Corals survive severe bleaching event in refuges related to taxa, colony size, and water depth.

Marine heatwaves are increasing in frequency and duration, threatening tropical reef ecosystems through intensified coral bleaching events. We examined a strikingly variable spatial pattern of bleaching in Moorea, French Polynesia following a heatwave that lasted from November 2018 to July 2019. In July 2019, four months after the onset of bleaching, we surveyed > 5000 individual colonies of the two dominant coral genera, Pocillopora and Acropora, at 10 m and 17 m water depths, at six forereef sites around the island where temperature was measured. We found severe bleaching increased with colony size for both coral genera, but Acropora bleached more severely than Pocillopora overall. Acropora bleached more at 10 m than 17 m, likely due to higher light availability at 10 m compared to 17 m, or greater daily temperature fluctuation at depth. Bleaching in Pocillopora corals did not differ with depth but instead varied with the interaction of colony size and Accumulated Heat Stress (AHS), in that larger colonies (> 30 cm) were more sensitive to AHS than mid-size (10-29 cm) or small colonies (5-9 cm). Our findings provide insight into complex interactions among coral taxa, colony size, and water depth that produce high spatial variation in bleaching and related coral mortality.

Microbiome ecological memory and responses to repeated marine heatwaves clarify variation in coral bleaching and mortality.

Microbiomes are essential features of holobionts, providing their hosts with key metabolic and functional traits like resistance to environmental disturbances and diseases. In scleractinian corals, questions remain about the microbiome's role in resistance and resilience to factors contributing to the ongoing global coral decline and whether microbes serve as a form of holobiont ecological memory. To test if and how coral microbiomes affect host health outcomes during repeated disturbances, we conducted a large-scale (32 exclosures, 200 colonies, and 3 coral species sampled) and long-term (28 months, 2018-2020) manipulative experiment on the forereef of Mo'orea, French Polynesia. In 2019 and 2020, this reef experienced the two most severe marine heatwaves on record for the site. Our experiment and these events afforded us the opportunity to test microbiome dynamics and roles in the context of coral bleaching and mortality resulting from these successive and severe heatwaves. We report unique microbiome responses to repeated heatwaves in Acropora retusa, Porites lobata, and Pocillopora spp., which included: microbiome acclimatization in A. retusa, and both microbiome resilience to the first marine heatwave and microbiome resistance to the second marine heatwave in Pocillopora spp. Moreover, observed microbiome dynamics significantly correlated with coral species-specific phenotypes. For example, bleaching and mortality in A. retusa both significantly increased with greater microbiome beta dispersion and greater Shannon Diversity, while P. lobata colonies had different microbiomes across mortality prevalence. Compositional microbiome changes, such as changes to proportions of differentially abundant putatively beneficial to putatively detrimental taxa to coral health outcomes during repeated heat stress, also correlated with host mortality, with higher proportions of detrimental taxa yielding higher mortality in A. retusa. This study reveals evidence for coral species-specific microbial responses to repeated heatwaves and, importantly, suggests that host-dependent microbiome dynamics may provide a form of holobiont ecological memory to repeated heat stress.

Priority effects in coral-macroalgae interactions can drive alternate community paths in the absence of top-down control.

The outcomes of species interactions can vary greatly in time and space with the outcomes of some interactions determined by priority effects. On coral reefs, benthic algae rapidly colonize disturbed substrate. In the absence of top-down control from herbivorous fishes, these algae can inhibit the recruitment of reef-building corals, leading to a persistent phase shift to a macroalgae-dominated state. Yet, corals may also inhibit colonization by macroalgae, and therefore the effects of herbivores on algal communities may be strongest following disturbances that reduce coral cover. Here, we report the results from experiments conducted on the fore reef of Moorea, French Polynesia, where we: (1) tested the ability of macroalgae to invade coral-dominated and coral-depauperate communities under different levels of herbivory, (2) explored the ability of juvenile corals (Pocillopora spp.) to suppress macroalgae, and (3) quantified the direct and indirect effects of fish herbivores and corallivores on juvenile corals. We found that macroalgae proliferated when herbivory was low but only in recently disturbed communities where coral cover was also low. When coral cover was <10%, macroalgae increased 20-fold within 1 year under reduced herbivory conditions relative to high herbivory controls. Yet, when coral cover was high (50%), macroalgae were suppressed irrespective of the level of herbivory despite ample space for algal colonization. Once established in communities with low herbivory and low coral cover, macroalgae suppressed recruitment of coral larvae, reducing the capacity for coral replenishment. However, when we experimentally established small juvenile corals (2 cm diameter) following a disturbance, juvenile corals inhibited macroalgae from invading local neighborhoods, even in the absence of herbivores, indicating a strong priority effect in macroalgae-coral interactions. Surprisingly, fishes that initially facilitated coral recruitment by controlling algae had a net negative effect on juvenile corals via predation. Corallivores reduced the growth rates of corals exposed to fishes by ~30% relative to fish exclosures, despite increased competition with macroalgae within the exclosures. These results highlight that different processes are important for structuring coral reef ecosystems at different successional stages and underscore the need to consider multiple ecological processes and historical contingencies to predict coral community dynamics.

Size-dependent mortality of corals during marine heatwave erodes recovery capacity of a coral reef.

For many long-lived taxa, such as trees and corals, older, and larger individuals often have the lowest mortality and highest fecundity. However, climate change-driven disturbances such as droughts and heatwaves may fundamentally alter typical size-dependent patterns of mortality and reproduction in these important foundation taxa. Working in Moorea, French Polynesia, we investigated how a marine heatwave in 2019, one of the most intense marine heatwaves at our sites over the past 30 years, drove patterns of coral bleaching and mortality. The marine heatwave drove island-wide mass coral bleaching that killed up to 76% and 65% of the largest individuals of the two dominant coral genera, Pocillopora and Acropora, respectively. Colonies of Pocillopora and Acropora ≥30 cm diameter were ~3.5× and ~1.3×, respectively, more likely to die than colonies <30-cm diameter. Typically, annual mortality in these corals is concentrated on the smallest size classes. Yet, this heatwave dramatically reshaped this pattern, with heat stress disproportionately killing larger coral colonies and equalizing annual mortality rates across the size spectrum. This shift in the size-mortality relationship reduced the overall fecundity of these genera by >60% because big corals are disproportionately important for reproduction on reefs. Additionally, the survivorship of microscopic coral recruits, critical for the recovery of corals following disturbances, declined to 2%, over an order of magnitude lower compared to a year without elevated thermal stress, where 33% of coral recruits survived. While other research has shown that larger corals can bleach more frequently than smaller corals, we show the severe impact this phenomenon can have at the reef-wide scale. As marine heatwaves become more frequent and intense, disproportionate mortality of the largest, most fecund corals and near-complete loss of entire cohorts of newly-settled coral recruits will likely reduce the recovery capacity of these iconic ecosystems.

Spatial covariation in nutrient enrichment and fishing of herbivores in an oceanic coral reef ecosystem.

Both natural and anthropogenic stressors are increasing on coral reefs, resulting in large-scale loss of coral and potential shifts from coral- to macroalgae-dominated community states. Two factors implicated in shifts to macroalgae are nutrient enrichment and fishing of reef herbivores. Although either of these factors alone could facilitate establishment of macroalgae, reefs may be particularly vulnerable to coral-to-algae phase shifts in which strong bottom-up forcing from nutrient enrichment is accompanied by a weakening of herbivore control of macroalgae via intense fishing. We explored spatial heterogeneity and covariance in these drivers on reefs in the lagoons of Moorea, French Polynesia, where the local fishery heavily targets herbivorous fishes and there are spatially variable inputs of nutrients from agricultural fertilizers and wastewater systems. Spatial patterns of fishing and nutrient enrichment were not correlated at the two landscape scales we examined: among the 11 interconnected lagoons around the island or among major habitats (fringing reef, mid-lagoon, back reef) within a lagoon. This decoupling at the landscape scale resulted from patterns of covariation between enrichment and fishing that differed qualitatively between cross-shore and long-shore directions. At the cross-shore scale, nutrient enrichment declined but fishing increased from shore to the crest of the barrier reef. By contrast, nutrient enrichment and fishing were positively correlated in the long-shore direction, with both increasing with proximity to a pass in the barrier reef. Contrary to widespread assumptions in the scientific literature that human coastal population density correlates with impact on marine ecosystems and that fishing effort declines linearly with distance from the shore, these local stressors produced a complex spatial mosaic of reef vulnerabilities. Our findings support spatially explicit management involving the control of anthropogenic nutrients and strategic reductions in fishing pressure on herbivores by highlighting specific areas to target for management actions.

Landscape-scale patterns of nutrient enrichment in a coral reef ecosystem: implications for coral to algae phase shifts.

Nutrient pollution is altering coastal ecosystems worldwide. On coral reefs, excess nutrients can favor the production of algae at the expense of reef-building corals, yet the role of nutrients in driving community changes such as shifts from coral to macroalgae is not well understood. Here we investigate the potential role of anthropogenic nutrient loading in driving recent coral-to-macroalgae phase shifts on reefs in the lagoons surrounding the Pacific island of Moorea, French Polynesia. We use nitrogen (N) tissue content and stable isotopes (δ15 N) in an abundant macroalga (Turbinaria ornata) together with empirical models of nutrient discharge to describe spatial and temporal patterns of nutrient enrichment in the lagoons. We then employ time series data to test whether recent increases in macroalgae are associated with nutrients. Our results revealed that patterns of N enrichment were linked to several factors, including rainfall, wave-driven circulation, and distance from anthropogenic nutrient sources, especially human sewage. Reefs near large watersheds, where inputs of N from sewage and agriculture are high, have been consistently enriched in N for at least the last decade. In many of these areas, corals have decreased and macroalgae have increased, while reefs with lower levels of N input have maintained high cover of coral and low cover of macroalgae. Importantly, these patchy phase shifts to macroalgae have occurred despite substantial island-wide increases in the density and biomass of herbivorous fishes over the time period. Together, these results indicate that nutrient loading may be an important driver of coral-to-macroalgae phase shifts in the lagoons of Moorea even though the reefs harbor an abundant and diverse herbivore assemblage. These results emphasize the important role that bottom-up factors can play in driving coral-to-macroalgae phase shifts and underscore the critical importance of watershed management for reducing inputs of nutrients and other land-based pollutants to coral reef ecosystems.

Intestinal microbes: an axis of functional diversity among large marine consumers.

Microbes are ubiquitous throughout the world's oceans, yet the manner and extent of their influence on the ecology and evolution of large, mobile fauna remains poorly understood. Here, we establish the intestinal microbiome as a hidden, and potentially important, 'functional trait' of tropical herbivorous fishes-a group of large consumers critical to coral reef resilience. Using field observations, we demonstrate that five common Caribbean fish species display marked differences in where they feed and what they feed on. However, in addition to space use and feeding behaviour-two commonly measured functional traits-we find that interspecific trait differences are even more pronounced when considering the herbivore intestinal microbiome. Microbiome composition was highly species specific. Phylogenetic comparison of the dominant microbiome members to all known microbial taxa suggest that microbiomes are comprised of putative environmental generalists, animal-associates and fish specialists (resident symbionts), the latter of which mapped onto host phylogeny. These putative symbionts are most similar to-among all known microbes-those that occupy the intestines of ecologically and evolutionarily related herbivorous fishes in more distant ocean basins. Our findings therefore suggest that the intestinal microbiome may be an important functional trait among these large-bodied consumers.

Nitrogen pollution interacts with heat stress to increase coral bleaching across the seascape.

Climate change is increasing the frequency and magnitude of temperature anomalies that cause coral bleaching, leading to widespread mortality of stony corals that can fundamentally alter reef structure and function. However, bleaching often is spatially variable for a given heat stress event, and drivers of this heterogeneity are not well resolved. While small-scale experiments have shown that excess nitrogen can increase the susceptibility of a coral colony to bleaching, we lack evidence that heterogeneity in nitrogen pollution can shape spatial patterns of coral bleaching across a seascape. Using island-wide surveys of coral bleaching and nitrogen availability within a Bayesian hierarchical modeling framework, we tested the hypothesis that excess nitrogen interacts with temperature anomalies to alter coral bleaching for the two dominant genera of branching corals in Moorea, French Polynesia. For both coral genera, Pocillopora and Acropora, heat stress primarily drove bleaching prevalence (i.e., the proportion of colonies on a reef that bleached). In contrast, the severity of bleaching (i.e., the proportion of an individual colony that bleached) was positively associated with both heat stress and nitrogen availability for both genera. Importantly, nitrogen interacted with heat stress to increase bleaching severity up to twofold when nitrogen was high and heat stress was relatively low. Our finding that excess nitrogen can trigger severe bleaching even under relatively low heat stress implies that mitigating nutrient pollution may enhance the resilience of coral communities in the face of mounting stresses from global climate change.

Identity of coral reef herbivores drives variation in ecological processes over multiple spatial scales.

Overexploitation of key species can negatively impact ecosystem processes, so understanding the ecological roles of individual species is critical for improving ecosystem management. Here, we use coral reefs and the process of herbivory as a model to examine how species identity of consumers influence ecosystem processes to inform management of these consumers. Herbivorous fishes can facilitate the recruitment, growth, and recovery of corals by controlling the fast-growing algae that can outcompete corals for space. However, herbivorous fish guilds are species rich with important differences among species in diet, movement, and habitat preferences. Yet, we lack a general understanding of (1) how these species-specific differences in feeding and behavior scale up to reef-wide rates of ecosystem processes and (2) how species identity and diversity impact these processes. To address these knowledge gaps, we used field observations to derive key species- and size-specific foraging parameters for nine herbivorous parrotfish species on coral reefs in the Florida Keys, USA. We then combined these foraging parameters with fish survey data spanning multiple spatial scales to estimate the rates of three ecosystem processes: area of reef grazed, amount of macroalgae removed, and rate of bioerosion. We found that predicted rates of ecological processes varied dramatically among habitats and among reef zones within habitats, driven primarily by variation in abundance among species with different foraging behaviors. In some cases, assemblages with similar levels of total biomass had different rates of ecological processes, and in others, assemblages with different biomass had similar rates of ecological processes. Importantly, our models of herbivory using species-specific parameters differed from those using genus-level parameters by up to 300% in rates of ecological processes, highlighting the importance of herbivore identity in this system. Our results indicate that there may be little overlap in the roles species play, suggesting that some systems may be vulnerable to loss of ecological function with the reduction or loss of just a few species. This work provides a framework that can be applied across the region to predict how changes in management may affect the ecological impact of these important herbivores.

Macroalgae size refuge from herbivory promotes alternative stable states on coral reefs.

Coral reef systems can undergo rapid transitions from coral-dominated to macroalgae-dominated states following disturbances, and models indicate that these may sometimes represent shifts between alternative stable states. While several mechanisms may lead to alternate stable states on coral reefs, only a few have been investigated theoretically. We explore a model that illustrates that reduced vulnerability of macroalgae to herbivory as macroalgae grow and mature could be an important mechanism: when macroalgae are palatable to herbivores as juveniles, but resistant as adults, coral-dominated and algae-dominated states are bistable across a wide range of parameter space. We compare two approaches to global sensitivity analysis to rank the relative importance of the model parameters in determining the presence and magnitude of alternative stable states, and find that the two most influential parameters are the death rate of coral and the rate of maturation of algae out of the vulnerable stage. The Random Forest approach for global sensitivity analysis, recently adopted by ecologists, provides a more efficient method for ranking the relative importance of parameters than a variance-based approach that has been used frequently by computer scientists and engineers. Our results suggest that managing reefs to reduce chronic stressors that cause coral mortality and/or enhance the growth rates of algae can help prevent reefs from becoming locked in a macroalgae-dominated state.

Recruitment Drives Spatial Variation in Recovery Rates of Resilient Coral Reefs.

Tropical reefs often undergo acute disturbances that result in landscape-scale loss of coral. Due to increasing threats to coral reefs from climate change and anthropogenic perturbations, it is critical to understand mechanisms that drive recovery of these ecosystems. We explored this issue on the fore reef of Moorea, French Polynesia, following a crown-of-thorns seastar outbreak and cyclone that dramatically reduced cover of coral. During the five-years following the disturbances, the rate of re-establishment of coral cover differed systematically around the triangular-shaped island; coral cover returned most rapidly at sites where the least amount of live coral remained after the disturbances. Although sites differed greatly in the rate of return of coral, all showed at least some evidence of re-assembly to their pre-disturbance community structure in terms of relative abundance of coral taxa and other benthic space holders. The primary driver of spatial variation in recovery was recruitment of sexually-produced corals; subsequent growth and survivorship were less important in shaping the spatial pattern. Our findings suggest that, although the coral community has been resilient, some areas are unlikely to attain the coral cover and taxonomic structure they had prior to the most recent disturbances before the advent of another landscape-scale perturbation.

The importance of individual and species-level traits for trophic niches among herbivorous coral reef fishes.

Resolving how species compete and coexist within ecological communities represents a long-standing challenge in ecology. Research efforts have focused on two predominant mechanisms of species coexistence: complementarity and redundancy. But findings also support an alternative hypothesis that within-species variation may be critical for coexistence. Our study focuses on nine closely related and ecologically similar coral reef fish species to test the importance of individual- versus species-level traits in determining the size of dietary, foraging substrate, and behavioural interaction niches. Specifically, we asked: (i) what level of biological organization best describes individual-level niches? and (ii) how are herbivore community niches partitioned among species, and are niche widths driven by species- or individual-level traits? Dietary and foraging substrate niche widths were best described by species identity, but no level of taxonomy explained behavioural interactions. All three niches were dominated by only a few species, contrasting expectations of niche complementarity. Species- and individual-level traits strongly drove foraging substrate and behavioural niches, respectively, whereas the dietary niche was described by both. Our findings underscored the importance of species-level traits for community-level niches, but highlight that individual-level trait variation within a select few species may be a key driver of the overall size of niches.

Coral Reef Resilience, Tipping Points and the Strength of Herbivory.

Coral reefs increasingly are undergoing transitions from coral to macroalgal dominance. Although the functional roles of reef herbivores in controlling algae are becoming better understood, identifying possible tipping points in the herbivory-macroalgae relationships has remained a challenge. Assessment of where any coral reef ecosystem lies in relation to the coral-to-macroalgae tipping point is fundamental to understanding resilience properties, forecasting state shifts, and developing effective management practices. We conducted a multi-year field experiment in Moorea, French Polynesia to estimate these properties. While we found a sharp herbivory threshold where macroalgae escape control, ambient levels of herbivory by reef fishes were well above that needed to prevent proliferation of macroalgae. These findings are consistent with previously observed high resilience of the fore reef in Moorea. Our approach can identify vulnerable coral reef systems in urgent need of management action to both forestall shifts to macroalgae and preserve properties essential for resilience.

Resource partitioning along multiple niche axes drives functional diversity in parrotfishes on Caribbean coral reefs.

The recent loss of key consumers to exploitation and habitat degradation has significantly altered community dynamics and ecosystem function across many ecosystems worldwide. Predicting the impacts of consumer losses requires knowing the level of functional diversity that exists within a consumer assemblage. In this study, we document functional diversity among nine species of parrotfishes on Caribbean coral reefs. Parrotfishes are key herbivores that facilitate the maintenance and recovery of coral-dominated reefs by controlling algae and provisioning space for the recruitment of corals. We observed large functional differences among two genera of parrotfishes that were driven by differences in diet. Fishes in the genus Scarus targeted filamentous algal turf assemblages, crustose coralline algae, and endolithic algae and avoided macroalgae, while fishes in the genus Sparisoma preferentially targeted macroalgae. However, species with similar diets were dissimilar in other attributes, including the habitats they frequented, the types of substrate they fed from, and the spatial scale at which they foraged. These differences indicate that species that appear to be functionally redundant when looking at diet alone exhibit high levels of complementarity when we consider multiple functional traits. By identifying key functional differences among parrotfishes, we provide critical information needed to manage parrotfishes to enhance the resilience of coral-dominated reefs and reverse phase shifts on algal-dominated reefs throughout the wider Caribbean. Further, our study provides a framework for predicting the impacts of consumer losses in other species rich ecosystems.

How will coral reef fish communities respond to climate-driven disturbances? Insight from landscape-scale perturbations.

Global climate change is rapidly altering disturbance regimes in many ecosystems including coral reefs, yet the long-term impacts of these changes on ecosystem structure and function are difficult to predict. A major ecosystem service provided by coral reefs is the provisioning of physical habitat for other organisms, and consequently, many of the effects of climate change on coral reefs will be mediated by their impacts on habitat structure. Therefore, there is an urgent need to understand the independent and combined effects of coral mortality and loss of physical habitat on reef-associated biota. Here, we use a unique series of events affecting the coral reefs around the Pacific island of Moorea, French Polynesia to differentiate between the impacts of coral mortality and the degradation of physical habitat on the structure of reef fish communities. We found that, by removing large amounts of physical habitat, a tropical cyclone had larger impacts on reef fish communities than an outbreak of coral-eating sea stars that caused widespread coral mortality but left the physical structure intact. In addition, the impacts of declining structural complexity on reef fish assemblages accelerated as structure became increasingly rare. Structure provided by dead coral colonies can take up to decades to erode following coral mortality, and, consequently, our results suggest that predictions based on short-term studies are likely to grossly underestimate the long-term impacts of coral decline on reef fish communities.

Ecology and evolution affect network structure in an intimate marine mutualism.

Elucidating patterns and causes of interaction among mutualistic species is a major focus of ecology, and recent meta-analyses of terrestrial networks show that network-level reciprocal specialization tends to be higher in intimate mutualisms than in nonintimate mutualisms. It is largely unknown, however, whether this pattern holds for and what factors affect specialization in marine mutualisms. Here we present the first analysis of network specialization ([Formula: see text]) for marine mutualistic networks. Specialization among eight Indo-Pacific networks of obligate mutualistic gobies and shrimps was indistinguishable from that among comparably intimate terrestrial mutualisms (ants-myrmecophytes) and higher than that among nonintimate ones (seed dispersers). Specialization was affected by variability in habitat use for both gobies and shrimps and by phylogenetic history for shrimps. Habitat use was phylogenetically conserved among shrimp, and thus effects of shrimp phylogeny on partner choice were mediated in part by habitat. By contrast, habitat use and pairing patterns in gobies were not related to phylogenetic history. This asymmetry appears to result from evolutionary constraints on partner use in shrimps and convergence among distantly related gobies to utilize burrows provided by multiple shrimp species. Results indicate that the evolution of mutualism is affected by life-history characteristics that transcend environments and that different factors constrain interactions in disparate ecosystems.

Mutualistic cleaner fish initiate trait-mediated indirect interactions by influencing the behaviour of coral predators.

1. Indirect interactions resulting from changes in organismal traits such as behaviour [i.e. trait-mediated indirect interactions (TMIIs)] are widespread in biological communities, yet few studies have explored the potential for mutualisms to initiate TMIIs. 2. This study used a combination of behavioural observations and manipulative field experiments to investigate potential TMIIs resulting from a mutualism between specialized cleaner fish and the 'clients' that visit cleaners for the removal of ectoparasites. 3. Behavioural observations indicate that the bluestreak cleaner wrasse, Labroides dimidiatus, increases local predation pressure on corals at cleaner stations by attracting corallivorous butterflyfish to their territories. 4. Observations of the ornate butterflyfish, Chaetodon ornatissimus, suggest a trade-off between seeking cleaning and foraging; individuals decreased their foraging rate at cleaner stations and shifted their diet to include a greater proportion of less preferred prey items. Nonetheless, predation pressure on corals was higher at cleaner stations because the spatial response of butterflyfish to cleaners more than compensated for their lower foraging rates. 5. The results of a field experiment suggest that the greater predation pressure observed at cleaner stations may be sufficient to reduce the growth rate of the unpreferred coral Porites rus. 6. Together, these results emphasize the need to consider mutualists as potential initiators of TMIIs and highlight the importance of integrating individual movement into conceptual analyses of TMIIs.

Herbivory, connectivity, and ecosystem resilience: response of a coral reef to a large-scale perturbation.

Coral reefs world-wide are threatened by escalating local and global impacts, and some impacted reefs have shifted from coral dominance to a state dominated by macroalgae. Therefore, there is a growing need to understand the processes that affect the capacity of these ecosystems to return to coral dominance following disturbances, including those that prevent the establishment of persistent stands of macroalgae. Unlike many reefs in the Caribbean, over the last several decades, reefs around the Indo-Pacific island of Moorea, French Polynesia have consistently returned to coral dominance following major perturbations without shifting to a macroalgae-dominated state. Here, we present evidence of a rapid increase in populations of herbivorous fishes following the most recent perturbation, and show that grazing by these herbivores has prevented the establishment of macroalgae following near complete loss of coral on offshore reefs. Importantly, we found the positive response of herbivorous fishes to increased benthic primary productivity associated with coral loss was driven largely by parrotfishes that initially recruit to stable nursery habitat within the lagoons before moving to offshore reefs later in life. These results underscore the importance of connectivity between the lagoon and offshore reefs for preventing the establishment of macroalgae following disturbances, and indicate that protecting nearshore nursery habitat of herbivorous fishes is critical for maintaining reef resilience.

High-quality habitat and facilitation ameliorate competitive effects of prior residents on new settlers.

Many species disperse during their lifetime. Two factors that can affect the performance of individuals following dispersal are the presence of conspecifics and intrinsic habitat quality at the settlement site. Detecting the influence of these factors can be difficult for at least two reasons: (1) the outcomes of interactions with conspecifics are often variable including both competition and facilitation, and (2) selection of high quality habitats often leads to positive covariance between habitat quality and density. In this study, I investigate positive and negative effects of resident blue streak cleaner wrasse (Labroides dimidiatus) on the growth and survival of recently settled conspecifics while accounting for habitat quality. Juvenile L. dimidiatus settle near adult conspecifics, but likely have to compete with resident adults for access to food. However, field experiments indicate that settlers have access to more resources at occupied sites, and as a result, grow faster despite evidence for competition with residents. This result is a direct consequence of two factors: (1) resident conspecifics facilitate settlers by attracting client fish, and (2) resident conspecifics are strongly associated with high quality habitat. These results highlight the need to simultaneously consider habitat quality and competitive and facilitative interactions between conspecifics when making inferences about ecological processes from spatial patterns of individual performance.