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The lively debate about speciation currently focuses on the relative importance of factors driving population differentiation. While many studies are increasingly producing results on the importance of selection, little is known about the interaction between drift and selection. Moreover, there is still little knowledge on the spatial-temporal scales at which speciation occurs, that is, arrangement of habitat patches, abruptness of habitat transitions, climate and habitat changes interacting with selective forces. To investigate these questions, we quantified variation on a fine geographical scale analysing morphological (shell) and genetic data sets coupled with environmental data in the land snail Murella muralis, endemic to the Mediterranean island of Sicily. Analysis of a fragment of the mitochondrial DNA cytochrome oxidase I gene (COI) and eight nuclear microsatellite loci showed that genetic variation is highly structured at a very fine spatial scale by local palaeogeographical events and historical population dynamics. Molecular clock estimates, calibrated here specifically for Tyrrhenian land snails, provided a framework of palaeogeographical events responsible for the observed geographical variations and migration routes. Finally, we showed for the first time well-documented lines of evidence of selection in the past, which explains divergence of land snail shell shapes. We suggest that time and palaeogeographical history acted as constraints in the progress along the ecological speciation continuum. Our study shows that testing for correlation among palaeogeography, morphology and genetic data on a fine geographical scale provides information fundamental for a detailed understanding of ecological speciation processes.
Talitrids are the only family within the order Amphipoda to have colonised supralittoral and terrestrial environments. They live in a variety of settings, from sandy to rocky and pebble beaches, to river and lake banks, and to leaf litter and caves. A common feature is the absence of a planktonic larval stage to facilitate passive dispersal over long-distances. However, some species have broad distributions. Genetic studies over the past 25 years have tried to explain this apparent contradiction by assessing patterns of species genetic structuring on different geographical scales. Here, we review the molecular studies available to date and focus on the population genetics of talitrids. Most of these studies considered populations in the Mediterranean area, but also along the Atlantic coast and in Canary Island caves. From this review, the group emerges as a potential model to understand processes of dispersal and divergence in non-highly-vagile supralittoral organisms. At the same time, studies on these issues are still too restricted geographically: a worldwide scale including different regions would provide us with a better perspective on these problems.