
A coral reef is an underwater
ecosystem
An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
characterized by reef-building
coral
Corals are colonial marine invertebrates within the subphylum Anthozoa of the phylum Cnidaria. They typically form compact Colony (biology), colonies of many identical individual polyp (zoology), polyps. Coral species include the important Coral ...
s. Reefs are formed of
colonies of
coral
Corals are colonial marine invertebrates within the subphylum Anthozoa of the phylum Cnidaria. They typically form compact Colony (biology), colonies of many identical individual polyp (zoology), polyps. Coral species include the important Coral ...
polyps held together by
calcium carbonate. Most coral reefs are built from
stony corals, whose polyps cluster in groups.
Coral belongs to the
class
Class, Classes, 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 d ...
Anthozoa
Anthozoa is one of the three subphyla of Cnidaria, along with Medusozoa and Endocnidozoa. It includes Sessility (motility), sessile marine invertebrates and invertebrates of brackish water, such as sea anemones, Scleractinia, stony corals, soft c ...
in the animal
phylum
In biology, a phylum (; : phyla) is a level of classification, or taxonomic rank, that is below Kingdom (biology), kingdom and above Class (biology), class. Traditionally, in botany the term division (taxonomy), division has been used instead ...
Cnidaria
Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
, which includes
sea anemone
Sea anemones ( ) are a group of predation, predatory marine invertebrates constituting the order (biology), order Actiniaria. Because of their colourful appearance, they are named after the ''Anemone'', a terrestrial flowering plant. Sea anemone ...
s and
jellyfish
Jellyfish, also known as sea jellies or simply jellies, are the #Life cycle, medusa-phase of certain gelatinous members of the subphylum Medusozoa, which is a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animal ...
. Unlike sea anemones, corals secrete hard carbonate
exoskeleton
An exoskeleton () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
s 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, displacing the microbial and
sponge reefs of the
Cambrian
The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
.
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
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
,
including
fish
A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
,
mollusk
Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The ...
s,
worms,
crustaceans
Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of Arthropod, arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquat ...
,
echinoderm
An echinoderm () is any animal of the phylum Echinodermata (), which includes starfish, brittle stars, sea urchins, sand dollars and sea cucumbers, as well as the sessile sea lilies or "stone lilies". While bilaterally symmetrical as ...
s,
sponges,
tunicates and other
cnidarians. 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.
Shallow tropical 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,
cyanide fishing,
spearfishing on
scuba), sunscreen use,
and harmful land-use practices, including
runoff and seeps (e.g., from
injection wells and cesspools).
Coral reefs deliver
ecosystem services 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).
Though the shallow water tropical coral reefs are best known, there are also deeper water reef-forming corals, which live in colder water and in temperate seas.
Formation
Most coral reefs were formed after the
Last Glacial Period when melting ice caused
sea level
Mean sea level (MSL, often shortened to sea level) is an mean, average surface level of one or more among Earth's coastal Body of water, bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical ...
to rise and flood
continental shelves. Most coral reefs are less than 10,000 years old. As communities established themselves, the reefs grew upwards, pacing rising
sea level
Mean sea level (MSL, often shortened to sea level) is an mean, average surface level of one or more among Earth's coastal Body of water, bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical ...
s. Reefs that rose too slowly could become drowned, without sufficient light. Coral reefs are also found in the deep sea away from
continental shelves, around
oceanic islands and
atoll
An atoll () is a ring-shaped island, including a coral rim that encircles a lagoon. There may be coral islands or cays on the rim. Atolls are located in warm tropical or subtropical parts of the oceans and seas where corals can develop. Most ...
s. The majority of these islands are
volcanic in origin. Others have
tectonic
Tectonics ( via Latin ) are the processes that result in the structure and properties of the Earth's crust and its evolution through time. The field of ''planetary tectonics'' extends the concept to other planets and moons.
These processes ...
origins where
plate movements lifted the deep ocean floor.
In ''
The Structure and Distribution of Coral Reefs'',
[Via Internet Archive]
/ref> Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
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 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 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 volcanic island 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'') an ...
inside.
File:Atoll forming-Atoll.png, Ultimately, the island sinks below the sea, and the barrier reef becomes an atoll
An atoll () is a ring-shaped island, including a coral rim that encircles a lagoon. There may be coral islands or cays on the rim. Atolls are located in warm tropical or subtropical parts of the oceans and seas where corals can develop. Most ...
enclosing an open lagoon.
Darwin predicted that underneath each 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'') an ...
would be a bedrock 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
The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
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.
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
The sea level has been rising from the end of the last ice age, which was around 20,000 years ago. Between 1901 and 2018, the average sea level rose by , with an increase of per year since the 1970s. This was faster than the sea level had e ...
, 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 to adequately photosynthesize, due to decreased light exposure.
The two main variables determining the geomorphology
Geomorphology () is the scientific study of the origin and evolution of topographic and bathymetric features generated by physical, chemical or biological processes operating at or near Earth's surface. Geomorphologists seek to understand wh ...
, 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
The Great Barrier Reef is the world's largest coral reef system, composed of over 2,900 individual reefs and 900 islands stretching for over over an area of approximately . The reef is located in the Coral Sea, off the coast of Queensland, ...
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 cays 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 and one of the three most common naturally occurring crystal forms of calcium carbonate (), the others being calcite and vaterite. It is formed by biological and physical processes, including precipitation fr ...
is formed. However, shell fragments and the remains of coralline algae such as the green-segmented genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
'' Halimeda'' 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.
In the geologic past
The times of maximum reef development were in the Middle Cambrian
Middle or The Middle may refer to:
* Centre (geometry), the point equally distant from the outer limits.
Places
* Middle (sheading), a subdivision of the Isle of Man
* Middle Bay (disambiguation)
* Middle Brook (disambiguation)
* Middle Creek (di ...
(513–501 Ma), Devonian
The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
(416–359 Ma) and Carboniferous
The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
(359–299 Ma), owing to extinct
Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
order Rugosa corals, and Late Cretaceous
The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
(100–66 Ma) and Neogene
The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...
(23 Ma–present), owing to order Scleractinia corals.
Not all reefs in the past were formed by corals: those in the Early Cambrian (542–513 Ma) resulted from calcareous algae
Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
and archaeocyathids (small animals with conical shape, probably related to sponges) and in the Late Cretaceous
The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
(100–66 Ma), when reefs formed by a group of bivalves called rudists 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.
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 lists "Four major forms of large-scale coral reefs" – the fringing reef, barrier reef, atoll and table reef based on Stoddart, D.R. (1969). Spalding ''et al.'' list four main reef types that can be clearly illustrated – the fringing reef, barrier reef, atoll, and "bank or platform reef"—and notes that many other structures exist which do not conform easily to strict definitions, including the "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 gravity, 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 ...
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, with outer regions pushed far out into the sea, the inner part is deepened by erosion and eventually forms a 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'') an ...
. 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
The Red Sea is a sea inlet of the Indian Ocean, lying between Africa and Asia. Its connection to the ocean is in the south, through the Bab-el-Mandeb Strait and the Gulf of Aden. To its north lie the Sinai Peninsula, the Gulf of Aqaba, and th ...
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 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'') an ...
.[ 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
The Great Barrier Reef is the world's largest coral reef system, composed of over 2,900 individual reefs and 900 islands stretching for over over an area of approximately . The reef is located in the Coral Sea, off the coast of Queensland, ...
.[''Types of Coral Reefs'']
at www.coral-reef-info.com. Retrieved 2 Feb 2018. Other major examples are the Mesoamerican Barrier Reef System and the New Caledonian Barrier Reef.[ Barrier reefs are also found on the coasts of Providencia,][ Mayotte, the ]Gambier Islands
The Gambier Islands ( or ) are an archipelago in French Polynesia, located at the southeast terminus of the Tuamotu archipelago. They cover an area of , and are made up of the Mangareva Islands, a group of high islands remnants of a caldera alo ...
, on the southeast coast of Kalimantan
Kalimantan (; ) is the Indonesian portion of the island of Borneo. It constitutes 73% of the island's area, and consists of the provinces of Central Kalimantan, East Kalimantan, North Kalimantan, South Kalimantan, and West Kalimantan. The non-Ind ...
, on parts of the coast of Sulawesi
Sulawesi ( ), also known as Celebes ( ), is an island in Indonesia. One of the four Greater Sunda Islands, and the List of islands by area, world's 11th-largest island, it is situated east of Borneo, west of the Maluku Islands, and south of Min ...
, southeastern New Guinea
New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
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 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 are typically situated within atolls, where they adopt the name "patch reefs" and often span a diameter of just a few dozen meters. In instances where platform reefs develop along elongated structures, such as old and weathered barrier reefs, they tend to arrange themselves in a linear formation. This is the case, for example, on the east coast of the ]Red Sea
The Red Sea is a sea inlet of the Indian Ocean, lying between Africa and Asia. Its connection to the ocean is in the south, through the Bab-el-Mandeb Strait and the Gulf of Aden. To its north lie the Sinai Peninsula, the Gulf of Aqaba, and th ...
near Jeddah
Jeddah ( ), alternatively transliterated as Jedda, Jiddah or Jidda ( ; , ), is a List of governorates of Saudi Arabia, governorate and the largest city in Mecca Province, Saudi Arabia, and the country's second largest city after Riyadh, located ...
. 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 are U-shaped, due to wind and water flow.
]
Atoll
Atolls or atoll reefs 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 Cook Islands, ]French Polynesia
French Polynesia ( ; ; ) is an overseas collectivity of France and its sole #Governance, overseas country. It comprises 121 geographically dispersed islands and atolls stretching over more than in the Pacific Ocean, South Pacific Ocean. The t ...
, the Marshall Islands and Micronesia.[
Atolls are found in the Indian Ocean, for example, in the ]Maldives
The Maldives, officially the Republic of Maldives, and historically known as the Maldive Islands, is an Archipelagic state, archipelagic country in South Asia located in the Indian Ocean. The Maldives is southwest of Sri Lanka and India, abou ...
, the Chagos Islands
The Chagos Archipelago (, ) or Chagos Islands (formerly , and later the Oil Islands) is a group of seven atolls comprising more than 60 islands in the Indian Ocean about south of the Maldives archipelago. This chain of islands is the southernmo ...
, the Seychelles
Seychelles (, ; ), officially the Republic of Seychelles (; Seychellois Creole: ), is an island country and archipelagic state consisting of 155 islands (as per the Constitution) in the Indian Ocean. Its capital and largest city, Victoria, ...
and around Cocos Island
Cocos Island () is a volcanic island in the Pacific Ocean administered by Costa Rica, approximately southwest of the Costa Rican mainland. It constitutes the 11th of the 15 districts of Puntarenas Canton of the Puntarenas Province, Province of ...
.[ 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 ]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'') an ...
or embayment, often circular and surrounded by sand or seagrass
Seagrasses are the only flowering plants which grow in marine (ocean), marine environments. There are about 60 species of fully marine seagrasses which belong to four Family (biology), families (Posidoniaceae, Zosteraceae, Hydrocharitaceae and ...
. 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.][
* Drying reef – a part of a reef which is above water at low tide but submerged at high tide
* Habili – reef specific to the ]Red Sea
The Red Sea is a sea inlet of the Indian Ocean, lying between Africa and Asia. Its connection to the ocean is in the south, through the Bab-el-Mandeb Strait and the Gulf of Aden. To its north lie the Sinai Peninsula, the Gulf of Aqaba, and th ...
; 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 Central Semitic languages, Central Semitic language of the Afroasiatic languages, Afroasiatic language family spoken primarily in the Arab world. The International Organization for Standardization (ISO) assigns lang ...
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
The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
sea level
Mean sea level (MSL, often shortened to sea level) is an mean, average surface level of one or more among Earth's coastal Body of water, bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical ...
s
* Cays – 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
The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is bounded by the cont ...
, Atlantic and Indian Ocean
The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or approximately 20% of the water area of Earth#Surface, Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia (continent), ...
s (including the Caribbean and on the Great Barrier Reef
The Great Barrier Reef is the world's largest coral reef system, composed of over 2,900 individual reefs and 900 islands stretching for over over an area of approximately . The reef is located in the Coral Sea, off the coast of Queensland, ...
and Belize Barrier Reef), where they provide habitable and agricultural land
* Seamount or guyot – formed when a coral reef on a volcanic island subsides; tops of seamounts are rounded and guyots are flat; flat tops of guyots, or ''tablemounts'', are due to erosion by waves, winds, and atmospheric processes
Zones
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
Seawater, or sea water, is water from a sea or ocean. On average, seawater in the world's oceans has a salinity of about 3.5% (35 g/L, 35 ppt, 600 mM). This means that every kilogram (roughly one liter by volume) of seawater has approximat ...
, sediment
Sediment is a solid material that is transported to a new location where it is deposited. It occurs naturally and, through the processes of weathering and erosion, is broken down and subsequently sediment transport, transported by the action of ...
s, 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 physical oceanography, 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 sur ...
does not occur, such as the Great Barrier Reef
The Great Barrier Reef is the world's largest coral reef system, composed of over 2,900 individual reefs and 900 islands stretching for over over an area of approximately . The reef is located in the Coral Sea, off the coast of Queensland, ...
. Others are found in the deep ocean surrounding islands or as atolls, such as in the Maldives
The Maldives, officially the Republic of Maldives, and historically known as the Maldive Islands, is an Archipelagic state, archipelagic country in South Asia located in the Indian Ocean. The Maldives is southwest of Sri Lanka and India, abou ...
. 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.
The reef surface is the shallowest part of the reef. It is subject to surge 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
Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
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 meadow
A seagrass meadow or seagrass bed is an underwater ecosystem formed by seagrasses. Seagrasses are marine (saltwater) plants found in shallow coastal waters and in the brackish waters of estuaries. Seagrasses are flowering plants with stems and ...
s 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
Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
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 algae provide complex habitats and areas that offer protection, such as cracks and crevices. Invertebrates and epiphytic
An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
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 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
The Indo-Pacific is a vast biogeographic region of Earth. In a narrow sense, sometimes known as the Indo-West Pacific or Indo-Pacific Asia, it comprises the tropical waters of the Indian Ocean, the western and central Pacific Ocean, and the ...
region (including the Red Sea
The Red Sea is a sea inlet of the Indian Ocean, lying between Africa and Asia. Its connection to the ocean is in the south, through the Bab-el-Mandeb Strait and the Gulf of Aden. To its north lie the Sinai Peninsula, the Gulf of Aqaba, and th ...
, Indian Ocean
The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or approximately 20% of the water area of Earth#Surface, Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia (continent), ...
, Southeast Asia
Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
and the Pacific
The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is bounded by the cont ...
) account for 91.9% of this total. Southeast Asia accounts for 32.3% of that figure, while the Pacific including Australia
Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
accounts for 40.8%. Atlantic and Caribbean
The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...
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 . When the net production by reef building corals no longer keeps pace with relative sea level and the reef structure permanently drowns a Darwin Point is reached. One such point exists at the northwestern end of the Hawaiian Archipelago; see Evolution of Hawaiian volcanoes#Coral atoll stage.
However, reefs in the Persian Gulf
The Persian Gulf, sometimes called the Arabian Gulf, is a Mediterranean seas, mediterranean sea in West Asia. The body of water is an extension of the Arabian Sea and the larger Indian Ocean located between Iran and the Arabian Peninsula.Un ...
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 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.
The northernmost coral reef on Earth is located near Eilat, Israel
Israel, officially the State of Israel, is a country in West Asia. It Borders of Israel, shares borders with Lebanon to the north, Syria to the north-east, Jordan to the east, Egypt to the south-west, and the Mediterranean Sea to the west. Isr ...
. Coral reefs are rare along the west coasts of the Americas
The Americas, sometimes collectively called America, are a landmass comprising the totality of North America and South America.''Webster's New World College Dictionary'', 2010 by Wiley Publishing, Inc., Cleveland, Ohio. When viewed as a sing ...
and Africa
Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
, due primarily to upwelling
Upwelling is an physical oceanography, 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 sur ...
and strong cold coastal currents that reduce water temperatures in these areas (the Humboldt, Benguela, and Canary Currents, 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
South Asia is the southern Subregion#Asia, subregion of Asia that is defined in both geographical and Ethnicity, ethnic-Culture, cultural terms. South Asia, with a population of 2.04 billion, contains a quarter (25%) of the world's populatio ...
—from the eastern tip of India (Chennai
Chennai, also known as Madras (List of renamed places in India#Tamil Nadu, its official name until 1996), is the capital city, capital and List of cities in Tamil Nadu by population, largest city of Tamil Nadu, the southernmost states and ...
) to the Bangladesh
Bangladesh, officially the People's Republic of Bangladesh, is a country in South Asia. It is the List of countries and dependencies by population, eighth-most populous country in the world and among the List of countries and dependencies by ...
and Myanmar
Myanmar, officially the Republic of the Union of Myanmar; and also referred to as Burma (the official English name until 1989), is a country in northwest Southeast Asia. It is the largest country by area in Mainland Southeast Asia and has ...
borders—as well as along the coasts of northeastern South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
and Bangladesh, due to the freshwater release from the Amazon
Amazon most often refers to:
* Amazon River, in South America
* Amazon rainforest, a rainforest covering most of the Amazon basin
* Amazon (company), an American multinational technology company
* Amazons, a tribe of female warriors in Greek myth ...
and Ganges Rivers respectively.
Significant coral reefs include:
* The Great Barrier Reef
The Great Barrier Reef is the world's largest coral reef system, composed of over 2,900 individual reefs and 900 islands stretching for over over an area of approximately . The reef is located in the Coral Sea, off the coast of Queensland, ...
—largest, comprising over 2,900 individual reefs and 900 islands stretching for over off Queensland, Australia
* The Mesoamerican Barrier Reef System—second largest, stretching from Isla Contoy at the tip of the Yucatán Peninsula 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 (, ) is the northernmost island of the Greece, 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 fruitful and ...
and Nassau
* The Red Sea
The Red Sea is a sea inlet of the Indian Ocean, lying between Africa and Asia. Its connection to the ocean is in the south, through the Bab-el-Mandeb Strait and the Gulf of Aden. To its north lie the Sinai Peninsula, the Gulf of Aqaba, and th ...
—includes 6,000-year-old fringing reefs located along a coastline
* The Florida Reef Tract—largest continental US reef and the third-largest coral barrier reef, extends from Soldier Key, located in Biscayne Bay, to the Dry Tortugas in the Gulf of Mexico
* Blake Plateau has the world's largest known deep-water coral reef, comprising a 6.4 million acre reef that stretches from Miami to Charleston, S. C. Its discovery was announced in January 2024.
* Pulley Ridge—deepest photosynthetic coral reef, Florida
Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
* Numerous reefs around the Maldives
The Maldives, officially the Republic of Maldives, and historically known as the Maldive Islands, is an Archipelagic state, archipelagic country in South Asia located in the Indian Ocean. The Maldives is southwest of Sri Lanka and India, abou ...
* The Philippines
The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
coral reef area, the second-largest in Southeast Asia, is estimated at 26,000 square kilometres. 915 reef fish species and more than 400 scleractinian coral species, 12 of which are endemic are found there.
* The Raja Ampat Islands in Indonesia
Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania, between the Indian Ocean, Indian and Pacific Ocean, Pacific oceans. Comprising over List of islands of Indonesia, 17,000 islands, including Sumatra, ...
's Southwest Papua province offer the highest known marine diversity.
* Bermuda
Bermuda is a British Overseas Territories, British Overseas Territory in the Atlantic Ocean, North Atlantic Ocean. The closest land outside the territory is in the American state of North Carolina, about to the west-northwest.
Bermuda is an ...
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 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 United States, then veers east near 36°N latitude (North Carolin ...
. Bermuda coral species represent a subset of those found in the greater Caribbean.
* The world's northernmost individual coral reef is located in the Finlayson Channel, in the inside passage of British Columbia, Canada.
* The world's southernmost coral reef is at Lord Howe Island
Lord Howe Island (; formerly Lord Howe's Island) is an irregularly crescent-shaped volcanic remnant in the Tasman Sea between Australia and New Zealand, part of the Australian state of New South Wales. It lies directly east of mainland Port ...
, in the Pacific Ocean off the east coast of Australia.
Coral
When alive, corals are colonies 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 corals live only in the photic zone (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 ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
( dinoflagellates) of the genus '' Symbiodinium'', 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
Glucose is a sugar with the Chemical formula#Molecular formula, molecular formula , which is often abbreviated as Glc. It is overall the most abundant monosaccharide, a subcategory of carbohydrates. It is mainly made by plants and most algae d ...
, glycerol
Glycerol () is a simple triol compound. It is a colorless, odorless, sweet-tasting, viscous liquid. The glycerol backbone is found in lipids known as glycerides. It is also widely used as a sweetener in the food industry and as a humectant in pha ...
and amino acids
Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the Proteinogenic amino acid, 22 α-amino acids incorporated into p ...
. 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 centimetre of coral, and provide a constant supply of the carbon dioxide
Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
they need for photosynthesis.
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 clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s of ''Symbiodinium'' phylotypes. 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 acids that are UV resistant, using a derivative of glycerin 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 urchin
Sea urchins or urchins () are echinoderms in the class (biology), class Echinoidea. About 950 species live on the seabed, inhabiting all oceans and depth zones from the intertidal zone to deep seas of . They typically have a globular body cove ...
s, 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 species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
are named by their resemblance to terrestrial objects such as wrinkled brains, cabbages, table tops, antlers, 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, 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 species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
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. 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 coralline algae, some sponges and bivalve
Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
s. Reefs are always built by the combined efforts of these different phyla
Phyla, the plural of ''phylum'', may refer to:
* Phylum, a biological taxon between Kingdom and Class
* by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another
Phy ...
, 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 geologic record, rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating stratum, strata ...
s.
Coralline algae
Coralline algae 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. Furthermore, in locations unfavorable to the growth of corals, coralline algae can be the primary builders of an algal reef.
Sponges
Sponge reefs are reefs produced by sea sponges. Hexactinellid sponges are known to form reefs off the coast of British Columbia
British Columbia is the westernmost Provinces and territories of Canada, province of Canada. Situated in the Pacific Northwest between the Pacific Ocean and the Rocky Mountains, the province has a diverse geography, with rugged landscapes that ...
, southeast Alaska
Alaska ( ) is a non-contiguous U.S. state on the northwest extremity of North America. Part of the Western United States region, it is one of the two non-contiguous U.S. states, alongside Hawaii. Alaska is also considered to be the north ...
and Washington state. Reefs discovered in Hecate Strait, British Columbia, have grown to up to 7 kilometres long and 20 metres high. Hexactinellid sponge reefs were first identified in the Middle Triassic
The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
(245–208 million years ago). The sponges reached their full extent in the late Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
(201–145 million years ago) when a discontinuous reef system 7,000 km long stretched across the northern Tethys and North Atlantic basins, but have since declined and were thought to be extinct until existing reefs were discovered in 1987–1988.
Archaeocyatha, an extinct clade of sponges, were the planet's first reef-building animals and are an index fossil for the Lower Cambrian worldwide. Similarly, Stromatoporoidea were another extinct clade of reef-building sponges. Unlike corals, stromatoporoids usually settled on soft substrates, so their ‘reefs’ occupied only a single level rather than a multi-tiered vertical framework of built-up skeletons.
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'' were once abundant in Chesapeake Bay and shorelines bordering the Atlantic coastal plain
The Atlantic Ocean is the second largest of the world's five borders of the oceans, oceanic divisions, with an area of about . It covers approximately 17% of Earth#Surface, Earth's surface and about 24% of its water surface area. During the ...
until the late nineteenth century. '' Ostrea angasi'' is a species of flat oyster that had also formed large reefs in South Australia.
Hippuritida, an extinct order of bivalves known as rudists, were major reef-building organisms during the Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
. 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.
Gastropods
Some gastropods, like family Vermetidae, are sessile and cement themselves to the substrate, contributing to the reef building.
Darwin's paradox
In '' The Structure and Distribution of Coral Reefs'', 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 plant
Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
s, 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.
One reason for the unusual clarity of tropical waters is their nutrient deficiency and drifting plankton
Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
. 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
Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...
(N), phosphorus
Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...
(P) and potassium
Potassium is a chemical element; it has Symbol (chemistry), symbol K (from Neo-Latin ) and atomic number19. It is a silvery white metal that is soft enough to easily cut with a knife. Potassium metal reacts rapidly with atmospheric oxygen to ...
(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
Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
, which powers carbon fixation
Biological carbon fixation, or сarbon assimilation, is the Biological process, process by which living organisms convert Total inorganic carbon, inorganic carbon (particularly carbon dioxide, ) to Organic compound, organic compounds. These o ...
, 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 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.]
Sponges live in crevices in the reefs. They are efficient filter feeders, and in the Red Sea
The Red Sea is a sea inlet of the Indian Ocean, lying between Africa and Asia. Its connection to the ocean is in the south, through the Bab-el-Mandeb Strait and the Gulf of Aden. To its north lie the Sinai Peninsula, the Gulf of Aqaba, and th ...
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.
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.
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 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
Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...
provide soluble nitrate
Nitrate is a polyatomic ion with the chemical formula . salt (chemistry), Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are solubility, soluble in wa ...
s via nitrogen fixation.
Coral reefs often depend on surrounding habitats, such as seagrass meadow
A seagrass meadow or seagrass bed is an underwater ecosystem formed by seagrasses. Seagrasses are marine (saltwater) plants found in shallow coastal waters and in the brackish waters of estuaries. Seagrasses are flowering plants with stems and ...
s 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
Sediment is a solid material that is transported to a new location where it is deposited. It occurs naturally and, through the processes of weathering and erosion, is broken down and subsequently sediment transport, transported by the action of ...
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 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 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 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, seabird
Seabirds (also known as marine birds) are birds that are adaptation, adapted to life within the marine ecosystem, marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent ...
s, sponges, cnidaria
Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
ns (which includes some types of corals and jellyfish
Jellyfish, also known as sea jellies or simply jellies, are the #Life cycle, medusa-phase of certain gelatinous members of the subphylum Medusozoa, which is a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animal ...
), worms, crustaceans (including shrimp, cleaner shrimp
Cleaner shrimp is a common name for a number of swimming Decapoda, decapod crustaceans that clean other organisms of parasites. Most are found in the family (biology), families Hippolytidae (including the Pacific cleaner shrimp, ''Lysmata amboi ...
, spiny lobsters and crab
Crabs are decapod crustaceans of the infraorder Brachyura (meaning "short tailed" in Greek language, Greek), which typically have a very short projecting tail-like abdomen#Arthropoda, abdomen, usually hidden entirely under the Thorax (arthropo ...
s), mollusk
Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The ...
s (including cephalopod
A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
s), echinoderm
An echinoderm () is any animal of the phylum Echinodermata (), which includes starfish, brittle stars, sea urchins, sand dollars and sea cucumbers, as well as the sessile sea lilies or "stone lilies". While bilaterally symmetrical as ...
s (including starfish
Starfish or sea stars are Star polygon, star-shaped echinoderms belonging to the class (biology), class Asteroidea (). Common usage frequently finds these names being also applied to brittle star, ophiuroids, which are correctly referred to ...
, sea urchin
Sea urchins or urchins () are echinoderms in the class (biology), class Echinoidea. About 950 species live on the seabed, inhabiting all oceans and depth zones from the intertidal zone to deep seas of . They typically have a globular body cove ...
s and sea cucumbers), sea squirts, sea turtles and sea snake
Sea snakes, or coral reef snakes, are Elapidae, elapid snakes that inhabit Marine (ocean), marine environments for most or all of their lives. They belong to two subfamilies, Hydrophiinae and Sea krait, Laticaudinae. Hydrophiinae also includes ...
s. Aside from humans, mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s are rare on coral reefs, with visiting cetacean
Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
s such as dolphins being 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 shark
Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
s.[
The great number and diversity of hiding places in coral reefs, i.e. refuges, are the most important factor causing the great diversity and high biomass of the organisms in coral reefs.]
Coral reefs also have a very high degree of microorganism diversity compared to other environments.
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, coralline algae and macro algae. Some sea urchins (such as '' Diadema antillarum'') eat these algae and could thus decrease the risk of algal encroachment.
Sponges
Sea sponges are an important component of coral reef communities. There are 420 species of sponges in coral reefs from Indonesia, 486 species in coral reefs from Indian waters, and 1500 species in the Great Barrier Reef
The Great Barrier Reef is the world's largest coral reef system, composed of over 2,900 individual reefs and 900 islands stretching for over over an area of approximately . The reef is located in the Coral Sea, off the coast of Queensland, ...
from Australia.
Sponges occupy an important role as detritivores in coral reef food webs by recycling detritus to higher trophic levels through their sponge loop.[ Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License ] For example, several sponge species are able to convert dissolved organic matter (DOM) derived from corals and algae into sponge detritus, which serves as food for species incapable of directly consuming DOM.
Sponges with photosynthesizing endosymbiont
An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...
s also produce up to three times more oxygen
Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...
, as well as more organic matter than they consume. Such contributions to their habitats' resources are significant along Australia's Great Barrier Reef but relatively minor in the Caribbeans.
Aside from producing nutrition, sponges also provide microhabitats for various invertebrates, as well as some fish species.
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 kilometre each year, but damaged reefs produce much less.
Invertebrates
Sea urchins, '' Dotidae'' and 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, '' Tripneustes gratilla'', for their potential as biocontrol agents to mitigate the spread of invasive algae species on coral reefs. '' Nudibranchia'' and sea anemone
Sea anemones ( ) are a group of predation, predatory marine invertebrates constituting the order (biology), order Actiniaria. Because of their colourful appearance, they are named after the ''Anemone'', a terrestrial flowering plant. Sea anemone ...
s 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, bivalve
Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
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
Seabirds (also known as marine birds) are birds that are adaptation, adapted to life within the marine ecosystem, marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent ...
species, some endangered. For example, Midway Atoll in Hawaii
Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
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 snake
Sea snakes, or coral reef snakes, are Elapidae, elapid snakes that inhabit Marine (ocean), marine environments for most or all of their lives. They belong to two subfamilies, Hydrophiinae and Sea krait, Laticaudinae. Hydrophiinae also includes ...
s feed exclusively on fish and their eggs. Marine birds, such as heron
Herons are long-legged, long-necked, freshwater and coastal birds in the family Ardeidae, with 75 recognised species, some of which are referred to as egrets or bitterns rather than herons. Members of the genus ''Botaurus'' are referred to as bi ...
s, gannets, pelicans and boobies, feed on reef fish. Some land-based reptile
Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s intermittently associate with reefs, such as monitor lizards, the marine crocodile and semiaquatic snakes, such as '' Laticauda colubrina''. Sea turtles, particularly hawksbill sea turtles, feed on sponges.
File:Prionurus laticlavius.jpg, 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 ecosystem services 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 kilometre 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
Tobago, officially the Ward of Tobago, is an List of islands of Trinidad and Tobago, island and Regions and municipalities of Trinidad and Tobago, ward within the Republic of Trinidad and Tobago. It is located northeast of the larger islan ...
, Belize
Belize is a country on the north-eastern coast of Central America. It is bordered by Mexico to the north, the Caribbean Sea to the east, and Guatemala to the west and south. It also shares a maritime boundary with Honduras to the southeast. P ...
, and the Dominican Republic
The Dominican Republic is a country located on the island of Hispaniola in the Greater Antilles of the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean. It shares a Maritime boundary, maritime border with Puerto Rico to the east and ...
. 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 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 kilometre. 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 canal
Canals or artificial waterways are waterways or engineered channels built for drainage management (e.g. flood control and irrigation) or for conveyancing water transport vehicles (e.g. water taxi). They carry free, calm surface ...
s and accesses into islands and bays, all of which can damage marine ecosystems if not done sustainably. Other localized threats include blast fishing, overfishing
Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing Fish stocks, fish stock), resu ...
, coral overmining, and marine pollution
Marine pollution occurs when substances used or spread by humans, such as industrial waste, industrial, agricultural pollution, agricultural, and municipal solid waste, residential waste; particle (ecology), particles; noise; excess carbon dioxi ...
, including use of the banned 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, resulting in widespread coral bleaching and loss of coral cover. Climate change
Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
causes more frequent and more severe storms, also changes ocean circulation patterns, which can destroy coral reefs. Ocean acidification also affects corals by decreasing calcification rates and increasing dissolution rates, although corals can adapt their calcifying fluids to changes in seawater
Seawater, or sea water, is water from a sea or ocean. On average, seawater in the world's oceans has a salinity of about 3.5% (35 g/L, 35 ppt, 600 mM). This means that every kilogram (roughly one liter by volume) of seawater has approximat ...
pH and carbonate levels to mitigate the impact. Volcanic and human-made 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 endosymbiont
An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...
s. 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 areas (MPAs) are areas designated because they provide various kinds of protection to ocean and/or estuarine areas. They are intended to promote responsible fishery management and 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
Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania, between the Indian Ocean, Indian and Pacific Ocean, Pacific oceans. Comprising over List of islands of Indonesia, 17,000 islands, including Sumatra, ...
, the Philippines
The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
and Papua New Guinea
Papua New Guinea, officially the Independent State of Papua New Guinea, is an island country in Oceania that comprises the eastern half of the island of New Guinea and offshore islands in Melanesia, a region of the southwestern Pacific Ocean n ...
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
Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing Fish stocks, fish stock), resu ...
needs to be stopped, especially fishing on species key to coral reefs, such as 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
Tourism is travel for pleasure, and the Commerce, commercial activity of providing and supporting such travel. World Tourism Organization, UN Tourism defines tourism more generally, in terms which go "beyond the common perception of tourism as ...
. 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 refugia, helps ensure the greatest chance of genetic diversity
Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species. It ranges widely, from the number of species to differences within species, and can be correlated to the span of survival for a species. It is d ...
, 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
The Great Barrier Reef is the world's largest coral reef system, composed of over 2,900 individual reefs and 900 islands stretching for over over an area of approximately . The reef is located in the Coral Sea, off the coast of Queensland, ...
, Henderson Island, Palau
Palau, officially the Republic of Palau, is an island country in the Micronesia subregion of Oceania in the western Pacific Ocean. The Republic of Palau consists of approximately 340 islands and is the western part of the Caroline Islands ...
and Papahānaumokuākea Marine National Monument
The Papahānaumokuākea Marine National Monument (PMNM) (roughly ) is a World Heritage Site, World Heritage listed National monument (United States), U.S. national monument encompassing of ocean waters, including ten islands and atolls of th ...
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
Papua New Guinea, officially the Independent State of Papua New Guinea, is an island country in Oceania that comprises the eastern half of the island of New Guinea and offshore islands in Melanesia, a region of the southwestern Pacific Ocean n ...
, 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
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 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 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.
Technology is becoming more popular in the coral farming process. Teams from the Reef Restoration and Adaptation Program (RRAP) have trialled coral counting technology utilizing a prototype robotic camera. The camera uses computer vision
Computer vision tasks include methods for image sensor, acquiring, Image processing, processing, Image analysis, analyzing, and understanding digital images, and extraction of high-dimensional data from the real world in order to produce numerical ...
and learning algorithms to detect and count individual coral babies and track their growth and health in real time. This technology, with research led by QUT, is intended to be used during annual coral spawning events and will provide researchers with control that is not currently possible when mass-producing 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 balls. Reefs grow unaided on marine structures such as oil rigs. In large restoration projects, propagated hermatypic coral 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
Steel is an alloy of iron and carbon that demonstrates improved mechanical properties compared to the pure form of iron. Due to steel's high Young's modulus, elastic modulus, Yield (engineering), yield strength, Fracture, fracture strength a ...
structures. The resultant white carbonate (aragonite
Aragonite is a carbonate mineral and one of the three most common naturally occurring crystal forms of calcium carbonate (), the others being calcite and vaterite. It is formed by biological and physical processes, including precipitation fr ...
) 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
An anode usually is an electrode of a polarized electrical device through which conventional current enters the device. This contrasts with a cathode, which is usually an electrode of the device through which conventional current leaves the devic ...
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 human-made substrates of both smooth and rough rocks with a surrounding fence, medium consisted of only the human-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
Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
. The University of Hawaii
A university () is an educational institution, institution of tertiary education and research which awards academic degrees in several Discipline (academia), academic disciplines. ''University'' is derived from the Latin phrase , which roughly ...
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 Hawaii Institute of Marine Biology on 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
The United States Army (USA) is the primary Land warfare, land service branch of the United States Department of Defense. It is designated as the Army of the United States in the United States Constitution.Article II, section 2, clause 1 of th ...
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 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 successfully generated 25,000 corals within a single year, subsequently transplanting 10,000 of them into the Florida Keys. 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.
See also
* Deep-water coral — Corals living in the cold waters of deeper, darker parts of the oceans
* Mesophotic coral reef — Corals living in the mesopelagic or twilight zone
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References
Further references
Coral Reef Protection: What Are Coral Reefs?
US EPA.
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External links
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Corals and Coral Reefs
overview at the Smithsonian Ocean Portal
About Corals
''Australian Institute of Marine Science''.
International Coral Reef Initiative
Moorea Coral Reef Long Term Ecological Research Site (US NSF)
ARC Centre of Excellence for Coral Reef Studies
NOAA's Coral-List Listserver for Coral Reef Information and News
NOAA's Coral Reef Conservation Program
NOAA's Coral Reef Information System
ReefBase: A Global Information System on Coral Reefs
National Coral Reef Institute
Nova Southeastern University
Marine Aquarium Council
NCORE National Center for Coral Reef Research
University of Miami
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Microdocs
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 IUCN
The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the status ...
. – , featuring the report.
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Animal products
Environmental impact of fishing
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Oceanographical terminology
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