Biology:Panthera

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Panthera[note 1] 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. Numerous extinct species are also named, including the cave lion and American lion.

Etymology

The word panther derives from Classical Latin panthēra, itself from the Ancient Greek pánthēr (πάνθηρ).[3]

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.[4] 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 9 mm (0.35 in) that provide a lower resistance to airflow; it was therefore proposed to be retained in the genus Uncia.[5] 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.[6]

Evolution

Genetic studies indicate that the pantherine cats diverged from the subfamily Felinae between six and ten million years ago.[7] The genus Neofelis is sister to Panthera.[7][8][9][10]

The geographic origin of the genus Panthera is uncertain, though the earliest known definitive species Panthera principialis is from Tanzania.[11] P. blytheae from northern Central Asia, originally described as the oldest known Panthera species, is suggested to have similar skull features to the snow leopard,[12] 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.[13][14][15] The tiger, snow leopard, and clouded leopard genetic lineages likely dispersed in Southeast Asia during the Late Miocene.[12] The clouded leopard appears to have diverged about 8.66 million years ago. Panthera diverged from other cat species about 11.3 million years ago and then evolved into the species tiger about 6.55 million years ago, snow leopard about 4.63 million years ago and leopard about 4.35 million years ago. 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 0.34 million years ago, and that both P. atrox and P. spelaea are most closely related to lions among living Panthera species.[16] The snow leopard is nested within Panthera and is the sister species of the tiger.[17]

The extinct species Panthera gombaszogensis, was probably closely related to the modern jaguar. The first fossil remains were excavated in Olivola, in Italy, and date to 1.6 million years ago.[18] Fossil remains found in South Africa that appear to belong within the Panthera lineage date to about 2 to 3.8 million years ago.[19]

Classification

Panthera was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group.[20][21] 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.[22][23][24] 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.[25] 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.[26][27] Subsequently, it was proposed to group all African leopard populations to P. p. pardus and retain eight subspecific names for Asian leopard populations.[28] 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.[7][29]

Based on genetic research, it was suggested to group all living sub-Saharan lion populations into P. l. leo.[30] 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.[31][32]

Black panthers do not form a distinct species, but are melanistic specimens of the genus, most often encountered in the leopard and jaguar.[33][34]

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:[35][36]

Species Subspecies IUCN Red List status and distribution
Lion P. leo (Linnaeus, 1758)[37]

P. l. leo (Linnaeus, 1758)[37] including:

P. l. melanochaita (Smith, 1842)[39] including:

VU[41]

frameless

Jaguar P. onca (Linnaeus, 1758)[37]

frameless

Monotypic[42][36] NT[43]

frameless

Leopard P. pardus (Linnaeus, 1758)[37]

frameless

African leopard P. p. pardus (Linnaeus, 1758)[37]

Indian leopard P. p. fusca (Meyer, 1794)[44]
Javan leopard P. p. melas (G. Cuvier, 1809)[45]
Arabian leopard P. p. nimr (Hemprich and Ehrenberg), 1833[46]
P. p. tulliana (Valenciennes, 1856),[47] syn. P. p. ciscaucasica (Satunin, 1914),[48] P. p. saxicolor Pocock, 1927[49]
Amur leopard P. p. orientalis (Schlegel, 1857),[50] syn. P. p. japonensis (Gray, 1862)[51]
Indochinese leopard P. p. delacouri Pocock, 1930[52]
Sri Lankan leopard P. p. kotiya Deraniyagala, 1956[53]

VU[54]

frameless

Tiger P. tigris (Linnaeus, 1758)[37]

frameless

P. t. tigris (Linnaeus, 1758) including:

Sunda Island tiger P. t. sondaica Temminck, 1844)[56] including

EN[61]

frameless

Snow leopard P. uncia[36] (Schreber, 1775)[62]

frameless

Monotypic[36] VU[29]

frameless

Extinct species and subspecies

Species and subspecies Fossil records Notes Images
Panthera atrox North America, 0.13 to 0.013 MYA, with dubious remains in South America.[63]
Panthera balamoides[64] Mexico, ~0.13 MYA Dubious, other authors suggest that the remains are actually of the extinct bear Arctotherium instead.[65]
Panthera fossilis[66] 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.[67]
Panthera gombaszogensis 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.[68] 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. gombaszogensis. It is occasionally classified as a subspecies of P. onca.[69][70]
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[71] At least three recent studies considered Panthera zdanskyi likely to be a synonym of P. palaeosinensis.[72][73][74]
Panthera principialis Tanzania, ~3.7 MYA Described in 2023.[11]
Panthera shawi Laetoli site in Tanzania, ~3 MYA A leopard-like cat[75]
Panthera spelaea Much of Eurasia, 0.6 to 0.013 MYA[76]
Panthera youngi[77] China, Japan, ~0.35 MYA
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.[71] But it is possibly synonymous with P. palaeosinensis.[11][78]
Panthera leo sinhaleyus Sri Lanka This lion subspecies was described on the basis of two teeth.[79]
Panthera onca augusta[80] North America May have lived in temperate forests across North America[81]
Panthera onca mesembrina[82] South America May have lived in grasslands in South America, unlike the modern jaguar
Panthera pardus spelaea Europe Closely related to Asiatic leopard subspecies,[83]
Panthera tigris acutidens Much of Asia Not closely related to modern tiger subspecies[84]
Panthera tigris soloensis Java, Indonesia Not closely related to modern tiger subspecies[84]
Panthera tigris trinilensis Java, Indonesia Not closely related to modern tiger subspecies[84]

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.[85] A "Panthera dhokpathanensis" was briefly referenced in 1986 in a report on apparent new carnivorans from the Dhok Patha region in the Siwaliks, but as no description was provided this name is a nomen nudum.[86]

Phylogeny

Results of a 2016 study based on analysis of biparental nuclear genomes suggest the following relationships of living Panthera species:[87]

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Two cladograms proposed for Panthera. The upper one is based on phylogenetic studies by Johnson et al. (2006),[7] and by Werdelin et al. (2010).[88] The lower cladogram is based on a study by Davis et al. (2010)[17] and by Mazák et al. (2011).[71]

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.[89]

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See also

Notes

  1. ↑ /ˈpænθərə, pænˈθiːrə/ PAN-thə-rə, pan-THEE-rə[1][2]

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References

  1. ↑ "Panthera" (in en). Collins Dictionary. Penguin Random House. 2005. https://www.collinsdictionary.com/dictionary/english/panthera. 
  2. ↑ Lua error: Internal error: The interpreter exited with status 1., spoken by Kallie Moore
  3. ↑ Liddell, H. G.; Scott, R. (1940). "πάνθηρ". A Greek-English Lexicon (Revised and augmented ed.). Oxford: Clarendon Press. https://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3D%2377441. Retrieved 21 February 2021. 
  4. ↑ Pocock, R. I. (1939). "Panthera". The fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1. London: Taylor and Francis. pp. 196–239. https://archive.org/stream/PocockMammalia1/pocock1#page/n243/mode/2up. 
  5. ↑ Hast, M. H. (1989). "The larynx of roaring and non-roaring cats". Journal of Anatomy 163: 117–121. PMID 2606766. 
  6. ↑ Weissengruber, G. E.; Forstenpointner, G.; Peters, G.; Kübber-Heiss, A.; Fitch, W. T. (2002). "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 201 (3): 195–209. doi:10.1046/j.1469-7580.2002.00088.x. PMID 12363272. 
  7. ↑ 7.0 7.1 7.2 7.3 Johnson, W.E.; Eizirik, E.; Pecon-Slattery, J.; Murphy, W.J.; Antunes, A.; Teeling, E.; O'Brien, S.J. (2006). "The Late Miocene radiation of modern Felidae: A genetic assessment". Science 311 (5757): 73–77. doi:10.1126/science.1122277. PMID 16400146. Bibcode: 2006Sci...311...73J. https://zenodo.org/record/1230866. Retrieved 28 June 2019. 
  8. ↑ Janczewski, D.N.; Modi, W.S.; Stephens, J.C.; O'Brien, S.J. (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. doi:10.1093/oxfordjournals.molbev.a040232. PMID 7544865. 
  9. ↑ Johnson, W. E.; O'Brien, S.J. (1997). "Phylogenetic reconstruction of the Felidae using 16S rRNA and NADH-5 mitochondrial genes". Journal of Molecular Evolution 44 (S1): S98–S116. doi:10.1007/PL00000060. PMID 9071018. Bibcode: 1997JMolE..44S..98J. https://zenodo.org/record/1232587. Retrieved 28 June 2019. 
  10. ↑ Yu, L.; Zhang, Y.P. (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. doi:10.1016/j.ympev.2005.01.017. PMID 15804417. Bibcode: 2005MolPE..35..483Y. 
  11. ↑ 11.0 11.1 11.2 Hemmer, H. (2023). "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. doi:10.1007/s12549-022-00542-2. Bibcode: 2023PdPe..103..465H. 
  12. ↑ 12.0 12.1 Tseng, Z.J.; Wang, X.; Slater, G.J.; Takeuchi, G.T.; Li, Q.; Liu, J.; Xie, G. (2014). "Himalayan fossils of the oldest known pantherine establish ancient origin of big cats". Proceedings of the Royal Society B: Biological Sciences 281 (1774). doi:10.1098/rspb.2013.2686. PMID 24225466. 
  13. ↑ Geraads, D.; Peigné, S (2017). "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. doi:10.1007/s10914-016-9349-6. https://hal.sorbonne-universite.fr/hal-01675275/document. 
  14. ↑ 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. doi:10.1007/s12549-023-00571-5. Bibcode: 2023PdPe..103..827H. 
  15. ↑ Jiangzuo, Q.; Madurell-Malapeira, J.; Li, X.; Estraviz-López, D.; Mateus, O.; Testu, A.; Li, S.; Wang, S. et al. (2025). "Insights on the evolution and adaptation toward high-altitude and cold environments in the snow leopard lineage". Science Advances 11 (3). doi:10.1126/sciadv.adp5243. PMID 39813339. Bibcode: 2025SciA...11P5243J. 
  16. ↑ Barnett, R.; Shapiro, B.; Barnes, I.; Ho, S.Y.W.; Burger, J.; Yamaguchi, N.; Higham, T.F.G.; Wheeler, H.T. et al. (2009). "Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity". Molecular Ecology 18 (8): 1668–1677. doi:10.1111/j.1365-294X.2009.04134.x. PMID 19302360. Bibcode: 2009MolEc..18.1668B. https://www.zin.ru/Labs/theriology/eng/staff/baryshnikov/references/barnett_et_al_2009.pdf. Retrieved 24 February 2019. 
  17. ↑ 17.0 17.1 Davis, B.W.; Li, G.; Murphy, W.J. (2010). "Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, Panthera (Carnivora: Felidae)". Molecular Phylogenetics and Evolution 56 (1): 64–76. doi:10.1016/j.ympev.2010.01.036. PMID 20138224. Bibcode: 2010MolPE..56...64D. 
  18. ↑ Hemmer, H.; Kahlke, R.D.; Vekua, A.K. (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. doi:10.1016/s0016-6995(01)80011-5. Bibcode: 2001Geobi..34..475H. 
  19. ↑ Turner, A. (1987). "New fossil carnivore remains from the Sterkfontein hominid site (Mammalia: Carnivora)". Annals of the Transvaal Museum 34 (15): 319–347. https://journals.co.za/content/nfi_annalstm/34/15/AJA00411752_121?crawler=true&mimetype=application/pdf. 
  20. ↑ Oken, L. (1816). "1. Art, Panthera". Lehrbuch der Zoologie. 2. Abtheilung. Jena: August Schmid & Comp.. p. 1052. https://books.google.com/books?id=S5o5AAAAcAAJ&pg=PA1052. 
  21. ↑ 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. http://digitallibrary.amnh.org/bitstream/handle/2246/509//v2/dspace/ingest/pdfSource/bul/B016a27.pdf?sequence=1&isAllowed=y. Retrieved 4 April 2019. 
  22. ↑ Pocock, R. I. (1930). "The panthers and ounces of Asia". Journal of the Bombay Natural History Society 34 (1): 65–82. 
  23. ↑ Pocock, R. I. (1932). "The leopards of Africa". Proceedings of the Zoological Society of London 102 (2): 543–591. doi:10.1111/j.1096-3642.1932.tb01085.x. 
  24. ↑ Pocock, R. I. (1939). "The races of jaguar (Panthera onca)". Novitates Zoologicae 41: 406–422. 
  25. ↑ 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. doi:10.1080/00222931608693854. https://archive.org/stream/ser8annalsmagazi18londuoft#page/314/mode/2up. 
  26. ↑ O'Brien, S. J.; Martenson, J. S.; Packer, C.; Herbst, L.; de Vos, V.; Joslin, P.; Ott-Joslin, J.; Wildt, D. E. et al. (1987). "Biochemical genetic variation in geographic isolates of African and Asiatic lions". National Geographic Research 3 (1): 114–124. http://www.cbs.umn.edu/sites/default/files/public/downloads/Biochemical_Genetic_Variation_in_Geographic_Isolates_of_African_and_Asiatic_Lions.pdf. 
  27. ↑ 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. doi:10.1046/j.1523-1739.1996.10041115.x. Bibcode: 1996ConBi..10.1115M. 
  28. ↑ 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. doi:10.1046/j.0962-1083.2001.01350.x. PMID 11883877. Bibcode: 2001MolEc..10.2617U. https://zenodo.org/record/1236516. Retrieved 28 June 2019. 
  29. ↑ 29.0 29.1 Lua error: Internal error: The interpreter exited with status 1.
  30. ↑ 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. doi:10.1007/s10592-004-7729-6. Bibcode: 2005ConG....6...15D. https://www.researchgate.net/publication/226066591. Retrieved 5 March 2019. 
  31. ↑ 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. et al. (2011). "Genetic diversity, evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa". Journal of Biogeography 38 (7): 1356–1367. doi:10.1111/j.1365-2699.2011.02500.x. Bibcode: 2011JBiog..38.1356B. http://dspace.learningnetworks.org/bitstream/1820/4311/1/2011_Bertola,Hooft,Vrieling,Weerd,York,Bauer,Prins,Haes,Iongh_GeneticDiversityEvolutionaryHistoryAndImplicationsForConservationOfTheLionInWestAndCentralAfrica.pdf. Retrieved 17 January 2017. 
  32. ↑ Bertola, L. D.; Jongbloed, H.; Van Der Gaag, K. J.; De Knijff, P.; Yamaguchi, N.; Hooghiemstra, H.; Bauer, H.; Henschel, P. et al. (2016). "Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (Panthera leo)". Scientific Reports 6. doi:10.1038/srep30807. PMID 27488946. Bibcode: 2016NatSR...630807B. 
  33. ↑ Robinson, R. (1970). "Inheritance of black form of the leopard Panthera pardus". Genetica 41 (1): 190–197. doi:10.1007/bf00958904. PMID 5480762. 
  34. ↑ Eizirik, E.; Yuhki, N.; Johnson, W. E.; Menotti-Raymond, M.; Hannah, S. S.; O'Brien, S. J. (2003). "Molecular Genetics and Evolution of Melanism in the Cat Family". Current Biology 13 (5): 448–453. doi:10.1016/S0960-9822(03)00128-3. PMID 12620197. Bibcode: 2003CBio...13..448E. 
  35. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named MSW3
  36. ↑ 36.0 36.1 36.2 36.3 Kitchener, A. C.; Breitenmoser-Würsten, C.; Eizirik, E.; Gentry, A.; Werdelin, L.; Wilting, A.; Yamaguchi, N.; Abramov, A. V. et al. (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. https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y. Retrieved 13 May 2018. 
  37. ↑ 37.0 37.1 37.2 37.3 37.4 37.5 Linnaeus, C. (1758). "Felis". Caroli Linnæi Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I (decima, reformata ed.). Holmiae: Laurentius Salvius. pp. 41−42. https://archive.org/stream/mobot31753000798865#page/41/mode/2up. 
  38. ↑ Lua error: Internal error: The interpreter exited with status 1.
  39. ↑ Smith, C. H. (1842). "Black maned lion Leo melanochaitus". in Jardine, W.. The Naturalist's Library. Vol. 15 Mammalia. London: Chatto and Windus. p. Plate X, 177. https://archive.org/stream/naturalistslibra15jardrich#page/176/mode/2up. 
  40. ↑ 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. 
  41. ↑ Lua error: Internal error: The interpreter exited with status 1.
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  43. ↑ Lua error: Internal error: The interpreter exited with status 1.
  44. ↑ Meyer, F. A. A. (1794). "Über de la Metheries schwarzen Panther". Zoologische Annalen. Erster Band. Weimar: Im Verlage des Industrie-Comptoirs. pp. 394–396. https://books.google.com/books?id=v4s-AAAAcAAJ&pg=PA394. 
  45. ↑ 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. 
  46. ↑ Hemprich, W.; Ehrenberg, C. G. (1830). "Felis, pardus?, nimr". in Dr. C. G. Ehrenberg. 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. pp. Plate 17. https://archive.org/details/SymbolaephysicaMammEhreA/page/n60. 
  47. ↑ Valenciennes, A. (1856). "Sur une nouvelles espèce de Panthère tué par M. Tchihatcheff à Ninfi, village situé à huit lieues est de Smyrne". Comptes rendus hebdomadaires des séances de l'Académie des sciences 42: 1035–1039. 
  48. ↑ Satunin, K. A. (1914). Opredelitel' mlekopitayushchikh Rossiiskoi Imperii. Tiflis: Tipographia Kantzelyarii Namestnichestva. 
  49. ↑ Pocock, R. I. (1927). "Description of two subspecies of leopards". Annals and Magazine of Natural History. Series 9 20 (116): 213–214. doi:10.1080/00222932708655586. 
  50. ↑ Schlegel, H. (1857). "Felis orientalis". Handleiding Tot de Beoefening der Dierkunde, Ie Deel. Breda: Boekdrukkerij van Nys. p. 23. 
  51. ↑ Gray, J. E. (1862). "Description of some new species of Mammalia". Proceedings of the Royal Zoological Society of London 30: 261−263, plate XXXIII. doi:10.1111/j.1469-7998.1862.tb06524.x. https://archive.org/details/proceedingsofgen62zool/page/262. 
  52. ↑ Pocock, R. I. (1930). "The Panthers and Ounces of Asia". Journal of the Bombay Natural History Society 34 (2): 307–336. 
  53. ↑ Deraniyagala, P. E. P. (1956). "The Ceylon leopard, a distinct subspecies". Spolia Zeylanica 28: 115–116. 
  54. ↑ Lua error: Internal error: The interpreter exited with status 1.
  55. ↑ Illiger, C. (1815). "Überblick der Säugethiere nach ihrer Verteilung über die Welttheile". Abhandlungen der Königlichen Preußischen Akademie der Wissenschaften zu Berlin 1804−1811: 39−159. http://bibliothek.bbaw.de/bbaw/bibliothek-digital/digitalequellen/schriften/anzeige/index_html?band=07-abh/18041811&seite:int=195. Retrieved 2 January 2021. 
  56. ↑ 56.0 56.1 56.2 Temminck, C. J. (1844). "Aperçu général et spécifique sur les Mammifères qui habitent le Japon et les Iles qui en dépendent". 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. Leiden: Lugduni Batavorum. https://archive.org/details/faunajaponicasi00sieb/page/43. 
  57. ↑ Hilzheimer, M. (1905). "Über einige Tigerschädel aus der Straßburger zoologischen Sammlung". Zoologischer Anzeiger 28: 594–599. https://archive.org/stream/zoologischeranze28deut#page/594/mode/2up. 
  58. ↑ Mazák, V. (1968). "Nouvelle sous-espèce de tigre provenant de l'Asie du sud-est". Mammalia 32 (1): 104−112. doi:10.1515/mamm.1968.32.1.104. 
  59. ↑ Luo, S. J.; Kim, J. H.; Johnson, W. E.; Walt, J. v. d.; Martenson, J.; Yuhki, N.; Miquelle, D. G. (2004). "Phylogeography and Genetic Ancestry of Tigers (Panthera tigris)". PLOS Biology 2 (12). doi:10.1371/journal.pbio.0020442. PMID 15583716. 
  60. ↑ Schwarz, E. (1912). "Notes on Malay tigers, with description of a new form from Bali". Annals and Magazine of Natural History Series 8 Volume 10 (57): 324–326. doi:10.1080/00222931208693243. https://archive.org/stream/annalsmagazineof8101912lond#page/324/mode/2up. 
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  63. ↑ 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. doi:10.1016/j.crpv.2017.06.009. Bibcode: 2017CRPal..16..850C. 
  64. ↑ 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. doi:10.1080/08912963.2018.1556649. 
  65. ↑ Blaine W. Schubert; James C. Chatters; Joaquin Arroyo-Cabrales; Joshua X. Samuels; Leopoldo H. Soibelzon; Francisco J. Prevosti; Christopher Widga; Alberto Nava et al. (2019). "Yucatán carnivorans shed light on the Great American Biotic Interchange". Biology Letters 15 (5). doi:10.1098/rsbl.2019.0148. PMID 31039726. 
  66. ↑ Lua error: Internal error: The interpreter exited with status 1.
  67. ↑ Sabo, Martin; Tomašových, Adam; Gullár, Juraj (August 2022). "Geographic and temporal variability in Pleistocene lion-like felids: Implications for their evolution and taxonomy" (in English). Palaeontologia Electronica 25 (2): 1–27. doi:10.26879/1175. ISSN 1094-8074. https://palaeo-electronica.org/content/2022/3681-lion-cranial-variability. Retrieved 11 March 2024. 
  68. ↑ 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. doi:10.1016/j.quaint.2013.12.029. Bibcode: 2014QuInt.326..105M. 
  69. ↑ 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. doi:10.1127/0077-7749/2010/0067. Bibcode: 2010NJGPA.257..115H. 
  70. ↑ 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. http://natuurtijdschriften.nl/download?type=document;docid=541752. Retrieved 28 September 2015. 
  71. ↑ 71.0 71.1 71.2 Mazák, J. H.; Christiansen, P.; Kitchener, A. C. (2011). "Oldest Known Pantherine Skull and Evolution of the Tiger". PLOS ONE 6 (10). doi:10.1371/journal.pone.0025483. ISSN 1932-6203. PMID 22016768. Bibcode: 2011PLoSO...625483M. 
  72. ↑ Hemmer, Helmut (2023). "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. doi:10.1007/s12549-022-00542-2. Bibcode: 2023PdPe..103..465H. 
  73. ↑ Jiangzuo, Qigao; Wang, Yuan; Ge, Junyi; Liu, Sizhao; Song, Yayun; Jin, Changzhu; Jiang, Hao; Liu, Jinyi (2023). "Discovery of jaguar from northeastern China middle Pleistocene reveals an intercontinental dispersal event". Historical Biology 35 (3): 293–302. doi:10.1080/08912963.2022.2034808. Bibcode: 2023HBio...35..293J. 
  74. ↑ Jiangzuo, Q.; Madurell-Malapeira, J.; Li, X.; Estraviz-López, D.; Mateus, O.; Testu, A.; Li, S.; Wang, S. et al. (2025). "Insights on the evolution and adaptation toward high-altitude and cold environments in the snow leopard lineage". Science Advances 11 (3). doi:10.1126/sciadv.adp5243. PMID 39813339. Bibcode: 2025SciA...11P5243J. 
  75. ↑ 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. https://www.researchgate.net/publication/287422796. 
  76. ↑ Stuart, A. J., Lister, A .M. (2011). "Extinction chronology of the cave lion Panthera spelaea". Quaternary Science Reviews 30 (17): 2329–2340. doi:10.1016/j.quascirev.2010.04.023. Bibcode: 2011QSRv...30.2329S. 
  77. ↑ Pei, W. C. (1934). "On the Carnivora from Locality 1 of Choukoutien". Palaeontologica Sinica Series C, Fascicle 1: 1−166. 
  78. ↑ Jiangzuo, Q.; Wang, Y.; Ge, J.; Liu, S.; Song, Y.; Jin, C.; Jiang, H.; Liu, J. (2023). "Discovery of jaguar from northeastern China middle Pleistocene reveals an intercontinental dispersal event". Historical Biology 35 (3): 293–302. doi:10.1080/08912963.2022.2034808. Bibcode: 2023HBio...35..293J. 
  79. ↑ 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. http://rmbr.nus.edu.sg/rbz/biblio/s12/s12rbz423-434.pdf. Retrieved 28 October 2013. 
  80. ↑ Ruiz-Garcia, M.; Payan, E.; Murillo, A.; Alvarez, D. (2006). "DNA microsatellite characterization of the jaguar (Panthera onca) in Colombia". Genes & Genetic Systems 81 (2): 115–127. doi:10.1266/ggs.81.115. PMID 16755135. https://www.jstage.jst.go.jp/article/ggs/81/2/81_2_115/_pdf. Retrieved 8 September 2015. 
  81. ↑ 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. http://revista.rebibio.net/v2n4/v02n04a14.pdf. Retrieved 4 April 2019. 
  82. ↑ Roth, S. (1899). "Descripción de los restos encontrados en la caverna de Última Esperanza". Revista del Museo la Plata 9: 381–388. 
  83. ↑ Paijmans, J. L. A.; Barlow, A.; Förster, D. W.; Henneberger, K.; Meyer, M.; Nickel, B.; Nagel, D.; Havmøller, R. W. et al. (2018). "Historical biogeography of the leopard (Panthera pardus) and its extinct Eurasian populations". BMC Evolutionary Biology 18 (1): 156. doi:10.1186/s12862-018-1268-0. PMID 30348080. Bibcode: 2018BMCEE..18..156P. 
  84. ↑ 84.0 84.1 84.2 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. 
  85. ↑ Turner, A. (1984). "Panthera crassidens Broom, 1948. The cat that never was?". South African Journal of Science 80 (5): 227–233. https://journals.co.za/docserver/fulltext/sajsci/80/5/7798.pdf?expires=1544536360&id=id&accname=guest&checksum=1BB15474E5A55CD94EC66579004AE19D. 
  86. ↑ Bakr, Abu (1986). "105: On a collection of Siwalik Carnivora". Proceedings of Fourth Pakistan Congress of Zoology held under the auspices of the Zoological Society of Pakistan, University of Peshawar, Peshawar, December 20-22, 1983. Biological Society of Pakistan. p. 51. OCLC 62447645. 
  87. ↑ Li, G.; Davis, B. W.; Eizirik, E.; Murphy, W. J. (2016). "Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae)". Genome Research 26 (1): 1–11. doi:10.1101/gr.186668.114. PMID 26518481. 
  88. ↑ Werdelin, L.; Yamaguchi, N.; Johnson, W. E.; O'Brien, S. J. (2010). "Phylogeny and evolution of cats (Felidae)". Biology and Conservation of Wild Felids. Oxford: Oxford University Press. pp. 59–82. ISBN 978-0-19-923445-5. https://www.researchgate.net/publication/266755142. Retrieved 3 February 2019. 
  89. ↑ Piras, P.; Silvestro, D.; Carotenuto, F.; Castiglione, S.; Kotsakis, A.; Maiorino, L.; Melchionna, M.; Mondanaro, A. et al. (2018). "Evolution of the sabertooth mandible: A deadly ecomorphological specialization". Palaeogeography, Palaeoclimatology, Palaeoecology 496: 166−174. doi:10.1016/j.palaeo.2018.01.034. Bibcode: 2018PPP...496..166P. 

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