Metapopulation genetic structure and migration pathways in the land snail Helix aspersa:: influence of landscape heterogeneity

The spatial genetic structuring of the land snail Helix aspersa was investigated for 32 colonies within an intensive agricultural area, the polders of the Bay of Mont-Saint-Michel ( France). Given the habitat patchiness and environmental instability, the setting of H. aspersa colonies meets the broader view of a metapopulation structure. The identification of extrinsic barriers to migration and their impact on the genetic distribution was addressed through the genotyping of 580 individuals using a combined set of enzyme and microsatellite loci. To evaluate the distance as well as the direction over which the spatial genetic arrangement occurs, two-dimensional spatial autocorrelation analyses, Mantel tests of association and multivariate Mantel correlograms were used. Different connectivity networks and geographical distances based on landscape features were constructed to evaluate the effect of environmental heterogeneity and to test the adequacy of an isolation by distance model on the distribution of the genetic variability. Genetic divergence was assessed using either classical IAM-based statistics, or SMM-based genetic distances specifically designed to accommodate the mutational processes thought to fit microsatellite evolution (IAM: Infinite Allele Model; SMM: Stepwise Mutation Model). Genetic distances based only on genetic drift yielded the most plausible biologically meaningful interpretation of the observed spatial structure. Applying a landscape-based geographical distance which postulates that migration arises along roadside verges, hedges or irrigation canal embankments gave a better fit to an isolation by distance model than did a simple Euclidean distance. The progressive decline of genetic similarity with physical distance appeared to be environmentally induced, leading to functional migration pathways.