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In recent years, extreme thermal events have become one of the main reasons for coral reef degradation. Determining the variation of coral skeletal geochemical proxies in response to extreme thermal events would be vital evidence for reconstructing historical extreme thermal events using corals. In this study, cores from modern Porites corals YSL24 that lived in 1987-1999 and YSL2A that lived in 1971-1999 were collected from the Yongshu Reef, Nansha Islands, southern South China Sea (SCS) for high-resolution analysis of skeletal Sr/Ca, δ18O, and δ13C to determine the responses of these geochemical proxies to past extreme warming events. There is evidence that extreme warming events occurred in 1973, 1983 and 1998 in the southern SCS during the period 1971-1999. The Sr/Ca ratios in the Porites corals showed a significant negative correlation with the variation in local sea surface temperature (SST), and the values in the bands of these three years were much lower. This suggested a clear record of higher temperatures than other years in response to the warming events in these episodes. Compared with the Sr/Ca ratios, skeletal δ18O proxies in the two Porites corals also revealed a negative correlation with SST, but the δ18O values in these three bands were likely to be less sensitive to the thermal events. This is mainly because the variation in this proxy was controlled by both local SST and sea surface salinity, while the two factors had opposite effects on the seasonal variation of coral δ18O in the SCS. After the removal of the SST contributions, the ∆δ18O series of the Porites corals showed clear annual cycles and a negative correlation with seasonal precipitation, but the records in the bands of those warming years did not reveal clear anomalous shifts due to thermal stress, which might be an inappropriate proxy for identifying extreme warming episodes. In the skeletal δ13C series of Porites corals, an abnormal negative shift was detected during the summers of 1973, 1983, and 1998. This was likely caused by a substantial reduction in photosynthesis of symbiotic zooxanthellae in the coral host cells, a biological reaction of corals responding to thermal stress in the three years. In conclusion, the lower Sr/Ca ratios and negative δ13C shifts in the summer that occurred in the bands of Porites corals in this study, revealed significant coincidence with past extreme warming events. As the Porites corals had a stronger tolerance to thermal stress and much lower bleaching rates than other corals, the abnormal geochemical records in the Porites corals are potential signals indicating local past mass coral bleaching events in recent years.

The growing threat of global warming on coral reefs underscores the urgency of identifying heat‐tolerant corals and discovering their adaptation mechanisms to high temperatures. Corals growing in intertidal rock pools that vary markedly in daily temperature may have improved heat tolerance. In this study, heat stress experiments were performed on scleractinian coral Porites lutea from subtidal habitat and intertidal rock pool of Weizhou Island in the northern South China Sea. Thermotolerance differences in corals from the two habitats and their mechanisms were explored through phenotype, physiological indicators, ITS2, 16S rRNA, and RNA sequencing. At the extremely high temperature of 34°C, rock pool P. lutea had a stronger heat tolerance than those in the subtidal habitat. The strong antioxidant capacity of the coral host and its microbial partners was important in the resistance of rock pool corals to high temperatures. The host of rock pool corals at 34°C had stronger immune and apoptotic regulation, downregulated host metabolism and disease‐infection‐related pathways compared to the subtidal habitat. P. lutea, in this habitat, upregulated Cladocopium C15 (Symbiodiniaceae) photosynthetic efficiency and photoprotection, and significantly increased bacterial diversity and coral probiotics, including ABY1, Ruegeria, and Alteromonas. These findings indicate that rock pool corals can tolerate high temperatures through the integrated response of coral holobionts. These corals may be 'touchstones' for future warming. Our research provides new insights into the complex mechanisms by which corals resist global warming and the theoretical basis for coral reef ecosystem restoration and selection of stress‐resistant coral populations. [ABSTRACT FROM AUTHOR]

Outbreaks of crown-of-thorn starfish (COTS) have caused dramatic declines in reefs through predation on corals, but the post-bloom effects of COTS may still potentially threaten the environment and living organisms due to massive organic decomposition. This stimulation experiment showed that the decomposition of COTS debris triggered an extra mineralization process and resulted in acidifying, hypoxic, and eutrophic seawater. Consequently, the photosynthetic efficiency of coral symbionts decreased by 83%, and coral bleached after removing the stress within two days, then the coral skeleton dissolved at rates of 0.02–0.05 mg cm−2 day−1. Within two weeks, the photosynthesis and growth of benthic algae were suppressed by 27–86% and 1.5–16%, respectively. The mortality of turf algae and coralline algae indicated compromised primary productivity and limited coral recruitment, respectively. However, macroalgae, as coral competitors, became the only survivors, with increasing chlorophyll content. This study suggests a continuing decline of reefs during the collapse phase of COTS outbreaks and highlights the need for improving control strategies for the COTS population. [ABSTRACT FROM AUTHOR]

Sexual reproduction plays an important role in population and community structure maintenance in scleractinian corals, particularly in growth and development after larvae settlement. Despite this, only a few studies have been conducted to monitor the post-settlement growth and development of scleractinian coral larvae. Our study aimed to inform this question by collecting adult Galaxea fascicularis colonies from Weizhou Island (21°00′–21°10′ N, 109°00′–109°15′ E) and recording their early life cycle, from fertilization to recruitment, for one year ex situ. The results demonstrated that G. fascicularis could reproduce sexually when reared in a pond for 2 y. The spawning occurred in April and May, in synchrony with wild corals. Larvae showed association with zooxanthellae 9 d after spawning (4 d after settlement) and completed metamorphosis after 1 month. The mean diameter of G. fascicularis recruits was 4.74 ± 1.12 mm in the first year, which was lower than that reported in the literature for other broadcast corals. The recruits' survival rate decreased continuously every month, with the most severe decline occurring in the second month, from 80.87% to 50.82%. By the end of the first year, the final survival rate was 5.46%, which may be attributed to their competition with algae. This study improved our understanding of the early life cycle of scleractinian corals. [ABSTRACT FROM AUTHOR]

Coral reef ecosystem, which is one of the most important marine ecosystems, is suffered from serious degradation due to climate change (such as global warming and ocean acidification) and human activities (such as overfishing, chemical pollution). Organic ultraviolet absorbents (OUVs) are a kind of emerging "pseudo-persistent" organic pollutants. Previous studies indicated that OUVs have an importantly negative influence on coral health and are ubiquitous in various coral reef regions. To systematically understand the toxicity mechanism and ecological risk of OUVs to corals, this paper reviewed the toxicological effects of OUVs on corals from the perspectives of toxicological experiment design and toxicological effect endpoint. The results show that OUVs are toxic to corals at individual, tissular and molecular levels, with the toxicological endpoints of lethality, bleaching, tentacle retraction, [ABSTRACT FROM AUTHOR]

Coral skeleton δ13C is a routinely measured indicator in modern insolation change research, but the knowledge of environmental and climatic signals recorded in its seasonality during geological time is sparse. In this study, we present eight Porites coral δ13C records from the mid‐late Holocene to the present from the northern South China Sea (NSCS). Compared with the main control factors for modern δ13C changes, coral δ13C seasonality in the NSCS since the mid‐Holocene shows a long‐term decreasing trend, which is consistent with the change trend as orbital‐induced (precession) insolation seasonal amplitude. By excluding other influencing factors, we speculate that the stronger coral δ13C seasonality (18.8%) over the mid‐Holocene than modern period is attributable to the metabolic effect, which predicts the stronger coral δ13C seasonality under stronger insolation seasonality. Our study has implications for coral δ13C seasonality as a potential indicator to record past insolation information under different climatic backgrounds. Plain Language Summary: Numerous studies have documented the seasonal features of fast‐growing modern coral skeleton δ13C in response to insolation variation. The environmental and climatic signals recorded by coral δ13C seasonality in the geological period remain unclear owing to the relatively short length of observation records. Here we present monthly resolved δ13C records in seven fossil Porites corals (5.6–3.6 ka BP, before 1950 CE) and one modern Porites coral (1987–2001 CE) from the northern South China Sea (NSCS). Compared with the widely accepted prevailing control factors for modern coral δ13C, our results indicated that the stronger coral δ13C seasonality (18.8%) over the mid‐Holocene compared to the modern period is consistent with the orbital‐induced (precession) insolation changes, which indicate a declining seasonality from the mid‐Holocene to present. By excluding other influencing factors, we infer that a tendency toward higher δ13C seasonality during the mid‐Holocene was primarily driven by the metabolic fractionation caused by the enhanced insolation seasonality. This study highlighted coral δ13C seasonality as a potential indicator for recording past insolation information. Key Points: The seasonal variation of modern coral skeleton δ13C in the northern South China Sea (NSCS) is primarily controlled by solar insolationCoral δ13C seasonality in the NSCS since the mid‐Holocene shows a long‐term decreasing trendThe decrease of orbital‐induced (precession) insolation seasonal amplitude led to the declining trend of δ13C seasonality [ABSTRACT FROM AUTHOR]

To better understand the long-term survival of modern coral reefs under global warming, the history of past coral bleaching events prior to modern marine ecological surveys must be investigated. Here, 11 dead Porites and 2 Porites with growth hiatuses growing in the 20th century were collected from the Nansha Islands, southern South China Sea (SCS) for monthly Sr/Ca, δ18O and δ13C analyses to explore the climatic conditions and metabolic information corresponding to these growth structures. Among the 13 episodes, the results showed that 8 mortalities occurred in the summers, coral MJ8 died in spring, and the other 2 mortalities and 2 growth hiatuses occurred in the cooling season. In addition to the dead surface of the coral mortalities, stress bands with significantly reduced growth rates could also be observed in the corals. Negative shifts in δ13C occurred prior to 5 coral mortalities and most stress bands were caused in summer, suggesting a sharp reduction in photosynthesis by decreased symbiotic zooxanthellae or photosynthetic pigments. Overall, the mortalities can be concluded to have occurred in summer and stress bands with abrupt δ13C decreases in the last century were mainly caused by thermal coral bleaching in the southern SCS. This in turn suggests that the repeated coral bleaching and mortalities occurred much earlier before the 1980s, which is a longer-term response to modern global warming during the last century. [ABSTRACT FROM AUTHOR]

Ecological surveys observe coral "winners" and "losers" in global coral bleaching events. However, the key contributors to holobiont tolerance and interactions between symbionts remain unclear. Herein, we compared bleaching and unbleaching Acropora pruinosa corals from Weizhou Island, during an extreme high-temperature event in the northern South China Sea in 2020. We found the dominant Symbiodiniaceae subclade in the bleaching and unbleaching corals to be C1; however, the density of Symbiodiniaceae in the latter was significantly higher than that in the former. Additionally, the symbiotic bacteria α diversity in the unbleaching coral was significantly higher than that in the bleaching coral, with a reorganized bacterial community structure. Core microbiome analyses revealed 55 bacterial core operational taxonomic units (OTUs), of which 10 were significantly differentially enriched between the two coral groups. The significantly enriched bacterial core OTUs in the unbleaching coral were primarily nitrogen cycling related, while those enriched in the bleaching coral were associated with antimicrobial activity. RNA-Seq analyses revealed that significantly upregulated genes in the bleaching coral were primarily associated with diseases and autophagy, while those in the unbleaching coral were associated with immune defense and maintenance of the symbiotic relationship between corals and symbionts. We propose that the differences in tolerance of A. pruinosa result from the cooperation between coral host, Symbiodiniaceae, and symbiotic bacteria. In extreme high-temperature events, unbleaching corals may maintain stable symbiotic relationships by increasing the diversity of symbiotic bacteria, regulating the structure of the symbiotic bacteria community, improving the interaction between coral host and symbiont and enhancing host immunity, thus avoiding coral bleaching. This study illuminates the relationship between the coral symbiont and tolerance differences of coral holobionts, providing new insights for further exploration into the adaptability of scleractinian corals in the context of global warming. [ABSTRACT FROM AUTHOR]

The seasonal and interannual variabilities of sea surface wind (SSW) in the South China Sea (SCS), especially in coral reef regions such as Nansha Islands, Xisha Islands, Zhongsha Islands and Dongsha Islands were investigated in detail using the Blended Sea Winds dataset (1988–2017). Annual and monthly variations of SSW and sea surface temperature (SST) in the four zones were investigated. Empirical Orthogonal Function (EOF) analysis of wind field was performed to aid in better understanding the different spatial patterns. The results indicate that, as observed in the spatial distribution of the first mode of monthly mean wind speed anomaly, the magnitudes in the four island zones are all negative and are similar to each other, showing that the variations of SSW in the four island zones are consistent. In the second mode, the magnitudes in Nansha Islands are opposite to those in the other three zones. The spatial distribution of the third mode reflects regional differences. The maximum annual SSW appears in Dongsha Islands, and the minimum appears in Nansha Islands. The interannual variations of SSW in all island zones are basically concurrent. The island zones with high SSW mostly have low SST, and vice versa. There may be an inverse relationship between SSW and SST in coral reef regions in the SCS. The multi-year monthly variations of SSW in the island zones present a 'W'-shaped structural variation. Each island undergoes two months of minimum SSW every year, one during March–May (MAM) and the other during September–November (SON). Both months are in monsoon transition periods. During the months with low SSW, high SST appears. The SST peaks almost correspond to the SSW troughs. This further indicates that SSW and SST may have opposite changes in coral reef regions. Coral bleaching events often correspond to years of high SST and low SSW. [ABSTRACT FROM AUTHOR]

Aim: In this study, we investigated whether environmental factors can effectively control the spatial distribution of various benthic algae and examined the critical ecological impacts of algae on corals across the South China Sea (SCS). Relationships between benthic algae and environmental factors were assessed, and potential ecological impacts of algae on coral communities were evaluated across spatial scales. Location: A total of 104 sites at 12 coral reefs in four coral reef regions (CRRs) of the SCS (latitude range of 9–22° N). Taxa: Turf algae, macroalgae (including fleshy macroalgae and Halimeda), crustose coralline algae (CCA), coral and juvenile coral. Methods: Using diver‐based surveys (2015–2018), we investigated four CRRs with different reef distance to mainland (RDM) across the SCS. We obtained field data on benthic algal composition and cover, coral cover and diversity, and juvenile coral density. We also measured and collected the environmental factors (including seawater environmental parameters and reef fish diversity). Results: The cover of turf algae and CCA gradually decreased and increased, respectively, with an increase in RDM, which had a strong relationship with the gradually changing environmental factors. Random forest models suggested that nutrients, reef fish diversity, seawater transparency and temperature were the most important factors for predicting turf algae and CCA cover. Linear regression analyses showed a significant relationship between the turf algae, macroalgae, and CCA covers, and juvenile coral density. Main conclusions: Our results showed that gradually changing environmental factors were correlated with the spatial distributions of turf algae and CCA. However, the natural biophysical relationships between macroalgae and environmental factors may be disrupted by turf algal overgrowth and nutrient subsidies. Increases in all benthic algal groups significantly impacted coral recruitment, highlighting the critical role of benthic algae in determining the recovery trajectory of the Indo‐Pacific reefs that are threatened by human activities. [ABSTRACT FROM AUTHOR]

Although many achievements have been made in studies on the relationship between Symbiodiniaceae and coral bleaching, there is little information on the responses of Symbiodiniaceae to coral thermal bleaching in the South China Sea (SCS). In this study, Symbiodiniaceae communities and the effective quantum yield (Ф PSII) of healthy and bleached massive corals from the SCS were compared. The results indicated that the Symbiodiniaceae communities and Ф PSII values showed coral bleaching-dependent variations. Specifically, the relative abundances of the rare putatively thermally tolerant Durusdinium sp. (D1) and the Symbiochlorum hainanensis increased significantly in the bleached corals. In addition, bleached Porites lutea was mainly associated with the putatively thermally tolerant Cladocopium sp. (C15 and C91) and exhibited the highest Ф PSII value compared with other bleached corals. These results highlight that the rare putatively thermally tolerant algae and coral species-specific algae may be important for understanding thermal bleaching of corals. • A snapshot of Symbiodiniaceae communities and photosynthesis following thermal bleaching of massive corals in the SCS. • Symbiodiniaceae communities and Ф PSII values showed coral bleaching-dependent variations. • The abundance of Durusdinium sp. (D1) and Symbiochlorum hainanensis were significantly increased in bleached corals. • Porites lutea showed smallest decrease in Ф PSII and hosted coral specific Cladocopium sp. (C15 and C91). [ABSTRACT FROM AUTHOR]

Abstract: Coral bleaching is becoming a serious issue for coral reefs under the stress of global warming. However, whether it has occurred in the past in times of thermal stress remains unclear. Moreover, an understanding of historic coral bleaching events would greatly improve our insight into the adaptive capabilities of corals under such stresses. It is known that Porites corals, a massive coral, have relatively high levels of symbiotic zooxanthellae and a strong thermal tolerance when compared with most other corals (and particularly branched corals). Thus, growth hiatuses and/or mortality surfaces of fossil Porites may be used to indicate past ecological or environmental stress events, such as severe bleaching. In this study, monthly geochemical and isotopic environmental proxies of four fossil Porites corals with well‐preserved growth hiatuses and mortality surfaces (aged 3,800–4,200 years before 2013 A.D.), collected from Wenchang fringing reef, Hainan Island, Northern South China Sea were analyzed. Specifically, the Sr/Ca, δ18O, and δ13C were measured with a monthly resolution for each sample. These environmental proxies were used to reconstruct the sea surface temperature (SST), sea surface salinity (SSS), and physiological activity of the coral at the time when the growth hiatuses and/or mortality surfaces occurred. These data were subsequently used to determine the causes of coral mortality and growth discontinuity in each case. The results show that growth hiatuses and mortalities mainly occurred in summer, with high SST (31–34°C) and SSS (32.8–38.4). In addition, abrupt negative shifts of 2–3‰ in δ13C were observed in almost all of the surfaces of growth hiatus and mortality, indicating a dramatically reduced level of photosynthetic activity in symbiotic zooxanthellae. Because of the above reasons, we conclude that the frequently observed mortality and growth discontinuity of Porites corals from the mid‐Holocene is evidence for thermal bleaching events in the past. That is, coral bleaching has occurred 3,800–4,200 years ago and is not a new phenomenon. [ABSTRACT FROM AUTHOR]

Coral bleaching heat stress products provide real-time and rapid coral bleaching alerts for coral reefs globally. However, geographical variations in the alert accuracy of multi-source coral bleaching heat stress products exist. Taking the coral reefs in the South China Sea (SCS) as the study area, we evaluated and improved the coral bleaching alert capabilities of two coral bleaching heat stress products: Coral Reef Watch (CRW) and Coral Reef Temperature Anomaly Database (CoRTAD). Using in situ coral bleaching survey data and evaluation indicators, the optimized thresholds of degree heating weeks (DHWs) for coral bleaching alerts were determined. The results in the SCS indicated that, first, CRW was better than CoRTAD for coral bleaching event alerts. However, both products underestimated coral bleaching events using the common DHW thresholds of 4°C-weeks and 8°C-weeks. Second, the DHW optimized threshold for CRW was 3.32°C-weeks for coral bleaching event alerts and 4.52°C-weeks for severe coral bleaching event alerts. For CoRTAD products, the DHW optimized threshold was 2.36°C-weeks for coral bleaching event alerts and 4.14°C-weeks for severe coral bleaching event alerts. This study proposed a method to evaluate and optimize the alert capability of multi-source coral bleaching heat stress products, which can provide more accurate basic data for coral reef ecosystem health assessment and contribute to global coral reef ecosystem protection and restoration. • A method to evaluate and optimize the alert capability of coral bleaching heat stress products in the SCS. • Optimized DHW thresholds for coral bleaching alerts were 3.32°C-weeks for CRW and 2.36°C-weeks for CoRTAD. • Optimized DHW thresholds for coral severe bleaching alerts were 4.52°C-weeks for CRW and 4.14°C-weeks for CoRTAD. • CRW was better than CoRTAD for coral bleaching event alerts in the SCS. [ABSTRACT FROM AUTHOR]

Global climate change has resulted in large-scale coral reef decline worldwide, for which the ocean warming has paid more attention. Coral is a typical mutually beneficial symbiotic organism with diverse symbiotic microorganisms, which maintain the stability of physiological functions. This study compared the responses of symbiotic microorganisms and host metabolism in a common coral species, Pavona minuta, under indoor simulated thermal and cold temperatures. The results showed that abnormal temperature stresses had unfavorable impact on the phenotypes of corals, resulting in bleaching and color change. The compositions of symbiotic bacteria and dinoflagellate communities only presented tiny changes under temperature stresses. However, some rare symbiotic members have been showed to be significantly influenced by water temperatures. Finally, by using ultra-performance liquid chromatography tandem mass spectrometry (UPLC–MS) method, we found that different temperature stresses had very different impacts on the metabolism of coral holobiont. The thermal and cold stresses induced the decrease of anti-oxidation metabolites, several monogalactosyldiacylglycerols (MGDGs), and the increase of lipotoxic metabolite, 10-oxo-nonadecanoic acid, in the coral holobiont, respectively. Our study indicated the response patterns of symbiotic microorganisms and host metabolism in coral to the thermal and cold stresses, providing theoretical data for the adaptation and evolution of coral to a different climate in the future. [ABSTRACT FROM AUTHOR]

Ciguatera poisoning is mainly caused by the consumption of reef fish that have accumulated ciguatoxins (CTXs) produced by the benthic dinoflagellates Gambierdiscus and Fukuyoa. China has a long history of problems with ciguatera, but research on ciguatera causative organisms is very limited, especially in the Beibu Gulf, where coral reefs have been degraded significantly and CTXs in reef fish have exceeded food safety guidelines. Here, five strains of Gambierdiscus spp. were collected from Weizhou Island, a ciguatera hotspot in the Beibu Gulf, and identified by light and scanning electron microscopy and phylogenetic analyses based on large and small subunit rDNA sequences. Strains showed typical morphological characteristics of Gambierdiscus caribaeus, exhibiting a smooth thecal surface, rectangular-shaped 2′, almost symmetric 4″, and a large and broad posterior intercalary plate. They clustered in the phylogenetic tree with G. caribaeus from other locations. Therefore, these five strains belonged to G. caribaeus, a globally distributed Gambierdiscus species. Toxicity was determined through the mouse neuroblastoma assay and ranged from 0 to 5.40 fg CTX3C eq cell−1. The low level of toxicity of G. caribaeus in Weizhou Island, with CTX-contaminated fish above the regulatory level in the previous study, suggests that the long-term presence of low toxicity G. caribaeus might lead to the bioaccumulation of CTXs in fish, which can reach dangerous CTX levels. Alternatively, other highly-toxic, non-sampled strains could be present in these waters. This is the first report on toxic Gambierdiscus from the Beibu Gulf and Chinese waters and will provide a basis for further research determining effective strategies for ciguatera management in the area. [ABSTRACT FROM AUTHOR]

Coral reef bleaching is usually characterized by expulsion of symbiotic zooxanthellae, loss of zooxanthellae pigmentation, or both. We collected 128 samples comprising 39 species of 21 genera of reef-building corals from Luhuitou and Xiaodonghai in Sanya of Hainan Island and Daya Bay of Guangdong Province, respectively, and analyzed the symbiotic zooxanthellae population density. The results show that: (1) the symbiotic zooxanthella density varies from 0.67×106 to 8.48×106 cell/cm2, displaying significant interspecies variability, with branch corals usually having relatively less zooxanthellae (ranging from 0.67×106 to 2.47×106 cell/cm2) than massive species (from 1.0×106 to 8.48×106 cell/cm2); (2) corals inhabiting within 4 m water depth have higher levels of symbiotic zooxanthellae than those living at the bottom (-.7 m depth) of the reef area; (3) there is no discernable difference in the zooxanthellae density between corals from relatively high latitude Daya Bay (-22°N) and those from relatively low latitude Sanya (-18°N) at comparable sea surface temperatures (SST); (4) in partially-bleached corals, the density of zooxanthellae shows the following order: healthy-looking part> semi-bleached part > bleached part. Based on the above results, we suggest that (1) the zooxanthellae density difference between branching and massive coral species is the main cause that branching corals are more vulnerable to bleaching than massive corals. For example, symbiotic zooxanthellae levels are low in branching Acropora and Pocillopora corals and thus these corals are more susceptible to bleaching and mortality; (2) symbiotic zooxanthellae density can also be affected by environmental conditions, such as sediment loads, diving-related turbidity, and aquaculture-related nitrate and phosphate input, and their increase may reduce symbiotic zooxanthellae density in corals. [ABSTRACT FROM AUTHOR]

Global warming and sea-level rise (SLR) induced by rising atmospheric CO2 concentrations can cause coral bleaching, death, and submergence of the world's coral reefs. Adopting the GIS and RS methods, we modeled how these two stressors combine to influence the future growth of the atolls and table reefs of three archipelagoes in the South China Sea (SCS), based on geomorphic and ecological zones. A large-scale survey of the coral communities in Xisha Islands in 2014, Dongsha Islands in 2014–2016 and Nansha Islands in 2007 provided zone-specific process datasets on the range of reef accretion rates. Sea surface temperature and extreme (minimum and maximum) SLR data above 1985–2005 levels by 2100 in the SCS were derived from the Intergovernmental Panel on Climate Change (IPCC) Fifth Assessment Report (AR5) models forced with the Representative Concentration Pathways (RCPs). Our model projected that: (1) the Xisha Islands and Dongsha Islands may have a better growth status, because the reef flat biotic sparse zone may be recolonized with hard coral and become a biotic dense zone; (2) the southern Nansha Islands reefs have a risk of stopping growing due to their earlier annual bleaching years. The increasing of water depths of these reefs is stronger in the RCP with more emissions. Our approach offers insights into the best-case and worst-case impacts of two global environmental pressures on potential future reef growth under a changing climate. [ABSTRACT FROM AUTHOR]

Scattered evidence indicates the occurrence of coral thermal bleaching events during the mid-Holocene. However, information on such occurrences is still very rare. Ten Porites coral cores were drilled from the Wenchang fringing reef of Hainan Island, northern South China Sea. These cores contain mortality surfaces and/or growth discontinuities dating from the mid-late Holocene between 3.4 and 5.7 thousand years ago (ka, before 1950 AD). Monthly Sr/Ca, δ18O and δ13C were analyzed to reconstruct sea-surface temperature (SST), sea-surface salinity (SSS), and photosynthesis intensity during the events. Results show that the mortality surfaces and growth discontinuities of five corals are associated with high SST of 30.3 °C–33.6 °C and high SSS of 34.3–38.3. The abrupt concurrent negative shifts of 1.8‰ or more in δ13C indicate a decrease in the photosynthesis intensity, most likely attributable to the loss of zooxanthellae. These lines of evidence imply that the mortality surfaces and growth discontinuities were the consequence of coral bleaching under abnormally high temperature. The observed recovery directly after bleaching events indicates that corals in the South China Sea are capable of rehabilitation after experiencing severe environmental stresses. • Ten Porites coral cores dated at 3.4–5.7 ka BP contain mortality surfaces/hiatuses. • Sr/Ca and δ18O suggest that mortality occurred in high SST and SSS environments. • Negative shifts of δ13C indicate a decrease in photosynthesis or zooxanthellae. • The geochemical proxies provide evidence of past coral bleaching and recovery. [ABSTRACT FROM AUTHOR]

Relatively high-latitude waters are supposed as a refuge for corals under ocean warming. A systematic assessment of the Weizhou Island reef in the northern South China Sea, a relatively high-latitude region, shows that the ecosystem restoration index decreased from 0.96 to 0.62 during the period between 1990 and 2015. Although the biotic community, supporting services, and regulating services remained at good or very good states, the provisioning services, cultural services, and especially habitat structure deteriorated to very poor or moderate states. Gray relational analysis showed that these ecological declines exhibited a strong relationship with human pressures from tourism activities and the petrochemical industry. The recoveries of the biotic community and supporting services that benefited from wintertime warming appeared to be partly offset by intensive human pressures. The long-term effects on ecosystem structure and functions suggest that anthropogenic disturbances have impaired the possibility of this area serving as a potential thermal refuge for reef-building corals in the South China Sea. This study thus provides an integrated approach for assessing the adaptive responses of coral reef ecosystems to climate change and local human activities. [ABSTRACT FROM AUTHOR]

Recent studies have indicated that coral mucus plays an important role in the bioaccumulation of a few organic pollutants by corals, but no relevant studies have been conducted on organochlorine pesticides (OCPs). Previous studies have also indicated that OCPs widely occur in a few coral reef ecosystems and have a negative effect on coral health. Therefore, this study focused on the occurrence and bioaccumulation of a few OCPs, such as dichlorodiphenyltrichloroethanes (DDTs), hexachlorobenzene (HCB) and p , p ′-methoxychlor (MXC), in the coral tissues and mucus as well as in plankton and seawater from a coastal reef ecosystem (Weizhou Island) in the South China Sea. The results indicated that DDTs were the predominant OCPs in seawater and marine biota. Higher concentrations of OCPs in plankton may contribute to the enrichment of OCPs by corals. The significantly higher total OCP concentration (∑ 8 OCPs) found in coral mucus than in coral tissues suggested that coral mucus played an essential role in resisting enrichment of OCPs by coral tissues. This study explored the different functions of coral tissues and mucus in OCP enrichment and biodegradation for the first time, highlighting the need for OCP toxicity experiments from both tissue and mucus perspectives. [Display omitted] • OCPs were firstly studied in coral mucus in the South China Sea. • Plankton may have an influence on coral enrichment of OCPs. • Obvious higher ∑ 8 OCPs occurred in coral mucus than in coral tissues. • Coral mucus showed lower metabolic capability of DDTs than coral tissues. • Coral mucus may resist the enrichment and metabolism of OCPs by coral tissues. [ABSTRACT FROM AUTHOR]

Coral bleaching events are becoming increasingly common worldwide, causing widespread coral mortality. However, not all colonies within the same coral taxa show sensitivity to bleaching events, and the current understanding of the metabolic mechanisms underlying thermal bleaching in corals remains limited. We used untargeted metabolomics to analyze the biochemical processes involved in the survival of two bleaching phenotypes of the common corals Pavona decussata and Acropora pruinosa , during a severe bleaching event in the northern South China Sea in 2020. During thermal bleaching, P. decussata and A. pruinosa significantly accumulated energy products such as succinate and EPA, antioxidants and inflammatory markers, and reduced energy storage substances like glutamate and thymidine. KEGG analysis revealed enrichment of energy production pathways such as ABC transporters, nucleotide metabolism and lipid metabolism, suggesting the occurrence of oxidative stress and energy metabolism disorders in bleached corals. Notably, heat stress exerted distinct effects on metabolic pathways in the two coral species, e.g., P. decussata activating carbohydrate metabolism pathways like glycolysis and the TCA cycle, along with amino acid metabolism pathways, whereas A. pruinosa significantly altered the content of multiple small peptides affected amino acid metabolism. Furthermore, the osmoregulatory potential of corals correlates with their ability to survive in heat-stress environments in the wild. This study provides valuable insights into the metabolic mechanisms linked to thermal tolerance in reef-building corals, contributes to the understanding of corals' adaptive potential to heat stress induced by global warming and lays the foundation for developing targeted conservation strategies in the future. • Metabolic profiles differed between unbleached and bleached corals. • Bleached corals consumed energy reserves like nucleotides and lipids. • Coral bleaching led to oxidative damage and immune response activation. • Osmotic substances were crucial in coral response to heat stress. • New insights into metabolic response of reef-building corals to heat stress. [ABSTRACT FROM AUTHOR]

Investigating the potential and mechanisms of thermal adaptability in scleractinian corals is critical for aiding corals in dealing with global warming and improving the efficiency of coral reef restoration efforts. Although thermal acclimation facilitates the resistance of branching corals to thermal stress, little is known about the physiological processes of massive and platy corals in the South China Sea or their ability to adapt to heat stress. In this study, we conducted indoor short-term thermal acclimation simulation experiments on three types of corals: massive Porites lutea , platy Pavona decussata , and branching Pocillopora damicornis. Subsequently, we studied the responses and adaptive mechanisms of the three corals to heat stress and explored the effects of short-term thermal acclimation using physiological and biochemical markers. The results showed that the three corals exhibited comparable phenotypic and physiological responses to heat stress, including tentacle retraction and decreases in zooxanthellae density, maximum quantum efficiency of photosystem II, and glutathione content, along with increases in antioxidant activity (catalase and superoxide dismutase), ammonium assimilation (glutamine synthetase), and apoptosis (lipid peroxide and caspase-3). Additionally, heat tolerance differed among the different species of coral. Furthermore, the physiological markers performed better in all three coral types after acclimation, and the effects of short-term acclimation decreased from P. lutea to P. damicornis to P. decussata. We propose that short-term thermal acclimation enhances the heat tolerance of corals by affecting their metabolism and antioxidant capacity. Leveraging short-term thermal acclimation in coral reef restoration efforts could help corals adapt to the threats posed by global warming and enhance restoration efficiency. • Coral reefs are in rapid decline in the context of global pressures. • Different types of scleractinian corals display similar responses to heat stress. • Short-term thermal acclimation enhances the coral antioxidant capacity. • Porites lutea and Pocillopora damicornis have better thermal acclimatization. [ABSTRACT FROM AUTHOR]

The El Niño-Southern Oscillation (ENSO) is the most influential climatic phenomenon on an interannual timescale. Studying the relationship between sea surface temperature (SST) and ENSO events during typical geological periods is helpful for understanding the trends of ENSO activities in the future, especially under the background of global warming. Here we quantitatively reconstructed a 115-year history of SST and ENSO activity around 282 ka during Marine Isotope Stage 8 (MIS8) by using the growth rate of Porites corals from Xisha Islands in the South China Sea. The results show that the average coral growth rate (6.0 ± 1.0 mm/yr (1σ)) around 282 ka was only 56.1% of the modern average value (10.7 ± 0.9 mm/yr; 1937–2014 CE), and the average SST (26.0 ± 0.3 °C (1σ)) was 0.7 ± 0.4 °C and 1.1 ± 0.4 °C lower than that in the Little Ice Age (LIA; 1520–1676 CE; 26.7 ± 0.2 °C) and the present (1937–2014; 27.1 ± 0.2 °C), respectively. The reconstructed results show that there was significant ENSO activity with moderate intensity or above at 282 ka, while the frequency of ENSO activity at that time was lower than that of the LIA and the present. This seems to indicate that the frequency of moderate intensity ENSO activity increases with rising SST. Given the current rapid warming, the future trend of ENSO activities deserves close attention. • The growth rate of Porites coral is a reliable paleoenvironmental proxy. • The coral growth rate at 282 ka was only equivalent to 56.1% of the present. • Coral–SST at 282 ka was 0.7 °C and 1.1 °C lower than the LIA and present, respectively. • The ENSO frequency at 282 ka was lower than the present. [ABSTRACT FROM AUTHOR]

Holocene coral reef development is valuable to understanding the behavior and impacts of paleoclimate and sea level changes. This study focuses on the Holocene carbonate sequences of a drill core extracted from Well CK2 in the northern South China Sea (SCS). This drill core consists of an 873.55 m thick carbonate layer overlying 55.20 m of volcanic basement. High precision U-series dating of 22 coral samples suggests that the Holocene coral reef, which has a thickness of 16.7 m, initiated at ~7800 yr BP (years before 1950 CE) and stopped vertical accretion at 3900 yr BP. The mean vertical accretion rate in the Holocene in Well CK2 was 3.48 m kyr−1, varying from 6.44 m kyr−1 during 7800–6000 yr BP to 0.87 m kyr−1 during 6000–3900 yr BP. After 3900 yr BP, the vertical development of the reef ceased, indicating that the reef development likely involved lateral accretion. The Holocene section mostly consists of unconsolidated coarse sediments, mainly including corals, coralline algae, large benthic foraminifera, and mollusks. Based on the relationship between coral reef development and sea level, together with the age profile, we determined that the relative sea level near Chenhang Island rose rapidly between 7800 and 6000 yr BP, slowed at ~6000 yr BP, and reached a position of about 2 m above the present-day mean sea level by 3900 yr BP. After 3900 yr BP, the sea level was stable or fell, resulting in a cessation in the reef's upward development. • 22 230Th ages of a SCS reef core suggest the Holocene reef initiated at 7800 yr BP. • Vertical development decreased from 6.44 to 0.87 m kyr−1 at 3900 yr BP. • Sea level at 7800 yr BP was ~13.8 m below present and rose rapidly. • Sea level has stabilized/fallen since 3900 yr BP, as indicated by reef development. [ABSTRACT FROM AUTHOR]

Coral probiotics can improve the tolerance of corals to heat stress, thus mitigating the process of coral thermal bleaching. Sensitive and specific detection of coral probiotics at low abundances is highly desirable but remains challenging, especially for rapid and on-site detection of coral probiotics. Since the electrochemical biosensor has been recently used in the field of environmental DNA (eDNA) detection, herein, an efficient electrochemical biosensor was developed based on CoS 2 /CoSe 2 -NC HNCs electrode material with a specific DNA probe for the C. marina detection. After optimization, the lower limit of detection (LOD) values of such biosensors for the target DNA and genomic DNA were 1.58 fM and 6.5 pM, respectively. On this basis, a portable device was constructed for the practical detection of C. marina eDNA, and its reliability and accuracy were verified by comparison with the ddPCR method (P > 0.05). For each analysis, the average cost was only ∼ $1.08 and could be completed within 100 min with reliable sensitivity and specificity. Overall, the biosensor could reflect the protective effect of probiotics on coral heat stress, and the proposed technique will put new insights into the rapid and on-site detection of coral probiotics to assist corals against global warming. [ABSTRACT FROM AUTHOR]

[Display omitted] • Blue light increases the thermal bleaching tolerance of coral holobiont. • Blue light decreases photosynthesis, respiration and ROS release of Symbiodiniaceae under heat stress. • Blue light reduces respiration and lipid peroxidation of host under heat stress. • The up-regulation of the DAO gene indicates a coral bleaching model that dependents on the host. Given the high degree of heterogeneity of coral bleaching, there is potential for abiotic approaches to improve the thermal tolerance of coral. However, there is limited knowledge about such factors and causes that regulating acclimation of coral to warming. Here, a study was conducted on the reef building coral Acropora tenuis to investigate the impact of light qualities on the physiological, cellular, and molecular biochemical responses of the coral holobiont to heat stress. The results showed that after subjecting the samples to heat stress at 32 ℃ for 9 days, over 90% corals exposed to compound light experienced bleaching on the fifth day, while corals exposed to blue light showed no significant signs of bleaching even on the ninth day. This indicates that blue light may enhanced the coral's thermal bleaching tolerance. Further analysis revealed blue light primarily increased coral's resistance to thermal bleaching by remodeling the interactions between Symbiodiniaceae (i.e. decreased photosynthesis, respiration and ROS release) and coral (i.e. decreased respiration and lipid peroxidation). The study also found that the host's of reactive oxygen species (ROS) played a role in causing coral bleaching besides the ROS released by Symbiodiniaceae. The up-regulation of the DAO gene (encoding D-amino-acid oxidase, up-regulated nearly 1000 times under warming conditions) in the host highlights the mechanism of coral bleaching, whereby the catalysis of amino acids by the D-amino-acid oxidase produces ROS. In summary, the present study reveals novelty insights into the molecular processes related to thermal bleaching of coral, identifies blue light as a positive environmental factor that enhances coral's tolerance to global warming. [ABSTRACT FROM AUTHOR]

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. • The most intense severe heat stress anomaly years were 2010 > 2014>2020 > 2015. • The anomaly duration of offshore islands was Xisha > Dongsha > Zhongsha > Nansha Islands. • High-latitude reefs were predicted not to be future thermal refuges for SCS reefs. • The optimized DHW threshold more accurately predicted spatiotemporal heat stress. [ABSTRACT FROM AUTHOR]

Widespread coral bleaching and mortality has occurred intensively and frequently under global warming conditions since the phenomenon was first documented in the early 1980s. However, abrupt and extensive mortality episodes, such as which occurred in the South China Sea (SCS) in mid-1991, cannot be explained solely on the basis of high summer sea surface temperatures. Here we examine this event in the light of time series rare earth elements, trace element Al/Ca ratios, and micro-domain images from coral samples collected in the SCS region. Results demonstrate that modern coral reefs were strongly imposed by a giant volcanic eruption, the 1991 Mount Pinatubo volcanic eruption. Our findings highlight that contemporary continental near-shore reefs are more susceptible than their open-ocean counterparts. [ABSTRACT FROM AUTHOR]