Cypriniformes/sɪˈprɪnɪfɔːrmiːz/ is anorder ofray-finned fish, which includes manyfamilies andgenera ofcyprinid (carps and their kin) fish, such asbarbs,gobies,loaches,botias, andminnows (among others). Cypriniformes is an "order-within-an-order", placed under thesuperorderOstariophysi—which is also made up of cyprinid,ostariophysin fishes. The order contains 11–12 families (with some authorities having listed as many as 23),[2] over 400 genera, and more than 4,250 namedspecies; new species are regularly described, and new genera are recognized frequently.[3][4] Cyprinids are most diverse inSouth andSoutheast Asia and are entirely absent fromAustralia andSouth America.[5] At 112 years old, the longest-lived cypriniform fish documented is thebigmouth buffalo.[6]
Like other orders of theOstariophysi, fishes of Cypriniformes possess aWeberian apparatus. They differ from most of their relatives in having only adorsal fin on their backs; most other fishes of Ostariophysi have a small, fleshyadipose fin behind the dorsal fin. Other differences are the Cypriniformes' uniquekinethmoid, a small median bone in the snout, and the lack of teeth in the mouth. Instead, they haveconvergent structures calledpharyngeal teeth in the throat. While other groups of fish, such ascichlids, also possess pharyngeal teeth, the cypriniformes' teeth grind against a chewing pad on the base of the skull, rather than an upper pharyngeal jaw.[5]
The most notable family placed here is theCyprinidae (carps andminnows), which make up two-thirds of the order's diversity. This is one of the largest families of fish, and is widely distributed acrossAfrica,Eurasia, andNorth America. Most species are strictly freshwater inhabitants, but some are found inbrackish water, such asroach andbream. At least one species is found in saltwater, the Pacific redfin,Tribolodon brandtii.[8] Brackish water and marine cyprinids are invariablyanadromous,[citation needed] swimming upstream into rivers to spawn. Sometimes separated as familyPsilorhynchidae, they seem to be specially adapted fishes of the Cyprinidae.[9]
TheBalitoridae andGyrinocheilidae are families of mountain-stream fishes feeding onalgae and smallinvertebrates. They are found only intropical andsubtropicalAsia. While the former are a speciose group, the latter contain only a handful ofspecies.[10] The suckers (Catostomidae) are found intemperateNorth America and eastern Asia.[citation needed] These large fishes are similar to carps in appearance and ecology. Members of theCobitidae are common acrossEurasia and parts ofNorth Africa.[citation needed] A midsized group like the suckers,[11] they are rather similar to catfish in appearance and behaviour, feeding primarily off the substrate and equipped with barbels to help them locate food at night or in murky conditions. Fishes in the families Cobitidae, Balitoridae, Botiidae, and Gyrinocheilidae are called loaches, although the last do not seem to belong to the lineage of "true" loaches, but are related to the suckers.[12]
Nemacheilus chrysolaimos is a stone loach. Closely related to true loaches, like these, they have barbels.The Chinese algae eater (Gyrinocheilus aymonieri) is one of the sucking loaches, which are distant from other "loaches".Erimyzon sucetta, a small sucker
Catostomoidea is usually treated as ajunior synonym of the Cobitoidei, but it could be split off the Catostomidae and Gyrinocheilidae in a distinct superfamily; the Catostomoidea might be closer relatives of the carps and minnows than of the "true" loaches. While the Cyprinioidea seem more "primitive" than the loach-like forms,[5] they were apparently successful enough never to shift from the originalecological niche of thebasalOstariophysi. Yet, from theecomorphologically conservative main lineage apparently at least two majorradiations branched off. These diversified from the lowlands into torrential riverhabitats, acquiring similarhabitus andadaptationsin the process.[12]
The mountain carps are the highlyapomorphic Cyprinidae, perhaps close totrue carps (Cyprininae), or maybe to thedanionins. While some details about thephylogenetic structures of this massively diverse family are known – e.g. thatCultrinae andLeuciscinae are rather close relatives and stand apart from Cyprininae – no good consensus exists yet on how the main lineages are interrelated. A systematic list, from the most ancient to the most modern lineages, can thus be given as:[15]
Cypriniformes include the most primitive of the Ostariophysi in the narrow sense (i.e. excluding theGonorynchiformes). This is evidenced not only by physiological details, but also by their great distribution, which indicates they had the longest time to spread. The earliest that Cypriniformes might have diverged fromCharaciphysi (Characiformes and relatives) is thought to be about theEarly Triassic, about 250 million years ago (mya).[22] However, their divergence probably occurred only with the splitting-up ofPangaea in theJurassic, maybe 160 million years ago (Mya). By 110 Mya, theplate tectonics evidence indicates that theLaurasian Cypriniformes must have been distinct from theirGondwanan relatives.[23]
The Cypriniformes are thought to have originated inSouth-east Asia, where the most diversity of this group is found today. The alternative hypothesis is that they began inSouth America, similar to the otherotophysans. If this were the case, they would have spread to Asia through Africa or North America before the continents split up, for these are purely freshwater fishes. As the Characiformes began to diversify and spread, they may have outcompeted South American basal cypriniforms in Africa, where more advanced cypriniforms survive and coexist with characiforms.[24]
The earliest cypriniform fossils are already assignable to the livingfamily Catostomidae; from thePaleocene ofAlberta, they are roughly 60 million years old. During theEocene (55–35 Mya), catostomids and cyprinids spread throughout Asia; the earliest members of the cyprinid subfamiliesBarbinae andDanioninae are known from the EoceneSangkarewang Formation ofIndonesia, in addition to possiblySmilogastrinae andLabeoninae.[25] The extinct familyJianghanichthyidae is known from the Eocene of China.[16] In theOligocene, around 30 Mya, advanced cyprinids began to outcompete catostomids wherever they weresympatric, causing a decline of the suckers. Cyprinids reached North America and Europe about the same time, and Africa in the earlyMiocene (some 23–20 Mya). The cypriniforms spread to North America through theBering land bridge, which formed and disappeared again several times during the many millions of years of cypriniformevolution.[24]
The Cyprinidae in particular are important in a variety of ways. Many species are important food fish, particularly in Europe and Asia. Some are also important as aquarium fish, of which thegoldfish andkoi are perhaps the most celebrated. The other families are of less commercial importance. The Catostomidae have some importance inangling, and some "loaches" are bred for the internationalaquarium fish trade.
Accidentally or deliberately introduced populations ofcommon carp (Cyprinus carpio) andgrass carp (Ctenopharyngodon idella) are found on allcontinents exceptAntarctica. In some cases, theseexotic species have a negative impact on the environment. Carp in particular stir up the riverbed, reducing the clarity of the water, making plant growth difficult.[26]
In science, one of the most famous members of the Cypriniformes is thezebrafish (Danio rerio). The zebrafish is one of the most important vertebratemodel organisms in biological and biochemical sciences, being used in many kinds of experiments. During early development, the zebrafish has a nearly transparent body, so it is ideal for studying developmental biology. It is also used for the elucidation of biochemical signaling pathways.[27] They are also good pets, but can be shy in bright light and crowded tanks.
Habitat destruction,damming of upland rivers,pollution, and in some casesoverfishing for food or the pet trade have driven some Cypriniformes to the brink ofextinction or even beyond. In particular, Cyprinidae of southwesternNorth America have been severely affected; a considerable number went entirely extinct after settlement by Europeans. For example, in 1900 thethicktail chub (Gila crassicauda) was the most common freshwater fish found inCalifornia; 70 years later, not a single living individual existed.
The well-knownred-tailed black shark (Epalzeorhynchos bicolor) from theMae Klong River ofThe Bridge on the River Kwai fame possibly only survives in captivity. Ironically, while pollution and other forms of overuse by humans have driven it from its native home, it is bred for the aquarium fish trade by the thousands. TheYarqon bleak (Acanthobrama telavivensis) from theYarqon River had to be rescued into captivity from imminent extinction; new populations have apparently been established again successfully from captive stock. The Balitoridae and Cobitidae, meanwhile, contain a very large number of species about which essentially nothing is known except how they look and where they were first found.[28]
^Thomas J. Near; Christine E. Thacker. "Phylogenetic Classification of Living and Fossil Ray-Finned Fishes (Actinopterygii)".Bulletin of the Peabody Museum of Natural History.65 (1):3–302.doi:10.3374/014.065.0101.
^Slechtová, V.; Bohlen, J.; Tan, H. H. (2007). "Families of Cobitoidea (Teleostei; Cypriniformes) as revealed from nuclear genetic data and the position of the mysterious genera Barbucca, Psilorhynchus, Serpenticobitis and Vaillantella".Molecular Phylogenetics and Evolution.44 (3):1358–65.Bibcode:2007MolPE..44.1358S.doi:10.1016/j.ympev.2007.02.019.PMID17433724.
^Chen, W.-J.; Lheknim, V.; Mayden, R. L. (2009). "Molecular phylogeny of the Cobitoidea (Teleostei: Cypriniformes) revisited: Position of enigmatic loachEllopostomaresolved with six nuclear genes".Journal of Fish Biology.75 (9):2197–2208.Bibcode:2009JFBio..75.2197C.doi:10.1111/j.1095-8649.2009.02398.x.PMID20738682.
^Jörg Bohlen, Vendula Šlechtová:Phylogenetic position of the fish genusEllopostoma (Teleostei: Cypriniformes) using molecular genetic data. Ichthyological Exploration of Freshwaters. Bd. 20, Nr. 2, 2009, S. 157–162 (PDFArchived 24 September 2015 at theWayback Machine; 1,8 MB)
He, Shunping; Gub, Xun; Mayden, Richard L.; Chen, Wei-Jen; Conway, Kevin W. & Chen, Yiyu (2008): Phylogenetic position of the enigmatic genusPsilorhynchus (Ostariophysi: Cypriniformes): Evidence from the mitochondrial genome.Mol. Phylogenet. Evol.47: 419–425.doi:10.1016/j.ympev.2007.10.012 (HTML abstract)
Helfman, G.; Collette, B. & Facey, D. (1997):The Diversity of Fishes. Blackwell Publishing.ISBN0-86542-256-7
Saitoh, Kenji; Miya, Masaki; Inoue, Jun G.; Ishiguro, Naoya B. & Nishida, Mutsuminame (2003): Mitochondrial Genomics of Ostariophysan Fishes: Perspectives on Phylogeny and Biogeography.J. Mol. Evol.56(4): 464–472.doi:10.1007/s00239-002-2417-yPMID12664166 (HTML abstract)
Tan, Milton & Armbruster, Jonathan W. (2018): Phylogenetic classification of extant genera of fishes of the order Cypriniformes (Teleostei: Ostariophysi).Zootaxa4476(1): 006–039.doi:10.11646/zootaxa.4476.1.4 (HTML abstract)
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