59-510: Labeobarbus is a mid-sized ray-finned fish genus in the family Cyprinidae . Its species are widely distributed throughout eastern Africa and especially southern Africa , but also in Lake Tana in Ethiopia . A common name , in particular for the southern species, is yellowfish . The scientific name refers to the fact that these large barbs recall the fairly closely related " carps " in
118-1054: A dagger (†); groups of uncertain placement are labelled with a question mark (?) and dashed lines (- - - - -). Jawless fishes (118 species: hagfish , lampreys ) [REDACTED] † Thelodonti , † Conodonta , † Anaspida [REDACTED] [REDACTED] [REDACTED] † Galeaspida [REDACTED] † Osteostraci [REDACTED] † Placodermi [REDACTED] † Acanthodii [REDACTED] (>1,100 species: sharks , rays , chimaeras ) [REDACTED] (2 species: coelacanths ) [REDACTED] Dipnoi (6 species: lungfish ) [REDACTED] Tetrapoda (>38,000 species, not considered fish: amphibians, reptiles, birds, mammals) [REDACTED] (14 species: bichirs , reedfish ) [REDACTED] (27 species: sturgeons , paddlefish ) [REDACTED] Ginglymodi (7 species: gars , alligator gars ) [REDACTED] Halecomorphi (2 species: bowfin , eyetail bowfin ) [REDACTED] (>32,000 species) [REDACTED] Fishes (without tetrapods) are
177-579: 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. 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
236-459: A free-swimming larval stage. However other patterns of ontogeny exist, with one of the commonest being sequential hermaphroditism . In most cases this involves protogyny , fish starting life as females and converting to males at some stage, triggered by some internal or external factor. Protandry , where a fish converts from male to female, is much less common than protogyny. Most families use external rather than internal fertilization . Of
295-436: A more spherical lens . Their retinas generally have both rods and cones (for scotopic and photopic vision ); many species have colour vision , often with three types of cone. Teleosts can see polarized light ; some such as cyprinids have a fourth type of cone that detects ultraviolet . Amongst jawless fish , the lamprey has well-developed eyes, while the hagfish has only primitive eyespots. Hearing too
354-419: A protective bony cover or operculum . They are able to oxygenate their gills using muscles in the head. 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
413-441: 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
472-485: 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. 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
531-726: A trait still present in Holostei ( bowfins and gars ). In some fish like the arapaima , the swim bladder has been modified for breathing air again, and in other lineages it have been completely lost. The teleosts have urinary and reproductive tracts that are fully separated, while the Chondrostei have common urogenital ducts, and partially connected ducts are found in Cladistia and Holostei. Ray-finned fishes have many different types of scales ; but all teleosts have leptoid scales . The outer part of these scales fan out with bony ridges, while
590-585: 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 . 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
649-422: 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. Since body tissue
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#1732851251812708-423: Is an aquatic , anamniotic , gill -bearing vertebrate animal with swimming fins and a hard skull , but lacking limbs with digits . Fish can be grouped into the more basal jawless fish and the more common jawed fish , the latter including all living cartilaginous and bony fish , as well as the extinct placoderms and acanthodians . Most fish are cold-blooded , their body temperature varying with
767-465: 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
826-431: 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
885-540: Is divided into the infraclasses Holostei and Teleostei . During the Mesozoic ( Triassic , Jurassic , Cretaceous ) and Cenozoic the teleosts in particular diversified widely. As a result, 96% of living fish species are teleosts (40% of all fish species belong to the teleost subgroup Acanthomorpha ), while all other groups of actinopterygians represent depauperate lineages. The classification of ray-finned fishes can be summarized as follows: The cladogram below shows
944-432: Is relatively rare and is found in about 6% of living teleost species; male care is far more common than female care. Male territoriality "preadapts" a species for evolving male parental care. There are a few examples of fish that self-fertilise. The mangrove rivulus is an amphibious, simultaneous hermaphrodite, producing both eggs and spawn and having internal fertilisation. This mode of reproduction may be related to
1003-465: 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 . 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
1062-500: The Cyprinidae (in goldfish and common carp as recently as 14 million years ago). Ray-finned fish vary in size and shape, in their feeding specializations, and in the number and arrangement of their ray-fins. In nearly all ray-finned fish, the sexes are separate, and in most species the females spawn eggs that are fertilized externally, typically with the male inseminating the eggs after they are laid. Development then proceeds with
1121-606: The Gnathostomata or (for bony fish) Osteichthyes , also contains the clade of tetrapods (four-limbed vertebrates, mostly terrestrial), which are usually not considered fish. Some tetrapods, such as cetaceans and ichthyosaurs , have secondarily acquired a fish-like body shape through convergent evolution . 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
1180-709: 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
1239-661: 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". Fishes are a paraphyletic group, since any clade containing all fish, such as
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#17328512518121298-469: 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 8,370 m (27,460 ft). In terms of temperature, Jonah's icefish live in cold waters of
1357-425: The end-Devonian extinction wiped out the apex placoderms. Bony fish are further divided into the lobe-finned and ray-finned fish . About 96% of all living fish species today are teleosts , a crown group of ray-finned fish that can protrude their jaws . The tetrapods , a mostly terrestrial clade of vertebrates that have dominated the top trophic levels in both aquatic and terrestrial ecosystems since
1416-400: The intertidal zone , are facultative air breathers, able to breathe air when out of water, as may occur daily at 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. Some catfish absorb air through their digestive tracts. The digestive system consists of a tube,
1475-405: The ostracoderms , had heavy bony plates that served as protective exoskeletons against invertebrate 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 swim bladders and later ossified endoskeletons , emerged as the dominant group of fish after
1534-481: The oviparous teleosts, most (79%) do not provide parental care. Viviparity , ovoviviparity , or some form of parental care for eggs, whether by the male, the female, or both parents is seen in a significant fraction (21%) of the 422 teleost families; no care is likely the ancestral condition. The oldest case of viviparity in ray-finned fish is found in Middle Triassic species of † Saurichthys . Viviparity
1593-651: The sister lineage of all other actinopterygians, the Acipenseriformes (sturgeons and paddlefishes) are the sister lineage of Neopterygii, and Holostei (bowfin and gars) are the sister lineage of teleosts. The Elopomorpha ( eels and tarpons ) appear to be the most basal teleosts. The earliest known fossil actinopterygian is Andreolepis hedei , dating back 420 million years ( Late Silurian ), remains of which have been found in Russia , Sweden , and Estonia . Crown group actinopterygians most likely originated near
1652-425: The stout infantfish . 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. 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
1711-412: The " wastebin genus " Barbus has recently been re-evaluated. Though hybrid introgression may confound studies based in mtDNA data alone, a number of these species appear to be so closely related to Labeobarbus as to warrant inclusion in the present genus outright, irrespective of whether Carasobarbus is considered distinct or not. These include L. ethiopicus . Labeobarbus at present contains
1770-944: The Devonian-Carboniferous boundary. The earliest fossil relatives of modern teleosts are from the Triassic period ( Prohalecites , Pholidophorus ), although it is suspected that teleosts originated already during the Paleozoic Era . The listing below is a summary of all extinct (indicated by a dagger , †) and living groups of Actinopterygii with their respective taxonomic rank . The taxonomy follows Phylogenetic Classification of Bony Fishes with notes when this differs from Nelson, ITIS and FishBase and extinct groups from Van der Laan 2016 and Xu 2021. [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] Fish A fish ( pl. : fish or fishes )
1829-610: The Late Paleozoic , evolved from lobe-finned fish during the Carboniferous , developing air-breathing lungs homologous to swim bladders. Despite the cladistic lineage, tetrapods are usually not considered to be fish, making "fish" a paraphyletic group. Fish have been an important natural resource for humans since prehistoric times, especially as food . Commercial and subsistence fishers harvest fish in wild fisheries or farm them in ponds or in breeding cages in
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1888-594: 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
1947-402: The adjacent diagram. The swim bladder is a more derived structure and used for buoyancy . Except from the bichirs , which just like the lungs of lobe-finned fish have retained the ancestral condition of ventral budding from the foregut , the swim bladder in ray-finned fishes derives from a dorsal bud above the foregut. In early forms the swim bladder could still be used for breathing,
2006-556: 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 . The body of
2065-463: The bichirs and holosteans (bowfin and gars) in having gone through a whole-genome duplication ( paleopolyploidy ). The WGD is estimated to have happened about 320 million years ago in the teleosts, which on average has retained about 17% of the gene duplicates, and around 180 (124–225) million years ago in the chondrosteans. It has since happened again in some teleost lineages, like Salmonidae (80–100 million years ago) and several times independently within
2124-550: 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. 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
2183-440: The bulkier, fleshy lobed fins of the sister class Sarcopterygii (lobe-finned fish). Resembling folding fans , the actinopterygian fins can easily change shape and wetted area , providing superior thrust-to-weight ratios per movement compared to sarcopterygian and chondrichthyian fins. The fin rays attach directly to the proximal or basal skeletal elements, the radials, which represent the articulation between these fins and
2242-706: The different actinopterygian clades (in millions of years , mya) are from Near et al., 2012. Jaw-less fishes ( hagfish , lampreys ) [REDACTED] Cartilaginous fishes ( sharks , rays , ratfish ) [REDACTED] Coelacanths [REDACTED] Lungfish [REDACTED] Amphibians [REDACTED] Mammals [REDACTED] Sauropsids ( reptiles , birds ) [REDACTED] Polypteriformes ( bichirs , reedfishes ) [REDACTED] Acipenseriformes ( sturgeons , paddlefishes ) [REDACTED] Teleostei [REDACTED] Amiiformes ( bowfins ) [REDACTED] Lepisosteiformes ( gars ) [REDACTED] The polypterids (bichirs and reedfish) are
2301-561: The exact root is unknown; some authorities reconstruct a Proto-Indo-European root * peysk- , attested only in Italic , Celtic , and Germanic . 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
2360-432: The fish's habit of spending long periods out of water in the mangrove forests it inhabits. Males are occasionally produced at temperatures below 19 °C (66 °F) and can fertilise eggs that are then spawned by the female. This maintains genetic variability in a species that is otherwise highly inbred. Actinopterygii is divided into the subclasses Cladistia , Chondrostei and Neopterygii . The Neopterygii , in turn,
2419-582: The following species : Ray-finned fish Actinopterygii ( / ˌ æ k t ɪ n ɒ p t ə ˈ r ɪ dʒ i aɪ / ; from actino- 'having rays' and Ancient Greek πτέρυξ (ptérux) 'wing, fins'), members of which are known as ray-finned fish or actinopterygians , is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of skin supported by radially extended thin bony spines called lepidotrichia , as opposed to
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2478-405: 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
2537-452: The genus Labeo in size and shape. As far as can be told, all Labeobarbus species are hexaploid . One species, L. microbarbis from Rwanda , is known to have gone extinct in recent times. Like many other "barbs", it was long included in Barbus . It appears to be a fairly close relative of the typical barbels and relatives – the genus Barbus proper – but closer still to
2596-412: 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
2655-532: 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. 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
2714-447: 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
2773-452: 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. Bichirs and lungfish have tetrapod-like paired lungs, requiring them to surface to gulp air, and making them obligate air breathers. Many other fish, including inhabitants of rock pools and
2832-544: The highest mountain streams . Extant species can range in size from Paedocypris , at 8 mm (0.3 in); to the massive ocean sunfish , at 2,300 kg (5,070 lb); and to the giant oarfish , at 11 m (36 ft). The largest ever known ray-finned fish, the extinct Leedsichthys from the Jurassic , has been estimated to have grown to 16.5 m (54 ft). Ray-finned fishes occur in many variant forms. The main features of typical ray-finned fish are shown in
2891-438: The inner part is crossed with fibrous connective tissue. Leptoid scales are thinner and more transparent than other types of scales, and lack the hardened enamel - or dentine -like layers found in the scales of many other fish. Unlike ganoid scales , which are found in non-teleost actinopterygians, new scales are added in concentric layers as the fish grows. Teleosts and chondrosteans (sturgeons and paddlefish) also differ from
2950-425: The internal skeleton (e.g., pelvic and pectoral girdles). The vast majority of actinopterygians are teleosts . By species count, they dominate the subphylum Vertebrata , and constitute nearly 99% of the over 30,000 extant species of fish . They are the most abundant nektonic aquatic animals and are ubiquitous throughout freshwater and marine environments from the deep sea to subterranean waters to
3009-455: 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. 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
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#17328512518123068-541: The large Near Eastern species nowadays separated in Carasobarbus . Barbus has been split to account for the improved phylogenetic knowledge which indicated it was highly paraphyletic in its wide circumscription; it may be that Carasobarbus and some other closely related "barbs" (e.g. Labeobarbus reinii ) are now included in Labeobarbus to avoid a profusion of very small genera. The taxonomy of many species in
3127-411: The main clades of living actinopterygians and their evolutionary relationships to other extant groups of fishes and the four-limbed vertebrates ( tetrapods ). The latter include mostly terrestrial species but also groups that became secondarily aquatic (e.g. whales and dolphins ). Tetrapods evolved from a group of bony fish during the Devonian period . Approximate divergence dates for
3186-419: 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
3245-443: 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. Fish range in size from the huge 16-metre (52 ft) whale shark to some tiny teleosts only 8-millimetre (0.3 in) long, such as the cyprinid Paedocypris progenetica and
3304-496: The ocean. Fish are caught for recreation , or raised by fishkeepers as ornaments for private and public exhibition in aquaria and garden ponds . Fish have had a role in 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 Proto-Germanic , and is related to German Fisch , the Latin piscis and Old Irish īasc , though
3363-402: The open ocean. Because the water all around is the same colour, reflecting an image of the water offers near-invisibility. 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
3422-502: The surrounding water, though some large active swimmers like white shark and tuna can hold a higher core temperature . Many fish can communicate acoustically with each other, such as during courtship displays . The earliest fish appeared during the Cambrian as small filter feeders ; they continued to evolve through the Paleozoic , diversifying into many forms. The earliest fish with dedicated respiratory gills and paired fins ,
3481-415: The taxon Osteichthyes as a clade, which now includes all tetrapods". The biodiversity of extant fish is unevenly distributed among the various groups; teleosts , bony fishes able to protrude their jaws , make up 96% of fish species. The cladogram shows the evolutionary relationships of all groups of living fishes (with their respective diversity ) and the tetrapods. Extinct groups are marked with
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