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Fish
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For fish as eaten by humans, see Fish as food. For the superclass containing the
vast majority of living fish, see Osteichthyes. For other uses, see Fish
(disambiguation).
Fish
Temporal range: 535–0 Ma
PreꞒꞒOSDCPTJKPgN
Middle Cambrian - Recent
Giant grouper swimming among schools of other fish
Giant grouper swimming among schools of other fish
Head-on view of a red lionfish
Head-on view of a red lionfish
Scientific classificationEdit this classification
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
†Conodonts
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.[1][2] Fish can
acoustically communicate with each other, most often in the context of feeding,
aggression or courtship.[3]

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.[4]
With 34,300 described species, fish exhibit greater species diversity than any
other group of vertebrates.[5]

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,[6][7] 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 timeperiods.

Contents
1 Etymology
2 Evolution
2.1 Phylogeny
2.2 Taxonomy
2.3 Diversity
3 Anatomy and physiology
3.1 Respiration
3.1.1 Gills
3.1.2 Air breathing
3.2 Circulation
3.3 Digestion
3.4 Excretion
3.5 Scales
3.6 Sensory and nervous system
3.6.1 Central nervous system
3.6.2 Sense organs
3.6.3 Vision
3.6.4 Hearing
3.6.5 Cognition
3.6.6 Capacity for pain
3.6.7 Emotion
3.7 Muscular system
3.7.1 Endothermy
3.8 Reproductive system
4 Acoustic communication
4.1 Stridulatory
4.2 Non-stridulatory
5 Diseases
5.1 Immune system
6 Conservation
6.1 Overfishing
6.2 Habitat destruction
6.3 Exotic species
7 Importance to humans
7.1 Economic
7.2 Recreation
7.2.1 Fishkeeping
7.2.2 Recreational fishing
7.3 Culture
8 Terminology
8.1 "Fish" or "fishes"
8.2 "True fish" or "finfish"
8.3 "Shoal" or "school"
9 See also
10 Notes
11 References
12 Further reading
13 External links
Etymology
The word for fish in English and the other Germanic languages (German Fisch; Gothic
fisks) 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 an
Proto-Indo-European root *peysk-, attested only in Italic, Celtic, and Germanic.[8]
[9][10][11]

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
Main article: Evolution of fish
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.

Drawing of animal with large mouth, long tail, very small dorsal fins, and pectoral
fins that attach towards the bottom of the body, resembling lizard legs in scale
and development.[12]
Dunkleosteus was a gigantic, 10-metre (33 ft) long prehistoric fish of class
Placodermi.

Lower jaw of the placoderm Eastmanosteus pustulosus, showing the shearing

structures ("teeth") on its oral surface; from the Devonian of Wisconsin
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), although
perhaps the reverse is the case.

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 (†).

Vertebrata/
 
Agnatha/
 
Hyperoartia (lampreys)Nejonöga, Iduns kokbok.jpg

 
 
Myxini (hagfish) Cuvier-120-Myxine116.jpg

 
Cyclostomes
 
 
†EuconodontaConodontZICA.png

 
 
 
†PteraspidomorphiAstraspis desiderata.png

 
 
†ThelodontiSphenonectris turnernae white background.jpg

 
 
 
†AnaspidaLasanius NT small.jpg

 
 
 
†Galeaspida Galeaspids NT.jpg

 
 
 
†PituriaspidaPituriaspida.gif

 
 
†OsteostraciCephalaspis Lyellii.jpg

 
Gnathostomata
 
"†Placodermi" (armoured fishes, paraphyletic)[13]Dunkleosteus intermedius.jpg

 
 
 
"†Acanthodii" ("spiny sharks", paraphyletic or polyphyletic)[14] Diplacanthus
reconstructed.jpg

 
 
Chondrichthyes
 
"†Acanthodii" ("spiny sharks", paraphyletic or polyphyletic)BrochoadmonesDB15.jpg

 
 
 
Holocephali (ratfish)Chimaera monstrosa1.jpg

 
 
Euselachii (sharks, rays)White shark (Duane Raver).pngMobula mobular.jpg

 
 
(cartilaginous fishes)
Euteleostomi/
 
"†Acanthodii" ("spiny sharks", paraphyletic or polyphyletic)Acanthodes BW.jpg

 
 
 
Actinopterygii
 
Cladistia (bichirs, reedfish) Cuvier-105-Polyptère.jpg

 
 
 
Chondrostei (sturgeons, paddlefish)Atlantic sturgeon flipped.jpg

 
 
Neopterygii (includes Teleostei, 96% of living fish species)Common carp (white
background).jpg

 
 
(ray-finned fishes)
Sarcopterygii
 
†Onychodontiformes OnychodusDB15 flipped.jpg

 
 
Actinistia (coelacanths) Coelacanth flipped.png

 
Rhipidistia
 
 
†PorolepiformesReconstruction of Porolepis sp flipped.jpg

 
 
Dipnoi (lungfishes) Barramunda coloured.jpg
 
 
Tetrapodomorpha/
 
 
†RhizodontimorphaGooloogongia loomesi reconstruction.jpg

 
 
 
†TristichopteridaeEusthenodon DB15 flipped.jpg

 
 
 
†TiktaalikTiktaalik BW white background.jpg

 
Tetrapoda
 
†IchthyostegaIchthyostega BW (flipped).jpg

 
 
crown-group tetrapods Salamandra salamandra (white background).jpg British
reptiles, amphibians, and fresh-water fishes (1920) (Lacerta agilis).jpg Meyers
grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908)
(Antwerpener Breiftaube).jpg Ruskea rotta.png

 
four-limbed vertebrates
 
 
 
(Choanata)
 
(lobe-finned fish)
 
 
Osteichthyes
 
 
(jawed vertebrates)
 
 
 
 
"Fishes"
Craniata
Taxonomy

Leedsichthys, of the subclass Actinopterygii, is the largest known fish, with

estimates in 2005 putting its maximum size at 16 metres (52 ft).
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:[15][16]

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 bony fishes)†
Class Osteichthyes (bony fish)
Subclass Actinopterygii (ray finned fishes)
Subclass Sarcopterygii (fleshy 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.[17]
[18]

The various fish groups account for more than half of vertebrate species. As of
2006,[19] 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.[20] Each year, new species are discovered and
scientifically described. As of 2016,[21] 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
Main article: Diversity of fish
Agnatha (Pacific hagfish)
Agnatha
(Pacific hagfish)

Chondrichthyes (Horn shark)

Chondrichthyes
(Horn shark)

Actinopterygii (Brown trout)

Actinopterygii
(Brown trout)

Sarcopterygii (Coelacanth)
Sarcopterygii
(Coelacanth)

Photo of fish with many narrow, straight appendages. Some are end in points, and
others are longer, ending in two or three approximately flat, triangular flaps,
each with a dark spot.
A relative of the seahorses, the leafy seadragon's appendages allow it to
camouflage (in the form of crypsis) with the surrounding seaweed.

The psychedelic mandarin dragonet is one of only two animal species known to have
blue colouring because of cellular pigment.[22]
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.[23] 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.[24][25] 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.[26]
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.[24] 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.[27] 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.[28] 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).[29] There are
even fish that live mostly on land or lay their eggs on land near water.[30]
Mudskippers feed and interact with one another on mudflats and go underwater to
hide in their burrows.[31] A single undescribed species of Phreatobius has been
called a true "land fish" as this worm-like catfish strictly lives among
waterlogged leaf litter.[32][33] Many species live in underground lakes,
underground rivers or aquifers and are popularly known as cavefish.[34]

Fish range in size from the huge 16-metre (52 ft) whale shark to the tiny 8-
millimetre (0.3 in) 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.[35]

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.[36]

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.[21] The following cladogram[37] shows the evolutionary
relationships of all groups of living fishes (with their respective diversity[21])
and the four-limbed vertebrates (tetrapods).

Diversity of various groups of fish (and other vertebrates) through time

Lungfish are the closest living relatives of tetrapods (four-limbed vertebrates).

The bowfin Amia calva is the sole survivor of the halecomorph clade.
Vertebrates
 
Jawless fish Petromyzon marinus Linnaeus 1758 Fig 106 (Matschie et al. 1909).svg
(118 living species: hagfish, lampreys)

 
Jawed vertebrates
 
Cartilaginous fishes Notorynchus cepedianus (Broadnose sevengill shark).gif (>1,100
living species: sharks, rays, chimaeras)

 
Bony fishes
 
Lobe-fin fish
Rhipidistia
 
Tetrapoda Salamender (PSF).png (>30,000 living species: amphibians, mammals,
reptiles, birds)

 
 
Dipnoi Lepidosiren paradoxa2.jpg (6 living species: lungfish)

 
 
 
Actinistia Coelacanth-bgiu.png (2 living species: coelacanths)

 
 
Ray-fin fish
 
Cladistia Polypterus bichir.jpg (14 living species: bichirs, reedfish)

 
Actinopteri
 
Chondrostei Acipenser gueldenstaedtii.jpg (27 living species: sturgeons,
paddlefish)

 
Neopterygii
 
Holostei
 
Ginglymodi Lepisosteus platostomus.jpg (7 living species: gars, alligator gars)

 
 
Halecomorphi Amia calva.jpg (1 living species: bowfin)

 
 
 
Teleostei Engraulis ringens.jpg (>32,000 living species)

 
 
 
 
 
 
 
 
 
Anatomy and physiology
Further information: Fish anatomy and Fish physiology

Organs: 1. Liver, 2. Gas bladder, 3. Roe, 4. Pyloric caeca, 5. Stomach, 6.

Intestine
Respiration
See also: Aquatic respiration
Gills
Photo of fish head split in half longitudinally with gill filaments crossing from
top to bottom
Tuna gills inside the head. The fish head is oriented snout-downwards, with the
view looking towards the mouth.
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,[dubious – discuss] 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.[38] 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 most 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.[38]

Circulation

Didactic model of a fish heart

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.[39] 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
Main article: Fish scale
The scales of fish originate from the mesoderm (skin); they may be similar in
structure to teeth.

Sensory and nervous system

Anatomical diagram showing the pairs of olfactory, telencephalon, and optic lobes,
followed by the cerebellum and the mylencephalon
Dorsal view of the brain of the rainbow trout
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.[40] 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.[41]

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.[40] 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.
[40] 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.[40] The pineal body
lies just above the diencephalon. This structure detects light, maintains circadian
rhythms, and controls color changes.[40]

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.[40]

The hindbrain (or metencephalon) is particularly involved in swimming and balance.

[40] The cerebellum is a single-lobed structure that is typically the biggest part
of the brain.[40] Hagfish and lampreys have relatively small cerebellae, while the
mormyrid cerebellum is massive and apparently involved in their electrical sense.
[40]

The brain stem (or myelencephalon) is the brain's posterior.[40] As well as

controlling some muscles and body organs, in bony fish at least, the brain stem
governs respiration and osmoregulation.[40]

Sense organs
Main article: Sensory systems in fish
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.[42]
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.[43] Some fish, such as catfish and
sharks, have the ampullae of Lorenzini, electroreceptors that detect weak electric
currents on the order of millivolt.[44] 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.[45]

Vision
Main article: Vision in fishes
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.[46] Fish vision
shows adaptation to their visual environment, for example deep sea fishes have eyes
suited to the dark environment.

Hearing
See also: Hearing in fish
Hearing is an important sensory system for most species of fish. Fish sense sound
using their lateral lines and their ears.
Cognition
Further information: Fish intelligence
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.[47]

Wrasses have also passed the mirror test in a 2018 scientific study.[48][49]

Cases of tool use have also been noticed, notably in the Choerodon family, in
archerfish and Atlantic cod.[50]

Capacity for pain

Further information: Pain in fish
Experiments done by William Tavolga provide evidence that fish have pain and fear
responses. For instance, in Tavolga's experiments, toadfish grunted when
electrically shocked and over time they came to grunt at the mere sight of an
electrode.[51]

In 2003, Scottish scientists at the University of Edinburgh and the Roslin

Institute concluded that rainbow trout exhibit behaviors often associated with pain
in other animals. Bee venom and acetic acid injected into the lips resulted in fish
rocking their bodies and rubbing their lips along the sides and floors of their
tanks, which the researchers concluded were attempts to relieve pain, similar to
what mammals would do.[52][53] Neurons fired in a pattern resembling human neuronal
patterns.[53]

Professor James D. Rose of the University of Wyoming claimed the study was flawed
since it did not provide proof that fish possess "conscious awareness, particularly
a kind of awareness that is meaningfully like ours".[54] Rose argues that since
fish brains are so different from human brains, fish are probably not conscious in
the manner humans are, so that reactions similar to human reactions to pain instead
have other causes. Rose had published a study a year earlier arguing that fish
cannot feel pain because their brains lack a neocortex.[55] However, animal
behaviorist Temple Grandin argues that fish could still have consciousness without
a neocortex because "different species can use different brain structures and
systems to handle the same functions."[53]

Animal welfare advocates raise concerns about the possible suffering of fish caused
by angling. Some countries, such as Germany, have banned specific types of fishing,
and the British RSPCA now formally prosecutes individuals who are cruel to fish.
[56]

Emotion
In 2019, scientists have shown that members of the monogamous species Amatitlania
siquia exhibit pessimistic behavior when they are prevented from being with their
partner.[57]

Muscular system
Main article: Fish locomotion

The anatomy of Lampanyctodes hectoris (1) operculum (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 (paired), (10) pectoral fins (paired)
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. 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.

Endothermy
Although most fish are exclusively ectothermic, there are exceptions. The only
known bony fishes (infraclass Teleostei) that exhibit endothermy are in the
suborder Scombroidei – which includes the billfishes, tunas, and the butterfly
kingfish, a basal species of mackerel[58] – and also the opah. The opah, a
lampriform, was demonstrated in 2015 to utilize "whole-body endothermy", generating
heat with its swimming muscles to warm its body while countercurrent exchange (as
in respiration) minimizes heat loss.[59] It is able to actively hunt prey such as
squid and swim for long distances due to the ability to warm its entire body,
including its heart,[60] which is a trait typically found in only mammals and birds
(in the form of homeothermy). In the cartilaginous fishes (class Chondrichthyes),
sharks of the families Lamnidae (porbeagle, mackerel, salmon, and great white
sharks) and Alopiidae (thresher sharks) exhibit endothermy. 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 in excess of
20 °C (68 °F) above ambient water temperatures.[58]

Endothermy, though metabolically costly, is thought to provide advantages such as

increased muscle strength, higher rates of central nervous system processing, and
higher rates of digestion.

Reproductive system
Further information: Fish reproduction and Spawn (biology)

Ovary of fish (Corumbatá)

Fish reproductive organs include testicles and ovaries. In most species, gonads are
paired organs of similar size, which can be partially or totally fused.[61] There
may also be a range of 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.[61]

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.[62] Gymnovaries are the primitive condition found in lungfish,
sturgeon, and bowfin. Cystovaries characterize most teleosts, where the ovary lumen
has continuity with the oviduct.[61] 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.[61]

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.[61]

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 all known fish are oviparous,[63] 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).

Egg of lamprey
Egg of lamprey

Egg of catshark (mermaids' purse)

Egg of catshark (mermaids' purse)

Egg of bullhead shark

Egg of bullhead shark

Egg of chimaera
Egg of chimaera

The newly hatched young of oviparous fish are called larvae. They are usually
poorly formed, carry a large yolk sac (for nourishment), and are very different in
appearance from juvenile and adult specimens. The larval period in oviparous fish
is relatively short (usually only several weeks), and larvae rapidly grow and
change appearance and structure (a process termed metamorphosis) 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.[64] 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.[64]

Aquarists commonly refer to ovoviviparous and viviparous fish as livebearers.

Acoustic communication
See also: Acoustic communication in aquatic animals
Acoustic communication in fish involves the transmission of acoustic signals from
one individual of a species to another. The production of sounds as a means of
communication among fish is most often used in the context of feeding, aggression
or courtship behaviour.[3] The sounds emitted can vary depending on 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.[65]

Stridulatory

French grunts – Haemulon flavolineatum

There are some species of fish that can produce sounds by rubbing or grinding their
bones together. These noises produced by bone-on-bone interactions are known as
'stridulatory sounds'.[65]

An example of this is seen in Haemulon flavolineatum, a species commonly referred

to as the 'French grunt fish', as it produces a grunting noise by grinding its
teeth together.[65] This behaviour is most pronounced when the H. flavolineatum is
in distress situations.[65] The grunts produced by this species of fishes generate
a frequency of approximately 700 Hz, and last approximately 47 milliseconds.[65]
The H. flavolineatum does not emit sounds with frequencies greater than 1000 Hz,
and does not detect sounds that have frequencies greater than 1050 Hz.[65]

In a study conducted by Oliveira et al. (2014), the longsnout seahorse, Hippocampus

reidi, was recorded producing two different categories of sounds; 'clicks' and
'growls'. The sounds emitted by the H. reidi are accomplished by rubbing their
coronet bone across the grooved section of their neurocranium.[66] 'Clicking'
sounds were found to be primarily produced during courtship and feeding, and the
frequencies of clicks were within the range of 50 Hz-800 Hz.[67] The frequencies
were noted to be on the higher end of the range during spawning periods, when the
female and male fishes were less than fifteen centimeters apart.[67] Growl sounds
were produced when the H. reidi encountered stressful situations, such as handling
by researchers.[67] The 'growl' sounds consist of a series of sound pulses and are
emitted simultaneously with body vibrations.[67]

Non-stridulatory

Oyster toadfish
Some fish species create noise by engaging specialized muscles that contract and
cause swimbladder vibrations.

Oyster toadfish produce loud grunting sounds by contracting muscles located along
the sides of their swim bladder, known as sonic muscles [68] Female and male
toadfishes emit short-duration grunts, often as a fright response.[69] In addition
to short-duration grunts, male toadfishes produce "boat whistle calls".[70] These
calls are longer in duration, lower in frequency, and are primarily used to attract
mates.[70] The sounds emitted by the O. tao have frequency range of 140 Hz to 260
Hz.[70] The frequencies of the calls depend on the rate at which the sonic muscles
contract.[71][68]

The red drum, Sciaenops ocellatus, produces drumming sounds by vibrating its
swimbladder.[72] Vibrations are caused by the rapid contraction of sonic muscles
that surround the dorsal aspect of the swimbladder.[72] These vibrations result in
repeated sounds with frequencies that range from 100 to >200 Hz.[72] The S.
ocellatus can produce different calls depending on the stimuli involved.[72] The
sounds created in courtship situations are different from those made during
distressing events such as predatorial attacks.[72] Unlike the males of the S.
ocellatus species, the females of this species do not produce sounds and lack
sound-producing (sonic) muscles.[72]

Diseases
Main article: Fish diseases and parasites
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.[73]
In recent years, vaccines have become widely used in aquaculture and also with
ornamental fish, for example furunculosis vaccines in farmed salmon and koi herpes
virus in koi.[74][75]

Some species use cleaner fish to remove external parasites. The best known of these
are the bluestreak cleaner wrasses of the genus Labroides found on coral reefs in
the Indian and Pacific oceans. These small fish maintain so-called "cleaning
stations" where other fish congregate and perform specific movements to attract the
attention of the cleaners.[76] 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.[77]

Immune system
Immune organs vary by type of fish.[78] In the jawless fish (lampreys and hagfish),
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 (or pronephros) and some areas
of the gut (where granulocytes mature.) They resemble primitive bone marrow in
hagfish. Cartilaginous fish (sharks and rays) have a more advanced immune system.
They have three specialized organs that are unique to Chondrichthyes; the epigonal
organs (lymphoid tissue similar to mammalian bone) that surround the gonads, the
Leydig's organ within the walls of their esophagus, and a spiral valve in their
intestine. These organs house typical immune cells (granulocytes, lymphocytes and
plasma cells). They also possess an identifiable thymus and a well-developed spleen
(their most important immune organ) where various lymphocytes, plasma cells and
macrophages develop and are stored. Chondrostean fish (sturgeons, paddlefish, and
bichirs) possess a major site for the production of granulocytes within a mass that
is associated with the meninges (membranes surrounding the central nervous system.)
Their heart is frequently covered with tissue that contains lymphocytes, reticular
cells and a small number of macrophages. The chondrostean kidney is an important
hemopoietic organ; where erythrocytes, granulocytes, lymphocytes and macrophages
develop.

Like chondrostean fish, the major immune tissues of bony fish (or teleostei)
include the kidney (especially the anterior kidney), which houses many different
immune cells.[79] 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.[80][81] In 2006, a lymphatic system similar to that
in mammals was described in one species of teleost fish, the zebrafish. Although
not confirmed as yet, this system presumably will be where naive (unstimulated) T
cells accumulate while waiting to encounter an antigen.[82]

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.[83]

Conservation
The 2006 IUCN Red List names 1,173 fish species that are threatened with
extinction.[84] Included are species such as Atlantic cod,[85] Devil's Hole
pupfish,[86] coelacanths,[87] and great white sharks.[88] 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.[89]

Overfishing
Photo of shark in profile surrounded by other, much smaller fish in bright sunlight
Whale sharks, the largest species of fish, are classified as vulnerable.
Main article: Overfishing
Overfishing is a major threat to edible fish such as cod and tuna.[90][91]
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.

One 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.[92]

The main tension between fisheries science and the fishing industry is that the two
groups 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.[93][94] On the other hand,
scientists and conservationists push for stringent protection, warning that many
stocks could be wiped out within fifty years.[95][96]

Habitat destruction
See also: 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.[97] 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.[98]

Exotic species
Introduction of non-native species has occurred in many habitats. One of the best
studied examples is the introduction of Nile perch into Lake Victoria in the 1960s.
Nile perch gradually exterminated the lake's 500 endemic cichlid species. Some of
them survive now in captive breeding programmes, but others are probably extinct.
[99] Carp, snakeheads,[100] tilapia, European perch, brown trout, rainbow trout,
and sea lampreys are other examples of fish that have caused problems by being
introduced into alien environments.
Importance to humans
Economic
Main articles: Fishing industry, Aquaculture, and Fish farming
A row of square artificial ponds, with trees on either side
These fish-farming ponds were created as a cooperative project in a rural village.
Throughout history, humans have utilized fish as a food source. Historically and
today, most fish protein has come by means of catching wild fish. However,
aquaculture, or fish farming, which has been practiced since about 3,500 BCE. in
China,[101] is becoming increasingly important in many nations. Overall, about one-
sixth of the world's protein is estimated to be provided by fish.[102] That
proportion is considerably elevated in some developing nations and regions heavily
dependent on the sea. In a similar manner, fish have been tied to trade.

Fish counter display at the Oulu Market Hall in Oulu, Finland.

Catching fish for the purpose of food or sport is known as fishing, while the
organized effort by humans to catch fish is called a fishery. Fisheries are a huge
global business and provide income for millions of people.[102] The annual yield
from all fisheries worldwide is about 154 million tons,[103] with popular species
including herring, cod, anchovy, tuna, flounder, and salmon. However, the term
fishery is broadly applied, and includes more organisms than just fish, such as
mollusks and crustaceans, which are often called "fish" when used as food.

Recreation

A Bengali fish vendor

Main articles: Fishkeeping, Recreational fishing, and Angling
Fishkeeping
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
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.[104][105] Angling is a method of fishing, specifically the practice of
catching fish by means of an "angle" (hook). Anglers must select the right hook,
cast accurately, and retrieve at the right speed while considering water and
weather conditions, species, fish response, time of the day, and other factors.

Culture
Main article: Fish in culture
Photo of painting showing blue-skinned, 4-armed upper body of man standing in the
opened mouth of a fish with bent tail with other, paler men facing him with hands
raised together
Avatar of Vishnu as a Matsya
Fish themes have symbolic significance in many religions. In ancient Mesopotamia,
fish offerings were made to the gods from the very earliest times.[106] Fish were
also a major symbol of Enki, the god of water.[106] 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.[106] 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.[106]
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.[106]
Fish were sacred to the Syrian goddess Atargatis[107] and, during her festivals,
only her priests were permitted to eat them.[107]

The ichthus is a Christian symbol of a fish signifying that the person who uses it
is a Christian.[107][108]
In the Book of Jonah, a work of Jewish literature probably written in the fourth
century BC, 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.[109][110]
[111] The fish later vomits Jonah out on shore after three days.[109][110][111]
This book was later included as part of the Hebrew Bible, or Christian Old
Testament,[112][113] and a version of the story it contains is summarized in Surah
37:139-148 of the Quran.[114] Early Christians used the ichthys, a symbol of a
fish, to represent Jesus,[107][108] 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".[107]
[108] The gospels also refer to "fishers of men"[115] and feeding the multitude. In
the dhamma of Buddhism, the fish symbolize happiness as they have complete freedom
of movement in the water. Often drawn in the form of carp which are regarded in the
Orient as sacred on account of their elegant beauty, size and life-span.

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.[116]

Fish feature prominently in art and literature, 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, most notably the novel
Jaws, which spawned a series of films of the same name that in turn inspired
similar films or parodies such as Shark Tale and Snakehead Terror. Piranhas are
shown in a similar light to sharks in films such as Piranha; however, contrary to
popular belief, the red-bellied piranha is actually a generally timid scavenger
species that is unlikely to harm humans.[117] Legends of half-human, half-fish
mermaids have featured in folklore, including the stories of Hans Christian
Andersen.

Terminology
"Fish" or "fishes"
Though often used interchangeably, in biology these words have different meanings.
Fish is used as a singular noun, or as a plural to describe multiple individuals
from a single species. Fishes is used to describe different species or species
groups.[118][119][120] Thus a pond would be said to contain 120 fish if all were
from a single species or 120 fishes if these included a mix of several species. The
distinction is similar to that between people and peoples.

"True fish" or "finfish"

In biology, the term fish is most strictly used to describe any animal with a
backbone, gills throughout life, and limbs (if any) in the shape of fins.[23] Many
types of aquatic animals with common names ending in "fish" are not fish in this
sense; 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.[26]
In fisheries, the term fish is used as a collective term, and includes mollusks,
crustaceans and any aquatic animal which is harvested.[121]
The strict biological definition of a fish, above, is sometimes called a true fish.
True fish are also referred to as finfish or fin fish to distinguish them from
other aquatic life harvested in fisheries or aquaculture.
"Shoal" or "school"
Main article: Shoaling and schooling
Photo of thousands of fish separated from each other by distances of 2 inches (51
mm) or less
These goldband fusiliers are schooling because their swimming is synchronised.
An assemblage of fish merely using some localised resource such as food or nesting
sites is known simply as an aggregation. When fish come together in an interactive,
social grouping, then they may be forming either a shoal or a school depending on
the degree of organisation. 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. Schools of fish are much more tightly organised,
synchronising their swimming so that all fish move at the same speed and in the
same direction. Shoaling and schooling behaviour is believed to provide a variety
of advantages.[122]

Examples:

Cichlids congregating at lekking sites form an aggregation.

Many minnows and characins form shoals.
Anchovies, herrings and silversides are classic examples of schooling fish.
The most common collective nouns for a group of fish in general are school and
shoal. Both the words have evolved from the same common Dutch root 'schole' meaning
a troop or crowd. While the words "school" and "shoal" have different meanings
within biology, the distinctions are often ignored by non-specialists who treat the
words as synonyms. Thus speakers of British English commonly use "shoal" to
describe any grouping of fish, and speakers of American English commonly use
"school" just as loosely.[123]

See also
Fish portal
Main article: Outline of fish
For a topical guide to sharks, see Outline of sharks.
Angling (sport fishing)
Aquaculture
Aquarium
Catch and release
Deep sea fish
Fish acute toxicity syndrome
Fish anatomy
Fish as food
Fish development
Fishing (fishing for food)
Fishkeeping
Forage fish
Ichthyology
List of fish common names
List of fish families
Marine biology
Marine vertebrates
Mercury in fish
Otolith (Bone used for determining the age of a fish)
Pregnancy (fish)
Seafood
Shoaling and schooling
Walking fish
Notes
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References
Helfman, G.; Collette, B.; Facey, D. (1997). The Diversity of Fishes (1st ed.).
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Further reading
Eschmeyer, William N.; Fong, Jon David (2013). "Catalog of Fishes". California
Academy of Sciences.
Helfman, G.; Collette, B.; Facey, D.; Bowen, B. (2009). The Diversity of Fishes:
Biology, Evolution, and Ecology (2nd ed.). Wiley-Blackwell. ISBN 978-1-4051-2494-2.
Moyle, Peter B. (1993) Fish: An Enthusiast's Guide University of California Press.
ISBN 978-0-520-91665-4 – good lay text.
Moyle, Peter B.; Cech, Joseph J. (2003). Fishes, An Introduction to Ichthyology
(5th ed.). Benjamin Cummings. ISBN 978-0-13-100847-2.
Scales, Helen (2018). Eye of the shoal: A Fishwatcher's Guide to Life, the Ocean
and Everything. Bloomsbury Sigma. ISBN 978-1-4729-3684-4.
Shubin, Neil (2009). Your inner fish: A journey into the 3.5 billion year history
of the human body. Vintage Books. ISBN 978-0-307-27745-9. UCTV interview
External links

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Wikisource has the text of the 1911 Encyclopædia Britannica article "Fish".
ANGFA – Illustrated database of freshwater fishes of Australia and New Guinea
Fischinfos.de – Illustrated database of the freshwater fishes of Germany at the
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FishBase online – Comprehensive database with information on over 29,000 fish
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Fisheries and Illinois Aquaculture Center – Data outlet for fisheries and
aquaculture research center in the central US at archive.today (archived 15
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United Nation – Fisheries and Aquaculture Department: Fish and seafood utilization
University of Washington Libraries Digital Collections – Digital collection of
freshwater and marine fish images
Davenport, Charles B.; Ingersoll, Ernest (1905). "Fish" . New International
Encyclopedia.
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Predatory billfishmackerelsalmontunaForage anchovyherringsardinespratsDemersal
codflatfishpollock
Major groups
Jawless hagfishlampreysCartilaginous chimaerassharksraysBony spiny-finnedfleshy-
finned
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Aquarium lifeBlindFish common namesFish familiesGlossary of
ichthyologyLargestSmallestThreatened rayssharksPrehistoric more lists...
Category WikiProject
vte
Evolution of fish

Taxon identifiers
Wikidata: Q152Wikispecies: PiscesAFD: PiscesPlazi: 21430E9C-6188-A40A-2DCF-
177F0B48AFD2WoRMS: 11676ZooBank: E48F627B-CFB0-4C41-8B2B-2DDA1EEF39D2
Authority control Edit this at Wikidata
Categories: FishAquatic ecologyFishingIchthyologySeafoodObsolete vertebrate
taxaParaphyletic groups
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This page was last edited on 7 August 2022, at 09:37 (UTC).
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