Comparative Functional and Phylogenomic Analyses of Host Association in the Remoras (Echeneidae), a Family of Hitchhiking Fishes

The family Echeneidae consists of eight species of marine fishes that hitchhike by adhering to a wide variety of vertebrate hosts via a sucking disc. While several studies have focused on the interrelationships of the echeneids and the adhesion performance of a single species, no clear phylogenetic hypothesis has emerged and the morphological basis of adhesion remains largely unknown. We first set out to resolve the interrelationships of the Echeneidae by taking a phylogenomic approach using ultraconserved elements. Then, within this framework, we characterized disc morphology through µ-CT analysis, evaluated host specificity through an analysis of host phylogenetic distance, and determined which axes of disc morphological variation are associated with host diversity, skin surface properties, mean pairwise phylogenetic distance (MPD obs.), and swimming regime. We recovered an extremely well-supported topology, found that the specificity of host choice is more variable in a pelagic group and less variable in a reef-generalist group than previously proposed, and that axes of disc morphospace are best explained by models that include host skin surface roughness, host MPD obs., and maximum host Reynolds number. This suggests that ecomorphological diversification was driven by the selection pressures of host skin surface roughness, host specialization, and hydrodynamic regime.