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

A coral "group" is a colony of very many polyps. Each polyp is a sac-like animal typically only a few millimeters in diameter and a few centimeters in height. A set of surround a central mouth opening. Each polyp excretes an near the base. Over many generations, the colony thus creates a skeleton characteristic of the species which can measure up to several meters in size. Individual colonies grow by asexual reproduction of polyps. Corals also breed sexually by : polyps of the same species release simultaneously overnight, often around a . Fertilized eggs form planulae, a mobile early form of the coral polyp which, when mature, settles to form a new colony.

Although some corals are able to catch and small using on their tentacles, most corals obtain the majority of their energy and nutrients from of the genus that live within their tissues. These are commonly known as and give the coral color. Such corals require sunlight and grow in clear, shallow water, typically at depths less than , but corals in the genus have been found as deep as . Corals are major contributors to the physical structure of the that develop in tropical and subtropical waters, such as the Great Barrier Reef off the coast of . These corals are increasingly at risk of events where polyps expel the zooxanthellae in response to stress such as high water temperature or toxins.

Other corals do not rely on zooxanthellae and can live globally in much deeper water, such as the cold-water which can survive as deep as . Some have been found as far north as the , northwest of , Scotland, and others off the coast of and the .


Taxonomy
The classification of corals has been discussed for millennia, owing to having similarities to both plants and animals. 's pupil described the , 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 .
(2025). 9781408836224, Bloomsbury.
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 ( tertia natura)". 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. Gyllius further noted, following Aristotle, how hard it was to define what was a plant and what was an animal.
(2025). 9783319074795, Springer. .
The Babylonian Talmud refers to coral among a list of types of trees, and the 11th-century French commentator describes it as "a type of tree (מין עץ) that grows underwater that goes by the (French) name 'coral'."Babylonian Talmud, Rosh Hashana 23a, and commentary of Rashi (24th narrow line)

The Persian polymath (d.1048) classified sponges and corals as animals, arguing that they respond to touch.

(2025). 9780520953635, University of California Press.
Nevertheless, people believed corals to be plants until the eighteenth century when used a microscope to establish that coral had the characteristic thin cell membranes of an .
(2020). 9781474771009, Capstone Global Library Ltd.

Presently, corals are classified as species of animals within the sub-classes and of the class in the . Hexacorallia includes the stony corals and these groups have polyps that generally have a 6-fold symmetry. Octocorallia includes and and species of Octocorallia have polyps with an eightfold symmetry, each polyp having eight tentacles and eight mesenteries. The group of corals is because the are also in the sub-class Hexacorallia.


Systematics
The delineation of coral species is challenging as hypotheses based on morphological traits contradict hypotheses formed via molecular tree-based processes. As of 2020, there are 2175 identified separate coral species, 237 of which are currently endangered, making distinguishing corals to be the utmost of importance in efforts to curb extinction.  and delineation continues to occur in species of coral in order to combat the dangers posed by the climate crisis. Corals are colonial modular organisms formed by asexually produced and genetically identical modules called polyps. Polyps are connected by living tissue to produce the full organism. The living tissue allows for inter module communication (interaction between each polyp), which appears in colony morphologies produced by corals, and is one of the main identifying characteristics for a species of coral.

There are two main classifications for corals: hard coral (scleractinian and stony coral) which form reefs by a calcium carbonate base, with polyps that bear six stiff tentacles, and soft coral (Alcyonacea and ahermatypic coral) which are pliable and formed by a colony of polyps with eight feather-like tentacles. These two classifications arose from differentiation in gene expressions in their branch tips and bases that arose through developmental signaling pathways such as , Hedgehog, Wnt, and BMP.

Scientists typically select Acropora as research models since they are the most diverse genus of hard coral, having over 120 species. Most species within this genus have polyps which are dimorphic:

(2025). 9781869775841, Magnolia Press.
axial polyps grow rapidly and have lighter coloration, while radial polyps are small and are darker in coloration.
(2025). 9780878932436, Sinauer Associates.
In the Acropora genus, and occur at the basal polyps, growth occurs mainly at the radial polyps. Growth at the site of the radial polyps encompasses two processes: asexual reproduction via mitotic cell proliferation, and skeleton deposition of the calcium carbonate via extracellular matrix (ECM) proteins acting as differentially expressed (DE) signaling genes between both branch tips and bases. These processes lead to colony differentiation, which is the most accurate distinguisher between coral species. In the Acropora genus, colony differentiation through up-regulation and down-regulation of DEs.

Systematic studies of soft coral species have faced challenges due to a lack of knowledge. Researchers have not found enough variability within the genus to confidently delineate similar species, due to a low rate in mutation of mitochondrial DNA.

Environmental factors, such as the rise of temperatures and acid levels in our oceans account for some of corals in the form of . Various coral species have heat shock proteins (HSP) that are also in the category of DE across species. These HSPs help corals combat the increased temperatures they are facing which lead to protein denaturing, growth loss, and eventually coral death. Approximately 33% of coral species are on the International Union for Conservation of Nature's endangered species list and at risk of species loss. Ocean acidification (falling pH levels in the oceans) is threatening the continued species growth and differentiation of corals. Mutation rates of shilonii, the reef responsible for , heavily outweigh the typical reproduction rates of coral colonies when pH levels fall. Thus, corals are unable to mutate their HSPs and other climate change preventative genes to combat the increase in temperature and decrease in pH at a competitive rate to these pathogens responsible for coral bleaching, resulting in species loss.


Anatomy
For most of their life corals are sessile animals of colonies of genetically identical polyps. Each polyp varies from millimeters to centimeters in diameter, and colonies can be formed from many millions of individual polyps. Stony coral, also known as hard coral, polyps produce a skeleton composed of calcium carbonate to strengthen and protect the organism. This is deposited by the polyps and by the , the living tissue that connects them. The polyps sit in cup-shaped depressions in the skeleton known as . Colonies of stony coral are markedly variable in appearance; a single species may adopt an encrusting, plate-like, bushy, columnar or massive solid structure, the various forms often being linked to different types of habitat, with variations in light level and water movement being significant.
(2025). 9788131501047, Cengage Learning.

The body of the polyp may be roughly compared in a structure to a , the wall of which is composed of two layers of cells. The outer layer is known technically as the , the inner layer as the . Between ectoderm and endoderm is a supporting layer of gelatinous substance termed , secreted by the cell layers of the body wall. The mesoglea can contain elements derived from cells from the ectoderm.

The sac-like body built up in this way is attached to a hard surface, which in hard corals are cup-shaped depressions in the skeleton known as . At the center of the upper end of the sac lies the only opening called the mouth, surrounded by a circle of which resemble glove fingers. The tentacles are organs which serve both for tactile sense and for the capture of food. Polyps extend their tentacles, particularly at night, often containing coiled stinging cells () which pierce, poison and firmly hold living prey paralyzing or killing them. Polyp prey includes plankton such as and fish larvae. Longitudinal muscular fibers formed from the cells of the ectoderm allow tentacles to contract to convey the food to the mouth. Similarly, circularly disposed muscular fibers formed from the endoderm permit tentacles to be protracted or thrust out once they are contracted. In both stony and soft corals, the polyps can be retracted by contracting muscle fibers, with stony corals relying on their hard skeleton and cnidocytes for defense. Soft corals generally secrete toxins to ward off predators.

In most corals, the tentacles are retracted by day and spread out at night to catch plankton and other small organisms. Shallow-water species of both stony and soft corals can be , the corals supplementing their plankton diet with the products of photosynthesis produced by these . The polyps interconnect by a complex and well-developed system of canals, allowing significant sharing of nutrients and symbionts.

The external form of the polyp varies greatly. The column may be long and slender, or may be so short in the axial direction that the body becomes disk-like. The tentacles may number many hundreds or may be very few, in rare cases only one or two. They may be simple and unbranched, or feathery in pattern. The mouth may be level with the surface of the peristome, or may be projecting and trumpet-shaped.


Soft corals
Soft corals have no solid exoskeleton as such. However, their tissues are often reinforced by small supportive elements known as made of calcium carbonate. The polyps of soft corals have eight-fold symmetry, which is reflected in the Octo in Octocorallia.

Soft corals vary considerably in form, and most are colonial. A few soft corals are , but the polyps of most are connected by sheets of tissue called coenosarc, and in some species these sheets are thick and the polyps deeply embedded in them. Some soft corals encrust other sea objects or form lobes. Others are tree-like or whip-like and have a central axial skeleton embedded at their base in the matrix of the supporting branch. These branches are composed of a fibrous protein called or of a calcified material.


Stony corals
The polyps of stony corals have six-fold symmetry. In stony corals, the tentacles are cylindrical and taper to a point, but in soft corals they are pinnate with side branches known as pinnules. In some tropical species, these are reduced to mere stubs and in some, they are fused to give a paddle-like appearance.
(1999). 9781883693091, Ricordea Publishing.

Coral skeletons are biocomposites (mineral + organics) of calcium 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. The organic matrices extracted from diverse species are acidic, and comprise proteins, sulphated sugars and lipids; they are species specific. The soluble organic matrices of the skeletons allow to differentiate and non-zooxanthellae specimens.


Ecology

Feeding
Polyps feed on a variety of small organisms, from microscopic zooplankton to small fish. The polyp's tentacles immobilize or kill prey using stinging cells called , commonly called nematocysts. These cells carry which they rapidly release in response to contact with another organism. A dormant nematocyst discharges in response to nearby prey touching the trigger. A stiff flap called an operculum opens and its stinging apparatus fires the barb into the prey. The venom is injected through the hollow filament to immobilise the prey; the tentacles then manoeuvre the prey into the stomach. Once the prey is digested the stomach reopens allowing the elimination of waste products and the beginning of the next hunting cycle.


Intracellular symbionts
Many corals, as well as other groups such as form a relationship with a class of , of the genus , which can form as much as 30% of the tissue of a polyp.
(2025). 9780878501380, The Darwin Press.
Typically, each polyp harbors one species of alga, and coral species show a preference for . Young corals are not born with zooxanthellae, but acquire the algae from the surrounding environment, including the water column and local sediment. The main benefit of the zooxanthellae is their ability to photosynthesize which supplies corals with the products of photosynthesis, including glucose, glycerol, also amino acids, which the corals can use for energy. Zooxanthellae also benefit corals by aiding in , for the coral skeleton, and waste removal.
(1986). 9789401083706, Geo Books.
In addition to the soft tissue, are also found in the coral's mucus and (in stony corals) the skeleton, with the latter showing the greatest microbial richness.

The zooxanthellae benefit from a safe place to live and consume the polyp's , phosphate and nitrogenous waste. Stressed corals will eject their zooxanthellae, a process that is becoming increasingly common due to strain placed on coral by rising ocean temperatures. Mass ejections are known as because the algae contribute to coral coloration; some 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 and if the stress subsides they can regain algae, possibly of a different species, at a later time. If the stressful conditions persist, the polyp eventually dies. Zooxanthellae are located within the coral 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. Stony Coral Tissue Loss Disease has been associated with the breakdown of host-zooxanthellae physiology.Landsberg et al., "Stony Coral Tissue Loss Disease in Florida Is Associated With Disruption of Host–Zooxanthellae Physiology". Moreover, Vibrio bacterium are known to have virulence traits used for host coral tissue damage and photoinhibition of algal symbionts.de O Santos et al., "Genomic and Proteomic Analyses of the Coral Pathogen Vibrio Coralliilyticus Reveal a Diverse Virulence Repertoire". Therefore, both coral and their symbiotic microorganisms could have evolved to harbour traits resistant to disease and transmission.


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


Sexual
Corals predominantly reproduce sexually. About 25% of (reef-building stony corals) form single-sex () colonies, while the rest are . It is estimated more than 67% of coral are simultaneous hermaphrodites.
(2011). 9780231527156, Columbia University Press. .


Broadcasters
About 75% of all hermatypic corals "broadcast spawn" by releasing —eggs and —into the water where they meet and fertilize to spread offspring. Corals often synchronize their time of spawning. This reproductive synchrony is essential so that male and female gametes can meet. Spawning frequently takes place in the evening or at night, and can occur as infrequently as once a year, and within a window of 10–30 minutes. Synchronous spawning is very typical on the coral reef, and often, all corals spawn on the same night even when multiple are present. Synchronous spawning may form hybrids and is perhaps involved in coral .

Environmental cues that influence the release of gametes into the water vary from species to species. The cues involve temperature change, , , and possibly chemical signalling.

(2025). 9780642322364, Australian Institute of Marine Sciences and CRR Qld.
Other factors that affect the rhythmicity of organisms in marine habitats include salinity, mechanical forces, and pressure or magnetic field changes.

Mass coral spawning often occurs at night on days following a full moon. A full moon is equivalent to four to six hours of continuous dim light exposure, which can cause light-dependent reactions in protein. Corals contain light-sensitive , proteins whose light-absorbing flavin structures are sensitive to different types of light. This allows corals such as Dipsastraea speciosa to detect and respond to changes in sunlight and moonlight.

Moonlight itself may actually suppress coral spawning. The most immediate cue to cause spawning appears to be the dark portion of the night between sunset and moonrise. Over the lunar cycle, moonrise shifts progressively later, occurring after sunset on the day of the full moon. The resulting dark period between day-light and night-light removes the suppressive effect of moonlight and enables coral to spawn.

The spawning event can be visually dramatic, clouding the usually clear water with gametes. Once released, gametes fertilize at the water's surface and form a microscopic called a , typically pink and elliptical in shape. A typical coral colony needs to release several thousand larvae per year to overcome the odds against formation of a new colony.

(1999). 9780865428348, Blackwell.

Studies suggest that light pollution desynchronizes spawning in some coral species. In areas such as the , as many as 10 out of 50 species may be showing spawning asynchrony, compared to 30 years ago. The establishment of new corals in the area has decreased and in some cases ceased. The area was previously considered a refuge for corals because mass bleaching events due to climate change had not been observed there. Coral restoration techniques for coral reef management are being developed to increase fertilization rates, larval development, and settlement of new corals.


Brooders
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 onto the waiting egg carriers that harbor unfertilized eggs for weeks. Synchronous spawning events sometimes occur even with these species. After fertilization, the corals release planula that are ready to settle.


Planulae
The time from spawning to larval settlement is usually two to three days but can occur immediately or up to two months.
(1973). 9780123896025, Harcourt Brace Jovanovich.
Broadcast-spawned larvae develop at the water's surface before descending to seek a hard surface on the benthos to which they can attach and begin a new colony. The larvae often need a biological cue to induce settlement such as specific crustose species or microbial biofilms. High failure rates afflict many stages of this process, and even though thousands of eggs are released by each colony, few new colonies form. During settlement, larvae are inhibited by physical barriers such as sediment, as well as chemical (allelopathic) barriers. The larvae metamorphose into a single polyp and eventually develops into a juvenile and then adult by asexual budding and growth.


Asexual
Within a coral head, the genetically identical polyps reproduce asexually, either by (gemmation) or by dividing, whether longitudinally or transversely.

Budding involves splitting a smaller polyp from an adult. As the new polyp grows, it forms . 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.

Division forms two polyps that each become as large as the original. Longitudinal division begins when a polyp broadens and then divides its coelenteron (body), effectively splitting along its length. The mouth divides and new tentacles form. The two polyps thus created then generate their missing body parts and exoskeleton. Transversal division occurs when polyps and the exoskeleton divide transversally into two parts. This means one has the basal disc (bottom) and the other has the oral disc (top); the new polyps must separately generate the missing pieces.

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

(1998). 9781566472210, Mutual Publishing.


Colony division
Whole colonies can reproduce asexually, forming two colonies with the same genotype. The possible mechanisms include fission, bailout and fragmentation. Fission occurs in some corals, especially among the family , where the colony splits into two or more colonies during early developmental stages. Bailout occurs when a single polyp abandons the colony and settles on a different substrate to create a new colony. Fragmentation involves individuals broken from the colony during storms or other disruptions. The separated individuals can start new colonies.
(2009). 9780191057342, OUP Oxford. .


Coral microbiomes
Corals are one of the more common examples of an animal host whose symbiosis with microalgae can turn to , and is visibly detected as bleaching. Coral have been examined in a variety of studies, which demonstrate how oceanic environmental variations, most notably temperature, light, and inorganic nutrients, affect the abundance and performance of the microalgal symbionts, as well as and physiology of the host.Dubinsky, Z. and Jokiel, P.L. (1994) "Ratio of energy and nutrient fluxes regulates symbiosis between zooxanthellae and corals". Pacific Science, 48(3): 313–324.Anthony, K.R., Kline, D.I., Diaz-Pulido, G., Dove, S. and Hoegh-Guldberg, O.(2008) "Ocean acidification causes bleaching and productivity loss in coral reef builders". Proceedings of the National Academy of Sciences, 105(45): 17442–17446. .Apprill, A. (2017) "Marine animal microbiomes: toward understanding host–microbiome interactions in a changing ocean". Frontiers in Marine Science, 4: 222. . Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.

Studies have also suggested that resident bacteria, archaea, and fungi additionally contribute to nutrient and organic matter cycling within the coral, with viruses also possibly playing a role in structuring the composition of these members, thus providing one of the first glimpses at a multi-domain marine animal symbiosis.Bourne, D.G., Morrow, K.M. and Webster, N.S. (2016) "Insights into the coral microbiome: underpinning the health and resilience of reef ecosystems". Annual Review of Microbiology, 70: 317–340. . The gammaproteobacterium is emerging as a central member of the coral's microbiome, with flexibility in its lifestyle.Neave, M.J., Apprill, A., Ferrier-Pagès, C. and Voolstra, C.R. (2016) "Diversity and function of prevalent symbiotic marine bacteria in the genus Endozoicomonas". Applied Microbiology and Biotechnology, 100(19): 8315–8324. .Neave, M.J., Michell, C.T., Apprill, A. and Voolstra, C.R. (2017) "Endozoicomonas genomes reveal functional adaptation and plasticity in bacterial strains symbiotically associated with diverse marine hosts". Scientific Reports, 7: 40579. . Given the recent mass bleaching occurring on reefs,Hughes, T.P., Kerry, J.T., Álvarez-Noriega, M., Álvarez-Romero, J.G., Anderson, K.D., Baird, A.H., Babcock, R.C., Beger, M., Bellwood, D.R., Berkelmans, R. and Bridge, T.C. (2017) "Global warming and recurrent mass bleaching of corals". Nature, 543(7645): 373–377. . corals will likely continue to be a useful and popular system for symbiosis and dysbiosis research.

Astrangia poculata, the northern star coral, is a temperate , widely documented along the eastern coast of the United States. The coral can live with and without (algal symbionts), making it an ideal to study microbial community interactions associated with symbiotic state. However, the ability to develop primers and to more specifically target key microbial groups has been hindered by the lack of full-length 16S rRNA sequences, since sequences produced by the Illumina platform are of insufficient length (approximately 250 base pairs) for the design of primers and probes. USGS scientists publish long-read microbiome sequences from temperate coral, providing community resource for probe and primer design, United States Geological Survey, 6 March 2019. Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License. In 2019, Goldsmith et al. demonstrated Sanger sequencing was capable of reproducing the biologically relevant diversity detected by deeper next-generation sequencing, while also producing longer sequences useful to the research community for probe and primer design (see diagram on right). Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.


Holobionts
Reef-building corals are well-studied that include the coral itself together with its symbiont (photosynthetic dinoflagellates), as well as its associated bacteria and viruses.Knowlton, N. and Rohwer, F. (2003) "Multispecies microbial mutualisms on coral reefs: the host as a habitat". The American Naturalist, 162(S4): S51-S62. . Co-evolutionary patterns exist for coral microbial communities and coral phylogeny.

It is known that the coral's and symbiont influence host health, however, the historic influence of each member on others is not well understood. Scleractinian corals have been diversifying for longer than many other symbiotic systems, and their microbiomes are known to be partially species-specific. It has been suggested that , a commonly highly abundant bacterium in corals, has exhibited codiversification with its host. This hints at an intricate set of relationships between the members of the coral holobiont that have been developing as of these members occurs.

A study published in 2018 revealed evidence of between corals and their tissue and skeleton microbiomes. The coral skeleton, which represents the most diverse of the three coral microbiomes, showed the strongest evidence of phylosymbiosis. Coral microbiome composition and were found to reflect coral . For example, interactions between bacterial and eukaryotic coral phylogeny influence the abundance of Endozoicomonas, a highly abundant bacterium in the coral holobiont. However, host-microbial appears to influence only a subset of coral-associated bacteria.


Reefs
Many corals in the order are , meaning that they are involved in building reefs. Most such corals obtain some of their energy from in the genus Symbiodinium. These are photosynthetic which require sunlight; reef-forming corals are therefore found mainly in shallow water. They secrete calcium carbonate to form hard skeletons that become the framework of the reef. However, not all reef-building corals in shallow water contain zooxanthellae, and some deep water species, living at depths to which light cannot penetrate, form reefs but do not harbour the symbionts.

There are various types of shallow-water coral reef, including fringing reefs, barrier reefs and atolls; most occur in tropical and subtropical seas. They are very slow-growing, adding perhaps one centimetre (0.4 in) in height each year. The Great Barrier Reef is thought to have been laid down about two million years ago. Over time, corals fragment and die, sand and rubble accumulates between the corals, and the shells of clams and other molluscs decay to form a gradually evolving calcium carbonate structure. Coral reefs are extremely diverse marine hosting over 4,000 species of fish, massive numbers of cnidarians, , , and many other animals.

(2025). 9780520232556, University of California Press and UNEP/WCMC. .


Evolution
At certain times in the geological past, corals were very abundant. Like modern corals, their ancestors built reefs, some of which ended as great structures in sedimentary rocks. Fossils of fellow reef-dwellers algae, sponges, and the remains of many , , , , and appear along with coral fossils. This makes some corals useful . Coral fossils are not restricted to reef remnants, and many solitary fossils are found elsewhere, such as Cyclocyathus, which occurs in England's formation.


Early corals
Corals first appeared in the about .
(2025). 9780231106139, Columbia University Press.
are extremely rare until the period, 100 million years later, when Heliolitida, , and corals became widespread. corals often contained numerous endobiotic symbionts.

Tabulate corals occur in and calcareous of the Ordovician period, with a gap in the fossil record due to extinction events at the end of the Ordovician. Corals reappeared some millions of years later during the period, and tabulate corals often form low cushions or branching masses of alongside rugose corals. Tabulate coral numbers began to decline during the middle of the Silurian period.

Rugose or horn corals became dominant by the middle of the Silurian period, and during the Devonian, corals flourished with more than 200 genera. The rugose corals existed in solitary and colonial forms, and were also composed of calcite. Both rugose and tabulate corals became extinct in the Permian–Triassic extinction eventXiang-Dong Wang and Xiao-Juan Wang (2007). "Extinction patterns of Late Permian (Lopingian) corals in China", Palaeoworld, 16, No. 1–3, 31–38 (along with 85% of marine species), and there is a gap of tens of millions of years until new forms of coral evolved in the .

File:Syringoporid.jpg| Tabulate coral (a syringoporid); Boone limestone (Lower ) near Hiwasse, Arkansas, scale bar is 2.0 cm File:AuloporaDevonianSilicaShale.jpg| Tabulate coral from the period File:RugosaOrdovician.jpg| Solitary rugose coral ( ) in three views; Ordovician, southeastern Indiana


Modern corals
The currently ubiquitous stony corals, , appeared in the to fill the niche vacated by the extinct rugose and tabulate orders and is not closely related to the earlier forms. Unlike the corals prevalent before the Permian extinction, which formed skeletons of a form of calcium carbonate known as , modern stony corals form skeletons composed of the . Their fossils are found in small numbers in rocks from the Triassic period, and become common in the and later periods. 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.


Status

Threats
Coral reefs are under stress around the world. In particular, coral mining, agricultural and , (organic and inorganic), , , disease, and the digging of 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. In 1998, 16% of the world's reefs died as a result of increased water temperature. Losing Our Coral Reefs – Eco Matters – State of the Planet. Blogs.ei.columbia.edu. Retrieved on 2011-11-01.

Approximately 10% of the world's coral reefs are dead.Save Our Seas, 1997 Summer Newsletter, Cindy Hunter and Alan Friedlander About 60% of the world's reefs are at risk due to human-related activities.

(2025). 9781569737620, World Resources Institute. .
The threat to reef health is particularly strong in , where 80% of reefs are . Over 50% of the world's may be destroyed by 2030; as a result, most nations protect them through environmental laws.

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 -soluble . Seaweed and algae proliferate given adequate and limited grazing by such as .

Water temperature changes of more than or changes can kill some species of coral. Under such environmental stresses, corals expel their ; without them, coral tissues reveal the white of their skeletons, an event known as .

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. 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.


Coral health
To assess the threat level of coral, scientists developed a coral imbalance ratio, Log (Average abundance of disease-associated taxa / Average abundance of healthy associated taxa). The lower the ratio the healthier the microbial community is. This ratio was developed after the microbial mucus of coral was collected and studied.


Climate change impacts
Increasing sea surface temperatures in tropical regions (~) the last century have caused major , death, and therefore shrinking coral populations. Although coral are able to adapt and acclimate, it is uncertain if this evolutionary process will happen quickly enough to prevent major reduction of their numbers. Climate change causes more frequent and more severe storms that can destroy .

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

Certain species form communities called , which are colonies whose top is dead and mostly above the water line, but whose perimeter is mostly submerged and alive. Average level limits their height. By analyzing the various growth morphologies, microatolls offer a low-resolution record of sea level change. Fossilized microatolls can also be dated using radiocarbon dating. Such methods can help to reconstruct .

Though coral have large sexually-reproducing populations, their evolution can be slowed by abundant asexual reproduction. is variable among coral species. According to the 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.

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

In the Gulf of Mexico, where sea temperatures are rising, cold-sensitive and have shifted in location. 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. 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. This by climatic barriers causes a to shrink greatly in comparison to the old fundamental niche.


Geochemistry
Corals are shallow, colonial organisms that integrate oxygen and trace elements into their skeletal (polymorph of ) crystalline structures as they grow. Geochemical anomalies within the crystalline structures of corals represent functions of temperature, salinity and oxygen isotopic composition. Such geochemical analysis can help with climate modeling. The ratio of oxygen-18 to oxygen-16 (δ18O), for example, is a proxy for temperature.


Strontium/calcium ratio anomaly
Time can be attributed to coral geochemistry anomalies by correlating / minimums with sea surface temperature (SST) maximums to data collected from NINO 3.4 SSTA.


Oxygen isotope anomaly
The comparison of coral strontium/calcium minimums with sea surface temperature maximums, data recorded from NINO 3.4 SSTA, time can be correlated to coral strontium/calcium and δ18O variations. To confirm the accuracy of the annual relationship between Sr/Ca and δ18O variations, a perceptible association to annual coral growth rings confirms the age conversion. is established by the blending of Sr/Ca data, growth rings, and stable isotope data. El Nino-Southern Oscillation (ENSO) is directly related to climate fluctuations that influence coral δ18O ratio from local salinity variations associated with the position of the South Pacific convergence zone (SPCZ) and can be used for modeling.


Sea surface temperature and sea surface salinity
The global moisture budget is primarily being influenced by tropical sea surface temperatures from the position of the Intertropical Convergence Zone (ITCZ). The Southern Hemisphere has a unique meteorological feature positioned in the southwestern Pacific Basin called the South Pacific Convergence Zone (SPCZ), which contains a perennial position within the Southern Hemisphere. During warm periods, the reverses orientation extending from the equator down south through , , and towards the French Polynesian Islands; and due east towards affecting geochemistry of corals in tropical regions.

Geochemical analysis of skeletal coral can be linked to sea surface salinity (SSS) and sea surface temperature (SST), from El Nino 3.4 SSTA data, of tropical oceans to seawater δ18O ratio anomalies from corals. phenomenon can be related to variations in sea surface salinity (SSS) and sea surface temperature (SST) that can help model tropical climate activities.


Limited climate research on current species
Climate research on live coral species is limited to a few studied species. Studying coral provides a stable foundation for geochemical interpretations that is much simpler to physically extract data in comparison to species where the complexity of species skeletal structure creates difficulty when physically sampled, which happens to be one of the only multidecadal living coral records used for coral modeling.


Protection
Marine Protected Areas, Biosphere reserves, , national monuments status, fishery management and habitat protection can protect reefs from anthropogenic damage.

Many governments now prohibit removal of coral from reefs, and inform coastal residents about reef protection and ecology. While local action such as habitat restoration and herbivore protection can reduce local damage, the longer-term threats of acidification, temperature change and sea-level rise remain a challenge.

Protecting networks of diverse and healthy reefs, not only climate refugia, helps ensure the greatest chance of genetic diversity, which is critical for coral to adapt to new climates. A variety of conservation methods applied across marine and terrestrial threatened ecosystems makes coral adaption more likely and effective.

To eliminate destruction of corals in their indigenous regions, projects have been started to grow corals in non-tropical countries. EcoDeco EcologicalTechnology . Ecodeco.nl. Retrieved on 2011-11-29. KoralenKAS project . Koraalwetenschap.nl. Retrieved on 2011-11-29.


Relation to humans
Local economies near major coral reefs benefit from an abundance of fish and other marine creatures as a food source. Reefs also provide recreational and tourism. These activities can damage coral but international projects such as that encourage dive and snorkel centres to follow a Code of Conduct have been proven to mitigate these risks.


Jewelry
Corals' many colors give it appeal for necklaces and other . Intensely red coral is prized as a gemstone. Sometimes called fire coral, it is not the same as . Red coral is very rare because of . 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".Welch, Patricia Bjaaland, Chinese Art: A Guide to Motifs and Visual Imagery. Tokyo, Rutland and Singapore: Tuttle, 2008, p. 61 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 (decorative miniature mineral trees). Coral was known as shanhu in Chinese. The "early-modern 'coral network' began the Mediterranean Sea and to Qing China via the English East India Company".Lacey, Pippa, "The Coral Network: The trade of red coral to the Qing imperial court in the eighteenth century" in The Global Lives of Things, ed. by Anne Gerritsen and Giorgio Aiello, London: Rutledge, 2016, p. 81 There were strict rules regarding its use in a code established by the in 1759.

By the middle of the 19th century, “coral fisheries” existed in the Mediterranean, the Red Sea, Persian Gulf and elsewhere. An instrument used to retrieve the coral consisted of two beams of heavy wood attached to each other at right angles. Heavy stones were added to make the apparatus sink and netting was attached beneath the device. It is then lowered by a strong rope over an outcrop of coral and the boat trawls over the coral causing it to break off and be caught in the netting. The device is then drawn to the surface by the boat crew.


Medicine
In medicine, chemical compounds from corals can potentially be used to treat cancer, neurological diseases, inflammation including arthritis, pain, bone loss, high blood pressure and for other therapeutic uses. Coral skeletons, e.g. are being researched for their potential near-future use for in humans. Coral Calx, known as Praval Bhasma in , is widely used in traditional system of as a supplement in the treatment of a variety of bone metabolic disorders associated with calcium deficiency. In classical times ingestion of pulverized coral, which consists mainly of the weak base calcium carbonate, was recommended for calming stomach ulcers by and .Pedanius Dioscorides – Der Wiener Dioskurides, Codex medicus Graecus 1 der Österreichischen Nationalbibliothek Graz: Akademische Druck- und Verlagsanstalt 1998 fol. 391 verso (Band 2), Kommentar S. 47 und 52.


Construction
Coral reefs in places such as the East African coast are used as a source of building material.
(2002). 9780521523097, Cambridge University Press. .
Ancient (fossil) coral limestone, notably including the Coral Rag Formation of the hills around (England), was once used as a building stone, and can be seen in some of the oldest buildings in that city including the Saxon tower of St Michael at the Northgate, St. George's Tower of , and the medieval walls of the city.


Shoreline protection
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.


Local economies
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. 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.


Aquaria
The saltwater fishkeeping hobby has expanded, over recent years, to include , fish tanks that include large amounts of on which coral is allowed to grow and spread. Aquarium Corals: Collection and Aquarium Husbandry of Northeast Pacific Non-Photosynthetic Cnidaria . Advancedaquarist.com (2011-01-14). Retrieved on 2016-06-13. These tanks are either kept in a natural-like state, with algae (sometimes in the form of an ) and a deep sand bed providing filtration, Reefkeeping 101 – Various Nutrient Control Methods. Reefkeeping.com. Retrieved on 2016-06-13. or as "show tanks", with the rock kept largely bare of the algae and that would normally populate it, Aquarium Substrate & Live Rock Clean Up Tips . Saltaquarium.about.com. Retrieved on 2016-06-13. in order to appear neat and clean.

The most popular kind of coral kept is , especially 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. Coral Reefs . Marinebio.org. Retrieved on 2016-06-13. More serious fishkeepers may keep small polyp , 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.


Aquaculture
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 , which have been declining around the world. 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. 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 hobbyists.


Gallery
Further images: and File:Mushroom Coral (Fungia) Top Macro 91.JPG| sp. skeleton File:Eusmilia fastigiata large.jpg|Polyps of Eusmilia fastigiata File:Dendrogyra cylindrus (pillar coral) (San Salvador Island, Bahamas) 1 (15513345363).jpg|, Dendrogyra cylindricus File:Brain coral.jpg|, Diploria labyrinthiformis File:Stony coral spawning 3.jpg|Brain coral releasing eggs File:EilatFringingReef.jpg|Fringing off the coast of , . File:000324-Corals-IMG 0418-2.jpg|Corals, Tis Beach, , File:000407-Corals-IMG 0693-2.jpg|Corals, Tis Beach, ,


See also


Sources


External links

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