Fish

For other uses, see Fish (disambiguation).

religious symbols, and as the subjects of art, books and

movies.

A sh is any member of a group of organisms that

consist of all gill-bearing aquatic craniate animals that
lack limbs with digits. They form a sister group to 1 Evolution
the tunicates, together forming the olfactores. Included
in this denition are the living hagsh, lampreys, and Main article: Evolution of sh
cartilaginous and bony sh as well as various extinct Fish do not represent a monophyletic group, and thererelated groups. Tetrapods emerged within lobe-nned
shes, so cladistically they are sh as well. However, traditionally sh are rendered obsolete or paraphyletic by excluding the tetrapods (i.e., the amphibians, reptiles, birds
and mammals which all descended from within the same
ancestry). Because in this manner the term sh is dened negatively as a paraphyletic group, it is not considered a formal taxonomic grouping in systematic biology.
The traditional term pisces (also ichthyes) is considered
a typological, but not a phylogenetic classication.
The earliest organisms that can be classied as sh
were soft-bodied chordates that rst 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 sh of the
Paleozoic developed external armor that protected them
from predators. The rst sh with jaws appeared in the
Silurian period, after which many (such as sharks) became formidable marine predators rather than just the
prey of arthropods.

Dunkleosteus was a gigantic, 10-metre (33 ft) long prehistoric

sh of class Placodermi.

fore the evolution of sh is not studied as a single

event.[4]
Early sh from the fossil record are represented by
a group of small, jawless, armored sh known as
ostracoderms. Jawless sh lineages are mostly extinct.
An extant clade, the lampreys may approximate ancient
pre-jawed sh. The rst jaws are found in Placodermi
fossils. 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.

Most sh 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 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.,
gulpers and anglersh). With 33,100 described species,
sh exhibit greater species diversity than any other group
of vertebrates.[3]

Fish may have evolved from a creature similar to a corallike Sea squirt, whose larvae resemble primitive sh in
important ways. The rst ancestors of sh may have kept
the larval form into adulthood (as some sea squirts do today), although perhaps the reverse is the case.

Fish are an important resource for humans worldwide,

especially as food. Commercial and subsistence shers
hunt sh in wild sheries (see shing) or farm them in
ponds or in cages in the ocean (see aquaculture). They
are also caught by recreational shers, kept as pets, raised
by shkeepers, and exhibited in public aquaria. Fish have
had a role in culture through the ages, serving as deities,

1.1 Taxonomy
Fish are a paraphyletic group: that is, any clade containing all sh also contains the tetrapods, which are not sh.
For this reason, groups such as the Class Pisces seen in
older reference works are no longer used in formal classications.
1

1 EVOLUTION

Traditional classication divide sh into three extant

classes, and with extinct forms sometimes classied
within the tree, sometimes as their own classes:[5][6]
Class Agnatha (jawless sh)
Subclass Cyclostomata (hagsh and lampreys)
Subclass Ostracodermi (armoured jawless
sh)
Class Chondrichthyes (cartilaginous sh)
Subclass Elasmobranchii (sharks and rays)
Subclass Holocephali (chimaeras and extinct
relatives)

Class Thelodonti
Class Anaspida
Class Petromyzontida or Hyperoartia
Petromyzontidae (lampreys)
Class Conodonta (conodonts)
Class Cephalaspidomorphi (early jawless sh)
(unranked) Galeaspida
(unranked) Pituriaspida
(unranked) Osteostraci
Infraphylum Gnathostomata (jawed vertebrates)

Class Placodermi (armoured sh)

Class Placodermi (armoured sh)

Class Acanthodii (spiny sharks, sometimes classied under bony shes)

Class Chondrichthyes (cartilaginous sh)

Class Acanthodii (spiny sharks)
Superclass Osteichthyes (bony sh)

Leedsichthys (left), of the subclass Actinopterygii, is the largest

known sh, with estimates in 2005 putting its maximum size at
16 metres (52 ft)

Class Osteichthyes (bony sh)

Subclass Actinopterygii (ray nned shes)

Class Actinopterygii (ray-nned sh)

Subclass Chondrostei
Order Acipenseriformes (sturgeons
and paddleshes)
Order Polypteriformes (reedshes
and bichirs).
Subclass Neopterygii
Infraclass Holostei (gars and
bowns)
Infraclass Teleostei (many orders of
common sh)
Class Sarcopterygii (lobe-nned sh)
Subclass Actinistia (coelacanths)
Subclass Dipnoi (lungsh)
indicates extinct taxon
Some palaeontologists contend that because Conodonta
are chordates, they are primitive sh. For a fuller treatment of this taxonomy, see the vertebrate article.

Subclass Sarcopterygii (eshy nned shes, The position of hagsh in the phylum Chordata is not settled. Phylogenetic research in 1998 and 1999 supported
ancestors of tetrapods)
the idea that the hagsh and the lampreys form a natural group, the Cyclostomata, that is a sister group of the
The above scheme is the one most commonly encounGnathostomata.[7][8]
tered in non-specialist and general works. Many of the
above groups are paraphyletic, in that they have given The various sh groups account for more than half of verrise to successive groups: Agnathans are ancestral to tebrate species. There are almost 28,000 known extant
Chondrichthyes, who again have given rise to Acanthodi- species, of which almost 27,000 are bony sh, with 970
ians, the ancestors of Osteichthyes. With the arrival of sharks, rays, and chimeras and about 108 hagsh and
phylogenetic nomenclature, the shes has been split up lampreys.[9] A third of these species fall within the nine
into a more detailed scheme, with the following major largest families; from largest to smallest, these families are Cyprinidae, Gobiidae, Cichlidae, Characidae,
groups:
Loricariidae, Balitoridae, Serranidae, Labridae, and
Scorpaenidae. About 64 families are monotypic, contain Class Myxini (hagsh)
ing only one species. The nal total of extant species may
Class Pteraspidomorphi (early jawless sh)
grow to exceed 32,500.[10]

Diversity

Examples of the major classes of sh

Agnatha
(Pacic hagsh)

Fish come in many shapes and sizes. This is a sea dragon, a close
relative of the seahorse. Their leaf-like appendages enable them
to blend in with oating seaweed.

lampreys, sharks and rays, ray-nned sh, coelacanths,

and lungsh.[12][13] Indeed, lungsh and coelacanths are
closer relatives of tetrapods (such as mammals, birds,
amphibians, etc.) than of other sh such as ray-nned sh
or sharks, so the last common ancestor of all sh is also
an ancestor to tetrapods. As paraphyletic groups are no
longer recognised in modern systematic biology, the use
Chondrichthyes
of the term sh as a biological group must be avoided.

(Horn shark)

Actinopterygii
(Brown trout)

Many types of aquatic animals commonly referred to as

sh are not sh in the sense given above; examples include shellsh, cuttlesh, starsh, craysh and jellysh.
In earlier times, even biologists did not make a distinction
sixteenth century natural historians classied also seals,
whales, amphibians, crocodiles, even hippopotamuses, as
well as a host of aquatic invertebrates, as sh.[14] However, according to the denition above, all mammals, including cetaceans like whales and dolphins, are not sh.
In some contexts, especially in aquaculture, the true sh
are referred to as nsh (or n sh) to distinguish them
from these other animals.
A typical sh 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 ns, usually one or two
(rarely three) dorsal ns, an anal n, and a tail n, has
jaws, has skin that is usually covered with scales, and lays
eggs.

(Coelacanth)
Main article: Diversity of sh
The term sh 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 ns.[11] Unlike groupings such as birds or mammals, sh are not a single clade
but a paraphyletic collection of taxa, including hagshes,

Each criterion has exceptions. Tuna, swordsh, and

some species of sharks show some warm-blooded adaptaSarcopterygii
tionsthey
can heat their bodies signicantly above ambient water temperature.[12] Streamlining and swimming
performance varies from sh such as tuna, salmon, and
jacks that can cover 1020 body-lengths per second to
species such as eels and rays that swim no more than 0.5
body-lengths per second.[15] Many groups of freshwater
sh extract oxygen from the air as well as from the water using a variety of dierent structures. Lungsh have
paired lungs similar to those of tetrapods, gouramis have
a structure called the labyrinth organ that performs a simi-

3 ANATOMY

lar function, while many catsh, such as corydoras extract

oxygen via the intestine or stomach.[16] Body shape and
the arrangement of the ns is highly variable, covering
such seemingly un-shlike forms as seahorses, puersh,
anglersh, and gulpers. Similarly, the surface of the skin
may be naked (as in moray eels), or covered with scales
of a variety of dierent types usually dened as placoid
(typical of sharks and rays), cosmoid (fossil lungsh and
coelacanths), ganoid (various fossil sh but also living
gars and bichirs), cycloid, and ctenoid (these last two are
found on most bony sh).[17] There are even sh that live
mostly on land. Mudskippers feed and interact with one
another on mudats and go underwater to hide in their
burrows.[18] The catsh Phreatobius cisternarum lives in
underground, phreatic habitats, and a relative lives in waterlogged leaf litter.[19][20]

3.1.1 Gills
Most sh exchange gases using gills on either side of
the pharynx. Gills consist of threadlike structures called
laments. Each lament contains a capillary network that
provides a large surface area for exchanging oxygen and
carbon dioxide. Fish exchange gases by pulling oxygenrich water through their mouths and pumping it over their
gills. In some sh, capillary blood ows 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 sh, like sharks
and lampreys, possess multiple gill openings. However,
bony sh have a single gill opening on each side. This
opening is hidden beneath a protective bony cover called
an operculum.

Fish range in size from the huge 16-metre (52 ft) whale Juvenile bichirs have external gills, a very primitive feashark to the tiny 8-millimetre (0.3 in) stout infantsh.
ture that they share with larval amphibians.
Fish species diversity is roughly divided equally between marine (oceanic) and freshwater ecosystems. Coral
reefs in the Indo-Pacic constitute the center of diversity for marine shes, whereas continental freshwater
shes are most diverse in large river basins of tropical
rainforests, especially the Amazon, Congo, and Mekong
basins. More than 5,600 sh species inhabit Neotropical
freshwaters alone, such that Neotropical shes represent
about 10% of all vertebrate species on the Earth. Exceptionally rich sites in the Amazon basin, such as Canto
State Park, can contain more freshwater sh species than
occur in all of Europe.[21]

Anatomy

Main article: Fish anatomy

The anatomy of Lampanyctodes hectoris

(1) operculum (gill cover), (2) lateral line, (3) dorsal n,
(4) fat n, (5) caudal peduncle, (6) caudal n, (7) anal
n, (8) photophores, (9) pelvic ns (paired), (10) pectoral
ns (paired)

3.1

Respiration

3.1.2 Air breathing

Fish from multiple groups can live out of the water for extended periods. Amphibious sh 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 sh 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. Catsh of the families Loricariidae,
Callichthyidae, and Scoloplacidae absorb air through
their digestive tracts.[22] Lungsh, with the exception of
the Australian lungsh, 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 bown have a vascularized swim bladder that
functions in the same way. Loaches, trahiras, and many
catsh breathe by passing air through the gut. Mudskippers breathe by absorbing oxygen across the skin (similar
to frogs). A number of sh have evolved so-called accessory breathing organs that extract oxygen from the
air. Labyrinth sh (such as gouramis and bettas) have a
labyrinth organ above the gills that performs this function.
A few other sh have structures resembling labyrinth
organs in form and function, most notably snakeheads,
pikeheads, and the Clariidae catsh family.
Breathing air is primarily of use to sh that inhabit shallow, seasonally variable waters where the waters oxygen
concentration may seasonally decline. Fish dependent
solely on dissolved oxygen, such as perch and cichlids,
quickly suocate, 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 sh are
able to survive in damp burrows for weeks without water,
entering a state of aestivation (summertime hibernation)
until water returns.
Air breathing sh can be divided into obligate air

3.4

Excretion

5
The intestine completes the process of digestion and nutrient absorption.

3.4 Excretion
As with many aquatic animals, most sh release their nitrogenous wastes as ammonia. Some of the wastes diuse
through the gills. Blood wastes are ltered by the kidneys.
Saltwater sh tend to lose water because of osmosis.
Their kidneys return water to the body. The reverse happens in freshwater sh: they tend to gain water osmotically. Their kidneys produce dilute urine for excretion.
Some sh have specially adapted kidneys that vary in
function, allowing them to move from freshwater to saltwater.
Tuna gills inside the head. The sh head is oriented snoutdownwards, with the view looking towards the mouth.

3.5 Scales
breathers and facultative air breathers. Obligate air
breathers, such as the African lungsh, must breathe air
periodically or they suocate. Facultative air breathers,
such as the catsh Hypostomus plecostomus, only breathe
air if they need to and will otherwise rely on their
gills for oxygen. Most air breathing sh are facultative
air breathers that avoid the energetic cost of rising to
the surface and the tness cost of exposure to surface
predators.[22]

Main article: Fish scale

3.2

Fish typically have quite small brains relative to body

size compared with other vertebrates, typically onefteenth the brain mass of a similarly sized bird or
mammal.[24] However, some sh have relatively large
brains, most notably mormyrids and sharks, which have
brains about as massive relative to body weight as birds
and marsupials.[25]

Circulation

Fish have a closed-loop circulatory system. The heart

pumps the blood in a single loop throughout the body.
In most sh, the heart consists of four parts, including
two chambers and an entrance and exit.[23] The rst part
is the sinus venosus, a thin-walled sac that collects blood
from the shs veins before allowing it to ow 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 thickwalled, muscular chamber and it pumps the blood, rst 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 ows to the gills for oxygenation.

3.3

Digestion

The scales of sh originate from the mesoderm (skin);

they may be similar in structure to teeth.

3.6 Sensory and nervous system

3.6.1 Central nervous system

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.[24] The olfactory lobes are very large in
sh that hunt primarily by smell, such as hagsh, sharks,
and catsh. Behind the olfactory lobes is the two-lobed
telencephalon, the structural equivalent to the cerebrum
in higher vertebrates. In sh the telencephalon is concerned mostly with olfaction.[24] 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.[24] The pineal body lies just above the
diencephalon. This structure detects light, maintains
circadian rhythms, and controls color changes.[24]

Jaws allow sh 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 sh, processed
in nger-shaped pouches called pyloric caeca, which secrete digestive enzymes and absorb nutrients. Organs The midbrain or mesencephalon contains the two optic
such as the liver and pancreas add enzymes and various lobes. These are very large in species that hunt by sight,
chemicals as the food moves through the digestive tract. such as rainbow trout and cichlids.[24]

3 ANATOMY
the order of millivolt.[27] Other sh, like the South American electric shes 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.[28]
Vision Main article: Vision in shes
Vision is an important sensory system for most species
of sh. 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 sh can see ultraviolet and some
can see polarized light. Amongst jawless sh, the lamprey
has well-developed eyes, while the hagsh has only primitive eyespots.[29] Fish vision shows adaptation to their visual environment, for example deep sea shes have eyes
suited to the dark environment.
Hearing See also: Sensory systems in sh Hearing
Hearing is an important sensory system for most species
of sh. Fish sense sound using their lateral lines and their
ears.

Dorsal view of the brain of the rainbow trout

The hindbrain or metencephalon is particularly involved

in swimming and balance.[24] The cerebellum is a singlelobed structure that is typically the biggest part of the
brain.[24] Hagsh and lampreys have relatively small cerebellae, while the mormyrid cerebellum is massive and apparently involved in their electrical sense.[24]

3.6.3 Capacity for pain

Further information: Pain in sh

Experiments done by William Tavolga provide evidence

that sh have pain and fear responses. For instance, in
Tavolgas experiments, toadsh grunted when electrically
shocked and over time they came to grunt at the mere
The brain stem or myelencephalon is the brains sight of an electrode.[30]
posterior.[24] As well as controlling some muscles and In 2003, Scottish scientists at the University of Edinburgh
body organs, in bony sh at least, the brain stem governs and the Roslin Institute concluded that rainbow trout exrespiration and osmoregulation.[24]
hibit behaviors often associated with pain in other animals. Bee venom and acetic acid injected into the lips
resulted in sh rocking their bodies and rubbing their lips
3.6.2 Sense organs
along the sides and oors of their tanks, which the reto relieve pain, simiMost sh possess highly developed sense organs. Nearly searchers concluded were attempts
[31][32]
lar
to
what
mammals
would
do.
Neurons red in a
all daylight sh have color vision that is at least as good
[32]
pattern
resembling
human
neuronal
patterns.
as a humans (see vision in shes). Many sh also have
chemoreceptors that are responsible for extraordinary Professor James D. Rose of the University of Wyoming
senses of taste and smell. Although they have ears, many claimed the study was awed since it did not provide
sh may not hear very well. Most sh have sensitive re- proof that sh possess conscious awareness, particularly
ceptors that form the lateral line system, which detects a kind of awareness that is meaningfully like ours.[33]
gentle currents and vibrations, and senses the motion of Rose argues that since sh brains are so dierent from hunearby sh and prey.[26] Some sh, such as catsh and man brains, sh are probably not conscious in the manner
sharks, have organs that detect weak electric currents on humans are, so that reactions similar to human reactions

3.8

Homeothermy

to pain instead have other causes. Rose had published a

study a year earlier arguing that sh cannot feel pain because their brains lack a neocortex.[34] However, animal
behaviorist Temple Grandin argues that sh could still
have consciousness without a neocortex because dierent species can use dierent brain structures and systems
to handle the same functions.[32]
Animal welfare advocates raise concerns about the possible suering of sh caused by angling. Some countries,
such as Germany have banned specic types of shing,
and the British RSPCA now formally prosecutes individuals who are cruel to sh.[35]

3.7

Muscular system

Main article: Fish locomotion

Most sh move by alternately contracting paired sets of

3.8 Homeothermy
Although most sh are exclusively ectothermic, there are
exceptions.
Certain species of sh maintain elevated body temperatures. Endothermic teleosts (bony sh) are all in the
suborder Scombroidei and include the billshes, tunas,
and one species of primitive mackerel (Gasterochisma
melampus). All sharks in the family Lamnidae shortn mako, long n mako, white, porbeagle, and salmon
shark are endothermic, and evidence suggests the trait
exists in family Alopiidae (thresher sharks). The degree
of endothermy varies from the billsh, which warm only
their eyes and brain, to bluen tuna and porbeagle sharks
who maintain body temperatures elevated in excess of
20 C above ambient water temperatures.[36] See also
gigantothermy. 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.

3.9 Reproductive system

Further information: Fish reproduction and Spawn (biology)
Fish reproductive organs include testicles and ovaries. In
Swim bladder of a rudd (Scardinius erythrophthalmus)

muscles on either side of the backbone. These contractions form S-shaped curves that move down the body. As
each curve reaches the back n, backward force is applied
to the water, and in conjunction with the ns, moves the
sh forward. The shs ns function like an airplanes
aps. Fins also increase the tails surface area, increasing speed. The streamlined body of the sh decreases the
amount of friction from the water. Since body tissue is
denser than water, sh must compensate for the dierence or they will sink. Many bony sh have an internal
organ called a swim bladder that adjusts their buoyancy
through manipulation of gases.
Organs: 1. Liver, 2. Gas bladder, 3. Roe, 4. Pyloric caeca, 5.
Stomach, 6. Intestine

most species, gonads are paired organs of similar size,

which can be partially or totally fused.[37] There may also
be a range of secondary organs that increase reproductive
tness.
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 sh they are conned to the distal portion of these structures. Fish can present cystic or semicystic spermatogenesis in relation to the release phase of
germ cells in cysts to the seminiferous tubules lumen.[37]
A great white shark o Isla Guadalupe

Fish ovaries may be of three types: gymnovarian, secondary gymnovarian or cystovarian. In the rst type,

8
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.[38] Gymnovaries are
the primitive condition found in lungsh, sturgeon, and
bown. Cystovaries characterize most teleosts, where
the ovary lumen has continuity with the oviduct.[37] Secondary gymnovaries are found in salmonids and a few
other teleosts.
Oogonia development in teleosts sh 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.[37]
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.[37]

3 ANATOMY

Egg of catshark (mermaids purse)

Egg of bullhead shark

Egg of chimaera
Some sh, like the California sheephead, are
hermaphrodites, having both testes and ovaries either at dierent phases in their life cycle or, as in
The newly hatched young of oviparous sh are called
hamlets, have them simultaneously.
larvae. They are usually poorly formed, carry a large yolk
Over 97% of all known sh are oviparous,[39] that is, sac (for nourishment), and are very dierent in appearthe eggs develop outside the mothers body. Examples ance from juvenile and adult specimens. The larval peof oviparous sh include salmon, goldsh, cichlids, tuna, riod in oviparous sh is relatively short (usually only sevand eels. In the majority of these species, fertilisation eral weeks), and larvae rapidly grow and change appeartakes place outside the mothers body, with the male and ance and structure (a process termed metamorphosis) to
female sh shedding their gametes into the surrounding become juveniles. During this transition larvae must
water. However, a few oviparous sh practice internal switch from their yolk sac to feeding on zooplankton prey,
fertilization, with the male using some sort of intromittent a process which depends on typically inadequate zooorgan to deliver sperm into the genital opening of the fe- plankton density, starving many larvae.
male, most notably the oviparous sharks, such as the horn
In ovoviviparous sh the eggs develop inside the mothers
shark, and oviparous rays, such as skates. In these cases,
body after internal fertilization but receive little or no
the male is equipped with a pair of modied pelvic ns
nourishment directly from the mother, depending instead
known as claspers.
on the yolk. Each embryo develops in its own egg. FamilMarine sh can produce high numbers of eggs which are iar examples of ovoviviparous sh include guppies, angel
often released into the open water column. The eggs have sharks, and coelacanths.
an average diameter of 1 millimetre (0.039 in).
Some species of sh are viviparous. In such species the
mother retains the eggs and nourishes the embryos. Typically, viviparous sh have a structure analogous to the
placenta seen in mammals connecting the mothers blood
supply with that of the embryo. Examples of viviparous
sh include the surf-perches, splitns, and lemon shark.
Some viviparous sh exhibit oophagy, in which the developing embryos eat other eggs produced by the mother.
This has been observed primarily among sharks, such as
the shortn mako and porbeagle, but is known for a few

Egg of lamprey
bony sh as well, such as the halfbeak Nomorhamphus
ebrardtii.[40] Intrauterine cannibalism is an even more un-

9
usual 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.[40]

thymus and a well-developed spleen (their most important

immune organ) where various lymphocytes, plasma cells
and macrophages develop and are stored. Chondrostean
sh (sturgeons, paddlesh, and bichirs) possess a major
site for the production of granulocytes within a mass that
Aquarists commonly refer to ovoviviparous and is associated with the meninges (membranes surrounding
the central nervous system.) Their heart is frequently covviviparous sh as livebearers.
ered with tissue that contains lymphocytes, reticular cells
and a small number of macrophages. The chondrostean
kidney is an important hemopoietic organ; where erythro4 Diseases
cytes, granulocytes, lymphocytes and macrophages develop.
Main article: Fish diseases and parasites
Like chondrostean sh, the major immune tissues of
bony sh (or teleostei) include the kidney (especially the
Like other animals, sh suer from diseases and par- anterior kidney), which houses many dierent immune
asites. To prevent disease they have a variety of de- cells.[47] In addition, teleost sh possess a thymus, spleen
fenses. Non-specic defenses include the skin and scales, and scattered immune areas within mucosal tissues (e.g.
as well as the mucus layer secreted by the epidermis in the skin, gills, gut and gonads). Much like the mamthat traps and inhibits the growth of microorganisms. malian immune system, teleost erythrocytes, neutrophils
If pathogens breach these defenses, sh can develop an and granulocytes are believed to reside in the spleen
inammatory response that increases blood ow to the whereas lymphocytes are the major cell type found in the
infected region and delivers white blood cells that at- thymus.[48][49] In 2006, a lymphatic system similar to that
tempt to destroy pathogens. Specic defenses respond in mammals was described in one species of teleost sh,
to particular pathogens recognised by the shs body, i.e., the zebrash. Although not conrmed as yet, this system
an immune response.[41] In recent years, vaccines have presumably will be where naive (unstimulated) T cells acbecome widely used in aquaculture and also with orna- cumulate while waiting to encounter an antigen.[50]
mental sh, for example furunculosis vaccines in farmed
B and T lymphocytes bearing immunoglobulins and T cell
salmon and koi herpes virus in koi.[42][43]
receptors, respectively, are found in all jawed shes. InSome species use cleaner sh to remove external para- deed, the adaptive immune system as a whole evolved in
sites. The best known of these are the Bluestreak cleaner an ancestor of all jawed vertebrate.[51]
wrasses of the genus Labroides found on coral reefs in the
Indian and Pacic oceans. These small sh maintain socalled cleaning stations where other sh congregate and 5 Conservation
perform specic movements to attract the attention of the
cleaners.[44] Cleaning behaviors have been observed in a
The 2006 IUCN Red List names 1,173 sh species
number of sh groups, including an interesting case be[52]
Included are
tween two cichlids of the same genus, Etroplus maculatus, that are threatened with extinction.
[53]
species such as Atlantic cod, Devils Hole pupsh,[54]
[45]
the cleaner, and the much larger Etroplus suratensis.
coelacanths,[55] and great white sharks.[56] Because sh
live underwater they are more dicult to study than terrestrial animals and plants, and information about sh
4.1 Immune system
populations is often lacking. However, freshwater sh
Immune organs vary by type of sh.[46] In the jawless seem particularly threatened because they often live in
sh (lampreys and hagsh), true lymphoid organs are relatively small water bodies. For example, the Devils
absent. These sh rely on regions of lymphoid tissue Hole pupsh occupies only a single 3 by 6 metres (10 by
[57]
within other organs to produce immune cells. For exam- 20 ft) pool.
ple, erythrocytes, macrophages and plasma cells are produced in the anterior kidney (or pronephros) and some
areas of the gut (where granulocytes mature.) They re- 5.1 Overshing
semble primitive bone marrow in hagsh. Cartilaginous
sh (sharks and rays) have a more advanced immune sys- Main article: Overshing
tem. They have three specialized organs that are unique
to Chondrichthyes; the epigonal organs (lymphoid tis- Overshing is a major threat to edible sh such as cod
sue similar to mammalian bone) that surround the go- and tuna.[58][59] Overshing eventually causes population
nads, the Leydigs organ within the walls of their esoph- (known as stock) collapse because the survivors cannot
agus, and a spiral valve in their intestine. These or- produce enough young to replace those removed. Such
gans house typical immune cells (granulocytes, lympho- commercial extinction does not mean that the species is
cytes and plasma cells). They also possess an identiable extinct, merely that it can no longer sustain a shery.

10

6 IMPORTANCE TO HUMANS

6.1 Economic importance

Main articles: Fishing industry, Aquaculture, and Fish
farming
Throughout history, humans have utilized sh as a food

A Whale shark, the worlds largest sh, is classied as

Vulnerable.

One well-studied example of shery collapse is the

Pacic sardine Sadinops sagax caerulues shery o 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 shery was no
longer economically viable.[60]
The main tension between sheries science and the shing
industry is that the two groups have dierent views on the
resiliency of sheries to intensive shing. In places such
as Scotland, Newfoundland, and Alaska the shing industry is a major employer, so governments are predisposed
to support it.[61][62] On the other hand, scientists and conservationists push for stringent protection, warning that
many stocks could be wiped out within fty years.[63][64]

5.2

Habitat destruction

See also: Environmental impact of shing

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.[65] An example of a sh
that has become endangered because of habitat change is
the pallid sturgeon, a North American freshwater sh that
lives in rivers damaged by human activity.[66]

5.3

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 lakes 500 endemic
cichlid species. Some of them survive now in captive
breeding programmes, but others are probably extinct.[67]
Carp, snakeheads,[68] tilapia, European perch, brown
trout, rainbow trout, and sea lampreys are other examples
of sh that have caused problems by being introduced into
alien environments.

Importance to humans

These sh-farming ponds were created as a cooperative project

in a rural village.

source. Historically and today, most sh protein has come

by means of catching wild sh. However, aquaculture, or
sh farming, which has been practiced since about 3,500
BCE. in China,[69] is becoming increasingly important in
many nations. Overall, about one-sixth of the worlds protein is estimated to be provided by sh.[70] That proportion is considerably elevated in some developing nations
and regions heavily dependent on the sea. In a similar
manner, sh have been tied to trade.
Catching sh for the purpose of food or sport is known
as shing, while the organized eort by humans to catch
sh is called a shery. Fisheries are a huge global business and provide income for millions of people.[70] The
annual yield from all sheries worldwide is about 154 million tons,[71] with popular species including herring, cod,
anchovy, tuna, ounder, and salmon. However, the term
shery is broadly applied, and includes more organisms
than just sh, such as mollusks and crustaceans, which
are often called sh when used as food.

6.2 Recreation
Main articles: Fishkeeping, Recreational shing, and
Angling
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 sh in ponds, and displayed in
aquariums in homes, oces, or public settings.
Recreational shing is shing for pleasure or competition;
it can be contrasted with commercial shing, which is
shing for prot. The most common form of recreational
shing is done with a rod, reel, line, hooks and any one
of a wide range of baits. Angling is a method of shing,
specically the practice of catching sh by means of an

11
angle (hook). Anglers must select the right hook, cast
accurately, and retrieve at the right speed while considering water and weather conditions, species, sh response,
time of the day, and other factors.

6.3

Culture

See also: Mermaid and Merman

Fish feature prominently in art and literature, in movies
The ichthus is a Christian symbol of a sh signifying that the
person who uses it is a Christian.[73]

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 sh are Ika-Roa of the Polynesians,
Dagon of various ancient Semitic peoples, the shark-gods
of Hawaii and Matsya of the Hindus.
The astrological symbol Pisces is based on a constellation
of the same name, but there is also a second sh constellation in the night sky, Piscis Austrinus.[75]

7 Terminology
7.1 Shoal or school
Main article: Shoaling and schooling
A random assemblage of sh merely using some lo-

Avatar of Vishnu as a Matsya

such as Finding Nemo and books such as The Old Man

and the Sea. Large sh, particularly sharks, have frequently been the subject of horror movies and thrillers,
most notably the novel Jaws, which spawned a series of
lms of the same name that in turn inspired similar lms
or parodies such as Shark Tale and Snakehead Terror.
Piranhas are shown in a similar light to sharks in lms
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.[72] In the Book
of Jonah a great sh swallowed Jonah the Prophet. Legends of half-human, half-sh mermaids have featured in
folklore, like the stories of Hans Christian Andersen.
Fish themes have symbolic signicance in many religions. The sh is used often as a symbol by Christians
to represent Jesus, or Christianity in general; the gospels
also refer to shers of men[74] and feeding the multitude. In the dhamma of Buddhism the sh symbolize
happiness as they have complete freedom of movement
in the water. Often drawn in the form of carp which are

These goldband fusiliers are schooling because their swimming

is synchronised

calised resource such as food or nesting sites is known

simply as an aggregation. When sh 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 sh 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 sh are much
more tightly organised, synchronising their swimming so

12

9 NOTES

that all sh move at the same speed and in the same di- 8 See also
rection. Shoaling and schooling behaviour is believed to
provide a variety of advantages.[76]
Main article: Outline of sh
Examples:
Cichlids congregating at lekking sites form an aggregation.

For a topical guide to sharks, see Outline of

sharks

Many minnows and characins form shoals.

Angling (sport shing)

Anchovies, herrings and silversides are classic examples of schooling sh.

Aquaculture

While the words school and shoal have dierent

meanings within biology, they are often treated as
synonyms by non-specialists, with speakers of British English using shoal to describe any grouping of sh, and
speakers of American English often using school just
as loosely.

7.2

Fish or shes

Though often used interchangeably, in biology these

words have dierent 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 dierent species or species groups.[77][78][79] Thus a pond that
contained a single species might be said to contain 120
sh. But if the pond contained a total of 120 sh from
three dierent species, it would be said to contain three
shes. The distinction is similar to that between people
and peoples.

7.3

Finsh

In biology, the term sh is most strictly used to

describe any animal with a backbone that has gills
throughout life and has limbs, if any, in the shape of
ns.[80] Many types of aquatic animals with common
names ending in sh are not sh in this sense; examples include shellsh, cuttlesh, starsh, craysh
and jellysh. In earlier times, even biologists did
not make a distinctionsixteenth century natural
historians classied also seals, whales, amphibians,
crocodiles, even hippopotamuses, as well as a host
of aquatic invertebrates, as sh.[14]
In sheries, the term sh is used as a collective term,
and includes mollusks, crustaceans and any aquatic
animal which is harvested.[81]
The strict biological denition of a sh, above, is
sometimes called a true sh. True sh are also referred to as nsh or n sh to distinguish them
from other aquatic life harvested in sheries or aquaculture.

Aquarium
Catch and release
Deep sea sh
Fish acute toxicity syndrome
Fish anatomy
Fish as food
Fish development
Fishing (shing for food)
Fish intelligence
Fishkeeping
Forage sh
Ichthyology
List of sh common names
List of sh families
Marine biology
Marine vertebrates
Mercury in sh
Otolith (Bone used for determining the age of a sh)
Pregnancy (sh)
Seafood
Walking sh

9 Notes
[1] Goldman, K.J. (1997). Regulation of body temperature in the white shark, Carcharodon carcharias". Journal of Comparative Physiology. B Biochemical Systemic and Environmental Physiology. 167 (6): 423
429. doi:10.1007/s003600050092. Retrieved 12 October 2011.

13

[2] Carey, F.G.; Lawson, K.D. (February 1973). Temperature regulation in free-swimming bluen tuna. Comparative Biochemistry and Physiology A. 44 (2): 375392.
doi:10.1016/0300-9629(73)90490-8.
[3] FishBase. FishBase. April 2015. Retrieved 29 August
2015.
[4] G. Lecointre & H. Le Guyader, 2007, The Tree of Life: A
Phylogenetic Classication, Harvard University Press Reference Library
[5] Romer, A.S. & T.S. Parsons. 1977. The Vertebrate Body.
5th ed. Saunders, Philadelphia. (6th ed. 1985)
[6] Benton, M. J. (1998) The quality of the fossil record of
vertebrates. Pp. 269303, in Donovan, S. K. and Paul,
C. R. C. (eds), The adequacy of the fossil record, Fig. 2.
Wiley, New York, 312 pp.
[7] Shigehiro Kuraku, Daisuke Hoshiyama, Kazutaka Katoh,
Hiroshi Suga, Takashi Miyata (1999) Monophyly of Lampreys and Hagshes Supported by Nuclear DNACoded
Genes J Mol Evol (1999) 49:729735

[23] Setaro, John F. (1999). Circulatory System. Microsoft

Encarta 99.
[24] Helfman, Collette & Facey 1997, pp. 4849
[25] Helfman, Collette & Facey 1997, p. 191
[26] Orr, James (1999). Fish. Microsoft Encarta 99. ISBN
0-8114-2346-8.
[27] Albert, J.S., and W.G.R. Crampton. 2005. Electroreception and electrogenesis. pp. 431472 in The Physiology
of Fishes, 3rd Edition. D.H. Evans and J.B. Claiborne
(eds.). CRC Press.
[28] Sciences, Journal of Undergraduate Life. Appropriate
maze methodology to study learning in sh (PDF). Retrieved 28 May 2009.
[29] Campbell, Neil A.; Reece, Jane B. (2005). Biology (Seventh ed.). San Francisco, California: Benjamin Cummings.
[30] Dunayer, Joan, Fish: Sensitivity Beyond the Captors
Grasp, The Animals Agenda, July/August 1991, pp. 12
18

[8] J. Mallatt, J. Sullivan (1998) 28S and 18S rDNA sequences

support the monophyly of lampreys and hagshes Molecular Biology and Evolution V 15, Issue 12, pp 17061718

[31] Kirby, Alex (30 April 2003). Fish do feel pain, scientists
say. BBC News. Retrieved 4 January 2010.

[9] Nelson 2006, pp. 45

[32] Grandin, Temple; Johnson, Catherine (2005). Animals in

Translation. New York, New York: Scribner. pp. 183
184. ISBN 0-7432-4769-8.

[10] Nelson 2006, p. 3

[11] Nelson 2006, p. 2
[12] Helfman, Collette & Facey 1997, p. 3

[33] Rose, J.D. 2003. A Critique of the paper: Do sh have

nociceptors: Evidence for the evolution of a vertebrate
sensory system"" (PDF). Retrieved 21 May 2011.

[13] Tree of life web project Chordates.

[34] Rose, James D. (2002). Do Fish Feel Pain?". Retrieved

27 September 2007.

[14] Cleveland P. Hickman, Jr.; Larry S. Roberts; Allan

L. Larson (2001). Integrated Principles of Zoology.
McGraw-Hill Publishing Co. ISBN 0-07-290961-7.

[35] The Times & The Sunday Times. Retrieved 30 December 2016.

[15] Helfman, Collette & Facey 1997, p. 103

[16] Helfman, Collette & Facey 1997, pp. 5357
[17] Helfman, Collette & Facey 1997, pp. 3336
[18] Froese, Rainer and Pauly, Daniel, eds.
(2006).
"Periophthalmus barbarus" in FishBase. November 2006
version.
[19] Froese, Rainer and Pauly, Daniel, eds.
(2006).
"Phreatobius cisternarum" in FishBase. November 2006
version.

[36] Block, BA; Finnerty, JR (1993). Endothermy in shes:

a phylogenetic analysis of constraints, predispositions,
and selection pressures (PDF). Environmental Biology of
Fishes. 40 (3): 283302. doi:10.1007/BF00002518.
[37] Guimaraes-Cruz, Rodrigo J., Rodrigo J.; Santos, Jos E.
dos; Santos, Gilmar B. (JulySeptember 2005). Gonadal structure and gametogenesis of Loricaria lentiginosa Isbrcker (Pisces, Teleostei, Siluriformes)". Rev.
Bras. Zool. 22 (3): 556564. doi:10.1590/S010181752005000300005. ISSN 0101-8175.

[20] Planet Catsh. Cat-eLog: Heptapteridae: Phreatobius:

Phreatobius sp. (1)". Planet Catsh. Retrieved 26
November 2006.

[38] Brito, M.F.G.; Bazzoli, N. (2003). Reproduction

of the surubim catsh (Pisces, Pimelodidae) in the
So Francisco River, Pirapora Region, Minas Gerais,
Brazil. Arquivo Brasileiro de Medicina Veterinria e
Zootecnia. 55 (5): 624633. doi:10.1590/S010209352003000500018. ISSN 0102-0935.

[21] Estudo das Espcies cticas do Parque Estadual do Canto,

sh species survey of Canto (in Portuguese)

[39] Peter Scott: Livebearing Fishes, p. 13. Tetra Press 1997.

ISBN 1-56465-193-2

[22] Modications of the Digestive Tract for Holding Air in

Loricariid and Scoloplacid Catshes (PDF). Copeia (3):
663675. 1998. doi:10.2307/1447796. Retrieved 25
June 2009.

[40] Meisner, A & Burns, J: Viviparity in the Halfbeak Genera Dermogenys and Nomorhamphus (Teleostei: Hemiramphidae)" Journal of Morphology 234, pp. 295317,
1997

14

9 NOTES

[41] Helfman, Collette & Facey 1997, pp. 9596

[60] Helfman, Collette & Facey 1997, p. 462

[42] R. C. Cipriano (2001), Furunculosis And Other Diseases

Caused By Aeromonas salmonicida. Fish Disease Leaet
66. U.S. Department of the Interior.

[61] UK 'must shield shing industry'". BBC News. 3 November 2006. Retrieved 18 January 2006.

[43] Hartman, K H; et al. (2004). Koi Herpes Virus

(KHV) Disease: Fact Sheet VM-149 (PDF). University
of Florida Institute of Food and Agricultural Sciences.
[44] Helfman, Collette & Facey 1997, p. 380
[45] Wyman, Richard L.; Ward, Jack A. (1972). A
Cleaning Symbiosis between the Cichlid Fishes Etroplus maculatus and Etroplus suratensis. I. Description
and Possible Evolution. Copeia. 1972 (4): 834838.
doi:10.2307/1442742.
[46] A.G. Zapata, A. Chiba and A. Vara. Cells and tissues of
the immune system of sh. In: The Fish Immune System:
Organism, Pathogen and Environment. Fish Immunology
Series. (eds. G. Iwama and T.Nakanishi,), New York,
Academic Press, 1996, pp. 155.
[47] D.P. Anderson. Fish Immunology. (S.F. Snieszko and
H.R. Axelrod, eds), Hong Kong: TFH Publications, Inc.
Ltd., 1977.
[48] Chilmonczyk, S. (1992). "The thymus in sh: development and possible function in the immune response". 2.
Annual Review of Fish Diseases: 181200.

[62] EU sh quota deal hammered out. BBC News. 21 December 2006. Retrieved 18 January 2006.
[63] Ocean study predicts the collapse of all seafood sheries
by 2050. Retrieved 13 January 2006.
[64] Atlantic bluen tuna could soon be commercially extinct. Archived from the original on 30 April 2007. Retrieved 18 January 2006.
[65] Helfman, Collette & Facey 1997, p. 463
[66] Threatened and Endangered Species: Pallid Sturgeon
Scaphirhynchus Fact Sheet. Archived from the original
on 26 November 2005. Retrieved 18 March 2016.
[67] Spinney, Laura (4 August 2005). The little sh ght
back. The Guardian. London. Retrieved 18 January
2006.
[68] Stop That Fish!". The Washington Post. 3 July 2002.
Retrieved 26 August 2007.
[69] Spalding, Mark (July 11, 2013). Sustainable Ancient
Aquaculture. National Geographic. Retrieved 13 August 2015.

[49] Hansen, J.D.; Zapata, A.G. (1998). "Lymphocyte development in sh and amphibians". 166. Immunological Reviews: 199220.

[70] Helfman, Gene S. (2007). Fish Conservation: A Guide

to Understanding and Restoring Global Aquatic Biodiversity and Fishery Resources. Island Press. p. 11. ISBN
1597267600.

[50] Kucher; et al. (2006). "Development of the zebrash lymphatic system requires VegFc signalling". 16. Current Biology: 12441248.

[71] World Review of Fisheries and Aquaculture (PDF).

fao.org. Food and Agriculture Organization of the United
Nations. Retrieved 13 August 2015.

[51] Flajnik, M. F., and M. Kasahara. Origin and evolution of

the adaptive immune system: genetic events and selective
pressures. Nature Reviews Genetics 11.1, 47-59 (2009).

[72] Zollinger, Sue Anne (3 July 2009). PiranhaFerocious

Fighter or Scavenging Softie?". A Moment of Science. Indiana Public Media. Retrieved 1 November 2015.

[52] Table 1: Numbers of threatened species by major groups

of organisms (19962004)". iucnredlist.org. Archived
from the original on 30 June 2006. Retrieved 18 January
2006.

[73] Coman, Elesha (August 8, 2008). What is the origin

of the Christian sh symbol?". Christianity Today. Retrieved 13 August 2015.

[53] Gadus morhua (Atlantic Cod)". Iucnredlist.org. Retrieved 21 May 2011.

[74] Matthew 4:19

Iuc-

[75] Piscis Austrinus. allthesky.com. The Deep Photographic Guide to the Constellations. Retrieved 1 November 2015.

[55] Latimeria chalumnae (Coelacanth, Gombessa)". Iucnredlist.org. Retrieved 21 May 2011.

[76] Helfman, G.; Collette, B.; Facey, D. (1997). The Diversity

of Fishes. Blackwell Publishing. p. 375. ISBN 0-86542256-7.

[54] Cyprinodon diabolis (Devils Hole Pupsh)".

nredlist.org. Retrieved 21 May 2011.

Iuc-

[77] Pauly, Daniel (13 May 2004). Fish(es)". Darwins

Fishes: An Encyclopedia of Ichthyology, Ecology, and
Evolution. Cambridge University Press. p. 77. ISBN
978-1-139-45181-9.

[58] Call to halt cod 'over-shing'". BBC News. 5 January

2007. Retrieved 18 January 2006.

[78] Nelson, Joseph S and Paetz, Martin Joseph (1992) The

Fishes of Alberta page 400, University of Alberta. ISBN
9780888642363.

[56] Carcharodon carcharias (Great White Shark)".

nredlist.org. Retrieved 21 May 2011.
[57] Helfman, Collette & Facey 1997, pp. 449450

[59] Tuna groups tackle overshing. BBC News. 26 January

2007. Retrieved 18 January 2006.

[79] Helfman, Collette & Facey 1997, p. 5

15

[80] Nelson, Joseph S. (2006). Fishes of the World. John Wiley & Sons, Inc. p. 2. ISBN 0-471-25031-7.

The Native Fish Conservancy Conservation and

study of North American freshwater shes

[81] FAO: Fisheries glossary

United Nation Fisheries and Aquaculture Department: Fish and seafood utilization

10

University of Washington Libraries Digital Collections Digital collection of freshwater and marine
sh images

References

Eschmeyer, William N.; Fong, Jon David (2013).

Catalog of Fishes. California Academy of Sciences.
Helfman, G.; Collette, B.; Facey, D. (1997). The Diversity of Fishes (1st ed.). Wiley-Blackwell. ISBN
978-0-86542-256-8.
Moyle, Peter B.; Cech, Joseph J. (2003). Fishes,
An Introduction to Ichthyology (5th ed.). Benjamin
Cummings. ISBN 978-0-13-100847-2.
Nelson, Joseph S. (2006). Fishes of the World
(PDF) (4th ed.). John Wiley & Sons. ISBN
9780471756446. Archived from the original (PDF)
on 5 March 2013. Retrieved 30 April 2013.

11

Further reading

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 Enthusiasts
Guide University of California Press.
ISBN
9780520916654 good lay text.
Shubin, Neil (2009) Your inner sh: A journey into
the 3.5 billion year history of the human body Vintage Books. ISBN 9780307277459. UCTV interview

12

External links

ANGFA Illustrated database of freshwater shes

of Australia and New Guinea
Fischinfos.de Illustrated database of the freshwater shes of Germany (German)
FishBase online Comprehensive database with information on over 29,000 sh species
Fisheries and Illinois Aquaculture Center Data
outlet for sheries and aquaculture research center
in the central US
Philippines Fishes Database with thousands of
Philippine Fishes photographed in natural habitat

Davenport, Charles B.; Ingersoll, Ernest (1905).

"Fish". New International Encyclopedia.
Live webcam feed of a goldsh tank

16

13

13
13.1

TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

Text and image sources, contributors, and licenses

Text

Fish Source: https://en.wikipedia.org/wiki/Fish?oldid=757842886 Contributors: Lee Daniel Crocker, Vicki Rosenzweig, Mav, The Anome,
Clasqm, Alex.tan, Josh Grosse, Danny, Rgamble, Rmhermen, PierreAbbat, Roadrunner, Graham, Heron, Montrealais, Sfdan, Stevertigo, JohnOwens, TimShell, Lexor, Dan Koehl, Dominus, Shyamal, Menchi, Ixfd64, Looxix~enwiki, Ams80, Ahoerstemeier, Stan Shebs,
Snoyes, Notheruser, Angela, War3rd, Julesd, Glenn, Stefan-S, Kils, Tkinias, Susurrus, Dpol, Evercat, Ritcheyrp, Hashar, Vanished user
5zariu3jisj0j4irj, Fuzheado, IceKarma, DJ Clayworth, Tpbradbury, Grendelkhan, Morwen, Jnc, Mowgli~enwiki, Phoebe, Traroth, Topbanana, JonathanDP81, HarryHenryGebel, Fvw, Wetman, Gakrivas, Jusjih, Johnleemk, Pollinator, PuzzletChung, Nufy8, Robbot, Astronautics~enwiki, Pigsonthewing, Moriori, Kizor, Altenmann, Kowey, Yelyos, Nurg, Arkuat, Lsy098~enwiki, Texture, Lisap, Hadal,
Wereon, Robinh, Rho~enwiki, Diberri, Xanzzibar, Pengo, Randyoo, Dina, Nebajoth, Alan Liefting, David Gerard, Enochlau, Gheather,
Huddy, Giftlite, Smjg, Christopher Parham, MPF, Ian Maxwell, Jyril, Sj, Fudoreaper, Tom harrison, Binadot, Obli, Peruvianllama, Everyking, No Guru, Curps, Michael Devore, Arnejohs, David Johnson, Frencheigh, Jfdwol, Tom-, Macrakis, Solipsist, Bobblewik, Jrdioko,
Stevietheman, Ato, Utcursch, Pgan002, Andycjp, Mike R, CryptoDerk, Gdr, Knutux, Slowking Man, Sonjaaa, LucasVB, Antandrus,
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13.2

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DBZFan30, Too Small a Fish to Fry, Bender the Bot, Lucas.Belo, WikiLE and Anonymous: 998

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File:Antennarius_striatus.jpg Source: https://upload.wikimedia.org/wikipedia/commons/3/3d/Antennarius_striatus.jpg License: Public

domain Contributors: ? Original artist: ?
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18

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TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

File:Latimeria_chalumnae01.jpg Source: https://upload.wikimedia.org/wikipedia/commons/3/3d/Latimeria_chalumnae01.jpg License:

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