Food intake rates of inactive fish are positively linked to boldness in three‐spined sticklebacks Gasterosteus aculeatus

To investigate the link between personality and maximum food intake of inactive individuals, food‐deprived three‐spined sticklebacks Gasterosteus aculeatus at rest in their home compartments were provided with ad libitum prey items. Bolder individuals ate considerably more than shyer individuals, even after accounting for body size, while sociability did not have an effect. These findings support pace‐of‐life theory predicting that life‐history strategies are linked to boldness.     

Personality and body condition have additive effects on motivation to feed in Zebra Finches Taeniopygia guttata

Several hypotheses have been proposed to account for the adaptive evolution of personality, defined as inter‐individual differences in behaviour that are consistent over time and across situations. For instance, the ‘pace‐of‐life syndrome’ hypothesis suggests that personality evolved as a behavioural correlate of life‐history trajectories that vary within populations. Thus, proactivity, corresponding to higher exploratory tendencies or higher boldness levels, has been linked to higher productivity or mortality rates. However, the extent to which proactivity is associated with a higher motivation to forage remains poorly understood. Moreover, although personality and its effects on foraging behaviour are usually considered to be independent of any motivational or nutritional state, few studies so far have challenged this. Here we show that personality traits, both individually or combined using a principal component analysis, and body condition have additive effects on latency to feed following food deprivation in the Zebra Finch Taeniopygia guttata, with personality accounting for 41% and body condition for about 20% of the total variation in latency to feed. In accordance with the pace‐of‐life syndrome hypothesis, latency to feed was negatively related to the degree of proactivity and positively related to body condition. Thus, proactive individuals and individuals in poorer condition were quicker to start feeding after a period of food deprivation. The absence of a significant interaction between personality and body condition further suggests that the effect of personality was independent of body condition. We discuss the relevance of our results in relation to the different factors influencing foraging in birds. Moreover, we place our results within a life‐history framework by emphasizing the correlated evolution of life‐history traits and personality.     

Behavioral plasticity and consistency in the naked-footed mouse (<Emphasis Type="Italic">Peromyscus nudipes</Emphasis>) with habitat disturbance

Very few studies have evaluated whether habitat disturbance affects behavioral consistency and plasticity. We measured shyness–boldness and exploration–avoidance for the first time in naked-footed mice (Peromyscus nudipes) in two adjacent habitats with differing level of disturbance in Costa Rica. Each habitat was measured in different, consecutive years. With these data, we explored the possibility of habitat differences in individual- and population-level behavioral plasticity, magnitude of behavior, and consistency. Mice in both habitats behaved consistently across time with high repeatability (i.e., showed personality). The strength of consistency of shyness–boldness and exploration–avoidance over time was higher in the disturbed habitat, and individuals in the undisturbed habitat were bolder and less exploratory. Behaviors were correlated with each other in both habitats, indicating that behavioral syndromes do not always break down in disturbed areas. Mice changed the magnitude of their response the second time they were tested, indicating population-level plasticity. There was also greater individual plasticity among mice in the undisturbed habitat. Our study suggests the possibility that habitat disturbance can affect behavioral plasticity and personality and that shyer/more exploratory individuals might colonize and/or persist in disturbed habitats. These results are preliminary and exploratory because we did not control for temporal differences between habitats.     

Do Female Zebra Finches,Taeniopygia guttata,Choose Their Mates Based on Their ‘Personality’?

A major challenge in behavioural and evolutionary ecology is to understand the evolution and maintenance of consistent behavioural differences among individuals within populations, often referred to as animal ‘personalities’. Here, we present evidence suggesting that sexual selection may act on such personality differences in zebra finches (Taeniopygia guttata), as females seem to choose males on the basis of their exploratory behaviour per se, while taking into account their own personality. After observing a pair of males, whose apparent levels of exploration were experimentally manipulated, females that exhibited low‐exploratory tendencies showed no preference during mate choice for males that had appeared to be either ‘exploratory’ or ‘unexploratory’. In contrast, intermediate and highly exploratory females preferred apparently exploratory males over apparently unexploratory ones. Our results suggest that behavioural or genetic compatibility for personality traits might be important for mate choice, at least for exploratory individuals.     

Environmental and genetic effects shape the development of personality traits in the mangrove killifish Kryptolebias marmoratus

Consistent individual differences in behaviour are well documented, for example, individuals can be defined as consistently bold or consistently shy. To date our understanding of the mechanisms underpinning consistent individual differences in behaviour (also termed behavioural types (BTs)) remains limited. Theoretical work suggests life‐history tradeoffs drive BT variation, however, empirical support is scarce. Moreover, whilst life‐history is known to be phenotypically plastic in response to environmental conditions during ontogeny, the extent to which such plasticity drives plasticity in behavioural traits and personality remains poorly understood. Using a natural clonal vertebrate, Kryptolebias marmoratus, we control for genetic variation and investigate developmental plasticity in life‐history and three commonly studied behavioural traits (exploration, boldness, aggression) in response to three ecologically relevant environments; conspecific presence, low food and perceived risk. Simulated predation risk was the only treatment that generated repeatable behaviour i.e. personality during ontogeny. Treatments differed in their effects on mean life‐history and behavioural scores. Specifically, low food fish exhibited reduced growth rate and exploration but did not differ from control fish in their boldness or aggression scores. Conspecific presence resulted in a strong negative effect on mean aggression, boldness and exploration during ontogeny but had minimal effect on life‐history traits. Simulated predation risk resulted in increased reproductive output but had minimal effect upon average behavioural scores. Together these results suggest that life‐history plasticity/variation may be insufficient in driving variation in personality during development. Finally, using offspring derived from each rearing environment we investigate maternal effects and find strong maternal influence upon offspring size, but not behaviour. These results highlight and support the current understanding that risk perception is important in shaping personality, and that social experience during ontogeny is a major influence upon behavioural expression.     

Does individual personality predict male mating preference for female body size in the Trinidadian guppy?

Within populations, individual animals vary considerably in their behaviour, including mate choice and personality. There is mounting interest in the potential covariation between these two behaviours within individuals, such that personality would influence mate choice. We experimentally tested this proposition under controlled laboratory conditions using male Trinidadian guppies (Poecilia reticulata) as a model study system. We assayed repeatedly the mating preference of individual males for females based on their body size. Additionally, we assayed repeatedly two ecologically relevant personality traits in males, namely exploration of a novel environment and boldness under a simulated predation threat. Finally, we analysed whether male mating preference and personality traits were repeatable, and tested whether the personality of individual males was correlated (covaried) with their mating preference scores. Although all but one of the measures of exploration and boldness behaviour were repeatable over time, male mating preference scores were not repeatable. Measures of male exploration and boldness were not inter-correlated among individuals, suggesting the absence of a behavioural syndrome between exploration and boldness. Unexpectedly, males did not exhibit on average a significant mating preference for larger females over smaller ones; they chose randomly between the paired stimulus females. Overall, we found no compelling evidence for a relationship between individual personality traits and mating preference in male guppies, suggesting that personality does not predict mate choice, at least in our study population and under our experimental conditions. We discuss potential factors, other than male personality and body length, that might maintain inter-individual variation in male mating preferences in the guppy in the wild.     

Absence of a Context‐General Behavioural Syndrome in a Solitary Predator

The correlation of seemingly unrelated behaviours into behavioural syndromes has been established in a wide range of species and taxa. However, most studies report on short‐term behavioural correlations without insight into individual consistency or temporal stability of the behavioural syndrome. Here, we examine the individual repeatability of single behaviours, and the presence and temporal stability of a context‐general behavioural syndrome in a solitary piscivorous predator, the pike (Esox lucius). Behavioural measurements on the same individuals were quantified independently through time and across three contexts: activity in the presence of a competitor, exploration of a novel environment and boldness under predation risk. There was no indication of a temporally stable behavioural syndrome, consisting of boldness, activity and exploration, nor were individuals consistent in the separate behaviours, contradicting the general assertion of its taxonomic prevalence. Furthermore, the study did not provide support for size or growth‐dependent behaviour in this size‐dimorphic species in conditions of limited food availability. The study highlights the importance of independent multiple observations of individual behaviour across time or contexts when measuring behavioural repeatability and covariation.     

Disruptive viability selection on adult exploratory behaviour in eastern chipmunks

Heterogeneous forces of selection associated with fluctuating environments are recognized as important factors involved in the maintenance of inter‐individual phenotypic variance within populations. Consistent behavioural differences over time and across situations (e.g. personality) are increasingly cited as examples of individual variation observed within populations. However, the suggestion that heterogeneous selective pressures target different animal personalities remains largely untested in the wild. In this 5‐year study, we investigated the dynamics of viability selection on a personality trait, exploration, in a population of eastern chipmunks (Tamias striatus) experiencing substantial seasonal variations in weather conditions and food availability associated with masting trees. Contrary to our expectations, we found no evidence of fluctuating selection on exploration. Instead, we found strong disruptive viability selection on adult exploration behaviour, independent of seasonal variations. Individuals with either low or high exploration scores were almost twice as likely to survive over a 6‐month period compared with individuals with intermediate scores. We found no evidence of viability selection on juvenile exploration. Our results highlight that disruptive selection might play an important role in the maintenance of phenotypic variance of wild populations through its effect on different personality types across temporally varying environmental conditions.     

Florivory by a floriphilic katydid,Phaneroptera brevis,induces changes in a leaf trait in Lantana camara

1. Plants can induce a response when they are attacked by herbivores. Although the induction of responses by herbivory in both flowers and leaves is relatively well studied, whether florivory (feeding of flowers) can also induce responses in flowers and leaves is less well explored and there are still unanswered questions. These include whether plants exhibit different levels of induced responses depending on the length of exposure to the florivores. 2. To address this knowledge gap, this study used a tropical floriphilic katydid, Phaneroptera brevis, and its non‐native food plant, Lantana camara. Nursery experiments were performed in which dry matter content and anthocyanin concentration of the flower (corolla and stamens), leaf dry matter content, and leaf blade punch resistance were measured at three time phases before and after exposing the plant to the katydid individuals for 0–7 days. 3. It was demonstrated that increasing the length (days) of exposure to the katydid individual leads to higher levels of induced plant response (leaf blade punch resistance), but only in the leaves. It was also shown that higher levels of induced plant response owing to the increase in the length of exposure to the katydid individual was not observed beyond the first set of leaves developed after the exposure. 4. These results address the knowledge gap and show that plants can exhibit different levels of induced responses depending on the length of exposure to florivores. This study thus highlights the far‐reaching importance of florivory on plants.     

Heritability and social brood effects on personality in juvenile and adult life‐history stages in a wild passerine

How has evolution led to the variation in behavioural phenotypes (personalities) in a population? Knowledge of whether personality is heritable, and to what degree it is influenced by the social environment, is crucial to understanding its evolutionary significance, yet few estimates are available from natural populations. We tracked three behavioural traits during different life‐history stages in a pedigreed population of wild house sparrows. Using a quantitative genetic approach, we demonstrated heritability in adult exploration, and in nestling activity after accounting for fixed effects, but not in adult boldness. We did not detect maternal effects on any traits, but we did detect a social brood effect on nestling activity. Boldness, exploration and nestling activity in this population did not form a behavioural syndrome, suggesting that selection could act independently on these behavioural traits in this species, although we found no consistent support for phenotypic selection on these traits. Our work shows that repeatable behaviours can vary in their heritability and that social context influences personality traits. Future efforts could separate whether personality traits differ in heritability because they have served specific functional roles in the evolution of the phenotype or because our concept of personality and the stability of behaviour needs to be revised.     

Adaptive strategies for managing uncertainty may explain personality‐related differences in behavioural plasticity

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.     

Between florivory and herbivory: inefficacy of decision‐making by generalist floriphilic katydids

1. Florivory, which is less studied but as equally widespread as herbivory, can have effects on plant and floral trait adaptations, individual fitness, and community interactions. However, there are still knowledge gaps in our understanding of florivory, including the role of neural constraints and diet specialisation of florivores in floral resource utilisation. 2. According to the neural constraint hypothesis, a generalist flower‐visiting katydid is expected to have lower decision‐making efficacy in feeding when subjected to the presence of both capitula and leaves, compared to the presence of either one of the two resources. 3. In the present study, experiments using the katydid, Phaneroptera brevis, and the plant, whiteweed, Ageratum conyzoides, were carried out to examine the foraging behaviour of the katydid to test our hypothesis. The results confirmed the prediction of the neural constraint hypothesis. 4. The decision‐making efficacy was generally lowered when the katydid was presented with the choice for both the resource that is preferred (capitula) and less preferred (leaves). It was also shown that Phaneroptera brevis is floriphilic, as it prefers capitula to leaves. 5. In conclusion, the first evidence of neural constraint in florivory is provided and suggests that more can be explored into the effects of neural constraints and diet specialisation in floral resource utilisation.     

Parenting styles in white‐rumped swallows (Tachycineta leucorrhoa) show a trade‐off between nest defense and chick feeding

Studies about personalities in wild animals usually focus on five categories of behavioral traits that do not easily accommodate all aspects of parental care, a class of behaviors with direct consequence to reproductive success. Parental care can vary consistently between individuals and constitute parenting styles. Here we investigate the consistency of four behaviors of parental care across two breeding seasons of white‐rumped swallows Tachycineta leucorrhoa in southern Brazil. These behaviors are a prospection of potential nest‐sites, nest defense against predators or conspecifics, and chick feeding. If these first three behaviors are consistent, they can be classified according to the standard categories of personality as exploration, boldness, and aggressiveness, respectively. We find that all behaviors, except exploration, are consistent between individuals in the long term. We also show that, besides individual identity, couple identity is an important component explaining variation in behavior. This pattern can arise if behavior influences pair formation or if behavior is a consequence of a common environment affecting the couple. We show that, depending on the parental task, males or females are more consistent, suggesting that sexes are adopting different strategies of parental care allocation. We then investigate if there is a trade‐off between nest defense and chick feeding. We find that birds that defend more also feed the brood less often and suggest that testosterone might be the mechanism modulating this trade‐off. Lastly, we discuss the implications of our results to mate choice and highlight the need for studies linking parenting styles to fitness in swallow species.     

Assessing Short‐ and Long‐Term Repeatability and Stability of Personality in Captive Zebra Finches Using Longitudinal Data

Assessing behavioural consistency is crucial to understand the evolution of personality traits. In the present study, we examined the short‐ and long‐term repeatability and stability of two unrelated personality traits – exploratory tendencies and struggling rate – using captive female zebra finches ( Taeniopygia guttata). We performed two experimental sessions of behavioural tests with a 7‐mo interval, which represents up to one quarter of a zebra finch's life expectancy. We showed that, overall, exploratory tendencies and struggling rate were significantly repeatable in the short term. However, only exploratory tendencies were repeatable in the long term. We found interindividual differences in short‐term stability of exploratory tendencies, but not struggling rate, providing evidence for differences in intraindividual variability. In the long term, struggling rate significantly decreased between the two experimental sessions, whereas exploratory tendencies remained stable. Finally, the amount of interindividual variation measured at both sessions did not differ. Our results suggest that short‐ and long‐term repeatability and stability of personality may vary between individuals, depending on the behavioural trait under scrutiny. As a consequence, deducing personality from measures realized earlier in a subject's life should be performed with caution. We discuss the implications of inter‐ and intraindividual variation in personality consistency on individual fitness.     

Personality and urbanization: behavioural traits and <Emphasis Type="Italic">DRD4</Emphasis> SNP830 polymorphisms in great tits in Barcelona city

Most examples of adaptation to the urban environment relate to plasticity processes rather than to natural selection. Personality, however, defined as consistent individual differences in behaviour related to exploration, caution, and neophobia, is a good behavioural candidate character to study natural selection in relation to the urban habitat due to its heritable variation. The aim of this paper was to analyse variation in personality by comparing urban and forest great tits Parus major using standard tests of exploratory behaviour and boldness. We studied personality in 130 wild great tits captured in Barcelona city and nearby forests and found that urban birds were more explorative and bolder towards a novel object than forest birds. Genotype frequencies of the DRD4 SNP830 polymorphism, a gene region often associated with personality variation, varied significantly between forest and urban birds. Behavioural scores, however, were not correlated with this polymorphism in our population. Exploration scores correlated to boldness for forest birds but not for urban birds. Our findings suggest that the novel selection pressures of the urban environment favour the decoupling of behavioural traits that commonly form behavioural syndromes in the wild.     

Additive and nonadditive genetic variation in avian personality traits

Individuals of all vertebrate species differ consistently in their reactions to mildly stressful challenges. These typical reactions, described as personalities or coping strategies, have a clear genetic basis, but the structure of their inheritance in natural populations is almost unknown. We carried out a quantitative genetic analysis of two personality traits (exploration and boldness) and the combination of these two traits (early exploratory behaviour). This study was carried out on the lines resulting from a two-directional artificial selection experiment on early exploratory behaviour (EEB) of great tits (Parus major) originating from a wild population. In analyses using the original lines, reciprocal F(1) and reciprocal first backcross generations, additive, dominance, maternal effects ands sex-dependent expression of exploration, boldness and EEB were estimated. Both additive and dominant genetic effects were important determinants of phenotypic variation in exploratory behaviour and boldness. However, no sex-dependent expression was observed in either of these personality traits. These results are discussed with respect to the maintenance of genetic variation in personality traits, and the expected genetic structure of other behavioural and life history traits in general.     

Shyness–Boldness,but not Exploration,Predicts Glucocorticoid Stress Response in Richardson's Ground Squirrels (Urocitellus richardsonii)

The relationship between stress and personality has often been studied using captive animals in a laboratory context, yet less often in wild populations. Wild populations, however, may reveal aspects of the personality–stress relationship that laboratory‐based studies cannot. Here, we assessed the personality and stress hormone response of adult females within a free‐living population of Richardson's ground squirrels (Urocitellus richardsonii). Personality was assessed by quantifying individual responses to a novel object, and physiological stress was measured from faecal glucocorticoid metabolites. Principal component and principal component regression analyses were performed to determine whether the behavioural and endocrine measures were related. Based on these analyses, shyness–boldness was found to best predict glucocorticoid levels, in that individuals expressing the greatest vigilance in response to the novel object also had the highest measured concentrations of faecal glucocorticoids. Exploration, however, was independent of measured glucocorticoid levels, consistent with a multidimensional interpretation of non‐human animal personality.     

Consistent inter‐individual differences in common marmosets (Callithrix jacchus) in Boldness‐Shyness,Stress‐Activity,and Exploration‐Avoidance

The study of animal personality, defined as consistent inter‐individual differences in correlated behavioral traits stable throughout time and/or contexts, has recently become one of the fastest growing areas in animal biology, with study species ranging from insects to non‐human primates. The latter have, however, only occasionally been tested with standardized experiments. Instead their personality has usually been assessed using questionnaires. Therefore, this study aimed to test 21 common marmosets (Callithrix jacchus) living in three family groups, in five different experiments, and their corresponding controls. We found that behavioral differences between our animals were not only consistent over time, but also across different contexts. Moreover, the consistent behaviors formed a construct of four major non‐social personality components: Boldness‐Shyness in Foraging, Boldness‐Shyness in Predation, Stress‐Activity, and Exploration‐Avoidance. We found no sex or age differences in these components, but our results did reveal differences in Exploration‐Avoidance between the three family groups. As social environment can have a large influence on behavior of individuals, our results may suggest group‐level similarity in personality (i.e., “group personality”) in common marmosets, a species living in highly cohesive social groups. Am. J. Primatol. 78:961–973, 2016. © 2016 The Authors. American Journal of Primatology published by Wiley Periodicals, Inc.     

An experimental test of density‐dependent selection on temperament traits of activity,boldness and sociability

Temperament traits are seen in many animal species, and recent evolutionary models predict that they could be maintained by heterogeneous selection. We tested this prediction by examining density‐dependent selection in juvenile common lizards Zootoca vivipara scored for activity, boldness and sociability at birth and at the age of 1 year. We measured three key life‐history traits (juvenile survival, body growth rate and reproduction) and quantified selection in experimental populations at five density levels ranging from low to high values. We observed consistent individual differences for all behaviours on the short term, but only for activity and one boldness measure across the first year of life. At low density, growth selection favoured more sociable lizards, whereas viability selection favoured less active individuals. A significant negative correlational selection on activity and boldness existed for body growth rate irrespective of density. Thus, behavioural traits were characterized by limited ontogenic consistency, and natural selection was heterogeneous between density treatments and fitness traits. This confirms that density‐dependent selection plays an important role in the maintenance of individual differences in exploration‐activity and sociability.     

An approach to incorporate individual personality in modeling fish dispersal across in‐stream barriers

Animal personalities are an important factor that affects the dispersal of animals. In the context of aquatic species, dispersal modeling needs to consider that most freshwater ecosystems are highly fragmented by barriers reducing longitudinal connectivity. Previous research has incorporated such barriers into dispersal models under the neutral assumption that all migrating animals attempt to ascend at all times. Modeling dispersal of animals that do not perform trophic or reproductive migrations will be more realistic if it includes assumptions of which individuals attempt to overcome a barrier. We aimed to introduce personality into predictive modeling of whether a nonmigratory invasive freshwater fish (the round goby, Neogobius melanostomus) will disperse across an in‐stream barrier. To that end, we experimentally assayed the personalities of 259 individuals from invasion fronts and established round goby populations. Based on the population differences in boldness, asociability, and activity, we defined a priori thresholds with bolder, more asocial, and more active individuals having a higher likelihood of ascent. We then combined the personality thresholds with swimming speed data from the literature and in situ measurements of flow velocities in the barrier. The resulting binary logistic regression model revealed probabilities of crossing a barrier which depended not only on water flow and fish swimming speed but also on animal personalities. We conclude that risk assessment through predictive dispersal modeling across fragmented landscapes can be advanced by including personality traits as parameters. The inclusion of behavior into modeling the spread of invasive species can help to improve the accuracy of risk assessments.