Phylogeny and Systematics of Cyathocotylid Digeneans (Digenea: Diplostomoidea) Parasitizing Snakes with Description of Three New Species of Gogatea from Australia and Vietnam

The Cyathocotylidae Mühling, 1896 is a small but broadly distributed family of digeneans parasitic in a wide range of vertebrate definitive hosts, from fish to mammals. Despite existing taxonomic questions, only a few studies have generated DNA sequence data from cyathocotylids, and only a single publication included sequence data from a cyathocotylid parasitic in snakes. Four genera are known to infect snakes: GogateaLutz, 1935, Szidatia Dubois, 1938, MesostephanoidesDubois, 1951, and SerpentostephanusSudarikov, 1961. Members of these genera were known from only Asia and Africa. In the present study, we describe 2 new species of Gogatea from snakes in Australia and 1 from Vietnam. The new species from Vietnam described herein is the first member of the genus that lacks a ventral sucker. We used partial sequences of the nuclear large ribosomal subunit (28S) and mitochondrial cytochrome c oxidase subunit I (COI) genes to explore phylogenetic relationships among cyathocotylids and species differentiation. In addition, this is the first report of a cyathocotylid from snakes in Australia, thus posing interesting questions regarding the dispersal and historical biogeography of these parasites. Cyathocotylid genera from snakes have a long, convoluted taxonomic history. The genera Gogatea, Mesostephanoides, and Szidatia were distinguished from each other based on very small morphological differences. Therefore, the validity of Szidatia and Mesostephanoides was often questioned in the literature. Based on the detailed morphological analysis of our freshly collected high-quality specimens and comparison with published information, we synonymize Mesostephanoides and Szidatia with Gogatea.