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Lay Summary Changes in space and group size for water striders (aquatic bugs) induced striking, reversible changes in mating behaviors within days or even hours. In large groups, no male could dominate the rest. Instead, males harassed females, drove them into hiding, mated frequently, and typically guarded females for several hours after mating. In contrast, in small groups, an alpha male often emerged, driving other males into hiding, mating only occasionally, and rarely harassing or guarding females.

International audience; A key challenge in invasion biology is identifying characteristics that allow some species to be repeatedly successful at invading novel environments. Invasions can often be disproportionately driven by a single sex, with differences in behavioural mechanisms between the sexes potentially underlying sex-biased invasiveness. Here, we took an animal personality approach to study the behaviour of two repeatedly successful congeneric invasive species, the western mosquitofish, Gambusia affinis, and the eastern mosquitofish, Gambusia holbrooki. In each species, we investigated whether males and females shared common personality traits (i.e. behavioural types and behavioural syndromes), with the aim of identifying possible behavioural mechanisms that could help explain why mosquitofish invasions are often characterised by sex-biased founder populations. We found sex-dependent personality, although sex differences varied between species. Specifically, male G. affinis were bolder and less social than female G. affinis, whereas we found no behavioural type differences between the sexes in G. holbrooki. We also found a consistent correlation between boldness and exploration in both sexes within G. affinis, but this correlation was weak in G. holbrooki. Finally, exploration was also correlated with sociability in male G. affinis, but not in females. Our results suggest that behavioural tendencies may diverge, both among species and between the sexes, because of adaptation experienced during different invasion pathways. Broadly, identifying the behavioural mechanisms that predict an individual's 'invasiveness' may be difficult to tease apart between species because each invasion is characterised by different abiotic and biotic interactions that likely require different suites of behaviours. Future studies are needed to elucidate whether, in fact, personality variation between the sexes can mediate the occurrence of sex-biased invasions.

International audience; Mosquitofish (Gambusia affinis) are a widespread, invasive species that frequently colonize habitats where they might encounter novel predators. Earlier work showed that asocial mosquitofish disperse more readily than social fish. Initial colonists to newly invaded, low density sites should thus be relatively asocial. Here, we tested the hypothesis that asocial mosquitofish should survive better than social fish when exposed to predators at low mosquitofish density. We used standardized behavioural assays to quantify the individual behavioral type (boldness, sociability, activity, exploratory tendency) of 224 mosquitofish, and then exposed them to predators in small groups. As predicted, asocial individuals survived exposure to predators better than social individuals. In addition, while body mass per se did not affect survival, males survived predators better than females. Overall, this study provides an early corroboration of the general prediction that behavioural types that disperse more readily might also be better at coping with predators at low density.

In systems of strong sexual conflict, male harassment can constrain female feeding, which can limit fecundity. Female responses to harassment can depend on the social context or differ between individuals, and could be based on social or intrinsic behavioral cues at different spatio-temporal scales. Using experimental groupings of stream water striders (Aquarius remigis), we examined individual behavioral differences and effects of sex ratio on 4 aspects of female behavior: 1) habitat use and 2) activity, which lead to differences in 3) feeding rates and 4) mating rates. We compared effects of the immediate social environment, immediate female habitat use and activity, and consistent female behavioral tendencies (personalities) on feeding and mating probabilities, and asked if population sex ratio affected the relationships among these behaviors. We found that individual females did differ significantly in all 4 behaviors. Population sex ratio strongly influenced average female habitat use, feeding, and mating behaviors, and female feeding and mating behaviors were predicted by a combination of moment-to-moment female behavioral state, moment-to-moment social factors, and consistent individual female behavioral differences. Furthermore, habitat use tendencies correlated significantly with activity tendencies, and habitat use and activity tendencies predicted mating probabilities, but not feeding probabilities. Our study elucidates the specific individual-level behavioral mechanisms that lead to observed population-level patterns and emphasizes the benefits of studying behavior at multiple spatial and temporal scales.

Dispersers are often not a random draw from a population, dispersal propensity being conditional on individual phenotypic traits and local contexts. This non-randomness consequently results in phenotypic differences between dispersers and non-dispersers and, in the context of biological invasions, in an invasion front made of individuals with a biased phenotype. This bias of phenotypes at the front may subsequently modulate the strength of ecological effects of an invasive species on invaded communities. We recently demonstrated that more asocial mosquitofish (Gambusia affinis), one of the 100 worst invasive species, disperse further, suggesting a sociability-biased invasion front. As behavioural types are related to the strength of interspecific interactions, an invasion by a biased subset of individuals should have important ecological implications for native communities. Here, we tested the impact of phenotypic biases in dispersing individuals (relative to non-dispersers) on prey communities in experimental mesocosms. We show that dispersers reduce prey abundance more than do non-dispersers during the first 4 weeks after introduction, and that the disperser's social types are likely drivers of these differences. These differences in prey communities disappeared after 8 weeks suggesting prey community resilience against predation in these mesocosm ecosystems. Consequently, we call for the integration of non-random dispersal, dispersal syndromes and more generally intraspecific variation into studies predicting the impacts of invasions.

Male expected reproductive success can be enhanced by increased mating success (mate number) or, when females can mate multiply, by increased mating exclusivity (i.e., reduced partner promiscuity). A positive or negative covariance between these two mating outcomes could substantially increase or decrease overall variation in male expected reproductive success, yet the relationship between these mating outcomes is relatively understudied. We examined this relationship and the influence of male personality traits, female personality traits, and the social environment on mating outcomes in stream water striders, Aquarius remigis, at two experimental sex ratios: equal and 2:1 male-biased. To our knowledge, this study is the first to quantify this full set of effects. We found that mating frequency (mating success) and mating exclusivity were positively correlated in the male-biased treatment, but were not related at equal sex ratios. At both sex ratios, males that were more active and aggressive had both higher mating frequency and higher mating exclusivity. A male’s effective mating (the product of mating frequency and mating exclusivity) was also higher if on average he mated with females that tended to hide in refuges (and were presumably less available for future matings). Finally, males that were more often in pools with extremely aggressive (“hyperaggressive”) males actually had increased mating exclusivity, potentially because hyperaggression reduced average female promiscuity. This study highlights the importance of considering mating outcomes beyond mating success and examining the simultaneous contribution of male behaviours, female behaviours and social factors to mating dynamics.

Summary 1.Despite a central line of research aimed at quantifying relationships between mating success and sexually dimorphic traits (e.g., ornaments), individual variation in sexually selected traits often explains only a modest portion of the variation in mating success. 2.Another line of research suggests that a significant portion of the variation in mating success observed in animal populations could be explained by correlational selection, where the fitness advantage of a given trait depends on other components of an individual's phenotype and/or its environment. We tested the hypothesis that interactions between multiple traits within an individual (phenotype-dependence) or between an individual's phenotype and its social environment (context-dependence) can select for individual differences in behaviour (i.e., personality) and social plasticity. 3.To quantify the importance of phenotype- and context-dependent selection on mating success, we repeatedly measured the behaviour, social environment, and mating success of about 300 male stream water striders, Aquarius remigis. Rather than explaining individual differences in long-term mating success, we instead quantified how the combination of a male's phenotype interacted with the immediate social context to explain variation in hour-by-hour mating decisions. We suggest that this analysis captures more of the mechanisms leading to differences in mating success. 4.Males differed consistently in activity, aggressiveness, and social plasticity. The mating advantage of these behavioural traits depended on male morphology and varied with the number of rival males in the pool, suggesting mechanisms selecting for consistent differences in behaviour and social plasticity. Accounting for phenotype- and context-dependence improved the amount of variation in male mating success we explained statistically by 30 – 274%. 5.Our analysis of the determinants of male mating success provides important insights into the evolutionary forces that shape phenotypic variation. In particular, our results suggest that sexual selection is likely to favour individual differences in behaviour, social plasticity (i.e. individuals adjusting their behaviour), niche preference (i.e. individuals dispersing to particular social conditions), or social niche construction (i.e. individuals modifying the social environment). The true effect of sexual traits can only be understood in interaction with the individual's phenotype and environment. This article is protected by copyright. All rights reserved.

Ecology Letters (2012) Abstract Interspecific trait variation has long served as a conceptual foundation for our understanding of ecological patterns and dynamics. In particular, ecologists recognise the important role that animal behaviour plays in shaping ecological processes. An emerging area of interest in animal behaviour, the study of behavioural syndromes (animal personalities) considers how limited behavioural plasticity, as well as behavioural correlations affects an individual’s fitness in diverse ecological contexts. In this article we explore how insights from the concept and study of behavioural syndromes provide fresh understanding of major issues in population ecology. We identify several general mechanisms for how population ecology phenomena can be influenced by a species or population’s average behavioural type, by within-species variation in behavioural type, or by behavioural correlations across time or across ecological contexts. We note, in particular, the importance of behavioural type-dependent dispersal in spatial ecology. We then review recent literature and provide new syntheses for how these general mechanisms produce novel insights on five major issues in population ecology: (1) limits to species’ distribution and abundance; (2) species interactions; (3) population dynamics; (4) relative responses to human-induced rapid environmental change; and (5) ecological invasions.

Understanding patterns of non-random mating is central to predicting the consequences of sexual selection. Most studies quantifying assortative mating focus on testing for correlations among partners' phenotypes in mated pairs. Few studies have distinguished between assortative mating arising from preferences for similar partners (expressed by all or a subset of the population) vs. from phenotypic segregation in the environment. Also, few studies have assessed the robustness of assortative mating against temporal changes in social conditions. We tracked multiple matings by stream water striders (Aquarius remigis) across variable social conditions to investigate mating patterns by both body size and behavioural type (personality). We documented temporal changes in partner availability and used a mixed model approach to analyse individual behaviours and changes in mating status recorded on an hourly basis. We assessed whether all or only a subset of individuals in the population expressed a tendency to mate with similar phenotypes. Our analyses took into account variation in the level of competition and in the phenotypes of available partners. Males and females exhibited significant assortative mating by body size: the largest males and females, and the smallest males and females mated together more often than random. However, individuals of intermediate size were equally likely to mate with small, intermediate or large partners. Individuals also displayed two contrasting patterns of assortative mating by personality (activity level). Individuals generally mated preferentially with partners of similar activity level. However, beyond that general trend, individuals with more extreme personalities tended to exhibit disassortative mating: the most active males mated disproportionately with less active females and the least active males tended to mate with more active females. Our analyses thus revealed multiple, distinct patterns of nonrandom mating. These mating patterns did not arise from differences in partner availability among individuals and were robust to temporal changes in social conditions. Hence, mating patterns likely reflect mate preferences or arise from male-male competition coupled with sexual conflict. Our study also stresses the importance of accounting for variation in partner availability and demonstrates the influence of behavioural variation on mating patterns.

Dispersal is a fundamental life-history trait for many ecological processes. Recent studies suggest that dispersers, in comparison to residents, display various phenotypic specializations increasing their dispersal inclination or success. Among them, dispersers are believed to be consistently more bold, exploratory, asocial or aggressive than residents. These links between behavioural types and dispersal should vary with the cause of dispersal. However, with the exception of one study, personality-dependent dispersal has not been studied in contrasting environments. Here, we used mosquitofish ( Gambusia affinis ) to test whether personality-dependent dispersal varies with predation risk, a factor that should induce boldness or sociability-dependent dispersal. Corroborating previous studies, we found that dispersing mosquitofish are less social than non-dispersing fish when there was no predation risk. However, personality-dependent dispersal is negated under predation risk, dispersers having similar personality types to residents. Our results suggest that adaptive dispersal decisions could commonly depend on interactions between phenotypes and ecological contexts.

In social species, individual and group fitness and social dynamics in groups often depend on group size and on the group's social composition (e.g. the mix of personality types within groups). In turn, the size and social composition of groups is an emergent outcome of the grouping tendencies of individuals. While grouping behaviour has often been studied at a species level, within-species variation in grouping tendency has rarely been studied. We examined the role of personality type in shoaling preferences in a social fish, the western mosquitofish, Gambusia affinis. After scoring individuals for their social personality types (sociability), we gave individuals the choice to associate with shoals that differed in size and in the average sociability of individuals in the shoal. Shoal preferences depended on both the individual's sociability and on shoal size and composition in sociability types, and on the interaction between the individual's sociability and shoal size and composition. On average, fish preferred large to small shoals and groups made up of social individuals to groups made of asocial individuals at least when shoal size was small. Individual sociability types were linked to the time individuals spent far from any shoal and to consistent differences in choosiness about social contexts. Asocial individuals shoaled less and swam more between shoals than social ones, but preferred large shoals during the short time that they shoaled. These results can help explain patterns of group size and composition and individual and population-level personality-dependent dispersal.

Interspecific trait variation has long served as a conceptual foundation for our understanding of ecological patterns and dynamics. In particular, ecologists recognise the important role that animal behaviour plays in shaping ecological processes. An emerging area of interest in animal behaviour, the study of behavioural syndromes (animal personalities) considers how limited behavioural plasticity, as well as behavioural correlations affects an individual's fitness in diverse ecological contexts. In this article we explore how insights from the concept and study of behavioural syndromes provide fresh understanding of major issues in population ecology. We identify several general mechanisms for how population ecology phenomena can be influenced by a species or population's average behavioural type, by within-species variation in behavioural type, or by behavioural correlations across time or across ecological contexts. We note, in particular, the importance of behavioural type-dependent dispersal in spatial ecology. We then review recent literature and provide new syntheses for how these general mechanisms produce novel insights on five major issues in population ecology: (1) limits to species' distribution and abundance; (2) species interactions; (3) population dynamics; (4) relative responses to human-induced rapid environmental change; and (5) ecological invasions.

Ecological invasions are a major worldwide problem exacting tremendous economic and ecological costs. Efforts to explain variability in invasion speed and impact by searching for combinations of ecological conditions and species traits associated with invasions have met with mixed success. We use a simulation model that integrates insights from life-history theory, animal personalities, network theory, and spatial ecology to derive a new mechanism for explaining variation in animal invasion success. We show that spread occurs most rapidly when (1) a species includes a mix of life-history or personality types that differ in density-dependent performance and dispersal tendencies, (2) the differences between types are of intermediate magnitude, and (3) patch connections are intermediate in number and widely spread. Within-species polymorphism in phenotype (e.g., life-history strategies or personality), a feature not included in previous models, is important for overcoming the fact that different traits are associated with success in different stages of the invasion process. Polymorphism in sociability (a personality type) increases the speed of the invasion front, since asocial individuals colonize empty patches and facilitate the local growth of social types that, in turn, induce faster dispersal by asocials at the invasion edge. The results hold implications for the prediction of invasion impacts and the classification of traits associated with invasiveness.

Dispersal is one of the most fundamental components of ecology, and affects processes as diverse as population growth, metapopulation dynamics, gene flow and adaptation. Although the act of moving from one habitat to another entails major costs to the disperser, empirical and theoretical studies suggest that these costs can be reduced by having morphological, physiological or behavioural specializations for dispersal. A few recent studies on different systems showed that individuals exhibit personality-dependent dispersal, meaning that dispersal tendency is associated with boldness, sociability or aggressiveness. Indeed, in several species, dispersers not only develop behavioural differences at the onset of dispersal, but display these behavioural characteristics through their life cycle. While personality-dependent dispersal has been demonstrated in only a few species, we believe that it is a widespread phenomenon with important ecological consequences. Here, we review the evidence for behavioural differences between dispersers and residents, to what extent they constitute personalities. We also examine how a link between personality traits and dispersal behaviours can be produced and how personality-dependent dispersal affects the dynamics of metapopulations and biological invasions. Finally, we suggest future research directions for population biologists, behavioural ecologists and conservation biologists such as how the direction and the strength of the relationship between personality traits and dispersal vary with ecological contexts.

Ecological invasions, where non-native species spread to new areas, grow to high densities and have large, negative impacts on ecological communities, are a major worldwide problem. Recent studies suggest that one of the key mechanisms influencing invasion dynamics is personality-dependent dispersal: the tendency for dispersers to have a different personality type than the average from a source population. We examined this possibility in the invasive mosquitofish ( Gambusia affinis ). We measured individual tendencies to disperse in experimental streams and several personality traits: sociability, boldness, activity and exploration tendency before and three weeks after dispersal. We found that mosquitofish display consistent behavioural tendencies over time, and significant positive correlations between all personality traits. Most notably, sociability was an important indicator of dispersal distance, with more asocial individuals dispersing further, suggesting personality-biased dispersal on an invasion front. These results could have important ecological implications, as invasion by a biased subset of individuals is likely to have different ecological impacts than invasion by a random group of colonists.

Arising from: M. Wolf, G. S. van Doorn, O. Leimar & F. J. Weissing , 581–584 (2007)10.1038/nature05835 ; Wolf et al. reply Wolf et al.1 propose a model to explain the existence of animal personalities, consistent with behavioural differences among individuals in various contexts2,3,4—their explanation is counter-intuitive and cogent. However, all models have their limits, and the particular life-history requirements of this one may be unclear. Here we analyse their model and clarify its organismal scope.