Phylogenetic patterns and phenotypic plasticity of molluscan sexual systems.

Molluscs show a wide diversity of sexual systems and strategies. There are both gastropod and bivalve families that are each primarily dioecious, simultaneous hermaphrodites, or sequential hermaphrodites, and other families in which almost every sexual strategy occurs. The multiple evolutionary transitions of sexual systems within molluscs would allow comparative analyses of the associated ecological factors, but data on all but a few groups are too sparse to draw many solid conclusions. The phylogenetic distribution of sexual systems in the Mollusca shows that gastropods and bivalves demonstrate different patterns, possibly associated with the presence/absence of copulation. The distribution of change of sex suggests that, in gastropods, sequential hermaphrodites do not evolve from simultaneous hermaphrodites, and that sex reversal (flip-flopping) occurs in free-spawners but not in copulators. Three well-studied protandrous gastropod groups (calyptraeids, coralliophilids, and patellogastropods) show similar responses to environmental conditions and associations with conspecifics. They all have the following attributes: (1) they are sedentary, (2) they live in groups, patches, or aggregates, and (3) size at sex change varies among sites and among aggregates. In addition the available experimental evidence suggests that (4) the presence of females or large individuals represses growth and sex change of males, and (5) behavior seems to mediate the repressive influence of large females. Available data from other species tend to support these patterns. Finally, the repression of growth of males by females in protandry likely facilitates the evolution of dwarf males.