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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]

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]

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]

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]

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]

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]

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]

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]

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]

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]

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]

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]