Overview of renicolid digeneans (Digenea, Renicolidae) from marine gulls of northern Holarctic with remarks on their species statuses, phylogeny and phylogeography

Renicolid digeneans parasitize aquatic birds. Their intramolluscan stages develop in marine and brackish-water gastropods, while metacercariae develop in molluscs and fishes. The systematics of renicolids is poorly developed, their life cycles are mostly unknown, and the statuses of many species require revision. Here, we establish based on integrated morphological and molecular data that adult renicolids from gulls Larus argentatus and Larus schistisagus and sporocysts and cercariae of Cercaria parvicaudata from marine snails Littorina spp. are life-cycle stages of the same species. We name it Renicola parvicaudatus and synonymized with it Renicola roscovitus. An analysis of the cox1 gene of R. parvicaudatus from Europe, North America and North Asia demonstrates a low genetic divergence, suggesting that this species has formed quite recently (perhaps during last glacial maximum) and that interregional gene flow is high. In Littorina saxatilis and L. obtusata from the Barents Sea, molecular analysis has revealed intramolluscan stages of Cercaria littorinae saxatilis VIII, a cryptic species relative to R. parvicaudatus. In the molecular trees, Renicola keimahuri from L. schistisagus belongs to another clade than R. parvicaudatus. We show that the species of this clade have cercariae of Rhodometopa group and outline morphological and behavioural transformations leading from xiphidiocercariae to these larvae. Molecular analysis has revealed 3 main phylogenetic branches of renicolids, differing in structure of adults, type of cercariae and host range. Our results elucidate the patterns of host colonization and geographical expansion of renicolids and pave the way to the solution of some long-standing problems of their classification.