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Phenological responses to climate change frequently vary among trophic levels, which can result in increasing asynchrony between the peak energy requirements of consumers and the availability of resources. Migratory birds use multiple habitats with seasonal food resources along migration flyways. Spatially heterogeneous climate change could cause the phenology of food availability along the migration flyway to become desynchronized. Such heterogeneous shifts in food phenology could pose a challenge to migratory birds by reducing their opportunity for food availability along the migration path and consequently influencing their survival and reproduction. We develop a novel graph‐based approach to quantify this problem and deploy it to evaluate the condition of the heterogeneous shifts in vegetation phenology for 16 migratory herbivorous waterfowl species in Asia. We show that climate change‐induced heterogeneous shifts in vegetation phenology could cause a 12% loss of migration network integrity on average across all study species. Species that winter at relatively lower latitudes are subjected to a higher loss of integrity in their migration network. These findings highlight the susceptibility of migratory species to climate change. Our proposed methodological framework could be applied to migratory species in general to yield an accurate assessment of the exposure under climate change and help to identify actions for biodiversity conservation in the face of climate‐related risks. [ABSTRACT FROM AUTHOR]

The social interactions that an individual experiences are a key component of its environment and can have important consequences for reproductive success. The dear enemy effect posits that having familiar neighbors at a territory boundary can reduce the need for territory defense and competition and potentially increase cooperation. Although fitness benefits of reproducing among familiar individuals are documented in many species, it remains unclear to what extent these relationships are driven by direct benefits of familiarity itself versus other socioecological covariates of familiarity. We use 58 years of great tit (Parus major) breeding data to disentangle the relationship between neighbor familiarity, partner familiarity, and reproductive success while simultaneously considering individual and spatiotemporal effects. We find that neighbor familiarity was positively associated with reproductive success for females but not males, while an individual's familiarity with their breeding partner was associated with fitness benefits for both sexes. There was strong spatial heterogeneity in all investigated fitness components, but our findings were robust and significant over and above these effects. Our analyses are consistent with direct effects of familiarity on individuals' fitness outcomes. These results suggest that social familiarity can yield direct fitness benefits, potentially driving the maintenance of long-term bonds and evolution of stable social systems. [ABSTRACT FROM AUTHOR]

Age shapes fundamental processes related to behaviour, survival and reproduction, where age influences reproductive success, non‐random mating with respect to age can magnify or mitigate such effects. Consequently, the correlation in partners' age across a population may influence its productivity. Despite widespread evidence for age‐assortative mating, little is known about what drives this assortment and its variation. Specifically, the relative importance of active (same‐age mate preference) and passive processes (assortment as a consequence of other spatial or temporal effects) in driving age assortment is not well understood.In this paper, we compare breeding data from a great tit and mute swan population (51‐ and 31‐year datasets, respectively) to tease apart the contributions of pair retention, cohort age structure and active age‐related mate selection to age assortment in species with contrasting life histories.Both species show age‐assortative mating and variable assortment between years. However, we demonstrate that the drivers of age assortment differ between the species, as expected from their life histories and resultant demographic differences. In great tits, pair fidelity has a weak effect on age‐assortative mating through pair retention; variation in age assortment is primarily driven by fluctuations in age structure from variable juvenile recruitment. Age‐assortative mating is, therefore, largely passive, with no evidence consistent with active age‐related mate selection. In mute swans, age assortment is partly explained by pair retention, but not population age structure, and evidence exists for active age‐assortative pairing.This difference is likely to result from shorter life‐spans in great tits compared with mute swans, leading to fundamental differences in their population age structure, whereby a larger proportion of great tit populations consist of a single age cohort. In mute swans, age‐assortative pairing through mate selection may also be driven by greater age‐dependent variation in fitness.The study highlights the importance of considering how different life histories and demographic differences arising from these affect population processes that appear congruent across species. We suggest that future research should focus on uncovering the proximate mechanisms that lead to variation in active age‐assortative mate selection (as seen in mute swans); and the consequences of variation in age structure on the ecological and social functioning of wild populations. [ABSTRACT FROM AUTHOR]

Identifying the environmental drivers of variation in fitness‐related traits is a central objective in ecology and evolutionary biology. Temporal fluctuations of these environmental drivers are often synchronized at large spatial scales. Yet, whether synchronous environmental conditions can generate spatial synchrony in fitness‐related trait values (i.e., correlated temporal trait fluctuations across populations) is poorly understood. Using data from long‐term monitored populations of blue tits (Cyanistes caeruleus, n = 31), great tits (Parus major, n = 35), and pied flycatchers (Ficedula hypoleuca, n = 20) across Europe, we assessed the influence of two local climatic variables (mean temperature and mean precipitation in February–May) on spatial synchrony in three fitness‐related traits: laying date, clutch size, and fledgling number. We found a high degree of spatial synchrony in laying date but a lower degree in clutch size and fledgling number for each species. Temperature strongly influenced spatial synchrony in laying date for resident blue tits and great tits but not for migratory pied flycatchers. This is a relevant finding in the context of environmental impacts on populations because spatial synchrony in fitness‐related trait values among populations may influence fluctuations in vital rates or population abundances. If environmentally induced spatial synchrony in fitness‐related traits increases the spatial synchrony in vital rates or population abundances, this will ultimately increase the risk of extinction for populations and species. Assessing how environmental conditions influence spatiotemporal variation in trait values improves our mechanistic understanding of environmental impacts on populations. [ABSTRACT FROM AUTHOR]

Understanding why individuals carry out behaviours that benefit others, especially genetically unrelated others, has been a major undertaking in many fields and particularly in biology. Here, we focus on the cooperation literature from natural populations and present the benefits of a social network approach in terms of how it can help to identify and understand factors that influence the maintenance and spread of cooperation, but are not easily captured when solely considering independent dyadic interactions. We describe how various routes to cooperation can be tested within the social network framework. Applying the social network approach to data from natural populations can help to uncover the evolutionary and ecological pressures that lead to differences in cooperation and other social processes. [ABSTRACT FROM AUTHOR]

The integration and synthesis of the data in different areas of science is drastically slowed and hindered by a lack of standards and networking programmes. Long‐term studies of individually marked animals are not an exception. These studies are especially important as instrumental for understanding evolutionary and ecological processes in the wild. Furthermore, their number and global distribution provides a unique opportunity to assess the generality of patterns and to address broad‐scale global issues (e.g. climate change).To solve data integration issues and enable a new scale of ecological and evolutionary research based on long‐term studies of birds, we have created the SPI‐Birds Network and Database (www.spibirds.org)—a large‐scale initiative that connects data from, and researchers working on, studies of wild populations of individually recognizable (usually ringed) birds. Within year and a half since the establishment, SPI‐Birds has recruited over 120 members, and currently hosts data on almost 1.5 million individual birds collected in 80 populations over 2,000 cumulative years, and counting.SPI‐Birds acts as a data hub and a catalogue of studied populations. It prevents data loss, secures easy data finding, use and integration and thus facilitates collaboration and synthesis. We provide community‐derived data and meta‐data standards and improve data integrity guided by the principles of Findable, Accessible, Interoperable and Reusable (FAIR), and aligned with the existing metadata languages (e.g. ecological meta‐data language).The encouraging community involvement stems from SPI‐Bird's decentralized approach: research groups retain full control over data use and their way of data management, while SPI‐Birds creates tailored pipelines to convert each unique data format into a standard format. We outline the lessons learned, so that other communities (e.g. those working on other taxa) can adapt our successful model. Creating community‐specific hubs (such as ours, COMADRE for animal demography, etc.) will aid much‐needed large‐scale ecological data integration. [ABSTRACT FROM AUTHOR]

Climate change has been shown to induce shifts in the timing of life‐history events. As a result, interactions between species can become disrupted, with potentially detrimental effects. Predicting these consequences has proven challenging. We apply structured population models to a well‐characterised great tit‐caterpillar model system and identify thresholds of temporal asynchrony, beyond which the predator population will rapidly go extinct. Our model suggests that phenotypic plasticity in predator breeding timing initially maintains temporal synchrony in the face of environmental change. However, under projections of climate change, predator plasticity was insufficient to keep pace with prey phenology. Directional evolution then accelerated, but could not prevent mismatch. Once predator phenology lagged behind prey by more than 24 days, rapid extinction was inevitable, despite previously stable population dynamics. Our projections suggest that current population stability could be masking a route to population collapse, if high greenhouse gas emissions continue. [ABSTRACT FROM AUTHOR]

Global climate change is altering the timing of life history events for species living in seasonal environments. These shifts in phenology can lead to the disruption of interspecific relationships with implications for individual fitness. Predicting phenological change and its population level consequences can provide insights into population persistence. Achieving this is challenging for labile traits as current structured population models do not explicitly distinguish between the roles of phenotypic plasticity and micro‐evolution, hindering realistic predictions of trait change. In this study we present the first empirical test of a new integral projection model (IPM) framework, which allows phenotypic plasticity and micro‐evolution to be teased apart by incorporating a quantitative genetic inheritance function. We parameterise this model for a population of wild great tits Parus major and test its predictive capabilities through K‐fold cross validation. We test the predictive accuracy of the quantitative genetic IPM in comparison to the standard IPM. We demonstrate that adding genetic inheritance rules maintains high accuracy of projections of phenological change, relative to the standard IPM. In addition, we find almost identical projections of population dynamics in this population for both IPMs, demonstrating that this model formulation allows researchers to investigate the contributions of phenotypic plasticity and micro‐evolution to trait change, without sacrificing predictive accuracy. Modelling in this way reveals that, under directional environmental change, both micro‐evolution and plasticity contribute to an advance of phenology, although the effect of plasticity is an order of magnitude higher than evolution. Despite this, synchrony between great tits and their caterpillar prey was reduced and population declines occurred. Our approach demonstrates that this model framework provides a promising avenue through which to explore the roles of phenotypic plasticity and evolution in trait changes and population dynamics. [ABSTRACT FROM AUTHOR]

An individual's fitness is not only influenced by its own phenotype, but by the phenotypes of interacting conspecifics. This is likely to be particularly true when considering fitness gains and losses caused by extrapair matings, as they depend directly on the social environment. While previous work has explored effects of dyadic interactions, limited understanding exists regarding how group-level characteristics of the social environment affect extrapair paternity (EPP) and cuckoldry. We use a wild population of great tits (Parus major) to examine how, in addition to the phenotypes of focal parents, two neighborhood-level traits—age and personality composition—predict EPP and cuckoldry. We used the well-studied trait "exploration behavior" as a measure of the reactive-proactive personality axis. Because breeding pairs inhabit a continuous "social landscape," we first established an ecologically relevant definition of a breeding "neighborhood" through genotyping parents and nestlings in a 51-ha patch of woodland and assessing the spatial predictors of EPP events. Using the observed decline in likelihood of EPP with increasing spatial separation between nests, we determined the relevant neighborhood boundaries, and thus the group phenotypic composition of an individual's neighborhood, by calculating the point at which the likelihood of EPP became negligible. We found no evidence that "social environment" effects (i.e. neighborhood age or personality composition) influenced EPP or cuckoldry. We did, however, find that a female's own age influenced the EPP of her social mate, with males paired to older females gaining more EPP, even when controlling for the social environment. These findings suggest that partner characteristics, rather than group phenotypic composition, influence mating activity patterns at the individual level. [ABSTRACT FROM AUTHOR]

Changes in the timing of life‐history events (phenology) are a widespread consequence of climate change. Predicting population resilience requires knowledge of how phenology is likely to change over time, which can be gained by identifying the specific environmental cues that drive phenological events. Cue identification is often achieved with statistical testing of candidate cues. As the number of methods used to generate predictions increases, assessing the predictive accuracy of different approaches has become necessary.This study aims to (a) provide an empirical illustration of the predictive ability of five commonly applied statistical methods for cue identification (absolute and relative sliding time window analyses, penalized signal regression, climate sensitivity profiles and a growing degree‐day model) and (b) discuss approaches for implementing cue identification methods in different systems.Using a dataset of mean clutch initiation timing in wild great tits (Parus major), we explored how the days of the year identified as most important, and the aggregate statistic identified as a cue, differed between statistical methods and with respect to the time span of data used. Each method's predictive capacity was tested using cross‐validation and assessed for robustness to varying sample size.We show that the identified critical time window of cue sensitivity was consistent across four of the five methods. The accuracy and precision of predictions differed by method with penalized signal regression resulting in the most accurate and most precise predictions in our case. Accuracy was maximal for near‐future predictions and showed a relationship with time. The difference between predictions and observations systematically shifted across the study from preceding observations to lagging.This temporal trend in prediction error suggests that the current statistical tools either fail to capture a key component of the cue–phenology relationship, or the relationship itself is changing through time in our system. These two influences need to be teased apart if we are to generate realistic predictions of phenological change. We recommend future phenological studies to challenge the idea of a static cue–phenology relationship and should cross‐validate results across multiple time periods. [ABSTRACT FROM AUTHOR]

For organisms living in seasonal environments, synchronizing the peak energetic demands of reproduction with peak food availability is a key challenge. Understanding the extent to which animals can adjust behavior to optimize reproductive timing, and the cues they use to do this, is essential for predicting how they will respond to future climate change. In birds, the timing of peak energetic demand is largely determined by the timing of clutch initiation; however, considerable alterations can still occur once egg laying has begun. Here, we use a wild population of great tits ( Parus major) to quantify individual variation in different aspects of incubation behavior (onset, duration, and daily intensity) and conduct a comprehensive assessment of the causes and consequences of this variation. Using a 54-year dataset, we demonstrate that timing of hatching relative to peak prey abundance (synchrony) is a better predictor of reproductive success than clutch initiation or clutch completion timing, suggesting adjustments to reproductive timing via incubation are adaptive in this species. Using detailed in-nest temperature recordings, we found that postlaying, birds improved their synchrony with the food peak primarily by varying the onset of incubation, with duration changes playing a lesser role. We then used a sliding time window approach to explore which spring temperature cues best predict variance in each aspect of incubation behavior. Variation in the onset of incubation correlated with mean temperatures just prior to laying; however, incubation duration could not be explained by any of our temperature variables. Daily incubation intensity varied in response to daily maximum temperatures throughout incubation, suggesting female great tits respond to temperature cues even in late stages of incubation. Our results suggest that multiple aspects of the breeding cycle influence the final timing of peak energetic demand. Such adjustments could compensate, in part, for poor initial timing, which has significant fitness impacts. [ABSTRACT FROM AUTHOR]

Many organisms rely on synchronizing the timing of their life-history events with those of other trophic levels-known as phenological matching-for survival or successful reproduction. In temperate deciduous forests, the extent of matching with the budburst date of key tree species is of particular relevance for many herbivorous insects and, in turn, insectivorous birds. In order to understand the ecological and evolutionary forces operating in these systems, we require knowledge of the factors influencing leaf emergence of tree communities. However, little is known about how phenology at the level of individual trees varies across landscapes, or how consistent this spatial variation is between different tree species. Here, we use field observations, collected over 2 years, to characterize within- and between-species differences in spring phenology for 825 trees of six species ( Quercus robur, Fraxinus excelsior, Fagus sylvatica, Betula pendula, Corylus avellana, and Acer pseudoplatanus) in a 385-ha woodland. We explore environmental predictors of individual variation in budburst date and bud development rate and establish how these phenological traits vary over space. Trees of all species showed markedly consistent individual differences in their budburst timing. Bud development rate also varied considerably between individuals and was repeatable in oak, beech, and sycamore. We identified multiple predictors of budburst date including altitude, local temperature, and soil type, but none were universal across species. Furthermore, we found no evidence for interspecific covariance of phenology over space within the woodland. These analyses suggest that phenological landscapes are highly complex, varying over small spatial scales both within and between species. Such spatial variation in vegetation phenology is likely to influence patterns of selection on phenology within populations of consumers. Knowledge of the factors shaping the phenological environments experienced by animals is therefore likely to be key in understanding how these evolutionary processes operate. [ABSTRACT FROM AUTHOR]

Population-level studies of how tit species ( Parus spp.) track the changing phenology of their caterpillar food source have provided a model system allowing inference into how populations can adjust to changing climates, but are often limited because they implicitly assume all individuals experience similar environments. Ecologists are increasingly using satellite-derived data to quantify aspects of animals' environments, but so far studies examining phenology have generally done so at large spatial scales. Considering the scale at which individuals experience their environment is likely to be key if we are to understand the ecological and evolutionary processes acting on reproductive phenology within populations. Here, we use time series of satellite images, with a resolution of 240 m, to quantify spatial variation in vegetation green-up for a 385-ha mixed-deciduous woodland. Using data spanning 13 years, we demonstrate that annual population-level measures of the timing of peak abundance of winter moth larvae ( Operophtera brumata) and the timing of egg laying in great tits ( Parus major) and blue tits ( Cyanistes caeruleus) is related to satellite-derived spring vegetation phenology. We go on to show that timing of local vegetation green-up significantly explained individual differences in tit reproductive phenology within the population, and that the degree of synchrony between bird and vegetation phenology showed marked spatial variation across the woodland. Areas of high oak tree ( Quercus robur) and hazel ( Corylus avellana) density showed the strongest match between remote-sensed vegetation phenology and reproductive phenology in both species. Marked within-population variation in the extent to which phenology of different trophic levels match suggests that more attention should be given to small-scale processes when exploring the causes and consequences of phenological matching. We discuss how use of remotely sensed data to study within-population variation could broaden the scale and scope of studies exploring phenological synchrony between organisms and their environment. [ABSTRACT FROM AUTHOR]

Associative learning is essential for resource acquisition, predator avoidance and reproduction in a wide diversity of species, and is therefore a key target for evolutionary and comparative cognition research. Automated operant devices can greatly enhance the study of associative learning and yet their use has been mainly restricted to laboratory conditions. We developed a portable, weatherproof, battery-operated operant device and conducted the first fully automated colour-associative learning experiment using free-ranging individuals in the wild. We used the device to run a colour discrimination task in a monitored population of tits (Paridae). Over two winter months, 80 individuals from four species recorded a total of 5,128 trials. Great tits (Parus major) were more likely than other species to visit the devices and engage in trials, but there were no sex or personality biases in the sample of great tits landing at the devices and registering key pecks. Juveniles were more likely than adults to visit the devices and to register trials. Individuals that were successful at solving a novel technical problem in captivity (lever-pulling) learned faster than non-solvers when at the operant devices in the wild, suggesting cross-contextual consistency in learning performance in very different tasks. There was no significant effect of personality or sex on learning rate, but juveniles’ choice accuracy tended to improve at a faster rate than adults. We discuss how customisable automated operant devices, such as the one described here, could prove to be a powerful tool in evolutionary ecology studies of cognitive traits, especially among inquisitive species such as great tits. [ABSTRACT FROM AUTHOR]

Summary: Cognition has been studied intensively for several decades, but the evolutionary processes that shape individual variation in cognitive traits remain elusive [1–3]. For instance, the strength of selection on a cognitive trait has never been estimated in a natural population, and the possibility that positive links with life history variation [1–5] are mitigated by costs [6] or confounded by ecological factors remains unexplored in the wild. We assessed novel problem-solving performance in 468 wild great tits Parus major temporarily taken into captivity and subsequently followed up their reproductive performance in the wild. Problem-solver females produced larger clutches than nonsolvers. This benefit did not arise because solvers timed their breeding better, occupied better habitats, or compromised offspring quality or their own survival. Instead, foraging range size and day length were relatively small and short, respectively, for solvers, suggesting that they were more efficient at exploiting their environment. In contrast to the positive effect on clutch size, problem solvers deserted their nests more often, leading to little or no overall selection on problem-solving performance. Our results are consistent with the idea that variation in cognitive ability is shaped by contrasting effects on different life history traits directly linked to fitness [1, 3]. [ABSTRACT FROM AUTHOR]

Behavioural traits generally and cognitive traits in particular are relatively understudied in an evolutionary ecological context. One reason for this is that such traits are often difficult to characterize among large numbers of individuals, without the influence of diverse environmental effects swamping intrinsic individual differences. We conducted standardized assays on a natural population of great tits, Parus major, to quantify and characterize individual variation in problem-solving performance, a simple cognitive trait often linked to innovative foraging ability. Forty-four per cent of 570 birds solved a food-motivated, lever-pulling problem and this proportion was consistent across three seasons. Individual performance was consistent within and across captivity sessions, across seasons, and between two different problem-solving tasks (lever and string pulling). Problem-solving performance was not explained by differences in latency to approach the empty task, nor latency to feed after human disturbance. Variation was unrelated to body condition, while age and natal origin explained significant but minimal amounts of variation, the importance of which varied between seasons. Problem-solving performance did not covary with exploration behaviour of a novel environment, suggesting that individual differences in problem solving represent an independent source of behavioural variation in our population. Rather than simply reflecting covariance with state or with other behavioural traits, our results suggest that variation in problem-solving performance represents inherent individual differences in the propensity to forage innovatively. We suggest that standardized problem-solving assays may prove ideal for studying the evolutionary ecology of simple cognitive traits. [ABSTRACT FROM AUTHOR]

Animal personalities can influence social interactions among individuals, and thus have major implications for population processes and structure. Few studies have investigated the significance of the social context of animal personalities, and such research has largely focused on the social organization of nonterritorial populations. Here we address the question of whether exploratory behaviour, a well-studied personality trait, is related to the social structure of a wild great tit, Parus major , population during the breeding season. We assayed the exploration behaviour of wild-caught great tits and then established the phenotypic spatial structure of the population over six consecutive breeding seasons. Network analyses of breeding proximity revealed that males, but not females, show positive assortment by behavioural phenotype, with males breeding closer to those of similar personalities. This assortment was detected when we used networks based on nearest neighbours, but not when we used the Thiessen polygon method where neighbours were defined from inferred territory boundaries. Further analysis found no relationship between personality assortment and local environmental conditions, suggesting that social processes may be more important than environmental variation in influencing male territory choice. This social organization during the breeding season has implications for the strength and direction of both natural and sexual selection on personality in wild animal populations. [ABSTRACT FROM AUTHOR]

Much of the evidence for the idea that individuals differ in their propensity to innovate and solve new problems has come from studies on captive primates. Increasingly, behavioural ecologists are studying innovativeness in wild populations, and uncovering links with functional behaviour and fitness-related traits. The relative importance of genetic and environmental factors in driving this variation, however, remains unknown. Here, we present the results of the first large-scale study to examine a range of causal factors underlying innovative problem-solving performance (PSP) among 831 great tits (Parus major) temporarily taken into captivity. Analyses show that PSP in this population: (i) was linked to a variety of individual factors, including age, personality and natal origin (immigrant or local-born); (ii) was influenced by natal environment, because individuals had a lower PSP when born in poor-quality habitat, or where local population density was high, leading to cohort effects. Links with many of the individual and environmental factors were present only in some years. In addition, PSP (iii) had little or no measurable heritability, as estimated by a Bayesian animal model; and (iv) was not influenced by maternal effects. Despite previous reports of links between PSP and a range of functional traits in this population, the analyses here suggest that innovativeness had weak if any evolutionary potential. Instead most individual variation was caused by phenotypic plasticity driven by links with other behavioural traits and by environmentally mediated developmental stress. Heritability estimates are population, time and context specific, however, and more studies are needed to determine the generality of these effects. Our results shed light on the causes of innovativeness within populations, and add to the debate on the relative importance of genetic and environmental factors in driving phenotypic variation within populations., (© 2016 The Author(s).)

Despite a growing body of evidence linking personality to life-history variation and fitness, the behavioural mechanisms underlying these relationships remain poorly understood. One mechanism thought to play a key role is how individuals respond to risk. Relatively reactive and proactive (or shy and bold) personality types are expected to differ in how they manage the inherent trade-off between productivity and survival, with bold individuals being more risk-prone with lower survival probability, and shy individuals adopting a more risk-averse strategy. In the great tit (Parus major), the shy-bold personality axis has been well characterized in captivity and linked to fitness. Here, we tested whether 'exploration behaviour', a captive assay of the shy-bold axis, can predict risk responsiveness during reproduction in wild great tits. Relatively slow-exploring (shy) females took longer than fast-exploring (bold) birds to resume incubation after a novel object, representing an unknown threat, was attached to their nest-box, with some shy individuals not returning within the 40 min trial period. Risk responsiveness was consistent within individuals over days. These findings provide rare, field-based experimental evidence that shy individuals prioritize survival over reproductive investment, supporting the hypothesis that personality reflects life-history variation through links with risk responsiveness., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)