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.     

Predation, habitat complexity, and variation in density-dependent mortality of temperate reef fishes

Density dependence in demographic rates can strongly affect the dynamics of populations. However, the mechanisms generating density dependence (e.g., predation) are also dynamic processes and may be influenced by local conditions. Understanding the manner in which local habitat features affect the occurrence and/or strength of density dependence will increase our understanding of population dynamics in heterogeneous environments. In this study I conducted two separate field experiments to investigate how local predator density and habitat complexity affect the occurrence and form of density-dependent mortality of juvenile rockfishes (Sebastes spp.). I also used yearly censuses of rockfish populations on nearshore reefs throughout central California to evaluate mortality of juvenile rockfish at large spatial scales. Manipulations of predators (juvenile bocaccio, S. paucispinus) and prey (kelp, gopher, and black-and-yellow [KGB] rockfish, Sebastes spp.) demonstrated that increasing the density of predators altered their functional response and thus altered patterns of density dependence in mortality of their prey. At low densities of predators, the number of prey consumed per predator was a decelerating function, and mortality of prey was inversely density dependent. However, at high densities of predators, the number of prey killed per predator became an accelerating response, and prey mortality was directly density dependent. Results of field experiments and large-scale surveys both indicated that the strength of density-dependent mortality may also be affected by the structural complexity of the habitat. In small-scale field experiments, increased habitat complexity increased the strength of density-dependent mortality. However, at large scales, increasing complexity resulted in a decrease in the strength of density dependence. I suggest that these differences resulted from scale-dependent changes in the predatory response that generated mortality. Whether increased habitat complexity leads to an increase or a decrease in the strength of density-dependent mortality may depend on how specific predatory responses (e.g., functional or aggregative) are altered by habitat complexity. Overall, the findings of this study suggest that rates of demographic density dependence and the resulting dynamics of local populations may largely depend upon attributes of the local habitat.     

Evaluating impacts of forage fish abundance on marine predators

Forage fish—small, low trophic level, pelagic fish such as herrings, sardines, and anchovies—are important prey species in marine ecosystems and also support large commercial fisheries. In many parts of the world, forage fish fisheries are managed using precautionary principles that target catch limits below the maximum sustainable yield. However, there are increasing calls to further limit forage fish catch to safeguard their fish, seabird, and marine mammal predators. The effectiveness of these extra-precautionary regulations, which assume that increasing prey abundance increases predator productivity, are under debate. In this study, we used prey-linked population models to measure the influence of forage fish abundance on the population growth rates of 45 marine predator populations representing 32 fish, seabird, and mammal species from 5 regions around the world. We used simulated data to confirm the ability of the statistical model to accurately detect prey influences under varying levels of influence strength and process variability. Our results indicate that predator productivity was rarely influenced by the abundance of their forage fish prey. Only 6 predator populations (13% of the total) were positively influenced by increasing prey abundance and the model exhibited high power to detect prey influences when they existed. These results suggest that additional limitation of forage fish harvest to levels well below sustainable yields would rarely result in detectable increases in marine predator populations.     

The influence of intraguild predation on prey suppression and prey release: a meta-analysis

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with IGP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates. Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature.     

Species identity dominates the relationship between predator biodiversity and herbivore suppression

Agricultural pest suppression is an important ecosystem service that may be threatened by the loss of predator diversity. This has stimulated interest in the relationship between predator biodiversity and biological control. Multiple-predator studies have shown that predators may complement or interfere with one another, but few experiments have determined if the resulting effects on prey are caused by changes in predator abundance, identity, species richness, or some combination of these factors. We experimentally isolated the effect of predator species richness on the biological control of an important agricultural pest, the green peach aphid. We found no evidence that increasing predator species richness affects aphid biological control; overall there was no strong complementarity or interference among predator species that altered the strength of aphid suppression. Instead, our experiments revealed strong effects of predator species identity, because predators varied dramatically in their per capita consumption rates. Our results are consistent with other multiple-predator studies finding strong species-identity effects and suggest that, for the biological control of aphids, conservation strategies that directly target key species will be more effective than those targeting predator biodiversity more broadly.     

Regulation of intertidal food webs by avian predators on New England rocky shores

Although there is a large body of research on food webs in rocky intertidal communities, most of the emphasis has been on the marine benthic components. Effects of avian predation on highly mobile predators such as crabs, remains practically unstudied in rocky shore ecosystems. The crab, Cancer borealis, is an important component of the diet of gulls (Larus marinus, L. argentatus) at the Isles of Shoals, Maine, USA. C. borealis prey include the predatory gastropod Nucella lapillus L., the herbivore Littorina littorea, and mussels Mytilus edulis L. We hypothesized that gulls reduce abundance of C. borealis in the low intertidal and shallow subtidal, thereby allowing C. borealis prey to persist in high numbers. A study of crab tidal migration showed that C. borealis density nearly doubled at high tide compared to low tide; thus, crabs from a large subtidal source population migrate into the intertidal zone during high tides and either emigrate or are removed by gulls during low tides. Results from a small-scale (1 m2) predator caging experiment in the low intertidal zone indicated that enclosed crabs significantly reduced L. littorea abundance when protected from gull predation. In a much larger-scale gull exclusion experiment, densities of C. borealis increased significantly during low and high tides in exclosures relative to the controls. C. borealis density was inversely correlated with changes in the abundance of two mesopredators Carcinus maenas and Nucella lapillus, and with the space-occupier M. edulis. There was a similar negative correlation between abundance of C. borealis and the change in abundance of the herbivore L. littorea, but the trend was not significant. Mortality of tethered L. littorea was associated with C. borealis density across sites. However, preferred algae did not change in response to L. littorea density during the experiment. Thus, we found suggestive, but not conclusive, evidence for a three-level cascade involving gulls, crabs, and L. littorea. Our studies strongly suggest that gulls, as apex predators, generate three-level trophic cascades in rocky intertidal food webs by preventing the highly mobile subtidal predator, C. borealis, from establishing substantial populations in the low-mid intertidal zone thereby indirectly enhancing densities of two key mesopredators (N. lapillus, Carcinus) and blue mussels (M. edulis).     

Simulated predator extinctions: predator identity affects survival and recruitment of oysters

The rate of species loss is increasing at a global scale, and human-induced extinctions are biased toward predator species. We examined the effects of predator extinctions on a foundation species, the eastern oyster (Crassostrea virginica). We performed a factorial experiment manipulating the presence and abundance of three of the most common predatory crabs, the blue crab (Callinectes sapidus), stone crab (Menippe mercenaria), and mud crab (Panopeus herbstii) in estuaries in the eastern United States. We tested the effects of species richness and identity of predators on juvenile oyster survival, oyster recruitment, and organic matter content of sediment. We also manipulated the density of each of the predators and controlled for the loss of biomass of species by maintaining a constant mass of predators in one set of treatments and simultaneously using an additive design. This design allowed us to test the density dependence of our results and test for functional compensation by other species. The identity of predator species, but not richness, affected oyster populations. The loss of blue crabs, alone or in combination with either of the other species, affected the survival rate of juvenile oysters. Blue crabs and stone crabs both affected oyster recruitment and sediment organic matter negatively. Mud crabs at higher than ambient densities, however, could fulfill some of the functions of blue and stone crabs, suggesting a level of ecological redundancy. Importantly, the strong effects of blue crabs in all processes measured no longer occurred when individuals were present at higher-than-ambient densities. Their role as dominant predator is, therefore, dependent on their density within the system and the density of other species within their guild (e.g., mud crabs). Our findings support the hypothesis that the effects of species loss at higher trophic levels are determined by predator identity and are subject to complex intraguild interactions that are largely density dependent. Understanding the role of biodiversity in ecosystem functioning or addressing practical concerns, such as loss of predators owing to overharvesting, remains complicated because accurate predictions require detailed knowledge of the system and should be drawn from sound experimental evidence, not based on observations or generalized models.     

Geographical gradients in diet affect population dynamics of Canada lynx

Geographical gradients in the stability of cyclic populations of herbivores and their predators may relate to the degree of specialization of predators. However, such changes are usually associated with transition from specialist to generalist predator species, rather than from geographical variation in dietary breadth of specialist predators. Canada lynx (Lynx canadensis) and snowshoe hare (Lepus americanus) populations undergo cyclic fluctuations in northern parts of their range, but cycles are either greatly attenuated or lost altogether in the southern boreal forest where prey diversity is higher. We tested the influence of prey specialization on population cycles by measuring the stable carbon and nitrogen isotope ratios in lynx and their prey, estimating the contribution of hares to lynx diet across their range, and correlating this degree of specialization to the strength of their population cycles. Hares dominated the lynx diet across their range, but specialization on hares decreased in southern and western populations. The degree of specialization correlated with cyclic signal strength indicated by spectral analysis of lynx harvest data, but overall variability of lynx harvest (the standard deviation of natural-log-transformed harvest numbers) did not change significantly with dietary specialization. Thus, as alternative prey became more important in the lynx diet, the fluctuations became decoupled from a regular cycle but did not become less variable. Our results support the hypothesis that alternative prey decrease population cycle regularity but emphasize that such changes may be driven by dietary shifts among dominant specialist predators rather than exclusively through changes in the predator community.     

Testing a stochastic version of the Beddington–DeAngelis functional response in foraging shore crabs

Current behaviour-based interference models assume that the predator population is infinitely large and that interference is weak. While the realism of the first assumption is questionable, the second assumption conflicts with the purpose of interference models. Here, we tested a recently developed stochastic version of the Beddington–DeAngelis functional response—which applies to a finite predator population without assuming weak interference—against experimental data of shore crabs (Carcinus maenas) foraging on mussels (Mytilus edulis). We present an approximate maximum likelihood procedure for parameter estimation when only one focal individual is observed, and introduce ‘correction factors’ that capture the average behaviour of the competing but unobserved individuals. We used the method to estimate shore crab handling time, interaction time, and searching rates for prey and competitor. Especially the searching rates were sensitive to variation in prey and competitor density. Incorporating constant parameter values in the model and comparing observed and predicted feeding rates revealed that the predictive power of the model is high. Our stochastic version of the Beddington–DeAngelis model better reflects reality than current interference models and is also amenable for modelling effects of interference on predator distributions.     

Predators of juvenile sea urchins and the effect of habitat refuges

We evaluated the effects of potential predators from intertidal habitats on Strongylocentrotus purpuratus survival using laboratory experiments and assessed abundances of main predatory species along the Pacific coast of North America. The interactive effects of urchins’ and predators’ sizes in mediating predation were quantified. Habitat complexity (substrate pits, adult spine canopy) was manipulated to examine its effects on predation of most susceptible individuals (<14 mm). Pachygrapsus crassipes was identified as a major predator of urchins up to ≈30 mm. A positive effect of predator size on consumption of progressively larger urchins was detected, probably due to a mechanical limitation on crabs’ ability to consume large prey. Larger claws of males with respect to females of comparable sizes facilitated the handling of larger prey. Substrate refuges significantly reduced mortality on juvenile urchins. These results show that crab predation may be important in organizing intertidal communities, despite multiple ecological mechanisms promoting sea urchin survival.     

Resource identity modifies the influence of predation risk on ecosystem function

It is well established that predators can scare as well as consume their prey. In many systems, the fear of being eaten causes trait-mediated cascades whose strength can rival or exceed that of more widely recognized density-mediated cascades transmitted by predators that consume their prey. Despite this progress it is only beginning to be understood how the influence of predation risk is shaped by environmental context and whether it can exert an important influence on ecosystem-level processes. This study used a factorial mesocosm experiment that manipulated basal-resource identity (either barnacles, Semibalanus balanoides, or mussels, Mytilus edulis) to determine how resources modify the influence of predation risk, cascade strength, and the efficiency of energy transfer in two, tritrophic, rocky-shore food chains containing the predatory green crab (Carcinus maenas) and an intermediate consumer (the snail, Nucella lapillus). The effect of predation risk and the strength of trait-mediated cascades (both in absolute and relative terms) were much stronger in the barnacle than in the mussel food chain. Moreover, predation risk strongly diminished the efficiency of energy transfer in the barnacle food chain but had no significant effect in the mussel food chain. The influence of resource identity on indirect-effect strength and energy transfer was likely caused by differences in how each resource shapes the degree of risk perceived by prey. We suggest that our understanding of the connection between trophic dynamics and ecosystem functioning will improve considerably once the effects of predation risk on individual behavior and physiology are considered.     

Predator identity and additive effects in a treehole community

Multiple predator species can interact as well as strongly affect lower trophic levels, resulting in complex, nonadditive effects on prey populations and community structure. Studies of aquatic systems have shown that interactive effects of predators on prey are not necessarily predictable from the direct effects of each species alone. To test for complex interactions, the individual and combined effects of a top and intermediate predator on larvae of native and invasive mosquito prey were examined in artificial analogues of water-filled treeholes. The combined effects of the two predators were accurately predicted from single predator treatments by a multiplicative risk model, indicating additivity. Overall survivorship of both prey species decreased greatly in the presence of the top predator Toxorhynchites rutilus. By itself, the intermediate predator Corethrella appendiculata increased survivorship of the native prey species Ochlerotatus triseriatus and decreased survivorship of the invasive prey species Aedes albopictus relative to treatments without predators. Intraguild predation did not occur until alternative prey numbers had been reduced by approximately one-half. Owing to changes in size structure accompanying its growth, T. rutilus consumed more prey as time progressed, whereas C. appendiculata consumed less. The intermediate predator, C. appendiculata, changed species composition by preferentially consuming A. albopictus, while the top predator, T. rutilus, reduced prey density, regardless of species. Although species interactions were in most cases predicted from pairwise interactions, risk reduction from predator interference occurred when C. appendiculata densities were increased and when the predators were similarly sized.     

A review of predator diet effects on prey defensive responses

Numerous studies have examined how predator diets influence prey responses to predation risk, but the role predator diet plays in modulating prey responses remains equivocal. We reviewed 405 predator–prey studies in 109 published articles that investigated changes in prey responses when predators consumed different prey items. In 54 % of reviewed studies, prey responses were influenced by predator diet. The value of responding based on a predator’s recent diet increased when predators specialized more strongly on particular prey species, which may create patterns in diet cue use among prey depending upon whether they are preyed upon by generalist or specialist predators. Further, prey can alleviate costs or accrue greater benefits using diet cues as secondary sources of information to fine tune responses to predators and to learn novel risk cues from exotic predators or alarm cues from sympatric prey species. However, the ability to draw broad conclusions regarding use of predator diet cues by prey was limited by a lack of research identifying molecular structures of the chemicals that mediate these interactions. Conclusions are also limited by a narrow research focus. Seventy percent of reviewed studies were performed in freshwater systems, with a limited range of model predator–prey systems, and 98 % of reviewed studies were performed in laboratory settings. Besides identifying the molecules prey use to detect predators, future studies should strive to manipulate different aspects of prey responses to predator diet across a broader range of predator–prey species, particularly in marine and terrestrial systems, and to expand studies into the field.     

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.     

Modelling predation by transient leopard seals for an ecosystem-based management of Southern Ocean fisheries

Correctly quantifying the impacts of rare apex marine predators is essential to ecosystem-based approaches to fisheries management, where harvesting must be sustainable for targeted species and their dependent predators. This requires modelling the uncertainty in such processes as predator life history, seasonal abundance and movement, size-based predation, energetic requirements, and prey vulnerability. We combined these uncertainties to evaluate the predatory impact of transient leopard seals on a community of mesopredators (seals and penguins) and their prey at South Georgia, and assess the implications for an ecosystem-based management. The mesopredators are highly dependent on Antarctic krill and icefish, which are targeted by regional fisheries. We used a state-space formulation to combine (1) a mark-recapture open-population model and individual identification data to assess seasonally variable leopard seal arrival and departure dates, numbers, and residency times; (2) a size-based bioenergetic model; and (3) a size-based prey choice model from a diet analysis. Our models indicated that prey choice and consumption reflected seasonal changes in leopard seal population size and structure, size-selective predation and prey vulnerability. A population of 104 (90–125) leopard seals, of which 64% were juveniles, consumed less than 2% of the Antarctic fur seal pup production of the area (50% of total ingested energy, IE), but ca. 12–16% of the local gentoo penguin population (20% IE). Antarctic krill (28% IE) were the only observed food of leopard seal pups and supplemented the diet of older individuals. Direct impacts on krill and fish were negligible, but the “escapement” due to leopard seal predation on fur seal pups and penguins could be significant for the mackerel icefish fishery at South Georgia. These results suggest that: (1) rare apex predators like leopard seals may control, and may depend on, populations of mesopredators dependent on prey species targeted by fisheries; and (2) predatory impacts and community control may vary throughout the predator's geographic range, and differ across ecosystems and management areas, depending on the seasonal abundance of the prey and the predator's dispersal movements. This understanding is important to integrate the predator needs as natural mortality of its prey in models to set prey catch limits for fisheries. Reliable estimates of the variability of these needs are essential for a precautionary interpretation in the context of an ecosystem-based management.     

A simple plant mutation abets a predator-diversity cascade

Resource consumption often increases with greater consumer biodiversity. This could result either from complementarity among consumers or the inclusion of particular key species, and it is often difficult to differentiate between these two mechanisms. We exploited a simple plant mutation (reduced production of surface waxes) to alter foraging within a community of aphid predators, and thus perhaps shift the nature of resulting predator diversity effects. We found that greater predator species richness dramatically increased prey suppression and plant biomass only on mutant, reduced-wax pea plants (Pisum sativum). On pea plants from a sister line with wild type, waxier plant surfaces, predator species richness did not influence predators' impacts on herbivores or plants. Thus, a change in plant surface structure acted to turn on, or off, the cascading effects of predator diversity. Greater predator richness encouraged higher densities of true predators but did not lead to greater reproduction by a parasitoid, Aphidius ervi; fecundity of each natural enemy species was similar for the two plant types. Behavioral observations indicated that although A. ervi was less likely to forage within species-rich predator communities, low-wax plants mitigated this interference by encouraging generally greater A. ervi foraging and thus high rates of aphid dislodgement (aphids dropped from plants to escape A. ervi, but not the other predators). Thus, only species-rich, low-wax plants simultaneously encouraged strong species-specific effects of A. ervi, and strong complementarity among the other predator species. In summary, our study provides evidence that diversity effects in predator assemblages are sensitive to habitat characteristics. Further, we show that a simple plant morphological trait, controlled by a single gene mutation, can dramatically alter the cascading effects of predator species richness on herbivores and plants.     

Consequences of patch reef spacing for density-dependent mortality of coral-reef fishes

The spatial configuration of habitat patches can profoundly affect a number of ecological interactions, including those between predators and prey. I examined the effects of reef spacing on predator-prey interactions within coral-reef fish assemblages in the Bahamas. Using manipulative field experiments, I determined that reef spacing influences whether and how density-dependent predation occurs. Mortality rates of juveniles of two ecologically dissimilar species (beaugregory damselfish and yellowhead wrasse) were similarly affected by reef spacing; for both species, mortality was density dependent on reef patches that were spatially isolated (separated by 50 m), and density independent on reef patches that were aggregated (separated by 5 m). A subsequent experiment with the damselfish demonstrated that a common resident predator (coney) caused a substantial proportion of the observed mortality, independent of reef spacing. Compared to isolated reefs, aggregated reefs were much more likely to be visited by transient predators (mostly yellowtail snappers), regardless of prey density, and on these reefs, mortality rates approached 100% for both prey species. Transient predators exhibited neither an aggregative response nor a type 3 functional response, and consequently were not the source of density dependence observed on the isolated reefs. These patterns suggest that resident predators caused density-dependent mortality in their prey through type 3 functional responses on all reefs, but on aggregated reefs, this density dependence was overwhelmed by high, density-independent mortality caused by transient predators. Thus, the spatial configuration of reef habitat affected both the magnitude of total predation and the existence of density-dependent mortality. The combined effects of the increasing fragmentation of coral reef habitats at numerous scales and global declines in predatory fish may have important consequences for the regulation of resident fish populations.     

An unusual trophic subsidy and species dominance in a tropical stream

In classical theory, species are assumed to achieve dominance through competitive exclusion, but if food resources are limiting, cross-habitat trophic subsidies could also underpin dominance. The impact of dominant species on community dynamics may depend on the energy base of population size. We report on an unusual, spatially subsidized population of a tropical, stream-dwelling crab that dominates the benthic fauna of a Kenyan stream. Diet and stable isotope analyses indicated that this crab is a true omnivore, with terrestrial subsidies dominating both plant and animal resources. Unusually, the animal prey included almost no aquatic invertebrates. Instead, a single species of ant constituted approximately 35% of the annual diet (stomach contents analysis) and up to 90% of assimilated nitrogen (estimates from stable isotope analysis). Ants may be pivotal to enabling crab dominance, and this crab may be largely disconnected from the local trophic network for its dietary needs. The paucity of other invertebrates in the stream community suggests that this super-dominant crab is a strong interactor that suppresses aquatic invertebrate populations. Common stabilizing attributes of spatially subsidized food webs (e.g., asynchronous prey availability, wide feeding niche, consumer migration) were absent from this system, and although apparently stable, it may be vulnerable to disturbance in the donor habitat.     

Potential effects of a non-indigenous predator in its expanded range: assessing green crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, prey preference in a productive coastal area of Atlantic Canada

The non-indigenous green crab (Carcinus maenas) is an important predator on bivalve wild beds in coastal areas worldwide. This study explored size-dependent green crab prey preference on American oysters (Crassostrea virginica), blue mussels (Mytilus edulis), and soft-shell clams (Mya arenaria) in a productive coastal system of Atlantic Canada. Using two sizes of prey and three different experimental manipulations, small, medium, and large green crabs were given a choice among these three bivalves, and their daily feeding rates were monitored over the course of 3 days. For both prey sizes, green crabs showed an early feeding preference for soft-shell clams and, only as they declined in numbers, a switch toward mussels and subsequently toward oysters. We found that such changes in the timing (order) of prey preference are related to prey differences in shell thickness, a fairly reliable indicator of prey shell strength.     

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.