Editing Panthera (section)

===Extinct species and subspecies===
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! Species and subspecies !! [[Fossil]] records!! Notes
!Images
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|''[[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 }}</ref>
|[[File:American lion National Park Service illustration.png|center|frameless]]
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|''[[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>
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|''[[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>
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|''[[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]]
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|''[[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]]
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|''[[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>
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|''[[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>
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|''[[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>
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|''[[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}}</ref>
|[[File:W. Gornig - P. spelaea spelaea.png|center|frameless]]
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|''[[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
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|[[File:Panthera youngi.webp|center|frameless]]
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|''[[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]]
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|''[[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>
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|''[[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>
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|''[[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]]
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|''[[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]]
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|''[[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>
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|''[[Panthera tigris soloensis]]''
|Java, Indonesia
|Not closely related to modern tiger subspecies<ref name="PBDB"/>
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|''[[Panthera tigris trinilensis]]''
|Java, Indonesia
|Not closely related to modern tiger subspecies<ref name="PBDB"/>
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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>