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We explored the potential influence of temperature and precipitation on the abundance of two nest-dwelling ectoparasites (blowflies and mites) of Eurasian blue tits (Cyanistes caeruleus) during a period of 21 years and compared the results with those of a shorter period. The abundance of blowflies was negatively related to precipitation, which could prevent flies from locating their host, and laying date. In addition, blowflies were positively related to brood size (more food implies more parasites) and the interaction between precipitation and temperature. The highest abundances of blowfly pupae were attained in conditions of increasing precipitation and decreasing temperature, which should be more common at the beginning of the bird breeding season. Mites were significantly and positively related to laying date and the interaction between average precipitation and temperature but only for the larger dataset. Higher abundances of mites were related to intermediate values of temperature and precipitations, conditions that are found at the end of the breeding season. These results imply that optimal conditions for both parasites differ, with blowflies preferring earlier breeders and colder and more humid conditions than mites. Thus, the effects of the climatic conditions studied on parasite abundances are non-monotonic and can vary with years and parasite species. Finally, the fact that average temperature and precipitation decreases across the years of study is probably due to the advancement in Eurasian blue tit laying date because we calculated those variables for the period of birds' reproduction. This earlier nesting does not affect parasite abundance.

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Climate models, and empirical observations, suggest that anthropogenic climate change is leading to changes in the occurrence and severity of extreme climatic events (ECEs). Effects of changes in mean climate on phenology, movement, and demography in animal and plant populations are well documented. In contrast, work exploring the impacts of ECEs on natural populations is less common, at least partially due to the challenges of obtaining sufficient data to study such rare events. Here, we assess the effect of changes in ECE patterns in a long-term study of great tits, near Oxford, over a 56-year period between 1965 and 2020. We document marked changes in the frequency of temperature ECEs, with cold ECEs being twice as frequent in the 1960s than at present, and hot ECEs being ~three times more frequent between 2010 and 2020 than in the 1960s. While the effect of single ECEs was generally quite small, we show that increased exposure to ECEs often reduces reproductive output, and that in some cases the effect of different types of ECE is synergistic. We further show that long-term temporal changes in phenology, resulting from phenotypic plasticity, lead to an elevated risk of exposure to low temperature ECEs early in reproduction, and hence suggest that changes in ECE exposure may act as a cost of plasticity. Overall, our analyses reveal a complex set of risks of exposure and effects as ECE patterns change and highlight the importance of considering responses to changes in both mean climate and extreme events. Patterns in exposure and effects of ECEs on natural populations remain underexplored and continued work will be vital to establish the impacts of ECEs on populations in a changing climate.

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Increasingly warmer springs have caused phenological shifts in both plants and animals. In birds, it is well established that mean laying date has advanced to match the earlier food peak. We know less about changes in the distribution of egg-laying dates within a population and the environmental variables that determine this variation. This could be an important component of how populations respond to climate change. We, therefore, used laying date and environmental data from 39 years (1983-2021) to determine how climate change affected laying date variation in blue tits (Cyanistes caeruleus) and marsh tits (Poecile palustris), two sympatric passerines with different life histories. Both species advanced mean laying date (0.19-0.24 days per year) and mean laying date showed a negative relationship with maximum spring temperature in both blue and marsh tits. In springs with no clear temperature increase during the critical time window (the time-window in which mean laying date was most sensitive to temperature) start of breeding in blue tits was distributed over a longer part of the season. However, there was no such pattern in marsh tits. Our findings suggest that temperature change, and not necessarily absolute temperature, can shape the variation in breeding phenology in a species-specific manner, possibly linked to variation in life-history strategies. This is an important consideration when predicting how climate change affects timing of breeding within a population.

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Background: Examining direct and indirect effects on reproduction at multiple scales allows for a broad understanding of species' resilience to environmental change. We examine how the fecundity of the mountain chickadee (Poecile gambeli), a secondary cavity-nesting, insectivorous bird, varied in relation to factors at three scales: regional weather conditions, regional- and site-level food availability, site-level community dynamics, and nest-level cavity characteristics. We hypothesized that earlier laying dates and higher fecundity (clutch size, nest survival, brood size) would be associated with milder climatic conditions, increased food from insect outbreaks, lower densities of conspecifics and nest predators (red squirrel; Tamiasciurus hudsonicus), and safer (smaller, higher) cavities. Methods: We collected data on laying date, clutch size, brood size, nest fate (success/failure), and cavity characteristics from 513 mountain chickadee nests in tree cavities in temperate mixed coniferous-broadleaf forest in interior British Columbia, Canada, from 2000 to 2011. We surveyed annual abundances of mountain chickadees and squirrels using repeated point counts, and mountain pine beetle (Dendroctonus ponderosae) and lepidopteran defoliators by monitoring host trees and by using regional-scale aerial overview forest insect survey data. We used weather data (temperature, rain, snow) from a local Environment and Climate Change Canada weather station. We modeled laying date, clutch size, daily nest survival, and brood size as a function of predictors at regional-, site-, and nest-scales. Results and Conclusions: Measures of fecundity varied dramatically across years and spatial scales. At the regional (study-wide) scale, chickadees laid earlier and larger first clutches in warmer springs with minimal storms, and daily nest survival (DSR) increased with a 2-year lag in growing season temperature. Despite a doubling of mountain chickadee density that roughly accompanied the outbreaks of mountain pine beetle and lepidopteran defoliators, we found little evidence at the site scale that fecundity was influenced by insect availability, conspecific density, or predator density. At the nest scale, DSR and brood size increased with clutch size but DSR declined with nest cavity size indicating a positive reproductive effect of small-bodied cavity excavators. Double-brooding, rare in chickadees, occurred frequently in 2005 and 2007, coinciding with early breeding, high food availability from insect outbreaks, and warm spring temperatures with 0-1 spring storms. Our results support the idea that fecundity in secondary cavity-nesting species is impacted directly and indirectly by weather, and indirectly through changes in community dynamics (via cavity resource supply). We stress the importance of adopting holistic, community-level study frameworks to refine our understanding of fecundity in opportunistic and climate-sensitive species in future.

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Knowledge of animal dispersal patterns is of great importance for the conservation and maintenance of natural populations. We here analyze juvenile dispersal of the poorly studied Black-shouldered Kite (Elanus caeruleus) monitored in southwestern Spain in an ongoing long-term study initiated in 2003. The European population of Black-shouldered kites is thought to be a recent one funded by colonizing African birds, as no kites have been found in the European fossil record, and the breeding population has progressively expanded to the North in the late 20th and 21st centuries. We obtained information on movements behavior during dispersal from 47 juveniles Kites after marking 384 nestlings with wing tags and three nestlings with radio transmitter. We have tested two competing hypotheses (i.e., the Resources Competition Hypothesis and the Wandering Hypothesis (WH)) that may explain the leptokurtic distribution of the natal dispersal distance in Elanus. After independence, juvenile females dispersed farther from the natal areas than males, as is common in birds. On average, males and females dispersed from their natal areas over 9 (i.e., 26.15 km) and 15 (i.e., 43.79 km) breeding territories, respectively. A male and two females dispersed further than 100 km from their natal nest. Our results indicated some evidence supporting the competition-for-resources hypotheses since nestlings hatched from high quality territories stayed closer from natal areas than nestlings hatched from low quality territories and also nestlings hatched first within the brood also tend to recruit closer to their natal area than later hatched nestlings which tend to disperse further away from their natal area. The information provided by these crucial demographic parameters will be used for the elaboration of future conservation plans for the management of this colonizing species in Europe.

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In recent decades, industrial emissions have been reduced in many countries, which provides an opportunity for the recovery of polluted ecosystems. However, our knowledge of the rate and factors facilitating the recovery of local bird populations after pollution abatement is incomplete. Long-term (1989-2021) annual observations on nest-box populations of a passerine bird, Ficedula hypoleuca, were used to analyze temporal dynamics of breeding parameters following a 50-fold reduction of industrial emissions from the Middle Ural copper smelter (MUCS) according to pollution zone, habitat, air temperature, and breeding density. In the heavily polluted (impact) zone (1-2 km of MUCS), egg and fledgling production were strongly impaired compared to the moderately polluted (buffer zone, 4-8 km of MUCS) and unpolluted control zone (16-27 km of MUCS). During the study period, the laying date advanced along with increasing spring air temperatures. The clutch size increased in the impact zone by 26%, in the buffer zone by 10%, and in control by 5%. The number of fledglings increased in the impact zone by 102% and the buffer zone by 17%. In the most recent year (2021), mean laying date, clutch size, fledgling production, and the frequency of nests with unhatched eggs in the impact zone did not reach the control level, whereas the frequency of nests with perished chicks did not differ among zones. Breeding parameters of birds in the impact zone improved slowly, likely due to the slow recovery of habitats. We conclude that bird reproduction may require many decades to recover fully in the heavily polluted zone.

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Earlier phenology induced by climate change, such as the passerines' breeding time, is observed in many natural populations. Understanding the nature of such changes is key to predict the responses of wild populations to climate change. Genetic changes have been rarely investigated for laying date, though it has been shown to be heritable and under directional selection, suggesting that the trait could evolve. In a Corsican blue tit population, the birds' laying date has significantly advanced over 40 years, and we here determine whether this response is of plastic or evolutionary origin, by comparing the predictions of the breeder's and the Robertson-Price (STS) equations, to the observed genetic changes. We compare the results obtained for two fitness proxies (fledgling and recruitment success), using models accounting for their zero inflation. Because the trait appears heritable and under directional selection, the breeder's equation predicts that genetic changes could drive a significant part of the phenological change observed. We, however, found that fitness proxies and laying date are not genetically correlated. The STS, therefore, predicts no evolution of the breeding time, predicting correctly the absence of trend in breeding values. Our results also emphasize that when investigating selection on a plastic trait under fluctuating selection, part of the fitness-trait phenotypic covariance can be due to within individual covariance. In the case of repeated measurements, splitting within and between individual covariance can shift our perspective on the actual intensity of selection over multiple selection episodes, shedding light on the potential for the trait to evolve.

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Nest building can represent an energetically costly activity for a variety of animal taxa. Besides, the determinants of within-species variation in the design of nests, notably with respect to natural and sexual selection, are still insufficiently documented. Based on an observational study, we examined the influence of nesting conditions (nesting-support quality, colony, laying date, and year) on male-built nest volume and also its potential role as a postmating sexually selected display in the whiskered tern Chlidonias hybrida. This tern species is a monogamous colonial bird with obligate biparental care breeding on aquatic vegetation. Hence, large nesting platforms are expected to be a selective advantage because they would better withstand adverse environmental conditions and provide a secure structure for eggs. Nest size may also serve as a postmating sexual trait, and variation in egg production would be positively associated with nest size. We found that nest volume was adjusted to different environmental cues. A positive relationship was found between nest volume and nesting-support quality, indicating that the leaf density of white waterlily is essential for nest stability. Variation in nest volume was not correlated to colony size but varied among colonies and years. Male-built nest volume was also positively associated with mean egg volume per clutch but not with clutch size. The fitness consequences of building a large nest are yet to be studied, and additional investigations are recommended to better understand whether the activity of males early during breeding season (e.g., nest building and courtship feeding performance) really serves as postmating sexually selected signals.

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The origin of natural selection is linked to environmental heterogeneity, which influences variation in relative fitness among phenotypes. However, individuals in wild populations are exposed to a plethora of biotic and abiotic environmental factors. Surprisingly, the relative influence of multiple environmental conditions on the relative fitness of phenotypes has rarely been tested in wild populations. Identifying the main selection agent(s) is crucial when the target phenotype is tightly linked to reproduction and when temporal variation in selection is expected to affect evolutionary responses. By using individual-based data from a 29-year study of a short-lived migratory songbird, the pied flycatcher (Ficedula hypoleuca), we studied the relative influence of 28 temperature- and precipitation-based factors at local and global scales on selection on breeding time (egg laying) at the phenotypic level. Selection, estimated using the number of recruits as a proxy for fitness, penalized late breeders. Minimum temperatures in April and May were the environmental drivers that best explained selection on laying date. In particular, there was negative directional selection on laying date mediated by minimum temperature in April, being strongest in cold years. In addition, nonlinear selection on laying date was influenced by minimum temperatures in May, with selection on laying date changing from null to negative as the breeding season progressed. The intensity of selection on late breeders increased when minimum temperatures in May were highest. Our results illustrate the complex influence of environmental factors on selection on laying date in wild bird populations. Despite minimum temperature in April being the only variable that changed over time, its increase did not induce a shift in laying date in the population. In this songbird population, stabilizing selection has led to a three-decade stasis in breeding time. We suggest that variation in the effects of multiple climatic variables on selection may constrain phenotypic change.

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Temperatures in mountain areas are increasing at a higher rate than the Northern Hemisphere land average, but how fauna may respond, in particular in terms of phenology, remains poorly understood. The aim of this study was to assess how elevation could modify the relationships between climate variability (air temperature and snow melt-out date), the timing of plant phenology and egg-laying date of the coal tit (Periparus ater). We collected 9 years (2011-2019) of data on egg-laying date, spring air temperature, snow melt-out date, and larch budburst date at two elevations (~1,300 m and ~1,900 m asl) on a slope located in the Mont-Blanc Massif in the French Alps. We found that at low elevation, larch budburst date had a direct influence on egg-laying date, while at high-altitude snow melt-out date was the limiting factor. At both elevations, air temperature had a similar effect on egg-laying date, but was a poorer predictor than larch budburst or snowmelt date. Our results shed light on proximate drivers of breeding phenology responses to interannual climate variability in mountain areas and suggest that factors directly influencing species phenology vary at different elevations. Predicting the future responses of species in a climate change context will require testing the transferability of models and accounting for nonstationary relationships between environmental predictors and the timing of phenological events.

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AbstractPhenological traits, such as the timing of reproduction, are often influenced by social interactions between paired individuals. Such partner effects may occur when pair members affect each other's prebreeding environment. Partner effects can be environmentally and/or genetically determined, and quantifying direct and indirect genetic effects is important for understanding the evolutionary dynamics of phenological traits. Here, using 26 years of data from a pedigreed population of a migratory seabird, the common tern (Sterna hirundo), we investigate male and female effects on female laying date. We find that female laying date harbors both genetic and environmental variation and is additionally influenced by the environmental and, to a lesser extent, genetic component of its mate. We demonstrate this partner effect to be largely explained by male arrival date. Interestingly, analyses of mating patterns with respect to arrival date show mating to be strongly assortative, and using simulations we show that assortative mating leads to overestimation of partner effects. Our study provides evidence for partner effects on breeding phenology in a long-distance migrant while uncovering the potential causal pathways underlying the observed effects and raising awareness for confounding effects resulting from assortative mating or other common environmental effects.

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Life-history traits are often plastic in response to environmental factors such as temperature or precipitation, and they also vary with age in many species. Trait variation during the lifetime could thus be partly driven by age-dependent plasticity in these traits. We study whether plasticity of a phenological trait-the egg-laying date-with respect to spring temperature, varies with age, and explore whether this variation relates to changes in breeding success throughout the life cycle. We use data from a four-decade long-term monitoring of a wild population of blue tits in Corsica, to estimate age-dependent plasticity of reproductive phenology and annual reproductive success. We show that both laying date plasticity and annual reproductive success vary with age: young and old females are less plastic, and fledge fewer offspring, than middle-age females. Furthermore, in contrast to young and prime-age females, in old females fledging success does not depend on laying date. Phenological plasticity is a major mechanism for coping with rapid environmental variation. Our results suggest that understanding its role in adaptation to climate change and population persistence requires integrating the age structure of the population.

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Within-population variation in the traits underpinning reproductive output has long been of central interest to biologists. Since they are strongly linked to lifetime reproductive success, these traits are expected to be subject to strong selection and, if heritable, to evolve. Despite the formation of durable pair bonds in many animal taxa, reproductive traits are often regarded as female-specific, and estimates of quantitative genetic variation seldom consider a potential role for heritable male effects. Yet reliable estimates of such social genetic effects are important since they influence the amount of heritable variation available to selection. Based on a 52-year study of a nestbox-breeding great tit (Parus major) population, we apply "extended" bivariate animal models in which the heritable effects of both sexes are modeled to assess the extent to which males contribute to heritable variation in seasonal reproductive timing (egg laying date) and clutch size, while accommodating the covariance between the two traits. Our analyses show that reproductive timing is a jointly expressed trait in this species, with (positively covarying) heritable variation for laydate being expressed in both members of a breeding pair, such that the total heritable variance is 50% larger than estimated by traditional models. This result was robust to explicit consideration of a potential male-biased environmental confound arising through sexually dimorphic dispersal. In contrast to laydate, males' contribution to heritable variation in clutch size was limited. Our study thus highlights the contrasting extent of social determination for two major components of annual reproductive success, and emphasizes the need to consider the social context of what are often considered individual-level traits.

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It has been suggested that extreme weather events may be treated as natural experiments that may unravel the mechanisms by which birds adjust their phenology and breeding parameters to environmental variability. In 2017, a sudden and heavy drop of temperatures for several days affected many European bird populations. This event occurred during the laying-early incubation period in the great tit (Parus major) population in central Poland, causing a large delay in hatching and had sustained reproductive consequences. This cold snap occurring once breeding activity had already started in 2017 was followed by the warm and invariable breeding season of 2018. This natural experiment had an essential influence on great tit reproductive parameters. We found a significant difference in hatching date, number of fledglings, hatching success, and fledging success between 2017 and 2018. In 2017, there were about two fledglings per nest fewer than in 2018. Fledging success was positively associated with hatching delay in 2017, while the relation was negative in 2018. Hatching success differed significantly between both years, being higher in 2018. Mean level of hemoglobin (used as index of body, physiological condition) in the blood of nestling great tits was higher in 2017 than in 2018. We argue that the moment of hatching may be (at least in some years) more tightly related to the moment of maximum food demand of tit nestlings than the traditionally used laying date. Also in extreme years, phenotypic plasticity of hatching delays may be insufficient to adjust the timing of breeding to the upcoming extreme weather events. Further examining its limits may be an important goal for future research.

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Phenological shifts of plants and animals due to climate change can vary among regions and species, requiring study of local ecosystems to understand specific impacts. The reproductive timing of insectivorous songbirds in temperate forests is tightly synchronized with peak prey abundance, and thus they can be susceptible to such shift in timing. We aimed to investigate the effect of future climate change on the egg-laying phenology of the Varied Tit (Sittiparus various), which is common and widely distributed in South Korean forests. We developed the predictive model by investigating their egg-laying dates in response to spring temperatures along geographical gradients, and our model indicated that the tits lay eggs earlier when the average of daily mean and daily maximum temperatures rise. We predicted future shifts in egg-laying dates based on the most recent climate change model published by the Intergovernmental Panel on Climate Change (IPCC), under a scenario with no climate change mitigation and under a scenario with moderate mitigation. Under this outcome, this species might be unable to adapt to rapid climate change due to asynchrony with prey species during the reproductive period. If no mitigation is undertaken, our model predicts that egg-laying dates will be advanced by more than 10 days compared to the present in 83.58% of South Korea. However, even moderate mitigation will arrest this phenomenon and maintain present egg-laying dates. These results demonstrate the first quantitative assessment for the effect of warming temperatures on the phenological response of insectivorous songbirds in South Korea.

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Establishing the cues or constraints that influence avian timing of breeding is the key to accurate prediction of future phenology. This study aims to identify the aspects of the environment that predict the timing of two measures of breeding phenology (nest initiation and egg laying date) in an insectivorous woodland passerine, the blue tit (Cyanistes caeruleus). We analyse data collected from a 220 km, 40-site transect over 3 years and consider spring temperatures, tree leafing phenology, invertebrate availability and photoperiod as predictors of breeding phenology. We find that mean night-time temperature in early spring is the strongest predictor of both nest initiation and lay date and suggest this finding is most consistent with temperature acting as a constraint on breeding activity. Birch budburst phenology significantly predicts lay date additionally to temperature, either as a direct cue or indirectly via a correlated variable. We use cross-validation to show that our model accurately predicts lay date in two further years and find that similar variables predict lay date well across the UK national nest record scheme. This work refines our understanding of the principal factors influencing the timing of tit reproductive phenology and suggests that temperature may have both a direct and indirect effect.

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Phenotypic plasticity is a major mechanism of response to global change. However, current plastic responses will only remain adaptive under future conditions if informative environmental cues are still available. We briefly summarize current knowledge of the evolutionary origin and mechanistic underpinnings of environmental cues for phenotypic plasticity, before highlighting the potentially complex effects of global change on cue availability and reliability. We then illustrate some of these aspects with a case study, comparing plasticity of blue tit breeding phenology in two contrasted habitats: evergreen and deciduous forests. Using long-term datasets, we investigate the climatic factors linked to the breeding phenology of the birds and their main food source. Blue tits occupying different habitats differ extensively in the cues affecting laying date plasticity, as well as in the reliability of these cues as predictors of the putative driver of selective pressure, the date of caterpillar peak. The temporal trend for earlier laying date, detected only in the evergreen populations, is explained by increased temperature during their cue windows. Our results highlight the importance of integrating ecological mechanisms shaping variation in plasticity if we are to understand how global change will affect plasticity and its consequences for population biology. This article is part of the theme issue 'The role of plasticity in phenotypic adaptation to rapid environmental change'.

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In seasonal environments, timing of reproduction is a trait with important fitness consequences, but we know little about the molecular mechanisms that underlie the variation in this trait. Recently, several studies put forward DNA methylation as a mechanism regulating seasonal timing of reproduction in both plants and animals. To understand the involvement of DNA methylation in seasonal timing of reproduction, it is necessary to examine within-individual temporal changes in DNA methylation, but such studies are very rare. Here, we use a temporal sampling approach to examine changes in DNA methylation throughout the breeding season in female great tits (Parus major) that were artificially selected for early timing of breeding. These females were housed in climate-controlled aviaries and subjected to two contrasting temperature treatments. Reduced representation bisulfite sequencing on red blood cell derived DNA showed genome-wide temporal changes in more than 40,000 out of the 522,643 CpG sites examined. Although most of these changes were relatively small (mean within-individual change of 6%), the sites that showed a temporal and treatment-specific response in DNA methylation are candidate sites of interest for future studies trying to understand the link between DNA methylation patterns and timing of reproduction.

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Many bird species have been observed shifting their laying date to earlier in the year in response to climate change. However, the vast majority of these studies were performed on non-threatened species, less impacted by reduced genetic diversity (which is expected to limit evolutionary response) as a consequence of genetic bottlenecks, drift and population isolation. Here, we study the relationship between lay date and fitness, as well as its genetic basis, to understand the evolutionary constraints on phenology faced by threatened species using a recently reintroduced population of the endangered New Zealand passerine, the hihi (Notiomystis cincta). A large discrepancy between the optimal laying date and the mode of laying date creates a strong selection differential of -11.24. The impact of this discrepancy on fitness is principally mediated through survival of offspring from hatchling to fledgling. This discrepancy does not seem to arise from a difference in female quality or a trade-off with lifetime breeding success. We find that start of breeding season depends on female age and average temperature prior to the breeding season. Laying date is not found to be significantly heritable. Overall, our research suggests that this discrepancy is a burden on hihi fitness, which will not be resolved through evolution or phenotypic plasticity. More generally, these results show that threatened species introduced to restored habitats might lack adaptive potential and plasticity to adjust their phenology to their new environment. This constraint is also likely to limit their ability to face future challenges, including climate change.

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In changing environments, phenotypic traits are shaped by numerous agents of selection. The optimal phenotypic value maximizing the fitness of an individual thus varies through time and space with various environmental covariates. Selection may differ between different life-cycle stages and act on correlated traits inducing changes in the distribution of several traits simultaneously. Despite increasing interests in environmental sensitivity of phenotypic selection, estimating varying selective optima on various traits throughout the life cycle, while considering (a)biotic factors as potential selective agents has remained challenging. Here, we provide a statistical model to measure varying selective optima from longitudinal data. We apply our approach to analyze environmental sensitivity of phenotypic selection on egg-laying date and clutch size throughout the life cycle of a white-throated dipper population. We show the presence of a joint optimal phenotype that varies over the 35-year period, being dependent on altitude and temperature. We also find that optimal laying date is density-dependent, with high population density favoring earlier laying dates. By providing a flexible approach, widely applicable to free-ranging populations for which long-term data on individual phenotypes, fitness, and environmental factors are available, our study improves the understanding of phenotypic selection in varying environments.

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