Late-glacial recolonization and phylogeography of European red deer (Cervus elaphus L.) MEIRAV MEIRI,* ADRIAN M. LISTER, † THOMAS F. G. HIGHAM, ‡ JOHN R. STEWART, § LAWRENCE G. STRAUS, ¶ ** HENRIETTE OBERMAIER, †† MANUEL R. GONZ  ALEZ MORALES,** ANA B. MAR  IN-ARROYO** and IAN BARNES ‡‡ *Department of Zoology, Institute of Archaeology, Tel Aviv University, Tel Aviv 69978, Israel, †Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK, ‡Research Lab for Archaeology and the History of Art, University of Oxford, Oxford OX1 3QY, UK, §School of Applied Sciences, Bournemouth University, Poole, Dorset BH12 5BB, UK, ¶Department of Anthropology, University of New Mexico, Albuquerque, NM 87131-0001, USA, **Instituto Internacional de Investigaciones Prehist oricas de Cantabria, Universidad de Cantabria, Santander 39005, Spain, ††Bavarian State Collection for Anthropology and Palaeoanatomy Munich, Munich 80539, Germany, ‡‡School of Biological Sciences, Royal Holloway, University of London, Egham, Surrey TW20 0EX, UK Abstract The Pleistocene was an epoch of extreme climatic and environmental changes. How individual species responded to the repeated cycles of warm and cold stages is a major topic of debate. For the European fauna and flora, an expansion–contraction model has been suggested, whereby temperate species were restricted to southern refugia during glacial times and expanded northwards during interglacials, including the present interglacial (Holocene). Here, we test this model on the red deer (Cervus elaphus)a large and highly mobile herbivore, using both modern and ancient mitochondrial DNA from the entire European range of the species over the last c. 40 000 years. Our results indicate that this species was sensitive to the effects of climate change. Prior to the Last Glacial Maximum (LGM) haplogroups restricted today to South-East Europe and Western Asia reached as far west as the UK. During the LGM, red deer was mainly restricted to southern refugia, in Iberia, the Balkans and possibly in Italy and South-Western Asia. At the end of the LGM, red deer expanded from the Iberian refu- gium, to Central and Northern Europe, including the UK, Belgium, Scandinavia, Ger- many, Poland and Belarus. Ancient DNA data cannot rule out refugial survival of red deer in North-West Europe through the LGM. Had such deer survived, though, they were replaced by deer migrating from Iberia at the end of the glacial. The Balkans served as a separate LGM refugium and were probably connected to Western Asia with genetic exchange between the two areas. Keywords: ancient DNA, contraction–expansion model, LGM, phylogeography, red deer, Southern refugia Received 15 March 2013; revision received 9 June 2013; accepted 11 June 2013 Introduction Climate oscillations during the Pleistocene have signifi- cantly affected the distribution of species. For Europe, in particular, a well-defined model has been established (e.g. Hewitt 1996; Taberlet et al. 1998; Lister 2004; Randi 2007; Provan & Bennett 2008; Hofreiter & Stewart 2009; Stewart et al. 2010). This model proposes that during glaciations, temperate species were restricted to south- ern refugia, principally in the Iberian, Italian and Bal- kan peninsulas (Hewitt 1996, 1999; Taberlet et al. 1998; Hewitt et al. 2004). According to this model, at the onset of glacial conditions, northern populations either went extinct or retreated southward, with southern popula- tions subsequently expanding northwards during inter- Correspondence: Meirav Meiri, Fax: 972 3 6409817; E-mail: meirav.meiri@gmail.com © 2013 John Wiley & Sons Ltd Molecular Ecology (2013) 22, 4711–4722 doi: 10.1111/mec.12420