Patella aspera and P-ulyssiponensis: genetic evidence of speciation in the North-east Atlantic

The group of subspecies of Patella ulyssiponensis, described by Christiaens, was widely known as Patella aspera until recently. The group extends throughout the Mediterranean, on all Macaronesian islands, along the North African coast, and in Europe, as far north as southern Norway. Throughout its range it displays great variation in shell sculpture and colour. The aim of this work was to re-examine the various subspecies proposed by Christiaens and, genetically, to test the hypothesis that European continental populations belong to a different species than the one composed by Macaronesian (north-west African) populations. In the present work, this group was studied by allozyme electrophoresis of 21 loci and by six morphological variables. The monomorphic locus of malate dehydrogenase (Mdh-1) was found to be diagnostic for distinguishing European continental populations from those of north-west African archipelagos, confirming the specific status of both groups. The allele observed at this locus in the Macaronesian populations was novel, while European continental populations showed a plesiomorphic one, shared with all other north-west Atlantic patellids with the exception of Patina pellucida, suggesting a more recent origin of the Macaronesian species from the continental forms. Both species showed a genetic identity of 0.730±0.061, which allowed rough estimations of 6.5-3 Mya since speciation. The subspecies Patella ulyssiponensis deserta described by Christiaens was not confirmed by our genetic data. Other loci (glyceraldehyde-3-phosphate dehydrogenase and the second locus of lactate dehydrogenase) were partially diagnostic, in which both species showed different most common alleles. Morphologically, both species are easily recognised by shell characters and the results agree with previous findings, that continental populations are more homogeneous in shell morphology and radula characters than populations from the Macaronesia. This work supports retention of the earliest valid name, Patella ulyssiponensis Gmelin, with Lisbon, Portugal as type locality, for the European continental species, and Patella aspera Roding for the Macaronesian populations. Population subdivision within species was measured by theta, the estimator of F-st, showing in both P. aspera and P. ulyssiponensis a high degree of genetic structuring (θ=0.226 and 0.182, respectively) mostly explained by the large distances separating the populations within species