Macrogeographic genetic variability in the gastropod mollusk Littorina sitkana from the northwest Pacific

Variation at four highly polymorphic allozyme loci (inorganic pyrophosphatase, peptidase, and two esterase loci) was examined in 25 settlements of the marine snail Littorina sitkana (Mollusca, Gastropoda). The sampling localities covered a wide part of the species range: from the Peter the Great Bay (the Sea of Japan) at the southwest to the Mednyi Island (Commander Islands) at the northeast. Like other littorines lacking the pelagic stage, L. sitkana was characterized by significant genetic differentiation (G(ST) for the pooled sample was 0.310). Cluster analysis and nonmetric multidimensional scaling conducted on a matrix of pairwise genetic distances between all of the settlements studied revealed four genetically different groups: southern Primorye, northern Prymorye, Sakhalin, and Kuril-Commanders. The population-genetic structure of the L. sitkana settlements is close to that described by the isolation-by-distance and stepping-stone models: the geographic and the genetic distances between the most settlements examined are distinctly correlated.