Nutrient enrichment can increase the severity of coral diseases

The prevalence and severity of marine diseases have increased over the last 20 years, significantly impacting a variety of foundation and keystone species. One explanation is that changes in the environment caused by human activities have impaired host resistance and/or have increased pathogen virulence. Here, we report evidence from field experiments that nutrient enrichment can significantly increase the severity of two important Caribbean coral epizootics: aspergillosis of the common gorgonian sea fan Gorgonia ventalina and yellow band disease of the reef‐building corals Montastraea annularis and M. franksii. Experimentally increasing nutrient concentrations by 2–5× nearly doubled host tissue loss caused by yellow band disease. In a separate experiment, nutrient enrichment significantly increased two measures of sea fan aspergillosis severity. Our results may help explain the conspicuous patchiness of coral disease severity, besides suggesting that minimizing nutrient pollution could be an important management tool for controlling coral epizootics.     

Turbid reefs moderate coral bleaching under climate‐related temperature stress

Thermal‐stress events that cause coral bleaching and mortality have recently increased in frequency and severity. Yet few studies have explored conditions that moderate coral bleaching. Given that high light and high ocean temperature together cause coral bleaching, we explore whether corals at turbid localities, with reduced light, are less likely to bleach during thermal‐stress events than corals at other localities. We analyzed coral bleaching, temperature, and turbidity data from 3,694 sites worldwide with a Bayesian model and found that K_d490, a measurement positively related to turbidity, between 0.080 and 0.127 reduced coral bleaching during thermal‐stress events. Approximately 12% of the world's reefs exist within this “moderating turbidity” range, and 30% of reefs that have moderating turbidity are in the Coral Triangle. We suggest that these turbid nearshore environments may provide some refuge through climate change, but these reefs will need high conservation status to sustain them close to dense human populations.     

Ocean acidification and warming will lower coral reef resilience

Ocean warming and acidification from increasing levels of atmospheric CO₂ represent major global threats to coral reefs, and are in many regions exacerbated by local‐scale disturbances such as overfishing and nutrient enrichment. Our understanding of global threats and local‐scale disturbances on reefs is growing, but their relative contribution to reef resilience and vulnerability in the future is unclear. Here, we analyse quantitatively how different combinations of CO₂ and fishing pressure on herbivores will affect the ecological resilience of a simplified benthic reef community, as defined by its capacity to maintain and recover to coral‐dominated states. We use a dynamic community model integrated with the growth and mortality responses for branching corals (Acropora) and fleshy macroalgae (Lobophora). We operationalize the resilience framework by parameterizing the response function for coral growth (calcification) by ocean acidification and warming, coral bleaching and mortality by warming, macroalgal mortality by herbivore grazing and macroalgal growth via nutrient loading. The model was run for changes in sea surface temperature and water chemistry predicted by the rise in atmospheric CO₂ projected from the IPCC's fossil‐fuel intensive A1FI scenario during this century. Results demonstrated that severe acidification and warming alone can lower reef resilience (via impairment of coral growth and increased coral mortality) even under high grazing intensity and low nutrients. Further, the threshold at which herbivore overfishing (reduced grazing) leads to a coral–algal phase shift was lowered by acidification and warming. These analyses support two important conclusions: Firstly, reefs already subjected to herbivore overfishing and nutrification are likely to be more vulnerable to increasing CO₂. Secondly, under CO₂ regimes above 450–500 ppm, management of local‐scale disturbances will become critical to keeping reefs within an Acropora‐rich domain.     

悬浮物对造礁石珊瑚营养方式的影响及其适应性研究进展

营养方式是造礁石珊瑚获取能量与营养物质的基础,影响其生长与分布。近年来珊瑚礁区悬浮物含量与组分结构发生显著变化,其对造礁石珊瑚营养方式的诸多影响正成为当前研究热点。研究系统综述了珊瑚礁区悬浮物变化特征、悬浮物对造礁石珊瑚营养方式的影响及其适应性研究现状。发现近年来人类活动加剧与强降雨事件频发是驱动珊瑚礁,尤其是近岸珊瑚礁区悬浮物含量递增、组分改变与变频加剧的主因;悬浮物变化对造礁石珊瑚光合自养与异养营养的影响存在显著的种间差异,这主要与悬浮物消光效应、生物可利用性及造礁石珊瑚种类密切相关。虽然少数种类造礁石珊瑚具光合可塑性或异养可塑性,能在高含量悬浮物存在的弱光环境中较好生长。然而对绝大多数造礁石珊瑚而言,其营养方式适应性较差,无法在悬浮物影响下较好地获取生命活动所需的能量与营养物质,进而难以生存。总体来说,悬浮物被认为是近年来影响造礁石珊瑚生长与分布的重要环境因子之一,而关于造礁石珊瑚营养方式对悬浮物变化的响应及其适应机制,当前研究仍较薄弱,需要进一步加强相关研究。     

Sunlight and water transparency: cornerstones in coral research

Reef-building corals throughout the world are considered endangered. The evidence is a decline in coral health and reduced coral cover. Competing hypotheses for the cause of coral loss include removal of grazers, nutrient enrichment, disease, coral bleaching, increase in temperature, and excess light/ultraviolet exposure. We suggest that light limitation as a second order effect of anthropogenic activity (e.g. sediment resuspension and nutrient enrichment) is a valid and tractable hypothesis. This experimental field and laboratory study demonstrates that corals of the Florida reefs are functioning close to the compensation point where respiration (of coral polyp plus zooxanthellae) consumes the products of photosynthesis of the zooxanthellae, with little if any remaining for growth. We extend this work into an optical nomograph that is useful for predicting coral loss and recovery. The nomograph is designed to elucidate compensation depth for waters of various transparencies.     

Nutrient Enrichment Coupled with Sedimentation Favors Sea Anemones over Corals

Fine sediments, which account for the majority of total fluvial sediment flux, have been suggested to degrade coral reefs on a global scale. Furthermore, sediment impacts can be exacerbated by extreme rainfall events associated with global climate change and anthropogenic nutrient enrichment. We report the findings from a series of mesocosm experiments exploring the effects of short-term sedimentation and nutrient enrichment on the interactions between the hard coral Acropora muricata, the sea anemone Mesactinia ganesis, and the green macroalga Codium edule. Mesocosms were manipulated to simulate either unimpacted reefs or reefs exposed to elevated levels of fine sediments for 10 or 14 days to simulate the effects of heavy rainfall. The first and second experiments were aimed to examine the effects of inorganic and organic sediments, respectively. The third experiment was designed to examine the interactive effects of nutrient enrichment and elevated sediment loads. Neither inorganic nor organic sediment loadings significantly affected the physiological performance of the coral, but, importantly, did reduce its ability to compete with other organisms. Photosynthetic efficiencies of both the green macroalga and the sea anemone increased in response to both sediment loadings when they were simultaneously exposed to nutrient enrichment. While organic sediment loading increased the nitrogen content of the green macroalga in the first experiment, inorganic sediment loading increased its phosphorus content in the second experiment. The coral mortality due to sea anemones attack was significantly greater upon exposure to enriched levels of organic sediments and nutrients. Our findings suggest that the combined effects of short-term sedimentation and nutrient enrichment could cause replacement of corals by sea anemones on certain coral reefs.     

Climate‐change refugia in the sheltered bays of Palau: analogs of future reefs

Coral bleaching and mortality are predicted to increase as climate change‐induced thermal‐stress events become more frequent. Although many studies document coral bleaching and mortality patterns, few studies have examined deviations from the expected positive relationships among thermal stress, coral bleaching, and coral mortality. This study examined the response of >30,000 coral colonies at 80 sites in Palau, during a regional thermal‐stress event in 2010. We sought to determine the spatial and taxonomic nature of bleaching and examine whether any habitats were comparatively resistant to thermal stress. Bleaching was most severe in the northwestern lagoon, in accordance with satellite‐derived maximum temperatures and anomalous temperatures above the long‐term averages. Pocillopora populations suffered the most extensive bleaching and the highest mortality. However, in the bays where temperatures were higher than elsewhere, bleaching and mortality were low. The coral‐community composition, constant exposure to high temperatures, and high vertical attenuation of light caused by naturally high suspended particulate matter buffered the corals in bays from the 2010 regional thermal‐stress event. Yet, nearshore reefs are also most vulnerable to land‐use change. Therefore, nearshore reefs should be given high conservation status because they provide refugia for coral populations as the oceans continue to warm.     

Assessing the effects of iron enrichment across holobiont compartments reveals reduced microbial nitrogen fixation in the Red Sea coral Pocillopora verrucosa

The productivity of coral reefs in oligotrophic tropical waters is sustained by an efficient uptake and recycling of nutrients. In reef‐building corals, the engineers of these ecosystems, this nutrient recycling is facilitated by a constant exchange of nutrients between the animal host and endosymbiotic photosynthetic dinoflagellates (zooxanthellae), bacteria, and other microbes. Due to the complex interactions in this so‐called coral holobiont, it has proven difficult to understand the environmental limitations of productivity in corals. Among others, the micronutrient iron has been proposed to limit primary productivity due to its essential role in photosynthesis and bacterial processes. Here, we tested the effect of iron enrichment on the physiology of the coral Pocillopora verrucosa from the central Red Sea during a 12‐day experiment. Contrary to previous reports, we did not see an increase in zooxanthellae population density or gross photosynthesis. Conversely, respiration rates were significantly increased, and microbial nitrogen fixation was significantly decreased. Taken together, our data suggest that iron is not a limiting factor of primary productivity in Red Sea corals. Rather, increased metabolic demands in response to iron enrichment, as evidenced by increased respiration rates, may reduce carbon (i.e., energy) availability in the coral holobiont, resulting in reduced microbial nitrogen fixation. This decrease in nitrogen supply in turn may exacerbate the limitation of other nutrients, creating a negative feedback loop. Thereby, our results highlight that the effects of iron enrichment appear to be strongly dependent on local environmental conditions and ultimately may depend on the availability of other nutrients.     

Global assessment of coral bleaching and required rates of adaptation under climate change

Elevated ocean temperatures can cause coral bleaching, the loss of colour from reef‐building corals because of a breakdown of the symbiosis with the dinoflagellate Symbiodinium. Recent studies have warned that global climate change could increase the frequency of coral bleaching and threaten the long‐term viability of coral reefs. These assertions are based on projecting the coarse output from atmosphere–ocean general circulation models (GCMs) to the local conditions around representative coral reefs. Here, we conduct the first comprehensive global assessment of coral bleaching under climate change by adapting the NOAA Coral Reef Watch bleaching prediction method to the output of a low‐ and high‐climate sensitivity GCM. First, we develop and test algorithms for predicting mass coral bleaching with GCM‐resolution sea surface temperatures for thousands of coral reefs, using a global coral reef map and 1985–2002 bleaching prediction data. We then use the algorithms to determine the frequency of coral bleaching and required thermal adaptation by corals and their endosymbionts under two different emissions scenarios. The results indicate that bleaching could become an annual or biannual event for the vast majority of the world's coral reefs in the next 30–50 years without an increase in thermal tolerance of 0.2–1.0°C per decade. The geographic variability in required thermal adaptation found in each model and emissions scenario suggests that coral reefs in some regions, like Micronesia and western Polynesia, may be particularly vulnerable to climate change. Advances in modelling and monitoring will refine the forecast for individual reefs, but this assessment concludes that the global prognosis is unlikely to change without an accelerated effort to stabilize atmospheric greenhouse gas concentrations.     

Changes in gametogenesis and fecundity of acroporid corals that were exposed to elevated nitrogen and phosphorus during the ENCORE experiment

Colonies of two scleractinian reef coral species, Acropora longicyathus and Acropora aspera were transplanted into patch reefs at One Tree Reef, Great Barrier Reef, Australia as part of the ENCORE experiment. These corals and colonies of A. aspera which were naturally present in the patch reefs were exposed to four treatments over two years: controls with normal seawater, elevated levels of nitrogen only, phosphorus only, or nitrogen plus phosphorus. These corals were sampled and used to determine whether gametogenic cycles and fecundity were affected by nutrient enrichment. Acropora longicyathus had a single annual gametogenic cycle. Corals exposed to elevated nitrogen produced significantly smaller and fewer eggs and contained less testes material than those which were not exposed to nitrogen. Exposure to elevated phosphorus only resulted in corals producing more but smaller eggs, and more testes material. Egg numbers of colonies from other treatments decreased as the gametogenic cycles continued, but those of the phosphorus colonies showed almost no reduction in egg numbers between the early and late stages of the gametogenic cycles. These results have important management implications for coral reefs as they demonstrate that small increases in concentrations of nitrogen and phosphorus can have severe effects on reproductive activity in these species of scleractinian corals.     

Suppressed recovery of functionally important branching Acropora drives coral community composition changes following mass bleaching in Indonesia

Mass coral bleaching events may have disproportionate effects on branching corals, leading to coral community restructuring, reduced biodiversity, and decreased structural complexity. This affects overall reef health and resilience. Functionally important, fast-growing branching Acropora corals were a historically dominant and vital component of Indonesian reefs throughout the twentieth century, yet the genus is also one of the most vulnerable to external stressors. This study used long-term annual reef monitoring data from Indonesia’s Wakatobi Marine National Park (WMNP) to investigate the effects of a mass bleaching event in 2010 on Acropora and other branching corals, evaluate their post-disturbance recovery trajectories, and analyse shifts in coral community composition. Post-bleaching scleractinian coral cover decreased across study sites, with losses in branching corals especially evident. Long-term branching Acropora cover decreased significantly and failed to demonstrate the significant post-disturbance recovery of other branching corals (especially Porites). In areas characterised by relatively high branching Acropora cover (> 15% mean cover) prior to bleaching, long-term coral community composition changes have trended predominately towards branching and massive Porites and branching Montipora. The novelty and key contribution of this study is that results suggest suppressed recovery of Acropora in the WMNP. Contributing factors may include the Allee effect (inhibition of reproduction at low population densities), other forms of inhibited larval recruitment, direct and indirect spatial competition, and changes in the physical reef habitat. These findings have critical implications for this functionally important taxon, future reef conservation efforts, and overall reef health and resilience in the park.

     

The distribution of scleractinian corals in the Bay of Biscay, NE Atlantic

Studies on the distribution of scleractinian corals in the Bay of Biscay were relatively numerous during the nineteenth and the first half of the twentieth century. Yet, recent reports are scattered, sometimes unpublished, and therefore knowledge about the current coral occurrence in the area is limited. This study aims at compiling the available historical and more recently collected information on the occurrence of scleractinian corals in the Bay of Biscay. Data from two recent cruises are included and compared with previous explored coral sites from as early as 1830 up to 1995. A database of 347 records including 34 described species of scleractinians highlights that the slope of the Bay of Biscay is an important habitat for scleractinians. This could be expected due to the high topographic relief providing the necessary hard substrate and accelerated bottom current flow that corals require. Further exploration of the occurrence and ecology of corals in the area is recommended to support the conservation of cold-water coral reefs along the European margin. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Identity and diversity of coral endosymbionts (zooxanthellae) from three Palauan reefs with contrasting bleaching, temperature and shading histories

The potential of corals to associate with more temperature-tolerant strains of algae (zooxanthellae, Symbiodinium) can have important implications for the future of coral reefs in an era of global climate change. In this study, the genetic identity and diversity of zooxanthellae was investigated at three reefs with contrasting histories of bleaching mortality, water temperature and shading, in the Republic of Palau (Micronesia). Single-stranded conformation polymorphism and sequence analysis of the ribosomal DNA internal transcribed spacer (ITS)1 region was used for genotyping. A chronically warm but partly shaded coral reef in a marine lake that is hydrographically well connected to the surrounding waters harboured only two single-stranded conformation polymorphism profiles (i.e. zooxanthella communities). It consisted only of Symbiodinium D in all 13 nonporitid species and two Porites species investigated, with the remaining five Porites harbouring C*. Despite the high temperature in this lake (> 0.5 degrees above ambient), this reef did not suffer coral mortality during the (1998) bleaching event, however, no bleaching-sensitive coral families and genera occur in the coral community. This setting contrasts strongly with two other reefs with generally lower temperatures, in which 10 and 12 zooxanthella communities with moderate to low proportions of clade D zooxanthellae were found. The data indicate that whole coral assemblages, when growing in elevated seawater temperatures and at reduced irradiance, can be composed of colonies associated with the more thermo-tolerant clade D zooxanthellae. Future increases in seawater temperature might, therefore, result in an increasing prevalence of Symbiodinium phylotype D in scleractinian corals, possibly associated with a loss of diversity in both zooxanthellae and corals.     

Bottlenecks to coral recovery in the Seychelles

Processes that affect recovery of coral assemblages require investigation because coral reefs are experiencing a diverse array of more frequent disturbances. Potential bottlenecks to coral recovery include limited larval supply, low rates of settlement, and high mortality of new recruits or juvenile corals. We investigated spatial variation in local abundance of scleractinian corals in the Seychelles at three distinct life history stages (recruits, juveniles, and adults) on reefs with differing benthic conditions. Following widespread coral loss due to the 1998 bleaching event, some reefs are recovering (i.e., relatively high scleractinian coral cover: ‘coral-dominated’), some reefs have low cover of living macrobenthos and unconsolidated rubble substrates (‘rubble-dominated’), and some reefs have high cover of macroalgae (‘macroalgal-dominated’). Rates of coral recruitment to artificial settlement tiles were similar across all reef conditions, suggesting that larval supply does not explain differential coral recovery across the three reef types. However, acroporid recruits were absent on macroalgal-dominated reefs (0.0 ± 0.0 recruits tile^?1) in comparison to coral-dominated reefs (5.2 ± 1.6 recruits tile^?1). Juvenile coral colony density was significantly lower on macroalgal-dominated reefs (2.4 ± 1.1 colonies m^?2), compared to coral-dominated reefs (16.8 ± 2.4 m^?2) and rubble-dominated reefs (33.1 ± 7.3 m^?2), suggesting that macroalgal-dominated reefs have either a bottleneck to successful settlement on the natural substrates or a high post-settlement mortality bottleneck. Rubble-dominated reefs had very low cover of adult corals (10.0 ± 1.7 %) compared to coral-dominated reefs (33.4 ± 3.6 %) despite no statistical difference in their juvenile coral densities. A bottleneck caused by low juvenile colony survivorship on unconsolidated rubble-dominated reefs is possible, or alternatively, recruitment to rubble-dominated reefs has only recently begun. This study identified bottlenecks to recovery of coral assemblages that varied depending on post-disturbance habitat condition.     

Seabird nutrient subsidies alter patterns of algal abundance and fish biomass on coral reefs following a bleaching event

Cross‐ecosystem nutrient subsidies play a key role in the structure and dynamics of recipient communities, but human activities are disrupting these links. Because nutrient subsidies may also enhance community stability, the effects of losing these inputs may be exacerbated in the face of increasing climate‐related disturbances. Nutrients from seabirds nesting on oceanic islands enhance the productivity and functioning of adjacent coral reefs, but it is unknown whether these subsidies affect the response of coral reefs to mass bleaching events or whether the benefits of these nutrients persist following bleaching. To answer these questions, we surveyed benthic organisms and fishes around islands with seabirds and nearby islands without seabirds due to the presence of invasive rats. Surveys were conducted in the Chagos Archipelago, Indian Ocean, immediately before the 2015–2016 mass bleaching event and, in 2018, two years following the bleaching event. Regardless of the presence of seabirds, relative coral cover declined by 32%. However, there was a post‐bleaching shift in benthic community structure around islands with seabirds, which did not occur around islands with invasive rats, characterized by increases in two types of calcareous algae (crustose coralline algae [CCA] and Halimeda spp.). All feeding groups of fishes were positively affected by seabirds, but only herbivores and piscivores were unaffected by the bleaching event and sustained the greatest difference in biomass between islands with seabirds versus those with invasive rats. By contrast, corallivores and planktivores, both of which are coral‐dependent, experienced the greatest losses following bleaching. Even though seabird nutrients did not enhance community‐wide resistance to bleaching, they may still promote recovery of these reefs through their positive influence on CCA and herbivorous fishes. More broadly, the maintenance of nutrient subsidies, via strategies including eradication of invasive predators, may be important in shaping the response of ecological communities to global climate change.     

Three-dimensional species distribution modelling reveals the realized spatial niche for coral recruitment on contemporary Caribbean reefs

The three-dimensional structure of habitats is a critical component of species' niches driving coexistence in species-rich ecosystems. However, its influence on structuring and partitioning recruitment niches has not been widely addressed. We developed a new method to combine species distribution modelling and structure from motion, and characterized three-dimensional recruitment niches of two ecosystem engineers on Caribbean coral reefs, scleractinian corals and gorgonians. Fine-scale roughness was the most important predictor of suitable habitat for both taxa, and their niches largely overlapped, primarily due to scleractinians' broader niche breadth. Crevices and holes at mm scales on calcareous rock with low coral cover were more suitable for octocorals than for scleractinian recruits, suggesting that the decline in scleractinian corals is facilitating the recruitment of octocorals on contemporary Caribbean reefs. However, the relative abundances of the taxa were independent of the amount of suitable habitat on the reef, emphasizing that niche processes alone do not predict recruitment rates.     

Algal contact as a trigger for coral disease

Diseases are causing alarming declines in reef‐building coral species, the foundation blocks of coral reefs. The emergence of these diseases has occurred simultaneously with large increases in the abundance of benthic macroalgae. Here, we show that physical contact with the macroalga Halimeda opuntia can trigger a virulent disease known as white plague type II that has caused widespread mortality in most Caribbean coral species. Colonies of the dominant coral Montastraea faveolata exposed to algal transplants developed the disease whereas unexposed colonies did not. The bacterium Aurantimonas coralicida, causative agent of the disease, was present on H. opuntia sampled close to, and away from diseased corals, indicating that the alga serves as a reservoir for this pathogen. Our results suggest that the spread of macroalgae on coral reefs could account for the elevated incidence of coral diseases over past decades and that reduction of macroalgal abundance could help control coral epizootics.     

Rehabilitation of coral reefs through removal of macroalgae: state of knowledge and considerations for management and implementation

Coral reef ecosystems are under increasing pressure by multiple stressors that degrade reef condition and function. Although improved management systems have yielded benefits in many regions, broad‐scale declines continue and additional practical and effective solutions for reef conservation and management are urgently needed. Ecological interventions to assist or enhance ecosystem recovery are standard practice in many terrestrial management regimes, and they are now increasingly being implemented in the marine environment. Intervention activities in coral reef systems include the control of coral predators (e.g. crown‐of‐thorns starfish), substrate modification, the creation of artificial habitats and the cultivation, transplantation, and assisted recruitment of corals. On many coastal reefs, corals face competition and overgrowth by fleshy macroalgae whose abundance may be elevated due to acute disturbance events, chronic nutrient enrichment, and reduced herbivory. Active macroalgae removal has been proposed and trialed as a management tool to reduce competition between algae and corals and provide space for coral recruitment, in the hope of restoring the spatial dominance of habitat‐forming corals. However, macroalgae removal has received little formal attention as a method of reef restoration. This review synthesizes available knowledge of the ecological role of macroalgae on coral reefs and the potential benefits and risks associated with their active removal.     

Role of host genetics and heat‐tolerant algal symbionts in sustaining populations of the endangered coral Orbicella faveolata in the Florida Keys with ocean warming

Identifying which factors lead to coral bleaching resistance is a priority given the global decline of coral reefs with ocean warming. During the second year of back‐to‐back bleaching events in the Florida Keys in 2014 and 2015, we characterized key environmental and biological factors associated with bleaching resilience in the threatened reef‐building coral Orbicella faveolata. Ten reefs (five inshore, five offshore, 179 corals total) were sampled during bleaching (September 2015) and recovery (May 2016). Corals were genotyped with 2bRAD and profiled for algal symbiont abundance and type. O. faveolata at the inshore sites, despite higher temperatures, demonstrated significantly higher bleaching resistance and better recovery compared to offshore. The thermotolerant Durusdinium trenchii (formerly Symbiondinium trenchii) was the dominant endosymbiont type region‐wide during initial (78.0% of corals sampled) and final (77.2%) sampling; >90% of the nonbleached corals were dominated by D. trenchii. 2bRAD host genotyping found no genetic structure among reefs, but inshore sites showed a high level of clonality. While none of the measured environmental parameters were correlated with bleaching, 71% of variation in bleaching resistance and 73% of variation in the proportion of D. trenchii was attributable to differences between genets, highlighting the leading role of genetics in shaping natural bleaching patterns. Notably, D. trenchii was rarely dominant in O. faveolata from the Florida Keys in previous studies, even during bleaching. The region‐wide high abundance of D. trenchii was likely driven by repeated bleaching associated with the two warmest years on record for the Florida Keys (2014 and 2015). On inshore reefs in the Upper Florida Keys, O. faveolata was most abundant, had the highest bleaching resistance, and contained the most corals dominated by D. trenchii, illustrating a causal link between heat tolerance and ecosystem resilience with global change.