Gymnotus

Gymnotus
	Gymnotus sp.
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Actinopterygii
Order:	Gymnotiformes
Family:	Gymnotidae
Genus:	Gymnotus; Linnaeus, 1758
Type species
	Gymnotus carapo; Linnaeus, 1758

Gymnotus is a genus of Neotropical freshwater fish in the family Gymnotidae found widely in South America, Central America and southern Mexico (36th parallel south to 18th parallel north).^[1] The greatest species richness is found in the Amazon basin.^[2] They are sometimes referred to by the English name banded knifefish,^[1] although this typically is reserved for the most widespread species, G. carapo.^[3]^[4] Overall Gymnotus is the most widespread genus in the order Gymnotiformes.^[5]

Although not commonly eaten by humans, some members of this genus are used locally as fishing bait,^[6] and occasionally kept in aquariums.^[7]

Habitat

Gymnotus occur in virtually any freshwater habitat in their range, even those with little oxygen (survives by breathing air directly from the water surface), areas affected by pollution,^[4] and for a period on land if their aquatic habitat dries out.^[8] Large species tend to live near floating vegetation along the edges of large rivers or floodplains, while smaller tend to live among leaf-litter or near banks of small streams.^[9] The genus includes both widespread and common species that occur in many different habitat types,^[4] and more restricted and rare species that occur in fewer habitats.^[10] There are species that remain in the same habitat throughout their lives, while others breed in specific habitats and spend the rest of their time elsewhere.^[11] At least as many as five species of Gymnotus may occur together in the same region and habitat.^[11]

Behavior

Gymnotus species are nocturnal and mainly feed on aquatic insects, crustaceans, small fish and other small animals,^[9] but may also take plant material.^[8] Being electric fish, they generate weak electric fields used for navigation, finding prey and communicating with other individuals of their species.^[8]^[12]^[13] At least some species are highly territorial and will react aggressively if detecting the electric field of another individual of their species, especially between conspecific males.^[13]^[14] The electric signal is species specific, and tends to differ between males and females.^[1] However, Gymnotus are not able to generate a strong electric field that can be used for incapacitating prey or enemies, like the related electric eel.^[9]

Nothing is known about the breeding behavior of most members of this genus, but in two species, G. carapo and G. mamiraua, males make a "nest" (a depression in the bottom in the former species and within vegetation in floating meadows in the latter) and guard the young. Additionally, males of at least G. carapo will mouthbrood.^[14]

Appearance

Gymnotus are generally brownish with a banded pattern, but this can also be more mottled or spotted in some species. Small scales are always present on these fish. The mouth is superior, meaning it is turned upwards. The anal fin terminates at a point near the tip of the tail.^[5]^[9] Like other Neotropical knifefish, they often lose their tail due to attacks by predators or aggressive encounters with conspecifics, but they are able to regenerate it.^[14] The largest Gymnotus are up to 100 cm (3.3 ft) in total length.^[5]^[9] Most species reach less than one-third that size and the smallest only around 10 cm (4 in) long.^[3]^[4]^[9]

Species

There are currently 46 recognized species divided into six subgenera in Gymnotus:^[3]^[15]^[16]

Subgenus Gymnotus (Gymnotus)

Gymnotus arapaima J. S. Albert & Crampton, 2001
Gymnotus ardilai Maldonado-Ocampo & J. S. Albert, 2004
Gymnotus bahianus Campos-da-Paz & W. J. E. M. Costa, 1996
Gymnotus capanema Milhomem, Crampton, Pieczarka, Shetka, D. S. Silva & Nagamachi, 2012^[17]
Gymnotus carapo Linnaeus, 1758 (Banded knifefish)
Gymnotus chaviro Maxime & J. S. Albert, 2009
Gymnotus chimarrao Cognato, Richer-de-Forges, J. S. Albert & Crampton, 2008^[18]
Gymnotus choco J. S. Albert, Crampton & Maldonado-Ocampo, 2003
Gymnotus cuia Craig, Malabarba, Crampton & J. S. Albert, 2018^[19]
Gymnotus curupira Crampton, Thorsen & J. S. Albert, 2005
Gymnotus darwini Campos-da-Paz & de Santana, 2019^[20]
Gymnotus diamantinensis Campos-da-Paz, 2002
Gymnotus eyra Craig, Correa-Roldán, Ortega, Crampton & J. S. Albert, 2018^[1]
Gymnotus interruptus Rangel-Pereira, 2012^[21]
Gymnotus mamiraua J. S. Albert & Crampton, 2001
Gymnotus obscurus Crampton, Thorsen & J. S. Albert, 2005
Gymnotus omarorum Richer-de-Forges, Crampton & J. S. Albert, 2009
Gymnotus pantanal Fernandes-Matioli, J. S. Albert, Daniel-Silva, C. E. Lopes, Crampton & Almeida-Toledo, 2005
Gymnotus riberalta Craig, Correa-Roldán, Ortega, Crampton & J. S. Albert, 2018^[1]
Gymnotus sylvius J. S. Albert & Fernandes-Matioli, 1999
Gymnotus ucamara Crampton, Lovejoy & J. S. Albert, 2003
Gymnotus varzea Crampton, Thorsen & J. S. Albert, 2005

Subgenus Gymnotus (Lamontiana)

Gymnotus anguillaris Hoedeman, 1962
Gymnotus cataniapo Mago-Leccia, 1994
Gymnotus coatesi La Monte, 1935
Gymnotus coropinae Hoedeman, 1962
Gymnotus pedanopterus Mago-Leccia, 1994
Gymnotus tiquie Maxime, F. C. T. Lima & J. S. Albert, 2011

Subgenus Gymnotus (Pantherus)

Gymnotus capitimaculatus Rangel-Pereira, 2014^[12]
Gymnotus pantherinus (Steindachner, 1908)
Gymnotus refugio Giora & L. R. Malabarba, 2016^[10]

Subgenus Gymnotus (Tijax)

Gymnotus cylindricus La Monte, 1935
Gymnotus maculosus J. S. Albert & R. R. Miller, 1995 (Spotted knifefish)
Gymnotus panamensis J. S. Albert & Crampton, 2003

Subgenus Gymnotus (Tigre)

Gymnotus esmeraldas J. S. Albert & Crampton, 2003
Gymnotus henni J. S. Albert, Crampton & Maldonado-Ocampo, 2003
Gymnotus inaequilabiatus (Valenciennes, 1839)^[22]
Gymnotus paraguensis J. S. Albert & Crampton, 2003
Gymnotus tigre J. S. Albert & Crampton, 2003

Subgenus Gymnotus (Tigrinus)

Gymnotus javari J. S. Albert, Crampton & Hagedorn, 2003
Gymnotus jonasi J. S. Albert & Crampton, 2001
Gymnotus melanopleura J. S. Albert & Crampton, 2001
Gymnotus onca J. S. Albert & Crampton, 2001
Gymnotus stenoleucus Mago-Leccia, 1994

References

^ ^a ^b ^c ^d ^e Craig, J.M.; V. Correa-Roldán; H. Ortega; W.G.R. Crampton; J.S. Albert (2018). "Revision of Gymnotus (Gymnotiformes: Gymnotidae) from the Upper Madeira Basin of Bolivia and Peru, with descriptions of two new species". Zootaxa. 4413 (1): 111–132. doi:10.11646/zootaxa.4413.1.3. PMID 29690122.
^ Albert, J.S.; R.R. Miller (1995). "Gymnotus maculosus, a new species of electric fish (Chordata: Teleostei: Gymnotoidei) from Middle America, with a key to species of Gymnotus". Proceedings of the Biological Society of Washington. 108 (4): 662–678.
^ ^a ^b ^c Froese, Rainer and Pauly, Daniel, eds. (2018). Species of Gymnotus in FishBase. May 2018 version.
^ ^a ^b ^c ^d Craig, J.M.; W.G.R. Crampton; J.S. Albert (2017). "Revision of the polytypic electric fish Gymnotus carapo (Gymnotiformes, Teleostei), with descriptions of seven subspecies". Zootaxa. 4318 (3): 401–438. doi:10.11646/zootaxa.4318.3.1.
^ ^a ^b ^c Nelson, J.S. (2006). Fishes of the World. John Wiley & Sons, Inc. ISBN 0-471-25031-7.
^ Medeiros, M. (translator) (12 September 2017). "Research identifies fish species used as live bait in Pantanal". Embrapa. Retrieved 11 May 2018. {{cite web}}: |author= has generic name (help)
^ "Gymnotus carapo". SeriouslyFish. Retrieved 11 May 2018.
^ ^a ^b ^c Froese, Rainer; Pauly, Daniel (eds.) (2018). "Gymnotus carapo" in FishBase. May 2018 version.
^ ^a ^b ^c ^d ^e ^f van der Sleen, P.; J.S. Albert, eds. (2017). Field Guide to the Fishes of the Amazon, Orinoco, and Guianas. Princeton University Press. pp. 332–334. ISBN 978-0-691-17074-9.
^ ^a ^b Giora, J.; L.R. Malabarba (2016). "Gymnotus refugio, a new and endangered species of electric fish of the Gymnotus pantherinus species-group from southern Brazil (Gymnotiformes: Gymnotidae)". Zootaxa. 4066 (5): 581–590. doi:10.11646/zootaxa.4066.5.6. PMID 27395856.
^ ^a ^b Albert, J.S.; Crampton, W.G.R. (2001). "Five new species of Gymnotus (Teleostei: Gymnotiformes) from an Upper Amazonian floodplain, with descriptions of electric organ discharges and ecology". Ichthyological Exploration of Freshwaters. 12 (3): 241–266.
^ ^a ^b Rangel-Pereira, G.S. (2014). "Gymnotus capitimaculatus, a new species of electric fish from rio Jucuruçu basin, northeastern Brazil (Ostariophysi: Gymnotiformes: Gymnotidae)". Vertebrate Zoology. 64 (2): 169–175. doi:10.3897/vz.64.e31474. S2CID 55651223.
^ ^a ^b Davis, E.A.; C.D. Hopkins (1988). "Behavioural analysis of electric signal localization in the electric fish, Gymnotus carapo (Gymnotiformes)". Animal Behaviour. 36 (6): 1658–1671. doi:10.1016/S0003-3472(88)80106-4. S2CID 53145791.
^ ^a ^b ^c Crampton, W.G.R.; C.D. Hopkins (2005). "Nesting and Paternal Care in the Weakly Electric Fish Gymnotus (Gymnotiformes: Gymnotidae) with Descriptions of Larval and Adult Electric Organ Discharges of Two Species". Copeia. 2005 (1): 48–60. doi:10.1643/CI-04-056R1. S2CID 16278706.
^ Eschmeyer, W.N.; R. Fricke; R. van der Laan (11 May 2018). "Catalog of Fishes". California Academy of Sciences. Retrieved 11 May 2018.
^ Craig, Jack M.; Kim, Lesley Y.; Tagliacollo, Victor A.; Albert, James S. (2019-11-07). "Phylogenetic revision of Gymnotidae (Teleostei: Gymnotiformes), with descriptions of six subgenera". PLOS ONE. 14 (11): e0224599. Bibcode:2019PLoSO..1424599C. doi:10.1371/journal.pone.0224599. ISSN 1932-6203. PMC 6837465. PMID 31697735.
^ Milhomem S.S.R.; Crampton W.G.R.; Pieczarka J.C.; Shetka G.H.; Silva D.S.; Nagamachi C.Y. (2012). "Gymnotus capanema, a new species of electric knife fish (Gymnotiformes, Gymnotidae) from eastern Amazonia, with comments on an unusual karyotype". Journal of Fish Biology. 80 (4): 802–815. doi:10.1111/j.1095-8649.2012.03219.x. PMID 22471800.
^ Cognato, D.P.; Richer-de-Forges, M.M.; Albert, J.S.; Crampton, W.G.R. (2008). "Gymnotus chimarrao, a new species of electric fish (Gymnotiformes: Gymnotidae) from Southern Brazil". Ichthyologial Exploration of Freshwaters. 18 (4): 375–382.
^ Craig, J.M.; L.R. Malabarba; W.G.R. Crampton; J.S. Albert (2018). "Revision of banded knifefishes of the Gymnotus carapo and G. tigre clades (Gymnotidae Gymnotiformes) from the Southern Neotropics". Zootaxa. 4379 (1): 47–73. doi:10.11646/zootaxa.4379.1.3. PMID 29689973.
^ Campos-da-Paz, R.; C.D. de Santana (2019). "A New Species of the Electric Knifefish Gymnotus Linnaeus (Gymnotiformes: Gymnotidae) from Northeastern Brazil". Copeia. 107 (1): 144–151. doi:10.1643/CI-18-141. S2CID 92304042.
^ Rangel-Pereira (2012). "Gymnotus interruptus, a new species of electric fish from the Rio de Contas basin, Bahia, Brazil (Teleostei: Gymnotiformes: Gymnotidae)" (PDF). Vertebrate Zoology. 62 (3): 363–370. doi:10.3897/vz.62.e31396. S2CID 85709855. Archived from the original (PDF) on 2018-05-07. Retrieved 2012-12-12.
^ Maxime E.L.; Albert J.S. (2014). "Redescription of the tuvirão, Gymnotus inaequilabiatus Valenciennes, 1839, using high-resolution X-ray computed tomography". Copeia. 2014 (3): 462–472. doi:10.1643/ci-13-054. S2CID 86328107.

[Craig2018a-1] Craig, J.M.; V. Correa-Roldán; H. Ortega; W.G.R. Crampton; J.S. Albert (2018). "Revision of Gymnotus (Gymnotiformes: Gymnotidae) from the Upper Madeira Basin of Bolivia and Peru, with descriptions of two new species". Zootaxa. 4413 (1): 111–132. doi:10.11646/zootaxa.4413.1.3. PMID 29690122.

[Albert1995-2] Albert, J.S.; R.R. Miller (1995). "Gymnotus maculosus, a new species of electric fish (Chordata: Teleostei: Gymnotoidei) from Middle America, with a key to species of Gymnotus". Proceedings of the Biological Society of Washington. 108 (4): 662–678.

[fishbase-3] Froese, Rainer and Pauly, Daniel, eds. (2018). Species of Gymnotus in FishBase. May 2018 version.

[Craig2017-4] Craig, J.M.; W.G.R. Crampton; J.S. Albert (2017). "Revision of the polytypic electric fish Gymnotus carapo (Gymnotiformes, Teleostei), with descriptions of seven subspecies". Zootaxa. 4318 (3): 401–438. doi:10.11646/zootaxa.4318.3.1.

[Nelson2006-5] Nelson, J.S. (2006). Fishes of the World. John Wiley & Sons, Inc. ISBN 0-471-25031-7.

[6] Medeiros, M. (translator) (12 September 2017). "Research identifies fish species used as live bait in Pantanal". Embrapa. Retrieved 11 May 2018. {{cite web}}: |author= has generic name (help)

[7] "Gymnotus carapo". SeriouslyFish. Retrieved 11 May 2018.

[FishbaseCarapo-8] Froese, Rainer; Pauly, Daniel (eds.) (2018). "Gymnotus carapo" in FishBase. May 2018 version.

[Sleen2017-9] ^ ^a ^b ^c ^d ^e ^f van der Sleen, P.; J.S. Albert, eds. (2017). Field Guide to the Fishes of the Amazon, Orinoco, and Guianas. Princeton University Press. pp. 332–334. ISBN 978-0-691-17074-9.

[Giora2016-10] Giora, J.; L.R. Malabarba (2016). "Gymnotus refugio, a new and endangered species of electric fish of the Gymnotus pantherinus species-group from southern Brazil (Gymnotiformes: Gymnotidae)". Zootaxa. 4066 (5): 581–590. doi:10.11646/zootaxa.4066.5.6. PMID 27395856.

[Albert2001-11] Albert, J.S.; Crampton, W.G.R. (2001). "Five new species of Gymnotus (Teleostei: Gymnotiformes) from an Upper Amazonian floodplain, with descriptions of electric organ discharges and ecology". Ichthyological Exploration of Freshwaters. 12 (3): 241–266.

[Rangel2014-12] Rangel-Pereira, G.S. (2014). "Gymnotus capitimaculatus, a new species of electric fish from rio Jucuruçu basin, northeastern Brazil (Ostariophysi: Gymnotiformes: Gymnotidae)". Vertebrate Zoology. 64 (2): 169–175. doi:10.3897/vz.64.e31474. S2CID 55651223.

[Davis1988-13] Davis, E.A.; C.D. Hopkins (1988). "Behavioural analysis of electric signal localization in the electric fish, Gymnotus carapo (Gymnotiformes)". Animal Behaviour. 36 (6): 1658–1671. doi:10.1016/S0003-3472(88)80106-4. S2CID 53145791.

[Crampton2005-14] Crampton, W.G.R.; C.D. Hopkins (2005). "Nesting and Paternal Care in the Weakly Electric Fish Gymnotus (Gymnotiformes: Gymnotidae) with Descriptions of Larval and Adult Electric Organ Discharges of Two Species". Copeia. 2005 (1): 48–60. doi:10.1643/CI-04-056R1. S2CID 16278706.

[CoF-15] Eschmeyer, W.N.; R. Fricke; R. van der Laan (11 May 2018). "Catalog of Fishes". California Academy of Sciences. Retrieved 11 May 2018.

[16] Craig, Jack M.; Kim, Lesley Y.; Tagliacollo, Victor A.; Albert, James S. (2019-11-07). "Phylogenetic revision of Gymnotidae (Teleostei: Gymnotiformes), with descriptions of six subgenera". PLOS ONE. 14 (11): e0224599. Bibcode:2019PLoSO..1424599C. doi:10.1371/journal.pone.0224599. ISSN 1932-6203. PMC 6837465. PMID 31697735.

[Milhomem2012-17] Milhomem S.S.R.; Crampton W.G.R.; Pieczarka J.C.; Shetka G.H.; Silva D.S.; Nagamachi C.Y. (2012). "Gymnotus capanema, a new species of electric knife fish (Gymnotiformes, Gymnotidae) from eastern Amazonia, with comments on an unusual karyotype". Journal of Fish Biology. 80 (4): 802–815. doi:10.1111/j.1095-8649.2012.03219.x. PMID 22471800.

[cognato2007-18] Cognato, D.P.; Richer-de-Forges, M.M.; Albert, J.S.; Crampton, W.G.R. (2008). "Gymnotus chimarrao, a new species of electric fish (Gymnotiformes: Gymnotidae) from Southern Brazil". Ichthyologial Exploration of Freshwaters. 18 (4): 375–382.

[Craig2018b-19] Craig, J.M.; L.R. Malabarba; W.G.R. Crampton; J.S. Albert (2018). "Revision of banded knifefishes of the Gymnotus carapo and G. tigre clades (Gymnotidae Gymnotiformes) from the Southern Neotropics". Zootaxa. 4379 (1): 47–73. doi:10.11646/zootaxa.4379.1.3. PMID 29689973.

[CamposDaPaz2019-20] Campos-da-Paz, R.; C.D. de Santana (2019). "A New Species of the Electric Knifefish Gymnotus Linnaeus (Gymnotiformes: Gymnotidae) from Northeastern Brazil". Copeia. 107 (1): 144–151. doi:10.1643/CI-18-141. S2CID 92304042.

[Rangel-Pereira2012-21] Rangel-Pereira (2012). "Gymnotus interruptus, a new species of electric fish from the Rio de Contas basin, Bahia, Brazil (Teleostei: Gymnotiformes: Gymnotidae)" (PDF). Vertebrate Zoology. 62 (3): 363–370. doi:10.3897/vz.62.e31396. S2CID 85709855. Archived from the original (PDF) on 2018-05-07. Retrieved 2012-12-12.

[Maxime2014-22] Maxime E.L.; Albert J.S. (2014). "Redescription of the tuvirão, Gymnotus inaequilabiatus Valenciennes, 1839, using high-resolution X-ray computed tomography". Copeia. 2014 (3): 462–472. doi:10.1643/ci-13-054. S2CID 86328107.

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Gymnotus

Habitat

Behavior

Appearance

Species

References

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