Diversification in gravel beaches: A radiation of interstitial clingfish (Gouania, Gobiesocidae) in the Mediterranean Sea.

We report a hitherto unknown radiation within the clingfishes (Gobiesocidae), discovered in one of the best-studied marine biomes, the Mediterranean Sea. The monotypic genus Gouania is a Mediterranean endemic inhabiting the interstices of gravel beaches. Using geometric morphometric analyses, we identified two distinct morphotypes (characterized by a slender and a stout body shape, respectively) among Gouania willdenowi sampled from the three major Mediterranean basins (Eastern, Western and Adriatic). Slender and stout G. willdenowi occurred sympatrically in the Adriatic and the Eastern Mediterranean basins. Only the stout morphotype was found in the Western Mediterranean. Morphotypes were further distinguished by relative eye size and number of vertebrae. Based on mitochondrial and nuclear gene sequences, five highly divergent clades (COI-based K2P distances of 8 to 15%) were identified. The clades represented, respectively, stout Adriatic, slender Adriatic, stout Eastern, slender Eastern and stout Western Mediterranean Gouania. This suggests that the genus Gouania comprises at least five different species. Phylogenetic relationships among clades furthermore imply repeated evolution of convergent morphotypes. The onset of the Gouania radiation was dated to 3.23 (95% HPD 2.08-5.90) mya, and may therefore have overlapped with or followed the Messinian salinity crisis.