Panthera

Panthera[1]
Temporal range: Late Miocene – present, 5.95–0 Ma
Tiger (Panthera tigris), the largest species of the genus Panthera
Radial bone of Panthera fossilis
Scientific classification
Kingdom:Animalia
Phylum:Chordata
Class:Mammalia
Order:Carnivora
Suborder:Feliformia
Family:Felidae
Subfamily:Pantherinae
Genus:Panthera
Oken, 1816
Type species
Felis pardus
Extant species

Panthera tigris
Panthera uncia
Panthera onca
Panthera leo
Panthera pardus

Panthera is a genus within the Felidae family that was named and first described by the German naturalist Oken in 1816.[2] The British taxonomist Pocock revised the classification of this genus in 1916 as comprising the species lion, tiger, jaguar, and leopard on the basis of cranial features.[3] Results of genetic analysis indicate that the snow leopard also belongs to the Panthera, a classification that was accepted by IUCN assessors in 2008.[4][5]

Only the tiger, lion, leopard, and jaguar have 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 it has an incomplete ossification of the hyoid bone, it lacks the special morphology of the larynx.[6]

Etymology

The word panther derives from classical Latin panthēra, itself from the ancient Greek pánthēr (πάνθηρ).[7] The word originated in antiquity in the Orient, probably from India to Persia to Greece.[8] A common folk etymology derives the word from Greek pan- (πάν), meaning "all", and thēr (θήρ), meaning "beast of prey".

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.[9] 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.[10]

Evolution

Panthera probably evolved in Asia, but the roots of the genus remain unclear. Genetic studies indicate that pantherine cats diverged from the subfamily Felinae between six and ten million years ago.[4] Fossil records that appear to belong within the genus Panthera reach only 2.0 to 3.8 million years back.[11]

The snow leopard was initially seen at the base of Panthera, but newer molecular studies suggest that it is nestled within Panthera and is a sister species of the tiger.[12] 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.[4][13][14][15] Since 2008, the IUCN Red List lists it as Panthera uncia using Uncia uncia as a synonym.[5]

The genus Neofelis is generally placed at the base of the Panthera group, but is not included in the genus itself.[4][14][15][16]

Results of a mitogenomic study suggest the phylogeny can be represented as Neofelis nebulosa (Panthera tigris (Panthera onca (Panthera pardus, (Panthera leo, Panthera uncia)))).[17] About 11.3 million years ago Panthera separated from other felid species and then evolved into the several species of the genus. N. nebulosa appears to have diverged about 8.66 million years ago, P. tigris about 6.55 million years ago, P. uncia about 4.63 million years ago and P. pardus about 4.35 million years ago. Mitochondrial sequence data from fossils suggest that the American lion (P. l. atrox) is a sister lineage to Upper Pleistocene Eurasian cave lion (P. l. spelaea) that diverged about 0.34 million years ago.[18]

The prehistoric cat Panthera onca gombaszogensis, often called European jaguar is probably closely related to the modern jaguar. The earliest evidence of the species was obtained at Olivola in Italy, and dates 1.6 million years.[19]

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.[20][21][22] 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.[23][24] Subsequently, it was proposed to group all African leopard populations to P. p. pardus and retain eight subspecific names for Asian leopard populations.[25]

Based on genetic research, it was suggested to group all living sub-Saharan lion populations into P. l. leo.[26] 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.[27][28]

Black panthers do not form a distinct species, but are melanistic specimens of the genus, most often encountered in the leopard and jaguar.[29][30]

Phylogeny

Two cladograms proposed for Panthera. The upper one is based on phylogenetic studies by Johnson et al. (2006),[4] and by Werdelin et al. (2010).[31] The lower cladogram is based on a study by Davis et al. (2010)[32] and by Mazák et al. (2011).[33]

The cladogram below follows Mazák, Christiansen and Kitchener (2011).[33]

Pantherinae

 

Neofelis

 
Panthera

 

Panthera uncia

 
 

 

†Panthera palaeosinensis

 
 

 

 

Panthera onca

 
 

 

†Panthera atrox

 
 

 

†Panthera spelaea

 
 

 

Panthera leo

 
 

Panthera pardus

 
 
 
 
 
 

 

Panthera tigris

 
 

†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:[34]

Panthera

 

†Panthera palaeosinensis

 
 

 

 

 

†Panthera blytheae

 
 

Panthera uncia

 
 
 
 

 

 

†Panthera zdanskyi

 
 

Panthera tigris

 
 
 
 
 

 

 

 

†Panthera gombaszoegensis

 
 

Panthera onca

 
 
 
 

 

 

Panthera pardus

 
 

 

Panthera leo

 
 

 

†Panthera spelaea

 
 

†Panthera atrox

 
 
 
 
 
 
 

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][35]

SpeciesSubspeciesCurrent distribution
Tiger P. tigris

Tigers of mainland Asia P. t. tigris including:[35]

Sunda Island tiger P. t. sondaica including[35]

 India, Nepal, Bangladesh, Bhutan, Myanmar, Thailand, Cambodia, eastern Russia and China, Indonesian island of Sumatra
Lion P. leo

P. l. leo including:[35]

P. l. melanochaita including:[35]

 India, West, Central, East and Southern Africa
Jaguar P. onca

Monotypic[42][35] Historic range stretches across the USA and Mexico to Chile and Argentina, including much of Amazonian Brazil.[43] Countries present in the modern range are Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica (particularly on the Osa Peninsula), Ecuador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, Paraguay, Peru, Suriname, the United States and Venezuela.
Leopard P. pardus

African leopard P. p. pardus

Arabian leopard P. p. nimr
Javan leopard P. p. melas
Indian leopard P. p. fusca
Sri Lankan leopard P. p. kotiya
Persian leopard and Anatolian leopard P. p. tulliana,[35] syn. P. p. ciscaucasica, P. p. saxicolor[1]
Indochinese leopard P. p. delacouri
Amur leopard P. p. orientalis, syn. P. p. japonensis[35]

 Africa, Middle East, South and Southeast Asia, East Asia
Snow leopard P. uncia[35]

Monotypic Kazakhstan, Kyrgyzstan, Tajikistan, and Uzbekistan to the Hindu Kush in eastern Afghanistan, Karakoram in northern Pakistan, in the Pamir Mountains, and in the high altitudes of the Himalayas in India, Nepal, and Bhutan, and the Tibetan Plateau.

Fossil species and subspecies

SpeciesSubspeciesFossil distributionNotes
†Panthera palaeosinensis Monotypic Northern China Originally thought to be an ancestral species of tiger, this species has been found in recent studies place it close to the base of the genus Panthera.[33][44]
†Panthera blytheae Monotypic Tibetan Plateau The oldest known species of Panthera closely related to the snow leopard.[45]
†Panthera zdanskyi Monotypic Gansu province of northwestern China Closely related to the tiger.[46]
Tiger P. tigris †P. t. acutidens[47]

†P. t. trinilensis[48]
†P. t. soloensis[49]

 Much of Asia (P. t. acutidens); Java, Indonesia (P. t. trinilensis and P. t. soloensis) These three subspecies are not closely related to the modern subspecies of tigers.[50]
†Panthera toscana

†P. t. toscana

†P. t. georgica[51]

 Italy (P. t. toscana) and Georgia (P. t. georgica) The oldest fossil species of jaguar (sometimes classified as fossil subspecies of the extant animal or P. gombaszoegensis).[51]
†Panthera gombaszoegensis

†P. g. schreuderi

†P. g. gombaszoegensis

 Europe P. g. schreuderi might be a junior synonym of P. g. gombaszoegensis.[52][51][53] Sometimes classified as subspecies of the P. onca.[51][53]
†Giant jaguars Panthera onca

†P. o. augusta[54]

†P. o. mesembrina[55]

 North America (P. o. augusta) and South America (P. o. mesembrina) P. onca represented the third wave of jaguar evolution,[51] with P. o. augusta in the temperate forests of North America, P. o. mesembrina in temperate grasslands of South America, and the smaller modern jaguar, P. o. onca, inhabiting the tropics of both continents during the Pleistocene.[43]
†European Panthera pardus spp. †P. p. spelaea[56]

†P. p. begoueni[57]
†P. p. sickenbergi[58]
†P. p. antiqua[59]
†P. p. begoueni[57]

 Europe Closely related to Persian leopard P. p. tulliana according to genetic work.[60]
†Panthera crassidens Monotypic South Africa A possible chimera of fossil leopards and cheetahs.[61]
†Panthera shawi Monotypic Laetoli site in Tanzania A leopard-like cat which is believed to be the oldest known species of lion.[62]
Lion P. leo †P. l. sinhaleyus Sri Lanka This lion subspecies is only known by two teeth.[63]
†Panthera spelaea †Panthera leo fossilis[64]

†Panthera leo spelaea

 Iberian peninsula, Southeast Europe, Great Britain, Central Europe, East European Plain, across most of northern Eurasia into Canada and Alaska[65] Originally spelaea and fossilis were classified as subspecies of the extant lion P. leo.[66] Results of recent genetic studies indicate that both belong to a distinct species, namely P. spelaea.[67][68] Other genetic results indicate that the fossilis cave lion warrants status of a species.[69][70]
†Panthera atrox

Monotypic North America, dubious remains in South America.[71] The American lion is thought to have descended from a basal spelaea cave lion population isolated south of the North American continental ice sheet, and then established a mitochondrial sister clade circa 200,000 BP.[70][72] It is sometimes considered either under the nomenclature of P. leo[72] or of P. spelaea.[73]
†Panthera youngi[74] Monotypic China, Japan

See also

References

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Further reading

  • Turner, A. and Antón, M. (1997). The Big Cats and Their Fossil Relatives: An Illustrated Guide to Their Evolution and Natural History. Columbia University Press. ISBN 978-0-231-10228-5.

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