Mitochondrial DNA variation in space and time in the northeastern Pacific gastropod, Littorina keenae

The present population structure of a species reflects the influence of population history as well as contemporary processes. To examine the relative importance of these factors in shaping the current population structure of Littorina keenae, we sequenced 762 base pairs of the mitochondrial ND6 and cytochrome b genes in 584 snails from 13 sites along the northeastern Pacific coast. Haplotype network analysis revealed a 'star-like' genealogy indicative of a recent population expansion. Nested clade and mismatch analyses also supported the hypothesis of sudden population expansion following a population bottleneck during the Last Glacial Maximum. Analysis of molecular variance and pairwise Phi(ST) showed no significant spatial population differentiation from Mexico to Oregon - not even across the recognized biogeographic boundary at Point Conception. This is probably due to high contemporary gene flow during the free-swimming larval stage of this snail. Surprisingly, we found a highly significant temporal population differentiation between a San Pedro sample from 1996 and one from 2005, which gave an estimate of effective population size (N(e)) of only 135. Nearly statistically significant changes in the frequency of a particular haplotype in three other populations over 2-3 years further support Hedgecock's 'sweepstakes' hypothesis. When by chance nearly all of the progeny from an aggregation of highly fecund sisters that possess a rare haplotype successfully recruit to become the next generation, the rare haplotype can become temporarily common across the entire species' range. This modification of the sweepstakes hypothesis can explain why the temporal variation that we observed was much greater than the spatial variation.