Phylogeny of the radula-less dorids (Mollusca, Nudibranchia), with the description of a new genus and a new family

This paper studies the phylogenetic relationships of the nudibranch dorids that lack a radula. Numerous specimens belonging to 29 different species, representing all genera of this group and other related taxa, were examined anatomically. Details of the foregut were examined with SEM. From this anatomical study a total of 62 characters were considered. These characters have been polarized using the genus Berthella as the outgroup. Eight species of dorids with a radula and the genus Armina have been also included in the analysis for comparative purposes. The phylogeny obtained supports the hypothesis that the radula has been lost once in the Doridina and that the radula-less dorids are a monophyletic group. Cryptobranchia is monophyletic when the radula-less dorids are included. The new genus Mandelia, introduced on the basis of a new species from South Africa, is the sister group of the rest of the radula-less dorids, and is described as a new family, Mandeliidae. The radula-less dorids (currently placed in a separate superfamily) actually constitute an internal branch of other cryptobranch dorids. Phyllidia has no synapomorphies that distinguish if from Fryeria, which has one autoapomorphy, the ventral position of the anus. Therefore, both genera are regarded as synonyms. The evolution of several characters among the radula-less dorids is discussed, and the present phylogeny is compared with prior studies.