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Fish
Temporal range: 535–0 Ma Middle Cambrian – Recent
Diversity of various fish including sharks, stingrays, bony fish, jawless fish, and coelacanths.
Diversity of various fish including sharks, stingrays, bony fish, jawless fish, and coelacanths.
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
Tetrapods
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A fish (pl.: fish or fishes) is an aquatic, gill-bearing animal with a hard skull that lacks limbs with digits. This includes hagfish, lampreys, and both cartilaginous and bony fish. Approximately 95% of living fish species are ray-finned bony fish; around 99% of those are teleosts. As a group, if tetrapods are excluded, fish are paraphyletic and so do not form a taxonomic group.

The earliest organisms that can be classified as fish were soft-bodied chordates that first appeared during the Cambrian period. Although they lacked a true spine, they possessed notochords which allowed them to be more agile than their invertebrate counterparts. Fish would continue to evolve through the Paleozoic era, diversifying into a wide variety of forms. Many fish of the Paleozoic developed external armor that protected them from predators. The first fish with jaws appeared in the Silurian period, after which many (such as sharks) became formidable marine predators rather than just the prey of arthropods.

Most fish are ectothermic ("cold-blooded"), allowing their body temperatures to vary as ambient temperatures change, though some of the large active swimmers like white shark and tuna can hold a higher core temperature. Fish can acoustically communicate with each other, most often in the context of feeding, aggression or courtship.

Fish are an important resource for humans worldwide, especially as food. Commercial and subsistence fishers hunt fish in wild fisheries or farm them in ponds or in cages in the ocean. They are also caught by recreational fishers, kept as pets, raised by fishkeepers, and exhibited in public aquaria. Fish have had a role in culture through the ages, serving as deities, religious symbols, and as the subjects of art, books and movies.

Etymology

The word fish is inherited from Proto-Germanic, and is related to the Latin piscis and 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]

Evolution

Further information: Evolution of fish

Fossil history

Dunkleosteus was a giant Devonian armoured Placoderm, c. 400 mya.

The evolution of fish began about 530 million years ago during the Cambrian explosion. At this time the early chordates developed their skull and vertebral column.[5] The first fish lineages belong to the Agnatha, or jawless fish, such as Haikouichthys.[6] During the late Cambrian, other jawless forms such as conodonts first appeared.[5]

Jawed vertebrates appear in the fossil record from the Silurian, with giant armoured placoderms such as Dunkleosteus,[7] and the Acanthodii or spiny sharks. The jawed fish still extant also appeared during the late Silurian: the cartilaginous Chondrichthyes and the bony Osteichthyes. The bony fish evolved into two separate groups: the ray-finned Actinopterygii and the fleshy-finned Sarcopterygii.

During the Devonian, fish diversity greatly increased, especially among the ostracoderms and placoderms, and among the lobe-finned fish and early sharks, earning the Devonian the name "age of fishes".

Phylogeny

Fishes are a paraphyletic group, since any clade containing all fish, such as the Gnathostomata or (for bony fish) Osteichthyes, also contains the clade of tetrapods (four-limbed vertebrates, mostly terrestrial), which are not considered to be fish.[8][9][10] Some tetrapods, such as cetaceans and ichthyosaurs, have secondarily acquired a fish-like body shape through convergent evolution.[11] 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".[10] The cladogram shows clades traditionally considered as "fishes" (cyan line), along with the tetrapods. Extinct groups are marked with a dagger (†).

Vertebrata/

†Euconodonta

†Pteraspidomorphi

†Thelodonti

†Anaspida

†Galeaspida

†Pituriaspida

†Osteostraci

Gnathostomata

"†Placodermi" (armoured fishes, paraphyletic)[12]

"†Acanthodii" ("spiny sharks", paraphyletic or polyphyletic)[13]

Chondrichthyes
(cartilaginous fishes)
Euteleostomi/

"†Acanthodii" ("spiny sharks", paraphyletic or polyphyletic)

Actinopterygii
(ray‑finned fishes)
Sarcopterygii
(lobe‑finned fish)
Osteichthyes
(jawed vertebrates)
"Fishes"
Craniata

Taxonomy

Main article: Taxonomy of fish

Fishes are a paraphyletic group and for this reason, the class Pisces seen in older reference works is no longer used in formal classifications. Traditional classification divides fish into three extant classes (Agnatha, Chondrichthyes, and Osteichthyes), and with extinct forms sometimes classified within those groups, sometimes as their own classes.[14]

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.[10]

Diversity

Main article: Diversity of fish

A typical fish is ectothermic, 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.[15]

Streamlining and 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.[16] Fish range in size from the huge 16-metre (52 ft) whale shark to some tiny teleosts only 8-millimetre (0.3 in), such as the cyprinid Paedocypris progenetica[17] and the stout infantfish.[18]

The biodiversity of extant fish is unevenly distributed among the various groups, with teleosts making up 96% of fish species.[10] The cladogram[19] shows the evolutionary relationships of all groups of living fishes (with their respective diversity[10][20]) and the four-limbed vertebrates (tetrapods).[21]

Vertebrates

Jawless fish (118 species: hagfish, lampreys)

Jawed vertebrates

Cartilaginous fishes (>1,100 species: sharks, rays, chimaeras)

Osteichthyes
Lobe-finned fish
Rhipidistia

Tetrapoda (>38,000 species: amphibians, reptiles, birds, mammals)

Dipnoi (6 species: lungfish)

Actinistia (2 species: coelacanths)

Ray-finned fish

Ecology

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

Fish species are roughly divided equally between marine (oceanic) and freshwater ecosystems. 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. Exceptionally rich sites in the Amazon basin, such as Cantão State Park, can contain more freshwater fish species than occur in all of Europe.[22]

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.[23] With 34,300 described species, fish exhibit greater species diversity than any other group of vertebrates.[24] 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 8,370 m (27,460 ft).[25][26]

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,[28] while desert pupfish live in desert springs, streams, and marshes, sometimes highly saline, with water temperatures as high as 36 C.[29][30]

A few fish live mostly on land or lay their eggs on land near water.[31] Mudskippers feed and interact with one another on mudflats and go underwater to hide in their burrows.[32] A single undescribed species of Phreatobius has been called a true "land fish" as this worm-like catfish strictly lives among waterlogged leaf litter.[33][34] Cavefish of multiple species live in underground lakes, underground rivers or aquifers.[35]

Anatomy and physiology

Main articles: Fish anatomy and Fish physiology

Respiration

Further information: Aquatic respiration

Gills

Tuna gills inside the head, seen from the rear. Water flows through the mouth and over the gills, either pumped or as the fish swims. Blood is circulated through the gills, picking up oxygen and releasing carbon dioxide.

Most fish exchange gases using gills on either side of the pharynx. Gills consist of threadlike 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. In some fish, capillary blood flows in the opposite direction to the water, causing countercurrent exchange. The gills push the oxygen-poor water out through openings in the sides of the pharynx. Some fish, like sharks and lampreys, possess multiple gill openings. However, bony fish have a single gill opening on each side. This opening is hidden beneath a protective bony cover called an operculum. Juvenile bichirs have external gills, a primitive feature shared with larval amphibians.

Air breathing

Fish from multiple groups can live out of the water for extended periods. Amphibious fish such as the mudskipper can move about on land and live in oxygen-depleted water. The skin of anguillid eels may absorb oxygen directly. The buccal cavity of the electric eel may breathe air. Catfish of the families Loricariidae, Callichthyidae, and Scoloplacidae absorb air through their digestive tracts.[36] Bichirs and lungfish (except for the Australian lungfish), have paired lungs similar to those of tetrapods; they have to surface to gulp fresh air through the mouth and pass spent air out through the gills. Gar and bowfin have a vascularized swim bladder that functions in the same way. Loaches, trahiras, and many catfish breathe by passing air through the gut. Mudskippers breathe by absorbing oxygen across the skin (similar to frogs). Some fish have evolved accessory breathing organs: labyrinth fish such as gouramis and bettas have a labyrinth organ above the gills, while snakeheads, pikeheads, and airbreathing catfish have similar structures. Some air-breathing fish are able to survive in damp burrows for weeks without water, entering a state of aestivation (summertime hibernation) until water returns. Air breathing fish can be divided into obligate air breathers and facultative air breathers. Obligate air breathers, such as the African lungfish, must breathe air periodically or they suffocate. Facultative air breathers, such as some catfish, only breathe air if they need to and will otherwise rely on their gills for oxygen. Most air breathing fish are facultative air breathers; they avoid the energetic cost of rising to the surface and the fitness cost of exposure to surface predators.[36]

Circulation

Fish heart (model shown) has two chambers (to a mammal's four), driving blood in a single loop around the body (compared to a mammal's two loops).

Fish have a closed-loop circulatory system. The heart pumps the blood in a single loop throughout the body, with two chambers; for comparison, the mammal heart has two loops through the body and four chambers.[37] The first part is the sinus venosus, a thin-walled sac that collects blood from the fish's veins before allowing it to flow to the second part, the atrium, which is a large muscular chamber. The atrium serves as a one-way antechamber, sends blood to the third part, ventricle. The ventricle is another thick-walled, muscular chamber and it pumps the blood, first to the fourth part, bulbus arteriosus, a large tube, and then out of the heart. The bulbus arteriosus connects to the aorta, through which blood flows to the gills for oxygenation.

Digestion

Jaws allow fish to eat a wide variety of food, including plants and other organisms. Fish ingest food through the mouth and break it down in the esophagus. In the stomach, food is further digested and, in many fish, processed in finger-shaped pouches called pyloric caeca, which secrete digestive enzymes and absorb nutrients. Organs such as the liver and pancreas add enzymes and various chemicals as the food moves through the digestive tract. The intestine completes the process of digestion and nutrient absorption.

Excretion

Most fish release their nitrogenous wastes as ammonia. This may be excreted through the gills or filtered by the kidneys.

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 to saltwater.

Brain

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.[38] 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.[39] Fish brains are divided into several regions. At the front are the olfactory lobes, a pair of structures that receive and process signals from the nostrils via the two olfactory nerves.[38] The olfactory lobes are very large in fish that hunt primarily by smell, such as hagfish, sharks, and catfish. Behind the olfactory lobes is the two-lobed telencephalon, the structural equivalent to the cerebrum in higher vertebrates. In fish the telencephalon is concerned mostly with olfaction. Together these structures form the forebrain.[38] Connecting the forebrain to the midbrain is the diencephalon (in the diagram, this structure is below the optic lobes and consequently not visible). The diencephalon performs functions associated with hormones and homeostasis.[38] The pineal body lies just above the diencephalon. This structure detects light, maintains circadian rhythms, and controls color changes.[38] The midbrain (or mesencephalon) contains the two optic lobes. These are very large in species that hunt by sight, such as rainbow trout and cichlids.[38] The hindbrain (or metencephalon) is particularly involved in swimming and balance.[38] The cerebellum is a single-lobed structure that is typically the biggest part of the brain.[38] Hagfish and lampreys have relatively small cerebellae, while the mormyrid cerebellum is massive and apparently involved in their electrical sense.[38] The brain stem or myelencephalon is the brain's posterior.[38] As well as controlling some muscles and body organs, in bony fish at least, the brain stem governs respiration and osmoregulation.[38]

Sensory systems

Main article: Sensory systems in fish

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.[40] 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.[41] Some fish, such as catfish and sharks, have the ampullae of Lorenzini, electroreceptors that detect weak electric currents on the order of millivolt.[42] Vision in fishes is an important sensory system. Fish eyes are similar to those of terrestrial vertebrates like birds and mammals, but have a more spherical lens. Their retinas generally have both rods and cones (for scotopic and photopic vision); most species have colour vision. Some fish can see ultraviolet, while others can see polarized light. Amongst jawless fish, the lamprey has well-developed eyes, while the hagfish has only primitive eyespots.[43] 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.[44] 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.[45][46] The mechanism of fish magnetoreception remains unknown;[47] experiments in birds imply a quantum radical pair mechanism.[48]

Cognition

Further information: Fish intelligence

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.[49][50][51] Choerodon wrasse, archerfish, and Atlantic cod can solve problems and invent tools.[52] The monogamous cichlid Amatitlania siquia exhibits pessimistic behavior when prevented from being with its partner.[53] Fish orient themselves using landmarks; they may use mental maps based on multiple landmarks. Fish behavior in mazes reveals that they possess spatial memory and visual discrimination.[54] Fish have pain and fear responses; toadfish grunt when electrically shocked and over time come to grunt at the mere sight of an electrode.[55] Rainbow trout rock their bodies and rub their lips when injected with bee venom and acetic acid, apparently attempting to relieve pain.[56][57] The wrasse's neurons fired in a pattern resembling human neuronal patterns.[57] The claims have been called flawed as they do not prove that fish possess conscious awareness, especially not human-like awareness,[58] and their brains are very different.[59]

Locomotion

Main article: Fish locomotion
Anatomy of a lanternfish: 1) gill cover 2) lateral line 3) dorsal fin 4) fat fin 5) caudal peduncle 6) caudal fin 7) anal fin 8) photophores 9) pelvic fins 10) pectoral fins
Photo of white bladder that consists of a rectangular section and a banana-shaped section connected by a much thinner element
Swim bladder of a rudd (Scardinius erythrophthalmus)

Most fish move 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 back fin, backward force is applied to the water, and in conjunction with the fins, moves the fish forward. The fish's fins function like an airplane's flaps. Fins also increase the tail's surface area, increasing speed.[60]

The streamlined body of the fish decreases the amount of friction from the water. 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 adjusts their buoyancy through manipulation of gases. The scales of fish originate from the mesoderm (skin); they may be similar in structure to teeth.

Electrogenesis

The elephantnose fish 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.[61]
Further information: Electroreception and electrogenesis

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.[62] Strongly electric fish like the electric eel can in addition use their electric organs to generate shocks powerful enough to stun their prey.[63]

Endothermy

Most fish are exclusively cold-blooded or ectothermic. However, the Scombroidei are warm-blooded (endothermic), including the billfishes and tunas.[64] The opah, a lampriform, uses whole-body endothermy, generating heat with its swimming muscles to warm its body while countercurrent exchange minimizes heat loss.[65] 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 20 °C (68 °F) above the ambient water.[64][66][67]

Reproduction

Main article: Fish reproduction
Ovary of fish (Corumbatá)

The primary reproductive organs are testicles and ovaries. In most species, gonads are paired organs of similar size, which can be partially or totally fused.[68] Some fish have secondary organs that increase reproductive fitness.

In terms of spermatogonia distribution, the structure of teleosts testes has two types: in the most common, spermatogonia occur all along the seminiferous tubules, while in atherinomorph fish they are confined to the distal portion of these structures. Fish can present cystic or semi-cystic spermatogenesis in relation to the release phase of germ cells in cysts to the seminiferous tubules lumen.[68]

Fish ovaries may be of three types: gymnovarian, secondary gymnovarian or cystovarian. In the first type, the oocytes are released directly into the coelomic cavity and then enter the ostium, then through the oviduct and are eliminated. Secondary gymnovarian ovaries shed ova into the coelom from which they go directly into the oviduct. In the third type, the oocytes are conveyed to the exterior through the oviduct.[69] Gymnovaries are the primitive condition found in lungfish, sturgeon, and bowfin. Cystovaries characterize most teleosts, where the ovary lumen has continuity with the oviduct.[68] Secondary gymnovaries are found in salmonids and a few other teleosts.

Oogonia development in teleosts fish varies according to the group, and the determination of oogenesis dynamics allows the understanding of maturation and fertilization processes. Changes in the nucleus, ooplasm, and the surrounding layers characterize the oocyte maturation process.[68] Postovulatory follicles are structures formed after oocyte release; they do not have endocrine function, present a wide irregular lumen, and are rapidly reabsorbed in a process involving the apoptosis of follicular cells. A degenerative process called follicular atresia reabsorbs vitellogenic oocytes not spawned. This process can also occur, but less frequently, in oocytes in other development stages.[68]

Some fish, like the California sheephead, are hermaphrodites, having both testes and ovaries either at different phases in their life cycle or, as in hamlets, have them simultaneously.

Over 97% of fish are oviparous,[70] that is, the eggs develop outside the mother's body. Examples of oviparous fish include salmon, goldfish, cichlids, tuna, and eels. In the majority of these species, fertilisation takes place outside the mother's body, with the male and female fish shedding their gametes into the surrounding water. However, a few oviparous fish practice internal fertilization, with the male using some sort of intromittent organ to deliver sperm into the genital opening of the female, most notably the oviparous sharks, such as the horn shark, and oviparous rays, such as skates. In these cases, the male is equipped with a pair of modified pelvic fins known as claspers.

Marine fish can produce high numbers of eggs which are often released into the open water column. The eggs have an average diameter of 1 millimetre (0.04 in).

The newly hatched young of oviparous fish are called larvae. They are usually poorly formed, carry a large yolk sac (for nourishment), and do not resemble juvenile or adult fish. The larval period in oviparous fish is usually only someweeks, and larvae rapidly grow and change in structure to become juveniles. During this transition, larvae must switch from their yolk sac to feeding on zooplankton prey, a process which depends on typically inadequate zooplankton density, starving many larvae.

In ovoviviparous fish the eggs develop inside the mother's body after internal fertilization but receive little or no nourishment directly from the mother, depending instead on the yolk. Each embryo develops in its own egg. Familiar examples of ovoviviparous fish include guppies, angel sharks, and coelacanths.

Some species of fish are viviparous. In such species the mother retains the eggs and nourishes the embryos. Typically, viviparous fish have a structure analogous to the placenta seen in mammals connecting the mother's blood supply with that of the embryo. Examples of viviparous fish include the surf-perches, splitfins, and lemon shark. Some viviparous fish exhibit oophagy, in which the developing embryos eat other eggs produced by the mother. This has been observed primarily among sharks, such as the shortfin mako and porbeagle, but is known for a few bony fish as well, such as the halfbeak Nomorhamphus ebrardtii.[71] Intrauterine cannibalism is an even more unusual mode of vivipary, in which the largest embryos eat weaker and smaller siblings. This behavior is also most commonly found among sharks, such as the grey nurse shark, but has also been reported for Nomorhamphus ebrardtii.[71]

Behavior

Shoaling and schooling

Main article: Shoaling and schooling
Fish such as these trumpetfishes school for safety from predators, and to spawn.[72]

An assemblage of fish merely using some localised resource such as food or nesting sites is called an aggregation. 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.[73] Schooling is sometimes an 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.[72]

Communication

See also: Acoustic communication in aquatic animals

Fish communicate by transmitting sounds, acoustic signals, to each other. This is most often in the context of feeding, aggression or courtship.[74] 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.[75]

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 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.[75] The longsnout seahorse, Hippocampus reidi produces two categories of sounds, 'clicks' and 'growls', by rubbing their coronet bone across the grooved section of their neurocranium.[76] 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.[77]

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.[78] Female and male toadfishes emit short-duration grunts, often as a fright response.[79] In addition to short-duration grunts, male toadfishes produce "boat whistle calls".[80] These calls are longer in duration, lower in frequency, and are primarily used to attract mates.[80] The sounds emitted by the O. tao have frequency range of 140 Hz to 260 Hz.[80] The frequencies of the calls depend on the rate at which the sonic muscles contract.[81][78]

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.[82]

Defense against disease

Further information: Fish diseases and parasites
A cleaner fish, a Hawaiian cleaner wrasse, cleaning parasites from a white-spotted puffer

Like other animals, fish suffer from diseases and parasites. To prevent disease they have a variety of defenses. 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, fish can develop an inflammatory response that increases blood flow to the infected region and delivers white blood cells that attempt to destroy pathogens. Specific defenses respond to particular pathogens recognised by the fish's body, i.e., an immune response.[83]

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.[84] 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 Etroplus suratensis.[85]

Immune organs vary by type of fish. In jawless fish, true lymphoid organs are absent. These fish rely on regions of lymphoid tissue within other organs to produce immune cells. For example, erythrocytes, macrophages and plasma cells are produced in the anterior kidney and some areas of the gut (where granulocytes mature). They resemble primitive bone marrow in hagfish. Cartilaginous fish have a more advanced immune system with three specialized organs: the epigonal organs around the gonads, Leydig's organ within the esophagus, and a spiral valve in their intestine. These organs house typical immune cells such as lymphocytes. They also possess a thymus and a well-developed spleen where lymphocytes, plasma cells and macrophages develop and are stored. Chondrostean fish produce granulocytes in tissue near the meninges. Their heart is covered with tissue that contains lymphocytes and reticular cells, while erythrocytes, granulocytes, lymphocytes and macrophages develop in the kidney.[86]

The major immune tissues of bony fish include the kidney, which houses many different immune cells.[87] In addition, teleost fish possess a thymus, spleen and scattered immune areas within mucosal tissues (e.g. in the skin, gills, gut and gonads). Much like the mammalian immune system, teleost erythrocytes, neutrophils and granulocytes are believed to reside in the spleen whereas lymphocytes are the major cell type found in the thymus.[88][89] An unconfirmed report claims a lymphatic system similar to that in mammals in teleosts, presumably where naive T cells accumulate while waiting to encounter an antigen.[90]

B and T lymphocytes bearing immunoglobulins and T cell receptors respectively are found in all jawed fishes. Indeed, the adaptive immune system as a whole evolved in an ancestor of all jawed vertebrates.[91]

Conservation

The 2006 IUCN Red List names 1,173 fish species that are threatened with extinction.[92] Included are species such as Atlantic cod,[93] Devil's Hole pupfish,[94] coelacanths,[95] and great white sharks.[96] 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.[97]

Overfishing

Main article: Overfishing
Collapse of the Atlantic northwest cod fishery

Overfishing is a major threat to edible fish such as cod and tuna.[98][99] Overfishing eventually causes population (known as stock) 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. A well-studied example of fishery collapse is the Pacific sardine Sadinops sagax caerulues fishery off the California coast. From a 1937 peak of 790,000 long tons (800,000 t) the catch steadily declined to only 24,000 long tons (24,000 t) in 1968, after which the fishery was no longer economically viable.[100]

Fisheries scientists and the fishing industry have different views on the resiliency of fisheries to intensive fishing. In places such as Scotland, Newfoundland, and Alaska the fishing industry is a major employer, so governments are predisposed to support it.[101][102] On the other hand, scientists and conservationists push for stringent protection, warning that many stocks could be wiped out within fifty years.[103][104]

Habitat destruction

Further information: Environmental impact of fishing

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.[105] An example of a fish that has become endangered because of habitat change is the pallid sturgeon, a North American freshwater fish that lives in rivers damaged by human activity.[106]

Exotic species

Introduction of non-native species occurs in many habitats. The Mediterranean Sea has become a major 'hotspot' of exotic invaders since the opening of the Suez Canal in 1869. A thousand marine species of all sorts – fishes, seaweeds, invertebrates – originating from the Red Sea and more broadly from the Indo-Pacific have crossed the Canal from south to north to settle in the eastern Mediterranean Basin. Nowadays many of these tropical or Lessepsian migrants, have extended their range towards the west, obviously favoured by the general warming of the Mediterranean. The resulting change in biodiversity is without precedent in human memory and is accelerating: a long-term cross-Basin survey engaged by the Mediterranean Science Commission recently documented that in just twenty years, from 2001 till 2021, no less than 107 alien fish species have reached the Mediterranean from both the tropical Atlantic and the Red Sea, which is more than the total recorded during the whole 130 preceding years.[107]

Another mode of introduction for marine species is transport across thousands of kms on ship hulls or in ballast waters. Examples abound of marine organisms being transported in ballast water, among them the invasive comb jelly Mnemiopsis leidyi, the dangerous bacterium Vibrio cholerae, or the fouling zebra mussel. The Mediterranean and Black Seas, with their high volume shipping from exotic harbors, are particularly impacted by this problem.[108]

Deliberate introductions of species with market potential are another frequent vector. A well-studied example is the introduction of the Nile perch into Lake Victoria in the 1960s. This predatory fish gradually exterminated the lake's 500 endemic cichlid species. Some of them now survive in captive breeding programmes, but others are probably extinct.[109]

Importance to humans

Economic

Main articles: Commercial fishing, Fishing industry, Aquaculture, and Fish farming
A trawler hauling in a large catch of cod, 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,[110] is becoming increasingly important in many nations. Overall, about one-sixth of the world's protein is estimated to be provided by fish.[111] Fishing is accordingly a large global business which provides income for millions of people.[111] The Environmental Defense Fund has a guide on which fish are safe to eat, given the state of pollution in today's world, and which fish are obtained in a sustainable way.[112] As of 2020, over 65 million tonnes (Mt) of marine fish and 10 Mt of freshwater fish were captured, while some 50 Mt of fish, mainly freshwater, were farmed. Of the marine species captured in 2020, 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.[113]

Recreation

Further information: Fishkeeping and Recreational fishing

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 is done with a rod, reel, line, hooks, and any one of a wide range of baits. Recreational fishing is particularly popular in North America and Europe and state, provincial, and federal government agencies actively management target fish species.[114][115]

Culture

Main article: Fish in culture

Fish themes have symbolic significance in many religions. In ancient Mesopotamia, fish offerings were made to the gods from the very earliest times.[116] Fish were also a major symbol of Enki, the god of water.[116] Fish frequently appear as filling motifs in cylinder seals from the Old Babylonian (c. 1830 BC – c. 1531 BC) and Neo-Assyrian (911–609 BC) periods.[116] Starting during the Kassite Period (c. 1600 BC – c. 1155 BC) and lasting until the early Persian Period (550–30 BC), healers and exorcists dressed in ritual garb resembling the bodies of fish.[116] During the Seleucid Period (312–63 BC), the legendary Babylonian culture hero Oannes, described by Berossus, was said to have dressed in the skin of a fish.[116] Fish were sacred to the Syrian goddess Atargatis[117] and, during her festivals, only her priests were permitted to eat them.[117] 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.[118] Early Christians used the ichthys, a symbol of a fish, to represent Jesus,[117][119] because the Greek word for fish, ΙΧΘΥΣ Ichthys, could be used as an acronym for "Ίησοῦς Χριστός, Θεοῦ Υἱός, Σωτήρ" (Iesous Christos, Theou Huios, Soter), meaning "Jesus Christ, Son of God, Saviour".[117][119] Among the deities said to take the form of a fish are Ika-Roa of the Polynesians, Dagon of various ancient Semitic peoples, the shark-gods of Hawaiʻi and Matsya of the Hindus. The astrological symbol Pisces is based on a constellation of the same name, but there is also a second fish constellation in the night sky, Piscis Austrinus.[120]

Fish feature prominently in art, in movies 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, in turn parodied in Shark Tale and Snakehead Terror. Piranhas are shown in a similar light to sharks in films such as Piranha.[121]

See also

Main article: Outline of fish

Notes

  1. ^ The temperature is often around 0 C. The freezing point of seawater at the surface is -1.85 C, falling to -2.62 C at a depth of 1000 metres. However, the water can be supercooled somewhat below these temperatures.[27]

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