Breeding Experience and not Age Modulates the Song Development of Pied Flycatchers (Ficedula hypoleuca)

In songbirds, the development of the species‐specific adult song involves a learning process that varies in extension. In species that incorporate new song elements throughout life (open‐ended learners), variation in male song composition could be the result of either age or breeding experience. Using data from 16 yr of fieldwork on pied flycatchers (Ficedula hypoleuca), we aimed to disclose the individual contribution of these two factors on the species song characteristics, as well as their relation with morphology and plumage color changes. Finally, we explored whether any of the song or physical features could predict the probability of males returning to the breeding site. We found that the song characteristics of the first‐time breeders did not differ between age classes, except for the total number of syllables per song, which was higher in the 1‐year‐old than in the 2‐year‐old males. However, we found that song variables associated with complexity (song and sample versatilities and repertoire size), increased significantly from the first to the second breeding season. Males showed delayed plumage maturation, with 1‐year‐old males being browner than the 2‐year‐old males independently of their breeding experience. Morphology, however, was not affected by age or breeding experience. The probability that males returned to the breeding site was not associated with song or physical features. Considering that some song learning occurs during the breeding period and that some males may skip the first breeding season, selective pressures may have been established for song complexity to be an honest indicator of breeding experience rather than age.     

Narrow hybrid zones in spite of very low population differentiation in neutral markers in an island bird species complex

Patterns of phenotypic and genic frequencies across hybrid zones provide insight into the origin and evolution of reproductive isolation. The Reunion grey white‐eye, Zosterops borbonicus, exhibits parapatrically distributed plumage colour forms across the lowlands of the small volcanic island of Reunion (Mascarene archipelago). These forms meet and hybridize in regions that are natural barriers to dispersal (rivers, lava fields). Here, we investigated the relationship among patterns of differentiation at neutral genetic (microsatellite) markers, phenotypic traits (morphology and plumage colour) and niche characteristics across three independent hybrid zones. Patterns of phenotypic divergence revealed that these hybrid zones are among the narrowest ever documented in birds. However, the levels of phenotypic divergence stand in stark contrast to the lack of clear population neutral genetic structure between forms. The position of the hybrid zones coincides with different natural physical barriers, yet is not associated with steep changes in vegetation and related climatic variables, and major habitat transitions are shifted from these locations by at least 18 km. This suggests that the hybrid zones are stabilized over natural dispersal barriers, independently of environmental boundaries, and are not associated with niche divergence. A striking feature of these hybrid zones is the very low levels of genetic differentiation in neutral markers between forms, suggesting that phenotypic divergence has a narrow genetic basis and may reflect recent divergence at a few linked genes under strong selection, with a possible role for assortative mating in keeping these forms apart.     

Sexual selection,speciation and constraints on geographical range overlap in birds

The role of sexual selection as a driver of speciation remains unresolved, not least because we lack a clear empirical understanding of its influence on different phases of the speciation process. Here, using data from 1306 recent avian speciation events, we show that plumage dichromatism (a proxy for sexual selection) does not predict diversification rates, but instead explains the rate at which young lineages achieve geographical range overlap. Importantly, this effect is only significant when range overlap is narrow (< 20%). These findings are consistent with a ‘differential fusion’ model wherein sexual selection reduces rates of fusion among lineages undergoing secondary contact, facilitating parapatry or limited co‐existence, whereas more extensive sympatry is contingent on additional factors such as ecological differentiation. Our results provide a more mechanistic explanation for why sexual selection appears to drive early stages of speciation while playing a seemingly limited role in determining broad‐scale patterns of diversification.     

Conditions on the Mexican moulting grounds influence feather colour and carotenoids in Bullock's orioles (Icterus bullockii)

Carotenoid‐based plumage coloration plays a critical role for both inter‐ and intrasexual communication. Habitat and diet during molt can have important consequences for the development of the ornamental signals used in these contexts. When molt occurs away from the breeding grounds (e.g., pre‐alternate molt on the wintering grounds, or stopover molt), discerning the influence of habitat and diet can be particularly important, as these effects may result in important carryover effects that influence territory acquisition or mate choice in subsequent seasons. Several species of songbirds in western North America, including the Bullock's oriole (Icterus bullockii), migrate from the breeding grounds to undergo a complete prebasic (post‐breeding) molt at a stopover site in the region affected by the Mexican monsoon climate pattern. This strategy appears to have evolved several times independently in response to the harsh, food‐limited late‐summer conditions in the arid West, which contrast strongly with the high productivity driven by heavy rains that is characteristic of the Mexican monsoon region. Within this region, individuals may be able to optimize plumage coloration by molting in favourable areas characterized by high resource abundance. We used stable isotope analysis (δ¹³C, δ¹⁵N) to ask whether the diet and molt habitat/location of Bullock's orioles influenced their expression of carotenoid‐based plumage coloration as well as plumage carotenoid content and composition. Bullock's orioles with lower feather δ¹⁵N values acquired more colorful plumage (orange‐shifted hue) but had feathers with lower total carotenoid concentration, lower zeaxanthin concentration, and marginally lower canthaxanthin and lutein concentration. Examining factors occurring throughout the annual cycle are critical for understanding evolutionary and ecological processes. Here, we demonstrate that conditions experienced during a stopover molt, occurring hundreds to thousands of kilometers from the breeding grounds, influence the production of ornamental plumage coloration, which may carryover to influence inter‐ and intrasexual signaling in subsequent seasons.     

Effects of carotenoid availability during laying on reproduction in the blue tit

Carotenoids are antioxidant pigments involved in several physiological processes and signalling in animals that cannot synthesise them and therefore must acquire them from food. We experimentally investigated the effects of carotenoid availability in the diet during egg laying on antioxidant deposition in egg yolk and the related effects on nestling condition, female body condition and parental investment in the blue tit (Parus caeruleus). Carotenoid supplementation of egg-laying females resulted in a significant increase in carotenoid concentration in egg yolk, but not in vitamin E or A concentration. There was no relationship between yellow plumage colour of adult females and carotenoid deposition in eggs, and no differential effect of feeding treatment depending on female colour. Nestlings from eggs laid by carotenoid supplemented females had longer tarsi, had faster development of the immune system as reflected by leukocyte concentration in blood, and grew brighter yellow feathers than nestlings from control females. However, nestlings from the two groups did not differ significantly in body mass, plasma antioxidants or plumage colour hue. At the time of chick rearing, carotenoid-fed females had increased plasma vitamin E levels compared to controls. However, females from the two treatment groups did not differ significantly in body condition or feeding rate. These results suggest that carotenoid availability is limiting during egg laying, and that females may have to balance the benefits of investing in egg quality against the potential costs of impairing their own future antioxidant protection. In addition, there may be considerable variation in carotenoid availability not only across seasons, but also among different stages of the breeding season.     

Reconstructing plumage evolution in orioles (Icterus): repeated convergence and reversal in patterns

Several empirical studies suggest that sexually selected characters, including bird plumage, may evolve rapidly and show high levels of convergence and other forms of homoplasy. However, the processes that might generate such convergence have not been explored theoretically. Furthermore, no studies have rigorously addressed this issue using a robust phylogeny and a large number of signal characters. We scored the appearance of 44 adult male plumage characters that varied across New World orioles (Icterus). We mapped the plumage characters onto a molecular phylogeny based on two mitochondrial genes. Reconstructing the evolution of these characters revealed evidence of convergence or reversal in 42 of the 44 plumage characters. No plumage character states are restricted to any groups of species higher than superspecies in the oriole phylogeny. The high frequency of convergence and reversal is reflected in the low overall retention index (RI = 0.66) and the low overall consistency index (CI = 0.28). We found similar results when we mapped plumage changes onto a total evidence tree. Our findings reveal that plumage patterns and colors are highly labile between species of orioles, but highly conserved within the oriole genus. Furthermore, there are at least two overall plumage types that have convergently evolved repeatedly in the three oriole clades. This overall convergence leads to significant conflict between the molecular and plumage data. It is not clear what evolutionary processes lead to this homoplasy in individual characters or convergence in overall pattern. However, evolutionary constraints such as developmental limitations and genetic correlations between characters are likely to play a role. Our results are consistent with the belief that avian plumage and other sexually selected characters may evolve rapidly and may exhibit high homoplasy. The overall convergence in oriole plumage patterns is an interesting evolutionary phenomenon, but it cautions against heavy reliance on plumage characters for constructing phylogenies.     

Evolutionary drivers of seasonal plumage colours: colour change by moult correlates with sexual selection,predation risk and seasonality across passerines

Some birds undergo seasonal colour change by moulting twice each year, typically alternating between a cryptic, non‐breeding plumage and a conspicuous, breeding plumage (‘seasonal plumage colours’). We test for potential drivers of the evolution of seasonal plumage colours in all passerines (N = 5901 species, c. 60% of all birds). Seasonal plumage colours are uncommon, having appeared on multiple occasions but more frequently lost during evolution. The trait is more common in small, ground‐foraging species with polygynous mating systems, no paternal care and strong sexual dichromatism, suggesting it evolved under strong sexual selection and high predation risk. Seasonal plumage colours are also more common in species predicted to have seasonal breeding schedules, such as migratory birds and those living in seasonal climates. We propose that seasonal plumage colours have evolved to resolve a trade‐off between the effects of natural and sexual selection on colouration, especially in seasonal environments.     

Selection based on the size of the black tie of the great tit may be reversed in urban habitats

A standard approach to model how selection shapes phenotypic traits is the analysis of capture–recapture data relating trait variation to survival. Divergent selection, however, has never been analyzed by the capture–recapture approach. Most reported examples of differences between urban and nonurban animals reflect behavioral plasticity rather than divergent selection. The aim of this paper was to use a capture–recapture approach to test the hypothesis that divergent selection can also drive local adaptation in urban habitats. We focused on the size of the black breast stripe (i.e., tie width) of the great tit (Parus major), a sexual ornament used in mate choice. Urban great tits display smaller tie sizes than forest birds. Because tie size is mostly genetically determined, it could potentially respond to selection. We analyzed capture/recapture data of male great tits in Barcelona city (N = 171) and in a nearby (7 km) forest (N = 324) from 1992 to 2008 using MARK. When modelling recapture rate, we found it to be strongly influenced by tie width, so that both for urban and forest habitats, birds with smaller ties were more trap‐shy and more cautious than their larger tied counterparts. When modelling survival, we found that survival prospects in forest great tits increased the larger their tie width (i.e., directional positive selection), but the reverse was found for urban birds, with individuals displaying smaller ties showing higher survival (i.e., directional negative selection). As melanin‐based tie size seems to be related to personality, and both are heritable, results may be explained by cautious personalities being favored in urban environments. More importantly, our results show that divergent selection can be an important mechanism in local adaptation to urban habitats and that capture–recapture is a powerful tool to test it.     

RECONSTRUCTING THE EVOLUTION OF SEXUAL DICHROMATISM: CURRENT COLOR DIVERSITY DOES NOT REFLECT PAST RATES OF MALE AND FEMALE CHANGE

Males of sexually dimorphic species often appear more divergent among taxa than do females, so it is often assumed that evolutionary changes have occurred primarily in males. Yet, sexual dimorphisms can result from historical changes in either or both of the sexes, and few previous studies have investigated such patterns using phylogenetic methods. Here, we describe the evolution of male and female plumage colors in the grackles and allies (Icteridae), a songbird clade with a broad range in levels of sexual dichromatism. Using a model of avian perceptual color space, we calculated color distances within and among taxa on a molecular phylogeny. Our results show that female plumage colors have changed more dramatically than male colors in the evolutionary past, yet male colors are significantly more divergent among species today. Historical increases in dichromatism have involved changes in both sexes, whereas decreases in dichromatism have nearly always involved females evolving rapidly to look like males. Dichromatism is also associated with mating system in this group, with monogamous taxa tending to exhibit relatively low levels of sexual dichromatism. Our findings suggest that, despite appearances, female plumage evolution plays a more prominent role in sexual dichromatism than is generally assumed.