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Identifying locations of refugia from the thermal stresses of climate change for coral reefs and better managing them is one of the key recommendations for climate change adaptation. We review and summarize approximately 30 years of applied research focused on identifying climate refugia to prioritize the conservation actions for coral reefs under rapid climate change. We found that currently proposed climate refugia and the locations predicted to avoid future coral losses are highly reliant on  excess heat metrics, such as degree heating weeks. However, many existing alternative environmental, ecological, and life-history variables could be used to identify other types of refugia that lead to the desired diversified portfolio for coral reef conservation. To improve conservation priorities for coral reefs, there is a need to evaluate and validate the predictions of climate refugia with long-term field data on coral abundance, diversity, and functioning. There is also the need to identify and safeguard locations displaying resistance toprolonged exposure to heat waves and the ability to recover quickly after thermal exposure. We recommend using more metrics to identify a portfolio of potential refugia sites for coral reefs that can avoid, resist, and recover from exposure to high ocean temperatures and the consequences of climate change, thereby shifting past efforts focused on avoidance to a diversified risk-spreading portfolio that can be used to improve strategic coral reef conservation in a rapidly warming climate., (© 2023 The Authors. Conservation Biology published by Wiley Periodicals LLC on behalf of Society for Conservation Biology.)

Scientists and managers rely on indicator taxa such as coral and macroalgal cover to evaluate the effects of human disturbance on coral reefs, often assuming a universally positive relationship between local human disturbance and macroalgae. Despite evidence that macroalgae respond to local stressors in diverse ways, there have been few efforts to evaluate relationships between specific macroalgae taxa and local human-driven disturbance. Using genus-level monitoring data from 1205 sites in the Indian and Pacific Oceans, we assess whether macroalgae percent cover correlates with local human disturbance while accounting for factors that could obscure or confound relationships. Assessing macroalgae at genus level revealed that no genera were positively correlated with all human disturbance metrics. Instead, we found relationships between the division or genera of algae and specific human disturbances that were not detectable when pooling taxa into a single functional category, which is common to many analyses. The convention to use percent cover of macroalgae as an indication of local human disturbance therefore likely obscures signatures of local anthropogenic threats to reefs. Our limited understanding of relationships between human disturbance, macroalgae taxa, and their responses to human disturbances impedes the ability to diagnose and respond appropriately to these threats., (© 2023 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Caribbean reefs have experienced unprecedented changes in the past four decades. Of great concern is the perceived widespread shift from coral to macroalgal dominance and the question of whether it represents a new, stable equilibrium for coral-reef communities. The primary causes of the shift-grazing pressure (top-down), nutrient loading (bottom-up) or direct coral mortality (side-in)-still remain somewhat controversial in the coral-reef literature. We have attempted to tease out the relative importance of each of these causes. Four insights emerge from our analysis of an early regional dataset of information on the benthic composition of Caribbean reefs spanning the years 1977-2001. First, although three-quarters of reef sites have experienced coral declines concomitant with macroalgal increases, fewer than 10% of the more than 200 sites studied were dominated by macroalgae in 2001, by even the most conservative definition of dominance. Using relative dominance as the threshold, a total of 49 coral-to-macroalgae shifts were detected. This total represents ~35% of all sites that were dominated by coral at the start of their monitoring periods. Four shifts (8.2%) occurred because of coral loss with no change in macroalgal cover, 15 (30.6%) occurred because of macroalgal gain without coral loss, and 30 (61.2%) occurred owing to concomitant coral decline and macroalgal increase. Second, the timing of shifts at the regional scale is most consistent with the side-in model of reef degradation, which invokes coral mortality as a precursor to macroalgal takeover, because more shifts occurred after regional coral-mortality events than expected by chance. Third, instantaneous observations taken at the start and end of the time-series for individual sites showed these reefs existed along a continuum of coral and macroalgal cover. The continuous, broadly negative relationship between coral and macroalgal cover suggests that in some cases coral-to-macroalgae phase shifts may be reversed by removing sources of perturbation or restoring critical components such as the herbivorous sea urchin Diadema antillarum to the system. The five instances in which macroalgal dominance was reversed corroborate the conclusion that macroalgal dominance is not a stable, alternative community state as has been commonly assumed. Fourth, the fact that the loss in regional coral cover and concomitant changes to the benthic community are related to punctuated, discrete events with known causes (i.e. coral disease and bleaching), lends credence to the hypothesis that coral reefs of the Caribbean have been under assault from climate-change-related maladies since the 1970s., (© 2020 Elsevier Ltd. All rights reserved.)

Temperature variability, habitat, coral communities, and fishing intensity are important factors influencing coral responses to climate change. Consequently, chronic and acute sea-surface temperatures (SSTs) and their interactions with habitat and fishing were studied along the East African coast (~400km) by evaluating changes over a ~25-year period in two major reef habitats-island and fringing reefs. These habitats had similar mean and standard deviation temperature measurements but differed in that islands had lower ocean heights and flatter and less right-skewed temperature distributions than fringing reefs. These patterns arise because islands are exposed to deep offshore water passing through deep channels while being protected from the open ocean storms and the strong inter-annual current temperature variability. Within these two seascapes, coral communities are shaped by population responses to the variable temperature distributions as determined by the taxa's associations with the competitive-stress-ruderal (CSR) life history groups. For example, competitive taxa were more abundant where temperature distributions were flat and lacked frequent warm water anomalies. In contrast, ruderal, weedy, and generalist taxa were more common where temperature distributions were centralized, standard deviations high, and warm water anomalies more frequent. Finally, stress-resistant taxa were more common in reefs with high temperature skew but flatter temperature distributions. The rare 1998 thermal anomaly impacted and disturbed the ruderal and stressed reef more than the competitive communities. Ruderal became more similar to stressed communities while the stressed community moved further from the mean before recovering towards the competitive community. Competitive taxa were more common on islands and the deeper fringing reef sites while ruderal were dominant in shallow fringing reef lagoons. Over time, islands were less disturbed than fringing reefs and maintained the highest coral cover, numbers of taxa, and most competitive or space-occupying taxa. However, some island reefs with a history of dynamite fishing aligned with the stress-resistant communities over the full study period. Compared to the in situ SST gauges at the study site, temperature proxies with global coverage were often good at estimating mean and standard deviations of the SSTs but much poorer at estimating the shape of the temperature distributions that reflect chronic and acute stress, as reflected by kurtosis and skewness metrics. Given that these stress variables were critical for understanding the impacts of rare climate disturbances, global climate models that use mean conditions are likely to be poor predictors of future impacts on corals, particularly their species and life history composition. Better predictions should be possible if appropriate chronic and acute stress metrics and their proxies are identified and used., (© 2020 Elsevier Ltd. All rights reserved.)

Without drastic efforts to reduce carbon emissions and mitigate globalized stressors, tropical coral reefs are in jeopardy. Strategic conservation and management requires identification of the environmental and socioeconomic factors driving the persistence of scleractinian coral assemblages-the foundation species of coral reef ecosystems. Here, we compiled coral abundance data from 2,584 Indo-Pacific reefs to evaluate the influence of 21 climate, social and environmental drivers on the ecology of reef coral assemblages. Higher abundances of framework-building corals were typically associated with: weaker thermal disturbances and longer intervals for potential recovery; slower human population growth; reduced access by human settlements and markets; and less nearby agriculture. We therefore propose a framework of three management strategies (protect, recover or transform) by considering: (1) if reefs were above or below a proposed threshold of >10% cover of the coral taxa important for structural complexity and carbonate production; and (2) reef exposure to severe thermal stress during the 2014-2017 global coral bleaching event. Our findings can guide urgent management efforts for coral reefs, by identifying key threats across multiple scales and strategic policy priorities that might sustain a network of functioning reefs in the Indo-Pacific to avoid ecosystem collapse.

Resilience underpins the sustainability of both ecological and social systems. Extensive loss of reef corals following recent mass bleaching events have challenged the notion that support of system resilience is a viable reef management strategy. While resilience-based management (RBM) cannot prevent the damaging effects of major disturbances, such as mass bleaching events, it can support natural processes that promote resistance and recovery. Here, we review the potential of RBM to help sustain coral reefs in the 21st century. We explore the scope for supporting resilience through existing management approaches and emerging technologies and discuss their opportunities and limitations in a changing climate. We argue that for RBM to be effective in a changing world, reef management strategies need to involve both existing and new interventions that together reduce stress, support the fitness of populations and species, and help people and economies to adapt to a highly altered ecosystem., (Copyright © 2018 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Coral bleaching is the detrimental expulsion of algal symbionts from their cnidarian hosts, and predominantly occurs when corals are exposed to thermal stress. The incidence and severity of bleaching is often spatially heterogeneous within reef-scales (<1 km), and is therefore not predictable using conventional remote sensing products. Here, we systematically assess the relationship between in situ measurements of 20 environmental variables, along with seven remotely sensed SST thermal stress metrics, and 81 observed bleaching events at coral reef locations spanning five major reef regions globally. We find that high-frequency temperature variability (i.e., daily temperature range) was the most influential factor in predicting bleaching prevalence and had a mitigating effect, such that a 1 °C increase in daily temperature range would reduce the odds of more severe bleaching by a factor of 33. Our findings suggest that reefs with greater high-frequency temperature variability may represent particularly important opportunities to conserve coral ecosystems against the major threat posed by warming ocean temperatures.

The reduction in coral cover on many contemporary tropical reefs suggests a different set of coral community assemblages will dominate future reefs. To evaluate the capacity of reef corals to persist over various time scales, we examined coral community dynamics in contemporary, fossil, and simulated future coral reef ecosystems. Based on studies between 1987 and 2012 at two locations in the Caribbean, and between 1981 and 2013 at five locations in the Indo-Pacific, we show that many coral genera declined in abundance, some showed no change in abundance, and a few coral genera increased in abundance. Whether the abundance of a genus declined, increased, or was conserved, was independent of coral family. An analysis of fossil-reef communities in the Caribbean revealed changes in numerical dominance and relative abundances of coral genera, and demonstrated that neither dominance nor taxon was associated with persistence. As coral family was a poor predictor of performance on contemporary reefs, a trait-based, dynamic, multi-patch model was developed to explore the phenotypic basis of ecological performance in a warmer future. Sensitivity analyses revealed that upon exposure to thermal stress, thermal tolerance, growth rate, and longevity were the most important predictors of coral persistence. Together, our results underscore the high variation in the rates and direction of change in coral abundances on contemporary and fossil reefs. Given this variation, it remains possible that coral reefs will be populated by a subset of the present coral fauna in a future that is warmer than the recent past.

Coral reefs are biodiverse ecosystems structured by abiotic and biotic factors operating across many spatial scales. Regional-scale interactions between climate change, biogeography and fisheries management remain poorly understood. Here, we evaluated large-scale patterns of coral communities in the western Indian Ocean after a major coral bleaching event in 1998. We surveyed 291 coral reef sites in 11 countries and over 30° of latitude between 2004 and 2011 to evaluate variations in coral communities post 1998 across gradients in latitude, mainland-island geography and fisheries management. We used linear mixed-effect hierarchical models to assess total coral cover, the abundance of four major coral families (acroporids, faviids, pocilloporids and poritiids), coral genus richness and diversity, and the bleaching susceptibility of the coral communities. We found strong latitudinal and geographic gradients in coral community structure and composition that supports the presence of a high coral cover and diversity area that harbours temperature-sensitive taxa in the northern Mozambique Channel between Tanzania, northern Mozambique and northern Madagascar. Coral communities in the more northern latitudes of Kenya, Seychelles and the Maldives were generally composed of fewer bleaching-tolerant coral taxa and with reduced richness and diversity. There was also evidence for continued declines in the abundance of temperature-sensitive taxa and community change after 2004. While there are limitations of our regional dataset in terms of spatial and temporal replication, these patterns suggest that large-scale interactions between biogeographic factors and strong temperature anomalies influence coral communities while smaller-scale factors, such as the effect of fisheries closures, were weak. The northern Mozambique Channel, while not immune to temperature disturbances, shows continued signs of resistance to climate disturbances and remains a priority for future regional conservation and management actions.

This study provides a descriptive analysis of the North Male, Maldives seven years after the 1998 bleaching disturbance to determine the state of the coral community composition, the recruitment community, evidence for recovery, and adaptation to thermal stress. Overall, hard coral cover recovered at a rate commonly reported in the literature but with high spatial variability and shifts in taxonomic composition. Massive Porites, Pavona, Synarea, and Goniopora were unusually common in both the recruit and adult communities. Coral recruitment was low and some coral taxa, namely Tubipora, Seriatopora, and Stylophora, were rarer than expected. A study of the bleaching response to a thermal anomaly in 2005 indicated that some taxa, including Leptoria, Platygyra, Favites, Fungia, Hydnophora, and Galaxea astreata, bleached as predicted while others, including Acropora, Pocillopora, branching Porites, Montipora, Stylophora, and Alveopora, bleached less than predicted. This indicates variable-adaptation potentials among the taxa and considerable potential for ecological reorganization of the coral community., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

There is an increasing need to evaluate the links between the social and ecological dimensions of human vulnerability to climate change. We use an empirical case study of 12 coastal communities and associated coral reefs in Kenya to assess and compare five key ecological and social components of the vulnerability of coastal social-ecological systems to temperature induced coral mortality [specifically: 1) environmental exposure; 2) ecological sensitivity; 3) ecological recovery potential; 4) social sensitivity; and 5) social adaptive capacity]. We examined whether ecological components of vulnerability varied between government operated no-take marine reserves, community-based reserves, and openly fished areas. Overall, fished sites were marginally more vulnerable than community-based and government marine reserves. Social sensitivity was indicated by the occupational composition of each community, including the importance of fishing relative to other occupations, as well as the susceptibility of different fishing gears to the effects of coral bleaching on target fish species. Key components of social adaptive capacity varied considerably between the communities. Together, these results show that different communities have relative strengths and weaknesses in terms of social-ecological vulnerability to climate change.

Climate change is reshaping biological communities against a background of existing human pressure. Evaluating the impacts of multiple stressors on community dynamics can be particularly challenging in species-rich ecosystems, such as coral reefs. Here, we investigate whether life-history strategies and cotolerance to different stressors can predict community responses to fishing and temperature-driven bleaching using a 20-year time series of coral assemblages in Kenya. We found that the initial life-history composition of coral taxa largely determined the impacts of bleaching and coral loss. Prior to the 1998 bleaching event, coral assemblages within no-take marine reserves were composed of three distinct life histories - competitive, stress-tolerant and weedy- and exhibited strong declines following bleaching with limited subsequent recovery. In contrast, fished reefs had lower coral cover, fewer genera and were composed of stress-tolerant and weedy corals that were less affected by bleaching over the long term. Despite these general patterns, we found limited evidence for cotolerance as coral genera and life histories were variable in their sensitivities to fishing and bleaching. Overall, fishing and bleaching have reduced coral diversity and led to altered coral communities of 'survivor' species with stress-tolerant and weedy life histories. Our findings are consistent with expectations that climate change interacting with existing human pressure will result in the loss of coral diversity and critical reef habitat., (© 2013 Blackwell Publishing Ltd.)

Classifying the biological traits of organisms can test conceptual frameworks of life-history strategies and allow for predictions of how different species may respond to environmental disturbances. We apply a trait-based classification approach to a complex and threatened group of species, scleractinian corals. Using hierarchical clustering and random forests analyses, we identify up to four life-history strategies that appear globally consistent across 143 species of reef corals: competitive, weedy, stress-tolerant and generalist taxa, which are primarily separated by colony morphology, growth rate and reproductive mode. Documented shifts towards stress-tolerant, generalist and weedy species in coral reef communities are consistent with the expected responses of these life-history strategies. Our quantitative trait-based approach to classifying life-history strategies is objective, applicable to any taxa and a powerful tool that can be used to evaluate theories of community ecology and predict the impact of environmental and anthropogenic stressors on species assemblages., (© 2012 Blackwell Publishing Ltd/CNRS.)

The risk of global extinction of reef-building coral species is increasing. We evaluated extinction risk using a biological trait-based resiliency index that was compared with Caribbean extinction during the Plio-Pleistocene, and with extinction risk determined by the International Union for Conservation of Nature (IUCN). Through the Plio-Pleistocene, the Caribbean supported more diverse coral assemblages than today and shared considerable overlap with contemporary Indo-Pacific reefs. A clear association was found between extant Plio-Pleistocene coral genera and our positive resilience scores. Regional extinction in the past and vulnerability in the present suggests that Pocillopora, Stylophora and foliose Pavona are among the most susceptible taxa to local and regional isolation. These same taxa were among the most abundant corals in the Caribbean Pliocene. Therefore, a widespread distribution did not equate with immunity to regional extinction. The strong relationship between past and present vulnerability suggests that regional extinction events are trait-based and not merely random episodes. We found several inconsistencies between our data and the IUCN scores, which suggest a need to critically re-examine what constitutes coral vulnerability.

In an effort to deliver better outcomes for people and the ecosystems they depend on, many governments and civil society groups are engaging natural resource users in collaborative management arrangements (frequently called comanagement). However, there are few empirical studies demonstrating the social and institutional conditions conducive to successful comanagement outcomes, especially in small-scale fisheries. Here, we evaluate 42 comanagement arrangements across five countries and show that: (i) comanagement is largely successful at meeting social and ecological goals; (ii) comanagement tends to benefit wealthier resource users; (iii) resource overexploitation is most strongly influenced by market access and users' dependence on resources; and (iv) institutional characteristics strongly influence livelihood and compliance outcomes, yet have little effect on ecological conditions.

Managing coral reefs for resilience to climate change is a popular concept but has been difficult to implement because the empirical scientific evidence has either not been evaluated or is sometimes unsupportive of theory, which leads to uncertainty when considering methods and identifying priority reefs. We asked experts and reviewed the scientific literature for guidance on the multiple physical and biological factors that affect the ability of coral reefs to resist and recover from climate disturbance. Eleven key factors to inform decisions based on scaling scientific evidence and the achievability of quantifying the factors were identified. Factors important to resistance and recovery, which are important components of resilience, were not strongly related, and should be assessed independently. The abundance of resistant (heat-tolerant) coral species and past temperature variability were perceived to provide the greatest resistance to climate change, while coral recruitment rates, and macroalgae abundance were most influential in the recovery process. Based on the 11 key factors, we tested an evidence-based framework for climate change resilience in an Indonesian marine protected area. The results suggest our evidence-weighted framework improved upon existing un-weighted methods in terms of characterizing resilience and distinguishing priority sites. The evaluation supports the concept that, despite high ecological complexity, relatively few strong variables can be important in influencing ecosystem dynamics. This is the first rigorous assessment of factors promoting coral reef resilience based on their perceived importance, empirical evidence, and feasibility of measurement. There were few differences between scientists' perceptions of factor importance and the scientific evidence found in journal publications but more before and after impact studies will be required to fully test the validity of all the factors. The methods here will increase the feasibility and defensibility of including key resilience metrics in evaluations of coral reefs, as well as reduce costs. Adaptation, marine protected areas, priority setting, resistance, recovery.

Background: The decline of coral reefs globally underscores the need for a spatial assessment of their exposure to multiple environmental stressors to estimate vulnerability and evaluate potential counter-measures., Methodology/principal Findings: This study combined global spatial gradients of coral exposure to radiation stress factors (temperature, UV light and doldrums), stress-reinforcing factors (sedimentation and eutrophication), and stress-reducing factors (temperature variability and tidal amplitude) to produce a global map of coral exposure and identify areas where exposure depends on factors that can be locally managed. A systems analytical approach was used to define interactions between radiation stress variables, stress reinforcing variables and stress reducing variables. Fuzzy logic and spatial ordinations were employed to quantify coral exposure to these stressors. Globally, corals are exposed to radiation and reinforcing stress, albeit with high spatial variability within regions. Based on ordination of exposure grades, regions group into two clusters. The first cluster was composed of severely exposed regions with high radiation and low reducing stress scores (South East Asia, Micronesia, Eastern Pacific and the central Indian Ocean) or alternatively high reinforcing stress scores (the Middle East and the Western Australia). The second cluster was composed of moderately to highly exposed regions with moderate to high scores in both radiation and reducing factors (Caribbean, Great Barrier Reef (GBR), Central Pacific, Polynesia and the western Indian Ocean) where the GBR was strongly associated with reinforcing stress., Conclusions/significance: Despite radiation stress being the most dominant stressor, the exposure of coral reefs could be reduced by locally managing chronic human impacts that act to reinforce radiation stress. Future research and management efforts should focus on incorporating the factors that mitigate the effect of coral stressors until long-term carbon reductions are achieved through global negotiations.

Many of the world's coral reefs suffered high coral mortality during the 1998 ENSO, with the highest mortality in the western Indian Ocean (WIO). A meta-analysis of field data on change in coral cover across the 1998 ENSO event was conducted for 36 major reef areas in the WIO, and relationship of the change with the historical sea-surface temperature (SST) variability investigated. WIO reefs were categorized into three major SST groups of differing coral cover change. Cover change was negatively associated with standard deviation (SD) SST until about SD 2.3, with increasing flatness of the SST frequency distributions. It increased with further increase in SD as the SST distributions became strongly bimodal in the Arabian/Persian Gulf area. The study indicates that environmental resistance/tolerance to extreme anomalous events could be predicted and management priorities directed accordingly for a warmer and more variable future climate., (Copyright 2010 Elsevier Ltd. All rights reserved.)

Coral reefs have emerged as one of the ecosystems most vulnerable to climate variation and change. While the contribution of a warming climate to the loss of live coral cover has been well documented across large spatial and temporal scales, the associated effects on fish have not. Here, we respond to recent and repeated calls to assess the importance of local management in conserving coral reefs in the context of global climate change. Such information is important, as coral reef fish assemblages are the most species dense vertebrate communities on earth, contributing critical ecosystem functions and providing crucial ecosystem services to human societies in tropical countries. Our assessment of the impacts of the 1998 mass bleaching event on coral cover, reef structural complexity, and reef associated fishes spans 7 countries, 66 sites and 26 degrees of latitude in the Indian Ocean. Using Bayesian meta-analysis we show that changes in the size structure, diversity and trophic composition of the reef fish community have followed coral declines. Although the ocean scale integrity of these coral reef ecosystems has been lost, it is positive to see the effects are spatially variable at multiple scales, with impacts and vulnerability affected by geography but not management regime. Existing no-take marine protected areas still support high biomass of fish, however they had no positive affect on the ecosystem response to large-scale disturbance. This suggests a need for future conservation and management efforts to identify and protect regional refugia, which should be integrated into existing management frameworks and combined with policies to improve system-wide resilience to climate variation and change.

The hypothesis that herbivory is higher in areas without fishing and will increase the rate at which hard coral communities return to pre-disturbance conditions was tested in and out of the marine protected areas (MPA) of Kenya after the 1998 El Niño Southern Oscillation (ENSO). Herbivory was estimated by assay and biomass methods, and both methods indicated higher herbivory in fishery closures. Despite higher herbivory, the effect of the ENSO disturbance was larger within these closures, with reefs undergoing a temporary transition from dominance by hard and soft coral to a temporary dominance of turf and erect algae that ended in the dominance of calcifying algae, massive Porites, Pocillopora and a few faviids six years after the disturbance. The fished reefs changed the least but had a greater cover of turf and erect algae and sponge shortly after the disturbance. Higher herbivory in the fishery closures reduced the abundance and persistence of herbivore-susceptible erect algae and created space and appropriate substratum for recruiting corals. Nonetheless, other post-settlement processes may have had strong influences such that annual rates of coral recovery were low ( approximately 2%) and not different between the management regimes. Recovery, as defined as and measured by the return to pre-disturbance coral cover and the dominant taxa, was slower in fishery closures than unmanaged reefs.

Nitrogen and phosphorous fertilizers were used to determine their short-term summer effects on algal colonization, abundance, and species composition in moderate herbivory treatments. Secondary succession of algae on coral skeletons was examined in four treatments: an untreated control, a pure phosphate fertilizer, a pure nitrogen fertilizer, and an equal mix of the two fertilizers. Turf algae cover was the only measure of algae abundance to respond significantly to fertilization. Turf cover was three times higher in treatments with added nitrogen when compared with the pure phosphorus treatment. These turfs were dominated by green and cyanobacteria taxa, namely Enteromorpha prolifera, Lyngbya confervoides, and two species of Cladophora. The phosphate treatment was dominated by encrusting corallines and the cyanobacteria L. confervoides, while the controls had the highest cover of frondose brown algae, namely Padina sanctae-crucis and two species of Dictyota. Results indicate that turf algae were co-limited by nitrogen and phosphorus but enrichment appeared to inhibit brown frondose algae that currently dominate these reefs. Number of species was lowest on the pure phosphorus and nitrogen treatments, highest in the controls and intermediate in the mixed treatments, which suggests that diversity is reduced most by an imbalanced nutrient ratio.

Identifying the rates of recovery of fish in no-take areas is fundamental to designing protected area networks, managing fisheries, estimating yields, identifying ecological interactions, and informing stakeholders about the outcomes of this management. Here we study the recovery of coral reef fishes through 37 years of protection using a space-for-time chronosequence of four marine national parks in Kenya. Using AIC model selection techniques, we assessed recovery trends using five ecologically meaningful production models: asymptotic, Ricker, logistic, linear, and exponential. There were clear recovery trends with time for species richness, total and size class density, and wet masses at the level of the taxonomic family. Species richness recovered rapidly to an asymptote at 10 years. The two main herbivorous families displayed differing responses to protection, scarids recovering rapidly, but then exhibiting some decline while acanthurids recovered more slowly and steadily throughout the study. Recovery of the two invertebrate-eating groups suggested competitive interactions over resources, with the labrids recovering more rapidly before a decline and the balistids demonstrating a slower logistic recovery. Remaining families displayed differing trends with time, with a general pattern of decline in smaller size classes or small-bodied species after an initial recovery, which suggests that some species- and size-related competitive and predatory control occurs in older closures. There appears to be an ecological succession of dominance with an initial rapid rise in labrids and scarids, followed by a slower rise in balistids and acanthurids, an associated decline in sea urchins, and an ultimate dominance in calcifying algae. Our results indicate that the unfished "equilibrium" biomass of the fish assemblage > 10 cm is 1100-1200 kg/ha, but these small parks (< 10 km2) are likely to underestimate pre-human influence values due to edge effects and the rarity of taxa with large area requirement, such as apex predators, including sharks.

Marine protected areas (MPAs) have been widely adopted as the leading tool for coral-reef conservation, but resource users seldom accept them , and many have failed to produce tangible conservation benefits [3]. Few studies have objectively and simultaneously examined the types of MPAs that are most effective in conserving reef resources and the socioeconomic factors responsible for effective conservation [4-6]. We simultaneously explored measures of reef and socioeconomic conservation success at four national parks, four comanaged reserves, and three traditionally managed areas in Indonesia and Papua New Guinea. Underwater visual censuses of key ecological indicators [7, 8] revealed that the average size and biomass of fishes were higher in all areas under traditional management and at one comanaged reserve when compared to nearby unmanaged areas. Socioeconomic assessments [6, 9, 10] revealed that this "effective conservation" was positively related to compliance, visibility of the reserve, and length of time the management had been in place but negatively related to market integration, wealth, and village population size. We suggest that in cases where the resources for enforcement are lacking, management regimes that are designed to meet community goals can achieve greater compliance and subsequent conservation success than regimes designed primarily for biodiversity conservation.

We studied the responses of algae, corals, and small fish to elevated inorganic fertilizer, organic matter, and their combination over a 49-day summer period in cages that simulated the coral reef in the remote Glovers reef atoll, Belize. The addition of organic matter reduced while fertilization had no effect on the numbers of herbivorous damsel and parrotfishes. All measures of algal biomass were influenced by fertilization. The combined inorganic and organic enrichment produced the highest algal biomass, which is most likely due to the combined effect of higher nutrients and lower herbivory. The cover of turf and total algae were influenced by all treatments and their interactions and most strongly and positively influenced by fertilization followed by organic matter and the combination of organic matter and inorganic fertilizer. The inorganic and combined treatments were both dominated by two turf algae, Enteromorpha prolifera and Digenia simplex, while the nonfertilized treatments were dominated by brown frondose algae Lobophora variegata, Padina sanctae, and Dictyota cervicornis. The organic matter treatment had greater cover of P. sanctae and D. cervicornis than the untreated control, which was dominated by Lobophora variegata, also the dominant algae on the nearby patch reefs. Crustose corallines grew slowly ( approximately 2.5 mm/49 days) and were not influenced by the treatments when grown on vertical surfaces but decreased on horizontal coral plates in the combined organic matter and fertilization treatment. No mortality occurred for the two coral species that were added to the cages. Porites furcata darkened in the fertilized cages while there was a mix of paling and darkening for a small amount of the coral tissue of Diploria labyrinthiformes. Inorganic fertilization stimulates small filamentous turf algae and Symbiodinium living in coral but inhibits brown frondose algae. Organic matter inhibits small herbivorous fish, L. variegata, and encrusting coralline algae when growing on horizontal surfaces.

The long-term response of coral reefs to climate change depends on the ability of reef-building coral symbioses to adapt or acclimatize to warmer temperatures, but there has been no direct evidence that such a response can occur. Here we show that corals containing unusual algal symbionts that are thermally tolerant and commonly associated with high-temperature environments are much more abundant on reefs that have been severely affected by recent climate change. This adaptive shift in symbiont communities indicates that these devastated reefs could be more resistant to future thermal stress, resulting in significantly longer extinction times for surviving corals than had been previously assumed.

We compared the bleaching and mortality response (BMI) of 19 common scleractinian corals to an anomalous warm-water event in 1998 to determine the degree of variation between depths, sites, and regions. Mombasa corals experienced a greater temperature anomaly than those on the Great Barrier Reef (GBR) sites and this was reflected in the greater BMI response of most taxa. Comparing coral taxa in different sites at the same depth produced high correlation coefficients in the bleaching response in Kenya at 2 m (r=0.86) and GBR at 6 m depth sites (r=0.80) but less in the GBR for shallow 2 m sites (r=0.49). The pattern of taxa susceptibility was remarkably consistent between the regions. Coral taxa explained 52% of the variation in the response of colonies to bleaching between these two regions (Kenya BMI=0.90 GBR BMI+26; F(1,19) - 18.3; p < 0.001; r2 = 0.52). Stylophora and Pocillopora were consistently susceptible while Cyphastrea, Goniopora Galaxea and Pavona were resistant in both regions. Three taxa behaved differently between the two regions; Acropora, and branching Porites were both moderately affected on the GBR but were highly affected in Kenya while the opposite was true for Pavona. These results suggest that a colonies response to bleaching is phylogenetically constrained, emphasizing the importance of features of the host's physiology or morphology in determining the response to thermal stress.

The proliferation of algae on disturbed coral reefs has often been attributed to (1) a loss of large-bodied herbivorous fishes, (2) increases in sea water nutrient concentrations, particularly phosphorus, and (3) a loss of hard coral cover or a combination of these and other factors. We performed replicated small-scale caging experiments in the offshore lagoon of Glovers Reef atoll, Belize where three treatments had closed-top (no large-bodied herbivores) and one treatment had open-top cages (grazing by large-bodied herbivores). Closed-top treatments simulated a reduced-herbivory situation, excluding large fishes but including small herbivorous fishes such as damselfishes and small parrotfishes. Treatments in the closed-top cages included the addition of high phosphorus fertilizer, live branches of Acropora cervicornis and a third unmanipulated control treatment. Colonization, algal biomass and species composition on dead A. palmata "plates" were studied weekly for 50 days in each of the four treatments. Fertilization doubled the concentration of phosphorus from 0.35 to 0.77 microM. Closed-top cages, particularly the fertilizer and A. cervicornis additions, attracted more small-bodied parrotfish and damselfish than the open-top cages such that there was moderate levels of herbivory in closed-top cages. The open-top cages did, however, have a higher abundance of the chemically and morphologically defended erect algal species including Caulerpa cupressoides, Laurencia obtusa, Dictyota menstrualis and Lobophora variegata. The most herbivore-resistant calcareous green algae (i.e. Halimeda) were, however, uncommon in all treatments. Algal biomass increased and fluctuated simultaneously in all treatments over time, but algal biomass, as measured by wet, dry and decalcified weight, did not differ greatly between the treatments with only marginally higher biomass (p < 0.06) in the fertilized compared to open-top cages. Algal species composition was influenced by all treatments with a maximum between-treatment Bray-Curtis similarity of only 29%. The fertilized cages showed rapid colonization by a mixed turf community largely composed of the filamentous brown (Hincksia mitchelliae) and green (Enteromorpha prolifera) species. Algal cover in the fertilized cages leveled at 80% after 20 days compared to less than 50% in the other treatments. There was no evidence that A. cervicornis suppressed algal colonization compared to the unmanipulated controls. Instead, the herbivore susceptible Padina sanctae-crucis was the most abundant algae followed by Jania capillacea in this treatment in contrast to the more chemically defended Dictyota menstrualis that dominated the unmanipulated controls. We conclude that A. cervicornis was not suppressing algae as a group and its loss cannot account for the observed changes in algal abundance in most reefs except for creating space. In contrast, A. cervicornis appears to attract aggressive damselfish that may reduce herbivory by larger herbivores. Phosphorus enrichment can lead to rapid colonization of space by filamentous turf communities but not high biomass and dominance of erect frondose algae within 50 days. Moderate levels of herbivory by large-bodied herbivores promoted moderately herbivore-resistant erect brown and green algae that are commonly reported on disturbed reefs. Consequently, all the studied factors influenced algal communities but seldom as commonly predicted.