Major histocompatibility complex class II variation in bottlenose dolphin from Adriatic Sea: inferences about the extent of balancing selection

The bottlenose dolphin (Tursiops truncatus) is the most common cetacean species worldwide and the only marine mammal species resident in the Croatian part of the Adriatic Sea. To gain insight into genetic diversity of bottlenose dolphins at adaptively important loci relevant to conservation, we analysed the polymorphism of major histocompatibility complex (MHC) genes, which play a key role in pathogen confrontation and clearance. Specifically, we examined the diversity of MHC class II DRA, DQA and DQB alleles in 50 bottlenose dolphins from the Adriatic Sea collected between 1997 and 2011 and in 12 animals from other Mediterranean locations. Notable variation in DQA, DQB and three-locus haplotypes was found, with all 10 DQA and 12 DQB alleles encoding unique protein products. Analysis of the ratio of non-synonymous to synonymous substitution rates suggests that positive selection acts at both highly variable loci. Phylogenetic analyses revealed trans-species polymorphism at the DQB locus, strongly indicating the influence of balancing selection in the long term. In fact, the balancing selection observed in bottlenose dolphins is higher than that reported for most other cetaceans and comparable to that seen in terrestrial mammals.