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Groupers are fish of any of a number of
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
in the subfamily Epinephelinae of the family Serranidae, in the order Perciformes. Not all serranids are called "groupers"; the family also includes the sea basses. The common name "grouper" is usually given to fish in one of two large
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
: '' Epinephelus'' and '' Mycteroperca''. In addition, the species classified in the small genera ''Anyperidon'', ''Cromileptes'', ''Dermatolepis'', ''Graciela'', ''Saloptia'', and ''Triso'' are also called "groupers." Fish in the genus '' Plectropomus'' are referred to as "coral groupers." These genera are all classified in the subfamily Epiphelinae. However, some of the hamlets (genus ''Alphestes''), the hinds (genus ''Cephalopholis''), the lyretails (genus ''Variola''), and some other small genera (''Gonioplectrus'', ''Niphon'', ''Paranthias'') are also in this subfamily, and occasional species in other serranid genera have common names involving the word "grouper." Nonetheless, the word "grouper" on its own is usually taken as meaning the subfamily Epinephelinae.


Description

Groupers are teleosts, typically having a stout body and a large mouth. They are not built for long-distance, fast swimming. They can be quite large: in length, over a meter. The largest is the Atlantic goliath grouper (''Epinephelus itajara'') which has been weighed at and a length of , though in such a large group, species vary considerably. They swallow prey rather than biting pieces off of them. They do not have many teeth on the edges of their jaws, but they have heavy crushing tooth plates inside the pharynx. They habitually eat fish,
octopus An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttle ...
es, and crustaceans. Some species prefer to ambush their prey, while others are active predators. Reports of fatal attacks on humans by the largest species, such as the giant grouper (''Epinephelus lanceolatus''), are unconfirmed. Their mouths and gills form a powerful vacuum that pulls their prey in from a distance. They also use their mouths to dig into sand to form their shelters under big rocks, jetting it out through their gills. Research indicates roving coralgroupers (''Plectropomus pessuliferus'') sometimes cooperate with giant morays in hunting. Groupers are also one of the only animals that eat invasive red lionfish.


Systematics


Etymology

The word "grouper" is from the Portuguese name, ''garoupa'', which has been speculated to come from an indigenous South American language. In Australia, "groper" is used instead of "grouper" for several species, such as the Queensland grouper (''Epinephelus lanceolatus''). In New Zealand, "groper" refers to a type of wreckfish, ''Polyprion oxygeneios'', which goes by the Māori name '' hāpuku''. In the Philippines, groupers are generally known as ''lapu-lapu'' in Luzon, while in the Visayas and Mindanao they are known as ''pugapo''. In the Middle East, the fish is known as ' hammour', and is widely eaten, especially in the Persian Gulf region. In Latin America, the fish is known as ' mero'. The species in the tribes Grammistini and Diploprionini secrete a mucus-like toxin in their skin called
grammistin Grammistins are peptide toxins synthesised by glands in the skin of soapfishes of the tribes Grammistini and Diploprionini which are both classified within the grouper subfamily Epinephelinae, a part of the family Serranidae. Grammistin has a hemoly ...
, and when they are confined in a restricted space and subjected to stress, the mucus produces a foam that is toxic to nearby fish. These fishes are often called soapfishes. They have been classified either as their own families or within subfamilies,, although they are classified by the 5th Edition of the ''Fishes of the World'', classifies these two groups as tribes within the subfamily Epinephelinae.


Classification

According to the 5th edition of ''
Fishes of the World ''Fishes of the World'' by the American ichthyologist Joseph S. Nelson (1937–2011) is a standard reference for fish systematics. Now in its fifth edition (2016), the work is a comprehensive overview of the diversity and classification of the ...
,'' the subfamily is divided up into 5 tribes containing a total of 32 genera and 234 species. Subfamily Epinephelinae
Bleeker Bleeker is a Dutch occupational surname. Bleeker is an old spelling of ''(linnen)bleker'' ("linen bleacher").Tribe
Niphonini The ara (''Niphon spinosus''), otherwise known as the saw-edged perch or the Dageumbari (다금바리) is a species of marine ray-finned fish from the monospecific genus ''Niphon'' which is in the monotypic tribe of the Niphonini which is part of ...
D.S. Jordan, 1923 ** '' Niphon''
Cuvier Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in nat ...
, 1828
*Tribe Epinephelini Bleeker, 1874 ** '' Aethaloperca'' Fowler, 1904 ** '' Alphestes''
Bloch Bloch is a surname of German origin. Notable people with this surname include: A–F * (1859-1914), French rabbi *Adele Bloch-Bauer (1881-1925), Austrian entrepreneur *Albert Bloch (1882–1961), American painter * (born 1972), German motor journal ...
&
Schneider Schneider may refer to: Hospital * Schneider Children's Medical Center of Israel People * Schneider (surname) Companies and organizations * G. Schneider & Sohn, a Bavarian brewery company * Schneider Rundfunkwerke AG, the former owner of th ...
, 1801
** '' Anyperodon'' Günther, 1859 ** '' Cephalopholis'' Bloch & Schneider, 1801 ** '' Chromileptes'' Swainson, 1839 ** '' Dermatolepis'' Gill, 1861 ** '' Epinephelus'' Bloch, 1793 ** '' Gonioplectrus'' Gill, 1862 ** '' Gracila''
Randall Randall may refer to the following: Places United States *Randall, California, former name of White Hall, California, an unincorporated community * Randall, Indiana, a former town *Randall, Iowa, a city *Randall, Kansas, a city *Randall, Minnesot ...
, 1964
** '' Hyporthodus'' Gill, 1861 ** '' Mycteroperca'' Gill, 1862 ** '' Paranthias'' Guichenot, 1868 ** '' Plectropomus'' Pken, 1817 ** '' Saloptia''
J.L.B. Smith James Leonard Brierley Smith (26 September 1897 – 8 January 1968) was a South African ichthyologist, organic chemist, and university professor. He was the first to identify a taxidermied fish as a coelacanth, at the time thought to be long e ...
, 1964
** '' Triso'' Randall, Johnson & Lowe, 1989 ** '' Variola'' Swainson, 1839 *Tribe
Diploprionini Diploprionini is one of the five tribes in the subfamily Epinephelinae, the groupers, which is part of the family Serranidae which also includes the anthias and the sea basses. Genera The following genera are included within the Diploprionini: ...
Bleeker, 1874 ** '' Aulacocephalus'' Temminck & Schlegel, 1843 ** ''
Belonoperca ''Belonoperca'' is a genus of marine ray-finned fish, related to the groupers and included in the subfamily Epinephelinae, part of the family Serranidae, which also includes the anthias and sea basses. They are found in the Indo-Pacific region. ...
'' Fowler & B.A. Bean, 1930 ** ''
Diploprion ''Diploprion'' is a genus of marine ray-finned fish, related to the groupers and included in the subfamily Epinephelinae, part of the family Serranidae, which also includes the anthias Anthias are members of the family Serranidae and make up ...
'' Cuvier, 1828 * Tribe
Liopropomini Liopropomini is one of the five tribes in the subfamily Epinephelinae, the group including the groupers, which is part of the family Serranidae which also includes the anthias and the sea basses. They are found mainly in the Indo-Pacific region a ...
Poey, 1867 ** ''
Bathyanthias ''Bathyanthias'' is a genus of marine ray-finned fish, related to the groupers and included in the subfamily Epinephelinae, part of the family Serranidae, which also includes the anthias and sea basses. They are found in the Western Atlantic O ...
'' Günther, 1880 ** ''
Liopropoma ''Liopropoma'' is a genus of marine ray-finned fish, related to the groupers and included in the subfamily Epinephelinae, part of the family Serranidae, which also includes the anthias and sea basses. They are sometimes seen in the mari ...
'' Gill, 1861 ** ''
Rainfordia The rainfordia (''Rainfordia opercularis''), also known as the flathead perch or Rainford's perch, is a species of marine ray-finned fish, related to the groupers and classified within the subfamily Epinephelinae of the Family (biology), family S ...
'' McCulloch, 1923 *Tribe Grammistini Bleeker, 1857 ** '' Aporops''
Schultz Schultz is a Germans, German surname derived from ''Schultheiß'', meaning village headman or constable/sheriff in the medieval sense (akin to today's office of mayor). It has many variations, such as Schuldt, Schulte, Schulten, Schultes, Schultheis ...
, 1943
** '' Grammistes'' Bloch & Schneider, 1801 ** ''
Grammistops The ocellate soapfish (''Grammistops ocellatus'') is a species of marine ray-finned fish with a wide Indo-Pacific distribution. It is the only species in the genus ''Grammistops''. It is also known as the ocellated soapfish, ocellated podge, o ...
'' Schultz 1953 ** '' Jeboehlkia'' Robins, 1967 ** ''
Pogonoperca ''Pogonoperca'' is a genus of marine ray-finned fish, related to the groupers and classified within the subfamily Epinephelinae of the family Serranidae. They are found in the Indo-Pacific region. Etymology The genus name ''Pogonoperca'' derives ...
'' Günther 1859 ** '' Pseudogramma'' Bleeker, 1875 ** ''
Rypticus ''Rypticus'' is a genus of marine ray-finned fish, related to the groupers and classified within the subfamily Epinephelinae of the family Serranidae. It is one of several genera of soapfishes. These fish live in the Atlantic and eastern Pacific ...
'' Cuvier, 1829 ** ''
Suttonia :''In plants, ''Suttonia'' is a junior synonym of the colicwood genus, ''Myrsine. ''Suttonia'' is a genus of marine ray-finned fish, related to the groupers and classified within the subfamily Epinephelinae of the family Serranidae. Species The ...
'' J.L.B. Smith, 1953


Reproduction

Groupers are mostly monandric protogynous hermaphrodites, i.e., they mature only as females and can change sex after sexual maturity.Erisman, B. E., M. T. Craig, and P. A. Hastings. 2009. A phylogenetic test of the size-advantage model: Evolutionary changes in mating behavior influence the loss of sex change in a fish lineage. American Naturalist 174:83-99.DeMartini, E. E., A. R. Everson and R. S. Nichols. 2011. Estimates of body sizes at maturation and sex change and the endemic Hawaiian grouper's spawning seasonality and sex ratio (''Hyporthodus queries'', f. Epinephelidae). Fishery Bulletin 109:123-134. Some species of groupers grow about a kilogram per year and are generally adolescents until they reach three kilograms when they become female. The largest males often control harems containing three to 15 females.Sadovy, Y. and P. L. Colin. 1995. Sexual development and sexuality in the Nassau grouper. Journal of Fish Biology 46:961-976. Groupers often pair spawn, which enables large males to competitively exclude smaller males from reproducing.Allsop, D. J. and S. A. West. 2003. Constant relative age and size at sex change for sequentially hermaphroditic fish. Journal of Evolutionary Biology 16:921-929.Munoz, R. C. and R. R. Warner. 2003. A new version of the size-advantage hypothesis for sex change: Incorporating sperm competition and size-fecundity skew. American Naturalist 161:749-761.Kuwamura, T. 2004. Sex change in fishes: Its process and evolutionary mechanism. Zoological Science 21:1248-1248. As such, if a small female grouper were to change sex before it could control a harem as a male, its fitness would decrease. If no male is available, the largest female that can increase fitness by changing sex will do so. However, some groupers are gonochoristic. Gonochorism, or a reproductive strategy with two distinct sexes, has evolved independently in groupers at least five times. The evolution of gonochorism is linked to group spawning high amounts of habitat cover.Erisman, B. E., J. A. Rosales-Cassian and P. A. Hastings. 2008. There is evidence of gonochorism in a grouper, ''Mycteroperca rosacea'', from the Gulf of California, Mexico. Environmental Biology of Fishes 82:23-33. Both group spawning and habitat cover increase the likelihood of a smaller male reproducing in the presence of large males. The fitness of male groupers in environments where competitive exclusion of smaller males is impossible is correlated with sperm production and thus testicle size.Molloy, P. P., N. B. Goodwin, I. M. Cote, J. D. Reynolds, and M. J. G. Gage. 2007. Sperm competition and sex change: A comparative analysis across fishes. Evolution 61:640-652. Gonochoristic groupers have larger testes than protogynous groupers (10% of body mass compared to 1% of body mass), indicating the evolution of gonochorism increased male grouper fitness in environments where large males were unable to competitively exclude small males from reproducing.


Parasites

Like other fish, groupers harbor parasites, including
digenea Digenea (Gr. ''Dis'' – double, ''Genos'' – race) is a class of trematodes in the Platyhelminthes phylum, consisting of parasitic flatworms (known as ''flukes'') with a syncytial tegument and, usually, two suckers, one ventral and one oral. ...
ns,
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s,
cestode Cestoda is a class of parasitic worms in the flatworm phylum (Platyhelminthes). Most of the species—and the best-known—are those in the subclass Eucestoda; they are ribbon-like worms as adults, known as tapeworms. Their bodies consist of man ...
s, monogeneans,
isopod Isopoda is an order of crustaceans that includes woodlice and their relatives. Isopods live in the sea, in fresh water, or on land. All have rigid, segmented exoskeletons, two pairs of antennae, seven pairs of jointed limbs on the thorax, an ...
s, and
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s. A study conducted in
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has shown that coral reef-associated groupers have about ten species of parasites per fish species. Species of ''
Pseudorhabdosynochus ''Pseudorhabdosynochus'' is a genus of monopisthocotylean monogeneans, included in the family Diplectanidae. The type-species of the genus is ''Pseudorhabdosynochus epinepheli'' ( Yamaguti, 1938). The genus includes more than 80 species, which a ...
'', monogeneans of the family Diplectanidae are typical of and especially numerous on groupers.


Modern use

Many groupers are important food fish; some are now
farmed Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people t ...
. Unlike most other fish species, which are chilled or frozen, groupers are usually sold live in markets. Many species are popular game fish for sea-angling. Some species are small enough to be kept in aquaria, though even the small species are inclined to grow rapidly. Groupers are commonly reported as a source of Ciguatera fish poisoning. DNA barcoding of grouper species might help control Ciguatera fish poisoning since fish are easily identified, even from meal remnants, with molecular tools.


Size

Malaysian newspaper ''
The Star ''The'' () is a grammatical article in English, denoting persons or things already mentioned, under discussion, implied or otherwise presumed familiar to listeners, readers, or speakers. It is the definite article in English. ''The'' is the m ...
'' reported a grouper being caught off the waters near
Pulau Sembilan Pulau may refer to: *Pulau virus (PuV), a novel strain of Nelson Bay orthoreovirus species *Pulau River, a river of West Papua and Papua New Guinea See also *List of islands of Malaysia – "Pulau" means "island" in the Malay language *Pulao (di ...
in the Strait of Malacca in January 2008. Shenzhen News in China reported that a grouper swallowed a whitetip reef shark at the Fuzhou Sea World aquarium. In September 2010, a Costa Rican newspaper reported a grouper in Cieneguita, Limón. The weight of the fish was , and it was lured using one kilogram of bait. In November 2013, a grouper had been caught and sold to a hotel in Dongyuan, China. In August 2014, off
Bonita Springs (beautiful), eng, beautiful springs , settlement_type = City , image_skyline = Bonita_Beach.JPG , imagesize = 250x200px , image_caption = Bonita Beach , image_flag ...
in Florida (USA), a big grouper took in one gulp a 4-foot shark that an angler had caught.


See also

*
Dusky grouper ''Epinephelus marginatus'' (), the dusky grouper, yellowbelly rock cod or yellowbelly grouper, is a species of marine ray-finned fish, a grouper from the subfamily Epinephelinae which is part of the family Serranidae, which also includes the an ...


References


External links

*ARKive
images and movies of the humpback grouper ''(Chromileptes altivelis)''
* {{Taxonbar, from=Q5559352 Epinephelinae Serranidae Hawaiian cuisine Commercial fish Fish common names Cuisine of the Southern United States