Cypriniformes is an order of ray-finned fish which contains many sister families and genera of cypriniform fish split into several suborders, including the barbs, loaches, botias, carps, danionins, and minnows, amongst others. Cypriniformes is an "order-within-an-order", placed under the superorder Ostariophysi—which is also made up of cyprinid, ostariophysin fishes. The order contains 11–12 families (with some authorities having listed as many as 23), over 400 genera, and more than 4,250 named species; new species are regularly described, and new genera are recognized frequently. Cyprinids are most diverse in South and Southeast Asia and are entirely absent from Australia and South America. At 112 years old, the longest-lived cypriniform fish documented is the bigmouth buffalo.
Their closest living relatives are the Characiformes (characins, tetras and their kin), the Gymnotiformes (electric eel and American knifefishes), and the Siluriformes (catfishes).
Description
Like other orders of the Ostariophysi, fishes of Cypriniformes possess a Weberian apparatus. They differ from most of their relatives in having only a dorsal fin on their backs; most other fishes of Ostariophysi have a small, fleshy adipose fin behind the dorsal fin. Other differences are the Cypriniformes' unique kinethmoid, a small median bone in the snout, and the lack of teeth in the mouth. Instead, they have convergent structures called pharyngeal teeth in the throat. While other groups of fish, such as cichlids, also possess pharyngeal teeth, the cypriniformes' teeth grind against a chewing pad on the base of the skull, rather than an upper pharyngeal jaw. Brackish water and marine cyprinids are invariably anadromous, swimming upstream into rivers to spawn. Sometimes separated as family Psilorhynchidae, they seem to be specially adapted fishes of the Cyprinidae.
The Balitoridae and Gyrinocheilidae are families of mountain-stream fishes feeding on algae and small invertebrates. They are found only in tropical and subtropical Asia. While the former are a speciose group, the latter contain only a handful of species. The suckers (Catostomidae) are found in temperate North America and eastern Asia. These large fishes are similar to carps in appearance and ecology. Members of the Cobitidae are common across Eurasia and parts of North Africa. A midsized group like the suckers, 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.
Systematics
thumb|A true loach – the [[spined loach, Cobitis taenia]]
thumb|right|[[Nemacheilus chrysolaimos is a stone loach. Closely related to true loaches, like these, they have barbels.]]
thumb|right|The Chinese algae eater ([[Gyrinocheilus aymonieri) is one of the sucking loaches, which are distant from other "loaches".]]
thumb|right|[[Erimyzon sucetta, a small sucker]]
Historically, these included all the forms now placed in the superorder Ostariophysi except the catfish, which were placed in the order Siluriformes. By this definition, the Cypriniformes were paraphyletic, so recently, the orders Gonorhynchiformes, Characiformes (characins and allies), and Gymnotiformes (knifefishes and electric eels) have been separated out to form their own monophyletic orders.
The families of Cypriniformes are traditionally divided into two suborders. Superfamily Cyprinioidea contains the carps and minnows (Cyprinidae) and also the mountain carps as the family Psilorhynchidae.
Catostomoidea is usually treated as a junior 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,thumb|Fossil of [[Jianghanichthys, the earliest known cypriniform]]
- Family †Jianghanichthyidae <small>Liu, Chang, Wilson & Murray, 2015</small> (Paleocene to Eocene of China)
- Suborder Gyrinocheiloidei <small>Betancur-R, et al., 2017</small>
- Family Gyrinocheilidae <small>Gill, 1907</small> (algae eaters)
- Suborder Catostomoidei <small>Betancur-R, et al., 2017</small>
- Family Catostomidae <small>Agassiz, 1850</small> (suckers)
- Suborder Cobitoidei <small>Fitzinger, 1832</small>
- Family Botiidae <small>Berg</small>, 1940 (pointface loaches)
- Family Vaillantellidae <small>Nalbant & Bănărescu, 1977</small> (longfin loaches)
- Family Cobitidae <small>Swainson, 1838</small> (spined loaches)
- Family Barbuccidae <small>Kottelat, 2012</small> (scooter loaches)
- Family Gastromyzontidae <small>Fowler, 1905</small> (hillstream loaches)
- Family Serpenticobitidae <small>Kottelat, 2012</small> (snake loaches)
- Family Balitoridae <small>Swainson, 1839</small> (river loaches)
- Family Ellopostomatidae <small>Bohlen & Šlechtová, 2009</small> (square-head loaches)
- Family Nemacheilidae <small>Regan, 1911</small> (brook loaches)
- Suborder Cyprinoidei <small>Fitzinger, 1832</small>
- Family Paedocyprididae <small>Mayden & W.J. Chen, 2010</small> (tiny carps)
- Family Psilorhynchidae <small>Hora, 1926</small> (mountain carps)
- Family Cyprinidae <small>Rafinesque, 1815</small> (carps)
- Family Sundadanionidae <small>Mayden & Chen, 2010</small> (tiny danios)
- Family Danionidae <small>Bleeker, 1863</small> (danionids)
- Family Leptobarbidae <small>Bleeker, 1864</small> (cigar barbs)
- Family Xenocyprididae <small>Günther, 1868</small> (East Asian minnows or sharpbellies)
- Family Tincidae <small>D. S. Jordan, 1878</small> (tenches)
- Family Acheilognathidae <small>Bleeker, 1863</small> (bitterlings)
- Family Gobionidae <small>Bleeker, 1863</small> (freshwater gudgeons)
- Family Tanichthyidae <small>Mayden & Chen, 2010</small> (mountain minnows)
- Family Leuciscidae <small>Bonaparte, 1835</small> (minnows)
Phylogeny
Phylogeny based on the work of the following works
Evolution
Cypriniformes include the most primitive of the Ostariophysi in the narrow sense (i.e. excluding the Gonorynchiformes). 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 from Characiphysi (Characiformes and relatives) is thought to be about the Early Triassic, about 250 million years ago (mya). However, their divergence probably occurred only with the splitting-up of Pangaea in the Jurassic, maybe 160 million years ago (Mya). By 110 Mya, the plate tectonics evidence indicates that the Laurasian Cypriniformes must have been distinct from their Gondwanan relatives.
The Cypriniformes are thought to have originated in South-east Asia, where the most diversity of this group is found today. The alternative hypothesis is that they began in South America, similar to the other otophysans. 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.
Until 2025, no cypriniform fossil remains were known predating the Cenozoic. In 2025, multiple fossil remains of an indeterminate cypriniform, including jaws, tooth plates, and vertebrae, were identified from the early Maastrichtian-aged (71–69 Mya) Prince Creek Formation of Alaska. This occurrence suggests that in contrast to previous hypotheses, cypriniforms may have originated in high-latitude regions and radiated southwards during the Cenozoic. The next-oldest cypriniform fossils are already assignable to the living family Catostomidae; from the Paleocene-aged Paskapoo Formation of Alberta, they are roughly 60 million years old. During the Eocene (55–35 Mya), catostomids and cyprinids spread throughout Asia; the earliest members of the cyprinid subfamilies Barbinae and Danioninae are known from the Eocene Sangkarewang Formation of Indonesia, in addition to possibly Smilogastrinae and Labeoninae. The extinct family Jianghanichthyidae is known from the Eocene of China.
In science, one of the most famous members of the Cypriniformes is the zebrafish (Danio rerio). The zebrafish is one of the most important vertebrate model 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. They are also good pets, but can be shy in bright light and crowded tanks.
Threats and extinction
thumb|left|The [[thicktail chub (Gila crassicauda) is globally extinct since about 1960.]]
Habitat destruction, damming of upland rivers, pollution, and in some cases overfishing for food or the pet trade have driven some Cypriniformes to the brink of extinction or even beyond. In particular, Cyprinidae of southwestern North America have been severely affected; a considerable number went entirely extinct after settlement by Europeans. For example, in 1900 the thicktail chub (Gila crassicauda) was the most common freshwater fish found in California; 70 years later, not a single living individual existed.
thumb|right|Few if any [[red-tailed black sharks (Epalzeorhynchos bicolor) remain in the wild today.]]
The well-known red-tailed black shark (Epalzeorhynchos bicolor) from the Mae Klong River of The 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. The Yarqon bleak (Acanthobrama telavivensis) from the Yarqon River had to be rescued into captivity from imminent extinction; new populations have apparently<!-- IUCN says no in 2006, but paper was published shortly thereafter --> 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.
- Acanthobrama hulensis
- Gökçe balığı, Alburnus akili
- Barbus microbarbis
- Snake River sucker, Chasmistes muriei
- Chondrostoma scodrense
- Cyprinus yilongensis
- Mexican dace, Evarra bustamantei
- Plateau chub, Evarra eigenmanni
- Endorheic chub, Evarra tlahuacensis
- Thicktail chub, Gila crassicauda
- Pahranagat spinedace, Lepidomeda altivelis
- Harelip sucker, Moxostoma lacerum
- Durango shiner, Notropis aulidion
- Phantom shiner, Notropis orca
- Salado shiner, Notropis saladonis
- Clear Lake splittail, Pogonichthys ciscoides
- Las Vegas dace, Rhinichthys deaconi
- Stumptooth minnow, Stypodon signifer
- Telestes ukliva
Notes
References
- (2005): The biogeography of otophysan fishes (Ostariophysi: Otophysi): a new appraisal. J. Biogeogr. 32(2): 287–294. <small></small> (HTML abstract)
- (2004a): Family Balitoridae - River loaches. Version of 2004-NOV-22. Retrieved 2007-03-05.
- (2004b): Family Catostomidae - Suckers. Version of 2004-NOV-22. Retrieved 2007-03-05.
- (2004c): Family Cobitidae - Loaches. Version of 2004-NOV-22. Retrieved 2007-03-05.
- (2004d): Family Cyprinidae - Minnows or carps. Version of 2004-NOV-22. Retrieved 2007-03-05.
- (2004e): Family Gyrinocheilidae - Algae eaters. Version of 2004-NOV-22. Retrieved 2007-03-05.
- (2004f): Family Psilorhynchidae - Mountain carps. Version of 2004-NOV-22. Retrieved 2007-03-05.
- (2005): Order Summary for Cypriniformes. Version of 2005-FEB-15. Retrieved 2007-03-05.
- (2006): Florida's Exotic Freshwater Fishes. Retrieved 2007-03-05.
- (2005): Cyprinus carpio (Linnaeus, 1758). Version of 2005-08-03. Retrieved 2007-05-03.
- (2008): Phylogenetic position of the enigmatic genus Psilorhynchus (Ostariophysi: Cypriniformes): Evidence from the mitochondrial genome. Mol. Phylogenet. Evol. 47: 419–425. <small></small> (HTML abstract)
- (1997): The Diversity of Fishes. Blackwell Publishing. <small></small>
- (2007): [www.iucnredlist.org 2007 IUCN Red List of Threatened Species].
- Kottelat, M. (2012): Conspectus cobitidum: an inventory of the loaches of the world (Teleostei: Cypriniformes: Cobitoidei). The Raffles Bulletin of Zoology, Suppl. No. 26: 1-199.
- (2006): Fishes of the World. John Wiley & Sons, Inc. <small></small>
- [2007]: FishBase - Tribolodon brandtii. Retrieved 2007-03-05.
- (2003): Mitochondrial Genomics of Ostariophysan Fishes: Perspectives on Phylogeny and Biogeography. J. Mol. Evol. 56(4): 464–472. <small> </small> (HTML abstract)
- (2018): Phylogenetic classification of extant genera of fishes of the order Cypriniformes (Teleostei: Ostariophysi). Zootaxa 4476(1): 006–039. <small></small> (HTML abstract)
External links
- Cypriniformes Tree of Life
