Acta Tropica 134 (2014) 43–51 Contents lists available at ScienceDirect Acta Tropica jo ur nal home p age: www.elsevier.com/locate/actatropica Deep intraspecific divergences in the medically relevant fat-tailed scorpions (Androctonus, Scorpiones) P. Coelho a , P. Sousa a,b , D.J. Harris a,b , A. van der Meijden a,∗ a CIBIO, InBio, Centro de Investigac ¸ ão em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de Vairão, Vairão 4485-661, Portugal b Departamento de Biologia, Faculdade de Ciências, Rua do Campo Alegre, Porto FC4 4169-007, Portugal a r t i c l e i n f o Article history: Received 9 October 2013 Received in revised form 28 January 2014 Accepted 1 February 2014 Available online 11 February 2014 Keywords: Scorpionism Androctonus Scorpions Phylogeny Biogeography Cryptic diversity a b s t r a c t The genus Androctonus, commonly known as fat-tailed scorpions, contains 22 species distributed from Togo and Mauritania in the west, North Africa, through the Middle East and to as far east as India. With 13 species, a substantial amount of this genus’ diversity occurs in North Africa, which is a major hotspot of scorpion sting incidents. Androctonus are among the most medically relevant animals in North Africa. Since venom composition within species is known to vary regionally, the improvement of therapeutic management depends on a correct assessment of the existing regional specific and sub-specific variation. In this study, we assessed the phylogeographical patterns in six species of Androctonus scorpions from North Africa using mitochondrial DNA markers. We sequenced COX1, 12S, 16S and ND1 genes from 110 individuals. Despite lacking basal resolution in the tree, we found taxonomical and geographically coherent clades. We discovered deep intraspecific variation in the widespread Androctonus amoreuxi and Androctonus australis, which consisted of several well-supported clades. Genetic distances between some of these clades are as high as those found between species. North African A. australis have a deep split in Tunisia around the Chott el-Djerid salt-lake. A novel split between A. amoreuxi scorpions was found in Morocco. We also found deep divergences in Androctonus mauritanicus, corresponding to areas attributed to invalidated subspecies. In addition we uncovered a clade of specimens from coastal south Morocco, which could not be ascribed to any know species using morphological characters. Based on these findings we recommend a reassessment of venom potency and anti-venom efficacy between these deep intraspecific divergent clades. © 2014 Elsevier B.V. All rights reserved. 1. Introduction Worldwide, 1.2 million people are stung by scorpions every year. Scorpionism, defined as the severe to lethal incident as a conse- quence of a scorpion sting (Lourenc ¸ o and Cuellar, 1995) may be responsible for 3250 global annual mortalities which are mostly concentrated in a few high-risk areas (Chippaux and Goyffon, 2008). North Africa in particular is considered a high-risk area for scorpionism (Chippaux and Goyffon, 2008), with the genera Leiu- rus Ehrenberg, 1828 and Androctonus Ehrenberg, 1828 being the foremost cause of serious envenomation in this area (Goyffon and Guette, 2005; Graham, 2011; Habermehl, 1994). Five Androctonus species are considered as dangerous to man, particularly Androc- tonus mauritanicus (Pocock, 1902) and thewidespread Androctonus ∗ Corresponding author. Tel.: +351 916712100. E-mail address: mail@arievandermeijden.nl (A. van der Meijden). australis (Linnaeus, 1758), which are the most dangerous Androc- tonus in the Maghreb region (Morocco, Algeria, Tunisia) (Goyffon and Guette, 2005). A. australis is known for envenomating humans and possessing a high toxicity (LD 50 = 0.32 mg/kg in mice; Watt and Simard, 1984). For this reason, A. australis was one of the first species of scorpions to have its venom purified for neurotoxin char- acterization (Miranda et al., 1966). As in snakes (Daltry et al., 1996; Prasad et al., 1999), scorpion venom is known to have consider- able intraspecific regional variation in composition (Devaux et al., 2004; El Ayeb and Rochat, 1985; Newton et al., 2007; Smertenko et al., 2001), and thus a different response to antivenom treatment (Omran and McVean, 2000). Furthermore, other species such as Androctonus amoreuxi (Audouin, 1826) may also cause more cases of scorpionism than currently thought (Goyffon et al., 2012). It is therefore important to study the phylogeographical patterns of Androctonus over a great part of their distribution as it may have direct applications in therapeutic management. Androctonus is present in deserts and semi-arid regions from Togo to Morocco in the Atlantic coast of Africa (Lourenc ¸ o and http://dx.doi.org/10.1016/j.actatropica.2014.02.002 0001-706X/© 2014 Elsevier B.V. All rights reserved.