Epinephelini

Groupers are a diverse group of marine ray-finned fish in the family Epinephelidae, in the order Perciformes.

Groupers were long considered a subfamily of the seabasses in Serranidae, but are now treated as distinct. Not all members of this family are called "groupers". The common name "grouper" is usually given to fish in one of two large genera: ''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 family Epinephelidae.

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, octopuses, 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 name '' hapuka'' (from the Māori language ). In the Philippines, groupers are generally known as ''lapu-lapu'' in Luzon, while in the Visayas and Mindanao they are known as ''pugapo''. It is known as ''kerapu'' in both Indonesian and Malay. 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, 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 the fifth Edition of the ''Fishes of the World'' classifies these two groups as tribes within the subfamily Epinephelinae. However, '' Eschmeyer's Catalog of Fishes'' presently treats these as distinct families.

Classification

Based on '' Eschmeyer's Catalog of Fishes:''

* Family Epinephelidae Bleeker, 1874 (groupers)
** '' Aethaloperca'' Fowler, 1904
** '' Alphestes'' Bloch & Schneider, 1801
** '' Anyperodon'' Günther, 1859
** '' Cephalopholis'' Bloch & Schneider, 1801
** '' Chromileptes'' Swainson, 1839
** '' Dermatolepis'' Gill, 1861
** ''Epinephelus'' Bloch, 1793
** '' Gonioplectrus'' Gill, 1862
** '' Gracila'' Randall, 1964
** '' Hyporthodus'' Gill, 1861
** '' Mycteroperca'' Gill, 1862
** '' Paranthias'' Guichenot, 1868
** '' Plectropomus'' Pken, 1817
** '' Saloptia'' J.L.B. Smith, 1964
** '' Triso'' Randall, Johnson & Lowe, 1989
** '' Variola'' Swainson, 1839

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 digeneans, nematodes, cestodes, monogeneans, isopods, and copepods. A study conducted in New Caledonia has shown that coral reef-associated groupers have about ten species of parasites per fish species. Species of '' Pseudorhabdosynochus'', 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. Unlike most other fish species, which are chilled or frozen, groupers are usually sold alive 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'' reported a grouper being caught off the waters near Pulau Sembilan 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 in Florida (USA), a big grouper took in one gulp a 4-foot shark that an angler had caught.

See also

* Dusky grouper

References

External links

The Ultimate Guide to Fly Fishing for Grouper
*ARKive �
images and movies of the humpback grouper ''(Chromileptes altivelis)''
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Epinephelinae
Serranidae
Commercial fish
Fish common names
Cuisine of the Southern United States