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Temporal range: 535–0 
Bala shark, a bony fish
Bala shark, a bony fish
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Olfactores
Subphylum: Vertebrata
Groups included
Jawless fish
Armoured fish
Spiny sharks
Cartilaginous fish
Bony fish
Ray-finned fish
Lobe-finned fish
Cladistically included but traditionally excluded taxa

A fish (pl.: fish or fishes) is an

core temperature. Many fish can communicate acoustically with each other, such as during courtship displays

The earliest fish appeared during the

predators. The first fish with jaws, the placoderms, appeared in the Silurian and greatly diversified during the Devonian
, the "Age of Fishes".

Bony fish, distinguished by the presence of


Fish have been an important

human culture
through the ages, serving as deities, religious symbols, and as the subjects of art, books and movies.


The word fish is inherited from

Old Irish īasc, though the exact root is unknown; some authorities reconstruct a Proto-Indo-European root *peysk-, attested only in Italic, Celtic, and Germanic.[1][2][3][4]


Fossil history


About 530 million years ago during the

fossil record.[5] During the late Cambrian, other jawless forms such as conodonts appear.[6][7]

placoderms such as Dunkleosteus.[8] Jawed fish, too, appeared during the Silurian:[9] the cartilaginous Chondrichthyes[10][11] and the bony Osteichthyes.[12]

During the Devonian, fish diversity greatly increased, including among the placoderms, lobe-finned fishes, and early sharks, earning the Devonian the epithet "the age of fishes".[13][14]


Fishes are a

Extinct groups are marked with a dagger (†); groups of uncertain placement[19]
are labelled with a question mark (?) and dashed lines (- - - - -).




Conodonta, †Anaspida








 (>1,100 species:


Lobe-finned fish

 (2 species: coelacanths)


Dipnoi (6 species: lungfish)

Tetrapoda (>38,000 species, not considered fish: amphibians, reptiles, birds, mammals)

Ray-finned fish

 (14 species:

bichirs, reedfish)


 (27 species:

sturgeons, paddlefish)


 (>32,000 species)



Fishes (without tetrapods) are a

extant classes (Agnatha, Chondrichthyes, and Osteichthyes), and with extinct forms sometimes classified within those groups, sometimes as their own classes.[21]

Fish account for more than half of vertebrate species. As of 2016, there are over 32,000 described species of bony fish, over 1,100 species of cartilaginous fish, and over 100 hagfish and lampreys. A third of these fall within the nine largest families; from largest to smallest, these are

monotypic, containing only one species.[16]


Fish range in size from the huge 16-metre (52 ft)

stout infantfish.[24]

Swimming performance varies from fish such as tuna, salmon, and jacks that can cover 10–20 body-lengths per second to species such as eels and rays that swim no more than 0.5 body-lengths per second.[25]

  • Fastest: e.g. salmon, 10-20 body lengths/second
    Fastest: e.g. salmon, 10-20 body lengths/second
  • Slowest: e.g. eel, 0.5 body lengths/second
    Slowest: e.g. eel, 0.5 body lengths/second

A typical fish is

streamlined body for rapid swimming, extracts oxygen from water using gills, has two sets of paired fins, one or two dorsal fins, an anal fin and a tail fin, jaws, skin covered with scales, and lays eggs. Each criterion has exceptions, creating a wide diversity in body shape and way of life. For example, some fast-swimming fish are warm-blooded, while some slow-swimming fish have abandoned streamlining in favour of other body shapes.[26]



Different fish species are adapted to a wide variety of freshwater and marine habitats.

Fish species are roughly divided equally between

Neotropical freshwaters alone, such that Neotropical fishes represent about 10% of all vertebrate species on the Earth.[30]

Fish are abundant in most bodies of water. They can be found in nearly all aquatic environments, from high

Puerto Rico trench at 8,370 m (27,460 ft).[32]

In terms of temperature, Jonah's icefish live in cold[a] waters of the Southern Ocean, including under the Filchner–Ronne Ice Shelf at a latitude of 79°S,[34] while desert pupfish live in desert springs, streams, and marshes, sometimes highly saline, with water temperatures as high as 36 C.[35][36]

A few fish live mostly on land or lay their eggs on land near water.

underground rivers or aquifers.[41]

Parasites and predators

Like other animals, fish suffer from

E. suratensis.[43]

Fish occupy many trophic levels in freshwater and marine food webs. Fish at the higher levels are predatory, and a substantial part of their prey consists of other fish.[44] In addition, mammals such as dolphins and seals feed on fish, alongside birds such as gannets and cormorants.[45]

Anatomy and physiology

Anatomy and locomotion

The body of a typical fish is adapted for efficient swimming by alternately contracting paired sets of muscles on either side of the backbone. These contractions form S-shaped curves that move down the body. As each curve reaches the tail fin, force is applied to the water, moving the fish forward. The other fins act as control surfaces like an aircraft's flaps, enabling the fish to steer in any direction.[46]

Since body tissue is denser than water, fish must compensate for the difference or they will sink. Many bony fish have an internal organ called a swim bladder that allows them to adjust their buoyancy by increasing or decreasing the amount of gas it contains.[47]

The scales of fish provide protection from predators at the cost of adding stiffness and weight.[48] Fish scales are often highly reflective; this silvering provides camouflage in the open ocean. Because the water all around is the same colour, reflecting an image of the water offers near-invisibility.[49]

  • Gas-filled swim bladder of a rudd helps maintain neutral buoyancy.
    rudd helps maintain neutral buoyancy
  • Silvered scales of a rohu provide protection and camouflage.
    Silvered scales of a rohu provide protection and camouflage.


The fish heart pumps blood to the gills, where it picks up oxygen. The blood then flows without further pumping to the body, from where it returns to the heart.

Fish have a closed-loop circulatory system. The heart pumps the blood in a single loop throughout the body; for comparison, the mammal heart has two loops, one for the lungs to pick up oxygen, one for the body to deliver the oxygen. In fish, the heart pumps blood through the gills. Oxygen-rich blood then flows without further pumping, unlike in mammals, to the body tissues. Finally, oxygen-depleted blood returns to the heart.[50]



Fish exchange gases using gills on either side of the pharynx. Gills consist of comblike structures called filaments. Each filament contains a capillary network that provides a large surface area for exchanging oxygen and carbon dioxide. Fish exchange gases by pulling oxygen-rich water through their mouths and pumping it over their gills. Capillary blood in the gills flows in the opposite direction to the water, resulting in efficient countercurrent exchange. The gills push the oxygen-poor water out through openings in the sides of the pharynx. Cartilaginous fish have multiple gill openings: sharks usually have five, sometimes six or seven pairs; they often have to swim to oxygenate their gills. Bony fish have a single gill opening on each side, hidden beneath a protective bony cover or operculum. They are able to oxygenate their gills using muscles in the head.[51]

Air breathing

Some 400 species of fish in 50 families can breathe air, enabling them to live in oxygen-poor water or to emerge on to land.

low tide, and to use their gills when in water. Some coastal fish like rockskippers and mudskippers choose to leave the water to feed in habitats temporarily exposed to the air.[52] Some catfish absorb air through their digestive tracts.[53]


The digestive system consists of a tube, the gut, leading from the mouth to the anus. The mouth of most fishes contains teeth to grip prey, bite off or scrape plant material, or crush the food. An

pyloric caeca, around the pylorus, of doubtful function. The pancreas secretes enzymes into the intestine to digest the food; other enzymes are secreted directly by the intestine itself. The liver produces bile which helps to break up fat into an emulsion which can be absorbed in the intestine.[54]


Most fish release their nitrogenous wastes as

filtered by the kidneys. Salt is excreted by the rectal gland.[55] Saltwater fish tend to lose water by osmosis; their kidneys return water to the body, and produce a concentrated urine. The reverse happens in freshwater fish: they tend to gain water osmotically, and produce a dilute urine. Some fish have kidneys able to operate in both freshwater and saltwater.[56]


Diagram showing the pairs of olfactory, telencephalon, and optic lobes, followed by the cerebellum and the mylencephalon
Diagram of rainbow trout brain, from above

Fish have small brains relative to body size compared with other vertebrates, typically one-fifteenth the brain mass of a similarly sized bird or mammal.

mormyrids, processing their electrical sense. The brain stem or myelencephalon controls some muscles and body organs, and governs respiration and osmoregulation.[57]

Sensory systems


electroreceptors that detect weak electric currents on the order of millivolt.[61]

jawless fish, the lamprey has well-developed eyes,[64] while the hagfish has only primitive eyespots.[65]

Hearing too is an important sensory system in fish. Fish sense sound using their lateral lines and otoliths in their ears, inside their heads. Some can detect sound through the swim bladder.[66]

Some fish, including salmon, are capable of


The cognitive capacities of fish include

monogamous cichlid Amatitlania siquia exhibits pessimistic behavior when prevented from being with its partner.[74] Fish orient themselves using landmarks; they may use mental maps based on multiple landmarks. Fish are able to learn to traverse mazes, showing that they possess spatial memory and visual discrimination.[75]


electric organ and then uses its electroreceptive organs to locate objects by the distortions they cause in its electric field.[76]

Electric fish such as elephantfishes, the African knifefish, and electric eels have some of their muscles adapted to generate electric fields. They use the field to locate and identify objects such as prey in the waters around them, which may be turbid or dark.[61] Strongly electric fish like the electric eel can in addition use their electric organs to generate shocks powerful enough to stun their prey.[77]


Most fish are exclusively cold-blooded or

porbeagle shark, which maintain body temperatures more than 20 °C (68 °F) above the ambient water.[78][80][81]

Reproduction and life-cycle

Salmon fry hatching from the egg, keeping its yolk sac

The primary reproductive organs are paired

viviparous or live-bearing, meaning that the mother retains the eggs and nourishes the embryos via a structure analogous to the placenta to connect the mother's blood supply with the embryo's.[85]

Defenses against disease

Fish have both non-specific and immune defenses against disease. Non-specific defenses include the skin and scales, as well as the mucus layer secreted by the

white blood cells that attempt to destroy pathogens, non-specifically. Specific defenses respond to particular antigens, such as proteins on the surfaces of pathogenic bacteria, recognised by the adaptive immune system.[86] Immune systems evolved in deuterostomes as shown in the cladogram.[87]

innate immunity

Immune organs vary by type of fish. The jawless fish have

T cell receptors respectively. This makes use of Variable–Diversity–Joining rearrangement (V(D)J) to create immunity to a wide range of antigens. This system evolved once and is basal to the jawed vertebrate clade.[87] Cartilaginous fish have three specialized organs that contain immune system cells: the epigonal organs around the gonads, Leydig's organ within the esophagus, and a spiral valve in their intestine, while their thymus and spleen have similar functions to those of the same organs in the immune systems of tetrapods.[88] Teleosts have lymphocytes in the thymus, and other immune cells in the spleen and other organs.[89][90]


Shoaling and schooling

Fish such as these trumpetfishes school for safety from predators, and to spawn.[91]

A shoal is a loosely organised group where each fish swims and forages independently but is attracted to other members of the group and adjusts its behaviour, such as swimming speed, so that it remains close to the other members of the group. A school is a much more tightly organised group, synchronising its swimming so that all fish move at the same speed and in the same direction.

antipredator adaptation, offering improved vigilance against predators. It is often more efficient to gather food by working as a group, and individual fish optimise their strategies by choosing to join or leave a shoal. When a predator has been noticed, prey fish respond defensively, resulting in collective shoal behaviours such as synchronised movements. Responses do not consist only of attempting to hide or flee; antipredator tactics include for example scattering and reassembling. Fish also aggregate in shoals to spawn.[91] The capelin migrates annually in large schools between its feeding areas and its spawning grounds.[93]


Fish communicate by transmitting acoustic signals (sounds) to each other. This is most often in the context of feeding, aggression or courtship.[94] The sounds emitted vary with the species and stimulus involved. Fish can produce either stridulatory sounds by moving components of the skeletal system, or can produce non-stridulatory sounds by manipulating specialized organs such as the swimbladder.[95]

French grunt fish makes sounds by grinding its teeth.

Some fish produce sounds by rubbing or grinding their bones together. These sounds are stridulatory. In

Hippocampus reidi produces two categories of sounds, 'clicks' and 'growls', by rubbing their coronet bone across the grooved section of their neurocranium.[96] Clicks are produced during courtship and feeding, and the frequencies of clicks were within the range of 50 Hz-800 Hz. The frequencies are at the higher end of the range during spawning, when the female and male fishes were less than fifteen centimeters apart. Growls are produced when the H. reidi are stressed. The 'growl' sounds consist of a series of sound pulses and are emitted simultaneously with body vibrations.[97]

Some fish species create noise by engaging specialized muscles that contract and cause swimbladder vibrations. Oyster toadfish produce loud grunts by contracting sonic muscles along the sides of the swim bladder.[98] Female and male toadfishes emit short-duration grunts, often as a fright response.[99] In addition to short-duration grunts, male toadfishes produce "boat whistle calls".[100] These calls are longer in duration, lower in frequency, and are primarily used to attract mates.[100] The various sounds have frequency range of 140 Hz to 260 Hz.[100] The frequencies of the calls depend on the rate at which the sonic muscles contract.[101][98]

The red drum,

Sciaenops ocellatus, produces drumming sounds by vibrating its swimbladder. Vibrations are caused by the rapid contraction of sonic muscles that surround the dorsal aspect of the swimbladder. These vibrations result in repeated sounds with frequencies from 100 to >200 Hz. S. ocellatus produces different calls depending on the stimuli involved, such as courtship or a predator's attack. Females do not produce sounds, and lack sound-producing (sonic) muscles.[102]


The 2024

Devil's Hole pupfish,[105] coelacanths,[106] and great white sharks.[107] Because fish live underwater they are more difficult to study than terrestrial animals and plants, and information about fish populations is often lacking. However, freshwater fish seem particularly threatened because they often live in relatively small water bodies. For example, the Devil's Hole pupfish occupies only a single 3 by 6 metres (10 by 20 ft) pool.[108]


Collapse of the Atlantic northwest cod fishery[109]


Pacific sardine fishery off the California coast, the catch steadily declined from a 1937 peak of 800,000 tonnes to an economically inviable 24,000 tonnes in 1968.[113] In the case of the Atlantic northwest cod fishery, overfishing reduced the fish population to 1% of its historical level by 1992.[109]
Fisheries scientists and the fishing industry have sharply differing views on the resiliency of fisheries to intensive fishing. In many coastal regions the fishing industry is a major employer, so governments are predisposed to support it.[114][115] On the other hand, scientists and conservationists push for stringent protection, warning that many stocks could be destroyed within fifty years.[116][117]

Other threats

A key stress on both freshwater and marine ecosystems is

comb jelly in the Black Sea damaged the anchovy fishery there.[123][122] The opening of the Suez Canal in 1869 has created Lessepsian migration, in which hundreds of species of fish, algae, and invertebrates have entered the Mediterranean Sea, changing its ecology.[124] The predatory Nile perch was deliberately introduced to Lake Victoria in the 1960s as a commercial and sports fish. The lake had high biodiversity, with some 500 endemic species of cichlid fish. It drastically altered the lake's ecology, and simplified the fishery from multi-species to just three: the Nile perch, the silver cyprinid, and another introduced fish, the Nile tilapia. The haplochromine cichlid populations have collapsed.[125][126]

Importance to humans


trawler hauling in a large catch of cod
, 2016

Throughout history, humans have used

anchoveta represented 4.9 Mt, Alaska pollock 3.5 Mt, skipjack tuna 2.8 Mt, and Atlantic herring and yellowfin tuna 1.6 Mt each; eight more species had catches over 1 Mt.[130]


Fish have been recognized as a source of beauty for almost as long as used for food, appearing in

ornamental fish in ponds, and displayed in aquariums in homes, offices, or public settings. Recreational fishing is fishing primarily for pleasure or competition; it can be contrasted with commercial fishing, which is fishing for profit, or artisanal fishing, which is fishing primarily for food. The most common form of recreational fishing employs a rod, reel, line, hooks, and a wide range of baits. Recreational fishing is particularly popular in North America and Europe; government agencies often actively manage target fish species.[131][132]


Fish themes have symbolic significance in many religions. In ancient

Oannes was said to have dressed in the skin of a fish.[133] Fish were sacred to the Syrian goddess Atargatis[134] and, during her festivals, only her priests were permitted to eat them.[134] In the Book of Jonah, the central figure, a prophet named Jonah, is swallowed by a giant fish after being thrown overboard by the crew of the ship he is travelling on.[135] Early Christians used the ichthys, a symbol of a fish, to represent Jesus.[134][136] Among the deities said to take the form of a fish are Ikatere of the Polynesians,[137]
the shark-god Kāmohoaliʻi of Hawaiʻi,[138] and Matsya of the Hindus.[139] The constellation Pisces ("The Fishes") is associated with a legend from Ancient Rome that Venus and her son Cupid were rescued by two fishes.[140]

Fish feature prominently in art,[141] in films such as Finding Nemo[142] and books such as The Old Man and the Sea.[143] Large fish, particularly sharks, have frequently been the subject of horror movies and thrillers, notably the novel Jaws, made into a film which in turn has been parodied and imitated many times.[144] Piranhas are shown in a similar light to sharks in films such as Piranha.[145]

See also


  1. supercooled somewhat below these temperatures.[33]


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