Cyprinidae

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Cyprinidae
Temporal range: Eocene - Holocene
Caprinus carpio Prague Vltava 1.jpg
The common carp, Cyprinus carpio
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Cypriniformes
Suborder: Cyprinoidei
Family: Cyprinidae
Rafinesque, 1815 [1]
Type genus
Cyprinus
Subfamilies

see text

Cyprinidae is a family of freshwater fish commonly called the carp or minnow family, including the carps, the true minnows, and their relatives the barbs and barbels, among others. Cyprinidae is the largest and most diverse fish family, and the largest vertebrate animal family overall, with about 3,000 species; only 1,270 of these remain extant, divided into about 200 valid genera. [2] [3] Cyprinids range from about 12 mm (0.5 in) in size to the 3 m (9.8 ft) giant barb (Catlocarpio siamensis). [4] By genus and species count, the family makes up more than two-thirds of the ostariophysian order Cypriniformes. [2] [3] [5] The family name is derived from the Greek word kyprînos ( κυπρῖνος 'carp').

Contents

Biology and ecology

Cyprinids are stomachless, or agastric, fish with toothless jaws. Even so, food can be effectively chewed by the gill rakers of the specialized last gill bow. These pharyngeal teeth allow the fish to make chewing motions against a chewing plate formed by a bony process of the skull. The pharyngeal teeth are unique to each species and are used to identify species. Strong pharyngeal teeth allow fish such as the common carp and ide to eat hard baits such as snails and bivalves.

Hearing is a well-developed sense in the cyprinids since they have the Weberian organ, three specialized vertebral processes that transfer motion of the gas 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 considered physostomes because the pneumatic 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. Giant Barb.jpg
Giant barbs (Catlocarpio siamensis) are the largest members of this family.

Cyprinids are native to North America, Africa, and Eurasia. The largest known cyprinid is the giant barb (Catlocarpio siamensis), which may grow up to 3 m (9.8 ft) in length and 300 kg (660 lb) in weight. [4] Other very large species that can surpass 2 m (6.6 ft) are the golden mahseer (Tor putitora) and mangar (Luciobarbus esocinus). [6] [7] The largest North American species is the Colorado pikeminnow (Ptychocheilus lucius), which can reach up to 1.8 m (5.9 ft) in length. [8] Conversely, many species are smaller than 5 cm (2 in). The smallest known fish is Paedocypris progenetica , reaching 10.3 mm (0.41 in) at the longest. [9]

All fish in this family are egg-layers and most do not guard their eggs; however, a few species build nests and/or guard the eggs. The bitterlings of subfamily Acheilognathinae are notable for depositing their eggs in bivalve molluscs, where the young develop until able to fend for themselves.

Cyprinids contain the first and only known example of androgenesis in a vertebrate, in the Squalius alburnoides allopolyploid complex. [10]

Most cyprinids feed mainly on invertebrates and vegetation, probably due to the lack of teeth and stomach; however, some species, like the asp, are predators that specialize in fish. Many species, such as the ide and the common rudd, prey on small fish when individuals become large enough. Even small species, such as the moderlieschen, are opportunistic predators that will eat larvae of the common frog in artificial circumstances.

Some cyprinids, such as the grass carp, are specialized herbivores; others, such as the common nase, eat algae and biofilms, while others, such as the black carp, specialize in snails, and some, such as the silver carp, are specialized filter 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 in eutrophic lakes. Here, they contribute towards positive feedback as they are efficient at eating the zooplankton that would otherwise graze on the algae, reducing its abundance.

Relationship with humans

Wild capture of cyprinids by species in million tonnes, 1950-2009, as reported by the FAO Cyprinids wild capture.png
Wild capture of cyprinids by species in million tonnes, 1950–2009, as reported by the FAO

Food

Cyprinids are highly important food fish; they are fished and farmed across Eurasia. In land-locked countries in particular, cyprinids are often the major species of fish eaten because they make the largest part of biomass 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 inexpensive frozen fish products made this less important now than it was in earlier times. Nonetheless, in certain places, they remain popular for food, as well as recreational fishing, for ornamental use, and have been deliberately stocked in ponds and lakes for centuries for this reason. [12]

Sport

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

As pest control

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

As a pest species

Carp in particular can stir up sediment, reducing the clarity of the water and making plant growth difficult. [13] [14]

In America and Australia, such as the Asian carp in the Mississippi Basin, they have become invasive species that compete with native fishes or disrupt the environment.

Cyprinus carpio is a major pest species in Australia impacting freshwater environments, amenity, and the agricultural economy, devastating biodiversity 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, the Murray-Darling Basin, they constitute 80–90 per cent of fish biomass. [15]

In 2016 the federal government announced A$15.2 million to fund the National Carp Control Plan to investigate using Cyprinid 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, [16] [17] [18] in 2020 this plan was found to be unlikely to work due to the high fecundity of the fish. [19]

Aquarium fish

Numerous cyprinids have become popular and important within the aquarium and fishpond hobbies, most famously the goldfish, which was bred in China from the Prussian carp (Carassius (auratus) gibelio). First imported into Europe around 1728, it was originally much-fancied by the Chinese nobility as early as 1150 AD and, after it arrived there in 1502, also in Japan. In addition to the goldfish, the common carp was bred in Japan into the colorful ornamental variety known as koi — or more accurately nishikigoi (錦鯉), as koi (鯉) simply means "common carp" in Japanese — from the 18th century until today.

Other popular aquarium cyprinids include danionins, rasborines and true barbs. [20] Larger species are bred by the thousands in outdoor ponds, particularly in Southeast Asia, and trade in these aquarium fishes is of considerable commercial importance. The small rasborines and danionines are perhaps only rivalled by characids (tetras) and poecilid livebearers in their popularity for community aquaria.[ citation needed ] Some of the most popular cyprinids among aquarists, other than goldfish and koi, include the cherry barb, Harlequin rasbora, pearl danios, rainbow sharks, tiger barbs, and the White Cloud Mountain minnow.

One particular species of these small and undemanding danionines is the zebrafish (Danio rerio). It has become the standard model species for studying developmental genetics of vertebrates, in particular fish. [21]

Threatened families

Habitat destruction and other causes have reduced the wild stocks of several cyprinids to dangerously low levels; some are already entirely extinct. In particular, the cyprinids of the subfamily Leuciscinae from southwestern North America have been hit hard by pollution and unsustainable water use in the early to mid-20th century; most globally extinct cypriniform species are in fact leuciscinid cyprinids from the southwestern United States and northern Mexico.

Systematics

The massive diversity of cyprinids has so far made it difficult to resolve their phylogeny in sufficient detail to make assignment to subfamilies more than tentative in many cases. Some distinct lineages obviously exist – for example, the Cultrinae and Leuciscinae, regardless of their exact delimitation, are rather close relatives and stand apart from Cyprininae  – but the overall systematics and taxonomy of the Cyprinidae remain a subject of considerable debate. A large number of genera are incertae sedis , too equivocal in their traits and/or too little-studied to permit assignment to a particular subfamily with any certainty. [22] [23] [24]

Part of the solution seems that the delicate rasborines are the core group, consisting of minor lineages that have not shifted far from their evolutionary niche, or have coevolved 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. [23]

The validity and circumscription of proposed subfamilies like the Labeoninae or Squaliobarbinae also remain doubtful, although the latter do appear to correspond to a distinct lineage. The sometimes-seen grouping of the large-headed carps (Hypophthalmichthyinae) with Xenocypris , though, seems quite in error. More likely, the latter are part of the Cultrinae. [23]

The entirely paraphyletic "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 in Barbus sensu stricto – indeed, they are as distant from the typical barbels and the typical carps (Cyprinus) as these are from Garra (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. [22] [25]

The tench (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. A cladistic analysis of DNA sequence data of the S7 ribosomal protein intron  1 supports the view that it is distinct enough to constitute a monotypic subfamily. It also suggests it may be closer to the small East Asian Aphyocypris , Hemigrammocypris , and Yaoshanicus . They would have diverged roughly at the same time from cyprinids of east-central Asia, perhaps as a result of the Alpide orogeny that vastly changed the topography of that region in the late Paleogene, when their divergence presumably occurred. [24]

A DNA-based analysis of these fish places the Rasborinae as the basal lineage with the Cyprininae as a sister clade to the Leuciscinae. [26] The subfamilies Acheilognathinae, Gobioninae, and Leuciscinae are monophyletic.

Subfamilies and genera

Rainbow shark, Epalzeorhynchos frenatum, a somewhat aggressive aquarium fish Fransenlipper.JPG
Rainbow shark, Epalzeorhynchos frenatum, a somewhat aggressive aquarium fish
Acheilognathus longipinnis: Acheilognathinae Acheilognathus longipinnis2.jpg
Acheilognathus longipinnis : Acheilognathinae
Blue danio, Danio kerri: Danioninae Danio kerri.jpg
Blue danio, Danio kerri: Danioninae
Pseudogobio esocinus , Gobioninae Pseudogobio esocinus(Hamamatsu,Shizuoka,Japan).jpg
Pseudogobio esocinus , Gobioninae
Silver carp, Hypophthalmichthys molitrix: Xenocyprinae, alternatively Hypophthalmichthyinae Hypophthalmichthys molitrix adult.jpg
Silver carp, Hypophthalmichthys molitrix: Xenocyprinae, alternatively Hypophthalmichthyinae
Rohu, Labeo rohita, of the disputed Labeoninae Labeo rohita.JPG
Rohu, Labeo rohita, of the disputed Labeoninae
The tench, Tinca tinca, is of unclear affiliations and often placed in a subfamily of its own. FemaleTench1.JPG
The tench, Tinca tinca, is of unclear affiliations and often placed in a subfamily of its own.

Eschmeyer's Catalog of Fishes sets out the subfamilies and genera within the family Cyprinidae as follows: [27] [1] [28]

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: [29]

Phylogeny

Phylogeny of living Cyprinoidei [29] [28] with clade names from van der Laan 2017. [30]




































See also

Related Research Articles

<span class="mw-page-title-main">Carp</span> Various species of cyprinid fishes

The term carp is a generic common name for numerous species of freshwater fish from the family Cyprinidae, a very large clade of ray-finned fish mostly native to Eurasia. While carp are prized quarries and are valued as both food and ornamental fish in many parts of the Old World, they are considered trash fish and invasive pests in many parts of Africa, Australia and most of the United States.

<span class="mw-page-title-main">Cypriniformes</span> Order of fishes

Cypriniformes is an order of ray-finned fish, which includes many families and genera of cyprinid fish, such as barbs, gobies, loaches, botias, and minnows. 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, 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, but are entirely absent from Australia and South America. At 112 years old, the longest-lived cypriniform fish documented is the bigmouth buffalo.

<span class="mw-page-title-main">Cobitidae</span> Family of fishes

Cobitidae, also known as the true loaches, is a family of Old World freshwater fish. They occur throughout Eurasia and in Morocco, and inhabit riverine ecosystems. Today, most "loaches" are placed in other families. The family includes about 260 described species. New species are being described regularly.

<i>Phoxinus</i> Genus of fishes

Phoxinus is a genus of freshwater fish in the family Leuciscidae of order Cypriniformes, and the only members of the subfamily Phoxininae, or Eurasian minnows. The type species is Phoxinus phoxinus. The other species in this genus are also commonly known as minnows. The name "minnow" was what early English fisherman used to describe "small and insignificant". The genus Phoxinus is found throughout Eurasia, and includes 21 known species. Previously, members of the North American genus Chrosomus were also believed to form part of this genus.

<i>Barbus</i> Genus of fishes

Barbus is a genus of ray-finned fish in the family Cyprinidae. The type species of Barbus is the common barbel, first described as Cyprinus barbus and now named Barbus barbus. Barbus is the namesake genus of the subfamily Barbinae, but given their relationships, that taxon is better included in the Cyprininae at least for the largest part.

<i>Puntius</i> Genus of fishes

Puntius is a genus of small freshwater fish in the family Cyprinidae native to South Asia and Mainland Southeast Asia, as well as Taiwan.

<i>Rhinichthys</i> Genus of fishes

Rhinichthys, known as the riffle daces, is a genus of freshwater fish in the carp family (Cyprinidae) of the order Cypriniformes. The type species is Rhinichthys atratulus, the blacknose dace. Rhinichthys species range throughout North America.

<span class="mw-page-title-main">Tench</span> Species of ray-finned fish

The tench or doctor fish is a fresh- and brackish-water fish of the order Cypriniformes found throughout Eurasia from Western Europe including the British Isles east into Asia as far as the Ob and Yenisei Rivers. It is also found in Lake Baikal. It normally inhabits slow-moving freshwater habitats, particularly lakes and lowland rivers.

<i>Microrasbora</i> Genus of fishes

Microrasbora is a genus of small fishes. The generic name means "small Rasbora", however these are more closely related to the danios than rasboras. They inhabit freshwater in Myanmar and Yunnan, China.

<span class="mw-page-title-main">Hitch (fish)</span> Genus of fishes

The hitch is a cyprinid fish endemic to central California, and was once very common. The common name may derive from a Pomoan word for this species. It is the only species in the monospecific genus Lavinia.

<i>Garra</i> Genus of fishes

Garra is a genus of fish in the family Cyprinidae. These fish are one example of the "log suckers", sucker-mouthed barbs and other cyprinids commonly kept in aquaria to keep down algae. The doctor fish of Anatolia and the Middle East belongs in this genus. The majority of the more than 160 species of garras are native to Asia, but about one-fifth of the species are from Africa.

<span class="mw-page-title-main">Labeoninae</span> Subfamily of fishes

Labeoninae is a doubtfully distinct subfamily of ray-finned fishes in the family Cyprinidae of order Cypriniformes. They inhabit fresh water and the largest species richness is in the region around southern China, but there are also species elsewhere in Asia, and some members of Garra and Labeo are from Africa. They are a generally very apomorphic group, perhaps the most "advanced" of the Cyprinidae. A common name for these fishes is labeonins or labeoins.

The redspot barb is a species of freshwater cyprinid fish found in East Africa. It is named for the large, orange-red spot found on each operculum.

<i>Hypselobarbus</i> Genus of fishes

Hypselobarbus is a genus of fish in the family Cyprinidae endemic to India.

<span class="mw-page-title-main">Leuciscinae</span> Subfamily of fishes

Leuciscinae is a subfamily of the freshwater fish family Cyprinidae, which contains the true minnows.

Osteochilus lini is a freshwater fish from Southeast Asia. It is found in the lower Mekong River basin, the Chao Phraya River basin, and some coastal drainages; it occurs in Laos, Vietnam, Cambodia, and Thailand. Its common name is dusky face carp.

<i>Osteochilus waandersii</i> Species of fish

Osteochilus waandersii is a cyprinid freshwater fish from Southeast Asia. It is found in Indochina as well as in Sumatra and Borneo. Its common name is Waanders's hard-lipped barb.

<span class="mw-page-title-main">Cyprininae</span> Subfamily of fishes

Cyprinae is a subfamily of largely freshwater ray-finned fishes, one of ten subfamilies belonging to the family Cyprinidae. This family comprises of the carps, minnows, barbs and related fishes.

<span class="mw-page-title-main">Squaliobarbinae</span> Subfamily of fishes

Squaliobarbinae is a small subfamily of the carp and minnow family, Cyprinidae, which consists of three monotypic genera which have their natural distributions in eastern Asia. Two species, the grass carp and the black carp, have been introduced to other parts of the world for weed control and aquaculture. They are large cyprinids characterised by an enlarged subtemporal fossa, the palate articulating with the supraethmoid, an enlarged intercalar bone in the cranial vault, and a divided levator posterior muscle.

<span class="mw-page-title-main">Xenocyprinae</span> Subfamily of fishes

Xenocyprinae, is a contentious subfamily of the family Cyprinidae, the carp and minnow family, originally from eastern Asia.

References

  1. 1 2 Richard van der Laan; William N. Eschmeyer & Ronald Fricke (2014). "Family-group names of recent fishes". Zootaxa. 3882 (2): 1–230. doi: 10.11646/zootaxa.3882.1.1 . PMID   25543675.
  2. 1 2 Froese, Rainer; Pauly, Daniel (eds.). "Family Cyprinidae". FishBase . July 2015 version.
  3. 1 2 Eschmeyer, William N.; Fricke, Ron & van der Laan, Richard (eds.). "Genera in the family Cyprinidae". Catalog of Fishes . California Academy of Sciences . Retrieved 2 July 2015.
  4. 1 2 Froese, Rainer; Pauly, Daniel (eds.). "Catlocarpio siamensis". FishBase . March 2015 version.
  5. ↑ Nelson, Joseph (2006). Fishes of the World. Chichester: John Wiley & Sons. ISBN   0-471-25031-7.
  6. ↑ Froese, Rainer; Pauly, Daniel (eds.). "Tor putitora". FishBase . March 2017 version.
  7. ↑ Froese, Rainer; Pauly, Daniel (eds.). "Luciobarbus esocinus". FishBase . March 2017 version.
  8. ↑ Froese, Rainer; Pauly, Daniel (eds.). "Ptychocheilus lucius". FishBase . March 2015 version.
  9. ↑ Froese, Rainer; Pauly, Daniel (eds.). "Paedocypris progenetica". FishBase . March 2015 version.
  10. ↑ Morgado-Santos, Miguel; Carona, Sara; Vicente, Luís; Collares-Pereira, Maria João (2017). "First empirical evidence of naturally occurring androgenesis in vertebrates". Royal Society Open Science. 4 (5): 170200. Bibcode:2017RSOS....470200M. doi: 10.1098/rsos.170200 . PMC   5451830 . PMID   28573029.
  11. ↑ Based on data sourced from the FishStat database
  12. ↑ MacMahon, Alexander Francis Magri (1946). Fishlore: British Freshwater Fishes. Pelican Books. Vol. 161. Penguin Books. pp. 149–152.
  13. ↑ Gulf States Marine Fisheries Commission (3 August 2005). "Cyprinus carpio (Linnaeus, 1758)". Archived from the original on 18 August 2007. Retrieved 3 May 2007.
  14. ↑ Florida Fish and Wildlife Conservation Commission (2006). "Exotic Freshwater Fishes". Archived from the original on 8 February 2007. Retrieved 5 March 2007.
  15. ↑ Peterson, Douglas; Pearson, James; Simpson, William (2022-05-30). "Effects of common carp on water quality and submerged vegetation: results from a short-term mesocosm experiment in an artificial wetland". Marine and Freshwater Research. 73 (7): 973–994. doi: 10.1071/mf22008 . ISSN   1323-1650. S2CID   249222934.
  16. ↑ Mankad, Aditi; Zhang, Airong; Carter, Lucy; Curnock, Matthew (March 2022). "A path analysis of carp biocontrol: effect of attitudes, norms, and emotion on acceptance". Biological Invasions. 24 (3): 709–723. Bibcode:2022BiInv..24..709M. doi:10.1007/s10530-021-02679-1. ISSN   1387-3547. S2CID   244518717.
  17. ↑ Kilvert, Nick; Thomas, Kerrin (1 May 2016). "Herpes virus to be used in fight against carp in Murray River, Christopher Pyne says". ABC News. Australian Broadcasting Corporation. Archived from the original on 5 May 2016.
  18. ↑ "Carp herpesvirus". Managing Water Ecosystems - CSIRO. 2018-04-26. Retrieved 2020-11-09.
  19. ↑ Mintram, Kate S.; Oosterhout, Cock; Lighten, Jackie (2020-10-14). Knutie, Sarah (ed.). "Genetic variation in resistance and high fecundity impede viral biocontrol of invasive fish". Journal of Applied Ecology. 58. Wiley: 148–157. doi: 10.1111/1365-2664.13762 . ISSN   0021-8901.
  20. ↑ Riehl, R.; Baensch, H. (1996). Aquarium Atlas Volume 1. Voyageur Press. p. 410.
  21. ↑ Helfman, Gene S.; Collette, Bruce B.; Facey, Douglas E. (1997). The diversity of fishes. Malden, Mass: Blackwell Science. p. 228. ISBN   978-0-86542-256-8. OCLC   299475257.
  22. 1 2 De Graaf, Martin; Megens, Hendrik-Jan; Samallo, Johannis; Sibbing, Ferdinand A. (2007). "Evolutionary origin of Lake Tana's (Ethiopia) small Barbus species: Indications of rapid ecological divergence and speciation". Animal Biology. 57: 39–48. doi:10.1163/157075607780002069.
  23. 1 2 3 He, Shunping; Mayden, Richard L.; Wang, Xuzheng; Wang, Wei; Tang, Kevin L.; Chen, Wei-Jen; Chen, Yiyu (2008). "Molecular phylogenetics of the family Cyprinidae (Actinopterygii: Cypriniformes) as evidenced by sequence variation in the first intron of S7 ribosomal protein-coding gene: Further evidence from a nuclear gene of the systematic chaos in the family" (PDF). Molecular Phylogenetics and Evolution. 46 (3): 818–29. Bibcode:2008MolPE..46..818H. doi:10.1016/j.ympev.2007.06.001. PMID   18203625. Archived (PDF) from the original on 2011-08-11.
  24. 1 2 He, Shunping; Gu, Xun; Mayden, Richard L.; Chen, Wei-Jen; Conway, Kevin W.; Chen, Yiyu (2008). "Phylogenetic position of the enigmatic genus Psilorhynchus (Ostariophysi: Cypriniformes): Evidence from the mitochondrial genome" (PDF). Molecular Phylogenetics and Evolution. 47 (1): 419–25. Bibcode:2008MolPE..47..419H. doi:10.1016/j.ympev.2007.10.012. PMID   18053751. Archived (PDF) from the original on 2020-07-29.
  25. ↑ 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.
  26. ↑ Tao, Wenjing; Mayden, Richard L.; He, Shunping (March 2013). "Remarkable phylogenetic resolution of the most complex clade of Cyprinidae (Teleostei: Cypriniformes): A proof of concept of homology assessment and partitioning sequence data integrated with mixed model Bayesian analyses". Molecular Phylogenetics and Evolution. 66 (3): 603–616. Bibcode:2013MolPE..66..603T. doi:10.1016/j.ympev.2012.09.024. ISSN   1055-7903. PMID   23044401. Archived from the original on 2020-06-08. Retrieved 2019-12-07.
  27. ↑ "Eschmeyer's Catalog of Fishes Classification". California Academy of Sciences . Retrieved 11 December 2024.
  28. 1 2 Lei Yang; et al. (2015). "Phylogeny and polyploidy: Resolving the classification of cyprinine fishes (Teleostei: Cypriniformes)". Molecular Phylogenetics and Evolution. 85: 97–116. doi:10.1016/j.ympev.2015.01.014. ISSN   1055-7903.
  29. 1 2 Betancur-Rodriguez, Ricardo; Edward O. Wiley; Gloria Arratia; Arturo Acero; Nicolas Bailly; Masaki Miya; Guillaume Lecointre; Guillermo Ortí (2017). "Phylogenetic classification of bony fishes". BMC Evolutionary Biology. 17 (162) (4 ed.): 162. Bibcode:2017BMCEE..17..162B. doi: 10.1186/s12862-017-0958-3 . PMC   5501477 . PMID   28683774.
  30. ↑ van der Laan, Richard (December 2017). Freshwater fish list (PDF) (23rd ed.). p. 997. ISSN   2468-9157.

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