Garra


Garra is a genus of fish in the family Cyprinidae. These fish are one example of the "log suckers", sucker-mouthed barbs and other cyprinids commonly kept in aquaria to keep down algae. The doctor fish of Anatolia and the Middle East belongs in this genus. The majority of the more than 160 species of garras are native to Asia, but about one-fifth of the species are from Africa.

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, the genus Garra can be distinguished as follows: their pharyngeal teeth are arranged in three rows, 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 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 into the rivers to spawn.

Taxonomy

The genus was established by Francis Buchanan-Hamilton in 1822 as a subgenus of Cyprinus ; 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 Hora, 1921Garra aethiopica Garra allostoma T. R. Roberts, 1990Garra alticaputus Arunachalam, Nandagopal & Mayden, 2013Garra alticauda L. Cao, Nyingi, Bart & E. Zhang, 2023Garra amirhosseini Esmaeili, Sayyadzadeh, Coad & Eagderi, 2016Garra andruzzii Garra annandalei Hora, 1921Garra apogon Garra arunachalami J. J. Johnson & R. Soranam, 2001Garra arunachalensis Nebeshwar & Vishwanath, 2013Garra arupi Nebeshwar, Vishwanath & D. N. Das, 2009Garra barreimiae Fowler & Steinitz, 1956Garra bicornuta Narayan Rao, 1920Garra biloborostris Roni & Vishwanath, 2017 Garra bimaculacauda Thoni, Gurung & Mayden, 2016Garra binduensis U. Das, Kosygin & Panigrahi, 2016Garra birostris Nebeshwar & Vishwanath, 2013Garra bispinosa E. Zhang, 2005Garra blanfordii Garra borneensis Garra bourreti Garra buettikeri Krupp, 1983Garra caudomaculata Garra centrala V. H. Nguyễn, T.H.N. Vũ & T.D.P. Nguyễn, 2015Garra ceylonensis Bleeker, 1863 Garra chakpiensis Nebeshwar & Vishwanath, 2015Garra chathensis Ezung, Shangningam & Pankaj, 2020Garra chaudhurii Hora, 1921Garra chebera Habteselassie, Mikschi, Ahnelt & Waidbacher, 2010Garra chindwinensis Premananda, Kosygin, Saidullah, 2017Garra chingaiensis Abonmai, Linthoingambi, Ngangbam, Thoidingjam & K. R. Singh, 2023Garra chivaensis Moyon & L. Arunkumar, 2020Garra clavirostris Roni, Sarbojit & Vishwanath, 2017Garra compressa Kosygin & Vishwanath, 1998Garra congoensis Poll, 1959 Garra cornigera Shangningam & Vishwanath, 2015 Garra culiciphaga Garra cyclostomata Đ. Y. Mai, 1978Garra cyrano Kottelat, 2000Garra dampaensis Lalronunga, Lalnuntluanga & Lalramliana, 2013Garra deccanensis Jadhav, Karuthapandi, Shangningam, Jaiswal & Shankar, 2022Garra dembecha Getahun & Stiassny, 2007Garra dembeensis Garra dengba S.-Q. Deng, L. Cao & E. Zhang, 2018Garra dohjei Marngar, Mawlong & Lokeshwor, 2024Garra dulongensis Garra dunsirei Banister, 1987Garra duobarbis Getahun & Stiassny, 2007Garra elegans Garra elongata Vishwanath & Kosygin, 2000Garra emarginata Kurup & Radhakrishnan, 2011Garra ethelwynnae Menon, 1958Garra festai Garra flavatra S. O. Kullander & F. Fang, 2004Garra fluviatilis Kangrang, Thoni, Mayden & Beamish, 2016Garra fuliginosa Fowler, 1934Garra gallagheri Krupp, 1988 Garra geba Getahun & Stiassny, 2007Garra ghorensis Krupp, 1982 Garra gotyla Garra gracilis Garra gravelyi Garra gymnothorax Berg, 1949 Garra hexagonarostris Nebeshwar, Rameshori & Prabina, 2024Garra hindii Garra hormuzensis Zamani-Faradonbe, E. Zhang & Keivany, 2021Garra hughi Silas, 1955 Garra ignestii Garra imberbis Garra irangensis Premananda, 2023Garra jaldhakaensis Kosygin, Shangningam, Singh & U. Das, 2021Garra jamila Moritz, Straube & Neumann, 2019Garra jenkinsonianum Hora, 1921Garra jerdoni F. Day, 1867Garra jordanica Hamidan, Geiger & Freyhof, 2014Garra joshuai Garra kalakadensis Rema Devi, 1993Garra kalpangi Nebeshwar, Bagra & D. N. Das, 2012Garra kangrae Prashad, 1919Garra kemali Garra kempi Hora, 1921Garra khawbungi Arunachalam, Nandagopal & Mayden, 2014Garra kimini Arunachalam, Nandagopal & Mayden, 2014Garra klatti Garra koladynensis Nebeshwar & Vishwanath, 2017Garra laishrami Surachita, Chowdhury & Palita, 2023Garra lamta Garra lancrenonensis Blache & Miton, 1960Garra langlungensis Ezung, Shangningam & Pankaj, 2021Garra lautior Banister, 1987Garra lissorhynchus Garra litanensis Vishwanath, 1993Garra longchuanensis Q. Yu, X. Z. Wang, H. Xiong & S. P. He, 2016Garra longipinnis Banister & M. A. Clarke, 1977Garra lorestanensis Mousavi-Sabet & Eagderi, 2016Garra lungongza Ngangbam & Lithoingambi, 2023Garra magnacavus Shangningam, Kosygin & Sinha, 2019Garra magnidiscus Tamang, 2013Garra makiensis Garra mamshuqa Krupp, 1983Garra manipurensis Vishwanath & Sarojnalini, 1988Garra matensis Nebeshwar & Vishwanath, 2017Garra mcclellandi Garra menderesensis Garra menoni Rema Devi & T. J. Indra, 1984Garra meymehensis Zamani-Faradonbe, Keivany, Dorafshan & E. Zhang, 2021Garra mini Rahman, Mollah, Norén & S. O. Kullander, 2016Garra minibarbata L. Cao, Nyingi, Bart & E. Zhang 2023Garra minimus Arunachalam, Nandagopal & Mayden, 2013Garra mirofrontis X. L. Chu & G. H. Cui, 1987Garra mlapparaensis Kurup & Radhakrishnan, 2011Garra mondica Sayyadzadeh, Esmaeili & Freyhof, 2015Garra montisalsi Hora, 1921Garra motuoensis Z. Gong, Freyhof, Jian [Wang (ichthyologist)|J. Wang], M. Liu, F. Liu, P. C. Lin, Y. Jiang & H. Z. Liu 2018Garra moyonkhulleni Moyon & Arunkumar, 2018Garra mullya Garra naganensis Hora, 1921Garra nambulica Vishwanath & Joyshree, 2005r.Garra namyaensis Shangningam & Vishwanath, 2012Garra nana Garra napata Moritz, Straube & Neumann, 2019Garra nasuta Garra nepalensis Rayamajhi & Arunachalam, 2017Garra nethravathiensis Arunachalam & Nandagopal, 2014Garra ngatangkha Arunkumar & Moyon, 2019Garra ngopi Tenali, Chandran, R. K. Singh & Sarkar, 2024Garra nigricauda Arunachalam, Nandagopal & Mayden, 2013Garra nigricollis S. O. Kullander & F. Fang, 2004Garra notata Garra nudiventris Garra orientalis Nichols, 1925Garra ornata Garra orontesi Bayçelebi, Kaya, Turan & Freyhof, 2021Garra palaniensis Garra palaruvica Arunachalam, Raja, Nandagopal & Mayden, 2013Garra panitvongi Tangjitjaroen, Z. S. Randall, Tongnunui, Boyd & Page, 2023Garra paralissorhynchus Vishwanath & K. Shanta Devi, 2005Garra parastenorhynchus Thoni, Gurung & Mayden, 2016Garra paratrilobata Roni, Chinglemba, Rameshori & Vishwanath, 2019Garra periyarensis K. C. Gopi, 2001Garra persica Berg, 1914Garra phewakholaensis Limbu, Rajbanshi & Raza 2024Garra phillipsi Deraniyagala, 1933 Garra platycephala Narayan Rao, 1920Garra poecilura S. O. Kullander & F. Fang 2004Garra poilanei Petit & T. L. Tchang, 1933Garra prashadi Hora, 1921Garra propulvinus S. O. Kullander & F. Fang 2004Garra qiaojiensis H. W. Wu & Yao, 1977Garra quadratirostris Nebeshwar & Vishwanath, 2013Garra quadrimaculata Garra quangtriensis V. H. Nguyễn, T.H.N. Vũ & T. D. P. Nguyễn, 2015Garra rakhinica S. O. Kullander & F. Fang 2004Garra ranganensis Tamang, Sinha, Abujam & Kumar, 2019Garra regressa Getahun & Stiassny, 2007Garra rezai Mousavi-Sabet, Eagderi, Saemi-Komsari, Kaya & Freyhof, 2022Garra rhynchota Koller, 1926Garra robertsi Thoni & Mayden, 2015Garra roseae Mousavi-Sabet, Saemi-Komsari, Doadrio & Freyhof, 2019Garra rossica Garra rotundinasus E. Zhang, 2006Garra rufa Garra rupecula Garra sahilia Krupp, 1983Garra salweenica Hora & Mukerji, 1934Garra sannarensis Moritz, Straube & Neumann, 2019Garra sauvagei Garra shamal Kirchner, Kruckenhauser, Pichler, Borkenhagen & Freyhof, 2020Garra sharq Kirchner, Kruckenhauser, Pichler, Borkenhagen & Freyhof, 2020Garra simbalbaraensis Rath, Shangningam & Kosygin, 2019Garra sindhi Lyon, Geiger & Freyhof, 2016.Garra songbangensis V. H. Nguyễn, T.H.N. Vũ & T. D. P. Nguyễn, 2015Garra spilota S. O. Kullander & F. Fang, 2004Garra stenorhynchus Jerdon, 1849Garra substrictorostris Roni & Vishwanath, 2018Garra surendranathanii C. P. Shaji, L. K. Arun & P. S. Easa, 1996Garra surgifrons W. Zhou & C. Sun, 2018Garra surinbinnani Page, Ray, Tongnunui, Boyd & Randall, 2019Garra tamangi Gurumayum & Kosygin, 2016Garra tana Getahun & Stiassny, 2007 Garra tashanensis Mousavi-Sabet, Vatandoust, Fatemi & Eagderi, 2016Garra tengchongensis E. Zhang & Y. Y. Chen, 2002Garra tezuensis Thoidingjam, Ngangbam, Linthoingambi & Kh. R. Singh, 2023Garra tiam Zamani-Faradonbe, Keivany, Dorafshan & E. Zhang, 2021Garra tibanica Trewavas, 1941Garra tibetana G. Zheng; S. Q. Deng; J. Wang; H. Z. Lui, 2018Garra trewavasae Monod, 1950Garra triangularis Shangningam, Rath & Kosygin, 2021Garra trilobata Shangningam & Vishwanath, 2015Garra turcica Karaman 1971Garra typhlops Garra ukhrulensis Nebeshwar & Vishwanath, 2015Garra variabilis Garra vinciguerrae Garra waensis Lothongkham, Arbsuwan & Musikasinthorn, 2014Garra wanae Garra waterloti Garra widdowsoni Garra yajiangensis Gong, Freyhof, Wang, Liu, Liu, Lin, Jiang & Liu, 2018Garra zubzaensis Tenali, Shangningam, Bhattacharjee, Patel & Kosygin, 2024