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{{Short description|Genus within Felidae}} {{Other uses}} {{Distinguish|Pantera}} {{Use dmy dates|date=May 2024}} {{Automatic taxobox | name = ''Panthera'' | fossil_range = {{geological range|Pliocene|Present|refs=<ref name="Mazák2011"/>}} | image = 4panthera3.0.png | image_caption = From top to bottom: [[tiger]], [[lion]], [[jaguar]], [[leopard]], [[snow leopard]] | taxon = Panthera | authority = [[Lorenz Oken|Oken]], 1816<ref name="MSW3">{{MSW3 Wozencraft |id=14000227 |pages=546–548 |heading=Genus ''Panthera''}}</ref> | type_species = ''[[Leopard|Felis pardus]]''<br/>(= ''Panthera pardus'') | type_species_authority = [[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]<ref name="MSW3"/> | subdivision_ranks = Species | subdivision = {{Linked species list |{{extinct}}Panthera atrox|(Leidy, 1853) |{{extinct}}Panthera balamoides|Stinnesbeck ''et al.'', 2019 |{{extinct}}Panthera dhokpathanensis|Bakr, 1986 |{{extinct}}Panthera fossilis|(Reichenau, 1906) |{{extinct}}Panthera gombaszoegensis|(Kretzoi, 1938) |Panthera leo|(Linnaeus, 1758) |Panthera onca|(Linnaeus, 1758) |{{extinct}}Panthera palaeosinensis|(Zdansky, 1924) |Panthera pardus|(Linnaeus, 1758) |{{extinct}}Panthera principialis|Hemmer, 2023 |{{extinct}}Panthera shawi|(Broom, 1948) |{{extinct}}Panthera spelaea|(Goldfuss, 1810) |Panthera tigris|(Linnaeus, 1758) |Panthera uncia|(Schreber, 1775) |{{extinct}}Panthera youngi|Pei, 1934 |{{extinct}}Panthera zdanskyi|Mazák, Christiansen & Kitchener, 2011 }} | synonyms = {{collapsible list |title=About 10 |titlestyle=text-align:center; font-weight:normal; |bullets=y |''Jaguarius'' <small>[[Nikolai Severtzov|Severtzov]], 1858</small> |''Leo'' <small>Frisch, 1775</small> |''Leonina'' <small>Greve, 1894</small> |''Leoninae'' <small>[[Johann Andreas Wagner|Wagner]], 1841</small> |''Pardotigris'' <small>Kretzoi, 1929</small> |''Pardus'' <small>[[Leopold Fitzinger|Fitzinger]], 1868</small> |''Tigrina'' <small>Greve, 1894</small> |''Tigrinae'' <small>Wagner, 1841</small> |''Tigris'' <small>[[John Edward Gray|Gray]], 1843</small> |''Tigris'' <small>Frisch, 1775</small> }} | synonyms_ref = <ref name="MSW3"/> }} '''''Panthera'''''{{notetag|{{IPAc-en|'|p|ae|n|T|ə|r|ə|,_|p|ae|n|'|T|i:|r|ə}} {{respell|PAN|thə|rə|,_|pan|THEE|rə}}<ref name="Collins">{{Cite encyclopedia |title=Panthera |encyclopedia=[[Collins Dictionary]] |year=2005 |publisher=[[Penguin Random House]] |url=https://www.collinsdictionary.com/dictionary/english/panthera |language=en}}</ref><ref>{{Harvard citation no brackets|Eons|2021|ref=Eons2021|loc=1:30}}, spoken by Kallie Moore</ref>}} is a [[genus]] within the [[family (biology)|family]] [[Felidae]], and one of two extant genera in the subfamily [[Pantherinae]]. It contains the largest living members of the cat family. There are five living species: the [[jaguar]], [[leopard]], [[lion]], [[snow leopard]] and [[tiger]]. Numerous extinct species are also named, including the [[Panthera spelaea|cave lion]] and [[American lion]]. ==Etymology== The word {{wikt-lang|en|panther}} derives from [[Classical Latin]] {{wikt-lang|la|panthēra}}, itself from the [[Ancient Greek]] {{translit|grc|pánthēr}} ({{wikt-lang|grc|πάνθηρ}}).<ref>{{cite book |last1=Liddell, H. G. |last2=Scott, R. |name-list-style=amp |year=1940 |chapter=πάνθηρ |chapter-url=https://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3D%2377441 |title=A Greek-English Lexicon |edition=Revised and augmented |location=Oxford |publisher=Clarendon Press |access-date=21 February 2021 |archive-date=11 April 2008 |archive-url=https://web.archive.org/web/20080411203109/http://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3D%2377441 |url-status=live }}</ref> ==Characteristics== In ''Panthera'' species, the dorsal profile of the [[skull]] is flattish or evenly convex. The frontal inter[[Orbit (anatomy)|orbit]]al area is not noticeably elevated, and the area behind the elevation is less steeply sloped. The basic [[Cranial cavity|cranial]] axis is nearly horizontal. The inner chamber of the [[Auditory bulla|bullae]] is large, the outer small. The partition between them is close to the [[external auditory meatus]]. The convexly rounded chin is sloping.<ref name=Pocock1939>{{cite book |author=Pocock, R. I. |year=1939 |chapter=''Panthera'' |chapter-url=https://archive.org/stream/PocockMammalia1/pocock1#page/n243/mode/2up |title=The fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1 |publisher=Taylor and Francis |location=London |pages=196–239}}</ref> All ''Panthera'' species have an incompletely ossified [[hyoid bone]] and a specially adapted [[larynx]] with large [[vocal folds]] covered in a fibro-elastic pad; these characteristics enable them to [[roar]]. Only the snow leopard cannot roar, as it has shorter vocal folds of {{cvt|9|mm}} that provide a lower resistance to airflow; it was therefore proposed to be retained in the genus ''Uncia''.<ref>{{Cite journal |title=The larynx of roaring and non-roaring cats |journal=Journal of Anatomy |date=1989 |pmc=1256521 |pmid=2606766 |pages=117–121 |volume=163 |first=M. H.|last=Hast}}</ref> ''Panthera'' species can [[prusten]], which is a short, soft, snorting sound; it is used during contact between friendly individuals. The roar is an especially loud call with a distinctive pattern that depends on the species.<ref>{{cite journal |last1=Weissengruber |first1=G. E. |last2=Forstenpointner, G. |last3=Peters, G. |last4= Kübber-Heiss, A. |last5=Fitch, W. T. |title=Hyoid apparatus and pharynx in the lion (''Panthera leo''), jaguar (''Panthera onca''), tiger (''Panthera tigris''), cheetah (''Acinonyx jubatus'') and the domestic cat (''Felis silvestris f. catus'') |journal=Journal of Anatomy |pages=195–209 | volume=201|issue=3 |year=2002 |doi=10.1046/j.1469-7580.2002.00088.x |pmc=1570911 |pmid=12363272}}</ref> ==Evolution== The geographic origin of the genus ''Panthera'' is uncertain, though the earliest known definitive species ''[[Panthera principialis]]'' is from [[Tanzania]].<ref name=principialis/> ''P. blytheae'' from northern [[Central Asia]], originally described as the oldest known ''Panthera'' species, is suggested to be similar in skull features to the snow leopard,<ref name=Tseng_al2014/> but subsequent studies have since agreed that it is not a member of or a related species of the snow leopard lineage and that it belongs to a different genus ''[[Palaeopanthera]]''.<ref>{{cite journal |author1=Geraads, D. |author2=Peigné, S |title=Re-appraisal of ''Felis pamiri'' Ozansoy 1959 (Carnivora, Felidae) from the upper Miocene of Turkey: the earliest pantherine cat? |journal=Journal of Mammalian Evolution |volume=24 |issue=4 |pages=415–425 |year=2017 |url=https://hal.sorbonne-universite.fr/hal-01675275/document |doi=10.1007/s10914-016-9349-6|s2cid=207195894}}</ref><ref>{{cite journal|last1=Hemmer|first1=H. |date=2023 |title=The evolution of the palaeopantherine cats, ''Palaeopanthera'' gen. nov. ''blytheae'' (Tseng et al., 2014) and ''Palaeopanthera pamiri'' (Ozansoy, 1959) comb. nov. (Mammalia, Carnivora, Felidae) |journal=Palaeobiodiversity and Palaeoenvironments |volume=103 |issue=4 |pages=827–839 |doi=10.1007/s12549-023-00571-5 |bibcode=2023PdPe..103..827H |s2cid=257842190}}</ref><ref>{{Cite journal|last1=Jiangzuo |first1=Q. |last2=Madurell-Malapeira |first2=J. |last3=Li |first3=X. |last4=Estraviz-López |first4=D. |last5=Mateus |first5=O. |last6=Testu |first6=A. |last7=Li |first7=S. |last8=Wang |first8=S. |last9=Deng |first9=T. |title=Insights on the evolution and adaptation toward high-altitude and cold environments in the snow leopard lineage |year=2025 |journal=Science Advances |volume=11 |issue=3 |pages=eadp5243 |doi=10.1126/sciadv.adp5243 |pmid=39813339 |pmc=11734717 |doi-access=free }}</ref> The tiger, snow leopard, and [[clouded leopard]] [[genetic lineage]]s likely dispersed in Southeast Asia during the [[Late Miocene]].<ref name=Tseng_al2014>{{cite journal |last1=Tseng |first1=Z.J. |last2=Wang |first2=X. |last3=Slater |first3=G.J. |last4=Takeuchi |first4=G.T. |last5=Li |first5=Q. |last6=Liu |first6=J. |last7=Xie |first7=G. |name-list-style=amp |year=2014 |title=Himalayan fossils of the oldest known pantherine establish ancient origin of big cats |journal=Proceedings of the Royal Society B: Biological Sciences |volume=281 |issue=1774 |page=20132686 |doi=10.1098/rspb.2013.2686 |pmid=24225466 |pmc=3843846}}</ref> Genetic studies indicate that the [[Pantherinae|pantherine cats]] diverged from the subfamily [[Felinae]] between six and ten million years ago.<ref name=Johnson2006>{{cite journal |last1=Johnson |first1=W.E. |last2=Eizirik |first2=E. |last3=Pecon-Slattery |first3=J. |last4=Murphy |first4=W.J. |last5=Antunes |first5=A. |last6=Teeling |first6=E. |last7=O'Brien |first7=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 |volume=311 |pages=73–77 |pmid=16400146 |issue=5757 |bibcode=2006Sci...311...73J |s2cid=41672825 |url=https://zenodo.org/record/1230866 |access-date=28 June 2019 |archive-date=4 October 2020 |archive-url=https://web.archive.org/web/20201004075725/https://zenodo.org/record/1230866 |url-status=live}}</ref> The genus ''[[Neofelis]]'' is [[sister group|sister]] to ''Panthera''.<ref name=Johnson2006/><ref name=Janczewski>{{cite journal |last1=Janczewski |first1=D.N. |last2=Modi |first2=W.S. |last3=Stephens |first3=J.C. |last4=O'Brien |first4=S.J. |name-list-style=amp |year=1996 |title=Molecular evolution of mitochondrial 12S RNA and cytochrome b sequences in the pantherine lineage of ''Felidae'' |journal=Molecular Biology and Evolution |volume=12 |issue=4 |pages=690–707 |doi=10.1093/oxfordjournals.molbev.a040232 |pmid=7544865 |doi-access=free}}</ref><ref name=Johnson1997>{{cite journal |last1=Johnson |first1=W. E. |last2=O'Brien |first2=S.J. |name-list-style=amp |year=1997 |title=Phylogenetic reconstruction of the ''Felidae'' using 16S rRNA and NADH-5 mitochondrial genes |journal=Journal of Molecular Evolution |volume=44 |issue=S1 |pages=S98–S116 |doi=10.1007/PL00000060 |pmid=9071018 |bibcode=1997JMolE..44S..98J |s2cid=40185850 |url=https://zenodo.org/record/1232587 |access-date=28 June 2019 |archive-date=4 October 2020 |archive-url=https://web.archive.org/web/20201004075723/https://zenodo.org/record/1232587 |url-status=live}}</ref><ref name=Yu1>{{cite journal |last1=Yu |first1=L. |last2=Zhang |first2=Y.P. |name-list-style=amp |year=2005 |title=Phylogenetic studies of pantherine cats (Felidae) based on multiple genes, with novel application of nuclear beta-fibrinogen intron 7 to carnivores |journal=Molecular Phylogenetics and Evolution |volume=35 |issue=2 |pages=483–495 |doi=10.1016/j.ympev.2005.01.017 |pmid=15804417|bibcode=2005MolPE..35..483Y }}</ref> The clouded leopard appears to have [[Genetic divergence|diverged]] about {{Ma|8.66}}. ''Panthera'' diverged from other cat species about {{Ma|11.3}} and then evolved into the species tiger about {{Ma|6.55}}, snow leopard about {{Ma|4.63}} and leopard about {{Ma|4.35}}. Mitochondrial sequence data from fossils suggest that the [[American lion]] (''P. atrox'') is a sister lineage to ''[[Panthera spelaea]]'' (the Eurasian cave or steppe lion) that diverged about {{Ma|0.34}}, and that both ''P. atrox'' and ''P. spelaea'' are most closely related to lions among living ''Panthera'' species.<ref name="Barnett, 2009">{{cite journal |last1=Barnett |first1=R. |last2=Shapiro |first2=B. |author-link2=Beth Shapiro |last3=Barnes |first3=I. |last4=Ho |first4=S.Y.W. |last5=Burger |first5=J. |author-link5=Joachim Burger |last6=Yamaguchi |first6=N. |last7=Higham |first7=T.F.G. |last8=Wheeler |first8=H.T. |last9=Rosendahl |first9=W. |last10=Sher |first10=A.V. |last11=Sotnikova |first11=M. |last12=Kuznetsova |first12=T. |last13=Baryshnikov |first13=G.F. |last14=Martin |first14=L.D. |last15=Harington |first15=C.R. |last16=Burns |first16=J.A. |last17=Cooper |first17=A. |name-list-style=amp |title=Phylogeography of lions (''Panthera leo'' ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity |journal=[[Molecular Ecology]] |year=2009 |volume=18 |issue=8 |pages=1668–1677 |pmid=19302360 |doi=10.1111/j.1365-294X.2009.04134.x |bibcode=2009MolEc..18.1668B |s2cid=46716748 |url=https://www.zin.ru/Labs/theriology/eng/staff/baryshnikov/references/barnett_et_al_2009.pdf |access-date=24 February 2019 |archive-date=8 August 2017 |archive-url=https://web.archive.org/web/20170808225555/https://www.zin.ru/Labs/theriology/eng/staff/baryshnikov/references/barnett_et_al_2009.pdf |url-status=live}}</ref> The snow leopard is nested within ''Panthera'' and is the [[sister species]] of the tiger.<ref name=Davis2010>{{cite journal |last1=Davis |first1=B.W. |title=Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, ''Panthera'' (''Carnivora'': ''Felidae'') |author2=Li, G. |author3=Murphy, W.J. |journal=Molecular Phylogenetics and Evolution |volume=56 |date=2010 |issue=1 |pages=64–76 |pmid=20138224 |doi=10.1016/j.ympev.2010.01.036|bibcode=2010MolPE..56...64D }}</ref> Results of a 2016 study based on analysis of biparental [[nuclear genome]]s suggest the following relationships of living ''Panthera'' species:<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 extinct species ''[[Panthera gombaszoegensis]]'', was probably closely related to the modern jaguar. The first fossil remains were [[Excavation (archaeology)|excavated]] in [[Olivola]], in Italy, and date to {{Ma|1.6}}.<ref>{{cite journal |last1=Hemmer |first1=H. |last2=Kahlke |first2=R.D. |last3=Vekua |first3=A.K. |name-list-style=amp |year=2001 |title=The Jaguar – ''Panthera onca gombaszoegensis'' (Kretzoi, 1938) (Carnivora: Felidae) in the late lower Pleistocene of Akhalkalaki (south Georgia; Transcaucasia) and its evolutionary and ecological significance | journal=Geobios |volume=34 |issue=4 |pages=475–486 |doi=10.1016/s0016-6995(01)80011-5 |bibcode=2001Geobi..34..475H}}</ref> Fossil remains found in South Africa that appear to belong within the ''Panthera'' lineage date to about {{Ma|2.0|3.8}}.<ref>{{cite journal |last1=Turner |first1=A. |year=1987 |title=New fossil carnivore remains from the Sterkfontein hominid site (''Mammalia'': ''Carnivora'') |journal=Annals of the Transvaal Museum |volume=34 |issue=15 |pages=319–347 |url=https://journals.co.za/content/nfi_annalstm/34/15/AJA00411752_121?crawler=true&mimetype=application/pdf}}</ref> ==Classification== ''Panthera'' was named and described by [[Lorenz Oken]] in 1816 who placed all the spotted cats in this group.<ref>{{cite book |last1=Oken |first1=L. |title=Lehrbuch der Zoologie. 2. Abtheilung |publisher=August Schmid & Comp. |year=1816 |location=Jena |page=1052 |chapter=1. Art, ''Panthera'' |chapter-url=https://books.google.com/books?id=S5o5AAAAcAAJ&pg=PA1052}}</ref><ref name="Allen1902">{{cite journal |last1=Allen |first1=J. A. |year=1902 |title=Mammal names proposed by Oken in his 'Lehrbuch der Zoologie' |url=http://digitallibrary.amnh.org/bitstream/handle/2246/509//v2/dspace/ingest/pdfSource/bul/B016a27.pdf?sequence=1&isAllowed=y |journal=Bulletin of the American Museum of Natural History |volume=16 |issue=27 |pages=373−379 |access-date=4 April 2019 |archive-date=9 May 2023 |archive-url=https://web.archive.org/web/20230509152416/https://digitallibrary.amnh.org/bitstream/handle/2246/509//v2/dspace/ingest/pdfSource/bul/B016a27.pdf?sequence=1&isAllowed=y |url-status=live }}</ref> During the 19th and 20th centuries, various explorers and staff of [[natural history museum]]s suggested numerous subspecies, or at times called "races", for all ''Panthera'' species. The taxonomist [[Reginald Innes Pocock]] reviewed skins and skulls in the zoological collection of the [[Natural History Museum, London]], and grouped subspecies described, thus shortening the lists considerably.<ref>{{cite journal | last1=Pocock | first1=R. I. | year=1930 | title=The panthers and ounces of Asia | journal=Journal of the Bombay Natural History Society | volume = 34 | issue = 1| pages = 65–82}}</ref><ref>{{cite journal |last1=Pocock | first1 = R. I. | year=1932 |title=The leopards of Africa | doi=10.1111/j.1096-3642.1932.tb01085.x | journal=Proceedings of the Zoological Society of London | volume = 102 | issue=2| pages = 543–591}}</ref><ref>{{cite journal | last1 = Pocock | first1 = R. I. | year = 1939 | title=The races of jaguar (''Panthera onca'') | journal=Novitates Zoologicae | volume=41 | pages = 406–422}}</ref> [[Reginald Innes Pocock]] revised the classification of this genus in 1916 as comprising the [[tiger]] (''P. tigris''), [[lion]] (''P. leo''), [[jaguar]] (''P. onca''), and [[leopard]] (''P. pardus'') on the basis of common features of their [[skull]]s.<ref name="Pocock1916">{{cite journal |last1=Pocock |first1=R. I. |year=1916 |title=The Classification and Generic Nomenclature of ''F. uncia'' and its Allies |url=https://archive.org/stream/ser8annalsmagazi18londuoft#page/314/mode/2up |journal=The Annals and Magazine of Natural History: Including Zoology, Botany, and Geology |series=Series 8 |volume=XVIII |issue=105 |pages=314–316 |doi=10.1080/00222931608693854}}</ref> Since the mid-1980s, several ''Panthera'' species became subjects of [[genetic research]], mostly using blood samples of captive individuals. Study results indicate that many of the lion and leopard subspecies are questionable because of insufficient genetic distinction between them.<ref name=OBrien1987>{{cite journal |author=O'Brien, S. J. |author2=Martenson, J. S. |author3=Packer, C. |author4=Herbst, L. |author5=de Vos, V. |author6=Joslin, P. |author7=Ott-Joslin, J. |author8=Wildt, D. E. |author9=Bush, M. |name-list-style=amp |year=1987 |url=http://www.cbs.umn.edu/sites/default/files/public/downloads/Biochemical_Genetic_Variation_in_Geographic_Isolates_of_African_and_Asiatic_Lions.pdf |title=Biochemical genetic variation in geographic isolates of African and Asiatic lions |journal=National Geographic Research |volume=3 |issue=1 |pages=114–124 |archive-url=https://web.archive.org/web/20130502101107/http://www.cbs.umn.edu/sites/default/files/public/downloads/Biochemical_Genetic_Variation_in_Geographic_Isolates_of_African_and_Asiatic_Lions.pdf |archive-date=2 May 2013}}</ref><ref>{{cite journal | last1 = Miththapala | first1 = S. | last2 = Seidensticker | first2 = J. | last3 = O'Brien | first3 = S. J. | year = 1996 | title = Phylogeographic subspecies recognition in leopards (''Panthera pardu''s): Molecular genetic variation | journal = Conservation Biology | volume = 10 | issue = 4| pages = 1115–1132 | doi=10.1046/j.1523-1739.1996.10041115.x| bibcode = 1996ConBi..10.1115M }}</ref> Subsequently, it was proposed to group all African leopard populations to ''P. p. pardus'' and retain eight [[scientific names|subspecific names]] for Asian leopard populations.<ref>{{cite journal | last1 = Uphyrkina | first1 = O. | last2 = Johnson | first2 = W. E. | last3 = Quigley | first3 = H. B. | last4 = Miquelle | first4 = D. G. | last5 = Marker | first5 = L. | last6 = Bush | first6 = M. E. | last7 = O'Brien | first7 = S. J. | year = 2001 | title = ''Phylogenetics, genome diversity and origin of modern leopard,'' Panthera pardus | url = https://zenodo.org/record/1236516 | journal = Molecular Ecology | volume = 10 | issue = 11 | pages = 2617–2633 | doi = 10.1046/j.0962-1083.2001.01350.x | pmid = 11883877 | bibcode = 2001MolEc..10.2617U | s2cid = 304770 | access-date = 28 June 2019 | archive-date = 6 August 2020 | archive-url = https://web.archive.org/web/20200806044827/https://zenodo.org/record/1236516 | url-status = live }}</ref> Results of [[genetic analysis]] indicate that the [[snow leopard]] (formerly ''Uncia uncia'') also belongs to the genus ''Panthera'' (''P. uncia''), a classification that was accepted by [[IUCN Red List]] assessors in 2008.<ref name="Johnson2006" /><ref name="iucn1" /> Based on [[genetic research]], it was suggested to group all living [[sub-Sahara]]n lion populations into ''P. l. leo''.<ref>{{cite journal | last1 = Dubach | first1 = J. | last2 = Patterson | first2 = B. D. | last3 = Briggs | first3 = M. B. | last4 = Venzke | first4 = K. | last5 = Flamand | first5 = J. | last6 = Stander | first6 = P. | last7 = Scheepers | first7 = L. | last8 = Kays | first8 = R. W. | year = 2005 | title = Molecular genetic variation across the southern and eastern geographic ranges of the African lion, ''Panthera leo'' | journal = Conservation Genetics | volume = 6 | issue = 1 | pages = 15–24 | doi = 10.1007/s10592-004-7729-6 | bibcode = 2005ConG....6...15D | s2cid = 30414547 | url = https://www.researchgate.net/publication/226066591 | access-date = 5 March 2019 | archive-date = 5 March 2024 | archive-url = https://web.archive.org/web/20240305201616/https://www.researchgate.net/publication/226066591_Molecular_genetic_variation_across_the_southern_and_eastern_geographic_ranges_of_the_African_lion_Panthera_leo | url-status = live }}</ref> Results of [[phylogeographic]] studies indicate that the [[West Africa|West]]ern and [[Central Africa]]n lion populations are more closely related to those in India and form a different [[clade]] than lion populations in [[Southern Africa|Southern]] and [[East Africa]]; southeastern Ethiopia is an [[Admixture (genetics)|admixture]] region between North African and East African lion populations.<ref>{{Cite journal |last1=Bertola |first1=L. D. |last2=Van Hooft |first2=W. F. |last3=Vrieling |first3=K. |last4=Uit De Weerd |first4=D. R. |last5=York |first5=D. S. |last6=Bauer |first6=H. |last7=Prins |first7=H. H. T. |last8=Funston |first8=P. J. |last9=Udo De Haes |first9=H. A. |last10=Leirs |first10=H. |last11=Van Haeringen |first11=W. A. |last12=Sogbohossou |first12=E. |last13=Tumenta |first13=P. N. |last14=De Iongh |first14=H. H. |title=Genetic diversity, evolutionary history and implications for conservation of the lion (''Panthera leo'') in West and Central Africa |doi=10.1111/j.1365-2699.2011.02500.x |journal=[[Journal of Biogeography]] |volume=38 |issue=7 |pages=1356–1367 |year=2011 |bibcode=2011JBiog..38.1356B |s2cid=82728679 |url=http://dspace.learningnetworks.org/bitstream/1820/4311/1/2011_Bertola,Hooft,Vrieling,Weerd,York,Bauer,Prins,Haes,Iongh_GeneticDiversityEvolutionaryHistoryAndImplicationsForConservationOfTheLionInWestAndCentralAfrica.pdf |access-date=17 January 2017 |archive-url=https://web.archive.org/web/20190608070014/http://dspace.learningnetworks.org/bitstream/1820/4311/1/2011_Bertola,Hooft,Vrieling,Weerd,York,Bauer,Prins,Haes,Iongh_GeneticDiversityEvolutionaryHistoryAndImplicationsForConservationOfTheLionInWestAndCentralAfrica.pdf |archive-date=8 June 2019 }}</ref><ref>{{cite journal |author=Bertola, L. D. |author2=Jongbloed, H. |author3=Van Der Gaag, K. J. |author4=De Knijff, P. |author5=Yamaguchi, N. |author6=Hooghiemstra, H. |author7=Bauer, H. |author8=Henschel, P. |author9=White, P. A. |author10=Driscoll, C. A. |author11=Tende, T. |year=2016 |title=Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (''Panthera leo'') |journal=Scientific Reports |volume=6 |page=30807 |doi=10.1038/srep30807|pmid=27488946 |pmc=4973251 |bibcode=2016NatSR...630807B}}</ref> [[Black panther]]s do not form a distinct species, but are [[Melanism|melanistic]] specimens of the genus, most often encountered in the leopard and jaguar.<ref>{{cite journal |author = Robinson, R. |year=1970 |title=Inheritance of black form of the leopard ''Panthera pardus'' |journal=Genetica |volume=41 |issue=1 |pages=190–197 |pmid=5480762 |doi=10.1007/bf00958904|s2cid=5446868 }}</ref><ref>{{cite journal |author=Eizirik, E. |author2=Yuhki, N. |author3=Johnson, W. E. |author4=Menotti-Raymond, M. |author5=Hannah, S. S. |author6=O'Brien, S. J. |title=Molecular Genetics and Evolution of Melanism in the Cat Family |journal=Current Biology |year=2003 |volume=13 |issue=5 |pages=448–453 |doi=10.1016/S0960-9822(03)00128-3 |pmid=12620197|s2cid=19021807 |doi-access=free |bibcode=2003CBio...13..448E }}</ref> ===Contemporary species=== The following list of the genus ''Panthera'' is based on the taxonomic assessment in ''[[Mammal Species of the World]]'' and reflects the [[Taxonomy (biology)|taxonomy]] revised in 2017 by the Cat Classification Task Force of the [[IUCN Species Survival Commission#Cat Specialist Group|Cat Specialist Group]]:<ref name="MSW3"/><ref name=CatSG2017>{{cite journal |author1=Kitchener, A. C. |author2=Breitenmoser-Würsten, C. |author3=Eizirik, E. |author4=Gentry, A. |author5=Werdelin, L. |author6=Wilting, A. |author7=Yamaguchi, N. |author8=Abramov, A. V. |author9=Christiansen, P. |author10=Driscoll, C. |author11=Duckworth, J. W. |author12=Johnson, W. |author13=Luo, S.-J. |author14=Meijaard, E. |author15=O'Donoghue, P. |author16=Sanderson, J. |author17=Seymour, K. |author18=Bruford, M. |author19=Groves, C. |author20=Hoffmann, M. |author21=Nowell, K. |author22=Timmons, Z. |author23=Tobe, S. |year=2017 |title=A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group |journal=Cat News |issue=Special Issue 11 |pages=66−75 |url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y |access-date=13 May 2018 |archive-date=17 January 2020 |archive-url=https://web.archive.org/web/20200117172708/https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y |url-status=live }}</ref> {| class="wikitable" |- ! Species !! Subspecies !! [[IUCN Red List]] status and distribution |- style="vertical-align: top;" |[[Lion]] ''P. leo'' {{small|(Linnaeus, 1758)}}<ref name="Linn1758" /> [[File:Ngorongoro Crater (26) (cropped).jpg|alt=|frameless]] |[[Panthera leo leo|''P. l. leo'']] {{small|(Linnaeus, 1758)}}<ref name="Linn1758" /> including: * †[[Barbary lion]] ''P. l. leo'' ''sensu stricto'' *[[Asiatic lion]] {{small|syn. ''P. l. persica'' (Johann Nepomuk Meyer, 1826)<ref>{{cite thesis |last1=Meyer |first1=J. N. |year=1826 |title=Dissertatio inauguralis anatomico-medica de genere felium |location=Vienna |type=Doctoral thesis |publisher=University of Vienna}}</ref>}} [[Panthera leo melanochaita|''P. l. melanochaita'']] {{small|([[Charles Hamilton Smith|Smith]], 1842)<ref name=Smith1842>{{cite book |author=Smith, C. H. |year=1842 |chapter=Black maned lion ''Leo melanochaitus'' |chapter-url=https://archive.org/stream/naturalistslibra15jardrich#page/176/mode/2up |page=Plate X, 177 |title=The Naturalist's Library. Vol. 15 Mammalia |editor1-last=Jardine |editor1-first=W. |location=London |publisher=Chatto and Windus}}</ref>}} including: * †[[Cape lion]] ''P. l. melanochaita'' ''sensu stricto''<ref name="Mazak1975">{{cite journal |author=Mazak, V. |year=1975 |title=Notes on the Black-maned Lion of the Cape, ''Panthera leo melanochaita'' (Ch. H. Smith, 1842) and a Revised List of the Preserved Specimens |journal=Verhandelingen Koninklijke Nederlandse Akademie van Wetenschappen |issue=64 |pages=1–44}}</ref> ||[[Vulnerable species|{{IUCN status|VU|15951}}]]<ref>{{cite iucn |title=''Panthera leo'' |author=Bauer, H. |author2=Packer, C. |author3=Funston, P. F. |author4=Henschel, P. |author5=Nowell, K. |name-list-style=amp |page=e.T15951A115130419 |year=2016}}</ref> [[File:Lion distribution.png|frameless]] |- style="vertical-align: top;" |[[Jaguar]] ''P. onca'' {{small|(Linnaeus, 1758)}}<ref name="Linn1758" /> [[File:Jagwar (cropped).jpg|frameless]] |Monotypic<ref>{{Cite journal|last=Larson|first=S. E.|date=1997 |title=Taxonomic re-evaluation of the jaguar |journal=Zoo Biology |language=en |volume=16 |issue=2 |pages=107–120 |doi=10.1002/(SICI)1098-2361(1997)16:2<107::AID-ZOO2>3.0.CO;2-E}}</ref><ref name="CatSG2017" /> |[[Near-threatened species|{{IUCN status|NT|15953}}]]<ref>{{cite iucn|title=''Panthera onca'' |author1=Quigley, H.|author2=Foster, R.|author3=Petracca, L.|author4=Payan, E.|author5=Salom, R. |author6=Harmsen, B. |name-list-style=amp |page=e.T15953A123791436 |year=2017}}</ref> [[File:Panthera onca distribution.svg|frameless]] |- style="vertical-align: top;" |[[Leopard]] ''P. pardus'' {{small|(Linnaeus, 1758)}}<ref name="Linn1758" /> [[File:Indian male leopard (cropped).jpg|frameless]] |[[African leopard]] ''P. p. pardus'' {{small|(Linnaeus, 1758)}}<ref name="Linn1758" /><br /> [[Indian leopard]] ''P. p. fusca'' {{small|([[Friedrich Albrecht Anton Meyer|Meyer]], 1794)}}<ref>{{cite book |author=Meyer, F. A. A. |year=1794 |title=Zoologische Annalen. Erster Band |location=Weimar |publisher=Im Verlage des Industrie-Comptoirs |pages=394–396 |chapter=Über de la Metheries schwarzen Panther |chapter-url=https://books.google.com/books?id=v4s-AAAAcAAJ&pg=PA394}}</ref><br />[[Javan leopard]] ''P. p. melas'' {{small|([[Georges Cuvier|G. Cuvier]], 1809)}}<ref>{{cite journal |last=Cuvier, G. |year=1809 |title=Recherches sur les espėces vivantes de grands chats, pour servir de preuves et d'éclaircissement au chapitre sur les carnassiers fossils |journal=Annales du Muséum National d'Histoire Naturelle |volume=Tome XIV |pages=136–164}}</ref><br />[[Arabian leopard]] ''P. p. nimr'' {{small|([[Wilhelm Hemprich|Hemprich]] and [[Christian Gottfried Ehrenberg|Ehrenberg]]), 1833}}<ref>{{cite book |last1=Hemprich |first1=W. |last2=Ehrenberg |first2=C. G. |year=1830 |chapter=''Felis, pardus?, nimr'' |pages=Plate 17 |title=Symbolae Physicae, seu Icones et Descriptiones Mammalium quae ex Itinere per Africam Borealem et Asiam Occidentalem Friderici Guilelmi Hemprich et Christiani Godofredi Ehrenberg. Decas Secunda. Zoologica I. Mammalia II |location=Berolini |publisher=Officina Academica |editor=Dr. C. G. Ehrenberg |chapter-url=https://archive.org/details/SymbolaephysicaMammEhreA/page/n60}}</ref><br />[[Panthera pardus tulliana|''P. p. tulliana'']] {{small|([[Achille Valenciennes|Valenciennes]], 1856),<ref name=Valenciennes>{{cite journal |last=Valenciennes, A. |date=1856 |title=Sur une nouvelles espèce de Panthère tué par M. Tchihatcheff à Ninfi, village situé à huit lieues est de Smyrne |journal=Comptes rendus hebdomadaires des séances de l'Académie des sciences |volume=42 |pages=1035–1039}}</ref> syn. ''P. p. ciscaucasica'' ([[Konstantin Alekseevich Satunin|Satunin]], 1914),<ref>{{cite book |last=Satunin, K. A. |year=1914 |title=Opredelitel' mlekopitayushchikh Rossiiskoi Imperii |trans-title=Guide to the mammals of the Russian Empire |location=Tiflis |publisher=Tipographia Kantzelyarii Namestnichestva }}</ref> ''P. p. saxicolor'' Pocock, 1927<ref>{{cite journal |last=Pocock |first=R. I. |year=1927 |title=Description of two subspecies of leopards |journal=Annals and Magazine of Natural History |volume=20 |series=Series 9 |issue=116 |pages=213–214|doi=10.1080/00222932708655586 }}</ref>}}<br />[[Amur leopard]] ''P. p. orientalis'' {{small|([[Hermann Schlegel|Schlegel]], 1857),<ref>{{cite book |author=Schlegel, H. |year=1857 |chapter=''Felis orientalis'' |page=23 |title=Handleiding Tot de Beoefening der Dierkunde, I<sup>e</sup> Deel |publisher=Boekdrukkerij van Nys |location=Breda}}</ref> syn. ''P. p. japonensis'' ([[John Edward Gray|Gray]], 1862)<ref>{{cite journal |author=Gray, J. E. |year=1862 |title=Description of some new species of Mammalia |journal=Proceedings of the Royal Zoological Society of London |volume=30 |pages=261−263, plate XXXIII |url=https://archive.org/details/proceedingsofgen62zool/page/262|doi=10.1111/j.1469-7998.1862.tb06524.x }}</ref>}}<br />[[Indochinese leopard]] ''P. p. delacouri'' {{small|Pocock, 1930}}<ref>{{cite journal |author=Pocock, R. I. |year=1930 |title=The Panthers and Ounces of Asia |journal=Journal of the Bombay Natural History Society |volume=34 |issue=2 |pages=307–336}}</ref><br />[[Sri Lankan leopard]] ''P. p. kotiya'' {{small|Deraniyagala, 1956}}<ref>{{cite journal |last=Deraniyagala |first=P. E. P. |year=1956 |title=The Ceylon leopard, a distinct subspecies |journal=Spolia Zeylanica |volume=28 |pages=115–116}}</ref> |{{IUCN status|VU|15954}}<ref>{{cite iucn |title=''Panthera pardus'' |author=Stein, A. B. |author2=Athreya, V. |author3=Gerngross, P. |author4=Balme, G. |author5=Henschel, P. |author6=Karanth, U. |author7=Miquelle, D. |author8=Rostro, S. |author9=Kamler, J. F. |author10=Laguardia, A. |name-list-style=amp |page=e.T15954A102421779|date=2016}}</ref> [[File:Leopard distribution.jpg|frameless]] |- style="vertical-align: top;" |[[Tiger]] ''P. tigris'' {{small|(Linnaeus, 1758)}}<ref name=Linn1758>{{cite book |author=Linnaeus, C. |year=1758 |title=Caroli Linnæi Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis |volume=Tomus I |edition=decima, reformata |location=Holmiae |publisher=Laurentius Salvius |pages=41−42 |chapter=''Felis'' |chapter-url=https://archive.org/stream/mobot31753000798865#page/41/mode/2up}}</ref> <span style="{{MirrorH}}">[[File:Sher Khan (cropped).jpg|frameless]]</span> |style="white-space: nowrap;"|[[Panthera tigris tigris|''P. t. tigris'']] {{small|(Linnaeus, 1758)}} including: * [[Bengal tiger]] ''P. t. tigris'' ''sensu stricto'' * †[[Caspian tiger]] {{small|[[syn.]] ''P. t. virgata'' ([[Johann Karl Wilhelm Illiger|Illiger]], 1815)<ref>{{cite journal |last1=Illiger |first1=C. |year=1815 |title=Überblick der Säugethiere nach ihrer Verteilung über die Welttheile |journal=Abhandlungen der Königlichen Preußischen Akademie der Wissenschaften zu Berlin |volume=1804−1811 |pages=39−159 |url=http://bibliothek.bbaw.de/bbaw/bibliothek-digital/digitalequellen/schriften/anzeige/index_html?band=07-abh/18041811&seite:int=195 |access-date=2 January 2021 |archive-date=8 June 2019 |archive-url=https://web.archive.org/web/20190608070026/http://bibliothek.bbaw.de/bbaw/bibliothek-digital/digitalequellen/schriften/anzeige/index_html?band=07-abh%2F18041811&seite%3Aint=195 }}</ref>}} * [[Siberian tiger]] {{small|syn. ''P. t. altaica'' ([[Coenraad Jacob Temminck|Temminck]], 1844)<ref name=Temminck>{{cite book |last=Temminck, C. J. |year=1844 |chapter=Aperçu général et spécifique sur les Mammifères qui habitent le Japon et les Iles qui en dépendent |title=Fauna Japonica sive Descriptio animalium, quae in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Batava imperium tenent, suscepto, annis 1825–1830 collegit, notis, observationibus et adumbrationibus illustravit Ph. Fr. de Siebold |location=Leiden |publisher=Lugduni Batavorum |editor1=Siebold, P. F. v. |editor2=Temminck, C. J. |editor3=Schlegel, H. |chapter-url=https://archive.org/details/faunajaponicasi00sieb/page/43}}</ref>}} * [[South China tiger]] {{small|syn. ''P. t. amoyensis'' ([[Max Hilzheimer|Hilzheimer]], 1905)<ref>{{cite journal |last1=Hilzheimer |first1=M. |year=1905 |title=Über einige Tigerschädel aus der Straßburger zoologischen Sammlung |journal=Zoologischer Anzeiger |volume=28 |pages=594–599 |url=https://archive.org/stream/zoologischeranze28deut#page/594/mode/2up}}</ref>}} * [[Indochinese tiger]] {{small|syn. ''P. t. corbetti'' [[Vratislav Mazák|Mazák]], 1968<ref>{{cite journal |last1=Mazák |first1=V. |year=1968 |title=Nouvelle sous-espèce de tigre provenant de l'Asie du sud-est |journal=Mammalia |volume=32 |issue=1 |pages=104−112 |doi=10.1515/mamm.1968.32.1.104|s2cid=84054536 }}</ref>}} * [[Malayan tiger]] {{small|syn. ''P. t. jacksoni'' Luo et al., 2004<ref>{{cite journal |last1=Luo, S. J. |last2=Kim, J. H. |last3=Johnson, W. E. |last4=Walt, J. v. d. |last5=Martenson, J. |last6=Yuhki, N. |last7=Miquelle, D. G. |year=2004 |title=Phylogeography and Genetic Ancestry of Tigers (''Panthera tigris'') |journal=PLOS Biology |volume=2 |issue=12 |pages=e442 |doi=10.1371/journal.pbio.0020442 |pmid=15583716 |pmc=534810 |doi-access=free }}</ref>}} [[Sunda Island tiger]] ''P. t. sondaica'' {{small|Temminck, 1844)<ref name=Temminck/>}} including * †[[Javan tiger]] ''P. t. sondaica'' ''sensu stricto'' * [[Sumatran tiger]] {{small|syn. ''P t. sumatrae'' (Temminck, 1844)<ref name=Temminck/>}} * †[[Bali tiger]] {{small|syn. ''P. t. balica'' [[Ernst Schwarz (zoologist)|Schwarz]], 1912)<ref>{{cite journal |last=Schwarz |first=E. |year=1912 |title=Notes on Malay tigers, with description of a new form from Bali |journal=Annals and Magazine of Natural History |pages=324–326 |volume=Series 8 Volume 10 |issue=57 |url=https://archive.org/stream/annalsmagazineof8101912lond#page/324/mode/2up |doi=10.1080/00222931208693243}}</ref>}} ||[[Endangered species|{{IUCN status|EN|15955}}]]<ref>{{Cite iucn |title=''Panthera tigris'' |author1=Goodrich, J. |author2=Lynam, A. |author3=Miquelle, D. |author4=Wibisono, H. |author5=Kawanishi, K. |author6=Pattanavibool, A. |author7=Htun, S. |author8=Tempa, T. |author9=Karki, J. |author10=[[Yadvendradev Vikramsinh Jhala|Jhala]], Y. |author11=Karanth, U. |name-list-style=amp | page=e.T15955A50659951 |date=2015}}</ref> [[File:Tiger map.svg|frameless]] |- style="vertical-align: top;" |[[Snow leopard]] ''P. uncia''<ref name=CatSG2017/> {{small|([[Johann Christian Daniel von Schreber|Schreber]], 1775)}}<ref>{{cite book |last1=Schreber |first1=J. C. D. |chapter=Die Unze |title=Die Säugethiere in Abbildungen nach der Natur mit Beschreibungen |date=1777 |pages=386–387 |location=Erlangen |publisher=Wolfgang Walther |chapter-url=https://archive.org/stream/SaYugthiereAbbiIIISchr#page/386/mode/2up}}</ref> [[File:Snow Leopards - Ghost of the Mountains for Ministry of Tourism (MOT) India 9 (cropped).png|frameless]] |[[Monotypic taxon|Monotypic]]<ref name=CatSG2017/> ||{{IUCN status|VU|22732}}<ref name=iucn1>{{cite iucn |title=''Panthera uncia'' |author1=McCarthy, T. |author2=Mallon, D. |author3=Jackson, R. |author4=Zahler, P. |author5=McCarthy, K. |name-list-style=amp |year= 2017 |page= e.T22732A50664030 |doi=10.2305/IUCN.UK.2017-2.RLTS.T22732A50664030.en |access-date=6 February 2023}}</ref> [[File:SnowLeopard distribution.jpg|frameless]] |- |} ===Extinct species and subspecies=== {| class="wikitable" |- ! Species and subspecies !! [[Fossil]] records!! Notes !Images |- style="vertical-align: top;" |''[[American lion|Panthera atrox]]'' |North America, 0.13 to 0.013 MYA, with dubious remains in South America.<ref name = "Chimento2017">{{Cite journal|last1= Chimento |first1= N. R.|last2= Agnolin|first2=F. L.|year= 2017 |title=The fossil American lion (''Panthera atrox'') in South America: Palaeobiogeographical implications |journal= Comptes Rendus Palevol |volume= 16|issue= 8|pages= 850–864 |doi= 10.1016/j.crpv.2017.06.009|bibcode= 2017CRPal..16..850C|doi-access= |hdl= 11336/65990|hdl-access= free}}</ref> |Commonly known as the American lion'', P. atrox'' is thought to have descended from a basal ''P. spelaea'' cave lion population isolated south of the [[Cordilleran Ice Sheet]], and then established a [[Mitochondrial DNA#Female inheritance|mitochondrial]] [[Sister group|sister clade]] circa [[Illinoian (stage)|200,000 BP.]]<ref name=Barnett2009>{{Cite journal |last1=Barnett |first1=R. |last2=Shapiro |first2=B. |last3=Barnes |first3=I. |last4=Ho |first4=S. Y. W. |last5=Burger |first5=J. |last6=Yamaguchi |first6=N. |last7=Higham |first7=T. F. G. |last8=Wheeler |first8=H. T. |last9=Rosendahl |first9=W. |date=2009 |title=Phylogeography of lions (''Panthera leo'' ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity |journal=Molecular Ecology |volume=18 |issue=8 |pages=1668–77 |doi=10.1111/j.1365-294X.2009.04134.x |pmid=19302360 |bibcode=2009MolEc..18.1668B |s2cid=46716748 |url=https://www.researchgate.net/publication/24216045 |access-date=7 March 2018 |archive-date=20 September 2018 |archive-url=https://web.archive.org/web/20180920011400/https://www.researchgate.net/publication/24216045 |url-status=live }}</ref> It was sometimes considered a subspecies either under the nomenclature of ''P. leo''<ref name=Barnett2009 /> or ''P. spelaea''.<ref>{{Cite journal |author1=Sotnikova, M. |author2=Nikolskiy, P. |date=2006 |title=Systematic position of the cave lion ''Panthera spelaea'' (Goldfuss) based on cranial and dental characters |journal=Quaternary International |language=en |volume=142–143 |pages=218–228 |doi=10.1016/j.quaint.2005.03.019 |bibcode=2006QuInt.142..218S |url=http://doc.rero.ch/record/13484/files/PAL_E260.pdf |access-date=24 January 2023 |archive-date=5 February 2023 |archive-url=https://web.archive.org/web/20230205061457/https://doc.rero.ch/record/13484/files/PAL_E260.pdf |url-status=live }}</ref> One of the largest ''Panthera'' species.<ref name="Christiansen2009">{{cite journal |last1=Christiansen |first1=P. |last2=Harris |first2=J. M. |year=2009 |title=Craniomandibular morphology and phylogenetic affinities of ''Panthera atrox'': implications for the evolution and paleobiology of the lion lineage |journal=Journal of Vertebrate Paleontology |volume=29 |issue=3 |pages=934–945 |bibcode=2009JVPal..29..934C |doi=10.1671/039.029.0314 |s2cid=85975640}}</ref> Became extinct around 13,000-12,000 years ago.<ref>{{Cite journal |last1=O’Keefe |first1=F. Robin |last2=Dunn |first2=Regan E. |last3=Weitzel |first3=Elic M. |last4=Waters |first4=Michael R. |last5=Martinez |first5=Lisa N. |last6=Binder |first6=Wendy J. |last7=Southon |first7=John R. |last8=Cohen |first8=Joshua E. |last9=Meachen |first9=Julie A. |last10=DeSantis |first10=Larisa R. G. |last11=Kirby |first11=Matthew E. |last12=Ghezzo |first12=Elena |last13=Coltrain |first13=Joan B. |last14=Fuller |first14=Benjamin T. |last15=Farrell |first15=Aisling B. |date=18 August 2023 |title=Pre–Younger Dryas megafaunal extirpation at Rancho La Brea linked to fire-driven state shift |url=https://www.science.org/doi/10.1126/science.abo3594 |journal=Science |language=en |volume=381 |issue=6659 |pages=eabo3594 |doi=10.1126/science.abo3594 |pmid=37590347 |s2cid=260956289 |issn=0036-8075 |access-date=11 March 2024 |archive-date=27 January 2024 |archive-url=https://web.archive.org/web/20240127153230/https://www.science.org/doi/10.1126/science.abo3594 |url-status=live |url-access=subscription }}</ref> |[[File:American lion National Park Service illustration.png|center|frameless]] |- style="vertical-align: top;" |''[[Panthera balamoides]]''<ref>{{Cite journal|last1=Stinnesbeck, S. R. |last2=Stinnesbeck, W. |last3=Frey, E. |last4=Olguín, J. A. |last5=Sandoval, C. R. |last6=Morlet, A. V. |author7=González, A. H. |year=2018 |title=''Panthera balamoides'' and other Pleistocene felids from the submerged caves of Tulum, Quintana Roo, Mexico |journal=Historical Biology: An International Journal of Paleobiology |volume=32 |issue= 7|pages=1–10 |doi=10.1080/08912963.2018.1556649|s2cid=92328512 }}</ref> |Mexico, ~0.13 MYA |Dubious, other authors suggest that the remains are actually of the extinct bear ''[[Arctotherium]]'' instead.<ref name=":3">{{Cite journal |author1=Blaine W. Schubert |author2=James C. Chatters |author3=Joaquin Arroyo-Cabrales |author4=Joshua X. Samuels |author5=Leopoldo H. Soibelzon |author6=Francisco J. Prevosti |author7=Christopher Widga |author8=Alberto Nava |author9=Dominique Rissolo |author10=Pilar Luna Erreguerena |year=2019 |title=Yucatán carnivorans shed light on the Great American Biotic Interchange |journal=Biology Letters |volume=15 |issue=5 |pages=Article ID 20190148 |doi=10.1098/rsbl.2019.0148 |pmc=6548739 |pmid=31039726}}</ref> | |- |''[[Panthera dhokpathanensis]]'' |[[Siwaliks]], ~2.7 MYA |Described in 1986.<ref>{{Cite journal|last=Bakr |first=A. |year=1986 |title=On a collection of Siwalik Carnivora |journal=Biological Society of Pakistan|volume=11|pages=1–64 }}</ref> | |- style="vertical-align: top;" |''[[Panthera fossilis]]''<ref>{{cite thesis |last1=Harington |first1=C. R. |year=1996 |title=Pleistocene mammals of the Yukon Territory |type=PhD |publisher=University of Alberta |location=Edmonton}}</ref> |Europe and Asia, 0.68 to 0.25 MYA |Extinct species of lion known from the Middle Pleistocene of Europe and Asia. One of the largest known species of ''Panthera''. Considered to be the ancestor of ''P.'' ''spelaea.''<ref name=":0">{{Cite journal |last1=Sabo |first1=Martin |last2=Tomašových |first2=Adam |last3=Gullár |first3=Juraj |date=August 2022 |title=Geographic and temporal variability in Pleistocene lion-like felids: Implications for their evolution and taxonomy |url=https://palaeo-electronica.org/content/2022/3681-lion-cranial-variability |journal=Palaeontologia Electronica |language=English |volume=25 |issue=2 |pages=1–27 |doi=10.26879/1175 |issn=1094-8074 |s2cid=251855356 |doi-access=free |access-date=11 March 2024 |archive-date=2 April 2023 |archive-url=https://web.archive.org/web/20230402015510/https://palaeo-electronica.org/content/2022/3681-lion-cranial-variability |url-status=live }}</ref> |[[File:W. Gornig - P. spelaea fossilis.png|center|frameless]] |- style="vertical-align: top;" |''[[Panthera gombaszoegensis]]'' |Europe, possibly Asia and Africa, 2.0 to 0.35 MYA |Ranged across Europe, as well as possibly Asia and Africa from around 2 million to 350,000 years ago.<ref name="Marciszak2014">{{cite journal |last1=Marciszak |first1=A. |year=2014 |title=Presence of ''Panthera gombaszoegensis'' (Kretzoi, 1938) in the late Middle Pleistocene of Biśnik Cave, Poland, with an overview of Eurasian jaguar size variability |journal=Quaternary International |volume=326-327 |pages=105–113 |bibcode=2014QuInt.326..105M |doi=10.1016/j.quaint.2013.12.029}}</ref> Often suggested to be the ancestor of the living jaguar (''Panthera onca''), and sometimes referred to as the "European jaguar". ''Panthera schreuderi'' and ''Panthera toscana'' are considered junior synonyms of ''P. gombaszoegensis''. It is occasionally classified as a subspecies of ''P. onca''.<ref name="hemmeretal2010">{{cite journal |last1=Hemmer, H. |last2=Kahlke, R. D. |last3=Vekua, A. K. |year=2010 |title=''Panthera onca georgica'' ssp. nov. from the Early Pleistocene of Dmanisi (Republic of Georgia) and the phylogeography of jaguars (Mammalia, Carnivora, Felidae) |journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen |volume=257|issue=1|pages=115–127 |doi=10.1127/0077-7749/2010/0067 }}</ref><ref>{{cite journal |author1=Mol, D. |author2=van Logchem, W. |author3=de Vos, J. |year=2011 |title=New record of the European jaguar, ''Panthera onca gombaszoegensis'' (Kretzoi, 1938), from the Plio-Pleistocene of Langenboom (The Netherlands) |journal=Cainozoic Research |volume=8 |issue=1–2 |pages=35–40 |url=http://natuurtijdschriften.nl/download?type=document;docid=541752 |access-date=28 September 2015 |archive-date=4 April 2019 |archive-url=https://web.archive.org/web/20190404200141/http://natuurtijdschriften.nl/download?type=document;docid=541752 |url-status=live }}</ref> |[[File:De vroeg-pleistocene sabeltandkat, Homotherium crenatidens (2008) Panthera onca gombaszoegensis.png|center|frameless]] | |- |''[[Panthera palaeosinensis]]'' |Northern China, ~3 MYA |Initially thought to be an ancestral tiger species, but several scientists place it close to the base of the genus ''Panthera''<ref name="Mazák2011">{{cite journal |last1=Mazák |first1=J. H. |last2=Christiansen |first2=P. |last3=Kitchener |first3=A. C. |year=2011 |title=Oldest Known Pantherine Skull and Evolution of the Tiger |journal=PLOS ONE |volume=6 |issue=10 |pages=e25483 |bibcode=2011PLoSO...625483M |doi=10.1371/journal.pone.0025483 |issn=1932-6203 |pmc=3189913 |pmid=22016768 |doi-access=free}}</ref> At least three recent studies considered ''Panthera zdanskyi'' likely to be a synonym of ''P. palaeosinensis''.<ref>{{cite journal |last1=Hemmer |first1=Helmut |title= The identity of the "lion", ''Panthera principialis'' sp. nov., from the Pliocene Tanzanian site of Laetoli and its significance for molecular dating the pantherine phylogeny, with remarks on Panthera shawi (Broom, 1948), and a revision of Puma incurva (Ewer, 1956), the Early Pleistocene Swartkrans "leopard" (Carnivora, Felidae)|url= |journal= Palaeobiodiversity and Palaeoenvironments|year=2023 |volume= 103|issue= 2|pages= 465–487|doi=10.1007/s12549-022-00542-2 |bibcode=2023PdPe..103..465H |access-date=}}</ref><ref>{{Cite journal|doi=10.1080/08912963.2022.2034808 |title=Discovery of jaguar from northeastern China middle Pleistocene reveals an intercontinental dispersal event |date=2023 |last1=Jiangzuo |first1=Qigao |last2=Wang |first2=Yuan |last3=Ge |first3=Junyi |last4=Liu |first4=Sizhao |last5=Song |first5=Yayun |last6=Jin |first6=Changzhu |last7=Jiang |first7=Hao |last8=Liu |first8=Jinyi |journal=Historical Biology |volume=35 |issue=3 |pages=293–302 |bibcode=2023HBio...35..293J |s2cid=246693903 }}</ref><ref>{{Cite journal|last1=Jiangzuo |first1=Q. |last2=Madurell-Malapeira |first2=J. |last3=Li |first3=X. |last4=Estraviz-López |first4=D. |last5=Mateus |first5=O. |last6=Testu |first6=A. |last7=Li |first7=S. |last8=Wang |first8=S. |last9=Deng |first9=T. |title=Insights on the evolution and adaptation toward high-altitude and cold environments in the snow leopard lineage |year=2025 |journal=Science Advances |volume=11 |issue=3 |pages=eadp5243 |doi=10.1126/sciadv.adp5243 |pmid=39813339 |pmc=11734717 |doi-access=free }}</ref> | |- |''[[Panthera principialis]]'' |Tanzania, ~3.7 MYA |Described in 2023.<ref name=principialis>{{Cite journal|last=Hemmer |first=H. |year=2023 |title=The identity of the "lion", ''Panthera principialis'' sp. nov., from the Pliocene Tanzanian site of Laetoli and its significance for molecular dating the pantherine phylogeny, with remarks on ''Panthera shawi'' (Broom, 1948), and a revision of ''Puma incurva'' (Ewer, 1956), the Early Pleistocene Swartkrans "leopard" (Carnivora, Felidae) |journal=Palaeobiodiversity and Palaeoenvironments |volume=103 |issue=2 |pages=465–487 |doi=10.1007/s12549-022-00542-2 |bibcode=2023PdPe..103..465H }}</ref> | |- style="vertical-align: top;" |''[[Panthera shawi]]'' |Laetoli site in Tanzania, ~3 MYA |A leopard-like cat<ref>{{cite journal|last=Sabol, M.|year=2011 |title=Masters of the lost world: a hypothetical look at the temporal and spatial distribution of lion-like felids |journal=Quaternaire |volume=4 |pages=229–236 |url=https://www.researchgate.net/publication/287422796}}</ref> | |- style="vertical-align: top;" |''[[Panthera spelaea]]'' |Much of Eurasia, 0.6 to 0.013 MYA<ref>{{cite journal |last=Stuart, A. J., Lister, A .M. |year=2011|title=Extinction chronology of the cave lion ''Panthera spelaea''|journal=Quaternary Science Reviews |volume=30 |issue=17 |pages=2329–2340 |doi=10.1016/j.quascirev.2010.04.023|bibcode=2011QSRv...30.2329S}}</ref> |Commonly known as the cave lion or steppe lion. Originally ''spelaea'' was classified as a subspecies of the extant lion ''P. leo''.<ref name=Sala>{{cite journal |last=Sala, B. |year=1990 |title=''Panthera leo fossilis'' (v. Reichenau, 1906) (Felidae) de Iserna la Pineta (Pléistocene moyen inférieur d'Italie) |journal=Géobios |volume=23 |issue=2 |pages=189–194 |doi=10.1016/S0016-6995(06)80051-3}}</ref> Results of recent genetic studies indicate that it belongs to a distinct species, namely ''P. spelaea'' that is most closely related to the modern lion among living ''Panthera'' species.<ref>{{Cite journal |last1=Marciszak |first1=A. |last2=Stefaniak |first2=K. |date=2010 |title=Two forms of cave lion: Middle Pleistocene ''Panthera spelaea fossilis'' Reichenau, 1906 and Upper Pleistocene ''Panthera spelaea spelaea'' Goldfuss, 1810 from the Bísnik Cave, Poland |journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen |volume=258 |issue=3 |pages=339–351 |doi=10.1127/0077-7749/2010/0117 |url=https://www.researchgate.net/publication/233669138 |access-date=7 March 2018 |archive-date=25 September 2018 |archive-url=https://web.archive.org/web/20180925142142/https://www.researchgate.net/publication/233669138 |url-status=live}}</ref><ref>{{Cite journal |last1=Marciszak |first1=A. |last2=Schouwenburg |first2=C. |last3=Darga |first3=R. |date=2014 |title=Decreasing size process in the cave (Pleistocene) lion ''Panthera spelaea'' (Goldfuss, 1810) evolution – A review |journal=Quaternary International |series=Fossil remains in karst and their role in reconstructing Quaternary paleoclimate and paleoenvironments |volume=339–340 |pages=245–257 |doi=10.1016/j.quaint.2013.10.008 |bibcode=2014QuInt.339..245M}}</ref> Other genetic results indicate that ''P. fossilis'' also warrants status as a species.<ref name=SotnikovaForonvoa2014>{{cite journal |author1=Sotnikova, M. V. |author2=Foronova, I. V. |title=First Asian record of ''Panthera (Leo) fossilis'' (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia. |year=2014|journal=Integrative Zoology |volume=9 |issue=4 |pages=517–530 |url=https://www.researchgate.net/publication/259545849|doi=10.1111/1749-4877.12082 |pmid=24382145}}</ref><ref name=Barnett2016>{{cite journal |author1=Barnett, R. |author2=Mendoza, M. L. Z. |author3= Soares, A. E. R. |author4=Ho, S. Y. W. |author5=Zazula, G. |author6=Yamaguchi, N. |author7=Shapiro, B. |author8=Kirillova, I. V. |author9=Larson, G. |author10=Gilbert, M. T. P. |title=Mitogenomics of the Extinct Cave Lion, ''Panthera spelaea'' (Goldfuss, 1810), resolve its position within the Panthera cats |year=2016 |journal=Open Quaternary |volume=2 |page=4 |doi=10.5334/oq.24|doi-access=free |hdl=10576/22920 |hdl-access=free}}</ref> It became extinct around 14,500-14,000 years ago.<ref>{{Cite journal |last1=Stuart |first1=Anthony J. |last2=Lister |first2=Adrian M. |date=August 2011 |title=Extinction chronology of the cave lion ''Panthera spelaea'' |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379110001320 |journal=Quaternary Science Reviews |language=en |volume=30 |issue=17–18 |pages=2329–2340 |doi=10.1016/j.quascirev.2010.04.023 |bibcode=2011QSRv...30.2329S |access-date=11 March 2024 |archive-date=24 April 2024 |archive-url=https://web.archive.org/web/20240424144240/https://linkinghub.elsevier.com/retrieve/pii/S0277379110001320 |url-status=live|url-access=subscription }}</ref> |[[File:W. Gornig - P. spelaea spelaea.png|center|frameless]] |- |''[[Panthera youngi]]''<ref>{{cite journal|last=Pei, W. C.|year=1934|title=On the Carnivora from Locality 1 of Choukoutien |journal=Palaeontologica Sinica Series C, Fascicle 1 |pages=1−166}}</ref> |China, Japan, ~0.35 MYA | |[[File:Panthera youngi.webp|center|frameless]] |- style="vertical-align: top;" |''[[Panthera zdanskyi]]'' |[[Gansu province]] of northwestern China, 2.55 to 2.16 MYA |It was initially considered to be a close relative of the tiger.<ref name=Mazák2011/> But it is possibly synonymous with ''P. palaeosinensis''.<ref name=principialis/><ref>{{Cite journal |doi=10.1080/08912963.2022.2034808 |title=Discovery of jaguar from northeastern China middle Pleistocene reveals an intercontinental dispersal event |date=2023 |last1=Jiangzuo |first1=Q. |last2=Wang |first2=Y. |last3=Ge |first3=J. |last4=Liu |first4=S. |last5=Song |first5=Y. |last6=Jin |first6=C. |last7=Jiang |first7=H. |last8=Liu |first8=J. |journal=Historical Biology |volume=35 |issue=3 |pages=293–302 |bibcode=2023HBio...35..293J |s2cid=246693903}}</ref> |[[File:Longdan tiger.png|center|frameless]] |- style="vertical-align: top;" |''[[Panthera leo sinhaleyus]]'' |Sri Lanka |This lion subspecies was described on the basis of two teeth.<ref>{{cite journal |author=Manamendra-Arachchi, K., Pethiyagoda, R., Dissanayake, R., Meegaskumbura, M. |year=2005 |title=A second extinct big cat from the late Quaternary of Sri Lanka |journal=The Raffles Bulletin of Zoology |issue=Supplement 12 |pages=423–434 |url=http://rmbr.nus.edu.sg/rbz/biblio/s12/s12rbz423-434.pdf |access-date=28 October 2013 |archive-date=7 August 2007 |archive-url=https://web.archive.org/web/20070807215533/http://rmbr.nus.edu.sg/rbz/biblio/s12/s12rbz423-434.pdf}}</ref> | |- style="vertical-align: top;" |''[[Panthera onca augusta]]''<ref name=Columbia>{{cite journal |author1=Ruiz-Garcia, M. |author2=Payan, E. |author3=Murillo, A. |author4=Alvarez, D. |name-list-style=amp |year=2006 |title=DNA microsatellite characterization of the jaguar (''Panthera onca'') in Colombia |journal=Genes & Genetic Systems |volume=81 |issue=2 |pages=115–127 |doi=10.1266/ggs.81.115 |pmid=16755135 |url=https://www.jstage.jst.go.jp/article/ggs/81/2/81_2_115/_pdf |access-date=8 September 2015 |doi-access=free |archive-date=6 June 2023 |archive-url=https://web.archive.org/web/20230606112430/https://www.jstage.jst.go.jp/article/ggs/81/2/81_2_115/_pdf |url-status=live}}</ref> |North America |May have lived in temperate forests across North America<ref name=":1">{{Cite journal |last1=Moreno, A. |last2=Lima-Ribeiro, M. |date=2015 |title=Ecological niche models, fossil record and the multi-temporal calibration for ''Panthera onca'' (Linnaeus, 1758) (Mammalia: Felidae) |url=http://revista.rebibio.net/v2n4/v02n04a14.pdf |journal=Brazilian Journal of Biological Sciences |volume=2 |issue=4 |pages=309–319 |access-date=4 April 2019 |archive-date=4 April 2019 |archive-url=https://web.archive.org/web/20190404195921/http://revista.rebibio.net/v2n4/v02n04a14.pdf |url-status=live}}</ref> | |- |''[[Panthera onca mesembrina]]''<ref>{{cite journal |last1=Roth |first1=S. |year=1899 |title=Descripción de los restos encontrados en la caverna de Última Esperanza |journal=Revista del Museo la Plata |volume=9 |pages=381–388}}</ref> |South America |May have lived in grasslands in South America, unlike the modern jaguar |[[File:Panthera onca mesembrina JF.png|center|frameless]] | |- style="vertical-align: top;" |''[[Panthera pardus spelaea]]'' |Europe |Closely related to Asiatic leopard subspecies,<ref>{{Cite journal |last1=Paijmans, J. L. A. |last2=Barlow, A. |last3=Förster, D. W. |last4=Henneberger, K. |last5=Meyer, M. |last6=Nickel, B. |last7=Nagel, D. |last8=Havmøller, R. W. |last9=Baryshnikov, G. F. |last10=Joger, U. |last11=Rosendahl, W. |last12=Hofreiter, M. |year=2018 |title=Historical biogeography of the leopard (''Panthera pardus'') and its extinct Eurasian populations |journal=BMC Evolutionary Biology |volume=18 |issue=1 |page=156 |doi=10.1186/s12862-018-1268-0 |pmid=30348080 |pmc=6198532 |doi-access=free |bibcode=2018BMCEE..18..156P}}</ref> |[[File:Chauvet leopard rock art (white background).png|center|frameless]] | |- style="vertical-align: top;" |''[[Panthera tigris acutidens]]'' |Much of Asia |Not closely related to modern tiger subspecies<ref name="PBDB">{{Cite journal |last1=Hasegawa |first1=Y. |last2=Tomida |first2=Y. |last3=Kohno |first3=N. |last4=Ono |first4=K. |last5=Nokariya |first5=H. |last6=Uyeno |first6=T. |year=1988 |title=Quaternary vertebrates from Shiriya area, Shimokita Pininsula, northeastern Japan |journal=Memoirs of the National Science Museum |volume=21 |pages=17–36}}</ref> | |- |''[[Panthera tigris soloensis]]'' |Java, Indonesia |Not closely related to modern tiger subspecies<ref name="PBDB"/> | |- |''[[Panthera tigris trinilensis]]'' |Java, Indonesia |Not closely related to modern tiger subspecies<ref name="PBDB"/> | |- |} Other, now invalid, species have also been described, such as ''Panthera crassidens'' from South Africa, which was later found to be based on a mixture of leopard and cheetah fossils.<ref>{{cite journal|author=Turner, A.|year=1984|title=''Panthera crassidens'' Broom, 1948. The cat that never was?|journal=South African Journal of Science|volume=80|issue=5|pages=227–233 |url=https://journals.co.za/docserver/fulltext/sajsci/80/5/7798.pdf?expires=1544536360&id=id&accname=guest&checksum=1BB15474E5A55CD94EC66579004AE19D}}</ref> ===Phylogeny=== [[File:Two cladograms for Panthera.svg|thumbnail|Two cladograms proposed for ''Panthera''. The upper one is based on phylogenetic studies by Johnson et al. (2006),<ref name=Johnson2006/> and by Werdelin et al. (2010).<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 |title=Biology and Conservation of Wild Felids |publisher=[[Oxford University Press]] |location=Oxford |editor-first1=D. W. |editor-last1=Macdonald |editor-first2=A. J. |editor-last2=Loveridge |pages=59–82 |chapter-url=https://www.researchgate.net/publication/266755142 |isbn=978-0-19-923445-5 |access-date=3 February 2019 |archive-date=25 September 2018 |archive-url=https://web.archive.org/web/20180925141956/https://www.researchgate.net/publication/266755142 |url-status=live }}</ref> The lower cladogram is based on a study by Davis et al. (2010)<ref name=Davis2010/> and by Mazák et al. (2011).<ref name="Mazák2011"/>]]In 2018, results of a [[phylogenetic]] study on living and [[fossil]] cats were published. This study was based on the [[Morphology (biology)|morphological]] diversity of the [[mandible]]s of [[saber-toothed cat]]s, their [[speciation]] and [[extinction rate]]s.<ref>{{cite journal |author1=Piras, P. |author2=Silvestro, D. |author3=Carotenuto, F. |author4=Castiglione, S.|author5=Kotsakis, A. |author6=Maiorino, L. |author7=Melchionna, M. |author8=Mondanaro, A. |author9=Sansalone, G. |author10=Serio, C. |author11=Vero, V.A. |author12=Raia, P. |year=2018 |title=Evolution of the sabertooth mandible: A deadly ecomorphological specialization |journal=[[Palaeogeography, Palaeoclimatology, Palaeoecology]] |volume=496 |pages=166−174 |doi=10.1016/j.palaeo.2018.01.034|bibcode=2018PPP...496..166P |hdl=2158/1268434 |hdl-access=free}}</ref> {{clade| style=font-size:85%;line-height:85% |label1='''''Panthera''''' |1={{Clade |1=†''[[Panthera palaeosinensis]]'' |2={{Clade |1={{Clade |label1= |1={{Clade |1=†''[[Panthera blytheae]]'' |2=Snow leopard [[File:Stamp-russia2014-save-russian-cats-(snow leopard).png|70 px]]}} }} |2={{Clade |label1= |1={{Clade |1=†''[[Panthera zdanskyi]]'' |2=Tiger [[File:Stamp-russia2014-save-russian-cats-(tiger).png|70 px]]}} }} }} |3={{Clade |1={{Clade |label1= |1={{Clade |1=†''[[Panthera gombaszoegensis]]'' (sometimes called the European jaguar) <span style="{{MirrorH}}">[[File:De vroeg-pleistocene sabeltandkat, Homotherium crenatidens (2008) Panthera onca gombaszoegensis.png|70 px]]</span> |2=Jaguar [[File:Felis onca - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70 px]]}} }} |2={{Clade |label1= |1={{Clade |1=Leopard [[File:Felis pardus - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70 px]] |2={{Clade |1=Lion <span style="{{MirrorH}}">[[File:W. Gornig - P. leo leo ♂.png|70 px]]</span> |2={{Clade |1=†''[[Panthera spelaea]]'' (cave lion or steppe lion) <span style="{{MirrorH}}">[[File:W. Gornig - P. spelaea spelaea.png|70 px]]</span> |2=†''[[American lion|Panthera atrox]]'' (American lion) <span style="{{MirrorH}}">[[File:W. Gornig - P. spelaea spelaea 02.png|70 px]]</span>}} }} }} }} }} }} }} ==See also== {{Portal|Cats|Mammals}} * [[Panthera hybrid|''Panthera'' hybrid]] * [[Panther (legendary creature)]] ==Notes== {{notefoot}} ==References== {{Reflist}} ==Further reading== * {{cite book |author1=Turner, A. |author2=Antón, M. |author2-link=Mauricio Antón |date=1997 |title=The Big Cats and Their Fossil Relatives: An Illustrated Guide to Their Evolution and Natural History |publisher=[[Columbia University Press]] |isbn=978-0-231-10228-5 |url=https://books.google.com/books?id=66mRJSxIAfoC}} ==External links== * {{Commons category-inline|Panthera|''Panthera''}} * {{cite AV media |title=The Ghostly Origins of the Big Cats |series=[[PBS Eons]] |date=16 May 2019 |url=https://www.youtube.com/watch?v=oPJnqWke5n8 |type=video |archive-url=https://ghostarchive.org/varchive/youtube/20211211/oPJnqWke5n8| archive-date=11 December 2021 |url-status=live|via=[[YouTube]] |language=en |ref=Eons2021}}{{cbignore}} {{Carnivora|Fe.}} {{Feliformia|Fel.|state=collapsed}} {{Taxonbar|from=Q127960}} {{Authority control}} [[Category:Panthera| ]] [[Category:Carnivoran genera]] [[Category:Extant Miocene first appearances]] [[Category:Taxa named by Lorenz Oken]] [[Category:Taxa described in 1816]]
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