Fish

The word fish is inherited from Proto-Germanic, and is related to German , the Latin , and Old Irish , though the exact root is unknown; some authorities reconstruct a Proto-Indo-European root , attested only in Italic, Celtic, and Germanic. Though often used interchangeably, in biology fish and fishes have different meanings. Fish is used as a singular noun, or as a plural to describe multiple individuals from a single species. Fishes is used to describe different species or species groups. == Evolution ==

Fossil history '' was a giant Devonian armoured placoderm, c. 400 mya. About 530 million years ago during the Cambrian explosion, fishlike animals with a notochord and eyes at the front of the body, such as Haikouichthys, appear in the fossil record. During the late Cambrian, other jawless forms such as conodonts appear. Jawed vertebrates appear in the Silurian, with giant armoured placoderms such as Dunkleosteus. Jawed fish, too, appeared during the Silurian: the cartilaginous Chondrichthyes and the bony Osteichthyes. 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". Phylogeny Fishes are a paraphyletic group, since any clade containing all jawed fish (Gnathostomata) or all bony fish (Osteichthyes) also contains the clade of tetrapods (four-limbed vertebrates, mostly terrestrial), which are usually not considered fish. On the other hand, Fishes of the World comments that "it is increasingly widely accepted that tetrapods, including ourselves, are simply modified bony fishes, and so we are comfortable with using the taxon Osteichthyes as a clade, which now includes all tetrapods". shows the evolutionary relationships of all groups of living fishes (with their respective diversity Extinct groups are marked with a dagger (†); groups of uncertain placement 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 Cyprinidae, Gobiidae, Cichlidae, Characidae, Loricariidae, Balitoridae, Serranidae, Labridae, and Scorpaenidae. About 64 families are monotypic, containing only one species. Diversity Fish range in size from the huge whale shark to some tiny teleosts only long, such as the cyprinid Paedocypris progenetica and the stout infantfish. File:Rhincodon typus fgbnms (cropped).jpg|Largest: whale shark File:Paedocypris progenetica 001.jpg|Smallest: e.g. Paedocypris progenetica 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. File:Salmo salar.jpg|Fastest: e.g. salmon, 10–20 body lengths/second File:Anguilla japonica 1856.jpg|Slowest: e.g. eel, 0.5 body lengths/second A typical fish is cold-blooded, has a 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. File:Humpback anglerfish.png|Ambush predator:anglerfish File:Atl mackerel photo3 exp.jpg|Streamlined, somewhat warm-blooded:mackerel File:Hippocampus hippocampus (cropped).jpg|Tail not used for swimming:seahorse File:Phycodurus eques P2023146 (cropped).JPG|Camouflaged:leafy seadragon File:Eastern Cleaner Clingfish (cropped).jpg|No scales:clingfish File:Cyphotilapia frontosa mouthbrooding.jpg|Mouthbrooder: front cichlid with young in mouth == Ecology ==

Habitats Fish species are roughly divided equally between freshwater and marine (oceanic) ecosystems; there are some 15,200 freshwater species and around 14,800 marine species. Coral reefs in the Indo-Pacific constitute the center of diversity for marine fishes, whereas continental freshwater fishes are most diverse in large river basins of tropical rainforests, especially the Amazon, Congo, and Mekong basins. More than 5,600 fish species inhabit Neotropical freshwaters alone, such that Neotropical fishes represent about 10% of all vertebrate species on the Earth. Fish are abundant in most bodies of water. They can be found in nearly all aquatic environments, from high mountain streams (e.g., char and gudgeon) to the abyssal and even hadal depths of the deepest oceans (e.g., cusk-eels and snailfish), although none have been found in the deepest 25% of the ocean. The deepest living fish in the ocean so far found is a cusk-eel, Abyssobrotula galatheae, recorded at the bottom of the Puerto Rico Trench at . In terms of temperature, Jonah's icefish live in cold waters of the Southern Ocean, including under the Filchner–Ronne Ice Shelf at a latitude of 79°S, while desert pupfish live in desert springs, streams, and marshes, sometimes highly saline, with water temperatures as high as 36 C. A few fish live mostly on land or lay their eggs on land near water. Mudskippers feed and interact with one another on mudflats and go underwater to hide in their burrows. A single undescribed species of Phreatobius has been called a true "land fish" as this worm-like catfish strictly lives among waterlogged leaf litter. Cavefish of multiple families live in underground lakes, underground rivers or aquifers. Parasites and predators Like other animals, fish suffer from parasitism. Some species use cleaner fish to remove external parasites. The best known of these are the bluestreak cleaner wrasses of coral reefs in the Indian and Pacific oceans. These small fish maintain cleaning stations where other fish congregate and perform specific movements to attract the attention of the cleaners. Cleaning behaviors have been observed in a number of fish groups, including an interesting case between two cichlids of the same genus, Etroplus maculatus, the cleaner, and the much larger E. suratensis. 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. In addition, mammals such as dolphins and seals feed on fish, alongside birds such as gannets and cormorants. File:Initial phase parrotfish feeding at Shaab Marsa Alam, Red Sea, Egypt -SCUBA (6336981391).jpg|A parrotfish feeding on algae on a coral reef File:Arothron hispidus is being cleaned by Hawaiian cleaner wrasses, Labroides phthirophagus 1.jpg|A cleaner fish removing parasites from its client, a pufferfish File:Barracuda with prey.jpg|A barracuda preying on a smaller fish File:1031 california sealion wright odfw (35281910502).jpg|Sea lion, a predatory mammal, eating a large salmonid File:Cormorant with fish (cropped).jpg|Cormorant with fish prey == Anatomy and physiology ==

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. File:Lampanyctodes hectoris (Hector's lanternfish).svg|Anatomy of a typical fish (lanternfish shown):1) gill cover 2) lateral line 3) dorsal fin 4) fat fin5) caudal peduncle 6) caudal fin 7) anal fin 8) photophores 9) pelvic fins 10) pectoral fins 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. The scales of fish provide protection from predators at the cost of adding stiffness and weight. 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. File:Swim bladder.jpg|Gas-filled swim bladder of a rudd helps maintain neutral buoyancy. File:Fish scales.jpg|Silvered scales of a rohu provide protection and camouflage. Circulation 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. Respiration Gills 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. 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. The ability of fish to do this is potentially limited by their single-loop circulation, as oxygenated blood from their air-breathing organ will mix with deoxygenated blood returning to the heart from the rest of the body. Lungfish, bichirs, ropefish, bowfins, snakefish, and the African knifefish have evolved to reduce such mixing, and to reduce oxygen loss from the gills to oxygen-poor water. Digestion 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 esophagus carries food to the stomach where it may be stored and partially digested. A sphincter, the pylorus, releases food to the intestine at intervals. Many fish have finger-shaped pouches, 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. Excretion Most fish release their nitrogenous wastes as ammonia. This may be excreted through the gills or filtered by the kidneys. Salt is excreted by the rectal gland. 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. Brain brain, from above|alt=Diagram showing the pairs of olfactory, telencephalon, and optic lobes, followed by the cerebellum and the mylencephalon 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. However, some fish have relatively large brains, notably mormyrids and sharks, which have brains about as large for their body weight as birds and marsupials. At the front of the brain are the olfactory lobes, a pair of structures that receive and process signals from the nostrils via the two olfactory nerves. Fish that hunt primarily by smell, such as hagfish and sharks, have very large olfactory lobes. Behind these is the telencephalon, which in fish deals mostly with olfaction. Together these structures form the forebrain. Connecting the forebrain to the midbrain is the diencephalon; it works with hormones and homeostasis. The pineal body is just above the diencephalon; it detects light, maintains circadian rhythms, and controls color changes. The midbrain contains the two optic lobes. These are very large in species that hunt by sight, such as rainbow trout and cichlids. The hindbrain controls swimming and balance. The single-lobed cerebellum is the biggest part of the brain; it is small in hagfish and lampreys, but very large in mormyrids, processing their electrical sense. The brain stem or myelencephalon controls some muscles and body organs, and governs respiration and osmoregulation. Sensory systems The lateral line system is a network of sensors in the skin which detects gentle currents and vibrations, and senses the motion of nearby fish, whether predators or prey. This can be considered both a sense of touch and of hearing. Blind cave fish navigate almost entirely through the sensations from their lateral line system. Some fish, such as catfish and sharks, have the ampullae of Lorenzini, electroreceptors that detect weak electric currents on the order of millivolt. Vision is an important sensory system in fish. Fish eyes are similar to those of terrestrial vertebrates like birds and mammals, but have a more spherical lens. some such as cyprinids have a fourth type of cone that detects ultraviolet. while the hagfish has only primitive eyespots. 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. Some fish, including salmon, are capable of magnetoreception; when the axis of a magnetic field is changed around a circular tank of young fish, they reorient themselves in line with the field. The mechanism of fish magnetoreception remains unknown; experiments in birds imply a quantum radical pair mechanism. Cognition The cognitive capacities of fish include self-awareness, as seen in mirror tests. Manta rays and wrasses placed in front of a mirror repeatedly check whether their reflection's behavior mimics their body movement. Choerodon wrasse, archerfish, and Atlantic cod can solve problems and invent tools. The monogamous cichlid Amatitlania siquia exhibits pessimistic behavior when prevented from being with its partner. 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. Behavioral research suggests that fish are sentient, capable of experiencing pain. Electrogenesis is a weakly electric fish which generates an electric field with its electric organ and then uses its electroreceptive organs to locate objects by the distortions they cause in its electric field. 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. Endothermy Most fish are exclusively cold-blooded or ectothermic. However, the Scombroidei are warm-blooded (endothermic), including the billfishes and tunas. The opah, a lampriform, uses whole-body endothermy, generating heat with its swimming muscles to warm its body while countercurrent exchange minimizes heat loss. Among the cartilaginous fishes, sharks of the families Lamnidae (such as the great white shark) and Alopiidae (thresher sharks) are endothermic. The degree of endothermy varies from the billfishes, which warm only their eyes and brain, to the bluefin tuna and the porbeagle shark, which maintain body temperatures more than above the ambient water. Reproduction and life-cycle fry hatching from the egg, keeping its yolk sac The primary reproductive organs are paired testicles and ovaries. Eggs are released from the ovary to the oviducts. Over 97% of fish, including salmon and goldfish, are oviparous, meaning that the eggs are shed into the water and develop outside the mother's body. The eggs are usually fertilized outside the mother's body, with the male and female fish shedding their gametes into the surrounding water. In a few oviparous fish, such as the skates, fertilization is internal: the male uses an intromittent organ to deliver sperm into the female's genital opening of the female. DNA repair Embryos of externally fertilized fish species are directly exposed during their development to environmental conditions that may damage their DNA, such as pollutants, UV light and reactive oxygen species. To deal with such DNA damages, a variety of different DNA repair pathways are employed by fish embryos during their development. 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 epidermis that traps and inhibits the growth of microorganisms. If pathogens breach these defenses, the innate immune system can mount an inflammatory response that increases blood flow to the infected region and delivers 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. Immune systems evolved in deuterostomes as shown in the cladogram. 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. Teleosts have lymphocytes in the thymus, and other immune cells in the spleen and other organs. == Behavior ==

Shoaling and schooling es school for safety from predators, and to spawn. The capelin migrates annually in large schools between its feeding areas and its spawning grounds. Communication Fish communicate by transmitting acoustic signals (sounds) to each other. This is most often in the context of feeding, aggression or courtship. 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. makes sounds by grinding its teeth. Some fish produce sounds by rubbing or grinding their bones together. These sounds are stridulatory. In Haemulon flavolineatum, the French grunt fish, as it produces a grunting noise by grinding its teeth together, especially when in distress. The grunts are at a frequency of around 700 Hz, and last approximately 47 milliseconds. 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. 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. Female and male toadfishes emit short-duration grunts, often as a fright response. In addition to short-duration grunts, male toadfishes produce "boat whistle calls". The frequencies of the calls depend on the rate at which the sonic muscles contract. == Conservation ==

The 2025 IUCN Red List names 2,367 fish species that are endangered or critically endangered. Included are species such as Atlantic cod, Devil's Hole pupfish, coelacanths, and great white sharks. Because fish live underwater they are more difficult to study than terrestrial animals and plants, and information about fish populations is often lacking. 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 pool. Overfishing . Overfishing is a major threat to edible fish such as cod and tuna. Overfishing eventually causes fish stocks to collapse, because the survivors cannot produce enough young to replace those removed. Such commercial extinction does not mean that the species is extinct, merely that it can no longer sustain a fishery. In the case of the 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. In the case of the Atlantic northwest cod fishery, overfishing reduced the fish population to 1% of its historical level by 1992. 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. On the other hand, scientists and conservationists push for stringent protection, warning that many stocks could be destroyed within fifty years. Other threats A key stress on both freshwater and marine ecosystems is habitat degradation including water pollution, the building of dams, removal of water for use by humans, and the introduction of exotic species including predators. Freshwater fish, especially if endemic to a region (occurring nowhere else), may be threatened with extinction for all these reasons, as is the case for three of Spain's ten endemic freshwater fishes. River dams, especially major schemes like the Kariba Dam (Zambezi river) and the Aswan Dam (River Nile) on rivers with economically important fisheries, have caused large reductions in fish catch. Industrial bottom trawling can damage seabed habitats, as has occurred on the Georges Bank in the North Atlantic. Introduction of aquatic invasive species is widespread. It modifies ecosystems, causing biodiversity loss, and can harm fisheries. Harmful species include fish but are not limited to them; the arrival of a comb jelly in the Black Sea damaged the anchovy fishery there. and ecology. 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. == Importance to humans ==

Economic hauling in hundreds of tons of Chilean jack mackerel, 2016 Throughout history, humans have used fish as a food source for dietary protein. Historically and today, most fish harvested for human consumption has come by means of catching wild fish. However, fish farming, which has been practiced since about 3,500 BCE in ancient China, is becoming increasingly important in many nations. In 2007, about one-sixth of the world's protein was estimated to be provided by fish. Fishing is a large global business which provides income for millions of people. Recreation Fish have been recognized as a source of beauty for almost as long as used for food, appearing in cave art, being raised as 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. Culture Fish themes have symbolic significance in many religions. In ancient Mesopotamia, fish offerings were made to the gods from the very earliest times. Fish were also a major symbol of Enki, the god of water. and, during her festivals, only her priests were permitted to eat them. Early Christians used the ichthys, a symbol of a fish, to represent Jesus. Among the deities said to take the form of a fish are Ikatere of the Polynesians, the shark-god Kāmohoaliʻi of Hawaii, and Matsya of the Hindus. The constellation Pisces ("The Fishes") is associated with a legend from Ancient Rome that Venus and her son Cupid were rescued by two fishes. Fish feature prominently in art, in films such as Finding Nemo and books such as The Old Man and the Sea. 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. Piranhas are shown in a similar light to sharks in films such as Piranha. File:Matsya painting.jpg|Avatar of Vishnu as a Matsya, India File:Bartolomeo Passerotti - The Fishmonger's Shop - WGA17072.jpg|''The Fishmonger's Shop'', Bartolomeo Passerotti, 1580s File:Goldfish Matisse.jpg|Goldfish by Henri Matisse, 1912 == See also ==