Panthera

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Panthera
Temporal range: 5.95–0  Ma
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Late Miocene – Present
4panthera3.0.png
From top to bottom: tiger, lion, jaguar, leopard, snow leopard
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Oken, 1816 [1]
Type species
Felis pardus
(= Panthera pardus)
Species
Synonyms [1]
About 10
  • Jaguarius Severtzov, 1858
  • LeoFrisch, 1775
  • LeoninaGreve, 1894
  • Leoninae Wagner, 1841
  • PardotigrisKretzoi, 1929
  • Pardus Fitzinger, 1868
  • TigrinaGreve, 1894
  • TigrinaeWagner, 1841
  • Tigris Gray, 1843
  • TigrisFrisch, 1775

Panthera is a genus within the family Felidae, it is one of two extant genera in the subfamily Pantherinae, and contains the largest living members of the cat family. There are 5 living species, the tiger, jaguar, lion, leopard and snow leopard and a number of extinct species.

Contents

Etymology

The word panther derives from classical Latin panthēra, itself from the ancient Greek pánthēr (πάνθηρ). [2]

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. [3] 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. [4] 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. [5]

Evolution

The geographic origin of the Panthera is most likely northern Central Asia. Panthera blytheae , the oldest known Panthera species, is similar in skull features to the snow leopard. The tiger, snow leopard, and clouded leopard genetic lineages dispersed in Southeast Asia during the Miocene. [6] Genetic studies indicate that the pantherine cats diverged from the subfamily Felinae between six and ten million years ago. [7] The genus Neofelis is sister to Panthera. [7] [8] [9] [10] 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. [11] The snow leopard is nested within Panthera and is the sister species of the tiger. [12]

Results of a 2016 study based on analysis of biparental nuclear genomes suggest the following relationships of living Panthera species: [13]

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. [14] Fossil remains found in South Africa that appear to belong within the Panthera lineage date to about 2 to 3.8 million years ago. [15]

Classification

Panthera was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group. [16] [17] 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. [18] [19] [20] 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. [21] 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. [22] [23] Subsequently, it was proposed to group all African leopard populations to P. p. pardus and retain eight subspecific names for Asian leopard populations. [24] 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. [7] [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]

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] [31]

SpeciesSubspecies IUCN Red List status and distribution
Lion P. leo(Linnaeus, 1758) [32]

Ngorongoro Crater (26) (cropped).jpg

P. l. leo (Linnaeus, 1758) [32] including:

P. l. melanochaita (Smith, 1842) [34] including:

 VU [36]

Lion distribution.png

Jaguar P. onca(Linnaeus, 1758) [32]

Jagwar (cropped).jpg

Monotypic [37] [31] NT [38]

Panthera onca distribution.svg

Leopard P. pardus(Linnaeus, 1758) [32]

Indian male leopard (cropped).jpg

African leopard P. p. pardus(Linnaeus, 1758) [32]

Indian leopard P. p. fusca(Meyer, 1794) [39]
Javan leopard P. p. melas(G. Cuvier, 1809) [40]
Arabian leopard P. p. nimr(Hemprich and Ehrenberg), 1833 [41]
P. p. tulliana (Valenciennes, 1856), [42] syn. P. p. ciscaucasica (Satunin, 1914), [43] P. p. saxicolor Pocock, 1927 [44]
Amur leopard P. p. orientalis(Schlegel, 1857), [45] syn. P. p. japonensis (Gray, 1862) [46]
Indochinese leopard P. p. delacouriPocock, 1930 [47]
Sri Lankan leopard P. p. kotiyaDeraniyagala, 1956 [48]

VU [49]

Leopard distribution.jpg

Tiger P. tigris(Linnaeus, 1758) [32]

Sher Khan (cropped).jpg

P. t. tigris (Linnaeus, 1758) including:

Sunda Island tiger P. t. sondaicaTemminck, 1844) [51] including

 EN [56]

Tiger map.svg

Snow leopard P. uncia [31] (Schreber, 1775) [57]

Snow Leopards - Ghost of the Mountains for Ministry of Tourism (MOT) India 9 (cropped).png

Monotypic [31] VU [25]

SnowLeopard distribution.jpg

Extinct species and subspecies

Species and subspecies Fossil recordsNotesImages
Panthera blytheae Tibetan PlateauOne of the oldest known Panthera species, possibly closely related to the snow leopard [6]
Panthera palaeosinensis Northern ChinaInitially thought to be an ancestral tiger species, but several scientists place it close to the base of the genus Panthera [58]
Panthera zdanskyi Gansu province of northwestern ChinaPossibly a close relative of the tiger [58]
Longdan tiger.png
Panthera youngi [59] China, Japan
Panthera atrox North America, dubious remains in South America. [60] 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. [61] It was sometimes considered a subspecies either under the nomenclature of P. leo [61] or P. spelaea. [62] One of the largest Panthera species. [63] Became extinct around 13,000-12,000 years ago. [64]
Panthera leo atrox Page.jpg
Panthera balamoides [65] MexicoDubious, other authors suggest that the remains are actually of the extinct bear Arctotherium instead. [66]
Panthera gombaszoegensis EuropeRanged across Europe from around 2 million to 350,000 years ago. [67] 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 fossilis [70] EuropeLikely the ancestor of P.spelaea [71]
Panthera spelaea Much of Eurasia [72] Commonly known as the cave lion or steppe lion. Originally spelaea was classified as a subspecies of the extant lion P. leo. [73] 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. [74] [75] Other genetic results indicate that P. fossilis also warrants status as a species. [76] [77] Became extinct around 14,500-14,000 years ago. [78]
Panthera leo spelaea in Vienna.jpg
Panthera leo sinhaleyus Sri LankaThis lion subspecies was described on the basis of two teeth. [79]
Panthera onca augusta [80] North AmericaMay have lived in temperate forests across North America [81]
Panthera onca mesembrina [82] South AmericaMay have lived in grasslands in South America, unlike the modern jaguar
Panthera pardus spelaea EuropeClosely related to Asiatic leopard subspecies, [83]
Panthera shawi Laetoli site in TanzaniaA leopard-like cat [84]
Panthera tigris acutidens Much of AsiaNot closely related to modern tiger subspecies [85]
Panthera tigris soloensis Java, IndonesiaNot closely related to modern tiger subspecies [85]
Panthera tigris trinilensis Java, IndonesiaNot closely related to modern tiger subspecies [85]
Panthera principialis TanzaniaDescribed in 2022. [86]
Panthera dhokpathanensis SiwaliksDescribed in 1986. [87]

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. [88]

Phylogeny

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

The cladogram below follows Mazák, Christiansen and Kitchener (2011). [58]

Pantherinae

Neofelis Studienblatt Felis macroscelis Nebelparder (white background).jpg

Panthera

Snow leopard Stamp-russia2014-save-russian-cats-(snow leopard).png

†Panthera palaeosinensis

Jaguar Felis onca - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg

†American lion

†Panthera spelaea W. Gornig - P. spelaea spelaea.png

Lion Felis leo - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(White Background).jpg

Leopard Felis pardus - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg

Tiger Stamp-russia2014-save-russian-cats-(tiger).png

†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: [90]

Panthera

†Panthera palaeosinensis

†Panthera blytheae

Snow leopard Stamp-russia2014-save-russian-cats-(snow leopard).png

†Panthera zdanskyi

Tiger Stamp-russia2014-save-russian-cats-(tiger).png

†European jaguar

Jaguar Felis onca - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg

Leopard Felis pardus - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg

Lion Felis leo - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(White Background).jpg

†Panthera spelaea W. Gornig - P. spelaea spelaea.png

†American lion

See also

Related Research Articles

<span class="mw-page-title-main">Felidae</span> Family of mammals

Felidae is the family of mammals in the order Carnivora colloquially referred to as cats. A member of this family is also called a felid. The term "cat" refers both to felids in general and specifically to the domestic cat.

<span class="mw-page-title-main">Jaguar</span> Large cat native to the Americas

The jaguar is a large cat species and the only living member of the genus Panthera native to the Americas. With a body length of up to 1.85 m and a weight of up to 158 kg (348 lb), it is the biggest cat species in the Americas and the third largest in the world. Its distinctively marked coat features pale yellow to tan colored fur covered by spots that transition to rosettes on the sides, although a melanistic black coat appears in some individuals. The jaguar's powerful bite allows it to pierce the carapaces of turtles and tortoises, and to employ an unusual killing method: it bites directly through the skull of mammalian prey between the ears to deliver a fatal blow to the brain.

<span class="mw-page-title-main">Lion</span> Large cat native to Africa and Asia

The lion is a large cat of the genus Panthera, native to Africa and India. It has a muscular, broad-chested body; a short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphic; adult male lions are larger than females and have a prominent mane. It is a social species, forming groups called prides. A lion's pride consists of a few adult males, related females, and cubs. Groups of female lions usually hunt together, preying mostly on large ungulates. The lion is an apex and keystone predator; although some lions scavenge when opportunities occur and have been known to hunt humans, lions typically do not actively seek out and prey on humans.

A black panther is the melanistic colour variant of the leopard and the jaguar. Black panthers of both species have excess black pigments, but their typical rosettes are also present. They have been documented mostly in tropical forests, with black leopards in Africa and Asia, and black jaguars in South America. Melanism is caused by a recessive allele in the leopard, and by a dominant allele in the jaguar.

<span class="mw-page-title-main">Leopard</span> Large spotted cat native to Africa and Asia

The leopard is one of the five extant species in the genus Panthera. It has a pale yellowish to dark golden fur with dark spots grouped in rosettes. Its body is slender and muscular reaching a length of 92–183 cm (36–72 in) with a 66–102 cm (26–40 in) long tail and a shoulder height of 60–70 cm (24–28 in). Males typically weigh 30.9–72 kg (68–159 lb), and females 20.5–43 kg (45–95 lb).

The term "big cat" is typically used to refer to any of the five living members of the genus Panthera, namely the tiger, lion, jaguar, leopard, and snow leopard, as well as the non-pantherine cheetah and cougar.

<i>Panthera spelaea</i> Extinct species of lion

Panthera spelaea, also known as the cave lion or steppe lion, is an extinct Panthera species that most likely evolved in Europe after the third Cromerian interglacial stage, less than 600,000 years ago. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion occurring in Africa and Asia, with the genetic divergence between the two species variously estimated between 1.9 million and 600,000 years ago. It is closely related and probably ancestral to the American lion. The species ranged from Western Europe to eastern Beringia in North America, and was a prominent member of the mammoth steppe fauna. It became extinct about 13,000 years ago.

<span class="mw-page-title-main">American lion</span> Extinct species of carnivore

Panthera atrox, better known as the American lion, also called the North American lion, or American cave lion, is an extinct pantherine cat. Panthera atrox lived in North America during the Pleistocene epoch, from around 340,000 to 12,800 years ago. The species was initially described by American paleontologist Joseph Leidy in 1853 based on a fragmentary mandible (jawbone) from Mississippi; the species name ('atrox') means "savage" or "cruel". The status of the species is debated, with some mammalogists and paleontologists considering it a distinct species or a subspecies of Panthera leo, which contains living lions. However, novel genetic evidence has shown that it is instead a distinct species derived from the Eurasian cave or steppe lion, evolving after its geographic isolation in North America. Its fossils have been excavated from Alaska to Mexico. It was about 25% larger than the modern lion, making it one of the largest known felids.

<span class="mw-page-title-main">Pantherinae</span> Subfamily of felids

The Pantherinae is a subfamily of the Felidae; it was named and first described by Reginald Innes Pocock in 1917 as only including the Panthera species. The Pantherinae genetically diverged from a common ancestor between 9.32 to 4.47 million years ago and 10.67 to 3.76 million years ago.

<span class="mw-page-title-main">African leopard</span> Leopard subspecies

The African leopard is the nominate subspecies of the leopard, native to many countries in Africa. It is widely distributed in most of sub-Saharan Africa, but the historical range has been fragmented in the course of habitat conversion. Leopards have also been recorded in North Africa as well.

<span class="mw-page-title-main">History of lions in Europe</span> Lions in prehistoric and historic Europe

The history of lions in Europe is based on fossils of Pleistocene and Holocene lions excavated in Europe since the early 19th century. The first lion fossil was excavated in southern Germany, and described by Georg August Goldfuss using the scientific name Felis spelaea. It probably dates to the Würm glaciation, and is 191,000 to 57,000 years old. Older lion skull fragments were excavated in Germany and described by Wilhelm von Reichenau under Felis fossilis in 1906. These are estimated at between 621,000 and 533,000 years old. The modern lion inhabited parts of Southern Europe since the early Holocene.

<i>Panthera gombaszoegensis</i> Extinct European jaguar species

Panthera gombaszoegensis, also known as the European jaguar, is a Panthera species that lived from about 2.0 to 0.35 million years ago in Europe The first fossils were excavated in 1938 in Gombasek, Slovakia. P. gombaszoegensis was medium-large sized species that formed an important part of the European carnivore guild for a period of over a million years. Many authors have posited that the species is the ancestor of the American jaguar, with some authors considering it the subspecies Panthera onca gombaszoegensis, though the close relationship between the two species has been questioned.

Panthera palaeosinensis was an early Pleistocene species from northern China. It is often incorrectly referenced as the ancestor of the tiger, Panthera tigris, although it shares features with all living large cats. Recent studies place it close to the base of the genus Panthera.

<i>Panthera fossilis</i> Fossil cat

Panthera fossilis, is an extinct species of cat belonging to the genus Panthera, known from remains found in Eurasia spanning the Middle Pleistocene and possibly into the Early Pleistocene. P. fossilis has sometimes been referred to by the common names steppe lion or cave lion, though these names are conventionally restricted to the later related species P. spelaea, to which P. fossilis is probably ancestral.

<span class="mw-page-title-main">Indian leopard</span> Leopard subspecies

The Indian leopard is a subspecies of the leopard that was first scientifically described in 1794. It is widely distributed on the Indian subcontinent. It is threatened by illegal trade of skins and body parts, and persecution due to human-leopard conflict and retaliation for livestock depredation.

<i>Panthera pardus tulliana</i> Leopard subspecies

Panthera pardus tulliana, also called Anatolian leopard, Persian leopard and Asia Minor leopard, is a leopard subspecies that was first described in 1856 based on a zoological specimen found in western Anatolia. It is native to the Iranian Plateau and the surrounding region from eastern Anatolia and the Caucasus to the Hindu Kush, where it inhabits foremost subalpine meadows, temperate broadleaf and mixed forests and rugged ravines at elevations of 600 to 3,800 m. It preys mostly on ungulates reliant on these habitats.

<i>Panthera onca mesembrina</i> Extinct subspecies of carnivore

Panthera onca mesembrina is an extinct subspecies of jaguar that was endemic to Patagonia in southern South America during the late Pleistocene epoch. It is known from several fragmentary specimens, the first of which found was in 1899 at "Cueva del Milodon" in Chile. These fossils were referred to a new genus and species "Iemish listai" by naturalist Santiago Roth, who thought they might be the bones of the mythological iemisch of Tehuelche folklore. A later expedition recovered more bones, including the skull of a large male that was described in detail by Angel Cabrera in 1934. Cabrera created a new name for the giant felid remains, Panthera onca mesembrina, after realizing that its fossils were near-identical to modern jaguars’. P. onca mesembrina's validity is disputed, with some paleontologists suggesting that it is a synonym of Panthera atrox.

<i>Panthera pardus spelaea</i> Pleistocene leopard subspecies

Panthera pardus spelaea, also known as the European Ice Age leopard and the Late Pleistocene leopard, is a fossil leopard subspecies which roamed Europe in the Late Pleistocene and possibly the Holocene.

<i>Panthera blytheae</i> Extinct species of mammal

Panthera blytheae is an extinct species of pantherine felid that lived during the late Messinian to early Zanclean ages approximately 5.95–4.1 million years ago.The first fossils were excavated in August 2010 in the Zanda Basin located in the Ngari Prefecture on the Tibetan Plateau; they were described and named in 2014.

Panthera uncia pyrenaica, also known as the Arago snow leopard or as the European snow leopard, is a subspecies of snow leopard that lived during the Middle Pleistocene in southern France.

References

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