SSP4.4 Late Quaternary mammal ecology: insight from new approaches (direct dating, stable isotopes, DNA) Conveners: Hervé Bocherens & Martina Pacher

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1 SSP4.4 Late Quaternary mammal ecology: insight from new approaches (direct dating, stable isotopes, DNA) Conveners: Hervé Bocherens & Martina Pacher ABSTRACTS

2 Vol. 12, EGU , 2010 Insights into the European glacial palaeoclimate from phosphate δ 18 O in mammoth tooth enamel Laura Arppe (1) and Juha Karhu (2) (1) University of Helsinki, Department of Geology, University of Helsinki, Finland (laura.arppe@helsinki.fi, ), (2) University of Helsinki, Department of Geology, University of Helsinki, Finland (juha.karhu@helsinki.fi, ) Quantitative palaeoclimatical and -ecological records on the European continent from the time period prior to the LGM in MIS 3 and 2 (ca ka) are scarce and fragmentary, and particularly poor in the core areas of the Scandinavian Ice Sheet (SIS) in Northern Europe. In this area, the palaeoenvironmental record contained in mammal skeletal remains can prove invaluable. This study makes use of the numerous finds of woolly mammoth (Mammuthus primigenius), which was common across most of Europe during the latest the Weichselian glacial age. Mammoth molars discovered from Estonia, Latvia, Lithuania, Poland and Denmark were analysed for the oxygen isotope composition of enamel phosphate to explore spatial patterns and temporal variations of glacial δ 18 O ppt (oxygen isotope value of precipitation) in northern Europe prior to and after the LGM during MIS 3 and 2. The molars (n=28) have all been radiocarbon dated (AMS), and have calendar dates from 49 to 11 ka. In order to extend the geographical coverage of the δ 18 O data, the new results were combined with compiled oxygen isotope data from prior investigations of mammoth enamel phosphate and palaeogroundwaters. The enamelderived long-term mean glacial δ 18 O ppt values agree remarkably well with local glacial palaeogroundwater oxygen isotope levels in several European locations. The combined dataset facilitates a first approximation of the geographic distribution of δ 18 O ppt values in glacial precipitation over much of Europe. The long-term mean values for the oxygen isotope composition of precipitation in Europe at ka were 1-4 lower than those for the present-day precipitation, with the largest changes recorded in the currently marine influenced southern Sweden and the Baltic region (Arppe and Karhu, 2010). The reconstructed geographic pattern of δ 18 O ppt in glacial precipitation reflects the progressive isotopic depletion of air masses moving northeast, consistent with a westerly source of moisture for the entire region, and a circulation pattern similar to that of the present day. The application of regionally varied δ/t-slopes, estimated from palaeogroundwater data and modern spatial correlations, yields reasonable estimates of glacial surface temperatures in Europe and imply 2-9 C lower long-term mean annual surface temperatures during the glacial period. The isotopic composition of carbon in enamel was also analysed and, as expected, it indicates a pure C3 diet for the European mammoths (Ukkonen et al., in prep.). A faint geographical gradient in the δ 13 C values of enamel is discernible, with more negative values in the northeast. References Arppe, L, Karhu, J.A., Oxygen isotope values of precipitation and the thermal climate in Europe during the middle to late Weichselian ice age. Quaternary Science Reviews, revision submitted. Ukkonen, P., Aaris-Sørensen, K., Arppe, L., Clark, P.U., Daugnora, L., Lister, A., Lõugas, L., Seppä, H.A., Stuart, A.J., Wojtal, P., Zupins, I. Northern Europe in the Middle to Late Weichselian: palaeoenvironment of the woolly mammoth (Mammuthus primigenius Blum.) In preparation.

3 Vol. 12, EGU , 2010 Stable isotopes in collagen and Late Quaternary carnivore palaeoecology Hervé Bocherens Universität Tübingen, Institute for Geosciences, Biogeology, Tübingen, Germany Several taxa of large carnivores co-occurred during the late Pleistocene in the steppe-tundra ecosystem, such as wolf Canis lupus, cave lion Panthera leo spelaea, cave hyaena Crocuta crocuta spelaea, brown bear Ursus arctos and cave bear Ursus spelaeus and Ursus ingressus. This abundance of taxa belonging to the same guild raises questions about niche partitioning, especially in terms of dietary specialization and prey selection. Observations of the dietary ecology of the extant relatives of these late Pleistocene carnivores does not provide unambiguous answers as these populations live under very different environmental conditions where other potential prey species are present, but it appears that most of these modern large carnivores are relatively flexible in their prey selection. Palaeontological investigations dealing with faunal associations and activity marks on fossil bones also have their limitations, such as taphonomic biases (palimpsests rather than biological associations) and do not allow the quantification of consumption of various preys. In contrast, carbon and nitrogen isotopic signatures of bone collagen depend directly on those of the average diet. Since different potential prey species occurring in the steppe-tundra exhibit consistent isotopic differences for these chemical elements, it is possible to relate the carbon and nitrogen isotopic signatures measured in fossil carnivores with the preferential consumption of some prey species. Some amount of quantification can be provided using modified versions of mixing models developed for modern ecosystems. In addition, this isotopic approach is individual-based and it is therefore possible to investigate intra- and inter-population differences in prey selection, as well as possible chronological trends and differences linked to genetic differences by combining isotopic and ancient DNA studies on the same material. The isotopic approach has already shown that among the tested large carnivores, cave bears of various genetic types are overwhelmingly vegetarian while coeval brown bears are essentially carnivores, cave lions have a marked preference for reindeer, and none of the large carnivores match Neanderthals in terms of megaherbivore consumption (i.e. woolly mammoth and woolly rhinoceros).

4 Vol. 12, EGU , 2010 Contribution of stable isotopes (C,N,S) in collagen of late Pleistocene large mammal trophic ecology and landscape use: a case study in Goyet and Scladina cave (30-40,000 years BP) Hervé Bocherens (1) and Mietje Germonpré (2) (1) Universität Tübingen, Institute for Geosciences, Biogeology, Tübingen, Germany (herve.bocherens@uni-tuebingen.de), (2) Department of Paleontology, Royal Belgian Institute of Natural Sciences, Brussels, Belgium (Mietje.Germonpre@naturalsciences.be) Two Belgian caves yielded very rich large mammal associations dating around 30 to 40,000 years ago: Goyet and Scladina cave (layer 1A). These sites are only 5 km apart but the cave entrances open on different valleys, in a quite diverse landscape ranging between open, unprotected uplands, steep cliffs and sheltered sun-exposed gorges, with the larger Meuse valley nearby. This mosaic scenery permitted during the Last Glacial a rich diversity of fossil flora and fauna. The faunal association includes a large diversity of taxa including Aurochs Bos primigenius, steppe bison Bison priscus, reindeer Rangifer tarandus, giant deer Megaloceros giganteus, horse Equus ferus, woolly rhinoceros Coelodonta antiquitatis, woolly mammoth Mammuthus primigenius, cave bear Ursus spelaeus, brown bear Ursus arctos, wolf Canis lupus, cave lion Panthera leo spelaea, and cave hyaena Crocuta crocuta spelaea. All the 90 studied bones and teeth yielded collagen with excellent collagen preservation, allowing reliable investigations of carbon, nitrogen and sulfur isotopic biogeochemistry. The combination of three different isotopic tracers allows to deciphering the effects of food selection and landscape use by different herbivorous and carnivorous taxa. This is the first study to include sulfur isotopic signatures in the study of late Quaternary large mammal palaeobiology. This new tracer yields evidence on mobility and differences in pasture areas, as different geological bedrock may exhibit various sulfur isotopic signatures that will pass on the herbivores and further on their predators. Using this feature in addition to the trophic information provided by carbon and nitrogen isotopic signatures, it appears that for some species present in both sites, such as horse and woolly rhinoceros, the individuals found in each site probably did not use the same pasture areas. This seems to also the case for the overwhelmingly vegetarian cave bears. In addition, individuals from the same species found in one site sometimes exhibit clear isotopic differences in the three isotopic tracers that suggest different pasture grounds as well, and therefore the possibility of different herds in the vicinity of the cave site. In the case of cave hyeanas, the isotopic signatures of individuals from both caves are not significantly different and suggest that these predators obtained their prey from a large territory including the pasture grounds of herbivores from both caves. In addition, direct radiocarbon dating of some of the studied collagen allows to investigating possible chronological trends. This study shows how new isotopic tracers can provide invaluable information on late Quaternary large mammal palaeobiology.

5 Vol. 12, EGU , 2010 Faunal reorganisation in terrestrial mammalian communities: evidence from France during the Lateglacial-Early Holocene transition anne BRIDAULT CNRS, UMR 7041(Archéologies environnementales), M.A.E., NANTERRE, France The Lateglacial-Early Holocene transition is characterized by rapid oscillations between warm and cold episodes. Their impact on ecosystem dynamics was particularly pronounced in north-western Europe where hunter-gatherer societies experienced a succession of environmental transformations, including the expansion and dispersal of biotic communities and changing herbivore habitats. Recent archaeozoological studies and AMS direct dating on mammalian bones/or bone collagen allow to map and precise this process at a supra-regional scale (France). At regional scales (i.e. Paris Basin & Jura-Northern French Alps), results indicate a rapid faunal reorganisation at the end of Lateglacial that will be presented in detail. Composition of faunal assemblages remains then unchanged during the Early Holocene. By contrast, significant herbivore habitat changes are recorded during the Early Holocene by other proxies (pollen data and isotopic data) and a decrease in Red Deer size through time is evidenced by osteometrical analyses. Hypotheses regarding the kind of adaptation process experienced by the faunal communities through time will be presented. Factors that may have controlled the observed changes will be discussed.

6 Vol. 12, EGU , 2010 Spotted hyena and steppe lion predation behaviours on cave bears of Europe?Late Quaternary cave bear extinction as result of predator stress Cajus G. Diedrich PaleoLogic, Geology/Palaeontology, Halle/Westph., Germany Cave bears hibernated in caves all over Eurasia (e.g. Rabeder et al., 2000) including alpine regions using mainly larger caves for this purpose. Late Quaternary spotted hyenas Crocuta crocuta spelaea instead occupied mainly areas close to the cave entrances as their dens (Diedrich and Žák 2006, Diedrich 2010). The largest predator, the steppe lion Panthera leo spelaea was only a sporadic cave dweller (Diedrich 2007b, 2009b). His presence and its remains from caves all over Europe can be recently explained best as result of imported carcasses after killing by their largest antagonists, the Late Quaternary spotted hyenas. In some cases the kill might have happened in the hyena den cave itself during the theft of prey remains by lions (Diedrich 2009a). Another reason of their remains in caves of Europe is the hunting onto the herbivorous cave bears, especially during hibernation times, when megafauna prey was less available in the open environments (Diedrich 2009c). These lion remains from caves of Europe, nearly all of which were from adult animals, provide evidence of active predation by lions onto cave bears even in medium high alpine regions (Diedrich 2009b, in review). Lion skeletons in European cave bear dens were therefore often found amongst originally articulated cave bear skeletons or scattered cave bear remains and even close to their hibernation nests (Diedrich et al. 2009c, in review). Not only lions fed on cave bears documented mainly by the large quantities of chewed, punctured and crushed cave bear long-bones; even damaged skulls reveal that hyenas scavenged primarily on cave bear carcasses which were mainly responsible for the destruction of their carcasses and bones (Diedrich 2005, 2009d). Predation and scavenging on cave bears by the two largest Late Quaternary predators C. c. spelaea and P. l. spelaea explains well the large quantity of fragmented cave bear bones over all European caves in low to medium high mountainous elevations, whereas in high alpine regions the leopard Panthera pardus seem to have used the ecological niche of the absent hyenas (Diedrich 2009d, in review a). At open air sites cave bear scavenging by the largest Late Quaternary predators were proven, too (Diedrich 2006, 2009e). The predation stress caused by the three main and largest Late Quaternary predators seem to have provided cave bears to hibernate often deeply in many European caves, and here especially in larger and longer cave systems (e.g. Diedrich et al 2009, in review, Diedrich and Moldovan 2010) to protect themselves against the largest Quaternary predators. In conflicts with those large felids must have been killed by adult cave bears, which explains why those predator carcasses remained as complete skeletons or as partly disarticulated ones even deep in caves all over Europe which were finally not scavenged by the herbivorous cave bears (Diedrich in review). In such cave bear den caves the amount of lion bones generally take only 1-3% of the total bone amount being highly dominated by cave bear bones (Diedrich 2009c, in review). Lions and hyenas seem to have focussed onto the hunt of cave bears all over Europe, especially with the reduction of the biomass in the open environments at the maximum cold period during around BP (Solutrean). The slow extinction of their largest prey was an important motor of the predation pressure onto cave bears. This might be one reason for the cave bear extinction in the final Late Quaternary (early Late Weichselian, maximum cold period) at least in northern Europe, but it seem to have happened in a combination of climatic change, and also human impact (cf. Diedrich, this volume). References Diedrich, C., Cracking and nibbling marks as indicators for the Upper Pleistocene spotted hyena as a scavenger of cave bear (Ursus spelaeus Rosenmüller, 1794) carcasses in the Perick Caves den of Northwest Germany. Abhandlungen der Naturhistorischen Gesellschaft Nürnberg, 45,

7 Diedrich, C., Ice age spotted hyenas?hunting or only scavenging on a cave bear Ursus spelaeus Rosenmüller at the Ice Age spotted hyena freeland den and prey deposit site Bad Wildungen-Biedensteg (Hessia, Germany). Scientific Annals, School of Geology Aristotle University of Thessaloniki (AUTH) special volume, 98, Diedrich, C., 2007b. The fairy tale about the "cave lions" Panthera leo spelaea (Goldfuss 1810) of Europe - Late Ice Age spotted hyenas and Ice Age steppe lions in conflict - lion killers and scavengers around Prague (Central Bohemia). Scripta Facultatis Scientiarum Universitatis Masarykianae Geology, 35 (2005), Diedrich, C., 2009a. Steppe lion remains imported by Ice Age spotted hyenas into the Late Pleistocene Perick Caves hyena den in Northern Germany. Quaternary Research, 71 (3), Diedrich, C., 2009b. Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from the Bilstein Caves (Sauerland Karst) and contribution to the steppe lion taphonomy, palaeobiology and sexual dimorphism. Annales de Paléontologie, 95, Diedrich, C. 2009c. Steppe lion predation another reason for cave bears to protect during hibernation against carnivores in deepest parts of large cave systems in Europe. Abstractc 15th International Cave Bear Symposium, Spisska Nova Ves Slovakia, 8 9. Diedrich, C., 2009d. Cave bear killers, scavengers between the Scandinavian and Alpine Ice shields the last hyenas and cave bears in antagonism - and the reason why cave bears hibernated deeply in caves. Stalactite, 58(2), Diedrich, C. 2009e. Late Pleistocene Cave bear remains from the open air hyena den Emscher River terrace site Bottrop (NW Germany). - Stalactite, 58 (2): Diedrich, C The Crocuta crocuta spelaea (GOLDFUSS 1823) population from the Late Pleistocene Rösenbeck Cave den in NW Germany and contribution to the sexual dimorphism and palaeobiogeography of the last hyenas of Europe. - Annales de Paléontologie. Diedrich, C. (in review). Cave bear killers and scavengers from the last European Ice Age. Palaeogeography Palaeoclimatology Palaeoecology. Diedrich, C., Žák, K 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), Diedrich, C., Robu, M., Dragusin, V., Constantin, S., Moldovan, O., New Upper Pleistocene steppe lion skeleton finds between the cave bear hibernation plateaus of the Ursilor Cave bear den, Romania. Abstractc 15th International Cave Bear Symposium, Spisska Nova Ves Slovakia, 10. Diedrich, C., Moldovan, O., Constantin, S Cave bear tracks, scratch marks, hair traces and hibernation nests in the Ursilor Cave (Transylvania, Romania) preliminary report from a famous European cave bear den. - Stalactite, 58 (2): Diedrich, C., Moldovan, O Ichnological and ethological studies in one of Europe s famous bear den in the Ursilor Cave (Carpathians, Romania). - Quarternary International (accepted). Rabeder, G., Nagel, D., Pacher, M., Der Höhlenbär. Stuttgart: Thorbecke.

8 Vol. 12, EGU , 2010 Disappearance of the last lions and hyenas of Europe in the Late Quaternary a chain reaction of large mammal prey migration, extinction and human antagonism Cajus G. Diedrich PaleoLogic, Geology/Palaeontology, Halle/Westph., Germany (cdiedri@gmx.net) In the Eemian to Early/Middle Weichselian (Late Pleistocene), when the Scandinavian and Alpine Glaciers were still small, and northern Germany under mammoth steppe to taiga palaoenvironment conditions, Late Quaternary steppe lions were well distributed in northern to central Germany, whereas generally all over Central Europe bones and rarely articulated skeletons were found less at open air but mainly at cave sites (Diedrich 2007a, 2008a-b, 2009a-b, 2010a-c, k, in review a-b; Diedrich and Rathgeber in review). A similar distribution, but more dense, is reported for the Late Quaternary Ice Age spotted hyenas (Diedrich 2005, 2006, 2007b-c, 2008a, c, 2010f-j, in review c-d, Diedrich and Žák 2006). The last lions of northern Europe were thought to have reached into the final Magdalénan (cf. Musil 1980). This can be not concluded with a restudy of the bone material from the Late Magdalenian (V-VI) Teufelsbrücke stone arch site near Saalfeld (Thuringia, Central Germany) and many other Magdalenian stations (open air and caves) in northern to central Germany (Münsterland Bay, Sauerland Karst, Harz Mountain Karst, Thuringian Karst). None of those sites yield remains of final Upper Pleistocene spotted hyenas or steppe lion bones anymore, nor in the few preserved Late Magdalenian mobile art can those be recognized in those regions. The last lion remains seem to reach into the Aurignacian or possibly into the Early Gravettian (early Late Weichselian) documented especially at the cave bear den, hyena den and overlapping Neandertalian to Modern human camp site Balve Cave (Sauerland Karst, cf. archaeology in Günther 1964) where still a mammoth fauna is documented for that time (Diedrich 2010a). The last and by archaeological layers dated hyena remains were also found in the Balve Cave and are from the Late Middle Palaeolithic cave site reaching a maximum Aurignacian age documenting an overlapping of hyena den and human camp site use (Diedrich 2010a, b). In northern Germany there are no records of indirectly dated hyena and lion remains being younger then even Aurignacian/Early Gravettian ( BP). Those largest Late Quaternary predators must have got extinct in northern Germany with the Late Weichselian/Wuermian extending Skandinavian Glacier, which reached northern Europe between Hamburg and Berlin its maximum extension about BP (Skupin et al. 1993). The two largest predators of the Late Quaternary of Europe seem to have been well represented in the Gravettian and up to Magdalénian Late Palaeolithic of southern Europe, in which mainly lions, but only rarely hyenas are well documented within the cave and mobile art (e.g Breuil 1952, Begouen and Clottes 1987, Chauvet et al. 1995, Diedrich and Rathgeber in review, Diedrich 2005). Hyenas and lions must have been represented in the Gravettian, Early and Middle and possibly even?late Magdalenian in southern Europe, which must have resulted from a southern migration of those predators during the Late Weichselian/Wuermian together with the megafauna and humans. The disappearance of hyenas and lions also correlate with the extinction of mammoth and woolly rhinoceros in northern Germany. The large mammals such as elephants and rhinoceroses were highly important for hyenas and lions during the Late Quaternary. Hyenas had a systematic scavenging strategy on both large prey which was even transferred until today compared to modern spotted hyenas and lions of Africa (Diedrich 2010d, e, in prep). Where those Late Quaternary giant mammals such as woolly mammoth and woolly rhinoceros were absent in middle mountainous regions (e.g. Sauerland Karst, Harz Mountain Karst, Bohemian Karst, Thuringian Karst) those had to kill other medium sized animals such as horse or steppe bison and those in larger amounts (Diedrich 2008, 2010c). Woolly rhinoceros and woolly mammoth seem to be not known in northern Germany after Aurignacian/?Early Gravettian times (Sauerland Karst and Münsterland Bay, Harz Mountain Karst) such as humans (archaeology cf. Günther 1988) but there are also lacking radiocarbon dating series or new excavations. Humans must have been in antagonism to the large predators about their prey and also their Late Palaeolithic

9 sanctuaries - the large caves especially in southern Europe, in which they left thousands of animal figurations mainly during the Gravettian to Late Magdalenian periods (e.g. Breuil 1952, Begouen and Clottes 1987, Chauvet et al. 1995). Therefore the extinction reason of the largest predators of the Late Quaternary seems to be a complex combination of climate change with resulting maximum glacier extension which finally caused a southern directed megafaunal migration and reduction of the available caves and habitat areas in general. Finally in those southern European regions a higher competition about larger caves as cave bear hibernation places, hyena den sites and human settlement/sanctuary sites must have happened. Therefore an active hunt onto lions and hyenas can not be excluded nor be proven well yet, even if there are finds of necklesses in Europe from Late Palaeolithic ages made of mainly carnivore teeth including rare lion and hyena teeth, which might have been only collected. The resettlement of the northern European region by humans and a reduced reindeer fauna and its main and largest predators such as wolves and wolverines after the maximum glacier extension in the middle Late Weichselian started not before the Late Magdalenian (Magdalénian V: e.g. Perick Caves, Martins Cave Sauerland Karst; Rübeland Caves Harz Mountain Karst, Teufelsbrücke stone arch Thuringian Karst). Humans occupied at that post-maximum glaciation time many caves in the Sauerland Karst region and even at open air sites in the Münsterland Bay or central Germany (cf. Bosinski 1987, Günther 1988). Within this Late Magdalénian V-VI times in the record of ten thousands of bones from more then 100 sites no hyena or lion remains are recorded anymore in northern and central Germany, such as cave bears. References Begouen R. and Clottes J Les Trois Frères after Breuil. Antiquity, 61: Bosinski, G Die große Zeit der Eiszeitjäger. Europa zwischen und v.chr. Jahrbuch des Römisch-Germanischen Zentralmuseums, 34: Breuil, H Four Hundred Centuries of Cave Art. Montignac, Dordogne, 414 pp. Chauvet, J.-M., Deschamps B.E, and Hillaire C Grotte Chauvet. Altsteinzeitliche Höhlenkunst im Tal der Ardèche. Thorbecke Speläo 1, Sigmaringen, 120 pp. Diedrich, C Eine oberpleistozäne Population von Crocuta crocuta spelaea (GOLDFUSS 1823) aus dem eiszeitlichen Fleckenhyänenhorst Perick-Höhlen von Hemer (Sauerland, NW Deutschland) und ihr Kannibalismus. Philippia, 12 (2): , Kassel. Diedrich, C The Crocuta crocuta spelaea (Goldfuss 1823) population from the early Upper Pleistocene hyena open air prey deposit site Biedensteg near Bad Wildungen (Hess, NW Germany) and the contribution to their phylogenetic position, coprolites and prey. Cranium, 23 (2): 39-53, Amsterdam. Diedrich, C. 2007a. Upper Pleistocene Panthera leo spelaea (Goldfuss 1810) skeleton remains from Praha-Podbaba and the contribution to other lion finds from loess and river terrace sites in Central Bohemia (Czech Republic). Bulletin of Geosciences, 82 (2), , Prague. Diedrich, C. 2007b. Crocuta crocuta spelaea (GOLDFUSS 1823) remains from the Upper Pleistocene hyaena Teufelskammer Cave den site near Hochdahl in the Neander valley (NRW, NW Germany). Cranium, 24 (2): 39-44, Amsterdam. Diedrich, C. 2007c. The Upper Pleistocene Crocuta crocuta spelaea (GOLDFUSS 1823) population and its prey from the gypsum karst den site Westeregeln near Magdeburg (Middle Germany). Abhandlungen und Berichte für Naturkunde, 30: 57-83, Magdeburg. Diedrich, C. 2008a. Late Pleistocene Hystrix (Acanthion) brachyura LINNAEUS 1758 from the Fuchsluken cave at the Rote Berg near Saalfeld (Thuringia, Germany) - a porcupine and hyena den and contribution to their palaeobiogeography. - The Open Palaeontological Journal, 2008 (1): Diedrich, C. 2008b. The rediscovered holotypes of the Upper Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss 1823) and the steppe lion Panthera leo spelaea (Goldfuss, 1810) and taphonomic discussion to the Zoolithen Cave hyena den at Geilenreuth (Bavaria, South-Germany). Zoological Journal of the Linnaean Society London, 154, Diedrich, C. 2008c. Late Pleistocene hyenas Crocuta crocuta spelaea (GOLDFUSS 1823) from Upper Rhine valley open air sites and the contribution to skull shape variability. Cranium, 25-2: (in press), Amsterdam. Diedrich, C. 2009a. Upper Pleistocene Panthera leo spelaea (GOLDFUSS 1810) remains from the Bilstein Caves (Sauerland Karst) and contribution to the steppe lion taphonomy, palaeobiology and sexual dimorphism. Annales de Paléontologie, 95 (2009) , Amsterdam. Diedrich, C. 2009b. Steppe lion remains imported by Ice Age spotted hyenas into the Late Pleistocene Perick Caves hyena den in Northern Germany. Quaternary Research 71 (3): , Amsterdam.

10 Diedrich, C. 2010a. Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from the Balve Cave (NW Germany) a hyena den and Middle Palaeolithic human site. - International Journal of Osteoarchaeology (acepted). Diedrich, C. 2010b. Ein Skelett einer kranken Löwin Panthera leo spelaea (Goldfuss 1810) und andere männliche Löwenreste aus Neumark-Nord. - Archäologie in Sachsen-Anhalt, Sonderbad (accepted). Diedrich, C. 2010c. Late Pleistocene lion Panthera leo spelaea (GOLDFUSS 1810) remains from the Keppler Cave (Sauerland Karst, NW Germany). Cranium 26, Amsterdam (in press). Diedrich, C. 2010c. Specialized horse killers in Europe foetal horse remains in the Late Pleistocene Srbsko Chlum-Komín Cave hyena den in the Bohemian Karst (Czech Republic) and actualistic comparisons to modern African spotted hyenas as zebra hunters. - Quaternary International. Diedrich, C. 2010d. Mammuthus primigenius (Blumenbach, 1799) carcass destructors, bone collectors, crackers and gnawers of the Late Pleistocene in Central Europe - the Ice Age spotted hyenas and their feeding strategies on their largest prey - the mammoth. Abstract Vth International Conference on Mammoths and their relatives, Le Puy-en-Velay, France. Diedrich, C. 2010e. Späteiszeiliche Fleckenhyänen-Fressstrategien und Steppenlöwen an ihrer größte Beute - dem Waldelefanten Palaeoloxodon antiquus Falconer & Cautley 1845 in Neumark-Nord. Archäologie in Sachsen-Anhalt, Sonderband (accepted). Diedrich, C. 2010f. Die späteiszeitlichen Fleckenhyänen und deren Exkremente aus Neumark-Nord. Archäologie in Sachsen-Anhalt, Sonderband (accepted). Diedrich, C. 2010g. Rare Upper Pleistocene Crocuta crocuta spelaea (GOLDFUSS 1823) remains from northern Czech Republic open air loess sites along the Elbe River. Cranium, (accepted). Diedrich, C. 2010h. Europe s first Upper Pleistocene Crocuta crocuta spelaea (GOLDFUSS 1823) skeleton from a small cannibalistic hyena population found in the main dome of the Koněprusy Caves a hyena cave prey depot site in the Bohemian Karst (Czech Republic). Acta Palaeontologica Polonica (in review). Diedrich, C. 2010i. Periodical use of the Balve Cave (NW Germany) as a Late Pleistocene Crocuta crocuta spelaea (GOLDFUSS 1823) den hyena occupations and bone accumulations versa human Middle Palaeolithic activity. Quaternary International. Diedrich, C. 2010j. The Crocuta crocuta spelaea (GOLDFUSS 1823) population from the Late Pleistocene Rösenbeck Cave den in NW Germany and contribution to the sexual dimorphism and palaeobiogeography of the last hyenas of Europe. Annales de Paléontologie. Diedrich, C. 2010k. Upper Pleistocene Panthera leo spelaea (Goldfuss 1810) remains from an open air loess bone accumulation site in Freyburg a. U. (Central Germany). Jahresschrift für mitteldeutsche Vorgeschichte (accepted). Diedrich, C. in prep. Coelodonta antiquitatis (Blumenbach 1799) hunters and scavengers - the Late Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss 1823) and its feeding strategy on its most important prey in Europe. Diedrich, C. in review a. A diseased Panthera leo spelaea (Goldfuss, 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord, Central Germany. Quaternary International. Diedrich, C. in review b. Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) remains from the open air hyena den Emscher River terrace site Bottrop and other sites of northern Germany new proves for hyena-lion antagonism. Palaeogeography, Palaeoclimatology, Palaeoecology. DIEDRICH, C. in review c. The Crocuta crocuta spelaea (GOLDFUSS 1823) population and its prey from the Upper Pleistocene Teufelskammer Cave hyena den site in the Neandertal (NRW, NW Germany). Annales de Paléontologie. Diedrich, C. in review d. The Late Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823) population from the Late Pleistocene hyena open air Emscher River terrace den near Bottrop (NWGermany) and other sites in the Westphalian Bay and its mammoth and woolly rhinoceros prey. Quaternary International. Diedrich, C. and Rathgeber, T. in review. Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) skeleton remains of the Upper Rhine valley (SW Germany) and contribution to their palaeobiogeography, sexual dimorphism and palaeoecology. Palaeogeography Palaeoclimatology Palaeoecology. Diedrich, C. AND Žák, K 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), Günther, K Die altsteinzeitlichen Funde der Balve Höhle. Bodenaltertümer Westfalens, 8, Günther, K Alt- und Mittelsteinzeitliche Fundplätze in Westfalen. Teil 2. Einführung in die Vor- und

11 Frühgeschichte Westfalens 6, Musil, R Die Großsäoger und Vögel der Teufelsbrücke. In: Feustel, R. Magdalénienstation Teufelsbrücke. II: Paläontologischer Teil. Weimarer Monographien zur Ur- und Frühgeschichte 3, Skupin, K., Speetzen, E. and Zandstra, J.G Die Eiszeit in Nordwestdeutschland. GLA Nordrhein-Westfalen, Krefeld, 143 pp.

12 Vol. 12, EGU , 2010 From climate change to diet change biochemistry investigations on Late Glacial and Early Holocene brown bear remains from caves in the Alpine region Doris Doeppes (1), Wilfried Rosendahl (1), Martina Pacher (2), and Hervé Bocherens (3) (1) Reiss-Engelhorn-Museen, Mannheim, Germany & (2) Austrian Academy of Science, Biologische Station Lunz am See and University of Vienna, Austria, (3) Department of Biogeology, University of Tübingen, Germany Bones of brown bears from caves in the Alpine region in Germany, Austria, Italy and Switzerland were examined and dated in the last years. The finds originate from the transition from the Bölling/Alleröd to the Early Holocene. In total we analyzed 15 samples from bones and teeth of directly radiocarbon dated brown bears from the alpine region for isotopic analyses. All collagen considered here exhibit carbon and nitrogen content similar to that of collagen extracted from fresh bones, and most of the bones and teeth contained almost the same quantity of collagen than fresh bone (around 25% weight). Atomic C/N ratios range from 3.1 to 3.4, well within the acceptable range ( ). The d13c values are rather high during the Late Glacial then a clear decrease is observed at the beginning of the Holocene. This trend coincides with the development of dense forests at low altitudes and the shift of timberline towards higher altitudes. The d15n values are relatively low in Bölling-Alleröd, then quite high during the Younger Dryas, and they decrease again during the Boreal and more recent periods. For the first time a more precise picture of the former habitat of the brown bears during the transition from the Bölling/Alleröd to the Early Holocene in the Alpine region could be reconstructed. The described investigation can also give an outlook of the coexistence of the herbivore cave bears and the omnivore/carnivore brown bears during the late Upper Pleistocene

13 Vol. 12, EGU , 2010 Palaeo-climatic information from isotopic signatures of fossil teeth in Late Pleistocene from Arkoudospilia Cave (Aridea, N. Greece) Elissavet Dotsika (1), Evangelia Tsoukala (2), Nikoleta Zisi (1), Dimitrios Poutoukis (3), and David Psomiadis (1) (1) NCSR Demokritos, Institute of Materials Science, Aghia Paraskevi, Attiki, Greece (2) Aristotle University of Thessaloniki, Department of Geology, School of Geology, Thessaloniki, Greece, (3) General Secretariat for Research and Technology, Mesogion 14-18, Athens, Greece, The O and C isotopic composition of enamel carbonate hydroxy-apatite in the teeth of certain animals reflects the oxygen isotope composition of the water they ingest. The isotopic composition of meteoric water is well-correlated with mean annual temperature so that there is potential for recovering palaeo-temperature of the regions where the animals lived. Analyses were made on enamel from fossil teeth of Ursus Ingressus from Arkoudospilia Cave in Northern Greece. Analyses were made also on modern teeth of Ursus from different areas in Greece. Oxygen and deuterium isotopic analyses of water were also made. Although the preservation of primary oxygen isotopic composition of enamel carbonate hydroxy-apatite was more difficult to assess, however the isotopic signals seem to have utility for the paleoenvironmental reconstructions of the studied area.

14 Vol. 12, EGU , 2010 Evolution of habitat and environment of deer during the Late-glacial and early Holocene: the case of red deer in French Jura. Dorothée Drucker (1), Anne Bridault (2), Alisa Hujic (1), and Hervé Bocherens (1) (1) Institut für Geowissenschaften, Biogeologie, Universität Tübingen, Germany (2) ArScAn-UMR7041, Archéologies environnementales, MAE, Nanterre, France The Late-glacial and early Holocene transition is a key period of environmental changes in a context of to a global warming. In northwestern Europe, extensive studies have documented the vegetation and faunal recomposition with the replacement of the cold steppe-tundra ecosystem by the forested temperate ecosystem we can still observe. Paleoecological interest focused on the extinct large mammals species like the Mammoth. In comparison, little has been done to decipher the ecological adaptation of the surviving species, especially those that are still present in the very same region than in the past. A better knowledge of the impact of changing environmental conditions on the ecology would be useful to define the degree of selective pressure. Thus, we have studied the habitat and environment evolution of red deer (Cervus elaphus) during the Late-glacial and early Holocene using stable isotopes and radiocarbon investigations. The analyzed bone material was selected from archaeological sites in French Jura. Performing direct radiocarbon dating on the bone collagen of the selected remains solved the problem of possible chronological uncertainties of the stratigraphical record of the sites. The same bone collagen samples were used for stable isotope measurements. We investigated the relative abundances in 13 C to examine changes in habitat closure (canopy effect), in 15 N to decipher changes in pedogenic activities (soil maturation) of the animals dwelling, and in 18 O to track changes in altitude and/or local temperatures of the occupied territories. The results demonstrate that the stable isotopic composition of red deer bone collagen can be a valuable and sensitive indicator of habitat use and environmental conditions. The associated direct dating allows us to reconstruct the chronology of ecological changes. The combined chronological and ecological results evidence local differences in red deer adaptation at a small geographical scale.

15 Vol. 12, EGU , 2010 Stable isotope time series and dentin increments elucidate Pleistocene proboscidean paleobiology Daniel Fisher (1), Adam Rountrey (2), Kathlyn Smith (3), and David Fox (4) (1) Museum of Paleontology, University of Michigan, Ann Arbor, Michigan, USA (2) Museum of Paleontology, University of Michigan, Ann Arbor, Michigan, USA (3) Department of Geological Sciences, University of Michigan, Ann Arbor, Michigan, USA (4) Department of Geology and Geophysics, University of Minnesota, Minneapolis, Minnesota, USA Investigations of stable isotope composition of mineralized tissues have added greatly to our knowledge of past climates and dietary behaviors of organisms, even when they are implemented through bulk sampling, in which a single assay yields a single, time-averaged value. Likewise, the practice of sclerochronology, which documents periodic structural increments comprising a growth record for accretionary tissues, offers insights into rates of growth and age data at a scale of temporal resolution permitted by the nature of structural increments. We combine both of these approaches to analyze dental tissues of late Pleistocene proboscideans. Tusk dentin typically preserves a record of accretionary growth consisting of histologically distinct increments on daily, approximately weekly, and yearly time scales. Working on polished transverse or longitudinal sections, we mill out a succession of temporally controlled dentin samples bounded by clear structural increments with a known position in the sequence of tusk growth. We further subject each sample (or an aliquot thereof) to multiple compositional analyses most frequently to assess δ 18 O and δ 13 C of hydroxyapatite carbonate, and δ 13 C and δ 15 N of collagen. This yields, for each animal and each series of years investigated, a set of parallel compositional time series with a temporal resolution of 1-2 months (or finer if we need additional precision). Patterns in variation of thickness of periodic sub-annual increments yield insight into intra-annual and inter-annual variation of tusk growth rate. This is informative even by itself, but it is still more valuable when coupled with compositional time series. Further, the controls on different stable isotope systems are sufficiently different that the data ensemble yields much more than the sum of its parts. By assessing how compositions and growth rates covary, we monitor with greater confidence changes in local climate, diet, behavior, and health status. We illustrate the potential of this approach with case studies that reveal: season of birth and age of weaning in juvenile mammoths; age of maturation in male mastodons; season of musth in mammoths and mastodons; and season of death and tests of simultaneity of death in mammoths and mastodons. The data provided by histological and stable isotope analyses rarely reveal cause of death directly, but they can, in concert with other observations, affect perceptions of the likelihood of competing interpretations of cause of death. Most important, paleobiological inferences based on these studies can be integrated over broad geographic and temporal scales to show how specific paleobiological traits changed through time, prior to extinction. These studies have great power for investigating causes of extinction because contrasting patterns of change are expected under different hypothesized drivers of extinction.

16 Vol. 12, EGU , 2010 AMS Radiocarbon Dating Individual Taxa and Individual Specimens: Implications for Small Mammal Paleoecology. Russell Graham (1), Thomas Stafford, Jr. (2), Holmes Semken, and Jr. (3) (1) Earth & Mineral Sciences Museum, The Pennsylvania State University, University Park, PA,16802, USA (2) Stafford Research, Inc., 200 Acadia Avenue, Lafayette, CO, 80026, USA, (3) Department of Geology, University of Iowa, Iowa City, IA, 52240, USA, Advances in AMS physics and organic geochemistry have revolutionized our ability to establish absolute chronologies on vertebrate fossils. Highly purified collagen, which provides extremely accurate 14C ages, can be extracted from single bones and teeth as small as 50 mg. Combined with measurement precisions of ±15 to 25 years for ages of < 20,000 yr, the direct AMS 14C technique enables fossil deposits to be chronologically dissected at the level of single animals. Analysis of data from a variety of sites in the United States indicates that most excavation levels (analysis units) as small as 10 cm can be time averaged by several thousand years at a minimum, even with the greatest care in excavation and processing of sediments. Time averaging of this magnitude has important implications for fine-scale paleoecological analysis of faunas, especially when compared to high-resolution climate records like those derived from speleothems, ice cores, or marine cores. To this end, we propose saturation dating of indicative taxa and plotting dates of individual specimens against high-resolution climate records rather than analysis of complete faunas or faunules. This technique provides even higher resolution of paleoenvironments than pollen spectra.

17 Vol. 12, EGU , 2010 On the palaeobiology of the extinct cave bear Ursus spelaeus ROSENMÜLLER. Insights from stable isotope analysis Aurora Grandal-d Anglade, Marta Pérez-Rama, and Daniel Fernández-Mosquera Instituto Universitario de Xeoloxia. Universidade da Coruña. Isotopic signatures (13C, 15N) of bone collagen are used more and more to obtain the paleobiological data of fossil species. By means of these signatures, for example, the diet type of an extint species may be inferred. Also, the climate in which this species developed may greatly influence on the isotopic signature of its bone collagen. This influence is firstly produced in the initial material of the trophic chain but also may produce variations due to physiological changes caused by climatic changes in the species involved in this trophic chain. The cave bear (Ursus spelaeus ROSENMÜLLER) is a species of broad distribution in the European Pleistocene sites that has been studied from the isotopic point of view, trying to establish its diet type. For the moment, the results vary: though in most cases the isotopic values indicate a preferably herbivore diet type, differences exist between sites of different geographic zones and chronologies. Taking into account that climate influences on the cave bear s physiology through the physiological mechanism of hibernation, it is expected that in bears that lived in different climatic phases, the isotopic signatures will be also different. During hibernation a recycling of nitrogenised compounds is produced for protein synthesis, including bone collagen, so it is expected that the isotopic signature, at least of Nitrogen, will be altered with respect to the synthesized collagen when the bear is active and feeds normally. However, it is difficult to establish up to what extent the isotopic signatures due to hibernation or diet are overlapped. To study the physiological effect of hibernation on isotopic signatures we have selected bone remains of cave bears from populations whose chronologies correspond to different climatic moments, and in different ontogenetic stages, coming from Galician caves (NW of the Iberian Peninsula). Adult individuals show different isotopic signatures depending on their chronology. Juvenile individuals show differences originated by the effect of a greater or less contribution of maternal milk in their diets. Finally, the neonate individuals or still in foetal stage show isotopic values that directly reflect their mother s physiology in the hibernation, during which foetal development is produced. Interestingly, all the individuals of this age class present similar isotopic values in spite of belonging to populations with different values in the adults. This indicates, on one hand, that there exists a physiological effect on the isotopic signals produced by hibernation, effect that is similar in all the studied populations, and on the other, that this physiological effect of hibernation only appears in adult individuals of populations corresponding to cold moments, in which the hibernation period is longer. According to the results, we suggest the use of isotopic values of adult cave bears as climatic proxy for European Pleistocene sites. This work is part of the Ph.D. Thesis of M.P.R. and is a contribution to the research project BTE-CGL of the Education and Science Ministry of Spain.

18 Vol. 12, EGU , 2010 Regional differences in bone collagen carbon- and nitrogen-isotope ratios of Pleistocene mammoths: Implications for paleoecology of the mammoth steppe Darren R. Grocke (1), Paul Szpak (2), and Hendrik N. Poinar (3) (1) Durham University, Earth Sciences, Durham, United Kingdom ), (2) Department of Anthropology, The University of Western Ontario, London, Ontario N6A 5C2, Canada, (3) McMaster Ancient DNA Centre, Departments of Anthropology and Biology, McMaster University, Hamilton, Ontario L8S 4K1, Canada In this study, we present bone collagen carbon- and nitrogen-isotope values from a large set of Pleistocene woolly mammoths (Mammuthus primigenius) from Siberia, Alaska and Yukon (n=58). Overall, results for mammoth specimens from eastern Beringia (Alaska and Yukon) significantly differ, for both carbon- and nitrogen-isotope values, from those from western Beringia (northeastern Siberia). In agreement with palynological, entomological, and physiographic data from the same regions, these isotopic differences strongly imply that the mammoth steppe, the extensive ice-free region spanning northern Eurasia and northwestern North America, was ecologically variable along its east-west axis to a significant degree. Prior to the Last Glacial Maximum (LGM), the high-latitude portions of Siberia and the Russian Far East appear to have been colder and more arid than central Alaska and Yukon, which were ecologically more diverse. During the LGM itself, however, isotopic signatures of mammoths from eastern Beringia support the argument that this region also experienced an extremely cold and arid climate. In terms of overall temporal trend, Beringia thus went from a condition prior to the LGM of greater ecological variability in the east to one of uniformly cold and dry conditions during the LGM.

19 Vol. 12, EGU , 2010 Palaeogenetics enlightening the ecology and evolution of species Michael Hofreiter For many years after its inception the study of ancient DNA was restricted to short fragments of mitochondrial DNA from single specimens, drastically limiting its usefulness. However, the last ten years have seen a dramatic increase in the amount of data, including an extension from mitochondrial to nuclear DNA as well as the advent of new analytical methods for interpreting the data. We are now in a position to not only decipher the phylogenetic position of extinct species using complete mitochondrial genomes but can reconstruct the population dynamics of species over time, investigate and resurrect functionally important genes, and even reconstruct complete genomes of extinct species. I will show how recovering the temporal patterns of population size in species allows more informative speculations about the contributions of environmental change versus human impact to species extinctions. I will also discuss the possibilities of deciphering phenotypic characteristics of extinct species via DNA analyses and evaluate the prospects and limitations of palaeogenomics.