Last updated

Panthera [1]
Temporal range: Late Miocene – present, 5.95–0  Ma
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Oken, 1816
Type species
Panthera pardus [2]
Extant species

Panthera tigris
Panthera uncia
Panthera onca
Panthera leo
Panthera pardus


Panthera is a genus within the family Felidae that was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group. [3] [2] Reginald Innes Pocock revised the classification of this genus in 1916 as comprising the species tiger (P. tigris), lion (P. leo), jaguar (P. onca), and leopard (P. pardus) on the basis of common cranial features. [4] Results of genetic analysis indicate that the snow leopard (formerly Uncia uncia) also belongs to the Panthera (P. uncia), a classification that was accepted by IUCN Red List assessors in 2008. [5] [6]

The tiger, lion, leopard, and jaguar are the only cat species with 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 its hyoid bone is incompletely ossified, it lacks the special morphology of the larynx. [7]


The word panther derives from classical Latin panthēra, itself from the ancient Greek pánthēr (πάνθηρ). [8] The phonetically similar Sanskrit word पाण्डर pând-ara means 'pale yellow, whitish, white'. [9]


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. [10] 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. [11] 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. [12]


The geographic origin of the Panthera is most likely northern Central Asia. Panthera blytheae , the oldest known Panthera species, is similar in skull features to the snow leopard. The tiger, snow leopard, and clouded leopard genetic lineages dispersed in Southeast Asia during the Miocene. [13] Genetic studies indicate that the pantherine cats diverged from the subfamily Felinae between six and ten million years ago. [5] The genus Neofelis is sister to Panthera. [5] [14] [15] [16] 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 P. spelaea that diverged about 0.34  million years ago. [17] The snow leopard is nestled within Panthera and is the sister species of the tiger. [18]

Results of a study based on analysis of biparental nuclear genomes suggest the following relationships of living Panthera species: [19]







Snow leopard



The prehistoric European jaguar Panthera onca gombaszogensis is 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. [20] Fossil remains found in South Africa that appear to belong within the Panthera lineage date to about 2 to 3.8 million years ago . [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 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] 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. [25] [26] Subsequently, it was proposed to group all African leopard populations to P. p. pardus and retain eight subspecific names for Asian leopard populations. [27]

Based on genetic research, it was suggested to group all living sub-Saharan lion populations into P. l. leo. [28] 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. [29] [30]

Black panthers do not form a distinct species, but are melanistic specimens of the genus, most often encountered in the leopard and jaguar. [31] [32]

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: [1] [33]

SpeciesSubspecies IUCN Red List status and distribution
Lion P. leo(Linnaeus, 1758) [34]

Lion d'Afrique.jpg

P. l. leo (Linnaeus, 1758) [34] including:

P. l. melanochaita (Smith, 1842) [36] including:

VU [38]

Lion distribution.png

Jaguar P. onca(Linnaeus, 1758) [34]

Jaguar (Panthera onca palustris) male Three Brothers River 2.jpg

Monotypic [39] [33] NT [40]

Panthera onca distribution.svg

Leopard P. pardus(Linnaeus, 1758) [34]

African leopard, Panthera pardus pardus, near Lake Panic, Kruger National Park, South Africa (19448654130).jpg

African leopard P. p. pardus(Linnaeus, 1758) [34]

Indian leopard P. p. fusca(Meyer, 1794) [41]
Javan leopard P. p. melas(G. Cuvier, 1809) [42]
Arabian leopard P. p. nimr(Hemprich and Ehrenberg), 1833 [43]
Anatolian leopard or Persian leopard P. p. tulliana(Valenciennes, 1856), [44] syn. P. p. ciscaucasica (Satunin, 1914), [45] P. p. saxicolor Pocock, 1927 [46]
Amur leopard P. p. orientalis(Schlegel, 1857), [47] syn. P. p. japonensis (Gray, 1862) [48]
Indochinese leopard P. p. delacouriPocock, 1930 [49]
Sri Lankan leopard P. p. kotiyaDeraniyagala, 1956 [50]

VU [51]

Leopard distribution.jpg

Tiger P. tigris(Linnaeus, 1758) [34]

Bengal tiger (Panthera tigris tigris) female 3 crop.jpg

Tigers of mainland Asia P. t. tigris(Linnaeus, 1758) including:

Sunda Island tiger P. t. sondaicaTemminck, 1844) [53] including

EN [58]

Tiger map.jpg

Snow leopard P. uncia [33] (Schreber, 1775) [59]

Schneeleopard Koeln.jpg

Monotypic [33] VU [60]

SnowLeopard distribution.jpg

Extinct species and subspecies

Species Fossil recordsNotes
Panthera blytheae Tibetan PlateauOne of the oldest known Panthera species, possibly closely related to the snow leopard. [13]
Panthera palaeosinensis Northern ChinaIt was initially thought to be an ancestral tiger species, but several scientists place it close to the base of the genus Panthera. [61]
Panthera zdanskyi Gansu province of northwestern China Possibly a close relative of the tiger. [61]
Panthera youngi [62] China, Japan
Panthera atrox North America, dubious remains in South America. [63] 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. [64] It was sometimes considered a subspecies either under the nomenclature of P. leo [64] or P. spelaea. [65]
Panthera balamoides [66] Mexico
Panthera crassidens South AfricaNo longer a valid species due to being described based on a mixture of leopard and cheetah fossils.
Panthera gombaszoegensis EuropePanthera schreuderi and Panthera toscana are considered junior synonyms of P. gombaszoegensis. It is occasionally classified as a subspecies of P. onca. [67] [68]
Panthera leo fossilis [69] Europe
Panthera spelaea Much of Eurasia [70] Originally spelaea was classified as a subspecies of the extant lion P. leo. [71] Results of recent genetic studies indicate that it belongs to a distinct species, namely P. spelaea. [72] [73] Other genetic results indicate that the fossilis cave lion also warrants status as a species. [74] [75]
Panthera leo sinhaleyus Sri LankaThis lion subspecies was described on the basis of two teeth. [76]
Panthera onca augusta [77] North AmericaMay have lived in temperate forests across North America. [78]
Panthera onca mesembrina [79] South AmericaMay have lived in grasslands in South America, unlike the modern jaguar.
Panthera pardus spelaea EuropeClosely related to Asiatic leopard subspecies, [80] with at least one study suggesting that it is closely related to the Persian leopard (P. p. tulliana) according to genetic work. [81]
Panthera shawi Laetoli site in TanzaniaA leopard-like cat. [82]
Panthera tigris acutidens Much of AsiaNot closely related to modern tiger subspecies. [83]
Panthera tigris soloensis Java, IndonesiaNot closely related to modern tiger subspecies. [83]
Panthera tigris trinilensis Java, IndonesiaNot closely related to modern tiger subspecies. [83]


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

The cladogram below follows Mazák, Christiansen and Kitchener (2011). [61]


Neofelis Studienblatt Felis macroscelis Nebelparder (white background).jpg


Snow leopard Stamp-russia2014-save-russian-cats-(snow leopard).png

Panthera palaeosinensis

Jaguar Felis onca - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg

†American lion

Panthera spelaea Stamps of Moldova 2010 Panthera leo spelaea (mod).jpg

Lion Felis leo - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(White Background).jpg

Leopard Felis pardus - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg

Tiger Stamp-russia2014-save-russian-cats-(tiger).png

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: [85]


Panthera palaeosinensis

Panthera blytheae

Snow leopard Stamp-russia2014-save-russian-cats-(snow leopard).png

Panthera zdanskyi

Tiger Stamp-russia2014-save-russian-cats-(tiger).png

†European jaguar

Jaguar Felis onca - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg

Leopard Felis pardus - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg

Lion Felis leo - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(White Background).jpg

Panthera spelaea Stamps of Moldova 2010 Panthera leo spelaea (mod).jpg

†American lion

See also

Related Research Articles

Felidae Family of mammals

Felidae is a family of mammals in the order Carnivora, colloquially referred to as cats, and constitutes a clade. A member of this family is also called a felid. The term "cat" refers both to felids in general and specifically to the domestic cat.

Jaguar A large cat native to the Americas

The jaguar is a large felid species and the only extant member of the genus Panthera native to the Americas. The jaguar's present range extends from the southwestern United States and Mexico in North America, across much of Central America, and south to Paraguay and northern Argentina in South America. Though there are single cats now living within Arizona, the species has largely been extirpated from the United States since the early 20th century. It is listed as Near Threatened on the IUCN Red List; and its numbers are declining. Threats include loss and fragmentation of habitat.

A black panther is the melanistic colour variant of any Panthera, particularly of the leopard in Asia and Africa, and the jaguar in the Americas. Black panthers of both species have excess black pigments, but their typical spotted markings are also present. Melanism in the leopard is caused by a recessive allele, and in the jaguar by a dominant allele.

Leopard A large cat native to Africa and Eurasia

The leopard is one of the five extant species in the genus Panthera, a member of the Felidae. It occurs in a wide range in sub-Saharan Africa, in small parts of Western and Central Asia, on the Indian subcontinent to Southeast and East Asia. It is listed as Vulnerable on the IUCN Red List because leopard populations are threatened by habitat loss and fragmentation, and are declining in large parts of the global range. In Hong Kong, Singapore, Kuwait, Syria, Libya, Tunisia and most likely in Morocco, leopard populations have already been extirpated. Contemporary records suggest that the leopard occurs in only 25% of its historical global range.

The term "bigcat" is typically used to refer to any of the five living members of the genus Panthera, namely tiger, lion, jaguar, leopard, and snow leopard. Except the snow leopard, these species are able to roar. A more liberal and expansive definition of the term includes species outside of Panthera including the cougar, clouded leopard, Sunda clouded leopard, cheetah and sometimes the several lynx species, although these added species also do not roar.


Neofelis is a genus comprising two extant cat species from Southeast Asia: the clouded leopard of mainland Asia, and the Sunda clouded leopard of Sumatra and Borneo.

<i>Panthera spelaea</i> Extinct cave lion species

Panthera spelaea, also known as the "Eurasian cave lion", "European cave lion", or "steppe lion", is an extinct Panthera species that evolved in Europe probably after the third Cromerian interglacial stage, less than 600,000 years ago. Phylogenetic analysis of fossil bone samples revealed that it was highly distinct and genetically isolated from the modern lion occurring in Africa and Asia. Analysis of morphological differences and mitochondrial data support the taxonomic recognition of Panthera spelaea as a distinct species that diverged from the lion about 1.9 million years ago. Nuclear genomic evidence suggests a more recent split approximately 500,000 years ago, with no subsequent interbreeding with the ancestors of the modern lion. The oldest known bone fragments were excavated in Yakutia and radiocarbon dated at least 62,400 years old. It became extinct about 13,000 years ago.

American lion

The American lion, also known as the "North American lion", or "American cave lion", is an extinct pantherine cat that lived in North America during the Pleistocene epoch and the early Holocene epoch, about 340,000 to 11,000 years ago. Its fossils have been excavated from Alaska to Mexico. Genetic analysis has shown that the American lion and the Late Pleistocene Eurasian cave lion are sister lineages. It was about 25% larger than the modern lion, making it one of the largest known felids.


Pantherinae is a subfamily within the family Felidae, which was named and first described by Reginald Innes Pocock in 1917. The Pantherinae and the Felinae diverged from a common ancestor between 10.8 and 11.5 million years ago.

African leopard Leopard subspecies

The African leopard is the nominate subspecies of the leopard, native to many countries in Africa. It is widely distributed in most of sub-Saharan Africa, but the historical range has been fragmented in the course of habitat conversion. Leopards have been recorded in North Africa as well.

Arabian leopard Leopard subspecies in the Arabian Peninsula

The Arabian leopard is a leopard subspecies native to the Arabian Peninsula. It has been listed as Critically Endangered on the IUCN Red List since 1996 as fewer than 200 wild individuals were estimated to be alive in 2006. The population is severely fragmented. Subpopulations are isolated and not larger than 50 mature individuals. The population is thought to decline continuously.

<i>Panthera gombaszoegensis</i> Extinct European jaguar species

Panthera gombaszoegensis, also known as the European jaguar, is a Panthera species that lived from about 2.0 to 0.35 million years ago in Europe. The first fossils were excavated in 1938 in Gombaszög, Slovakia.

Panthera youngi is a fossil cat species that was described in 1934; fossil remains of this cat were excavated in a Sinanthropus formation in Choukoutien, northeastern China. Upper and lower jaws excavated in Japan's Yamaguchi Prefecture were also attributed to this species. It is estimated to have lived about 350,000 years ago in the Pleistocene epoch. It was suggested that it was conspecific with Panthera atrox and P. spelaea due to their extensive similarities. Some dental similarities were also noted with the older P. fossilis, however, Panthera youngi showed more derived features.

Persian leopard Leopard subspecies

The Persian leopard, also known as the Caucasian leopard is a leopard population in the Caucasus, Iran, Afghanistan and Central Asia. The Persian leopard was previously considered a distinct subspecies, Panthera pardus saxicolor or Panthera pardus ciscaucasica, but is now assigned to the subspecies Panthera pardus tulliana, which also includes the Anatolian leopard in Turkey. The Persian leopard is listed as Endangered on the IUCN Red List, as the population is estimated at fewer than 871–1,290 mature individuals and considered declining.

Panthera leo fossilis is a fossil cat of the genus Panthera, which was first excavated near Mauer in Germany, and lived during the Upper Pleistocene. Bone fragments of P. l. fossilis were also excavated near Pakefield in the United Kingdom, which are estimated at 680,000 years old. Bone fragments excavated near Isernia in Italy are estimated at between 600,000 and 620,000 years old. The first Asian record of a fossilis lion was found in the Kuznetsk Basin in western Siberia and dates to the late Early Pleistocene.

Indian leopard Leopard subspecies

The Indian leopard is a leopard subspecies widely distributed on the Indian subcontinent. The species Panthera pardus is listed as Vulnerable on the IUCN Red List because populations have declined following habitat loss and fragmentation, poaching for the illegal trade of skins and body parts, and persecution due to conflict situations.

Amur leopard Leopard subspecies in Far East Asia

The Amur leopard is a leopard subspecies native to the Primorye region of southeastern Russia and northern China. It is listed as Critically Endangered on the IUCN Red List. In 2007, only 19–26 wild leopards were estimated to survive in southeastern Russia and northeastern China. It was considered as one of the rarest cats on Earth.

Panthera pardus spelaea, sometimes called the European Ice Age leopard or Late Pleistocene Ice Age leopard, is a fossil leopard subspecies, which roamed Europe in the Late Pleistocene. The youngest known bone fragments date to about 32,000 to 26,000 years ago, and are similar in size to modern leopard bones.

Chinese leopard

The term Chinese leopard refers to any of the following three leopard subspecies occurring in China:

South American jaguar

The South American jaguar is a jaguar population in South America. Though a number of subspecies of jaguar have been proposed for South America, morphological and genetic research did not reveal any evidence for subspecific differentiation.


  1. 1 2 Wozencraft, W.C. (2005). "Genus Panthera". In Wilson, D.E.; Reeder, D.M (eds.). 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. 1 2 Allen, J. A. (1902). "Mammal names proposed by Oken in his 'Lehrbuch der Zoologie'" (PDF). Bulletin of the American Museum of Natural History. 16 (27): 373−379.
  3. Oken, L. (1816). "1. Art, Panthera". Lehrbuch der Zoologie. 2. Abtheilung. Jena: August Schmid & Comp. p. 1052.
  4. 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.
  5. 1 2 3 4 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. S2CID   41672825.
  6. McCarthy, T.; Mallon, D.; Jackson, R.; Zahler, P.; McCarthy, K. (2017). "Panthera uncia". IUCN Red List of Threatened Species . 2017: e.T22732A50664030. doi: 10.2305/IUCN.UK.2017-2.RLTS.T22732A50664030.en .
  7. Peters, G.; Hast, M. H. (1994). "Hyoid structure, laryngeal anatomy, and vocalization in felids (Mammalia: Carnivora: Felidae)" (PDF). Zeitschrift für Säugetierkunde. 59 (2): 87−104.
  8. Liddell, H. G. & Scott, R. (1940). "πάνθηρ". A Greek-English Lexicon (Revised and augmented ed.). Oxford: Clarendon Press.
  9. Macdonell, A. A. (1929). "पाण्डर pând-ara". A practical Sanskrit dictionary with transliteration, accentuation, and etymological analysis throughout. London: Oxford University Press. p. 95.
  10. Pocock, R. I. (1939). "Panthera". The fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1. London: Taylor and Francis. pp. 196–239.
  11. 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.
  12. 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. PMC   1570911 . PMID   12363272.
  13. 1 2 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): 20132686. doi:10.1098/rspb.2013.2686. PMC   3843846 . PMID   24225466.
  14. 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.
  15. 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. Bibcode:1997JMolE..44S..98J. doi:10.1007/PL00000060. PMID   9071018. S2CID   40185850.
  16. 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.
  17. Barnett, R.; Shapiro, B.; Barnes, I.; Ho, S.Y.W.; Burger, J.; 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. (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.
  18. 1 2 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.
  19. 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. PMC   4691742 . PMID   26518481.
  20. 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.
  21. Turner, A. (1987). "New fossil carnivore remains from the Sterkfontein hominid site (Mammalia: Carnivora)". Annals of the Transvaal Museum. 34 (15): 319–347.
  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. 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.
  26. 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.
  27. 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.
  28. 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. S2CID   30414547.
  29. 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. Archived from the original (PDF) on 8 June 2019. Retrieved 17 January 2017.
  30. 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. Bibcode:2016NatSR...630807B. doi:10.1038/srep30807. PMC   4973251 . PMID   27488946.CS1 maint: uses authors parameter (link)
  31. Robinson, R. (1970). "Inheritance of black form of the leopard Panthera pardus". Genetica. 41 (1): 190–197. doi:10.1007/bf00958904. PMID   5480762. S2CID   5446868.
  32. 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. S2CID   19021807.
  33. 1 2 3 4 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): 66−75.
  34. 1 2 3 4 5 6 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.
  35. Meyer, J. N. (1826). Dissertatio inauguralis anatomico-medica de genere felium (Doctoral thesis). Vienna: University of Vienna.
  36. Smith, C. H. (1842). "Black maned lion Leo melanochaitus". In Jardine, W. (ed.). The Naturalist's Library. Vol. 15 Mammalia. London: Chatto and Windus. p. Plate X, 177.
  37. 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.
  38. Bauer, H.; Packer, C.; Funston, P. F.; Henschel, P. & Nowell, K. (2016). "Panthera leo". IUCN Red List of Threatened Species . 2016: e.T15951A115130419.
  39. 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.
  40. Quigley, H.; Foster, R.; Petracca, L.; Payan, E.; Salom, R. & Harmsen, B. (2017). "Panthera onca". IUCN Red List of Threatened Species . 2017: e.T15953A123791436.
  41. Meyer, F. A. A. (1794). "Über de la Metheries schwarzen Panther". Zoologische Annalen. Erster Band. Weimar: Im Verlage des Industrie-Comptoirs. pp. 394–396.
  42. 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.
  43. Hemprich, W.; Ehrenberg, C. G. (1830). "Felis, pardus?, nimr". In Dr. C. G. Ehrenberg (ed.). 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.
  44. 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.
  45. Satunin, K. A. (1914). Opredelitel' mlekopitayushchikh Rossiiskoi Imperii[Guide to the mammals of the Russian Empire]. Tiflis: Tipographia Kantzelyarii Namestnichestva.
  46. 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.
  47. Schlegel, H. (1857). "Felis orientalis". Handleiding Tot de Beoefening der Dierkunde, Ie Deel. Breda: Boekdrukkerij van Nys. p. 23.
  48. 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.
  49. Pocock, R. I. (1930). "The Panthers and Ounces of Asia". Journal of the Bombay Natural History Society. 34 (2): 307–336.
  50. Deraniyagala, P. E. P. (1956). "The Ceylon leopard, a distinct subspecies". Spolia Zeylanica. 28: 115–116.
  51. Stein, A. B.; Athreya, V.; Gerngross, P.; Balme, G.; Henschel, P.; Karanth, U.; Miquelle, D.; Rostro, S.; Kamler, J. F. & Laguardia, A. (2016). "Panthera pardus". IUCN Red List of Threatened Species . 2016: e.T15954A102421779.
  52. 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.
  53. 1 2 3 Temminck, C. J. (1844). "Aperçu général et spécifique sur les Mammifères qui habitent le Japon et les Iles qui en dépendent". In Siebold, P. F. v.; Temminck, C. J.; Schlegel, H. (eds.). 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.
  54. Hilzheimer, M. (1905). "Über einige Tigerschädel aus der Straßburger zoologischen Sammlung". Zoologischer Anzeiger. 28: 594–599.
  55. 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. S2CID   84054536.
  56. 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): e442. doi:10.1371/journal.pbio.0020442. PMC   534810 . PMID   15583716.
  57. 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.
  58. Goodrich, J.; Lynam, A.; Miquelle, D.; Wibisono, H.; Kawanishi, K.; Pattanavibool, A.; Htun, S.; Tempa, T.; Karki, J.; Jhala, Y. & Karanth, U. (2015). "Panthera tigris". IUCN Red List of Threatened Species . 2015: e.T15955A50659951.
  59. Schreber, J. C. D. (1777). "Die Unze". Die Säugethiere in Abbildungen nach der Natur mit Beschreibungen. Erlangen: Wolfgang Walther. pp. 386–387.
  60. McCarthy, T.; Mallon, D.; Jackson, R.; Zahler, P. & McCarthy, K. (2017). "Panthera uncia". IUCN Red List of Threatened Species . 2017: e.T22732A50664030.
  61. 1 2 3 4 Mazák, J. H.; Christiansen, P.; Kitchener, A. 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.
  62. Pei, W. C. (1934). "On the Carnivora from Locality 1 of Choukoutien". Palaeontologica Sinica Series C, Fascicle 1: 1−166.
  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 .
  64. 1 2 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. doi:10.1111/j.1365-294X.2009.04134.x. PMID   19302360.
  65. 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. Bibcode:2006QuInt.142..218S. doi:10.1016/j.quaint.2005.03.019.CS1 maint: uses authors parameter (link)
  66. 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. S2CID   92328512.
  67. 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.
  68. 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 28 September 2015.
  69. Harington, C. R. (1996). Pleistocene mammals of the Yukon Territory (PhD). Edmonton: University of Alberta.
  70. Stuart, A. J., Lister, A .M. (2011). "Extinction chronology of the cave lion Panthera spelaea". Quaternary Science Reviews. 30 (17): 2329–2340. Bibcode:2011QSRv...30.2329S. doi:10.1016/j.quascirev.2010.04.023.CS1 maint: multiple names: authors list (link)
  71. Sala, B. (1990). "Panthera leo fossilis (v. Reichenau, 1906) (Felidae) de Iserna la Pineta (Pléistocene moyen inférieur d'Italie)". Géobios. 23 (2): 189–194. doi:10.1016/S0016-6995(06)80051-3.
  72. 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.
  73. 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. Bibcode:2014QuInt.339..245M. doi:10.1016/j.quaint.2013.10.008.
  74. 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". Integrative Zoology. 9 (4): 517–530. doi:10.1111/1749-4877.12082. PMID   24382145.
  75. 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: 4. doi: 10.5334/oq.24 .
  76. Manamendra-Arachchi, K., Pethiyagoda, R., Dissanayake, R., Meegaskumbura, M. (2005). "A second extinct big cat from the late Quaternary of Sri Lanka" (PDF). The Raffles Bulletin of Zoology (Supplement 12): 423–434.CS1 maint: multiple names: authors list (link)
  77. 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 . Retrieved 8 September 2015.
  78. Moreno, A.; Lima-Ribeiro, M. (2015). "Ecological niche models, fossil record and the multi-temporal calibration for Panthera onca (Linnaeus, 1758) (Mammalia: Felidae)" (PDF). Brazilian Journal of Biological Sciences. 2 (4): 309–319.
  79. Roth, S. (1899). "Descripción de los restos encontrados en la caverna de Última Esperanza". Revista del Museo la Plata. 9: 381–388.
  80. 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. doi:10.1186/s12862-018-1268-0. PMC   6198532 . PMID   30348080.
  81. 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: 167–193. Bibcode:2013QSRv...76..167D. doi:10.1016/j.quascirev.2013.05.009.
  82. 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.
  83. 1 2 3 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.
  84. Werdelin, L.; Yamaguchi, N.; Johnson, W. E. & O'Brien, S. J. (2010). "Phylogeny and evolution of cats (Felidae)". In Macdonald, D. W. & Loveridge, A. J. (eds.). Biology and Conservation of Wild Felids. Oxford: Oxford University Press. pp. 59–82. ISBN   978-0-19-923445-5.
  85. 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. Bibcode:2018PPP...496..166P. doi:10.1016/j.palaeo.2018.01.034.CS1 maint: multiple names: authors list (link)

Further reading