The Cave Lion (Panthera spelaea)

Taxonomy

Panthera spelaea usually called the ‘The Cave Lion’ or ‘Steppe lion’ is a species or subspecies of Felidae and belongs to the genus Panthera which includes most of the extant big cats. There are 3 subspecies recognized based on morphology occurring in succession. The earliest specimens dating to the middle Pleistocene are assigned to the subspecies Panthera spelaea fossilis and late-middle Pleistocene specimens assigned to the recently described Panthera spelaea intermedia (2). Late Pleistocene populations have been assigned to Panthera spelaea spelaea, though an additional contemporary subspecies Panthera spelaea vereshchagini is sometimes recognized as a unique Beringian population (29). This split is based on genetic evidence which indicates the two as belonging to distinct haplogroups (24, 29).

Recent DNA discoveries have shown that the Cave Lion is most closely related to the American Lion (Panthera atrox) separated by about 340,000 years (3). Amongst extant species, the modern lion (Panthera leo) is the closest relative according to both morphological and genetic analyses (3, 4, 8, 12, 22, 24). The divergence date from the lion is contentious, with studies suggesting either a late split about 475,000-550,000 years ago (3, 12) or an early split 1.85-1.89 million years ago (4, 24). The huge difference hinges on the inclusion of Panthera spelaea fossilis into the calibration of molecular clocks. As a result, there is no consensus on whether the cave lion should be treated as a separate species or subspecies of Panthera leo.

Distribution and Age

The earliest putative remains date to about 800-1200kya in Western Siberia and are usually assigned to P. s. fossilis, though not without controversy (23). A more definite earliest set of fossils can be dated to various locations in Europe around 700kya (13). P. s. intermedia is dated to 150-350kya (13). Panthera spelaea spelaea appears in the fossil record around 180-200kya (13). The latest reliably dated fossils in Europe are around 14,000-14,500 years old and coincide with the extinction of most of the cold-adapted European megafauna (25), the disappearance of the Cave Lion from Siberia is concurrent with Europe (25). Alaska appears to have been the last holdout of the species with the youngest fossil dated of the species at 13,200 (25).

The Late Pleistocene sub-species is known from sites ranging from Western Europe (1) to Alaska, which was connected to Siberia via a landbridge (3). Latitudinally, the species seems to have ranged as far North as the Eurasian glaciers allowed. A few exceptions to this appear to have been North-West Siberia and Ireland. Its southern distribution is less clear, finds support a presence in the Cantabrian mountains (1) and alps (Pacher 2018). A few sites in the Italian peninsula (18) and a single site in Northern Greece (5) are attributed to the Late Pleistocene cave lion as well. Late Pleistocene and Early Holocene remains from Iberia south of the Cantabrian mountains belong to either P. leo or P. spelaea, but are too fragmented and rare to be classified, nor are they adequately dated (16). In the East, the North American distribution is exclusive to Alaska and North-West Yukon as glaciers likely blocked movement further to the East and South, not to mention the presence of the American lion (25). In Asia Late Pleistocene fossil remains seem to be restricted to modern-day Russia (25).

Morphology and Ecology

The exact size of the late Pleistocene varient P. s. spelaea has not been studied in detail, Sherani 2016 provides an estimate on the maximum known body size of the species at 339kg (21) and Marciszak et al 2021 suggests 400kg for a large male, but a reliable average is lacking in literature. Late Pleistocene specimen were considerably smaller than earlier subspecies (13). Additionally, the species falls into two distinct contemporary size classes, these are widely interpreted as being due to sexual dimorphism, with males considerably larger than females (11). Supporting evidence of this hypothesis is a cave painting displaying a male (as indicated by a scrotum) beside a much smaller female (28), furthermore, sexual dimorphism has been demonstrated in the closely related American lion (27).

The skeletal structure of the cave lion highly resembles that of other members of Panthera, and most especially the modern lion. The most notable morphological difference between the two is the difference in the limb structure, cave lions have relatively short limbs which resemble those of felids associated with closed habitats. This is in stark contrast to the savannah-adapted modern lion and the lions of the American steppe, which have the longest limbs of any late Pleistocene pantherines (20).

The cave lion appears to have been a temperate (1) and polar (10) taxon. Remains in Mediterranean climate are rare and probably represent the climatic fringe of Panthera spelaea spelaea. As an adaptation to this cold-climate niche the cave lion possessed a thick and compact undercoat of downy hair for heat retention, with a layer of thicker ‘guard hairs’ on the outside probably to protect the undercoat from the elements and/or provide additional insulation (9). The color of the guard hair different to the modern lion, lacking the light bands present in the top of lion hairs indicating a darker overall colour, though some hair did possess a yellow-ish top (9). The habitats occupied by the cave lion appears to have been wide, ranging from mountains (17) to closed and open woodland (1), to the Mammoth steppe (10).

Cave lions hunted in prides, at least on occasion. Clear evidence of this is a cave painting depicting a pride of 8 lions observing a herd of bison (25). It is unclear if prides were a universal behavior. Male cave lions lack manes, as evidenced by paintings at Chauvet cave.  This has been suggested to indicate a solitary lifestyle as larger manes in extant lions correlate with pride size, but mane size is also linked to temperature (28). More likely, manes are a sexual trait unique to the modern lion and evolved after the two taxa diverged (28).

If they exhibited co-operative hunting, Cave lions were probably capable of taking down bigger prey than any of the contemporary predators within its range. A study by Van Valkenburgh et al 2016 estimated that the max prey size of a cave lion pride would have reached 6700kg, which would have precluded only the straight-tusked elephant (Palaeoloxodon antiquus) and possibly adult male woolly mammoths (Mammuthus primigenius)(26). The typical prey range of a cave lion pride more likely falls between 200 and 900kg (26). Prey choice in the modern lion is of course very variable, depending on the abundance of different prey items in their territory, this was true for the cave lions as well.

In Western and Central Europe cave lions appear to have fed primarily on Reindeer (Rangifer tarandus) (6, 7) and Muskoxen (7), these are both relatively small species for a whole pride of giant lions, this could be considered evidence for solitary hunting, small pride size or a need to hunt frequently. The diet was however highly variable, more so than any other predator in the region. Other common megafauna prey (6, 7) included the wild horse (Equus ferus) (6), large bovines, (6) probably Aurochs (Bos primigenius) and members of the bison species complex (Bison spp.), and other cervids, notably red deer (Cervus elaphus) and Irish elk (megaloceros giganteus)(6).

The cave bear (Ursus spelaea), wooly mammoth (Mammuthus primigenius), and wooly rhinoceros (Coelodonta antiquitatis) also appear to have been part of the diet (6, 7), in these cases, it may be that juvenile prey and/or scavenging was preferred as these constitute exceptionally dangerous prey items (6). Nevertheless, it is commonly thought that the prevalent occurrence of cave lion fossils in caves was due to the hunting of U. spelaea which sheltered there. Evidence of this comes from damages to cave bear bones (both juvenile and adult) inflicted by a large predator (15), whilst some of this could be chalked up to scavenging there are bones that exhibit signs of subsequent healing (14).

Antagonistic relationships with other species certainly existed. Most notably, the cave hyena (Crocuta crocuta spelaea) would come into conflict with the lions, probably over carcasses much like their African equivalents. In fact, hyena bitemarks are common on the bones of the cave lion, with some injuries show signs of healing, suggesting cave hyenas fought and probably killed cave lions (10, 19). Hyenas may have pushed cave lions away from a generalist niche as lions rely much more heavily on reindeer and cave bears in their presence (6). Though antagonistic relationship in other species is not recorded there were several other megafaunal predators in the range, the giant brown bear (Ursus arctos priscus), the scimitar-tooth cat (Homotherium latidens), leopard (Panthera pardus), and grey wolf (Canis lupus) which all likely competed with the cave lion to some degree.

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References

1.       Alvarez-Lao, D. J., Alvarez-Vena, A., Ballestros, D., Garcia, N., Laplana, C.. (2020). A cave lion (Panthera spelaea) skeleton from Torca del Leon (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3). Quarternary Science Reviews 229, 106123.

2.       Argant, A., Brugal, J-P.. (2017). The cave lion Panthera (Leo) spelaea and its evolution: Panthera spelaea intermedia nov. subspecies. Acta zoologica cracoviensia 60(2), 60-98.

3.       Barnett, R., Shapiro, B., Barnes, I., Ho, S. Y. W., Burger, J., Yamaguchi, N., Higham, T. F. G., Wheeler, H. T., Rosendahl, W., Sher, A. V., Sotnikova, M., Kuznetsova, T., Baryshnikov, G. F., Martin, L. D., Harington, R., Burns, J. A., Cooper, A.. (2009). Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity. Molecular Ecology 18, 1668-1677.

4.       Barnett, R., Mendoza, M. L. Z., Soares, A. E. R., Ho, S. Y. W., Zazula, G., Yamaguchi, N., Shapiro, B., Kirillova, I. V., Larson, G., Gilbert, M. T. P..(2016). Mitogenomics of the Extinct Cave Lion, Panthera spelaea (Goldfuss, 1810), Resolve its Position within the Panthera Cats. Open Quaternary, 2 (4), 1–11

5.       Baryshnikov, G. F., Tsoukala, E.. (2010). New analysis of the Pleistocene carnivores from Petralona Cave (Macedonia, Greece) based on the Collection of the Thessaloniki Aristotle University. Geobios 43, 389-402.

6.       Bocherens, H., Drucker, D. G., Bonjean, D., Bridault, A., Conard, N. J., Cupillard, C., Germonpre, M., Honeisen, M., Munzel, S. C., Napierala, H., Patou-Mathis, M., Stephan, E., Uerpmann, H-P., Ziegler, R.. (2011). Isotopic evidence for dietary ecology of cave lion (Panthera spelaea) in North-Western Europe: Prey choice, competition and implications for extinction. Quarternary International 245, 249-261.

7.       Bocherens, H., Drucker, D. G., Germonpre, M., Laznickova-Galetova, M., Naito, Y. I., Wissing, C., Bruzek, J., Oliva, M.. (2015). Reconstruction of the Gravettian food-web at Predmostí I using multi-isotopic tracking (13C, 15N, 34S) of bone collagen. Quarternary International 359-360, 211-228.

8.       Burger, J., Rosendahl, W., Loreille, O., Hemmer, H., Eriksson, T., Gotherstrom, A., Hiller, J., Collins, M. J., Wess, T., Alt, K. W.. (2004). Molecular phylogeny of the extinct cave lion Panthera leo spelaea. Molecular Phylogenetics and Evolution 30, 841-849.

9.       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) Quarternary Science Reviews 142, 61-73

10.   Diedrich, C. G.. (2011). Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) footprints and bone records from open air sites in northern Germany - Evidence of hyena-lion antagonism and scavenging in Europe. Quarternary Science Reviews 30, 1883-1906.

11.   Diedrich, C. G., Rathgeber, T.. (2012). Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss 1810) skeleton remains of the Upper Rhine Valley (SW Germany) and contributions to their sexual dimorphism, taphonomy and habitus, Historical Biology, 24(1), 1-28

12.   Manuel, M., Barnett, R., Sandoval-Velasco, M., Yamaguchi, N., Vieira, F. G., Mendoza, M. L. Z., Liu, S., Martin, M. D., Sinding, M-H. S., Mak, S. S. T., Eizirik, E., Koepfli, K-P., Johnson, W. E., Antunes, A., Sicheritz-Ponten, T., Gopalakrishnan, S., Larson, G., Yang, H., O’Brien, S. J., Hansen, A. J., Zhang, G., Marques-Bonet, T., Gilbert, M. T. P.. (2020). The evolutionary history of extinct and living lions. PNAS 117 (20), 10927-10934.

13.   Marciszak, A., Schouwenvurg, C., Gornig, W., Lipecki, G.,Mackiewicz, P.. (2019). Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka. Quarternary Science Reviews 223, 105950.

14.   Marciszak, A., Sobczyk, A., Kasprzak, M., Gornig, W., Ratajczak, U., Wisniewski, A., Stefaniak, K..(2020). Taphonomic and paleoecological aspects of large mammals from Sudety Mts (Silesia, Sw Poland), with particular interest to the carnivores. Quarternary International 546, 42-63

15.   Marciszak, A., Lipecki, G., Pawlowska, K., Jakubowski, G., Ratajczak-Skrzatek, U., Zarzecka-Szubinska, K., Nadachowski, A..(2021). The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland – A review. Quarternary International Unpublished

16.   Masseti, M., Mazza, P. P. A..(2013). Western European Quarternary lions: new working hypotheses. Biological Journal of the Linnaen Society 109, 66-77

17.   Pacher, M.. (2018). Prowling the Mountains – Alpine Cave Lion (Panthera spelaea) distribution and metrics..

18.   Persico, D.. (2021).  First fossil record of cave lion (Panthera (Leo) spelaea intermedia) from alluvial deposits of the Po River in northern Italy. Quarternary International 586, 14-23.

19.   Rothschild, B. M., Diedrich, C. G.. (2012). Pathologies in the extinct Pleistocene Eurasian steppe lion Panthera leo spelaea (Goldfuss, 1810)—Results of fights with hyenas, bears and lions and other ecological stresses. International Journal of Paleopathology 2, 187-198

20.   Schellhorn, R., Sanmugaraja, M.. (2015). Habitat adaptations in the felid forearm. Palaontol Z 89, 261-269.

21.   Sherani, S.. (2016). A new specimen-dependent method of estimating felid body mass. PeerJ unpublished.

22.   Sotnikova, M., Nikolskiy, P.. (2006). Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters. Quarternary International 142-143, 218-228

23.   Sotnikova, M., 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, 517-530

24.   Stanton, D. W. G., Alberti, F., Plotnikov, V., Androsov, S., Grigoriev, S., Fedorov, S., Kosintsev, P., Nagel, D., Vartanyan, S., Barnes, I., Barnett, R., Ersmark, E., Doppes, D., Germonpre, M., Hofreiter, M., Rosendahl, W., Skoglund, P., Dalen, L.. (2020). Scientific Reports 19, 12621

25.   Stuart, A. J., Lister, A. M.. (2011). Extinction chronology of the cave lion Panthera spelaea. Quarternary Science Reviews 30, 2329-2340

26.  Van Valkenburgh, B., Hayward, M. W., Ripple, W. J., Meloro, C., Roth, V. L.. (2016). The impact of large terrestrial carnivores on Pleistocene ecosystems. PNAS 113(4). 862-867.

27.  Wheeler, H. T., Jefferson, G. T.. (2009). Panthera Atrox: Body proportions, size, sexual dimorphism, and behaviour of the cursorial lion of the North American Plains.

28.   Yamaguchi, N., Cooper, A., Werdelin, L., Macdonald, D. W.. (2004). Evolution of the mane and group-living in the lion (Panthera leo): a review

29.   Ersmark, E., Orlando, L., Sandoval-Castellanos, E., Barnes, I., Barnett, I., Stuart, A., Lister, A., Dalen, L.. (2015). Population demography and genetic diversity in the Pleistocene cave lion. Open Quarternary 1 (4), 1-14