The biogeography of non-marine molluscs in the Tuscan Archipelago reveals combined effects of current eco-geographical drivers and paleogeography

We investigated the role of present (Recent) and historical (Pleistocene, i.e., Wurmian Last Glacial Maximum) eco-geographical variables on the richness and diversity of non-marine molluscs in the Tuscan Archipelago, as well as inter-island faunal dissimilarity and relationships with source pools (Sardinia and Corsica, Tuscany). The association between species richness and present and historical eco-geographical variables were assessed with Spearman's rank correlation test, while faunal dissimilarity both between islands and with their source pools was analyzed through beta-diversity partitioning (SOrensen index and its nestedness and turnover component) with UPGMA clustering tested with a multiscale bootstrap procedure. Non-metric multidimensional scaling in RGB color space was also used. Multiple regressions on distance matrices were then applied to explain assemblage composition between islands. Analyses were performed on all species and on all species except aliens. The overall framework showed the combined effects of current eco-geographical and paleogeographical imprints on non-marine malacofauna in the Tuscan Archipelago. However, excluding aliens, differences in species spatial turnover showed a clear correlation with Pleistocene inter-island distances, evidence of stronger historical biogeographical relationships between islands. This may indicate that widespread native species established their distribution during the Pleistocene, while alien species spread into the Tuscan Archipelago through stochastic and human-mediated dispersion events in recent times. Interestingly, Giglio's relationships do not agree with the most accepted paleogeographical model, suggesting that this island might have been connected to the Tuscan mainland during the Wurmian Last Glacial Maximum. An in-depth revision of the paleogeographic framework of the northern Tyrrhenian is therefore called for.