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{{short description|Marine invertebrates of the class Anthozoa}}{{other uses}}File:Coral Outcrop Flynn Reef.jpg|thumb|upright=1.25|A coral outcrop on the Great Barrier Reef, AustraliaAustraliaCorals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically live in compact colonies of many identical individual polyps. Corals species include the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.A coral "group" is a colony of myriad genetically identical polyps. Each polyp is a sac-like animal typically only a few millimeters in diameter and a few centimeters in length. A set of tentacles surround a central mouth opening. An exoskeleton is excreted near the base. Over many generations, the colony thus creates a large skeleton characteristic of the species. Individual heads grow by asexual reproduction of polyps. Corals also breed sexually by spawning: polyps of the same species release gametes simultaneously over a period of one to several nights around a full moon.Although some corals are able to catch small fish and plankton using stinging cells on their tentacles, most corals obtain the majority of their energy and nutrients from photosynthetic unicellular dinoflagellates in the genus Symbiodinium that live within their tissues. These are commonly known as zooxanthellae. Such corals require sunlight and grow in clear, shallow water, typically at depths less than {{convert|60|m}}. Corals are major contributors to the physical structure of the coral reefs that develop in tropical and subtropical waters, such as the Great Barrier Reef off the coast of Queensland, 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 {{convert|3300|m}}.JOURNAL, Squires, D.F., 1959, Deep sea corals collected by the Lamont Geological Observatory. 1. Atlantic corals,weblink American Museum Novitates, 1965, 23, Some have been found as far north as the Darwin Mounds, northwest of Cape Wrath, Scotland, and others off the coast of Washington State and the Aleutian Islands.{{TOC limit|3}}


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.BOOK, Leroi, Armand Marie, Armand Marie Leroi, The Lagoon: How Aristotle Invented Science, Aristotle's Lagoon, Bloomsbury, 2014, 978-1-4088-3622-4, 271, 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)". 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. Gyllius further noted, following Aristotle, how hard it was to define what was a plant and what was an animal.BOOK, Bowen, James, The Coral Reef Era: From Discovery to Decline: A history of scientific investigation from 1600 to the Anthropocene Epoch,weblink 2015, Springer, 978-3-319-07479-5, 5–7, The Persian polymath Al-Biruni (d. 1048) classified sponges and corals as animals, arguing that they respond to touch.BOOK, Egerton, Frank N., Roots of Ecology: Antiquity to Haeckel, 2012, University of California Press, 978-0-520-95363-5, 24, Nevertheless, people believed corals to be plants until the eighteenth century, when William Herschel used a microscope to establish that coral had the characteristic thin cell membranes of an animal.The Light of Reason 8 August 2006 02:00 BBC Four{{better source needed|date=January 2018}}Presently, corals are classified as certain species of animals within the sub-classes Hexacorallia and Octocorallia of the class Anthozoa in the phylum Cnidaria.WORMS, Hoeksema, Bert, 2015, Anthozoa, 1292, 2015-04-24, 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.WORMS, Schuchert, Peter, 2015, Milleporidae Fleming, 1828, 196235, 2015-04-24,


(File:Coral polyp.jpg|thumb|Anatomy of a stony coral polyp)Corals are sessile animals and differ from most other cnidarians in not having a medusa stage in their life cycle. The body unit of the animal is a polyp. Most corals are colonial, the initial polyp budding to produce another and the colony gradually developing from this small start. In stony corals, also known as hard corals, the polyps produce a skeleton composed of calcium carbonate to strengthen and protect the organism. This is deposited by the polyps and by the coenosarc, the living tissue that connects them. The polyps sit in cup-shaped depressions in the skeleton known as corallites. Colonies of stony coral are very 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.BOOK, Invertebrate Zoology, 7th edition, Ruppert, Edward E., Fox, Richard, S., Barnes, Robert D., 2004, Cengage Learning, 978-81-315-0104-7, 132–48,

Soft corals

Soft corals have no solid exoskeleton per se. However, their tissues are often reinforced by small supportive elements known as "sclerites" made of calcium carbonate. Soft corals vary considerably in form, and most are colonial. A few soft corals are stolonate, but the polyps of most are connected by sheets of coenosarc [def], 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 chem a central axial skeleton embedded at its base in the matrix of the supporting branch.WEB,weblink existing and potential value of coral ecosystems with respect to income and other economic values, Administration, US Department of Commerce, National Oceanic and Atmospheric,, EN-US, 2018-02-04, These branches are composed either of a fibrous protein called gorgonin or of a calcified material. In both stony and soft corals, the polyps can be retracted, with stony corals relying on their hard skeleton and cnidocytes for defence. Soft corals generally secrete terpenoid toxins to ward off predators.

Stony corals

File:Montastrea cavernosa.jpg|thumb|Montastraea cavernosaMontastraea cavernosaThe polyps of stony corals have six-fold symmetry while those of soft corals have eight. The mouth of each polyp is surrounded by a ring of tentacles. In stony corals these 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.BOOK, Corals: A quick reference guide, Sprung, Julian, 1999, Ricordea Publishing, 978-1-883693-09-1, 145, 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 zooxanthellate, the corals supplementing their plankton diet with the products of photosynthesis produced by these symbionts. The polyps interconnect by a complex and well-developed system of gastrovascular canals, allowing significant sharing of nutrients and symbionts.JOURNAL, D. Gateno, A. Israel, Y. Barki, B. Rinkevich, Gastrovascular Circulation in an Octocoral: Evidence of Significant Transport of Coral and Symbiont Cells, The Biological Bulletin, 1998, 178–86, 194, 2,weblink 10.2307/1543048, 28570841, 1543048, 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.JOURNAL, Cuif, J.P., Dauphin, Y., 1998, Microstructural and physico-chemical characterization of 'centers of calcification' in septa of some Recent scleractinian corals, Paläontologische Zeitschrift, 72, 3–4, 257–269, 10.1007/bf02988357, 0031-0220, JOURNAL, Cuif, J.P., Dauphin, Y., Doucet, J., Salomé, M., Susini, J., 2003, XANES mapping of organic sulfate in three scleractinian coral skeletons, Geochimica et Cosmochimica Acta, 67, 1, 75–83, 10.1016/s0016-7037(02)01041-4, 0016-7037, The organic matrices extracted from diverse species are acidic, and comprise proteins, sulphated sugars and lipids; they are species specific.JOURNAL, Dauphin, Y., Cuif, J.P., Williams, C. T., 2008, Soluble organic matrices of aragonitic skeletons of Merulinidae (Cnidaria, Anthozoa), Comparative Biochemistry and Physiology Part B: Biochemistry and Molecular Biology, 150, 1, 10–22, 10.1016/j.cbpb.2008.01.002, 18325807, 1096-4959, The soluble organic matrices of the skeletons allow to differentiate zooxanthellae and non zooxanthellae specimens.JOURNAL, Cuif, J.P., Dauphin, Y., Freiwald, A., Gautret, P., Zibrowius, H., 1999, Biochemical markers of zooxanthellae symbiosis in soluble matrices of skeleton of 24 Scleractinia species, Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 123, 3, 269–278, 10.1016/s1095-6433(99)00059-8, 1095-6433,


(File:nematocyst discharge.png|left|thumb|Discharge mechanism of a stinging cell (nematocyst))


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 nematocysts. These cells carry venom which they rapidly release in response to contact with another organism. A dormant nematocyst discharges in response to nearby prey touching the trigger (cnidocil). A flap (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.{{rp|24}}

Intracellular symbionts

Many corals, as well as other cnidarian groups such as sea anemones form a symbiotic relationship with a class of dinoflagellate algae, zooxanthellae of the genus Symbiodinium, which can form as much as 30% of the tissue of a polyp.BOOK, Coral Reefs: Cities Under The Seas, Murphy, Richard C., 2002, 978-0-87850-138-0, The Darwin Press, {{rp|23-24}} Typically, each polyp harbors one species of alga, and coral species show a preference for Symbiodinium.JOURNAL, Yuyama, Ikuko, Comparing the Effects of Symbiotic Algae (Symbiodinium) Clades C1 and D on Early Growth Stages of Acropora tenuis, PLOS One, 2014, 9, 6, e98999, 10.1371/journal.pone.0098999, 24914677, 4051649, 2014PLoSO...998999Y, Young corals are not born with zooxanthellae, but acquire the algae from the surrounding environment, including the water column and local sediment.JOURNAL, Yamashita, Hiroshi, Establishment of Coral–Algal Symbiosis Requires Attraction and Selection, PLOS One, 2014, 9, 5, e97003, 10.1371/journal.pone.0097003, 24824794, 4019531, 2014PLoSO...997003Y, The main benefit of the zooxanthellae is their ability to photosynthesize which supplies corals with the products of photosynthesis, including glucose, glycerol, and amino acids, which the corals can use for energy.WEB, Zooxanthellae...What's That?,weblink NOAA Ocean Service Education, National Oceanic and Atmospheric Administration, 1 December 2017, Zooxanthellae also benefit corals by aiding in calcification, for the coral skeleton, and waste removal.WEB, Madl, P., Yip, M., 2000,weblink Field Excursion to Milne Bay Province – Papua New Guinea, 2006-03-31, BOOK, van de Plaasche, Orson, Sea-level research: a manual for the collection and evaluation of data, 1986, Geo Books, Norwich, UK, 978-94-010-8370-6, 196, In addition to the soft tissue, microbiomes are also found in the coral's mucus and (in stony corals) the skeleton, with the latter showing the greatest microbial richness.Corals and their microbiomes evolved together | Penn State UniversityThe zooxanthellae benefit from a safe place to live and consume the polyp's carbon dioxide, phosphate 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 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.JOURNAL, W. W. Toller, R. Rowan, N. Knowlton, Repopulation of Zooxanthellae in the Caribbean Corals Montastraea annularis and M. faveolata following Experimental and Disease-Associated Bleaching, The Biological Bulletin, 2001, 360–73, 201,weblink 10.2307/1543614, 11751248, 3, 1543614, 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 JOURNAL, Brownlee, Colin, pH regulation in symbiotic anemones and corals: A delicate balancing act, Proceedings of the National Academy of Sciences of the United States of America, 2009, 106, 39, 16541–16542, 10.1073/pnas.0909140106, 19805333, 2757837, 2009PNAS..10616541B,


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.


(File:Coral Life Cycles ZP.svg|thumb|upright=1.75|Life cycles of broadcasters and brooders)Corals predominantly reproduce sexually. About 25% of hermatypic corals (stony corals) form single sex (gonochoristic) colonies, while the rest are hermaphroditic.BOOK, Veron, J.E.N., 2000, Corals of the World. Vol 3, 3rd, Australian Institute of Marine Sciences and CRR Qld, Australia, 978-0-642-32236-4,


About 75% of all hermatypic corals "broadcast spawn" by releasing gametes—eggs and sperm—into the water to spread offspring. The gametes fuse during fertilization to form a microscopic larva called a planula, typically pink and elliptical in shape. A typical coral colony forms several thousand larvae per year to overcome the odds against formation of a new colony.BOOK, Barnes, R. and, R., Hughes, 1999, An Introduction to Marine Ecology, 3rd, 117–41, Blackwell, Malden, MA, 978-0-86542-834-8, File:Stony coral spawning 2.jpg|thumb|A male great star coralgreat star coralSynchronous 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. Synchronous spawning may form hybrids and is perhaps involved in coral speciation.JOURNAL, Hatta, M., Fukami, H., Wang, W., Omori, M., Shimoike, K., Hayashibara, T., Ina, Y., Sugiyama, T., Reproductive and genetic evidence for a reticulate evolutionary theory of mass spawning corals, Molecular Biology and Evolution, 1999, 1607–13, 16, 11,weblink 10555292, 10.1093/oxfordjournals.molbev.a026073, The immediate cue is most often sunset, which cues the release. 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. After fertilization, the corals release planula that are ready to settle.(File:Life Cycle of Corals.svg|thumb|left|Generalized life cycle of corals via sexual reproduction.)


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.JOURNAL, Vermeij, Mark J. A., Marhaver, Kristen L., Huijbers, Chantal M., Nagelkerken, Ivan, Simpson, Stephen D., 2010, Coral Larvae Move toward Reef Sounds, PLoS ONE, 5, 5, e10660, 20498831, 10.1371/journal.pone.0010660,weblink ScienceDaily, May 16, 2010, 2871043, 2010PLoSO...510660V, 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.BOOK, Jones, O.A., Endean, R., 1973, Biology and Geology of Coral Reefs, 205–45, Harcourt Brace Jovanovich, New York, USA, 978-0-12-389602-5, The larva grows into a polyp and eventually becomes a coral head by asexual budding and growth.


File:Orbicella annularis - calices.jpg|thumb|right|Basal plates (calices) of Orbicella annularisOrbicella annularisWithin a coral head, the genetically identical polyps reproduce asexually, either by budding (gemmation) or by dividing, whether longitudinally or transversely.Budding involves splitting a smaller polyp from an adult. As the new polyp grows, it forms
missing image!
- Coral polyp.jpg -
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.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.BOOK, Hawaiian Coral Reef Ecology, 1998, Gulko, David, Mutual Publishing, Honolulu, Hawaii, 978-1-56647-221-0, 10,

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 Fungiidae, 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.BOOK, Sheppard, Charles R.C., Davy, Simon K., Pilling, Graham M., The Biology of Coral Reefs,weblink 25 June 2009, OUP Oxford, 978-0-19-105734-2, 78–81,


{{see also|Coral reef fish|List of reefs}}(File:Coral reef locations.jpg|thumb|upright=1.5|Locations of coral reefs around the world)Many corals in the order Scleractinia are hermatypic, meaning that they are involved in building reefs. Most such corals obtain some of their energy from zooxanthellae in the genus Symbiodinium. These are symbiotic photosynthetic dinoflagellates 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.JOURNAL, Schuhmacher, Helmut, Zibrowius, Helmut, 1985, What is hermatypic?, Coral Reefs, 4, 1, 1–9, 10.1007/BF00302198, 1985CorRe...4....1S, File:Hertshoon.jpg|thumb|left|Staghorn coralStaghorn coralThere 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.ENCYCLOPEDIA, MSN Encarta, 2006, Great Barrier Reef,weblink April 25, 2015,weblink" title="">weblink October 28, 2009, dead, Coral reefs are extremely diverse marine ecosystems hosting over 4,000 species of fish, massive numbers of cnidarians, molluscs, crustaceans, and many other animals.BOOK, Spalding, Mark, Ravilious, Corinna, Green, Edmund, 2001, World Atlas of Coral Reefs, 205–45, University of California Press and UNEP/WCMC, Berkeley, CA, 978-0-520-23255-6,

Evolutionary history

(File:RugosaOrdovician.jpg|thumb|Solitary rugose coral (Grewingkia) in three views; Ordovician, southeastern Indiana)Although corals first appeared in the Cambrian period,BOOK, Pratt, B.R., Spincer, B.R., R.A., Wood, A.Yu., Zhuravlev, Ecology of the Cambrian Radiation, 2001,weblink 2007-04-06, Columbia University Press, 978-0-231-10613-9, 259, 12: Ecology and Evolution of Cambrian Reefs, {{dead link|date=January 2018 |bot=InternetArchiveBot |fix-attempted=yes }} some {{Ma|535}}, 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.JOURNAL, 10.1666/07-056.1, The earliest endosymbiotic mineralized tubeworms from the Silurian of Podolia, Ukraine, 2008, Vinn, O., Mõtus, M.-A., Journal of Paleontology, 82, 2, 409–14,weblink 2014-06-11, JOURNAL, Diverse early endobiotic coral symbiont assemblage from the Katian (Late Ordovician) of Baltica, 2012, Vinn, O., Mõtus, M.-A., Palaeogeography, Palaeoclimatology, Palaeoecology, 321–322, 137–41, 10.1016/j.palaeo.2012.01.028, File:Syringoporid.jpg|thumb|left|Tabulate coral (a syringoporid); Boone limestone (Lower CarboniferousCarboniferousFile:AuloporaDevonianSilicaShale.jpg|thumb|right|Tabulate coral Aulopora from the DevonianDevonianTabulate corals occur in limestones and calcareous shales of the Ordovician and Silurian periods, and often form low cushions or branching masses of calcite alongside rugose corals. Their numbers began to decline during the middle of the Silurian period, and they became extinct at the end of the Permian period, {{Ma|250}}.WEB, Introduction to the Tabulata,weblink UCMP Berkeley, 25 April 2015, Rugose or horn corals became dominant by the middle of the Silurian period, and became extinct early in the Triassic period. The rugose corals existed in solitary and colonial forms, and were also composed of calcite.WEB, Introduction to the Rugosa,weblink UCMP Berkeley, 25 April 2015, 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.WEB, Evolutionary history,weblink AIMS, 25 April 2015, Scleractinian skeletons are composed of a form of calcium carbonate known as aragonite.JOURNAL, Ries JB, Stanley SM, Hardie LA, July 2006, Scleractinian corals produce calcite, and grow more slowly, in artificial Cretaceous seawater, Geology, 34, 7, 525–28, 10.1130/G22600.1, 2006Geo....34..525R, 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.{{Coral fossil record timeline}}At certain times in the geological past, corals were very abundant. Like modern corals, these 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 echinoids, brachiopods, bivalves, gastropods, and trilobites appear along with coral fossils. This makes some corals useful index fossils.WEB, Alden, Andrew, Index Fossils,weblink About education, 25 April 2015, Coral fossils are not restricted to reef remnants, and many solitary fossils may be found elsewhere, such as Cyclocyathus, which occurs in England's Gault clay formation.



(File:Reef0484.jpg|thumb|upright|A healthy coral reef has a striking level of biodiversity in many forms of marine life.)Coral reefs are under stress around the world.NEWS,weblink Coral reefs around the world,, 2 September 2009, 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.WEB, Threats to Coral Reefs,weblink Coral Reef Alliance, 2010, 5 December 2011, dead,weblink" title="">weblink 1 December 2011, 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. Retrieved on 2011-11-01.Approximately 10% of the world's coral reefs are dead.JOURNAL, Kleypas, J.A., R.A., Feely, V.J., Fabry, C., Langdon, C.L., Sabine, L.L., Robbins, 2006, Impacts of Ocean Acidification on Coral Reefs and Other Marine Calcifiers: A guide for Future Research, National Science Foundation, NOAA, & United States Geological Survey,weblink April 7, 2011, dead,weblink" title="">weblink July 20, 2011, Save Our Seas, 1997 Summer Newsletter, Dr. Cindy Hunter and Dr. Alan FriedlanderBOOK, Status of Coral Reefs, Coral Reef Monitoring and Management in Southeast Asia, 2004, Tun, K., L.M., Chou, A., Cabanban, V.S., Tuan, S., Philreefs, T., Yeemin, Suharsono, K., Sour, D., Lane, 2004, 235–76, C., Wilkinson, Status of Coral Reefs of the world: 2004, Australian Institute of Marine Science, Townsville, Queensland, Australia, 2019-04-23,weblink About 60% of the world's reefs are at risk due to human-related activities.BOOK, Burke, Lauretta, Reefs at risk revisited, 2011, World Resources Institute, Washington, DC, 978-1-56973-762-0, 38, Reytar, K., Spalding, M., Perry, A., The threat to reef health is particularly strong in Southeast Asia, where 80% of reefs are endangered.WEB, Bryant, Dirk, Burke, Lauretta, McManus, John, Spalding, Mark, Reefs at Risk: A Map-Based Indicator of Threats to the World's Coral Reef,weblink NOAA, 25 April 2015,weblink" title="">weblink 2013-02-18, dead, Over 50% of the world's coral reefs may be destroyed by 2030; as a result, most nations protect them through environmental laws.JOURNAL, Norlander, Coral crisis! Humans are killing off these bustling underwater cities. Can coral reefs be saved? (Life science: corals), Science World, 8 December 2003,weblink 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.JOURNAL, Rasher DB, Hay ME, Chemically rich seaweeds poison corals when not controlled by herbivores, Proceedings of the National Academy of Sciences of the United States of America, 107, 21, 9683–88, May 2010, 20457927, 2906836, 10.1073/pnas.0912095107, 2010PNAS..107.9683R, Seaweed and algae proliferate given adequate nutrients and limited grazing by herbivores such as parrotfish.Water temperature changes of more than 1–2 Â°C (1.8–3.6 Â°F) or salinity changes can kill some species of coral. Under such environmental stresses, corals expel their Symbiodinium; without them coral tissues reveal the white of their skeletons, an event known as coral bleaching.JOURNAL, Hoegh-Guldberg, O., Climate change, coral bleaching and the future of the world's coral reefs, Marine and Freshwater Research, 1999, 839–66, 50, 8,weblink 10.1071/MF99078, Ove Hoegh-Guldberg (biologist), dead,weblink" title="">weblink 2012-04-26, 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.WEB, Stephens, Tim, Submarine springs offer preview of ocean acidification effects on coral reefs,weblink University of California Santa Cruz, 28 November 2011, 25 April 2015, 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.


Marine Protected Areas, Biosphere reserves, marine parks, national monuments world heritage status, fishery management and habitat protection can protect reefs from anthropogenic damage.MAGAZINE, Phoenix Rising, National Geographic Magazine, January 2011, April 30, 2011,weblink 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.To eliminate destruction of corals in their indigenous regions, projects have been started to grow corals in non-tropical countries.EcoDeco EcologicalTechnology {{webarchive|url= |date=2011-03-07 }}. Retrieved on 2011-11-29.KoralenKAS project {{webarchive|url= |date=2012-04-26 }}. 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 scuba diving and snorkeling tourism. These activities can damage coral but international projects such as Green Fins that encourage dive and snorkel centres to follow a Code of Conduct have been proven to mitigate these risks.JOURNAL, Hunt, Chloe V., Harvey, James J., Miller, Anne, Johnson, Vivienne, Phongsuwan, Niphon, 2013, The Green Fins approach for monitoring and promoting environmentally sustainable scuba diving operations in South East Asia, Ocean & Coastal Management, 78, 35–44, 10.1016/j.ocecoaman.2013.03.004,


File:6-Strand Necklace, Navajo (Native American), ca. 1920s, cropped.jpg|thumb|upright|6-strand necklace, Navajo (Native American), ca. 1920s, Brooklyn MuseumBrooklyn MuseumCorals' 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.NEWS, Coral makes a splash, Melissa, Magsaysay,weblink Los Angeles Times, June 21, 2009, January 12, 2013, 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 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".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 Qianlong Emperor in 1759.


missing image!
- ViennaDioscoridesCoral.jpg -
upright|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.Folio 391, Juliana Anicia Codex
In medicine, chemical compounds from corals can potentially be used to treat cancer, AIDS, pain, and for other therapeutic uses.JOURNAL, Copper, Edwin, Hirabayashi, K., Strychar, K. B., Sammarco, P. W., 2014, Corals and their Potential Applications to Integrative Medicine, Evidence-Based Complementary and Alternative Medicine, 9, ProQuest, 3976867, JOURNAL, Senthilkumar, Kalimuthu, Se-Kwon, Kim, 2013, Marine Invertebrate Natural Products for Anti-Inflammatory and Chronic Diseases., Evidence-Based Complementary and Alternative Medicine, ProQuest, 3893779, Coral skeletons, e.g. Isididae are also used for bone grafting in humans.JOURNAL, Biomaterial structure in deep‐sea bamboo coral (Anthozoa: Gorgonacea: Isididae): perspectives for the development of bone implants and templates for tissue engineering, Materialwissenschaft und Werkstofftechnik, 37, 6, 552–57, 10.1002/mawe.200600036, 2006, Ehrlich, H., Etnoyer, P., Litvinov, S. D., Olennikova, M.M., Domaschke, H., Hanke, T., Born, R., Meissner, H., Worch, H., 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.JOURNAL, Reddy PN, Lakshmana M, Udupa UV, Effect of Praval bhasma (Coral calx), a natural source of rich calcium on bone mineralization in rats, Pharmacological Research, 48, 6, 593–99, December 2003, 14527824, 10.1016/S1043-6618(03)00224-X, 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.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. {{ISBN|3-201-01725-6}}


Coral reefs in places such as the East African coast are used as a source of building material.BOOK, Pouwels, Randall L., Horn and Crescent: Cultural Change and Traditional Islam on the East African Coast, 800–1900,weblink 6 June 2002, Cambridge University Press, 978-0-521-52309-7, 26, Ancient (fossil) coral limestone, notably including the Coral Rag Formation of the hills around Oxford (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 Oxford Castle, and the medieval walls of the city.WEB, Strategic Stone Study: A Building Stone Atlas of Oxfordshire,weblink English Heritage, 23 April 2015, March 2011,

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.JOURNAL, Ferrario, F., Beck, M.W., Storlazzi, C.D., Micheli, F., Shepard, C.C., Airoldi, L., The effectiveness of coral reefs for coastal hazard risk reduction and adaptation, Nature Communications, 2014, 5, 3794, 3794, 10.1038/ncomms4794, 24825660, 4354160,

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.WEB,weblink Status of Coral Reefs of the World: 2004 Volume 1, Global Coral Reef Monitoring Network, 2019-01-14, 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 research

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.WEB,weblink National Oceanic and Atmospheric Administration – New Deep-Sea Coral Discovered on NOAA-Supported Mission,, 2009-05-11, 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.JOURNAL, Schrag, D.P., Linsley, B.K., Corals, chemistry, and climate, Science, 2002, 277–78, 296, 8, 11951026, 10.1126/science.1071561, Certain species form communities called microatolls, which are colonies whose top is dead and mostly above the water line, but whose perimeter is mostly submerged and alive. Average tide 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 Holocene sea levels.JOURNAL, Smithers, Scott G., Woodroffe, Colin D., 2000, Microatolls as sea-level indicators on a mid-ocean atoll, Marine Geology, 168, 1–4, 61–78, 10.1016/S0025-3227(00)00043-8, 2000MGeol.168...61S, Increasing sea temperatures in tropical regions (~1 degree C) the last century have caused major coral bleaching, death, and therefore shrinking coral populations since although they are able to adapt and acclimate, it is uncertain if this evolutionary process will happen quickly enough to prevent major reduction of their numbers.JOURNAL, Hoegh-Guldberg O., Climate change, coral bleaching and the future of the world's coral reefs, Marine and Freshwater Research, 1999, 839–99, 50, 8, 10.1071/mf99078, Though coral have large sexually-reproducing populations, their evolution can be slowed by abundant asexual reproduction.JOURNAL, Hughes, T., Baird, A., Bellwood, D., Card, M., Connolly, S., Folke, C., Grosberg, R., Hoegh-Guldberg, O., Jackson, J., Klepas, J., Lough, J., Marshall, P., Nystrom, M., Palumbi, S., Stephen Palumbi, Pandolfi, J., Rosen, B., and Roughgarden, J., Climate change, human impacts, and the resilience of coral reefs, Science, 2003, 929–33, 301, 5635, 10.1126/science.1085046, 12920289, 2003Sci...301..929H, Gene flow is variable among coral species. 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.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.JOURNAL, Parmesan, C., Ecological and evolutionary responses to recent climate change, Annual Review of Ecology, Evolution, and Systematics, 2006, 637–69, 37, 10.1146/annurev.ecolsys.37.091305.110100, Scientists found that a certain scleractinian zooxanthella is becoming more common where sea temperature is high.JOURNAL, Baker, A., Corals' adaptive response to climate change, Nature, 2004, 741, 430, 7001, 10.1038/430741a, 2004Natur.430..741B, JOURNAL, Donner, S., Skirving, W., Little, C., Oppenheimer, M., Hoegh-Guldberg, O., Global assessment of coral bleaching and required rates of adaptation under climate change, Global Change Biology, 2005, 2251–65, 11, 12, 10.1111/j.1365-2486.2005.01073.x,weblink 2005GCBio..11.2251D,, 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 staghorn and elkhorn coral 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.JOURNAL, Baskett, M., Gaines, S., Nisbet, R., yes, Symbiont diversity may help coral reefs survive moderate climate change, Ecological Applications, 2009, 3–17, 19, 1, 10.1890/08-0139.1, 19323170, 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.JOURNAL, McClanahan, T., Ateweberhan, M., Muhando, C., Maina, J., Mohammed, M., yes, Effects of Climate and Seawater Temperature Variation on Coral Bleaching and Morality, Ecological Monographs, 2007, 503–25, 77, 4, 10.1890/06-1182.1,, This separation of populations by climatic barriers causes a realized niche to shrink greatly in comparison to the old fundamental niche.


Corals are shallow, colonial organisms that integrate δ18O and trace elements into their skeletal aragonite (polymorph of calcite) crystalline structures, as they grow. Geochemistry anomalies within the crystalline structures of corals represent functions of temperature, salinity and oxygen isotopic composition. Such geochemical analysis can help with climate modeling.JOURNAL, Kilbourne, K. Halimeda, Quinn, Terrence M., Taylor, Frederick W., Delcroix, Thierry, Gouriou, Yves, 2004, El Niño-Southern Oscillation-related salinity variations recorded in the skeletal geochemistry of a Porites coral from Espiritu Santo, Vanuatu, Paleoceanography, 19, 4, PA4002, 10.1029/2004PA001033, 2004PalOc..19.4002K,

Strontium/calcium ratio anomaly

Time can be attributed to coral geochemistry anomalies by correlating strontium/calcium minimums with sea surface temperature (SST) maximums to data collected from NINO 3.4 SSTA.JOURNAL, Ren, Lei, Linsley, Braddock K., Wellington, Gerard M., Schrag, Daniel P., Hoegh-guldberg, Ove, 2003, Deconvolving the δ18O seawater component from subseasonal coral δ18O and Sr/Ca at Rarotonga in the southwestern subtropical Pacific for the period 1726 to 1997, Geochimica et Cosmochimica Acta, 67, 9, 1609–21, 10.1016/S0016-7037(02)00917-1, 2003GeCoA..67.1609R,

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 accuracy of the annual relationship between Sr/Ca and δ18O variations, a perceptible association to annual coral growth rings confirms the age conversion. Geochronology 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 ENSO modeling.

Sea surface temperature and sea surface salinity

(File:Global Sea Surface Temperature - GPN-2003-00032.jpg|thumb|Global sea surface temperature (SST))The global moisture budget is primarily being influenced by tropical sea surface temperatures from the position of the Intertropical Convergence Zone (ITCZ).JOURNAL, Wu, Henry C., Linsley, Braddock K., Dassié, Emilie P., Schiraldi, Benedetto, deMenocal, Peter B., 2013, Oceanographic variability in the South Pacific Convergence Zone region over the last 210 years from multi-site coral Sr/Ca records, Geochemistry, Geophysics, Geosystems, 14, 5, 1435–53, 10.1029/2012GC004293, 2013GGG....14.1435W, 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 ENSO warm periods, the SPCZ reverses orientation extending from the equator down south through Solomon Islands, Vanuatu, Fiji and towards the French Polynesian Islands; and due east towards South America affecting geochemistry of corals in tropical regions.JOURNAL, Kiladis, George N., von Storch, Hans, van Loon, Harry, 1989, Origin of the South Pacific Convergence Zone, Journal of Climate, 2, 10, 1185–95, 10.1175/1520-0442(1989)0022.0.CO;2, 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. ENSO phenomenon can be related to variations in sea surface salinity (SSS) and sea surface temperature (SST) that can help model tropical climate activities.JOURNAL, Lukas, Roger, Lindstrom, Eric, 1991, The mixed layer of the western equatorial Pacific Ocean, Journal of Geophysical Research, 96, S1, 3343–58, 10.1029/90JC01951, 1991JGR....96.3343L,

Limited climate research on current species

(File:Porites lutea.jpg|thumb|Porites lutea)Climate research on live coral species is limited to a few studied species. Studying Porites coral provides a stable foundation for geochemical interpretations that is much simpler to physically extract data in comparison to Platygyra species where the complexity of Platygyra species skeletal structure creates difficulty when physically sampled, which happens to be one of the only multidecadal living coral records used for coral paleoclimate modeling.


(File:Zoanthus-dragon-eye.jpg|right|thumb|This dragon-eye zoanthid is a popular source of color in reef tanks.)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.Aquarium Corals: Collection and Aquarium Husbandry of Northeast Pacific Non-Photosynthetic Cnidaria. (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 algae scrubber) and a deep sand bed providing filtration,Reefkeeping 101 – Various Nutrient Control Methods. Retrieved on 2016-06-13. or as "show tanks", with the rock kept largely bare of the algae and microfauna that would normally populate it,Aquarium Substrate & Live Rock Clean Up Tips. Retrieved on 2016-06-13. 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.Coral Reefs {{webarchive|url= |date=2013-01-21 }}. Retrieved on 2016-06-13. 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.JOURNAL, Horoszowski-Fridman YB, Izhaki I, Rinkevich B, 2011, Engineering of coral reef larval supply through transplantation of nursery-farmed gravid colonies, Journal of Experimental Marine Biology and Ecology, 399, 2, 162–66, 10.1016/j.jembe.2011.01.005, JOURNAL, Pomeroy, Robert S., Parks, John E., Balboa, Cristina M., 2006, Farming the reef: Is aquaculture a solution for reducing fishing pressure on coral reefs?, Marine Policy, 30, 2, 111–30, 10.1016/j.marpol.2004.09.001, JOURNAL, Rinkevich B, 2008, Management of coral reefs: We have gone wrong when neglecting active reef restoration,weblinkweblink" title="">weblink dead, 2013-05-23, Marine Pollution Bulletin, 56, 11, 1821–24, 10.1016/j.marpolbul.2008.08.014, 18829052, 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.JOURNAL, Ferse, Sebastian C.A., 2010, Poor Performance of Corals Transplanted onto Substrates of Short Durability, Restoration Ecology, 18, 4, 399–407, 10.1111/j.1526-100X.2010.00682.x, 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)File:Mushroom Coral (Fungia) Top Macro 91.JPG|Fungia sp. skeletonFile:Eusmilia fastigiata large.jpg|Polyps of Eusmilia fastigiataFile:Dendrogyra cylindrus (pillar coral) (San Salvador Island, Bahamas) 1 (15513345363).jpg|Pillar coral, Dendrogyra cylindricusFile:Brain coral.jpg|Brain coral, Diploria labyrinthiformisFile:Brain coral spawning.jpg|Brain coral spawningFile:Stony coral spawning 3.jpg|Brain coral releasing eggsFile:EilatFringingReef.jpg|Fringing coral reef off the coast of Eilat, Israel.




  • BOOK, Allen, G.R, R. Steene, Indo-Pacific Coral Reef Field Guide, 1994, 978-981-00-5687-2,
  • BOOK, Calfo, Anthony, Book of Coral Propagation, 978-0-9802365-0-7, 2007,
  • BOOK, Colin, P.L., C. Arneson, Tropical Pacific Invertebrates
    • BOOK, Fagerstrom, J.A., The Evolution of Reef Communities, 1987, 978-0-471-81528-0,
    • BOOK, Gosliner, T., D. Behrens, G. Williams, Coral Reef Animals of the Indo-Pacific, Animals Life from Africa to Hawai'i (invertebrates), 1996, 978-0-930118-21-1,
    • BOOK, Nybakken, J.W., Marine Biology, An Ecological Approach, 2004, 978-0-8053-4582-7,
    • BOOK, Redhill, Surrey, Corals of the World: Biology and Field Guide,
    • BOOK, Segaloff, Nat, Paul Erickson, A Reef Comes to Life. Creating an Undersea Exhibit, 1991, 978-0-531-10994-6, registration,weblink
    • BOOK, Sheppard, Charles R.C., Davy, Simon K., Pilling, Graham M., The Biology of Coral Reefs,weblink 25 June 2009, OUP Oxford, 978-0-19-105734-2,
    • BOOK, Veron, J.E.N., Corals of Australia and the Indo-Pacific, 1993, 978-0-8248-1504-2,
    • BOOK, Wells, Susan, Coral Reefs of the World,

    External links

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