Multi-isotopic tracking (δ13C, δ15N, δ34S) of ancient trophic webs around the time of Neandertal replacement by anatomically modern humans in North-Western Europe Dissertation der Mathematisch-Naturwissenschaftlichen Fakultät der Eberhard Karls Universität Tübingen zur Erlangung des Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.) vorgelegt von Magister Christoph Wißing aus Melle Tübingen 2016 Gedruckt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Eberhard Karls Universität Tübingen. Tag der mündlichen Qualifikation: 09.06.2016 Dekan: Prof. Dr. Wolfgang Rosenstiel 1. Berichterstatter: Prof. Dr. Hervé Bocherens 2. Berichterstatter: Prof. Dr. Ronny Schönberg Contents Abstract 1 1.Introduction 5 1.1. Climatological conditions in Europe 5 1.2. Ecological context in Europe 8 1.3. Hominids in Europe around 40,000 years ago 9 1.4. Principles of isotopic tracking in a Pleistocene 13 context13 2. Objectives 14 3. Material 16 3.1. Ziegeleigrube Coenen 17 3.2. “Troisième caverne” of Goyet 18 3.3. Spy Cave 20 3.4. Scladina Cave 20 3.5. Lommersum 21 4. Methods 22 4.1. Collagen preparation and isotopic analysis 22 4.2. Statistical methods 24 4.2.1. Cluster analysis 24 4.2.2. Canonical discriminant analysis 24 4.2.3. Bayesian mixing model 24 5. Isotopic results 25 5.1. Belgium sites 25 5.1.1. Hominids 25 5.1.2. Faunal remains 27 5.2. Ziegeleigrube Coenen 31 5.3. Lommersum 33 6. Discussion 34 6.1. Impact of climatic fluctuations and the impact of hominids on the mammothsteppe ecosystem 34 6.1.1. Impact of climatic fluctuation on the structure of the mammothsteppe 34 6.1.2. Impact of human activity on the 38 mammothsteppe ecosystem 6.2. Niche partitioning in the late Pleistocene 40 6.2.1. 2D-scatter plot (13C versus 15N) 41 6.2.1.1. Herbivores 42 6.2.1.2. Carnivores 43 6.2.1.3. Hominids 44 6.2.2. Discriminant analysis 45 6.2.3. Cluster Analysis 47 6.2.3.1. Herbivores 47 6.2.3.2. Carnivores 49 6.2.4. Summary niche partitioning 50 6.3. Hominid prey preference 51 6.3.1. Neandertals 52 6.3.2. Modern humans 54 6.3.3. The role of mammoth as prey 55 6.3.4. Review of isotopic signatures of late Neandertals 57 6.3.5. Review of isotopic signatures of ancient modern humans 59 6.3.6. Summary diet reconstruction for hominids 60 6.4. Aspects of habitat and mobility 61 6.4.1. Context 61 6.4.2. late Pleistocene faunal remains Belgium 61 6.4.3. late Pleistocene human remains Belgium 63 6.4.3.1. Neandertals 6.4.3.2. Modern humans 64 7. Conclusions 65 8. Acknowledgements 67 9. References 68-82 Publications published or preparation to fulfil the requirements for a cumulative dissertation. Percentages of the own contribution to the articles or manuscripts are listed in parentheses (original idea/data collection/data analysis/writing and publication). Paper I (75/100/80/75) Wißing, C., Matzerath, S., Turner, E., Bocherens, H., 2015a. Paleoecological and climatic implications of stable isotope results from late Pleistocene bone collagen, Ziegeleigrube Coenen, Germany. Quaternary Research 84, 96-105. Paper II (70/40/80/75) Wißing, C., Rougier, H., Crevecoeur, I., Germonpré, M., Naito, Y.I., Semal, P., Bocherens, H., 2015b. Isotopic evidence for dietary ecology of late Neandertals in North-Western Europe. Quaternary International (in press). Paper III (75/40/80/75) Wißing, C., Rougier, H., Crevecoeur, I.,Drucker, D. G., Germonpré, M., Krause, J., Naito, Y.I., Posth, C., Schönberg, R., Semal, P., Bocherens, H. Different landscape use but no dietary change between early modern humans and last Neandertals in NW-Europe (close to submission) List of publications not included but relevant to the topic of the thesis Bocherens, H., Drucker, D., Wißing, C., 2014. Die Mammutsteppe. Isotopenuntersuchungen in einem vergangenen Ökosystem. Natur – Forschung – Museum 144 (7/8) 2014, 226–231. Bocherens, H., Drucker, D.G., Geronpré, M., Lázničková-Galetová, M., Naito, Y.I., Wißing, C., Brůžek, J., Oliva, M., 5. Reconstruction of the Gravettian food-web at Předostí I using ulti-isotopic tracking (13C, 15N, 34S) of bone collagen. Quaternary International 359-360, 211-228. Naito, Y.I., Bocherens, H., Chikaraishi, Y., Drucker, D.G., Wißing, C., Yoneda, M., Ohkouchi, N., 2015. An overview of methods used for the detection of aquatic resource consumption by humans: Compound-specific delta N-15 analysis of amino acids in archaeological materials. Journal of Archaeological Science: Reports (in press). Matzerath, S., Turner, E., Fischer, P., van der Plicht, J., Bocherens, H., Wißing, C., Kels, H., Interdisziplinäre Forschungen zu eiszeitlichen Großsäugern aus der Ziegeleigrube Coenen, Kreis Düren. Archäologie im Rheinland 2012, 67–69. Naito, Y.I., Chikaraishi, Y., Drucker, D.G., Ohkouchi, N., Semal, P., Wißing, C., Bocherens, H., 2016. Ecological niche of Neanderthals from Spy Cave revealed by nitrogen isotopes of individual amino acids in collagen. Journal of Human Evolution 93, 82-90. Posth, C., Renaud, G., Mittnik, A., Drucker, Dorothée G., Rougier, H., Cupillard, C., Valentin, F., Thevenet, C., Furtwängler, A., Wißing, C., Francken, M., Malina, M., Bolus, M., Lari, M., Gigli, E., Capecchi, G., Crevecoeur, I., Beauval, C., Flas, D., Germonpré, M., van der Plicht, J., Cottiaux, R., Gély, B., Ronchitelli, A., Wehrberger, K., Grigourescu, D., Svoboda, J., Semal, P., Caramelli, D., Bocherens, H., Harvati, K., Conard, Nicholas J., Haak, W., Powell, A., Krause, J., 2016. Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe. Current Biology 26. Rougier, H., Crevecoeur, I., Beauval, C., Flas, D., H., B., Wißing, C., Germonpé, M., Semal, P., Plicht, J., 2013. The First Upper Paleolithic Human Remains from Belgium: Aurignacian, Gravettian and Magdalenian Fossils at the "Troisième caverne" of Goyet. Poster at the Paleoanthropology Society Meeting , Hawai :A 33. Rougier, H., Crevecoeur, I., Beauval, C., Posth, C., Flas, D., Wißing, C., Furtwängler, A., Germonpré,M., Gómez-Olivencia, A., Semal, P., van der Plicht, J., Bocherens, H., Krause, J. First evidence of Neandertal cannibalism in Northern Europe, accepted in Scientific Reports. Multi-isotopic tracking (δ13C, δ15N, δ34S) of ancient trophic webs around the time of Neandertal replacement by anatomically modern humans in North-Western Europe Abstract The study investigated ecological aspects of the mammoth steppe ecosystem in NW Europe with a special focus on the role of late Neandertals and early modern humans through the analysis of stable isotopes in bone collagen (13C, 15N and 34S). Around the time of Neandertal extinction and the arrival of modern humans a diverse community of large herbivorous and carnivorous mammal lived under cold, fluctuating environmental conditions in NW Europe. The study reconstructed the ecological robustness of this ancient ecosystem and its response to oscillating climatic conditions as well as in regard of the ecological participation of Neandertals and early modern humans. The trophic web, the niche partitioning, the ecological flexibility of single species, as well as the general buffer capacity of this biocenosis are objects of investigation. The Belgium site Troisième caverne of Goyet provided a broad set of directly dated late Neandertal and very early modern human skeletal remains associated with a considerably assemblage of Pleistocene faunal remains. The occurrence of late Neandertals and early modern human skeletal remains at the same place are unique circumstance and allowed a direct comparison of several ecological aspects during a time span with a special relevance for the human evolution. The site Ziegeleigrube Coenen is contemporaneous with the occurrence of late Neandertals and reflects the ecosystem during or shortly after a cold spell. Even during such phases the structure of the mammoth steppe ecosystem was intact and provided consequently an adequate basis for late Neandertal subsidence. In an ecological context the Neandertals were part of there are no indications for any stress on the structure of the ecosystem. Through the analysis of 13C and 15N isotopic ratios in bone collagen of faunal and hominid remains the trophic web was evaluated. Processing of the isotopic data 1 through several statistical approaches allowed a detailed reconstruction of ecological niche partitioning of late Neandertals and early modern humans. Both hominid species occupied the same distinct ecological niche in terms of diet with a clear preference for mammoth and reindeer. The investigation of the 34S composition in Pleistocene bone collagen is a relatively novel approach and provided insights into spatial hominid procurement. The Goyet Neandertals do not reflect the local fauna 34S congruously their main prey had an origin different from the neighborhood of the Belgium sites (Scladina, Spy and Goyet). In contrast the 34S signal of the Spy Neandertals and the Goyet ancient modern humans corresponds to the values obtained on the local fauna. Potentially different mobility strategies among Pleistocene hominids while exploring the same prey species is the drafted scenario. The site Lommersum reflects the ecosystem at an initial phase of occupation by modern humans in the area, when Neandertals became already extinct. At this time the regional mammoth population was declining, since the horses entered their niche in distinct region of Western Europe. This study hypothesized that increasing hunting pressure on mammoth by modern humans was the main reason for this, since so far no significant environmental changes on terrestrial ecosystems could be observed in NW Europe. This study indicates that no distinct ecological change happened through the time of the last Neandertals, even during colder phases of the OIS 3. Potential ecological reasons could not be served as arguments for Neandertal extinction in NW Europe. With the capture of the area by representants of the Aurignacian the mammoth steppe ecosystem started to show evidences for stress, in this case at least partly independently of environmental fluctuations.