Perciformes (), also called the Acanthopteri, is an

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

or superorder 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 ...

in the clade

Percomorpha Percomorpha () is an extremely large and diverse clade of ray-finned fish. With more than 17,000 known species (including Scombroidei, tuna, Syngnathiformes, seahorses, gobies, Cichlidae, cichlids, flatfish, Labridae, wrasse, Perciformes, perches ...

. ''Perciformes'' means "

perch Perch is a common name for freshwater fish from the genus ''Perca'', which belongs to the family Percidae of the large order Perciformes. The name comes from , meaning the type species of this genus, the European perch (''P. fluviatilis'') ...

-like". Among the well-known members of this group are perches and darters (

Percidae The Percidae are a family of ray-finned fish, part of the order Perciformes, which are found in fresh and brackish waters of the Northern Hemisphere. The majority are Nearctic, but there are also Palearctic species. The family contains nearly 250 ...

), and also sea basses and groupers (

Serranidae Serranidae is a large family (biology), family of fishes belonging to the order Perciformes. The family contains about 450 species in 65 genera, including the sea basses and the groupers (subfamily Epinephelinae). Although many species are small, ...

). This order contains many familiar freshwater temperate and tropical marine fish groups, but also

extremophiles An extremophile () is an organism that is able to live (or in some cases thrive) in extreme environments, i.e., environments with conditions approaching or stretching the limits of what known life can adapt to, such as extreme temperature, pres ...

that have successfully colonized both the North and South Poles, as well as the deepest depths of the ocean.

Taxonomy

Formerly, this group was thought to be even more diverse than it is thought to be now, containing about 41% of all

bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...

(about 10,000 species) and about 160 families, which is the most of any order within the vertebrates. However, many of these other families have since been reclassified within their own orders within the clade Percomorpha, significantly reducing the size of the group. In contrast to this splitting, other groups formerly considered distinct, such as the

Scorpaeniformes The Scorpaeniformes are a diverse Order (biology), order of Actinopterygii, ray-finned fish, including the lionfishes and sculpins, but have also been called the Scleroparei. It is one of the five largest orders of bony fishes by number of spec ...

, are now classified in the Perciformes.

Evolution

The earliest fossil perciform is the extinct

stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...

-perciform '' Paleoserranus'' (originally considered an early serranid) from the

Early Paleocene The Danian is the oldest age (geology), age or lowest stage (stratigraphy), stage of the Paleocene Epoch or series (stratigraphy), Series, of the Paleogene Period or system (stratigraphy), System, and of the Cenozoic Era or Erathem. The beginnin ...

Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...

, but potential records of "percoids" are known from the

Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...

, including '' Eoserranus'' and '' Prolates'', although their exact taxonomic identity remains uncertain. The earliest crown-group perciform fossils are known from the

Early Eocene In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...

, including the scorpaenoid ''

Eosynanceja ''Eosynanceja'' ("dawn '' Synanceia''") is an extinct genus of prehistoric scorpionfish that lived during the Eocene. It contains a single species, ''E. brabantica'', known from the Early Eocene of Belgium. It is the oldest known scorpaenoid� ...

'' and platycephalid otoliths from New Zealand.

Classification

Classification of this group has long been controversial, with various families being placed in and out of Perciformes depending on the study. Only in recent decades, with the advent of

molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

, has the classification of the family been largely resolved. Based on these studies, many suborders formerly placed within the Perciformes are better placed elsewhere in the Percomorpha, but former members of the

Gasterosteiformes 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 a ...

, and some members of the

Trachiniformes Trachiniformes is an Order (taxonomy), order of Percomorpha, percomorph bony fish, whose contents are traditionally placed in suborder Trachinoidei of Perciformes. However, Trachiniformes is recovered as polyphyletic in recent large scale molec ...

(including the type genus) are now considered true perciforms.

Present classification

The following classification is based on ''

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 ...

'': * Order Perciformes ** Genus †'' Paleoserranus'' Cantalice, Alvarado-Ortega & Alaniz-Galvan, 2018 ** Suborder

Percoidei Percoidei is a suborder of bony fishes in the order Perciformes. Many commercially harvested fish species are considered to be contained in this suborder, including the groupers, Serranidae, seabasses and perches. Divisions The following classif ...

*** Family

Swainson, 1839 (sea basses) *** Family

Anthiadidae Anthias are members of the family Anthiadidae in the order Perciformes. The group has also been called Anthiidae or Anthiinae, but these names are preoccupied by a subfamily of ground beetles in the family Carabidae erected by Bonelli in 1813 ...

Poey, 1861 (fairy basslets or streamer basses) *** Family Epinephelidae Bleeker, 1874 (groupers) *** Family Liopropomatidae Poey, 1867 (painted basslets) *** Family Grammistidae Bleeker, 1857 (soapfishes) *** Family

Rafinesque Constantine Samuel Rafinesque-Schmaltz (; 22 October 178318 September 1840) was a French early 19th-century polymath born near Constantinople in the Ottoman Empire and self-educated in France. He traveled as a young man in the United States, ult ...

, 1815 ( perches and darters) *** Family Niphonidae

Jordan Jordan, officially the Hashemite Kingdom of Jordan, is a country in the Southern Levant region of West Asia. Jordan is bordered by Syria to the north, Iraq to the east, Saudi Arabia to the south, and Israel and the occupied Palestinian ter ...

, 1923 (Ara groupers) *** Family Trachinidae Rafinesque, 1815 (weeverfishes) *** Family Bembropidae Regan, 1913 (flatheads or duckbill flatheads) ** 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 ...

*** Family Percophidae Swainson, 1839 (Brazilian flatheads) *** Family

Bovichtidae Bovichtidae, the temperate icefishes or thornfishes, is a family of marine ray-finned fishes, classified in the suborder Notothenioidei of the order Perciformes. They are native to coastal waters off Australia, New Zealand, and South America ...

Gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...

, 1862 (thornfishes) *** Family

Pseudaphritidae The congoli (''Pseudaphritis urvillii''), also known as the freshwater flathead, marble fish, marbled flathead, sand trout, sanding, sandy, sandy whiting or tupong, is a species of marine ray-finned fish. It is the only species of fish in the ...

McCulloch, 1929 (congollis) *** Family

Eleginopidae The Patagonian blennie (''Eleginops maclovinus''), also known as the rock cod, is a species of marine ray-finned fish, belonging to the monotypic family Eleginopidae and monotypic genus ''Eleginops''. It is found in coastal and estuarine habita ...

Gill, 1893 (Patagonian blennies) *** Family Nototheniidae Günther, 1861 (cod icefishes) *** Family Harpagiferidae Gill, 1861 (plunderfishes) *** Family Bathydraconidae Regan, 1913 (Antarctic dragonfishes) *** Family

Channichthyidae The crocodile icefish or white-blooded fish comprise a family (biology), family (Channichthyidae) of notothenioid fishes, fish found in the Southern Ocean around Antarctica. They are the only known vertebrates to lack hemoglobin in their blood as ...

Gill, 1861 (crocodile icefishes) ** 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 ...

*** Family

Platycephalidae The Platycephalidae are a Family (biology), family of marine ray-finned fish, most commonly referred to as flatheads. They are relatives of the lionfish, and belong to the order Perciformes. Taxonomy Platycephalidae was first proposed as a famil ...

Swainson, 1839 (flatheads) *** Family Hoplichthyidae Kaup, 1873 (spiny flatheads) *** Family Triglidae Rafinesque, 1815 (searobins) *** Family

Bembridae Bembridae, the deep-water flatheads, are a family of bottom-dwelling ray-finned fishes. They are found in the Indian and western Pacific Oceans. Taxonomy Bembridae was first proposed as a family in 1873 by the German zoologist Johann Jakob Kaup ...

Kaup, 1873 (deepwater flatheads) *** Family

Synanceiidae Synanceiinae is a subfamily of Venom, venomous ray-finned fishes, waspfishes, which is classification of life, classified as part of the family Scorpaenidae, the scorpionfishes and their relatives. These fishes are found in the Indo-Pacific ocean ...

Swainson, 1839 (stonefishes) *** Family Neosebastidae

Matsubara is a city located in Osaka Prefecture, Japan. , the city had an estimated population of 117,811 in 57351 households and a population density of 7100 persons per km2. The total area of the city is . Geography Matsubara is located in the center ...

, 1943 (gurnard scorpionfishes) *** Family Plectrogeniidae Fowler, 1938 (stinger flatheads) *** Family

Scorpaenidae The Scorpaenidae (also known as scorpionfish) are a family (biology), family of mostly ocean, marine fish that includes many of the world's most venomous species. As their name suggests, scorpionfish have a type of "sting" in the form of sharp ...

Risso, 1827 (scorpionfishes) *** Family Congiopodidae Gill, 1889 (racehorses or pigfishes) *** Family Zanclorhynchidae Andriashev, 1993 (horsefishes) *** Family Normanichthyidae Clark, 1937 (barehead scorpionfishes ) ** 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 ...

*** Family Anoplopomatidae Jordan & Gilbert ,1883 (sablefishes) *** Family

Zaniolepididae The Zaniolepididae is a Family (biology), family of marine ray-finned fishes classified within the suborder Cottoidei of the Order (biology), order Perciformes. They are found in the North Pacific Ocean. Taxonomy ''Zaniolepididae'' was first pro ...

Jordan & Gilbert, 1883 (combfishes) *** Family

Hexagrammidae Hexagrammidae, the greenlings, is a Family (biology), family of marine ray-finned fishes belonging to the suborder Cottoidei in the Order (biology), order Perciformes. These fishes are found in the North Pacific Ocean. Taxonomy Hexagrammidae wa ...

Jordan, 1888 (greenlings) *** Family

Rhamphocottidae Rhamphocottidae is a family of ray-finned fishes belonging to the superfamily Cottoidea, the sculpins. The species in this family occur in the North Pacific Ocean. Taxonomy Rhamphocottidae was first proposed as a family by the American icht ...

Jordan & Gilbert, 1883 (horsehead sculpins) *** Family

Jordaniidae Jordaniidae is a small Family (biology), family of marine ray-finned fishes belonging to the Order (biology), order Perciformes. These fishes are found in the eastern North Pacific Ocean. Taxonomy Jordaniidae was first proposed as a subfamily, Jo ...

Jordan & Evermann, 1898 (longfin sculpins) *** Family

Cottidae The Cottidae are a family of fish in the superfamily Cottoidea, the sculpins. It is the largest sculpin family, with about 275 species in 70 genera.Kane, E. A. and T. E. Higham. (2012)Life in the flow lane: differences in pectoral fin morphology ...

Bonaparte, 1831 (sculpins) *** Family

Psychrolutidae The fish family Psychrolutidae (commonly known as blobfishes, flathead sculpins, or tadpole sculpins) contains over 35 recognized species in 8 genera. This family consists of bottom-dwelling marine sculpins shaped like tadpoles, with large heads ...

Günther, 1861 (marine sculpins) *** Family

Agonidae Agonidae is a family of small, bottom-dwelling, cold-water marine fish. Common names for members of this family include poachers, Irish lords, sea ravens, alligatorfishes, starsnouts, hooknoses, and rockheads. They are notable for having elongate ...

Swainson, 1839 (poachers) *** Family

Trichodontidae Trichodontidae, the sandfishes, is a small Family (biology), family of ray-finned fishes from the Order (biology), order Perciformes. The species in this family are found in the Pacific Ocean, North Pacific Ocean. Taxonomy Trichodontidae was f ...

Bleeker, 1859 (sandfishes) *** Family Cyclopteridae Bonaparte, 1831 (lumpfishes) *** Family

Liparidae The snailfishes or sea snails (not to be confused with invertebrate sea snails), are a family of marine ray-finned fishes. These fishes make up the Liparidae, a family classified within the order Scorpaeniformes. Widely distributed from the ...

Gill, 1861 (snailfishes) ** 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 ...

*** Family

Hypoptychidae The Korean sandlance (''Hypoptychus dybowskii'') is a species of marine ray-finned fish belonging to the family Hypoptychidae. The Korean sandlance is the only species in this monotypic family and genus and is found in the northwestern Pacific O ...

Steindachner, 1880 (sand-eels) *** Family Gasterosteidae Bonaparte, 1831 (sticklebacks) *** Family

Aulorhynchidae Aulorhynchidae, the tube-snouts, is a small Family (biology), family of marine ray-finned fishes belonging to the suborder Gasterosteoidei in the Order (biology), order Perciformes. These fishes are found in the northern Pacific Ocean. Taxonomy ...

Gill, 1861 (tubesnouts) ** 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 ...

*** Family Zoarcidae Swainson, 1839 (eelpouts) *** Family

Anarhichadidae Anarhichadidae, the wolffishes, sea wolves or wolf eels, is a family of marine ray finned fishes belonging to the Order (biology), order Perciformes. These are predatory, eel shaped fishes which are native to the cold waters of the Arctic, North ...

Bonaparte, 1835 (wolffishes) *** Family Neozoarcidae Jordan & Snyder, 1902 (largemouth kissing eelpouts) *** Family Eulophiidae Smith, 1902 (spinous eelpouts) *** Family

Stichaeidae Stichaeidae, the pricklebacks or shannies, are a Family (biology), family of marine ray-finned fishes in the suborder Zoarcoidei of the Order (biology), order Scorpaeniformes. Most species are found in the North Pacific Ocean with a few in the N ...

Gill, 1864 (pricklebacks) *** Family Lumpenidae Jordan & Evermann, 1898 (eel pricklebacks) *** Family Opisthocentridae Jordan & Evermann, 1898 (rearspined fin pricklebacks) *** Family

Pholidae ''Pholidae'' is a Family (biology), family of marine ray-finned fishes, known as gunnels, in the Scorpaeniformes, scorpaeniform suborder Zoarcoidei. These are fishes of the littoral zone and are mainly found in North Pacific Ocean, with two speci ...

Gill, 1893 (gunnels) *** Family Ptilichthyidae Jordan & Gilbert, 1883 (quillfishes) *** Family Zaproridae Jordan, 1896 (prowfishes) *** Family Cryptacanthodidae Gill, 1861 (wrymouths) *** Family Cebidichthyidae Gill, 1862 (monkeyface pricklebacks) *** Family Scytalinidae Jordan & Starks, 1895 (graveldivers) *** Family

Bathymasteridae Ronquils (sometimes spelt ronchils) is a small Family (biology), family marine ray-finned fish, the Bathymasteridae. These fishes are found only in Arctic Ocean, Arctic and Pacific Ocean, North Pacific waters. This family contains just seven spec ...

Jordan & Gilbert, 1883 (ronquils)

Past classifications

As traditionally defined before the introduction of

cladistics Cladistics ( ; from Ancient Greek 'branch') is an approach to Taxonomy (biology), biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesiz ...

, the Perciformes are almost certainly

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

. Other orders that should possibly be included as suborders are the

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 ...

, and Pleuronectiformes. Of the presently recognized suborders, several may be paraphyletic, as well. These are grouped by suborder/superfamily, generally following the text ''

Fishes of the World ''Fishes of the World'' is a standard reference for the systematics of fishes. It was first written in 1976 by the American ichthyologist Joseph S. Nelson (1937–2011). Now in its fifth edition (2016), the work is a comprehensive overview of t ...

''.

Characteristics

The

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...

and anal fins are divided into

anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...

spiny and posterior soft-rayed portions, which may be partially or completely separated. The

pelvic fins Pelvic fins or ventral fins are paired fins located on the ventral (belly) surface of fish, and are the lower of the only two sets of paired fins (the other being the laterally positioned pectoral fins). The pelvic fins are homologous to the hi ...

usually have one spine and up to five soft rays, positioned unusually far forward under the chin or under the belly. Scales are usually ctenoid (rough to the touch), although sometimes they are

cycloid In geometry, a cycloid is the curve traced by a point on a circle as it Rolling, rolls along a Line (geometry), straight line without slipping. A cycloid is a specific form of trochoid and is an example of a roulette (curve), roulette, a curve g ...

(smooth to the touch) or otherwise modified.

References

{{Authority control Ray-finned fish orders Taxa named by Pieter Bleeker Extant Danian first appearances