Rate of movement of juvenile lemon sharks in a novel open field, are we measuring activity or reaction to novelty?

ArticleinAnimal Behaviour 116:75-82 · June 2016with 282 Reads
Abstract
Personality differences are widespread throughout the animal kingdom and can have important ecological and evolutionary consequences. Despite a rapidly increasing body of literature, large (marine) vertebrates remain underrepresented in personality research. Given their unique life history traits (e.g. slow growth rate, slow reproduction rate, long life span) and their pivotal role in ecosystem processes, this is an important gap in our current knowledge. Here we investigated consistency and plasticity in movement behaviour of wild juvenile lemon sharks, Negaprion brevirostris, by repeatedly subjecting sharks to open field tests. First, we investigated the presence of interindividual differences in movement behaviour in a novel open field. Second, we investigated the effect of trial repetition on movement behaviour to understand whether movement in a novel open field reflects a reaction to novelty, or general activity. Third, we estimated individual differences in habituation/sensitization rates over trial repetition and studied how the habituation rate was predicted by the initial movement rate. We found consistent individual differences in movement behaviour during the open field tests. Sharks showed habituation in movement behaviour (i.e. decrease) over repeated trials indicating that the movement behaviour during the first trials is a reaction to novelty, and not general activity. Individuals, however, differed in their rate of habituation (i.e. plasticity) and this rate was negatively related to an individual's movement behaviour in the first open field trial. In addition to showing individual differences in consistency and plasticity in juvenile lemon sharks, our study emphasizes the importance of examining the validity of personality tests when adapting them to new species.

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  • Thesis
    Unevenness within a population is challenging to explain. It appears hazardous to interpret inter-individual dissimilarities in behavior, mainly due to a lack of information about the underlying mechanisms responsible for such expression. The key component of this study was the focus on the relationship between an intrinsic decision-making mechanism and the expression of individual movements. The uniqueness of this research laid in the study of how personality in juvenile lemon sharks, Negaprion brevirostris (Poey, 1868), may influence their natural behavior, providing a correlative analysis between personality and movement ecology. Twelve individuals were preliminarily exposed to a novel open field test to quantify a personality trait. Afterwards, the sharks were fitted with acoustic transmitters and monitored inside their nursery area, using an array of fifteen acoustic receivers, over an eight-month period. Movement patterns were assessed using active tracking. Home range and core area were measured using Minimum Convex Polygon (MCP) and Kernel Utilization Density (KUD). Although the two analyses produced different outputs, both revealed high individual differences in term of location and size. The results suggested an extensive use of the mangrove by the juveniles. The home range varied from 568.52m2 to 1296.01 m2 whilst using MCP approach, and ranged from 770.10 m2 to 1474.51 m2 based on the kernel-bivariate analysis. Similarly, core area estimates ranged from 85.88 m2 to 323.67 m2 (KUD). The estimation of the distance from the nearest shore captured a similar pattern and ranged from 38.16 m to 155.38 m. These inter-individual differences persisted even after effects of body size, sex or monitoring features were removed. However, multiple correlations revealed a strong relationship between personality traits and the spatial metrics (home range, Rs = 0.71; core area, Rs = 0.84; distance from the shore, Rs = 0.69). The results uncovered the likelihood of an influence of personality on the movement ecology of juvenile lemon sharks. Identifying mechanisms driving the expression of movement patterns provided crucial insight into decision-making processes at an individual level. Such observation should encourage further investigations to consider individual-based analyses for conservation purposes and advocate for the integration of behavioral ecology and movement ecology into a common framework to enhance the understanding of evolutionary and ecological processes.
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    Understanding the manner by which individual differences in personality arise and are maintained in animal populations is currently a topic of considerable research interest. This is particularly the case when it comes to developmental processes and understanding how behaviors change over ontogeny. Such developmental perspectives are essential given that the vast majority of animal species possess complex life cycles or undergo some form of metamorphosis. Yet, in spite of the broad taxonomic relevance and the obvious potential importance of metamorphosis for understanding the basis of consistency in personality over ontogeny, almost no research has been done on this topic. Using the lake frog (Rana ridibunda) as a study organism, we tested whether individual-level differences in personality (activity, exploration and boldness) were consistent within both larval and juvenile frog life-history stages and across metamorphosis. We found that most behaviors of interest were highly consistent within a given life-history stage and at least some traits were consistent across metamorphosis (e. g., activity and exploration). Generally, more active, exploratory individuals in novel experimental arenas were also bolder and more likely to spend time in more risky open areas of a familiar environment. To our knowledge, our study is the first to both characterize personality traits across anuran development and provide evidence of consistency in behavior across metamorphosis in a vertebrate species.
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    Although mixed effects models are widely used in ecology and evolution, their application to standardized traits that change within season or across ontogeny remains limited. Mixed models offer a robust way to standardize individual quantitative traits to a common condition such as body mass at a certain point in time (within a year or across ontogeny), or parturition date for a given climatic condition. Currently, however, most researchers use simple linear models to accomplish this task. We use both empirical and simulated data to underline the application of mixed models for standardizing trait values to a common environment for each individual. We show that mixed model standardizations provide more accurate estimates of mass parameters than linear models for all sampling regimes and especially for individuals with few repeated measures. Our simulations and analyses on empirical data both confirm that mixed models provide a better way to standardize trait values for individuals with repeated measure
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    There is growing evidence that individuals within populations show consistent differences in their behaviour across contexts (personality), and that personality is associated with the extent to which individuals adjust their behaviour as function of changing conditions (behavioural plasticity). We propose an evolutionary explanation for a link between personality and plasticity based upon how individuals manage uncertainty. Individuals can employ three categories of tactics to manage uncertainty. They can 1) gather information (sample) to reduce uncertainty, 2) show strategic (state-dependent) preferences for options that differ in their associated variances in rewards (i.e. variance-sensitivity), or 3) invest in insurance to mitigate the consequences of uncertainty. We explicitly outline how individual differences in the use of any of these tactics can generate personality-related differences in behavioural plasticity. For example, sampling effort is likely to co-vary with individual activity and exploration behaviours, while simultaneously creating population variation in reactions to changes in environmental conditions. Individual differences in the use of insurance may be associated with differences in risk-taking behaviours, such as boldness in the face of predation, thereby influencing the degree of adaptive plasticity across individuals. Population variation in responsiveness to environmental changes may also reflect individual differences in variance-sensitivity, because stochastic change in the environment increases variances in rewards, which may both attract and benefit variance-prone individuals, but not variance-averse individuals. We review the existing evidence that individual variation in strategies for managing uncertainty exist, and describe how positive-feedbacks between sampling, variance-sensitivity and insurance can maintain and exaggerate even small initial differences between individuals in the relative use of these tactics. Given the pervasiveness of the problem of uncertainty, alternative strategies for managing uncertainty may provide a powerful explanation for consistent differences in behaviour and behavioural plasticity for a wide range of traits.
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    Personality research has begun to take hold in the animal kingdom as psychologists turn to animal models to investigate various aspects of personality. Similarly, behavioral ecologists and related fields have begun to explore the idea that individual variation in behavior is more than just noise around an average for a given population or group of interest. As a result, many have begun to turn to personality-related questions to explain individual differences in animal behavior. Collectively, psychologists, ecologists and related fields have created a boom in animal personality-related research. This interest has expanded to a variety of fish species, with many studies focused on an important axis of behavior in humans: the shy-bold axis. Unfortunately, there has been very little consideration for the methodology employed. We review both the experimental and statistical methodology found in a body of research on fish species, for which personality-related research has been conducted. Our aim is to shed light on many important considerations that are often overlooked in order to facilitate research concerned with the reliability and validity of the many methods used. The classic approach to behavioral and evolutionary ecology seeks answers based on averaged behavior and fails to consider unique variation between individuals and the functional importance of such variability (Mather, 1998; Slater, 1981). The importance of finer scale investigations at the level of the individual is only beginning to emerge as researchers have begun explore Darwin's less accepted views that evolution acts on the individual and may not be limited to only physical traits (Darwin, 1998; Gosling, 2001). This burgeoning interest has given rise to research in areas of animal personality and temperament in a wide range of taxa and has led to recent recognition of the potential implications in ecological studies of animal behavior. If something similar to personality is found in animals and individual differences in behavior reflect more than just noise around an average, then traditional approaches to behavioral ecology, behavioral biology and how we approach questions about evolution may be challenged. The boom of interest in these new directions combined with the fear of anthropomorphism seems to have placed the search for "individual differences" at the forefront of investigations with little regard to ensuring adequate measures and methodology. In a complex and controversial topic such as animal personality, these considerations are vital to research in this area. This article aims to present some inconsistencies and considerations by reviewing a subset of studies focused on an important shy-bold axis, which has been a growing focus in research with many species of fish.
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    1. Body size determines rates of respiration and production, energy requirements, mortality rates, patterns of predation and vulnerability to mortality. Body size distribu-tions are often used to describe structure and energy flux in communities and ecosystems. 2. If clear relationships can be established between body size and trophic level in fishes, they may provide a basis for integrating community and ecosystem analyses based on size spectra, food webs and life histories. 3. We investigated relationships between the body sizes (weight and length) of north-east Atlantic fishes and their trophic level. The abundance of 15 N, as determined by stable isotope analysis, was used as an index of trophic level. 4. Cross-species and comparative analyses demonstrated that body size was unrelated or weakly related to trophic level. Thus allometric relationships between body size and trophic level could not be used to predict the trophic structure of fish communities. 5. The results of the cross-species analyses contrasted with patterns in the size and trophic structure of entire fish communities. When fish communities were divided into size classes, there were strong positive relationships between size class and trophic level. The slope suggested a mean predator : prey body mass ratio of 80 : 1. 6. Our results suggest that body size does not provide a useful surrogate of trophic level for individual species, but that body size is an excellent predictor of trophic level within the community, providing an empirical basis for integrating community analyses based on models of trophic structure and body size distributions.
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    Personality differences are a widespread phenomenon throughout the animal kingdom. Past research has focused on the characterization of such differences and a quest for their proximate and ultimate causation. However, the consequences of these differences for ecology and evolution received much less attention. Here, we strive to fill this gap by providing a comprehensive inventory of the potential implications of personality differences, ranging from population growth and persistence to species interactions and community dynamics, and covering issues such as social evolution, the speed of evolution, evolvability, and speciation. The emerging picture strongly suggests that personality differences matter for ecological and evolutionary processes (and their interaction) and, thus, should be considered a key dimension of ecologically and evolutionarily relevant intraspecific variation.
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    Boldness and shyness were investigated as ‘personality’ traits in hatchery-reared rainbow trout Oncorhynchus mykiss. Bold fish spent more time in an open area and were more active than shy fish and these behaviours could be used as indicators of boldness and shyness. These differences were related to learning ability in a simple conditioning task. Bold fish learned the task more quickly than shy fish.
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    Animal temperament describes behavioural differences between individuals that are consistent across time and contexts. Variation in animal temperament is rapidly gaining interest and attention within behavioural and evolutionary ecology. If we are to understand the causes and consequences of temperament variation within and between populations we need to determine the selection pressures that affect temperament in natural environments. To date, however, the vast majority of temperament studies have been carried out on captive-bred individuals. This review highlights potential problems that arise from using captive animals to elucidate the ecological and evolutionary functions of temperament in wild populations. For example, development, learning and environmental variability can all affect behaviour. Thus, both environment and gene-by environment interactions can affect the fitness functions of different temperaments, and hence selection. We stress the need for measurements of repeatability and heritability, and the importance of biological and ecological validation of temperament tests in wild animals. We describe the limited evidence from wild populations of the fitness consequences of temperament variation, and the use of intra- and inter-specific comparisons to prove adaptation. To identify multiple axes of behavioural variation, and how these interact with environments that vary spatially and temporally, we need long-term studies on wild populations – yet few studies of this nature currently exist. Finally, and perhaps counter-intuitively, we suggest that there is much to be gained from incorporating some of the approaches and statistics employed in the much longer established field of human personality.
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    1. Interest in measuring individual variation in reaction norms using mixed-effects and, more specifically, random regression models have grown apace in the last few years within evolution and ecology. However, these are data hungry methods, and little effort to date has been put into understanding how much and what kind of data we need to collect in order to apply these models usefully and reliably. 2. We conducted simulations to address three central questions. First, what is the best sampling strategy to collect sufficient data to test for individual variation using random regression models? Second, on occasions when precision is difficult to assess, can we be confident that a failure to detect significant variance in plasticity using random regression represents a biological reality rather than a lack of statistical power? Finally, does the common practice of censoring individuals with one or few repeated measures improve or reduce power to estimate individual variation in random regressions? 3. We have also developed a series of easy-to-use functions in the ‘pamm’ statistical package for R, which is freely available, that will allow researchers to conduct similar power analyses tailored more specifically to their own data. 4. Our results reveal potentially useful rules of thumb: large data sets (N > 200) are needed to evaluate the variance of individual-specific slopes; a number of individuals/number of observations per individual ratio of approximately 0·5 consistently yielded the highest power to detect random effects; individuals with one or few observations should not generally be censored as this reduces power to detect variance in plasticity. 5. We discuss the wider implications of these simulations and remaining challenges and suggest a new way to standardize results that would better facilitate the comparison of findings across empirical studies.
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    Intrapopulational polymorphism in habitat use is widely reported in many animal species. The phenomenon has recently also been recognized in adult female loggerhead sea turtles Caretta caretta, with small females tending to inhabit oceanic areas (where water depths are >200 m) while presumably feeding pelagically and large females tending to inhabit neritic areas (where depths are <200 m) while presumably feeding benthically. In this study, dive recording satellite telemetry units were used to verify their foraging and diving behaviours in these habitats. Two females that nested on Yakushima Island, Japan, were tracked for 124 and 197 days. The small female wandered in the oceanic Pacific, and spent most of the time at 0–25 m depths regardless of day or night, implying that she foraged pelagically at the surface and shallow depths. Her mean dive durations were significantly longer at night than during the day. The large female moved into the neritic East China Sea, and spent most of the time over the continental shelf at 100–150 m depths during the day and at 0–25 m depths at night, suggesting that she alternated between diurnal benthic foraging and nocturnal resting within the depths where she could attain neutral buoyancy. Her mean dive durations were not significantly different between day and night. The increase in dive duration for both turtles coincided with a seasonal decrease in water temperature. The small female sometimes showed midwater dormancy at 0–25 m depths with a duration of >5 h that was in contrast with bottom dormancy by sea turtles inhabiting other regions. The diving behaviours observed during this study were consistent with their estimated main feeding habits, which demonstrated resource polymorphism in a marine reptile.
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    A review of past behavioral ultrasonic telemetry studies of sharks and rays is presented together with previously unpublished material on the behavior of the lemon shark, Negaprion brevirostris, around the Bimini Islands, Bahamas. The review, focusing on movement behaviors of 20 shark and three ray species, reveals that elasmobranchs exhibit a variety of temporal and spatial patterns in terms of rates-of-movement and vertical as well as horizontal migrations. The lack of an apparent pattern in a few species is probably attributable to the scarcity of tracking data. Movements are probably governed by several factors, some still not studied, but data show that food, water temperature, bottom type, and magnetic gradient play major roles in a shark's decision of where and when to swim. A few species exhibit differences in behavior between groups of sharks within the same geographical area. This interesting finding warrants further research to evaluate the causes of these apparent differences and whether these groups constitute different subpopulations of the same species. The lack of telemetry data on batoids and some orders of sharks must be addressed before we can gain a more comprehensive understanding of the behavior of elasmobranch fishes. Previously unpublished data from 47 smaller and 38 larger juvenile lemon sharks, collected over the decade 1988–1998, provide new results on movement patterns, habitat selection, activity rhythms, swimming speed, rate-of-movement, and homing behavior. From these results we conclude that the lemon shark is an active predator with a strong, apparently innate homing mechanism. This species shows ontogenetic differences in habitat selection and behavior, as well as differences in movements between groups of individuals within the same area. We suggest three hypotheses for future research on related topics that will help to understand the enigmatic behavior of sharks.
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    An experimental group of lemon sharks received 100 daily presentations of light flash as the conditioned stimulus (CS) and electric shock as the unconditioned stimulus (US) in a classical conditioning situation. The conditioned responses (CRs) and unconditioned responses (URs) under observation consisted of extensions of the nictitating membrane. Separate control groups received either (a) no CS or US, (b) CS-alone, or (c) completely random presentations of CS and US. Few CRs occurred in the experimental group at the outset of conditioning, but the percentage of CRs during the second half of the first acquisition session exceeded 95%. Conditioning stabilized above 95% CRs during Acquisition Sessions 3 through 7. These responses could not be attributed to pseudoconditioning, sensitization, or other nonassociative factors. When the experimental group was subsequently given six CS-alone sessions, the course of extinction was gradual. Most results seemed similar to those previously obtained during classical conditioning of the nictitating membrane in rabbits.
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    Underwater field observations in a non-reproductive period revealed enormous variation in schooling tendencies among individual european wrasses, Symphodus ocellatus. The fish occupying vegetated habitats displayed much more pronounced schooling tendencies than those from open areas. Sixteen solitary and schooling males were caught with a small seine and observed in a novel environment. The former individuals turned out to be much more bold and active than the latter in this test, which would suggest lower fear and higher exploratory tendency. It was hypothesized that the bolder wrasses might appear as territorial nest-builders in the reproductive period, whereas the shyer ones are likely to play sneaky and satellite tactics.
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    We surgically implanted ultrasonic transmitters in 38 lemon sharks,Negaprion brevirostris, and manually tracked the sharks for 1–153 days. This yielded 2281 positional fixes recorded at 15-min intervals. We used these positional data with availability data of four environmental variables (water depth, temperature, salinity, and bottom type), sampled at 213 stations along 15 transects, to examine usage of habitat. All sharks used contours of water depth, water temperature, and bottom type disproportionately to the availability of these variables in the study site. Specifically, juvenile lemon sharks selected shallower, warmer water with an underlying rocky or sandy substrate, perhaps for predator avoidance. This is the first report on habitat selection by any elasmobranch.
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    Understanding the foraging behavior and spatial distribution of top predators is crucial to gaining a complete understanding of communities. However, studies of top predators are often logistically difficult and it is important to develop appropriate methods for identifying factors influencing their spatial distribution. Sharks are top predators in many marine communities, yet no studies have quantified the habitat use of large predatory sharks or determined the factors that might influence shark spatial distributions. We used acoustic telemetry and animal-borne video cameras ("Crittercam") to test the hypothesis that tiger shark (Galeocerdo cuvier) habitat use is determined by the availability of their prey. We also used Crittercam to conduct the first investigation of foraging behavior of tiger sharks. To test for habitat preferences of sharks, the observed proportion of time in each habitat for each individual was compared to the predicted values for that individual based on correlated random walk and track randomization methods. Although there was individual variation in habitat use, tiger sharks preferred shallow seagrass habitats, where their prey is most abundant. Despite multiple encounters with potential prey, sharks rarely engaged in prolonged high-speed chases, and did not attack prey that were vigilant. We propose that the tiger sharks' foraging tactic is one of stealth, and sharks rely upon close approaches to prey in order to be successful. This study shows that using appropriate analysis techniques and a variety of field methods it is possible to elucidate the factors influencing habitat use and gain insights into the foraging behavior of elusive top predators.
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    The existence of individual prey specializations has been reported for an ever-growing number of taxa, and has important ramifications for our understanding of predator–prey dynamics. We use the California sea otter population as a case study to validate the use of archival time–depth data to detect and measure differences in foraging behaviour and diet. We collected observational foraging data from radio-tagged sea otters that had been equipped with Mk9 time depth recorders (TDRs, Wildlife Computers, Redmond, WA). After recapturing the study animals and retrieving the TDRs it was possible to compare the two data types, by matching individual dives from the TDR record with observational data and thus examining behavioural correlates of capture success and prey species. Individuals varied with respect to prey selection, aggregating into one of three distinct dietary specializations. A number of TDR-derived parameters, particularly dive depth and post-dive surface interval, differed predictably between specialist types. A combination of six dive parameters was particularly useful for discriminating between specialist types, and when incorporated into a multivariate cluster analysis, these six parameters resulted in classification of 13 adult female sea otters into three clusters that corresponded almost perfectly to the diet-based classification (1 out of 13 animals was misclassified). Thus based solely on quantifiable traits of time–depth data that have been collected over an appropriate period (in this case 1 year per animal), it was possible to assign female sea otters to diet type with >90% accuracy. TDR data can thus be used as a tool to measure the degree of individual specialization in sea otter populations, a conclusion that will likely apply to other diving marine vertebrates as well. Our ultimate goals must be both to understand the causes of individual specialization, and to incorporate such variation into models of population- and community-level food web dynamics.
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    Group living in sharks is a widespread phenomenon but relatively little is known about the composition and organization of these groups. In binary choice field experiments juvenile lemon sharks were attracted to conspecifics presumably to form groups. Experiments investigating size assortment preferences indicated that lemon sharks aged 2–3 years (but not 0–1 years) preferred to spend more time with a group of size-matched individuals than unmatched ones. Furthermore, in species association tests lemon sharks spent significantly more time associating with conspecifics than with a sympatric heterospecific, the nurse shark, Ginglymostoma cirratum. These findings enhance our knowledge of group-joining decisions in sharks indicating that active mechanisms can play a role in the formation and composition of shark groups.
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    Hunting by humans played a major role in extirpating terrestrial megafauna on several continents and megafaunal loss continues today in both terrestrial and marine ecosystems. Recent declines of large marine vertebrates that are of little or no commercial value, such as sea turtles, seabirds and marine mammals, have focused attention on the ecological impacts of incidental take, or bycatch, in global fisheries. In spite of the recognition of the problem of bycatch, few comprehensive assessments of its effects have been conducted. Many vulnerable species live in pelagic habitats, making surveys logistically complex and expensive. Bycatch data are sparse and our understanding of the demography of the affected populations is often rudimentary. These factors, combined with the large spatial scales that pelagic vertebrates and fishing fleets cover, make accurate and timely bycatch assessments difficult. Here, we review the current research that addresses these challenging questions in the face of uncertainty, analytical limitations and mounting conservation crises.
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    Until recently, large apex consumers were ubiquitous across the globe and had been for millions of years. The loss of these animals may be humankind’s most pervasive influence on nature. Although such losses are widely viewed as an ethical and aesthetic problem, recent research reveals extensive cascading effects of their disappearance in marine, terrestrial, and freshwater ecosystems worldwide. This empirical work supports long-standing theory about the role of top-down forcing in ecosystems but also highlights the unanticipated impacts of trophic cascades on processes as diverse as the dynamics of disease, wildfire, carbon sequestration, invasive species, and biogeochemical cycles. These findings emphasize the urgent need for interdisciplinary research to forecast the effects of trophic downgrading on process, function, and resilience in global ecosystems.
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    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.