Fish evolution.png

Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of living fish species are ray-finned fish, belonging to the class Actinopterygii, with around 99% of those being teleosts.

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 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 no species has yet been documented in the deepest 25% of the ocean. With 34,300 described species, fish exhibit greater species diversity than any other group of vertebrates.

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 (in aquaculture). 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.

Tetrapods ( amphibians, reptiles, birds and mammals) emerged within lobe-finned fishes, so cladistically they are fish as well. However, traditionally fish (pisces or ichthyes) are rendered paraphyletic by excluding the tetrapods, and are therefore not considered a formal taxonomic grouping in systematic biology, unless it is used in the cladistic sense, including tetrapods, although usually "vertebrate" is preferred and used for this purpose (fish plus tetrapods) instead. Furthermore, cetaceans, although mammals, have often been considered fish by various cultures and time periods.

Etymology

The word for ''fish'' in English and the other Germanic languages (German ; Gothic ) is inherited from Proto-Germanic, and is related to 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.

The English word once had a much broader usage than its current biological meaning. Names such as starfish, jellyfish, shellfish and cuttlefish attest to almost any fully aquatic animal (including whales) once being ''fish''. "Correcting" such names (e.g. to ''sea star'') is an attempt to retroactively apply the current meaning of ''fish'' to words that were coined when it had a different meaning.

Evolution

Fish, as vertebrata, developed as sister of the tunicata. As the tetrapods emerged deep within the fishes group, as sister of the lungfish, characteristics of fish are typically shared by tetrapods, including having vertebrae and a cranium.

Early fish from the fossil record are represented by a group of small, jawless, armored fish known as ostracoderms. Jawless fish lineages are mostly extinct. An extant clade, the lampreys may approximate ancient pre-jawed fish. The first jaws are found in Placodermi fossils. They lacked distinct teeth, having instead the oral surfaces of their jaw plates modified to serve the various purposes of teeth. The diversity of jawed vertebrates may indicate the evolutionary advantage of a jawed mouth. It is unclear if the advantage of a hinged jaw is greater biting force, improved respiration, or a combination of factors.

Fish may have evolved from a creature similar to a coral-like sea squirt, whose larvae resemble primitive fish in important ways. The first ancestors of fish may have kept the larval form into adulthood (as some sea squirts do today).

Phylogeny

Fishes are a paraphyletic group: that is, any clade containing all fish also contains the tetrapods, which are not fish (though they include fish-shaped forms, such as Whales and Dolphins or the extinct ichthyosaurs, which acquired a fish-like body shape due to secondary aquatic adaptation, see evolution of cetaceans).

The following cladogram shows clades - some with, some without extant relatives - that are traditionally considered as "fishes" ( cyan line) and the tetrapods (four-limbed vertebrates), which are mostly terrestrial. Extinct groups are marked with a dagger (†).

Taxonomy

Fishes are a paraphyletic group and for this reason, groups such as the class ''Pisces'' seen in older reference works are no longer used in formal classifications. Traditional classification divides fish into three extant classes, and with extinct forms sometimes classified within the tree, sometimes as their own classes: Romer, A.S. & T.S. Parsons. 1977. ''The Vertebrate Body.'' 5th ed. Saunders, Philadelphia. (6th ed. 1985)
* Class Agnatha (jawless fish)
** Subclass Cyclostomata (hagfish and lampreys)
** Subclass Ostracodermi (armoured jawless fish) †
* Class Chondrichthyes (cartilaginous fish)
** Subclass Elasmobranchii (sharks and rays)
** Subclass Holocephali ( chimaeras and extinct relatives)
* Class Placodermi (armoured fish) †
* Class Acanthodii ("spiny sharks", sometimes classified under Actinopterygii)
* Class Osteichthyes (bony fish)
** Subclass Actinopterygii (ray finned fishes)
** Subclass Sarcopterygii (lobe finned fishes, ancestors of tetrapods)

The above scheme is the one most commonly encountered in non-specialist and general works. Many of the above groups are paraphyletic, in that they have given rise to successive groups: Agnathans are ancestral to Chondrichthyes, who again have given rise to Acanthodiians, the ancestors of Osteichthyes. With the arrival of phylogenetic nomenclature, the fishes has been split up into a more detailed scheme, with the following major groups:
* Class Myxini (hagfish)
* Class Pteraspidomorphi † (early jawless fish)
* Class Thelodonti †
* Class Anaspida †
* Class Petromyzontida or Hyperoartia
** Petromyzontidae (lampreys)
* Class Conodonta (conodonts) †
* Class Cephalaspidomorphi † (early jawless fish)
** (unranked) Galeaspida †
** (unranked) Pituriaspida †
** (unranked) Osteostraci †
* Infraphylum Gnathostomata (jawed vertebrates)
** Class Placodermi † (armoured fish)
** Class Chondrichthyes (cartilaginous fish)
** Class Acanthodii † (spiny sharks)
** Superclass Osteichthyes (bony fish)
*** Class Actinopterygii (ray-finned fish)
**** Subclass Chondrostei
***** Order Acipenseriformes ( sturgeons and paddlefishes)
***** Order Polypteriformes (reedfishes and bichirs).
**** Subclass Neopterygii
***** Infraclass Holostei (gars and bowfins)
***** Infraclass Teleostei (many orders of common fish)
*** Class Sarcopterygii (lobe-finned fish)
**** Subclass Actinistia (coelacanths)
**** Subclass Dipnoi (lungfish, sister group to the tetrapods)

† – indicates extinct taxon
Some palaeontologists contend that because Conodonta are chordates, they are primitive fish. For a fuller treatment of this taxonomy, see the vertebrate article.

The position of hagfish in the phylum Chordata is not settled. Phylogenetic research in 1998 and 1999 supported the idea that the hagfish and the lampreys form a natural group, the Cyclostomata, that is a sister group of the Gnathostomata.

The various fish groups account for more than half of vertebrate species. As of 2006, there are almost 28,000 known extant species, of which almost 27,000 are bony fish, with 970 sharks, rays, and chimeras and about 108 hagfish and lampreys. A third of these species fall within the nine largest families; from largest to smallest, these families are Cyprinidae, Gobiidae, Cichlidae, Characidae, Loricariidae, Balitoridae, Serranidae, Labridae, and Scorpaenidae. About 64 families are monotypic, containing only one species. The final total of extant species may grow to exceed 32,500. Each year, new species are discovered and scientifically described. As of 2016, there are over 32,000 documented species of bony fish and over 1,100 species of cartilaginous fish. Species are lost through extinction (see biodiversity crisis). Recent examples are the Chinese paddlefish or the smooth handfish.

Diversity

Pacific_hagfish_Myxine.jpg, Agnatha
( Pacific hagfish)
Hornhai (Heterodontus francisci).JPG, Chondrichthyes
(Horn shark)
Salmo trutta.jpg, Actinopterygii
( Brown trout)
Latimeria chalumnae01.jpg, Sarcopterygii
(Coelacanth)

The term "fish" most precisely describes any non-tetrapod craniate (i.e. an animal with a skull and in most cases a backbone) that has gills throughout life and whose limbs, if any, are in the shape of fins. Unlike groupings such as birds or mammals, fish are not a single clade but a paraphyletic collection of taxa, including hagfishes, lampreys, sharks and rays, ray-finned fish, coelacanths, and lungfish. Indeed, lungfish and coelacanths are closer relatives of tetrapods (such as mammals, birds, amphibians, etc.) than of other fish such as ray-finned fish or sharks, so the last common ancestor of all fish is also an ancestor to tetrapods. As paraphyletic groups are no longer recognised in modern systematic biology, the use of the term "fish" as a biological group must be avoided.

Many types of aquatic animals commonly referred to as "fish" are not fish in the sense given above; examples include shellfish, cuttlefish, starfish, crayfish and jellyfish. In earlier times, even biologists did not make a distinction – sixteenth century natural historians classified also seals, whales, amphibians, crocodiles, even hippopotamuses, as well as a host of aquatic invertebrates, as fish. However, according to the definition above, all mammals, including cetaceans like whales and dolphins, are not fish. In some contexts, especially in aquaculture, the true fish are referred to as finfish (or fin fish) to distinguish them from these other animals.

A typical fish is ectothermic, has a streamlined body for rapid swimming, extracts oxygen from water using gills or uses an accessory breathing organ to breathe atmospheric oxygen, has two sets of paired fins, usually one or two (rarely three) dorsal fins, an anal fin, and a tail fin, has jaws, has skin that is usually covered with scales, and lays eggs.

Each criterion has exceptions. Tuna, swordfish, and some species of sharks show some warm-blooded adaptations – they can heat their bodies significantly above ambient water temperature. 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. Many groups of freshwater fish extract oxygen from the air as well as from the water using a variety of different structures. Lungfish have paired lungs similar to those of tetrapods, gouramis have a structure called the labyrinth organ that performs a similar function, while many catfish, such as ''Corydoras'' extract oxygen via the intestine or stomach. Body shape and the arrangement of the fins is highly variable, covering such seemingly un-fishlike forms as seahorses, pufferfish, anglerfish, and gulpers. Similarly, the surface of the skin may be naked (as in moray eels), or covered with scales of a variety of different types usually defined as placoid (typical of sharks and rays), cosmoid (fossil lungfish and coelacanths), ganoid (various fossil fish but also living gars and bichirs), cycloid, and ctenoid (these last two are found on most bony fish). There are even fish that 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. Many species live in underground lakes, underground rivers or aquifers and are popularly known as cavefish.

Fish range in size from the huge whale shark to the tiny stout infantfish.

Fish species diversity is 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.

The deepest living fish in the ocean so far found is the Mariana snailfish (''Pseudoliparis swirei'') which lives at deeps of 8,000 meters (26,200 feet) along the Mariana Trench near Guam.

The diversity of living fish (finfish) is unevenly distributed among the various groups, with teleosts making up the bulk of living fishes (96%), and over 50% of all vertebrate species. The following cladogram shows the evolutionary relationships of all groups of living fishes (with their respective diversity) and the four-limbed vertebrates (tetrapods).

Anatomy and physiology

Respiration

Gills

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 very primitive feature that they share 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 live and move about on land for up to several days, or live in stagnant or otherwise oxygen-depleted water. Many such fish can breathe air via a variety of mechanisms. 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. Lungfish, with the exception of the Australian lungfish, and bichirs have paired lungs similar to those of tetrapods and must 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). A number of fish have evolved so-called accessory breathing organs that extract oxygen from the air. Labyrinth fish (such as gouramis and bettas) have a labyrinth organ above the gills that performs this function. A few other fish have structures resembling labyrinth organs in form and function, most notably snakeheads, pikeheads, and the Clariidae catfish family.

Breathing air is primarily of use to fish that inhabit shallow, seasonally variable waters where the water's oxygen concentration may seasonally decline. Fish dependent solely on dissolved oxygen, such as perch and cichlids, quickly suffocate, while air-breathers survive for much longer, in some cases in water that is little more than wet mud. At the extreme, 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 the catfish ''Hypostomus plecostomus'', only breathe air if they need to and will otherwise rely on their gills for oxygen. Most air breathing fish are facultative air breathers that avoid the energetic cost of rising to the surface and the fitness cost of exposure to surface predators.

Circulation

Fish have a closed-loop circulatory system. The heart pumps the blood in a single loop throughout the body. In most fish, the heart consists of four parts, including two chambers and an entrance and exit. 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

As with many aquatic animals, most fish release their nitrogenous wastes as ammonia. Some of the wastes diffuse through the gills. Blood wastes are filtered by the kidneys.

Saltwater fish tend to lose water because of osmosis. Their kidneys return water to the body. The reverse happens in freshwater fish: they tend to gain water osmotically. Their kidneys produce dilute urine for excretion. Some fish have specially adapted kidneys that vary in function, allowing them to move from freshwater to saltwater.

Scales

The scales of fish originate from the mesoderm (skin); they may be similar in structure to teeth.

Sensory and nervous system

Central nervous system

Fish typically have quite 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, most notably mormyrids and sharks, which have brains about as massive relative to body weight as birds and marsupials.

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. 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.

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. The pineal body lies just above the diencephalon. This structure detects light, maintains circadian rhythms, and controls color changes.

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.

The hindbrain (or metencephalon) is particularly involved in swimming and balance. The cerebellum is a single-lobed structure that is typically the biggest part of the brain. Hagfish and lampreys have relatively small cerebellae, while the mormyrid cerebellum is massive and apparently involved in their electrical sense.

The brain stem (or myelencephalon) is the brain's posterior. As well as controlling some muscles and body organs, in bony fish at least, the brain stem governs respiration and osmoregulation.

Sense organs

Most fish possess highly developed sense organs. Nearly all daylight fish have color vision that is at least as good as a human's (see vision in fishes). Many fish also have chemoreceptors that are responsible for extraordinary senses of taste and smell. Although they have ears, many fish may not hear very well. Most fish have sensitive receptors that form the lateral line system, which detects gentle currents and vibrations, and senses the motion of nearby fish and prey. The sense information obtained from the lateral line system can be considered both a sense of touch and 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. Other fish, like the South American electric fishes Gymnotiformes, can produce weak electric currents, which they use in navigation and social communication.

Fish orient themselves using landmarks and may use mental maps based on multiple landmarks or symbols. Fish behavior in mazes reveals that they possess spatial memory and visual discrimination.

Vision

Vision is an important sensory system for most species of fish. 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), and most species have colour vision. Some fish can see ultraviolet and some can see polarized light. Amongst jawless fish, the lamprey has well-developed eyes, while the hagfish has only primitive eyespots. Fish vision shows adaptation to their visual environment, for example deep sea fishes have eyes suited to the dark environment.

Hearing

Hearing is an important sensory system for most species of fish. Fish sense sound using their lateral lines and their ears.

Cognition

New research has expanded preconceptions about the cognitive capacities of fish. For example, manta rays have exhibited behavior linked to self-awareness in mirror test cases. Placed in front of a mirror, individual rays engaged in contingency testing, that is, repetitive behavior aiming to check whether their reflection's behavior mimics their body movement.

Wrasses have also passed the mirror test in a 2018 scientific study.

Cases of tool use have also been noticed, notably in the Choerodon family, in archerfish and Atlantic cod

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