Editing Cypriniformes (section)

== Systematics ==
[[File:Nemac fasci 080519 9380 ckoep.jpg|thumb|right|''[[Nemacheilus chrysolaimos]]'' is a stone loach. Closely related to true loaches, like these, they have barbels.]]
[[File:Chinese algae eater.jpg|thumb|right|The Chinese algae eater (''[[Gyrinocheilus aymonieri]]'') is one of the sucking loaches, which are distant from other "loaches".]]
[[File:Erimyzon sucetta.jpg|thumb|right|''[[Erimyzon sucetta]]'', a small sucker]]
Historically, these included all the forms now placed in the [[superorder]] [[Ostariophysi]] except the catfish, which were placed in the order [[Siluriformes]]. By this definition, the Cypriniformes were [[paraphyletic]], so recently, the orders [[Gonorhynchiformes]], [[Characiformes]], ([[characin]]s and allies), and [[Gymnotiformes]] ([[Gymnotiformes|knifefishes]] and [[electric eel]]s) have been separated out to form their own [[Monophyly|monophyletic]] orders.<ref>Helfman ''et al.'' (1997): pp.228-229</ref>

The [[family (biology)|families]] of Cypriniformes are traditionally divided into two [[Order (biology)#Hierarchy of ranks|suborders]]. Superfamily [[Cyprinioidea]] contains the carps and minnows ([[Cyprinidae]]) and also the mountain carps as the family [[Psilorhynchidae]].<ref name=Nelson/> In 2012, [[Maurice Kottelat]] reviewed the superfamily [[Cobitoidei]] 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]]).<ref name=Kottelat2012>Kottelat, M. (2012)</ref>

[[Catostomoidea]] is usually treated as a [[Synonym (taxonomy)#Zoology|junior synonym]] of the Cobitoidei, but it could be split off the Catostomidae and Gyrinocheilidae in a distinct superfamily; the Catostomoidea might be closer relatives of the carps and minnows than of the "true" loaches. While the Cyprinioidea seem more "primitive" than the loach-like forms,<ref name=Nelson/> they were apparently successful enough never to shift from the original [[ecological niche]] of the [[Basal (phylogenetics)|basal]] [[Ostariophysi]]. Yet, from the [[ecomorphological]]ly conservative main lineage apparently at least two major [[evolutionary radiation|radiation]]s branched off. These diversified from the lowlands into torrential river [[habitat]]s, acquiring similar [[Morphology (biology)|habitus]] and [[adaptation]]s [[convergent evolution|in the process]].<ref name = heetal2008 />

The mountain carps are the highly [[Cladistics#apomorphy|apomorph]]ic Cyprinidae, perhaps close to [[Cyprininae|true carps]] (Cyprininae), or maybe to the [[danionin]]s. While some details about the [[Phylogenetics|phylogenetic]] structures of this massively diverse family are known &ndash; e.g. that [[Cultrinae]] and [[Leuciscinae]] are rather close relatives and stand apart from Cyprininae &ndash;  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:<ref name=ECoF>{{cite web |url=https://www.calacademy.org/scientists/catalog-of-fishes-classification |title=Eschmeyer's Catalog of Fishes Classification |access-date=5 November 2024 |publisher=[[California Academy of Sciences]]}}</ref>[[File:Amyzon hunanensis 01.jpg|thumb|Fossil of ''[[Jianghanichthys]]'', the earliest known cypriniform]]
* Family †[[Jianghanichthyidae]] <small>Liu, Chang, Wilson & Murray, 2015</small> ([[Paleocene]] to [[Eocene]] of China)<ref name=":1">{{Cite journal |last1=Liu |first1=Juan |last2=Chang |first2=Mee-Mann |last3=Wilson |first3=Mark V. H. |last4=Murray |first4=Alison M. |date=2015-11-02 |title=A new family of Cypriniformes (Teleostei, Ostariophysi) based on a redescription of † Jianghanichthys hubeiensis (Lei, 1977) from the Eocene Yangxi Formation of China |url=http://www.tandfonline.com/doi/full/10.1080/02724634.2015.1004073 |journal=Journal of Vertebrate Paleontology |language=en |volume=35 |issue=6 |pages=e1004073 |doi=10.1080/02724634.2015.1004073 |bibcode=2015JVPal..35E4073L |issn=0272-4634}}</ref>
* Suborder [[Gyrinocheilus|Gyrinocheiloidei]] <small>[[Ricardo Betancur-R|Betancur-R]], ''et al.'', 2017</small>
** Family [[Gyrinocheilus|Gyrinocheilidae]] <small>[[Theodore Gill|Gill]], 1907</small> (algae eaters)
* Suborder [[Catostomidae|Catostomoidei]] <small>Betancur-R, et al., 2017</small>
** Family [[Catostomidae]] <small>[[Louis Agassiz|Agassiz]], 1850</small> (suckers)
* Suborder [[Cobitoidei]] <small>[[Leopold  Fitzinger|Fitzinger]], 1832</small>
** Family [[Botiidae]] <small>[[Lev Berg|Berg]]</small>, 1940 (pointface loaches)
** Family [[Vaillantellidae]] <small>Nalbant & [[Petru Mihai Bănărescu|Bănărescu]], 1977</small> (longfin loaches)
** Family [[Cobitidae]] <small>[[William Swainson|Swainson]], 1838</small> (spined loaches)
** Family [[Barbuccidae]] <small>[[Maurice Kottelat|Kottelat]], 2012</small> (scooter loaches)
** Family [[Gastromyzontidae]] <small>[[Henry Weed Fowler|Fowler]], 1905</small> (hillstream loaches)
** Family [[Serpenticobitidae]] <small>Kottelat, 2012</small> (snake loaches)
** Family [[Balitoridae]] <small>Swainson, 1839</small> (river loaches)
** Family [[Ellopostomatidae]] <small>[[Jörg Bohlen|Bohlen]] & [[Vendula Šlechtová|Šlechtová]], 2009</small> (square-head loaches)
** Family [[Nemacheilidae]] <small>[[Charles Tate Regan|Regan]], 1911</small> (brook loaches)
* Suborder [[Cyprinoidei]] <small>Fitzinger, 1832</small>
** Family [[Paedocyprididae]] <small>[[Richard L. Mayden|Mayden]] & [[Wei-Jen Chen|W.J. Chen]], 2010</small> (tiny carps)
** Family [[Psilorhynchidae]] <small>[[Sunder Lal Hora|Hora]], 1926</small> (mountain carps)
** Family [[Cyprinidae]] <small>[[Constantine Samuel Rafinesque|Rafinesque]], 1815</small> (carps)
** Family [[Sundadanionidae]] <small>Mayden & Chen, 2010</small> (tiny danios)
** Family [[Danionin|Danionidae]] <small>[[Pieter Bleeker|Bleeker]], 1863</small> (danionids)
** Family [[Leptobarbidae]] <small>Bleeker, 1864</small> (cigar barbs)
** Family [[Xenocyprididae]] <small>[[Albert Günther|Günther]], 1868</small> (East Asian minnows or sharpbellies)
** Family [[Tincidae]] <small>[[David Starr Jordan|D. S. Jordan]], 1878</small> (tenches)
** Family [[Acheilognathidae]] <small>Bleeker, 1863</small> (bitterlings)
** Family [[Gobioninae|Gobionidae]] <small>Bleeker, 1863</small> (freshwater gudgeons)
** Family [[Tanichthyidae]] <small>Mayden & Chen, 2010</small> (mountain minnows)
** Family [[Leuciscidae]] <small>[[Charles Lucien Bonaparte|Bonaparte]], 1835</small> (minnows)

===Phylogeny===
Phylogeny based on the work of the following works<ref name="slech">{{cite journal|pmid=17433724|year=2007|last1=Slechtová|first1=V.|title=Families of Cobitoidea (Teleostei; Cypriniformes) as revealed from nuclear genetic data and the position of the mysterious genera Barbucca, Psilorhynchus, Serpenticobitis and Vaillantella|journal=Molecular Phylogenetics and Evolution|volume=44|issue=3|pages=1358–65|last2=Bohlen|first2=J.|last3=Tan|first3=H. H.|doi=10.1016/j.ympev.2007.02.019|bibcode=2007MolPE..44.1358S }}</ref><ref name="chen">{{cite journal|doi=10.1111/j.1095-8649.2009.02398.x|pmid=20738682|title=Molecular phylogeny of the Cobitoidea (Teleostei: Cypriniformes) revisited: Position of enigmatic loach ''Ellopostomaresolved'' with six nuclear genes|journal=Journal of Fish Biology|volume=75|issue=9|pages=2197–2208|year=2009|last1=Chen|first1=W.-J.|last2=Lheknim|first2=V.|last3=Mayden|first3=R. L.|bibcode=2009JFBio..75.2197C }}</ref><ref name="bohlen">Jörg Bohlen, Vendula Šlechtová: ''Phylogenetic position of the fish genus ''Ellopostoma'' (Teleostei: Cypriniformes) using molecular genetic data.'' Ichthyological Exploration of Freshwaters. Bd.&nbsp;20, Nr.&nbsp;2, 2009, S.&nbsp;157–162 ([http://www.pfeil-verlag.de/04biol/pdf/ief20_2_05.pdf PDF] {{Webarchive|url=https://web.archive.org/web/20150924072138/http://www.pfeil-verlag.de/04biol/pdf/ief20_2_05.pdf |date=24 September 2015 }}; 1,8 MB)</ref><ref>{{cite web |last1 = Mikko Haaramo |title=Cobitoidei – loach-like cypriniforms |website=Mikko's Phylogeny Archive |access-date= 2013-10-26 |url=http://www.helsinki.fi/~mhaaramo/metazoa/deuterostoma/chordata/actinopterygii/cypriniformes/cobitoidei.html}}</ref><ref>{{Cite web|url=https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=638796#null|title=ITIS Standard Report Page: Cyprinoidea|website=www.itis.gov|access-date=2019-07-26}}</ref>

{{clade|style=font-size:90%;line-height:100%;width:700px;
|label1=Cypriniformes
|1={{clade
  |label1=Cyprinoidei
  |1={{clade
    |label1=Cyprinoidea
    |1={{clade
      |1=[[Cyprinidae]] (carps & minnows) [[File:Common carp (white background).jpg|70 px]]
      |2=[[Psilorhynchidae]] <span style="{{MirrorH}}">[[File:Psilorhynchus balitora Day Mintern 122.jpg|70 px]]</span>
    }}
  }}
  |label2=Cobitoidei
  |2={{clade
    |label1=Catostomoidea (suckers)
    |1=[[Catostomidae]] [[File:Ictiobus niger (white background).jpg|70 px]]
    |2={{clade
      |label1=Gyrinocheiloidea
      |1=[[Gyrinocheilidae]]
      |label2=[[Cobitoidea]] (loaches)
      |2={{clade
        |1=[[Botiidae]] <span style="{{MirrorH}}">[[File:Chromobotia macracanthus Bleeker.jpg|70 px]]</span>
        |2={{clade
          |1=[[Vaillantellidae]] <span style="{{MirrorH}}">[[File:Vaillantella euepiptera.png|70 px]]</span>
          |2={{clade
            |1=[[Cobitidae]] <span style="{{MirrorH}}">[[File:Acantopsis choirorhynchos Bleeker.jpg|70 px]]</span>
            |2={{clade
              |1=[[Balitoridae]] <span style="{{MirrorH}}">[[File:Homaloptera zollingeri Bleeker (cropped).jpg|70 px]]</span>
              |2={{clade
                |1=[[Ellopostomatidae]]
                |2=[[Nemacheilidae]] [[File:Nemachilus barbatulus (white background).jpg|70 px]]
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
   }}
}}

===Evolution===
Cypriniformes include the most primitive of the Ostariophysi in the narrow sense (i.e. excluding the [[Gonorynchiformes]]). This is evidenced not only by physiological details, but also by their great distribution, which indicates they had the longest time to spread. The earliest that Cypriniformes might have diverged from [[Characiphysi]] ([[Characiformes]] and relatives) is thought to be about the [[Early Triassic]], about 250 million years ago ([[mya (unit)|mya]]).<ref>Saitoh ''et al.'' (2003)</ref> However, their divergence probably occurred only with the splitting-up of [[Pangaea]] in the [[Jurassic]], maybe 160 million years ago (Mya). By 110 Mya, the [[plate tectonics]] evidence indicates that the [[Laurasia]]n Cypriniformes must have been distinct from their [[Gondwana]]n relatives.<ref>Briggs (2005), Nelson (2006)</ref>

The Cypriniformes are thought to have originated in [[South-east Asia]], where the most diversity of this group is found today. The alternative hypothesis is that they began in [[South America]], similar to the other [[Otophysi|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.<ref name=briggs>Briggs (2005)</ref>

Until 2025, no cypriniform fossil remains were known predating the [[Cenozoic]]. In 2025, multiple fossil remains of an indeterminate cypriniform, including jaws, tooth plates, and vertebrae, were identified from the early [[Maastrichtian]]-aged (71-69 Mya) [[Prince Creek Formation]] of Alaska. This occurrence suggests that in contrast to previous hypotheses, cypriniforms may have originated in high-latitude regions and radiated southwards during the Cenozoic.<ref name="Brinkman2025">{{Cite journal |last=Brinkman |first=Donald B. |last2=López |first2=J. Andrés |last3=Erickson |first3=Gregory M. |last4=Eberle |first4=Jaelyn J. |last5=Muñoz |first5=Xochitl |last6=Wilson |first6=Lauren N. |last7=Perry |first7=Zackary R. |last8=Murray |first8=Alison M. |last9=Van Loon |first9=Lisa |last10=Banerjee |first10=Neil R. |last11=Druckenmiller |first11=Patrick S. |date=2025 |title=Fishes from the Upper Cretaceous Prince Creek Formation, North Slope of Alaska, and their palaeobiogeographical significance |url=https://onlinelibrary.wiley.com/doi/full/10.1002/spp2.70014 |journal=Papers in Palaeontology |language=en |volume=11 |issue=3 |pages=e70014 |doi=10.1002/spp2.70014 |issn=2056-2802}}</ref> The next-oldest cypriniform fossils are already assignable to the living [[family (biology)|family]] Catostomidae; from the [[Paleocene]]-aged [[Paskapoo Formation]] of [[Alberta]], they are roughly 60 million years old. During the [[Eocene]] (55–35 Mya), catostomids and cyprinids spread throughout Asia; the earliest members of the cyprinid subfamilies [[Barbinae]] and [[Danionin|Danioninae]] are known from the Eocene [[Sangkarewang Formation]] of [[Indonesia]], in addition to possibly [[Smilogastrinae]] and [[Labeoninae]].<ref>{{Cite journal |last=Murray |first=Alison M. |date=2020-01-02 |title=Early Cenozoic Cyprinoids (Ostariophysi: Cypriniformes: Cyprinidae and Danionidae) from Sumatra, Indonesia |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2020.1762627 |journal=Journal of Vertebrate Paleontology |language=en |volume=40 |issue=1 |pages=e1762627 |doi=10.1080/02724634.2020.1762627 |bibcode=2020JVPal..40E2627M |issn=0272-4634}}</ref> The extinct family [[Jianghanichthyidae]] is known from the Eocene of China.<ref name=":1" /> In the [[Oligocene]], around 30 Mya, advanced cyprinids began to outcompete catostomids wherever they were [[Sympatry|sympatric]], causing a decline of the suckers. Cyprinids reached North America and Europe about the same time, and Africa in the early [[Miocene]] (some 23–20 Mya). The cypriniforms spread to North America through the [[Beringia|Bering land bridge]], which formed and disappeared again several times during the many millions of years of cypriniform [[evolution]].<ref name=briggs/>