Traditional Taxonomic Groupings Mask Evolutionary History: A Molecular Phylogeny and New Classification of the Chromodorid Nudibranchs

Chromodorid nudibranchs (16 genera, 300+ species) are beautiful, brightly colored sea slugs found primarily in tropical coral reef habitats and subtropical coastal waters. The chromodorids are the most speciose family of opisthobranchs and one of the most diverse heterobranch clades. Chromodorids have the potential to be a model group with which to study diversification, color pattern evolution, are important source organisms in natural products chemistry and represent a stunning and widely compelling example of marine biodiversity. Here, we present the most complete molecular phylogeny of the chromodorid nudibranchs to date, with a broad sample of 244 specimens (142 new), representing 157 (106 new) chromodorid species, four actinocylcid species and four additional dorid species utilizing two mitochondrial markers (16s and COI). We confirmed the monophyly of the Chromodorididae and its sister group relationship with the Actinocyclidae. We were also able to, for the first time, test generic monophyly by including more than one member of all 14 of the non-monotypic chromodorid genera. Every one of these 14 traditional chromodorid genera are either non-monophyletic, or render another genus paraphyletic. Additionally, both the monotypic genera Verconia and Diversidoris are nested within clades. Based on data shown here, there are three individual species and five clades limited to the eastern Pacific and Atlantic Oceans (or just one of these ocean regions), while the majority of chromodorid clades and species are strictly Indo-Pacific in distribution. We present a new classification of the chromodorid nudibranchs. We use molecular data to untangle evolutionary relationships and retain a historical connection to traditional systematics by using generic names attached to type species as clade names

The Genera of Pleurobemini (Bivalvia: Unionidae: Ambleminae)

The unionid tribe Pleurobemini is diverse but poorly-understood phylogenetically. Current classification recognizes two highly diverse genera, Elliptio Rafinesque, 1819 and Pleurobema Rafinesque, 1820, besides the moderately diverse Fusconaia Simpson, 1900, and several genera with one to three species. However, classification at the species and genus level has been problematic. Molecular data and re-examination of shell morphology and anatomy indicate new groupings of these taxa. Several genera proposed by early workers such as Rafinesque and Swainson are available, but are poorly-characterized and are often overlooked. We analyzed two mitochondrial genes, cox1 and nad1, for 50 species assigned to Pleurobemini, including the type species of each genus and as many other species as possible. Although the majority of studied species in Elliptio, Pleurobema, and Fusconaia show close affinities to the respective type species, the affinities of others are problematic. Genera or subgenera such as Eurynia Rafinesque, 1819, SintoxiaRafinesque, 1820, and Pleuronaia Frierson, 1927, generally regarded as subjective synonyms, apply to some clades. Other clades or unaffiliated species have no available name. Quincuncina burkei (Walker, 1922), the type of the genus Quincuncina Ortmann, 1922, is assigned to Fusconaia Simpson, 1900. Fusconaia apalachicola (Williams and Fradkin, 1999), F. ebenus (Lea, 1831), F. rotulata (Wright, 1899) (also listed as Obovaria rotulata), F. succissa (Lea, 1852), Cyclonaias tuberculatus (Rafinesque, 1820), Uniomerus Conrad, 1853, and the Quincuncina infucata (Conrad, 1834) complex are all excluded from Pleurobemini. The first three are placed in the new genus Reginaia Campbell and Lydeard; Uniomerus Conrad, 1853 is assigned to Quadrulini; and the remainder belong in the pustulosa Lea, 1831 group of Quadrulini (genus RotundariaRafinesque, 1820)