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Whether an individual disperses or remains site‐faithful between breeding seasons can have important impacts on individual fitness and population dynamics. While several studies have identified factors influencing the probability of breeding dispersal, the consequences of dispersal are much less certain, particularly over an individual’s lifetime. Here, we use 81 cases (13 paired and 55 single dispersals) of breeding dispersal across 55 yr of breeding and re‐sighting data from an individually marked population of Canada jays (Perisoreus canadensis) at the southern edge of their range in Algonquin Provincial Park, Ontario to determine both the short‐ (year after dispersal) and long‐term (lifetime) consequences of breeding dispersal. In the year following dispersal, adults had larger brood sizes and higher nest success compared to the year prior to dispersal. However, when adults dispersed during the fall/winter, they had significantly later lay dates and lower rates of nest success than adults that dispersed during the summer. Additionally, most breeders dispersed to territories of higher quality and individuals that dispersed to a territory of lesser quality experienced lower rates of nest success. Importantly, individuals that dispersed at least once in their lifetime produced an average of 2.7 more young and recruited an average of 0.9 more juveniles into the population compared to individuals that remained site‐faithful. Our study provides rare evidence of both the short‐ and long‐term benefits of breeding dispersal in Canada jays and demonstrates how the timing of dispersal can also have consequences for individual reproductive performance.

Changes in the frequency of extreme climatic events (ECEs) can have profound impacts on individual fitness by degrading habitat quality. Organisms may respond to such changes through habitat selection, favouring those areas less affected by ECEs; however, documenting habitat selection in response to ECEs is difficult in the wild due to the rarity of such events and the long-term biological data required. Sea level rise and changing weather patterns over the past decades have led to an increase in the frequency of coastal flooding events, with serious consequences for ground nesting shorebirds. Shorebirds therefore present a useful natural study system to understand habitat selection as a response to ECEs. We used a 32-year study of the Eurasian oystercatcher (Haematopus ostralegus) to investigate whether habitat selection can lead to an increase in nest elevation and minimize the impacts of coastal flooding. The mean nest elevation of H. ostralegus has increased during the last three decades. We hypothesized that this change has been driven by changes in H. ostralegus territory settlement patterns over time. We compared various possible habitat selection cues to understand what information H. ostralegus might use to inform territory settlement. There was a clear relationship between elevation and territory settlement in H. ostralegus. In early years, settlements were more likely at low elevations but in more recent years the likelihood of settlement was similar between high and low elevation areas. Territory settlement was associated with conspecific fledgling output and conspecific density. Settlement was more likely in areas of high density and areas with high fledgling output. This study shows that habitat selection can minimize the effects of increasingly frequent ECEs. However, it seems unlikely that the changes we observe will fully alleviate the consequences of anthropogenic climate change. Rates of nest elevation increase were insufficient to track current increases in maximum high tide (0.5 vs. 0.8 cm/year). Furthermore, habitat selection cues that rely on information from previous breeding seasons (e.g. conspecific fledgling output) may become ineffective as ECEs become more frequent and environmental predictability is diminished.

1. Understanding individual-level variation in response to the environment is fundamental to understanding life-history evolution and population dynamics. Telomeres, the protective caps at the ends of chromosomes, shorten in response to oxidative stress, and telomere shortening is correlated with reduced survival and life span. Investigating telomere dynamics may help us quantify individual variation in the costs experienced from social and ecological factors, and enhance our understanding of the dynamics of natural populations.2. Here, we study spatio-temporal variation in lifelong telomere dynamics in the Seychelles warbler, Acrocephalus sechellensis. We combine long-term life history and ecological data with a large longitudinal dataset of mean telomere lengths, consisting of 1,808 samples from 22 cohorts born between 1993 and 2014. We provide a detailed analysis of how telomere dynamics vary over individual life spans and cohorts, and with spatio-temporal variation in the social and ecological environment.3. We found that telomere length decreases with cross-sectional and longitudinal measures of age, and most rapidly very early in life. However, both cross-sectional and longitudinal data suggested that against this overall pattern of shortening, bouts of telomere length increase occur in some individuals. Using a large number of repeated measurements we show statistically that these increases are unlikely to be explained solely by qPCR measurement error.4. Telomere length varied markedly among cohorts. Telomere length was positively associated with temporal variation in island-wide insect abundancea key resource for the insectivorous Seychelles warblersuggesting that the costs associated with living in harsher environments can be studied by investigating telomere dynamics. We also found evidence for sex-specific relationships between telomeres and tarsus length, potentially reflecting differential costs of growth.5. Our long-term data show that in a natural population, telomere dynamics vary in a complex manner over individual life spans, and across space and time. Variance in telomere dynamics among individuals is the product of a wide array of genetic, parental and environmental factors. Explaining this variation more fully will require the integration of comprehensive long-term ecological and genetic data from multiple populations and species.

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.

Why sexually mature individuals stay in groups as nonreproductive subordinates is central to the evolution of sociality and cooperative breeding. To understand such delayed dispersal, its costs and benefits need to be compared with those of permanently leaving to float through the population. However, comprehensive comparisons, especially regarding differences in future breeding opportunities, are rare. Moreover, extraterritorial prospecting by philopatric individuals has generally been ignored, even though the factors underlying this route to independent breeding may differ from those of strict philopatry or floating. We use a comprehensive predictive framework to explore how various costs, benefits and intrinsic, environmental and social factors explain philopatry, prospecting, and floating in Seychelles warblers (Acrocephalus sechellensis). Not only floaters more likely obtained an independent breeding position before the next season than strictly philopatric individuals, but also suffered higher mortality. Prospecting yielded similar benefits to floating but lower mortality costs, suggesting that it is overall more beneficial than floating and strict philopatry. While prospecting is probably individual-driven, although limited by resource availability, floating likely results from eviction by unrelated breeders. Such differences in proximate and ultimate factors underlying each route to independent breeding highlight the need for simultaneous consideration when studying the evolution of delayed dispersal.

It is well understood that females may gain direct benefits from breeding with attractive males. However, the direct fitness effects of mate-choice are rarely considered with respect to mating between different species (hybridization), a field dominated by discussion of indirect costs of producing unfit hybrid offspring. Hybridizing females may also gain by the types of direct benefits that are important for intraspecific mate choice, and in addition may have access to certain benefits that are restricted to mating with males of an ecologically diverged sister-taxon. We investigate possible direct benefits and costs female Ficedula flycatchers gain from breeding with a heterospecific male, and demonstrate that hybridizing female collared flycatchers (F. albicollis) breed in territories that do not suffer the seasonal decline in habitat quality experienced by females breeding with conspecifics. We exclude the hypotheses that heterospecific males provide alternative food-types or assume a greater amount of the parental workload. In fact, the diets of the two species (F. albicollis and F. hypoleuca) were highly similar, suggesting possible interspecific competition over food resources in sympatry. We discuss the implications of direct fitness effects of hybridization, and why there has been such a disparity in the attention paid to such benefits and costs with regard to intraspecific and interspecific mate-choice.

Long-term effects of conditions during early development on fitness are important for life history evolution and population ecology. Using multistrata mark-recapture models on 20 years of data, we quantified the relation between rearing conditions and lifetime fitness in a long-lived shorebird, the oystercatcher (Haematopus ostralegus). We addressed specifically the relative contribution of short- and long-term effects of rearing conditions to overall fitness consequences. Rearing conditions were defined by differences in natal habitat quality, in which there is a clear dichotomy in our study population. In the first year of life, fledglings from high-quality natal origin had a 1.3 times higher juvenile survival. Later in life (age 3-11), individuals of high-quality natal origin had a 1.6 times higher adult prebreeder survival. The most striking effect of natal habitat quality was that birds that were reared on high-quality territories had a higher probability of settling in high-quality habitat (44% vs. 6%). Lifetime reproductive success of individuals born in high-quality habitat was 2.2 times higher than that of individuals born in low-quality habitat. This difference increased further when fitness was calculated over several generations, due to a correlation between the quality of rearing conditions of parents and their offspring. Long-term effects of early conditions contributed more to overall fitness differences as short-term consequences, contrary to common conceptions on this issue. This study illustrates that investigating only short-term effects of early conditions can lead to the large underestimation of fitness consequences. We discuss how long-term consequences of early conditions may affect settlement decisions and source-sink population interactions.

The continental African Reed Warbler Acrocephalus baeticatus, like its relative the Seychelles Warbler Acrocephalus sechellensis, breeds in isolated patches. We studied the mating system of the African Reed Warbler to see whether this species, like the Seychelles Warbler, shows co-operative breeding. The African Reed Warbler is not polygynous. The majority breed monogamously (88Y0, n = 65), however in 12% of the territories three adult unrelated birds (mostly males) were observed participating in the brooding and feeding of nestlings, suggesting a polyandrous breeding system. Multilocus DNA fingerprinting revealed that the helping bird was unrelated to the pair birds. The percentage of nests with helpers was low compared to rates found in the Seychelles Warbler or Henderson Reed Warbler Acrocephalus vuughani taiti. This could be due to the scarcity of potential helpers or to the fact that, although limited, birds still had the opportunity to disperse within a meta-population structure in search of vacant territories. The presence of helpers was associated with increased hatching success due to lower predation rates, but not with increased fledging success. Another possible benefit of helping behaviour in this species could be improved predator detection and mobbing. Nest predation was high and warblers tended to build their nests in the highest, most dense reed patches available in their territory. There was no relation between habitat quality, measured as insect food availability, and the occurrence of helpers. Co-operative breeding is a reproductive system in which individuals other than the main male-female pair also exhibit parent-like (‘helping’) behaviour toward young of a single nest or brood. Numerous variations have been identified including helping at the nest by non-breeding offspring that delay dispersal and remain with their parents, and various forms of cooperative polygamy or plural breeding in which more than a single male or female share breeding status within the same social unit. Helping generally consists of feeding nestlings or fledglings, but can also include incubation and nest or territorial defence (Brown 1987). Co-operative breeding, where more birds than just the breeding pair raise the young, occurs in about

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.

1. Oystercatchers that breed on the saltmarsh of the Frisian island of Schiermon-nikoog occupy two different types of territory. In 'resident' territories the nesting area and the feeding area are adjacent. In 'leapfrog' territories the nesting area and the feeding area are separated by distances of 200-500 m.2. Between 1984 and 1989, residents fledged on average 3.5 times as many chicks per year as leapfrogs.3. The discrepancy in fledgling production arose primarily through increased mortality of leapfrog chicks due to starvation. This was indicated by low body weights of leapfrog chicks found dead and successful enhancement of leapfrog chick growth through supplementary food. In addition, the presence of a sibling reduced the probability of fledging for leapfrog chicks, but not for resident chicks, providing further evidence that leapfrog chicks competed for a limited food supply.4. Leapfrog parents fledged fewer chicks, not because of poor feeding conditions in their feeding territory (intake rates of leapfrogs exceeded those of residents), but because they failed to transport a sufficient amount of food to the chicks. To supply their chicks with the same amount of food as did residents, leapfrog parents should have spent c. 4000 s per low water period in transport flights: no leapfrog parent ever reached this level of effort.5. Our data show that it is not correct to equate parental effort to the number of chicks raised, as territories of different quality require different levels of effort for successful reproduction.