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Panthera spelaea, also known as the Eurasian cave lion, European cave lion, or steppe lion,[1] is an extinct Panthera species that evolved in Europe probably after the third Cromerian interglacial stage, less than 600,000 years ago. Phylogenetic analysis of fossil bone samples revealed that it was highly distinct and genetically isolated from the modern lion (Panthera leo) occurring in Africa and Asia.[2] Analysis of morphological differences and mitochondrial data support the taxonomic recognition of Panthera spelaea as a distinct species that diverged from the lion about 1.9 million years ago.[3][4] The oldest known bone fragments were excavated in Yakutia and radiocarbon dated at least 62,400 years old. It became extinct about 13,000 years ago.[5]

Panthera spelaea
Temporal range: Middle Pleistocene to Early Holocene, 0.1–0.011 Ma
Panthera leo spelaea in Vienna.jpg
Skeleton in Natural History Museum, Vienna
Scientific classification edit
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
P. spelaea
Binomial name
Panthera spelaea
Goldfuss, 1810
Cave lion range.png
Red indicates the maximal range of Panthera spelaea, blue Panthera atrox, and green Panthera leo.

Panthera leo spelaea M. Boule & L. De Villeneuve, 1927; Panthera leo vereshchagini Baryshnikov & Boeskorov, 2001


Cave lion skull exhibited in the Muséum de Toulouse

Several authors regarded Panthera spelaea as a subspecies of the modern lion and therefore Panthera leo spelaea.[6][7][8][2] However, results from morphological studies indicate that it is distinct in cranial and dental anatomy to justify specific status of Panthera spelaea.[9][10] Results of phylogenetic studies also support this assessment.[11][3][4]

In 2001, the subspecies P. spelaea vereshchagini was proposed for seven specimens found in Siberia and Yukon, which have smaller skulls and teeth than average member of P. spelaea.[12] Genetic analysis using ancient DNA provided no evidence for their distinct subspecific status; DNA signatures from P. spelaea from Europe and Alaska were indistinguishable, suggesting one large panmictic population.[3]

One author considered the cave lion to be more closely related to the tiger based on a comparison of skull shapes, and proposed the scientific name Panthera tigris spelaea.[13]


Lions and bisons depicted in the Chauvet Cave, France[14]

Panthera spelaea evolved in the late Saalian glaciation or early Eemian from the earlier P. fossilis, which first appeared in Eurasia about 700,000 years ago.[2] P. spelaea bone fragments excavated in Poland were radiocarbon dated to between the early and late Weichselian glaciation, and are between 109,000 and 57,000 years old.[15] In Eurasia, it became extinct between 14,900 and 14,100 years ago, and survived in Beringia until 13,800 to 13,300 years ago as the Weichselian glaciation receded.[5]

Mitochondrial DNA sequence data from fossil lion remains show that the American lion represents a sister group of P. spelaea, and likely arose when an early P. spelaea population became isolated south of the Cordilleran Ice Sheet about 340,000 years ago.[3] The following cladogram shows the genetic relationship between P. spelaea and other pantherine cats.[4]

Felis catus 

Neofelis nebulosa 

Panthera tigris sondaica

Panthera tigris tigris 

Panthera onca 

Panthera uncia 

Panthera pardus 

Panthera pardus orientalis 

Panthera leo leo 

Panthera leo melanochaita

Panthera spelaea 


Skeletal diagram
Cave lions, Chamber of Felines, Lascaux caves

P. spelaea is known from Paleolithic cave paintings, ivory carvings, and clay figurines. These archaeological artifacts indicate that it may have been featured in Paleolithic religious rituals.[16][17] A drawing in France's Chauvet cave, estimated to be 30,000 years old, depicts two cave lions walking together. The one in the foreground is slightly smaller than the one in the background, which has been drawn with a scrotum and without a mane.[18] This suggests male cave lions completely lacked manes, or had very small manes.[5]

Prior to its split from Panthera leo, cave lions were thought to have been one of the largest lion subspecies. The skeleton of an adult male found in 1985 near Siegsdorf in Germany had a shoulder height of around 1.2 m (3.9 ft) and a head-body length of 2.1 m (6.9 ft) without the tail, similar in size to large modern lions. The size of this male was exceeded by other specimens. This suggests, P. spelaea may have been over 10% larger than modern lions, but smaller than the earlier Panthera fossilis or the later American lion.[19] Like modern lions, females were smaller than males.[20]

In 2016, hair found near the Maly Anyuy River was identified as cave lion hair through DNA analysis. Comparison with hair of a modern lion revealed that cave lion hair was probably similar in colour as that of the modern lion, though slightly lighter. In addition, the cave lion is thought to have had a very thick and dense undercoat comprising closed and compressed yellowish-to-white wavy downy hair with a smaller mass of darker-coloured guard hairs, possibly an adaptation to the Ice Age climate.[21]

Distribution and habitatEdit

Cave lions and other Ice Age fauna in northern Spain, by Mauricio Antón

P. spelaea formed a contiguous population from Europe to Alaska over the Bering land bridge.[22] It was widely distributed from the Iberian peninsula, Southeast Europe, Great Britain, Central Europe, the East European Plain, and across most of northern Eurasia into Canada and Alaska. The oldest known fossils were excavated in northeastern Yakutia and were radiocarbon dated at 62,400 years old. The youngest known fossils are dated 11,925 years old and originated near Fairbanks, Alaska.[5]Phalanx bones excavated in Spain's La Garma cave complex were radiocarbon dated to 14,300–14,000 years old.[23] In Slovakia, skull, femur and pelvis remains were excavated in ten Karst caves in hilly and montane areas at elevations from 240 to 1,133 m (787 to 3,717 ft).[24] In Yakutia's Khayrgas Cave, bones of P. spelaea were found together with remains of humans, wolf, reindeer, Pleistocene horse and fish in a layer dated 13,200–21,500 years old.[25] In 2008, a well-preserved mature cave lion specimen was unearthed near the Maly Anyuy River in Chukotka Autonomous Okrug in Russia, which still retained some clumps of hair.[26]


P. spelaea was first described by the German paleontologist Georg August Goldfuss in 1810 under the scientific name Felis spelaea. The first fossil lion skull was excavated in a cave in southern Germany, research suggests that it may date to the Würm glaciation [27][28][5][2].

In 2015, two frozen cave lion cubs, estimated to be between 25,000 and 55,000 years old, were discovered close to the Uyandina River in Yakutia, Siberia in permafrost.[29][30][31] Research results indicate that the cubs were likely barely a week old at the time of their deaths, as their milk teeth had not fully erupted. Further evidence shows the cubs were hidden at a den site until they were old enough to join the pride. Researchers think that the cubs were trapped and killed by a landslide, and that without air, the cubs were preserved in such good condition. A second expedition to the site where the cubs were found was planned for 2016, in hopes of finding either the remains of a third cub or possibly the cubs' mother.[32]

In 2017, another frozen specimen, thought to be a lion cub, was found in Yakutia on the banks of the Tirekhtyakh River (Russian: Тирехтях), a tributary of the Indigirka River. Named 'Boris',[33] this cat was believed to be slightly older than the 2015 cubs at the time of its death; it is estimated to have been around one and a half to two months.[34] In 2018, another preserved carcass of a cub was found in a location 15 m (49 ft) away. Named 'Spartak', it was considered to be around a month old when it died approximately 50,000 years ago, and presumed to be a sibling to Boris.[33]


Cave lion with a reindeer. Painting by Heinrich Harder[35]

P. spelaea inhabited open environment like mammoth steppe and boreal forest. Large amounts of bones belonging to P. spelaea were excavated in caves, where bones of cave hyena, cave bear and Paleolithic artefacts were also found.[36][37] It was most likely a solitary hunter.[38]

Isotopic analyses of bone collagen samples extracted from fossils indicate that cave bear cubs, reindeer and other cervids were prominent in the diet of cave lions. Later cave lions seem to have preyed foremost on reindeer, up to the brink of local extinction or extirpation of both species.[39] Other possible prey species were giant deer, red deer, musk ox, bison, young woolly rhino and young mammoth.[5] It likely competed for prey with the European Ice Age leopard (P. pardus spelaea).[40]

See alsoEdit


  1. ^ Diedrich, C. G. (2014). "Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition about Prey". Paleontology Journal. 2014: 1–34. doi:10.1155/2014/106203.
  2. ^ a b c d Burger, J.; Rosendahl, W.; Loreille, O.; Hemmer, H.; Eriksson, T.; Götherström, A.; Hiller, J.; Collins, M. J.; Wess, T.; Alt, K. W. (2004). "Molecular phylogeny of the extinct cave lion Panthera leo spelaea". Molecular Phylogenetics and Evolution. 30 (3): 841–849. doi:10.1016/j.ympev.2003.07.020. PMID 15012963.
  3. ^ a b c d Barnett, R., Shapiro, B., Barnes, I. A. N., Ho, S. Y., Burger, J., Yamaguchi, N., Higham, T. F., Wheeler, H., Rosendahl, W., Sher, A. V., Sotnikova, M. (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. doi:10.1111/j.1365-294X.2009.04134.x. PMID 19302360. Archived from the original (PDF) on 21 January 2012.CS1 maint: uses authors parameter (link)
  4. ^ a b c Barnett, R.; Mendoza, M. L. Z.; Soares, A. E. R.; Ho, S. Y. W.; Zazula, G.; Yamaguchi, N.; Shapiro, B.; Kirillova, I. V.; Larson, G.; Gilbert, M. T. P. (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.
  5. ^ a b c d e f 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.CS1 maint: uses authors parameter (link)
  6. ^ Kurtén, B., 1968. Pleistocene Mammals of Europe. Weidenfeld and Nicolson, London.
  7. ^ Hemmer, H. (1974). "Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae) Teil 3. Zur Artgeschichte des Löwen Panthera (Panthera) leo (Linnaeus, 1758)". Veröffentlichungen der Zoologischen Staatssammlung 17: 167–280.
  8. ^ Turner, A. (1984). "Dental sex dimorphism in European lions (Panthera leo L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications". Annales Zoologici Fennici. 21: 1–8.
  9. ^ Spassov, N., Iliev, N. (1994). "Animal remains from the submerged Late Eneolithic – early Bronze Age settlements in Sozopol (South Bulgarian Black Sea Coast)". Proceedings of the International Symposium VI. Thracia Pontica. pp. 287–314.CS1 maint: multiple names: authors list (link)
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  11. ^ Christiansen, P. (2008). "Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters". Cladistics. 24 (6): 977–992. doi:10.1111/j.1096-0031.2008.00226.x.
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  14. ^ Chauvet, J.-M.; Brunel, D. E.; Hillaire, C. (1996). Dawn of Art: The Chauvet Cave. The oldest known paintings in the world. New York: Harry N. Abrams.
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