Panthera spelaea

Panthera spelaea Temporal range: Middle-Late Pleistocene, 0.6–0.013  Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Skeleton in Natural History Museum, Vienna
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Suborder:	Feliformia
Family:	Felidae
Subfamily:	Pantherinae
Genus:	Panthera
Species:	†P. spelaea
Binomial name
†Panthera spelaea Goldfuss, 1810
Subspecies
Panthera spelaea vereshchagini(Baryshnikov & Boeskorov, 2001) Panthera spelaea spelaeaGoldfuss, 1810
Red indicates the maximal range of Panthera spelaea, blue Panthera atrox , and green Panthera leo .
Synonyms
Panthera leo spelaeaM. Boule & L. De Villeneuve, 1927

Panthera spelaea, commonly known as the cave lion (or less commonly as the steppe lion), is an extinct Panthera species that was native to Eurasia and northwest North America during the Pleistocene epoch. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion (Panthera leo), ^[1] with the genetic divergence between the two species estimated at around 500,000 years ago. ^[2] The earliest fossils of the P. spelaea lineage (either regarded as the separate species Panthera fossilis or the subspecies P. spelaea fossilis) in Eurasia date to around 700,000 years ago (with possible late Early Pleistocene records). ^[3] It is closely related and probably ancestral to the American lion (Panthera atrox). ^[2] The species ranged from Western Europe to eastern Beringia in North America, and was a prominent member of the mammoth steppe fauna, and an important apex predator across its range. It became extinct about 13,000 years ago. ^[4] It closely resembled living lions with a coat of yellowish-grey fur though unlike extant lions, males appear to have lacked manes.

Panthera spelaea interacted with both Neanderthals and modern humans, who used their pelts and in the case of the latter, depicted them in artistic works.

Taxonomy

Cave lion skull exhibited in the Muséum de Toulouse, France

In 1774, Zoolithenhöhle cave near the village of Burggaillenreuth in Bavaria, southern Germany was brought to scientific attention by Johan Friedrich Esper, who realised that the bones of extinct animals were present in the cave. ^[5] In 1810, a fossil skull from the cave was given the scientific name Felis spelaea by Georg August Goldfuss. ^[6] It possibly dates to the Last Glacial Period. ^{[7] [1] [4]}

Several anatomical studies of remains of Panthera spelaea were conducted during the early-mid 19th century, who generally agreed that the species had lion affinities. ^[4] During the 20th century and the first decade of the 21st century, Panthera spelaea was often regarded as a subspecies of the modern lion, and therefore as Panthera leo spelaea. ^{[8] [9] [10] [1]} 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. ^[11]

Results from morphological studies showed that it is distinct in cranial and dental anatomy to justify the specific status of Panthera spelaea. ^{[12] [13]} Results of phylogenetic studies also support this assessment. ^{[14] [15] [16]}

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. ^[17] 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. ^{[15] [18]} However, analysis of mitochondrial genome sequences from 31 cave lions showed that they fall into two monophyletic clades. 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. ^{[18] [19]}

Evolution

Lion-like pantherine felids first appeared in the Tanzanian Olduvai Gorge about 1.7 to 1.2 million years ago. These cats dispersed into Eurasia from East Africa around the end of the Early Pleistocene and the beginning of the Middle Pleistocene, giving rise to Panthera fossilis. The oldest widely accepted fossils of P. fossilis in Europe date to around 700,000 years ago, such as that from Pakefield in England, ^{[20] [21] [3] [22]} with possible older fossils from Western Siberia dating to the late Early Pleistocene, ^[23] with a 2024 study suggesting a presence in Spain by 1 million years ago. ^[24] Different authors considered Panthera fossils as either a distinct species ancestral to P. spelaea, ^[25] or as a subspecies of P. spelaea. ^{[22] [26]} 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. ^{[20] [1] [2]}

The following cladogram shows the genetic relationship between P. spelaea and other pantherine cats. ^[16]

	Felis catus Pantherinae Neofelis nebulosa (clouded leopard) Panthera Panthera tigris (tiger) Panthera onca (jaguar) Panthera uncia (snow leopard) Panthera pardus (leopard) Panthera leo Panthera leo leo (Asian, North and West African lions) Panthera leo melanochaita (Southern and East African lions) †Panthera spelaea

The arrival of Panthera (spelaea) fossilis in Europe was part of a faunal turnover event around the Early-Middle Pleistocene transition in which many of the species that characterised the preceeding late Villafranchian became extinct. In the carnivore guild, this notably included the giant hyena Pachycrocuta and the sabertooth cat Megantereon . Following the arrival of Panthera (spelaea) fossilis the lion-sized sabertooth cat Homotherium and the "European jaguar" Panthera gombaszoegensis became much rarer, ^[24] ultimately becoming extinct in Europe during the late Middle Pleistocene, with competition with lions suggested to be a likely important factor. ^{[27] [28]}

Specimens intermediate between P. fossilis and Late Pleistocene P. spelaea are referred to as the subspecies P. s. intermedia. ^[21] The transition from P. fossilis to Late Pleistocene P. spelaea shows significant reduction in body size, as well as changes in skull and tooth morphology. ^[29] Mitochondrial DNA sequence data from fossil lion remains show that the American lion represents a sister group of Late Pleistocene P. spelaea, and likely arose when an early P. spelaea population became isolated south of the Cordilleran Ice Sheet. Initially this was suggested to be around 340,000 years ago, ^[15] but later studies suggested that the split between the two species was probably younger, around 165,000 years ago, consistent with the late first appearance of P. spelaea in Eastern Beringia (now Alaska and adjacent regions) during the Illinoian (around 190-130,000 years ago). ^[30]

Characteristics

Cave lions and bison depicted in the Chauvet Cave, France

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. ^{[31] [32]} 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. ^[33] Such cave paintings suggest that male cave lions completely lacked manes, or at most had very small manes. ^[4]

Size comparison of various specimens attributed to the P. spelaea/fossilis lineage. 1 =P. (spelaea) fossilis male (M) from Château, France MIS (Marine Isotope Stage) 15 ~ 600,000 years ago 2= P. spelaea from Niedźwiedzia Cave, Poland, MIS 5 (130-80,000 years ago). 3 = M from Siegsdorf, Germany MIS 3 (57-29,000 years ago) 4= female (F) from Zandobbio, Italy, Last Interglacial (130-115,000 years ago) 5= F from Kryshtaleva Cave, Ukraine (21-22,000 years ago). Scale bar = 10 cm

Size comparison of Middle Pleistocene Panthera (spelaea) fossilis and Late Pleistocene Panthera spelaea compared to a human

Early members of the cave lion lineage assigned to Panthera (spelaea) fossilis during the Middle Pleistocene were considerably larger than individuals of P. spelaea from the Last Glacial Period and modern lions, with some of these individuals having an estimated length of 2.5–2.9 metres (8.2–9.5 ft), shoulder height of 1.4–1.5 metres (4.6–4.9 ft) and body mass of 400–500 kilograms (880–1,100 lb), respectively, making them among the largest cats to have ever lived. The Late Pleistocene Panthera spelaea spelaea was noticeably smaller though still large relative to living cats, with an estimated length of 2–2.1 metres (6.6–6.9 ft) and shoulder height of 1.1–1.2 metres (3.6–3.9 ft), respectively, The species showed a progressive size reduction over the course of the Last Glacial Period up until its extinction, with the last P. spelaea populations comparable in size to small-sized modern lions, with a body mass of only 70–90 kilograms (150–200 lb), a body length of 1.2–1.3 metres (3.9–4.3 ft) and shoulder height of 70–75 centimetres (2.30–2.46 ft) respectively. ^{[26] [34]}

P. spelaea had a relatively longer and narrower muzzle compared to that of the extant lion, with the zygomatic region being strongly arched, with the carnassial teeth having differences in cusp morphology (displaying preparastyles). ^[4] Like modern lions, females were smaller than males. ^[35] Compared to the earlier P. (spelaea) fossilis, Late Pleistocene P. spelaea spelaea differs (in addition to previously mentioned size differences) in having larger incisor teeth, more narrow and flattened canines, as well narrower upper and lower third and fourth premolars, which display some differences in cusp morphology, with the lower first molar being narrower and more elongate. ^[26] The orbits (eye sockets) of P. spelaea spelaea are also relatively larger and muzzle marginally narrower compared to P. (spelaea) fossilis, with the nasal region also being proportionally narrower, while the postorbital and mastoid regions of the skull are wider, with the tympanic bullae being more inflated. ^[36]

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. ^[37] While juveniles fur coat colour was yellowish, adult cave lions are suggested to have had grey fur. ^[19]

Distribution and habitat

Cave lions feeding on reindeer in a mammoth steppe landscape in northern Spain also including woolly rhinoceros, woolly mammoth and wild horse. Art by Mauricio Antón

During the Last Glacial Period, P. spelaea formed a contiguous population across the mammoth steppe, from Western Europe to northwest North America. ^{[4] [38]} It was widely distributed in the Iberian Peninsula, ^[39] Italian Peninsula, ^[40] Southeast Europe, ^[41] Great Britain, ^[4] Central Europe, ^{[42] [43] [44]} the East European Plain, ^[4] the Ural Mountains, ^[45] most of Northeast Asia (ranging as far south as Northeast China ^[46] and possibly the Korean peninsula ^[47] ), and across the Bering land bridge into Alaska and Yukon. ^[4] The cave lion had a wide elevation range, with finds extending up over 2,000 metres (6,600 ft) above sea level in the European Alps and in Buryatia in Northern Asia, though they probably did not occupy mountainous habitats all-year round. ^[48] 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. ^[4] While during the Last Glacial Period it was often associated with cold environments, the species also inhabited temperate environments, ^[49] such as in Europe during the Last Interglacial/Eemian. ^[50]

Paleobiology

Ecology

A cave lion with a reindeer, painting by Heinrich Harder

P. spelaea 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. ^[52] 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. ^{[53] [54]} Despite their common name, "cave lions" probably only infrequently if ever used caves, and were present in regions where caves were absent. ^[55] Some of these accumulations of cave lion bones in cave hyena dens have been attributed to confrontations between cave hyenas and cave lions over carcasses, with the remains of cave lions killed in these confrontations subsequently transported to the dens. ^{[56] [55]}

Isotopic analyses of bone collagen samples extracted from remains in Europe ^{[57] [52]} and East Beringia ^[58] indicate that reindeer were particularly prominent in the diet of cave lions in these regions during the Last Glacial Period. ^[52] Cave lions also seem to have opportunistically preyed on the cubs of cave bears. ^{[52] [57]} Isotopic analysis of other European specimens suggests a diet including wild horse, woolly mammoth and cave bears for these individuals. ^[52] It may have sought out hibernating bears in montane caves as a food source during the winter. ^[59] Bite marks found on the bones of straight-tusked elephants in Neumark Nord, Germany, dating to the Last Interglacial, have been suggested to be the result of scavenging by cave lions. ^[60]

The mummy of Blue Babe, a 36,000 year old male Steppe Bison found in Alaska with claw marks on its rear and tooth punctures in its skin, which indicate that the bison was killed by a cave lion.

Other possible prey species were giant deer, red deer, muskox, aurochs, wisent, steppe bison, and young woolly rhinoceros. It likely competed for prey with the European leopards, cave hyenas, brown bears and grey wolves in Eurasia, ^[62] along with short-faced bears, Homotherium , and Beringian wolves in Beringia. ^[52]

Social behavior

Whether or not cave lions existed in prides like modern lions is unclear. ^[19] Isotopic analysis on cave lions by Hervé Bocherens and colleagues lead them to suggest that cave lions may have been solitary, due to cave lions shifting their diets after the disappearance of cave hyenas, carcasses being consumed the cave hyenas as well, suggests they were at a competitive disadvantage, and the scattering of isotopic data between individuals. ^{[57] [52] [63]} Some other authors have also argued that the absence of manes in cave lions suggests that cave lions did not live in prides, given the importance of manes in the social hierarchy of modern lions. ^[19] Boeskorov et al. 2021 suggested both European and Beringian cave lions may have hunted in larger prides than modern lions because sexual dimorphism in cave lions was more pronounced than in modern African lions and solitary big cats. However, they admitted the data is insufficient to come down to a certain conclusion. ^[19]

Cave lion cubs appear to have lived in dens during their earliest stages of life, like modern lion cubs and were likely solely raised by females, like living Panthera species. ^[19]

Relationship with humans

Drawing of a now-lost Palaeolithic sculpture of a cave lion from Isturitz, France

Drawing of cave lions in the Chamber of Felines, Lascaux caves

Carving of a Panthera spelaea head from Vogelherd Cave, dating to around 35–40,000 years ago

Anthropomorphic Lion-man (Löwenmensch) figurine, around 35-41,000 years old

Cave lions were hunted and their pelts exploited in Europe by Neanderthals during the Middle Paleolithic, ^[64] and during the Upper Paleolithic by modern humans in Spain as evidenced in the La Garma site dating to the Magdalenian. ^[65] Modern humans also drew cave paintings of cave lions, engraved their likeness on bones and created sculptures of them, including the famous anthropomorphic lion-man figure from Hohlenstein-Stadel cave in Germany dating to around 41-35,000 years ago with the body of a human and the head of a lion. Cave lion canines with perforated holes may have been worn as personal ornaments. ^[64] Decorated stones with engravings representing cave lions have been found in southern Italy. ^[66]

Extinction

See also: Late Pleistocene extinctions

Radiocarbon dating suggests that the species went extinct approximately simultaneously across its range during the last few thousand years of the Late Pleistocene, around 14-15,000 years ago, possibly surviving around 1000 years later in the far east North American portion of its range. This timing roughly corresponds to the onset of the Bølling–Allerød Interstadial warm period and the consequent collapse of the mammoth steppe ecosystem. The precise cause of its extinction is unclear, but possibly involved environmental change from open habitats to closed forests, changes in prey abundance, as well as human impact, though it is difficult to distentangle the precise causes of its extinction. ^[4] Cave lions appear to have undergone a population bottleneck that considerably reduced their genetic diversity between 47,000 and 18,000 years ago, probably driven at least in part by climatic instability. ^[67]

Mummified specimens

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. ^[68]

"Sparta", a 28,000 year old mummified female cave lion cub from the banks of the Semyuelyakh River in Siberia.

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. ^{[69] [70] [71]} 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. ^[72]

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. ^[73] 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. ^[74] 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. ^[19] Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the Ice age. ^[75]

See also

• History of lions in Europe
• Megafauna
• Panthera atrox
• Panthera blytheae
• Panthera gombaszoegensis
• Panthera palaeosinensis
• Panthera shawi
• Panthera youngi
• Panthera zdanskyi
• Panthera leo sinhaleyus
• Panthera leo fossilis

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External links

• Prehistoric cats and prehistoric cat-like creatures, from the Messybeast Cat Resource Archive
• The mammoth and the flood, volume 5, chapter 1, by Hans Krause.
• Hoyle and cavetigers Archived 13 January 2016 at the Wayback Machine , from the Dinosaur Mailing List. (Groiss)
• Photo-reconstruction of Panthera spelaea, by paleoartist Roman Uchytel