Lesley J. Morrell

From Hawks and Doves to Self‐Consistent Games of Territorial Behavior

Explaining the “prior‐residence effect” (automatic owner status of individuals who arrived first in an area) was one of the very first applications of game theory in animal behavior. These models, however, predict paradoxical solutions where intruders always win, with no satisfactory explanation for the absence of such cases in nature. We propose a solution based on new developments in evolutionary game theory. A self‐consistent model with feedbacks between individual behavior and population dynamics produces qualitatively different frequency‐dependent selection on intruders (floaters) than on territory owners. Starting with an ancestral population with no respect for ownership, the most likely evolutionary end point is complete or partial respect. Conventional rules of conflict resolution thus can rely on “uncorrelated asymmetries” without differences in resource‐holding power or territory value, although they will be strengthened by such differences. We also review the empirical literature on animal contests, testing whether asymmetries in resource‐holding power are required to explain the observations. Despite much empirical effort, results remain inconclusive, because experiments are often unable to distinguish between the motivation of individuals to fight and the behavioral outcome of a contest. To help arrive at conclusive answers, we suggest a standardized empirical approach to quantify prior‐residence effects.

Predation risk as a driving force for sexual segregation: a cross-population comparison.

Sexual segregation is widespread throughout the animal kingdom. Although a number of hypotheses have been proposed to account for observed patterns, the generality of the mechanisms remains debated. One possible reason for this is the focus on segregation patterns in large mammals such as ungulates, where the majority of studies are descriptions of a single population. Here, we present the results of a cross‐population comparison of patterns of sexual segregation in the Trinidadian guppy, Poecilia reticulata. We relate observed patterns to experimental quantification of predation risk and sexual harassment of females by males in eight populations. We find that the degree of segregation increases with predation risk, with deeper waters becoming increasingly female biased. Furthermore, we observed that levels of male harassment are lower in deeper water but only in those rivers that contain major guppy predators. We conclude that sexual segregation in guppies is consistent with the predation risk hypothesis: sexual segregation results from a combination of predation risk driving males (the more vulnerable sex) into less risky habitats and females gaining benefits of reduced sexual harassment by remaining in high‐predation environments.

Social network correlates of food availability in an endangered population of killer whales, Orcinus orca

For the majority of social species, group composition is dynamic, and individuals are interconnected in a heterogeneous social network. Social network structure has far-reaching implications for the ecology of individuals and populations. However, we have little understanding of how ecological variables shape this structure. We used a long-term data set (1984e2007) to examine the relationship between food availability and social network structure in the endangered southern resident killer whales. During the summer months individuals in this population feed primarily on chinook salmon, Oncorhynchus tshawytscha, which show annual variation in abundance. We tested the hypothesis that temporal variation in chinook salmon will correlate with variation in social network structure. Using a null model that controlled for population demography, group size and sampling effort, we found a significant relationship between the connectivity of the social network and salmon abundance, with a more interconnected social network in years of high salmon abundance. Our results demonstrate that resource availability may be an important determinant of social network structure. Given the central importance of the social network for population processes such as the maintenance of cooperation and the transmission of information and disease, a change in social network structure caused by a change in food availability may have significant ecological and evolutionary consequences. 2012 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Why are small males aggressive

Aggression is ubiquitous in the animal kingdom, whenever the interests of individuals conflict. In contests between animals, the larger opponent is often victorious. However, counter intuitively, an individual that has little chance of winning (generally smaller individuals) sometimes initiates contests. A number of hypotheses have been put forward to explain this behaviour, including the ‘desperado effect’ according to which, the likely losers initiate aggression due to lack of alternative options. An alternative explanation suggested recently is that likely losers attack due to an error in perception: they mistakenly perceive their chances of winning as being greater than they are. We show that explaining the apparently maladaptive aggression initiated by the likely loser can be explained on purely economic grounds, without requiring either the desperado effect or perception errors. Using a game-theoretical model, we show that if smaller individuals can accurately assess their chance of winning, if this chance is less than, but close to, a half, and if resources are scarce (or the contested resource is of relatively low value), they are predicted to be as aggressive as their larger opponents. In addition, when resources are abundant, and small individuals have some chance of winning, they may be more aggressive than their larger opponents, as it may benefit larger individuals to avoid the costs of fighting and seek alternative uncontested resources.

Predation Risk Shapes Social Networks in Fission-Fusion Populations

Predation risk is often associated with group formation in prey, but recent advances in methods for analysing the social structure of animal societies make it possible to quantify the effects of risk on the complex dynamics of spatial and temporal organisation. In this paper we use social network analysis to investigate the impact of variation in predation risk on the social structure of guppy shoals and the frequency and duration of shoal splitting (fission) and merging (fusion) events. Our analyses revealed that variation in the level of predation risk was associated with divergent social dynamics, with fish in high-risk populations displaying a greater number of associations with overall greater strength and connectedness than those from low-risk sites. Temporal patterns of organisation also differed according to predation risk, with fission events more likely to occur over two short time periods (5 minutes and 20 minutes) in low-predation fish and over longer time scales (>1.5 hours) in high-predation fish. Our findings suggest that predation risk influences the fine-scale social structure of prey populations and that the temporal aspects of organisation play a key role in defining social systems.

The Effect of Prey Density on Predators: Conspicuousness and Attack Success Are Sensitive to Spatial Scale

In contrast to the numerous studies that have examined the response of predators to prey group size, little is known about how prey density affects prey detection and the accuracy of attacks. We demonstrate that increasing the density of Daphnia magna swarms increases conspicuousness to a natural predator, the three‐spined stickleback. Denser areas of groups were more conspicuous, as the fish attacked prey in denser parts of the group than would be expected if they attacked the nearest prey upon entering the feeding chamber. The spatial error of attacks also increased with the density around the target; hence, different stages of predation (searching for vs. successfully attacking prey) seem to select for opposing responses to prey density. However, whereas the effect of density on target selection only occurred using a global measure of density (average interindividual distance), the effect on attack error was only significant using a local measure of density (Voronoi polygon area). We believe this effect of spatial scale reflects the reduction in the number of prey in the visual field of the predator as an attack progresses, providing a perceptual basis for the importance of spatial scale in density‐dependent processes.

RISK‐SENSITIVE ANTIPREDATOR BEHAVIOR IN THE TRINIDADIAN GUPPY, POECILIA RETICULATA

The comparative approach has become a powerful tool for understanding how predation has shaped prey behavior. In this study we recorded the occurrence of common aquatic predator species and their densities in seven natural populations of Trinidadian guppies, Poecilia reticulata. We then exposed shoals of guppies from each of these populations to a series of predator treatments. Predator treatments differed in the species of predator used (pike cichlids, Crenicichla frenata; rivulus, Rivulus hartii; and freshwater prawns, Macrobrachium carcinus) and thus in the level of risk posed. We recorded the antipredator responses of guppies in each of these predator treatments. The strength of antipredator behavior shown by guppies was affected by both the type of predator they were exposed to and the level of predation risk they experienced naturally in the wild. Importantly, we found that guppies from high-risk populations showed a heightened response, compared to those from lower risk populations, only when exposed to the predator species that posed the greatest risk. Our results show the importance of individual predator species in shaping the behavioral traits of prey species at the population level. This has implications for prey movement between habitats that are geographically close but differ greatly in predator fauna.

Association patterns and foraging behaviour in natural and artificial guppy shoals

Animal groups are often nonrandom assemblages of individuals that tend to be assorted by factors such as sex, body size, relatedness and familiarity. Laboratory studies using fish have shown that familiarity among shoal members confers a number of benefits to individuals, such as increased foraging success. However, it is unclear whether fish in natural shoals obtain these benefits through association with familiars. We investigated whether naturally occurring shoals of guppies, Poecilia reticulata, are more adept at learning a novel foraging task than artificial (in which we selected shoal members randomly) shoals. We used social network analysis to compare the structures of natural and artificial shoals and examined whether shoal organization predicts patterns of foraging behaviour. Fish in natural shoals benefited from increased success in the novel foraging task compared with fish in artificial shoals. Individuals in natural shoals showed a reduced latency to approach the novel feeder, followed more and formed smaller subgroups compared to artificial shoals. Our findings show that fish in natural shoals do gain foraging benefits and that this may be facilitated by a reduced perception of risk among familiarized individuals and/or enhanced social learning mediated by following other individuals and small group sizes. Although the structure of shoals was stable over time, we found no direct relationship between shoal social structure and patterns of foraging behaviour.

Balancing the Dilution and Oddity Effects: Decisions Depend on Body Size

Background Grouping behaviour, common across the animal kingdom, is known to reduce an individuals risk of predation; particularly through dilution of individual risk and predator confusion (predator inability to single out an individual for attack). Theory predicts greater risk of predation to individuals more conspicuous to predators by difference in appearance from the group (the ‘oddity’ effect). Thus, animals should choose group mates close in appearance to themselves (eg. similar size), whilst also choosing a large group. Methodology and Principal Findings We used the Trinidadian guppy (Poecilia reticulata), a well known model species of group-living freshwater fish, in a series of binary choice trials investigating the outcome of conflict between preferences for large and phenotypically matched groups along a predation risk gradient. We found body-size dependent differences in the resultant social decisions. Large fish preferred shoaling with size-matched individuals, while small fish demonstrated no preference. There was a trend towards reduced preferences for the matched shoal under increased predation risk. Small fish were more active than large fish, moving between shoals more frequently. Activity levels increased as predation risk decreased. We found no effect of unmatched shoal size on preferences or activity. Conclusions and Significance Our results suggest that predation risk and individual body size act together to influence shoaling decisions. Oddity was more important for large than small fish, reducing in importance at higher predation risks. Dilution was potentially of limited importance at these shoal sizes. Activity levels may relate to how much sampling of each shoal was needed by the test fish during decision making. Predation pressure may select for better decision makers to survive to larger size, or that older, larger fish have learned to make shoaling decisions more efficiently, and this, combined with their size relative to shoal-mates, and attractiveness as prey items influences shoaling decisions.

Behavioural consequences of sensory plasticity in guppies

Sensory plasticity, whereby individuals compensate for sensory deprivation in one sense by an improvement in the performance of an alternative sense, is a well-documented phenomenon in nature. Despite this, the behavioural and ecological consequences of sensory plasticity have not been addressed. Here we show experimentally that some components (vision and chemoreception) of the sensory system of guppies are developmentally plastic, and that this plasticity has important consequences for foraging behaviour. Guppies reared under low light conditions had a significantly stronger response to chemical food cues encountered in isolation than fish reared at higher light levels. Conversely, they exhibited a weaker response to visual-only cues. When visual and olfactory/gustatory cues were presented together, no difference between the strength of response for fish reared at different light intensities was evident. Our data suggest that guppies can compensate for experience of a visually poor, low light environment via a sensory switch from vision to olfaction/gustation. This switch from sight to chemoreception may allow individuals to carry out the foraging behaviour that is essential to their survival in a visually poor environment. These considerations are especially important given the increasing frequency of anthropogenic changes to ecosystems. Compensatory phenotypic plasticity as demonstrated by our study may provide a hitherto unconsidered buffer that could allow animals to perform fundamental behaviours in the face of considerable change to the sensory environment.