A misinterpreted disjunction: the phylogenetic relationships of the North African land snail Gyrostomella (Gastropoda: Stylommatophora: Helicidae)

Abstract

Abstract The North African helicid Gyrostomella has been considered closely related to Levantina, suggesting a disjunct distribution with a distribution gap of almost 2000 km in North Africa. We studied this disjunction and the systematics of Helicinae using molecular markers. Our analyses indicate an affiliation of Gyrostomella with the radiation of Helicinae in the Maghreb, especially with Maurohelix. The occurrence of Gyrostomella in Libya does not represent a western outpost of Helicini in North Africa, but an eastern outpost of the Maghreb radiation of Helicinae. The disjunction of Gyrostomella in Libya and Maurohelix in Algeria may be the result of a long-distance dispersal or may represent relicts of a larger range of the lineage in the past as a consequence of aridification processes in North Africa. Taking into account biogeographic patterns, we suggest a subdivision of Helicinae into four tribes: Allognathini, Helicini, Maculariini trib. nov. and Thebini. The Eastern Mediterranean Helicini diverged from the Western Mediterranean clade in the Eocene. The Western Mediterranean clade then split into Allognathini in the Iberian Peninsula, Maculariini in the region of the future western Alps and Thebini on terranes probably including the Rif-Betic and Kabylies blocks that now form part of the Maghreb region.