A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of

colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...

of coral polyps held together by calcium carbonate. Most coral reefs are built from

stony coral Scleractinia, also called stony corals or hard corals, are marine animals in the phylum Cnidaria that build themselves a hard skeleton. The individual animals are known as polyps and have a cylindrical body crowned by an oral disc in which a mo ...

s, whose polyps cluster in groups. Coral belongs to the

class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differently ...

Anthozoa Anthozoa is a subphylum of marine invertebrates which includes the sea anemones, stony corals and soft corals. Adult anthozoans are almost all attached to the seabed, while their larvae can disperse as part of the plankton. The basic unit of ...

in the animal phylum

Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...

, which includes sea anemones and jellyfish. Unlike sea anemones, corals secrete hard carbonate exoskeletons that support and protect the coral. Most reefs grow best in warm, shallow, clear, sunny and agitated water. Coral reefs first appeared 485 million years ago, at the dawn of the

Early Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. The ...

, displacing the microbial and

sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through ...

reefs of the

Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...

. Sometimes called ''rainforests of the sea'', shallow coral reefs form some of Earth's most diverse ecosystems. They occupy less than 0.1% of the world's ocean area, about half the area of France, yet they provide a home for at least 25% of all marine

species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...

, including fish, mollusks, worms,

crustaceans Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group c ...

, echinoderms,

s, tunicates and other

cnidarians Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...

. Coral reefs flourish in ocean waters that provide few nutrients. They are most commonly found at shallow depths in tropical waters, but deep water and cold water coral reefs exist on smaller scales in other areas. Coral reefs have declined by 50% since 1950, partly because they are sensitive to water conditions. They are under threat from excess nutrients (nitrogen and phosphorus), rising ocean heat content and acidification, overfishing (e.g., from

blast fishing Blast fishing, fish bombing, dynamite fishing or grenade fishing is a destructive fishing practice using explosives to stun or kill schools of fish for easy collection. This often illegal practice is extremely destructive to the surrounding eco ...

, cyanide fishing, spearfishing on scuba), sunscreen use, and harmful land-use practices, including

runoff Runoff, run-off or RUNOFF may refer to: * RUNOFF, the first computer text-formatting program * Runoff or run-off, another name for bleed, printing that lies beyond the edges to which a printed sheet is trimmed * Runoff or run-off, a stock marke ...

and seeps (e.g., from

injection well An injection well is a device that places fluid deep underground into porous rock formations, such as sandstone or limestone, or into or below the shallow soil layer. The fluid may be water, wastewater, brine (salt water), or water mixed with indus ...

s and cesspools). Coral reefs deliver

ecosystem services Ecosystem services are the many and varied benefits to humans provided by the natural environment and healthy ecosystems. Such ecosystems include, for example, agroecosystems, forest ecosystem, grassland ecosystems, and aquatic ecosystems. ...

for tourism, fisheries and shoreline protection. The annual global economic value of coral reefs has been estimated at anywhere from US$30–375 billion (1997 and 2003 estimates) to US$2.7 trillion (a 2020 estimate) to US$9.9 trillion (a 2014 estimate).

Formation

Most coral reefs were formed after the Last Glacial Period when melting ice caused sea level to rise and flood

continental shelves A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an island ...

. Most coral reefs are less than 10,000 years old. As communities established themselves, the reefs grew upwards, pacing rising sea levels. Reefs that rose too slowly could become drowned, without sufficient light. Coral reefs are also found in the deep sea away from

, around

oceanic islands Oceanic may refer to: *Of or relating to the ocean *Of or relating to Oceania **Oceanic climate **Oceanic languages **Oceanic person or people, also called "Pacific Islander(s)" Places * Oceanic, British Columbia, a settlement on Smith Island, ...

and

atoll An atoll () is a ring-shaped island, including a coral rim that encircles a lagoon partially or completely. There may be coral islands or cays on the rim. Atolls are located in warm tropical or subtropical oceans and seas where corals can gro ...

s. The majority of these islands are volcanic in origin. Others have tectonic origins where plate movements lifted the deep ocean floor. In ''

The Structure and Distribution of Coral Reefs ''The Structure and Distribution of Coral Reefs, Being the first part of the geology of the voyage of the Beagle, under the command of Capt. Fitzroy, R.N. during the years 1832 to 1836'', was published in 1842 as Charles Darwin's first monogr ...

'',Via Internet Archive
/ref>

Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...

set out his theory of the formation of atoll reefs, an idea he conceived during the voyage of the ''Beagle''. He theorized that uplift and

subsidence Subsidence is a general term for downward vertical movement of the Earth's surface, which can be caused by both natural processes and human activities. Subsidence involves little or no horizontal movement, which distinguishes it from slope move ...

of Earth's crust under the oceans formed the atolls. Darwin set out a sequence of three stages in atoll formation. A fringing reef forms around an extinct

volcanic island Geologically, a high island or volcanic island is an island of volcanic origin. The term can be used to distinguish such islands from low islands, which are formed from sedimentation or the uplifting of coral reefs (which have often formed ...

as the island and ocean floor subside. As the subsidence continues, the fringing reef becomes a barrier reef and ultimately an atoll reef. File:Atoll forming-volcano.png, Darwin's theory starts with a

which becomes extinct File:Atoll forming-Fringing reef.png, As the island and ocean floor subside, coral growth builds a fringing reef, often including a shallow lagoon between the land and the main reef. File:Atoll forming-Barrier reef.png, As the subsidence continues, the fringing reef becomes a larger barrier reef further from the shore with a bigger and deeper

lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'' ...

inside. File:Atoll forming-Atoll.png, Ultimately, the island sinks below the sea, and the barrier reef becomes an

enclosing an open lagoon. Darwin predicted that underneath each

would be a

bedrock In geology, bedrock is solid rock that lies under loose material (regolith) within the crust of Earth or another terrestrial planet. Definition Bedrock is the solid rock that underlies looser surface material. An exposed portion of bedro ...

base, the remains of the original volcano. Subsequent research supported this hypothesis. Darwin's theory followed from his understanding that coral polyps thrive in the tropics where the water is agitated, but can only live within a limited depth range, starting just below low tide. Where the level of the underlying earth allows, the corals grow around the coast to form fringing reefs, and can eventually grow to become a barrier reef. Coral atoll formation animation

Where the bottom is rising, fringing reefs can grow around the coast, but coral raised above sea level dies. If the land subsides slowly, the fringing reefs keep pace by growing upwards on a base of older, dead coral, forming a barrier reef enclosing a lagoon between the reef and the land. A barrier reef can encircle an island, and once the island sinks below sea level a roughly circular atoll of growing coral continues to keep up with the sea level, forming a central lagoon. Barrier reefs and atolls do not usually form complete circles but are broken in places by storms. Like sea level rise, a rapidly subsiding bottom can overwhelm coral growth, killing the coral and the reef, due to what is called ''coral drowning''. Corals that rely on zooxanthellae can die when the water becomes too deep for their

symbionts Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...

to adequately

photosynthesize Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...

, due to decreased light exposure. The two main variables determining the

geomorphology Geomorphology (from Ancient Greek: , ', "earth"; , ', "form"; and , ', "study") is the scientific study of the origin and evolution of topographic and bathymetric features created by physical, chemical or biological processes operating at or ...

, or shape, of coral reefs are the nature of the substrate on which they rest, and the history of the change in sea level relative to that substrate. The approximately 20,000-year-old Great Barrier Reef offers an example of how coral reefs formed on continental shelves. Sea level was then lower than in the 21st century. As sea level rose, the water and the corals encroached on what had been hills of the Australian coastal plain. By 13,000 years ago, sea level had risen to lower than at present, and many hills of the coastal plains had become continental islands. As sea level rise continued, water topped most of the continental islands. The corals could then overgrow the hills, forming

cay A cay ( ), also spelled caye or key, is a small, low-elevation, sandy island on the surface of a coral reef. Cays occur in tropical environments throughout the Pacific, Atlantic, and Indian Oceans, including in the Caribbean and on the Great ...

s and reefs. Sea level on the Great Barrier Reef has not changed significantly in the last 6,000 years. The age of living reef structure is estimated to be between 6,000 and 8,000 years. Although the Great Barrier Reef formed along a continental shelf, and not around a volcanic island, Darwin's principles apply. Development stopped at the barrier reef stage, since Australia is not about to submerge. It formed the world's largest barrier reef, from shore, stretching for . Healthy tropical coral reefs grow horizontally from per year, and grow vertically anywhere from per year; however, they grow only at depths shallower than because of their need for sunlight, and cannot grow above sea level.

Material

As the name implies, coral reefs are made up of coral skeletons from mostly intact coral colonies. As other chemical elements present in corals become incorporated into the calcium carbonate deposits,

aragonite Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including pr ...

is formed. However, shell fragments and the remains of

coralline algae Coralline algae are red algae in the order Corallinales. They are characterized by a thallus that is hard because of calcareous deposits contained within the cell walls. The colors of these algae are most typically pink, or some other shade of re ...

such as the green-segmented genus ''

Halimeda ''Halimeda'' is a genus of green macroalgae. The algal body (thallus) is composed of calcified green segments. Calcium carbonate is deposited in its tissues, making it inedible to most herbivores. However one species, '' Halimeda tuna'', was desc ...

'' can add to the reef's ability to withstand damage from storms and other threats. Such mixtures are visible in structures such as

Eniwetok Atoll Enewetak Atoll (; also spelled Eniwetok Atoll or sometimes Eniewetok; mh, Ānewetak, , or , ; known to the Japanese as Brown Atoll or Brown Island; ja, ブラウン環礁) is a large coral atoll of 40 islands in the Pacific Ocean and with it ...

Types

Since Darwin's identification of the three classical reef formations – the fringing reef around a volcanic island becoming a barrier reef and then an atoll – scientists have identified further reef types. While some sources find only three, Thomas and Goudie list four "principal large-scale coral reef types" – the fringing reef, barrier reef, atoll and table reef – while Spalding ''et al.'' list five "main types" – the fringing reef, barrier reef, atoll, "bank or platform reef" and patch reef.

Fringing reef

A fringing reef, also called a shore reef, is directly attached to a shore, or borders it with an intervening narrow, shallow channel or lagoon. It is the most common reef type.''Types of Coral Reef Formations''
at coral.org. Retrieved 2 Feb 2018. Fringing reefs follow coastlines and can extend for many kilometres. They are usually less than 100 metres wide, but some are hundreds of metres wide. Fringing reefs are initially formed on the shore at the

low water Tides are the rise and fall of sea levels caused by the combined effects of the gravitational forces exerted by the Moon (and to a much lesser extent, the Sun) and are also caused by the Earth and Moon orbiting one another. Tide tables ...

level and expand seawards as they grow in size. The final width depends on where the sea bed begins to drop steeply. The surface of the fringe reef generally remains at the same height: just below the waterline. In older fringing reefs, whose outer regions pushed far out into the sea, the inner part is deepened by erosion and eventually forms a

. Fringing reef lagoons can become over 100 metres wide and several metres deep. Like the fringing reef itself, they run parallel to the coast. The fringing reefs of the Red Sea are "some of the best developed in the world" and occur along all its shores except off sandy bays.

Barrier reef

Barrier reefs are separated from a mainland or island shore by a deep channel or

. They resemble the later stages of a fringing reef with its lagoon but differ from the latter mainly in size and origin. Their lagoons can be several kilometres wide and 30 to 70 metres deep. Above all, the offshore outer reef edge formed in open water rather than next to a shoreline. Like an atoll, it is thought that these reefs are formed either as the seabed lowered or sea level rose. Formation takes considerably longer than for a fringing reef, thus barrier reefs are much rarer. The best known and largest example of a barrier reef is the Australian Great Barrier Reef.''Types of Coral Reefs''
at www.coral-reef-info.com. Retrieved 2 Feb 2018. Other major examples are the

Belize Barrier Reef The Belize Barrier Reef is a series of coral reefs straddling the coast of Belize, roughly offshore in the north and in the south within the country limits. The Belize Barrier Reef is a long section of the Mesoamerican Barrier Reef System, ...

and the

New Caledonian Barrier Reef The New Caledonian barrier reef is a barrier reef located in New Caledonia in the South Pacific, being the longest continuous barrier reef in the world and the third largest after the Great Barrier Reef of Australia and the Mesoamerican Barrier ...

. Barrier reefs are also found on the coasts of Providencia, Mayotte, the Gambier Islands, on the southeast coast of Kalimantan, on parts of the coast of Sulawesi, southeastern New Guinea and the south coast of the Louisiade Archipelago.

Platform reef

Platform reefs, variously called bank or table reefs, can form on the continental shelf, as well as in the open ocean, in fact anywhere where the seabed rises close enough to the surface of the ocean to enable the growth of zooxanthemic, reef-forming corals. Platform reefs are found in the southern Great Barrier Reef, the Swain and Capricorn Group on the continental shelf, about 100–200 km from the coast. Some platform reefs of the northern

Mascarenes The Mascarene Islands (, ) or Mascarenes or Mascarenhas Archipelago is a group of islands in the Indian Ocean east of Madagascar consisting of the islands belonging to the Republic of Mauritius as well as the French department of Réunion. Their ...

are several thousand kilometres from the mainland. Unlike fringing and barrier reefs which extend only seaward, platform reefs grow in all directions. They are variable in size, ranging from a few hundred metres to many kilometres across. Their usual shape is oval to elongated. Parts of these reefs can reach the surface and form sandbanks and small islands around which may form fringing reefs. A lagoon may form In the middle of a platform reef. Platform reefs can be found within atolls. There they are called patch reefs and may reach only a few dozen metres in diameter. Where platform reefs form on an elongated structure, e. g. an old, eroded barrier reef, they can form a linear arrangement. This is the case, for example, on the east coast of the Red Sea near Jeddah. In old platform reefs, the inner part can be so heavily eroded that it forms a pseudo-atoll. These can be distinguished from real atolls only by detailed investigation, possibly including core drilling. Some platform reefs of the

Laccadives Lakshadweep (), also known as Laccadives (), is a union territory of India. It is an archipelago of 36 islands in the Arabian sea, located off the Malabar Coast. The name ''Lakshadweep'' means "one lakh islands" in Sanskrit, though the Lacc ...

are U-shaped, due to wind and water flow.

Atoll

Atolls or

atoll reef An atoll () is a ring-shaped island, including a coral rim that encircles a lagoon partially or completely. There may be coral islands or cays on the rim. Atolls are located in warm tropical or subtropical oceans and seas where corals can grow ...

s are a more or less circular or continuous barrier reef that extends all the way around a lagoon without a central island. They are usually formed from fringing reefs around volcanic islands. Over time, the island erodes away and sinks below sea level. Atolls may also be formed by the sinking of the seabed or rising of the sea level. A ring of reefs results, which enclose a lagoon. Atolls are numerous in the South Pacific, where they usually occur in mid-ocean, for example, in the

Caroline Islands The Caroline Islands (or the Carolines) are a widely scattered archipelago of tiny islands in the western Pacific Ocean, to the north of New Guinea. Politically, they are divided between the Federated States of Micronesia (FSM) in the centra ...

, the

Cook Islands ) , image_map = Cook Islands on the globe (small islands magnified) (Polynesia centered).svg , capital = Avarua , coordinates = , largest_city = Avarua , official_languages = , langu ...

French Polynesia )Territorial motto: ( en, "Great Tahiti of the Golden Haze") , anthem = , song_type = Regional anthem , song = " Ia Ora 'O Tahiti Nui" , image_map = French Polynesia on the globe (French Polynesia centered).svg , map_alt = Location of Frenc ...

, the Marshall Islands and Micronesia. Atolls are found in the Indian Ocean, for example, in the

Maldives Maldives (, ; dv, ދިވެހިރާއްޖެ, translit=Dhivehi Raajje, ), officially the Republic of Maldives ( dv, ދިވެހިރާއްޖޭގެ ޖުމްހޫރިއްޔާ, translit=Dhivehi Raajjeyge Jumhooriyyaa, label=none, ), is an archipela ...

, the

Chagos Islands The Chagos Archipelago () or Chagos Islands (formerly the Bassas de Chagas, and later the Oil Islands) is a group of seven atolls comprising more than 60 islands in the Indian Ocean about 500 kilometres (310 mi) south of the Maldives archi ...

, the

Seychelles Seychelles (, ; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic state consisting of 115 islands in the Indian Ocean. Its capital and largest city, ...

and around Cocos Island. The entire Maldives consist of 26 atolls.

Other reef types or variants

* Apron reef – short reef resembling a fringing reef, but more sloped; extending out and downward from a point or peninsular shore. The initial stage of a fringing reef.National Oceanic and Atmospheric Administration. ''Coral Reef Information System Glossary'', 2014. * Bank reef – isolated, flat-topped reef larger than a patch reef and usually on mid-shelf regions and linear or semi-circular in shape; a type of platform reef. * Patch reef – common, isolated, comparatively small reef outcrop, usually within a

embayment A bay is a recessed, coastal body of water that directly connects to a larger main body of water, such as an ocean, a lake, or another bay. A large bay is usually called a gulf, sea, sound, or bight. A cove is a small, circular bay with a na ...

, often circular and surrounded by sand or

seagrass Seagrasses are the only flowering plants which grow in marine environments. There are about 60 species of fully marine seagrasses which belong to four families ( Posidoniaceae, Zosteraceae, Hydrocharitaceae and Cymodoceaceae), all in the ...

. Can be considered as a type of platform reef or as features of fringing reefs, atolls and barrier reefs. The patches may be surrounded by a ring of reduced seagrass cover referred to as a ''grazing halo''. * Ribbon reef – long, narrow, possibly winding reef, usually associated with an atoll lagoon. Also called a shelf-edge reef or sill reef. * Habili – reef specific to the Red Sea; does not reach near enough to the surface to cause visible surf; may be a hazard to ships (from the

Arabic Arabic (, ' ; , ' or ) is a Semitic language spoken primarily across the Arab world.Semitic languages: an international handbook / edited by Stefan Weninger; in collaboration with Geoffrey Khan, Michael P. Streck, Janet C. E.Watson; Walte ...

for "unborn") * Microatoll – community of species of corals; vertical growth limited by average tidal height; growth morphologies offer a low-resolution record of patterns of sea level change; fossilized remains can be dated using radioactive carbon dating and have been used to reconstruct Holocene sea levels *

Cay A cay ( ), also spelled caye or key, is a small, low-elevation, sandy island on the surface of a coral reef. Cays occur in tropical environments throughout the Pacific, Atlantic, and Indian Oceans, including in the Caribbean and on the Great ...

s – small, low-elevation, sandy islands formed on the surface of coral reefs from eroded material that piles up, forming an area above sea level; can be stabilized by plants to become habitable; occur in tropical environments throughout the Pacific, Atlantic and Indian Oceans (including the Caribbean and on the Great Barrier Reef and Belize Barrier Reef), where they provide habitable and agricultural land *

Seamount A seamount is a large geologic landform that rises from the ocean floor that does not reach to the water's surface (sea level), and thus is not an island, islet or cliff-rock. Seamounts are typically formed from extinct volcanoes that rise abr ...

guyot In marine geology, a guyot (pronounced ), also known as a tablemount, is an isolated underwater volcanic mountain (seamount) with a flat top more than below the surface of the sea. The diameters of these flat summits can exceed . Coral reef ecosystems contain distinct zones that host different kinds of habitats. Usually, three major zones are recognized: the fore reef, reef crest, and the back reef (frequently referred to as the reef lagoon). The three zones are physically and ecologically interconnected. Reef life and oceanic processes create opportunities for the exchange of seawater, sediments, nutrients and marine life. Most coral reefs exist in waters less than 50 m deep. Some inhabit tropical continental shelves where cool, nutrient-rich

upwelling Upwelling is an oceanographic phenomenon that involves wind-driven motion of dense, cooler, and usually nutrient-rich water from deep water towards the ocean surface. It replaces the warmer and usually nutrient-depleted surface water. The nutr ...

does not occur, such as the Great Barrier Reef. Others are found in the deep ocean surrounding islands or as atolls, such as in the

. The reefs surrounding islands form when islands subside into the ocean, and atolls form when an island subsides below the surface of the sea. Alternatively, Moyle and Cech distinguish six zones, though most reefs possess only some of the zones. Propagation du tsunami en profondeur variable

Propagation du tsunami en profondeur variable

The reef surface is the shallowest part of the reef. It is subject to

surge Surge means a sudden transient rush or flood, and may refer to: Science * Storm surge, the onshore gush of water associated with a low-pressure weather system * Surge (glacier), a short-lived event where a glacier can move up to velocities 100 ...

and tides. When waves pass over shallow areas, they shoal, as shown in the adjacent diagram. This means the water is often agitated. These are the precise condition under which corals flourish. The light is sufficient for photosynthesis by the symbiotic zooxanthellae, and agitated water brings plankton to feed the coral. The off-reef floor is the shallow sea floor surrounding a reef. This zone occurs next to reefs on continental shelves. Reefs around tropical islands and atolls drop abruptly to great depths and do not have such a floor. Usually sandy, the floor often supports seagrass meadows which are important foraging areas for reef fish. The reef drop-off is, for its first 50 m, habitat for reef fish who find shelter on the cliff face and plankton in the water nearby. The drop-off zone applies mainly to the reefs surrounding oceanic islands and atolls. The reef face is the zone above the reef floor or the reef drop-off. This zone is often the reef's most diverse area. Coral and

calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an adje ...

algae provide complex habitats and areas that offer protection, such as cracks and crevices. Invertebrates and epiphytic algae provide much of the food for other organisms. A common feature on this forereef zone is spur and groove formations that serve to transport sediment downslope. The reef flat is the sandy-bottomed flat, which can be behind the main reef, containing chunks of coral. This zone may border a lagoon and serve as a protective area, or it may lie between the reef and the shore, and in this case is a flat, rocky area. Fish tend to prefer it when it is present. The reef lagoon is an entirely enclosed region, which creates an area less affected by wave action and often contains small reef patches. However, the "topography of coral reefs is constantly changing. Each reef is made up of irregular patches of algae,

sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...

invertebrates, and bare rock and sand. The size, shape and relative abundance of these patches change from year to year in response to the various factors that favor one type of patch over another. Growing coral, for example, produces constant change in the fine structure of reefs. On a larger scale, tropical storms may knock out large sections of reef and cause boulders on sandy areas to move."

Locations

Coral reefs are estimated to cover 284,300 km² (109,800 sq mi), just under 0.1% of the oceans' surface area. The Indo-Pacific region (including the Red Sea, Indian Ocean, Southeast Asia and the Pacific) account for 91.9% of this total. Southeast Asia accounts for 32.3% of that figure, while the Pacific including Australia accounts for 40.8%. Atlantic and

Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean S ...

coral reefs account for 7.6%.Spalding, Mark, Corinna Ravilious, and Edmund Green (2001). ''World Atlas of Coral Reefs''. Berkeley, CA: University of California Press and UNEP/WCMC . Although corals exist both in temperate and tropical waters, shallow-water reefs form only in a zone extending from approximately 30° N to 30° S of the equator. Tropical corals do not grow at depths of over . The optimum temperature for most coral reefs is , and few reefs exist in waters below . However, reefs in the Persian Gulf have adapted to temperatures of in winter and in summer. 37 species of scleractinian corals inhabit such an environment around Larak Island.

Deep-water coral The habitat of deep-water corals, also known as cold-water corals, extends to deeper, darker parts of the oceans than tropical corals, ranging from near the surface to the abyss, beyond where water temperatures may be as cold as . Deep-water co ...

inhabits greater depths and colder temperatures at much higher latitudes, as far north as Norway. Although deep water corals can form reefs, little is known about them. Coral reefs are rare along the west coasts of the

Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with t ...

and

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

, due primarily to

and strong cold coastal currents that reduce water temperatures in these areas (the Peru,

Benguela Benguela (; Umbundu: Luombaka) is a city in western Angola, capital of Benguela Province. Benguela is one of Angola's most populous cities with a population of 555,124 in the city and 561,775 in the municipality, at the 2014 census. History Po ...

and

Canary Current The Canary Current is a wind-driven surface current that is part of the North Atlantic Gyre. This eastern boundary current branches south from the North Atlantic Current and flows southwest about as far as Senegal where it turns west and later ...

s respectively).Nybakken, James. 1997. ''Marine Biology: An Ecological Approach.'' 4th ed. Menlo Park, CA: Addison Wesley. Corals are seldom found along the coastline of South Asia—from the eastern tip of India (

Chennai Chennai (, ), formerly known as Madras (List of renamed Indian cities and states#Tamil Nadu, the official name until 1996), is the capital city of Tamil Nadu, the southernmost states and territories of India, Indian state. The largest city ...

) to the

Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mos ...

and Myanmar borders—as well as along the coasts of northeastern South America and Bangladesh, due to the freshwater release from the

Amazon Amazon most often refers to: * Amazons, a tribe of female warriors in Greek mythology * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon River, in South America * Amazon (company), an American multinational technology co ...

and Ganges Rivers respectively. Significant coral reefs include: * The Great Barrier Reef—largest, comprising over 2,900 individual reefs and 900 islands stretching for over off Queensland, Australia * The

Mesoamerican Barrier Reef System The Mesoamerican Barrier Reef System (MBRS), also popularly known as the Great Mayan Reef or Great Maya Reef, is a marine region that stretches over along the coasts of four countries – Mexico, Belize, Guatemala, and Honduras – from Isla Con ...

—second largest, stretching from Isla Contoy at the tip of the

Yucatán Peninsula The Yucatán Peninsula (, also , ; es, Península de Yucatán ) is a large peninsula in southeastern Mexico and adjacent portions of Belize and Guatemala. The peninsula extends towards the northeast, separating the Gulf of Mexico to the north ...

down to the Bay Islands of Honduras * The New Caledonia Barrier Reef—second longest double barrier reef, covering * The Andros, Bahamas Barrier Reef—third largest, following the east coast of Andros Island, Bahamas, between

Andros Andros ( el, Άνδρος, ) is the northernmost island of the Greek Cyclades archipelago, about southeast of Euboea, and about north of Tinos. It is nearly long, and its greatest breadth is . It is for the most part mountainous, with many ...

and Nassau * The Red Sea—includes 6,000-year-old fringing reefs located along a coastline * The

Florida Reef Tract The Florida Reef (also known as the Great Florida Reef, Florida reefs, Florida Reef Tract and Florida Keys Reef Tract) is the only living coral barrier reef in the continental United States. It is the third largest coral barrier reef system in ...

—largest continental US reef and the third-largest coral barrier reef, extends from

Soldier Key Soldier Key is an island in Biscayne National Park in Miami-Dade County, Florida. It is located between Biscayne Bay and the Atlantic Ocean, about three miles north of the Ragged Keys, five miles south of Cape Florida on Key Biscayne, seven-and ...

, located in

Biscayne Bay Biscayne Bay () is a lagoon with characteristics of an estuary located on the Atlantic coast of South Florida. The northern end of the lagoon is surrounded by the densely developed heart of the Miami metropolitan area while the southern end is l ...

, to the Dry Tortugas in the Gulf of Mexico *

Pulley Ridge Pulley Ridge is a mesophotic coral reef system off the shores of the continental United States. The reef rests on sunken barrier islands and lies 100 miles west of the Tortugas Ecological Reserve and stretches north about 60 miles at depths rangi ...

—deepest photosynthetic coral reef, Florida * Numerous reefs around the

* The Philippines coral reef area, the second-largest in Southeast Asia, is estimated at 26,000 square kilometers. 915 reef fish species and more than 400 scleractinian coral species, 12 of which are endemic are found there. * The

Raja Ampat Islands Raja Ampat, or the ''Four Kings'', is an archipelago located off the northwest tip of Bird's Head Peninsula on the island of New Guinea, in Indonesia's Southwest Papua province. It comprises over 1,500 small islands, cays, and shoals surrounding ...

in Indonesia's West Papua province offer the highest known marine diversity. *

Bermuda ) , anthem = "God Save the King" , song_type = National song , song = " Hail to Bermuda" , image_map = , map_caption = , image_map2 = , mapsize2 = , map_caption2 = , subdivision_type = Sovereign state , subdivision_name = , e ...

is known for its northernmost coral reef system, located at . The presence of coral reefs at this high latitude is due to the proximity of the

Gulf Stream The Gulf Stream, together with its northern extension the North Atlantic Drift, is a warm and swift Atlantic ocean current that originates in the Gulf of Mexico and flows through the Straits of Florida and up the eastern coastline of the Unit ...

. Bermuda coral species represent a subset of those found in the greater Caribbean. * The world's northernmost individual coral reef is located within a bay of Japan's Tsushima Island in the Korea Strait. * The world's southernmost coral reef is at Lord Howe Island, in the Pacific Ocean off the east coast of Australia.

Coral

When alive, corals are

of small animals embedded in calcium carbonate shells. Coral heads consist of accumulations of individual animals called polyps, arranged in diverse shapes. Polyps are usually tiny, but they can range in size from a pinhead to across. Reef-building or

hermatypic coral Hermatypic corals are those corals in the order Scleractinia which build reefs by depositing hard calcareous material for their skeletons, forming the stony framework of the reef. Corals that do not contribute to coral reef development are refer ...

s live only in the

photic zone The photic zone, euphotic zone, epipelagic zone, or sunlight zone is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological pro ...

(above 70 m), the depth to which sufficient sunlight penetrates the water.

Zooxanthellae

Coral polyps do not photosynthesize, but have a symbiotic relationship with microscopic

algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...

( dinoflagellates) of the genus ''

Symbiodinium : ''This is about the genus sometimes called Zoox. For the company, see Zoox (company)'' ''Symbiodinium'' is a genus of dinoflagellates that encompasses the largest and most prevalent group of endosymbiotic dinoflagellates known. These unicellul ...

'', commonly referred to as zooxanthellae. These organisms live within the polyps' tissues and provide organic nutrients that nourish the polyp in the form of glucose, glycerol and

amino acids Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...

. Because of this relationship, coral reefs grow much faster in clear water, which admits more sunlight. Without their symbionts, coral growth would be too slow to form significant reef structures. Corals get up to 90% of their nutrients from their symbionts. In return, as an example of mutualism, the corals shelter the zooxanthellae, averaging one million for every cubic centimeter of coral, and provide a constant supply of the carbon dioxide they need for photosynthesis. Coral detail

The varying pigments in different species of zooxanthellae give them an overall brown or golden-brown appearance and give brown corals their colors. Other pigments such as reds, blues, greens, etc. come from colored proteins made by the coral animals. Coral that loses a large fraction of its zooxanthellae becomes white (or sometimes pastel shades in corals that are pigmented with their own proteins) and is said to be bleached, a condition which, unless corrected, can kill the coral. There are eight clades of ''Symbiodinium''

phylotype In taxonomy, a phylotype is an observed similarity used to classify a group of organisms by their phenetic relationship. This phenetic similarity, particularly in the case of asexual organisms, may reflect the evolutionary relationships. The term ...

s. Most research has been conducted on clades A–D. Each clade contributes their own benefits as well as less compatible attributes to the survival of their coral hosts. Each photosynthetic organism has a specific level of sensitivity to photodamage to compounds needed for survival, such as proteins. Rates of regeneration and replication determine the organism's ability to survive. Phylotype A is found more in the shallow waters. It is able to produce

mycosporine-like amino acid Mycosporine-like amino acids (MAAs) are small secondary metabolites produced by organisms that live in environments with high volumes of sunlight, usually marine environments. The exact number of compounds within this class of natural products is ye ...

s that are UV resistant, using a derivative of

glycerin Glycerol (), also called glycerine in British English and glycerin in American English, is a simple triol compound. It is a colorless, odorless, viscous liquid that is sweet-tasting and non-toxic. The glycerol backbone is found in lipids known ...

to absorb the UV radiation and allowing them to better adapt to warmer water temperatures. In the event of UV or thermal damage, if and when repair occurs, it will increase the likelihood of survival of the host and symbiont. This leads to the idea that, evolutionarily, clade A is more UV resistant and thermally resistant than the other clades. Clades B and C are found more frequently in deeper water, which may explain their higher vulnerability to increased temperatures. Terrestrial plants that receive less sunlight because they are found in the undergrowth are analogous to clades B, C, and D. Since clades B through D are found at deeper depths, they require an elevated light absorption rate to be able to synthesize as much energy. With elevated absorption rates at UV wavelengths, these phylotypes are more prone to coral bleaching versus the shallow clade A. Clade D has been observed to be high temperature-tolerant, and has a higher rate of survival than clades B and C during modern bleaching events.

Skeleton

Reefs grow as polyps and other organisms deposit calcium carbonate, the basis of coral, as a skeletal structure beneath and around themselves, pushing the coral head's top upwards and outwards. Waves, grazing fish (such as parrotfish), sea urchins, sponges and other forces and organisms act as bioeroders, breaking down coral skeletons into fragments that settle into spaces in the reef structure or form sandy bottoms in associated reef lagoons. Typical shapes for coral

are named by their resemblance to terrestrial objects such as wrinkled brains, cabbages,

table tops ''Table Tops'' was a free daily newspaper produced for Australian Army personnel, published in the Atherton Tableland Training Area in Queensland, Australia,{{cite web, title= Provincial publishing proliferated in Australia's colonial era, ...

antlers Antlers are extensions of an animal's skull found in members of the Cervidae (deer) family. Antlers are a single structure composed of bone, cartilage, fibrous tissue, skin, nerves, and blood vessels. They are generally found only on male ...

, wire strands and pillars. These shapes can depend on the life history of the coral, like light exposure and wave action, and events such as breakages.

Reproduction

Corals reproduce both sexually and asexually. An individual polyp uses both reproductive modes within its lifetime. Corals reproduce sexually by either internal or external fertilization. The reproductive cells are found on the

mesenteries In zoology, a mesentery is a membrane inside the body cavity of an animal. The term identifies different structures in different phyla: in vertebrates it is a double fold of the peritoneum enclosing the intestines; in other organisms it forms co ...

, membranes that radiate inward from the layer of tissue that lines the stomach cavity. Some mature adult corals are hermaphroditic; others are exclusively male or female. A few

change sex as they grow. Internally fertilized eggs develop in the polyp for a period ranging from days to weeks. Subsequent development produces a tiny larva, known as a planula. Externally fertilized eggs develop during synchronized spawning. Polyps across a reef simultaneously release eggs and sperm into the water en masse. Spawn disperse over a large area. The timing of spawning depends on time of year, water temperature, and tidal and lunar cycles. Spawning is most successful given little variation between high and low tide. The less water movement, the better the chance for fertilization. Ideal timing occurs in the spring. The release of eggs or planula usually occurs at night and is sometimes in phase with the lunar cycle (three to six days after a full moon). The period from release to settlement lasts only a few days, but some planulae can survive afloat for several weeks. During this process, the larvae may use several different cues to find a suitable location for settlement. At long distances sounds from existing reefs are likely important, while at short distances chemical compounds become important. The larvae are vulnerable to predation and environmental conditions. The lucky few planulae that successfully attach to substrate then compete for food and space.

Gallery of reef-building corals

Other reef builders

Corals are the most prodigious reef-builders. However many other organisms living in the reef community contribute skeletal calcium carbonate in the same manner as corals. These include

, some sponges and

bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, biva ...

s. Reefs are always built by the combined efforts of these different phyla, with different organisms leading reef-building in different

geological period The geologic time scale, or geological time scale, (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating strata to time) and geochrono ...

Coralline algae

Coralline algae Coralline algae are red algae in the order Corallinales. They are characterized by a thallus that is hard because of calcareous deposits contained within the cell walls. The colors of these algae are most typically pink, or some other shade of re ...

are important contributors to reef structure. Although their mineral deposition rates are much slower than corals, they are more tolerant of rough wave-action, and so help to create a protective crust over those parts of the reef subjected to the greatest forces by waves, such as the reef front facing the open ocean. They also strengthen the reef structure by depositing limestone in sheets over the reef surface.

Sponges

Sclerosponge Demosponges (Demospongiae) are the most diverse class in the phylum Porifera. They include 76.2% of all species of sponges with nearly 8,800 species worldwide (World Porifera Database). They are sponges with a soft body that covers a hard, ...

" is the descriptive name for all ''Porifera'' that build reefs. In the early

Cambrian period The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...

, ''Archaeocyatha'' sponges were the world's first reef-building organisms, and sponges were the only reef-builders until the

Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. The ...

s still assist corals building modern reefs, but like

are much slower-growing than corals and their contribution is (usually) minor. In the northern Pacific Ocean

cloud sponge The cloud sponge ''(Aphrocallistes vastus)'' is a species of sea sponge in the class Hexactinellida. It is a deep-water reef-forming animal. The species was first described by F.E. Schulze in 1886. Description The cloud sponge takes the form o ...

s still create deep-water mineral-structures without corals, although the structures are not recognizable from the surface like tropical reefs. They are the only

extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extin ...

organisms known to build reef-like structures in cold water.

Bivalves

Oyster reefs are dense aggregations of oysters living in colonial communities. Other regionally-specific names for these structures include oyster beds and oyster banks. Oyster larvae require a hard substrate or surface to attach on, which includes the shells of old or dead oysters. Thus reefs can build up over time as new larvae settle on older individuals. ''

Crassostrea virginica The eastern oyster (''Crassostrea virginica'')—also called the Atlantic oyster, American oyster, or East Coast oyster—is a species of true oyster native to eastern North and South America. Other names in local or culinary use include the We ...

'' were once abundant in

Chesapeake Bay The Chesapeake Bay ( ) is the largest estuary in the United States. The Bay is located in the Mid-Atlantic region and is primarily separated from the Atlantic Ocean by the Delmarva Peninsula (including the parts: the Eastern Shore of Maryland / E ...

and shorelines bordering the

Atlantic coastal plain The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the "Old World" of Africa, Europe an ...

until the late nineteenth century. ''

Ostrea angasi The southern mud oyster, Australian flat oyster, native flat oyster, native mud oyster, or angasi oyster (''Ostrea angasi''), is endemic to southern Australia, ranging from Western Australia to southeast New South Wales and around Tasmania. '' ...

'' is a species of flat oyster that had also formed large reefs in South Australia. Hippuritida, an extinct order of bivalves known as

rudists Rudists are a group of extinct box-, tube- or ring-shaped marine heterodont bivalves belonging to the order Hippuritida that arose during the Late Jurassic and became so diverse during the Cretaceous that they were major reef-building organis ...

, were major reef-building organisms during the Cretaceous. By the mid-Cretaceous, rudists became the dominant tropical reef-builders, becoming more numerous than scleractinian corals. During this period, ocean temperatures and saline levels—which corals are sensitive to—were higher than it is today, which may have contributed to the success of rudist reefs.

Darwin's paradox

In ''

'', published in 1842, Darwin described how coral reefs were found in some tropical areas but not others, with no obvious cause. The largest and strongest corals grew in parts of the reef exposed to the most violent surf and corals were weakened or absent where loose sediment accumulated. Tropical waters contain few nutrients yet a coral reef can flourish like an "oasis in the desert". This has given rise to the ecosystem conundrum, sometimes called "Darwin's paradox": "How can such high production flourish in such nutrient poor conditions?"De Goeij, Jasper M (2009
"Element cycling on tropical coral reefs: the cryptic carbon shunt revealed"
PhD thesis, page 13. University of Groningen. Coral reefs support over one-quarter of all marine species. This diversity results in complex food webs, with large predator fish eating smaller forage fish that eat yet smaller zooplankton and so on. However, all food webs eventually depend on plants, which are the primary producers. Coral reefs typically produce 5–10 grams of carbon per square meter per day (gC·m⁻²·day⁻¹) Biomass (ecology), biomass. One reason for the unusual clarity of tropical waters is their nutrient deficiency and drifting plankton. Further, the sun shines year-round in the tropics, warming the surface layer, making it less dense than subsurface layers. The warmer water is separated from deeper, cooler water by a stable thermocline, where the temperature makes a rapid change. This keeps the warm surface waters floating above the cooler deeper waters. In most parts of the ocean, there is little exchange between these layers. Organisms that die in aquatic environments generally sink to the bottom, where they decompose, which releases nutrients in the form of nitrogen (N), phosphorus (P) and potassium (K). These nutrients are necessary for plant growth, but in the tropics, they do not directly return to the surface. Plants form the base of the food chain and need sunlight and nutrients to grow. In the ocean, these plants are mainly microscopic phytoplankton which drift in the water column. They need sunlight for photosynthesis, which powers carbon fixation, so they are found only relatively near the surface, but they also need nutrients. Phytoplankton rapidly use nutrients in the surface waters, and in the tropics, these nutrients are not usually replaced because of the thermocline.

Explanations

Around coral reefs, lagoons fill in with material eroded from the reef and the island. They become havens for marine life, providing protection from waves and storms. Most importantly, reefs Biogeochemical cycle, recycle nutrients, which happens much less in the open ocean. In coral reefs and lagoons, producers include phytoplankton, as well as seaweed and coralline algae, especially small types called turf algae, which pass nutrients to corals. The phytoplankton form the base of the food chain and are eaten by fish and crustaceans. Recycling reduces the nutrient inputs needed overall to support the community. Corals also absorb nutrients, including inorganic nitrogen and phosphorus, directly from water. Many corals extend their tentacles at night to catch zooplankton that pass near. Zooplankton provide the polyp with nitrogen, and the polyp shares some of the nitrogen with the zooxanthellae, which also require this element.Castro, Peter and Huber, Michael (2000) ''Marine Biology.'' 3rd ed. Boston: McGraw-Hill. Multy color corals

Sponges live in crevices in the reefs. They are efficient filter feeders, and in the Red Sea they consume about 60% of the phytoplankton that drifts by. Sponges eventually excrete nutrients in a form that corals can use. The roughness of coral surfaces is key to coral survival in agitated waters. Normally, a boundary layer of still water surrounds a submerged object, which acts as a barrier. Waves breaking on the extremely rough edges of corals disrupt the boundary layer, allowing the corals access to passing nutrients. Turbulent water thereby promotes reef growth. Without the access to nutrients brought by rough coral surfaces, even the most effective recycling would not suffice. Deep nutrient-rich water entering coral reefs through isolated events may have significant effects on temperature and nutrient systems. This water movement disrupts the relatively stable thermocline that usually exists between warm shallow water and deeper colder water. Temperature regimes on coral reefs in the Bahamas and Florida are highly variable with temporal scales of minutes to seasons and spatial scales across depths. Polyps (PSF)

Water can pass through coral reefs in various ways, including current rings, surface waves, internal waves and tidal changes. Movement is generally created by tides and wind. As tides interact with varying bathymetry and wind mixes with surface water, internal waves are created. An internal wave is a gravity wave that moves along density stratification within the ocean. When a water parcel encounters a different density it oscillates and creates internal waves. While internal waves generally have a lower frequency than surface waves, they often form as a single wave that breaks into multiple waves as it hits a slope and moves upward. This vertical breakup of internal waves causes significant diapycnal mixing and turbulence. Internal waves can act as nutrient pumps, bringing plankton and cool nutrient-rich water to the surface. Sea Cotton

The irregular structure characteristic of coral reef bathymetry may enhance mixing and produce pockets of cooler water and variable nutrient content. Arrival of cool, nutrient-rich water from depths due to internal waves and tidal bores has been linked to growth rates of suspension feeders and benthic algae as well as plankton and larval organisms. The seaweed Codium, ''Codium isthmocladum'' reacts to deep water nutrient sources because their tissues have different concentrations of nutrients dependent upon depth. Aggregations of eggs, larval organisms and plankton on reefs respond to deep water intrusions. Similarly, as internal waves and bores move vertically, surface-dwelling larval organisms are carried toward the shore. This has significant biological importance to cascading effects of food chains in coral reef ecosystems and may provide yet another key to unlocking the paradox. Cyanobacteria provide soluble nitrates via nitrogen fixation. Coral reefs often depend on surrounding habitats, such as seagrass meadows and mangrove forests, for nutrients. Seagrass and mangroves supply dead plants and animals that are rich in nitrogen and serve to feed fish and animals from the reef by supplying wood and vegetation. Reefs, in turn, protect mangroves and seagrass from waves and produce sediment in which the mangroves and seagrass can root.

Biodiversity

Coral reefs form some of the world's most productive ecosystems, providing complex and varied marine habitats that support a wide range of other organisms. Fringing reefs just below low tide level have a mutually beneficial relationship with mangrove forests at high tide level and sea grass meadows in between: the reefs protect the mangroves and seagrass from strong currents and waves that would damage them or erosion, erode the sediments in which they are rooted, while the mangroves and sea grass protect the coral from large influxes of silt, fresh water and pollution, pollutants. This level of variety in the environment benefits many coral reef animals, which, for example, may feed in the sea grass and use the reefs for protection or breeding. Reefs are home to a variety of animals, including fish, seabirds, sponges, cnidarians (which includes some types of corals and jellyfish), worms, crustaceans (including shrimp, cleaner shrimp, spiny lobsters and crabs), mollusks (including cephalopods), echinoderms (including starfish, sea urchins and Holothuroidea, sea cucumbers), sea squirts, sea turtles and sea snakes. Aside from humans, mammals are rare on coral reefs, with visiting cetaceans such as dolphins the main exception. A few species feed directly on corals, while others graze on algae on the reef. Reef biomass is positively related to species diversity. The same hideouts in a reef may be regularly inhabited by different species at different times of day. Nighttime predators such as cardinalfish and squirrelfish hide during the day, while damselfish, surgeonfish, triggerfish, wrasses and parrotfish hide from eels and sharks. The great number and diversity of hiding places in coral reefs, i.e. Refuge (ecology), refuges, are the most important factor causing the great diversity and high Biomass (ecology), biomass of the organisms in coral reefs.

Algae

Reefs are chronically at risk of algal encroachment. Overfishing and excess nutrient supply from onshore can enable algae to outcompete and kill the coral. Increased nutrient levels can be a result of sewage or chemical fertilizer runoff. Runoff can carry nitrogen and phosphorus which promote excess algae growth. Algae can sometimes out-compete the coral for space. The algae can then smother the coral by decreasing the oxygen supply available to the reef. Decreased oxygen levels can slow down calcification rates, weakening the coral and leaving it more susceptible to disease and degradation. Algae inhabit a large percentage of surveyed coral locations. The algal population consists of turf algae,

and macroalgae, macro algae. Some sea urchins (such as ''Diadema antillarum'') eat these algae and could thus decrease the risk of algal encroachment.

Sponges

Sponges are essential for the functioning of the coral reef that system. Algae and corals in coral reefs produce organic material. This is filtered through sponges which convert this organic material into small particles which in turn are absorbed by algae and corals. Sponges are essential to the coral reef system however, they are quite different from corals. While corals are complex and many celled while sponges are very simple organisms with no tissue. They are alike in that they are both immobile aquatic vertebrates but otherwise are completely different. Types of sponges- There are several different species of sea sponge. They come in multiple shapes and sizes and all have unique characteristics. Some types of sea sponges include; the tube sponge, vase sponge, yellow sponge, bright red tree sponge, painted tunicate sponge, and the sea squirt sponge. Medicinal Qualities of Sea Sponges- Sea sponges have provided the base for many life saving medications. Scientists began to study them in the 1940's and after a few years, discovered that sea sponges contain properties that can stop viral infections. The first drug developed from sea sponges was released in 1969.

Fish

Over 4,000 species of fish inhabit coral reefs. The reasons for this diversity remain unclear. Hypotheses include the "lottery", in which the first (lucky winner) recruit to a territory is typically able to defend it against latecomers, "competition", in which adults compete for territory, and less-competitive species must be able to survive in poorer habitat, and "predation", in which population size is a function of postsettlement piscivore mortality. Healthy reefs can produce up to 35 tons of fish per square kilometer each year, but damaged reefs produce much less.

Invertebrates

Sea urchins, ''Dotidae'' and Nudibranch, sea slugs eat seaweed. Some species of sea urchins, such as ''Diadema antillarum'', can play a pivotal part in preventing algae from overrunning reefs. Researchers are investigating the use of native collector urchins, ''Collector urchin, Tripneustes gratilla'', for their potential as biocontrol agents to mitigate the spread of invasive algae species on coral reefs. ''Nudibranchia'' and sea anemones eat sponges. A number of invertebrates, collectively called "cryptofauna," inhabit the coral skeletal substrate itself, either boring into the skeletons (through the process of bioerosion) or living in pre-existing voids and crevices. Animals boring into the rock include sponges,

mollusks, and sipunculans. Those settling on the reef include many other species, particularly crustaceans and polychaete worms.

Seabirds

Coral reef systems provide important habitats for seabird species, some endangered. For example, Midway Atoll in Hawaii supports nearly three million seabirds, including two-thirds (1.5 million) of the global population of Laysan albatross, and one-third of the global population of black-footed albatross. Each seabird species has specific sites on the atoll where they nest. Altogether, 17 species of seabirds live on Midway. The short-tailed albatross is the rarest, with fewer than 2,200 surviving after excessive feather hunting in the late 19th century.

Other

Sea snakes feed exclusively on fish and their eggs. Marine birds, such as herons, gannets, pelicans and boobies, feed on reef fish. Some land-based reptiles intermittently associate with reefs, such as monitor lizards, the Crocodylus porosus, marine crocodile and semiaquatic snakes, such as ''Laticauda colubrina''. Sea turtles, particularly hawksbill sea turtles, feed on sponges. File:Prionurus laticlavius.jpg, Shoaling and schooling, Schooling reef fish File:Caribbean reef squid.jpg, Caribbean reef squid File:Stenopus hispidus (high res).jpg, Banded coral shrimp File:Triaenodon obesus moc.jpg, Whitetip reef shark File:Green turtle John Pennekamp.jpg, Green turtle File:Red sea-reef 3627.jpg, Giant clam File:Nephtheidae komodo.jpg, Soft coral, cup coral, sponges and ascidians File:Laticauda colubrina (Wakatobi).jpg, Banded sea krait File:Latiaxis wormaldi 002.jpg, The shell of ''Latiaxis wormaldi'', a coral snail

Ecosystem services

Coral reefs deliver

to tourism, fisheries and coastline protection. The global economic value of coral reefs has been estimated to be between US$29.8 billion (pdf
link
and $375 billion per year. About 500 million people benefit from ecosystem services provided by coral reefs. The economic cost over a 25-year period of destroying one square kilometer of coral reef has been estimated to be somewhere between $137,000 and $1,200,000. To improve the management of coastal coral reefs, the World Resources Institute (WRI) developed and published tools for calculating the value of coral reef-related tourism, shoreline protection and fisheries, partnering with five Caribbean countries. As of April 2011, published working papers covered St. Lucia, Tobago, Belize, and the Dominican Republic. The WRI was "making sure that the study results support improved coastal policies and management planning". The Belize study estimated the value of reef and mangrove services at $395–559 million annually. Bermuda's coral reefs provide economic benefits to the Island worth on average $722 million per year, based on six key ecosystem services, according to Sarkis ''et al'' (2010).

Shoreline protection

Coral reefs protect shorelines by absorbing wave energy, and many small islands would not exist without reefs. Coral reefs can reduce wave energy by 97%, 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. Reefs can attenuate waves as well as or better than artificial structures designed for coastal management, coastal defence such as breakwaters. An estimated 197 million people who live both below 10 m elevation and within 50 km of a reef consequently may receive risk reduction benefits from reefs. Restoring reefs is significantly cheaper than building artificial breakwaters in tropical environments. Expected damages from flooding would double, and costs from frequent storms would triple without the topmost meter of reefs. For 100-year storm events, flood damages would increase by 91% to $US 272 billion without the top meter.

Fisheries

About six million tons of fish are taken each year from coral reefs. Well-managed reefs have an average annual yield of 15 tons of seafood per square kilometer. Southeast Asia's coral reef fisheries alone yield about $2.4 billion annually from seafood.

Threats

Since their emergence 485 million years ago, coral reefs have faced many threats, including disease, predation, invasive species, bioerosion by grazing fish, algal blooms, and geologic hazards. Recent human activities present new threats. From 2009 to 2018, coral reefs worldwide declined 14%. Human activities that threaten coral include coral mining, bottom trawling, and the digging of canals and accesses into islands and bays, all of which can damage marine ecosystems if not done sustainably. Other localized threats include

, overfishing, coral overmining, and marine pollution, including use of the banned anti-fouling paint, anti-fouling biocide tributyltin; although absent in developed countries, these activities continue in places with few environmental protections or poor regulatory enforcement. Chemicals in sunscreens may awaken latent viral infections in zooxanthellae and impact reproduction. However, concentrating tourism activities via offshore platforms has been shown to limit the spread of coral disease by tourists. Greenhouse gas emissions present a broader threat through sea temperature rise and sea level rise, though corals homeostasis, adapt their calcifying fluids to changes in seawater pH and dissolved inorganic carbon, carbonate levels and are not directly threatened by ocean acidification. Volcanic and manmade aerosol pollution can modulate regional sea surface temperatures. In 2011, two researchers suggested that "extant marine invertebrates face the same synergistic effects of multiple stressors" that occurred during the end-Permian extinction, and that genera "with poorly buffered respiratory physiology and calcareous shells", such as corals, were particularly vulnerable. Corals respond to stress by "bleaching," or expelling their colorful zooxanthellate endosymbionts. Corals with Clade C zooxanthellae are generally vulnerable to heat-induced bleaching, whereas corals with the hardier Clade A or D are generally resistant, as are tougher coral genera like Porites and Montipora. Every 4–7 years, an El Niño event causes some reefs with heat-sensitive corals to bleach, with especially widespread bleachings in 1998 and 2010. However, reefs that experience a severe bleaching event become resistant to future heat-induced bleaching, due to rapid directional selection. Similar rapid adaption may protect coral reefs from global warming. A large-scale systematic study of the Jarvis Island coral community, which experienced ten El Niño-coincident coral bleaching events from 1960 to 2016, found that the reef recovered from almost complete death after severe events.

Protection

Marine Protected Area, Marine protected areas (MPAs) are areas designated because they provide various kinds of protection to ocean and/or estuarine areas. They are intended to promote responsible Fisheries management, fishery management and habitat (ecology), habitat protection. MPAs can also encompass social and biological objectives, including reef restoration, aesthetics, biodiversity and economic benefits. The effectiveness of MPAs is still debated. For example, a study investigating the success of a small number of MPAs in Indonesia, the Philippines and Papua New Guinea found no significant differences between the MPAs and unprotected sites. Furthermore, in some cases they can generate local conflict, due to a lack of community participation, clashing views of the government and fisheries, effectiveness of the area and funding. In some situations, as in the Phoenix Islands Protected Area, MPAs provide revenue to locals. The level of income provided is similar to the income they would have generated without controls. Overall, it appears the MPA's can provide protection to local coral reefs, but that clear management and sufficient funds are required. The Caribbean Coral Reefs - Status Report 1970–2012, states that coral decline may be reduced or even reversed. For this overfishing needs to be stopped, especially fishing on Coral reef fish#Herbivores, species key to coral reefs, such as Parrotfish#Economic importance, parrotfish. Direct human pressure on coral reefs should also be reduced and the inflow of sewage should be minimised. Measures to achieve this could include restricting coastal settlement, development and tourism. The report shows that healthier reefs in the Caribbean are those with large, healthy populations of parrotfish. These occur in countries that protect parrotfish and other species, like sea urchins. They also often ban fish trapping and spearfishing. Together these measures help creating "resilient reefs". Protecting networks of diverse and healthy reefs, not only climate Refugium (population biology), 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. Designating a reef as a biosphere reserve, marine park, national monument or world heritage site can offer protections. For example, Belize's barrier reef, Sian Ka'an, the Galapagos islands, Great Barrier Reef, Henderson Island (Pitcairn Islands), Henderson Island, Palau and Papahānaumokuākea Marine National Monument are world heritage sites. In Australia, the Great Barrier Reef is protected by the Great Barrier Reef Marine Park Authority, and is the subject of much legislation, including a biodiversity action plan. Australia compiled a Coral Reef Resilience Action Plan. This plan consists of adaptive management strategies, including reducing carbon footprint. A public awareness plan provides education on the "rainforests of the sea" and how people can reduce carbon emissions. Inhabitants of Ahus Island, Manus Province, Papua New Guinea, have followed a generations-old practice of restricting fishing in six areas of their reef lagoon. Their cultural traditions allow line fishing, but no net or spear fishing. Both biomass and individual fish sizes are significantly larger than in places where fishing is unrestricted. Increased levels of atmospheric CO₂ contribute to ocean acidification, which in turn damages coral reefs. To help combat ocean acidification, several countries have put laws in place to reduce greenhouse gases such as carbon dioxide. Many land use laws aim to reduce CO₂ emissions by limiting deforestation. Deforestation can release significant amounts of CO₂ absent sequestration via active follow-up forestry programs. Deforestation can also cause erosion, which flows into the ocean, contributing to ocean acidification. Incentives are used to reduce miles traveled by vehicles, which reduces carbon emissions into the atmosphere, thereby reducing the amount of dissolved CO₂ in the ocean. State and federal governments also regulate land activities that affect coastal erosion. High-end satellite technology can monitor reef conditions. The United States Clean Water Act puts pressure on state governments to monitor and limit run-off of polluted water.

Restoration

Coral reef restoration has grown in prominence over the past several decades because of the unprecedented reef die-offs around the planet. Coral stressors can include pollution, warming ocean temperatures, extreme weather events, and overfishing. With the deterioration of global reefs, fish nurseries, biodiversity, coastal development and livelihood, and natural beauty are under threat. Fortunately, researchers have taken it upon themselves to develop a new field, coral restoration, in the 1970s-1980s

Coral farming

NOAA_coral_nurseries_help_support_coral_restoration

Coral aquaculture, also known as coral farming or coral gardening, is showing promise as a potentially effective tool for restoring coral reefs. The "gardening" process bypasses the early growth stages of corals when they are most at risk of dying. Coral seeds are grown in nurseries, then replanted on the reef. Coral is farmed by coral farmers whose interests range from reef Conservation movement, conservation to increased income. Due to its straight forward process and substantial evidence of the technique having a significant effect on coral reef growth, coral nurseries became the most widespread and arguably the most effective method for coral restoration. Coral planting and reef restoration may 11

Coral planting and reef restoration may 11

Coral gardens take advantage of a coral's natural ability to fragment and continuing to grow if the fragments are able to anchor themselves onto new substrates. This method was first tested by Baruch Rinkevich in 1995 which found success at the time. By today's standards, coral farming has grown into a variety of different forms, but still has the same goals of cultivating corals. Consequently, coral farming quickly replaced previously used transplantation methods or the act of physically moving sections or whole colonies of corals into a new area. Transplantation has seen success in the past and decades of experiments have led to a high success and survival rate. However, this method still requires the removal of corals from existing reefs. With the current state of reefs, this kind of method should generally be avoided if possible. Saving healthy corals from eroding substrates or reefs that are doomed to collapse could be a major advantage of utilizing transplantation. Coral gardens generally take on the safe forms no matter where you go. It begins with the establishment of a nursery where operators can observe and care for coral fragments. It goes without saying that nurseries should be established in areas that are going to maximize growth and minimize mortality. Floating offshore coral trees or even aquariums are possible locations where corals can grow. After a location has been determined, collection and cultivation can occur. The major benefit of using coral farms is it lowers polyp and juvenile mortality rates. By removing predators and recruitment obstacles, corals are able to mature without much hindrance. However, nurseries cannot stop climate stressors. Warming temperatures or hurricanes can still disrupt or even kill nursery corals.

Creating substrates

Efforts to expand the size and number of coral reefs generally involve supplying substrate to allow more corals to find a home. Substrate materials include discarded vehicle tires, scuttled ships, subway cars and formed concrete, such as Reef Ball, reef balls. Reefs grow unaided on marine structures such as Oil platform, oil rigs. In large restoration projects, propagated

on substrate can be secured with metal pins, superglue or milliput. Needle and thread can also attach A-hermatype coral to substrate. Biorock is a substrate produced by a patented process that runs low voltage electrical currents through seawater to cause dissolved minerals to precipitate onto steel structures. The resultant white carbonate (

) is the same mineral that makes up natural coral reefs. Corals rapidly colonize and grow at accelerated rates on these coated structures. The electrical currents also accelerate the formation and growth of both chemical limestone rock and the skeletons of corals and other shell-bearing organisms, such as oysters. The vicinity of the anode and cathode provides a high-pH environment which inhibits the growth of competitive filamentous and fleshy algae. The increased growth rates fully depend on the accretion activity. Under the influence of the electric field, corals display an increased growth rate, size and density. Simply having many structures on the ocean floor is not enough to form coral reefs. Restoration projects must consider the complexity of the substrates they are creating for future reefs. Researchers conducted an experiment near Ticao Island in the Philippines in 2013 where several substrates in varying complexities were laid in the nearby degraded reefs. Large complexity consisted of plots that had both a man-made substrates of both smooth and rough rocks with a surrounding fence, medium consisted of only the man-made substrates, and small had neither the fence or substrates. After one month, researchers found that there was a positive correlation between structure complexity and recruitment rates of larvae. The medium complexity performed the best with larvae favoring rough rocks over smooth rocks. Following one year of their study, researchers visited the site and found that many of the sites were able to support local fisheries. They came to the conclusion that reef restoration could be done cost-effectively and will yield long term benefits given they are protected and maintained.

Relocation

One case study with coral reef restoration was conducted on the island of Oahu in Hawaii. The University of Hawaii operates a Coral Reef Assessment and Monitoring Program to help relocate and restore coral reefs in Hawaii. A boat channel from the island of Oahu to the Hawaiʻi Institute of Marine Biology, Hawaii Institute of Marine Biology on Coconut Island (Oahu Island), Coconut Island was overcrowded with coral reefs. Many areas of coral reef patches in the channel had been damaged from past dredging in the channel. Dredging covers corals with sand. Coral larvae cannot settle on sand; they can only build on existing reefs or compatible hard surfaces, such as rock or concrete. Because of this, the University decided to relocate some of the coral. They transplanted them with the help of United States Army divers, to a site relatively close to the channel. They observed little if any damage to any of the colonies during transport and no mortality of coral reefs was observed on the transplant site. While attaching the coral to the transplant site, they found that coral placed on hard rock grew well, including on the wires that attached the corals to the site. No environmental effects were seen from the transplantation process, recreational activities were not decreased, and no scenic areas were affected. As an alternative to transplanting coral themselves, juvenile fish can also be encouraged to relocate to existing coral reefs by auditory simulation. In damaged sections of the Great Barrier Reef, loudspeakers playing recordings of healthy reef environments were found to attract fish twice as often as equivalent patches where no sound was played, and also increased species biodiversity by 50%.

Heat-tolerant symbionts

Another possibility for coral restoration is gene therapy: inoculating coral with genetically modified bacterium, genetically modified bacteria, or naturally-occurring heat-tolerant varieties of coral symbiotes, may make it possible to grow corals that are more resistant to climate change and other threats. Warming oceans are forcing corals to adapt to unprecedented temperatures. Those that do not have a tolerance for the elevated temperatures experience coral bleaching and eventually mortality. There is already research that looks to create genetically modified corals that can withstand a warming ocean. Madeleine J. H. van Oppen, James K. Oliver, Hollie M. Putnam, and Ruth D. Gates described four different ways that gradually increase in human intervention to genetically modify corals. These methods focus on altering the genetics of the zooxanthellae within coral rather than the alternative. The first method is to induce acclimatization of the first generation of corals. The idea is that when adult and offspring corals are exposed to stressors, the zooxanthellae will gain a mutation. This method is based mostly on the chance that the zooxanthellae will acquire the specific trait that will allow it to better survive in warmer waters. The second method focuses on identifying what different kinds of zooxanthellae are within the coral and configuring how much of each zooxanthella lives within the coral at a given age. Use of zooxanthellae from the previous method would only boost success rates for this method. However, this method would only be applicable to younger corals, for now, because previous experiments of manipulation zooxanthellae communities at later life stages have all failed. The third method focuses on selective breeding tactics. Once selected, corals would be reared and exposed to simulated stressors in a laboratory. The last method is to genetically modify the zooxanthellae itself. When preferred mutations are acquired, the genetically modified zooxanthellae will be introduced to an aposymbiotic poly and a new coral will be produced. This method is the most laborious of the fourth, but researchers believe this method should be utilized more and holds the most promise in genetic engineering for coral restoration.

Invasive algae

Hawaiian coral reefs smothered by the spread of invasive algae were managed with a two-prong approach: divers manually removed invasive algae, with the support of super-sucker barges. Grazing pressure on invasive algae needed to be increased to prevent the regrowth of the algae. Researchers found that native collector urchins were reasonable candidate grazers for algae biocontrol, to extirpate the remaining invasive algae from the reef.

Invasive algae in Caribbean reefs

Macroalgae, or better known as seaweed, has to potential to cause reef collapse because they can outcompete many coral species. Macroalgae can overgrow on corals, shade, block recruitment, release biochemicals that can hinder spawning, and potentially form bacteria harmful to corals. Historically, algae growth was controlled by herbivorous fish and sea urchins. Parrotfish are a prime example of reef caretakers. Consequently, these two species can be considered as keystone species for reef environments because of their role in protecting reefs. Before the 1980s, Jamaica's reefs were thriving and well cared for, however, this all changed after Hurricane Allen occurred in 1980 and an unknown disease spread across the Caribbean. In the wake of these events, massive damage was caused to both the reefs and sea urchin population across Jamaican's reefs and into the Caribbean Sea. As little as 2% of the original sea urchin population survived the disease. Primary macroalgae succeeded the destroyed reefs and eventually larger, more resilient macroalgae soon took its place as the dominant organism. Parrotfish and other herbivorous fish were few in numbers because of decades of overfishing and bycatch at the time. Historically, the Jamaican coast had 90% coral cover and was reduced to 5% in the 1990s. Eventually, corals were able to recover in areas where sea urchin populations were increasing. Sea urchins were able to feed and multiply and clear off substrates, leaving areas for coral polyps to anchor and mature. However, sea urchin populations are still not recovering as fast as researchers predicted, despite being highly fecundate. It is unknown whether or not the mysterious disease is still present and preventing sea urchin populations from rebounding. Regardless, these areas are slowly recovering with the aid of sea urchin grazing. This event supports an early restoration idea of cultivating and releasing sea urchins into reefs to prevent algal overgrowth.

Microfragmentation and fusion

In 2014, Christopher Page, Erinn Muller, and David Vaughan from the International Center for Coral Reef Research & Restoration at Mote Marine Laboratory in Summerland Key, Florida developed a new technology called "microfragmentation," in which they use a specialized diamond band saw to cut corals into 1 cm² fragments instead of 6 cm² to advance the growth of brain, boulder, and star corals. Corals ''Orbicella faveolata'' and ''Montastraea cavernosa'' were outplanted off the Florida's shores in several microfragment arrays. After two years, ''O. faveolata'' had grown 6.5x its original size while ''M. cavernosa'' had grown nearly twice its size. Under conventional means, both corals would have required decades to reach the same size. It is suspected that if predation events had not occurred near the beginning of the experiment ''O. faveolata'' would have grown at least ten times its original size. By using this method, Mote Marine Laboratory produced 25,000 corals and planted 10,000 in the Florida Keys in only one year. Shortly after, they discovered that these microfragments fused with other microfragments from the same parent coral. Typically, corals that are not from the same parent fight and kill nearby corals in an attempt to survive and expand. This new technology is known as "fusion" and has been shown to grow coral heads in just two years instead of the typical 25–75 years. After fusion occurs, the reef will act as a single organism rather than several independent reefs. Currently, there has been no published research into this method.

History

The times of maximum reef development were in the Middle Cambrian (513–501 Annum, Ma), Devonian (416–359 Ma) and Carboniferous (359–299 Ma), owing to order Rugosa extinction, extinct corals and Late Cretaceous (100–66 Ma) and all Neogene (23 Ma–present), owing to Order (biology), order Scleractinia corals. Not all reefs in the past were formed by corals: those in the Early Cambrian (542–513 Ma) resulted from calcareous

and Archaeocyatha, archaeocyathids (small animals with conical shape, probably related to sponges) and in the Late Cretaceous (100–66 Ma), when reefs formed by a group of bivalves called

existed; one of the valves formed the main conical structure and the other, much smaller valve acted as a cap. Measurements of the oxygen isotopic composition of the aragonitic skeleton of coral reefs, such as ''Porites'', can indicate changes in sea surface temperature and sea surface salinity conditions during the growth of the coral. This technique is often used by climate scientists to infer a region's paleoclimate.

References

Further references

Coral Reef Protection: What Are Coral Reefs?
US EPA.

* [http://www.unepscs.org/SCS_Documents/Download/19_-_Technical_Publications_and_Guidelines/Technical_Publication_05_-_Coral_Reef_Demonstration_Sites_in_the_South_China_Sea.html UNEP. 2007. Coral Reefs Demonstration Sites in the South China Sea. UNEP/GEF/SCS Technical Publication No. 5.]
UNEP, 2007. National Reports on Coral Reefs in the Coastal Waters of the South China Sea. UNEP/GEF/SCS Technical Publication No. 11.

External links

*
Corals and Coral Reefs
overview at the Smithsonian Ocean Portal
About Corals
''Australian Institute of Marine Science''.
International Coral Reef InitiativeMoorea Coral Reef Long Term Ecological Research Site (US NSF)ARC Centre of Excellence for Coral Reef StudiesNOAA's Coral-List Listserver for Coral Reef Information and NewsNOAA's Coral Reef Conservation ProgramNOAA's Coral Reef Information SystemReefBase: A Global Information System on Coral ReefsNational Coral Reef Institute
Nova Southeastern University
Marine Aquarium CouncilNCORE National Center for Coral Reef Research
University of Miami
Science and Management of Coral Reefs in the South China Sea and Gulf of ThailandMicrodocs

Reef Relief
Active Florida environmental non-profit focusing on coral reef education and protection
Global Reef Record
– Catlin Seaview Survey of reef, a database of images and other information

(archived). Nancy Knowlton, ''iBioSeminars'', 2011.
Nancy Knowlton's Seminar: "Corals and Coral Reefs
. Nancy Knowlton, ''iBioSeminars'', 2011.
About coral reefs
Living Reefs Foundation, Bermuda
Caribbean Coral Reefs - Status Report 1970-2012
' by the International Union for Conservation of Nature, IUCN. - , featuring the report. {{Biomes Coral reefs, Animal products Environmental impact of fishing Coastal and oceanic landforms Ecosystems Oceanographical terminology Oceanography

Formation

Material

Types

Fringing reef

Barrier reef

Platform reef

Atoll

Other reef types or variants

Locations

Coral

Zooxanthellae

Skeleton

Reproduction

Gallery of reef-building corals

Other reef builders

Coralline algae

Sponges

Bivalves

Darwin's paradox

Explanations

Biodiversity

Algae

Sponges

Fish

Invertebrates

Seabirds

Other

Ecosystem services

Shoreline protection

Fisheries

Threats

Protection

Restoration

Coral farming

Creating substrates

Relocation

Heat-tolerant symbionts

Invasive algae

Invasive algae in Caribbean reefs

Microfragmentation and fusion

History

See also

References

Further references

External links