title: Phylogenetic and morphometric analyses reveal ecophenotypic plasticity in freshwater mussels Obovaria jacksoniana and Villosa arkansasensis (Bivalvia: Unionidae).
abstract: Freshwater mollusk shell morphology exhibits clinal variation along a stream continuum that has been termed the Law of Stream Distribution. We analyzed phylogenetic relationships and morphological similarity of two freshwater mussels (Bivalvia: Unionidae), Obovaria jacksoniana and Villosa arkansasensis, throughout their ranges. The objectives were to investigate phylogenetic structure and evolutionary divergence of O. jacksoniana and V. arkansasensis and morphological similarity between the two species. Our analyses were the first explicit tests of phenotypic plasticity in shell morphologies using a combination of genetics and morphometrics. We conducted phylogenetic analyses of mitochondrial DNA (1416 bp; two genes) and morphometric analyses for 135 individuals of O. jacksoniana and V. arkansasensis from 12 streams. We examined correlations among genetic, morphological, and spatial distances using Mantel tests. Molecular phylogenetic analyses revealed a monophyletic relationship between O. jacksoniana and V. arkansasensis. Within this O. jacksoniana/V. arkansasensis complex, five distinct clades corresponding to drainage patterns showed high genetic divergence. Morphometric analysis revealed relative differences in shell morphologies between the two currently recognized species. We conclude that morphological differences between the two species are caused by ecophenotypic plasticity. A series of Mantel tests showed regional and local genetic isolation by distance. We observed clear positive correlations between morphological and geographic distances within a single drainage. We did not observe correlations between genetic and morphological distances. Phylogenetic analyses suggest O. jacksoniana and V. arkansasensis are synonomous and most closely related to a clade composed of O. retusa, O. subrotunda, and O. unicolor. Therefore, the synonomous O. jacksoniana and V. arkansasensis should be recognized as Obovaria arkansasensis (Lea 1862) n. comb. Phylogenetic analyses also showed relative genetic isolation among drainages, suggesting no current gene flow. Further investigation of in-progress speciation and/or cryptic species within O. arkansasensis is warranted followed by appropriate revision of conservation management designations. In this study, we found Obovaria jacksoniana and Villosa arkansasensis are synonomous. We suggest that morphological differences between the two species are caused by ecophenotypic plasticity, where V. arkansasensis is the upstream morphotype and O. jacksoniana is the downstream morphotype of a single species.
authors.affiliation: Environmental Sciences Graduate Program, Arkansas State University P.O. Box 877, State University, Arkansas, 72467 ; Department of Zoology, Miami University 700 High Street, 212 Pearson Hall, Oxford, Ohio, 45056.
authors.forename: Kentaro
authors.initials: K
authors.lastname: Inoue
authors.affiliation: Environmental Sciences Graduate Program, Arkansas State University P.O. Box 877, State University, Arkansas, 72467 ; Department of Biological Sciences, Eastern Kentucky University 521 Lancaster Avenue, 235 Moore Building, Richmond, Kentucky, 40475.
authors.forename: David M
authors.initials: DM
authors.lastname: Hayes
authors.affiliation: Department of Biological Sciences, Arkansas State University P.O Box 599, State University, Arkansas, 72467.
authors.forename: John L
authors.initials: JL
authors.lastname: Harris
authors.affiliation: Environmental Sciences Graduate Program, Arkansas State University P.O. Box 877, State University, Arkansas, 72467 ; Department of Biological Sciences, Arkansas State University P.O Box 599, State University, Arkansas, 72467 ; Biology Department, University of Massachusetts at Boston 100 Morrissey Boulevard, Boston, Massachusetts, 02125.
authors.forename: Alan D
authors.initials: AD
authors.lastname: Christian
keywords: Isolation by distance; Law of Stream Distribution; Mantel test; Ortmann's law; mitochondrial DNA; phenotypic plasticity