Patterns and correlates of extrapair paternity in American redstarts (Setophaga ruticilla)

We examined correlates and hypotheses pertaining to extrapairfertilizations in socially monogamous American redstarts (Setophagaruticilla). DNA fingerprinting revealed extrapair fertilizationin 59% of broods (19 of 32), involving 40% of nestlings (43of 108). Fewer broods than expected had mixed paternity, asdetermined from a binomial distribution of extrapair young inthe population. This result is consistent with the "good genes"hypothesis, but not with the "genetic diversity" hypothesis.There was a negative association between the age of putativefathers and the proportion of extrapair young in their broods.Irrespective of age, males with prior residency were cuckoldedless often than males new to the study area. Extrapair fatherswere' immediate neighbors in 7 of 10 cuckolded broods whereall neighbors were sampled. Males were more likely to sire offspringin the territories of younger neighbors than in those of olderneighbors. Plumage characteristics of adult males, breedingsynchrony of females, and breeding densities were not significantlyassociated with cuckoldry. Realized reproductive gain from cuckoldrywas small because of high nest predation in our area. Extrapairfertilizations allowed one-quarter of males whose own nestshad failed to achieve some reproductive success. Only 2 of 17males whose own nests were successful also had extrapair young.There was no egg dumping by females. We conclude that male ageand prior residency were predictors of cuckoldry in Americanredstarts. In the context of the heavy predation experiencedby our birds, extrapair fertilizations allowed many males tosalvage some reproductive success and did not increase the varianceof success across males     

Diurnal and individual variability in the foraging behavior of American redstarts (Setophaga ruticilla)

Summary The foraging behavior of American redstarts, Setophaga ruticilla (Aves: Parulidae), was examined from dawn to dusk in the nestling phase of their breeding cycle in a northern hardwoods forest in New Hampshire, USA. Based on a sample size of nearly 3000 foraging acts, we found that redstarts hawked more and foraged lower at midday than early in the morning, coincident with a statistically significant increase in the abundance of flying insects in the lower strata of the forest at that time of day. Furthermore, individual redstarts of like sex on adjacent territories and members of mated pairs on the same territory often foraged at significantly different heights, used significantly different combinations of foraging methods, or both. A complex of factors, including the diurnal changes in insect distribution and availability, influences how birds search for and capture prey, and results in intraspecific variability which complicates studies of species-typical foraging patterns.     

Prevalence of hematozoa in overwintering American redstarts (Setophaga ruticilla): no evidence for local transmission

We examined American redstarts (Setophaga ruticilla) for protozoan blood parasites on their wintering grounds to determine whether transmission of these parasites occurs prior to spring migration. A total of 73 blood smears from 37 birds were examined for presence and intensity of infection. Thirty-six birds were sampled in the fall, soon after arriving from northern breeding grounds, and the spring prior to departure. Two (5%) of the samples collected in the fall were positive for Haemoproteus fringillae and one (3%) had detectable infections of Trypanosoma avium. Individuals infected with H. fringillae were hatching year redstarts sampled in September and October. Intensity of infection was 78 and < 1 infected erythrocytes per 10,000 erythrocytes, respectively. None of the birds had detectable infections when resampled prior to spring migration the following March.     

Songs of American Redstarts (Setophaga ruticilla): Sequencing Rules and their Relationships to Repertoire Size

In order to determine the rules of sequencing of songs used by American redstarts, we related Markovian and hierarchical models to recordings obtained from free-living males. In the smaller repertoires of three or four songs, low order Markov chain models fitted the data. 9 of the 10 sequences so examined were first-order, and the last was second-order. Larger repertoires of 6 and 8 songs were hierarchical in organization with subsets of songs having independent sequencing rules. Most samples of singing were stationary in their transition rules over periods of several days: non-stationarity was sometimes associated with a change in the number of songs forming the sequence, or in repetitions of songs. We examine causal models of song sequencing and conclude that our results generally favor competition models, although some sequential dependencies may also apply. Hierarchical organization in the serial repertoires of American redstarts may reflect developmental influences rather than effects of repertoire size itself.     

Short‐term changes in body condition in relation to habitat and rainfall abundance in American redstarts Setophaga ruticilla during the non‐breeding season

In migratory birds, environmental conditions during the stationary period of the non‐breeding season are crucial to consider because they ultimately affect the fitness of individuals by influencing their subsequent migration, breeding success and survival. Although a few studies have investigated the influence of non‐breeding habitat on the capacity of individuals to cope with long‐term seasonal rainfall fluctuations, it remains unknown how habitat quality and variations in rainfall abundance–at a monthly scale–interact to affect non‐breeding condition of migrating birds. In this study, we examined the influence of monthly changes in rainfall abundance on body condition of non‐breeding female redstarts Setophaga ruticilla living either in a high quality habitat (mangrove) or in a low quality habitat (scrub). Body condition of both mangrove and scrub redstarts showed important variations over the study period, demonstrating for the first time that body condition of non‐breeding female redstarts can change rapidly in response to short‐term fluctuations in rainfall. Importantly, we found that female redstarts living in mangrove were usually in better condition during periods of low rainfall compared to females living in scrub. However, body condition did not differ between mangrove females and scrub females during an episode of frequent, heavy precipitation. Importantly, our study also demonstrated that the duration of a perturbation is an important determinant of body condition in redstarts since a prolonged drought resulted in similar low body condition for birds from both habitats. Age was not correlated with body condition whatever the habitat and the rainfall conditions. Our results demonstrate that high quality habitat can temporarily reduce the deleterious effect of a short‐term drought on body condition, but also, that a habitat of low quality does not constrain individuals when climatic conditions are optimal.     

Plumage colour is related to ectosymbiont load during moult in the serin, Serinus serinus: an experimental study

Various hypotheses propose that plumage ornamentation is a reliable indicator of the health or resistance to parasites and illness of individuals. The impact of endoparasites on plumage brightness has only recently been demonstrated experimentally. We tested the impact of ectosymbionts, in particular feather mites, on plumage brightness, using 2 years of observational data and experiments in the field. The abundance of feather mites during moult was negatively correlated with brightness and saturation of plumage coloration developed by male serins. The application of an insecticide before moult resulted in experimental males developing a brighter plumage than control individuals. Experimental adult males, but not juveniles, also developed more saturated plumages in one of the years, but did not differ from controls in the other year. This is the first experimental demonstration that ectosymbionts (including mites) have a negative impact on the characteristics of the plumage developed and consequently can signal the healthiness of their hosts. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Melanocortin 1-receptor (MC1R) mutations are associated with plumage colour in chicken

The co-segregation of plumage colour and sequence polymorphism in the melanocortin 1-receptor gene (MC1R) was investigated using an intercross between the red junglefowl and White Leghorn chickens. The results provided compelling evidence that the Extended black (E) locus controlling plumage colour is equivalent to MC1R. E/MC1R was assigned to chromosome 11 with overwhelming statistical support. Sequence analysis indicated that the E92K substitution, causing a constitutively active receptor in the sombre mouse, is the most likely causative mutation for the Extended black allele carried by the White Leghorn founders in this intercross. The MC1R sequence associated with the recessive buttercup (ebc) allele indicated that this allele evolved from a dominant Extended black allele as it shared the E92K and M71T substitutions with some E alleles. It also carried a third missense mutation H215P which thus may interfere with the constitutive activation of the receptor caused by E92K (and possibly M71T).     

Missense SNP of the MC1R gene is associated with plumage variation in the Gyrfalcon (Falco rusticolus)

A single nucleotide polymorphism (MC1R: c.376A>G) in the MC1R gene was found to be highly correlated with pigment phenotype in the Gyrfalcon. Homozygous genotypes c.376GG and c.376AA were found to dominate the extreme white and dark plumage types respectively, and heterozygotes occurred mainly in intermediate phenotypes. However, some heterozygotes were associated with extreme phenotypes, indicating that melanism/albinism might also involve other loci.     

Reconciling ecogeographical rules: rainfall and temperature predict global colour variation in the largest bird radiation

Ecogeographical rules that associate climate with organismal form and function can reveal patterns of climatic adaptation. Two rules link animal coloration with climate: Gloger's rule (darker coloration where wet and warm), and Bogert's rule (darker coloration where cold). Whereas Gloger's rule was proposed for endotherms, and Bogert's rule for ectotherms, both rules may apply more broadly, despite their seemingly opposing effects. Here, we test this contradiction on a global scale across passerine birds. Consistent with Gloger's rule, birds were darker in wetter areas and, following Bogert's rule, lighter where warm, although birds became lighter again at very low temperatures. Rainfall and temperature had antagonistic or additive effects depending on their pattern of covariation, and this predicted whether birds followed the rules. We integrate both rules into a general framework to explain heterogeneity in climatic effects on coloration, which has implications to understand patterns of diversification, climatic adaptation and climate change impacts.     

Predation is associated with variation in colour pattern, but not body shape or colour reflectance, in a rainbowfish (Melanotaenia australis)

1. In freshwater fishes, inter-population variation in male phenotype is often associated with differences in predation intensity, but these effects can be difficult to disentangle from environmental influences. 2. The western rainbowfish Melanotaenia australis exhibits marked sexual dimorphism - females are plain with a slender body, while males have striking coloration and are deeper in the body. Male traits differ in expression among populations, but this has not been described or explored in the literature. 3. This paper describes a study designed to test for geographic structuring of male phenotype in M. australis and to determine whether between-population variation in male phenotype is attributable to variation in predation regime, after accounting for environment. 4. We collected data describing habitat, and the size, activity and abundance of predators at sites containing M. australis populations. We then used photography, spectrometry and geometric morphometrics to describe colour pattern, spectral reflectance and body shape in males from these populations. Finally, we used permutation-based multivariate statistics to partition variance in these traits according to environment and predation regime. 5. Downstream environments posed higher predation risk to M. australis. Furthermore, males from these sites consistently exhibited larger cheek spots and fewer coloured lateral stripes than those from upstream sites. Variation in predation regime accounted for a significant proportion of the total variance in these traits (30·9%), after controlling for the effects of environment. 6. Variation in predation regime did not explain variation in reflectance or shape. Environmental variation, however, explained a significant portion of the total variance in reflectance (74·9%), and there was a strong trend towards it explaining a portion of the total variance in body shape (34·9%). 7. We conclude that natural selection by predators may be an important determinant of the evolution of colour pattern variation in M. australis, but not of that of body shape or colour reflectance. 8. Further study of M. australis will complement existing models, which show complex relationships between predation regime, environment and phenotype. Understanding these relationships is prerequisite to predicting the evolution of phenotypic variation in natural systems.     

Geographic patterns of genetic differentiation and plumage colour variation are different in the pied flycatcher (Ficedula hypoleuca)

The pied flycatcher is one of the most phenotypically variable bird species in Europe. The geographic variation in phenotypes has often been attributed to spatial variation in selection regimes that is associated with the presence or absence of the congeneric collared flycatcher. Spatial variation in phenotypes could however also be generated by spatially restricted gene flow and genetic drift. We examined the genetic population structure of pied flycatchers across the breeding range and applied the phenotypic Q _ST ( P _ST)– F _ST approach to detect indirect signals of divergent selection on dorsal plumage colouration in pied flycatcher males. Allelic frequencies at neutral markers were found to significantly differ among populations breeding in central and southern Europe whereas northerly breeding pied flycatchers were found to be one apparently panmictic group of individuals. Pairwise differences between phenotypic ( P _ST) and neutral genetic distances ( F _ST) were positively correlated after removing the most differentiated Spanish and Swiss populations from the analysis, suggesting that genetic drift may have contributed to the observed phenotypic differentiation in some parts of the pied flycatcher breeding range. Differentiation in dorsal plumage colouration however greatly exceeded that observed at neutral genetic markers, which indicates that the observed pattern of phenotypic differentiation is unlikely to be solely maintained by restricted gene flow and genetic drift.     

Genetic structure and differentiation of the Japanese extremely long-tailed chicken breed (Onagadori), associated with plumage colour variation: suggestions for its management and conservation

The Onagadori is a distinguished chicken breed that is characterized by an extremely long tail in the male. In this breed, three different plumage colour varieties have been developed (black-breasted white, black-breasted red and white) in which the black-breasted white is believed to be the original colour of the Onagadori, based on historical records. To establish a conservation strategy, 176 birds were genotyped for autosomal microsatellites. Significant genetic distinctness was found between the original (black-breasted white) and two derivative varieties ( F _ST = 0.091 and 0.093). At the same time, a Bayesian model-based clustering revealed that the majority of individuals belonging to the black-breasted red and white varieties had an extremely low proportion of the genome shared with the original type (black-breasted white). This suggests that derivative varieties were created by crossing with other breeds, with low introgression of the original-type genome. We propose that the three plumage colour varieties should be treated as separate genetic units in a conservation programme.     

Sequence variation in the coding region of the melanocortin-1 receptor gene (MC1R) is not associated with plumage variation in the blue-crowned manakin (Lepidothrix coronata)

Avian plumage traits are the targets of both natural and sexual selection. Consequently, genetic changes resulting in plumage variation among closely related taxa might represent important evolutionary events. The molecular basis of such differences, however, is unknown in most cases. Sequence variation in the melanocortin-1 receptor gene (MC1R) is associated with melanistic phenotypes in many vertebrate taxa, including several avian species. The blue-crowned manakin (Lepidothrix coronata), a widespread, sexually dichromatic passerine, exhibits striking geographic variation in male plumage colour across its range in southern Central America and western Amazonia. Northern males are black with brilliant blue crowns whereas southern males are green with lighter blue crowns. We sequenced 810 bp of the MC1R coding region in 23 individuals spanning the range of male plumage variation. The only variable sites we detected among L. coronata sequences were four synonymous substitutions, none of which were strictly associated with either plumage type. Similarly, comparative analyses showed that L. coronata sequences were monomorphic at the three amino acid sites hypothesized to be functionally important in other birds. These results demonstrate that genes other than MC1R underlie melanic plumage polymorphism in blue-crowned manakins.     

The lavender plumage colour in Japanese quail is associated with a complex mutation in the region of MLPH that is related to differences in growth, feed consumption and body temperature

ABSTRACT: BACKGROUND: The lavender phenotype in quail is a dilution of both eumelanin and phaeomelanin in feathers that produces a blue-grey colour on a wild-type feather pattern background. It has been previously demonstrated by intergeneric hybridization that the lavender mutation in quail is homologous to the same phenotype in chicken, which is caused by a single base-pair change in exon 1 of MLPH. RESULTS: In this study, we have shown that a mutation of MLPH is also associated with feather colour dilution in quail, but that the mutational event is extremely different. In this species, the lavender phenotype is associated with a non-lethal complex mutation involving three consecutive overlapping chromosomal changes (two inversions and one deletion) that have consequences on the genomic organization of four genes (MLPH and the neighbouring PRLH, RAB17 and LRRFIP1). The deletion of PRLH has no effect on the level of circulating prolactin. Lavender birds have lighter body weight, lower body temperature and increased feed consumption and residual feed intake than wild-type plumage quail, indicating that this complex mutation is affecting the metabolism and the regulation of homeothermy. CONCLUSIONS: An extensive overlapping chromosome rearrangement was associated with a non-pathological Mendelian trait and minor, non deleterious effects in the lavender Japanese quail which is a natural knockout for PRLH.     

Global changes and animal phenotypic responses: melanin-based plumage redness of scops owls increased with temperature and rainfall during the last century

The ecological effects of global climate changes include shifts of species'' distribution and changes in migration strategies and phenotype. Colour polymorphism, which can be envisaged as a species'' evolutionary response to alternating conditions or to a wide range of habitats, may be affected by climate changes as well. The scops owl (Otus scops) shows two main colour morphs, dark- and pale-reddish, as well as intermediate morphs. We investigated temporal trends in an index of plumage colour of Italian scops owls from museum collections (1870–2007). We found a significant increase in plumage redness over the last century, which was correlated with an increase in temperature and rainfall of the years before specimen collection. However, the temporal increase in plumage redness persisted after controlling for climatic variables, suggesting that other environmental factors could be involved. Our study indicates that ongoing climate changes might have either shifted the selective balance between colour morphs, or differentially affected migration and movement patterns of colour morphs.     

Evolutionary significance of geographic variation in a plumage-based foraging adaptation: an experimental test in the slate-throated redstart (Myioborus miniatus)

Geographic variation in the plumage pattern of birds is widespread but poorly understood, and in very few cases has its evolutionary significance been investigated experimentally. Neotropical warblers of the genus Myioborus use their contrasting black-and-white plumage to flush insect prey during animated foraging displays. Although previous experimental work has demonstrated that white plumage patches are critical to flush-pursuit foraging success, the amount of white in the plumage shows considerable interspecific and intraspecific geographic variation. We investigated the evolutionary significance of this geographic variation by experimentally decreasing or increasing the amount of white in the tail of slate-throated redstarts (Myioborus miniatus comptus) from Monteverde, Costa Rica, to mimic the natural extremes of tail pattern variation in this species. In addition to measuring the effects of plumage manipulation on foraging performance, we performed field experiments measuring the escape response of a common insect prey species (an asilid fly) using model redstarts representing four different Myioborus plumage patterns. Our experiments were designed to test four hypotheses that could explain geographic variation in plumage pattern. Compared to controls, experimental birds with reduced-white tails that mimic the plumage pattern of M. miniatus hellmayri of Guatemala showed significant reductions in flush-pursuit foraging performance. In contrast, the addition of white to the tail to mimic the plumage pattern of M. miniatus verticalis of Bolivia had no significant effect on foraging performance of Costa Rican redstarts. In field experiments with asilid flies, model redstarts simulating the plumage of M. miniatus comptus of Costa Rica and M. miniatus verticalis of Bolivia elicited greater responses than did models of other Myioborus taxa with either less or more white in the plumage. The results of our experiments with both birds and insects allow us to reject two hypotheses for geographic variation in plumage pattern: (1) that geographic variation is a nonadaptive result of genetic drift, and (2) that selection for enhanced flush-pursuit foraging performance generally favors increased white in the plumage, but evolutionary trade-offs constrain the evolution of extensive patches of white in some geographic regions. Instead, our results suggest that geographic variation in the plumage pattern of Myioborus redstarts reflects adaptation to regional habitat characteristics that enhances flush-pursuit foraging performance.     

Genome‐wide variation in DNA methylation is associated with stress resilience and plumage brightness in a wild bird

Individuals often differ in their ability to cope with challenging environmental and social conditions. Evidence from model systems suggests that patterns of DNA methylation are associated with variation in coping ability. These associations could arise directly if methylation plays a role in controlling the physiological response to stressors by, among other things, regulating the release of glucocorticoids in response to challenges. Alternatively, the association could arise indirectly if methylation and resilience have a common cause, such as early‐life conditions. In either case, methylation might act as a biomarker for coping ability. At present, however, relatively little is known about whether variation in methylation is associated with organismal performance and resilience under natural conditions. We studied genome‐wide patterns of DNA methylation in free‐living female tree swallows (Tachycineta bicolor) using methylated DNA immunoprecipitation (MeDIP) and a tree swallow genome that was assembled for this study. We identified areas of the genome that were differentially methylated with respect to social signal expression (breast brightness) and physiological traits (ability to terminate the glucocorticoid stress response through negative feedback). We also asked whether methylation predicted resilience to a subsequent experimentally imposed challenge. Individuals with brighter breast plumage and higher stress resilience had lower methylation at differentially methylated regions across the genome. Thus, widespread differences in methylation predicted both social signal expression and the response to future challenges under natural conditions. These results have implications for predicting individual differences in resilience, and for understanding the mechanistic basis of resilience and its environmental and social mediators.