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Percomorpha () is an extremely large and diverse
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
of
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of sk ...
. With more than 17,000 known species (including
tuna A tuna (: tunas or tuna) is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae ( mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bul ...
, seahorses,
gobies Gobiidae or gobies is a family of bony fish in the order Gobiiformes, one of the largest fish families comprising over 2,000 species in more than 200 genera. Most of gobiid fish are relatively small, typically less than in length, and the fam ...
,
cichlids Cichlids () are a large, diverse, and widespread family of Percomorpha, percomorph fish in the family (biology), family Cichlidae, order Cichliformes. At least 1,760 species have been Binomial nomenclature, scientifically described, making ...
,
flatfish A flatfish is a member of the Ray-finned fish, ray-finned demersal fish Order (biology), suborder Pleuronectoidei, also called the Heterosomata. In many species, both eyes lie on one side of the head, one or the other migrating through or around ...
,
wrasse The wrasses are a family, Labridae, of marine ray-finned fish, many of which are brightly colored. The family is large and diverse, with over 600 species in 81 genera, which are divided into nine subgroups or tribes. They are typically small, ...
, perches,
anglerfish The anglerfish are ray-finned fish in the order Lophiiformes (). Both the order's common name, common and scientific name comes from the characteristic mode of predation, in which a modified dorsal Fish fin#Ray-fins, fin ray acts as a Aggressiv ...
, and
pufferfish Tetraodontidae is a family of marine and freshwater fish in the order Tetraodontiformes. The family includes many familiar species variously called pufferfish, puffers, balloonfish, blowfish, blowers, blowies, bubblefish, globefish, swellfis ...
) known from both marine and freshwater ecosystems, it is the most speciose clade of extant
vertebrates Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
.


Evolution

Percomorpha are the most diverse group of
teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
fish today. Teleosts, and percomorphs in particular, thrived during the
Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...
era. Fossil evidence shows that there was a major increase in size and abundance of teleosts immediately after the mass extinction event at the Cretaceous-Paleogene boundary ago. The oldest known percomorph fossils are of the early tetraodontiforms '' Protriacanthus'' and Cretatriacanthidae from the
Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya ( million years ago) and 83.6 ± 0.7 m ...
to
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
of
Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...
and
Slovenia Slovenia, officially the Republic of Slovenia, is a country in Central Europe. It borders Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the south and southeast, and a short (46.6 km) coastline within the Adriati ...
. A higher diversity of early percomorphs is also known from the
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
of Nardò, Italy, and these also show some level of diversification into modern orders, with representatives of the Syngnathiformes and Tetraodontiformes known. Possibly the oldest percomorph is '' Plectocretacicus'' from the
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
of
Lebanon Lebanon, officially the Republic of Lebanon, is a country in the Levant region of West Asia. Situated at the crossroads of the Mediterranean Basin and the Arabian Peninsula, it is bordered by Syria to the north and east, Israel to the south ...
, which may be a stem-tetraodontiform; however, some morphological analyses indicate that it shows similarities with non-percomorph groups.


Taxonomy

Many of the orders placed in Percomorpha today were originally placed in an expanded
Perciformes Perciformes (), also called the Acanthopteri, is an order or superorder of ray-finned fish in the clade Percomorpha. ''Perciformes'' means " perch-like". Among the well-known members of this group are perches and darters ( Percidae), and als ...
, hence many sources often referring to that order as the most diverse vertebrate clade. However, more recent studies have found such a placement to be paraphyletic, and many have thus been moved to their orders within Percomorpha. The Percomorpha are an extremely diverse group with at least 22 orders according to
Eschmeyer's Catalog of Fishes Catalog of Fishes is a comprehensive on-line database and reference work on the scientific names of fish species and genera. It is global in its scope and is hosted by the California Academy of Sciences. It has been compiled and is continuously up ...
. Other authorities find a different number of orders depending on the classification used. The following taxonomy is based on Eschmeyer's Catalog of Fishes (2025), with additional suborders based on other studies: * Clade Percomorpha ** Subseries Ophidiida *** Order Ophidiiformes **** Suborder Ophidioidei
cusk-eels The cusk-eel family, Ophidiidae, is a group of marine bony fishes in the Ophidiiformes order. The scientific name is from the Greek ''ophis'' meaning "snake", and refers to their eel-like appearance. True eels diverged from other ray-finn ...
, pearlfishes and allies (4 families) **** Suborder Bythoidei
viviparous In animals, viviparity is development of the embryo inside the body of the mother, with the maternal circulation providing for the metabolic needs of the embryo's development, until the mother gives birth to a fully or partially developed juve ...
and false brotulas (2 families) ** Subseries Batrachoidida *** Order Batrachoidiformestoadfishes (1 family) ** Subseries Gobiida *** Order
Gobiiformes Gobiiformes (meaning "goby-like") is an order of percomorph fish containing three suborders: Apogonoidei, Trichonotoidei, and Gobioidei. The order was formerly defined as containing only the gobies (now placed within the Gobioidei). However, ...
**** Suborder Apogonoidei (=order
Kurtiformes The Apogonoidei is a suborder of Gobiiformes, gobiiform fish consisting of two families: the Indo-Pacific Kurtidae (consisting solely of two species in the genus ''Kurtus'') and the much more diverse and widespread Apogonidae (the cardinalfishes) ...
'' sensu auct.'') – nurseryfishes and cardinalfishes (2 families) **** Suborder Trichonotoideisand divers (1 family) **** Suborder Gobioidei
gobies Gobiidae or gobies is a family of bony fish in the order Gobiiformes, one of the largest fish families comprising over 2,000 species in more than 200 genera. Most of gobiid fish are relatively small, typically less than in length, and the fam ...
, sleepers and allies (7 families) ** Subseries Syngnatharia *** Order Syngnathiformes **** Suborder Dactylopteroideiseamoths and flying gurnards (2 families) **** Suborder Mulloideigoatfishes (1 family) **** Suborder Callionymoidei
dragonets Dragonets are small percomorph marine fish of the diverse family Callionymidae (from the Greek ''kallis'', "beautiful" and ', "name") found mainly in the tropical waters of the western Indo-Pacific. They are benthic organisms, spending most of ...
(2 families) **** Suborder
Syngnathoidei The Syngnathiformes are an order of ray-finned fishes that includes the leafy seadragons, sea moths, trumpetfishes and seahorses, among others.FishBase (2005)Order Summary for Syngnathiformes Version of 2005-FEB-15. Retrieved 19 Aug 2008. Th ...
seahorses, pipefish, trumpetfish, cornetfish and shrimpfishes (5 families) ** Subseries Pelagiaria *** Order Scombriformes **** Suborder Stromateoideimedusafishes, driftfishes, butterfishes, squaretails, ariommas and allies (6 families) **** Suborder Scombroidei
mackerel Mackerel is a common name applied to a number of different species of pelagic fish, mostly from the family Scombridae. They are found in both temperate and tropical seas, mostly living along the coast or offshore in the oceanic environment. ...
, tunas, manefishes, pomfrets, cutlassfishes, bluefish, swallowers, ragfish and allies (10 families) ** Subseries Anabantaria *** Order
Synbranchiformes Synbranchiformes, often called swamp eels, though that name can also refer specifically to Synbranchidae, is an order of ray-finned fishes that are eel-like but have spiny rays, indicating that they belong to the superorder Acanthopterygii. ...
**** Suborder Mastacembeloideifreshwater spiny eels and earthworm eels (2 families) **** Suborder Indostomoideiarmored sticklebacks (1 family) **** Suborder Synbranchoideiswamp eels (1 family) *** Order
Anabantiformes The Anabantiformes , is an Order (biology), order of bony fish (Teleostei) proposed in 2009. They are collectively known as labyrinth fish, are an Order (biology), order of air-breathing freshwater Actinopterygii, ray-finned fish with three sub ...
**** Suborder
Anabantoidei The Anabantoidei are a suborder of Anabantiformes, anabantiform ray-finned fish, ray-finned freshwater fish distinguished by their possession of a lung-like labyrinth organ, which enables them to breathe air. The fish in the Anabantoidei suborder ...
gouramies and fighting fishes (3 families) **** Suborder Channoideisnakeheads (2 families) **** Suborder NandoideiAsian leaffishes, chameleonfishes, and mudperches (3 families) ** Subseries Carangaria *** Order
Carangiformes Carangiformes is a large, diverse order of ray-finned fishes within the clade Percomorpha. It is part of a sister clade to the Ovalentaria, alongside its sister group, the Anabantaria (including Anabantiformes and Synbranchiformes). The Cara ...
**** Suborder Centropomoideibarracudas, lates perches, snooks and false trevally (4 families) **** Suborder Pleuronectoideiflatfishes and threadfins (17 families) **** Suborder Toxotoideiarcherfishes and beachsalmons (2 families) **** Suborder Nematistioideiroosterfish (1 family) **** Suborder
Menoidei Menoidei is a suborder of ray-finned fishes belonging to the class Actinopterygii, a diverse group of vertebrates characterized by their bony skeletons. The suborder Menoidei includes the family Menidae, family Xiphiidae and family Istiophorida ...
moonfish and billfishes (3 families) **** Suborder Carangoideijacks, trevallies, pompanos, remoras, cobias and dolphinfishes (4 families) ** Subseries Ovalentaria *** Order
Atheriniformes The Atheriniformes, also known as the silversides, are an order of ray-finned fishes that includes the Old & New World silversides, the rainbowfishes, and several less-familiar families, including the unusual Phallostethidae. The order incl ...
**** Suborder AtherinopsoideiNeotropical silversides (1 family) **** Suborder AtherinoideiOld World silversides, rainbowfishes, priapumfishes and allies (10 families) *** Order
Beloniformes Beloniformes is an order composed of six families (and about 264 species) of freshwater and marine ray-finned fish: * Adrianichthyidae (ricefish and medakas) * Belonidae (needlefish) * Exocoetidae (flyingfishes) * Hemiramphidae (halfb ...
**** Suborder
Adrianichthyoidei The ricefishes are a family (biology), family (Adrianichthyidae) of small ray-finned fish that are found in fresh and brackish waters from India to Japan and out into the Malay Archipelago, most notably Sulawesi (where the Lake Poso and Lore Lin ...
ricefishes and buntingis (1 family) **** Suborder Belonoideisauries, needlefishes, halfbeaks and flyingfishes (5 families) *** Order
Cyprinodontiformes Cyprinodontiformes is an order (biology), order of Actinopterygii, ray-finned fish, comprising mostly small, freshwater fish. Many popular aquarium fish, such as killifish and Poeciliidae, live-bearers, are included. They are closely related t ...
**** Suborder
Aplocheiloidei Aplocheiloidei is a suborder of the Order (biology), order Cyprinodontiformes consisting of three Family (biology), families which are commonly known as killifishes. Families The three families within the suborder Aplocheiloidei are: * Aplochei ...
– Old and New World rivulines (3 families) **** Suborder
Cyprinodontoidei Cyprinodontoidei is a suborder of fishes, one of the two suborders in the order Cyprinodontiformes. The Cyprinodontoidei consists of eleven families which are found in the Americas, the Mediterranean and in Africa, including Madagascar. Classi ...
killifishes, pupfishes, splitfins, livebearers, lampeyes and four-eyed fishes (11 families) *** Order Cichliformes
cichlids Cichlids () are a large, diverse, and widespread family of Percomorpha, percomorph fish in the family (biology), family Cichlidae, order Cichliformes. At least 1,760 species have been Binomial nomenclature, scientifically described, making ...
, leaffishes and convict blenny (3 families) *** Order Mugiliformesmullets and Asiatic glassfishes (2 families) *** Order Blenniiformes **** Unranked clades – dottybacks,
roundheads Roundheads were the supporters of the Parliament of England during the English Civil War (1642–1651). Also known as Parliamentarians, they fought against King Charles I of England and his supporters, known as the Cavaliers or Royalists, who ...
, damselfishes, clownfishes, surfperches and jawfishes (6 families) **** Suborder Gobiesocoideiclingfishes (1 family) **** Suborder Blennioideiblennies (6 families) ** Subseries Eupercaria *** Order
Perciformes Perciformes (), also called the Acanthopteri, is an order or superorder of ray-finned fish in the clade Percomorpha. ''Perciformes'' means " perch-like". Among the well-known members of this group are perches and darters ( Percidae), and als ...
**** Suborder Percoideiperches, darters, seabasses, groupers, anthias, weeverfishes and duckbills (9 families) **** Suborder
Notothenioidei Notothenioidei is one of 19 suborders of the order Perciformes. The group is found mainly in Antarctic and Subantarctic waters, with some species ranging north to southern Australia and southern South America. Notothenioids constitute approximat ...
icefishes, toothfishes, Antarctic dragonfishes and allies (8 families) **** Suborder
Scorpaenoidei Scorpaenoidei is a suborder of Actinopterygii, ray-finned fishes, part of the Order (biology), order Perciformes, that includes the scorpionfishes, lionfishes and velvetfishes. This suborder is at its most diverse in the Pacific and Indian Oceans ...
scorpionfishes, stonefishes, rockfishes, flatheads, searobins and allies (11 families) **** Suborder
Cottoidei Cottoidei is a suborder of ray-finned fishes which, according to the 5th edition of ''Fishes of the World'', is placed within the order Scorpaeniformes, alongside the scorpionfishes, flatheads, eelpouts, sticklebacks and related fishes. Taxon ...
sculpins, psychrolutes, snailfishes, sablefishes, greenlings,
poachers Poaching is the illegal hunting or capturing of wild animals, usually associated with land use rights. Poaching was once performed by impoverished peasants for subsistence purposes and to supplement meager diets. It was set against the hunti ...
, sandfishes, lumpfishes and allies (11 families) **** Suborder
Gasterosteoidei Gasterosteoidei is a suborder of ray-finned fishes that includes the sticklebacks and relatives, the 5th edition of ''Fishes of the World'' classifies this suborder within the order Scorpaeniformes. Systematics Gasterosteoidei is treated as ...
sticklebacks, Korean sandlance and tubesnouts (3 families) **** Suborder
Zoarcoidei Zoarcoidei is a suborder of marine ray-finned fishes belonging to the Order (biology), order Perciformes. The suborder includes the wolffishes, Pholidae, gunnels and eelpouts. The suborder includes about 400 species. These fishes are predominantl ...
eelpouts, wolffishes, pricklebacks, gunnels, prowfish and allies (14 families) *** Order
Centrarchiformes Centrarchiformes is an order of ray-finned fish, previously included amongst the perciformes. Froese, R. and D. Pauly. Editors. (2023). FishBase. Centrarchiformes. Accessed through: World Register of Marine Species at: https://www.marinespecies. ...
**** Suborder PercalatoideiAustralian basses (1 undescribed family) **** Suborder Terapontoideigrunters,
sea chubs The sea chubs, also known as rudderfish and pilot fish and in Hawaiian language, Hawaiian as ''enenue'' or ''nenue'', are a family (biology), family, Kyphosidae, (from Greek, ''kyphos'' = hump) of fishes in the order Perciformes native to the At ...
, flagtails, knifejaws, stripeys and allies (9 families) **** Suborder Centrarchoideifreshwater sunfish, Chinese perches, oldwives, temperate perch, jutjaws and allies (7 families) **** Suborder Cirrhitioideihawkfishes, kelpfishes, marblefishes, fingerfins and trumpeters (5 families) *** Order
Labriformes Labriformes is an Order (biology), order of ray-finned fishes within the clade Percomorpha. Some authors include the Labriformes as the clade Labroidei within the Perciformes while others include more Family (biology), families within the Labrif ...
**** Suborder Labroideiwrasses, parrotfishes, false scorpionfish and weed whitings (3 families) **** Suborder Uranoscopoideisandlances, stargazers, southern sandfishes, sandperches and torrentfish (5 families) *** Order Acropomatiformes (=Pempheriformes) – lanternbellies, sweepers, gnomefishes, wreckfishes, banjofish, oceanic basslets, armorheads and allies (20 families) *** Order Acanthuriformessurgeonfishes, butterflyfishes, rabbitfishes, marine angelfishes, drumfish, grunts, ponyfishes, mojarras, snappers, temperate basses, spadefishes, porgies, moonyfish, tripletails,
tilefish file:Malacanthus latovittatus.jpg, 250px, Blue blanquillo, ''Malacanthus latovittatus'' Tilefishes are mostly small perciform marine fish comprising the family (biology), family Malacanthidae. They are usually found in sandy areas, especially n ...
, scats, bigeyes, boarfishes and allies (30 families) *** Order
Lophiiformes The anglerfish are ray-finned fish in the order Lophiiformes (). Both the order's common and scientific name comes from the characteristic mode of predation, in which a modified dorsal fin ray acts as a lure for prey (akin to a human angler, ...
anglerfish The anglerfish are ray-finned fish in the order Lophiiformes (). Both the order's common name, common and scientific name comes from the characteristic mode of predation, in which a modified dorsal Fish fin#Ray-fins, fin ray acts as a Aggressiv ...
**** Suborder
Lophioidei Goosefishes, sometimes called anglers or monkfishes, are a family, the Lophiidae, of marine ray-finned fishes belonging to the order Lophiiformes, the anglerfishes. The family includes 30 recognized species. These fishes are found in all the ...
goosefishes (1 family) **** Suborder Ogcocephaloideibatfishes (1 family) **** Suborder Antennarioideifrogfishes and handfishes (7 families) **** Suborder Chaunacoideisea toads (1 family) **** Suborder Ceratioidei
deep-sea anglerfish Ceratioidei, the deep-sea anglerfishes or pelagic anglerfishes, is a suborder of marine Actinopterygii, ray-finned fishes, one of five suborders in the Order (biology), order Lophiiformes, the anglerfishes. These fishes are found in tropical and t ...
(11 families) *** Order
Tetraodontiformes Tetraodontiformes (), also known as the Plectognathi, is an order of ray-finned fishes which includes the pufferfishes and related taxa. This order has been classified as a suborder of the order Perciformes, although recent studies have found ...
**** Suborder Triacanthoideispikefishes and triplespines (2 families) **** Suborder Tetraodontoideipufferfishes, porcupinefishes, ocean sunfishes and allies (4 families) **** Suborder Balistoideiboxfishes, filefishes, triggerfishes and allies (4 families)


Phylogeny


External relationships

The two
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
s below are based on Betancur-R ''et al.'', 2017. Percomorphs are a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
of
teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
fishes. The first cladogram shows the interrelationships of percomorphs with other living groups of teleosts.


Internal relationships

The following cladogram shows the evolutionary relationships of the various groups of
extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
percomorph fishes:


References

{{Taxonbar, from= Q22109111, from2=Q258278 Vertebrate unranked clades Taxa named by Edward Drinker Cope