Global warming, regional trends and inshore environmental conditions influence coral bleaching in Hawaii

Hawaiian waters show a trend of increasing temperature over the past several decades that are consistent with observations in other coral reef areas of the world. The first documented large‐scale coral bleaching occurred in the Hawaii region during late summer of 1996, with a second in 2002. The bleaching events in Hawaii were triggered by a prolonged regional positive oceanic sea surface temperature (SST) anomaly greater than 1°C that developed offshore during the time of annual summer temperature maximum. High solar energy input and low winds further elevated inshore water temperature by 1–2°C in reef areas with restricted water circulation (bays, reef flats and lagoons) and in areas where mesoscale eddies often retain water masses close to shore for prolonged periods of time. Data and observations taken during these events illustrate problems in predicting the phenomena of large‐scale bleaching. Forecasts and hind‐casts of these events are based largely on offshore oceanic SST records, which are only a first approximation of inshore reef conditions. The observed oceanic warming trend is the ultimate cause of the increase in the frequency and severity of bleaching events. However, coral reefs occur in shallow inshore areas where conditions are influenced by winds, orographic cloud cover, complex bathymetry, waves and inshore currents. These factors alter local temperature, irradiance, water motion and other physical and biological variables known to influence bleaching.     

Tropical cyclone cooling combats region‐wide coral bleaching

Coral bleaching has become more frequent and widespread as a result of rising sea surface temperature (SST). During a regional scale SST anomaly, reef exposure to thermal stress is patchy in part due to physical factors that reduce SST to provide thermal refuge. Tropical cyclones (TCs – hurricanes, typhoons) can induce temperature drops at spatial scales comparable to that of the SST anomaly itself. Such cyclone cooling can mitigate bleaching across broad areas when well‐timed and appropriately located, yet the spatial and temporal prevalence of this phenomenon has not been quantified. Here, satellite SST and historical TC data are used to reconstruct cool wakes (n=46) across the Caribbean during two active TC seasons (2005 and 2010) where high thermal stress was widespread. Upon comparison of these datasets with thermal stress data from Coral Reef Watch and published accounts of bleaching, it is evident that TC cooling reduced thermal stress at a region‐wide scale. The results show that during a mass bleaching event, TC cooling reduced thermal stress below critical levels to potentially mitigate bleaching at some reefs, and interrupted natural warming cycles to slow the build‐up of thermal stress at others. Furthermore, reconstructed TC wave damage zones suggest that it was rare for more reef area to be damaged by waves than was cooled (only 12% of TCs). Extending the time series back to 1985 (n = 314), we estimate that for the recent period of enhanced TC activity (1995–2010), the annual probability that cooling and thermal stress co‐occur is as high as 31% at some reefs. Quantifying such probabilities across the other tropical regions where both coral reefs and TCs exist is vital for improving our understanding of how reef exposure to rising SSTs may vary, and contributes to a basis for targeting reef conservation.     

Opposite latitudinal gradients in projected ocean acidification and bleaching impacts on coral reefs

Coral reefs and the services they provide are seriously threatened by ocean acidification and climate change impacts like coral bleaching. Here, we present updated global projections for these key threats to coral reefs based on ensembles of IPCC AR5 climate models using the new Representative Concentration Pathway (RCP) experiments. For all tropical reef locations, we project absolute and percentage changes in aragonite saturation state (Ωarag) for the period between 2006 and the onset of annual severe bleaching (thermal stress >8 degree heating weeks); a point at which it is difficult to believe reefs can persist as we know them. Severe annual bleaching is projected to start 10–15 years later at high‐latitude reefs than for reefs in low latitudes under RCP8.5. In these 10–15 years, Ωarag keeps declining and thus any benefits for high‐latitude reefs of later onset of annual bleaching may be negated by the effects of acidification. There are no long‐term refugia from the effects of both acidification and bleaching. Of all reef locations, 90% are projected to experience severe bleaching annually by 2055. Furthermore, 5% declines in calcification are projected for all reef locations by 2034 under RCP8.5, assuming a 15% decline in calcification per unit of Ωarag. Drastic emissions cuts, such as those represented by RCP6.0, result in an average year for the onset of annual severe bleaching that is ~20 years later (2062 vs. 2044). However, global emissions are tracking above the current worst‐case scenario devised by the scientific community, as has happened in previous generations of emission scenarios. The projections here for conditions on coral reefs are dire, but provide the most up‐to‐date assessment of what the changing climate and ocean acidification mean for the persistence of coral reefs.     

The past,present, and future of coral heat stress studies

The global loss and degradation of coral reefs, as a result of intensified frequency and severity of bleaching events, is a major concern. Evidence of heat stress affecting corals through loss of symbionts and consequent coral bleaching was first reported in the 1930s. However, it was not until the 1998 major global bleaching event that the urgency for heat stress studies became internationally recognized. Current efforts focus not only on examining the consequences of heat stress on corals but also on finding strategies to potentially improve thermal tolerance and aid coral reefs survival in future climate scenarios. Although initial studies were limited in comparison with modern technological tools, they provided the foundation for many of today's research methods and hypotheses. Technological advancements are providing new research prospects at a rapid pace. Understanding how coral heat stress studies have evolved is important for the critical assessment of their progress. This review summarizes the development of the field to date and assesses avenues for future research.     

Cumulative bleaching undermines systemic resilience of the Great Barrier Reef

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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。     

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.     

Downscaled projections of Caribbean coral bleaching that can inform conservation planning

Projections of climate change impacts on coral reefs produced at the coarse resolution (~1°) of Global Climate Models (GCMs) have informed debate but have not helped target local management actions. Here, projections of the onset of annual coral bleaching conditions in the Caribbean under Representative Concentration Pathway (RCP) 8.5 are produced using an ensemble of 33 Coupled Model Intercomparison Project phase‐5 models and via dynamical and statistical downscaling. A high‐resolution (~11 km) regional ocean model (MOM4.1) is used for the dynamical downscaling. For statistical downscaling, sea surface temperature (SST) means and annual cycles in all the GCMs are replaced with observed data from the ~4‐km NOAA Pathfinder SST dataset. Spatial patterns in all three projections are broadly similar; the average year for the onset of annual severe bleaching is 2040–2043 for all projections. However, downscaled projections show many locations where the onset of annual severe bleaching (ASB) varies 10 or more years within a single GCM grid cell. Managers in locations where this applies (e.g., Florida, Turks and Caicos, Puerto Rico, and the Dominican Republic, among others) can identify locations that represent relative albeit temporary refugia. Both downscaled projections are different for the Bahamas compared to the GCM projections. The dynamically downscaled projections suggest an earlier onset of ASB linked to projected changes in regional currents, a feature not resolved in GCMs. This result demonstrates the value of dynamical downscaling for this application and means statistically downscaled projections have to be interpreted with caution. However, aside from west of Andros Island, the projections for the two types of downscaling are mostly aligned; projected onset of ASB is within ±10 years for 72% of the reef locations.     

Refugia under threat: Mass bleaching of coral assemblages in high‐latitude eastern Australia

Environmental anomalies that trigger adverse physiological responses and mortality are occurring with increasing frequency due to climate change. At species' range peripheries, environmental anomalies are particularly concerning because species often exist at their environmental tolerance limits and may not be able to migrate to escape unfavourable conditions. Here, we investigated the bleaching response and mortality of 14 coral genera across high‐latitude eastern Australia during a global heat stress event in 2016. We evaluated whether the severity of assemblage‐scale and genus‐level bleaching responses was associated with cumulative heat stress and/or local environmental history, including long‐term mean temperatures during the hottest month of each year (SST_LTMAX), and annual fluctuations in water temperature (SST_VAR) and solar irradiance (PARZ_VAR). The most severely‐bleached genera included species that were either endemic to the region (Pocillopora aliciae) or rare in the tropics (e.g. Porites heronensis). Pocillopora spp., in particular, showed high rates of immediate mortality. Bleaching severity of Pocillopora was high where SST_LTMAX was low or PARZ_VAR was high, whereas bleaching severity of Porites was directly associated with cumulative heat stress. While many tropical Acropora species are extremely vulnerable to bleaching, the Acropora species common at high latitudes, such as A. glauca and A. solitaryensis, showed little incidence of bleaching and immediate mortality. Two other regionally‐abundant genera, Goniastrea and Turbinaria, were also largely unaffected by the thermal anomaly. The severity of assemblage‐scale bleaching responses was poorly explained by the environmental parameters we examined. Instead, the severity of assemblage‐scale bleaching was associated with local differences in species abundance and taxon‐specific bleaching responses. The marked taxonomic disparity in bleaching severity, coupled with high mortality of high‐latitude endemics, point to climate‐driven simplification of assemblage structures and progressive homogenisation of reef functions at these high‐latitude locations.     

Turbid reefs moderate coral bleaching under climate‐related temperature stress

Thermal‐stress events that cause coral bleaching and mortality have recently increased in frequency and severity. Yet few studies have explored conditions that moderate coral bleaching. Given that high light and high ocean temperature together cause coral bleaching, we explore whether corals at turbid localities, with reduced light, are less likely to bleach during thermal‐stress events than corals at other localities. We analyzed coral bleaching, temperature, and turbidity data from 3,694 sites worldwide with a Bayesian model and found that K_d490, a measurement positively related to turbidity, between 0.080 and 0.127 reduced coral bleaching during thermal‐stress events. Approximately 12% of the world's reefs exist within this “moderating turbidity” range, and 30% of reefs that have moderating turbidity are in the Coral Triangle. We suggest that these turbid nearshore environments may provide some refuge through climate change, but these reefs will need high conservation status to sustain them close to dense human populations.     

New insights into global patterns of ocean temperature anomalies: implications for coral reef health and management

Aim Coral reefs are widely considered to be particularly vulnerable to changes in ocean temperatures, yet we understand little about the broad‐scale spatio‐temporal patterns that may cause coral mortality from bleaching and disease. Our study aimed to characterize these ocean temperature patterns at biologically relevant scales. Location Global, with a focus on coral reefs. Methods We created a 4‐km resolution, 21‐year global ocean temperature anomaly (deviations from long‐term means) database to quantify the spatial and temporal characteristics of temperature anomalies related to both coral bleaching and disease. Then we tested how patterns varied in several key metrics of disturbance severity, including anomaly frequency, magnitude, duration and size. Results Our analyses found both global variation in temperature anomalies and fine‐grained spatial variability in the frequency, duration and magnitude of temperature anomalies. However, we discovered that even during major climatic events with strong spatial signatures, like the El Niño–Southern Oscillation, areas that had high numbers of anomalies varied between years. In addition, we found that 48% of bleaching‐related anomalies and 44% of disease‐related anomalies were less than 50 km², much smaller than the resolution of most models used to forecast climate changes. Main conclusions The fine‐scale variability in temperature anomalies has several key implications for understanding spatial patterns in coral bleaching‐ and disease‐related anomalies as well as for designing protected areas to conserve coral reefs in a changing climate. Spatial heterogeneity in temperature anomalies suggests that certain reefs could be targeted for protection because they exhibit differences in thermal stress. However, temporal variability in anomalies could complicate efforts to protect reefs, because high anomalies in one year are not necessarily predictive of future patterns of stress. Together, our results suggest that temperature anomalies related to coral bleaching and disease are likely to be highly heterogeneous and could produce more localized impacts of climate change.     

Boring sponges,an increasing threat for coral reefs affected by bleaching events

Coral bleaching is a stress response of corals induced by a variety of factors, but these events have become more frequent and intense in response to recent climate‐change‐related temperature anomalies. We tested the hypothesis that coral reefs affected by bleaching events are currently heavily infested by boring sponges, which are playing a significant role in the destruction of their physical structure. Seventeen reefs that cover the entire distributional range of corals along the Mexican Pacific coast were studied between 2005/2006, and later between 2009/2010. Most of these coral reefs were previously impacted by bleaching events, which resulted in coral mortalities. Sponge abundance and species richness was used as an indicator of bioerosion, and coral cover was used to describe the present condition of coral reefs. Coral reefs are currently highly invaded (46% of the samples examined) by a very high diversity of boring sponges (20 species); being the coral reef framework the substrate most invaded (56%) followed by the rubbles (45%), and the living colonies (36%). The results also indicated that boring sponges are promoting the dislodgment of live colonies and large fragments from the framework. In summary, the eastern coral reefs affected by bleaching phenomena, mainly provoked by El Niño, present a high diversity and abundance of boring sponges, which are weakening the union of the colony with the reef framework and promoting their dislodgment. These phenomena will probably become even more intense and severe, as temperatures are projected to continue to rise under the scenarios for future climate change, which could place many eastern coral reefs beyond their survival threshold.     

Thermal refugia against coral bleaching throughout the northern Red Sea

Tropical reefs have been impacted by thermal anomalies caused by global warming that induced coral bleaching and mortality events globally. However, there have only been very few recordings of bleaching within the Red Sea despite covering a latitudinal range of 15° and consequently it has been considered a region that is less sensitive to thermal anomalies. We therefore examined historical patterns of sea surface temperature (SST) and associated anomalies (1982–2012) and compared warming trends with a unique compilation of corresponding coral bleaching records from throughout the region. These data indicated that the northern Red Sea has not experienced mass bleaching despite intensive Degree Heating Weeks (DHW) of >15°C‐weeks. Severe bleaching was restricted to the central and southern Red Sea where DHWs have been more frequent, but far less intense (DHWs <4°C‐weeks). A similar pattern was observed during the 2015–2016 El Niño event during which time corals in the northern Red Sea did not bleach despite high thermal stress (i.e. DHWs >8°C‐weeks), and bleaching was restricted to the central and southern Red Sea despite the lower thermal stress (DHWs < 8°C‐weeks). Heat stress assays carried out in the northern (Hurghada) and central (Thuwal) Red Sea on four key reef‐building species confirmed different regional thermal susceptibility, and that central Red Sea corals are more sensitive to thermal anomalies as compared to those from the north. Together, our data demonstrate that corals in the northern Red Sea have a much higher heat tolerance than their prevailing temperature regime would suggest. In contrast, corals from the central Red Sea are close to their thermal limits, which closely match the maximum annual water temperatures. The northern Red Sea harbours reef‐building corals that live well below their bleaching thresholds and thus we propose that the region represents a thermal refuge of global importance.     

The impact of rising temperatures on the prevalence of coral diseases and its predictability: A global meta-analysis

Coral reefs are under threat from disease as climate change alters environmental conditions. Rising temperatures exacerbate coral disease, but this relationship is likely complex as other factors also influence coral disease prevalence. To better understand this relationship, we meta-analytically examined 108 studies for changes in global coral disease over time alongside temperature, expressed using average summer sea surface temperature (SST) and cumulative heat stress as weekly sea surface temperature anomalies (WSSTAs). We found that both rising average summer SST and WSSTA were associated with global increases in the mean and variability in coral disease prevalence. Global coral disease prevalence tripled, reaching 9.92% in the 25 years examined, and the effect of ‘year’ became more stable (i.e. prevalence has lower variance over time), contrasting the effects of the two temperature stressors. Regional patterns diverged over time and differed in response to average summer SST. Our model predicted that, under the same trajectory, 76.8% of corals would be diseased globally by 2100, even assuming moderate average summer SST and WSSTA. These results highlight the need for urgent action to mitigate coral disease. Mitigating the impact of rising ocean temperatures on coral disease is a complex challenge requiring global discussion and further study.     

The cumulative impact of annual coral bleaching can turn some coral species winners into losers

Mass coral bleaching events caused by elevated seawater temperatures result in extensive coral loss throughout the tropics, and are projected to increase in frequency and severity. If bleaching becomes an annual event later in this century, more than 90% of coral reefs worldwide may be at risk of long‐term degradation. While corals can recover from single isolated bleaching and can acclimate to recurring bleaching events that are separated by multiple years, it is currently unknown if and how they will survive and possibly acclimatize to annual coral bleaching. Here, we demonstrate for the first time that annual coral bleaching can dramatically alter thermal tolerance in Caribbean corals. We found that high coral energy reserves and changes in the dominant algal endosymbiont type (Symbiodinium spp.) facilitated rapid acclimation in Porites divaricata, whereas low energy reserves and a lack of algal phenotypic plasticity significantly increased susceptibility in Porites astreoides to bleaching the following year. Phenotypic plasticity in the dominant endosymbiont type of Orbicella faveolata did not prevent repeat bleaching, but may have facilitated rapid recovery. Thus, coral holobiont response to an isolated single bleaching event is not an accurate predictor of its response to bleaching the following year. Rather, the cumulative impact of annual coral bleaching can turn some coral species ‘winners’ into ‘losers’, and can also facilitate acclimation and turn some coral species ‘losers’ into ‘winners’. Overall, these findings indicate that cumulative impact of annual coral bleaching could result in some species becoming increasingly susceptible to bleaching and face a long‐term decline, while phenotypically plastic coral species will acclimatize and persist. Thus, annual coral bleaching and recovery could contribute to the selective loss of coral diversity as well as the overall decline of coral reefs in the Caribbean.     

Thermally tolerant corals have limited capacity to acclimatize to future warming

Thermal stress affects organism performance differently depending on the ambient temperature to which they are acclimatized, which varies along latitudinal gradients. This study investigated whether differences in physiological responses to temperature are consistent with regional differences in temperature regimes for the stony coral Oculina patagonica. To resolve this question, we experimentally assessed how colonies originating from four different locations characterized by >3 °C variation in mean maximum annual temperature responded to warming from 20 to 32 °C. We assessed plasticity in symbiont identity, density, and photosynthetic properties, together with changes in host tissue biomass. Results show that, without changes in the type of symbiont hosted by coral colonies, O. patagonica has limited capacity to acclimatize to future warming. We found little evidence of variation in overall thermal tolerance, or in thermal optima, in response to spatial variation in ambient temperature. Given that the invader O. patagonica is a relatively new member of the Mediterranean coral fauna, our results also suggest that coral populations may need to remain isolated for a long period of time for thermal adaptation to potentially take place. Our study indicates that for O. patagonica, mortality associated with thermal stress manifests primarily through tissue breakdown under moderate but prolonged warming (which does not impair symbiont photosynthesis and, therefore, does not lead to bleaching). Consequently, projected global warming is likely to cause repeat incidents of partial and whole colony mortality and might drive a gradual range contraction of Mediterranean corals.     

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.     

Resolving spatial and temporal patterns of coral bleaching risk using image analysis: an active learning experience to improve climate change literacy in college students

Recent studies indicate poor understanding of the causes and consequences of climate change among college students. In an effort to improve climate change literacy, we have developed an authentic research experience for upper level undergraduate students focused on resolving spatial and temporal patterns of coral reef bleaching, an ecologically and economically important consequence of climate warming. In the research, students use a public archive of maps generated by the United States National Oceanographic and Atmospheric Association (NOAA) that use coloration to depict ocean areas experiencing above-average surface temperatures and where corals are at an increased risk of bleaching. Students are required to quantify the total area of coloration on individual maps using open-source image analysis software called Image J. By quantifying coloration (ie bleaching risk) over a large number of maps in a chronological sequence, students can test hypotheses regarding the relationship between ongoing climate warming and coral bleaching risk. Students are required to summarise their findings in a scientific journal-style report that incorporates graphical representations and statistical tests of their coral bleaching risk data. The research activity is cost-effective, repeatable, requires little specialised knowledge and addresses common programmatic learning outcomes that target scientific communication, quantitative reasoning and sustainability.     

Corals escape bleaching in regions that recently and historically experienced frequent thermal stress

The response of coral-reef ecosystems to contemporary thermal stress may be in part a consequence of recent or historical sea-surface temperature (SST) variability. To test this hypothesis, we examined whether: (i) there was a relationship between the historical frequency of SST variability and stress experienced during the most recent thermal-stress events (in 1998 and 2005–2006) and (ii) coral reefs that historically experienced frequent thermal anomalies were less likely to experience coral bleaching during these recent thermal-stress events. Examination of nine detrended coral δ¹⁸O and Sr/Ca anomaly records revealed a high- (5.7-year) and low-frequency (>54-year) mode of SST variability. There was a positive relationship between the historical frequency of SST anomalies and recent thermal stress; sites historically dominated by the high-frequency mode experienced greater thermal stress than other sites during both events, and showed extensive coral bleaching in 1998. Nonetheless, in 2005–2006, corals at sites dominated by high-frequency variability showed reduced bleaching, despite experiencing high thermal stress. This bleaching resistance was most likely a consequence of rapid directional selection that followed the extreme thermal event of 1998. However, the benefits of regional resistance could come at the considerable cost of shifts in coral species composition.