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Panthera is a within the family , and one of two extant genera in the subfamily . It contains the largest living members of the cat family. There are five living species: the , , , and . Numerous extinct species are also named, including the and .


Etymology
The word derives from , itself from the ().


Characteristics
In Panthera species, the dorsal profile of the 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 axis is nearly horizontal. The inner chamber of the is large, the outer small. The partition between them is close to the external auditory meatus. The convexly rounded chin is sloping. All Panthera species have an incompletely ossified and a specially adapted with large covered in a fibro-elastic pad; these characteristics enable them to . Only the snow leopard cannot roar, as it has shorter vocal folds of that provide a lower resistance to airflow; it was therefore proposed to be retained in the genus Uncia. Panthera species can , 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.


Evolution
The geographic origin of the genus Panthera is uncertain, though the earliest known definitive species Panthera principialis is from . P. blytheae from northern , originally described as the oldest known Panthera species, is suggested to be similar in skull features to the snow leopard, 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 . The tiger, snow leopard, and likely dispersed in Southeast Asia during the . Genetic studies indicate that the diverged from the subfamily between six and ten million years ago. The genus is to Panthera. The clouded leopard appears to have diverged about . Panthera diverged from other cat species about and then evolved into the species tiger about , snow leopard about and leopard about . Mitochondrial sequence data from fossils suggest that the ( P. atrox) is a sister lineage to (the Eurasian cave or steppe lion) that diverged about , and that both P. atrox and P. spelaea are most closely related to lions among living Panthera species. The snow leopard is nested within Panthera and is the of the tiger.

Results of a 2016 study based on analysis of biparental suggest the following relationships of living Panthera species:

The extinct species Panthera gombaszogensis, was probably closely related to the modern jaguar. The first fossil remains were excavated in , in Italy, and date to . Fossil remains found in South Africa that appear to belong within the Panthera lineage date to about .


Classification
Panthera was named and described by in 1816 who placed all the spotted cats in this group. 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. Reginald Innes Pocock revised the classification of this genus in 1916 as comprising the ( P. tigris), ( P. leo), ( P. onca), and ( P. pardus) on the basis of common features of their . Since the mid-1980s, several Panthera species became subjects of , 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. Subsequently, it was proposed to group all African leopard populations to P. p. pardus and retain eight for Asian leopard populations. Results of indicate that the (formerly Uncia uncia) also belongs to the genus Panthera ( P. uncia), a classification that was accepted by IUCN Red List assessors in 2008.

Based on , it was suggested to group all living lion populations into P. l. leo. Results of studies indicate that the and lion populations are more closely related to those in India and form a different than lion populations in and ; southeastern Ethiopia is an admixture region between North African and East African lion populations.

do not form a distinct species, but are specimens of the genus, most often encountered in the leopard and jaguar.


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:
P. leo P. l. leo including: P. l. melanochaita including:
  • P. l. melanochaita sensu stricto
P. onca Monotypic
P. pardus P. p. pardus
P. p. fusca
P. p. melas
P. p. nimr
P. p. tulliana
P. p. orientalis
Indochinese leopard P. p. delacouri
Sri Lankan leopard P. p. kotiya
P. tigris P. t. tigris including: Sunda Island tiger P. t. sondaica including
P. uncia


Extinct species and subspecies
North America, 0.13 to 0.013 MYA, with dubious remains in South America.Commonly known as the American lion , P. atrox is thought to have descended from a basal P. spelaea cave lion population isolated south of the Cordilleran Ice Sheet, and then established a mitochondrial circa 200,000 BP. It was sometimes considered a subspecies either under the nomenclature of P. leo or P. spelaea. One of the largest Panthera species. Became extinct around 13,000-12,000 years ago.
Panthera balamoidesMexico, ~0.13 MYADubious, other authors suggest that the remains are actually of the extinct bear instead.
Panthera fossilisEurope and Asia, 0.68 to 0.25 MYAExtinct 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.
Panthera gombaszogensisEurope, possibly Asia and Africa, 2.0 to 0.35 MYARanged across Europe, as well as possibly Asia and Africa from around 2 million to 350,000 years ago. 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.
Panthera palaeosinensisNorthern China, ~3 MYAInitially thought to be an ancestral tiger species, but several scientists place it close to the base of the genus Panthera At least three recent studies considered Panthera zdanskyi likely to be a synonym of P. palaeosinensis.
Panthera principialisTanzania, ~3.7 MYADescribed in 2023.
Laetoli site in Tanzania, ~3 MYAA leopard-like cat
Much of Eurasia, 0.6 to 0.013 MYACommonly known as the cave lion or steppe lion. Originally spelaea was classified as a subspecies of the extant lion P. leo. Results of recent genetic studies indicate that it belongs to a distinct species, namely P. spelaea that is most closely related to the modern lion among living Panthera species. Other genetic results indicate that P. fossilis also warrants status as a species. It became extinct around 14,500-14,000 years ago.
China, Japan, ~0.35 MYA
Panthera zdanskyi of northwestern China, 2.55 to 2.16 MYAIt was initially considered to be a close relative of the tiger. But it is possibly synonymous with P. palaeosinensis.
Panthera leo sinhaleyusSri LankaThis lion subspecies was described on the basis of two teeth.
Panthera onca augustaNorth AmericaMay have lived in temperate forests across North America
Panthera onca mesembrinaSouth AmericaMay have lived in grasslands in South America, unlike the modern jaguar
Panthera pardus spelaeaEuropeClosely related to Asiatic leopard subspecies,
Panthera tigris acutidensMuch of AsiaNot closely related to modern tiger subspecies
Panthera tigris soloensisJava, IndonesiaNot closely related to modern tiger subspecies
Panthera tigris trinilensisJava, IndonesiaNot closely related to modern tiger subspecies
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. A " Panthera dhokpathanensis" was briefly referenced in 1986 in a report on apparent new carnivorans from the Dhok Patha region in the , but as no description was provided this name is a .


Phylogeny
2018, results of a study on living and cats were published. This study was based on the morphological diversity of the of saber-toothed cats, their and .


See also


Notes

Further reading


External links
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