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The global impacts of climate change are evident in every marine ecosystem. On coral reefs, mass coral bleaching and mortality have emerged as ubiquitous responses to ocean warming, yet one of the greatest challenges of this epiphenomenon is linking information across scientific disciplines and spatial and temporal scales. Here we review some of the seminal and recent coral-bleaching discoveries from an ecological, physiological, and molecular perspective. We also evaluate which data and processes can improve predictive models and provide a conceptual framework that integrates measurements across biological scales. Taking an integrative approach across biological and spatial scales, using for example hierarchical models to estimate major coral-reef processes, will not only rapidly advance coral-reef science but will also provide necessary information to guide decision-making and conservation efforts. To conserve reefs, we encourage implementing mesoscale sanctuaries (thousands of km2 ) that transcend national boundaries. Such networks of protected reefs will provide reef connectivity, through larval dispersal that transverse thermal environments, and genotypic repositories that may become essential units of selection for environmentally diverse locations. Together, multinational networks may be the best chance corals have to persist through climate change, while humanity struggles to reduce emissions of greenhouse gases to net zero. published

Corals nucleate and grow aragonite crystals, organizing them into intricate skeletal structures that ultimately build the world’s coral reefs. Crystallography and chemistry have profound influence on the material properties of these skeletal building blocks, yet gaps remain in our knowledge about coral aragonite on the atomic scale. Across a broad diversity of shallow-water and deep-sea scleractinian corals from vastly different environments, coral aragonites are remarkably similar to one another, confirming that corals exert control on the carbonate chemistry of the calcifying space relative to the surrounding seawater. Nuances in coral aragonite structures relate most closely to trace element chemistry and aragonite saturation state, suggesting the primary controls on aragonite structure are ionic strength and trace element chemistry, with growth rate playing a secondary role. We also show how coral aragonites are crystallographically indistinguishable from synthetic abiogenic aragonite analogs precipitated from seawater under conditions mimicking coral calcifying fluid. In contrast, coral aragonites are distinct from geologically formed aragonites, a synthetic aragonite precipitated from a freshwater solution, and mollusk aragonites. Crystallographic signatures have future applications in understanding the material properties of coral aragonite and predicting the persistence of coral reefs in a rapidly changing ocean.

Rising temperatures and extreme climate events are propelling tropical species into temperate marine ecosystems, but not all species can persist. Here, we used the heatwave-driven expatriation of tropical Black Rabbitfish (Siganus fuscescens) to the temperate environments of Western Australia to assess the ecological and evolutionary mechanisms that may entail their persistence. Population genomic assays for this rabbitfish indicated little genetic differentiation between tropical residents and vagrants to temperate environments due to high migration rates, which were likely enhanced by the marine heatwave. DNA metabarcoding revealed a diverse diet for this species based on phytoplankton and algae, as well as an ability to feed on regional resources, including kelp. Irrespective of future climate scenarios, these macroalgae-consuming vagrants may self-recruit in temperate environments and further expand their geographic range by the year 2100. This expansion may compromise the health of the kelp forests that form Australia’s Great Southern Reef. Overall, our study demonstrates that projected favourable climate conditions, continued large-scale genetic connectivity between populations, and diet versatility are key for tropical range-shifting fish to establish in temperate ecosystems., Laura Gajdzik et al. used population genomics, dietary DNA metabarcoding, and climate models to understand the influence of the 2010/2011 marine heatwave on the migration of rabbitfish from tropical environments into temperate environments in Western Australia. Their results reveal (i) little genetic differentiation and high migration rates between tropical residents and temperate vagrants, (ii) an overall diverse diet (including phytoplankton, local and widespread algae, and kelp), and (iii) projected suitable climate conditions for vagrants to establish in temperate marine waters.

Intensified coastal development is compromising the health and functioning of marine ecosystems. A key example of this is the Red Sea, a biodiversity hotspot subjected to increasing local human pressures. While some marine‐protected areas (MPAs) were placed to alleviate these stressors, it is unclear whether these MPAs are managed or enforced, thus providing limited protection. Yet, most importantly, MPAs in the Red Sea were not designed using climate considerations, likely diminishing their effectiveness against global stressors. Here, we propose to tailor the design of MPAs in the Red Sea by integrating approaches to enhance climate change mitigation and adaptation. First, including coral bleaching susceptibility could produce a more resilient network of MPAs by safeguarding reefs from different thermal regions that vary in spatiotemporal bleaching responses, reducing the risk that all protected reefs will bleach simultaneously. Second, preserving the basin‐wide genetic connectivity patterns that are assisted by mesoscale eddies could further ensure recovery of sensitive populations and maintain species potential to adapt to environmental changes. Finally, protecting mangrove forests in the northern and southern Red Sea that act as major carbon sinks could help offset greenhouse gas emissions. If implemented with multinational cooperation and concerted effort among stakeholders, our portfolio of climate‐tailored approaches may help build a network of MPAs in the Red Sea that protects more effectively its coastal resources against escalating coastal development and climate instability. Beyond the Red Sea, we anticipate this study to serve as an example of how to improve the utility of tropical MPAs as climate‐informed conservation tools., A marine protected area (MPA) design for the Red Sea that buffers against the effects of global climate change includes coral reefs that vary in bleaching susceptibility over time: from unbleached/healthy, to heat‐stress‐ and upwelling‐induced bleaching, to recovering‐from‐bleaching reefs. Crucially, such MPA design also accounts for seascape genetics by ensuring these reefs are genetically and physically (through mesoscale eddies) connected to promote recovery and adaptation. Finally, a climate‐informed MPA design helps offset carbon dioxide emissions (CO2) by protecting high blue carbon areas, which are large mangrove forests that uptake CO2 from the atmosphere via photosynthesis and sequester it in sediments.

The structure and function of coral reef ecosystems is increasingly compromised by multiple stressors, even in the most remote locations. Severe, acute disturbances such as volcanic eruptions represent extreme events that can annihilate entire reef ecosystems, but also provide unique opportunities to examine ecosystem resilience and recovery. Here, we examine the destruction, persistence and initial recovery of reefs associated with the hydro-magmatic eruption that created Earth’s newest landmass, the Hunga Tonga–Hunga Ha’apai volcanic island. Despite extreme conditions associated with the eruption, impacts on nearby reefs were spatially variable. Importantly, even heavily affected reefs showed signs of rapid recovery driven by high recruitment, likely from local refuges. The remote location and corresponding lack of additional stressors likely contribute to the resilience of Hunga’s reefs, suggesting that in the absence of chronic anthropogenic stressors, coral reefs can be resilient to one of the largest physical disturbances on Earth.

Coral reefs are experiencing a dramatic loss of hard coral abundance and associated habitat structure from a myriad of local and global factors. Here, utilizing U–Th radiometric age-dating of coral death assemblages, we investigated patterns of coral mortality from the eastern margin of the Red Sea along a latitudinal gradient (Yanbu, 24o N; Thuwal, 22o N; Al-Lith, 19o N; Farasan Banks, 18o N) in 2018 and 2019. In all four regions, radiometric ages of in situ dead Acropora and Pocillopora colonies were largely confined to the late twentieth and early twenty-first century. During the early twenty-first century, coral mortality was found to be synchronous with previously documented bleaching events in 2010 and 2015 and, at one site (Farasan Banks), an outbreak of crown-of-thorns starfish (COTS) in 2009. The most northern site, Yanbu, had the highest relative percentage of live coral (42 ± 4%) and of living Acropora, and may serve as a refugium under climate warming scenarios. For the three southern regions (Thuwal, Al-Lith, Farasan Banks) benthic structure was mostly comprised of dead corals. The southernmost survey site, Farasan Banks, underwent a dramatic change in coral benthic structure associated with a COTS outbreak in 2009 and a bleaching event in 2015, and had the lowest relative percentage of live coral (6 ± 2%), comprised mostly of massive Porites, with no live Acropora or Pocillopora. Our results highlight the asynchronous impact of disturbance events on eastern Red Sea coral reefs and emphasize regional differences in recovery and ecosystem state.

Bioerosion on turbid inshore reefs is expected to increase with global climate change reducing reef stability and accretionary potential. Most studies investigating bioerosion have focused on external grazers, such as parrotfish and urchins, whose biomass is more easily measured. Yet, cryptic endolithic bioeroders such as macroboring (worms, sponges and bivalves) and microboring taxa (fungus and algae) have the potential to be the dominant bioeroders, especially among inshore reef systems exposed to increased nutrient supply. We measured bioerosion rates of three bioeroder groups (microborers, macroborers, and grazers), and their response to environmental parameters (temperature, turbidity, light, chlorophyll a, salinity, pH) across two inshore reefs of north Western Australia. Total bioerosion rates were low (0.152 ± 0.012 kg m-1 yr-1) compared to average global rates, but were comparable to other inshore turbid reefs. Macroboring worms were the dominant source of bioerosion and displayed a significant negative relationship with temperature (r2= 0.24, P=0.008) and light (r2= 0.16, P=0.037). A global assessment of environmental influences on bioerosion further identified chlorophyll-a as a significant driver of macroboring (r2= 0.60, P=

Significance The growth of coral reefs is threatened by the dual stressors of ocean warming and acidification. Despite a wealth of studies assessing the impacts of climate change on individual taxa, projections of their impacts on coral reef net carbonate production are limited. By projecting impacts across 233 different locations, we demonstrate that the majority of coral reefs will be unable to maintain positive net carbonate production globally by the year 2100 under representative concentration pathways RCP4.5 and 8.5, while even under RCP2.6, coral reefs will suffer reduced accretion rates. Our results provide quantitative projections of how different climate change stressors will influence whole ecosystem carbonate production across coral reefs in all major ocean basins., Ocean warming and acidification threaten the future growth of coral reefs. This is because the calcifying coral reef taxa that construct the calcium carbonate frameworks and cement the reef together are highly sensitive to ocean warming and acidification. However, the global-scale effects of ocean warming and acidification on rates of coral reef net carbonate production remain poorly constrained despite a wealth of studies assessing their effects on the calcification of individual organisms. Here, we present global estimates of projected future changes in coral reef net carbonate production under ocean warming and acidification. We apply a meta-analysis of responses of coral reef taxa calcification and bioerosion rates to predicted changes in coral cover driven by climate change to estimate the net carbonate production rates of 183 reefs worldwide by 2050 and 2100. We forecast mean global reef net carbonate production under representative concentration pathways (RCP) 2.6, 4.5, and 8.5 will decline by 76, 149, and 156%, respectively, by 2100. While 63% of reefs are projected to continue to accrete by 2100 under RCP2.6, 94% will be eroding by 2050 under RCP8.5, and no reefs will continue to accrete at rates matching projected sea level rise under RCP4.5 or 8.5 by 2100. Projected reduced coral cover due to bleaching events predominately drives these declines rather than the direct physiological impacts of ocean warming and acidification on calcification or bioerosion. Presently degraded reefs were also more sensitive in our analysis. These findings highlight the low likelihood that the world’s coral reefs will maintain their functional roles without near-term stabilization of atmospheric CO2 emissions.

Ocean acidification (OA) is a major threat to coral reefs, which are built by calcareous species. However, long-term assessments of the impacts of OA are scarce, limiting the understanding of the capacity of corals and coralline algae to acclimatize to high partial pressure of carbon dioxide ( $${p}_{\mathrm{CO}_{2}}$$ ) levels. Species-specific sensitivities to OA are influenced by its impacts on chemistry within the calcifying fluid (CF). Here, we investigate the capacity of multiple coral and calcifying macroalgal species to acclimatize to elevated $${p}_{\mathrm{CO}_{2}}$$ by determining their chemistry in the CF during a year-long experiment. We found no evidence of acclimatization to elevated $${p}_{\mathrm{CO}_{2}}$$ across any of the tested taxa. The effects of increasing seawater $${p}_{\mathrm{CO}_{2}}$$ on the CF chemistry were rapid and persisted until the end of the experiment. Our results show that acclimatization of the CF chemistry does not occur within one year, which confirms the threat of OA for future reef accretion and ecological function. Calcifying marine species are threatened by ocean acidification. A year-long study of calcifying fluid chemistry shows that reef species, both corals and calcifying macroalgal species, were not able to adapt to ocean acidification.

Ocean warming is negatively impacting coral reef ecosystems and considerable effort is currently invested in projecting coral reef futures under 21st century climate change. A limiting factor in these projections is lack of quantitative data on the thermal thresholds of different reef communities, due in large part to spatial and temporal gaps in bleaching observations. Here we apply a coral bleaching proxy, skeletal stress bands, to reconstruct the history of bleaching on eight coral reefs in the central equatorial Pacific (CEP) and use this information to constrain the thermal thresholds of their coral communities. First, three genera of massive corals collected on both Pacific and Caribbean reefs are used to derive a calibration between the proportion of corals that form stress bands during a bleaching event, and the total observed bleaching incidence in the community of mixed coral taxa. The correlation is highly significant, indicating that stress bands in massive corals reflect community-level bleaching severity (R2 = 0.945, p

X-ray computed tomography (CT) is a non-destructive imaging technique that provides three-dimensional (3D) visualisation and high-resolution quantitative data in the form of CT numbers. CT numbers are derived as a function of the X-ray energy, effective atomic number and density of the sample. The sensitivity of the CT number to changes in material density allows it to successfully identify facies changes within sediment cores by detecting downcore shifts in sediment properties, and quantify skeletal linear extension rates and the volume of internal voids from biological erosion of coral cores. Here we present two algorithms to analyse CT scan images specific to geoscience research packaged within an open source MATLAB application (Core-CT). The first algorithm facilitates the computation of representative CT numbers from a user-defined region of interest to identify boundaries of density change (e.g. sedimentary facies, laminations, coral growth bands). The second algorithm enables the segmentation of regions with major density contrast (e.g. internal void space or biogenic material) and the geometric measurements of these irregularities. The versatility of Core-CT for geoscience applications is then demonstrated by utilising CT scans from a range of environmental settings comprising both sediment (Lake Huelde, Chile and Kallang River Basin, Singapore) and coral cores (Thuwal region of Red Sea, Saudi Arabia). Analysis of sediment cores show the capabilities of Core-CT to: 1) locate tsunami deposits from lacustrine sediments, 2) provide rapid and detailed measurement of varved sediments, and 3) identify sedimentary facies from an unsplit shallow marine sediment core. Analysis of coral cores allow us to successfully measure skeletal linear extension from annual growth bands, and provide volumetric quantification and 3D visualisation of internal bioerosion. Core-CT is an accessible, multi-use MATLAB based program that is freely available at GitHub (https://github.com/yuting-yan/Core-CT).

Coral reef ecosystems are highly sensitive to thermal anomalies, making them vulnerable to ongoing global warming. Yet, a variety of cooling mechanisms, such as upwelling, offer some respite to cer...

Coral reefs provide critical ecological and geomorphic (e.g. sediment production for reef-fronted shoreline maintenance) services, which interact in complex and dynamic ways. These services are under threat from climate change, requiring dynamic modelling approaches that predict how reef systems will respond to different future climate scenarios. Carbonate budgets, which estimate net reef calcium carbonate production, provide a comprehensive ‘snap-shot’ assessment of reef accretionary potential and reef stability. These budgets, however, were not intended to account for the full suite of processes that maintain coral reef services or to provide predictive capacity on longer timescales (decadal to centennial). To respond to the dual challenges of enhancing carbonate budget assessments and advancing their predictive capacity, we applied a novel model elicitation and review method to create a qualitative geo-ecological carbonate reef system model that links geomorphic, ecological and physical processes. Our approach conceptualizes relationships between net carbonate production, sediment transport and landform stability, and rates knowledge confidence to reveal major knowledge gaps and critical future research pathways. The model provides a blueprint for future coral reef research that aims to quantify net carbonate production and sediment dynamics, improving our capacity to predict responses of reefs and reef-fronted shorelines to future climate change.

Nutrients become a death knell 5 for corals as the ocean warms., With predictions that mass coral bleaching will occur annually within this century, conservation efforts must focus their limited resources based on an accurate understanding of the drivers of bleaching. Here, we provide spatial and temporal evidence that excess nutrients exacerbate the detrimental effects of heat stress to spark mass coral bleaching in the Red Sea. Exploiting this region’s unique oceanographic regime, where nutrients and heat stress vary independently, we demonstrate that the world’s third largest coral reef system historically suffered from severe mass bleaching only when exposed to both unusually high temperature and nutrients. Incorporating nutrient-supplying ocean currents and their variability into coral bleaching forecasts will be critical for effectively guiding efforts to safeguard the reefs most likely to persist in the Anthropocene.

Crustose coralline algae play a crucial role in the building of reefs in the photic zones of nearshore ecosystems globally, and are highly susceptible to ocean acidification1–3. Nevertheless, the extent to which ecologically important crustose coralline algae can gain tolerance to ocean acidification over multiple generations of exposure is unknown. We show that, while calcification of juvenile crustose coralline algae is initially highly sensitive to ocean acidification, after six generations of exposure the effects of ocean acidification disappear. A reciprocal transplant experiment conducted on the seventh generation, where half of all replicates were interchanged across treatments, confirmed that they had acquired tolerance to low pH and not simply to laboratory conditions. Neither exposure to greater pH variability, nor chemical conditions within the micro-scale calcifying fluid internally, appeared to play a role in fostering this capacity. Our results demonstrate that reef-accreting taxa can gain tolerance to ocean acidification over multiple generations of exposure, suggesting that some of these cosmopolitan species could maintain their critical ecological role in reef formation. Crustose coralline algae help build coral reef structures through calcification, a process threatened under ocean acidification. Juvenile algae were highly sensitive on initial exposure to ocean acidification, but continued exposure over six generations showed a gain of tolerance.

In a recent review, “Bioerosion: the other ocean acidification problem,” Schönberg et al. claim that studies of bioerosion across natural chemical gradients are “flawed” or “compromised” by co-variation among environmental factors. Their discussion falls largely on two publications, Silbiger et al. and DeCarlo et al. Here, we demonstrate that critical errors in plotting, statistical analysis, and data selection in Schönberg et al.’s reanalysis, result in a gross misrepresentation of these studies. Further, we argue three key points regarding field-based studies that require broader discussion within the bioerosion community and marine scientists in general: (1) that natural variability in field studies is not a flaw, (2) interpretations must be supported by mechanistic understanding, and (3) field-based studies play an essential role in elucidating interactions between OA and natural variability that is not captured by laboratory CO2-manipulation experiments. Our goal with this comment is to encourage open discussion of the advantages and caveats of field-based studies in general, and ultimately, advance our understanding of bioerosion patterns observed in nature.

Coral barium to calcium (Ba/Ca) ratios have been used to reconstruct records of upwelling, river and groundwater discharge, and sediment and dust input to the coastal ocean. However, this proxy has not yet been explicitly tested to determine if Ba inclusion in the coral skeleton is directly proportional to seawater Ba concentration and to further determine how additional factors such as temperature and calcification rate control coral Ba/Ca ratios. We measured the inclusion of Ba within aquaria reared juvenile corals ( Favia fragum ) at three temperatures (∼27.7, 24.6 and 22.5 °C) and three seawater Ba concentrations (73, 230 and 450 nmol kg −1 ). Coral polyps were settled on tiles conditioned with encrusting coralline algae, which complicated chemical analysis of the coral skeletal material grown during the aquaria experiments. We utilized Sr/Ca ratios of encrusting coralline algae (as low as 3.4 mmol mol −1 ) to correct coral Ba/Ca for this contamination, which was determined to be 26 ± 11% using a two end member mixing model. Notably, there was a large range in Ba/Ca across all treatments, however, we found that Ba inclusion was linear across the full concentration range. The temperature sensitivity of the distribution coefficient is within the range of previously reported values. Finally, calcification rate, which displayed large variability, was not correlated to the distribution coefficient. The observed temperature dependence predicts a change in coral Ba/Ca ratios of 1.1 μmol mol −1 from 20 to 28 °C for typical coastal ocean Ba concentrations of 50 nmol kg −1 . Given the linear uptake of Ba by corals observed in this study, coral proxy records that demonstrate peaks of 10–25 μmol mol −1 would require coastal seawater Ba of between 60 and 145 nmol kg −1 . Further validation of the coral Ba/Ca proxy requires evaluation of changes in seawater chemistry associated with the environmental perturbation recorded by the coral as well as verification of these results for Porites species, which are widely used in paleo reconstructions.

From air to shining sea Nitrogen is an essential nutrient for phytoplankton growth. Nitrogen is primarily supplied to the surface ocean by mixing from below. However, as fertilizer use and combustion of fossil fuels rise, the atmosphere is expected to become an increasingly important source. Ren et al. measured nitrogen isotopes in organic matter from a South China Sea coral (see the Perspective by Boyle). Their findings suggest that atmospheric deposition of anthropogenic nitrogen began right at the end of the 20th century. This pathway now supplies nearly one quarter of the annual nitrogen input to the surface ocean in this region. Science , this issue p. 749 ; see also p. 700

A major challenge in modeling the circulation over coral reefs is uncertainty in the drag coefficient because existing estimates span two orders of magnitude. Current and pressure measurements from five coral reefs are used to estimate drag coefficients based on depth-average flow, assuming a balance between the cross-reef pressure gradient and the bottom stress. At two sites wind stress is a significant term in the cross-reef momentum balance and is included in estimating the drag coefficient. For the five coral reef sites and a previous laboratory study, estimated drag coefficients increase as the water depth decreases consistent with open channel flow theory. For example, for a typical coral reef hydrodynamic roughness of 5 cm, observational estimates, and the theory indicate that the drag coefficient decreases from 0.4 in 20 cm of water to 0.005 in 10 m of water. Synthesis of results from the new field observations with estimates from previous field and laboratory studies indicate that coral reef drag coefficients range from 0.2 to 0.005 and hydrodynamic roughnesses generally range from 2 to 8 cm. While coral reef drag coefficients depend on factors such as physical roughness and surface waves, a substantial fraction of the scatter in estimates of coral reef drag coefficients is due to variations in water depth.

The skeletons of many reef-building corals are accreted with rhythmic structural patterns that serve as valuable sclerochronometers. Annual high- and low-density band couplets, visible in X-radiographs or computed tomography scans, are used to construct age models for paleoclimate reconstructions and to track variability in coral growth over time. In some corals, discrete, anomalously high-density bands, called “stress bands,” preserve information about coral bleaching. However, the mechanisms underlying the formation of coral skeletal density banding remain unclear. Dissepiments—thin, horizontal sheets of calcium carbonate accreted by the coral to support the living polyp—play a key role in the upward growth of the colony. Here, we first conducted a vital staining experiment to test whether dissepiments were accreted with lunar periodicity in Porites coral skeleton, as previously hypothesized. Over 6, 15, and 21 months, dissepiments consistently formed in a 1:1 ratio to the number of full moons elapsed over each study period. We measured dissepiment spacing to reconstruct multiple years of monthly skeletal extension rates in two Porites colonies from Palmyra Atoll and in another from Palau that bleached in 1998 under anomalously high sea temperatures. Spacing between successive dissepiments exhibited strong seasonality in corals containing annual density bands, with narrow (wide) spacing associated with high (low) density, respectively. A high-density “stress band” accreted during the 1998 bleaching event was associated with anomalously low dissepiment spacing and missed dissepiments, implying that thermal stress disrupts skeletal extension. Further, uranium/calcium ratios increased within stress bands, indicating a reduction in the carbonate ion concentration of the coral’s calcifying fluid under stress. Our study verifies the lunar periodicity of dissepiments, provides a mechanistic basis for the formation of annual density bands in Porites, and reveals the underlying cause of high-density stress bands.

Coral skeletons are valuable archives of past ocean conditions. However, interpretation of coral paleotemperature records is confounded by uncertainties associated with single element-ratio thermometers, including Sr/Ca. A new approach, Sr-U, uses U/Ca to constrain the influence of Rayleigh fractionation on Sr/Ca [DeCarlo et al., 2016]. Here, we build on the initial Pacific Porites Sr-U calibration to include multiple Atlantic and Pacific coral genera from multiple coral reef locations spanning a temperature range of 23.15-30.12 °C. Accounting for the wintertime growth cessation of one Bermuda coral, we show that Sr-U is strongly correlated with the average water temperature at each location (r2 = 0.91, P

Fine scale temperature structures, which are commonly found in the top few meters of shallow water columns, may result in deviations of the remotely sensed night-time sea surface temperatures (SST) by the MODIS-Aqua sensor (SSTsat) from the bulk sea surface temperatures (SSTbulk) that they purport to represent. The discrepancies between SSTsat and SSTbulk recorded by temperature loggers at eight stations with bottom depths of 2 - 20 m around the Dongsha Atoll (DSA) between June 2013 and May 2015 were examined. The SSTsat had an average cool bias error of -0.43 ± 0.59°C. The bias error was larger in the warmer (> 26°C) waters which were presumably more strongly stratified. The root mean square error (RMSE) between SSTsat and SSTbulk, ±0.73°C, was 25% larger than that reported in the open northern South China Sea. An operational calibration algorithm was developed to increase the accuracy in the estimation of SSTbulk from SSTsat. In addition to removing the cool bias error, this algorithm also reduced the RMSE to virtually the same level as that found in the open northern South China Sea. With the application of the algorithm, in June 2015, the average SST in the lagoon of the DSA was raised by about 0.5°C to 31.1 ± 0.4°C, and the area of lagoon with SSTbulk above 31°C, the median value of the physiological temperature threshold of reef organisms, was increased by 69% to about three quarters of the lagoon.

Coral reefs are built of calcium carbonate (CaCO3) produced biogenically by a diversity of calcifying plants, animals, and microbes. As the ocean warms and acidifies, there is mounting concern that declining calcification rates could shift coral reef CaCO3 budgets from net accretion to net dissolution. We quantified net ecosystem calcification (NEC) and production (NEP) on Dongsha Atoll, northern South China Sea, over a 2 week period that included a transient bleaching event. Peak daytime pH on the wide, shallow reef flat during the nonbleaching period was ∼8.5, significantly elevated above that of the surrounding open ocean (∼8.0–8.1) as a consequence of daytime NEP (up to 112 mmol C m−2 h−1). Diurnal-averaged NEC was 390 ± 90 mmol CaCO3 m−2 d−1, higher than any other coral reef studied to date despite comparable calcifier cover (25%) and relatively high fleshy algal cover (19%). Coral bleaching linked to elevated temperatures significantly reduced daytime NEP by 29 mmol C m−2 h−1. pH on the reef flat declined by 0.2 units, causing a 40% reduction in NEC in the absence of pH changes in the surrounding open ocean. Our findings highlight the interactive relationship between carbonate chemistry of coral reef ecosystems and ecosystem production and calcification rates, which are in turn impacted by ocean warming. As open-ocean waters bathing coral reefs warm and acidify over the 21st century, the health and composition of reef benthic communities will play a major role in determining on-reef conditions that will in turn dictate the ecosystem response to climate change.

This study investigates the impact of extreme heat wave events on long‐lived massive corals (Porites spp.) from the central Saudi Arabian Red Sea using trace element (Sr/Ca, Li/Mg, Mg/Ca, U/Ca, B/Ca, and Li/Ca) records preserved in the coral skeleton for the period between 1992 and 2012. Prior to 1998, the trace element records show strong correlations with sea surface temperature. However, during the prolonged high temperature phase associated with the 1998 El Niño event, the seasonal trace element signals were disrupted, which also coincided with a reduction in extension rates. This disruption in normally highly correlated seasonal trace element ratios‐sea surface temperature relationships was unusually long, lasting for approximately 2 years in the inner‐shelf reef site and nearly 4 years in the outer‐shelf reef site. Although the seasonal signal of trace element ratios in both cores eventually stabilized, for the inner‐shelf core the amplitude and absolute values in most trace element ratios remained significantly different compared to pre‐1998 levels. This suggests that prolonged thermal stress can induce subtle but potentially long‐lasting physiological changes that affect the elemental composition of the coral's calcifying fluid. The lack of indication of stress in the core records during later bleaching events (2003, 2005, and 2010) suggests that some of these physiological changes could have induced increased thermal tolerance, particularly for inner‐shelf corals, lending support to the capacity for corals to acclimatize.Plain Language SummaryThe future of coral reefs is jeopardized by global warming, particularly by marine heat waves and mass bleaching events, as evidenced by the 2016 and 2017 events. In this study the geochemical composition of the skeleton of two long‐lived massive corals from the Red Sea was used to evaluate possible long‐term acclimatization or changes in sensitivity to thermal stress. We detected a clear disturbance in the biomineralization process of the two corals following the 1998 bleaching event. However, posterior thermal stress events of similar magnitude were not registered in the skeletal growth or geochemical signature of the same corals hinting toward a possible long‐term acclimatization following the exposure to the 1998 event. published

Ocean warming threatens the functioning of coral reef ecosystems by inducing mass coral bleaching and mortality events. The link between temperature and coral bleaching is now well-established based on observations that mass bleaching events usually occur when seawater temperatures are anomalously high. However, times of high heat stress but without coral bleaching are equally important because they can inform an understanding of factors that regulate temperature-induced bleaching. Here, we investigate the absence of mass coral bleaching on the Great Barrier Reef (GBR) during austral summer 2004. Using four gridded sea surface temperature data products, validated with in situ temperature loggers, we demonstrate that the summer of 2004 was among the warmest summers of the satellite era (1982–2017) on the GBR. At least half of the GBR experienced temperatures that were high enough to initiate bleaching in other years, yet mass bleaching was not reported during 2004. The absence of bleaching is not fully explained by wind speed or cloud cover. Rather, 2004 is clearly differentiated from bleaching years by the slow speed of the East Australian Current (EAC) offshore of the GBR. An anomalously slow EAC during summer 2004 may have dampened the upwelling of nutrient-rich waters onto the GBR shelf, potentially mitigating bleaching due to the lower susceptibility of corals to heat stress in low-nutrient conditions. Although other factors such as irradiance or acclimatization may have played a role in the absence of mass bleaching, 2004 remains a key case study for demonstrating the dynamic nature of coral responses to marine heatwaves.

Reef-building corals typically live close to the upper limits of their thermal tolerance and even small increases in summer water temperatures can lead to bleaching and mortality. Projections of coral reef futures based on forecasts of ocean temperatures indicate that by the end of this century, corals will experience their current thermal thresholds annually, which would lead to the widespread devastation of coral reef ecosystems. Here, we use skeletal cores of long-lived Porites corals collected from 14 reefs across the northern Great Barrier Reef, the Coral Sea, and New Caledonia to evaluate changes in their sensitivity to heat stress since 1815. High-density ‘stress bands’—indicative of past bleaching—first appear during a strong pre-industrial El Niño event in 1877 but become significantly more frequent in the late twentieth and early twenty-first centuries in accordance with rising temperatures from anthropogenic global warming. However, the proportion of cores with stress bands declines following successive bleaching events in the twenty-first century despite increasing exposure to heat stress. Our findings demonstrate an increase in the thermal tolerance of reef-building corals and offer a glimmer of hope that at least some coral species can acclimatize fast enough to keep pace with global warming.

Ocean acidification poses a serious threat to marine calcifying organisms, yet experimental and field studies have found highly diverse responses among species and environments. Our understanding of the underlying drivers of differential responses to ocean acidification is currently limited by difficulties in directly observing and quantifying the mechanisms of bio‐calcification. Here, we present Raman spectroscopy techniques for characterizing the skeletal mineralogy and calcifying fluid chemistry of marine calcifying organisms such as corals, coralline algae, foraminifera, and fish (carbonate otoliths). First, our in vivo Raman technique is the ideal tool for investigating non‐classical mineralization pathways. This includes calcification by amorphous particle attachment, which has recently been controversially suggested as a mechanism by which corals resist the negative effects of ocean acidification. Second, high‐resolution ex vivo Raman mapping reveals complex banding structures in the mineralogy of marine calcifiers, and provides a tool to quantify calcification responses to environmental variability on various timescales from days to years. We describe the new insights into marine bio‐calcification that our techniques have already uncovered, and we consider the wide range of questions regarding calcifier responses to global change that can now be proposed and addressed with these new Raman spectroscopy tools., Newly developed in vivo and high‐resolution Raman spectroscopy analyses of marine calcifying organisms enable characterization of their mineralogy and calcification mechanisms. Insights from these new Raman spectroscopy tools are helping us understand the sensitivity of these important organisms to ocean warming and acidification.

The sensitivity of corals to ocean acidification depends on the extent to which they can buffer their calcifying fluid aragonite saturation state (Ωcf) from declines in seawater pH. While the seasonal response of the coral calcifying fluid Ωcf to seawater pH has been studied previously, relatively little is known about Ωcf dynamics on shorter (daily) timescales, particularly whether it is sensitive to seawater pH. Here, we use alizarin dye to mark 4 days of skeletal growth in the corals Acropora nasuta and Pocillopora damicornis living in situ on Ningaloo Reef in Western Australia. Exploiting newly developed confocal Raman spectroscopy techniques, we imaged the alizarin stains and quantified Ωcf between them. We report the first observations of diurnal Ωcf cycles, which were found in both species. Our results are consistent with either external control of Ωcf by seawater pH or light, or alternatively that Ωcf follows an endogenous circadian rhythm.

Natural variability in pH in the diffusive boundary layer (DBL), the discrete layer of seawater between bulk seawater and the outer surface of organisms, could be an important factor determining the response of corals and coralline algae to ocean acidification (OA). Here, two corals with different morphologies and one coralline alga were maintained under two different regimes of flow velocities, pH, and light intensities in a 12 flumes experimental system for a period of 27 weeks. We used a combination of geochemical proxies, physiological and micro-probe measurements to assess how these treatments affected the conditions in the DBL and the response of organisms to OA. Overall, low flow velocity did not ameliorate the negative effect of low pH and therefore did not provide a refugia from OA. Flow velocity had species-specific effects with positive effects on calcification for two species. pH in the calcifying fluid (pHcf) was reduced by low flow in both corals at low light only. pHcf was significantly impacted by pH in the DBL for the two species capable of significantly modifying pH in the DBL. The dissolved inorganic carbon in the calcifying fluid (DICcf) was highest under low pH for the corals and low flow for the coralline, while the saturation state in the calcifying fluid and its proxy (FWHM) were generally not affected by the treatments. This study therefore demonstrates that the effects of OA will manifest most severely in a combination of lower light and lower flow habitats for sub-tropical coralline algae. These effects will also be greatest in lower flow habitats for some corals. Together with existing literature, these findings reinforce that the effects of OA are highly context dependent, and will differ greatly between habitats, and depending on species composition.

Measurements of B/Ca ratios in marine carbonates have been suggested to record seawater carbonate chemistry, however experimental calibration of such proxies based on inorganic partitioning remains limited. Here we conducted a series of synthetic aragonite precipitation experiments to evaluate the factors influencing the partitioning of B/Ca between aragonite and seawater. Our results indicate that the B/Ca ratio of synthetic aragonites depends primarily on the relative concentrations of borate and carbonate ions in the solution from which the aragonite precipitates; not on bicarbonate concentration as has been previously suggested. The influence of temperature was not significant over the range investigated (20–40 °C), however, partitioning may be influenced by saturation state (and/or growth rate). Based on our experimental results, we suggest that aragonite B/Ca ratios can be utilized as a proxy of [CO 3 2– ]. Boron isotopic composition (δ 11 B) is an established pH proxy, thus B/Ca and δ 11 B together allow the full carbonate chemistry of the solution from which the aragonite precipitated to be calculated. To the extent that aragonite precipitation by marine organisms is affected by seawater chemistry, B/Ca may also prove useful in reconstructing seawater chemistry. A simplified boron purification protocol based on amberlite resin and the organic buffer TRIS is also described.

Coral skeletons archive past climate variability with unrivaled temporal resolution. However, extraction of accurate temperature information from coral skeletons has been limited by “vital effects,” which confound, and sometimes override, the temperature dependence of geochemical proxies. We present a new approach to coral paleothermometry based on results of abiogenic precipitation experiments interpreted within a framework provided by a quantitative model of the coral biomineralization process. DeCarlo et al. (2015a) investigated temperature and carbonate chemistry controls on abiogenic partitioning of Sr/Ca and U/Ca between aragonite and seawater and modeled the sensitivity of skeletal composition to processes occurring at the site of calcification. The model predicts that temperature can be accurately reconstructed from coral skeleton by combining Sr/Ca and U/Ca ratios into a new proxy, which we refer to hereafter as the Sr-U thermometer. Here we test the model predictions with measured Sr/Ca and U/Ca ratios of 14 Porites sp. corals collected from the tropical Pacific Ocean and the Red Sea, with a subset also analyzed using the boron isotope (δ11B) pH proxy. Observed relationships among Sr/Ca, U/Ca, and δ11B agree with model predictions, indicating that the model accounts for the key features of the coral biomineralization process. By calibrating to instrumental temperature records, we show that Sr-U captures 93% of mean annual temperature variability (26–30°C) and has a standard deviation of prediction of 0.5°C, compared to 1°C using Sr/Ca alone. The Sr-U thermometer may offer significantly improved reliability for reconstructing past ocean temperatures from coral skeletons.

Coral Sr/Ca is widely used to reconstruct past ocean temperatures. However, some studies report different Sr/Ca-temperature relationships for conspecifics on the same reef, with profound implications for interpretation of reconstructed temperatures. We assess whether these differences are attributable to small-scale oceanographic variability or “vital effects” associated with coral calcification and quantify the effect of intercolony differences on temperature estimates and uncertainties. Sr/Ca records from four massive Porites colonies growing on the east and west sides of Jarvis Island, central equatorial Pacific, were compared with in situ logger temperatures spanning 2002–2012. In general, Sr/Ca captured the occurrence of interannual sea surface temperature events but their amplitude was not consistently recorded by any of the corals. No long-term trend was identified in the instrumental data, yet Sr/Ca of one coral implied a statistically significant cooling trend while that of its neighbor implied a warming trend. Slopes of Sr/Ca-temperature regressions from the four different colonies were within error, but offsets in mean Sr/Ca rendered the regressions statistically distinct. Assuming that these relationships represent the full range of Sr/Ca-temperature calibrations in Jarvis Porites, we assessed how well Sr/Ca of a nonliving coral with an unknown Sr/Ca-temperature relationship can constrain past temperatures. Our results indicate that standard error of prediction methods underestimate the actual error as we could not reliably reconstruct the amplitude or frequency of El Nino–Southern Oscillation events as large as ± 2°C. Our results underscore the importance of characterizing the full range of temperature-Sr/Ca relationships at each study site to estimate true error.

Identifying the long-term effects of ocean acidification (OA) and global warming on coral calcification has proven elusive yet has major implications for the continuing viability of coral reefs in the face of climate change. Here we address this question using seasonally and annually resolved boron proxies (11B/10B and B/Ca) of calcifying fluid (cf) pHcf and carbonate ion concentrations ([CO 3 2 − ]cf) preserved in a long-lived Porites coral from the Great Barrier Reef (GBR). From 1939 to 2013 we find that the coral pHcf closely followed the decline in seawater pH of ∼0.1 units, but at a reduced rate of ∼60%, indicative of biological buffering. Of the decline in pHcf ∼82% is attributed to OA and ∼17% to the ∼0.5 °C long-term warming observed over this period. This long-term warming induced change in pHcf is consistent with the much larger seasonally modulated changes in pHcf where ∼4 to 6 °C seasonal changes in temperatures are accompanied by relatively large antithetic ∼0.1 changes in pHcf. Furthermore, we find that although the supply of dissolved inorganic carbon (DIC) of the coral cf has remained at constant elevated levels of ∼1.5 × seawater, there has been a significant long-term decline (4 to 11%) in the [CO 3 2 − ]cf, due primarily to the OA-induced change in pHcf. This decline in [CO 3 2 − ]cf, a critical parameter controlling calcification, is thus likely responsible for the ∼15% decline in coral calcification observed since 1939 and across the GBR generally.

The processes that occur at the micro-scale site of calcification are fundamental to understanding the response of coral growth in a changing world. However, our mechanistic understanding of chemical processes driving calcification is still evolving. Here, we report the results of a long-term in situ study of coral calcification rates, photo-physiology, and calcifying fluid (cf) carbonate chemistry (using boron isotopes, elemental systematics, and Raman spectroscopy) for seven species (four genera) of symbiotic corals growing in their natural environments at tropical, subtropical, and temperate locations in Western Australia (latitudinal range of ~11°). We find that changes in net coral calcification rates are primarily driven by pHcf and carbonate ion concentration [ CO 3 2 - ]cf in conjunction with temperature and DICcf . Coral pHcf varies with latitudinal and seasonal changes in temperature and works together with the seasonally varying DICcf to optimize [ CO 3 2 - ]cf at species-dependent levels. Our results indicate that corals shift their pHcf to adapt and/or acclimatize to their localized thermal regimes. This biological response is likely to have critical implications for predicting the future of coral reefs under CO2 -driven warming and acidification.