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, as well as a number of extinct species.

Etymology

The word panther derives from classical Latin panthēra, itself from the ancient Greek pánthēr (πάνθηρ).[3]

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.[4] 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.[5] 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.[6]

Evolution

The geographic origin of the Panthera is most likely northern Central Asia. Panthera blytheae, possibly the
oldest known Panthera species, is similar in skull features to the snow leopard,[7] though its taxonomic
placement has been disputed by other researchers who suggest that the species likely belongs to a different
genus.[8][9] The tiger, snow leopard, and clouded leopard genetic lineages dispersed in Southeast Asia during
the Miocene.[7] Genetic studies indicate that the pantherine cats diverged from the subfamily Felinae between
six and ten million years ago.[10] The genus Neofelis is sister to Panthera.[10][11][12][13] The clouded leopard
appears to have diverged about 8.66 (https://geoltime.github.io/?Ma=8.66) million years ago. Panthera
diverged from other cat species about 11.3 (https://geoltime.github.io/?Ma=11.3) million years ago and then
evolved into the species tiger about 6.55 (https://geoltime.github.io/?Ma=6.55) million years ago, snow
leopard about 4.63 (https://geoltime.github.io/?Ma=4.63) million years ago and leopard about
4.35 (https://geoltime.github.io/?Ma=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 (https://geoltime.github.io/?Ma=0.34) million years ago, and that both P. atrox
and P. spelaea are most closely related to lions among living
Panthera
Panthera species.[14] The snow leopard is nested within
Temporal range:
Panthera and is the sister species of the tiger.[15]

Results of a 2016 study based on analysis of biparental

nuclear genomes suggest the following relationships of
living Panthera species:[16]

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 (https://geoltime.github.io/?Ma=1.6) million years ago.
[17]
Fossil remains found in South Africa that appear to

From top to bottom: tiger, lion, jaguar, leopard,

snow leopard

Scientific classification

Domain: Eukaryota

Kingdom: Animalia
belong within the Panthera lineage date to about Phylum: Chordata

Class: Mammalia

Order: Carnivora

Suborder: Feliformia

Family: Felidae

Subfamily: Pantherinae

Genus: Panthera
Oken, 1816[2]

Type species

Felis pardus
(= Panthera pardus)
Linnaeus, 1758[2]

Species

†Panthera atrox (Leidy, 1853)

†Panthera balamoides Stinnesbeck et al.,

2019

†Panthera blytheae Tseng et al., 2014

†Panthera dhokpathanensis Bakr, 1986

†Panthera fossilis (Reichenau, 1906)

†Panthera gombaszoegensis (Kretzoi,

1938)

Panthera leo (Linnaeus, 1758)

Panthera onca (Linnaeus, 1758)

†Panthera palaeosinensis (Zdansky, 1924)

Panthera pardus (Linnaeus, 1758)

†Panthera principialis Hemmer, 2023

†Panthera shawi (Broom, 1948)

†Panthera spelaea (Goldfuss, 1810)

Panthera tigris (Linnaeus, 1758)

 Panthera uncia (Schreber, 1775)

†Panthera youngi Pei, 1934

†Panthera zdanskyi Mazák, Christiansen &

Kitchener, 2011

Synonyms[2]

About 10 [show]

2 to 3.8 (https://geoltime.github.io/?Ma=2.0%E2%80%933.8) million years ago.[18]

Classification

Panthera was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group.[19][20]
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.[21][22][23] 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.[24] 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.[25][26] Subsequently, it was proposed to group all African leopard populations to P. p.
pardus and retain eight subspecific names for Asian leopard populations.[27] 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.[10][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:[2][34]
 IUCN Red List status
Species Subspecies
and distribution

Lion P. leo (Linnaeus, P. l. leo (Linnaeus, 1758)[35] including: VU [39]

[35]
1758) Barbary lion P. l. leo sensu stricto

Asiatic lion syn. P. l. persica (Johann Nepomuk Meyer, 1826)[36]

P. l. melanochaita (Smith, 1842)[37] including:

†Cape lion P. l. melanochaita sensu stricto[38]

Jaguar P. onca Monotypic[40][34] NT [41]

[35]
(Linnaeus, 1758)

Leopard P. pardus African leopard P. p. pardus (Linnaeus, 1758)[35] VU [52]

[35]
(Linnaeus, 1758)
Indian leopard P. p. fusca (Meyer, 1794)[42]
Javan leopard P. p. melas (G. Cuvier, 1809)[43]
Arabian leopard P. p. nimr (Hemprich and Ehrenberg), 1833[44]
P. p. tulliana (Valenciennes, 1856),[45] syn. P. p. ciscaucasica (Satunin, 1914),[46] P. p.
saxicolor Pocock, 1927[47]

Amur leopard P. p. orientalis (Schlegel, 1857),[48] syn. P. p. japonensis (Gray,

1862)[49]

Indochinese leopard P. p. delacouri Pocock, 1930[50]

Sri Lankan leopard P. p. kotiya Deraniyagala, 1956[51]

Tiger P. tigris P. t. tigris (Linnaeus, 1758) including: EN [59]

[35]
(Linnaeus, 1758) Bengal tiger P. t. tigris sensu stricto

†Caspian tiger syn. P. t. virgata (Illiger, 1815)[53]

Siberian tiger syn. P. t. altaica (Temminck, 1844)[54]

South China tiger syn. P. t. amoyensis (Hilzheimer, 1905)[55]

Indochinese tiger syn. P. t. corbetti Mazák, 1968[56]

Malayan tiger syn. P. t. jacksoni Luo et al., 2004[57]

Sunda Island tiger P. t. sondaica Temminck, 1844)[54] including

†Javan tiger P. t. sondaica sensu stricto

Sumatran tiger syn. P t. sumatrae (Temminck, 1844)[54]

†Bali tiger syn. P. t. balica Schwarz, 1912)[58]

Snow leopard P. Monotypic[34] VU [28]

uncia[34] (Schreber,
 [60]
1775)
Extinct species and subspecies

Species and
Fossil records Notes Images
subspecies

Intitally described as one of the oldest known Panthera species, possibly

Panthera blytheae Tibetan Plateau closely related to the snow leopard,[7] though other researchers consider it as
a species of a different genus.[8][9]

Panthera Initially thought to be an ancestral tiger species, but several scientists place it
Northern China
palaeosinensis close to the base of the genus Panthera[1]

Panthera zdanskyi Gansu province of Possibly a close relative of the tiger[1]

northwestern
China

Panthera youngi[61] China, Japan

Panthera atrox North America, Commonly known as the American lion, P. atrox is thought to have descended
dubious remains from a basal P. spelaea cave lion population isolated south of the Cordilleran
in South Ice Sheet, and then established a mitochondrial sister clade circa 200,000
[62]
America. BP.[63] It was sometimes considered a subspecies either under the
nomenclature of P. leo[63] or P. spelaea.[64] One of the largest Panthera
species.[65] Became extinct around 13,000-12,000 years ago.[66]

Panthera Mexico Dubious, other authors suggest that the remains are actually of the extinct
[67]
balamoides bear Arctotherium instead.[68]

Panthera Europe Ranged across Europe from around 2 million to 350,000 years ago.[69]
gombaszoegensis Panthera schreuderi and Panthera toscana are considered junior synonyms of
P. gombaszoegensis. It is occasionally classified as a subspecies of P.
onca.[70][71]

Panthera fossilis[72] Europe Likely the ancestor of P. spelaea [73]

Panthera spelaea Much of Commonly known as the cave lion or steppe lion. Originally spelaea was
Eurasia[74] classified as a subspecies of the extant lion P. leo.[75] 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.[76][77] Other genetic results indicate that P. fossilis also warrants
status as a species.[78][79] Became extinct around 14,500-14,000 years ago.[80]

Panthera leo Sri Lanka This lion subspecies was described on the basis of two teeth.[81]
sinhaleyus

Panthera onca North America May have lived in temperate forests across North America[83]
augusta[82]

Panthera onca
South America May have lived in grasslands in South America, unlike the modern jaguar
mesembrina[84]

Panthera pardus Europe Closely related to Asiatic leopard subspecies,[85]

spelaea
Panthera shawi Laetoli site in A leopard-like cat[86]
Tanzania

Panthera tigris Much of Asia Not closely related to modern tiger subspecies[87]
acutidens

Panthera tigris
Java, Indonesia Not closely related to modern tiger subspecies[87]
soloensis

Panthera tigris
Java, Indonesia Not closely related to modern tiger subspecies[87]
trinilensis

Panthera
Tanzania Described in 2022.[88]
principialis

Panthera
Siwaliks Described in 1986.[89]
dhokpathanensis

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.[90]

Phylogeny

Two cladograms proposed for

Panthera. The upper one is based on
phylogenetic studies by Johnson et
al. (2006),[10] and by Werdelin et al.
(2010).[91] The lower cladogram is
based on a study by Davis et al.
(2010)[15] and by Mazák et al.
(2011).[1]

The cladogram below follows Mazák, Christiansen and Kitchener (2011).[1]

 Neofelis

Snow leopard

†Panthera palaeosinensis

Jaguar

†American lion

†Panthera spelaea
Pantherinae
Panthera
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:[92]

†Panthera palaeosinensis

†Panthera blytheae

Snow leopard

†Panthera zdanskyi

Tiger

†Panthera gombaszoegensis (sometimes called the European jaguar)

Jaguar
Panthera

Leopard

Lion

†Panthera spelaea (cave lion or steppe lion)

†American lion (Panthera atrox)

See also

Panthera hybrid
 Panther (legendary creature)
Cats portal

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