Diversity and ancestry of flatworms infecting blood of nontetrapod craniates "fishes".

We herein review all published molecular studies (life history, taxonomy, and phylogeny) and summarize all GenBank sequences and primer sets for the "fish blood flukes". Further, by analysing new and all available sequence data for the partial D1-D2 domains of 28S from 83 blood fluke taxa, we explore the evolutionary expansion of flatworm parasitism in the blood of craniates. Based on this analysis, the blood flukes infecting marine bony fishes (Euteleostei) are monophyletic. The clade comprising the chondrichthyan blood fluke plus the marine euteleost blood flukes is the sister group to tetrapod blood flukes (spirorchiids and schistosomes). The innominate blood fluke cercariae from freshwater gastropods were monophyletic and sister to the clade comprising spirorchiids and schistosomes, but low nodal support indicated that they may represent a distinct blood fluke lineage with phylogenetic affinities also to fish blood flukes. Blood flukes that utilize gastropod intermediate hosts were monophyletic (unidentified gastropod cercariae+tetrapod blood flukes) and those utilizing bivalves and polychaetes were monophyletic (marine fish blood flukes). Low or no taxon sampling among blood flukes of basal fish lineages and primary division freshwater fish lineages are significant data gaps needing closure. We also note that no record of an infection exists in a hagfish (Myxiniformes), lamprey (Petromyzontiformes), or nontetrapod sarcopterygiian, i.e., coelacanth (Coelacanthimorpha) or lungfish (Dipnoi). The present phylogenetic analysis reiterated support for monophyly of Schistosomatidae and paraphyly of spirorchiids, with the blood flukes of freshwater turtles basal to those of marine turtles and schistosomes.