PATTERNS OF MORPHOLOGICAL DIVERSITY AMONG AND WITHIN ARCID BIVALVE SPECIES PAIRS SEPARATED BY THE ISTHMUS OF PANAMA

Abstract Geminate species are morphologically similar sister-species found on either side of the Isthmus of Panama. The existence of all geminates in the tropical Eastern Pacific ocean and the Caribbean Sea is most often explained by vicariance: closure of the Central American Seaway 3.1 to 3.5 Ma simultaneously isolated populations of species with amphi-American distributions. In this paper, we test the potential of morphological measurements for discriminating between Recent geminate species pairs from three genera (Arca, Arcopsis, and Barbatia) in the bivalve family Arcidae and examine the prospects for distinguishing nominal species in the fossil record. Fourteen morphological variables were used to characterize shell shape and multivariate methods were used to discriminate between five Recent species pairs. Collection sites were also used as a priori groups for discrimination to describe patterns of intraspecific morphological variation and to evaluate differences among samples from different geographic regions. On average, 84 percent of specimens within geminate pairs are classified correctly following five separate discriminant analyses with nominal species as the grouping variable. Although all but one arcid species pair are discriminated with high statistical significance, some collection sites within species are highly morphologically distinct. Overall, a large proportion of specimens from each collection locality (79 percent on average) can be classified correctly to site although no single site possessed a multivariate centroid that was significantly different from all other conspecific centroids. The distinctiveness of some collection sites, however, raises the possibility that some nominal species may harbor cryptic species, indicating the need for wider geographic surveys of both molecular and morphological variation within geminate species pairs. The eigenvalue coefficients derived from the Recent samples of one geminate pair (Arca mutabilis and A. imbricata) were used to assess the potential for identifying arcid species in the fossil record. Discriminant analyses of fossil Arca indicate that the forms that characterize Recent A. mutabilis and A. imbricata are present in the fossil record as far back as the Late Early Miocene, in the Cantaure Formation of Venezuela. Because a deep water connection between the Eastern Pacific and Western Atlantic existed until the Middle Miocene, the morphological differences associated with Recent A. mutabilis and A. imbricata likely existed well before the rising Isthmus affected ocean circulation patterns in tropical America. Therefore, despite great overall morphological similarity, these putative geminate species likely have a time of divergence that is at least four times older than final seaway closure. The geographic distribution of fossils also suggests that morphological forms associated with each Recent species had amphi-American distributions both before and after isthmus formation but are now geographically restricted to either side of the isthmus in the Recent fauna.