Actinopterygii Klein, 1885

Actinopterygii (; ), members of which are known as ray-finned fishes, is a clade (traditionally class or subclass) of the bony fishes. They comprise over 50% of living vertebrate species. 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.(Davis, Brian 2010). 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 , to the massive ocean sunfish, at , and the long-bodied oarfish, at . The vast majority of Actinopterygii (~99%) are teleosts.

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

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. 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. Ray-finned and lobe-finned fishes, including tetrapods, possessed lungs used for aerial respiration. Only bichirs retain ventrally budding lungs.

Actinopterygii is divided into the classes Cladistia and Actinopteri. The latter comprises the subclasses Chondrostei and Neopterygii. The Neopterygii, in turn, is divided into the infraclasses Holostei and Teleostei. During the Mesozoic (Triassic, Jurassic, Cretaceous) and Cenozoic the teleosts in particular diversified widely. As a result, 96% of living fish species are teleosts (40% of all fish species belong to the teleost subgroup Acanthomorpha), while all other groups of actinopterygians represent depauperate lineages. The classification of ray-finned fishes can be summarized as follows:

Cladistia, which include bichirs and reedfish Actinopteri, which include: Chondrostei, which include Acipenseriformes (paddlefishes and sturgeons) Neopterygii, which include: Teleostei (most living fishes) Holostei, which include: Lepisosteiformes (gars) Amiiformes (bowfin)

The cladogram below shows the main clades of living actinopterygians and their evolutionary relationships to other extant groups of fishes and the four-limbed vertebrates (tetrapods). The latter include mostly terrestrial species but also groups that became secondarily aquatic (e.g. Whales and Dolphins). Tetrapods evolved from a group of bony fish during the Devonian period. Approximate divergence dates for the different actinopterygian clades (in millions of years, mya) are from Near et al., 2012.

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. The earliest known fossil actinopterygian is Andreolepis hedei, dating back 420 million years (Late Silurian). Remains have been found in Russia, Sweden, and Estonia. The earliest fossil relatives of modern teleosts are from the Triassic period (Prohalecites, Pholidophorus), although it is suspected that teleosts originated already during the Paleozoic Era.

Chondrostei 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. 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. Neopterygii 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.

Fossil of the Devonian cheirolepidiform Cheirolepis canadensis

Fossil of the Carboniferous elonichthyiform Elonichthys peltigerus

Fossil of the Permian aeduelliform Aeduella blainvillei

Fossil of the Permian palaeonisciform Palaeoniscum freieslebeni

Fossil of the Triassic bobasatraniiform Bobasatrania canadensis

Fossil of the Triassic perleidiform Thoracopterus magnificus

Fossils of the Triassic prohaleciteiform Prohalecites sp., the earliest teleosteomorph

Fossil of the Jurassic aspidorhynchiform Aspidorhynchus sp.

Fossil of the Jurassic pachycormiform Pachycormus curtus

Fossil of the Cretaceous acipenseriform Yanosteus longidorsalis

Fossil of the Cretaceous aulopiform Nematonotus longispinus

Fossil of the Cretaceous ichthyodectiform Thrissops formosus

Fossil of the Cretaceous carangiform Mene oblonga

Fossil of the Cretaceous pleuronectiform Amphistium paradoxum

Fossil of a ray-finned perch (Priscacara serrata) from the Lower Eocene about 50 million years ago

Fossil of the Miocene syngnathiform Nerophis zapfei

Skeleton of the angler fish, Lophius piscatorius. The first spine of the dorsal fin of the anglerfish is modified so it functions like a fishing rod with a lure

Skeleton of another ray-finned fish, the lingcod



Taxonomy The listing below is a summary of all extinct (indicated by a dagger, †) and living groups of Actinopterygii with their respcective taxonomic rank. The taxonomy follows Phylogenetic Classification of Bony Fishes with notes when this differs from Nelson, ITIS and FishBase and extinct groups from Van der Laan 2016 and Xu 2021.

Order †?Asarotiformes Schaeffer 1968 Order †?Discordichthyiformes Minikh 1998 Order †?Paphosisciformes Grogan & Lund 2015 Order †?Scanilepiformes Selezneya 1985 Order †Cheirolepidiformes Kazantseva-Selezneva 1977 Order †Paramblypteriformes Heyler 1969 Order †Rhadinichthyiformes Order †Palaeonisciformes Hay 1902 Order †Tarrasiiformes sensu Lund & Poplin 2002 Order †Ptycholepiformes Andrews et al. 1967 Order †Haplolepidiformes Westoll 1944 Order †Aeduelliformes Heyler 1969 Order †Platysomiformes Aldinger 1937 Order †Dorypteriformes Cope 1871 Order †Eurynotiformes Sallan & Coates 2013 Class Cladistia Pander 1860 Order †Guildayichthyiformes Lund 2000 Order Polypteriformes Bleeker 1859 (bichirs and reedfishes)In Nelson, Polypteriformes is placed in its own subclass Cladistia. Class Actinopteri Cope 1972 s.s. Order †Elonichthyiformes Kazantseva-Selezneva 1977 Order †Phanerorhynchiformes Order †Bobasatraniiformes Berg 1940 Order †Saurichthyiformes Aldinger 1937 Subclass Chondrostei Müller, 1844 Order †Birgeriiformes Heyler 1969 Order †Chondrosteiformes Aldinger, 1937 Order Acipenseriformes Berg 1940 (includes sturgeons and paddlefishes) Subclass Neopterygii Regan 1923 sensu Xu & Wu 2012 Order †Pholidopleuriformes Berg 1937 Order †Redfieldiiformes Berg 1940 Order †Platysiagiformes Brough 1939 Order †Polzbergiiformes Griffith 1977 Order †Perleidiformes Berg 1937 Order †Louwoichthyiformes Xu 2021 Order †Peltopleuriformes Lehman 1966 Order †Luganoiiformes Lehman 1958 Order †Pycnodontiformes Berg 1937 Infraclass Holostei Müller 1844 Division Halecomorphi Cope 1872 sensu Grande & Bemis 1998 Order †Parasemionotiformes Lehman 1966 Order †Ionoscopiformes Grande & Bemis 1998 Order Amiiformes Huxley 1861 sensu Grande & Bemis 1998 (bowfins) Division Ginglymodi Cope 1871 Order †Dapediiformes Thies & Waschkewitz 2015 Order †Semionotiformes Arambourg & Bertin 1958 Order Lepisosteiformes Hay 1929 (gars) Clade Teleosteomorpha Arratia 2000 sensu Arratia 2013 Order †Prohaleciteiformes Arratia 2017 Division Aspidorhynchei Nelson, Grand & Wilson 2016 Order †Aspidorhynchiformes Bleeker 1859 Order †Pachycormiformes Berg 1937 Infraclass Teleostei Müller 1844 sensu Arratia 2013 Order †?Araripichthyiformes Order †?Ligulelliiformes Taverne 2011 Order †?Tselfatiiformes Nelson 1994 Order †Pholidophoriformes Berg 1940 Order †Dorsetichthyiformes Nelson, Grand & Wilson 2016 Order †Leptolepidiformes Order †Crossognathiformes Taverne 1989 Order †Ichthyodectiformes Bardeck & Sprinkle 1969 Teleocephala de Pinna 1996 s.s. Megacohort Elopocephalai Patterson 1977 sensu Arratia 1999 (Elopomorpha Greenwood et al. 1966) Order Elopiformes Gosline 1960 (ladyfishes and tarpon) Order Albuliformes Greenwood et al. 1966 sensu Forey et al. 1996 (bonefishes) Order Notacanthiformes Goodrich 1909 (halosaurs and spiny eels) Order Anguilliformes Jarocki 1822 sensu Goodrich 1909 (true eels) Megacohort Osteoglossocephalai sensu Arratia 1999 Supercohort Osteoglossocephala sensu Arratia 1999 (Osteoglossomorpha Greenwood et al. 1966) Order †Lycopteriformes Chang & Chou 1977 Order Hiodontiformes McAllister 1968 sensu Taverne 1979 (mooneye and goldeye) Order Osteoglossiformes Regan 1909 sensu Zhang 2004 (bony-tongued fishes) Supercohort Clupeocephala Patterson & Rosen 1977 sensu Arratia 2010 Cohort Otomorpha Wiley & Johnson 2010 (Otocephala; Ostarioclupeomorpha) Subcohort Clupei Wiley & Johnson 2010 (Clupeomorpha Greenwood et al. 1966) Order †Ellimmichthyiformes Grande 1982 Order Clupeiformes Bleeker 1859 (herrings and anchovies) Subcohort Alepocephali Order Alepocephaliformes Marshall 1962 Subcohort Ostariophysi Sagemehl 1885 Section Anotophysa (Rosen & Greenwood 1970) Sagemehl 1885 Order †Sorbininardiformes Taverne 1999 Order Gonorynchiformes Regan 1909 (milkfishes) Section Otophysa Garstang 1931 Order Cypriniformes Bleeker 1859 sensu Goodrich 1909 (barbs, carp, danios, goldfishes, loaches, minnows, rasboras) Order Characiformes Goodrich 1909 (characins, pencilfishes, hatchetfishes, piranhas, tetras, dourado / golden (genus Salminus) and pacu) Order Gymnotiformes Berg 1940 (electric eels and knifefishes) Order Siluriformes Cuvier 1817 sensu Hay 1929 (catfishes) Cohort Euteleosteomorpha (Greenwood et al. 1966) (Euteleostei Greenwood 1967 sensu Johnson & Patterson 1996) Subcohort Lepidogalaxii Lepidogalaxiiformes Betancur-Rodriguez et al. 2013 (salamanderfish) Subcohort Protacanthopterygii Greenwood et al. 1966 sensu Johnson & Patterson 1996 Order Argentiniformes (barreleyes and slickheads) (formerly in Osmeriformes) Order Galaxiiformes Order Salmoniformes Bleeker 1859 sensu Nelson 1994 (salmon and trout) Order Esociformes Bleeker 1859 (pike) Subcohort Stomiati Order Osmeriformes (smelts) Order Stomiatiformes Regan 1909 (bristlemouths and marine hatchetfishes) Subcohort Neoteleostei Nelson 1969 Infracohort Ateleopodia Order Ateleopodiformes (jellynose fish) Infracohort Eurypterygia Rosen 1973 Section Aulopa [Cyclosquamata Rosen 1973] Order Aulopiformes Rosen 1973 (Bombay duck and lancetfishes) Section Ctenosquamata Rosen 1973 Subsection Myctophata [Scopelomorpha] Order Myctophiformes Regan 1911 (lanternfishes) Subsection Acanthomorpha Betancur-Rodriguez et al. 2013 Division Lampridacea Betancur-Rodriguez et al. 2013 [Lampridomorpha; Lampripterygii] Order Lampriformes Regan 1909 (oarfish, opah and ribbonfishes) Division Paracanthomorphacea sensu Grande et al. 2013 (Paracanthopterygii Greenwood 1937) Order Percopsiformes Berg 1937 (cavefishes and trout-perches) Order †Sphenocephaliformes Rosen & Patterson 1969 Order Zeiformes Regan 1909 (dories) Order Stylephoriformes Miya et al. 2007 Order Gadiformes Goodrich 1909 (cods) Division Polymixiacea Betancur-Rodriguez et al. 2013 (Polymyxiomorpha; Polymixiipterygii) Order †Pattersonichthyiformes Gaudant 1976 Order †Ctenothrissiformes Berg 1937 Order Polymixiiformes Lowe 1838 (beardfishes) Division Euacanthomorphacea Betancur-Rodriguez et al. 2013 (Euacanthomorpha sensu Johnson & Patterson 1993; Acanthopterygii Gouan 1770 sensu]) Subdivision Berycimorphaceae Betancur-Rodriguez et al. 2013 Order Beryciformes (fangtooths and pineconefishes) (incl. Stephanoberyciformes; Cetomimiformes) Subdivision Holocentrimorphaceae Betancur-Rodriguez et al. 2013 Order Holocentriformes (Soldierfishes) Subdivision Percomorphaceae Betancur-Rodriguez et al. 2013 (Percomorpha sensu Miya et al. 2003; Acanthopteri) Series Ophidiimopharia Betancur-Rodriguez et al. 2013 Order Ophidiiformes (pearlfishes) Series Batrachoidimopharia Betancur-Rodriguez et al. 2013 Order Batrachoidiformes (toadfishes) Series Gobiomopharia Betancur-Rodriguez et al. 2013 Order Kurtiformes(Nurseryfishes and cardinalfishes) Order Gobiiformes(Sleepers and gobies) Series Scombrimopharia Betancur-Rodriguez et al. 2013 Order Syngnathiformes (seahorses, pipefishes, sea moths, cornetfishes and flying gurnardsIn Nelson and ITIS, Syngnathiformes is placed as the suborder Syngnathoidei of the order Gasterosteiformes.) Order Scombriformes (Tunas and (mackerels) Series Carangimopharia Betancur-Rodriguez et al. 2013 Subseries Anabantaria Betancur-Rodriguez et al. 2014 Order Synbranchiformes (swamp eels) Order Anabantiformes (Labyrinthici) (gouramies, snakeheads, ) Subseries Carangaria Betancur-Rodriguez et al. 2014 Carangaria incertae sedis Order Istiophoriformes Betancur-Rodriguez 2013 (Marlins, swordfishes, billfishes) Order Carangiformes (Jack mackerels, pompanos) Order Pleuronectiformes Bleeker 1859 (flatfishes) Subseries Ovalentaria Smith & Near 2012 (Stiassnyiformes sensu Li et al. 2009) Ovalentaria incertae sedis Order Cichliformes Betancur-Rodriguez et al. 2013 (Cichlids, Convict blenny, leaf fishes) Order Atheriniformes Rosen 1964 (silversides and rainbowfishes) Order Cyprinodontiformes Berg 1940 (livebearers, killifishes) Order Beloniformes Berg 1940 (flyingfishes and ricefishes) Order Mugiliformes Berg 1940 (mullets) Order Blenniiformes Springer 1993 (Blennies) Order Gobiesociformes Gill 1872 (Clingfishes) Series Eupercaria Betancur-Rodriguez et al. 2014 (Percomorpharia Betancur-Rodriguez et al. 2013) Eupercaria incertae sedis Order Gerreiformes (Mojarras) Order Labriformes (Wrasses and Parrotfishes) Order Caproiformes (Boarfishes) Order Lophiiformes Garman 1899 (Anglerfishes) Order Tetraodontiformes Regan 1929 (Filefishes and pufferfish) Order Centrarchiformes Bleeker 1859 (Sunfishes and mandarin fishes) Order Gasterosteiformes (Sticklebacks and relatives) Order Scorpaeniformes (Lionfishes and relatives) Order Perciformes Bleeker 1859

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. 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. Most families use external rather than internal fertilization. Of the oviparous teleosts, most (79%) do not provide parental care. 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. The oldest case of viviparity in ray-finned fish is found in Middle Triassic species of Saurichthys. Viviparity is relatively rare and is found in about 6% of living teleost species; male care is far more common than female care. Male territoriality "preadapts" a species for evolving male parental care. 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 and can fertilise eggs that are then spawned by the female. This maintains genetic variability in a species that is otherwise highly inbred.