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Animal coloration is one of the most conspicuous aspects of human-perceived organismal diversity, yet also one of the least understood. In particular, explaining why species have specific colors (e.g., blue vs. red) has proven elusive. Here, we quantify for nearly all bird species, the proportion of the body covered by each of 12 human-visible color categories, and test whether existing theory can predict the direction of color evolution. The most common colors are black, white, gray and brown, while the rarest are green, blue, purple, and red. Males have more blue, purple, red, or black, whereas females have more yellow, brown, or gray. Sexual dichromatism is partly due to sexual selection favoring ornamental colors in males but not in females. However, sexual selection also correlated positively with brown in both sexes. Strong social selection favors red and black, colors used in agonistic signaling, with the strongest effects in females. Reduced predation risk selects against cryptic colors (e.g., brown) and favors specific ornamental colors (e.g., black). Nocturnality is mainly associated with brown. The effects of habitat use support the sensory drive theory for camouflage and signaling. Darker colors are more common in species living in wet and cold climates, matching ecogeographical rules. Our study unambiguously supports existing theories of color evolution across an entire class of vertebrates, but much variation remains unexplained.

Migratory birds undertake long and challenging journeys that have selected for a suite of adaptations from sensory mechanisms that facilitate orientation to extreme feats of endurance that push physiological limits. Recent work on two distantly related species revealed that migrating individuals increase their flight altitude dramatically during the day compared to at night 1 , 2 . These studies suggested that the phenomenon is driven by thermoregulation: the ascent to cooler heights during the day may offset heat generated by absorption of solar radiation. If thermoregulation is an important selective force on migratory species, migrants should have evolved lighter, more reflective plumage to avoid overheating. Here we show, across the entire avian radiation, that migratory species are indeed lighter coloured., (Copyright © 2021 Elsevier Inc. All rights reserved.)

The integration and synthesis of the data in different areas of science is drastically slowed and hindered by a lack of standards and networking programmes. Long-term studies of individually marked animals are not an exception. These studies are especially important as instrumental for understanding evolutionary and ecological processes in the wild. Furthermore, their number and global distribution provides a unique opportunity to assess the generality of patterns and to address broad-scale global issues (e.g. climate change). To solve data integration issues and enable a new scale of ecological and evolutionary research based on long-term studies of birds, we have created the SPI-Birds Network and Database (www.spibirds.org)-a large-scale initiative that connects data from, and researchers working on, studies of wild populations of individually recognizable (usually ringed) birds. Within year and a half since the establishment, SPI-Birds has recruited over 120 members, and currently hosts data on almost 1.5 million individual birds collected in 80 populations over 2,000 cumulative years, and counting. SPI-Birds acts as a data hub and a catalogue of studied populations. It prevents data loss, secures easy data finding, use and integration and thus facilitates collaboration and synthesis. We provide community-derived data and meta-data standards and improve data integrity guided by the principles of Findable, Accessible, Interoperable and Reusable (FAIR), and aligned with the existing metadata languages (e.g. ecological meta-data language). The encouraging community involvement stems from SPI-Bird's decentralized approach: research groups retain full control over data use and their way of data management, while SPI-Birds creates tailored pipelines to convert each unique data format into a standard format. We outline the lessons learned, so that other communities (e.g. those working on other taxa) can adapt our successful model. Creating community-specific hubs (such as ours, COMADRE for animal demography, etc.) will aid much-needed large-scale ecological data integration., (© 2020 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Heterozygosity-fitness correlations (HFCs) have been used to monitor the effects of inbreeding in threatened populations. HFCs can also be useful to investigate the potential effects of inbreeding in isolated relict populations of long-term persistence and to better understand the role of inbreeding and outbreeding as drivers of changes in genetic diversity. We studied a continental island population of thorn-tailed rayadito (Aphrastura spinicauda) inhabiting the relict forest of Fray Jorge National Park, north-central Chile. This population has experienced a long-term, gradual process of isolation since the end of the Tertiary. Using 10 years of field data in combination with molecular techniques, we tested for HFCs to assess the importance of inbreeding depression. If inbreeding depression is important, we predict a positive relationship between individual heterozygosity and fitness-related traits. We genotyped 183 individuals at 12 polymorphic microsatellite loci and used 7 measures of reproductive success and estimates of apparent survival to calculate HFCs. We found weak to moderate statistical support (P-values between 0.05 and 0.01) for a linear effect of female multi-locus heterozygosity (MLH) on clutch size and nonlinear effects on laying date and fledging success. While more heterozygous females laid smaller clutches, nonlinear effects indicated that females with intermediate values of MLH started laying earlier and had higher fledging success. We found no evidence for effects of MLH on annual fecundity or on apparent survival. Our results along with the long-term demographic stability of the study population contradict the hypothesis that inbreeding depression occurs in this population., (© The American Genetic Association. 2020. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Temporal variation in natural selection is predicted to strongly impact the evolution and demography of natural populations, with consequences for the rate of adaptation, evolution of plasticity, and extinction risk. Most of the theory underlying these predictions assumes a moving optimum phenotype, with predictions expressed in terms of the temporal variance and autocorrelation of this optimum. However, empirical studies seldom estimate patterns of fluctuations of an optimum phenotype, precluding further progress in connecting theory with observations. To bridge this gap, we assess the evidence for temporal variation in selection on breeding date by modeling a fitness function with a fluctuating optimum, across 39 populations of 21 wild animals, one of the largest compilations of long-term datasets with individual measurements of trait and fitness components. We find compelling evidence for fluctuations in the fitness function, causing temporal variation in the magnitude, but not the direction of selection. However, fluctuations of the optimum phenotype need not directly translate into variation in selection gradients, because their impact can be buffered by partial tracking of the optimum by the mean phenotype. Analyzing individuals that reproduce in consecutive years, we find that plastic changes track movements of the optimum phenotype across years, especially in bird species, reducing temporal variation in directional selection. This suggests that phenological plasticity has evolved to cope with fluctuations in the optimum, despite their currently modest contribution to variation in selection., Competing Interests: The authors declare no competing interest.

Variation in extra-pair paternity (EPP) among individuals of the same population could result from stochastic demography or from individual differences in mating strategies. Although the adaptive value of EPP has been widely studied, much less is known about the characteristics of the social environment that drive the observed patterns of EPP. Here, we demonstrate how concepts and well-developed tools for the study of social behaviour (such as social network analysis) can enhance the study of extra-pair mating decisions (focussing in particular on avian mating systems). We present several hypotheses that describe how characteristics of the social environment in which individuals are embedded might influence the levels of EPP in a socially monogamous population. We use a multi-level social approach (Hinde, 1976) to achieve a detailed description of the social structure and social dynamics of individuals in a group. We propose that the pair-bond, the direct (local) social environment and the indirect (extended) social environment, can contribute in different ways to the variation observed in the patterns of EPP, at both the individual and the population level. A strength of this approach is that it integrates into the analysis (indirect) interactions with all potential mates in a population, thus extending the current framework to study extra-pair mating behaviour. We also encourage the application of social network methods such as temporal dynamic analysis to depict temporal changes in the patterns of interactions among individuals in a group, and to study how this affects mating behaviour. We argue that this new framework will contribute to a better understanding of the proximate mechanisms that drive variation in EPP within populations in socially monogamous species, and might ultimately provide insights into the evolution and maintenance of mating systems., (© 2018 Cambridge Philosophical Society.)

Circadian clocks are centrally involved in the regulation of daily behavioural and physiological processes. These clocks are synchronized to the 24 h day by external cues (Zeitgeber), the most important of which is the light-dark cycle. In polar environments, however, the strength of the Zeitgeber is greatly reduced around the summer and winter solstices (continuous daylight or continuous darkness). How animals time their behaviour under such conditions has rarely been studied in the wild. Using a radio-telemetry-based system, we investigated daily activity rhythms under continuous daylight in Barrow, Alaska, throughout the breeding season in four bird species that differ in mating system and parental behaviour. We found substantial diversity in daily activity rhythms depending on species, sex and breeding stage. Individuals exhibited either robust, entrained 24 h activity cycles, were continuously active (arrhythmic) or showed 'free-running' activity cycles. In semipalmated sandpipers, a shorebird with biparental incubation, we show that the free-running rhythm is synchronized between pair mates. The diversity of diel time-keeping under continuous daylight emphasizes the plasticity of the circadian system, and the importance of the social and life-history context. Our results support the idea that circadian behaviour can be adaptively modified to enable species-specific time-keeping under polar conditions.

In many species that form socially monogamous pair bonds, a considerable proportion of the offspring is sired by extrapair males. This observation has remained a puzzle for evolutionary biologists: although mating outside the pair bond can obviously increase the offspring production of males, the benefits of such behavior to females are less clear, yet females are known to actively solicit extrapair copulations. For more than two decades adaptionist explanations have dominated the discussions, yet remain controversial, and genetic constraint arguments have been dismissed without much consideration. An intriguing but still untested hypothesis states that extrapair mating behavior by females may be affected by the same genetic variants (alleles) as extrapair mating behavior by males, such that the female behavior could evolve through indirect selection on the male behavior. Here we show that in the socially monogamous zebra finch, individual differences in extrapair mating behavior have a hereditary component. Intriguingly, this genetic basis is shared between the sexes, as shown by a strong genetic correlation between male and female measurements of extrapair mating behavior. Hence, positive selection on males to sire extrapair young will lead to increased extrapair mating by females as a correlated evolutionary response. This behavior leads to a fundamentally different view of female extrapair mating: it may exist even if females obtain no net benefit from it, simply because the corresponding alleles were positively selected in the male ancestors.

It has been suggested that positive selection, in particular selection favoring a change in the protein sequence, plays a role in the evolution of olfactory receptor (OR) gene repertoires in fish and mammals. ORs are 7-transmembrane domain (TM) proteins, members of the G-protein-coupled receptor superfamily in vertebrate genomes, and responsible for odorant binding and discrimination. OR gene repertoires in birds are surprisingly large and diverse, suggesting that birds have a keen olfactory sense. The aim of this study is to investigate signatures of positive selection in an expanded OR clade (group-gamma-c) that seems to be a characteristic of avian genomes. Using maximum-likelihood methods that estimate the d(N)/d(S) ratios and account for the effects of recombination, we show here that there is evidence for positive selection in group-gamma-c partial OR coding sequences of 9 bird species that are likely to have different olfactory abilities: the blue tit (Cyanistes caeruleus), the black coucal (Centropus grillii), the brown kiwi (Apteryx australis), the canary (Serinus canaria), the galah (Eolophus roseicapillus), the kakapo (Strigops habroptilus), the mallard (Anas platyrhynchos), the red jungle fowl (Gallus gallus), and the snow petrel (Pagodroma nivea). Positively selected codons were predominantly located in TMs, which in other vertebrates are involved in odorant binding. Our data suggest that 1) at least some avian OR genes have been subjected to adaptive evolution, 2) the extent of such adaptive evolution differs between bird species, and 3) positive selective pressures may have been stronger on the group-gamma-c OR genes of species that have well-developed olfactory abilities.

Among vertebrates, the sense of smell is mediated by olfactory receptors (ORs) expressed in sensory neurons within the olfactory epithelium. Comparative genomic studies suggest that the olfactory acuity of mammalian species correlates positively with both the total number and the proportion of functional OR genes encoded in their genomes. In contrast to mammals, avian olfaction is poorly understood, with birds widely regarded as relying primarily on visual and auditory inputs. Here, we show that in nine bird species from seven orders (blue tit, Cyanistes caeruleus; black coucal, Centropus grillii; brown kiwi, Apteryx australis; canary, Serinus canaria; galah, Eolophus roseicapillus; red jungle fowl, Gallus gallus; kakapo, Strigops habroptilus; mallard, Anas platyrhynchos; snow petrel, Pagodroma nivea), the majority of amplified OR sequences are predicted to be from potentially functional genes. This finding is somewhat surprising as one previous report suggested that the majority of OR genes in an avian (red jungle fowl) genomic sequence are non-functional pseudogenes. We also show that it is not the estimated proportion of potentially functional OR genes, but rather the estimated total number of OR genes that correlates positively with relative olfactory bulb size, an anatomical correlate of olfactory capability. We further demonstrate that all the nine bird genomes examined encode OR genes belonging to a large gene clade, termed gamma-c, the expansion of which appears to be a shared characteristic of class Aves. In summary, our findings suggest that olfaction in birds may be a more important sense than generally believed.

Ultraviolet (UV) reflectance of the plumage is common in birds and plays an important role in sexual signalling. Recently, it has been proposed that birds are able to modify plumage UV reflectance by the application of uropygial gland secretion. Based on a survey of the optical properties of this secretion from 51 species belonging to 12 avian orders, we show that two main types of uropygial secretions exist, one predominantly found in passerines and one in non-passerines, both reducing relative UV reflectance of a white background (Teflon tape). We quantified how each type of secretion (exemplified by blue tit and mallard) affected feather UV reflectance. Both secretions reduced overall brightness and relative UV reflectance of white mallard feathers but hardly affected the reflectance of UV/blue blue tit crown feathers. According to models of avian colour vision, changes in reflectance due to application of the secretion were at or below the discrimination threshold of most birds. We conclude that the uropygial secretion is unlikely to play a major role in modifying plumage UV reflectance. However, the optical properties of the uropygial secretion may have been selected to interfere as little as possible with visual signaling through plumage reflectance.

The steroid hormone testosterone (T) plays a central role in the regulation of breeding in males, because many physiological, morphological and behavioural traits related to reproduction are T dependent. Moreover, in many seasonally breeding vertebrates, male plasma T levels typically show a pronounced peak during the breeding season. While such population-level patterns are fairly well worked out, the sources and the implications of the large variability in individual T levels within the seasonal cycle remain surprisingly little understood. Understanding the potential sources of individual variation in T levels is important for behavioural and evolutionary ecologists, for at least two reasons. First, in 'honest signalling' theory, T is hypothesized to play a critical role as the assumed factor that enforces honesty of the expression of sexually selected quality indicators. Second, T is often considered a key mediator of central life-history trade-offs, such as investment in survival versus reproduction or in mating versus parental care. Here, we discuss the patterns of within- and between-individual variation in male plasma T levels in free-living populations of birds. We argue that it is unclear whether this variability mainly reflects differences in underlying individual quality (intrinsic factors such as genetic or maternal effects) or in the environment (extrinsic factors including time of day, individual territorial status and past experience). Research in avian behavioural endocrinology has mainly focused on the effects of extrinsic factors, while other sources of variance are often ignored. We suggest that studies that use an integrative approach and investigate the relative importance of all potential sources of variation are essential for the interpretation of data on individual plasma T levels.

Colorful plumages are conspicuous social signals in birds, and the expression of these colors often reflects the quality of their bearers. Since mature feathers are dead structures, plumage color is often considered a static signal that does not change after molt. Feathers, however, can and do deteriorate between molts, and birds need to invest heavily in plumage maintenance. Here we argue that this need for preserving plumage condition and hence signaling content might have given rise to a novel type of sexual signal: cosmetic coloration. Cosmetic coloration occurs when the substances used for plumage maintenance change the color of the feathers, thereby becoming a signal themselves. Our review of cosmetic coloration in birds demonstrates that it is more widespread than currently realized, occurring in at least 13 bird families. Cosmetics have varied origins: they can be produced by the bird itself (uropygial and skin secretions, feather powder) or obtained from the environment (soil, iron oxide). Intraspecific patterns of cosmetic use (sex, age, and seasonal dimorphism) suggest that in many cases it may act as a sexual signal. However, more information is required on function, mechanisms, and costs to understand the evolution of cosmetic coloration and to confirm its signaling role.

Steroid hormones are predicted to vary with mating systems, degree of paternal care and aggression, as proposed in the "challenge hypothesis." We measured plasma concentrations of testosterone (T) and 5alpha-dihydrotestosterone (DHT) in two high Arctic breeding and closely related shorebird species, the polygynous pectoral sandpiper (Calidris melanotos) and the monogamous semipalmated sandpiper (Calidris pusilla) to examine whether the hormonal findings corroborate the predictions of the challenge hypothesis. In both species, males showed significantly higher levels of DHT and T than females, but in pectoral sandpipers median T levels were 34 times greater in males than in females, whereas in semipalmated sandpiper there was only a 4.9-fold difference. T and DHT concentrations correlated in semipalmated sandpipers and in male, but not in female, pectoral sandpipers. In semipalmated sandpipers, androgen levels were highest in the beginning of the breeding season and then declined, whereas male pectoral sandpipers showed extremely high androgen levels which were sustained throughout the breeding season. In both species, androgen levels were independent of body condition. Several incubating male semipalmated sandpipers had high circulating T levels, suggesting that low T is not required to exhibit paternal care. Our results are consistent with the challenge hypothesis in that androgen concentrations were higher throughout the breeding season in the polygynous compared to the monogamous species. Our study also supports previous studies suggesting that the short breeding season in the high Arctic and the importance of male care to nest success may be factors leading to behavioral T insensitivity.

Matings between close relatives often reduce the fitness of offspring, probably because homozygosity leads to the expression of recessive deleterious alleles. Studies of several animals have shown that reproductive success is lower when genetic similarity between parents is high, and that survival and other measures of fitness increase with individual levels of genetic diversity. These studies indicate that natural selection may favour the avoidance of matings with genetically similar individuals. But constraints on social mate choice, such as a lack of alternatives, can lead to pairing with genetically similar mates. In such cases, it has been suggested that females may seek extra-pair copulations with less related males, but the evidence is weak or lacking. Here we report a strong positive relationship between the genetic similarity of social pair members and the occurrence of extra-pair paternity and maternity ('quasi-parasitism') in three species of shorebirds. We propose that extra-pair parentage may represent adaptive behavioural strategies to avoid the negative effects of pairing with a genetically similar mate.

Under many circumstances, it might be adaptive for parents to bias the investment in offspring in relation to sex. Recently developed molecular techniques that allow sex determination of newly hatched offspring have caused a surge in studies of avian sex allocation. Whether females bias the primary brood sex ratio in relation to factors such as environmental and parental quality is debated. Progress is hampered because the mechanisms for primary sex ratio manipulation are unknown. Moreover, publication bias against non-significant results may distort our view of adaptive sex ratio manipulation. Despite this, there is recent experimental evidence for adaptive brood sex ratio manipulation in birds. Parental care is a particularly likely candidate to affect the brood sex ratio because it can have strong direct effects on the fitness of both parents and their offspring. We investigate and make predictions of factors that can be important for adaptive brood sex ratio manipulation under different patterns of parental care. We encourage correlational studies based on sufficiently large datasets to ensure high statistical power, studies identifying and experimentally altering factors with sex-differential fitness effects that may cause brood sex ratio skew, and studies that experimentally manipulate brood sex ratio and investigate fitness effects.

Undeveloped eggs occur frequently in birds and are often considered infertile, and discarded. However, the majority of undeveloped eggs may in fact have been fertilised and embryos might have died at an early stage. Such eggs contain valuable information, for example about offspring sex and paternity, and level of inbreeding. Obtaining such information may also give insight into the patterns and causes of early embryo mortality. Here we describe a simple technique for removing embryo cells from the blastoderm to obtain DNA to genotype the offspring and unequivocally ascertain fertilisation status, while retaining the overlying perivitelline layer (PVL) for sperm counts over the entire membrane. We tested this method on freshly collected eggs (high‐quality material), as well as on eggs from abandoned clutches and unhatched eggs (potentially deteriorated material) of blue tits (Cyanistes caeruleus). We sampled a total of 707 eggs from a wild population of blue tits, extracted DNA from the eggs' blastoderm using a Qiagen kit, and genotyped the samples with 14 polymorphic microsatellite markers, plus one sexing marker. Overall, we successfully genotyped 97% of all eggs. Our study is the most extensive dataset of genotyped undeveloped eggs to date and demonstrates that one can reliably genotype undeveloped fertile eggs as well as retain the PVL for observations of sperm and embryo cells. [ABSTRACT FROM AUTHOR]

The germline-restricted chromosome (GRC) is likely present in all songbird species but differs widely in size and gene content. This extra chromosome has been described as either a microchromosome with only limited basic gene content or a macrochromosome with enriched gene functions related to female gonad and embryo development. Here, we assembled, annotated, and characterized the first micro-GRC in the blue tit (Cyanistes caeruleus) using high-fidelity long-read sequencing data. Although some genes on the blue tit GRC show signals of pseudogenization, others potentially have important functions, either currently or in the past. We highlight the GRC gene paralog BMP15 , which is among the highest expressed GRC genes both in blue tits and in zebra finches (Taeniopygia guttata) and is known to play a role in oocyte and follicular maturation in other vertebrates. The GRC genes of the blue tit are further enriched for functions related to the synaptonemal complex. We found a similar functional enrichment when analyzing published data on GRC genes from two nightingale species (Luscinia spp.). We hypothesize that these genes play a role in maintaining standard maternal inheritance or in recombining maternal and paternal GRCs during potential episodes of biparental inheritance. [ABSTRACT FROM AUTHOR]

To link to full-text access for this article, visit this link: http://dx.doi.org/10.1016/j.anbehav.2009.01.012 Byline: Alain Jacot, Mihai Valcu, Kees van Oers, Bart Kempenaers Abstract: In resource defence mating systems, males monopolize a resource that is of primary importance for breeding females. For secondary cavity nesters, the availability of suitable nesting sites is important in determining the strength of intrasexual competition, whereby phenotypic and behavioural traits will be favoured that enable individuals to gain access to these sites. The traits that are important in male competition may additionally affect mate choice decisions and a female's investment in the current brood. In a field study on blue tits, Cyanistes caeruleus, we increased intrasexual competition by experimentally limiting nest sites in experimental plots and compared these plots to control plots. Birds breeding in experimental plots did not differ phenotypically from birds in control plots. However, females that bred in the nest site-limited plots fed their offspring at a higher rate than control females. This result indicates that increased competition for limited resources led to more investment in current reproduction, either because successful females were of higher intrinsic quality or because they adjusted their investment in relation to superior territory or male characteristics. Author Affiliation: Max Planck Institute for Ornithology, Department of Behavioural Ecology and Evolutionary Genetics, Germany Article History: Received 18 March 2008; Revised 30 May 2008; Accepted 16 January 2009 Article Note: (miscellaneous) MS. number: 08-00191R

The evolutionary consequences of individual genetic diversity are frequently studied by assessing heterozygosity–fitness correlations (HFCs). The prevalence of positive and negative HFCs and the predominance of general versus local effects in wild populations are far from understood, partly because comprehensive studies testing for both inbreeding and outbreeding depression are lacking. We studied a genetically diverse population of blue tits in southern Germany using a genome‐wide set of 87 microsatellites to investigate the relationship between proxies of reproductive success and measures of multilocus and single‐locus individual heterozygosity (MLH and SLH). We used complimentary measures of MLH and partitioned markers into functional categories according to their position in the blue tit genome. HFCs based on MLH were consistently negative for functional loci, whereas correlations were rather inconsistent for loci found in nonfunctional areas of the genome. Clutch size was the only reproductive variable showing a general effect. We found evidence for local effects for three measures of reproductive success: arrival date at the breeding site, the probability of breeding at the study site and male reproductive success. For these, we observed consistent, and relatively strong, negative effects at one functional locus. Remarkably, this marker had a similar effect in another blue tit population from Austria (~400 km to the east). We suggest that a genetic local effect on timing of arrival might be responsible for most negative HFCs detected, with carry‐over effects on other reproductive traits. This effect could reflect individual differences in the distance between overwintering areas and breeding sites. [ABSTRACT FROM AUTHOR]

Studies on the relationship between behavioral traits and dispersal are necessary to understand the evolution of dispersal syndromes. Empirical studies have mainly focused on natal dispersal, even though behavioral differences between dispersers and philopatric individuals are suspected to hold through the whole life cycle, potentially affecting breeding dispersal propensity. Using capture–mark–recapture data and behavioral trials in a forest passerine, the thorn‐tailed rayadito Aphrastura spinicauda, we describe inter‐individual differences in exploratory behavior and aggressiveness, and investigate the relationship between those traits and breeding dispersal. Our study took place in Fray Jorge National Park, north‐central Chile, where a relatively isolated population of rayaditos inhabits a naturally fragmented environment. We found that scores for behavioral traits were consistent between years. Exploratory behavior was similar between sexes, while males showed higher levels of aggression towards a conspecific male intruder. Only exploratory behavior was related to breeding dispersal propensity, with fast‐exploring rayaditos being more likely to have dispersed between seasons. This finding provides indirect evidence for the existence of a dispersal strategy that could reduce dispersal costs in the fragmented landscape of Fray Jorge. To our knowledge, this is the first study documenting an association between breeding dispersal and exploratory behavior in a wild bird population. A longitudinal individual‐based study will help determining whether this association constitutes a behavioral syndrome. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

It is often claimed that pair bonds preferentially form between individuals that resemble one another. Such assortative mating appears to be widespread throughout the animal kingdom. Yet it is unclear whether the apparent ubiquity of assortative mating arises primarily from mate choice (“like attracts like”), which can be constrained by same-sex competition for mates; from spatial or temporal separation; or from observer, reporting, publication, or search bias. Here, based on a conventional literature search, we find compelling meta-analytical evidence for size-assortative mating in birds (r = 0.178, 95% CI 0.142–0.215, 83 species, 35,591 pairs). However, our analyses reveal that this effect vanishes gradually with increased control of confounding factors. Specifically, the effect size decreased by 42% when we used previously unpublished data from nine long-term field studies, i.e., data free of reporting and publication bias (r = 0.103, 95% CI 0.074–0.132, eight species, 16,611 pairs). Moreover, in those data, assortative mating effectively disappeared when both partners were measured by independent observers or separately in space and time (mean r = 0.018, 95% CI −0.016–0.057). Likewise, we also found no evidence for assortative mating in a direct experimental test for mutual mate choice in captive populations of Zebra finches (r = −0.020, 95% CI −0.148–0.107, 1,414 pairs). These results highlight the importance of unpublished data in generating unbiased meta-analytical conclusions and suggest that the apparent ubiquity of assortative mating reported in the literature is overestimated and may not be driven by mate choice or mating competition for preferred mates. [ABSTRACT FROM AUTHOR]

Most bird species exhibit biparental care, but the type of care provided by each sex may differ substantially. In particular, during the incubation phase in passerines, females perform most or all of the incubation, while the male cares for the brood indirectly by feeding the female. However, detailed descriptions of this male investment during the incubation period are missing. Here, we quantitatively describe female nest attendance and male incubation feeding throughout the ~ 14‐day incubation period in a population of Eurasian Blue Tits Cyanistes caeruleus breeding in nestboxes. Males and females progressively increased their daily activity at the nest over the incubation period. The amount of day‐time incubation, measured as the proportion of the active day (time interval between first nestbox exit in the morning and last entry in the evening) a female spent inside the nestbox, varied between 52 and 60% with an average of 55% per day. The frequency of male incubation feeding varied between 0 and 74 times per day with an average of 12 feeds per day. Both male feeding rate and female nest attendance were highest in the morning and declined rapidly throughout the day. Females were more likely to be off the nest during the warmest periods (15–21 °C), as expected based on thermal needs of the developing embryos, but also during the coldest periods (2–5 °C), presumably due to the energetic needs of the female. This was despite the fact that males fed their females more often at the nest when ambient temperatures were low. Females that received more feeds incubated more and their off‐nest bouts were shorter after a feed. The observed variation in female incubation and in male feeding rate was not linked to individual age or to variation in measures of reproductive success. However, direct observations showed that in some pairs a substantial amount of feeding by males occurred outside the nestbox. This suggests that the true male investment might have been underestimated, here and in previous studies. [ABSTRACT FROM AUTHOR]

Abstract: A central goal in evolutionary ecology is to identify factors that explain variation in reproductive success, i.e. in the number of offspring produced. In altricial birds, a substantial part of this variation is determined by the number of nestlings that die before fledging, but surprisingly little is known about the proximate causes of offspring mortality during the nestling period. We used a uniquely comprehensive dataset of parental nestbox visits from seven breeding seasons to investigate the association between parental behaviour and nestling mortality in a population of blue tits (Cyanistes caeruleus). In almost all nests that suffered complete brood mortality one of the parents had suddenly disappeared during the nestling stage. In contrast, parental disappearance in nests with partial brood mortality was rare and equally common as in nests with no brood loss. With few exceptions, parents that disappeared during the nestling stage were never observed again and never returned to breed. In contrast, parents that remained after their partner disappeared were equally likely to be observed again or return to breed as parents of nests where both parents stayed. Visit rates at nests where a parent would disappear did not differ from those at nests where both parents stayed. Taken together, our results show that ‐ in contrast to partial brood failure ‐ complete brood failure is almost always associated with the sudden and permanent disappearance of one of the parents, probably due to predation. Partial and complete brood mortality should be treated as distinct processes that have different underlying causes. [ABSTRACT FROM AUTHOR]

Colorful plumage ornaments may evolve because they play a role in mate choice or in intrasexual competition, acting as signals of species identity or of individual quality. The zebra finch (Taeniopygia guttata) is a model organism for the study of mate choice and its colorful plumage ornaments are thought to be used in both of these contexts. Numerous genetic color variants have been described for this species, but they are rare in the wild. This raises the question whether discrimination against deviant phenotypes maintains the species' uniform plumage color (rare-mate disadvantage). Furthermore, comparison to closely related species suggests that the lack of colorful ornaments in female zebra finches is a derived condition. Male preferences for less-ornamented females may have led to sexual dichromatism in the zebra finch. Here, we test the role of plumage ornaments experimentally by altering male and female coloration to mimic 2 types of naturally occurring genetic color variants. We estimated effects on social dominance and reproductive success in large breeding aviaries in one domesticated and 2 recently wild-derived populations. Hypotheses, methods, and analyses were preregistered to ensure maximal objectivity of the results presented. Despite a fairly drastic manipulation and a powerful experimental design, we found no effect of the treatment on social dominance or on reproductive success. Our results suggest that mate choice is not the mechanism that maintains homogeneity of zebra finch plumage coloration, or that can explain the loss of ornaments in females. [ABSTRACT FROM AUTHOR]

Parents provisioning their offspring can adopt different tactics to meet increases in offspring demand. In this study, we experimentally manipulated brood demand in free living great tits (Parus major) via brood size manipulations and compared the tactics adopted by parents in 2 successive years (2010 and 2011) with very different ecological conditions. In 2011, temperatures were warmer, there were fewer days with precipitation, and caterpillars (the preferred prey of great tits) made up a significantly larger proportion of the diet. In this "good" year, parents responded to experimental increases in brood demand by decreasing mean inter-visit intervals (IVIs) and reducing prey selectivity, which produced equal average long-term delivery of food to nestlings across the brood size treatments. In 2010, there was no evidence for effects of brood size manipulations on mean IVIs or prey selectivity. Consequently, nestlings from enlarged broods experienced significantly lower long-term average delivery rates compared with nestlings from reduced broods. In this "bad" year, parents also exhibited changes in the variance in inter-visit intervals (IVIs) as a function of treatment that were consistent with variance-sensitive foraging theory: variance in IVIs tended to be lowest for reduced broods and highest for enlarged broods. Importantly, this pattern differed significantly from that observed in the "good" year. We therefore found some support for variancesensitive provisioning in the year with more challenging ecological conditions. Taken together, our results show that variation in brood demand can result in markedly different parental foraging tactics depending on ecological conditions. [ABSTRACT FROM AUTHOR]

The distribution of suitable habitat influences natal and breeding dispersal at small spatial scales, resulting in strong microgeographic genetic structure. Although environmental variation can promote interpopulation differences in dispersal behavior and local spatial patterns, the effects of distinct ecological conditions on within-species variation in dispersal strategies and in fine-scale genetic structure remain poorly understood. We studied local dispersal and fine-scale genetic structure in the thorn-tailed rayadito ( Aphrastura spinicauda), a South American bird that breeds along a wide latitudinal gradient. We combine capture-mark-recapture data from eight breeding seasons and molecular genetics to compare two peripheral populations with contrasting environments in Chile: Navarino Island, a continuous and low density habitat, and Fray Jorge National Park, a fragmented, densely populated and more stressful environment. Natal dispersal showed no sex bias in Navarino but was female-biased in the more dense population in Fray Jorge. In the latter, male movements were restricted, and some birds seemed to skip breeding in their first year, suggesting habitat saturation. Breeding dispersal was limited in both populations, with males being more philopatric than females. Spatial genetic autocorrelation analyzes using 13 polymorphic microsatellite loci confirmed the observed dispersal patterns: a fine-scale genetic structure was only detectable for males in Fray Jorge for distances up to 450 m. Furthermore, two-dimensional autocorrelation analyzes and estimates of genetic relatedness indicated that related males tended to be spatially clustered in this population. Our study shows evidence for context-dependent variation in natal dispersal and corresponding local genetic structure in peripheral populations of this bird. It seems likely that the costs of dispersal are higher in the fragmented and higher density environment in Fray Jorge, particularly for males. The observed differences in microgeographic genetic structure for rayaditos might reflect the genetic consequences of population-specific responses to contrasting environmental pressures near the range limits of its distribution. [ABSTRACT FROM AUTHOR]

In species with biparental care and lifetime monogamy, the fecundity of a male's partner can be a major component of his fitness but it is unclear whether males can assess female fecundity before breeding. We carried out an experiment in which we measured variation in female fecundity (repeatability 39%, 213 females) in a captive zebra finch population and tested whether males preferred unfamiliar females of high fecundity (approximately top 10% of the population; 30 eggs laid on average) over those of low fecundity (bottom 10%; 6 eggs). We first tested whether naïve human observers could identify the high-fecundity female when confronted with duos of high and low fecundity. Humans guessed correctly in 58% of the cases (95% confidence interval [CI] 50-66%) indicating that differences in female condition were not highly obvious to humans. Zebra finch males preferred the high-fecundity female in 59% of choice tests that lasted 20 min (CI 52-66%). When extending such choice tests over several days, male "success" in associating with the high-fecundity female was still modest (61% correct choices, CI 44-76%). Overall, male zebra finches seem to have only limited abilities to identify the better mate when faced with a choice between extremes in terms of female fecundity. We found no male preference for heavier females. We speculate that such a preference may not have evolved because, in contrast to many ectothermic species, predicting fecundity from female weight is not sufficiently accurate (r² = 0.04) for the benefits to outweigh the costs of increased male-male competition for heavy females. [ABSTRACT FROM AUTHOR]

Identifying causal genetic variants underlying heritable phenotypic variation is a long-standing goal in evolutionary genetics. We previously identified several quantitative trait loci ( QTL) for five morphological traits in a captive population of zebra finches ( Taeniopygia guttata) by whole-genome linkage mapping. We here follow up on these studies with the aim to narrow down on the quantitative trait variants ( QTN) in one wild and three captive populations. First, we performed an association study using 672 single nucleotide polymorphisms ( SNPs) within candidate genes located in the previously identified QTL regions in a sample of 939 wild-caught zebra finches. Then, we validated the most promising SNP-phenotype associations ( n = 25 SNPs) in 5228 birds from four populations. Genotype-phenotype associations were generally weak in the wild population, where linkage disequilibrium ( LD) spans only short genomic distances. In contrast, in captive populations, where LD blocks are large, apparent SNP effects on morphological traits (i.e. associations) were highly repeatable with independent data from the same population. Most of those SNPs also showed significant associations with the same trait in other captive populations, but the direction and magnitude of these effects varied among populations. This suggests that the tested SNPs are not the causal QTN but rather physically linked to them, and that LD between SNPs and causal variants differs between populations due to founder effects. While the identification of QTN remains challenging in nonmodel organisms, we illustrate that it is indeed possible to confirm the location and magnitude of QTL in a population with stable linkage between markers and causal variants. [ABSTRACT FROM AUTHOR]

Males of socially monogamous species can increase their siring success via within-pair and extra-pair fertilizations. In this study, we focused on the different sources of (co)variation between these siring routes, and asked how each contributes to total siring success. We quantified the fertilization routes to siring success, as well as behaviors that have been hypothesized to affect siring success, over a five-year period for a wild population of great tits Parus major. We considered siring success and its fertilization routes as 'interactive phenotypes' arising from phenotypic contributions of both members of the social pair. We show that siring success is strongly affected by the fecundity of the social (female) partner. We also demonstrate that a strong positive correlation between extra-pair fertilization success and paternity loss likely constrains the evolution of these two routes. Moreover, we show that more explorative and aggressive males had less extra-pair fertilizations, whereas more explorative females laid larger clutches. This study thus demonstrates that (co)variation in siring routes is caused by multiple factors not necessarily related to characteristics of males. We thereby highlight the importance of acknowledging the multilevel structure of male fertilization routes when studying the evolution of male mating strategies. [ABSTRACT FROM AUTHOR]

Decoding genomic sequences and determining their variation within populations has potential to reveal adaptive processes and unravel the genetic basis of ecologically relevant trait variation within a species. The blue tit Cyanistes caeruleus - a long-time ecological model species - has been used to investigate fitness consequences of variation in mating and reproductive behaviour. However, very little is known about the underlying genetic changes due to natural and sexual selection in the genome of this songbird. As a step to bridge this gap, we assembled the first draft genome of a single blue tit, mapped the transcriptome of five females and five males to this reference, identified genomewide variants and performed sex-differential expression analysis in the gonads, brain and other tissues. In the gonads, we found a high number of sex-biased genes, and of those, a similar proportion were sex-limited (genes only expressed in one sex) in males and females. However, in the brain, the proportion of female-limited genes within the female-biased gene category (82%) was substantially higher than the proportion of male-limited genes within the male-biased category (6%). This suggests a predominant on-off switching mechanism for the female-limited genes. In addition, most male-biased genes were located on the Z-chromosome, indicating incomplete dosage compensation for the male-biased genes. We called more than 500 000 SNPs from the RNA-seq data. Heterozygote detection in the single reference individual was highly congruent between DNA-seq and RNA-seq calling. Using information from these polymorphisms, we identified potential selection signals in the genome. We list candidate genes which can be used for further sequencing and detailed selection studies, including genes potentially related to meiotic drive evolution. A public genome browser of the blue tit with the described information is available at . [ABSTRACT FROM AUTHOR]

Classical sexual selection theory provides a well-supported conceptual framework for understanding the evolution and signalling function of male ornaments. It predicts that males obtain greater fitness benefits than females through multiple mating because sperm are cheaper to produce than eggs. Sexual selection should therefore lead to the evolution of male-biased secondary sexual characters. However, females of many species are also highly ornamented. The view that this is due to a correlated genetic response to selection on males was widely accepted as an explanation for female ornamentation for over 100 years and current theoretical and empirical evidence suggests that genetic constraints can limit sex-specific trait evolution. Alternatively, female ornamentation can be the outcome of direct selection for signalling needs. Since few studies have explored interspecific patterns of both male and female elaboration, our understanding of the evolution of animal ornamentation remains incomplete, especially over broad taxonomic scales. Here we use a new method to quantify plumage colour of all ~6,000 species of passerine birds to determine the main evolutionary drivers of ornamental colouration in both sexes. We found that conspecific male and female colour elaboration are strongly correlated, suggesting that evolutionary changes in one sex are constrained by changes in the other sex. Both sexes are more ornamented in larger species and in species living in tropical environments. Ornamentation in females (but not males) is increased in cooperative breeders-species in which female-female competition for reproductive opportunities and other resources related to breeding may be high. Finally, strong sexual selection on males has antagonistic effects, causing an increase in male colouration but a considerably more pronounced reduction in female ornamentation. Our results indicate that although there may be genetic constraints to sexually independent colour evolution, both female and male ornamentation are strongly and often differentially related to morphological, social and life-history variables. [ABSTRACT FROM AUTHOR]

Most studies on extra-pair paternity ( EPP) focus either on a specific male's extra-pair gains or his extra-pair losses. For an individual bird however, mate choice or mate availability may underlie strong spatial restrictions. Disregarding this spatial aspect may underestimate or mask effects of parameters influencing observed EPP patterns., Here, we propose a spatially explicit model for investigating the probability of having extra-pair offspring ( EPO) within local networks of breeding pairs. The data set includes all realized and unrealized potential extra-pair matings. This method is biologically meaningful because it allows (a) considering both members of an extra-pair mating and their social mates, and (b) direct modelling of the spatial context in which extra-pair behaviour occurs. The method has the advantage that it can provide inference about the relative contribution of spatial and non-spatial parameters, and about the relative importance of male and female neighbourhoods., We apply this method to parentage data from 1025 broods collected over 12 breeding seasons in two independent study populations of blue tits ( Cyanistes caeruleus). We investigate a set of predictions based on the EPP literature, namely that EPP depends on male age and body size, breeding density and breeding synchrony. In all analyses, we control for breeding distance, a parameter that is expected to influence EPP even under random mating., The results show that older and larger males were more likely to sire EPO, but both effects decreased with increasing breeding distance. Local breeding density but not synchrony predicted whether a particular male-female combination had EPO, at least in one of the study areas. Apart from breeding distance, male age had the strongest effect on EPP, followed by a measure of breeding density. The method thus allows a comprehensive assessment of the relative importance of different types of spatial and non-spatial parameters to explain variation in the occurrence of EPP, while controlling for the fact that individuals that breed further apart are less likely to have EPO., The proposed approach is not limited to investigate EPP, but can be applied to other behavioural interactions between two individuals, such as dominance, competition and (social) mating. [ABSTRACT FROM AUTHOR]

Indirect benefits of mate choice result from increased offspring genetic quality and may be important drivers of female behaviour. 'Good-genes-for-viability' models predict that females prefer mates of high additive genetic value, such that offspring survival should correlate with male attractiveness. Mate choice may also vary with genetic diversity (e.g. heterozygosity) or compatibility (e.g. relatedness), where the female's genotype influences choice. The relative importance of these nonexclusive hypotheses remains unclear. Leks offer an excellent opportunity to test their predictions, because lekking males provide no material benefits and choice is relatively unconstrained by social limitations. Using 12 years of data on lekking lance-tailed manakins, Chiroxiphia lanceolata, we tested whether offspring survival correlated with patterns of mate choice. Offspring recruitment weakly increased with father attractiveness (measured as reproductive success, RS), suggesting attractive males provide, if anything, only minor benefits via offspring viability. Both male RS and offspring survival until fledging increased with male heterozygosity. However, despite parent-offspring correlation in heterozygosity, offspring survival was unrelated to its own or maternal heterozygosity or to parental relatedness, suggesting survival was not enhanced by heterozygosity per se. Instead, offspring survival benefits may reflect inheritance of specific alleles or nongenetic effects. Although inbreeding depression in male RS should select for inbreeding avoidance, mates were not less related than expected under random mating. Although mate heterozygosity and relatedness were correlated, selection on mate choice for heterozygosity appeared stronger than that for relatedness and may be the primary mechanism maintaining genetic variation in this system despite directional sexual selection. [ABSTRACT FROM AUTHOR]