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Cooperative breeding occurs when helpers provide alloparental care to the offspring of a breeding pair. One hypothesis of why helping occurs is that helpers gain valuable skills that may increase their own future reproductive success. However, research typically focuses on the effect of helping on short-term measures of reproductive success. Fewer studies have considered how helping affects long-term fitness measures. Here, we analyze how helping experience affects key breeding and fitness-related parameters in the Seychelles warbler (Acrocephalus sechellensis). Importantly, we control for females that have cobred (reproduced as a subordinate by laying an egg within a territory in which they are not a dominant breeder), as they already have experience with direct reproduction. Helping experience had no significant association with any of the metrics considered, except that helpers had an older age at first dominance. Accounting for helping experience, females that had cobred produced more adult offspring (≥1 year) after acquiring dominance and had a higher lifetime reproductive success (LRS) than females that had never cobred. Our results suggest that, in the Seychelles warbler, helping experience alone does not increase the fitness of helpers in any of the metrics considered, and highlights the importance of separating the effects of helping from cobreeding. Our findings also emphasize the importance of analyzing the effect of helping at various life-history stages, as higher short-term fitness may not translate to an overall increase in LRS., (© The Author(s) 2023. Published by Oxford University Press on behalf of The Society for the Study of Evolution (SSE).)

In cooperatively breeding birds, why do some individuals breed independently but others have to help at home? This question has been rarely addressed despite its fundamental importance for understanding the evolution of social cooperation. We address it using 15 years of data from Tibetan ground tits Pseudopodoces humilis where helpers consist of younger males. Since whether younger males successfully breed depends critically on their chances to occupy territories nearby home, our analytic strategy is to identify the determinants of individual differences in gaining territory ownership among these ready-to-breed males. Across widowed, last-year helper and yearling males, an age advantage was evident in inheriting resident territories, occupying adjacent vacancies and budding off part of adjacent territories, which left some last-year helpers and most yearling males to take the latter two routes. These males were more likely to acquire a territory if they were genetically related to the previous or current territory owners; otherwise they remained on natal territories as helpers. The relatedness effect can arise from the prior residence advantage established in the preceding winter when younger males followed their parents to perform kin-directed off-territory forays. Our research highlights the key role of local kinship in determining younger males' territory acquisition and thus their fate in terms of independent reproduction versus help. This finding provides insight into the formation of kin-based, facultative cooperative societies prevailing among vertebrates., (© 2023 The Authors. Journal of Animal Ecology © 2023 British Ecological Society.)

Parental care patterns differ enormously among and even within species. This is exemplified by Chinese penduline tits Remiz consobrinus, where biparental care, female-only care, male-only care and biparental desertion all occur in the same population; moreover, the distribution of these care patterns differs systematically between populations. The eco-evolutionary determinants of this diversity are largely unknown. We developed an individual-based model that allows us to investigate the effects of season length and offspring needs (expressed by the efficacy with which a clutch can be raised by a single parent) on the evolution of parental care patterns. The model is largely conceptual, aiming at general conclusions. However, to keep the model realistic, its set-up and the choice of parameters are motivated by field studies on Chinese penduline tits. Exploring a wide range of parameters, we investigate how parental care patterns are affected by season length and offspring needs and whether and under what conditions diverse parental care patterns can stably coexist. We report five main findings. First, under a broad range of conditions, different care patterns (e.g. male care and biparental care) coexist at equilibrium. Second, for the same parameters, alternative evolutionary equilibria are possible; this can explain differences in care patterns across populations. Third, rapid evolutionary transitions can occur between alternative equilibria; this can explain the often-reported evolutionary lability of parental care patterns. Fourth, season length has a strong but nonmonotonic effect on the evolved care patterns. Fifth, when uniparental care efficacy is low, biparental care tends to evolve; however, in many scenarios uniparental care is still common at equilibrium. In addition, our study sheds new light on Trivers' hypothesis that the sex with the highest prezygotic investment is predestined to invest more postzygotically as well. Our study highlights that diversity in parental care can readily evolve and it shows that even in the absence of environmental change parental care patterns can be evolutionary labile. In the presence of directional environmental change, systematic shifts in care patterns are to be expected., (© 2023 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Individual variation in telomere length is predictive of health and mortality risk across a range of species. However, the relative influence of environmental and genetic variation on individual telomere length in wild populations remains poorly understood. Heritability of telomere length has primarily been calculated using parent-offspring regression which can be confounded by shared environments. To control for confounding variables, quantitative genetic "animal models" can be used, but few studies have applied animal models in wild populations. Furthermore, parental age at conception may also influence offspring telomere length, but most studies have been cross-sectional. We investigated within- and between-parental age at conception effects and heritability of telomere length in the Seychelles warbler using measures from birds caught over 20 years and a multigenerational pedigree. We found a weak negative within-paternal age at conception effect (as fathers aged, their offspring had shorter telomeres) and a weak positive between-maternal age at conception effect (females that survived to older ages had offspring with longer telomeres). Animal models provided evidence that heritability and evolvability of telomere length were low in this population, and that variation in telomere length was not driven by early-life effects of hatch period or parental identities. Quantitative polymerase chain reaction plate had a large influence on telomere length variation and not accounting for it in the models would have underestimated heritability. Our study illustrates the need to include and account for technical variation in order to accurately estimate heritability, as well as other environmental effects, on telomere length in natural populations., (© 2021 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

In socially monogamous species, extra-pair paternity (EPP) is predicted to increase variance in male reproductive success (RS) beyond that resulting from genetic monogamy, thus, increasing the "opportunity for selection" (maximum strength of selection that can act on traits). This prediction is challenging to investigate in wild populations because lifetime reproduction data are often incomplete. Moreover, age-specific variances in reproduction have been rarely quantified. We analyzed 21 years of near-complete social and genetic reproduction data from an insular population of Seychelles warblers (Acrocephalus sechellensis). We quantified EPP's contribution to lifetime and age-specific opportunities for selection in males. We compared the variance in male genetic RS vs social ("apparent") RS (RS ap ) to assess if EPP increased the opportunity for selection over that resulting from genetic monogamy. Despite not causing a statistically significant excess (19%) of the former over the latter, EPP contributed substantially (27%) to the variance in lifetime RS, similarly to within-pair paternity (WPP, 39%) and to the positive WPP-EPP covariance (34%). Partitioning the opportunity for selection into age-specific (co)variance components, showed that EPP also provided a substantial contribution at most ages, varying with age. Therefore, despite possibly not playing the main role in shaping sexual selection in Seychelles warblers, EPP provided a substantial contribution to the lifetime and age-specific opportunity for selection, which can influence evolutionary processes in age-structured populations., (© 2022 The Authors. Evolution © 2022 The Society for the Study of Evolution.)

The social environment in which individuals live affects their fitness and in turn population dynamics as a whole. Birds with facultative cooperative breeding can live in social groups with dominants, subordinate helpers that assist with the breeding of others, and subordinate non-helpers. Helping behaviour benefits dominants through increased reproductive rates and reduced extrinsic mortality, such that cooperative breeding might have evolved in response to unpredictable, harsh conditions affecting reproduction and/or survival of the dominants. Additionally, there may be different costs and benefits to both helpers and non-helpers, depending on the time-scale. For example, early-life costs might be compensated by later-life benefits. These differential effects are rarely analysed in the same study. We examined whether helping behaviour affects population persistence in a stochastic environment and whether there are direct fitness consequences of different life-history tactics adopted by helpers and non-helpers. We parameterised a matrix population model describing the population dynamics of female Seychelles warblers Acrocephalus sechellensis, birds that display facultative cooperative breeding. The stochastic density-dependent model is defined by a (st)age structure that includes life-history differences between helpers and non-helpers and thus can estimate the demographic mechanisms of direct benefits of helping behaviour. We found that population dynamics are strongly influenced by stochastic variation in the reproductive rates of the dominants, that helping behaviour promotes population persistence and that there are only early-life differences in the direct fitness of helpers and non-helpers. Through a matrix population model, we captured multiple demographic rates simultaneously and analysed their relative importance in determining population dynamics of these cooperative breeders. Disentangling early-life versus lifetime effects of individual tactics sheds new light on the costs and benefits of helping behaviour. For example, the finding that helpers and non-helpers have similar lifetime reproductive outputs and that differences in reproductive values between the two life-history tactics arise only in early life suggests that overall, helpers and non-helpers have a similar balance of costs and benefits when analysing direct benefits. We recommend analysing the consequence of different life-history tactics, during both early life and over the lifetime, as analyses of these different time frames may produce conflicting results., (© 2021 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Understanding where genetic variation exists, and how it influences fitness within populations is important from an evolutionary and conservation perspective. Signatures of past selection suggest that pathogen-mediated balancing selection is a key driver of immunogenetic variation, but studies tracking contemporary evolution are needed to help resolve the evolutionary forces and mechanism at play. Previous work in a bottlenecked population of Seychelles warblers (Acrocephalus sechellensis) show that functional variation has been maintained at the viral-sensing Toll-like receptor 3 (TLR3) gene, including one nonsynonymous SNP, resulting in two alleles. Here, we characterise evolution at this TLR3 locus over a 25-year period within the original remnant population of the Seychelles warbler, and in four other derived, populations. Results show a significant and consistent temporal decline in the frequency of the TLR3 C allele in the original population, and that similar declines in the TLR3 C allele frequency occurred in all the derived populations. Individuals (of both sexes) with the TLR3 CC genotype had lower survival, and males - but not females - that carry the TLR3 C allele had significantly lower lifetime reproductive success than those with only the TLR3 A allele. These results indicate that positive selection on the TLR3 A allele, caused by an as yet unknown agent, is driving TLR3 evolution in the Seychelles warbler. No evidence of heterozygote advantage was detected. However, whether the positive selection observed is part of a longer-term pattern of balancing selection (through fluctuating selection or rare-allele advantage) cannot be resolved without tracking the TLR3 C allele over an extended time period., (© 2021 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

Senescence is widespread in nature, often resulting in diminishing survival or reproduction with age, but its role in age-dependent variation in sexual traits is often poorly understood. One reason is that few studies of sexual traits consider non-linear relationships with age, or only consider a narrow range of years relative to the life span of the species. Birdsong has evolved to allow assessment of conspecific quality in numerous bird species. Whilst theory and empirical work suggests that song may become more elaborate with age, there are a paucity of long-term studies testing whether song is associated with age or longevity. In particular, the occurrence of song senescence has rarely been demonstrated. Using an exceptional long-term dataset for the Seychelles warbler (Acrocephalus sechellensis), we analysed relationships between male song, age, survival, and longevity. This species is a long-lived songbird with early life increases, followed by senescent declines, in survival and reproduction. The study population (Cousin Island, Seychelles) is a closed population, with no depredation of adults, providing an excellent opportunity to study senescence in free-living animals. We tested whether song traits were related to age at recording, future survival, longevity, and territory quality. We found age-dependent changes in five song traits (duration, maximum frequency, peak frequency of songs, and duration and frequency bandwidth of trills). Relationships with age were quadratic, indicating reversal in the expression of song coinciding with the onset of senescence in reproduction and survival in this species. One song trait (trill bandwidth) had a quadratic relationship with future survival, but no song traits were related to longevity, suggesting age-related patterns were not the result of selective disappearance. Our study provides one of the first examples of functional senescence in song, offering new insights into avian senescence. Late-life declines in avian song, and possibly other sexual traits, may be more common than currently known, and may play a fundamental role in age-dependent changes in reproductive success.

Extra-pair paternity (EPP) is often linked to male age in socially monogamous vertebrates; that is, older males are more likely to gain EPP and less likely to be cuckolded. However, whether this occurs because males improve at gaining paternity as they grow older, or because "higher quality" males that live longer are preferred by females, has rarely been tested, despite being central to our understanding of the evolutionary drivers of female infidelity. Moreover, how extra-pair reproduction changes with age within females has received even less attention. Using 18 years of longitudinal data from an individually marked population of Seychelles warblers (Acrocephalus sechellensis), we found considerable within-individual changes in extra-pair reproduction in both sexes: an early-life increase and a late-life decline. Furthermore, males were cuckolded less as they aged. Our results indicate that in this species age-related patterns of extra-pair reproduction are determined by within-individual changes with age, rather than differences among individuals in longevity. These results challenge the hypothesis-based on longevity reflecting intrinsic quality-that the association between male age and EPP is due to females seeking high-quality paternal genes for offspring. Importantly, EPP accounted for up to half of male reproductive success, emphasizing the male fitness benefits of this reproductive strategy. Finally, the occurrence of post-peak declines in extra-pair reproduction provides explicit evidence of senescence in infidelity in both males and females., (© 2020 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

In many cooperatively breeding animals, a combination of ecological constraints and benefits of philopatry favours offspring taking a subordinate position on the natal territory instead of dispersing to breed independently. However, in many species individuals disperse to a subordinate position in a non-natal group ("subordinate between-group" dispersal), despite losing the kin-selected and nepotistic benefits of remaining in the natal group. It is unclear which social, genetic and ecological factors drive between-group dispersal. We aim to elucidate the adaptive significance of subordinate between-group dispersal by examining which factors promote such dispersal, whether subordinates gain improved ecological and social conditions by joining a non-natal group, and whether between-group dispersal results in increased lifetime reproductive success and survival. Using a long-term dataset on the cooperatively breeding Seychelles warbler (Acrocephalus sechellensis), we investigated how a suite of proximate factors (food availability, group composition, age and sex of focal individuals, population density) promote subordinate between-group dispersal by comparing such dispersers with subordinates that dispersed to a dominant position or became floaters. We then analysed whether subordinates that moved to a dominant or non-natal subordinate position, or became floaters, gained improved conditions relative to the natal territory and compared fitness components between the three dispersal strategies. We show that individuals that joined another group as non-natal subordinates were mainly female and that, similar to floating, between-group dispersal was associated with social and demographic factors that constrained dispersal to an independent breeding position. Between-group dispersal was not driven by improved ecological or social conditions in the new territory and did not result in higher survival. Instead, between-group dispersing females often became cobreeders, obtaining maternity in the new territory, and were likely to inherit the territory in the future, leading to higher lifetime reproductive success compared to females that floated. Males never reproduced as subordinates, which may be one explanation why subordinate between-group dispersal by males is rare. Our results suggest that subordinate between-group dispersal is used by females to obtain reproductive benefits when options to disperse to an independent breeding position are limited. This provides important insight into the additional strategies that individuals can use to obtain reproductive benefits., (© 2018 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Because virtually all organisms compete with others in their social environment, mechanisms that reduce conflict between interacting individuals are crucial for the evolution of stable families, groups, and societies. Here, we tested whether costs of social conflict over territorial space between Seychelles warblers ( Acrocephalus sechellensis ) are mitigated by kin-selected (genetic relatedness) or mutualistic (social familiarity) mechanisms. By measuring longitudinal changes in individuals' body mass and telomere length, we demonstrated that the fitness costs of territoriality are driven by a complex interplay between relatedness, familiarity, local density, and sex. Physical fights were less common at territory boundaries shared between related or familiar males. In line with this, male territory owners gained mass when living next to related or familiar males and also showed less telomere attrition when living next to male kin. Importantly, these relationships were strongest in high-density areas of the population. Males also had more rapid telomere attrition when living next to unfamiliar male neighbors, but mainly when relatedness to those neighbors was also low. In contrast, neither kinship nor familiarity was linked to body mass or telomere loss in female territory owners. Our results indicate that resolving conflict over territorial space through kin-selected or mutualistic pathways can reduce both immediate energetic costs and permanent somatic damage, thus providing an important mechanism to explain fine-scale population structure and cooperation between different social units across a broad range of taxa., Competing Interests: The authors declare no conflict of interest.

Understanding why individuals delay dispersal and become subordinates within a group is central to studying the evolution of sociality. Hypotheses predict that dispersal decisions are influenced by costs of extra-territorial prospecting that are often required to find a breeding vacancy. Little is known about such costs, partly because it is complicated to demonstrate them empirically. For example, prospecting individuals may be of inferior quality already before prospecting and/or have been evicted. Moreover, costs of prospecting are mainly studied in species where prospectors suffer from predation risk, so how costly prospecting is when predators are absent remains unclear. Here, we determine a cost of prospecting for subordinate Seychelles warblers, Acrocephalus sechellensis, in a population where predators are absent and individuals return to their resident territory after prospecting. Prospecting individuals had 5.2% lower body mass than non-prospecting individuals. Our evidence suggests this may be owing to frequent attacks by resident conspecifics, likely leading to reduced food intake by prospectors. These results support the hypothesis that energetic costs associated with dispersal opportunities are one factor influencing dispersal decisions and shaping the evolution of delayed dispersal in social animals., (© 2016 The Authors.)

Desertion of clutch (or brood) by both parents often leads to breeding failure, since in vast majority of birds care by at least one parent is required for any young to fledge. Recent works in a highly polygamous passerine bird, the Eurasian penduline tit (Remiz pendulinus), suggest that biparental clutch desertion is due to intense sexual conflict over care. However, an alternative yet untested hypothesis for biparental desertion is low offspring viability so that the parents abandon the offspring that have poor prospect for survival. Here we test the latter hypothesis in a common garden experiment by comparing the viability of deserted and cared for eggs. We show that embryonic development does not differ between deserted and cared for eggs. Therefore, sexual conflict over care remains the best supported hypothesis for biparental clutch desertion in penduline tits. Our work points out that conflict over care is a potential - yet rarely considered - cause of biparental nest desertion, and a strong alternative for the traditional explanations of low offspring viability, human disturbance or deteriorating ambient environment. Apart from a handful of species, the intensity of sexual conflict has not been quantified, and we call for further studies to consider sexual conflict as a cause of nest desertion., (Copyright © 2015 Elsevier B.V. All rights reserved.)

Eurasian penduline tits (Remiz pendulinus) have an unusually diverse breeding system consisting of frequent male and female polygamy, and uniparental care by the male or the female. Intriguingly, 30 to 40 per cent of all nests are deserted by both parents. To understand the evolution of this diverse breeding system and frequent clutch desertion, we use 6 years of field data to derive fitness expectations for males and females depending on whether or not they care for their offspring. The resulting payoff matrix corresponds to an asymmetric Snowdrift Game with two alternative evolutionarily stable strategies (ESSs): female-only and male-only care. This, however, does not explain the polymorphism in care strategies and frequent biparental desertion, because theory predicts that one of the two ESSs should have spread to fixation. Using a bootstrapping approach, we demonstrate that taking account of individual variation in payoffs explains the patterns of care better than a model based on the average population payoff matrix. In particular, a model incorporating differences in male attractiveness closely predicts the observed frequencies of male and female desertion. Our work highlights the need for a new generation of individual-based evolutionary game-theoretic models.

In many iteroparous species individual fitness components, such as reproductive output, first increase with age and then decline during late-life. However, individuals differ greatly in reproductive lifespan, but reproductive declines may only occur in the period just before their death as a result of an age-independent decline in physiological condition. To fully understand reproductive senescence it is important to investigate to what extent declines in late-life reproduction can be explained by age, time until death, or both. However, the study of late-life fitness performance in natural populations is challenging as the exact birth and death dates of individuals are often not known, and most individuals succumb to extrinsic mortality before reaching old age. Here, we used an exceptional long-term longitudinal dataset of individuals from a natural, closed, and predator-free population of the Seychelles warbler (Acrocephalus sechellensis) to investigate reproductive output, both in relation to age and to the time until the death of an individual (reverse-age approach). We observed an initial age-dependent increase in reproductive output that was followed by a decline in old age. However, we found no significant decline in reproductive output in the years directly preceding death. Although post-peak reproductive output declined with age, this pattern differed between terminal and non-terminal reproductive attempts, and the age-dependence of the terminal breeding attempt explained much of the variation in age-specific reproductive output. In fact, terminal declines in reproductive output were steeper in very old individuals. These results indicate that not only age-dependent, but also age-independent factors, such as physiological condition, need to be considered to understand reproductive senescence in wild-living animals.

The major histcompatibility complex (MHC) is a vital component of the adaptive immune system in all vertebrates. This study is the first to characterize MHC class I (MHC-I) in blue tits (Cyanistes caeruleus), and we use MHC-I exon 3 sequence data from individuals originating from three locations across Europe: Spain, the Netherlands to Sweden. Our phylogeny of the 17 blue tit MHC-I alleles contains one allele cluster with low nucleotide diversity compared to the remaining more diverse alleles. We found a significant evidence for balancing selection in the peptide-binding region in the diverse allele group only. No separation according to geographic location was found in the phylogeny of alleles. Although the number of MHC-I loci of the blue tit is comparable to that of other passerine species, the nucleotide diversity of MHC-I appears to be much lower than that of other passerine species, including the closely related great tit (Parus major) and the severely inbred Seychelles warbler (Acrocephalus sechellensis). We believe that this initial MHC-I characterization in blue tits provides an important step towards understanding the mechanisms shaping MHC-I diversity in natural populations.

In many socially monogamous animals, females engage in extrapair copulation (EPC), causing some broods to contain both within-pair and extrapair young (EPY). The proportion of all young that are EPY varies across populations and species. Because an EPC that does not result in EPY leaves no forensic trace, this variation in the proportion of EPY reflects both variation in the tendency to engage in EPC and variation in the extrapair fertilization (EPF) process across populations and species. We analyzed data on the distribution of EPY in broods of four passerines (blue tit, great tit, collared flycatcher, and pied flycatcher), with 18,564 genotyped nestlings from 2,346 broods in two to nine populations per species. Our Bayesian modeling approach estimated the underlying probability function of EPC (assumed to be a Poisson function) and conditional binomial EPF probability. We used an information theoretical approach to show that the expected distribution of EPC per female varies across populations but that EPF probabilities vary on the above-species level (tits vs. flycatchers). Hence, for these four passerines, our model suggests that the probability of an EPC mainly is determined by ecological (population-specific) conditions, whereas EPF probabilities reflect processes that are fixed above the species level.

Background: The evolutionary interests of males and females rarely coincide (sexual conflict), and these conflicting interests influence morphology, behavior and speciation in various organisms. We examined consequences of variation in sexual conflict in two closely-related passerine birds with contrasting breeding systems: the Eurasian penduline tit Remiz pendulinus (EPT) exhibiting a highly polygamous breeding system with sexually antagonistic interests over parental care, and the socially monogamous Cape penduline tit Anthoscopus minutus (CPT). We derived four a priori predictions from sexual conflict theory and tested these using data collected in Central Europe (EPT) and South Africa (CPT). Firstly, we predicted that EPTs exhibit more sexually dimorphic plumage than CPTs due to more intense sexual selection. Secondly, we expected brighter EPT males to provide less care than duller males. Thirdly, since song is a sexually selected trait in many birds, male EPTs were expected to exhibit more complex songs than CPT males. Finally, intense sexual conflict in EPT was expected to lead to low nest attendance as an indication of sexually antagonistic interests, whereas we expected more cooperation between parents in CPT consistent with their socially monogamous breeding system., Results: Consistent with our predictions EPTs exhibited greater sexual dimorphism in plumage and more complex song than CPTs, and brighter EPT males provided less care than duller ones. EPT parents attended the nest less frequently and less simultaneously than CPT parents., Conclusions: These results are consistent with sexual conflict theory: species in which sexual conflict is more manifested (EPT) exhibited a stronger sexual dimorphism and more elaborated sexually selected traits than species with less intense sexual conflict (CPT). Our results are also consistent with the notion that EPTs attempt to force their partner to work harder as expected under sexual conflict: each member of the breeding pair attempts to shift the costs of care to the other parent. More brightly colored males benefit more from desertion than dull ones, because they are more likely to remate with a new female. Taken together, the comparison between two closely related species with contrasting breeding systems suggest that sexual conflict over care has influenced the evolution of behavior and morphology in penduline tits.

The explanation for extra-pair mating in female birds remains poorly understood and contentious [1-7]. Several leading hypotheses propose that females benefit indirectly by enhancing the genetic quality of their offspring, through good genes or genetic compatibility effects [1, 8, 9]. Supporting this idea, recent studies have identified a range of fitness-related traits for which extra-pair offspring (EPO) are superior to their within-pair (WP) half-siblings [10-21]. However, such performance differences may result from nongenetic maternal effects if EPO are positioned earlier in the laying order and benefit from the advantages of earlier hatching [22, 23]. Here we show that EPO are larger, heavier, and more likely to fledge than their WP half-siblings in a population of blue tits, Cyanistes caeruleus. However, extra-pair paternity declined markedly with laying order, resulting in EPO generally hatching earlier. After correcting for variation in hatch time, none of the observed disparities between EPO and their WP half-siblings remained significant. These findings indicate that phenotypic comparisons between maternal half-siblings must consider potential hatching-order effects and suggest that the evidence for genetic benefits from extra-pair copulation may be less compelling than currently accepted. Moreover, the overrepresentation of EPO early in the laying order may help explain female extra-pair mating.

Temporal variation in survival, fecundity, and dispersal rates is associated with density-dependent and density-independent processes. Stable natural populations are expected to be regulated by density-dependent factors. However, detecting this by investigating natural variation in density is difficult because density-dependent and independent factors affecting population dynamics may covary. Therefore, experiments are needed to assess the density dependence of demographic rates. In this study, we investigate the effect of density on demographic rates of the Seychelles Warbler (Acrocephalus sechellensis). This species, endemic to a few islands in the Indian Ocean, went through a severe population bottleneck in the middle of the last century, with only approximately 30 individuals left on one small island, but has since recovered. Our monitoring shows that since reaching the island's carrying capacity, population density has remained stable. However, we detected neither density-dependent reproduction nor survival on the basis of natural density variation during this stable period. For conservation reasons, new populations have been established by transferring birds to nearby suitable islands. Using the change of numbers during the process of saturation as a natural experiment, we investigated whether we can detect regulation of numbers via density-dependent survival and reproduction within these new populations. We found that populations were mainly regulated by density-dependent reproduction, and not survival. Variation in density between islands can be explained by food abundance, measured as insect density. Islands with the highest insect densities also had the highest bird densities and the largest breeding groups. Consequently, we suggest that the density-dependent effect on reproduction is caused by competition for food.

Background: The trade-off between current and future parental investment is often different between males and females. This difference may lead to sexual conflict between parents over care provisioning in animals that breed with multiple mates. One of the most obvious manifestations of sexual conflict over care is offspring desertion whereby one parent deserts the young to increase its reproductive success at the expense of its mate. Offspring desertion is a wide-spread behavior, and its frequency often varies within populations. We studied the consistency of offspring desertion in a small passerine bird, the Eurasian penduline tit Remiz pendulinus, that has an extremely variable breeding system. Both males and females are sequentially polygamous, and a single parent (either the male or the female) incubates the eggs and rears the young. About 28-40% of offspring are abandoned by both parents, and these offspring perish. Here we investigate whether the variation in offspring desertion in a population emerges either by each individual behaving consistently between different broods, or it is driven by the environment., Results: Using a three-year dataset from Southern Hungary we show that offspring desertion by females is consistent between nests. Male desertion, however, depends on ambient environment, because all males desert their nests early in the season and some of them care late in the season. Therefore, within-population variation in parental care emerges by sexually different mechanisms; between-individual variation was responsible for the observed pattern of offspring desertion in females, whereas within-individual variation was responsible for the observed pattern in males., Conclusion: To our knowledge, our study is the first that investigates repeatability of offspring desertion behavior in nature. The contrasting strategies of the sexes imply complex evolutionary trajectories in breeding behavior of penduline tits. Our results raise an intriguing question whether the sexual difference in caring/deserting decisions explain the extreme intensity of sexual conflict in penduline tits that produces a high frequency of biparentally deserted (and thus wasted) offspring.

Sex biases in the allocation of resources to offspring occur in a broad range of taxa. Parents have been shown to achieve such biases either by producing numerically more of one sex or by providing the individuals of one sex with a greater quantity of resources. In addition, skews in allocation could occur if the offspring of one sex receive resources of higher quality (greater nutritional or energetic value by weight or volume), although this mode of adjustment has, to our knowledge, never been demonstrated. We compared the types of prey and the metabolizable energy provisioned to male and female nestlings in one of the most sexually size dimorphic of all birds, the brown songlark, Cinclorhamphus cruralis. Within broods, we found that males not only received more prey than their smaller sisters, but also prey of apparently higher quality. This dietary disparity could result either from mothers actively discriminating between the sexes when providing prey or from competition among siblings. We suggest that sex differences in offspring diet quality may occur in a wide range of other taxa and function as an additional mechanism of sex allocation adjustment.

Oxidative damage, caused by reactive oxygen species during aerobic respiration, is thought to be an important mediator of life-history trade-offs. To mitigate oxidative damage, antioxidant defence mechanisms are deployed, often at the cost of resource allocation to other body functions. Both reduced resource allocation to body functions and direct oxidative damage may decrease individual fitness, through reducing survival and/or reproductive output., The oxidative costs of reproduction have gained much attention recently, but few studies have investigated the long-term consequences of oxidative damage on survival and (future) reproductive output under natural conditions., Using a wild population of the cooperatively breeding Seychelles warbler ( Acrocephalus sechellensis), we tested the prediction that high levels of reactive oxygen species, or high antioxidant investments to avoid oxidative damage, have fitness consequences because they reduce survival and/or reproductive output., We found that individuals with higher circulating non-enzymatic antioxidant capacity had a lower probability of surviving until the next year. However, neither current reproductive output, nor future reproductive output in the surviving individuals, was associated with circulating non-enzymatic antioxidant capacity or oxidative damage., The negative relationship between antioxidant capacity and survival that we observed concurs with the findings of an extensive comparative study on birds, however the mechanisms underlying this association remain to be resolved., A is available for this article. [ABSTRACT FROM AUTHOR]

Many studies have assessed the costs of sibling rivalry in systems where offspring always have competitors, but conclusions about sibling rivalry in these species are restricted to interpreting the cost of changes in the relative level of competition and are often complicated by the expression of potentially costly rivalry related traits. Additionally, the majority of studies focus on early-life sibling rivalry, but the costs of competition can also affect later-life performance. We test a suite of hypothesized immediate (early-life body mass, telomere length, and survival) and delayed (adult reproductive potential and lifespan) costs of sibling rivalry for offspring of differing competitive ability in Seychelles warblers, where most offspring are raised singly and hence competitor success can be compared to a competition-free scenario. Compared to those raised alone, all competing nestlings had lower body mass and weaker competitors experienced reduced survival. However, the stronger competitors appeared to have longer adult breeding tenures and lifespan than those raised alone. We propose that comparisons with competition-free groups, as well as detailed fitness measures across entire lifetimes, are needed to understand the evolution of sibling rivalry and thus individual reproductive strategy in wild systems. [ABSTRACT FROM AUTHOR]

Individuals that retain their former partners often perform better than conspecifics that switch partners. This may be due to high-quality individuals being more faithful to their partners and more productive. Investigations of the fitness benefits of mate retention that also control for potential confounding effects of individuals are scarce. We studied the influence of mate retention and breeding experience on breeding performance of the hair-crested drongo (Dicrurus hottentottus) by carrying out both cross-sectional and longitudinal analyses. Pairs with longer pair-bond duration did not fledge more young or fledglings of better body condition nor did they produce more or better fledglings than newly formed pairs consisting of at least 1 experienced breeder, that is, individuals that had bred before. Individuals produced fewer fledglings when they were paired with an inexperienced breeder, especially when females were paired with inexperienced males. Although clutch size was not affected by mate retention or breeding experience, pairs consisting of inexperienced breeder(s) had a relatively higher predation rate of eggs and/or nestlings because they may be less effective in nest defense. The onset of breeding was advanced in the year following mate retention, but not in the second year thereafter, when pairs still remained together. Furthermore, only the breeding experience of the male determined the onset of breeding: Pairs consisting of inexperienced males bred later in the season. Our results suggest that breeding experience, and particularly the breeding experience of the male, but not mate retention, is important in determining the breeding performance of hair-crested drongos. [ABSTRACT FROM AUTHOR]

Inbreeding results in more homozygous offspring that should suffer reduced fitness, but it can be difficult to quantify these costs for several reasons. First, inbreeding depression may vary with ecological or physiological stress and only be detectable over long time periods. Second, parental homozygosity may indirectly affect offspring fitness, thus confounding analyses that consider offspring homozygosity alone. Finally, measurement of inbreeding coefficients, survival and reproductive success may often be too crude to detect inbreeding costs in wild populations. Telomere length provides a more precise measure of somatic costs, predicts survival in many species and should reflect differences in somatic condition that result from varying ability to cope with environmental stressors. We studied relative telomere length in a wild population of Seychelles warblers ( Acrocephalus sechellensis) to assess the lifelong relationship between individual homozygosity, which reflects genome-wide inbreeding in this species, and telomere length. In juveniles, individual homozygosity was negatively associated with telomere length in poor seasons. In adults, individual homozygosity was consistently negatively related to telomere length, suggesting the accumulation of inbreeding depression during life. Maternal homozygosity also negatively predicted offspring telomere length. Our results show that somatic inbreeding costs are environmentally dependent at certain life stages but may accumulate throughout life. [ABSTRACT FROM AUTHOR]

Parental care is costly, and in many organisms, the male or the female parent benefits from reducing its own care which may be compensated for by its mate. One of the parents may even face all costs of parental care if its mate deserts and leaves him/her to care for the offspring alone. Theoretical models have generated contrasting predictions as to how parents negotiate a resolution of this sexual conflict over care, although empirical tests are largely lacking. We investigated pre-desertion behaviour (nest attendance) of a highly polygamous passerine, the Eurasian penduline tit ( Remiz pendulinus) that exhibits intense sexual conflict over care culminating in clutch desertion by the male, by the female or by both parents. We conjectured that nest attendance of parents should predict desertion, so that the lack of care provided by the deserting parent may be compensated for. By analysing over 200 000 video frames (2.50 ± 1.36 d per pair, 302 ± 170 min/d) of 20 pairs, we show that nest desertion is not a gradual process and cannot be predicted by nest attendance. These results are consistent with the argument that the predictable desertion may not be evolutionarily stable, and suggest that male and female penduline tits do not negotiate clutch desertion. [ABSTRACT FROM AUTHOR]

Species that pass repeatedly through narrow population bottlenecks (<100 individuals) are likely to have lost a large proportion of their genetic variation. Having genotyped 92 Raso larks Alauda razae, a Critically Endangered single-island endemic whose world population in the Cape Verdes over the last 100 years has fluctuated between about 15 and 130 pairs, we found variation at 7 of 21 microsatellite loci that successfully amplified, the remaining loci being monomorphic. At 6 of the polymorphic loci variation was sex-linked, despite the fact that these microsatellites were not sex-linked in the other passerine birds where they were developed. Comparative analysis strongly suggests that material from several different autosomes has been recently transferred to the sex chromosomes in larks. Sex-linkage might plausibly allow some level of heterozygosity to be maintained, even in the face of persistently small population sizes. [ABSTRACT FROM AUTHOR]

Knowing why certain individuals refrain from reproduction is fundamental to our understanding of the evolution of cooperative breeding. Although many studies have focused on the role of life history and ecological factors, the proximate mechanisms underlying this behaviour have received considerably less attention. We determined the natural variation in plasma testosterone (T) level and cloacal protuberance (CP) size – an indication of sperm storage – in successfully reproducing primary male and in reproductively inactive subordinate male Seychelles warblers ( Acrocephalus sechellensis). Primary males had significantly higher T levels during female’s fertile period and significantly larger CPs throughout the breeding season than subordinate males. To test if the lower T levels of subordinates were caused by their social status per se, we experimentally promoted subordinates to primary males, by permanent removal of primary males from their territories. The experimentally promoted males increased T production and CP sizes significantly, suggesting former suppression by primary males. However, T levels of the promoted males only marginally increased. As this could not be explained by their younger age, we conclude that recovery from social suppression may take a substantial time period. [ABSTRACT FROM AUTHOR]

Do the two parents at a nest make simultaneous decisions whether to care for their offspring or to desert? If a single parent is sufficient for rearing young, one parent (typically, the male) may desert and reproduce with a new mate within the same breeding season, leaving the other parent with the brunt of care. As each parent is expected to maximize its own reproductive success, the interests of the two parents do not necessarily coincide, and a sexual conflict over care may emerge. Here we investigate the process of clutch desertion in a small passerine bird, the Penduline Tit Remiz pendulinus. Among birds, this species has a remarkably variable breeding system, because a single parent (either the male or the female) may provide the full care of the young, whereas about 30% of clutches are abandoned by both parents. First, we show that biparental desertion occurs within a single day in 73.7% of the clutches ( n = 14), whereas desertion decisions are sequential in 26.3% of the clutches ( n = 5) (male first: 10.5% ( n = 2); female first: 15.8% ( n = 3); n = 19 clutches in total). Secondly, we observed the behaviour of both parents before desertion, and investigated whether desertion can be predicted from their behaviour. However, neither singing nor nest-building behaviour predicted whether the male or the female would desert. We therefore suggest that biparental desertion may be simultaneous by male and female in our population of Penduline Tits. Furthermore, the parents do not appear to signal their intention to desert their mate. We argue that the parents’ interest may be actually to disguise their intention to desert. [ABSTRACT FROM AUTHOR]

1. Variation in survival, a major determinant of fitness, may be caused by individual or environmental characteristics. Furthermore, interactions between individuals may influence survival through the negative feedback effects of density dependence. Compared to species in temperate regions, we have little knowledge about population processes and variation in fitness in tropical bird species. 2. To investigate whether variation in survival could be explained by population size or climatic variables we used capture–recapture models in conjunction with a long-term data set from an island population of the territorial, cooperatively breeding Seychelles warbler ( Acrocephalus sechellensis). The lack of migration out of the study population means that our results are not confounded by dispersal. 3. Annual survival was high, both for adults (84%) and juveniles (61%), and did not differ between the sexes. Although there was significant variation in survival between years, this variation could not be explained by overall population size or weather variables. 4. For territorial species, resource competition will work mainly on a local scale. The size of a territory and number of individuals living in it will therefore be a more appropriate measure of density than overall population density. Consequently, both an index of territory quality per individual (food availability) and local density, measured as group size, were included as individual covariates in our analyses. 5. Local density had a negative effect on survival; birds living in larger groups had lower survival probabilities than those living in small groups. Food availability did not affect survival. 6. Our study shows that, in a territorial species, although density-dependent effects might not be detectable at the population level they can be detected at the individual territory level − the scale at which individuals compete. These results will help to provide a better understanding of the small-scale processes involved in the dynamics of a population in general, but in particular in tropical species living in relatively stable environments. [ABSTRACT FROM AUTHOR]

Discusses the results of experiments on the transfers of Seychelles warblers, Acrocephalus sechellensis, to the unoccupied islands of Aride and Cousine to test the ecological constraints model and the benefits of philopatry model for cooperative breeding. Effect of habitat saturation and variation in cooperative breeding on the frequency of delayed dispersal.

Sex-ratio theory states that if the fitness costs to the parents of producing one offspring’s sex relative to the other are higher, parents should discount these costs by producing fewer individuals of the more costly sex. In the co-operatively breeding Seychelles warbler (Acrocephalus sechellensis) mothers adaptively modify the sex of their single egg toward daughters, the helping sex, when living on territories with rich resources where helpers increase parental reproductive success, but toward sons, the dispersing sex, when living on territories where resources are scarce and/or no helping benefits accrue. By modifying offspring sex ratio, parents maximize their inclusive fitness benefits. Pairs in high-quality territories gained significantly more inclusive fitness benefits (through helping and reproducing offspring) from the production of daughters than from sons, and vice versa in low-quality territories (through reproducing offspring). Experimental manipulation of the offspring’s sex shows that the consequences of sex allocation are adaptive for parents on high-quality territories. On high-quality territories with female production, breeding pairs raising step-daughters gained significantly higher inclusive benefits (through indirect and direct fitness gains) than by raising step-sons. [ABSTRACT FROM AUTHOR]

The process of nest leaving (fledging) in hole-breeding passerines is largely unexplored, although it is potentially an important facet of reproduction. We used the great tit, Parus major, to investigate whether fledging timing and order were affected by nestling development and sex, as well as the sex ratio in the brood the nestlings were raised in. Because of the difference in size and competitiveness between male and female nestlings, we expected to find an effect of sex and brood sex ratio (BSR) on the process of fledging. To explore its effect on fledging experimentally, we manipulated BSR by swapping 6-day-old nestlings of both sexes. We implanted transponders in all 14-day-old nestlings to determine timing and order of fledging. The brood age at fledging was best explained by average wing length of the brood (negative), average body mass of the brood (positive) and hatching date (negative). In contrast to our hypothesis, BSR did not affect fledging time. Also, the asynchrony of fledging within broods did not depend on BSR. Within broods, fledging order was not affected by sex or by the interaction between sex and BSR. Nestlings with long wings on day 14 fledged earlier than nestmates with shorter wings. Although females were lighter at day 14, they had similar length wings as their male nestmates. Nestlings should keep up with their nestmates during development, because developmental status relative to nestmates, rather than sex-specific competition, influenced the process of fledging. [Copyright &y& Elsevier]

Senescence – the progressive age-dependent decline in performance – occurs in most organisms. There is considerable variation in the onset and rate of senescence between and within species. Yet the causes of this variation are still poorly understood, despite being central to understanding the evolution of senescence. Long-term longitudinal studies on wild animals are extremely well-suited to studying the impact of environmental and individual characteristics (and the interaction between the two) on senescence, and can help us to understand the mechanisms that shape the evolution of senescence. In this review, we summarize and discuss the insights gained from our comprehensive long-term individual-based study of the Seychelles warbler ( Acrocephalus sechellensis ). This species provides an excellent model system in which to investigate the evolution of senescence in the wild. We found that Seychelles warblers show senescent declines in survival and reproduction, and discuss how individual characteristics (body condition, body size) and environmental effects (low- versus high-quality environments) may affect the onset and rate of senescence. Further, we highlight the evidence for trade-offs between early-life investment and senescence. We describe how key cellular and physiological processes (oxidative stress and telomere shortening) underpinning senescence are affected by individual and environmental characteristics in the Seychelles warbler (e.g. food availability, reproductive investment, disease) and we discuss how such physiological variation may mediate the relationship between environmental characteristics and senescence. Based on our work using Seychelles warblers as a model system, we show how insights from long-term studies of wild animals may help unravel the causes of the remarkable variation in senescence observed in natural systems, and highlight areas for promising future research. [ABSTRACT FROM AUTHOR]