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 160 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).
Description and ecology
These species are slim cyprinids with a flat belly and a sucking mouth; their shape indicates that they are at least in tendency rheophilic. They are distinguished from other cyprinids by a combination of features: As in their closest relatives, their lower lip is expanded at its posterior rim to form a round or oval sucking pad, the vomero-palatine organ is much reduced or completely lost, the pectoral fins have at least the first two rays enlarged and usually unbranched, the supraethmoid is wider than long when seen from above, and the cleithrum is narrow and elongated to the front.
From other Garrini (or Garraina), the genus Garra can be distinguished as follows: their pharyngeal teeth are arranged in three rows (like 2,4,5âÂÂ5,4,2), the dorsal fin has 10 or 11 rays and starts slightly anterior to the pelvic fins, while the anal fin starts well behind the pelvic fins and has 8 or 9 rays. As far as is known, the diploid karyotype of garras is 2n = 50.
Garras are not or barely noticeably sexually dimorphic and generally cryptically coloured benthic freshwater fish. Six species in the genus (G. andruzzii, G. dunsirei, G. lorestanensis, G. tashanensis, G. typhlops and some populations of G. barreimiae) are cave-adapted, lacking pigmentation and/or eyes. Garras are omnivorous, eating alga, plankton and small invertebrates that they suck off substrate like rocks or logs. The food is scraped off with the sharp keratinized borders of the jaws and ingested via suction, created by contracting and relaxing the buccopharynx. As typical for Cypriniformes, the garras lack a stomach entirely, their oesophagus leading directly to the sphincter of the intestine. Different Garra species eat animal and vegetable matter in different proportions, which can – as typical for vertebrates – usually be recognized by the length of their intestine compared to related species: more herbivorous species have a longer intestine. Indeed, intestinal length in this genus is remarkably constant within species and varies a lot between species, meaning that it is useful to distinguish species and that dietary shifts have played a significant role in the evolution of garras.
When the females are ready to spawn, they are markedly plump and swollen; the ripe roe may fill almost four-fifths of their body cavity. The testicles of reproductive males are large too. The average Garra egg is 1.77 mm in diameter and a clutch contains several hundred eggs – up to a thousand or so in large females. The breeding behaviour is generally not well known and breeding is not often achieved in the aquarium; presumably, like many of their relatives they migrate upstream or (if they otherwise inhabit lakes) into the rivers to spawn.
Taxonomy
The genus was established by Francis Buchanan-Hamilton in 1822 as a subgenus of Cyprinus (which at that time was a "basket genus" for carp-like cyprinids); he did not designate a type species, though as the only member of this subgenus was Garra lamta it became the type species of the genus after Pieter Bleeker did a taxonomic revision 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 was considered a close relative, but this is no longer thought to be the case.
Phylogeny
The following maximum likelihood phylogenetic tree is based on a dataset of nuclear and mitochondrial DNA, from Yang et al. 2012, with updated binomial names:
"Garra" imberba and "G." micropulvinus were recovered in a more derived position than Garra sensu stricto; the two were recovered within the same clade as Discogobio and Pseudocrossocheilus, so they were subsequently reassigned to new genera.
Species
These are the currently recognized species in this genus:
- Garra abhoyai <small>Hora, 1921</small>
- Garra aethiopica <small>(Pellegrin, 1927)</small>
- Garra allostoma <small>T. R. Roberts, 1990</small>
- Garra alticaputus <small>Arunachalam, Nandagopal & Mayden, 2013</small>
- Garra alticauda <small>L. Cao, Nyingi, Bart & E. Zhang, 2023</small>
- Garra amirhosseini <small>Esmaeili, Sayyadzadeh, Coad & Eagderi, 2016</small>
- Garra andruzzii <small>(Vinciguerra, 1924)</small> (Somalian cavefish)
- Garra annandalei <small>Hora, 1921</small>
- Garra apogon <small>(Norman, 1925)</small>
- Garra arunachalami <small>J. J. Johnson & R. Soranam, 2001</small>
- Garra arunachalensis <small>Nebeshwar & Vishwanath, 2013</small>
- Garra arupi <small>Nebeshwar, Vishwanath & D. N. Das, 2009</small>
- Garra barreimiae <small>Fowler & Steinitz, 1956</small>
- Garra bicornuta <small>Narayan Rao, 1920</small>
- Garra biloborostris <small>Roni & Vishwanath, 2017</small>
- Garra bimaculacauda <small>Thoni, Gurung & Mayden, 2016</small>
- Garra binduensis <small>U. Das, Kosygin & Panigrahi, 2016</small>
- Garra birostris <small>Nebeshwar & Vishwanath, 2013</small>
- Garra bispinosa <small>E. Zhang, 2005</small>
- Garra blanfordii <small>(Boulenger, 1901)</small>
- Garra borneensis <small>(Vaillant, 1902)</small>
- Garra bourreti <small>(Pellegrin, 1928)</small>
- Garra buettikeri <small>Krupp, 1983</small>
- Garra caudomaculata <small>(Battalgil, 1942)</small> (Antakya minnow)
- Garra centrala <small>V. H. Nguyá»Â
n, T.H.N. VÃ
© & T.D.P. Nguyá»Â
n, 2015</small>
- Garra ceylonensis <small>Bleeker, 1863</small> (Ceylon logsucker)
- Garra chakpiensis <small>Nebeshwar & Vishwanath, 2015</small>
- Garra chathensis <small>Ezung, Shangningam & Pankaj, 2020</small>
- Garra chaudhurii <small>Hora, 1921</small>
- Garra chebera <small>Habteselassie, Mikschi, Ahnelt & Waidbacher, 2010</small>
- Garra chindwinensis <small>Premananda, Kosygin, Saidullah, 2017</small>
- Garra chingaiensis <small>Abonmai, Linthoingambi, Ngangbam, Thoidingjam & K. R. Singh, 2023</small>
- Garra chivaensis <small>Moyon & L. Arunkumar, 2020</small>
- Garra clavirostris <small>Roni, Sarbojit & Vishwanath, 2017</small>
- Garra compressa <small>Kosygin & Vishwanath, 1998</small>
- Garra congoensis <small>Poll, 1959</small>
- Garra cornigera <small>Shangningam & Vishwanath, 2015</small>
- Garra culiciphaga <small>(Pellegrin, 1927)</small> (Redstripe barb)
- Garra cyclostomata <small>ÃÂ. Y. Mai, 1978</small>
- Garra cyrano <small>Kottelat, 2000</small>
- Garra dampaensis <small>Lalronunga, Lalnuntluanga & Lalramliana, 2013</small>
- Garra deccanensis <small>Jadhav, Karuthapandi, Shangningam, Jaiswal & Shankar, 2022</small>
- Garra dembecha <small>Getahun & Stiassny, 2007</small>
- Garra dembeensis <small>(Rüppell, 1835)</small> (Dembea stone lapper)
- Garra dengba <small>S.-Q. Deng, L. Cao & E. Zhang, 2018</small>
- Garra dohjei <small>Marngar, Mawlong & Lokeshwor, 2024</small>
- Garra dulongensis <small>(Chen, X. F. Pan, X. Hong & J. X. Yang, 2006)</small>
- Garra dunsirei <small>Banister, 1987</small>
- Garra duobarbis <small>Getahun & Stiassny, 2007</small>
- Garra elegans <small>(Günther, 1868)</small>
- Garra elongata <small>Vishwanath & Kosygin, 2000</small>
- Garra emarginata <small>Kurup & Radhakrishnan, 2011</small>
- Garra ethelwynnae <small>Menon, 1958</small>
- Garra festai <small>(Tortonese, 1939)</small>
- Garra flavatra <small>S. O. Kullander & F. Fang, 2004</small>
- Garra fluviatilis <small>Kangrang, Thoni, Mayden & Beamish, 2016</small>
- Garra fuliginosa <small>Fowler, 1934</small>
- Garra gallagheri <small>Krupp, 1988</small> (Black Garra)
- Garra geba <small>Getahun & Stiassny, 2007</small>
- Garra ghorensis <small>Krupp, 1982</small>
- Garra gotyla <small>(J. E. Gray, 1830)</small> (Sucker head)
- Garra gracilis <small>(Pellegrin & Chevey, 1936)</small>
- Garra gravelyi <small>(Annandale, 1919)</small>
- Garra gymnothorax <small>Berg, 1949</small> (Chest scaleless garra)
- Garra hexagonarostris <small>Nebeshwar, Rameshori & Prabina, 2024</small>
- Garra hindii <small>(Boulenger, 1905)</small>
- Garra hormuzensis <small>Zamani-Faradonbe, E. Zhang & Keivany, 2021</small>
- Garra hughi <small>Silas, 1955</small>
- Garra ignestii <small>(Gianferrari, 1925)</small>
- Garra imberbis <small>(Vinciguerra, 1890)</small>
- Garra irangensis <small>Premananda, 2023</small>
- Garra jaldhakaensis <small>Kosygin, Shangningam, Singh & U. Das, 2021</small>
- Garra jamila <small>Moritz, Straube & Neumann, 2019</small>
- Garra jenkinsonianum <small>Hora, 1921</small>
- Garra jerdoni <small>F. Day, 1867</small>
- Garra jordanica <small>Hamidan, Geiger & Freyhof, 2014</small>
- Garra joshuai <small>(Silas, 1954)</small>
- Garra kalakadensis <small>Rema Devi, 1993</small>
- Garra kalpangi <small>Nebeshwar, Bagra & D. N. Das, 2012</small>
- Garra kangrae <small>Prashad, 1919</small>
- Garra kemali <small>(Hankó, 1925)</small> (Eregil minnow)
- Garra kempi <small>Hora, 1921</small>
- Garra khawbungi <small>Arunachalam, Nandagopal & Mayden, 2014</small>
- Garra kimini <small>Arunachalam, Nandagopal & Mayden, 2014</small>
- Garra klatti <small>(Kosswig, 1950)</small>
- Garra koladynensis <small>Nebeshwar & Vishwanath, 2017</small>
- Garra laishrami <small>Surachita, Chowdhury & Palita, 2023</small>
- Garra lamta <small>(F. Hamilton, 1822)</small>
- Garra lancrenonensis <small>Blache & Miton, 1960</small>
- Garra langlungensis <small>Ezung, Shangningam & Pankaj, 2021</small>
- Garra lautior <small>Banister, 1987</small>
- Garra lissorhynchus <small>(McClelland, 1842)</small>
- Garra litanensis <small>Vishwanath, 1993</small>
- Garra longchuanensis <small>Q. Yu, X. Z. Wang, H. Xiong & S. P. He, 2016</small>
- Garra longipinnis <small>Banister & M. A. Clarke, 1977</small>
- Garra lorestanensis <small>Mousavi-Sabet & Eagderi, 2016</small>
- Garra lungongza <small>Ngangbam & Lithoingambi, 2023</small>
- Garra magnacavus <small>Shangningam, Kosygin & Sinha, 2019</small>
- Garra magnidiscus <small>Tamang, 2013</small>
- Garra makiensis <small>(Boulenger, 1904)</small>
- Garra mamshuqa <small>Krupp, 1983</small>
- Garra manipurensis <small>Vishwanath & Sarojnalini, 1988</small>
- Garra matensis <small>Nebeshwar & Vishwanath, 2017</small>
- Garra mcclellandi <small>(Jerdon, 1849)</small>
- Garra menderesensis <small>(Küçük, Bayçelebi, Güçlü & Gülle, 2015)</small>
- Garra menoni <small>Rema Devi & T. J. Indra, 1984</small>
- Garra meymehensis <small>Zamani-Faradonbe, Keivany, Dorafshan & E. Zhang, 2021</small>
- Garra mini <small>Rahman, Mollah, Norén & S. O. Kullander, 2016</small>
- Garra minibarbata <small>L. Cao, Nyingi, Bart & E. Zhang 2023</small>
- Garra minimus <small>Arunachalam, Nandagopal & Mayden, 2013</small>
- Garra mirofrontis <small>X. L. Chu & G. H. Cui, 1987</small>
- Garra mlapparaensis <small>Kurup & Radhakrishnan, 2011</small>
- Garra mondica <small>Sayyadzadeh, Esmaeili & Freyhof, 2015</small>
- Garra montisalsi <small>Hora, 1921</small>
- Garra motuoensis <small>Z. Gong, Freyhof, J. Wang, M. Liu, F. Liu, P. C. Lin, Y. Jiang & H. Z. Liu 2018</small>
- Garra moyonkhulleni <small>Moyon & Arunkumar, 2018</small>
- Garra mullya <small>(Sykes, 1839)</small> (sucker fish)
- Garra naganensis <small>Hora, 1921</small>
- Garra nambulica <small>Vishwanath & Joyshree, 2005</small>r.
- Garra namyaensis <small>Shangningam & Vishwanath, 2012</small>
- Garra nana <small>(Heckel, 1843)</small>
- Garra napata <small>Moritz, Straube & Neumann, 2019</small>
- Garra nasuta <small>(McClelland, 1838)</small>
- Garra nepalensis <small>Rayamajhi & Arunachalam, 2017</small>
- Garra nethravathiensis <small>Arunachalam & Nandagopal, 2014</small>
- Garra ngatangkha <small>Arunkumar & Moyon, 2019</small>
- Garra ngopi <small>Tenali, Chandran, R. K. Singh & Sarkar, 2024</small>
- Garra nigricauda <small>Arunachalam, Nandagopal & Mayden, 2013</small>
- Garra nigricollis <small>S. O. Kullander & F. Fang, 2004</small>
- Garra notata <small>(Blyth, 1860)</small>
- Garra nudiventris <small>(Berg, 1905)</small>
- Garra orientalis <small>Nichols, 1925</small>
- Garra ornata <small>(Nichols & Griscom, 1917)</small>
- Garra orontesi <small>Bayçelebi, Kaya, Turan & Freyhof, 2021</small>
- Garra palaniensis <small>(Rema Devi & Menon, 1994)</small>
- Garra palaruvica <small>Arunachalam, Raja, Nandagopal & Mayden, 2013</small>
- Garra panitvongi <small>Tangjitjaroen, Z. S. Randall, Tongnunui, Boyd & Page, 2023</small>
- Garra paralissorhynchus <small>Vishwanath & K. Shanta Devi, 2005</small>
- Garra parastenorhynchus <small>Thoni, Gurung & Mayden, 2016</small>
- Garra paratrilobata <small>Roni, Chinglemba, Rameshori & Vishwanath, 2019</small>
- Garra periyarensis <small>K. C. Gopi, 2001</small>
- Garra persica <small>Berg, 1914</small>
- Garra phewakholaensis <small>Limbu, Rajbanshi & Raza 2024</small>
- Garra phillipsi <small>Deraniyagala, 1933</small> (Phillips's garra)
- Garra platycephala <small>Narayan Rao, 1920</small>
- Garra poecilura <small>S. O. Kullander & F. Fang 2004</small>
- Garra poilanei <small>Petit & T. L. Tchang, 1933</small>
- Garra prashadi <small>Hora, 1921</small>
- Garra propulvinus <small>S. O. Kullander & F. Fang 2004</small>
- Garra qiaojiensis <small>H. W. Wu & Yao, 1977</small>
- Garra quadratirostris <small>Nebeshwar & Vishwanath, 2013</small>
- Garra quadrimaculata <small>(Rüppell 1835)</small>
- Garra quangtriensis <small>V. H. Nguyá»Â
n, T.H.N. VÃ
© & T. D. P. Nguyá»Â
n, 2015</small>
- Garra rakhinica <small>S. O. Kullander & F. Fang 2004</small>
- Garra ranganensis <small>Tamang, Sinha, Abujam & Kumar, 2019</small>
- Garra regressa <small>Getahun & Stiassny, 2007</small>
- Garra rezai <small>Mousavi-Sabet, Eagderi, Saemi-Komsari, Kaya & Freyhof, 2022</small>
- Garra rhynchota <small>Koller, 1926</small>
- Garra robertsi <small>Thoni & Mayden, 2015</small>
- Garra roseae <small>Mousavi-Sabet, Saemi-Komsari, Doadrio & Freyhof, 2019</small>
- Garra rossica <small>(A. M. Nikolskii, 1900)</small>
- Garra rotundinasus <small>E. Zhang, 2006</small>
- Garra rufa <small>(Heckel, 1843)</small> (Red garra)
- Garra rupecula <small>(McClelland, 1839)</small>
- Garra sahilia <small>Krupp, 1983</small>
- Garra salweenica <small>Hora & Mukerji, 1934</small>
- Garra sannarensis <small>Moritz, Straube & Neumann, 2019</small>
- Garra sauvagei <small>(Lortet, 1883)</small>
- Garra shamal <small>Kirchner, Kruckenhauser, Pichler, Borkenhagen & Freyhof, 2020</small>
- Garra sharq <small>Kirchner, Kruckenhauser, Pichler, Borkenhagen & Freyhof, 2020</small>
- Garra simbalbaraensis <small>Rath, Shangningam & Kosygin, 2019</small>
- Garra sindhi <small>Lyon, Geiger & Freyhof, 2016</small>.
- Garra songbangensis <small>V. H. Nguyá»Â
n, T.H.N. VÃ
© & T. D. P. Nguyá»Â
n, 2015</small>
- Garra spilota <small>S. O. Kullander & F. Fang, 2004</small>
- Garra stenorhynchus <small>Jerdon, 1849</small>
- Garra substrictorostris <small>Roni & Vishwanath, 2018</small>
- Garra surendranathanii <small>C. P. Shaji, L. K. Arun & P. S. Easa, 1996</small>
- Garra surgifrons <small>W. Zhou & C. Sun, 2018</small>
- Garra surinbinnani <small>Page, Ray, Tongnunui, Boyd & Randall, 2019</small>
- Garra tamangi <small>Gurumayum & Kosygin, 2016</small>
- Garra tana <small>Getahun & Stiassny, 2007</small>
- Garra tashanensis <small>Mousavi-Sabet, Vatandoust, Fatemi & Eagderi, 2016</small>
- Garra tengchongensis <small>E. Zhang & Y. Y. Chen, 2002</small>
- Garra tezuensis <small>Thoidingjam, Ngangbam, Linthoingambi & Kh. R. Singh, 2023</small>
- Garra tiam <small>Zamani-Faradonbe, Keivany, Dorafshan & E. Zhang, 2021</small>
- Garra tibanica <small>Trewavas, 1941</small>
- Garra tibetana <small>G. Zheng; S. Q. Deng; J. Wang; H. Z. Lui, 2018</small>
- Garra trewavasae <small>Monod, 1950</small>
- Garra triangularis <small>Shangningam, Rath & Kosygin, 2021</small>
- Garra trilobata <small>Shangningam & Vishwanath, 2015</small>
- Garra turcica <small>Karaman 1971</small>
- Garra typhlops <small>(Bruun & E. W. Kaiser, 1944)</small> (Iran cave barb)
- Garra ukhrulensis <small>Nebeshwar & Vishwanath, 2015</small>
- Garra variabilis <small>(Heckel, 1843)</small> (Variable garra)
- Garra vinciguerrae <small>(Boulenger 1901)</small> (Cameroon logsucker)
- Garra waensis <small>Lothongkham, Arbsuwan & Musikasinthorn, 2014</small>
- Garra wanae <small>(Regan, 1914)</small>
- Garra waterloti <small>(Pellegrin, 1936)</small>
- Garra widdowsoni <small>(Trewavas, 1955)</small> (Iraq blind barb)
- Garra yajiangensis <small>Gong, Freyhof, Wang, Liu, Liu, Lin, Jiang & Liu, 2018</small>
- Garra zubzaensis <small>Tenali, Shangningam, Bhattacharjee, Patel & Kosygin, 2024</small>
References