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Genes of the major histocompatibility complex (MHC) encode receptor molecules that are responsible for recognition of intracellular and extracellular pathogens (class I and class II genes, respectively) in vertebrates. Given the different roles of class I and II MHC genes, one might expect the strength of selection to differ between these two classes. Different selective pressures may also promote different rates of gene conversion at each class. Despite these predictions, surprisingly few studies have looked at differences between class I and II genes in terms of both selection and gene conversion. Here, we investigated the molecular evolution of MHC class I and II genes in five closely related species of prairie grouse (Centrocercus and Tympanuchus) that possess one class I and two class II loci. We found striking differences in the strength of balancing selection acting on MHC class I versus class II genes. More than half of the putative antigen-binding sites (ABS) of class II were under positive or episodic diversifying selection, compared with only 10% at class I. We also found that gene conversion had a stronger role in shaping the evolution of MHC class II than class I. Overall, the combination of strong positive (balancing) selection and frequent gene conversion has maintained higher diversity of MHC class II than class I in prairie grouse. This is one of the first studies clearly demonstrating that macroevolutionary mechanisms can act differently on genes involved in the immune response against intracellular and extracellular pathogens.

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Females often possess ornaments that appear smaller and duller than homologous traits in males. These ornaments may arise as nonfunctional by-products of sexual selection in males and cause negative viability or fecundity selection in females in proportion to the cost of their production and maintenance. Alternatively, female ornaments may function as signals of quality that are maintained by sexual or social selection. In a 4-year study of 83 female common yellowthroats (Geothlypis trichas) and their 222 young, we found strong viability and fecundity selection on the yellow bib, a carotenoid-based plumage ornament that is a target of sexual selection in males. Females with larger bibs were older, larger and more fecund than females with smaller bibs. However, bib size positively covaried with bib total brightness and carotenoid chroma, aspects of bib coloration that were under negative viability and fecundity selection. Females with more colourful bibs laid fewer eggs in their first clutch, were more likely to suffer total brood loss due to predation and were less likely to return to the study area. Selection against bib coloration limits the value of bib size as a quality indicator in females and may constrain the elaboration of bib attributes in males.

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In socially monogamous species, extra-pair paternity may increase the strength of intersexual selection by allowing males with preferred phenotypes to monopolize matings. Several studies have found relationships between male signals and extra-pair mating, but many others fail to explain variation in extra-pair mating success. A greater appreciation for the role that ecological contingencies play in structuring behavioural processes may help to reconcile contradictory results. We studied extra-pair mating in a spatial context in the common yellowthroat (Geothlypis trichas), a territorial wood warbler. Over the course of 6 years, we observed 158 breeding attempts by 99 males, resulting in a total of 369 nests and 520 sampled nestlings. The spatial distribution of territories varied greatly, with males having between 0 and 10 close neighbours and between three and 39 neighbouring nestlings close enough to represent extra-pair siring opportunities. Both within-pair and extra-pair reproductive success increased with breeding density, but the opportunity for sexual selection and strength of selection varied with density. Total variance in reproductive success was highest at low density and was mostly explained by variation in within-pair success. In contrast, at high density, both within-pair and extra-pair successes contributed substantially to variance in reproductive success. The relationships between plumage and extra-pair mating also varied by density; plumage was under strong sexual selection via extra-pair mating success at high density, but no selection was detected at low density. Thus, ecological factors that structure social interactions can drive patterns of sexual selection by facilitating or constraining the expression of mating preferences.

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The major histocompatibility complex (MHC) plays a central role in innate and adaptive immunity, but relatively little is known about the evolution of the number and arrangement of MHC genes in birds. Insights into the evolution of the MHC in birds can be gained by comparing the genetic architecture of the MHC between closely related species. We used a fosmid DNA library to sequence a 60.9-kb region of the MHC of the greater prairie chicken (Tympanuchus cupido), one of five species of Galliformes with a physically mapped MHC. Greater prairie chickens have the smallest core MHC yet observed in any bird species, and major changes are observed in the number and arrangement of MHC loci. In particular, the greater prairie chicken differs from other Galliformes in the deletion of an important class I antigen binding gene. Analysis of the remaining class IA gene in a population of greater prairie chickens in Wisconsin, USA revealed little evidence for selection at the region responsible for antigen binding.

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Population bottlenecks may reduce genetic variation and potentially increase the risk of extinction. Here, we present the first study to use historic samples to analyse loss of variation at the major histocompatibility complex (MHC), which plays a central role in vertebrate disease resistance. Balancing selection acts on the MHC and could moderate the loss of variation expected from drift; however, in a Wisconsin population of greater prairie-chickens (Tympanuchus cupido), the number of MHC class II B alleles per individual declined by 44% following a population bottleneck, compared to a loss of only 8% at microsatellites. Simulations indicate that drift likely reduced MHC variation at the population level, as well as within individuals by reducing the number of gene copies per individual or by fixing the same alleles across multiple loci. These multiple effects of genetic drift on MHC variation could have important implications for immunity and fitness.

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Females often mate with several different males, which may promote sperm competition and increase offspring viability. However, the potential benefits of polyandry remain controversial, particularly in birds where recent reviews have suggested that females gain few genetic benefits from extra-pair mating. In tree swallows (Tachycineta bicolor), we found that females with prior breeding experience had more sires per brood when paired to genetically similar social mates, and, among experienced females, broods with more sires had higher hatching success. Individual females breeding in two consecutive years also produced broods with more sires when they were more genetically similar to their mate. Thus, experienced females were able to avoid the costs of mating with a genetically similar social mate and realize fitness benefits from mating with a relatively large number of males. This is one of the first studies to show that female breeding experience influences polyandry and female fitness in a natural population of vertebrates. Our results suggest that the benefits of polyandry may only be clear when considering both the number of mates females acquire and their ability to modify the outcome of sexual conflict.

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Comparative analyses suggest that a variety of ecological and behavioural factors contribute to the tremendous variability in extrapair mating among birds. In an analysis of 1010 species of birds, we examined several ecological and behavioural factors in relation to testes size; an index of sperm competition and the extent of extrapair mating. In univariate and multivariate analyses, testes size was significantly larger in species that breed colonially than in species that breed solitarily, suggesting that higher breeding density is associated with greater sperm competition. After controlling for phylogenetic effects and other ecological variables, testes size was also larger in taxa that did not participate in feeding their offspring. In analyses of both the raw species data and phylogenetically independent contrasts, monogamous taxa had smaller testes than taxa with multiple social mates, and testes size tended to increase with clutch size, which suggests that sperm depletion may play a role in the evolution of testes size. Our results suggest that traditional ecological and behavioural variables, such as social mating system, breeding density and male parental care can account for a significant portion of the variation in sperm competition in birds.

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We studied polymorphism in all species of birds that are presently known to show intraspecific variation in plumage colour. At least three main mechanisms have been put forward to explain the maintenance of polymorphism: apostatic, disruptive and sexual selection. All of them make partly different predictions. Our aims were to investigate evolutionary causes and adaptive functions of colour polymorphism by taking into account a number of ecological and morphological features of polymorphic species. Overall, we found 334 species showing colour polymorphism, which is 3.5% of all bird species. The occurrence of colour polymorphism was very high in Strigiformes, Ciconiiformes, Cuculiformes and Galliformes. Phylogenetically corrected analysis using independent contrasts revealed that colour polymorphism was maximally expressed in species showing a daily activity rhythm extended to day/night, living in both open and closed habitats. All these findings support the hypothesis that colour polymorphism probably evolved under selective pressures linked to bird detectability as affected by variable light conditions during activity period. Thus, we conclude that selective agents may be prey, predators and competitors, and that colour polymorphism in birds may be maintained by disruptive selection.

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The evidence that exposure to polychlorinated biphenyls (PCBs) leads to mutations is equivocal and controversial. Using multilocus DNA fingerprinting, we compared the mutation rate of tree swallows (Tachycineta bicolor) nesting at sites with high and low levels of contamination with PCBs. The upper Hudson River, USA, is highly contaminated with PCBs as a result of releases from two capacitor manufacturing plants in Hudson Falls and Fort Edward, New York, USA. Tree swallows nesting nearby have some of the highest known concentrations of PCBs in their tissues of any contemporary bird population (up to 114,000 ng PCB/g tissue). We found no difference in mutation rates between sites in New York with high PCB contamination and reference sites in Wisconsin, USA, and Ontario and Alberta, Canada, with known or presumably low levels of contamination. Thus, the mechanism behind altered reproductive behavior of tree swallows along the upper Hudson River is most likely physiological impairment, such as endocrine disruption, rather than mutation.

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Comparative analyses suggest that a variety of factors influence the evolution of sexual dimorphism in birds. We analyzed the relative importance of social mating system and sperm competition to sexual differences in plumage and body size (mass and tail and wing length) of more than 1,000 species of birds from throughout the world. In these analyses we controlled for phylogeny and a variety of ecological and life-history variables. We used testis size (corrected for total body mass) as an index of sperm competition in each species, because testis size is correlated with levels of extrapair paternity and is available for a large number of species. In contrast to recent studies, we found strong and consistent effects of social mating system on most forms of dimorphism. Social mating system strongly influenced dimorphism in plumage, body mass, and wing length and had some effect on dimorphism in tail length. Sexual dimorphism was relatively greater in species with polygynous or lekking than monogamous mating systems. This was true when we used both species and phylogenetically independent contrasts for analysis. Relative testis size was also related positively to dimorphism in tail and wing length, but in most analyses it was a poorer predictor of plumage dimorphism than social mating system. There was no association between relative testis size and mass dimorphism. Geographic region and life history were also associated with the four types of dimorphism, although their influence varied between the different types of dimorphism. Although there is much interest in the effects of sperm competition on sexual dimorphism, we suggest that traditional explanations based on social mating systems are better predictors of dimorphism in birds.

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Most studies of condition-dependent sexual ornaments have treated such ornaments as single traits. However, sexual ornaments are often composites of several components, each produced by partially independent developmental pathways. Depending on environmental and individual condition, components of these ornaments may reflect different behavioral or physiological properties of an individual. One of the best-known, condition-dependent ornaments is carotenoid-based plumage coloration, which has at least four distinct components: pigment elaboration, patch area, pigment symmetry, and patch area symmetry. Here we examined fitness consequences of variation in individual components of carotenoid ornamentation in male house finches (Carpodacus mexicanus). Over 5 yr and several selection episodes, we studied variation in the plumage components in a large sample (n = 498) of males from a Montana population. The ornament components were partially independent of each other and had distinct fitness consequences. Selection for higher fecundity favored an increase in redness of coloration and a decrease in pigment asymmetry and patch area asymmetry but did not act on patch area itself. In contrast, viability selection favored larger and more symmetrical ornamental patches but did not act on pigment elaboration. Developmental and functional interrelationships among individual components of ornamentation strongly differed between house finch populations. Distinct patterns of selection on individual components of condition-dependent ornaments, combined with partially independent development of components, should favor the evolution of composite sexual traits whose components reliably reflect condition across a wide array of environments.

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Organisms are expected to adjust the sex ratio of their offspring in relation to the relative fitness benefits of sons and daughters. We used a molecular sexing technique that amplifies an intron of the CHD1 gene in birds to examine the sex ratio at egg-laying in socially monogamous tree swallows (Tachycineta bicolor). We examined all individuals in 40 broods (210 young), including all unhatched eggs and nestlings. Thus, the sex ratio we measured was the same as the sex ratio at laying. Overall, the mean sex ratio per brood (+/- SD) was biased significantly towards males (57 +/- 2% male). Within broods, male-biased sex ratios were associated with females in better body condition, and these females were more likely to produce sons in better condition. Tree swallows have one of the highest known levels of extra-pair paternity in birds (38-76% extra-pair young), and, as a consequence, variance in male reproductive success is greater than that of females. Thus, in tree swallows, investment in sons has the potential for higher fitness returns than investment in daughters, assuming that sons in better condition have greater reproductive success.

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