Clade of ray-finned fishes
Percomorpha (from Latin perca ' perch ' and Ancient Greek μορφή (morphḗ) ' shape, appearance ' ) is an extremely large and diverseclade ofray-finned fish . With more than 17,000 known species (includingtuna ,seahorses ,gobies ,cichlids ,flatfish ,wrasse ,perches ,anglerfish , andpufferfish ) known from both marine and freshwater ecosystems, it is the most speciose clade of extantvertebrates .[ 1] [ 2] [ 3] [ 4] [ 5] [ 6]
Percomorpha are the mostdiverse group ofteleost fish today. Teleosts, and percomorphs in particular, thrived during theCenozoic era . Fossil evidence shows that there was a major increase in size and abundance of teleosts immediately after themass extinction event at theCretaceous-Paleogene boundary c. 66Ma ago.[ 7] The oldest known percomorph fossils are of the early tetraodontiforms Protriacanthus andCretatriacanthidae from theSantonian toCampanian ofItaly andSlovenia .[ 8] A higher diversity of early percomorphs is also known from theCampanian ofNardò , Italy, and these also show some level of diversification into modern orders, with representatives of theSyngnathiformes and Tetraodontiformes known.[ 9] Possibly the oldest percomorph isPlectocretacicus from theCenomanian ofLebanon , which may be a stem-tetraodontiform; however, some morphological analyses indicate that it shows similarities with non-percomorph groups.[ 8] [ 10]
Evolution of ray-finned fishes,Actinopterygii , from theDevonian 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 expandedPerciformes , 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 toEschmeyer'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 theEocene , and many afterwards, cannot be confidently assigned to extant percomorph lineages.[ 12] In the past they were placed within an expanded Perciformes, as with many extant groups, a treatment that is now known to be paraphyletic.[ 13]
The following taxonomy is based on Eschmeyer's Catalog of Fishes (2025), with additional suborders based on other studies:[ 5] [ 11]
CladePercomorpha SubseriesOphidiida SubseriesBatrachoidida SubseriesGobiida SubseriesSyngnatharia SubseriesPelagiaria OrderScombriformes SuborderStromateoidei –medusafishes ,driftfishes ,butterfishes ,squaretails ,ariommas and allies (6 families) SuborderScombroidei –mackerel ,tunas ,manefishes ,pomfrets ,cutlassfishes ,bluefish ,swallowers ,ragfish and allies (10 families) SubseriesAnabantaria SubseriesCarangaria SubseriesOvalentaria SubseriesEupercaria OrderPerciformes SuborderPercoidei –perches ,darters ,seabasses ,groupers ,anthias ,weeverfishes andduckbills (9 families) SuborderNotothenioidei –icefishes ,toothfishes ,Antarctic dragonfishes and allies (8 families) SuborderScorpaenoidei –scorpionfishes ,stonefishes ,rockfishes ,flatheads ,searobins and allies (11 families) SuborderCottoidei –sculpins ,psychrolutes ,snailfishes ,sablefishes ,greenlings ,poachers ,sandfishes ,lumpfishes and allies (11 families) SuborderGasterosteoidei –sticklebacks ,Korean sandlance andtubesnouts (3 families) SuborderZoarcoidei –eelpouts ,wolffishes ,pricklebacks ,gunnels ,prowfish and allies (14 families) OrderCentrarchiformes SuborderPercalatoidei –Australian basses (1 undescribed family) SuborderTerapontoidei –grunters ,sea chubs ,flagtails ,knifejaws ,stripeys and allies (9 families) SuborderCentrarchoidei –freshwater sunfish ,Chinese perches ,oldwives ,temperate perch ,jutjaws and allies (7 families) SuborderCirrhitioidei –hawkfishes ,kelpfishes ,marblefishes ,fingerfins andtrumpeters (5 families) OrderLabriformes OrderAcropomatiformes (=Pempheriformes) –lanternbellies ,sweepers ,gnomefishes ,wreckfishes ,banjofish ,oceanic basslets ,armorheads and allies (20 families) OrderAcanthuriformes –surgeonfishes ,butterflyfishes ,rabbitfishes ,marine angelfishes ,drumfish ,grunts ,ponyfishes ,mojarras ,snappers ,temperate basses ,spadefishes ,porgies ,moonyfish ,tripletails ,tilefish ,scats ,bigeyes ,boarfishes and allies (30 families) OrderLophiiformes –anglerfish OrderTetraodontiformes External relationships [ edit ] The twocladograms below are based on Betancur-Ret al. , 2017.[ 5] Percomorphs are aclade ofteleost fishes. The first cladogram shows the interrelationships of percomorphs with other living groups of teleosts.
Internal relationships [ edit ] The following cladogram shows theevolutionary relationships of the various groups ofextant percomorph fishes:
^ 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 . ^ 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 . ^ 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 . ^ 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 . ^a b c d e 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 .ISSN 1471-2148 .PMC 5501477 .PMID 28683774 . ^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 . Archived fromthe original on 2019-04-08. Retrieved2019-10-16 . ^ 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 . ^a b Arcila, Dahiana; Alexander Pyron, R.; Tyler, James C.; Ortí, Guillermo; Betancur-R., Ricardo (2015-01-01)."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 .ISSN 1055-7903 .PMID 25462998 . ^ Friedman, Matt; V. Andrews, James; Saad, Hadeel; El-Sayed, Sanaa (2023-06-16)."The Cretaceous–Paleogene transition in spiny-rayed fishes: surveying "Patterson's Gap" in the acanthomorph skeletal record André Dumont medalist lecture 2018" .Geologica Belgica .doi :10.20341/gb.2023.002 .ISSN 1374-8505 . ^ Carnevale, Giorgio; Johnson, G. David (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 .ISSN 0045-8511 . ^a b Fricke, R.; Eschmeyer, W. N.; Van der Laan, R. (2025)."ESCHMEYER'S CATALOG OF FISHES: CLASSIFICATION" .California Academy of Sciences . Retrieved2025-02-10 . ^ Friedman, Matt; V. Andrews, James; Saad, Hadeel; El-Sayed, Sanaa (2023-06-16)."The Cretaceous–Paleogene transition in spiny-rayed fishes: surveying "Patterson's Gap" in the acanthomorph skeletal record André Dumont medalist lecture 2018" .Geologica Belgica .doi :10.20341/gb.2023.002 .ISSN 1374-8505 . ^ 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 .ISSN 1471-2148 .PMC 5501477 .PMID 28683774 . ^ Wong, Man-Kwan; Chen, Wei-Jen (2024-10-01)."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 .ISSN 1055-7903 .
Percomorphaceae Percomorpha