Cypriniformes

Cypriniformes is an order of ray-finned fish, including the carps, minnows, loaches and relatives. This order contains 11-12 families, over 400 genera, and more than 4,250 species, with new species being described every few months or so, and new genera being recognized frequently.[2][3] They are most diverse in southeastern Asia, and are entirely absent from Australia and South America.[4]

Their closest living relatives are the Characiformes (characins and allies), the Gymnotiformes (electric eel and American knifefishes) and the Siluriformes (catfishes).[5]

Cypriniformes
Temporal range: Paleocene–Recent
Karper 59326
A wild-type common carp (Cyprinus carpio, Cyprinidae: Cyprininae)
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
(unranked): Cypriniphysae
Order: Cypriniformes
Bleeker, 1859
Families[2]

Balitoridae
Barbuccidae[1]
Botiidae[1]
Catostomidae
Cobitidae
Cyprinidae
Ellopostomatidae[1]
Gastromyzontidae[1]
Gyrinocheilidae
Nemacheilidae
Psilorhynchidae
Serpenticobitidae[1]
Vaillantellidae[1]
and see text

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 back; most other fishes of Ostariophysi have a small fleshy adipose fin behind the dorsal fin. Further 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, instead of an upper pharyngeal jaw.[4]

Steinbeisser 001
A true loach: the spined loach, Cobitis taenia

The most notable family placed here is Cyprinidae (carps and minnows) which make up two-thirds of the order's diversity. This is one of the largest families of fish, and is widely distributed across Africa, Eurasia, and North America. Most species are strictly freshwater inhabitants, but a considerable number are found in brackish water, such as roach and bream. At least one species is found in saltwater, the Pacific redfin, Tribolodon brandtii.[6] 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 Cyprinidae.[7]

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.[8] 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 Cobitidae common across Eurasia and parts of North Africa. A mid-sized group like the suckers,[9] 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 it seems that the last do not belong to the lineage of "true" loaches but are related to the suckers.[10]

Systematics

Nemac fasci 080519 9380 ckoep
Nemacheilus chrysolaimos is a stone loach. Closely related to true loaches, like these they have barbels.
Chinese algae eater
The Chinese algae eater (Gyrinocheilus aymonieri), one of the sucking loaches which are distant from other "loaches"
Erimyzon sucetta
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.[11]

The families of Cypriniformes are traditionally divided into two superfamilies. Superfamily Cyprinioidea contains the carps and minnows (Cyprinidae) and also the mountain carps as the family Psilorhynchidae.[4] In 2012 Maurice Kottelat reviewed the superfamily Cobitioidea and under his revision it now consists of the following families: hillstream loaches (Balitoridae), Barbuccidae, Botiidae, suckers (Catostomidae), true loaches (Cobitidae), Ellopostomatidae, Gastromyzontidae, sucking loaches (Gyrinocheilidae), stone loaches (Nemacheilidae), Serpenticobitidae and long-finned loaches (Vaillantellidae).[1]

Catostomoidea is usually treated as a junior synonym of Cobitioidea. But it seems that 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,[4] they were apparently successful enough never to shift from the original ecological niche of the basal Ostariophysi. Yet, from the ecomorphologically conservative main lineage apparently at least two major radiations branched off. These diversified from the lowlands into torrential river habitats, acquiring similar habitus and adaptations in the process.[10]

The mountain carps are highly apomorphic Cyprinidae, perhaps close to true carps (Cyprininae), or maybe to the danionins. While some details about the phylogenetic structures of this massively diverse family are known – e.g. that Cultrinae and Leuciscinae are rather close relatives and stand apart from Cyprininae – there is no good consensus 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:[10]

Phylogeny

Phylogeny based on the work of the following works[12][13][14][15]

Cypriniformes
Cyprinoidei

Cyprinidae

Cobitoidei
Catostomoidea

Catostomidae

Gyrinocheiloidea

Gyrinocheilidae

Cobitoidea

Botiidae

Vaillantellidae

Cobitidae

Balitoridae

Ellopostomatidae

Nemacheilidae

Evolution

Cypriniformes include the most primitive of the Ostariophysi in the narrow sense (i.e. excluding Gonorynchiformes). This is evidenced not only by physiological details, but 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).[16] However, their divergence probably occurred only with the splitting-up of Pangaea in the Jurassic, maybe 160 million years ago. By 110 mya, the plate tectonics evidence indicates that the Laurasian Cypriniformes must have been distinct from their Gondwanan relatives.[17]

Cypriniformes is 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 out-competed South American basal cypriniforms in Africa, where more advanced cypriniforms survive and coexist with characiforms.[18]

The earliest cypriniform fossils are already assignable to the living family Catostomidae; from the Paleocene of Alberta, they are roughly 60 million years old. During the Eocene (55-35 mya), catostomids and cyprinids spread throughout Asia. In the Oligocene, around 30 mya, advanced cyprinids began to out-compete catostomids wherever they were sympatric, causing a decline of the suckers. Cyprinids reached North America and Europe by about the same time, and Africa in the early Miocene (some 23-20 mya). The cypriniforms spread to North America through the Bering land bridge, which formed and disappeared again several times during the many millions of years of cypriniform evolution.[18]

Relationship with humans

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 the goldfish and koi are perhaps the most celebrated. The other families are of less commercial importance. The Catostomidae have some importance in angling, and some "loaches" are bred for the international aquarium fish trade.

Accidentally or deliberately introduced populations of common carp (Cyprinus carpio) and grass carp (Ctenopharyngodon idella) are found on all continents except Antarctica. In some cases, these exotic species have a negative impact on the environment. Carp in particular stir up the riverbed reducing the clarity of the water, making it difficult for plants to grow.[19]

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 experiment. As, during early development, the zebrafish has a nearly transparent body, it is ideal for studying developmental biology. It is also used for the elucidation of biochemical signaling pathways.[20] They are also good pets, but can be shy in bright light and crowded tanks.

Threats and extinction

Thicktail Chub
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.

Epalzeorhynchos bicolor
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 been established again successfully from captive stock. 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.[21]

Globally extinct Cypriniformes species are:[21]

Notes

  1. ^ a b c d e f g Kottelat, M. (2012)
  2. ^ a b Froese, Rainer, and Daniel Pauly, eds. (2012). "Cypriniformes" in FishBase. December 2012 version.
  3. ^ Eschmeyer, W.N., Fong, J.D. (2015) Species by family/subfamily in the Catalog of Fishes, California Academy of Sciences (retrieved 2 July 2015)
  4. ^ a b c d Nelson (2006)
  5. ^ Saitoh et al. (2003), Briggs (2005)
  6. ^ Orlov & Sa-a {2007]
  7. ^ FishBase (2004d,f), He et al. (2008)
  8. ^ FishBase (2004a,e)
  9. ^ FishBase (2004b,c)
  10. ^ a b c He et al. (2008)
  11. ^ Helfman et al. (1997): pp.228-229
  12. ^ Vendula Šlechtová, Jörg Bohlen, Heok Hui Tan: 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. Bd. 44, Nr. 3, 2007, S. 1358–1365
  13. ^ Wei-Jen Chen, V. Lheknim, Richard L. Mayden: Molecular phylogeny of the Cobitoidea (Teleostei: Cypriniformes) revisited: position of enigmatic loach Ellopostoma resolved with six nuclear genes. Journal of Fish Biology. Bd. 75, Nr. 9, 2009 S. 2197-2208, doi:10.1111/j.1095-8649.2009.02398.x
  14. ^ Jörg Bohlen, Vendula Šlechtová: Phylogenetic position of the fish genus Ellopostoma (Teleostei: Cypriniformes) using molecular genetic data. Ichthyological Exploration of Freshwaters. Bd. 20, Nr. 2, 2009, S. 157-162 (PDF; 1,8 MB)
  15. ^ Mikko Haaramo. "Cobitoidei – loach-like cypriniforms". Mikko's Phylogeny Archive. Retrieved 2013-10-26.
  16. ^ Saitoh et al. (2003)
  17. ^ Briggs (2005), Nelson (2006)
  18. ^ a b Briggs (2005)
  19. ^ GSMFC (2005), FFWCC [2008]
  20. ^ "Biochemical Signaling Pathways". ZFIN.
  21. ^ a b IUCN (2007)

References

External links

Barbinae

Barbinae are a subfamily of fish included in the family Cyprinidae. The taxonomy for this group has not been entirely worked out as some genera historically considered within it are still considered incertae sedis with respect to being a member of the family, and may be included here, while others may be moved to other subfamilies.

Catostomidae

The Catostomidae are the suckers of the order Cypriniformes, with about 78 species in this family of freshwater fishes. The Catostomidae are primarily native to North America, but Catostomus catostomus is found in both North America and Russia, and Myxocyprinus asiaticus is from China. They are not usually fished recreationally; they are not highly prized in North America for their flesh, although they are a fairly popular target with spear fisherman, and in some areas, such as the Ozarks, they are a common food fish.

Cobitidae

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 (see below). The family includes about 260 described species. New species are being described regularly.

Cyprinidae

The Cyprinidae are the family of freshwater fishes, 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 known fish family and the largest vertebrate animal family in general, with about 3,000 living and extinct species but only 1,270 remain extant. They range from about 12 mm to 3meters Catlocarpio siamensis. This certain family of fish is one of the only that do not take care of their eggs. In about 370 genera. The family belongs to the ostariophysian order Cypriniformes, of whose genera and species the cyprinids make more than two-thirds. The family name is derived from the Ancient Greek kyprînos (κυπρῖνος, "carp").

Cyprininae

The Cyprininae are one of at least 11 subfamilies of cyprinid fish. It contains three genera in its strictest definition but many more are included depending on which authority is defining it, especially if the Labeobarbinae is not considered to be a valid grouping.

Garra

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 140 species of garras are native to Asia, but about one-fifth of the species are from Africa (East, Middle and West, but by far the highest species richness in Ethiopia).The genus was established by Francis Buchanan-Hamilton in 1822 as a subgenus of Cyprinus (which at that time was a "wastebin genus" for carp-like cyprinids); he did not designate a type species. But as no other garras except the newly discovered G. lamta were known to science in 1822, this was designated as the type species by Pieter Bleeker in 1863. The garras and their closest relatives are sometimes placed in a subfamily Garrinae, but this seems hardly warranted. More often, this group is included in the Labeoninae, or together with these in the Cyprininae. In the former case, the garras are members of the labeonine tribe Garrini, in the latter they are in the subtribe Garraina of tribe Labeonini. The genus Discogobio is a close relative.

Gastromyzon

Gastromyzon is a genus of gastromyzontid loaches native to Borneo.

Gastromyzontidae

The Gastromyzontidae are a family of loaches native to China and Southeast Asia, where typically found in streams and rivers with a fast current. The family includes about 137 species in eighteen genera. This family was resurrected by M. Kottelat in his review and revision of the loaches in 2012.

Hillstream loach

The hillstream loaches or river loaches are a family, the Balitoridae, of small fish from South, Southeast and East Asia. The family includes about 99 species. They are sometimes sold as "lizardfish" or (in Germany) "flossensaugers". Many of the species are popular for aquaria. They have a number of similarities with the Cobitidae, their sibling family of "loaches", such as multiple barbels around the mouth. They should not be confused with the loricariids, which look similar but are a family of catfish.

Most species are rheophilic, living in swift, clear and well-oxygenated streams. Several species of the family live in fast-flowing streams or torrents and have modified ventral fins used for clinging to rocks.The subfamily Nemacheilinae has recently been separated as a distinct family, Nemacheilidae (stone loaches) and several genera have been separated into the family Gastromyzontidae.

Ictiobus

Ictiobus, also known as buffalo fish or simply buffalo, is a genus of freshwater fish common in the United States, but also found in Canada, Mexico, and Guatemala. They are the largest North American suckers, reaching up to 1.23 m (4.0 ft) in length. They are sometimes mistaken for carp because of the flat face and large, silver scales running along the body, though they lack the whisker-like barbels common to carp. Buffalo fish live in most types of freshwater bodies where panfish are found, such as ponds, creeks, rivers, and lakes. Ictiobus fish were caught by the Lewis and Clark Expedition.

From a fishermen's point of view, the buffalo fish is not a popular game fish because it is difficult to catch. Yet, once on the line, it can put up a good fight. The preferred method of catch is by the use of gill nets. These nets are set by hand during the night, when they are most effective.

Labeoninae

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 (when considered a distinct subfamily) or labeoins (when included in subfamily Cyprininae).

They include the group sometimes separated as Garrinae, but these do not seem to be that distinct. In fact, the entire Labeoninae is merged into the Cyprininae by a number of authors; in any case, these two and the former "Barbinae" form a close-knit group whose internal phylogeny is far from resolved. If the subfamily is considered distinct, it is typically split in the tribes Labeonini (which are able to swim well in open water) and Garrini (which are mostly benthic), and sometimes in addition the Banganini (which are somewhat intermediate in habitus) If the labeo lineage is included in the Cyprininae, it becomes the tribe Labeonini, while its two (or three) subdivisions are the subtribes Labeoina, Garraina and perhaps Banganina.Notable genera are Crossocheilus, Epalzeorhynchos and Garra, which contain some of the popular aquarium fishes often called "algae eaters", e.g. the Siamese algae-eater (Crossocheilus siamensis). Labeo – the type genus of this subfamily – contains many sizeable species which are often used as food.

Anatomically, the labeonins are distinguished by the Weberian apparatus contacting the skull with the supraneural bones, and its basioccipital process being concave in cross-section. The first vertebra has a parapophysis that is elongated to forward and partially overlaps the basioccipital process. The fourth vertebra, meanwhile, has a short but stout transverse process that is prominently elongated bellywards; the os suspensorium is often hidden behind if viewed from the side. In the skull, the frontal and sphenotic bones have prominent foramina. In the anal fin, the first pterygiophore is elongated and has well-developed anterior and posterior flanges, with the former very large and concave at the distal end. Most labeonins have the skinny flap of the underside of the snout well-developed into a fleshy cap that at least partially hides the upper lip except when feeding, and a similar structure at the lower lip.

Moxostoma

Moxostoma is a genus of North American ray-finned fish in the Catostomidae family.

Nemacheilidae

The Nemacheilidae, or stone loaches, are a family of cypriniform fishes that inhabit stream environments, mostly in Eurasia, with one genus, Afronemacheilus found in Africa. The family includes about 630 species.

Nemacheilus

Nemacheilus is a genus of stone loaches native to Asia.

Schistura

Schistura is a genus of fish in the stone loach family Nemacheilidae native to the streams and rivers of the southern and eastern Asia. Some of these species are troglobitic.

Shiner (fish)

Shiner is a common name used in North America for any of several kinds of small, usually silvery fish, in particular a number of cyprinids, but also e.g. the shiner perch (Cymatogaster aggregata).

Cyprinid shiners are:

Eastern shiners, genus Notropis

Finescale shiners, genus Lythrurus

Flagfin shiners, genus Pteronotropis

Golden shiner, Notemigonus crysoleucas (a monotypic genus)

Highscale shiners, genus Luxilus

Redside shiners, genus Richardsonius

Satinfin shiners, genus Cyprinella

Suckermouth

In fishes, a suckermouth is a ventrally-oriented (inferior) mouth adapted for grazing on algae and small organisms that grow on submerged objects.

All Loricariidae possess a suckermouth as do the cypriniform algae eaters of the genus Gyrinocheilus and other genera. The 'False Siamensis' (Epalzeorhynchus sp. or Garra taeniata) also has this feature. Three genera in Mochokidae, Atopochilus, Chiloglanis, and Euchilichthys, also possess an oral sucker, formed by the lips and part of the barbels. The Loricariidae and members of the genus Gyrinocheilus are further adapted by having a special opening on the gill cover so that the fish can breathe without using its mouth.Additionally many other fish of the order Cypriniformes have developed a sucking ability to a lesser or greater extent:

the Hillstream loaches (family Balitoridae) have modified fins to attach themselves to a surface and a ventrally located mouth.

The sucker fish (family Catostomidae) also have a modified mouth.In general many benthic dwelling or feeding fish will have a suckermouth feature. Many of these fishes originate from fast-moving waters, where the sucker allows the fish to 'stick' itself down without too much difficulty.

Whilst all these fish have a limited sucking ability – they are able to swallow their food – it is not necessarily correct to assume that they can attach themselves to a submerged object by suction; though their behaviour may give that impression, the orientation of their fins and a flow of water can give sufficient downward force to temporarily attach themselves to an object.

Triplophysa

Triplophysa is a genus of fish in the family Nemacheilidae found mainly in and around the Qinghai-Tibet Plateau in China. Currently, the genus is a mixed assemblage of species. Some lineages have been identified and treated as subgenera (Hedinichthys, Indotriplophysa, Labiatophysa, Qinghaichthys and Tarimichthys), but as Wikipedia follows Fishbase for fish species these have been treated as subgenera in Wikipedia, although Kottelat and the Catalog of Fishes treat them as genera. FishBase, however, includes these in Triplophysa without specifying subgenera and treats the names given by Kottelat as synonyms.

Yunnanilus

Yunnanilus is a genus of small stone loaches that are endemic to southeastern China, especially Guangxi and Yunnan. They are found in rivers, streams and lakes; some species are restricted to caves.

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