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Superregnum : Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Superclassis/Classis: Actinopterygii
Unranked clades: Palaeoniscimorpha - Teleosteomorpha
Classis/Subclassis: Actinopteri - Cladistia
Subclassis/Infraclassis: Chondrostei - Neopterygii
Superordines: Palaeonisci
Ordines inc. sed.: †Discordichthyiformes - †Docopteri - †Eurynotoidiformes - †Lysopteri - †Merospondyli - †Pachycormiformes - †Palaeonisciformes - †Pycnodontiformes
Genera inc. sed.: †Angatubichthys – †Melanecta – †Paphosiscus – †Pteronisculus – †Raynerius – †Woodichthys

Actinopterygii Klein, 1885
Actinopterygii sensu Goodrich, 1930 (according to DOI: 10.26028/cybium/2020-441-001)


Template:Klein, 1885

Betancur-R., R., Broughton, R.E., Wiley, E.O., Carpenter, K., Andrés López, J., Li, C., Holcroft, N.I., Arcila, D., Sanciangco, M., Cureton II, J.C., Zhang, F., Buser, T., Campbell, M.A., Ballesteros, J.A., Roa-Varon, A., Willis, S., Borden, W.C., Rowley, T., Reneau, P.C., Hough, D.J., Lu, G., Grande, T., Arratia, G. & Ortí, G. 2013. The tree of life and a new classification of bony fishes. (PDF) PLOS Currents Tree of Life 2013 Apr 18: 1–45, downloadable Appendix 2 (new classification): 1–21, and downloadable Figure S1 (complete cladogram with annotated classification). DOI: 10.1371/currents.tol.53ba26640df0ccaee75bb165c8c26288 Reference page.
Do Nascimiento, C., Cárdenas-Bautista, J-S., Borja Acosta, K.G., González-Alvarado, A. & Medina-Uribe, C.A. 2016. Illustrated and online catalog of type specimens of freshwater fishes in the Colección de Peces Dulceacuícolas of Instituto de Investigación de Recursos Biológicos Alexander von Humboldt (IAvH-P), Colombia. Zootaxa 4171(3): 401–438. DOI: 10.11646/zootaxa.4171.3.1. Reference page.
Eder, E.B., Marin, M.R. & Lewis, M.N. 2017. Demersal and pelagic species of fish and squid from the Patagonian shelf. ZooKeys 668: 139—145. Dataset DOI: 10.3897/zookeys.668.11826. Reference page.
Nelson, J.S. 2006: Fishes of the world. 4th edition. John Wiley & Sons Inc., New York. ISBN 0-471-25031-7 ISBN 978-0-471-25031-9 Google books
Roberts, C.D.; Paulin, C.D.; Stewart, A.L.; McPhee, R.P.; McDowall, R.M. (compilers) 2009: Checklist of New Zealand Chordata: living lancelets, jawless fishes, cartilaginous fishes, and bony fishes. Pp. 527-536 in Gordon, D.P. (ed.) New Zealand inventory of biodiversity. Volume 1. Kingdom Animalia. Radiata, Lophotrochozoa, Deuterostomia. Canterbury University Press, Christchurch, New Zealand. ISBN 978-1-877257-72-8
Melo, B.F., Benine, R.C., Britzke, R., Gama, C.S. & Oliveira, C. 2016. An inventory of coastal freshwater fishes from Amapá highlighting the occurrence of eight new records for Brazil. ZooKeys 606: 127–140. DOI: 10.3897/zookeys.606.9297. Reference page.
Miyazaki, Y., Teramura, A. & Senou, S. 2019. Preliminary report on bycatch fish species collected from the Tokyo Submarine Canyon, Japan. Zookeys, 843: 117–128. DOI: 10.3897/zookeys.843.32410 Reference page.
Silva, A.S., Groz, M.P., Leandro, P., Assis, C.A. & Figueira, R. 2017. Ichthyological collection of the Museu Oceanográfico D. Carlos I. ZooKeys 752: 137–148. DOI: 10.3897/zookeys.752.20086. Reference page.
Stringer, G.L. 2016. Late Cretaceous actinopterygians represented by otoliths from the Coon Creek Site in southwest Tennessee. Pp 78–95 In Ehret, D.J., Harrell, T.L., Jr. & Ebersole, S.M. (eds.) 2016. The Paleontology of the Cretaceous Coon Creek Formation. Bulletin Alabama Museum of natural history, Bull. 33, vol. 2: 1-121. Reference page. . Reference page.
Torres-Hernández, E., Palacios-Morales, G., Romero-Gallardo, S., Salazar-Araujo, P., García-Meraz, A., Madrigal-Guridi, Z., Del Moral-Flores, L.F. & Dominguez-Domínguez, O. 2016. Annotated checklist of the coastal ichthyofauna from Michoacán State, Mexico. ZooKeys 606: 99–126. DOI: 10.3897/zookeys.606.9004. Reference page.
Wiley, E.O.; Johnson, G.D.; Dimmick, W.W. 2000: The interrelationships of acanthomorph fishes: a total evidence approach using morphological and molecular data. Biochemical systematics and ecology, 28(4): 319–350. DOI: 10.1016/S0305-1978(99)00069-1

Vernacular names
беларуская: Прамянёвапёрыя рыбы
čeština: Paprskoploutví
Deutsch: Strahlenflosser
English: Ray-finned Fishes
Esperanto: Aktinopterigoj
eesti: Kiiruimsed, Aktinopterüügid
suomi: Viuhkaeväiset kalat
français: Actinoptérygiens
עברית: מקריני סנפיר
hrvatski: Zrakoperke
magyar: Sugarasúszójú halak
日本語: 条鰭綱
한국어: 조기어강
latviešu: Starspures
polski: promieniopłetwe
português: Actinopterígio
русский: Лучепёрые рыбы
svenska: Taggfeniga fiskar
ไทย: ปลาที่มีก้านครีบ
Türkçe: Işınsal yüzgeçliler
українська: Променепері
Tiếng Việt: Lớp Cá vây tia
中文: 辐鳍鱼纲

Actinopterygii /ˌæktɪnɒptəˈrɪdʒiaɪ/ (New Latin actino- ('having rays') + Greek πτέρυξ (ptérux 'wing, fins')), members of which are known as ray-finned fishes, is a clade (traditionally class or subclass) of the bony fishes.[1]

The ray-finned fishes are so-called because their fins are webs of skin supported by bony or horny spines (rays), as opposed to the fleshy, lobed fins that characterize the class Sarcopterygii (lobe-finned fish). These actinopterygian fin rays attach directly to the proximal or basal skeletal elements, the radials, which represent the link or connection between these fins and the internal skeleton (e.g., pelvic and pectoral girdles).

By species count, actinopterygians dominate the vertebrates, and they comprise nearly 99% of the over 30,000 species of fish.[2] They are ubiquitous throughout freshwater and marine environments from the deep sea to the highest mountain streams. Extant species can range in size from Paedocypris, at 8 mm (0.3 in), to the massive ocean sunfish, at 2,300 kg (5,070 lb), and the long-bodied oarfish, at 11 m (36 ft). The vast majority of Actinopterygii (~95%) are teleosts.

Anatomy of a typical ray-finned fish (cichlid)
A: dorsal fin, B: fin rays, C: lateral line, D: kidney, E: swim bladder, F: Weberian apparatus, G: inner ear, H: brain, I: nostrils, L: eye, M: gills, N: heart, O: stomach, P: gall bladder, Q: spleen, R: internal sex organs (ovaries or testes), S: ventral fins, T: spine, U: anal fin, V: tail (caudal fin). Possible other parts not shown: barbels, adipose fin, external genitalia (gonopodium)

Ray-finned fishes occur in many variant forms. The main features of a typical ray-finned fish are shown in the adjacent diagram. The swim bladder is the more derived structure.[3]

Ray-finned fishes have many different types of scales; but all teleosts, the most advanced actinopterygians, have leptoid scales. The outer part of these scales fan out with bony ridges while the inner part is crossed with fibrous connective tissue. Leptoid scales are thinner and more transparent than other types of scales, and lack the hardened enamel or dentine-like layers found in the scales of many other fish. Unlike ganoid scales, which are found in non-teleost actinopterygians, new scales are added in concentric layers as the fish grows.[4]

Ray-finned and lobe-finned fishes, including tetrapods, possessed lungs used for aerial respiration. Only bichirs retain ventrally budding lungs.[3]
Body shapes and fin arrangements
Further information: Fish fin
Further information: Diversity of fish

Ray-finned fish vary in size and shape, in their feeding specializations, and in the number and arrangement of their ray-fins.

Tuna are streamlined for straight line speed with a deeply forked tail

The swordfish is even faster and more streamlined than the tuna

Salmon generate enough thrust with their powerful tail fin to jump obstacles during river migrations

Cod have three dorsal and two anal fins, which give them great maneuverability

Flatfish have developed partially symmetric dorsal and pelvic fins


Elongated bristlemouth

Fangtooth are indifferent swimmers who try to ambush their prey

The first spine of the dorsal fin of anglerfish is modified like a fishing rod with a lure


King of herrings

European conger are ray-finned fish

Hawaiian turkeyfish

The benthic batfish Ogcocephalus notatus

The deep sea eel Saccopharynx ampullaceus

The freshwater elephant fish Campylomormyrus curvirostris

The sturgeon Acipenser oxyrhynchus has a cartilaginous endoskeleton

The ambush predator needlefish Belone belone


Mirror dory


The "flying fish" Exocoetus obtusirostris has specialized pectoral fins for gliding

The hoodwinker sunfish Mola tecta has no caudal fin

The Jurassic †Leedsichthys was a giant filter-feeder

Three-spined stickleback males (red belly) build nests and compete to attract females to lay eggs in them. Males then defend and fan the eggs. Painting by Alexander Francis Lydon, 1879

In nearly all ray-finned fish, the sexes are separate, and in most species the females spawn eggs that are fertilized externally, typically with the male inseminating the eggs after they are laid. Development then proceeds with a free-swimming larval stage.[5] However other patterns of ontogeny exist, with one of the commonest being sequential hermaphroditism. In most cases this involves protogyny, fish starting life as females and converting to males at some stage, triggered by some internal or external factor. Protandry, where a fish converts from male to female, is much less common than protogyny.[6]

Most families use external rather than internal fertilization.[7] Of the oviparous teleosts, most (79%) do not provide parental care.[8] Viviparity, ovoviviparity, or some form of parental care for eggs, whether by the male, the female, or both parents is seen in a significant fraction (21%) of the 422 teleost families; no care is likely the ancestral condition.[8] The oldest case of viviparity in ray-finned fish is found in Middle Triassic species of †Saurichthys.[9] Viviparity is relatively rare and is found in about 6% of living teleost species; male care is far more common than female care.[8][10] Male territoriality "preadapts" a species for evolving male parental care.[11][12]

There are a few examples of fish that self-fertilise. The mangrove rivulus is an amphibious, simultaneous hermaphrodite, producing both eggs and spawn and having internal fertilisation. This mode of reproduction may be related to the fish's habit of spending long periods out of water in the mangrove forests it inhabits. Males are occasionally produced at temperatures below 19 °C (66 °F) and can fertilise eggs that are then spawned by the female. This maintains genetic variability in a species that is otherwise highly inbred.[13]
Fossil record
Evolution of ray-finned fish.png
See also: Evolution of fish

The earliest known fossil actinopterygian is Andreolepis hedei, dating back 420 million years (Late Silurian). Remains have been found in Russia, Sweden, and Estonia.[14]

Actinopterygii is divided into the classes Cladistia and Actinopteri. The latter comprised subclasses Chondrostei and Neopterygii. The Neopterygii, in turn, is divided into the infraclasses Holostei and Teleostei. During the Mesozoic and Cenozoic the teleosts in particular diversified widely, and as a result, 96% of all known fish species are teleosts. The cladogram shows the major groups of actinopterygians and their relationship to the terrestrial vertebrates (tetrapods) that evolved from a related group of fish.[15][16][17] Approximate dates are from Near et al., 2012.[15]


Coelacanths, Lungfish Coelacanth flipped.png


Amphibians Deutschlands Amphibien und Reptilien (Salamandra salamdra).jpg


Mammals Phylogenetic tree of marsupials derived from retroposon data (Paucituberculata).png

Sauropsids (reptiles, birds) Zoology of Egypt (1898) (Varanus griseus).png


Polypteriformes (bichirs, reedfishes) Cuvier-105-Polyptère.jpg


Acipenseriformes (sturgeons, paddlefishes) Atlantic sturgeon flipped.jpg


Lepisosteiformes (gars) Longnose gar flipped.jpg

Amiiformes (bowfins) Amia calva 1908 flipped.jpg

275 mya

Teleostei Cyprinus carpio3.jpg

The polypterids (bichirs and reedfish) are the sister lineage of all other actinopterygians, the Acipenseriformes (sturgeons and paddlefishes) are the sister lineage of Neopterygii, and Holostei (bowfin and gars) are the sister lineage of teleosts. The Elopomorpha (eels and tarpons) appear to be the most basal teleosts.[15]

Chondrostei Atlantic sturgeon flipped.jpg
Atlantic sturgeon
Chondrostei (cartilage bone) is a subclass of primarily cartilaginous fish showing some ossification. Earlier definitions of Chondrostei are now known to be paraphyletic, meaning that this subclass does not contain all the descendants of their common ancestor. There were 52 species divided among two orders, the Acipenseriformes (sturgeons and paddlefishes) and the Polypteriformes (reedfishes and bichirs). Reedfish and birchirs are now separated from the Chondrostei into their own sister lineage, the Cladistia. It is thought that the chondrosteans evolved from bony fish but lost the bony hardening of their cartilaginous skeletons, resulting in a lightening of the frame. Elderly chondrosteans show beginnings of ossification of the skeleton, suggesting that this process is delayed rather than lost in these fish.[18] This group had once been classified with the sharks: the similarities are obvious, as not only do the chondrosteans mostly lack bone, but the structure of the jaw is more akin to that of sharks than other bony fish, and both lack scales (excluding the Polypteriforms). Additional shared features include spiracles and, in sturgeons, a heterocercal tail (the vertebrae extend into the larger lobe of the caudal fin). However the fossil record suggests that these fish have more in common with the Teleostei than their external appearance might suggest.[18]
Neopterygii Salmo salar flipped.jpg
Atlantic salmon
Neopterygii (new fins) is a subclass of ray-finned fish that appeared somewhere in the Late Permian. There were only few changes during its evolution from the earlier actinopterygians. Neopterygians are a very successful group of fishes because they can move more rapidly than their ancestors. Their scales and skeletons began to lighten during their evolution, and their jaws became more powerful and efficient. While electroreception and the ampullae of Lorenzini is present in all other groups of fish, with the exception of hagfish, neopterygians have lost this sense, though it later re-evolved within Gymnotiformes and catfishes, who possess nonhomologous teleost ampullae.[19]

The listing below follows Phylogenetic Classification of Bony Fishes[16][20] with notes when this differs from Nelson,[21] ITIS[22] and FishBase[23] and extinct groups from Van der Laan 2016 and Xu 2021.[24][25]


Kardong, Kenneth (2015). Vertebrates: Comparative Anatomy, Function, Evolution. New York: McGraw-Hill Education. pp. 99–100. ISBN 978-0-07-802302-6.
(Davis, Brian 2010).
Funk, Emily; Breen, Catriona; Sanketi, Bhargav; Kurpios, Natasza; McCune, Amy (2020). "Changing in Nkx2.1, Sox2, Bmp4, and Bmp16 expression underlying the lung-to-gas bladder evolutionary transition in ray-finned fishes". Evolution & Development. 22 (5): 384–402. doi:10.1111/ede.12354. PMC 8013215. PMID 33463017.
"Actinopterygii Klein, 1885". Retrieved 20 September 2021.
Dorit, R.L.; Walker, W.F.; Barnes, R.D. (1991). Zoology. Saunders College Publishing. p. 819. ISBN 978-0-03-030504-7.
Avise, J.C.; Mank, J.E. (2009). "Evolutionary perspectives on hermaphroditism in fishes". Sexual Development. 3 (2–3): 152–163. doi:10.1159/000223079. PMID 19684459. S2CID 22712745.
Pitcher, T (1993). The Behavior of Teleost Fishes. London: Chapman & Hall.
Reynolds, John; Nicholas B. Goodwin; Robert P. Freckleton (19 March 2002). "Evolutionary Transitions in Parental Care and Live Bearing in Vertebrates". Philosophical Transactions of the Royal Society B: Biological Sciences. 357 (1419): 269–281. doi:10.1098/rstb.2001.0930. PMC 1692951. PMID 11958696.
Maxwell; et al. (2018). "Re‐evaluation of the ontogeny and reproductive biology of the Triassic fish Saurichthys (Actinopterygii, Saurichthyidae)". Palaeontology. 61: 559–574. doi:10.5061/dryad.vc8h5.
Clutton-Brock, T. H. (1991). The Evolution of Parental Care. Princeton, NJ: Princeton UP.
Werren, John; Mart R. Gross; Richard Shine (1980). "Paternity and the evolution of male parentage". Journal of Theoretical Biology. 82 (4): 619–631. doi:10.1016/0022-5193(80)90182-4. PMID 7382520. Retrieved 15 September 2013.
Baylis, Jeffrey (1981). "The Evolution of Parental Care in Fishes, with reference to Darwin's rule of male sexual selection". Environmental Biology of Fishes. 6 (2): 223–251. doi:10.1007/BF00002788. S2CID 19242013.
Wootton, Robert J.; Smith, Carl (2014). Reproductive Biology of Teleost Fishes. Wiley. ISBN 978-1-118-89139-1.
"Fossilworks: Andreolepis". Archived from the original on 12 February 2010. Retrieved 14 May 2008.
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. PMC 3644299. PMID 23653398. Archived from the original on 13 October 2013.
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.
"Chondrosteans: Sturgeon Relatives". Archived from the original on 25 December 2010.
Theodore Holmes Bullock; Carl D. Hopkins; Arthur N. Popper (2005). Electroreception. Springer Science+Business Media, Incorporated. p. 229. ISBN 978-0-387-28275-6.
Betancur-Rodriguez; et al. (2017). "Phylogenetic Classification of Bony Fishes Version 4". BMC Evolutionary Biology. 17 (1): 162. doi:10.1186/s12862-017-0958-3. PMC 5501477. PMID 28683774.
Nelson, Joseph, S. (2016). Fishes of the World. John Wiley & Sons, Inc. ISBN 978-1-118-34233-6.
"Actinopterygii". Integrated Taxonomic Information System. Retrieved 3 April 2006.
R. Froese and D. Pauly, editors (February 2006). "FishBase". Archived from the original on 5 July 2018. Retrieved 8 January 2020.
Van der Laan, Richard (2016). Family-group names of fossil fishes. doi:10.13140/RG.2.1.2130.1361.
Xu, Guang-Hui (9 January 2021). "A new stem-neopterygian fish from the Middle Triassic (Anisian) of Yunnan, China, with a reassessment of the relationships of early neopterygian clades". Zoological Journal of the Linnean Society. 191 (2): 375–394. doi:10.1093/zoolinnean/zlaa053. ISSN 0024-4082.
In Nelson, Polypteriformes is placed in its own subclass Cladistia.
In Nelson and ITIS, Syngnathiformes is placed as the suborder Syngnathoidei of the order Gasterosteiformes.

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