Labeonins or labeoins are fish of the subfamily Labeoninae (or the tribe Labeonini, when subsumed into subfamily Cyprininae), which are ray-finned fishes in the family Cyprinidae of order Cypriniformes. They are freshwater fish, with the greatest species richness in the region around southern China, though they are present throughout subtropical and tropical Asia as well as equivalent regions in Africa. They are a generally very apomorphic group, perhaps the most "advanced" or derived of the Cyprinidae.

Anatomically, the labeonins are distinguished by traits of 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 ventrally; 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.

Notable genera include Crossocheilus, Epalzeorhynchos and Garra, which contain some of the popular aquarium fishes often called "algae eaters", e.g. the Siamese algae-eater (Crossocheilus siamensis) along with "pedicure fish" such as Garra rufa. Labeo – the type genus of this subfamily – contains many sizeable species which are often used as food.

Taxonomy

This clade 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); this classification is however not based on genetic studies. 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.[verification needed]

Genera

Labeoninae contains the following genera:

Phylogeny

The following is a maximum likelihood phylogenetic tree from a study of a combined genetic dataset, from Zheng et. al 2012, with binomial names updated and well-recovered clades labeled with A, B, C, and D:

LabeoninaeA Labeo bata Incisilabeo behri Labeo boga Cirrhinus microlepis Cirrhinus mrigalaLabeo spp.Labeo rohita Labeo stolizkae Labeo calbasu Labeo chrysophekadion Labeo pierrei Labeo batesii Labeo weeksii Labeo senegalensis B Ceratogarra fasciacaudaGarra spp.Garra flavatra Garra spilota Garra annandalei Garra gotyla gotyla Garra fuliginosa Garra orientalis Garra mirofrontis Garra ceylonensis Garra congoensis Garra makiensis Garra tengchongensis Tariqilabeo burmanicus Tariqilabeo bicornis C Labiobarbus lineatus Labiobarbus siamensisOsteochilus spp.Osteochilus melanopleurus Osteochilus microcephalus Osteochilus waandersii Osteochilus lini Osteochilus salsburyi Cirrhinus molitorella Lobocheilos melanotaeniaHenicorhynchus spp.Henicorhynchus lineatus Henicorhynchus lobatus Henicorhynchus siamensis Crossocheilus reticulatusEpalzeorhynchos spp.Epalzeorhynchos kalopterus Epalzeorhynchos bicolor Epalzeorhynchos frenatum D D1 Decorus lemassoni D2 Supradiscus micropulvinus Ageneiogarra imberba Pseudoplacocheilus cryptonema Mekongina erythrospila Altigena tonkinensis Altigena lippa D3 Parasinilabeo assimilis Semilabeo obscurus Rectoris posehensis Stenorynchoacrum xijiangensisPseudocrossocheilus spp.Pseudocrossocheilus liuchengensis Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus Pseudogyrinocheilus prochilus Qianlabeo striatus Ptychidio jordani Sinocrossocheilus labiatusHongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
ALabeo bata Incisilabeo behri Labeo boga Cirrhinus microlepis Cirrhinus mrigalaLabeo spp.Labeo rohita Labeo stolizkae Labeo calbasu Labeo chrysophekadion Labeo pierrei Labeo batesii Labeo weeksii Labeo senegalensis
Labeo bata Incisilabeo behri Labeo boga
Labeo bata
Incisilabeo behri Labeo boga
Incisilabeo behri
Labeo boga
Cirrhinus microlepis Cirrhinus mrigalaLabeo spp.Labeo rohita Labeo stolizkae Labeo calbasu Labeo chrysophekadion Labeo pierrei Labeo batesii Labeo weeksii Labeo senegalensis
Cirrhinus microlepis
Cirrhinus mrigalaLabeo spp.Labeo rohita Labeo stolizkae Labeo calbasu Labeo chrysophekadion Labeo pierrei Labeo batesii Labeo weeksii Labeo senegalensis
Cirrhinus mrigala
Labeo spp.Labeo rohita Labeo stolizkae Labeo calbasu Labeo chrysophekadion Labeo pierrei Labeo batesii Labeo weeksii Labeo senegalensis
Labeo spp.Labeo rohita Labeo stolizkae Labeo calbasu Labeo chrysophekadion Labeo pierrei Labeo batesii Labeo weeksii Labeo senegalensis
Labeo rohita Labeo stolizkae Labeo calbasu Labeo chrysophekadion
Labeo rohita Labeo stolizkae
Labeo rohita
Labeo stolizkae
Labeo calbasu Labeo chrysophekadion
Labeo calbasu
Labeo chrysophekadion
Labeo pierrei Labeo batesii Labeo weeksii Labeo senegalensis
Labeo pierrei
Labeo batesii Labeo weeksii Labeo senegalensis
Labeo batesii
Labeo weeksii Labeo senegalensis
Labeo weeksii
Labeo senegalensis
B Ceratogarra fasciacaudaGarra spp.Garra flavatra Garra spilota Garra annandalei Garra gotyla gotyla Garra fuliginosa Garra orientalis Garra mirofrontis Garra ceylonensis Garra congoensis Garra makiensis Garra tengchongensis Tariqilabeo burmanicus Tariqilabeo bicornis C Labiobarbus lineatus Labiobarbus siamensisOsteochilus spp.Osteochilus melanopleurus Osteochilus microcephalus Osteochilus waandersii Osteochilus lini Osteochilus salsburyi Cirrhinus molitorella Lobocheilos melanotaeniaHenicorhynchus spp.Henicorhynchus lineatus Henicorhynchus lobatus Henicorhynchus siamensis Crossocheilus reticulatusEpalzeorhynchos spp.Epalzeorhynchos kalopterus Epalzeorhynchos bicolor Epalzeorhynchos frenatum D D1 Decorus lemassoni D2 Supradiscus micropulvinus Ageneiogarra imberba Pseudoplacocheilus cryptonema Mekongina erythrospila Altigena tonkinensis Altigena lippa D3 Parasinilabeo assimilis Semilabeo obscurus Rectoris posehensis Stenorynchoacrum xijiangensisPseudocrossocheilus spp.Pseudocrossocheilus liuchengensis Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus Pseudogyrinocheilus prochilus Qianlabeo striatus Ptychidio jordani Sinocrossocheilus labiatusHongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
BCeratogarra fasciacaudaGarra spp.Garra flavatra Garra spilota Garra annandalei Garra gotyla gotyla Garra fuliginosa Garra orientalis Garra mirofrontis Garra ceylonensis Garra congoensis Garra makiensis Garra tengchongensis Tariqilabeo burmanicus Tariqilabeo bicornis
Ceratogarra fasciacauda
Garra spp.Garra flavatra Garra spilota Garra annandalei Garra gotyla gotyla Garra fuliginosa Garra orientalis Garra mirofrontis Garra ceylonensis Garra congoensis Garra makiensis Garra tengchongensis Tariqilabeo burmanicus Tariqilabeo bicornis
Garra spp.Garra flavatra Garra spilota Garra annandalei Garra gotyla gotyla Garra fuliginosa Garra orientalis Garra mirofrontis Garra ceylonensis Garra congoensis Garra makiensis
Garra spp.Garra flavatra Garra spilota Garra annandalei Garra gotyla gotyla Garra fuliginosa Garra orientalis Garra mirofrontis Garra ceylonensis Garra congoensis Garra makiensis
Garra flavatra
Garra spilota Garra annandalei Garra gotyla gotyla Garra fuliginosa Garra orientalis Garra mirofrontis Garra ceylonensis Garra congoensis Garra makiensis
Garra spilota
Garra annandalei Garra gotyla gotyla Garra fuliginosa Garra orientalis Garra mirofrontis Garra ceylonensis Garra congoensis Garra makiensis
Garra annandalei Garra gotyla gotyla
Garra annandalei
Garra gotyla gotyla
Garra fuliginosa Garra orientalis Garra mirofrontis Garra ceylonensis Garra congoensis Garra makiensis
Garra fuliginosa Garra orientalis Garra mirofrontis
Garra fuliginosa
Garra orientalis Garra mirofrontis
Garra orientalis
Garra mirofrontis
Garra ceylonensis Garra congoensis Garra makiensis
Garra ceylonensis
Garra congoensis Garra makiensis
Garra congoensis
Garra makiensis
Garra tengchongensis Tariqilabeo burmanicus Tariqilabeo bicornis
Garra tengchongensis
Tariqilabeo burmanicus Tariqilabeo bicornis
Tariqilabeo burmanicus
Tariqilabeo bicornis
C Labiobarbus lineatus Labiobarbus siamensisOsteochilus spp.Osteochilus melanopleurus Osteochilus microcephalus Osteochilus waandersii Osteochilus lini Osteochilus salsburyi Cirrhinus molitorella Lobocheilos melanotaeniaHenicorhynchus spp.Henicorhynchus lineatus Henicorhynchus lobatus Henicorhynchus siamensis Crossocheilus reticulatusEpalzeorhynchos spp.Epalzeorhynchos kalopterus Epalzeorhynchos bicolor Epalzeorhynchos frenatum D D1 Decorus lemassoni D2 Supradiscus micropulvinus Ageneiogarra imberba Pseudoplacocheilus cryptonema Mekongina erythrospila Altigena tonkinensis Altigena lippa D3 Parasinilabeo assimilis Semilabeo obscurus Rectoris posehensis Stenorynchoacrum xijiangensisPseudocrossocheilus spp.Pseudocrossocheilus liuchengensis Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus Pseudogyrinocheilus prochilus Qianlabeo striatus Ptychidio jordani Sinocrossocheilus labiatusHongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
CLabiobarbus lineatus Labiobarbus siamensisOsteochilus spp.Osteochilus melanopleurus Osteochilus microcephalus Osteochilus waandersii Osteochilus lini Osteochilus salsburyi Cirrhinus molitorella Lobocheilos melanotaeniaHenicorhynchus spp.Henicorhynchus lineatus Henicorhynchus lobatus Henicorhynchus siamensis Crossocheilus reticulatusEpalzeorhynchos spp.Epalzeorhynchos kalopterus Epalzeorhynchos bicolor Epalzeorhynchos frenatum
Labiobarbus lineatus Labiobarbus siamensisOsteochilus spp.Osteochilus melanopleurus Osteochilus microcephalus Osteochilus waandersii Osteochilus lini Osteochilus salsburyi
Labiobarbus lineatus Labiobarbus siamensis
Labiobarbus lineatus
Labiobarbus siamensis
Osteochilus spp.Osteochilus melanopleurus Osteochilus microcephalus Osteochilus waandersii Osteochilus lini Osteochilus salsburyi
Osteochilus spp.Osteochilus melanopleurus Osteochilus microcephalus Osteochilus waandersii Osteochilus lini Osteochilus salsburyi
Osteochilus melanopleurus
Osteochilus microcephalus Osteochilus waandersii Osteochilus lini Osteochilus salsburyi
Osteochilus microcephalus
Osteochilus waandersii Osteochilus lini Osteochilus salsburyi
Osteochilus waandersii
Osteochilus lini Osteochilus salsburyi
Osteochilus lini
Osteochilus salsburyi
Cirrhinus molitorella Lobocheilos melanotaeniaHenicorhynchus spp.Henicorhynchus lineatus Henicorhynchus lobatus Henicorhynchus siamensis Crossocheilus reticulatusEpalzeorhynchos spp.Epalzeorhynchos kalopterus Epalzeorhynchos bicolor Epalzeorhynchos frenatum
Cirrhinus molitorella
Lobocheilos melanotaeniaHenicorhynchus spp.Henicorhynchus lineatus Henicorhynchus lobatus Henicorhynchus siamensis Crossocheilus reticulatusEpalzeorhynchos spp.Epalzeorhynchos kalopterus Epalzeorhynchos bicolor Epalzeorhynchos frenatum
Lobocheilos melanotaeniaHenicorhynchus spp.Henicorhynchus lineatus Henicorhynchus lobatus Henicorhynchus siamensis
Lobocheilos melanotaenia
Henicorhynchus spp.Henicorhynchus lineatus Henicorhynchus lobatus Henicorhynchus siamensis
Henicorhynchus spp.Henicorhynchus lineatus Henicorhynchus lobatus Henicorhynchus siamensis
Henicorhynchus lineatus
Henicorhynchus lobatus Henicorhynchus siamensis
Henicorhynchus lobatus
Henicorhynchus siamensis
Crossocheilus reticulatusEpalzeorhynchos spp.Epalzeorhynchos kalopterus Epalzeorhynchos bicolor Epalzeorhynchos frenatum
Crossocheilus reticulatus
Epalzeorhynchos spp.Epalzeorhynchos kalopterus Epalzeorhynchos bicolor Epalzeorhynchos frenatum
Epalzeorhynchos spp.Epalzeorhynchos kalopterus Epalzeorhynchos bicolor Epalzeorhynchos frenatum
Epalzeorhynchos kalopterus
Epalzeorhynchos bicolor Epalzeorhynchos frenatum
Epalzeorhynchos bicolor
Epalzeorhynchos frenatum
DD1 Decorus lemassoni D2 Supradiscus micropulvinus Ageneiogarra imberba Pseudoplacocheilus cryptonema Mekongina erythrospila Altigena tonkinensis Altigena lippa D3 Parasinilabeo assimilis Semilabeo obscurus Rectoris posehensis Stenorynchoacrum xijiangensisPseudocrossocheilus spp.Pseudocrossocheilus liuchengensis Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus Pseudogyrinocheilus prochilus Qianlabeo striatus Ptychidio jordani Sinocrossocheilus labiatusHongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
D1Decorus lemassoni
D2 Supradiscus micropulvinus Ageneiogarra imberba Pseudoplacocheilus cryptonema Mekongina erythrospila Altigena tonkinensis Altigena lippa D3 Parasinilabeo assimilis Semilabeo obscurus Rectoris posehensis Stenorynchoacrum xijiangensisPseudocrossocheilus spp.Pseudocrossocheilus liuchengensis Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus Pseudogyrinocheilus prochilus Qianlabeo striatus Ptychidio jordani Sinocrossocheilus labiatusHongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
D2Supradiscus micropulvinus Ageneiogarra imberba Pseudoplacocheilus cryptonema Mekongina erythrospila Altigena tonkinensis Altigena lippa
Supradiscus micropulvinus Ageneiogarra imberba
Supradiscus micropulvinus
Ageneiogarra imberba
Pseudoplacocheilus cryptonema Mekongina erythrospila Altigena tonkinensis Altigena lippa
Pseudoplacocheilus cryptonema
Mekongina erythrospila Altigena tonkinensis Altigena lippa
Mekongina erythrospila
Altigena tonkinensis Altigena lippa
Altigena tonkinensis
Altigena lippa
D3Parasinilabeo assimilis Semilabeo obscurus Rectoris posehensis Stenorynchoacrum xijiangensisPseudocrossocheilus spp.Pseudocrossocheilus liuchengensis Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus Pseudogyrinocheilus prochilus Qianlabeo striatus Ptychidio jordani Sinocrossocheilus labiatusHongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
Parasinilabeo assimilis Semilabeo obscurus Rectoris posehensis Stenorynchoacrum xijiangensis
Parasinilabeo assimilis
Semilabeo obscurus Rectoris posehensis Stenorynchoacrum xijiangensis
Semilabeo obscurus
Rectoris posehensis Stenorynchoacrum xijiangensis
Rectoris posehensis
Stenorynchoacrum xijiangensis
Pseudocrossocheilus spp.Pseudocrossocheilus liuchengensis Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus Pseudogyrinocheilus prochilus Qianlabeo striatus Ptychidio jordani Sinocrossocheilus labiatusHongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
Pseudocrossocheilus spp.Pseudocrossocheilus liuchengensis Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus
Pseudocrossocheilus spp.Pseudocrossocheilus liuchengensis Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus
Pseudocrossocheilus liuchengensis
Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus
Pseudocrossocheilus tridentis Pseudocrossocheilus papillolabrus
Pseudocrossocheilus tridentis
Pseudocrossocheilus papillolabrus
Pseudocrossocheilus longibullus Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus
Pseudocrossocheilus longibullus
Pseudocrossocheilus bamaensis Pseudocrossocheilus nigrovittatus
Pseudocrossocheilus bamaensis
Pseudocrossocheilus nigrovittatus
Pseudogyrinocheilus prochilus Qianlabeo striatus Ptychidio jordani Sinocrossocheilus labiatusHongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
Pseudogyrinocheilus prochilus Qianlabeo striatus
Pseudogyrinocheilus prochilus
Qianlabeo striatus
Ptychidio jordani Sinocrossocheilus labiatusHongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
Ptychidio jordani
Sinocrossocheilus labiatusHongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
Sinocrossocheilus labiatus
Hongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoliDiscogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
Hongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoli
Hongshuia spp.Hongshuia megalophthalmus Hongshuia microstomatus Hongshuia paoli
Hongshuia megalophthalmus
Hongshuia microstomatus Hongshuia paoli
Hongshuia microstomatus
Hongshuia paoli
Discogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
Discogobio & Discocheilus spp.Discogobio yunnanensis Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
Discogobio yunnanensis
Discocheilus wui Discocheilus wuluoheensis Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
Discocheilus wui Discocheilus wuluoheensis
Discocheilus wui
Discocheilus wuluoheensis
Discogobio tetrabarbatus Discogobio brachyphysallidos Discogobio macrophysallidos
Discogobio tetrabarbatus
Discogobio brachyphysallidos Discogobio macrophysallidos
Discogobio brachyphysallidos
Discogobio macrophysallidos