Sex, size, and prey caloric value affect diet specialization and consumption of an invasive prey by a native predator.

Escaping the control of natural enemies is thought to heavily influence the establishment success and impact of non-native species. Here, we examined how the profitability of alternative prey in combination with the presence of a competitor and predator aggressive behavior explain individual differences in diet specialization and the consumption of the invasive green porcelain crab Petrolisthes armatus by the native mud crab predator Panopeus herbstii. Results from bomb calorimetry estimates show that invasive P. armatus has high caloric value relative to alternative native prey. Laboratory assays indicated that specialization and consumption of invasive P. armatus was mostly exhibited by large, female P. herbstii, but the presence of a competitor and predator aggressiveness did not influence diet and the consumption of P. armatus. Thus, intrinsic factors (e.g., sex and body size) seem to explain consumption of P. armatus and dietary specialization in P. herbstii, more generally. Although there are still many predator individuals that do not consume P. armatus, the proportion of individuals that have begun to specialize on P. armatus suggests that for some, it has become more profitable relative to alternative native prey. Given the high caloric value of P. armatus, we suggest that it is likely that differences in the cost of its consumption, including attack, capture, and handling times relative to alternative prey, determine its net profitability to individual predators.

Behavioral differences within and among populations of an African cichlid found in divergent and extreme environments.

Animals are increasingly faced with human-induced stressors that vary in space and time, thus we can expect population-level divergence in behaviors that help animals to cope with environmental change. However, empirical evidence of behavioral trait divergence across environmental extremes is lacking. We tested for variation in behavioral traits among 2 populations of an African cichlid fish (Pseudocrenilabrus multicolor victoriae Seegers, 1990) that experience extremes of dissolved oxygen (DO) and turbidity and are known to vary in a number of physiological and life history traits associated with these stressors. Using a common garden rearing experiment, F1 progeny from wild-caught parents originating from a swamp (low DO, clear) and a river (high DO, turbid) were reared in high DO, clear water. Predator simulation assays were conducted to test for (1) variation in boldness, general activity, and foraging activity between populations, (2) differences in correlations between behaviors within and across populations, and (3) repeatability of behaviors. There was strong evidence for divergence between populations, with swamp fish being more bold (i.e., leaving refuge sooner after a simulated predator attack) and active (i.e., spent more time out of refuge) than river fish. Across populations there were positive correlations between foraging activity and both boldness and general activity; however, within populations, there was only a strong positive relationship between foraging activity and boldness in the river population. Here, we have demonstrated that populations that originate from drastically different environments can produce progeny that exhibit measurable differences in behaviors and their correlated relationships even when reared under common conditions.

Do native predators benefit from non-native prey?

Despite knowledge on invasive species' predatory effects, we know little of their influence as prey. Non-native prey should have a neutral to positive effect on native predators by supplementing the prey base. However, if non-native prey displace native prey, then an invader's net influence should depend on both its abundance and value relative to native prey. We conducted a meta-analysis to quantify the effect of non-native prey on native predator populations. Relative to native prey, non-native prey similarly or negatively affect native predators, but only when studies employed a substitutive design that examined the effects of each prey species in isolation from other prey. When native predators had access to non-native and native prey simultaneously, predator abundance increased significantly relative to pre-invasion abundance. Although non-native prey may have a lower per capita value than native prey, they seem to benefit native predators by serving as a supplemental prey resource.

Individual variation in foraging behavior reveals a trade-off between flexibility and performance of a top predator.

There is increasing evidence that behavioral flexibility is associated with the ability to adaptively respond to environmental change. Flexibility can be advantageous in some contexts such as exploiting novel resources, but it may come at a cost of accuracy or performance in ecologically relevant tasks, such as foraging. Such trade-offs may, in part, explain why individuals within a species are not equally flexible. Here, we conducted a reversal learning task and predation experiment on a top fish predator, the Northern pike (Esox lucius), to examine individual variation in flexibility and test the hypothesis that an individual's behavioral flexibility is negatively related with its foraging performance. Pikes were trained to receive a food reward from either a red or blue cup and then the color of the rewarded cup was reversed. We found that pike improved over time in how quickly they oriented to the rewarded cup, but there was a bias toward the color red. Moreover, there was substantial variation among individuals in their ability to overcome this red bias and switch from an unrewarded red cup to the rewarded blue cup, which we interpret as consistent variation among individuals in behavioral flexibility. Furthermore, individual differences in behavioral flexibility were negatively associated with foraging performance on ecologically relevant stickleback prey. Our data indicate that individuals cannot be both behaviorally flexible and efficient predators, suggesting a trade-off between these two traits.

Reciprocal behavioral plasticity and behavioral types during predator-prey interactions.

How predators and prey interact has important consequences for population dynamics and community stability. Here we explored how predator-prey interactions are simultaneously affected by reciprocal behavioral plasticity (i.e., plasticity in prey defenses countered by plasticity in predator offenses and vice versa) and consistent individual behavioral variation (i.e., behavioral types) within both predator and prey populations. We assessed the behavior of a predator species (northern pike) and a prey species (three-spined stickleback) during one-on-one encounters. We also measured additional behavioral and morphological traits in each species. Using structural equation modeling, we found that reciprocal behavioral plasticity as well as predator and prey behavioral types influenced how individuals behaved during an interaction. Thus, the progression and ultimate outcome of predator-prey interactions depend on both the dynamic behavioral feedback occurring during the encounter and the underlying behavioral type of each participant. We also examined whether predator behavioral type is underlain by differences in metabolism and organ size. We provide some of the first evidence that behavioral type is related to resting metabolic rate and size of a sensory organ (the eyes). Understanding the extent to which reciprocal behavioral plasticity and intraspecific behavioral variation influence the outcome of species interactions could provide insight into the maintenance of behavioral variation as well as community dynamics.

Climate and pH predict the potential range of the invasive apple snail (Pomacea insularum) in the southeastern United States.

Predicting the potential range of invasive species is essential for risk assessment, monitoring, and management, and it can also inform us about a species' overall potential invasiveness. However, modeling the distribution of invasive species that have not reached their equilibrium distribution can be problematic for many predictive approaches. We apply the modeling approach of maximum entropy (MaxEnt) that is effective with incomplete, presence-only datasets to predict the distribution of the invasive island apple snail, Pomacea insularum. This freshwater snail is native to South America and has been spreading in the USA over the last decade from its initial introductions in Texas and Florida. It has now been documented throughout eight southeastern states. The snail's extensive consumption of aquatic vegetation and ability to accumulate and transmit algal toxins through the food web heighten concerns about its spread. Our model shows that under current climate conditions the snail should remain mostly confined to the coastal plain of the southeastern USA where it is limited by minimum temperature in the coldest month and precipitation in the warmest quarter. Furthermore, low pH waters (pH <5.5) are detrimental to the snail's survival and persistence. Of particular note are low-pH blackwater swamps, especially Okefenokee Swamp in southern Georgia (with a pH below 4 in many areas), which are predicted to preclude the snail's establishment even though many of these areas are well matched climatically. Our results elucidate the factors that affect the regional distribution of P. insularum, while simultaneously presenting a spatial basis for the prediction of its future spread. Furthermore, the model for this species exemplifies that combining climatic and habitat variables is a powerful way to model distributions of invasive species.

Maternal exposure to predation risk decreases offspring antipredator behaviour and survival in threespined stickleback.

1. Adaptive maternal programming occurs when mothers alter their offspring's phenotype in response to environmental information such that it improves offspring fitness. When a mother's environment is predictive of the conditions her offspring are likely to encounter, such transgenerational plasticity enables offspring to be better-prepared for this particular environment. However, maternal effects can also have deleterious effects on fitness.2. Here, we test whether female threespined stickleback fish exposed to predation risk adaptively prepare their offspring to cope with predators. We either exposed gravid females to a model predator or not, and compared their offspring's antipredator behaviour and survival when alone with a live predator. Importantly, we measured offspring behaviour and survival in the face of the same type of predator that threatened their mothers (Northern pike).3. We did not find evidence for adaptive maternal programming; offspring of predator-exposed mothers were less likely to orient to the predator than offspring from unexposed mothers. In our predation assay, orienting to the predator was an effective antipredator behaviour and those that oriented, survived for longer.4. In addition, offspring from predator-exposed mothers were caught more quickly by the predator on average than offspring from unexposed mothers. The difference in antipredator behaviour between the maternal predator-exposure treatments offers a potential behavioural mechanism contributing to the difference in survival between maternal treatments.5. However, the strength and direction of the maternal effect on offspring survival depended on offspring size. Specifically, the larger the offspring from predator-exposed mothers, the more vulnerable they were to predation compared to offspring from unexposed mothers.6. Our results suggest that the predation risk perceived by mothers can have long-term behavioural and fitness consequences for offspring in response to the same predator. These stress-mediated maternal effects can have nonadaptive consequences for offspring when they find themselves alone with a predator. In addition, complex interactions between such maternal effects and offspring traits such as size can influence our conclusions about the adaptive nature of maternal effects.

Evolutionary game theory as a framework for studying biological invasions.

Although biological invasions pose serious threats to biodiversity, they also provide the opportunity to better understand interactions between the ecological and evolutionary processes structuring populations and communities. However, ecoevolutionary frameworks for studying species invasions are lacking. We propose using game theory and the concept of an evolutionarily stable strategy (ESS) as a conceptual framework for integrating the ecological and evolutionary dynamics of invasions. We suggest that the pathways by which a recipient community may have no ESS provide mechanistic hypotheses for how such communities may be vulnerable to invasion and how invaders can exploit these vulnerabilities. We distinguish among these pathways by formalizing the evolutionary contexts of the invader relative to the recipient community. We model both the ecological and the adaptive dynamics of the interacting species. We show how the ESS concept provides new mechanistic hypotheses for when invasions result in long- or short-term increases in biodiversity, species replacement, and subsequent evolutionary changes.

Behavioral correlations provide a mechanism for explaining high invader densities and increased impacts on native prey.

The fact that superabundant invasive pests are also sometimes highly aggressive represents an interesting paradox. Strong intraspecific aggression should result in high intraspecific competition and limit the densities reached by exotic species. One mechanism that can allow invaders to attain high densities despite high intraspecific aggression, involves positive correlations between aggression and other behaviors such as foraging activity. We conducted a mesocosm experiment to quantify the ecological implications of correlations between aggressiveness and foraging activity among groups of exotic signal crayfish (Pacifastacus leniusculus) at low and high densities. Our results showed that high invader densities increased intraspecific aggression and per capita interactions between crayfish, but also increased foraging activity and impacts on preferred prey. As a result, exotic crayfish did not show density-dependent reductions in per capita feeding or growth rates. We suggest that the positive correlation between aggression and activity is part of an aggression syndrome whereby some individuals are generally more aggressive/active than others across situations. An aggression syndrome can couple aggressive behaviors important to population establishment of invasive species with foraging activity that enhances the ability of invaders to attain high densities and have large impacts on invaded communities.