History as a cause of area effects: an illustration from Cerion on Great Inagua, Bahamas

The two parts of this paper work towards the common aim of setting contexts for and documenting explanations based on historical contingencies. The first part is a review of area effects in Cepaea. We discuss the original definitions and explanations, emphasizing the debate of adaptationist vsstochastic approaches, but arguing that the contrast of historical contingency vs. selective fit to environment forms a more fruitful and fundamental context in discussing the origin of area effects. We argue that contingencies of bottlenecks and opening of formerly unsuited habitats may explain the classic area effects of Cepaea better than selectionist accounts originally proposed. The second part is a documentation of an area effect within Cerion columnaon the northern coast of Great Inagua, Bahamas. Historical explanations are often plagued by insufficiency of preserved information, but the Inagua example provides an unusual density of data, with several independent criteria all pointing to the same conclusion. Shells in the area effect are squat and flat-topped in contrast with typical populations of long, thin, tapering shells living both east and west of the area effect. The flat-topped area effect is a result of introgression with a propagule of the C. dimidiatum stock (living on nearby Cuba, and most apically flattened of all Cerion). Fossils of this propagule were found fully cemented into highly indurated fossil soil crusts within the region of the current area effect. Multivariate morphometry, based on complex patterns of covariation, not just intermediacy in single characters, identifies the area effect samples as hybrids between this propagule and typical C. columna. Genetic analysis has identified three unexpected alleles in area effect samples only, and in no other snails of any other Cerion taxon anywhere else on Inagua. We hypothesize that the flattopped area effect did not arise as a selective response to local environments within C. columna, but by introgression from a fortuitously introduced propagule of the C. dimidiaium complex. The unexpected alleles therefore represent genetic phantoms of C. dimidiatum's former presence or are hybrizymes—novel alleles produced by interspecific hybridization