# Panthera

Panthera is a genus within the family Felidae, and one of two extant genera in the subfamily Pantherinae. It contains the largest living members of the cat family. There are five living species: the jaguar, leopard, lion, snow leopard and tiger. Numerous extinct species are also named, including the cave lion and American lion.

## Etymology

The word panther derives from classical Latin panthēra, itself from the ancient Greek pánthēr (πάνθηρ).

## 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. All Panthera species have an incompletely ossified hyoid bone and a specially adapted larynx with large vocal folds covered in a fibro-elastic pad; these characteristics enable them to roar. Only the snow leopard cannot roar, as it has shorter vocal folds of 9 mm (0.35 in) that provide a lower resistance to airflow; it was therefore proposed to be retained in the genus Uncia. 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.

## Evolution

The geographic origin of the genus Panthera is uncertain, though the earliest known definitive species Panthera principialis is from Tanzania. 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, 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. The tiger, snow leopard, and clouded leopard genetic lineages likely dispersed in Southeast Asia during the Late Miocene. Genetic studies indicate that the pantherine cats diverged from the subfamily Felinae between six and ten million years ago. The genus Neofelis is sister to Panthera. The clouded leopard appears to have diverged about 8.66 million years ago. Panthera diverged from other cat species about 11.3 million years ago and then evolved into the species tiger about 6.55 million years ago, snow leopard about 4.63 million years ago and leopard about 4.35 million years ago. Mitochondrial sequence data from fossils suggest that the American lion (P. atrox) is a sister lineage to Panthera spelaea (the Eurasian cave or steppe lion) that diverged about 0.34 million years ago, and that both P. atrox and P. spelaea are most closely related to lions among living Panthera species. The snow leopard is nested within Panthera and is the sister species of the tiger.
Results of a 2016 study based on analysis of biparental nuclear genomes suggest the following relationships of living Panthera species:
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 1.6 million years ago. Fossil remains found in South Africa that appear to belong within the Panthera lineage date to about 2 to 3.8 million years ago.

## Classification

Panthera was named and described by Lorenz Oken 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 tiger (P. tigris), lion (P. leo), jaguar (P. onca), and leopard (P. pardus) on the basis of common features of their skulls. 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. Subsequently, it was proposed to group all African leopard populations to P. p. pardus and retain eight subspecific names for Asian leopard populations. 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.
Based on genetic research, it was suggested to group all living sub-Saharan lion populations into P. l. leo. 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.
Black panthers do not form a distinct species, but are melanistic 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:
| Species | Subspecies | IUCN Red List status and distribution |
| Lion P. leo (Linnaeus, 1758) | P. l. leo (Linnaeus, 1758) including: †Barbary lion P. l. leo sensu stricto Asiatic lion syn. P. l. persica (Johann Nepomuk Meyer, 1826) P. l. melanochaita (Smith, 1842) including: †Cape lion P. l. melanochaita sensu stricto | VU |
| Jaguar P. onca (Linnaeus, 1758) | Monotypic | NT |
| Leopard P. pardus (Linnaeus, 1758) | African leopard P. p. pardus (Linnaeus, 1758) Indian leopard P. p. fusca (Meyer, 1794) Javan leopard P. p. melas (G. Cuvier, 1809) Arabian leopard P. p. nimr (Hemprich and Ehrenberg), 1833 P. p. tulliana (Valenciennes, 1856), syn. P. p. ciscaucasica (Satunin, 1914), P. p. saxicolor Pocock, 1927 Amur leopard P. p. orientalis (Schlegel, 1857), syn. P. p. japonensis (Gray, 1862) Indochinese leopard P. p. delacouri Pocock, 1930 Sri Lankan leopard P. p. kotiya Deraniyagala, 1956 | VU |
| Tiger P. tigris (Linnaeus, 1758) | P. t. tigris (Linnaeus, 1758) including: Bengal tiger P. t. tigris sensu stricto †Caspian tiger syn. P. t. virgata (Illiger, 1815) Siberian tiger syn. P. t. altaica (Temminck, 1844) South China tiger syn. P. t. amoyensis (Hilzheimer, 1905) Indochinese tiger syn. P. t. corbetti Mazák, 1968 Malayan tiger syn. P. t. jacksoni Luo et al., 2004 Sunda Island tiger P. t. sondaica Temminck, 1844) including †Javan tiger P. t. sondaica sensu stricto Sumatran tiger syn. P t. sumatrae (Temminck, 1844) †Bali tiger syn. P. t. balica Schwarz, 1912) | EN |
| Snow leopard P. uncia (Schreber, 1775) | Monotypic | VU |

### Extinct species and subspecies

| Species and subspecies | Fossil records | Notes |
| Panthera atrox | 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 sister clade 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 balamoides | Mexico, ~0.13 MYA | Dubious, other authors suggest that the remains are actually of the extinct bear Arctotherium instead. |
| Panthera dhokpathanensis | Siwaliks, ~2.7 MYA | Described in 1986. |
| Panthera fossilis | 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. |
| 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. 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. |
| 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 At least three recent studies considered Panthera zdanskyi likely to be a synonym of P. palaeosinensis. |
| Panthera principialis | Tanzania, ~3.7 MYA | Described in 2023. |
| Panthera shawi | Laetoli site in Tanzania, ~3 MYA | A leopard-like cat |
| Panthera spelaea | Much of Eurasia, 0.6 to 0.013 MYA | Commonly 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. |
| Panthera youngi | China, Japan, ~0.35 MYA | |
| Panthera zdanskyi | Gansu province of northwestern China, 2.55 to 2.16 MYA | It was initially considered to be a close relative of the tiger. But it is possibly synonymous with P. palaeosinensis. |
| Panthera leo sinhaleyus | Sri Lanka | This lion subspecies was described on the basis of two teeth. |
| Panthera onca augusta | North America | May have lived in temperate forests across North America |
| Panthera onca mesembrina | South America | May have lived in grasslands in South America, unlike the modern jaguar |
| Panthera pardus spelaea | Europe | Closely related to Asiatic leopard subspecies, |
| Panthera tigris acutidens | Much of Asia | Not closely related to modern tiger subspecies |
| Panthera tigris soloensis | Java, Indonesia | Not closely related to modern tiger subspecies |
| Panthera tigris trinilensis | Java, Indonesia | Not 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.

### Phylogeny

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.