MHC structuring and divergent allele advantage in a urodele amphibian: a hierarchical multi-scale approach.

Proteins encoded by extraordinarily polymorphic major histocompatibility complex (MHC) genes are involved in the adaptive immune response. Balancing selection is believed to maintain MHC polymorphism in the long term, although neutral processes also play a role in shaping MHC diversity. However, the relative contribution of these processes is poorly understood. Here we characterized MHC class II variation of a low-dispersal, pond-breeding newt (Triturus carnifex) over a restricted, geographically structured area. We aimed to (1) evaluate the contribution of selection and neutral processes to shaping MHC diversity at two geographic scales, and (2) test for signatures of divergent allele advantage (DAA), which is a potentially important mechanism of balancing selection. The dominant role of selection in shaping MHC variation was suggested by the lack of correlation between MHC and neutral (microsatellite) variation. Although most variation occurred within populations for both types of markers, they differed in the extent of structuring at the two spatial scales. MHC structuring was more pronounced at local scales, suggesting the role of local selection, while structuring was not detectable at a larger scale, possibly due to the effect of balancing selection. Microsatellites showed the opposite pattern. As expected under DAA, the observed genotypes combined more sequence diversity than expected under a random association of alleles. Thus, DAA may contribute to maintaining MHC polymorphism, which is ancient, as supported by signatures of historical positive selection and trans-species polymorphism. Our results point to the importance of a multi-scale approach in studying MHC variation, especially in low-dispersal taxa, which are genetically structured at fine spatial scales.

Rutting behaviour of territorial and nonterritorial male chamois: is there a home advantage?

Males using alternative male mating tactics (AMTs) may express their mating effort in a variety of ways. In polygynous species with limited sexual dimorphism, differences in male aggressiveness may affect mating opportunities. We recorded the behaviour of 8 territorial and 7 nonterritorial male Alpine chamois Rupicapra rupicapra, a nearly monomorphic ungulate, during the 2011 rut in the Gran Paradiso National Park (Italy), to analyse differences in mating effort and mating opportunities between AMTs. The chamois showed a rich behavioural repertoire (31 behavioural patterns), with a prevalence of indirect aggression. Territorial males had higher frequency of aggressive and courtship behaviour than nonterritorial males over the early rut. Later, nonterritorials increased their mating effort, possibly because of reduced competition with dominant males. Territorial males monopolised all observed mating events. Our results support the hypothesis that chamois may assert dominance through intense aggressiveness rather than through horn size or body mass as found in other polygynous ungulates. Most important, differences in mating effort mediated by AMTs resulted in different mating opportunities; these benefits, however, are traded off against greater costs, due to higher levels of hormone metabolites and parasitism. Data on AMTs flexibility, lifetime reproductive success and survival are needed to clarify the mechanisms underlying the evolution and maintenance of AMTs within chamois populations.