Panthera leo spelaea M. Boule & L. De Villeneuve, 1927
Panthera spelaea, also known as the cave lion or steppe lion,[1] is an
extinctPanthera species that most likely evolved in
Europe after the third
Cromerian interglacial stage, less than 600,000 years ago. Genetic analysis of
ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern
lion (Panthera leo) occurring in Africa and Asia,[2] with the
genetic divergence between the two species variously estimated between 1.9 million[3][4] and 600,000 years ago.[5] It is closely related and probably ancestral to the
American lion (Panthera atrox).[5] The species ranged from Western Europe to eastern
Beringia in North America, and was a prominent member of the
mammoth steppe fauna. It became extinct about 13,000 years ago.[6]
Several authors regarded Panthera spelaea as a
subspecies of the modern
lion, and therefore as Panthera leo spelaea.[9][10][11][2]
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.[12]
Results from morphological studies showed that it is distinct in cranial and dental
anatomy to justify the
specific status of Panthera spelaea.[13][14] Results of
phylogenetic studies also support this assessment.[15][3][4]
In 2001, the subspecies Panthera spelaea vereshchagini was proposed for seven specimens found in
Siberia and
Yukon, which have smaller skulls and teeth than the average P. spelaea.[16] Before 2020, 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][17] However, analysis of
mitochondrial genome sequences from 31 cave lions showed that they fall into two
monophyleticclades. One lived across western Europe and the other was restricted to Beringia during the
Pleistocene. For this reason, the Beringian population is considered a distinct subspecies, P. s. vereshchagini.[17][18]
Evolution
Lion-like
pantherine felids first appeared in the
TanzanianOlduvai Gorge about 1.7 to 1.2 million years ago. These cats dispersed to Europe from
East Africa in the first half of the Middle Pleistocene, giving rise to P. fossilis in
Central Europe by 610,000 years ago.[20]Panthera spelaea evolved from P. fossilis about 460,000 years ago in central Europe during the late
Saalian glaciation or early
Eemian and would have been common throughout
Eurasia from 450,000 to 14,000 years ago. Recent nuclear genomic evidence suggest that interbreeding between modern lions and all Eurasian fossil lions took place up until 500,000 years ago, but by 470,000 years ago, no subsequent interbreeding between the two lineages occurred.[2][20][5]
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.[21] 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.[6][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]
Carvings and
cave paintings of cave lions, which were discovered in the
Lascaux and
Chauvet Caves in France, were dated to 15,000 to 17,000 years old.[19][22] A drawing in the Chauvet cave 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.[23] Such cave paintings suggest that male cave lions completely lacked manes, or at most had very small manes.[6]
P. spelaea was thought to have been one of the largest lion species. The skeleton of an adult male found in 1985 near
Siegsdorf in Germany had a shoulder height of around 1.2 m (3 ft 11 in) and a head-body length of 2 m (6 ft 7 in) without the tail, similar in size to large modern lions.[26] The size of this male was exceeded by other specimens, with another male reaching 2.5 m (8 ft 2 in) long without the tail.[citation needed] Similarly, footprints attributed to a male cave lion measured 15 cm (6 in) across. The heaviest Panthera spelaea was estimated to weigh 339 kg (747 lb).[27]
This shows that P. spelaea would have been up to or over 12% larger than modern lions, but still smaller than the earlier Panthera fossilis or the
American lion (P. atrox).
Cave paintings almost exclusively show hunting animals without a mane, suggesting that males were indeed maneless.[28][29]P. spelaea had a relatively longer and narrower muzzle compared to that of the extant lion. Despite this, the two species do not exhibit major differences in morphology.[6] Like modern lions, females were smaller than males.[30]
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.[31]
The oldest known fossils dated with
radiocarbon dating were excavated in northeastern Russia and were radiocarbon dated at 62,400 years old.[6]Phalanx bones excavated in Spain's
La Garma cave complex were radiocarbon dated to 14,300–14,000 years old.[40] The youngest known fossils are dated 11,925 years old and originated near
Fairbanks, Alaska.[6] In Western Europe, however, P. spelaea may have survived into the Early Holocene.[41]
In 2008, a well-preserved mature cave lion specimen was unearthed near the
Maly Anyuy River in
Chukotka Autonomous Okrug, which still retained some clumps of hair.[42] In the
Sakha Republic'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.[43]
The cave lion probably inhabited predominantly open habitats such as
steppe and grasslands although it would have also have occurred in open woodlands as well.[6] It also lived in mountainous regions, likely because it sought out hibernating bears in montane caves as a food source during the winter.[44]
Discoveries
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.[45][46][47]
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 suggests the cubs were hidden at a den site until they were strong enough to follow their mother back to the pride, as with modern lions. Researchers believe that the cubs were trapped and killed by a landslide, and that the absence of oxygen underground hindered their decomposition and allowed the cubs to be 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.[48]
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. This male cub was thought 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.[49] In 2018, another preserved carcass of a cub was found in a location 15 m (50 ft) away. It was considered to be around a month old when it died approximately 50,000 years ago, and presumed to be a sibling of the male cub.[50] However,
carbon dating showed them to have lived about 15,000 years apart, with the female estimated to have lived 28,000 years ago, and the male 43,448 years ago.[18] Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the
Ice age.[51]
Paleobiology
P. spelaea inhabited colder environments such as
mammoth steppe and
boreal forest. It was one of the
keystone species of the mammoth steppe, being one of the main
apex predators alongside the
gray wolf,
cave hyena and
brown bear.[53] 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.[54][55] Some of these accumulations of cave lion bones have been attributed to hoarding of meat from cave lion carcasses by cave hyenas in caves occupied by the latter.[56] It is unclear whether P. spelaea was social like the modern lion; some evidence indicates that it may have been solitary.[53]
The extinction of the Eurasian cave lion was multifaceted, being attributable to a combination of human activity, the decline of the mammoth steppe, and changes in the herbivore guild.[60] There is direct evidence of human hunting of cave lions and exploitation of their pelts for ritualistic purposes during the
Magdalenian, which was likely an important factor contributing to their ultimate demise.[40]
^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.
^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.{{
cite book}}: CS1 maint: location missing publisher (
link)
^
abSotnikova, 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.
PMID24382145.
^Marciszak, A. & Stefaniak, K. (2010). "Two forms of cave lion: Middle Pleistocene Panthera spelaea fossilis Reichenau, 1906 and Upper Pleistocene Panthera spelaea spelaea Goldfuss, 1810 from the Bisnik Cave, Poland". Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 258 (3): 339–351.
doi:
10.1127/0077-7749/2010/0117.
^Yamaguchi, N.; Cooper, A.; Werdelin, L.; MacDonald, D. W. (2004). "Evolution of the mane and group-living in the lion (Panthera leo): a review". Journal of Zoology. 263 (4): 329–342.
doi:
10.1017/S0952836904005242.
^Guthrie, R. D. (2005). The Nature of Paleolithic Art. Chicago: University of Chicago Press.
^Hussain, S.T.; Floss, H. (2015). "Sharing the world with mammoths, cave lions and other beings: linking animal-human interactions and the Aurignacian "belief world"". Quartär. 65: 85–120.
doi:
10.7485/QU62_4.
^Chernova, O. F.; Kirillova, I. V.; Shapiro, B.; Shidlovskiy, F. K.; Soares, A. E. R.; Levchenko, V. A.; Bertuch, F. (2016). "Morphological and genetic identification and isotopic study of the hair of a cave lion (Panthera spelaea Goldfuss, 1810) from the Malyi Anyui River (Chukotka, Russia)". Quaternary Science Reviews. 142: 61–73.
Bibcode:
2016QSRv..142...61C.
doi:
10.1016/j.quascirev.2016.04.018.
^Diedrich, C. G. (2011). "The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe". Historical Biology. 23 (2–3): 271–311.
Bibcode:
2011HBio...23..271D.
doi:
10.1080/08912963.2010.546529.
S2CID86638786.
^Diedrich, C. G. (2013). "Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art". Quaternary Science Reviews. 76: 167–193.
Bibcode:
2013QSRv...76..167D.
doi:
10.1016/j.quascirev.2013.05.009.