Advances in Spermatological Characters in the Digenea: Review and Proposal of Spermatozoa Models and Their Phylogenetic Importance.

The wide biodiversity and economic importance of digeneans have motivated a great deal of research in the last decade, focussing on their phylogenetic positions. Molecular research was instrumental for our understanding of phylogeny in the Digenea, but spermatological studies have also provided many results, which are potentially useful for phylogeny; however, the complete spermatological data set has never been reviewed in a whole phylogenetic perspective. Spermatological data are now available for more than 100 species, belonging to 15 superfamilies and 46 families. In this paper, we try to summarize the current knowledge about sperm structure in the digeneans and propose a classification of digenean spermatozoa into five basic models. The main ultrastructural characters used are (1) the type of axoneme, (2) the lateral expansion, (3) the association 'external ornamentation of the plasma membrane + cortical microtubules', (4) the field of cortical microtubules and its number, (5) the location of the external ornamentation, (6) the location of the maximum number of cortical microtubules and (7) the number of mitochondria. We also outline the most interesting features for phylogenetic inference and their possible value in the context of digenean systematics, phylogeny and evolution. Associations between sperm models and superfamilies were found as follows: Type 1 in the Schistosomatoidea; Type 2 in the Hemiuroidea; Type 3 in the Opecoeloidea, Lepocreadioidea, Haploporoidea and Opisthorchioidea; Type 4 in the Gorgoderoidea, Microphalloidea, Plagiorchioidea and Gymnophalloidea; Type 5 in the Echinostomatoidea, Microscaphidioidea, Paramphistomoidea, Pronocephaloidea and Brachylaimoidea.