BackgroundReconstructing the higher relationships of pulmonate gastropods has been difficult. The use of morphology is problematic due to high homoplasy. Molecular studies have suffered from low taxon sampling. Forty-eight complete mitochondrial genomes are available for gastropods, ten of which are pulmonates. Here are presented the new complete mitochondrial genomes of the ten following species of pulmonates: Salinator rhamphidia (Amphiboloidea); Auriculinella bidentata, Myosotella myosotis, Ovatella vulcani, and Pedipes pedipes (Ellobiidae); Peronia peronii (Onchidiidae); Siphonaria gigas (Siphonariidae); Succinea putris (Stylommatophora); Trimusculus reticulatus (Trimusculidae); and Rhopalocaulis grandidieri (Veronicellidae). Also, 94 new pulmonate-specific primers across the entire mitochondrial genome are provided, which were designed for amplifying entire mitochondrial genomes through short reactions and closing gaps after shotgun sequencing.ResultsThe structural features of the 10 new mitochondrial genomes are provided. All genomes share similar gene orders. Phylogenetic analyses were performed including the 10 new genomes and 17 genomes from Genbank (outgroups, opisthobranchs, and other pulmonates). Bayesian Inference and Maximum Likelihood analyses, based on the concatenated amino-acid sequences of the 13 protein-coding genes, produced the same topology. The pulmonates are paraphyletic and basal to the opisthobranchs that are monophyletic at the tip of the tree. Siphonaria, traditionally regarded as a basal pulmonate, is nested within opisthobranchs. Pyramidella, traditionally regarded as a basal (non-euthyneuran) heterobranch, is nested within pulmonates. Several hypotheses are rejected, such as the Systellommatophora, Geophila, and Eupulmonata. The Ellobiidae is polyphyletic, but the false limpet Trimusculus reticulatus is closely related to some ellobiids.ConclusionsDespite recent efforts for increasing the taxon sampling in euthyneuran (opisthobranchs and pulmonates) molecular phylogenies, several of the deeper nodes are still uncertain, because of low support values as well as some incongruence between analyses based on complete mitochondrial genomes and those based on individual genes (18S, 28S, 16S, CO1). Additional complete genomes are needed for pulmonates (especially for Williamia, Otina, and Smeagol), as well as basal heterobranchs closely related to euthyneurans. Increasing the number of markers for gastropod (and more broadly mollusk) phylogenetics also is necessary in order to resolve some of the deeper nodes -although clearly not an easy task. Step by step, however, new relationships are being unveiled, such as the close relationships between the false limpet Trimusculus and ellobiids, the nesting of pyramidelloids within pulmonates, and the close relationships of Siphonaria to sacoglossan opisthobranchs. The additional genomes presented here show that some species share an identical mitochondrial gene order due to convergence.
Species of the limpet genus Siphonaria (Gastropoda: Euthyneura) are commonly found in the rocky intertidal, worldwide, except in the Arctic. In total, 205 species-group names are available and not permanently invalid. However, estimating the actual species diversity of Siphonaria has remained challenging, mainly because past authors have interpreted differently the variation of shell characters, resulting in different taxonomic accounts. Species diversity of Siphonaria is evaluated for the first time here based on DNA sequence data (three mitochondrial gene fragments: COI, 12S, and 16S) and a large sampling focusing on the tropical and subtropical Indo-West Pacific (from eastern Africa to Hawaii): new sequences are provided for 153 individuals, 123 of which were collected from 93 locations throughout the Indo-West Pacific. In total, 41 species (molecular units) are recognized worldwide (31 from the Indo-West Pacific), all of which are strongly supported. Potential names are discussed for those 41 species, based on traditional taxonomy. The shells of 66 of the individuals from which DNA was extracted are illustrated: intra- and inter-specific variation is documented in detail and discussed in the light of new molecular results. It is shown that many species could hardly be identified based on the shell only, because the variation of shell characters is too high and overlaps between species. Geographically, no species is found across the entire Indo-West Pacific, where quite a few species seem to be endemic to restricted areas. The biogeography of Siphonaria in the Indo-West Pacific is compared to other groups.
Corallimorpharia is currently considered an order of hexacorallian anthozoans. Being skeletonless, its members are sometimes referred to as sea anemones, but they are morphologically more similar to members of Scleractinia than to members of Actiniaria. We describe two new species of corallimorpharians from deep water off the west coast of North America as Corallimorphus denhartogi, n. sp. and Corallimorphus pilatus, n. sp. The former occurs at depths of 2550-4300 m from Oregon to Baja California, and the latter at depths of 198-900 m from British Columbia to southernmost California. The average size of individuals of C. denhartogi is greater than that of C. pilatus, and tentacles of the latter are more densely arrayed and relatively longer than those of the former. The distribution and sizes of their cnidae distinguish them from one another as well as from their four congeners, which are widely distributed in the world's oceans. In the collections we examined, specimens of C. denhartogi are more common than those of C. pilatus.CORALLIMORPHARIA IS CURRENTLY considered an order of hexacorallian anthozoans. Its members are skeletonless, and so are sometimes referred to as sea anemones, but are characterized by anatomy and nematoeysts that more closely ally them with "true" or I
Not since Hubendick’s 1946 Systematic Monograph of the Patelliformia has a comprehensive taxonomic work been com-pleted on the genus of false-limpets Siphonaria, distributed worldwide, except in the northern Atlantic. A checklist of allsupra-specific names associated with Siphonaria is established. A checklist of all species-group names is provided as welland their nomenclatural status is discussed (available, not available, permanently invalid). Of the 270 species-group namesincluded in the checklist, 205 are available and not permanently invalid. All other names are either not available or avail-able but permanently invalid. The geographic distribution of the type localities is also presented; of the 205 species-group names that are available and not permanently invalid, 62 are from the tropical and subtropical Indo-West Pacific.
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