Fish are aquatic
-bearing animals that lack limbs
. They form a sister group
to the tunicate
s, together forming the olfactores
. Included in this definition are the living hagfish
s, and cartilaginous
and bony fish
as well as various extinct related groups. Around 99% of living fish species are ray-finned fish, belonging to the class Actinopterygii
, with over 95% belonging to the teleost
The earliest organisms that can be classified as fish were soft-bodied chordate
s that first appeared during the Cambrian
period. Although they lacked a true spine
, they possessed notochord
s 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 jaw
s appeared in the Silurian
period, after which many (such as shark
s) became formidable marine predators rather than just the prey of arthropod
Most fish are ectotherm
ic ("cold-blooded"), allowing their body temperatures to vary as ambient temperatures change, though some of the large active swimmers like white shark
can hold a higher core temperature
Fish can communicate in their underwater environments through the use of acoustic communication. Acoustic communication in fish involves the transmission of acoustic signals from one individual of a species
to another. The production of sounds as a means of communication among fish is most often used in the context of feeding, aggression or courtship behaviour.
The sounds emitted by fish can vary depending on the species and stimulus involved. They can produce either stridulatory sounds by moving components of the skeletal system, or can produce non-stridulatory sounds by manipulating specialized organs such as the swimbladder
Fish are abundant in most bodies of water. They can be found in nearly all aquatic environments, from high mountain streams (e.g., char
) to the abyssal
and even hadal
depths of the deepest oceans (e.g., cusk-eels
), although no species has yet been documented in the deepest 25% of the ocean. With 34,300 described species, fish exhibit greater species diversity than any other group of vertebrates.
Fish are an important resource for humans worldwide, especially as food
. Commercial and subsistence fishers hunt fish in wild fisheries
them in ponds or in cages in the ocean (in aquaculture
). They are also caught by recreational fishers
, kept as pets, raised by fishkeepers
, and exhibited in public aquaria
. Fish have had a role in culture through the ages, serving as deities, religious symbols, and as the subjects of art, books and movies.
s emerged within lobe-finned fishes
, so cladistically
they are fish as well. However, traditionally fish are rendered paraphyletic
by excluding the tetrapods (i.e., the amphibian
s and mammal
s which all descended from within the same ancestry). Because in this manner the term "fish" is defined negatively as a paraphyletic group, it is not considered a formal taxonomic grouping in systematic biology
, unless it is used in the cladistic
sense, including tetrapods. The traditional term pisces (also ichthyes) is considered a typological
, but not a phylogenetic
The word for ''fish'' in English and the other Germanic languages
''fisks'') is inherited from Proto-Germanic
, and is related to the Latin
''piscis'' and Old Irish
''īasc'', though the exact root is unknown; some authorities reconstruct an Proto-Indo-European
root ''*peysk-'', attested only in Italic
, and Germanic
The English word once had a much broader usage than its current biological meaning. Names such as starfish
attest to almost any fully aquatic animal (including whales) once being 'fish'. "Correcting" such names (e.g. to 'sea star') is an attempt to retroactively apply the current meaning of 'fish' to words that were coined when it had a different meaning.
Fish, as vertebrata, developed as sister of the tunicata. As the tetrapods emerged deep within the fishes group, as sister of the lungfish, characteristics of fish are typically shared by tetrapods, including having vertebrae and a cranium.
pustulosus'', showing the shearing structures ("teeth") on its oral surface; from the Devonian
Early fish from the fossil record are represented by a group of small, jawless, armored fish known as ostracoderm
s. Jawless fish lineages are mostly extinct. An extant clade, the lamprey
s may approximate ancient pre-jawed fish. The first jaws are found in Placodermi
fossils. They lacked distinct teeth, having instead the oral surfaces of their jaw plates modified to serve the various purposes of teeth. The diversity of jawed vertebrates may indicate the evolutionary advantage of a jawed mouth
. It is unclear if the advantage of a hinged jaw is greater biting force, improved respiration, or a combination of factors.
Fish may have evolved from a creature similar to a coral-like sea squirt
, whose larvae resemble primitive fish in important ways. The first ancestors of fish may have kept the larval form into adulthood
(as some sea squirts do today), although perhaps the reverse is the case.
Fish are a paraphyletic
group: that is, any clade
containing all fish also contains the tetrapod
s, 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 divides fish into three extant classes
, and with extinct forms sometimes classified within the tree, sometimes as their own classes:
[Romer, A.S. & T.S. Parsons. 1977. ''The Vertebrate Body.'' 5th ed. Saunders, Philadelphia. (6th ed. 1985)]
* Class Agnatha
** Subclass Cyclostomata
** Subclass Ostracodermi
(armoured jawless fish) †
* Class Chondrichthyes
** Subclass Elasmobranchii
s and rays
** Subclass Holocephali
s and extinct relatives)
* Class Placodermi
(armoured fish) †
* Class Acanthodii
("spiny sharks", sometimes classified under bony fishes)†
* Class Osteichthyes
** Subclass Actinopterygii
(ray finned fishes)
** Subclass Sarcopterygii
(fleshy finned fishes, ancestors of tetrapods)
The above scheme is the one most commonly encountered in non-specialist and general works. Many of the above groups are paraphyletic, in that they have given rise to successive groups: Agnathans are ancestral to Chondrichthyes, who again have given rise to Acanthodiians, the ancestors of Osteichthyes. With the arrival of phylogenetic nomenclature
, the fishes has been split up into a more detailed scheme, with the following major groups:
* Class Myxini (hagfish
* Class Pteraspidomorphi
† (early jawless fish)
* Class Thelodonti
* Class Anaspida
* Class Petromyzontida or Hyperoartia
** Petromyzontidae (lamprey
* Class Conodont
a (conodonts) †
* Class Cephalaspidomorphi
† (early jawless fish)
** (unranked) Galeaspida
** (unranked) Pituriaspida
** (unranked) Osteostraci
* Infraphylum Gnathostomata
** Class Placodermi
† (armoured fish)
** Class Chondrichthyes
** Class Acanthodii
† (spiny sharks)
** Superclass Osteichthyes
*** Class Actinopterygii
**** Subclass Chondrostei
***** Order Acipenseriformes
s and paddlefish
***** Order Polypteriformes
es and bichir
**** Subclass Neopterygii
***** Infraclass Holostei
s and bowfin
***** Infraclass Teleostei
(many orders of common fish)
*** Class Sarcopterygii
**** Subclass Actinistia
**** Subclass Dipnoi (lungfish
, sister group to the tetrapods
† – indicates extinct taxon
Some palaeontologists contend that because Conodonta
s, they are primitive fish. For a fuller treatment of this taxonomy, see the vertebrate
The position of hagfish
in the phylum Chordata is not settled. Phylogenetic research in 1998 and 1999 supported the idea that the hagfish and the lampreys form a natural group, the Cyclostomata
, that is a sister group of the Gnathostomata.
The various fish groups account for more than half of vertebrate species. There are almost 28,000 known extant
species, of which almost 27,000 are bony fish, with 970 sharks, rays, and chimeras
and about 108 hagfish and lampreys. A third of these species fall within the nine largest families; from largest to smallest, these families are Cyprinidae
, and Scorpaenidae
. About 64 families are monotypic
, containing only one species. The final total of extant species may grow to exceed 32,500.
Hornhai (Heterodontus francisci).JPG|Chondrichthyes
'' is one of only two animal species known to have blue colouring because of cellular pigment.]]
The term "fish" most precisely describes any non-tetrapod craniate
(i.e. an animal with a skull and in most cases a backbone) that has gill
s throughout life and whose limbs, if any, are in the shape of fins. Unlike groupings such as birds or mammal
s, fish are not a single clade
but a paraphyletic
collection of taxa
, including hagfish
s, sharks and rays
, ray-finned fish
s, and lungfish
. Indeed, lungfish and coelacanths are closer relatives of tetrapod
s (such as mammal
s, birds, amphibians
, etc.) than of other fish such as ray-finned fish
or sharks, so the last common ancestor
of all fish is also an ancestor to tetrapods. As paraphyletic groups are no longer recognised in modern systematic biology
, the use of the term "fish" as a biological group must be avoided.
Many types of aquatic animal
s commonly referred to as "fish" are not fish in the sense given above; examples include shellfish
. In earlier times, even biologists did not make a distinction – sixteenth century natural historians classified also seals
, whales, amphibian
s, even hippopotamus
es, as well as a host of aquatic invertebrates, as fish.
However, according to the definition above, all mammals, including cetacean
s like whales and dolphins, are not fish. In some contexts, especially in aquaculture
, the true fish are referred to as finfish (or fin fish) to distinguish them from these other animals.
A typical fish is ectothermic
, has a streamlined
body for rapid swimming, extracts oxygen from water using gills or uses an accessory breathing organ to breathe atmospheric oxygen, has two sets of paired fins, usually one or two (rarely three) dorsal fins, an anal fin, and a tail fin, has jaws, has skin that is usually covered with scales
, and lays eggs.
Each criterion has exceptions. Tuna
, and some species of shark
s show some warm-blooded adaptations
– they can heat their bodies significantly above ambient water temperature. Streamlining and swimming performance varies from fish such as tuna
, and jacks
that can cover 10–20 body-lengths per second to species such as eel
s and rays
that swim no more than 0.5 body-lengths per second. Many groups of freshwater fish extract oxygen from the air as well as from the water using a variety of different structures. Lungfish
have paired lungs similar to those of tetrapods, gourami
s have a structure called the labyrinth organ
that performs a similar function, while many catfish, such as ''Corydoras
'' extract oxygen via the intestine or stomach. Body shape and the arrangement of the fins is highly variable, covering such seemingly un-fishlike forms as seahorse
, and gulpers
. Similarly, the surface of the skin may be naked (as in moray eel
s), 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 gar
s and bichir
, and ctenoid
(these last two are found on most bony fish
). There are even fish that live mostly on land or lay their eggs on land near water. Mudskipper
s feed and interact with one another on mudflats and go underwater to hide in their burrows. A single, undescribed species
'', has been called a true "land fish" as this worm-like catfish strictly lives among waterlogged leaf litter
. Many species live in underground lake
s, underground river
s or aquifer
s and are popularly known as cavefish
Fish range in size from the huge whale shark
to the tiny stout infantfish
diversity is roughly divided equally between marine (oceanic) and freshwater
ecosystems. Coral reef
s in the Indo-Pacific
constitute the center of diversity for marine fishes, whereas continental freshwater fishes are most diverse in large river basin
s of tropical rainforest
s, especially the Amazon
, and Mekong
basins. More than 5,600 fish species inhabit Neotropic
al freshwaters alone, such that Neotropical fish
es represent about 10% of all vertebrate
species on the Earth. Exceptionally rich sites in the Amazon basin, such as Cantão State Park
, can contain more freshwater fish species than occur in all of Europe.
The deepest living fish in the ocean so far found is the Mariana snailfish (''Pseudoliparis swirei
'') which lives at deeps of 8,000 meters (26,200 feet) along the Mariana Trench near Guam.
Anatomy and physiology
Most fish exchange gases using gill
s on either side of the pharynx
. Gills consist of threadlike structures called filaments
. Each filament contains a capillary
network that provides a large surface area
for exchanging oxygen
and carbon dioxide
. Fish exchange gases by pulling oxygen-rich water through their mouths and pumping it over their gills. In some fish, capillary blood flows in the opposite direction to the water, causing countercurrent exchange
. The gills push the oxygen-poor water out through openings in the sides of the pharynx. Some fish, like shark
s and lamprey
s, possess multiple gill openings. However, bony fish
have a single gill opening on each side. This opening is hidden beneath a protective bony cover called an operculum
s have external gills, a very primitive feature that they share with larval amphibian
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
, and Scoloplacidae
absorb air through their digestive tracts.
, with the exception of the Australian lungfish
, and bichir
s have paired lungs similar to those of tetrapod
s and must surface to gulp fresh air through the mouth and pass spent air out through the gills. Gar
have a vascularized swim bladder that functions in the same way. Loaches
, 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 gourami
s and betta
s) 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
, and the Clariidae
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 cichlid
s, quickly suffocate, while air-breathers survive for much longer, in some cases in water that is little more than wet mud. At the most extreme, some air-breathing fish are able to survive in damp burrows for weeks without water, entering a state of aestivation
(summertime hibernation) until water returns.
Air breathing fish can be divided into obligate air breathers and facultative air breathers. Obligate air breathers, such as the African lungfish
, ''must'' breathe air periodically or they suffocate. Facultative air breathers, such as the catfish ''Hypostomus plecostomus
'', only breathe air if they need to and will otherwise rely on their gills for oxygen. Most air breathing fish are facultative air breathers that avoid the energetic cost of rising to the surface and the fitness cost of exposure to surface predators.
Fish have a closed-loop circulatory system
. The heart
pumps the blood in a single loop throughout the body. In most fish, the heart consists of four parts, including two chambers and an entrance and exit.
The first part is the sinus venosus
, a thin-walled sac that collects blood from the fish's vein
s before allowing it to flow to the second part, the atrium
, which is a large muscular chamber. The atrium serves as a one-way antechamber, sends blood to the third part, ventricle
. The ventricle is another thick-walled, muscular chamber and it pumps the blood, first to the fourth part, bulbus arteriosus
, a large tube, and then out of the heart. The bulbus arteriosus connects to the aorta
, through which blood flows to the gills for oxygenation.
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 enzyme
s and absorb nutrients. Organs such as the liver
add enzymes and various chemicals as the food moves through the digestive tract. The intestine completes the process of digestion and nutrient absorption.
As with many aquatic animals, most fish release their nitrogenous wastes as ammonia
. Some of the wastes diffuse
through the gills. Blood wastes are filtered
by the kidney
Saltwater fish tend to lose water because of osmosis
. Their kidneys return water to the body. The reverse happens in freshwater fish
: they tend to gain water osmotically. Their kidneys produce dilute urine for excretion. Some fish have specially adapted kidneys that vary in function, allowing them to move from freshwater to saltwater.
The scales of fish originate from the mesoderm
(skin); they may be similar in structure to teeth.
Sensory and nervous system
Central nervous system
Fish typically have quite small brains relative to body size compared with other vertebrates, typically one-fifteenth the brain mass of a similarly sized bird or mammal. However, some fish have relatively large brains, most notably mormyrids
s, which have brains about as massive relative to body weight as birds and marsupial
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 nostril
s via the two olfactory nerve
s. The olfactory lobes are very large in fish that hunt primarily by smell, such as hagfish, sharks, and catfish. Behind the olfactory lobes is the two-lobed telencephalon
, the structural equivalent to the cerebrum
in higher vertebrates
. In fish the telencephalon is concerned mostly with olfaction
. Together these structures form the forebrain.
Connecting the forebrain to the midbrain is the diencephalon
(in the diagram, this structure is below the optic lobes and consequently not visible). The diencephalon performs functions associated with hormone
s and homeostasis
. The pineal body
lies just above the diencephalon. This structure detects light, maintains circadian
rhythms, and controls color changes.
(or mesencephalon) contains the two optic lobes
. These are very large in species that hunt by sight, such as rainbow trout
The hindbrain (or metencephalon
) is particularly involved in swimming and balance. The cerebellum is a single-lobed structure that is typically the biggest part of the brain. Hagfish and lamprey
s have relatively small cerebellae, while the mormyrid
cerebellum is massive and apparently involved in their electrical sense
The brain stem (or myelencephalon
) is the brain's posterior. As well as controlling some muscles and body organs, in bony fish at least, the brain stem governs respiration
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.
Some fish, such as catfish and sharks, have the ampullae of Lorenzini
s that detect weak electric currents on the order of millivolt. Other fish, like the South American electric fishes Gymnotiformes
, can produce weak electric currents, which they use in navigation and social communication.
Fish orient themselves using landmarks and may use mental maps based on multiple landmarks or symbols. Fish behavior in mazes reveals that they possess spatial memory and visual discrimination.
is an important sensory system
for most species of fish. Fish eyes are similar to those of terrestrial vertebrate
s like birds
and mammals, but have a more spherical lens
. Their retina
s generally have both rods
and photopic vision
), and most species have colour vision
. Some fish can see ultraviolet
and some can see polarized light
. Amongst jawless fish
, the lamprey
has well-developed eyes, while the hagfish
has only primitive eyespots
. Fish vision shows adaptation
to their visual environment, for example deep sea fish
es have eyes suited to the dark environment.
is an important sensory system for most species of fish. Fish sense sound using their lateral line
s and their ears
New research has expanded preconceptions about the cognitive capacities of fish. For example, manta rays
have exhibited behavior linked to self-awareness
in mirror test
cases. Placed in front of a mirror, individual rays engaged in contingency testing, that is, repetitive behavior aiming to check whether their reflection's behavior mimics their body movement.
have also passed the mirror test in a 2018 scientific study.
Cases of tool use have also been noticed, notably in the Choerodon
family, in archerfish
and Atlantic cod
Capacity for pain
Experiments done by William Tavolga provide evidence that fish have pain
and fear responses. For instance, in Tavolga's experiments, toadfish
grunted when electrically shocked and over time they came to grunt at the mere sight of an electrode.
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.
Neurons fired in a pattern resembling human neuronal patterns.
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". Rose argues that since fish brains are so different from human brains, fish are probably not conscious in the manner humans are, so that reactions similar to human reactions to pain instead have other causes. Rose had published a study a year earlier arguing that fish cannot feel pain because their brains lack a neocortex
. However, animal behaviorist Temple Grandin
argues that fish could still have consciousness without a neocortex because "different species can use different brain structures and systems to handle the same functions."
Animal welfare advocates raise concerns about the possible suffering
of fish caused by angling. Some countries, such as Germany have banned specific types of fishing, and the British RSPCA
now formally prosecutes individuals who are cruel to fish.
In 2019, scientists have shown that members of the monogamous
species Amatitlania siquia
exhibit pessimistic behavior when they are prevented from being with their partner.
thumb|left|Swim bladder of a rudd (''alt=Photo of white bladder that consists of a rectangular section and a banana-shaped section connected by a much thinner element
Most fish move by alternately contracting paired sets of muscles on either side of the backbone. These contractions form S-shaped curves that move down the body. As each curve reaches the back fin, backward force is applied to the water, and in conjunction with the fins, moves the fish forward. The fish's fins function like an airplane's flaps. Fins also increase the tail's surface area, increasing speed. The streamlined body of the fish decreases the amount of friction from the water. Since body tissue is denser than water, fish must compensate for the difference or they will sink. Many bony fish have an internal organ called a swim bladder
that adjusts their buoyancy through manipulation of gases.
Although most fish are exclusively ectothermic
, there are exceptions. The only known bony fishes (infraclass Teleostei
) that exhibit endothermy
are in the suborder Scombroidei
– which includes the billfish
es, tunas, and the butterfly kingfish
, a basal
species of mackerel
– and also the opah
. The opah, a lampriform
, was demonstrated in 2015 to utilize "whole-body endothermy", generating heat with its swimming muscles to warm its body while countercurrent exchange (as in respiration) minimizes heat loss. It is able to actively hunt prey such as squid and swim for long distances due to the ability to warm its entire body, including its heart, which is a trait typically found in only mammals and birds (in the form of homeothermy
). In the cartilaginous fishes (class Chondrichthyes
), sharks of the families Lamnidae
(porbeagle, mackerel, salmon, and great white sharks) and Alopiidae
(thresher sharks) exhibit endothermy. The degree of endothermy varies from the billfishes, which warm only their eyes and brain, to the bluefin tuna
and the porbeagle shark
, which maintain body temperatures in excess of above ambient water temperatures.
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
Fish reproductive organs include testicle
s and ovaries
. In most species, gonads are paired organs of similar size, which can be partially or totally fused.
There may also be a range of secondary organs that increase reproductive fitness.
In terms of spermatogonia
distribution, the structure of teleost
s testes has two types: in the most common, spermatogonia occur all along the seminiferous tubule
s, while in atherinomorph
fish they are confined to the distal
portion of these structures. Fish can present cystic or semi-cystic spermatogenesis
in relation to the release phase of germ cells in cysts to the seminiferous tubules lumen
Fish ovaries may be of three types: gymnovarian, secondary gymnovarian or cystovarian. In the first type, the oocyte
s are released directly into the coelom
ic 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
Gymnovaries are the primitive condition found in lungfish
, and bowfin
. Cystovaries characterize most teleosts, where the ovary lumen has continuity with the oviduct.
Secondary gymnovaries are found in salmonid
s and a few other teleosts.
development in teleosts fish varies according to the group, and the determination of oogenesis dynamics allows the understanding of maturation and fertilization processes. Changes in the nucleus
, ooplasm, and the surrounding layers characterize the oocyte maturation process.
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.
Some fish, like the California sheephead
, are hermaphrodite
s, having both testes and ovaries either at different phases in their life cycle or, as in hamlets
, have them simultaneously.
Over 97% of all known fish are oviparous
[Peter Scott: ''Livebearing Fishes'', p. 13. Tetra Press 1997. ]
that is, the eggs develop outside the mother's body. Examples of oviparous fish include salmon
, and eel
s. In the majority of these species, fertilisation takes place outside the mother's body, with the male and female fish shedding their gamete
s into the surrounding water. However, a few oviparous fish practice internal fertilization, with the male using some sort of intromittent organ
to deliver sperm into the genital opening of the female, most notably the oviparous sharks, such as the horn shark
, and oviparous rays, such as skates
. In these cases, the male is equipped with a pair of modified pelvic fin
s known as clasper
Marine fish can produce high numbers of eggs which are often released into the open water column. The eggs have an average diameter of .
File:Oeufs002b,57.png|Egg of lamprey
File:Oeufs002b,54.png|Egg of catshark (mermaids' purse)
File:Oeufs002b,55.png|Egg of bullhead shark
File:Oeufs002b,56.png|Egg of chimaera
The newly hatched young of oviparous fish are called larvae
. They are usually poorly formed, carry a large yolk sac
(for nourishment), and are very different in appearance from juvenile and adult specimens. The larval period in oviparous fish is relatively short (usually only several weeks), and larvae rapidly grow and change appearance and structure (a process termed metamorphosis
) to become juveniles. During this transition larvae must switch from their yolk sac to feeding on zooplankton
prey, a process which depends on typically inadequate zooplankton density, starving many larvae.
fish the eggs develop inside the mother's body after internal fertilization but receive little or no nourishment directly from the mother, depending instead on the yolk
. Each embryo develops in its own egg. Familiar examples of ovoviviparous fish include guppies
, angel shark
s, and coelacanth
Some species of fish are viviparous
. In such species the mother retains the eggs and nourishes the embryos. Typically, viviparous fish have a structure analogous to the placenta
seen in mammals
connecting the mother's blood supply with that of the embryo. Examples of viviparous fish include the surf-perches
, and lemon shark
. Some viviparous fish exhibit oophagy
, in which the developing embryos eat other eggs produced by the mother. This has been observed primarily among sharks, such as the shortfin mako
, but is known for a few bony fish as well, such as the halfbeak
[Meisner, A & Burns, J: Viviparity in the Halfbeak Genera ''Dermogenys'' and ''Nomorhamphus'' (Teleostei: Hemiramphidae)" ''Journal of Morphology'' 234, pp. 295–317, 1997] Intrauterine cannibalism
is an even more unusual mode of vivipary, in which the largest embryos eat weaker and smaller siblings. This behavior is also most commonly found among sharks, such as the grey nurse shark
, but has also been reported for ''Nomorhamphus ebrardtii''.
commonly refer to ovoviviparous and viviparous fish as livebearer
Acoustic communication in fish
Acoustic communication in fish involves the transmission of acoustic signals from one individual of a species to another. The production of sounds as a means of communication among fish is most often used in the context of feeding, aggression or courtship behaviour.
The sounds emitted can vary depending on the species and stimulus involved. Fish can produce either stridulatory sounds by moving components of the skeletal system, or can produce non-stridulatory sounds by manipulating specialized organs such as the swimbladder.
Stridulatory sound producing mechanisms
There are some species of fish that can produce sounds by rubbing or grinding their bones together. These noises produced by bone-on-bone interactions are known as 'stridulatory sounds'.
An example of this is seen in ''Haemulon flavolineatum
'', a species commonly referred to as the 'French grunt fish', as it produces a grunting noise by grinding its teeth together.
This behaviour is most pronounced when the ''H. flavolineatum
'' is in distress situations.
The grunts produced by this species of fishes generate a frequency of approximately 700 Hz, and last approximately 47 milliseconds.
The ''H. flavolineatum
'' does not emit sounds with frequencies greater than 1000 Hz, and does not detect sounds that have frequencies greater than 1050 Hz.
In a study conducted by Oliveira et al. (2014), the longsnout seahorse, ''Hippocampus reidi
'', was recorded producing two different categories of sounds; ‘clicks’ and ‘growls’. The sounds emitted by the ''H. reidi
'' are accomplished by rubbing their coronet bone across the grooved section of their neurocranium. ‘Clicking’ sounds were found to be primarily produced during courtship and feeding, and the frequencies of clicks were within the range of 50 Hz-800 Hz.
The frequencies were noted to be on the higher end of the range during spawning periods, when the female and male fishes were less than fifteen centimeters apart.
Growl sounds were produced when the ''H. reidi
'' encountered stressful situations, such as handling by researchers.
The 'growl' sounds consist of a series of sound pulses and are emitted simultaneously with body vibrations.
Non-stridulatory sound producing mechanisms
Some fish species create noise by engaging specialized muscles that contract and cause swimbladder vibrations.
Oyster toadfish produce loud grunting sounds by contracting muscles located along the sides of their swim bladder, known as sonic muscles
Female and male toadfishes emit short-duration grunts, often as a fright response. In addition to short-duration grunts, male toadfishes produce “boat whistle calls”. These calls are longer in duration, lower in frequency, and are primarily used to attract mates. The sounds emitted by the ''O. tao'' have frequency range of 140 Hz to 260 Hz. The frequencies of the calls depend on the rate at which the sonic muscles contract.
The red drum, ''Sciaenops ocellatus'', produces drumming sounds by vibrating its swimbladder. Vibrations are caused by the rapid contraction of sonic muscles that surround the dorsal aspect of the swimbladder. These vibrations result in repeated sounds with frequencies that range from 100 to >200 Hz. The ''S. Ocellatus'' can produce different calls depending on the stimuli involved. The sounds created in courtship situations are different from those made during distressing events such as predatorial attacks. Unlike the males of the ''S. Ocellatus'' species, the females of this species don't produce sounds and lack sound-producing (sonic) muscles.
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. 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.
Some species use cleaner fish to remove external parasites. The best known of these are the Bluestreak cleaner wrasses of the genus ''Labroides'' found on coral reefs in the Indian and Pacific oceans. These small fish maintain so-called "cleaning stations" where other fish congregate and perform specific movements to attract the attention of the cleaners. 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''.
Immune organs vary by type of fish. In the jawless fish (lampreys and hagfish), true lymphoid organs are absent. These fish rely on regions of lymphoid tissue within other organs to produce immune cells. For example, erythrocytes, macrophages and plasma cells are produced in the anterior kidney (or pronephros) and some areas of the gut (where granulocytes mature.) They resemble primitive bone marrow in hagfish. Cartilaginous fish (sharks and rays) have a more advanced immune system. They have three specialized organs that are unique to Chondrichthyes; the epigonal organs (lymphoid tissue similar to mammalian bone) that surround the gonads, the Leydig's organ within the walls of their esophagus, and a spiral valve in their intestine. These organs house typical immune cells (granulocytes, lymphocytes and plasma cells). They also possess an identifiable thymus and a well-developed spleen (their most important immune organ) where various lymphocytes, plasma cells and macrophages develop and are stored. Chondrostean fish (sturgeons, paddlefish, and bichirs) possess a major site for the production of granulocytes within a mass that is associated with the meninges (membranes surrounding the central nervous system.) Their heart is frequently covered with tissue that contains lymphocytes, reticular cells and a small number of macrophages. The chondrostean kidney is an important hemopoietic organ; where erythrocytes, granulocytes, lymphocytes and macrophages develop.
Like chondrostean fish, the major immune tissues of bony fish (or teleostei) include the kidney (especially the anterior kidney), which houses many different immune cells. In addition, teleost fish possess a thymus, spleen and scattered immune areas within mucosal tissues (e.g. in the skin, gills, gut and gonads). Much like the mammalian immune system, teleost erythrocytes, neutrophils and granulocytes are believed to reside in the spleen whereas lymphocytes are the major cell type found in the thymus. In 2006, a lymphatic system similar to that in mammals was described in one species of teleost fish, the zebrafish. Although not confirmed as yet, this system presumably will be where naive (unstimulated) T cells accumulate while waiting to encounter an antigen.
B and T lymphocytes bearing immunoglobulins and T cell receptors, respectively, are found in all jawed fishes. Indeed, the adaptive immune system as a whole evolved in an ancestor of all jawed vertebrate.
The 2006 IUCN Red List names 1,173 fish species that are threatened with extinction. Included are species such as Atlantic cod, Devil's Hole pupfish, coelacanths, and great white sharks. 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 pool.
Overfishing is a major threat to edible fish such as cod and tuna. Overfishing eventually causes population (known as stock) collapse because the survivors cannot produce enough young to replace those removed. Such commercial extinction does not mean that the species is extinct, merely that it can no longer sustain a fishery.
One well-studied example of fishery collapse is the Pacific sardine ''Sadinops sagax caerulues'' fishery off the California coast. From a 1937 peak of the catch steadily declined to only in 1968, after which the fishery was no longer economically viable.
The main tension between fisheries science and the fishing industry is that the two groups have different views on the resiliency of fisheries to intensive fishing. In places such as Scotland, Newfoundland, and Alaska the fishing industry is a major employer, so governments are predisposed to support it. On the other hand, scientists and conservationists push for stringent protection, warning that many stocks could be wiped out within fifty years.
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. 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.
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. Carp, snakeheads, tilapia, European perch, brown trout, rainbow trout, and sea lampreys are other examples of fish that have caused problems by being introduced into alien environments.
Importance to humans
Throughout history, humans have utilized fish as a food source. Historically and today, most fish protein has come by means of catching wild fish. However, aquaculture, or fish farming, which has been practiced since about 3,500 BCE. in China, is becoming increasingly important in many nations. Overall, about one-sixth of the world's protein is estimated to be provided by fish.
That proportion is considerably elevated in some developing nations and regions heavily dependent on the sea. In a similar manner, fish have been tied to trade.
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. The annual yield from all fisheries worldwide is about 154 million tons, 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.
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 primarily for pleasure or competition; it can be contrasted with commercial fishing, which is fishing for profit, or artisanal fishing, which is fishing primarily for food. The most common form of recreational fishing is done with a rod, reel, line, hooks, and any one of a wide range of baits. Recreational fishing is particularly popular in North America and Europe and state, provincial, and federal government agencies actively management target fish species. Angling is a method of fishing, specifically the practice of catching fish by means of an "angle" (hook). Anglers must select the right hook, cast accurately, and retrieve at the right speed while considering water and weather conditions, species, fish response, time of the day, and other factors.
Fish themes have symbolic significance in many religions. In ancient Mesopotamia, fish offerings were made to the gods from the very earliest times.
Fish were also a major symbol of Enki, the god of water. Fish frequently appear as filling motifs in cylinder seals from the Old Babylonian ( 1830 BC – 1531 BC) and Neo-Assyrian (911–609 BC) periods. Starting during the Kassite Period ( 1600 BC – 1155 BC) and lasting until the early Persian Period (550–30 BC), healers and exorcists dressed in ritual garb resembling the bodies of fish. During the Seleucid Period (312–63 BC), the legendary Babylonian culture hero Oannes, described by Berossus, was said to have dressed in the skin of a fish. Fish were sacred to the Syrian goddess Atargatis and, during her festivals, only her priests were permitted to eat them.
In the Book of Jonah, a work of Jewish literature probably written in the fourth century BC, the central figure, a prophet named Jonah, is swallowed by a giant fish after being thrown overboard by the crew of the ship he is travelling on. The fish later vomits Jonah out on shore after three days. This book was later included as part of the Hebrew Bible, or Christian Old Testament, and a version of the story it contains is summarized in Surah 37:139-148 of the Quran. Early Christians used the ''ichthys'', a symbol of a fish, to represent Jesus, because the Greek word for fish, ΙΧΘΥΣ Ichthys, could be used as an acronym for "Ίησοῦς Χριστός, Θεοῦ Υἱός, Σωτήρ" (Iesous Christos, Theou Huios, Soter), meaning "Jesus Christ, Son of God, Saviour". The gospels also refer to "fishers of men" and feeding the multitude. In the dhamma of Buddhism, the fish symbolize happiness as they have complete freedom of movement in the water. Often drawn in the form of carp which are regarded in the Orient as sacred on account of their elegant beauty, size and life-span.
Among the deities said to take the form of a fish are Ika-Roa of the Polynesians, Dagon of various ancient Semitic peoples, the shark-gods of 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 fish constellation in the night sky, Piscis Austrinus.
Fish feature prominently in art and literature, in movies such as ''Finding Nemo'' and books such as ''The Old Man and the Sea''. Large fish, particularly sharks, have frequently been the subject of horror movies and thrillers, most notably the novel ''Jaws'', which spawned a series of films of the same name that in turn inspired similar films or parodies such as ''Shark Tale'' and ''Snakehead Terror''. Piranhas are shown in a similar light to sharks in films such as ''Piranha''; however, contrary to popular belief, the red-bellied piranha is actually a generally timid scavenger species that is unlikely to harm humans. Legends of half-human, half-fish mermaids have featured in folklore, including the stories of Hans Christian Andersen.
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.
Thus a pond would be said to contain 120 fish if all were from a single species or 120 fishes if these included a mix of several species. The distinction is similar to that between people and peoples.
True fish and 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. 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.
* In fisheries, the term ''fish'' is used as a collective term, and includes mollusks, crustaceans and any aquatic animal which is harvested.
* The strict biological definition of a fish, above, is sometimes called a ''true fish''. True fish are also referred to as ''finfish'' or ''fin fish'' to distinguish them from other aquatic life harvested in fisheries or aquaculture.
Shoal or school
A random assemblage of fish merely using some localised resource such as food or nesting sites is known simply as an ''aggregation''. When fish come together in an interactive, social grouping, then they may be forming either a ''shoal'' or a ''school'' depending on the degree of organisation. A ''shoal'' is a loosely organised group where each fish swims and forages independently but is attracted to other members of the group and adjusts its behaviour, such as swimming speed, so that it remains close to the other members of the group. ''Schools'' of fish are much more tightly organised, synchronising their swimming so that all fish move at the same speed and in the same direction. Shoaling and schooling behaviour is believed to provide a variety of advantages.
* Cichlids congregating at lekking sites form an ''aggregation''.
* Many minnows and characins form ''shoals''.
* Anchovies, herrings and silversides are classic examples of ''schooling'' fish.
While the words "school" and "shoal" have different meanings within biology, the distinctions are often ignored by non-specialists who treat the words as synonyms. Thus speakers of British English commonly use "shoal" to describe any grouping of fish, and speakers of American English commonly use "school" just as loosely.
* Moyle, Peter B. (1993
''Fish: An Enthusiast's Guide''
University of California Press. – good lay text.
– Illustrated database of freshwater fishes of Australia and New Guinea
– Comprehensive database with information on over 29,000 fish species
– Database with thousands of Philippine Fishes photographed in natural habitat
The Native Fish Conservancy
– Conservation and study of North American freshwater fishes
– Fisheries and Aquaculture Department: Fish and seafood utilization
University of Washington Libraries Digital Collections
– Digital collection of freshwater and marine fish images
Category:Obsolete vertebrate taxa