ORIGINAL MANUSCRIPT Phylogeographic patterns of Buthus scorpions (Scorpiones: Buthidae) in the Maghreb and South-Western Europe based on CO1 mtDNA sequences P. Sousa 1,3 , D. J. Harris 1,3 , E. Froufe 2 & A. van der Meijden 1 1 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Vila do Conde, Portugal 2 CIIMAR, Centro Interdisciplinar de Investigação Marinha e Ambiental, Rua dos Bragas, Porto, Portugal 3 Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, Porto, Portugal Keywords Buthus; Scorpiones; Western Mediterranean; CO1; cryptic diversity; phylogeography. Correspondence Pedro Sousa, CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Campus Agrário de Vairão, Rua Padre Armando Quintas, 7 – Crasto, Vairão 4485-661, Portugal. Email: prsousa@gmail.com Editor: Mark-Oliver Rödel Received 31 December 2011; revised 13 April 2012; accepted 13 April 2012 doi:10.1111/j.1469-7998.2012.00925.x Abstract The genus Buthus is a medium diverse scorpion genus, with 35 species distributed from Portugal and Morocco ranging eastward to Yemen in the Arabic Peninsula. The bulk of the genus’ known species diversity occurs in the Western Mediterra- nean area. A recent molecular study started to elucidate the patterns of diversity of this genus in the Iberian Peninsula and the Maghreb. Since then, the taxonomy of the genus has changed substantially, with several new species having been described, and with the elevation of former subspecies to species-level. In this study, we assessed the patterns of diversity of Buthus scorpions from across the Maghreb region of North Africa using CO1 DNA sequence data. Based on our dataset of 147 sequences, including 67 new sequences, we recovered four well- supported deep clades within Buthus scorpions from the Maghreb and Southern Europe. This further strengthens the support for cryptic diversity in the Maghreb region. The broader sampling of the Maghreb permitted a better understanding of the phylogeographic structure in this area. Three clades were restricted to Morocco and appear to have originated at the Atlantic Coast of this country, while the fourth was found throughout the region. We propose a model with two colonizing events to explain the distribution patterns across the Strait of Gibral- tar, with an initial colonization from North Africa to Iberia followed by a rein- vasion of the Rif Mountains region in Morocco. Introduction The scorpion genus Buthus Leach, 1815 (Scorpiones, Buthi- dae) received very little taxonomic attention during the last half of the 20th century. In 1952, Vachon greatly modified the genus’ taxonomy, transforming a poorly defined and hetero- geneous group into a morphologically uniform coherent genus. Few changes were then made until Fet & Lowe (2000) suppressed all infrasubspecific varieties created by Vachon, reducing the number of taxa to five species and 11 subspecies (two of B. atlantis Pocock, 1889 and nine of B. occitanus Amoreux, 1789). However, since then, the genus has gained 30 new species, with the elevation of several subspecies to species level, the redescription of suppressed varieties and with the discovery of new material. The genus is now known from Southern Europe, Iberia and the South of France, Africa, in all countries bordering the Mediterranean, plus Guinea, Mau- ritania, Niger, Senegal and Sudan (south of the Sahara Desert) and Eritrea, Ethiopia and Somalia in the Horn of Africa, and the Middle East, in Cyprus, Egypt (Sinai), Israel, Jordan and Yemen (Fet & Lowe, 2000; Lourenço, 2002, 2003, 2005a,b, 2008; Lourenço & Slimani, 2004; Lourenço & Vachon, 2004; Lourenço & Geniez, 2005; Kovar ˇík, 2006, 2011; Lourenço & Qi, 2006; Lourenço, Sun & Zhu, 2009; Lourenço, Yag ˘mur & Duhem, 2010; Touloun & Boumez- zough, 2011; Yag ˘ mur, Koç & Lourenço, 2011). The genus has been cited for other countries such as Djibouti, Gambia, Ghana, Iraq and Lebanon, but these records require addi- tional confirmation (Fet & Lowe, 2000). The distribution of the 35 species of Buthus is not uniform across the range of the genus. Twenty-one species are endemic to the Western Mediterranean area alone, although this is also historically the most surveyed area were the genus is known to occur. Morocco has 13 endemic species and shares two more with Algeria and Tunisia, with the latter country adding two endemic species to this region (Lourenço, 2002; Kovar ˇík, 2006). Additionally, three more are known from the Iberian Peninsula and South of France. Still much remains to be Journal of Zoology Journal of Zoology. Print ISSN 0952-8369 Journal of Zoology •• (2012) ••–•• © 2012 The Authors. Journal of Zoology © 2012 The Zoological Society of London 1