CORAL REEFS

Coral reefs are underwater structures made from calcium

carbonate secreted by corals. Corals are colonies of tiny living animals
found in marine waters containing few nutrients. Most coral reefs are
built from stony corals, and are formed by polyps that live together in
groups. The polyps secrete a hard carbonate exoskeleton which
provides support and protection for the body of each polyp. Reefs grow
best in warm, shallow, clear, sunny and agitated waters.

Often called “rainforests of the sea”, coral reefs form some of

the most diverse ecosystems on earth. They occupy less than one
percent of the world ocean surface, about half the area of France, yet
they provide a home for twenty-five percent of all marine species,
including fish, molluscs, echinoderms andsponges.[1]

Paradoxically, coral reefs flourish even though they are surrounded by ocean waters that provide few
nutrients. They are most commonly found at shallow depths in tropical waters, particularly in the Pacific Ocean,
but deep water and cold water corals also exist on smaller scales.
Coral reefs deliver ecosystem services to tourism, fisheries and shoreline protection. The annual global
economic value of coral reefs has been estimated at $30 billion. However, coral reefs are fragile ecosystems,
partly because they are very sensitive to water temperature. They are under threat from climate change, ocean
acidification, blast fishing, cyanide fishing for aquarium fish, overuse of reef resources, and harmful land-use
practices. High nutrient levels such as those found in runoff from agricultural areas can harm reefs by
encouraging excess algae growth.[2]

Formation

Most coral reefs were formed after the last glacial period when melting ice caused the sea level to rise
and flood the continental shelves. This means that most coral reefs are less than 10,000 years old. As coral reef
communities were established on the shelves, they built reefs that grew upwards, keeping pace with the rise in
sea level. Reefs that didn't keep pace could become drowned reefs, covered by so much water that there was
insufficient light for further survival.[3]

Coral reefs are also found in the deep sea away from the continental shelves, around oceanic
islands and as atolls. The vast majority of these ocean coral islands are volcanic in origin. The few exceptions
have tectonic origins where plate movements have lifted the deep ocean floor on the surface.
In 1842 Charles Darwin published his first monograph, The Structure and Distribution of Coral Reefs.[4] There he
set out his theory of the formation of atoll reefs, an idea he conceived during the voyage of the Beagle. His
theory was that atolls were formed by the uplift and subsidence of the Earth's crust under the oceans.[5] Darwin’s
theory sets out a sequence of three stages in atoll formation. It starts with a fringing reef forming around an
extinct volcanic island as the island and ocean floor subsides. As the subsidence continues, the fringing reef
becomes a barrier reef, and ultimately an atoll reef.

As the subsidence Ultimately the island sinks

As the island and ocean below the sea, and the
floor subside, coral growth continues the fringing reef
becomes a larger barrier barrier reef becomes an
Darwin’s theory starts builds a fringing reef, often atoll enclosing an open
including a shallow lagoon reef further from the shore
with a volcanic with a bigger and deeper lagoon
island which becomes between the land and the
main reef lagoon inside
extinct

Darwin predicted that underneath each lagoon would be a bed rock base, the remains of the original
volcano. Subsequent drilling has proved this correct. Darwin's theory followed from his understanding that
coral polyps thrive in the clean seas of the tropics where the water is agitated, but can only live within a limited
depth of water, starting just below low tide. Where the level of the underlying land stays the same, the corals
grow around the coast to form what he called fringing reefs, and can eventually grow out from the shore to
become a barrier reef.

A fringing reef can take ten thousand years to form, and an atoll can take up to 30 million years[6]
Where the land is rising, fringing reefs can grow around the coast, but coral raised above sea level dies and
becomes white limestone. If the land subsides slowly, the fringing reefs keep pace by growing upwards on a
base of dead coral, forming a barrier reef enclosing a lagoon between the reef and the land. A barrier reef can
encircle an island, and once the island sinks below sea level a roughly circular atoll of growing coral continues to
keep up with the sea level, forming a central lagoon. Barrier reefs and atolls don't usually form complete circles,
but are broken in places by storms. Should the land subside too quickly or sea level rise too fast, the coral dies
[5][7]
as it is below its habitable depth.

In general, the two main variables determining the geomorphology, or shape, of coral reefs are the
nature of the underlying substrate on which they rest, and the history of the change in sea level relative to that
substrate.
 As an example of how coral reefs have formed on continental shelves, the current living reef structure of
the Great Barrier Reef began growing about 20,000 years ago. The sea level was then 120 metres (390 ft) lower
[8][9]
than it is today. As the sea level rose, the water and the corals encroached on what had been the hills of the
coastal plain. By 13,000 years ago the sea level was 60 metres (200 ft) lower than at present, and the hills of
the coastal plains were, by then,continental islands. As the sea level rise continued most of the continental
islands were submerged. The corals could then overgrow the hills, forming the present cays and reefs. The sea
level on the Great Barrier Reef has not changed significantly in the last 6,000 years, [9] and the age of the
present living reef structure is estimated to be between 6,000 and 8,000 years. [10] Although the Great Barrier
Reef formed along a continental shelf, and not around a volcanic island, the same principles apply as outlined
by Darwin's theory above. The Great Barrier Reef development has stopped at the barrier reef stage, since
Australia is not about to submerge. It has formed the world's largest barrier reef, 300–1000 metres (330-1100
[11]
yards) from shore, and 2,000 kilometres (1,200 mi) long.

Healthy coral reefs grow horizontally from 1 to 3 centimetres (0.39 to 1.2 in) per year, and grow
vertically anywhere from 1 to 25 centimetres (0.4–12 in) per year; however, they are limited to growing above a
depth of 150 metres (490 ft) due to their need for sunlight, and cannot grow above sea level.[12]

Types

The three principal reef types, as detailed in the previous section, are:
 Fringing reef – a reef that is directly attached to a shore or borders it with an intervening shallow channel or
lagoon.
 Barrier reef – a reef separated from a mainland or island shore by a deep lagoon.
 Atoll reef – a more or less circular or continuous barrier reef extending all the way around a lagoon without
a central island.

A small atoll in the Maldives. Inhabited cay in the Maldives

Other reef types or variants are:
 Patch reef – an isolated, comparatively small reef outcrop, usually within a lagoon or embayment, often
circular and surrounded by sand or seagrass. Patch reefs are common.
 Apron reef – a short reef resembling a fringing reef, but more sloped; extending out and downward from a
point or peninsular shore.
 Bank reef – a linear or semi-circular shaped-outline, larger than a patch reef.
 Ribbon reef – a long, narrow, somewhat winding reef, usually associated with an atoll lagoon.
 Table reef – an isolated reef, approaching an atoll type, but without a lagoon.
 Microatolls – certain species of corals form communities called microatolls. The vertical growth of
microatolls is limited by average tidal height. By analysing the various growth morphologies, microatolls can
be used as a low resolution record of patterns of sea level change. Fossilized microatolls can also be dated
using radioactive carbon dating. Such methods have been used to reconstruct Holocene sea levels.[13]
 Cays – small, low-elevation, sandy islands formed on the surface of a coral reef. Material eroded from the
reef piles up on parts of the reef or lagoon, forming an area above sea level. Plants can stabilize cays
enough for them to be habitable by humans. Cays occur in tropical environments throughout the
Pacific, Atlantic and Indian Oceans (including in the Caribbean and on the Great Barrier Reef and Belize
Barrier Reef), where they provide habitable and agricultural land for hundreds of thousands of people. Their
surrounding reef ecosystems also provide food and building materials for island inhabitants.
 When a coral reef cannot keep up with the sinking of a volcanic island, a seamount or guyot is formed.
Seamounts and guyots are below the surface of the ocean and may host many species, depending on their
location and depth. Seamounts are rounded at the top and guyots are flat. The flat top of the guyot, also
called a tablemount, is due to erosion by waves, winds, and atmospheric processes.

Zones

The three major zones of a coral reef: the fore reef, reef crest, and the back reef

Coral reef ecosystems contain a number of distinctive zones that represent different kinds of habitats for
fishes and invertebrate animals. Usually, three major zones are recognized: the fore reef (outermost and
deepest part of the reef), reef crest (shallowest and narrowest zone over which ocean waves break), and the
back reef (behind the reef crest and closest to shore, with calm protected waters) which is also frequently
referred to as the reef lagoon.

All three zones are physically and ecologically interconnected to some extent, with reef life and oceanic
processes creating ample opportunities for exchange of ocean water, sediments, nutrients, and marine life
between one another. Thus, they should properly be viewed as integrated components of the coral reef
ecosystem, each playing a role in the support of the abundant and diverse fish assemblages that characterize
coral reefs.

Most coral reefs exist in shallow waters less than fifty metres deep. Some are found on tropical
continental shelves where cool, nutrient rich upwelling does not occur, such as Great Barrier Reef. Others are
found in the deep ocean surrounding islands or as atolls, such as in theMaldives. The reefs surrounding islands
form when islands subside into the ocean, and atolls form when an island subsides below the surface of the
sea. Moyle and Cech[14] distinguish six zones, though most reefs possess only some of the zones.

Water in the reef surface zone is often agitated. This diagram represents a reef on a continental shelf. The
water waves at the left travel over the off-reef floor until they encounter the reef slope or fore reef. Then the
waves pass over the shallow reef crest. When a wave enters shallow water itshoals, that is, it slows down and
the wave height increases.

 The reef surface is the shallowest part of the reef. It is subject to the constant surge of waves and the rise
and fall of tides. When water waves pass over shallow areas, they shoal, as shown in the diagram at the
right. This means that the water in the reef surface zone is often agitated. These are the precise condition
under which coral flourish. Shallowness means there is plenty of light for photosynthesis, and agitated water
promotes the ability of coral to feed on plankton. However other organisms, such as fish and invertebrates,
must be able to withstand the robust conditions to flourish in this zone.

 The off-reef floor is the shallow sea floor surrounding a reef. This zone applies to reefs on continental
shelves. Reefs around tropical islands and atolls drop abruptly to great depths, and don't have an off-reef
floor. Usually sandy, the off-reef floor often supports seagrass meadows which are important foraging areas
for reef fish.
 The reef drop-off is, for its first 50 metres, habitat for many reef fish who find shelter on the cliff face and
plankton in the water nearby. The drop-off zone applies mainly to the reefs surrounding oceanic islands and
atolls.

 The reef face is the zone above the reef floor or the reef drop-off. "It is usually the richest habitat for fish
and invertebrates. Its complex growths of coral and calcareous algae provide innumerable cracks and
crevices for protection, and the abundant invertebrates and epiphytic algae provide an ample source of
food."[14]

 The reef flat – sandy bottomed flat can be behind the main reef, containing chunks of coral. "The reef flat
may be a protective area bordering a lagoon, or it may be a flat, rocky area between the reef and the shore.
In the former case, the number of fish species living in the area often is the highest of any reef zone." [14]
 The reef lagoon – "many coral reefs completely enclose an area, thereby creating a quiet-water lagoon that
usually contains small patched of reef."[14]

However, the "topography of coral reefs is constantly changing. Each reef is made up of irregular patches of
algae, sessile invertebrates, and bare rock an sand. The size, shape and relative abundance of these patches
changes from year to year in response to the various factors that favour one type of patch over another.
Growing coral, for example, produces constant change in the fine structure of reefs. On a larger scale, tropical
storms may knock out large sections of reef and cause boulders on sandy areas to move." (Connell 1978)

Distribution

Locations of coral reefs.

Boundary for 20 °C isotherms. Most corals live within

this boundary. Note the cooler waters caused by
upwelling on the south west coast of Africa and off the
coast of Peru.
 This map shows areas of upwelling in red. Coral reefs are
not found in coastal areas where colder and nutrient-rich
upwellings occur

Coral reefs are estimated to cover 284,300 square kilometers (109,800 sq mi), which is just under one
percent of the surface area occupied by the world oceans. The Indo-Pacific region (including the Red
Sea, Indian Ocean, Southeast Asia and the Pacific) account for 91.9% of this total. Southeast Asia accounts for
32.3% of that figure, while the Pacific including Australia accounts for 40.8%. Atlantic and Caribbean coral reefs
only account for 7.6%.[15]

Although corals exist both in temperate and tropical waters, shallow-water reefs form only in a zone
extending from 30° N to 30° S of the equator. Tropical corals do not grow at depths of over 50 meters (160 ft).
The optimum temperature for most coral reefs is 26–27 °C, and few reefs exist in waters below
18 °C.[16] However reefs in the Persian Gulf have adapted to temperatures of 13 °C in winter and 38 °C in
summer.[17]

Deep water coral is more still exceptional since it can exist at greater depths and colder temperatures.
Although deep water corals can form reefs, very little is known about them.

Coral reefs are rare along the American west coast, as well as along the African west coast. This is due
primarily to upwelling and strong cold coastal currents that reduce water temperatures in these areas
(respectively the Peru, Benguela and Canary streams).[18] Corals are seldom found along the coastline of South
Asia from the eastern tip of India (Madras) to the border of Bangladesh and Myanmar. [15] They are also rare
along the coast around north-eastern South America and Bangladesh due to the freshwater release from
the Amazon and Ganges Rivers respectively.

Principal coral reefs and reef areas of the world

 The Great Barrier Reef - largest coral reef system in the world, Queensland, Australia.
 The Belize Barrier Reef - second largest in the world, stretching from southern Quintana Roo, Mexico along
the coast of Belize to the Bay Islands of Honduras.
 The New Caledonia Barrier Reef - second longest double barrier reef in the world, with a length of about
1,500 kilometers (930 mi).
 The Andros, Bahamas Barrier Reef - third largest in the world, following the east coast of Andros Island,
Bahamas, between Andros and Nassau.
 The Red Sea Coral Reef - located off the coast of Israel, Egypt, Sudan, Eritrea, Djibouti, Somalia, Jordan,
Saudi Arabia, and Yemen.
 Pulley Ridge - deepest photosynthetic coral reef, Florida.
 Numerous reefs scattered over the Maldives.
 Ghe Raja Ampat Islands in Indonesia's West Papua province offer the highest known marine diversity.[19]

Biology

Anatomy of a coral polyp.

Live coral can be thought of as small live animals embedded in calcium carbonate. It is a mistake to
think of coral as plants or rocks. Coral consists of accumulations of individual animals called polyps, arranged in
diverse shapes.[20] Polyps are usually tiny, but they can range in size from a pinhead to a foot across. Reefs
[21][22]
grow as polyps along with other organisms deposit calcium carbonate, the basis of coral, as a skeletal
structure beneath and around themselves, pushing the coral's "head" or polyps upwards and
[23]
towards. Waves, grazing fish (such as parrotfish), sea urchins, sponges, and other forces and organisms act
as bioeroders, breaking down coral skeletons into fragments that settle into spaces in the reef structure or form
sandy bottoms in associated reef lagoons. Many other organisms living in the reef community contribute
skeletal calcium carbonate in the same manner.Coralline algae are important contributors to reef structure in
those parts of the reef subjected to the greatest forces by waves (such as the reef front facing the open ocean).
These algae deposit limestone in sheets over the reef surface, thereby strengthening it.

Reef-building or hermatypic corals are found only in the photic zone (above 50 m depth), the depth to
which sufficient sunlight penetrates the water forphotosynthesis to occur. Coral polyps do not photosynthesize,
but have a symbiotic relationship with single-celled organisms called zooxanthellae; these cells within the
tissues of the coral polyps carry out photosynthesis and produce excess organic nutrients that are then used by
the coral polyps. Because of this relationship, coral reefs grow much faster in clear water, which admits more
sunlight. Indeed, the relationship is responsible for coral reefs in the sense that without their symbionts, coral
growth would be too slow for the corals to form impressive reef structures. Corals get up to 90% of their
[24]
nutrients from their zooxanthellae symbionts.

Table coral

Close up of polyps arrayed on a coral, waving their tentacles. There can

be thousands of polyps on a single coral branch.

Corals can reproduce both sexually and asexually. An individual polyp may use both reproductive
modes within its lifetime. Corals reproduce sexually by either internal or external fertilization. The reproductive
cells are found on the mesentery membranes that radiate inward from the layer of tissue that lines the stomach
cavity. Some mature adult corals are hermaphroditic; others are exclusively male or female. A few even change
sex as they grow.

Internally fertilized eggs develop in the polyp for a period ranging from days to weeks. Subsequent
development produces a tiny larva, known as a planula. Externally fertilized eggs develop during synchronized
spawning. Polyps release eggs and sperm into the water simultaneously. Eggs disperse over a large area.
Spawning depends on four factors: time of year, water temperature, and tidal and lunar cycles. Spawning is
most successful when there is little variation between high and low tides. The less water movement, the better
the chance for fertilization. Ideal timing occurs in the spring. Release of eggs or planula larvae usually occurs at
night and is sometimes in phase with the lunar cycle (3–6 days after a full moon). The period from release to
settlement lasts only a few days, but some planulae can survive afloat for several weeks (7, 14). They are
vulnerable to heavy predation and adverse environmental conditions. For the lucky few who survive to attach to
substrate, the challenge comes from competition for food and space.
There are about one thousand species of coral, which build different shapes such as wrinkled brains,
cabbages, table tops, stag antlers, wire strands and pillars.

Staghorn coral

Brain coral Spiral wire coral Pillar coral

Darwin's paradox

Darwin's paradox
Coral... seems to proliferate when ocean waters are warm, poor, clear and agitated, a fact which Darwin
had already noted when he passed through Tahiti in 1842.

This constitutes a fundamental paradox, shown quantitatively by the apparent impossibility of balancing
input and output of the nutritive elements which control the coral polyp metabolism.
Recent oceanographic research has brought to light the reality of this paradox by confirming that
theoligotrophy of the ocean euphotic zone persists right up to the swell-battered reef crest. When you approach
the reef edges and atolls from the quasi-desert of the open sea, the near absence of living matter suddenly
becomes a plethora of life, without transition. So why is there something rather than nothing, and more
precisely, where do the necessary nutrients for the functioning of this extraordinary coral reef machine come
[25]
from ? — Francis Rougerie

During his voyage on the Beagle, Darwin described tropical coral reefs as oases in the desert of the
ocean. He reflected on the paradox that tropical coral reefs, which are among the richest and
most diverse ecosystems on earth, flourish when they are surrounded and supported by tropical ocean waters
that provide hardly any nutrients. It has been a challenge for scientists to explain this paradox.
Coral reefs cover just under one percent of the surface of the world’s ocean, yet they support over one-quarter
of all marine species. This huge number of species results in complex food webs, with large predator fish eating
smaller forage fish that eat yet smaller zooplankton and so on. However, all food webs eventually depend
on plants, which are the primary producers. The primary productivity on a coral reef is very high, resulting in a
−2 −1 [26]
typical lbiomass production of 5-10g C m day .
 Tropical waters are often described as crystal clear. This is because they are deficient in nutrients and
drifting plankton. The sun shines year round in the tropics, warming the surface ocean layer so it is less dense
than subsurface layers. The warmer water is separated from the cooler water by a stable thermocline, where the
temperature makes a rapid change. This keeps the warm surface waters floating above the cooler deeper
waters. There is little exchange between these layers. Organisms that die in aquatic environments generally
sink to the bottom where they decompose. This decomposition releases nutrients in the form
of nitrogen (N), phosphorus (P) and potassium (K). These nutrients are necessary for plant growth, but in the
tropics they are not directly recycled back to the surface.[7]

Plants are the base of the food chain, and need sunlight and nutrients if they are to grow. In the ocean
these plants are mainly a type of plankton, microscopic phytoplankton which drift in the water column. They
need sunlight for photosynthesis, which powers carbon fixation, so they are found only in the surface waters.
But they also need nutrients. Phytoplankton rapidly use any nutrients in the surface waters, and in the tropics
these nutrients are not usually replaced because of the thermocline.[7]

Coral polyps

The situation with coral reefs is different. The lagoons that are formed by the upward growth of coral
reefs fill in with material eroded from the reef and the island. They become havens for marine life, providing
protection from waves and storms.

Most importantly, nutrients are recycled, and not lost like they are in the open ocean. In coral reefs and
lagoons, the producers include phytoplankton as well as marine worms, seaweed, and coralline algae,
[27]
especially small types called turf algae, which pass their nutrients to the corals. The phytoplankton are eaten
by fish and crustaceans, who also pass nutrients along the food web. Recycling ensures that fewer nutrients are
needed overall to support the community.

Corals harbour numerous symbiotic organisms. In particular, there is a remarkable symbiosis between
coral and a microscopic algae, the single celldinoflagellate known as a zooxanthella. The zooxanthella forms
an endosymbiosis with a coral polyp, that is, it lives within the tissues of the polyp. There it absorbs solar energy
with special pigments, using photosynthesis to supply the polyp with organic nutrients in the form of glucose,
glycerol, and amino acids.[28] Zooxanthellae can provide up to 90% of a coral’s energy requirements. [29] In return,
as an example of mutualism, the coral provides the zooxanthellae, averaging one million for every cubic
centimetre of coral, with a relatively safe place to live and a constant supply of the carbon dioxide it needs for
photosynthesis.

The colour of corals depends on the type of zooxanthella they host.

Corals also absorb nutrients, including inorganic nitrogen and phosphorus, directly from the water. Many
corals extend their tentacles at night to catchzooplankton that brush them when the water is agitated.

Zooplankton provides the polyp with nitrogen, and the polyp shares some of the nitrogen with the
zooxanthellae, which also require this element.[27] The pigments in different species of zooxanthellae give corals
their different colours. Coral which loses its zooxanthellae becomes white and is said to be bleached, a
condition which unless corrected can lead to the death of the coral.

A 2001 paper reported that sponges are another key to explaining Darwin’s paradox. These sponges
live in crevices in the coral reefs. They are efficient filter feeders, and in the Red Sea they consume about sixty
percent of the phytoplankton that drifts by. The nutrients the sponges absorb from the phytoplankton are then
excreted in a form the corals can use.[30]

Most coral polyps are nocturnal feeders. Here, in the dark, polyps have
extended their tentacles to feed on zooplankton

Researchers in 2002 explained why coral thrives better in agitated waters. They found the roughness of
coral surfaces is the key. Normally there is a boundary layer of still water around a submerged object which acts
as a barrier. But when waves break on the extremely rough edges of corals the boundary layer is disrupted,
allowing the corals access to the few nutrients that are there. The researchers claim that turbulent water
promotes rapid reef growth and lots of branching. Although coral ecosystems are great at recycling, with the
wastes of one species becoming the food of another, the researchers also claim that, without the nutritional
gains achieved by rough coral surfaces, even the most effective recycling would leave corals wanting in
nutrients.[31]

In 2004, another symbiotic organism, a bacterium called Cyanobacteria, was discovered to provide
[32]
soluble nitrates for the reef via nitrogen fixation. Coral reefs also often depend on other habitats, such
as seagrass meadows and mangrove forests in the surrounding area for the supply of nutrients. Seagrass and
mangroves supply dead plants and animals which are rich in nitrogen and also serve to feed fish and animals
from the reef by supplying wood and vegetation. Reefs in turn protect mangroves and seagrass from waves and
produce sediment for the mangroves and seagrass to root in.[33]

Biodiversity

Tube sponges attracting cardinal fishes, glassfishes andwrasses.

Reefs are also home to a large variety of other organisms, including

fish, seabirds, sponges, Cnidarians (which includes some types of
corals and jellyfish), worms, crustaceans (including shrimp, cleaner
shrimp, spiny
lobsters and crabs), molluscs (including cephalopods), echinoderms
(including starfish, sea urchins and sea cucumbers), sea
squirts, sea turtles and sea snakes. Aside from
humans, mammals are rare on coral reefs, with
visiting cetaceans such as dolphins being the main exception. A few
of these varied species feed directly on corals, while others graze
on algae on the reef and participate in complex food webs.[15][27]
Researchers have found evidence of algae dominance in locations
of healthy coral reefs. In surveys done around largely uninhabited
[34]
US Pacific islands, algae inhabit a large percentage of surveyed coral locations. The algae population
consists of turf algae, coralline algae, and macroalgae.

Fish
Coral reefs are home to a variety of tropical or reef fish which can be distinguished. These include:
 fish that adjust the coral (such as Labridae and parrotfish) These types of fish feed either on small animals
living near the coral, seaweed, or on the coral itself. Fish that feed on small animals include cleaner
fish (these fish feed between the jaws of larger predatory fish), bullet fish and Balistidae (these eat sea
urchins) while seaweed eating fish include the Pomacentridae (damselfishes). Serranidae cultivate the
seaweed by removing creatures feeding on it (as sea urchins), and they remove inedible seaweeds. Fish
that eat coral include parrotfish andbutterflyfish.
 fish that cruise the boundaries of the reef or nearby seagrass meadows. These include predatory fish such
as pompanos, groupers, Horse mackerels, certain types of shark, Epinephelus
marginatus, barracudas, snappers, ...) They also include herbivorous and plankton-eating fish. Fish eating
seagrass include Horse mackerel, snapper, Pagellus, Conodon, ... Fish eating plankton include Caesio,
manta ray, chromis, Holocentridae, pterapogon kauderni, ...

Organisms can cover every square inch of a coral reef. Generally, fish that
swim in coral reefs are as colourful as the reef itself. Examples are the
beautiful parrotfish, angelfish, damselfish, Pomacanthus
paru, Clinidae and butterflyfish. At night, some change to a less vivid color.
Besides colorful fish matching their environment, other fish (e.g., predatory
and herbivorous fish such as Lampanyctodes
hectoris, Holocentridae, Pterapogon kauderni, ...) as well as aquatic
animals (Comatulida, Crinoidea, Ophiuroidea, ...) emerge and become
active while others rest.

Other fish groups found on coral reefs

include groupers, grunts and wrasses. Over 4,000 species of fish inhabit
[15]
coral reefs. It has been suggested that the fish species that inhabit coral reefs are able to coexist in such high
numbers because any free living space is inhabited by the first planktonic fish larvae that find it in what has been
termed "a lottery for living space".[35]

Seabirds
Coral reef systems provide important habitats for seabird species, some endangered. For example, Midway
Atoll supports nearly three million seabirds, including two-thirds (1.5 million) of the global population of Laysan
Albatross, and one-third of the global population of black-footed albatross.[36] Each seabird species have specific
sites on the atoll where they nest. Altogether, 17 species of seabirds live on Midway. The short-tailed
albatross is the rarest, with fewer than 2,200 surviving after excessive feather hunting in the late nineteenth
century.[37]

Invertebrates
Invertebrates have their part in the food-chain of the reef. For example, sea urchins, Dotidae and sea slugs eat
seaweed. Some species of sea urchins, such as Diadema antillarum, can play a pivotal part in preventing algae
[38]
overrunning reefs. Hawksbill turtles, Nudibranchia and sea anemones eat sponges.
A number of invertebrates, collectively called cryptofauna, inhabit the coral skeletal substrate itself, either
boring into the skeletons (through the process of bioerosion) or living in pre-existing voids and crevices. Those
animals boring into the rock include sponges, bivalve molluscs, and sipunculans. Those settling on the reef
include many other species, particularly crustaceans and polychaete worms.[18]
Other
Sea snakes feed exclusively on fish and their eggs. Many tropical birds forage on reef fish, such
as herons, gannets, pelicans and boobies. Some land based reptiles can be intermittently associated with reefs,
such as monitor lizards, the marine crocodile and semi-aquatic snakes like Laticauda colubrina.

Schooling reef fish The whitetip reef sharkalmost

Caribbean reef
squid exclusively inhabits coral reefs
Banded coral shrimp

Green turtle Giant clam Soft coral, cup coral, sponges

and ascidians
Banded sea krait

Economic value

Coral reefs deliver ecosystem services to tourism, fisheries and coastline protection. The global
economic value of coral reefs has been estimated at $30 billion.[39] Coral reefs protect shorelines by absorbing
wave energy, and many small islands would not exist without their reef to protect them. According to the WWF,
the economic cost over a 25 year period of destroying one kilometre of coral reef is somewhere between
$137,000 and $1,200,000.[40] About 6 million tons of fish are taken each year from coral reefs. Well managed
coral reefs have an annual yield of 15 tons seafood on average per square kilometre. Southeast Asia's coral
reef fisheries alone yield about $ 2.4 billion annually from seafood.[40]

Issues

Island with fringing reef off Yap,Micronesia. Coral reefs are dying
[41]
around the world.
 Coral reefs are dying around the world.[41] In particular, coral mining, pollution (organic and non-
organic), overfishing, blast fishing and the digging of canalsand access into islands and bays are serious threats
to these ecosystems. Coral reefs also face high dangers from pollution, diseases, destructive fishing practices
[42]
and warming oceans." In order to find answers for these problems, researchers study the various factors that
impact reefs. The list of factors is long, including the ocean's role as a carbon dioxide sink, atmospheric
changes, ultraviolet light, ocean acidification, biological virus, impacts of dust stormscarrying agents to far flung
reefs, pollutants, algal blooms and others. Reefs are threatened well beyond coastal areas.
General estimates show approximately 10% world's coral reefs are already dead. [43][44][45] It is estimated
that about 60% of the world's reefs are at risk due to destructive, human-related activities. The threat to the
health of reefs is particularly strong in Southeast Asia, where 80% of reefs are endangered.

Protection

Coral reefs and fish in Papua New Guinea

Marine Protected Areas (MPAs) have become increasingly prominent for reef management. MPAs in
Southeast Asia and elsewhere around the world attempt to promote responsible fishery
management and habitat protection. Much like national parks and wild life refuges, MPAs prohibit potentially
damaging extraction activities. The objectives of MPAs are both social and biological, including reef restoration,
aesthetics, increased and protected biodiversity, and economic benefits. Conflicts surrounding MPAs involve
lack of participation, clashing views and perceptions of effectiveness, and funding.

Biosphere reserves are other protected areas that may protect reefs. Also, Marine parks, as well
as world heritage sites can protect reefs. World heritage designation can also play a vital role. For example
the Chagos archipelago, Sian Ka'an, the Great Barrier Reef, Henderson Island, theGalapagos islands, Belize's
Barrier reef and Palau have been designated as protected by nomination as a world heritage site.
In Australia, the Great Barrier Reef is protected by the Great Barrier Reef Marine Park Authority, and is the
subject of much legislation, including a Biodiversity Action Plan.

Inhabitants of Ahus Island, Manus Province, Papua New Guinea, have followed a generations-old
practice of restricting fishing in six areas of their reef lagoon. Their cultural traditions allow line fishing but not net
and spear fishing. The result is that both the biomass and individual fish sizes are significantly larger in these
areas than in places where fishing is unrestricted.[46][47]

Restoration

Low voltage electrical currents applied through seawater crystallize dissolved minerals onto steel
structures. The resultant white carbonate (aragonite) is the same mineral that makes up natural coral reefs.
Corals rapidly colonize and grow at accelerated rates on these coated structures. The electrical currents also
accelerate formation and growth of both chemical limestone rock and the skeletons of corals and other shell-
bearing organisms. The vicinity of the anode and cathode provides a high pH environment which inhibits the
growth of filamentous and fleshy algae, which compete with coral for space. The increased growth rates cease
[48]
when the mineral accretion process stops.

During mineral accretion, the settled corals display an increased growth rate, and size, and density, but
after the process is complete, growth rate and density return to levels that are comparable to naturally growing
corallites, and are about the same size or slightly smaller.[48]

In large restoration projects, depending on the type of coral, placement of propagated hermatypic
coral unto substrate is often done with metal pins, superglue or milliput.[49] Needle and thread can also attach A-
hermatype coral to substrate.[50] Concrete has also been used to restore large sections of broken coral reef.
Finally, special structures as reef balls can be placed to provide corals a base to grow on.

Organizations

Organizations which currently undertake coral reef/atoll restoration projects using simple methods of plant
propagation:
 Coral Cay
 Counterpart[51]
 U.S. Coral Reef Task Force (CRTF)
 National Coral Reef Institute (NCRI)
 US Department of Commerce’s National Oceanic and Atmospheric Administration (NOAA): Coral Reef
Conservation Program
 National Center for Coral Reef Research (NCORE)
 Reef Ball
 Southeast Florida Coral Reef Initiative (SEFCRI)
 Foundation of the peoples of the South Pacific
 WorldFishCenter: promotes sustainable mariculture techniques to grow reef organisms as tridacnidae
 Coral Restoration Foundation (CRF) : Adopt a Coral
Organizations which promote interest, provide knowledge bases about coral reef survival, and promote activities
to protect and restore coral reefs:
 Australian Coral Reef Society
 Beautiful Oceans
[52]
 Biosphere Foundation
 Chagos Conservation Trust
 Conservation Society of Pohnpei
 Coral Cay Conservation
 Coral Reef Care
 Coral Reef Alliance (CORAL)
 Coral Reef Targeted Research and Capacity Building for Management
 Coral Triangle Initiative
 Cousteau Society
 Crusoe Reef Society
 CEDAM International
 Earthwatch
 Environmental Defense Fund
 Environmental Solutions International
 Friends of Saba Marine Park
 Global Coral Reef Alliance (GCRA)
 Global Coral Reef Monitoring Network[53]
 Great Barrier Reef Marine Park Authority
 ICRAN Mesoamerican Reef Alliance
 International Marinelife Alliance
 International Society for Reef Studies
 Intercoast Network
 Kosrae Conservation and Safety Organization
 Marine Conservation Group
 Marine Conservation Society
 Mesoamerican Reef Tourism Initiative (MARTI)
[54]
 NSF Moorea Coral Reef Long-term Ecological Research site
 Nature Conservancy
 Ocean Voice International
 PADI
 Planetary Coral Reef Foundation[55]
[56]
 Practical Action
 Project Reefkeeper
 ReefBase
 Reef Check
  Reef Relief [57]
 Reefwatch
 Seacology
 SECORE
 Singapore Underwater Federation
 Society for Andaman and Nicobar Ecology
 Tubbataha Foundation
 Wildlife Conservation International
 WWF

Reefs in the past

Ancient coral reefs

Throughout Earth history, from a few thousand years after hard skeletons were developed by marine
organisms, there were almost always reefs. The times of maximum development were in the Middle
Cambrian (513–501 Ma), Devonian (416–359 Ma) and Carboniferous (359–299 Ma), due to
OrderRugosa extinct corals, and Late Cretaceous (100-65 Ma) and all Neogene (23 Ma–present), due to
Order Scleractinia corals.
Not all reefs in the past were formed by corals: in the Early Cambrian (542–513 Ma) resulted from
calcareous algae and archaeocyathids (small animals with conical shape, probably related to sponges) and in
the Late Cretaceous (100–65 Ma), when there also existed reefs formed by a group ofbivalves called rudists;
one of the valves formed the main conical structure and the other, much smaller valve acted as a cap.

See also

Ecology portal
 Census of Coral Reefs
 Marine biology
[edit]Notes
1. ^ Mulhall M (2007) Saving rainforests of the sea: An analysis of international efforts to conserve coral
reefs Duke Environmental Law and Policy Forum 19:321–351.
2. ^ "Corals reveal impact of land use". ARC Centre of Excellence for Coral Reef Studies. Retrieved 12
July 2007.
3. ^ Coral reef The Encyclopedia of Earth, Updated February 27, 2009.
4. ^ Darwin, Charles (1842). The Structure and Distribution of Coral Reefs. Being the first part of the
geology of the voyage of the Beagle, under the command of Capt. Fitzroy, R.N. during the years 1832
to 1836. London: Smith Elder and Co
a b
5. ^ Gordon Chancellor (2008). Introduction to Coral reefs. Darwin Online. Retrieved 2009-01-20
6. ^ Animation of coral atoll formation NOAA Ocean Education Service. Retrieved 9 January 2010.
7. ^ a b c Anderson, G (2003) Coral Reef Formation Marine Science.
8. ^ Great Barrier Reef Marine Park Authority (2006). "A "big picture" view of the Great Barrier
Reef" (PDF). Reef Facts for Tour Guides. Retrieved 18 June 2007.
9. ^ a b Tobin, Barry (1998, revised 2003). "How the Great Barrier Reef was formed". Australian Institute of
Marine Science. Retrieved 22 November 2006.
10. ^ CRC Reef Research Centre Ltd. "What is the Great Barrier Reef?". Retrieved 28 May 2006.
11. ^ Four Types of Coral Reef Microdocs, Stanford Education. Retrieved 10 January 2010.
12. ^ MSN Encarta (2006). "Great Barrier Reef". Great Barrier Reef. Retrieved 11 December 2006.
13. ^ Smithers, S.G. and Woodroffe, C.D. (August 2000). "Microatolls as sea-level indicators on a mid-
ocean atoll.". Marine Geology 168 (1–4): 61–78. doi:10.1016/S0025-3227(00)00043-8.
14. ^ a b c d Moyle and Cech, 2003, p. 556.
15. ^ a b c d Spalding, Mark, Corinna Ravilious, and Edmund Green. 2001. World Atlas of Coral Reefs.
Berkeley, CA: University of California Press and UNEP/WCMC.
16. ^ Achituv, Y. and Dubinsky, Z. 1990. Evolution and Zoogeography of Coral Reefs Ecosystems of the
World. Vol. 25:1-8.
17. ^ The Greenpeace Book of Coral Reefs
18. ^ a b Nybakken, James. 1997. Marine Biology: An Ecological Approach. 4th ed. Menlo Park, CA:
Addison Wesley.
19. ^ [1] Ultra Marine: In far eastern Indonesia, the Raja Ampat islands embrace a phenomenal coral
wilderness, by David Doubilet, National Geographic, September 2007
20. ^ Sherman, C.D.H. "The Importance of Fine-scale Environmental Heterogeneity in Determining Levels
of Genotypic Diversity and Local Adaption." University of Wollongong Ph.D. Thesis. 2006. Accessed
2009-06-07.
21. ^ Stacy, J., Marion, G., McCulloch, M. and Hoegh-Guldberg, O. "Long-term changes to Mackay
Whitsunday water quality and connectivity between terrestrial, mangrove and coral reef ecosystems:
Clues from coral proxies and remote sensing records - Synthesis of research from an ARC Linkage
Grant (2004-2007)." University of Queensland - Centre for Marine Studies. May 2007. Accessed 2009-
06-07.
22. ^ Nothdurft, L.D. "Microstructure and early diagensis of recent reef building scleractinian corals, Heron
Reef, Great Barrier Reef: Implications for palaeoclimate analysis." Queensland University of
Technology Ph.D. Thesis. 2007. Accessed 2009-06-07.
23. ^ Wilson, R.A. "The Biological Notion of Individual." Stanford Encyclopedia of Philosophy. August 9,
2007. Accessed 2009-06-07.
24. ^ Paul Marshall and Heidi Schuttenberg.; Marshall, Paul; Schuttenberg, Heidi. (2006). A Reef
Manager’s Guide to Coral Bleaching. Townsville, Australia: Great Barrier Reef Marine Park
Authority,. ISBN 1 876945 40 0.
25. ^ Rougerier, F The functioning of coral reefs and atolls: from paradox to paradigm ORSTOM, Papeete.
26. ^ Sorokin, Y. I. Coral Reef Ecology. Germany. Sringer-Herlag, Berlin Heidelberg. 1993.
27. ^ a b c Castro, Peter and Michael Huber. 2000. Marine Biology. 3rd ed. Boston: McGraw-Hill.
28. ^ http://oceanservice.noaa.gov/education/kits/corals/coral02_zooxanthellae.html
29. ^ A Reef Manager’s Guide to Coral Bleaching. Townsville, Australia: Great Barrier Reef Marine Park
Authority,. 2006. ISBN 1 876945 40 0.
30. ^ Rich Coral Reefs in Nutrient-Poor Water: Paradox Explained? National Geographic News, November
7, 2001.
31. ^ Corals play rough over Darwin's paradox New Scientist, 21 September 2002.
32. ^ Wilson E (2004) "Coral’s Symbiotic Bacteria Fluoresce, Fix Nitrogen" Chemical and engineering
news, 82(33): 7.
33. ^ Greenpeace Book of Coral Reefs
34. ^ Vroom, Peter S.; Page, Kimberly N.; Kenyon, Jean C.; Brainard, Russell E. (2006). "Algae-Dominated
Reefs". American Scientist 94 (5): 430–437.
35. ^ Coexistence of coral reef fishes—a lottery for living space PF Sale 1978 - Environmental Biology of
Fishes, 1978
36. ^ http://the.honoluluadvertiser.com/article/2005/Jan/17/ln/ln23p.html
37. ^ "U.S. Fish & Wildlife Service - Birds of Midway Atoll". Retrieved August 19, 2009.
38. ^ Osborne, Patrick L. (2000). Tropical Ecosystem and Ecological Concepts. Cambridge: Cambridge
University Press. pp. 464. ISBN 0 521 64523 9.
39. ^ Testimony of Dr. Lara J. Hansen before the Senate Committee on Commerce, Science and
Transportation, May 10, 2007.
a b
40. ^ The Importance of Coral to People WWF. Retrieved 12 January 2009.
a b
41. ^ Coral reefs around the world Guardian.co.uk, 2 September 2009.
42. ^ Space Daily etal. (2009). "Coral reefs tough it out against seaweed."science"".
43. ^ Kleypas, J.A., R.A. Feely, V.J. Fabry, C. Langdon, C.L. Sabine, and L.L. Robbins, 2006, Impacts of
Ocean Acidification on Coral Reefs and Other Marine Calcifiers: A guide for Future Research, NSF,
NOAA, & USGS, 88 pp.
44. ^ Save Our Seas, 1997 Summer Newsletter, Dr. Cindy Hunter and Dr. Alan Friedlander
45. ^ Tun, K., L.M. Chou, A. Cabanban, V.S. Tuan, Philreefs, T. Yeemin, Suharsono, K.Sour, and D. Lane,
2004, p:235-276 in C. Wilkinson (ed.), Status of Coral Reefs of the world: 2004.
 46. ^ Cinner, J. et al. (2005). Conservation and community benefits from traditional coral reef management
at Ahus Island, Papua New Guinea. Conservation Biology 19 (6), 1714-1723
47. ^ "Coral Reef Management, Papua New Guinea". Nasa's Earth Observatory. Retrieved 2 November
2006.
48. ^ a b Sabater, Marlowe G.; Yap, Helen T. 2004. "Long-term effects of induced mineral accretion on
growth, survival, and corallite properties of Porites cylindrica Dana." Journal of Experimental Marine
Biology and Ecology. Vol. 311:355-374.
49. ^ Superglue used for placement of coral
50. ^ Needle and thread use with soft coral
51. ^ 'The Coral Gardener'-documentary on coral gardening by Counterpart
52. ^ [2]
53. ^ Global Coral Reef Monitoring Network Status of Coral Reefs of the World 2008
54. ^ [3]
55. ^ [4]
56. ^ Practical Action coral reef restoration
57. ^ [5]

References

 Butler, Steven. 1996. "Rod? Reel? Dynamite? A tough-love aid program takes aim at the devastation of
the coral reefs". U.S. News and World Report, 25 November 1996.
 Christie, P. 2005a. University of Washington, Lecture. 18 May 2005.
 Christie, P. 2005b. University of Washington, Lecture. 4 May 2005.
 Clifton, Julian. 2003. Prospects for Co-Management in Indonesia's Marine Protected Areas. Marine
Policy, 27(5): 389-395.
 Courtney, Catherine and Alan White. 2000. Integrated Coastal Management in the Philippines. Coastal
Management; Taylor and Francis.
 Fox, Helen. 2005. Experimental Assessment of Coral Reef Rehabilitation Following Blast Fishing. The
Nature Conservancy Coastal and Marine Indonesia Program. Blackwell Publishers Ltd, February 2005.
 Gjertsen, Heidi. 2004. Can Habitat Protection Lead to Improvements in Human Well-Being? Evidence
from Marine Protected Areas in the Philippines.
 Moyle, PB and Cech, JJ (2003) Fishes, An Introduction to Ichthyology. 5th Ed, Benjamin
Cummings.ISBN 978-0131008472
 Sadovy, Y.J. Ecological Issues and the Trades in Live Reef Fishes, Part 1
 USEPA.
 UNEP. 2004. Coral Reefs in the South China Sea. UNEP/GEF/SCS Technical Publication No. 2.
 UNEP. 2007. Coral Reefs Demonstration Sites in the South China Sea. UNEP/GEF/SCS Technical
Publication No. 5.
  UNEP, 2007. National Reports on Coral Reefs in the Coastal Waters of the South China Sea.
UNEP/GEF/SCS Technical Publication No. 11.

External links

External images

Coral Reefs: Rainforests of the

SeaORG Educational films.

Wikimedia Commons
has media related
to: Coral reefs

Wikisource has the

text of the1911
Encyclopædia
Britannicaarticle Coral-
reefs.

 Coral Reefs- At the Smithsonian Ocean Portal

 How Coral Reefs Work
 International Year of the Reef in 2008
 Moorea Coral Reef Long Term Ecological Research Site (US NSF)
 ARC Centre of Excellence for Coral Reef Studies
 NOAA's Coral-List Listserver for Coral Reef Information and News
 NOAA's Coral Reef Conservation Program
 Exhibition of the Mexican Caribbean coral reef biodiversity aquarium in Xcaret Mexico
 NOAA's Coral Reef Information System
 ReefBase: A Global Information System on Coral Reefs
 National Coral Reef Institute Nova Southeastern University
 Marine Aquarium Council
 NCORE National Center for Coral Reef Research University of Miami
 Science and Management of Coral Reefs in the South China Sea and Gulf of Thailand
 NBII portal on coral reefs
 Microdocs: 4 kinds of Reef & Reef structure
 [6] Reef Relief Founders site: Coral reef resources, images, education, threats, solutions
Organisations
 Reef Fest Concert Series Non-profit effort to benefit coral reef conservation globally, starting in 2008, the
International Year of the Reef
 The Coral Reef Alliance (CORAL)
 Global Coral Reef Alliance
 Global Coral Reef Monitoring Network (GCRMN)
SCLERACTINIA

Brain Coral (Diploria labyrinthiformis)

Scientific classification

Kingdom: Animalia
Phylum: Cnidaria
Class: Anthozoa
Order: Scleractinia (Bourne, 1900)
Families

Suborder Astrocoeiina Suborder Suborder

 Acroporidae Caryophylliina Dendrophylliina
 Astrocoeniidae  Caryophylliidae  Dendrophylliidae
 Pocilloporidae

Suborder Faviina Suborder Oculina Suborder Fungiina

 Rhizangiidae  Pectiniidae  Agariciidae
 Faviidae  Trachyphyllidae  Fungiidae
 Meandrinidae  Poritidae
 Merulinidae  Siderastreidae
 Mussidae  Thamnasteriidae
 Oculinidae

Scleractinia, also called stony corals, are exclusively marine animals; they are very similar to sea
anemones but generate a hard skeleton. They first appeared in the Middle Triassic and
replaced tabulate and rugose corals that went extinct at the end of the Permian. Much of the framework of coral
reefs is formed by scleractinians.

There are two groups of Scleractinia:

 Colonial corals found in clear, shallow tropical waters; they are the world's primary reef-builders.
 Solitary corals are found in all regions of the oceans and do not build reefs. Some live
in temperate, polar waters, or below the photic zone down to 6000 meters.

Anatomy

Stony corals may be solitary or compound. Most have very small polyps, ranging from 1 to 3 millimetres
[1]
(0.04 to 0.1 in) in diameter, although some solitary species may be as large as 25 centimetres (9.8 in). The
most common forms include conical and horn-shaped polyps. Colonies can reach considerable size, consisting
of a large number of individual polyps.
 In a colonial Scleractinia, the repeated asexual division of the polyps causes the corallites to be
interconnected, thus forming the colonies. There are also cases in which the adjacent colonies of the same
species form a single colony by fusing. The living polyps are connected by horizontal sheets of tissue extending
over the outer surface of the skeleton and completely covering it. These sheets are outgrowths of the main body
of the polyp, and include extensions of the gastrovascular cavity, so that food and water can constantly circulate
between all the different members of the colony.[1]

Skeleton
The modern scleractinian skeleton, which lies external to the polyps that make it, is composed
of calcium carbonate in the form of aragonite. However, a prehistoric scleractinian (Coelosimilia) had a non-
[2]
aragonite calcium carbonate skeletal structure. The structure of both simple and compound scleractinians is
light and porous, rather than solid as in the Rugosa.

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 thin sheets of living tissue termed mesenteries.[1]
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 that is termed serial and that produces a
bilateral symmetry. Scleractinians are also distinguished from the Rugosa by their pattern of septal
insertion[vague]. They secrete a stony exoskeleton in which the septa are inserted between the mesenteries in
multiples of six.

In scleractinians, there are two main secondary structures:

 Stereome is an adherent layer of secondary tissue, which covers the septal surface. It consists of
transverse bundles of aragonitic needles and protects the polyps. However, its function can be nullified by
the thickening of the septa itself.
 Coenosteum is a perforated complex tissue that separates individual corallites in a compound
scleractinians.

At the beginning of Scleractinia’s development four groups with different microstructure can distinguished.
These are:
 Pachytecal: Corals having very thick walls and rudimentary septa. This is the group which probably
originated from Rugosa corals.
 Thick Trabecular: Corals with septa built from thick structures, resembling little beams, called trabecules.
 Minitrabecular: Corals with septa built from thin trabecules.
 Fascilcular or non-trabecular: Corals with septa not built from trabecules, but from columns composed of
bunches of aragonite fibres.

Ecology and life history

Hard coral Favites extends its polyps at night to feed

Scleractinians fall into one of two main categories:

 Zooxanthellate (hermatypic)
 Non-zooxanthellate (ahermatypic)

In hermatypic corals, the endodermal cells are replete with zooxanthellae symbiotic algae. These symbionts
benefit the corals because nearly 95% of the organic carbons produced by zooxanthellae are used as food by
the polyps. The oxygen byproduct of photosynthesis and additional energy derived from sugars produced by
zooxanthallae enable these corals to grow as much as three times faster as those without symbionts. These
corals are restricted to shallow (less than 200 feet - 60 meters), 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 feed on zooplankton, but those with larger polyps take correspondingly larger prey,
including various invertebrates and even small fish.[3] In addition to capturing prey with their tentacles, many
stony corals also produce mucus films that they can move over their bodies using cilia; these trap small organic
particles and are then pulled into the mouth. In a few stony corals, this is the primary method of feeding, and the
[1]
tentacles are reduced or absent.

Stony corals are generally nocturnal, with the polyps retreating into their skeletons during the day, but there
are a number of exceptions to this general rule.[1]

Non-zooxanthellate corals are usually non-reef formers and can be found most abundantly beneath about
500 m 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 calciferous and more susceptible to mechanical damage than that of zooxanthellate corals. 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 10 centimetres (3.9 in) a year (about the same
rate at which human hair grows). Other corals, like the dome and plate species are more bulky and may only
[4]
grow 0.3 to 2 centimetres (0.12 to 0.79 in) per year.

Life history

A Stony Coral

Stony corals can reproduce both sexually and asexually. Many

species have separate sexes, but others are hermaphroditic. Sexual
reproduction results in the birth of a free-swimming planula larva that
eventually settles to form a polyp. In colonial species, this initial polyp
then repeatedly divides asexually, to give rise to the entire colony. [1]

There are two main controls on the form of a scleractinian colony. One is the mode of budding and the
other is the relative growth rate. There are two types of budding: intratentacular and extratentacular. In
an intratentacular budding, polyps are divided by simple fission across the stomodaeum, and each bud retains
part of the original stomodaeum and regenerates the rest. Extratentacular budding takes place outside the
tentacular ring of the parent. These daughter buds do not share any part in the functions within the parent
scleractinians as do the products of intratentacular budding.

Evolutionary history

Scleractinian coral from the Matmor Formation (Jurassic) of Makhtesh

Gadol,Israel.

There are two main hypotheses about the origin of Scleractinia. The closest scleractinian analog in
the Paleozoic is the Rugosa, which suggests direct, possibly polyphyletic, descent, with different scleractinian
suborders having originated in different rugosan families. The second hypothesis suggests the similarities of
scleractinians to rugosans are due to a common non-skeletalized ancestor in the early Paleozoic. Recently
discovered Paleozoic corals with aragonitic skeletons and cyclic septal insertion - two features that characterize
Scleractinia - have strengthened the hypothesis for an independent origin of the Scleractinia.

Classification

The evolutionary relationships among stony corals were first examined in the 19th and early 20th
centuries. Early classification schemes used anatomical features of the polyps to propose evolutionary
relationships. The two most advanced 19th century classifications both used complex skeletal characters; Milne
Edwards and Haime’s 1857 classification was based on macroscopic skeletal characters, while Ogilvie’s 1897
scheme was developed using observations of skeletal microstructures, with particular attention to the structure
[5]
and pattern of the septal trabeculae .

Vaughan and Wells in 1943, and Wells 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[5]. Alloiteau’s 1952 classification built off of
these earlier studies but included more microstructural observations and did not involve the anatomical
characters of the polyp. Alloiteau recognized eight suborders[5]. Bryan and Hill, in 1942, 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[5]. Alloiteau later showed that established morphological classifications were unbalanced and that
the comparison of micro and macrostructural characters uncovered many convergences (convergent evolution)
between fossils and recent taxa.

The rise of molecular techniques at 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[5].
Through Romano and Palumbi’s 1996 analysis of mitochondrial RNA, it was found that molecular data
supported the assembling of species into the existing families (biology), 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.

Veron et al. analyzed ribosomal RNA in 1996 to obtain similar results to Romano and Palumbi, again
concluding that the traditional families were plausible but that the suborders were incorrect. Veron et al. also
established that stony corals are monophyletic, including all the descendants of a common ancestor, but that
[5]
they are divided into two groups, the robust and complex clades . He suggested that both morphological and
molecular systems be used in future classification schemes.
References

a b c d e f
1. ^ Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders
International. pp. 158–164. ISBN 0-03-056747-5.
2. ^ Stolarski, Jaroslaw; Anders Meibom, Radoslaw Przenioslo and Maciej Mazur (2007-10-05). "A
Cretaceous Scleractinian Coral with a Calcitic Skeleton". Science (USA: American Association for the
Advancement of Science) 318 (5847): 92–94. doi:10.1126/science.1149237. PMID 17916731.
Retrieved 2007-10-11.
3. ^ Witzany G, Madl P. (2009) Biocommunication of corals. International Journal of Integrative Biology
5(3): 152-163.
4. ^ Piper, Ross (2007), Extraordinary Animals: An Encyclopedia of Curious and Unusual
Animals, Greenwood Press.
5. ^ a b c d e f Stolarski, J., and Roniewicz, E., 2001, Towards a new synthesis of evolutionary relationships
and classification of Scleractinia: Journal of Paleontology, v. 75, p. 1090-1108
 Ezaki, Yoichi (1998). "Paleozoic Scleractinia: progenitors or extinct
experiments?". Palaeobiology (Paleontological Society) 24 (2): 227–234.

Alcyonacea

Soft coral
 Cladiella sp.

Scientific classification

Kingdom: Animalia

Phylum: Cnidaria

Class: Anthozoa

Subclass: Octocorallia

Order: Alcyonacea

Suborders

Alcyoniina
Calcaxonia
Holaxonia
Protoalcyonaria
Scleraxonia
Stolonifera