Coral guard crabs.

Adrian Stier and Craig Osenberg introduce Trapeziid crabs, which live in close symbiosis with corals.

Variation in body size drives spatial and temporal variation in lobster-urchin interaction strength.

How strongly predators and prey interact is both notoriously context dependent and difficult to measure. Yet across taxa, interaction strength is strongly related to predator size, prey size and prey density, suggesting that general cross-taxonomic relationships could be used to predict how strongly individual species interact. Here, we ask how accurately do general size-scaling relationships predict variation in interaction strength between specific species that vary in size and density across space and time? To address this question, we quantified the size and density dependence of the functional response of the California spiny lobster Panulirus interruptus, foraging on a key ecosystem engineer, the purple sea urchin Strongylocentrotus purpuratus, in experimental mesocosms. Based on these results, we then estimated variation in lobster-urchin interaction strength across five sites and 9 years of observational data. Finally, we compared our experimental estimates to predictions based on general size-scaling relationships from the literature. Our results reveal that predator and prey body size has the greatest effect on interaction strength when prey abundance is high. Due to consistently high urchin densities in the field, our simulations suggest that body size-relative to density-accounted for up to 87% of the spatio-temporal variation in interaction strength. However, general size-scaling relationships failed to predict the magnitude of interactions between lobster and urchin; even the best prediction from the literature was, on average, an order of magnitude (+18.7×) different than our experimental predictions. Harvest and climate change are driving reductions in the average body size of many marine species. Anticipating how reductions in body size will alter species interactions is critical to managing marine systems in an ecosystem context. Our results highlight the extent to which differences in size-frequency distributions can drive dramatic variation in the strength of interactions across narrow spatial and temporal scales. Furthermore, our work suggests that species-specific estimates for the scaling of interaction strength with body size, rather than general size-scaling relationships, are necessary to quantitatively predict how reductions in body size will alter interaction strengths.

Material legacies can degrade resilience: Structure-retaining disturbances promote regime shifts on coral reefs.

Standing dead structures of habitat-forming organisms (e.g., dead trees, coral skeletons, oyster shells) killed by a disturbance are material legacies that can affect ecosystem recovery processes. Many ecosystems are subject to different types of disturbance that either remove biogenic structures or leave them intact. Here we used a mathematical model to quantify how the resilience of coral reef ecosystems may be differentially affected following structure-removing and structure-retaining disturbance events, focusing in particular on the potential for regime shifts from coral to macroalgae. We found that dead coral skeletons could substantially diminish coral resilience if they provided macroalgae refuge from herbivory, a key feedback associated with the recovery of coral populations. Our model shows that the material legacy of dead skeletons broadens the range of herbivore biomass over which coral and macroalgae states are bistable. Hence, material legacies can alter resilience by modifying the underlying relationship between a system driver (herbivory) and a state variable (coral cover).

Glimmers of hope in large carnivore recoveries.

In the face of an accelerating extinction crisis, scientists must draw insights from successful conservation interventions to uncover promising strategies for reversing broader declines. Here, we synthesize cases of recovery from a list of 362 species of large carnivores, ecologically important species that function as terminal consumers in many ecological contexts. Large carnivores represent critical conservation targets that have experienced historical declines as a result of direct exploitation and habitat loss. We examine taxonomic and geographic variation in current extinction risk and recovery indices, identify conservation actions associated with positive outcomes, and reveal anthropogenic threats linked to ongoing declines. We find that fewer than 10% of global large carnivore populations are increasing, and only 12 species (3.3%) have experienced genuine improvement in extinction risk, mostly limited to recoveries among marine mammals. Recovery is associated with species legislation enacted at national and international levels, and with management of direct exploitation. Conversely, ongoing declines are robustly linked to threats that include habitat modification and human conflict. Applying lessons from cases of large carnivore recovery will be crucial for restoring intact ecosystems and maintaining the services they provide to humans.

Avoiding critical thresholds through effective monitoring.

A major challenge in sustainability science is identifying targets that maximize ecosystem benefits to humanity while minimizing the risk of crossing critical system thresholds. One critical threshold is the biomass at which populations become so depleted that their population growth rates become negative-depensation. Here, we evaluate how the value of monitoring information increases as a natural resource spends more time near the critical threshold. This benefit emerges because higher monitoring precision promotes higher yield and a greater capacity to recover from overharvest. We show that precautionary buffers that trigger increased monitoring precision as resource levels decline may offer a way to minimize monitoring costs and maximize profits. In a world of finite resources, improving our understanding of the trade-off between precision in estimates of population status and the costs of mismanagement will benefit stakeholders that shoulder the burden of these economic and social costs.

Fertilization by coral-dwelling fish promotes coral growth but can exacerbate bleaching response.

Many corals form close associations with a diverse assortment of coral-dwelling fishes and other fauna. As coral reefs around the world are increasingly threatened by mass bleaching events, it is important to understand how these biotic interactions influence corals' susceptibility to bleaching. We used dynamic energy budget modeling to explore how nitrogen excreted by coral-dwelling fish affects the physiological performance of host corals. In our model, fish presence influenced the functioning of the coral-Symbiodiniaceae symbiosis by altering nitrogen availability, and the magnitude and sign of these effects depended on environmental conditions. Although our model predicted that fish-derived nitrogen can promote coral growth, the relationship between fish presence and coral tolerance of photo-oxidative stress was non-linear. Fish excretions supported denser symbiont populations that provided protection from incident light through self-shading. However, these symbionts also used more of their photosynthetic products for their own growth, rather than sharing with the coral host, putting the coral holobiont at a higher risk of becoming carbon-limited and bleaching. The balance between the benefits of increased symbiont shading and costs of reduced carbon sharing depended on environmental conditions. Thus, while there were some scenarios under which fish presence increased corals' tolerance of light stress, fish could also exacerbate bleaching and slow or prevent subsequent recovery. We discuss how the contrast between the potentially harmful effects of fish predicted by our model and results of empirical studies may relate to key model assumptions that warrant further investigation. Overall, this study provides a foundation for future work on how coral-associated fauna influence the bioenergetics of their host corals, which in turn has implications for how these corals respond to bleaching-inducing stressors.

Detrital supply suppresses deforestation to maintain healthy kelp forest ecosystems.

Herbivores can reach extraordinary abundances in many ecosystems. When herbivore abundance is high, heavy grazing can severely defoliate primary producers and, in some cases, even drive ecosystem to undergo regime shifts from a high productivity state to a denuded, low productivity state. While the phenomenon of herbivore-driven regime shifts is well documented, we only partially understand the mechanisms underlying these events. Here, we combine herbivory experiments with 21 years of long-term monitoring data of kelp forest ecosystems to test the hypothesis that herbivores drive regime shifts when herbivory exceeds primary production. To test this hypothesis, we quantified how the foraging habits of an important group of marine herbivores-sea urchins-change with increases in sea urchin biomass and trigger regime shifts to a foundation species, giant kelp (Macrocystis pyrifera). Using experiments, we quantified how the grazing capacity of urchins increases as urchin biomass increases, then we combined these estimates of urchin grazing capacity with estimates of kelp production to predict when and where urchin grazing capacity exceeded kelp production. When grazing capacity exceeded kelp production, sea urchins caused a 50-fold reduction in giant kelp biomass. Our findings support the hypothesis that the balance between herbivory and production underlies herbivore-driven regime shifts in Southern California kelp forests and provides insight into when and where urchins are likely to force regime shifts in kelp forest ecosystems.

Remoteness does not enhance coral reef resilience.

Remote coral reefs are thought to be more resilient to climate change due to their isolation from local stressors like fishing and pollution. We tested this hypothesis by measuring the relationship between local human influence and coral community resilience. Surprisingly, we found no relationship between human influence and resistance to disturbance and some evidence that areas with greater human development may recover from disturbance faster than their more isolated counterparts. Our results suggest remote coral reefs are imperiled by climate change, like so many other geographically isolated ecosystems, and are unlikely to serve as effective biodiversity arks. Only drastic and rapid cuts in greenhouse gas emissions will ensure coral survival. Our results also indicate that some reefs close to large human populations were relatively resilient. Focusing research and conservation resources on these more accessible locations has the potential to provide new insights and maximize conservation outcomes.

Variation in disturbance to a foundation species structures the dynamics of a benthic reef community.

Disturbance and foundation species can both have strong impacts on ecosystem structure and function, but studies of their interacting effects are hindered by the long life spans and slow growth of most foundation species. Here, we investigated the extent to which foundation species may mediate the impacts of disturbance on ecological communities, using the kelp forest ecosystem as a study system. Giant kelp (Macrocystis pyrifera) grows rapidly and experiences wave disturbance from winter storms. We developed and analyzed a model of the effects of variable storm regimes on giant kelp population dynamics and of the cascading effects on kelp-mediated competition between benthic community members in kelp forests. Simulations of severe storm regimes resulted in a greater abundance of understory macroalgae and a lower abundance of sessile invertebrates than did milder regimes. Both the cascading effects of periodic loss of giant kelp as well as the degree to which storms directly impacted the benthos (in the form of scouring) influenced the outcome of competition between benthic community members. The model's qualitative predictions were consistent with empirical data from a 20-yr time series of community dynamics, suggesting that interannual variability in disturbance that affects giant kelp abundance can have strong consequences for benthic community structure. Our findings point to the value of long-term studies in elucidating the interacting effects of disturbance and foundation species.

Evidence that spillover from Marine Protected Areas benefits the spiny lobster (Panulirus interruptus) fishery in southern California.

Marine Protected Areas (MPAs) are designed to enhance biodiversity and ecosystem services. Some MPAs are also established to benefit fisheries through increased egg and larval production, or the spillover of mobile juveniles and adults. Whether spillover influences fishery landings depend on the population status and movement patterns of target species both inside and outside of MPAs, as well as the status of the fishery and behavior of the fleet. We tested whether an increase in the lobster population inside two newly established MPAs influenced local catch, fishing effort, and catch-per-unit-effort (CPUE) within the sustainable California spiny lobster fishery. We found greater build-up of lobsters within MPAs relative to unprotected areas, and greater increases in fishing effort and total lobster catch, but not CPUE, in fishing zones containing MPAs vs. those without MPAs. Our results show that a 35% reduction in fishing area resulting from MPA designation was compensated for by a 225% increase in total catch after 6-years, thus indicating at a local scale that the trade-off of fishing ground for no-fishing zones benefitted the fishery.

Ecological impacts of human-induced animal behaviour change.

A growing body of literature has documented myriad effects of human activities on animal behaviour, yet the ultimate ecological consequences of these behavioural shifts remain largely uninvestigated. While it is understood that, in the absence of humans, variation in animal behaviour can have cascading effects on species interactions, community structure and ecosystem function, we know little about whether the type or magnitude of human-induced behavioural shifts translate into detectable ecological change. Here we synthesise empirical literature and theory to create a novel framework for examining the range of behaviourally mediated pathways through which human activities may affect different ecosystem functions. We highlight the few empirical studies that show the potential realisation of some of these pathways, but also identify numerous factors that can dampen or prevent ultimate ecosystem consequences. Without a deeper understanding of these pathways, we risk wasting valuable resources on mitigating behavioural effects with little ecological relevance, or conversely mismanaging situations in which behavioural effects do drive ecosystem change. The framework presented here can be used to anticipate the nature and likelihood of ecological outcomes and prioritise management among widespread human-induced behavioural shifts, while also suggesting key priorities for future research linking humans, animal behaviour and ecology.

Foundation species promote community stability by increasing diversity in a giant kelp forest.

Foundation species structure communities, promote biodiversity, and stabilize ecosystem processes by creating locally stable environmental conditions. Despite their critical importance, the role of foundation species in stabilizing natural communities has seldom been quantified. In theory, the stability of a foundation species should promote community stability by enhancing species richness, altering the population fluctuations of individual species, or both. Here we tested the hypothesis that the stability of a marine foundation species, the giant kelp Macrocystis pyrifera, increased the stability of the aggregate biomass of a phylogenetically diverse assemblage of understory algae and sessile invertebrates that compete for space beneath the giant kelp canopy. To achieve this goal, we analyzed an 18-yr time series of the biomass of giant kelp and its associated benthic community collected from 32 plots distributed among nine shallow reefs in the Santa Barbara Channel, USA. We showed that the stability of understory algae and sessile invertebrates was positively and indirectly related to the stability of giant kelp, which primarily resulted from giant kelp's direct positive association with species richness. The stability of all community types was positively related to species richness via increased species stability and species asynchrony. The stabilizing effects of richness were three to four times stronger when algae and invertebrates were considered separately rather than in combination. Our finding that diversity-stability relationships were stronger in communities consisting of species with similar resource requirements suggests that competition for shared resources rather than differential responses to environmental conditions played a more important role in stabilizing the community. Increasing threats to structure-forming foundation species worldwide necessitates a detailed understanding of how they influence their associated community. This study is among the first to show that dampened temporal fluctuations in the biomass of a foundation species is an important determinant of the stability of the complex communities it supports.

Spatial variation in exploited metapopulations obscures risk of collapse.

Unanticipated declines among exploited species have commonly occurred despite harvests that appeared sustainable prior to collapse. This is particularly true in the oceans where spatial scales of management are often mismatched with spatially complex metapopulations. We explore causes, consequences, and potential solutions for spatial mismatches in harvested metapopulations in three ways. First, we generate novel theory illustrating when and how harvesting metapopulations increases spatial variability and in turn masks local-scale volatility. Second, we illustrate why spatial variability in harvested metapopulations leads to negative consequences using an empirical example of a Pacific herring metapopulation. Finally, we construct a numerical management strategy evaluation model to identify and highlight potential solutions for mismatches in spatial scale and spatial variability. Our results highlight that spatial complexity can promote stability at large scales, however, ignoring spatial complexity produces cryptic and negative consequences for people and animals that interact with resources at small scales. Harvesting metapopulations magnifies spatial variability, which creates discrepancies between regional and local trends while increasing risk of local population collapses. Such effects asymmetrically impact locally constrained fishers and predators, which are more exposed to risks of localized collapses. Importantly, we show that dynamically optimizing harvest can minimize local risk without sacrificing yield. Thus, multiple nested scales of management may be necessary to avoid cryptic collapses in metapopulations and the ensuing ecological, social, and economic consequences.

Strong Evidence for an Intraspecific Metabolic Scaling Coefficient Near 0.89 in Fish.

As an example of applying the evidential approach to statistical inference, we address one of the longest standing controversies in ecology, the evidence for, or against, a universal metabolic scaling relationship between metabolic rate and body mass. Using fish as our study taxa, we curated 25 studies with measurements of standard metabolic rate, temperature, and mass, with 55 independent trials and across 16 fish species and confronted this data with flexible random effects models. To quantify the body mass - metabolic rate relationship, we perform model selection using the Schwarz Information Criteria (ΔSIC), an established evidence function. Further, we formulate and justify the use of ΔSIC intervals to delineate the values of the metabolic scaling relationship that should be retained for further consideration. We found strong evidence for a metabolic scaling coefficient of 0.89 with a ΔSIC interval spanning 0.82 to 0.99, implying that mechanistically derived coefficients of 0.67, 0.75, and 1, are not supported by the data. Model selection supports the use of a random intercepts and random slopes by species, consistent with the idea that other factors, such as taxonomy or ecological or lifestyle characteristics, may be critical for discerning the underlying process giving rise to the data. The evidentialist framework applied here, allows for further refinement given additional data and more complex models.

Ocean recoveries for tomorrow's Earth: Hitting a moving target.

Growing scientific awareness, strong regulations, and effective management have begun to fulfill the promise of recovery in the ocean. However, many efforts toward ocean recovery remain unsuccessful, in part because marine ecosystems and the human societies that depend upon them are constantly changing. Furthermore, recovery efforts are embedded in marine social-ecological systems where large-scale dynamics can inhibit recovery. We argue that the ways forward are to (i) rethink an inclusive definition of recovery that embraces a diversity of stakeholder perspectives about acceptable recovery goals and ecosystem outcomes; (ii) encourage research that enables anticipation of feasible recovery states and identifies pathways toward resilient ecosystems; and (iii) adopt policies that are sufficiently nimble to keep pace with rapid change and governance that works seamlessly from local to regional scales. Application of these principles can facilitate successful recoveries in a world where environmental conditions and social imperatives are constantly shifting.

Predator-induced selection on urchin activity level depends on urchin body size.

Temporally consistent individual differences in behavior impact many ecological processes. We simultaneously examined the effects of individual variation in prey activity level, covering behavior, and body size on prey survival with predators using an urchin-lobster system. Specifically, we tested the hypothesis that slow-moving purple sea urchins (Strongylocentrotus purpuratus) and urchins who deploy extensive substrate (pebbles and stones) covering behavior will out-survive active urchins that deploy little to no covering behavior when pitted against a predator, the California spiny lobster (Panulirus interruptus). We evaluated this hypothesis by first confirming whether individual urchins exhibit temporally consistent differences in activity level and covering behavior, which they did. Next, we placed groups of four urchins in mesocosms with single lobster and monitored urchin survival for 108 hours. High activity level was negatively associated with survival, whereas urchin size and covering behavior independently did not influence survival. The negative effect of urchin activity level on urchin survival was strong for smaller urchins and weaker for large urchins. Taken together, these results suggest that purple urchin activity level and size jointly determine their susceptibility to predation by lobsters. This is potentially of great interest, because predation by recovering lobster populations could alter the stability of kelp forests by culling specific phenotypes, like foraging phenotypes, from urchin populations.

Rapid and direct recoveries of predators and prey through synchronized ecosystem management.

One of the twenty-first century's greatest environmental challenges is to recover and restore species, habitats and ecosystems. The decision about how to initiate restoration is best-informed by an understanding of the linkages between ecosystem components and, given these linkages, an appreciation of the consequences of choosing to recover one ecosystem component before another. However, it remains difficult to predict how the sequence of species' recoveries within food webs influences the speed and trajectory of restoration, and what that means for human well-being. Here, we develop theory to consider the ecological and social implications of synchronous versus sequential (species-by-species) recovery in the context of exploited food webs. A dynamical systems model demonstrates that synchronous recovery of predators and prey is almost always more efficient than sequential recovery. Compared with sequential recovery, synchronous recovery can be twice as fast and produce transient fluctuations of much lower amplitude. A predator-first strategy is particularly slow because it counterproductively suppresses prey recovery. An analysis of real-world predator-prey recoveries shows that synchronous and sequential recoveries are similarly common, suggesting that current practices are not ideal. We highlight policy tools that can facilitate swift and steady recovery of ecosystem structure, function and associated services.

Ecosystem context and historical contingency in apex predator recoveries.

Habitat loss, overexploitation, and numerous other stressors have caused global declines in apex predators. This "trophic downgrading" has generated widespread concern because of the fundamental role that apex predators can play in ecosystem functioning, disease regulation, and biodiversity maintenance. In attempts to combat declines, managers have conducted reintroductions, imposed stricter harvest regulations, and implemented protected areas. We suggest that full recovery of viable apex predator populations is currently the exception rather than the rule. We argue that, in addition to well-known considerations, such as continued exploitation and slow life histories, there are several underappreciated factors that complicate predator recoveries. These factors include three challenges. First, a priori identification of the suite of trophic interactions, such as resource limitation and competition that will influence recovery can be difficult. Second, defining and accomplishing predator recovery in the context of a dynamic ecosystem requires an appreciation of the timing of recovery, which can determine the relative density of apex predators and other predators and therefore affect competitive outcomes. Third, successful recovery programs require designing adaptive sequences of management strategies that embrace key environmental and species interactions as they emerge. Consideration of recent research on food web modules, alternative stable states, and community assembly offer important insights for predator recovery efforts and restoration ecology more generally. Foremost among these is the importance of a social-ecological perspective in facilitating a long-lasting predator restoration while avoiding unintended consequences.

Intraspecific variation in body size does not alter the effects of mesopredators on prey.

As humans continue to alter the species composition and size structure of marine food webs, it is critical to understand size-dependent effects of predators on prey. Yet, how shifts in predator body size mediate the effect of predators is understudied in tropical marine ecosystems, where anthropogenic harvest has indirectly increased the density and size of small-bodied predators. Here, we combine field surveys and a laboratory feeding experiment in coral reef fish communities to show that small and large predators of the same species can have similar effects. Specifically, surveys show that the presence of a small predator (Paracirrhites arcatus) was correlated with lower chances of prey fish presence, but these correlations were independent of predator size. Experimental trials corroborated the size-independent effect of the predator; attack rates were indistinguishable between small and large predators, suggesting relatively even effects of hawkfish in various size classes on the same type of prey. Our results indicate that the effects of small predators on coral reefs can be size-independent, suggesting that variation in predator size-structure alone may not always affect the functional role of these predators.