Behavioral repeatability and personality in pit-building antlion larvae under differing environmental contexts

Over the last decades, there has been growing interest among behavioral ecologists in exploring animal personalities. However, while the foraging behavior of active foragers has been extensively studied, only little is known about that of sit-and-wait predators within the personality framework. We investigated the existence of repeatability and personality in pit-building antlion larvae in the context of foraging (pit construction) and habitat selection (relocation distance and direction) over time and under three environmental contexts: thermal conditions, sand depth, and soil type. Over time, repeatability was much stronger for relocation distance than for movement directionality. Additionally, we observed positive correlations across the two levels of sand depth and soil type but not between thermal conditions. Change in substrate type may induce faster decision-making in these sand-dwelling insects or could be perceived by such insects as a more drastic alteration in their habitat. We suggest that different individuals indeed possess distinct personalities. We also suggest that repeatability should be measured at two levels: the amount of energy expenditure (distances and pit construction) and the pattern of energy expenditure (directionality). Finally, our study illustrates how differing environmental conditions can result in differing levels of plasticity, while largely preserving individual personalities.     

Habitat structure helps guide the emergence of colony-level personality in social spiders

Studying the environmental factors that guide the emergence of collective behaviors is instrumental to understanding the ecology and evolution of animal societies. Although recent work has provided insights into the demographic factors that influence inter-colony variation in collective behavior (i.e., colony-level personality or collective personality), relatively few studies have investigated how the physical environment (e.g., habitat structure) affects colony-level personality. Here, we study the emergence of collective personality in prey capture behavior in the social spider, Stegodyphus dumicola. We measured collective prey capture behavior four times over 36 days in a classic repeated measures design. We used four different artificial habitat (web support) structures in three different treatments: habitat structure was either (1) fixed and undisturbed, (2) disturbed with a complete removal of webbing between each measurement, or (3) disturbed with changes of habitat structure between each measurement. Our results revealed that repeatability in colony-level personality was retained as long as habitat structure was not altered. However, the repeatability of colony-level personality declined precipitously when groups were forced to build their webs on novel habitat structures. Furthermore, habitat structure affected collective capture behavior, that is, latency to attack and the number of attackers differed among colonies on different habitat structures. Collectively, our data demonstrate that habitat structure is instrumental in shaping both the mean and repeatability of the collective behavior of colonies and may influence overall foraging success.     

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.     

Risk-taking behavior in the lesser wax moth: disentangling within- and between-individual variation

Behaviors that appear to be plastic may well be determined by environmental influences during development. Being able to produce a wide range of variants of one kind of behavior, e.g., a very short and a very long response time to a stimulus under different environmental conditions, can be described as behavioral plasticity. How such behavioral reaction norms develop for individuals is poorly understood, but several factors are likely to play a role. We investigated what factors may affect how the risk-taking behavior of the lesser wax moth, Achroia grisella, is shaped during ontogeny. We manipulated larval density to represent the potential intensity of future competition for females in a lek of males, determined adult moths’ reaction to predator signals, and tested for plasticity in the silence response, i.e., the acoustic evasion behavior of the moths during the experiments. While we found no effect of larval density on either the probability or the duration of the silence response, 11 % of the variance in duration could be explained by differences between families, and 30 % of the variance was the result of differences between individuals. We found evidence for habituation to the predator signal, clearly indicating that the silence response is a plastic-enough trait to be adjustable to the immediate environment. These results suggest that the degree to which individuals take risks in the context of acoustic signaling depends more on the immediate context and, possibly, genetic differentiation than it is a product of adaptive developmental plasticity.     

Network position: a key component in the characterization of social personality types

In recent years, animal social interactions have received much attention in terms of personality research (e.g. aggressive or cooperative interactions). However, other components of social behaviour such as those describing the intensity, frequency, directedness and individual repeatability of interactions in animal groups have largely been neglected. Network analysis offers a valuable opportunity to characterize individual consistency of traits in labile social groups and therein provide novel insights to personality research in ways previously not possible using traditional techniques. Should individual network positions be consistently different between individuals under changing conditions, they might reflect expressions of an individual's personality. Here, we discuss a conceptual framework for using network analyses to infer the presence of individual differences and present a statistical test based on randomization techniques for testing the consistency of network positions in individuals. The statistical tools presented are useful because if particular individuals consistently occupy key positions in social networks, then this is also likely to have consequences for their fitness as well as for that of others in the population. These consequences may be particularly significant since individual network position has been shown to be important for the transmission of diseases, socially learnt information and genetic material between individuals and populations.     

Bibliometric investigation of the integration of animal personality in conservation contexts

Consistent individual differences in behavior, commonly termed animal personality, are a widespread phenomenon across taxa that have important consequences for fitness, natural selection, and trophic interactions. Animal personality research may prove useful in several conservation contexts, but which contexts remains to be determined. We conducted a structured literature review of 654 studies identified by combining search terms for animal personality and various conservation subfields. We scored the relevance of personality and conservation issues for each study to identify which studies meaningfully integrated the 2 fields as opposed to surface-level connections or vague allusions. We found a taxonomic bias toward mammals (29% of all studies). Very few amphibian or reptile studies applied personality research to conservation issues (6% each). Climate change (21%), invasive species (15%), and captive breeding and reintroduction (13%) were the most abundant conservation subfields that occurred in our search, though a substantial proportion of these papers weakly integrated conservation and animal personality (climate change 54%, invasive species 51%, captive breeding and reintroduction 40%). Based on our results, we recommend that researchers strive for consistent and broadly applicable terminology when describing consistent behavioral differences to minimize confusion and improve the searchability of research. We identify several gaps in the literature that appear to be promising and fruitful avenues for future research, such as disease transmission as a function of sociability or exploration as a driver of space use in protected areas. Practitioners can begin informing future conservation efforts with knowledge gained from animal personality research.     

Patterns in swimming by a scyphomedusa: a novel approach to quantifying behavior in individuals

Behavior is commonly studied at the group level using several individuals, but there is increasing evidence that the behavior of a few individuals often has a disproportionate effect on the response of a population to its environment. The present study used a suite of statistical techniques, random series analysis, analysis of variance, spectral analysis, and goodness-of-fit tests of frequency histograms, to quantitatively describe the time-dependent changes in individual behavior. Each technique reveals a different facet of the behavior and, when simultaneously applied to the data, distinguishes significant differences among the behaviors of several individuals. The approach was developed and tested on the swimming behavior of four specimens of the scyphomedusa Aurelia aurita (Linnaeus, 1758), which were observed for 19 days, beginning 16 January 1998, and videotaped under identical environmental conditions during that period. The analyses showed that each medusa swam in a unique pattern, varying swimming at characteristic frequencies. Application of the approach to individual-based numerical modeling, to the role of endogenous stimuli in the behavioral repertoire, and to in situ studies of animal behavior is discussed.Communicated by J.P. Grassle, New Brunswick     

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.     

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.     

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.     

Economic and Social Dimensions of Environmental Behavior: Balancing Conservation and Development in Bhutan

Abstract: One of the primary approaches to environmental conservation emphasizes economic development. This conservation‐and‐development approach often ignores how development affects sociocultural characteristics that may motivate environmental behaviors (actions that actively benefit or limit one's negative impacts on the environment). Evolutionary anthropologists espouse a theoretical perspective that supports the conservation‐and‐development approach. Others believe sociocultural factors are the foundation of environmental behavior and worry that development will erode the values and norms that may shape such behavior. My research assistants and I surveyed 170 individuals from eight villages in two communities in Bhutan to explore whether economic (wealth, market integration) or social (religious behaviors, environmental values, social capital) factors are better indicators of environmental behavior. I used multilevel modeling to analyze use of fuelwood, use of agricultural chemicals, and tree planting, and to determine whether social norms were associated with these behaviors. Although economic factors were more often associated with these behaviors than social factors, local conditions and control variables were the best indicators of behaviors. Furthermore, economic factors were not always associated with positive environmental outcomes. Instead, farmers attempted to make the best economic decisions given their circumstances rather than seeking to conserve resources. Although religion was not a strong predictor of any of the behaviors I examined, I found evidence that the understanding of Buddhist philosophy is growing, which suggests that social factors may play a more prominent role as Bhutan's development progresses. My results highlight the need for conservation planners to be aware of local conditions when planning and implementing policies aimed at motivating environmental behaviors and that economic and social motivations for conservation may not be mutually exclusive.     

Population Constraints Associated with the Use of Black Rhinos as an Umbrella Species for Desert Herbivores

The numerous tactics used to conserve biodiversity include the designation of protected areas, political change, and research and education, the latter involving paradigms such as insular biogeography and the "umbrella species concept." In Namibia lands removed from national park status in 1970 and currently under the jurisdiction of indigenous people now contain one of the few unfenced populations of black rhinos (Diceros bicornis) remaining in Africa. Theory predicts that the protection of umbrella species will ensure the survival of other biota that require(s) less space. To gauge how well biodiversity might be retained by examining the spatial needs of a small population of black rhinos, I used data gathered under various ecological conditions to estimate mean and minimum population sizes of six large herbivores of the Namib Desert ranging in size from giraffe (Giraffa camelopardalis) to springbok (Antidorcas marsupialis) and ostrich (Struthio camelus). My results indicate that annual differences in rainfall, both within and between seasons, resulted in wide fluctuations in herbivore population sizes for all species except rhino. Although other herbivores switched to areas of higher rainfall, rhinos did not. The data suggest that under conditions of extreme environmental variance the space used by rhinos alone was unlikely to assure the existence of populations of other species in excess of 250 individuals. Fifty percent of the species failed to exceed 150 individuals 50% of the time and one third of the species never attained populations in excess of 50 individuals. However, by employing assumptions about the spatial needs of rhino populations numbering up to 100 individuals, the mean minimum population sizes attained by any of four desert herbivores is 535. A future challenge in using rhinos and other large-bodied species as umbrellas for organisms of either similar or dissimilar trophic levels will be the refinement of estimates of population viability.     

Exploring the effects of individual traits and within-colony variation on task differentiation and collective behavior in a desert social spider

Social animals are extraordinarily diverse and ecologically abundant. In understanding the success of complex animal societies, task differentiation has been identified as a central mechanism underlying the emergence and performance of adaptive collective behaviors. In this study, we explore how individual differences in behavior and body size determine task allocation in the social spider Stegodyphus dumicola. We found that individuals with high body condition indices were less likely to participate in prey capture, and individuals’ tendency to engage in prey capture was not associated with either their behavioral traits or body size. No traits were associated with individuals’ propensity to participation in web repair, but small individuals were more likely to engage in standard web-building. We also discovered consistent, differences among colonies in their collective behavior (i.e., colony-level personality). At the colony level, within-colony variation in behavior (aggressiveness) and body size were positively associated with aggressive foraging behavior. Together, our findings reveal a subtly complex relationship between individual variation and collective behavior in this species. We close by comparing the relationship between individual variation and social organization in nine species of social spider. We conclude that intraspecific variation is a major force behind the social organization of multiple independently derived lineages of social spider.     

Adaptive phenotypic differentiation across the intertidal gradient in the alga Silvetia compressa

Populations of intertidal species span a steep environmental gradient driven by differences in emersion time. In spite of strong differential selection on traits related to this gradient, the small spatial scale over which differences occur may prevent local adaptation, and instead may favor a single intermediate phenotype, or nongenetic mechanisms of differentiation. Here I examine whether a common macroalga, Silvetia compressa, exhibits phenotypic differentiation across the intertidal gradient and evaluate how local adaptation, developmental plasticity, and maternal effects may interact to shape individual phenotypes. Reciprocal transplants of both adults and embryos showed a "home-height advantage" in two of the three populations tested. In laboratory trials, the progeny of upper-limit individuals survived exposure to air significantly better than lower-limit progeny from the same population. I compared the emersion tolerance of full-sib families generated from gametes produced in the field to those produced under common garden conditions. The relative advantage of upper-limit lineages was robust to maternal environment during gametogenesis; this pattern is consistent with genetic differentiation. The possible role of local adaptation has historically been ignored in studies of intertidal zonation. In S. compressa, phenotypic differentiation may have important consequences for vertical range, both within and among sites.     

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.     

Can modeling improve estimation of desert tortoise population densities?

The federally listed desert tortoise (Gopherus agassizii) is currently monitored using distance sampling to estimate population densities. Distance sampling, as with many other techniques for estimating population density, assumes that it is possible to quantify the proportion of animals available to be counted in any census. Because desert tortoises spend much of their life in burrows, and the proportion of tortoises in burrows at any time can be extremely variable, this assumption is difficult to meet. This proportion of animals available to be counted is used as a correction factor (g0) in distance sampling and has been estimated from daily censuses of small populations of tortoises (6-12 individuals). These censuses are costly and produce imprecise estimates of go due to small sample sizes. We used data on tortoise activity from a large (N = 150) experimental population to model activity as a function of the biophysical attributes of the environment, but these models did not improve the precision of estimates from the focal populations. Thus, to evaluate how much of the variance in tortoise activity is apparently not predictable, we assessed whether activity on any particular day can predict activity on subsequent days with essentially identical environmental conditions. Tortoise activity was only weakly correlated on consecutive days, indicating that behavior was not repeatable or consistent among days with similar physical environments.     

Sex-specific plasticity in brain morphology depends on social environment of the guppy, Poecilia reticulata

The vertebrate brain is a remarkably plastic organ, which responds quickly to environmental changes. However, to date, studies investigating plasticity in brain morphology have focused mostly on the physical properties of the surrounding environment, and little is known about brain plasticity in response to the social environment. Moreover, sex differences in brain plasticity remain virtually unexplored. Here, we tested how the social environment influenced brain morphology in adult males and females using experimental manipulation of the sex composition of social pairs (same sex vs. mixed sex) in the guppy (Poecilia reticulata). We detected substantial sex-specific plasticity in both the overall brain size (controlling for body size) and separate brain structures. The brain size was larger in males that interacted with females, and female optic tectum was larger in female-only groups. Overall, females had larger olfactory bulbs and cerebellum in comparison to males. While net sexual dimorphism in the brain structure can be explained in light of the known differences in boldness and foraging behaviour between the sexes, our results also support that cognitive demands associated with courtship behaviour can lead to plastic changes in the brain size. Our findings demonstrate that not only social environment can generate rapid, plastic responses in the vertebrate brain but also that such responses can depend strongly on sex.     

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.     

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.     

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.