Maintenance of Behavioral Variation under Predation Risk: Effects on Personality, Plasticity, and Predictability.

AbstractClassic evolutionary theory predicts that predation will shift trait means and erode variance within prey species; however, several studies indicate higher behavioral trait variance and trait integration in high-predation populations. These results come predominately from field-sampled animals comparing low- and high-predation sites and thus cannot isolate the role of predation from other ecological factors, including density effects arising from higher predation. Here, we study the role of predation on behavioral trait (co)variation in experimental populations of guppies (Poecilia reticulata) living with and without a benthic ambush predator (Jaguar cichlid) to better evaluate the role of predation and where density was equalized among replicates twice per year. At 2.5 years after introduction of the predators (∼10 overlapping generations), 40 males were sampled from each of the six replicate populations and extensively assayed for activity rates, water column use, and latency to feed following disturbance. Individual variation was pronounced in both treatments, with substantial individual variation in means, temporal plasticity, and predictability (inverse residual variance). Predators had little effect on mean behavior, although there was some evidence for greater use of the upper water column in predator-exposed fish. There was greater variance among individuals in water column use in predator-exposed fish, and they habituated more quickly over time; individuals higher in the water column fed slower and had a reduced positive correlation with activity, although again this effect was time specific. Predators also affected the integration of personality and plasticity-among-individual variances in water column use increased, and those in activity decreased, through time-which was absent in controls. Our results contrast with the extensive guppy literature showing rapid evolution in trait means, demonstrating either increases or maintenance of behavioral variance under predation.

Contaminant-by-environment interactive effects on animal behavior in the context of global change: Evidence from avian behavioral ecotoxicology.

The potential for chemical contaminant exposure to interact with other stressors to affect animal behavioral responses to environmental variability is of mounting concern in the context of anthropogenic environmental change. We systematically reviewed the avian literature to evaluate evidence for contaminant-by-environment interactive effects on animal behavior, as birds are prominent models in behavioral ecotoxicology and global change research. We found that only 17 of 156 (10.9 %) avian behavioral ecotoxicological studies have explored contaminant-by-environment interactions. However, 13 (76.5 %) have found evidence for interactive effects, suggesting that contaminant-by-environment interactive effects on behavior are understudied but important. We draw on our review to develop a conceptual framework to understand such interactive effects from a behavioral reaction norm perspective. Our framework highlights four patterns in reaction norm shapes that can underlie contaminant-by-environment interactive effects on behavior, termed exacerbation, inhibition, mitigation and convergence. First, contamination can render individuals unable to maintain critical behaviors across gradients in additional stressors, exacerbating behavioral change (reaction norms steeper) and generating synergy. Second, contamination can inhibit behavioral adjustment to other stressors, antagonizing behavioral plasticity (reaction norms shallower). Third, a second stressor can mitigate (antagonize) toxicological effects of contamination, causing steeper reaction norms in highly contaminated individuals, with improvement of performance upon exposure to additional stress. Fourth, contamination can limit behavioral plasticity in response to permissive conditions, such that performance of more and less contaminated individuals converges under more stressful conditions. Diverse mechanisms might underlie such shape differences in reaction norms, including combined effects of contaminants and other stressors on endocrinology, energy balance, sensory systems, and physiological and cognitive limits. To encourage more research, we outline how the types of contaminant-by-environment interactive effects proposed in our framework might operate across multiple behavioral domains. We conclude by leveraging our review and framework to suggest priorities for future research.

The evolution of social philopatry in female primates.

The transition from solitary life to sociality is considered one of the major transitions in evolution. In primates, this transition is currently not well understood. Traditional verbal models appear insufficient to unravel the complex interplay of environmental and demographic factors involved in the evolution of primate sociality, and recent phylogenetic reconstructions have produced conflicting results. We therefore analyze a theoretical model for the evolution of female social philopatry that sheds new light on the question why most primates live in groups. In individual-based simulations, we study the evolution of dispersal strategies of both resident females and their offspring. The model reveals that social philopatry can evolve through kin selection, even if retention of offspring is costly in terms of within-group resource competition and provides no direct benefits. Our model supports the role of predator avoidance as a selective pressure for group-living in primates, but it also suggests that a second benefit of group-living, communal resource defense, might be required to trigger the evolution of sizable groups. Lastly, our model reveals that seemingly small differences in demographic parameters can have profound effects on primate social evolution.

A guide for studying among-individual behavioral variation from movement data in the wild.

Animal tracking and biologging devices record large amounts of data on individual movement behaviors in natural environments. In these data, movement ecologists often view unexplained variation around the mean as "noise" when studying patterns at the population level. In the field of behavioral ecology, however, focus has shifted from population means to the biological underpinnings of variation around means. Specifically, behavioral ecologists use repeated measures of individual behavior to partition behavioral variability into intrinsic among-individual variation and reversible behavioral plasticity and to quantify: a) individual variation in behavioral types (i.e. different average behavioral expression), b) individual variation in behavioral plasticity (i.e. different responsiveness of individuals to environmental gradients), c) individual variation in behavioral predictability (i.e. different residual within-individual variability of behavior around the mean), and d) correlations among these components and correlations in suites of behaviors, called 'behavioral syndromes'. We here suggest that partitioning behavioral variability in animal movements will further the integration of movement ecology with other fields of behavioral ecology. We provide a literature review illustrating that individual differences in movement behaviors are insightful for wildlife and conservation studies and give recommendations regarding the data required for addressing such questions. In the accompanying R tutorial we provide a guide to the statistical approaches quantifying the different aspects of among-individual variation. We use movement data from 35 African elephants and show that elephants differ in a) their average behavior for three common movement behaviors, b) the rate at which they adjusted movement over a temporal gradient, and c) their behavioral predictability (ranging from more to less predictable individuals). Finally, two of the three movement behaviors were correlated into a behavioral syndrome (d), with farther moving individuals having shorter mean residence times. Though not explicitly tested here, individual differences in movement and predictability can affect an individual's risk to be hunted or poached and could therefore open new avenues for conservation biologists to assess population viability. We hope that this review, tutorial, and worked example will encourage movement ecologists to examine the biology of individual variation in animal movements hidden behind the population mean.

Repeatability of thermal reaction norms for spontaneous locomotor activity in juvenile newts.

Locomotor activity is a major attribute of animals. Although this trait determines important ecological processes, such as dispersal and species interactions, the sources of its variation are not fully understood. We examined the influence of body temperature (13, 18, 23, and 28 °C) and individual identity on spontaneous locomotor activity in juvenile alpine newts, Ichthyosaura alpestris, over three consecutive weeks. Locomotor activity was characterized by four parameters: distance covered, mean velocity, frequency of movements, and total activity rate (all directional and non-directional movements). Apart from total activity rate, thermal reaction norms for locomotor parameters had convex or concave curvilinear shapes. During the first trial series, i.e. across the four body temperatures that were tested, individual identity explained less variation in thermal reaction norms than during the second series. Individual means, i.e. the vertical positions of individual thermal reaction norms, were repeatable between trial series in all locomotor activity parameters but the frequency of movements. We conclude that spontaneous locomotor activity is a complex trait, which can be characterized by several parameters with varying individual repeatability and thermal dependency. This information should be considered for planning further locomotor activity experiments, conservation strategies, and modeling ectotherm responses to climate change.

Dynamics of among-individual behavioral variation over adult lifespan in a wild insect.

Investigating patterns of among and within-individual trait variation in populations is essential to understanding how selection shapes phenotypes. Behavior is often the most flexible aspect of the phenotype, and to understand how it is affected by selection, we need to examine how consistent individuals are. However, it is not well understood whether among-individual differences tend to remain consistent over lifetimes, or whether the behavior of individuals relative to one another varies over time. We examined the dynamics of 4 behavioral traits (tendency to leave a refuge, shyness, activity, and exploration) in a wild population of field crickets (Gryllus campestris). We tagged individuals and then temporarily removed them from their natural environment and tested them under laboratory conditions. All 4 traits showed among-individual variance in mean levels of expression across the adult lifespan, but no significant differences in how rapidly expression changed with age. For all traits, among-individual variance increased as individuals got older. Our findings reveal seldom examined changes in variance components over the adult lifetime of wild individuals. Such changes will have important implications for the relationship between behavioral traits, life-histories, and fitness and the consequences of selection on wild individuals.