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The trade winds cause strong upwelling in the eastern equatorial Pacific, and create the eastern Pacific Cold Tongue (EPCT) that has far‐reaching impacts on global climate. However, large discrepancies persist in quantifying 20th‐century EPCT sea surface temperature (SST) changes across different instrumental data sets. Here we synthesize four coral Sr/Ca‐SST records from the tropical central‐eastern Pacific to develop a Cold Tongue Index (CTI) reconstruction for 1887–1997. The coral CTI record shows a rapid 20th century warming of the EPCT, suggesting an underestimation of warming trends in instrumental CTI records. The decadal to multidecadal changes in reconstructed EPCT SST show an association with the Walker Circulation. Our reconstruction indicates that recent EPCT cooling during the global warming hiatus is not unusual in the context of the 20th century. Our results provide new evidence for 20th century EPCT SST changes and an observational constraint for predicting future tropical climate changes. Plain Language Summary: The tropical Pacific exhibits an asymmetric pattern of sea surface temperatures (SST) along the equator, with a warm pool in the west and a cold tongue in the east. The Eastern Pacific Cold Tongue (EPCT) plays a crucial role in influencing global climate. To date, however, considerable disagreement persists about changes and causes of the EPCT SST over the 20th century due to the sparse instrumental observations before ∼1960. Here we synthesize published coral Sr/Ca‐SST records from the tropical central‐eastern Pacific to develop a coral Cold Tongue Index (CTIcoral) for the period 1887–1997. The CTIcoral record correlates significantly with instrumental data since 1960, and thus allows reliable assessment of 20th century EPCT SST changes. Our record exhibits a rapid 20th century warming trend of the EPCT, indicating an underestimation of warming trends in instrumental CTI records. The decadal to multidecadal changes in CTIcoral also show an association with the Walker Circulation. Comparing the magnitude of the 1992–2011 trend in instrumental CTI with our reconstruction reveals that recent EPCT cooling during the global warming hiatus is not anomalous in the context of 20th century. Our findings have implications for predicting future EPCT SST changes by offering a constraint for model simulations. Key Points: Synthesizing coral Sr/Ca‐SST records yields robust estimates of 20th century warming trend in the eastern Pacific Cold Tongue (EPCT)A coral‐based reconstruction indicates an underestimation of 20th century EPCT warming trends in instrumental SST recordsRecent EPCT cooling during the global warming hiatus is not anomalous in the context of the 20th century [ABSTRACT FROM AUTHOR]

Environmental conditions at different water depths on tropical atolls are substantially different, but few studies have investigated the depth distribution and adaptation in the microbiomes of juvenile and adult corals. We collected samples of juvenile and adult Pocillopora verrucosa from different depths on an atoll in the central South China Sea and analyzed their communities of symbiotic Symbiodiniaceae and associated bacteria. Results showed that the Symbiodiniaceae communities were significantly different at different depths in both juvenile and adult P. verrucosa. Symbiodiniaceae community changed with increasing depth and was dominated by Durusdinium and Cladocopium at 1-m and 20-m depth, respectively. Furthermore, the composition and diversity of Symbiodiniaceae in juvenile and adult corals were different at the same depths. The compositions of associated bacterial communities at different depths also differed significantly between juvenile and adult P. verrucosa. The relative abundance of Endozoicomonas in juvenile and adult P. verrucosa increased with increasing depth, while the Thermus decreased. Our study suggests that juvenile and adult P. verrucosa corals in shallow-water regions associate with high abundances of thermotolerant Symbiodiniaceae and bacteria in the shallow reef flats with high temperature and intense solar radiation, but associate with Symbiodiniaceae and associated bacteria with high photosynthetic efficiency at greater depths. [ABSTRACT FROM AUTHOR]

Tropical cyclones (TCs) lead to huge economic losses and casualties in coastal regions, but multi‐decadal variation of TC remains poorly understood due to the limited length of observational records and low‐resolution reconstructions. Porites corals provide continuous and monthly resolved records that allow for the determination of weather‐timescale (weekly to monthly) extreme events, including TC, to overcome the above limitations. Here we synthesize five coral Δδ18O records from the South China Sea (SCS) and develop a coral TC frequency index, in good agreement with post‐1980 instrumental records and sediment‐based TC reconstructions over the past century. Our results indicate that TC frequency in the SCS exhibits pronounced decadal variations, with active TCs corresponding to the negative phase of the Pacific Decadal Oscillation. Our findings here provide novel evidence for past TC changes in the SCS and a constraint for predicting future TC changes. Plain Language Summary: The impact of tropical cyclones (TCs) on coastal regions results in significant economic losses and casualties. Due to the relatively short observations or low‐resolution records, assessing long‐term changes of TCs has suffered from large uncertainties. Porites corals are sensitive to environmental changes and can provide century‐long, monthly resolved records, which could serve as a potential archive for reconstructing past TCs. Here we present five coral Δδ18O records from the South China Sea (SCS) and develop two TC frequency indexes. Both the TCs generated in the SCS and those entering the SCS from the Western North Pacific are reflected in our coral‐based TCs reconstruction. In addition, our reconstruction suggests that the TCs in the SCS were increased in 1878–1888, 1894–1904, 1934–1952, 1971–1978, and 1995–2004, which corresponded to the negative phase of the Pacific Decadal Oscillation over the past century. Our findings provide new evidence for historical changes in TCs in the SCS and a valuable constraint for predicting future TC variations. Key Points: The low residual oxygen isotope values of coral strongly respond to tropical cyclones in the South China SeaExcluding temperature, we reconstruct tropical cyclone activities in the South China Sea over the past century using coral oxygen isotopesPacific Decadal Oscillation modulates tropical cyclones in the South China Sea over the past century [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]

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]

Significant differences in environmental conditions between nearshore and offshore coral reefs lead to different symbiotic zooxanthellae density (ZD) of corals. From the perspective of energy supply, different ZD lead to different energy provided by the photosynthesis of zooxanthellae to the coral host. However, few studies have conducted a comparative analysis of coral trophic status (i.e., autotrophic photosynthesis of zooxanthellae and heterotrophic feeding of the host) between nearshore and offshore reefs. In this study, 70 coral samples of Favia palauensis were collected from the nearshore Sanya reefs, and offshore Xisha, Nansha reefs in the South China Sea (SCS). The ZD, δ15N of seawater particulate organic matter (δ15NPOM), host tissue (δ15Nh), zooxanthellae (δ15Nz) and the difference between δ15Nh and δ15Nz (i.e., △h−z 15N = δ15Nh−δ15Nz) were measured and statistically analyzed. The purpose is to elucidate the differences in the energy maintenance mechanisms of corals between nearshore and offshore reefs. Results show that ZD is significantly positively correlated with δ15Nz in all three reefs (p < 0.05). Corals in nearshore Sanya have higher autotrophic abilities than corals from offshore Xisha and Nansha reefs. By contrast, corals in offshore reefs are more dependent on heterotrophic feeding of the host to obtain energy. Our research demonstrate that corals can adjust their trophic status, especially improve their heterotrophic feeding intensity to meet energy needs under different environmental conditions. This dynamic adjustment of energy supply patterns is of great significance to improve coral's environmental adaptability. Plain Language Summary: Coral reefs are mainly divided into nearshore fringing reefs, barrier reefs and offshore atolls according to geographical distribution. Therein, ambient seawater in nearshore reefs are significantly affected by human activities and has relatively higher nutrient levels. In contrast, corals in offshore atolls are less affected by human activities as they are far away from the mainland, but they will be potentially affected by abnormal natural conditions. Different environmental conditions lead to significant differences in the physiological parameters of corals, such as symbiotic zooxanthellae density (ZD). From the perspective of energy supply, different ZD may lead to different energy provided by the photosynthesis of zooxanthellae to the coral host. However, few studies have conducted a comparative analysis of coral trophic status between nearshore and offshore reefs. Based on the δ15N labeling technology, we find that corals in nearshore fringing reefs have higher autotrophic abilities than corals from offshore reefs. By contrast, corals in the offshore reefs are more dependent on heterotrophic feeding of the host to obtain energy. This means that corals can adjust their trophic status to meet energy needs under different environmental conditions. This dynamic adjustment of energy supply patterns is of great significance to improve coral's environmental adaptability. Key Points: The δ15N can record coral trophic status changes between nearshore and offshore reefsCorals in nearshore reefs have higher autotrophic photosynthesis and heterotrophic energy than corals in offshore reefsCoral can independently adjust trophic status under different environmental conditions between nearshore and offshore [ABSTRACT FROM AUTHOR]

In recent decades, vanadium (V) levels in the environment have dramatically increased and will continue to rise alarmingly in the future. Conventional methods for assessing oceanic V typically only involved the collection of seawater and sediments, which significantly limited our understanding of the dynamics of oceanic V and its relationship to environment and climate. Here, the interannual temporal variations in surface seawater V levels in the northern South China Sea, were reconstructed utilizing a 137‐year record of V/Ca ratios from Porites lutea coral from Xiaodonghai Bay off Sanya. We observed that anthropogenic activities had a limited impact on the V levels of surface seawater in the study area, whereas V variability exhibited significant disparities across different time periods. Precipitation‐related submarine groundwater discharge (SGD) recharge was the primary driver of V variation prior to 1990, with tropical cyclones (TCs) playing a detectable role in total precipitation during this period. Conversely, changes in surface seawater V levels after 1990 were predominantly linked to increasingly intense upwelling driven by summer winds. The recharge processes in unconfined groundwater systems within the study area might generate favorable oxic and alkaline conditions, which facilitated mineral dissolution and/or desorption reactions that elevated V concentrations along the groundwater flow path, which were recorded in coral skeletal V/Ca ratios. This study confirmed the seawater V was dominated by SGD and upwelling near Sanya, providing essential evidence for comprehending the geochemical behavior of V and the impact of ocean‐atmospheric climate interactions on long‐term variations in seawater V levels. Plain Language Summary: Recently, the amount of vanadium (V) in our environment has skyrocketed and will only continue to increase. The traditional method of checking oceanic V levels involved collecting seawater and sediment, which provided an incomplete understanding of V behavior in different environments and its relationship to climate change. Therefore, we conducted a detailed study on surface seawater V levels in the northern South China Sea over several years. We observed that human activities had little impact on the surface seawater V levels in our study area, but there were significant variations in V changes across different temporal intervals. Before 1990, precipitation‐induced submarine groundwater discharge was the main cause of changes in V, while tropical cyclones played a minor role in rainfall. However, after 1990, surface seawater V levels were significantly affected by stronger upwelling caused by summer winds. Groundwater systems in the study area may have caused mineral dissolution or desorption reactions, leading to increased V concentrations detected in corals. We looked into where V comes from in surface seawater and what factors affect it, giving us some solid proof on how V behaves chemically and how ocean‐atmospheric climate changes can impact its levels over time. Key Points: Coral vanadium (V)/Ca ratios exhibit a remarkable correlation with submarine groundwater discharge (SGD) prior to 1990Enhanced upwelling activity may modify the relationship between surface seawater V levels and SGDTropical cyclones amplify the impact of SGD on seawater V level by causing heavy precipitation [ABSTRACT FROM AUTHOR]

The strontium‐to‐calcium (Sr/Ca) ratio of coral is widely used as a paleothermometer for tropical sea surface temperature (SST) at subannual resolutions. The fidelity of Sr/Ca‐derived SST estimates largely depends on the properties of the monthly age model selected. Here, we develop a revised monthly age model for coral Sr/Ca estimation from the northern South China Sea based on specific tie points and evaluate the impacts of different age models on SST reconstruction. Our results indicate that our revised monthly age model reduces conversion errors by up to nearly 20%–30% relative to previously reported values, thereby improving the precision of seasonal SST estimates and enabling better paleoclimate interpretations, such as El Niño–Southern Oscillation variability. The revised monthly age model provides a generalized approach for developing more robust seasonal resolution Sr/Ca–SST reconstructions in other regions. Key Points: We develop a revised monthly age model for coral Sr/Ca‐derived sea surface temperature estimatesThe new age model reduces errors by up to 20%–30% relative to previously reportedThe new age model improves the accuracy of the South China Sea climate and El Niño–Southern Oscillation reconstructions [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]

High-latitude coral communities have attracted much attention due to their potential as refuges during global climate change. However, this function is being constrained by the combined pressure of global climate and anthropogenic activities. To determine how the reef has developed, we conducted a long-term monitoring study on coral communities along the Bailong Peninsula in the northern South China Sea. The results showed that the distribution area of corals was 3.67 km2 and that corals extended about 4.7 km along the coastline. The coral distribution pattern is scattered and uneven. Our results showed that the growth of reef-building corals and coral recruitment are improving, indicating that coral recruitment plays an important role in regulating the structure of adult coral populations and promoting the development of coral communities. Bailong Peninsula is expected to become a refuge for corals provided that human activities impacting coral restoration potential are controlled. [ABSTRACT FROM AUTHOR]

As a typical contaminant in fossil fuels, vanadium has undergone dramatically increase due to the impacts from human activities. The excessive anthropogenic vanadium emissions have altered natural geochemical processes. However, research on vanadium geochemistry in the ocean, which is the major sink for vanadium mobilized on land, has lagged much behind relative to those of other contaminants. Here we reconstruct historical patterns of vanadium in surface seawater utilizing a high-resolution temporal record (1984–2015 CE) of geochemical proxies (vanadium/calcium (V/Ca), δ18O, δ13C, etc.) in a Porites coral colony from an offshore non-estuarine island in the northern South China Sea (SCS). We find a close association between the surface winds and the temporal patterns of coral V/Ca ratios, revealing great potential for documenting prehistoric changes in tropical cyclones. Moreover, the coral V/Ca record remarkably captured the temporal patterns of anthropogenic activities, offering a valuable and novel tool for testing the efficacy of specific legislation on pollution prevention. Combined with the other reported coral vanadium data, we recovered the historical changes in vanadium emissions in the northern SCS and identified that changing industrial practices should be responsible for the differences in anthropogenic vanadium emissions at each stage. Our results demonstrated that the coral V/Ca ratio at a high temporal resolution has great potential as a new and direct geochemical proxy for tropical cyclones after removing artificial disturbances in the ocean. [ABSTRACT FROM AUTHOR]

Life-history strategies play a critical role in susceptibility to environmental stresses for Scleractinia coral. Metabolomics, which is capable of determining the metabolic responses of biological systems to genetic and environmental changes, is competent for the characterization of species' biological traits. In this study, two coral species (Pocillopora meandrina and Seriatopora hystrix in the South China Sea) with different life-history strategies ("competitive" and "weedy") were targeted, and untargeted mass spectrometry metabolomics combined with molecular networking was applied to characterize their differential metabolic pathways. The results show that lyso-platelet activating factors (lyso-PAFs), diacylglyceryl carboxyhydroxymethylcholine (DGCC), aromatic amino acids, and sulfhydryl compounds were more enriched in P. meandrina, whereas new phospholipids, dehydrated phosphoglycerol dihydroceramide (de-PG DHC), monoacylglycerol (MAG), fatty acids (FA) (C < 18), short peptides, and guanidine compounds were more enriched in S. hystrix. The metabolic pathways involved immune response, energy metabolism, cellular membrane structure regulation, oxidative stress system, secondary metabolite synthesis, etc. While the immune system (lysoPAF) and secondary metabolite synthesis (aromatic amino acids and sulfhydryl compounds) facilitates fast growth and resistance to environmental stressors of P. meandrina, the cell membrane structure (structural lipids), energy storage (storage lipids), oxidative stress system (short peptides), and secondary metabolite synthesis (guanidine compounds) are beneficial to the survival of S. hystrix in harsh conditions. This study contributes to the understanding of the potential molecular traits underlying life-history strategies of different coral species. [ABSTRACT FROM AUTHOR]

Cadmium (Cd) geochemical cycle plays a significant role in the composition and function of the marine ecosystem. Skeletal cadmium‐to‐calcium (Cd/Ca) ratios in hermatypic corals have been applied to reconstruct the historical changes of oceanic and climatic processes, yet there was no systematic evaluation of this tracer's natural variability in high resolution over time. Here, we reported a coral skeletal Cd/Ca record in monthly resolution from 1999 to 2008 CE and reconstructed the history of Cd contents in surface seawater in the northern South China Sea. A significant seasonal variation (higher in the winter but lower in the summer) of Cd contents in surface seawater can be identified. We found that the seasonal variations in coral skeletal Cd/Ca ratios exhibited significant trends coupled with the surface wind speeds, indicating that strong winds had likely driven the vertical seawater mixing process and then induced the process of sediment remobilization on the shelf, which significantly increased Cd contents in surface seawater. The reduction in Cd contents in surface seawater due to biological processes might be masked by the impacts of surface winds. Importantly, we also observed that coral skeletal Cd/Ca records in the winter showed significant correlations with the winter monsoon index, highlighting the possibility as a new proxy of winter monsoon in the non‐upwelling shelf environments. Plain Language Summary: Cadmium (Cd) is a nutrient element absorbed by phytoplankton in surface seawater, showing a distribution of low surface content and high bottom content. In order to determine the seasonal variations of Cd content in surface seawater in the northern South China Sea, we analyzed the coral skeletal Cd/Ca ratios in monthly resolution (1999–2008 CE) and finally concluded that the process of sediment resuspension and vertical mixing of seawater driven by winter monsoons is the main reason. The reduction of Cd contents in surface seawater due to biological processes was masked by the impacts of surface winds. In the past, Cd content in seawater/coral has been used to track the history of upwelling and El Niño Southern Oscillation, but the potential indicative on winter monsoon has not been reported so far. Considering that the trend between coral skeletal Cd/Ca ratios and average wind speeds showed a great agreement, we suggested that coral skeletal Cd/Ca ratios in the winter might be a potential proxy of winter monsoon. Key Points: Coral skeletal Cd/Ca ratios in monthly resolution were first reportedSeawater mixing process and sediments resuspension caused by winter monsoon are responsible for Cd seasonal variations in surface seawaterCoral skeletal Cd/Ca ratios have potential to be a novel proxy for winter monsoon [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]

Keywords: coral holobiont; physiological regulation; homeostasis; heat stress; nutrient; microbiome; Symbiodiniaceae; bleaching EN coral holobiont physiological regulation homeostasis heat stress nutrient microbiome Symbiodiniaceae bleaching 1 4 4 05/12/22 20220510 NES 220510 Scleractinian corals form the biological and physical framework for the most productive marine ecosystem in the oligotrophic oceans and play an important role in oceanographic processes ([3]). This paper provides further evidence that corals and their associated Symbiodiniaceae and bacteria communities engage in highly coordinated metabolic interactions that are crucial for coral holobiont health, homeostasis, and heat tolerance. Data further showed that coral-associated diazotrophs were common among coral demonstrate the potential importance of N SB 2 sb -fixing bacteria as the source of N-nutrient for corals, which live in the oligotrophic environment. These factors have important effects on coral holobionts tolerance; however, there are some differences between regions, coral species and even coral individuals ([26]; [27]; [29]). [Extracted from the article]

Ocean acidification is one of many stressors that coral reef ecosystems are currently contending with. Thus, understanding the response of key symbiotic microbes to ocean acidification is of great significance for understanding the adaptation mechanism and development trend of coral holobionts. Here, high-throughput sequencing technology was employed to investigate the coral-associated bacteria and Symbiodiniaceae of the ecologically important coral Acropora valida exposed to different pH gradients. After 30 days of acclimatization, we set four acidification gradients (pH 8.2, 7.8, 7.4, and 7.2, respectively), and each pH condition was applied for 10 days, with the whole experiment lasting for 70 days. Although the Symbiodiniaceae density decreased significantly, the coral did not appear to be bleached, and the real-time photosynthetic rate did not change significantly, indicating that A. valida has strong tolerance to acidification. Moreover, the Symbiodiniaceae community composition was hardly affected by ocean acidification, with the C1 subclade (Cladocopium goreaui) being dominant among the Symbiodiniaceae dominant types. The relative abundance of the Symbiodiniaceae background types was significantly higher at pH 7.2, indicating that ocean acidification might increase the stability of the community composition by regulating the Symbiodiniaceae rare biosphere. Furthermore, the stable symbiosis between the C1 subclade and coral host may contribute to the stability of the real-time photosynthetic efficiency. Finally, concerning the coral-associated bacteria, the stable symbiosis between Endozoicomonas and coral host is likely to help them adapt to ocean acidification. The significant increase in the relative abundance of Cyanobacteria at pH 7.2 may also compensate for the photosynthesis efficiency of a coral holobiont. In summary, this study suggests that the combined response of key symbiotic microbes helps the whole coral host resist the threats of ocean acidification. [ABSTRACT FROM AUTHOR]

Beachrock is considered a good archive for past sea-levels because of its unique formation position (intertidal zone). To evaluate sea-level history in the northern South China Sea, three well-preserved beachrock outcrops (Beigang, Gongshanbei, and Hengling) at Weizhou Island, northern South China Sea were selected to examine their relative elevation, sedimentological, mineralogical, and geochemical characteristics. Acropora branches with well-preserved surface micro-structures were selected from the beachrocks and used to determine the ages of these beachrocks via U-series dating. The results show that the beachrocks are composed of coral reef sediments, terrigenous clastics, volcanic clastics, and various calcite cements. These sediments accumulated in the intertidal zone of Weizhou Island were then cemented in a meteoric water environment. The U-series ages of beachrocks from Beigang, Gongshanbei, and Hengling are 1712–768 ca. BP, 1766–1070 ca. BP, and 1493–604 ca. BP (before 1950 AD) respectively. Their elevations are 0.91–1.16 m, 0.95–1.24 m, and 0.82–1.17 m higher than the modern homologous sedimentary zones, respectively. Therefore, we concluded that the sea-level in the Meghalayan age (1766–604 ca. BP) was 0.82–1.24 m higher than the present, and that the sea-level over this period showed a declining trend. [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]

Summary: Environmental conditions between the outer reef slope (ORS) and lagoon in tropical atolls are significantly different, but the variations of juvenile coral‐microbiomes in the two environments and their relationship with coral thermal acclimatization are poorly understood. We explored this issue based on local water conditions and the microbiome of juvenile corals in the ORS and lagoon in the central South China Sea. Coral‐symbiotic Symbiodiniaceae showed significant differences among coral species; Pocillopora verrucosa and Pachyseris rugosa in the ORS, and Acropora formosa in the lagoon were dominated by Durusdinium, but other corals were dominated by Cladocopium. Although A. formosa in the ORS were dominated by Cladocopium (C3u), they were dominated by Durusdinium (D1/D1a) and Cladocopium (C50) in the lagoon. Other coral species were both dominated by Cladocopium in the lagoon and ORS. The relative abundance of bacteria in the Deinococcus–Thermus was generally higher in the lagoon corals than in the ORS corals. Our study indicates that P. verrucosa, P. rugosa and Porites lutea may have high thermal tolerance based on the relatively high abundance of heat‐tolerant Durusdinium and Thermus scotoductus. Likewise, A. formosa in the lagoon may acclimatize to the thermal environment based on a high relative abundance of heat‐tolerant Durusdinium. [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]

Scleractinian corals exhibit a dual trophic pattern of autotrophic photosynthesis and heterotrophic predation. However, whether corals can adjust their trophic status under contrasting environmental conditions remains unclear. In our study, 70 scleractinian corals (Favia palauensis) were collected from Sanya and the Xisha and Nansha Islands in the South China Sea. We measured the zooxanthellae density (ZD) and δ13C of zooxanthellae (δ13Cz) and host tissue (δ13Ch) and analyzed the difference between δ13Ch and δ13Cz (i.e., Δh−z 13C). The relatively high ZD and δ13Cz values in the samples from Sanya indicate that these corals might have higher photosynthetic rates and autotrophic abilities than those from Xisha and Nansha. In contrast, the relatively low δ13Ch and Δh−z 13C values in the samples from Xisha and Nansha suggest that these corals might have a higher heterotrophic ability than those from Sanya. In addition, we tested the coral tissue biomass and skeletal δ13C (δ13Cs) in the samples from Sanya and examined their correlations with Δh−z 13C. The results showed a negative correlation, indicating that the more the organic material produced by the coral, the stronger its heterotrophic ability. Our results show that corals can adjust their trophic status under different environmental and physiological conditions, which is essential for increasing their adaptability to different environmental conditions. [ABSTRACT FROM AUTHOR]

The El Niño–Southern Oscillation (ENSO) dominates interannual climate variability worldwide and has important environmental and socio‐economic consequences. However, determining the evolution of ENSO variability and its long‐term response to climate forcing remains an ongoing challenge owing to the limited instrumental records. In this study, we quantified ENSO variability via an empirically calibrated threshold and sliding variance windows using monthly sea‐surface temperature (SST) anomalies based on Porites coral Sr/Ca records from the Xisha Islands in the northern South China Sea. Instrumental SST anomalies from the Xisha Islands correctly captured increasing ENSO variability in the twentieth century, with ENSO detection skills similar to those for Niño3.4 regions. Coral Sr/Ca‐SST anomalies can also serve as sensitive and robust proxies for ENSO variability. Sub‐fossil coral Sr/Ca‐SST anomalies indicated intensified ENSO variability at the end of the Medieval Climate Anomaly (MCA) from 1149 to 1205 ± 4.9 (2σ) Common Era (CE). Combining our records with other ENSO‐sensitive proxy reconstructions from the tropical Pacific, we observed fluctuating ENSO variability during the MCA and intensified ENSO variability for the late MCA. Considering the fewer and low intensity fluctuations associated with external climate forcing and the absence of a coherent temporal correspondence of ENSO activity with solar irradiance and volcanic eruption during the MCA, we hypothesized that the internal dynamics of the climate system play a prominent role in modulating ENSO variability and its evolution, which is supported by unforced climate model simulations and coral reconstructions across the tropical Pacific. Key Points: Coral Sr/Ca‐sea‐surface temperature variations from the northern South China Sea are a reliable proxy of El Niño–Southern Oscillation variabilityIntensified El Niño–Southern Oscillation variability occurred during the late Medieval Climate AnomalyUnforced internal variability within the climate system likely play a prominent role in modulating El Niño–Southern Oscillation variability [ABSTRACT FROM AUTHOR]

Epigenetic modifications based on DNA methylation can rapidly improve the potential of corals to adapt to environmental pressures by increasing their phenotypic plasticity, a factor important for scleractinian corals to adapt to future global warming. However, the extent to which corals develop similar adaptive mechanisms and their specific adaptation processes remain unclear. Here, to reveal the regulatory mechanism by which DNA methylation improves thermal tolerance in Pocillopora damicornis under fluctuating environments, we analyzed genome-wide DNA methylation signatures in P. damicornis and compared the differences in the methylation and transcriptional responses of P. damicornis from fluctuating and stable environments using whole-genome bisulfite sequencing and nanopore-based RNA sequencingtranscriptome sequencing. We discovered low methylation levels in P. damicornis (average methylation 4.14%), with CpG accounting for 74.88%, CHH for 13.27%, and CHG for 11.85% of this methylation. However, methylation levels did not change between coral samples from the fluctuating and stable environments. The varied methylation levels in different regions of the gene revealed that the overall methylation level of the gene body was relatively high and showed a bimodal methylation pattern. Methylation occurs primarily in exons rather than introns within the gene body In P. damicornis , there was only a weak correlation between methylation and transcriptional changes at the individual gene level, and the methylation and gene expression levels generally exhibited a bell-shaped relationship, which we speculate may be due to the specificity of cnidarian species. Correlation analysis between methylation levels and the transcriptome revealed that the highest proportion of the top 20 enriched KEGG pathways was related to immunity. Additionally, P. damicornis collected from a high-temperature pool had a lower metabolic rate than those collected from a low-temperature pool. We hypothesize that the dynamic balance of energy–expenditure costs between immunity and metabolism is an important strategy for increasing P. damicornis tolerance. The fluctuating environment of high-temperature pools may increase the heat tolerance in corals by increasing their immunity and thus lowering their metabolism. • Scleractinian corals are more tolerant in fluctuating reef-dwelling environments. • Low methylation levels in P. damicornis. • Thermal acclimation increase tolerance in corals by increasing coral immunity and thus lowering their metabolism. • Dynamic balance of energy–expenditure costs between immunity and metabolism is an important strategy. [ABSTRACT FROM AUTHOR]

Great efforts have been made to understand the impacts of the 1997 Indonesian wildfires on land ecology and human health, yet its influences on marine environments are poorly understood owing to sparse observations. Here we present weekly to monthly resolution coral δ13C records across the South China Sea in conjunction with a regional ocean general circulation model to study the effects of these cataclysmic wildfires on nearby oceans. Large and persistent negative δ13C excursions were found in corals from the interior South China Sea several months after the wildfires. This delay indicates that the 13C‐depleted ash fallout was distributed by ocean currents rather than direct atmospheric transport, which is supported by the results of our numerical model. Thus, the impacts of tropical wildfires on marine ecology may last for 1 year and extend thousands of kilometers from the burning center. Plain Language Summary: Tropical wildfires, intensified by extreme weather events and land use change, have continued to increase during this century, causing serious damage to the rainforests and local economies as well as human suffering and death. The threats of tropical wildfires to nearby oceans have been given little attention and are largely unknown because of limited observations. Here we use multiple coral records with weekly to monthly resolution from five sites near and around the South China Sea and a computer model to study the influences of the 1997 Indonesian wildfires on the ocean. We find a previously unrecognized ash fallout signal recorded in the corals in the interior of the South China Sea. Our model results indicate that the penetration of ash fallout into the South China Sea occurred through ocean currents rather than direct atmospheric transport. This study shows that the impacts of tropical wildfires on oceans may last for 1 year and can extend for thousands of kilometers from the source. We also show the potential for using coral records with computer modeling to identify and study the frequency and extent of past megafires. Key Points: A large negative excursion in δ13C values during summer–winter 1998 was detected in corals from the interior of the South China SeaModel shows that ash fallout from the 1997 Indonesian wildfires distributed by currents was responsible for the negative δ13C excursionNetworks of coral δ13C have great potential to record past pantropical wildfires [ABSTRACT FROM AUTHOR]

Coral reef ecosystems cannot operate normally without an effective nitrogen cycle. For oligotrophic coral reef areas, coral-associated diazotrophs are indispensable participants in the nitrogen cycle. However, the distribution of these diazotrophs and the correlation with the physical and chemical variables of the surrounding seawater remain unclear. To this end, 68 scleractinian coral colonies were sampled from 6 coral reef areas with different environmental variables in the South China Sea to investigate the composition of associated diazotrophs based on nifH gene amplification using high-throughput sequencing. The six coral reefs can be clearly divided into two types (fringing reefs and island reefs), are affected by varying degrees of human activities and are located at different latitudes from 9°20'06"N to 22°34'55"N with different seawater temperatures. Alpha- and beta-diversity analyses showed that the distribution of diazotrophs among coral reefs exhibited significant geographical fluctuations (p ≤ 0.05) and non-significant interspecific fluctuations (p > 0.05). The predominant bacterial phyla included Proteobacteria, Chlorobi, Cyanobacteria, and two unclassified phyla. Chlorobi exhibited a relative abundance of 47–96% in coral samples from the high-latitude Daya Bay fringing reef affected by eutrophication. Unclassified bacteria II, with a relative abundance of 28–87%, was found in all coral samples from the midlatitude Luhuitou fringing reef affected by eutrophication. However, unclassified bacteria I and Proteobacteria dominated (>80% relative abundance) in most of the coral samples from the Weizhou Island fringing reef, which is far from land, and three island reefs (Huangyan Island, Xinyi Reef, and Sanjiao Reef) at relatively low latitudes. At the genus level, some core diazotrophs were found in different coral sample groups. In addition, correlation analysis with various environmental variables revealed that the variables were positively or negatively correlated with different diazotrophic genera. Coral-associated diazotrophs were common among coral individuals. However, their composition was closely related to the different environmental variables. These results provide insights into the geographical distribution characteristics of coral-associated diazotrophs and their evolutionary trends in response to environmental change in the South China Sea. [ABSTRACT FROM AUTHOR]

El Niño-Southern Oscillation (ENSO) is the largest source of interannual climate variability over the globe. Knowledge of ENSO variability back to the pre-instrumental period (and earlier) can help to enhance our understanding of its mechanisms and impacts. Here we use the coral record from the northern South China Sea, where the interannual climate variability is associated with ENSO activity, to reconstruct ENSO variability over the past 150 years. Our record, together with other ENSO chronologies, indicated not only the weakened ENSO activity between ~1930s and ~1960s but also the larger variations of El Niño activity relative to that of La Niña based on the sliding window method. The comparison between the SCS and Pacific ENSO band coral records revealed the relatively stronger climatic coupling between the SCS and the equatorial central-eastern Pacific compared with that between SCS and western Pacific, indicating the reliability of teleconnection relationship between the SCS climate and ENSO. Our result indicated that the coral δ 18O record from the SCS could contribute to the pan-Pacific ENSO reconstructions. [ABSTRACT FROM AUTHOR]

The knowledges of trophic status (i.e., autotrophy/heterotrophy) of corals are helpful for understanding the mechanisms driving their environmental stress tolerance and bleaching resilience. We conducted this study to elucidate the differences in trophic status among corals of different genera and then explore the implication of trophic status for their different environmental stress tolerance. In this study, 117 coral samples including five genera (i.e., relatively stress‐tolerant massive genera Porites, Favites, and Favia and relatively vulnerable plating genus Pavona and branching genus Acropora) were collected from Weizhou Island, northern South China Sea. The symbiotic zooxanthellae density, δ13C of host tissues (δ13Ch), and symbiotic zooxanthellae (δ13Cz) were measured. Besides, the difference in values between δ13Ch and δ13Cz (i.e., △h‐z 13C) was analyzed. Our results indicated that both zooxanthellae density and δ13Cz values of Porites, Favites, and Favia are higher than that of Pavona and Acropora and indicate that autotrophic photosynthesis intensity of symbiotic zooxanthellae of Porites, Favites, and Favia is higher than that of Pavona and Acropora. Besides, both δ13Ch and △h‐z 13C of Porites, Favites, and Favia are lower than that of Pavona and Acropora and indicate a higher contribution of heterotrophic predation to the coral host of Porites, Favites, and Favia. These comparisons suggest that trophic status of relatively stress‐tolerant Porites, Favites, and Favia is more plasticity than vulnerable Pavona and Acropora, which is conducive to maintain a positive energy budget. This is one of important factors leading to different environmental stress tolerance and bleaching resilience among corals of different genera. Key Points: Autotrophic photosynthesis and heterotrophic predation of massive corals are higher than that of plating and branching coralsTrophic status of relatively stress‐tolerant massive corals is more plasticity than that of vulnerable plating and branching coralsDifferent trophic status is one of important factors leading to different environmental stress tolerance and bleaching resilience [ABSTRACT FROM AUTHOR]

The decadal variability of ocean primary productivity (PP) has mainly been studied using relatively short term satellite data, limiting its in‐depth analysis. Here, by removing the influence of the 13C Suess effect, we extract the PP signal on decadal timescales using a record of coral δ13C values from the northern South China Sea (NSCS), where the physical‐biogeochemical conditions are influenced largely by the winter monsoon. Our results show that the PP in the NSCS increased from 1851 to the 1920s and decreased from the 1920s to 2007, and large superimposed decadal‐scale changes were observed. The in‐phase relationship observed between the PP reconstruction and winter monsoon records suggests that the changes in the PP in the NSCS are linked to decadal‐scale changes in the winter monsoon. Considering that the winter monsoon intensity may weaken in the future under global warming, the PP in the NSCS might decrease in the coming decades. Plain Language Summary: Marine phytoplankton contributes roughly half of the global primary production and thus constitutes a vital component of the marine ecosystem that serves as the foundation of the marine food web. Due to the relatively short length of observation records, evaluations of the long‐term changes in ocean primary productivity (PP) have suffered from large uncertainties, which have hampered further discussion of the decadal variability of PP in a long‐term context. To discuss the decadal variability of the PP beyond the range of satellite observations, we present a coral‐based PP reconstruction since 1851 CE from the northern South China Sea (NSCS), where the physical‐biogeochemical conditions are influenced largely by the winter monsoon. Our results show considerable decadal variability of the PP in the NSCS superimposed on a pre‐1920s increase and a post‐1920s decrease. We also found an in‐phase relationship between the decadal variability of the reconstructed PP and the winter monsoon records over the past 150 years, indicating that the PP in the NSCS is linked to changes in the winter monsoon forcing on decadal timescales. Considering that the winter monsoon intensity may weaken in the future under global warming scenarios, the PP in the NSCS might decrease in the coming decades. Key Points: The decadal variability of primary productivity was extracted from a coral δ13C record by removing the influence of the 13C Suess effectThe coral‐based primary productivity reconstruction for the past 150 years showed a pre‐1920s increase and a post‐1920s decreaseThe primary productivity of the northern South China Sea varied in phase with the intensity of the winter monsoon on decadal timescales [ABSTRACT FROM AUTHOR]

"Refuge theory" suggests that global warming would be beneficial to corals in high latitude waters. In theory, then, the Weizhou Island reef (21°00′–21°10′N, 109°00′–109°15′E), which is located in a relatively high latitude area in the northern South China Sea, is a refuge for corals under global warming. Yet, the corals here have degenerated significantly. We investigated the ecological status of the Weizhou Island reef in 2015 and recorded 11 families, 22 genera, and 41 coral species. The mean living coral cover has decreased from ~ 42% in 1984 to ~ 10% in 2015 and there are many dead Acropora in the study area, especially at the reef flat. Coral assemblage structure has undergone degradation with the dominant group shifting from high complexity branching, foliaceous and massive colonies to a simpler group of massive morphologies. The only sign indicating the corals here benefiting from global warming is the occurrence of a large amount of juvenile Porites lutea (31.91% of the total population), which represents the recovery potential of the Weizhou Island reefs. Further analysis concludes that the main reason for the rapid degeneration is escalating anthropogenic impact, such as seawater pollution, unsustainable tourism activities, ongoing overfishing, all of which degrade the local ecological environment. It seems that intensive anthropogenic activities have weakened the "refuge" function significantly. Decreasing living coral cover as well as degraded assemblage structure all suggest that Weizhou Island offers limited potential as refuge habitat for corals in the context of global warming and intensive human activities. [ABSTRACT FROM AUTHOR]

It is well-known that the adaptability of coral-Symbiodiniaceae symbiosis to thermal stress varies among coral species, but the cause and/or mechanism behind it are not well-understood. In this study, we aimed to explore this issue based on zooxanthellae density (ZD) and Symbiodiniaceae genus/subclade. Hemocytometry and next-generation sequencing of the internal transcribed spacer region 2 (ITS2) marker gene were used to observe ZDs and Symbiodiniaceae genera/subclades associated with 15 typical coral species in the southern South China Sea (SCS). Average ZDs of all corals were in low levels, ranging from 0.84 to 1.22 × 106 cells cm−2, with a total of five Symbiodiniaceae genera, Symbiodinium, Cladocopium, Durusdinium, Fugacium , and Gerakladium , as well as 24 dominant subclades, were detected and varied among these coral species. Pocillopora verrucosa was dominated by Durusdinium (subclade D1/D1a), and other colonial corals were dominated by Cladocopium , but the subclades were varied among these species. Porites lutea and Montipora efflorescens were dominated by C15, and Echinopora lamellosa, Hydnophora exesa , and Coscinaraea exesa were dominated by C40. Acropora corymbosa, Merulina ampliata , and five species of Faviidae were mainly associated with Cladocopium types of C3u and Cspc. In contrast to other colonial corals, the dominant subclade of solitary Fungia fungites was C27, with high host specificity. Our study indicates that coral thermal stress adaptability is mainly affected by dominant Symbiodiniaceae type instead of ZD in the southern SCS. Some heat-sensitive corals, such as P. verrucosa corals, have acquired a high abundance of heat-tolerant Durusdinium to adapt to thermal stress. This could be the main reason for these corals becoming the dominant corals in this reef region. Background subclades analyses showed significant differences among coral species in subclade quantity and diversity. These suggest that numbers of coral species may have adapted to high environmental temperature by adopting various symbionts and/or associating with heat-tolerant Symbiodiniaceae. [ABSTRACT FROM AUTHOR]

Manmade antibiotics are emerging organic pollutants widely detected in the marine environment. In this study, 14 out of 19 target antibiotics were detected in corals collected from coastal and offshore regions in the South China Sea. The average total antibiotic concentrations (∑ 19 ABs) in the two regions were similar: 28 ng/g for coastal corals and 31 ng/g for offshore corals, based on dry tissue weight (dw). Fluoroquinolones (FQs) were predominant antibiotics in the coastal corals (mean ∑FQs: 18 ng/g dw), while sulfonamides (SAs) predominated in the offshore corals (mean ∑SAs: 23 ng/g dw). However, corals living in coastal regions tend to excrete more mucus than corals in offshore habitat. We found 53% by average of ∑ 19 ABs in the mucus of the coastal corals; while in offshore corals, most antibiotics (88% by average) were accumulated in the tissues. In addition, the tissue-mucus mass distribution differs among individual antibiotics. Sulfonamides were mainly accumulated in tissues while fluoroquinolones were present mainly in mucus. The results of this study suggest that mucus played an important role in the bioaccumulation of antibiotics by corals. It may resist the bioaccumulation of antibiotics by coral tissue, especially for the coastal corals. Additionally, corals were compared with other marine biotas in the study area and found to be more bioaccumulative towards antibiotics. Image 1 • The study first studied man-made antibiotic contamination in coral tissue and mucus. • Fourteen antibiotics were detected in corals at the level of 10−2 - 101 ng/g dw. • Tissue-mucus distribution of antibiotics differs between coastal and offshore corals. • Coastal coral mucus may resist accumulation of antibiotics by coral tissue. • Corals have high bioaccumulation factors because of their large surface area. Man-made antibiotics were first reported in coral tissue and mucus, which paly different roles in bioaccumulation of antibiotics. [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]

Coral reefs are continuing to decline worldwide due to anthropogenic climate change. The study of the molecular diversity and biogeographical patterns of Symbiodiniaceae, is essential to understand the adaptive potential and resilience of coral–algal symbiosis. Next generation sequencing was used to analyze the Symbiodiniaceae rDNA internal transcribed spacer 2 marker genes from 178 reef-building coral samples in eight coral habitats across approximately 13° of latitude in the South China Sea (SCS). A total of three Symbiodiniaceae genera, Cladocopium , Durusdinium , and Gerakladium , as well as 31 dominant Symbiodiniaceae types, were identified. Symbiodiniaceae richness, diversity, and community composition varied according to latitude; intermediate and low latitude coral reefs (IR and LR) have higher Symbiodiniaceae richness and diversity than high latitude coral habitats (HC and HR). A PERMANOVA analysis found significant differences in the Symbiodiniaceae community composition in the SCS (F = 14.75, R 2 = 0.20, p = 0.001 < 0.01). The major dominant Symbiodiniaceae types were C1 in the HC and the HR, C1/Cspc/C50/C15 and D1 in the IR, and C3u and C15 in the LR. Canonical correspondence analysis showed that the relative abundance of different Symbiodiniaceae types is affected by multiple environmental factors. Phylogenetic analysis indicated that the Symbiodiniaceae type Cladocopium , which shared common ancestors, shows similar environmental adaptability. Based on these results, we suggest that coral host species played a relatively small role in the identity of the dominant Symbiodiniaceae type. Therefore, the biogeographical patterns of Symbiodiniaceae may be mainly affected by environmental factors. Our research provides a comprehensive overview of the biogeography of Symbiodiniaceae in the SCS, where coral communities and reefs are widely distributed across different latitude regions and have variable environmental conditions. Our data will provide support for further study of the regional diversification of Symbiodiniaceae and the ecological resilience of the coral-Symbiodiniaceae symbioses. [ABSTRACT FROM AUTHOR]

Abstract As a natural laboratory for testing various paleoclimatological and paleoceanographic hypotheses, the Pliocene-Pleistocene stratigraphic sequence from a full-coring major scientific drilling (Well CK2 with total length: 928.75 m) in Yongle atoll of Xisha Islands, northern South China Sea was studied. The Petrography of studied section of the core was dominated by reef carbonate limestones which contain abundant shallow marine organisms, e.g., coralline algae, corals, and foraminifera, which almost consisted of pure carbonates and preserved the primary geochemical message since Pliocene. According to the isolation of Well CK2 and the provenance of elements, the systematic changes and abrupt points of isotope and element geochemical compositions (rare earth element parameters, SiO 2 content, Al/Ti ratio, etc.) from coral reef carbonate record presented the elevated terrigenous influences related to the East Asian monsoon after ~2.6 Ma BP, as part of the Pliocene-Pleistocene climate transition. Our results demonstrated that the continental weathering controlled by the East Asian summer monsoon dominated the surface seawater compositions of study area before ~2.6 Ma BP, when the winter monsoon was not strong enough to exert significant influences. Then, the effects of eolian dust associated with the intensity of East Asian winter monsoon, the precipitation driven by summer monsoon, and the global temperature, which played different roles at different periods after ~2.6 Ma BP, should be considered. Our results confirmed that the potentials and significances of shallow-water carbonate deposition from coral reefs, and provided a unique perspective to shed light on the carbonate record of climate changes. Highlights • Shallow-water carbonates from coral reefs can contribute to record of climate change. • Continental weathering dominated the surface seawater composition before ~2.6 Ma BP. • Eolian dusts contributed distinctly to surface seawater composition after ~2.6 Ma BP. [ABSTRACT FROM AUTHOR]

In this study, we examined radioactivity simultaneously in surface marine sediments and coral skeletons collected from 12 locations of the fringing and atoll reefs in the South China Sea. Radioactive level declined from the fringing reefs to atoll reefs because of input of terrigenous minerals in the fringing reefs. Radioactivity was higher in coral skeletons than in marine sediments because of the high 228Ra activity in coral skeletons. Additionally, an abnormally low 226Ra/238U activity ratio (<0.1) of marine sediments in coral reefs was attributed to the biological process of active uptake of 226Ra and 238U from seawater by coral polyps rather than the ingrowth process in the 238U-230Th-226Ra decay chain. Several radiological indices were evaluated in coral reefs and significantly lower than recommended values. Particularly, the average Ra eq in the atoll reefs was <5% of the world's average of Ra eq. Our results displayed typically radioactive status in coral reefs without close-in fallout of anthropogenic radionuclides. Unlabelled Image • Radioactivity in marine sediments declined from the fringing reefs to atoll reefs. • Radioactivity was higher in coral skeletons than in marine sediments. • The average Ra eq in the atoll reefs was <5% of the world's average Ra eq. • An abnormally low 226Ra/238U activity ratio (<0.1) occurred in coral reefs. [ABSTRACT FROM AUTHOR]

The coral symbiont Symbiodinium plays important roles in the adaptation of coral to environmental changes. However, coral- Symbiodinium symbiotic associations are not well-understood in the South China Sea (SCS) whilst considering environmental factors and host taxa. In this study, next-generation sequencing of the internal transcribed spacer region 2 (ITS2) marker gene was used to explore the symbiotic associations between Symbiodinium and five typical coral species across tropical and subtropical reef regions of the SCS. The results showed that Acropora sp., Galaxea fascicularis , Platygyra lamellina , and Sarcophyton glaucum exhibited distinct Symbiodinium compositions between tropical and subtropical reef regions, whereas Porites lutea had stable Symbiodinium compositions. More heterogeneous Symbiodinium compositions among different coral species were observed in the tropical region, but there were no statistically significant differences in Symbiodinium compositions among different coral species in subtropical reef regions. There was a correlation between the Symbiodinium compositions and environmental factors, except for the composition of P. lutea. Symbiodinium subclades D1, D2, C71, C71a, C21, C3b, and C161 were primarily explained by the seawater temperature, nitrate, ammonia, and phosphate. Several host-specific Symbiodinium subclades (e.g., C15, C15.6, and C91) were observed in P. lutea as well. The findings of this study demonstrate the relationship of Symbiodinium diversity with coral hosts and the environment are helpful for elucidating the adaptation of corals to global climate change and anthropogenic disturbance. [ABSTRACT FROM AUTHOR]

The importance of submarine groundwater discharge (SGD) is becoming increasingly recognized because of its potential significance as a source of dissolved species. To explore the probable coral geochemical signal of SGD and verify the validity of potential reliable proxies, multiple geochemical proxies over the last 137 years were identified from a Porites coral near a subterranean estuary in the northern South China Sea, where the SGD was reported to be the predominant flux of terrestrial waters to the coastal ocean. Results indicated that the SGD in the coastal zone was the dominant source of trace elements, especially REE and Ba, due to the various dissolution reactions occurring during groundwater flow in the karst terrain. The time- and frequency-domain comparison between the coral geochemical proxy and the local/regional precipitation indicated that coral REE/Ca ratios are predominantly impacted by the SGD associated with local precipitation, while coral Ba/Ca ratios are also affected by the primary productivity and allochthonous seawater Ba from surrounding areas. The REE signal from coral allows us to reconstruct the coastal surface seawater REE concentrations and the SGD rates on the coast of Sanya during 1870–2006. In a novel approach to developing a proxy for historic SGD to coastal waters, this study provides evidence that the coral REE/Ca record from the karst coast with large SGD has potentials to be a promising paleohydrological indicator. [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]

Although it is well established that different coral species have different susceptibilities to thermal stress, the reasons behind this variation are still unclear. In this study, 384 samples across five dominant coral species were collected seasonally between September 2013 and August 2014 at Luhuitou fringing reef in Sanya, Hainan Island, northern South China Sea, and their algal symbiont density and effective photochemical efficiency ( Φ ) were measured. The results indicated that both the Symbiodinium density and Φ of corals were subject to significant interspecies and seasonal variations. Stress-tolerant coral species, including massive Porites lutea and plating Pavona decussata, had higher symbiont densities but lower Φ compared to the vulnerable branching species of Acropora over the course of all four seasons. Seasonally, coral symbiont densities were the lowest during winter, while during the same period, Φ of corals was at the highest point. Further analysis suggested that dissolved inorganic nutrients and upwelling in the reef area were probably responsible for the observed seasonal variations in symbiont density. The fact that Porites lutea has the lowest Φ during all four seasons is likely related to their symbionts' lower capacity to provide required photosynthates for calcification. These results suggest that a coral's thermal tolerance is primarily and positively dependent on its symbiont density and is less related to its effective photochemical efficiency. [ABSTRACT FROM AUTHOR]

Coral bleaching has generally been recognized as the main reason for tropical coral reef degradation, but there are few long-term records of coral bleaching events. In this study, trace metals including chromium (Cr), copper (Cu), molybdenum (Mo), manganese (Mn), lead (Pb), tin (Sn), titanium (Ti), vanadium (V), and yttrium (Y), were analyzed in two Porites corals collected from Meiji Reef in the tropical South China Sea (SCS) to assess differences in trace metal concentrations in bleached compared with unbleached coral growth bands. Ti, V, Cr, and Mo generally showed irregular fluctuations in both corals. Bleached layers contained high concentrations of Mn, Cu, Sn, and Pb. Unbleached layers showed moderately high concentrations of Mn and Cu only. The different distribution of trace metals in Porites may be attributable to different selectivity on the basis of vital utility or toxicity. Ti, V, Cr, and Mo are discriminated against by both coral polyps and zooxanthellae, but Mn, Cu, Sn, and Pb are accumulated by zooxanthellae and only Mn and Cu are accumulated by polyps as essential elements. The marked increase in Cu, Mn, Pb, and Sn are associated with bleaching processes, including mucus secretion, tissue retraction, and zooxanthellae expulsion and occlusion. Variation in these trace elements within the coral skeleton can be used as potential tracers of short-lived bleaching events. [ABSTRACT FROM AUTHOR]

In this study, the contamination levels and seasonal variation of 22 PFASs were investigated in coastal reef-building corals (n = 68) from the northern South China Sea (SCS) during wet and dry seasons. Perfluorohexane sulfonate (PFHxS) was the predominant PFASs in all coral samples, representing 43% of the total PFAS. Long-chain PFASs, as well as PFAS alternatives, were frequently detected above the MQL (>88%) but showed relatively low concentrations compared to short-chain PFASs in most species and seasons. Seasonal variation of PFAS concentrations were observed in branching corals, indicating that the accumulation of PFASs may be associated with coral morphological structures. Interspecies differences in PFAS levels agree well with different bioaccumulation potentials among coral species. Redundancy analysis (RDA) showed that seasonal factor and coral genus could partly influence PFAS concentrations in coral tissues. In summary, our study firstly reported the occurrence of PFASs in coral communities from the SCS and highlights the necessity for future investigations on more toxicity data for coral communities. [Display omitted] • This is the first study about PFAS occurrence in coral reef ecosystem. • PFHxS was the predominant PFAS in all coral species. • Short-chain PFASs presented low detection frequencies but relatively high levels. • Branching species are particularly susceptible to PFAS exposure. • Seasonal factor and coral genus have certain effects on the accumulation of PFASs. [ABSTRACT FROM AUTHOR]

Compared with other ocean basins, annual resolution long-term series of sea-level records in the South China Sea (SCS) are still rare. Here we first analyzed the correlation mechanism between the oxygen stable isotopes (δ18O) of Porites coral and instrumental sea-level, sea surface salinity (SSS), sea surface temperature (SST) and rainfall of the SCS, and NINO 3.4 SST during 1980–2015 CE (Common Era); and we then quantitatively reconstructed sea-level at an annual resolution based on the coral δ18O in the SCS. The results show that the sea-level fell slightly (−0.73 ± 0.27 mm/yr) during 1850–1900 CE; then has continuously risen (1.31 ± 0.06 mm/yr) with a total increase of 152 ± 7 mm from 1900 to 2015 CE. On the inter-annual timescale, ENSO by mediating the overflow effect of seawater in the Western Pacific Warm Pool and the location of storm genesis in the western Pacific, indirectly affects sea-level, rainfall and SSS, which is finally reflected in changes in coral δ18O in the SCS. On the long-term timescale, especially since the 1900s, the gradual increase in SST and glaciers and ice sheets meltwater injected into the ocean due to the continuous warming mainly caused by human activities has not only promoted the rise of sea-level, but also caused a gradual negative bias of δ18O seawater (i.e. a gradual decrease of SSS), these processes are faithfully recorded by coral δ18O. Compared with SST, SSS or δ18O seawater may play a more important role in promoting the negative bias of coral δ18O in the SCS. Our study not only provides a better understanding of sea-level history and relationship mechanism around the SCS during 1850–2015 CE, but also may have the potential to provide a valuable proxy for the accurate quantitative reconstruction of sea-level at an annual resolution over a longer time scale for recent past periods. • On the interannual timescale, ENSO is the main driver of coral δ18O via its indirect effects on seawater δ18O in the SCS • On the long-term timescale, SST and injection of meltwater are the main driver of changes in coral δ18O and sea-level • Both SSS and SST are bridges between coral δ18O and sea-level • Coral δ18O is an ideal proxy to indicate sea-level, which has risen by 152 ± 7 mm in total from 1900 to 2015 CE in the SCS [ABSTRACT FROM AUTHOR]

Reconstructing the precise timing of sea level fall is important for understanding earth system during the geologic time. However, previous studies are limited by the absence of effective indicators for sea level fall. To overcome this limitation, we investigated the 87Sr/86Sr and petrologic features of carbonate strata in a 928.75-m-long coral reef core (CK2) from Xisha Islands, northern South China Sea and their relationship to sea level fluctuations. Ten exposed surfaces were identified by their petrologic features. The Mn and Sr content, Mn/Sr, Sr/Ca and δ18O suggest that most of the coral reef carbonate in the CK2 core preserved the original seawater 87Sr/86Sr except at intervals of 672–616, 596–575, 521–491, 423–414, and 192.5–161 m, which suffered alteration of meteoric water and showed higher 87Sr/86Sr during low sea level period. The 160 carbonate 87Sr/86Sr showed a continuously increasing trend, which is consistent with that of seawater over the same period. The continuously increasing 87Sr/86Sr curve of CK2 was divided into seven units by six 87Sr/86Sr hiatuses. Inconsistency between the sedimentation rate of CK2 and tectonic subsidence rate in the Xisha area suggest that the continuously increasing 87Sr/86Sr was controlled by the coupled effect of tectonic subsidence and sea level change, and the hiatus and higher 87Sr/86Sr interval in the 87Sr/86Sr curve were controlled by sea level fall. The higher 87Sr/86Sr usually produced during low sea level (<5 m), 87Sr/86Sr hiatus usually produced when sea level changed from deeper to shallower. The depth of hiatuses and higher intervals of 87Sr/86Sr correspond well with the exposed surface in CK2 core, verifying that the hiatuses and higher intervals of 87Sr/86Sr in the coral reef carbonate succession provide an effective indicator of sea level fall. Using this indicator and the CK2 87Sr/86Sr profile, we identified eleven large-scale sea level falls and three dramatic regressions during 19.6–0.2 Ma and constrained the age based on strontium isotope stratigraphy. These interpreted sea level falls are mostly consistent with other records of global sea level changes, confirming the reliability of our records. The 87Sr/86Sr profile of a coral reef carbonate is an efficient indicator of past sea level falls, including their timing. • 87Sr/86Sr ratios of carbonate in a coral reef core from Xisha Islands. • The core indicated five hiatuses and six anomalously high ratios in Sr isotope. • Hiatus and high ratios in reef carbonate Sr isotope were produced by sea level fall. • Eleven major lower sea level and three dramatic regressions found during 19.6–0.2 Ma. • This method provides an effective indicator of past sea level fluctuation. [ABSTRACT FROM AUTHOR]

Coral ingestion by crown-of-thorns starfish (COTS) is an important cause of coral reef degradation, although the impacts of COTS feeding on coral-associated microbial communities are not well understood. Therefore, in this study, we analyzed the coral tissue-weight, Symbiodiniaceae density (SD), bacterial community composition, and the predicted functions of bacterial genes associated with Pocillopora corals in healthy portions and feeding scars, following COTS feeding. Coral tissue-weight loss rate in the feeding scars was 71.3–94.95%. The SDs were significantly lower in the feeding scars, and the SD-loss rate was 92.05% ± 2.12%. The relative abundances of bacterial communities associated with Pocillopora corals after COTS feeding changed significantly and were almost completely reorganized at the phylum and genus levels. Analysis of the microbial metagenomic-functional capacities showed that numerous physiological functions of the coral-bacterial holobionts in the feeding scars were different, including amino acid metabolism, xenobiotic biodegradation and metabolism, lipid metabolism, membrane transport, signal transduction, and cell motility, and all these capacities could be corroborated based on metagenomic, transcriptomic or proteomic technologies. Overall, our research suggests that coral holobionts may be destroyed by COTS, and our findings imply that bacterial communities in feeding scars could affect the health of Pocillopora corals. [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]

We have previously shown that the coralline algal assemblages (CAAs) of the South China Sea may indicate post-Pliocene paleo-water depth changes; however, the pre-Pliocene coralline algal compositions and related sea-level significance remain to be elucidated. In the present study, we explore the Miocene coralline algal compositions and their role in recording the sea level, and show the distribution, classification, type, abundance, and diversity of Miocene coralline algae from 213 petrologic thin sections using a coral-reef carbonate sequence from the northern South China Sea. The diversity and abundance of coralline algae in the early and late Miocene are higher than those in the middle Miocene. Eleven genera were identified and grouped into seven CAAs, showing a water depth range from <5 m to >25 m. At 18.67–17.98 Ma, the coral reef was in a stagnation/drowning stage, with rising sea levels exceeding the development of the coral reef. At 17.98–16.78 Ma, the coral reef was in a rapid development during a rapid rise in sea level. Additionally, at 16.78–14.79 Ma, low coralline algal abundance was observed, and the corals were sporadically distributed at a relatively stable sedimentation rate, suggesting a decline in coral reef development. At 14.79–10 Ma, the coral reef development was remarkably slow or stagnant. Furthermore, at 10–5.3 Ma, the coral reefs developed slowly, indicating a gradual rise in the sea level. The sea-level changes indicated by coral reef development are consistent with the long-term global sea-level changes, indicating that the composition and assemblage of Miocene coralline algae can be used to accurately relay the sea-level history. • Eleven coralline algal genera were identified in a Miocene carbonate sequence. • Miocene coralline algae were divided into 7 coralline algal assemblages (CAAs). • Sea-level changes due to coral reef development were consistent with global values. [ABSTRACT FROM AUTHOR]

Few reconstructions of sea-surface temperature (SST) have focused on seasonal and interannual variability, two major components of the global climate system, due to the limited temporal resolution of proxies. This study presents a combined 228-year-long, monthly resolved strontium to calcium ratios (Sr/Ca) record covering the period of 2070–1740 a BP (years before 1950 CE (Common Era)) extracted from three U-series-based sub-fossil Porites corals located on the Xisha Islands, northern South China Sea (SCS). The composite time series allowed for accurate assessment of the natural range of SST variations during snapshots of the Roman Warm Period (RWP). Reconstructed SST revealed that the RWP was characterized by cooler conditions compared with the 20th century, consistent with climatic variations in the western Pacific and East Asia. The amplitude of SST seasonality was within the modern range, except for a lower value at 1980–1928 a BP. Interannual variability associated with El Niño–Southern Oscillation (ENSO) activity was enhanced by 39% relative to 1980–2014 CE. The results of the sliding window demonstrate that ENSO variability persistently strengthened during 2070–2010 a BP, followed by an overall fluctuating attenuation during 1980–1928 a BP. Then, the trends of rising first before descending twice during 1852–1800 a BP appeared. Furthermore, ENSO activity played a leading role in steering short-term changes in SST seasonality in the northern SCS, manifested as stronger ENSO activity with more frequent El Niño events and decreased SST seasonality. Considering that the frequency of extreme ENSO events may strengthen in the future under global warming, the climate in the northern SCS might become more variable and complex. • 228-year monthly SST derived from coral Sr/Ca for snapshots of the Roman Warm Period. • Cooler conditions and variable SST seasonality characterize the 2070–1740 a BP. • Stronger interannual variability and ENSO evolution for the RWP are investigated. • ENSO activity modulates changes of SST seasonality in the northern South China Sea. [ABSTRACT FROM AUTHOR]