The evolution of reproductive isolation in a simultaneous hermaphrodite, the freshwater snail Physa.

BACKGROUND: The cosmopolitan freshwater snail Physa acuta has recently found widespread use as a model organism for the study of mating systems and reproductive allocation. Mitochondrial DNA phylogenies suggest that Physa carolinae, recently described from the American southeast, is a sister species of P. acuta. The divergence of the acuta/carolinae ancestor from the more widespread P. pomilia appears to be somewhat older, and the split between a hypothetical acuta/carolinae/pomilia ancestor and P. gyrina appears older still. RESULTS: Here we report the results of no-choice mating experiments yielding no evidence of hybridization between gyrina and any of four other populations (pomilia, carolinae, Philadelphia acuta, or Charleston acuta), nor between pomilia and carolinae. Crosses between pomilia and both acuta populations yielded sterile F1 progeny with reduced viability, while crosses between carolinae and both acuta populations yielded sterile F1 hybrids of normal viability. A set of mate-choice tests also revealed significant sexual isolation between gyrina and all four of our other Physa populations, between pomilia and carolinae, and between pomilia and Charleston acuta, but not between pomilia and the acuta population from Philadelphia, nor between carolinae and either acuta population. These observations are consistent with the origin of hybrid sterility prior to hybrid inviability, and a hypothesis that speciation between pomilia and acuta may have been reinforced by selection for prezygotic reproductive isolation in sympatry. CONCLUSIONS: We propose a two-factor model for the evolution of postzygotic reproductive incompatibility in this set of five Physa populations consistent with the Dobzhansky-Muller model of speciation, and a second two-factor model for the evolution of sexual incompatibility. Under these models, species trees may be said to correspond with gene trees in American populations of the freshwater snail, Physa.

Phylogenetic analysis of the Lancinae (Gastropoda, Lymnaeidae) with a description of the U.S. federally endangered Banbury Springs lanx.

We examined the patelliform snails of the subfamily Lancinae, endemic to northwestern North America, to test whether morphological variation correlated with genetic and anatomical differences. Molecular analyses using cox1, 16S, calmodulin intron, and 28S rDNA partial sequences and anatomical data supported recognition of four species in three genera. The relationships of lancines within Lymnaeidae are not yet well-resolved. The federally endangered Banbury Springs lanx is described as a new genus and species, Idaholanx fresti, confirming its distinctiveness and narrow endemicity.

Identification of 'extinct' freshwater mussel species using DNA barcoding.

Freshwater mollusks are highly imperiled, with 70% of the North American species extinct, endangered, or at risk of extinction. Impoundments and other human impacts on the Coosa River of Alabama, Georgia and Tennessee of the southeastern USA alone are believed to have caused 50 mollusk species extinctions, but uncertainty over boundaries among several putatively closely related species makes this number preliminary. Our examination of freshwater mussels collected during an extensive survey of the upper-drainage basin, DNA barcoding and molecular phylogenetic analyses confirm the rediscovery of four morphospecies in the genus Pleurobema (Unionidae) previously thought to be extinct from the upper Coosa basin. A fifth 'extinct' form was found in an adjoining basin. Molecular data show that the Coosa morphologies represent at least three species-level taxa: Pleurobema decisum, P. hanleyianum and P. stabile. Endemism is higher than currently recognized, both at the species level and for multispecies clades. Prompt conservation efforts may preserve some of these taxa and their ecosystem.

Molecular phylogeny and biogeography of spring-associated hydrobiid snails of the Great Artesian Basin, Australia.

The Great Artesian Basin (GAB) of Australia underlies some of the driest parts of South Australia and Queensland and feeds numerous freshwater springs. Prominent and endangered components of the GAB spring community are snails of the family Hydrobiidae. This paper examines the evolutionary relationships of the entire hydrobiid fauna associated with the GAB, and includes appropriate non-GAB species to place the GAB fauna in a broader phylogenetic context. The Queensland genus Jardinella is a focus of this paper, providing a fine scale examination of relationships between spring supergroups in the northeastern regions of the GAB. Maximum parsimony and Bayesian analyses performed on 16S, CO1, and combined sequence data from 40 hydrobiid taxa found four major clades of Australian taxa. The analysis revealed that at least three separate colonization events of the GAB spring fauna have occurred. Two of these are represented by considerable radiations, (1) Jardinella to the north and east and (2) Caldicochlea, Fonscochlea, and possibly Trochidrobia in South Australia. The phylogenetic position of the latter is uncertain so it may represent yet another invasion. The third definite invasion is represented by a single species of the speciose SE Australian genus Austropyrgus in the Dalhousie Springs in South Australia. Jardinella is found to be monophyletic, and with one exception, its members in each of the Queensland spring supergroups are found to be monophyletic.

Complete mtDNA sequence of the North American freshwater mussel, Lampsilis ornata (Unionidae): an examination of the evolution and phylogenetic utility of mitochondrial genome organization in Bivalvia (Mollusca).

Molluscs in general, and bivalves in particular, exhibit an extraordinary degree of mitochondrial gene order variation when compared with other metazoans. Two factors inhibiting our understanding the evolution of gene rearrangement in bivalves are inadequate taxonomic sampling and failure to examine gene order in a phylogenetic framework. Here, we report the first complete nucleotide sequence (16,060 bp) of the mitochondrial (mt) genome of a North American freshwater bivalve, Lampsilis ornata (Mollusca: Paleoheterodonta: Unionidae). Gene order and mt genome content is examined in a comparative phylogenetic framework for Lampsilis and five other bivalves, representing five families. Mitochondrial genome content is shown to vary by gene duplication and loss among taxa and between male and female mitotypes within a species. Although mt gene arrangement is highly variable among bivalves, when optimized on an independently derived phylogenetic hypothesis, it allows for the reconstruction of ancestral gene order states and indicates the potential phylogenetic utility of the data. However, the interpretation of reconstructed ancestral gene order states must take in to account both the accuracy of the phylogenetic estimation and the probability of character state change across the topology, such as the presence/absence of atp8 in bivalve lineages. We discuss what role, if any, doubly uniparental inheritance (DUI) and recombination between sexual mitotypes may play in influencing gene rearrangement of the mt genome in some bivalve lineages.

Phylogeny, taxonomy, genetics and global heritage ranks of an imperilled, freshwater snail genus Lithasia (Pleuroceridae).

Numerous aquatic species are threatened with extinction from habitat elimination or modification. One particularly imperilled group is the freshwater gastropod family Pleuroceridae. Pleurocerids reach their greatest diversity in the southeastern United States, and many species are currently considered extinct, endangered or threatened. One issue hindering efforts to implement conservation management plans for imperilled pleurocerid species is that the taxonomy is in an abysmal state. The taxonomy of pleurocerids is currently based on late 19th- and early 20th-century studies, which used a typological or morphospecies concept. Most biologists today doubt the validity of many of the currently recognized species; however, this does not stop them from assigning conservation ranks in an attempt to determine which species are imperilled or currently stable. We conducted a phylogenetic analysis of the pleurocerid genus Lithasia using morphological and mitochondrial DNA sequence (mtDNA) data in an attempt to delimit species boundaries and test previous taxonomic schemes. We found that the current taxonomy of Lithasia does not reflect species diversity adequately within the genus, with two new undescribed species being discovered. The conservation status ranks of the new, undescribed species are imperilled and would have been overlooked had we relied on the conventional taxonomy. Additionally, the undescribed species' conservation ranks that were previously apparently secure became vulnerable due to being inappropriately assigned as members of formerly widely distributed species instead of the imperilled status they warrant and vice versa, as some taxa that were considered imperilled are now thought to be modestly stable. Our study suggests that conservation ranks should be considered suspect at best in taxonomically poorly known groups until the taxa are reviewed using modern systematic methods.

Molecular systematics of the North American freshwater bivalve genus Quadrula (Unionidae: Ambleminae) based on mitochondrial ND1 sequences.

A molecular phylogeny of the bivalve genus Quadrula (Unionidae) was constructed based on nucleotide sequences of the mitochondrial ND1 gene. Phylogenetic analysis of 66 specimens representing 17 of the 20 currently recognized taxa within Quadrula, three closely allied species, and 16 outgroup taxa reveals a non-monophyletic Quadrula due to the placement of Tritogonia verrucosa, 'Fusconaia' succissa, and 'Quincuncina' infucata. We suggest that the taxonomic description of the genus Quadrula be expanded to include these species. Within the genus, we continue to recognize three monophyletic species groups (the quadrula, metanvera, and pustulosa species groups), as historically described; however, the pustulosa species group must include 'F.' succissa and 'Quincuncina' infucata. Finally, while our findings place the monotypic genus Tritogonia within Quadrula, its relationship to members within the genus Quadrula remains unresolved.

Molecular phylogeny of a circum-global, diverse gastropod superfamily (Cerithioidea: Mollusca: Caenogastropoda): pushing the deepest phylogenetic limits of mitochondrial LSU rDNA sequences.

The Cerithioidea is a very diverse group of gastropods with ca. 14 extant families and more than 200 genera occupying, and often dominating, marine, estuarine, and freshwater habitats. While the composition of Cerithioidea is now better understood due to recent anatomical and ultrastructural studies, the phylogenetic relationships among families remain chaotic. Morphology-based studies have provided conflicting views of relationships among families. We generated a phylogeny of cerithioideans based on mitochondrial large subunit rRNA and flanking tRNA gene sequences (total aligned data set 1873 bp). Nucleotide evidence and the presence of a unique pair of tRNA genes (i.e., threonine + glycine) between valine-mtLSU and the mtSSU rRNA gene support conclusions based on ultrastructural data that Vermetidae and Campanilidae are not Cerithioidea, certain anatomical similarities being due to convergent evolution. The molecular phylogeny shows support for the monophyly of the marine families Cerithiidae [corrected], Turritellidae, Batillariidae, Potamididae, and Scaliolidae as currently recognized. The phylogenetic data reveal that freshwater taxa evolved on three separate occasions; however, all three recognized freshwater families (Pleuroceridae, Melanopsidae, and Thiaridae) are polyphyletic. Mitochondrial rDNA sequences provide valuable data for testing the monophyly of cerithioidean [corrected] families and relationships within families, but fail to provide strong evidence for resolving relationships among families. It appears that the deepest phylogenetic limits for resolving caenogastropod relationships is less than about 245--241 mya, based on estimates of divergence derived from the fossil record.