Cotylea (Polycladida): a cladistic analysis of morphology

Polyclad flatworms are acoelomate bilaterians found in benthic communities worldwide, predominantly in marine environments. Current polyclad systematics is unstable, with two non-concordant classification schemes resulting in a poor understanding of within-group relationships. Here we present the first phylogenetic framework for the suborder Cotylea using a morphological matrix. Representatives of 34 genera distributed among all cotylean families (except four, excluded due to their dubious taxonomic status) were investigated. The number of families included ranges from a conservative eight to a revisionary 11. Outgroup analysis indicated that the suborder is monophyletic and defined by the presence of a ventral adhesive structure, a short posteriorly positioned vagina, and cement glands. Of the eight to 11 families included, we confirmed that three were monophyletic: Boniniidae, Prosthiostomidae, and Pseudocerotidae. Boniniidae was consistently recovered as the sister group to other Cotylea, based on the retention of the plesiomorphic Lang\u27s vesicle. The clade consisting of Anonymus, Marcusia, and Pericelis is sister to the Boniniidae and the rest of the Cotylea. Above this clade there is little resolution at the base of the sister group. The Euryleptidae are found to be paraphyletic and give rise to the Pseudocerotidae. Neither classification scheme received unequivocal support. The intrafamilial relationships of the diverse Pseudocerotidae and Euryleptidae were examined. Color pattern characters (used for species identification) were highly homoplasious but increased cladogram resolution within genera. The monophyly of seven genera within the Pseudocerotidae and Euryleptidae was not supported and many genera showed no autapomorphies, highlighting the need for taxonomic revision of these families