Panthera Explained
Panthera is a genus within the 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, as well as a number of extinct species.
Etymology
The word panther derives from classical Latin panthēra, itself from the ancient Greek pánthēr (πάνθηρ).[1]
Characteristics
In Panthera species, the dorsal profile of the skull is flattish or evenly convex. The frontal interorbital area is not noticeably elevated, and the area behind the elevation is less steeply sloped. The basic cranial axis is nearly horizontal. The inner chamber of the bullae is large, the outer small. The partition between them is close to the external auditory meatus. The convexly rounded chin is sloping.[2] 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 that provide a lower resistance to airflow; it was therefore proposed to be retained in the genus Uncia.[3] 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.[4]
Evolution
The geographic origin of the Panthera is most likely northern Central Asia. Panthera blytheae, possibly the oldest known Panthera species, is similar in skull features to the snow leopard, though its taxonomic placement has been disputed by other researchers who suggest that the species likely belongs to a different genus.[5] [6] The tiger, snow leopard, and clouded leopard genetic lineages dispersed in Southeast Asia during the Miocene.[7] Genetic studies indicate that the pantherine cats diverged from the subfamily Felinae between six and ten million years ago.[8] The genus Neofelis is sister to Panthera.[8] [9] [10] [11] The clouded leopard appears to have diverged about . Panthera diverged from other cat species about and then evolved into the species tiger about, snow leopard about and leopard about . 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, and that both P. atrox and P. spelaea are most closely related to lions among living Panthera species.[12] The snow leopard is nested within Panthera and is the sister species of the tiger.[13]
Results of a 2016 study based on analysis of biparental nuclear genomes suggest the following relationships of living Panthera species:[14]
The extinct species Panthera gombaszoegensis, was probably closely related to the modern jaguar. The first fossil remains were excavated in Olivola, in Italy, and date to .[15] Fossil remains found in South Africa that appear to belong within the Panthera lineage date to about .[16]
Classification
Panthera was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group.[17] [18] During the 19th and 20th centuries, various explorers and staff of natural history museums 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.[19] [20] [21] 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 skulls.[22] 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.[23] [24] Subsequently, it was proposed to group all African leopard populations to P. p. pardus and retain eight subspecific names for Asian leopard populations.[25] 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.
Based on genetic research, it was suggested to group all living sub-Saharan lion populations into P. l. leo.[26] Results of phylogeographic studies indicate that the Western and Central African lion populations are more closely related to those in India and form a different clade than lion populations in Southern and East Africa; southeastern Ethiopia is an admixture region between North African and East African lion populations.[27] [28]
Black panthers do not form a distinct species, but are melanistic specimens of the genus, most often encountered in the leopard and jaguar.[29] [30]
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 revised in 2017 by the Cat Classification Task Force of the Cat Specialist Group:[31]
Extinct species and subspecies
Species and subspecies | Fossil records | Notes | Images |
---|
Panthera blytheae | Tibetan Plateau | Intitally described as one of the oldest known Panthera species, possibly closely related to the snow leopard, though other researchers consider it as a species of a different genus. | |
Panthera palaeosinensis | Northern China | Initially thought to be an ancestral tiger species, but several scientists place it close to the base of the genus Panthera | |
Panthera zdanskyi | Gansu province of northwestern China | Possibly a close relative of the tiger | |
Panthera youngi[46] | China, Japan | | |
Panthera atrox | North America, dubious remains in South America.[47] | 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 sister clade circa 200,000 BP.[48] It was sometimes considered a subspecies either under the nomenclature of P. leo or P. spelaea.[49] One of the largest Panthera species.[50] Became extinct around 13,000-12,000 years ago.[51] | |
Panthera balamoides[52] | Mexico | Dubious, other authors suggest that the remains are actually of the extinct bear Arctotherium instead.[53] | |
Panthera gombaszoegensis | Europe | Ranged across Europe from around 2 million to 350,000 years ago.[54] Panthera schreuderi and Panthera toscana are considered junior synonyms of P. gombaszoegensis. It is occasionally classified as a subspecies of P. onca.[55] [56] | |
Panthera fossilis[57] | Europe | Likely the ancestor of P. spelaea[58] | |
Panthera spelaea | Much of Eurasia[59] | Commonly known as the cave lion or steppe lion. Originally spelaea was classified as a subspecies of the extant lion P. leo.[60] 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.[61] [62] Other genetic results indicate that P. fossilis also warrants status as a species.[63] [64] Became extinct around 14,500-14,000 years ago.[65] | |
Panthera leo sinhaleyus | Sri Lanka | This lion subspecies was described on the basis of two teeth.[66] | |
Panthera onca augusta[67] | North America | May have lived in temperate forests across North America[68] | |
Panthera onca mesembrina[69] | South America | May have lived in grasslands in South America, unlike the modern jaguar | |
Panthera pardus spelaea | Europe | Closely related to Asiatic leopard subspecies,[70] | |
Panthera shawi | Laetoli site in Tanzania | A leopard-like cat[71] | |
Panthera tigris acutidens | Much of Asia | Not closely related to modern tiger subspecies[72] | |
Panthera tigris soloensis | Java, Indonesia | Not closely related to modern tiger subspecies | |
Panthera tigris trinilensis | Java, Indonesia | Not closely related to modern tiger subspecies | |
Panthera principialis | Tanzania | Described in 2022.[73] | |
Panthera dhokpathanensis | Siwaliks | Described in 1986.[74] | |
|
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.
[75] Phylogeny
The cladogram below follows Mazák, Christiansen and Kitchener (2011).[76]
In 2018, results of a phylogenetic study on living and fossil cats were published. This study was based on the morphological diversity of the mandibles of saber-toothed cats, their speciation and extinction rates. The generated cladogram indicates a different relation of the Panthera species, as shown below:[77]
See also
Further reading
External links
Notes and References
- Book: Liddell, H. G. . Scott, R. . amp . 1940 . πάνθηρ . https://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3D%2377441 . A Greek-English Lexicon . Revised and augmented . Oxford . Clarendon Press . 21 February 2021 . 11 April 2008 . https://web.archive.org/web/20080411203109/http://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3D%2377441 . live .
- Book: Pocock, R. I. . 1939 . Panthera . https://archive.org/stream/PocockMammalia1/pocock1#page/n243/mode/2up . The fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1 . Taylor and Francis . London . 196–239.
- The larynx of roaring and non-roaring cats . Journal of Anatomy . 1989 . 1256521 . 2606766 . 117–121 . 163 . M. H.. Hast.
- Weissengruber . G. E. . Forstenpointner, G. . Peters, G. . Kübber-Heiss, A. . Fitch, W. T. . 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 of Anatomy . 195–209 . 201. 3 . 2002 . 10.1046/j.1469-7580.2002.00088.x . 1570911 . 12363272.
- Geraads, D. . Peigné, S . Re-appraisal of Felis pamiri Ozansoy 1959 (Carnivora, Felidae) from the upper Miocene of Turkey: the earliest pantherine cat? . Journal of Mammalian Evolution . 24 . 4 . 415–425 . 2017 . 10.1007/s10914-016-9349-6. 207195894.
- Hemmer. H. . 2023 . The evolution of the palaeopantherine cats, Palaeopanthera gen. nov. blytheae (Tseng et al., 2014) and Palaeopanthera pamiri (Ozansoy, 1959) comb. nov. (Mammalia, Carnivora, Felidae) . Palaeobiodiversity and Palaeoenvironments . 103 . 4 . 827–839 . 10.1007/s12549-023-00571-5 . 2023PdPe..103..827H . 257842190.
- Tseng . Z.J. . Wang . X. . Slater . G.J. . Takeuchi . G.T. . Li . Q. . Liu . J. . Xie . G. . amp . 2014 . Himalayan fossils of the oldest known pantherine establish ancient origin of big cats . Proceedings of the Royal Society B: Biological Sciences . 281 . 1774 . 20132686 . 10.1098/rspb.2013.2686 . 24225466 . 3843846.
- Johnson . W.E. . Eizirik . E. . Pecon-Slattery . J. . Murphy . W.J. . Antunes . A. . Teeling . E. . O'Brien . S.J. . amp . 2006 . 10.1126/science.1122277 . The Late Miocene radiation of modern Felidae: A genetic assessment . Science . 311 . 73–77 . 16400146 . 5757 . 2006Sci...311...73J . 41672825 . 28 June 2019 . 4 October 2020 . https://web.archive.org/web/20201004075725/https://zenodo.org/record/1230866 . live.
- Janczewski . D.N. . Modi . W.S. . Stephens . J.C. . O'Brien . S.J. . amp . 1996 . Molecular evolution of mitochondrial 12S RNA and cytochrome b sequences in the pantherine lineage of Felidae . Molecular Biology and Evolution . 12 . 4 . 690–707 . 10.1093/oxfordjournals.molbev.a040232 . 7544865 . free.
- Johnson . W. E. . O'Brien . S.J. . amp . 1997 . Phylogenetic reconstruction of the Felidae using 16S rRNA and NADH-5 mitochondrial genes . Journal of Molecular Evolution . 44 . S1 . S98–S116 . 10.1007/PL00000060 . 9071018 . 1997JMolE..44S..98J . 40185850 . 28 June 2019 . 4 October 2020 . https://web.archive.org/web/20201004075723/https://zenodo.org/record/1232587 . live.
- Yu . L. . Zhang . Y.P. . amp . 2005 . Phylogenetic studies of pantherine cats (Felidae) based on multiple genes, with novel application of nuclear beta-fibrinogen intron 7 to carnivores . Molecular Phylogenetics and Evolution . 35 . 2 . 483–495 . 10.1016/j.ympev.2005.01.017 . 15804417. 2005MolPE..35..483Y .
- Barnett . R. . Shapiro . B. . Beth Shapiro . Barnes . I. . Ho . S.Y.W. . Burger . J. . Joachim Burger . Yamaguchi . N. . Higham . T.F.G. . Wheeler . H.T. . Rosendahl . W. . Sher . A.V. . Sotnikova . M. . Kuznetsova . T. . Baryshnikov . G.F. . Martin . L.D. . Harington . C.R. . Burns . J.A. . Cooper . A. . amp . Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity . . 2009 . 18 . 8 . 1668–1677 . 19302360 . 10.1111/j.1365-294X.2009.04134.x . 2009MolEc..18.1668B . 46716748 . 24 February 2019 . 8 August 2017 . https://web.archive.org/web/20170808225555/https://www.zin.ru/Labs/theriology/eng/staff/baryshnikov/references/barnett_et_al_2009.pdf . live.
- Davis . B.W. . Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, Panthera (Carnivora: Felidae) . Li, G. . Murphy, W.J. . Molecular Phylogenetics and Evolution . 56 . 2010 . 1 . 64–76 . 20138224 . 10.1016/j.ympev.2010.01.036. 2010MolPE..56...64D .
- Li, G. . Davis, B. W. . Eizirik, E. . amp . Murphy, W. J. . 2016 . Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae) . Genome Research . 26 . 1 . 1–11 . 10.1101/gr.186668.114 . 26518481 . 4691742.
- Hemmer . H. . Kahlke . R.D. . Vekua . A.K. . amp . 2001 . 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 . Geobios . 34 . 4 . 475–486 . 10.1016/s0016-6995(01)80011-5 . 2001Geobi..34..475H.
- Turner . A. . 1987 . New fossil carnivore remains from the Sterkfontein hominid site (Mammalia: Carnivora) . Annals of the Transvaal Museum . 34 . 15 . 319–347 .
- Book: Oken . L. . Lehrbuch der Zoologie. 2. Abtheilung . August Schmid & Comp. . 1816 . Jena . 1052 . 1. Art, Panthera . https://books.google.com/books?id=S5o5AAAAcAAJ&pg=PA1052.
- Allen . J. A. . 1902 . Mammal names proposed by Oken in his 'Lehrbuch der Zoologie' . Bulletin of the American Museum of Natural History . 16 . 27 . 373−379 . 4 April 2019 . 9 May 2023 . 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 . live .
- Pocock . R. I. . 1930 . The panthers and ounces of Asia . Journal of the Bombay Natural History Society . 34 . 1. 65–82.
- Pocock . R. I. . 1932 . The leopards of Africa . 10.1111/j.1096-3642.1932.tb01085.x . Proceedings of the Zoological Society of London . 102 . 2. 543–591.
- Pocock . R. I. . 1939 . The races of jaguar (Panthera onca) . Novitates Zoologicae . 41 . 406–422.
- Pocock . R. I. . 1916 . The Classification and Generic Nomenclature of F. uncia and its Allies . The Annals and Magazine of Natural History: Including Zoology, Botany, and Geology . Series 8 . XVIII . 105 . 314–316 . 10.1080/00222931608693854.
- O'Brien, S. J. . Martenson, J. S. . Packer, C. . Herbst, L. . de Vos, V. . Joslin, P. . Ott-Joslin, J. . Wildt, D. E. . Bush, M. . amp . 1987 . Biochemical genetic variation in geographic isolates of African and Asiatic lions . National Geographic Research . 3 . 1 . 114–124 . 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 . 2 May 2013.
- Miththapala . S. . Seidensticker . J. . O'Brien . S. J. . 1996 . Phylogeographic subspecies recognition in leopards (Panthera pardus): Molecular genetic variation . Conservation Biology . 10 . 4. 1115–1132 . 10.1046/j.1523-1739.1996.10041115.x. 1996ConBi..10.1115M .
- Uphyrkina . O. . Johnson . W. E. . Quigley . H. B. . Miquelle . D. G. . Marker . L. . Bush . M. E. . O'Brien . S. J. . 2001 . Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus . Molecular Ecology . 10 . 11 . 2617–2633 . 10.1046/j.0962-1083.2001.01350.x . 11883877 . 2001MolEc..10.2617U . 304770 . 28 June 2019 . 6 August 2020 . https://web.archive.org/web/20200806044827/https://zenodo.org/record/1236516 . live .
- Dubach . J. . Patterson . B. D. . Briggs . M. B. . Venzke . K. . Flamand . J. . Stander . P. . Scheepers . L. . Kays . R. W. . 2005 . Molecular genetic variation across the southern and eastern geographic ranges of the African lion, Panthera leo . Conservation Genetics . 6 . 1 . 15–24 . 10.1007/s10592-004-7729-6 . 2005ConG....6...15D . 30414547 . 5 March 2019 . 5 March 2024 . 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 . live .
- Bertola . L. D. . Van Hooft . W. F. . Vrieling . K. . Uit De Weerd . D. R. . York . D. S. . Bauer . H. . Prins . H. H. T. . Funston . P. J. . Udo De Haes . H. A. . Leirs . H. . Van Haeringen . W. A. . Sogbohossou . E. . Tumenta . P. N. . De Iongh . H. H. . Genetic diversity, evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa . 10.1111/j.1365-2699.2011.02500.x . . 38 . 7 . 1356–1367 . 2011 . 2011JBiog..38.1356B . 82728679 . 17 January 2017 . 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 . 8 June 2019 .
- Bertola, L. D. . Jongbloed, H. . Van Der Gaag, K. J. . De Knijff, P. . Yamaguchi, N. . Hooghiemstra, H. . Bauer, H. . Henschel, P. . White, P. A. . Driscoll, C. A. . Tende, T. . 2016 . Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (Panthera leo) . Scientific Reports . 6 . 30807 . 10.1038/srep30807. 27488946 . 4973251 . 2016NatSR...630807B.
- Robinson, R. . 1970 . Inheritance of black form of the leopard Panthera pardus . Genetica . 41 . 1 . 190–197 . 5480762 . 10.1007/bf00958904. 5446868 .
- Eizirik, E. . Yuhki, N. . Johnson, W. E. . Menotti-Raymond, M. . Hannah, S. S. . O'Brien, S. J. . Molecular Genetics and Evolution of Melanism in the Cat Family . Current Biology . 2003 . 13 . 5 . 448–453 . 10.1016/S0960-9822(03)00128-3 . 12620197. 19021807 . free . 2003CBio...13..448E .
- Kitchener, A. C. . Breitenmoser-Würsten, C. . Eizirik, E. . Gentry, A. . Werdelin, L. . Wilting, A. . Yamaguchi, N. . Abramov, A. V. . Christiansen, P. . Driscoll, C. . Duckworth, J. W. . Johnson, W. . Luo, S.-J. . Meijaard, E. . O'Donoghue, P. . Sanderson, J. . Seymour, K. . Bruford, M. . Groves, C. . Hoffmann, M. . Nowell, K. . Timmons, Z. . Tobe, S. . 2017 . A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group . Cat News . Special Issue 11 . 66−75 . 13 May 2018 . 17 January 2020 . https://web.archive.org/web/20200117172708/https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y . live .
- Mazak, V. . 1975 . Notes on the Black-maned Lion of the Cape, Panthera leo melanochaita (Ch. H. Smith, 1842) and a Revised List of the Preserved Specimens . Verhandelingen Koninklijke Nederlandse Akademie van Wetenschappen . 64 . 1–44.
- Panthera leo . Bauer, H. . Packer, C. . Funston, P. F. . Henschel, P. . Nowell, K. . amp . e.T15951A115130419 . 2016.
- Larson. S. E.. 1997 . Taxonomic re-evaluation of the jaguar . Zoo Biology . en . 16 . 2 . 107–120 . 10.1002/(SICI)1098-2361(1997)16:2<107::AID-ZOO2>3.0.CO;2-E.
- Panthera onca . Quigley, H.. Foster, R.. Petracca, L.. Payan, E.. Salom, R. . Harmsen, B. . amp . e.T15953A123791436 . 2017.
- Book: Meyer, F. A. A. . 1794 . Zoologische Annalen. Erster Band . Weimar . Im Verlage des Industrie-Comptoirs . 394–396 . Über de la Metheries schwarzen Panther . https://books.google.com/books?id=v4s-AAAAcAAJ&pg=PA394.
- Cuvier, G. . 1809 . Recherches sur les espėces vivantes de grands chats, pour servir de preuves et d'éclaircissement au chapitre sur les carnassiers fossils . Annales du Muséum National d'Histoire Naturelle . Tome XIV . 136–164.
- Book: Hemprich . W. . Ehrenberg . C. G. . 1830 . Felis, pardus?, nimr . Plate 17 . 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 . Berolini . Officina Academica . Dr. C. G. Ehrenberg . https://archive.org/details/SymbolaephysicaMammEhreA/page/n60.
- Pocock, R. I. . 1930 . The Panthers and Ounces of Asia . Journal of the Bombay Natural History Society . 34 . 2 . 307–336.
- Deraniyagala . P. E. P. . 1956 . The Ceylon leopard, a distinct subspecies . Spolia Zeylanica . 28 . 115–116.
- Panthera pardus . Stein, A. B. . Athreya, V. . Gerngross, P. . Balme, G. . Henschel, P. . Karanth, U. . Miquelle, D. . Rostro, S. . Kamler, J. F. . Laguardia, A. . amp . e.T15954A102421779. 2016.
- Book: Linnaeus, C. . 1758 . Caroli Linnæi Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis . Tomus I . decima, reformata . Holmiae . Laurentius Salvius . 41−42 . Felis . https://archive.org/stream/mobot31753000798865#page/41/mode/2up.
- Panthera tigris . Goodrich, J. . Lynam, A. . Miquelle, D. . Wibisono, H. . Kawanishi, K. . Pattanavibool, A. . Htun, S. . Tempa, T. . Karki, J. . Jhala, Y. . Karanth, U. . amp . e.T15955A50659951 . 2015.
- Book: Schreber . J. C. D. . Die Unze . Die Säugethiere in Abbildungen nach der Natur mit Beschreibungen . 1777 . 386–387 . Erlangen . Wolfgang Walther . https://archive.org/stream/SaYugthiereAbbiIIISchr#page/386/mode/2up.
- Panthera uncia . McCarthy, T. . Mallon, D. . Jackson, R. . Zahler, P. . McCarthy, K. . amp . 2017 . e.T22732A50664030 . 10.2305/IUCN.UK.2017-2.RLTS.T22732A50664030.en . 6 February 2023.
- Pei, W. C.. 1934. On the Carnivora from Locality 1 of Choukoutien . Palaeontologica Sinica Series C, Fascicle 1. 1−166.
- Chimento . N. R.. Agnolin. F. L.. 2017 . The fossil American lion (Panthera atrox) in South America: Palaeobiogeographical implications . Comptes Rendus Palevol . 16. 8. 850–864 . 10.1016/j.crpv.2017.06.009. 2017CRPal..16..850C. 11336/65990. free.
- Barnett . R. . Shapiro . B. . Barnes . I. . Ho . S. Y. W. . Burger . J. . Yamaguchi . N. . Higham . T. F. G. . Wheeler . H. T. . Rosendahl . W. . 2009 . Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity . Molecular Ecology . 18 . 8 . 1668–77 . 10.1111/j.1365-294X.2009.04134.x . 19302360 . 2009MolEc..18.1668B . 46716748 . 7 March 2018 . 20 September 2018 . https://web.archive.org/web/20180920011400/https://www.researchgate.net/publication/24216045 . live .
- Sotnikova, M. . Nikolskiy, P. . 2006 . Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters . Quaternary International . en . 142–143 . 218–228 . 10.1016/j.quaint.2005.03.019 . 2006QuInt.142..218S . 24 January 2023 . 5 February 2023 . https://web.archive.org/web/20230205061457/https://doc.rero.ch/record/13484/files/PAL_E260.pdf . live .
- Christiansen . P. . Harris . J. M. . 2009 . Craniomandibular morphology and phylogenetic affinities of Panthera atrox: implications for the evolution and paleobiology of the lion lineage . Journal of Vertebrate Paleontology . 29 . 3 . 934–945 . 2009JVPal..29..934C . 10.1671/039.029.0314 . 85975640.
- O’Keefe . F. Robin . Dunn . Regan E. . Weitzel . Elic M. . Waters . Michael R. . Martinez . Lisa N. . Binder . Wendy J. . Southon . John R. . Cohen . Joshua E. . Meachen . Julie A. . DeSantis . Larisa R. G. . Kirby . Matthew E. . Ghezzo . Elena . Coltrain . Joan B. . Fuller . Benjamin T. . Farrell . Aisling B. . 18 August 2023 . Pre–Younger Dryas megafaunal extirpation at Rancho La Brea linked to fire-driven state shift . Science . en . 381 . 6659 . 10.1126/science.abo3594 . 37590347 . 260956289 . 0036-8075 . 11 March 2024 . 27 January 2024 . https://web.archive.org/web/20240127153230/https://www.science.org/doi/10.1126/science.abo3594 . live .
- Stinnesbeck, S. R. . Stinnesbeck, W. . Frey, E. . Olguín, J. A. . Sandoval, C. R. . Morlet, A. V. . González, A. H. . 2018 . Panthera balamoides and other Pleistocene felids from the submerged caves of Tulum, Quintana Roo, Mexico . Historical Biology: An International Journal of Paleobiology . 32 . 7. 1–10 . 10.1080/08912963.2018.1556649. 92328512 .
- Blaine W. Schubert . James C. Chatters . Joaquin Arroyo-Cabrales . Joshua X. Samuels . Leopoldo H. Soibelzon . Francisco J. Prevosti . Christopher Widga . Alberto Nava . Dominique Rissolo . Pilar Luna Erreguerena . 2019 . Yucatán carnivorans shed light on the Great American Biotic Interchange . Biology Letters . 15 . 5 . Article ID 20190148 . 10.1098/rsbl.2019.0148 . 6548739 . 31039726.
- Marciszak . A. . 2014 . Presence of Panthera gombaszoegensis (Kretzoi, 1938) in the late Middle Pleistocene of Biśnik Cave, Poland, with an overview of Eurasian jaguar size variability . Quaternary International . 326-327 . 105–113 . 2014QuInt.326..105M . 10.1016/j.quaint.2013.12.029.
- Hemmer, H. . Kahlke, R. D. . Vekua, A. K. . 2010 . Panthera onca georgica ssp. nov. from the Early Pleistocene of Dmanisi (Republic of Georgia) and the phylogeography of jaguars (Mammalia, Carnivora, Felidae) . Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen . 257. 1. 115–127 . 10.1127/0077-7749/2010/0067 .
- Mol, D. . van Logchem, W. . de Vos, J. . 2011 . New record of the European jaguar, Panthera onca gombaszoegensis (Kretzoi, 1938), from the Plio-Pleistocene of Langenboom (The Netherlands) . Cainozoic Research . 8 . 1–2 . 35–40 . 28 September 2015 . 4 April 2019 . https://web.archive.org/web/20190404200141/http://natuurtijdschriften.nl/download?type=document;docid=541752 . live .
- Harington . C. R. . 1996 . Pleistocene mammals of the Yukon Territory . PhD . University of Alberta . Edmonton.
- Sabo . Martin . Tomašových . Adam . Gullár . Juraj . August 2022 . Geographic and temporal variability in Pleistocene lion-like felids: Implications for their evolution and taxonomy . Palaeontologia Electronica . English . 25 . 2 . 1–27 . 10.26879/1175 . 1094-8074 . 251855356 . free . 11 March 2024 . 2 April 2023 . https://web.archive.org/web/20230402015510/https://palaeo-electronica.org/content/2022/3681-lion-cranial-variability . live .
- Stuart, A. J., Lister, A .M. . 2011. Extinction chronology of the cave lion Panthera spelaea. Quaternary Science Reviews . 30 . 17 . 2329–2340 . 10.1016/j.quascirev.2010.04.023. 2011QSRv...30.2329S.
- Sala, B. . 1990 . Panthera leo fossilis (v. Reichenau, 1906) (Felidae) de Iserna la Pineta (Pléistocene moyen inférieur d'Italie) . Géobios . 23 . 2 . 189–194 . 10.1016/S0016-6995(06)80051-3 .
- Marciszak . A. . Stefaniak . K. . 2010 . 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 . Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen . 258 . 3 . 339–351 . 10.1127/0077-7749/2010/0117 . 7 March 2018 . 25 September 2018 . https://web.archive.org/web/20180925142142/https://www.researchgate.net/publication/233669138 . live .
- Marciszak . A. . Schouwenburg . C. . Darga . R. . 2014 . Decreasing size process in the cave (Pleistocene) lion Panthera spelaea (Goldfuss, 1810) evolution – A review . Quaternary International . Fossil remains in karst and their role in reconstructing Quaternary paleoclimate and paleoenvironments . 339–340 . 245–257 . 10.1016/j.quaint.2013.10.008. 2014QuInt.339..245M .
- Sotnikova, M. V. . Foronova, I. V. . First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia. . 2014. Integrative Zoology . 9 . 4 . 517–530 . 10.1111/1749-4877.12082 . 24382145 .
- Barnett, R. . Mendoza, M. L. Z. . Soares, A. E. R. . Ho, S. Y. W. . Zazula, G. . Yamaguchi, N. . Shapiro, B. . Kirillova, I. V. . Larson, G. . Gilbert, M. T. P. . Mitogenomics of the Extinct Cave Lion, Panthera spelaea (Goldfuss, 1810), resolve its position within the Panthera cats . 2016 . Open Quaternary . 2 . 4 . 10.5334/oq.24. free . 10576/22920 . free .
- Stuart . Anthony J. . Lister . Adrian M. . August 2011 . Extinction chronology of the cave lion Panthera spelaea . Quaternary Science Reviews . en . 30 . 17–18 . 2329–2340 . 10.1016/j.quascirev.2010.04.023 . 2011QSRv...30.2329S . 11 March 2024 . 24 April 2024 . https://web.archive.org/web/20240424144240/https://linkinghub.elsevier.com/retrieve/pii/S0277379110001320 . live .
- Manamendra-Arachchi, K., Pethiyagoda, R., Dissanayake, R., Meegaskumbura, M. . 2005 . A second extinct big cat from the late Quaternary of Sri Lanka . The Raffles Bulletin of Zoology . Supplement 12 . 423–434 . 28 October 2013 . 7 August 2007 . https://web.archive.org/web/20070807215533/http://rmbr.nus.edu.sg/rbz/biblio/s12/s12rbz423-434.pdf .
- Ruiz-Garcia, M. . Payan, E. . Murillo, A. . Alvarez, D. . amp . 2006 . DNA microsatellite characterization of the jaguar (Panthera onca) in Colombia . Genes & Genetic Systems . 81 . 2 . 115–127 . 10.1266/ggs.81.115 . 16755135 . 8 September 2015 . free . 6 June 2023 . https://web.archive.org/web/20230606112430/https://www.jstage.jst.go.jp/article/ggs/81/2/81_2_115/_pdf . live .
- Moreno, A. . Lima-Ribeiro, M. . 2015 . Ecological niche models, fossil record and the multi-temporal calibration for Panthera onca (Linnaeus, 1758) (Mammalia: Felidae) . Brazilian Journal of Biological Sciences . 2 . 4 . 309–319 . 4 April 2019 . 4 April 2019 . https://web.archive.org/web/20190404195921/http://revista.rebibio.net/v2n4/v02n04a14.pdf . live .
- Roth . S. . 1899 . Descripción de los restos encontrados en la caverna de Última Esperanza . Revista del Museo la Plata . 9 . 381–388 .
- Paijmans, J. L. A. . Barlow, A. . Förster, D. W. . Henneberger, K. . Meyer, M. . Nickel, B. . Nagel, D. . Havmøller, R. W. . Baryshnikov, G. F. . Joger, U. . Rosendahl, W. . Hofreiter, M. . 2018 . Historical biogeography of the leopard (Panthera pardus) and its extinct Eurasian populations . BMC Evolutionary Biology . 18 . 1 . 156 . 10.1186/s12862-018-1268-0 . 30348080 . 6198532 . free . 2018BMCEE..18..156P .
- Sabol, M.. 2011 . Masters of the lost world: a hypothetical look at the temporal and spatial distribution of lion-like felids . Quaternaire . 4 . 229–236 .
- Hasegawa . Y. . Tomida . Y. . Kohno . N. . Ono . K. . Nokariya . H. . Uyeno . T. . 1988 . Quaternary vertebrates from Shiriya area, Shimokita Pininsula, northeastern Japan . Memoirs of the National Science Museum . 21 . 17–36.
- Hemmer . H. . 2022 . 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) . Palaeobiodiversity and Palaeoenvironments . 103 . 2 . 465–487 . 10.1007/s12549-022-00542-2 . 2023PdPe..103..465H .
- Bakr . A. . 1986 . On a collection of Siwalik Carnivora . Biological Society of Pakistan. 11. 1–64 .
- Turner, A.. 1984. Panthera crassidens Broom, 1948. The cat that never was?. South African Journal of Science. 80. 5. 227–233 .
- Mazák . J. H.. Christiansen. P. . Kitchener. A. C. . Oldest Known Pantherine Skull and Evolution of the Tiger . PLOS ONE. 6. 10. 2011. e25483 . 1932-6203 . 10.1371/journal.pone.0025483 . 22016768 . 3189913 . 2011PLoSO...625483M. free.
- Piras, P. . Silvestro, D. . Carotenuto, F. . Castiglione, S.. Kotsakis, A. . Maiorino, L. . Melchionna, M. . Mondanaro, A. . Sansalone, G. . Serio, C. . Vero, V.A. . Raia, P. . 2018 . Evolution of the sabertooth mandible: A deadly ecomorphological specialization . . 496 . 166−174 . 10.1016/j.palaeo.2018.01.034. 2018PPP...496..166P . 2158/1268434 . free.