Diverse, continuous, and plastic sexual systems in barnacles.

Barnacles (Crustacea: Thoracica) show diverse sexual systems, including simultaneous hermaphroditism, androdioecy (hermaphrodites + males), and dioecy (females + males). When males occur, they are always much smaller (called dwarf males) than conspecific hermaphrodites or females. Ever since Darwin made this discovery, many scientists have been fascinated by such diversity. In this study, we provide an overview of (1) the diversity of sexual systems in barnacles, (2) the continuity between different sexual systems in some genera or species, and (3) the plasticity in sexual expression in several species. First, although most barnacles are hermaphroditic, both theoretical and empirical studies suggest that females and dwarf males tend to occur in species with small mating groups. Low sperm competition among hermaphrodites and little chance to act as a male are both associated with small group sizes and identified as the forces promoting the evolution of dwarf males and pure females, respectively. Second, in some groups of barnacles, the distinction between hermaphrodites and dwarf males is unclear because of the potential of dwarf males to become hermaphrodites. As many barnacle species tend toward protandric simultaneous hermaphroditism (develop male function first and then add female function without discarding male function), the dwarf males in such cases are best described as potential hermaphrodites that arrest growth and emphasize male function much earlier because of attachment to conspecifics. This is presumably advantageous in fertilizing the eggs of the host individuals. The distinction between hermaphrodites and females may also be obscured in some species. Third, sex allocation and penial morphology are plastic in some species. We also report the results of a transplanting experiment on small individuals of the pedunculate barnacle Octolasmis angulata, which suggests that individuals transplanted onto conspecifics developed longer and broader penises than did control individuals. Overall, the diversity, continuity, and plasticity in the sexual systems of barnacles are sources of important insights into the evolution and maintenance of the diversity of sexual systems.