Coral reef bleaching: facts, hypotheses and implications

Coral reef bleaching, the temporary or permanent loss of photosynthetic microalgae (zooxanthellae) and/or their pigments by a variety of reef taxa, is a stress response usually associated with anthropogenic and natural disturbances. Degrees of bleaching, within and among coral colonies and across reef communities, are highly variable and difficult to quantify, thus complicating comparisons of different bleaching events. Small-scale bleaching events can often be correlated with specific disturbances (e.g. extreme low/high temperatures, low/high solar irradiance, subaerial exposure, sedimentation, freshwater dilution, contaminants, and diseases), whereas large scale (mass) bleaching occurs over 100s to 1000s of km², which is more difficult to explain. Debilitating effects of bleaching include reduced/no skeletal growth and reproductive activity, and a lowered capacity to shed sediments, resist invasion of competing species and diseases. Severe and prolonged bleaching can cause partial to total colony death, resulting in diminished reef growth, the transformation of reef-building communities to alternate, non-reef building community types, bioerosion and ultimately the disappearance of reef structures. Present evidence suggests that the leading factors responsible for large-scale coral reef bleaching are elevated sea temperatures and high solar irradiance (especially ultraviolet wavelengths), which may frequently act jointly.     

Coral reef bleaching in the 1980s and possible connections with global warming

Scleractinian corals and their symbiotic dinoflagellate algae build massive, wave-resistant coral reefs that are pre-eminent in shallow tropical seas. This mutualism is especially sensitive to numerous environmental stresses, and has been disrupted frequently during the past decade. Increased seawater temperatures have been proposed as the most likely cause of coral reef bleaching, and it has been suggested that the recent large-scale disturbances are the first biological indication of global warming. This article describes recent bleaching events and their possible link with sea warming and other environmental stresses, and offers some speculation on the fate of coral reefs if the Earth enters a sustained period of warming.     

Coral bleaching, reef fish community phase shifts and the resilience of coral reefs

The 1998 global coral bleaching event was the largest recorded historical disturbance of coral reefs and resulted in extensive habitat loss. Annual censuses of reef fish community structure over a 12-year period spanning the bleaching event revealed a marked phase shift from a prebleach to postbleach assemblage. Surprisingly, we found that the bleaching event had no detectable effect on the abundance, diversity or species richness of a local cryptobenthic reef fish community. Furthermore, there is no evidence of regeneration even after 5–35 generations of these short-lived species. These results have significant implications for our understanding of the response of coral reef ecosystems to global warming and highlight the importance of selecting appropriate criteria for evaluating reef resilience.     

Coral bleaching: causes and consequences

It has been over 10 years since the phenomenon of extensive coral bleaching was first described. In most cases bleaching has been attributed to elevated temperature, but other instances involving high solar irradiance, and sometimes disease, have also been documented. It is timely, in view of our concern about worldwide reef condition, to review knowledge of physical and biological factors involved in bleaching, the mechanisms of zooxanthellae and pigment loss, and the ecological consequences for coral communities. Here we evaluate recently acquired data on temperature and irradiance-induced bleaching, including long-term data sets which suggest that repeated bleaching events may be the consequence of a steadily rising background sea temperature that will in the future expose corals to an increasingly hostile environment. Cellular mechanisms of bleaching involve a variety of processes that include the degeneration of zooxanthellae in situ, release of zooxanthellae from mesenterial filaments and release of algae within host cells which become detached from the endoderm. Photo-protective defences (particularly carotenoid pigments) in zooxanthellae are likely to play an important role in limiting the bleaching response which is probably elicited by a combination of elevated temperature and irradiance in the field. The ability of corals to respond adaptively to recurrent bleaching episodes is not known, but preliminary evidence suggests that phenotypic responses of both corals and zooxanthellae may be significant.     

Coral reef management and conservation in light of rapidly evolving ecological paradigms

The decline of many coral reef ecosystems in recent decades surprised experienced managers and researchers. It shattered old paradigms that these diverse ecosystems are spatially uniform and temporally stable on the scale of millennia. We now see reefs as heterogeneous, fragile, globally stressed ecosystems structured by strong positive or negative feedback processes. We review the causes and consequences of reef decline and ask whether management practices are addressing the problem at appropriate scales. We conclude that both science and management are currently failing to address the comanagement of extractive activities and ecological processes that drive ecosystems (e.g. productivity and herbivory). Most reef conservation efforts are directed toward reserve implementation, but new approaches are needed to sustain ecosystem function in exploited areas.     

Coral and algal changes after the 1998 coral bleaching: interaction with reef management and herbivores on Kenyan reefs

Interaction between the El Niño and Indian Ocean dipole ocean-atmosphere quasi-periodic oscillations produced one of the warmest seawater temperatures on record in 1998. During the warm northeast monsoon in March and April, Kenya's shallow coral reefs experienced water temperatures between 30 and 31 °C and low winds. This caused large-scale bleaching of hard and soft corals at the end of March, which extended into the cooler months of May and June. Direct observations of coloration in the Mombasa Marine National Park found that the coral genera Acropora, Millepora, Pocillopora, branching Porites and Stylophora showed rapid bleaching and high mortality by the end of May 1998. Other hard coral genera that bleached significantly included Echinopora, Favia, Favites, Galaxea, Hydnophora, Goniopora, Leptoria, Montipora, Platygyra and massive Porites, but mortality was variable among these genera. Astreopora, Coscinarea, Cyphastrea and Pavona were the least responsive genera, with some paling, but little evidence of full bleaching or significant mortality. We compared changes in reef ecology in four national parks (protected from fishing) with four non-park areas (heavy fishing) to determine how coral mortality and herbivory interact under the two management regimes. Benthic studies using line transects in 16 sites spread across ~150 km of coastline were completed before and 6 to 13 months after the bleaching event and found that the cover of nine hard coral genera including Acropora, Alveopora, Favites, Goniopora, Platygyra, Pocillopora, branching Porites, Stylophora and Tubipora decreased significantly (p<0.04) after the event, usually by >85%, and soft coral cover decreased by ~75%. One year after the bleaching, sites in the national parks experienced 88 and 115% increases in turf and fleshy algal cover, respectively, while reefs outside the parks had a 220% increase in fleshy algal cover with no appreciable change in turf-forming algal cover. There was, however, high spatial variation and no statistically significant difference in the change in fleshy algal cover between sites in and out of the national parks. Estimates of herbivory by both fish and sea urchins at each site were a good predictor of the change in fleshy algal cover over the 1-year post-bleaching period in about half of the sites. The other half of the sites did not exhibit large changes in fleshy algal cover, and algal cover at these sites was not clearly influenced by herbivory. The number of coral genera per transect at each site decreased significantly by 31 and 44% in and out of the national parks respectively. Larger-scale search sampling to determine the presence/absence of genera in study sites found consistent losses of coral genera from sites, but produced smaller differences than the line-transect method, suggesting that some genera persisted, but at very low population densities and small colony sizes.     

Coral reef fisheries stock assessment

Summary Although all fisheries are multispecies and spatially heterogeneous, coral reef fisheries are an extreme in both respects. The two main approaches to stock assessment have been either to consider individual species separately or to lump all species together. Both are limited in their predictive power. The lack of ecological knowledge and the large number of parameters required make methods based on single species often impractical or expensive. However, where the appropriate information is available, ecological studies do not provide significant improvements on yield-per-recruit or surplus yield models currently used.Alternative community models based on aggregates of species lack predictive power, empirical support and relative data. Models are further limited because they do not address various economic aspects, fish movement and recruitment, all of which must be spatially resolved. However, the ECOPATH model, based on trophic compartments, represents a new approach useful to multispecies assessment, and the only way at present to include predation in stock models. In practice the data available will be the most important factor in the choice of stock assessment model.The major concern for management of many coral reefs is conservation of the habitat and stocks. In some cases this has been achieved with little reference to stock assessment, by using community management and closed areas, which are receiving increasing support. However, once the objective of conservation is achieved, stock assessment should have an important role in improving the economic performance of the fisheries. The wider problems of management have no simple solutions, but managers should look to adaptive management, designing their own experiments to choose between management models.     

Coral bleaching: the winners and the losers

Sea surface temperatures were warmer throughout 1998 at Sesoko Island, Japan, than in the 10 preceding years. Temperatures peaked at 2.8 °C above average, resulting in extensive coral bleaching and subsequent coral mortality. Using random quadrat surveys, we quantitatively documented the coral community structure one year before and one year after the bleaching event. The 1998 bleaching event reduced coral species richness by 61% and reduced coral cover by 85%. Colony morphology affected bleaching vulnerability and subsequent coral mortality. Finely branched corals were most susceptible, while massive and encrusting colonies survived. Most heavily impacted were the branched Acropora and pocilloporid corals, some of which showed local extinction. We suggest two hypotheses whose synergistic effect may partially explain observed mortality patterns (i.e. preferential survival of thick-tissued species, and shape-dependent differences in colony mass-transfer efficiency). A community-structural shift occurred on Okinawan reefs, resulting in an increase in the relative abundance of massive and encrusting coral species.     

Coral reef primary productivity: A beggar's banquet

Flashy venue, high attendance and much activity: but the food is plain and hard to find, and at the end of the day nobody leaves with much. Primary production in coral reef ecosystems proceeds rapidly through small packages. Maximum areal rates rival the best agricultures, but most of the organic production is conserved and recycled within component organisms and communities. Interactions with animals serve to maximize the productivity of both individual plants and extensive plant assemblages. The diverse aspects of the topic have attracted research for over 40 years. In this, the first of a two-part review, I describe the components of coral reef primary productivity at several levels of organization, and discuss recent approaches to their studs.     

Coral reef fish communities: a compromise view

Synopsis The fishes associated with coral reefs offer excellent opportunities for the study of the factors that determine the species composition of complex, highly interactive communities. Amenable to sampling and direct observations many patch reefs are small enough to be studied as entire units and yet diverse enough to include a wide range of interactions.Coral reef fishes appear to be highly specialized in morphology, color, behavior and life cycles and yet colonization experiments and repeated censusing have shown a surprisingly high variation in the fishes that are associated in similar habitats or in the same habitat at different times. This has led to two different views: (i) the chaos view that holds that the species composition is due to random factors and chance colonization, and (ii) the order view that resource sharing adaptations determine which species can live together.This paper reviews some of the obvious adaptations of reef fishes. An examination of the maximum sizes reached by infaunal reef dwellers shows that the largest individual of each species differs by a constant proportion from the next larger and next smaller species. This suggests growth limitations by interspecies competition. A hypothetical model showing how this might work is offered and it is shown that as long as there are more species than can be accommodated in the community at any one time the number of combinations is great enough to give the appearance of randomness even though the individual species may have precise environmental requirements.This paper forms part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Hellman, ed.).     

Coral growth and reef growth: a brief review

The growth potential of modern zooxanthellate corals from the major reef provinces is reviewed with respect to Holocene reef growth. Both coral growth and reef growth is enhanced globally at the beginning of the Holocene and is maintained regionally in the Caribbean Sea up to the present in contrast to reefs of the Indo-Pacific Ocean. This regional difference is mainly caused by the siphoning effect of the tropical Atlantic, which is characterised still by a rising sea level in contrast to global ocean. Hence, Indo-Pacific reefs exhibit a well-cemented reef crest and reef roof barren of living corals. The evaluation of reef growth rates throughout the Phanerozoic shows reduced growth rates of more than one order of magnitude in comparison to their modern counterparts. This is a result of compaction and diagenesis but also strongly biased by uncertainties in absolute dating. Point counting of individual framebuilders with known growth rate may result in more comparative figures for growth rates of fossil reefs with respect to modern ones.     

Coral gardens: paternity and drug testing on the reef

An international team has used molecular genetics and chemical tagging to trace how baby clownfish travel from their mother's nest through the ocean to the anemone they will live on. More than one out of five juveniles came from nests that were only meters away, despite spending over a week drifting in ocean currents. Such surprising fidelity to a small area of the coral reef bodes well for efforts to preserve coral reef diversity with reserves.     

Coral reef resilience differs among islands within the Gulf of Mannar,southeast India,following successive coral bleaching events

Coral Reefs - We used a 12-yr data set of benthic cover (2005–2017), spanning two bleaching events, to assess changes in benthic cover and coral community composition along 21 islands within...     

Coral reef environmental science: truth versus the Cassandra syndrome

In 1970, coral reef science was warned that the crown-of-thorns starfish, Acanthaster planci, might cause the extinction of scleractinian corals in the Pacific Ocean. Now, 20 years later we can fortunately say that this alarm was almost certainly too severe. Many reefs were devastated by the starfish, but none are extinct, none have disappeared and many are in various stages of recovery. But now in the 1990's a new alarm is being sounded. This time the concern is over widespread destruction of coral reefs by elevated surface temperatures. Once again a few scientists have issued a dire warning that these events may represent a harbinger of ocean warming caused by the Greenhouse Effect. Has not Acanthaster taught coral reef science a lesson? The debate is far from over but this time the mood in general is not one of over-reaction. This time the Cassandras will be tested by the truth of careful experimentation, long-term monitoring and objective interpretation. Coral reef science appears to have come of age.     

Coral bleaching response index: a new tool to standardize and compare susceptibility to thermal bleaching

As coral bleaching events become more frequent and intense, our ability to predict and mitigate future events depends upon our capacity to interpret patterns within previous episodes. Responses to thermal stress vary among coral species; however the diversity of coral assemblages, environmental conditions, assessment protocols, and severity criteria applied in the global effort to document bleaching patterns creates challenges for the development of a systemic metric of taxon‐specific response. Here, we describe and validate a novel framework to standardize bleaching response records and estimate their measurement uncertainties. Taxon‐specific bleaching and mortality records (2036) of 374 coral taxa (during 1982–2006) at 316 sites were standardized to average percent tissue area affected and a taxon‐specific bleaching response index (taxon‐BRI) was calculated by averaging taxon‐specific response over all sites where a taxon was present. Differential bleaching among corals was widely variable (mean taxon‐BRI = 25.06 ± 18.44%, ±SE). Coral response may differ because holobionts are biologically different (intrinsic factors), they were exposed to different environmental conditions (extrinsic factors), or inconsistencies in reporting (measurement uncertainty). We found that both extrinsic and intrinsic factors have comparable influence within a given site and event (60% and 40% of bleaching response variance of all records explained, respectively). However, when responses of individual taxa are averaged across sites to obtain taxon‐BRI, differential response was primarily driven by intrinsic differences among taxa (65% of taxon‐BRI variance explained), not conditions across sites (6% explained), nor measurement uncertainty (29% explained). Thus, taxon‐BRI is a robust metric of intrinsic susceptibility of coral taxa. Taxon‐BRI provides a broadly applicable framework for standardization and error estimation for disparate historical records and collection of novel data, allowing for unprecedented accuracy in parameterization of mechanistic and predictive models and conservation plans.     

珊瑚礁白化研究进展

珊瑚礁白化是由于珊瑚失去体内共生的虫黄藻和（或）共生的虫黄藻失去体内色素而导致五彩缤纷的珊瑚礁变白的生态现象。近年来,频繁发生的珊瑚礁白化导致了珊瑚礁生态系统严重退化,并已经影响到全球珊瑚礁生态系统的平衡,受到了人们的高度重视。研究认为：（1）大范围珊瑚礁白化主要是全球环境变化引起的,尤其是全球变暖和紫外辐射增强;（2）导致珊瑚礁白化的机制主要在于细胞机制和光抑制机制;（3）珊瑚礁白化后的恢复与白化程度有关,大范围白化的珊瑚礁完全恢复需要几年到几十年;（4）珊瑚礁白化的后果在于降低珊瑚繁殖能力、减缓珊瑚礁生长、改变礁栖生物的群落结构,导致大面积珊瑚死亡和改变珊瑚礁生态类型,如变为海藻型等;（5）与珊瑚共生的D系群虫黄藻更适应高温环境,珊瑚礁有可能通过D系群逐渐取代C系群的方式适应全球环境变化。