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Fitness is at the core of evolutionary theory, but it is difficult to measure accurately. One way to measure long-term fitness is by calculating the individual’s reproductive value, which represents the expected number of allele copies an individual passes on to distant future generations. However, this metric of fitness is scarcely used because the estimation of individual’s reproductive value requires long-term pedigree data, which is rarely available in wild populations where following individuals from birth to death is often impossible. Wild study systems therefore use short-term fitness metrics as proxies, such as the number of offspring produced. This study compared two frequently used short-term metrics for fitness obtained at different offspring life stages (eggs, hatchlings, fledglings and recruits), and compared their ability to predict reproductive values derived from the genetic pedigree of a wild passerine bird population. We used twenty years of precise field observations and a near-complete genetic pedigree to calculate reproductive success, individual growth rate and de-lifed fitness as lifetime fitness measures, and as annual de-lifed fitness. We compared the power of these metrics to predict reproductive values and lineage survival to the end of the study period. The three short-term fitness proxies predict the reproductive values and lineage survival only when measured at the recruit stage. There were no significant differences between the different fitness proxies at the same offspring stages in predicting the reproductive values and lineage survival. Annual fitness at one year old predicted reproductive values equally well as lifetime de-lifed fitness. However, none of the short-term fitness proxies were strongly associated with the reproductive values. The commonly used short-term fitness proxies best predict long-term fitness when measured at recruitment stage. Thus, because lifetime fitness measured at recruit stage and annual fitness in the first year of life were the best proxies of long-term fitness in short-lived birds, we encourage their future use.

The question of why socially monogamous females engage in extra‐pair behaviour is long‐standing in evolutionary biology. Due to a lack of empirical support among passerine birds, recent work has moved away from the indirect‐benefits hypothesis to explain extra‐pair mating behaviour by females, instead favouring the hypothesis that this is the result of a pleiotropic effect. That is, a trait under strong positive selection in either or both sexes are genetically linked with another, potentially unrelated, trait. For example, genes beneficial to female fecundity (that promote within‐pair solicitation of mating from a male partner) might also lead to extra‐pair behaviour (by also promoting solicited copulations from extra‐pair males). Here, we test two predictions from this hypothesis: We test the prediction that female divorce, measured as the number of social mates within a given year, is linked with 1) the number of extra‐pair males engaged by the female and 2) the proportion of the female's offspring that are extra‐pair. Our results show that females who divorce their social partner are more likely to produce extra‐pair offspring than those who maintain social monogamy, supporting the pleiotropy hypothesis. However, those females did not also have a higher proportion of extra‐pair offspring. The number of broods initiated was also positively correlated with the number of extra‐pair males that sired a female's offspring, probably through increased opportunity for extra‐pair males to sire offspring over a longer breeding season. Our results support the intrasexual pleiotropy hypothesis as a driver of female extra‐pair behaviour.

Fitness in birds has been shown to be negatively associated with anthropogenic noise, but the underlying mechanisms remain obscure. It is however crucial to understand the mechanisms of how urban noise impinges on fitness to obtain a better understanding of the role of chronic noise in urban ecology. Here, we examine three hypotheses on how noise might reduce reproductive output in passerine birds: (H1) by impairing mate choice, (H2) by reducing territory quality and (H3) by impeding chick development. We used long-term data from an island population of house sparrows, Passer domesticus, in which we can precisely estimate fitness. We found that nests in an area affected by the noise from large generators produced fewer young, of lower body mass, and fewer recruits, even when we corrected statistically for parental genetic quality using a cross-fostering set-up, supporting H3. Also, individual females provided their young with food less often when they bred in the noisy area compared to breeding attempts by the same females elsewhere. Furthermore, we show that females reacted flexibly to increased noise levels by adjusting their provisioning rate in the short term, which suggests that noise may be a causal factor that reduces reproductive output. We rejected H1 and H2 because nestbox occupancy, parental body mass, age and reproductive investment did not differ significantly between noisy and quiet areas. OUR RESULTS SUGGEST A PREVIOUSLY UNDESCRIBED MECHANISM TO EXPLAIN HOW ENVIRONMENTAL NOISE CAN REDUCE FITNESS IN PASSERINE BIRDS: by acoustically masking parent-offspring communication. More importantly, using a cross-fostering set-up, our results demonstrate that birds breeding in a noisy environment experience significant fitness costs. Chronic noise is omnipresent around human habitation and may produces similar fitness consequences in a wide range of urban bird species.

Helping by group members is predicted to lead to delayed senescence by affecting the trade-off between current reproduction and future survival for dominant breeders. Here we investigate this prediction in the Seychelles warbler, Acrocephalus sechellensis , in which mainly female subordinate helpers (both co-breeders and non-breeding helpers) often help dominants raise offspring. We find that the late-life decline in survival usually observed in this species is greatly reduced in female dominants when a helper is present. Female dominants with a female helper show reduced telomere attrition, a measure that reflects biological ageing in this and other species. Finally, the probability of having female, but not male, helpers increases with dominant female age. Our results suggest that delayed senescence is a key benefit of cooperative breeding for elderly dominants and support the idea that sociality and delayed senescence are positively self-reinforcing. Such an effect may help explain why social species often have longer lifespans. Sociality explains substantial variation in ageing across species, but less is known about this relationship within species. Here, the authors show that female dominant Seychelles warblers with helpers at the nest have higher late-life survival and lower telomere attrition and the probability of having helpers increases with age.

In a rapidly changing world, it is of fundamental importance to understand processes constraining or facilitating adaptation through microevolution. As different traits of an organism covary, genetic correlations are expected to affect evolutionary trajectories. However, only limited empirical data are available. We investigate the extent to which multivariate constraints affect the rate of adaptation, focusing on four morphological traits often shown to harbour large amounts of genetic variance and considered to be subject to limited evolutionary constraints. Our data set includes unique long-term data for seven bird species and a total of 10 populations. We estimate population-specific matrices of genetic correlations and multivariate selection coefficients to predict evolutionary responses to selection. Using Bayesian methods that facilitate the propagation of errors in estimates, we compare (1) the rate of adaptation based on predicted response to selection when including genetic correlations with predictions from models where these genetic correlations were set to zero and (2) the multivariate evolvability in the direction of current selection to the average evolvability in random directions of the phenotypic space. We show that genetic correlations on average decrease the predicted rate of adaptation by 28%. Multivariate evolvability in the direction of current selection was systematically lower than average evolvability in random directions of space. These significant reductions in the rate of adaptation and reduced evolvability were due to a general nonalignment of selection and genetic variance, notably orthogonality of directional selection with the size axis along which most (60%) of the genetic variance is found. These results suggest that genetic correlations can impose significant constraints on the evolution of avian morphology in wild populations. This could have important impacts on evolutionary dynamics and hence population persistence in the face of rapid environmental change.

Theory predicts that individuals behave altruistically towards their relatives. Hence, some form of kin recognition is useful for individuals to optimize their behavior. In species that display bi-parental care and are subject to extra-pair matings, kin recognition theoretically can allow cuckolded fathers to reduce their parental investment, and thus optimize their fitness. Whether this is possible remains unclear in birds. This study investigates whether males provide differential parental care depending on relatedness, as a proxy to recognizing chicks in their nest as kin or not. We cross-fostered House sparrow (Passer domesticus) chicks after hatching, and then expected that fathers would show a decrease in their parental efforts when tending to a clutch of unrelated offspring. House sparrow males are able to adjust their parental care to the identity of their partner, making them an ideal study species. However, there was no significant effect of relatedness on provisioning rates. This suggests that sparrows may not be capable of kin recognition, or at least do not display kin discrimination despite its apparent evolutionary advantage.

Telomeres, DNA structures located at the end of eukaryotic chromosomes, shorten with each cellular cycle. The shortening rate is affected by factors associated with stress, and, thus telomere length has been used as a biomarker of ageing, disease, and different life history trade-offs. Telomere research has received much attention in the last decades, however there is still a wide variety of factors that may affect telomere measurements and to date no study has thoroughly evaluated the possible long-term effect of a storage medium on telomere measurements. In this study we evaluated the long-term effects of ethanol on relative telomere length (RTL) measured by qPCR, using blood samples of magpies collected over twelve years and stored in absolute ethanol at room temperature. We firstly tested whether storage time had an effect on RTL and secondly we modelled the effect of time of storage (from 1 to 12 years) in differences in RTL from DNA extracted twice in consecutive years from the same blood sample. We also tested whether individual amplification efficiencies were influenced by storage time, and whether this could affect our results. Our study provides evidence of an effect of storage time on telomere length measurements. Importantly, this effect shows a pattern of decreasing loss of telomere sequence with storage time that stops after approximate 4 years of storage, which suggests that telomeres may degrade in blood samples stored in ethanol. Our method to quantify the effect of storage time could be used to evaluate other storage buffers and methods. Our results highlight the need to evaluate the long-term effects of storage on telomere measurements, particularly in long-term studies.

Life-history theory, central to our understanding of diversity in morphology, behaviour, and senescence, describes how traits evolve through the optimisation of trade-offs in investment. Despite considerable study, there is only minimal support for trade-offs within species between the two traits most closely linked to fitness – reproductive effort and survival – questioning the theory’s general validity. We used a meta-analysis to separate the effects of individual quality (positive survival/reproduction correlation) from the costs of reproduction (negative survival/reproduction correlation) using studies of reproductive effort and parental survival in birds. Experimental enlargement of brood size caused reduced parental survival. However, the effect size of brood size manipulation was small and opposite to the effect of phenotypic quality, as we found that individuals that naturally produced larger clutches also survived better. The opposite effects on parental survival in experimental and observational studies of reproductive effort provide the first meta-analytic evidence for theory suggesting that quality differences mask trade-offs. Fitness projections using the overall effect size revealed that reproduction presented negligible costs, except when reproductive effort was forced beyond the maximum level observed within species, to that seen between species. We conclude that there is little support for the most fundamental life-history trade-off, between reproductive effort and survival, operating within a population. We suggest that within species the fitness landscape of the reproduction–survival trade-off is flat until it reaches the boundaries of the between-species fast–slow life-history continuum. Our results provide a quantitative explanation as to why the costs of reproduction are not apparent and why variation in reproductive effort persists within species.

Measuring parental care behaviour in the wild is central to the study of animal ecology and evolution, but it is often labour‐ and time‐intensive. Efficient open‐source tools have recently emerged that allow animal behaviour to be quantified from videos using machine learning and computer vision techniques, but there is limited appraisal of how these tools perform compared to traditional methods. To gain insight into how different methods perform in extracting data from videos taken in the field, we compared estimates of the parental provisioning rate of wild house sparrows Passer domesticus from video recordings. We compared four methods: manual annotation by experts, crowd‐sourcing, automatic detection based on the open‐source software DeepMeerkat, and a hybrid annotation method. We found that the data collected by the automatic method correlated with expert annotation (r = 0.62) and further show that these data are biologically meaningful as they predict brood survival. However, the automatic method produced largely biased estimates due to the detection of non‐visitation events, while the crowd‐sourcing and hybrid annotation produced estimates that are equivalent to expert annotation. The hybrid annotation method takes approximately 20% of annotation time compared to manual annotation, making it a more cost‐effective way to collect data from videos. We provide a successful case study of how different approaches can be adopted and evaluated with a pre‐existing dataset, to make informed decisions on the best way to process video datasets. If pre‐existing frameworks produce biased estimates, we encourage researchers to adopt a hybrid approach of first using machine learning frameworks to preprocess videos, and then to do manual annotation to save annotation time. As open‐source machine learning tools are becoming more accessible, we encourage biologists to make use of these tools to cut annotation time but still get equally accurate results without the need to develop novel algorithms from scratch.

Chromosomal inversions frequently underlie major phenotypic variation maintained by divergent selection within and between sexes. Here we examine whether and how intralocus conflicts contribute to balancing selection stabilizing an autosomal inversion polymorphism in the ruff Calidris pugnax . In this lekking shorebird, three male mating morphs (Independents, Satellites and Faeders) are controlled by an inversion-based supergene. We show that in a captive population, Faeder females, who are smaller and whose inversion haplotype has not undergone recombination, have lower average reproductive success in terms of laying rate, egg size, and offspring survival than Independent females, who lack the inversion. Satellite females, who carry a recombined inversion haplotype and have intermediate body size, more closely resemble Independent than Faeder females in reproductive performance. We inferred that the lower reproductive output of Faeder females is most likely balanced by higher than average reproductive success of individual Faeder males. These findings suggest that intralocus conflicts may play a major role in the evolution and maintenance of supergene variants. ‘A supergene that underlies variation in male mating phenotypes has consequences for female reproduction. Here, the authors use evolutionary models to show that the rarest variant of this supergene is maintained by disproportionally high male reproductive success.’