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Coral bleaching occurs when coralpolyps expel algae that live inside their tissues. Normally, coral polyps live in an endosymbiotic relationship with the algae and that relationship is crucial for the coral and hence for the health of the whole reef. Bleached corals might continue to live for some time. But as the algae provide the coral with up to 90% of its energy, after expelling the algae the coral begins to starve. After bleaching, some corals may recover, while some die. Above-average sea water temperatures caused by global warming have been identified as a leading cause for coral bleaching worldwide. Between 2014 and 2016, the longest global bleaching events ever were recorded. According to the United Nations Environment Programme, these bleaching events killed coral on an unprecedented scale. In 2016, bleaching of coral on the Great Barrier Reef killed between 29 and 50 percent of the reef's coral. In 2017, the bleaching further expanded to areas of the reef that were previously spared, such as the central one. The average interval between bleaching events has halved between 1980 and 2016.
Coral and microscopic algae have a symbiotic relationship. When water temperatures get too high, the algae leave the coral tissue and the coral begins to starve.
The corals that form the great reefecosystems of tropical seas depend upon a symbiotic relationship with algae-like single-celled flagellateprotozoa called zooxanthellae that live within their tissues and give the coral its coloration. The zooxanthellae provide the coral with nutrients through photosynthesis, a crucial factor in the clear and nutrient-poor tropical waters. In exchange, the coral provide the zooxanthellae with the carbon dioxide and ammonium needed for photosynthesis. Negative environmental conditions thwart the coral's ability to provide for the zooxanthellae's needs. To ensure short-term survival, the coral-polyp then expels the zooxanthellae. This leads to a lighter or completely white appearance, hence the term "bleached". As the zooxanthellae provide up to 90% of the coral's energy needs through products of photosynthesis, after expelling, the coral may begin to starve.
Healthy coral at left and bleached, but still living, coral to right
Coral can survive short-term disturbances, but if the conditions that lead to the expulsion of the zooxanthellae persist, the coral's chances of survival diminish. In order to recover from bleaching, the zooxanthellae have to re-enter the tissues of the coral polyps and restart photosynthesis to sustain the coral as a whole and the ecosystem that depends on it. If the coral polyps die of starvation after bleaching, they will decay. The hard coral species will then leave behind their calcium carbonateskeletons, which will be taken over by algae, effectively blocking coral re-growth. Eventually, the coral skeletons will erode, causing the reef structure to collapse.
Coral bleaching may be caused by a number of factors. While localized triggers lead to localized bleaching, the large scale coral bleaching events of the recent years have been triggered by increased sea surface temperatures. Coral reefs located in warm, shallow water with low water flow have been more affected than reefs located in areas with higher water flow.
ocean acidification due to elevated levels of CO2 caused by air pollution
Mass bleaching events
Bleached Acropora coral (foreground) and normal colony (background), Keppel Islands, Great Barrier Reef
Elevated sea water temperatures are the main cause of mass bleaching events. Sixty major episodes of coral bleaching have occurred between 1979 and 1990, with the associated coral mortality affecting reefs in every part of the world. In 2016, the longest coral bleaching event was recorded.
Factors that influence the outcome of a bleaching event include stress-resistance which reduces bleaching, tolerance to the absence of zooxanthellae, and how quickly new coral grows to replace the dead. Due to the patchy nature of bleaching, local climatic conditions such as shade or a stream of cooler water can reduce bleaching incidence. Coral and zooxanthellae health and genetics also influence bleaching.
Large coral colonies such as Porites are able to withstand extreme temperature shocks, while fragile branching corals such Acropora are far more susceptible to stress following a temperature change. Corals consistently exposed to low stress levels may be more resistant to bleaching.
Scientists believe that the oldest known bleaching was that of the Late Devonian (Frasnian/Famennian), also triggered by the rise of sea surface temperatures. It resulted in the demise of the largest coral reefs in the Earth's history 
Two images of the Great Barrier Reef showing that the warmest water (top picture) coincides with the coral reefs (lower picture), setting up conditions that can cause coral bleaching
In the 2012–2040 period, coral reefs are expected to experience more frequent bleaching events. The Intergovernmental Panel on Climate Change (IPCC) sees this as the greatest threat to the world's reef systems. Coral reefs worldwide were lost by 19% and 60% of the remaining reefs are at immediate risk of being lost. There are a few ways to tell the impacts of coral bleaching on reefs. First by the coral cover, the more coral that is covering the ground the less of an impact bleaching had. Second, coral abundance, which is the number of different living species on the coral reef. (Shidqi et al. 2017)
Great Barrier Reef
The Great Barrier Reef along the coast of Australia experienced bleaching events in 1980, 1982, 1992, 1994, 1998, 2002, 2006, 2016 and 2017. Some locations suffered severe damage, with up to 90% mortality. The most widespread and intense events occurred in the summers of 1998 and 2002, with 42% and 54% respectively of reefs bleached to some extent, and 18% strongly bleached. However coral losses on the reef between 1995 and 2009 were largely offset by growth of new corals. An overall analysis of coral loss found that coral populations on the Great Barrier Reef had declined by 50.7% from 1985 to 2012, but with only about 10% of that decline attributable to bleaching, and the remaining 90% caused about equally by tropical cyclones and by predation by crown-of-thorns starfishes. A global mass coral bleaching has been occurring since 2014 because of the highest recorded temperatures plaguing oceans. These temperatures have caused the most severe and widespread coral bleaching ever recorded in the Great Barrier reef. The most severe bleaching in 2016 occurred near Port Douglas. In late November 2016 surveys of 62 reefs showed that long term heat stress from climate change caused a 29% loss of shallow water coral. The highest coral death and reef habitat loss was inshore and mid-self reefs around Cape Grenville and Princess Charlotte Bay.  The IPCC's moderate warming scenarios (B1 to A1T, 2 °C by 2100, IPCC, 2007, Table SPM.3, p. 13) forecast that corals on the Great Barrier Reef are very likely to regularly experience summer temperatures high enough to induce bleaching.
According to a 2017 Japanese government report, almost 75% of Japan's largest coral reef in Okinawa has died from bleaching.
Coral reef provinces have been permanently damaged by warm sea temperatures, most severely in the Indian Ocean. Up to 90% of coral cover has been lost in the Maldives, Sri Lanka, Kenya and Tanzania and in the Seychelles during the massive 1997-98 bleaching event.
More than 60% of the coral in the Maldives has suffered from bleaching in 2016.
Thailand experienced a severe mass bleaching in 2010 which affected 70% of the coral in the Andaman Sea. Between 30% and 95% of the bleached coral died.
In 2017 there was a study done on two islands in Indonesia to see how their coral cover was. One of the places was Melinjo Islands and the other was Saktu Islands. In Saktu Island the lifeform conditions were categorized as bad, with an average coral cover of 22.3%. In Melinjo Islands the lifeform conditions were categorized as bad, with an average coral cover of 22.2%. (Shidqi et al. 2017)
The first recorded mass bleaching event that took place in the Belize Barrier Reef was in 1998, where sea level temperatures reached up to 31.5 °C (88.7 °F) from 10 August to 14 October. For a few days, Hurricane Mitch brought in stormy weather on October 27 but only reduced temperatures by 1 degree or less. During this time period, mass bleaching in the fore-reef and lagoon occurred. While some fore reef colonies suffered some damage, coral mortality in the lagoon was catastrophic.
The most prevalent coral in the reefs of Belize in 1998 was the lettuce coral, Agaricia tenuifolia. On 22 and 23 October, surveys were conducted at two sites and the findings were devastating. Virtually all the living coral was bleached white and their skeletons indicated that they had died recently. At the lagoon floor, complete bleaching was evident among Agaricia tenuifolia. Furthermore, surveys done in 1999 and 2000 showed a near total mortality of Agaricia tenuifolia at all depths. Similar patterns occurred in other coral species as well. Measurements on water turbidity suggest that these mortalities were attributed to rising water temperatures rather than solar radiation.
Since 1970 the coral reef cover in the Caribbean has decreased from 50% to only 10%. In 2013 there was a study done to see how the coral was doing after the mass bleaching event in 2010. Only one year later in 2011, the majority of the dominant species declined by about 62%. As well as the dominant species the coral abundance declined by about 50%. However, between 2011 and 2013 the cover of coral did increase for 10 of the 26 dominant species but declined for 5 other populations (Buglass, et al. 2016).
Coral in the south Red Sea does not bleach despite summer water temperatures up to 34 °C (93 °F). Coral bleaching in the Red Sea is more common in the northern section of the reefs, the southern part of the reef has been plagued by coral eating starfish, dynamite fishing and human impacts on the environment. In 1988 there was a massive bleaching event that affected the reefs in Saudi Arabia and in Sudan, the southern reefs were more resilient and affected them very little. Previously it was thought that the North suffers more from coral bleaching but they show a fast turnover of coral and the southern reef was thought to not suffer from bleaching as harshly, they show more consistency. However, new research shows where the south reef should be bigger and healthier than the north it was not. This is believed to be because of major disturbances in recent history from bleaching events, and coral eating starfish. In 2010, coral bleaching occurred in Saudi Arabia and Sudan, where the temperature rose 10 to 11 degrees. Certain taxa experienced 80% to 100% of their colonies bleaching, while some showed on average 20% of that taxa bleaching.
Economic and political impact
According to Brian Skoloff of The Christian Science Monitor, "If the reefs vanished, experts say, hunger, poverty and political instability could ensue." Since countless sea life depend on the reefs for shelter and protection from predators, the extinction of the reefs would ultimately create a domino effect that would trickle down to the many human societies that depend on those fish for food and livelihood. There has been a 44% decline over the last 20 years in the Florida Keys, and up to 80% in the Caribbean alone.
Coral reefs provide various ecosystem services, one of which is being a natural fishery, as many frequently consumed commercial fish spawn or live out their juvenile lives in coral reefs around the tropics. Thus, reefs are a popular fishing site and are an important source of income for fishers, especially small, local fisheries. As coral reef habitat decreases due to bleaching, reef associated fish populations also decrease, which affects fishing opportunities. A model from one study by Speers et al. calculated direct losses to fisheries from decreased coral cover to be around $49 – $69 billion, if human societies continue to emit high levels of greenhouse gases. But, these losses could be reduced for a consumer surplus benefit of about $14 – $20 billion, if societies chose to emit a lower level of greenhouse gases instead. These economic losses also have important political implications, as they fall disproportionately on developing countries where the reefs are located, namely in Southeast Asia and around the Indian Ocean. It would cost more for countries in these areas to respond to coral reef loss as they would need to turn to different sources of income and food, in addition to losing other ecosystem services such as ecotourism. A study completed by Chen et al. suggested that the commercial value of reefs decreases by almost 4% every time coral cover decreases by 1% because of losses in ecotourism and other potential outdoor recreational activities.
Coral reefs also act as a protective barrier for coastlines by reducing wave impact, which lowers the damage from storms, erosions, and flooding. Countries that lose this natural protection will lose more money because of the increased susceptibility of storms. This indirect cost, combined with the lost revenue in tourism, will result in enormous economic effects.
Monitoring reef sea surface temperature
The US National Oceanic and Atmospheric Administration (NOAA) monitors for bleaching "hot spots", areas where sea surface temperature rises 1 °C or more above the long-term monthly average. The "hot spots" are the location in which thermal stress is measured and with the development of Degree Heating Week (DHW), the coral reef's thermal stress is monitored . Global coral bleaching is being detected earlier due to the satellite remote sensing the rise of sea temperatures. It is necessary to monitor the high temperatures because coral bleaching events are affecting coral reef reproduction and normal growth capacity, as well as it weakening corals, eventually leading to their mortality. This system detected the worldwide 1998 bleaching event, that corresponded to the 1997–98 El Niño event. Currently, 190 reef sites around the globe are monitored by the NOAA, and send alerts to research scientists and reef managers via NOAA Coral Reef Watch (CRW) website. By monitoring the warming of sea temperatures, the early warnings of coral bleaching, alerts reef managers to prepare and draw awareness to future bleaching events.
Changes in ocean chemistry
Increasing ocean acidification due to rises in carbon dioxide levels exacerbates the bleaching effects of thermal stress. Acidification affects the corals' ability to create calcareous skeletons, essential to their survival. This is because ocean acidification decreases the amount of carbonate ion in the water, making it more difficult for corals to absorb the calcium carbonate they need for the skeleton. As a result, the resilience of reefs goes down, while it becomes easier for them to erode and dissolve. In addition, the increase in CO2 allows herbivore overfishing and nutrification to change coral-dominated ecosystems to algal-dominated ecosystems. A recent study from the Atkinson Center for a Sustainable Future found that with the combination of acidification and temperature rises, the levels of CO2 could become too high for coral to survive in as little as 50 years.
Infectious bacteria of the species Vibrio shiloi are the bleaching agent of Oculina patagonica in the Mediterranean Sea, causing this effect by attacking the zooxanthellae.V. shiloi is infectious only during warm periods. Elevated temperature increases the virulence of V. shiloi, which then become able to adhere to a beta-galactoside-containing receptor in the surface mucus of the host coral.V. shiloi then penetrates the coral's epidermis, multiplies, and produces both heat-stable and heat-sensitive toxins, which affect zooxanthellae by inhibiting photosynthesis and causing lysis.
During the summer of 2003, coral reefs in the Mediterranean Sea appeared to gain resistance to the pathogen, and further infection was not observed. The main hypothesis for the emerged resistance is the presence of symbiotic communities of protective bacteria living in the corals. The bacterial species capable of lysing V. shiloi had not been identified as of 2011.
In 2010, researchers at Penn State discovered corals that were thriving while using an unusual species of symbiotic algae in the warm waters of the Andaman Sea in the Indian Ocean. Normal zooxanthellae cannot withstand temperatures as high as was there, so this finding was unexpected. This gives researchers hope that with rising temperatures due to global warming, coral reefs will develop tolerance for different species of symbiotic algae that are resistant to high temperature, and can live within the reefs. In 2010, researchers from Stanford University also found corals around the Samoan Islands that experience a drastic temperature increase for about four hours a day during low tide. The corals do not bleach or die regardless of the high heat increase. Studies showed that the corals off the coast of Ofu Island near America Samoa have become trained to withstand the high temperatures. Researchers are now asking a new question: can we condition corals, that are not from this area, in this manner and slowly introduce them to higher temperatures for short periods of time and make them more resilient against rising ocean temperatures.
Recovery and macroalgal regime shifts
After corals experience a bleaching event to increased temperature stress some reefs are able to return to their original, pre-bleaching state. Reefs either recover from bleaching, where they are recolonized by zooxanthellae, or they experience a regime shift, where previously flourishing coral reefs are taken over by thick layers of macroalgae. This inhibits further coral growth because the algae produces antifouling compounds to deter settlement and competes with corals for space and light. As a result, macroalgae forms stable communities that make it difficult for corals to grow again. Reefs will then be more susceptible to other issues, such as declining water quality and removal of herbivore fish, because coral growth is weaker. Discovering what causes reefs to be resilient or recover from bleaching events is of primary importance because it helps inform conservation efforts and protect coral more effectively.
Corals have shown to be resilient to short-term disturbances. Recovery has been shown in after storm disturbance and crown of thorns starfish invasions. Fish species tend to fare better following reef disturbance than coral species as corals show limited recovery and reef fish assemblages have shown little change as a result of short-term disturbances. In contrast, fish assemblages in reefs that experience bleaching exhibit potentially damaging changes. One study by Bellwood et al. notes that while species richness, diversity, and abundance did not change, fish assemblages contained more generalist species and less coral dependent species. Responses to coral bleaching are diverse between reef fish species, based on what resources are affected. Rising sea temperature and coral bleaching do not directly impact adult fish mortality, but there are many indirect consequences of both. Coral-associated fish populations tend to be in decline due to habitat loss; however, some herbivorous fish populations have seen a drastic increase due to the increase of algae colonization on dead coral. Studies note that better methods are needed to measure the effects of disturbance on the resilience of corals.
The lemon damselfish (Pomacentrus moluccensis) is a coral associated species that has been shown to decline dramatically following coral bleaching.
Until recently, the factors mediating the recovery of coral reefs from bleaching were not well studied. Research by Graham et al. (2005) studied 21 reefs around Seychelles in the Indo-Pacific in order to document the long-term effects of coral bleaching. After the loss of more than 90% of corals due to bleaching in 1998 around 50% of the reefs recovered and roughly 40% of the reefs experienced regime shifts to macroalgae dominated compositions. After an assessment of factors influencing the probability of recovery, the study identified five major factors: density of juvenile corals, initial structural complexity, water depth, biomass of herbivorous fishes, and nutrient conditions on the reef. Overall, resilience was seen most in coral reef systems that were structurally complex and in deeper water.
The ecological roles and functional groups of species also play a role in the recovery of regime shifting potential in reef systems. Coral reefs are affected by bioeroding, scraping, and grazing fish species. Bioeroding species remove dead corals, scraping species remove algae and sediment to further future growth, grazing species remove algae. The presence of each type of species can influence the ability for normal levels of coral recruitment which is an important part of coral recovery. Lowered numbers of grazing species after coral bleaching in the Caribbean has been likened to sea-urchin-dominated systems which do not undergo regime shifts to fleshy macroalgae dominated conditions.
There is always the possibility of unobservable changes, or cryptic losses or resilience, in a coral community's ability to perform ecological processes. These cryptic losses can result in unforeseen regime changes or ecological flips. More detailed methods for determining the health of coral reefs that take into account long-term changes to the coral ecosystems and better-informed conservation policies are necessary to protect coral reefs in the years to come.
Economic value of coral reefs
Coral reefs provide goods and services to an estimated quarter of all ocean species.  Experts estimate that coral reef services are worth up to $1.2 million per hectare which translates to an average of $172 billion per year. The benefits of coral reefs include providing physical structures such as coastal shoreline protection, biotic services within and between ecosystems, biogeochemical services such as maintaining nitrogen levels in the ocean, climate records, and recreational and commercial (tourism) services.
Cost benefit analysis of reducing loss of coral reefs
In 2010, the Convention on Biological Diversity's (CBD) Strategic Plan for Biodiversity 2011-2020 created twenty distinct targets for sustainable development for post-2015. Target 10 indicates the goal of minimizing "anthropogenic pressures on coral reefs". Two programs were looked at, one that reduces coral reef loss by 50% that has a capital cost of $684 million and a recurrent cost of $81 million. The other program reduces coral reef loss by 80% and has a capital cost of $1,036 million with recurring costs of $130 million. CBD acknowledges that they may be underestimating the costs and resources needed to achieve this target due to lack of relevant data but nonetheless, the cost-benefit analysis shows that the benefits outweigh the costs by a great enough amount for both programs (Benefit Cost Ratio of 95.3 and 98.5) that "there is ample scope to increase outlays on coral protection and still achieve a benefit to cost ratio that is well over one.
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