Panthera spelaea

, France 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. Analysis of cave lion mitochondrial genomes published in 2004 supported the modern lion as the closest relative of P. 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. Mitochondrial genomic evidence suggests P. spelaea and modern lions split off around 1.85 Ma. }}}}}}}}}}}}}}}} 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 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 (spelaea) fossilis the lion-sized sabertooth cat Homotherium and the "European jaguar" Panthera gombaszoegensis became much rarer, Specimens intermediate between P. fossilis and Late Pleistocene P. spelaea are referred to as the subspecies P. s. intermedia. 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, ==Description==

depicted in the Chauvet Cave, France 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. 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). 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. 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==

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. It was widely distributed in the Iberian Peninsula, Italian Peninsula, Southeast Europe, Great Britain, the East European Plain, most of Northeast Asia (ranging as far south as Northeast China, Japan, and possibly the Korean peninsula), and across the Bering land bridge into Alaska, Yukon, 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. such as in Europe during the Last Interglacial/Eemian. ==Paleobiology==

Ecology 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. 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. Despite their common name, "cave lions" probably only infrequently if ever used caves, and were present in regions where caves were absent. Analysis on the ecology of cave lions found that they may have been diurnal predators. 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.Isotopic analyses of bone collagen samples extracted from remains in Europe indicate that reindeer were particularly prominent in the diet of cave lions in these regions during the Last Glacial Period. 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. Other possible prey species were giant deer, red deer, muskox, aurochs, wisent, steppe bison, Competitors It likely competed for prey with the European leopards, cave hyenas, brown bears and grey wolves in Eurasia, along with short-faced bears, the sabertooth cat Homotherium, and Beringian wolves in Beringia. Some caves, such as Zoolithen cave, had a high concentration of males cave lions. This suggests much like modern male lions, male cave lions may have ventured into caves and actively killed cave hyenas. Isotopic analysis suggests cave hyenas and cave lions may have hunted different prey species due to competitive exclusion. In contrast to tigers, cave lions coexisted with large, robust wolves, which would have been serious competitors, After MIS 3, with the appearance of smaller and even dwarf individuals made cave lions increasingly vulnerable to attacks from wolves, which became most notable during MIS 2. 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. Within mountain ranges and higher altitude environments, cave lions may have also been solitary or hunt in mating pairs, much like today's lions. Lions tend to hunt in prides in altitudes below 1,500 meters, but within higher altitudes, they tend to be solitary or hunt in mating pairs. This proposed behavior for cave lions is further supported by the ratios of males and females from Moravian Karst being 1:1. == Relationship with humans ==

Both Neanderthals and modern humans interacted with cave lions. Findings of cave lion phalanges (hand bones) in archaeological layers from Einhornhöhle in Germany, dating to at least 190,000 years ago, are suggested to represent remains of a skinned cave lion pelt that had been transported to the site by Neanderthals. Cave lion bones at Scladina Cave in Belgium, dating to around 130,000 years ago, were fashioned by Neanderthals into tools that were used to retouch stone artifacts. At Siegsdorf in Germany, a cave lion skeleton dating to around 48,000 years ago has a puncture wound on the underside of a rib, suggested to have been produced when the cave lion was killed by a wooden spear thrust into its chest, again probably by Neanderthals, with cut marks found on other bones suggesting that it was subsequently butchered. Like Neanderthals, modern humans also utilized their pelts, as evidenced by phalanges found at the La Garma site in Spain dating to the Magdalenian, around 16,800 years ago. Some bones of cave lions found in Upper Paleolithic layers such as the Aurignacian and Gravettian of the Swabian Jura in southern Germany display cut marks and modification into tools. This may represent evidence of hunting, though evidence of hunting cave lions by modern humans during the Upper Paleolithic is uncommon. == Extinction ==

Radiocarbon dating suggests that the species went extinct approximately simultaneously across its range during the last few thousand years of the Late Pleistocene, around 14,000-15,000 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 distantangle the precise causes of its extinction. The extinction formed part of the broader end-Pleistocene extinction event, in which most large terrestrial mammals globally became extinct, including many large carnivores. == 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. 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. 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. In 2017, another frozen specimen, thought to be a lion cub, was found in Yakutia on the banks of the Tirekhtyakh River (), 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. In 2018, another preserved carcass of a cub was found in a location 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. 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. == Notes ==