Panthera

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]

Not to be confused with Pantera.

Panthera[1]
Temporal range: Late Miocene – present, 5.95–0 Ma
From top to bottom: Tiger, Lion, Jaguar, Leopard, Snow Leopard
Scientific classification
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

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]

Etymology

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]

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 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]

Evolution

The geographic origin of the Panthera is most likely northern Central Asia. Panthera blytheae, the oldest known Panthera species, exhibits similar skull features as 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 generally placed at the base of the Panthera group, but is not included in the genus itself.[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 a sister species of the tiger.[18] 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.[5][19][14][15] Results of a mitogenomic study suggest the phylogeny can be represented as Neofelis nebulosa (Panthera tigris (Panthera onca (Panthera pardus, (Panthera leo, Panthera uncia)))).[20]

The prehistoric Panthera onca gombaszogensis, often called the European jaguar, is probably closely related to the modern jaguar. The first fossil remains were excavated in Olivola in Italy and date 1.6 million years.[21] Fossil remains found in South Africa that appear to belong within the Panthera are about 2.0 to 3.8 million years old.[22]

Classification

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.[23][24][25] 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.[26][27] Subsequently, it was proposed to group all African leopard populations to P. p. pardus and retain eight subspecific names for Asian leopard populations.[28]

Based on genetic research, it was suggested to group all living sub-Saharan lion populations into P. l. leo.[29] 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.[30][31]

Black panthers do not form a distinct species, but are melanistic specimens of the genus, most often encountered in the leopard and jaguar.[32][33]

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][34]

SpeciesSubspeciesIUCN Red List status and distribution
Tiger P. tigris (Linnaeus, 1758)[35]

Tigers of mainland Asia P. t. tigris (Linnaeus, 1758) including:
  • Bengal tiger P. t. tigris sensu stricto
  • †Caspian tiger syn. P. t. virgata (Illiger, 1815)[36]
  • Siberian tiger syn. P. t. altaica (Temminck, 1844)[37]
  • South China tiger syn. P. t. amoyensis (Hilzheimer, 1905)[38]
  • Indochinese tiger syn. P. t. corbetti Mazák, 1968[39]
  • Malayan tiger syn. P. t. jacksoni Luo et al., 2004[40]

Sunda Island tiger P. t. sondaica Temminck, 1844)[37] including

  • †Javan tiger P. t. sondaica sensu stricto
  • Sumatran tiger syn. P t. sumatrae (Temminck, 1844)[37]
  • †Bali tiger syn. P. t. balica Schwarz, 1912)[41]
 EN[42]

Lion P. leo (Linnaeus, 1758)[35]

P. l. leo (Linnaeus, 1758)[35] including:

P. l. melanochaita (Smith, 1842)[44] including:

  • †Cape lion P. l. melanochaita sensu stricto[45]
 VU[46]

Jaguar P. onca (Linnaeus, 1758)[35]

Monotypic[47][34] NT[48]

Leopard P. pardus (Linnaeus, 1758)[35]

African leopard P. p. pardus (Linnaeus, 1758)[35]

Indian leopard P. p. fusca (Meyer, 1794)[49]
Javan leopard P. p. melas (G. Cuvier, 1809)[50]
Arabian leopard P. p. nimr (Hemprich and Ehrenberg), 1833[51]
Anatolian leopard or Persian leopard P. p. tulliana (Valenciennes, 1856),[52] syn. P. p. ciscaucasica (Satunin, 1914),[53] P. p. saxicolor Pocock, 1927[54]
Amur leopard P. p. orientalis (Schlegel, 1857),[55] syn. P. p. japonensis (Gray, 1862)[56]
Indochinese leopard P. p. delacouri Pocock, 1930[57]
Sri Lankan leopard P. p. kotiya Deraniyagala, 1956[58]

 VU[59]

Snow leopard P. uncia[34] (Schreber, 1775)[60]

Monotypic[34] VU[61]

Extinct species and subspecies
SpeciesFossil recordsNotes
Panthera blytheae Tibetan Plateau One of the oldest known Panthera species, possibly closely related to the snow leopard.[13]
Panthera palaeosinensis Northern China It was initially thought to be an ancestral tiger species, but several scientists place it close to the base of the genus Panthera.[62]
Panthera zdanskyi Gansu province of northwestern China Possibly a close relative of the tiger.[62]
Panthera youngi[63] China, Japan
Panthera atrox North America, dubious remains in South America.[64] 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.[65] It was sometimes considered a subspecies either under the nomenclature of P. leo[65] or P. spelaea.[66]
Panthera balamoides[67] Mexico
Panthera crassidens South Africa No longer a valid species due to being described based on a mixture of leopard and cheetah fossils.
Panthera gombaszoegensis Europe Panthera schreuderi and Panthera toscana are considered junior synonyms of P. gombaszoegensis. It is occasionally classified as a subspecies of P. onca.[68][69]
Panthera leo fossilis[70] Europe
Panthera spelaea Much of Eurasia[71] Originally spelaea was classified as a subspecies of the extant lion P. leo.[72] Results of recent genetic studies indicate that it belongs to a distinct species, namely P. spelaea.[73][74] Other genetic results indicate that the fossilis cave lion also warrants status as a species.[75][76]
Panthera leo sinhaleyus Sri Lanka This lion subspecies was described on the basis of two teeth.[77]
Panthera onca augusta[78] North America May have lived in temperate forests across North America.[79]
Panthera onca mesembrina[80] South America May have lived in grasslands in South America, unlike the modern jaguar.
Panthera pardus spelaea Europe Closely related to Asiatic leopard subspecies,[81] with at least one study suggesting that it is closely related to the Persian leopard (P. p. tulliana) according to genetic work.[82]
Panthera shawi Laetoli site in Tanzania A leopard-like cat.[83]
Panthera tigris acutidens Much of Asia Not closely related to modern tiger subspecies.[84]
Panthera tigris soloensis Java, Indonesia Not closely related to modern tiger subspecies.[84]
Panthera tigris trinilensis Java, Indonesia Not closely related to modern tiger subspecies.[84]

Phylogeny
Two cladograms proposed for Panthera. The upper one is based on phylogenetic studies by Johnson et al. (2006),[5] and by Werdelin et al. (2010).[85] The lower cladogram is based on a study by Davis et al. (2010)[18] and by Mazák et al. (2011).[62]

The cladogram below follows Mazák, Christiansen and Kitchener (2011).[62]

Pantherinae
 

Neofelis

 
Panthera
 

Snow leopard

 
 
 

Panthera palaeosinensis

 
 
 
 

Jaguar

 
 
 

American lion

 
 
 

Panthera spelaea

 
 
 

Lion

 
 

Leopard

 
 
 
 
 
 
 

Tiger

 
 

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:[86]

Panthera
 

Panthera palaeosinensis

 
 
 
 
 

Panthera blytheae

 
 

Snow leopard

 
 
 
 
 
 

Panthera zdanskyi

 
 

Tiger

 
 
 
 
 
 
 
 

European jaguar

 
 

Jaguar

 
 
 
 
 
 

Leopard

 
 
 

Lion

 
 
 

Panthera spelaea

 
 

American lion

 
 
 
 
 
 
 

See also
  • Pantherinae
  • Panthera hybrid
  • Panther (legendary creature)

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