1 3 Mitochondrial phylogeny shows multiply independent ecological 4 transitions and northern dispersion despite of Pleistocene glaciations 5 in meadow and steppe vipers (Vipera ursinii and Vipera renardi) 6 7 8 Oleksandr Zinenko a,b,c,⇑ Q1 , Nikolaus Stümpel b , Lyudmila Mazanaeva d , Andrey Bakiev e , 9 Konstantin Shiryaev f , Aleksey Pavlov g , Tatiana Kotenko h , Oleg Kukushkin i , Yury Chikin j , 10 Tatiana Duisebajeva k , Göran Nilson l , Nikolai L. Orlov m , Sako Tuniyev n , Natalia B. Ananjeva m , 11 Robert W. Murphy o,p , Ulrich Joger b 12 a The Museum of Nature at V.N. Karazin Kharkiv National University, Trinkler str. 8, Kharkiv 61058, Ukraine Q2 13 b Staatliches Naturhistorisches Museum Braunschweig, Gausstrasse 22, Braunschweig D-38106, Germany 14 c National Park ‘‘Dvorichansky’’, Privokzalna str. 51, Dvorichna, Kharkiv Oblast 62701, Ukraine 15 d Dagestan State University, Faculty of Biology, Department of Zoology, apt. 13, 37a, M. Gadzhiyeva st., Makhachkala, Dagestan 367025, Russia 16 e Institute of Ecology of the Volga River Basin of Russian Academy of Science, Komzina str. 10, Togliatti 445003, Russia 17 f Tula Regional Exotarium, Oktyabr’skaya str. 26, Tula 300002, Russia 18 g Volzhsko-Kamsky National Nature Biosphere Rezerve, Vekchnik str., 1, Sadovyi set. Zelenodolsk distr., Tatarstan Republic 42253, Russia 19 h The Shmalgauzen Institute of Zoology, National Academy of Science of Ukraine, B. Khemlnits’kogo st., 15, Kyiv-30, 01601, Ukraine 20 i Karadagh Nature Reserve of Ukrainian National Academy of Sciences, Nauki str., 24, Theodosia 98188, AR Crimea, Ukraine 21 j Institute of Genofond of Animals and Plants, Durmon-yuli str., 32, Toshkent, Uzbekistan 22 k Institute of Zoology, al-Farabi Av., 93, Almaty 050060, Kazakhstan 23 l Göteborg Natural History Museum, Box 7283, SE-402 35 Göteborg, Sweden 24 m Zoological Institute, Russian Academy of Science, Universitetskaya nab., 1, St. Petersburg 199034, Russia 25 n Federal State Institution Sochi National Park, Ul. Moskovskaya, 13, Sochi, Krasnodarsky Krai 354000, Russia 26 o State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, The Chinese Academy of Sciences, Kunming 650223, China 27 p Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6, Canada 28 29 30 32 article info 33 Article history: 34 Received 22 November 2013 35 Revised 20 October 2014 36 Accepted 9 December 2014 37 Available online xxxx 38 Keywords: 39 Mitochondrial phylogeny 40 Historical demography 41 Pleistocene 42 Vipera ursinii complex 43 Vipera renardi complex 44 45 abstract 46 The phylogeny and historical demography of small Eurasian vipers of the Vipera ursinii and V. renardi 47 complexes were studied using mitochondrial DNA sequences analysed with Bayesian inference, Maxi- 48 mum Likelihood and Maximum Parsimony approaches, and mismatch distributions. Diversification in 49 the group resulted from an initial dispersion in the later Pliocene – Pleistocene in two directions: 50 north-westwards via the Balkans (V. ursinii complex) and north-eastwards from Asia Minor via the Cau- 51 casus (V. renardi complex). An independent, comparatively recent transition occurred from montane hab- 52 itats to lowland grasslands in different mitochondrial lineages during the Late Pleistocene, when 53 representatives of both complexes had reached lowland steppes to the north. Effective population size 54 showed clear signs of rapid growth in eastern V. renardi, triggered by colonization of vast lowland 55 steppes, but in western V. ursinii complex grew during the Last Glaciation and experienced stabilization 56 in Holocene. Expansion and population growth in lowland lineages of V. renardi was not strongly affected 57 by Pleistocene climatic oscillations, when cold, dry conditions could have favoured species living in open 58 grasslands. The high diversity of closely related haplotypes in the Caucasus and Tien-Shan could have 59 resulted from repetitive expansion-constriction-isolation events in montane regions during Pleistocene 60 climate fluctuations. The mitochondrial phylogeny pattern conflicts with the current taxonomy. 61 Ó 2014 Elsevier Inc. All rights reserved. 62 63 64 65 1. Q4 Introduction 66 Pleistocene climatic fluctuations profoundly affected the distri- 67 butions and structure of most European species (Hewitt, 1999; 68 Taberlet et al., 1998; Nieto, 2011). Traditional biogeographic http://dx.doi.org/10.1016/j.ympev.2014.12.005 1055-7903/Ó 2014 Elsevier Inc. All rights reserved. ⇑ Corresponding author at: The Museum of Nature at V.N. Karazin Kharkiv National University, Trinkler str. 8, Kharkiv 61058, Ukraine. E-mail address: zinenkoa@yahoo.com (O. Zinenko). Molecular Phylogenetics and Evolution xxx (2014) xxx–xxx Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev YMPEV 5085 No. of Pages 17, Model 5G 20 December 2014 Please cite this article in press as: Zinenko, O., et al. Mitochondrial phylogeny shows multiply independent ecological transitions and northern dispersion despite of Pleistocene glaciations in meadow and steppe vipers (Vipera ursinii and Vipera renardi). Mol. Phylogenet. Evol. (2014), http://dx.doi.org/10.1016/ j.ympev.2014.12.005