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Teleost

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Teleost
Temporal range:Early Triassic–Recent[1][2]
Teleosts of different orders, painted byCastelnau,1856 (left to right, top to bottom):Fistularia tabacaria(Syngnathiformes),Mylossoma duriventre(Characiformes),Mesonauta acora(Cichliformes),Corydoras splendensandPseudacanthicus spinosus(Siluriformes),Acanthurus coeruleus(Acanthuriformes),Stegastes pictus(Incertae sedis,Pomacentridae)
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Infraclass: Teleosteomorpha
Division: Teleostei
J. P. Müller,1845[3]
Subdivisions

See text

Teleostei(/ˌtɛliˈɒstiaɪ/;Greekteleios"complete" +osteon"bone" ), members of which are known asteleosts(/ˈtɛliɒsts,ˈtiːli-/),[4]is, by far, the largestinfraclassin the classActinopterygii,the ray-finned fishes,[a]and contains 96% of allextantspecies offish.Teleosts are arranged into about 40ordersand 448families.Over 26,000specieshave been described. Teleosts range fromgiant oarfishmeasuring 7.6 m (25 ft) or more, andocean sunfishweighing over 2 t (2.0 long tons; 2.2 short tons), to the minute maleanglerfishPhotocorynus spiniceps,just 6.2 mm (0.24 in) long. Including not only torpedo-shaped fish built for speed, teleosts can be flattened vertically or horizontally, be elongated cylinders or take specialised shapes as in anglerfish andseahorses.

The difference between teleosts and other bony fish lies mainly in their jaw bones; teleosts have a movablepremaxillaand corresponding modifications in the jaw musculature which make it possible for them toprotrude their jaws outwards from the mouth.This is of great advantage, enabling them tograb preyanddraw it into the mouth.In morederivedteleosts, the enlarged premaxilla is the main tooth-bearing bone, and the maxilla, which is attached to the lower jaw, acts as a lever, pushing and pulling the premaxilla as the mouth is opened and closed. Other bones further back in the mouth serve to grind and swallow food. Another difference is that the upper and lower lobes of thetail (caudal) finare about equal in size. Thespineends at thecaudal peduncle,distinguishing this group from other fish in which the spine extends into the upper lobe of the tail fin.

Teleosts have adopted a range ofreproductive strategies.Most use external fertilisation: the female lays a batch of eggs, the male fertilises them and thelarvaedevelop without any further parental involvement. A fair proportion of teleosts are sequentialhermaphrodites,starting life as females and transitioning to males at some stage, with a few species reversing this process. A small percentage of teleosts areviviparousand some provide parental care with typically the male fish guarding a nest and fanning the eggs to keep them well-oxygenated.

Teleosts are economically important to humans, as is shown by theirdepiction in artover the centuries. Thefishing industryharvests them for food, andanglersattempt to capture themfor sport.Some species arefarmedcommercially, and this method of production is likely to be increasingly important in the future. Others are kept inaquariumsor used in research, especially in the fields ofgeneticsanddevelopmental biology.

Anatomy

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Main articles:Fish anatomyandFish jaw
Teleost skull and jaw anatomy enables them both to suck in prey, and to close the mouth without expelling the prey again.[5]

Distinguishingfeatures of the teleosts are mobilepremaxilla,elongatedneural archesat the end of thecaudal finand unpairedbasibranchialtoothplates.[6]The premaxilla is unattached to theneurocranium(braincase); it plays a role in protruding the mouth and creating a circular opening. This lowers the pressure inside the mouth, sucking the prey inside. The lower jaw andmaxillaare then pulled back to close the mouth, and the fishis able to grasp the prey.By contrast, mere closure of the jaws would risk pushing food out of the mouth. In more advanced teleosts, the premaxilla is enlarged and has teeth, while the maxilla is toothless. The maxilla functions to push both the premaxilla and the lower jaw forward. To open the mouth, anadductor musclepulls back the top of the maxilla, pushing the lower jaw forward. In addition, the maxilla rotates slightly, which pushes forward a bony process that interlocks with the premaxilla.[5]

Caudal skeleton showing symmetrical (homocercal) tail

Thepharyngeal jawsof teleosts, a second set of jaws contained within the throat, are composed of fivebranchial arches,loops of bone which support thegills.The first three arches include a single basibranchial surrounded by two hypobranchials, ceratobranchials, epibranchials and pharyngobranchials. The median basibranchial is covered by a toothplate. The fourth arch is composed of pairs of ceratobranchials and epibranchials, and sometimes additionally, some pharyngobranchials and a basibranchial. The base of the lower pharyngeal jaws is formed by the fifth ceratobranchials while the second, third and fourth pharyngobranchials create the base of the upper. In the morebasalteleosts the pharyngeal jaws consist of well-separated thin parts that attach to the neurocranium,pectoral girdle,andhyoid bar.Their function is limited to merely transporting food, and they rely mostly on lower pharyngeal jaw activity. In more derived teleosts the jaws are more powerful, with left and right ceratobranchials fusing to become one lower jaw; the pharyngobranchials fuse to create a large upper jaw that articulates with the neurocranium. They have also developed a muscle that allows the pharyngeal jaws to have a role in grinding food in addition to transporting it.[7]

The caudal fin ishomocercal,meaning the upper and lower lobes are about equal in size. The spine ends at the caudal peduncle, the base of the caudal fin, distinguishing this group from those in which the spine extends into the upper lobe of the caudal fin, such as most fish from thePaleozoic(541 to 252 million years ago). The neural arches are elongated to form uroneurals which provide support for this upper lobe.[5]

Teleosts tend to be quicker and more flexible than more basal bony fishes. Their skeletal structure has evolved towards greater lightness. While teleost bones are wellcalcified,they are constructed from a scaffolding of struts, rather than the densecancellous bonesofholosteanfish. In addition, the lower jaw of the teleost is reduced to just three bones; thedentary,theangular boneand thearticular bone.[8]Thegenitalandurinary tractsend behind theanusin thegenital papilla;this is observed tosexteleosts.[9]

Evolution and phylogeny

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Main article:Evolution of fish

External relationships

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The teleosts were first recognised as a distinct group by the GermanichthyologistJohannes Peter Müllerin 1845.[10]The name is fromGreekteleios,"complete" +osteon,"bone".[11]Müller based this classification on certain soft tissue characteristics, which would prove to be problematic, as it did not take into account the distinguishing features of fossil teleosts. In 1966, Greenwood et al. provided a more solid classification.[10][12]The oldest fossils of teleosteomorphs (thestem groupfrom which teleosts later evolved) date back to theTriassicperiod(Prohalecites,Pholidophorus).[13][14]However, it has been suggested that teleosts probably first evolved already during thePaleozoicera.[15]During theMesozoicandCenozoiceras they diversified widely, and as a result, 96% of all living fish species are teleosts.[16]

Thecladogrambelow shows theevolutionary relationshipsof the teleosts to otherextantcladesof bony fish,[15]and to the four-limbed vertebrates (tetrapods) thatevolvedfrom a related group of bony fish during theDevonianperiod.[17][18]Approximatedivergence dates(in millions of years,mya) are from Near et al., 2012.[15]

Euteleostomi/
Osteichthyes

Internal relationships

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The phylogeny of the teleosts has been subject to long debate, without consensus on either theirphylogenyor the timing of the emergence of the major groups before the application of modernDNA-based cladistic analysis. Near et al. (2012) explored the phylogeny and divergence times of every major lineage, analysing the DNA sequences of 9 unlinked genes in 232 species. They obtained well-resolved phylogenies with strong support for the nodes (so, the pattern of branching shown is likely to be correct). They calibrated (set actual values for) branching times in this tree from 36 reliable measurements of absolute time from the fossil record.[15]The teleosts are divided into the major clades shown on the cladogram,[19]with dates, following Near et al.[15]More recent research divide the teleosts into two major groups: Eloposteoglossocephala (Elopomorpha + Osteoglossomorpha) and Clupeocephala (the rest of the teleosts).[20][21]

Teleostei
Eloposteoglossocephala
Clupeocephala
Otocephala
230 mya
Euteleostei
Lepidogalaxii

Lepidogalaxiiformes(salamanderfish)

240 mya
310 mya

The mostdiversegroup of teleost fish today are the Percomorpha, which include, among others, thetuna,seahorses,gobies,cichlids,flatfish,wrasse,perches,anglerfish,andpufferfish.[22]Teleosts, and percomorphs in particular, thrived during theCenozoicera.Fossil evidence shows that there was a major increase in size and abundance of teleosts immediately after themass extinction eventat theCretaceous-Paleogene boundaryca. 66mya.[23]

Evolution of ray-finned fishes,Actinopterygii,from theDevonianto the present as a spindle diagram. The width of the spindles are proportional to the number of families as a rough estimate of diversity. The diagram is based on Benton, M. J. (2005) Vertebrate Palaeontology, Blackwell, 3rd edition, Fig 7.13 on page 185.
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Aspidorhynchus acustirostris,an early teleost from theMiddle Jurassic

The first fossils assignable to this diverse group appear in theEarly Triassic,[24]after which teleosts accumulated novel body shapes predominantly gradually for the first 150 million years of their evolution[24](Early Triassicthroughearly Cretaceous).

The most basal of the living teleosts are theElopomorpha(eels and allies) and theOsteoglossomorpha(elephantfishes and allies). There are 800 species of elopomorphs. They have thin leaf-shaped larvae known asleptocephali,specialised for a marine environment. Among the elopomorphs, eels have elongated bodies with lost pelvic girdles and ribs and fused elements in the upper jaw. The 200 species of osteoglossomorphs are defined by a bony element in the tongue. This element has a basibranchial behind it, and both structures have large teeth which are paired with the teeth on the parasphenoid in the roof of the mouth. The cladeOtocephalaincludes theClupeiformes(herrings) andOstariophysi(carps, catfishes and allies). Clupeiformes consists of 350 living species of herring and herring-like fishes. This group is characterised by an unusual abdominalscuteand a different arrangement of the hypurals. In most species, the swim bladder extends to the braincase and plays a role in hearing. Ostariophysi, which includes most freshwater fishes, includes species that have developed some unique adaptations.[5]One is theWeberian apparatus,an arrangement of bones (Weberian ossicles) connecting the swim bladder to the inner ear. This enhances their hearing, as sound waves make the bladder vibrate, and the bones transport the vibrations to the inner ear. They also have achemical alarm system;when a fish is injured, the warning substance gets in the water, alarming nearby fish.[25]

The majority of teleost species belong to the cladeEuteleostei,which consists of 17,419 species classified in 2,935 genera and 346 families. Shared traits of the euteleosts include similarities in the embryonic development of the bony or cartilaginous structures located between the head and dorsal fin (supraneural bones), an outgrowth on the stegural bone (a bone located near the neural arches of the tail), and caudal median cartilages located between hypurals of the caudal base. The majority of euteleosts are in the cladeNeoteleostei.A derived trait of neoteleosts is a muscle that controls the pharyngeal jaws, giving them a role in grinding food. Within neoteleosts, members of theAcanthopterygiihave a spiny dorsal fin which is in front of the soft-rayed dorsal fin.[26]This fin helps provide thrust in locomotion[27]and may also play a role in defense. Acanthomorphs have developed spinyctenoid scales(as opposed to thecycloid scalesof other groups), tooth-bearing premaxilla and greater adaptations to high speed swimming.[5]

Theadipose fin,which is present in over 6,000 teleost species, is often thought to have evolved once in the lineage and to have been lost multiple times due to its limited function. A 2014 study challenges this idea and suggests that the adipose fin is an example ofconvergent evolution.InCharaciformes,the adipose fin develops from an outgrowth after the reduction of the larval fin fold, while inSalmoniformes,the fin appears to be a remnant of the fold.[28]

Diversity

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Further information:Diversity of fish
Predatoryteleost: the flesh-cutting teeth of a piranha (Serrasalmidae)

There are over 26,000 species of teleosts, in about 40ordersand 448families,[29]making up 96% of allextantspecies offish.[16]Approximately 12,000 of the total 26,000 species are found in freshwater habitats.[30]Teleosts are found in almost every aquatic environment and have developed specializations to feed in a variety of ways as carnivores, herbivores,filter feedersandparasites.[31]The longest teleost is thegiant oarfish,reported at 7.6 m (25 ft) and more,[32]but this is dwarfed by the extinctLeedsichthys,one individual of which has been estimated to have a length of 27.6 m (91 ft).[33]The heaviest teleost is believed to be theocean sunfish,with a specimen landed in 2003 having an estimated weight of 2.3 t (2.3 long tons; 2.5 short tons),[34]while the smallest fully mature adult is the male anglerfishPhotocorynus spinicepswhich can measure just 6.2 mm (0.24 in), though the female at 50 mm (2 in) is much larger.[32]Thestout infantfishis the smallest and lightest adult fish and is in fact the smallest vertebrate in the world; the females measures 8.4 mm (0.33 in) and the male just 7 mm (0.28 in).[35]

A raregiant oarfish(Regalecus glesne), 7-metre (23 ft) long, captured in 1996

Open water fish are usually streamlined liketorpedoesto minimize turbulence as they move through the water. Reef fish live in a complex, relatively confined underwater landscape and for them, manoeuvrability is more important than speed, and many of them have developed bodies which optimize their ability to dart and change direction. Many have laterally compressed bodies (flattened from side to side) allowing them to fit into fissures and swim through narrow gaps; some use theirpectoral finsfor locomotion and others undulate their dorsal and anal fins.[36]Some fish have grown dermal (skin) appendages forcamouflage;theprickly leather-jacketis almost invisible among the seaweed it resembles and thetasselled scorpionfishinvisibly lurks on the seabed ready toambush prey.Some like thefoureye butterflyfishhave eyespots to startle or deceive, while others such aslionfishhaveaposematic colorationto warn that they are toxic or havevenomousspines.[37]

Flatfish aredemersal fish(bottom-feeding fish) that show a greater degree of asymmetry than any other vertebrates. The larvae are at firstbilaterally symmetricalbut they undergometamorphosisduring the course of their development, with one eye migrating to the other side of the head, and they simultaneously start swimming on their side. This has the advantage that, when they lie on the seabed, both eyes are on top, giving them a broad field of view. The upper side is usuallyspeckled and mottledfor camouflage, while the underside is pale.[38]

Thewinter flounderis asymmetrical, with both eyes lying on the same side of the head.
Commensalfish: aremoraholds on to its host with a sucker-like organ (detail inset)

Some teleosts are parasites.Remorashave their front dorsal fins modified into large suckers with which they cling onto ahost animalsuch as awhale,sea turtle,sharkorray,but this is probably acommensalrather than parasitic arrangement because both remora and host benefit from the removal ofectoparasitesand loose flakes of skin.[39]More harmful are thecatfishthat enter the gill chambers of fish and feed on their blood and tissues.[40]Thesnubnosed eel,though usually ascavenger,sometimes bores into the flesh of a fish, and has been found inside the heart of ashortfin mako shark.[41]

The knifefishGymnarchus niloticusgenerates weak electric fieldsenabling it todetect and locate preyin turbid water.

Some species, such aselectric eels,can produce powerful electric currents, strong enough to stun prey. Other fish, such asknifefish,generate and sense weak electric fieldsto detect their prey; they swim with straight backs to avoid distorting their electric fields. These currents are produced by modified muscle or nerve cells.[25]

Distribution

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Teleosts are found worldwide and in most aquatic environments, including warm and cold seas, flowing and stillfreshwater,and even, in the case of thedesert pupfish,isolated and sometimes hot andsaline bodies of waterin deserts.[42][43]Teleost diversity becomes low at extremely high latitudes; atFranz Josef Land,up to82°N,ice cover and water temperatures below 0 °C (32 °F) for a large part of the year limit the number of species; 75 percent of the species found there are endemic to the Arctic.[44]

Fish in a hot desert: thedesert pupfish

Of the major groups of teleosts, the Elopomorpha, Clupeomorpha and Percomorpha (perches, tunas and many others) all have a worldwide distribution and aremainly marine;the Ostariophysi and Osteoglossomorpha are worldwide butmainly freshwater,the latter mainly in the tropics; the Atherinomorpha (guppies, etc.) have a worldwide distribution, both fresh and salt, but are surface-dwellers. In contrast, the Esociformes (pikes) are limited to freshwater in the Northern Hemisphere, while the Salmoniformes (salmon,trout) are found in both Northern and Southern temperate zones in freshwater, some speciesmigratingto and from the sea. The Paracanthopterygii (cods, etc.) are Northern Hemisphere fish, with both salt and freshwater species.[43]

Some teleosts are migratory; certain freshwater species move within river systems on an annual basis; other species are anadromous, spending their lives at sea and moving inland tospawn,salmon andstriped bassbeing examples. Others, exemplified by theeel,arecatadromous,doing the reverse.[45]The fresh waterEuropean eelmigrates across the Atlantic Ocean as an adult to breed in floating seaweed in theSargasso Sea.The adults spawn here and then die, but the developing young are swept by theGulf Streamtowards Europe. By the time they arrive, they are small fish and enter estuaries and ascend rivers, overcoming obstacles in their path to reach the streams and ponds where they spend their adult lives.[46]

Teleosts including thebrown troutand thescaly osmanare found in mountain lakes inKashmirat altitudes as high as 3,819 m (12,530 ft).[47]Teleosts are found at extreme depths in the oceans; thehadal snailfishhas been seen at a depth of 7,700 m (25,300 ft), and a related (unnamed) species has been seen at 8,145 m (26,720 ft).[48][49]

Physiology

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Further information:Fish physiology

Respiration

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Gills
Further information:Fish respirationandFish gill

The major means of respiration in teleosts, as in most other fish, is the transfer of gases over the surface of the gills as water is drawn in through the mouth and pumped out through the gills. Apart from theswim bladder,which contains a small amount of air, the body does not have oxygen reserves, and respiration needs to be continuous over the fish's life. Some teleosts exploit habitats where the oxygen availability is low, such as stagnant water or wet mud; they have developed accessory tissues and organs to support gas exchange in these habitats.[50]

Several genera of teleosts have independently developed air-breathing capabilities, and some have becomeamphibious.Somecombtooth blenniesemerge to feed on land, and freshwater eels are able to absorb oxygen through damp skin.Mudskipperscan remain out of water for considerable periods, exchanging gases through skin andmucous membranesin the mouth and pharynx.Swamp eelshave similar well-vascularised mouth-linings, and can remain out of water for days and go into a resting state (aestivation) in mud.[51]Theanabantoidshave developed an accessory breathing structure known as thelabyrinth organon the first gill arch and this is used for respiration in air, andairbreathing catfishhave a similar suprabranchial organ. Certain other catfish, such as theLoricariidae,are able to respire through air held in their digestive tracts.[52]

Sensory systems

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A stickleback stained to show thelateral lineelements (neuromasts)
Further information:Sensory systems in fish,Electroreception,andMagnetoreception

Teleosts possess highly developed sensory organs. Nearly all daylightfish have colour visionat least as good as a normal human's. Many fish also havechemoreceptorsresponsible for acute senses of taste and smell. Most fish have sensitive receptors that form thelateral line system,which detects gentle currents and vibrations, and senses the motion of nearby fish and prey.[53]Fish sense sounds in a variety of ways, using the lateral line, the swim bladder, and in some species the Weberian apparatus. Fish orient themselves using landmarks, and may usemental mapsbased on multiple landmarks or symbols. Experiments with mazes show that fish possess thespatial memoryneeded to make such a mental map.[54]

Osmoregulation

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Osmoticchallenge:American eelsspawn in theseabut spend most of their adult life infreshwater,returning only to spawn.

The skin of a teleost is largely impermeable to water, and the main interface between the fish's body and its surroundings is the gills. In freshwater, teleost fish gain water across their gills byosmosis,while in seawater they lose it. Similarly, saltsdiffuseoutwards across the gills in freshwater and inwards in salt water. TheEuropean flounderspends most of its life in the sea but often migrates into estuaries and rivers. In the sea in one hour, it can gain Na+ions equivalent to forty percent of its total freesodiumcontent, with 75 percent of this entering through the gills and the remainder through drinking. By contrast, in rivers there is an exchange of just two percent of the body Na+content per hour. As well as being able to selectively limit salt and water exchanged by diffusion, there is an active mechanism across the gills for the elimination of salt in sea water and its uptake in fresh water.[55]

Thermoregulation

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Fish arecold-blooded,and in general their body temperature is the same as that of their surroundings. They gain and lose heat through their skin, and regulate their circulation in response to changes in water temperature by increasing or reducing the blood flow to the gills. Metabolic heat generated in the muscles or gut is quickly dissipated through the gills, with blood being diverted away from the gills during exposure to cold.[56]Because of their relative inability to control their blood temperature, most teleosts can only survive in a small range of water temperatures.[57]

Teleost species that inhabit colder waters have a higher proportion of unsaturated fatty acids in brain cell membranes compared to fish from warmer waters, which allows them to maintain appropriatemembrane fluidityin the environments in which they live.[58]When cold acclimated, teleost fish show physiological changes in skeletal muscle that include increased mitochondrial and capillary density.[59]This reduces diffusion distances and aids in the production of aerobicATP,which helps to compensate for the drop inmetabolic rateassociated with colder temperatures.

Tunaand otherfast-swimmingocean-goingfish maintain their muscles at higher temperatures than their environment for efficient locomotion.[60]Tuna achieve muscle temperatures 11 °C (19 °F) or even higher above the surroundings by having acounterflow systemin which themetabolic heatproduced by the muscles and present in the venous blood, pre-warms the arterial blood before it reaches the muscles. Other adaptations of tuna for speed include a streamlined, spindle-shaped body, fins designed to reducedrag,[60]and muscles with a raisedmyoglobincontent, which gives these a reddish colour and makes for a more efficient use of oxygen.[61]Inpolar regionsand in thedeep ocean,where the temperature is a few degrees above freezing point, some large fish, such as theswordfish,marlinand tuna, have a heating mechanism which raises the temperature of the brain and eye, allowing them significantly better vision than their cold-blooded prey.[62]

Buoyancy

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A teleostswim bladder

The body of a teleost is denser than water, so fish must compensate for the difference, or they will sink. A defining feature ofActinopteri(Chondrostei, Holostei and teleosts) is theswim bladder.[63][64]Originally present in the last common ancestor of the teleosts, it has since been lost independently at least 30–32 times in at least 79 of 425 families of teleosts where the swim bladder is absent in one or more species. This absence is often the case in fast-swimming fishes such as the tuna andmackerel.[65]The swim bladder helps fish adjusting their buoyancy through manipulation of gases, which allows them to stay at the current water depth, or ascend or descend without having to waste energy in swimming. In the more primitive groups like someminnows,the swim bladder is open (physostomous) to theesophagus.In fish where the swim bladder is closed (physoclistous), the gas content is controlled through therete mirabilis,a network of blood vessels serving as a countercurrent gas exchanger between the swim bladder and the blood.[66]

Locomotion

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Flying fishcombine swimming movements with the ability toglidein air using their longpectoral fins.
Main article:Fish locomotion

A typical teleost fish has a streamlined body for rapid swimming, and locomotion is generally provided by a lateral undulation of the hindmost part of the trunk and the tail, propelling the fish through the water.[67]There are many exceptions to this method of locomotion, especially where speed is not the main objective; among rocks and oncoral reefs,slow swimming with great manoeuvrability may be a desirable attribute.[68]Eels locomote by wiggling their entire bodies. Living amongseagrassesandalgae,theseahorseadopts an upright posture and moves by fluttering its pectoral fins, and the closely relatedpipefishmoves by rippling its elongated dorsal fin.Gobies"hop" along the substrate, propping themselves up and propelling themselves with their pectoral fins.[69]Mudskippers move in much the same way on terrestrial ground.[70]In some species, a pelvic sucker allows them to climb, and theHawaiian freshwater gobyclimbs waterfalls while migrating.[69]Gurnardshave three pairs of free rays on theirpectoral finswhich have a sensory function but on which they can walk along the substrate.[71]Flying fishlaunch themselves into the air and canglideon their enlarged pectoral fins for hundreds of metres.[72]

Sound production

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The ability to produce sound for communication appears to haveevolved independentlyin several teleost lineages.[73]Sounds are produced either bystridulationor by vibrating the swim bladder. In theSciaenidae,the muscles that attach to the swim bladder cause it to oscillate rapidly, creating drumming sounds. Marine catfishes, sea horses andgruntsstridulate by rubbing together skeletal parts, teeth or spines. In these fish, the swim bladder may act as aresonator.Stridulation sounds are predominantly from 1000–4000Hz,though sounds modified by the swim bladder have frequencies lower than 1000 Hz.[74][75]

Reproduction and lifecycle

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Sockeye salmon spawns, which breed only once and then die soon afterwards
Further information:Fish reproduction

Most teleost species areoviparous,havingexternal fertilisationwith both eggs and sperm being released into the water for fertilisation.Internal fertilisationoccurs in 500 to 600 species of teleosts but is more typical forChondrichthyesand many tetrapods. This involves the male inseminating the female with anintromittent organ.[76]Fewer than one in a million of externally fertilised eggs survives to develop into a mature fish, but there is a much better chance of survival among the offspring of members of about a dozen families which areviviparous.In these, the eggs are fertilised internally and retained in the female during development. Some of these species, like thelive-bearing aquarium fishin the familyPoeciliidae,areovoviviparous;each egg has ayolk sacwhich nourishes the developing embryo, and when this is exhausted, the egg hatches and the larva is expelled into thewater column.Other species, like the splitfins in the familyGoodeidae,are fully viviparous, with the developing embryo nurtured from the maternal blood supply via a placenta-like structure that develops in theuterus.Oophagyis practised by a few species, such asNomorhamphus ebrardtii;the mother lays unfertilised eggs on which the developing larvae feed in the uterus, and intrauterinecannibalismhas been reported in somehalfbeaks.[77]

There are two major reproductive strategies of teleosts;semelparity and iteroparity.In the former, an individual breeds once after reaching maturity and then dies. This is because the physiological changes that come with reproduction eventually lead to death.[78]Salmon of the genusOncorhynchusare well known for this feature; they hatch in fresh water and then migrate to the sea for up to four years before travelling back to their place of birth where they spawn and die. Semelparity is also known to occur in some eels and smelts. The majority of teleost species have iteroparity, where mature individuals can breed multiple times during their lives.[79]

Sex identity and determination

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Clownfishareprotandrous hermaphrodites;when the female of a breeding pair dies, the male changes sex and a subordinate male takes his place as the breeding male.

88 percent of teleost species aregonochoristic,having individuals that remain either male or female throughout their adult lives. The sex of an individual can be determinedgeneticallyas in birds and mammals, or environmentally as in reptiles. In some teleosts, both genetics and the environment play a role in determining sex.[80]For species whose sex is determined by genetics, it can come in three forms. In monofactorial sex determination, a single-locus determines sex inheritance. Both theXY sex-determination systemandZW sex-determination systemexist in teleost species. Some species, such as thesouthern platyfish,have both systems and a male can be determined by XY or ZZ depending on the population.[81]

Multifactorial sex determination occurs in numerousNeotropicalspecies and involves both XY and ZW systems. Multifactorial systems involve rearrangements of sex chromosomes and autosomes. For example, thedarter characinehas a ZW multifactorial system where the female is determined by ZW1W2and the male by ZZ. Thewolf fishhas a XY multifactorial system where females are determined by X1X1X2X2and the male by X1X2Y.[82]Some teleosts, such aszebrafish,have a polyfactorial system, where there are several genes which play a role in determining sex.[83]Environment-dependent sex determination has been documented in at least 70 species of teleost.Temperatureis the main factor, but pH levels, growth rate, density and social environment may also play a role. For theAtlantic silverside,spawning in colder waters creates more females, while warmer waters create more males.[84]

Hermaphroditism

[edit]

Some teleost species arehermaphroditic,which can come in two forms: simultaneous and sequential. In the former, both spermatozoa and eggs are present in the gonads.Simultaneous hermaphroditismtypically occurs in species that live in the ocean depths, where potential mates are sparsely dispersed.[85][86]Self-fertilisation is rare and has only been recorded in two species,Kryptolebias marmoratusandKryptolebias hermaphroditus.[86]With sequential hermaphroditism, individuals may function as one sex early in their adult life and switch later in life. Species with this condition includeparrotfish,wrasses,sea basses,flatheads,sea breamsandlightfishes.[85]

Protandry is when an individual starts out male and becomes female while the reverse condition is known as protogyny, the latter being more common. Changing sex can occur in various contexts. In thebluestreak cleaner wrasse,where males have harems of up to ten females, if the male is removed the largest and most dominant female develops male-like behaviour and eventually testes. If she is removed, the next ranking female takes her place. In the speciesAnthias squamipinnis,where individuals gather into large groups and females greatly outnumber males, if a certain number of males are removed from a group, the same number of females change sex and replace them. Inclownfish,individuals live in groups and only the two largest in a group breed: the largest female and the largest male. If the female dies, the male switches sexes and the next largest male takes his place.[87]

In deep-seaanglerfish(sub-order Ceratioidei), the much smaller male becomes permanently attached to the female and degenerates into a sperm-producing attachment. The female and their attached male become a "semi-hermaphroditic unit".[88]

Mating tactics

[edit]
Maledesert gobycourting a female

There are several different mating systems among teleosts. Some species arepromiscuous,where both males and females breed with multiple partners and there are no obvious mate choices. This has been recorded inGuppies,Baltic herring,Nassau groupers,humbug damselfish,cichlids andcreole wrasses.Polygamy,where one sex has multiple partners can come in many forms.Polyandryconsists of one adult female breeding with multiple males, which only breed with that female. This is rare among teleosts, and fish in general, but is found in the clownfish. In addition, it may also exist to an extent among anglerfish, where some females have more than one male attached to them.Polygyny,where one male breeds with multiple females, is much more common. This is recorded insunfish,sculpins,darters,damselfishand cichlids where multiple females may visit a territorial male that guards and takes care of eggs and young. Polygyny may also involve a male guarding aharemof several females. This occurs in coral reef species, such as damselfishes, wrasses, parrotfishes,surgeonfishes,triggerfishesandtilefishes.[79]

Lek breeding,where males congregate to display to females, has been recorded in at least one speciesCyrtocara eucinostomus.Lek-like breeding systems have also been recorded in several other species. Inmonogamousspecies, males and females may form pair bonds and breed exclusively with their partners. This occurs in North American freshwater catfishes, manybutterflyfishes,sea horses and several other species.[79]Courtship in teleosts plays a role in species recognition, strengthening pair bonds, spawning site position and gamete release synchronisation. This includes colour changes, sound production and visual displays (fin erection, rapid swimming, breaching), which is often done by the male. Courtship may be done by a female to overcome a territorial male that would otherwise drive her away.[89]

Male (top) and female humphead parrotfish, showing sexual dimorphism

Sexual dimorphismexists in some species. Individuals of one sex, usually males developsecondary sexual characteristicsthat increase their chances ofreproductive success.Indolphinfish,males have larger and blunter heads than females. In several minnow species, males develop swollen heads and small bumps known asbreeding tuberclesduring the breeding season.[90]The malegreen humphead parrotfishhas a more well-developed forehead with an "ossified ridge"which plays a role in ritualised headbutting.[91]Dimorphism can also take the form of differences in coloration. Again, it is usually the males that are brightly coloured; inkillifishes,rainbowfishesand wrasses the colours are permanent while in species like minnows, sticklebacks, darters and sunfishes, the colour changes with seasons. Such coloration can be very conspicuous to predators, showing that the drive to reproduce can be stronger than that to avoid predation.[90]

Males that have been unable to court a female successfully may try to achieve reproductive success in other ways. In sunfish species, like thebluegill,larger, older males known as parental males, which have successfully courted a female, construct nests for the eggs they fertilise. Smaller satellite males mimic female behaviour and coloration to access a nest and fertilise the eggs. Other males, known as sneaker males, lurk nearby and then quickly dash to the nest, fertilising on the run. These males are smaller than satellite males. Sneaker males also exist inOncorhynchussalmon, where small males that were unable to establish a position near a female dash in while the large dominant male is spawning with the female.[92]

Spawning sites and parental care

[edit]
Three-spined sticklebackmales (red belly) build nests and compete to attract females to lay eggs in them. Males then defend and fan the eggs. Painting byAlexander Francis Lydon,1879
Further information:Parental care

Teleosts may spawn in the water column or, more commonly, on the substrate. Water column spawners are mostly limited to coral reefs; the fish will rush towards the surface and release their gametes. This appears to protect the eggs from some predators and allow them to disperse widely via currents. They receive noparental care.Water column spawners are more likely than substrate spawners to spawn in groups. Substrate spawning commonly occurs in nests, rock crevices or even burrows. Some eggs can stick to various surfaces like rocks, plants, wood or shells.[93]

"Pregnant"male seahorse

Of the oviparous teleosts, most (79 percent) do not provide parental care.[94]Male care is far more common than female care.[94][95]Male territoriality"preadapts"a species to evolve male parental care.[96][97]One unusual example of female parental care is indiscuses,which provide nutrients for their developing young in the form of mucus.[98]Some teleost species have their eggs or young attached to or carried in their bodies. Forsea catfishes,cardinalfishes,jawfishesand some others, the egg may be incubated or carried in the mouth, a practice known asmouthbrooding.In some African cichlids, the eggs may be fertilised there. In species like thebanded acara,young are brooded after they hatch and this may be done by both parents. The timing of the release of young varies between species; some mouthbrooders release new-hatched young while other may keep then until they are juveniles. In addition to mouthbrooding, some teleost have also developed structures to carry young. Malenurseryfishhave a bony hook on their foreheads to carry fertilised eggs; they remain on the hook until they hatch. For seahorses, the male has a brooding pouch where the female deposits the fertilised eggs and they remain there until they become free-swimming juveniles. Femalebanjo catfisheshave structures on their belly to which the eggs attach.[99]

In some parenting species, young from a previous spawning batch may stay with their parents and help care for the new young. This is known to occur in around 19 species of cichlids inLake Tanganyika.These helpers take part in cleaning and fanning eggs and larvae, cleaning the breeding hole and protecting the territory. They have reduced growth rate but gain protection from predators.Brood parasitismalso exists among teleosts; minnows may spawn in sunfish nests as well as nests of other minnow species. Thecuckoo catfishis known for laying eggs on the substrate as mouthbrooding cichclids collect theirs and the young catfish will eat the cichlid larvae.Filial cannibalismoccurs in some teleost families and may have evolved to combat starvation.[100]

Growth and development

[edit]
Newly hatched Atlantic salmon with yolk sac

Teleosts have four major life stages: the egg, the larva, the juvenile and the adult. Species may begin life in a pelagic environment or ademersalenvironment (near the seabed). Most marine teleosts have pelagic eggs, which are light, transparent and buoyant with thin envelopes. Pelagic eggs rely on the ocean currents to disperse and receive no parental care. When they hatch, the larvae areplanktonicand unable to swim. They have a yolk sac attached to them which provides nutrients. Most freshwater species produce demersal eggs which are thick, pigmented, relatively heavy and able to stick to substrates. Parental care is much more common among freshwater fish. Unlike their pelagic counterparts, demersal larvae are able to swim and feed as soon as they hatch.[85]Larval teleosts often look very different from adults, particularly in marine species. Some larvae were even considered different species from the adults. Larvae have high mortality rates, most die from starvation or predation within their first week. As they grow, survival rates increase and there is greater physiological tolerance and sensitivity, ecological and behavioural competence.[101]

At the juvenile stage, a teleost looks more like its adult form. At this stage, itsaxial skeleton,internal organs, scales, pigmentation and fins are fully developed. The transition from larvae to juvenile can be short and fairly simple, lasting minutes or hours as in some damselfish, while in other species, like salmon,squirrelfish,gobies and flatfishes, the transition is more complex and takes several weeks to complete.[102]At the adult stage, a teleost is able to produce viable gametes for reproduction. Like many fish, teleosts continue to grow throughout their lives. Longevity depends on the species with some gamefish likeEuropean perchandlargemouth bassliving up to 25 years.Rockfishappear to be the longest living teleosts with some species living over 100 years.[103]

Shoaling and schooling

[edit]
Schooling predatorybluefin trevallysizing up schoolinganchovies
Main article:Shoaling and schooling

Many teleosts formshoals,which serve multiple purposes in different species. Schooling is sometimes anantipredator adaptation,offering improved vigilance against predators. It is often more efficient to gather food by working as a group, and individual fish optimise their strategies by choosing to join or leave a shoal. When a predator has been noticed, prey fish respond defensively, resulting in collective shoal behaviours such as synchronised movements. Responses do not consist only of attempting to hide or flee; antipredator tactics include for example scattering and reassembling. Fish also aggregate in shoals to spawn.[104]

Relationship with humans

[edit]
Main article:Fish in culture

Economic importance

[edit]
Fish farmingin the sea offScotland
Main article:Fishing

Teleosts are economically important in different ways. They arecaptured for foodaround the world. A small number of species such asherring,cod,pollock,anchovy,tuna andmackerelprovide people with millions of tons of food per year, while many other species are fished in smaller amounts.[105]They provide a large proportion of thefish caught for sport.[106]Commercial and recreational fishing together provide millions of people with employment.[107]

A small number of productive species including carp, salmon,[108]tilapiaandcatfisharefarmed commercially,producing millions of tons of protein-rich food per year. The UN'sFood and Agriculture Organizationexpects production to increase sharply so that by 2030, perhaps sixty-two percent of food fish will be farmed.[109]

Fish are consumed fresh, or may be preserved by traditional methods, which include combinations of drying,smoking,andsalting,orfermentation.[110]Modern methods of preservation include freezing,freeze-drying,and heat processing (as incanning). Frozen fish products include breaded orbatteredfillets,fish fingersandfishcakes.Fish meal is used as a food supplement for farmed fish and for livestock. Fish oils are made either from fish liver, especially rich invitamins AandD,or from the bodies of oily fish such as sardine and herring, and used as food supplements and to treat vitamin deficiencies.[111]

Some smaller and more colourful species serve asaquariumspecimens andpets.Sea wolvesare used in the leather industry.Isinglassis made from thread fish and drum fish.[106]

Impact on stocks

[edit]
Capture of Atlantic Cod 1950–2005 (FAO)

Human activities have affected stocks of many species of teleost, throughoverfishing,[112]pollutionandglobal warming.Among many recorded instances, overfishing caused the complete collapse of theAtlantic codpopulation offNewfoundlandin 1992, leading to Canada's indefinite closure of the fishery.[113]Pollution, especially in rivers and along coasts, has harmed teleosts as sewage, pesticides and herbicides have entered the water. Many pollutants, such asheavy metals,organochlorines,andcarbamatesinterfere with teleost reproduction, often by disrupting theirendocrinesystems. In theroach,river pollution has caused the intersex condition, in which an individual's gonads contain both cells that can make male gametes (such asspermatogonia) and cells that can make female gametes (such asoogonia). Since endocrine disruption also affects humans, teleosts are used to indicate the presence of such chemicals in water. Water pollution caused local extinction of teleost populations in many northern European lakes in the second half of the twentieth century.[114]

The effects of climate change on teleosts could be powerful but are complex. For example, increased winter precipitation (rain and snow) could harm populations of freshwater fish in Norway, whereas warmer summers could increase growth of adult fish.[115]In the oceans, teleosts may be able to cope with warming, as it is simply an extension of natural variation in climate.[116]It is uncertain howocean acidification,caused by rising carbon dioxide levels, might affect teleosts.[117]

Other interactions

[edit]
Service to science:zebrafishbeing bred in a research institute

A few teleosts are dangerous. Some, like eeltail catfish (Plotosidae), scorpionfish (Scorpaenidae) or stonefish (Synanceiidae) have venomous spines that can seriously injure or kill humans. Some, like theelectric eeland theelectric catfish,cangive a severe electric shock.Others, such as thepiranhaandbarracuda,have a powerful bite and have sometimes attacked human bathers.[106]Reports indicate that some of thecatfishfamily can be large enough toprey on human bathers.

Medakaand zebrafish are used as research models for studies ingeneticsanddevelopmental biology.The zebrafish is the most commonly used laboratory vertebrate,[106]offering the advantages of genetic similarity to mammals, small size, simple environmental needs, transparent larvae permitting non-invasive imaging, plentiful offspring, rapid growth, and the ability to absorbmutagensadded to their water.[118]

In art

[edit]

Teleost fishes have been frequent subjects in art, reflecting their economic importance, for at least 14,000 years. They were commonly worked into patterns inAncient Egypt,acquiringmythological significanceinAncient GreeceandRome,and from there intoChristianityas areligious symbol;artists in China and Japan similarly use fish images symbolically. Teleosts became common inRenaissance art,withstill lifepaintings reaching a peak of popularity in theNetherlands in the 17th century.In the 20th century, different artists such asKlee,Magritte,MatisseandPicassoused representations of teleosts to express radically different themes, from attractive to violent.[119]The zoologist and artistErnst Haeckelpainted teleosts and other animals in his 1904Kunstformen der Natur.Haeckel had become convinced byGoetheandAlexander von Humboldtthat by making accurate depictions of unfamiliar natural forms, such as from the deep oceans, he could not only discover "the laws of their origin and evolution but also to press into the secret parts of their beauty by sketching and painting".[120]

Notes

[edit]
  1. ^The other three groups are theHolostei(bowfinsandgars), theChondrostei(sturgeonsandpaddlefish), and theCladistia(bichirsandreedfish).

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Bibliography

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