Ecological diversity and speciation in land snails of the genus Mandarina from the Bonin Islands

Endemic land snails of the genus Mandarina of the oceanic Bonin Islands offer an example of habitat and character divergence among closely related species. The molecular phylogenies of Mandarina show that a divergence of arboreal, semiarboreal, and ground-dwelling species has occurred repeatedly in different times, areas, and lineages. Ecological diversification is suggested to be important for the coexistence of Mandarina species based on the facts that sympatric species are typically highly differentiated ecologically and morphologically, and that species occupying similar habitats do not coexist. The ecological diversification of Mandarina has occurred without much genetic divergence compared with that of its mainland relatives. This suggests that morphological and ecological diversifications are accelerated in depauperate environments where there are fewer competitors and predators. Although the details of the reproductive isolation mechanisms are not understood and further examination is needed, the rapid evolution of prezygotic isolation is the main cause of speciation in Mandarina. In particular, ecological diversification may be an effective barrier to gene exchange between two species. Because of incomplete post-mating isolation and the lower genetic divergence among species of Mandarina, breakdowns of reproductive isolation have frequently occurred as a result of habitat change. It is important to estimate the effect of hybridization on species diversification in future studies.