Resilience of coral reefs

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The resilience of coral reefs is the biological ability of coral reefs to recover from natural and anthropogenic disturbances such as storms and bleaching episodes. [1] Resilience refers to the ability of biological or social systems to overcome pressures and stresses by maintaining key functions through resisting or adapting to change. [2] Reef resistance measures how well coral reefs tolerate changes in ocean chemistry, sea level, and sea surface temperature. [3] Reef resistance and resilience are important factors in coral reef recovery from the effects of ocean acidification. Natural reef resilience can be used as a recovery model for coral reefs and an opportunity for management in marine protected areas (MPAs).

Contents

Thermal tolerance

Many corals rely on a symbiotic algae called zooxanthellae for nutrient uptake through photosynthesis. Corals obtain about 60-85% of their total nutrition from symbiotic zooxanthellae. [4] Slight increases in sea surface temperature can cause zooxanthellae to die. Coral hosts become bleached when they lose their zooxanthellae. Differences in symbionts, determined by genetic groupings (clades A-H), may explain thermal tolerance in corals. [5] Research has shown that some corals contain thermally-resistant clades of zooxanthellae. Corals housing primarily clade D symbionts, and certain types of thermally resistant clade C symbionts, allow corals to avoid bleaching as severely as others experiencing the same stressor. [6] Scientists remain in debate if thermal resistance in corals is due to mixing or shifting of symbionts, or thermally resistant vs. thermally-sensitive types of zooxanthellae. Species of coral housing multiple types of zooxanthellae can withstand a 1-1.5 °C change in temperature. [6] However, few species of coral are known to house multiple types of zooxanthellae. Corals are more likely to contain clade D symbionts after multiple coral bleaching events. [6]

Reef recovery

Corals "seeded" with thermally-resistant zooxanthellae may become more resilient and able to resist coral bleaching Keppelbleaching.jpg
Corals "seeded" with thermally-resistant zooxanthellae may become more resilient and able to resist coral bleaching

Research studies of the Mediterranean species of coral Oculina patagonica [7] reveal that the presence of endolithic algae in coral skeletons may provide additional energy which can result in post-bleaching recovery. [8] During bleaching, the loss of zooxanthellae decreases the amount of light absorbed by coral tissue, which allows increased amounts of photosynthetically active radiation to penetrate the coral skeleton. Greater amounts of photosynthetically active radiation in coral skeletons cause an increase in endolithic algae biomass and production of photoassimilates. [8] During bleaching, the energy input to the coral tissue of phototrophic endoliths expand as the energy input of the zooxanthellae dwindles. This additional energy could explain the survival and rapid recovery of O. patagonica after bleaching events. [8] A study by the Australian Research Council proposed that the loss of fast-growing coral could lead to less resilience of the remaining coral. The study was undertaken in both the Caribbean and the Indo-Pacific and reached the conclusion that the latter may be more resilient than the former based on several factors; the process of herbivory and the rates of algal blooms forming. [9]

Coral bleaching effects on biodiversity

Parrot Fish Parrot fish herbivory.jpg
Parrot Fish

Coral bleaching is a major consequence of stress on coral reefs. Bleaching events due to distinct temperature changes, pollution, and other shifts of environmental conditions are detrimental to coral health, but corals can restore from bleaching events if the stress is not chronic. [10] When corals are exposed to a long period of severe stress, death may occur due to the loss of zooxanthellae, which are vital to the coral's survival because of the nutrients they supply. [11] Coral bleaching, degradation, and death have a great effect on the surrounding ecosystem and biodiversity. Coral reefs are important, diverse ecosystems that host a plethora of organisms that contribute different services to maintain reef health. For example, herbivorous reef fish, like the parrotfish, maintain levels of macro algae. The upkeep of seaweed contributes to decreasing space competition for substrate-seeking organisms, like corals, to establish and propagate, creating a stronger, more resilient reef. [12] However, when corals become bleached, organisms often leave the coral reef habitat which in turn takes away the services that they were previously supplying. Reefs also administer many ecosystem services such as food provision for many people around the world who are dependent on fishing reefs to sustain themselves. There is evidence that some species of coral are resilient to elevated sea surface temperatures for a short period of time. [13]

Natural disturbances

Natural forces such as disease and storms degrade corals. The frequency of coral disease caused by microbial pathogens has increased over the years, contributing to coral reef mortality. [14] Bacterial, fungal, viral, and parasitic infections can result in physiological and morphological effects. Some of the most common coral diseases include black band disease, white pox disease, white plague, and white band disease, all of which involve tissue degradation and exposure of the coral skeleton. [15] Diseases such as these can quickly spread among healthy coral reefs, potentially making them more susceptible to injury from disturbances like storms. Storms, including cyclones and hurricanes, can cause mechanical destruction to reefs and a change in sedimentation. [16] The strong waves that result from these disturbances can strike corals, causing them to dislodge, and can also cause the reef to come into contact with released sediments and freshwater.

Anthropogenic disturbances

Anthropogenic forces contribute to coral reef degradation, reducing their resiliency. Some anthropogenic forces that degrade corals include pollution, sedimentation from coastal development, and ocean acidification due to increased fossil fuel emissions. Carbon emissions cause ocean surface waters to warm and acidify. [17] The combustion of fossil fuels results in the emission of greenhouse gases, such as carbon dioxide into the atmosphere. The ocean uptakes some of the emitted carbon dioxide, injurious to the natural processes that occur in the ocean. Ocean acidification results in a lower ocean water pH, negatively affecting the formation of calcium carbonate structures which are imperative to coral development. [18] Developing coastal areas has the potential for chemical and nutrient pollution to run off into surrounding waters. Nutrient pollution causes the overgrowth of aquatic vegetation which has the ability to out-compete corals for space, nutrients, and other resources. [19] Overfishing can also have devastating effects on coral reefs. Due to the food security that reefs hold, they are often overfished, which can cause reef ecosystems to be unable to reconstruct after damage has been done. [20] Restoration can be challenging due to the direct harm that fishing activities can have on coral reefs through damage caused by fishing gear, including nets, lines, and traps. Additionally, noticeable changes in marine life, such as the loss of herbivorous fish that offer valuable services to coral reefs, can reduce ecosystem function as a whole. [21] Another anthropogenic force that degrades coral reefs is bottom trawling; a fishing practice that scrapes coral reef habitats and other bottom substrate-dwelling organisms off the ocean floor. Bottom trawling results in physical wreckage and stress that leads to corals being broken and zooxanthellae expelled. Similar to bottom trawling, rock anchoring used for fishing can cause physical damage to these fragile reefs due to the heavy weight of the anchor, cables, and chains. [22] If coral reefs are exposed to physical damage like rock anchoring regularly, it can result in less resiliency to ocean acidification. Ecotourism is another anthropogenic factor that contributes to coral reef degradation. During ecotourism, humans can cause stress to the corals by accidentally touching, polluting, or breaking off parts of the reef, often resulting in coral bleaching as they attempt to fight off the intrusion. [23] However, ecotourism is not only harmful when humans are close enough to touch the coral. Less direct impacts, such as harmful chemicals in sunscreen and sedimentation driven by the tourism industry, can have irreversible effects as well. [23]

Managing coral reefs

In an attempt to prevent coral bleaching, scientists are experimenting by "seeding" corals that can host multiple types of zooxanthellae with thermally-resistant zooxanthellae. [1] MPAs have begun to apply reef resilience management techniques in order to improve the 'immune system' of coral reefs and promote reef recovery after bleaching. [3]

The Nature Conservancy has developed, and is continually refining, a model to help manage and promote reef resilience. Although this model does not guarantee reef resilience, it is a comprehensible management model to follow. The principles outlined in their model are: [3]

Scientists have also developed a new technique by Smithsonian’s National Zoo and Conservation Biology Institute and funded by a conservation organization called Revive and Restore. This technique is referred to as cryopreservation and involves freezing and thawing entire fragments of coral, resulting in slowing the loss of coral species and restoring damaged reefs. Previous coral cryopreservation techniques relied on largely freezing sperm and larvae, making collection difficult, as spawning events only occur a few days a year. This previous technique was also difficult because frequent marine heatwaves and warm waters can cause corals to be biologically stressed, resulting in their reproductive material being too weak to be frozen or thawed. The new technique is easier and works more rapidly, as it allows researchers and preservations to work throughout the year, rather than waiting for a certain species to spawn and put stress on coral's reproductive materials. [24] Scientists have also looked deeper into energy reserves and coral feeding. Feeding on zooplankton, brine shrimp, and algae may serve as a buffer for the harsh effects of climate change. Feeding corals can help them sustain tissue biomass and energy reserves and enhance nitrogen content, allowing for a higher zooxanthellae concentration and increased photosynthesis. [25] [26] Increased feeding rates can also allow certain species of bleached and recovering coral to exceed their daily metabolic energy requirements. These results suggest that coral species with a high CHAR (percent contribution of heterotrophically acquired carbon to daily animal respiration) capability may be more resilient to bleaching events, becoming the dominant species, and helping to safeguard affected reefs from extinction. [27]

Related Research Articles

<span class="mw-page-title-main">Coral</span> Marine invertebrates of the class Anthozoa

Corals are colonial marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.

<span class="mw-page-title-main">Coral reef</span> Outcrop of rock in the sea formed by the growth and deposit of stony coral skeletons

A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in groups.

<span class="mw-page-title-main">Coral bleaching</span> Phenomenon where coral expel algae tissue

Coral bleaching is the process when corals become white due to various stressors, such as changes in temperature, light, or nutrients. Bleaching occurs when coral polyps expel the zooxanthellae that live inside their tissue, causing the coral to turn white. The zooxanthellae are photosynthetic, and as the water temperature rises, they begin to produce reactive oxygen species. This is toxic to the coral, so the coral expels the zooxanthellae. Since the zooxanthellae produce the majority of coral colouration, the coral tissue becomes transparent, revealing the coral skeleton made of calcium carbonate. Most bleached corals appear bright white, but some are blue, yellow, or pink due to pigment proteins in the coral.

<span class="mw-page-title-main">Zooxanthellae</span> Dinoflagellates in symbiosis with coral, jellyfish and nudibranchs

Zooxanthellae is a colloquial term for single-celled dinoflagellates that are able to live in symbiosis with diverse marine invertebrates including demosponges, corals, jellyfish, and nudibranchs. Most known zooxanthellae are in the genus Symbiodinium, but some are known from the genus Amphidinium, and other taxa, as yet unidentified, may have similar endosymbiont affinities. The true Zooxanthella K.brandt is a mutualist of the radiolarian Collozoum inerme and systematically placed in Peridiniales. Another group of unicellular eukaryotes that partake in similar endosymbiotic relationships in both marine and freshwater habitats are green algae zoochlorellae.

<span class="mw-page-title-main">Southeast Asian coral reefs</span> Marine ecosystem

Southeast Asian coral reefs have the highest levels of biodiversity for the world's marine ecosystems. They serve many functions, such as forming the livelihood for subsistence fishermen and even function as jewelry and construction materials. Corals inhabit coastal waters off of every continent except Antarctica, with an abundance of reefs residing along Southeast Asian coastline in several countries including Indonesia, the Philippines, and Thailand. Coral reefs are developed by the carbonate-based skeletons of a variety of animals and algae. Slowly and overtime, the reefs build up to the surface in oceans. Coral reefs are found in shallow, warm salt water. The sunlight filters through clear water and allows microscopic organisms to live and reproduce. Coral reefs are actually composed of tiny, fragile animals known as coral polyps. Coral reefs are significantly important because of the biodiversity. Although the number of fish are decreasing, the remaining coral reefs contain more unique sea creatures. The variety of species living on a coral reef is greater than anywhere else in the world. An estimation of 70-90% of fish caught are dependent on coral reefs in Southeast Asia and reefs support over 25% of all known marine species. However, those sensitive coral reefs are facing detrimental effects on them due to variety of factors: overfishing, sedimentation and pollution, bleaching, and even tourist-related damage.

<span class="mw-page-title-main">White band disease</span> Disease affecting marine corals

White band disease is a coral disease that affects acroporid corals and is distinguishable by the white band of exposed coral skeleton that it forms. The disease completely destroys the coral tissue of Caribbean acroporid corals, specifically elkhorn coral and staghorn coral. The disease exhibits a pronounced division between the remaining coral tissue and the exposed coral skeleton. These symptoms are similar to white plague, except that white band disease is only found on acroporid corals, and white plague has not been found on any acroporid corals. It is part of a class of similar disease known as "white syndromes", many of which may be linked to species of Vibrio bacteria. While the pathogen for this disease has not been identified, Vibrio carchariae may be one of its factors. The degradation of coral tissue usually begins at the base of the coral, working its way up to the branch tips, but it can begin in the middle of a branch.

<span class="mw-page-title-main">Elkhorn coral</span> Species of coral

Elkhorn coral is an important reef-building coral in the Caribbean. The species has a complex structure with many branches which resemble that of elk antlers; hence, the common name. The branching structure creates habitat and shelter for many other reef species. Elkhorn coral is known to grow quickly with an average growth rate of 5 to 10 cm per year. They can reproduce both sexually and asexually, though asexual reproduction is much more common and occurs through a process called fragmentation.

<i>Symbiodinium</i> Genus of dinoflagellates (algae)

Symbiodinium is a genus of dinoflagellates that encompasses the largest and most prevalent group of endosymbiotic dinoflagellates known and have photosymbiotic relationships with many species. These unicellular microalgae commonly reside in the endoderm of tropical cnidarians such as corals, sea anemones, and jellyfish, where the products of their photosynthetic processing are exchanged in the host for inorganic molecules. They are also harbored by various species of demosponges, flatworms, mollusks such as the giant clams, foraminifera (soritids), and some ciliates. Generally, these dinoflagellates enter the host cell through phagocytosis, persist as intracellular symbionts, reproduce, and disperse to the environment. The exception is in most mollusks, where these symbionts are intercellular. Cnidarians that are associated with Symbiodinium occur mostly in warm oligotrophic (nutrient-poor), marine environments where they are often the dominant constituents of benthic communities. These dinoflagellates are therefore among the most abundant eukaryotic microbes found in coral reef ecosystems.

<span class="mw-page-title-main">Coral Triangle</span> Ecoregion of Asia

The Coral Triangle (CT) is a roughly triangular area in the tropical waters around the Philippines, Indonesia, Malaysia, Papua New Guinea, the Solomon Islands and Timor-Leste. This area contains at least 500 species of reef-building corals in each ecoregion. The Coral Triangle is located between the Pacific and Indian oceans and encompasses portions of two biogeographic regions: the Indonesian-Philippines Region, and the Far Southwestern Pacific Region. As one of eight major coral reef zones in the world, the Coral Triangle is recognized as a global centre of marine biodiversity and a global priority for conservation. Its biological resources make it a global hotspot of marine biodiversity. Known as the "Amazon of the seas", it covers 5.7 million square kilometres (2,200,000 sq mi) of ocean waters. It contains more than 76% of the world's shallow-water reef-building coral species, 37% of its reef fish species, 50% of its razor clam species, six out of seven of the world's sea turtle species, and the world's largest mangrove forest. In 2014, the Asian Development Bank (ADB) reported that the gross domestic product of the marine ecosystem in the Coral Triangle is roughly $1.2 trillion per year and provides food to over 120 million people. According to the Coral Triangle Knowledge Network, the region annually brings in about $3 billion in foreign exchange income from fisheries exports, and another $3 billion from coastal tourism revenues.

<span class="mw-page-title-main">Coral reef protection</span> Modifying human activities to reduce impact on coral reefs.

Coral reef protection is the process of modifying human activities to avoid damage to healthy coral reefs and to help damaged reefs recover. The key strategies used in reef protection include defining measurable goals and introducing active management and community involvement to reduce stressors that damage reef health. One management technique is to create Marine Protected Areas (MPAs) that directly limit human activities such as fishing.

<span class="mw-page-title-main">Environmental issues with coral reefs</span> Factors which adversely affect tropical coral reefs

Human activities have substantial impact on coral reefs, contributing to their worldwide decline.[1] Damaging activities encompass coral mining, pollution, overfishing, blast fishing, as well as the excavation of canals and access points to islands and bays. Additional threats comprise disease, destructive fishing practices, and the warming of oceans.[2] Furthermore, the ocean's function as a carbon dioxide sink, alterations in the atmosphere, ultraviolet light, ocean acidification, viral infections, the repercussions of dust storms transporting agents to distant reefs, pollutants, and algal blooms represent some of the factors exerting influence on coral reefs. Importantly, the jeopardy faced by coral reefs extends far beyond coastal regions. The ramifications of climate change, notably global warming, induce an elevation in ocean temperatures that triggers coral bleaching—a potentially lethal phenomenon for coral ecosystems.

<span class="mw-page-title-main">Aquaculture of coral</span> Cultivation of coral for commercial purposes

Coral aquaculture, also known as coral farming or coral gardening, is the cultivation of corals for commercial purposes or coral reef restoration. Aquaculture is showing promise as a tool for restoring coral reefs, which are dying off around the world. The process protects young corals while they are most at risk of dying. Small corals are propagated in nurseries and then replanted on the reef.

<i>Pocillopora verrucosa</i> Species of coral

Pocillopora verrucosa, commonly known as cauliflower coral, rasp coral, or knob-horned coral, is a species of stony coral in the family Pocilloporidae. It is native to tropical and subtropical parts of the Indian and Pacific Oceans.

<span class="mw-page-title-main">Mesophotic coral reef</span>

A Mesophotic coral reef or mesophotic coral ecosystem (MCE), originally from the Latin word meso (meaning middle) and photic (meaning light), is characterised by the presence of both light-dependent coral and algae, and organisms that can be found in water with low light penetration. Mesophotic Coral Ecosystem (MCEs) is a new, widely-adopted term used to refer to mesophotic coral reefs, as opposed to other similar terms like "deep coral reef communities" and "twilight zone", since those terms sometimes are confused due to their unclear, interchangeable nature.

<span class="mw-page-title-main">Ocean acidification in the Great Barrier Reef</span> Threat to the reef which reduces the viability and strength of reef-building corals

Ocean acidification threatens the Great Barrier Reef by reducing the viability and strength of coral reefs. The Great Barrier Reef, considered one of the seven natural wonders of the world and a biodiversity hotspot, is located in Australia. Similar to other coral reefs, it is experiencing degradation due to ocean acidification. Ocean acidification results from a rise in atmospheric carbon dioxide, which is taken up by the ocean. This process can increase sea surface temperature, decrease aragonite, and lower the pH of the ocean. The more humanity consumes fossil fuels, the more the ocean absorbs released CO₂, furthering ocean acidification.

Coral diseases are transmissible pathogens that cause the degradation of coral colonies. Coral cover in reef ecosystems has decreased significantly for a diverse set of reasons, ranging from variable environmental conditions to mechanical breakdowns from storms. In recent years, diseases that infect and kill coral have shown to be a threat to the health of coral reefs. Since the first coral disease was reported in 1965, many different kinds of diseases have popped up in mostly Caribbean waters. These diseases are diverse, including pathogens of bacteria, fungi, viruses, and protozoans. Coral diseases have widespread implications, impacting entire ecosystems and communities of organisms. Researchers are working to understand these diseases, and how potential treatments could stop these pathogens from causing the widespread death of corals in a way that permanently impacts the community structure of reefs.

<span class="mw-page-title-main">Human impact on marine life</span>

Human activities affect marine life and marine habitats through overfishing, habitat loss, the introduction of invasive species, ocean pollution, ocean acidification and ocean warming. These impact marine ecosystems and food webs and may result in consequences as yet unrecognised for the biodiversity and continuation of marine life forms.

<span class="mw-page-title-main">Coral reefs of Jamaica</span>

Jamaica, an island located within the Caribbean Sea, known for being a popular tourist destination because of its pristine white sand beaches, is now faced with the issue of mass coral depletion. Both environmental and human factors contribute to the destruction of these corals, which inevitably affect Jamaica's environmental sustainability and economy. Actions have been put in place to counteract the negative consequences associated with the loss of the corals, which act as a symbol of hope for the revival of Jamaica's environment.

<span class="mw-page-title-main">Marine microbiome</span>

All animals on Earth form associations with microorganisms, including protists, bacteria, archaea, fungi, and viruses. In the ocean, animal–microbial relationships were historically explored in single host–symbiont systems. However, new explorations into the diversity of marine microorganisms associating with diverse marine animal hosts is moving the field into studies that address interactions between the animal host and a more multi-member microbiome. The potential for microbiomes to influence the health, physiology, behavior, and ecology of marine animals could alter current understandings of how marine animals adapt to change, and especially the growing climate-related and anthropogenic-induced changes already impacting the ocean environment.

<span class="mw-page-title-main">Coral reef restoration</span> Restoration of coral reefs

Coral reef restoration strategies use natural and anthropogenic processes to restore damaged coral reefs. Reefs suffer damage from a number of natural and man-made causes, and efforts are being made to rectify the damage and restore the reefs. This involves the fragmentation of mature corals, the placing of the living fragments on lines or frames, the nurturing of the fragments as they recover and grow, and the transplantation of the pieces into their final positions on the reef when they are large enough.

References

  1. 1 2 Coral reef conservation program: Addressing key threats NOAA. Retrieved 7 December 2011.
  2. Holling, C.S. (1973) "Resilience and stability of ecological systems" Annual Review of Ecology and Systematics, 4: 1–23.
  3. 1 2 3 Reef resilience toolkit model: Introduction The Nature Conservancy Retrieved 7 December 2011.
  4. Fujise, L., Yamashita, H., Suzuki, G., Sasaki, K., Liao, L.M., Koike, K. (2014) Moderate thermal stress causes active and immediate expulsion of photosynthetically damaged zooxanthellae (Symbiodinium) from corals PLoS ONE,9(12): 1-18.
  5. Sampayo, E.M., Ridgway, T., Bongaerts, P., Hoegh-Guldberg, O. (2008) "Bleaching susceptibility and mortality of corals are determined by fine-scale differences in symbiont type" PNAS Environmental Sciences,105 (30): 10444–10449.
  6. 1 2 3 Berkelmans, R. and M.J.H. van Oppen (2006) "The role of zooxanthellae in the thermal tolerance of corals: a 'nugget of hope' for coral reefs in an era of climate change" Proceedings of the Royal Society of London Series B, 273: 2305–2312
  7. Palomares ML, Pauly D, eds. (2011). "Oculina patagonica" in SeaLifeBase. December 2011 version.
  8. 1 2 3 Fine, Maoz, Loya, Yossi (2002) "Endolithic algae: an alternative source of photoassimilates during coral bleaching" Proceedings of the Royal Society,269 (1497): 1205–1210.
  9. Roff, George; Mumby, Peter J. (2012-07-01). "Global disparity in the resilience of coral reefs". Trends in Ecology & Evolution. 27 (7): 404–413. doi:10.1016/j.tree.2012.04.007. ISSN   0169-5347. PMID   22658876.
  10. US Department of Commerce, National Oceanic and Atmospheric Administration. "What is coral bleaching?". oceanservice.noaa.gov. Retrieved 2020-08-25.
  11. "Zooxanthellae and Coral Bleaching | Smithsonian Ocean". ocean.si.edu. Retrieved 2020-08-25.
  12. Pratchett, Morgan S.; Hoey, Andrew S.; Wilson, Shaun K.; Messmer, Vanessa; Graham, Nicholas A. J. (September 2011). "Changes in Biodiversity and Functioning of Reef Fish Assemblages following Coral Bleaching and Coral Loss". Diversity. 3 (3): 424–452. doi: 10.3390/d3030424 . hdl: 10754/334624 .
  13. Curran, Sara R.; Agardy, Tundi (June 2002). "Common Property Systems, Migration, and Coastal Ecosystems". Ambio: A Journal of the Human Environment. 31 (4): 303–305. Bibcode:2002Ambio..31..303C. doi:10.1579/0044-7447-31.4.303. ISSN   0044-7447. PMID   12174600. S2CID   24074727.
  14. Sisney, Marsha A.; Cummins, R. Hays; Wolfe, Christopher R. (2018-12-15). "Incidence of black band disease, brown band disease, and white syndrome in branching corals on the Great Barrier Reef". Estuarine, Coastal and Shelf Science. 214: 1–9. Bibcode:2018ECSS..214....1S. doi:10.1016/j.ecss.2018.09.005. ISSN   0272-7714. S2CID   133646016.
  15. Sharma, Diksha; Ravindran, Chinnarajan (2020-06-01). "Diseases and pathogens of marine invertebrate corals in Indian reefs". Journal of Invertebrate Pathology. 173: 107373. doi:10.1016/j.jip.2020.107373. ISSN   0022-2011. PMID   32272136. S2CID   215726552.
  16. Harmelin-Vivien, Mireille L. (1994). "The Effects of Storms and Cyclones on Coral Reefs: A Review". Journal of Coastal Research: 211–231. ISSN   0749-0208. JSTOR   25735600.
  17. Brace, Claire (2018). "Climate Change Below the Surface: The Impact of Ocean Acidification on Reef Corals". Reinvention. 11 (2): 1 via Academic Search Complete.
  18. US EPA, OW (2017-01-30). "Threats to Coral Reefs". US EPA. Retrieved 2020-08-25.
  19. US Department of Commerce, National Oceanic and Atmospheric Administration. "Anthropogenic Threats to Corals - Corals: NOAA's National Ocean Service Education". oceanservice.noaa.gov. Retrieved 2020-08-25.
  20. Cinner, Joshua (2014-04-01). "Coral reef livelihoods". Current Opinion in Environmental Sustainability. Environmental change issues. 7: 65–71. Bibcode:2014COES....7...65C. doi:10.1016/j.cosust.2013.11.025. ISSN   1877-3435.
  21. McLean, Matthew; Cuetos-Bueno, Javier; Nedlic, Osamu; Luckymiss, Marston; Houk, Peter (2016-11-30). "Local Stressors, Resilience, and Shifting Baselines on Coral Reefs". PLOS ONE. 11 (11): e0166319. Bibcode:2016PLoSO..1166319M. doi: 10.1371/journal.pone.0166319 . ISSN   1932-6203. PMC   5130202 . PMID   27902715.
  22. Maynard, Jeffrey A.; Anthony, Kenneth R. N.; Afatta, Siham; Dahl-Tacconi, Nancy; Hoegh-Guldberg, Ove (2010-03-19). "Making a Model Meaningful to Coral Reef Managers in a Developing Nation: a Case Study of Overfishing and Rock Anchoring in Indonesia: Overfishing and Rock Anchoring in Indonesia". Conservation Biology. 24 (5): 1316–1326. doi:10.1111/j.1523-1739.2010.01487.x. PMID   20337685. S2CID   11512144.
  23. 1 2 Cossio, Camila (2016-04-18). "Coral Reefs and the Unintended Impact of Tourism". Earthjustice. Retrieved 2023-10-25.
  24. "New Technique Could Facilitate Rapid Cryopreservation of All Coral Species". Smithsonian's National Zoo. 2023-08-23. Retrieved 2023-10-25.
  25. Grottoli, Andréa G.; Martins, Paula Dalcin; Wilkins, Michael J.; Johnston, Michael D.; Warner, Mark E.; Cai, Wei-Jun; Melman, Todd F.; Hoadley, Kenneth D.; Pettay, D. Tye; Levas, Stephen; Schoepf, Verena (2018-01-16). "Coral physiology and microbiome dynamics under combined warming and ocean acidification". PLOS ONE. 13 (1): e0191156. Bibcode:2018PLoSO..1391156G. doi: 10.1371/journal.pone.0191156 . ISSN   1932-6203. PMC   5770069 . PMID   29338021.
  26. Grottoli, Andréa G. (2002-06-01). "Effect of light and brine shrimp on skeletal δ13C in the Hawaiian coral Porites compressa: a tank experiment". Geochimica et Cosmochimica Acta. 66 (11): 1955–1967. Bibcode:2002GeCoA..66.1955G. doi:10.1016/S0016-7037(01)00901-2. ISSN   0016-7037.
  27. Grottoli, Andréa G.; Rodrigues, Lisa J.; Palardy, James E. (April 2006). "Heterotrophic plasticity and resilience in bleached corals". Nature. 440 (7088): 1186–1189. Bibcode:2006Natur.440.1186G. doi:10.1038/nature04565. ISSN   1476-4687. PMID   16641995. S2CID   4422247.

Further references