Solar Ultraviolet and the Evolutionary History of Cyanobacteria

On the basis of photobiological, evolutionary, paleontological, paleoenvironmental and physiological arguments, a time course for the role of solar ultraviolet radiation (UVR, wavelengths below 400 nm) in the ecology and evolution of cyanobacteria is proposed in which three main periods can be distinguished. An initial stage, before the advent of oxygenic photosynthesis, when high environmental fluxes of UVC (wavelengths below 280 nm) and UVB (280–320 nm) may have depressed the ability of protocyanobacteria to develop large populations or restricted them to UVR refuges. A second stage lasting between 500 and 1500 Ma (million years), started with the appearance of true oxygen-evolving cyanobacteria and the concomitant formation of oxygenated (micro)environments under an oxygen free-atmosphere. In this second stage, the age of UV, the overall importance of UVR must have increased substantially, since the incident fluxes of UVC and UVB remained virtually unchanged, but additionally the UVA portion of the spectrum (320–400 nm) suddenly became biologically injurious and extremely reactive oxygen species must have formed wherever oxygen and UVR spatially coincided. The last period began with the gradual oxygenation of the atmosphere and the formation of the stratospheric ozone shield. The physiological stress due to UVC all but disappeared and the effects of UVB were reduced to a large extent. Evidence in support of this dynamics is drawn from the phylogenetic distribution of biochemical UV-defense mechanisms among cyanobacteria and other microorganisms. The specific physical characteristics of UVR and oxygen exposure in planktonic, sedimentary and terrestrial habitats are used to explore the plausible impact of UVR in each of the periods on the ecological distribution of cyanobacteria.     

海洋酸化对珊瑚礁生态系统的影响研究进展

目前,大气CO2浓度的升高已导致海水pH值比工业革命前下降了约0.1,海水碳酸盐平衡体系随之变化,进而影响珊瑚礁生态系统的健康。近年来的研究表明海洋酸化导致造礁石珊瑚幼体补充和群落恢复更加困难,造礁石珊瑚和其它造礁生物(Reef-building organisms)钙化率降低甚至溶解,乃至影响珊瑚礁鱼类的生命活动。虽然海洋酸化对造礁石珊瑚光合作用的影响不显著,但珊瑚-虫黄藻共生体系会受到一定影响。建议选择典型海区进行长期系统监测,结合室内与原位模拟试验,从个体、种群、群落到系统不同层面,运用生理学和分子生物学技术,结合生态学研究手段,综合研究珊瑚的相应响应,以期深入认识海洋酸化对珊瑚礁生态系统健康(例如珊瑚白化)的影响及其效应。     

Importance of live coral habitat for reef fishes

Live corals are the key habitat forming organisms on coral reefs, contributing to both biological and physical structure. Understanding the importance of corals for reef fishes is, however, restricted to a few key families of fishes, whereas it is likely that a vast number of fish species will be adversely affected by the loss of live corals. This study used data from published literature together with independent field based surveys to quantify the range of reef fish species that use live coral habitats. A total of 320 species from 39 families use live coral habitats, accounting for approximately 8 % of all reef fishes. Many of the fishes reported to use live corals are from the families Pomacentridae (68 spp.) and Gobiidae (44 spp.) and most (66 %) are either planktivores or omnivores. 126 species of fish associate with corals as juveniles, although many of these fishes have no apparent affiliation with coral as adults, suggesting an ontogenetic shift in coral reliance. Collectively, reef fishes have been reported to use at least 93 species of coral, mainly from the genus Acropora and Porities and associate predominantly with branching growth forms. Some fish associate with a single coral species, whilst others can be found on more than 20 different species of coral indicating there is considerable variation in habitat specialisation among coral associated fish species. The large number of fishes that rely on coral highlights that habitat degradation and coral loss will have significant consequences for biodiversity and productivity of reef fish assemblages.     

Upper Permian coral reef and colonial rugose corals in northwest Hunan, South China

Summary The roles of Permian colonial corals in forming organic reefs have not been adequately assessed, although they are common fossils in the Permian strata. It is now known that colonial corals were important contributors to reef framework during the middle and late Permian such as those in South China, northeast Japan, Oman and Thailand. A coral reef occurs in Kanjia-ping, Cili County, Hunan, South China. It is formed by erect and unscathed colonies ofWaagenophyllum growing on top of one anotherin situ to form a baffle and framework. Paleontological data of the Cili coral reef indicates a middle to late Changhsing age (Late Permian), corresponding to thePalaeofusulina zone. The coral reef exposure extends along the inner platform margin striking in E-S direction for nearly 4 km laterally and generally 35 to 57 m thick. The Cili coral reef exhibits a lateral differentiation into three main reef facies; reef core facies, fore-reef facies, and marginal slope facies. The major reef-core facies is well exposed in Shenxian-wan and Guanyin-an sections where it rests on the marginal slope facies. Colonial corals are dispersed and preserved in non-living position easward. Sponges become major stabilizing organisms in the eastern part of Changhsing limestone outcrop in Kanjia-ping, but no read sponge reefs were formed. Coral reefs at Cili County in Human are different distinctly from calcisponge reefs in South China in their palaeogeography, lithofacies development, organic constitutuents, palaeoecology and diagenesis. The Cili coral reef also shows differences in age, depositional facies association, reef organisms and diagenesis from coral reefs in South Kitakami of Japan, Khorat Plateau of Thailand, and Saih Hatat of Oman. Although some sponge reefs and mounds can reach up to the unconformable Permian/Triassic boundary, coral reef at Kanjia-ping, Cili County, is the latest Permian reef known. This reef appears to had been formed in a palaeoenvironment that is different from that of the sponge reefs and provides an example of new and unique Permian reef type in South China, and could help us to: 1) understand the significance of colonial corals in Permian carbonate buildups; 2) evaluate the importance of coral community evolution prior to the collapse of reef ecosystems at the Permian/Triassic boundary; 3) better understand the effects of the biotic extinction events in Palaeotethys realm; 4) look for environmental factors that may have controlled reefs through time and space, and 5) provide valuable data for the study of Permian palaeoclimate and global evolutionary changes of Permian reefs and reef community.     

Shifting paradigms in restoration of the world's coral reefs

Many ecosystems around the world are rapidly deteriorating due to both local and global pressures, and perhaps none so precipitously as coral reefs. Management of coral reefs through maintenance (e.g., marine‐protected areas, catchment management to improve water quality), restoration, as well as global and national governmental agreements to reduce greenhouse gas emissions (e.g., the 2015 Paris Agreement) is critical for the persistence of coral reefs. Despite these initiatives, the health and abundance of corals reefs are rapidly declining and other solutions will soon be required. We have recently discussed options for using assisted evolution (i.e., selective breeding, assisted gene flow, conditioning or epigenetic programming, and the manipulation of the coral microbiome) as a means to enhance environmental stress tolerance of corals and the success of coral reef restoration efforts. The 2014–2016 global coral bleaching event has sharpened the focus on such interventionist approaches. We highlight the necessity for consideration of alternative (e.g., hybrid) ecosystem states, discuss traits of resilient corals and coral reef ecosystems, and propose a decision tree for incorporating assisted evolution into restoration initiatives to enhance climate resilience of coral reefs.     

Indicators of UV Exposure in Corals and Their Relevance to Global Climate Change and Coral Bleaching

A compelling aspect of the deterioration of coral reefs is the phenomenon of coral bleaching. Through interactions with other factors such as sedimentation, pollution, and bacterial infection, bleaching can impact large areas of a reef with limited recovery, and it might be induced by a variety of stressors including temperature and salinity extremes, and ultraviolet light. Under conditions of ocean warming, often associated with calm and stratified waters, photobleaching of UV-absorb-ing chromophoric dissolved organic matter (CDOM) is increased, and penetration of both UV-B and UV-A is greatly enhanced. Indices of UV-specific effects in coral tissue are needed to test whether UV increases, associated with global climate change, are harmful to corals. To address this challenge, we have evaluated UV-specific effects in corals and have characterized factors that alter penetration of UV radiation over coral reefs. An immunoblotting assay was developed to examine UV-specific lesions (thymine dimers) in coral and zooxanthellae DNA. We observed dose-dependent increases of thymine dimers in coral (Porites porites var porites) exposed to artificial solar irradiance in a solar simulator, although effects were not strictly proportional. UV measurements were made in July 1999 at Eastern Sambo reef and nearby sites, including profiling along transects from reef to shore. Results of these analyses indicate that the coral at Eastern Sambo reef (at 3-4 meters) were receiving UV-B radiation that was equivalent to 25 to 30% of surface UV irradiance. However, the water just inside the reef in Hawk Channel (located closer to land) was considerably more opaque to UV. This water photobleached with loss of UV absorbance and fluorescence when it was exposed to simulated solar radiation. These results indicate that photobleaching of the DOM and transport of near-shore water out over the reefs might play a key role in controlling UV penetration to the reef surface.     

Exposure to ultraviolet radiation (290-400 nm) causes oxidative stress, DNA damage, and expression of p53/p73 in laboratory experiments on embryos of the spotted salamander, Ambystoma maculatum

Developing embryos of the spotted salamander, Ambystoma maculatum, exposed to ultraviolet radiation (UVR; 290-400 nm) in the laboratory show a significant sensitivity to UVB (290-320 nm) radiation. Embryos in laboratory experiments exhibited significant DNA damage during exposures to UVR despite a significant increase in the production of the protective pigment melanin in response to UVR exposure. DNA damage occurs as a result of both the direct effects of exposure to UVR, and the indirect effects are mediated by the production of reduced oxygen intermediates. The production of reactive oxygen species initiates the expression of p53/p73 that leads to either DNA repair or apoptosis. When similar experiments are conducted on salamander embryos exposed to solar UVR in vernal pools, the embryos show significantly less sensitivity and higher survivorship. The differences between laboratory and field experiments are a result of the attenuation of UVR caused by the accumulation of dissolved organic carbon within the pools of these wooded areas. These findings suggest that northeastern populations of spotted salamanders are sensitive to UVR but are not significantly affected by present-day irradiances of UVR in the field. These results do suggest that continued decreases in stratospheric ozone over temperate latitudes have the potential to affect spotted salamanders in their natural habitats.     

Will increased storm disturbance affect the biodiversity of intertidal,nonscleractinian sessile fauna on coral reefs?

Relatively little is known about how the future effects of climatic change, including increases in sea level, temperature and storm severity and frequency, will impact on patterns of biodiversity on coral reefs, with the notable exception of recent work on corals and fish in tropical reef ecosystems. Sessile invertebrates such as ascidians, sponges and bryozoans occupying intertidal rubble habitats on coral reefs contribute significantly to the overall biodiversity and ecosystem function, but there is little or no information available on the likely impacts on these species from climate change. The existing strong physical gradients in these intertidal habitats will be exacerbated under predicted climatic change. By examining the distribution and abundance of nonscleractinian, sessile invertebrate assemblages exposed to different levels of wave action and at different heights on the shore around a coral reef, we show that coral reef intertidal biodiversity is particularly sensitive to physical disturbance. As physical disturbance regimes increase due to more intense storms and wave action associated with global warming, we can expect to see a corresponding decrease in the diversity of these cryptic sessile assemblages. This could impact negatively on the future health and productivity of coral reef ecosystems, given the ecosystem services these organisms provide.     

Comments on coral reef regeneration, bioerosion, biogeography, and chemical ecology: future directions

This paper represents a brief review of three processes operating on coral reefs and the results of studies of a fourth process, and how those results may be applied to the benefit of humankind. The areas are reef regeneration processes; bioerosion; dispersal, recruitment and biogeography of corals; and chemical ecology and natural products chemistry of reef organisms. Possible future directions for research will also be considered in each area. Regarding reef regeneration processes, coral reefs are degrading rapidly on a global scale due to over-fishing, fishing techniques causing habitat destruction, deforestation, mass mortalities of key reef species, nutrient enrichment and sedimentation. Seeding of reefs with the larvae of corals and other key reef organisms, such as echinoids, may help to promote and enhance reef regeneration in the future. Such techniques will be made possible by studies of the embryology, larval ecology, dispersal and recruitment processes, and related local physical oceanographic processes. Regarding bioerosion, both internal and external bioerosion are affected by grazers and predators. Bioerosion is also affected by nutrient enrichment, as shown through correlative studies (Great Barrier Reef) and studies of opportunity (Kaneohe Bay). Ongoing experiments such as ENCORE will help to answer questions about the role of dissolved nutrients in enhancing internal bioerosion. Questions still remain, however, regarding the role of particulates in promoting internal bioerosion and the resultant weakening of and negative growth in the reef framework. Regarding dispersal, recruitment and the biogeography of corals, it is now known that most species of coral reproduce via broadcasting, although there appear to be proportionally more brooders in the Caribbean than in the western Pacific. Differential extinctions in the western Pacific vs. the western Atlantic have contributed to the biogeographic distribution of corals we observe today and the concentric isoclines of species diversity in numerous reef organisms in the western Pacific. The role of reproductive mode in contributing to these patterns is, however, still not understood. Investigations into the roles of different larval longevities and reproductive modes may help us answer questions regarding their differential distribution and the potential effects of major perturbations such as global warming on future distributions. With respect to the chemical ecology of alcyonacean octocorals (soft corals), four functions have been determined thus far for secondary metabolites in this group, anti-predation, anti-competition (allelopathy), anti-fouling, and enhancement of reproductive success. Investigations of alcyonacean reproduction has revealed that it may be necessary for several secondary metabolites to be present simultaneously before a function may be realized or fully manifested. This raises questions regarding the manner in which novel compounds are tested by medical laboratories for bioactivity using a single compound. Simultaneously testing of multiple compounds derived from a single organism may be necessary in the future to reveal potential valuable synergistic bioactivity. Also, some novel secondary metabolites may have other valuable commercial applications, as is the case with the UV-absorbing compounds of corals and other reef organisms found on the Great Barrier Reef. In order to avoid overlooking medically or commercially valuable functions of these compounds, broader testing may be necessary.     

Ultraviolet radiation and consumer effects on a field-grown intertidal macroalgal assemblage in Antarctica

Ultraviolet radiation (UVR) research on marine macroalgae has hithero focussed on physiological effects at the organism level, while little is known on the impact of UV radiation on macroalgal assemblages and even less on interactive effects with other community drivers, e.g. consumers. Field experiments on macrobenthos are scarce, particularly in the Antarctic region. Therefore, the effects of UVR and consumers (mainly limpets were excluded) on early successional stages of a hard bottom macroalgal community on King George Island, Antarctica, were studied. In a two‐factorial design experimental units [(1) ambient radiation, 280–700 nm; (2) ambient minus UVB, 320–700 nm and (3) ambient minus UVR, 400–700 nm vs. consumer–no consumer] were installed between November 2004 and March 2005 (n= 4 plus controls). Dry mass, species richness, diversity and composition of macroalgal assemblages developing on ceramic tiles were followed. Consumers significantly suppressed green algal recruits and total algal biomass but increased macroalgal richness and diversity. Both UVA and UVB radiation negatively affected macroalgal succession. UVR decreased the density of Monostroma hariotii germlings in the first 10 weeks of the experiment, whereas the density of red algal recruits was significantly depressed by UVR at the end of the study. After 106 days macroalgal diversity was significantly higher in UV depleted than in UV‐exposed assemblages. Furthermore, species richness was significantly lower in the UV treatments and species composition differed significantly between the UV‐depleted and the UV‐exposed treatment. Marine macroalgae are very important primary producers in coastal ecosystems, serving as food for herbivores and as habitat for many organisms. Both, UVR and consumers significantly shape macroalgal succession in the Antarctic intertidal. Consumers, particularly limpets can mediate negative effects of ambient UVR on richness and diversity till a certain level. UVB radiation in general and an increase of this short wavelength due to stratospheric ozone depletion in particular may have the potential to affect the zonation, composition and diversity of Antarctic intertidal seaweeds altering trophic interactions in this system.     

Status and progress in coral reef disease research

Recent findings on the ecology, etiology and pathology of coral pathogens, host resistance mechanisms, previously unknown disease/syndromes and the global nature of coral reef diseases have increased our concern about the health and future of coral reef communities. Much of what has been discovered in the past 4 years is presented in this special issue. Among the significant findings, the role that various Vibrio species play in coral disease and health, the composition of the 'normal microbiota' of corals, and the possible role of viruses in the disease process are important additions to our knowledge. New information concerning disease resistance and vectors, variation in pathogen composition for both fungal diseases of gorgonians and black band disease across oceans, environmental effects on disease susceptibility and resistance, and temporal and spatial disease variations among different coral species is presented in a number of papers. While the Caribbean may still be the 'disease hot spot' for coral reefs, it is now clear that diseases of coral reef organisms have become a global threat to coral reefs and a major cause of reef deterioration.     

大型海藻在珊瑚礁退化过程中的作用

随着全球范围珊瑚礁的退化,大型海藻在珊瑚礁区的覆盖度呈增多的趋势。大型海藻的大量生长,妨碍了珊瑚的生长、繁殖、恢复等过程。概括起来,大型海藻对珊瑚生长、繁殖及恢复过程所产生的不利影响主要包括:(1)大型海藻通过与珊瑚竞争空间和光照而影响珊瑚生长;(2)大型海藻与珊瑚直接接触时,通过摩擦作用及释放化感物质而影响珊瑚生长;(3)大型海藻的大量生长打破了珊瑚与海藻的竞争平衡,珊瑚为应对大型海藻的入侵而把用于生长和繁殖的能量转移到组织修复与防御上,进而造成珊瑚繁殖能量的减少;(4)大型海藻通过影响珊瑚幼虫的附着及附着后的存活率,而阻碍珊瑚群落的发展;(5)海藻还能通过富集沉积物、释放病原体及扰乱珊瑚共生微生物的生长等而间接影响珊瑚生长。明确的竞争机制有利于研究海藻与珊瑚的相互作用过程。在总结前人对海藻与珊瑚的竞争机制研究的基础上,把两者的竞争机制划分成物理机制、化学机制、微生物机制三大类,物理机制是研究得比较透彻的竞争机制,而化学机制与微生物机制则需要更深入的研究,是当前研究的热点。目前,我国对珊瑚礁中底栖海藻与珊瑚的相互作用研究甚少;鉴于此,对底栖海藻功能群的划分类型以及三大类型底栖海藻对珊瑚的作用特点做了简要介绍,并对珊瑚礁退化的现状和退化珊瑚礁区内海藻的表现做了概述。在此基础上,再综述国外关于大型海藻对珊瑚的影响研究进展,指出我国应该加强对南海珊瑚礁区大型海藻的种类分布及丰富度等的调查,评价大型海藻对南海珊瑚礁的影响现状;并结合生理学、分子生物学技术和生态学研究手段,在细胞与分子水平上探索海藻对珊瑚的影响机制,以期为珊瑚礁生态系统的保护提供参考。     

Coral Skeletons Defend against Ultraviolet Radiation

Background

Many coral reef organisms are photosynthetic or have evolved in tight symbiosis with photosynthetic symbionts. As such, the tissues of reef organisms are often exposed to intense solar radiation in clear tropical waters and have adapted to trap and harness photosynthetically active radiation (PAR). High levels of ultraviolet radiation (UVR) associated with sunlight, however, represent a potential problem in terms of tissue damage.

Methodology/Principal Findings

By measuring UVR and PAR reflectance from intact and ground bare coral skeletons we show that the property of calcium carbonate skeletons to absorb downwelling UVR to a significant extent, while reflecting PAR back to the overlying tissue, has biological advantages. We placed cnidarians on top of bare skeletons and a UVR reflective substrate and showed that under ambient UVR levels, UVR transmitted through the tissues of cnidarians placed on top of bare skeletons were four times lower compared to their counterparts placed on a UVR reflective white substrate. In accordance with the lower levels of UVR measured in cnidarians on top of coral skeletons, a similar drop in UVR damage to their DNA was detected. The skeletons emitted absorbed UVR as yellow fluorescence, which allows for safe dissipation of the otherwise harmful radiation.

Conclusions/Significance

Our study presents a novel defensive role for coral skeletons and reveals that the strong UVR absorbance by the skeleton can contribute to the ability of corals, and potentially other calcifiers, to thrive under UVR levels that are detrimental to most marine life.     

珊瑚及共生藻在白化过程中的适应机制研究进展

珊瑚礁生态系统具有非常重要的生态学功能。但是随着全球气候变暖和CO2浓度的升高,珊瑚白化事件越来越频繁,珊瑚礁生态系统面临严重的危机。影响珊瑚白化的重要因子主要有海水温度的异常（过高或过低）,太阳辐射与紫外线辐射,海水盐度的偏离,珊瑚疾病,海洋污染,长棘海星的爆发,人类的过度捕鱼和全球CO2浓度升高等。其中,海洋表面水体温度（SST）的异常升高为珊瑚白化的主要因素。珊瑚主要是通过珊瑚与共生藻的生理适应机制以及更换共生藻基因型机制两种方式来适应环境胁迫的。生理适应机制主要通过叶黄素循环、珊瑚色素荧光（热）、活性氧清除系统（自由基）、分泌紫外线吸收物质MAAs（紫外光）、产生热休克蛋白HspS（热）来实现的。珊瑚共生藻基因型更换适应机制是指珊瑚的适应性白化假说。珊瑚的适应性白化假说还有很多争议,还需要更多的实验证据提供支持。未来的研究重点将在珊瑚白化过程中共生藻-珊瑚共生功能体作为整体性的研究,尤其是珊瑚宿主在白化过程中对共生功能体作出贡献的研究。     

Experimental biology of coral reef ecosystems

Coral reef ecosystems are at the crossroads. While significant gaps still exist in our understanding of how “normal” reefs work, unprecedented changes in coral reef systems have forced the research community to change its focus from basic research to understand how one of the most diverse ecosystems in the world works to basic research with strong applied implications to alleviate damage, save, or restore coral reef ecosystems. A wide range of stressors on local, regional, and global spatial scales including over fishing, diseases, large-scale disturbance events, global climate change (e.g., ozone depletion, global warming), and over population have all contributed to declines in coral cover or phase shifts in community structure on time scales never observed before. Many of these changes are directly or indirectly related to anthropogenically induced changes in the global support network that affects all ecosystems. This review focuses on some recent advances in the experimental biology of coral reef ecosystems, and in particular scleractinian corals, at all levels of biological organization. Many of the areas of interest and techniques discussed reflect a progression of technological advances in biology and ecology but have found unique and timely application in the field of experimental coral reef biology. The review, by nature, will not be exhaustive and reflects the author's interests to a large degree. Because of the voluminous literature available, an attempt has been made to capture the essential elements and references for each topic discussed.     

Effects of ultraviolet radiation and water motion on the reef coral, Porites compressa Dana: a transplantation experiment

Scleractinian corals have adapted to live in habitats were the level of ultraviolet radiation (UVR, 280–400 nm) is extremely high. The putative photoprotective molecules called mycosporine-like amino acids (MAAs) contained in the corals' tissues absorb UVR and release it harmlessly as heat. MAA concentration in corals is quite plastic and correlates well with UVR dose, but other ecological factors such as water motion may influence MAA production as well. In this study, the effects of ambient UVR and water motion on MAA concentration and several physiological parameters of the reef coral Porites compressa Dana were investigated in a two by two factorial transplantation experiment. Replicate branches from nine morphologically distinct colonies were transplanted from the windward side of Coconut Island (Kaneohe Bay, HI) to a control area on the windward side (ambient water motion) and to an area on the leeward side (low water motion). The transplanted corals were placed under UV-opaque (UVO) or UV-transparent (UVT) filters fixed to the reef. Initially and at 3 and 6 weeks, coral branches were weighed to determine calcification rate and tissues were extracted in methanol for photosynthetic pigment and MAA analysis via high performance liquid chromatography (HPLC). UVR was a significant factor determining MAA concentration. When UVR was screened from the corals' environment, total MAA concentration decreased by 33% over 6 weeks. However, UVR-exposed corals moved to low water motion also decreased MAA levels, while UVR-exposed corals moved to the control area retained initial levels. Photosynthetic pigments and calcification rate were also significantly reduced in corals moved to low water motion. There was no UVR effect on photosynthetic pigments or calcification rate. This study provides evidence that water motion is important for the maintenance of MAAs. However, there were interesting colony-specific patterns in MAA composition and response to the UVR treatment; some colonies had high total concentrations of MAAs in all treatments, while others displayed a pronounced UVR effect. Also, each genotype seemed to have its own signature MAA composition. These findings indicate a genetic (host, zooxanthellae or both) component to UVR resistance in this population of P. compressa.     

Reef coral diversity and global change

Regional anthropogenic processes such as pollution, dredging, and overfishing on coral reefs currently threaten the biodiversity of stony corals and other reef-associated organisms. Global climate change may interact with anthropogenic processes to create additional impacts on coral diversity in the near future. In order to predict these changes, it is necessary to understand the magnitude and causes of variation in scleractinian coral diversity throughout their 240 million year history. The fossil record documents long periods of speciation in corals, interrupted repeatedly by events of mass extinction. Some of these events relate clearly to changes in global climate. Diversity in reef corals has increased since their last period of extinction at the end of the Cretaceous (65 My bp ), and is still rising. During the last 8 million years, the fragmentation of the once pantropical Tethys Sea separated corals into two major biogeographical provinces. Periods of glaciation also have caused major changes in sea level and temperature. Accumulated evidence supports the theory that relative habitat area and changing patterns of oceanic circulation are mainly responsible for the two observed centres of recent coral diversity at the western tropical margins of the Atlantic and Pacific oceans. At predicted rates of climate change in the near future, coral reefs are likely to survive as an ecosystem. Increases in sea level may actually benefit corals and lead to regional increases in diversity if new habitat area on back reefs is opened to increased water circulation and thus coral dispersal. Rising temperature may cause higher rates of coral mortality and even local extinction in isolated, small populations such as those on oceanic islands. The effects of increases in ultraviolet radiation (UV) are largely unknown, but likely to be negative. UV may damage planktonic coral propagules in oceanic surface waters and thus decrease rates of gene flow between coral populations. This may result in increased local extinctions, again with the strongest impact on widely separated reefs with small coral populations. The largest threats to coral diversity are regional anthropogenic impacts, which may interact with global climate change to exacerbate rates of local species extinctions. Centres of high reef coral diversity coincide with human population centres in south-east Asia and the Caribbean, and thus the greatest potential for species loss lies in these geographical areas.     

On the potential for ocean acidification to be a general cause of ancient reef crises

Anthropogenic rise in the carbon dioxide concentration in the atmosphere leads to global warming and acidification of the oceans. Ocean acidification (OA) is harmful to many organisms but especially to those that build massive skeletons of calcium carbonate, such as reef corals. Here, we test the recent suggestion that OA leads not only to declining calcification of reef corals and reduced growth rates of reefs but may also have been a trigger of ancient reef crises and mass extinctions in the sea. We analyse the fossil record of biogenic reefs and marine organisms to (1) assess the timing and intensity of ancient reef crises, (2) check which reef crises were concurrent with inferred pulses of carbon dioxide concentrations and (3) evaluate the correlation between reef crises and mass extinctions and their selectivity in terms of inferred physiological buffering. We conclude that four of five global metazoan reef crises in the last 500 Myr were probably at least partially governed by OA and rapid global warming. However, only two of the big five mass extinctions show geological evidence of OA.     

Coral Community Adaptability to Environmental Change at the Scales of Regions, Reefs and Reef Zones

SYNOPSIS. Projected global increases in temperature, sea level,storminess and atmospheric carbon dioxide (CO₂) are likely tocause changes in reef coral communities which the present humangeneration will view as deleterious. It is likely coral communitytrajectories will be influenced as much by the reduction inintervals between extreme events as the projected increasesin means of environmental parameters such as temperature, atmosphericCO₂ and sea-level. Depressed calcification rates in corals causedby reduced aragonite saturation state of water may increasevulnerability of corals to storms. Moreover, reduction in intervalsbetween storms and other extreme events causing mass mortalityin corals (coral predators, diseases, bleaching) are likelyto more frequently "set back" reef coral communities to earlysuccessional stages or alternate states characterized by non-calcifyingbenthos (plants, soft corals, sponges). The greater the areaand the longer the duration of dominance of putative "coral/corallinealgae" zones of coral reefs by non-calcifying stages, the lesswill be the reefs capacity to accrete limestone bulk lockedup in the big skeletal units of late successional stages (i.e.,very large old corals). Averaged over decades to centuries,the effects of such changes on the coral community's carryingcapacity for other biota such as fish are unpredictable. A "shiftingsteady-state mosaic" null model may provide a useful conceptualtool for defining a baseline and tracking changes from thatbaseline through time.     

Global assessment of coral bleaching and required rates of adaptation under climate change

Elevated ocean temperatures can cause coral bleaching, the loss of colour from reef‐building corals because of a breakdown of the symbiosis with the dinoflagellate Symbiodinium. Recent studies have warned that global climate change could increase the frequency of coral bleaching and threaten the long‐term viability of coral reefs. These assertions are based on projecting the coarse output from atmosphere–ocean general circulation models (GCMs) to the local conditions around representative coral reefs. Here, we conduct the first comprehensive global assessment of coral bleaching under climate change by adapting the NOAA Coral Reef Watch bleaching prediction method to the output of a low‐ and high‐climate sensitivity GCM. First, we develop and test algorithms for predicting mass coral bleaching with GCM‐resolution sea surface temperatures for thousands of coral reefs, using a global coral reef map and 1985–2002 bleaching prediction data. We then use the algorithms to determine the frequency of coral bleaching and required thermal adaptation by corals and their endosymbionts under two different emissions scenarios. The results indicate that bleaching could become an annual or biannual event for the vast majority of the world's coral reefs in the next 30–50 years without an increase in thermal tolerance of 0.2–1.0°C per decade. The geographic variability in required thermal adaptation found in each model and emissions scenario suggests that coral reefs in some regions, like Micronesia and western Polynesia, may be particularly vulnerable to climate change. Advances in modelling and monitoring will refine the forecast for individual reefs, but this assessment concludes that the global prognosis is unlikely to change without an accelerated effort to stabilize atmospheric greenhouse gas concentrations.