Identifying zones of phenetic compression in West Mediterranean butterflies (Satyrinae): refugia, invasion and hybridization

Aim Distinct insular populations are generally considered important units for conservation. In islandmainland situations, unidirectional introgressive gene flow from the most abundant, typically continental, populations into the smaller island populations can erase native insular genetic units. As an indication of threat, the concept of phenetic slope is developed, a measure proportional to differentiation and to geographical proximity. Location The Western Mediterranean, including the following islands: Sardinia, Sicily, Corsica, Balearics, circum-Italian, circum-Sicilian and circum-Sardo-Corsican archipelagos. Eastern Europe is included for comparison. Methods Geometric morphometrics was applied to 2392 male genitalia of seven butterfly species groups. Geographic Information System techniques were used to depict the pattern in the distribution of morphotypes. The slope of variation in genital shape was computed to highlight geographical areas showing abrupt morphological changes. Correlation analyses were performed between the mean slope values across sea straits separating islands and nearest sources and ecological traits of the species that underlie their colonization and migration capacity. Results Phenetic slope analysis has revealed that the strait of Messina and the northern Tyrrhenian Sea support particularly contrasting populations. In these areas, mean slopes for species also correlated with certain ecological traits of the species. Sardinia emerges as the most stable refugium for ancestral mediterranean populations. Main conclusions There is strong support for the hypothesis that Italy has experienced invasion by populations from Eastern Europe with postglacial expansion of these populations across Italy. However, propagules are impeded from invading islands by the expanse of sea straits. Even so, sea straits are not invariably barriers. Our results suggest that wind direction in combination with habitat occupancy may have maintained ancestral insular populations in key locations distinguished by phenetic compression. We conclude that native insular populations acting as barriers to introgression in the areas showing particularly steep phenetic slopes deserve attention in conservation programmes.