ORIGINAL ARTICLE Subspecies dynamics in space and time: A study of the red deer complex using ancient and modern DNA and morphology Meirav Meiri 1,2 | Pavel Kosintsev 3 | Keziah Conroy 4 | Shai Meiri 1,5 | Ian Barnes 6 | Adrian Lister 6 1 The Steinhardt Museum of Natural History, Tel Aviv University, Tel Aviv, Israel 2 Institute of Archaeology, Tel Aviv University, Tel Aviv, Israel 3 Institute of Plant and Animal Ecology, Urals Branch of the Russian Academy of Sciences, Ekaterinburg 620144, Russia 4 Leverhulme Centre for Human Evolutionary Studies, University of Cambridge, Cambridge, UK 5 Department of Zoology, Tel Aviv University, Tel Aviv 69978, Israel 6 Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK Correspondence Adrian Lister, Department of Earth Sciences, Natural History Museum, London, UK. Email: a.lister@nhm.ac.uk Funding information NERC, Grant/Award Number: NE/G00269X/ 1, NF/2009/1/18; NERC/AHRC/ORAU, Grant/Award Number: NE/G00269X/1, NF/ 2009/1/18 Editor: Jenny McGuire Abstract Aim: The status of geographical units within species and species complexes is debated for many taxa, with many molecular studies failing to detect phenotypically defined subspecies. The history and longevity of geographical patterns are also gen- erally very poorly understood. We examine Holarctic red deer (Cervus elaphus and related forms), incorporating ancient DNA to ask whether the present phylogeogra- phy has persisted through climatic perturbations or is a relatively recent phe- nomenon. Location: Holarctic (Europe, northern Asia and North America). Methods: We obtained 21 modern and 30 Holocene and Late Pleistocene samples, which together with published data resulted in 180 individuals spanning 21 nominal extant and one extinct subspecies. Phylogenetic analyses were carried out on 748- bp of mitochondrial DNA (cytochrome b and control region). Where possible, the morphology of DNA-yielding ancient samples was examined to assess subspecies identity. Results: Major clades within the red deer complex are upheld, but subspecies within them receive varying support. The ancient phylogeographical structure con- forms in significant part to the modern situation, but some haplogroups no longer survive. Moreover, there have been substantial shifts in geographical ranges through time. Wapitoids spread as far west as Romania in the last glaciation, and elaphoids reached eastward to the Ural Mountains. A possible contact zone between the two lineages stretched from the Urals through the Crimea to Eastern Europe. Main conclusions: Ancient DNA and morphology are strongly complementary in elucidating population history. Through the past 50 kyr, the major lineages of red deer, and some of the subspecies groups within them, have maintained their genetic and morphological integrity and their core geographical distributions, despite range expansions and contractions and likely contact between the haplogroups (with potential for hybridization). KEYWORDS ancient DNA, Cervus canadensis, Cervus elaphus, Cervus hanglu, mitochondrial DNA, morphology, phylogeography, red deer DOI: 10.1111/jbi.13124 Journal of Biogeography. 2018;45:367–380. wileyonlinelibrary.com/journal/jbi © 2017 John Wiley & Sons Ltd | 367