In complex food webs, interactions among species in different trophic levels can generate cascading indirect effects that couple top predators with primary producers, thereby affecting ecosystem functioning. Natural selection imposed by top predators on intermediate predators may play a role in shaping the strength of these trophic cascades, but this conjecture remains largely untested. To determine the effects of natural selection on the strength of trophic cascades, we conducted a two‐part experiment in a four‐level aquatic trophic system involving a top predator (fish), an intermediate predator (damselflies), herbivores (zooplankton) and primary producers (algae). We first quantified how predation by fish generated selection on damselfly activity levels after controlling for phenotypic plasticity. We then measured the indirect effects of this selection on primary production (phytoplankton biomass). In both experiments, we varied the density of predators, allowing us to elucidate both trait‐mediated and density‐mediated indirect effects. We found that as fish density increased, damselfly survivorship declined, which generated natural selection favouring less active damselflies. These results are robust after taking into account latent effects of plasticity in response to fish predator cues. The surviving damselflies likely foraged less, freeing herbivores from predation, which in turn reduced primary production. This selection‐driven trait‐mediated indirect effect was only apparent at low damselfly densities, because the consumptive effect of damselflies at high densities overwhelmed the effects of past selection. These results demonstrate how the past action of natural selection imposed by predators affects the phenotypes of prey that consume herbivores, which ultimately influences primary productivity—a selective trophic cascade. Natural selection can therefore act as a mechanism coupling ecological dynamics across trophic levels, which ultimately influences ecosystem functioning. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13102/suppinfo is available for this article.
Predators have a key role shaping competitor dynamics in food webs. Perhaps the most obvious way this occurs is when predators reduce competitor densities. However, consumption could also generate phenotypic selection on prey that determines the strength of competition, thus coupling consumptive and trait-based effects of predators. In a mesocosm experiment simulating fish predation on damselflies, we found that selection against high damselfly activity ratesa phenotype mediating predation and competitionweakened the strength of density dependence in damselfly growth rates. A field experiment corroborated this finding and showed that increasing damselfly densities in lakes with high fish densities had limited effects on damselfly growth rates but generated a precipitous growth rate decline where fish densities were lowera pattern expected because of spatial variation in selection imposed by predation. These results suggest that accounting for both consumption and selection is necessary to determine how predators regulate prey competitive interactions.
Despite the ubiquitous nature of parasitism, how parasitism alters the outcome of host–species interactions such as competition, mutualism and predation remains unknown. Using a phylogenetically informed meta‐analysis of 154 studies, we examined how the mean and variance in the outcomes of species interactions differed between parasitized and non‐parasitized hosts. Overall, parasitism did not significantly affect the mean or variance of host–species interaction outcomes, nor did the shared evolutionary histories of hosts and parasites have an effect. Instead, there was considerable variation in outcomes, ranging from strongly detrimental to strongly beneficial for infected hosts. Trophically‐transmitted parasites increased the negative effects of predation, parasites increased and decreased the negative effects of interspecific competition for parasitized and non‐parasitized heterospecifics, respectively, and parasites had particularly strong negative effects on host species interactions in freshwater and marine habitats, yet were beneficial in terrestrial environments. Our results illuminate the diverse ways in which parasites modify critical linkages in ecological networks, implying that whether the cumulative effects of parasitism are considered detrimental depends not only on the interactions between hosts and their parasites but also on the many other interactions that hosts experience.
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