Actinopterygii

Etymology and Definition

The scientific name Actinopterygii derives from Ancient Greek aktis, meaning ray or beam, and pteryx or pterygion, meaning wing or fin.^[1] This designation refers directly to the anatomical construction of their paired and median fins, in which flexible membranes of skin are supported and stiffened by radiating rays.^[1] Ray-finned fishes form one of the two primary evolutionary lineages of bony fishes (Osteichthyes), existing as the sister group to the lobe-finned fishes (Sarcopterygii), which also gave rise to terrestrial tetrapods.^[1][2][3]

By species count, ray-finned fishes make up over half of all living vertebrate species.^[4] Another count puts them at nearly half of all vertebrate species..^[3][4] Living species range from the miniature cyprinid Paedocypris progenetica, which can mature at less than 8 mm in length, to massive ocean sunfishes (Mola) weighing up to 2,300 kg or 2,700 kg, and the giant oarfish (Regalecus glesne), which reaches lengths of 8 to 11 m.^{[1][5][6][7][8]} The largest ray-finned fish ever known was the extinct filter-feeding Leedsichthys problematicus from the Jurassic, which reached estimated lengths of 16 to 16.5 m.^[1][9][10]

Characteristics and Anatomy

Actinopterygian fishes share a collection of distinct morphological synapomorphies that separate them from other gnathostomes.^[2][3] The fins are supported by dermal lepidotrichia that articulate with skeletal basal elements situated predominantly within the body wall, leaving the external fin membrane light, thin, and highly mobile.^[2] Unlike sarcopterygians, ray-finned fishes possess teeth tipped with a specialized hypermineralized cap known as acrodin, lack internal nostrils (choanae), exhibit an everted forebrain, and possess a unique secondary vascular network.^[2][11][12]

Fins and Locomotor Mechanics

The fin web of actinopterygians is supported by segmented, flexible soft rays or unsegmented rigid spines, termed lepidotrichia, which develop through dermal ossification of connective tissue.^[2][13][14] The proximal skeletal elements that articulate these rays with the pectoral and pelvic girdles are termed radials (radialia).^[15][16] In modern teleosts, cartilaginous basal elements (basalia) are lost, and the radials in the pectoral girdle are reduced to a simple vertical row of typically four bones, whereas in the pelvic fins radials may be absent, with the lepidotrichia articulating directly onto the pelvic girdle (basipterygium).^{[2][13][15][17]} In contrast, basal actinopterygians retain more extensive radials; bichirs (Polypteridae) retain elongated radials articulating through two basalia surrounding a partially ossified cartilaginous plate, along with substantial muscle extending into the fleshy fin base.^{[13][14][18][19]}

Because intrinsic fin musculature is attached at the skeletal base of each ray, ray-finned fishes can control the abduction, adduction, orientation, and wetted surface area of their fins with high precision, resembling the folding and unfolding of a fan.^[20][21] This arrangement produces superior thrust-to-weight ratios per fin movement compared to the bulkier fleshy fins of sarcopterygian fishes and the rigid fins of chondrichthyans.^[21] Primitive actinopterygians originally possessed a single dorsal fin, whereas ancestral sarcopterygians possessed two; secondary modifications across different teleost lineages have since multiplied, segmented, or lost the dorsal fin.^[2] The caudal fin varies across clades: it is internally and externally symmetrical (diphycercal) in bichirs, asymmetrical with the vertebral column extending into the upper lobe (heterocercal) in sturgeons and early fossil forms, and externally symmetrical with an upturned terminal vertebral column (homocercal) in teleosts.^[2][22]

Integument and Scales

The skin of ray-finned fishes consists of a multi-layered non-keratinizing epidermis resting on a collagenous dermis.^[23] Numerous unicellular epidermal glands produce a protective mucus layer that acts as a physical barrier, contains antibacterial agents, and reduces hydrodynamic drag during swimming.^[23] Chromatophores situated within or directly beneath the dermis generate diverse colour patterns, while specialized cells containing guanine crystals impart a silvery reflective sheen.^[23]

Ancestral actinopterygians possessed heavy rhombic ganoid scales, composed of a basal plate of laminar bone, a middle layer of spongy vascular bone, and an outer coating of multi-layered ganoin, a highly mineralized enamel-like tissue.^{[2][3][11][24]} These ganoid scales interlocked through an articulating peg-and-socket system.^[3] Today ganoid scales are found only in basal lineages of ray-finned fishes, namely bichirs, sturgeons and gars (Lepisosteidae), while the scales of bowfins have practically no ganoin.^[3][22] During the evolution of modern teleosts, the heavy mineral layers were eliminated, giving rise to thin, transparent, and flexible elasmoid (leptoid) scales composed of uncalcified laminar bone and fibrous connective tissue.^[2][24][25] Elasmoid scales grow by adding concentric layers as the fish grows and occur as either smooth-edged cycloid scales or spine-edged ctenoid scales.^[23][24] Some actinopterygian lineages have replaced scales with bony plates or have lost scale coverings completely.^[26][27]

Skeleton and Cranial Morphology

The endoskeleton of actinopterygians is predominantly ossified, although cartilaginous skeletons with delayed or reduced ossification occur secondarily in chondrosteans such as sturgeons and paddlefishes.^[23][28] Ray-finned fishes exhibit the greatest number of dermal bones covering the skull among all vertebrates.^[2] The neurocranium is formed from both chondral bones (basioccipital, exoccipitals, supraoccipital, sphenoids, and otic bones) and dermal roofing bones (frontals, parietals, parasphenoid, and vomer).^[2] The jaws exhibit high mobility: the maxilla is freed from the cheek, and in teleosts the premaxilla forms the upper margin of the mouth and can protrude forward via specialized ligaments, enabling suction feeding.^[21][26]

The vertebral column consists of amphicoelous (biconcave) vertebrae that constrict or replace the embryonic notochord, though sturgeons lack vertebral centra and retain an unrestricted notochord.^[2][29] Each vertebra gives rise dorsally to neural arches that fuse to form the neural canal enclosing the spinal cord and terminate in a neural spine.^[2][29] Ventrally, abdominal vertebrae articulate with ribs, whereas caudal vertebrae form haemal arches that fuse to encase the caudal artery and vein, extending into ventral haemal spines.^[2][29] In many teleosts, intermuscular bones (epipleurals and epineurals) form by ossification of the myosepta.^[30] The pectoral girdle is anchored to the back of the skull through the posttemporal bone, which attaches to the cleithrum, supracleithrum, scapula, and coracoid.^[30]

The branchial skeleton comprises five pairs of branchial arches.^[2][23] The first four arches typically bear two rows of vascularized gill filaments externally and gill rakers internally.^[2][23] Gill rakers prevent mechanical damage to the delicate respiratory surfaces and aid in prey retention or filtration.^[2] The fifth branchial arch typically lacks respiratory filaments and is modified to process food, often bearing robust pharyngeal teeth.^[2][23] The entire branchial chamber is protected externally by a single pair of bony gill covers, the opercula, supported by branchiostegal rays.^[2][23] Teeth are widely distributed, occurring not only on the premaxillae and dentaries, but also on the vomer, palatines, pterygoids, and pharyngeal bones.^[2][23] At the tip of each tooth crown sits an acrodin cap composed of hard enameloid.^[2][11][12]

Nervous System and Brain Eversion

The actinopterygian brain is organized into five primary divisions: telencephalon (forebrain), diencephalon, mesencephalon (midbrain), cerebellum, and myelencephalon (medulla oblongata).^[2] The telencephalon develops through a unique structural process known as eversion.^[2][31][32] While in other vertebrates the dorsal walls of the embryonic telencephalic hemispheres invaginate inward to enclose two lateral ventricles, in actinopterygians the lateral walls bend outward and down, resulting in an unevaginated, solid neural mass with a single T-shaped ventricle situated dorsally and covered by a non-neural epithelial roof.^[2][31][32] The mesencephalon (midbrain) and the cerebellum are well developed.^[2]

Circulatory and Vascular Architecture

The actinopterygian circulatory system is closed and driven by a single-circuit, venous heart containing one atrium and one ventricle.^[2][33] Deoxygenated blood returns from the body tissues into the sinus venosus, moves into the muscular atrium, fills the thick-walled ventricle, and is pumped into the ventral aorta toward the branchial arches.^[2][33] In basal actinopterygians (bichirs, sturgeons, gars), the outflow tract consists of a contractile conus arteriosus containing multiple rows of valves, whereas in teleosts the conus is reduced and replaced by a non-contractile, elastic bulb, the bulbus arteriosus, formed from vascular smooth muscle and connective tissue.^[2] Blood undergoes oxygenation across the capillary beds of the four pairs of gill arches before flowing into the dorsal aorta for systemic distribution.^[2] Red blood cells (erythrocytes) are nucleated.^[34]

Actinopterygians possess a specialized secondary vascular system in addition to the standard primary arterial and venous circuits.^[2][35] This secondary system is composed of fine capillaries and vessels connected to the primary arteries via narrow, coiled interarterial anastomoses measuring 7 to 25 micrometres in diameter.^[2][35] Cells at the entrances of these anastomoses feature elongated cytoplasmic projections.^[35] The secondary system receives blood that has a much lower hematocrit (erythrocyte concentration) than primary blood and is concentrated in specific areas of the head, oral mucosa, and axial musculature, where it may participate in volume regulation, cutaneous respiration, or regional fluid balance.^[2][35]

Respiration and the Swim Bladder

Aquatic respiration takes place primarily through the branchial apparatus.^[2][33] Water is pulled into the mouth by expansion of the buccal and opercular cavities and is subsequently forced outward over the gill filaments when the mouth closes and the opercula contract.^[33] Secondary gas exchange mechanisms occur across the skin in many species, across specialized vascularized areas of the digestive tract, or through suprabranchial organs like the labyrinth organ found in anabantoids.^[2][33]

The swim bladder is a derived hydrostatic organ that enables actinopterygians to maintain neutral buoyancy at variable water depths.^{[2][33][36][37]} Except in bichirs, which retain the ancestral vertebrate condition of paired ventral lung sacs budding from the foregut, the actinopterygian swim bladder develops as an unpaired dorsal outpocketing of the digestive tract.^[2][36][37] In basal lineages such as bowfins and gars, the vascularized swim bladder can function as an air-breathing organ, a capacity secondarily evolved in some teleosts like the arapaima (Arapaima gigas), while in bottom-dwelling or deep-sea lineages it has been lost completely.^{[36][38][39][40]} In physostomous fishes, the bladder remains permanently connected to the gut through a pneumatic duct (ductus pneumaticus), allowing gas to be swallowed or expelled directly; in physoclistous teleosts, the duct is closed, and gas volume is regulated entirely by a specialized vascular gas gland and oval window.^[2][33]

Digestion, Osmoregulation, and Endocrine Function

The digestive tract begins at the oral cavity, which opens into an indistinct pharynx followed by an esophagus.^[2][23] A distinct muscular stomach is present in predatory species, but may be absent or rudimentary in others.^[2][23] In non-teleost actinopterygians, surface area in the intestine is increased by an internal spiral valve, whereas in teleosts the spiral valve is lost and the intestine is elongated, frequently bearing blind tubular diverticula called pyloric caeca near the gastroduodenal junction.^[2][23] Ray-finned fishes have a liver with a gallbladder, and their pancreas is made up of separate small islets. Unlike sarcopterygians and chondrichthyans, modern teleosts lack a cloaca; the anus opens independently, in front of an urogenital papilla carrying separate urinary and reproductive openings.^[2][26][41]

The kidneys are ribbon-shaped and lie near the vertebral column. In freshwater ray-finned fishes, water continuously diffuses into the body while salts are lost; their large kidneys produce large volumes of dilute urine, while specialized ionocytes in the gills actively take up ions.^[23] In marine actinopterygians, which lose water through osmosis, the kidneys are reduced or aglomerular, conserving water by producing minimal volumes of concentrated urine, while excess salts are actively secreted by the gills.^[23] Endocrine regulation involves the pituitary, thyroid, adrenal homologues, endocrine pancreas, and gonads.^[2] In addition, actinopterygians possess unique endocrine tissues: ultimobranchial bodies derived from the posterior branchial epithelium secrete calcitonin, and the urophysis, a neurosecretory swelling at the caudal end of the spinal cord, produces urotensins that regulate osmoregulation and blood pressure.^[2]

Sensory Systems and Acoustic Communication

Ray-finned fishes perceive their surroundings through diverse sensory organs. The eyes lack eyelids and tear glands; accommodation is achieved not by changing the curvature of the spherical lens, but by shifting its distance from the retina via the retractor lentis muscle.^[2][23] Both rods and cones are present in the retina, providing color vision, though cones are reduced or lost in deep-sea taxa.^[29] Olfaction is mediated by paired blind olfactory sacs opening onto the snout via incurrent and excurrent nares that do not penetrate into the mouth.^[2][23] Taste buds occur throughout the oral cavity and pharynx, and in many lineages they extend externally onto barbels, lips, fins, and body skin.^[2][23]

The acoustico-lateralis system includes the inner ear and the lateral line system.^[2][23] The inner ear contains three semicircular canals and three otolithic chambers (sacculus, utriculus, and lagena), each containing a calcium carbonate (aragonite) otolith: the sagitta, asteriscus, and lapillus.^[2][29][42] Otoliths register linear accelerations and gravity, record annual growth increments, and provide morphological characters in taxonomy and paleontology.^[29] In otocephalan fishes, acoustic sensitivity is enhanced by connections between the swim bladder and inner ear; members of the superorder Otophysa possess the Weberian apparatus, an anatomical chain of modified anterior vertebrae and ossicles (claustrum, intercalarium, scaphium, and tripus) that mechanically transmits swim bladder sound vibrations directly to the inner ear.^[2][43] The lateral line system comprises neuromasts located in subdermal canals or surface pits along the trunk and head, detecting water displacement, local currents, and low-frequency vibrations.^[2][23]

Acoustic signalling is widespread among ray-finned fishes. Active sound production used for intraspecific communication and defensive deterrence has been confirmed in 172 out of 470 actinopterygian families.^[44] Fishes produce sounds by stridulating pectoral or cranial bones, grating pharyngeal teeth, or vibrating sonic muscles attached to the walls of the swim bladder.

Bioelectrogenesis

Specialized bioelectric organs capable of generating electric fields have evolved independently in several actinopterygian clades.^[45][46] These organs typically consist of stacks of modified, multinucleated muscle cells (electrocytes) or, in the electric catfish (Malapterurus electricus), converted dermal glandular cells.^[46] Weak electric fields are generated by knifefishes (Gymnotiformes) and elephantfishes (Mormyridae) for navigation and social communication in turbid waters.^[45][46] Strongly electric fishes use high-voltage discharges to stun prey and deter predators.^[45][46] One electric organ of the electric eel (Electrophorus electricus) contains about 6,000 plates, whose voltage can rise above 800 volts, whereas Malapterurus electricus produces about 50 volts, and 100 volts in exceptional cases, with 500 plates.^[46] While ancestral actinopterygians possessed ampullary electroreceptors (ampullae of Lorenzini), these were lost in Neopterygii and subsequently re-evolved independently as non-homologous teleost ampullary organs in Gymnotiformes and Siluriformes.^[47]

Reproduction and Development

In the majority of ray-finned fishes, the sexes are separate (gonochorism), and fertilization is external, occurring when males release milt over eggs spawned into open water or onto substrates.^[2][48] Fertilization takes place through a small opening in the egg membrane, the micropyle.^[2][49] Some teleosts exhibit internal fertilization, utilizing modified fins as intromittent organs, such as the gonopodium derived from the anal fin in Poeciliidae or priapium structures in phallostethids.^[2][50] Several clades undergo sequential hermaphroditism, most frequently manifesting as protogyny (transition from female to male), whereas protandry (male to female) is less common.^[51] Simultaneous hermaphroditism and self-fertilization occur in the mangrove rivulus (Kryptolebias marmoratus), which maintains genetic variability by periodically producing males at temperatures below 19 °C.^[52]

Parental care is absent in approximately seventy-nine percent of oviparous teleost families.^[53][54][55] In the remaining twenty-one percent of teleost families, parental care occurs in diverse forms, including egg burying, nest construction, mouthbrooding, and internal gestation (viviparity and ovoviviparity).^[53][54][55] Male parental care is far more frequent than female care, often evolving in species where males defend breeding territories.^{[53][54][56][57][58][59]} Viviparity is relatively rare, present in about six percent of living teleost species.^[53][54][55] The earliest fossil record of viviparity among ray-finned fishes is found in Middle Triassic fossils of the genus Saurichthys.^[60][61]

Post-hatching ontogeny begins with an embryonic stage during which the free-swimming larva subsists on yolk reserves in an attached yolk sac.^[2][33] The embryonic period ends, and the larval period begins, only when the young fish starts taking external food. Larvae often exhibit temporary specialized adaptations, such as the pelagic leptocephalus larvae characteristic of Elopomorpha.^[33] Metamorphosis into the juvenile form involves the resorption of larval structures and the acquisition of the adult body plan, culminating in sexual maturity.^[33]

Distribution and Habitats

Ray-finned fishes inhabit almost every body of water on the planet.^[1][3] Approximately 40 to 45 percent of all ray-finned fish species live in fresh water.^[62] Around 40 to 45 percent of species reside in marine continental shelf waters, 10 to 15 percent inhabit deep oceanic waters, and only about 1.5 percent reside in the upper epipelagic zones of the open ocean.^[62] A small percentage are diadromous, moving between marine and freshwater habitats to feed or spawn, including anadromous species like salmon and catadromous species like freshwater eels.^[62][63]

Actinopterygians display adaptations across extraordinary environmental gradients.^[2][3] In highly acidic habitats, the cyprinid Tribolodon hakonensis survives in volcanic caldera lakes with water pH as low as 3.4 to 3.8.^[64] In contrast, Lake Magadi in Kenya is characterized by hyper-alkaline water (pH around 10.5), high salinity (up to 40 parts per thousand), and water temperatures near 40 °C, supporting the cichlid Alcolapia grahami.^[65] Extreme salinity tolerance is also found in pupfishes (Cyprinodontidae), which can endure salinities reaching 140 parts per thousand, four times that of sea water, and temperatures of 44 °C in desert ephemeral waters.^[65] In sub-zero Antarctic marine waters, notothenioid icefishes survive at temperatures of −1.8 °C to −2.2 °C by producing glycoprotein antifreeze compounds that arrest the formation of ice crystals in their tissues.^[3][65]

In the abyssal and hadal ocean depths, where ambient hydrostatic pressure exceeds hundreds of atmospheres, specialized actinopterygians persist.^[65][66] The deepest living fishes recorded in oceanic trenches are the snailfish Pseudoliparis swirei, filmed at depths of 8,178 m in the Mariana Trench, and the cusk-eel Abyssobrotula galatheae, dredged from 7,965 m in the Puerto Rico Trench.^[67] In freshwater, deep-water sculpins of the family Abyssocottidae dwell at depths of up to 1,600 m in Lake Baikal.^[68] Conversely, at high altitudes, the loach Triplophysa stolickai inhabits thermal springs in western Tibet at elevations of roughly 5,200 m above sea level, while its congener Triplophysa zhaoi lives in marshes 50 m below sea level in the Turpan Depression.^[69] Subterranean caves shelter numerous blind, depigmented species, including cave morphs of the Mexican tetra (Astyanax mexicanus) and the catfish Horaglanis krishnai.^[65]

Evolution and Fossil Record

The earliest known fossils attributed to ray-finned fishes date to the Late Silurian, roughly 420 million years ago.^[70][71] Isolated scales and microvertebrate remains from this period, including Andreolepis hedei and Lophosteus superbus, were found in Russia, Sweden, and Estonia.^[70][72] More recent analyses suggest that Andreolepis and Lophosteus represent stem osteichthyans rather than crown actinopterygians.^[73][74][75] The oldest well-documented anatomical fossil showing clear ray-finned cranial affinities is Meemannia, an early Devonian fish from China dated to approximately 415 million years ago, originally described as a sarcopterygian before high-resolution scans revealed ray-finned internal anatomy.^{[76][77][78][79]}

Throughout the Devonian, actinopterygians were overshadowed by sarcopterygians and placoderms, being represented by comparatively few genera such as Cheirolepis, Moythomasia, Stegotrachelus, and Raynerius.^[3][77][79] Cheirolepis, from the Middle and Late Devonian of Europe and Canada, possessed a long, low skull, deep heterocercal tail, and small ganoid scales.^[3][77] Following the end-Devonian extinction, actinopterygians underwent a radiation at the Devonian-Carboniferous boundary, rapidly becoming dominant nektonic vertebrates in freshwater and marine habitats.^[45][77][80]

During the Carboniferous and Permian, ray-finned fishes diversified into a wide array of morphological designs historically referred to as palaeoniscoids (Palaeonisciformes), an evolutionary grade characterized by ganoid scales, anterior eyes, and heterocercal tails.^[81][82] Specialized body plans emerged, including deep-bodied forms like Cheirodus, eel-like forms like Tarrasius, and elongated pursuit predators like the Permian Saurichthys.^[82][83] In the Triassic, Thoracopterus developed expanded wing-like pectoral fins and an asymmetrical lower caudal lobe that enabled gliding above the water, paralleling modern flying fishes.^[82]

Crown Neopterygii appeared in the Late Permian and expanded rapidly across the Triassic and Jurassic.^[47][84] Triassic and Jurassic neopterygians included deep-bodied, crushing-toothed Pycnodontiformes, shoaling Semionotiformes, Parasemionotiformes, and stem-teleosts.^[85] Stem teleosteomorphs like Prohalecites, Aspidorhynchiformes (elongated predatory fishes with rostral spikes), and Pachycormiformes flourished.^[86][87] Pachycormids included the massive filter-feeding Leedsichthys.^[1][9] The first true teleosts arose in the Mesozoic, characterized by homocercal tails, mobile premaxillae, and light elasmoid scales.^[87][88]

Several lineages of ray-finned fishes have gone through whole-genome duplication (paleopolyploidy) events.^[89][90][91] A whole-genome duplication occurred in ancestral teleosts approximately 320 million years ago, with roughly seventeen percent of duplicate genes retained over evolutionary time.^[89][90][91] An independent genome duplication occurred in chondrostean ancestors roughly 180 million years ago (ranging from 124 to 225 million years ago).^[89][90][91] Additional lineage-specific genome duplications took place subsequently in salmonids (80 to 100 million years ago) and multiple times within cyprinids, including carps and goldfish within the last 14 million years.^{[89][90][91][92][93][94][95][96][97][98][99]}

Taxonomy and Classification

The classification of actinopterygians has experienced substantial revision.^{[4][100][101]} Historically, ray-finned fishes were partitioned into three grades: Chondrostei (cartilage-bone fishes, including sturgeons, paddlefishes, and bichirs), Holostei (bony ganoids, including gars and bowfins), and Teleostei (modern bony fishes).^[28] The historical Chondrostei proved paraphyletic, as bichirs and reedfish represent the most basal extant actinopterygian clade, now classified as the subclass Cladistia.^[102][103] The remaining actinopterygians form the clade Actinopteri, which contains Chondrostei in the strict sense (Acipenseriformes) and Neopterygii.^{[101][102][103]} Holostei was once viewed as paraphyletic, but morphological re-evaluations and molecular phylogenomics have confirmed that gars (Ginglymodi / Lepisosteiformes) and bowfins (Halecomorphi / Amiiformes) form a monophyletic sister group to Teleostei.^{[104][105][106][107]}

Within Teleostei, representing over ninety-nine percent of living actinopterygians, modern classifications recognize three primary basal lineages: Elopomorpha (tarpons, bonefishes, spiny eels, true eels, and gulpers), Osteoglossomorpha (bony-tongued fishes, mooneyes, and elephantfishes), and Clupeocephala.^[100][101] Molecular studies demonstrate that Elopomorpha and Osteoglossomorpha together form a monophyletic assemblage named Oseanacephala (or Eloposteoglossocephala), sister to Clupeocephala.^{[101][108][109][110]} Clupeocephala is divided into Otomorpha (Otocephala) and Euteleostei.^[100][101] Otomorpha includes Clupeiformes (herrings and anchovies), Alepocephaliformes (slickheads), and Ostariophysi (Gonorynchiformes, Cypriniformes, Characiformes, Gymnotiformes, and Siluriformes).^[100][101]

Euteleostei comprises the remainder of modern bony fishes, consisting of basal clades such as Lepidogalaxiiformes, Protacanthopterygii (Argentiniformes, Salmoniformes, Esociformes, and Galaxiiformes), Stomiati (Osmeriformes and Stomiiformes), and Neoteleostei.^[100][101] Within Neoteleostei, the vast majority of diversity sits within Acanthomorpha (spiny-rayed fishes).^[88] Acanthomorphs include Lampriformes, Paracanthopterygii (Percopsiformes, Zeiformes, Gadiformes, and allies), Polymixiiformes, and Acanthopterygii. Modern phylogenomic studies have disassembled the historically polyphyletic Perciformes into clearly demarcated monophyletic clades, including Ophidiiformes, Batrachoidiformes, Gobiiformes, Syngnathiformes, Scombriformes, Anabantiformes, Carangiformes, Ovalentaria (Cichliformes, Atheriniformes, Beloniformes, Cyprinodontiformes, Blenniiformes), and Eupercaria (Labriformes, Acanthuriformes, Lophiiformes, Tetraodontiformes, Centrarchiformes, and Perciformes sensu stricto).^[100][101]

Relationship with Humans

Ray-finned fishes provide a primary source of animal protein, micronutrients, and economic livelihood for human populations worldwide.^[111] In 2019, global wild capture fisheries harvested nearly 80 million metric tons of fish, according to data from the Food and Agriculture Organization (FAO).^[112] Inland freshwater fisheries yielded over 11 million tons, dominated by cyprinids (over 1.87 million tons) and cichlids like tilapias (nearly 820,000 tons).^[112] Marine capture fisheries accounted for 66.5 million tons, dominated by schooling pelagic clupeiforms (anchovies, herrings, sardines), gadiforms (cods, pollocks), and scombrids (tunas, mackerels).^[112] Diadromous fishes such as shads, salmon and smelts caught at sea or in rivers amounted to just under 2 million tons.^[112]

The single most harvested wild fish species in 2019 was the Peruvian anchoveta (Engraulis ringens), with catches exceeding 4.2 million metric tons, down from 7.0 million tons in 2018.^[112] Other major wild-caught species include Alaska pollock (Gadus chalcogrammus, 3.49 million tons), skipjack tuna (Katsuwonus pelamis, 3.44 million tons), yellowfin tuna (Thunnus albacares, 1.57 million tons), Atlantic herring (Clupea harengus, 1.55 million tons), blue whiting (Micromesistius poutassou, 1.51 million tons), and European pilchard (Sardina pilchardus, 1.49 million tons).^[112] In freshwater capture fisheries, the silver cyprinid (Rastrineobola argentea) in Lake Victoria yielded 335,500 tons, while Nile tilapia (Oreochromis niloticus) and Nile perch (Lates niloticus) produced 280,000 and 250,000 tons, respectively.^[112]

Aquaculture has expanded to match or exceed capture fisheries, producing nearly 56.5 million metric tons of finfish in 2019.^[112] Freshwater aquaculture accounted for over 47 million tons, diadromous farming produced almost 6 million tons, and marine farming yielded over 3 million tons.^[112] The most heavily cultivated fish species globally is the grass carp (Ctenopharyngodon idella), with annual production reaching 5.73 million tons in 2019, followed by silver carp (Hypophthalmichthys molitrix, 4.83 million tons), Nile tilapia (Oreochromis niloticus, 4.59 million tons), common carp (Cyprinus carpio, 4.41 million tons), catla (Catla catla, 3.29 million tons), bighead carp (Hypophthalmichthys nobilis, 3.15 million tons), striped catfish (Pangasianodon hypophthalmus, 2.68 million tons), Atlantic salmon (Salmo salar, 2.62 million tons), roho labeo (Labeo rohita, 1.99 million tons), and milkfish (Chanos chanos, 1.54 million tons). In addition to meat, sturgeons are exploited for black caviar, while salmonids provide red caviar.^[113]

Beyond food production, ray-finned fishes are an important object of sport fishing and of great importance for aquarium keeping. In freshwater aquarium keeping, the most important fishes are otophysans such as cyprinids, tetras and various catfishes, followed by livebearers, labyrinth fishes and cichlids. Marine ornamental species include damselfishes, clownfishes, wrasses, tangs, and angelfishes.^[21]

Conservation Status

. According to the International Union for Conservation of Nature (IUCN) as of 2025, out of 27,481 assessed actinopterygian species, 90 are listed as Extinct (EX), 11 as Extinct in the Wild (EW), 697 as Critically Endangered (CR), 1,341 as Endangered (EN), and 1,507 as Vulnerable (VU), totaling 3,620 threatened species.^[114] An additional 5,220 species are listed as Data Deficient (DD) and cannot currently be assessed.^[114]

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