Editing Leopard cat (section)

==Taxonomy==
[[File:Tsushima Cat 001.jpg|thumb|Tsushima leopard cat]]

''Felis bengalensis'' was the [[scientific name]] proposed by [[Robert Kerr (writer)|Robert Kerr]] in 1792 for a leopard cat from [[Bengal]].<ref>{{cite book |last1=Kerr |first1=R. |title=The Animal Kingdom or zoological system of the celebrated Sir Charles Linnaeus. Class I. Mammalia |date=1792 |publisher=A. Strahan & T. Cadell |location=Edinburgh & London |page=151−152 |chapter=Bengal Tiger-Cat ''Felis bengalensis'' |chapter-url=https://archive.org/details/animalkingdomorz00linn/page/150}}</ref> In the subsequent decades, 20 more leopard cat [[zoological specimen|specimen]]s were described and named, including:<ref name="ems66">{{cite book |author1=Ellerman, J. R. |author2=Morrison-Scott, T. C. S. |year=1966 |title=Checklist of Palaearctic and Indian mammals 1758 to 1946 |edition= Second |publisher=British Museum of Natural History |location=London |pages=312–313 |chapter=''Felis bengalensis'' Kerr, 1792 Leopard Cat |chapter-url=https://archive.org/stream/checklistofindia00elle#page/312/mode/2up}}</ref>

* ''Felis nipalensis'' ([[Thomas Horsfield|Horsfield]] & [[Nicholas Aylward Vigors|Vigors]], 1829) from Nepal
* ''Felis chinensis'' ([[John Edward Gray|Gray]], 1837) from [[Guangdong|Canton Province]], China
* ''Leopardus ellioti'' (Gray, 1842) from the area of [[Bombay Presidency]]
* ''Felis horsfieldi'' (Gray, 1842) from [[Bhutan]]
* ''Felis wagati'' (Gray, 1867) and ''Felis tenasserimensis'' (Gray, 1867) from [[Tanintharyi Region|Tenasserim]]
* ''Felis microtis'' ([[Alphonse Milne-Edwards|Milne-Edwards]], 1872) from the [[Peking]] area; and also from [[Tsushima Island]].<ref>{{cite journal |last1=Thomas |first1=O. |year=1908 |title=The Duke of Bedford's zoological exploration in Eastern Asia. – VII List of mammals from the Tsushima Islands |journal=Proceedings of the Zoological Society of London |volume=1908 |issue=January – April |pages=47–54 |doi=10.1111/j.1096-3642.1908.tb01833.x |url=https://archive.org/stream/proceedingsofzoo19081430zool#page/52/mode/2up}}</ref>
* ''Felis euptilura'' ([[Daniel Giraud Elliot|Elliot]], 1871) based on two skins from Siberia. One was depicted in [[Gustav Radde]]'s illustration cum description of a wild cat; the other was part of a collection at the [[Regent's Park Zoo]]. The ground colour of both was light brownish-yellow, strongly mixed with grey and covered with reddish-brown spots, head grey with a dark-red stripe across the cheek.<ref>{{cite journal |last1=Elliott |first1=D. G. |year=1871 |title=Remarks on Various Species of Felidae, with a Description of a Species from North-Western Siberia |journal=Proceedings of the Scientific Meetings of the Zoological Society of London |pages=765–761 |url=https://archive.org/stream/proceedingsofgen71zool#page/n939/mode/2up}}</ref> The initial binomial ''euptilura'' given by Elliott has been incorrectly changed to "''euptilurus''" by some later authors, but under the [[International Code of Zoological Nomenclature]] Article 31.2.1, nouns and noun phrases are not subject to gender agreement;<ref>{{cite web|url=http://www.nhm.ac.uk/hosted-sites/iczn/code/|title=International Code of Zoological Nomenclature|website=www.nhm.ac.uk|access-date=2017-03-07|archive-date=2015-12-26|archive-url=https://web.archive.org/web/20151226171112/http://www.nhm.ac.uk/hosted-sites/iczn/code/|url-status=live}}</ref> at present, both terms appear in use, but only the spelling "''euptilura''" is correct.
* ''Felis manchurica'' ([[Tamezo Mori|Mori]], 1922) from the vicinity of [[Mukden]] in Manchuria was a light grey spotted skin.<ref>{{cite journal |last1=Mori |first1=T. |year=1922 |title=On some new Mammals from Korea and Manchuria |journal=The Annals and Magazine of Natural History: Including Zoology, Botany, and Geology |series=Ninth  |volume=X |pages=609–610 |url=https://archive.org/stream/ser9annalsmagazi10londuoft#page/608/mode/2up}}</ref>
In 1939, [[Reginald Innes Pocock]] subordinated them to the genus ''[[Prionailurus]]''. The collection of the [[Natural History Museum, London]] comprised several skulls and large numbers of skins of leopard cats from various regions. Based on this broad variety of skins, he proposed to differentiate between a southern [[subspecies]] ''P. bengalensis bengalensis'' from warmer [[latitude]]s to the west and east of the [[Bay of Bengal]], and a northern ''P. bengalensis horsfieldi'' from the [[Himalayas]], having a fuller winter coat than the southern. His description of leopard cats from the areas of [[Gilgit]] and [[Karachi]] under the [[trinomen]] ''Prionailurus bengalensis trevelyani'' is based on seven skins that had longer, paler and more greyish fur than those from the Himalayas. He assumed that ''trevelyani'' inhabits more rocky, less forested habitats than ''bengalensis'' and ''horsfieldi''.<ref name="pocock">{{cite book |author=Pocock, R. I. |year=1939 |title=The Fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1 |publisher=Taylor and Francis, Ltd. |location=London |chapter=''Prionailurus bengalensis'' |pages=266–276 |chapter-url=https://archive.org/stream/PocockMammalia1/pocock1#page/n339/mode/2up}}</ref>

Two more subspecies were proposed and described:

* ''P. b. alleni'' ([[:nl:Henri Jacob Victor Sody|Sody]], 1949) from [[Hainan]] Island<ref>{{cite journal |author=Sody, H. J. V. |year=1949 |title=Notes on some Primates, Carnivora and the babirusa from the Indo-Malayan and Indo-Australian regions |journal=Treubia |volume=20 |pages=121–190}}</ref>
* [[Iriomote cat]] ''P. b. iriomotensis'' (Imaizumi, 1967) from the island of [[Iriomote]], one of the [[Ryukyu Islands]] in the [[Japanese Archipelago]];<ref name="imaizumi67">{{cite journal |author=Imaizumi, Y. |year=1967 |title=A new genus and species of cat from Iriomote, Ryukyu Islands |journal=Journal of Mammalian Society Japan |volume=3 |issue=4 |page=74}}</ref> Initially, the Iriomote cat was recognised as a distinct species, but following [[mtDNA]] analysis in the 1990s was considered a leopard cat subspecies.<ref name="masuda95">{{cite journal |last1=Masuda |first1=R. |last2=Yoshida |first2=M. C. |year=1995 |title=Two Japanese wildcats, the Tsushima cat and the Iriomote cat, show the same mitochondrial DNA lineage as the leopard cat ''Felis bengalensis'' |journal=Zoological Science |volume=12 |issue=5 |pages=655–659 |doi=10.2108/zsj.12.655 |pmid=8590833|s2cid=31111378 |doi-access=free }}</ref>

In the 1970s and 1980s, the Russian zoologists [[Vladimir Geptner|Geptner]], Gromov and Baranova disagreed with this classification. They emphasized the differences of skins and skulls at their disposal and the ones originating in Southeast Asia, and coined the term '''Amur forest cat''', which they regarded as a distinct species.<ref>{{cite journal |author=Heptner, V. G. |year=1971 |title=[On the systematic position of the Amur forest cat and some other east Asian cats placed in ''Felis bengalensis'' Kerr, 1792.] |journal=Zoologicheskii Zhurnal |volume=50 |pages=1720–1727 |language=ru}}</ref><ref>Gromov, I.M., Baranova, G.I., Baryšnikov, G. F. (eds.) (1981). ''Katalog mlekopitaûŝih SSSR : pliocen--sovremennostʹ'' Zoologičeskij Institut "Nauka." Leningradskoe otdelenie, Leningrad</ref> In 1987, Chinese zoologists pointed out the affinity of leopard cats from northern China, Amur cats and leopard cats from southern latitudes. In view of the [[Morphology (biology)|morphological]] similarities they did not support classifying the Amur cat as a species.<ref>Gao, Y.; Wang, S.; Zhang, M.L.; Ye, Z.Y.; Zhou, J.D.; eds. (1987). [Fauna Sinica. Mammalia 8: Carnivora.] Science Press, Beijing. (in Chinese)</ref>

[[Molecular genetics|Molecular analysis]] of 39 leopard cat tissue samples clearly showed three [[clade]]s: a northern lineage and southern lineages 1 and 2. The northern lineage comprises leopard cats from [[Tsushima Islands]], the [[Korean Peninsula]], the continental [[Far East]], Taiwan, and [[Iriomote Island]]. Southern lineage 1, comprising Southeast Asian populations, showed higher [[genetic diversity]]. Southern lineage 2 is genetically distant from the other lineages.<ref name=Tamada>{{cite journal |author1=Tamada, T. |author2=Siriaroonrat, B. |author3=Subramaniam, V. |author4=Hamachi, M. |author5=Lin, L.-K. |author6=Oshida, T. |author7=Rerkamnuaychoke, W. |author8=Masuda, R. |year=2006 |title=Molecular Diversity and Phylogeography of the Asian Leopard Cat, ''Felis bengalensis'', Inferred from Mitochondrial and Y-Chromosomal DNA Sequences |journal=Zoological Science |volume=25 |issue=2 |pages=154–163 |doi=10.2108/zsj.25.154 |pmid=18533746 |citeseerx=10.1.1.332.7592 |s2cid=16057327 |url=http://www2.thu.edu.tw/~biology/files/writing_journal/12/121_808f5b32.pdf |access-date=2011-02-19 |archive-date=2011-07-21 |archive-url=https://web.archive.org/web/20110721081858/http://www2.thu.edu.tw/~biology/files/writing_journal/12/121_808f5b32.pdf |url-status=live }}</ref>

Following a revision of Felidae taxonomy in 2017, two leopard cat [[species]] are now recognised, based on molecular analyses, morphological differences, and biogeographic separation:<ref name=catsg/>

* the mainland leopard cat (''P. bengalensis'') is widely distributed on mainland Asia, from Pakistan to Southeast Asia, China, and the [[Russian Far East]].
* the [[Sunda leopard cat]] (''P. javanensis'') is native to Java, Bali, Borneo, Sumatra, Palawan, Negros, Cebu, Panay, and possibly the [[Malay Peninsula]].

Two mainland leopard cat subspecies are currently recognised:<ref name=catsg/>

* ''P. b. bengalensis''  (Kerr, 1792) ranges in South and East Asia, from Pakistan to China, and probably the Malay Peninsula; and 
* ''P. b. euptilura'' (Elliott, 1871) is native to the Russian Far East, [[Manchuria]], Korea, Taiwan, Iriomote and Tsushima Islands.

=== Phylogeny ===
[[Phylogenetic]] analysis of the [[nuclear DNA]] in tissue samples from all Felidae species revealed that the [[evolutionary radiation]] of the Felidae began in Asia in the [[Miocene]] around {{mya|14.45|8.38}}.<ref name=Johnson2006>{{cite journal |last1=Johnson |first1=W. E. |last2=Eizirik |first2=E. |last3=Pecon-Slattery |first3=J. |last4=Murphy |first4=W. J. |author5=Antunes, A. |author6=Teeling, E. |author7=O'Brien, S. J. |name-list-style=amp |year=2006 |doi=10.1126/science.1122277 |title=The Late Miocene radiation of modern Felidae: A genetic assessment |journal=[[Science (journal)|Science]] |volume=311 |pages=73–77 |pmid=16400146 |issue=5757 |url=https://zenodo.org/record/1230866 |bibcode=2006Sci...311...73J |s2cid=41672825 |access-date=2020-04-30 |archive-date=2020-10-04 |archive-url=https://web.archive.org/web/20201004075725/https://zenodo.org/record/1230866 |url-status=live }}</ref><ref name=Werdelin2010>{{cite book |last1=Werdelin |first1=L. |last2=Yamaguchi |first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien |first4=S. J. |name-list-style=amp |chapter=Phylogeny and evolution of cats (Felidae) |year=2010 |pages=59–82 |chapter-url=https://www.researchgate.net/publication/266755142 |editor1-last=Macdonald |editor1-first=D. W. |editor2-last=Loveridge |editor2-first=A. J. |title=Biology and Conservation of Wild Felids |publisher=Oxford University Press |location=Oxford, UK |isbn=978-0-19-923445-5 |access-date=2020-04-30 |archive-date=2018-09-25 |archive-url=https://web.archive.org/web/20180925141956/https://www.researchgate.net/publication/266755142 |url-status=live }}</ref> Analysis of [[mitochondrial DNA]] of all Felidae species indicates a radiation at around {{mya|16.76|6.46}}.<ref name=Li_al2016>{{cite journal |author=Li, G. |author2=Davis, B. W. |author3=Eizirik, E. |name-list-style=amp |author4=Murphy, W. J. |year=2016 |title=Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae) |journal=Genome Research |volume=26 |issue=1 |pages=1–11 |doi=10.1101/gr.186668.114 |pmid=26518481 |pmc=4691742}}</ref>

The ''Prionailurus'' species are estimated to have had a [[common ancestor]] between {{mya|8.16|4.53}},<ref name=Johnson2006/> and {{mya|8.76|0.73}}.<ref name=Li_al2016/>
Both models agree in the [[rusty-spotted cat]] (''P. rubiginosus'') having been the first cat of this [[evolution]]ary [[Lineage (genetic)|lineage]] that [[Genetic divergence|genetically diverged]], followed by the [[flat-headed cat]] (''P. planiceps'') and then the [[fishing cat]] (''P. viverrinus'').<ref name=Johnson2006/><ref name=Li_al2016/> It is estimated to have diverged together with the leopard cat between {{mya|4.31|1.74}}<ref name=Johnson2006/> and {{mya|4.25|0.02}}.<ref name=Li_al2016/>

The following [[cladogram]] shows the phylogenetic relationships of the leopard cat as derived through analysis of nuclear DNA:<ref name=Johnson2006/><ref name=Werdelin2010/>
{{cladogram |align=left |style=font-size:90%;line-height:100%;width:500px;
 |cladogram={{clade |label1=[[Felidae]]
 |1={{clade |label1=[[Felinae]]
   |1={{clade
    |1={{clade
       |1={{clade |label1=''Prionailurus''
         |1={{clade
           |1={{clade
            |1={{clade
             |1={{clade
               |1='''Leopard cat'''
               |2=Fishing cat }}
             |2=Flat-headed cat }}
           |2=Rusty-spotted cat }} }}
       |2=[[Pallas's cat]] (''O. manul'') |label2=''Otocolobus''}} }}
    |2=other Felinae lineages }}
   |2=[[Pantherinae]] }} }} }}
{{clear}}