Panthera

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PantheraTemplate:Notetag 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 Template:Wikt-lang derives from Classical Latin Template:Wikt-lang, itself from the Ancient Greek Template:Translit (Template:Wikt-lang).<ref>Template:Cite book</ref>

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.<ref name=Pocock1939>Template:Cite book</ref> 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 Template:Cvt that provide a lower resistance to airflow; it was therefore proposed to be retained in the genus Uncia.<ref>Template:Cite journal</ref> 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.<ref>Template:Cite journal</ref>

Evolution

The geographic origin of the genus Panthera is uncertain, though the earliest known definitive species Panthera principialis is from Tanzania.<ref name=principialis/> P. blytheae from northern Central Asia, originally described as the oldest known Panthera species, is suggested to be similar in skull features to the snow leopard,<ref name=Tseng_al2014/> 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.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> The tiger, snow leopard, and clouded leopard genetic lineages likely dispersed in Southeast Asia during the Late Miocene.<ref name=Tseng_al2014>Template:Cite journal</ref> Genetic studies indicate that the pantherine cats diverged from the subfamily Felinae between six and ten million years ago.<ref name=Johnson2006>Template:Cite journal</ref> The genus Neofelis is sister to Panthera.<ref name=Johnson2006/><ref name=Janczewski>Template:Cite journal</ref><ref name=Johnson1997>Template:Cite journal</ref><ref name=Yu1>Template:Cite journal</ref> The clouded leopard appears to have diverged about Template:Ma. Panthera diverged from other cat species about Template:Ma and then evolved into the species tiger about Template:Ma, snow leopard about Template:Ma and leopard about Template:Ma. 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 Template:Ma, and that both P. atrox and P. spelaea are most closely related to lions among living Panthera species.<ref name="Barnett, 2009">Template:Cite journal</ref> The snow leopard is nested within Panthera and is the sister species of the tiger.<ref name=Davis2010>Template:Cite journal</ref>

Results of a 2016 study based on analysis of biparental nuclear genomes suggest the following relationships of living Panthera species:<ref name=Li_al2016>Template:Cite journal</ref>

The extinct species Panthera gombaszoegensis, was probably closely related to the modern jaguar. The first fossil remains were excavated in Olivola, in Italy, and date to Template:Ma.<ref>Template:Cite journal</ref> Fossil remains found in South Africa that appear to belong within the Panthera lineage date to about Template:Ma.<ref>Template:Cite journal</ref>

Classification

Panthera was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group.<ref>Template:Cite book</ref><ref name="Allen1902">Template:Cite journal</ref> 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.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> 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.<ref name="Pocock1916">Template:Cite journal</ref> 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.<ref name=OBrien1987>Template:Cite journal</ref><ref>Template:Cite journal</ref> Subsequently, it was proposed to group all African leopard populations to P. p. pardus and retain eight subspecific names for Asian leopard populations.<ref>Template:Cite journal</ref> 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.<ref name="Johnson2006" /><ref name="iucn1" />

Based on genetic research, it was suggested to group all living sub-Saharan lion populations into P. l. leo.<ref>Template:Cite journal</ref> 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.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>

Black panthers do not form a distinct species, but are melanistic specimens of the genus, most often encountered in the leopard and jaguar.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>

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:<ref name="MSW3"/><ref name=CatSG2017>Template:Cite journal</ref>

Species	Subspecies	IUCN Red List status and distribution
Lion P. leo Template:Small<ref name="Linn1758" />	P. l. leo Template:Small<ref name="Linn1758" /> including: †Barbary lion P. l. leo sensu stricto Asiatic lion Template:Small P. l. melanochaita Template:Small including: †Cape lion P. l. melanochaita sensu stricto<ref name="Mazak1975">Template:Cite journal</ref>	[[Vulnerable species|Template:IUCN status]]<ref>Template:Cite iucn</ref>
Jaguar P. onca Template:Small<ref name="Linn1758" />	Monotypic<ref>Template:Cite journal</ref><ref name="CatSG2017" />	[[Near-threatened species|Template:IUCN status]]<ref>Template:Cite iucn</ref>
Leopard P. pardus Template:Small<ref name="Linn1758" />	African leopard P. p. pardus Template:Small<ref name="Linn1758" /> Indian leopard P. p. fusca Template:Small<ref>Template:Cite book</ref> Javan leopard P. p. melas Template:Small<ref>Template:Cite journal</ref> Arabian leopard P. p. nimr Template:Small<ref>Template:Cite book</ref> P. p. tulliana Template:Small Amur leopard P. p. orientalis Template:Small Indochinese leopard P. p. delacouri Template:Small<ref>Template:Cite journal</ref> Sri Lankan leopard P. p. kotiya Template:Small<ref>Template:Cite journal</ref>	Template:IUCN status<ref>Template:Cite iucn</ref>
Tiger P. tigris Template:Small<ref name=Linn1758>Template:Cite book</ref> File:Sher Khan (cropped).jpg	P. t. tigris Template:Small including: Bengal tiger P. t. tigris sensu stricto †Caspian tiger Template:Small Siberian tiger Template:Small South China tiger Template:Small Indochinese tiger Template:Small Malayan tiger Template:Small Sunda Island tiger P. t. sondaica Template:Small including †Javan tiger P. t. sondaica sensu stricto Sumatran tiger Template:Small †Bali tiger Template:Small	[[Endangered species|Template:IUCN status]]<ref>Template:Cite iucn</ref>
Snow leopard P. uncia<ref name=CatSG2017/> Template:Small<ref>Template:Cite book</ref>	Monotypic<ref name=CatSG2017/>	Template:IUCN status<ref name=iucn1>Template:Cite iucn</ref>

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.<ref name = "Chimento2017">Template:Cite journal</ref>	Commonly known as the American lion, P. atrox is thought to have descended from a basal P. spelaea cave lion population isolated south of the Cordilleran Ice Sheet, and then established a mitochondrial sister clade circa 200,000 BP.<ref name=Barnett2009>Template:Cite journal</ref> It was sometimes considered a subspecies either under the nomenclature of P. leo<ref name=Barnett2009 /> or P. spelaea.<ref>Template:Cite journal</ref> One of the largest Panthera species.<ref name="Christiansen2009">Template:Cite journal</ref> Became extinct around 13,000-12,000 years ago.<ref>Template:Cite journal</ref>	File:American lion National Park Service illustration.png
Panthera balamoides<ref>Template:Cite journal</ref>	Mexico, ~0.13 MYA	Dubious, other authors suggest that the remains are actually of the extinct bear Arctotherium instead.<ref name=":3">Template:Cite journal</ref>
Panthera dhokpathanensis	Siwaliks, ~2.7 MYA	Described in 1986.<ref>Template:Cite journal</ref>
Panthera fossilis<ref>Template:Cite thesis</ref>	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.<ref name=":0">Template:Cite journal</ref>	File:W. Gornig - P. spelaea fossilis.png
Panthera gombaszoegensis	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.<ref name="Marciszak2014">Template:Cite journal</ref> 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. gombaszoegensis. It is occasionally classified as a subspecies of P. onca.<ref name="hemmeretal2010">Template:Cite journal</ref><ref>Template:Cite journal</ref>	File:De vroeg-pleistocene sabeltandkat, Homotherium crenatidens (2008) Panthera onca gombaszoegensis.png
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<ref name="Mazák2011">Template:Cite journal</ref> At least three recent studies considered Panthera zdanskyi likely to be a synonym of P. palaeosinensis.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>
Panthera principialis	Tanzania, ~3.7 MYA	Described in 2023.<ref name=principialis>Template:Cite journal</ref>
Panthera shawi	Laetoli site in Tanzania, ~3 MYA	A leopard-like cat<ref>Template:Cite journal</ref>
Panthera spelaea	Much of Eurasia, 0.6 to 0.013 MYA<ref>Template:Cite journal</ref>	Commonly known as the cave lion or steppe lion. Originally spelaea was classified as a subspecies of the extant lion P. leo.<ref name=Sala>Template:Cite journal</ref> 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.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> Other genetic results indicate that P. fossilis also warrants status as a species.<ref name=SotnikovaForonvoa2014>Template:Cite journal</ref><ref name=Barnett2016>Template:Cite journal</ref> It became extinct around 14,500-14,000 years ago.<ref>Template:Cite journal</ref>	File:W. Gornig - P. spelaea spelaea.png
Panthera youngi<ref>Template:Cite journal</ref>	China, Japan, ~0.35 MYA		File:Panthera youngi.webp
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.<ref name=Mazák2011/> But it is possibly synonymous with P. palaeosinensis.<ref name=principialis/><ref>Template:Cite journal</ref>	File:Longdan tiger.png
Panthera leo sinhaleyus	Sri Lanka	This lion subspecies was described on the basis of two teeth.<ref>Template:Cite journal</ref>
Panthera onca augusta<ref name=Columbia>Template:Cite journal</ref>	North America	May have lived in temperate forests across North America<ref name=":1">Template:Cite journal</ref>
Panthera onca mesembrina<ref>Template:Cite journal</ref>	South America	May have lived in grasslands in South America, unlike the modern jaguar	File:Panthera onca mesembrina JF.png
Panthera pardus spelaea	Europe	Closely related to Asiatic leopard subspecies,<ref>Template:Cite journal</ref>	File:Chauvet leopard rock art (white background).png
Panthera tigris acutidens	Much of Asia	Not closely related to modern tiger subspecies<ref name="PBDB">Template:Cite journal</ref>
Panthera tigris soloensis	Java, Indonesia	Not closely related to modern tiger subspecies<ref name="PBDB"/>
Panthera tigris trinilensis	Java, Indonesia	Not closely related to modern tiger subspecies<ref name="PBDB"/>

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.<ref>Template:Cite journal</ref>

Phylogeny

File:Two cladograms for Panthera.svg

Two cladograms proposed for Panthera. The upper one is based on phylogenetic studies by Johnson et al. (2006),<ref name=Johnson2006/> and by Werdelin et al. (2010).<ref name=Werdelin2010>Template:Cite book</ref> The lower cladogram is based on a study by Davis et al. (2010)<ref name=Davis2010/> and by Mazák et al. (2011).<ref name="Mazák2011"/>

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.<ref>Template:Cite journal</ref>

See also

• Panthera hybrid
• Panther (legendary creature)

Further reading

• Template:Cite book

External links

• Template:Commons category-inline
• Template:Cite AV mediaTemplate:Cbignore