Cyprinidae

The Cyprinidae are the family of freshwater fish, collectively called cyprinids, that includes the carps, the true minnows, and their relatives (for example, the barbs and barbels). Also commonly called the "carp family", or "minnow family", Cyprinidae is the largest and most diverse fish family and the largest vertebrate animal family in general, with about 3,000 species of which only 1,270 remain extant, divided into about 370 genera.[1][2] They range from about 12 mm to the 3-m Catlocarpio siamensis.[3] The family belongs to the ostariophysian order Cypriniformes, of whose genera and species the cyprinids make up more than two-thirds.[1][2][4] The family name is derived from the Ancient Greek kyprînos (κυπρῖνος, "carp").

Cyprinidae
Temporal range: Eocene - Holocene
The common carp, Cyprinus carpio
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Cypriniformes
Superfamily: Cyprinoidea
Family: Cyprinidae
Subfamilies

and see text

Biology and ecology

Cyprinids are stomachless 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 by scientists 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.

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.[5] 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.

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[10]

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, and have been deliberately stocked in ponds and lakes for centuries for this reason.[11]

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.

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. In some cases, such as the Asian carp in the Mississippi Basin, they have become invasive species that compete with native fishes or disrupt the environment. Carp in particular can stir up sediment, reducing the clarity of the water and making plant growth difficult.[12][13]

Numerous cyprinids have become important in 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 much fancied by Chinese nobility as early as 1150 AD and after it arrived there in 1502, also in Japan. In the latter country, from the 18th century onwards, the common carp was bred into the ornamental variety known as koi – or more accurately nishikigoi (錦鯉), as koi (鯉) simply means "common carp" in Japanese.

Other popular aquarium cyprinids include danionins, rasborines, and true barbs.[14] 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 and poecilid livebearers in their popularity for community aquaria.

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.[15]

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.[16][17][18]

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.[17]

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.[17]

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.[16][19][20]

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.[18]

A DNA-based analysis of these fish places the Rasborinae as the basal lineage with the Cyprininae as a sister clade to the Leuciscinae.[21] The subfamilies Acheilognathinae, Gobioninae, and Leuciscinae are monophyletic.

Subfamilies and genera
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, alternatively Hypophthalmichthyinae
Rohu, Labeo rohita, of the disputed Labeoninae
The tench, Tinca tinca, is of unclear affiliations and often placed in a subfamily of its own.

The 5th Edition of Fishes of the World sets out the following subfamilies:[22]

Flame chub Hemitremia flammea, one of the chubs in the Leuciscinae)
Ide, Leuciscus idus , one of the Eurasian daces
Sailfin shiner, Notropis hypselopterus, a small and colorful shiner of the Leuciscinae
Rhynchocypris oxycephalus, a minnow related to some North American daces
Sarmarutilus rubilio, a European roach

Subfamily Leuciscinae
Trigonostigma somphongsi, a rasbora, a relative of the blue danio above
Black carp, Mylopharyngodon piceus: Squaliobarbinae

Incertae sedis
Hemigrammocypris rasborella, of uncertain relationship:
Possibly related to Aphyocypris.

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, foe example:[34]

Phylogeny
Phylogeny of living Cyprinoidei[34][35] with clade names from van der Laan 2017.[36]
 
 
Psilorhynchidae

Psilorhynchus

 
Cyprinidae
 

Probarbinae

 
 
Labeoninae
 

Parapsilorhynchini

 
 

Labeonini

 
 

Garrini

 
 
 
 

Torinae

 
 
 

Smiliogastrinae

 
Cyprininae
 
 

Cyprinini

 
 

Rohteichthyini

 
 
 
 
 

Acrossocheilini

 
 
 

Spinibarbini

 
 

Schizothoracini

 
 
 
 
 

Schizopygopsini

 
 

Barbini

 
 
 
 
 
 
 
 
 
Leuciscidae
Danioninae
 

?Paedocypridini

 
 

?Sundadanionini

 
 

Rasborini

 
 

Danionini

 
 

Chedrini

 
 
 
Leptobarbinae

Leptobarbus

 
 
Xenocyprinae
 

Squaliobarbini

 
 

Opsariichthyini

 
 

Oxygastrini

 
 

Hypophthalmichthyini

 
 

Xenocypridini

 
 
 
Tincinae

Tinca

 
 
 
 

Acheilognathinae

 
Gobioninae
 

Hemibarbus-Squalidus clade

 
 

Sarcocheilichthyini

 
 

Gobionini

 
 
 
 
Tanichthyinae

Tanichthys

 
Leuciscinae
 

Phoxinini

 
 

Laviniini

 
 

Plagiopterini

 
 

Leuciscini

 
 

Pogonichthyini

 
 
 
 
 
 
 
 
 

Subfamily Probarbinae

Subfamily Labeoninae

Subfamily Torinae

Subfamily Smiliogastrinae

Subfamily Cyprininae [incl. Barbinae]

Subfamily Danioninae

Subfamily Leptobarbinae

Flame chub Hemitremia flammea, one of the chubs in the Leuciscinae)
Ide, Leuciscus idus , one of the Eurasian daces
Sailfin shiner, Notropis hypselopterus, a small and colorful shiner of the Leuciscinae
Rhynchocypris oxycephalus, a minnow related to some North American daces
Sarmarutilus rubilio, a European roach
Trigonostigma somphongsi, a rasbora, a relative of the blue danio above
Black carp, Mylopharyngodon piceus: Squaliobarbinae

Subfamily Xenocypridinae [incl. Cultrinae & Squaliobarbinae]

Subfamily Tincinae

Subfamily Acheilognathinae (bitterlings)

Subfamily Gobioninae

Subfamily Tanichthyinae

Subfamily Leuciscinae [incl. Alburninae]

Incertae sedis
Hemigrammocypris rasborella, of uncertain relationship:
Possibly related to Aphyocypris.
gollark: No, i mean why is heavpoot utilizing C++, despite it being wrong?
gollark: That is an oddly specific time constraint, and C++?
gollark: Well, it seems incomprehensible, so who knows.
gollark: If it compiles, it runs, thus use C with all warnings disabled.
gollark: > they were probably too busy patting themselves on the back that their 100% type-safe code with type level computations, lenses and optimized memoization fucking compiled correctly to remember to test and make sure the program actually works;yroc;.

See also

References
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  2. Eschmeyer, W.N.; Fong, J.D. (2015). "Species by family/subfamily". Catalog of Fishes. California Academy of Science. Retrieved 2 July 2015.
  3. Fishbase-Catlocarpio-siamensis
  4. Nelson, Joseph (2006). Fishes of the World. Chichester: John Wiley & Sons. ISBN 0-471-25031-7.
  5. Froese, Rainer and Pauly, Daniel, eds. (2015). "Catlocarpio siamensis" in FishBase. March 2015 version.
  6. Froese, Rainer and Pauly, Daniel, eds. (2017). "Tor putitora" in FishBase. March 2017 version.
  7. Froese, Rainer and Pauly, Daniel, eds. (2017). "Luciobarbus esocinus" in FishBase. March 2017 version.
  8. Froese, Rainer and Pauly, Daniel, eds. (2015). "Ptychocheilus lucius" in FishBase. March 2015 version.
  9. Froese, Rainer and Pauly, Daniel, eds. (2015). "Paedocypris progenetica" in FishBase. March 2015 version.
  10. Based on data sourced from the FishStat database
  11. MacMahon, Alexander Francis Magri (1946). Fishlore: British Freshwater Fishes. Pelican Books. 161. Penguin Books. pp. 149–152.
  12. Gulf States Marine Fisheries Commission (3 August 2005). "Cyprinus carpio (Linnaeus, 1758)". Archived from the original on 18 August 2007. Retrieved 3 May 2007.
  13. Florida Fish and Wildlife Conservation Commission (2006). "Exotic Freshwater Fishes". Archived from the original on 8 February 2007. Retrieved 5 March 2007.
  14. Riehl, R.; Baensch, H. (1996). Aquarium Atlas Volume 1. Voyageur Press. p. 410.
  15. 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.
  16. 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.
  17. 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. doi:10.1016/j.ympev.2007.06.001. PMID 18203625.
  18. 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. doi:10.1016/j.ympev.2007.10.012. PMID 18053751.
  19. 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.
  20. IUCN (2009). "2009 IUCN Red List of Threatened Species". Archived from the original on 27 June 2014. Retrieved 20 September 2009.
  21. 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. doi:10.1016/j.ympev.2012.09.024. ISSN 1055-7903. PMID 23044401.
  22. J. S. Nelson; T. C. Grande; M. V. H. Wilson (2016). Fishes of the World (5th ed.). Wiley. pp. 181–186. ISBN 978-1-118-34233-6.
  23. Pethiyagoda, R.; Meegaskumbura, M.; Maduwage, K. (2012). "A synopsis of the South Asian fishes referred to Puntius (Pisces: Cyprinidae)". Ichthyological Exploration of Freshwaters. 23 (1): 69–95. ISSN 0936-9902.
  24. Kottelat, M. (2013). "The Fishes of the Inland Waters of Southeast Asia: A Catalogue and Core Bibliography of the Fishes Known to Occur in Freshwaters, Mangroves and Estuaries" (PDF). The Raffles Bulletin of Zoology (Supplement No. 27): 1–663. Archived from the original (PDF) on 2 December 2013.
  25. Pethiyagoda, R (2013). "Haludaria, a replacement generic name for Dravidia (Teleostei: Cyprinidae)" (PDF). Zootaxa. 3646 (2): 199. doi:10.11646/zootaxa.3646.2.9. PMID 26213759.
  26. Chang, Meemann; et al. (2008). "Extraordinarily thick-boned fish linked to the aridification of the Qaidam Basin (northern Tibetan Plateau)". Proceedings of the National Academy of Sciences. 105 (36): 13246–13251. doi:10.1073/pnas.0805982105. PMC 2533176. PMID 18757732.
  27. Borkenhagen, K (2014). "A new genus and species of cyprinid fish (Actinopterygii, Cyprinidae) from the Arabian Peninsula, and its phylogenetic and zoogeographic affinities". Environmental Biology of Fishes. 97 (10): 1179–1195. doi:10.1007/s10641-014-0315-y.
  28. Britz, Ralf; Kottelat, Maurice; Tan, Heok (1 December 2011). "Fangfangia spinicleithralis, a new genus and species of miniature cyprinid fish from the peat swamp forests of Borneo (Teleostei: Cyprinidae)". Ichthyological Exploration of Freshwaters. 22 (4): 327–335.
  29. Zhao, H.-T.; Sullivan, J.P.; Zhang, Y.-G.; Peng, Z.-G. (2014). "Paraqianlabeo lineatus, a new genus and species of labeonine fishes (Teleostei: Cyprinidae) from South China". Zootaxa. 3841 (2): 257–270. doi:10.11646/zootaxa.3841.2.5.
  30. Zhang, E; Zhou, W (2012). "Sinigarra napoense, a new genus and species of labeonin fishes (Teleostei: Cyprinidae) from Guangxi Province, South China" (PDF). Zootaxa. 3586: 17–25. doi:10.11646/zootaxa.3586.1.4.
  31. Huang, Y.; Yang, J.; Chen, X. (2014). "Stenorynchoacrum xijiangensis, a new genus and a new species of Labeoninae fish from Guangxi, China (Teleostei: Cyprinidae)". Zootaxa. 3793 (3): 379–386. doi:10.11646/zootaxa.3793.3.6.
  32. Nguyen, V.H.; Nguyen, H.D.; Nguyen, T.D.P. (2016). "Vinalabeo, a new generic name for Vinalabeo tonkinensis (Cyprinidae, Teleostei)". Journal of Science of Hnue, Natural Sciences. 61 (9): 140–144. doi:10.18173/2354-1059.2016-0066.
  33. Bianco, P.G.; Ketmaier, V. (2014). "A revision of the Rutilus complex from Mediterranean Europe with description of a new genus, Sarmarutilus, and a new species, Rutilus stoumboudae (Teleostei: Cyprinidae)" (PDF). Zootaxa. 3841 (3): 379–402. doi:10.11646/zootaxa.3841.3.4.
  34. 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 (4 ed.). 17 (162): 162. doi:10.1186/s12862-017-0958-3. PMC 5501477. PMID 28683774.
  35. Lei Yang; Tetsuya Sado; M. Vincent Hirt; Emmanuel Pasco-Viel; M. Arunachalam; Junbing Li; Xuzhen Wang; Jörg Freyhof; Kenji Saitoh; Andrew M. Simons; Masaki Miya; Shunping He; Richard L. Mayden (2015). "Phylogeny and Polyploidy: Resolving the Classification of Cyprinine Fishes (Teleostei: Cypriniformes)". Molecular Phylogenetics and Evolution. 85 (February 2015): 97. doi:10.1016/j.ympev.2015.01.014. PMID 25698355.
  36. van der Laan, Richard (December 2017). Freshwater fish list (PDF) (23rd ed.). p. 997. ISSN 2468-9157.
  37. Zheng, Lan‐Ping; Chen, Xiao‐Yong; Yang, Jun‐Xing (2019). "Molecular phylogeny and systematic revision of Bangana sensu lato (Teleostei, Cyprinidae)". Journal of Zoological Systematics and Evolutionary Research. 57 (4): 884–891. doi:10.1111/jzs.12294.
  38. Zhao, H.-T.; Sullivan, J.P.; Zhang, Y.-G.; Peng, Z.-G. (2014). "Paraqianlabeo lineatus, a new genus and species of labeonine fishes (Teleostei: Cyprinidae) from South China". Zootaxa. 3841 (2): 257–270. doi:10.11646/zootaxa.3841.2.5.
  39. Zhang, E; Zhou, W (2012). "Sinigarra napoense, a new genus and species of labeonin fishes (Teleostei: Cyprinidae) from Guangxi Province, South China" (PDF). Zootaxa. 3586: 17–25. doi:10.11646/zootaxa.3586.1.4.
  40. Huang, Y.; Yang, J.; Chen, X. (2014). "Stenorynchoacrum xijiangensis, a new genus and a new species of Labeoninae fish from Guangxi, China (Teleostei: Cyprinidae)". Zootaxa. 3793 (3): 379–386. doi:10.11646/zootaxa.3793.3.6.
  41. Pethiyagoda, R (2013). "Haludaria, a replacement generic name for Dravidia (Teleostei: Cyprinidae)" (PDF). Zootaxa. 3646 (2): 199. doi:10.11646/zootaxa.3646.2.9. PMID 26213759.
  42. Yang J, He S, Freyhof J, Witte K, Liu H (2006). "The phylogenetic relationships of the Gobioninae (Teleostei: Cyprinidae) inferred from mitochondrial cytochrome b gene sequences". Hydrobiologia. 553: 255–66. doi:10.1007/s10750-005-1301-3.
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