Panthera leo spelaea M. Boule & L. De Villeneuve, 1927
Panthera spelaea, commonly known as the cave lion (or less commonly as the steppe lion) is an
extinctPanthera species 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 living 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.
Several authors regarded Panthera spelaea as a
subspecies of the modern
lion, and therefore as Panthera leo spelaea.[7][8][9][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.[10] Some authors regard the larger Middle Pleisto
Results from morphological studies showed that it is distinct in cranial and dental
anatomy to justify the
specific status of Panthera spelaea.[11][12] Results of
phylogenetic studies also support this assessment.[13][14][15]
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.[16] 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.[14][17] However, analysis of
mitochondrial genome sequences from 31 cave lions showed that they fall into two
monophyleticclades. 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.[17][18]
Evolution
Lion-like
pantherine felids first appeared in the
TanzanianOlduvai Gorge about 1.7 to 1.2 million years ago. These cats dispersed to Europe from
East Africa in the first half 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.[19][20][3][21] Different authors considered Panthera fossils as either a distinct species ancestral to P. spelaea,[22] or as a subspecies of P. spelaea.[21][23] 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.[19][1][2]
The following cladogram shows the genetic relationship between P. spelaea and other pantherine cats.[15]
Specimens intermediate between P. fossilis and Late Pleistocene P. spelaea are referred to as the subspecies P. s. intermedia.[20] The transition from P. fossilis to Late Pleistocene P. spelaea involved significant changes in skull and tooth morphology.[24]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,[14] 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).[25]
Characteristics
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.[26][27] 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.[28] Such cave paintings suggest that male cave lions completely lacked manes, or at most had very small manes.[4]
Some individuals of P. spelaea were considerably larger than modern lions. The skeleton of an adult male found in 1985 near
Siegsdorf in Germany had a shoulder height of around 1.2 m (3 ft 11 in) and a head-body length of 2 m (6 ft 7 in) without the tail, similar in size to large modern lions.[29] Similarly, footprints attributed to a male cave lion measured 15 cm (6 in) across. The heaviest Panthera spelaea was estimated to weigh 339 kg (747 lb).[30] This shows that some individuals of P. spelaea would have been up to or over 12% larger than modern lions, but still smaller than the earlier Panthera fossilis or the
American lion (P. atrox).[31][32] Early members of the cave lion lineage (including those assigned Panthera fossils) were considerably larger than individuals of P. spelaea from the
Last Glacial Period, which were comparable in size to modern lions, with the species showing 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.[23]
P. spelaea had a relatively longer and narrower muzzle compared to that of the extant lion. Despite this, the two species do not exhibit major differences in morphology.[4] Like modern lions, females were smaller than males.[33]
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.[34] While juveniles fur coat colour was yellowish, adult cave lions are suggested to have had grey fur.[18]
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] It also lived in mountainous regions, likely because it sought out hibernating bears in montane caves as a food source during the winter.[46] While during the Last Glacial Period it was often associated with cold environments, the species also inhabited temperate environments,[47] such as in Europe during the
Last Interglacial/Eemian.[48]
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.[50] 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.[51][52] Some of these accumulations of cave lion bones have been attributed to hoarding of meat from cave lion carcasses by cave hyenas in caves occupied by the latter.[53]
Whether or not cave lions existed in prides like modern lions is unclear. Isotopic analysis done by Hervé Bocherns in 2015, suggested cave lions in Europe may have been solitary, due to scattering of individual data which was more similar to individualistic behavior compared to modern day lion populations.[50] 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.[18] 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.[18]
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.[18]
Relationship with humans
There is direct evidence of hunting of cave lions and exploitation of their pelts in Europe by both
Neanderthals during the
Middle Paleolithic,[56] as well as modern humans during
Upper Paleolithic (as evidenced by the
Magdalenian aged La Garma site in Spain).[57] 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 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 oraments.[56]
Paleolithic artwork of Panthera spelaea
Carving of a Panthera spelaea head from
Vogelherd Cave, Germany, dating to around 35-40,000 years ago
Drawing of a now-lost Paleolithic sculpture of a cave lion from Isturitz, France
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.[58]
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.[60][61][62]
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.[63]
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.[64] 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.[65] 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.[18] Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the
Ice age.[66]
^Turner, A. (1984). "Dental sex dimorphism in European lions (Panthera leo L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications". Annales Zoologici Fennici. 21: 1–8.
^Spassov, N. & Iliev, N. (1994). "Animal remains from the submerged Late Eneolithic – early Bronze Age settlements in Sozopol (South Bulgarian Black Sea Coast)". Proceedings of the International Symposium VI. Thracia Pontica. pp. 287–314.{{
cite book}}: CS1 maint: location missing publisher (
link)
^
abSotnikova, M.V. & Foronova, I.V. (2014). "First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia". Integrative Zoology. 9 (4): 517–530.
doi:
10.1111/1749-4877.12082.
PMID24382145.
^Yamaguchi, N.; Cooper, A.; Werdelin, L.; MacDonald, D. W. (2004). "Evolution of the mane and group-living in the lion (Panthera leo): a review". Journal of Zoology. 263 (4): 329–342.
doi:
10.1017/S0952836904005242.
^Hussain, S.T.; Floss, H. (2015). "Sharing the world with mammoths, cave lions and other beings: linking animal-human interactions and the Aurignacian "belief world"". Quartär. 65: 85–120.
doi:
10.7485/QU62_4.
^Chernova, O. F.; Kirillova, I. V.; Shapiro, B.; Shidlovskiy, F. K.; Soares, A. E. R.; Levchenko, V. A.; Bertuch, F. (2016). "Morphological and genetic identification and isotopic study of the hair of a cave lion (Panthera spelaea Goldfuss, 1810) from the Malyi Anyui River (Chukotka, Russia)". Quaternary Science Reviews. 142: 61–73.
Bibcode:
2016QSRv..142...61C.
doi:
10.1016/j.quascirev.2016.04.018.
^Iannucci, Alessio, Mecozzi, Beniamino, Pineda, Antonio, Sardella, Raffaele, Carpentieri, Marco, Rabinovich, Rivka, and Moncel Marie-Helene. "Early Occurence of (Panthera spelaea) at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy). Journal of Quarternary Sciences 39:3 (2024): 4.
^Diedrich, C. G. (2011). "The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe". Historical Biology. 23 (2–3): 271–311.
Bibcode:
2011HBio...23..271D.
doi:
10.1080/08912963.2010.546529.
S2CID86638786.
^Diedrich, C. G. (2013). "Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art". Quaternary Science Reviews. 76: 167–193.
Bibcode:
2013QSRv...76..167D.
doi:
10.1016/j.quascirev.2013.05.009.
Panthera leo spelaea M. Boule & L. De Villeneuve, 1927
Panthera spelaea, commonly known as the cave lion (or less commonly as the steppe lion) is an
extinctPanthera species 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 living 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.
Several authors regarded Panthera spelaea as a
subspecies of the modern
lion, and therefore as Panthera leo spelaea.[7][8][9][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.[10] Some authors regard the larger Middle Pleisto
Results from morphological studies showed that it is distinct in cranial and dental
anatomy to justify the
specific status of Panthera spelaea.[11][12] Results of
phylogenetic studies also support this assessment.[13][14][15]
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.[16] 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.[14][17] However, analysis of
mitochondrial genome sequences from 31 cave lions showed that they fall into two
monophyleticclades. 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.[17][18]
Evolution
Lion-like
pantherine felids first appeared in the
TanzanianOlduvai Gorge about 1.7 to 1.2 million years ago. These cats dispersed to Europe from
East Africa in the first half 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.[19][20][3][21] Different authors considered Panthera fossils as either a distinct species ancestral to P. spelaea,[22] or as a subspecies of P. spelaea.[21][23] 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.[19][1][2]
The following cladogram shows the genetic relationship between P. spelaea and other pantherine cats.[15]
Specimens intermediate between P. fossilis and Late Pleistocene P. spelaea are referred to as the subspecies P. s. intermedia.[20] The transition from P. fossilis to Late Pleistocene P. spelaea involved significant changes in skull and tooth morphology.[24]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,[14] 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).[25]
Characteristics
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.[26][27] 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.[28] Such cave paintings suggest that male cave lions completely lacked manes, or at most had very small manes.[4]
Some individuals of P. spelaea were considerably larger than modern lions. The skeleton of an adult male found in 1985 near
Siegsdorf in Germany had a shoulder height of around 1.2 m (3 ft 11 in) and a head-body length of 2 m (6 ft 7 in) without the tail, similar in size to large modern lions.[29] Similarly, footprints attributed to a male cave lion measured 15 cm (6 in) across. The heaviest Panthera spelaea was estimated to weigh 339 kg (747 lb).[30] This shows that some individuals of P. spelaea would have been up to or over 12% larger than modern lions, but still smaller than the earlier Panthera fossilis or the
American lion (P. atrox).[31][32] Early members of the cave lion lineage (including those assigned Panthera fossils) were considerably larger than individuals of P. spelaea from the
Last Glacial Period, which were comparable in size to modern lions, with the species showing 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.[23]
P. spelaea had a relatively longer and narrower muzzle compared to that of the extant lion. Despite this, the two species do not exhibit major differences in morphology.[4] Like modern lions, females were smaller than males.[33]
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.[34] While juveniles fur coat colour was yellowish, adult cave lions are suggested to have had grey fur.[18]
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] It also lived in mountainous regions, likely because it sought out hibernating bears in montane caves as a food source during the winter.[46] While during the Last Glacial Period it was often associated with cold environments, the species also inhabited temperate environments,[47] such as in Europe during the
Last Interglacial/Eemian.[48]
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.[50] 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.[51][52] Some of these accumulations of cave lion bones have been attributed to hoarding of meat from cave lion carcasses by cave hyenas in caves occupied by the latter.[53]
Whether or not cave lions existed in prides like modern lions is unclear. Isotopic analysis done by Hervé Bocherns in 2015, suggested cave lions in Europe may have been solitary, due to scattering of individual data which was more similar to individualistic behavior compared to modern day lion populations.[50] 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.[18] 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.[18]
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.[18]
Relationship with humans
There is direct evidence of hunting of cave lions and exploitation of their pelts in Europe by both
Neanderthals during the
Middle Paleolithic,[56] as well as modern humans during
Upper Paleolithic (as evidenced by the
Magdalenian aged La Garma site in Spain).[57] 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 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 oraments.[56]
Paleolithic artwork of Panthera spelaea
Carving of a Panthera spelaea head from
Vogelherd Cave, Germany, dating to around 35-40,000 years ago
Drawing of a now-lost Paleolithic sculpture of a cave lion from Isturitz, France
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.[58]
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.[60][61][62]
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.[63]
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.[64] 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.[65] 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.[18] Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the
Ice age.[66]
^Turner, A. (1984). "Dental sex dimorphism in European lions (Panthera leo L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications". Annales Zoologici Fennici. 21: 1–8.
^Spassov, N. & Iliev, N. (1994). "Animal remains from the submerged Late Eneolithic – early Bronze Age settlements in Sozopol (South Bulgarian Black Sea Coast)". Proceedings of the International Symposium VI. Thracia Pontica. pp. 287–314.{{
cite book}}: CS1 maint: location missing publisher (
link)
^
abSotnikova, M.V. & Foronova, I.V. (2014). "First Asian record of Panthera (Leo) fossilis (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia". Integrative Zoology. 9 (4): 517–530.
doi:
10.1111/1749-4877.12082.
PMID24382145.
^Yamaguchi, N.; Cooper, A.; Werdelin, L.; MacDonald, D. W. (2004). "Evolution of the mane and group-living in the lion (Panthera leo): a review". Journal of Zoology. 263 (4): 329–342.
doi:
10.1017/S0952836904005242.
^Hussain, S.T.; Floss, H. (2015). "Sharing the world with mammoths, cave lions and other beings: linking animal-human interactions and the Aurignacian "belief world"". Quartär. 65: 85–120.
doi:
10.7485/QU62_4.
^Chernova, O. F.; Kirillova, I. V.; Shapiro, B.; Shidlovskiy, F. K.; Soares, A. E. R.; Levchenko, V. A.; Bertuch, F. (2016). "Morphological and genetic identification and isotopic study of the hair of a cave lion (Panthera spelaea Goldfuss, 1810) from the Malyi Anyui River (Chukotka, Russia)". Quaternary Science Reviews. 142: 61–73.
Bibcode:
2016QSRv..142...61C.
doi:
10.1016/j.quascirev.2016.04.018.
^Iannucci, Alessio, Mecozzi, Beniamino, Pineda, Antonio, Sardella, Raffaele, Carpentieri, Marco, Rabinovich, Rivka, and Moncel Marie-Helene. "Early Occurence of (Panthera spelaea) at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy). Journal of Quarternary Sciences 39:3 (2024): 4.
^Diedrich, C. G. (2011). "The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe". Historical Biology. 23 (2–3): 271–311.
Bibcode:
2011HBio...23..271D.
doi:
10.1080/08912963.2010.546529.
S2CID86638786.
^Diedrich, C. G. (2013). "Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art". Quaternary Science Reviews. 76: 167–193.
Bibcode:
2013QSRv...76..167D.
doi:
10.1016/j.quascirev.2013.05.009.