Panthera is a
genus within the
familyFelidae, 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.
In Panthera species, the dorsal profile of the
skull is flattish or evenly convex. The frontal inter
orbital 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 leopardgenetic 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]
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
mitochondrialsister 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]
Ranged 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]
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]
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
The
cladogram below follows Mazák, Christiansen and Kitchener (2011).[58]
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