Balancing yield with resilience and conservation objectives in harvested predator–prey communities

The global overexploitation of fish stocks is endangering many marine food webs. Scientists and managers now call for an ecosystem‐based fisheries management, able to take into account the complexity of marine ecosystems and the multiple ecosystem services they provide. By contrast, many fishery management plans only focus on maximizing the productivity of harvested stocks. Such practices are suggested to affect other ecosystem services, altering the integrity and resilience of natural communities. Here we show that while yield‐maximizing policies can allow for coexistence and resilience in predator–prey communities, they are not optimal in a multi‐objective context. We find that although total prey and predator maximum yields are higher with a prey‐oriented harvest, focusing on the predator improves species coexistence. Also, moderate harvesting of the predator can enhance resilience. Furthermore, increasing maximum yields by changing catchabilities improves resilience in predator‐oriented systems, but reduces it in prey‐oriented systems. In a multi‐objective context, optimal harvesting strategies involve a general tradeoff between yield and resilience. Resilience‐maximizing strategies are however compatible with quite high yields, and should often be favored. Our results further suggest that balancing harvest between trophic levels is often best at maintaining simultaneously species coexistence, resilience and yield.     

Stability of complex food webs: resilience, resistance and the average interaction strength

In the face of stochastic climatic perturbations, the overall stability of an ecosystem will be determined by the balance between its resilience and its resistance, but their relative importance is still unknown. Using aquatic food web models we study ecosystem stability as a function of food web complexity. We measured three dynamical stability properties: resilience, resistance, and variability. Specifically, we evaluate how a decrease in the strength of predator-prey interactions with food web complexity, reflecting a decrease in predation efficiency with the number of prey per predator, affects the overall stability of the ecosystem. We find that in mass conservative ecosystems, a lower interaction strength slows down the mass cycling rate in the system and this increases its resistance to perturbations of the growth rate of primary producers. Furthermore, we show that the overall stability of the food webs is mostly given by their resistance, and not by their resilience. Resilience and resistance display opposite trends, although they are shown not to be simply opposite concepts but rather independent properties. The ecological implication is that weaker predator-prey interactions in closed ecosystems can stabilize food web dynamics by increasing its resistance to climatic perturbations.     

Perturbation and abrupt shift in trophic control of biodiversity and productivity

Ecology is founded on the view that ecosystem properties like biodiversity and productivity change smoothly with changing environmental conditions. However, emerging theory predicts that environmental change may cause abrupt shifts to alternate states. In many ecosystems, top predators play a pivotal role in controlling plant productivity and diversity. Yet it remains uncertain if altering this control shifts systems to alternate states. I report on a test of the hypothesis that loss of predator control of ecosystem function causes abrupt state changes in diversity and productivity. In this meadow ecosystem, predators enhance plant diversity by causing a highly productive, competitively dominant plant species to be suppressed by herbivores. Experimental predator removal caused rapid proliferation of the competitively dominant plant. Moreover, temporally staggered predator reintroductions failed to restore the ecosystem. This loss of resilience confirmed that the ecosystem crossed a critical threshold and entrained into an alternate state.     

Do experiments exploring plant diversity–ecosystem functioning relationships inform how biodiversity loss impacts natural ecosystems?

An enormous recent research effort focused on how plant biodiversity (notably species richness) influences ecosystem functioning, usually through experiments in which diversity is varied through random draws of species from a species pool. Such experiments are increasingly used to predict how species losses influence ecosystem functioning in ‘real’ ecosystems. However, this assumes that comparisons of experimental communities with low vs high species richness are analogous to comparisons of natural communities from which species either have or have not been lost. I explore the validity of this assumption, and highlight difficulties in using such experiments to draw conclusions about the ecosystem consequences of biodiversity loss in natural systems. Notably, these experiments do not mimic what happens in real ecosystems either when local extinctions occur or when species losses are offset by gains of new species. Despite limitations, this single experimental approach for studying how biodiversity loss affects ecosystems has often been advocated and implemented at the expense of other approaches; this limits understanding of how natural ecosystems respond to biodiversity loss. I conclude that a broader spectrum of approaches, and more explicit consideration of how species losses and gains operate in concert to influence ecosystems, will help progress this field.     

Diversity in thermal affinity among key piscivores buffers impacts of ocean warming on predator–prey interactions

Asymmetries in responses to climate change have the potential to alter important predator–prey interactions, in part by altering the location and size of spatial refugia for prey. We evaluated the effect of ocean warming on interactions between four important piscivores and four of their prey in the U.S. Northeast Shelf by examining species overlap under historical conditions (1968–2014) and with a doubling in CO₂. Because both predator and prey shift their distributions in response to changing ocean conditions, the net impact of warming or cooling on predator–prey interactions was not determined a priori from the range extent of either predator or prey alone. For Atlantic cod, an historically dominant piscivore in the region, we found that both historical and future warming led to a decline in the proportion of prey species’ range it occupied and caused a potential reduction in its ability to exert top‐down control on these prey. In contrast, the potential for overlap of spiny dogfish with prey species was enhanced by warming, expanding their importance as predators in this system. In sum, the decline in the ecological role for cod that began with overfishing in this ecosystem will likely be exacerbated by warming, but this loss may be counteracted by the rise in dominance of other piscivores with contrasting thermal preferences. Functional diversity in thermal affinity within the piscivore guild may therefore buffer against the impact of warming on marine ecosystems, suggesting a novel mechanism by which diversity confers resilience.     

Predator diversity and abundance provide little support for the enemies hypothesis in forests of high tree diversity

Predatory arthropods can exert strong top-down control on ecosystem functions. However, despite extensive theory and experimental manipulations of predator diversity, our knowledge about relationships between plant and predator diversity--and thus information on the relevance of experimental findings--for species-rich, natural ecosystems is limited. We studied activity abundance and species richness of epigeic spiders in a highly diverse forest ecosystem in subtropical China across 27 forest stands which formed a gradient in tree diversity of 25-69 species per plot. The enemies hypothesis predicts higher predator abundance and diversity, and concomitantly more effective top-down control of food webs, with increasing plant diversity. However, in our study, activity abundance and observed species richness of spiders decreased with increasing tree species richness. There was only a weak, non-significant relationship with tree richness when spider richness was rarefied, i.e. corrected for different total abundances of spiders. Only foraging guild richness (i.e. the diversity of hunting modes) of spiders was positively related to tree species richness. Plant species richness in the herb layer had no significant effects on spiders. Our results thus provide little support for the enemies hypothesis--derived from studies in less diverse ecosystems--of a positive relationship between predator and plant diversity. Our findings for an important group of generalist predators question whether stronger top-down control of food webs can be expected in the more plant diverse stands of our forest ecosystem. Biotic interactions could play important roles in mediating the observed relationships between spider and plant diversity, but further testing is required for a more detailed mechanistic understanding. Our findings have implications for evaluating the way in which theoretical predictions and experimental findings of functional predator effects apply to species-rich forest ecosystems, in which trophic interactions are often considered to be of crucial importance for the maintenance of high plant diversity.     

Diversity enhances community recovery, but not resistance, after drought

1.  There is growing concern that the current loss of biodiversity may negatively affect ecosystem functioning and stability. Although it has been shown that species loss may reduce biomass production and increase temporal variability, experimental evidence that species loss affects ecosystem resistance and resilience after perturbation is limited.
2.  Here, we use the response of experimental plant communities – which differ in diversity – to a natural drought to disentangle the effects of diversity and biomass on resistance, recovery and resilience.
3.  Resistance to drought decreased with diversity, but this pattern was highly dependent upon pre-drought biomass. When corrected for biomass, no relationship between diversity and resistance was observed: at each level of diversity, biomass production was reduced by approximately 30%.
4.  In contrast, recovery (change in biomass production after drought) increased with diversity and was independent of biomass. Resilience (measured as the ratio of post- to pre-drought biomass) was similar at each level of diversity.
5.   Synthesis . On the one hand, our results confirm earlier findings that a positive relationship between diversity and resistance is mainly driven by pre-perturbation performance rather than by diversity. However, the results also show that recovery after drought strongly increased with diversity, independent of performance. We conclude that it is this diversity-dependent recovery which allowed diverse, productive communities to reach the same level of resilience as less diverse (and productive) communities. This finding provides strong experimental evidence for the insurance hypothesis.     

Predator Assemblage Structure and Temporal Variability of Species Richness and Abundance in Forests of High Tree Diversity

Predators significantly affect ecosystem functions, but our understanding of to what extent findings can be transferred from experiments and low‐diversity systems to highly diverse, natural ecosystems is limited. With a particular threat of biodiversity loss at higher trophic levels, however, knowledge of spatial and temporal patterns in predator assemblages and their interrelations with lower trophic levels is essential for assessing effects of trophic interactions and advancing biodiversity conservation in these ecosystems. We analyzed spatial and temporal variability of spider assemblages in tree species‐rich subtropical forests in China, across 27 study plots varying in woody plant diversity and stand age. Despite effects of woody plant richness on spider assemblage structure, neither habitat specificity nor temporal variability of spider richness and abundance were influenced. Rather, variability increased with forest age, probably related to successional changes in spider assemblages. Our results indicate that woody plant richness and theory predicting increasing predator diversity with increasing plant diversity do not necessarily play a major role for spatial and temporal dynamics of predator assemblages in such plant species‐rich forests. Diversity effects on biotic or abiotic habitat conditions might be less pronounced across our gradient from medium to high plant diversity than in previously studied less diverse systems, and bottom‐up effects might level out at high plant diversity. Instead, our study highlights the importance of overall (diversity‐independent) environmental heterogeneity in shaping spider assemblages and, as indicated by a high species turnover between plots, as a crucial factor for biodiversity conservation at a regional scale in these subtropical forests.     

Modelling Disease Introduction as Biological Control of Invasive Predators to Preserve Endangered Prey

Invasive species are a significant cause of bio-diversity loss particularly in island ecosystems. It has been suggested to release pathogenic parasites as an efficient control measure of these mostly immune-naïve populations. In order to explore the potential impacts of such bio-control approach, we construct and investigate mathematical models describing disease dynamics in a host population that acts as a predator embedded in a simple food chain. The consequences of Feline Immunodeficiency Virus (FIV) introduction into a closed ecosystem are addressed using a bi-trophic system, comprising an indigenous prey (birds) and an introduced predator (cats). Our results show that FIV is unlikely to fully eradicate cats on sub-Antarctic islands, but it can be efficient in depressing their population size, allowing for the recovery of the endangered prey. Depending on the ecological setting and disease transmission mode (we consider proportionate mixing as well as mass action), successful pathogen invasion can induce population oscillations that are not possible in the disease-free predator–prey system. These fluctuations can be seen as a mixed blessing from a management point of view. On the one hand, they may increase the extinction risk of the birds. On the other hand, they provide an opportunity to eradicate cats more easily in combination with other methods such as trapping or culling.     

Species richness and trophic diversity increase decomposition in a co-evolved food web

Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.     

Your worst enemy could be your best friend: predator contributions to invasion resistance and persistence of natives

Native predators are postulated to have an important role in biotic resistance of communities to invasion and community resilience. Effects of predators can be complex, and mechanisms by which predators affect invasion success and impact are understood for only a few well-studied communities. We tested experimentally whether a native predator limits an invasive species’ success and impact on a native competitor for a community of aquatic insect larvae in water-filled containers. The native mosquito Aedes triseriatus alone had no significant effect on abundance of the invasive mosquito Aedes albopictus. The native predatory midge Corethrella appendiculata, at low or high density, significantly reduced A. albopictus abundance. This effect was not caused by trait-mediated oviposition avoidance of containers with predators, but instead was a density-mediated effect caused by predator-induced mortality. The presence of this predator significantly reduced survivorship of the native species, but high predator density also significantly increased development rate of the native species when the invader was present, consistent with predator-mediated release from interspecific competition with the invader. Thus, a native predator can indirectly benefit its native prey when a superior competitor invades. This shows the importance of native predators as a component of biodiversity for both biotic resistance to invasion and resilience of a community perturbed by successful invasion.     

Context-dependent effects of predator removal from experimental microcosm communities

The loss of a predator from an ecological community can cause large changes in community structure and ecosystem processes, or have very little consequence for the remaining species and ecosystem. Understanding when and why the loss of a predator causes large changes in community structure and ecosystem processes is critical for understanding the functional consequences of biodiversity loss. We used experimental microbial communities to investigate how the removal of a large generalist predator affected the extinction frequency, population abundance and total biomass of its prey. We removed this predator in the presence or absence of an alternative, more specialist, predator in order to determine whether the specialist predator affected the outcome of the initial species removal. Removal of the large generalist predator altered some species' populations but many were unaffected and no secondary extinctions were observed. The specialist predator, though rare, altered the response of the prey community to the removal of the large generalist predator. In the absence of the specialist predator, the effects of the removal were only measurable at the level of individual species. However, when the specialist predator was present, the removal of the large generalist predator affected the total biomass of prey species. The results demonstrate that the effect of species loss from high trophic levels may be very context-dependent, as rare species can have disproportionately large effects in food webs.     

Loss of Rare Fish Species from Tropical Floodplain Food Webs Affects Community Structure and Ecosystem Multifunctionality in a Mesocosm Experiment

Experiments with realistic scenarios of species loss from multitrophic ecosystems may improve insight into how biodiversity affects ecosystem functioning. Using 1000 L mesocoms, we examined effects of nonrandom species loss on community structure and ecosystem functioning of experimental food webs based on multitrophic tropical floodplain lagoon ecosystems. Realistic biodiversity scenarios were developed based on long-term field surveys, and experimental assemblages replicated sequential loss of rare species which occurred across all trophic levels of these complex food webs. Response variables represented multiple components of ecosystem functioning, including nutrient cycling, primary and secondary production, organic matter accumulation and whole ecosystem metabolism. Species richness significantly affected ecosystem function, even after statistically controlling for potentially confounding factors such as total biomass and direct trophic interactions. Overall, loss of rare species was generally associated with lower nutrient concentrations, phytoplankton and zooplankton densities, and whole ecosystem metabolism when compared with more diverse assemblages. This pattern was also observed for overall ecosystem multifunctionality, a combined metric representing the ability of an ecosystem to simultaneously maintain multiple functions. One key exception was attributed to time-dependent effects of intraguild predation, which initially increased values for most ecosystem response variables, but resulted in decreases over time likely due to reduced nutrient remineralization by surviving predators. At the same time, loss of species did not result in strong trophic cascades, possibly a result of compensation and complexity of these multitrophic ecosystems along with a dominance of bottom-up effects. Our results indicate that although rare species may comprise minor components of communities, their loss can have profound ecosystem consequences across multiple trophic levels due to a combination of direct and indirect effects in diverse multitrophic ecosystems.     

Species richness facilitates ecosystem resilience in aquatic food webs

1. Many studies indicate that biodiversity in ecosystems affects stability, either by promoting temporal stability of ecosystem attributes or by enhancing ecosystem resistance and resilience to perturbation. The effects on temporal stability are reasonably well understood and documented but effects on resistance and resilience are not. 2. Here, we report results from an aquatic mesocosm experiment in which we manipulated the species richness and composition of aquatic food webs (macrophytes, macro‐herbivores and invertebrate predators), imposed a pulse disturbance (acidification), and monitored the resistance (initial response) and resilience (recovery) of ecosystem productivity and respiration. 3. We found that species‐rich macroinvertebrate communities had higher resilience of whole‐ecosystem respiration, but were not more resistant to perturbations. We also found that resilience and resistance were unaffected by species composition, despite the strong role composition is known to play in determining mean levels of function in these communities. 4. Biodiversity’s effects on resilience were probably mediated through complex pathways affecting phytoplankton and microbial communities (e.g. via changes in nutrient regeneration, grazing or compositional changes) rather than through simpler effects (e.g. insurance effects, enhanced facilitation) although these simpler mechanisms probably played minor roles in enhancing respiration resilience. 5. Current mechanisms for understanding biodiversity’s effects on ecosystem stability have been developed primarily in the context of single‐trophic level communities. These mechanisms may be overly simplistic for understanding the consequences of species richness on ecosystem stability in complex, multi‐trophic food webs where additional factors such as indirect effects and highly variable life‐history traits of species may also be important.     

Intraspecific diversity loss in a predator species alters prey community structure and ecosystem functions

Loss in intraspecific diversity can alter ecosystem functions, but the underlying mechanisms are still elusive, and intraspecific biodiversity–ecosystem function (iBEF) relationships have been restrained to primary producers. Here, we manipulated genetic and functional richness of a fish consumer (Phoxinus phoxinus) to test whether iBEF relationships exist in consumer species and whether they are more likely sustained by genetic or functional richness. We found that both genotypic and functional richness affected ecosystem functioning, either independently or interactively. Loss in genotypic richness reduced benthic invertebrate diversity consistently across functional richness treatments, whereas it reduced zooplankton diversity only when functional richness was high. Finally, losses in genotypic and functional richness altered functions (decomposition) through trophic cascades. We concluded that iBEF relationships lead to substantial top-down effects on entire food chains. The loss of genotypic richness impacted ecological properties as much as the loss of functional richness, probably because it sustains “cryptic” functional diversity.

Global change is expected to generate a loss of intraspecific diversity worldwide. This mesocosm study explores whether loss of genetic and functional diversity in a predator species affects community and ecosystem functioning of lower trophic levels in pond ecosystems, revealing that diversity loss in a single consumer species can impact an entire ecosystem, reducing its functionality.     

A cross‐system meta‐analysis reveals coupled predation effects on prey biomass and diversity

Predator diversity and abundance are under strong human pressure in all types of ecosystems. Whereas predator potentially control standing biomass and species interactions in food webs, their effects on prey biomass and especially prey biodiversity have not yet been systematically quantified. Here, we test the effects of predation in a cross‐system meta‐analysis of prey diversity and biomass responses to local manipulation of predator presence. We found 291 predator removal experiments from 87 studies assessing both diversity and biomass responses. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey across ecosystems. Predation effects were highly similar between ecosystem types, whereas previous studies had shown that herbivory or decomposition effects differed fundamentally between terrestrial and aquatic systems based on different stoichiometry of plant material. Such stoichiometric differences between systems are unlikely for carnivorous predators, where effect sizes on species richness strongly correlated to effect sizes on biomass. However, the negative predation effect on prey biomass was ameliorated significantly with increasing prey richness and increasing species richness of the manipulated predator assemblage. Moreover, with increasing richness of the predator assemblage present, the overall negative effects of predation on prey richness switched to positive effects. Our meta‐analysis revealed strong general relationships between predator diversity, prey diversity and the interaction strength between trophic levels in terms of biomass. This study indicates that anthropogenic changes in predator abundance and diversity will potentially have strong effects on trophic interactions across ecosystems. Synthesis The past centuries we have experienced a dramatic loss of top–predator abundance and diversity in most types of ecosystems. To understand the direct consequences of predator loss on a global scale, we quantitatively summarized experiments testing predation effects on prey communities in a cross‐system meta‐analysis. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey, and predation effects were highly similar. However, with increasing predator richness, the overall negative effects of predation on prey richness switched to positive ones. Anthropogenic changes in predator communities will potentially have strong effects on prey diversity, biomass, and trophic interactions across ecosystems.     

Effects of perturbation on the predator–prey system in a heterogeneous landscape

Environmental perturbations occur in ecosystems as the result of disturbance, which is closely related to ecosystem stability and resilience. To understand how perturbations can affect ecosystems, we constructed a spatially explicit lattice model to simulate the integrative predator–prey–grass relationships. In this model, a predator (or prey) gives birth to offspring, according to a specific birth probability, when it is able to feed on prey (or grass). When a predator or prey animal was initially introduced or newly born, its health state was set at a given high value. This state decreased by 1 with each time step. When the state of an animal decreased to zero, the animal was considered dead and was removed from the system. In this model, the perturbation was defined as the sudden death of some portion of the population. The heterogeneous landscape was characterized by a parameter, H, which controlled the degree of heterogeneity. When H ≥ 0.6, the predator population size was positively influenced by the perturbation. However, the perturbation had little effect upon the population sizes of prey or grass, regardless of the value of H.     

Resilience to Stress and Disturbance,and Resistance to Bromus tectorum L. Invasion in Cold Desert Shrublands of Western North America

Alien grass invasions in arid and semi-arid ecosystems are resulting in grass–fire cycles and ecosystem-level transformations that severely diminish ecosystem services. Our capacity to address the rapid and complex changes occurring in these ecosystems can be enhanced by developing an understanding of the environmental factors and ecosystem attributes that determine resilience of native ecosystems to stress and disturbance, and resistance to invasion. Cold desert shrublands occur over strong environmental gradients and exhibit significant differences in resilience and resistance. They provide an excellent opportunity to increase our understanding of these concepts. Herein, we examine a series of linked questions about (a) ecosystem attributes that determine resilience and resistance along environmental gradients, (b) effects of disturbances like livestock grazing and altered fire regimes and of stressors like rapid climate change, rising CO₂, and N deposition on resilience and resistance, and (c) interacting effects of resilience and resistance on ecosystems with different environmental conditions. We conclude by providing strategies for the use of resilience and resistance concepts in a management context. At ecological site scales, state and transition models are used to illustrate how differences in resilience and resistance influence potential alternative vegetation states, transitions among states, and thresholds. At landscape scales management strategies based on resilience and resistance—protection, prevention, restoration, and monitoring and adaptive management—are used to determine priority management areas and appropriate actions.     

Phenotypic differentiation within a foundation grass species correlates with species richness in a subalpine community

The effects of species loss on ecosystems depend on the community’s functional diversity (FD). However, how FD responds to environmental changes is poorly understood. This applies particularly to higher trophic levels, which regulate many ecosystem processes and are strongly affected by human-induced environmental changes. We analyzed how functional richness (FRic), evenness (FEve), and divergence (FDiv) of important generalist predators—epigeic spiders—are affected by changes in woody plant species richness, plant phylogenetic diversity, and stand age in highly diverse subtropical forests in China. FEve and FDiv of spiders increased with plant richness and stand age. FRic remained on a constant level despite decreasing spider species richness with increasing plant species richness. Plant phylogenetic diversity had no consistent effect on spider FD. The results contrast with the negative effect of diversity on spider species richness and suggest that functional redundancy among spiders decreased with increasing plant richness through non-random species loss. Moreover, increasing functional dissimilarity within spider assemblages with increasing plant richness indicates that the abundance distribution of predators in functional trait space affects ecological functions independent of predator species richness or the available trait space. While plant diversity is generally hypothesized to positively affect predators, our results only support this hypothesis for FD—and here particularly for trait distributions within the overall functional trait space—and not for patterns in species richness. Understanding the way predator assemblages affect ecosystem functions in such highly diverse, natural ecosystems thus requires explicit consideration of FD and its relationship with species richness.     

Not worth the risk: apex predators suppress herbivory on coral reefs

Apex predators are known to exert strong ecological effects, either through direct or indirect predator–prey interactions. Indirect interactions have the potential to influence ecological communities more than direct interactions as the effects are propagated throughout the population as opposed to only one individual. Indirect effects of apex predators are well documented in terrestrial environments, however there is a paucity of information for marine environments. Furthermore, manipulative studies, as opposed to correlative observations, isolating apex predator effects are lacking. Coral reefs are one of the most diverse ecosystems, providing a useful model system for investigating the ecological role of apex predators and their influence on lower trophic levels. Using predator models and transplanted macroalgae we examined the indirect effects of predators on herbivore foraging behaviour. We show that the presence of a model reef shark or large coral‐grouper led to a substantial reduction in bite rate and species richness of herbivorous fishes and an almost absolute localized cessation of macroagal removal, due to the perceived risk of predation. A smaller‐sized coral‐grouper also reduced herbivore diversity and activity but to a lesser degree than the larger model predators. These indirect effects of apex predators on the foraging behaviour of herbivores may have flow‐on effects on the biomass and distribution of macroalgae, and the functioning of coral reef ecosystems. This highlights that the ecological interactions and processes that contribute to ecosystem resilience may be more complex than previously assumed.