Black coral

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Black coral
Blackcoral colony 600.jpg
Black coral colony
CITES Appendix II (CITES)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Cnidaria
Class: Hexacorallia
Order: Antipatharia
Milne-Edwards & Haime, 1857
Families [1]

Antipatharians, also known as black corals or thorn corals, [2] are an order of soft deep-water corals. These corals can be recognized by their jet-black or dark brown chitin skeletons, which are surrounded by their colored polyps (part of coral that is alive). Antipatharians are a cosmopolitan order, existing in nearly every oceanic location and depth, with the sole exception of brackish waters. However, they are most frequently found on continental slopes under 50 m (164 ft) deep. A black coral reproduces both sexually and asexually throughout its lifetime. Many black corals provide housing, shelter, food, and protection for other animals.

Contents

Black corals were originally classified in the subclass Ceriantipatharia along with ceriantharians (tube-dwelling anemones), but were later reclassified under Hexacorallia. Though they have historically been used by Pacific Islanders for medical treatment and in rituals, its only modern use is making jewelry. Black corals have been declining in numbers and are expected to continue declining due to the effects of poaching, ocean acidification and climate change.

Etymology

Despite its name, a black coral is rarely black, and depending on the species can be white, red, green, yellow, or brown. The corals derive their name from their black skeletons, which are composed of protein and chitin. [3] Black corals are also known as thorn corals due to the microscopic spines lining their skeletons. [4]

The name Antipatharia comes from the Ancient Greek word antipathes ("against disease"). In the Hawaiian language, a black coral is called ʻēkaha kū moana ("hard bush growing in the sea"); it is the official state gem of Hawaii. [5] In Malay, the corals are called akah bahar ("root of the sea"), likely named for their tendency to grow at low-light depths. [6]

Taxonomy and classification

Black corals have historically been difficult to classify due to poor-quality specimens. They have few distinguishing morphological characteristics, and the few that there are vary across species, similar to other corals. When black corals were first documented by Henri Milne-Edwards and Jules Haime, two French zoologists in 1857, all species of Antipatharia were placed in the family Antipathidae. [4] From 2001 to 2006, marine biologists Dennis Opresko and Tina Molodtsova helped transform the taxonomic system into what it is today. [1] A 2007 phylogenetic study confirmed the new taxonomic system. [7]

Blacks corals are classified in the order Antipatharia with 7 families, 44 genera, and 280 distinct species. [2] The families are Antipathidae, Aphanipathidae, Cladopathidae, Leiopathidae, Myriopathidae, Schizopathidae, and Stylopathidae. [8] Black corals can be distinguished from other corals by their black, flexible skeletons and near-total lack of any kind of protection from sediment. All black corals have small or medium-sized polyps and a chitin skeleton, lined with small spines. [9]

Genera

List of genera according to the World Register of Marine Species: [10]

Physical characteristics

A sample of coral next to its skeleton, showing the minuscule spines all along it Stylopathes adinocrada (10.3897-zoologia.36.e28714) Figures 8-10.jpg
A sample of coral next to its skeleton, showing the minuscule spines all along it

The skeletons of these corals grow in many patterns unique to this order, such as whips, trees, fans, or coils. These range in size from 10 to 300 cm (3.94 to 118 in), though polyps can be as small as 1 mm (0.0394 in) in size. [4] [11] Skeletons are also lined with tiny spikes. [4] These spikes are roughly 0.5 mm (0.0197 in) in size, and vary widely in terms of size, length, proportion, and sharpness. [2] A layer of "bark" forms around the skeleton as the coral grows. The polyps that live inside this bark are less than 2 mm (0.0787 in) [12] and are gelatinous and have six tentacles (the same as hard corals and unlike soft corals, which have eight). [13] These polyps can be nearly any color. [3] Some corals also have "sweeper tentacles", which can grow up to 15 mm (0.591 in) long. [12] Though individual polyps are either male or female, entire colonies are typically hermaphroditic. [14]

Unlike the vast majority of other corals, black corals have no protection against abrasive materials such as sand and rocks and lack muscular development which can help the corals to hide. These factors can lead to sediment tearing the soft tissue, resulting in death. In response, corals live near crevices, which allows much of their body to be protected. [9]

Ecology

Habitat

A Cirrhipathes coral, sometimes known as wire coral Cirrhipathes sp - coral-wire-groovy.jpg
A Cirrhipathes coral, sometimes known as wire coral

Black corals occur throughout all the oceans from the surface down to the deep-sea, though nearly 75% of species are only found at depths below 50 m (164 ft). The sole oceanic area in which black corals have not been found are brackish waters, though they can inhabit areas with decreased salinity. [15] Black corals are found on reefs, and may contribute to overall reef building, but are also often found as solitary colonies on isolated outcroppings. Most individuals require a hard surface for attachment. They will frequently grow where undersea currents flow, which allows them to feed on the meiofauna that is swept by. Since undersea currents benefit the corals, they will often grow on or by geographic structures that cause currents, such as continental slopes, cliffs, caves, or undersea plateaus. [9] Species distributions of black corals are poorly understood, and while many deep sea black corals have large distributions, more recent work has indicated that shallow black coral species—such as Antipathes grandis—can be found spanning from the Indian to the Pacific Ocean. [16]

Diet

Black corals are carnivorous, with the coral's polyps allowing it to feed mostly on meiofauna such as zooplankton. [17] [4] The polyps of cnidarians have an oral disk in their center which serves as the mouth for the coral. The disk is surrounded by the tentacles, which stings and digests food. [9] The reason many corals are fan-shaped is to catch meiofauna. Many corals only have polyps on the downstream side of the coral, [17] allowing them to catch nearly the same number of animals without wasting energy keeping unnecessary polyps alive. [17]

A princess parrotfish, one of the few predators of black coral Princess-parrotfish.png
A princess parrotfish, one of the few predators of black coral

Predators

Vertebrate predation is not a major threat to black corals. [15] There are rare reports of parrotfish and butterflyfish gnawing and eating at the polyps of black corals, but even if a polyp is gnawed off, it will not affect the coral as a whole. The skeleton of a black coral is hard and inert, due to its composition of protein and chitin, making it nearly inedible. Though black coral skeletons have been found in the stomachs of green sea turtles and sharks, these incidents are rare; it has thus been suggested that black corals are not a major part of any vertebrate diets. [9]

However, invertebrates such as muricids and ovulids [9] feed on black corals and similar corals regularly. These mollusks mimic the polyps that the coral typically feeds on and is taken inside of the coral. They will then consume the polyps from the inside out. [9] Various mollusks, such as Coralliophila kaofitorum and Phenacovolva carneptica live solely where various species of black corals are found, suggesting that they prey exclusively on the species. [15]

Interactions

Black corals around the world provide a unique environment for crustaceans, bivalves, and fish. Some species, such as Dascyllus albisella and Centropyge potteri inhabit specific coral trees. Due to this abundance of species, nighttime predation around the coral beds has been observed. [18] [19] [20]

Life cycle and reproduction

Due to the slow life cycle and deep-water habitats of black coral, little is known about their life cycle and reproduction. [8] As with other cnidarians, the life cycle of these corals involves both asexual and sexual reproduction. Asexual reproduction (also known as budding), is the first method of reproduction used by a black coral during their lifespan. [17] Once a polyp is anchored, it builds a colony by creating a skeleton, growing new branches and making it thicker, similar to the growth of a tree. This method of growing creates "growth rings" which can be used to estimate the age of a colony. [21] Asexual reproduction can also occur if a branch breaks off and a replacement is needed. [17] Though light is not required for growth or development, mature colonies will grow towards light. Why they do so is unknown. [22]

Antipathes dendrochristos growing several hundred meters down in the ocean Antipathes dendrochristos.jpg
Antipathes dendrochristos growing several hundred meters down in the ocean

Sexual reproduction in these corals remains largely unstudied. It occurs after the coral colony is established; a colony will produce eggs and sperm, which meet in the water to create larvae that use currents to disperse and settle in new areas. [17] The larval stage of the coral, called a planula, will drift along until it finds a surface on which it can grow. Once it settles, it metamorphoses into its polyp form and creates skeletal material to attach itself to the seafloor. It will then begin to bud, which will create new polyps and eventually form a colony. [17] In areas with ideal conditions, black coral colonies can grow to be extremely dense, creating beds. [9] In some black corals that have been closely examined, colonies will grow roughly 6.4 cm (2.52 in) every year. Sexual reproduction occurs after 10 to 12 years of growth; the colony will then reproduce annually for the rest of its life. The male to female polyp ratio is 1:1, with females producing anywhere from 1.2 million to 16.9 million oocytes. [23] A large 1.8 m (5.91 ft) tall coral tree is somewhere between 30 and 40 years old. [17]

The estimated natural lifespan of a black coral colony in the epipelagic zone is 70 years. However, in March 2009 around 4,265 years old specimens of Leiopathes glaberrima were found at depths of nearly 300 to 3,000 m (984 to 9,840 ft), making them some of the oldest living organisms on earth. The researchers showed that the "individual colony longevities are on the order of thousands of years." [24] [25] Rarely, black corals will grow too large to support their own weight, and collapse. [9]

Human use and harvesting

Black coral bracelet Image of black coral bracelet.jpg
Black coral bracelet

Black corals have historically been associated with mystical and medicinal properties in Indonesian, Chinese, and Hawaiian culture. [9] [26] More recent harvesting has been for use as jewelry. [26] [27] Many Indo-Pacific peoples believed that black coral has curative and anti-evil powers and made them into necklace and bracelets; however, black corals are not ideal for jewelry-making due to it being soft as opposed to stony, [4] causing jewelry made with it to dry out and break. [4] If a real black coral is boiled in milk, it will smell of myrrh; this test can be used to determine if a sample is genuine. [28]

The best studied and regulated black coral fisheries are in Hawaii, where they have been harvested since the 1960s. [26] [29] In the Caribbean harvesting is typically done to produce jewelry for sale to tourists, and has followed a boom-and-bust cycle, where new coral populations are discovered and overexploited leading to rapid declines. [26] For example, Cozumel, Mexico, was famed for dense black coral beds that have been harvested since the 1960s [30] leading to widespread black coral population declines. [31] Despite improvements in management in Cozumel, including no harvesting permits issued since the mid-1990s, the black coral population had failed to recover when assessed in 2016. [32] Though it is illegal to move black corals across international borders without authorization, as they are listed in Appendix II of the Convention on International Trade in Endangered Species (CITES), it is still possible to buy them. [33]

Though various methods have been proposed for quicker and more efficient rejuvenation of black coral colonies, none have worked to the point where these corals could be commercially farmed. [34]

Threats

A Bathypathes species growing in a brush shape Bathypathes y cangrejos.jpg
A Bathypathes species growing in a brush shape

Though black corals are not listed on the IUCN Red List, a number of factors threaten them today. The largest single threat is poaching— though the majority of black coral fisheries are heavily regulated, there is still a black market for the corals. [35] Particularly on tropical islands and Madagascar, the market for illegally-harvested black corals is large. [35] [36] Due to overfishing of mature corals, in some areas nearly 90% of corals are juveniles (less than 50 cm (19.7 in) tall.) [37]

Global warming is the primary threat to black corals worldwide, as well as all other corals. [38] Though black corals rarely builds reefs (the most threatened areas), threats caused by climate change such as coral bleaching, rising sea temperatures, changing underwater currents, and changing salinity and pH also affect deep-sea corals. [39] Invasive species such as Carijoa riisei , which were introduced to Hawaiian waters by humans, may pose a significant threat to black corals. [36]

Related Research Articles

<span class="mw-page-title-main">Cnidaria</span> Aquatic animal phylum having cnydocytes

Cnidaria is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, including jellyfish, hydroids, sea anemone, corals and some of the smallest marine parasites. Their distinguishing feature is the cnidocytes, specialized cells with ejectable flagella used mainly for envenomation and capturing prey. Their bodies consist of mesoglea, a non-living jelly-like substance, sandwiched between two layers of epithelium that are mostly one cell thick.

<span class="mw-page-title-main">Coral</span> Marine invertebrates of the class Anthozoa

Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.

<span class="mw-page-title-main">Anthozoa</span> Class of cnidarians without a medusa stage

Anthozoa is a class of marine invertebrates which includes the sea anemones, stony corals and soft corals. Adult anthozoans are almost all attached to the seabed, while their larvae can disperse as part of the plankton. The basic unit of the adult is the polyp; this consists of a cylindrical column topped by a disc with a central mouth surrounded by tentacles. Sea anemones are mostly solitary, but the majority of corals are colonial, being formed by the budding of new polyps from an original, founding individual. Colonies are strengthened by calcium carbonate and other materials and take various massive, plate-like, bushy or leafy forms.

<span class="mw-page-title-main">Zoantharia</span> Order of hexacorallians with marginal tentacles

Zoanthids are an order of cnidarians commonly found in coral reefs, the deep sea and many other marine environments around the world. These animals come in a variety of different colonizing formations and in numerous different colors. They can be found as individual polyps, attached by a fleshy stolon or a mat that can be created from small pieces of sediment, sand and rock. The term "zoanthid" refers to all animals within this order Zoantharia, and should not be confused with "Zoanthus", which is one genus within Zoantharia.

<span class="mw-page-title-main">Alcyonacea</span> Order of octocorals that do not produce massive calcium carbonate skeletons

Alcyonacea are a species of sessile colonial cnidarians that are found throughout the oceans of the world, especially in the deep sea, polar waters, tropics and subtropics. Whilst not in a strict taxonomic sense, Alcyonacea are commonly known as "soft corals" (Octocorallia) that are quite different from "true" corals (Scleractinia). The term “soft coral” generally applies to organisms in the two orders Pennatulacea and Alcyonacea with their polyps embedded within a fleshy mass of coenenchymal tissue. Consequently, the term “gorgonian coral” is commonly handed to multiple species in the order Alcyonacea that produce a mineralized skeletal axis composed of calcite and the proteinaceous material gorgonin only and corresponds to only one of several families within the formally accepted taxon Gorgoniidae (Scleractinia). These can be found in order Malacalcyonacea (taxonomic synonyms of include : Alcyoniina, Holaxonia, Protoalcyonaria, Scleraxonia, and Stolonifera. They are sessile colonial cnidarians that are found throughout the oceans of the world, especially in the deep sea, polar waters, tropics and subtropics. Common names for subsets of this order are sea fans and sea whips; others are similar to the sea pens of related order Pennatulacea. Individual tiny polyps form colonies that are normally erect, flattened, branching, and reminiscent of a fan. Others may be whiplike, bushy, or even encrusting. A colony can be several feet high and across, but only a few inches thick. They may be brightly coloured, often purple, red, or yellow. Photosynthetic gorgonians can be successfully kept in captive aquaria.

<span class="mw-page-title-main">Hexacorallia</span> Class of cnidarians with 6-fold symmetry

Hexacorallia is a class of Anthozoa comprising approximately 4,300 species of aquatic organisms formed of polyps, generally with 6-fold symmetry. It includes all of the stony corals, most of which are colonial and reef-forming, as well as all sea anemones, and zoanthids, arranged within five extant orders. The hexacorallia are distinguished from another class of Anthozoa, Octocorallia, in having six or fewer axes of symmetry in their body structure; the tentacles are simple and unbranched and normally number more than eight. These organisms are formed of individual soft polyps which in some species live in colonies and can secrete a calcite skeleton. As with all Cnidarians, these organisms have a complex life cycle including a motile planktonic phase and a later characteristic sessile phase. Hexacorallia also include the significant extinct order of rugose corals.

<span class="mw-page-title-main">Sea anemone</span> Marine animals of the order Actiniaria

Sea anemones are a group of predatory marine invertebrates constituting the order Actiniaria. Because of their colourful appearance, they are named after the Anemone, a terrestrial flowering plant. Sea anemones are classified in the phylum Cnidaria, class Anthozoa, subclass Hexacorallia. As cnidarians, sea anemones are related to corals, jellyfish, tube-dwelling anemones, and Hydra. Unlike jellyfish, sea anemones do not have a medusa stage in their life cycle.

<span class="mw-page-title-main">Antipathidae</span> Family of corals

Antipathidae is a family of corals in the order Antipatharia, commonly known as black corals. They are generally considered a deep-water taxon; however, some of the most diverse communities are known from tropical shallow waters.

Allopathes is a genus of corals in the family Antipathidae. It is characterized by several long stems protruding from a short, thick base with spines arranged vertically around the stem. Its polyps are arranged in a single row that run the length of the coral. This genus was initially a subgenus of Cirrhipathes and Stichopathes, although it also displayed similarities to Antipathes verticillata. However, the presence of branched growth forms excludes it from Stichopathes or Cirrhipathes and the unique morphology of its spines meant that it could not be included in Allopathes. Because they seemed to have a combination of characteristics of different genera, the two species in question, Allopathes desbonni and Allopathes robillardi were given their own genus.

<i>Antipathes</i> Genus of corals

Antipathes is a genus of coral in the order Antipatharia, composed of black coral. Distinct features vary greatly within this genus: it contains symmetrically aligned as well as irregularly shaped corals, a range of different colors, and colonies that can be either sparsely branched or closely packed. polyps for these corals have six tentacles that are each lined with stinging cells. Unlike their reef-building cousins, these coral lack photosynthesizing algae and are not restricted to the lighter surface regions. They prefer to live in deeper waters near currents so they can catch and eat passing zooplankton.

<i>Aphanipathes</i> Genus of corals

Aphanipathes is a diverse genus of black corals in the family Aphanipathidae, typified by large polypar spines. However, there are some disagreement in the correct taxonomic classification of this genus. The Global Biodiversity Information Facility (GBIF) classifies Aphanipathes as being a genus of the family Aphanipathidae while the Integrated Taxonomic Information System (ITIS) classifies it as a genus of the family Antipathidae.

<i>Antipathes dichotoma</i> Species of coral

Antipathes dichotoma is a species of colonial coral in the order Antipatharia, the black corals, so named because their calcareous skeletons are black. It was first described by the German zoologist and botanist Peter Simon Pallas in 1766, from a single specimen he received from near Marseilles in the Mediterranean Sea.

<i>Cirrhipathes</i> Genus of corals

Cirrhipathes is a genus of black coral from the family Antipathidae. Coral species in this genus are commonly known as whip or wire corals because they often exhibit a twisted or coiled morphology. In addition to their colorful appearance, with colors ranging from yellow to red passing through blue and green, these species possess a dark skeleton that is characteristic to every black coral.

<i>Savalia savaglia</i> Species of coral

Savalia savaglia, commonly known as gold coral, is a species of colonial false black coral in the family Parazoanthidae. It is native to the northeastern Atlantic Ocean and the Mediterranean Sea where it often grows in association with a gorgonian. It is extremely long-lived, with a lifespan of 2,700 years, and develops into a large tree-like colony.

<i>Porites cylindrica</i> Species of coral

Porites cylindrica, commonly known as hump coral, is a stony coral belonging to the subclass Hexacorallia in the class Anthozoa. Hexacorallia differ from other subclasses in that they have 6 or fewer axes of symmetry. Members of this class possess colonial polyps which can be reef-building, secreting a calcium carbonate skeleton. They are dominant in both inshore reefs and midshelf reefs.

<i>Plumapathes pennacea</i> Species of coral

Plumapathes pennacea is a species of black coral in the order Antipatharia. It is found in the tropical Indian, Pacific and Atlantic Oceans in deep reef habitats where it forms part of a biologically diverse community.

<i>Leiopathes glaberrima</i> Species of cnidarian

Leiopathes glaberrima is a species of black coral of the order Antipatharia found in the northern Atlantic Ocean and the Mediterranean Seas deep water habitats. A very slow-growing species, it is among the oldest living animals on the planet.

<i>Antipathella fiordensis</i> Species of coral

Antipathella fiordensis is a species of colonial coral in the order Antipatharia, the black corals, so named because their calcareous skeletons are black. It was first described as Antipathes fiordensis by the New Zealand zoologist Ken R. Grange in 1990, from material collected in the steep-sided fiords of Fiordland in the southeastern South Island, New Zealand. A 2001 revision of the Antipatheria put this species in the newly-created genus Antipathella.

Bathypathes is a genus of black coral in the family Schizopathidae.

Alternipathes bipinnata is a species of coral belonging to the family Schizopathidae. It has been found off the Pacific coast of Mexico.

References

  1. 1 2 Tina Molodtsova, Dennis Opresko (2020). "Antipatharia". WoRMS. World Register of Marine Species . Retrieved 13 May 2020.
  2. 1 2 3 Opresko, Dennis. "Spotlight on Antipatharians (Black Corals)". NMNH. Retrieved 4 May 2020.
  3. 1 2 Bo, Marzia (21 April 2012). "Isolation and identification of chitin in the black coral Parantipathes larix (Anthozoa: Cnidaria)". International Journal of Biological Macromolecules. 51 (1–2): 129–137. doi:10.1016/j.ijbiomac.2012.04.016. hdl: 11567/802206 . PMID   22546360.
  4. 1 2 3 4 5 6 7 "Spotlight on antipatharians". NMNH. 18 April 2016. Retrieved 4 September 2019.
  5. Grigg, Richard W. (1993). "Precious Coral Fisheries of Hawaii and the U.S. Pacific Islands" (PDF). Marine Fisheries Review. 55 (2): 54. Retrieved 29 September 2010.
  6. Skeat, Walter William (1906). Pagan Races of the Malay Peninsula: pt. 3. Religion. pt. 4. Language. Appendix. Comparative vocabulary of aboriginal dialects. Index of subjects. Index of proper names. Index of native words. Macmillen and company. ISBN   1149951974.
  7. Brugler, Mercer, R.; France, Scott C. (March 2007). "The complete mitochondrial genome of the black coral Chrysopathes formosa (Cnidaria:Anthozoa:Antipatharia) supports classification of antipatharians within the subclass Hexacorallia". Molecular Phylogenetics and Evolution. 42 (3): 776–778. doi:10.1016/j.ympev.2006.08.016. PMID   17049278 . Retrieved 4 May 2020.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  8. 1 2 NOAA. "Black Corals of Hawaii". oceanexplorer.noaa.gov.
  9. 1 2 3 4 5 6 7 8 9 10 Wagner, Daniel (December 2011). The biology and ecology of Hawaiian black corals (Cnidaria : Anthozoa: Hexacorallia: Antipatharia) (PhD). University of Hawaii at Manoa.
  10. Dennis Opresko (2019). "Antipatharia". WoRMS. World Register of Marine Species . Retrieved 25 November 2019.
  11. "Black Coral: Hawaii State Gem". State Symbols USA. 21 September 2014. Retrieved 13 September 2019.
  12. 1 2 Goldberg, W. M.; Grange, K. R.; Zuniga, A. L. (August 1990). "The Structure of Sweeper Tentacles in the Black Coral Antipathes fiordensis". The Biological Bulletin. 179 (1): 96–104. doi:10.2307/1541743. JSTOR   1541743. PMID   29314907.
  13. Milne-Edwards and Haine. "Antipatharia sp (Milne-Edwards & Haime, 1857): "Black Coral"". EdwardsLabs. Archived from the original on 30 October 2019. Retrieved 30 October 2019.
  14. Bo, Marzia; Wijgerde, Tim (19 November 2014). "Black corals". Reefs. Retrieved 4 May 2020.
  15. 1 2 3 Wagner, Daniel; Toonen, R. J. (2012). "The biology and ecology of black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia)". Advanced Marine Biology. 63 (132): 63–67. doi:10.1016/B978-0-12-394282-1.00002-8. PMID   22877611.
  16. Gress, Erika; Opresko, Dennis M.; Brugler, Mercer R.; Wagner, Daniel; Eeckhaut, Igor; Terrana, Lucas (2020-12-09). "Widest geographic distribution of a shallow and mesophotic antipatharian coral (Anthozoa: Hexacorallia): Antipathes grandis VERRILL, 1928 – confirmed by morphometric and molecular analyses". Marine Biodiversity Records. 13 (1): 12. doi: 10.1186/s41200-020-00195-0 . ISSN   1755-2672.
  17. 1 2 3 4 5 6 7 8 "Black Coral". Waikiki Aquarium. 2013-11-21.
  18. Boland, Raymond C.; Parrish, Frank A. (1 July 2005). "A Description of Fish Assemblages in the Black Coral Beds off Lahaina, Maui, Hawai'i". Pacific Science. 59 (3): 411–420. doi:10.1353/psc.2005.0032. hdl: 10125/24187 . S2CID   41135750.
  19. Murphy, Richard C. (2002). Coral Reefs: Cities Under The Seas. The Darwin Press, Inc. ISBN   978-0-87850-138-0.
  20. Bo, Marzia; Baker, Andrew C.; Gaino, Elda; Wirsching, Herman H.; Scoccia, Francesca; Bavestrello, Giorgio (2011). "First description of algal mutualistic endosymbiosis in a black coral (Anthozoa: Antipatharia)". Marine Ecology Progress Series. 435 (13): 1–11. Bibcode:2011MEPS..435....1B. doi: 10.3354/meps09228 .
  21. Goldberg, Walter M. (1991). "Chemistry and structure of skeletal growth rings in the black coral Antipathes fiordensis (Cnidaria, Antipatharia)". Coelenterate Biology: Recent Research on Cnidaria and Ctenophora. pp. 403–409. doi:10.1007/978-94-011-3240-4_58. ISBN   978-94-010-5428-7.{{cite book}}: |journal= ignored (help)
  22. Grigg, Richard (April 1965). "Ecological Studies of Black Coral in Hawaii". Pacific Studies. 19: 244–260. hdl:10125/4416 . Retrieved 12 November 2019.
  23. Parker, N. R.; Mladenov, P. V.; Grange, K. R. (November 1997). "Reproductive biology of the antipatharian black coral Antipathes fiordensis in Doubtful Sound, Fiordland, New Zealand". Marine Biology. 130 (130): 11–22. doi:10.1007/s002270050220. S2CID   85999468.
  24. Roark, E. B.; Guilderson, T. P.; Dunbar, R. B.; Fallon, S. J.; Mucciarone, D. A. (10 February 2009). "Extreme longevity in proteinaceous deep-sea corals". Proceedings of the National Academy of Sciences of the United States of America. 106 (13): 5204–5208. doi: 10.1073/pnas.0810875106 . PMC   2663997 . PMID   19307564.
  25. Graczyk, Michael (25 March 2009). "Scientists ID living coral as 4,265 years old". The Associated Press.
  26. 1 2 3 4 Bruckner, Andrew W. (2016), "Advances in Management of Precious Corals to Address Unsustainable and Destructive Harvest Techniques", The Cnidaria, Past, Present and Future, Springer International Publishing, pp. 747–786, doi:10.1007/978-3-319-31305-4_46, ISBN   9783319313030
  27. Wagner, Daniel; Luck, Daniel G.; Toonen, Robert J. (1 January 2012). The Biology and Ecology of Black Corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia). pp. 67–132. doi:10.1016/B978-0-12-394282-1.00002-8. ISBN   9780123942821. ISSN   0065-2881. PMID   22877611.{{cite book}}: |journal= ignored (help)
  28. Hickson, Sydney J. (July 1922). "Black Coral". Nature. 110 (2754): 207–208. Bibcode:1922Natur.110..217H. doi: 10.1038/110217a0 . Retrieved 28 October 2019.
  29. Grigg, Richard W. (1 July 2001). "Black Coral: History of a Sustainable Fishery in Hawai'i" (PDF). Pacific Science. 55 (3): 291–299. doi:10.1353/psc.2001.0022. hdl: 10125/2453 . ISSN   1534-6188. S2CID   38992352.
  30. Kenyon, J. (1984). "Black coral off Cozumel". Sea Frontiers. 30: 267–272.
  31. Padilla, C., & Lara, M. (2003). Banco Chinchorro: the last shelter for black coral in the Mexican Caribbean. Bulletin of Marine Science, 73(1), 197–202.
  32. Gress, Erika; Andradi-Brown, Dominic A. (4 July 2018). "Assessing population changes of historically overexploited black corals (Order: Antipatharia) in Cozumel, Mexico". PeerJ. 6: e5129. doi: 10.7717/peerj.5129 . ISSN   2167-8359. PMC   6035717 . PMID   30013832.
  33. "Appendices". CITES. Retrieved 29 October 2019.
  34. Montgomery, Anthony D. (March 2002). "The feasibility of transplanting black coral (Order Antipatharia)". Hydrobiologia. 471 (4711): 157–164. doi:10.1023/A:1016573926566. S2CID   12598714.
  35. 1 2 Terrana, Lucas; Todinanahary, Gildas Georges Boleslas; Eeckhaut, Igor (24 June 2016). Illegal harvesting and trading of black corals (Antipatharia) in Madagascar: the necessity of field studies. 13th International Coral Reef Symposium.
  36. 1 2 "Case Study for Black Coral from Hawaii" (PDF). CITES. Retrieved 4 November 2019.
  37. Grange, K. R. (18 Feb 1985). "Distribution, standing crop, population structure, and growth rates of black coral in the southern fiords of New Zealand". New Zealand Journal of Marine and Freshwater Research. 19 (4): 467–475. doi: 10.1080/00288330.1985.9516111 .
  38. "How does Climate Change Affect Coral Reefs?". NOAA. Retrieved 28 October 2019.
  39. Guinotte, John (2005). "Climate Change and Deep-sea Corals" (PDF). The Journal of Marine Education. 21 (4). Retrieved 4 November 2019.