Open main menu

Wikipedia β

Panthera is a genus within the Felidae family that was named and first described by the German naturalist Oken in 1816.[2] The British taxonomist Pocock revised the classification of this genus in 1916 as comprising the species lion, tiger, jaguar, and leopard on the basis of cranial features.[3] Results of genetic analysis indicate that the snow leopard also belongs to the Panthera, a classification that was accepted by IUCN assessors in 2008.[4][5]

Temporal range: Late Miocene – present, 5.95–0 Ma
An Indian tiger in the wild. Royal, Bengal tiger (27466438332).jpg
Tiger (Panthera tigris), the largest species of the Panthera genus
Female Liger 1904.jpg
The largest member of the genus Panthera, the liger, between its two parent species, the tiger (left) and lion (right)
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Oken, 1816
Type species
Felis pardus
Linnaeus, 1758
Extant species

Panthera tigris
Panthera uncia
Panthera onca
Panthera leo
Panthera pardus

Only the tiger, lion, leopard, and jaguar have the anatomical structure that enables them to roar. The primary reason for this was formerly assumed to be the incomplete ossification of the hyoid bone. However, new studies show the ability to roar is due to other morphological features, especially of the larynx. The snow leopard does not roar. Although it has an incomplete ossification of the hyoid bone, it lacks the special morphology of the larynx.[6]



The word panther derives from classical Latin panthēra, itself from the ancient Greek pánthēr (πάνθηρ).[7] The Greek pan- (πάν), meaning "all", and thēr (θήρ), meaning "prey" bears the meaning of "predator of all animals". Use of the word for a beast originated in antiquity in the Orient, probably from India to Persia to Greece.[8]


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 basicranial 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.[9] All Panthera species have an incompletely ossified hyoid bone. Specially adapted larynx with proportionally larger vocal folds are covered in a large fibro-elastic pad. These characteristics enable all Panthera species except snow leopard to roar.[10]


Panthera probably evolved in Asia, but the roots of the genus remain unclear. Genetic studies indicate that pantherine cats diverged from the subfamily Felinae between six and ten million years ago.[4] Fossil records that appear to belong within the Panthera genus reach only 2.0 to 3.8 million years back.[11]

The snow leopard was initially seen at the base of Panthera, but newer molecular studies suggest that it is nestled within Panthera and is a sister species of the tiger.[12] Many place the snow leopard within the genus Panthera, but there is currently no consensus as to whether the snow leopard should retain its own genus Uncia or be moved to Panthera uncia.[4][13][14][15] Since 2008, the IUCN Red List lists it as Panthera uncia using Uncia uncia as a synonym.[5]

The genus Neofelis is generally placed at the base of the Panthera group, but is not included in the genus itself.[4][14][15][16]

Results of a mitogenomic study suggest the phylogeny can be represented as Neofelis nebulosa (Panthera tigris (Panthera onca (Panthera pardus, (Panthera leo, Panthera uncia)))).[17] About 11.3 million years ago Panthera separated from other felid species and then evolved into the several species of the genus. N. nebulosa appears to have diverged about 8.66 million years ago, P. tigris about 6.55 million years ago, P. uncia about 4.63 million years ago and P. pardus about 4.35 million years ago. Mitochondrial sequence data from fossils suggest that the American lion (P. l. atrox) is a sister lineage to Upper Pleistocene Eurasian cave lion (P. l. spelaea) that diverged about 0.34 million years ago.[18]

The prehistoric cat Panthera onca gombaszogensis, often called European jaguar is probably closely related to the modern jaguar. The earliest evidence of the species was obtained at Olivola in Italy, and dates 1.6 million years.[19]


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 Pocock reviewed skins and skulls in the zoological collection of the Natural History Museum, London and grouped subspecies described, thus shortening the lists considerably.[20][21][22] Since the mid-1980s, several Panthera species became subject 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]

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]


Two cladograms proposed for Panthera. The upper one is based on phylogenetic studies by Johnson et al. (2006),[4] and by Werdelin et al. (2010).[31] The lower cladogram is based on a study by Davis et al. (2010)[32] and by Mazák et al. (2011).[33]

The cladogram below follows Mazák, Christiansen and Kitchener (2011).[33]




Panthera uncia 

Panthera palaeosinensis

Panthera onca 

Panthera atrox

Panthera spelaea

Panthera leo 

Panthera pardus 

Panthera tigris 

Panthera zdanskyi

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:[34]


Panthera palaeosinensis

Panthera blytheae

Panthera uncia  

Panthera zdanskyi

Panthera tigris  

Panthera gombaszoegensis

Panthera onca  

Panthera pardus  

Panthera leo  

Panthera spelaea

Panthera atrox

Contemporary speciesEdit

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:[1][35]

Species Subspecies Current distribution
Tiger P. tigris


Tigers of mainland Asia P. t. tigris including:[35]

Sunda tiger P. t. sondaica including[35]

India, Nepal, Bangladesh, Bhutan, Myanmar, Thailand, Cambodia, eastern Russia and China, Indonesian island of Sumatra
Lion P. leo


Northern lion P. l. leo including:[35]

Southern African lion P. l. melanochaita including:[35]

West, East, Central and Southern Africa, India
Jaguar P. onca


None[42][35] Historic range stretches across the USA and Mexico to Chile and Argentina, including much of Amazonian Brazil.[43] Countries present in the modern range are Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica (particularly on the Osa Peninsula), Ecuador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, Paraguay, Peru, Suriname, the United States and Venezuela.
Leopard P. pardus


African leopard P. p. pardus

Arabian leopard P. p. nimr
Javan leopard P. p. melas
Indian leopard P. p. fusca
Sri Lankan leopard P. p. kotiya
Persian leopard and Anatolian leopard P. p. tulliana,[35] syn. P. p. ciscaucasica, P. p. saxicolor[1]
Indochinese leopard P. p. delacouri
Amur leopard P. p. orientalis, including P. p. japonensis[35]

Africa, Middle East, South and Southeast Asia, East Asia
Snow leopard P. uncia[35]


Kazakhstan, Kyrgyzstan, Tajikistan, and Uzbekistan to the Hindu Kush in eastern Afghanistan, Karakoram in northern Pakistan, in the Pamir Mountains, and in the high altitudes of the Himalayas in India, Nepal, and Bhutan, and the Tibetan Plateau.

Fossil species and subspeciesEdit

Species Subspecies Fossil distribution Notes
Panthera palaeosinensis None Northern China Originally thought to be an ancestral species of tiger, this species has been found in recent studies place it close to the base of the genus Panthera.[33][44]
Panthera blytheae None Tibetan Plateau The oldest known species of Panthera closely related to the snow leopard.[45]
Panthera zdanskyi None Gansu province of northwestern China Closely related to the tiger.[46]
Tiger P. tigris P. t. acutidens[47]

P. t. trinilensis[48]
P. t. soloensis[49]

Much of Asia (P. t. acutidens); Java, Indonesia (P. t. trinilensis and P. t. soloensis) These three subspecies are not closely related to the modern subspecies of tigers.[50]
Panthera toscana


P. t. toscana

P. t. georgica[51]

Italy (P. t. toscana) and Georgia (P. t. georgica) The oldest fossil species of jaguar (sometimes classified as fossil subspecies of the extant animal or P. gombaszoegensis).[51]
Panthera gombaszoegensis


P. g. schreuderi

P. g. gombaszoegensis

Europe P. g. schreuderi might be a junior synonym of P. g. gombaszoegensis.[52][51][53] Sometimes classified as subspecies of the P. onca.[51][53]
†Giant jaguars Panthera onca


P. o. augusta[54]

P. o. mesembrina[55]

North America (P. o. augusta) and South America (P. o. mesembrina) P. onca represented the third wave of jaguar evolution,[51] with P. o. augusta in the temperate forests of North America, P. o. mesembrina in temperate grasslands of South America, and the smaller modern jaguar, P. o. onca, inhabiting the tropics of both continents during the Pleistocene.[43]
European leopards P. pardus P. p. begoueni[56]

P. p. sickenbergi[57]
P. p. antiqua[58]
P. p. begoueni[56]
P. p. spelaea[59]

Europe Closely related to Persian leopard P. p. tulliana according to genetic work.[59]
Panthera crassidens None South Africa A possible chimera of fossil leopards and cheetahs.[60]
Panthera shawi None Laetoli site in Tanzania A leopard-like cat which is believed to be the oldest known species of lion.[61]
Lion P. leo P. l. sinhaleyus Sri Lanka This subspecies of lion is only known by two teeth.[62]
P. spelaea


P. s. fossilis[63]

P. s. youngi[64]
P. s. spelaea

Iberian peninsula, Southeast Europe, Great Britain, Central Europe, East European Plain, across most of northern Eurasia into Canada and Alaska[65] Originally spelaea and fossilis were classified as subspecies of the extant lion P. leo.[66] Results of recent genetic studies indicate that both belong to a distinct species, namely P. spelaea.[67][68] Other genetic results indicate that the fossilis cave lion warrants status of a species.[69][70]
Panthera atrox


None North America, dubious remains in South America.[71] The American lion is thought to have descended from a basal spelaea cave lion population isolated south of the North American continental ice sheet, and then established a mitochondrial sister clade circa 200,000 BP.[70][72] It is sometimes considered either under the nomenclature of P. leo[72] or of P. spelaea.[73]

See alsoEdit


  1. ^ a b c d e f g h i j k l m n o p q r Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 546–548. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  2. ^ Oken, L. (1816). Lehrbuch der Zoologie, 2. Abtheilung. August Schmid & Comp., Jena.
  3. ^ 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, Volume XVIII: 314–316.
  4. ^ a b c d e 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. Bibcode:2006Sci...311...73J. doi:10.1126/science.1122277. PMID 16400146. 
  5. ^ a b Jackson, R.; Mallon, D.; McCarthy, T.; Chundaway, R. A.; Habib, B. (2008). "Panthera uncia". IUCN Red List of Threatened Species. Version 2017-1. International Union for Conservation of Nature. 
  6. ^ Nowak, R. M. (1999). Walker's Mammals of the World. Johns Hopkins University Press. ISBN 0-8018-5789-9. 
  7. ^ Liddell, H. G. & R. Scott (1940). "πάνθηρ". A Greek-English Lexicon, revised and augmented. Oxford: Clarendon Press. 
  8. ^ OUP (2002). "Panther". Oxford English Dictionary. Oxford: Oxford University Press. 
  9. ^ Pocock, R. I. (1939). "Panthera". The fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1. London: Taylor and Francis. pp. 196–239. 
  10. ^ Hast, M. H. (1989). "The larynx of roaring and non-roaring cats". Journal of Anatomy. 163: 117–121. ISSN 0021-8782. PMC 1256521 . PMID 2606766. 
  11. ^ Turner, A. (1987). "New fossil carnivore remains from the Sterkfontein hominid site (Mammalia: Carnivora)". Annals of the Transvaal Museum. 34 (15): 319–347. 
  12. ^ Davis, B. W.; Li, G..; Murphy, W.J. (Jul 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. 
  13. ^ 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. 
  14. ^ a b 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. Retrieved 2006-08-06. 
  15. ^ a b 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: S98–S116. doi:10.1007/PL00000060. PMID 9071018. 
  16. ^ Yu L & Zhang YP (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. 
  17. ^ Wei L, Wu X, Zhu L, Jiang Z (2010). "Mitogenomic analysis of the genus Panthera". Science China Life Sciences. 54 (10): 917–930. doi:10.1007/s11427-011-4219-1. PMID 22038004. 
  18. ^ Barnett, R.; Shapiro, B.; Barnes, I.; Yo, 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" (PDF). Molecular Ecology. 18 (8): 1668–1677. doi:10.1111/j.1365-294X.2009.04134.x. PMID 19302360. Archived from the original (PDF) on 21 January 2012. Retrieved 23 November 2011. 
  19. ^ 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. 
  20. ^ Pocock, R. I. (1930). "The panthers and ounces of Asia". Journal of the Bombay Natural History Society. 34 (1): 65–82. 
  21. ^ 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. 
  22. ^ Pocock, R. I. (1939). "The races of jaguar (Panthera onca)". Novitates Zoologicae. 41: 406–422. 
  23. ^ O'Brien, S. J.; Martenson, J. S.; Packer, C.; Herbst, L.; de Vos, V.; Joslin, P.; Ott-Joslin, J.; Wildt, D. E. & Bush, M. (1987). "Biochemical genetic variation in geographic isolates of African and Asiatic lions" (PDF). National Geographic Research. 3 (1): 114–124. Archived from the original (PDF) on 2 May 2013. 
  24. ^ 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. 
  25. ^ 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. 
  26. ^ 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. 
  27. ^ 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. (2011). "Genetic diversity, evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa" (PDF). Journal of Biogeography. 38 (7): 1356–1367. doi:10.1111/j.1365-2699.2011.02500.x. 
  28. ^ 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. and Tende, T. (2016). "Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (Panthera leo)". Scientific Reports. 6: 30807. doi:10.1038/srep30807. 
  29. ^ Robinson, R. (1970). "Inheritance of black form of the leopard Panthera pardus". Genetica. 41 (1): 190–197. doi:10.1007/bf00958904. PMID 5480762. 
  30. ^ 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. 
  31. ^ Werdelin, L.; Yamaguchi, N.; Johnson, W.E.; O'Brien, S.J. (2010). "Phylogeny and evolution of cats (Felidae)" (PDF). Biology and Conservation of Wild Felids: 59–82. 
  32. ^ Davis, B.W., Li, G. and 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. 
  33. ^ a b c Mazák, Ji H.; Christiansen, Per; Kitchener, Andrew C. (2011). "Oldest Known Pantherine Skull and Evolution of the Tiger". Plos One. 6 (10): e25483. Bibcode:2011PLoSO...625483M. doi:10.1371/journal.pone.0025483. ISSN 1932-6203. PMC 3189913 . PMID 22016768. 
  34. ^ Piras, P.; Silvestro, D.; Carotenuto, F.; Castiglione, S.; Kotsakis, A.; Maiorino, L.; Melchionna, M.; Mondanaro, A.; Sansalone, G., Serio, C. and Vero, V.A. (2018). "Evolution of the sabertooth mandible: A deadly ecomorphological specialization". Palaeogeography, Palaeoclimatology, Palaeoecology. 496: 166−174. doi:10.1016/j.palaeo.2018.01.034. 
  35. ^ a b c d e f g h i 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" (PDF). Cat News. Special Issue 11. 
  36. ^ Jackson, P.; Nowell, K. (2008). "Panthera tigris ssp. virgata". IUCN Red List of Threatened Species. Version 2017-1. International Union for Conservation of Nature. 
  37. ^ Luo, S.J.; Kim, J.H.; Johnson, W.E.; Walt, J.v.d.; Martenson, J.; Yuhki, N.; Miquelle, D.G.; et al. (2004). "Phylogeography and Genetic Ancestry of Tigers (Panthera tigris)". PLOS Biology. 2 (12): e442. doi:10.1371/journal.pbio.0020442. PMC 534810 . PMID 15583716. 
  38. ^ Jackson, P.; Nowell, K. (2008). "Panthera tigris ssp. sondaica". IUCN Red List of Threatened Species. Version 2017-1. International Union for Conservation of Nature. 
  39. ^ Jackson, P. Nowell, K. (2008). "Panthera tigris ssp. balica". IUCN Red List of Threatened Species. Version 2017-1. International Union for Conservation of Nature. 
  40. ^ Bauer, H.; Packer, C.; Funston, P. F.; Henschel, P. & Nowell, K. (2016). "Panthera leo". The IUCN Red List of Threatened Species. IUCN. 2016: e.T15951A115130419. doi:10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en. Retrieved 15 January 2018. 
  41. ^ 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. 
  42. ^ Larson, S. E. (1997). "Taxonomic re-evaluation of the jaguar". Zoo Biology. 16 (2): 107–120. doi:10.1002/(SICI)1098-2361(1997)16:2<107::AID-ZOO2>3.0.CO;2-E. ISSN 1098-2361. 
  43. ^ a b MORENO, LIMA-RIBEIRO, Ana, Matheus (31 December 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): p. 309–319. ISSN 2358-2731. 
  44. ^ Tseng, Z. J.; Wang, X.; Slater, G. J.; Takeuchi, G. T.; Li, Q.; Liu, J.; Xie, G. (2013). "Himalayan fossils of the oldest known pantherine establish ancient origin of big cats". Proceedings of the Royal Society B: Biological Sciences. 281 (1774): 20132686–20132686. doi:10.1098/rspb.2013.2686. ISSN 0962-8452. 
  45. ^ Tseng, Z. Jack; Wang, Xiaoming; Slater, Graham J.; Takeuchi, Gary T.; Li, Qiang; Liu, Juan; Xie, Guangpu (2014-01-07). "Himalayan fossils of the oldest known pantherine establish ancient origin of big cats". Proc. R. Soc. B. 281 (1774): 20132686. doi:10.1098/rspb.2013.2686. ISSN 0962-8452. PMC 3843846 . PMID 24225466. 
  46. ^ Ji H. Mazák, Per Christiansen and Andrew C. Kitchener (2011). "Oldest Known Pantherine Skull and Evolution of the Tiger". PLoS ONE. 6 (10): e25483. doi:10.1371/journal.pone.0025483. PMC 3189913 . PMID 22016768. 
  47. ^ Hooijer, D. A. (1947). Pleistocene remains of Panthera tigris (Linnaeus) subspecies from Wanhsien, Szechwan, China, compared with fossil and recent tigers from other localities. American Museum Novitates no. 1346.
  48. ^ Brongersma, L. D. (1935). "Notes on some recent and fossil cats, chiefly from the Malay Archipelago". Zoologische Mededelingen 18: 1–89.
  49. ^ Koenigswald, G. H. R. von (1933). "Beitrag zur Kenntnis der fossilen Wirbeltiere Javas". Wetenschappelijke Mededeelingen Dienst Mijnbouw Nederlansch Oost-Indie 23: 1–127. 
  50. ^ Hasegawa, Y. Tomida, Y. Kohno, N. Ono, K. Nokariya, H., UyenoT. (1988). "Quaternary vertebrates from Shiriya area, Shimokita Pininsula, northeastern Japan". Memoirs of the National Science Museum. 21: 17–36. 
  51. ^ a b c d e 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. 
  52. ^ O'Regan, H.; Turner, A (2004). "Biostratigraphic and palaeoecological implications of new fossil felid material from the Plio-Pleistocene site of Tegelen, the Netherlands". Palaeontology. 47 (5): 1181–1193. doi:10.1111/j.0031-0239.2004.00400.x. 
  53. ^ a b 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. Retrieved 2015-09-28. 
  54. ^ 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. Retrieved 2015-09-08. 
  55. ^ Roth, S., 1899. Descripción de los restos encontrados en la caverna de Última Esperanza. Revista del Museo La Plata 9, 381–388.
  56. ^ a b Fraipont, C. (1923). "Crane de Panthère ou de Lynx géant provenent de la caverne de Trois-Frères (Ariège)". Revue d'Anthropologie 33: 42. 
  57. ^ Schütt, Von G. (1969). "Panthera pardus sickenbergi n. subsp. Aus den Mauerer Sanden". Neues Jahrbuch für Geologie und Paläontologie: 299–310. 
  58. ^ Cuvier, G. (1835). Recherches sur les ossemens fossiles ou l'on retablit les caractères de plusieurs animaux dont les revolutions du globe ont détruit les espèces. Paris: Dufour et E. d'Ocagne. 
  59. ^ a b Diedrich, C. G. (2013). "Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art". Quaternary Science Reviews 76. 76: 167–193. doi:10.1016/j.quascirev.2013.05.009. 
  60. ^ Turner, A (1984). "Panthera crassidens Broom, 1948. The cat that never was?". South African Journal of Science. 80 (5): 227–233. 
  61. ^ Sabol, M. (2011). "Masters of the lost world: a hypothetical look at the temporal and spatial distribution of lion-like felids" (PDF). Quaternaire. 4: 229–236. 
  62. ^ 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.
  63. ^ Harington, C. R. (1996). Pleistocene mammals of the Yukon Territory. Ph.D. dissertation, University of Alberta, Edmonton.
  64. ^ Harington, C. R. (1969). "Pleistocene remains of the lion-like cat (Panthera atrox) from the Yukon Territory and northern Alaska". Canadian Journal of Earth Sciences. 6 (5): 1277–1288. doi:10.1139/e69-127. 
  65. ^ Stuart, A.J., Lister, A.M. (2011). "Extinction chronology of the cave lion Panthera spelaea". Quaternary Science Reviews 30 (17): 2329–2340. 
  66. ^ Sala, B. (1990). "Panthera leo fossilis (v. Reichenau, 1906) (Felidae) de Iserna la Pineta (Pléistocene moyen inférieur d'Italie)". Géobios (23): 189–194. 
  67. ^ 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. doi:10.1127/0077-7749/2010/0117. 
  68. ^ 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. doi:10.1016/j.quaint.2013.10.008. 
  69. ^ Sotnikova, M. V.; Foronova, I. V. (2014). "First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia" (PDF). Integrative zoology. 9 (4): 517–530. 
  70. ^ a b 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. (2016). "Mitogenomics of the Extinct Cave Lion, Panthera spelaea (Goldfuss, 1810), resolve its position within the Panthera cats". Open Quaternary 2. 2: 4. doi:10.5334/oq.24. 
  71. ^ 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. 
  72. ^ a b 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" (18): 1668−1677. 
  73. ^ Sotnikova, M. and Nikolskiy, P. (2006). "Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters". Quaternary International. 142–143: 218–228. doi:10.1016/j.quaint.2005.03.019. ISSN 1040-6182. 

Further readingEdit

  • A. Turner: The big cats and their fossil relatives. Columbia University Press, 1997.ISBN 0-231-10229-1