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Abstract: Predators can impact competition among prey by altering prey density via consumption or by causing prey to modify their traits or foraging behavior. Yet, differences between these two mechanisms may lead to different cascading impacts on lower trophic levels. Using a crab‐snail‐barnacle rocky intertidal food chain, we tested the effects of predation risk from crabs (top predators) on intraspecific competition among snails (intermediate consumers) and emergent indirect effects on the density of and competition among barnacles (basal resources). The per capita foraging and growth rates of snails declined with high conspecific density. Predation risk from crabs, which caused even larger reductions in snail foraging and growth, weakened competition among snails, whereas a 45% increase in barnacle density had no detectable effect on snail competition. Intraspecific competition between barnacles, however, depended on the interactive effects of barnacle density, snail density, and crab predation risk. Barnacles developed hummocking morphologies as they grew and competed for space. Hummock formation (a proxy for competition) increased as a result of either greater initial barnacle density or reduced snail foraging pressure, but these effects depended on predation risk. The effects of crab predation risk on snail foraging behavior weakened an otherwise strong relationship between barnacle density and hummock development: hummocking increased with barnacle density in the absence of crabs but remained relatively high when crabs were present. In communities with similar final barnacle densities, hummocking was more common in those with crabs than those without crabs. The extent to which predators can drive trophic cascades by suppressing the foraging rates of their prey is highly context‐dependent: the positive trait‐mediated indirect effect of predators on basal resource abundance is stronger when many prey respond simultaneously to the threat of predation. However, our results demonstrate that top predators can also enhance competition among basal resources even when their indirect effect on resource abundance is relatively weak. Hence, the cascading effects of predators on competition within lower trophic levels may play an important but under‐appreciated role in the dynamics of basal resource populations and the communities they support. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

In simple, linear food chains, top predators can have positive indirect effects on basal resources by causing changes in the traits (e.g. behaviour, feeding rates) of intermediate consumers. Although less is known about trait-mediated indirect interactions (TMIIs) in more complex food webs, it has been suggested that such complexity dampens trophic cascades. We examined TMIIs between a predatory crab (Carcinus maenas) and two ecologically important basal resources, fucoid algae (Ascophyllum nodosum) and barnacles (Semibalanus balanoides), which are consumed by herbivorous (Littorina littorea) and carnivorous (Nucella lapillus) snails, respectively. Because crab predation risk suppresses snail feeding rates, we hypothesized that crabs would also shape direct and indirect interactions among the multiple consumers and resources. We found that the magnitude of TMIIs between the crab and each resource depended on the suite of intermediate consumers present in the food web. Carnivorous snails (Nucella) transmitted TMIIs between crabs and barnacles. However, crab-algae TMIIs were transmitted by both herbivorous (Littorina) and carnivorous (Nucella) snails, and these TMIIs were additive. By causing Nucella to consume fewer barnacles, crab predation risk allowed fucoids that had settled on or between barnacles to remain in the community. Hence, positive interactions between barnacles and algae caused crab-algae TMIIs to be strongest when both consumers were present. Studies of TMIIs in more realistic, reticulate food webs will be necessary for a more complete understanding of how predation risk shapes community dynamics. [ABSTRACT FROM AUTHOR]

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, Seinibalanus 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 inaenas) 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. [ABSTRACT FROM AUTHOR]

Food chain length is an important property of ecosystems, but the mechanisms maintaining it remain elusive. Classical views suggest that energetic inefficiencies (the ‘energy-flow hypothesis’) limit food chain length, but others have argued that better explanations reside in more complex scenarios that consider the stability of food webs or the combined effects of productivity and ecosystem size. We argue that abandonment of the energy-flow hypothesis is premature. For a simple tritrophic rocky intertidal food chain, we show that the efficiency of energy transfer is strongly influenced by predation risk and consumer density. Effects tied to predation risk were particularly strong, explaining 32% of the variation in growth efficiency (compared to 15% for density effects) and reducing it by 44–76%. Hence, the stress (fear of being eaten) that predators impose on prey may be instrumental in limiting energy transfer up the food chain and thus contribute to the shortening of food chains. [ABSTRACT FROM AUTHOR]

Classical views of trophic cascades emphasize the primacy of consumptive predator effects on prey populations to the transmission of indirect effects [density-mediated indirect interactions (DMIIs)]. However, trophic cascades can also emerge without changes in the density of interacting species because of non-consumptive predator effects on prey traits such as foraging behaviour [trait-mediated indirect interactions (TMIIs)]. Although ecologists appreciate this point, measurements of the relative importance of each indirect predator effect are rare. Experiments with a three-level, rocky shore food chain containing an invasive predatory crab ( Carcinus maenas), an intermediate consumer (the snail, Nucella lapillus) and a basal resource (the barnacle, Semibalanus balanoides) revealed that the strength of TMIIs is comparable with, or exceeds, that of DMIIs. Moreover, the sign and strength of each indirect predator effect depends on whether it is measured in risky or refuge habitats. Because habitat shifts are often responsible for the emergence of TMIIs, attention to the sign and strength of these interactions in both habitats will improve our understanding of the link between individual behaviour and community dynamics. [ABSTRACT FROM AUTHOR]

Studies on the implications of food web interactions to community structure have often focused on density-mediated interactions between predators and their prey. This approach emphasizes the importance of predator regulation of prey density via consumption (i.e. lethal effects), which, in turn, leads to cascading effects on the prey’s resources. A more recent and contrasting view emphasizes the importance of non-lethal predator effects on prey traits (e.g. behaviour, morphology), or trait-mediated interactions. On rocky intertidal shores in New England, green crab (Carcinus maenas ) predation is thought to be important to patterns of algal abundance and diversity by regulating the density of herbivorous snails (Littorina littorea ). We found, however, that risk cues from green crabs can dramatically suppress snail grazing, with large effects on fucoid algal communities. Our results suggest that predator-induced changes in prey behaviour may be an important and under-appreciated component of food web interactions and community dynamics on rocky intertidal shores. [ABSTRACT FROM AUTHOR]