Editing Dhole

{{Short description|Species of mammal}}
{{Redirect|Mountain wolf}}
{{Distinguish|Red wolf|Northern Rocky Mountain wolf|African wild dog}}
{{Use dmy dates|date=April 2020}}
{{Speciesbox
| name = Dhole
| fossil_range = [[Middle Pleistocene]] – Recent {{fossilrange|0.78|0}}
| image = Dhole in its habitat (cropped).jpg
| image_caption = 
| image_upright = 0.9
| status = EN
| status_system = IUCN3.1
| status_ref = <ref name=iucn>{{cite iucn |title=''Cuon alpinus'' |author=Kamler, J.F. |author2=Songsasen, N. |author3=Jenks, K. |author4=Srivathsa, A. |author5=Sheng, L. |author6=Kunkel, K. |date=2015 |page=e.T5953A72477893 |doi=10.2305/IUCN.UK.2015-4.RLTS.T5953A72477893.en |access-date=24 January 2022}}</ref>
| status2 = CITES_A2 
| status2_system = CITES 
| status2_ref = <ref name=iucn/>
| genus = Cuon
| display_parents = 3
| parent_authority = [[Brian Houghton Hodgson|Hodgson]], 1838
| species = alpinus
| authority = ([[Peter Simon Pallas|Pallas]], 1811)
| subdivision_ranks = Subspecies
| subdivision = * ''C. a. adjustus''
* ''[[Ussuri dhole|C. a. alpinus]]''
* ''C. a. fumosus''
*† ''[[Tien Shan dhole|C. a. hesperius]]''
* ''C. a. laniger''
* ''C. a.  lepturus''
* ''C. a. sumatrensis''
*† ''C. a. antiquus''
*† ''C. a. caucasicus''
*† ''[[European dhole|C. a. europaeus]]''
| range_map = Cuon alpinus range.png
| range_map_caption = {{legend0|#ee161f|&nbsp;dhole range}}
| synonyms = ''Canis alpinus''
}}

The '''dhole''' ({{IPAc-en|d|oʊ|l}} {{respell|dohl}};<ref name=MWeb>{{cite web|url= https://www.merriam-webster.com/dictionary/dhole|title= "dhole"|author= <!--Not stated-->|website= Merriam-Webster|access-date= 7 January 2024|archive-date= 7 January 2024|archive-url= https://web.archive.org/web/20240107071959/https://www.merriam-webster.com/dictionary/dhole|url-status= live}}</ref><ref name=ColDict>{{cite web|url= https://www.collinsdictionary.com/dictionary/english/dhole|title= "dhole"|author= <!--Not stated-->|website= Collins Dictionary|access-date= 7 January 2024|archive-date= 1 May 2021|archive-url= https://web.archive.org/web/20210501140052/https://www.collinsdictionary.com/dictionary/english/dhole|url-status= live}}</ref> '''''Cuon alpinus''''') is a [[Canidae|canid]] native to [[South Asia|South]], [[East Asia|East]] and [[Southeast Asia]]. It is anatomically distinguished from members of the genus ''[[Canis]]'' in several aspects: its [[skull]] is convex rather than concave in profile, it lacks a third lower [[molar (tooth)|molar]], and the upper molars possess only a single [[Cusp (dentistry)|cusp]] as opposed to between two and four. During the [[Pleistocene]], the dhole ranged throughout [[Asia]], with its range also extending into [[Europe]] (with a single putative, controversial record also reported from [[North America]]) but became restricted to its historical range 12,000–18,000&nbsp;years ago. It is now extinct in [[Central Asia]], parts of [[Southeast Asia]], and possibly the [[Korean peninsula]] and [[Russia]].

Genetic evidence indicates that the dhole was the result of [[reticulate evolution]], emerging from the [[Hybrid (biology)|hybridization]] between a species closely related to genus ''Canis'' and one from a lineage closely related to the [[African wild dog]] (''Lycaon pictus'').

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&nbsp;individuals, but groups of over 40 are known. It is a diurnal pack hunter which preferentially targets large and medium-sized [[ungulate]]s. In tropical forests, the dhole competes with the [[tiger]] (''Panthera tigris'') and the [[leopard]] (''Panthera pardus''), targeting somewhat different prey species, but still with substantial dietary overlap.

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

==Etymology and naming==
The [[etymology]] of "dhole" is unclear. The possible earliest written use of the word in English occurred in 1808 by soldier Thomas Williamson, who encountered the animal in [[Ramgarh district|Ramghur district]], [[India]]. He stated that ''dhole'' was a common local name for the species.<ref>{{cite book |author=Williamson, T. |year=1808 |title=Oriental field sports: being a complete, detailed, and accurate description of the wild sports of the East|volume=II|publisher=Orme |location=London |url=https://books.google.com/books?id=1BlcAAAAcAAJ&pg=PA1}}</ref> In 1827, [[Charles Hamilton Smith]] claimed that it was derived from a language spoken in 'various parts of the East'.<ref name="smith1827">{{cite book |author=Smith, C. H. |year=1827 |title=The class Mammalia |publisher=Geo. B. Whittaker |location=London |url=https://archive.org/stream/classmammalia21827cuvi#page/326/mode/2up/search/dhole}}</ref>

Two years later, Smith connected this word with {{langx|tr|[[wikt:deli#Turkish|deli]]}} 'mad, crazy', and erroneously compared the Turkish word with {{langx|osx|[[wikt:dol#Old English|dol]]}} and {{langx|nl|[[wikt:dol#Dutch|dol]]}} (cfr. also {{langx|en|[[wikt:dull#English|dull]]}}; {{langx|de|[[wikt:toll#German|toll]]}}),<ref name="smith1839">{{cite book |author1=Smith, C. H. |author2=Jardine, W. |year=1839 |title=The natural history of dogs: Canidae or genus canis of authors; including also the genera hyaena and proteles |volume=I |publisher=W.H. Lizars |location=Edinburgh, UK |url=https://archive.org/stream/naturalhistoryof139smit#page/166/mode/2up}}</ref> which are in fact from the [[Proto-Germanic language|Proto-Germanic]] *''dwalaz'' 'foolish, stupid'.<ref>{{Citation |title=A Handbook of Germanic Etymology |last=Orel |first=V. |author-link=Vladimir Orel |year=2003 |publisher=Brill |location=Leiden, DE; Boston, MA |isbn=978-90-04-12875-0 |page=[https://archive.org/details/handbookofgerman0000orel/page/81 81] |url=https://archive.org/details/handbookofgerman0000orel/page/81 }}</ref> [[Richard Lydekker]] wrote nearly 80&nbsp;years later that the word was not used by the natives living within the species' range.<ref name="lydekker1907">{{cite book |last=Lydekker |first=R. |author-link=Richard Lydekker |url=https://archive.org/stream/gameanimalsofind00lyde#page/360/mode/2up |title=The game animals of India, Burma, Malaya, and Tibet |publisher=R. Ward Limited |year=1907 |location=London, UK}}</ref> The [[Merriam-Webster]] ''Dictionary'' theorises that it may have come from the {{langx|kn|ತೋಳ|tōḷa|wolf}}.<ref>[http://www.merriam-webster.com/dictionary/dhole dhole] {{Webarchive|url=https://web.archive.org/web/20161224095139/https://www.merriam-webster.com/dictionary/dhole |date=24 December 2016 }}. Merriam-Webster Dictionary.</ref>

Other English names for the species include '''Asian wild dog''', '''Asiatic wild dog''',<ref name=f>{{Harvnb|Fox|1984}}</ref> '''Indian wild dog''',<ref name="lydekker1907"/> '''whistling dog''', '''red dog''',<ref name=cohen1978>{{cite journal |last1=Cohen |first1=James A. |title=Cuon alpinus |journal=Mammalian Species |date=1978 |issue=100 |pages=1–3 |doi=10.2307/3503800 |jstor=3503800 |doi-access=free}}</ref> and '''red wolf'''.<ref name=heptner1998>{{cite book |editor1=Heptner, V. G. |editor2=Naumov, N. P. |year=1998 |chapter=Genus ''Cuon'' Hodgson, 1838 |chapter-url=https://archive.org/stream/mammalsofsov211998gept#page/566/mode/2up |title=Mammals of the Soviet Union |volume=(II. Part 1A: Sirenia and Carnivora (Sea Cows, Wolves, and Bears)) |pages=566–586 |publisher=Smithsonian Institution and National science Foundation |location=Washington, DC |isbn=1-886106-81-9}}</ref>

==Taxonomy and evolution==
[[File:Cuon alpinus Cova Negra.jpg|thumb|Skeletal remains of a [[European dhole]] dating back to the upper [[Last Glacial Period#Würm glaciation (Alps)|Würm]] period from Cova Negra de [[Xàtiva]], [[Valencia]], Spain]]
[[File:Leopold v. Schrenck - Cuon alpinus.png|thumb|Illustration (1859) by [[Leopold von Schrenck]], one of the first accurate depictions of the species, based on a single skin purchased in the village of Dshare on the [[Amur]]<ref name= "schrenk1859"/>]]

<!-- The species was first described in European literature in 1794 by the Russian traveler and surveyor Pesterev, who encountered dholes during his travels in the [[Russian Far East]]. He described the animal as being a regular pack hunter of [[Alpine ibex]], and of bearing many similarities with the [[golden jackal]].<ref>{{cite journal |last1=Pesterev |first1=E. |year=1794 |journal=Journ. von Russland |page=24 }}</ref> is not relevant for taxonomy-->
''Canis alpinus'' was the [[binomial name]] proposed by [[Peter Simon Pallas]] in 1811, who described its range as encompassing the upper levels of Udskoi Ostrog in [[Amurland]], towards the eastern side and in the region of the upper [[Lena River]], around the [[Yenisei]] River and occasionally crossing into [[China]].<ref name=Pallas1811>{{cite book| last=Pallas| first= P. S.| year= 1811| chapter=''Canis alpinus''| pages= 34–35 |chapter-url= https://archive.org/stream/zoographiarossoa11831pall#page/34/mode/2up |title=Zoographia Rosso-Asiatica: Sistens omnium animalium in extenso Imperio Rossico, et adjacentibus maribus observatorum recensionem, domicilia, mores et descriptiones, anatomen atque icones plurimorum| location=Petropoli|publisher=In officina Caes. Acadamiae Scientiarum Impress| language=la}}</ref><ref name= "auto">{{cite book |editor1=Heptner, V. G. |editor2=Naumov, N. P. |title= Mammals of the Soviet Union |volume= II |at=Part 1A: Sirenia and Carnivora (Sea Cows, Wolves, and Bears), pp. 571–586 |location= Washington, DC |publisher= Smithsonian Institution and National Science Foundation |year=1998 |chapter=Red Wolf ''Cuon alpinus'' Pallas, 1811 |chapter-url=https://archive.org/details/mammalsofsov211998gept/page/570}}</ref> This northern Russian range reported by Pallas during the 18th and 19th centuries is "considerably north" of where this species occurs today.<ref name="auto"/>

''Canis primaevus'' was a name proposed by [[Brian Houghton Hodgson]] in 1833 who thought that the dhole was a primitive ''Canis'' form and the [[progenitor]] of the [[domestic dog]].<ref>{{cite journal | last=Hodgson | first= B. H. |year=1833 |title=Description and Characters of the Wild Dog of the Himalaya (''Canis primævus'') |journal=Asiatic Researches |volume=XVIII |issue=2 |pages=221–237, 235 |url= https://archive.org/stream/asiaticresearche181833cal#page/220/mode/2up}}</ref> Hodgson later took note of the dhole's physical distinctiveness from the genus ''Canis'' and proposed the genus ''Cuon''.<ref>{{cite journal | last=Hodgson | first= B. H. |year=1842 |url=https://archive.org/stream/calcuttajournalo02mccl#page/204 |title=European notices of Indian canines, with further illustrations of the new genus ''Cuon'' vel ''Chrysæus'' |journal=Calcutta Journal of Natural History |volume=II |pages=205–209}}</ref>

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.<ref name= "thenius1954">{{cite journal | last= Thenius | first= E. |year=1955 |title=Zur Abstammung der Rotwölfe (Gattung ''Cuon'' Hodgson) |trans-title=On the origins of the dholes (Genus ''Cuon'' Hodgson) |journal= Österreichische Zoologische Zeitschrift |volume=5 |pages=377–388 |url= http://www.landesmuseum.at/pdf_frei_remote/OEZ_05_0377-0387.pdf |archive-url=https://web.archive.org/web/20141110114203/http://www.landesmuseum.at/pdf_frei_remote/OEZ_05_0377-0387.pdf |archive-date=2014-11-10 |url-status=live |language=de}}</ref> 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]] era [[Valdarno]], Italy and in China—was almost indistinguishable from the genus ''Canis''. In comparison, the modern species has greatly reduced [[Molar (tooth)|molar]]s and the [[Cusp (anatomy)|cusp]]s have developed into sharply trenchant points. During the Early [[Middle Pleistocene]] there arose both ''Canis majori stehlini'' that was the size of a large [[wolf]], and the early dhole ''Canis alpinus'' Pallas 1811 which first appeared at [[Hundsheim]] and [[Mosbach]] in Germany. In the [[Late Pleistocene]] era the [[European dhole]] (''C. a. europaeus'') was modern-looking and the transformation of the lower molar into a single cusped, slicing tooth had been completed; however, its size was comparable with that of a wolf. This subspecies became extinct in Europe at the end of the late [[Last Glacial Period#Würm glaciation (Alps)|Würm]] period, but the species as a whole still inhabits a large area of Asia.<ref name="kurten1968"/> The European dhole may have survived up until the early [[Holocene]] in the [[Iberian Peninsula]],<ref name="ripoll2010">{{cite journal |title=Presence of the genus ''Cuon'' in upper Pleistocene and initial Holocene sites of the Iberian Peninsula: New remains identified in archaeological contexts of the Mediterranean region |journal=Journal of Archaeological Science |doi= 10.1016/j.jas.2009.10.008 |volume=37 |issue=3 |pages=437–450 |year=2010 |name-list-style=amp |last1=Ripoll |first1=M.P.R. |last2=Morales Pérez |first2=J.V. |last3=Sanchis Serra |first3= A. |last4=Aura Tortosa |first4= J.E. |last5=Montañana |first5=I.S N.|bibcode=2010JArSc..37..437R | display-authors= 3}}</ref> and what is believed to be dhole remains have been found at Riparo Fredian in northern [[Italy]] dated 10,800 years old.<ref>{{cite journal |author1= Petrucci, M. |author2=Romiti, S. |author3= Sardella, R. |name-list-style=amp |year= 2012 |title=The Middle-Late Pleistocene ''Cuon'' Hodgson, 1838 (Carnivora, Canidae) from Italy |journal=Bollettino della Società Paleontologica Italiana |volume=51 |issue=2 |pages=146 |url=http://paleoitalia.org/media/u/archives/07.Petrucci_et_al._2012_BSPI_512_1.pdf |archive-url=https://web.archive.org/web/20150102034611/http://paleoitalia.org/media/u/archives/07.Petrucci_et_al._2012_BSPI_512_1.pdf |archive-date=2015-01-02 |url-status=usurped}}</ref><ref name=Taron2021>{{cite journal| doi= 10.3390/genes12020144|title=Ancient DNA from the Asiatic Wild Dog (''Cuon alpinus'') from Europe |year= 2021| last1=Taron|first1=U.H.|last2=Paijmans|first2=J.L.A. |last3=Barlow |first3=A. |last4=Preick |first4=M. |last5=Iyengar |first5=A. |last6=Drăgușin|first6=V. |last7=Vasile|first7=Ș. |last8=Marciszak |first8=A. |last9=Roblíčková|first9=M. |last10=Hofreiter|first10=M. | display-authors= 3 |journal=Genes |volume=12 |issue=2 |page=144 |pmid=33499169 |pmc=7911384 |doi-access=free}}</ref>

The vast Pleistocene range of this species also included numerous islands in Asia that this species no longer inhabits, such as [[Sri Lanka]], [[Borneo]] and possibly [[Palawan]] in the [[Philippines]].<ref>{{cite book |author=Nowak, R. M. |title=Walker's Carnivores of the World |location= Baltimore, Maryland |publisher=Johns Hopkins University Press |year=2005 |pages=110–111 |chapter=''Cuon'' |chapter-url= https://archive.org/details/walkerscarnivore0000nowa/page/110/mode/2up |isbn=9780801880322}}</ref><ref>{{cite journal |last=Cranbrook |first=Earl of |year=1988 |title=The contribution of archaeology to the zoogeography of Borneo : with the first record of a wild canid of Early Holocene Age ; a contribution in celebration of the distinguished scholarship of Robert F. Inger on the occasion of his sixty-fifth birthday |url=https://archive.org/details/contributionofar42earl/page/n9 |journal=Fieldiana Zoology |series=42 |pages=6–24}}</ref><ref>{{Cite journal |last1=Ochoa |first1=J. |last2=Paz |first2=V. |last3=Lewis |first3=H. |last4=Carlos |first4=J. |last5=Robles |first5=E. |last6=Amano |first6=N. |last7=Ferreras |first7=M. R. |last8=Myra |first8=L. |last9=Vallejo |first9=B. Jr. |last10=Velarde |first10=G. |last11=Villaluz |first11=S. A. |last12=Ronquillo |first12=W. |last13=Solheim |first13=W. |display-authors=3 |year=2014 |title=The archaeology and palaeobiological record of Pasimbahan-Magsanib Site, northern Palawan, Philippines |journal=Philippine Science Letters |volume=7 |issue=1 |pages=22–36 |issn=2094-2818 |hdl-access=free |hdl=10197/5394|s2cid=55862985}}</ref><ref>{{cite book |author1=Dennell, R. |author2=Parr, M. |title=Southern Asia, Australia, and the Search for Human Origins |location=New York |publisher=Cambridge University Press |year=2014 |page=139 |url=https://books.google.com/books?id=Qq-kAgAAQBAJ&pg=PA139 |isbn=9781107729131 |access-date=29 July 2016 |archive-date=15 April 2021 |archive-url=https://web.archive.org/web/20210415090456/https://books.google.com/books?id=Qq-kAgAAQBAJ&pg=PA139 |url-status=live }}</ref><ref>{{cite book |last=Tarling |first=N. |title=The Cambridge History of Southeast Asia |volume=(Volume One: From Early Times to ca. 1800) |location=Cambridge, UK |publisher=Cambridge University Press |year=1992 |page=84 |isbn=978-0-5213-5505-6 |url=https://books.google.com/books?id=rOw8AAAAIAAJ&pg=PA84 |access-date=30 August 2015 |archive-date=3 February 2021 |archive-url=https://web.archive.org/web/20210203060937/https://books.google.com/books?id=rOw8AAAAIAAJ&pg=PA84 |url-status=live }}</ref><ref>{{cite journal |author1=Piper, P.J. |name-list-style=amp |author2=Ochoa, J.B. |author3=Robles, E.C. |author4=Lewis, H. |author5=Paz, V. |display-authors=3 |year=2011 |title=Palaeozoology of Palawan Island, Philippines |journal=Quaternary International |volume=233 |issue=2 |pages=142–158 |doi=10.1016/j.quaint.2010.07.009 |bibcode=2011QuInt.233..142P |url=https://www.researchgate.net/publication/222656323 |access-date=29 July 2021 |archive-date=3 March 2024 |archive-url=https://web.archive.org/web/20240303095556/https://www.researchgate.net/publication/222656323_Palaeozoology_of_Palawan_Island_Philippines |url-status=live }}</ref> Middle Pleistocene dhole fossils have also been found in the Matsukae Cave in northern [[Kyushu]] Island in western [[Japan]] and in the Lower Kuzuu fauna in [[Tochigi Prefecture]] in [[Honshu]] Island, east Japan.<ref>{{cite journal |last1=Ogino |first1=S. |last2=Otsuka |first2=H. |last3= Harunari |first3=H. |title=The Middle Pleistocene Matsugae Fauna, Northern Kyushu, West Japan |journal= Paleontological Research |date= December 2009 |volume=13 |issue=4 |pages=367–384 |doi=10.2517/1342-8144-13.4.367 |bibcode=2009PalRe..13..367O |s2cid=53691225}}</ref> 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]] in [[south China]] where they no longer exist.<ref name=Turvey_al2019>{{cite journal |last1=Turvey |first1=S.T. |last2=Walsh |first2=C. |last3=Hansford |first3=J.P. |last4=Crees |first4=J.J. |last5=Bielby |first5=J. |last6=Duncan |first6=C. |last7=Hu |first7=K. |last8=Hudson |first8=M.A. | display-authors= 3 |title= Complementarity, completeness and quality of long-term faunal archives in an Asian biodiversity hotspot |journal= Philosophical Transactions of the Royal Society B: Biological Sciences |date= 2019 |volume=374 |issue=1788 |page= 20190217 |doi=10.1098/rstb.2019.0217 |pmid=31679488 |pmc=6863502 |doi-access=free}}</ref> Additionally, fossils of canidae possibly belonging to dhole have been excavated from [[Dajia River]] in [[Taichung County]], Taiwan.<ref>{{Cite web |date=October 2009 |title=疑豺、狼化石 大甲溪床現蹤 |trans-title=Suspected dhole and wolf fossils found in the bed of Dajia River |url=https://twgeoref.moeacgs.gov.tw/GipOpenWeb/wSite/ct?xItem=97564&ctNode=1326&mp=6 |url-status=dead |archive-url=https://web.archive.org/web/20230422094352/https://twgeoref.moeacgs.gov.tw/GipOpenWeb/wSite/ct?xItem=97564&ctNode=1326&mp=6 |archive-date=22 April 2023 |access-date=1 September 2022 |website=Taiwan Geoscience Portal |language=zh}}</ref>

A single record of the dhole is known from North America. This consists of a jaw fragment and teeth of [[Late Pleistocene]] age found in San Josecito Cave in northeast Mexico, dating to around 27–11,000 years ago.<ref>{{Cite journal |last1=Tedford |first1=Richard H. |last2=Wang |first2=Xiaoming |last3=Taylor |first3=Beryl E. |date=2009-09-03 |title=Phylogenetic Systematics of the North American Fossil Caninae (Carnivora: Canidae) |url=http://www.bioone.org/doi/abs/10.1206/574.1 |journal=Bulletin of the American Museum of Natural History |language=en |volume=325 |pages=1–218 |doi=10.1206/574.1 |issn=0003-0090}}</ref> Other researchers have either considered this record as "insufficient" or suggested that further corroboration is required for the definitive taxonomic attribution of these specimens.<ref>{{cite journal|author1=Arias-Alzate, A.|author2=rroyave, F.J.|author3=Romero Goyeneche, O.Y.|author4=Hurtado Heredia, R.G.|author5=González-Maya, J.F.|author6=Arroyo-Cabrales, J.|author7=Peterson, A.T.|author8=Martínez-Meyer, E.|year=2022|title=Functional niche constraints on carnivore assemblages (Mammalia: Carnivora) in the Americas: What facilitates coexistence through space and time?|journal=Journal of Biogeography|volume=49|issue=3|pages=497–510|doi=10.1111/jbi.14319|bibcode=2022JBiog..49..497A }}</ref><ref>{{Cite journal |last1=Ferrusquía-Villafranca |first1=Ismael |last2=Arroyo-Cabrales |first2=Joaquín |last3=Martínez-Hernández |first3=Enrique |last4=Gama-Castro |first4=Jorge |last5=Ruiz-González |first5=José |last6=Polaco |first6=Oscar J. |last7=Johnson |first7=Eileen |date=2010-04-15 |title=Pleistocene mammals of Mexico: A critical review of regional chronofaunas, climate change response and biogeographic provinciality |url=https://www.sciencedirect.com/science/article/pii/S104061820900442X |journal=Quaternary International |series=Faunal Dynamics and Extinction in the Quaternary: Studies in Honor of Ernest L. Lundelius, Jr. |language=en |volume=217 |issue=1 |pages=53–104 |doi=10.1016/j.quaint.2009.11.036 |bibcode=2010QuInt.217...53F |issn=1040-6182}}</ref>

{{Cladogram|align=right|title=[[Phylogenetic tree]] of the [[Canina (subtribe)|wolf-like canids]] with timing in millions of years{{refn|group=lower-alpha|name=mtdna_seq_desc|For a full set of supporting references refer to the note (a) in the phylotree at [[Evolution of the wolf#Wolf-like canids]]}}
|cladogram={{clade | style = font-size:85%;line-height:80%;width:400px;
 |label1=[[Caninae]] 3.5 [[Megaannum|Ma]]
 |1={{clade
  |label1=3.0
  |1={{clade
   |label1=2.5
   |1={{clade
    |label1=2.0
    |1={{clade
     |label1=0.96
     |1={{clade
       |label1=0.6
       |1={{clade
        |label1=0.38
        |1={{clade
         |1={{clade
          |1={{clade
           |1=[[Domestic dog]] <span style="{{MirrorH}}">[[File:202104 Dog.svg|50 px]]</span>
           |2=[[Wolf|Gray wolf]] [[File:Dogs, jackals, wolves, and foxes (Plate I).jpg|50 px]]
}}
}}
         |2=[[Coyote]] [[File:Dogs, jackals, wolves, and foxes (Plate IX).jpg|50 px]]
}}
        |2=[[African wolf]] [[File:Dogs, jackals, wolves, and foxes (Plate XI).jpg|50 px]]
}}
       |2=[[Golden jackal]] [[File:Dogs, jackals, wolves, and foxes (Plate X).jpg|50 px]]
}}
      |2=[[Ethiopian wolf]] [[File:Dogs, jackals, wolves, and foxes (Plate VI).jpg|50 px]]
}}
     |2=Dhole [[File:Dogs, jackals, wolves, and foxes (Plate XLI).jpg|50 px]]
}}
     |2=[[African wild dog]] [[File:Dogs, jackals, wolves, and foxes (Plate XLIV).jpg|50 px]]
}}
  |2={{clade
   |1={{clade
    |1=[[Side-striped jackal]] [[File:Dogs, jackals, wolves, and foxes (Plate XIII).jpg|50 px]]
    |2=[[Black-backed jackal]] [[File:Dogs, jackals, wolves, and foxes (Plate XII).jpg|50 px]]
}}
    |label1=2.6
}}
}}
}}
}}

Dholes are also known from the Middle and Late Pleistocene fossil record of Europe.<ref>{{Cite journal |last1=Ripoll |first1=Manuel Pérez |last2=Morales Pérez |first2=Juan V. |last3=Sanchis Serra |first3=Alfred |last4=Aura Tortosa |first4=J. Emili |last5=Montañana |first5=Inocencio Sarrión |date=1 March 2010 |title=Presence of the genus Cuon in upper Pleistocene and initial Holocene sites of the Iberian Peninsula: new remains identified in archaeological contexts of the Mediterranean region |url=https://www.sciencedirect.com/science/article/pii/S0305440309003756 |journal=[[Journal of Archaeological Science]] |volume=37 |issue=3 |pages=437–450 |doi=10.1016/j.jas.2009.10.008 |bibcode=2010JArSc..37..437R |issn=0305-4403 |access-date=16 January 2024 |via=Elsevier Science Direct |archive-date=24 September 2015 |archive-url=https://web.archive.org/web/20150924171734/http://www.sciencedirect.com/science/article/pii/S0305440309003756 |url-status=live }}</ref> In 2021, the analyses of the [[Mitochondrial DNA|mitochondrial]] [[genomes]] extracted from the fossil remains of two extinct European dhole specimens from the Jáchymka cave, [[Czech Republic]] dated 35,000–45,000 years old indicate that these were genetically [[Basal (phylogenetics)|basal]] to modern dholes and possessed much greater genetic diversity.<ref name= Taron2021/>

The dhole's distinctive morphology has been a source of much confusion in determining the species' systematic position among the Canidae. [[George Gaylord Simpson|George Simpson]] placed the dhole in the [[subfamily]] Simocyoninae alongside the [[African wild dog]] and the [[bush dog]], on account of all three species' similar dentition.<ref name="simpson1945">{{Cite journal |last=Simpson |first=G. G. |year=1945 |title=The principles of classification and a classification of mammals |journal=Bulletin of the American Museum of Natural History |volume=85 |pages=1–350 |hdl-access=free |hdl=2246/1104}}</ref> Subsequent authors, including [[Juliet Clutton-Brock]], noted greater morphological similarities to canids of the genera ''Canis'', ''[[Dusicyon]]'' and ''[[Arctic fox|Alopex]]'' than to either ''[[Speothos]]'' or ''[[Lycaon (genus)|Lycaon]]'', with any resemblance to the latter two being due to [[convergent evolution]].<ref name=cluttonbrock1976>{{cite journal |author1=Clutton-Brock, J. |author2=Corbet, G. G. |author3=Hills, M. |name-list-style=amp |year=1976 |url=http://biostor.org/reference/65860 |title=A review of the family Canidae, with a classification by numerical methods |journal=Bulletin of the British Museum of Natural History |volume=29 |pages=179–180 |access-date=8 December 2014 |archive-url=https://web.archive.org/web/20131217052727/http://biostor.org/reference/65860 |archive-date=17 December 2013 |url-status=dead}}</ref>

Some authors consider the extinct ''Canis'' [[subgenus]] ''[[Xenocyon]]'' as ancestral to both the genus ''Lycaon'' and the genus ''Cuon''.<ref name=moulle2006/><ref name= baryshnikov2012/><ref name= cherin2013>{{cite journal |doi= 10.1007/s10914-013-9227-4 |title=Re-Defining ''Canis etruscus'' (Canidae, Mammalia): A New Look into the evolutionary history of Early Pleistocene dogs resulting from the outstanding fossil record from Pantalla (Italy) |journal=Journal of Mammalian Evolution |volume=21|pages=95–110 |year=2013 |last1=Cherin|first1=M. |last2=Bertè |first2=D.F.| last3= Rook|first3=L. |last4=Sardella|first4=R. |s2cid=17083040}}</ref><ref name= wang2008/>{{rp|p149}} Subsequent studies on the canid [[genome]] revealed that the dhole and African wild dog are closely related to members of the genus ''Canis''.<ref name="lindblad-toh2005">{{cite journal |last1=Lindblad-Toh |first1=K. |last2=Wade |first2=C.M. |last3=Mikkelsen |first3=T.S. |last4=Karlsson |first4=E.K. |last5=Jaffe |first5=D.B. |last6=Kamal |first6=M. |last7=Clamp |first7=M. |last8=Chang |first8=J.L. |last9=Kulbokas |first9=E.J. |last10=Zody |first10=M.C. |last11=Mauceli |first11=E. |last12=Xie |first12=X. |last13=Breen |first13=M. |last14=Wayne |first14=R.K. |last15=Ostrander |first15=E.A. |display-authors=29 |year=2005 |title=Genome sequence, comparative analysis, and haplotype structure of the domestic dog |journal=Nature |volume=438 |issue=7069 |pages=803–819 |bibcode=2005Natur.438..803L |doi=10.1038/nature04338 |pmid=16341006 |doi-access=free |last16=Ponting |first16=C.P. |last17=Galibert |first17=F. |last18=Smith |first18=D.R. |last19=Dejong |first19=P.J. |last20=Kirkness |first20=E. |last21=Alvarez |first21=P. |last22=Biagi |first22=T. |last23=Brockman |first23=W. |last24=Butler |first24=J. |last25=Chin |first25=C.W. |last26=Cook |first26=A. |last27=Cuff |first27=J. |last28=Daly |first28=M.J. |last29=Decaprio |first29=D. |last30=Gnerre |first30=S.}}</ref> This closeness to ''Canis'' may have been confirmed in a menagerie in [[Madras]], where according to zoologist [[Reginald Innes Pocock]] there is a record of a dhole that interbred with a golden jackal.<ref name=Pocock1941>{{cite book |author-link=Reginald Innes Pocock| last=Pocock| first= R. I. |year=1941 |title=Fauna of British India: Mammals |volume=2 |publisher=Taylor & Francis |chapter=Genus ''Cuon'' Hodgs. |pages=146–163 |chapter-url= https://archive.org/stream/PocockMammalia2/pocock2#page/n163/mode/2up}}</ref> DNA sequencing of the [[Sardinian dhole]] (''Cynotherium sardous'') an extinct small canine species formerly native to the island of Sardinia in the Mediterranean, and which has often been suggested to have descended from ''Xenocyon'', has found that it is most closely related to the living dhole among canines.<ref name=":0">{{Cite journal |last1=Ciucani |first1=Marta Maria |last2=Jensen |first2=Julie Kragmose |last3=Sinding |first3=Mikkel-Holger S. |last4=Smith |first4=Oliver |last5=Lucenti |first5=Saverio Bartolini |last6=Rosengren |first6=Erika |last7=Rook |first7=Lorenzo |last8=Tuveri |first8=Caterinella |last9=Arca |first9=Marisa |last10=Cappellini |first10=Enrico |last11=Galaverni |first11=Marco |last12=Randi |first12=Ettore |last13=Guo |first13=Chunxue |last14=Zhang |first14=Guojie |last15=Sicheritz-Pontén |first15=Thomas |date=December 2021 |title=Evolutionary history of the extinct Sardinian dhole |url=https://linkinghub.elsevier.com/retrieve/pii/S0960982221013154 |journal=Current Biology |language=en |volume=31 |issue=24 |pages=5571–5579.e6 |doi=10.1016/j.cub.2021.09.059 |pmid=34655517 |bibcode=2021CBio...31E5571C |s2cid=238996621 |hdl-access=free |hdl=2158/1252046}}</ref>

=== Admixture with the African wild dog ===
In 2018, [[whole genome sequencing]] was used to compare all members (apart from the black-backed and side-striped jackals) of the genus ''Canis'', along with the dhole and the [[African wild dog]] (''Lycaon pictus''). There was strong evidence of ancient [[genetic admixture]] between the dhole and the African wild dog. Today, their ranges are remote from each other; however, during the [[Pleistocene]] era the dhole could be found as far west as Europe. The study proposes that the dhole's distribution may have once included the [[Middle East]], from where it may have admixed with the African wild dog in [[North Africa]]. However, there is no evidence of the dhole having existed in the Middle East nor North Africa,<ref name=gopalakrishnan2018>{{cite journal |doi=10.1016/j.cub.2018.08.041 |pmid=30344120 |pmc=6224481 |title=Interspecific gene flow shaped the evolution of the Genus ''Canis'' |journal=Current Biology |volume=28 |issue=21 |pages=3441–3449.e5 |year=2018|last1=Gopalakrishnan|first1=S. |last2=Sinding|first2=M.-H.S. |last3=Ramos-Madrigal|first3=J. |last4= Niemann |first4=J. |last5= Samaniego Castruita|first5=J.A. |last6=Vieira|first6=F.G. |last7= Carøe|first7=C. |last8=Montero |first8= M.d.M. |last9=Kuderna| first9=L. |last10= Serres|first10=A. |last11= González-Basallote |first11=V.M. |last12=Liu|first12=Y.-H. |last13=Wang |first13=G.-D. |last14=Marques-Bonet |first14=T. |last15=Mirarab |first15=S. |last16=Fernandes|first16=C. |last17= Gaubert|first17=P. |last18= Koepfli|first18=K.-P. |last19=Budd |first19=J. |last20= Rueness| first20= E.K. |last21=Heide-Jørgensen| first21= M.P. |last22= Petersen |first22=B. |last23=Sicheritz-Ponten|first23=T. |last24= Bachmann| first24=L. |last25= Wiig|first25=Ø. |last26=Hansen |first26= A.J. |last27= Gilbert| first27= M.T.P. |bibcode=2018CBio...28E3441G | display-authors= 3}}</ref> though the ''Lycaon'' was present in Europe during the Early Pleistocene, with its last record in the region dating to 830,000 years ago. Genetic evidence from the Sardinan dhole suggests that both Sardinian and modern dholes (which are estimated to have split from each other around 900,000 years ago) share ancestry from the ''Lycaon'' lineage, but this ancestry is significantly higher in modern dholes than in the Sardinian dhole.<ref name=":0"/>
{{clear}}

===Subspecies===
Historically, up to ten subspecies of dholes have been recognised.<ref name="ellerman1966">{{cite book |author1=Ellerman, J.R. |author2= Morrison-Scott, T.C.S. |year=1966 |title=Checklist of Palaearctic and Indian mammals |url= https://archive.org/stream/checklistofindia00elle#page/232/mode/2up |publisher=British Museum (Natural History) |location=London, UK}}</ref> {{As of|2005}}, seven subspecies are recognised.<ref name= Wozencraft2004/><ref name= Orrell2019/>

However, studies on the dhole's [[mtDNA]] and [[microsatellite]] genotype showed no clear subspecific distinctions. Nevertheless, two major phylogeographic groupings were discovered in dholes of the Asian mainland, which likely diverged during a [[Glacial period|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, {{as of|2005|lc=y}}, 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]] of the [[International Union for the Conservation of Nature]] (IUCN) states that further research is needed because all of the samples were from the southern part of this species' range and the [[Tien Shan]] subspecies has distinct morphology.<ref>{{Cite web| url= https://www.canids.org/species/view/PREKQW171301| title= Dhole &#124; Canids| website= canids.org| publisher= | access-date= | archive-date= 23 September 2021| archive-url= https://web.archive.org/web/20210923000058/https://canids.org/species/view/PREKQW171301| url-status= live}}</ref>

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.<ref>{{cite journal |last1=Iyengar |first1=A. |last2= Babu |first2=V. N. |last3=Hedges |first3=S. |last4=Venkataraman |first4=A.B. |last5=Maclean |first5=N. |last6=Morin |first6=P. A. | display-authors= 3 |title= Phylogeography, genetic structure, and diversity in the dhole (''Cuon alpinus'') |journal= Molecular Ecology |date=2005 |volume=14 |issue=8 |pages=2281–2297 |doi=10.1111/j.1365-294X.2005.02582.x |pmid=15969714 |bibcode=2005MolEc..14.2281I |s2cid=8280811 |url= http://eprints.iisc.ac.in/3543/1/Phylogeography_genetic.pdf |archive-url=https://web.archive.org/web/20200307041249/http://eprints.iisc.ac.in/3543/1/Phylogeography_genetic.pdf |archive-date=2020-03-07 |url-status=live}}</ref> Fossils of dhole from the early [[Middle Pleistocene]] have been found in [[Java]].<ref>{{cite journal |last1= Volmer |first1=R. |last2=van der Geer |first2=A.A.E. |last3=Cabrera |first3= P.A. |last4=Wibowo |first4=U. P. |last5=Kurniawan |first5=I.| display-authors= 3 |title=When did Cuon reach Java? – Reinvestigation of canid fossils from Homo erectus faunas |journal= Geobios |date=September 2019 |volume=55 |pages=89–102 |doi= 10.1016/j.geobios.2019.06.004 |bibcode=2019Geobi..55...89V |doi-access=free}}</ref>

{| class="wikitable sortable" width=90% font=90%
|- bgcolor="#115a6c"
!Subspecies
!Image
!Trinomial authority
!Description
!Distribution
!Synonyms
|----
|''C. a. adjustus'' '''Burmese dhole''',<ref name=Pocock1941/> Indian dhole
|[[File:2010-kabini-dhole.jpg|150 px]]
||Pocock, 1941<ref name=Pocock1941/>
|Reddish coat, short hair on the paws and black whiskers<ref name=Durbin2004>{{cite book |last1=Durbin |first1=D.L. |last2=Venkataraman |first2=A. |last3=Hedges |first3=S.|last4=Duckworth|first4=W. |name-list-style=amp |title=Canids: Foxes, Wolves, Jackals, and Dogs:Status Survey and Conservation Action Plan|publisher=IUCN The World Conservation Union|editor1-last=Sillero-Zubiri|editor1-first=C. |editor2-last=Hoffmann|editor2-first=M. |editor3-last=Macdonald|editor3-first=D.W. |year=2004 |chapter=8.1–Dhole |pages=211 |isbn=978-2831707860|chapter-url=http://www.carnivoreconservation.org/files/actionplans/canids.pdf |archive-url=https://web.archive.org/web/20061013041056/http://www.carnivoreconservation.org/files/actionplans/canids.pdf |archive-date=2006-10-13 |url-status=live|access-date=31 January 2020}}</ref>
|Northeastern India and south of the [[Ganges]] River, northern Myanmar<ref name=Durbin2004/>
|<small>''antiquus'' (Matthew & Granger, 1923), ''dukhunensis'' (Sykes, 1831)</small>
|----
|''[[Ussuri dhole|C. a. alpinus]]'' '''Ussuri dhole'''<ref name="heptner1998"/>
([[nominate subspecies]])
|[[File:Leopold v. Schrenck - Cuon alpinus.png|150 px]]
|[[Peter Simon Pallas|Pallas]], 1811<ref name=Pallas1811/>
|Thick tawny red coat, greyish neck and ochre muzzle<ref name=Durbin2004/>
|East of the eastern [[Sayan Mountains]], eastern Russia, northeastern Asia<ref name=Durbin2004/>
| {{center|–}}
|----
|''C. a. fumosus''<ref name=Pocock1936/>
|
|Pocock, 1936<ref name=Pocock1936/>
|Luxuriant yellowish-red coat, dark back and grey neck<ref name=Durbin2004/>
|Western [[Sichuan]], China and Mongolia. Southern Myanmar, Thailand, Laos, Cambodia, Vietnam, Malaysia and [[Java]], Indonesia<ref name=Durbin2004/>
|<small>''infuscus'' (Pocock, 1936), ''javanicus'' (Desmarest, 1820)</small>
|----
|†''[[Tien Shan dhole|C. a. hesperius]]'' '''Tien Shan dhole'''<ref name="heptner1998"/>
|[[File:Keulmans Cuon alpinus alpinus.png|150 px]]
|Afanasjev and Zolotarev, 1935<ref name=Afanasjev1935/>
|Long yellow tinted coat, white underside and pale whiskers<ref name=Durbin2004/> Smaller than ''C. a. alpinus'', with wider skull and lighter-coloured winter fur.<ref name="heptner1998"/>
|Altai, Tian Shan and Pamir mountain ranges.<ref name=iucn/> Currently considered to be extinct since 1946.<ref name=Makenov-2018/><ref name=iucn/>
|<small>''jason'' (Pocock, 1936)</small>
|----
|''C. a. laniger''<ref name=Pocock1936/>
|
|Pocock, 1936<ref name=Pocock1936/>
|Full, yellowish-grey coat, tail not black but same colour as body<ref name="Durbin2004"/>
|Southern Tibet, Himalayan Nepal, Sikkim, Bhutan and Kashmir<ref name=Durbin2004/>
|<small>''grayiformis'' (Hodgson, 1863), ''primaevus'' (Hodgson, 1833)</small>
|----
|''C. a. lepturus''<ref name=Pocock1936/>
|[[File:Zoo Osnabrück, Dhole (4413112187).jpg|150 px]]
|Heude, 1892<ref name=Heude1892/>
|Uniform red coat with thick underfur<ref name=Durbin2004/>
|South of the [[Yangtze]] River, China<ref name=Durbin2004/>
|<small>''clamitans'' (Heude, 1892), ''rutilans'' (Müller, 1839), ''sumatrensis'' (Hodgson, 1833)</small>
|----
|Sumatran dhole and Javan dhole ''C. a. sumatrensis''<ref name=Castello2018/>
|[[File:Keulemans Cuon alpinus javanicus.png|150 px]]
|[[Thomas Hardwicke|Hardwicke]], 1821<ref name=Hardwicke1821/>
|Red coat and dark whiskers<ref name=Durbin2004/>
|[[Sumatra]], Indonesia<ref name=Durbin2004/> Its range is highly fragmented with multiple protected areas in Sumatra and Java.<ref name=iucn/>
|
|----
|}

==Characteristics==
[[File:Dhole Kanha.jpg|thumb|A dhole in [[Kanha National Park]], India]]
[[File:Dholeskull.jpg|thumb|Dhole skull and molars illustrated by [[St. George Mivart]] (1890)]]
[[File:Dhole in Ueno, Tokyo.jpg|thumb|Captive adult dhole]]
The dhole's 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.<ref name=heptner1998/> The dorsal and lateral [[guard hair]]s in adults measure {{cvt|20–30|mm}} in length. Dholes in the [[Moscow Zoo]] moult once a year from March to May.<ref name=cohen1978/> A [[Melanism|melanistic]] individual was recorded in the northern [[Coimbatore]] Forest Division in [[Tamil Nadu]].<ref>{{cite journal |author1=Mahabal, A. |name-list-style=amp |author2=Sharma, R.M. |author3=Patil, R.N. |author4=Jadhav, S. |year=2019 |title=Colour aberration in Indian mammals: a review from 1886 to 2017 |journal=Journal of Threatened Taxa |volume=11 |issue=6 |pages=13690–13719 |doi=10.11609/jott.3843.11.6.13690-13719 |doi-access=free}}</ref>

The dhole has a wide and massive skull with a well-developed [[sagittal crest]],<ref name=heptner1998/> and its [[masseter muscle]]s are highly developed compared to other canid species, giving the face an almost [[hyena]]-like appearance.<ref name="f61_2">{{Harvnb|Fox|1984|pp=61–2}}</ref> The [[rostrum (anatomy)|rostrum]] is shorter than that of domestic dogs and most other canids.<ref name=cohen1978/> It has six rather than seven lower [[Molar (tooth)|molars]].<ref name=f41>{{Harvnb|Fox|1984|pp=41}}</ref> 
The upper molars are weak, being one third to one half the size of those of wolves and have only one [[Cusp (dentistry)|cusp]] as opposed to between two and four, as is usual in canids,<ref name=heptner1998/> an adaptation thought to improve shearing ability and thus speed of prey consumption. This may allow dholes to compete more successfully with [[kleptoparasite]]s.<ref name=Durbin2004/> In terms of size, dholes average about {{cvt|88|-|113|cm}} in length (excluding a {{cvt|41|-|50|cm}} long tail), and stand around {{cvt|42|-|50|cm}} at the shoulders.<ref name=cohen1978/> Adult females can weigh {{cvt|10|-|17|kg}}, while the slightly larger male may weigh {{cvt|15|-|21|kg}}. The mean weight of adults from three small samples was {{cvt|15.1|kg}}.<ref name=Durbin2004/><ref name=Kamler_al2012>{{cite journal |author=Kamler J. F. |author2=Johnson A. |name-list-style=amp |author3=Vongkhamheng C. |author4=Bousa A. |year=2012 |title=The diet, prey selection, and activity of dholes (''Cuon alpinus'') in northern Laos |journal=Journal of Mammalogy |volume=93 |issue=3 |pages=627–633 |doi=10.1644/11-mamm-a-241.1 |s2cid=54996917 |url=https://www.researchgate.net/publication/257297107}}</ref>

In appearance, the dhole has been variously described as combining the physical characteristics of the [[gray wolf]] and the [[red fox]],<ref name=heptner1998/> and as being "cat-like" on account of its long backbone and slender limbs.<ref name=thenius1954/>

==Distribution and habitat==
[[File:Dhole Mudhumalai.jpg|thumb|Lone dhole strolling through the jungle in [[Mudumalai National Park]]]]
[[File:Dhole at Tadoba.jpg|thumb|Dhole in [[Tadoba Andhari Tiger Reserve]], Maharashtra, India]]
Historically, the dhole lived in [[Singapore]] and throughout [[Central Asia]] including [[Afghanistan]], [[Kyrgyzstan]], [[Kazakhstan]], [[Mongolia]], [[Tajikistan]] and [[Uzbekistan]], though it is now considered to be regionally extinct in these regions.<ref name=iucn/> Historical record in [[South Korea]] from the [[Veritable Records of the Joseon Dynasty]] also indicate that the dhole once inhabited [[Yangju]] in [[Gyeonggi Province]],<ref>{{Cite web |title=조선왕조실록 – 양주 등지에 출몰하는 승냥이와 범을 잡도록 전교하다 |trans-title=Veritable Records of the Joseon Dynasty – Ordered to capture the dholes and tigers in Yangju and other areas |url=https://sillok.history.go.kr/id/kob_11004008_005 |access-date=7 September 2024 |website=[[Veritable Records of the Joseon Dynasty]] |language=ko}}</ref> but it is now also extinct in South Korea,<ref name=iucn/> with the last known capture reports in 1909 and 1921 from [[Yeoncheon]] of [[Gyeonggi Province]].<ref>{{Cite web |title=기묘한 생김새의 개과 동물, 승냥이|trans-title=Dhole, a canid with an odd appearance|access-date=10 September 2024|website=usjournal.kr|date=27 August 2020 |language=ko|url=https://m.usjournal.kr/news/newsview.php?ncode=1065574756016281}}</ref> The current presence of dholes in [[North Korea]] and [[Pakistan]] is considered uncertain.<ref name=iucn/> The dholes also once inhabited the alpine steppes extending into [[Kashmir]] to the [[Ladakh]] area, though they disappeared from 60% of their historic range in India during the past century.<ref name=iucn/> In India, Myanmar, Indochina, Indonesia and China, it prefers forested areas in [[alpine zone]]s and is occasionally sighted in [[plain]]s regions.<ref name="heptner1998"/>

In the [[Bek-Tosot Conservancy]] of southern [[Kyrgyzstan]], the possible presence of the dholes was considered likely based on genetic samples collected in 2019.<ref name=Cancellare-2022>{{cite journal |author1=Cancellare, I.A. |name-list-style=amp |author2=Kachel, S.M. |author3=Kubanychbekov, Z. |author4=Kulenbekov, R. |author5=Pilgrim, K.L. |author6=McCarthy, K.P. |author7=Weckworth, B.V. |year=2022 |title=New distribution record of dhole from southern Kyrgyzstan using non-invasive genetic sampling |journal=Canid Biology & Conservation |volume=24 |issue=1 |pages=1–3 |url=https://www.canids.org/CBC/24/Dhole_distribution_Kyrgyzstan.pdf |access-date=22 February 2022 |archive-date=2 February 2022 |archive-url=https://web.archive.org/web/20220202160121/https://www.canids.org/CBC/24/Dhole_distribution_Kyrgyzstan.pdf |url-status=live}}</ref> This was the first record of dholes from the country in almost three decades.<ref name=Cancellare-2022/>

The dhole might still be present in the [[Tunkinsky National Park]] in extreme southern [[Siberia]] near [[Lake Baikal]].<ref>{{cite journal |author=Williams, M. |name-list-style=amp |author2=Troitskaya, N. |year=2007 |url=http://www.wild-russia.org/pubs/pdfs/42.pdf |archive-url=https://web.archive.org/web/20120128043346/http://www.wild-russia.org/pubs/pdfs/42.pdf |archive-date=2012-01-28 |url-status=live |title=Then and Now: Updates from Russia's Imperiled Zapovedniks |journal=Russian Conservation News |volume=42 |page=14}}</ref> It possibly still lives in the [[Primorsky Krai]] province in far eastern Russia, where it was considered a rare and endangered species in 2004, with unconfirmed reports in the Pikthsa-Tigrovy Dom protected forest area; no sighting was reported in other areas since the late 1970s.<ref>{{Cite book |last=Newell |first=J. |url=https://archive.org/details/russianfareastre0000newe |title=The Russian Far East: A Reference Guide for Conservation and Development |publisher=Daniel & Daniel |year=2004 |edition=Second |location=McKinleyville}}</ref>{{page needed|date=March 2024}}
Currently, no other recent reports are confirmed of dholes being present in [[Russia]],<ref name=Makenov-2018>{{Cite journal |last=Makenov |first=M. |date=2018 |title=Extinct or extant? A review of dhole (''Cuon alpinus'' Pallas, 1811) distribution in the former USSR and modern Russia |journal=Mammal Research |volume=63 |issue=1 |pages=1–9 |doi=10.1007/s13364-017-0339-8 |s2cid=20037994}}</ref> so the IUCN considered them to be extinct in Russia.<ref name=iucn/> However, the dhole might be present in the eastern [[Sayan Mountains]] and in the [[Transbaikal]] region; it has been sighted in [[Tofalaria]] in the [[Irkutsk Oblast]], the [[Republic of Buryatia]] and [[Zabaykalsky Krai]].<ref name=Makenov-2018/>

One pack was sighted in the [[Qilian Mountains]] in 2006.<ref>{{cite journal |author=Harris, R. B. |year=2006 |title= Attempted predation on blue sheep ''Pseudois nayaur'' by dholes ''Cuon alpinus'' |journal=Journal of the Bombay Natural History Society |volume=103 |pages=95–97}}</ref>
In 2011 to 2013, local government officials and herders reported the presence of several dhole packs at elevations of {{cvt|2000|to|3500|m}} near [[Taxkorgan Nature Reserve]] in the [[Xinjiang]] Autonomous Region. Several packs and a female adult with pups were also recorded by [[camera trap]]s at elevations of around {{cvt|2500|to|4000|m}} in Yanchiwan National Nature Reserve in the northern [[Gansu Province]] in 2013–2014.<ref name=Riordan2015>{{cite journal |author=Riordan, P. |year=2015 |title=New evidence of dhole ''Cuon alpinus'' populations in northwest China |journal=Oryx |volume=49 |issue=2 |pages=203–204 |doi=10.1017/s0030605315000046 |doi-access=free}}</ref>
Dholes have been also reported in the [[Altyn-Tagh]] Mountains.<ref>{{cite journal |author1=Yadong, X. |author2=Diqiang, L. |author3=Wenfa, X. |author4=Yuguang, Z. |author5=Bin, F. |author6=Heng, J. |year=2015 |title=Records of the dhole (''Cuon alpinus'') in an arid region of the Altun Mountains in western China |journal=European Journal of Wildlife Research |volume=61 |issue=6 |pages=903–907 |doi=10.1007/s10344-015-0947-z |bibcode=2015EJWR...61..903X |s2cid=16752357}}</ref>

In China's [[Yunnan]] Province, dholes were recorded in [[Baima Xueshan Nature Reserve]] in 2010–2011.<ref>{{cite journal |author1=Li, X. |name-list-style=amp |author2=Buzzard, P. |author3=Chen, Y. |author4=Jiang, X. |year=2013 |title=Patterns of livestock predation by carnivores: human-wildlife conflict in northwest Yunnan, China |journal=Environ Manage |volume=52 |issue=6 |pages=1334–1340 |doi=10.1007/s00267-013-0192-8 |pmid=24202281 |bibcode=2013EnMan..52.1334L |s2cid=5273403}}</ref> Dhole samples were obtained in [[Jiangxi]] Province in 2013.<ref>{{cite book |author=Canid and Hyaenid Taxon Advisory Group |year=2017 |title=Best Practice Guideline Dhole (''C. alpinus'') |location=Amsterdam |publisher=European Association of Zoos and Aquaria |url=https://www.eaza.net/assets/Uploads/CCC/2017-Dhole-EAZA-Best-Practice-Guidelines-Approved.pdf |archive-url=https://web.archive.org/web/20200206040139/https://www.eaza.net/assets/Uploads/CCC/2017-Dhole-EAZA-Best-Practice-Guidelines-Approved.pdf |archive-date=2020-02-06 |url-status=live}}</ref>
Confirmed records by camera-trapping since 2008 have occurred in southern and western [[Gansu]] province, southern [[Shaanxi]] province, southern [[Qinghai]] province, southern and western [[Yunnan]] province, western [[Sichuan]] province, the southern [[Xinjiang]] Autonomous Region and in the Southeastern [[Tibet]] Autonomoous Region.<ref>Kao, J., N. Songsasen, K. Ferraz and K. Traylor-Holzer (Eds.) (2020). Range-wide Population and Habitat Viability Assessment for the Dhole, ''Cuon alpinus''. IUCN SSC Conservation Planning Specialist Group, Apple Valley, MN, USA. p8. https://www.canids.org/resources/Dhole_PHVA_Report_2020.pdf {{Webarchive|url=https://web.archive.org/web/20200606174930/https://www.canids.org/resources/Dhole_PHVA_Report_2020.pdf |date=6 June 2020 }}</ref> 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.<ref name=Turvey_al2019/>

The dhole occurs in most of India south of the Ganges, particularly in the Central Indian Highlands and the [[Western Ghats|Western]] and [[Eastern Ghats]]. It is also present in [[Arunachal Pradesh]], [[Assam]], [[Meghalaya]] and [[West Bengal]] and in the [[Indo-Gangetic Plain]]'s [[Terai]] region. Dhole populations in the [[Himalaya]]s and northwest India are fragmented.<ref name=iucn/>

In 2011, dhole packs were recorded by camera traps in the [[Chitwan National Park]].<ref>{{cite journal |author=Thapa, K. |author2=Kelly, M. J. |author3=Karki, J. B. |author4=Subedi, N. |name-list-style=amp |year=2013 |title=First camera trap record of pack hunting dholes in Chitwan National Park, Nepal |journal=Canid Biology & Conservation |volume=16 |issue=2 |pages=4–7 |url=http://www.canids.org/CBC/16/Dholes_in_Nepal.pdf |archive-url=https://web.archive.org/web/20141221190135/http://www.canids.org/CBC/16/Dholes_in_Nepal.pdf |archive-date=2014-12-21 |url-status=live}}</ref> Its presence was confirmed in the [[Kanchenjunga Conservation Area]] in 2011 by camera traps.<ref>{{Cite journal |doi=10.3126/init.v4i0.5531 |title=The Pack Hunter (Dhole): Received Little Scientific Attention |journal=The Initiation |volume=4 |pages=8–13 |year=2011 |last1=Khatiwada |first1=A. P. |last2=Awasthi |first2=K. D. |last3=Gautam |first3=N. P. |last4=Jnawali |first4=S. R. |last5=Subedi |first5=N. |last6=Aryal |first6=A. |name-list-style=amp |url=http://www.nepjol.info/index.php/INIT/article/download/5531/4560 |doi-access=free |access-date=20 April 2018 |archive-date=22 September 2017 |archive-url=https://web.archive.org/web/20170922103658/https://www.nepjol.info/index.php/INIT/article/download/5531/4560 |url-status=live}}</ref> In February 2020, dholes were sighted in the [[Vansda National Park]], with camera traps confirming the presence of two individuals in May of the same year. This was the first confirmed sighting of dholes in [[Gujarat]] since 1970.<ref>{{Cite news |last=Parmar |first=V. |date=2020 |title=Rare whistling dogs spotted in Gujarat after 50 years |work=[[The Times of India]] |url=https://timesofindia.indiatimes.com/city/surat/whistling-dog-spotted-in-gujarat-after-50-years/articleshow/75905282.cms |access-date=1 June 2020 |archive-date=1 June 2020 |url-status=live |archive-url=https://web.archive.org/web/20200601141034/https://timesofindia.indiatimes.com/city/surat/whistling-dog-spotted-in-gujarat-after-50-years/articleshow/75905282.cms}}</ref>

In [[Bhutan]], the dhole is present in [[Jigme Dorji National Park]].<ref>{{cite journal |author=Wangchuk, T. |year=2004 |title= Predator-prey dynamics: the role of predators in the control of problem species |journal= Journal of Bhutan Studies |volume=10 |pages=68–89 |url=http://www.bhutanstudies.org.bt/publicationFiles/JBS/JBS_Vol10/v10-7.pdf |archive-url=https://web.archive.org/web/20150224070842/http://www.bhutanstudies.org.bt/publicationFiles/JBS/JBS_Vol10/v10-7.pdf |archive-date=2015-02-24 |url-status=live}}</ref><ref>{{cite journal |author1=Thinley, P. |author2=Kamler, J. F. |author3=Wang, S. W. |author4=Lham, K. |author5=Stenkewitz, U. |year=2011 |title=Seasonal diet of dholes (''Cuon alpinus'') in northwestern Bhutan |journal=Mammalian Biology |volume=76 |issue=4 |pages=518–520 |doi=10.1016/j.mambio.2011.02.003 |bibcode=2011MamBi..76..518T}}</ref>

In [[Bangladesh]], it inhabits forest reserves in the [[Sylhet]] area, as well the [[Chittagong Hill Tracts]] in the southeast. Recent camera trap photos in the Chittagong in 2016 showed the continued presence of the dhole.<ref>{{Cite web|url=https://www.theguardian.com/environment/radical-conservation/2016/mar/01/tiger-country-scientists-uncover-wild-surprises-in-tribal-bangladesh|title=Tiger country? Scientists uncover wild surprises in tribal Bangladesh |work=[[The Guardian]] |date=2016 |access-date=6 June 2020|archive-date=29 July 2020|archive-url=https://web.archive.org/web/20200729204731/https://www.theguardian.com/environment/radical-conservation/2016/mar/01/tiger-country-scientists-uncover-wild-surprises-in-tribal-bangladesh|url-status=live}}</ref> These regions probably do not harbour a viable population, as mostly small groups or solitary individuals were sighted.<ref name=iucn/>

In [[Myanmar]], the dhole is present in several protected areas.<ref name=iucn/> In 2015, dholes and tigers were recorded by camera-traps for the first time in the hill forests of [[Karen State]].<ref>{{cite journal |author=Saw Sha Bwe Moo |author2=Froese, G.Z.L. |author3=Gray, T. N. E. |year=2017 |title=First structured camera-trap surveys in Karen State, Myanmar, reveal high diversity of globally threatened mammals |journal=Oryx |volume=52 |issue=3 |doi=10.1017/S0030605316001113 |pages=1–7|doi-access=free}}</ref>

Its range is highly fragmented in the [[Malaysian Peninsula]], [[Sumatra]], [[Java]], [[Vietnam]] and [[Thailand]], with the Vietnamese population considered to be possibly extinct.<ref name=iucn/> In 2014, camera trap videos in the montane tropical forests at {{Cvt|2000|m}} in the [[Kerinci Seblat National Park]] in Sumatra revealed its continued presence.<ref>{{cite web |url=http://www.fauna-flora.org/sumatran-secrets-revealed-high-altitude-camera-trapping/ |title=Sumatran secrets start to be revealed by high altitude camera trapping |website=Flora and Fauna International |access-date=5 September 2015|archive-url=https://web.archive.org/web/20170501235901/http://www.fauna-flora.org/sumatran-secrets-revealed-high-altitude-camera-trapping/|archive-date=1 May 2017|url-status=dead}}</ref> A camera trapping survey in the [[Khao Ang Rue Nai Wildlife Sanctuary]] in Thailand from January 2008 to February 2010 documented one healthy dhole pack.<ref>{{cite journal |author1=Jenks, K. E. |author2=Songsasen, N. |author3=P. Leimgruber |name-list-style=amp |year=2012 |title=Camera trap records of dholes in Khao Ang Rue Nai Wildlife Sanctuary, Thailand |journal=Canid News |pages=1–5 |url=http://www.canids.org/canidnews/15/Camera_trap_records_of_dholes_in_Thailand.pdf |archive-url=https://web.archive.org/web/20141221202145/http://www.canids.org/canidnews/15/Camera_trap_records_of_dholes_in_Thailand.pdf |archive-date=2014-12-21 |url-status=live}}</ref> In northern [[Laos]], dholes were studied in [[Nam Et-Phou Louey]] National Protected Area.<ref name=Kamler_al2012/> Camera trap surveys from 2012 to 2017 recorded dholes in the same Nam Et-Phou Louey National Protected Area.<ref>{{cite journal |last1=Rasphone |first1=Akchousanh |last2=Kéry |first2=Marc |last3=Kamler |first3=Jan F. |last4=Macdonald |first4=David W. |title=Documenting the demise of tiger and leopard, and the status of other carnivores and prey, in Lao PDR's most prized protected area: Nam Et - Phou Louey |journal=Global Ecology and Conservation |date=October 2019 |volume=20 |pages=e00766 |doi=10.1016/j.gecco.2019.e00766 |s2cid=202920288 |doi-access=free |bibcode=2019GEcoC..2000766R}}</ref>

In [[Vietnam]], dholes were sighted only in [[Pu Mat National Park]] in 1999, in [[Yok Don National Park]] in 2003 and 2004; and in [[Ninh Thuan Province]] in 2014.<ref>{{cite journal |last1=Hoffmann |first1=M. |last2=Abramov |first2=A. |last3=Duc |first3=H. M. |last4=Trai |first4=L. T. |last5=Long |first5=B. |last6=Nguyen |first6=A. |last7=Son |first7=N. T. |last8=Rawson |first8=B. |last9=Timmins |first9=R. |last10=Bang |first10=T. V. |last11=Willcox |first11=D. |title=The status of wild canids (Canidae, Carnivora) in Vietnam |journal=Journal of Threatened Taxa |date=2019 |volume=11 |issue=8 |pages=13951–13959 |doi=10.11609/jott.4846.11.8.13951-13959 |s2cid=198272874 |doi-access=free |name-list-style=amp}}</ref>

A disjunct dhole population was reported in the area of [[Trabzon]] and [[Rize]] in northeastern [[Turkey]] near the border with [[Georgia (country)|Georgia]] in the 1990s.<ref>{{cite journal |author1=Serez, M. |name-list-style=amp |author2=Eroðlu, M. |year=1994 |title=A new threatened wolf species, ''Cuon alpinus hesperius'' Afanasiev and Zolatarev, 1935 in Turkey |journal=Council of Europe Environmental Encounters Series |volume=17 |pages=103–106}}</ref> This report was not considered to be reliable.<ref name=iucn/> One single individual was claimed to have been shot in 2013 in the nearby [[Kabardino-Balkaria]] Republic of Russia in the central [[Caucasus]]; its remains were analysed in May 2015 by a biologist from the [[Kabardino-Balkarian State University]], who concluded that the skull was indeed that of a dhole.<ref>{{cite journal |author=Khatukhov, A.M. |title=Красный волк (''Cuon alpinus'' Pallas, 1811) на Центральном Кавказе |trans-title=The Dhole (''Cuon alpinus'' Pallas 1811) in the Central Caucasus |journal=Современные проблемы науки и образования |volume=3 |pages=574–581 |year=2015 |url=http://www.science-education.ru/pdf/2015/3/195.pdf |archive-url=https://web.archive.org/web/20151208144903/http://www.science-education.ru/pdf/2015/3/195.pdf |archive-date=2015-12-08 |url-status=live}}</ref> In August 2015, researchers from the [[National Museum of Natural History (Bulgaria)|National Museum of Natural History]] and the [[Karadeniz Technical University]] started an expedition to track and document possible Turkish population of dhole.<ref>{{cite web |date=2015 |title=NMNHS expedition went on the trail of an unknown population of the rare dhole in Turkey |url=http://www.nmnhs.com/15081601-news_en.html |url-status=live |archive-url=https://web.archive.org/web/20150909061531/http://www.nmnhs.com/15081601-news_en.html |archive-date=9 September 2015 |access-date=4 September 2015 |work=National Museum of Natural History, Sofia (NMNHS)}}</ref> In October 2015, they concluded that two skins of alleged dholes in Turkey probably belonged to dogs, pending DNA analysis of samples from the skins, and, having analysed photos of the skull of alleged dhole in Kabardino-Balkaria Republic of Russia, they concluded it was a grey wolf.<ref>{{cite web |last=Coel |first=C. |date=12 October 2015 |title=[UPDATE] Strongly endangered and undescribed subspecies of dhole discovered? Dhole NOT less endangered than previously thought, according to NMNHS (Bulgaria) |url=http://cetaf.org/strongly-endangered-and-undescribed-subspecies-of-dhole-discovered |url-status=dead |archive-url=https://web.archive.org/web/20151208061158/http://cetaf.org/strongly-endangered-and-undescribed-subspecies-of-dhole-discovered |archive-date=8 December 2015 |access-date=15 October 2015 |website=Consortium of European Taxonomic Facilities}}</ref>

==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.<ref name="f93">{{Harvnb|Fox|1984|p=93}}</ref> Other sounds include whines (food soliciting), growls (warning), screams, chatterings (both of which are alarm calls) and yapping cries.<ref name="f95">{{Harvnb|Fox|1984|p=95}}</ref> In contrast to wolves, dholes do not [[Howling|howl]] or bark.<ref name="heptner1998"/>

Dholes have a complex [[body language]]. Friendly or submissive greetings are accompanied by horizontal lip retraction and the lowering of the tail, as well as licking. Playful dholes 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.<ref name="f97">{{Harvnb|Fox|1984|p=97}}</ref>

===Social and territorial behaviour===
Dholes are more social than [[gray wolves]],<ref name="heptner1998"/> 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.<ref name="f85">{{Harvnb|Fox|1984|p=85}}</ref> They live in [[clan]]s rather than [[pack (canine)|pack]]s, 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.<ref name="f81">{{Harvnb|Fox|1984|pp=81–2}}</ref> 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.<ref name="f86"/> Intragroup fighting is rarely observed.<ref name="nowak1983"/>

[[File:20140303 7687 Pench Dhole.jpg|thumb|Dholes playing, [[Pench National Park]]]]
[[File:Asiatic Wild Dogs (43518944044).jpg|thumb|A pack of Asiatic wild dog]]

Dholes are far less [[territory (animal)|territorial]] than wolves, with pups from one clan often joining another without trouble once they mature sexually.<ref name="f92">{{Harvnb|Fox|1984|p=92}}</ref> Clans typically number 5 to 12 individuals in India, though clans of 40 have been reported. In [[Thailand]], clans rarely exceed three individuals.<ref name="cohen1978"/> Unlike other canids, there is no evidence of dholes using [[urine]] to mark their territories or travel routes. When urinating, dholes, especially males, may [[Raised-leg urination|raise one hind leg]] or both to result in a handstand. Handstand urination is also seen in [[bush dog]]s (''Speothos venaticus'')<ref>{{cite journal |title=Einige beobachtungen zum verhalten des Dekkan-Rothundes (Cuon alpinus dukhunensis Sykes) im Kanha National Park |trans-title=Some observations on the behavior of the Deccan Red Dog (Cuon alpinus dukhunensis Sykes) in Kanha National Park |last1=Keller |first1=R. |journal= Vierteljahresschrift. Naturf. Ges. Zürich |volume=118 |year=1973|pages=129–135 |language=de |url= https://www.ngzh.ch/archiv/1973_118/118_1/118_13.pdf |archive-url=https://web.archive.org/web/20190812095239/https://www.ngzh.ch/archiv/1973_118/118_1/118_13.pdf |archive-date=2019-08-12 |url-status=live }}</ref> and domestic dogs.<ref>{{cite journal |last1=Wirant |first1=Sharon Cudd |first2=Betty |last2=McGuire |url=https://www.researchgate.net/publication/248335168 |title=Urinary behavior of female domestic dogs (''Canis familiaris''): influence of reproductive status, location, and age |journal=Applied Animal Behaviour Science |volume=85 |issue=3–4 |year=2004 |pages=335–348|doi=10.1016/j.applanim.2003.09.012 }}</ref> They may defecate in conspicuous places, though a territorial function is unlikely, as [[faeces]] are mostly deposited within the clan's territory rather than the periphery. Faeces are often deposited in what appear to be communal [[Animal latrine|latrines]]. They do not scrape the earth with their feet, as other canids do, to mark their territories.<ref name="f97"/>

===Denning===
Four kinds of den have been described; simple earth dens with one entrance (usually remodeled [[striped hyena]] or [[porcupine]] dens); complex cavernous earth dens with more than one entrance; simple cavernous dens excavated under or between rocks; and complex cavernous dens with several other dens in the vicinity, some of which are interconnected. Dens are typically located under dense scrub or on the banks of dry rivers or creeks. The entrance to a dhole den can be almost vertical, with a sharp turn three to four feet down. The tunnel opens into an antechamber, from which extends more than one passage. Some dens may have up to six entrances leading up to {{cvt|30|m}} of interconnecting tunnels. These "cities" may be developed over many generations of dholes, and are shared by the clan females when raising young together.<ref name="f43">{{Harvnb|Fox|1984|pp=43–49}}</ref> Like [[African wild dog]]s and [[dingo]]es, dholes will avoid killing prey close to their dens.<ref name="f80"/>

===Reproduction and development===
[[File:Cuon alpinus alpinus puppy.jpg|thumb|Dhole pup, [[Kolmården Wildlife Park]]]]
In India, the [[mating season]] occurs between mid-October and January, while captive dholes in the [[Moscow Zoo]] breed mostly in February.<ref name="cohen1978"/> Unlike wolf packs, dhole clans may contain more than one breeding female.<ref name="f86"/> More than one female dhole may den and rear their litters together in the same den.<ref name="nowak1983"/> During [[mating]], the female assumes a crouched, cat-like position. There is no [[copulatory tie]] characteristic of other canids when the male dismounts. Instead, the pair lie on their sides facing each other in a semicircular formation.<ref name="f79">{{Harvnb|Fox|1984|p=79}}</ref> The [[gestation period]] lasts 60–63 days, with litter sizes averaging four to six pups.<ref name="cohen1978"/> Their growth rate is much faster than that of wolves, being similar in rate to that of [[coyote]]s.

The [[hormone]] metabolites of five males and three females kept in Thai zoos was studied. The breeding males showed an increased level of [[testosterone]] from October to January. The [[oestrogen]] level of captive females increases for about two weeks in January, followed by an increase of [[progesterone]]. They displayed sexual behaviours during the oestrogen peak of the females.<ref>{{cite journal |author1=Khonmee, J. |name-list-style=amp |author2= Rojanasthien, S. |author3=Thitaram, C. |author4=Sumretprasong, J. |author5= Aunsusin, A. |author6=Chaisongkram, C. |author7=Songsasen, N. |title=Non-invasive endocrine monitoring indicates seasonal variations in gonadal hormone metabolites in dholes (''Cuon alpinus'') |date=2017 |journal=Conservation Physiology |volume=5 |issue=1 |pages=cox001 |doi= 10.1093/conphys/cox001 |pmid= 28852505|pmc=5570072 |doi-access=free }}</ref>

Pups are suckled at least 58 days. During this time, the pack feeds the mother at the den site. Dholes do not use [[wikt:rendezvous|rendezvous]] sites to meet their pups as wolves do, though one or more adults will stay with the pups at the den while the rest of the pack hunts. Once [[weaning]] begins, the adults of the clan will regurgitate food for the pups until they are old enough to join in hunting. They remain at the den site 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 deer|sambar]] by the age of eight months.<ref name="f80">{{Harvnb|Fox|1984|p=80}}</ref> Maximum longevity in captivity is 15–16 years.<ref name="nowak1983"/>

===Hunting behaviour===
Before embarking on a hunt, clans go through elaborate prehunt social rituals involving nuzzling, body rubbing and mounting.<ref name="f100">{{Harvnb|Fox|1984|pp=100–101}}</ref> Dholes are primarily [[Diurnality|diurnal]] hunters, hunting in the early hours of the morning. They rarely hunt at night, except on moonlit nights, indicating they greatly rely on [[Visual perception|sight]] when hunting.<ref name="f50">{{Harvnb|Fox|1984|p=50}}</ref> They can chase their prey for many hours.<ref name="heptner1998"/> During a pursuit, one or more dholes takes over chasing the prey, while the rest of the pack keeps up at a steadier pace behind, taking over once the other group tires. Most chases are short, lasting only {{cvt|500|m}}.<ref name="f73">{{Harvnb|Fox|1984|p=73}}</ref> When chasing fleet-footed prey, they run at a pace of {{cvt|30|mph|km/h|-1|order=flip}}.<ref name="heptner1998"/> Dholes frequently drive their prey into water bodies, where the targeted animal's movements are hindered.<ref name="f67">{{Harvnb|Fox|1984|p=67}}</ref>

[[File:2012-bandipur-dhole-sambar.jpg|thumb|Dholes attacking a [[sambar (deer)|sambar]], [[Bandipur National Park]]]]

Once large prey is caught, one dhole grabs the prey's nose, while the rest of the pack pulls the animal down by the flanks and hindquarters. They do not use a killing bite to the throat.<ref name="f61">{{Harvnb|Fox|1984|p=61}}</ref> They occasionally blind their prey by attacking the eyes.<ref>{{cite journal |author=Grassman, L. I. Jr. |author2=M. E. Tewes |author3=N. J. Silvy |author4=K. Kreetiyutanont |year=2005 |title=Spatial ecology and diet of the dhole ''Cuon alpinus'' (Canidae, Carnivora) in north central Thailand |name-list-style= amp| journal=Mammalia |volume=69 |issue=1 |pages=11–20 |s2cid= 85198149 |doi= 10.1515/mamm.2005.002 |url= http://www.mnhn.fr/museum/front/medias/publication/6555_m05n1a2 |url-status=dead |archive-url=https://web.archive.org/web/20061123090326/http://www.mnhn.fr/museum/front/medias/publication/6555_m05n1a2 |archive-date=23 November 2006}}</ref> [[Serow]]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.<ref name="lydekker1907"/> Dholes tear open their prey's flanks and [[Disembowelment|disembowel]] it, eating the [[heart]], [[liver]], [[lungs]] and some sections of the [[intestines]]. The [[stomach]] and [[rumen]] are usually left untouched.<ref name="f63">{{Harvnb|Fox|1984|p=63}}</ref> Prey weighing less than {{cvt|50|kg|lb}} is usually killed within two minutes, while large stags may take 15 minutes to die. Once prey is secured, dholes tear off pieces of the carcass and eat in seclusion.<ref name="f70">{{Harvnb|Fox|1984|p=70}}</ref> They give the pups access to a kill.<ref name="f86">{{Harvnb|Fox|1984|pp=86–87}}</ref> They are generally tolerant of [[scavenger]]s at their kills.<ref name="f51">{{Harvnb|Fox|1984|p=51}}</ref> Both mother and young are provided with regurgitated food by other pack members.<ref name="nowak1983"/>
{{clear}}

===Feeding ecology===
Prey animals in India include [[chital]], [[sambar deer]], [[muntjac]], [[mouse deer]], [[barasingha]], [[wild boar]], [[gaur]], [[water buffalo]]es, [[banteng]], [[cattle]], [[nilgai]], [[goat]]s, [[Indian hare]]s, [[Himalayan field rat]]s and [[langur]]s.<ref name="cohen1978"/><ref name=Pocock1941/><ref name="f58">{{Harvnb|Fox|1984|pp=58–60}}</ref> There is one record of a pack bringing down an [[Indian elephant]] calf in [[Assam]], despite desperate defense of the mother, resulting in numerous losses to the pack.<ref name="perry1965"/> In Kashmir, they prey on [[markhor]],<ref name=Pocock1941/> and [[thamin]] in Myanmar,<ref name="cohen1978"/> [[Malayan tapir]], [[Sumatran serow]] in [[Sumatra]] and the [[Malay Peninsula]] and [[Javan rusa]] in Java.<ref name=Durbin2004/> In the [[Tian Shan]] and [[Tarbagatai Mountains]], dholes prey on [[Siberian ibex]]es, [[arkhar]], [[roe deer]], [[Caspian red deer]] and [[wild boar]]. In the [[Altai Mountains|Altai]] and [[Sayan Mountains]], they prey on [[musk deer]] and [[reindeer]]. In eastern Siberia, they prey on roe deer, [[Manchurian wapiti]], wild pig, musk deer and reindeer, while in [[Primorye]] they feed on [[sika deer]] and [[goral]]. In Mongolia, they prey on [[argali]] and rarely Siberian ibex.<ref name="heptner1998"/>

[[File:2012-bandipur-dhole-chital.jpg|thumb|Dholes feeding on a [[chital]], Bandipur National Park]]
[[File:Dhole feeding Khao Yai NP.jpg|thumb|Dhole feeding on [[sambar deer]] carcass, [[Khao Yai National Park]]]]

Like African wild dogs, but unlike wolves, dholes are not known to actively hunt people.<ref name="heptner1998"/><ref name=Pocock1941/> They are known to eat [[insect]]s and [[lizard]]s.<ref>{{Cite web |last=Chacon |first=Raquel |title=Cuon alpinus (dhole) |url=https://animaldiversity.org/accounts/Cuon_alpinus/ |access-date=2023-04-04 |website=Animal Diversity Web |language=en |archive-date=27 January 2023 |archive-url=https://web.archive.org/web/20230127081104/https://animaldiversity.org/accounts/cuon_alpinus/ |url-status=live }}</ref> 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.<ref name="mivart1890">{{cite book |last=Mivart |first=George |title=Dogs, Jackals, Wolves and Foxes: A Monograph of the Canidæ |publisher=R.H. Porter |year=1890 |location=London |pages=177–188 |chapter=Genus ''Cyon'', Hodgson (1838) |chapter-url=https://archive.org/stream/dogsjackalswolve00mivauoft#page/n307/mode/2up}}</ref> In summertime in the Tian Shan Mountains, dholes eat large quantities of mountain [[rhubarb]].<ref name="heptner1998"/> Although opportunistic, dholes have a seeming aversion to hunting cattle and their calves.<ref name="f71">{{Harvnb|Fox|1984|p=71}}</ref> [[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 [[cow]]s, probably because they are given less protection.<ref>{{Cite journal |last1=Johnsingh |first1=A.J.T. |last2=Yonten |first2=D. |last3=Wangchuck |first3=S. |year=2007 |title=Livestock-Dhole Conflict in Western Bhutan |url=https://www.biodiversitylibrary.org/page/48380954 |journal=Journal of the Bombay Natural History Society |volume=104 |issue=2 |pages=201–202 |via=Biodiversity Heritage Library}}</ref>

===Enemies and competitors===
[[File:Dhole killed by leopard.jpg|thumb|Dhole killed and cached in a tree by a [[leopard]], India]]
In some areas, dholes are [[wiktionary:sympatric|sympatric]] to [[tiger]]s and [[leopard]]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 {{cvt|30|–|175|kg}} range (mean weights of {{cvt|35.3|kg}} for dhole and {{cvt|23.4|kg}} for leopard), while tigers selected for prey animals heavier than {{cvt|176|kg}} (but their mean prey weight was {{cvt|65.5|kg}}). Also, other characteristics of the prey, such as sex, arboreality and aggressiveness, may play a role in prey selection. For example, dholes preferentially select male chital, whereas leopards kill both sexes more evenly (and tigers prefer larger prey altogether), dholes and tigers kill langurs rarely compared to leopards due to the leopards' greater arboreality, while leopards kill wild boar infrequently due to the inability of this relatively light predator to tackle aggressive prey of comparable weight.<ref name="jstor.org">{{cite journal |author1=Karanth, K. U. |author2=Sunquist, M. E. |jstor=5647 |title=Prey selection by tiger, leopard and dhole in tropical forests |journal=[[Journal of Animal Ecology]] |year=1995 |volume=64 |issue=4 |pages=439–450|doi=10.2307/5647 |bibcode=1995JAnEc..64..439K |name-list-style=amp}}</ref>

Tigers are dangerous opponents for dholes, as they have sufficient strength to kill a dhole with a single paw strike.<ref name="perry1965">{{cite book |author=Perry, R. |year=1964 |title=The World of the Tiger |publisher=Cassell |location=London}}</ref> Dhole packs are smaller in areas with higher tiger densities due to tigers directly killing dholes and stealing kills they made. The kleptoparasitism causes dholes to prefer hunting smaller animals because they can eat more of a smaller carcass before a tiger arrives to steal it. Direct predation can lead to lower reproductive and recruitment rates, lower hunting success rates and less food for the pups when a helper is killed, and potentially pack destabilization if one member of the breeding pair is killed.<ref>{{cite journal |last1=Bhandari |first1=Aishwarya |last2=Ghaskadbi |first2=Pallavi |last3=Nigam |first3=Parag |last4=Habib |first4=Bilal |name-list-style=and |year=2021 |title=Dhole pack size variation: Assessing the effect of prey availability and apex predator |url= |journal=Ecology and Evolution |volume=11 |issue=9 |pages=4774–4785 |bibcode=2021EcoEv..11.4774B |doi=10.1002/ece3.7380 |pmc=8093734 |pmid=33976847}}</ref>

Dhole packs may steal leopard kills, while leopards may kill dholes if they encounter them singly or in pairs.<ref name=Pocock1941/> There are numerous records of leopards being treed by dholes.<ref name="nowak1983">{{cite book |author1=Walker, E. P. |author2=Nowak, R. M. |author3=Warnick, F. |year=1983 |title=Walker's Mammals of the World |edition=Fourth |publisher=Johns Hopkins University Press |location=Baltimore |name-list-style=amp}}</ref> 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.<ref name="fin1929">{{cite book |author=Finn, F. |year=1929 |title=Sterndale's Mammalia of India |publisher=Thacker, Spink & Co. |location=London}}</ref> 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.<ref name="venkataraman">{{cite journal |author=Venkataraman, A. |year=1995 |title=Do dholes (''Cuon alpinus'') live in packs in response to competition with or predation by large cats? |journal=Current Science |volume=69 |issue=11 |pages=934–936 |url=http://eprints.iisc.ac.in/25085/1/article_id_069_11_0934_0936_0.pdf |archive-url=https://web.archive.org/web/20190713030406/http://eprints.iisc.ac.in/25085/1/article_id_069_11_0934_0936_0.pdf |archive-date=2019-07-13 |url-status=live}}</ref>

Dhole packs occasionally attack [[Asiatic black bear]]s, [[snow leopard]]s and [[sloth bear]]s. When attacking bears, dholes will attempt to prevent them from seeking refuge in caves and lacerate their hindquarters.<ref name=Pocock1941/>
Although usually antagonistic toward [[wolves]],<ref name=heptner1998/> they may hunt and feed alongside one another.<ref name=shretha1997>{{cite book |author=Shrestha, T. J. |year=1997 |title=Mammals of Nepal: (with reference to those of India, Bangladesh, Bhutan and Pakistan) |publisher=Bimala Shrestha |location=Kathmandu |isbn=978-0-9524390-6-6}}</ref>

The dhole is also sympatric with the [[Indian wolf]] (''Canis lupus pallipes'') in parts of its range.<ref>{{cite journal |author=Mukherjee, S. |name-list-style=amp |author2=Zelcer, M. |author3=Kotler, B.P. |year=2009 |title=Patch use in time and space for a meso-predator in a risky world |journal=Oecologia |volume=159 |issue=3| pages=661–668 |doi=10.1007/s00442-008-1243-3 |pmid=19082629 |bibcode=2009Oecol.159..661M |s2cid=24051254}}</ref><ref>{{cite journal |author=Afik, D. |author2=Pinshow, B. |year=1993 |title=Temperature regulation and water economy in desert wolves |journal=Journal of Arid Environments |volume=24 |issue=2 |pages=197–209 |doi=10.1006/jare.1993.1017 |bibcode=1993JArEn..24..197A}}</ref> There is at least one record of a lone wolf associating with a pair of dholes in [[Debrigarh Wildlife Sanctuary]],<ref>{{cite journal |author1=Nair M. V. |name-list-style=amp |author2=Panda S. K. |year=2013 |title=Just Friends |journal=Sanctuary Asia |volume=XXXIII |page=3 |url=http://www.sanctuaryasia.com/magazines/features/9390-just-friends.html |access-date=23 December 2014 |archive-date=24 July 2019 |archive-url=https://web.archive.org/web/20190724195645/https://www.sanctuaryasia.com/magazines/features/9390-just-friends.html |url-status=dead }}</ref> and two observations in [[Satpura Tiger Reserve]].<ref>{{cite journal |author1=Ghaskadbi, P. |name-list-style=amp |author2=Das, J. |author3=Mahadev, V. |author4=Habib, B. |year=2021 |title=First record of mixed species association between dholes and a wolf from Satpura Tiger Reserve, India |journal=Canid Biology & Conservation |volume=23 |issue=4 |pages=15–17 |url=https://www.canids.org/CBC/23/Dhole_wolf_association.pdf |archive-url=https://web.archive.org/web/20211017142640/https://www.canids.org/CBC/23/Dhole_wolf_association.pdf |archive-date=2021-10-17 |url-status=live}}</ref> They infrequently associate in mixed groups with [[golden jackal]]s. Domestic dogs may kill dholes, though they will feed alongside them on occasion.<ref name=humphrey1990>{{cite book |author1=Humphrey, S. R. |author2=Bain, J. R. |name-list-style=amp |year=1990 |title=Endangered Animals of Thailand |publisher=Sandhill Crane Press |location=Gainesville |isbn=978-1-877743-07-8}}</ref>

===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 [[endoparasite]]s such as [[cestode]]s and [[roundworm]]s.<ref name=Durbin2004/>

==Threats==
[[Habitat loss]] is thought to amount to 60% of the dhole's historical range in India. The fragmentation and isolation of dhole populations has resulted in inbreeding and the [[Allee effect]], which threaten its long-term viability.<ref>{{cite journal |last1=Modi |first1=S. |last2=Mondol |first2=S. |last3=Nigam |first3=P. |last4=Habib |first4=B. |date=2021 |title=Genetic analyses reveal demographic decline and population differentiation in an endangered social carnivore, Asiatic wild dog |journal=Nature |volume=11 |issue=1 |page=16371 |doi=10.1038/s41598-021-95918-3 |doi-access=free|pmid=34385570 |pmc=8361113 |bibcode=2021NatSR..1116371M }}</ref>

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.<ref name=nowak1983/> 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.<ref name=Durbin2004/> Bounties paid for dholes used to be 25 [[rupee]]s, though this was reduced to 20 in 1926 after the number of presented dhole carcasses became too numerous to maintain the established reward.<ref name=f109>{{Harvnb|Fox|1984|p=109}}</ref> The Indochinese dhole population suffers heavily from nonselective hunting techniques such as [[snaring]].<ref name=Durbin2004/>
The [[fur trade]] does not pose a significant threat to the dhole.<ref name=Durbin2004/> The people of India do not eat dhole flesh and their fur is not considered overly valuable.<ref name=mivart1890/> 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 [[ruble]]s. In the early 20th century, dhole pelts reached eight rubles in [[Manchuria]]. In [[Jetisu|Semirechye]], [[fur coat]]s made from dhole skin were considered the warmest, but were very costly.<ref name=heptner1998/>

== 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]].<ref>[http://www.thehindu.com/news/cities/Visakhapatnam/zoo-to-have-conservation-breeding-centre-for-dhole/article6326027.ece Zoo to have conservation breeding centre for ‘dhole’] {{Webarchive|url=https://web.archive.org/web/20240303095610/https://www.thehindu.com/fragment/latestnews?count=4 |date=3 March 2024 }}, ''The Hindu'' (18 August 2014)</ref> 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.<ref name=iucn/>

In 2016, the Korean company Sooam Biotech was reported to be attempting to clone the dhole using dogs as [[surrogate mother]]s to help conserve the species.<ref name="NewScientist500">{{cite magazine|last1=Zastrow|first1=M.|title=Inside the cloning factory that creates 500 new animals a day|url=https://www.newscientist.com/article/2076681-inside-the-cloning-factory-that-creates-500-new-animals-a-day/|access-date=23 February 2016|magazine=New Scientist|date=8 February 2016|archive-date=22 February 2016|archive-url=https://web.archive.org/web/20160222043834/https://www.newscientist.com/article/2076681-inside-the-cloning-factory-that-creates-500-new-animals-a-day/|url-status=live}}</ref>

==In culture and literature==
{{multiple image |perrow=1 |image1=RR5109-0073R 1 рубль серебро 2005 Красная книга Красный волк реверс.gif |caption1 = [[Russian ruble]] |image2=Coin of Kazakhstan 500RedWolf reverse.jpg |caption2=[[Kazakhstani tenge]]}}
[[File:Imperial Encyclopaedia - Animal Kingdom - pic171 - 豺圖.svg|thumb|alt=In ancient China, dholes were known and were also present in mythology|Drawing of a dhole from the Imperial Chinese Encyclopedia]]
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.<ref>van der Geer, A. A. E. (2008), ''Animals in stone: Indian mammals sculptured through time'', BRILL, p. 188, {{ISBN|90-04-16819-2}}</ref> 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]]".<ref name="perry1965"/>

[[Leopold von Schrenck]] had trouble obtaining dhole specimens during his exploration of [[Amurland]], as the local [[Gilyaks]] greatly feared the species. This fear and superstition was not, however, shared by neighbouring [[Tungusic peoples]]. It was speculated that this differing attitude towards the dhole was due to the Tungusic peoples' more nomadic, hunter-gatherer lifestyle.<ref name="schrenk1859">{{cite book |last=Schrenk |first=L. v. |author-link=Leopold von Schrenck |title=Reisen und Forschungen im Amur-lande in den jahren 1854–1856 |publisher=Kaiserliche Akademie der Wissenschaften |year=1859 |location=St. Petersburg |pages=48–50 |language=de |chapter=''Canis alpinus'' |chapter-url=https://archive.org/stream/reisenundforschu0112schr#page/48/mode/2up}}</ref>

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 {{Lang|ja|犬物語}} ({{Transliteration|ja|Inu monogatari}}; 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.<ref>Skabelund, A. H. (2011). ''Empire of Dogs: Canines, Japan, and the Making of the Modern Imperial World''. Cornell University Press, p. 85, {{ISBN|0801463246}}</ref>

A fictional version of the dhole, imbued with supernatural abilities, appears in a sixth-season 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]] ({{Lang|zh|睚眦}}), which was believed to be a creature that was part-dhole, part-dragon. In modern times, however, the Chinese word for dhole ({{Lang-zh|c=豺|p=Chái|labels=no}}) is often confused with 'jackal' or 'wolf', resulting in many confusions and mistranslations of dholes as jackals or wolves.<ref>{{Cite journal |last=Schafer |first=Edward H. |date=1991 |title=Brief Note: The Chinese Dhole |url=https://www.jstor.org/stable/41645459 |journal=Asia Major |volume=4 |issue=1 |pages=1–6 |jstor=41645459 |issn=0004-4482}}</ref>

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 kept captured dholes in captivity, and found, with the exception of one animal, they remained shy and vicious even after 10 months.<ref>{{cite journal |last1=Hodgson |first1=B.H. |title=Description of the Wild Dog of the Himalaya |journal=Asiatic Researches |date=1833 |volume=XVIII |page=226 |url=https://www.biodiversitylibrary.org/item/137991#page/584/mode/1up |access-date=14 January 2021 |archive-date=29 January 2021 |archive-url=https://web.archive.org/web/20210129081829/https://www.biodiversitylibrary.org/item/137991#page/584/mode/1up |url-status=live}}</ref> 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.<ref name=lydekker1907/> A dhole may have been presented as a gift to the Akkadian king [[Ibbi-Sin]] as tribute referred to in the inscription as the "red dog of Meluhha" or [[Indus Valley Civilisation]] of [[Pakistan]] suggesting a once greater range of the dhole.<ref>{{cite book |author=McIntosh, J. |year=2008 |title=The ancient Indus Valley: new perspectives |publisher=ABC-CLIO |isbn=978-1-57607-907-2}}</ref>

==See also==
* ''[[Wild Dog Diaries]]''

== Notes ==
{{Notelist}}

==References==
{{Reflist|refs=
<ref name=Afanasjev1935>{{cite journal |last1=Афанасьев |first1=А.В. |last2=Золотарев |first2=Н.Т. |title=Новые данные по систематике и распространению красного волка |trans-title=Contribution to the systematics and distribution of red wolf |language=ru |journal=Известия Академии наук СССР. |series=VII Отделение математических и естественных наук |year=1935 |issue=3 |pages=425–429 |url=http://mi.mathnet.ru/izv4677 |access-date=27 July 2021 |archive-date=3 March 2024 |archive-url=https://web.archive.org/web/20240303095446/https://www.mathnet.ru/php/archive.phtml?wshow=paper&jrnid=im&paperid=4677&option_lang=rus |url-status=live }}</ref>
<ref name=baryshnikov2012>{{cite journal |author=Baryshnikov, G.F. |year=2012 | title=Pleistocene Canidae (Mammalia, Carnivora) from the Paleolithic Kudaro caves in the Caucasus | journal=Russian Journal of Theriology |volume=11 |issue=2| pages=77–120 |doi=10.15298/rusjtheriol.11.2.01|doi-access=free}}</ref>

<ref name=Castello2018>{{cite book |last=Castelló |first=J.R. |title=Canids of the World: Wolves, Wild Dogs, Foxes, Jackals, Coyotes, and Their Relatives |publisher=[[Princeton University Press]] |year=2018 |chapter=Ch.2 - Wolf-like Canids |pages=155 |isbn=978-0-691-18372-5 |chapter-url=https://books.google.com/books?id=islgDwAAQBAJ&pg=PA155 |access-date=14 June 2021 |archive-date=14 June 2021 |archive-url=https://web.archive.org/web/20210614010200/https://books.google.com/books?id=islgDwAAQBAJ&pg=PA155 |url-status=live }}</ref>
<ref name=Hardwicke1821>{{cite journal |author=Hardwicke, T. |journal=Transactions of the Linnean Society of London |volume=3 |pages=235–238 |year=1821 |title=Descriptions of the Wild Dog of Sumatra, a new Species of Viverra, and a new Species of Pheasant |url=https://www.biodiversitylibrary.org/page/754977#page/263/mode/1up |access-date=3 March 2024 |archive-date=23 October 2023 |archive-url=https://web.archive.org/web/20231023225157/https://www.biodiversitylibrary.org/page/754977#page/263/mode/1up |url-status=live }}</ref>

<ref name=Heude1892>Heude, Mém. Hist. Nat. Empire Chinois, II, pt2, p. 102 footnote 1892</ref>

<ref name="kurten1968">{{cite book|author=Kurtén, B. |author-link=Bjorn Kurten|year=1968 |publisher=Weidenfeld and Nicolson |pages=111–114 |url=https://books.google.com/books?id=OsPBXSNL8ZkC&pg=PA111 |title=Pleistocene mammals of Europe|isbn=9781412845144}}</ref>
<ref name=moulle2006>{{cite journal |author1=Moulle, P.E. |author2=Echassoux, A. |author3=Lacombat, F. |year=2006 |title=Taxonomie du grand canidé de la grotte du Vallonnet (Roquebrune-Cap-Martin, Alpes-Maritimes, France) |journal=L'Anthropologie |volume=110 |issue=5 |pages=832–836 |url=http://cat.inist.fr/?aModele=afficheN&cpsidt=18357587 |doi=10.1016/j.anthro.2006.10.001 |access-date=30 July 2017 |archive-date=14 March 2012 |archive-url=https://web.archive.org/web/20120314061933/http://cat.inist.fr/?aModele=afficheN&cpsidt=18357587 |url-status=dead }} (in French)</ref>

<ref name=Orrell2019>{{cite web|url=http://www.catalogueoflife.org/annual-checklist/2019/details/species/id/59a93bdf80e76056b2e314cf0b036f9c/source/tree|title=Cuon alpinus (Pallas, 1811) (accepted name)|last1=Orrell|first1=T.|veditors=Nicolson D, Roskov Y, Abucay L, Orrell T, Nicolson D, Bailly N, Kirk PM, Bourgoin T, DeWalt RE, Decock W, De Wever A, van Nieukerken E, Zarucchi J, Penev L|year=2020|website=Catalogue of Life: 2019 Annual Checklist|publisher=Catalogue of Life|access-date=5 February 2020|archive-date=6 February 2020|archive-url=https://web.archive.org/web/20200206094126/http://www.catalogueoflife.org/annual-checklist/2019/details/species/id/59a93bdf80e76056b2e314cf0b036f9c/source/tree|url-status=live}}</ref>

<ref name=Pocock1936>{{cite journal|doi=10.1111/j.1096-3642.1936.tb02278.x|title=The Asiatic Wild Dog or Dhole (''Cuon javanicus'')|journal=Proceedings of the Zoological Society of London|volume=106|pages=33–55 |year=1936|last1=Pocock|first1=R.I.}}</ref>

<ref name="wang2008">{{Cite book |last1=Wang |first1=X. |title=Dogs: Their Fossil Relatives and Evolutionary History |last2=Tedford |first2=R.H. |publisher=Columbia University Press |year=2008 |isbn=9780231509435 |location=New York |doi=10.7312/wang13528}}</ref>

<ref name=Wozencraft2004>{{cite book|last1=Wozencraft|first1=C. W.|title=Mammal Species of the World: A Taxonomic and Geographic Reference|publisher=Johns Hopkins University Press|editor1-last=Wilson|editor1-first=D. E.|editor2-last=Reader|editor2-first=D. M.|volume=1|edition=3rd|year=2005|chapter=Order Carnivora|pages=578|isbn=978-0-8018-8221-0|chapter-url=https://books.google.com/books?id=JgAMbNSt8ikC&pg=PA578|access-date=3 March 2024|archive-date=3 March 2024|archive-url=https://web.archive.org/web/20240303095624/https://books.google.com/books?id=JgAMbNSt8ikC&pg=PA578#v=onepage&q&f=false|url-status=live}}</ref>  
}}

===Bibliography===
*{{cite book |last=Fox |first=M.W.|title=The Whistling Hunters: Field Studies of the Asiatic Wild Dog (''Cuon alpinus'') |publisher=State University of New York Press |year=1984 |location=Albany |isbn=978-0-9524390-6-6 |url=https://books.google.com/books?id=V8qMs6x-PhUC}}   
*{{cite book |last=Karanth|first=U.K.|title=A View From the Machan|publisher=Permanent Black |year=2006 |location=Karnataka |isbn=978-8-1782413-7-1 |url=https://books.google.com/books?id=W6ks4b0l7NgC}}

==External links==
{{Wiktionary}}
*[http://www.cuon.net/dholes/ Dhole Home Page] ([https://web.archive.org/web/20170919112743/http://www.cuon.net:80/dholes Archive])
*ARKive – [https://web.archive.org/web/20060313170514/http://www.arkive.org/species/GES/mammals/Cuon_alpinus/ images and movies of the dhole]
*[https://www.theguardian.com/environment/radical-conservation/2015/jun/25/dhole-asia-endangered-tiger-ignored Saving the dhole: The forgotten 'badass' Asian dog more endangered than tigers], ''The Guardian'' (25 June 2015)
*[https://web.archive.org/web/20110608190250/http://public.fotki.com/msandeep/wild-dogs---bandipur/ Photos of dhole in Bandipur]

{{Carnivora|Ca.}}
{{Canidae extinct nav|W.}}
{{Taxonbar|from=Q132585}}
{{Authority control}}

[[Category:Dhole| ]]
[[Category:Canina (subtribe)]]
[[Category:Taxa named by Peter Simon Pallas]]
[[Category:Mammals described in 1811]]
[[Category:Carnivorans of Asia]]
[[Category:Mammals of South Asia]]
[[Category:Mammals of Southeast Asia]]
[[Category:Endangered fauna of Asia]]
[[Category:Pleistocene mammals of Asia]]
[[Category:Extant Middle Pleistocene first appearances]]
[[Category:Species that are or were threatened by habitat fragmentation]]
[[Category:Endangered Fauna of China]]