Out of Africa: Phylogeny and biogeography of the widespread genus Acanthodactylus (Reptilia: Lacertidae) Karin Tamar a,b,⇑ , Salvador Carranza c , Roberto Sindaco d , Jir ˇí Moravec e , Jean-François Trape f , Shai Meiri a,b a Department of Zoology, George S. Wise Faculty of Life Sciences, Tel Aviv University, Tel Aviv 6997801, Israel b The Steinhardt Museum of Natural History, Israel National Center for Biodiversity Studies, Tel Aviv University, Tel Aviv 6997801, Israel c Institut de Biologia Evolutiva (CSIC-Universitat Pompeu Fabra), Passeig Maritím de la Barceloneta, 37-49, 08003 Barcelona, Spain d Museo Civico de Storia Naturale, via San Francesco di Sales 188, I-10022 Carmagnola, Torino, Italy e Department of Zoology, National Museum, Cirkusová 1740, 19300 Prague 9, Czech Republic f Laboratoire de paludologie et zoologie médicale, Institut de Recherche pour le Développement, UMR MIVEGEC, B.P. 1386, Dakar, Senegal article info Article history: Received 15 February 2016 Revised 1 June 2016 Accepted 5 July 2016 Available online 6 July 2016 Keywords: Arabia Diversification Molecular clock Phylogeography Systematics Taxonomy abstract Acanthodactylus lizards are among the most diverse and widespread diurnal reptiles in the arid regions spanning from North Africa across to western India. Acanthodactylus constitutes the most species-rich genus in the family Lacertidae, with over 40 recognized species inhabiting a wide variety of dry habitats. The genus has seldom undergone taxonomic revisions, and although there are a number of described spe- cies and species-groups, their boundaries, as well as their interspecific relationships, remain largely unre- solved. We constructed a multilocus phylogeny, combining data from two mitochondrial (12S, cytb) and three nuclear (MC1R, ACM4, c-mos) markers for 302 individuals belonging to 36 known species, providing the first large-scale time-calibrated molecular phylogeny of the genus. We evaluated phylogenetic rela- tionships between and within species-groups, and assessed Acanthodactylus biogeography across its known range. Acanthodactylus cladogenesis is estimated to have originated in Africa due to vicariance and dispersal events from the Oligocene onwards. Radiation started with the separation into three clades: the Western and scutellatus clades largely distributed in North Africa, and the Eastern clade occurring mostly in south-west Asia. Most Acanthodactylus species diverged during the Miocene, possibly as a result of regional geological instability and climatic changes. We support most of the current taxonomic classi- fications and phylogenetic relationships, and provide genetic validity for most species. We reveal a new distinct blanfordii species-group, suggest new phylogenetic positions (A. hardyi, A. masirae), and syn- onymize several species and subspecies (A. lineomaculatus, A. boskianus khattensis and A. b. nigeriensis) with their phylogenetically closely-related species. We recommend a thorough systematic revision of taxa, such as A. guineensis, A. grandis, A. dumerilii, A. senegalensis and the pardalis and erythrurus species-groups, which exhibit high levels of intraspecific variability, and clear evidence of phylogenetic complexity. Ó 2016 Elsevier Inc. All rights reserved. 1. Introduction The Old World genus Acanthodactylus Fitzinger, 1834 is the most species-rich genus in the family Lacertidae, comprising over 40 currently recognized species (Uetz and Hošek, 2016). These lizards are commonly known as fringe-fingered lizards due to their distinctive lateral finger scalation. Acanthodactylus is a member of the Saharo-Eurasian clade within the Eremiadini tribe (Mayer and Pavlicev, 2007), and along with its phylogenetically closest members (i.e., Eremias, Mesalina, Ophisops; Pyron et al., 2013), it inhabits mostly xeric habitats in North Africa and Asia (Sindaco and Jeremc ˇenko, 2008). Acanthodactylus ranges from the Iberian Peninsula, across North Africa (including the Sahel) and the Arabian Peninsula towards western India, and northward to Cyprus and southern Turkey (Fig. 1; Salvador, 1982; Arnold, 1983; Schleich et al., 1996; Sindaco and Jeremc ˇenko, 2008). Acanthodactylus are diurnal, ground-dwelling, medium-sized lizards, occurring in several climatic regions, from the Mediterranean ecoregion to the harsh desert environments of the Sahara and Arabia, occupying a wide array of arid ecosystems from open woodland, shrubland and savanna to sand dune deserts (Salvador, 1982; Arnold, 1983; Sindaco and Jeremc ˇenko, 2008). http://dx.doi.org/10.1016/j.ympev.2016.07.003 1055-7903/Ó 2016 Elsevier Inc. All rights reserved. ⇑ Corresponding author at: Department of Zoology, George S. Wise Faculty of Life Sciences, Tel Aviv University, Tel Aviv 6997801, Israel. E-mail address: karintmr@gmail.com (K. Tamar). Molecular Phylogenetics and Evolution 103 (2016) 6–18 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev