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Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young., This meta-analysis was financed by the grant of the Polish National Science Centre (Narodowe Centrum Nauki) (no. 2017/27/B/NZ8/00465) awarded to Lucyna Hałupka.

International audience; Climate warming is considered to be among the most serious of anthropogenic stresses to the environment, because it not only has direct effects on biodiversity, but it also exacerbates the harmful effects of other human-mediated threats. The associated consequences are potentially severe, particularly in terms of threats to species preservation, as well as in the preservation of an array of ecosystem services provided by biodiversity. Among the most affected groups of animals are insects—central components of many ecosystems—for which climate change has pervasive effects from individuals to communities. In this contribution to the scientists' warning series, we summarize the effect of the gradual global surface temperature increase on insects, in terms of physiology, behavior, phenology, distribution, and species interactions, as well as the effect of increased frequency and duration of extreme events such as hot and cold spells, fires, droughts, and floods on these parameters. We warn that, if no action is taken to better understand and reduce the action of climate change on insects, we will drastically reduce our ability to build a sustainable future based on healthy, functional ecosystems. We discuss perspectives on relevant ways to conserve insects in the face of climate change, and we offer several key recommendations on management approaches that can be adopted, on policies that should be pursued, and on the involvement of the general public in the protection effort.

Climate change may influence animal population dynamics through reproduction and mortality. However, attributing changes in mortality to specific climate variables is challenging because the exact time of death is usually unknown in the wild. Here, we investigated climate effects on adult mortality in Australian superb fairy-wrens ( Malurus cyaneus ). Over a 27-year period, mortality outside the breeding season nearly doubled. This nonbreeding season mortality increased with lower minimum (night-time) and higher maximum (day-time) winter temperatures and with higher summer heat wave intensity. Fine-scale analysis showed that higher mortality in a given week was associated with higher maxima 2 weeks prior and lower minima in the current fortnight, indicating costs of temperature drops. Increases in summer heat waves and in winter maximum temperatures collectively explained 62.6% of the increase in mortality over the study period. Our results suggest that warming climate in both summer and winter can adversely affect survival, with potentially substantial population consequences.

Body condition is an important concept in behaviour, evolution and conservation, commonly used as a proxy of an individual's performance, for example in the assessment of environmental impacts. Although body condition potentially encompasses a wide range of health state dimensions (nutritional, immune or hormonal status), in practice most studies operationalize body condition using a single (univariate) measure, such as fat storage. One reason for excluding additional axes of variation may be that multivariate descriptors of body condition impose statistical and analytical challenges. Structural equation modelling (SEM) is used in many fields to study questions relating multidimensional concepts, and we here explain how SEM is a useful analytical tool to describe the multivariate nature of body condition. In this 'Research Methods Guide' paper, we show how SEM can be used to resolve different challenges in analysing the multivariate nature of body condition, such as (a) variable reduction and conceptualization, (b) specifying the relationship of condition to performance metrics, (c) comparing competing causal hypothesis and (d) including many pathways in a single model to avoid stepwise modelling approaches. We illustrated the use of SEM on a real-world case study and provided R-code of worked examples as a learning tool. We compared the predictive power of SEM with conventional statistical approaches that integrate multiple variables into one condition variable: multiple regression and principal component analyses. We show that model performance on our dataset is higher when using SEM and led to more accurate and precise estimates compared to conventional approaches. We encourage researchers to consider SEM as a flexible framework to describe the multivariate nature of body condition and thus understand how it affects biological processes, thereby improving the value of body condition proxies for predicting organismal performance. Finally, we highlight that it can be useful for other multidimensional ecological concepts as well, such as immunocompetence, oxidative stress and environmental conditions.

Individual variation in disturbance vulnerability (i.e. the likelihood that disturbance negatively affects an individual's fitness) can affect how disturbance impacts animal populations, as even at low disturbance levels some individuals could be severely affected and die. Individual variation in vulnerability can arise due to different responses to disturbance. We propose a new hypothesis that even when individuals respond similarly to disturbance, time-limited individuals are more at risk that their condition deteriorates since they have limited ability to extend their foraging time to compensate for disturbance. We investigate this ‘time-limitation hypothesis' both empirically and mathematically by studying how individuals that differ in their average foraging time (presumably due to differences in foraging efficiency) are affected by disturbance. We used tracking data of 22 wintering Eurasian oystercatchers Haematopus ostralegus to compare time budgets between disturbed and undisturbed tidal periods. In three tidal periods with severe disturbance by transport airplanes, more than a third of the variation in additional flight time and foraging time loss was explained by individual differences. Inefficient individuals that foraged longer in undisturbed tidal periods experienced higher costs in disturbed tidal periods, since they lost more foraging time. We next used an analytical energy balance model to study how time-limited individuals differed in their maximum disturbance thresholds. Both our theoretical model and empirical study suggest that inefficient individuals in a time-limited environment may be unable to increase their foraging time to compensate for the effects of disturbance. Consequently, the magnitude of individual variation in time budgets strongly determines what proportion of the population is at risk that their condition deteriorates due to disturbance. Our hypothesis implies that, when assessing disturbance effects on a population, it is not only important to consider individual variation in disturbance responses, but also variation in time budgets that determine the consequences of disturbance.

Assignment of parentage with molecular markers is most difficult when the true parents have close relatives in the adult population. Here, we present an efficient solution to that problem by extending simple exclusion approaches to parentage analysis with single nucleotide polymorphic markers (SNPs). We augmented the previously published homozygote opposite test (hot), which counts mismatches due to the offspring and candidate parent having different homozygous genotypes, with an additional test. In this case, parents homozygous for the same SNP are incompatible with heterozygous offspring (i.e., “Homozygous Identical Parents, Heterozygous Offspring are Precluded”: hiphop). We tested this approach in a cooperatively breeding bird, the superb fairy-wren, Malurus cyaneus, where rates of extra-pair paternity are exceptionally high, and where paternity assignment is challenging because breeding males typically have first-order adult relatives in their neighbourhood. Combining the tests and conditioning on the maternal genotype with a set of 1376 autosomal SNPs always allowed us to distinguish a single most likely sire from his relatives, and also to identify cases where the true sire must have been unsampled. In contrast, if just the hot test was used, we failed to identify a single most-likely sire in 2.5% of cases. Resampling enabled us to create guidelines for the number of SNPs required when first-order relatives coexist in the mating pool. Our method, implemented in the R package hiphop, therefore provides unambiguous parentage assignments even in systems with complex social organisation. We also identified a suite of Z- and W-linked SNPs that always identified sex correctly.

Many wild populations are experiencing temporal changes in life-history and other phenotypic traits, and these changes are frequently assumed to be driven by climate change rather than nonclimatic drivers. However, this assumption relies on three conditions: that local climate is changing, traits are sensitive to climate variability, and other drivers are not also changing over time. Although many studies acknowledge one or more of these conditions, all three are rarely checked simultaneously. Consequently, the relative contribution of climate change to trait change, and the variation in this contribution across traits and species, remain unclear. We used long-term datasets on 60 bird species in Europe to test the three conditions in laying date, offspring number, and body condition and used a method that quantifies the contribution of warming temperatures to changes in traits relative to other effects. Across species, approximately half of the magnitude of changes in traits could be attributed to rising mean temperature, suggesting that increasing temperatures are likely the single most important contributor to temporal trends and emphasizes the impact that global warming is having on natural populations. There were also substantial nontemperature-related temporal trends (presumably due to other changes such as urbanization), which generally caused trait change in the same direction as warming. Attributing temporal trends solely to warming thus overestimates the impact of warming. Furthermore, contributions from nontemperature drivers explained most of the interspecific variation in trait changes, raising concerns about comparative studies that attribute differences in temporal trends to species differences in climate-change sensitivity.

Feeding specialization is a common cause of individual variation. Fitness payoffs of specialization vary with environmental conditions, but the underlying behavioral mechanisms are poorly understood. Such mechanistic knowledge, however, is crucial to reliably predict responses of heterogeneous populations to environmental change. We quantified spatiotemporal allocation of foraging behavior in wintering Eurasian oystercatchers (Haematopus ostralegus), a species in which feeding specialization can be inferred from bill shape. We combined global positioning system (GPS) and accelerometer data to quantify foraging time of 64 individuals for every tidal period in one or two winter seasons. Individuals varied widely in foraging time (3.7-6.5 h per tidal period) and individuals that spend more time foraging had lower inferred survival. Feeding specialization appeared a major determinant of individual variation in foraging time and its spatiotemporal allocation. Visually hunting worm specialists foraged more during day time and complemented intertidal foraging with grassland foraging when the exposure of intertidal flats was limited and nights were well illuminated. Shellfish specialists increased total foraging time in cold weather, whereas foraging time of worm specialists decreased as frosty grasslands became inaccessible. Our results imply that worm specialists may be most sensitive to cold snaps and daytime disturbance, whereas shellfish specialists are most sensitive to high water levels. These behavioral responses can be implemented in population models to predict the vulnerability of heterogeneous populations to environmental change and, thereby, provide a shortcut to long-term population studies that require fitness data across many years and conditions to make similar projections. Lay Summary: Populations consist of different types of individuals. Predicting how heterogeneous populations of long-lived species respond to environmental change is hard as it takes decades to quantify how fitness varies among individuals in different environmental conditions. We studied the behavior of GPS-tracked oystercatchers and showed how foraging time is differentially affected by environmental conditions among individuals that specialize in different prey. This knowledge can be implemented in models to predict the vulnerability of heterogeneous populations to environmental change.

AIM: In many species, density‐dependent effects on reproduction are an important driver of population dynamics. However, it is rarely considered that the direction of density dependence is expected to vary over space and time depending on anti‐predator behaviour and predator community. Aggregation may allow for effective group mobbing against avian nest predators while aggregation may also attract mammalian predators, causing negative density dependence. We aim to quantify spatial variation in the effect of conspecific breeding density on nest survival in a mobbing bird species (Eurasian oystercatcher; Haematopus ostralegus) and identify whether this variation in density dependence can be explained by the predator community. LOCATION: Country‐wide (The Netherlands). METHODS: We integrated reproductive data with breeding territory maps of Eurasian oystercatchers and occupancy maps of avian and mammalian predator species across the Netherlands for a 10‐year period. RESULTS: Spatial variation in the composition of the predator community explained the effects of neighbour density, showing decreasing nest survival when both conspecific density and mammalian dominance increased. Also, heterospecific density (from breeding godwits and lapwing) has an additional effect on the oystercatcher nest survival. Strikingly, this pattern did not extend to mammal‐free island populations. MAIN CONCLUSIONS: Our study provides evidence that both the strength and sign of density dependence can vary spatially within species, implying that it is dangerous to generalize results from a single local population to large‐scale management implications and modelling exercises. The study also suggests that conservation actions that aim to attract breeding birds should be prioritized in areas with fewer mammalian predators, but this idea requires further testing on island populations.

To understand the consequences of anthropogenic and environmental changes for wildlife populations, it is important to study how individuals differ in their sensitivity to environmental change and whether this depends on individual characteristics. An individual's reproductive performance may provide an integrative, unidimensional proxy of an individual's characteristics. In this study, we define an individual's characteristics by three such reproductive states, namely successful, failed and non-breeders in the previous year. We used a 16-year dataset of individually marked breeding Eurasian Oystercatchers Haematopus ostralegus to examine the interannual fluctuations in reproductive success and survival among breeding states, and their state-dependent sensitivity to environmental conditions. Environmental conditions included available biomass of the main prey species of breeding Oystercatchers (Ragworm, Baltic Tellin and Lugworm), tidal height, which reflects one of the main causes of nest loss (flooding), and conditions that may impact the energetic requirements during incubation, such as temperature. We also included environmental variables measured in winter, including available biomass of the main winter prey species (Blue Mussel and Common Cockle) along with factors that may affect food availability and energetic requirements for homeostasis, namely bivalve weight loss, windchill, winter severity and precipitation. Breeding birds that were successful the previous year had higher survival and were more likely to remain successful, in comparison with failed or non-breeders. The effects of environmental conditions acted in the same direction on reproductive success but had opposite effects on survival among the three breeding states, especially for windchill and Blue Mussel biomass. The contrasting state-dependent effects of the environment on survival thus averaged out when examining consequences for lifetime reproductive nest success (LRnS); instead, LRnS was largely influenced by environmental conditions acting upon reproduction. Our study indicates that an individual's previous breeding state provides an integrative measure of heterogeneity in individuals' sensitivity of reproduction and survival to environmental change. Incorporating previous breeding state as a source of individual heterogeneity in population modelling may improve predictions of future population dynamics in a rapidly changing world.

Shorebird populations, especially those feeding on shellfish, have strongly declined in recent decades and identifying the drivers of these declines is important for conservation. Changing food stocks are thought to be a key driver of these declines and may also explain why trends have not been uniform across Europe's largest estuary. We therefore investigated how winter population trends of Eurasian oystercatchers (Haematopus ostralegus) were linked to food availability in the Dutch Wadden Sea. Our analysis incorporated two spatial scales, a smaller scale focused on roost counting areas and food available to birds in these areas and a larger spatial scale of tidal basins. A novelty in our study is that we quantify the connectivity between roosting and foraging areas, identified from GPS tracking data. This allowed us to estimate food available to roosting birds and thus how food availability may explain local population trends. At the smaller spatial scale of roost counting areas, there was no clear relationship between available food and the number of roosting oystercatchers, indicating that other factors may drive population fluctuations at finer spatial scales. At the scale of tidal basins, however, there was a significant relationship between population trends and available food, especially cockle Cerastoderma edule,. Mortality and recruitment alone could not account for the large fluctuations in bird counts, suggesting that the site choice of wintering migratory oystercatchers may primarily drive these large fluctuations. Furthermore, the relationship between oystercatcher abundance and benthic food stocks, suggests winter shorebird counts could act as ecological indicators of ecosystem health, informing about the winter status of food stocks at a spatial scale of tidal basins.

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.

Changes in climate are shifting the timing of life cycle events in the natural world. Compared to northern temperate areas, these effects are relatively poorly understood in tropical and southern regions, where there is limited information on how timing of breeding and food availability are affected by climatic factors, and where patterns of breeding activity are more unpredictable within and between years. Combining a new statistical modelling approach with 5 years of continuous individual-based monitoring of a monsoonal tropical insectivorous bird, we quantified (a) the proximate climatic drivers at two trophic levels: timing of breeding and abundance of arthropod prey; (b) the effect of climate variation on reproductive output and (c) the role of individual plasticity. Rainfall was identified as the main determinant of phenology at both trophic levels. Throughout the year, likelihood of egg laying increased very rapidly in response to even small amounts of rain during the preceding 0–3 weeks. Adult body mass and male sperm storage also increased rapidly after rain, suggesting high breeding preparedness. Additionally, females were flexible, since they were more likely to nest whether their previous attempt was longer ago and unsuccessful. Arthropod abundance also increased after rainfall, but more slowly, with a peak around 10 weeks. Therefore, the peak food availability coincided with the presence of dependent fledglings. Fitness benefits of nesting after more rain appeared to be linked to offspring quantity rather than quality: nest attempts following higher rainfall produced larger clutches, but showed no improvement in nestling mass or relative fledging success. The response of clutch size to rainfall was plastic, since repeated sampling showed that individual females laid larger clutches after more rain, possibly mediated by improved body mass. Rapid, individually flexible breeding in response to rainfall and slower increase in arthropod abundance also as a response to rainfall, might buffer insectivorous species living in tropical seasonal environments from climate change-induced phenological trophic mismatches.

Background Anthropogenic disturbance can negatively affect an animal’s energy budget by evoking movement responses. Existing research focuses mainly on immediate displacement as a disturbance effect, since this can be easily observed in the field. However, effects on movement over longer timescales are poorly examined and it is largely unknown if and to what extent they reflect immediate responses. Longer-term responses could for example be larger than immediate responses if birds, after disturbance, return to the original location and thereby travel twice the immediate disturbed distance. Methods We combined GPS tracking data with observational data to quantify the effects of anthropogenic (air force and walkers) and non-anthropogenic disturbances on distances travelled by roosting Eurasian Oystercatchers (Haematopus ostralegus) during the non-breeding season. We compared immediate displacement after a disturbance with distance travelled during the entire high tide period (longer-term response), while accounting for environmental factors. Additionally, we calculated energy expenditure due to disturbance based on observed disturbance frequencies. Results Disturbance resulted in an immediate displacement response of ~ 200 m (median). Air force disturbances tended to yield larger immediate responses than walker and, especially, than non-anthropogenic disturbances. Longer-term responses and immediate responses were approximately similar, suggesting that, over longer timescales, spatial disturbance effects in the study area remain confined to immediate effects. However, disturbances were infrequent (0.17 disturbances per bird per hour) and most disturbances were of natural origin (62%). Consequently, anthropogenic disturbance of roosting oystercatchers in the study area on average costs 0.08% of the daily energy expenditure. Conclusions Our results suggest that immediate spatial responses to disturbance can be a useful proxy for spatial responses over longer timescales. Over the non-exhaustive range of conditions investigated, energetic consequences of spatial disturbance responses for an oystercatcher in the study area are marginal due to low disturbance levels.

Climate change has caused a clear and univocal trend towards advancement in spring phenology. Changes in autumn phenology are much more diverse, with advancement, delays, and ‘no change' all occurring frequently. For migratory birds, patterns in autumn migration phenology trends have been identified based on ecological and life‐history traits. Explaining interspecific variation has nevertheless been challenging, and the underlying mechanisms have remained elusive. Radar studies on non‐species‐specific autumn migration intensity have repeatedly suggested that there are strong links with weather. In long‐term species‐specific studies, the variance in autumn migration phenology explained by weather has, nevertheless, been rather low, or a relationship was even lacking entirely. We performed a spatially explicit time window analysis of weather effects on mean autumn passage of four trans‐Saharan and six intra‐European passerines to gain insights into this apparent contradiction. We analysed data from standardized daily captures at the Heligoland island constant‐effort site (Germany), in combination with gridded daily temperature, precipitation and wind data over a 55‐year period (1960–2014), across northern Europe. Weather variables at the breeding and stopover grounds explained up to 80% of the species‐specific interannual variability in autumn passage. Overall, wind conditions were most important. For intra‐European migrants, wind was even twice as important as either temperature or precipitation, and the pattern also held in terms of relative contributions of each climate variable to the temporal trends in autumn phenology. For the trans‐Saharan migrants, however, the pattern of relative trend contributions was completely reversed. Temperature and precipitation had strong trend contributions, while wind conditions had only a minor impact because they did not show any strong temporal trends. As such, understanding species‐specific effects of climate on autumn phenology not only provides unique insights into each species' ecology but also how these effects shape the observed interspecific heterogeneity in autumn phenological trends.

An increasing number of empirical studies aim to quantify individual variation in demographic parameters because these patterns are key for evolutionary and ecological processes. Advanced approaches to estimate individual heterogeneity are now using a multivariate normal distribution with correlated individual random effects to account for the latent correlations among different demographic parameters occurring within individuals. Despite the frequent use of multivariate mixed models, we lack an assessment of their reliability when applied to Bernoulli variables. Using simulations, we estimated the reliability of multivariate mixed effect models for estimating correlated fixed individual heterogeneity in demographic parameters modelled with a Bernoulli distribution. We evaluated both bias and precision of the estimates across a range of scenarios that investigate the effects of life-history strategy, levels of individual heterogeneity and presence of temporal variation and state dependence. We also compared estimates across different sampling designs to assess the importance of study duration, number of individuals monitored and detection probability. In many simulated scenarios, the estimates for the correlated random effects were biased and imprecise, which highlight the challenge in estimating correlated random effects for Bernoulli variables. The amount of fixed among-individual heterogeneity was frequently overestimated, and the absolute value of the correlation between random effects was almost always underestimated. Simulations also showed contrasting performances of mixed models depending on the scenario considered. Generally, estimation bias decreases and precision increases with slower pace of life, large fixed individual heterogeneity and large sample size. We provide guidelines for the empirical investigation of individual heterogeneity using correlated random effects according to the life-history strategy of the species, as well as, the volume and structure of the data available to the researcher. Caution is warranted when interpreting results regarding correlated individual random effects in demographic parameters modelled with a Bernoulli distribution. Because bias varies with sampling design and life history, comparisons of individual heterogeneity among species is challenging. The issue addressed here is not specific to demography, making this warning relevant for all research areas, including behavioural and evolutionary studies.

Biological processes exhibit complex temporal dependencies due to the sequential nature of allocation decisions in organisms' life cycles, feedback loops and two-way causality. Consequently, longitudinal data often contain cross-lags: the predictor variable depends on the response variable of the previous time step. Although statisticians have warned that regression models that ignore such covariate endogeneity in time series are likely to be inappropriate, this has received relatively little attention in biology. Furthermore, the resulting degree of estimation bias remains largely unexplored. We use a graphical model and numerical simulations to understand why and how regression models that ignore cross-lags can be biased, and how this bias depends on the length and number of time series. Ecological and evolutionary examples are provided to illustrate that cross-lags may be more common than is typically appreciated and that they occur in functionally different ways. We show that routinely used regression models that ignore cross-lags are asymptotically unbiased. However, this offers little relief, as for most realistically feasible lengths of time-series conventional methods are biased. Furthermore, collecting time series on multiple subjects—such as populations, groups or individuals—does not help to overcome this bias when the analysis focusses on within-subject patterns (often the pattern of interest). Simulations, a literature search and a real-world empirical example together suggest that approaches that ignore cross-lags are likely biased in the direction opposite to the sign of the cross-lag (e.g. towards detecting density dependence of vital rates and against detecting life-history trade-offs and benefits of group living). Next, we show that multivariate (e.g. structural equation) models can dynamically account for cross-lags, and simultaneously address additional bias induced by measurement error, but only if the analysis considers multiple time series. We provide guidance on how to identify a cross-lag and subsequently specify it in a multivariate model, which can be far from trivial. Our tutorials with data and R code of the worked examples provide step-by-step instructions on how to perform such analyses. Our study offers insights into situations in which cross-lags can bias analysis of ecological and evolutionary time series and suggests that adopting dynamical models can be important, as this directly affects our understanding of population regulation, the evolution of life histories and cooperation, and possibly many other topics. Determining how strong estimation bias due to ignoring covariate endogeneity has been in the ecological literature requires further study, also because it may interact with other sources of bias.

Many wild populations are showing changes in phenotypic traits. However, the common assumption that such changes are driven by climate change relies on three conditions: that local climate is changing over time, that trait(s) are sensitive to climate variability, and that other causal agents are not also changing. We used long-term datasets on 60 bird species to test these conditions and to quantify the contribution of warming temperatures to changes in three important life-history traits. Across species, approximately half of the magnitude of changes in traits could be attributed to a single temperature window, with averages of 50% for laying date, 40% for body condition and 56% for offspring numbers. Thus, although warming temperatures were a key driver of change, other unknown factors contributed substantially to temporal trends (typically reinforcing change). Further analyses showed that these non-temperature-driven contributions explained most of the inter-specific variation in trait changes.

The phenology of many species shows strong sensitivity to climate change; however, with few large scale intra-specific studies it is unclear how such sensitivity varies over a species’ range. We document large intra-specific variation in phenological sensitivity to temperature using laying date information from 67 populations of two European songbirds covering a large part of their breeding range. Populations inhabiting deciduous habitats showed stronger phenological sensitivity compared with those in evergreen and mixed habitats. Strikingly, however, the lowest sensitivity was seen in populations that had experienced the greatest change in climate. Therefore, we predict that the strongest phenological advancement will not occur in those populations with the highest sensitivity. Our results show that to effectively assess the impact of climate change on phenology across a species range it will be necessary to account for intra-specific variation in phenological sensitivity, climate change exposure, and the ecological characteristics of a population.

Group living can be beneficial when individuals reproduce or survive better in the presence of others, but, simultaneously, there might be costs due to competition for resources. Positive and negative effects on various fitness components might thus counteract each other, so integration is essential to determine their overall effect. Here, we investigated how an integrated fitness measure (reproductive values [RVs]) based on six fitness components varied with group size among group members in cooperatively breeding red-winged and superb fairy wrens (Malurus elegans and Malurus cyaneus, respectively). Despite life-history differences between the species, patterns of RVs were similar, suggesting that the same behavioral mechanisms are important. Group living reduced RVs for dominant males, but for other group members, this was true only in large groups. Decomposition analyses showed that our integrated fitness proxy was most strongly affected by group size effects on survival and was amplified through carryover effects between years. Our study shows that integrative consideration of fitness components and subsequent decomposition analysis provide much needed insights into the key behavioral mechanisms shaping the costs and benefits of group living. Such attribution is crucial if we are to synthesize the relative importance of the myriad group size costs and benefits currently reported in the literature.

Climate change has strong effects on traits such as phenology and physiology. Studies typically assume that climate-induced trait changes will have consequences for population dynamics, but explicit tests are rare. Body condition reflects energy storage and may directly affect how much can be invested in reproduction and survival. However, the causal pathway by which decreased body condition impacts population dynamics has never been quantified across multiple populations and species. Therefore, we lack a general understanding of the consequences of changes in condition for variables more relevant for conservation, such as population size. Using structural equation modeling, we investigate how temperature-induced changes in body condition affect reproduction, and the subsequent impact on population growth rates of 19 bird species across 80 Dutch sites over a 21-year period. Warmer temperatures were associated with decreased body condition, which led to both decreased and increased reproduction at different sites, cancelling out any overall effect. The indirect effect of temperature on population growth (via body condition and reproduction) only explained within-species variation in the total effects of temperature on population growth. Instead, the direct effect of temperature on population growth (unrelated to condition and reproduction) was the most important pathway underlying the total effects of temperature on population growth, suggesting that unknown variables are mediating this effect. About half of the species are expected to increase under global warming, but this variation was not associated with any species characteristic. Overall, body condition responses to global warming are common, but their consequences on reproduction and subsequently population growth contribute relatively little to the total temperature impacts on population dynamics. Given that warming temperatures have strong effects on population dynamics, understanding the pathways via which temperature impacts population dynamics will be crucial for our ability to predict climate change effects in the future and improve conservation efforts.

Assessing the impacts of disturbance over large areas and long time periods is crucial for nature management, but also challenging since impacts depend on both wildlife responses to disturbance and on the spatiotemporal distribution of disturbance sources. Combined tracking of animals and disturbance sources enables quantification of wildlife responses as a function of the distance to a disturbance source. We provide a framework to derive such distance–response curves and combine those with disturbance source presence data to quantify energetic costs of disturbance at a landscape scale. We tracked 90 Eurasian Oystercatchers Haematopus ostralegus and all aircraft in a military training area in the Dutch Wadden Sea. We quantified distance–response curves estimating flight probability and additional displacement for five types of aircraft activities, by comparing bird movement prior to aircraft presence with movement during aircraft presence. We then used the distance–response curves to map mean and variation in additional daily energy expenditure due to cumulative aircraft disturbance across the landscape for a 700‐day period. Flight probability and displacement responses differed strongly among aircraft activities and decreased from transport aeroplanes, through bombing jets, helicopters, jets to small civil aeroplanes. Since the most disturbing aircraft activities were also the rarest ones, mean additional daily energy expenditure did not exceed 0.25%. However, days with substantial (>1%) additional expenditure occurred between 0.1% and 3.7% of all days across high‐tide roosts in the tidal basin. Notably, expenditure particularly spiked on days with transport aeroplane activity (up to 8.5%). Synthesis and applications. We quantified cumulative energetic flight costs due to aircraft disturbance and found that these were low and unlikely to impact survival of oystercatchers in our study area. Our results provide evidence that the legal minimum flight height of 450 m for small civil aeroplanes effectively limits the disturbance of oystercatchers. Mitigation should focus on limiting the number of days when disturbance has a high impact by reducing rare but highly disturbing activities, especially transport aeroplanes. Our approach can be applied to other species and disturbance sources that are automatically tracked, for example boats and walkers, ultimately to quantify the entire anthropogenic disturbance landscape.

L-M.C. and P.d.V. acknowledge support from the European Research Council (ERC) (Grant 678140-FluctEvol). The Montpellier tit group acknowledges the long-term support of the Observatoire des Sciences de l’Univers – Obser-vatoire de REcherche Montpelliérain de l’Environnement (OSU-OREME). The bighorn, mountain goat, and eastern gray kangaroo studies were supported by Natural Sciences and Engineering Research Council (NSERC) of Canada. Recent data collection for Wytham has been provided by grants from Biotechnology and Biological Sciences Research Council (BB/L006081/1), ERC (AdG250164), and the UK Natural Environment Research Council (NERC) (NE/K006274/1, NE/S010335/1). The Columbian ground squirrel study was supported by the National Science Foundation (Grant DEB-0089473). Trait and fitness data for hihi were collected/managed by John Ewen under New Zealand Department of Conservation hihi management contracts and research permits AK/15073/RES, AK-24128-FAU, 36186-FAU, and 44300-FAU and with additional financial support via NERC UK, The Leverhulme Trust UK, Marsden Fund New Zealand, and the Hihi Conservation Charitable Trust. The data on reindeer were made available through the Reindeer husbandry in a Globalizing North Nordic Center of Excellence, and the crew at Kutuharju Experimental Reindeer Research Station in the Reindeer Herder’s Association are thanked for their valuable assistance and logistic support in data collection. The red deer, Silwood blue tit, and Soay sheep datasets were supported by UK NERC. Lundy sparrow data were supported by NERC, a Marie Skłodowska-Curie Action, and Volkswagenstiftung. The red squirrel project was funded by NSERC of Canada and the National Science Foundation. J.C.S. was supported by a grant from the Ministry of Economy and Competitivity, Spanish Research Council (CGL-2016-79568-C3-3-P). J.T., T.K., and M.G. were supported by the Research Council of Norway through its Centers for Excellence funding scheme, Project 223257. Research on fairy wrens has been supported by the Australian Research Council. The Northern wheatear and the flycatcher studies were supported by grants from the Swedish Research Council VR. Temporal variation in natural selection is predicted to strongly impact the evolution and demography of natural populations, with consequences for the rate of adaptation, evolution of plasticity, and extinction risk. Most of the theory underlying these predictions assumes a moving optimum phenotype, with predictions expressed in terms of the temporal variance and autocorrelation of this optimum. However, empirical studies seldom estimate patterns of fluctuations of an optimum phenotype, precluding further progress in connecting theory with observations. To bridge this gap, we assess the evidence for temporal variation in selection on breeding date by modeling a fitness function with a fluctuating optimum, across 39 populations of 21 wild animals, one of the largest compilations of long-term datasets with individual measurements of trait and fitness components. We find compelling evidence for fluctuations in the fitness function, causing temporal variation in the magnitude, but not the direction of selection. However, fluctuations of the optimum phenotype need not directly translate into variation in selection gradients, because their impact can be buffered by partial tracking of the optimum by the mean phenotype. Analyzing individuals that reproduce in consecutive years, we find that plastic changes track movements of the optimum phenotype across years, especially in bird species, reducing temporal variation in directional selection. This suggests that phenological plasticity has evolved to cope with fluctuations in the optimum, despite their currently modest contribution to variation in selection. Postprint

1. Much is known about the brief adult phase of fireflies. However, fireflies spend a relatively long developmental period under the soil surface. Climatic and soil conditions may directly affect the eggs, larvae, and pupae, and also affect them indirectly through predators, competitors, and prey items. Climatic conditions during the early life stages of this iconic species are therefore relevant to their hypothesised decline within the context of global warming. 2. Data on the abundance of fireflies were extracted from the publicly available citizen dataset across North America over a period of 9 years. The effects of weather in the 24 months before the observations of firefly abundance were documented based on 6761 observations. 3. Climatic conditions during both the larval and adult phases have a non-linear effect on adult firefly abundance. Maximum winter and spring temperatures and mean precipitation in the 20-month period before the observations had the greatest impact on the abundance of firefly adults. Low maximum soil moisture during the 5–19 months preceding the observations affected the adult abundance negatively, and high maximum soil moisture affected it positively. 4. After correcting the firefly abundance for these weather effects, it is estimated that the abundance of fireflies increased over the time period of this study. 5. This study suggests that early life climatic conditions have a small but significant impact on adult firefly abundance with a total R 2 of 0.017.

To understand the effects of climate change and predict its future impacts, biologists relate variation in biological variables to spatial or temporal variation in weather variables. It is often unclear a priori which weather variables are important, over which period they act, and in what way they affect biological responses. Recently, multiple methods have been proposed to determine climatic sensitivity. Yet there has been little critical comparison of these different methods, making it difficult to determine the most appropriate method for different situations. Furthermore, evolutionary and behavioural biologists investigate individual variation in phenotypically plastic responses to weather, while ecologists and conservation biologists investigate variation among populations or species in climatic sensitivity. How do we quantify climatic sensitivities in such a way that they can be used effectively for such comparative studies? We provide an overview of systematic approaches, the pros and cons of different methods available, and outstanding challenges.

Kubelka et al. (Science, 9 November 2018, p. 680-683) claim that climate change has disrupted patterns of nest predation in shorebirds. They report that predation rates have increased since the 1950s, especially in the Arctic. We describe methodological problems with their analyses and argue that there is no solid statistical support for their claims.

Background Ring wear and loss may have important consequences for mark-recapture studies that aim to estimate survival trends. Our study quantifies the rates of wear and loss from a long-running colour-ringing project of the Eurasian Oystercatcher (Haematopus ostralegus) in the Netherlands. Methods Our analysis included 8909 colour-ringed oystercatchers with 118,071 resightings, predominantly ringed and observed by citizen scientists. We quantified how ring wear and loss may vary with ring colour and age, and how this may impact resighting probabilities. We then determined how ring loss may impact survival estimates and resighting probabilities of mark-recapture models by simulating four scenarios of how citizen scientists may resight and report birds with lost colour rings. Results Annual rates of colour ring loss averaged 2.5% compared with 1% for ring wear, but lost rings also had a higher reporting probability (31.2%) than worn rings (10.3%). Lost rings may not directly impact survival estimates since 50% of oystercatchers with lost rings could still be uniquely identified. Ring wear and loss rapidly increased between 10 and 15 years after ringing. Rates of ring loss were comparable amongst ring colours, but the wear rate appeared higher for red and white rings compared to other colours. Rates of ring wear and loss varied significantly amongst different regions in our study, which were characterised as having different habitat features. Differential rates of ring wear may have important implications for studies conducted over large geographical scales or of multiple species. Conclusions Based on simulations of ring loss in our population, estimates of apparent survival may be 0.3–1.2% lower whilst the impact of ring wear was deemed even lower. We developed a table of recommendations containing advice for ring fitting, data collection, study design, and mark-recapture analyses, so that future colour-ringing studies can benefit from our experiences in collecting and analysing data of ring wear and loss through citizen science.

Kubelka et al. (Reports, 9 November 2018, p. 680) claim that climate change has disrupted patterns of nest predation in shorebirds. They report that predation rates have increased since the 1950s, especially in the Arctic. We describe methodological problems with their analyses and argue that there is no solid statistical support for their claims. Fil: Bulla, Martin. Nioz Royal Netherlands Institute For Sea Research; Países Bajos. Max Planck Institute For Ornithology; Alemania. Czech University of Life Sciences; República Checa Fil: Reneerkens, Jeroen. Nioz Royal Netherlands Institute For Sea Research; Países Bajos. University of Groningen; Países Bajos Fil: Weiser, Emily L.. Upper Midwest Environmental Sciences Center; Estados Unidos Fil: Sokolov, Aleksandr. Institute Of Plant And Animal Ecology; Rusia Fil: Taylor, Audrey R.. University of Alaska; Estados Unidos Fil: Sittler, Benoît. Albert Ludwigs University of Freiburg; Alemania. Arctic Ecology Research Group; Francia Fil: McCaffery, Brian J.. U.S. Fish and Wildlife Service; Estados Unidos Fil: Ruthrauff, Dan R.. U.S. Geological Survey; Estados Unidos Fil: Catlin, Daniel H.. Virginia Tech University; Estados Unidos Fil: Payer, David C.. National Park Service; Estados Unidos Fil: Ward, David H.. U.S. Geological Survey; Estados Unidos Fil: Solovyeva, Diana V.. Institute of Biological Problems of the North; Rusia Fil: Santos, Eduardo S. A.. Universidade de São Paulo; Brasil Fil: Rakhimberdiev, Eldar. Lomonosov Moscow State University; Rusia. University of Groningen; Países Bajos Fil: Nol, Erica. Trent University; Canadá Fil: Kwon, Eunbi. Virginia Tech; Estados Unidos Fil: Brown, Glen S.. Ministry of Natural Resources and Forestry; Canadá Fil: Hevia, Glenda Denise. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Centro para el Estudio de Sistemas Marinos; Argentina Fil: Gates, H. River. National Audubon Society; Estados Unidos Fil: Johnson, James A.. Fish and Wildlife Service; Estados Unidos Fil: van Gils, Jan A.. Department of Coastal Systems and Utrecht University; Países Bajos Fil: Hansen, Jannik. Aarhus University. Aarhus Institute Of Advanced Studies.; Dinamarca Fil: Lamarre, Jean-François. Polar Knowledge Canada; Canadá Fil: Rausch, Jennie. Environment and Climate Change Canada; Canadá Fil: Conklin, Jesse R.. University of Groningen; Países Bajos Fil: Liebezeit, Joe. Audubon Society of Portland; Estados Unidos Fil: Bêty, Joël. University of Quebec; Canadá Fil: Lang, Johannes. Arctic Ecology Research Group ; Francia. Giessen University; Alemania Fil: Alves, José A.. University of Aveiro; Portugal. University of Iceland; Islandia Fil: Fernández-Elipe, Juan. No especifíca; Fil: Exo, Klaus-Michael. Institute of Avian Research “Vogelwarte Helgoland"; Alemania Fil: Bollache, Loïc. Universite de Franche-Comte; Francia Fil: Bertellotti, Néstor Marcelo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Centro para el Estudio de Sistemas Marinos; Argentina Fil: Giroux, Marie-Andrée. Université de Moncton; Canadá Fil: van de Pol, Martijn. Netherlands Institute of Ecology ; Países Bajos Fil: Johnson, Matthew. USDA Forest Service; Estados Unidos Fil: Boldenow, Megan L.. University of Alaska; Estados Unidos Fil: Valcu, Mihai. Max Planck Institute für Ornithologie; Alemania Fil: Soloviev, Mikhail. Lomonosov Moscow State University; Alemania Fil: Sokolova, Natalya. Arctic Research Station; Rusia Fil: Senner, Nathan R.. University of South Carolina; Estados Unidos Fil: Lecomte, Nicolas. Université de Moncton; Canadá Fil: Meyer, Nicolas. Universite de Franche-Comte; Francia. Arctic Ecology Research Group; Francia Fil: Schmidt, Niels Martin. Aarhus University. Department of Bioscience; Dinamarca Fil: Gilg, Olivier. Arctic Ecology Research Group; Francia. Universite de Franche-Comte; Francia Fil: Smith, Paul A.. Environment and Climate Change Canada; Canadá Fil: Machín, Paula. No especifíca; Fil: McGuire, Rebecca L.. Wildlife Conservation Society; Estados Unidos Fil: Cerboncini, Ricardo A. S.. Universidade Federal do Paraná; Brasil Fil: Ottvall, Richard. No especifíca; Fil: van Bemmelen, Rob S. A.. Wageningen Marine Research; Países Bajos Fil: Swift, Rose J.. Cornell University; Estados Unidos Fil: Saalfeld, Sarah T.. No especifíca; Fil: Jamieson,Sarah E.. Simon Fraser University; Canadá Fil: Brown, Stephen. No especifíca; Fil: Piersma, Theunis. NIOZ Royal Netherlands Institute for Sea Research; Países Bajos. Conservation Ecology Group; Países Bajos Fil: Albrecht, Tomas. Czech Academy of Sciences; República Checa. Charles University; República Checa Fil: D’Amico, Verónica. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Centro para el Estudio de Sistemas Marinos; Argentina Fil: Lanctot, Richard B.. U.S. Fish and Wildlife Service; Estados Unidos Fil: Kempenaers, Bart. Max Planck Institute für Ornithologie; Alemania

Migratory connectivity describes linkages between breeding and non-breeding areas. An ongoing challenge is tracking avian species between breeding and non-breeding areas and hence estimating migratory connectivity and seasonal survival. Collaborative color-ringing projects between researchers and citizen scientists provide opportunities for tracking the annual movements of avian species. Our study describes seasonal survival and migratory connectivity using data from more than 4,600 individuals with over 51,000 observations, predominantly collected by citizen scientists. Our study focuses on the Eurasian Oystercatcher (Haematopus ostralegus), a species that has experienced a substantial and ongoing decline in recent decades. Multiple threats have been described, and given that these threats vary in space and time, there is an urgent need to estimate demographic rates at the appropriate spatio-temporal scale. We performed a seasonal multi-state (5 geographical areas within The Netherlands) live- and dead-recoveries analysis under varying model structures to account for biological and data complexity. Coastal breeding populations were largely sedentary, while inland breeding populations were migratory and the direction of migration varied among areas, which has not been described previously. Our results indicated that survival was lower during winter than summer and that survival was lower in inland areas compared with coastal areas. A concerning result was that seasonal survival of individuals over-wintering in the Wadden Sea, an internationally important site for over-wintering shorebirds, appeared to decline during the study period. We discuss the outcomes of our study, and how citizen science was integral for conducting this study. Our findings identify how the demographic rates of the oystercatcher vary in space and time, knowledge that is vital for generating hypotheses and prioritizing future research into the causes of decline.

Extra‐pair paternity (EPP) has been suggested to improve the genetic quality of offspring, but evidence has been equivocal. Benefits of EPP may be only available to specific individuals or under certain conditions. Red‐winged fairy‐wrens have extremely high levels of EPP, suggesting fitness benefits might be large and available to most individuals. Furthermore, extreme philopatry commonly leads to incestuous social pairings, so inbreeding avoidance may be an important selection pressure. Here, we quantified the fitness benefits of EPP under varying conditions and across life‐stages. Extra‐pair offspring (EPO) did not appear to have higher fitness than within‐pair offspring (WPO), neither in poor years nor in the absence of helpers‐at‐the‐nest. However, EPP was beneficial for closely related social pairs, because inbred WPO suffered an overall 75% reduction in fitness. Inbreeding depression was nonlinear and reduced nestling body condition, first year survival and reproductive success. Our comprehensive study indicates that EPP should be favored for the 17% of females paired incestuously, but cannot explain the widespread infidelity in this species. Furthermore, our finding that fitness benefits of EPP only become apparent for a small part of the population could potentially explain the apparent absence of fitness differences in population wide comparisons of EPO and WPO.

Fitness can be profoundly influenced by the age at first reproduction (AFR), but to date the AFR–fitness relationship only has been investigated intraspecifically. Here, we investigated the relationship between AFR and average lifetime reproductive success (LRS) across 34 bird species. We assessed differences in the deviation of the Optimal AFR (i.e., the species-specific AFR associated with the highest LRS) from the age at sexual maturity, considering potential effects of life history as well as social and ecological factors. Most individuals adopted the species-specific Optimal AFR and both the mean and Optimal AFR of species correlated positively with life span. Interspecific deviations of the Optimal AFR were associated with indices reflecting a change in LRS or survival as a function of AFR: a delayed AFR was beneficial in species where early AFR was associated with a decrease in subsequent survival or reproductive output. Overall, our results suggest that a delayed onset of reproduction beyond maturity is an optimal strategy explained by a long life span and costs of early reproduction. By providing the first empirical confirmations of key predictions of life-history theory across species, this study contributes to a better understanding of life-history evolution.

Species' responses to environmental changes such as global warming are affected not only by trends in mean conditions, but also by natural and human-induced environmental fluctuations. Methods are needed to predict how such environmental variation affects ecological and evolutionary processes, in order to design effective strategies to conserve biodiversity under global change. Here, we review recent theoretical and empirical studies to assess: (1) how populations respond to changes in environmental variance, and (2) how environmental variance affects population responses to changes in mean conditions. Contrary to frequent claims, empirical studies show that increases in environmental variance can increase as well as decrease long-term population growth rates. Moreover, environmental variance can alter and even reverse the effects of changes in the mean environment, such that even if environmental variance remains constant, omitting it from population models compromises their ability to predict species' responses to changes in mean conditions. Drawing on theory relating these effects of environmental variance to the curvatures of population growth responses to the environment, we outline how species' traits such as phylogenetic history and body mass could be used to predict their responses to global change under future environmental variability.

1.Environmental variation can induce life history changes that can last over a large part of the lifetime of an organism. If multiple demographic traits are affected, expected changes in climate may influence environmental covariances among traits in a complex manner. Thus, examining the consequences of environmental fluctuations requires that individual information at multiple life stages are available, which is particularly challenging in long-lived species. 2.Here we analyse how variation in climatic conditions occurring in the year of hatching of female goshawks Accipiter gentilis (L.) affects age-specific variation in demographic traits and lifetime reproductive success (LRS). LRS decreased with increasing temperature in April in the year of hatching, due to lower breeding frequency and shorter reproductive lifespan. In contrast, the probability for a female to successfully breed was higher in years with a warm April, but lower LRS of the offspring in these years generated a negative covariance among fecundity rates among generations. 3.The mechanism by which climatic conditions generated cohort effects was likely through influencing the quality of the breeding segment of the population in a given year, as the proportion of pigeons in the diet during the breeding period was positively related to annual and lifetime reproductive success, and the diet of adult females that hatched in warm years contained fewer pigeons. 4.Climatic conditions experienced during different stages of individual life histories caused complex patterns of environmental covariance among demographic traits even across generations. Such environmental covariances may either buffer or amplify impacts of climate change on population growth, emphasizing the importance of considering demographic changes during the complete life history of individuals when predicting the effect of climatic change on population dynamics of long-lived species. This article is protected by copyright. All rights reserved.

More extreme climatic events (ECEs) are among the most prominent consequences of climate change. Despite a long-standing recognition of the importance of ECEs by paleo-ecologists and macro-evolutionary biologists, ECEs have only recently received a strong interest in the wider ecological and evolutionary community. However, as with many rapidly expanding fields, it lacks structure and cohesiveness, which strongly limits scientific progress. Furthermore, due to the descriptive and anecdotal nature of many ECE studies it is still unclear what the most relevant questions and long-term consequences are of ECEs. To improve synthesis, we first discuss ways to define ECEs that facilitate comparison among studies. We then argue that biologists should adhere to more rigorous attribution and mechanistic methods to assess ECE impacts. Subsequently, we discuss conceptual and methodological links with climatology and disturbance-, tipping point- and paleo-ecology. These research fields have close linkages with ECE research, but differ in the identity and/or the relative severity of environmental factors. By summarizing the contributions to this theme issue we draw parallels between behavioural, ecological and evolutionary ECE studies, and suggest that an overarching challenge is that most empirical and theoretical evidence points towards responses being highly idiosyncratic, and thus predictability being low. Finally, we suggest a roadmap based on the proposition that an increased focus on the mechanisms behind the biological response function will be crucial for increased understanding and predictability of the impacts of ECE. This article is part of the themed issue ‘Behavioural, ecological and evolutionary responses to extreme climatic events’.

Extra-pair paternity (EPP), where offspring are sired by a male other than the social male, varies enormously both within and among species. Trying to explain this variation has proved difficult because the majority of the interspecific variation is phylogenetically based. Ideally, variation in EPP should be investigated in closely related species, but clades with sufficient variation are rare. We present a comprehensive multifactorial test to explain variation in EPP among individuals in 20 populations of nine species over 89years from a single bird family (Maluridae). Females had higher EPP in the presence of more helpers, more neighbours or if paired incestuously. Furthermore, higher EPP occurred in years with many incestuous pairs, populations with many helpers and species with high male density or in which males provide less care. Altogether, these variables accounted for 48% of the total and 89% of the interspecific and interpopulation variation in EPP. These findings indicate why consistent patterns in EPP have been so challenging to detect and suggest that a single predictor is unlikely to account for the enormous variation in EPP across levels of analysis. Nevertheless, it also shows that existing hypotheses can explain the variation in EPP well and that the density of males in particular is a good predictor to explain variation in EPP among species when a large part of the confounding effect of phylogeny is excluded.

Fairy-wrens (genus Malurus) maintain territories year round, and breed cooperatively, with all members of the social group provisioning young. Despite living with several adult males, the breeding female typically cuckolds all of them, seeking fertilisations from extra-group males that provide no care to her offspring, instead caring for the young reared on their own territory. We trace the evolutionary origins and persistence of this extraordinary combination of traits. We argue that the high rate of infidelity in some fairy-wrens facilitates social pairing among nuclear family relatives, rather than being an evolutionary response to avoid inbreeding. It seems likely that females mate with extra-group males to improve the genetic quality of their offspring. The ability of males to maintain breeding plumage for long periods is the primary criterion for female choice; only older males can do so. Several features of the mating system undermine the accuracy of female choice, and low-quality males exploit this uncertainty. Extra-group matings by low-quality males can help stabilise the mating system but may leave it vulnerable to collapse under certain circumstances. Nonetheless, sexual selection in most species is very strong, confirming the utility of fairy-wrens as model organisms for the study of mate choice and intersexual selection.

To understand which populations and species are most sensitive to climate change, studies correlate time series of climate variables with those of traits important for population dynamics, and subsequently compare which aspects of a species' ecology or life-history best explain variation in climate sensitivity. Often large-scale oceanic climate indices (LOCIs) are used as a proxy for local climatic drivers, with many studies reporting geographic gradients in climate sensitivity to LOCIs (e.g. suggesting that species living further from the equator are relatively climate sensitive). However, the relationship between LOCIs and local weather variables also varies geographically, raising the possibility that apparent intra- and inter-specific differences in climate sensitivity to LOCIs could also reflect geographic variation in how well LOCIs function as a proxy for local climatic drivers. This hypothesis is rarely tested due to lack of knowledge about the specific local climatic drivers. Here we show, using reproductive and climate data from 16 long-term population studies of 7 Australian fairy-wren species (Malurus genus), that the use of LOCIs can result in 1) strong overestimation of the amount of inter-specific variation in climate sensitivity and 2) spurious patterns, particularly geographic gradients. Consequently a paradox emerges: LOCIs often explain much of the temporal variation in traits important for population dynamics, but the common usage of LOCIs may prevent meaningful intra- and inter-specific comparisons of climate sensitivities over large spatial scales. Our results thus may offer an alternative interpretation of the widely reported geographic gradients in sensitivity to LOCIs. Future progress will likely require better knowledge about the identity and temporal features of local environmental drivers of population dynamics. © 2013 The Authors.

The evolution of cooperation has been a focus of interest for evolutionary biologists for over a hundred years (Darwin 1859; Kropotkin 1908; Williams 1966). While all forms of cooperation challenge the centrality of competition in the process of evolution, cooperative and eusocial breeding systems, where offspring produced by a small number of breeding individuals are reared by nonbreeding helpers or workers, raise some of the most fundamental questions about the level at which selection operates the measurement of....

Summary Extreme climatic events (ECEs) are predicted to become more frequent as the climate changes. A rapidly increasing number of studies – though few on animals – suggest that the biological consequences of ECEs can be severe. However, ecological research on the impacts of ECEs has been limited by a lack of cohesiveness and structure. ECEs are often poorly defined and have often been confusingly equated with climatic variability, making comparison between studies difficult. In addition, a focus on short-term studies has provided us with little information on the long-term implications of ECEs, and the descriptive and anecdotal nature of many studies has meant it is still unclear what the key research questions are. Synthesizing the current state of work is essential to identify ways to make progress. We conduct a synthesis of the literature and discuss conceptual and practical challenges faced by research on ECEs. We consider three steps to advance research. First, we discuss the importance of choosing an ECE definition and identify the pros and cons of ‘climatological’ and ‘biological’ definitions of ECEs. Secondly, we advocate research beyond short-term descriptive studies to address questions concerning the long-term implications of ECEs, focussing on selective pressures and phenotypically plastic responses and how they might differ from responses to a changing climatic mean. Finally, we encourage a greater focus on multi-event studies that help us understand the implications of changing patterns of ECEs, through the combined use of modelling, experimental and observational field studies. This study aims to open a discussion on the definitions, questions and methods currently used to study ECEs, which will lead to a more cohesive approach to future ECE research.

1.Ecologists and many evolutionary biologists relate variation in physiological, behavioral, life-history, demographic, population and community traits to variation in weather, a key environmental driver. However, identifying which weather variables (e.g. rain, temperature, El Niño index), over which time period (e.g. recent weather, spring or year-round weather) and in what ways (e.g. mean, threshold of temperature) they affect biological responses is by no means trivial, particularly when traits are expressed at different times among individuals. 2.A literature review shows that a systematic approach for identifying weather signals is lacking and that the majority of studies select weather variables from a small number of competing hypotheses that are founded on unverified a priori assumptions. This is worrying because studies that investigate the nature of weather signals in detail suggest that signals can be complex. Using suboptimal or wrongly-identified weather signals may lead to unreliable projections and management decisions. 3.We propose a four-step approach which allows for more rigorous identification and quantification of weather signals (or any other predictor variable for which data is available at high temporal resolution), easily implementable with our new R package ‘climwin’. We compare our approach with conventional approaches and provide worked examples. 4.Although our more exploratory approach also has some drawbacks–such as the risk of overfitting and bias that our simulations show can occur at low sample and effect sizes—these issues can be addressed with the right knowledge and tools. 5.By developing both the methods to fit critical weather windows to a wide range of biological responses and the tools to validate them and determine sample size requirements, our approach facilitates the exploration and quantification of the biological effects of weather in a rigorous, replicable, and comparable way, while also providing a benchmark performance to compare other approaches to.