Coral reefs: corals' adaptive response to climate change

The long-term response of coral reefs to climate change depends on the ability of reef-building coral symbioses to adapt or acclimatize to warmer temperatures, but there has been no direct evidence that such a response can occur. Here we show that corals containing unusual algal symbionts that are thermally tolerant and commonly associated with high-temperature environments are much more abundant on reefs that have been severely affected by recent climate change. This adaptive shift in symbiont communities indicates that these devastated reefs could be more resistant to future thermal stress, resulting in significantly longer extinction times for surviving corals than had been previously assumed.     

涠洲岛西局部造礁石珊瑚盖度和大小频率分布

本研究通过分析珊瑚大小频率分布,探讨涠洲岛西局部造礁石珊瑚群落结构特点和种群动态,以了解珊瑚的生活史和所经历的变化。所分析的珊瑚大小数据来自2021年2月24-26日开展的涠洲岛西珊瑚礁的造礁石珊瑚样带调查,利用对数转换将珊瑚大小变换为珊瑚迭代,然后作直方图对珊瑚大小频率分布加以分析。分析结果表明,片状的牡丹珊瑚属Pavona和块状的滨珊瑚属Porites、角蜂巢珊瑚属Favites、盘星珊瑚属Dipsastraea等在涠洲岛西局部珊瑚群落中占优势(珊瑚面积和珊瑚个数)。对数转换的大小频率分布呈正态分布,偏斜不明显,变异系数小于0.5,表明这些珊瑚目前处于正常的自然补充、生长和死亡过程。涠洲岛西局部造礁石珊瑚盖度降低,主要是因为个体大的枝状珊瑚锐减。     

Effects of temperature,hypoxia, ammonia and nitrate on the bleaching among three coral species

Coral bleaching, which is defined as the loss of colour in corals due to the loss of their symbiotic algae (commonly called zooxanthellae) or pigments or both, is occurring globally at increasing rates, and its harm becomes more and more serious during these two decades. The significance of these bleaching events to the health of coral reef ecosystems is extreme, as bleached corals exhibited high mortality, reduced fecundity and productivity and increased susceptibility to diseases. This decreased coral fitness is easily to lead to reef degradation and ultimately to the breakdown of the coral reef ecosystems. Recently, the reasons leading to coral bleaching are thought to be as follows: too high or too low temperature, excess ultraviolet exposure, heavy metal pollution, cyanide poison and seasonal cycle. To date there has been little knowledge of whether mariculture can result in coral bleaching and which substance has the worst effect on corals. And no research was conducted on the effect of hypoxia on corals. To address these questions, effects of tem- perature, hypoxia, ammonia and nitrate on bleaching of three coral species were studied through examination of morphology and the measurement of the number of symbiotic algae of three coral species Acropora nobilis, Palythoa sp. and Alveopora verrilliana. Results showed that increase in temperature and decrease in dissolved oxygen could lead to increasing number of symbiotic algae and more serious bleaching. In addition, the concentration of 0.001 mmol/L ammonia or nitrate could increase significantly the expulsion of the symbiotic algae of the three coral species. Except for Acropora nobilis, the numbers of symbiotic algae of other two corals did not significantly increase with the increasing concentration of ammonia and nitrate. Furthermore, different hosts have different stress susceptibilities on coral bleaching.     

涠洲岛珊瑚礁特点、演变及保护与修复对策的思考

涠洲岛珊瑚礁最早形成于约7 000年前的末次海侵,是中国大陆沿海成礁珊瑚分布最北缘的典型边缘珊瑚礁（Marginal Coral Reefs）,与区域内其他珊瑚礁缺少生态关联,其特征不同于热带珊瑚礁,即块状珊瑚占优势。20多年来,涠洲岛珊瑚礁持续退化,枝状珊瑚大量死亡,石珊瑚盖度急剧降低,反映珊瑚礁良好状况的指标生物如鹦嘴鱼、蝴蝶鱼等的数量大幅减少。近10年来,珊瑚虽未出现异常死亡现象,但珊瑚自然修复不明显,表明珊瑚补充不足。功能性植食性鱼类的缺乏,显示珊瑚礁恢复力不足。涠洲岛珊瑚礁国家级海洋公园的建立,体现了保护涠洲岛珊瑚礁的政治意愿。只有恢复鹦嘴鱼等功能性植食性鱼类的数量,才能恢复和增强珊瑚礁固有恢复力,进而促进涠洲岛珊瑚礁的自然恢复。人为修复可采取移植珊瑚断枝的方式,并以实现生态、经济和社会增益为目标,构建能应对人为干预且能持续演化的珊瑚礁。     

Using the Acropora digitifera genome to understand coral responses to environmental change

Despite the enormous ecological and economic importance of coral reefs, the keystone organisms in their establishment, the scleractinian corals, increasingly face a range of anthropogenic challenges including ocean acidification and seawater temperature rise. To understand better the molecular mechanisms underlying coral biology, here we decoded the approximately 420-megabase genome of Acropora digitifera using next-generation sequencing technology. This genome contains approximately 23,700 gene models. Molecular phylogenetics indicate that the coral and the sea anemone Nematostella vectensis diverged approximately 500 million years ago, considerably earlier than the time over which modern corals are represented in the fossil record (～240 million years ago). Despite the long evolutionary history of the endosymbiosis, no evidence was found for horizontal transfer of genes from symbiont to host. However, unlike several other corals, Acropora seems to lack an enzyme essential for cysteine biosynthesis, implying dependency of this coral on its symbionts for this amino acid. Corals inhabit environments where they are frequently exposed to high levels of solar radiation, and analysis of the Acropora genome data indicates that the coral host can independently carry out de novo synthesis of mycosporine-like amino acids, which are potent ultraviolet-protective compounds. In addition, the coral innate immunity repertoire is notably more complex than that of the sea anemone, indicating that some of these genes may have roles in symbiosis or coloniality. A number of genes with putative roles in calcification were identified, and several of these are restricted to corals. The coral genome provides a platform for understanding the molecular basis of symbiosis and responses to environmental changes.     

Emerged fossil corals on the coast of northwestern Hainan Island, China: Implications for mid-Holocene sea level change and tectonic uplift

Fossil corals are widespread on the coast of northwestern Hainan Island, China. Most of these corals are exposed during low tide levels, indicating that a higher sea level may have existed during their life time. The radiocarbon data introduced by this paper in combination with those from other sources show that the corals were alive mainly during the midto late-Holocene. Mid-Holocene sea levels 1-3 m higher than the present level were confirmed for adjacent coasts; however, the emerged corals on the investigated coast are not necessarily indicators of a higher eustatic sea level. They do predict relative sea levels of 1.5-3.4 m when using 1 m below the tidal datum as the upper limit of coral growth. However, using the Mean Lower Low Water as the coral’s upper growth limit, the relative sea level on the non-volcanic coast was almost as high as the present level, whereas that on the volcanic coast was 0.4-2.0 m higher than present. Therefore, the exposure of these dead coral heads and their discrepancy in elevation may be the result of tectonic uplift caused by volcanic activities. From these results, the highest sea level of upward coral growth must be defined first when using fossil corals to reconstruct past sea levels. Meanwhile, vertical tectonics have to be taken into account as a significant factor when conducting high-resolution sea level reconstruction, although the South China coast is located in a "tectonically stable" region.     

Assessment of coral bleaching using symbiotic zooxanthellae density and satellite remote sensing data in the Nansha Islands,South China Sea

Coral bleaching,characterized by a significant loss of symbiotic zooxanthellae,is the primary cause of mass coral mortality and reef degradation throughout the world.The characteristics,processes,and resistance of corals to bleaching varies significantly and is dependent on environmental conditions.We documented a mass coral bleaching event in June 2007 at the Meiji and Zhubi Reefs,Nansha Islands (NS),South China Sea using ecological surveys and measurement of coral zooxanthellae density and sea surface temperatures (SST).More than 35 species of corals (between 0-20 m in depth) were bleached.These bleached corals accounted for 15.6% of total corals in the investigated quadrats.The branching corals Pocillopora and Acropora were the most vulnerable species whereas the massive corals Porites and Favia were more tolerant of the high SSTs.Surprisingly,we found no evidence of bleaching in Agariciidae corals suggesting that this family is resistant to thermal stresses.The bleached corals had lost 72%-90% of their symbiotic zooxanthellae.Furthermore,corals that had no visual signs of bleaching had also lost 31%-53% of their zooxanthellae suggesting that most corals were experiencing the early stage of bleaching.The monthly mean SST during June 2007 was 30.8°C,the highest since 1998.Based on measurements of SST and the Hotspots and DHW data (NOAA),we conclude that it the extremely high SSTs triggered this coral bleaching event.Our results suggest that the previously accepted temperature thresholds used to predict coral bleaching based on satellite data are likely to underestimate the extent and intensity of coral bleaching,at least in the NS.     

涠洲岛珊瑚健康及其影响因子分析

根据2007年秋至2008年春间,涠洲岛6条主剖面15条断面的珊瑚和底质覆盖率、珊瑚补充量、死亡指数的一次两阶段的调查数据和同期4个季度月的水环境调查数据,结合以往的研究成果,初步分析了涠洲岛珊瑚健康状况及其影响因子。结果表明,调查断面珊瑚死亡指数平均为40.90%,珊瑚死亡发生于本次调查的两年前;珊瑚有恢复的迹象,但珊瑚补充量小,珊瑚补充量与活珊瑚+死珊瑚覆盖率显著相关(r=0.520,P<0.05);除汞、铅、锌和油类外,涠洲岛珊瑚礁海区水体的理化、生物因子符合一类海水水质标准,大体上呈微营养水平,春季的水体营养指数明显高于其他季节。涠洲岛珊瑚死亡率高和恢复慢,表明珊瑚处于退化中的亚健康状态。     

Heterotrophic plasticity and resilience in bleached corals

Mass coral bleaching events caused by elevated seawater temperatures have resulted in extensive coral mortality throughout the tropics over the past few decades. With continued global warming, bleaching events are predicted to increase in frequency and severity, causing up to 60% coral mortality globally within the next few decades. Although some corals are able to recover and to survive bleaching, the mechanisms underlying such resilience are poorly understood. Here we show that the coral host has a significant role in recovery and resilience. Bleached and recovering Montipora capitata (branching) corals met more than 100% of their daily metabolic energy requirements by markedly increasing their feeding rates and CHAR (per cent contribution of heterotrophically acquired carbon to daily animal respiration), whereas Porites compressa (branching) and Porites lobata (mounding) corals did not. These findings suggest that coral species with high-CHAR capability during bleaching and recovery, irrespective of morphology, will be more resilient to bleaching events over the long term, could become the dominant coral species on reefs, and may help to safeguard affected reefs from potential local and global extinction.     

Coral mucus functions as an energy carrier and particle trap in the reef ecosystem

Zooxanthellae, endosymbiotic algae of reef-building corals, substantially contribute to the high gross primary production of coral reefs, but corals exude up to half of the carbon assimilated by their zooxanthellae as mucus. Here we show that released coral mucus efficiently traps organic matter from the water column and rapidly carries energy and nutrients to the reef lagoon sediment, which acts as a biocatalytic mineralizing filter. In the Great Barrier Reef, the dominant genus of hard corals, Acropora, exudes up to 4.8 litres of mucus per square metre of reef area per day. Between 56% and 80% of this mucus dissolves in the reef water, which is filtered through the lagoon sands. Here, coral mucus is degraded at a turnover rate of at least 7% per hour. Detached undissolved mucus traps suspended particles, increasing its initial organic carbon and nitrogen content by three orders of magnitude within 2 h. Tidal currents concentrate these mucus aggregates into the lagoon, where they rapidly settle. Coral mucus provides light energy harvested by the zooxanthellae and trapped particles to the heterotrophic reef community, thereby establishing a recycling loop that supports benthic life, while reducing loss of energy and nutrients from the reef ecosystem.     

Communication arising. Is coral bleaching really adaptive?

From an experiment in which corals are transplanted between two depths on a Panamanian coral reef, Baker infers that bleaching may sometimes help reef corals to survive environmental change. Although Baker's results hint at further mechanisms by which reef-building corals may acclimatize to changing light conditions, we do not consider that the evidence supports his inference.     

Coral community changes in response to a high sedimentation event: A case study in southern Hainan Island

A high sedimentation event caused by dredging and dumping of sediment was recorded on Xiaodonghai Reef in Yulin Bay,southern Hainan Island,China.Significantly high sedimentation and constant light shading were observed during the sediment dumping event(SD Event).Using long-term video transects,we quantified coral community changes and responses to the SD Event between 2008 and 2010.The SD Event caused severe coral mortality on Xiaodonghai Reef at a depth of 6 m,while corals at 3 m were less affected.Total live coral cover at 6 m decreased from 54.3% to 14.8%,and Diploastrea heliopora replaced Galaxea fascicularis as the dominant coral species at 6 and 9 m.The density of juvenile corals also declined after the SD Event,especially for the genera Galaxea and Platygyra.However,the density of juvenile Porites and Pocillopora spp.slightly increased.Monitoring for 11 months after the SD Event indicated no recovery of coral communities on Xiaodonghai Reef.Long-term video transect data also revealed that mean live coral cover dramatically declined,from 30.5% in 2008 to 9% in 2010,while the dominant corals in Yulin Bay shifted to more tolerant coral species,such as massive Porites spp.and D.heliopora.The rapid coral community degradation in Yulin Bay between 2008 and 2010 was probably caused by high sediment deposition resulting from intensive dredging and land-clearing activities.These results highlight the necessity for an integrated watershed management to control sediment deposition on near-shore coral reefs.     

涠洲岛珊瑚礁健康评估

【目的】科学地判断涠洲岛珊瑚礁的现状,并找出造成目前现状的主要原因,为及时采取有效措施以维持涠洲岛珊瑚礁的健康提供有力依据。【方法】根据历年来有关涠洲岛珊瑚礁生长、演变、环境变化与社会发展的研究成果和最新的调查研究资料,采用HRHPI的指标体系法,从生态结构、生态功能、压力及社会经济等几个特征属性对珊瑚礁健康进行评估。【结果】涠洲岛的自然环境仍适合石珊瑚的生存,石珊瑚的种类丰度仍正常;珊瑚礁的覆盖率降低、分布范围缩小、生物多样性下降,尤其在浅海和局部人为影响大的区域,珊瑚死亡率大于补充率。【结论】涠洲岛的珊瑚礁处于衰退中的亚健康状态。过度捕捞、污染、物理损伤（踩踏、抛锚、水下工程等）和大尺度环境变化（如气候异常）等是造成涠洲岛珊瑚礁衰退的主要原因。     

Ecological consequences of major hydrodynamic disturbances on coral reefs

A recent tsunami and an apparent increase in the frequency of severe tropical storms underscore the need to understand and predict the ecological consequences of major hydrodynamic disturbances. Reef corals provide the habitat structure that sustains the high biodiversity of tropical reefs, and thus provide the foundation for the ecosystem goods and services that are critical to many tropical societies. Here we integrate predictions from oceanographic models with engineering theory, to predict the dislodgement of benthic reef corals during hydrodynamic disturbances. This generalizes earlier work, by incorporating colonies of any shape and by explicitly examining the effects of hydrodynamic gradients on coral assemblage structure. A field test shows that this model accurately predicts changes in the mechanical vulnerability of coral colonies, and thus their size and shape, with distance from the reef crest. This work provides a general framework for understanding and predicting the effects of hydrodynamic disturbances on coral reef communities; such disturbances have a major role in determining species zonation and coexistence on coral reefs, and are critical determinants of how coral assemblages will respond to changes in the frequency and intensity of tropical storms associated with a changing climate.     

Twenty-five years of change in scleractinian coral communities of Daya Bay （northern South China Sea） and its response to the 2008 AD extreme cold climate event

Coral reefs worldwide are becoming increasingly and detrimentally impacted upon by a variety of factors including significant climate changes, such as global warming and increased El Nino-Southern Oscillation activity. Generally, the persistence of coral reefs, especially at low-latitudes, is governed, in part, by sea surface temperatures not exceeding the critical limit （-30℃） at which mass mortality can occur. Thus, it is thought that corals living at high-latitudes （i.e., currently cooler sea surface temperatures） will likely respond more favourably to hypothesized future temperature increases than corals living at low-latitudes （i.e., currently warmer sea surface temperatures）. Consequently, high-latitude coral communities may have the potential to act as regions of refugia for many coral species in the face of potential future global warming. The Daya Bay （22°31′--22°50′N）, northern South China Sea, contains several high-latitude non-reefal coral communities and represents one of the most northerly distributions of scleractinian corals within the region. Significantly, Daya Bay has experienced dramatic warming in both air and sea surface temperatures throughout the past 50 years. In this paper, we analyze 25 years of change in the Daya Bay coral communities, based both on historic surveys and our latest 2006--2008 regional ecological surveys. Our results suggest that, contrary to predictions, there have been significant declines in coral cover within the Daya Bay during the past 25 years （i.e., 76.6% coral cover in 1983/1984 to only 15.3% coral cover by 2008）. Such changes also reflect a significant shift in the most abundant coral species, from Acropora pruinosa to Favites abdita. Most of the modern coral communities became established between 15 and 30 years ago, corresponding to a period of increased winter sea surface temperature. However, very few colonies have become established within the last 15 years, despite a more intense period of warming. By taking into account additional factors, we hypothesize that direct anthropogenic impacts, rather than climatic events, have both restricted the development, and drove the decline, of Daya Bay coral communities in the last 15 years. The Daya Bay has also been subjected to occasional extreme cold events during the past 50 years, with the most recent occurring in early 2008 （13 January-13 February）. During the 2008 cold event, the lowest air temperature reaches only 6.6℃, and the mean sea surface temperature for February fall to 〈 14℃, including six continuous days at 12.3℃. Significantly, the sea surface temperatures fall below the hypothesized critical lower temperature threshold （-13℃） that commonly leads to mass mortality in scleractinian coral communities. Surprisingly, our coral community surveys, conducted both before （August 2007） and after （late February 2008） the extreme 2008 cold event, demonstrate that the Daya Bay coral ecosystems are barely impacted upon during the cold period. Those observations suggest that the Daya Bay scleractinian coral communities have developed adaptations to low sea surface temperatures. Overall, our data support the hypothesis that high-latitude coral communities, such as Daya Bay, have the potential to act as areas of refugia for scleractinian corals in the advent of potential future global warming.     

Conventional taxonomy obscures deep divergence between Pacific and Atlantic corals

Only 17% of 111 reef-building coral genera and none of the 18 coral families with reef-builders are considered endemic to the Atlantic, whereas the corresponding percentages for the Indo-west Pacific are 76% and 39%. These figures depend on the assumption that genera and families spanning the two provinces belong to the same lineages (that is, they are monophyletic). Here we show that this assumption is incorrect on the basis of analyses of mitochondrial and nuclear genes. Pervasive morphological convergence at the family level has obscured the evolutionary distinctiveness of Atlantic corals. Some Atlantic genera conventionally assigned to different families are more closely related to each other than they are to their respective Pacific 'congeners'. Nine of the 27 genera of reef-building Atlantic corals belong to this previously unrecognized lineage, which probably diverged over 34 million years ago. Although Pacific reefs have larger numbers of more narrowly distributed species, and therefore rank higher in biodiversity hotspot analyses, the deep evolutionary distinctiveness of many Atlantic corals should also be considered when setting conservation priorities.     

Coral cover as a proxy of disturbance:A case study of the biodiversity of the hermatypic corals in Yongxing Island, Xisha Islands in the South China Sea

The Yongxing Island is the biggest island of Xisha Islands in the middle part of the South China Sea. It occupies 1.8 square kilometers of land area and has permanent residence on it. There are only a few papers on the coral community of the Xisha Islands, reporting the species composition, structure and zonations of the hermatypic coral community in 1970s. The present study describes the hermatypic coral community based on the quadrat survey after almost 30 years in Yongxing Island in August 2002. It was the first time to present the percent cover data of live corals of Yongxing Island via a systematic scheme of quantitative quadrat sampling. The average total percent cover of the live coral is 68.4%. We found the total percent cover (TPC) could be an idea proxy of the overall disturbance regime impinging on the coral community. TPC can be used as an integrated measure of disturbance to coral reef. Using this surrogate, the best fit relation between TPC of the live corals and species richness, species diversity, or the species evenness is the unimodal second-order polymorphic parabola equation. This result is expected by the intermediate disturbance hypothesis (IDH). From the fitted parabola equations, we calculated the best TPCs corresponding to the maximal species richness, the highest species diversity, or the most evenness respectively. They are surprisingly stabilized between 55% and 58%. Coralcover is the basic parameter widely available for most observation, research or monitoring programs. The good qualitative characteristics of percent cover provide powerful tool for the experimental, theoretical and modeling studies of coral reef in response to the disturbance.     

Estimate of carbonate production by scleractinian corals at Luhuitou fringing reef, Sanya, China

Carbonate production by scleractinian corals not only maintains coral reef growth, but also represents an important source of atmospheric carbon dioxide. In this paper the carbonate production by scler-actinian corals at Luhuitou fringing reef, Sanya, Hainan Island, China, is investigated with an ecological census-based method. Averaged carbonate production is 1.16 ± 0.55 kg·m-2·a-1 and 3.52 ± 1.32 kg·m-2·a-1 on the reef flat and reef slope, respectively, depending on the composition and distribution of cor...     

Fluorescent pigments in corals are photoprotective

All reef-forming corals depend on the photosynthesis performed by their algal symbiont, and such corals are therefore restricted to the photic zone. The intensity of light in this zone declines over several orders of magnitude--from high and damaging levels at the surface to extreme shade conditions at the lower limit. The ability of corals to tolerate this range implies effective mechanisms for light acclimation and adaptation. Here we show that the fluorescent pigments (FPs) of corals provide a photobiological system for regulating the light environment of coral host tissue. Previous studies have suggested that under low light, FPs may enhance light availability. We now report that in excessive sunlight FPs are photoprotective; they achieve this by dissipating excess energy at wavelengths of low photosynthetic activity, as well as by reflecting of visible and infrared light by FP-containing chromatophores. We also show that FPs enhance the resistance to mass bleaching of corals during periods of heat stress, which has implications for the effect of environmental stress on the diversity of reef-building corals, such as enhanced survival of a broad range of corals allowing maintenance of habitat diversity.     

珊瑚礁相关病毒的研究进展

珊瑚礁被称为海洋中的“热带雨林”,是海洋生态系统中的核心组成部分之一。珊瑚礁病毒包含珊瑚本身感染的病毒和珊瑚共生生物（如藻类、细菌等）感染的病毒,对于珊瑚礁健康与疾病、生长与死亡起着独特而重要的作用,在维持珊瑚礁生态系统的物质循环和能量流动等生态学功能方面起着潜在作用。由于技术的局限以及关注度较低等原因,有关珊瑚礁病毒的研究较少。近年来,随着高通量测序技术的发展,宏转录组和宏基因组逐渐成为研究珊瑚礁病毒多样性及其生态学功能的重要方法之一。本文对国内外相关研究进行总结和归纳,拟为珊瑚礁病毒后续研究提供新的方向和思路,同时指导我国珊瑚礁的保护以及修复。