Molecular phylogeny and trait evolution of Madeiran land snails: radiation of the Geomitrini (Stylommatophora: Helicoidea: Geomitridae).

The Geomitrini is the most species-rich group of land snails in the Madeiran Archipelago. The phylogeny of the group is reconstructed based on mitochondrial and nuclear genetic markers. The timing of diversification, the colonisation history of the islands of the Madeiran Archipelago and the evolution of characters of the dart apparatus are studied. The results of the phylogenetic analyses confirm the sister group relationship of Geomitrini and Cochlicellini, but also show that several previously accepted genus-group taxa are not monophyletic. A new classification for the Geomitrini is proposed, including the description of two new genera, Domunculifex Brozzo, De Mattia, Harl & Neiber, n. gen. and Testudodiscula Brozzo, De Mattia, Harl & Neiber, n. gen. The onset of diversification of Geomitrini was dated in our analysis at 13 Ma, which largely coincides with the emergence of the present-day islands. The ancestral state estimation recovered the presence of two appendiculae in the reproductive system as the ancestral state in Geomitrini. One appendicula was lost three times independently within the tribe and is even missing completely in one group. The ancestral area estimation suggests recurrent colonisations of Madeira (and the Ilhas Desertas) from the older island Porto Santo.