Predation Regimes Research Articles

Environmental drivers of demographics, habitat use, and behavior during a post-Pleistocene radiation of Bahamas mosquitofish (Gambusia hubbsi)

A fundamental goal of evolutionary ecology is to understand the environmental drivers of ecological divergence during the early stages of adaptive diversification. Using the model system of the post-Pleistocene radiation of Bahamas mosquitofish (Gambusia hubbsi) inhabiting blue holes, we used a comparative field study to examine variation in density, age structure, tertiary (adult) sex ratio, habitat use, as well as adult feeding and social behaviors in relation to environmental features including predation risk, interspecific competition, productivity (e.g. chlorophyll a, zooplankton density), and abiotic factors (e.g. salinity, surface diameter). The primary environmental factor associated with ecological differentiation in G. hubbsi was the presence of piscivorous fish. Gambusia hubbsi populations coexisting with predatory fish were less dense, comprised of a smaller proportion of juveniles, and were more concentrated in shallow, near-shore regions of blue holes. In addition to predation risk, the presence of a competitor fish species was associated with G. hubbsi habitat use, and productivity covaried with both age structure and habitat use. Feeding and social behaviors differed considerably between sexes, and both sexes showed behavioral differences between predator regimes by exhibiting more foraging behaviors in the absence of predators and more sexual behaviors in their presence. Males additionally exhibited more aggressive behaviors toward females in the absence of predators, but were more aggressive toward other males in the presence of predators. These results largely matched a priori predictions, and several findings are similar to trends in other related systems. Variation in predation risk appears to represent the primary driver of ecological differentiation in this system, but other previously underappreciated factors (interspecific competition, resource availability) are notable contributors as well. This study highlights the utility of simultaneously evaluating multiple environmental factors and multiple population characteristics within a natural system to pinpoint environmental drivers of ecological differentiation.

Divergent reproductive life histories in Haida Gwaii stickleback (Gasterosteusspp.)

To assess intraspecific variability in fecundity of threespine stickleback (Gasterosteus aculeatus L., 1758) as a test of life-history trade-offs, we quantified egg traits, morphological characters, and habitat variables in 43 allopatric and morphologically diverse populations from Haida Gwaii off the west coast of Canada. Mean mature egg size and total egg count (12 eggs per female and 8 gravid females per locality) were both positively and significantly correlated with standard length (SL) of the fish (r = 0.58 and 0.84, respectively). Egg size was ∼10% larger in freshwater localities than in similar-sized stickleback in adjacent marine localities (P < 0.02). Multiple regression and Akaike’s information criterion (AIC) analyses of residual egg size against morphological and lake habitat variables yields a negative correlation with lake pH (partial r = –0.34, P < 0.05) and no association with lake size, aquatic spectra, gill raker number, defense armor, or predation regime. Relative to stickleback from continental regions, Haida Gwaii stickleback life histories appear to be K-shifted (large but few eggs) possibly because of cool temperatures, low aquatic productivity, low community diversity, predation regime, and occasional iteroparity.

High predation risk for a small mammal: the eastern pygmy-possum (Cercartetus nanus)

We report high rates of predation within a population of eastern pygmy-possum (Cercartetus nanus) near Sydney, observed during a study of the impacts of logging on the species. Predation by reptiles was observed on six of 61 radio-tracked pygmy-possums, while two were suspected of being taken by raptors. We further investigated predation by sooty owls (Tyto tenebricosa) by identifying remains in pellets. Pygmy-possums comprised 15% of the 126 dietary items identified. Predation occurred at both logged and unlogged sites in our study. We suggest that documented high rates of predation in our study could make the species vulnerable to altered predation regimes, such as influxes of feral predators, and highlight the need for a better understanding of any influence of logging on predator activity.

Predation’s Role in Life-History Evolution of a Livebearing Fish and a Test of the Trexler-DeAngelis Model of Maternal Provisioning

Populations experiencing consistent differences in predation risk and resource availability are expected to follow divergent evolutionary trajectories. For example, live-history theory makes specific predictions for how predation should drive life-history evolution, and according to the Trexler-DeAngelis model for the evolution of matrotrophy, postfertilization maternal provisioning is most likely to evolve in environments with consistent, high levels of resource availability. Using the model system of Bahamas mosquitofish (Gambusia hubbsi) inhabiting blue holes with and without the piscivorous bigmouth sleeper (Gobiomorus dormitor), we provide some of the strongest tests of these predictions to date, as resource availability does not covary with predation regime in this system, and we examine numerous (14) isolated natural populations. We found clear evidence for the expected life-history divergence between predation regimes and empirical support of the Trexler-DeAngelis model. Moreover, based on molecular and lab-rearing data, our study offers strong evidence for convergent evolution of similar life histories in similar predation regimes, largely matching previous phenotypic patterns observed in other poeciliid lineages (Brachyrhaphis spp., Poecilia reticulata), and further supports the notion that matrotrophy is most likely to evolve in stable high-resource environments.

Experimental Evidence for Density-Dependent Regulation and Selection on Trinidadian Guppy Life Histories

Recent study of feedbacks between ecological and evolutionary processes has renewed interest in population regulation and density-dependent selection because they represent black-box descriptions of these feedbacks. The roles of population regulation and density-dependent selection in life-history evolution have received a significant amount of theoretical attention, but there are few empirical examples demonstrating their importance. We address this challenge in natural populations of the Trinidadian guppy (Poecilia reticulata) that differ in their predation regimes. First, we tested whether natural populations of guppies are regulated by density dependence and quantified in which phases of the life cycle the effects of density are important. We found that guppies from low-predation (LP) environments are tightly regulated and that the density-dependent responses disproportionately affected some size classes. Second, we tested whether there are differences in density-dependent selection between guppies from LP or high-predation (HP) environments. We found that the fitness of HP guppies is more sensitive to the depressant effects of density than the fitness of LP guppies. Finally, we used an evolutionary invasion analysis to show that, depending on the effect of density on survival of the HP phenotype, this greater sensitivity of the HP phenotype to density can partially explain the evolution of the LP phenotype. We discuss the relevance of these findings to the study of feedbacks between ecology and evolution.

ResurrectedCeriodaphnia quadrangulahighlight differences between pheno- and genotypic expressions

The hatching of cladoceran ephippia from a 15-cm long sediment core was investigated, and Ceriodaphnia quadrangula clones were isolated from different sediment layers. Bosmina microfossil data were also analyzed, and compared with the corresponding data from a Pb210 dated core, which allowed us to infer the age of the sediment layers. Using changes in Bosmina microfossil morphologies, we were, furthermore, able to infer the presence of different regimes of fish predation. C. quadrangula was found to hatch in layers with an inferred age of approximately a century. Newly hatched individuals had smaller eye-size in sediment layers corresponding to high predation by young-of-the-year perch. Newly hatched individuals also generally had a marked neck-spine. In contrast, morphological characters of C. quadrangula clones reared in the laboratory over several generations showed no variation in relation to predation regime, indicating the absence of fixed genotype level changes. Furthermore, the laboratory grown clones only rarely produced a neck-spine. The results suggest phenotypic variation in response to the regime under which ephippia were produced.

Adding parasites to the guppy-predation story: insights from field surveys

Studies of phenotypic variation in nature often consider only a single potential selective agent. In such cases, it remains an open question as to whether variation attributed to that single measured agent might be influenced by some other unmeasured agent. Previous research has shown that phenotypic variation in the Trinidadian guppy (Poecilia reticulata) is strongly influenced by predation regime, and we here ask whether parasitism might represent an additional important selective agent shaping this variation. We performed a field survey of 26 natural guppy populations of known predation regime in northern Trinidad. We quantified levels of parasitism of guppies by the monogenean ecotoparasite, Gyrodactylus, and examined whether this parasite was associated with guppy body size or male colour. Spatial variation in Gyrodactylus parasitism was consistent between years, and parasite prevalence was generally, but not always, higher at high-predation sites than at low-predation sites. Consistent with previous work, predation regime was related to guppy size and some aspects of male colour, whereas parasitism showed few and only minor associations with the same traits. Moreover, a consideration of parasitism did not alter any interpretations regarding associations between guppy traits and predation regimes. These results suggest that parasitism, at least as quantified in the present study, does not play a major role in shaping variation in guppy body size or colour. Nevertheless, considerable variation in these traits, even within a predation regime, suggests the likely importance of other selective agents beyond just predation regime.

Ontogeny and sex alter the effect of predation on body shape in a livebearing fish: sexual dimorphism, parallelism, and costs of reproduction

Predation can cause morphological divergence among populations, while ontogeny and sex often determine much of morphological diversity among individuals. We used geometric morphometrics to characterize body shape in the livebearing fish Brachyrhaphis rhabdophora to test for interactions between these three major shape-determining factors. We assessed shape variation between juveniles and adults of both sexes, and among adults for populations from high- and low-predation areas. Shape differed significantly between predation regimes for all juveniles regardless of sex. As males grew and matured into adults, ontogenetic shape trajectories were parallel, thus maintaining shape differences in adult males between predation environments. However, shape of adult females between predation environments followed a different pattern. As females grew and matured, ontogenetic shape trajectories converged so that shape differences were less pronounced between mature females in predator and nonpredator environments. Convergence in female body shape may indicate a trade-off between optimal shape for predator evasion versus shape required for the livebearing mode of reproduction.

Philopatric predisposition to predation-induced ecological traps: habitat-dependent mortality of breeding eiders

Because population size is sensitive to changes in adult survival, adult survival may be buffered against environmental variability. Philopatry may be adaptive in changing environments, but it could also constrain breeding habitat selection under changing conditions such as shifting predation regimes. Habitat preference and quality could become decoupled in long-lived philopatric species that evolved in stable environments when suddenly faced by increased adult predation risk, as dispersal may be triggered by past reproductive failure. We evaluated whether the Baltic eider (Somateria m. mollissima) population may currently face a predation-induced ecological trap. Eiders are philopatric and nest on open and forested islands. We hypothesized that open-nesting females would be disproportionately affected by increased predation. We compared female annual survival in these two habitats in 1996-2010. We also tested for effects of time trends, winter severity (NAO), female body condition, and habitat-specific predation pressure on survival. Our results revealed the lowest survival recorded for this species (Φ=0.720), and survival on open islands was significantly lower (Φ=0.679) than on forested islands (Φ=0.761). Nonetheless, only 0.7% of females changed breeding habitat type despite ample availability of alternative islands, and breeding phenology in both habitats was similar. Female survival increased with body condition, while it was unrelated to winter climate and stable over time. Open islands had a higher predation pressure on incubating females. Breeding philopatry results in a predator-mediated ecological trap for open-nesting eiders. Our results contribute to explaining the drastic decline of the Baltic eider population.

Boldness in two perch populations – long‐term differences and the effect of predation pressure

1. Populations of the same species often display different behaviours, for example, in their response to predators. The question is whether this difference is developed as part of a divergent selection caused by differences in predation pressure, or as a result of phenotypic responses to current environmental conditions. 2. Two populations of Eurasian perch were investigated over a time span of 6 years to see whether risk-taking behaviour in young-of-the-year perch were consistent across cohorts, or if behaviour varied over time with changes in predation regime. 3. Boldness was estimated in aquarium studies by looking at how the fish made trade-offs between foraging in a risky area and staying in shelter. Predation risk of each year and lake was estimated from fishing surveys, using an individual-based model calculating attack rates for cannibalistic perch. 4. The average boldness scores were consistently lower in perch from Fisksjön compared with those in Ängersjön, although the magnitude of the difference varied among years. Variance component analyses showed that differences between lakes in boldness scores only explained 12 per cent of the total variation. Differences between years were contributing at least similarly or more to the total variance, and the variation was higher in Fisksjön than in Ängersjön. 5. The observed risk-taking behaviour of young-of-the-year perch, compared across cohorts, was significantly correlated with the year-specific estimates of cannibalistic attack rates, with lower boldness scores in years with higher predation pressure. In Fisksjön, with significant changes over the years in population structure, the range of both predation risk and boldness scores was wider than in Ängersjön. 6. By following the two perch populations over several years, we have been able to show that the differences in risk-taking behaviour mainly are due to direct phenotypic responses to recent experience of predation risk. Long-term differences in behaviour among perch populations thus reflect consistent differences in predation regime rather than diverging inherent traits.

Predation and habitat isolation influence the community composition-area relationship in dytiscid beetles (Coleoptera: Dytiscidae)

A major goal in ecology remains the understanding of patterns in diversity and distributions of species in natural communities. The species-area relationship is an important tool for investigating differences among communities, and may be also influenced by habitat isolation and dominant predator presence. In this two-year study, we evaluated the influences of habitat area, isolation and predation on community composition of dytiscids in two geographical regions dominated by different top predators (large predaceous dragonfly larvae or fish). Contrary to expectations, surface area, isolation and predator presence /absence alone did not significantly influence dytiscid species richness and total abundance, but in association with other environmental variables, such as submerged macrophyte growth forms. Components of habitat heterogeneity likely outweighed effects of area and predation regime on prey species diversity. However, differences in the set of abundant species were best explained by habitat surface area. Thus, in contrast to species diversity, gradients in community composition were not out-weighed by components of heterogeneity. In this study, predator presence was not correlated with habitat isolation. Instead, our results revealed that the effect of predator presence/absence on prey community composition and the resulting set of coexisting species may depend on habitat isolation. Within regions, the effect of large predaceous dragonfly larvae on species composition may depend on pond surface area, whereas the presence of fish influenced species composition alone and in association with area. We advocate that regional differences in environmental gradients be considered when community composition is evaluated.

Chaoborus and Gasterosteus Anti-Predator Responses in Daphnia pulex Are Mediated by Independent Cholinergic and Gabaergic Neuronal Signals

Many prey species evolved inducible defense strategies that protect effectively against predation threats. Especially the crustacean Daphnia emerged as a model system for studying the ecology and evolution of inducible defenses. Daphnia pulex e.g. shows different phenotypic adaptations against vertebrate and invertebrate predators. In response to the invertebrate phantom midge larvae Chaoborus (Diptera) D. pulex develops defensive morphological defenses (neckteeth). Cues originating from predatory fish result in life history changes in which resources are allocated from somatic growth to reproduction. While there are hints that responses against Chaoborus cues are transmitted involving cholinergic neuronal pathways, nothing is known about the neurophysiology underlying the transmission of fish related cues. We investigated the neurophysiological basis underlying the activation of inducible defenses in D. pulex using induction assays with the invertebrate predator Chaoborus and the three-spined stickleback Gasterosteus aculeatus. Predator-specific cues were combined with neuro-effective substances that stimulated or inhibited the cholinergic and gabaergic nervous system. We show that cholinergic-dependent pathways are involved in the perception and transmission of Chaoborus cues, while GABA was not involved. Thus, the cholinergic nervous system independently mediates the development of morphological defenses in response to Chaoborus cues. In contrast, only the inhibitory effect of GABA significantly influence fish-induced life history changes, while the application of cholinergic stimulants had no effect in combination with fish related cues. Our results show that cholinergic stimulation mediates signal transmission of Chaoborus cues leading to morphological defenses. Fish cues, which are responsible for predator-specific life history adaptations involve gabaergic control. Our study shows that both pathways are independent and thus potentially allow for adjustment of responses to variable predation regimes.

Little evidence for a selective advantage of armour-reduced threespined stickleback individuals in an invertebrate predation experiment

The repeated colonization of freshwater habitats by the ancestrally marine threespined stickleback Gasterosteus aculeatus has been associated with many instances of parallel reduction in armour traits, most notably number of lateral plates. The change in predation regime from marine systems, dominated by gape-limited predators such as piscivorous fishes, to freshwater habitats where grappling invertebrate predators such as insect larvae can dominate the predation regime, has been hypothesized as a driving force. Here we experimentally test the hypothesis that stickleback with reduced armour possess a selective advantage in the face of predation by invertebrates, using a natural population of stickleback that is highly polymorphic for armour traits and a common invertebrate predator from the same location. Our results provide no compelling evidence for selection in this particular predator–prey interaction. We suggest that the postulated selective advantage of low armour in the face of invertebrate predation may not be universal.

Widespread intraspecific organismal stoichiometry among populations of the Trinidadian guppy

Summary 1. Ecological stoichiometry expresses ecological interactions as the balance between multiple elements. It relates the ecological function of organisms to their elemental composition, or their organismal stoichiometry. Organismal stoichiometry is thought to reflect elemental investments in life history and morphology acting in concert with variability in abiotic or environmental conditions, but the relative contribution of these factors to natural variability in organismal stoichiometry is poorly understood. 2. We assessed the relative contribution of stream identity, predation, body size and sex to the organismal stoichiometry of guppies (Poecilia reticulata) in six streams in Trinidad. In this system, guppy life-history phenotype evolves in response to predation. Guppies adapted to high-predation (HP) pressure grow faster, mature earlier, produce fewer and smaller offspring and eat a higher-quality diet than guppies adapted to low-predation (LP) pressure. This pattern of life-history evolution is repeated in many rivers encompassing a wide range of abiotic conditions. 3. Organismal stoichiometry of guppies was widely variable, spanning up to ∼70% of the range of variability reported across freshwater fish taxa. Streams from where guppies were sampled were the most important predictor of organismal stoichiometry. In many cases, guppy populations from sites within the same stream varied as much as from sites in different streams. 4. Surprisingly, predation regime was a minor predictor of % C, C : P and C : N in female guppies, despite its strong correlation with life-history phenotype and other organismal traits in this species. Body size and sex were not significant predictors of organismal stoichiometry. 5. Guppies from HP sites were more stoichiometrically balanced with their diets than guppies from LP sites. The latter appeared to be more vulnerable to phosphorus limitation than the former, suggesting that dietary specialization associated with guppy life-history phenotype may have stoichiometric consequences that can affect guppy physiology and nutrient recycling. 6. Our findings suggest that local environmental conditions are a stronger predictor of organismal stoichiometry than organismal traits. We recommend that future work should explicitly consider correlations between organismal traits and organismal stoichiometry in the context of environmental heterogeneity.

Are host–parasite interactions influenced by adaptation to predators? A test with guppies and Gyrodactylus in experimental stream channels

Natural populations often face multiple mortality sources. Adaptive responses to one mortality source might also be beneficial with respect to other sources of mortality, resulting in "reinforcing adaptations"; or they might be detrimental with respect to other sources of mortality, resulting in "conflicting adaptations". We explored these possibilities by testing experimentally if the responses of guppies (Poecilia reticulata) to the monogenean ectoparasitic worm Gyrodactylus differed between populations adapted to different predation regimes. In experimental stream channels designed to replicate the natural environment, we exposed eight guppy populations (high-predation and low-predation populations from each of four separate rivers) either to their local Gyrodactylus parasites (infection treatment) or to the absence of those parasites (control). We found that infection dynamics varied dramatically among populations in a repeatable fashion, but that this variation was not related to the predation regime of origin. Consistent with previous work, high-predation guppy females gained more mass, had lower reproductive investment, and had more but smaller embryos than did low-predation females. Relative to control (no parasite) channels, guppies from treatment (infected) channels gained less mass but produced similar numbers and sizes of embryos-and thus had a higher reproductive effort. However, no interaction was evident between infection treatment and predation regime. We conclude that parasitism by Gyrodactylus and predation are both likely selective forces for guppies, but that adaptation to predation does not have an obvious deterministic effect on host-parasite dynamics or on life-history traits of female guppies.

A test of the coupling of predator defense morphology and behavior variation in two threespine stickleback populations

Abstract Among-population differences in morphology and behaviors such as boldness have been shown to co-vary with ecological conditions, including predation regime. However, between- and within-population covariation of predator defense morphology with variation in behaviors relevant to ecology and evolution (boldness, exploration, activity, sociability and aggressiveness, often defined as personality traits when they are consistent across time and contexts) have never been quantified together in a single study in juvenile fish from populations found in contrasting environments. We measured predator defense morphology differences between adults from two freshwater populations of threespine sticklebacks with different ecological conditions. We then quantified five behaviors in juveniles from both populations raised in a common environment. Wild-caught adults showed significant differences in predator defense morphology. One population had significantly lower lateral plate number, shorter dorsal spine, pelvic spine and pelvic girdle. Furthermore, 61% of individuals from that population showed an absence of pelvic spine and girdle. At the population level, we found that differences in defense morphology in adults between the two lakes were coupled with differences in behaviors in juveniles raised in a common environment. Levels of activity, aggressiveness and boldness were higher in juveniles from the population lacking predator defense structures. At the individual level, anti-predator morphology of adult females could not predict their offspring’s behavior, but juvenile coloration predicted individual boldness in a population-specific manner. Our results suggest that ecological conditions, as reflected in adult predator defense morphology, also affect juvenile behavior in threespine sticklebacks, resulting in trait co-specialization, and that there is a genetic or epigenetic component to these behavioral differences.

PREDATION-ASSOCIATED DIFFERENCES IN SEX LINKAGE OF WILD GUPPY COLORATION

Evolutionary theory predicts that the sex linkage of sexually selected traits can influence the direction and rate of evolutionary change, and also itself be subject to selection. Theory abounds on how sex-specific selection, mate choice, or other phenomena should favor different types of sex-linked inheritance, yet evidence in nature remains limited. Here, we use hormone assays in Trinidadian guppies to explore the extent to which linkage of male coloration differs among populations adapted to varying predation regimes. Results show there is consistently higher degree of X- and autosomal linkage in body coloration among populations adapted to low-predation environments. More strikingly, analyses of an introduced population of guppies from a high- to a low-predation environment suggest that this difference can change in 50 years or less.

Does sexual selection evolve following introduction to new environments?

The rate of evolution of mating preferences and mate signalling traits can influence local adaptation and diversification under environmental change. However, the rate of evolution of female preferences has not been directly examined in natural populations. An opportunity to do so arose through the introduction of high-predation Trinidadian guppies, Poecilia reticulata, from the Yarra River into high- and low-predation environments in the Damier River. Nine years (13–26 guppy generations) after the introduction, we tested whether female preferences for key aspects of male colour differed between the rivers and between introduced high- and low-predation populations. Based on two independent laboratory experiments, we conclude that little divergence in female preferences has apparently occurred between any of the populations, or between predation regimes. In combination with previous work, these results suggest that the evolution of guppy colour and female preferences are influenced by factors in addition to just predation, and that female preferences may generally take longer to evolve than other types of traits, particularly in populations that experience weak or fluctuating sexual selection gradients.

Predator-Induced Morphological Plasticity Across Local Populations of a Freshwater Snail

The expression of anti-predator adaptations may vary on a spatial scale, favouring traits that are advantageous in a given predation regime. Besides, evolution of different developmental strategies depends to a large extent on the grain of the environment and may result in locally canalized adaptations or, alternatively, the evolution of phenotypic plasticity as different predation regimes may vary across habitats. We investigated the potential for predator-driven variability in shell morphology in a freshwater snail, Radix balthica, and whether found differences were a specialized ecotype adaptation or a result of phenotypic plasticity. Shell shape was quantified in snails from geographically separated pond populations with and without molluscivorous fish. Subsequently, in a common garden experiment we investigated reaction norms of snails from populations' with/without fish when exposed to chemical cues from tench (Tinca tinca), a molluscivorous fish. We found that snails from fish-free ponds had a narrow shell with a well developed spire, whereas snails that coexisted with fish had more rotund shells with a low spire, a shell morphology known to increase survival rate from shell-crushing predators. The common garden experiment mirrored the results from the field survey and showed that snails had similar reaction norms in response to chemical predator cues, i.e. the expression of shell shape was independent of population origin. Finally, we found significant differences for the trait means among populations, within each pond category (fish/fish free), suggesting a genetic component in the determination of shell morphology that has evolved independently across ponds.

The influence of experience on risk taking: results from a common-garden experiment on populations of Eurasian perch

Predation is often thought of as an unforgiving and strong selective force, quickly selecting against maladaptive behaviour in the prey. It is argued that experience is likely to have low influence on the phenotypic response to predation, as failing to react correctly to a predator may mean death to the prey and no second chance to learn and correct the behaviour. Individuals from different populations of Eurasian perch are known to differ in risk-taking behaviour. Variation in predation pressure has been suggested as a key factor causing these differences, but little is known about the underlying mechanism by which predation generates risk-taking phenotypes in perch. We compared the degree of boldness between two natural populations of Eurasian perch, living under different predation regimes, and the same populations hatched and reared under identical conditions, free from predation. By this common-garden approach, we sought to investigate patterns in the influence of inheritance and experience on boldness phenotype. The wild fish differed in risk taking, with fish from the low predation-risk population acting bolder than fish from the high-risk environment. In the reared fish, both populations behaved equally bold. Only the fish originating from the high predation population showed different behaviour when comparing wild and reared ecotypes. Our results suggest that experience has an important impact on the response to predators and that geographic variation in risk taking between populations of Eurasian perch to a high degree is shaped by adjustments to the current environment. Habituation had an effect of risk-taking behaviour over the experimental period, but consistent differences between individuals were also found. Furthermore, we also show, by the estimation of variance components, that the behaviour we observe is affected by a range of random effects, such as aquaria and group membership, that in concert shapes the behaviour of an individual perch.