coral
   HOME

TheInfoList



OR:

Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton. A coral "group" is a colony of very many genetically identical polyps. Each polyp is a sac-like animal typically only a few millimeters in diameter and a few centimeters in height. A set of
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic ...
s surround a central mouth opening. Each polyp excretes an exoskeleton near the base. Over many generations, the colony thus creates a skeleton characteristic of the species which can measure up to several meters in size. Individual colonies grow by asexual reproduction of polyps. Corals also breed sexually by spawning: polyps of the same species release gametes simultaneously overnight, often around a
full moon The full moon is the lunar phase when the Moon appears fully illuminated from Earth's perspective. This occurs when Earth is located between the Sun and the Moon (when the ecliptic longitudes of the Sun and Moon differ by 180°). This me ...
. Fertilized eggs form planulae, a mobile early form of the coral polyp which, when mature, settles to form a new colony. Although some corals are able to catch
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind Wind is the natural movement of air or other gases relative to a planet's surface. Winds occu ...
and small
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% ...
using stinging cells on their tentacles, most corals obtain the majority of their energy and nutrients from photosynthetic unicellular dinoflagellates of the genus '' Symbiodinium'' that live within their tissues. These are commonly known as zooxanthellae and give the coral color. Such corals require sunlight and grow in clear, shallow water, typically at depths less than . Corals are major contributors to the physical structure of the
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in groups ...
s that develop in tropical and subtropical waters, 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 ...
off the coast of
Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country b ...
. These corals are increasingly at risk of bleaching events where polyps expel the zooxanthellae in response to stress such as high water temperature or toxins. Other corals do not rely on zooxanthellae and can live globally in much deeper water, such as the cold-water
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as virus A virus is a submicroscopic infectious agent that replicates only inside the living cells of an o ...
'' Lophelia'' which can survive as deep as . Some have been found as far north as the Darwin Mounds, northwest of
Cape Wrath Cape Wrath ( gd, Am Parbh, known as ' in Lewis) is a cape in the Durness parish of the county of Sutherland in the Highlands of Scotland Scotland (, ) is a country that is part of the United Kingdom The United Kingdom of G ...
, Scotland, and others off the coast of Washington state and the Aleutian Islands.


Taxonomy

The classification of corals has been discussed for millennia, owing to having similarities to both plants and animals.
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
's pupil Theophrastus described the red coral, ''korallion'', in his book on stones, implying it was a mineral, but he described it as a deep-sea plant in his ''Enquiries on Plants'', where he also mentions large stony plants that reveal bright flowers when under water in the Gulf of Heroes.
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire The Roman Empire ( la, Imperium Romanum ; grc-gre, Βασ ...
stated boldly that several sea creatures including sea nettles and sponges "are neither animals nor plants, but are possessed of a third nature (''tertia natura'')". Petrus Gyllius copied Pliny, introducing the term ''zoophyta'' for this third group in his 1535 book ''On the French and Latin Names of the Fishes of the Marseilles Region''; it is popularly but wrongly supposed that Aristotle created the term. Gyllius further noted, following Aristotle, how hard it was to define what was a plant and what was an animal. The
Babylonian Talmud The Talmud (; he, , Talmūḏ) is the central text of Rabbinic Judaism and the primary source of Jewish religious law (''halakha ''Halakha'' (; he, הֲלָכָה, ), also transliterated as ''halacha'', ''halakhah'', and ''halocho'' ( ) ...
refers to coral among a list of types of trees, and the 11th century French commentator
Rashi Shlomo Yitzchaki ( he, רבי שלמה יצחקי; la, Salomon Isaacides; french: Salomon de Troyes, 22 February 1040 – 13 July 1105), today generally known by the acronym Rashi (see below), was a medieval French rabbi and author of a comp ...
describes it as "a type of tree (מין עץ) that grows underwater that goes by the (French) name "coral." The Persian polymath
Al-Biruni Abu Rayhan Muhammad ibn Ahmad al-Biruni (973 – after 1050) commonly known as al-Biruni, was a Khwarazmian Iranian in scholar and polymath during the Islamic Golden Age. He has been called variously the "founder of Indology", "Father ...
(d.1048) classified sponges and corals as animals, arguing that they respond to touch. Nevertheless, people believed corals to be plants until the eighteenth century, when William Herschel used a microscope to establish that coral had the characteristic thin cell membranes of an
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
. Presently, corals are classified as species of animals within the sub-classes Hexacorallia and Octocorallia of the class Anthozoa in the phylum Cnidaria. Hexacorallia includes the stony corals and these groups have polyps that generally have a 6-fold symmetry. Octocorallia includes blue coral and soft corals and species of Octocorallia have polyps with an eightfold symmetry, each polyp having eight tentacles and eight mesenteries. The group of corals is paraphyletic because the sea anemones are also in the sub-class Hexacorallia.


Systematics

Th
delineation
of coral species is challenging as hypotheses based on morphological traits contradict hypotheses formed via molecular tree-based processes.   As of 2020, there are 2175 identified separate coral species, 237 of which are currently endangered, making distinguishing corals to be the utmost of importance in efforts to curb extinction.  Adaptation and delineation continues to occur in species of coral in order to combat the dangers posed by the climate crisis. Corals ar
colonial modular organisms
formed by asexually produced and genetically identical modules calle
polyps
Polyps are connected by living tissue to produce the full organism.  The living tissue allows for inter module communication (interaction between each polyp), which appears in colony morphologies produced by corals, and is one of the main identifying characteristics for a species of coral.   There are 2 main classifications for corals: 1. Hard coral (scleractinian and stony coral) which form reefs by a calcium carbonate base, with polyps with 6 stiff tentacles, and 2. Soft coral (Alcyonacea and ahermatypic coral) which are bendable and formed by a colony of polyps with 8 feather like tentacles.  These two classifications arose fro
differentiation in gene expressions
in their branch tips and bases that arose through developmental signaling pathways such as Hox, Hedgehog, Wnt
BMP
etc. Scientists typically select ''Acropora'' as research models since they are the most diverse genus of hard coral, having over 120 species.  Most species within this genus have polyps which ar
axial
polyps grow rapidly and have lighter coloration, whil
radial
polyps are small and are darker in coloration. In the ''Acropora'' genus, gamete synthesis and
photosynthesis Photosynthesis is a process used by plant Plants are predominantly photosynthetic eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. Th ...
occur at th
basal
polyps, growth occurs mainly at the radial polyps. Growth at the site of the radial polyps encompasses two processes: asexual reproduction vi
mitotic cell proliferation
and skeleton deposition of the calcium carbonate via
extra cellular matrix In biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that pro ...
(EMC) proteins acting a
differentially expressed (DE) signaling genes
ref name=":2" /> between both branch tips and bases.  These processes lead t
colony differentiation
which is the most accurate distinguisher between coral species. In the Acropora genus, colony differentiation through up-regulation and down-regulation of DEs. Systematic studies of soft coral species have faced challenges due to a lack of taxonomic knowledge.  Researchers have not found enough variability within the genus to confidently delineate similar species, due to a low rate in mutation of mitochondrial DNA. Environmental factors, such as the rise of temperatures and acid levels in our oceans account for some speciation of corals in the form of species lost.  Various coral species have
heat shock protein Heat shock proteins (HSP) are a family of protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including cat ...
s (HSP) that are also in the category of DE across species.  These HSPs help corals combat the increased temperatures they are facing which lead to protein denaturing, growth loss, and eventually coral death.  Approximately 33% of coral species are on the International Union for Conservation of Nature’s endangered species list and at risk of species loss.  Ocean acidification (falling pH levels in the oceans) is threatening the continued species growth and differentiation of corals.  Mutation rates of '' Vibrio shilonii'', the reef pathogen responsible for coral bleaching, heavily outweigh the  typical reproduction rates of coral colonies when pH levels fall. Thus, corals are unable to mutate their HSPs and other climate change preventative genes to combat the increase in temperature and decrease in pH at a competitive rate to these pathogens responsible for coral bleaching, resulting in species loss.


Anatomy

For most of their life corals are sessile animals of colonies of genetically identical polyps. Each polyp varies from millimeters to centimeters in diameter, and colonies can be formed from many millions of individual polyps. Stony coral, also known as hard coral, polyps produce a skeleton composed of calcium carbonate to strengthen and protect the organism. This is deposited by the polyps and by the coenosarc, the living tissue that connects them. The polyps sit in cup-shaped depressions in the skeleton known as corallites. Colonies of stony coral are markedly variable in appearance; a single species may adopt an encrusting, plate-like, bushy, columnar or massive solid structure, the various forms often being linked to different types of habitat, with variations in light level and water movement being significant. The body of the polyp may be roughly compared in a structure to a sac, the wall of which is composed of two layers of cells. The outer layer is known technically as the
ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastr ...
, the inner layer as the endoderm. Between ectoderm and endoderm is a supporting layer of gelatinous substance termed mesoglea, secreted by the cell layers of the body wall. The mesoglea can contain skeletal elements derived from cells migrated from the ectoderm. The sac-like body built up in this way is attached to a hard surface, which in hard corals are cup-shaped depressions in the skeleton known as corallites. At the center of the upper end of the sac lies the only opening called the mouth, surrounded by a circle of
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic ...
s which resemble glove fingers. The tentacles are organs which serve both for tactile sense and for the capture of food. Polyps extend their tentacles, particularly at night, often containing coiled stinging cells (
cnidocyte A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this ...
s) which pierce, poison and firmly hold living prey paralyzing or killing them. Polyp prey includes plankton such as copepods and fish larvae. Longitudinal muscular fibers formed from the cells of the ectoderm allow tentacles to contract to convey the food to the mouth. Similarly, circularly disposed muscular fibres formed from the endoderm permit tentacles to be protracted or thrust out once they are contracted. In both stony and soft corals, the polyps can be retracted by contracting muscle fibres, with stony corals relying on their hard skeleton and cnidocytes for defense. Soft corals generally secrete terpenoid toxins to ward off predators. In most corals, the tentacles are retracted by day and spread out at night to catch plankton and other small organisms. Shallow-water species of both stony and soft corals can be zooxanthellate, the corals supplementing their plankton diet with the products of photosynthesis produced by these symbionts. The polyps interconnect by a complex and well-developed system of gastrovascular canals, allowing significant sharing of nutrients and symbionts. The external form of the polyp varies greatly. The column may be long and slender, or may be so short in the axial direction that the body becomes disk-like. The tentacles may number many hundreds or may be very few, in rare cases only one or two. They may be simple and unbranched, or feathery in pattern. The mouth may be level with the surface of the peristome, or may be projecting and trumpet-shaped.


Soft corals

Soft corals have no solid exoskeleton as such. However, their tissues are often reinforced by small supportive elements known as sclerites made of calcium carbonate. The polyps of soft corals have eight-fold symmetry, which is reflected in the ''Octo'' in Octocorallia. Soft corals vary considerably in form, and most are colonial. A few soft corals are stolonate, but the polyps of most are connected by sheets of tissue called coenosarc, and in some species these sheets are thick and the polyps deeply embedded in them. Some soft corals encrust other sea objects or form lobes. Others are tree-like or whip-like and have a central axial skeleton embedded at their base in the matrix of the supporting branch. These branches are composed of a fibrous protein called gorgonin or of a calcified material.


Stony corals

The polyps of stony corals have six-fold symmetry. In stony corals, the tentacles are cylindrical and taper to a point, but in soft corals they are pinnate with side branches known as pinnules. In some tropical species, these are reduced to mere stubs and in some, they are fused to give a paddle-like appearance. Coral skeletons are biocomposites (mineral + organics) of calcium carbonate, in the form of calcite or aragonite. In scleractinian corals, "centers of calcification" and fibers are clearly distinct structures differing with respect to both morphology and chemical compositions of the crystalline units. The organic matrices extracted from diverse species are acidic, and comprise proteins, sulphated sugars and lipids; they are species specific. The soluble organic matrices of the skeletons allow to differentiate zooxanthellae and non-zooxanthellae specimens.


Ecology


Feeding

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


Intracellular symbionts

Many corals, as well as other cnidarian groups such as sea anemones form a symbiotic relationship with a class of dinoflagellate algae, zooxanthellae of the genus '' Symbiodinium'', which can form as much as 30% of the tissue of a polyp. Typically, each polyp harbors one species of alga, and coral species show a preference for '' Symbiodinium''. Young corals are not born with zooxanthellae, but acquire the algae from the surrounding environment, including the water column and local sediment. The main benefit of the zooxanthellae is their ability to photosynthesize which supplies corals with the products of photosynthesis, including glucose, glycerol, and amino acids, which the corals can use for energy. Zooxanthellae also benefit corals by aiding in calcification, for the coral skeleton, and waste removal. In addition to the soft tissue, microbiomes are also found in the coral's mucus and (in stony corals) the skeleton, with the latter showing the greatest microbial richness. The zooxanthellae benefit from a safe place to live and consume the polyp's
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
, phosphate and nitrogenous waste. Stressed corals will eject their zooxanthellae, a process that is becoming increasingly common due to strain placed on coral by rising ocean temperatures. Mass ejections are known as coral bleaching because the algae contribute to coral coloration; some colors, however, are due to host coral pigments, such as green fluorescent proteins (GFPs). Ejection increases the polyp's chance of surviving short-term stress and if the stress subsides they can regain algae, possibly of a different species, at a later time. If the stressful conditions persist, the polyp eventually dies. Zooxanthellae are located within the coral cytoplasm and due to the algae's photosynthetic activity the internal pH of the coral can be raised; this behavior indicates that the zooxanthellae are responsible to some extent for the metabolism of their host corals. Stony Coral Tissue Loss Disease has been associated with the breakdown of host-zooxanthellae physiology. Moreover, Vibrio bacterium are known to have virulence traits used for host coral tissue damage and photoinhibition of algal symbionts.Therefore, both coral and their symbiotic microorganisms could have evolved to harbour traits resistant to disease and transmission.


Reproduction

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


Sexual

Corals predominantly reproduce sexually. About 25% of hermatypic corals (reef building stony corals) form single sex ( gonochoristic) colonies, while the rest are hermaphroditic. It is estimated more than 67% of coral are simultaneous hermaphrodites.


Broadcasters

About 75% of all hermatypic corals "broadcast spawn" by releasing gametes— eggs and sperm—into the water to spread offspring. A genetic ability to detect blue light allows these sightless organisms to detect a full moon, triggering the simultaneous release. The gametes fertilize at the water's surface to form a microscopic larva called a planula, typically pink and elliptical in shape. A typical coral colony forms several thousand larvae per year to overcome the odds against formation of a new colony. Synchronous spawning is very typical on the coral reef, and often, even when multiple
species In biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that ...
are present, all corals spawn on the same night. This synchrony is essential so male and female gametes can meet. Corals rely on environmental cues, varying from species to species, to determine the proper time to release gametes into the water. The cues involve temperature change, lunar cycle, day length, and possibly chemical signalling. Synchronous spawning may form hybrids and is perhaps involved in coral speciation. The immediate cue is most often sunset, which cues the release. The spawning event can be visually dramatic, clouding the usually clear water with gametes.


Brooders

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


Planulae

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


Asexual

Within a coral head, the genetically identical polyps reproduce asexually, either by budding (gemmation) or by dividing, whether longitudinally or transversely. Budding involves splitting a smaller polyp from an adult. As the new polyp grows, it forms its body parts. The distance between the new and adult polyps grows, and with it, the coenosarc (the common body of the colony). Budding can be intratentacular, from its oral discs, producing same-sized polyps within the ring of tentacles, or extratentacular, from its base, producing a smaller polyp. Division forms two polyps that each become as large as the original. Longitudinal division begins when a polyp broadens and then divides its coelenteron (body), effectively splitting along its length. The mouth divides and new tentacles form. The two polyps thus created then generate their missing body parts and exoskeleton. Transversal division occurs when polyps and the exoskeleton divide transversally into two parts. This means one has the basal disc (bottom) and the other has the oral disc (top); the new polyps must separately generate the missing pieces. Asexual reproduction offers the benefits of high reproductive rate, delaying senescence, and replacement of dead modules, as well as geographical distribution.


Colony division

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


Coral microbiome

Corals are one of the more common examples of an animal host whose symbiosis with microalgae can turn to dysbiosis, and is visibly detected as bleaching. Coral microbiomes have been examined in a variety of studies, which demonstrate how oceanic environmental variations, most notably temperature, light, and inorganic nutrients, affect the abundance and performance of the microalgal symbionts, as well as calcification and physiology of the host.Apprill, A. (2017) "Marine animal microbiomes: toward understanding host–microbiome interactions in a changing ocean". ''Frontiers in Marine Science'', 4: 222. . Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
Studies have also suggested that resident bacteria, archaea, and fungi additionally contribute to nutrient and organic matter cycling within the coral, with viruses also possibly playing a role in structuring the composition of these members, thus providing one of the first glimpses at a multi-domain marine animal symbiosis. The gammaproteobacterium '' Endozoicomonas'' is emerging as a central member of the coral's microbiome, with flexibility in its lifestyle.Neave, M.J., Apprill, A., Ferrier-Pagès, C. and Voolstra, C.R. (2016) "Diversity and function of prevalent symbiotic marine bacteria in the genus ''Endozoicomonas''". ''Applied Microbiology and Biotechnology'', 100(19): 8315–8324. . Given the recent mass bleaching occurring on reefs, corals will likely continue to be a useful and popular system for symbiosis and dysbiosis research. '' Astrangia poculata'', the northern star coral, is a temperate stony coral, widely documented along the eastern coast of the United States. The coral can live with and without zooxanthellae (algal symbionts), making it an ideal
model organism A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the work ...
to study microbial community interactions associated with symbiotic state. However, the ability to develop primers and probes to more specifically target key microbial groups has been hindered by the lack of full length 16S rRNA sequences, since sequences produced by the Illumina platform are of insufficient length (approximately 250 base pairs) for the design of primers and probes. In 2019, Goldsmith et al demonstrated Sanger sequencing was capable of reproducing the biologically-relevant diversity detected by deeper next-generation sequencing, while also producing longer sequences useful to the research community for probe and primer design (see diagram on right). Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License


Holobiont

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


Cophylogeny and phylosymbiosis

It is known that the coral's microbiome and symbiont influence host health, however the historic influence of each member on others is not well understood. Scleractinian corals have been diversifying for longer than many other symbiotic systems, and their microbiomes are known to be partially species-specific. It has been suggested that '' Endozoicomonas'', a commonly highly abundant bacterium in corals, has exhibited codiversification with its host. This hints at an intricate set of relationships between the members of the coral holobiont that have been developing as
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of gene In biology, the word gene (from , ; "... Wilhelm Johannsen coined the ...
of these members occurs. A study published in 2018 revealed evidence of phylosymbiosis between corals and their tissue and skeleton microbiomes. The coral skeleton, which represents the most diverse of the three coral microbiomes, showed the strongest evidence of phylosymbiosis. Coral microbiome composition and richness were found to reflect coral phylogeny. For example, interactions between bacterial and eukaryotic coral phylogeny influence the abundance of ''Endozoicomonas'', a highly abundant bacterium in the coral holobiont. However, host-microbial cophylogeny appears to influence only a subset of coral-associated bacteria.


Reefs

Many corals in the order Scleractinia are hermatypic, meaning that they are involved in building reefs. Most such corals obtain some of their energy from zooxanthellae in the genus ''Symbiodinium''. These are symbiotic photosynthetic dinoflagellates which require sunlight; reef-forming corals are therefore found mainly in shallow water. They secrete calcium carbonate to form hard skeletons that become the framework of the reef. However, not all reef-building corals in shallow water contain zooxanthellae, and some deep water species, living at depths to which light cannot penetrate, form reefs but do not harbour the symbionts. There are various types of shallow-water coral reef, including fringing reefs, barrier reefs and atolls; most occur in tropical and subtropical seas. They are very slow-growing, adding perhaps one centimetre (0.4 in) in height each year. The
Great Barrier Reef 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 ...
is thought to have been laid down about two million years ago. Over time, corals fragment and die, sand and rubble accumulates between the corals, and the shells of clams and other molluscs decay to form a gradually evolving calcium carbonate structure. Coral reefs are extremely diverse marine ecosystems hosting over 4,000 species of fish, massive numbers of cnidarians, molluscs,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod Arthropods (, (gen. ποδός)) are invertebrate animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals ...
s, and many other animals.


Evolution

At certain times in the geological past, corals were very abundant. Like modern corals, these ancestors built reefs, some of which ended as great structures in sedimentary rocks. Fossils of fellow reef-dwellers algae, sponges, and the remains of many echinoids, brachiopods,
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, biv ...
s,
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from lan ...
s, and trilobites appear along with coral fossils. This makes some corals useful index fossils. Coral fossils are not restricted to reef remnants, and many solitary fossils are found elsewhere, such as ''Cyclocyathus'', which occurs in England's Gault clay formation.


Early corals

Corals first appeared in the Cambrian about .
Fossil A fossil (from Classical Latin Classical Latin is the form of Literary Latin recognized as a literary standard by writers of the late Roman Republic and early Roman Empire The Roman Empire ( la, Imperium Romanum ; grc-gre, ...
s are extremely rare until the Ordovician period, 100 million years later, when Heliolitida, rugose, and tabulate corals became widespread. Paleozoic corals often contained numerous endobiotic symbionts. Tabulate corals occur in
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms w ...
s and calcareous shales of the Ordovician period, with a gap in the fossil record due to extinction events at the end of the Ordovician. Corals reappeared some millions of years later during the Silurian period, and tabulate corals often form low cushions or branching masses of calcite alongside rugose corals. Tabulate coral numbers began to decline during the middle of the Silurian period. Rugose or horn corals became dominant by the middle of the Silurian period, and during the Devonian, corals flourished with more than 200 genera. The rugose corals existed in solitary and colonial forms, and were also composed of calcite. Both rugose and tabulate corals became extinct in the Permian–Triassic extinction event (along with 85% of marine species), and there is a gap of tens of millions of years until new forms of coral evolved in the Triassic. File:Syringoporid.jpg, Tabulate coral (a syringoporid); Boone limestone (Lower
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonif ...
) near Hiwasse, Arkansas, scale bar is 2.0 cm File:AuloporaDevonianSilicaShale.jpg, Tabulate coral '' Aulopora'' from the
Devonian The Devonian ( ) is a period (geology), geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Dev ...
period File:RugosaOrdovician.jpg, Solitary rugose coral ('' Grewingkia'') in three views; Ordovician, southeastern Indiana


Modern corals

The currently ubiquitous stony corals, Scleractinia, appeared in the Middle Triassic to fill the niche vacated by the extinct rugose and tabulate orders, and is not closely related to the earlier forms. Unlike the corals prevalent before the Permian extinction, which formed skeletons of a form of calcium carbonate known as calcite, modern stony corals form skeletons composed of the aragonite. Their fossils are found in small numbers in rocks from the Triassic period, and become common in the
Jurassic The Jurassic ( ) is a geologic period The geologic time scale, or geological time scale, (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the proce ...
and later periods. Although they are geologically younger than the tabulate and rugose corals, the aragonite of their skeletons is less readily preserved, and their fossil record is accordingly less complete.


Status


Threats

Coral reefs are under stress around the world. In particular, coral mining, agricultural and urban runoff, pollution (organic and inorganic), overfishing, blast fishing, disease, and the digging of canals and access into islands and bays are localized threats to coral ecosystems. Broader threats are sea temperature rise, sea level rise and pH changes from ocean acidification, all associated with greenhouse gas emissions. In 1998, 16% of the world's reefs died as a result of increased water temperature. Approximately 10% of the world's coral reefs are dead. About 60% of the world's reefs are at risk due to human-related activities. The threat to reef health is particularly strong in
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia Asia (, ) is one of the world's most notable geographi ...
, where 80% of reefs are endangered. Over 50% of the world's
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in groups ...
s may be destroyed by 2030; as a result, most nations protect them through environmental laws. In the Caribbean and tropical Pacific, direct contact between ~40–70% of common seaweeds and coral causes bleaching and death to the coral via transfer of
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids in ...
-soluble metabolites. Seaweed and algae proliferate given adequate nutrients and limited grazing by herbivores such as parrotfish. Water temperature changes of more than or
salinity Salinity () is the saltiness or amount of salt dissolved in a body of water, called saline water (see also soil salinity). It is usually measured in g/L or g/kg (grams of salt per liter/kilogram of water; the latter is dimensionless and equa ...
changes can kill some species of coral. Under such environmental stresses, corals expel their Symbiodinium; without them coral tissues reveal the white of their skeletons, an event known as coral bleaching. Submarine springs found along the coast of Mexico's Yucatán Peninsula produce water with a naturally low pH (relatively high acidity) providing conditions similar to those expected to become widespread as the oceans absorb carbon dioxide. Surveys discovered multiple species of live coral that appeared to tolerate the acidity. The colonies were small and patchily distributed, and had not formed structurally complex reefs such as those that compose the nearby
Mesoamerican Barrier Reef System The Mesoamerican Barrier Reef System (MBRS), also popularly known as the Great Mayan Reef or Great Maya Reef, is a marine region that stretches over along the coasts of four countries – Mexico Mexico ( Spanish: México), officially the ...
.


Coral health

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


Climate change impacts

Increasing sea surface temperatures in tropical regions (~) the last century have caused major coral bleaching, death, and therefore shrinking coral populations. Although coral are able to adapt and acclimate, it is uncertain if this evolutionary process will happen quickly enough to prevent major reduction of their numbers. Annual growth bands in some corals, such as the
deep sea The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of 200 metres (656 feet) or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combin ...
bamboo corals (''Isididae''), may be among the first signs of the effects of ocean acidification on marine life. The growth rings allow
geologist A geologist is a scientist A scientist is a person who conducts scientific research to advance knowledge in an area of the natural sciences. In classical antiquity Classical antiquity (also the classical era, classical period or ...
s to construct year-by-year chronologies, a form of incremental dating, which underlie high-resolution records of past climatic and environmental changes using geochemical techniques. Certain species form communities called microatolls, which are colonies whose top is dead and mostly above the water line, but whose perimeter is mostly submerged and alive. Average tide level limits their height. By analyzing the various growth morphologies, microatolls offer a low resolution record of sea level change. Fossilized microatolls can also be dated using
radiocarbon dating Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for determining the age of an object containing organic material by using the properties of radiocarbon, a radioactive isotope of carbon. The method was ...
. Such methods can help to reconstruct
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene toge ...
sea level Mean sea level (MSL, often shortened to sea level) is an average surface level of one or more among Earth Earth is the third planet from the Sun and the only astronomical object known to harbor life. While large volumes of wate ...
s. Though coral have large sexually-reproducing populations, their evolution can be slowed by abundant asexual reproduction. Gene flow is variable among coral species. According to the
biogeography Biogeography is the study of the distribution of species In biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent ...
of coral species, gene flow cannot be counted on as a dependable source of adaptation as they are very stationary organisms. Also, coral longevity might factor into their adaptivity. However, adaptation to climate change has been demonstrated in many cases, which is usually due to a shift in coral and zooxanthellae genotypes. These shifts in allele frequency have progressed toward more tolerant types of zooxanthellae. Scientists found that a certain scleractinian zooxanthella is becoming more common where sea temperature is high. Symbionts able to tolerate warmer water seem to photosynthesise more slowly, implying an evolutionary trade-off. In the Gulf of Mexico, where sea temperatures are rising, cold-sensitive staghorn and elkhorn coral have shifted in location. Not only have the symbionts and specific species been shown to shift, but there seems to be a certain growth rate favorable to selection. Slower-growing but more heat-tolerant corals have become more common. The changes in temperature and acclimation are complex. Some reefs in current shadows represent a refugium location that will help them adjust to the disparity in the environment even if eventually the temperatures may rise more quickly there than in other locations. This separation of populations by climatic barriers causes a realized niche to shrink greatly in comparison to the old fundamental niche.


Geochemistry

Corals are shallow, colonial organisms that integrate oxygen and trace elements into their skeletal aragonite ( polymorph of calcite) crystalline structures as they grow. Geochemical anomalies within the crystalline structures of corals represent functions of temperature, salinity and oxygen isotopic composition. Such geochemical analysis can help with climate modeling. The ratio of oxygen-18 to oxygen-1618O), for example, is a proxy for temperature.


=Strontium/calcium ratio anomaly

= Time can be attributed to coral geochemistry anomalies by correlating strontium/
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar ...
minimums with sea surface temperature (SST) maximums to data collected fro
NINO 3.4 SSTA


=Oxygen isotope anomaly

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


=Sea surface temperature and sea surface salinity

= The global moisture budget is primarily being influenced by tropical sea surface temperatures from the position of the Intertropical Convergence Zone (ITCZ). The Southern Hemisphere has a unique meteorological feature positioned in the southwestern Pacific Basin called the South Pacific Convergence Zone (SPCZ), which contains a perennial position within the Southern Hemisphere. During ENSO warm periods, the SPCZ reverses orientation extending from the equator down south through
Solomon Islands Solomon Islands is an island country consisting of six major islands and over 900 smaller islands in Oceania Oceania (, , ) is a region, geographical region that includes Australasia, Melanesia, Micronesia, and Polynesia. Spanning t ...
,
Vanuatu Vanuatu ( or ; ), officially the Republic of Vanuatu (french: link=no, République de Vanuatu; bi, Ripablik blong Vanuatu), is an island country located in the South Pacific Ocean. The archipelago An archipelago ( ), sometimes call ...
,
Fiji Fiji ( , ,; fj, Viti, ; Fiji Hindi: फ़िजी, ''Fijī''), officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean The Pacific Ocean is the largest and deepest o ...
and towards the French Polynesian Islands; and due east towards
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sou ...
affecting geochemistry of corals in tropical regions. Geochemical analysis of skeletal coral can be linked to sea surface salinity (SSS) and
sea surface temperature Sea surface temperature (SST), or ocean surface temperature, is the ocean temperature close to the surface. The exact meaning of ''surface'' varies according to the measurement method used, but it is between and below the sea surface. Air ma ...
(SST), fro
El Nino 3.4 SSTA
data, of tropical oceans to seawater δ18O ratio anomalies from corals. ENSO phenomenon can be related to variations in sea surface salinity (SSS) and sea surface temperature (SST) that can help model tropical climate activities.


=Limited climate research on current species

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


Protection

Marine Protected Areas, Biosphere reserves, marine parks, national monuments world heritage status, fishery management and habitat protection can protect reefs from anthropogenic damage. Many governments now prohibit removal of coral from reefs, and inform coastal residents about reef protection and ecology. While local action such as habitat restoration and herbivore protection can reduce local damage, the longer-term threats of acidification, temperature change and sea-level rise remain a challenge. Protecting networks of diverse and healthy reefs, not only climate refugia, helps ensure the greatest chance of
genetic diversity 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 In biology Biology is the scientific study of life. It ...
, which is critical for coral to adapt to new climates. A variety of conservation methods applied across marine and terrestrial threatened ecosystems makes coral adaption more likely and effective. To eliminate destruction of corals in their indigenous regions, projects have been started to grow corals in non-tropical countries.


Relation to humans

Local economies near major coral reefs benefit from an abundance of fish and other marine creatures as a food source. Reefs also provide recreational
scuba diving Scuba diving is a mode of underwater diving whereby divers use breathing equipment that is completely independent of a surface air supply. The name "scuba", an acronym for " Self-Contained Underwater Breathing Apparatus", was coined by Ch ...
and
snorkeling Snorkeling ( British and Commonwealth English spelling: snorkelling) is the practice of swimming on or through a body of water while equipped with a diving mask, a shaped breathing tube called a snorkel, and usually swimfins. In cooler waters ...
tourism. These activities can damage coral but international projects such as Green Fins that encourage dive and snorkel centres to follow a Code of Conduct have been proven to mitigate these risks.


Jewelry

Corals' many colors give it appeal for necklaces and other jewelry. Intensely red coral is prized as a gemstone. Sometimes called fire coral, it is not the same as fire coral. Red coral is very rare because of overharvesting. In general, it is inadvisable to give coral as gifts since they are in decline from stressors like climate change, pollution, and unsustainable fishing. Always considered a precious mineral, "the Chinese have long associated red coral with auspiciousness and longevity because of its color and its resemblance to deer antlers (so by association, virtue, long life, and high rank". It reached its height of popularity during the Manchu or Qing Dynasty (1644-1911) when it was almost exclusively reserved for the emperor's use either in the form of coral beads (often combined with pearls) for court jewelry or as decorative Penjing (decorative miniature mineral trees). Coral was known as ''shanhu'' in Chinese. The "early-modern 'coral network' egan inthe Mediterranean Sea nd found its wayto Qing China via the English
East India Company The East India Company (EIC) was an English, and later British, joint-stock company founded in 1600 and dissolved in 1874. It was formed to trade in the Indian Ocean region, initially with the East Indies (the Indian subcontinent Th ...
". There were strict rules regarding its use in a code established by the Qianlong Emperor in 1759.


Medicine

In medicine, chemical compounds from corals can potentially be used to treat cancer, AIDS, pain, and for other therapeutic uses. Coral skeletons, e.g. '' Isididae'' are also used for bone grafting in humans. Coral Calx, known as Praval Bhasma in
Sanskrit Sanskrit (; attributively , ; nominally , , ) is a classical language belonging to the Indo-Aryan branch of the Indo-European languages. It arose in South Asia South Asia is the southern subregion of Asia Asia (, ) is one ...
, is widely used in traditional system of Indian medicine as a supplement in the treatment of a variety of bone metabolic disorders associated with calcium deficiency. In classical times ingestion of pulverized coral, which consists mainly of the weak base calcium carbonate, was recommended for calming stomach ulcers by Galen and Dioscorides.


Construction

Coral reefs in places such as the East African coast are used as a source of
building material Building material is material used for construction Construction is a general term meaning the art and science Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and ...
. Ancient (fossil) coral limestone, notably including the Coral Rag Formation of the hills around
Oxford Oxford () is a city in England. It is the county town and only city of Oxfordshire. In 2020, its population was estimated at 151,584. It is north-west of London London is the capital and largest city of England and the United Kin ...
(England), was once used as a building stone, and can be seen in some of the oldest buildings in that city including the Saxon tower of St Michael at the Northgate, St. George's Tower of Oxford Castle, and the medieval walls of the city.


Shoreline protection

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


Local economies

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


Aquaria

The saltwater fishkeeping hobby has expanded, over recent years, to include reef tanks, fish tanks that include large amounts of live rock on which coral is allowed to grow and spread. These tanks are either kept in a natural-like state, with algae (sometimes in the form of an algae scrubber) and a deep sand bed providing filtration, or as "show tanks", with the rock kept largely bare of the algae and microfauna that would normally populate it, in order to appear neat and clean. The most popular kind of coral kept is soft coral, especially zoanthids and mushroom corals, which are especially easy to grow and propagate in a wide variety of conditions, because they originate in enclosed parts of reefs where water conditions vary and lighting may be less reliable and direct.Coral Reefs
. Marinebio.org. Retrieved on 2016-06-13.
More serious fishkeepers may keep small polyp stony coral, which is from open, brightly lit reef conditions and therefore much more demanding, while large polyp stony coral is a sort of compromise between the two.


Aquaculture

Coral aquaculture, also known as ''coral farming'' or ''coral gardening'', is the cultivation of corals for commercial purposes or coral reef restoration. Aquaculture is showing promise as a potentially effective tool for restoring
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in groups ...
s, which have been declining around the world. The process bypasses the early growth stages of corals when they are most at risk of dying. Coral fragments known as "seeds" are grown in nurseries then replanted on the reef. Coral is farmed by coral farmers who live locally to the reefs and farm for reef conservation or for income. It is also farmed by scientists for research, by businesses for the supply of the live and ornamental coral trade and by private aquarium hobbyists.


Gallery

''Further images: commons:Coral reefs and commons:Corals'' File:Mushroom Coral (Fungia) Top Macro 91.JPG, '' Fungia'' sp. skeleton File:Eusmilia fastigiata large.jpg, Polyps of '' Eusmilia fastigiata'' File:Dendrogyra cylindrus (pillar coral) (San Salvador Island, Bahamas) 1 (15513345363).jpg, Pillar coral, ''Dendrogyra cylindricus'' File:Brain coral.jpg, Brain coral, '' Diploria labyrinthiformis'' File:Brain coral spawning.jpg, Brain coral spawning File:Stony coral spawning 3.jpg, Brain coral releasing eggs File:EilatFringingReef.jpg, Fringing
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in groups ...
off the coast of Eilat,
Israel Israel (; he, יִשְׂרָאֵל, ; ar, إِسْرَائِيل, ), officially the State of Israel ( he, מְדִינַת יִשְׂרָאֵל, label=none, translit=Medīnat Yīsrāʾēl; ), is a country in Western Asia. It is situated ...
. File:000324-Corals-IMG 0418-2.jpg, Corals, Tis Beach, Chabahar,
Iran Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq Iraq,; ku, عێراق, translit=Êraq officially the Republic of Iraq, '; ku, کۆما ...
File:000407-Corals-IMG 0693-2.jpg, Corals, Tis Beach, Chabahar,
Iran Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq Iraq,; ku, عێراق, translit=Êraq officially the Republic of Iraq, '; ku, کۆما ...


See also

* Keystone species * Ringstead Coral Bed


References


Sources

* * * * * * * * * * *


External links


Coral Reefs
The Ocean Portal by the Smithsonian Institution * NOAA
Coral Reef Conservation Program
* NOAA CoRIS
Coral Reef Biology
* NOAA Office for Coastal Management
Fast Facts - Coral Reefs
* NOAA Ocean Service Education
Corals
* {{Authority control Anthozoa