Panthera

[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.

[8] Genetic studies indicate that the pantherine cats diverged from the subfamily Felinae between six and ten million years ago.

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

[21][22] 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.

[23][24][25] 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.

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

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.

Two cladograms proposed for Panthera . The upper one is based on phylogenetic studies by Johnson et al. (2006), [ 12 ] and by Werdelin et al. (2010). [ 96 ] The lower cladogram is based on a study by Davis et al. (2010) [ 17 ] and by Mazák et al. (2011). [ 1 ]