Many marine species, including mussels in the Mytilus edulis species group (i.e. M. edulis L., M. galloprovincialis Lamarck, and M. trossulus Gould), have an antitropical distribution pattern, with closely related taxa occurring in high latitudes of the northern and southern hemispheres but being absent from the tropics. We tested four hypotheses to explain the timing and route of transequatorial migration by species with antitropical distributions. These hypotheses yield dierent predictions for the phylogenetic relationship of southern hemisphere taxa relative to their northern counter-parts. The three Mytilus species were used to test these hypotheses since they exhibit a typical antitropical distribution and representative taxa occur in both the Paci®c and Atlantic. Two types of mtDNA lineages were found among populations of mussels collected from the southern hemisphere between 1988 and 1996; over 90% of the mtDNA lineages formed a distinct subclade which, on average, had 1.4% divergence from haplotypes found exclusively in northern Atlantic populations of M. galloprovincialis. These data indicate that southern hemisphere mussels arose from a migration event from the northern hemisphere during the Pleistocene via an Atlantic route. The remainder of the southern hemisphere lineages (<10%) were very closely related to mtDNA haplotypes found in both M. edulis and M. galloprovincialis in the northern hemisphere, suggesting a second, more recent migration to the southern hemisphere. There was no evidence that southern hemisphere mussels arose from Paci®c populations of mussels.

Geographically isolated populations may accumulate alleles that function well on their own genetic backgrounds but poorly on the genetic backgrounds of other populations. Consequently, interpopulation hybridization may produce offspring of low fitness as a result of incompatibilities arising in allopatry. Genes participating in these epistatic incompatibility systems remain largely unknown. In fact, despite the widely recognized importance of epistatic interactions among gene products, few data directly address the functional consequences of such interactions among natural genetic variants. In the marine copepod, Tigriopus californicus, we found that the cytochrome c variants isolated from two different populations each had significantly higher activity with the cytochrome c oxidase derived from their respective source population. Three amino acid substitutions in the cytochrome c protein appear to be sufficient to confer population specificity. These results suggest that electron transport system (ETS) proteins form coadapted sets of alleles within populations and that disruption of the coadapted ETS gene complex leads to functional incompatibilities that may lower hybrid fitness.

Abstract. The North Atlantic intertidal community provides a rich set of organismal and environmental material for the study of ecological genetics. Clearly defined environmental gradients exist at multiple spatial scales: there are broad latitudinal trends in temperature, meso-scale changes in salinity along estuaries, and smaller scale gradients in desiccation and temperature spanning the intertidal range. The geology and geography of the American and European coasts provide natural replication of these gradients, allowing for population genetic analyses of parallel adaptation to environmental stress and heterogeneity. Statistical methods have been developed that provide genomic neutrality tests of population differentiation and aid in the process of candidate gene identification. In this paper, we review studies of marine organisms that illustrate associations between an environmental gradient and specific genetic markers. Such highly differentiated markers become candidate genes for adaptation to the environmental factors in question, but the functional significance of genetic variants must be comprehensively evaluated. We present a set of predictions about locus-specific selection across latitudinal, estuarine, and intertidal gradients that are likely to exist in the North Atlantic. We further present new data and analyses that support and contradict these simple selection models. Some taxa show pronounced clinal variation at certain loci against a background of mild clinal variation at many loci. These cases illustrate the procedures necessary for distinguishing selection driven by internal genomic vs. external environmental factors. We suggest that the North Atlantic intertidal community provides a model system for identifying genes that matter in ecology due to the clarity of the environmental stresses and an extensive experimental literature on ecological function. While these organisms are typically poor genetic and genomic models, advances in comparative genomics have provided access to molecular tools that can now be applied to taxa with well-defined ecologies. As many of the organisms we discuss have tight physiological limits driven by climatic factors, this synthesis of molecular population genetics with marine ecology could provide a sensitive means of assessing evolutionary responses to climate change.

Two blue mussel sibling species, Mytilus edulis and Mytilus galloprovincialis, have been imported to Pacific northwest waters in separate attempts to circumvent early mortality experienced by the native species (Mytilus trossulus) when cultured. Here we describe two novel markers based on the polymerase chain reaction that discriminate the two non-native, or alien, mussel species from the native species. The two markers are inherited in a Mendelian fashion, show no evidence of close linkage, and are highly diagnostic. To screen British Columbian mussel populations for the presence of the alien species, we assayed genotypes at the two marker loci in 482 subtidal mussels collected at 12 sites. Animals with at least one alien allele were present at 9 of 12 sampling sites with an overall frequency of 6.2%. The distribution of alien alleles at the two marker loci differed among sampling sites, suggesting differential introgression. The widespread incidence of alien alleles, combined with evidence of extensive hybridization between the alien and native species, indicates that the introduced alleles will probably persist in British Columbia mussels for some time.

We have used the polymerase chain reaction (PCR) and restriction fragment length polymorphism (RFLP) techniques to design two genetic markers for blue mussels in the Mytilus edulis species complex. Both of these markers target the gene encoding the mussel polyphenolic adhesive protein. The first marker, Glu-5', is highly differentiated among and can be used to identify the three blue mussel species, M edulis, M galloprovincialis and M trossulus. The second marker, Glu-3', can identify M edulis and M galloprovincialis. Using these markers we have demonstrated that hybrid mussels from Whitsand Bay, UK carry alleles for this gene that are the products of intragenic recombination. The high frequency (10 per cent) of these recombinant alleles within the hybrid population suggests that recombination is fairly frequent within this gene or that hybridization between M. edulis and M. galloprovincialis is substantial and has been occurring over considerable evolutionary time. The two novel genetic markers, Glu-5' and Glu-3' will be invaluable in additional studies regarding the importance of hybrization among blue mussels.