Scleractinia


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 across but in colonial species 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, 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 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 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 the effects of global warming and ocean acidification.

Anatomy

Scleractinian corals may be solitary or colonial. Colonies can reach considerable size, consisting of a large number of individual polyps.

Soft parts

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 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 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 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 the adjacent colonies of the same species form a single colony by fusing. Most colonial species have very small polyps, ranging from in diameter, although some solitary species may be as large as.

Skeleton

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 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 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 the form of crystals of aragonite, however, a prehistoric scleractinian 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.

Growth

In colonial corals, growth results from 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 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 the species, but some of the branching species can increase in height or length by around a year. Other corals, like the dome and plate species, are more bulky and may only grow 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 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 long and have more than a thousand septa.

Distribution

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 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.

Ecology

Scleractinians fall into one of two main categories:
  • Reef-forming or hermatypic corals, which mostly contain zooxanthellae;
  • Non-reef-forming or ahermatypic corals, which mostly do not contain zooxanthellae
In reef-forming corals, 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 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 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 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 of water. They thrive at much colder temperatures and can live in total darkness, deriving their energy from 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, the pyrgomatids, but a recent study recorded evidence of living pyrgomatids in stylasterids, casting doubt on this idea.