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A coral outcrop on the Great Barrier Reef, Australia




Other corals do not rely on zooxanthellae and can live in much deeper water, with the cold-water genus Lophelia surviving as deep as 3,300 metres (10,800 ft).[1] Some have been found on the Darwin Mounds, northwest of Cape Wrath, Scotland, and others as far north as off the coast of Washington State and the Aleutian Islands.



Aristotle's pupil Theophrastus described the red coral, korallion, in his book on stones, implying it was a mineral, but he described it as a deep-sea plant in his Enquiries on Plants, where he also mentions large stony plants that reveal bright flowers when under water in the Gulf of Heroes.[2] Pliny the Elder stated boldly that several sea creatures including sea nettles and sponges "are neither animals nor plants, but are possessed of a third nature (tertius natura)".[3] Petrus Gyllius copied Pliny, introducing the term zoophyta for this third group in his 1535 book On the French and Latin Names of the Fishes of the Marseilles Region; it is popularly but wrongly supposed that Aristotle created the term.[3] Gyllius further noted, following Aristotle, how hard it was to define what was a plant and what was an animal.[3]


Presently, corals are classified as certain species of animals within the sub-classes Hexacorallia and Octocorallia of the class Anthozoa in the phylum Cnidaria.[6] Hexacorallia includes the stony corals and these groups have polyps that generally have a 6-fold symmetry. Octocorallia includes blue coral and soft corals and species of Octocorallia have polyps with an eightfold symmetry, each polyp having eight tentacles and eight mesenteries.

Fire corals are not true corals, being in the order Anthomedusae (sometimes known as Anthoathecata) of the class Hydrozoa.[7]


Anatomy of a stony coral polyp


Soft coralsEdit


Stony coralsEdit

Montastraea cavernosa polyps with tentacles extended


Coral skeletons are biocomposites (mineral + organics) Ca carbonate, in the form of calcite or aragonite. In scleractinian corals, "centers of calcification" and fibers are clearly distinct structures differing with respect to both morphology and chemical compositions of the crystalline units.[12] [13][1] [2] The organic matrices extracted from diverse species are acidic, and comprize proteins, sulphated sugars and lipids; they are species specific. [14] The soluble organic matrices of the skeletons allow to differentiate zooxanthellae and non zooxanthellae specimens.[15]





Intracellular symbiontsEdit

Many corals, as well as other cnidarian groups such as Aiptasia (a sea anemone) form a symbiotic relationship with a class of dinoflagellate algae, zooxanthellae of the genus Symbiodinium.[18]:24 Aiptasia, a familiar pest among coral reef aquarium hobbyists, serves as a valuable model organism in the study of cnidarian-algal symbiosis.[19] Typically, each polyp harbors one species of alga, and coral species show a preference for Symbiodinium.[20] Young corals are not born with zooxanthellae, but acquire the algae from the surrounding environment, including the water column and local sediment.[21] Via photosynthesis, these provide energy for the coral, and aid in calcification.[22] The main benefit of the zooxanthellae is their ability to photosynthesize. By using this technique, zooxanthellae are able to supply corals with the products of photosynthesis, including glucose, glycerol, and amino acids, which the corals can use for energy.[23] As much as 30% of the tissue of a polyp may be algal material.[18]:23 Zooxanthellae also benefit corals by aiding in waste removal.[24]

The algae benefit from a safe place to live and consume the polyp's carbon dioxide and nitrogenous waste. Due to the strain the algae can put on the polyp, stress on the coral often drives them to eject the algae. Mass ejections are known as coral bleaching, because the algae contribute to coral's brown coloration; other colors, however, are due to host coral pigments, such as green fluorescent proteins (GFPs). Ejection increases the polyp's chance of surviving short-term stress—they can regain algae, possibly of a different species at a later time. If the stressful conditions persist, the polyp eventually dies.[25] Zooxanthellae are located within the corals' cytoplasm and due to the algae's photosynthetic activity, the internal pH of the coral can be raised; this behavior indicates that the zooxanthellae are responsible to some extent for the metabolism of their host corals [26]


Corals can be both gonochoristic (unisexual) and hermaphroditic, each of which can reproduce sexually and asexually. Reproduction also allows coral to settle in new areas. Reproduction is coordinated by chemical communication.


Life cycles of broadcasters and brooders





Synchronous spawning is very typical on the coral reef, and often, even when multiple species are present, all corals spawn on the same night. This synchrony is essential so male and female gametes can meet. Corals rely on environmental cues, varying from species to species, to determine the proper time to release gametes into the water. The cues involve temperature change, lunar cycle, day length, and possibly chemical signalling.[27] Synchronous spawning may form hybrids and is perhaps involved in coral speciation.[29] The immediate cue is most often sunset, which cues the release.[27] The spawning event can be visually dramatic, clouding the usually clear water with gametes.


Brooding species are most often ahermatypic (not reef-building) in areas of high current or wave action. Brooders release only sperm, which is negatively buoyant, sinking on to the waiting egg carriers who harbor unfertilized eggs for weeks. Synchronous spawning events sometimes occur even with these species.[27] After fertilization, the corals release planula that are ready to settle.[22]


Planula larvae exhibit positive phototaxis, swimming towards light to reach surface waters, where they drift and grow before descending to seek a hard surface to which they can attach and begin a new colony. They also exhibit positive sonotaxis, moving towards sounds that emanate from the reef and away from open water.[30] High failure rates afflict many stages of this process, and even though millions of gametes are released by each colony, few new colonies form. The time from spawning to settling is usually two to three days, but can be up to two months.[31] The larva grows into a polyp and eventually becomes a coral head by asexual budding and growth.


Tabulate coral Aulopora (Devonian) showing initial budding


Budding involves splitting a smaller polyp from an adult.[28] As the new polyp grows, it forms its body parts. The distance between the new and adult polyps grows, and with it, the coenosarc (the common body of the colony). Budding can be intratentacular, from its oral discs, producing same-sized polyps within the ring of tentacles, or extratentacular, from its base, producing a smaller polyp.


Asexual reproduction offers the benefits of high reproductive rate, delaying senescence, and replacement of dead modules, as well as geographical distribution.[32]

Colony divisionEdit



Locations of coral reefs around the world




Evolutionary historyEdit


Although corals first appeared in the Cambrian period,[37] some 535 million years ago, fossils are extremely rare until the Ordovician period, 100 million years later, when rugose and tabulate corals became widespread. Paleozoic corals often contained numerous endobiotic symbionts.[38][39]



The scleractinian corals filled the niche vacated by the extinct rugose and tabulate species. Their fossils may be found in small numbers in rocks from the Triassic period, and became common in the Jurassic and later periods.[42] Scleractinian skeletons are composed of a form of calcium carbonate known as aragonite.[43] Although they are geologically younger than the tabulate and rugose corals, the aragonite of their skeletons is less readily preserved, and their fossil record is accordingly less complete.

RugosaScleractiniaTabulataEdiacaranCambrianCambrianOrdovicianOrdovicianSilurianSilurianDevonianDevonianCarboniferousCarboniferousPermianPermianTriassicTriassicJurassicCretaceousTertiaryPrecambrianPaleozoicMesozoicCenozoicPermian-Triassic extinctionLate Devonian extinctionCothoniidamya (unit) 

Timeline of the major coral fossil record and developments from 650 m.y.a. to present.[44][45]





Coral reefs are under stress around the world.[47] In particular, coral mining, agricultural and urban runoff, pollution (organic and inorganic), overfishing, blast fishing, disease, and the digging of canals and access into islands and bays are localized threats to coral ecosystems. Broader threats are sea temperature rise, sea level rise and pH changes from ocean acidification, all associated with greenhouse gas emissions.[48] In 1998, 16% of the world's reefs died as a result of increased water temperature.[49]

Approximately 10% of the world's coral reefs are dead.[50][51][52] About 60% of the world's reefs are at risk due to human-related activities.[53] The threat to reef health is particularly strong in Southeast Asia, where 80% of reefs are endangered.[54] Over 50% of the world's coral reefs may be destroyed by 2030; as a result, most nations protect them through environmental laws.[55]

In the Caribbean and tropical Pacific, direct contact between ~40–70% of common seaweeds and coral causes bleaching and death to the coral via transfer of lipid-soluble metabolites.[56] Seaweed and algae proliferate given adequate nutrients and limited grazing by herbivores such as parrotfish.


Submarine springs found along the coast of Mexico's Yucatán Peninsula produce water with a naturally low pH (relatively high acidity) providing conditions similar to those expected to become widespread as the oceans absorb carbon dioxide.[58] Surveys discovered multiple species of live coral that appeared to tolerate the acidity. The colonies were small and patchily distributed, and had not formed structurally complex reefs such as those that compose the nearby Mesoamerican Barrier Reef System.[58]


Marine Protected Areas, Biosphere reserves, marine parks, national monuments world heritage status, fishery management and habitat protection can protect reefs from anthropogenic damage.[59]


To eliminate destruction of corals in their indigenous regions, projects have been started to grow corals in non-tropical countries.[60][61]

Relation to humansEdit




Corals' many colors give it appeal for necklaces and other jewelry. Intensely red coral is prized as a gemstone. Sometimes called fire coral, it is not the same as fire coral. Red coral is very rare because of overharvesting.[63] In general, it is inadvisable to give coral as gifts since they are in decline from stressors like climate change, pollution, and unsustainable fishing.

Always considered a precious mineral, "the Chinese have long associated red coral with auspiciousness and longevity because of its color and its resemblance to deer antlers (so by association, virtue, long life, and high rank".[64] It reached its height of popularity during the Manchu or Qing Dynasty (1644-1911) when it was almost exclusively reserved for the emperor's use either in the form of coral beads (often combined with pearls) for court jewelry or as decorative Penjing (decorative miniature mineral trees). Coral was known as shanhu in Chinese. The "early-modern 'coral network' [began in] the Mediterranean Sea [and found its way] to Qing China via the English East India Company".[65] There were strict rules regarding its use in a code established by the Qianlong Emperor in 1759.


Depiction of coral in the Juliana Anicia Codex, a 6th-century copy of Dioscorides' De Materia Medica. The facing page states that coral can be used to treat ulcers.[66]

In medicine, chemical compounds from corals are used to treat cancer, AIDS and pain, and for other uses.[citation needed] Coral skeletons, e.g. Isididae are also used for bone grafting in humans.[67] Coral Calx, known as Praval Bhasma in Sanskrit, is widely used in traditional system of Indian medicine as a supplement in the treatment of a variety of bone metabolic disorders associated with calcium deficiency.[68] In classical times ingestion of pulverized coral, which consists mainly of the weak base calcium carbonate, was recommended for calming stomach ulcers by Galen and Dioscorides.[69]




Shoreline protectionEdit

Healthy coral reefs absorb 97 percent of a wave’s energy, which buffers shorelines from currents, waves, and storms, helping to prevent loss of life and property damage. Coastlines protected by coral reefs are also more stable in terms of erosion than those without.[72]

Local EconomiesEdit

Coastal communities near coral reefs rely heavily on them. Worldwide, more than 500 million people depend on coral reefs for food, income, coastal protection, and more.[73] The total economic value of coral reef services in the United States - including fisheries, tourism, and coastal protection - is more than $3.4 billion a year.

Climate researchEdit

Annual growth bands in some corals, such as the deep sea bamboo corals (Isididae), may be among the first signs of the effects of ocean acidification on marine life.[74] The growth rings allow geologists to construct year-by-year chronologies, a form of incremental dating, which underlie high-resolution records of past climatic and environmental changes using geochemical techniques.[75]



Though coral have large sexually-reproducing populations, their evolution can be slowed by abundant asexual reproduction.[78] Gene flow is variable among coral species.[78] According to the biogeography of coral species gene flow cannot be counted on as a dependable source of adaptation as they are very stationary organisms. Also, coral longevity might factor into their adaptivity.[78]

However, adaptation to climate change has been demonstrated in many cases. These are usually due to a shift in coral and zooxanthellae genotypes. These shifts in allele frequency have progressed toward more tolerant types of zooxanthellae.[79] Scientists found that a certain scleractinian zooxanthella is becoming more common where sea temperature is high.[80][81] Symbionts able to tolerate warmer water seem to photosynthesise more slowly, implying an evolutionary trade-off.[81]

In the Gulf of Mexico, where sea temperatures are rising, cold-sensitive staghorn and elkhorn coral have shifted in location.[79] Not only have the symbionts and specific species been shown to shift, but there seems to be a certain growth rate favorable to selection. Slower-growing but more heat-tolerant corals have become more common.[82] The changes in temperature and acclimation are complex. Some reefs in current shadows represent a refugium location that will help them adjust to the disparity in the environment even if eventually the temperatures may rise more quickly there than in other locations.[83] This separation of populations by climatic barriers causes a realized niche to shrink greatly in comparison to the old fundamental niche.



Strontium/calcium ratio anomalyEdit


Oxygen isotope anomalyEdit


Sea surface temperature and sea surface salinityEdit
Global sea surface temperature (SST)



Limited climate research on current speciesEdit
Porites lutea




The saltwater fishkeeping hobby has expanded, over recent years, to include reef tanks, fish tanks that include large amounts of live rock on which coral is allowed to grow and spread.[89] These tanks are either kept in a natural-like state, with algae (sometimes in the form of an algae scrubber) and a deep sand bed providing filtration,[90] or as "show tanks", with the rock kept largely bare of the algae and microfauna that would normally populate it,[91] in order to appear neat and clean.

The most popular kind of coral kept is soft coral, especially zoanthids and mushroom corals, which are especially easy to grow and propagate in a wide variety of conditions, because they originate in enclosed parts of reefs where water conditions vary and lighting may be less reliable and direct.[92] More serious fishkeepers may keep small polyp stony coral, which is from open, brightly lit reef conditions and therefore much more demanding, while large polyp stony coral is a sort of compromise between the two.


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 potentially effective tool for restoring coral reefs, which have been declining around the world.[93][94][95] The process bypasses the early growth stages of corals when they are most at risk of dying. Coral fragments known as "seeds" are grown in nurseries then replanted on the reef.[96] Coral is farmed by coral farmers who live locally to the reefs and farm for reef conservation or for income. It is also farmed by scientists for research, by businesses for the supply of the live and ornamental coral trade and by private aquarium hobbyists.


Further images: commons:Category:Coral reefs and commons:Category:Corals

See alsoEdit


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