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.
For 40 years, paleontological studies of marine gastropods have suggested that species selection favors lineages with short-lived (lecithotrophic) larvae, which are less dispersive than long-lived (planktotrophic) larvae. Although lecithotrophs appeared to speciate more often and accumulate over time in some groups, lecithotrophy also increased extinction rates, and tests for state-dependent diversification were never performed. Molecular phylogenies of diverse groups instead suggested lecithotrophs accumulate without diversifying due to frequent, unidirectional character change. Although lecithotrophy has repeatedly originated in most phyla, no adult trait has been correlated with shifts in larval type. Thus, both the evolutionary origins of lecithotrophy and its consequences for patterns of species richness remain poorly understood. Here, we test hypothesized links between development mode and evolutionary rates using likelihood-based methods and a phylogeny of 202 species of gastropod molluscs in Sacoglossa, a clade of herbivorous sea slugs. Evolutionary quantitative genetics modeling and stochastic character mapping supported 27 origins of lecithotrophy. Tests for correlated evolution revealed lecithotrophy evolved more often in lineages investing in extra-embryonic yolk, the first adult trait associated with shifts in development mode across a group. However, contrary to predictions from paleontological studies, species selection actually favored planktotrophy; most extant lecithotrophs originated through recent character change, and did not subsequently diversify. Increased offspring provisioning in planktotrophs thus favored shifts to short-lived larvae, which led to short-lived lineages over macroevolutionary time scales. These findings challenge long-standing assumptions about the effects of alternative life histories in the sea. Species selection can explain the long-term persistence of planktotrophy, the ancestral state in most clades, despite frequent transitions to lecithotrophy.
BackgroundThe impact of predator-prey interactions on the evolution of many marine invertebrates is poorly understood. Since barriers to genetic exchange are less obvious in the marine realm than in terrestrial or freshwater systems, non-allopatric divergence may play a fundamental role in the generation of biodiversity. In this context, shifts between major prey types could constitute important factors explaining the biodiversity of marine taxa, particularly in groups with highly specialized diets. However, the scarcity of marine specialized consumers for which reliable phylogenies exist hampers attempts to test the role of trophic specialization in evolution. In this study, RNA-Seq data is used to produce a phylogeny of Cladobranchia, a group of marine invertebrates that feed on a diverse array of prey taxa but mostly specialize on cnidarians. The broad range of prey type preferences allegedly present in two major groups within Cladobranchia suggest that prey type shifts are relatively common over evolutionary timescales.ResultsIn the present study, we generated a well-supported phylogeny of the major lineages within Cladobranchia using RNA-Seq data, and used ancestral state reconstruction analyses to better understand the evolution of prey preference. These analyses answered several fundamental questions regarding the evolutionary relationships within Cladobranchia, including support for a clade of species from Arminidae as sister to Tritoniidae (which both preferentially prey on Octocorallia). Ancestral state reconstruction analyses supported a cladobranchian ancestor with a preference for Hydrozoa and show that the few transitions identified only occur from lineages that prey on Hydrozoa to those that feed on other types of prey.ConclusionsThere is strong phylogenetic correlation with prey preference within Cladobranchia, suggesting that prey type specialization within this group has inertia. Shifts between different types of prey have occurred rarely throughout the evolution of Cladobranchia, indicating that this may not have been an important driver of the diversity within this group.Electronic supplementary materialThe online version of this article (10.1186/s12862-017-1066-0) contains supplementary material, which is available to authorized users.
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