Panthera

Panthera[1]
Temporal range: Late Miocene - Present, 5.95–0 Ma
Tiger (Panthera tigris), the largest of the Panthera genus
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Oken, 1816
Type species
Felis pardus
Linnaeus, 1758
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]

Name

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

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 and this along with a specially adapted larynx that has proportionally larger vocal folds that are covered in a large fibro-elastic pad, is what enables 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 Panthera genus 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] More recent genetic research indicates that the Western and Central African lion populations form a different clade and are more related to the Asian lion than to lions from southern or eastern Africa. These populations have been largely ignored in previous studies.[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 cladogram is based on a 2006 phylogenetic study by Warren E. Johnson (of the National Cancer Institute) and colleagues, and a 2009 study by Lars Werdelin and colleagues. The lower cladogram is based on a 2010 study by Brian W. Davis (of the Texas A&M University) and colleagues and a 2011 study by Ji H. Mazák (of the Shanghai Science and Technology Museum) and colleagues.

The cladogram below follows Mazák, Christiansen and Kitchener (2011).[31]

Pantherinae


Neofelis


Panthera


Panthera uncia


Panthera palaeosinensis†



Panthera onca


Panthera atrox†


Panthera spelaea†


Panthera leo

Panthera pardus


Panthera tigris

Panthera zdanskyi†







Species

The genus Panthera comprises:[1]

Taxonomic placing is uncertain for the extinct fossil Panthera species:

See also

References

  1. 1 2 Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 546–548. ISBN 978-0-8018-8221-0. OCLC 62265494.
  2. ↑ Oken, L. (1816). Lehrbuch der Zoologie, 2. Abtheilung. August Schmid & Comp., Jena.
  3. ↑ Pocock, R. I. (1916). The Classification and Generic Nomenclature of F. uncia and its Allies. The Annals and Magazine of Natural History: zoology, botany, and geology. Series 8, Volume XVIII: 314–316.
  4. 1 2 3 4 5 Johnson, W. E.; Eizirik, E.; Pecon-Slattery, J.; Murphy, W. J.; Antunes, A.; Teeling, E. & O'Brien, S. J. (2006). "The Late Miocene radiation of modern Felidae: A genetic assessment". Science. 311 (5757): 73–77. Bibcode:2006Sci...311...73J. PMID 16400146. doi:10.1126/science.1122277.
  5. 1 2 Jackson, R.; Mallon, D.; McCarthy, T.; Chundaway, R. A.; Habib, B. (2008). "Panthera uncia". IUCN Red List of Threatened Species. Version 2017-1. International Union for Conservation of Nature.
  6. ↑ Nowak, R. M. (1999). Walker's Mammals of the World. Johns Hopkins University Press. ISBN 0-8018-5789-9.
  7. ↑ Liddell, H. G. & R. Scott (1940). "πάνθηρ". A Greek-English Lexicon, revised and augmented. Oxford: Clarendon Press.
  8. ↑ OUP (2002). "Panther". Oxford English Dictionary. Oxford: Oxford University Press.
  9. ↑ Pocock, R. I. (1939). The fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1. Taylor and Francis, London.
  10. ↑ Hast, M. H. (1989). "The larynx of roaring and non-roaring cats". Journal of Anatomy. 163: 117–121. ISSN 0021-8782. PMC 1256521 Freely accessible. PMID 2606766.
  11. ↑ Turner, A. (1987). "New fossil carnivore remains from the Sterkfontein hominid site (Mammalia: Carnivora)". Annals of the Transvaal Museum. 34 (15): 319–347.
  12. ↑ Davis, B. W.; Li, G..; Murphy, W.J. (Jul 2010). "Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, Panthera (Carnivora: Felidae)". Molecular Phylogenetics and Evolution. 56 (1): 64–76. PMID 20138224. doi:10.1016/j.ympev.2010.01.036.
  13. ↑ Yu, L.; Zhang, Y. P. (2005). "Phylogenetic studies of pantherine cats (Felidae) based on multiple genes, with novel application of nuclear beta-fibrinogen intron 7 to carnivores". Molecular Phylogenetics and Evolution. 35 (2): 483–495. PMID 15804417. doi:10.1016/j.ympev.2005.01.017.
  14. 1 2 Janczewski, D. N.; Modi, W. S.; Stephens, J. C.; O'Brien, S. J. (1996). "Molecular Evolution of Mitochondrial 12S RNA and Cytochrome b Sequences in the Pantherine Lineage of Felidae". Molecular Biology and Evolution. 12 (4): 690–707. PMID 7544865. doi:10.1093/oxfordjournals.molbev.a040232. Retrieved 2006-08-06.
  15. 1 2 Johnson, W. E.; O'Brien, S. J. (1997). "Phylogenetic reconstruction of the Felidae using 16S rRNA and NADH-5 mitochondrial genes". Journal of Molecular Evolution. 44: S98–S116. PMID 9071018. doi:10.1007/PL00000060.
  16. ↑ Yu L & Zhang YP (2005). "Phylogenetic studies of pantherine cats (Felidae) based on multiple genes, with novel application of nuclear beta-fibrinogen intron 7 to carnivores". Molecular Phylogenetics and Evolution. 35 (2): 483–495. PMID 15804417. doi:10.1016/j.ympev.2005.01.017.
  17. ↑ Wei L, Wu X, Zhu L, Jiang Z (2010). "Mitogenomic analysis of the genus Panthera". Science China Life Sciences. 54 (10): 917–930. PMID 22038004. doi:10.1007/s11427-011-4219-1.
  18. ↑ Barnett, R.; Shapiro, B.; Barnes, I.; Yo, S. Y.W.; Burger, J.; Yamaguchi, N.; Higham, T. F.G.; Wheeler, H. T.; et al. (2009). "Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity" (PDF). Molecular Ecology. 18 (8): 1668–1677. PMID 19302360. doi:10.1111/j.1365-294X.2009.04134.x. Retrieved 2011-11-23.
  19. ↑ Hemmer, H.; Kahlke, R. D.; Vekua, A. K. (2001). "The Jaguar – Panthera onca gombaszoegensis (Kretzoi, 1938) (Carnivora: Felidae) in the late lower pleistocene of Akhalkalaki (south Georgia; Transcaucasia) and its evolutionary and ecological significance". Geobios. 34 (4): 475–486. doi:10.1016/s0016-6995(01)80011-5.
  20. ↑ Pocock, R. I. (1930). "The panthers and ounces of Asia". Journal of the Bombay Natural History Society. 34 (1): 65–82.
  21. ↑ Pocock, R. I. (1932). [10.1111/j.1096-3642.1932.tb01085.x "The leopards of Africa"] Check |url= value (help). Proceedings of the Zoological Society of London. 102 (2): 543–591.
  22. ↑ Pocock, R. I. (1939). "The races of jaguar (Panthera onca)". Novitates Zoologicae. 41: 406–422.
  23. ↑ O’Brien, S. J.; Martenson, J. S.; Packer, C.; Herbst, L.; de Vos, V.; Joslin, P.; Ott-Joslin, J.; Wildt, D. E. & Bush, M. (1987). "Biochemical genetic variation in geographic isolates of African and Asiatic lions" (PDF). National Geographic Research. 3 (1): 114–124.
  24. ↑ Miththapala, S.; Seidensticker, J.; O'Brien, S. J. (1996). "Phylogeographic subspecies recognition in leopards (Panthera pardus): Molecular genetic variation". Conservation Biology. 10 (4): 1115–1132. doi:10.1046/j.1523-1739.1996.10041115.x.
  25. ↑ Uphyrkina, O.; Johnson, W. E.; Quigley, H. B.; Miquelle, D. G.; Marker, L.; Bush, M. E.; O'Brien, S. J. (2001). "Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus". Molecular Ecology. 10 (11): 2617–2633. PMID 11883877. doi:10.1046/j.0962-1083.2001.01350.x.
  26. ↑ Dubach, J.; Patterson, B. D.; Briggs, M. B.; Venzke, K.; Flamand, J.; Stander, P.; Scheepers, L.; Kays, R. W. (2005). "Molecular genetic variation across the southern and eastern geographic ranges of the African lion, Panthera leo". Conservation Genetics. 6 (1): 15–24. doi:10.1007/s10592-004-7729-6.
  27. ↑ Bertola, L., de Iongh, H., Vrieling, K. (2011). Researchers confirm West and Central African lion is different from other lions. University of Leiden. Institute of Environmental Sciences (CML). Faculty of Science.
  28. ↑ Bertola, L. D.; Van Hooft, W. F.; Vrieling, K.; Uit De Weerd, D. R.; York, D. S.; Bauer, H.; Prins, H. H. T.; Funston, P. J.; Udo De Haes, H. A.; Leirs, H.; Van Haeringen, W. A.; Sogbohossou, E.; Tumenta, P. N.; De Iongh, H. H. (2011). "Genetic diversity, evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa" (PDF). Journal of Biogeography. 38 (7): 1356–1367. doi:10.1111/j.1365-2699.2011.02500.x.
  29. ↑ Robinson, R. (1970). "Inheritance of black form of the leopard Panthera pardus". Genetica. 41 (1): 190–197. PMID 5480762. doi:10.1007/bf00958904.
  30. ↑ Eizirik, E.; Yuhki, N.; Johnson, W. E.; Menotti-Raymond, M.; Hannah, S. S.; O'Brien, S. J. (2003). "Molecular Genetics and Evolution of Melanism in the Cat Family". Current Biology. 13 (5): 448–453. PMID 12620197. doi:10.1016/S0960-9822(03)00128-3.
  31. ↑ Mazák, Ji H.; Christiansen, Per; Kitchener, Andrew C. (2011). "Oldest Known Pantherine Skull and Evolution of the Tiger". PLoS ONE. 6 (10): e25483. Bibcode:2011PLoSO...625483M. PMC 3189913 Freely accessible. PMID 22016768. doi:10.1371/journal.pone.0025483.
  32. ↑ Luo, S.J.; Kim, J.H.; Johnson, W.E.; Walt Jvd; Martenson, J.; et al. (2004). "Phylogeography and Genetic Ancestry of Tigers (Panthera tigris)". PLoS Biol. 2 (12): e442. PMC 534810 Freely accessible. PMID 15583716. doi:10.1371/journal.pbio.0020442.
  33. ↑ Jackson, P.; Nowell, K. (2008). "Panthera tigris ssp. virgata". IUCN Red List of Threatened Species. Version 2012.2. International Union for Conservation of Nature.
  34. ↑ Jackson, P. Nowell, K. (2008). "Panthera tigris ssp. balica". IUCN Red List of Threatened Species. Version 2012.2. International Union for Conservation of Nature.
  35. ↑ Jackson, P.; Nowell, K. (2008). "Panthera tigris ssp. sondaica". IUCN Red List of Threatened Species. Version 2012.2. International Union for Conservation of Nature.
  36. ↑ Hooijer, D. A. (1947). Pleistocene remains of Panthera tigris (Linnaeus) subspecies from Wanhsien, Szechwan, China, compared with fossil and recent tigers from other localities. American Museum Novitates no. 1346.
  37. ↑ Brongersma, L. D. (1935). "Notes on some recent and fossil cats, chiefly from the Malay Archipelago". Zoologische Mededelingen 18: 1–89.
  38. 1 2 Barnett, R.; Yamaguchi, N.; Barnes, I.; Cooper, A. (2006). "Lost populations and preserving genetic diversity in the lion Panthera leo: Implications for its ex situ conservation" (PDF). Conservation Genetics. 7 (4): 507–514. doi:10.1007/s10592-005-9062-0.
  39. ↑ Manamendra-Arachchi, K., Pethiyagoda, R., Dissanayake, R., Meegaskumbura, M. (2005). A second extinct big cat from the late Quaternary of Sri Lanka. The Raffles Bulletin of Zoology, Supplement 12: 423–434.
  40. ↑ Tchernov, E.; Tsoukala, E. (1997). "Middle Pleistocene (early Toringian) carnivore remains from northern Israel". Quaternary Research. 48: 122–136. Bibcode:1997QuRes..48..122T. doi:10.1006/qres.1997.1901.
  41. 1 2 Harington, C. R. (1996). Pleistocene mammals of the Yukon Territory. Ph.D. dissertation, University of Alberta, Edmonton.
  42. ↑ Nowell, K.; Jackson, P., eds. (1996). "Panthera Onca". Wild Cats. Status Survey and Conservation Action Plan (PDF). Gland, Switzerland: IUCN/SSC Cat Specialist Group. IUCN. pp. 118–122. Retrieved 2015-09-04.
  43. ↑ Khorozyan, I. (2008). "Panthera pardus ssp. saxicolor". IUCN Red List of Threatened Species. Version 2013.1. International Union for Conservation of Nature.
  44. ↑ O'Regan, H.; Turner, A (2004). "Biostratigraphic and palaeoecological implications of new fossil felid material from the Plio-Pleistocene site of Tegelen, the Netherlands". Palaeontology. 47 (5): 1181–1193. doi:10.1111/j.0031-0239.2004.00400.x.

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