Predator hunting mode and habitat domain alter nonconsumptive effects in predator-prey interactions

Predators can affect prey populations through changes in traits that reduce predation risk. These trait changes (nonconsumptive effects, NCEs) can be energetically costly and cause reduced prey activity, growth, fecundity, and survival. The strength of nonconsumptive effects may vary with two functional characteristics of predators: hunting mode (actively hunting, sit-and-pursue, sit-and-wait) and habitat domain (the ability to pursue prey via relocation in space; can be narrow or broad). Specifically, cues from fairly stationary sit-and-wait and sit-and-pursue predators should be more indicative of imminent predation risk, and thereby evoke stronger NCEs, compared to cues from widely ranging actively hunting predators. Using a meta-analysis of 193 published papers, we found that cues from sit-and-pursue predators evoked stronger NCEs than cues from actively hunting predators. Predator habitat domain was less indicative of NCE strength, perhaps because habitat domain provides less reliable information regarding imminent risk to prey than does predator hunting mode. Given the importance of NCEs in determining the dynamics of prey communities, our findings suggest that predator characteristics may be used to predict how changing predator communities translate into changes in prey. Such knowledge may prove particularly useful given rates of local predator change due to habitat fragmentation and the introduction of novel predators.     

Modulation of predator-prey interactions by the Allee effect

An Allee effect arising from density-dependent mating success can have significant impacts at the ecosystem level when considered in the context of predator-prey interactions. These are captured by a mathematical model for the exchange of biomass between a structured predator population (continuous weight distribution) and a resource. Because the predator’s mating success affects the amount of resources required for the production of offsprings and their future growth into mature organisms, it influences the flux of biomass between trophic levels. Under simple assumptions, the equations can be reduced to an equivalent unstructured predator-prey model in which the Allee effect modulates the predation rate: the mating probability multiplies the rate of predator growth as well as the rate of resource depletion. Implications of the Allee effect for the bifurcation structure and equilibrium densities are examined. The model is compared to a modified version in which the Allee effect instead modulates the assimilation efficiency, hence the mating probability does not appear in the dynamical equation for the resource density. Both models exhibit qualitatively similar dynamics. However, compared to the model in which the Allee effect modulates predation, the model in which the Allee effect modulates assimilation efficiency predicts (i) unrealistically inefficient resource assimilation when predator density is low, (ii) a higher risk of catastrophic extinction resulting from a change in the parameter controlling the strength of the Allee effect, and (iii) no possibility of an increase in population size when the density dependence is enhanced.     

The influence of size-specific indirect interactions in predator-prey systems

Nonlethal indirect interactions between predators often lead to nonadditive effects of predator number on prey survival and growth. Previous studies have focused on systems with at least two different predator species and one prey species. However, most predators undergo extreme ontological changes in phenotype such that interactions between different-sized cohorts of a predator and its prey could lead to nonadditive effects in systems with only two species. This may be important since different-sized individuals of the same species can differ more in their ecology than similar-sized individuals of different species. This study examined trait-mediated indirect effects in a two-species system including a cannibalistic predator with different-sized cohorts and its prey. I tested for these effects using larvae of two stream salamanders, Gyrinophilus porphyriticus (predator) and Eurycea cirrigera (prey), by altering the densities and combinations of predator size classes in experimental streams. Results showed that the presence of large individuals can significantly reduce the impact of density changes of smaller conspecifics on prey survival through nonlethal means. In the absence of large conspecifics, an increase in the relative frequency of small predators significantly increased predation rates, thereby reducing prey survival. However, with large conspecifics present, increasing the density of small predators did not decrease prey survival, resulting in a 14.3% lower prey mortality than predicted from the independent effects of both predator size classes. Small predators changed their microhabitat use in the presence of larger conspecifics. Prey individuals reduced activity in response to large predators but did not respond to small predators. Both predators reduced prey growth. These results demonstrate that the impact of a predator can be significantly altered by two different types of trait-mediated indirect effects in two-species systems: between different-sized cohorts and between different cohorts and prey. This study demonstrates that predictions based on simple numerical changes that assume independent effects of different size classes or ignore size structure can be strongly misleading. We need to account for the size structure within predator populations in order to predict how changes in predator abundance will affect predator-prey dynamics.     

Landscape of fear influences the relative importance of consumptive and nonconsumptive predator effects

Predators can initiate trophic cascades by consuming and/or scaring their prey. Although both forms of predator effect can increase the overall abundance of prey's resources, nonconsumptive effects may be more important to the spatial and temporal distribution of resources because predation risk often determines where and when prey choose to forage. Our experiment characterized temporal and spatial variation in the strength of consumptive and nonconsumptive predator effects in a rocky intertidal food chain consisting of the predatory green crab (Carcinus maenas), an intermediate consumer (the dogwhelk, Nucella lapillus), and barnacles (Semibalanus balanoides) as a resource. We tracked the survival of individual barnacles through time to map the strength of predator effects in experimental communities. These maps revealed striking spatiotemporal patterns in Nucella foraging behavior in response to each predator effect. However, only the nonconsumptive effect of green crabs produced strong spatial patterns in barnacle survivorship. Predation risk may play a pivotal role in determining the small-scale distribution patterns of this important rocky intertidal foundation species. We suggest that the effects of predation risk on individual foraging behavior may scale up to shape community structure and dynamics at a landscape level.     

Consumptive and nonconsumptive effects of predators on metacommunities of competing prey

Although predators affect prey both via consumption and by changing prey migration behavior, the interplay between these two effects is rarely incorporated into spatial models of predator-prey dynamics and competition among prey. We develop a model where generalist predators have consumptive effects (i.e., altering the likelihood of local prey extinction) as well as nonconsumptive effects (altering the likelihood of colonization) on spatially separated prey populations (metapopulations). We then extend this model to explore the effects of predators on competition among prey. We find that generalist predators can promote persistence of prey metapopulations by promoting prey colonization, but predators can also hasten system-wide extinction by either increasing local extinction or reducing prey migration. By altering rates of prey migration, predators in one location can exert remote control over prey dynamics in another location via predator-mediated changes in prey flux. Thus, the effect of predators may extend well beyond the proportion of patches they visit. In the context of prey metacommunities, predator-mediated shifts in prey migration and mortality can shift the competition-colonization trade-off among competing prey, leading to changes in the prey community as well as changes in the susceptibility of prey species to habitat loss. Consequently, native prey communities may be susceptible to invasion not only by exotic prey species that experience reduced amounts of mortality from resident predators, but also by exotic prey species that exhibit strong dispersal in response to generalist native predators. Ultimately, our work suggests that the consumptive and nonconsumptive effects of generalist predators may have strong, yet potentially cryptic, effects on competing prey capable of mediating coexistence, fostering invasion, and interacting with anthropogenic habitat alteration.     

Impact of cannibalism on predator-prey dynamics: size-structured interactions and apparent mutualism

Direct and indirect interactions between two prey species can strongly alter the dynamics of predator-prey systems. Most predators are cannibalistic, and as a consequence, even systems with only one predator and one prey include two prey types: conspecifics and heterospecifics. The effects of the complex direct and indirect interactions that emerge in such cannibalistic systems are still poorly understood. This study examined how the indirect interaction between conspecific and heterospecific prey affects cannibalism and predation rates and how the direct interactions between both species indirectly alter the effect of the cannibalistic predator. I tested for these effects using larvae of the stream salamanders Eurycea cirrigera (prey) and Pseudotriton ruber (cannibalistic predator) by manipulating the relative densities of the conspecific and heterospecific prey in the presence and absence of the predator in experimental streams. The rates of cannibalism and heterospecific predation were proportional to the respective densities and negatively correlated, indicating a positive indirect interaction between conspecific and heterospecific prey, similar to "apparent mutualism." Direct interactions between prey species did not alter the effect of the predator. Although both types of prey showed a similar 30% reduction in night activity and switch in microhabitat use in response to the presence of the predator, cannibalism rates were three times higher than heterospecific predation rates irrespective of the relative densities of the two types of prey. Cumulative predation risks differed even more due to the 48% lower growth rate of conspecific prey. Detailed laboratory experiments suggest that the 3:1 difference in cannibalism and predation rate was due to the higher efficiency of heterospecific prey in escaping immediate attacks. However, no difference was observed when the predator was a closely related salamander species, Gyrinophilus porphyriticus, indicating that this difference is species specific. This demonstrates that cannibalism can result in the coupling of predator and prey mortality rates that strongly determines the dynamics of predator-prey systems.     

Tidal regime dictates the cascading consumptive and nonconsumptive effects of multiple predators on a marsh plant

Prey perception of predators can dictate how prey behaviorally balance the need to avoid being eaten with the need to consume resources, and this perception and consequent behavior can be strongly influenced by physical processes. Physical factors, however, can also alter the density and diversity of predators that pursue prey. Thus, it remains uncertain to what extent variable risk perception and antipredator behavior vs. variation in predator consumption of prey underlie prey-resource dynamics and give rise to large-scale patterns in natural systems. In an experimental food web where tidal inundation of marsh controls which predators access prey, crab and conch (predators) influenced the survivorship and antipredator behavior of snails (prey) irrespective of whether tidal inundation occurred on a diurnal or mixed semidiurnal schedule. Specifically, cues of either predator caused snails to ascend marsh leaves; snail survivorship was reduced more by unrestrained crabs than by unrestrained conchs; and snail survivorship was lowest with multiple predators than with any single predator despite interference. In contrast to these tidally consistent direct consumptive and nonconsumptive effects, indirect predator effects differed with tidal regime: snail grazing of marsh leaves in the presence of predators increased in the diurnal tide but decreased in the mixed semidiurnal tidal schedule, overwhelming the differences in snail density that resulted from direct predation. In addition, results suggest that snails may increase their foraging to compensate for stress-induced metabolic demand in the presence of predator cues. Patterns from natural marshes spanning a tidal inundation gradient (from diurnal to mixed semidiurnal tides) across 400 km of coastline were consistent with experimental results: despite minimal spatial variation in densities of predators, snails, abiotic stressors, and marsh productivity, snail grazing on marsh plants increased and plant biomass decreased on shorelines exposed to a diurnal tide. Because both the field and experimental results can be explained by tidal-induced variation in risk perception and snail behavior rather than by changes in snail density, this study reinforces the importance of nonconsumptive predator effects in complex natural systems and at large spatial scales.     

The role of turbidity as a constraint on predator-prey interactions in aquatic environments

Many of the world's most productive aquatic ecosystems usually contain turbid water. Paradoxically, many fish species that live in these habitats are also those that often rely on vision to detect their predators and their prey. For these fish, turbidity will reduce the distance at which predator-prey interactions occur, and there should be a reduction in the opportunity for behavioural modification to control the risk of predation. Under these conditions, we predict that most antipredator behaviour will become ineffective and that predator-prey interactions in turbid water will be primarily characterised by the direct effects of predator consumption of prey, rather than behavioural modification reducing the growth rates of prey. This hypothesis was tested in two laboratory experiments. The first experiment investigated how water turbidity, risk of predation, and their interaction affect habitat choice decisions by fathead minnows (Pimephales promelas). These data demonstrate that fathead minnows reduce their use of dangerous habitats, but that this effect is diminished in turbid water. A second experiment examined mortality patterns when these fish were preyed upon by yellow perch (Perca flavescens) in clear and turbid water. Absolute mortality rates were unaffected by visibility, but patterns of mortality were random in turbid water and skewed towards smaller individuals in clear water. Combined, these results support our hypothesis and suggest that the impact of predation risk will be reduced in turbid aquatic ecosystems. Received: 23 May 1996 / Accepted after revision: 18 November 1996     

Revisiting the evolution of ecological specialization, with emphasis on insect-plant interactions

Ecological specialization is a fundamental and well-studied concept, yet its great reach and complexity limit current understanding in important ways. More than 20 years after the publication of D. J. Futuyma and G. Moreno's oft-cited, major review of the topic, we synthesize new developments in the evolution of ecological specialization. Using insect-plant interactions as a model, we focus on important developments in four critical areas: genetic architecture, behavior, interaction complexity, and macroevolution. We find that theory based on simple genetic trade-offs in host use is being replaced by more subtle and complex pictures of genetic architecture, and multitrophic interactions have risen as a necessary framework for understanding specialization. A wealth of phylogenetic data has made possible a more detailed consideration of the macroevolutionary dimension of specialization, revealing (among other things) bidirectionality in transitions between generalist and specialist lineages. Technological advances, including genomic sequencing and analytical techniques at the community level, raise the possibility that the next decade will see research on specialization spanning multiple levels of biological organization in non-model organisms, from genes to populations to networks of interactions in natural communities. Finally, we offer a set of research questions that we find to be particularly pressing and fruitful for future research on ecological specialization.     

Aggressive mimicry,prey-specific predatory behaviour and predator-recognition in the predator-prey interactions of Portia fimbriata and Euryattus sp., jumping spiders from Queensland

Summary Adults and large juveniles of Queensland Portia fimbriata, a salticid spider known to prey on other spiders (including other salticids), are shown to use prey-specific predatory behaviour against Euryattus sp., one of the salticids on which it feeds. Euryattus females are unusual because they nest inside suspended rolled-up leaves. P. fimbriata used vibratory displays to lure Euryattus females from their nests. These displays seem to mimic the courtship displays of Euryattus males. Other species of Portia and other populations of P. fimbriata, in habitats in which Euryattus is not known to occur, did not practise this prey-specific behaviour. In the laboratory, Euryattus — but not Jacksonoides queenslandica, another salticid on which P. fimbriata is known to feed — readily recognized approaching Portia as a potential predator. A possible evolutionary arms race between Portia and Euryattus is discussed. Offrint request to: R.R. Jackson     

Biological interactions and environmental effects in the economics of pest control

The problem of pest control is tackled in a context of an ecosystem that consists of prey-predator populations with human interaction through pesticide application. The control, aimed at reducing pest damage, results in two undesirable external effects: reduction of beneficial predator population and environmental contamination. The untapped natural equilibrium is compared with equilibrium resulting from decentralized and centralized economic decision making. It is shown that, under certain conditions, myopic decision rules increase rather than decrease the pest damage. The “user cost” (or benefit) is shown to be crucial in determining the optimal centralized policy, and its relations to the various components of the system are analyzed. The components of the user cost are analyzed to determine the level of taxes or subsidies that will yield the optimal policy.     

Demographic stochasticity reduces the synchronizing effect of dispersal in predator-prey metapopulations

Dispersal may affect predator-prey metapopulations by rescuing local sink populations from extinction or by synchronizing population dynamics across the metapopulation, increasing the risk of regional extinction. Dispersal is likely influenced by demographic stochasticity, however, particularly because dispersal rates are often very low in metapopulations. Yet the effects of demographic stochasticity on predator-prey metapopulations are not well known. To that end, I constructed three models of a two-patch predator-prey system. The models constitute a hierarchy of complexity, allowing direct comparisons. Two models included demographic stochasticity (pure jump process [PJP] and stochastic differential equations [SDE]), and the third was deterministic (ordinary differential equations [ODE]). One stochastic model (PJP) treated population sizes as discrete, while the other (SDE) allowed population sizes to change continuously. Both stochastic models only produced synchronized predator-prey dynamics when dispersal was high for both trophic levels. Frequent dispersal by only predators or prey in the PJP and SDE spatially decoupled the trophic interaction, reducing synchrony of the non-dispersive species. Conversely, the ODE generated synchronized predator-prey dynamics across all dispersal rates, except when initial conditions produced anti-phase transients. These results indicate that demographic stochasticity strongly reduces the synchronizing effect of dispersal, which is ironic because demographic stochasticity is often invoked post hoc as a driver of extinctions in synchronized metapopulations.     

Understanding the effects of rainfall on elephant-vegetation interactions around waterholes

The distribution of surface water affects herbivore-vegetation interactions in arid and semi-arid regions. Limited access to surface water typically results in the emergence of vegetation gradients around natural and artificial water sources. In particular, African elephants can create large-scale gradients of woody vegetation. Understanding the dynamics of these gradients is of particular importance for the conservation of other, less mobile herbivores that depend on woody vegetation in areas close to water. While rainfall is known to be a key determinant of herbivore-vegetation interactions in dry areas, we only have limited understanding on how it impacts woody vegetation gradients around waterholes. To address this problem, we developed a deterministic simulation model that describes the interplay of rainfall, elephants and woody vegetation in the vicinity of waterholes. The model is based on elephant telemetry data and the ecological conditions in Etosha National Park (ENP), Namibia. We found that decreasing amounts of rainfall led to an increased degradation of woody vegetation, which was particularly severe in areas close to water. Based on this result we conclude that low rainfall was an important driver of recently observed patterns of vegetation degradation in ENP. More generally, rainfall appears to be a key factor that determines elephant-vegetation interactions and thus dynamics of woody vegetation gradients around waterholes. Using long-term rainfall data from ENP, we also demonstrate that an increase in the number of water sources during periods of low rainfall can mitigate the destructive impact of elephants in areas close to water. However, more research is required to assess the sustainability and effectiveness of rainfall-adapted strategies of artificial water provisioning in more detail. In particular it is important to investigate potential effects on elephant population dynamics.     

Harvesting creates ecological traps: consequences of invisible mortality risks in predator-prey metacommunities

Models of two-patch predator-prey metacommunities are used to explore how the global predator population changes in response to additional mortality in one of the patches. This could describe the dynamics of a predator in an environment that includes a refuge area where that predator is protected and a spatially distinct ("risky") area where it is harvested. The predator's movement is based on its perceived fitness in the two patches, but the risk from the additional mortality is potentially undetectable; this often occurs when the mortality is from human harvesting or from a novel type of top predator. Increases in undetected mortality in the risky area can produce an abrupt collapse of either the refuge population or of the entire predator population when the mortality rate exceeds a threshold level. This is due to the attraction of the risky patch, which has abundant prey due to its high predator mortality. Extinction of the refuge predator population does not occur when the refuge patch has a higher maximum per capita predator growth rate than the exploited patch because the refuge is then more attractive when the predator is rare. The possibility of abrupt extinction of one or both patches from high densities in response to a small increase in harvest is often associated with alternative states. In such cases, large reductions in mortality may be needed to avoid extinction in a collapsing predator population, or to reestablish an extinct population. Our analysis provides a potential explanation for sudden collapses of harvested populations, and it argues for more consideration of adaptive movement in designing protected areas.     

Ecosystem-based management of predator-prey relationships: piscivorous birds and salmonids.

Predator-prey relationships are often altered as a result of human activities. Where prey are legally protected, conservation action may include lethal predator control. In the Columbia River basin (Pacific Northwest, USA and Canada), piscivorous predators have been implicated in contributing to a lack of recovery of several endangered anadromous salmonids (Oncorhynchus spp.), and lethal and nonlethal control programs have been instituted against both piscine and avian species. To determine the consequences of avian predation, we used a bioenergetics approach to estimate the consumption of salmonid smolts by waterbirds (Common Merganser, California and Ring-billed Gull, Caspian Tern, Double-crested Cormorant) found in the mid-Columbia River from April through August, 2002-2004. We used our model to explore several predator-prey scenarios, including the impact of historical bird abundance, and the effect of preserving vs. removing birds, on smolt abundance. Each year, <1% of the estimated available salmonid smolts (interannual range: 44,830-109,209; 95% CI = 38,000-137,000) were consumed, 85-98% away from dams. Current diet data combined with historical gull abundance at dams suggests that past smolt consumption may have been 1.5-3 times current numbers, depending on the assumed distribution of gulls along the reaches. After the majority (80%) of salmonid smolts have left the study area, birds switch their diet to predominantly juvenile northern pikeminnow (Ptychocheilus oregonensis), which as adults are significant native salmonid predators in the Columbia River. Our models suggest that one consequence of removing birds from the system may be increased pikeminnow abundance, which--even assuming 80% compensatory mortality in juvenile pikeminnow survival--would theoretically result in an annual average savings of just over 180,000 smolts, calculated over a decade. Practically, this suggests that smolt survival could be maximized by deterring birds from the river when smolts are present, allowing bird presence after the diet switch to act as a tool for salmonid-predator control, and conducting adult-pikeminnow control throughout. Our analysis demonstrates that identifying the strength of ecosystem interactions represents a top priority when attempting to manage the abundance of a particular ecosystem constituent, and that the consequences of a single-species view may be counterintuitive, and potentially counterproductive.     

Stormwater treatment: examples of computational fluid dynamics modeling

Control of rainfall-runoff particulate matter (PM) and PM-bound chemical loads is challenging; in part due to the wide gradation of PM complex geometries of many unit operations and variable flow rates. Such challenges and the expense associated with resolving such challenges have led to the relatively common examination of a spectrum of unit operations and processes. This study applies the principles of computational fluid dynamics (CFD) to predict the particle and pollutant clarification behavior of these systems subject to dilute multiphase flows, typical of rainfall-runoff, within computationally reasonable limits, to a scientifically acceptable degree of accuracy. The Navier-Stokes (NS) system of nonlinear partial differential equations for multiphase hydrodynamics and separation of entrained particles are solved numerically over the unit operation control volume with the boundary and initial conditions defined and then solved numerically until the desired convergence criteria are met. Flow rates examined are scaled based on sizing of common unit operations such as hydrodynamic separators (HS), wet basins, or filters, and are examined from 1 to 100 percent of the system maximum hydraulic operating flow rate. A standard turbulence model is used to resolve flow, and a discrete phase model (DPM) is utilized to examine the particle clarification response. CFD results closely follow physical model results across the entire range of flow rates. Post-processing the CFD predictions provides an in-depth insight into the mechanistic behavior of unit operations by means of three dimensional (3-D) hydraulic profiles and particle trajectories. Results demonstrate the role of scour in the rapid degradation of unit operations that are not maintained. Comparisons are provided between measured and CFD modeled results and a mass balance error is identified. CFD is arguably the most powerful tool available for our profession since continuous simulation modeling.     

From interference to predation: type and effects of direct interspecific interactions of small mammals

Indirect exploitative competition, direct interference and predation are important interactions affecting species coexistence. These interaction types may overlap and vary with the season and life-history state of individuals. We studied effects of competition and potential nest predation by common shrews (Sorex araneus) on lactating bank voles (Myodes glareolus) in two seasons. The species coexist and may interact aggressively. Additionally, shrews can prey on nestling voles. We studied bank vole mothers’ spatial and temporal adaptations to shrew presence during summer and autumn. Further, we focused on fitness costs, e.g. decreased offspring survival, which bank voles may experience in the presence of shrews. In summer, interference with shrews decreased the voles’ home ranges and they spent more time outside the nest, but there were no effects on offspring survival. In autumn, we found decreased offspring survival in enclosures with shrews, potentially due to nest predation by shrews or by increased competition between species. Our results indicate a shift between interaction types depending on seasonal constraints. In summer, voles and shrews seem to interact mainly by interference, whereas resource competition and/or nest predation by shrews gain importance in autumn. Different food availability, changing environmental conditions and the energetic constraints in voles and shrews later in the year may be the reasons for the varying combinations of interaction types and their increasing effects on the inclusive fitness of bank voles. Our study provides evidence for the need of studies combining life history with behavioural measurements and seasonal constraints.