Anatomy, functional morphology, evolutionary ecology and systematics of the invasive gastropod Cipangopaludina japonica (Viviparidae: Bellamyinae)

Abstract

The anatomy, functional morphology and evolutionary ecology of the Viviparidae, and the subfamily Bellamyinae in particular, are incompletely known. Partly as a result, genealogical relationships within the family remain poorly understood. Because of this lack in knowledge, few informed hypotheses exist on ancestral states, how differences in body plans between the subfamilies evolved, and how the peculiar biogeographic distribution patterns of viviparids have arisen. Here we document the anatomy, morphology, life history and systematics of Cipangopaludina japonica, a Japanese species that has been introduced into North America, to resolve taxonomic confusion and to improve our understanding of how form and function are related in bellamyines. Anatomical and histological examinations demonstrate marked differences between C. japonicaand other bellamyines in the radula, salivary gland, kidney, nerve ring and reproductive organs. Substantial differences also exist between male and female body organization, but conchological differences between sexes in semi-landmark morphometric analyses are limited. The volume of the brood pouch of females, and hence body and shell size, appear to be good predictors of reproductive success, and the species’ ecological versatility may relate to high fecundity and the ability to alternate between feeding modes. Comparing our observations on C. japonicawith other viviparids and basal Architaenioglossa, we identify several persistent misinterpretations in the literature on how form and function are related in viviparids, not in the least as to female reproductive anatomy. Our reinterpretations improve understanding of the evolution of Viviparidae and its subfamilies, and hopefully will allow future workers to isolate key traits that shaped the evolution of viviparids at the taxonomic levels of their interest for more detailed studies.