult with pups were also recorded by
camera trap A camera trap is a camera that is automatically triggered by a change in some activity in its vicinity, like presence of an animal or a human being. It is typically equipped with a motion sensor – usually a passive infrared (PIR) sensor ...
s at elevations of around in Yanchiwan National Nature Reserve in the northern
Gansu Province Gansu (, ; alternately romanized as Kansu) is a province in Northwest China. Its capital and largest city is Lanzhou, in the southeast part of the province. The seventh-largest administrative district by area at , Gansu lies between the Tibeta ...
in 2013–2014. Dholes have been also reported in the
Altyn-Tagh Altyn-Tagh (also Altun Mountains, Altun Shan; , Pinyin: ''A'erjin Shan'', Wade–Giles: ''A-erh-chin Shan;'' Uyghur'':ئالتۇن تاغ'') is a mountain range in Northwestern China that separates the Eastern Tarim Basin from the Tibetan P ...
Mountains. In China's
Yunnan Yunnan , () is a landlocked Provinces of China, province in Southwest China, the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is ...
Province, dholes were recorded in
Baima Xueshan Nature Reserve Baima may refer to: * Baima, Sierra Leone town in Kenema District, Eastern Province, Sierra Leone China Baima ( unless otherwise noted) may refer to: * Baima people, subgroup of Tibetans * Baima language, language spoken by 10,000 people of Tibe ...
in 2010–2011. Dhole samples were obtained in
Jiangxi Jiangxi (; ; formerly romanized as Kiangsi or Chianghsi) is a landlocked province in the east of the People's Republic of China. Its major cities include Nanchang and Jiujiang. Spanning from the banks of the Yangtze river in the north int ...
Province in 2013. Confirmed records by camera-trapping since 2008 have occurred in southern and western
Gansu Gansu (, ; alternately romanized as Kansu) is a province in Northwest China. Its capital and largest city is Lanzhou, in the southeast part of the province. The seventh-largest administrative district by area at , Gansu lies between the Tibet ...
province, southern
Shaanxi Shaanxi (alternatively Shensi, see #Name, § Name) is a landlocked Provinces of China, province of China. Officially part of Northwest China, it borders the province-level divisions of Shanxi (NE, E), Henan (E), Hubei (SE), Chongqing (S), Sichu ...
province, southern
Qinghai Qinghai (; alternately romanized as Tsinghai, Ch'inghai), also known as Kokonor, is a landlocked province in the northwest of the People's Republic of China. It is the fourth largest province of China by area and has the third smallest po ...
province, southern and western
Yunnan Yunnan , () is a landlocked Provinces of China, province in Southwest China, the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is ...
province, western
Sichuan Sichuan (; zh, c=, labels=no, ; zh, p=Sìchuān; alternatively romanized as Szechuan or Szechwan; formerly also referred to as "West China" or "Western China" by Protestant missions) is a province in Southwest China occupying most of the ...
province, the southern
Xinjiang Xinjiang, SASM/GNC: ''Xinjang''; zh, c=, p=Xīnjiāng; formerly romanized as Sinkiang (, ), officially the Xinjiang Uygur Autonomous Region (XUAR), is an autonomous region of the People's Republic of China (PRC), located in the northwest ...
Autonomous Region and in the Southeastern
Tibet Tibet (; ''Böd''; ) is a region in East Asia, covering much of the Tibetan Plateau and spanning about . It is the traditional homeland of the Tibetan people. Also resident on the plateau are some other ethnic groups such as Monpa people, ...
Autonomoous Region. 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. The dhole occurs in most of India south of the Ganges, particularly in the Central Indian Highlands and the
Western Western may refer to: Places *Western, Nebraska, a village in the US *Western, New York, a town in the US *Western Creek, Tasmania, a locality in Australia *Western Junction, Tasmania, a locality in Australia *Western world, countries that id ...
and
Eastern Ghats The Eastern Ghats are a discontinuous range of mountains along India's eastern coast. The Eastern Ghats pass through Odisha, Andhra Pradesh to Tamil Nadu in the south passing some parts of Karnataka as well as Telangana. They are eroded and cut ...
. It is also present in
Arunachal Pradesh Arunachal Pradesh (, ) is a state in Northeastern India. It was formed from the erstwhile North-East Frontier Agency (NEFA) region, and became a state on 20 February 1987. It borders the states of Assam and Nagaland to the south. It shares int ...
,
Assam Assam (; ) is a state in northeastern India, south of the eastern Himalayas along the Brahmaputra and Barak River valleys. Assam covers an area of . The state is bordered by Bhutan and Arunachal Pradesh to the north; Nagaland and Manipur ...
,
Meghalaya Meghalaya (, or , meaning "abode of clouds"; from Sanskrit , "cloud" + , "abode") is a states and union territories of India, state in northeastern India. Meghalaya was formed on 21 January 1972 by carving out two districts from the state of As ...
and
West Bengal West Bengal (, Bengali: ''Poshchim Bongo'', , abbr. WB) is a state in the eastern portion of India. It is situated along the Bay of Bengal, along with a population of over 91 million inhabitants within an area of . West Bengal is the fourt ...
and in the
Indo-Gangetic Plain The Indo-Gangetic Plain, also known as the North Indian River Plain, is a fertile plain encompassing northern regions of the Indian subcontinent, including most of northern and eastern India, around half of Pakistan, virtually all of Bangla ...
's
Terai The Terai or Tarai is a lowland region in northern India and southern Nepal that lies south of the outer foothills of the Himalayas, the Sivalik Hills, and north of the Indo-Gangetic Plain. This lowland belt is characterised by tall grasslands, scr ...
region. Dhole populations in the
Himalaya The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 100 ...
s and northwest India are fragmented. In 2011, dhole packs were recorded by camera traps in the
Chitwan National Park , iucn_category = II , location = Central Terai of Nepal , established = 1973 , nearest_city = Bharatpur , map = Nepal Bagmati Province#Nepal#India#South Asia , relief = 1 , label = Chitwan National Park , label_position = top , coordina ...
. Its presence was confirmed in the
Kanchenjunga Conservation Area Kanchenjunga Conservation Area is a protected area in the Himalayas of eastern Nepal that was established in 1997. It covers in the Taplejung District and comprises two peaks of Kanchenjunga. In the north it adjoins the Qomolangma National Natur ...
in 2011 by camera traps. In February 2020, dholes were sighted in the
Vansda National Park Vansda National Park, also known as Bansda National Park, is a protected area which represents the thick woodlands of the Dangs and southern Gujarat, and is situated in the Vansda tehsil, Navsari District of Gujarat state, India. Riding on the ...
, 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 Gujarat (, ) is a state along the western coast of India. Its coastline of about is the longest in the country, most of which lies on the Kathiawar peninsula. Gujarat is the fifth-largest Indian state by area, covering some ; and the ninth ...
since 1970. In
Bhutan Bhutan (; dz, འབྲུག་ཡུལ་, Druk Yul ), officially the Kingdom of Bhutan,), is a landlocked country in South Asia. It is situated in the Eastern Himalayas, between China in the north and India in the south. A mountainous ...
, the dhole is present in
Jigme Dorji National Park The Jigme Dorji National Park (JDNP), named after the late Jigme Dorji Wangchuck, is the second-largest National Park of Bhutan. It occupies almost the entire Gasa District, as well as the northern areas of Thimphu District, Paro District, Puna ...
. In
Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mos ...
, it inhabits forest reserves in the
Sylhet Sylhet ( bn, সিলেট) is a metropolitan city in northeastern Bangladesh. It is the administrative seat of the Sylhet Division. Located on the north bank of the Surma River at the eastern tip of Bengal, Sylhet has a subtropical climate an ...
area, as well the
Chittagong Hill Tracts The Chittagong Hill Tracts ( bn, পার্বত্য চট্টগ্রাম, Parbotto Chottogram), often shortened to simply the Hill Tracts and abbreviated to CHT, are group of districts within the Chittagong Division in southeast ...
in the southeast. Recent camera trap photos in the Chittagong in 2016 showed the continued presence of the dhole. These regions probably do not harbour a viable population, as mostly small groups or solitary individuals were sighted. In
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
, the dhole is present in several protected areas. In 2015, dholes and tigers were recorded by camera-traps for the first time in the hill forests of
Karen State Kayin State ( my, ကရင်ပြည်နယ်, ; kjp, ဖၠုံခါန်ႋကၞင့်, italics=no; ksw, ကညီကီၢ်စဲၣ်, ), also known by the endonyms Kawthoolei and Karen State, is a state of Myanmar. The ca ...
. Its range is highly fragmented in the
Malaysian Peninsula The Malay Peninsula ( Malay: ''Semenanjung Tanah Melayu'') is a peninsula in Mainland Southeast Asia. The landmass runs approximately north–south, and at its terminus, it is the southernmost point of the Asian continental mainland. The area ...
,
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
,
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's List ...
,
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
and
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
. In 2014, camera trap videos in the montane tropical forests at in the
Kerinci Seblat National Park Kerinci Seblat National Park is the largest national park on the island of Sumatra, Indonesia. It has a total area of 13,791 km2, and spans four provinces: West Sumatra, Jambi, Bengkulu, and South Sumatra. Geography It is located between 1 ...
in Sumatra revealed its continued presence. A camera trapping survey in the
Khao Ang Rue Nai Wildlife Sanctuary The Khao Ang Rue Nai Wildlife Sanctuary ( th, เขตรักษาพันธุ์สัตว์ป่าเขาอ่างฤๅไน) is a protected area at the western extremities of the Cardamom Mountains in Chachoengsao Province, T ...
in Thailand from January 2008 to February 2010 documented one healthy dhole pack. In northern
Laos Laos (, ''Lāo'' )), officially the Lao People's Democratic Republic ( Lao: ສາທາລະນະລັດ ປະຊາທິປະໄຕ ປະຊາຊົນລາວ, French: République démocratique populaire lao), is a socialist ...
, dholes were studied in Nam Et-Phou Louey National Protected Area. Camera trap surveys from 2012 to 2017 recorded dholes in the same Nam Et-Phou Louey National Protected Area. In
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
, dholes were sighted only in Pu Mat National Park in 1999, in
Yok Don National Park ट or YOK may refer to: *Yok weaving technique, see Thai-style dresses, Tstyle dresses *Yok, a Turkish copula, negation of "to be" *YOK or YÖK, an abbreviation for the Commission for Higher Education, Turkey *yok, the ISO 639-3 code for the Yoku ...
in 2003 and 2004; and in Ninh Thuan Province in 2014. In 2019, scat samples collected in the Bek-Tosot Conservancy in
Kyrgyzstan Kyrgyzstan,, pronounced or the Kyrgyz Republic, is a landlocked country in Central Asia. Kyrgyzstan is bordered by Kazakhstan to the north, Uzbekistan to the west, Tajikistan to the south, and the People's Republic of China to the east. ...
confirmed the continued presence of dholes in the area. This was the first record of dholes from the country in almost three decades. A disjunct dhole population was reported in the area of
Trabzon Trabzon (; Ancient Greek: Tραπεζοῦς (''Trapezous''), Ophitic Pontic Greek: Τραπεζούντα (''Trapezounta''); Georgian: ტრაპიზონი (''Trapizoni'')), historically known as Trebizond in English, is a city on the B ...
and
Rize Rize (Greek language, Greek: ρίζα, Laz language, Laz: რიზინი, Georgian language, Georgian: რიზე, , Ottoman Turkish: ريزه) is the capital city of Rize Province in the eastern part of the Black Sea Region of Turkey. Rize ...
in northeastern
Turkey Turkey ( tr, Türkiye ), officially the Republic of Türkiye ( tr, Türkiye Cumhuriyeti, links=no ), is a list of transcontinental countries, transcontinental country located mainly on the Anatolia, Anatolian Peninsula in Western Asia, with ...
near the border with
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
in the 1990s. This report was not considered to be reliable. One single individual was claimed to have been shot in 2013 in the nearby
Kabardino-Balkaria The Kabardino-Balkarian Republic (russian: Кабарди́но-Балка́рская Респу́блика, ''Kabardino-Balkarskaya Respublika''; kbd, Къэбэрдей-Балъкъэр Республикэ, ''Ķêbêrdej-Baĺķêr Respublik ...
Republic in the central
Caucasus The Caucasus () or Caucasia (), is a region between the Black Sea and the Caspian Sea, mainly comprising Armenia, Azerbaijan, Georgia, and parts of Southern Russia. The Caucasus Mountains, including the Greater Caucasus range, have historically ...
; its remains were analysed in May 2015 by a biologist from the
Kabardino-Balkarian State University Kabardino-Balkarian State University (KBSU); russian: Кабардино-Балкарский государственный университет имени Х. М. Бербекова ''Kabardino_Balkarskii gosudarstvennii universitet imeni H. ...
, who concluded that the skull was indeed that of a dhole. In August 2015, researchers from the
National Museum of Natural History The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. In 2021, with 7 ...
and the
Karadeniz Technical University Karadeniz Technical University ( tr, Karadeniz Teknik Üniversitesi or KTÜ) is a public research university in Trabzon, in the Black Sea Region of Turkey. Established in 1955, it is the fourth oldest university in Turkey. Karadeniz Technical U ...
started an expedition to track and document this possible Turkish population of dhole. In October 2015, they concluded that no real evidence exists of a living dhole population in Turkey or in the Kabardino-Balkaria Republic, pending DNA analysis of samples from the original 1994 skins.


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. Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries. In contrast to wolves, dholes do not
howl Howl most often refers to: *Howling, an animal vocalization in many canine species *Howl (poem), a 1956 poem by Allen Ginsberg Howl may also refer to: Film * ''The Howl'', a 1970 Italian film * ''Howl'' (2010 film), a 2010 American arthouse b ...
or bark. Dholes have a complex
body language Body language is a type of communication in which physical behaviors, as opposed to words, are used to express or convey information. Such behavior includes facial expressions, body posture, gestures, eye movement, touch and the use of space. Th ...
. 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.


Social and territorial behaviour

Dholes are more social than
gray wolves The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly un ...
, and have less of a dominance hierarchy, as seasonal scarcity of food is not a serious concern for them. In this manner, they closely resemble African wild dogs in social structure. They live in
clan A clan is a group of people united by actual or perceived kinship and descent. Even if lineage details are unknown, clans may claim descent from founding member or apical ancestor. Clans, in indigenous societies, tend to be endogamous, meaning ...
s 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 three to five animals, particularly during the spring season, as this is the optimal number for catching fawns. Dominant dholes are hard to identify, as they do not engage in dominance displays as wolves do, though other clan members will show submissive behaviour toward them. Intragroup fighting is rarely observed. Dholes are far less
territorial A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or a ...
than wolves, with pups from one clan often joining another without trouble once they mature sexually. Clans typically number 5 to 12 individuals in India, though clans of 40 have been reported. In
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
, clans rarely exceed three individuals. Unlike other canids, there is no evidence of dholes using
urine Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excretion, excreted from the body through the urethra. Cel ...
to mark their territories or travel routes. When urinating, dholes, especially males, may raise one hind leg or both to result in a handstand. Handstand urination is also seen in
bush dog The bush dog (''Speothos venaticus'') is a canine found in Central and South America. In spite of its extensive range, it is very rare in most areas except in Suriname, Guyana and Peru; it was first identified by Peter Wilhelm Lund from fossi ...
s (''Speothos venaticus''). They may defecate in conspicuous places, though a territorial function is unlikely, as
faeces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
are mostly deposited within the clan's territory rather than the periphery. Faeces are often deposited in what appear to be communal
latrines A latrine is a toilet or an even simpler facility that is used as a toilet within a sanitation system. For example, it can be a communal trench in the earth in a camp to be used as emergency sanitation, a hole in the ground (pit latrine), or m ...
. They do not scrape the earth with their feet, as other canids do, to mark their territories.


Denning

Four kinds of den have been described; simple earth dens with one entrance (usually remodeled striped hyena or
porcupine Porcupines are large rodents with coats of sharp spines, or quills, that protect them against predation. The term covers two families of animals: the Old World porcupines of family Hystricidae, and the New World porcupines of family, Erethizont ...
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 of interconnecting tunnels. These "cities" may be developed over many generations of dholes, and are shared by the clan females when raising young together. Like
African wild dog The African wild dog (''Lycaon pictus''), also called the painted dog or Cape hunting dog, is a wild canine which is a native species to sub-Saharan Africa. It is the largest wild canine in Africa, and the only extant member of the genus '' Ly ...
s and
dingo The dingo (''Canis familiaris'', ''Canis familiaris dingo'', ''Canis dingo'', or ''Canis lupus dingo'') is an ancient (Basal (phylogenetics), basal) lineage of dog found in Australia (continent), Australia. Its taxonomic classification is de ...
es, dholes will avoid killing prey close to their dens.


Reproduction and development

In India, the
mating season Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and ch ...
occurs between mid-October and January, while captive dholes in the
Moscow Zoo The Moscow Zoo or Moskovsky Zoopark (russian: Московский зоопарк) is a zoo founded in 1864 by professor-biologists, K.F. Rulje, S.A. Usov and A.P. Bogdanov, from the Moscow State University. In 1919, the zoo was nationalized. In ...
breed mostly in February. Unlike wolf packs, dhole clans may contain more than one breeding female. More than one female dhole may den and rear their litters together in the same den. During
mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reproduc ...
, the female assumes a crouched, cat-like position. There is no
copulatory tie Canine reproduction is the process of sexual reproduction in domestic dogs, wolves, coyotes and other canine species. Canine sexual anatomy and development Male reproductive system Erectile tissue As with all mammals, a dog's penis is made up o ...
characteristic of other canids when the male dismounts. Instead, the pair lie on their sides facing each other in a semicircular formation. The
gestation period In mammals, pregnancy is the period of reproduction during which a female carries one or more live offspring from implantation in the uterus through gestation. It begins when a fertilized zygote implants in the female's uterus, and ends once it ...
lasts 60–63 days, with litter sizes averaging four to six pups. Their growth rate is much faster than that of wolves, being similar in rate to that of
coyote The coyote (''Canis latrans'') is a species of canis, canine native to North America. It is smaller than its close relative, the wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecologica ...
s. The
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...
metabolites of five males and three females kept in Thai zoos was studied. The breeding males showed an increased level of
testosterone Testosterone is the primary sex hormone and anabolic steroid in males. In humans, testosterone plays a key role in the development of Male reproductive system, male reproductive tissues such as testes and prostate, as well as promoting secondar ...
from October to January. The
oestrogen Estrogen or oestrogen is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three major endogenous estrogens that have estrogenic hormonal acti ...
level of captive females increases for about two weeks in January, followed by an increase of
progesterone Progesterone (P4) is an endogenous steroid and progestogen sex hormone involved in the menstrual cycle, pregnancy, and embryogenesis of humans and other species. It belongs to a group of steroid hormones called the progestogens and is the m ...
. They displayed sexual behaviours during the oestrogen peak of the females. 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 Weaning is the process of gradually introducing an infant human or another mammal to what will be its adult diet while withdrawing the supply of its mother's milk. The process takes place only in mammals, as only mammals produce milk. The infan ...
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 by the age of eight months. Maximum longevity in captivity is 15–16 years.


Hunting behaviour

Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and homo- and heterosexual mounting. Dholes are primarily diurnal hunters, hunting in the early hours of the morning. They rarely hunt
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
ly, except on moonlit nights, indicating they greatly rely on
sight Visual perception is the ability to interpret the surrounding Biophysical environment, environment through photopic vision (daytime vision), color vision, scotopic vision (night vision), and mesopic vision (twilight vision), using light in the ...
when hunting. Although not as fast as jackals and foxes, they can chase their prey for many hours. During a pursuit, one or more dholes may take over chasing their prey, while the rest of the pack keeps up at a steadier pace behind, taking over once the other group tires. Most chases are short, lasting only . When chasing fleet-footed prey, they run at a pace of . Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered. Once large prey is caught, one dhole will grab the prey's nose, while the rest of the pack pulls the animal down by the flanks and hindquarters. They do not use a killing bite to the throat. They occasionally blind their prey by attacking the eyes.
Serow The serows ( or ) are four species of medium-sized goat-like or antelope-like mammals of the genus ''Capricornis''. All four species of serow were until recently also classified under ''Naemorhedus'', which now only contains the gorals. Extant ...
s are among the only ungulate species capable of effectively defending themselves against dhole attacks, due to their thick, protective coats and short, sharp horns capable of easily impaling dholes. They will tear open their prey's flanks and
disembowel Disembowelment or evisceration is the removal of some or all of the organs of the gastrointestinal tract (the bowels, or viscera), usually through a horizontal incision made across the abdominal area. Disembowelment may result from an accident ...
it, eating the
heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...
,
liver The liver is a major Organ (anatomy), organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the Protein biosynthesis, synthesis of proteins and biochemicals necessary for ...
,
lungs The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of th ...
and some sections of the
intestines The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans ...
. The
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
and
rumen The rumen, also known as a paunch, is the largest stomach compartment in ruminants and the larger part of the reticulorumen, which is the first chamber in the alimentary canal of ruminant animals. The rumen's microbial favoring environment al ...
are usually left untouched. Prey weighing less than is usually killed within two minutes, while large stags may take 15 minutes to die. Once prey is secured, dholes will tear off pieces of the carcass and eat in seclusion. Unlike wolf packs, in which the breeding pair monopolises food, dholes give access to the pups at a kill. They are generally tolerant of
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
s at their kills. Both mother and young are provided with regurgitated food by other pack members.


Feeding ecology

Prey animals in India include
chital The chital or cheetal (''Axis axis''; ), also known as the spotted deer, chital deer, and axis deer, is a deer species native to the Indian subcontinent. It was first described and given a binomial name by German naturalist Johann Christian Po ...
, sambar deer,
muntjac Muntjacs ( ), also known as the barking deer or rib-faced deer, (URL is Google Books) are small deer of the genus ''Muntiacus'' native to South Asia and Southeast Asia. Muntjacs are thought to have begun appearing 15–35 million years ago, ...
,
mouse deer Chevrotains, or mouse-deer, are small even-toed ungulates that make up the family Tragulidae, the only extant members of the infraorder Tragulina. The 10  extant species are placed in three genera, but several species also are known only ...
,
barasingha The barasingha (''Rucervus duvaucelii''), also known as the swamp deer, is a deer species distributed in the Indian subcontinent. Populations in northern and central India are fragmented, and two isolated populations occur in southwestern Nepal. ...
,
wild boar The wild boar (''Sus scrofa''), also known as the wild swine, common wild pig, Eurasian wild pig, or simply wild pig, is a suid native to much of Eurasia and North Africa, and has been introduced to the Americas and Oceania. The species is ...
,
gaur The gaur (''Bos gaurus''; ), also known as the Indian bison, is a bovine native to South Asia and Southeast Asia, and has been listed as Vulnerable on the IUCN Red List since 1986. The global population was estimated at a maximum of 21,000 m ...
,
water buffalo The water buffalo (''Bubalus bubalis''), also called the domestic water buffalo or Asian water buffalo, is a large bovid originating in the Indian subcontinent and Southeast Asia. Today, it is also found in Europe, Australia, North America, So ...
es, banteng,
cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult mal ...
,
nilgai The nilgai (''Boselaphus tragocamelus'') (, literally meaning "blue cow") is the largest Asian antelope and is ubiquitous across the northern Indian subcontinent. It is the sole member of the genus ''Boselaphus'' and was described by Peter S ...
,
goat The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...
s,
Indian hare The Indian hare (''Lepus nigricollis''), also known as the black-naped hare, is a common species of hare native to the Indian subcontinent, and Java. Introductions It has been introduced to Madagascar, Comoro Islands, Andaman Islands, Irian Jay ...
s,
Himalayan field rat The Himalayan field rat (''Rattus nitidus''), sometimes known as the white-footed Indo-Chinese rat, is a species of rodent in the family Muridae. It has a wide range, being found in India, Bangladesh, Nepal, Bhutan, China, Myanmar, Laos, Thai ...
s and
langur The Colobinae or leaf-eating monkeys are a subfamily of the Old World monkey family that includes 61 species in 11 genera, including the black-and-white colobus, the large-nosed proboscis monkey, and the gray langurs. Some classifications sp ...
s. There is one record of a pack bringing down an
Indian elephant The Indian elephant (''Elephas maximus indicus'') is one of four extant recognised subspecies of the Asian elephant and native to mainland Asia. Since 1986, the Asian elephant has been listed as Endangered on the IUCN Red List as the wild pop ...
calf in
Assam Assam (; ) is a state in northeastern India, south of the eastern Himalayas along the Brahmaputra and Barak River valleys. Assam covers an area of . The state is bordered by Bhutan and Arunachal Pradesh to the north; Nagaland and Manipur ...
, despite desperate defense of the mother, resulting in numerous losses to the pack. In Kashmir, they prey on
markhor The markhor (''Capra falconeri'') is a large ''Capra'' species native to Central Asia, the Karakoram, and the Himalayas. It is listed on the IUCN Red List as Near Threatened since 2015. The markhor is the national animal of Pakistan, where it ...
, and thamin in Myanmar,
Malayan tapir The Malayan tapir (''Tapirus indicus''), also called Asian tapir, Asiatic tapir and Indian tapir, is the only tapir species native to Southeast Asia from the Malay Peninsula to Sumatra. It has been listed as Endangered on the IUCN Red List since ...
,
Sumatran serow The Sumatran serow (''Capricornis sumatraensis sumatraensis''), also known as the southern serow, is a subspecies of the mainland serow native to mountain forests in the Thai-Malay Peninsula and on the Indonesian island of Sumatra. It was previo ...
in
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
and the
Malay Peninsula The Malay Peninsula (Malay: ''Semenanjung Tanah Melayu'') is a peninsula in Mainland Southeast Asia. The landmass runs approximately north–south, and at its terminus, it is the southernmost point of the Asian continental mainland. The area ...
and Javan rusa in Java. In the
Tian Shan The Tian Shan,, , otk, 𐰴𐰣 𐱅𐰭𐰼𐰃, , tr, Tanrı Dağı, mn, Тэнгэр уул, , ug, تەڭرىتاغ, , , kk, Тәңіртауы / Алатау, , , ky, Теңир-Тоо / Ала-Тоо, , , uz, Tyan-Shan / Tangritog‘ ...
and
Tarbagatai Mountains The Tarbagatai Mountains ( mn, , , translit. ''Tarvagatai nuruu'', literally: "range with marmots"; ; kk, Тарбағатай жотасы, ''Tarbağatai jotasy'') are a range of mountains located in the north-western parts of Xinjiang, China, ...
, dholes prey on
Siberian ibex The Siberian ibex (''Capra sibirica''), also known as the Altai ibex, Central Asia(n) ibex, Gobi ibex, Himalayan ibex, Mongolian ibex or Tian Shan ibex, is a species of ibex that lives in central Asia. It has traditionally been treated as a subs ...
es,
arkhar The urial ( ; ''Ovis vignei''), also known as the arkars or shapo, is a wild sheep native to Central and South Asia. It is listed as Vulnerable on the IUCN Red List. Characteristics Urial males have large horns, curling outwards from the t ...
,
roe deer The roe deer (''Capreolus capreolus''), also known as the roe, western roe deer, or European roe, is a species of deer. The male of the species is sometimes referred to as a roebuck. The roe is a small deer, reddish and grey-brown, and well-adapt ...
,
Caspian red deer The Caspian red deer (''Cervus elaphus maral''), is one of the easternmost subspecies of red deer that is native to areas between the Black Sea and Caspian Sea such as Crimea, Asia Minor, the Caucasus Mountains region bordering Europe and Asia, ...
and
wild boar The wild boar (''Sus scrofa''), also known as the wild swine, common wild pig, Eurasian wild pig, or simply wild pig, is a suid native to much of Eurasia and North Africa, and has been introduced to the Americas and Oceania. The species is ...
. In the Altai and
Sayan Mountains The Sayan Mountains (russian: Саяны ''Sajany''; mn, Соёны нуруу, ''Soyonï nurû''; otk, 𐰚𐰇𐰏𐰢𐰤, Kögmen) are a mountain range in southern Siberia, Russia ( Buryatia, Irkutsk Oblast, Krasnoyarsk Krai, Tuva Republic ...
, they prey on
musk deer Musk deer can refer to any one, or all seven, of the species that make up ''Moschus'', the only extant genus of the family Moschidae. Despite being commonly called deer, they are not true deer belonging to the family Cervidae, but rather their fa ...
and
reindeer Reindeer (in North American English, known as caribou if wild and ''reindeer'' if domesticated) are deer in the genus ''Rangifer''. For the last few decades, reindeer were assigned to one species, ''Rangifer tarandus'', with about 10 subspe ...
. In eastern Siberia, they prey on roe deer,
Manchurian wapiti The Manchurian wapiti (''Cervus canadensis xanthopygus'') is a subspecies of the wapiti native to East Asia. Description The Manchurian wapiti's coat is reddish brown during summer, and brownish gray in winter. It has dark hairs on the neck and ...
, wild pig, musk deer and reindeer, while in
Primorye Primorsky Krai (russian: Приморский край, r=Primorsky kray, p=prʲɪˈmorskʲɪj kraj), informally known as Primorye (, ), is a federal subject (a krai) of Russia, located in the Far East region of the country and is a part of t ...
they feed on
sika deer The sika deer (''Cervus nippon''), also known as the Northern spotted deer or the Japanese deer, is a species of deer native to much of East Asia and introduced to other parts of the world. Previously found from northern Vietnam in the south to ...
and goral. In Mongolia, they prey on argali and rarely Siberian ibex. Like African wild dogs, but unlike wolves, dholes are not known to attack people. Dholes eat fruit and vegetable matter more readily than other canids. In captivity, they eat various kinds of grasses, herbs and leaves, seemingly for pleasure rather than just when ill. In summertime in the Tian Shan Mountains, dholes eat large quantities of mountain rhubarb. Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves. Livestock predation by dholes has been a problem in Bhutan since the late 1990s, as domestic animals are often left outside to graze in the forest, sometimes for weeks at a time. Livestock stall-fed at night and grazed near homes are never attacked. Oxen are killed more often than cows, probably because they are given less protection.


Enemies and competitors

In some areas, dholes are wiktionary:sympatric, sympatric to
tiger The tiger (''Panthera tigris'') is the largest living cat species and a member of the genus '' Panthera''. It is most recognisable for its dark vertical stripes on orange fur with a white underside. An apex predator, it primarily preys on u ...
s and
leopard The leopard (''Panthera pardus'') is one of the five extant species in the genus '' Panthera'', a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, a ...
s. 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 range (mean weights of for dhole and for leopard), while tigers selected for prey animals heavier than (but their mean prey weight was ). 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. Tigers are dangerous opponents for dholes, as they have sufficient strength to kill a dhole with a single paw strike. 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 destabilisation if one member of the breeding pair is killed. Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs. There are numerous records of leopards being treed by dholes. 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. 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. Dhole packs occasionally attack Asiatic black bears, snow leopards and sloth bears. When attacking bears, dholes will attempt to prevent them from seeking refuge in caves and lacerate their hindquarters. Although usually antagonistic toward wolves, they may hunt and feed alongside one another. There is at least one record of a lone wolf associating with a pair of dholes in Debrigarh Wildlife Sanctuary, and two observations in Satpura Tiger Reserve. They infrequently associate in mixed groups with
golden jackal The golden jackal (''Canis aureus''), also called common jackal, is a wolf-like canid that is native to Southeast Europe, Southwest Asia, South Asia, and regions of Southeast Asia. The golden jackal's coat varies in color from a pale creamy y ...
s. Domestic dogs may kill dholes, though they will feed alongside them on occasion.


Diseases and parasites

Dholes are vulnerable to a number of different diseases, particularly in areas where they are wiktionary:Sympatric, 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.


Threats

The dhole only rarely takes domestic livestock. Some ethnic groups like the Kuruba and Mon Khmer-speaking tribes will appropriate dhole kills; some Indian villagers welcome the dhole because of this appropriation of dhole kills. Dholes were persecuted throughout India for bounties until they were given protection by the Wildlife Protection Act of 1972. Methods used for dhole hunting included poisoning, snaring, shooting and clubbing at den sites. Native Indian people killed dholes primarily to protect livestock, while British sporthunters during the British Raj did so under the conviction that dholes were responsible for drops in Game (hunting), game populations. Persecution of dholes still occurs with varying degrees of intensity according to the region. Bounties paid for dholes used to be 25 rupees, though this was reduced to 20 in 1926 after the number of presented dhole carcasses became too numerous to maintain the established reward. In Indochina, dholes suffer heavily from nonselective hunting techniques such as snaring. The fur trade does not pose a significant threat to dholes. The people of India do not eat dhole flesh and their fur is not considered overly valuable. Due to their rarity, dholes were never harvested for their skins in large numbers in the Soviet Union and were sometimes accepted as dog or wolf pelts (being labeled as "half wolf" for the latter). The winter fur was prized by the Chinese, who bought dhole pelts in Ussuriysk during the late 1860s for a few silver rubles. In the early 20th century, dhole pelts reached eight rubles in Manchuria. In Semirechye, fur coats made from dhole skin were considered the warmest, but were very costly.


Conservation

In India, the dhole is protected under Schedule 2 of the Wildlife Protection Act, 1972. The creation of reserves under Project Tiger provided some protection for dhole populations sympatric with tigers. In 2014, the Indian government sanctioned its first dhole conservation breeding centre at the Indira Gandhi Zoological Park (IGZP) in Visakhapatnam. The dhole has been protected in Russia since 1974, though it is vulnerable to poison 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, the dhole is protected from all hunting, while conservation laws in Vietnam limit extraction and utilisation. In 2016, the Korean company Sooam Biotech was reported to be attempting to clone the dhole using dogs as surrogate mothers to help conserve the species.


In culture and literature

Three dhole-like animals are featured on the Coping (architecture), coping stone of the Bharhut stupa dating from 100 BC. They are shown waiting by a tree, with a woman or spirit trapped up it, a scene reminiscent of dholes treeing tigers. The animal's fearsome reputation in India is reflected by the number of pejorative names it possesses in Hindi, which variously translate as "red devil", "devil dog", "jungle devil", or "hound of Kali". Leopold von Schrenck had trouble obtaining dhole specimens during his exploration of
Amurland Outer Manchuria (russian: Приаму́рье, translit=Priamurye; zh, s=外满洲, t=外滿洲, p=Wài Mǎnzhōu), or Outer Northeast China ( zh, s=外东北, t=外東北, p=Wài Dōngběi), refers to a territory in Northeast Asia that is now ...
, as the local Gilyaks greatly feared the species. This fear and superstition was not, however, shared by neighbouring Tungusic peoples. It was speculated that this differing attitude towards the dhole was due to the Tungusic people's more nomadic, hunter-gatherer lifestyle. Dholes appear in Rudyard Kipling's ''Red Dog (Kipling short story), Red Dog'', where they are portrayed as aggressive and bloodthirsty animals which descend from the Deccan Plateau into the Seoni, Madhya Pradesh, Seeonee Hills inhabited by Mowgli and his adopted wolf pack to cause carnage among the jungle's denizens. They are described as living in packs numbering hundreds of individuals, and that even Shere Khan and Hathi make way for them when they descend into the jungle. The dholes are despised by the wolves because of their destructiveness, their habit of not living in dens and the hair between their toes. With Mowgli and Kaa's help, the Seeonee wolf pack manages to wipe out the dholes by leading them through bee hives and torrential waters before finishing off the rest in battle. Japanese author Uchida Roan wrote (; A dog's tale) in 1901 as a Japanese nationalism, nationalistic critique of the declining popularity of indigenous dog breeds, which he asserted were descended from the dhole. A fictional version of the dhole, imbued with supernatural abilities, appears in the season 6 episode of TV series ''The X-Files'', titled "Alpha (The X-Files), Alpha". In China, the dhole were widely known throughout history and mythology. One notable legendary creature is the Nine sons of the dragon, Yazi (), which was believed to be a creature that was part-dhole part-dragon. In modern times, however, the Chinese word for dhole () is often confused with 'jackal' or 'wolf', resulting in many confusions and mistranslations of dholes as jackals or wolves. Dholes also appear as enemies in the video game ''Far Cry 4'', alongside other predators such as the Bengal tiger, honey badger, snow leopard, clouded leopard, Tibetan wolf and Asian black bear. They can be found hunting the player and other non-player character, NPCs across the map, but are easily killed, being one of the weakest enemies in the game. They once again appear in the video game ''Far Cry Primal'', where they play similar roles as their counterparts in the previous game, but can now also be tamed and used in combat by Takkar, the main protagonist of the game.


Tameability

Brian Houghton Hodgson Brian Houghton Hodgson (1 February 1800 or more likely 1801 – 23 May 1894) was a pioneer naturalist and ethnologist working in India and Nepal where he was a British Resident. He described numerous species of birds and mammals from the Himala ...
kept captured dholes in captivity, and found, with the exception of one animal, they remained shy and vicious even after 10 months. According to
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker, ...
, adult dholes are nearly impossible to tame, though pups are docile and can even be allowed to play with domestic dog pups until they reach early adulthood. A dhole may have been presented as a gift to Ibbi-Sin as tribute.McIntosh, J. (2008). ''The ancient Indus Valley: new perspectives'', p. 130, ABC-CLIO,


See also

* ''Wild Dog Diaries''


Notes


References


Bibliography

*


External links


Dhole Home PageArchive
*ARKive
images and movies of the dholeSaving the dhole: The forgotten 'badass' Asian dog more endangered than tigers
''The Guardian'' (25 June 2015)
Photos of dhole in Bandipur
{{Taxonbar, from=Q132585 Dhole, Canina (subtribe) Taxa named by Peter Simon Pallas Mammals described in 1811 Carnivorans of Asia Mammals of South Asia Mammals of Southeast Asia Endangered fauna of Asia Pleistocene mammals of Asia Extant Middle Pleistocene first appearances Species endangered by habitat fragmentation