Molecular genetics of peripheral populations of Nova Scotian Unionidae (Mollusca: Bivalvia)

Peripheral populations of eight species of freshwater bivalves (Unionidae.) extending their geographic ranges into Nova Scotia, Canada, were examined electrophoretically to determine both the extent of genetic variability within such populations, and whether the hypothesized pathway of colonization across the Isthmus of Chignecto is reflected in patterns of genetic resemblance among these populations. The Nova Scotian species examined could be separated into two groups based on levels of observed heterozygosity and levels of variability in allele frequencies. The first group is characterized by low levels of heterozygosity and polymorphism compared with north-eastern American populations, and in the case of one species, Elliptio complanala, considerable variability in allele frequencies among populations occurring in similar habitats in different drainages. Populations of E. complanata from Nova Scotia can be differentiated from conspecific populations on the southern Atlantic Slope by possession of fast alleles at two loci. Multivariate analyses define subgroups within populations of E. complanata consistent with hypothesis that the species invaded Nova Scotia by way of the Isthmus of Chignecto, and then split into two groups, one of which colonized Cape Breton to the north and the other of which colonized southern areas of the Province. The second group of Nova Scotian species is characterized by little reduction in heterozygosity and polymorphism compared with values observed among north-eastern American conspecifics or congeners, little variability in allele frequencies from population to population, and little evidence to suggest that these species were dependent on the land bridge to invade the Province. The type of dispersal is hypothesized to be responsible, in part, for these differences: larvae of species in the first group rely on a parasitic attachment to fish with territorial habits limited to fresh water, and are thus likely to invade new drainages separated by salt water by chance, in small numbers, and in stepping-stone fashion. Species in the second group parasitize anadromous or saltwater tolerant hosts, are likely to be introduced into new habitats in greater numbers and/or receive greater amounts of gene flow subsequent to colonization, and seem less dependent on land-bridges to colonize new habitats.     

PARASITISM AND THE UNIONACEA (BIVALVIA)

It is proposed that the incorporation of a unique parasitic stage in the life-cycle of unionaceans which involves an obligate relationship between a vertebrate host, usually a fish, and a highly modified larval stage, the glochidium, has had far-reaching consequences with respect to overall morphology, extent of species' geographic ranges, and rate of speciation in the group. Glochidia are separable into three main types with respect to overall shape and attachment features, and are retained in variously modified brood pouches. When mature, glochidia are released in several different ways which reflect various adaptations involved in either attracting the fish host and/or increasing the probability of attachment. Glochidia do not seem capable of host selection, and the reaction of the host to the parasite seems to be the main factor in determining specificity. Release of glochidia is synchronized to correspond to periods of predictable host availability, such as during spawing migrations and nesting behaviour. Other adaptations include modifications of glochidial conglutinates to mimic host food items, and modifications of the unionacean mantle edges to attract hosts. In all cases, a good correlation exists between the type of lure used and host food preferences, but, despite these adaptations, host specificity among unionaceans seems low. Parasitism among unionaceans is postulated to be mainly advantageous in terms of predictability of dispersal by habitat-specific hosts, but parasitism is hypothesized to entail constraints in terms of the degree to which shell shape and life-habit can be diversified among unionaceans. The type of host parasitized is considered to affect the rate of diversification among populations and speciation among unionaceans: those that parasitize strictly freshwater hosts are more likely to exhibit highly individualistic populations in different drainages with respect to molecular genetic and soft-part characters, while those that parasitize anadromous or saltwater-tolerant hosts show little differentiation among widely distributed populations.     

Paul David Pavy II: A Correction

Careful analysis of Nacireman academies does not bear out the conclusion of Walker (1970) that these are dysfunctional. Indeed, they are an integral part of Nacireman age-grade structure and play an important role in initiation into a Nacireman adult age-grade.