A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia)

Abstract

The phylogenetic relationships of the cryptobranch dorids are studied based on morphological characters of species belonging to all previously described genera. The phylogenetic hypothesis supports the cryptobranch dorids as a monophyletic group. There are two major clades within the Cryptobranchia: the radula-less dorids (Porostomata), and the radula-bearing dorids (Labiostomata new taxon). Labiostomata consists of those taxa sharing a more recent common ancestor with Actinocyclus than with Mandelia, and includes several monophyletic groups: Actinocyclidae, Chromodorididae, Dorididae and Discodorididae. The traditional group Phanerobranchia is probably paraphyletic. The new classification proposed for the Cryptobranchia addresses concepts of phylogenetic nomenclature, but is in accordance with the rules of the International Code of Zoological Nomenclature. The following genera of cryptobranch dorids are regarded as valid: Doris Linnaeus, 1758, Asteronotus Ehrenberg, 1831, Atagema J. E. Gray, 1850, Jorunna Bergh, 1876, Discodoris Bergh, 1877, Platydoris Bergh, 1877, Thordisa Bergh, 1877, Diaulula Bergh, 1878, Aldisa Bergh, 1878, Rostanga Bergh, 1879, Aphelodoris Bergh, 1879, Halgerda Bergh, 1880, Peltodoris Bergh, 1880, Hoplodoris Bergh, 1880, Paradoris Bergh, 1884, Baptodoris Bergh, 1884, Geitodoris Bergh, 1891, Gargamella Bergh, 1894, Alloiodoris Bergh, 1904, Sclerodoris Eliot, 1904, Otinodoris White, 1948, Taringa Er. Marcus, 1955 , Sebadoris Er. Marcus & Ev. Marcus, 1960, Conualevia Collier & Farmer, 1964, Thorybopus Bouchet, 1977, Goslineria Valdés, 2001, Pharodoris Valdés, 2001, Nophodoris Valdés & Gosliner, 2001. Several genera previously considered as valid are here regarded as synonyms of other names: Doridigitata d’Orbigny, 1839, Doriopsis Pease, 1860, Staurodoris Bergh, 1878, Fracassa Bergh, 1878, Archidoris Bergh, 1878, Anoplodoris Fischer, 1883, Etidoris Ihering, 1886, Phialodoris Bergh, 1889, Montereina MacFarland, 1905, Ctenodoris Eliot, 1907, Carryodoris Vayssière, 1919, Austrodoris Odhner, 1926, Guyonia Risbec, 1928, Erythrodoris Pruvot-Fol, 1933, Neodoris Baba, 1938, Siraius Er. Marcus, 1955, Tayuva Ev. Marcus & Er. Marcus, 1967, Nuvuca Ev. Marcus & Er. Marcus, 1967, Doriorbis Kay & Young, 1969, Pupsikus Er. Marcus & Ev. Marcus, 1970, Percunas Ev. Marcus, 1970, Verrillia Ortea & Ballesteros, 1981 . The genera Artachaea Bergh, 1882, Carminodoris Bergh, 1889 and Homoiodoris Bergh, 1882 have been poorly described and no type material is known to exist. They are regarded as incertae sedis until more material becomes available. The genus names Xenodoris Odhner in Franc, 1968 and Cryptodoris Ostergaard, 1950 are unavailable within the meaning of the Code. Hexabranchus Ehrenberg, 1831 is not a cryptobranch dorid, as suggested by other authors, because of the lack of a retractile gill. Other nomenclatural and taxonomic problems are discussed, and several type species, neotypes and lectotypes are selected. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society, 2002, 136, 535−636.