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。     

Coral bleaching: thermal adaptation in reef coral symbionts

Many corals bleach as a result of increased seawater temperature, which causes them to lose their vital symbiotic algae (Symbiodinium spp.) - unless these symbioses are able to adapt to global warming, bleaching threatens coral reefs worldwide. Here I show that some corals have adapted to higher temperatures, at least in part, by hosting specifically adapted Symbiodinium. If other coral species can host these or similar Symbiodinium taxa, they might adapt to warmer habitats relatively easily.     

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.     

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.     

Thresholds and the resilience of Caribbean coral reefs

The deteriorating health of the world's coral reefs threatens global biodiversity, ecosystem function, and the livelihoods of millions of people living in tropical coastal regions. Reefs in the Caribbean are among the most heavily affected, having experienced mass disease-induced mortality of the herbivorous urchin Diadema antillarum in 1983 and two framework-building species of coral. Declining reef health is characterized by increases in macroalgae. A critical question is whether the observed macroalgal bloom on Caribbean reefs is easily reversible. To answer this question, we must resolve whether algal-dominated reefs are an alternative stable state of the ecosystem or simply the readily reversible result of a phase change along a gradient of some environmental or ecological parameter. Here, using a fully parameterized simulation model in combination with a simple analytical model, we show that Caribbean reefs became susceptible to alternative stable states once the urchin mortality event of 1983 confined the majority of grazing to parrotfishes. We reveal dramatic hysteresis in a natural system and define critical thresholds of grazing and coral cover beyond which resilience is lost. Most grazing thresholds lie near the upper level observed for parrotfishes in nature, suggesting that reefs are highly sensitive to parrotfish exploitation. Ecosystem thresholds can be combined with stochastic models of disturbance to identify targets for the restoration of ecosystem processes. We illustrate this principle by estimating the relationship between current reef state (coral cover and grazing) and the probability that the reef will withstand moderate hurricane intensity for two decades without becoming entrained in a shift towards a stable macroalgal-dominated state. Such targets may help reef managers face the challenge of addressing global disturbance at local scales.     

世界珊瑚礁现状和威胁研究进展

鉴于珊瑚礁巨大的价值和面临全球气候变化严峻的威胁,珊瑚礁退化和发展前景一直是珊瑚礁研究的最中心议题。人类活动和气候变化已经导致珊瑚礁全球性衰退损失40%~50%。1998年全球珊瑚礁严重白化和2015—2017年延时最长全球珊瑚礁白化事件,一再证实全球变暖已经成为珊瑚礁可持续发展的主要威胁,现在扭转导致海洋水温升高的全球变暖的趋势是保护珊瑚礁的唯一希望。由科学家和慈善机构发起的2011年XL Catlin海景调查和2017年50礁倡议成为珊瑚礁管理保护的创新举措。但仍然需要更多的人认识珊瑚礁,热爱珊瑚礁,研究珊瑚礁,保护珊瑚礁。     

Confronting the coral reef crisis

The worldwide decline of coral reefs calls for an urgent reassessment of current management practices. Confronting large-scale crises requires a major scaling-up of management efforts based on an improved understanding of the ecological processes that underlie reef resilience. Managing for improved resilience, incorporating the role of human activity in shaping ecosystems, provides a basis for coping with uncertainty, future changes and ecological surprises. Here we review the ecological roles of critical functional groups (for both corals and reef fishes) that are fundamental to understanding resilience and avoiding phase shifts from coral dominance to less desirable, degraded ecosystems. We identify striking biogeographic differences in the species richness and composition of functional groups, which highlight the vulnerability of Caribbean reef ecosystems. These findings have profound implications for restoration of degraded reefs, management of fisheries, and the focus on marine protected areas and biodiversity hotspots as priorities for conservation.     

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.     

Permanent El Niño during the Pliocene warm period not supported by coral evidence

The El Ni?o/Southern Oscillation (ENSO) system during the Pliocene warm period (PWP; 3-5 million years ago) may have existed in a permanent El Ni?o state with a sharply reduced zonal sea surface temperature (SST) gradient in the equatorial Pacific Ocean. This suggests that during the PWP, when global mean temperatures and atmospheric carbon dioxide concentrations were similar to those projected for near-term climate change, ENSO variability--and related global climate teleconnections-could have been radically different from that today. Yet, owing to a lack of observational evidence on seasonal and interannual SST variability from crucial low-latitude sites, this fundamental climate characteristic of the PWP remains controversial. Here we show that permanent El Ni?o conditions did not exist during the PWP. Our spectral analysis of the δ(18)O SST and salinity proxy, extracted from two 35-year, monthly resolved PWP Porites corals in the Philippines, reveals variability that is similar to present ENSO variation. Although our fossil corals cannot be directly compared with modern ENSO records, two lines of evidence suggest that Philippine corals are appropriate ENSO proxies. First, δ(18)O anomalies from a nearby live Porites coral are correlated with modern records of ENSO variability. Second, negative-δ(18)O events in the fossil corals closely resemble the decreases in δ(18)O seen in the live coral during El Ni?o events. Prior research advocating a permanent El Ni?o state may have been limited by the coarse resolution of many SST proxies, whereas our coral-based analysis identifies climate variability at the temporal scale required to resolve ENSO structure firmly.     

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。     

Interspecies and spatial diversity in the symbiotic zooxanthellae density in corals from northern South China Sea and its relationship to coral reef bleaching

Coral reef bleaching is usually characterized by expulsion of symbiotic zooxanthellae, loss of zooxanthellae pigmentation, or both. We collected 128 samples comprising 39 species of 21 genera of reef-building corals from Luhuitou and Xiaodonghai in Sanya of Hainan Island and Daya Bay of Guangdong Province, respectively, and analyzed the symbiotic zooxanthellae population density. The results show that： （1） the symbiotic zooxanthella density varies from 0.67×10^6 to 8.48×10^6 cell/cm^2, displaying significant interspecies variability, with branch corals usually having relatively less zooxanthellae （ranging from 0.67×10^6 to 2.47×10^6 cell/cm^2） than massive species （from 1.0×10^6 to 8.48×10^6 cell/cm^2）; （2） corals inhabiting within 4 m water depth have higher levels of symbiotic zooxanthellae than those living at the bottom （-7 m depth） of the reef area; （3） there is no discernable difference in the zooxanthellae density between corals from relatively high latitude Daya Bay （-22°N） and those from relatively low latitude Sanya （-18°N） at comparable sea surface temperatures （SST）; （4） in partially-bleached corals, the density of zooxanthellae shows the following order： healthy-looking part〉 semi-bleached part 〉 bleached part. Based on the above results, we suggest that （1） the zooxanthellae density difference between branching and massive coral species is the main cause that branching corals are more vulnerable to bleaching than massive corals. For example, symbiotic zooxanthellae levels are low in branching Acropora and Pocillopora corals and thus these corals are more susceptible to bleaching and mortality; （2） symbiotic zooxanthellae density can also be affected by environmental conditions, such as sediment loads, diving-related turbidity, and aquaculture-related nitrate and phosphate input, and their increase may reduce symbiotic zooxanthellae density in corals.     

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。     

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.     

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.     

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.     

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。     

砗磲人工繁育、资源恢复与南海岛礁生态牧场建设

南海珊瑚岛礁是海洋生态系统的重要组成部分,是区域特色海洋和渔业生物资源及其多样性的保障。砗磲是珊瑚岛礁的构成物种,具有优良的造礁、固礁护礁功能,并为其它岛礁生物提供栖息地、繁育和庇护场所；近二十多年来,在人类活动和全球环境变化影响下,砗磲及珊瑚等岛礁生物资源受到比较严重的破坏,其岛礁生态牧场天然参与者和构建者的功能受损明显。我们突破砗磲幼虫虫黄藻植入难点和变态率极低的瓶颈问题,率先在海南成功研发了砗磲规模化人工繁育和中培技术,并进行了放流增殖试验,取得了良好的效果。但目前和今后砗磲及珊瑚等生物资源情况依然严峻,期望今后开展更系统、科学、更大规模的基于砗磲、珊瑚及其它重要礁栖生物的放流增殖与生态牧场构建,全面恢复南海珊瑚岛礁的生态环境和生物资源,推动南海的可持续发展。     

Mangroves enhance the biomass of coral reef fish communities in the Caribbean

Mangrove forests are one of the world's most threatened tropical ecosystems with global loss exceeding 35% (ref. 1). Juvenile coral reef fish often inhabit mangroves, but the importance of these nurseries to reef fish population dynamics has not been quantified. Indeed, mangroves might be expected to have negligible influence on reef fish communities: juvenile fish can inhabit alternative habitats and fish populations may be regulated by other limiting factors such as larval supply or fishing. Here we show that mangroves are unexpectedly important, serving as an intermediate nursery habitat that may increase the survivorship of young fish. Mangroves in the Caribbean strongly influence the community structure of fish on neighbouring coral reefs. In addition, the biomass of several commercially important species is more than doubled when adult habitat is connected to mangroves. The largest herbivorous fish in the Atlantic, Scarus guacamaia, has a functional dependency on mangroves and has suffered local extinction after mangrove removal. Current rates of mangrove deforestation are likely to have severe deleterious consequences for the ecosystem function, fisheries productivity and resilience of reefs. Conservation efforts should protect connected corridors of mangroves, seagrass beds and coral reefs.     

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.     

石斑鱼类的分子系统学研究进展

石斑鱼类是世界上最重要的海洋经济鱼类之一，其种类繁多，大部分营定居性生活．由于种间缺乏明显的外部形态区别特征．石斑鱼的分类一直是鱼类系统分类学中的一个难题，存在着很多争议和混淆．通过DNA序列分析在分子水平重建物种的系统进化关系是当前系统学研究的一个新热点，是传统系统学研究的重要补充和佐证．本文简要综述了石斑鱼类传统分类研究状况、存在问题及当前石斑鱼类的分子系统学研究进展，并对石斑鱼类的系统学研究发展作了展望．