111-615: See text Acropora is a genus of small polyp stony coral in the phylum Cnidaria . Some of its species are known as table coral, elkhorn coral , and staghorn coral . Over 149 species are described. Acropora species are some of the major reef corals responsible for building the immense calcium carbonate substructure that supports the thin living skin of a reef. Depending on the species and location, Acropora species may grow as plates or slender or broad branches. Like other corals, Acropora corals are colonies of individual polyps, which are about 2 mm across and share tissue and
222-477: A full moon . 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 and small fish 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
333-603: A monophyletic group. The earliest scleractinians were not reef builders, but were small, phaceloid or solitary individuals. Scleractinian corals were probably at their greatest diversity in the Jurassic and all but disappeared in the mass extinction event at the end of the Cretaceous, about 18 out of 67 genera surviving. Recently discovered Paleozoic corals with aragonitic skeletons and cyclic septal insertion – two features that characterize Scleractinia – have strengthened
444-467: A nerve net . The polyps can withdraw back into the coral in response to movement or disturbance by potential predators, but when undisturbed, they protrude slightly. The polyps typically extend further at night to help capture plankton and organic matter from the water. The species are distributed in the Indo-Pacific (over 100 species) and Caribbean (3 species). However, the true number of species
555-453: 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
666-420: A calcified material. 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
777-491: A calcium carbonate base, with polyps that bear six stiff tentacles, and soft coral (Alcyonacea and ahermatypic coral) which are pliable and formed by a colony of polyps with eight feather-like tentacles. These two classifications arose from differentiation in gene expressions in their branch tips and bases that arose through developmental signaling pathways such as Hox , Hedgehog , Wnt , BMP etc. Scientists typically select Acropora as research models since they are
888-417: 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 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
999-403: A cup surrounding this part of the polyp. The interior of the cup contains radially aligned plates, or septa , projecting upwards from the base. Each of these plates is flanked by a pair of mesenteries. The septa are secreted by the mesenteries, and are therefore added in the same order as the mesenteries are. As a result, septa of different ages are adjacent to one another, and the symmetry of
1110-548: 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 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
1221-503: A dwindling of populations of Acropora , along with other coral species. Acropora is especially susceptible to bleaching when stressed. Bleaching is due to the loss of the coral's zooxanthellae, which are a golden-brown color. Bleached corals are stark white and may die if new Symbiodinium cells cannot be assimilated. Common causes of bleaching and coral death include pollution, abnormally warm water temperatures, increased ocean acidification , sedimentation, and eutrophication. In 2014
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#17327733566221332-459: A few centimeters in height. A set of tentacles 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
1443-520: A full moon. A full moon is equivalent to four to six hours of continuous dim light exposure, which can cause light-dependent reactions in protein. Corals contain light-sensitive cryptochromes , proteins whose light-absorbing flavin structures are sensitive to different types of light. This allows corals such as Dipsastraea speciosa to detect and respond to changes in sunlight and moonlight. Moonlight itself may actually suppress coral spawning. The most immediate cue to cause spawning appears to be
1554-441: 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. 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
1665-640: A rate up to three times faster than similar species without symbionts. These corals typically grow in shallow, well-lit, warm water with moderate to brisk turbulence and abundant oxygen, and prefer firm, non-muddy surfaces on which to settle. Most stony corals extend their tentacles to feed on zooplankton , but those with larger polyps take correspondingly larger prey, including various invertebrates and even small fish. In addition to capturing prey in this way, many stony corals also produce mucus films they can move over their bodies using cilia ; these trap small organic particles which are then pulled towards and into
1776-689: A result of teaming up with symbiotic algae . Nine of the sub-orders were in existence by the end of the Triassic and three more had appeared by the Jurassic (200 million years ago), with a further suborder appearing in the Middle Cretaceous (100 million years ago). Some may have developed from a common ancestor, either an anemone-like coral without a skeleton, or a rugose coral . A rugose coral seems an unlikely common ancestor because these corals had calcite rather than aragonite skeletons, and
1887-408: A ring of tentacles . The base of the polyp secretes the stony material from which the coral skeleton is formed. The body wall of the polyp consists of mesoglea sandwiched between two layers of epidermis. The mouth is at the centre of the oral disc and leads into a tubular pharynx which descends for some distance into the body before opening into the gastrovascular cavity that fills the interior of
1998-400: A single polyp and eventually develops into a juvenile and then adult by asexual budding and growth. 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
2109-415: 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,
2220-453: A subset of coral-associated bacteria. 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
2331-531: Is paraphyletic because the sea anemones are also in the sub-class Hexacorallia. The delineation of coral species is challenging as hypotheses based on morphological traits contradict hypotheses formed via molecular tree-based processes. As of 2020, there are 2175 identified separate coral species, 237 of which are currently endangered, making distinguishing corals to be the utmost of importance in efforts to curb extinction. Adaptation and delineation continues to occur in species of coral in order to combat
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#17327733566222442-409: Is coordinated by chemical communication. 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 . About 75% of all hermatypic corals "broadcast spawn" by releasing gametes — eggs and sperm —into
2553-415: 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. 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
2664-449: Is particularly challenging. Many species were described before the advent of scuba diving , with little realisation by the authors that coral species could have varying morphologies in different habitats. Collectors were mostly limited to observing corals on reef flats, and were unable to observe the changes in morphology that occurred in more turbid, deeper-water conditions. More than 2,000 nominal species were described in this era, and by
2775-449: 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 , crustaceans , and many other animals. At certain times in
2886-402: Is unknown: firstly, the validity of many of these species is questioned as some have been shown to represent hybrids, for example Acropora prolifera ; and secondly, some species have been shown to represent cryptic species complexes. Symbiodinium , symbiotic algae , live in the corals' cells and produce energy for the animals through photosynthesis . Environmental destruction has led to
2997-531: The Convention on International Trade in Endangered Species (CITES) meaning that their international trade (including in parts and derivatives) is regulated. The World Register of Marine Species lists the following families as being included in the order Scleractinia. Some species have not been placeable ( Incertae sedis ): Coral Corals are colonial marine invertebrates within
3108-530: The Darwin Mounds , northwest of Cape Wrath , Scotland, and others off the coast of Washington state and the Aleutian Islands . The classification of corals has been discussed for millennia, owing to having similarities to both plants and animals. Aristotle 's pupil Theophrastus described the red coral , korallion , in his book on stones, implying it was a mineral, but he described it as
3219-451: The effects of global warming and ocean acidification . Scleractinian corals may be solitary or colonial . Colonies can reach considerable size, consisting of a large number of individual polyps. Stony corals are members of the class Anthozoa and like other members of the group, do not have a medusa stage in their life cycle. The individual animals are known as polyps and have a cylindrical body crowned by an oral disc surrounded by
3330-461: The endodermal cells are usually replete with symbiotic unicellular dinoflagellates known as zooxanthellae . There are sometimes as many as five million cells of these per 1 square centimetre (0.16 sq in) of coral tissue. Up to 50% of organic compounds produced by symbionts are used as food by polyps. The oxygen byproduct of photosynthesis and the additional energy derived from sugars produced by zooxanthellae enable these corals to grow at
3441-488: The pyrgomatids , but a recent study recorded evidence of living pyrgomatids in stylasterids , casting doubt on this idea. Stony corals have a great range of reproductive strategies and can reproduce both sexually and asexually. Many species have separate sexes, the whole colony being either male or female, but others are hermaphroditic , with individual polyps having both male and female gonads. Some species brood their eggs but in most species, sexual reproduction results in
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3552-426: The subphylum 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
3663-420: The (French) name 'coral'." The Persian polymath Al-Biruni (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 . Presently, corals are classified as species of animals within
3774-478: The 19th and early 20th centuries. The two most advanced 19th century classifications both used complex skeletal characters; The 1857 classification of the French zoologists Henri Milne-Edwards and Jules Haime 's was based on macroscopic skeletal characters, while Francis Grant Ogilvie 's 1897 scheme was developed using observations of skeletal microstructures, with particular attention to the structure and pattern of
3885-455: 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
3996-619: 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 refers to coral among a list of types of trees, and the 11th-century French commentator Rashi describes it as "a type of tree (מין עץ) that grows underwater that goes by
4107-536: 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). Reef-building corals are well-studied holobionts that include
4218-605: 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
4329-404: The U.S. Fish and Wildlife Service listed ten Acropora species as 'threatened'. Most Acropora species are brown or green, but a few are brightly colored, and those rare corals are prized by aquarists. Captive propagation of Acropora is widespread in the reef-keeping community. Given the right conditions, many Acropora species grow quickly, and individual colonies can exceed a meter across in
4440-405: The abundance and performance of the microalgal symbionts, as well as calcification and physiology of the host. 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
4551-421: The adjacent colonies of the same species form a single colony by fusing. Most colonial species have very small polyps, ranging from 1 to 3 mm (0.04 to 0.12 in) in diameter, although some solitary species may be as large as 25 cm (10 in). The skeleton of an individual scleractinian polyp is known as a corallite. It is secreted by the epidermis of the lower part of the body, and initially forms
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4662-525: The age of the coral. Solitary corals do not bud. They gradually increase in size as they deposit more calcium carbonate and produce new whorls of septa. A large Ctenactis echinata for example normally has a single mouth, may be about 25 cm (10 in) long and have more than a thousand septa. Stony corals occur in all the world's oceans. There are two main ecological groups. Hermatypic corals are mostly colonial corals which tend to live in clear, oligotrophic, shallow tropical waters; they are
4773-407: 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
4884-477: The amount of water movement and other factors. Many shallow-water corals contain symbiont unicellular organisms known as zooxanthellae within their tissues. These give their colour to the coral which thus may vary in hue depending on what species of symbiont it contains. Stony corals are closely related to sea anemones , and like them are armed with stinging cells known as cnidocytes . Corals reproduce both sexually and asexually. Most species release gametes into
4995-487: The area has decreased and in some cases ceased. The area was previously considered a refuge for corals because mass bleaching events due to climate change had not been observed there. Coral restoration techniques for coral reef management are being developed to increase fertilization rates, larval development, and settlement of new corals. Brooding species are most often ahermatypic (not reef-building) in areas of high current or wave action. Brooders release only sperm, which
5106-503: The assembling of species into the existing families , but not into the traditional suborders. For example, some genera affiliated with different suborders were now located on the same branch of a phylogenetic tree . In addition, there is no distinguishing morphological character that separates clades , only molecular differences. The Australian zoologist John Veron and his co-workers analyzed ribosomal RNA in 1996 to obtain similar results to Romano and Palumbi, again concluding that
5217-452: The body and tentacles. Unlike other cnidarians however, the cavity is subdivided by a number of radiating partitions, thin sheets of living tissue, known as mesenteries . The gonads are also located within the cavity walls. The polyp is retractable into the corallite , the stony cup in which it sits, being pulled back by sheet-like retractor muscles. The polyps are connected by horizontal sheets of tissue known as coenosarc extending over
5328-404: The budding of new polyps. There are two types of budding, intratentacular and extratentacular. In intratentacular budding, a new polyp develops on the oral disc, inside the ring of tentacles. This can form individual, separate polyps or a row of partially separated polyps sharing an elongate oral disc with a series of mouths. Tentacles grow around the margin of this elongated oral disc and not around
5439-458: The capture of food. Polyps extend their tentacles, particularly at night, often containing coiled stinging cells ( cnidocytes ) 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
5550-414: The capture of plankton and suspended organic particles. The growth rates of most species of non-zooxanthellate corals are significantly slower than those of their counterparts, and the typical structure for these corals is less calcified and more susceptible to mechanical damage than that of zooxanthellate corals. Scleratinians were previously believed to be obligatory hosts of another group of barnacles,
5661-507: 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 tentacles which resemble glove fingers. The tentacles are organs which serve both for tactile sense and for
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#17327733566225772-428: The coast of Australia . 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 Lophelia which can survive as deep as 3,300 metres (10,800 feet; 1,800 fathoms). Some have been found as far north as
5883-440: The colony during storms or other disruptions. The separated individuals can start new colonies. 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
5994-476: The coral color. Such corals require sunlight and grow in clear, shallow water, typically at depths less than 60 metres (200 feet; 33 fathoms), but corals in the genus Leptoseris have been found as deep as 172 metres (564 feet; 94 fathoms). Corals are major contributors to the physical structure of the coral reefs that develop in tropical and subtropical waters, such as the Great Barrier Reef off
6105-797: 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. 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
6216-591: The coral itself together with its symbiont zooxanthellae (photosynthetic dinoflagellates), as well as its associated bacteria and viruses. Co-evolutionary patterns exist for coral microbial communities and coral phylogeny. 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 ,
6327-598: 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
6438-578: The dangers posed by the climate crisis. Corals are colonial modular organisms formed by asexually produced and genetically identical modules called polyps. Polyps are connected by living tissue to produce the full organism. The living tissue allows for inter module communication (interaction between each polyp), which appears in colony morphologies produced by corals, and is one of the main identifying characteristics for a species of coral. There are two main classifications for corals: hard coral (scleractinian and stony coral) which form reefs by
6549-434: The dark portion of the night between sunset and moonrise. Over the lunar cycle, moonrise shifts progressively later, occurring after sunset on the day of the full moon. The resulting dark period between day-light and night-light removes the suppressive effect of moonlight and enables coral to spawn. The spawning event can be visually dramatic, clouding the usually clear water with gametes. Once released, gametes fertilize at
6660-483: The eastern coast of the United States. The coral can live with and without zooxanthellae (algal symbionts), making it an ideal model organism 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
6771-589: The end of the 20th century prompted new evolutionary hypotheses that were different from ones founded on skeletal data. Results of molecular studies explained a variety of aspects of the evolutionary biology of the Scleractinia, including connections between and within extant taxa, and supplied support for hypotheses about extant corals that are founded on the fossil record. The 1996 analysis of mitochondrial RNA undertaken by American zoologists Sandra Romano and Stephen Palumbi found that molecular data supported
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#17327733566226882-525: 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 ,
6993-426: 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. Whole colonies can reproduce asexually, forming two colonies with
7104-676: The form of "polyp bail-out", which may allow polyps to survive even though the parent colony dies. It involves the growth of the coenosarc to seal off the polyps, detachment of the polyps and their settlement on the seabed to initiate new colonies. In other species, small balls of tissue detach themselves from the coenosarc, differentiate into polyps and start secreting calcium carbonate to form new colonies, and in Pocillopora damicornis , unfertilised eggs can develop into viable larvae. The overwhelming majority of scleractinian taxa are hermaphroditic in their adult colonies. In temperate regions,
7215-400: 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
7326-444: The form of crystals of aragonite , however, a prehistoric scleractinian ( Coelosimilia ) had a non-aragonite skeletal structure which was composed of calcite . The structure of both simple and compound scleractinians is light and porous, rather than solid as is the case in the prehistoric order Rugosa . Scleractinians are also distinguished from rugosans by their pattern of septal insertion. In colonial corals, growth results from
7437-498: 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
7548-598: The genus Acropora: Scleractinia About 35, see text . Scleractinia , also called stony corals or hard corals , are marine animals in the phylum Cnidaria that build themselves a hard skeleton . The individual animals are known as polyps and have a cylindrical body crowned by an oral disc in which a mouth is fringed with tentacles. Although some species are solitary, most are colonial . The founding polyp settles and starts to secrete calcium carbonate to protect its soft body. Solitary corals can be as much as 25 cm (10 in) across but in colonial species
7659-638: The geological past, corals were very abundant. Like modern corals, their ancestors built reefs, some of which ended as great structures in sedimentary rocks . Fossils of fellow reef-dwellers algae, sponges, and the remains of many echinoids , brachiopods , bivalves , gastropods , 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. Corals first appeared in
7770-521: The hypothesis for an independent origin of the Scleractinia. Whether the early scleractinian corals were zooxanthellate is an open question. The phenomenon seems to have evolved independently on numerous occasions during the Tertiary, and the genera Astrangia , Madracis , Cladocora and Oculina , all in different families, each have both zooxanthellate and non-zooxanthellate members. The fact that zooxanthellate coral make up only about half of
7881-436: The increase in temperature and decrease in pH at a competitive rate to these pathogens responsible for coral bleaching, resulting in species loss. 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
7992-430: The individual mouths. This is surrounded by a single corallite wall, as is the case in the meandroid corallites of brain corals. Extratentacular budding always results in separate polyps, each with its own corallite wall. In the case of bushy corals such as Acropora , lateral budding from axial polyps form the basis of the trunk and branches. The rate at which a stony coral colony lays down calcium carbonate depends on
8103-463: The most diverse genus of hard coral, having over 120 species. Most species within this genus have polyps which are dimorphic: axial polyps grow rapidly and have lighter coloration, while radial polyps are small and are darker in coloration. In the Acropora genus, gamete synthesis and photosynthesis occur at the basal polyps, growth occurs mainly at the radial polyps. Growth at the site of
8214-418: 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
8325-644: The mouth. In a few stony corals, this is the primary method of feeding, and the tentacles are reduced or absent, an example being Acropora acuminata . Caribbean stony corals are generally nocturnal, with the polyps retracting into their skeletons during the day, thus maximising the exposure of the zooxanthellae to the light, but in the Indo-Pacific region, many species feed by day and night. Non-zooxanthellate corals are usually not reef-formers; they can be found most abundantly beneath about 500 m (1,600 ft) of water. They thrive at much colder temperatures and can live in total darkness, deriving their energy from
8436-637: 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
8547-645: The order is unusual, as symbiosis is almost always an all-or-nothing phenomenon. This symbiotic equilibrium suggests that there must be evolutionary processes simultaneously maintaining and limiting symbiotic relationships. This is likely because despite the energetic benefits it provides, photosymbiosis appears to be an evolutionary disadvantage during mass extinctions. Traits that generally enable corals to survive mass extinction include deep-water or large habitat range, non-symbiotic, solitary or small colonies, and bleaching resistance, all of which tend to characterize azooxanthellate (non-symbiotic) corals. Endosymbionts, on
8658-523: The other hand, which rely on specialized conditions and access to light to survive, are especially vulnerable to prolonged darkness, temperature change, and eutrophication, all of which have been hallmarks of past mass extinctions. This makes zooxanthellate coral especially vulnerable to unstable conditions. Therefore, it is possible that coral and zooxanthellate coevolved loosely, with the relationship dissolving when advantages decreased, then reforming when conditions stabilized. The taxonomy of Scleractinia
8769-428: The outer surface of the skeleton and completely covering it. These sheets are continuous with the body wall of the polyps, and include extensions of the gastrovascular cavity, so that food and water can circulate between all the different members of the colony. In colonial species, the repeated asexual division of the polyps causes the corallites to be interconnected, thus forming the colonies. Also, cases exist in which
8880-399: The polyps are usually only a few millimetres in diameter. These polyps reproduce asexually by budding , but remain attached to each other, forming a multi-polyp colony of clones with a common skeleton, which may be up to several metres in diameter or height according to species. The shape and appearance of each coral colony depends not only on the species, but also on its location, depth,
8991-416: 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
9102-463: The production of a free-swimming planula larva that eventually settles on the seabed to undergo metamorphosis into a polyp. In colonial species, this initial polyp then repeatedly divides asexually, to give rise to the entire colony. The most common means of asexual reproduction in colonial stony corals is by fragmentation. Pieces of branching corals may get detached during storms, by strong water movement or by mechanical means, and fragments fall to
9213-412: The radial polyps encompasses two processes: asexual reproduction via mitotic cell proliferation , and skeleton deposition of the calcium carbonate via extra cellular matrix (EMC) proteins acting as differentially expressed (DE) signaling genes between both branch tips and bases. These processes lead to colony differentiation , which is the most accurate distinguisher between coral species. In
9324-440: 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 proteins (HSP) that are also in the category of DE across species. These HSPs help corals combat the increased temperatures they are facing which lead to protein denaturing, growth loss, and eventually coral death. Approximately 33% of coral species are on
9435-410: The risk of self-fertilization. Immediately after spawning, the eggs are delayed in their capability for fertilization until after the release of polar bodies. This delay, and possibly some degree of self-incompatibility, likely increases the chance of cross-fertilization. A study of four species of Scleractinia found that cross-fertilization was actually the dominant mating pattern, although three of
9546-642: The rules of nomenclature, the name given to the first described species has precedence over the rest, even when that description is poor, and the environment and even sometimes the country of the type specimen is unknown. Even the concept of "the species" is suspect, with regard to corals which have large geographical ranges with a number of sub-populations; their geographic boundaries merge with those of other species; their morphological boundaries merge with those of other species; and there are no definite distinctions between species and subspecies. The evolutionary relationships among stony corals were first examined in
9657-406: 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
9768-563: The same night even when multiple species are present. Synchronous spawning may form hybrids and is perhaps involved in coral speciation . Environmental cues that influence the release of gametes into the water vary from species to species. The cues involve temperature change, lunar cycle , day length , and possibly chemical signalling. Other factors that affect the rhythmicity of organisms in marine habitats include salinity, mechanical forces, and pressure or magnetic field changes. Mass coral spawning often occurs at night on days following
9879-470: The scleractinian skeleton is radial or biradial . This pattern of septal insertion is termed "cyclic" by paleontologists. By contrast, in some fossil corals, adjacent septa lie in order of increasing age, a pattern termed serial and produces a bilateral symmetry . Scleractinians secrete a stony exoskeleton in which the septa are inserted between the mesenteries in multiples of six. All modern scleractinian skeletons are composed of calcium carbonate in
9990-469: The sea bed. In suitable conditions, these are capable of adhering to the substrate and starting new colonies. Even such massive corals as Montastraea annularis have been shown to be capable of forming new colonies after fragmentation. This process is used in the reef aquarium hobby to increase stock without the necessity to harvest corals from the wild. Under adverse conditions, certain species of coral resort to another type of asexual reproduction in
10101-481: The sea where fertilisation takes place, and the planula larvae drift as part of the plankton , but a few species brood their eggs. Asexual reproduction is mostly by fragmentation , when part of a colony becomes detached and reattaches elsewhere. Stony corals occur in all the world's oceans. Much of the framework of modern coral reefs is formed by scleractinians. Reef-building or hermatypic corals are mostly colonial; most of these are zooxanthellate and are found in
10212-507: The septa were arranged serially rather than cyclically. However, it may be that similarities of scleractinians to rugosans are due to a common non-skeletalized ancestor in the early Paleozoic. Alternatively, scleractinians may have developed from a Corallimorpharia -like ancestor. It seems that skeletogenesis may have been associated with the development of symbiosis and reef formation, and may have occurred on more than one occasion. DNA sequencing appears to indicate that scleractinian corals are
10323-486: The septal trabeculae . In 1943, the American zoologists Thomas Wayland Vaughan and John West Wells , and Wells again in 1956, used the patterns of the septal trabeculae to divide the group into five suborders . In addition, they considered polypoid features such as the growth of the tentacles. They also distinguished families by wall type and type of budding . The 1952 classification by French zoologist J. Alloiteau
10434-529: The shallow waters into which sunlight penetrates. Other corals that do not form reefs may be solitary or colonial; some of these occur at abyssal depths where no light reaches. Stony corals first appeared in the Middle Triassic , but their relationship to the tabulate and rugose corals of the Paleozoic is currently unresolved. In modern times stony corals numbers are expected to decline due to
10545-409: 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 , 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
10656-433: The skeletons allow to differentiate zooxanthellae and non-zooxanthellae specimens. 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
10767-511: The species were also capable of self-fertilization to varying extents. There is little evidence on which to base a hypothesis about the origin of the scleractinians; plenty is known about modern species but very little about fossil specimens, which first appeared in the record in the Middle Triassic ( 240 million years ago ). It was not until 25 million years later that they became important reef builders, their success likely
10878-498: The species, but some of the branching species can increase in height or length by around 10 cm (4 in) a year (about the same rate as human hair grows). Other corals, like the dome and plate species, are more bulky and may only grow 0.3 to 2 cm (0.1 to 0.8 in) per year. The rate of aragonite deposition varies diurnally and seasonally. Examination of cross sections of coral can show bands of deposition indicating annual growth. Like tree rings, these can be used to estimate
10989-413: 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
11100-558: The surface of the peristome, or may be projecting and trumpet-shaped. 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
11211-449: The traditional families were plausible but that the suborders were incorrect. They also established that stony corals are monophyletic, including all the descendants of a common ancestor , but that they are divided into two groups, the robust and complex clades. Veron suggested that both morphological and molecular systems be used in future classification schemes. All Scleractinian corals (excluding fossils) are listed under Appendix II of
11322-454: 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. Many corals, as well as other cnidarian groups such as sea anemones form
11433-441: The usual pattern is synchronized release of eggs and sperm into the water during brief spawning events, often related to the phases of the moon. In tropical regions, reproduction may occur throughout the year. In many cases, as in the genus Acropora , the eggs and sperm are released in buoyant bundles which rise to the surface. This increases the concentration of sperm and eggs and thus the likelihood of fertilization , and reduces
11544-466: 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 inner layer as the endoderm . Between ectoderm and endoderm is a supporting layer of gelatinous substance termed mesoglea , secreted by
11655-427: The water where they meet and fertilize to spread offspring. Corals often synchronize their time of spawning. This reproductive synchrony is essential so that male and female gametes can meet. Spawning frequently takes place in the evening or at night, and can occur as infrequently as once a year, and within a window of 10–30 minutes. Synchronous spawning is very typical on the coral reef, and often, all corals spawn on
11766-542: The water's surface and form a microscopic larva called a planula , typically pink and elliptical in shape. A typical coral colony needs to release several thousand larvae per year to overcome the odds against formation of a new colony. Studies suggest that light pollution desynchronizes spawning in some coral species. In areas such as the Red Sea , as many as 10 out of 50 species may be showing spawning asynchrony, compared to 30 years ago. The establishment of new corals in
11877-559: 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
11988-738: The wild. In a well-maintained reef aquarium, finger-sized fragments can grow into medicine ball-sized colonies in one to two years. Captive specimens are steadily undergoing changes due to selection which enable them to thrive in the home aquarium. In some cases, fragments of captive specimens are used to repopulate barren reefs in the wild. Acropora species are challenging to keep in a home aquarium. They require bright light, stable temperatures, regular addition of calcium and alkalinity supplements, and clean, turbulent water. Common parasites of colonies in reef aquariums are "Acropora-eating flatworms" Amakusaplana acroporae , and "red bugs" ( Tegastes acroporanus ). The following species are recognised in
12099-408: The world's primary reef -builders. Ahermatypic corals are either colonial or solitary and are found in all regions of the ocean and do not build reefs. Some live in tropical waters but some inhabit temperate seas, polar waters, or live at great depths, from the photic zone down to about 6,000 m (20,000 ft). Scleractinians fall into one of two main categories: In reef-forming corals,
12210-410: The zooxanthellae is their ability to photosynthesize which supplies corals with the products of photosynthesis, including glucose, glycerol, also amino acids, which the corals can use for energy. Zooxanthellae also benefit corals by aiding in 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)
12321-697: Was built on these earlier systems but included more microstructural observations and did not involve the anatomical characters of the polyp. Alloiteau recognized eight suborders. In 1942, W.H. Bryan and D. Hill stressed the importance of microstructural observations by proposing that stony corals begin skeletal growth by configuring calcification centers, which are genetically derived. Therefore, diverse patterns of calcification centers are vital to classification. Alloiteau later showed that established morphological classifications were unbalanced and that there were many examples of convergent evolution between fossils and recent taxa . The rise of molecular techniques at
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