Environmental Predation Research Articles

Morphological divergence driven by predation environment within and between species of Brachyrhaphis fishes.

Natural selection often results in profound differences in body shape among populations from divergent selective environments. Predation is a well-studied driver of divergence, with predators having a strong effect on the evolution of prey body shape, especially for traits related to escape behavior. Comparative studies, both at the population level and between species, show that the presence or absence of predators can alter prey morphology. Although this pattern is well documented in various species or population pairs, few studies have tested for similar patterns of body shape evolution at multiple stages of divergence within a taxonomic group. Here, we examine morphological divergence associated with predation environment in the livebearing fish genus Brachyrhaphis. We compare differences in body shape between populations of B. rhabdophora from different predation environments to differences in body shape between B. roseni and B. terrabensis (sister species) from predator and predator free habitats, respectively. We found that in each lineage, shape differed between predation environments, consistent with the hypothesis that locomotor function is optimized for either steady swimming (predator free) or escape behavior (predator). Although differences in body shape were greatest between B. roseni and B. terrabensis, we found that much of the total morphological diversification between these species had already been achieved within B. rhabdophora (29% in females and 47% in males). Interestingly, at both levels of divergence we found that early in ontogenetic development, females differed in shape between predation environments; however, as females matured, their body shapes converged on a similar phenotype, likely due to the constraints of pregnancy. Finally, we found that body shape varies with body size in a similar way, regardless of predation environment, in each lineage. Our findings are important because they provide evidence that the same source of selection can drive similar phenotypic divergence independently at multiple divergence levels.

Ontogenetic shifts in phenotype-environment associations in Nile perch,Lates niloticus(Perciformes: Latidae) from Lake Nabugabo, Uganda

Habitat-associated trait divergence may vary across ontogeny if there are strong size-related shifts in selection pressures. We quantified patterns of phenotypic divergence in Nile perch (Lates niloticus) from ecologically distinct wetland edge and forest edge habitats in Lake Nabugabo, Uganda, and we compared patterns of divergence across three size classes to determine whether trends are consistent through Nile perch ontogeny. We predicted that inter-habitat variation in biotic (e.g. vegetation structure) and abiotic (e.g. dissolved oxygen concentration) variables may create divergent selective regimes. We compared body morphology using geometric morphometrics and found substantial differences between habitats, although not all trends were consistent across size classes. The most striking aspects of divergence in small Nile perch were in mouth orientation, head size, and development of the caudal region. Medium-sized Nile perch also showed differences in mouth orientation. Differences in large individuals were related to eye size and orientation, as well as caudal length. The observed patterns of divergence are consistent with functional morphological predictions for fish across divergent trophic regimes, high and low predation environments, and complex and simple habitats. Although this suggests adaptive divergence, the source of phenotypic variation is unknown and may reflect phenotypic plasticity and/or genetic differences. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110, 449–465.

The influence of body size on the foraging behaviour of European hake after settlement to the bottom

The trophic niche width and feeding strategy (FS) of juvenile European hake were studied during three trawl surveys in two different areas of the Mediterranean Sea. Both of these aspects of trophic behaviour were analysed as function of different biotic and abiotic factors. Prey were grouped by factors such as size, swimming behaviour and energy density. The trophic niche breadth values observed (2.25–2.89) were very similar regardless of the geographic area (geomorphology) and season. The FSs analysed were also quite similar regardless of the geographic area (geomorphology) and season, or predation environment (bottom or water column). Conversely, the FSs were linked with predator size. Increased predator size allowed changes in predatory behaviour through the development of preferences for larger and more energy-rich prey. The changes in FS from generalist to specialist began at a total length (TL) of approximately 12cm in both populations analysed. At larger sizes, the juveniles of both populations became more specialised, as confirmed by the reduction in the breadth of the individual trophic niche. The observed simultaneous changes in the FS, which occurred regardless of the geomorphology and seasonal characteristics, suggest that this shift is driven primarily by endogenous factors. The influence of body size on the foraging behaviour of European hake in the early life stages after bottom recruitment is a crucial information for the application of ecosystem mass-balance models.

Geographic Variation of Movement and Display Behavior of Side-blotched Lizards (Uta stansburiana) Related to Predation Environment

Abstract Animals in high predation environments tend to react to predators more quickly and effectively compared to animals in low predation environments. Because antipredator behavior can be increased by predator introductions and decreased by predator losses, we examined whether populations occurring outside of a predator's range are associated with behavioral changes. Genetic evidence indicates that Side-blotched Lizards (Uta stansburiana) radiated into the northern portions of their range relatively recently and rapidly. As such, Uta populations exist in the presence or absence of various predators. We studied four populations of Uta in Oregon and Nevada to test for behavioral differences among populations that co-occur with different predators. We measured several behavioral traits thought to be associated with predator exposure and used predation-pressure estimates for each population to show that movement and display behavior vary among sites but tend to be negatively affected by predation environm...

Effects of predation environment and food availability on somatic growth in theLivebearingFishBrachyrhaphis rhabdophora(Pisces:Poeciliidae)

Variation in somatic growth rates is of great interest to biologists because of the relationship between growth and other fitness-determining traits, and it results from both genetic and environmentally induced variation (i.e. plasticity). Theoretical predictions suggest that mean somatic growth rates and the shape of the reaction norm for growth can be influenced by variation in predator-induced mortality rates. Few studies have focused on variation in reaction norms for growth in response to resource availability between high-predation and low-predation environments. We used juvenile Brachyrhaphis rhabdophora from high-predation and low-predation environments to test for variation in mean growth rates and for variation in reaction norms for growth at two levels of food availability in a common-environment experiment. To test for variation in growth rates in the field, we compared somatic growth rates in juveniles in high-predation and low-predation environments. In the common-environment experiment, mean growth rates did not differ between fish from differing predation environments, but the interaction between predation environment and food level took the form of a crossing reaction norm for both growth in length and mass. Fish from low-predation environments exhibited no significant difference in growth rate between high and low food treatments. In contrast, fish from high-predation environments exhibited variation in growth rates between high and low food treatments, with higher food availability resulting in higher growth rates. In the field, individuals in the high-predation environment grow at a faster rate than those in low-predation environments at the smallest sizes (comparable to sizes in the common-environment experiment). These data provide no evidence for evolved differences in mean growth rates between predation environments. However, fish from high-predation environments exhibited greater plasticity in growth rates in response to resource availability suggesting that predation environments may exhibit increased variation in food availability for prey fish and consequent selection for plasticity.

Soldiers with large weapons in predator-abundant midsummer: phenotypic plasticity in a eusocial aphid

Eusocial aphids produce sterile individuals (“soldiers”) that specialize behaviorally and morphologically to protect their colony from predators, while production of soldiers can negatively affect colony growth because of reproductive allocation and opportunity cost. Hence, a cost-saving soldier production strategy is expected to be favored. Here, we hypothesize that, to save the cost, a eusocial aphid Ceratovacuna japonica produces soldiers with smaller weapon in the season when predators are not abundant. The abundance of two specialist lepidopteran predators (i.e., Taraka hamada and Atkinsonia ignipicta) of C. japonica dramatically increased, and aphid colony size significantly decreased, from July to August. In line with these, the soldiers in August had larger weapons (i.e., frontal horns) than those in June, indicating a correlational increase in weapon size with predation pressure. We predict that a reliable prospective signal indicating the coming of midsummer (environmental temperature) induces mother aphids to produce soldiers with larger weapons. Experiments clarified that soldiers produced at 20 °C (typical temperature of July to August) had larger weapons than those produced at 15 °C (typical temperature of May to July). Such phenotypic plasticity appears to be adaptive to maximize the fitness of C. japonica under a temporally variable but predictable predation environment. These results indicate that C. japonica aphids not merely have distinctive reproductive—and soldier castes, but also produce differentially armed soldiers in a habitat with temporally changing predation risks.

The Effect of Exploration on the Use of Producer-Scrounger Tactics

Individuals foraging in groups can use two different tactics for obtaining food resources. Individuals can either search for food sources themselves (producing) or they can join food discoveries of others (scrounging). In this study we use a genetic algorithm in a spatially explicit producer-scrounger game to explore how individuals compromise between exploration (an important axis of animal personality) and scrounging and how characteristics of the environment affect this compromise. Agents varied in exploration and scrounging and a genetic algorithm searched for the optimal combination of exploration and scrounging. The foraging environments featured different levels of patch richness, predation and patch density. Our simulations show that under conditions of low patch densities slow exploring scroungers were favored whereas high patch density favored fast exploring individuals that either produced (at low patch richness) or scrounged (at high patch richness). In high predation environments fast exploring individuals were selected for but only at low to intermediate patch densities. Predation did not affect scrounging behavior. We did not find a divergence of exploration ‘types’ within a given environment, but there was a general association between exploration and scrounging across different environments: high rates of scrounging were observed over nearly the full spectrum of exploration values, whereas high rates of producing were only observed at high exploration values, suggesting that cases in which slow explorers start producing should be rare. Our results indicate that the spatial arrangement of food resources can affect the optimal social attraction rules between agents, the optimality of foraging tactic and the interaction between both.

Assessing behavioural and morphological responses of frog tadpoles to temporal variability in predation risk

AbstractFinely tuned adjustment of an individual's phenotype can offer substantial fitness benefits when it is closely matched with environmental change. For instance, prey may be safeguarded against unnecessary costs to growth or development when their responses to temporally variable predation risk include plastic anti‐predator defences. Yet, the correspondence between perceived predation risk and related responses should differ between behavioural and morphological phenotypes when risk fluctuates because behaviour can be modified quickly, whereas morphological phenotypes require time to build. Theoretical models predict intermediate expression when risk fluctuates rapidly relative to the time required to mount a response, whereas traits that can be modified relatively quickly should more closely track current conditions. Using a tadpole‐dragonfly larva system, we sought to compare the expression of behavioural and morphological defences following exposure to constant versus variable predation risk. By varying the pattern and total duration of predator cue exposure, but not cue concentration, we quantified phenotypic plasticity and trait reversibility. Our results show that strong behavioural responses were limited to early ontogeny but closely matched current level of risk. The morphology of prey experiencing a weekly changing predator environment was intermediate to that of prey in the no‐predator and constantly exposed treatments. Yet, prey exposed to a predator environment for the same total duration as the weekly changing environment, but in a different exposure pattern, was morphologically unresponsive to the onset of predation risk. Finally, unexposed tadpoles gained deeper tails and smaller relative body size in late development, coincident with limb bud development. Such changes are consistent with anti‐predator response and represent either an innate response when prey are more vulnerable or shape optimization when faced with increased drag. We conclude that phenotypic expression depends critically on patterns of temporal variability in the environment, although the actual extent of expression depends on the specific trait in question.

Correlating context-specific boldness and physiological condition of female sand fiddler crabs (Uca pugilator)

Consistent individual behavioral differences across ecological contexts are a recognized feature of animal populations. These differences can be expressed in two ways: context-specifically or context-generally. The former is characterized by consistent responses in one context (i.e. repeatability), whereas the latter by consistency that spans contexts (i.e. behavioral syndromes). The proximate causes of behavioral consistency remain unclear, yet there is evidence that physiology may couple the expression of some behavioral traits in unrelated contexts. We therefore explored the correlation between bold behavior of female sand fiddler crabs (Uca pugilator) and the condition of the hepatopancreas, an organ vital to crustacean metabolism and reproduction. We did this by taking replicate measurements of two risk-taking behaviors per individual in the contexts of predator avoidance and environment exploration, and examining correlations within and between these observations. We then determined the relationship of behavior with hepatopancreas mass and lipid content. Individual crabs responded consistently within each context. However, across-context correlations were absent, indicating that boldness is isolated, at least in the selected scenarios. Additionally, anti-predator and exploratory behaviors were significantly influenced by size but not linked to hepatopancreas physiology. Our results show that context-specific trait expression may occur in the absence of a physiological correlate.

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.

An approach for quantifying effects of multiple predators that forage on different time scales

We used a comprehensive experimental design, consisting of a full suite of predator environments, behavioural observations, and calculation of per capita consumption rates, to examine multiple predator effects on prey when green crabs (Carcinus maenas) and dogwhelks (Nucella lapillus) foraged on blue mussels (Mytilus edulis). Predator environments (i.e., isolated predators, conspecific pairs, and interspecific pair) were used to identify multiple predator effects on prey by comparing observed predation to that predicted from the multiplicative risk model. Behavioural data and per capita consumption rates provided insight into mechanisms underlying observed multiple predator effects on prey. Experiments were conducted at various spatial scales in the laboratory and field. In all experiments, dogwhelks reduced or ceased feeding when a green crab was present compared to isolated dogwhelks, although overall predation was not influenced. Independent multiple predator effects on prey, where observed and predicted consumption did not differ, were usually maintained because crabs and whelks forage on dissimilar time scales. Contrary to other studies, interference competition through aggressive encounters was not observed for dogwhelk and green crab conspecifics, or the interspecific pair, and non-independent multiple predator effects on prey in the form of risk reduction did not result. Spatial scale appeared to influence some outcomes of predator interactions. In large tanks, green crab conspecifics and the interspecific pair caused risk enhancement for prey. This resulted from increased search time, increased per capita consumption rate, and decreased handling time per prey of crabs, and was likely induced by longer residence time of water in the large tanks compared to smaller tanks. In small tanks, dogwhelks with a conspecific had higher per capita consumption rates than isolated dogwhelks, suggesting risk enhancement for prey. However, dogwhelks were unable to consume enough prey over the experiment to allow detection of a non-independent multiple predator effect on prey by comparing observed to predicted consumption. Predator species that operate on different time scales of predation may have independent multiple predator effects on prey even when changes to individual predator behaviours or per capita consumption rates occur. Examination of predator behaviours and per capita consumption rates, in addition to the full suite of predator environments, is essential to fully understand dynamics among multiple conspecific and interspecific predators, particularly when predator species forage on dissimilar time scales.

Structural consistency of behavioural syndromes: does predator training lead to multi-contextual behavioural change?

Behavioural syndromes are suites of behaviours that are correlated across multiple contexts. Syndromes may occur in populations because behaviours are tightly linked by underlying mechanisms, such as genetics or physiology, which constrain flexibility and preclude multi-contextual plasticity. Alternatively, correlated behaviours may not share a common mechanism and may be able to change independently, allowing for potentially maladaptive combinations of traits to be broken apart. We tested these two hypotheses by training Trinidadian guppies (Poecilia reticulata) which possessed a behavioural syndrome encompassing three contexts, to avoid a potential predator. While we found no difference in magnitude of behavioural change between the trained and control groups, we did find that all subjects generally became shyer toward a potential predator following training and hypothesise that this resulted from sensitization and a predisposition to quickly recognize and adjust behaviour to predator-like stimuli. Importantly, behavioural changes in response to a potential predator did not generate changes in ‘general activity’ or ‘exploration’, and a tri-contextual syndrome broke apart. Our results suggest that in this population of guppies, individuals differ in behavioural plasticity in terms of their response to experience with predation risk and behaviours across contexts are domain specific and are able to change independently of each other. Future research should focus on populations that evolved in high predation environments and, therefore, may possess more rigid syndromes, to determine whether behavioural flexibility is limited.

Morphological convergence during pregnancy among predator and nonpredator populations of the livebearing fishBrachyrhaphis rhabdophora(Teleostei: Poeciliidae)

Predation can drive morphological divergence in prey populations, although examples of divergent selection are typically limited to nonreproductive individuals. In livebearing females, shape often changes drastically during pregnancy, reducing speed and mobility and enhancing susceptibility to predation. In the present study, we document morphological divergence among populations of nonreproductive female livebearing fish (Brachyrhaphis rhabdophora) in predator and nonpredator environments. We then test the hypothesis that shape differences among nonreproductive females are maintained among reproductive females between predator and nonpredator environments. Nonreproductive females in predator environments had larger caudal regions and more fusiform bodies than females in nonpredator environments; traits that are associated with burst speed in fish. Shape differences were maintained in reproductive females, although the magnitude of this difference declined relative to nonreproductive females, suggesting morphological convergence during pregnancy. Phenotypic change vector analysis revealed that females in predator environments became more similar to females in nonpredator environments in the transition from nonreproductive to reproductive. Furthermore, the level of reproductive allocation affected shape similarly between predator environments. These results suggest a life-history constraint on morphology, in which predator-driven morphological divergence among nonreproductive B. rhabdophora is not maintained at the same level during pregnancy. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104, 386–392. ADDITIONAL KEYWORDS: geometric morphometrics – life history – morphology – predation – trade-off.

Green Iguanas (Iguana iguana) in Puerto Rico: is it time for management?

Iguana iguana is native to Central and South America, and was introduced into Puerto Rico in the 1970s as a result of pet trade. The invasive biology of this reptile has not been studied in Puerto Rico, where its negative effects may threaten local biodiversity. The purposes of this study were to: (1) estimate population densities of I. iguana; (2) describe some aspects of its reproductive biology; and (3) assess its potential impacts. Visual-encounter surveys were performed at Parque Lineal in San Juan and Canal Blasina in Carolina, while nesting activity data were collected at Las Cabezas de San Juan in Fajardo. Densities of I. iguana in Puerto Rico reached a maximum of 223 individuals ha−1, higher than in any known locality in its native range, and showed fluctuations related to seasonality. Our 2008–2009 observations at the nesting sites document that this population of I. iguana is a reproductively successful species, producing more than 100 egg clutches and 2,558 eggs with a 91.4% egg viability. The ability to proliferate in a low predation environment and the absence of good competitors are the major drivers of the population densities observed in Puerto Rico. We found evidence that I. iguana is threatening native biodiversity and impacting infrastructure, agriculture and human safety. Thus, a management program to control the species must soon be developed to prevent this invasive reptile from becoming more widespread and dominant in other localities around the island.

Colonization of novel White Sands habitat is associated with changes in lizard anti-predator behaviour

Colonization of novel habitats is often associated with differences in ecological community composition. For small diurnal animals, differences in predator diversity and abundance can lead to behavioural shifts in the novel habitat. The eastern fence lizard Sceloporus undulatus (Bosc and Daudin, 1801) recently colonized the gypsum dunes of White Sands, a predator-poor community relative to the predator-rich community of the surrounding Chihuahuan dark-soil habitat. We used field experiments to assess S. undulatus anti-predator behaviour in white-sand versus dark-soil habitats, and used laboratory assays to determine whether behavioural differences could be mediated by hormonal regulation. Overall, we found that white-sand lizards were less vigilant but more wary than their dark-soil counterparts; it took them longer to detect a simulated predator, but once detected they were more likely to retreat from their perches than dark-soil lizards. At the proximate level, differences in anti-predator behaviour could not be explained by differences in plasma hormone levels (corticosterone and testosterone); we detected elevated corticosterone for lizards in our stress treatment relative to control treatment, but found no differences between habitats in baseline or acute corticosterone levels. At the evolutionary level, we suggest that differences in anti-predator behaviour may be explained by differences across habitats in predation environment, habituation, and/or the cost of retreating. Our study implicates changes in predator community composition in mediating ecological divergence in behaviour.

Guppy populations differ in cannibalistic degree and adaptation to structural environments

There is considerable variation in cannibalism between different species and also between individuals of different species, although relatively little is known about what creates this variation. We investigated the degree of cannibalism in guppy (Poecilia reticulata) populations originating from high and low predation environments in Trinidad, and also how cannibalism was affected by the presence of refuges. Females from two populations were allowed to feed on juveniles from two populations in aquaria trials. The cannibalism was size-dependent and varied depending on both juvenile and female origin. Low predation females were more efficient cannibals and low predation juveniles were better at avoiding cannibalism compared to high predation guppies when no refuges were present. The high predation females were superior cannibals and the high predation juveniles were better at escaping cannibalism than the low predation guppies when refuges were present. We discuss whether the differences in cannibalism and response to refuge addition relate to predation-induced habitat shifts and differences in the guppies' natural environment.

A small increase in UV-B increases the susceptibility of tadpoles to predation

Increased ultraviolet-B (UV-B) radiation as a consequence of ozone depletion is one of the many potential drivers of ongoing global amphibian declines. Both alone and in combination with other environmental stressors, UV-B is known to have detrimental effects on the early life stages of amphibians, but our understanding of the fitness consequences of these effects remains superficial. We examined the independent and interactive effects of UV-B and predatory chemical cues (PCC) on a suite of traits of Limnodynastes peronii embryos and tadpoles, and assessed tadpole survival time in a predator environment to evaluate the potential fitness consequences. Exposure to a 3 to 6 per cent increase in UV-B, which is comparable to changes in terrestrial UV-B associated with ozone depletion, had no effect on any of the traits measured, except survival time in a predator environment, which was reduced by 22 to 28 per cent. Exposure to PCC caused tadpoles to hatch earlier, have reduced hatching success, have improved locomotor performance and survive for longer in a predator environment, but had no effect on tadpole survival, behaviour or morphology. Simultaneous exposure to UV-B and PCC resulted in no interactive effects. These findings demonstrate that increased UV-B has the potential to reduce tadpole fitness, while exposure to PCCs improves their fitness.

Effect of predation and habitat quality on growth and reproduction of a stream fish

Billman EJ, Tjarks BJ, Belk MC. Effect of predation and habitat quality on growth and reproduction of a stream fish. Ecology of Freshwater Fish 2011: 20: 102–113. © 2010 John Wiley & Sons A/SAbstract – Anthropogenic disturbances are rarely independent, requiring native fishes to respond to multiple factors to persist in changing environments. We examined the interaction of predation environment (presence of introduced brown trout, Salmo trutta) and habitat quality on growth and reproduction of southern leatherside chub, Lepidomeda aliciae, a small‐bodied stream fish native to central Utah, USA. Southern leatherside chub were sampled from four streams representing a complete two‐factor cross of predation environment and habitat quality. Growth was estimated using increment analysis of annuli on otoliths, and reproductive traits were measured for both sexes. Southern leatherside chub growth was greater in high‐quality than in low‐quality habitats, and greater in predator than in nonpredator environments. However, fish exhibited a greater growth response to presence of brown trout in low‐quality habitats. Southern leatherside chub growth followed predictions of plastic responses to resource availability based on habitat quality and predation environment (lethal vs. nonlethal effects). Reproductive allocation (gonad wet mass) was significantly greater in low‐quality versus high‐quality habitats, but was unaffected by predation environment. Other female life‐history traits were affected either by both effects or their interaction. Reproductive responses to habitat quality and predation environment were consistent with predictions based on differential mortality. Southern leatherside chub growth and reproduction responded differently to the combination of habitat quality and predation environment, thus demonstrating the importance of assessing interacting effects of anthropogenic disturbances to more fully comprehend impacts on native species and to appropriately manage, recover and restore these species and their habitats.

Effects of species‐specific interactions with predation risk on the relative species sensitivities to a pesticide in water boatmen (Corixidae)

There is increasing concern that synergistic interactions between stressors may result in accelerated biodiversity loss. Yet, the prevalence and magnitude of these interactions remain one of the largest uncertainties in projections of future ecological change. Synergistic interactions between pesticide stress and predation risk are receiving increasing attention because they indicate that standard pesticide tests in the absence of predation risk may underestimate effects that occur under natural conditions. We questioned whether synergisms (or interactions in general), by differently shaping individual species’ sensitivities, can modulate species sensitivity rankings (SSRs) in survival and growth rate. Using laboratory and outdoor mesocosm experiments with five species of water boatmen (Hemiptera: Corixidae) we studied mortality and growth rate over seven days of exposure. We evaluated the presence and nature of interactions between the pesticide endosulfan and predation risk and tested to what extent this affected the SSRs to endosulfan. The combined exposure to the pesticide and predation risk resulted in synergistic effects for survival (Sigara lateralis) and growth rate (Sigara iactans) and in an antagonistic effect for growth rate (Hesperocorixa linnaei and female Sigara striata). These results suggest that standard tests may underestimate the pesticide effect in a natural predator environment for some species. While the effects of these interactions on SSRs were not strong and SSRs remained largely similar in the absence and presence of predation risk, some obvious species rank shifts occurred for growth rate which may potentially affect community structure through changed competitive strength. The study of SSRs and their dependency on biotic stressors may provide a simple conceptual and predictive framework to increase our understanding of how stressors like pesticides may differentially affect community structure in the absence and presence of another stressor.

Variation in aggressive behaviour in the poeciliid fish Brachyrhaphis episcopi: Population and sex differences

Aggression is often positively correlated with other behavioural traits such as boldness and activity levels. Comparisons across populations can help to determine factors that promote the evolution of such traits. We quantified these behaviours by testing the responses of wild-caught poeciliid fish, Brachyrhaphis episcopi, to mirror image stimuli. This species occurs in populations that experience either high or low levels of predation pressure. Previous studies have shown that B. episcopi from low predation environments are less bold than those that occur with many predators. We therefore predicted that fish from high predation populations would be more aggressive and more active than fish from low predation populations. However, we found the opposite - low predation fish approached a mirror and a novel object more frequently than high predation fish suggesting that 'boldness' and aggression were higher in low predation populations, and that population-level boldness measures may vary depending on context. When tested individually, low predation fish inspected their mirror image more frequently. Females, but not males, from low predation sites were also more aggressive towards their mirror image. Variation in female aggression may be driven by a trade-off between food availability and predation risk. This suggests that the relationship between aggression and boldness has been shaped by adaptation to environmental conditions, and not genetic constraints.