Fish
1 Evolution
For other uses, see Fish (disambiguation).
For an explanation of very similar terms, see Craniata.
Main article: evolution of fish
Fish do not represent a monophyletic group, and thereA fish is any member of a paraphyletic group of organisms that consist of all gill-bearing aquatic craniate 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.
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 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 anglerfish). With 33,100 described species,
fish exhibit greater species diversity than any other group
of vertebrates.[3]
Outdated evolutionary view of continual gradation
Fish are an important resource for humans worldwide,
especially as food. Commercial and subsistence fishers
hunt fish in wild fisheries (see fishing) or farm them in
ponds or in cages in the ocean (see 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.
Because the term “fish” is defined negatively, and excludes the tetrapods (i.e., the amphibians, reptiles, birds
and mammals) which descend from within the same ancestry, it is paraphyletic, and is not considered a proper
grouping in systematic biology. The traditional term
pisces (also ichthyes) is considered a typological, but not
Leedsichthys (left) the largest known fish.
a phylogenetic classification.
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.
fore the “evolution of fish” is not studied as a single
event.[4]
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. 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 corallike Sea squirt, whose larvae resemble primitive fish in
1
2
1 EVOLUTION
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 †
Dunkleosteus was a gigantic, 10 meter (33 feet) long prehistoric
fish.
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.
1.1
Taxonomy
Fish are a paraphyletic group: that is, any clade containing all fish also contains the tetrapods, which are not fish.
For this reason, groups such as the “Class Pisces” seen in
older reference works are no longer used in formal classifications.
Traditional classification divide fish into three extant
classes, and with extinct forms sometimes classified
within the tree, sometimes as their own classes:[5][6]
• 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 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)
• Class Acanthodii (“spiny sharks”, sometimes classified under bony fishes)†
† – indicates extinct taxon
Some palaeontologists contend that because Conodonta
• Class Osteichthyes (bony fish)
are chordates, they are primitive fish. For a fuller treatment of this taxonomy, see the vertebrate article.
• Subclass Actinopterygii (ray finned fishes)
• Subclass Sarcopterygii (fleshy finned fishes, The position of hagfish in the phylum chordata is not settled. Phylogenetic research in 1998 and 1999 supported
ancestors of tetrapods)
the idea that the hagfish and the lampreys form a natuthat is a sister group of the
The above scheme is the one most commonly encoun- ral group, the Cyclostomata,
[7][8]
Gnathostomata.
tered in non-specialist and general works. Many of the
above groups are paraphyletic, in that they have given The various fish groups account for more than half of verrise to successive groups: Agnathans are ancestral to tebrate species. There are almost 28,000 known extant
3
species, of which almost 27,000 are bony fish, with 970
sharks, rays, and chimeras and about 108 hagfish and
lampreys.[9] 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.[10]
2
Diversity
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.[14] 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.
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 floating seaweed.
Main article: Diversity of fish
The term “fish” most precisely describes any non-
Feeding the koi. The koi are ornamental varieties of domesticated
common carp and are kept in garden ponds.
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.[11] 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.[12][13] Indeed, lungfish and coelacanths are
closer relatives of tetrapods (such as mammals, birds,
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.[12] 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.[12] 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.[15] 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).[16] There are even
fish that live mostly on land. Mudskippers feed and interact with one another on mudflats and go underwater
to hide in their burrows.[17] The catfish Phreatobius cisternarum lives in underground, phreatic habitats, and a
relative lives in waterlogged leaf litter.[18][19]
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
4
3 ANATOMY
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.[20]
3
Anatomy
Main article: Fish anatomy
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)
3.1
3.1.1
3.1.2 Air breathing
Fish from multiple groups can live out of the water for extended periods. Amphibious fish such as the mudskipper
can live and move about on land for up to several days,
or live in stagnant or otherwise oxygen depleted water. Many such fish can breathe air via a variety of
mechanisms. The skin of anguillid eels may absorb
oxygen directly. The buccal cavity of the electric eel
may breathe air. Catfish of the families Loricariidae,
Callichthyidae, and Scoloplacidae absorb air through
their digestive tracts.[21] 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.
Respiration
Gills
Most fish exchange gases using gills on either side of
the pharynx. Gills consist of threadlike structures called
filaments. Each filament contains a capillary network that
provides a large surface area for exchanging oxygen and
carbon dioxide. Fish exchange gases by pulling oxygenrich 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 Tuna gills inside the head. The fish head is oriented snoutopening is hidden beneath a protective bony cover called downwards, with the view looking towards the mouth.
an operculum.
Juvenile bichirs have external gills, a very primitive fea- Air breathing fish can be divided into obligate air
ture that they share with larval amphibians.
breathers and facultative air breathers. Obligate air
3.5
Scales
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.[21]
5
3.5 Scales
Main article: Fish scale
The scales of fish originate from the mesoderm (skin);
they may be similar in structure to teeth.
3.6 Sensory and nervous system
3.2
Circulation
Fish have a closed-loop circulatory system. The heart
pumps the blood in a single loop throughout the body.
In most fish, the heart consists of four parts, including
two chambers and an entrance and exit.[22] 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 thickwalled, 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.
3.3
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.
Dorsal view of the brain of the rainbow trout
3.4
Excretion
3.6.1 Central nervous system
As with many aquatic animals, most fish release their ni- Fish typically have quite small brains relative to body
trogenous wastes as ammonia. Some of the wastes diffuse size compared with other vertebrates, typically onethrough the gills. Blood wastes are filtered by the kidneys. fifteenth the brain mass of a similarly sized bird or
[23]
However, some fish have relatively large
Saltwater fish tend to lose water because of osmosis. mammal.
mormyrids and sharks, which have
brains,
most
notably
Their kidneys return water to the body. The reverse hapbrains
about
as
massive
relative to body weight as birds
pens in freshwater fish: they tend to gain water osmoti[24]
marsupials.
and
cally. Their kidneys produce dilute urine for excretion.
Some fish have specially adapted kidneys that vary in Fish brains are divided into several regions. At the
function, allowing them to move from freshwater to salt- front are the olfactory lobes, a pair of structures that receive and process signals from the nostrils via the two
water.
6
3 ANATOMY
olfactory nerves.[23] 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.[23] Together these structures form the forebrain.
spherical lens. Their retinas generally have both rod
cells and cone cells (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.[28] Fish vision
shows adaptation to their visual environment, for example
Connecting the forebrain to the midbrain is the deep sea fishes have eyes suited to the dark environment.
diencephalon (in the diagram, this structure is below the
optic lobes and consequently not visible). The diencephalon performs functions associated with hormones
and homeostasis.[23] The pineal body lies just above the Hearing See also: Sensory systems in fish § Hearing
diencephalon. This structure detects light, maintains
circadian rhythms, and controls color changes.[23]
Hearing is an important sensory system for most species
The midbrain or mesencephalon contains the two optic of fish. Fish sense sound using their lateral lines and their
lobes. These are very large in species that hunt by sight, ears.
such as rainbow trout and cichlids.[23]
The hindbrain or metencephalon is particularly involved
in swimming and balance.[23] The cerebellum is a single- 3.6.3 Capacity for pain
lobed structure that is typically the biggest part of the
brain.[23] Hagfish and lampreys have relatively small cerebellae, while the mormyrid cerebellum is massive and ap- Further information: Pain in fish
parently involved in their electrical sense.[23]
The brain stem or myelencephalon is the brain’s Experiments done by William Tavolga provide evidence
posterior.[23] As well as controlling some muscles and that fish have pain and fear responses. For instance, in
body organs, in bony fish at least, the brain stem governs Tavolga’s experiments, toadfish grunted when electrically
shocked and over time they came to grunt at the mere
respiration and osmoregulation.[23]
sight of an electrode.[29]
3.6.2
Sense organs
Most fish possess highly developed sense organs. Nearly
all daylight fish have color vision that is at least as good
as a human’s (see vision in fishes). Many fish also have
chemoreceptors that are responsible for extraordinary
senses of taste and smell. Although they have ears, many
fish may not hear very well. Most fish have sensitive receptors that form the lateral line system, which detects
gentle currents and vibrations, and senses the motion of
nearby fish and prey.[25] Some fish, such as catfish and
sharks, have organs that detect weak electric currents on
the order of millivolt.[26] Other fish, like the South American electric fishes Gymnotiformes, can produce weak
electric currents, which they use in navigation and social
communication.
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.[30][31] Neurons fired in a
pattern resembling human neuronal patterns.[31]
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”.[32]
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
Fish orient themselves using landmarks and may use men- study a year earlier arguing that fish cannot feel pain betal maps based on multiple landmarks or symbols. Fish cause their brains lack a neocortex.[33] However, animal
behavior in mazes reveals that they possess spatial mem- behaviorist Temple Grandin argues that fish could still
have consciousness without a neocortex because “differory and visual discrimination.[27]
ent species can use different brain structures and systems
to handle the same functions.”[31]
Vision Main article: Vision in fishes
Animal welfare advocates raise concerns about the possible suffering of fish caused by angling. Some countries,
Vision is an important sensory system for most species such as Germany have banned specific types of fishing,
of fish. Fish eyes are similar to those of terrestrial and the British RSPCA now formally prosecutes individvertebrates like birds and mammals, but have a more uals who are cruel to fish.[34]
3.9
3.7
Reproductive system
7
Muscular system
20 °C above ambient water temperatures.[35] See also
gigantothermy. Endothermy, though metabolically costly,
is thought to provide advantages such as increased muscle
Main article: Fish locomotion
Most fish move by alternately contracting paired sets of strength, higher rates of central nervous system processing, and higher rates of digestion.
3.9 Reproductive system
Swim bladder of a Rudd (Scardinius erythrophthalmus)
Further information: Fish reproduction and Spawn (biology)
Fish reproductive organs include testes and ovaries. In
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.
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.[36] There may also
be a range of secondary organs that increase reproductive
fitness.
A great white shark off Isla Guadalupe
3.8
Homeothermy
Although most fish are exclusively ectothermic, there are
exceptions.
Certain species of fish maintain elevated body temperatures. Endothermic teleosts (bony fish) are all in the
suborder Scombroidei and include the billfishes, tunas,
and one species of “primitive” mackerel (Gasterochisma
melampus). All sharks in the family Lamnidae – shortfin mako, long fin 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 billfish, which warm only
their eyes and brain, to bluefin tuna and porbeagle sharks
who maintain body temperatures elevated in excess of
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 semicystic spermatogenesis in relation to the release phase of
germ cells in cysts to the seminiferous tubules lumen.[36]
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.[37] Gymnovaries are
the primitive condition found in lungfish, sturgeon, and
bowfin. Cystovaries characterize most teleosts, where
the ovary lumen has continuity with the oviduct.[36] 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
8
4
DISEASES
the surrounding layers characterize the oocyte maturation The newly hatched young of oviparous fish are called
process.[36]
larvae. They are usually poorly formed, carry a large yolk
Postovulatory follicles are structures formed after oocyte sac (for nourishment) and are very different in appearrelease; they do not have endocrine function, present a ance from juvenile and adult specimens. The larval pewide irregular lumen, and are rapidly reabsorbed in a riod in oviparous fish is relatively short (usually only sevprocess involving the apoptosis of follicular cells. A de- eral weeks), and larvae rapidly grow and change appeargenerative process called follicular atresia reabsorbs vitel- ance and structure (a process termed metamorphosis) to
logenic oocytes not spawned. This process can also oc- become juveniles. During this transition larvae must
switch from their yolk sac to feeding on zooplankton prey,
cur, but less frequently, in oocytes in other development
a process which depends on typically inadequate zoo[36]
stages.
plankton density, starving many larvae.
Some fish, like the California sheephead, are
hermaphrodites, having both testes and ovaries ei- In ovoviviparous fish the eggs develop inside the mother’s
ther at different phases in their life cycle or, as in body after internal fertilization but receive little or no
nourishment directly from the mother, depending instead
hamlets, have them simultaneously.
on the yolk. Each embryo develops in its own egg. FamilOver 97% of all known fish are oviparous,[38] that is, iar examples of ovoviviparous fish include guppies, angel
the eggs develop outside the mother’s body. Examples sharks, and coelacanths.
of oviparous fish include salmon, goldfish, cichlids, tuna,
and eels. In the majority of these species, fertilisation Some species of fish are viviparous. In such species the
takes place outside the mother’s body, with the male and mother retains the eggs and nourishes the embryos. Typfemale fish shedding their gametes into the surrounding ically, viviparous fish have a structure analogous to the
water. However, a few oviparous fish practice internal placenta seen in mammals connecting the mother’s blood
supply with that of the embryo. Examples of viviparous
fertilization, with the male using some sort of intromittent
organ to deliver sperm into the genital opening of the fe- fish include the surf-perches, splitfins, and lemon shark.
Some viviparous fish exhibit oophagy, in which the demale, most notably the oviparous sharks, such as the horn
shark, and oviparous rays, such as skates. In these cases, veloping embryos eat other eggs produced by the mother.
This has been observed primarily among sharks, such as
the male is equipped with a pair of modified pelvic fins
the shortfin mako and porbeagle, but is known for a few
known as claspers.
bony fish as well, such as the halfbeak Nomorhamphus
Marine fish can produce high numbers of eggs which are ebrardtii.[39] Intrauterine cannibalism is an even more unoften released into the open water column. The eggs have usual mode of vivipary, in which the largest embryos
an average diameter of 1 millimetre (0.039 in).
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 Nomorham• Egg of lamprey
phus ebrardtii.[39]
• Egg of catshark (mermaids’ purse)
Aquarists commonly refer to ovoviviparous and
• Egg of bullhead shark
• Egg of chimaera
viviparous fish as livebearers.
4 Diseases
Main article: Fish diseases and parasites
An example of zooplankton
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.[40] 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.[41][42]
9
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 socalled “cleaning stations” where other fish congregate and
perform specific movements to attract the attention of the
cleaners.[43] 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.[44]
4.1
Immune system
Immune organs vary by type of fish.[45] 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.
deed, the adaptive immune system as a whole evolved in
an ancestor of all jawed vertebrate.[50]
5 Conservation
The 2006 IUCN Red List names 1,173 fish species
that are threatened with extinction.[51] Included are
species such as Atlantic cod,[52] Devil’s Hole pupfish,[53]
coelacanths,[54] and great white sharks.[55] 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.[56]
5.1 Overfishing
A Whale shark, the world’s largest fish, is classified as
Vulnerable.
Main article: Overfishing
Overfishing is a major threat to edible fish such as cod
and tuna.[57][58] Overfishing eventually causes population
(known as stock) collapse because the survivors cannot
produce enough young to replace those removed. Such
Like chondrostean fish, the major immune tissues of commercial extinction does not mean that the species is
extinct, merely that it can no longer sustain a fishery.
bony fish (or teleostei) include the kidney (especially the
anterior kidney), which houses many different immune One well-studied example of fishery collapse is the
cells.[46] In addition, teleost fish possess a thymus, spleen Pacific sardine Sadinops sagax caerulues fishery off the
and scattered immune areas within mucosal tissues (e.g. California coast. From a 1937 peak of 790,000 long tons
in the skin, gills, gut and gonads). Much like the mam- (800,000 t) the catch steadily declined to only 24,000 long
malian immune system, teleost erythrocytes, neutrophils tons (24,000 t) in 1968, after which the fishery was no
and granulocytes are believed to reside in the spleen longer economically viable.[59]
whereas lymphocytes are the major cell type found in the The main tension between fisheries science and the fishing
thymus.[47][48] In 2006, a lymphatic system similar to that industry is that the two groups have different views on the
in mammals was described in one species of teleost fish, resiliency of fisheries to intensive fishing. In places such
the zebrafish. Although not confirmed as yet, this system as Scotland, Newfoundland, and Alaska the fishing induspresumably will be where naive (unstimulated) T cells ac- try is a major employer, so governments are predisposed
cumulate while waiting to encounter an antigen.[49]
to support it.[60][61] On the other hand, scientists and conB and T lymphocytes bearing immunoglobulins and T cell servationists push for stringent protection, warning that
receptors, respectively, are found in all jawed fishes. In- many stocks could be wiped out within fifty years.[62][63]
10
5.2
6 IMPORTANCE TO HUMANS
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.[64] 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.[65]
5.3
and regions heavily dependent on the sea. In a similar
manner, fish have been tied to trade.
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.[69] The
annual yield from all fisheries worldwide is about 154 million tons,[70] 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.
Exotic species
6.2 Recreation
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.[66]
Carp, snakeheads,[67] 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.
Main articles: Fishkeeping, Recreational fishing 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 fish in ponds, and displayed in
aquariums in homes, offices, or public settings.
Recreational fishing is fishing for pleasure or competition;
it can be contrasted with commercial fishing, which is
fishing for profit. The most common form of recreational
fishing is done with a rod, reel, line, hooks and any one
6 Importance to humans
of a wide range of baits. Angling is a method of fishing,
specifically the practice of catching fish by means of an
6.1 Economic importance
“angle” (hook). Anglers must select the right hook, cast
accurately, and retrieve at the right speed while considerMain articles: Fishing industry, Aquaculture and Fish ing water and weather conditions, species, fish response,
farming
time of the day, and other factors.
Throughout history, humans have utilized fish as a food
6.3 Culture
See also: Mermaid and Merman
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
These fish-farming ponds were created as a cooperative project
species that is unlikely to harm humans.[71] In the Book
in a rural village.
of Jonah a “great fish” swallowed Jonah the Prophet. Legsource. Historically and today, most fish protein has come ends of half-human, half-fish mermaids have featured in
by means of catching wild fish. However, aquaculture, or folklore, like the stories of Hans Christian Andersen.
fish farming, which has been practiced since about 3,500 Fish themes have symbolic significance in many reliBCE. in China,[68] is becoming increasingly important in gions. The fish is used often as a symbol by Christians
many nations. Overall, about one-sixth of the world’s pro- to represent Jesus or Christianity; the gospels also refer
tein is estimated to be provided by fish.[69] That propor- to “fishers of men”[73] and feeding the multitude. In the
tion is considerably elevated in some developing nations dhamma of Buddhism the fish symbolize happiness as
7.1
Shoal or school
11
7.1 Shoal or school
Main article: Shoaling and schooling
A random assemblage of fish merely using some lo-
These goldband fusiliers are schooling because their swimming
is synchronised
Avatar of Vishnu as a Matsya
calised 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.[75]
Examples:
The ichthus is a Christian symbol of a fish signifying that the
person who uses it is a Christian.[72]
• Cichlids congregating at lekking sites form an aggregation.
• Many minnows and characins form shoals.
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.
• Anchovies, herrings and silversides are classic examples of schooling fish.
While school and shoal have different meanings within
biology, they are often treated as synonyms by nonspecialists, with speakers of British English using “shoal”
to describe any grouping of fish, while speakers of
American English often using “school” just as loosely.
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.[74]
7.2
7
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.[76][77][78] Thus a pond that
12
9 NOTES
contained a single species might be said to contain 120
fish. But if the pond contained a total of 120 fish from
three different species, it would be said to contain three
fishes. The distinction is similar to that between people
and peoples.
• List of fish common names
• List of fish families
• Marine biology
• Marine vertebrates
7.3
Finfish
• In biology, the term fish 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
fins.[79] 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.[14]
• In fisheries, the term fish is used as a collective term,
and includes mollusks, crustaceans and any aquatic
animal which is harvested.[80]
• 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.
8
See also
For a topical guide to sharks, see Outline of sharks
• Angling (sport fishing)
• Aquaculture
• Aquarium
• Catch and release
• Deep sea fish
• Fish Acute Toxicity Syndromes
• Fish anatomy
• Fish as food
• Mercury in fish
• Otolith (Bone used for determining the age of a fish)
• Pregnancy (fish)
• Seafood
• Walking fish
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.
[2] Carey, F.G.; Lawson, K.D. (February 1973). “Temperature regulation in free-swimming bluefin tuna”. Comparative Biochemistry and Physiology Part A: Physiology 44
(2): 375–392. 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 Classification, 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. 269–303, 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 Hagfishes Supported by Nuclear DNA–Coded
Genes J Mol Evol (1999) 49:729–735
• Fish development
[8] J. Mallatt, J. Sullivan (1998) 28S and 18S rDNA sequences
support the monophyly of lampreys and hagfishes Molecular Biology and Evolution V 15, Issue 12, pp 1706–1718
• Fishing (fishing for food)
[9] Nelson 2006, pp. 4–5
• Fish intelligence
[10] Nelson 2006, p. 3
• Fishkeeping
[11] Nelson 2006, p. 2
• Forage fish
[12] Helfman, Collette & Facey 1997, p. 3
• Ichthyology
[13] Tree of life web project – Chordates.
13
[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.
[15] Helfman, Collette & Facey 1997, pp. 53–57
[16] Helfman, Collette & Facey 1997, pp. 33–36
[17] Froese, Rainer and Pauly, Daniel, eds.
(2006).
"Periophthalmus barbarus" in FishBase. November 2006
version.
[18] Froese, Rainer and Pauly, Daniel, eds.
(2006).
"Phreatobius cisternarum" in FishBase. November 2006
version.
[35] Block, BA; Finnerty, JR (1993). “Endothermy in fishes:
a phylogenetic analysis of constraints, predispositions,
and selection pressures” (PDF). Environmental Biology of
Fishes 40 (3): 283–302. doi:10.1007/BF00002518.
[36] Guimaraes-Cruz, Rodrigo J., Rodrigo J.; Santos, José E.
dos; Santos, Gilmar B. (July–September 2005). “Gonadal structure and gametogenesis of Loricaria lentiginosa Isbrücker (Pisces, Teleostei, Siluriformes)". Rev.
Bras. Zool. 22 (3): 556–564. doi:10.1590/S010181752005000300005. ISSN 0101-8175.
[19] Planet Catfish. “Cat-eLog: Heptapteridae: Phreatobius:
Phreatobius sp. (1)". Planet Catfish. Retrieved 26
November 2006.
[37] Brito, M.F.G.; Bazzoli, N. (2003). “Reproduction
of the surubim catfish (Pisces, Pimelodidae) in the
São Francisco River, Pirapora Region, Minas Gerais,
Brazil”. Arquivo Brasileiro de Medicina Veterinária
e Zootecnia 55 (5): 624–633. doi:10.1590/S010209352003000500018. ISSN 0102-0935.
[20] Estudo das Espécies Ícticas do Parque Estadual do Cantão,
fish species survey of Cantão (in Portuguese)
[38] Peter Scott: Livebearing Fishes, p. 13. Tetra Press 1997.
ISBN 1-56465-193-2
[21] “Modifications of the Digestive Tract for Holding Air in
Loricariid and Scoloplacid Catfishes” (PDF). Copeia (3):
663–675. 1998. doi:10.2307/1447796. Retrieved 25
June 2009.
[22] Setaro, John F. (1999). Circulatory System. Microsoft
Encarta 99.
[23] Helfman, Collette & Facey 1997, pp. 48–49
[24] Helfman, Collette & Facey 1997, p. 191
[25] Orr, James (1999). Fish. Microsoft Encarta 99. ISBN
0-8114-2346-8.
[26] Albert, J.S., and W.G.R. Crampton. 2005. Electroreception and electrogenesis. pp. 431–472 in The Physiology
of Fishes, 3rd Edition. D.H. Evans and J.B. Claiborne
(eds.). CRC Press.
[27] Journal of Undergraduate Life Sciences. “Appropriate
maze methodology to study learning in fish” (PDF). Retrieved 28 May 2009.
[28] N. A. Campbell and J. B. Reece (2005). Biology (Seventh
ed.). San Francisco, California: Benjamin Cummings.
[29] Dunayer, Joan, “Fish: Sensitivity Beyond the Captor’s
Grasp,” The Animals’ Agenda, July/August 1991, pp. 12–
18
[30] Kirby, Alex (30 April 2003). “Fish do feel pain, scientists
say”. BBC News. Retrieved 4 January 2010.
[31] Grandin, Temple; Johnson, Catherine (2005). Animals in
Translation. New York, New York: Scribner. pp. 183–
184. ISBN 0-7432-4769-8.
[32] “Rose, J.D. 2003. A Critique of the paper: “Do fish have
nociceptors: Evidence for the evolution of a vertebrate
sensory system"" (PDF). Retrieved 21 May 2011.
[33] James D. Rose (2002). “Do Fish Feel Pain?". Retrieved
27 September 2007.
[34] Leake, J. “Anglers to Face RSPCA Check,” The Sunday
Times – Britain, 14 March 2004
[39] Meisner, A & Burns, J: Viviparity in the Halfbeak Genera Dermogenys and Nomorhamphus (Teleostei: Hemiramphidae)" Journal of Morphology 234, pp. 295–317,
1997
[40] Helfman, Collette & Facey 1997, pp. 95–96
[41] R. C. Cipriano (2001), Furunculosis And Other Diseases
Caused By Aeromonas salmonicida. Fish Disease Leaflet
66. U.S. Department of the Interior.
[42] K H Hartman et al. (2004), Koi Herpes Virus (KHV) Disease. Fact Sheet VM-149. University of Florida Institute
of Food and Agricultural Sciences.
[43] Helfman, Collette & Facey 1997, p. 380
[44] 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): 834–838.
doi:10.2307/1442742.
[45] A.G. Zapata, A. Chiba and A. Vara. Cells and tissues of
the immune system of fish. In: The Fish Immune System:
Organism, Pathogen and Environment. Fish Immunology
Series. (eds. G. Iwama and T.Nakanishi,), New York,
Academic Press, 1996, pp. 1–55.
[46] D.P. Anderson. Fish Immunology. (S.F. Snieszko and
H.R. Axelrod, eds), Hong Kong: TFH Publications, Inc.
Ltd., 1977.
[47] S. Chilmonczyk (1992). "The thymus in fish: development
and possible function in the immune response" 2. Annual
Review of Fish Diseases: 181–200.
[48] J.D. Hansen and A.G. Zapata (1998). "Lymphocyte development in fish and amphibians" 166. Immunological
Reviews: 199–220.
[49] Kucher et al., (2006). "Development of the zebrafish lymphatic system requires VegFc signalling" 16. Current Biology: 1244–1248.
14
11 FURTHER READING
[50] 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).
[71] Zollinger, Sue Anne (3 July 2009). “Piranha–Ferocious
Fighter or Scavenging Softie?". A Moment of Science. Indiana Public Media. Retrieved 1 November 2015.
[51] “Table 1: Numbers of threatened species by major groups
of organisms (1996–2004)". iucnredlist.org. Archived
from the original on 30 June 2006. Retrieved 18 January
2006.
[72] Coffman, Elesha (August 8, 2008). “What is the origin
of the Christian fish symbol?". Christianity Today. Retrieved 13 August 2015.
[52] “Gadus morhua (Atlantic Cod)". Iucnredlist.org. Retrieved 21 May 2011.
[74] “Piscis Austrinus”. allthesky.com. The Deep Photographic Guide to the Constellations. Retrieved 1 November 2015.
[53] “Cyprinodon diabolis (Devils Hole Pupfish)".
nredlist.org. Retrieved 21 May 2011.
Iuc-
[54] “Latimeria chalumnae (Coelacanth, Gombessa)". Iucnredlist.org. Retrieved 21 May 2011.
[55] “Carcharodon carcharias (Great White Shark)".
nredlist.org. Retrieved 21 May 2011.
Iuc-
[56] Helfman, Collette & Facey 1997, pp. 449–450
[73] Matthew 4:19
[75] Helfman G., Collette B., & Facey D. (1997). The Diversity of Fishes. Blackwell Publishing. p. 375. ISBN 086542-256-7.
[76] Pauly, Daniel (13 May 2004). “Fish(es)". Darwin’s
Fishes: An Encyclopedia of Ichthyology, Ecology, and
Evolution. Cambridge University Press. p. 77. ISBN
978-1-139-45181-9.
[57] “Call to halt cod 'over-fishing'". BBC News. 5 January
2007. Retrieved 18 January 2006.
[77] Nelson, Joseph S and Paetz, Martin Joseph (1992) The
Fishes of Alberta page 400, University of Alberta. ISBN
9780888642363.
[58] “Tuna groups tackle overfishing”. BBC News. 26 January
2007. Retrieved 18 January 2006.
[78] Helfman, Collette & Facey 1997, p. 5
[59] Helfman, Collette & Facey 1997, p. 462
[79] Nelson, Joseph S. (2006). Fishes of the World. John Wiley & Sons, Inc. p. 2. ISBN 0-471-25031-7.
[60] “UK 'must shield fishing industry'". BBC News. 3 November 2006. Retrieved 18 January 2006.
[80] FAO: Fisheries glossary
[61] “EU fish quota deal hammered out”. BBC News. 21 December 2006. Retrieved 18 January 2006.
10 References
[62] “Ocean study predicts the collapse of all seafood fisheries
by 2050”. Retrieved 13 January 2006.
[63] “Atlantic bluefin tuna could soon be commercially extinct”. Archived from the original on 30 April 2007. Retrieved 18 January 2006.
[64] Helfman, Collette & Facey 1997, p. 463
[65] “Threatened and Endangered Species: Pallid Sturgeon
Scaphirhynchus Fact Sheet”. Retrieved 18 January 2006.
[66] Spinney, Laura (4 August 2005). “The little fish fight
back”. The Guardian (London). Retrieved 18 January
2006.
[67] “Stop That Fish!". The Washington Post. 3 July 2002.
Retrieved 26 August 2007.
[68] Spalding, Mark (July 11, 2013). “Sustainable Ancient
Aquaculture”. National Geographic. Retrieved 13 August 2015.
• 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
[69] Helfman, Gene S. (2007). Fish Conservation: A Guide
to Understanding and Restoring Global Aquatic Biodiversity and Fishery Resources. Island Press. p. 11. ISBN
1597267600.
• 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.
[70] “World Review of Fisheries and Aquaculture” (PDF).
fao.org. Food and Agriculture Organization of the United
Nations. Retrieved 13 August 2015.
• Moyle, Peter B. (1993) Fish: An Enthusiast’s
ISBN
Guide University of California Press.
9780520916654 – good lay text.
15
• Shubin, Neil (2009) Your inner fish: 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 fishes
of Australia and New Guinea
• Fischinfos.de – Illustrated database of the freshwater fishes of Germany (German)
• FishBase online – Comprehensive database with information on over 29,000 fish species
• Fisheries and Illinois Aquaculture Center – Data
outlet for fisheries and aquaculture research center
in the central US
• Philippines Fishes – Database with thousands of
Philippine Fishes photographed in natural habitat
• The Native Fish Conservancy – Conservation and
study of North American freshwater fishes
• United Nation – Fisheries and Aquaculture Department: Fish and seafood utilization
• University of Washington Libraries Digital Collections – Digital collection of freshwater and marine
fish images
• Charles B. Davenport; Ernest Ingersoll (1905).
"Fish". New International Encyclopedia.
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=695509940 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,
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