Cyprinidae

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Cyprinidaeis afamilyoffreshwater fishcommonly called thecarporminnow family,including thecarps,the trueminnows,and their relatives thebarbsandbarbels,among others. Cyprinidae is the largest and most diverse fish family, and the largestvertebrate animalfamily overall, with about 3,000species;only 1,270 of these remain extant, divided into about 200 validgenera.[1][2]Cyprinidsrange from about 12 mm (0.5 in) in size to the 3 m (9.8 ft)giant barb(Catlocarpio siamensis).[3]By genus and species count, the family makes up more than two-thirds of theostariophysianorderCypriniformes.[1][2][4]The family name is derived from the Greek wordkyprînos(κυπρῖνος'carp').

Cyprinidae
Temporal range:Eocene-Holocene
Thecommon carp,Cyprinus carpio
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Cypriniformes
Superfamily: Cyprinoidea
Family: Cyprinidae
Rafinesque,1815
Type genus
Cyprinus
Subfamilies

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Biology and ecology

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Cyprinids are stomachless, oragastric,fish with toothless jaws. Even so, food can be effectively chewed by thegill rakersof the specialized last gill bow. Thesepharyngeal teethallow the fish to make chewing motions against a chewing plate formed by abony processof theskull.The pharyngeal teeth are unique to each species and are used by scientists to identify species. Strong pharyngeal teeth allow fish such as thecommon carpandideto eat hard baits such assnailsandbivalves.

Hearing is a well-developed sense in the cyprinids since they have theWeberian organ,three specialized vertebral processes that transfer motion of thegas bladderto the inner ear. The vertebral processes of the Weberian organ also permit a cyprinid to detect changes in motion of the gas bladder due to atmospheric conditions or depth changes. The cyprinids are consideredphysostomesbecause thepneumatic ductis retained in adult stages and the fish are able to gulp air to fill the gas bladder, or they can dispose of excess gas to the gut.

Giant barbs(Catlocarpio siamensis) are the largest members of this family.

Cyprinids are native toNorth America,Africa,andEurasia.The largest known cyprinid is thegiant barb(Catlocarpio siamensis), which may grow up to 3 m (9.8 ft) in length and 300 kg (660 lb) in weight.[3]Other very large species that can surpass 2 m (6.6 ft) are thegolden mahseer(Tor putitora) andmangar(Luciobarbus esocinus).[5][6]The largest North American species is theColorado pikeminnow(Ptychocheilus lucius), which can reach up to 1.8 m (5.9 ft) in length.[7]Conversely, many species are smaller than 5 cm (2 in). Thesmallest known fishisPaedocypris progenetica,reaching 10.3 mm (0.41 in) at the longest.[8]

All fish in this family areegg-layersand most do not guard their eggs; however, a few species build nests and/or guard the eggs. The bitterlings of subfamilyAcheilognathinaeare notable for depositing their eggs in bivalvemolluscs,where the young develop until able to fend for themselves.

Cyprinids contain the first and only known example ofandrogenesisin a vertebrate, in theSqualius alburnoidesallopolyploidcomplex.[9]

Most cyprinids feed mainly oninvertebratesandvegetation,probably due to the lack of teeth and stomach; however, some species, like theasp,are predators that specialize in fish. Many species, such as theideand thecommon rudd,prey on small fish when individuals become large enough. Even small species, such as themoderlieschen,are opportunistic predators that will eat larvae of thecommon frogin artificial circumstances.

Some cyprinids, such as thegrass carp,are specialized herbivores; others, such as thecommon nase,eat algae andbiofilms,while others, such as theblack carp,specialize in snails, and some, such as thesilver carp,are specializedfilter feeders.For this reason, cyprinids are often introduced as a management tool to control various factors in the aquatic environment, such as aquatic vegetation and diseases transmitted by snails.

Unlike most fish species, cyprinids generally increase in abundance ineutrophiclakes. Here, they contribute towards positive feedback as they are efficient at eating thezooplanktonthat would otherwise graze on the algae, reducing its abundance.

Relationship with humans

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Wild capture of cyprinids by species in million tonnes, 1950–2009, as reported by the FAO[10]

Food

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Cyprinids are highly important food fish; they arefishedandfarmedacrossEurasia.Inland-lockedcountries in particular, cyprinids are often the major species of fish eaten because they make the largest part ofbiomassin most water types except for fast-flowing rivers. In Eastern Europe, they are often prepared with traditional methods such as drying and salting. The prevalence of inexpensivefrozenfish products made this less important now than it was in earlier times. Nonetheless, in certain places, they remain popular for food, as well asrecreational fishing,for ornamental use, and have been deliberately stocked in ponds and lakes for centuries for this reason.[11]

Sport

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Cyprinids are popular for angling especially formatch fishing(due to their dominance in biomass and numbers) and fishing for common carp because of its size and strength.

As pest control

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Several cyprinids have been introduced to waters outside their natural ranges to provide food, sport, or biological control for somepestspecies. The common carp (Cyprinus carpio) and the grass carp (Ctenopharyngodon idella) are the most important of these, for example inFlorida.

As a pest species

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Carp in particular can stir upsediment,reducing the clarity of the water and making plant growth difficult.[12][13]

In America and Australia, such as theAsian carpin theMississippi Basin,they have becomeinvasive speciesthat compete with native fishes or disrupt the environment.

Cyprinus carpiois a major pest species inAustraliaimpacting freshwater environments, amenity, and the agricultural economy, devastatingbiodiversityby decimating native fish populations where they first became established as a major pest in the wild in the 1960s. In the major river system of eastern Australia, theMurray-Darling Basin,they constitute 80-90 per cent of fish biomass.[14]

In 2016 the federal government announced A$15.2 million to fund the National Carp Control Plan to investigate usingCyprinid herpesvirus 3(carp virus) as a biological control agent while minimising impacts on industry and environment should a carp virus release go ahead. Despite initial, favourable assessment,[15][16][17]in 2020 this plan was found to be unlikely to work due to the high fecundity of the fish.[18]

Aquarium fish

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Numerous cyprinids have become popular and important within theaquarium and fishpondhobbies, most famously thegoldfish,which was bred in China from thePrussian carp(Carassius (auratus) gibelio). First imported into Europe around 1728, it was originally much-fancied by the Chinese nobility as early as 1150AD and, after it arrived there in 1502, also inJapan.In addition to the goldfish, the common carp was bred in Japan into the colorful ornamental variety known askoi— or more accuratelynishikigoi(Cẩm lí),askoi()simply means "common carp" inJapanese— from the 18th century until today.

Other popular aquarium cyprinids includedanionins,rasborinesandtrue barbs.[19]Larger species are bred by the thousands in outdoor ponds, particularly inSoutheast Asia,and trade in these aquarium fishes is of considerable commercial importance. The small rasborines and danionines are perhaps only rivalled bycharacids(tetras) andpoecilid livebearersin their popularity forcommunity aquaria.[citation needed]Some of the most popular cyprinids amongaquarists,other than goldfish and koi, include thecherry barb,Harlequin rasbora,pearl danios,rainbow sharks,tiger barbs,and theWhite Cloud Mountain minnow.

One particular species of these small and undemanding danionines is thezebrafish(Danio rerio). It has become the standardmodel speciesfor studying developmental genetics ofvertebrates,in particular fish.[20]

Threatened families

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Habitat destructionand other causes have reduced the wild stocks of several cyprinids to dangerously low levels; some are already entirelyextinct.In particular, the cyprinids of the subfamilyLeuciscinaefrom southwestern North America have been hit hard bypollutionand unsustainable water use in the early to mid-20th century; most globally extinctcypriniformspecies are in fact leuciscinid cyprinids from the southwestern United States and northern Mexico.

Systematics

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The massive diversity of cyprinids has so far made it difficult to resolve theirphylogenyin sufficient detail to make assignment tosubfamiliesmore than tentative in many cases. Some distinct lineages obviously exist – for example, theCultrinaeand Leuciscinae, regardless of their exact delimitation, are rather close relatives and stand apart fromCyprininae– but the overallsystematicsandtaxonomyof the Cyprinidae remain a subject of considerable debate. A large number ofgeneraareincertae sedis,too equivocal in their traits and/or too little-studied to permit assignment to a particular subfamily with any certainty.[21][22][23]

Part of the solution seems that the delicate rasborines are the core group, consisting of minor lineages that have not shifted far from theirevolutionary niche,or havecoevolvedfor millions of years. These are among the most basal lineages of living cyprinids. Other "rasborines" are apparently distributed across the diverse lineages of the family.[22]

The validity and circumscription of proposed subfamilies like theLabeoninaeorSqualiobarbinaealso remain doubtful, although the latter do appear to correspond to a distinct lineage. The sometimes-seen grouping of thelarge-headed carps(Hypophthalmichthyinae) withXenocypris,though, seems quite in error. More likely, the latter are part of the Cultrinae.[22]

The entirelyparaphyletic"Barbinae" and the disputed Labeoninae might be better treated as part of the Cyprininae, forming a close-knit group whose internal relationships are still little known. The small African "barbs"do not belong inBarbussensu stricto– indeed, they are as distant from the typicalbarbelsand thetypical carps(Cyprinus) as these are fromGarra(which is placed in the Labeoninae by most who accept the latter as distinct) and thus might form another as yet unnamed subfamily. However, as noted above, how various minor lineages tie into this has not yet been resolved; therefore, such a radical move, though reasonable, is probably premature.[21][24]

Thetench(Tinca tinca), a significant food species farmed in western Eurasia in large numbers, is unusual. It is most often grouped with the Leuciscinae, but even when these were rather loosely circumscribed, it always stood apart. Acladisticanalysis ofDNA sequencedata of the S7ribosomal proteinintron1 supports the view that it is distinct enough to constitute amonotypicsubfamily. It also suggests it may be closer to the smallEast AsianAphyocypris,Hemigrammocypris,andYaoshanicus.They would have diverged roughly at the same time from cyprinids of east-central Asia, perhaps as a result of theAlpide orogenythat vastly changed thetopographyof that region in the latePaleogene,when their divergence presumably occurred.[23]

A DNA-based analysis of these fish places the Rasborinae as the basal lineage with the Cyprininae as a sister clade to the Leuciscinae.[25]The subfamiliesAcheilognathinae,Gobioninae,andLeuciscinaeare monophyletic.

Subfamilies and genera

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Rainbow shark,Epalzeorhynchos frenatum,a somewhat aggressive aquarium fish
Acheilognathus longipinnis:Acheilognathinae
Blue danio,Danio kerri:Danioninae
Pseudogobio esocinus,Gobioninae
Silver carp,Hypophthalmichthys molitrix:Xenocyprinae,alternativelyHypophthalmichthyinae
Rohu,Labeo rohita,of the disputedLabeoninae
Thetench,Tinca tinca,is of unclear affiliations and often placed in asubfamilyof its own.

The 5th Edition ofFishes of the Worldsets out the following subfamilies:[26]

Flame chubHemitremia flammea,one of the chubs in theLeuciscinae)
Ide,Leuciscus idus,one of theEurasiandaces
Sailfin shiner,Notropis hypselopterus,a small and colorfulshinerof theLeuciscinae
Rhynchocypris oxycephalus,aminnowrelated to some North American daces
Sarmarutilus rubilio,aEuropeanroach
Trigonostigma somphongsi,arasbora,a relative of theblue danioabove
Black carp,Mylopharyngodon piceus:Squaliobarbinae
Hemigrammocypris rasborella,of uncertain relationship:
Possibly related toAphyocypris.

With such a large and diverse family the taxonomy and phylogenies are always being worked on so alternative classifications are being created as new information is discovered, for example:[37]

Phylogeny

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Phylogeny of living Cyprinoidei[37][38]with clade names from van der Laan 2017.[39]

SubfamilyProbarbinae

SubfamilyLabeoninae

SubfamilyTorinae

SubfamilySmiliogastrinae

SubfamilyCyprininae[incl.Barbinae]

SubfamilyDanioninae

SubfamilyLeptobarbinae

Flame chubHemitremia flammea,one of the chubs in theLeuciscinae)
Ide,Leuciscus idus,one of theEurasiandaces
Sailfin shiner,Notropis hypselopterus,a small and colorfulshinerof theLeuciscinae
Rhynchocypris oxycephalus,aminnowrelated to some North American daces
Sarmarutilus rubilio,aEuropeanroach
Trigonostigma somphongsi,arasbora,a relative of theblue danioabove
Black carp,Mylopharyngodon piceus:Squaliobarbinae

SubfamilyXenocypridinae[incl.Cultrinae&Squaliobarbinae]

SubfamilyTincinae

SubfamilyAcheilognathinae(bitterlings)

SubfamilyGobioninae

SubfamilyTanichthyinae

SubfamilyLeuciscinae[incl.Alburninae]

Hemigrammocypris rasborella,of uncertain relationship:
Possibly related toAphyocypris.

See also

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References

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  2. ^abEschmeyer, William N.;Fricke, Ron & van der Laan, Richard (eds.)."Genera in the family Cyprinidae".Catalog of Fishes.California Academy of Sciences.Retrieved2 July2015.
  3. ^abFroese, Rainer;Pauly, Daniel (eds.)."Catlocarpio siamensis".FishBase.March 2015 version.
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  6. ^Froese, Rainer;Pauly, Daniel (eds.)."Luciobarbus esocinus".FishBase.March 2017 version.
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