Erhaia (Gastropoda: Rissooidea): phylogenetic relationships and the question of Paragonimus coevolution in Asia.

The human lung fluke Paragonimus is transmitted by gastropod taxa of two superfamilies: Ceritheoidea and Rissooidea. The question whether or not Paragonimus shows the same specificity of host-parasite coevolved relationship as the human blood fluke Schistosoma was inspired by the finding of two sympatric snail species as hosts for Paragonimus skrjabini in Fujian Province, China: Gammatricula and Erhaia. The former species can clearly be classified as Pomatiopsidae: Triculinae. The latter has previously been classified as Pomatiopsidae: Pomatiopsinae. However, this classification based on anatomical characteristics is uncertain. In order to obtain a robust phylogenetic hypothesis for Erhaia, we have studied three gene fragments from this taxon as well as from twelve related taxa. The data show that the species involved represent four families: Pomatiopsidae, Hydrobiidae, Cochliopidae (here raised to family status), and Amnicolidae. Erhaia fits securely into the Amnicolidae. This indicates that P. skrjabini has not coevolved with snail lineages. However, P. skrjabini has so far only been reported from rissooidean snails, whereas members of the Paragonimus westermani complex have only been found in ceritheoidean snails. The implication is that there is a host specificity on the superfamily level. However, Asian freshwater species of the Ceritheoidea and Rissooidea usually are not sympatric and often prefer different habitats. It is therefore possible that ecological niche partitioning plays the primary role for Paragonimus evolution.