Difference between revisions of "Coral reef" - New World Encyclopedia

From New World Encyclopedia
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* '''[[Atoll reef]]''' – a more or less circular or continuous barrier reef extending all the way around a lagoon without a central island
 
* '''[[Atoll reef]]''' – a more or less circular or continuous barrier reef extending all the way around a lagoon without a central island
  
[[File:Maldives small island.jpg|thumb|right|A small [[atoll]] in the [[Maldives]].]]
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[[Image:Maldives small island.jpg|thumb|right|A small [[atoll]] in the [[Maldives]].]]
[[File:Maldives - Kurumba Island.jpg|thumb|right|Inhabited [[cay]] in the [[Maldives]]]]
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[[Image:Maldives - Kurumba Island.jpg|thumb|right|Inhabited [[cay]] in the [[Maldives]]]]
  
 
Other reef types or variants are:
 
Other reef types or variants are:
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<gallery>
 
<gallery>
File:Atoll forming-volcano.png|Darwin’s theory starts with a [[volcanic island]] which becomes extinct
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Image:Atoll forming-volcano.png|Darwin’s theory starts with a [[volcanic island]] which becomes extinct
File:Atoll forming-Fringing reef.png|As the island and ocean floor subside, coral growth builds a [[fringing reef]], often including a shallow lagoon between the land and the main reef
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Image:Atoll forming-Fringing reef.png|As the island and ocean floor subside, coral growth builds a [[fringing reef]], often including a shallow lagoon between the land and the main reef
File:Atoll forming-Barrier reef.png|As the subsidence continues the fringing reef becomes a larger barrier reef further from the shore with a bigger and deeper [[lagoon]] inside
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Image:Atoll forming-Barrier reef.png|As the subsidence continues the fringing reef becomes a larger barrier reef further from the shore with a bigger and deeper [[lagoon]] inside
File:Atoll forming-Atoll.png|Ultimately the island sinks below the sea, and the barrier reef becomes an [[atoll]] enclosing an open lagoon
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Image:Atoll forming-Atoll.png|Ultimately the island sinks below the sea, and the barrier reef becomes an [[atoll]] enclosing an open lagoon
 
</gallery>
 
</gallery>
  
 
Darwin predicted that underneath each lagoon would be a [[bed rock]] base, the remains of the original volcano. Subsequent drilling 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.
 
Darwin predicted that underneath each lagoon would be a [[bed rock]] base, the remains of the original volcano. Subsequent drilling 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.
  
[[File:Coral atoll formation animation.gif|thumb|right|A fringing reef can take ten thousand years to form, and an atoll can take up to 30 million years<ref>[http://oceanservice.noaa.gov/education/kits/corals/media/supp_coral04a.html Animation of coral atoll formation] [[NOAA]] Ocean Education Service. Retrieved 9 January 2010.</ref>]]
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[[Image:Coral atoll formation animation.gif|thumb|right|A fringing reef can take ten thousand years to form, and an atoll can take up to 30 million years<ref>[http://oceanservice.noaa.gov/education/kits/corals/media/supp_coral04a.html Animation of coral atoll formation] [[NOAA]] Ocean Education Service. Retrieved 9 January 2010.</ref>]]
  
 
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 as it is below its habitable depth.<ref name=cr/><ref name=Anderson2003 />
 
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 as it is below its habitable depth.<ref name=cr/><ref name=Anderson2003 />
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===Zones===
 
===Zones===
[[File:Coral reef diagram.jpg|thumb|300px|left|The three major zones of a coral reef: the fore reef, reef crest, and the back reef]]
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[[Image:Coral reef diagram.jpg|thumb|300px|left|The three major zones of a coral reef: the fore reef, reef crest, and the back reef]]
  
 
Coral reef ecosystems contain distinct zones that represent different kinds of habitats. Usually three major zones are recognized: the fore reef, reef crest, and the back reef (frequently referred to as the reef lagoon).
 
Coral reef ecosystems contain distinct zones that represent different kinds of habitats. Usually three major zones are recognized: the fore reef, reef crest, and the back reef (frequently referred to as the reef lagoon).
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Alternatively, Moyle and Cech distinguish six zones, though most reefs possess only some of the zones.<ref name=MoyleCech556>{{harvnb|Moyle|Cech|2003|p=556}}</ref>
 
Alternatively, Moyle and Cech distinguish six zones, though most reefs possess only some of the zones.<ref name=MoyleCech556>{{harvnb|Moyle|Cech|2003|p=556}}</ref>
  
[[File:Propagation du tsunami en profondeur variable.gif|thumb|right|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 it [[Wave shoaling|shoals]], that is, it slows down and the wave height increases.]]
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[[Image:Propagation du tsunami en profondeur variable.gif|thumb|right|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 it [[Wave shoaling|shoals]], 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 [[tidal surge|surge]] and the rise and fall of [[tide]]s. When waves pass over shallow areas, they [[Wave shoaling|shoal]], as shown in the diagram at the right. This means that the water is often agitated. These are the precise condition under which coral flourish. Shallowness means there is plenty of light for [[photosynthesis]] by the symbiotic [[zooxanthellae]], and agitated water promotes the ability of coral to feed on [[plankton]]. However other organisms must be able to withstand the robust conditions to flourish in this zone.
 
* '''The reef surface''' is the shallowest part of the reef. It is subject to the [[tidal surge|surge]] and the rise and fall of [[tide]]s. When waves pass over shallow areas, they [[Wave shoaling|shoal]], as shown in the diagram at the right. This means that the water is often agitated. These are the precise condition under which coral flourish. Shallowness means there is plenty of light for [[photosynthesis]] by the symbiotic [[zooxanthellae]], and agitated water promotes the ability of coral to feed on [[plankton]]. However other organisms must be able to withstand the robust conditions to flourish in this zone.
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==Distribution==
 
==Distribution==
[[File:Coral reef locations.jpg|thumb|300px|Locations of coral reefs.]]
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[[Image:Coral reef locations.jpg|thumb|300px|Locations of coral reefs.]]
[[File:20 Grad Isotherme.png|thumb|300px|Boundary for 20 °C [[Contour line|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.]]
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[[Image:20 Grad Isotherme.png|thumb|300px|Boundary for 20 °C [[Contour line|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.]]
[[File:Upwelling image1.jpg|thumb|right|300px|This map shows areas of [[upwelling]] in red. Coral reefs are not found in coastal areas where colder and nutrient-rich upwellings occur]]
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[[Image:Upwelling image1.jpg|thumb|right|300px|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 {{convert|284300|km2|sp=us}},<ref>[[UNEP]] (2001) [http://coral.unep.ch/atlaspr.htm UNEP-WCMC World Atlas of Coral Reefs] Coral Reef Unit</ref> which is just under one tenth of one percent of the oceans' surface area. 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 account for 7.6%.<ref name="Spalding">Spalding, Mark, Corinna Ravilious, and Edmund Green. 2001. ''World Atlas of Coral Reefs''. Berkeley, CA: University of California Press and UNEP/WCMC.</ref>
 
Coral reefs are estimated to cover {{convert|284300|km2|sp=us}},<ref>[[UNEP]] (2001) [http://coral.unep.ch/atlaspr.htm UNEP-WCMC World Atlas of Coral Reefs] Coral Reef Unit</ref> which is just under one tenth of one percent of the oceans' surface area. 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 account for 7.6%.<ref name="Spalding">Spalding, Mark, Corinna Ravilious, and Edmund Green. 2001. ''World Atlas of Coral Reefs''. Berkeley, CA: University of California Press and UNEP/WCMC.</ref>
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==Coral biology==
 
==Coral biology==
[[File:Coral polyp.jpg|thumb|240px|right|Anatomy of a coral polyp.]]
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[[Image:Coral polyp.jpg|thumb|240px|right|Anatomy of a coral polyp.]]
 
{{See also|Coral}}
 
{{See also|Coral}}
  
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The colonies of the one thousand coral [[species]] assume a characteristic shape such as [[Brain coral|wrinkled brains]], cabbages, [[Acropora|table tops]], [[Staghorn coral|antlers]], wire strands and [[Pillar coral|pillars]].{{citation needed|date=December 2010}}
 
The colonies of the one thousand coral [[species]] assume a characteristic shape such as [[Brain coral|wrinkled brains]], cabbages, [[Acropora|table tops]], [[Staghorn coral|antlers]], wire strands and [[Pillar coral|pillars]].{{citation needed|date=December 2010}}
  
[[File:FFS Table bottom.jpg|thumb|180px|left|[[Acropora|Table coral]]]]
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[[Image:FFS Table bottom.jpg|thumb|180px|left|[[Acropora|Table coral]]]]
[[File:Coral detail.jpg|thumb|180px|right|Close up of polyps arrayed on a coral, waving their tentacles. There can be thousands of polyps on a single coral branch.]]
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[[Image:Coral detail.jpg|thumb|180px|right|Close up of polyps arrayed on a coral, waving their tentacles. There can be thousands of polyps on a single coral branch.]]
  
 
===Reproduction===
 
===Reproduction===
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<gallery>
 
<gallery>
File:Brain coral.jpg|[[Brain coral]]
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Image:Brain coral.jpg|[[Brain coral]]
File:Staghorn-coral-1.jpg|[[Staghorn coral]]
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Image:Staghorn-coral-1.jpg|[[Staghorn coral]]
File:Cirripathes sp (Spiral Wire Coral).jpg|Spiral wire coral
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Image:Cirripathes sp (Spiral Wire Coral).jpg|Spiral wire coral
File:PillarCoral.jpg|[[Pillar coral]]
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Image:PillarCoral.jpg|[[Pillar coral]]
 
</gallery>
 
</gallery>
  
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Plants form the base of the food chain, and need sunlight and nutrients to grow. In the ocean these plants are mainly microscopic [[phytoplankton]] which drift in the [[water column]]. They need sunlight for [[photosynthesis]], which powers [[carbon fixation]], so they are found only relatively near the surface. But they also need nutrients. Phytoplankton rapidly use nutrients in the surface waters, and in the tropics these nutrients are not usually replaced because of the [[thermocline]].<ref name=Anderson2003 />
 
Plants form the base of the food chain, and need sunlight and nutrients to grow. In the ocean these plants are mainly microscopic [[phytoplankton]] which drift in the [[water column]]. They need sunlight for [[photosynthesis]], which powers [[carbon fixation]], so they are found only relatively near the surface. But they also need nutrients. Phytoplankton rapidly use nutrients in the surface waters, and in the tropics these nutrients are not usually replaced because of the [[thermocline]].<ref name=Anderson2003 />
  
[[File:Polyps (PSF).png|thumb|left|Coral polyps]]
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[[Image:Polyps (PSF).png|thumb|left|Coral polyps]]
  
 
====Solution: retention and recycling====
 
====Solution: retention and recycling====
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   | isbn = 1 876945 40 0  }}</ref> In return, as an example of [[Mutualism (biology)|mutualism]], the coral shelter the zooxanthellae, averaging one million for every cubic centimetre of coral, with and provide a constant supply of the [[carbon dioxide]] it needs for photosynthesis.
 
   | isbn = 1 876945 40 0  }}</ref> In return, as an example of [[Mutualism (biology)|mutualism]], the coral shelter the zooxanthellae, averaging one million for every cubic centimetre of coral, with and provide a constant supply of the [[carbon dioxide]] it needs for photosynthesis.
  
[[File:Multy color corals.JPG|thumb|right|The colour of corals depends on the type of [[zooxanthella]] they host]]
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[[Image:Multy color corals.JPG|thumb|right|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 catch [[zooplankton]] that brush them when the water is agitated. Zooplankton provide the polyp with nitrogen, and the polyp shares some of the nitrogen with the zooxanthellae, which also require this element.<ref name="Castro">Castro, Peter and Michael Huber. 2000. ''Marine Biology.'' 3rd ed. Boston: McGraw-Hill.</ref> The varying pigments in different species of zooxanthellae give corals their different colours. Coral which loses its zooxanthellae becomes white and is said to be [[Coral bleaching|bleached]], a condition which unless corrected can kill the coral.
 
Corals also absorb nutrients, including inorganic nitrogen and phosphorus, directly from the water. Many corals extend their tentacles at night to catch [[zooplankton]] that brush them when the water is agitated. Zooplankton provide the polyp with nitrogen, and the polyp shares some of the nitrogen with the zooxanthellae, which also require this element.<ref name="Castro">Castro, Peter and Michael Huber. 2000. ''Marine Biology.'' 3rd ed. Boston: McGraw-Hill.</ref> The varying pigments in different species of zooxanthellae give corals their different colours. Coral which loses its zooxanthellae becomes white and is said to be [[Coral bleaching|bleached]], a condition which unless corrected can kill the coral.
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Sponges are another key to explaining Darwin’s paradox. They live in crevices in the coral reefs. They are efficient [[filter feeder]]s, and in the [[Red Sea]] they consume about sixty percent of the phytoplankton that drifts by. The sponges eventually excrete nutrients in a form the corals can use.<ref>[http://news.nationalgeographic.com/news/2001/11/1107_keyholecoral.html Rich Coral Reefs in Nutrient-Poor Water: Paradox Explained?] ''[[National Geographic News]]'', November 7, 2001.</ref>
 
Sponges are another key to explaining Darwin’s paradox. They live in crevices in the coral reefs. They are efficient [[filter feeder]]s, and in the [[Red Sea]] they consume about sixty percent of the phytoplankton that drifts by. The sponges eventually excrete nutrients in a form the corals can use.<ref>[http://news.nationalgeographic.com/news/2001/11/1107_keyholecoral.html Rich Coral Reefs in Nutrient-Poor Water: Paradox Explained?] ''[[National Geographic News]]'', November 7, 2001.</ref>
  
[[File:Sea Cotton.jpg|thumb|left|Most coral polyps are nocturnal feeders. Here, in the dark, polyps have extended their tentacles to feed on zooplankton]]
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[[Image:Sea Cotton.jpg|thumb|left|Most coral polyps are nocturnal feeders. Here, in the dark, polyps have extended their tentacles to feed on zooplankton]]
  
 
The roughness of coral surfaces is the key to coral survival in agitated waters. Normally a boundary layer of still water surrounds a submerged object, which acts as a barrier. Waves breaking on the extremely rough edges of corals disrupt the boundary layer, allowing the corals access to nutrients. Turbulent water thereby promotes rapid reef growth and lots of branching. Without the nutritional gains brought by rough coral surfaces, even the most effective recycling would leave corals wanting in nutrients.<ref>[http://www.newscientist.com/article/mg17523612.100-corals-play-rough-over-darwins-paradox.html Corals play rough over Darwin's paradox] ''New Scientist'', 21 September 2002.</ref>
 
The roughness of coral surfaces is the key to coral survival in agitated waters. Normally a boundary layer of still water surrounds a submerged object, which acts as a barrier. Waves breaking on the extremely rough edges of corals disrupt the boundary layer, allowing the corals access to nutrients. Turbulent water thereby promotes rapid reef growth and lots of branching. Without the nutritional gains brought by rough coral surfaces, even the most effective recycling would leave corals wanting in nutrients.<ref>[http://www.newscientist.com/article/mg17523612.100-corals-play-rough-over-darwins-paradox.html Corals play rough over Darwin's paradox] ''New Scientist'', 21 September 2002.</ref>
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===Biodiversity===
 
===Biodiversity===
[[File:Callyspongia sp. (Tube sponge).jpg|thumb|300px|right|Tube sponges attracting [[cardinal fish]]es, [[Asiatic glassfish|glassfish]]es and [[wrasse]]s]]
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[[Image:Callyspongia sp. (Tube sponge).jpg|thumb|300px|right|Tube sponges attracting [[cardinal fish]]es, [[Asiatic glassfish|glassfish]]es and [[wrasse]]s]]
  
 
Reefs are home to a large variety of organisms, including fish, [[seabird]]s, [[sea sponge|sponges]], [[Cnidaria]]ns (which includes some types of corals and [[jellyfish]]), [[worm]]s, [[crustacean]]s (including [[shrimp]], [[cleaner shrimp]], [[spiny lobster]]s and [[crab]]s), [[mollusc]]s (including [[cephalopod]]s), [[echinoderm]]s (including [[starfish]], sea urchins and [[Holothuroidea|sea cucumber]]s), [[sea squirt]]s, [[sea turtle]]s and [[sea snake]]s. Aside from humans, [[mammal]]s are rare on coral reefs, with visiting [[cetacean]]s such as [[dolphin]]s being the main exception. A few of these varied species feed directly on corals, while others graze on algae on the reef.<ref name="Spalding" /><ref name="Castro" />
 
Reefs are home to a large variety of organisms, including fish, [[seabird]]s, [[sea sponge|sponges]], [[Cnidaria]]ns (which includes some types of corals and [[jellyfish]]), [[worm]]s, [[crustacean]]s (including [[shrimp]], [[cleaner shrimp]], [[spiny lobster]]s and [[crab]]s), [[mollusc]]s (including [[cephalopod]]s), [[echinoderm]]s (including [[starfish]], sea urchins and [[Holothuroidea|sea cucumber]]s), [[sea squirt]]s, [[sea turtle]]s and [[sea snake]]s. Aside from humans, [[mammal]]s are rare on coral reefs, with visiting [[cetacean]]s such as [[dolphin]]s being the main exception. A few of these varied species feed directly on corals, while others graze on algae on the reef.<ref name="Spalding" /><ref name="Castro" />
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*Fish that cruise the boundaries of the reef or nearby seagrass meadows. These include predatory fish such as [[pompano]]s, [[grouper]]s, [[Horse mackerel]]s, certain types of shark, ''[[Epinephelus marginatus]]'', [[barracuda]]s and [[Lutjanidae|snappers]]).  Herbivorous and plankton-eating fish also populate reefs.  Seagrass-eating fish include Horse mackerel, snapper, Pagellus and Conodon.  Plankton-eating fish include Caesio, manta ray, chromis, Holocentridae and  pterapogon kauderni.
 
*Fish that cruise the boundaries of the reef or nearby seagrass meadows. These include predatory fish such as [[pompano]]s, [[grouper]]s, [[Horse mackerel]]s, certain types of shark, ''[[Epinephelus marginatus]]'', [[barracuda]]s and [[Lutjanidae|snappers]]).  Herbivorous and plankton-eating fish also populate reefs.  Seagrass-eating fish include Horse mackerel, snapper, Pagellus and Conodon.  Plankton-eating fish include Caesio, manta ray, chromis, Holocentridae and  pterapogon kauderni.
  
[[File:Coral reef PloS.jpg|thumb|right|Organisms can cover every square inch of a coral reef.]]
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[[Image:Coral reef PloS.jpg|thumb|right|Organisms can cover every square inch of a coral reef.]]
  
 
Fish that swim in coral reefs can be as colorful as the reef. Examples are the parrotfish, [[Pomacanthidae|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]]'' and ''[[Pterapogon kauderni]]'') as well as aquatic animals (Comatulida, [[Crinoidea]] and [[Ophiuroidea]]) emerge and become active while others rest.
 
Fish that swim in coral reefs can be as colorful as the reef. Examples are the parrotfish, [[Pomacanthidae|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]]'' and ''[[Pterapogon kauderni]]'') as well as aquatic animals (Comatulida, [[Crinoidea]] and [[Ophiuroidea]]) emerge and become active while others rest.
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<gallery>
 
<gallery>
File:Prionurus laticlavius.jpg|[[Shoaling and schooling|Schooling]] [[reef fish]]
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Image:Prionurus laticlavius.jpg|[[Shoaling and schooling|Schooling]] [[reef fish]]
File:Caribbean reef squid.jpg|[[Caribbean reef squid]]
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Image:Caribbean reef squid.jpg|[[Caribbean reef squid]]
File:Stenopus hispidus.jpg|[[Banded coral shrimp]]
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Image:Stenopus hispidus.jpg|[[Banded coral shrimp]]
File:Triaenodon obesus moc.jpg|The [[whitetip reef shark]] almost exclusively inhabits coral reefs
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Image:Triaenodon obesus moc.jpg|The [[whitetip reef shark]] almost exclusively inhabits coral reefs
File:Green turtle John Pennekamp.jpg|[[Green turtle]]
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Image:Green turtle John Pennekamp.jpg|[[Green turtle]]
File:Red sea-reef 3627.jpg|[[Giant clam]]
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Image:Red sea-reef 3627.jpg|[[Giant clam]]
File:Nephtheidae komodo.jpg|Soft coral, cup coral, sponges and ascidians
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Image:Nephtheidae komodo.jpg|Soft coral, cup coral, sponges and ascidians
File:Laticauda colubrina (Wakatobi).jpg|[[Banded sea krait]]
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Image:Laticauda colubrina (Wakatobi).jpg|[[Banded sea krait]]
 
</gallery>
 
</gallery>
  
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==Threats==
 
==Threats==
[[File:Mvey0290.jpg|thumb|right|Island with [[fringing reef]] off [[Yap]], [[Micronesia]]. Coral reefs are dying around the world.<ref name="Coral reefs around the world">[http://www.guardian.co.uk/environment/interactive/2009/sep/02/coral-world-interactive Coral reefs around the world] ''[[Guardian.co.uk]]'', 2 September 2009.</ref>]]
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[[Image:Mvey0290.jpg|thumb|right|Island with [[fringing reef]] off [[Yap]], [[Micronesia]]. Coral reefs are dying around the world.<ref name="Coral reefs around the world">[http://www.guardian.co.uk/environment/interactive/2009/sep/02/coral-world-interactive Coral reefs around the world] ''[[Guardian.co.uk]]'', 2 September 2009.</ref>]]
 
{{Main|Environmental issues with coral reefs}}
 
{{Main|Environmental issues with coral reefs}}
  
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==Protection==
 
==Protection==
 
{{Main|Coral reef protection}}
 
{{Main|Coral reef protection}}
[[File:Coral Outcrop Flynn Reef.jpg|thumb|250px|A diversity of corals]]
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[[Image:Coral Outcrop Flynn Reef.jpg|thumb|250px|A diversity of corals]]
  
 
[[Marine Protected Area|Marine Protected Areas (MPAs)]] have become increasingly prominent for reef management. MPAs in Southeast Asia and elsewhere around the world attempt to promote responsible [[Fisheries management|fishery management]] and [[habitat (ecology)|habitat protection]]. Much like [[national park]]s and wildlife 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.{{citation needed|date=December 2010}}
 
[[Marine Protected Area|Marine Protected Areas (MPAs)]] have become increasingly prominent for reef management. MPAs in Southeast Asia and elsewhere around the world attempt to promote responsible [[Fisheries management|fishery management]] and [[habitat (ecology)|habitat protection]]. Much like [[national park]]s and wildlife 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.{{citation needed|date=December 2010}}
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== Origins ==
 
== Origins ==
[[File:Ancient coral reefs.jpg|thumb|260px||left|Ancient coral reefs]]
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[[Image:Ancient coral reefs.jpg|thumb|260px||left|Ancient coral reefs]]
  
 
Beginning a few thousand years after hard skeletons were developed by marine organisms, coral reefs emerged. The times of maximum development were in the [[Middle Cambrian]] (513–501 [[Annum|Ma]]), [[Devonian]] (416–359 Ma) and [[Carboniferous]] (359–299 Ma), due to [[Order (biology)|Order]] [[Rugosa]] [[extinction|extinct]] corals, and [[Late Cretaceous]] (100–65 Ma) and all [[Neogene]] (23 Ma–present), due to Order [[Scleractinia]] corals.
 
Beginning a few thousand years after hard skeletons were developed by marine organisms, coral reefs emerged. The times of maximum development were in the [[Middle Cambrian]] (513–501 [[Annum|Ma]]), [[Devonian]] (416–359 Ma) and [[Carboniferous]] (359–299 Ma), due to [[Order (biology)|Order]] [[Rugosa]] [[extinction|extinct]] corals, and [[Late Cretaceous]] (100–65 Ma) and all [[Neogene]] (23 Ma–present), due to Order [[Scleractinia]] corals.

Revision as of 00:36, 4 March 2011

Ocean habitats
Some of the biodiversity of a coral reef, in this case the Great Barrier Reef

Some of the biodiversity of a coral reef, in this case the Great Barrier Reef

Littoral zone
Intertidal zone
Estuaries
Kelp forests
Coral reefs
Ocean banks
Continental shelf
Neritic zone
Straits
Pelagic zone
Oceanic zone
Seamounts
Hydrothermal vents
Cold seeps
Demersal zone
Benthic zone

Coral reefs are underwater structures made from calcium carbonate secreted by corals. Corals are colonies of tiny living animals found in marine waters that contain 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.Also coral reef is a reef consisting of coral consolidated into limestone

Often called “rainforests of the sea”, coral reefs form some of the most diverse ecosystems on Earth. They occupy less than one tenth of 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,[1] including fish, molluscs, worms, crustaceans, echinoderms, sponges, tunicates and other cnidarians.[2] 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, but deep water and cold water corals also exist on smaller scales in other areas.

Coral reefs deliver ecosystem services to tourism, fisheries and shoreline protection. The annual global economic value of coral reefs has been estimated at $US375 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, including urban and agricultural runoff and water pollution, which can harm reefs by encouraging excess algae growth.[3][4][5]

Reef structure

Types

The three principal reef types 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 channel or 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, possibly winding reef, usually associated with an atoll lagoon
  • Table reef – an isolated reef, approaching an atoll type, but without a lagoon.
  • Habili - reef in the Red Sea that does not reach the surface near enough to cause visible surf, although it may a hazard to ships (from the Arabic for "unborn")
  • Microatolls – certain species of corals form communities called microatolls. The vertical growth of microatolls is limited by average tidal height. By analysing growth morphologies, microatolls offer 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.[6]
  • 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 to become habitable by humans. Cays occur in tropical environments throughout the Pacific, Atlantic and Indian Oceans (including 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.
  • When a coral reef cannot keep up with the sinking of a volcanic island, a seamount or guyot is formed. The tops of seamounts and guyots are below the surface. 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.

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.[7] 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 in his first monograph, The Structure and Distribution of Coral Reefs[8] Charles Darwin 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.[9] 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.

Darwin predicted that underneath each lagoon would be a bed rock base, the remains of the original volcano. Subsequent drilling 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[10]

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 as it is below its habitable depth.[9][11]

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. Sea level was then 120 meters (390 ft) lower than it is today.[12][13] As sea level rose, the water and the corals encroached on what had been hills of the Australian coastal plain. By 13,000 years ago sea level had risen to 60 meters (200 ft) lower than at present, and the hills of the coastal plains were, by then, continental islands. As the sea level rise continued, water topped most of the continental islands. The corals could then overgrow the hills, forming the present cays and reefs. Sea level on the Great Barrier Reef has not changed significantly in the last 6,000 years,[13] and the age of the present living reef structure is estimated to be between 6,000 and 8,000 years.[14] Although the Great Barrier Reef formed along a continental shelf, and not around a volcanic island, Darwin's principles apply. The Great Barrier Reef development stopped at the barrier reef stage, since Australia is not about to submerge. It formed the world's largest barrier reef, {{safesubst:#invoke:convert|convert}} from shore, stretching for 2,000 kilometers (1,200 mi).[15]

Healthy coral reefs grow horizontally from 1 to 3 centimeters (0.39 to 1.2 in) per year, and grow vertically anywhere from 1 to 25 centimeters (0.39 to 9.8 in) per year; however, they grow only at depths shallower than 150 meters (490 ft) due to their need for sunlight, and cannot grow above sea level.[16]

Zones

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

Coral reef ecosystems contain distinct zones that represent different kinds of habitats. Usually three major zones are recognized: the fore reef, reef crest, and the back reef (frequently referred to as the reef lagoon).

All three zones are physically and ecologically interconnected. Reef life and oceanic processes create opportunities for exchange of seawater, sediments, nutrients, and marine life among one another.

Thus, they are integrated components of the coral reef ecosystem, each playing a role in the support of the reefs' abundant and diverse fish assemblages.

Most coral reefs exist in shallow waters less than fifty metres deep. Some inhabit 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 the Maldives. The reefs surrounding islands form when islands subside into the ocean, and atolls form when an island subsides below the surface of the sea.

Alternatively, Moyle and Cech distinguish six zones, though most reefs possess only some of the zones.[17]

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 it shoals, 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 surge and the rise and fall of tides. When waves pass over shallow areas, they shoal, as shown in the diagram at the right. This means that the water is often agitated. These are the precise condition under which coral flourish. Shallowness means there is plenty of light for photosynthesis by the symbiotic zooxanthellae, and agitated water promotes the ability of coral to feed on plankton. However other organisms 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 occurs by reefs on continental shelves. Reefs around tropical islands and atolls drop abruptly to great depths, and don't have a floor. Usually sandy, the 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. Its complex growths of coral and calcareous algae provide cracks and crevices for protection, and the abundant invertebrates and epiphytic algae provide an ample source of food."[17]
  • 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."[17]
  • The reef lagoon – "many coral reefs completely enclose an area, thereby creating a quiet-water lagoon that usually contains small patches of reef."[17]

However, the "topography of coral reefs is constantly changing. Each reef is made up of irregular patches of algae, sessile invertebrates, and bare rock and sand. The size, shape and relative abundance of these patches 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."[18]

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),[19] which is just under one tenth of one percent of the oceans' surface area. 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 account for 7.6%.[20]

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 {{safesubst:#invoke:convert|convert}}, and few reefs exist in waters below 18 °C (64 °F).[21] However reefs in the Persian Gulf have adapted to temperatures of 13 °C (55 °F) in winter and 38 °C (100 °F) in summer.[22]

Deep water coral 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).[23] Corals are seldom found along the coastline of South Asia from the eastern tip of India (Madras) to the border of Bangladesh and Myanmar.[20] 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

  • The Great Barrier Reef - largest, comprising over 2,900 individual reefs and 900 islands stretching for over 2,600 kilometers (1,615.6 mi) off Queensland, Australia
  • The Mesoamerican Barrier Reef System - second largest, stretching 1,000 kilometers (621.4 mi) from Isla Contoy at the tip of the Yucatán Peninsula down to the Bay Islands of Honduras
  • The New Caledonia Barrier Reef - second longest double barrier reef, covering 1,500 kilometers (930 mi)
  • The Andros, Bahamas Barrier Reef - third largest, following the east coast of Andros Island, Bahamas, between Andros and Nassau
  • The Red Sea - includes 6000 year old fringing reefs located around a 2,000 km (1,240 mi) coastline.
  • 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.[24]

Coral biology

Anatomy of a coral polyp.
See also: Coral

Live coral are small animals embedded in calcium carbonate shells. It is a mistake to think of coral as plants or rocks. Coral heads consist of accumulations of individual animals called polyps, arranged in diverse shapes.[25] Polyps are usually tiny, but they can range in size from a pinhead to a foot across. Reef-building or hermatypic corals live only in the photic zone (above 50 m depth), the depth to which sufficient sunlight penetrates the water for photosynthesis to occur. Coral polyps do not themselves photosynthesize, but have a symbiotic relationship with single-celled organisms called zooxanthellae; these organisms live within the tissues of polyps and provide organic nutrients that nourish the polyp. 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 significant reef structures. Corals get up to 90% of their nutrients from their zooxanthellae symbionts.[26]

Reefs grow as polyps and other organisms deposit calcium carbonate,[27][28] the basis of coral, as a skeletal structure beneath and around themselves, pushing the coral head's top upwards and outwards.[29] 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.

The colonies of the one thousand coral species assume a characteristic shape such as wrinkled brains, cabbages, table tops, antlers, wire strands and pillars.[citation needed]

Table coral
Close up of polyps arrayed on a coral, waving their tentacles. There can be thousands of polyps on a single coral branch.

Reproduction

Corals reproduce both sexually and asexually. Individual polyp uses both reproductive modes within its lifetime. Corals reproduce sexually by either internal or external fertilization. The reproductive cells are found on the 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 species 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 en masse, simultaneously. Eggs disperse over a large area. The timing of spawning depends on time of year, water temperature, and tidal and lunar cycles. Spawning is most successful 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 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. They are vulnerable to predation and environmental conditions. The lucky few who attach to substrate next confront competition for food and space.[citation needed]

Ecology

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 the oligotrophy 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 from ? — Francis Rougerie[30]
—{{{source}}}

The nutrient paradox

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 surrounded by tropical ocean waters that provide hardly any nutrients.

Coral reefs cover less than one tenth of 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. Coral reefs' primary productivity is very high, typically producting 5-10g C m−2 day−1 biomass.[31]

One reason for the startling clarity of tropical waters is that they are deficient in nutrients and drifting plankton. Further, the sun shines year round in the tropics, warming the surface layer, making it less dense than subsurface layers. The warmer water is separated from 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. In most parts of the ocean there is little exchange between these layers. Organisms that die in aquatic environments generally sink to the bottom where they decompose. 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.[11]

Plants form the base of the food chain, and need sunlight and nutrients to grow. In the ocean these plants are mainly microscopic phytoplankton which drift in the water column. They need sunlight for photosynthesis, which powers carbon fixation, so they are found only relatively near the surface. But they also need nutrients. Phytoplankton rapidly use nutrients in the surface waters, and in the tropics these nutrients are not usually replaced because of the thermocline.[11]

Coral polyps

Solution: retention and recycling

Around coral reefs, lagoons fill in with material eroded from the reef and the island. They become havens for marine life, providing protection from waves and storms.

Most importantly, reefs recycle nutrients, which happens much less in the open ocean. In coral reefs and lagoons, producers include phytoplankton as well as seaweed and coralline algae, especially small types called turf algae, which pass nutrients to corals.[32] 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.

Coral reefs support many symbiotic relationships. In particular, zooxanthellae provides energy to coral in the form of glucose, glycerol, and amino acids.[33] Zooxanthellae can provide up to 90% of a coral’s energy requirements.[34] In return, as an example of mutualism, the coral shelter the zooxanthellae, averaging one million for every cubic centimetre of coral, with and provide 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 catch zooplankton that brush them when the water is agitated. Zooplankton provide the polyp with nitrogen, and the polyp shares some of the nitrogen with the zooxanthellae, which also require this element.[32] The varying 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 kill the coral.

Sponges are another key to explaining Darwin’s paradox. They 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 sponges eventually excrete nutrients in a form the corals can use.[35]

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

The roughness of coral surfaces is the key to coral survival in agitated waters. Normally a boundary layer of still water surrounds a submerged object, which acts as a barrier. Waves breaking on the extremely rough edges of corals disrupt the boundary layer, allowing the corals access to nutrients. Turbulent water thereby promotes rapid reef growth and lots of branching. Without the nutritional gains brought by rough coral surfaces, even the most effective recycling would leave corals wanting in nutrients.[36]

Cyanobacteria provide soluble nitrates for the reef via nitrogen fixation.[37]

Coral reefs also often depend on surrounding habitats, such as seagrass meadows and mangrove forests, for 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.[38]

Biodiversity

Tube sponges attracting cardinal fishes, glassfishes and wrasses

Reefs are home to a large variety of 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.[20][32]

Fish

Coral reefs are home to a wide variety of tropical or reef fish, among them are the following:

  • Fish that influence 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 on organisms that inhabit larger 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 (such as sea urchins), and they remove inedible seaweeds. Fish that eat coral itself include parrotfish and butterflyfish.
  • 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 and snappers). Herbivorous and plankton-eating fish also populate reefs. Seagrass-eating fish include Horse mackerel, snapper, Pagellus and Conodon. Plankton-eating fish include Caesio, manta ray, chromis, Holocentridae and pterapogon kauderni.
Organisms can cover every square inch of a coral reef.

Fish that swim in coral reefs can be as colorful as the reef. Examples are the 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 and Pterapogon kauderni) as well as aquatic animals (Comatulida, Crinoidea and 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 coral reefs.[20] It has been suggested that the fish species that inhabit coral reefs are able to coexist in such high numbers because the reef is "full" in that 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".[39] Healthy reefs can produce up to 35 tons of fish per square kilometer each year, but damaged reefs produce much less.[40]

Invertebrates

Sea urchins, Dotidae and sea slugs eat seaweed. Some species of sea urchins, such as Diadema antillarum, can play a pivotal part in preventing algae overrunning reefs.[41] 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 mollusks, and sipunculans. Those settling on the reef include many other species, particularly crustaceans and polychaete worms.[23]

Algae

Researchers have found evidence of algae dominance in locations of healthy coral reefs. In surveys done around largely uninhabited US Pacific islands, algae inhabit a large percentage of surveyed coral locations.[42] The algae population consists of turf algae, coralline algae, and macroalgae.

Seabirds

Coral reef systems provide important habitats for seabird species, some endangered. For example, Midway Atoll in Hawaii 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.[43] Each seabird species has specific sites on the atoll where they nest. Altogether, 17 species of seabirds live on Midway. The short-tailed albatross is the rarest, with fewer than 2,200 surviving after excessive feather hunting in the late nineteenth century.[44]

Other

Sea snakes feed exclusively on fish and their eggs. Tropical birds such as herons, gannets, pelicans and boobies feed on reef fish. Some land based reptiles intermittently associate with reefs, such as monitor lizards, the marine crocodile and semi-aquatic snakes like Laticauda colubrina. Sea turtles eat sponges.[citation needed]

Economic value

Coral reefs deliver ecosystem services to tourism, fisheries and coastline protection. The global economic value of coral reefs has been estimated at as much as $US375 billion per year.[45] 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.[46] 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.[46]

To improve the management of coastal coral reefs the WRI (World Resource Institute) has been trying to retrieve reliable information on the actual value of services possible due to the existence of coral reefs. Currently they are working partners in five Caribbean countries and have come up with a detailed estimate of the value of coral reef related tourism, shoreline protection and fisheries. The WRI is also making sure that the study results actually support the improved coastal policies and management planning recently set up them. They have completed economic valuation in St. Lucia, Tobago, Belize, and the Dominican Republic and are now conducting an economic valuation in Jamaica. They have also created a excel-based program called the “Economic Valuation Tool” that helps guide users through step by step instructions of getting data, checking assumptions and coming up with appropriate results.[47]

Threats

Island with fringing reef off Yap, Micronesia. Coral reefs are dying around the world.[48]


Coral reefs are dying around the world.[48] In particular, coral mining, agricultural and urban runoff, pollution (organic and non-organic), overfishing, blast fishing, disease, and the digging of canals and access into islands and bays are localized threats to coral ecosystems. Broader threats are sea temperature rise, sea level rise and pH changes from ocean acidification, all associated with greenhouse gas emissions.[citation needed]

In El Nino-year 2010, preliminary reports show global coral bleaching reached its worst level since another El Nino year, 1998, when 16 percent of the world's reefs died as a result of excessive water temperature. In Indonesia's Aceh province, surveys showed some 80 percent of bleached corals died. In July, Malaysia closed several dive sites after virtually all the corals In some areas were damaged by bleaching.[49][50]

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 storms carrying agents to far flung reefs, pollutants, algal blooms and others. Reefs are threatened well beyond coastal areas.[citation needed]

General estimates show approximately 10% world's coral reefs are already dead.[51][52][53] 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.[citation needed]

Protection

A diversity of corals

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 wildlife 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.[citation needed]

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 Belize's Barrier reef, Chagos archipelago, Sian Ka'an, the Galapagos islands, Great Barrier Reef, Henderson Island, Palau and Papahānaumokuākea Marine National Monument have been designated as world heritage sites.

In Australia, the Great Barrier Reef is protected by the Great Barrier Reef Marine Park Authority, and is the subject of much legislation, including a Biodiversity Action Plan.

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.[54][55]

Organizations

Organizations which currently undertake coral reef/atoll restoration projects using simple methods of plant propagation:


Organizations which promote interest, provide knowledge bases about coral reef survival, and promote activities to protect and restore coral reefs:

  • Australian Coral Reef Society
  • Biosphere Foundation[57]
  • 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[58]
  • Great Barrier Reef Marine Park Authority
  • Green Fins
  • ICRAN Mesoamerican Reef Alliance
  • International Coral Reef Initiative (ICRI)[59]
  • 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)
  • NSF Moorea Coral Reef Long-term Ecological Research site[60]
  • Nature Conservancy
  • Ocean Voice International
  • PADI
  • Planetary Coral Reef Foundation[61]
  • Practical Action[62]
  • Project Reefkeeper
  • ReefBase
  • Reef Check
  • Reef Relief[63]
  • Reefwatch
  • Save Our Seas Foundation[64]
  • Seacology
  • SECORE
  • Singapore Underwater Federation
  • Society for Andaman and Nicobar Ecology
  • Tubbataha Foundation
  • Wildlife Conservation International
  • WWF


Artificial reefs

Efforts to expand the size and number of coral reefs generally involve supplying substrate to allow more corals to find a home. Substrate materials include discarded vehicle tires, scuttled ships, subway cars, and formed concrete such as reef balls. Reefs also grow unaided on marine structures such as oil rigs.

In large restoration projects, propagated hermatypic coral on substrate can be secured with metal pins, superglue or milliput.[65] Needle and thread can also attach A-hermatype coral to substrate.[66]

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 competitive filamentous and fleshy algae. The increased growth rates fully depend on the accretion activity.[67]

During 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.[67]

Origins

Ancient coral reefs

Beginning a few thousand years after hard skeletons were developed by marine organisms, coral reefs emerged. The times of maximum development were in the Middle Cambrian (513–501 Ma), Devonian (416–359 Ma) and Carboniferous (359–299 Ma), due to Order Rugosa 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: Early Cambrian (542–513 Ma) reefs resulted from calcareous algae and archaeocyathids (small animals with conical shape, probably related to sponges) and rudists, a type of bivalve, built Late Cretaceous (100–65 Ma) reefs.

See also

Template:Portal box

  • Census of Coral Reefs
  • Marine biology

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. Hoover, John (November, 2007). Hawaiʻi's Sea Creatures. Mutual. Retrieved December, 2010. 
  3. Corals reveal impact of land use. ARC Centre of Excellence for Coral Reef Studies. Retrieved 12 July 2007.
  4. Minato, Charissa (July 1, 2002). Urban runoff and coastal water quality being researched for effects on coral reefs. Retrieved December, 2010.
  5. Coastal Watershed Factsheets - Coral Reefs and Your Coastal Watershed. Environmental Protection Agency Office of Water (July 1998). Retrieved December, 2010.
  6. 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.
  7. Coral reef The Encyclopedia of Earth, Updated February 27, 2009.
  8. Template:Cite document
  9. 9.0 9.1 Template:Cite document
  10. Animation of coral atoll formation NOAA Ocean Education Service. Retrieved 9 January 2010.
  11. 11.0 11.1 11.2 Anderson, G (2003) Coral Reef Formation Marine Science.
  12. 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.
  13. 13.0 13.1 Tobin, Barry (1998, revised 2003). How the Great Barrier Reef was formed. Australian Institute of Marine Science. Retrieved 22 November 2006.
  14. CRC Reef Research Centre Ltd. What is the Great Barrier Reef?. Retrieved 28 May 2006.
  15. Four Types of Coral Reef Microdocs, Stanford Education. Retrieved 10 January 2010.
  16. MSN Encarta. (2006). "Great Barrier Reef".. Retrieved on 11 December 2006.
  17. 17.0 17.1 17.2 17.3 Moyle & Cech 2003, p. 556
  18. Connell, Joseph H. (March 24, 1978). Diversity in Tropical Rain Forests and Coral Reefs. Science 199 (4335 pp).
  19. UNEP (2001) UNEP-WCMC World Atlas of Coral Reefs Coral Reef Unit
  20. 20.0 20.1 20.2 20.3 Spalding, Mark, Corinna Ravilious, and Edmund Green. 2001. World Atlas of Coral Reefs. Berkeley, CA: University of California Press and UNEP/WCMC.
  21. Achituv, Y. and Dubinsky, Z. 1990. Evolution and Zoogeography of Coral Reefs Ecosystems of the World. Vol. 25:1-8.
  22. The Greenpeace Book of Coral Reefs
  23. 23.0 23.1 Nybakken, James. 1997. Marine Biology: An Ecological Approach. 4th ed. Menlo Park, CA: Addison Wesley.
  24. [1] Ultra Marine: In far eastern Indonesia, the Raja Ampat islands embrace a phenomenal coral wilderness, by David Doubilet, National Geographic, September 2007
  25. 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.
  26. Paul Marshall and Heidi Schuttenberg. and 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. 
  27. Stacy, J., Marion, G., McCulloch, M. and Hoegh-Guldberg, O. "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.
  28. 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.
  29. Wilson, R.A. "The Biological Notion of Individual."Stanford Encyclopedia of Philosophy. August 9, 2007. Accessed 2009-06-07.
  30. Rougerier, F The functioning of coral reefs and atolls: from paradox to paradigm ORSTOM, Papeete.
  31. Sorokin, Y. I. Coral Reef Ecology. Germany. Sringer-Herlag, Berlin Heidelberg. 1993.
  32. 32.0 32.1 32.2 Castro, Peter and Michael Huber. 2000. Marine Biology. 3rd ed. Boston: McGraw-Hill.
  33. Zooxanthellae… What's That?
  34. (2006) A Reef Manager’s Guide to Coral Bleaching. Townsville, Australia: Great Barrier Reef Marine Park Authority,. ISBN 1 876945 40 0. 
  35. Rich Coral Reefs in Nutrient-Poor Water: Paradox Explained? National Geographic News, November 7, 2001.
  36. Corals play rough over Darwin's paradox New Scientist, 21 September 2002.
  37. Wilson E (2004) "Coral’s Symbiotic Bacteria Fluoresce, Fix Nitrogen" Chemical and engineering news, 82(33): 7.
  38. Greenpeace Book of Coral Reefs
  39. Coexistence of coral reef fishes—a lottery for living space PF Sale 1978 - Environmental Biology of Fishes, 1978
  40. Coral degradation through destructive fishing practices. Encyclopedia of Earth (2008).
  41. Osborne, Patrick L. (2000). Tropical Ecosystem and Ecological Concepts. Cambridge: Cambridge University Press, 464. ISBN 0 521 64523 9. 
  42. (2006). Algae-Dominated Reefs. American Scientist 94 (5): 430–437..
  43. http://the.honoluluadvertiser.com/article/2005/Jan/17/ln/ln23p.html
  44. U.S. Fish & Wildlife Service - Birds of Midway Atoll. Retrieved August 19, 2009.
  45. Heat Stress to Caribbean Corals in 2005 Worst on Record National Oceanic and Atmospheric Administration, NOAA. 15 November 2010.
  46. 46.0 46.1 The Importance of Coral to People WWF. Retrieved 12 January 2009.
  47. "Coastal Capital: Economic Valuation of Coastal Ecosystems in the Caribbean." World Resources Institute. World Resources Institute, n.d. Web. 21 Jan 2011. <http://www.wri.org/project/valuation-caribbean-reefs>.
  48. 48.0 48.1 Coral reefs around the world Guardian.co.uk, 2 September 2009.
  49. RITTER, KARL (December 8, 2010). goal-coral-reefs.html Climate goal may spell end for some coral reefs. Associated Press. Retrieved December, 2010.
  50. Error on call to template:cite web: Parameters url and title must be specifiedMarkey, Sean (May 16, 2006). . National Geographic News. Retrieved December, 2010.
  51. 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.
  52. Save Our Seas, 1997 Summer Newsletter, Dr. Cindy Hunter and Dr. Alan Friedlander
  53. 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.
  54. 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
  55. Coral Reef Management, Papua New Guinea. Nasa's Earth Observatory. Retrieved 2 November 2006.
  56. 'The Coral Gardener'-documentary about Counterpart scientist Austin Bowden-Kerby
  57. Biosphere Foundation
  58. Global Coral Reef Monitoring Network Status of Coral Reefs of the World 2008
  59. International Coral Reef Initiative
  60. NSF Moorea Coral Reef Long-term Ecological Research site
  61. Planetary Coral Reef Foundation
  62. Action coral reef restoration Practical Action
  63. Reef Relief
  64. Save Our Seas Foundation
  65. Superglue used for placement of coral
  66. Needle and thread use with soft coral
  67. 67.0 67.1 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.

References
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