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IntroductionChanges in temperature can fundamentally transform how species interact, causing wholesale shifts in ecosystem dynamics and stability. Yet we still have a limited understanding of how temperature-dependence in physiology drives temperature-dependence in species-interactions. For predator-prey interactions, theory predicts that increases in temperature drive increases in metabolism and that animals respond to this increased energy expenditure by ramping up their food consumption to meet their metabolic demand. However, if consumption does not increase as rapidly with temperature as metabolism, increases in temperature can ultimately cause a reduction in consumer fitness and biomass via starvation.MethodsHere we test the hypothesis that increases in temperature cause more rapid increases in metabolism than increases in consumption using the California spiny lobster (Panulirus interruptus) as a model system. We acclimated individual lobsters to temperatures they experience sacross their biogeographic range (11, 16, 21, or 26°C), then measured whether lobster consumption rates are able to meet the increased metabolic demands of rising temperatures.Results and discussionWe show positive effects of temperature on metabolism and predation, but in contrast to our hypothesis, rising temperature caused lobster consumption rates to increase at a faster rate than increases in metabolic demand, suggesting that for the mid-range of temperatures, lobsters are capable of ramping up consumption rates to increase their caloric demand. However, at the extreme ends of the simulated temperatures, lobster biology broke down. At the coldest temperature, lobsters had almost no metabolic activity and at the highest temperature, 33% of lobsters died. Our results suggest that temperature plays a key role in driving the geographic range of spiny lobsters and that spatial and temporal shifts in temperature can play a critical role in driving the strength of species interactions for a key predator in temperate reef ecosystems.

The impacts of invasive lionfish (Pterois volitans/miles) on native coral reef populations in the Western Atlantic Ocean and Caribbean Sea can be enormous. However, how much lionfish differ from native predators and whether their effects outweigh the abundant mesopredators that occupy many reefs invite continued examination. Here, we present empirical evidence from Caribbean Panama and beyond suggesting that lionfish are less abundant than native mesopredators. Furthermore, we show that their direct impacts on survivorship and size distributions of one native prey species are similar to those of a native mesopredator. These results support calls for lionfish management that considers evolving local ecological and social dynamics, including prey community composition, the roles of native mesopredators, and regional goals for conservation and fisheries. Recognition of regional context creates the potential for synergies between conservation actions aimed both at the invasion and other consequential problems such as overexploitation and climate change.

A huge proportion of the world’s population resides in urban areas along the coast. As cities expand, the ability of coastal ecosystems to provide the benefits people derive from nature, ranging from food from fisheries to coastal defense to maritime transportation and beyond, is in question. While it is well understood that coastal development changes ecosystems, quantitative insights about how terrestrial urbanization fundamentally alters ecosystem structure and function in adjacent freshwater and downstream coastal marine habitats remain rare, though a general expectation is that impacts of terrestrial urbanization will attenuate from land to freshwater to coastal marine habitats. Empirical assessments of these phenomena are especially important for species that rely on freshwater and coastal marine habitats at multiple points in their life cycles, including endangered and threatened Pacific salmon (Oncorhynchus spp.). We investigated associations between landscape-scale urbanization and ecosystem structure (biodiversity of epibenthic invertebrate taxa) and function (benthic net primary productivity and decomposition) in freshwater and coastal marine habitats across six pairs of more and less urbanized, coastal watersheds in Puget Sound, WA, USA, using principal components analysis, analysis of covariance, and Mantel tests. Greater upland urbanization was associated with greater reductions in freshwater biodiversity, measured as the density and evenness of epibenthic invertebrate families. In contrast and surprisingly, however, coastal marine biodiversity (measured as the density and evenness of epibenthic invertebrate families) tended to be higher at more urbanized sites, suggesting the potential role of low to moderate levels of urbanization-related disturbance in determining coastal marine biodiversity patterns. We found no statistical association between urbanization and freshwater and coastal marine ecosystem functions, estimated from changes in accumulated algal biomass on tiles (benthic net primary productivity) and loss of biomass from litter bags (decomposition). In addition, there was no evidence that changes in ecosystem structure and function with urbanization were more severe in freshwater than coastal marine habitats, as might be expected if the land-sea boundary diminished effects of landscape-scale urbanization. Our results suggest that the effects of urbanization can be complex and that attention to terrestrial, freshwater, and coastal marine systems in concert will produce more effective, ecosystem-based management.

A suite of processes drive variation in coral populations in space and time, yet our understanding of how variation in coral density affects coral performance is limited. Theory predicts that reductions in density can send coral populations into a predator pit, where concentrated corallivory maintains corals at low densities. In reality, how variation in coral density alters corallivory rates is poorly resolved. Here, we experimentally quantified the effects of corallivory and coral density on growth and survival of small colonies of the staghorn coral Acropora pulchra. Our findings suggest that coral density and corallivory have strong but independent effects on coral performance. In the presence of corallivores, corals suffered high but density-independent mortality. When corallivores were excluded, however, vertical extension rates of colonies increased with increasing densities. While we found no evidence for a predator pit, our results suggest that spatio-temporal variation in corallivore and coral densities can fundamentally alter population dynamics via strong effects on juvenile corals.

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.

Conflict is a common feature in conservation and resource management. Environmental conflicts are frequently attributed to differences in values; however, variability in the perception of facts, rooted in social and cultural differences also underlies conflicts. Such differences in perception have been termed the Rashomon effect after the Kurosawa film. In the present article, we explore a conservation Rashomon effect—a phenomenon that results from a combination of differences in perspective, plausible alternative perspectives of a conservation issue, and the absence of evidence to elevate one perspective above others. As a remedy to the Rashomon effect, policy-makers have turned to scientists as honest brokers who share a common environmental reality. We evaluate this supposition and suggest that scientists, themselves, display Rashomon effects. We suggest that Rashomon effects can be reduced by acknowledging the plurality of reality, embracing epistemic pluralism, and prioritizing an inclusive process of resource management.

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.

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.

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.

Herbivorous fish can increase coral growth and survival by grazing down algal competitors. With coral reefs in global decline, maintaining adequate herbivory has become a primary goal for many managers. However, herbivore biomass targets assume grazing behavior is consistent across different reef systems, even though relatively few have been studied. We document grazing behavior of two scarid species in Antigua, Barbuda, and Bonaire. Our analyses show significant differences in intraspecific feeding rates, time spent grazing, and intensity of grazing across sites, which may alter the ecological impact of a given scarid population. We suggest several hypothesized mechanisms for these behavioral variations that would benefit from explicit testing in future research. As managers set targets to enhance herbivory on reefs, it is critical that we understand potential differences in scarid grazing impact. Our findings demonstrate the variability of grazing behavior across different reef sites and call for further investigation of the drivers and ecological implications of these inconsistencies.

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.

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.

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.

Interspecific competition is often asymmetric, and it can limit the spatial distributions of competitively inferior species within a community. When asymmetric competition involves 2 or more component species, the ranking of species' competitive abilities may form competitive hierarchies (all species of higher rank out-compete all species of lower rank) or competitive networks (at least 1 species of lower rank out-competes =1 species of higher rank). Expectations of resource monopolization and patterns of distribution and abundance among species in competitive networks are expected to differ from those in competitive hierarchies. We conducted a field experiment to evaluate the relative competitive abilities of juveniles of 3 closely related species of reef fish (bird wrasse Gomphosus varius, fivestripe wrasse Thalassoma quinquevittatum and the sixbar wrasse T. hardwicke) on Moorea, French Polynesia. We controlled for intrinsic variation in survivorship among species and found that competition among these 3 species was highly asymmetric, resulting in a simple competitive hierarchy (sequence of competitive ability from superior to inferior competitors): fivestripe wrasse > bird wrasse > sixbar wrasse. We surveyed densities of the 3 reef fish species on 55 patch reefs and observed significant negative spatial covariation between superior and inferior competitors, consistent with competitive hierarchies that limit the spatial distributions of the inferior species (sixbar wrasse). Our work demonstrates that intense asymmetric competition and the formation of competitive hierarchies may be an important determinant of resource monopolization and patterns of distribution and abundance in reef fishes. © Inter-Research 2013.

A growing body of literature has documented myriad effects of human activities on animal behavior, yet the ultimate ecological consequences of these behavioral shifts remain largely uninvestigated. While it is understood that, in the absence of humans, variation in animal behavior can have cascading effects on species interactions, community structure, and ecosystem function, we know little about whether the type or magnitude of human-induced behavioral shifts translate into meaningful ecological change. Here we synthesize empirical literature and theory to create a novel framework for examining the range of behaviorally mediated pathways through which human activities may affect different ecosystem functions. We highlight the few empirical studies that show the potential realization 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 behavioral effects with little ecological relevance, or conversely mismanaging situations in which behavioral effects do drive ecosystem change. The framework presented here can be used to anticipate the nature and likelihood of ecological outcomes and prioritize management among widespread human-induced behavioral shifts, while also suggesting key priorities for future research linking humans, animal behavior, and ecology.

Recent warnings from scientists suggest there is limited time to enact policies to avert wide‐ranging ecological and social damage from climate change. In the United States, discussions about comprehensive national policies to avert climate change have begun, with “Green New Deal” proposals and climate plans put forth by members of Congress and presidential candidates. Oceans are largely absent or separate from these nascent policy proposals. Here, we highlight a policy framework to develop terrestrial and ocean‐integrated policies that can complement and enhance terrestrial‐focused initiatives focused on four specific sectors: 1) energy; 2) transportation; 3) food security; and 4) habitat restoration. Given political friction and constrained budgets, an integrated policy framework offers greater potential to achieve a portfolio of mitigation and adaptation goals in a cost‐effective manner, beyond what could be realized with marine or terrestrial policy solutions alone.

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.

Ecosystems often shift abruptly and dramatically between different regimes in response to human or natural disturbances. When ecosystems tip from one regime to another, the suite of available ecosystem benefits changes, impacting the stakeholders who rely on these benefits. These changes often create some groups who stand to incur large losses if an ecosystem returns to a previous regime. When the participation cost in the decision-making process is extremely high, this can “lock in” ecosystem regimes, making it harder for policy and management to shift ecosystems out of what the majority of society views as the undesirable regime. Public stakeholder meetings often have high costs of participation, thus economic theory predicts they will be dominated by extreme views and often lead to decisions that do not represent the majority viewpoint. Such extreme viewpoints can create strong inertia even when there is broad consensus to manage an ecosystem towards a different regime. In the same manner that reinforcing ecological feedback loops make it harder to exit an ecosystem regime, there are decision-making feedback loops that contribute additional inertia.

Researchers share tips for transforming your group with open data science and teamwork. Researchers share tips for transforming your group with open data science and teamwork.

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.

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 (SIC), an established evidence function. Further, we formulate and justify the use of SIC 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 SIC interval spanning 0.81 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 are moving targets As the human population has grown, our demands on the ocean have increased rapidly. These demands have similarly increased the pressure we place on these systems, and we now cause considerable damage globally. If we want to maintain healthy ocean ecosystems into the future, we must learn to use ocean resources in a sustainable way and facilitate recovery in regions that have declined. Determining how to make these goals a reality, however, is no small challenge. Ingeman et al. review the challenge presented by attempting both to recover and to use ecosystems simultaneously and discuss several approaches for facilitating this essential dual goal. Science , this issue p. eaav1004

The sequence of species colonization is increasingly recognized as an important determinant of community structure, yet the significance of sequence of arrival relative to colonizer abundance is seldom assessed. We manipulated the magnitude and timing of coral-reef fish settlement to investigate whether the competitive dominance of early-arriving Ambon damselfish (i.e., a priority effect) decreased in strength with increasing abundance of late-arriving lemon damselfish. Sequence of arrival had a stronger effect on survival than the number of competing individuals. Relative to when both species arrived simultaneously, lemon damselfish were less aggressive, avoided competitive interactions more frequently and experienced depressed survival when they arrived later than Ambon damselfish, with these effects occurring independently of lemon damselfish abundance. These results suggest priority effects are more important than colonizer abundance and should motivate the integration of priority effects into future studies of density dependence to determine their relative importance.

Decision-makers often rely on expert knowledge, especially in complex and data-poor social-ecological systems (SESs). However, expert knowledge and perceptions of SES structure and function vary; therefore, understanding how these perceptions differ is critical to building knowledge and developing sustainability solutions. Here, we quantify how scientific, local, and traditional knowledge experts vary in their perceptions of food webs centered on Pacific herring—a valuable ecological, economic, and cultural resource in Haida Gwaii, BC, Canada. Expert perceptions of the herring food web varied markedly in structure, and a simulated herring recovery with each of these unique mental models demonstrated wide variability in the perceived importance of herring to the surrounding food web. Using this general approach to determine the logical consequences of expert perceptions of SES structure in the context of potential future management actions, decision-makers can work explicitly toward filling knowledge gaps while embracing a diversity of perspectives.

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.

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.

Variation in the density of organisms among habitat patches is often attributed to variation in inherent patch properties. For example, higher quality patches might have higher densities because they attract more colonists or confer better post-colonization survival. However, variation in occupant density can also be driven by landscape configuration if neighboring patches draw potential colonists away from the focal habitat (a phenomenon we call propagule redirection). Here, we develop and analyze a stochastic model to quantify the role of landscape configuration and propagule redirection on occupant density patterns. We model a system with a dispersive larval stage and a sedentary adult stage. The model includes sensing and decision-making in the colonization stage and density-dependent mortality (a proxy for patch quality) in the post-colonization stage. We demonstrate that spatial variation in colonization is retained when the supply of colonists is not too high, post-colonization density-dependent survival is not too strong, and colonization events are not too frequent. Using a reef fish system, we show that the spatial variation produced by propagule redirection is comparable to spatial variation expected when patch quality varies. Thus, variation in density arising from the spatial patterning of otherwise identical habitat can play an important role in shaping long-term spatial patterns of organisms occupying patchy habitats. Propagule redirection is a potentially powerful mechanism by which landscape configuration can drive variation in occupant densities, and may therefore offer new insights into how populations may shift as landscapes change in response to natural and anthropogenic forces.

As climatic changes and human uses intensify, resource managers and other decision makers are taking actions to either avoid or respond to ecosystem tipping points, or dramatic shifts in structure and function that are often costly and hard to reverse. Evidence indicates that explicitly addressing tipping points leads to improved management outcomes. Drawing on theory and examples from marine systems, we distill a set of seven principles to guide effective management in ecosystems with tipping points, derived from the best available science. These principles are based on observations that tipping points (1) are possible everywhere, (2) are associated with intense and/or multifaceted human use, (3) may be preceded by changes in early‐warning indicators, (4) may redistribute benefits among stakeholders, (5) affect the relative costs of action and inaction, (6) suggest biologically informed management targets, and (7) often require an adaptive response to monitoring. We suggest that early action to preserve system resilience is likely more practical, affordable, and effective than late action to halt or reverse a tipping point. We articulate a conceptual approach to management focused on linking management targets to thresholds, tracking early‐warning signals of ecosystem instability, and stepping up investment in monitoring and mitigation as the likelihood of dramatic ecosystem change increases. This approach can simplify and economize management by allowing decision makers to capitalize on the increasing value of precise information about threshold relationships when a system is closer to tipping or by ensuring that restoration effort is sufficient to tip a system into the desired regime.

Predators are critical components of ecosystems. Globally, conservation efforts have targeted depleted populations of top predators for legal protection, and in many cases, this protection has helped their recoveries. Where the recovery of individual species is the goal, these efforts can be seen as largely successful. From an ecosystem perspective, however, predator recovery can introduce significant new conservation and legal challenges. We highlight three types of conflicts created by a single-species focus: (1) recovering predator populations that increase competition with humans for the same prey, (2) new tradeoffs that emerge when protected predators consume protected prey, and (3) multiple predator populations that compete for the same limited prey. We use two food webs with parallel conservation challenges, the Northeast Pacific Ocean and the Greater Yellowstone Ecosystem, to demonstrate legal/policy conflicts and the policy levers that exist to ameliorate conflicts. In some cases, scientific uncertainty about the ecological interaction hinders progress towards resolving conflicts. In others, available policy options are insufficient. In all cases, management decisions must be made in the face of an unknown future. We suggest a framework that incorporates multispecies science, policy tools, and tradeoff analyses into management.

The oceans are changing more rapidly than ever before. Unprecedented climatic variability is interacting with unmistakable long‐term trends, all against a backdrop of intensifying human activities. What remains unclear, however, is how to evaluate whether conditions have changed sufficiently to provoke major responses of species, habitats, and communities. We developed a framework based on multimodel inference to define ecosystem‐based thresholds for human and environmental pressures in the California Current marine ecosystem. To demonstrate how to apply the framework, we explored two decades of data using gradient forest and generalized additive model analyses, screening for nonlinearities and potential threshold responses of ecosystem states (n = 9) across environmental (n = 6) and human (n = 10) pressures. These analyses identified the existence of threshold responses of five ecosystem states to four environmental and two human pressures. Both methods agreed on threshold relationships in two cases: (1) the winter copepod anomaly and habitat modification, and (2) sea lion pup production and the summer mode of the Pacific Decadal Oscillation (PDO). Considered collectively, however, these alternative analytical approaches imply that as many as five of the nine ecosystem states may exhibit threshold changes in response to negative PDO values in the summer (copepods, scavengers, groundfish, and marine mammals). This result is consistent with the idea that the influence of the PDO extends across multiple trophic levels, but extends current knowledge by defining the nonlinear nature of these responses. This research provides a new way to interpret changes in the intensities of human and environmental pressures as they relate to the ecological integrity of the California Current ecosystem. These insights can be used to make more informed assessments of when and under what conditions intervention, preparation, and mitigation may enhance progress toward ecosystem‐based management goals.

Trophic island biogeography theory predicts that the effects of predators on prey diversity are context dependent in heterogeneous landscapes. Specifically, models predict that the positive effect of habitat area on prey diversity should decline in the presence of predators, and that predators should modify the partitioning of alpha and beta diversity across patchy landscapes. However, experimental tests of the predicted context dependency in top-down control remain limited. Using a factorial field experiment we quantify the effects of a focal predatory fish species (grouper) and habitat characteristics (patch size, fragmentation) on the partitioning of diversity and assembly of coral reef fish communities. We found independent effects of groupers and patch characteristics on prey communities. Groupers reduced prey abundance by 50% and gamma diversity by 45%, with a disproportionate removal of rare species relative to common species (64% and 36% reduction, respectively; an oddity effect). Further, there was a 77% reduction in beta diversity. Null model analysis demonstrated that groupers increased the importance of stochastic community assembly relative to patches without groupers. With regard to patch size, larger patches contained more fishes, but a doubling of patch size led to a modest (36%) increase in prey abundance. Patch size had no effect on prey diversity; however, fragmented patches had 50% higher species richness and modified species composition relative to unfragmented patches. Our findings suggest two different pathways (i.e., habitat or predator shifts) by which natural and/or anthropogenic processes can drive variation in fish biodiversity and community assembly.

Predation is a key process driving coral reef fish population dynamics, with higher per capita prey mortality rates on reefs with more predators. Reef predators often forage together, and at high densities, they may either cooperate or antagonize one another, thereby causing prey mortality rates to be substantially higher or lower than one would expect if predators did not interact. However, we have a limited mechanistic understanding of how prey mortality rates change with predator densities. We re-analyzed a previously published observational dataset to investigate how the foraging response of the coney grouper (Cephalopholis fulva) feeding on the bluehead wrasse (Thalassoma bifasciatum) changed with shifts in predator and prey densities. Using a model-selection approach, we found that per-predator feeding rates were most consistent with a functional response that declines as predator density increases, suggesting either antagonistic interactions among predators or a shared antipredator behavioral response by the prey. Our findings suggest that variation in predator density (natural or anthropogenic) may have substantial consequences for coral reef fish population dynamics.

Marine organisms occasionally settle at exceptional densities, whereby thousands of individuals arrive concurrently. High levels of mortality, which has historically been attributed to predation or competition, often follow this episodic settlement of reef fishes. Here, however, we observed large numbers of newly settled surgeonfish (Ctenochaetus striatus) with white lesions lying dead on the sand amongst patch reefs following separate episodic settlement events in 2006 and 2009 in Moorea, French Polynesia. Pathogens have been identified as an important driver of population dynamics in other marine organisms but less so for reef fishes. Our observations suggest that disease outbreaks may play an underappreciated role as a mechanism of mortality following episodic settlement events in reef fishes.

Coral reefs are the most diverse marine systems in the world, yet our understanding of the processes that maintain such extraordinary diversity remains limited and taxonomically biased toward the most conspicuous species. Cryptofauna that live deeply embedded within the interstitial spaces of coral reefs make up the majority of reef diversity, and many of these species provide important protective services to their coral hosts. However, we know very little about the processes governing the diversity and composition of these less conspicuous but functionally important species. Here, we experimentally quantify the role of predation in driving the community organization of small fishes and decapods that live embedded within Pocillopora eydouxi, a structurally complex, reef-building coral found widely across the Indo-Pacific. We use surveys to describe the natural distribution of predators, and then, factorially manipulate two focal predator species to quantify the independent and combined effects of predator density and identity on P. eydouxi-dwelling cryptofauna. Predators reduced abundance (34 %), species richness (20 %), and modified species composition. Rarefaction revealed that observed reductions in species richness were primarily driven by changes in abundance. Additionally, the two predator species uniquely affected the beta diversity and composition of the prey assemblage. Predators reduced the abundance and modified the composition of a number of mutualist fishes and decapods, whose benefit to the coral is known to be both diversity- and density-dependent. We predict that the density and identity of predators present within P. eydouxi may substantially alter coral performance in the face of an increased frequency and intensity of natural and anthropogenic stressors.

Illegal shark fishing is thought to occur globally, including within so-called ''shark sanctuaries'', marine reserves and even inside UNESCO World Heritage sites, such as the Galapagos Islands. Presumably, this is due to poor local enforcement coupled with the growing international demand (and high economic incentives) for shark and other wildlife products. Understanding illegal shark fishing practices, and specifically catch composition, is important as poaching is identified as a causal factor of global declines in shark populations. Unfortunately, reliable quantitative data on illegal shark fishing are scarce. Here, the catch onboard an illegal shark fishing vessel seized within the borders of the Galapagos Marine Reserve was documented. A total of 379 sharks from seven shark species were found onboard the vessel. A large fraction of the illegal catch was comprised of both female and juvenile sharks (64% and 89%, respectively). Despite numerous recent advances in shark conservation worldwide, this study demonstrates illegal shark fishing is an ongoing concern and that stricter enforcement and legislation is urgently needed, particularly in areas of high biodiversity.

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.

Using rarefaction to isolate the effects of patch size and sampling effort on beta diversity A DRIAN C. S TIER , 1, ! B ENJAMIN M. B OLKER , 2 AND C RAIG W. O SENBERG3 Department of Ecology, Evolution and Marine Biology, University of California, Santa Barbara, California 93106 USA Departments of Mathematics & Statistics and Biology, McMaster University, Hamilton, Ontario L8S 4K1 Canada Odum School of Ecology, University of Georgia, Athens, Georgia 30602 USA Citation: Stier, A. C., B. M. Bolker, and C. W. Osenberg. 2016. Using rarefaction to isolate the effects of patch size and sampling effort on beta diversity. Ecosphere 7(12):e01612. 10.1002/ecs2.1612 Abstract. Beta diversity describes how species composition varies across space and through time. Current estimators of beta diversity typically ignore the effects of within-patch sample size, determined jointly by local abundance and sampling effort. Many ecological processes such as immigration, predation, or nutrient limitation affect abundance and asymptotic beta diversity concurrently; thus, existing metrics may confound changes in asymptotic beta diversity with changes that result from differences in abundance or sampling. Results from a stochastic simulation model illustrate how decreasing within-patch sample size may either increase or decrease observed beta diversity, depending on the type of metric, sample size, and community properties; these changes are easy to understand, and predict, by considering the effects of sampling on variance. A modified, patch-level form of rarefaction controls for variation in within-patch sample size; two case studies illustrate the utility of this approach. Studies seeking a mechanistic link between ecological process and beta diversity will continue to benefit from explicit consideration of sampling effects. Key words: beta diversity; biodiversity; rarefaction; sampling effects. Received 6 October 2016; accepted 10 October 2016. Corresponding Editor: Debra P. C. Peters. Copyright: © 2016 Stier et al. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. ! E-mail: adrian.stier@lifesci.ucsb.edu I NTRODUCTION resource managers to design reserve networks that maximize species complementarity across sites (Kattan et al. 2006). Similarly, beta diversity can be used to describe the rate at which species turn over through time in the presence of human impacts (Dornelas et al. 2014). Knowledge of the factors maintaining diversity among patches can also facilitate biodiversity preservation or recov- ery of diversity following the restoration of a degraded ecosystem (McKnight et al. 2007). Beta diversity is typically examined at two spatial scales: (1) large-scale variation in beta diversity that links species compositional differ- ences across gradients (e.g., with latitude or alti- tude) and encompasses multiple species pools (Condit et al. 2002, McKnight et al. 2007, He and Zhang 2009), and (2) local-scale variation in beta Within-patch (alpha) and among-patch (beta) diversities combine to produce regional patterns of diversity. Traditionally, ecologists have focused on local processes and their influence on alpha diversity. More recently, ecologists have turned to studying the drivers of beta diversity; patterns of beta diversity provide insight into the role of factors that generate species boundaries (Condit et al. 2002), facilitate regional diversity via environmental heterogeneity, and produce alternate community states (Fukami and Naka- jima 2011). From a conservation perspective, beta diversity is arguably as important as alpha diver- sity (Socolar et al. 2016): For example, quantify- ing the spatial scale of beta diversity can allow ❖ www.esajournals.org December 2016 ❖ Volume 7(12) ❖ Article e01612

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.

Synthesis Predation risk experienced by individuals living in groups depends on the balance between predator dilution, competition for refuges, and predator interference or synergy. These interactions operate between prey species as well: the benefits of group living decline in the presence of an alternative prey species. We apply a novel model-fitting approach to data from field experiments to distinguish among competing hypotheses about shifts in predator foraging behavior across a range of predator and prey densities. Our study provides novel analytical tools for analyzing predator foraging behavior and offers insight into the processes driving the dynamics of coral reef fish. Studies of predator foraging behavior typically focus on single prey species and fixed predator densities, ignoring the potential importance of complexities such as predator dilution; predator-mediated effects of alternative prey; heterospecific competition; or predator–predator interactions. Neglecting the effects of prey density is particularly problematic for prey species that live in mixed species groups, where the beneficial effects of predator dilution may swamp the negative effects of heterospecific competition. Here we use field experiments to investigate how the mortality rates of a shoaling coral reef fish (a wrasse: Thalassoma amblycephalum), change as a result of variation in: 1) conspecific density, 2) density of a predator (a hawkfish: Paracirrhites arcatus), and 3) presence of an alternative prey species that competes for space (a damselfish: Pomacentrus pavo). We quantify changes in prey mortality rates from the predator's perspective, examining the effects of added predators or a second prey species on the predator's functional response. Our analysis highlights a model-fitting approach that discriminates amongst multiple hypotheses about predator foraging in a community context. Wrasse mortality decreased with increasing conspecific density (i.e. mortality was inversely density-dependent). The addition of a second predator doubled prey mortality rates, without significantly changing attack rate or handling time – i.e. there was no evidence for predator interference. The presence of a second prey species increased wrasse mortality by 95%; we attribute this increase either to short-term apparent competition (predator aggregation) or to a decrease in handling time of the predator (e.g. through decreased wrasse vigilance). In this system, 1) prey benefit from intraspecific group living though a reduced predation risk, and 2) the benefit of group living is reduced in the presence of an alternative prey species.

The majority of our understanding of mutualisms comes from studies of pairwise interactions. However, many hosts support mutualist guilds, and interactions among mutualists make the prediction of aggregate effects difficult. Here, we apply a factorial experiment to interactions of 'guard' crustaceans that defend their coral host from seastar predators. Predation was reduced by the presence of mutualists (15% reduction in predation frequency and 45% in volume of coral consumed). The frequency of attacks with both mutualists was lower than with a single species, but it did not differ significantly from the expected frequency of independent effects. In contrast, the combined defensive efficacy of both mutualist species reduced the volume of coral tissue lost by 73%, significantly more than the 38% reduction expected from independent defensive efforts, suggesting the existence of a cooperative synergy in defensive behaviors of 'guard' crustaceans. These emergent 'multiple defender effects' are statistically and ecologically analogous to the emergent concept of 'multiple predator effects' known from the predation literature.

One of the most compelling questions in evolutionary biology is why some animals can regenerate injured structures while others cannot. Appendage regeneration appears to be common when viewed across the metazoan phylogeny, yet this ability has been lost in many taxa to varying degrees. Within species, the capacity for regeneration also can vary ontogenetically among individuals. Here we argue that appendage regeneration along the secondary body axis may be constrained by fundamental traits such as body size, aging, life stage, and growth pattern. Studies of the molecular mechanisms affecting regeneration have been conducted primarily with small organisms at early life stages. Such investigations disregard the dramatic shifts in morphology and physiology that organisms undergo as they age, grow, and mature. To help explain interspecific and intraspecific constraints on regeneration, we link particular fundamental traits to specific molecular mechanisms that control regeneration. We present a new synthesis for how these fundamental traits may affect the molecular mechanisms of regeneration at the tissue, cellular, and genomic levels of biological organization. Future studies that explore regeneration in organisms across a broad phylogenetic scale, and within an ontogenetic framework, will help elucidate the proximate mechanisms that modulate regeneration and may reveal new biomedical applications for use in regenerative medicine.