Cyprinidae is afamily offreshwater fish commonly called thecarp orminnow family, including thecarps and their relatives thebarbs andbarbels, among others. Cyprinidae is the largest and most diverse fish family, and the largestvertebrate animal family overall, with about 1,780 species divided into 166 validgenera.[2] Cyprinids range from about 12 mm (0.5 in) in size[needs update] to the 3 m (9.8 ft)giant barb (Catlocarpio siamensis).[3] The family name is derived from the Greek wordkyprînos (κυπρῖνος 'carp').
Cyprinids are stomachless, oragastric, fish with toothless jaws. Even so, food can be effectively chewed by thegill rakers of the specialized last gill bow. Thesepharyngeal teeth allow the fish to make chewing motions against a chewing plate formed by abony process of theskull. The pharyngeal teeth are unique to each species and are used to identify species. Strong pharyngeal teeth allow fish such as thecommon carp andide to eat hard baits such assnails andbivalves.
Hearing is a well-developed sense in the cyprinids since they have theWeberian organ, three specialized vertebral processes that transfer motion of thegas bladder to 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 consideredphysostomes because thepneumatic duct is 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).[4][5]
All fish in this family areegg-layers and most do not guard their eggs; however, a few species build nests and/or guard the eggs.
Most cyprinids feed mainly oninvertebrates andvegetation, probably due to the lack of teeth and stomach; however, some species, like theasp, are predators that specialize in fish. Many species, such as theide and 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 frog in 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 ineutrophic lakes. Here, they contribute towards positive feedback as they are efficient at eating thezooplankton that would otherwise graze on the algae, reducing its abundance.
Cyprinids are highly important food fish; they arefished andfarmed acrossEurasia. Inland-locked countries in particular, cyprinids are often the major species of fish eaten because they make the largest part ofbiomass in 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 inexpensivefrozen fish 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.[8]
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.
Several cyprinids have been introduced to waters outside their natural ranges to provide food, sport, or biological control for somepest species. The common carp (Cyprinus carpio) and the grass carp (Ctenopharyngodon idella) are the most important of these, for example inFlorida.
Cyprinus carpio is a major pest species inAustralia impacting freshwater environments, amenity, and the agricultural economy, devastatingbiodiversity by 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.[11]
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,[12][13][14] in 2020 this plan was found to be unlikely to work due to the high fecundity of the fish.[15]
Numerous cyprinids have become popular and important within theaquarium and fishpond hobbies, most famously thegoldfish, which was bred in China from wild Carassius species (Carassius auratus). 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 amur carp was bred in Japan into the colorful ornamental variety known askoi — or more accuratelynishikigoi (錦鯉), askoi (鯉) simply means "common carp" inJapanese — from the 18th century until today.
One particular species of these small and undemanding danionines is thezebrafish (Danio rerio). It has become the standardmodel species for studying developmental genetics ofvertebrates, in particular fish.[17]
Habitat destruction and other causes have reduced the wild stocks of several cyprinids to dangerously low levels; some are already entirelyextinct. In particular, the cyprinids of the subfamilyLeuciscinae from southwestern North America have been severely affected bypollution and unsustainable water use in the early to mid-20th century. The majority of globally extinctcypriniform species in fact belong to the leuciscinid cyprinids from the southwestern United States and northern Mexico.
The massive diversity of cyprinids has so far made it difficult to resolve theirphylogeny in sufficient detail to make assignment tosubfamilies more than tentative in many cases. Some distinct lineages obviously exist – for example, theCultrinae and Leuciscinae, regardless of their exact delimitation, are rather close relatives and stand apart fromCyprininae – but the overallsystematics andtaxonomy of the Cyprinidae remain a subject of considerable debate. A large number ofgenera areincertae sedis, too equivocal in their traits and/or too little-studied to permit assignment to a particular subfamily with any certainty.[18][19][20]
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 havecoevolved for 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.[19]
The validity and circumscription of proposed subfamilies like theLabeoninae orSqualiobarbinae also 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.[19]
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 typicalbarbels and 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.[18][21]
Thetench,Tinca tinca, is of unclear affiliations and often placed in asubfamily or family of its own.
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. Acladistic analysis ofDNA sequence data of the S7ribosomal proteinintron1 supports the view that it is distinct enough to constitute amonotypic subfamily. 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 orogeny that vastly changed thetopography of that region in the latePaleogene, when their divergence presumably occurred.[20]
A DNA-based analysis of these fish places the Rasborinae as the basal lineage with the Cyprininae as a sister clade to the Leuciscinae.[22] The subfamiliesAcheilognathinae,Gobioninae, andLeuciscinae are monophyletic.
More recent studies split theleuciscids,danionids,xenocyprids, and many others out of the family. However, even with these splits, Cyprinidae still remains the largest fish family.[2]
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. The following is a phylogeny of Cyprinoidei,[25][24] with clade names from van der Laan 2017:[26]
^Riehl, R.; Baensch, H. (1996).Aquarium Atlas Volume 1. Voyageur Press. p. 410.
^Helfman, Gene S.; Collette, Bruce B.; Facey, Douglas E. (1997).The diversity of fishes. Malden, Mass: Blackwell Science. p. 228.ISBN978-0-86542-256-8.OCLC299475257.
^abDe Graaf, Martin; Megens, Hendrik-Jan; Samallo, Johannis; Sibbing, Ferdinand A. (2007). "Evolutionary origin of Lake Tana's (Ethiopia) smallBarbus species: Indications of rapid ecological divergence and speciation".Animal Biology.57:39–48.doi:10.1163/157075607780002069.
^Howes, G.I. (1991). "Systematics and biogeography: an overview". In Winfield, I.J.; Nelson, J.S. (eds.).Biology of Cyprinids. London: Chapman and Hall Ltd. pp. 1–33.
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