Cypriniformes is an

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of sk ...

, which includes many

families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...

and

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

of cyprinid ( carps and their kin) fish, such as barbs,

loaches Loaches are ray-finned fishes of the suborder Cobitoidei. They are freshwater, benthic (bottom-dwelling) fish found in rivers and creeks throughout Eurasia and northern Africa. Loaches are among the most diverse groups of fish; the 1249 known sp ...

, botias, and

minnow Minnow is the common name for a number of species of small freshwater fish, belonging to several genus, genera of the family Cyprinidae and in particular the subfamily Leuciscinae. They are also known in Ireland as wikt:pinkeen, pinkeens. While ...

s (among others). Cypriniformes is an "order-within-an-order", placed under the

superorder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...

Ostariophysi Ostariophysi is the second-largest superorder of fish. Members of this superorder are called ostariophysians. This diverse group contains 10,758 species, about 28% of known fish species in the world and 68% of freshwater species, and are present ...

—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 A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

; new species are regularly described, and new genera are recognized frequently.Eschmeyer, W.N., Fong, J.D. (2015
Species by family/subfamily
in the

Catalog of Fishes Catalog of Fishes is a comprehensive on-line database and reference work on the scientific names of fish species and genera. It is global in its scope and is hosted by the California Academy of Sciences. It has been compiled and is continuously up ...

, California Academy of Sciences (retrieved 2 July 2015) Cyprinids are most diverse in

South South is one of the cardinal directions or compass points. The direction is the opposite of north and is perpendicular to both west and east. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Proto-Germanic ''*sunþa ...

and

Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...

and are entirely absent from

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

.Nelson (2006) At 112 years old, the longest-lived cypriniform fish documented is the

bigmouth buffalo The bigmouth buffalo (''Ictiobus cyprinellus'') is a fish native to North America that is in decline. It is the largest North American species in the Catostomidae or "sucker" family, and is one of the List of longest-living organisms, longest-liv ...

. Their closest living relatives are the

Characiformes Characiformes is an order of ray-finned fish, comprising the characins and their allies. Grouped in 18 recognized families, more than 2000 different species are described, including the well-known piranha and tetras.; Buckup P.A.: "Relationsh ...

(

characin Characiformes is an order of ray-finned fish, comprising the characins and their allies. Grouped in 18 recognized families, more than 2000 different species are described, including the well-known piranha and tetras.; Buckup P.A.: "Relationsh ...

s, tetras and their kin), the

Gymnotiformes The Gymnotiformes are an order of teleost bony fishes commonly known as Neotropical knifefish or South American knifefish. They have long bodies and swim using undulations of their elongated anal fin. Found almost exclusively in fresh water (the ...

(

electric eel The electric eels are a genus, ''Electrophorus'', of neotropical freshwater fish from South America in the family Gymnotidae, of which they are the only members of the subfamily Electrophorinae. They are known for their electric fish, ability ...

and

American knifefish The Gymnotiformes are an order of teleost bony fishes commonly known as Neotropical knifefish or South American knifefish. They have long bodies and swim using undulations of their elongated anal fin. Found almost exclusively in fresh water (the ...

es), and the

Siluriformes Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Catfish are named for their prominent barbels, which resemble a cat's whiskers, though not all catfish have prominent barbels or "whis ...

( catfishes).

Description

Like other orders of the

, fishes of Cypriniformes possess a

Weberian apparatus The Weberian apparatus is an anatomical structure that connects the swim bladder to the auditory system in fishes belonging to the superorder Ostariophysi. When it is fully developed in adult fish, the elements of the apparatus are sometimes coll ...

. They differ from most of their relatives in having only a

dorsal fin A dorsal fin is a fin on the back of most marine and freshwater vertebrates. Dorsal fins have evolved independently several times through convergent evolution adapting to marine environments, so the fins are not all homologous. They are found ...

on their backs; most other fishes of Ostariophysi have a small, fleshy

adipose fin Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the back bone and are supported only b ...

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 Pharyngeal teeth are teeth in the pharyngeal arch of the throat of cyprinids, suckers, and a number of other fish species otherwise lacking teeth.cichlid Cichlids () are a large, diverse, and widespread family of percomorph fish in the family Cichlidae, order Cichliformes. At least 1,760 species have been scientifically described, making it one of the largest vertebrate families, with on ...

s, also possess pharyngeal teeth, the cypriniformes' teeth grind against a chewing pad on the base of the skull, rather than an upper pharyngeal jaw. Steinbeisser 001

The most notable family placed here is the

Cyprinidae 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 t ...

(

carp The term carp (: carp) is a generic common name for numerous species of freshwater fish from the family (biology), family Cyprinidae, a very large clade of ray-finned fish mostly native to Eurasia. While carp are prized game fish, quarries and a ...

s and

s), 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 Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...

Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...

, and

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

. Most species are strictly freshwater inhabitants, but some are found in

brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...

water, such as roach and

bream Bream (, ) are species of freshwater fish belonging to a variety of genera including '' Abramis'' (e.g., ''A. brama'', the common bream), '' Ballerus'', '' Blicca'', '' Brama'', '' Chilotilapia'', '' Etelis'', '' Lepomis'', '' Gymnocranius'', ...

. At least one species is found in saltwater, the Pacific redfin, '' Tribolodon brandtii''. Brackish water and marine cyprinids are invariably

anadromous Fish migration is mass relocation by fish from one area or body of water to another. Many types of fish migrate on a regular basis, on time scales ranging from daily to annually or longer, and over distances ranging from a few metres to thousa ...

, swimming upstream into rivers to spawn. Sometimes separated as family Psilorhynchidae, they seem to be specially adapted fishes of the Cyprinidae. The

Balitoridae Balitoridae, the hillstream loaches or river loaches, is a family, of small fish from South, Southeast and East Asia. The family includes about 202 species. They are sometimes sold as "lizardfish" or (in Germany) "flossensaugers". Many of the spe ...

and

Gyrinocheilidae ''Gyrinocheilus'' is the single genus in the family Gyrinocheilidae, a family of small Southeast Asian cypriniform fishes that live in fast-flowing freshwater mountain streams. The species in this genus are commonly called algae eaters. They ho ...

are families of mountain-stream fishes feeding on

alga Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular microalgae, suc ...

e and small

invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...

s. They are found only in

tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...

and

subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...

Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...

. While the former are a speciose group, the latter contain only a handful of

. The suckers (

Catostomidae The Catostomidae are the suckers of the order (biology), order Cypriniformes, with about 78 species in this family (biology), family of freshwater fishes. The Catostomidae are almost exclusively native to North America. The only exceptions are ' ...

) are found in

temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...

and eastern Asia. These large fishes are similar to carps in appearance and ecology. Members of the

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 includ ...

are common across

and parts of

North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...

. 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.He ''et al.'' (2008)

Systematics

Historically, these included all the forms now placed in the

except the catfish, which were placed in the order

. By this definition, the Cypriniformes were

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

, so recently, the orders Gonorhynchiformes,

, (

s and allies), and

( knifefishes and

s) have been separated out to form their own

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

orders. The

of Cypriniformes are traditionally divided into two suborders. Superfamily Cyprinioidea contains the carps and minnows (

) and also the mountain carps as the family Psilorhynchidae. In 2012,

Maurice Kottelat Maurice Kottelat (born 16 July 1957 in Delémont, SwitzerlandC ...

reviewed the superfamily Cobitoidei and under his revision it now consists of the following families: hillstream loaches (

), Barbuccidae,

Botiidae Botiidae, the pointface loaches, is a family of cypriniform ray-finned fishes from South, Southeast, and East Asia. Until recently they were placed in the true loach family Cobitidae, until Maurice Kottelat revised the loaches and re-elevated t ...

, suckers (

), true loaches (

), Ellopostomatidae,

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 Genus (; : genera ) is a taxonomic ran ...

, sucking loaches (

), stone loaches (

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 Africa is the world's second-largest and second-most populous ...

Serpenticobitidae ''Serpenticobitis'', popularly known as serpent loaches, is a small genus of loaches found in the Mekong River Basin in Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern r ...

, and long-finned loaches ( Vaillantellidae).Kottelat, M. (2012) Catostomoidea is usually treated as a

junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...

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, they were apparently successful enough never to shift from the original

ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of Resource (biology), resources an ...

of the basal

. Yet, from the

ecomorphological Ecomorphology or ecological morphology is the study of the relationship between the ecological role of an individual and its morphological adaptations. The term "morphological" here is in the anatomical context. Both the morphology and ecology ex ...

ly conservative main lineage apparently at least two major

radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or a material medium. This includes: * ''electromagnetic radiation'' consisting of photons, such as radio waves, microwaves, infr ...

s branched off. These diversified from the lowlands into torrential river

habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...

s, acquiring similar habitus and

adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...

s in the process. The mountain carps are the highly

apomorph In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ha ...

ic Cyprinidae, perhaps close to true carps (Cyprininae), or maybe to the danionins. While some details about the

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

structures of this massively diverse family are known – e.g. that Cultrinae and

Leuciscinae Leuciscinae is a subfamily of freshwater Actinopterygii, ray-finned fishes belonging to the Family (biology), family Leuciscidae, which includes the fishes known as daces, chubs, shiners and minnows. The fishes in this subfamily are mainly found i ...

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: Amyzon hunanensis 01

* Family †

Jianghanichthyidae ''Jianghanichthys'' is an extinct genus of freshwater Cypriniformes, cypriniform fish from the Thanetian, Late Paleocene to Ypresian, Early Eocene of central and southern China, and the only member of the family Jianghanichthyidae. It is the olde ...

Liu, Chang, Wilson & Murray, 2015 (

Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...

Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...

of China) * Suborder Gyrinocheiloidei Betancur-R, ''et al.'', 2017 ** Family

Gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...

, 1907 (algae eaters) * Suborder Catostomoidei Betancur-R, et al., 2017 ** Family

Agassiz Jean Louis Rodolphe Agassiz ( ; ) FRS (For) FRSE (May 28, 1807 – December 14, 1873) was a Swiss-born American biologist and geologist who is recognized as a scholar of Earth's natural history. Spending his early life in Switzerland, he recei ...

, 1850 (suckers) * Suborder Cobitoidei Fitzinger, 1832 ** Family

Berg Berg may refer to: People *Berg (surname), a surname (including a list of people with the name) * General Berg (disambiguation) * Berg Ng (born 1960), Hong Kong actor * Berg (footballer, born 1963), Ninimbergue dos Santos Guerra, Brazilian footba ...

, 1940 (pointface loaches) ** Family Vaillantellidae Nalbant & Bănărescu, 1977 (longfin loaches) ** Family

Swainson, 1838 (spined loaches) ** Family Barbuccidae Kottelat, 2012 (scooter loaches) ** Family

Fowler, 1905 (hillstream loaches) ** Family

Kottelat, 2012 (snake loaches) ** Family

Swainson, 1839 (river loaches) ** Family Ellopostomatidae Bohlen & Šlechtová, 2009 (square-head loaches) ** Family

Regan, 1911 (brook loaches) * Suborder

Cyprinoidei Cyprinoidei is a suborder, or superfamily, of mostly freshwater Actinopterygii, ray-finned fishes within the large order (biology), order Cypriniformes. Cyprinoidei was formerly a monotypic grouping only containing a single large family, the Cyprin ...

Fitzinger, 1832 ** Family Paedocyprididae Mayden & W.J. Chen, 2010 (tiny carps) ** Family Psilorhynchidae Hora, 1926 (mountain carps) ** Family

Rafinesque Constantine Samuel Rafinesque-Schmaltz (; 22 October 178318 September 1840) was a French early 19th-century polymath born near Constantinople in the Ottoman Empire and self-educated in France. He traveled as a young man in the United States, ult ...

, 1815 (carps) ** Family

Sundadanionidae Sundadanionidae, the tiny danios, is a family of freshwater ray-finned fishes belongong to the suborder Cyprinoidei of the order Cypriniformes. The fishes in this family are endemic Endemism is the state of a species being found only in a si ...

Mayden & Chen, 2010 (tiny danios) ** Family

Danionidae The danionins are a group of small, minnow-type fish belonging to the family Danionidae. Species of this group are in the genera clades ''Danio'' and '' Devario'' (which also includes ''Chela'', ''Laubuka'', ''Microdevario'', and ''Microrasbora ...

Bleeker, 1863 (danionids) ** Family Leptobarbidae Bleeker, 1864 (cigar barbs) ** Family

Xenocyprididae Xenocyprididae, is a family of freshwater ray-finned fishes commonly called the East Asian minnows or sharpbellies with a natural distribution in Asia. This taxon, sometimes spelt Xenocypridae, was previously regarded to be a subfamily, Xenocypri ...

Günther, 1868 (East Asian minnows or sharpbellies) ** Family Tincidae D. S. Jordan, 1878 (tenches) ** Family Acheilognathidae Bleeker, 1863 (bitterlings) ** Family

Gobionidae Gobioninae is a monophyletic family of Eurasian cyprinoid fishes. This is a species rich clade which, as a subfamily of the Cyprinidae was divided into five tribes: Gobionini, Pseudogobionini, Hemibarbini, Coreiini, and Sarcocheilichthyini. T ...

Bleeker, 1863 (freshwater gudgeons) ** Family Tanichthyidae Mayden & Chen, 2010 (mountain minnows) ** Family

Leuciscidae Leuciscidae is a family of freshwater ray-finned fishes, formerly classified as a subfamily of the Cyprinidae, which contains the true minnows. Members of the Old World (OW) clade of minnows within this subfamily are known as European minnow ...

Bonaparte, 1835 (minnows)

Phylogeny

Phylogeny based on the work of the following worksJörg 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)

Evolution

Cypriniformes include the most primitive of the Ostariophysi in the narrow sense (i.e. excluding the

Gonorynchiformes The Gonorynchiformes are an order of ray-finned fish that includes the important food source, the milkfish (''Chanos chanos'', family Chanidae), and a number of lesser-known types, both marine and freshwater. The alternate spelling "Gonorhy ...

). 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 (

and relatives) is thought to be about the

Early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between 251.9 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which ...

, about 250 million years ago ( mya). However, their divergence probably occurred only with the splitting-up of

Pangaea Pangaea or Pangea ( ) was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous period approximately 335 mi ...

in the

Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...

, maybe 160 million years ago (Mya). By 110 Mya, the

plate tectonics Plate tectonics (, ) is the scientific theory that the Earth's lithosphere comprises a number of large tectonic plates, which have been slowly moving since 3–4 billion years ago. The model builds on the concept of , an idea developed durin ...

evidence indicates that the

Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...

n Cypriniformes must have been distinct from their

Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...

n relatives. The Cypriniformes are thought to have originated in

South-east Asia Southeast Asia is the geographical southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Australian mainland, which is part of Oceania. Southeast Asia i ...

, where the most diversity of this group is found today. The alternative hypothesis is that they began in

, 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.Briggs (2005) Until 2025, no cypriniform fossil remains were known predating the

Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...

. In 2025, multiple fossil remains of an indeterminate cypriniform, including jaws, tooth plates, and vertebrae, were identified from the early

Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...

-aged (71-69 Mya)

Prince Creek Formation The Prince Creek Formation is a geological Formation (geology), formation in Alaska with strata dating to the Late Campanian stage of the Late Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weishamp ...

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 Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

Catostomidae; from the

-aged

Paskapoo Formation The Paskapoo Formation is a stratigraphic unit of Middle to Late Paleocene age in the Western Canada Sedimentary Basin. The Paskapoo underlies much of southwestern Alberta, and takes the name from the Blindman River ( means 'He is blind' in Cr ...

Alberta Alberta is a Provinces and territories of Canada, province in Canada. It is a part of Western Canada and is one of the three Canadian Prairies, prairie provinces. Alberta is bordered by British Columbia to its west, Saskatchewan to its east, t ...

, they are roughly 60 million years old. During the

(55–35 Mya), catostomids and cyprinids spread throughout Asia; the earliest members of the cyprinid subfamilies

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 membe ...

and Danioninae are known from the Eocene Sangkarewang Formation of

Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania, between the Indian Ocean, Indian and Pacific Ocean, Pacific oceans. Comprising over List of islands of Indonesia, 17,000 islands, including Sumatra, ...

, in addition to possibly Smilogastrinae and

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 else ...

. The extinct family

is known from the Eocene of China. In the

Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...

, around 30 Mya, advanced cyprinids began to outcompete catostomids wherever they were

sympatric In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...

, causing a decline of the suckers. Cyprinids reached North America and Europe about the same time, and Africa in the early

Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...

(some 23–20 Mya). The cypriniforms spread to North America through the

Bering land bridge Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72° north latitude in the Chukchi Sea; and on the south by the tip of the ...

, which formed and disappeared again several times during the many millions of years of cypriniform

evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...

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 The goldfish (''Carassius auratus'') is a freshwater fish in the family Cyprinidae of the order Cypriniformes. It is commonly kept as a pet in indoor aquariums, and is one of the most popular aquarium fish. Goldfish released into the w ...

and koi are perhaps the most celebrated. The other families are of less commercial importance. The Catostomidae have some importance in

angling Angling (from Old English ''angol'', meaning "hook") is a fishing technique that uses a fish hook attached to a fishing line to tether individual fish in the mouth. The fishing line is usually manipulated with a fishing rod, although rodless te ...

, and some "loaches" are bred for the international

aquarium An aquarium (: aquariums or aquaria) is a vivarium of any size having at least one transparent side in which aquatic plants or animals are kept and displayed. fishkeeping, Fishkeepers use aquaria to keep fish, invertebrates, amphibians, aquati ...

fish trade. Accidentally or deliberately introduced populations of

common carp The common carp (''Cyprinus carpio''), also known as European carp, Eurasian carp, or simply carp, is a widespread freshwater fish of eutrophic waters in lakes and large rivers in Europe and Asia.Fishbase''Cyprinus carpio'' Linnaeus, 1758/ref>Ark ...

(''Cyprinus carpio'') and

grass carp The grass carp (''Ctenopharyngodon idella'') is a species of large herbivorous freshwater fish in the family Cyprinidae, native to the Pacific Far East, with a native range stretching from northern Vietnam to the Amur River on the Sino-Russian ...

(''Ctenopharyngodon idella'') are found on all

continent A continent is any of several large geographical regions. Continents are generally identified by convention (norm), convention rather than any strict criteria. A continent could be a single large landmass, a part of a very large landmass, as ...

s except

Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...

. In some cases, these

exotic species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived ther ...

have a negative impact on the environment. Carp in particular stir up the riverbed, reducing the clarity of the water, making plant growth difficult. In science, one of the most famous members of the Cypriniformes is the

zebrafish The zebrafish (''Danio rerio'') is a species of freshwater ray-finned fish belonging to the family Danionidae of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (an ...

(''Danio rerio''). The zebrafish is one of the most important vertebrate

model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...

s 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

Habitat destruction Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...

, damming of upland rivers,

pollution Pollution is the introduction of contaminants into the natural environment that cause harm. Pollution can take the form of any substance (solid, liquid, or gas) or energy (such as radioactivity, heat, sound, or light). Pollutants, the component ...

, and in some cases

overfishing Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing Fish stocks, fish stock), resu ...

for food or the pet trade have driven some Cypriniformes to the brink of

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

ion or even beyond. In particular, Cyprinidae of southwestern

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 California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...

; 70 years later, not a single living individual existed.

The well-known red-tailed black shark (''Epalzeorhynchos bicolor'') from the

Mae Klong The Mae Klong (, , ), sometimes spelled Meklong, is a river A river is a natural stream of fresh water that flows on land or inside Subterranean river, caves towards another body of water at a lower elevation, such as an ocean, lake, o ...

River of ''

The Bridge on the River Kwai ''The Bridge on the River Kwai'' is a 1957 epic war film directed by David Lean and based on the novel ''The Bridge over the River Kwai'', written by Pierre Boulle. Boulle's novel and the film's screenplay are almost entirely fictional but u ...

'' 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. 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. Globally extinct Cypriniformes species are:IUCN (2007) * '' 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 ''Telestes ukliva'', the Cetina dace or Ukliva dace, is a species of ray-finned fish belonging to the family Leuciscidae, whch includes the daces, Eurasian minnows and related species. Endemic to the Cetina river in Croatia and reported as exti ...

Notes

References

* (2005): The biogeography of otophysan fishes (Ostariophysi: Otophysi): a new appraisal. '' J. Biogeogr.'' 32(2): 287–294. (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)

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. (HTML abstract) * (1997): ''The Diversity of Fishes''. Blackwell Publishing. * (2007): '' ww.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 ''Fishes of the World'' is a standard reference for the systematics of fishes. It was first written in 1976 by the American ichthyologist Joseph S. Nelson (1937–2011). Now in its fifth edition (2016), the work is a comprehensive overview of t ...

''. John Wiley & Sons, Inc. *

007 The ''James Bond'' franchise focuses on James Bond (literary character), the titular character, a fictional Secret Intelligence Service, British Secret Service agent created in 1953 by writer Ian Fleming, who featured him in twelve novels ...

FishBase FishBase is a global species database of fish species (specifically finfish). It is the largest and most extensively accessed online database on adult finfish on the web.

''Tribolodon brandtii''
Retrieved 2007-03-05. * (2003): Mitochondrial Genomics of Ostariophysan Fishes: Perspectives on Phylogeny and Biogeography. '' J. Mol. Evol.'' 56(4): 464–472. (HTML abstract) * (2018): Phylogenetic classification of extant genera of fishes of the order Cypriniformes (Teleostei: Ostariophysi). ''

Zootaxa ''Zootaxa'' is a peer-reviewed scientific mega journal for animal taxonomists. It is published by Magnolia Press (Auckland, New Zealand). The journal was established by Zhi-Qiang Zhang in 2001 and new issues are published multiple times a week ...

'' 4476(1): 006–039. (HTML abstract)

External links

Cypriniformes Tree of Life
{{Authority control Ray-finned fish orders Ostariophysi Fish of Africa Fish of Europe Fish of Asia Fish of North America Fish of Central America Extant Maastrichtian first appearances