Stable isotope analysis of well-preserved 120,000-year-old herbivore bone collagen from the Middle Palaeolithic site of Neumark-Nord 2, Germany reveals niche separation between bovids and equids Kate Britton a, b, ⁎ , 1 , Sabine Gaudzinski-Windheuser c , Wil Roebroeks d , Lutz Kindler c , Michael P. Richards a, e a Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig, Germany b Department of Archaeology, University of Aberdeen, St. Mary's Building, Elphinstone Road, Aberdeen, AB24 3UF, UK c Römisch-Germanisches Zentralmuseum, Forschungsbereich Altsteinzeit Schloss Monrepos, 56567 Neuwied, Germany and Johannes Gutenberg–Universität Mainz, Institut für Vor- und Frühgeschichte, Germany d Faculty of Archaeology, Leiden University, P.O. Box 9515, 2300 RA Leiden, The Netherlands e Department of Anthropology, 6303 NW Marine Drive, Vancouver, BC, Canada V6T 1Z1 abstract article info Article history: Received 9 August 2011 Received in revised form 18 March 2012 Accepted 20 March 2012 Available online 29 March 2012 Keywords: Stable isotopes Herbivores Bone collagen Palaeoecology Diet Resource partitioning Herbivores from the Neumark-Nord 2 archaeological site, Germany, were analysed for bone collagen stable carbon (δ 13 C) and nitrogen (δ 15 N) isotope ratios in order to investigate feeding ecology at this early Last Interglacial (Eemian) shallow-lake site. Of 42 faunal samples selected, 23 yielded collagen, demonstrating re- markable preservation for material of this age. The results indicate clear inter-specific differences in δ 15 N and δ 13 C values, notably between equids (Equus) and bovids (Bos/Bison), with mean difference Δ 15 N of +2‰ measured in the bovids compared to the equids. The potential reasons for these differences are explored, in- cluding physiology, herbivore feeding ecology, biogeography and resource partitioning within the local envi- ronment. The data are compared to previously published archaeological data, and modern experimental and ecological data, suggesting that these inter-specific differences are not consistent and therefore unlikely to be solely the product of physiology or habitual forage preference. Data from this study are compared to the local vegetation (as reconstructed from pollen profiles), and it is suggested that these trends are likely the result of niche partitioning at the shallow lake site, reflecting the local diversity in vegetational zones. The evidence for resource partitioning amongst Pleistocene herbivore communities at Neumark-Nord 2 and elsewhere is dis- cussed. This study represents one of the largest data sets for collagen of this age, and the implications for our understanding of Late Pleistocene herbivore ecology, local herbivore community behaviour and hominin palaeodietary studies are explored. © 2012 Elsevier B.V. All rights reserved. 1. Introduction The landscapes of Mid- and Late Pleistocene Europe were domi- nated by large herbivores, including pachyderms (e.g. Mammuthus, Elephas), equids (Equus), cervids (Cervus), and bovids such as bison (Bison) and aurochs (Bos). The mechanisms that allow for the co- existence of different modern mid- and large-sized grazing herbivore guilds have been debated, but are largely thought to lie in their differ- ent digestive systems that permit them to adopt alternative foraging strategies where ranges overlap (Janis, 1976), and/or through the selective feeding on different plant communities or portions of plants within the same biome—known as resource partitioning (Krysl et al., 1984; Menard et al., 2002). These mechanisms lead to niche separa- tion, through the occupation of different sub-habitats by the selection of different plants within the same habitat, or different parts of the same plant; and therefore facilitating biodiversity in herbivore species (Janis, 1976). It has previously been suggested that the high diversity of herbivore taxa living in the steppic biomes of Pleistocene North Europe can be explained by such dietary specialisation (Guthrie, 1982), with a proposed high floral biodiversity allowing niche separation and the avoidance of direct inter-specific competition (Drucker et al., 2003). In this study, we investigate the feeding palaeoecology of equids (Equus) and bovids (Bos/Bison) through stable isotope data, and explore the possibility (and nature) of such niche separation in Late Pleistocene Europe. The stable isotope analysis of fossil animal remains is a power- ful tool for investigating ancient environments, ecosystems and feeding behaviours, and is a method that can be used to directly investigate dif- ferences in herbivore dietary specialisation. Stable isotope techniques Palaeogeography, Palaeoclimatology, Palaeoecology 333-334 (2012) 168–177 ⁎ Corresponding author at: Department of Archaeology, University of Aberdeen, St. Mary's Building, Elphinstone Road, Aberdeen, AB24 3UF, UK. Tel.: + 44 1224 273823. E-mail addresses: kate_britton@eva.mpg.de, k.britton@abdn.ac.uk (K. Britton), gaudzinski@rgzm.de (S. Gaudzinski-Windheuser), w.roebroeks@arch.leidenuniv.nl (W. Roebroeks), kindler@rgzm.de (L. Kindler), richards@eva.mpg.de, michael.richards@ubc.ca (M.P. Richards). 1 Current address: Department of Archaeology, University of Aberdeen, St. Mary's Building, Elphinstone Road, Aberdeen, AB24 3UF, UK. 0031-0182/$ – see front matter © 2012 Elsevier B.V. All rights reserved. doi:10.1016/j.palaeo.2012.03.028 Contents lists available at SciVerse ScienceDirect Palaeogeography, Palaeoclimatology, Palaeoecology journal homepage: www.elsevier.com/locate/palaeo