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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.

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.

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.

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.

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

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.

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.

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.

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.

Increased habitat availability or quality can alter production of habitat-dependent organisms in two contrasting ways: (1) by enhancing input of new colonists to the new sites (the Field-of-Dreams Hypothesis); and (2) by drawing colonists away from existing sites (the Propagule Redirection Hypothesis), and thus reducing the deleterious effects of density. We conducted a field experiment on coral reef fishes in Moorea, French Polynesia, to quantify how differing levels of habitat availability (controlling for quality) increased and/or redirected colonizing larval fish. Focal reefs without neighboring reefs received two to four times more settlers than reefs with adjacent habitat, demonstrating that increased habitat redirected larval fish. At the scale of the entire reef array, total colonization increased 1.3-fold in response to a sixfold increase in reef area (and a 2.75-fold increase in adjusted habitat availability). Thus, propagules were both increased and redirected, a result midway between the Field-of-Dreams and Propagule Redirection Hypotheses. A recruitment model using our data and field estimates of density-dependent recruitment predicts that habitat addition increases recruitment primarily by ameliorating the negative effects of competition at existing sites rather than increasing colonization at the new sites per se. Understanding long-term implications of these effects depends upon the interplay among habitat dynamics, population connectivity, colonization dynamics, and density dependence.

Research in eco-evolutionary dynamics and community genetics has demon- strated that variation within a species can have strong impacts on associated communities and ecosystem processes. Yet, these studies have centred around individual focal species and at single trophic levels, ignoring the role of phenotypic variation in multiple taxa within an ecosystem. Given the ubi- quitous nature of local adaptation, and thus intraspecific variation, we sought to understand how combinations of intraspecific variation in multiple species within an ecosystem impacts its ecology. Using two species that co-occur and demonstrate adaptation to their natal environments, black cottonwood (Populus trichocarpa) and three-spined stickleback (Gasterosteus aculeatus), we investigated the effects of intraspecific phenotypic variation on both top-down and bottom-up forces using a large-scale aquatic mesocosm experiment. Black cottonwood genotypes exhibit genetic variation in their productivity and consequently their leaf litter subsidies to the aquatic system, which mediates the strength of top-down effects from stickleback on prey abundances. Abundances of four common invertebrate prey species and available phosphorous, the most critically limiting nutrient in freshwater systems, are dictated by the interaction between genetic variation in cotton- wood productivity and stickleback morphology. These interactive effects fit with ecological theory on the relationship between productivity and top- down control and are comparable in strength to the effects of predator addition. Our results illustrate that intraspecific variation, which can evolve rapidly, is an under-appreciated driver of community structure and eco- system function, demonstrating that a multi-trophic perspective is essential to understanding the role of evolution in structuring ecological patterns. Fil: Rudman, Seth M.. University Of British Columbia; Canadá Fil: Rodriguez Cabal, Mariano Alberto. University Of British Columbia; Canadá. Universidad Nacional del Comahue; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Patagonia Norte. Instituto de Investigación en Biodiversidad y Medioambiente; Argentina Fil: Stier, Adrian. University Of British Columbia; Canadá. Nnational Center for Ecological Analysis and Synthesis; Estados Unidos Fil: Sato, Takuya . Kobe University. Graduate school of Science. Department of Biology; Japón Fil: Heavyside, Julian. University Of British Columbia; Canadá Fil: El Sabaawi, Rana W. . University Of Victoria; Canadá Fil: Crutsinger, Gregory M. . University Of British Columbia; Canadá

Top predators are a critical part of healthy ecosystems. Yet, these species are often absent from spatially isolated habitats leading to the pervasive view that fragmented ecological communities collapse from the top down. Here we study reef fish from coral reef communities across the Pacific Ocean. Our analysis shows that species richness of reef fish top predators is relatively stable across habitats that vary widely in spatial isolation and total species richness. In contrast, species richness of prey reef fish declines rapidly with increasing isolation. By consequence, species-poor communities from isolated islands have three times as many predator species per prey species as near-shore communities. We develop and test a colonization-extinction model to reveal how larval dispersal patterns shape this ocean-scale gradient in trophic structure.

Twenty-first century conservation is centered on negotiating trade-offs between the diverse needs of people and the needs of the other species constituting coupled human-natural ecosystems. Marine forage fishes, such as sardines, anchovies and herring, are a nexus for such trade-offs because they are both central nodes in marine food webs and targeted by fisheries. An important example is Pacific herring, Clupea pallisii in the Northeast Pacific. Herring populations are subject to two distinct fisheries: one that harvests adults and one that harvests spawned eggs. We develop stochastic, age-structured models to assess the interaction between fisheries, herring populations and the persistence of predators reliant on herring populations. We show that egg- and adult-fishing have asymmetric effects on herring population dynamics - herring stocks can withstand higher levels of egg harvest before becoming depleted. Second, ecosystem thresholds proposed to ensure the persistence of herring predators do not necessarily pose more stringent constraints on fisheries than conventional, fishery driven harvest guidelines. Our approach provides a general template to evaluate ecosystem trade-offs between stage-specific harvest practices in relation to environmental variability, the risk of fishery closures and the risk of exceeding ecosystem thresholds intended to ensure conservation goals are met.

Introduction Marine protected areas (MPAs) are increasingly used as tools to conserve biodiversity, manage fishing effort, and facilitate recovery of degraded ecosystems (Roberts et al ., 2001; Sale, 2002; Hastings and Botsford, 2003; Cinner et al ., 2006). Marine protected areas are expected to produce long-lasting local increases in the density, size, diversity, and/or productivity of marine organisms within MPA boundaries (Russ and Alcala 1996; Claudet et al ., 2006; Chapter 2), as well as regional increases outside of the MPA via spillover from the MPA to sites that continue to be fished (Chapter 3). Assessment of the actual effects of MPAs relative to these goals is essential for adaptive management and decision-making. Despite a large number of assessments, however, there remains considerable uncertainty about the actual effects of MPAs (e.g., Osenberg et al ., 2006). The central question (and challenge) that underlies the assessment of any MPA is easy to express but difficult to measure: “How does the state of the system (e.g., density or size of a target organism) within the MPA compare to the state that would have existed had the MPA never been established?” The former can be directly observed; however, the latter cannot and must therefore be estimated through indirect means (Stewart-Oaten et al ., 1986; Osenberg and Schmitt, 1996; Stewart-Oaten, 1996a, 1996b; Stewart-Oaten and Bence, 2001; Osenberg et al ., 2006). Several different approaches have been proposed to estimate this baseline (Box 6.1; see also Osenberg et al ., 2006), yet, considerable controversy still exists (Box 6.2).

The global decline in estuarine and coastal ecosystems (ECEs) is affecting a number of critical benefits, or ecosystem services. We review the main ecological functions and their services across a variety of ECEs, including marshes, mangroves, nearshore coral reefs, seagrass beds, and sand beaches and dunes. We cite estimates of the key economic values arising from these services, and discuss how the natural variability of ECE impacts their benefits, the synergistic relationships of ECE across seascapes, and the management implications.

Coral and rocky reef fish populations are widely used as model systems for the experimental exploration of density-dependent vital rates, but patterns of density-dependent mortality in these systems are not yet fully understood. In particular, the paradigm for strong, directly density-dependent (DDD) postsettlement mortality stands in contrast to recent evidence for inversely density-dependent (IDD) mortality. We review the processes responsible for DDD and IDD per capita mortality in reef fishes, noting that the pattern observed depends on predator and prey behavior, the spatial configuration of the reef habitat, and the spatial and temporal scales of observation. Specifically, predators tend to produce DDD prey mortality at their characteristic spatial scale of foraging, but prey mortality is IDD at smaller spatial scales due to attack-abatement effects (e.g., risk dilution). As a result, DDD mortality may be more common than IDD mortality on patch reefs, which tend to constrain predator foraging to the same scale as prey aggregation, eliminating attack-abatement effects. Additionally, adjacent groups of prey on continuous reefs may share a subset of refuges, increasing per capita refuge availability and relaxing DDD mortality relative to prey on patch reefs, where the patch edge could prevent such refuge sharing. These hypotheses lead to a synthetic framework to predict expected mortality patterns for a variety of scenarios. For nonsocial, nonaggregating species and species that aggregate in order to take advantage of spatially clumped refuges, IDD mortality is possible but likely superseded by DDD refuge competition, especially on patch reefs. By contrast, for species that aggregate socially, mortality should be IDD at the scale of individual aggregations but DDD at larger scales. The results of nearly all prior reef fish studies fit within this framework, although additional work is needed to test many of the predicted outcomes. This synthesis reconciles some apparent contradictions in the recent reef fish literature and suggests the importance of accounting for the scale-sensitive details of predator and prey behavior in any study system.

Many communities experience repeated periods of colonization due to seasonally regenerating habitats or pulsed arrival of young-of-year. When an individual's persistence in a community depends upon the strength of competitive interactions, changes in the timing of arrival relative to the arrival of a competitor can modify competitive strength and, ultimately, establishment in the community. We investigated whether the strength of intracohort competitive interactions between recent settlers of the reef fishes Thalassoma hardwicke and T. quinquevittatum are dependent on the sequence and temporal separation of their arrival into communities. To achieve this, we manipulated the sequence and timing of arrival of each species onto experimental patch reefs by simulating settlement pulses and monitoring survival and aggressive interactions. Both species survived best in the absence of competitors, but when competitors were present, they did best when they arrived at the same time. Survival declined as each species entered the community progressively later than its competitor and as aggression by its competitor increased. Intraspecific effects of resident T. hardwicke were similar to interspecific effects. This study shows that the strength of competition depends not only on the identity of competitors, but also on the sequence and timing of their interactions, suggesting that when examining interaction strengths, it is important to identify temporal variability in the direction and magnitude of their effects. Furthermore, our findings provide empirical evidence for the importance of competitive lotteries in the maintenance of species diversity in demographically open marine systems.