The paper describes morphometric and allozymic differences among four European species of the family Viviparidae: Viviparus contectus (Millet, 1813), V. viviparus (Linnaeus, 1758), V. acerosus Bourguignat, 1862, and V. ater (Cristofori et Jan, 1832). Fourteen continuous biometrical characters were measured. Incremental discriminant-function analysis, principal-component analysis, and non-metric multidimensional scaling were applied to analysis of the morphometric differences. All the techniques confirmed a similar picture: a slight morphometric differentiation, with the variability ranges of the species overlapping. On the other hand, the allozymic differentiation, studied at 12 loci, eight of them intra and/or interspecific polymorphic, is much better marked, the intraspecific Nei's distances among the four V. contectus populations ranging from 0.0014 to 0.0397, mean 0.0166, and interspecific distances ranging from 0.2306 (V. ater–V. acerosus) to 0.9888 (V. contectus 2 and V. viviparus), mean 0.6871. The allele frequencies and genetic distances (Nei's distance and Cavalli-Sforza and Edwards’chord distance) were used to compute maximum likelihood, additive Fitch–Margoliash and ultrametric Fitch–Margoliash trees. All the trees presented a similar pattern, but the maximum-likelihood and additive trees, based on Cavalli-Sforza and Edwards’distance, seem to reflect the phylogeny best. The results are compared with the most parsimonious phylogenies inferred for radular, soft-part morphology and anatomy, and opercular data from other papers by us, and the inferred phylogenies are also compared. Although the inferred molecular and morphological phylogenies are little different in topology, the amount of evolution along the corresponding branches (measured as the number of changes averaged over all reconstructions) is very different, the value of correlation coefficient between the two phylogenies being statistically insignificant. The occurrence of interspecific hybrids is discussed, and the isolation-by-competition mechanism is postulated. The probable origin of V. viviparus from a founder population extremely restricted in number is stressed. The possible history of the group is briefly discussed. The species is suggested to have originated in an unusual habitat of melt water at a glacier foreland that could have promoted genotypic differentiation and sympatric speciation.
Read full abstract