Panthera spelaea
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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),<ref name="Burger">Template:Cite journal</ref> with the genetic divergence between the two species estimated at around 500,000 years ago.<ref name="Manuel_al2020">Template:Cite journal</ref>
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).<ref name=":2">Template:Cite journal</ref> It is closely related and probably ancestral to the American lion (Panthera atrox).<ref name="Manuel_al2020" /> 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 unlike extant lions, males appear to have lacked manes. Whether or not cave lions lived in social groups like living lions is uncertain, but they are frequently suggested to have been largely solitary, similar to living tigers.<ref name=":7">Template:Cite journal</ref>
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, 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.<ref name="Stuart2011">Template:Cite journal</ref>
Research history and taxonomyEdit
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.<ref name=":6">Template:Cite journal</ref> In 1810, a fossil skull from the cave was given the scientific name Felis spelaea by Georg August Goldfuss.<ref>Template:Cite book</ref> It possibly dates to the Last Glacial Period.<ref>Template:Cite journal</ref><ref name=Burger /><ref name=Stuart2011 />
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.<ref name=":6" /> 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.<ref>Template:Cite book</ref><ref name=Hemmer>Template:Cite journal</ref><ref>Template:Cite journal</ref><ref name=Burger/> However, other authors considered the cave lion to be more closely related to the tiger,<ref name=":6" /> 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.<ref>Template:Cite journal</ref> Analysis of cave lion mitochondrial genomes published in 2004 supported the modern lion as the closest relative of P. spelaea,<ref name="Burger" /> with this result being later confirmed by analysis of the nuclear genome.<ref name="Manuel_al2020" /> Results from morphological studies showed that it is distinct in cranial and dental anatomy to justify the specific status of Panthera spelaea.<ref>Template:Cite book</ref><ref>Template:Cite journal</ref> Results of genetic studies also support this assessment.<ref>Template:Cite journal</ref><ref name="Barnett2009">Template:Cite journal</ref><ref name="Barnett2016">Template:Cite journal</ref>
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.<ref>Template:Cite journal</ref> 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.<ref name=Barnett2009/><ref name=Stanton2020>Template:Cite journal</ref> 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.<ref name=Stanton2020 /><ref name="Boeskorov_al.2021">Template:Cite journal</ref>
EvolutionEdit
Lion-like pantherine felids first appeared in the Tanzanian Olduvai Gorge about Template:Mya. 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,<ref name=Sotnikova2014>Template:Cite journal</ref><ref name=":0">Template:Cite journal</ref><ref name=":2" /><ref name="Ianucci_al.2024">Template:Cite journal</ref> with possible older fossils from Western Siberia dating to the late Early Pleistocene,<ref>Template:Cite journal</ref> 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.<ref name=":3">Template:Cite journal</ref> Different authors considered Panthera fossils as either a distinct species ancestral to P. spelaea,<ref name=":02">Template:Cite journal</ref> or as a subspecies of P. spelaea.<ref name="Ianucci_al.2024" /><ref name=":1">Template:Cite journal</ref> 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.<ref name=Sotnikova2014/><ref name=Burger/><ref name=Manuel_al2020/>
The following cladogram shows the genetic relationship between P. spelaea and other pantherine cats.<ref name="Barnett2016" />Template:CladeThe 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,<ref name=":3" /> ultimately becoming extinct in Europe during the late Middle Pleistocene, with competition with lions suggested to be a likely important factor.<ref>Template:Cite journal</ref><ref name="Marciszak2022">Template:Cite journal</ref>
Specimens intermediate between P. fossilis and Late Pleistocene P. spelaea are referred to as the subspecies P. s. intermedia.<ref name=":0" /> 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.<ref>Template:Cite journal</ref> 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,<ref name="Barnett2009" /> 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).<ref>Template:Cite journal</ref>
CharacteristicsEdit
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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.<ref name=Chauvet1996 /><ref>Template:Cite book</ref> 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.<ref name=Yamaguchietal2004>Template:Cite journal</ref> Such cave paintings suggest that male cave lions completely lacked manes, or at most had very small manes.<ref name=Stuart2011/>
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 Template:Convert, shoulder height of Template:Convert and body mass of Template:Convert, 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 Template:Convert and shoulder height of Template:Convert, 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 Template:Convert, a body length of Template:Convert and shoulder height of Template:Convert respectively.<ref name=":1" /><ref name=":5">Template:Cite journal</ref>
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).<ref name="Stuart2011" /> Like modern lions, females were smaller than males.<ref>Template:Cite journal</ref> 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.<ref name=":1" /> 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.<ref>SABOL, Martin. Panthera fossilis (REICHENAU, 1906) (Felidae, Carnivora) from Za Hájovnou Cave (Moravia, the Czech Republic): a fossil record from 1987–2007. Fossil Imprint / Acta Musei Nationalis Pragae, Series B – Historia Naturalis Prague: National Museum, 2014, 70(1-2), 59-70. ISSN 2533-4050 (tisk), 2533-4069 (online).</ref>
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.<ref>Template:Cite journal</ref> While juveniles fur coat colour was yellowish, adult cave lions are suggested to have had grey fur.<ref name="Boeskorov_al.2021" />
Distribution and habitatEdit
During the Last Glacial Period, P. spelaea formed a contiguous population across the mammoth steppe, from Western Europe to northwest North America.<ref name="Stuart2011" /><ref>Template:Cite journal</ref> It was widely distributed in the Iberian Peninsula,<ref>Template:Cite journal</ref> Italian Peninsula,<ref>Iannucci, Alessio, Mecozzi, Beniamino, Pineda, Antonio, Sardella, Raffaele, Carpentieri, Marco, Rabinovich, Rivka, and Moncel Marie-Helene. "Early Occurrence of lion (Panthera spelaea) at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy). Journal of Quarternary Sciences 39:3 (2024): 4.</ref> Southeast Europe,<ref>Template:Cite journal</ref> Great Britain,<ref name="Stuart2011" /> Central Europe,<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> the East European Plain,<ref name="Stuart2011" /> the Ural Mountains,<ref>Template:Cite journal</ref> most of Northeast Asia (ranging as far south as Northeast China<ref>Template:Cite journal</ref> and possibly the Korean peninsula<ref>Template:Cite journal</ref>), and across the Bering land bridge into Alaska and Yukon.<ref name="Stuart2011" /> The cave lion had a wide elevation range, with finds extending up over Template:Convert above sea level in the European Alps and in Buryatia in Northern Asia, though they probably did not occupy mountainous habitats all-year round.<ref>Template:Cite journal</ref>
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.<ref name="Stuart2011" /> While during the Last Glacial Period it was often associated with cold environments, the species also inhabited temperate environments,<ref>Template:Cite journal</ref> such as in Europe during the Last Interglacial/Eemian.<ref>Template:Cite journal</ref>
PaleobiologyEdit
EcologyEdit
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.<ref name="Bocherens2015">Template:Cite journal</ref> 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.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> Despite their common name, "cave lions" probably only infrequently if ever used caves, and were present in regions where caves were absent.<ref name="diedrich&zak">Diedrich, C.G. & ŽÁK, K. 2006. Prey deposits and den sites of the Upper Pleistocene hyena Crocuta crocuta spelaea (Goldfuss, 1823) in horizontal and vertical caves of the Bohemian Karst (Czech Republic). Bulletin of Geosciences 81(4), 237–276 (25 figures). Czech Geological Survey, Prague. ISSN 1214-1119.</ref> 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.<ref>Template:Cite journal</ref><ref name="diedrich&zak" />
Isotopic analyses of bone collagen samples extracted from remains in Europe<ref name="Bocherens">Template:Cite journal</ref><ref name="Bocherens2015" /> and East Beringia<ref>Template:Cite journal</ref> indicate that reindeer were particularly prominent in the diet of cave lions in these regions during the Last Glacial Period.<ref name="Bocherens2015" /> Cave lions also seem to have opportunistically preyed on the cubs of cave bears,<ref name="Bocherens2015" /><ref name="Bocherens"/> with adult cave bears also likely being targeted at least on occasion.<ref name=":5" /> Isotopic analysis of other European specimens suggests a diet including wild horse, woolly mammoth and cave bears for these individuals.<ref name="Bocherens2015" /> It may have sought out hibernating bears in montane caves as a food source during the winter.<ref name=":4">Template:Cite journal</ref> 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.<ref>Template:Cite journal</ref>
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,<ref name="Diedrich 2013">Template:Cite journal</ref> along with short-faced bears, the sabertooth cat Homotherium, and Beringian wolves in Beringia.<ref name="Bocherens2015" />
Social behaviorEdit
Whether or not cave lions were gregarious like modern lions is unclear.<ref name="Boeskorov_al.2021" /> Sabol and Puzachenko in their 2024 paper stated that the cave lions are "generally thought" to have been solitary, but that "opinions on this vary" and that the question remained open.<ref name=":7" /> Some experts, such as Guthrie argue that the cave lion lived solitarily or only in small family groups, based primarily on the lack of manes (which are highly important to the social hierarchy of modern lions) depicted on Paleolithic cave art.<ref>Guthrie, R.D. Frozen Fauna of the Mammoth Steppe. The Story of Blue Babe; University of Chicago Press: Chicago, IL, USA, 1990; pp. 1–323. </ref><ref name="Boeskorov_al.2021" /> This argument was criticized by Yamaguichi and colleagues, they argued that the emergence of group living in lions probably predates the split between cave lions and modern lions and the evolution of manes in modern lions. On the other hand, authors argued gregariousness via sexual dimorphism. Sexual dimorphism within canine dimensions suggests cave lions had similar sexual dimorphism to lions, and in evolutionary context may suggests they lived in groups according to Yamaguichi and colleagues.<ref name="Yamaguchietal2004" /> Boeskorov and their colleagues suggested that the more extreme sexual dimorphism seen in cave lions could suggest that they lived in larger prides. However they admitted that there isn’t enough evidence to conclude on their social structure.<ref name="Boeskorov_al.2021" /> On the contrary, Valkenburgh and Sacco warned that sexual dimorphism isn’t a reliable way to determine group behavior, as leopards are also very sexually dimorphic in canine dimensions but live a solitary lifestyle.<ref name="Van Valenburgh">Template:Cite journal</ref>
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.<ref name="Bocherens" /><ref name="Bocherens2015" /><ref name="Bocherens and Kirillova 2024">Template:Cite journal</ref> 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.<ref name=":4" /><ref>Template:Cite journal</ref>
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.<ref name="Boeskorov_al.2021" />
Relationship with humansEdit
Template:Multiple image Cave lions were hunted and their pelts exploited in Europe by Neanderthals during the Middle Paleolithic,<ref name=Russo>Template:Cite journal</ref> and during the Upper Paleolithic by modern humans in Spain as evidenced in the La Garma site dating to the Magdalenian.<ref name=UnderTheSkinOfLion>Template:Cite journal</ref> 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 (Löwenmensch) 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.<ref name=Russo/> Decorated stones with engravings representing cave lions have been found in southern Italy.<ref>Template:Cite journal</ref>
ExtinctionEdit
Template:See also 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.<ref name="Stuart2011" /> Competition with wolves may have been an important factor in its extinction.<ref name=":5" /> 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.<ref>Template:Cite journal</ref>
Mummified specimensEdit
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.<ref>Template:Cite journal</ref>
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.<ref>Template:Cite news</ref><ref>Template:Cite news</ref><ref>Template:Cite news</ref> 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.<ref>Template:Cite news</ref>
In 2017, another frozen specimen, thought to be a lion cub, was found in Yakutia on the banks of the Tirekhtyakh River (Template:Langx), 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.<ref>Template:Cite news</ref> In 2018, another preserved carcass of a cub was found in a location Template:Cvt 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.<ref>Template:Cite news</ref> 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.<ref name="Boeskorov_al.2021" /> Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the Ice age.<ref>Template:Cite news</ref>
NotesEdit
See alsoEdit
- 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
ReferencesEdit
External linksEdit
Template:Sister project Template:Sister project
- 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 Template:Webarchive, from the Dinosaur Mailing List. (Groiss)
- Photo-reconstruction of Panthera spelaea, by paleoartist Roman Uchytel