Assisted sexual coral recruits show high thermal tolerance to the 2023 Caribbean mass bleaching event.

Assisted sexual coral propagation, resulting in greater genet diversity via genetic recombination, has been hypothesized to lead to more adaptable and, hence, resilient restored populations compared to more common clonal techniques. Coral restoration efforts have resulted in substantial populations of 'Assisted sexual Recruits' (i.e., juvenile corals derived from assisted sexual reproduction; AR) of multiple species outplanted to reefs or held in in situ nurseries across many locations in the Caribbean. These AR populations provided context to evaluate their relative resilience compared to co-occurring coral populations during the 2023 marine heat wave of unprecedented duration and intensity that affected the entire Caribbean. Populations of six species of AR, most ranging in age from 1-4 years, were surveyed across five regions during the mass bleaching season in 2023 (Aug-Dec), alongside co-occurring groups of corals to compare prevalence of bleaching and related mortality. Comparison groups included conspecific adult colonies as available, but also the extant co-occurring coral assemblages in which conspecifics were rare or lacking, as well as small, propagated coral fragments. Assisted sexual recruits had significantly lower prevalence of bleaching impacts (overall pooled ~ 10%) than conspecific coral populations typically comprised of larger colonies (~ 60-100% depending on species). In addition, small corals derived from fragmentation (rather than sexual propagation) in two regions showed bleaching susceptibility intermediate between AR and wild adults. Overall, AR exhibited high bleaching resistance under heat stress exposure up to and exceeding Degree Heating Weeks of 20°C-weeks. As coral reefs throughout the globe are subject to increasingly frequent and intense marine heatwaves, restoration activities that include sexual reproduction and seeding can make an important contribution to sustain coral populations.

Severe cold-water bleaching of a deep-water reef underscores future challenges for Mesophotic Coral Ecosystems.

Elevated sea surface temperatures are causing an increase in coral bleaching events worldwide, and represent an existential threat to coral reefs. Early studies of Mesophotic Coral Ecosystems (MCEs) highlighted their potential as thermal refuges for shallow-water coral species in the face of predicted 21st century warming. However, recent genetic evidence implies that limited ecological connectivity between shallow- and deep-water coral communities inhibits their effectiveness as refugia; instead MCEs host distinct endemic communities that are ecologically significant in and of themselves. In either scenario, understanding the response of MCEs to climate change is critical given their ecological significance and widespread global distribution. Such an understanding has so far eluded the community, however, because of the challenges associated with long-term field monitoring, the stochastic nature of climatic events that drive bleaching, and the paucity of deep-water observations. Here we document the first observed cold-water bleaching of a mesophotic coral reef at Clipperton Atoll, a remote Eastern Tropical Pacific (ETP) atoll with high coral cover and a well-developed MCE. The severe bleaching (>70 % partially or fully bleached coral cover at 32 m depth) was driven by an anomalously shallow thermocline, and highlights a significant and previously unreported challenge for MCEs. Prompted by these observations, we compiled published cold-water bleaching events for the ETP, and demonstrate that the timing of past cold-water bleaching events in the ETP coincides with decadal oscillations in mean zonal wind strength and thermocline depth. The latter observation suggests any future intensification of easterly winds in the Pacific could be a significant concern for its MCEs. Our observations, in combination with recent reports of warm-water bleaching of Red Sea and Indian Ocean MCEs, highlight that 21st century MCEs in the Eastern Pacific face a two-pronged challenge: warm-water bleaching from above, and cold-water bleaching from below.

Corals that survive repeated thermal stress show signs of selection and acclimatization.

Climate change is transforming coral reefs by increasing the frequency and intensity of marine heatwaves, often leading to coral bleaching and mortality. Coral communities have demonstrated modest increases in thermal tolerance following repeated exposure to moderate heat stress, but it is unclear whether these shifts represent acclimatization of individual colonies or mortality of thermally susceptible individuals. For corals that survive repeated bleaching events, it is important to understand how past bleaching responses impact future growth potential. Here, we track the bleaching responses of 1,832 corals in leeward Maui through multiple marine heatwaves and document patterns of coral growth and survivorship over a seven-year period. While we find limited evidence of acclimatization at population scales, we document reduced bleaching over time in specific individuals that is indicative of acclimatization, primarily in the stress-tolerant taxa Porites lobata. For corals that survived both bleaching events, we find no relationship between bleaching response and coral growth in three of four taxa studied. This decoupling suggests that coral survivorship is a better indicator of future growth than is a coral's bleaching history. Based on these results, we recommend restoration practitioners in Hawai'i focus on colonies of Porites and Montipora with a proven track-record of growth and survivorship, rather than devote resources toward identifying and cultivating bleaching-resistant phenotypes in the lab. Survivorship followed a latitudinal thermal stress gradient, but because this gradient was small, it is likely that local environmental factors also drove differences in coral performance between sites. Efforts to reduce human impacts at low performing sites would likely improve coral survivorship in the future.

Connectivity modelling identifies sources and sinks of coral recruitment within reef clusters.

Connectivity aids the recovery of populations following disturbances, such as coral bleaching and tropical cyclones. Coral larval connectivity is a function of physical connectivity and larval behaviour. In this study, we used OceanParcels, a particle tracking simulator, with 2D and 3D velocity outputs from a high resolution hydrodynamic-biogeochemical marine model (RECOM) to simulate the dispersal and settlement of larvae from broadcast spawning Acropora corals in the Moore Reef cluster, northern Great Barrier Reef, following the annual spawning events in 2015, 2016 and 2017. 3D velocity simulations showed 19.40-68.80% more links and sinks than those of 2D simulations. Although the patterns of connectivity among sites vary over days and years, coral larvae consistently dispersed from east to west in the cluster domain, with some sites consistently acting as sources or sinks for local larval recruitment. Results can inform coral reef intervention plans for climate change, such as the design of marine protected areas and the deployment of proposed interventions within reef clusters. For example, the wider benefits of interventions (e.g., deployment of heat adapted corals) may be optimised when deployed at locations that are a source of larvae to others within comparable habitats across the reef cluster.

Cumulative risk of future bleaching for the world's coral reefs.

Spatial and temporal patterns of future coral bleaching are uncertain, hampering global conservation efforts to protect coral reefs against climate change. Our analysis of daily projections of ocean warming establishes the severity, annual duration, and onset of severe bleaching risk for global coral reefs this century, pinpointing vital climatic refugia. We show that low-latitude coral regions are most vulnerable to thermal stress and will experience little reprieve from climate mitigation. By 2080, coral bleaching is likely to start on most reefs in spring, rather than late summer, with year-round bleaching risk anticipated to be high for some low-latitude reefs regardless of global efforts to mitigate harmful greenhouse gasses. By identifying Earth's reef regions that are at lowest risk of accelerated bleaching, our results will prioritize efforts to limit future loss of coral reef biodiversity.

Should I stay or should I go? Coral bleaching from the symbionts' perspective.

Coral bleaching, the stress-induced breakdown of coral-algal symbiosis, threatens reefs globally. Paradoxically, despite adverse fitness effects, corals bleach annually, even outside of abnormal temperatures. This generally occurs shortly after the once-per-year mass coral spawning. Here, we propose a hypothesis linking annual coral bleaching and the transmission of symbionts to the next generation of coral hosts. We developed a dynamic model with two symbiont growth strategies, and found that high sexual recruitment and low adult coral survivorship and growth favour bleaching susceptibility, while the reverse promotes bleaching resilience. Otherwise, unexplained trends in the Indo-Pacific align with our hypothesis, where reefs and coral taxa exhibiting higher recruitment are more bleaching susceptible. The results from our model caution against interpreting potential shifts towards more bleaching-resistant symbionts as evidence of climate adaptation-we predict such a shift could also occur in declining systems experiencing low recruitment rates, a common scenario on today's reefs.

Mitigation of Vibrio coralliilyticus-induced coral bleaching through bacterial dysbiosis prevention by Ruegeria profundi.

Vibrio species are prevalent in ocean ecosystems, particularly Vibrio coralliilyticus, and pose a threat to corals and other marine organisms under global warming conditions. While microbiota manipulation is considered for coral disease management, understanding the role of commensal bacteria in stress resilience remains limited. Here, a single bacterial species (Ruegeria profundi) rather than a consortium of native was used to combat pathogenic V. coralliilyticus and protect corals from bleaching. R. profundi showed therapeutic activity in vivo, preventing a significant reduction in bacterial diversity in bleached corals. Notably, the structure of the bacterial community differed significantly among all the groups. In addition, compared with the bleached corals caused by V. coralliilyticus, the network analysis revealed that complex interactions and positive correlations in the bacterial community of the R. profundi protected non-bleached corals, indicating R. profundi's role in fostering synergistic associations. Many genera of bacteria significantly increased in abundance during V. coralliilyticus infection, including Vibrio, Alteromonas, Amphritea, and Nautella, contributing to the pathogenicity of the bacterial community. However, R. profundi effectively countered the proliferation of these genera, promoting potential probiotic Endozoicomonas and other taxa, while reducing the abundance of betaine lipids and the type VI section system of the bacterial community. These changes ultimately influenced the interactive relationships among symbionts and demonstrated that probiotic R. profundi intervention can modulate coral-associated bacterial community, alleviate pathogenic-induced dysbiosis, and preserve coral health. These findings elucidated the relationship between the behavior of the coral-associated bacterial community and the occurrence of pathological coral bleaching.IMPORTANCEChanges in the global climate and marine environment can influence coral host and pathogen repartition which refers to an increased likelihood of pathogen infection in hosts. The risk of Vibrio coralliilyticus-induced coral disease is significantly heightened, primarily due to its thermos-dependent expression of virulent and populations. This study investigates how coral-associated bacterial communities respond to bleaching induced by V. coralliilyticus. Our findings demonstrate that Ruegeria profundi exhibits clear evidence of defense against pathogenic bacterial infection, contributing to the maintenance of host health and symbiont homeostasis. This observation suggests that bacterial pathogens could cause dysbiosis in coral holobionts. Probiotic bacteria display an essential capability in restructuring and manipulating coral-associated bacterial communities. This restructuring effectively reduces bacterial community virulence and enhances the pathogenic resistance of holobionts. The study provides valuable insights into the correlation between the health status of corals and how coral-associated bacterial communities may respond to both pathogens and probiotics.

Algal symbiont diversity in Acropora muricata from the extreme reef of Bouraké associated with resistance to coral bleaching.

Widespread coral bleaching has generally been linked to high water temperatures at larger geographic scales. However, the bleaching response can be highly variable among individual of the same species, between different species, and across localities; what causes this variability remains unresolved. Here, we tracked bleached and non-bleached colonies of Acropora muricata to see if they recovered or died following a stress event inside the semi-enclosed lagoon of Bouraké (New Caledonia), where corals are long-term acclimatized to extreme conditions of temperature, pH and dissolved oxygen, and at a nearby control reef where conditions are more benign. We describe Symbiodiniaceae community changes based on next-generation sequencing of the ITS2 marker, metabolic responses, and energetic reserve measures (12 physiological traits evaluated) during the La Niña warm and rainy summer in 2021. Widespread coral bleaching (score 1 and 2 on the coral colour health chart) was observed only in Bouraké, likely due to the combination of the high temperatures (up to 32°C) and heavy rain. All colonies (i.e., Bouraké and reference site) associated predominantly with Symbiodinaceae from the genera Cladocopium. Unbleached colonies in Bouraké had a specific ITS2-type profile (proxies for Symbiodiniaceae genotypes), while the bleached colonies in Bouraké had the same ITS2-type profile of the reef control colonies during the stress event. After four months, the few bleached colonies that survived in Bouraké (B2) acquired the same ITS2 type profiles of the unbleached colonies in Bouraké. In terms of physiological performances, all bleached corals showed metabolic depression (e.g., Pgross and Rdark). In contrast, unbleached colonies in Bouraké maintained higher metabolic rates and energetic reserves compared to control corals. Our study suggests that Acropora muricata enhanced their resistance to bleaching thanks to specific Symbiodiniaceae associations, while energetic reserves may increase their resilience after stress.

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

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

Global algal outbreaks are crowding out corals.

Reefs already stressed by bleaching are being smothered by crust-forming red algae.

Mesophotic coral bleaching associated with changes in thermocline depth.

As global temperatures continue to rise, shallow coral reef bleaching has become more intense and widespread. Mesophotic coral ecosystems reside in deeper (30-150 m), cooler water and were thought to offer a refuge to shallow-water reefs. Studies now show that mesophotic coral ecosystems instead have limited connectivity with shallow corals but host diverse endemic communities. Given their extensive distribution and high biodiversity, understanding their susceptibility to warming oceans is imperative. In this multidisciplinary study of an atoll in the Chagos Archipelago in the central Indian Ocean, we show evidence of coral bleaching at 90 m, despite the absence of shallow-water bleaching. We also show that the bleaching was associated with sustained thermocline deepening driven by the Indian Ocean Dipole, which might be further enhanced by internal waves whose influence varied at a sub-atoll scale. Our results demonstrate the potential vulnerability of mesophotic coral ecosystems to thermal stress and highlight the need for oceanographic knowledge to predict bleaching susceptibility and heterogeneity.

Standardization of in situ coral bleaching measurements highlights the variability in responses across genera, morphologies, and regions.

Marine heatwaves and regional coral bleaching events have become more frequent and severe across the world's oceans over the last several decades due to global climate change. Observational studies have documented spatiotemporal variation in the responses of reef-building corals to thermal stress within and among taxa across geographic scales. Although many tools exist for predicting, detecting, and quantifying coral bleaching, it remains difficult to compare bleaching severity (e.g., percent cover of bleached surface areas) among studies and across species or regions. For this review, we compiled over 2,100 in situ coral bleaching observations representing 87 reef-building coral genera and 250 species of common morphological groups from a total of 74 peer-reviewed scientific articles, encompassing three broad geographic regions (Atlantic, Indian, and Pacific Oceans). While bleaching severity was found to vary by region, genus, and morphology, we found that both genera and morphologies responded differently to thermal stress across regions. These patterns were complicated by (i) inconsistent methods and response metrics across studies; (ii) differing ecological scales of observations (i.e., individual colony-level vs. population or community-level); and (iii) temporal variability in surveys with respect to the onset of thermal stress and the chronology of bleaching episodes. To improve cross-study comparisons, we recommend that future surveys prioritize measuring bleaching in the same individual coral colonies over time and incorporate the severity and timing of warming into their analyses. By reevaluating and standardizing the ways in which coral bleaching is quantified, researchers will be able to track responses to marine heatwaves with increased rigor, precision, and accuracy.

Turbidity buffers coral bleaching under extreme wind and rainfall conditions.

Coral reefs in turbid waters have been hypothesized to be a refuge from climate change. These naturally occurring communities were brought into the spotlight because some of their species exhibited record levels of resistance to marine heatwaves (MHWs) by disturbance-tolerant corals. However, long-term monitoring data on the drivers of coral bleaching in these extreme reef habitats are scarce. Here, we describe the population structure and bleaching rates of a widespread and resilient coral (Siderastrea stellata). We examine the links between environmental factors, namely, rainfall, wind speed, turbidity, solar irradiance, sea surface temperature, MHWs, and coral bleaching status under the worst recorded drought cycle in the Tropical South Atlantic (2013-2015). We examined 2880 colonies, most of which (∼93%) fit in the size group of 2-10 cm, with a small number (∼1%) of larger and older colonies (>20 cm). The results indicated the absence of MHWs and normal sea surface temperature variations (between 26.6 °C and 29.3 °C), however, we detected an extreme rainfall deficit (30-40% less annual volume precipitation). In general, a high proportion (44-84%) of bleached colonies was found throughout the months when turbidity decreased. Siderastrea is the only reef-building coral that comprises this seascape with encrusting and low-relief colonies. During drought periods, cloudiness is reduced, turbidity and wind speed are reduced, and solar irradiance increase, driving coral bleaching in turbid reefs. However, episodic rainfall and higher wind speeds increase turbidity and decrease coral bleaching. Our hypothesis is that turbidity decreases during drought periods which increases bleaching risk to corals even without thermal stress. Our results suggest that turbidity may have related to wind and rainfall to provoke the coral bleaching phenomenon.

Integrated metagenomic and metaproteomic analyses reveal bacterial micro-ecological mechanisms in coral bleaching.

IMPORTANCE: Coral reefs worldwide are facing rapid decline due to coral bleaching. However, knowledge of the physiological characteristics and molecular mechanisms of coral symbionts respond to stress is scarce. Here, metagenomic and metaproteomic approaches were utilized to shed light on the changes in the composition and functions of coral symbiotic bacteria during coral bleaching. The results demonstrated that coral bleaching significantly affected the composition of symbionts, with bacterial communities dominating in bleached corals. Through differential analyses of gene and protein expression, it becomes evident that symbionts experience functional disturbances in response to heat stress. These disturbances result in abnormal energy metabolism, which could potentially compromise the health and resilience of the symbionts. Furthermore, our findings highlighted the highly diverse microbial communities of coral symbionts, with beneficial bacteria providing critical services to corals in stress responses and pathogenic bacteria driving coral bleaching. This study provides comprehensive insights into the complex response mechanisms of coral symbionts under heat stress from the micro-ecological perspective and offers fundamental data for future monitoring of coral health.

Optimized spatial and temporal pattern for coral bleaching heat stress alerts for China's coral reefs.

Most studies on coral bleaching alerts use common Degree Heating Week (DHW) thresholds; however, these may underestimate historical patterns of heat stress for coral reef ecosystems. Taking an optimized DHW threshold for coral bleaching alerts for Coral Reef Watch (CRW) and Coral Reef Temperature Anomaly Database (CoRTAD) products, we analyzed the precise spatial and temporal pattern of heat stress on China's coral reefs from 2010 to 2021 in the South China Sea (SCS) and the Beibu Gulf (BG). We compared acute heat stress using common and optimized thresholds. Results indicated that the ocean warming rate in 2010-2021 was approximately 0.43 ± 0.22 °C/10a, showing a significant increase in the northern SCS and the BG. More severe bleaching events were predicted by the optimized thresholds and the high-frequency areas were mainly in the northern SCS. The number and intensity of years with severe heat stress anomalies was in the order 2020 > 2014 > 2010 > 2015. Heat stress duration was the longest in the Xisha Islands among offshore archipelagos, and longest in 2020-2021 in Weizhou Island in BG in the relative high-latitude inshore reefs. These abnormal events were mainly caused by El Niño, but La Niña was also involved in 2020.

Genomic exploration of coral-associated bacteria: identifying probiotic candidates to increase coral bleaching resilience in Galaxea fascicularis.

BACKGROUND: Reef-building corals are acutely threatened by ocean warming, calling for active interventions to reduce coral bleaching and mortality. Corals associate with a wide diversity of bacteria which can influence coral health, but knowledge of specific functions that may be beneficial for corals under thermal stress is scant. Under the oxidative stress theory of coral bleaching, bacteria that scavenge reactive oxygen (ROS) or nitrogen species (RNS) are expected to enhance coral thermal resilience. Further, bacterial carbon export might substitute the carbon supply from algal photosymbionts, enhance thermal resilience and facilitate bleaching recovery. To identify probiotic bacterial candidates, we sequenced the genomes of 82 pure-cultured bacteria that were isolated from the emerging coral model Galaxea fascicularis. RESULTS: Genomic analyses showed bacterial isolates were affiliated with 37 genera. Isolates such as Ruegeria, Muricauda and Roseovarius were found to encode genes for the synthesis of the antioxidants mannitol, glutathione, dimethylsulfide, dimethylsulfoniopropionate, zeaxanthin and/or ß-carotene. Genes involved in RNS-scavenging were found in many G. fascicularis-associated bacteria, which represents a novel finding for several genera (including Pseudophaeobacter). Transporters that are suggested to export carbon (semiSWEET) were detected in seven isolates, including Pseudovibrio and Roseibium. Further, a range of bacterial strains, including strains of Roseibium and Roseovarius, revealed genomic features that may enhance colonisation and association of bacteria with the coral host, such as secretion systems and eukaryote-like repeat proteins. CONCLUSIONS: Our work provides an in-depth genomic analysis of the functional potential of G. fascicularis-associated bacteria and identifies novel combinations of traits that may enhance the coral's ability to withstand coral bleaching. Identifying and characterising bacteria that are beneficial for corals is critical for the development of effective probiotics that boost coral climate resilience. Video Abstract.

Sedimentary Vibrio Blooms in the Xisha Islands May Associate with the 2020 Coral Bleaching Event.

Coral reefs are among the most biodiverse ecosystems, providing habitats for various organisms. Studies on coral bleaching have been increasing recently, but little is known about the distribution and community assembly of coral pathogenic bacteria (e.g., several Vibrio species). We elucidated the distribution pattern and interaction relationships of total bacteria and Vibrio spp. in sediments from the Xisha Islands, which are characterized by their high coverage and diversity of coral resources. Vibrio spp. showed significantly higher relative abundance values in the Xisha Islands (1.00 × 108 copies/g) than in other areas (approximately 1 × 104 to 9.04 × 105 copies/g), indicating that the coral bleaching event of 2020 may have promoted the bloom of vibrios. A spatial shift in community composition was observed between the northern (Photobacterium rosenbergii and Vibrio ponticus) and southern (Vibrio ishigakensis and Vibrio natriegens) sites, accompanied by a clear distance-decay pattern. The spatial distance and coral species (e.g., Acroporidae and Fungiidae) had much greater correlations with the Vibrio community than did environmental factors. However, complex mechanisms may exist in the community assembly of Vibrio spp. due to the large proportion of unexplained variation. Stochastic processes may play an important role, as shown by the neutral model. Vibrio harveyi had the highest relative abundance (77.56%) and niche breadth, compared to other species, and it was negatively correlated with Acroporidae, likely reflecting its strong competitive ability and adverse effects on specific corals. Our study provides insights into the bloom and underlying assembly mechanisms of sedimentary vibrios in the Xisha Islands, thereby contributing to identify the potential indicator of coral bleaching and provide inspiration for the environmental management of coral reef areas. IMPORTANCE Coral reefs exert important roles in maintaining the sustainability of marine ecosystems but decline worldwide due to various drivers, especially pathogenic microorganisms. Here, we investigated the distribution pattern and interactions of total bacteria and Vibrio spp. in the sediments from Xisha Islands during the coral bleaching event of 2020. Our results showed that the abundances of Vibrio (1.00 × 108 copies/g) were high across the whole sites, indicating the bloom of sedimentary Vibrio spp. Coral pathogenic Vibrio species were abundant in the sediments, likely reflecting adverse effects on several kinds of corals. The compositions of the Vibrio spp. were separated by geographical location, which was mainly attributable to the spatial distance and coral species. Overall, this work contributes by providing evidence for the outbreak of coral pathogenic vibrios. The pathogenic mechanism of the dominant species (especially V. harveyi) should be comprehensively considered by laboratory infection experiments in the future.

The 2022 summer marine heatwaves and coral bleaching in China's Greater Bay Area.

From July to August 2022, scleractinian coral communities in China's Greater Bay Area (GBA) in the northern South China Sea (nSCS) experienced an unprecedented bleaching event, despite the fact that coral communities in this area are often considered coral thermal refugia due to their high latitude distribution. Field surveys of six sites covering three main coral distribution areas of the GBA revealed that coral bleaching occurred at all sites. Bleaching was more severe in shallow water (1-3 m) than in deep water (4-6 m), as indicated by both percent bleached cover (51.80 ± 10.04% vs. 7.09 ± 7.37%) and bleached colonies (45.86 ± 11.22% vs. 6.58 ± 6.53%). Coral genera Acropora, Favites, Montipora, Platygyra, Pocillopora, and Porites showed high susceptibility to bleaching, and Acropora and Pocillopora suffered high post-bleaching mortality. In the three areas surveyed, analysis of oceanographic data detected marine heatwaves (MHWs) during the summer, with mean intensities between 1.62 and 1.97 °C and durations between 5 and 22 days. These MHWs were primarily driven by increased shortwave radiation due to strong western Pacific Subtropical High (WPSH), combined with reduced mixing between the surface and deep upwelling waters due to reduced wind speed. Comparing with histological oceanographic data showed that the 2022 MHWs were unprecedented, and there was a significant increase in the frequency, intensity, and total days of MHWs during 1982-2022. Furthermore, the heterogeneous distribution of summer MHW characteristics indicates that the coastal upwelling may modulate the spatial distribution of summer MHWs in nSCS through its cooling effect. Overall, our study indicates that MHWs may have affected the structure of the subtropical coral communities in the nSCS, and impaired their potential as thermal refugia.

Baseline dynamics of Symbiodiniaceae genera and photochemical efficiency in corals from reefs with different thermal histories.

Ocean warming and marine heatwaves induced by climate change are impacting coral reefs globally, leading to coral bleaching and mortality. Yet, coral resistance and resilience to warming are not uniform across reef sites and corals can show inter- and intraspecific variability. To understand changes in coral health and to elucidate mechanisms of coral thermal tolerance, baseline data on the dynamics of coral holobiont performance under non-stressed conditions are needed. We monitored the seasonal dynamics of algal symbionts (family Symbiodiniaceae) hosted by corals from a chronically warmed and thermally variable reef compared to a thermally stable reef in southern Taiwan over 15 months. We assessed the genera and photochemical efficiency of Symbiodiniaceae in three coral species: Acropora nana, Pocillopora acuta, and Porites lutea. Both Durusdinium and Cladocopium were present in all coral species at both reef sites across all seasons, but general trends in their detection (based on qPCR cycle) varied between sites and among species. Photochemical efficiency (i.e., maximum quantum yield; Fv/Fm) was relatively similar between reef sites but differed consistently among species; no clear evidence of seasonal trends in Fv/Fm was found. Quantifying natural Symbiodiniaceae dynamics can help facilitate a more comprehensive interpretation of thermal tolerance response as well as plasticity potential of the coral holobiont.

Moderate chlorophyll-a environments reduce coral bleaching during thermal stress in Yap, Micronesia.

Thermal-stress events on coral reefs lead to coral bleaching, mortality, and changes in species composition. The coral reefs of Yap, in the Federated States of Micronesia, however, remained largely unaffected by major thermal-stress events until 2020, when temperatures were elevated for three months. Twenty-nine study sites were examined around Yap to determine geographical and taxonomic patterns of coral abundance, bleaching susceptibility, and environmental predictors of bleaching susceptibility. Island-wide, 21% (± 14%) of the coral cover was bleached in 2020. Although inner reefs had a greater proportion of thermally-tolerant Porites corals, the prevalence of bleaching was consistently lower on inner reefs (10%) than on outer reefs (31%) for all coral taxa. Corals on both inner and outer reefs along the southwestern coast exhibited the lowest prevalence of coral bleaching and had consistently elevated chlorophyll-a concentrations. More broadly, we revealed a negative relationship between bleaching prevalence and (moderate) chlorophyll-a concentrations that may have facilitated resistance to thermal stress by reducing irradiance and providing a heterotrophic energy source to benefit some corals exposed to autotrophic stress. Southwestern reefs also supported a high but declining fish biomass, making these bleaching-resistant and productive reefs a potential climate-change refuge and a prime target for conservation.