How stable are personalities? A multivariate view of behavioural variation over long and short timescales in the sheepshead swordtail,Xiphophorus birchmanni

Many studies have revealed repeatable (among-individual) variance in behavioural traits consistent with variation in animal personality; however, these studies are often conducted using data collected over single sampling periods, most commonly with short time intervals between observations. Consequently, it is not clear whether population-level patterns of behavioural variation are stable across longer timescales and/or multiple sampling periods or whether individuals maintain consistent ranking of behaviours (and/or personality) over their lifetimes. Here, we address these questions in a captive-bred population of a tropical freshwater poeciliid fish, Xiphophorus birchmanni. Using a multivariate approach, we estimate the among-individual variance-covariance matrix (I), for a set of behavioural traits repeatedly assayed in two different experimental contexts (open-field trials, emergence and exploration trials) over long-term (56 days between observations) and short-term (4-day observation interval) time periods. In both long- and short-term data sets, we find that traits are repeatable and the correlation structure of I is consistent with a latent axis of variation in boldness. While there are some qualitative differences in the way individual traits contribute to boldness and a tendency towards higher repeatabilities in the short-term study, overall, we find that population-level patterns of among-individual behavioural (co)variance to be broadly similar over both time frames. At the individual level, we find evidence that short-term studies can be informative for an individual’s behavioural phenotype over longer (e.g. lifetime) periods. However, statistical support is somewhat mixed and, at least for some observed behaviours, relative rankings of individual performance change significantly between data sets.     

Behavioral correlations across activity, mating, exploration, aggression, and antipredator contexts in the European house cricket, Acheta domesticus

Recently, there has been increasing interest in behavioral syndrome research across a range of taxa. Behavioral syndromes are suites of correlated behaviors that are expressed either within a given behavioral context (e.g., mating) or between different contexts (e.g., foraging and mating). Syndrome research holds profound implications for animal behavior as it promotes a holistic view in which seemingly autonomous behaviors may not evolve independently, but as a “suite” or “package.” We tested whether laboratory-reared male and female European house crickets, Acheta domesticus, exhibited behavioral syndromes by quantifying individual differences in activity, exploration, mate attraction, aggressiveness, and antipredator behavior. To our knowledge, our study is the first to consider such a breadth of behavioral traits in one organism using the syndrome framework. We found positive correlations across mating, exploratory, and antipredatory contexts, but not aggression and general activity. These behavioral differences were not correlated with body size or condition, although age explained some of the variation in motivation to mate. We suggest that these across-context correlations represent a boldness syndrome as individual risk-taking and exploration was central to across-context mating and antipredation correlations in both sexes.     

A systematic review of animal personality in conservation science

Although animal personality research may have applied uses, this suggestion has yet to be evaluated by assessing empirical studies examining animal personality and conservation. To address this knowledge gap, we performed a systematic review of the peer-reviewed literature relating to conservation science and animal personality. Criteria for inclusion in our review included access to full text, primary research articles, and relevant animal conservation or personality focus (i.e., not human personality studies). Ninety-two articles met these criteria. We summarized the conservation contexts, testing procedures (including species and sample size), analytical approach, claimed personality traits (activity, aggression, boldness, exploration, and sociability), and each report's key findings and conservation-focused suggestions. Although providing evidence for repeatability in behavior is crucial for personality studies, repeatability quantification was implemented in only half of the reports. Nonetheless, each of the 5 personality traits were investigated to some extent in a range of conservations contexts. The most robust studies in the field showed variance in how personality relates to other ecologically important variables across species and contexts. Moreover, many studies were first attempts at using personality for conservation purposes in a given study system. Overall, it appears personality is not yet a fully realized tool for conservation. To apply personality research to conservation problems, we suggest researchers think about where individual differences in behavior may affect conservation outcomes in their system, assess where there are opportunities for repeated measures, and follow the most current methodological guides on quantifying personality.     

Behavioral variation shows heritability in juvenile brown trout Salmo trutta

Animal personalities (sometimes referred also as coping styles) and their fitness consequences are currently among the most intensively explored subjects in behavioral ecology. To estimate the evolvability and adaptability of individually consistent behavioral variation, there is a crucial need to quantify the genetics underlying personality. Here, we experimentally studied the repeatability of various individual behaviors and then estimated heritability of formed boldness, exploration, and aggression components in juvenile brown trout Salmo trutta in standardized laboratory environment. Principal component analysis indicated that individually recorded behaviors were described by two personality axes: the first reflecting boldness, exploration, and aggression and the second tendency to freeze. These personality components, as well as the originally recorded behaviors, were statistically significantly repeatable over time. The latter PC, but not the first one, was statistically significantly heritable, though at low level (h ²?=?0.142?±?0.096). These results suggest that additive genetic variation underlies phenotypically consistent behavioral patterns, proposing that any selection acting on behavior, stress tolerance, or correlated traits has a potential to induce evolution in fish personality.     

Are behavioral syndromes invariant? Spatiotemporal variation in shy/bold behavior in squid

Behavioral syndromes are correlated suites of behavior, analogous to human personality traits. Most work to date has been taken from limited “snapshots” in space and time, with the implicit assumption that a behavioral syndrome is an invariant property, fixed by evolutionary constraints or adaptations. However, directional selection on two mechanistically independent traits (selective covariance) could also result in correlated behaviors. Previously, we have shown that shy/bold behavior in Southern dumpling squid (Euprymna tasmanica) across predator encounter and feeding risk contexts is genetically and phenotypically uncoupled, and hence potentially free to vary independently. Here, we collected data on shy/bold behaviors from two independent wild populations of squid in two different years to test whether behavioral correlations across these same two functional contexts vary through time and space. We detected significant influences of population, sex, and body size on the expression of boldness in squid within each functional context, and this was coupled with significant differences in relative population density and adult sex ratio. Despite these changes in behavior and demographic parameters, we found that correlations between boldness scores across the two functional contexts were largely absent in both wild populations of squid in both years. Our work suggests that some animal groups may be largely characterized by context-specific behavioral expression. A theoretical framework which conceptualizes behavioral syndromes resulting from context-specific behavioral rules may be needed to fully understand why behaviors are sometimes correlated, and why sometimes they are not.     

Shark personalities? Repeatability of social network traits in a widely distributed predatory fish

Interest in animal personalities has generated a burgeoning literature on repeatability in individual traits such as boldness or exploration through time or across different contexts. Yet, repeatability can be influenced by the interactive social strategies of individuals, for example, consistent inter-individual variation in aggression is well documented. Previous work has largely focused on the social aspects of repeatability in animal behaviour by testing individuals in dyadic pairings. Under natural conditions, individuals interact in a heterogeneous polyadic network. However, the extent to which there is repeatability of social traits at this higher order network level remains unknown. Here, we provide the first empirical evidence of consistent and repeatable animal social networks. Using a model species of shark, a taxonomic group in which repeatability in behaviour has yet to be described, we repeatedly quantified the social networks of ten independent shark groups across different habitats, testing repeatability in individual network position under changing environments. To understand better the mechanisms behind repeatable social behaviour, we also explored the coupling between individual preferences for specific group sizes and social network position. We quantify repeatability in sharks by demonstrating that despite changes in aggregation measured at the group level, the social network position of individuals is consistent across treatments. Group size preferences were found to influence the social network position of individuals in small groups but less so for larger groups suggesting network structure, and thus, repeatability was driven by social preference over aggregation tendency.     

Testing for between individual correlations of personality and physiological traits in a wild bird

Recently, integration of personality traits into a ‘pace-of-life syndrome’ (POLS) context has been advocated. To be able to understand how an individual’s behavioural, physiological and life history traits may coevolve, we need to jointly quantify these traits in order to study their covariance. Few studies have established links between personality and immunity properties of an individual. We here examined covariation of a measure of skeletal size (tarsus length), three behavioural traits (activity, handling aggression and breath rate) and two immunological traits (IgG level and haematocrit), in 592 wild caught blue tits. Many individuals (201) were tested more than once, allowing quantification of individual consistency of all traits and partition of the covariances between the traits, using a multivariate mixed model, into between individual and residual covariances. We find individual consistency of all behavioural traits, indicating that these capture aspects of blue tit adult personality and also the physiological measures are repeatable. Contrary to the POLS expectation, we find no overall significant individual level correlation structure between these traits and a factor analytical approach confirmed that between individual correlations across traits were not due to a common (POLS) factor or driven by size (tarsus length). Based on a published power study, we conclude that there is no common syndrome of individual level covariance between personality and physiological traits in wild blue tits or that the effect sizes, such a syndrome generates, are too low (r?<?0.3) to detect. Future field-based work should be designed to explore low effect sizes and strive to measure specific traits whose involvement is implicated to have large effect sizes as based on, e.g. laboratory findings.     

Heritable and experiential effects on boldness in a tropical poeciliid

Consistent differences in human behaviour are often explained with reference to personality traits. Recent evidence suggests that similar traits are widespread across the entire animal kingdom and that they may have substantial fitness consequences. One of the major components of personality is the shyness–boldness continuum. Little is known about the relative contributions of genes and the environment in the development of boldness in wild animal populations. Here, we bred wild-caught fish (Brachyraphis episcopi) collected from regions of high- and low-predation pressure, reared their offspring in the laboratory under varying conditions and tested boldness utilising an open-field paradigm. First-generation laboratory-reared fish showed similar behaviour to their wild parents suggesting that boldness has a heritable component. In addition, repeated chasing with a net increased boldness in both high- and low-predation offspring, showing that boldness is also heavily influenced by life experiences. Differences between males and females were also sustained in the laboratory-reared generation indicating that sex differences in boldness are also heritable. We discuss these results with reference to the potential underlying genetic and hormonal mechanisms as well as the environmental influences that may be responsible for expression of boldness in wild animals.     

Nymphal density, behavioral development, and life history in a field cricket

Population density regulates the strength of intraspecific competition and may thereby be reflected in life-history variables, such as development time, growth rate, or investment in immune defense. However, population density may also affect the fitness payoffs of different behaviors and consequently shape the development of personality. Here we studied if population density during nymphal development (one, four, or ten individuals raised together) affects the level of boldness, measured as the latency time to recover from freezing and emerge from a shelter, aggressiveness towards conspecifics or their correlation at the adult stage in the field crickets, Gryllus integer. In addition, we tested if individuals invest more resources in immune function or speed up their development in response to a high conspecifics density during ontogeny. Nymphal density did not affect adult boldness or aggressiveness towards conspecific males per se, but these variables showed a negative association, i.e., indicated an unconventional behavioral syndrome in the highest density treatment. Supporting the effectiveness of density treatments in inducing plastic responses, individuals reached maturity sooner and invested more resources in immune function in the highest nymphal density group compared to groups consisting of one or four individuals. Our results suggest that population density may play an important role in shaping both the realized life history and development of behavioral syndromes.     

Laterality is linked to personality in the black-lined rainbowfish,Melanotaenia nigrans

Emotions such as fear in vertebrates are often strongly lateralised, that is, a single cerebral hemisphere tends to be dominant when processing emotive stimuli. Boldness is a measure of an individual’s propensity to take risks and it has obvious connections with fear responses. Given the emotive nature of this well-studied personality trait, there is good reason to suspect that it is also likely to be expressed in a single hemisphere. Here, we examined the link between laterality and boldness in wild and captive-reared rainbowfish, Melanotaenia nigrans. We found that fish from the wild were bolder than those from captivity, which might be a reflection of the differences in the level of predation pressure experienced by the two populations. Secondly, we found that non-lateralised fish were bolder than strongly lateralised fish. In addition, differences in boldness scores between left- and right-biased fish were revealed. We suggest that variation in cerebral lateralisation contributes to the persistence of individual differences in boldness scores in animal populations.     

Do brain parasites alter host personality? — Experimental study in minnows

Despite that the existence of animal personalities is widely recognized, no consensus has been reached on the relative importance of different ecological factors behind their expression. Recently, it has been suggested that parasites may have a crucial role in shaping animal personalities, but only a very few studies have experimentally tested the idea. We infected Eurasian minnows (Phoxinus phoxinus) with the brain-encysted trematode parasite, Diplostomum phoxini, and studied whether infection could modify the personality of their hosts. Our results show that D. phoxini infection did not affect the mean levels of boldness, activity or exploration, but infected minnows showed higher repeatability in boldness and activity, and reduced repeatability in exploration. We also found that D. phoxini may be able to break the associations (behavioral syndromes) between behavioral traits, but that this effect may be dependent on parasite intensity. Furthermore, the effect of D. phoxini infection on personality of the hosts was found to be nonlinearly dependent on infection intensity. Taken together, our results suggest that D. phoxini parasites may shape the personality of their hosts, but that behavioral consequences of ecologically relevant infection levels may be rather subtle and easily remain undetected if only the mean trait expressions are compared.     

Evaluating animal personalities: do observer assessments and experimental tests measure the same thing?

The animal personality literature uses three approaches to assess personality. However, two of these methods, personality ratings and experimentation, have been little compared in captivity and never compared in the wild. We assessed the boldness of wild chacma baboons Papio ursinus using both ratings and experimental methods. Boldness was experimentally assessed when individuals were presented with a novel food item during natural foraging. The boldness of the same individuals was rated on a five-point scale by experienced observers. The ratings and experimental assessments of boldness were found to correlate positively and in a linear fashion. When considered categorically the two approaches showed variable agreement depending on the number of categories assigned and the cut-off criteria adopted. We suggest that the variation between approaches arises because each method captures different aspects of personality; ratings consider personality in absolute terms (using predefined criteria) and multiple contexts, while experimental assessments consider personality in relative terms (using experimental scores relative to the population average) and in limited contexts. We encourage animal personality researchers to consider adopting both methodologies in future studies. We also propose that future studies restrict their analyses to continuous data, since the greatest comparability between methods was found with these data. However, if individuals must be categorised, we suggest that researchers either (a) analyse only those individuals categorised as bold or shy by both ratings and experimental approaches or, if these methods cannot be employed simultaneously, (b) do not use approach-specific criteria but choose a cut-off that can be compared by both approaches.     

Habituation to low-risk predators improves body condition in lizards

Habituation to nonlethal predation stimuli may provide benefits for animals living in areas with frequent encounters with low-risk predators. On the other hand, individuals can be very consistent in their antipredator responses, with shy individuals showing greater degree of responsiveness than bold individuals. However, the link between habituation or boldness and individual benefits has not been thoroughly investigated. We established whether and how two behavioral components associated with antipredator responses (habituation and boldness, and their interaction) would influence body condition, which is a parameter related to fitness. We conducted an outdoor semi-natural experiment with Iberian wall lizards (Podarcis hispanica). Individual boldness was consistent across contexts, but we did not find any effect of boldness or the interaction between boldness and habituation on body condition. However, those individuals that habituated more readily to a frequent predatory stimulus were able to increase their body condition more relative to lizards that habituated less. This finding highlights the importance of individual differences in behavioral plasticity, which could influence traits related to fitness. Habituation can provide benefits for individuals exposed to low-risk predators; however, individuals more prone to habituation could also experience mortality costs by wrongly habituating to a dangerous predator.     

Personality trait differences between mainland and island populations in the common frog (Rana temporaria)

Understanding and predicting species range expansions is an important challenge in modern ecology because of rapidly changing environments. Recent studies have revealed that consistent within-species variation in behavior (i.e., animal personality) can be imperative for dispersal success, a key process in range expansion. Here we investigate how habitat isolation can mediate differentiation of personality traits between recently founded island populations and the main population. We performed laboratory studies of boldness and exploration across life stages (tadpoles and froglets) using four isolated island populations and four mainland populations of the common frog (Rana temporaria). Both tadpoles and froglets from isolated populations were bolder and more exploratory than conspecifics from the mainland. Although the pattern can be influenced by possible differences in predation pressure, we suggest that this behavioral differentiation might be the result of a disperser-dependent founder effect brought on by an isolation-driven environmental filtering of animal personalities. These findings can have important implications for both species persistence in the face of climate change (i.e., range expansions) and ecological invasions as well as for explaining rapid speciation in isolated patches.     

Causes and consequences of contest outcome: aggressiveness, dominance and growth in the sheepshead swordtail, Xiphophorus birchmanni

Although our understanding of how animal personality affects fitness is incomplete, one general hypothesis is that personality traits (e.g. boldness and aggressiveness) contribute to competitive ability. If so, then under resource limitation, personality differences will generate variation in life history traits crucial to fitness, like growth. Here, we test this idea using data from same-sex dyadic interaction trials of sheepshead swordtails (Xiphophorus birchmanni). In males, there was evidence of repeatable variation across a suite of agonistic contest behaviours, while repeatable opponent effects on focal behaviour were also detected. A single vector explains 80 % of the among-individual variance in multivariate phenotype and can be viewed as aggressiveness. We also find that aggressiveness predicts dominance—the repeatable tendency to win food in competition—and dominant individuals show faster post-trial weight gain (independently of initial size). In females, a dominance hierarchy predictive of weight gain was also found, but there was no evidence of variation in aggressiveness. While size often predicts contest outcome, our results show that individuals may sometimes grow larger because they are behaviourally dominant rather than vice versa. When resources are limited, personality traits such as aggression can influence growth, life history, and fitness through impacts on resource acquisition.     

Boldness by habituation and social interactions: a model

Most studies of animal personality attribute personality to genetic traits. But a recent study by Magnhagen and Staffan (Behav Ecol Sociobiol 57:295–303, 2005) on young perch in small groups showed that boldness, a central personality trait, is also shaped by social interactions and by previous experience. The authors measured boldness by recording the duration that an individual spent near a predator and the speed with which it fed there. They found that duration near the predator increased over time and was higher the higher the average boldness of other group members. In addition, the feeding rate of shy individuals was reduced if other members of the same group were bold. The authors supposed that these behavioral dynamics were caused by genetic differences, social interactions, and habituation to the predator. However, they did not quantify exactly how this could happen. In the present study, we therefore use an agent-based model to investigate whether these three factors may explain the empirical findings. We choose an agent-based model because this type of model is especially suited to study the relation between behavior at an individual level and behavioral dynamics at a group level. In our model, individuals were either hiding in vegetation or feeding near a predator, whereby their behavior was affected by habituation and by two social mechanisms: social facilitation to approach the predator and competition over food. We show that even if we start the model with identical individuals, these three mechanisms were sufficient to reproduce the behavioral dynamics of the empirical study, including the consistent differences among individuals. Moreover, if we start the model with individuals that already differ in boldness, the behavioral dynamics produced remained the same. Our results indicate the importance of previous experience and social interactions when studying animal personality empirically.     

Sex differences in the repeatability of boldness over metamorphosis

Consistent individual differences in boldness have been identified in many species and can have important effects on fitness. In most animals, juveniles face different costs and benefits of risk-taking behavior than do adults. Furthermore, profound changes in hormones, morphology and environment often occur when juveniles become adults. Therefore, the boldness of individuals might change with ontogeny. In field crickets, adult males call to attract sexually receptive females, and male calling increases predation risk. We measured the repeatability of boldness (latency to emerge from a safe refuge) in both male and female crickets. Each cricket was tested once as a small nymph and once as an adult. We found that boldness was repeatable across metamorphosis in females, but not in males. Males became less bold with maturation, a result that we predicted because of the risk associated with calling for mates. We also found that in general, nymphs were bolder than adults and that individuals spent more time immobile in response to a predator cue when they were nymphs, versus when they were adults.     

Prospecting at conspecific nests and exploration in a novel environment are associated with reproductive success in the jackdaw

Consistent behavioural differences between individuals of the same population (“personality” variation) might arise if individuals follow different life-history strategies. Thus, it would be important to determine how personality variation relates to behaviours potentially associated with life-history strategies, such as those related to the use of information about the state of the environment. Little is, however, known about how personality is associated with information use and reproductive success. We tested whether wild social jackdaws, Corvus monedula, show consistent behavioural differences in their exploratory behaviour (in a novel environment in the lab and in their reaction towards a novel object in the wild) and prospecting behaviour (number of visits to conspecific nests). We furthermore examined whether these behavioural traits are linked with each other and predictors of reproductive success. Breeding jackdaws were consistent in their exploratory behaviour within, and in their prospecting behaviour between, years. Exploratory behaviour in the novel environment was correlated with the latency to approach a novel object in the wild but not with the frequency of prospecting at conspecific nests. Highly exploratory males and females and frequently prospecting males produced fewer fledglings than less exploratory individuals or less prospecting males, respectively. We discuss the importance of consistent individual differences in exploration and information sampling on individual fitness.     

Retention of acquired predator recognition among shy versus bold juvenile rainbow trout

The ability to acquire information about predators allows prey to better balance threat-sensitive tradeoffs by responding only to ecologically relevant predation threats. However, predation risk is highly variable through time and responding to predators that no longer represent a threat would likely prove costly to prey. While a wealth of studies have examined the way in which prey learn, little attention has been paid to retention of acquired information. Recent studies suggest that retention is indeed plastic and shaped by a suite of intrinsic factors such as strength of initial conditioning and individual growth rate. Here, we investigated if the duration of retention of acquired information is influenced by individual behavioral tactics (i.e., ‘personality’). We recorded latency to escape an opaque acclimation chamber of juvenile rainbow trout (Oncorhynchus mykiss) as a measure of behavioral tactic. We then immediately conditioned individual trout to recognize pumpkinseed (Lepomis gibbosus) and tested for recognition 24 h or 8 days postconditioning. Our results demonstrate that while shy versus bold trout exhibited no difference in the strength of conditioned response to pumpkinseed odor during conditioning trials or when tested for recognition 24 h postconditioning, there was a significant effect of individual behavioral tactic on the retention of learned predator recognition. While shy trout continued to exhibit a learned response to pumpkinseed odor when tested 8 days postconditioning, bold trout were not different from our pseudoconditioned controls. These data suggest that the behavioral tactic employed at the time of conditioning influences the ‘memory window’ of acquired information.     

Correlates of boldness in three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>)

Behavioural variation is known to occur between individuals of the same population competing for resources. Individuals also vary with respect to their boldness or shyness. An individuals position along the shy-bold axis may be defined as the extent to which it is willing to trade off potentially increased predation risks for possible gains in resources. Similarly, group living may be interpreted as a trade-off between anti-predatory tactics and foraging efficiency. The responses of three-spined sticklebacks (Gasterosteus aculeatus) were tested across four social contexts to assess relative boldness or shyness and to further examine whether their behaviour would be consistent within and between contexts. Individuals displayed consistent responses within and between the first two contexts: those individuals which resumed foraging rapidly after a simulated aerial predator attack also displayed low shoaling tendencies. Such fish were deemed to be bold, whilst those which displayed the converse behaviour, slow resumption of foraging and a high shoaling tendency, were deemed to be shy. In a third context, bold individuals out-competed shy conspecifics for food. Boldness was also positively correlated with growth over a 6-week period. The position adopted by an individual within a group is usually interpreted as a trade-off between predation risk and foraging efficiency—both are greater at the front of a mobile group. Bold individuals showed significantly stronger tendencies towards front positions than shy conspecifics. The results suggest that, contrary to some previous studies on other animals, bold or shy behaviour in sticklebacks is consistent between contexts.Communicated by T. Czeschlik