Biology:Panthera

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Short description: Genus within Felidae

Panthera
Temporal range: 5.95–0 Ma
Late Miocene – Present
4panthera3.0.png
From top to bottom: tiger, lion, jaguar, leopard, snow leopard
Scientific classification e
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Oken, 1816[1]
Type species
Panthera pardus
Species
Synonyms[1]

Panthera is a genus within the family Felidae that was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group.[2][3] 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.[4] 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.[5][6]

Etymology

The word panther derives from classical Latin panthēra, itself from the ancient Greek pánthēr (πάνθηρ).[7]

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.[8] 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.[9] 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.[10]

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.[11] Genetic studies indicate that the pantherine cats diverged from the subfamily Felinae between six and ten million years ago.[5] The genus Neofelis is sister to Panthera.[5][12][13][14] 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.[15] The snow leopard is nested within Panthera and is the sister species of the tiger.[16]

Results of a 2016 study based on analysis of biparental nuclear genomes suggest the following relationships of living Panthera species:[17]

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.[18] Fossil remains found in South Africa that appear to belong within the Panthera lineage date to about 2 to 3.8 million years ago.[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 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.[20][21][22] 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.[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] 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]

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

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. delacouri Pocock, 1930[47]
Sri Lankan leopard P. p. kotiya Deraniyagala, 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. sondaica Temminck, 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[6]

SnowLeopard distribution.jpg

Extinct species and subspecies

Species and subspecies Fossil records Notes
Panthera blytheae Tibetan Plateau One of the oldest known Panthera species, possibly closely related to the snow leopard[11]
Panthera palaeosinensis Northern China Initially 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 China Possibly a close relative of the tiger[58]
Panthera youngi[59] China, Japan
Panthera atrox North America, dubious remains in South America.[60] 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]
Panthera balamoides[63] Mexico
Panthera gombaszoegensis Europe Panthera schreuderi and Panthera toscana are considered junior synonyms of P. gombaszoegensis. It is occasionally classified as a subspecies of P. onca.[64][65]
Panthera fossilis[66] Europe
Panthera spelaea Much of Eurasia[67] Originally spelaea was classified as a subspecies of the extant lion P. leo.[68] Results of recent genetic studies indicate that it belongs to a distinct species, namely P. spelaea.[69][70] Other genetic results indicate that P. fossilis also warrants status as a species.[71][72]
Panthera leo sinhaleyus Sri Lanka This lion subspecies was described on the basis of two teeth.[73]
Panthera onca augusta[74] North America May have lived in temperate forests across North America[75]
Panthera onca mesembrina[76] South America May have lived in grasslands in South America, unlike the modern jaguar
Panthera pardus spelaea Europe Closely related to Asiatic leopard subspecies,[77] with at least one study suggesting that it is closely related to the Persian leopard (P. p. tulliana) according to genetic work.[78]
Panthera shawi Laetoli site in Tanzania A leopard-like cat[79]
Panthera tigris acutidens Much of Asia Not closely related to modern tiger subspecies[80]
Panthera tigris soloensis Java, Indonesia Not closely related to modern tiger subspecies[80]
Panthera tigris trinilensis Java, Indonesia Not closely related to modern tiger subspecies[80]
Panthera principialis Tanzania Described in 2022.[81]
Panthera dhokpathanensis Siwaliks Described in 1986.[82]

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.[83]

Phylogeny

Two cladograms proposed for Panthera. The upper one is based on phylogenetic studies by Johnson et al. (2006),[5] and by Werdelin et al. (2010).[84] The lower cladogram is based on a study by Davis et al. (2010)[16] and by Mazák et al. (2011).[58]

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:[85]

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

References

  1. ↑ 1.0 1.1 1.2 1.3 Wozencraft, W.C. (2005). "Genus Panthera". 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. http://www.departments.bucknell.edu/biology/resources/msw3/browse.asp?id=14000227. 
  2. ↑ Oken, L. (1816). "1. Art, Panthera". Lehrbuch der Zoologie. 2. Abtheilung. Jena: August Schmid & Comp.. p. 1052. https://books.google.com/books?id=S5o5AAAAcAAJ&pg=PA1052. 
  3. ↑ Allen, J. A. (1902). "Mammal names proposed by Oken in his 'Lehrbuch der Zoologie'". Bulletin of the American Museum of Natural History 16 (27): 373−379. http://digitallibrary.amnh.org/bitstream/handle/2246/509//v2/dspace/ingest/pdfSource/bul/B016a27.pdf?sequence=1&isAllowed=y. 
  4. ↑ Pocock, R. I. (1916). "The Classification and Generic Nomenclature of F. uncia and its Allies". The Annals and Magazine of Natural History: Including Zoology, Botany, and Geology. Series 8 XVIII (105): 314–316. doi:10.1080/00222931608693854. https://archive.org/stream/ser8annalsmagazi18londuoft#page/314/mode/2up. 
  5. ↑ 5.0 5.1 5.2 5.3 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. doi:10.1126/science.1122277. PMID 16400146. Bibcode: 2006Sci...311...73J. https://zenodo.org/record/1230866. 
  6. ↑ 6.0 6.1 McCarthy, T.; Mallon, D.; Jackson, R.; Zahler, P.; McCarthy, K. (2017). "Panthera uncia". IUCN Red List of Threatened Species 2017: e.T22732A50664030. doi:10.2305/IUCN.UK.2017-2.RLTS.T22732A50664030.en. https://www.iucnredlist.org/species/22732/50664030. Retrieved 6 February 2023. 
  7. ↑ Liddell, H. G.; Scott, R. (1940). "πάνθηρ". A Greek-English Lexicon (Revised and augmented ed.). Oxford: Clarendon Press. https://www.perseus.tufts.edu/cgi-bin/ptext?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3D%2377441. 
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  9. ↑ Hast, M. H. (1989). "The larynx of roaring and non-roaring cats". Journal of Anatomy 163: 117–121. PMID 2606766. 
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  13. ↑ 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 (S1): S98–S116. doi:10.1007/PL00000060. PMID 9071018. Bibcode: 1997JMolE..44S..98J. https://zenodo.org/record/1232587. 
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  16. ↑ 16.0 16.1 Davis, B.W.; Li, G.; Murphy, W.J. (2010). "Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, Panthera (Carnivora: Felidae)". Molecular Phylogenetics and Evolution 56 (1): 64–76. doi:10.1016/j.ympev.2010.01.036. PMID 20138224. 
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  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). "The leopards of Africa". Proceedings of the Zoological Society of London 102 (2): 543–591. doi:10.1111/j.1096-3642.1932.tb01085.x. 
  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. et al. (1987). "Biochemical genetic variation in geographic isolates of African and Asiatic lions". National Geographic Research 3 (1): 114–124. http://www.cbs.umn.edu/sites/default/files/public/downloads/Biochemical_Genetic_Variation_in_Geographic_Isolates_of_African_and_Asiatic_Lions.pdf. 
  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. doi:10.1046/j.0962-1083.2001.01350.x. PMID 11883877. https://zenodo.org/record/1236516. 
  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. https://www.researchgate.net/publication/226066591. 
  27. ↑ 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. et al. (2011). "Genetic diversity, evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa". Journal of Biogeography 38 (7): 1356–1367. doi:10.1111/j.1365-2699.2011.02500.x. http://dspace.learningnetworks.org/bitstream/1820/4311/1/2011_Bertola,Hooft,Vrieling,Weerd,York,Bauer,Prins,Haes,Iongh_GeneticDiversityEvolutionaryHistoryAndImplicationsForConservationOfTheLionInWestAndCentralAfrica.pdf. Retrieved 17 January 2017. 
  28. ↑ Bertola, L. D.; Jongbloed, H.; Van Der Gaag, K. J.; De Knijff, P.; Yamaguchi, N.; Hooghiemstra, H.; Bauer, H.; Henschel, P. et al. (2016). "Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (Panthera leo)". Scientific Reports 6: 30807. doi:10.1038/srep30807. PMID 27488946. Bibcode: 2016NatSR...630807B. 
  29. ↑ Robinson, R. (1970). "Inheritance of black form of the leopard Panthera pardus". Genetica 41 (1): 190–197. doi:10.1007/bf00958904. PMID 5480762. 
  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. doi:10.1016/S0960-9822(03)00128-3. PMID 12620197. 
  31. ↑ 31.0 31.1 31.2 31.3 Kitchener, A. C.; Breitenmoser-Würsten, C.; Eizirik, E.; Gentry, A.; Werdelin, L.; Wilting, A.; Yamaguchi, N.; Abramov, A. V. et al. (2017). "A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group". Cat News (Special Issue 11): 66−75. https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y. 
  32. ↑ 32.0 32.1 32.2 32.3 32.4 32.5 Linnaeus, C. (1758). "Felis". Caroli Linnæi Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I (decima, reformata ed.). Holmiae: Laurentius Salvius. pp. 41−42. https://archive.org/stream/mobot31753000798865#page/41/mode/2up. 
  33. ↑ Meyer, J. N. (1826). Dissertatio inauguralis anatomico-medica de genere felium (Doctoral thesis). Vienna: University of Vienna.
  34. ↑ Smith, C. H. (1842). "Black maned lion Leo melanochaitus". in Jardine, W.. The Naturalist's Library. Vol. 15 Mammalia. London: Chatto and Windus. p. Plate X, 177. https://archive.org/stream/naturalistslibra15jardrich#page/176/mode/2up. 
  35. ↑ Mazak, V. (1975). "Notes on the Black-maned Lion of the Cape, Panthera leo melanochaita (Ch. H. Smith, 1842) and a Revised List of the Preserved Specimens". Verhandelingen Koninklijke Nederlandse Akademie van Wetenschappen (64): 1–44. 
  36. ↑ Bauer, H.; Packer, C.; Funston, P. F.; Henschel, P.; Nowell, K. (2016). "Panthera leo". IUCN Red List of Threatened Species 2016: e.T15951A115130419. https://www.iucnredlist.org/species/15951/115130419. 
  37. ↑ Larson, S. E. (1997). "Taxonomic re-evaluation of the jaguar" (in en). Zoo Biology 16 (2): 107–120. doi:10.1002/(SICI)1098-2361(1997)16:2<107::AID-ZOO2>3.0.CO;2-E. 
  38. ↑ Quigley, H.; Foster, R.; Petracca, L.; Payan, E.; Salom, R.; Harmsen, B. (2017). "Panthera onca". IUCN Red List of Threatened Species 2017: e.T15953A123791436. https://www.iucnredlist.org/species/15953/123791436. 
  39. ↑ Meyer, F. A. A. (1794). "Über de la Metheries schwarzen Panther". Zoologische Annalen. Erster Band. Weimar: Im Verlage des Industrie-Comptoirs. pp. 394–396. https://books.google.com/books?id=v4s-AAAAcAAJ&pg=PA394. 
  40. ↑ Cuvier, G. (1809). "Recherches sur les espėces vivantes de grands chats, pour servir de preuves et d'éclaircissement au chapitre sur les carnassiers fossils". Annales du Muséum National d'Histoire Naturelle Tome XIV: 136–164. 
  41. ↑ Hemprich, W.; Ehrenberg, C. G. (1830). "Felis, pardus?, nimr". in Dr. C. G. Ehrenberg. Symbolae Physicae, seu Icones et Descriptiones Mammalium quae ex Itinere per Africam Borealem et Asiam Occidentalem Friderici Guilelmi Hemprich et Christiani Godofredi Ehrenberg. Decas Secunda. Zoologica I. Mammalia II. Berolini: Officina Academica. pp. Plate 17. https://archive.org/details/SymbolaephysicaMammEhreA/page/n60. 
  42. ↑ Valenciennes, A. (1856). "Sur une nouvelles espèce de Panthère tué par M. Tchihatcheff à Ninfi, village situé à huit lieues est de Smyrne". Comptes rendus hebdomadaires des séances de l'Académie des sciences 42: 1035–1039. 
  43. ↑ Satunin, K. A. (1914). Opredelitel' mlekopitayushchikh Rossiiskoi Imperii. Tiflis: Tipographia Kantzelyarii Namestnichestva. 
  44. ↑ Pocock, R. I. (1927). "Description of two subspecies of leopards". Annals and Magazine of Natural History. Series 9 20 (116): 213–214. doi:10.1080/00222932708655586. 
  45. ↑ Schlegel, H. (1857). "Felis orientalis". Handleiding Tot de Beoefening der Dierkunde, Ie Deel. Breda: Boekdrukkerij van Nys. p. 23. 
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  49. ↑ Stein, A. B.; Athreya, V.; Gerngross, P.; Balme, G.; Henschel, P.; Karanth, U.; Miquelle, D.; Rostro, S. et al. (2016). "Panthera pardus". IUCN Red List of Threatened Species 2016: e.T15954A102421779. https://www.iucnredlist.org/species/15954/102421779. 
  50. ↑ Illiger, C. (1815). "Überblick der Säugethiere nach ihrer Verteilung über die Welttheile". Abhandlungen der Königlichen Preußischen Akademie der Wissenschaften zu Berlin 1804−1811: 39−159. http://bibliothek.bbaw.de/bbaw/bibliothek-digital/digitalequellen/schriften/anzeige/index_html?band=07-abh/18041811&seite:int=195. Retrieved 2 January 2021. 
  51. ↑ 51.0 51.1 51.2 Temminck, C. J. (1844). "Aperçu général et spécifique sur les Mammifères qui habitent le Japon et les Iles qui en dépendent". Fauna Japonica sive Descriptio animalium, quae in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Batava imperium tenent, suscepto, annis 1825–1830 collegit, notis, observationibus et adumbrationibus illustravit Ph. Fr. de Siebold. Leiden: Lugduni Batavorum. https://archive.org/details/faunajaponicasi00sieb/page/43. 
  52. ↑ Hilzheimer, M. (1905). "Über einige Tigerschädel aus der Straßburger zoologischen Sammlung". Zoologischer Anzeiger 28: 594–599. https://archive.org/stream/zoologischeranze28deut#page/594/mode/2up. 
  53. ↑ Mazák, V. (1968). "Nouvelle sous-espèce de tigre provenant de l'Asie du sud-est". Mammalia 32 (1): 104−112. doi:10.1515/mamm.1968.32.1.104. 
  54. ↑ Luo, S. J.; Kim, J. H.; Johnson, W. E.; Walt, J. v. d.; Martenson, J.; Yuhki, N.; Miquelle, D. G. (2004). "Phylogeography and Genetic Ancestry of Tigers (Panthera tigris)". PLOS Biology 2 (12): e442. doi:10.1371/journal.pbio.0020442. PMID 15583716. 
  55. ↑ Schwarz, E. (1912). "Notes on Malay tigers, with description of a new form from Bali". Annals and Magazine of Natural History Series 8 Volume 10 (57): 324–326. doi:10.1080/00222931208693243. https://archive.org/stream/annalsmagazineof8101912lond#page/324/mode/2up. 
  56. ↑ Goodrich, J.; Lynam, A.; Miquelle, D.; Wibisono, H.; Kawanishi, K.; Pattanavibool, A.; Htun, S.; Tempa, T. et al. (2015). "Panthera tigris". IUCN Red List of Threatened Species 2015: e.T15955A50659951. https://www.iucnredlist.org/species/15955/50659951. 
  57. ↑ Schreber, J. C. D. (1777). "Die Unze". Die Säugethiere in Abbildungen nach der Natur mit Beschreibungen. Erlangen: Wolfgang Walther. pp. 386–387. https://archive.org/stream/SaYugthiereAbbiIIISchr#page/386/mode/2up. 
  58. ↑ 58.0 58.1 58.2 58.3 Mazák, J. H.; Christiansen, P.; Kitchener, A. C. (2011). "Oldest Known Pantherine Skull and Evolution of the Tiger". PLOS ONE 6 (10): e25483. doi:10.1371/journal.pone.0025483. ISSN 1932-6203. PMID 22016768. Bibcode: 2011PLoSO...625483M. 
  59. ↑ Pei, W. C. (1934). "On the Carnivora from Locality 1 of Choukoutien". Palaeontologica Sinica Series C, Fascicle 1: 1−166. 
  60. ↑ Chimento, N. R.; Agnolin, F. L. (2017). "The fossil American lion (Panthera atrox) in South America: Palaeobiogeographical implications". Comptes Rendus Palevol 16 (8): 850–864. doi:10.1016/j.crpv.2017.06.009. Bibcode: 2017CRPal..16..850C. 
  61. ↑ 61.0 61.1 Barnett, R.; Shapiro, B.; Barnes, I.; Ho, 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". Molecular Ecology 18 (8): 1668–77. doi:10.1111/j.1365-294X.2009.04134.x. PMID 19302360. https://www.researchgate.net/publication/24216045. 
  62. ↑ Sotnikova, M.; Nikolskiy, P. (2006). "Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters" (in en). Quaternary International 142–143: 218–228. doi:10.1016/j.quaint.2005.03.019. Bibcode: 2006QuInt.142..218S. http://doc.rero.ch/record/13484/files/PAL_E260.pdf. 
  63. ↑ Stinnesbeck, S. R.; Stinnesbeck, W.; Frey, E.; Olguín, J. A.; Sandoval, C. R.; Morlet, A. V.; González, A. H. (2018). "Panthera balamoides and other Pleistocene felids from the submerged caves of Tulum, Quintana Roo, Mexico". Historical Biology: An International Journal of Paleobiology 32 (7): 1–10. doi:10.1080/08912963.2018.1556649. 
  64. ↑ Hemmer, H.; Kahlke, R. D.; Vekua, A. K. (2010). "Panthera onca georgica ssp. nov. from the Early Pleistocene of Dmanisi (Republic of Georgia) and the phylogeography of jaguars (Mammalia, Carnivora, Felidae)". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 257 (1): 115–127. doi:10.1127/0077-7749/2010/0067. 
  65. ↑ Mol, D.; van Logchem, W.; de Vos, J. (2011). "New record of the European jaguar, Panthera onca gombaszoegensis (Kretzoi, 1938), from the Plio-Pleistocene of Langenboom (The Netherlands)". Cainozoic Research 8 (1–2): 35–40. http://natuurtijdschriften.nl/download?type=document;docid=541752. Retrieved 28 September 2015. 
  66. ↑ Harington, C. R. (1996). Pleistocene mammals of the Yukon Territory (PhD). Edmonton: University of Alberta.
  67. ↑ Stuart, A. J., Lister, A .M. (2011). "Extinction chronology of the cave lion Panthera spelaea". Quaternary Science Reviews 30 (17): 2329–2340. doi:10.1016/j.quascirev.2010.04.023. Bibcode: 2011QSRv...30.2329S. 
  68. ↑ Sala, B. (1990). "Panthera leo fossilis (v. Reichenau, 1906) (Felidae) de Iserna la Pineta (Pléistocene moyen inférieur d'Italie)". Géobios 23 (2): 189–194. doi:10.1016/S0016-6995(06)80051-3. 
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  70. ↑ Marciszak, A.; Schouwenburg, C.; Darga, R. (2014). "Decreasing size process in the cave (Pleistocene) lion Panthera spelaea (Goldfuss, 1810) evolution – A review". Quaternary International. Fossil remains in karst and their role in reconstructing Quaternary paleoclimate and paleoenvironments 339–340: 245–257. doi:10.1016/j.quaint.2013.10.008. Bibcode: 2014QuInt.339..245M. 
  71. ↑ Sotnikova, M. V.; Foronova, I. V. (2014). "First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia.". Integrative Zoology 9 (4): 517–530. doi:10.1111/1749-4877.12082. PMID 24382145. https://www.researchgate.net/publication/259545849. 
  72. ↑ Barnett, R.; Mendoza, M. L. Z.; Soares, A. E. R.; Ho, S. Y. W.; Zazula, G.; Yamaguchi, N.; Shapiro, B.; Kirillova, I. V. et al. (2016). "Mitogenomics of the Extinct Cave Lion, Panthera spelaea (Goldfuss, 1810), resolve its position within the Panthera cats". Open Quaternary 2: 4. doi:10.5334/oq.24. 
  73. ↑ 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. http://rmbr.nus.edu.sg/rbz/biblio/s12/s12rbz423-434.pdf. Retrieved 28 October 2013. 
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  76. ↑ Roth, S. (1899). "Descripción de los restos encontrados en la caverna de Última Esperanza". Revista del Museo la Plata 9: 381–388. 
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  79. ↑ Sabol, M. (2011). "Masters of the lost world: a hypothetical look at the temporal and spatial distribution of lion-like felids". Quaternaire 4: 229–236. https://www.researchgate.net/publication/287422796. 
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  81. ↑ Hemmer, H. (2022). "The identity of the "lion", Panthera principialis sp. nov., from the Pliocene Tanzanian site of Laetoli and its significance for molecular dating the pantherine phylogeny, with remarks on Panthera shawi (Broom, 1948), and a revision of Puma incurva (Ewer, 1956), the Early Pleistocene Swartkrans "leopard" (Carnivora, Felidae)". Palaeobiodiversity and Palaeoenvironments 103 (2): 465–487. doi:10.1007/s12549-022-00542-2. 
  82. ↑ Bakr, A. (1986). "On a collection of Siwalik Carnivora". Biological Society of Pakistan 11: 1–64. 
  83. ↑ Turner, A. (1984). "Panthera crassidens Broom, 1948. The cat that never was?". South African Journal of Science 80 (5): 227–233. https://journals.co.za/docserver/fulltext/sajsci/80/5/7798.pdf?expires=1544536360&id=id&accname=guest&checksum=1BB15474E5A55CD94EC66579004AE19D. 
  84. ↑ Werdelin, L.; Yamaguchi, N.; Johnson, W. E.; O'Brien, S. J. (2010). "Phylogeny and evolution of cats (Felidae)". Biology and Conservation of Wild Felids. Oxford: Oxford University Press. pp. 59–82. ISBN 978-0-19-923445-5. https://www.researchgate.net/publication/266755142. 
  85. ↑ Piras, P.; Silvestro, D.; Carotenuto, F.; Castiglione, S.; Kotsakis, A.; Maiorino, L.; Melchionna, M.; Mondanaro, A. et al. (2018). "Evolution of the sabertooth mandible: A deadly ecomorphological specialization". Palaeogeography, Palaeoclimatology, Palaeoecology 496: 166−174. doi:10.1016/j.palaeo.2018.01.034. Bibcode: 2018PPP...496..166P. 

Further reading

External links

Wikidata ☰ Q127960 entry



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