Panthera spelaea
Panthera spelaea, commonly known as the cave 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, with the genetic divergence between the two species estimated at around 500,000 years ago.
The earliest fossils of the P. spelaea lineage in Eurasia date to around 700,000 years ago. It is closely related and probably ancestral to the American lion. 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 along with other large carnivores like cave hyenas, which cave lions came into conflict with.
It closely resembled living lions with a coat of yellowish-grey fur, though depictions in cave art have led to suggestions that males lacked substantial manes unlike those present in living male lions. Whether or not cave lions lived in social groups like living lions is uncertain, but they are often suggested to have been largely solitary, similar to living tigers.
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.
Cave lions became extinct about 13,000 years ago as part of the end-Pleistocene extinction event, the precise cause of which is unknown, though climatic change, changes in prey abundance, and competition with other carnivores and humans have been suggested as possible causal factors.
Research history and taxonomy
In 1774, the 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. In 1810, a fossil skull from the cave was given the scientific name Felis spelaea by Georg August Goldfuss. It possibly dates to the Last Glacial Period.Several anatomical studies of remains of Panthera spelaea were conducted during the early-mid 19th century, who found the morphology of the species most similar to lions, tigers and jaguars. A monograph by W. Dawkins and W. Sandford studying P. spelaea published in 1868 found that it had closest affinities with the modern lion. At the end of the 19th century, the earliest remains of P. spelaea from Siberia were reported by M. Tschersky, who mistook them for those of a tiger. 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. However, other authors considered the cave lion to be more closely related to the tiger, based in part on a comparison of skull shapes, with some proposing that it should be considered a subspecies of the tiger as Panthera tigris spelaea. Analysis of cave lion mitochondrial genomes published in 2004 supported the modern lion as the closest relative of P. spelaea, with this result being later confirmed by analysis of the nuclear genome. Results from morphological studies showed that it is distinct in cranial and dental anatomy to justify the specific status of Panthera spelaea. Results of genetic studies also support this assessment.
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. 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. 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.
Evolution
Lion-like pantherine felids first appeared in the Tanzanian Olduvai Gorge about. 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 700,000-600,000 years ago, such as that from Pakefield in England, with possible older fossils from Western Siberia dating to the late Early Pleistocene, with a 2024 study suggesting a presence in Spain by 1 million years ago during the latest Early Pleistocene around the same age as the Siberian specimen. Different authors considered Panthera fossils as either a distinct species ancestral to P. spelaea, or as a subspecies of P. spelaea. 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.Cladogram after Tseng et al. :
The arrival of Panthera 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 preceding 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 fossilis the lion-sized sabertooth cat Homotherium and the "European jaguar" Panthera gombaszoegensis became much rarer, ultimately becoming extinct in Europe during the late Middle Pleistocene, with competition with lions suggested to be a likely important factor.
Specimens intermediate between P. fossilis and Late Pleistocene P. spelaea are referred to as the subspecies P. s. intermedia. 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. 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, 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 during the Illinoian.
Description
Carvings and cave paintings of cave lions, which were discovered in the Lascaux and Chauvet Caves in France, dating to the Upper Paleolithic. 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. Such cave paintings suggest that male cave lions completely lacked manes, or at most had very small manes.Early members of the cave lion lineage assigned to Panthera ''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, shoulder height of and body mass of, 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 and shoulder height of, 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, a body length of and shoulder height of respectively.
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. Like modern lions, females were smaller than males. Compared to the earlier P. fossilis, Late Pleistocene P. spelaea spelaea differs 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. The orbits of P. spelaea spelaea are also relatively larger and muzzle marginally narrower compared to P. 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.
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. While juveniles fur coat colour was yellowish, adult cave lions are suggested to have had grey fur.
Distribution and habitat
During the Last Glacial Period, P. spelaea formed a contiguous population across the mammoth steppe, from Western Europe to northwest North America. It was widely distributed in the Iberian Peninsula, Italian Peninsula, Southeast Europe, Great Britain, Central Europe, the East European Plain, the Ural Mountains, most of Northeast Asia, and across the Bering land bridge into Alaska, Yukon, and possibly Alberta. The cave lion had a wide elevation range, with finds extending up over above sea level in the European Alps and in Buryatia in Northern Asia, though they probably did not occupy mountainous habitats all-year round.The cave lion probably inhabited predominantly open habitats such as steppe and grasslands although it would have also occurred in open woodlands as well. While during the Last Glacial Period it was often associated with cold environments, the species also inhabited temperate environments, such as in Europe during the Last Interglacial/Eemian.