Pattern and process of diversification in an ecologically diverse epifaunal bivalve group Pterioidea (Pteriomorphia, Bivalvia).

The pterioid bivalves (superfamily Pterioidea) are epifaunal filter feeders that attach to various substrata, including rocks, corals, gorgonians, and sponges. An intriguing question is how different substratum types have affected the diversification of pterioid bivalves. To elucidate the evolutionary pathway of Pterioidea, we conducted a phylogenetic analysis of 49 individuals belonging to 18 pterioid and 5 outgroup species using 18S and 28S ribosomal RNA genes. The results supported the monophyly of superfamily Pterioidea and recovered three major clades within Pterioidea: Malleus, Pteria and Electroma (Pterelectroma) zebra and the rest. This result contradicts the current circumscription of the pterioid families, Pteriidae, Malleidae, and Isognomonidae, and suggests that hinge morphology, which has long been used as a diagnostic character, does not reflect phylogenetic relationships. The monophyly of most genera, however, was confirmed. Mapping substrata types on the phylogenetic tree indicated that the ancestors of pterioid bivalves were epifaunal on rocks, and that epizoic life on sessile cnidarians had a single origin. Although we could not ascertain whether endozoic life in sponges evolved once or twice, our results suggest that colonization of biotic substrata resulted in diversification and morphological and ecological adaptation to epi-/endozoic life. We estimated that the genus Pteria has diversified since the late Cretaceous in response to gorgonian diversification. These results emphasize the importance of substratum shifts in speciation and diversification of pterioid bivalves.