Coral reef bleaching: ecological perspectives

Coral reef bleaching, the whitening of diverse invertebrate taxa, results from the loss of symbiotic zooxanthellae and/or a reduction in photosynthetic pigment concentrations in zooxanthellae residing within the gastrodermal tissues of host animals. Of particular concern are the consequences of bleaching of large numbers of reef-building scleractinian corals and hydrocorals. Published records of coral reef bleaching events from 1870 to the present suggest that the frequency (60 major events from 1979 to 1990), scale (co-occurrence in many coral reef regions and often over the bathymetric depth range of corals) and severity (>95% mortality in some areas) of recent bleaching disturbances are unprecedented in the scientific literature. The causes of small scale, isolated bleaching events can often be explained by particular stressors (e.g., temperature, salinity, light, sedimentation, aerial exposure and pollutants), but attempts to explain large scale bleaching events in terms of possible global change (e.g., greenhouse warming, increased UV radiation flux, deteriorating ecosystem health, or some combination of the above) have not been convincing. Attempts to relate the severity and extent of large scale coral reef bleaching events to particular causes have been hampered by a lack of (a) standardized methods to assess bleaching and (b) continuous, long-term data bases of environmental conditions over the periods of interest. An effort must be made to understand the impact of bleaching on the remainder of the reef community and the long-term effects on competition, predation, symbioses, bioerosion and substrate condition, all factors that can influence coral recruitment and reef recovery. If projected rates of sea warming are realized by mid to late AD 2000, i.e. a 2°C increase in high latitude coral seas, the upper thermal tolerance limits of many reef-building corals could be exceeded. Present evidence suggests that many corals would be unable to adapt physiologically or genetically to such marked and rapid temperature increases.     

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 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.     

Sequestration: causes and consequences

Adhesion of Plasmodium falciparum-infected erythrocytes to endothelial cells and to syncytiotrophoblasts lining the placenta is a key feature of malaria pathogenesis. P. falciparum erythrocyte membrane protein 1, a family of variable proteins, mediates adhesion to CD36 and intercellular adhesion molecule 1 in the systemic vasculature, and to chondroitin sulphate A and hyaluronic acid in the placenta. Recent studies of the pathology of fatal cerebral malaria and of placental malaria that follow such sequestration suggest that coagulation disturbances may have a greater role in pathogenesis than previously realized, and that monocyte infiltrates in response to malaria may initiate some of these changes. Chemokines such as macrophage inflammatory protein 1 alpha and beta and monocyte chemoattractant protein 1 may play a key role in attracting monocytes to the placenta and other organs, but the stimulus to chemokine secretion is not presently known.     

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.