Garra Explained
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).
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); though it didn't lead to an act of him to designate a type species by the time. However, as no other garras except the newly discovered G. lamta were known to science in 1822, this was designated as the type species by Pieter Bleeker 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 is a close relative.[1]
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 2,4,5–5,4,2), the dorsal fin has 10-11 rays and starts slightly anterior to the pelvic fins, while the anal fin starts well behind the pelvic fins and has 8-9 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.[2] 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.
Species
These are the currently recognized species in this genus:
- Garra abhoyai Hora, 1921
- Garra aethiopica (Pellegrin, 1927)
- Garra allostoma T. R. Roberts, 1990
- Garra alticaputus Arunachalam, Nandagopal & Mayden, 2013 [3]
- Garra amirhosseini Esmaeili, Sayyadzadeh, Coad & Eagderi, 2016 [4]
- Garra andruzzii (Vinciguerra, 1924) (Somalian cavefish)
- Garra annandalei Hora, 1921
- Garra apogon (Norman, 1925)
- Garra arunachalami Johnson & Soranam, 2001
- Garra arunachalensis Nebeshwar & Vishwanath, 2013 [5]
- Garra arupi Nebeshwar, Vishwanath & D. N. Das, 2009
- Garra barreimiae Fowler & Steinitz, 1956
- Garra bicornuta Narayan Rao, 1920
- Garra biloborostris Roni & Vishwanath, 2017
- Garra bimaculacauda Thoni, Gurung & Mayden, 2016 [6]
- Garra binduensis Das, Kosygin & Panigrahi, 2016
- Garra birostris Nebeshwar & Vishwanath, 2013 [5]
- Garra bisangularis Chen, Wu, & Xiao, 2010
- Garra bispinosa E. Zhang, 2005
- Garra blanfordii (Boulenger, 1901)
- Garra borneensis (Vaillant, 1902)
- Garra bourreti (Pellegrin, 1928)
- Garra buettikeri Krupp, 1983
- Garra cambodgiensis (Tirant, 1883) (Cambodian logsucker)
- Garra caudomaculata (Battalgil, 1942) (Antakya minnow)
- Garra centrala V.H. Nguyen, T.H.N. Vu & T.D.P. Nguyen, 2015
- Garra ceylonensis Bleeker, 1863 (Ceylon logsucker)
- Garra chakpiensis Nebeshwar & Vishwanath, 2015 [7]
- Garra chathensis Ezung, Shangningam & Pankaj, 2020
- Garra chebera Habteselassie, Mikschi, Ahnelt & Waidbacher, 2010
- Garra chindwinensis Premananda, Kosygin, Saidullah, 2017[8]
- Garra compressus Kosygin & Vishwanath, 1998
- Garra congoensis Poll, 1959
- Garra cornigera Shangningam & Vishwanath, 2015 [9]
- Garra cryptonema (G. H. Cui & Z. Y. Li, 1984)
- Garra culiciphaga (Pellegrin, 1927) [10] (Red stripe barb)
- Garra cyclostomata Đ. Y. Mai, 1978
- Garra cyrano Kottelat, 2000
- Garra dampaensis Lalronunga, Lalnuntluanga & Lalramliana, 2013 [11]
- Garra dembecha Getahun & Stiassny, 2007
- Garra dembeensis (Rüppell, 1835) (Dembea stone lapper)
- Garra dulongensis (Chen, Pan, Kong & Yang, 2006)
- Garra dunsirei Banister, 1987
- Garra duobarbis Getahun & Stiassny, 2007
- Garra elegans (Günther, 1868) [12]
- Garra elongata Vishwanath & Kosygin, 2000
- Garra emarginata Kurup & Radhakrishnan, 2011
- Garra ethelwynnae Menon, 1958
- Garra fasciacauda Fowler, 1937
- Garra festai (Tortonese, 1939)
- Garra fisheri (Fowler, 1937)
- Garra flavatra S. O. Kullander & F. Fang, 2004
- Garra fluviatilis Kangrang, Thoni, Mayden & Beamish, 2016 [13]
- Garra fuliginosa Fowler, 1934
- Garra gallagheri Krupp, 1988 (Black Garra)
- Garra geba Getahun & Stiassny, 2007
- Garra ghorensis Krupp, 1982
- Garra gotyla (J. E. Gray, 1830) (Sucker head)
- Garra gracilis (Pellegrin & Chevey, 1936)
- Garra gravelyi (Annandale, 1919)
- Garra gymnothorax Berg, 1949 (Chest scaleless garra)
- Garra hainanensis Y. R. Chen & C. Y. Zheng, 1983
- Garra hindii (Boulenger, 1905)
- Garra hughi Silas, 1955
- Garra ignestii (Gianferrari, 1925)
- Garra imbarbatus (Nguyen, 2001)
- Garra imberba Garman, 1912
- Garra imberbis (Vinciguerra, 1890)
- Garra incisorbis L. P. Zheng, J. X. Yang & X. Y. Chen, 2016 [14]
- Garra jaldhakaensis, Kosygin, Bungdon Shangningam, Pratima Singh, Ujjal Das, 2021[15]
- Garra jerdoni F. Day, 1867
- Garra jordanica Hamidan, Geiger & Freyhof, 2014 [16]
- Garra kalakadensis Rema Devi, 1993
- Garra kalpangi Nebeshwar, Bagra & D. N. Das, 2012 [17]
- Garra kemali (Hankó, 1925) [10]
- Garra kempi Hora, 1921
- Garra khawbungi Arunachalam, Nandagopal & Mayden, 2014 [18]
- Garra kimini Arunachalam, Nandagopal & Mayden, 2013 [3]
- Garra lamta (F. Hamilton, 1822)
- Garra lancrenonensis Blache & Miton, 1960
- Garra lautior Banister, 1987
- Garra lissorhynchus (McClelland, 1842)
- Garra litanensis Vishwanath, 1993
- Garra longchuanensis Q. Yu, X. Z. Wang, H. Xiong & S. P. He, 2016 [19]
- Garra longipinnis Banister & M. A. Clarke, 1977
- Garra lorestanensis Mousavi-Sabet & Eagderi, 2016 [20]
- Garra magnidiscus Tamang, 2013 [21]
- Garra makiensis (Boulenger, 1904)
- Garra mamshuqa Krupp, 1983
- Garra manipurensis Vishwanath & Sarojnalini, 1988
- Garra mcclellandi (Jerdon, 1849)
- Garra menderesensis (Küçük, Bayçelebi, Güçlü & Gülle, 2015) [10]
- Garra menoni Rema Devi & T. J. Indra, 1984
- Garra micropulvinus W. Zhou, X. F. Pan & Kottelat, 2005
- Garra mini Rahman, Mollah, Norén & Kullander, 2016
- Garra minimus Arunachalam, Nandagopal & Mayden, 2013 [3]
- Garra mirofrontis X. L. Chu & G. H. Cui, 1987
- Garra mlapparaensis Kurup & Radhakrishnan, 2011 [22]
- Garra mondica Sayyadzadeh, Esmaeili & Freyhof, 2015 [23]
- Garra mullya (Sykes, 1839) (sucker fish)
- Garra naganensis Hora, 1921
- Garra nambulica Vishwanath & Joyshree, 2005
- Garra namyaensis Shangningam & Vishwanath, 2012 [24] [25]
- Garra nana (Heckel, 1843) [10]
- Garra nasuta (McClelland, 1838)
- Garra nethravathiensis Arunachalam & Nandagopal, 2014 [26]
- Garra nigricauda Arunachalam, Nandagopal & Mayden, 2013 [3]
- Garra nigricollis S. O. Kullander & F. Fang, 2004
- Garra nkhruletisis Nebeshwar & Vishwanath, 2015 [7]
- Garra notata (Blyth, 1860)
- Garra nujiangensis Z. M. Chen, S. Zhao & J. X. Yang, 2009
- Garra orientalis Nichols, 1925
- Garra ornata (Nichols & Griscom, 1917)
- Garra palaniensis Rema Devi & Menon, 1994
- Garra palaruvica Arunachalam, Raja, Nandagopal & Mayden, 2013
- Garra panitvongi Tangjitjaroen, Z. S. Randall, Tongnunui, Boyd & Page, 2023[27]
- Garra paralissorhynchus Vishwanath & K. Shanta Devi, 2005
- Garra parastenorhynchus Thoni, Gurung & Mayden, 2016 [6]
- Garra periyarensis K. C. Gopi, 2001
- Garra persica L. S. Berg, 1914
- Garra phillipsi Deraniyagala, 1933 (Phillips's garra)
- Garra platycephala Narayan Rao, 1920 [26]
- Garra poecilura S. O. Kullander & F. Fang, 2004
- Garra poilanei Petit & T. L. Tchang, 1933
- Garra propulvinus S. O. Kullander & F. Fang, 2004
- Garra qiaojiensis H. W. Wu & Yao, 1977
- Garra quadratirostris Nebeshwar & Vishwanath, 2013 [5]
- Garra quadrimaculata (Rüppell, 1835)
- Garra rakhinica S. O. Kullander & F. Fang, 2004
- Garra regressus Getahun & Stiassny, 2007
- Garra robertsi Thoni & Mayden, 2015 [28]
- Garra rossica (A. M. Nikolskii, 1900)
- Garra rotundinasus E. Zhang, 2006
- Garra rufa (Heckel, 1843) (Red garra)
- Garra rupecula (McClelland, 1839)
- Garra sahilia Krupp, 1983
- Garra salweenica Hora & Mukerji, 1934
- Garra sindhi Lyon, Geiger & Freyhof, 2016 [29]
- Garra smarti Krupp & Budd, 2009
- Garra spilota S. O. Kullander & F. Fang, 2004
- Garra stenorhynchus Jerdon, 1849
- Garra surendranathanii C. P. Shaji, L. K. Arun & P. S. Easa, 1996
- Garra surinbinnani Page, Ray, Tongnunui, Boyd & Randall, 2019
- Garra tamangi Gurumayum & Kosygin, 2016 [30]
- Garra tana Getahun & Stiassny, 2007
- Garra tangjitjaroeni
- Garra tashanensis Mousavi-Sabet, Vatandoust, Fatemi & Eagderi, 2016[31]
- Garra tengchongensis E. Zhang & Y. Y. Chen, 2002
- Garra theunensis Kottelat, 1998
- Garra tibanica Trewavas, 1941 [16]
- Garra trilobata Shangningam & Vishwanath, 2015 [9]
- Garra tyao Arunachalam, Nandagopal & Mayden, 2014 [18]
- Garra typhlops (Bruun & E. W. Kaiser, 1944)[32] (Iran cave barb)
- Garra ukhrulensis Nebeshwar & Vishwanath, 2015
- Garra variabilis (Heckel, 1843) (Variable garra)
- Garra vittatula S. O. Kullander & F. Fang, 2004
- Garra waensis Lothongkham, Arbsuwan & Musikasinthorn, 2014 [33]
- Garra wanae (Regan, 1914)
- Garra waterloti (Pellegrin, 1935)
- Garra widdowsoni (Trewavas, 1955) (Iraq blind barb)
- Garra yiliangensis H. W. Wu & Q. Z. Chen, 1977
Notes and References
- Stiassny, M.L.J. & Getahun, A. (2007): An overview of labeonin relationships and the phylogenetic placement of the Afro-Asian genus Garra Hamilton, 1922 (Teleostei: Cyprinidae), with the description of five new species of Garra from Ethiopia, and a key to all African species. Zoological Journal of the Linnean Society, 150 (1): 41–83.
- Romero, A., editor (2001). The Biology of Hypogean Fishes, p. 17. Developments in Environmental Biology of Fishes.
- Arunachalam, M., Nandagopal, S. & Mayden, R.L. (2013): Morphological diagnoses of Garra (Cypriniformes: Cyprinidae) from North-Eastern India with four new species description from Brahmaputra River. Journal of Fisheries and Aquaculture, 4 (3): 121-138.
- Esmaeili, H.R., Sayyadzadeh, G., Coad, B.W. & Eagderi, S. (2016): Review of the genus Garra Hamilton, 1822 in Iran with description of a new species: a morpho-molecular approach (Teleostei: Cyprinidae). Iranian Journal of Ichthyology, 3 (2): 82-121.
- Nebeshwar, K. & Vishwanath, W. (2013): Three new species of Garra (Pisces: Cyprinidae) from north-eastern India and redescription of G. gotyla. Ichthyological Exploration of Freshwaters, 24 (2): 97-120.
- Thoni, R.J., Gurung, D.B. & Mayden, R.L. (2016): A review of the genus Garra Hamilton 1822 of Bhutan, including the descriptions of two new species and three additional records (Cypriniformes: Cyprinidae). Zootaxa, 4169 (1): 115-132.
- Nebeshwar, K. & Vishwanath, W. (2015): Two new species of Garra (Pisces: Cyprinidae) from the Chindwin River basin in Manipur, India, with notes on some nominal Garra species of the Himalayan foothills. Ichthyological Exploration of Freshwaters, 25 (4): 305-321.
- http://recordsofzsi.com/index.php/zsoi/article/view/120968/84207| Nongthombam Premananda, Laishram Kosygin, Bano Saidullah. 2017. Garra chindwinensis, a New Species of Cyprinid Fish (Teleostei: Cypriniformes) from Manipur, Northeastern India
- Shangningam, B. & Vishwanath, W. (2015): Two new species of Garra from the Chindwin basin, India (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters, 26 (3): 263–272.
- Behrens-Chapuis, S., Herder, F., Esmaeili, H.R., Freyhof, J., Hamidan, N.A., Özuluğ, M., Šanda, R. & Geiger, M.F. (2015): Adding nuclear rhodopsin data where mitochondrial COI indicates discrepancies – can this marker help to explain conflicts in cyprinids? DNA Barcodes, 3 (1): 187-199.
- Lalronunga, S., Lalnuntluanga & Lalramliana (2013): Garra dampaensis, a new ray-finned fish species (Cypriniformes: Cyprinidae) from Mizoram, northeastern India. Journal of Threatened Taxa, 5 (9): 4368–4377.
- Freyhof, J. (2016): Redescription of Garra elegans (Günther, 1868), a poorly known species from the Tigris River drainage (Teleostei: Cyprinidae). Zootaxa, 4173 (5): 496-500.
- Kangrang, P., Thoni, R.J., Mayden, R.L. & Beamish, F.W.H. (2016): Garra fluviatilis, a new hillstream fish species (Cypriniformes: Cyprinidae) from the Kwai Noi River system, Mae Khlong basin, Thailand. Zootaxa, 4175 (4): 335-344.
- Zheng, L.-P., Yang, J.-X. & Chen, X.-Y. (2016): Garra incisorbis, a new species of labeonine from Pearl River basin in Guangxi, China (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters, 26 (4): 299-304.
- http://recordsofzsi.com/index.php/zsoi/article/view/158415/| Garra jaldhakaensis, a new cyprinid fish (Teleostei: Cyprinidae) from West Bengal, India
- Hamidan, N.A., Geiger, M.F. & Freyhof, J. (2014): Garra jordanica, a new species from the Dead Sea basin with remarks on the relationship of G. ghorensis, G. tibanica and G. rufa (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters, 25 (3): 223-236.
- Nebeshwar, K., Bagra, K. & Das, D.N. (2012): Garra kalpangi, a new cyprinid fish species (Pisces: Teleostei) from upper Brahmaputra basin in Arunachal Pradesh, India. Journal of Threatened Taxa, 4 (2): 2353–2362.
- Arunachalam, M., Nandagopal, S. & Mayden, R.L. (2014): Two new Species of Garra from Mizoram, India (Cypriniformes: Cyprinidae) and a General Comparative Analyses of Indian Garra. Species, 10 (24): 58-78.
- Yu, Q., Wang, X., Xiong, H. & He, S. (2016): Garra longchuanensis, a new cyprinid (Teleostei: Cypriniformes) from southern China. Zootaxa, 4126 (2): 295-300.
- Mousavi-Sabet, H. & Eagderi, S. (2016): Garra lorestanensis, a new cave fish from the Tigris River drainage with remarks on the subterranean fishes in Iran (Teleostei: Cyprinidae). FishTaxa, 1 (1): 45-54.
- Tamang, L. (2013): Garra magnidiscus, a new species of cyprinid fish (Teleostei: Cypriniformes) from Arunachal Pradesh, northeastern India. Ichthyological Exploration of Freshwaters, 24 (1): 31-40.
- Kurup, B.M. & Radhakrishnan, K.V. (2011): Two new cyprinid fishes under the genus Garra (Hamilton) from Kerala, southern India. Journal of the Bombay Natural History Society, 107 (3): 220-223.
- Sayyadzadeh, G., Esmaeili, H.R. & Freyhof, J. (2015): Garra mondica, a new species from the Mond River drainage with remarks on the genus Garra from the Persian Gulf basin in Iran (Teleostei: Cyprinidae). Zootaxa, 4048 (1): 75–89.
- Shangningam, B. & Vishwanath, W. (2012): A New Species of the Genus Garra Hamilton, 1822 from the Chindwin Basin of Manipur, India (Teleostei: Cyprinidae: Labeoninae). International Scholarly Research Network (ISRN Zoology), 2012: 1-6.
- Shangningam, B. & Vishwanath, W. (2012): Validation of Garra namyaensis Shangningam & Vishwanath, 2012 (Teleostei: Cyprinidae: Labeoninae). Ichthyological Exploration of Freshwaters, 23 (1): 10-10.
- Arunachalam, M. & Nandagopal, S. (2014): A New Species of the Genus Garra Hamilton, (Cypriniformes: Cyprinidae) from Nethravathi River, Western Ghats, India. Species, 10 (24): 43-57.
- [Weerapongse Tangjitjaroen|Tangjitjaroen]
- Thoni, R.J. & Mayden, R.L. (2015): Garra robertsi, a new cyprinid (Cypriniformes: Cyprinidae) fish species from Borneo. Zootaxa, 3985 (2): 284–290.
- Lyon, R.G., Geiger, M.F. & Freyhof, J. (2016): Garra sindhi, a new species from the Jebel Samhan Nature Reserve in Oman (Teleostei: Cyprinidae). Zootaxa, 4154 (1): 79-88.
- Gurumayum, S.D. & Kosygin, L. (2016): Garra tamangi, a new species of cyprinid fish (Teleostei: Cypriniformes) from Arunachal Pradesh, northeastern India. Species, 17 (55): 84-93.
- MOUSAVI-SABET, Hamed, et al. “Tashan Cave a New Cave Fish Locality for Iran; and Garra Tashanensis, a New Blind Species from the Tigris River Drainage (Teleostei: Cyprinidae).” FISHTAXA(2016) 1(3): 133-148, http://www.fishtaxa.com/index.php/ft/article/view/1-3-3.
- Farashi, A., Kaboli, M., Rezaei, H.R., Naghavi, M.R., Rahimian, H. & Coad, B.W. (2014): Reassessment of the taxonomic position of Iranocypris typhlops Bruun & Kaiser, 1944 (Actinopterygii, Cyprinidae). ZooKeys, 374: 69-77.
- Lothongkham, A., Arbsuwan, S. & Musikasinthorn, P. (2014): Garra waensis, a new cyprinid fish (Actinopterygii: Cypriniformes) from the Nan River basin of the Chao Phraya River system, northern Thailand. Zootaxa, 3790 (4): 543–554.