Phylogeny, biogeography, and evolution of two Mediterranean snakes, Malpolon monspessulanus and Hemorrhois hippocrepis (Squamata, Colubridae), using mtDNA sequences

Variation in 815 bp of mitochondrial DNA from two gene fragments (300 bp of cytochrome b and 395–515 bp of 12S rRNA) for 26 Malpolon monspessulanus, and cytochrome b for a further 21 individuals, indicates that this species originated in the Maghreb area of Northwest Africa. Here, an estimated 3.5–6 Mya, it divided into the western M. m. monspessulanus, and an eastern clade including M. m. insignitus and M. m. fuscus. The very limited genetic differentiation between Maghreb and Southwest European populations of this form suggests that it arrived in the Iberian Peninsula only recently. Population genetics and demographic tests indicate subsequent expansion in this area around 83,000–168,000 year ago. Because present populations of Malpolon arrived recently, mid-Pliocene and at least some Pleistocene fossils of the genus Malpolon in Southwest Europe are probably derived from an earlier invasion from the Maghreb, possibly as early as the end of the Miocene period, 5.3–5.9 Mya, when there was a temporary land bridge across the site of the Strait of Gibraltar and the Mediterranean Sea desiccated. The descendants of this earlier invasion must have eventually become extinct, perhaps during one of the Pleistocene glaciations. In contrast to the western M. m. monspessulanus, the greater genetic divergence found in the eastern clade of M. monspessulanus suggests that it dispersed at an earlier date and probably over a longer period, spreading eastwards through northern Libya and Egypt to Syria, Iraq, and Iran, and around the Mediterranean Sea through Turkey into the Aegean archipelagos and the Balkan peninsula. The western and eastern units of M. monspessulanus have different dorsal color pattern, differences in skull structure and exhibit an 8.4% uncorrected genetic divergence in the combined gene fragments investigated here. It is consequently recommended that they should be treated as separate species: M. monspessulanus (sensu stricto) and Malpolon insignitus stat. nov., the latter including the subspecies Malpolon insignitus fuscus comb. nov. The same combined mitochondrial gene fragments used in Malpolon were investigated in 20 individuals of Hemorrhois hippocrepis, and of cytochrome b alone in a further 17. They indicate that this species also originated in the Maghreb and again invaded the Iberian Peninsula quite recently. Some of the most recent invasions of the Iberian Peninsula by reptiles and amphibian taxa could probably be anthropogenic in origin. Some other species including M. monspessulanus and H. hippocrepis, may have crossed naturally, by 'hopping' across the Strait of Gibraltar via temporary islands on the shallowest parts that were exposed during sea-level fall associated with Pleistocene glaciations.