Heterogeneous selection on exploration behavior within and among West European populations of a passerine bird.

Heterogeneous selection is often proposed as a key mechanism maintaining repeatable behavioral variation ("animal personality") in wild populations. Previous studies largely focused on temporal variation in selection within single populations. The relative importance of spatial versus temporal variation remains unexplored, despite these processes having distinct effects on local adaptation. Using data from >3,500 great tits (Parus major) and 35 nest box plots situated within five West-European populations monitored over 4 to 18 y, we show that selection on exploration behavior varies primarily spatially, across populations, and study plots within populations. Exploration was, simultaneously, selectively neutral in the average population and year. These findings imply that spatial variation in selection may represent a primary mechanism maintaining animal personalities, likely promoting the evolution of local adaptation, phenotype-dependent dispersal, and nonrandom settlement. Selection also varied within populations among years, which may counteract local adaptation. Our study underlines the importance of combining multiple spatiotemporal scales in the study of behavioral adaptation.

Artificial selection for reversal learning reveals limited repeatability and no heritability of cognitive flexibility in great tits (Parus major).

Cognitive flexibility controls how animals respond to changing environmental conditions. Individuals within species vary considerably in cognitive flexibility but the micro-evolutionary potential in animal populations remains enigmatic. One prerequisite for cognitive flexibility to be able to evolve is consistent and heritable among-individual variation. Here we determine the repeatability and heritability of cognitive flexibility among great tits (Parus major) by performing an artificial selection experiment on reversal learning performance using a spatial learning paradigm over three generations. We found low, yet significant, repeatability (R = 0.15) of reversal learning performance. Our artificial selection experiment showed no evidence for narrow-sense heritability of associative or reversal learning, while we confirmed the heritability of exploratory behaviour. We observed a phenotypic, but no genetic, correlation between associative and reversal learning, showing the importance of prior information on reversal learning. We found no correlation between cognitive and personality traits. Our findings emphasize that cognitive flexibility is a multi-faceted trait that is affected by memory and prior experience, making it challenging to retrieve reliable values of temporal consistency and assess the contribution of additive genetic variation. Future studies need to identify what cognitive components underlie variation in reversal learning and study their between-individual and additive genetic components.

Individual variation in the avian gut microbiota: The influence of host state and environmental heterogeneity.

The gut microbiota have important consequences for host biological processes and there is some evidence that they also affect fitness. However, the complex, interactive nature of ecological factors that influence the gut microbiota has scarcely been investigated in natural populations. We sampled the gut microbiota of wild great tits (Parus major) at different life stages allowing us to evaluate how microbiota varied with respect to a diverse range of key ecological factors of two broad types: (1) host state, namely age and sex, and the life history variables, timing of breeding, fecundity and reproductive success; and (2) the environment, including habitat type, the distance of the nest to the woodland edge, and the general nest and woodland site environments. The gut microbiota varied with life history and the environment in many ways that were largely dependent on age. Nestlings were far more sensitive to environmental variation than adults, pointing to a high degree of flexibility at an important time in development. As nestlings developed their microbiota from one to two weeks of life, they retained consistent (i.e., repeatable) among-individual differences. However these apparent individual differences were driven entirely by the effect of sharing the same nest. Our findings point to important early windows during development in which the gut microbiota are most sensitive to a variety of environmental drivers at multiple scales, and suggest reproductive timing, and hence potentially parental quality or food availability, are linked with the microbiota. Identifying and explicating the various ecological sources that shape an individual's gut bacteria is of vital importance for understanding the gut microbiota's role in animal fitness.

Handedness and individual roll-angle specialism when plunge diving in the northern gannet.

Many vertebrates show lateralized behaviour, or handedness, where an individual preferentially uses one side of the body more than the other. This is generally thought to be caused by brain lateralization and allows functional specializations such as sight, locomotion, and decision-making among other things. We deployed accelerometers on 51 northern gannets, Morus bassanus, to test for behavioural lateralization during plunge dives. When plunge diving, gannets 'roll' to one side, and standard indices indicated that 51% of individuals were left-sided, 43% right-sided, and 6% 'non-lateralized'. Lateralization indices provide no measure of error and do not account for environmental covariance, so we conducted two repeatability analyses on individuals' dive roll direction and angle. Dive side lateralization was highly repeatable among individuals over time at the population level (R = 0.878, p < 0.001). Furthermore, roll angle was also highly repeatable in individuals (R = 0.751, p < 0.001) even after controlling for lateralized state. Gannets show individual specializations in two different parts of the plunge diving process when attempting to catch prey. This is the first demonstration of lateralization during prey capture in a foraging seabird. It is also one of the few demonstrations of behavioural lateralization in a mixed model approach, providing a structure for further exploring behavioural lateralization.

Inhibitory control, exploration behaviour and manipulated ecological context are associated with foraging flexibility in the great tit.

Organisms are constantly under selection to respond effectively to diverse, sometimes rapid, changes in their environment, but not all individuals are equally plastic in their behaviour. Although cognitive processes and personality are expected to influence individual behavioural plasticity, the effects reported are highly inconsistent, which we hypothesise is because ecological context is usually not considered. We explored how one type of behavioural plasticity, foraging flexibility, was associated with inhibitory control (assayed using a detour-reaching task) and exploration behaviour in a novel environment (a trait closely linked to the fast-slow personality axis). We investigated how these effects varied across two experimentally manipulated ecological contexts-food value and predation risk. In the first phase of the experiment, we trained great tits Parus major to retrieve high value (preferred) food that was hidden in sand so that this became the familiar food source. In the second phase, we offered them the same familiar hidden food at the same time as a new alternative option that was visible on the surface, which was either high or low value, and under either high or low perceived predation risk. Foraging flexibility was defined as the proportion of choices made during 4-min trials that were for the new alternative food source. Our assays captured consistent differences among individuals in foraging flexibility. Inhibitory control was associated with foraging flexibility-birds with high inhibitory control were more flexible when the alternative food was of high value, suggesting they inhibited the urge to select the familiar food and instead selected the new food option. Exploration behaviour also predicted flexibility-fast explorers were more flexible, supporting the information-gathering hypothesis. This tendency was especially strong under high predation risk, suggesting risk aversion also influenced the observed flexibility because fast explorers are risk prone and the new unfamiliar food was perceived to be the risky option. Thus, both behaviours predicted flexibility, and these links were at least partly dependent on ecological conditions. Our results demonstrate that an executive cognitive function (inhibitory control) and a behavioural assay of a well-known personality axis are both associated with individual variation in the plasticity of a key functional behaviour. That their effects on foraging flexibility were primarily observed as interactions with food value or predation risk treatments also suggest that the population-level consequences of some behavioural mechanisms may only be revealed across key ecological conditions.

Cognitive flexibility in the wild: Individual differences in reversal learning are explained primarily by proactive interference, not by sampling strategies, in two passerine bird species.

Behavioural flexibility allows animals to adjust to changes in their environment. Although the cognitive processes that explain flexibility have been relatively well studied in psychology, this is less true for animals in the wild. Here we use data collected automatically during self-administered discrimination-learning trials for two passerine species, and during four phases (habituation, initial learning, first reversal and second reversal) in order to decompose sources of consistent among-individual differences in reversal learning, a commonly used measure for cognitive flexibility. First, we found that, as expected, proactive interference was significantly repeatable and had a negative effect on reversal learning, confirming that individuals with poor ability to inhibit returning to a previously rewarded feeder were also slower to reversal learn. Second, to our knowledge for the first time in a natural population, we examined how sampling of non-rewarding options post-learning affected reversal-learning performance. Sampling quantity was moderately repeatable in blue tits but not great tits; sampling bias, the variance in the proportion of visits to each non-rewarded feeder, was not repeatable for either species. Sampling behaviour did not predict variation in reversal-learning speed to any significant extent. Finally, the repeatability of reversal learning was explained almost entirely by proactive interference for blue tits; in great tits, the effects of proactive interference and sampling bias on the repeatability of reversal learning were indistinguishable. Our results highlight the value of proactive interference as a more direct measurement of cognitive flexibility and shed light on how animals respond to changes in their environment.

A time-lagged association between the gut microbiome, nestling weight and nestling survival in wild great tits.

Natal body mass is a key predictor of viability and fitness in many animals. While variation in body mass and therefore juvenile viability may be explained by genetic and environmental factors, emerging evidence points to the gut microbiota as an important factor influencing host health. The gut microbiota is known to change during development, but it remains unclear whether the microbiome predicts fitness, and if it does, at which developmental stage it affects fitness traits. We collected data on two traits associated with fitness in wild nestling great tits Parus major: weight and survival to fledging. We characterised the gut microbiome using 16S rRNA sequencing from nestling faeces and investigated temporal associations between the gut microbiome and fitness traits across development at Day-8 (D8) and Day-15 (D15) post-hatching. We also explored whether particular microbial taxa were 'indicator species' that reflected whether nestlings survived or not. There was no link between mass and microbial diversity on D8 or D15. However, we detected a time-lagged relationship where weight at D15 was negatively associated with the microbial diversity at D8, controlling for weight at D8, therefore reflecting relative weight gain over the intervening period. Indicator species analysis revealed that specificity values were high and fidelity values were low, suggesting that indicator taxa were primarily detected within either the survived or not survived groups, but not always detected in birds that either survived or died. Therefore these indicator taxa may be sufficient, but not necessary for determining either survival or mortality, perhaps owing to functional overlap in microbiota. We highlight that measuring microbiome-fitness relationships at just one time point may be misleading, especially early in life. Instead, microbial-host fitness effects may be best investigated longitudinally to detect critical development windows for key microbiota and host traits associated with neonatal weight. Our findings should inform future hypothesis testing to pinpoint which features of the gut microbial community impact on host fitness, and when during development this occurs. Such confirmatory research will shed light on population level processes and could have the potential to support conservation.

Cognition and covariance in the producer-scrounger game.

The producer-scrounger game is a key element of foraging ecology in many systems. Producing and scrounging typically covary negatively, but partitioning this covariance into contributions of individual plasticity and consistent between individual differences is key to understanding population-level consequences of foraging strategies. Furthermore, little is known about the role cognition plays in the producer-scrounger game. We investigated the role of cognition in these alternative foraging tactics in wild mixed-species flocks of great tits and blue tits, using a production learning task in which we measured individuals' speed of learning to visit the single feeder in an array that would provide them with a food reward. We also quantified the proportion of individuals' feeds that were scrounges ('proportion scrounged'); scrounging was possible if individuals visited immediately after a previous rewarded visitor. Three learning experiments-initial and two reversal learning-enabled us to estimate the repeatability and covariance of each foraging behaviour. First, we examined whether individuals learned to improve their scrounging success (i.e. whether they obtained food by scrounging when there was an opportunity to do so). Second, we quantified the repeatability of proportion scrounged, and asked whether proportion scrounged affected production learning speed among individuals. Third, we used multivariate analyses to partition within- and among-individual components of covariance between proportion scrounged and production learning speed. Individuals improved their scrounging success over time. Birds with a greater proportion scrounged took longer to learn their own rewarding feeder. Although multivariate analyses showed that covariance between proportion scrounged and learning speed was driven primarily by within-individual variation, that is, by behavioural plasticity, among-individual differences also played a role for blue tits. This is the first demonstration of a cognitive trait influencing producing and scrounging in the same wild system, highlighting the importance of cognition in the use of alternative resource acquisition tactics. The results of our covariance analyses suggest the potential for genetic differences in allocation to alternative foraging tactics, which are likely species- and system-dependent. They also point to the need to control for different foraging tactics when studying individual cognition in the wild.

Tidal drift removes the need for area-restricted search in foraging Atlantic puffins.

Understanding how animals forage is a central objective in ecology. Theory suggests that where food is uniformly distributed, Brownian movement ensures the maximum prey encounter rate, but when prey is patchy, the optimal strategy resembles a Lévy walk where area-restricted search (ARS) is interspersed with commuting between prey patches. Such movement appears ubiquitous in high trophic-level marine predators. Here, we report foraging and diving behaviour in a seabird with a high cost of flight, the Atlantic puffin ( Fratercula arctica), and report a clear lack of Brownian or Levy flight and associated ARS. Instead, puffins foraged using tides to transport them through their feeding grounds. Energetic models suggest the cost of foraging trips using the drift strategy is 28-46% less than flying between patches. We suggest such alternative movement strategies are habitat-specific, but likely to be far more widespread than currently thought.

Breeding phenology, provisioning behaviour, and unusual patterns of life history variation across an anthropogenic heterogeneous landscape.

Understanding how resource use and life history variation influence a population's response to modified, fragmented landscapes is a major challenge for ecologists. We investigated the phenology, life history decisions and provisioning behaviour of a generalist passerine-the great tit-across a heavily managed woodland landscape. Contrary to most previous studies on this species, reproductive investment and success were lower in deciduous than in coniferous woodland fragments. This could not be explained by differences in provisioning behaviour; instead population density was considerably higher in deciduous woodlands, suggesting birds did not follow an ideal free distribution. Clutch size declined with lay date amongst populations breeding in coniferous woodland fragments, but these populations also displayed pronounced seasonal declines in the proportion of fledglings produced per egg and fledgling mass. In contrast, and against patterns observed in other similar study systems, clutch size did not change with lay date in mixed-species deciduous woodland fragments. Furthermore, the proportion of young fledged and fledgling condition remained stable throughout the season, even though the quality of food provisioned to nestlings increased over the season. Local recruitment was negligible, suggesting that plasticity rather than natural selection played a key role in driving the patterns observed. The unusual patterns we report are likely explained by the fragmented nature of the landscape, and unreliable phenological cues in a mixed-species tree community coupled with low food availability. They contrast with those reported from most other populations situated in continuous woodland, demonstrating that caution is needed when generalising across different contexts and ecosystems.

Replicated analysis of the genetic architecture of quantitative traits in two wild great tit populations.

Currently, there is much debate on the genetic architecture of quantitative traits in wild populations. Is trait variation influenced by many genes of small effect or by a few genes of major effect? Where is additive genetic variation located in the genome? Do the same loci cause similar phenotypic variation in different populations? Great tits (Parus major) have been studied extensively in long-term studies across Europe and consequently are considered an ecological 'model organism'. Recently, genomic resources have been developed for the great tit, including a custom SNP chip and genetic linkage map. In this study, we used a suite of approaches to investigate the genetic architecture of eight quantitative traits in two long-term study populations of great tits--one in the Netherlands and the other in the United Kingdom. Overall, we found little evidence for the presence of genes of large effects in either population. Instead, traits appeared to be influenced by many genes of small effect, with conservative estimates of the number of contributing loci ranging from 31 to 310. Despite concordance between population-specific heritabilities, we found no evidence for the presence of loci having similar effects in both populations. While population-specific genetic architectures are possible, an undetected shared architecture cannot be rejected because of limited power to map loci of small and moderate effects. This study is one of few examples of genetic architecture analysis in replicated wild populations and highlights some of the challenges and limitations researchers will face when attempting similar molecular quantitative genetic studies in free-living populations.

Animal personality meets community ecology: founder species aggression and the dynamics of spider communities.

Silken web-reef created by the spider Anelosimus studiosus (main picture) and close-up (insert picture) of multi-female, adult colony of the same species. (photographs: T. Jones, J. Pruitt and A. Wild) In Focus: Pruitt, J.N. & Modlmeier, A.P. (2015) Animal personality in a foundation species drives community divergence and collapse in the wild. Journal of Animal Ecology, 84 Interspecific interactions form the cornerstone of niche theory in community ecology. The 7-year study In Focus here supports the view that variation within species could also be crucially important. Spider communities created experimentally in the wild, with either aggressive or docile individuals of the same founder species, were highly divergent in patterns of community succession for several years. Eventually, they converged on the same community composition only to collapse entirely shortly after, apparently because of the specific mix of aggression phenotypes within and between species just before collapse. These results suggest numerous avenues of research for behavioural ecology and evolutionary community ecology in metapopulations, and could help to resolve differences between competing theories.

Large-scale movements in European badgers: has the tail of the movement kernel been underestimated?

Characterizing patterns of animal movement is a major aim in population ecology, and yet doing so at an appropriate spatial scale remains a major challenge. Estimating the frequency and distances of movements is of particular importance when species are implicated in the transmission of zoonotic diseases. European badgers (Meles meles) are classically viewed as exhibiting limited dispersal, and yet their movements bring them into conflict with farmers due to their potential to spread bovine tuberculosis in parts of their range. Considerable uncertainty surrounds the movement potential of badgers, and this may be related to the spatial scale of previous empirical studies. We conducted a large-scale mark-recapture study (755 km(2); 2008-2012; 1935 capture events; 963 badgers) to investigate movement patterns in badgers, and undertook a comparative meta-analysis using published data from 15 European populations. The dispersal movement (>1 km) kernel followed an inverse power-law function, with a substantial 'tail' indicating the occurrence of rare long-distance dispersal attempts during the study period. The mean recorded distance from this distribution was 2.6 km, the 95 percentile was 7.3 km and the longest recorded was 22.1 km. Dispersal frequency distributions were significantly different between genders; males dispersed more frequently than females, but females made proportionally more long-distance dispersal attempts than males. We used a subsampling approach to demonstrate that the appropriate minimum spatial scale to characterize badger movements in our study population was 80 km(2), substantially larger than many previous badger studies. Furthermore, the meta-analysis indicated a significant association between maximum movement distance and study area size, while controlling for population density. Maximum long-distance movements were often only recorded by chance beyond the boundaries of study areas. These findings suggest that the tail of the badger movement distribution is currently underestimated. The implications of this for understanding the spatial ecology of badger populations and for the design of disease intervention strategies are potentially significant.

Shy birds play it safe: personality in captivity predicts risk responsiveness during reproduction in the wild.

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.

Larger groups of passerines are more efficient problem solvers in the wild.

Group living commonly helps organisms face challenging environmental conditions. Although a known phenomenon in humans, recent findings suggest that a benefit of group living in animals generally might be increased innovative problem-solving efficiency. This benefit has never been demonstrated in a natural context, however, and the mechanisms underlying improved efficiency are largely unknown. We examined the problem-solving performance of great and blue tits at automated devices and found that efficiency increased with flock size. This relationship held when restricting the analysis to naive individuals, demonstrating that larger groups increased innovation efficiency. In addition to this effect of naive flock size, the presence of at least one experienced bird increased the frequency of solving, and larger flocks were more likely to contain experienced birds. These findings provide empirical evidence for the "pool of competence" hypothesis in nonhuman animals. The probability of success also differed consistently between individuals, a necessary condition for the pool of competence hypothesis. Solvers had a higher probability of success when foraging with a larger number of companions and when using devices located near rather than further from protective tree cover, suggesting a role for reduced predation risk on problem-solving efficiency. In contrast to traditional group living theory, individuals joining larger flocks benefited from a higher seed intake, suggesting that group living facilitated exploitation of a novel food source through improved problem-solving efficiency. Together our results suggest that both ecological and social factors, through reduced predation risk and increased pool of competence, mediate innovation in natural populations.

Haplotype structure, adaptive history and associations with exploratory behaviour of the DRD4 gene region in four great tit (Parus major) populations.

The assessment of genetic architecture and selection history in genes for behavioural traits is fundamental to our understanding of how these traits evolve. The dopamine receptor D4 (DRD4) gene is a prime candidate for explaining genetic variation in novelty seeking behaviour, a commonly assayed personality trait in animals. Previously, we showed that a single nucleotide polymorphism in exon 3 of this gene is associated with exploratory behaviour in at least one of four Western European great tit (Parus major) populations. These heterogeneous association results were explained by potential variable linkage disequilibrium (LD) patterns between this marker and the causal variant or by other genetic or environmental differences among the populations. Different adaptive histories are further hypothesized to have contributed to these population differences. Here, we genotyped 98 polymorphisms of the complete DRD4 gene including the flanking regions for 595 individuals of the four populations. We show that the LD structure, specifically around the original exon 3 SNP is conserved across the four populations and does not explain the heterogeneous association results. Study-wide significant associations with exploratory behaviour were detected in more than one haplotype block around exon 2, 3 and 4 in two of the four tested populations with different allele effect models. This indicates genetic heterogeneity in the association between multiple DRD4 polymorphisms and exploratory behaviour across populations. The association signals were in or close to regions with signatures of positive selection. We therefore hypothesize that variation in exploratory and other dopamine-related behaviour evolves locally by occasional adaptive shifts in the frequency of underlying genetic variants.

Decadal increase in vessel interactions by a scavenging pelagic seabird across the North Atlantic.

Fisheries waste is used by many seabirds as a supplementary source of food,1 but interacting with fishing vessels to obtain this resource puts birds at risk of entanglement in fishing gear and mortality.2 As a result, bycatch is one of the leading contributors to seabird decline worldwide,3 and this risk may increase over time as birds increasingly associate fishing vessels with food. Light-level geolocators mounted on seabirds can detect light emitted from vessels at night year-round.4 We used a 16-year time series of geolocator data from 296 northern fulmars (Fulmarus glacialis) breeding at temperate and arctic colonies to investigate trends of nocturnal vessel interactions in this scavenging pelagic seabird. Vessel attendance has progressively increased over the study period despite no corresponding increase in the number of vessels or availability of discards over the same time frame. Fulmars are highly mobile generalist surface feeders,5 so this may signal a reduction in available prey biomass in the upper water column, leading to increased reliance on anthropogenic food subsidies6 and increased risk of bycatch mortality in already threatened seabird populations. Individuals were consistent in the extent to which they interacted with vessels, as shown in other species,7 suggesting that population-level increases may be due to a higher proportion of fulmars following vessels rather than changes at an individual level. Higher encounter rates were correlated with lower time spent foraging and a geographically restricted overwintering distribution, suggesting an energetic advantage for these scavenging strategists compared with foraging for natural prey.

Personality and problem-solving performance explain competitive ability in the wild.

Competitive ability is a major determinant of fitness, but why individuals vary so much in their competitiveness remains only partially understood. One increasingly prevalent view is that realized competitive ability varies because it represents alternative strategies that arise because of the costs associated with competitiveness. Here we use a population of great tits (Parus major) to explore whether individual differences in competitive ability when foraging can be explained by two traits that have previously been linked to alternative behavioural strategies: the personality trait 'exploration behaviour' and a simple cognitive trait, 'innovative problem-solving performance'. We assayed these traits under standardized conditions in captivity and then measured competitive ability at feeders with restricted access in the wild. Competitive ability was repeatable within individual males across days and correlated positively with exploration behaviour, representing the first such demonstration of a link between a personality trait and both competitive ability and food intake in the wild. Competitive ability was also simultaneously negatively correlated with problem-solving performance; individuals who were poor competitors were good at problem-solving. Rather than being the result of variation in 'individual quality', our results support the hypothesis that individual variation in competitive ability can be explained by alternative behavioural strategies.

Promiscuity, paternity and personality in the great tit.

Understanding causes of variation in promiscuity within populations remain a major challenge. While most studies have focused on quantifying fitness costs and benefits of promiscuous behaviour, an alternative possibility--that variation in promiscuity within populations is maintained because of linkage with other traits-has received little attention. Here, we examine whether promiscuity in male and female great tits (Parus major)--quantified as extra-pair paternity (EPP) within and between nests--is associated with variation in a well-documented personality trait: exploration behaviour in a novel environment. Exploration behaviour has been shown to correlate with activity levels, risk-taking and boldness, and these are behaviours that may plausibly influence EPP. Exploration behaviour correlated positively with paternity gained outside the social pair among males in our population, but there was also a negative correlation with paternity in the social nest. Hence, while variation in male personality predicted the relative importance of paternity gain within and outside the pair bond, total paternity gained was unrelated to exploration behaviour. We found evidence that males paired with bold females were more likely to sire extra-pair young. Our data thus demonstrate a link between personality and promiscuity, with no net effects on reproductive success, suggesting personality-dependent mating tactics, in contrast with traditional adaptive explanations for promiscuity.

Variation in personality and behavioural plasticity across four populations of the great tit Parus major.

1. Interest in the evolutionary origin and maintenance of individual behavioural variation and behavioural plasticity has increased in recent years. 2. Consistent individual behavioural differences imply limited behavioural plasticity, but the proximate causes and wider consequences of this potential constraint remain poorly understood. To date, few attempts have been made to explore whether individual variation in behavioural plasticity exists, either within or between populations. 3. We assayed 'exploration behaviour' among wild-caught individual great tits Parus major when exposed to a novel environment room in four populations across Europe. We quantified levels of individual variation within and between populations in average behaviour, and in behavioural plasticity with respect to (i) repeated exposure to the room (test sequence), (ii) the time of year in which the assays were conducted and (iii) the interval between successive tests, all of which indicate habituation to novelty and are therefore of functional significance. 4. Consistent individual differences ('I') in behaviour were present in all populations; repeatability (range: 0.34-0.42) did not vary between populations. Exploration behaviour was also plastic, increasing with test sequence - but less so when the interval between subsequent tests was relatively large - and time of year; populations differed in the magnitude of plasticity with respect to time of year and test interval. Finally, the between-individual variance in exploration behaviour increased significantly from first to repeat tests in all populations. Individuals with high initial scores showed greater increases in exploration score than individuals with low initial scores; individual by environment interaction ('I × E') with respect to test sequence did not vary between populations. 5. Our findings imply that individual variation in both average level of behaviour and behavioural plasticity may generally characterize wild great tit populations and may largely be shaped by mechanisms acting within populations. Experimental approaches are now needed to confirm that individual differences in behavioural plasticity (habituation) - not other hidden biological factors - caused the observed patterns of I × E. Establishing the evolutionary causes and consequences of this variation in habituation to novelty constitutes an exciting future challenge.