Editing Dhole (section)

==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/>
|
|----
|}