Percomorpha
Temporal range: Santonian–present
PreꞒ
O
S
D
C
P
T
J
K
Pg
N
Possible Cenomanian record
Percomorph diversity (clockwise from top right): Epinephelus marginatus (Perciformes), Sciaenochromis fryeri (Cichliformes), Scomber australasicus (Scombriformes), Valenciennea strigata (Gobiiformes), Scarus niger (Labriformes), Pygoplites diacanthus (Acanthuriformes)
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Superorder: Acanthopterygii
Clade: Percomorpha
Cope, 1871
Subgroups

See text

Synonyms
  • Percomorphaceae Betancur-Rodriguez et al., 2013
  • Acanthopteri

Percomorpha (from Latin perca 'perch' and Ancient Greek μορφή (morphḗ) 'shape, appearance') is an extremely large and diverse clade of ray-finned fish. With more than 17,000 known species (including tuna, seahorses, gobies, cichlids, flatfish, wrasse, perches, anglerfish, and pufferfish) known from both marine and freshwater ecosystems, it is the most speciose clade of extant vertebrates.[1][2][3][4][5][6]

Evolution

Percomorpha are the most diverse group of teleost fish today. Teleosts, and percomorphs in particular, thrived during the Cenozoic 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 c. 66 Ma ago.[7] The oldest known percomorph fossils are of the early tetraodontiforms Protriacanthus and Cretatriacanthidae from the Santonian to Campanian of Italy and Slovenia.[8] A higher diversity of early percomorphs is also known from the Campanian of Nardò, Italy, and these also show some level of diversification into modern orders, with representatives of the Syngnathiformes and Tetraodontiformes known.[9] Possibly the oldest percomorph is Plectocretacicus from the Cenomanian of Lebanon, which may be a stem-tetraodontiform; however, some morphological analyses indicate that it shows similarities with non-percomorph groups.[8][10]

Taxonomy

Evolution of ray-finned fishes, Actinopterygii, from the Devonian to the present as a spindle diagram. The width of the spindles are proportional to the number of families as a rough estimate of diversity. The diagram is based on Benton, M. J. (2005) Vertebrate Palaeontology, Blackwell, 3rd edition, Fig 7.13 on page 185.

Many of the orders placed in Percomorpha today were originally placed in an expanded Perciformes, 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.[6]

The Percomorpha are an extremely diverse group with at least 22 orders according to Eschmeyer's Catalog of Fishes.[11] Other authorities find a different number of orders depending on the classification used.

Numerous fossil percomorphs are known from the Late Cretaceous onwards, but most of these prior to the Eocene, and many afterwards, cannot be confidently assigned to extant percomorph lineages.[9] In the past they were placed within an expanded Perciformes, as with many extant groups, a treatment that is now known to be paraphyletic.[5]

The following taxonomy is based on Eschmeyer's Catalog of Fishes (2025), with additional suborders based on other studies:[5][11]

  • Clade Percomorpha
    • Subseries Ophidiida
      • Order Ophidiiformes
        • Suborder Ophidioideicusk-eels, pearlfishes and allies (4 families)[12]
        • Suborder Bythoideiviviparous and false brotulas (2 families)
    • Subseries Batrachoidida
      • Order Batrachoidiformestoadfishes (1 family)
    • Subseries Gobiida
      • Order Gobiiformes
        • Suborder Apogonoidei (=order Kurtiformes sensu auct.) – nurseryfishes and cardinalfishes (2 families)
        • Suborder Trichonotoideisand divers (1 family)
        • Suborder Gobioideigobies, sleepers and allies (7 families)
    • Subseries Syngnatharia
      • Order Syngnathiformes
        • Suborder Dactylopteroideiseamoths and flying gurnards (2 families)
        • Suborder Mulloideigoatfishes (1 family)
        • Suborder Callionymoideidragonets (2 families)
        • Suborder Syngnathoideiseahorses, pipefish, trumpetfish, cornetfish and shrimpfishes (5 families)
    • Subseries Pelagiaria
      • Order Scombriformes
        • Suborder Stromateoideimedusafishes, driftfishes, butterfishes, squaretails, ariommas and allies (6 families)
        • Suborder Scombroideimackerel, tunas, manefishes, pomfrets, cutlassfishes, bluefish, swallowers, ragfish and allies (10 families)
    • Subseries Anabantaria
      • Order Synbranchiformes
        • Suborder Mastacembeloideifreshwater spiny eels and earthworm eels (2 families)
        • Suborder Indostomoideiarmored sticklebacks (1 family)
        • Suborder Synbranchoideiswamp eels (1 family)
      • Order Anabantiformes
        • Suborder Anabantoideigouramies and fighting fishes (3 families)
        • Suborder Channoideisnakeheads (2 families)
        • Suborder NandoideiAsian leaffishes, chameleonfishes, and mudperches (3 families)
    • Subseries Carangaria
      • Order Carangiformes
        • 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 Menoideimoonfish and billfishes (3 families)
        • Suborder Carangoideijacks, trevallies, pompanos, remoras, cobias and dolphinfishes (4 families)
    • Subseries Ovalentaria
      • Order Atheriniformes[5]
        • Suborder AtherinopsoideiNeotropical silversides (1 family)
        • Suborder AtherinoideiOld World silversides, rainbowfishes, priapumfishes and allies (10 families)
      • Order Beloniformes[5]
        • Suborder Adrianichthyoideiricefishes and buntingis (1 family)
        • Suborder Belonoideisauries, needlefishes, halfbeaks and flyingfishes (5 families)
      • Order Cyprinodontiformes
        • Suborder Aplocheiloidei – Old and New World rivulines (3 families)
        • Suborder Cyprinodontoideikillifishes, pupfishes, splitfins, livebearers, lampeyes and four-eyed fishes (11 families)
      • Order Cichliformescichlids, leaffishes and convict blenny (3 families)
      • Order Mugiliformesmullets and Asiatic glassfishes (2 families)
      • Order Blenniiformes
        • Unranked clades – dottybacks, roundheads, damselfishes, clownfishes, surfperches and jawfishes (6 families)
        • Suborder Gobiesocoideiclingfishes (1 family)
        • Suborder Blennioideiblennies (6 families)
    • Subseries Eupercaria
      • Order Perciformes
        • Suborder Percoideiperches, darters, seabasses, groupers, anthias, weeverfishes and duckbills (9 families)
        • Suborder Notothenioideiicefishes, toothfishes, Antarctic dragonfishes and allies (8 families)
        • Suborder Scorpaenoideiscorpionfishes, stonefishes, rockfishes, flatheads, searobins and allies (11 families)
        • Suborder Cottoideisculpins, psychrolutes, snailfishes, sablefishes, greenlings, poachers, sandfishes, lumpfishes and allies (11 families)
        • Suborder Gasterosteoideisticklebacks, Korean sandlance and tubesnouts (3 families)
        • Suborder Zoarcoideieelpouts, wolffishes, pricklebacks, gunnels, prowfish and allies (14 families)
      • Order Centrarchiformes
        • Suborder PercalatoideiAustralian basses (1 undescribed family)
        • Suborder Terapontoideigrunters, sea chubs, 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
        • 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, scats, bigeyes, boarfishes and allies (30 families)
      • Order Lophiiformesanglerfish
        • Suborder Lophioideigoosefishes (1 family)
        • Suborder Ogcocephaloideibatfishes (1 family)
        • Suborder Antennarioideifrogfishes and handfishes (1 family)
        • Suborder Chaunacoideisea toads (1 family)
        • Suborder Ceratioideideep-sea anglerfish (11 families)
      • Order Tetraodontiformes
        • 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 cladograms below are based on Betancur-R et al., 2017.[5] Percomorphs are a clade of teleost fishes. The first cladogram shows the interrelationships of percomorphs with other living groups of teleosts.


Teleostei

Elopomorpha (Elopiformes, Albuliformes, Notacanthiformes, Anguilliformes)

Osteoglossocephala

Osteoglossomorpha (Hiodontiformes, Osteoglossiformes)

Clupeocephala
Otocephala

Clupei (Clupeiformes)

Alepocephali (Alepocephaliformes)

Ostariophysi (Gonorynchiformes, Cypriniformes, Characiformes, Gymnotiformes, Siluriformes)

Euteleostei
Lepidogalaxii

Lepidogalaxiiformes (salamanderfish)

Protacanthopterygii (Argentiniformes, Galaxiiformes, Esociformes, Salmoniformes)

Stomiati (Stomiiformes, Osmeriformes)

Neoteleostei
Ateleopodia

Ateleopodidae (jellynoses)

Eurypterygia
Aulopa

Aulopiformes (lizardfish)

Ctenosquamata
Scopelomorpha

Myctophiformes (lanternfish)

Acanthomorpha

Lampripterygii (Lampriformes)

Paracanthopterygii (Percopsiformes, Zeiformes, Stylephoriformes, Gadiformes)

Polymixiipterygii

Polymixiiformes (beardfish)

Acanthopterygii
Berycimorphaceae

Beryciformes (alfonsinos, whalefishes)

Trachichthyiformes (pinecone fishes, slimeheads)

Holocentrimorphaceae

Holocentriformes (squirrelfish, soldier fishes)

Percomorpha


Internal relationships

The following cladogram shows the evolutionary relationships of the various groups of extant percomorph fishes:


Percomorpha
Ophidiaria

Ophidiiformes (cusk-eels)

Batrachoidaria

Batrachoididae (toadfishes)

Pelagiaria

Scombriformes (tunas, mackerel, etc.)

Syngnatharia

Syngnathiformes (seahorses, seadragon, etc.)

Gobiaria

Gobiiformes (gobies)

Kurtiformes (nurseryfishes, cardinalfishes)

Anabantaria

Anabantiformes (snakeheads, Siamese fighting fish, gouramies)

Synbranchiformes

Carangaria

Polynemidae (threadfins)

Lactariidae (false trevally)

Menidae (moonfish)

Part of "Carangiformes"

Nematistiidae (roosterfish)

Echeneidae (remoras)

Coryphaenidae (dolphinfish)

Rachycentridae (cobia)

Sphyraenidae (barracudas)

Centropomidae (snooks)

Pleuronectiformes (flatfish)

Part of "Carangiformes"

Carangidae (jacks)

Istiophoriformes (billfish)

Leptobramidae (beachsalmons)

Toxotidae (archerfish)

Ovalentaria
Cichlomorphae

Cichliformes (cichlids, convict blennies)

Polycentridae (leaffish)

Atherinomorphae

Beloniformes (needlefish, flying fish, halfbeaks)

Atheriniformes (silversides, rainbowfish, etc.)

Cyprinodontiformes (tooth-carps)

Ambassidae (Asiatic glassfishes)

Congrogadidae (eel blenny)

Pomacentridae (damselfishes, clownfish)

Embiotocidae (surfperches)

Mugilomorphae

Mugiliformes (mullets)

Lipogramma

Plesiopidae (roundheads)

Pseudochromidae (dottybacks)

Grammatidae (basslets)

Opistognathidae (jawfishes)

Blenniimorphae

Blenniiformes (blennies, clinids, sand stargazers)

Gobiesociformes (clingfishes)

Eupercaria

Gerreiformes (mojarras)

Uranoscopiformes (stargazers)

Centrogenyidae (false scorpionfish)

Labriformes (wrasses, cales, parrotfish)

Perciformes (perches, sticklebacks, scorpionfishes, etc.)

Centrarchiformes (black basses, temperate perches, etc.)

Pempheriformes (sweepers, banjofish, etc.)

Moronidae (temperate basses)

Sillaginidae (smelt-whitings)

Ephippiformes (spadefishes, batfishes)

Chaetodontiformes (butterflyfishes)

Sciaenidae (drums, croakers)

Acanthuriformes (surgeonfishes, ponyfishes)

Monodactylidae (moonyfishes, fingerfishes)

Emmelichthyidae (rovers)

Pomacanthidae (angelfishes)

Lutjaniformes (snappers)

Callanthiidae (splendid perches)

Malacanthidae (tilefishes)

Lobotiformes (tripletails)

Spariformes

Siganidae (rabbitfishes)

Priacanthiformes (bigeyes, bandfishes)

Scatophagidae (scats)

Caproiformes (boarfishes)

Lophiiformes (anglerfishes)

Tetraodontiformes (pufferfishes, triggerfishes, etc.)

References

  1. ^ Harvey, Virginia L.; Keating, Joseph N.; Buckley, Michael (August 2021). "Phylogenetic analyses of ray-finned fishes (Actinopterygii) using collagen type I protein sequences". Royal Society Open Science. 8 (8) 201955. Bibcode:2021RSOS....801955H. doi:10.1098/rsos.201955. PMC 8355665. PMID 34430038.
  2. ^ Thomas J. Near; et al. (2012). "Resolution of ray-finned fish phylogeny and timing of diversification". PNAS. 109 (34): 13698–13703. Bibcode:2012PNAS..10913698N. doi:10.1073/pnas.1206625109. PMC 3427055. PMID 22869754.
  3. ^ Betancur-R, Ricardo; et al. (2013). "The Tree of Life and a New Classification of Bony Fishes". PLOS Currents Tree of Life. 5 (Edition 1). doi:10.1371/currents.tol.53ba26640df0ccaee75bb165c8c26288. hdl:2027.42/150563. PMC 3644299. PMID 23653398.
  4. ^ Laurin, M.; Reisz, R.R. (1995). "A reevaluation of early amniote phylogeny". Zoological Journal of the Linnean Society. 113 (2): 165–223. doi:10.1111/j.1096-3642.1995.tb00932.x.
  5. ^ a b c d e f Betancur-R, Ricardo; Wiley, Edward O.; Arratia, Gloria; Acero, Arturo; Bailly, Nicolas; Miya, Masaki; Lecointre, Guillaume; Ortí, Guillermo (6 July 2017). "Phylogenetic classification of bony fishes". BMC Evolutionary Biology. 17 (1): 162. Bibcode:2017BMCEE..17..162B. doi:10.1186/s12862-017-0958-3. PMC 5501477. PMID 28683774.
  6. ^ a b Nelson, Joseph S.; Grande, Terry C.; Wilson, Mark V. H. (2016). Fishes of the World (5th ed.). Hoboken: John Wiley and Sons. pp. 314–526. doi:10.1002/9781119174844. ISBN 978-1-118-34233-6.
  7. ^ Sibert, E. C.; Norris, R. D. (2015-06-29). "New Age of Fishes initiated by the Cretaceous−Paleogene mass extinction". PNAS. 112 (28): 8537–8542. Bibcode:2015PNAS..112.8537S. doi:10.1073/pnas.1504985112. PMC 4507219. PMID 26124114.
  8. ^ a b Arcila, Dahiana; Alexander Pyron, R.; Tyler, James C.; Ortí, Guillermo; Betancur-R., Ricardo (January 2015). "An evaluation of fossil tip-dating versus node-age calibrations in tetraodontiform fishes (Teleostei: Percomorphaceae)". Molecular Phylogenetics and Evolution. 82: 131–145. doi:10.1016/j.ympev.2014.10.011. PMID 25462998.
  9. ^ a b Friedman, Matt; Andrews, James V.; Saad, Hadeel; El-Sayed, Sanaa (31 August 2023). "The Cretaceous–Paleogene transition in spiny-rayed fishes: surveying 'Patterson's Gap' in the acanthomorph skeletal record". Geologica Belgica. 26 (1–2): 1–23. doi:10.20341/gb.2023.002.
  10. ^ Carnevale, Giorgio; Johnson, G. David (December 2015). "A Cretaceous Cusk-Eel (Teleostei, Ophidiiformes) from Italy and the Mesozoic Diversification of Percomorph Fishes". Copeia. 103 (4): 771–791. doi:10.1643/CI-15-236.
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  12. ^ Wong, Man-Kwan; Chen, Wei-Jen (October 2024). "Exploring the phylogeny and depth evolution of cusk eels and their relatives (Ophidiiformes: Ophidioidei)". Molecular Phylogenetics and Evolution. 199 108164. doi:10.1016/j.ympev.2024.108164. PMID 39084413.