Predators can strongly influence prey populations not only through consumptive effects (CE) but also through non-consumptive effects (NCE) imposed by predation risk. Yet, the impact of NCE on bioenergetic and stoichiometric body contents of prey, traits that are shaping life histories, population and food web dynamics, is largely unknown. Moreover, the degree to which NCE can evolve and can drive evolution in prey populations is rarely studied. A 6-week outdoor mesocosm experiment with Caged-Fish (NCE) and Free-Ranging-Fish (CE and NCE) treatments was conducted to quantify and compare the effects of CE and NCE on population densities, bioenergetic and stoichiometric body contents of Daphnia magna, a keystone species in freshwater ecosystems. We tested for evolution of CE and NCE by using experimental populations consisting of D. magna clones from two periods of a resurrected natural pond population: a pre-fish period without fish and a high-fish period with high predation pressure. Both Caged-Fish and Free-Ranging-Fish treatments decreased the body size and population densities, especially in Daphnia from the high-fish period. Only the Free-Ranging-Fish treatment affected bioenergetic variables, while both the Caged-Fish and Free-Ranging-Fish treatments shaped body stoichiometry. The effects of CE and NCE were different between both periods indicating their rapid evolution in the natural resurrected population. Both the Caged-Fish and Free-Ranging-Fish treatments changed the clonal frequencies of the experimental Daphnia populations of the pre-fish as well as the high-fish period, indicating that not only CE but also NCE induced clonal sorting, hence rapid evolution during the mesocosm experiment in both periods. Our results demonstrate that CE as well as NCE have the potential to change not only the body size and population density but also the bioenergetic and stoichiometric characteristics of prey populations. Moreover, we show that these responses not only evolved in the studied resurrected population, but that CE and NCE also caused differential rapid evolution in a time frame of 6 weeks (ca. four to six generations). As NCE can evolve as well as can drive evolution, they may play an important role in shaping eco-evolutionary dynamics in predator-prey interactions.

Predator-prey interactions are a fundamental part of community ecology, yet the relative importance of consumptive and nonconsumptive effects (NCEs) (defined as a risk-induced response that alters prey fitness) has not been resolved. Theory suggests that the emergence and subsequent predominance of consumptive or NCEs depend on the given habitat\'s complexity as well as predator hunting mode and spatial domain sizes of both predator and prey, but their relative influence on the outcome of predator-prey interactions is unknown. We built agent-based models in NetLogo to simulate predator-prey interactions for three hunting modes-sit-and-wait, sit-and-pursue, and active-while concurrently simulating large versus small spatial domain sizes for both predators and prey. We studied (1) how hunting mode and spatial domain size interact to influence the emergence of consumptive or NCEs and (2) how, when NCEs do dominate, hunting mode and spatial domain separately or additively determine prey shifts in time, space, and habitat use. Our results indicate consumptive effects only dominate for active predators when prey habitat domains overlap completely with the predator\'s spatial domain and when sit-and-wait and sit-and-pursue predators and their prey both have large spatial domains. Prey are most likely to survive when they shift their time but most frequently shift their habitat. Our paper helps to better understand the underlying mechanisms that drive consumptive or NCEs to be most dominant.

Arctic terrestrial herbivores influence tundra carbon and nutrient dynamics through their consumption of resources, waste production, and habitat-modifying behaviors. The strength of these effects is likely to change spatially and temporally as climate change drives shifts in herbivore abundance, distribution, and activity timing. Here, we review how herbivores influence tundra carbon and nutrient dynamics through their consumptive and nonconsumptive effects. We also present evidence for herbivore responses to climate change and discuss how these responses may alter the spatial and temporal distribution of herbivore impacts. Several current knowledge gaps limit our understanding of the changing functional roles of herbivores; these include limited characterization of the spatial and temporal variability in herbivore impacts and of how herbivore activities influence the cycling of elements beyond carbon. We conclude by highlighting approaches that will promote better understanding of herbivore effects on tundra ecosystems, including their integration into existing biogeochemical models, new applications of remote sensing techniques, and the continued use of distributed experiments.

Omnivory is ubiquitous in ecological communities. Yet, we lack a consensus of how plant alternative resources impact the ability of omnivores to suppress prey populations. Previous work suggests that plant alternative resources can increase, decrease, or have no effect on the magnitude of omnivore-prey interactions. This discrepancy may arise from (1) the ability of omnivore populations to respond to plant alternative resources and (2) identity-specific effects of plant alternative resources. We used a meta-analysis to examine how omnivore population responses and the identity of plant alternative resources affect (1) omnivore predation rates (mainly reported as per capita predation rate) and (2) omnivore impacts on prey population density. Plant alternative resources reduced omnivore predation rate regardless of identity. The suppression of the predation rate by flowers and flowering plants was magnified when pollen alone was tested as the alternative resource. Surprisingly, plant alternative resource availability reduced prey density, suggesting that omnivore predation increased with plant alternative resources. This discrepancy (plant alternative resources not only decreased omnivore predation rates but also decreased prey density) resulted from experimental differences in the ability of omnivore populations to respond to plant alternative resources. In the presence of plant alternative resources, allowing omnivore population responses decreased prey density, while not allowing population responses increased prey density. Because omnivores commonly suppress prey density in the presence of plant alternative resources when population responses of omnivores are allowed, the effectiveness of biological control may depend upon the availability of such resources and the facilitation of population responses.

Invasive predators are known to have negative consumptive and non-consumptive effects on native species, but few examples show how the abundance of native prey may influence an established invasive predator. We compared invasive brown treesnakes (Boiga irregularis; BTS) found in caves occupied by endangered Mariana swiftlets (Aerodramus bartschi) to snakes found in nearby forests and caves without birds to quantify how the abundance of native avian prey impacts BTS abundance and behavior on Guam. From 2011 to 2017 we removed 151 BTS in caves occupied by swiftlets and never observed BTS in caves without birds. Notable locations included snakes foraging near swiftlets and in holes that allowed cave access and escape from capture. Of 43 BTS with gut contents, 27 (63%) contained swiftlets. BTS in swiftlet-occupied caves had greater fat mass compared to forests, indicating access to swiftlets may increase body condition and promote reproduction. Number of ovarian follicles was significantly greater in female snakes from swiftlet-occupied caves compared to those from ravine, but not limestone forests; evidence of male BTS being more capable of reproduction was limited (i.e., fewer non-discernible but not significantly larger testes in snakes from caves). Assuming other limiting factors are considered, altering the functional response of predators through the modification of caves or interdiction lures to exclude or hinder the largest BTS could bolster swiftlet populations by increasing nesting refugia in currently-occupied caves and facilitate recolonization of historical caves.

Indirect effects are ubiquitous in nature, and have received much attention in terrestrial plant-insect herbivore-enemy systems. In such tritrophic systems, changes in plant quality can have consequential effects on the behavior and abundance of insect predators and parasitoids. Plant quality as perceived by insect herbivores may vary for a range of reasons, including because of infection by plant pathogens. However, plant diseases vary in their origin (viral, bacterial or fungal) and as a result may have differing effects on plant physiology. To investigate if the main groups of plant pathogens differ in their indirect effects on higher trophic levels, we performed a meta-analysis using 216 measured responses from 29 primary studies. There was no overall effect of plant pathogens on natural enemy traits as differences between pathogen types masked their effects. Infection by fungal plant pathogens showed indirect negative effects on the performance and preference of natural enemies via both chewing and piercing-sucking insect herbivore feeding guilds. Infection by bacterial plant pathogens had a positive effect on the natural enemies (parasitoids) of chewing herbivores. Infection by viral plant pathogens showed no clear effect, although parasitoid preference may be positively affected by their presence. It is important to note that given the limited volume of studies to date on such systems, this work should be considered exploratory. Plant pathogens are very common in nature, and tritrophic systems provide an elegant means to examine the consequences of indirect interactions in ecology. We suggest that further studies examining how plant pathogens affect higher trophic levels would be of considerable value.

Trophic cascades have become a dominant paradigm in ecology, yet considerable debate remains about the relative strength of density- (consumptive) and trait-mediated (non-consumptive) effects in trophic cascades. This debate may, in part, be resolved by considering prey experience, which shapes prey traits (through genetic and plastic change) and influences prey survival (and therefore density). Here, we investigate the cascading role of prey experience through the addition of mosquitofish (Gambusia affinis) from predator-experienced or predator-naïve sources to mesocosms containing piscivorous largemouth bass (Micropterus salmoides), zooplankton, and phytoplankton. These two sources were positioned along a competition-defense tradeoff. Results show that predator-naïve mosquitofish suffered higher depredation rates, which drove a density-mediated cascade, whereas predator-experienced mosquitofish exhibited higher survival but fed less, which drove a trait-mediated cascade. Both cascades were similar in strength, leading to indistinguishable top-down effects on lower trophic levels. Therefore, the accumulation of prey experience with predators can cryptically shift cascade mechanisms from density- to trait-mediated.

Many studies have demonstrated that the nonconsumptive effect (NCE) of predators on prey traits can alter prey demographics in ways that are just as strong as the consumptive effect (CE) of predators. Less well studied, however, is how the CE and NCE of multiple predator species can interact to influence the combined effect of multiple predators on prey mortality. We examined the extent to which the NCE of one predator altered the CE of another predator on a shared prey and evaluated whether we can better predict the combined impact of multiple predators on prey when accounting for this influence. We conducted a set of experiments with larval dragonflies, adult newts (a known keystone predator), and their tadpole prey. We quantified the CE and NCE of each predator, the extent to which NCEs from one predator alters the CE of the second predator, and the combined effect of both predators on prey mortality. We then compared the combined effect of both predators on prey mortality to four predictive models. Dragonflies caused more tadpoles to hide under leaf litter (a NCE), where newts spend less time foraging, which reduced the foraging success (CE) of newts. Newts altered tadpole behavior but not in a way that altered the foraging success of dragonflies. Our study suggests that we can better predict the combined effect of multiple predators on prey when we incorporate the influence of interactions between the CE and NCE of multiple predators into a predictive model. In our case, the threat of predation to prey by one predator reduced the foraging efficiency of a keystone predator. Consequently, the ability of a predator to fill a keystone role could be compromised by the presence of other predators.