Panthera Spelaea
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''Panthera spelaea'', commonly known as the cave lion (or less commonly as the steppe lion), is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
''
Panthera ''Panthera'' is a genus within the family (biology), family Felidae, and one of two extant genera in the subfamily Pantherinae. It contains the largest living members of the cat family. There are five living species: the jaguar, leopard, lion, ...
'' species that was native to Eurasia and northwest North America during the
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
epoch. Genetic analysis of
ancient DNA Ancient DNA (aDNA) is DNA isolated from ancient sources (typically Biological specimen, specimens, but also environmental DNA). Due to degradation processes (including Crosslinking of DNA, cross-linking, deamination and DNA fragmentation, fragme ...
has revealed that while closely related, it was a distinct species genetically isolated from the modern
lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'', native to Sub-Saharan Africa and India. It has a muscular, broad-chested body (biology), body; a short, rounded head; round ears; and a dark, hairy tuft at the ...
(''Panthera leo''), with the
genetic divergence Genetic divergence is the process in which two or more populations of an ancestral species accumulate independent genetic changes ( mutations) through time, often leading to reproductive isolation and continued mutation even after the populations h ...
between the two species estimated at around 500,000 years ago. The earliest fossils of the ''P. spelaea'' lineage (either regarded as the separate species ''
Panthera fossilis ''Panthera fossilis'' (also known as ''Panthera leo fossilis'' or ''Panthera spelaea fossilis'') is an extinct species of Felidae, cat belonging to the genus ''Panthera,'' known from remains found in Eurasia spanning the Middle Pleistocene and po ...
'' or the subspecies ''P. spelaea fossilis'') in Eurasia date to around 700,000 years ago (with possible late Early Pleistocene records). It is closely related and probably ancestral to the
American lion The American lion (''Panthera atrox'' (), with the species name meaning "savage" or "cruel", also called the North American lion) is an extinct pantherine cat native to North America during the Late Pleistocene from around 129,000 to 12,800 y ...
(''Panthera atrox''). The species ranged from Western Europe to eastern
Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 70th parallel north, 72° north latitude in the Chukchi Sea; and on the south ...
in North America, and was a prominent member of the
mammoth steppe The mammoth steppe, also known as steppe-tundra, was once the Earth's most extensive biome. During glacial periods in the later Pleistocene, it stretched east-to-west, from the Iberian Peninsula in the west of Europe, then across Eurasia and thr ...
fauna, and an important
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
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. ''Panthera spelaea'' interacted with both
Neanderthals Neanderthals ( ; ''Homo neanderthalensis'' or sometimes ''H. sapiens neanderthalensis'') are an extinction, extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle Pleistocene, Middle to Late Plei ...
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.


Research history and taxonomy

In 1774, the Zoolithenhöhle cave near the village of Burggaillenreuth in
Bavaria Bavaria, officially the Free State of Bavaria, is a States of Germany, state in the southeast of Germany. With an area of , it is the list of German states by area, largest German state by land area, comprising approximately 1/5 of the total l ...
, 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 In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
''Felis spelaea'' by
Georg August Goldfuss Georg August Goldfuß (18 April 1782 – 2 October 1848) was a German palaeontologist, zoologist and botanist. He became a professor of zoology at the University of Erlangen and later at the University of Bonn. He coined the terms "protozoa" an ...
. 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 A monograph is generally a long-form work on one (usually scholarly) subject, or one aspect of a subject, typically created by a single author or artist (or, sometimes, by two or more authors). Traditionally it is in written form and published a ...
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 In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
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 The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
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 Anatomy () is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old scien ...
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 Siberia ( ; , ) is an extensive geographical region comprising all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has formed a part of the sovereign territory of Russia and its predecessor states ...
and
Yukon Yukon () is a Provinces and territories of Canada, territory of Canada, bordering British Columbia to the south, the Northwest Territories to the east, the Beaufort Sea to the north, and the U.S. state of Alaska to the west. It is Canada’s we ...
, 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 In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s. One lived across western Europe and the other was restricted to Beringia during the
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
. For this reason, the Beringian population is considered a distinct subspecies, ''P. s. vereshchagini''.


Evolution

Lion-like pantherine felids first appeared in the
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
n
Olduvai Gorge The Olduvai Gorge or Oldupai Gorge in Tanzania is one of the most important paleoanthropology, paleoanthropological localities in the world; the many sites exposed by the gorge have proven invaluable in furthering understanding of early human evo ...
about . These cats dispersed into Eurasia from
East Africa East Africa, also known as Eastern Africa or the East of Africa, is a region at the eastern edge of the Africa, African continent, distinguished by its unique geographical, historical, and cultural landscape. Defined in varying scopes, the regi ...
around the end of the
Early Pleistocene The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
and the beginning of the
Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...
, giving rise to ''
Panthera fossilis ''Panthera fossilis'' (also known as ''Panthera leo fossilis'' or ''Panthera spelaea fossilis'') is an extinct species of Felidae, cat belonging to the genus ''Panthera,'' known from remains found in Eurasia spanning the Middle Pleistocene and po ...
.'' The oldest widely accepted fossils of ''P. fossilis'' in Europe date to around 700,000-600,000 years ago, such as that from
Pakefield Pakefield is a suburb of the town of Lowestoft, in the East Suffolk district of Suffolk, England. It is located around south of the centre of the town. It 2011 the ward had a population of 6,563. Pakefield has boundaries with Carlton Colvill ...
in England, with possible older fossils from
Western Siberia Western Siberia or West Siberia ( rus, Западная Сибирь, p=ˈzapədnəjə sʲɪˈbʲirʲ; , ) is a region in North Asia. It is part of the wider region of Siberia that is mostly located in the Russia, Russian Federation, with a Sout ...
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. The following cladogram shows the genetic relationship between ''P. spelaea'' and other pantherine cats.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 ''Homotherium'' is an extinct genus of Homotherini, scimitar-toothed cat belonging to the extinct subfamily Machairodontinae that inhabited North America, Eurasia, and Africa, as well as possibly South America during the Pliocene and Pleistocene ...
'' 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 Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
sequence data from fossil lion remains show that the
American lion The American lion (''Panthera atrox'' (), with the species name meaning "savage" or "cruel", also called the North American lion) is an extinct pantherine cat native to North America during the Late Pleistocene from around 129,000 to 12,800 y ...
represents a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
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 Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 70th parallel north, 72° north latitude in the Chukchi Sea; and on the south ...
(now Alaska and adjacent regions) during the Illinoian (around 190-130,000 years ago).


Characteristics

Carvings and cave paintings of cave lions, which were discovered in the
Lascaux Lascaux ( , ; , "Lascaux Cave") is a network of caves near the village of Montignac, Dordogne, Montignac, in the Departments of France, department of Dordogne in southwestern France. Over 600 Parietal art, parietal cave painting, wall paintin ...
and
Chauvet Cave The Chauvet-Pont-d'Arc Cave ( ) in the Ardèche department of southeastern France is a cave that contains some of the best-preserved figurative cave paintings in the world, as well as other evidence of Upper Paleolithic life.Clottes (2003b), p. ...
s in France, were dated to 15,000 to 17,000 years old. 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 In most terrestrial mammals, the scrotum (: scrotums or scrota; possibly from Latin ''scortum'', meaning "hide" or "skin") or scrotal sac is a part of the external male genitalia located at the base of the penis. It consists of a sac of skin ...
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 ''Panthera'' is a genus within the family (biology), family Felidae, and one of two extant genera in the subfamily Pantherinae. It contains the largest living members of the cat family. There are five living species: the jaguar, leopard, lion, ...
'' (''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 , 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 A cusp is the most pointed end of a curve. It often refers to cusp (anatomy), a pointed structure on a tooth. Cusp or CUSP may also refer to: Mathematics * Cusp (singularity), a singular point of a curve * Cusp catastrophe, a branch of bifu ...
morphology (displaying preparastyles). Like modern lions, females were smaller than males. Compared to the earlier ''P.'' (''spelaea'') ''fossilis'', Late Pleistocene ''P. spelaea spelaea'' differs (in addition to previously mentioned size differences) in having larger
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
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 In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an physical body, object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an satellite, artificia ...
(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. In 2016, hair found near the Maly Anyuy River was identified as cave lion hair through
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
analysis. Comparison with hair of a modern
lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'', native to Sub-Saharan Africa and India. It has a muscular, broad-chested body (biology), body; a short, rounded head; round ears; and a dark, hairy tuft at the ...
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 An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages, and g ...
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 The mammoth steppe, also known as steppe-tundra, was once the Earth's most extensive biome. During glacial periods in the later Pleistocene, it stretched east-to-west, from the Iberian Peninsula in the west of Europe, then across Eurasia and thr ...
, from Western Europe to northwest North America. It was widely distributed in the
Iberian Peninsula The Iberian Peninsula ( ), also known as Iberia, is a peninsula in south-western Europe. Mostly separated from the rest of the European landmass by the Pyrenees, it includes the territories of peninsular Spain and Continental Portugal, comprisin ...
, Italian Peninsula,
Southeast Europe Southeast Europe or Southeastern Europe is a geographical sub-region of Europe, consisting primarily of the region of the Balkans, as well as adjacent regions and Archipelago, archipelagos. There are overlapping and conflicting definitions of t ...
,
Great Britain Great Britain is an island in the North Atlantic Ocean off the north-west coast of continental Europe, consisting of the countries England, Scotland, and Wales. With an area of , it is the largest of the British Isles, the List of European ...
,
Central Europe Central Europe is a geographical region of Europe between Eastern Europe, Eastern, Southern Europe, Southern, Western Europe, Western and Northern Europe, Northern Europe. Central Europe is known for its cultural diversity; however, countries in ...
, the East European Plain, the
Ural Mountains The Ural Mountains ( ),; , ; , or simply the Urals, are a mountain range in Eurasia that runs north–south mostly through Russia, from the coast of the Arctic Ocean to the river Ural (river), Ural and northwestern Kazakhstan.
, most of
Northeast Asia Northeast Asia or Northeastern Asia is a geographical Subregion#Asia, subregion of Asia. Its northeastern landmass and islands are bounded by the Pacific Ocean, North Pacific Ocean. The term Northeast Asia was popularized during the 1930s by Ame ...
(ranging as far south as
Northeast China Northeast China () is a geographical region of China, consisting officially of three provinces Liaoning, Jilin and Heilongjiang. The heartland of the region is the Northeast China Plain, the largest plain in China with an area of over . The regi ...
and possibly the
Korean peninsula Korea is a peninsular region in East Asia consisting of the Korean Peninsula, Jeju Island, and smaller islands. Since the end of World War II in 1945, it has been politically divided at or near the 38th parallel between North Korea (Dem ...
), and across the
Bering land bridge Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72° north latitude in the Chukchi Sea; and on the south by the tip of the ...
into
Alaska Alaska ( ) is a non-contiguous U.S. state on the northwest extremity of North America. Part of the Western United States region, it is one of the two non-contiguous U.S. states, alongside Hawaii. Alaska is also considered to be the north ...
and
Yukon Yukon () is a Provinces and territories of Canada, territory of Canada, bordering British Columbia to the south, the Northwest Territories to the east, the Beaufort Sea to the north, and the U.S. state of Alaska to the west. It is Canada’s we ...
. The cave lion had a wide elevation range, with finds extending up over above sea level in the European Alps and in
Buryatia Buryatia, officially the Republic of Buryatia, is a republic of Russia located in the Russian Far East. Formerly part of the Siberian Federal District, it has been administered as part of the Far Eastern Federal District since 2018. To its nort ...
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 In physical geography, a steppe () is an ecoregion characterized by grassland plains without closed forests except near rivers and lakes. Steppe biomes may include: * the montane grasslands and shrublands biome * the tropical and subtropica ...
and grasslands although it would have also have 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.


Paleobiology


Ecology

''P. spelaea'' was one of the
keystone species A keystone species is a species that has a disproportionately large effect on its natural environment relative to its abundance. The concept was introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in main ...
of the mammoth steppe, being one of the main
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
s alongside the
gray wolf The wolf (''Canis lupus''; : wolves), also known as the grey wolf or gray wolf, is a canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, including the dog and dingo, though gr ...
, cave hyena and
brown bear The brown bear (''Ursus arctos'') is a large bear native to Eurasia and North America. Of the land carnivorans, it is rivaled in size only by its closest relative, the polar bear, which is much less variable in size and slightly bigger on av ...
. Large amounts of bones belonging to ''P. spelaea'' were excavated in caves, where bones of cave hyena,
cave bear The cave bear (''Ursus spelaeus'') is a prehistoric species of bear that lived in Europe and Asia during the Pleistocene and became extinct about 24,000 years ago during the Last Glacial Maximum. Both the word ''cave'' and the scientific name '' ...
and
Paleolithic The Paleolithic or Palaeolithic ( years ago) ( ), also called the Old Stone Age (), is a period in human prehistory that is distinguished by the original development of stone tools, and which represents almost the entire period of human prehist ...
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.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.
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. Isotopic analyses of bone
collagen Collagen () is the main structural protein in the extracellular matrix of the connective tissues of many animals. It is the most abundant protein in mammals, making up 25% to 35% of protein content. Amino acids are bound together to form a trip ...
samples extracted from remains in Europe and East Beringia indicate that
reindeer The reindeer or caribou (''Rangifer tarandus'') is a species of deer with circumpolar distribution, native to Arctic, subarctic, tundra, taiga, boreal, and mountainous regions of Northern Europe, Siberia, and North America. It is the only re ...
were particularly prominent in the diet of cave lions in these regions during the Last Glacial Period. Cave lions also seem to have opportunistically preyed on the cubs of cave bears, with adult cave bears also likely being targeted at least on occasion. Isotopic analysis of other European specimens suggests a diet including
wild horse The wild horse (''Equus ferus'') is a species of the genus Equus (genus), ''Equus'', which includes as subspecies the modern domestication of the horse, domesticated horse (''Equus ferus caballus'') as well as the Endangered species, endangered ...
,
woolly mammoth The woolly mammoth (''Mammuthus primigenius'') is an extinct species of mammoth that lived from the Middle Pleistocene until its extinction in the Holocene epoch. It was one of the last in a line of mammoth species, beginning with the African ...
and cave bears for these individuals. It may have sought out hibernating bears in montane caves as a food source during the winter. Bite marks found on the bones of straight-tusked elephants in Neumark Nord, Germany, dating to the
Last Interglacial The Last Interglacial, also known as the Eemian, was the interglacial period which began about 130,000 years ago at the end of the Penultimate Glacial Period and ended about 115,000 years ago at the beginning of the Last Glacial Period. It cor ...
, have been suggested to be the result of scavenging by cave lions. Other possible prey species were giant deer,
red deer The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or Hart (deer), hart, and a female is called a doe or hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Ir ...
, muskox,
aurochs The aurochs (''Bos primigenius''; or ; pl.: aurochs or aurochsen) is an extinct species of Bovini, bovine, considered to be the wild ancestor of modern domestic cattle. With a shoulder height of up to in bulls and in cows, it was one of t ...
,
wisent The European bison (: bison) (''Bison bonasus'') or the European wood bison, also known as the wisent ( or ), the zubr (), or sometimes colloquially as the European buffalo, is a European species of bison. It is one of two extant species of bi ...
,
steppe bison The steppe bison (''Bison'' ''priscus'', also less commonly known as the steppe wisent and the primeval bison) is an extinct species of bison which lived from the Middle Pleistocene to the Holocene. During the Late Pleistocene, it was widely dist ...
, and young
woolly rhinoceros The woolly rhinoceros (''Coelodonta antiquitatis'') is an extinct species of rhinoceros that inhabited northern Eurasia during the Pleistocene epoch. The woolly rhinoceros was a member of the Pleistocene megafauna. The woolly rhinoceros was larg ...
. 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 ''Homotherium'' is an extinct genus of Homotherini, scimitar-toothed cat belonging to the extinct subfamily Machairodontinae that inhabited North America, Eurasia, and Africa, as well as possibly South America during the Pliocene and Pleistocene ...
'', and Beringian wolves in
Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 70th parallel north, 72° north latitude in the Chukchi Sea; and on the south ...
.


Social behavior

Whether or not cave lions were gregarious like modern lions is unclear. 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. 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. 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. 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. 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. 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. 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.


Relationship with humans

Cave lions were hunted and their pelts exploited in Europe by
Neanderthal Neanderthals ( ; ''Homo neanderthalensis'' or sometimes ''H. sapiens neanderthalensis'') are an extinction, extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle Pleistocene, Middle to Late Plei ...
s during the
Middle Paleolithic The Middle Paleolithic (or Middle Palaeolithic) is the second subdivision of the Paleolithic or Old Stone Age as it is understood in Europe, Africa and Asia. The term Middle Stone Age is used as an equivalent or a synonym for the Middle P ...
, and during the
Upper Paleolithic The Upper Paleolithic (or Upper Palaeolithic) is the third and last subdivision of the Paleolithic or Old Stone Age. Very broadly, it dates to between 50,000 and 12,000 years ago (the beginning of the Holocene), according to some theories ...
by modern humans in Spain as evidenced in the La Garma site dating to the
Magdalenian Magdalenian cultures (also Madelenian; ) are later cultures of the Upper Paleolithic and Mesolithic in western Europe. They date from around 17,000 to 12,000 years before present. It is named after the type site of Abri de la Madeleine, a ro ...
. 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 The figurine, also called the Lion-man of , is a prehistoric ivory sculpture discovered in Hohlenstein-Stadel, a German cave, part of the Caves and Ice Age Art in the Swabian Jura UNESCO World Heritage Site, in 1939. The German name, , meaning ...
(''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. Decorated stones with engravings representing cave lions have been found in southern Italy.


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 14-15,000 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 The mammoth steppe, also known as steppe-tundra, was once the Earth's most extensive biome. During glacial periods in the later Pleistocene, it stretched east-to-west, from the Iberian Peninsula in the west of Europe, then across Eurasia and thr ...
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. Competition with wolves may have been an important factor in its extinction. Cave lions appear to have undergone a
population bottleneck A population bottleneck or genetic bottleneck is a sharp reduction in the size of a population due to environmental events such as famines, earthquakes, floods, fires, disease, and droughts; or human activities such as genocide, speciocide, wid ...
that considerably reduced their genetic diversity between 47,000 and 18,000 years ago, probably driven at least in part by climatic instability.


Mummified specimens

In 2008, a well-preserved mature cave lion specimen was unearthed near the Maly Anyuy River in
Chukotka Autonomous Okrug Chukotka ( ; ), officially the Chukotka Autonomous Okrug, is the easternmost federal subjects of Russia, federal subject of Russia. It is an Autonomous okrugs of Russia, autonomous okrug situated in the Russian Far East, and shares a border wi ...
, 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 Siberia ( ; , ) is an extensive geographical region comprising all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has formed a part of the sovereign territory of Russia and its predecessor states ...
in
permafrost Permafrost () is soil or underwater sediment which continuously remains below for two years or more; the oldest permafrost has been continuously frozen for around 700,000 years. Whilst the shallowest permafrost has a vertical extent of below ...
. 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 The Indigirka (; ) is a river in the Sakha Republic in Russia between the Yana to the west and the Kolyma to the east. It is long. The area of its basin is . History The isolated village of Russkoye Ustye, located on the delta of the Indigi ...
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 Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for determining the age of an object containing organic material by using the properties of radiocarbon, a radioactive isotope of carbon. The method was ...
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. Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the
Ice age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages, and g ...
.


Notes


See also

* History of lions in Europe *
Megafauna In zoology, megafauna (from Ancient Greek, Greek μέγας ''megas'' "large" and Neo-Latin ''fauna'' "animal life") are large animals. The precise definition of the term varies widely, though a common threshold is approximately , this lower en ...
*'' Panthera atrox'' *'' Panthera blytheae'' *'' Panthera gombaszoegensis'' *'' Panthera palaeosinensis'' *'' Panthera shawi'' *'' Panthera youngi'' *'' Panthera zdanskyi'' *'' Panthera leo sinhaleyus'' *'' Panthera leo fossilis''


References


External links


Prehistoric cats and prehistoric cat-like creatures
from the Messybeast Cat Resource Archive

volume 5, chapter 1, by Hans Krause.

, from the Dinosaur Mailing List. (Groiss)
Photo-reconstruction of Panthera spelaea
by paleoartis
Roman Uchytel
{{Authority control spelaea spelaea Pleistocene carnivorans Pleistocene extinctions Pleistocene mammals of Asia Pleistocene mammals of Europe Pleistocene mammals of North America Fossil taxa described in 1810 Taxa named by Georg August Goldfuss Apex predators Species that are or were threatened by climate change History of lions in Europe