Phylogeny and evolution of the land snail tribe Clausiliini (Gastropoda: Clausiliidae)

The phylogenetic relationships within the land snail group Clausiliini were analysed based on an almost complete taxon sampling (28 of 29 extant species) and a set of mitochondrial and nuclear markers. The phylogenetic analyses and character state reconstructions revealed several trends in the group that resulted in repeated convergences. The N-type closing apparatus was convergently transformed into a G-type clausilial apparatus in Graciliaria, Ruthenica + Micridyla and probably three times within Neostyriaca. There were parallel changes in the length of the male copulatory organs and repeated reductions of the basal keel in several lineages. Such trends resulting in repeated convergences are characteristic for several radiations of stylommatophoran land snails. They may reflect parallel adaptations of related lineages to similar environmental constraints. However, it is also possible that some of these transformations are not adaptive, but represent random changes in a restricted morphospace. We explored the timing of the Clausiliini radiation using different calibration strategies. Despite large numbers of fossils of Clausiliidae from Western and Central Europe, the results of these analyses and the fossil evidence for a Cretaceous origin of the Clausiliidae indicate that there are large gaps in the fossil record of the Clausiliini. Especially the origin and early evolution of the Clausiliini in Southeastern Europe is obscured by our insufficient knowledge of Neogene deposits there.