Scale dependence of coral reef oases and their environmental correlates.

Identifying relatively intact areas within ecosystems and determining the conditions favoring their existence is necessary for effective management in the context of widespread environmental degradation. In this study, we used 3766 surveys of randomly selected sites in the United States and U.S. Territories to identify the correlates of sites categorized as "oases" (defined as sites with relatively high total coral cover). We used occupancy models to evaluate the influence of 10 environmental predictors on the probability that an area (21.2-km2 cell) would harbor coral oases defined at four spatial extents: cross-basin, basin, region, and subregion. Across all four spatial extents, oases were more likely to occur in habitats with high light attenuation. The influence of the other environmental predictors on the probability of oasis occurrence were less consistent and varied with the scale of observation. Oases were most likely in areas of low human population density, but this effect was evident only at the cross-basin and subregional extents. At the regional and subregional extents oases were more likely where sea-surface temperature was more variable, whereas at the larger spatial extents the opposite was true. By identifying the correlates of oasis occurrence, the model can inform the prioritization of reef areas for management. Areas with biophysical conditions that confer corals with physiological resilience, as well as limited human impacts, likely support coral reef oases across spatial extents. Our approach is widely applicable to the development of conservation strategies to protect biodiversity and ecosystems in an era of magnified human disturbance.

Bright spots among the world's coral reefs.

Ongoing declines in the structure and function of the world's coral reefs require novel approaches to sustain these ecosystems and the millions of people who depend on them3. A presently unexplored approach that draws on theory and practice in human health and rural development is to systematically identify and learn from the 'outliers'­places where ecosystems are substantially better ('bright spots') or worse ('dark spots') than expected, given the environmental conditions and socioeconomic drivers they are exposed to. Here we compile data from more than 2,500 reefs worldwide and develop a Bayesian hierarchical model to generate expectations of how standing stocks of reef fish biomass are related to 18 socioeconomic drivers and environmental conditions. We identify 15 bright spots and 35 dark spots among our global survey of coral reefs, defined as sites that have biomass levels more than two standard deviations from expectations. Importantly, bright spots are not simply comprised of remote areas with low fishing pressure; they include localities where human populations and use of ecosystem resources is high, potentially providing insights into how communities have successfully confronted strong drivers of change. Conversely, dark spots are not necessarily the sites with the lowest absolute biomass and even include some remote, uninhabited locations often considered near pristine6. We surveyed local experts about social, institutional, and environmental conditions at these sites to reveal that bright spots are characterized by strong sociocultural institutions such as customary taboos and marine tenure, high levels of local engagement in management, high dependence on marine resources, and beneficial environmental conditions such as deep-water refuges. Alternatively, dark spots are characterized by intensive capture and storage technology and a recent history of environmental shocks. Our results suggest that investments in strengthening fisheries governance, particularly aspects such as participation and property rights, could facilitate innovative conservation actions that help communities defy expectations of global reef degradation.

Recovery potential of the world's coral reef fishes.

Continuing degradation of coral reef ecosystems has generated substantial interest in how management can support reef resilience. Fishing is the primary source of diminished reef function globally, leading to widespread calls for additional marine reserves to recover fish biomass and restore key ecosystem functions. Yet there are no established baselines for determining when these conservation objectives have been met or whether alternative management strategies provide similar ecosystem benefits. Here we establish empirical conservation benchmarks and fish biomass recovery timelines against which coral reefs can be assessed and managed by studying the recovery potential of more than 800 coral reefs along an exploitation gradient. We show that resident reef fish biomass in the absence of fishing (B0) averages ∼1,000 kg ha(-1), and that the vast majority (83%) of fished reefs are missing more than half their expected biomass, with severe consequences for key ecosystem functions such as predation. Given protection from fishing, reef fish biomass has the potential to recover within 35 years on average and less than 60 years when heavily depleted. Notably, alternative fisheries restrictions are largely (64%) successful at maintaining biomass above 50% of B0, sustaining key functions such as herbivory. Our results demonstrate that crucial ecosystem functions can be maintained through a range of fisheries restrictions, allowing coral reef managers to develop recovery plans that meet conservation and livelihood objectives in areas where marine reserves are not socially or politically feasible solutions.

Gravity of human impacts mediates coral reef conservation gains.

Coral reefs provide ecosystem goods and services for millions of people in the tropics, but reef conditions are declining worldwide. Effective solutions to the crisis facing coral reefs depend in part on understanding the context under which different types of conservation benefits can be maximized. Our global analysis of nearly 1,800 tropical reefs reveals how the intensity of human impacts in the surrounding seascape, measured as a function of human population size and accessibility to reefs ("gravity"), diminishes the effectiveness of marine reserves at sustaining reef fish biomass and the presence of top predators, even where compliance with reserve rules is high. Critically, fish biomass in high-compliance marine reserves located where human impacts were intensive tended to be less than a quarter that of reserves where human impacts were low. Similarly, the probability of encountering top predators on reefs with high human impacts was close to zero, even in high-compliance marine reserves. However, we find that the relative difference between openly fished sites and reserves (what we refer to as conservation gains) are highest for fish biomass (excluding predators) where human impacts are moderate and for top predators where human impacts are low. Our results illustrate critical ecological trade-offs in meeting key conservation objectives: reserves placed where there are moderate-to-high human impacts can provide substantial conservation gains for fish biomass, yet they are unlikely to support key ecosystem functions like higher-order predation, which is more prevalent in reserve locations with low human impacts.

Parsing human and biophysical drivers of coral reef regimes.

Coral reefs worldwide face unprecedented cumulative anthropogenic effects of interacting local human pressures, global climate change and distal social processes. Reefs are also bound by the natural biophysical environment within which they exist. In this context, a key challenge for effective management is understanding how anthropogenic and biophysical conditions interact to drive distinct coral reef configurations. Here, we use machine learning to conduct explanatory predictions on reef ecosystems defined by both fish and benthic communities. Drawing on the most spatially extensive dataset available across the Hawaiian archipelago-20 anthropogenic and biophysical predictors over 620 survey sites-we model the occurrence of four distinct reef regimes and provide a novel approach to quantify the relative influence of human and environmental variables in shaping reef ecosystems. Our findings highlight the nuances of what underpins different coral reef regimes, the overwhelming importance of biophysical predictors and how a reef's natural setting may either expand or narrow the opportunity space for management interventions. The methods developed through this study can help inform reef practitioners and hold promises for replication across a broad range of ecosystems.

Seascape models reveal places to focus coastal fisheries management.

To design effective marine reserves and support fisheries, more information on fishing patterns and impacts for targeted species is needed, as well as better understanding of their key habitats. However, fishing impacts vary geographically and are difficult to disentangle from other factors that influence targeted fish distributions. We developed a set of fishing effort and habitat layers at high resolution and employed machine learning techniques to create regional-scale seascape models and predictive maps of biomass and body length of targeted reef fishes for the main Hawaiian Islands. Spatial patterns of fishing effort were shown to be highly variable and seascape models indicated a low threshold beyond which targeted fish assemblages were severely impacted. Topographic complexity, exposure, depth, and wave power were identified as key habitat variables that influenced targeted fish distributions and defined productive habitats for reef fisheries. High targeted reef fish biomass and body length were found in areas not easily accessed by humans, while model predictions when fishing effort was set to zero showed these high values to be more widely dispersed among suitable habitats. By comparing current targeted fish distributions with those predicted when fishing effort was removed, areas with high recovery potential on each island were revealed, with average biomass recovery of 517% and mean body length increases of 59% on Oahu, the most heavily fished island. Spatial protection of these areas would aid recovery of nearshore coral reef fisheries.

Environmental conditions and herbivore biomass determine coral reef benthic community composition: implications for quantitative baselines.

Our ability to understand natural constraints on coral reef benthic communities requires quantitative assessment of the relative strengths of abiotic and biotic processes across large spatial scales. Here, we combine underwater images, visual censuses and remote sensing data for 1566 sites across 34 islands spanning the central-western Pacific Ocean, to empirically assess the relative roles of abiotic and grazing processes in determining the prevalence of calcifying organisms and fleshy algae on coral reefs. We used regression trees to identify the major predictors of benthic composition and to test whether anthropogenic stress at inhabited islands decouples natural relationships. We show that sea surface temperature, wave energy, oceanic productivity and aragonite saturation strongly influence benthic community composition; overlooking these factors may bias expectations of calcified reef states. Maintenance of grazing biomass above a relatively low threshold (~ 10-20 kg ha-1) may also prevent transitions to algal-dominated states, providing a tangible management target for rebuilding overexploited herbivore populations. Biophysical relationships did not decouple at inhabited islands, indicating that abiotic influences remain important macroscale processes, even at chronically disturbed reefs. However, spatial autocorrelation among inhabited reefs was substantial and exceeded abiotic and grazing influences, suggesting that natural constraints on reef benthos were superseded by unmeasured anthropogenic impacts. Evidence of strong abiotic influences on reef benthic communities underscores their importance in specifying quantitative targets for coral reef management and restoration that are realistic within the context of local conditions.

Fishing degrades size structure of coral reef fish communities.

Fishing pressure on coral reef ecosystems has been frequently linked to reductions of large fishes and reef fish biomass. Associated impacts on overall community structure are, however, less clear. In size-structured aquatic ecosystems, fishing impacts are commonly quantified using size spectra, which describe the distribution of individual body sizes within a community. We examined the size spectra and biomass of coral reef fish communities at 38 US-affiliated Pacific islands that ranged in human presence from near pristine to human population centers. Size spectra 'steepened' steadily with increasing human population and proximity to market due to a reduction in the relative biomass of large fishes and an increase in the dominance of small fishes. Reef fish biomass was substantially lower on inhabited islands than uninhabited ones, even at inhabited islands with the lowest levels of human presence. We found that on populated islands size spectra exponents decreased (analogous to size spectra steepening) linearly with declining biomass, whereas on uninhabited islands there was no relationship. Size spectra were steeper in regions of low sea surface temperature but were insensitive to variation in other environmental and geomorphic covariates. In contrast, reef fish biomass was highly sensitive to oceanographic conditions, being influenced by both oceanic productivity and sea surface temperature. Our results suggest that community size structure may be a more robust indicator than fish biomass to increasing human presence and that size spectra are reliable indicators of exploitation impacts across regions of different fish community compositions, environmental drivers, and fisheries types. Size-based approaches that link directly to functional properties of fish communities, and are relatively insensitive to abiotic variation across biogeographic regions, offer great potential for developing our understanding of fishing impacts in coral reef ecosystems.

Natural bounds on herbivorous coral reef fishes.

Humans are an increasingly dominant driver of Earth's biological communities, but differentiating human impacts from natural drivers of ecosystem state is crucial. Herbivorous fish play a key role in maintaining coral dominance on coral reefs, and are widely affected by human activities, principally fishing. We assess the relative importance of human and biophysical (habitat and oceanographic) drivers on the biomass of five herbivorous functional groups among 33 islands in the central and western Pacific Ocean. Human impacts were clear for some, but not all, herbivore groups. Biomass of browsers, large excavators, and of all herbivores combined declined rapidly with increasing human population density, whereas grazers, scrapers, and detritivores displayed no relationship. Sea-surface temperature had significant but opposing effects on the biomass of detritivores (positive) and browsers (negative). Similarly, the biomass of scrapers, grazers, and detritivores correlated with habitat structural complexity; however, relationships were group specific. Finally, the biomass of browsers and large excavators was related to island geomorphology, both peaking on low-lying islands and atolls. The substantial variability in herbivore populations explained by natural biophysical drivers highlights the need for locally appropriate management targets on coral reefs.

Investigating functional redundancy versus complementarity in Hawaiian herbivorous coral reef fishes.

Patterns of species resource use provide insight into the functional roles of species and thus their ecological significance within a community. The functional role of herbivorous fishes on coral reefs has been defined through a variety of methods, but from a grazing perspective, less is known about the species-specific preferences of herbivores on different groups of reef algae and the extent of dietary overlap across an herbivore community. Here, we quantified patterns of redundancy and complementarity in a highly diverse community of herbivores at a reef on Maui, Hawaii, USA. First, we tracked fish foraging behavior in situ to record bite rate and type of substrate bitten. Second, we examined gut contents of select herbivorous fishes to determine consumption at a finer scale. Finally, we placed foraging behavior in the context of resource availability to determine how fish selected substrate type. All species predominantly (73-100 %) foraged on turf algae, though there were differences among the types of macroalgae and other substrates bitten. Increased resolution via gut content analysis showed the composition of turf algae consumed by fishes differed across herbivore species. Consideration of foraging behavior by substrate availability revealed 50 % of herbivores selected for turf as opposed to other substrate types, but overall, there were variable foraging portfolios across all species. Through these three methods of investigation, we found higher complementarity among herbivorous fishes than would be revealed using a single metric. These results suggest differences across species in the herbivore "rain of bites" that graze and shape benthic community composition.

Sustainable reference points for multispecies coral reef fisheries.

Sustainably managing fisheries requires regular and reliable evaluation of stock status. However, most multispecies reef fisheries around the globe tend to lack research and monitoring capacity, preventing the estimation of sustainable reference points against which stocks can be assessed. Here, combining fish biomass data for >2000 coral reefs, we estimate site-specific sustainable reference points for coral reef fisheries and use these and available catch estimates to assess the status of global coral reef fish stocks. We reveal that >50% of sites and jurisdictions with available information have stocks of conservation concern, having failed at least one fisheries sustainability benchmark. We quantify the trade-offs between biodiversity, fish length, and ecosystem functions relative to key benchmarks and highlight the ecological benefits of increasing sustainability. Our approach yields multispecies sustainable reference points for coral reef fisheries using environmental conditions, a promising means for enhancing the sustainability of the world's coral reef fisheries.

Re-creating missing population baselines for Pacific reef sharks.

Sharks and other large predators are scarce on most coral reefs, but studies of their historical ecology provide qualitative evidence that predators were once numerous in these ecosystems. Quantifying density of sharks in the absence of humans (baseline) is, however, hindered by a paucity of pertinent time-series data. Recently researchers have used underwater visual surveys, primarily of limited spatial extent or nonstandard design, to infer negative associations between reef shark abundance and human populations. We analyzed data from 1607 towed-diver surveys (>1 ha transects surveyed by observers towed behind a boat) conducted at 46 reefs in the central-western Pacific Ocean, reefs that included some of the world's most pristine coral reefs. Estimates of shark density from towed-diver surveys were substantially lower (<10%) than published estimates from surveys along small transects (<0.02 ha), which is not consistent with inverted biomass pyramids (predator biomass greater than prey biomass) reported by other researchers for pristine reefs. We examined the relation between the density of reef sharks observed in towed-diver surveys and human population in models that accounted for the influence of oceanic primary productivity, sea surface temperature, reef area, and reef physical complexity. We used these models to estimate the density of sharks in the absence of humans. Densities of gray reef sharks (Carcharhinus amblyrhynchos), whitetip reef sharks (Triaenodon obesus), and the group "all reef sharks" increased substantially as human population decreased and as primary productivity and minimum sea surface temperature (or reef area, which was highly correlated with temperature) increased. Simulated baseline densities of reef sharks under the absence of humans were 1.1-2.4/ha for the main Hawaiian Islands, 1.2-2.4/ha for inhabited islands of American Samoa, and 0.9-2.1/ha for inhabited islands in the Mariana Archipelago, which suggests that density of reef sharks has declined to 3-10% of baseline levels in these areas.

Incorporating reef fish avoidance behavior improves accuracy of species distribution models.

Species distribution models (SDMs) are used to interpret and map fish distributions based on habitat variables and other drivers. Reef fish avoidance behavior has been shown to vary in the presence of divers and is primarily driven by spearfishing pressure. Diver avoidance behavior or fish wariness may spatially influence counts and other descriptive measures of fish assemblages. Because fish assemblage metrics are response variables for SDMs, measures of fish wariness may be useful as predictors in SDMs of fishes targeted by spearfishing. We used a diver operated stereo-video system to conduct fish surveys and record minimum approach distance (MAD) of targeted reef fishes inside and outside of two marine reserves on the island of O'ahu in the main Hawaiian Islands. By comparing MAD between sites and management types we tested the assumption that it provides a proxy for fish wariness related to spearfishing pressure. We then compared the accuracy of SDMs which included MAD as a predictor with SDMs that did not. Individual measures of MAD differed between sites though not management types. When included as a predictor, MAD averaged at the transect level greatly improved the accuracy of SDMs of targeted fish biomass.

Meeting fisheries, ecosystem function, and biodiversity goals in a human-dominated world.

The worldwide decline of coral reefs necessitates targeting management solutions that can sustain reefs and the livelihoods of the people who depend on them. However, little is known about the context in which different reef management tools can help to achieve multiple social and ecological goals. Because of nonlinearities in the likelihood of achieving combined fisheries, ecological function, and biodiversity goals along a gradient of human pressure, relatively small changes in the context in which management is implemented could have substantial impacts on whether these goals are likely to be met. Critically, management can provide substantial conservation benefits to most reefs for fisheries and ecological function, but not biodiversity goals, given their degraded state and the levels of human pressure they face.

Effects of SCUBA bubbles on counts of roving piscivores in a large remote marine protected area.

This study examined the effects of SCUBA bubbles on fish counts in underwater visual surveys conducted in the Papahanaumokuakea Marine National Monument (PMNM). Specifically, paired fish surveys were conducted at each survey site, utilizing two different gear types: open-circuit SCUBA (OC) and closed-circuit rebreather (CCR). Bubble exhaust released from the OC equipment is a potential source of bias for in-situ fish observations, as the associated audio and visual disturbances could either attract or repel fishes depending on whether their behavior is more driven by curiosity or caution. The study area, is a large (~1.5 million km2) and extremely remote marine protected area in which the response of coral reef fishes to divers represent natural behavior of naive fishes with little or no previous contact with humans. Historically, surveys conducted on OC in this area have shown an abundance of large roving piscivores and this study set out to determine the extant, if any, the audible and visual disturbances of OC bubbles have. The species typically seen in these prior surveys were Caranx ignobilis, Caranx melampygus, Aprion virescens, and a couple of species of sharks. We found differences in counts for some roving piscivores, including significantly more jacks observed on OC than CCR (Caranx ignobilis 57% more, and Caranx melampygus 113% more). Instance of first encounter, i.e. the time when a fish was first observed during a survey, also varied for some species. Higher numbers of Aprion virescens (p = 0.04), and C. melampygus (p = <0.001) were observed in the first 5-minutes of counts by divers on OC (i.e. when they were using breathing apparatus that produced noises that could be heard over long distances). Although not the focus of the study, we also assessed differences between OC and CCR counts for other groups of fishes. Estimated abundance of benthic damselfish was higher on OC than CCR, and counts of butterflyfish were lower on OC; but there were no significant differences for the other groups considered. This is an important control study that documents the natural responses of coral reef fishes to SCUBA bubbles generated by in-situ surveys.

Estimating nearshore coral reef-associated fisheries production from the main Hawaiian Islands.

Currently, information on nearshore reef-associated fisheries is frequently disparate or incomplete, creating a challenge for effective management. This study utilized an existing non-commercial fishery dataset from Hawai'i, covering the period 2004-13, to estimate a variety of fundamental fishery parameters, including participation, effort, gear use, and catch per unit effort. We then used those data to reconstruct total catches per island. Non-commercial fisheries in this case comprise recreational, subsistence, and cultural harvest, which may be exchanged, but are not sold. By combining those data with reported commercial catch data, we estimated annual catch of nearshore reef-associated fisheries in the main Hawaiian Islands over the study period to be 1,167,758 ± 43,059 kg year-1 (mean ± standard error). Average annual commercial reef fish catch over the same time period-184,911 kg year-1-was 16% of the total catch, but that proportion varied greatly among islands, ranging from 23% on O'ahu to 5% on Moloka'i. These results emphasize the importance of reef fishing in Hawai'i for reasons beyond commerce, such as food security and cultural practice, and highlight the large differences in fishing practices across the Hawaiian Islands.

Combining fish and benthic communities into multiple regimes reveals complex reef dynamics.

Coral reefs worldwide face an uncertain future with many reefs reported to transition from being dominated by corals to macroalgae. However, given the complexity and diversity of the ecosystem, research on how regimes vary spatially and temporally is needed. Reef regimes are most often characterised by their benthic components; however, complex dynamics are associated with losses and gains in both fish and benthic assemblages. To capture this complexity, we synthesised 3,345 surveys from Hawai'i to define reef regimes in terms of both fish and benthic assemblages. Model-based clustering revealed five distinct regimes that varied ecologically, and were spatially heterogeneous by island, depth and exposure. We identified a regime characteristic of a degraded state with low coral cover and fish biomass, one that had low coral but high fish biomass, as well as three other regimes that varied significantly in their ecology but were previously considered a single coral dominated regime. Analyses of time series data reflected complex system dynamics, with multiple transitions among regimes that were a function of both local and global stressors. Coupling fish and benthic communities into reef regimes to capture complex dynamics holds promise for monitoring reef change and guiding ecosystem-based management of coral reefs.

An Assessment of Mobile Predator Populations along Shallow and Mesophotic Depth Gradients in the Hawaiian Archipelago.

Large-bodied coral reef roving predators (sharks, jacks, snappers) are largely considered to be depleted around human population centers. In the Hawaiian Archipelago, supporting evidence is primarily derived from underwater visual censuses in shallow waters (≤30 m). However, while many roving predators are present or potentially more abundant in deeper strata (30-100 m+), distributional information remains sparse. To partially fill that knowledge gap, we conducted surveys in the remote Northwestern Hawaiian Islands (NWHI) and populated Main Hawaiian Islands (MHI) from 2012-2014 using baited remote underwater stereo-video. Surveys between 0-100 m found considerable roving predator community dissimilarities between regions, marked conspicuous changes in species abundances with increasing depth, and largely corroborated patterns documented during shallow water underwater visual censuses, with up to an order of magnitude more jacks and five times more sharks sampled in the NWHI compared to the MHI. Additionally, several species were significantly more abundant and larger in mesophotic versus shallow depths, which remains particularly suggestive of deep-water refugia effects in the MHI. Stereo-video extends the depth range of current roving predator surveys in a more robust manner than was previously available, and appears to be well-suited for large-scale roving predator work in the Hawaiian Archipelago.

Long-term monitoring of coral reef fish assemblages in the Western central pacific.

Throughout the tropics, coral reef ecosystems, which are critically important to people, have been greatly altered by humans. Differentiating human impacts from natural drivers of ecosystem state is essential to effective management. Here we present a dataset from a large-scale monitoring program that surveys coral reef fish assemblages and habitats encompassing the bulk of the US-affiliated tropical Pacific, and spanning wide gradients in both natural drivers and human impact. Currently, this includes >5,500 surveys from 39 islands and atolls in Hawaii (including the main and Northwestern Hawaiian Islands) and affiliated geo-political regions of American Samoa, the Commonwealth of the Northern Mariana Islands, Guam, and the Pacific Remote Islands Areas. The dataset spans 2010-2017, during which time, each region was visited at least every three years, and ~500-1,000 surveys performed annually. This standardised dataset is a powerful resource that can be used to understand how human, environmental and oceanographic conditions influence coral reef fish community structure and function, providing a basis for research to support effective management outcomes.

Responses of Herbivorous Fishes and Benthos to 6 Years of Protection at the Kahekili Herbivore Fisheries Management Area, Maui.

In response to concerns about declining coral cover and recurring macroalgal blooms, in 2009 the State of Hawaii established the Kahekili Herbivore Fisheries Management Area (KHFMA). Within the KHFMA, herbivorous fishes and sea urchins are protected, but other fishing is allowed. As part of a multi-agency monitoring effort, we conducted surveys at KHFMA and comparison sites around Maui starting 19 months before closure, and over the six years since implementation of herbivore protection. Mean parrotfish and surgeonfish biomass both increased within the KHFMA (by 139% [95%QR (quantile range): 98-181%] and 28% [95%QR: 3-52%] respectively). Most of those gains were of small-to-medium sized species, whereas large-bodied species have not recovered, likely due to low levels of poaching on what are preferred fishery targets in Hawaii. Nevertheless, coincident with greater biomass of herbivores within the KHFMA, cover of crustose coralline algae (CCA) has increased from ~2% before closure to ~ 15% in 2015, and macroalgal cover has remained low throughout the monitoring period. Strong evidence that changes in the KHFMA were a consequence of herbivore management are that (i) there were no changes in biomass of unprotected fish families within the KHFMA; and that (ii) there were no similar changes in parrotfish or CCA at comparison sites around Maui. It is not yet clear how effective herbivore protection might eventually be for the KHFMA's ultimate goal of coral recovery. Coral cover declined over the first few years of surveys-from 39.6% (SE 1.4%) in 2008, to 32.9% (SE 0.8%) in 2012, with almost all of that loss occurring by 2010 (1 year after closure), i.e. before meaningful herbivore recovery had occurred. Coral cover subsequently stabilized and may have slightly increased from 2012 through early 2015. However, a region-wide bleaching event in 2015 had already led to some coral mortality by the time surveys were conducted in late 2015, at which time cover had dropped back to levels recorded in the KHFMA in 2012.