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Understanding the relative role of different evolutionary forces in shaping the level and distribution of functional genetic diversity among natural populations is a key issue in evolutionary and conservation biology. To do so accurately genetic data must be analysed in conjunction with an unambiguous understanding of the historical processes that have acted upon the populations. Here, we focused on diversity at toll-like receptor ( TLR) loci, which play a key role in the vertebrate innate immune system and, therefore, are expected to be under pathogen-mediated selection. We assessed TLR variation within and among 13 island populations (grouped into three archipelagos) of Berthelot's pipit, Anthus berthelotii, for which detailed population history has previously been ascertained. We also compared the variation observed with that found in its widespread sister species, the tawny pipit, Anthus campestris. We found strong evidence for positive selection at specific codons in TLR1 LA, TLR3 and TLR4. Despite this, we found that at the allele frequency level, demographic history has played the major role in shaping patterns of TLR variation in Berthelot's pipit. Levels of diversity and differentiation within and across archipelagos at all TLR loci corresponded very closely with neutral microsatellite variation and with the severity of the bottlenecks that occurred during colonization. Our study shows that despite the importance of TLRs in combating pathogens, demography can be the main driver of immune gene variation within and across populations, resulting in patterns of functional variation that can persist over evolutionary timescales. [ABSTRACT FROM AUTHOR]

Genes of the major histocompatibility complex (MHC) code for receptors that are central to the adaptive immune response of vertebrates. These genes are therefore important genetic markers with which to study adaptive genetic variation in the wild. Next-generation sequencing (NGS) has increasingly been used in the last decade to genotype the MHC. However, NGS methods are highly prone to sequencing errors, and although several methodologies have been proposed to deal with this, until recently there have been no standard guidelines for the validation of putative MHC alleles. In this study, we used the 454 NGS platform to screen MHC class I exon 3 variation in a population of the island endemic Berthelot's pipit ( Anthus berthelotii). We were able to characterise MHC genotypes across 309 individuals with high levels of repeatability. We were also able to determine alleles that had low amplification efficiencies, whose identification within individuals may thus be less reliable. At the population level we found lower levels of MHC diversity in Berthelot's pipit than in its widespread continental sister species the tawny pipit ( Anthus campestris), and observed trans-species polymorphism. Using the sequence data, we identified signatures of gene conversion and evidence of maintenance of functionally divergent alleles in Berthelot's pipit. We also detected positive selection at 10 codons. The present study therefore shows that we have an efficient method for screening individual MHC variation across large datasets in Berthelot's pipit, and provides data that can be used in future studies investigating spatio-temporal patterns and scales of selection on the MHC. [ABSTRACT FROM AUTHOR]

The fauna of oceanic islands provide exceptional models with which to examine patterns of dispersal, isolation and diversification, from incipient speciation to species level radiations. Here, we investigate recent differentiation and microevolutionary change in Berthelot's pipit ( Anthus berthelotii), an endemic bird species inhabiting three Atlantic archipelagos. Mitochondrial DNA sequence data and microsatellite markers were used to deduce probable colonization pathway, genetic differentiation, and gene flow among the 12 island populations. Phenotypic differentiation was investigated based on eight biologically important morphological traits. We found little mitochondrial DNA variability, with only one and four haplotypes for the control region and cytochrome b, respectively. However, microsatellite data indicated moderate population differentiation ( FST = 0.069) between the three archipelagos that were identified as genetically distinct units with limited gene flow. Both results, combined with the estimated time of divergence (2.5 millions years ago) from the Anthus campestris (the sister species), suggest that this species has only recently dispersed throughout these islands. The genetic relationships, patterns of allelic richness and exclusive alleles among populations suggest the species originally colonized the Canary Islands and only later spread from there to the Madeiran archipelago and Selvagen Islands. Differentiation has also occurred within archipelagos, although to a lesser degree. Gene flow was observed more among the eastern and central islands of the Canaries than between these and the western islands or the Madeiran Islands. Morphological differences were also more important between than within archipelagos. Concordance between morphological and genetic differentiation provided ambiguous results suggesting that genetic drift alone was not sufficient to explain phenotypic differentiation. The observed genetic and morphological differences may therefore be the result of differing patterns of selection pressures between populations, with Berthelot's pipit undergoing a process of incipient differentiation. [ABSTRACT FROM AUTHOR]