A phylogeny of planorbid snails, with implications for the evolution of Schistosoma parasites

The Planorbidae represent one of the most important families of freshwater snails. They have a wide distribution and are significant both medically and economically as intermediate hosts for trematode worms. Digenetic trematodes of the genus Schistosoma cause schistosomiasis, a disease that infects 200 million people, and domestic animals throughout the tropics. Three of the four recognized species groups of Schistosoma rely on snails of the family Planorbidae to complete their life cycles. Each species group requires a specific planorbid genus-Bulinus, Biomphalaria, or Indoplanorbis. Our understanding of the relationships among the genera within the Planorbidae is rudimentary and based solely on internal anatomy and shell morphology. Two molecular markers, ribosomal 28S and actin exon 2, were sequenced and a phylogeny constructed for 38 taxa representing 16 planorbid genera. The phylogeny supports the division of the Planorbidae into two subfamilies, the Bulininae and Planorbinae. Interestingly, two representatives of the family Ancylidae fall within the Planorbidae highlighting the need for further analysis and possible reclassification of this group. A molecular based phylogeny of the genus Schistosoma was then mapped against the snail tree. The trees indicate that planorbid-transmitted Schistosoma appear not to be co-speciating with their current snail host lineages. Rather, host switching was prominent, including a switch involving two distantly related planorbid genera, Biomphalaria and Bulinus. Our study of the Planorbidae poses fundamental questions regarding how and when Schistosoma acquired new snail hosts, including how switches to relatively distant hosts are accomplished and why some available planorbids were not colonized.