Editing Subspecies of Canis lupus (section)

==Disputed subspecies==
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===Global===
In 2019, a workshop hosted by the [[IUCN]]/SSC Canid Specialist Group considered the New Guinea singing dog and the dingo to be feral dogs (''Canis familiaris'').<ref name=Alvares2019>{{cite report |title=Old World ''Canis'' spp. with taxonomic ambiguity: Workshop conclusions and recommendations, 28th–30th May 2019 |first1=F. |last1=Alvares |first2=W. |last2=Bogdanowicz |first3=L.A.D. |last3=Campbell |first4=R. |last4=Godinho |first5=J. |author6-link=Yadvendradev Vikramsinh Jhala |last5=Hatlauf |first6=Y.V. |last6=Jhala |first7=A. C. |last7=Kitchener|first8=K.-P. |last8=Koepfli |first9=M. |last9=Krofel |first10=P. D. |last10=Moehlman |first11=H. |last11=Senn |first12=C. |last12=Sillero-Zubiri |first13=S. |last13=Viranta |first14=G. |last14=Werhahn |year=2019 |publisher=IUCN/SSC Canid Specialist Group, Centro de Investigação em Biodiversidade e Recursos Genéticos |location=Vairão, Portugal |url=https://www.canids.org/CBC/Old_World_Canis_Taxonomy_Workshop.pdf |archive-url=https://web.archive.org/web/20200212111856/https://www.canids.org/CBC/Old_World_Canis_Taxonomy_Workshop.pdf |archive-date=2020-02-12 |url-status=live |access-date=6 March 2020}}</ref> In 2020, a literature review of canid domestication stated that modern dogs were not descended from the same ''Canis'' lineage as modern wolves, and proposed that dogs may be descended from a Pleistocene wolf closer in size to a village dog.<ref>{{cite journal|doi=10.1016/j.tree.2019.10.011|pmid=31810775|title=The History of Farm Foxes Undermines the Animal Domestication Syndrome|journal=Trends in Ecology & Evolution|volume=35|issue=2|pages=125–136|year=2020|last1=Lord|first1=Kathryn A.|last2=Larson|first2=Greger|last3=Coppinger|first3=Raymond P.|last4=Karlsson|first4=Elinor K.|doi-access=free|bibcode=2020TEcoE..35..125L }}</ref> In 2021, the [[American Society of Mammalogists]] also considered dingos a feral dog (''Canis familiaris'') population.<ref name="ASMdog"/>

===Eurasia===
[[File:Vertebrati del Molise e dell'Abruzzo (1925) C. l. lupus & C. l. italicus.jpg|thumb|Giuseppe Altobello's 1925 comparative illustration of the skulls and dentition of ''[[Canis lupus lupus|C. l. lupus]]'' ('''a''') and ''[[Canis lupus italicus|C. l. italicus]]'' ('''b'''). The distinct status of the latter is currently unrecognized by MSW3.]]

====Italian wolf====
{{Further|Italian wolf}}
The [[Italian wolf]] (or Apennine wolf) was first recognised as a distinct subspecies (''Canis lupus italicus'') in 1921 by zoologist Giuseppe Altobello.<ref name="altobello1921">{{in lang|it}} Altobello, G. (1921), ''[http://www.storiadellafauna.it/scaffale/testi/alto/Carnivo.htm Fauna dell'Abruzzo e del Molise. Mammiferi. IV. I Carnivori (Carnivora)] {{Webarchive|url=https://web.archive.org/web/20160504133849/http://www.storiadellafauna.it/scaffale/testi/alto/Carnivo.htm |date=2016-05-04 }}'', Colitti e Figlio, Campobasso, pp. 38-45</ref>  Altobello's classification was later rejected by several authors, including [[Reginald Innes Pocock]], who [[Synonym (taxonomy)|synonymised]] ''C. l. italicus'' with ''C. l. lupus''.<ref name="nowak2002">{{cite journal | last1 = Nowak | first1 = R. M. | last2 = Federoff | first2 = N. E. | year = 2002 | title = The systematic status of the Italian wolf Canis lupus | journal = Acta Theriologica | volume = 47 | issue = #3| pages = 333–338 | doi=10.1007/bf03194151| bibcode = 2002AcTh...47..333N | s2cid = 366077 }}</ref> In 2002, the noted paleontologist R.M. Nowak reaffirmed the morphological distinctiveness of the Italian wolf and recommended the recognition of ''Canis lupus italicus''.<ref name="nowak2002"/> A number of DNA studies have found the Italian wolf to be genetically distinct.<ref name="wayne1992">{{Cite journal | doi = 10.1046/j.1523-1739.1992.06040559.x| title = Mitochondrial DNA Variability of the Gray Wolf: Genetic Consequences of Population Decline and Habitat Fragmentation| journal = Conservation Biology| volume = 6| issue = #4| pages = 559–569| year = 1992| last1 = Wayne | first1 = R. K. | last2 = Lehman | first2 = N. | last3 = Allard | first3 = M. W. | last4 = Honeycutt | first4 = R. L. | bibcode = 1992ConBi...6..559W}}</ref><ref name="randi2000">{{Cite journal | doi = 10.1046/j.1523-1739.2000.98280.x| title = Mitochondrial DNA Variability in Italian and East European Wolves: Detecting the Consequences of Small Population Size and Hybridization| journal = Conservation Biology| volume = 14| issue = #2| pages = 464–473| year = 2000| last1 = Randi | first1 = E. | last2 = Lucchini | first2 = V. | last3 = Christensen | first3 = M. F. | last4 = Mucci | first4 = N. | last5 = Funk | first5 = S. M. | last6 = Dolf | first6 = G. | last7 = Loeschcke | first7 = V. | bibcode = 2000ConBi..14..464R| s2cid = 86614655}}</ref> In 2004, the genetic distinction of the Italian wolf subspecies was supported by analysis which consistently assigned all the wolf [[genotype]]s of a sample in [[Italy]] to a single group. This population also showed a unique [[mitochondrial DNA]] control-region [[haplotype]], the absence of private [[alleles]] and lower [[Heterozygous|heterozygosity]] at microsatellite [[Locus (genetics)|loci]], as compared to other wolf populations.<ref>V. LUCCHINI, A. GALOV and E. RANDI'' Evidence of genetic distinction and long-term population decline in wolves (Canis lupus) in the Italian Apennines''. Molecular Ecology (2004) 13, 523–536. [https://archive.today/20120604094904/http://www3.interscience.wiley.com/journal/118794860/abstract abstract online]</ref> In 2010, a genetic analysis indicated that a single wolf haplotype (w22) unique to the Apennine Peninsula and one of the two haplotypes (w24, w25), unique to the Iberian Peninsula, belonged to the same haplogroup as the prehistoric wolves of Europe. Another haplotype (w10) was found to be common to the Iberian peninsula and the Balkans. These three populations with geographic isolation exhibited a near lack of gene flow and spatially correspond to three glacial refugia.<ref name=pilot2010/>

The taxonomic reference ''[[Mammal Species of the World]]'' (3rd edition, 2005) does not recognize ''Canis lupus italicus''; however, [[National Center for Biotechnology Information|NCBI]]/[[Genbank]] publishes research papers under that name.<ref>{{cite web|title=NCBI search Canis lupus italicus|url=https://pubmed.ncbi.nlm.nih.gov/?term=canis+lupus+italicus}}</ref>

====Iberian wolf====
{{Further|Iberian wolf}}
The [[Iberian wolf]] was first recognised as a distinct subspecies (''Canis lupus signatus'') in 1907 by zoologist [[Ángel Cabrera (naturalist)|Ángel Cabrera]]. The wolves of the Iberian peninsula have [[Morphology (biology)|morphologically]] distinct features from other [[Eurasian wolf|Eurasian wolves]] and each are considered by their researchers to represent their own subspecies.<ref>{{cite web| url = http://www.signatus.org/docs/situation.pdf| title = The wolf in Spain}}</ref><ref>{{cite journal |doi=10.1111/j.1469-7998.2000.tb00801.x|title=Food habits and livestock depredation of two Iberian wolf packs (Canis lupus signatus) in the north of Portugal|journal=Journal of Zoology|volume=251|issue=#4|pages=457–462|year=2000 |last1=Vos|first1=J.}}</ref>

The taxonomic reference ''[[Mammal Species of the World]]'' (3rd edition, 2005) does not recognize ''Canis lupus signatus''; however, [[National Center for Biotechnology Information|NCBI]]/[[Genbank]] does list it.<ref>{{cite web|title=Canis lupus signatus|url=https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=425934}}</ref>

====Himalayan wolf====
{{Further|Himalayan wolf}}
{{Cladogram|align=right|title=[[Phylogenetic tree]] with timing in years for ''Canis lupus''{{refn|group=lower-alpha|name=mtdna_seq_desc|For a full set of supporting references, refer to note (a) in the phylotree at [[Evolution of the wolf#Wolf-like canids|Evolution of the wolf]]}}
|cladogram={{clade | style = font-size:85%;line-height:80%;width:375px;
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    |label1=31,000
    |1={{clade
     |1=[[Dog|Domestic dog]] [[File:Tibetan mastiff (transparent background).png|50 px]]
     |2=Holarctic [[Wolf|gray wolf]] [[File:Dogs, jackals, wolves, and foxes (Plate I).png|50 px]]
     }}
    |2=[[Megafaunal wolf|Late Pleistocene wolf]]† [[File:The American Museum journal (c1900-(1918)) (Aenocyon dirus) transparent.png|60 px]]
    }}
   |2=[[Indian wolf#Canis indica|Indian plains wolf]] [[File:Dogs, jackals, wolves, and foxes (Plate I).png|50 px]]
   }}
   |2=[[Himalayan wolf]] [[File:Dogs, jackals, wolves, and foxes (Plate I).png|50 px]]
  }}
 }}
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The Himalayan wolf is distinguished by its [[mitochondrial DNA]], which is basal to all other wolves. The taxonomic name of this wolf is disputed, with the species ''Canis himalayensis'' being proposed based on two limited DNA studies.<ref name=agga2007/><ref name=sharma2004/><ref name=shrotriya2012/> In 2017, a study of [[mitochondrial DNA]], [[X-chromosome]] (maternal lineage) markers and [[Y-chromosome]] (male lineage) markers found that the Himalayan wolf was genetically [[Basal (phylogenetics)|basal]] to the Holarctic grey wolf and has an association with the [[African golden wolf]].<ref name=werhahn2017/>

In 2019, a workshop hosted by the [[IUCN]]/SSC Canid Specialist Group noted that the Himalayan wolf's distribution included the Himalayan range and the Tibetan Plateau. The group recommends that this wolf lineage be known as the "Himalayan wolf" and classified as ''Canis lupus chanco'' until a genetic analysis of the holotypes is available.<ref name=Alvares2019/> In 2020, further research on the Himalayan wolf found that it warranted species-level recognition under the Unified Species Concept, the Differential Fitness Species Concept, and the Biological Species Concept. It was identified as an Evolutionary Significant Unit that warranted assignment onto the [[IUCN Red List]] for its protection.<ref>{{cite journal|doi=10.1111/jbi.13824|title=Himalayan wolf distribution and admixture based on multiple genetic markers|year=2020|last1=Werhahn|first1=Geraldine|last2=Liu|first2=Yanjiang|last3=Meng|first3=Yao|last4=Cheng|first4=Chen|last5=Lu|first5=Zhi|last6=Atzeni|first6=Luciano|last7=Deng|first7=Zhixiong|last8=Kun|first8=Shi|last9=Shao|first9=Xinning|last10=Lu|first10=Qi|last11=Joshi|first11=Jyoti|last12=Man Sherchan|first12=Adarsh|last13=Karmacharya|first13=Dibesh|last14=Kumari Chaudhary|first14=Hemanta|last15=Kusi|first15=Naresh|last16=Weckworth|first16=Byron|last17=Kachel|first17=Shannon|last18=Rosen|first18=Tatjana|last19=Kubanychbekov|first19=Zairbek|last20=Karimov|first20=Khalil|last21=Kaden|first21=Jennifer|last22=Ghazali|first22=Muhammad|last23=MacDonald|first23=David W.|last24=Sillero-Zubiri|first24=Claudio|last25=Senn|first25=Helen|journal=Journal of Biogeography|volume=47|issue=6|pages=1272–1285|doi-access=free|bibcode=2020JBiog..47.1272W }}</ref>

====Indian plains wolf====
{{Further|Indian wolf#Canis indica}}
The [[Indian wolf#Canis indica|Indian plains wolf]] is a proposed clade within the Indian wolf (''Canis lupus pallipes'') that is distinguished by its mitochondrial DNA, which is basal to all other wolves except for the Himalayan wolf. The taxonomic status of this wolf clade is disputed, with the separate species ''Canis indica'' being proposed based on two limited DNA studies.<ref name=agga2007/><ref name=sharma2004/> The proposal has not been endorsed because it relied on a limited number of museum and zoo samples that may not have been representative of the wild population, and a call for further fieldwork has been made.<ref name=shrotriya2012/>

The taxonomic reference ''Mammal Species of the World'' (3rd edition, 2005) does not recognize ''Canis indica''; however, [[National Center for Biotechnology Information|NCBI]]/[[Genbank]] lists it as a new subspecies, ''Canis lupus indica''.<ref>{{cite web|title=Canis lupus indica|url=https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=242524}}</ref>

====Southern Chinese wolf====
In 2017, a comprehensive study found that the gray wolf was present across all of mainland China, both in the past and today. It exists in southern China, which refutes claims made by some researchers in the Western world that the wolf had never existed in southern China.<ref name=wang2017/><ref name=larson2017/> This wolf has not been taxonomically classified.<ref name=smith2008/><ref name=wang2003/>

In 2019, a genomic study on the wolves of China included museum specimens of wolves from southern China that were collected between 1963 and 1988. The wolves in the study formed three clades: northern Asian wolves that included those from northern China and eastern Russia, [[Himalayan wolf|Himalayan wolves]] from the Tibetan Plateau, and a unique population from southern China. One specimen from [[Zhejiang]] Province in eastern China shared gene flow with the wolves from southern China; however, its genome was 12–14 percent admixed with a canid that may be the dhole or an unknown canid that predates the genetic divergence of the [[dhole]]. The wolf population from southern China is believed to still exist in that region.<ref name=Wang2019/>

===North America===

====Coastal wolves====
A study of the three coastal wolves indicates a close phylogenetic relationship across regions that are geographically and ecologically contiguous, and the study proposed that ''Canis lupus ligoni'' (the [[Alexander Archipelago wolf]]), ''Canis lupus columbianus'' (the [[British Columbia wolf|British Columbian wolf]]), and ''Canis lupus crassodon'' (the [[Vancouver Coastal Sea wolf]]) should be recognized as a single subspecies of ''Canis lupus'', synonymized as ''Canis lupus crassodon''.<ref name=weckworth2011/> They share the same habitat and prey species, and form one study's six identified North American [[ecotype]]s – a genetically and ecologically distinct population separated from other populations by their different types of habitat.<ref name=schweitzer2016a/><ref name=schweitzer2016b/>

====Eastern wolf====
{{Further|Eastern wolf}}
The eastern wolf has two proposals over its origin. One is that the eastern wolf is a distinct species (''C. lycaon'') that evolved in North America, as opposed to the gray wolf that evolved in the Old World, and is related to the red wolf. The other is that it is derived from admixture between gray wolves, which inhabited the Great Lakes area and coyotes, forming a hybrid that was classified as a distinct species by mistake.<ref name=wayne2016/>

The taxonomic reference ''[[Mammal Species of the World]]'' (3rd edition, 2005) does not recognize ''Canis lycaon''; however, [[National Center for Biotechnology Information|NCBI]]/[[Genbank]] does list it.<ref>{{cite web|title=Canis lycaon|url=https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=228401}}</ref> In 2021, the [[American Society of Mammalogists]] also considered ''Canis lycaon'' a valid species.<ref>{{BioRef|asm|id=1005944|title=''Canis lycaon'' |version=1.5 |access-date=17 September 2021}}</ref>

====Red wolf====
{{Further|Red wolf}}
The red wolf is an enigmatic taxon, of which there are two proposals over its origin. One is that the red wolf is a distinct species (''C. rufus'') that has undergone human-influenced admixture with coyotes. The other is that it was never a distinct species but was derived from past admixture between coyotes and gray wolves, due to the gray wolf population being eliminated by humans.<ref name=wayne2016>{{cite journal|doi=10.1111/mec.13642|pmid=27064931|title=Hybridization and endangered species protection in the molecular era|journal=Molecular Ecology|volume=25|issue=#11|pages=2680–9|year=2016|last1=Wayne|first1=Robert K.|last2=Shaffer|first2=H. Bradley|s2cid=15939116|doi-access=free|bibcode=2016MolEc..25.2680W }}</ref>

The taxonomic reference ''[[Mammal Species of the World]]'' (3rd edition, 2005) does not recognize ''Canis rufus''; however, [[National Center for Biotechnology Information|NCBI]]/[[Genbank]] does list it.<ref>{{cite web|title=Canis rufus|url=https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=45781}}</ref> In 2021, the [[American Society of Mammalogists]] also considered ''Canis rufus'' a valid species.<ref>{{BioRef|asm|id=1005945|title=''Canis rufus'' |version=1.5 |access-date=17 September 2021}}</ref>