Elsevier

Quaternary International

Volume 245, Issue 2, 6 December 2011, Pages 249-261
Quaternary International

Isotopic evidence for dietary ecology of cave lion (Panthera spelaea) in North-Western Europe: Prey choice, competition and implications for extinction

https://doi.org/10.1016/j.quaint.2011.02.023Get rights and content

Abstract

The prey choice of extinct cave lions Panthera spelaea was determined using bone collagen isotopic signatures in the Belgian Ardennes and the Swabian Jura between 40,000 and 25,000 years ago as well as in the Late-glacial of the northwestern Alp foreland and of the Paris Basin. More than 370 specimens of large carnivorous and herbivorous mammals from 25 sites coeval with cave lion were analyzed. The isotopic results point to an individualistic prey choice for cave lions, with some individuals more oriented on reindeer and others on young cave bears. The isotopic signatures and therefore dietary choice of cave lions did not overlap with those of cave hyenas, indicating competitive exclusion between the large predators. The most recent western European cave lions seem to have been consuming mainly reindeer until the local extirpation of this prey species, which coincides chronologically with their own extinction. This restricted prey choice may be involved in the extinction of this large predator in Western Europe.

Introduction

Determining the actual dietary preferences of extinct species is a difficult task even when closely related modern representatives are still extant. This is especially true when the environmental conditions, potential prey and competitor suite, as well as the genetic structure were different in the past than for the modern populations used for comparison. This is the case for cave lions, considered sometimes as a subspecies of modern lions Panthera leo spelaea (e.g., Kurtén, 1985, Burger et al., 2004, Barnett et al., 2009), or treated as a distinct species, Panthera spelaea (e.g., Argant, 1996, Stuart and Lister, 2011). This paper follows the latter option in ranking the cave lion as a separate species. Investigations of ancient DNA have demonstrated a long genetic separation between the modern lions from subtropical Africa and Asia, and the Eurasian and American lions adapted to cold climates (Burger et al., 2004, Barnett et al., 2009). A similar conclusion was based on distinctive morphological characters from the skull and teeth (Sotnikova and Nikolskiy, 2006). Although modern lions have survived quite successfully until the advent of firearms, cave lions became extinct in their whole distribution area by around 12,000 years ago (Stuart and Lister, 2011). Understanding the reasons behind this extinction implies a detailed knowledge of the ecology of this species, including its diet, more precisely the prey preferences in case of a predator such as the cave lion.

The main approaches used to infer prey preferences in extinct predators such as the cave lion are analogy with modern relatives and evidence of predation activity on preys in the fossil record. In the case of the cave lion, fortunately there is a very close modern relative abundantly studied for its dietary ecology (e.g., Breuer, 2005, Funston and Mills, 2006) and for competition with other large predators (Trinkel and Kastberger, 2005, Hayward, 2006). Some predators are also closely related to the ones formerly sharing the same environment with cave lions, such as cave hyenas Crocuta crocuta spelaea (e.g., Rohland et al., 2005), even if behavioural differences are to be expected between modern and Pleistocene hyenas, the latter being most probably a more active predator than modern hyenas (e.g., Baryshnikov, 1999). Since modern lions consume essentially large ungulates such as zebras and large antelopes but are unable to tackle adults of very large species such as elephants and rhinoceros, it is generally assumed that cave lions should have had a similar prey choice shifted to larger sized prey, as the average size of cave lions was larger than that of modern lions (e.g., Hemmer, 2004). This leads to the assumption that cave lions were primarily consuming horse and bison, as well as reindeer, muskoxen and young individuals of mammoths and woolly rhinoceros (e.g.. Guthrie, 1968, Turner, 2009, Stuart and Lister, 2011). Based on such an assumption, a link has been suggested between the decline of genetic diversity in bison around 45,000 years ago in North America and the decrease of genetic diversity that occurred in fossil lions around the same time, as cave lions were supposed to be preferentially feeding on this large herbivore (Barnett et al., 2009). However, extant lion populations exhibit important variations in their diet and prey choice, even within a given ecosystem, as documented in the Serengeti (Hopcraft et al., 2005). Although the habitats used by modern lions range from desert to closed woodland (Patterson, 2007), none of the modern lion populations dwells under ecological conditions even remotely comparable to the steppe-tundra of the late Pleistocene, making the ecological analogy between modern and fossil lions very difficult.

Another source of information is the direct predation traces left by cave lions on the fossil remains of their prey. Most spectacular are those found on the mummy of an Alaskan bison, bite marks and a tip of lion canine embedded in the carcass, showing clearly that cave lions happened to prey on bison (Guthrie, 1990). However, such evidence yields information about which prey species was consumed by cave lions, but does not provide quantitative results. In contrast, approaches such as stable isotopic tracking may provide such direct quantitative evidence at an individual level.

Based on the analysis of the faunal composition and taphonomical aspects of some upper Pleistocene cave deposits, some inferences have been made on possible antagonism between cave lions and cave hyenas (e.g.. Diedrich, 2008, Diedrich, 2009b). Here again, it is very difficult to infer quantitatively this type of ecological interaction between predators in the past with such evidence, while stable isotope approaches allow to test dietary competition between predators by comparing their signatures: different isotopic signatures indicate that the average source of proteins was different for both predators (e.g., Lavin et al., 2003, Zhao et al., 2004, Lewis et al., 2006, Mitani et al., 2009).

Finally, prehistoric cave paintings depicting cave lions together with potential prey species have sometimes been interpreted as reflecting naturalistic behaviour. For instance, in Chauvet Cave (Ardèche, France), a whole panel includes several lions and bisons that may represent a hunting event (Packer and Clottes, 2000). Such an interpretation implies that cave lions hunted cooperatively in pack as modern lions do in savanna environments, and that they hunted bison. However, there is no certainty in the meaning of such representations and, although anatomical details of such cave paintings are usually very accurate, the species association may have other meanings than predator–prey interactions. Even if such a hunting event actually happened, it is not clear how representative it would be in the predatory behaviour and the palaeobiology of cave lions in terms of occurrence in time and across the distribution range of the species.

The present paper aims at considering quantitatively how much of different prey species were consumed by individual cave lions, and to which extent competition with other large predators impacted on the dietary ecology of cave lions. It considers exemplary sites in North Western Europe, where cave lions co-occur with potential prey and competitors, during two different time periods, before and after the Last Glacial Maximum (LGM). Pre-Last Glacial Maximum (pre-LGM) assemblages of large mammals from sites in the Ardennes (Belgium) and the Swabian Jura (Southern Germany), dated between around 40,000 and 24,000 14C years ago, were used. These sites have yielded, besides cave lions, other predators that were potential competitors of the lions, such as leopards, cave hyenas, wolves, wolverines and brown bears, as well as a diverse assemblage of herbivores that represent the potential prey, such as reindeer, bison, horse, woolly rhinoceros and mammoth, as well as cave bears. Although cave bears belong to the order Carnivora, the isotopic data gathered for specimens from the studied sites clearly point to a vegetarian diet, ruling them out from dietary competition with lions (e.g., Bocherens et al., 1997, Bocherens et al., 2011; Münzel et al., 2008, Bocherens et al., in press). Predation of cave lions on cave bears has also been suggested based on taphonomical analyses (Weinstock, 1999, Diedrich, 2009b), which led to the inclusion of cave bears in the list of potential prey of cave lions. The sites dated from the late-glacial (post-LGM) considered here include some of the most recently dated cave lions, in the Paris basin (France) and in the French and Swabian Jura (France, Switzerland, Germany) (Stuart and Lister, 2007, Stuart and Lister, 2011).

Section snippets

Material and methods

Faunal remains from 25 sites were studied in the present work. Five of these sites yielded cave lions and other mammalian species while the other sites provided additional specimens of species of coeval herbivores and carnivores in the same area (Fig. 1). Altogether, 375 individual carbon and nitrogen isotopic signatures are reported here, of which 33 from Scladina cave were previously published in Bocherens et al. (1997). Nine were published as “horse” from the Swabian Jura by Stevens and

Results and discussion

In the Ardennes and the Swabian Jura during the pre-LGM period, around 40,000 to 24,000 14C BP, a similar pattern of δ13C and δ15N values was exhibited by ungulates: reindeer presented the least negative δ13C values and rather low δ15N values, while woolly mammoth and woolly rhinoceros exhibited the highest δ15N values of all herbivores and mammoth the most negative δ13C values. Horses were relatively low in δ13C values and their δ15N values were variable. When present, large bovines (Bos or

Conclusion

The carbon and nitrogen isotopic signatures of bone collagen from the pre-LGM sites pointed to a large scatter of the individual data reflecting individualistic predatory behaviour with long lasting prey preferences, with some individuals strongly oriented on reindeer and others possibly on juvenile cave bears. In the post-LGM sites, the isotopic data point to a preference on reindeer for cave lion. It appears that cave lion was the most reindeer oriented large predator during the late

Acknowledgments

B. Steinhilber, H. Taubald, A. Orendi, and C. Wissing are warmly thanked for their technical support. The Alexander von Humboldt Foundation, the Deutsche Forschungsgemeinschaft (DFG, grant CO 226/14-1), the region Franche-Comté (France), and the French Ministry of Culture (grant PCR P08) provided significant financial support for this research.

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