Quaternary International 245 (2011) 183–185
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Guest Editorial
Late Quaternary mammal ecology: Insight from new approaches
During the last 40,000 years, the terrestrial communities of large mammals have been subject to dramatic changes not only in their taxonomic composition, but also in their ecological distribution. Many large terrestrial mammal species became extinct during this period, while surviving ones were extirpated in large portions of their distribution range and may have changed their ecological requirements (e.g. Stuart and Lister, 2007; Drucker et al., 2008; Bridault and Chaix, 2009). This period was a time of major climate changes, with the occurrence of the Last Glacial Maximum (LGM) between around 25,000 and 20,000 years ago and the global warming trend of the Late-Glacial and early Holocene, between around 18,000 and 8000 years ago. In parallel, it was also the period during which modern humans expanded their distribution range in the Middle and High latitudes, and colonized new continents such as the Americas and Australasia (e.g., Barnosky et al., 2004). To address the respective impact of climatic change and modern human expansion on terrestrial large mammals, we need an accurate knowledge of the following aspects: (i) a detailed chronological framework of changes in distribution ranges (e.g. Barnosky et al., 2004; Stuart and Lister, 2007), (ii) a good knowledge of the genetic structure of the ancient populations of large mammals that allows to document changes in genetic diversity and possibly leads to the reconstruction of the population dynamics through this troubled time (e.g. Drummond et al., 2005; Stiller et al., 2010; Campos et al., 2010a, 2010b), (iii) and a direct record of how the species reacted to the changing environment through phenotypic tracers related to individuals, such as stable isotope compositions in bones and teeth and tooth wear patterns (e.g. Bocherens et al., 1999; Drucker et al., 2008; Pushkina et al., 2010). This last type of approach is essential to address palaeodiets and ancient habitats at an individual level, independently of modern analogues which are often misleading since populations of recent large mammals are often relicts that do not reflect the whole range of ecological possibilities of the species when it was much more abundant, and exhibit also a restricted genetic diversity compared to the Pleistocene species members (e.g. Hofreiter and Barnes, 2010). These three essential aspects of ancient large terrestrial mammals, i.e. chronology, genetic diversity and ecological requirements, can be evaluated using new methods that have been developed during the last decades. With regard to establishing an accurate chronological framework, AMS radiocarbon dating allows the direct dating of identified bones during the last 45,000 years (e.g. Stuart and Lister, 2007; Pacher and Stuart, 2009). To evaluate genetic diversity of large terrestrial mammals during the late 1040-6182/$ – see front matter Ó 2011 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2011.07.030
Quaternary, the analysis of ancient DNA retrieved from fossil bones allows the reconstruction of phylogeography and population dynamics (e.g., Barnett et al., 2009; Stiller et al., 2010). The palaeoecology of large mammals leaves some distinct traces in their tissues, which can be retrieved from their fossil remains. Different food resources lead to different tooth wear patterns in modern herbivores, it is therefore possible to reconstruct which plant types were consumed shortly before death by fossil ungulates using the tooth microwear patterns (e.g. Merceron and Madelaine, 2006; Rivals et al., 2010). Different isotopic compositions from fossil tissues can be used to reconstruct different aspects of the palaeoecology of late Quaternary large mammals. Carbon stable isotopic compositions in collagen or bioapatite allow habitat reconstruction and niche partitioning especially among herbivores (e.g. Bocherens et al., 1999; Drucker et al., 2008; Pushkina et al., 2010), carbon and nitrogen stable isotopes in collagen allow palaeodietary reconstruction, such as food preferences in omnivores and prey choice of predators (e.g., Bocherens et al., 2005, 2006; Fox-Dobbs et al., 2008), while oxygen stable isotopes in bioapatite allow climatic reconstruction (e.g. Tütken et al., 2007). In addition, choosing tissues growing at different moments of an individual life history for isotopic analysis allows the reconstruction of some aspects of the ontogenic development of extinct species (e.g. Bocherens, 2004; Bocherens et al., 2004; Feranec, 2008). Combined with a good palaeontological background, these different approaches have already yielded invaluable information about the palaeobiology of late Quaternary mammals. However, few studies have studied late Quaternary mammal palaeoecology using a combination of these approaches on the same material, and it was the goal of the symposium “Late Quaternary mammal ecology: insight from new approaches”, organized at the EGU Annual meeting in Vienna on May 6th, 2010, to bring together specialists of these different new approaches as well as palaeontologists to integrate these various complementary approaches to provide better answers to questions related to the palaeoecology of Late Quaternary mammals. From all the manuscripts submitted, fifteen peer-reviewed papers on the outcome of the conference are being published in this issue of Quaternary International under the commendable and creditable guidance of Prof. Norm Catto. The volume starts with four articles dedicated to more accurate dating of late Quaternary species in Europe (Döppes et al., this issue; Nadachowski et al., this issue) and in South America (Kerber et al., this issue; Prevosti et al., this issue). These papers focused on a given region or a species provided some surprising results, such as the later survival of woolly mammoth in southern Poland than in more northern
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Guest Editorial / Quaternary International 245 (2011) 183–185
regions (Nadachowski et al., this issue), while the South American fox Dusicyon avus became extinct well into the Holocene and not around the transition between the late Pleistocene and Holocene as much of the other extinct large mammals (Prevosti et al., this issue). To get a better understanding of the processes leading to the extirpation or the survival of large mammals during the late Quaternary, the next two articles combined direct radiocarbon dating with palaeoecological investigations using stable isotopic tracking for reindeer (Drucker et al., this issue-a) and for cave bears (Muenzel et al., this issue). Moreover, the genetic changes associated to these changes were investigated for the cave bears from Southwestern Germany (Muenzel et al., this issue). These studies confirm that reindeer survived well into the Holocene in Southern Germany and could somehow adapt to the changing ecological conditions (Drucker et al., this issue-a). In contrast, no evidence for surviving of cave bears could be found into and after the Last Glacial Maximum in Southwestern Germany (Muenzel et al., this issue). This last paper also showed that genetic replacement was not as clear-cut as previously thought, and that it was not associated to any significant ecological change. Interestingly, the extirpation of cave bear seems to have opened a vegetarian ecological niche for the surviving brown bears, which shifted their diet from carnivorous when they coexisted with vegetarian cave bears to much more vegetarian after the cave bears were not present any more. In the Austrian Alps, a combined investigation of the genetic structure and the palaeoecological isotopic tracking of cave bears could document a case of genetic isolation associated to niche partitioning which was possibly linked to short-term climatic oscillations during the late Pleistocene and altitudinal shifts of foraging (Bocherens et al., this issuea). In addition, a comprehensive study of the existing radiometric data as well as morphological and genetic analysis of cave bear remains from ten high-alpine cave sites allow climatic implications for the period between 65,000– 30,000 years before present, and conditions warmer than today are suggested in order to provide herbivorous cave bears with sufficient food resources (Döppes et al., this issue). These examples show how it becomes possible to track the dynamics of late Quaternary mammals at different temporal and geographical scales by combining direct dating, palaeogenetic and isotopic tracking approaches on the same faunal material. Another important aspect of terrestrial ecosystems dynamics is to evaluate the prey-predator relationships in ancient ecosystems. The isotopic study focused on cave lions, studied together with coeval fauna of potential prey and competitors illustrate the possibilities offered by stable isotopic tracking to reconstruct palaeotrophic webs (Bocherens et al., this issueb). Using well identified individuals from different regions at different times, it was possible to establish an individual behaviour for this large predator, in contrast with cave hyena, and to support the hypothesis of competitive exclusion of cave lions by cave hyenas. Different cave lion individuals seem to have focused their prey choice either on reindeer or on young cave bears, and the fact that Late-glacial cave lions strongly focused their predation on reindeer suggests that the demise of this cervid may be involved in the extirpation of its predator at the end of the Pleistocene in Western Europe. Not only extinct species were investigated with new approaches but also surviving ones, such as red deer and bison. For red deer, one palaeogenetic study (Stankovi c et al., this issue) and one palaeoecological study using stable isotopic tracking (Drucker et al., this issue-b) showed how the phylogeography and the palaeoecology of this species changed between the Pleistocene and the Holocene, implying that using modern populations as analogues for ancient ones might prove problematic in such cases. Another case where modern analogues were not fully representative of the variations
exhibited by populations of ancient relatives was the tooth microwear study of Rivals and Semprebon (this issue) on North American late Pleistocene bisons. Such examples further show that using modern representatives of fossil species may be misleading and phenotypic ecological indicators, such as stable isotopes and tooth microwear, should be widely used in the palaeoecological studies of late Quaternary mammals. Late Quaternary large mammals can also become indicators of ancient climatic conditions, since their isotopic signatures depend on the environment they lived in. Two studies published in the present volume provided clear examples, one on European woolly mammoths (Arppe et al., this issue) and one of Greek cave bears (Dotsika et al., this issue). Interestingly, the correlations observed between the oxygen isotopic composition of bears and environmental water were nearly identical between the modern populations from Greece (Dotsika et al., this issue) and those from western Canada (Bocherens et al., this issuea), showing that despite hibernation, the oxygen isotopic signatures of bears may be used as climatic indicators. At a finer level, it is also possible to evaluate ontogenetic changes during the development of an extinct species, as demonstrated by the isotopic study of (Pérez-Rama et al., this issue) on cave bears from Spain. Finally, a paper dealing with the fundamentals of isotopic biogeochemistry of bone collagen helped to refine the origin of measurement errors in this approach (Rumpelmayr et al., this issue). It seems that, after a phase of independent development, the different approaches that may provide direct chronological, genetical and ecological information on late Quaternary large mammals are now in a position to be used jointly with morphological investigations to provide more detailed information about the palaeobiology of these extinct ecosystems and their dynamics in a time of climate change and increasing human pressure. References Arppe, L., Aaris-Sørensen, K., Daugnora, L., Lõugas, L., Wojtal, P., Zupins, I., The paleoenvironmental d13C record in European woolly mammoth tooth enamel. Quaternary International, this issue. Barnett, R., Shapiro, B., Barnes, I., Ho, S.Y.W., Burger, J., Yamaguchi, N., Higham, T.F.G., Wheeler, 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. Barnosky, A.D., Koch, P.L., Feranec, R.S., Wing, S.C., Shabel, A.B., 2004. Assessing the causes of Late Pleistocene extinctions on the continents. Science 306, 70–75. Bocherens, H., 2004. Cave bear palaeoecology and stable isotopes: checking the rules of the game. In: Philippe, M., Argant, A., Argant, J. (Eds.), Proceedings of the 9th International Cave Bear Conference, Cahiers scientifiques du Centre de Conservation et d’Etude des Collections. Muséum d’Histoire naturelle de Lyon, pp. 183–188. Hors Série No 2. Bocherens, H., Billiou, D., Patou-Mathis, M., Otte, M., Bonjean, D., Toussaint, M., Mariotti, A., 1999. Palaeoenvironmental and palaeodietary implications of isotopic biogeochemistry of late interglacial Neandertal and mammal bones in Scladina Cave (Belgium). Journal of Archaeological Science 26, 599–607. Bocherens, H., Argant, A., Argant, J., Billiou, D., Crégut-Bonnoure, E., DonatAyache, B., Philippe, M., Thinon, M., 2004. Diet reconstruction of ancient brown bear (Ursus arctos) from Mont Ventoux (France) using bone collagen stable isotope biogeochemistry (13C, 15N. Canadian Journal of Zoology 82, 576–586. Bocherens, H., Drucker, D.G., Billiou, D., Patou-Mathis, M., Vandermeersch, B., 2005. Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: review and use of a multi-source mixing model. Journal of Human Evolution 49, 71–87. Bocherens, H., Drucker, D.G., Billiou, D., Geneste, J.-M., van der Plicht, J., 2006. Bears and humans in Chauvet cave (Vallon-Pont-d’Arc, Ardèche, France): Insights from stable isotopes and radiocarbon dating of bone collagen. Journal of Human Evolution 50, 370–376. Bocherens, H., Hobson, K.A., Pacher, M., Rabeder, G., Stiller, M., Tütken, T., Hofreiter, M., Niche partition between two coeval genetically distinct cave bears from Austria (Ursus spelaeus and Ursus ingressus): isotopic evidence from fossil bones. Quaternary International, this issuea. Bocherens, H., Drucker, D., Bonjean, D., Bridault, A., Conard, N.J., Cupillard, C., Germonpré, M., Höneisen, M., Münzel, S.C., Napierala, H., Patou-Mathis, M.,
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Hervé Bocherens* Fachbereich Geowissenschaften, Biogeologie, Universität Tübingen, Sigwartstraße 10, 72076 Tübingen, Germany Martina Pacher Austrian Academy of Sciences, Commission for Quaternary Research, Project “F.A.C.E.” and Institute for Palaeontology, University of Vienna, Althanstr. 14, A-1090 Vienna, Austria E-mail address:
[email protected] * Corresponding author. E-mail address:
[email protected] Available online 22 July 2011
