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{{Short description|Order of fish}} | |||
{{Other uses}} | |||
{{For|the deceptive online actions by one to another|Catfishing}} | |||
{{Taxobox | |||
{{About|the fish||Catfish (disambiguation)|}} | |||
| name = Catfish | |||
{{Redirect|Kaari|other uses|Kaari (disambiguation)}} | |||
| fossil_range = Late ] – present | |||
{{pp-move|small=yes}} | |||
{{Use dmy dates|date=August 2020}} | |||
{{Automatic taxobox | |||
| fossil_range = {{fossilrange|Campanian|Present|refs=<ref name=NT24>{{cite journal |last1=Near |first1=Thomas J |last2=Thacker |first2=Christine E |date=18 April 2024 |title=Phylogenetic classification of living and fossil ray-finned fishes (Actinopterygii) |journal=Bulletin of the Peabody Museum of Natural History |volume=65 |doi=10.3374/014.065.0101 |doi-access=free}}</ref>}}<small>Possible earlier ] records<ref name=Patterson1993/><ref name=Afrocascudo>{{Cite journal |last1=Brito |first1=P. M. |last2=Dutheil |first2=D. B. |last3=Gueriau |first3=P. |last4=Keith |first4=P. |last5=Carnevale |first5=G. |last6=Britto |first6=M. |last7=Meunier |first7=F. J. |last8=Khalloufi |first8=B. |last9=King |first9=A. |last10=de Amorim |first10=P. F. |last11=Costa |first11=W. J. E. M. |title=A saharan fossil and the dawn of Neotropical armoured catfishes in Gondwana |year=2024 |journal=Gondwana Research |volume=132 |pages=103–112 |doi=10.1016/j.gr.2024.04.008 |bibcode=2024GondR.132..103B }}</ref></small> | |||
| image = Ameiurus melas by Duane Raver.png | | image = Ameiurus melas by Duane Raver.png | ||
| image_caption = ] | | image_caption = ] | ||
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| image_upright = 1.1 | ||
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| taxon = Siluriformes | ||
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| authority = ], 1817 | ||
| subdivision_ranks = Families | |||
| superclassis = ] | |||
| subdivision_ref = <ref>{{FishBase order | order = Siluriformes| month = December | year = 2011}}</ref> | |||
| classis = ] | |||
| subdivision = '''Extant families:''' | |||
| subclassis = ] | |||
*]<ref name=Wang2016>{{cite journal|doi=10.1371/journal.pone.0145675|title=Phylogenetic Relationships of Five Asian Schilbid Genera Including ''Clupisoma'' (Siluriformes: Schilbeidae)|year=2016 |doi-access=free |last1=Wang |first1=Jing |last2=Lu |first2=Bin |last3=Zan |first3=Ruiguang |last4=Chai |first4=Jing |last5=Ma |first5=Wei |last6=Jin |first6=Wei |last7=Duan |first7=Rongyao |last8=Luo |first8=Jing |last9=Murphy |first9=Robert W. |last10=Xiao |first10=Heng |last11=Chen |first11=Ziming |journal=PLOS ONE |volume=11 |issue=1 |pages=e0145675 |pmid=26751688 |pmc=4713424 |bibcode=2016PLoSO..1145675W }}</ref> | |||
| infraclassis = ] | |||
*] | |||
| superordo = ] | |||
*] | |||
| ordo = '''Siluriformes''' | |||
*] | |||
| ordo_authority = ], 1817 | |||
*] | |||
| subdivision_ranks = Families<ref>{{FishBase order | order = Siluriformes| month = December | year = 2011}}</ref> | |||
*] | |||
| subdivision = | |||
*] | |||
- Extant Families -<br /> | |||
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*]<!-- Zoosystema 22 (4): 847-852 --> | |||
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]<br /><!-- Zoosystema 22 (4): 847-852 --> | |||
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* '']'': | |||
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**'']'' | |||
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'''Extinct family:''' | |||
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| type_species = '']'' | |||
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| type_species_authority = ], 1758 | |||
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'']''<br /> | |||
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- Extinct Family -<br /> | |||
] ] | |||
}} | }} | ||
'''Catfishes''' (] '''Siluriformes''') are a diverse group of ] ]. Named for their prominent ]s, which resemble a cat's ]s, catfish range in size and behavior from the heaviest and longest, the ] from Southeast Asia and the second longest, the ] of Eurasia, to ]s (species that eat dead material on the bottom), and even to a tiny ] species commonly called the ], ''Vandellia cirrhosa''. There are armour-plated types and also naked types, neither having scales. Despite their name, not all catfish have prominent barbels; members of the Siluriformes order are defined by features of the skull and ]. Catfish are of considerable commercial importance; many of the larger species are ] or fished for food. Many of the smaller species, particularly the genus '']'', are important in the ]. Catfish are ].<ref name="aworld"></ref><ref name="samer"></ref> | |||
==Distribution and habitat== | |||
Extant catfish species live inland or in coastal waters of every continent except ]. Catfish have inhabited all continents at one time or another.<ref name="Nelson"/> Catfish are most diverse in ] South America, Africa, and Asia.<ref name=tol/> More than half of all catfish species live in the Americas. They are the only ]s that have entered ] habitats in ], Australia, and ].<ref name="Bruton">{{cite journal|url=http://www.edpsciences.org/articles/alr/pdf/1996/05/alr96hs02.pdf?access=ok|format=PDF|title=Alternative life-history strategies of catfishes|last=Bruton|first=Michael N.|journal=Aquat. Living Resour.|year=1996|volume=9|pages=35–41|doi=10.1051/alr:1996040|accessdate=2009-06-22}}</ref> | |||
They are found in freshwater environments, though most inhabit shallow, running water.<ref name="Bruton"/> Representatives of at least eight families are ] (live underground) with three families that are also ] (inhabiting caves).<ref>{{cite journal|title=Morphological Adaptations of the Texas Blind Catfishes ''Trogloglanis pattersoni'' and ''Satan eurystomus'' (Siluriformes: Ictaluridae) to Their Underground Environment|first=Thomas G.|last=Langecker|coauthors=Longley, Glenn|journal=]|year=1993|pages=976–986|doi=10.2307/1447075|volume=1993|issue=4|jstor=1447075}}</ref><ref>{{cite journal|title=Mexican blindcats genus ''Prietella'' (Siluriformes: Ictaluridae): an overview of recent explorations|first=Dean A.|last=Hendrickson|coauthors=Krejca, Jean K.; Martinez, Juan Manuel Rodríguez|journal=Environmental Biology of Fishes|volume=62|pages=315–337|year=2001|doi=10.1023/A:1011808805094}}</ref> One such species is '']'', known to live underground in ] habitats.<ref name=pcisternarum>{{FishBase species|genus=Phreatobius|species=cisternarum|year=2007|month=Apr}}</ref> Numerous species from the families ] and ], and a few species from among the ] and ], are found in salt water.<ref>Monks N. (editor): ''Brackish Water Fishes'', TFH 2006, ISBN 0-7938-0564-3</ref><ref>Schäfer F: ''Brackish Water Fishes'', Aqualog 2005, ISBN 3-936027-82-X</ref> | |||
In the United States, catfish species may be known by a variety of slang names, just as ''mud cat'', ''polliwogs'', or ''chuckleheads''.<ref></ref> These nicknames are not standardized, so one area may call a Bullhead catfish by the nickname ''chucklehead'', while in another state or region, that nickname refers to the Blue catfish. | |||
==Physical characteristics== | |||
===External anatomy of catfish=== | |||
]''.]] | |||
Most catfish are ]. In general, they are ], which means that they will usually sink rather than float due to a reduced ] and a heavy, bony head.<ref name="Bruton"/> Catfish have a variety of body shapes, though most have a cylindrical body with a flattened ] to allow for benthic feeding.<ref name="Bruton"/> | |||
A flattened head allows for digging through the substrate as well as perhaps serving as a ]. Most have a mouth that can expand to a large size and contains no ] teeth; catfish generally feed through ] or gulping rather than biting and cutting prey.<ref name="Bruton"/> However, some families, notably ] and ], have a ] that allows them to fasten themselves to objects in fast-moving water. Catfish also have a ] reduced to a support for ]; this means that they are unable to protrude their mouths as other fish such as ].<ref name="Bruton"/> | |||
Catfish may have up to four pairs of barbels: nasal, maxillary (on each side of mouth), and two pairs of chin barbels, even though pairs of barbels may be absent depending on the species. Catfish also have ]s across their entire bodies, which means they "taste" anything they touch and "smell" any chemicals in the water. "In catfish, ] plays a primary role in the orientation and location of food".<ref>Atema, Jelle (1980) pp. 57–101. In: Bardach, JE ''Fish behavior and its use in the capture and culture of fishes', The WorldFish Center, ISBN 978-971-02-0003-0.</ref> Because their barbels and chemoreception are more important in detecting food, the eyes on catfish are generally small. Like other ]s, they are characterized by the presence of a ].<ref name="Nelson"/> Their well-developed Weberian apparatus and reduced ] allow for improved ] as well as sound production.<ref name="Bruton"/> | |||
Catfish have no ]s; their bodies are often naked. In some species, the ]-covered ] is used in cutaneous ], where the fish breathes through its skin.<ref name="Bruton"/> In some catfish, the skin is covered in bony plates called ]; some form of body armor appears in various ways within the order. In ] and in the Asian genus '']'', the armor is primarily made up of one or more rows of free ] plates. Similar plates are found in large specimens of '']''. These plates may be supported by ]l ]es, as in ] and in ''Sisor'', but the processes never fuse to the plates or form any external armor. By contrast, in the subfamily Doumeinae (family ]) and in hoplomyzontines (]), the armor is formed solely by expanded vertebral processes that form plates. Finally, the lateral armor of ], ''Sisor'', and hoplomyzontines consists of hypertrophied ] ]s with dorsal and ventral ].<ref>{{cite journal|author=J P Friel, J G Lundberg|year=1996|title=''Micromyzon akamai'', gen. et sp. nov., a small and eyeless banjo catfish (Siluriformes: Aspredinidae) from the river channels of the lower Amazon basin|journal=]|issue=3|pages=641–648|jstor=1447528|volume=1996}}</ref> | |||
] has four pairs of ].]] | |||
All catfish, except members of ] (electric catfish), possess a strong, hollow, bonified leading spine-like ray on their ] and ]s. As a defense, these spines may be locked into place so that they stick outwards, which can inflict severe wounds.<ref name=tol/> In several species catfish can use these fin rays to deliver a stinging ] if the fish is irritated.<ref name=fin>{{cite web| url = http://www.fcps.k12.va.us/StratfordLandingES/Ecology/mpages/channel_catfish.htm|title=Channel Catfish |accessdate=2006-12-02|publisher=Fairfax County Public Schools}}</ref> This ] is produced by ]ular cells in the ] tissue covering the spines.<ref name="Nelson"/> In members of the family ], and of the genus '']'', this protein is so strong it may hospitalize humans, those unfortunate enough to receive a sting; in '']'', the stings may result in death.<ref name="Nelson"/> | |||
Juvenile catfish, like most fish, have relatively large heads, eyes and posterior median fins in comparison to larger, more mature individuals. These juveniles can be readily placed in their families, particularly those with highly derived fin or body shapes; in some cases identification of the genus is possible. As far as known for most catfish, features that are often characteristic of species such as mouth and fin positions, fin shapes, and barbel lengths show little difference between juveniles and adults. For many species, pigmentation pattern is also similar in juveniles and adults. Thus, juvenile catfishes generally resemble and develop smoothly into their adult form without distinct juvenile specializations. Exceptions to this are the ariid catfishes, where the young retain yolk sacs late into juvenile stages, and many pimelodids, which may have elongated barbels and fin filaments or coloration patterns.<ref>{{cite journal|url=http://www.mansfield.ohio-state.edu/~tberra/pdf-files/Diplomystes.pdf|title=First description of small juveniles of the primitive catfish ''Diplomystes'' (Siluriformes: Diplomystidae)|first=John G.|last=Lundberg|coauthors=Berra, Tim M.; Friel, John P.|journal=Ichthyol. Explor. Freshwaters|volume=15|issue=1|pages=71–82|month=March | year=2004|format=PDF|accessdate=2009-06-22}} {{Dead link|date=September 2010|bot=H3llBot}}</ref> | |||
] is reported in about half of all families of catfish.<ref>{{cite journal|url=http://www.mapress.com/zootaxa/2006f/zt01125p056.pdf|title=''Synodontis acanthoperca'', a new species from the Ogôoué River system, Gabon with comments on spiny ornamentation and sexual dimorphism in mochokid catfishes (Siluriformes: Mochokidae)|first=John P.|last=Friel|coauthors=Vigliotta, Thomas R.|journal=]|volume=1125|pages=45–56|year=2006|format=PDF|accessdate=2009-06-22}}</ref> The modification of the ] into an ] (in internal fertilizers) as well as accessory structures of the reproductive apparatus (in both internal and external fertilizers) have been described in species belonging to 11 different families.<ref name="Mazzoldi">{{cite journal|title=Variation of male reproductive apparatus in relation to fertilization modalities in the catfish families Auchenipteridae and Callichthyidae (Teleostei: Siluriformes)|first=C.|last=Mazzoldi|coauthors=Lorenzi, V.; Rasotto, M. B.|journal=Journal of Fish Biology|year=2007|volume=70|pages=243–256|doi=10.1111/j.1095-8649.2006.01300.x}}</ref> | |||
===Size=== | |||
]'', may be fatal.]] | |||
Catfish have one of the greatest ranges in size within a single order of ].<ref name="Bruton"/> Many catfish have a maximum length of under 12 cm.<ref name="Nelson"/> Some of the smallest species of ] and ] reach sexual maturity at only {{convert|1|cm|in}}.<ref name=tol/> | |||
The ], ''Silurus glanis'', is the only native catfish species of Europe, besides the much smaller related ] found in Greece. ] and literature record wels catfish of astounding proportions, yet to be proven scientifically. The average size of the species is about 1.2–1.6 m (3.9–5.2 ft), and fish more than {{convert|2|m|ft}} are very rare. The largest specimens on record measure more than {{convert|2.5|m|ft}} in length and sometimes exceeded {{convert|100|kg|lb}}. | |||
'''Catfish''' (or '''catfishes'''; ] '''Siluriformes''' {{IPAc-en|s|ɪ|ˈ|lj|ʊər|ᵻ|f|ɔːr|m|iː|z}} or '''Nematognathi''') are a diverse group of ]. Named for their prominent ]s, which resemble a ]'s ]s, catfish range in size and behavior from the three largest species alive, the ] from ], the ] of ], and the ] of ], to ]s (species that eat dead material on the bottom), and even to a tiny ] species commonly called the ], ''Vandellia cirrhosa''. Neither the armour-plated types nor the naked types have scales. Despite their name, not all catfish have prominent barbels or "whiskers". Members of the Siluriformes order are defined by features of the skull and ]. Catfish are of considerable commercial importance; many of the larger species are ] or fished for food. Many of the smaller species, particularly the genus '']'', are important in the ]. Many catfish are ],<ref name="aworld"> {{Webarchive|url=https://web.archive.org/web/20120417234946/http://animal-world.com/encyclo/fresh/catfish/catfish.htm |date=17 April 2012 }}. animal-world.com</ref><ref name="samer">Wong, Kate (6 June 2001) {{Webarchive|url=https://web.archive.org/web/20110320110006/https://www.scientificamerican.com/article.cfm?id=how-nocturnal-catfish-sta |date=20 March 2011 }}. ''Scientific American''.</ref> but others (many ]) are ] or ] (most ] or ], for example). | |||
The largest '']'', caught in the ] on July 20, 2010, weighed {{convert|130|lb|kg}}. The largest ], ''Pylodictis olivaris'', ever caught was in ], weighing 123 lb 9 oz (56.0 kg). In July 2009, a catfish weighing 193 pounds was caught in the ], Spain, by an 11-year old British schoolgirl.<ref>{{cite news| url=http://www.telegraph.co.uk/news/newstopics/howaboutthat/5834754/Schoolgirl-nets-9ft-monster-fish.html | work=] | location=London | title=Schoolgirl nets 9ft monster fish Schoolgirl angler Jessica Wanstall netted a record when she hooked this monster fish that was more than twice the size of her. | date=2009-07-15 | accessdate=2010-04-28}}</ref> However, these records pale in comparison to a ] caught in northern ] on May 1, 2005 and reported to the press almost 2 months later that weighed {{convert|293|kg|lb}}. This is the largest giant Mekong catfish caught since Thai officials started keeping records in 1981.<ref name=Mekong>{{cite web|url=http://news.nationalgeographic.com/news/2005/06/0629_050629_giantcatfish.html|title=Grizzly Bear-Size Catfish Caught in Thailand |accessdate=2006-07-14|publisher=National Geographic News}}</ref> The giant Mekong catfish are not well studied since they live in developing countries and it is quite possible that they can grow even larger. | |||
===Internal anatomy=== | |||
In many catfish, the ''humeral process'' is a bony process extending backward from the ] immediately above the base of the pectoral fin. It lies beneath the skin where its outline may be determined by dissecting the skin or probing with a needle.<ref>{{cite web|url=http://filaman.ifm-geomar.de/Glossary/Glossary.cfm?TermEnglish=humeral%20process|title=Term : humeral process|publisher=]|year=2007}}</ref> | |||
The ] of catfish are composed of single ]s and large ]s. Many catfish have a ] which may help enhance ] capture and increase low-light sensitivity. ]s, though present in most ]s, are absent from catfish.<ref name="Douglas">{{cite journal|last=Douglas|first=Ron H.|coauthors=Collin, Shaun P.; Corrigan, Julie|date=15 November 2002|url=http://jeb.biologists.org/cgi/reprint/205/22/3425|title=The eyes of suckermouth armoured catfish (Loricariidae, subfamily Hypostomus): pupil response, lenticular longitudinal spherical aberration and retinal topography|publisher=The Journal of Experimental Biology|volume=205|issue=22|pages=3425–3433|format=PDF|journal=Journal of Experimental Biology|pmid=12364396}}</ref> | |||
The anatomical organization of the ] in catfish is variable among the families of catfish, but the majority of them present fringed testis: Ictaluridae, Claridae, Auchenipteridae, Doradidae, Pimelodidae, and Pseudopimelodidae.<ref name="Barros"/> In the testes of some species of Siluriformes, organs and structures such as a spermatogenic cranial region and a secretory caudal region are observed, in addition to the presence of seminal vesicles in the caudal region.<ref name="Brito"/> The total number of fringes and their length are different in the ] and ] portions between species.<ref name="Barros">{{cite journal|title=Reproductive apparatus and gametogenesis of ''Lophiosilurus alexandri'' Steindachner (Pisces, Teleostei, Siluriformes)|first=Marcelo D. M.|last=Barros|coauthors=Guimarães-Cruz, Rodrigo J.; Veloso-Júnior, Vanderlei C.; Santos, José E. dos|journal=Revista Brasileira de Zoologia|volume=24|issue=1|pages=213–221|year=2007|doi=10.1590/S0101-81752007000100028}}</ref> Fringes of the caudal region may present tubules, in which the lumen is filled by secretion and ].<ref name="Barros"/> Spermatocysts are formed from cytoplasmic extensions of ]s; the release of spermatozoa is allowed by breaking of the cyst walls.<ref name="Barros"/> | |||
The occurrence of ]s, in spite of their interspecific variability in size, gross morphology and function, has not been related to the mode of fertilization. They are typically paired, multi-chambered, and connected with the ], and have been reported to play a glandular and a storage function. Seminal vesicle secretion may include ]s and steroid glucuronides, with hormonal and pheromonal functions, but it appears to be primarily constituted of mucoproteins, acid mucopolysaccharides, and phospholipids.<ref name="Mazzoldi"/> | |||
Fish ovaries may be of two types: gymnovarian or cystovarian. In the first type, the oocytes are released directly into the ]ic cavity and then eliminated. In the second type, the oocytes are conveyed to the exterior through the ].<ref name="Brito"/> Many catfish are cystovarian in type, including '']'', '']'', '']'', and '']''.<ref name="Barros"/><ref name="Brito">{{cite journal|title=Reproduction of the surubim catfish (Pisces, Pimelodidae) in the São Francisco River, Pirapora Region, Minas Gerais, Brazil|last=Brito|first=M.F.G.|coauthors=Bazzoli, N.|journal=Arquivo Brasileiro de Medicina Veterinária e Zootecnia|volume=55|issue=5|year=2003|doi=10.1590/S0102-09352003000500018|pages=624}}</ref> | |||
==Catfish as food== | |||
] | |||
] | |||
Catfish have widely been caught and farmed for food for hundreds of years in Africa, Asia, Europe, and North America. Judgments as to the quality and flavor vary, with some food critics considering catfish as being excellent food, while others dismiss them as watery and lacking in flavor.<ref name=Baker>Jenny Baker (1988), ''Simply Fish'' p 36-37. Faver & Faber, London.</ref> In ], catfish were often viewed as a delicacy to be enjoyed on feast days and holidays. Migrants from Europe and Africa to the United States brought along this tradition, and in the ], catfish is an extremely popular food. The most commonly eaten species in the United States are the ] and the ], both of which are common in the wild and increasingly widely farmed. Farm-raised catfish became such a staple of the diet of the United States that on June 25, 1987, President ] established ] to recognize "the value of farm-raised catfish." | |||
Catfish is eaten in a variety of ways. In Europe it is often cooked in similar ways to ], but in the United States it is popularly crumbed with ] and fried.<ref name=Baker/> | |||
In ], catfish is usually served grilled in street stalls called '']'' and eaten with vegetables and ]; the dish is called ''pecel lele''. Catfish can also be eaten with ] as ''lele penyet'' (minced catfish). (''Lele'' is the ]n word for catfish.) | |||
In ] catfish, called "ikan keli", is fried with spices or grilled and eaten with tamarind and Thai chillies gravy and also is often eaten with steamed rice. | |||
In ] and the Indian states of ], ] and ] catfish (locally known as Magur) is eaten as a favored delicacy during the monsoons. Catfish, locally known as ''thedu''' or ''etta''' in Malayalam, is very famous in the Indian state ]. In the inland ponds in Kerala, 2 varieties of catfish is abundant- Muzhi and Kari while "Etta" is a basically a salt water fish. The smaller, slender Kari is notorious for its ability to sting, and Muzhi is much bigger and easy to catch, especially during Monsoon when this seems to literally walk where very little water is present from the rain water. All the catfish are eaten as curry and their extra-large eggs, especially that of Etta, is fried and is a delicacy. It is also believed that catfish meat helps in blood purification. Catfish curry is consumed in these parts to promote faster recovery to patients suffering from fever or other ailments. | |||
In Hungary catfish is often cooked in paprika sauce (Harcsapaprikás) typical of Hungarian cuisine. It is traditionally served with pasta smothered with curd cheese (]). | |||
Catfish is high in ].<ref name = FactD>{{cite web|url=http://www.health.state.ny.us/diseases/conditions/osteoporosis/vitd.htm|title=Vitamin D and Healthy Bones|publisher=New York State Department of Health|accessdate=2007-07-13 }}</ref> Farm-raised catfish contains low levels of ]s and a much higher proportion of ]s.<ref>Reuters, . Source: Journal of the American Dietetic Association, July 2008</ref> | |||
Vietnamese catfish cannot be legally marketed as catfish in the United States, and is subsequently referred to as ] or ] <ref name = FactSwai>{{cite web|url=http://www.unionfishco.com/swai/details.htm|title=Union Fish Company – Basa/Swai Details|accessdate=2007-11-11 |archiveurl = http://web.archive.org/web/20071109150512/http://www.unionfishco.com/swai/details.htm |archivedate = 2007-11-09}}</ref> Only fish of the family Ictaluridae may be marketed as catfish in the United States.<ref name = 21USC343>{{cite web|url=http://www.law.cornell.edu/uscode/21/usc_sec_21_00000343----000-.html|title=United States Code, Title 21,343. Misbranded food}}</ref> | |||
As catfish lack scales, they are judged not to be ] and may not be eaten by observant ], some ] who follow the ]'s ], and observant ] of various schools. | |||
==Aquaculture== | |||
{{Main|Aquaculture of catfish}} | |||
Catfish are easy to farm in warm climates, leading to inexpensive and safe food at local grocers. About 60% of U.S. farm-raised catfish are grown within a 65-mile (100-km) radius of ].<ref>{{Cite journal | author= J.E. Morris|title= Pond Culture of Channel Catfish in the North Central Region | publisher= North Central Regional Aquaculture Center| date= October 1993| format= PDF| url= http://www.extension.iastate.edu/Publications/NCR444.pdf| accessdate= 2006-06-28}}</ref> ] (''Ictalurus punctatus'') supports a $450 million/yr aquaculture industry.<ref name=tol/> | |||
Catfish raised in inland tanks or channels are considered safe for the environment, since their waste and disease should be contained and not spread to the wild.<ref>{{cite journal | first = Paul | last = Rogers | title = Economy of Scales | work = Stanford Magazine | publisher = ] | issue = March / April 2006 | url = http://www.stanfordalumni.org/news/magazine/2006/marapr/features/fishfarming.html | accessdate = 2008-02-14}}</ref> | |||
In Asia, many catfish species are important as food. Several ] (Clariidae) and ] (Pangasiidae) species are heavily cultured in Africa and Asia. Exports of one particular shark catfish species from ], '']'', has met with pressures from the U.S. catfish industry. In 2003, The ] passed a law preventing the imported fish from being labeled as catfish.<ref></ref> As a result, the Vietnamese exporters of this fish now label their products sold in the U.S. as "basa fish." Trader Joe's has labeled frozen fillets of Vietnamese '']'' as "striper."<ref></ref> | |||
There is a large and growing ornamental fish trade, with hundreds of species of catfish, such as '']'' and ] (often called ]s), being a popular component of many ]. Other catfish commonly found in the aquarium trade are ], ], and ]. | |||
==Catfish as invasive species== | |||
] is an ] in Florida.]] | |||
Representatives of the ] '']'' have been introduced into European waters in the hope of obtaining a sporting and food resource. However, the European stock of American catfishes has not achieved the dimensions of these fish in their native waters, and have only increased the ecological pressure on native European ]. ] have also been introduced in the freshwaters of Florida, with the voracious catfish becoming a major alien pest there. ], ''Pylodictis olivaris'', is also a North American pest on Atlantic slope drainages.<ref name=tol/> '']'' species, released by aquarium fishkeepers, have also established ] populations in many warm waters around the world.<ref>{{cite journal|title=The South American Suckermouth Armored Catfish, ''Pterygoplichthys anisitsi'' (Pisces: Loricaridae), in Texas, with Comments on Foreign Fish Introductions in the American Southwest|first=Leo G.|last=Nico|coauthors=Martin, R. Trent|journal=The Southwestern Naturalist|volume=46|issue=1|month=March | year=2001|pages=98–104|doi=10.2307/3672381|jstor=3672381}}</ref><ref>{{cite journal|title=Amazon Sailfin Catfish, ''Pterygoplichthys pardalis'' (Castelnau, 1855) (Loricariidae), Another Exotic Species Established in Southeastern Mexico|first=Armando T.|last=Wakida-Kusunokia|coauthors=Ruiz-Carusb, Ramon; Amador-del-Angelc, Enrique|journal=The Southwestern Naturalist|pages=141–144|volume=52|issue=1|month=March | doi=10.1894/0038-4909(2007)522.0.CO;2|year=2007|issn=0038-4909}}</ref><ref>{{cite journal|url=http://www.mapress.com/zootaxa/2006f/zt01109p068.pdf|title=New Philippine record of South American sailfin catfishes (Pisces: Loricariidae)|first=Joel M.|last=Chavez|coauthors=de la Paz, Reynaldo M.; Manohar, Surya Krishna; Pagulayan, Roberto C.; Carandang Vi, Jose R.|journal=]|volume=1109|pages=57–68|year=2006|format=PDF|accessdate=2009-06-22}}</ref><ref>{{cite journal|url=http://academic.uprm.edu/publications/cjs/VOL30/P090-094.PDF|title=The South American Sailfin Armored Catfish, ''Liposarcus multiradiatus'' (Hancock), a New Exotic Established in Puerto Rican Fresh Waters|first=Lucy|last=Bunkley-Williams|coauthors=Williams, Ernest H., Jr.; Lilystrom, Craig G.; Corujo-Flores, Iris; Zerbi, Alfonso J.; Aliaume, Catherine; Churchill, Timothy N.|journal=Caribbean Journal of Science|volume=30|issue=1–2|pages=90–94|year=1994|format=PDF|accessdate=2009-06-22}}</ref><ref>{{cite journal|url=http://zoolstud.sinica.edu.tw/Journals/44.2/252.pdf|journal=Zoological Studies|volume=44|issue=2|pages=252–259|year=2005|title=Size Structure, Reproductive Phenology, and Sex Ratio of an Exotic Armored Catfish (''Liposarcus multiradiatus'') in the Kaoping River of Southern Taiwan|first=Shih-Hsiung|last=Liang|coauthors=Wu, Hsiao-Ping; Shieh, Bao-Sen|format=PDF|accessdate=2009-06-22}}</ref> | |||
==Dangers to humans== | |||
While the vast majority of catfish are harmless to humans, a few species are known to present some risk. Perhaps the most notorious of these is the ], due to the way it is reputed to parasitize the urethra, though there is only ]. | |||
Since 2007, the ] has also gained attention<ref>{{cite news|last=Cox|first=Emma|title=Humans Scoffed by Mutant Fish|url=http://www.thesun.co.uk/sol/homepage/news/weird/article1784470.ece|accessdate=1 October 2010|newspaper=The Sun|date=09 October, 2008}}</ref><ref>{{cite web|title=Corpse-Eating Mutant Fish Caught|url=http://news.sky.com/skynews/Home/World-News/Flesh-Eating-Catfish-Goonch-Tracked-Down-By-Angler-Jeremy-Wade-For-Channel-Five/Article/200810215117069|work=News.sky.com|publisher=Sky.com|accessdate=1 October 2010}}</ref> following a ] which have been alleged by biologist ] to have been from unusually large goonch. | |||
The ] has also been reputed to kill humans (especially young children), and while there are no documented cases of fatalities, larger specimens are known to cause serious injuries in rare instances.<ref>{{cite web|title=Wels Catfish|url=http://animal.discovery.com/fish/river-monsters/wels-catfish/|work=River Monsters|publisher=Animal Planet|accessdate=1 October 2010}}</ref> In addition, other species are reputed to be dangerous to humans as well, but with less definitive evidence. | |||
Many catfish species have “stings” (actually non-venomous in most cases) embedded behind their fins; thus precautions must be taken when handling them. | |||
==Taxonomy== | ==Taxonomy== | ||
Molecular evidence suggests that in spite of the great morphological diversity in the order, all catfish form a ] group.<ref name="Sullivan">{{cite journal |last=Sullivan |first=JP |author2=Lundberg JG |author3=Hardman M |year=2006 |title=A phylogenetic analysis of the major groups of catfish (Teleostei: Siluriformes) using rag1 and rag2 nuclear gene sequences |journal=Mol Phylogenet Evol |volume=41 |issue=3 |pages=636–62 |doi=10.1016/j.ympev.2006.05.044 |pmid=16876440|bibcode=2006MolPE..41..636S }}</ref> Catfish belong to a superorder called the ], which also includes the ] (carps and minnows), ] (characins and tetras), ] (milkfish and beaked salmons) and ] (South American knifefish), a superorder characterized by the ]. Some place Gymnotiformes as a sub-order of Siluriformes; however, this is not as widely accepted. Currently, the Siluriformes are said to be the ] to the Gymnotiformes, though this has been debated due to more recent molecular evidence.<ref name="Nelson">{{cite book |last=Nelson |first=Joseph S. |title=Fishes of the World |title-link=Fishes of the World |publisher=], Inc |year=2006 |isbn=978-0-471-25031-9}}</ref> {{As of|2007}} there were about thirty-six ] catfish families, and about 3,093 extant species have been described.<ref name="ferraris">{{cite journal |last1=Ferraris |first1=Carl J. Jr. |last2=Miya |first2=M |last3=Azuma |first3=Y |last4=Nishida |first4=M |year=2007 |title=Checklist of catfish, recent and fossil (Osteichthyes: Siluriformes), and catalogue of siluriform primary types |url=http://mapress.com/zootaxa/2007f/zt01418p300.pdf |url-status=live |journal=] |volume=1418 |pages=1–628 |citeseerx=10.1.1.232.798 |doi=10.11646/zootaxa.1418.1.1 |archive-url=https://web.archive.org/web/20080414082401/http://mapress.com/zootaxa/2007f/zt01418p300.pdf |archive-date=2008-04-14 |access-date=22 June 2009}}</ref> This makes the catfish order the second or third most diverse ] order; in fact, one out of every twenty vertebrate species is a catfish.<ref name="tol">{{cite web |last=Lundberg |first=John G. |author2=Friel, John P. |date=20 January 2003 |title=Siluriformes |url=http://tolweb.org/tree?group=Siluriformes&contgroup=Ostariophysi |url-status=live |archive-url=https://web.archive.org/web/20070128012752/http://tolweb.org/tree?group=Siluriformes&contgroup=Ostariophysi |archive-date=28 January 2007 |access-date=18 April 2007 |publisher=]}}</ref> | |||
The catfishes are a ] group. This is supported by molecular evidence.<ref name="Sullivan"/> | |||
Catfish belong to a superorder called the ], which also includes the ], ], ] and ], a superorder characterized by the ]. Some place Gymnotiformes as a sub-order of Siluriformes, however this is not as widely accepted. Currently, the Siluriformes are said to be the ] to the Gymnotiformes, though this has been debated due to more recent molecular evidence.<ref name="Nelson"/> {{As of|2007}} there are about 36 ] catfish families, and about 3,093 extant species have been described.<ref name=ferraris5>{{cite journal|url=http://mapress.com/zootaxa/2007f/zt01418p300.pdf|title=Checklist of catfishes, recent and fossil (Osteichthyes: Siluriformes), and catalogue of siluriform primary types|first=Carl J., Jr.|last=Ferraris|journal=]|volume=1418|page=5|year=2007|format=PDF|accessdate=2011-10-05|last2=Miya|first2=M|last3=Azuma|first3=Y|last4=Nishida|first4=M}}</ref> This makes the catfish order the second or third most diverse ] order; in fact, 1 out of every 20 vertebrate species is a catfish.<ref name=tol>{{cite web|url=http://tolweb.org/tree?group=Siluriformes&contgroup=Ostariophysi|title=Siluriformes|last=Lundberg|first=John G.|coauthors=Friel, John P.|publisher=]|date=2003-01-20|accessdate=2007-04-18}}</ref> | |||
The taxonomy of catfishes is quickly changing. In a 2007 and 2008 paper, '']'', '']'', and '']'' were not classified under any current catfish families.<ref name=ferraris>{{cite journal|url=http://mapress.com/zootaxa/2007f/zt01418p300.pdf|title=Checklist of catfishes, recent and fossil (Osteichthyes: Siluriformes), and catalogue of siluriform primary types|first=Carl J., Jr.|last=Ferraris|journal=]|volume=1418|pages=1–628|year=2007|format=PDF|accessdate=2009-06-22|last2=Miya|first2=M|last3=Azuma|first3=Y|last4=Nishida|first4=M}}</ref> There is disagreement on the family status of certain groups; for example, Nelson (2006) lists Auchenoglanididae and Heteropneustidae as separate families, while the All Catfish Species Inventory (ACSI) includes them under other families. Also, ] and the ] lists Parakysidae as a separate family, while this group is included under ] by both Nelson (2006) and ACSI.<ref name="Nelson"/><ref>{{cite web|url=http://silurus.acnatsci.org/ACSI/taxa/Families.html|title=Catfish Families|publisher=All Catfish Species Inventory|accessdate=2007-04-28}}</ref><ref>{{FishBase family|family=Parakysidae|year=2007|month=Apr}}</ref><ref>{{ITIS |id=553185 |taxon=Parakysidae |accessdate=April 2007}}</ref> Many sources do not list the recently revised family ].<ref name=Anchariidae>{{cite journal|url=http://www.pfeil-verlag.de/04biol/pdf/ief16_4_02.pdf|title=Revision of the endemic Malagasy catfish family Anchariidae (Teleostei: Siluriformes), with descriptions of a new genus and three new species|first=Heok Hee|last=Ng|coauthors=Sparks, John S.|year=2005|journal=Ichthyol. Explor. Freshwaters|volume=16|issue=4|pages=303–323}}</ref> The family ], including ''Horabagrus'', '']'', and '']'', is also not shown by some authors but presented by others as a true group.<ref name="Sullivan"/> Thus, the actual number of families differs between authors. The species count is in constant flux due to ] work as well as description of new species. On the other hand, our understanding of catfishes should increase in the next few years due to work by the ACSI.<ref name="Nelson">{{cite book|title=]|last=Nelson|first=Joseph S.|publisher=], Inc|year=2006|isbn=0-471-25031-7}}</ref> | |||
Catfish are believed to have a ]n origin primarily centered around South America, as the most ] living catfish groups are known from there. The earliest known definitive members lived in the ] from the ] to ] stages of the ], including the ], ''Vorhisia vulpes'' and possibly '']''.<ref name=NT24/><ref>{{Cite journal |last1=Stringer |first1=Gary |last2=Schwarzhans |first2=Werner |date=2021-09-01 |title=Upper Cretaceous teleostean otoliths from the Severn Formation (Maastrichtian) of Maryland, USA, with an unusual occurrence of Siluriformes and Beryciformes and the oldest Atlantic coast Gadiformes |journal=Cretaceous Research |volume=125 |pages=104867 |doi=10.1016/j.cretres.2021.104867 |issn=0195-6671|doi-access=free |bibcode=2021CrRes.12504867S }}</ref><ref name=Cavin2017/> A potential fossil record is known from the earlier ]-] stages in ] of ],<ref name=Patterson1993>{{cite book|author=Patterson, C.|year=1993|chapter=Osteichthyes: Teleostei|editor=Benton, M.J.|title=The Fossil Record 2|pages=621-656|publisher=]|location=London}}</ref> though this has been considered unreliable,<ref name=Cavin2017>{{Citation |last=Cavin |first=Lionel |title=Evolutionary Histories of Freshwater Fishes |date=2017 |work=Freshwater Fishes: 250 Million Years of Evolutionary History |pages=53–125 |url=http://dx.doi.org/10.1016/b978-1-78548-138-3.50004-2 |access-date=2024-05-08 |publisher=Elsevier |isbn=978-1-78548-138-3}}</ref> and the putative earliest ] known from the fossil record, '']'', lived during the ] age of the ] in ] of ] (]).<ref name="Afrocascudo" /> The describers of ''Afrocascudo'' claimed that the presence of a derived loricariid so early on would indicate the extensive diversification of catfish, or at least loricarioids, prior to the beginning of the Late Cretaceous. As extant loricariids are only known from South America, much of this diversification must have occurred on the supercontinent of ] prior to its fragmentation into South America and Africa.<ref name="Afrocascudo" /> Britz and colleagues suggested that ''Afrocascudo'' instead represents a juvenile ] ], possibly a junior synonym of ''Obaichthys''.<ref>{{cite journal|author1=Britz, R.|author2=Pinion, Amanda K.|author3=Kubicek, Kole M.|author4=Conway, Kevin W.|year=2024|title=Comment on “A Saharan fossil and the dawn of Neotropical armoured catfishes in Gondwana” by Brito et al|journal=Gondwana Research|doi=10.1016/j.gr.2024.06.014}}</ref> The authors of the original study still stood by their original conclusion based on the absence of important ] characters, and noted that it could not be a juvenile, since the bones were completely ossified.<ref>{{cite journal|last1=Brito |first1=Paulo M. |last2=Dutheil |first2=Didier B. |last3=Keith |first3=Philippe |last4=Carnevale |first4=Giorgio |last5=Meunier |first5=François J. |last6=Khalloufi |first6=Bouziane |last7=Gueriau |first7=Pierre |year=2024|title=A reply to a comment on Brito et al., 2024, A Saharan fossil and the dawn of the Neotropical armoured catfishes in Gondwana by Britz, Pinion, Kubicek and Conway|journal=Gondwana Research|doi=10.1016/j.gr.2024.06.013}}</ref> | |||
The rate of description of new catfishes is at an all-time high. Between 2003 and 2005, over 100 species have been named, a rate three times faster than that of the past century.<ref name=neotropdiversity>{{cite journal|url=http://www.ufrgs.br/ni/vol3num4%5Cpreface.pdf|title=Neotropical catfish diversity: an historical perspective|first=Carl J., Jr.|last=Ferraris|coauthors=]|journal=Neotropical Ichthyology|volume=3|issue=4|pages=453–454|year=2005|doi=10.1590/S1679-62252005000400001|format=PDF}}</ref> In June, 2005, researchers named the newest family of catfish, ], only the third new family of fish distinguished in the last 70 years (others being the ] in 1938 and the ] in 1983). The new species in ], '']'', was found in the ] in the Mexican state of ].<ref name=rodiles>{{cite journal|title=''Lacantunia enigmatica'' (Teleostei: Siluriformes) a new and phylogenetically puzzling freshwater fish from Mesoamerica|last=Rodiles-Hernández|first=Rocío|coauthors=Hendrickson, Dean A.; Lundberg, John G.; Humphries, Julian M.|journal=]|pages=1–24|year=2005|volume=1000|format=PDF|url=http://biostor.org/reference/15994|accessdate=2009-06-22}}</ref> | |||
])'' skeleton on display at the ].]] | |||
The taxonomy of catfish is quickly changing. In a 2007 and 2008 paper, '']'', '']'', and '']'' were not classified under any current catfish families.<ref name="ferraris" /> There is disagreement on the family status of certain groups; for example, Nelson (2006) lists Auchenoglanididae and Heteropneustidae as separate families, while the All Catfish Species Inventory (ACSI) includes them under other families. ] and the ] lists Parakysidae as a separate family, while this group is included under ] by both Nelson (2006) and ACSI.<ref name="Nelson" /><ref>{{cite web |title=Catfish Families |url=http://silurus.acnatsci.org/ACSI/taxa/Families.html |url-status=live |archive-url=https://web.archive.org/web/20070502195241/http://silurus.acnatsci.org/ACSI/taxa/Families.html |archive-date=2 May 2007 |access-date=28 April 2007 |publisher=All Catfish Species Inventory}}</ref><ref>{{FishBase family|family=Parakysidae|year=2007|month=April}}</ref><ref>{{ITIS|id=553185|taxon=Parakysidae|access-date=10 September 2016}}</ref> Many sources do not list the recently revised family ].<ref name="Anchariidae">{{cite journal |last=Ng |first=Heok Hee |author2=Sparks, John S. |year=2005 |title=Revision of the endemic Malagasy catfish family Anchariidae (Teleostei: Siluriformes), with descriptions of a new genus and three new species |url=http://www.pfeil-verlag.de/04biol/pdf/ief16_4_02.pdf |url-status=live |journal=Ichthyol. Explor. Freshwaters |volume=16 |issue=4 |pages=303–323 |archive-url=https://web.archive.org/web/20071215172203/http://www.pfeil-verlag.de/04biol/pdf/ief16_4_02.pdf |archive-date=2007-12-15}}</ref> The family ], including ''Horabagrus'', '']'', and '']'', is not shown by some authors but presented by others as a true group.<ref name="Sullivan" /> Thus, the actual number of families differs between authors. The species count is in constant flux due to ] work as well as description of new species.<ref name="Nelson" /> Between 2003 and 2005, over one hundred species were named, a rate three times faster than that of the past century.<ref name="neotropdiversity">{{cite journal |last=Ferraris |first=Carl J. Jr. |author2=Reis, Roberto E. |author-link2=Roberto Esser dos Reis |year=2005 |title=Neotropical catfish diversity: an historical perspective |journal=Neotropical Ichthyology |volume=3 |issue=4 |pages=453–454 |doi=10.1590/S1679-62252005000400001 |doi-access=free}}</ref> In June 2005, researchers named the newest family of catfish, ], only the third new family of fish distinguished in the last seventy years, the others being the ] in 1938 and the ] in 1983. The new species in ], '']'', was found in the ] in the Mexican state of ].<ref name="rodiles">{{cite journal |last=Rodiles-Hernández |first=Rocío |author2=Hendrickson, Dean A. |author3=Lundberg, John G. |author4=Humphries, Julian M. |year=2005 |title=''Lacantunia enigmatica'' (Teleostei: Siluriformes) a new and phylogenetically puzzling freshwater fish from Mesoamerica |url=http://biostor.org/reference/15994 |url-status=live |journal=] |volume=1000 |pages=1–24 |doi=10.11646/zootaxa.1000.1.1 |archive-url=https://web.archive.org/web/20121025015806/http://biostor.org/reference/15994 |archive-date=25 October 2012 |access-date=22 June 2009 |doi-access=free}}</ref> | |||
The higher-level phylogeny of Siluriformes has gone through several recent changes, mainly due to ] studies. While most studies, both morphological and molecular, agree that catfishes are arranged into three main ], the relationship among these lineages has been a contentious point in which these studies, performed for example by ], differ.<ref name=":0">{{Cite journal |last1=Arcila |first1=Dahiana |last2=Ortí |first2=Guillermo |last3=Vari |first3=Richard |last4=Armbruster |first4=Jonathan W. |last5=Stiassny |first5=Melanie L. J. |last6=Ko |first6=Kyung D. |last7=Sabaj |first7=Mark H. |last8=Lundberg |first8=John |last9=Revell |first9=Liam J. |date=2017-01-13 |title=Genome-wide interrogation advances resolution of recalcitrant groups in the tree of life |journal=Nature Ecology & Evolution |volume=1 |issue=2 |pages=0020 |bibcode=2017NatEE...1...20A |doi=10.1038/s41559-016-0020 |pmid=28812610 |s2cid=16535732}}</ref><ref name=":1">{{Cite journal |last1=Chen |first1=Wei-Jen |last2=Lavoué |first2=Sébastien |last3=Mayden |first3=Richard L. |date=2013-04-09 |title=Evolutionary Origin and Early Biogeography of Otophysan Fishes (Ostariophysi: Teleostei) |journal=Evolution |volume=67 |issue=8 |pages=2218–2239 |doi=10.1111/evo.12104 |pmid=23888847 |s2cid=40056087 |doi-access=free}}</ref><ref name=":2">{{Cite journal |last1=Rivera-Rivera |first1=Carlos J. |last2=Montoya-Burgos |first2=Juan I. |date=October 2018 |title=Back to the roots: Reducing evolutionary rate heterogeneity among sequences gives support for the early morphological hypothesis of the root of Siluriformes (Teleostei: Ostariophysi) |journal=Molecular Phylogenetics and Evolution |volume=127 |pages=272–279 |doi=10.1016/j.ympev.2018.06.004 |pmid=29885935 |s2cid=47014511 |doi-access=free|bibcode=2018MolPE.127..272R }}</ref><ref name=":3">{{Cite journal |last=Diogo |first=Rui |date=2004-11-01 |title=Phylogeny, origin and biogeography of catfishes: support for a Pangean origin of 'modern teleosts' and reexamination of some Mesozoic Pangean connections between the Gondwanan and Laurasian supercontinents |journal=Animal Biologyn |volume=54 |issue=4 |pages=331–351 |doi=10.1163/1570756042729546}}</ref><ref name=":4">{{Cite book |last=Rui. |first=Diogo |title=The origin of higher clades : osteology, myology, phylogeny and evolution of bony fishes and the rise of tetrapods |date=2007 |publisher=Science Publishers |isbn=9781578085590 |location=Enfield, NH |oclc=680560456}}</ref> The three main lineages in Siluriformes are the family ], the denticulate catfish suborder ] (containing the Neotropical "suckermouth" catfishes), and the suborder Siluroidei, which contains the remaining families of the order. According to ] data, ] is usually considered to be the earliest branching catfish lineage and the ] to the other two lineages, Loricarioidei and Siluroidei.<ref name=":3" /><ref name=":4" /><ref>{{Cite journal |last=Yang |first=Lei |date=April 2011 |title=GONORYNCHIFORMES AND OSTARIOPHYSAN RELATIONSHIPS: A COMPREHENSIVE REVIEW - Edited by T. Grande, F. J. Poyato-Ariza and R. Diogo |journal=Journal of Fish Biology |volume=78 |issue=4 |pages=1277–1278 |bibcode=2011JFBio..78.1277Y |doi=10.1111/j.1095-8649.2011.02907.x}}</ref> Molecular evidence usually contrasts with this hypothesis, and shows the suborder Loricarioidei as the earliest branching catfish lineage, and sister to a ] that includes the Diplomystidae and Siluroidei; this phylogeny has been obtained in numerous studies based on genetic data.<ref name="Sullivan" /><ref name=":0" /><ref name=":1" /><ref>{{Cite journal |last1=Nakatani |first1=Masanori |last2=Miya |first2=Masaki |last3=Mabuchi |first3=Kohji |last4=Saitoh |first4=Kenji |last5=Nishida |first5=Mutsumi |date=2011-06-22 |title=Evolutionary history of Otophysi (Teleostei), a major clade of the modern freshwater fishes: Pangaean origin and Mesozoic radiation |journal=BMC Evolutionary Biology |volume=11 |issue=1 |pages=177 |bibcode=2011BMCEE..11..177N |doi=10.1186/1471-2148-11-177 |pmc=3141434 |pmid=21693066 |doi-access=free}}</ref> However, it has been suggested that these molecular results are errors as a result of ], incorrectly placing Loricarioidei as the earliest-branching catfish lineage.<ref name=":2" /> When a data filtering method<ref>{{cite journal |last1=Rivera-Rivera |first1=Carlos J. |last2=Montoya-Burgos |first2=Juan I. |date=2019-08-13 |title=LSX: automated reduction of gene-specific lineage evolutionary rate heterogeneity for multi-gene phylogeny inference |journal=BMC Bioinformatics |publisher=Springer Science and Business Media LLC |volume=20 |issue=1 |page=420 |biorxiv=10.1101/220053 |doi=10.1186/s12859-019-3020-1 |pmc=6693147 |pmid=31409290 |doi-access=free}}</ref> was used to reduce lineage rate heterogeneity (the potential source of bias) on their dataset, a final phylogeny was recovered which showed the ] are the earliest-branching catfish, followed by ] and Siluroidei as sister lineages, providing both morphological and molecular support for ] being the earliest branching catfish.<ref name=":2"/> | |||
According to ] data, ] is usually considered to be the most primitive of catfishes and the ] to the remaining catfishes, grouped in a ] called '''Siluroidei'''. Recent molecular evidence contrasts the prevailing hypothesis, where the suborder Loricarioidei are the sister group to all catfishes, including Diplomystidae (Diplomystoidei) and Siluroidei; though they were not able to reject the past hypothesis, the new hypothesis is not unsupported. Siluroidei was found to be monophyletic without Loricarioid families or Diplomystidae with molecular evidence; morphological evidence is unknown that supports Siluroidei without Loricarioidea.<ref name="Sullivan"/> | |||
Below is a list of family relationships by different authors. Lacantuniidae is included in the Sullivan scheme based on recent evidence that places it sister to ].<ref>{{cite journal| |
Below is a list of family relationships by different authors. Lacantuniidae is included in the Sullivan scheme based on recent evidence that places it sister to ].<ref>{{cite journal |last=Lundberg |first=John G. |author2=Sullivan, John P. |author3=Rodiles-Hernández, Rocío |author4=Hendrickson, Dean A. |year=2007 |title=Discovery of African roots for the Mesoamerican Chiapas catfish, ''Lacantunia enigmatica'', requires an ancient intercontinental passage |url=https://webspace.utexas.edu/deanhend/www/pdfs/Lundberg_2007_African_roots_Lacantunia.pdf |url-status=dead |journal=Proceedings of the Academy of Natural Sciences of Philadelphia |volume=156 |pages=39–53 |doi=10.1635/0097-3157(2007)1562.0.CO;2 |s2cid=4171034 |archive-url=https://web.archive.org/web/20090326130159/https://webspace.utexas.edu/deanhend/www/pdfs/Lundberg_2007_African_roots_Lacantunia.pdf |archive-date=26 March 2009 |access-date=22 June 2009}}</ref> | ||
{| | {| | ||
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{| class="wikitable |
{| class="wikitable" | ||
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! colspan=2 | Nelson, 2006<ref name="Nelson"/> | ! colspan="2" | Nelson, 2006<ref name="Nelson" /> | ||
! colspan="2" | Sullivan et al., 2006<ref name="Sullivan" /> | |||
! colspan=2 | Sullivan et al., 2006<ref name="Sullivan">{{cite journal|journal=Mol Phylogenet Evol.|year=2006|volume=41|issue=3|pages=636–62|title=A phylogenetic analysis of the major groups of catfishes (Teleostei: Siluriformes) using rag1 and rag2 nuclear gene sequences|last=Sullivan|first=JP|coauthors=Lundberg JG; Hardman M|doi=10.1016/j.ympev.2006.05.044|pmid=16876440}}</ref> | |||
|- style="vertical-align:top;" | |- style="vertical-align:top;" | ||
| |
| style="width:25%;" | | ||
*Unresolved families | *Unresolved families | ||
**Cetopsidae | **Cetopsidae | ||
Line 198: | Line 101: | ||
**Doradidae | **Doradidae | ||
**Auchenipteridae | **Auchenipteridae | ||
| |
| style="width:25%;" | | ||
*Siluroidea | *Siluroidea | ||
**Siluridae | **Siluridae | ||
Line 215: | Line 118: | ||
**Bagridae | **Bagridae | ||
**Pimelodidae | **Pimelodidae | ||
| |
| style="width:25%;" | | ||
*Unresolved families | *Unresolved families | ||
**Cetopsidae | **Cetopsidae | ||
Line 243: | Line 146: | ||
**Ictaluridae | **Ictaluridae | ||
**Cranoglanididae | **Cranoglanididae | ||
| |
| style="width:25%;" | | ||
*Doradoidea (sister to Aspredinidae) | *Doradoidea (sister to Aspredinidae) | ||
**Doradidae | **Doradidae | ||
Line 255: | Line 158: | ||
**'']'' + '']'' (Asian schilbeids) | **'']'' + '']'' (Asian schilbeids) | ||
**Horabagridae ('']'' + '']'' + '']'') | **Horabagridae ('']'' + '']'' + '']'') | ||
**Bagridae (without '']'') | **Bagridae (without '']'') | ||
*" |
*"Africa" | ||
**Mochokidae | **Mochokidae | ||
**Malapteruridae | **Malapteruridae | ||
Line 266: | Line 169: | ||
|} | |} | ||
=== Phylogeny === | |||
Phylogeny of living Siluriformes based on 2017<ref>{{cite journal |last=Betancur-Rodriguez |first=Ricardo |author2=Edward O. Wiley |author3=Gloria Arratia |author4=Arturo Acero |author5=Nicolas Bailly |author6=Masaki Miya |author7=Guillaume Lecointre |author8=Guillermo Ortí |year=2017 |title=Phylogenetic classification of bony fishes |journal=BMC Evolutionary Biology |edition=4 |volume=17 |issue=162 |pages=162 |bibcode=2017BMCEE..17..162B |doi=10.1186/s12862-017-0958-3 |pmc=5501477 |pmid=28683774 |doi-access=free}}</ref> and extinct families based on Nelson, Grande & Wilson 2016.<ref>{{cite book |last=Nelson |first=Joseph S. |title=Fishes of the World |author2=Terry C. Grande |author3=Mark V. H. Wilson |publisher=John Wiley & Sons |year=2016 |isbn=9781118342336 |edition=5}}</ref> | |||
{{clade| style=font-size:100%;line-height:80% | |||
|label1=Siluriformes | |||
|1={{clade | |||
|1=†] | |||
|2={{clade | |||
|label1=Loricaroidei | |||
|1={{clade | |||
|1={{clade | |||
|1=] ] | |||
|2=] <span style="{{MirrorH}}">]</span> | |||
}} | |||
|2={{clade | |||
|1=] ] | |||
|2={{clade | |||
|1=] ] | |||
|2=] ] | |||
}} | |||
}} | |||
}} | |||
|2={{clade | |||
|label1=Diplomystoidei | |||
|1={{clade | |||
|1=] | |||
|2=†] | |||
}} | |||
|label2=Siluroidei | |||
|2={{clade | |||
|label1=Hypsidoroidea | |||
|1=†] | |||
|2={{clade | |||
|label1=Cetopsoidea | |||
|1=] ] | |||
|2={{clade | |||
|label1=Siluroidea | |||
|1=] ] | |||
|2={{clade | |||
|1={{clade | |||
|1={{clade | |||
|label1=Arioidea | |||
|1=] | |||
|label2=Big African<br>catfishes | |||
|2={{clade | |||
|1=] <span style="{{MirrorH}}">]</span> | |||
|2=] | |||
}} | |||
}} | |||
|2={{clade | |||
|label1=Ictaluroidea | |||
|1={{clade | |||
|1=] | |||
|2=] ] }} | |||
|2={{clade | |||
|label1=Clarioidea | |||
|1=] ] | |||
|label2=Sisoroidea | |||
|2={{clade | |||
|1=] | |||
|2={{clade | |||
|1=] ] | |||
|2=] <span style="{{MirrorH}}">]</span> | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
|2={{clade | |||
|label1=Doradoidea | |||
|1={{clade | |||
|1=] | |||
|2={{clade | |||
|1=] <span style="{{MirrorH}}">]</span> | |||
|2=] ] | |||
}} | |||
}} | |||
|label2=Pimelodoidea | |||
|2={{clade | |||
|1=] <span style="{{MirrorH}}">]</span> | |||
|2={{clade | |||
|1=] ] | |||
|2=] ] | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
}} | |||
Unassigned families: | |||
* ]] | |||
* ] (Loricarioidei) | |||
* ] (Sisoroidea) | |||
* ] (Sisoroidea) | |||
* ] (Arioidea) | |||
* ] (Arioidea) | |||
* ] (Big African catfishes) | |||
* ] (Arioidea) | |||
* ] (Siluroidei) | |||
* '']'' (Pimelodoidea) | |||
* ] (Ictaluroidea) | |||
* ] (Clarioidea) | |||
* ] (Sisoroidea) | |||
* ] (Siluroidea) | |||
* ] (Big African catfishes) | |||
* ] (Big African catfishes) | |||
* ] (Pimelodoidea) | |||
* '']'' (Sisoroidea) | |||
* ] (Big African catfishes) | |||
==Ecology== | |||
===Distribution and habitat=== | |||
Extant catfish species live inland or in coastal waters of every continent except ]. Catfish have inhabited all continents at one time or another.<ref name="Nelson"/> They are most diverse in ] South America, Asia, and Africa, with one family native to North America and one family in Europe.<ref name=tol/> More than half of all catfish species live in the Americas. They are the only ]s that have entered ] habitats in ], Australia, and ].<ref name="Bruton">{{cite journal|title=Alternative life-history strategies of catfishes|last=Bruton|first=Michael N.|journal=Aquat. Living Resour.|year=1996|volume=9|pages=35–41|doi=10.1051/alr:1996040|s2cid=85428351 |doi-access=free}}</ref> | |||
They are found in fresh water/] environments, though most inhabit shallow, running water.<ref name="Bruton"/> Representatives of at least eight families are ] (live underground) with three families that are also ] (inhabiting caves).<ref>{{cite journal|title=Morphological Adaptations of the Texas Blind Catfishes ''Trogloglanis pattersoni'' and ''Satan eurystomus'' (Siluriformes: Ictaluridae) to Their Underground Environment|first=Thomas G.|last=Langecker|author2=Longley, Glenn|journal=]|year=1993|pages=976–986|doi=10.2307/1447075|volume=1993|issue=4|jstor=1447075}}</ref><ref>{{cite journal|title=Mexican blindcats genus ''Prietella'' (Siluriformes: Ictaluridae): an overview of recent explorations|first=Dean A.|last=Hendrickson|author2=Krejca, Jean K. |author3=Martinez, Juan Manuel Rodríguez |journal=Environmental Biology of Fishes|volume=62|pages=315–337|year=2001|issue=1–3|doi=10.1023/A:1011808805094|bibcode=2001EnvBF..62..315H |s2cid=19962442}}</ref> One such species is '']'', known to live underground in ] habitats.<ref name=pcisternarum>{{FishBase|genus=Phreatobius|species=cisternarum|year=2007|month=Apr}}</ref> Numerous species from the families ] and ], and a few species from among the ] and ], are found in salt water.<ref>Monks N. (editor): ''Brackish Water Fishes'', TFH 2006, {{ISBN|0-7938-0564-3}}</ref><ref>Schäfer F: ''Brackish Water Fishes'', Aqualog 2005, {{ISBN|3-936027-82-X}}</ref> | |||
In the Southern United States, catfish species may be known by a variety of slang names, such as "mud cat", "polliwogs", or "chuckleheads".<ref>{{cite web| url = http://www.tpwd.state.tx.us/publications/pwdpubs/media/pwd_br_t3200_0236.pdf| title = Texas Dept. Wildlife| access-date = 3 January 2012| archive-date = 21 February 2012| archive-url = https://web.archive.org/web/20120221021704/https://www.tpwd.state.tx.us/publications/pwdpubs/media/pwd_br_t3200_0236.pdf| url-status = live}}</ref> These nicknames are not standardized, so one area may call a bullhead catfish by the nickname "chucklehead", while in another state or region, that nickname refers to the blue catfish.<ref>{{Cite web |title=Catfish |url=https://www.friendsofwoodlandpark.org/catfish |access-date=2023-06-14 |website=Friends of Woodland Park |language=en-US |archive-date=14 June 2023 |archive-url=https://web.archive.org/web/20230614015117/https://www.friendsofwoodlandpark.org/catfish |url-status=live }}</ref> | |||
===As invasive species=== | |||
Representatives of the ] '']'' have been introduced into European waters in the hope of obtaining a sporting and food resource, but the European stock of American catfishes has not achieved the dimensions of these fish in their native waters and have only increased the ecological pressure on native European ]. ] have also been introduced in the freshwater areas of Florida, with the voracious catfish becoming a major alien pest there. ], ''Pylodictis olivaris'', is also a North American pest on Atlantic slope drainages.<ref name=tol/> '']'' species, released by aquarium fishkeepers, have also established ] populations in many warm waters around the world.<ref>{{cite journal|title=The South American Suckermouth Armored Catfish, ''Pterygoplichthys anisitsi'' (Pisces: Loricaridae), in Texas, with Comments on Foreign Fish Introductions in the American Southwest|first=Leo G.|last=Nico|author2=Martin, R. Trent|journal=The Southwestern Naturalist|volume=46|issue=1|year=2001|pages=98–104|doi=10.2307/3672381|jstor=3672381}}</ref><ref>{{cite journal|title=Amazon Sailfin Catfish, ''Pterygoplichthys pardalis'' (Castelnau, 1855) (Loricariidae), Another Exotic Species Established in Southeastern Mexico|first=Armando T.|last=Wakida-Kusunokia|author2=Ruiz-Carusb, Ramon |author3=Amador-del-Angelc, Enrique |journal=The Southwestern Naturalist|pages=141–144|volume=52|issue=1|doi=10.1894/0038-4909(2007)522.0.CO;2|year=2007|s2cid=86847378 }}</ref><ref>{{cite journal|url=http://www.mapress.com/zootaxa/2006f/zt01109p068.pdf |archive-url=https://web.archive.org/web/20061031235535/http://www.mapress.com/zootaxa/2006f/zt01109p068.pdf |archive-date=2006-10-31 |url-status=live|title=New Philippine record of South American sailfin catfishes (Pisces: Loricariidae)|first=Joel M.|last=Chavez|author2=de la Paz, Reynaldo M.|author3= Manohar, Surya Krishna|author4= Pagulayan, Roberto C.|author5= Carandang Vi, Jose R.|journal=]|volume=1109|pages=57–68|year=2006|access-date=22 June 2009|doi=10.11646/zootaxa.1109.1.6}}</ref><ref>{{cite journal|url=http://academic.uprm.edu/publications/cjs/VOL30/P090-094.PDF|archive-url=https://web.archive.org/web/20090304213101/http://academic.uprm.edu/publications/cjs/VOL30/P090-094.PDF|archive-date=4 March 2009|title=The South American Sailfin Armored Catfish, ''Liposarcus multiradiatus'' (Hancock), a New Exotic Established in Puerto Rican Fresh Waters|first=Lucy|last=Bunkley-Williams|author2=Williams, Ernest H. Jr.|author3= Lilystrom, Craig G.|author4= Corujo-Flores, Iris|author5= Zerbi, Alfonso J.|author6= Aliaume, Catherine|author7= Churchill, Timothy N.|journal=Caribbean Journal of Science|volume=30|issue=1–2|pages=90–94|year=1994|access-date=22 June 2009}}</ref><ref>{{cite journal|url=http://zoolstud.sinica.edu.tw/Journals/44.2/252.pdf |archive-url=https://web.archive.org/web/20060506010509/http://zoolstud.sinica.edu.tw/Journals/44.2/252.pdf |archive-date=2006-05-06 |url-status=live|journal=Zoological Studies|volume=44|issue=2|pages=252–259|year=2005|title=Size Structure, Reproductive Phenology, and Sex Ratio of an Exotic Armored Catfish (''Liposarcus multiradiatus'') in the Kaoping River of Southern Taiwan|first=Shih-Hsiung|last=Liang|author2=Wu, Hsiao-Ping |author3=Shieh, Bao-Sen |access-date=22 June 2009}}</ref> | |||
==Physical characteristics== | |||
===External anatomy of catfish=== | |||
Most catfish are ]s. In general, they are negatively ], which means that they usually sink rather than float due to a reduced ] and a heavy, bony head.<ref name="Bruton"/> Catfish have a variety of body shapes, though most have a cylindrical body with a flattened ] to allow for benthic feeding.<ref name="Bruton"/> | |||
A flattened head allows for digging through the substrate, as well as perhaps serving as a ]. Some have a mouth that can expand to a large size and contains no ] teeth; catfish generally feed through ] or gulping rather than biting and cutting prey.<ref name="Bruton"/> Some families, though, notably the ] and ], have a ] that allows them to fasten themselves to objects in fast-moving water. Catfish also have a ] reduced to a support for ]; this means that they are unable to protrude their mouths as other fish such as ].<ref name="Bruton"/> | |||
] has four pairs of ].]] | |||
Catfish may have up to four pairs of barbels - nasal, maxillary (on each side of mouth), and two pairs of chin barbels, though pairs of barbels may be absent depending on the species. Catfish barbels always occur in pairs. Many larger catfish also have ]s across their entire bodies, which means they "taste" anything they touch and "smell" any chemicals in the water. "In catfish, ] plays a primary role in the orientation and location of food".<ref>Atema, Jelle (1980) pp. 57–101. In: Bardach, JE ''Fish behavior and its use in the capture and culture of fishes'', The WorldFish Center, {{ISBN|978-971-02-0003-0}}.</ref> Because their barbels and chemoreception are more important in detecting food, the eyes on catfish are generally small. Like other ]s, they are characterized by the presence of a ].<ref name="Nelson"/> Their well-developed Weberian apparatus and reduced gas bladder allow for improved ] and sound production.<ref name="Bruton"/> | |||
Catfish do not have ]s; their bodies are often naked. In some species, their ]-covered ] is used in ], where the fish breathes through its skin.<ref name="Bruton"/> In some catfish, the skin is covered in bony plates called ]s; some form of body armor appears in various ways within the order. In ] and in the Asian genus '']'', the armor is primarily made up of one or more rows of free ] plates. Similar plates are found in large specimens of '']''. These plates may be supported by ]l ]es, as in ] and in ''Sisor'', but the processes never fuse to the plates or form any external armor. By contrast, in the subfamily Doumeinae (family ]) and in hoplomyzontines (]), the armor is formed solely by expanded vertebral processes that form plates. Finally, the lateral armor of ], ''Sisor'', and hoplomyzontines consists of hypertrophied ] ossicles with dorsal and ventral ].<ref>{{cite journal|author1=Friel, J P |author2=Lundberg, J G |year=1996|title=''Micromyzon akamai'', gen. et sp. nov., a small and eyeless banjo catfish (Siluriformes: Aspredinidae) from the river channels of the lower Amazon basin|journal=]|issue=3|pages=641–648|jstor=1447528|volume=1996|doi=10.2307/1447528}}</ref> | |||
All catfish other than members of the ] (]), possess a strong, hollow, bony, leading spine-like ray on their ] and ]s. As a defense, these spines may be locked into place so that they stick outwards, enabling them to inflict severe wounds.<ref name=tol/> In numerous catfish species, these fin rays can be used to deliver a stinging ] if the fish is irritated;<ref name=fin>{{cite web| url = http://www.fcps.k12.va.us/StratfordLandingES/Ecology/mpages/channel_catfish.htm| archive-url = https://web.archive.org/web/20060603225626/http://www.fcps.k12.va.us/StratfordLandingES/Ecology/mpages/channel_catfish.htm| archive-date = 3 June 2006|title=Channel Catfish |access-date=2 December 2006|publisher=Fairfax County Public Schools}}</ref> as many as half of all catfish species may be venomous in this fashion, making the Siluriformes overwhelmingly the vertebrate order with the largest number of venomous species.<ref>{{cite journal |last1=Wright |first1=Jeremy J |title=Diversity, phylogenetic distribution, and origins of venomous catfishes |journal=BMC Evolutionary Biology |date=4 December 2009 |volume=9 |issue=1 |pages=282 |doi=10.1186/1471-2148-9-282 |pmid=19961571 |pmc=2791775 |bibcode=2009BMCEE...9..282W |doi-access=free }}</ref> This ] is produced by ]ular cells in the ] tissue covering the spines.<ref name="Nelson"/> In members of the family ] and of the genus '']'', this protein is so strong it may hospitalize humans who receive a sting; in '']'', the stings can be lethal.<ref name="Nelson"/> The dorsal- and pectoral-fin spines are two of the most conspicuous features of siluriforms, and differ from those in other fish groups.<ref name='Ballen'>{{cite journal|author1=Ballen, Gustavo A.|author2=De Pinna, Mario C. C.|title=A standardized terminology of spines in the order Siluriformes (Actinopterygii: Ostariophysi)|year=2022|journal=Zoological Journal of the Linnean Society|volume=194|issue=2|pages=601–625|doi=10.1093/zoolinnean/zlab008|url=https://doi.org/10.1093/zoolinnean/zlab008|access-date=10 February 2022|archive-date=23 September 2023|archive-url=https://web.archive.org/web/20230923095540/https://academic.oup.com/zoolinnean/article/194/2/601/6191677|url-status=live}}</ref> Despite the widespread use of the spines for taxonomic and phylogenetic studies the fields have struggled to effectively use the information due to a lack of consistency in the nomenclature, with a general standard for the descriptive anatomy of catfish spines proposed in 2022 to try and resolve this problem.<ref name='Ballen'/> | |||
Juvenile catfish, like most fish, have relatively large heads, eyes, and posterior median fins in comparison to larger, more mature individuals. These juveniles can be readily placed in their families, particularly those with highly derived fin or body shapes; in some cases, identification of the genus is possible. As far as known for most catfish, features that are often characteristic of species, such as mouth and fin positions, fin shapes, and barbel lengths, show little difference between juveniles and adults. For many species, pigmentation pattern is also similar in juveniles and adults. Thus, juvenile catfish generally resemble and develop smoothly into their adult form without distinct juvenile specializations. Exceptions to this are the ariid catfish, where the young retain yolk sacs late into juvenile stages, and many pimelodids, which may have elongated barbels and fin filaments or coloration patterns.<ref>{{cite journal|title=First description of small juveniles of the primitive catfish ''Diplomystes'' (Siluriformes: Diplomystidae)|first=John G.|last=Lundberg|author2=Berra, Tim M.|author3=Friel, John P.|journal=Ichthyol. Explor. Freshwaters|volume=15|issue=1|pages=71–82|year=2004|url=https://mansfield.osu.edu/assets/mansfield/tberra/pdf/Diplomystes.pdf|access-date=27 March 2023|archive-date=11 December 2021|archive-url=https://web.archive.org/web/20211211152811/https://mansfield.osu.edu/assets/mansfield/tberra/pdf/Diplomystes.pdf|url-status=live}}</ref> | |||
] is reported in about half of all families of catfish.<ref>{{cite journal|url=http://www.mapress.com/zootaxa/2006f/zt01125p056.pdf |archive-url=https://web.archive.org/web/20061031234339/http://www.mapress.com/zootaxa/2006f/zt01125p056.pdf |archive-date=2006-10-31 |url-status=live|title=''Synodontis acanthoperca'', a new species from the Ogôoué River system, Gabon with comments on spiny ornamentation and sexual dimorphism in mochokid catfishes (Siluriformes: Mochokidae)|first=John P.|last=Friel|author2=Vigliotta, Thomas R.|journal=]|volume=1125|pages=45–56|year=2006|access-date=22 June 2009|doi=10.11646/zootaxa.1125.1.3}}</ref> The modification of the ] into an ] (in internal fertilizers) as well as accessory structures of the reproductive apparatus (in both internal and external fertilizers) have been described in species belonging to 11 different families.<ref name="Mazzoldi">{{cite journal|title=Variation of male reproductive apparatus in relation to fertilization modalities in the catfish families Auchenipteridae and Callichthyidae (Teleostei: Siluriformes)|first=C.|last=Mazzoldi|author2=Lorenzi, V. |author3=Rasotto, M. B. |journal=Journal of Fish Biology|year=2007|volume=70|issue=1 |pages=243–256|doi=10.1111/j.1095-8649.2006.01300.x|bibcode=2007JFBio..70..243M }}</ref> | |||
===Size=== | |||
]'' (goonch) caught in India. Some goonch in the Kali River grow large enough to attack humans and water buffalo]] | |||
Catfish have one of the largest ranges in size within a single order of ].<ref name="Bruton"/> Many catfish have a maximum length of under {{convert|12|cm|in|abbr=on}}.<ref name="Nelson" /> Some of the smallest species of the ] and ] reach sexual maturity at only {{convert|1|cm|in|abbr=on}}.<ref name=tol/> | |||
The ], ''Silurus glanis'', and the much smaller related ], are the only catfish indigenous to ]; the former ranges throughout Europe, and the latter is restricted to ]. ] and literature record wels catfish of astounding proportions yet are to be proven scientifically. The typical size of the species is about {{convert|1.2–1.6|m|ft|abbr=on}}, and fish more than {{convert|2|m|ft|abbr=on}} are rare. However, they are known to exceed {{convert|2.5|m|ft|abbr=on}} in length and {{convert|100|kg|lb|abbr=on}} in weight. In July 2009, a catfish weighing {{convert|88|kg|lb}} was caught in the ], Spain, by an 11-year-old British schoolgirl.<ref>{{cite news| url=https://www.telegraph.co.uk/news/newstopics/howaboutthat/5834754/Schoolgirl-nets-9ft-monster-fish.html |archive-url=https://ghostarchive.org/archive/20220111/https://www.telegraph.co.uk/news/newstopics/howaboutthat/5834754/Schoolgirl-nets-9ft-monster-fish.html |archive-date=11 January 2022 |url-access=subscription |url-status=live | work=] | location=London | title=Schoolgirl nets 9ft monster fish | date=15 July 2009 | access-date=28 April 2010}}{{cbignore}}</ref> | |||
In North America, the largest '']'' (blue catfish) caught in the ] on 20 July 2010, weighed {{convert|59|kg|lb|abbr=on}}. The largest ], ''Pylodictis olivaris'', ever caught was in ], weighing {{convert|56|kg|lb|abbr=on}}. | |||
These records pale in comparison to a ] caught in northern ] on 1 May 2005, and reported to the press almost 2 months later, that weighed {{convert|293|kg|lb}}. This is the largest giant Mekong catfish caught since Thai officials started keeping records in 1981.<ref name=Mekong>{{cite web|url=http://news.nationalgeographic.com/news/2005/06/0629_050629_giantcatfish.html|title=Grizzly Bear-Size Catfish Caught in Thailand|date=29 June 2005|access-date=14 July 2006|publisher=National Geographic News|archive-date=30 June 2005|archive-url=https://web.archive.org/web/20050630233848/http://news.nationalgeographic.com/news/2005/06/0629_050629_giantcatfish.html|url-status=dead}}</ref> Also in Asia, ] caught a {{convert|75.5|kg|lb|1|adj=on}} ] following ] in the ] on the ]-] border. Wade was of the opinion that the offending fish must have been significantly larger than this to have taken an 18-year-old boy, as well as a ].{{citation needed|reason=This claim needs a reliable source.|date=March 2020}} | |||
Piraíba ''(])'' can grow exceptionally large and are native to the Amazon Basin. They can occasionally grow to {{convert|200|kg|lb|abbr=on}}, as evidenced by numerous catches. Deaths from being swallowed by these fish have been reported in the region. | |||
===Internal anatomy=== | |||
]'' (glass catfish) have transparent bodies lacking both scales and pigments. Most of the internal organs are located near the head.]] | |||
In many catfish, the "humeral process" is a bony process extending backward from the ] immediately above the base of the pectoral fin. It lies beneath the skin, where its outline may be determined by dissecting the skin or probing with a needle.<ref>{{cite web|url=http://filaman.ifm-geomar.de/Glossary/Glossary.cfm?TermEnglish=humeral%20process|archive-url=https://web.archive.org/web/20071217212539/http://filaman.ifm-geomar.de/Glossary/Glossary.cfm?TermEnglish=humeral%20process|archive-date=17 December 2007|title=Term : humeral process|publisher=]|year=2007}}</ref> | |||
The ]e of catfish are composed of single ]s and large ]s. Many catfish have a ], which may help enhance ] capture and increase low-light sensitivity. ]s, though present in most ]s, are absent from catfish.<ref name="Douglas">{{cite journal|last=Douglas|first=Ron H.|author2=Collin, Shaun P.|author3=Corrigan, Julie|date=15 November 2002|url=http://jeb.biologists.org/cgi/reprint/205/22/3425|title=The eyes of suckermouth armoured catfish (Loricariidae, subfamily Hypostomus): pupil response, lenticular longitudinal spherical aberration and retinal topography|publisher=The Journal of Experimental Biology|volume=205|issue=22|pages=3425–3433|journal=Journal of Experimental Biology|doi=10.1242/jeb.205.22.3425|pmid=12364396|access-date=9 June 2007|archive-date=30 September 2007|archive-url=https://web.archive.org/web/20070930044830/http://jeb.biologists.org/cgi/reprint/205/22/3425|url-status=live}}</ref> | |||
The anatomical organization of the ] in catfish is variable among the families of catfish, but the majority of them present fringed testis: Ictaluridae, Claridae, Auchenipteridae, Doradidae, Pimelodidae, and Pseudopimelodidae.<ref name="Barros"/> In the testes of some species of Siluriformes, organs and structures such as a spermatogenic cranial region and a secretory caudal region are observed, in addition to the presence of seminal vesicles in the caudal region.<ref name="Brito"/> The total number of fringes and their length are different in the ] and ] portions between species.<ref name="Barros">{{cite journal|title=Reproductive apparatus and gametogenesis of ''Lophiosilurus alexandri'' Steindachner (Pisces, Teleostei, Siluriformes)|first=Marcelo D. M.|last=Barros|author2=Guimarães-Cruz, Rodrigo J. |author3=Veloso-Júnior, Vanderlei C. |author4= Santos, José E. dos |journal=Revista Brasileira de Zoologia|volume=24|issue=1|pages=213–221|year=2007|doi=10.1590/S0101-81752007000100028|doi-access=free}}</ref> Fringes of the caudal region may present tubules, in which the lumen is filled by secretion and ].<ref name="Barros"/> Spermatocysts are formed from cytoplasmic extensions of ]s; the release of spermatozoa is allowed by breaking of the cyst walls.<ref name="Barros"/> | |||
The occurrence of ]s, in spite of their interspecific variability in size, gross morphology, and function, has not been related to the mode of fertilization. They are typically paired, multichambered, and connected with the ], and have been reported to play glandular and storage functions. Seminal vesicle secretion may include ]s and steroid glucuronides, with hormonal and pheromonal functions, but it appears to be primarily constituted of mucoproteins, acid mucopolysaccharides, and phospholipids.<ref name="Mazzoldi"/> | |||
Fish ovaries may be of two types - gymnovarian or cystovarian. In the first type, the oocytes are released directly into the ]ic cavity and then eliminated. In the second type, the oocytes are conveyed to the exterior through the ].<ref name="Brito"/> Many catfish are cystovarian in type, including '']'', '']'', '']'', and '']''.<ref name="Barros"/><ref name="Brito">{{cite journal|title=Reproduction of the surubim catfish (Pisces, Pimelodidae) in the São Francisco River, Pirapora Region, Minas Gerais, Brazil|last=Brito|first=M.F.G.|author2=Bazzoli, N.|journal=Arquivo Brasileiro de Medicina Veterinária e Zootecnia|volume=55|issue=5|year=2003|doi=10.1590/S0102-09352003000500018|page=624|doi-access=free}}</ref> | |||
== Communication == | |||
Catfish can produce different types of sounds and also have well-developed auditory reception used to discriminate between sounds with different pitches and velocities. They are also able to determine the distance of the sound's origin and from what direction it originated.<ref name="General Sounds Kasumayan">{{cite journal|last=Kasumayan|first=A. O.|title=Sounds and Sound Production in Fishes|journal=Journal of Ichthyology|year=2008|volume=48|issue=11|pages=981–1030|doi=10.1134/S0032945208110039|s2cid=23223714}}</ref> This is a very important fish communication mechanism, especially during ] and distress behaviors. Catfish are able to produce a variety of sounds for communication that can be classified into two groups: drumming sounds and ] sounds. The variability in catfish sound signals differs due to a few factors: the mechanism by which the sound is produced, the function of the resulting sound, and physiological differences such as size, sex, and age.<ref name="Stridulatory">{{cite journal|last=Vance|first=Theresa L.|title=Variation in Stridulatory Sound Production in the Channel Catfish, ''Ictalurus punctatus''|jstor=4608557|journal=BIOS|year=2000|volume=71|issue=3|pages=79–84}}</ref> To create a drumming sound, catfish use an indirect vibration mechanism using a ]. In these fishes, sonic muscles insert on the ramus Mulleri, also known as the elastic spring. The sonic muscles pull the elastic spring forward and extend the swimbladder. When the muscles relax, the tension in the spring quickly returns the swimbladder to its original position, which produces the sound.<ref name="Sound Generating Mechs">{{cite journal|last=Ladich|first=Friedrich|author2=Michael L. Fine|title=Sound-Generating Mechanisms in Fishes: A Unique Diversity in Vertebrates|journal=Communication in Fishes|year=2006|volume=1|pages=3–43}}</ref> | |||
Catfish also have a sound-generating mechanism in their ]. Many species in the catfish family possess an enhanced first pectoral fin ray, called the spine, which can be moved by large ] and ] muscles. The base of the catfishes' spines has a sequence of ridges, and the spine normally slides within a groove on the fish's pelvic girdle during routine movement; but, pressing the ridges on the spine against the pelvic girdle groove creates a series of short pulses.<ref name="General Sounds Kasumayan" /><ref name="Sound Generating Mechs" /> The movement is analogous to a finger moving down the teeth of a comb, and consequently a series of sharp taps is produced.<ref name="Stridulatory" /> | |||
Sound-generating mechanisms are often different between the sexes. In some catfish, pectoral fins are longer in males than in females of similar length, and differences in the characteristic of the sounds produced were also observed.<ref name="Sound Generating Mechs" /> Comparison between families of the same order of catfish demonstrated family and species-specific patterns of vocalization, according to a study by Maria Clara Amorim. During courtship behavior in three species of ''Corydoras'' catfish, all males actively produced stridulation sounds before egg fertilization, and the species' songs were different in pulse number and sound duration.<ref name="Amorim article">{{cite journal|last=Amorim|first=Maria Clara P.|title=Diversity of Sound Production in Fish|journal=Communication in Fish|year=2006|volume=1|pages=71–105}}</ref> | |||
Sound production in catfish may also be correlated with fighting and alarm calls. According to a study by Kaatz, sounds for disturbance (e.g. alarm) and agonistic behavior were not significantly different, which suggests distress sounds can be used to sample variation in agonistic sound production.<ref name="Amorim article" /> However, in a comparison of a few different species of tropical catfish, some fish put under distress conditions produced a higher intensity of stridulatory sounds than drumming sounds.<ref name="Agonistic behavior"/> Differences in the proportion of drumming versus stridulation sounds depend on ] constraints, such as different sizes of drumming muscles and pectoral spines. Due to these constraints, some fish may not even be able to produce a specific sound. In several different species of catfish, aggressive sound production occurs during cover site defense or during threats from other fish. More specifically, in long-whiskered catfish, drumming sounds are used as a threatening signal and stridulations are used as a defense signal. Kaatz investigated 83 species from 14 families of catfish, and determined that catfish produce more stridulatory sounds in disturbance situations and more swimbladder sounds in intraspecific conflicts.<ref name="Agonistic behavior">{{cite journal|last=Ladich|first=Friedrich|author2=Myrberg, Arthur A Jr.|title=Agonistic Behavior and Acoustic Communication|journal=Communication in Fishes|year=2006|volume=1|pages=121–148}}</ref> | |||
==Economic importance== | |||
===Aquaculture=== | |||
] | |||
{{Main|Aquaculture of catfish}} | |||
Catfish are easy to farm in warm climates, leading to inexpensive and safe food at local grocers. About 60% of U.S. farm-raised catfish are grown within a 65-mile (100-km) radius of ].<ref>{{Cite web | author= Morris, J.E. |title= Pond Culture of Channel Catfish in the North Central Region | publisher= North Central Regional Aquaculture Center| date= October 1993| url= http://www.extension.iastate.edu/Publications/NCR444.pdf| archive-url= https://web.archive.org/web/20070206022434/http://www.extension.iastate.edu/Publications/NCR444.pdf| archive-date= 6 February 2007| access-date= 28 June 2006}}</ref> ] (''Ictalurus punctatus'') supports a $450 million/yr aquaculture industry.<ref name=tol/> The largest producers are located in the ], including ], ], and ].<ref>{{Cite web|url=http://usda.mannlib.cornell.edu/usda/current/CatfProd/CatfProd-07-21-2017.pdf |archive-url=https://web.archive.org/web/20170915113343/http://usda.mannlib.cornell.edu/usda/current/CatfProd/CatfProd-07-21-2017.pdf |archive-date=2017-09-15 |url-status=live|title=Catfish Production|date=21 July 2017|website=www.nass.usda.gov|access-date=14 September 2017}}</ref> | |||
Catfish raised in inland tanks or channels are usually considered safe for the environment, since their waste and disease should be contained and not spread to the wild.<ref>{{cite journal | first = Paul | last = Rogers | title = Economy of Scales | journal = Stanford Magazine | publisher = ] | issue = March / April 2006 | url = http://www.stanfordalumni.org/news/magazine/2006/marapr/features/fishfarming.html | access-date = 14 February 2008 | archive-url = https://web.archive.org/web/20080611123839/http://www.stanfordalumni.org/news/magazine/2006/marapr/features/fishfarming.html | archive-date = 11 June 2008 | url-status = dead}}</ref> | |||
In Asia, many catfish species are important as food. Several ] (Clariidae) and ] (Pangasiidae) species are heavily cultured in Africa and Asia. Exports of one particular shark catfish species from ], '']'', have met with pressures from the U.S. catfish industry. In 2003, The ] passed a law preventing the imported fish from being labeled as catfish.<ref>{{cite web| url = http://www.latimes.com/business/la-fi-catfish28nov28,0,6595048.story?coll=la-home-business| title = "'Catfish' bred in Asia move up on U.S. food chain"$, Associated Press via L.A. Times, 28 November 2006| website = ]| date = 28 November 2006| access-date = 5 December 2006| archive-date = 23 September 2023| archive-url = https://web.archive.org/web/20230923095543/https://www.latimes.com/archives/la-xpm-2006-nov-28-fi-catfish28-story.html| url-status = live}}</ref> As a result, the Vietnamese exporters of this fish now label their products sold in the U.S. as "basa fish." Trader Joe's has labeled frozen fillets of Vietnamese '']'' as "striper."<ref>Cole, Nancy (27 January 2006) . Northwest Arkansas News</ref> | |||
There is a large and growing ornamental fish trade, with hundreds of species of catfish, such as '']'' and ] (often called plecos), being a popular component of many ]. Other catfish commonly found in the aquarium trade are ], ], and ]. | |||
===Catfish as food=== | |||
{{Cookbook|Catfish}}]]] | |||
Catfish have widely been caught and farmed for food for thousands of years in Africa, Asia, Europe, and North America. Judgments as to the quality and flavor vary, with some food critics considering catfish excellent to eat, while others dismiss them as watery and lacking in flavor.<ref name=Baker>Jenny Baker (1988), ''Simply Fish'' p 36–37. Faver & Faber, London.</ref> Catfish is high in ].<ref name = FactD>{{cite web|url=http://www.health.state.ny.us/diseases/conditions/osteoporosis/vitd.htm|title=Vitamin D and Healthy Bones|publisher=New York State Department of Health|access-date=13 July 2007|archive-date=18 August 2010|archive-url=https://web.archive.org/web/20100818173653/http://www.health.state.ny.us/diseases/conditions/osteoporosis/vitd.htm|url-status=live}}</ref> Farm-raised catfish contains low levels of ]s and a much higher proportion of ]s.<ref> {{Webarchive|url=https://web.archive.org/web/20120321233925/http://www.fitfunctional.com/index.php/articles/nutrition/60-fatty-fish-not-equal-in-qgoodq-fats-study |date=21 March 2012 }}. Reuters. Source: Journal of the American Dietetic Association, July 2008</ref> | |||
In ], catfish were often viewed as a ] to be enjoyed on ]s and holidays. Migrants from Europe and Africa to the United States brought along this tradition, and in the ], catfish is an extremely popular food. | |||
The most commonly eaten species in the United States are the ] and the ], both of which are common in the wild and increasingly widely farmed. Farm-raised catfish became such a staple of the U.S. diet that President ] proclaimed National Catfish Day on June 25, 1987, to recognize "the value of farm-raised catfish."<ref>{{Cite web |date=2011-09-07 |title=The Rehab Archipelago {{!}} Forced Labor and Other Abuses in Drug Detention Centers in Southern Vietnam |url=https://www.hrw.org/report/2011/09/07/rehab-archipelago/forced-labor-and-other-abuses-drug-detention-centers-southern |access-date=2019-06-25 |website=Human Rights Watch |language=en}}</ref> | |||
Catfish is eaten in a variety of ways. In Europe, it is often cooked in similar ways to ], but in the United States it is popularly crumbed with ] and fried.<ref name=Baker/> | |||
] served with '']'', ] and '']'' vegetables in a tent '']'' in Jakarta, Indonesia]] | |||
In ], catfish is usually served fried or grilled in street stalls called '']'' and eaten with vegetables, ] (a spicy ] or sauce), and usually '']'' (traditional ]). The dish is called {{lang|id|pecel lele}} or {{lang|id|pecak lele}}. {{lang|id|Lele}} is the ] word for catfish. The same dish can also be called as {{lang|id|lele penyet}} (squashed catfish) if the fish is lightly squashed along with sambal with a stone ]. The {{lang|id|pecel}} or {{lang|id|pecak}} version presents the fish in a separate plate while the mortar is solely for sambal. | |||
In ], catfish is called ''ikan keli'' and is fried with spices or grilled and eaten with ] and ] gravy and is also often eaten with ]. | |||
In ] and the ]n states of ], ] and ], catfish (locally known as ''magur'') is eaten as a favored delicacy during the ]s. In the Indian state of ], the local catfish, known as ''thedu''' or ''etta'' in ], is also popular. | |||
In ], catfish is often cooked in ] sauce (Harcsapaprikás) typical of ]. It is traditionally served with ] smothered with ] (]). | |||
In ] (formally Burma), catfish is usually used in '']'', a traditional noodle ] cooked with ], ], ], pepper, banana stem, onions, and other local ingredients. | |||
] fried ''hito'' (catfish) with vinegar and '']'' dip sauce]] | |||
Vietnamese catfish, of the genus '']'', cannot be legally marketed as catfish in the United States, and so is referred to as '']'' or '']''.<ref name = FactSwai>{{cite web|url=http://www.unionfishco.com/swai/details.htm|title=Union Fish Company – Basa/Swai Details|access-date=11 November 2007 |archive-url = https://web.archive.org/web/20071109150512/http://www.unionfishco.com/swai/details.htm |archive-date = 9 November 2007}}</ref> Only fish of the family '']'' may be marketed as catfish in the United States.<ref name = PL107171>Public Law 107-171, § 10806, 116 ] 526-527, codified in {{cite web|url=https://www.law.cornell.edu/uscode/text/21/321d|title=United States Code, Title 21, section 321d. Market names for catfish and ginseng|mode=cs2|access-date=28 October 2020|archive-date=17 April 2021|archive-url=https://web.archive.org/web/20210417123630/https://www.law.cornell.edu/uscode/text/21/321d|url-status=live}} and {{cite web|url=https://www.law.cornell.edu/uscode/text/21/343-|title=United States Code, Title 21, section 343 (t). Misbranded food|access-date=9 May 2017|archive-date=23 September 2023|archive-url=https://web.archive.org/web/20230923095542/https://www.law.cornell.edu/uscode/text/21/343|url-status=live}}</ref><ref>''See'' '' {{Webarchive|url=https://web.archive.org/web/20230923095542/https://scholar.google.com/scholar_case?case=14897106900736461981&q=ictaluridae |date=23 September 2023 }}'', 448 F. 3d 744 (5th Cir. 2006), ''citing'' Kerrilee E. Kobbeman, "Legislative Note, Hook, Line and Sinker: How Congress Swallowed the Domestic Catfish Industry's Narrow Definition of this Ubiquitous Bottomfeeder," 57 ARK. L.REV. 407, 411-18 (2004).</ref> In the UK, Vietnamese catfish is sometimes sold as "Vietnamese river cobbler", although more commonly as Basa.<ref>{{cite web| url = http://www.legislation.gov.uk/uksi/2006/506/pdfs/uksi_20060506_en.pdf| publisher = ]| title = Fish Labelling (Amendment) (England) Regulations 2006| date = 26 May 2007| access-date = 23 May 2013| archive-date = 31 January 2012| archive-url = https://web.archive.org/web/20120131042544/http://www.legislation.gov.uk/uksi/2006/506/pdfs/uksi_20060506_en.pdf| url-status = live}}</ref> | |||
In ], catfish is often cooked in a variety of ]s. It is particularly cooked in a delicacy popularly known as "catfish pepper soup" which is enjoyed throughout the nation.<ref>{{Cite news|url=http://www.pulse.ng/lifestyle/food-travel-arts-culture/how-to-cook-the-irresistible-catfish-pepper-soup-id8377325.html|title=How to cook the irresistible catfish pepper soup|last=Oreva|first=Duke|date=2018-05-14|access-date=16 May 2018|archive-date=16 May 2018|archive-url=https://web.archive.org/web/20180516175134/http://www.pulse.ng/lifestyle/food-travel-arts-culture/how-to-cook-the-irresistible-catfish-pepper-soup-id8377325.html|url-status=live}}</ref> | |||
In Jewish dietary law, known as ], fish must have fins and scales to be ].<ref>{{Cite web |title=Kosher Spirit: Fins and Scales |url=https://www.ok.org/article/fins-scales/ |access-date=2022-11-29 |website=OK Kosher Certification |archive-date=29 November 2022 |archive-url=https://web.archive.org/web/20221129062913/https://www.ok.org/article/fins-scales/ |url-status=live }}</ref> Since catfish lack scales, they are not kosher.<ref>{{Cite web|url=https://fisheries.tamu.edu/pond-management/species/channel-catfish/|title=Channel Catfish|website=fisheries.tamu.edu|access-date=2019-11-14|archive-date=1 May 2019|archive-url=https://web.archive.org/web/20190501061619/https://fisheries.tamu.edu/pond-management/species/channel-catfish/|url-status=live}}</ref> | |||
==Mythology== | |||
In the mythology of the Japanese ] natural phenomenon are caused by '']''. ] are caused by a giant catfish called ]. There are other ''kami'' associated with earthquakes. In ] it's usually an eel, but after the ] {{Nihongo|'''''Namazu-e'''''|鯰絵||"catfish prints"}} were printed giving more popularity to the catfish ''kami'' that has been known since the 16th century ].<ref>{{cite book |last1=Rosen |first1=Brenda |title=The Mythical Creatures Bible |publisher=Sterling |date=2009 |page=370}}</ref> In one catfish print the divine white horse of ] is depicted knocking down the earthquake-causing catfish.<ref>{{cite book |last1=Smits |first1=Gregory |title=Seismic Japan: The Long History and Continuing Legacy of the Ansei Edo Earthquake |date=2013 |publisher=University of Hawaii Press |page=158|bibcode=2013sjlh.book.....S }}</ref> | |||
==Dangers to humans== | |||
]'', may be fatal.]] | |||
While the vast majority of catfish are harmless to humans, a few species are known to present some risk. Many catfish species have "stings" (actually non-venomous in most cases) embedded behind their fins; thus precautions must be taken when handling them. Stings by the venomous ] have killed people in rare cases.<ref>{{FishBase|genus=Plotosus|species=lineatus|month=November|year=2014}}</ref> | |||
==Catfish fishing records== | |||
By information from International Game Fish Association ] the most outstanding record:<ref>{{cite web |url=http://wrec.igfa.org/ |title=IGFA World Records |publisher=International Game Fish Association |access-date=1 November 2015 |archive-date=1 November 2015 |archive-url=https://web.archive.org/web/20151101064327/http://wrec.igfa.org/ |url-status=live }}</ref> | |||
* The biggest flathead catfish caught was by Ken Paulie in the ] in Kansas, US on 19 May 1998 that weighed {{convert|55.79|kg|lboz|abbr=on}} | |||
==See also== | |||
* ] | |||
==References== | ==References== | ||
{{clear}} | {{clear}} | ||
{{Reflist |
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==External links== | ||
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{{Commons category|Siluriformes}} | {{Commons category|Siluriformes}} | ||
{{Wikispecies|Siluriformes}} | {{Wikispecies|Siluriformes}} | ||
{{EB1911 poster|Cat-fish}} | |||
* Catfish clonk fishing method ] | |||
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* {{Webarchive|url=https://web.archive.org/web/20141129032305/http://archive.thedailystar.net/newDesign/news-details.php?nid=87845 |date=29 November 2014 }} in ''The Daily Star'' (Bangladesh), 12 May 2009 | |||
* Skelton, Paul H. and Teugels, Guy G. 1992. . J.L.B. Smith Institute of Ichthyology, Rhodes University, Grahamstown, South Africa | |||
{{Actinopterygii}} | {{Actinopterygii}} | ||
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Revision as of 02:19, 6 December 2024
Order of fish For the deceptive online actions by one to another, see Catfishing. This article is about the fish. For other uses, see Catfish (disambiguation). "Kaari" redirects here. For other uses, see Kaari (disambiguation).
Catfish Temporal range: Campanian–Present PreꞒ Ꞓ O S D C P T J K Pg N Possible earlier Late Cretaceous records | |
---|---|
Black bullhead | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Actinopterygii |
(unranked): | Otophysi |
Order: | Siluriformes G. Cuvier, 1817 |
Type species | |
Silurus glanis Linnaeus, 1758 | |
Families | |
Extant families:
Extinct family: |
Catfish (or catfishes; order Siluriformes /sɪˈljʊərɪfɔːrmiːz/ or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive, the Mekong giant catfish from Southeast Asia, the wels catfish of Eurasia, and the piraíba of South America, to detritivores (species that eat dead material on the bottom), and even to a tiny parasitic species commonly called the candiru, Vandellia cirrhosa. Neither the armour-plated types nor the naked types have scales. Despite their name, not all catfish have prominent barbels or "whiskers". Members of the Siluriformes order are defined by features of the skull and swimbladder. Catfish are of considerable commercial importance; many of the larger species are farmed or fished for food. Many of the smaller species, particularly the genus Corydoras, are important in the aquarium hobby. Many catfish are nocturnal, but others (many Auchenipteridae) are crepuscular or diurnal (most Loricariidae or Callichthyidae, for example).
Taxonomy
Molecular evidence suggests that in spite of the great morphological diversity in the order, all catfish form a monophyletic group. Catfish belong to a superorder called the Ostariophysi, which also includes the Cypriniformes (carps and minnows), Characiformes (characins and tetras), Gonorynchiformes (milkfish and beaked salmons) and Gymnotiformes (South American knifefish), a superorder characterized by the Weberian apparatus. Some place Gymnotiformes as a sub-order of Siluriformes; however, this is not as widely accepted. Currently, the Siluriformes are said to be the sister group to the Gymnotiformes, though this has been debated due to more recent molecular evidence. As of 2007 there were about thirty-six extant catfish families, and about 3,093 extant species have been described. This makes the catfish order the second or third most diverse vertebrate order; in fact, one out of every twenty vertebrate species is a catfish.
Catfish are believed to have a Gondwanan origin primarily centered around South America, as the most basal living catfish groups are known from there. The earliest known definitive members lived in the Americas from the Campanian to Maastrichtian stages of the Late Cretaceous, including the Andinichthyidae, Vorhisia vulpes and possibly Arius. A potential fossil record is known from the earlier Coniacian-Santonian stages in Niger of West Africa, though this has been considered unreliable, and the putative earliest armored catfish known from the fossil record, Afrocascudo, lived during the Cenomanian age of the Late Cretaceous in Morocco of North Africa (Kem Kem Group). The describers of Afrocascudo claimed that the presence of a derived loricariid so early on would indicate the extensive diversification of catfish, or at least loricarioids, prior to the beginning of the Late Cretaceous. As extant loricariids are only known from South America, much of this diversification must have occurred on the supercontinent of West Gondwana prior to its fragmentation into South America and Africa. Britz and colleagues suggested that Afrocascudo instead represents a juvenile obaichthyid lepisosteiform, possibly a junior synonym of Obaichthys. The authors of the original study still stood by their original conclusion based on the absence of important holostean characters, and noted that it could not be a juvenile, since the bones were completely ossified.
The taxonomy of catfish is quickly changing. In a 2007 and 2008 paper, Horabagrus, Phreatobius, and Conorhynchos were not classified under any current catfish families. There is disagreement on the family status of certain groups; for example, Nelson (2006) lists Auchenoglanididae and Heteropneustidae as separate families, while the All Catfish Species Inventory (ACSI) includes them under other families. FishBase and the Integrated Taxonomic Information System lists Parakysidae as a separate family, while this group is included under Akysidae by both Nelson (2006) and ACSI. Many sources do not list the recently revised family Anchariidae. The family Horabagridae, including Horabagrus, Pseudeutropius, and Platytropius, is not shown by some authors but presented by others as a true group. Thus, the actual number of families differs between authors. The species count is in constant flux due to taxonomic work as well as description of new species. Between 2003 and 2005, over one hundred species were named, a rate three times faster than that of the past century. In June 2005, researchers named the newest family of catfish, Lacantuniidae, only the third new family of fish distinguished in the last seventy years, the others being the coelacanth in 1938 and the megamouth shark in 1983. The new species in Lacantuniidae, Lacantunia enigmatica, was found in the Lacantun river in the Mexican state of Chiapas.
The higher-level phylogeny of Siluriformes has gone through several recent changes, mainly due to molecular phylogenetic studies. While most studies, both morphological and molecular, agree that catfishes are arranged into three main lineages, the relationship among these lineages has been a contentious point in which these studies, performed for example by Rui Diogo, differ. The three main lineages in Siluriformes are the family Diplomystidae, the denticulate catfish suborder Loricarioidei (containing the Neotropical "suckermouth" catfishes), and the suborder Siluroidei, which contains the remaining families of the order. According to morphological data, Diplomystidae is usually considered to be the earliest branching catfish lineage and the sister group to the other two lineages, Loricarioidei and Siluroidei. Molecular evidence usually contrasts with this hypothesis, and shows the suborder Loricarioidei as the earliest branching catfish lineage, and sister to a clade that includes the Diplomystidae and Siluroidei; this phylogeny has been obtained in numerous studies based on genetic data. However, it has been suggested that these molecular results are errors as a result of long branch attraction, incorrectly placing Loricarioidei as the earliest-branching catfish lineage. When a data filtering method was used to reduce lineage rate heterogeneity (the potential source of bias) on their dataset, a final phylogeny was recovered which showed the Diplomystidae are the earliest-branching catfish, followed by Loricarioidei and Siluroidei as sister lineages, providing both morphological and molecular support for Diplomystidae being the earliest branching catfish.
Below is a list of family relationships by different authors. Lacantuniidae is included in the Sullivan scheme based on recent evidence that places it sister to Claroteidae.
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Phylogeny
Phylogeny of living Siluriformes based on 2017 and extinct families based on Nelson, Grande & Wilson 2016.
Siluriformes |
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Unassigned families:
- Bachmanniidae†
- Scoloplacidae (Loricarioidei)
- Akysidae (Sisoroidea)
- Amblycipitidae (Sisoroidea)
- Anchariidae (Arioidea)
- Ariidae (Arioidea)
- Amphiliidae (Big African catfishes)
- Austroglanididae (Arioidea)
- Chacidae (Siluroidei)
- Conorhynchos (Pimelodoidea)
- Cranoglanididae (Ictaluroidea)
- Heteropneustidae (Clarioidea)
- Horabagridae (Sisoroidea)
- Kryptoglanidae (Siluroidea)
- Lacantuniidae (Big African catfishes)
- Malapteruridae (Big African catfishes)
- Phreatobiidae (Pimelodoidea)
- Rita (Sisoroidea)
- Schilbeidae (Big African catfishes)
Ecology
Distribution and habitat
Extant catfish species live inland or in coastal waters of every continent except Antarctica. Catfish have inhabited all continents at one time or another. They are most diverse in tropical South America, Asia, and Africa, with one family native to North America and one family in Europe. More than half of all catfish species live in the Americas. They are the only ostariophysans that have entered freshwater habitats in Madagascar, Australia, and New Guinea.
They are found in fresh water/brackish water environments, though most inhabit shallow, running water. Representatives of at least eight families are hypogean (live underground) with three families that are also troglobitic (inhabiting caves). One such species is Phreatobius cisternarum, known to live underground in phreatic habitats. Numerous species from the families Ariidae and Plotosidae, and a few species from among the Aspredinidae and Bagridae, are found in salt water.
In the Southern United States, catfish species may be known by a variety of slang names, such as "mud cat", "polliwogs", or "chuckleheads". These nicknames are not standardized, so one area may call a bullhead catfish by the nickname "chucklehead", while in another state or region, that nickname refers to the blue catfish.
As invasive species
Representatives of the genus Ictalurus have been introduced into European waters in the hope of obtaining a sporting and food resource, but the European stock of American catfishes has not achieved the dimensions of these fish in their native waters and have only increased the ecological pressure on native European fauna. Walking catfish have also been introduced in the freshwater areas of Florida, with the voracious catfish becoming a major alien pest there. Flathead catfish, Pylodictis olivaris, is also a North American pest on Atlantic slope drainages. Pterygoplichthys species, released by aquarium fishkeepers, have also established feral populations in many warm waters around the world.
Physical characteristics
External anatomy of catfish
Most catfish are bottom feeders. In general, they are negatively buoyant, which means that they usually sink rather than float due to a reduced gas bladder and a heavy, bony head. Catfish have a variety of body shapes, though most have a cylindrical body with a flattened ventrum to allow for benthic feeding.
A flattened head allows for digging through the substrate, as well as perhaps serving as a hydrofoil. Some have a mouth that can expand to a large size and contains no incisiform teeth; catfish generally feed through suction or gulping rather than biting and cutting prey. Some families, though, notably the Loricariidae and Astroblepidae, have a suckermouth that allows them to fasten themselves to objects in fast-moving water. Catfish also have a maxilla reduced to a support for barbels; this means that they are unable to protrude their mouths as other fish such as carp.
Catfish may have up to four pairs of barbels - nasal, maxillary (on each side of mouth), and two pairs of chin barbels, though pairs of barbels may be absent depending on the species. Catfish barbels always occur in pairs. Many larger catfish also have chemoreceptors across their entire bodies, which means they "taste" anything they touch and "smell" any chemicals in the water. "In catfish, gustation plays a primary role in the orientation and location of food". Because their barbels and chemoreception are more important in detecting food, the eyes on catfish are generally small. Like other ostariophysans, they are characterized by the presence of a Weberian apparatus. Their well-developed Weberian apparatus and reduced gas bladder allow for improved hearing and sound production.
Catfish do not have scales; their bodies are often naked. In some species, their mucus-covered skin is used in cutaneous respiration, where the fish breathes through its skin. In some catfish, the skin is covered in bony plates called scutes; some form of body armor appears in various ways within the order. In loricarioids and in the Asian genus Sisor, the armor is primarily made up of one or more rows of free dermal plates. Similar plates are found in large specimens of Lithodoras. These plates may be supported by vertebral processes, as in scoloplacids and in Sisor, but the processes never fuse to the plates or form any external armor. By contrast, in the subfamily Doumeinae (family Amphiliidae) and in hoplomyzontines (Aspredinidae), the armor is formed solely by expanded vertebral processes that form plates. Finally, the lateral armor of doradids, Sisor, and hoplomyzontines consists of hypertrophied lateral line ossicles with dorsal and ventral lamina.
All catfish other than members of the Malapteruridae (electric catfish), possess a strong, hollow, bony, leading spine-like ray on their dorsal and pectoral fins. As a defense, these spines may be locked into place so that they stick outwards, enabling them to inflict severe wounds. In numerous catfish species, these fin rays can be used to deliver a stinging protein if the fish is irritated; as many as half of all catfish species may be venomous in this fashion, making the Siluriformes overwhelmingly the vertebrate order with the largest number of venomous species. This venom is produced by glandular cells in the epidermal tissue covering the spines. In members of the family Plotosidae and of the genus Heteropneustes, this protein is so strong it may hospitalize humans who receive a sting; in Plotosus lineatus, the stings can be lethal. The dorsal- and pectoral-fin spines are two of the most conspicuous features of siluriforms, and differ from those in other fish groups. Despite the widespread use of the spines for taxonomic and phylogenetic studies the fields have struggled to effectively use the information due to a lack of consistency in the nomenclature, with a general standard for the descriptive anatomy of catfish spines proposed in 2022 to try and resolve this problem.
Juvenile catfish, like most fish, have relatively large heads, eyes, and posterior median fins in comparison to larger, more mature individuals. These juveniles can be readily placed in their families, particularly those with highly derived fin or body shapes; in some cases, identification of the genus is possible. As far as known for most catfish, features that are often characteristic of species, such as mouth and fin positions, fin shapes, and barbel lengths, show little difference between juveniles and adults. For many species, pigmentation pattern is also similar in juveniles and adults. Thus, juvenile catfish generally resemble and develop smoothly into their adult form without distinct juvenile specializations. Exceptions to this are the ariid catfish, where the young retain yolk sacs late into juvenile stages, and many pimelodids, which may have elongated barbels and fin filaments or coloration patterns.
Sexual dimorphism is reported in about half of all families of catfish. The modification of the anal fin into an intromittent organ (in internal fertilizers) as well as accessory structures of the reproductive apparatus (in both internal and external fertilizers) have been described in species belonging to 11 different families.
Size
Catfish have one of the largest ranges in size within a single order of bony fish. Many catfish have a maximum length of under 12 cm (4.7 in). Some of the smallest species of the Aspredinidae and Trichomycteridae reach sexual maturity at only 1 cm (0.39 in).
The wels catfish, Silurus glanis, and the much smaller related Aristotle's catfish, are the only catfish indigenous to Europe; the former ranges throughout Europe, and the latter is restricted to Greece. Mythology and literature record wels catfish of astounding proportions yet are to be proven scientifically. The typical size of the species is about 1.2–1.6 m (3.9–5.2 ft), and fish more than 2 m (6.6 ft) are rare. However, they are known to exceed 2.5 m (8.2 ft) in length and 100 kg (220 lb) in weight. In July 2009, a catfish weighing 88 kilograms (194 lb) was caught in the River Ebro, Spain, by an 11-year-old British schoolgirl.
In North America, the largest Ictalurus furcatus (blue catfish) caught in the Missouri River on 20 July 2010, weighed 59 kg (130 lb). The largest flathead catfish, Pylodictis olivaris, ever caught was in Independence, Kansas, weighing 56 kg (123 lb).
These records pale in comparison to a Mekong giant catfish caught in northern Thailand on 1 May 2005, and reported to the press almost 2 months later, that weighed 293 kilograms (646 lb). This is the largest giant Mekong catfish caught since Thai officials started keeping records in 1981. Also in Asia, Jeremy Wade caught a 75.5-kilogram (166.4 lb) goonch following three fatal attacks on humans in the Kali River on the India-Nepal border. Wade was of the opinion that the offending fish must have been significantly larger than this to have taken an 18-year-old boy, as well as a water buffalo.
Piraíba (Brachyplatystoma filamentosum) can grow exceptionally large and are native to the Amazon Basin. They can occasionally grow to 200 kg (440 lb), as evidenced by numerous catches. Deaths from being swallowed by these fish have been reported in the region.
Internal anatomy
In many catfish, the "humeral process" is a bony process extending backward from the pectoral girdle immediately above the base of the pectoral fin. It lies beneath the skin, where its outline may be determined by dissecting the skin or probing with a needle.
The retinae of catfish are composed of single cones and large rods. Many catfish have a tapetum lucidum, which may help enhance photon capture and increase low-light sensitivity. Double cones, though present in most teleosts, are absent from catfish.
The anatomical organization of the testis in catfish is variable among the families of catfish, but the majority of them present fringed testis: Ictaluridae, Claridae, Auchenipteridae, Doradidae, Pimelodidae, and Pseudopimelodidae. In the testes of some species of Siluriformes, organs and structures such as a spermatogenic cranial region and a secretory caudal region are observed, in addition to the presence of seminal vesicles in the caudal region. The total number of fringes and their length are different in the caudal and cranial portions between species. Fringes of the caudal region may present tubules, in which the lumen is filled by secretion and spermatozoa. Spermatocysts are formed from cytoplasmic extensions of Sertoli cells; the release of spermatozoa is allowed by breaking of the cyst walls.
The occurrence of seminal vesicles, in spite of their interspecific variability in size, gross morphology, and function, has not been related to the mode of fertilization. They are typically paired, multichambered, and connected with the sperm duct, and have been reported to play glandular and storage functions. Seminal vesicle secretion may include steroids and steroid glucuronides, with hormonal and pheromonal functions, but it appears to be primarily constituted of mucoproteins, acid mucopolysaccharides, and phospholipids.
Fish ovaries may be of two types - gymnovarian or cystovarian. In the first type, the oocytes are released directly into the coelomic cavity and then eliminated. In the second type, the oocytes are conveyed to the exterior through the oviduct. Many catfish are cystovarian in type, including Pseudoplatystoma corruscans, P. fasciatum, Lophiosilurus alexandri, and Loricaria lentiginosa.
Communication
Catfish can produce different types of sounds and also have well-developed auditory reception used to discriminate between sounds with different pitches and velocities. They are also able to determine the distance of the sound's origin and from what direction it originated. This is a very important fish communication mechanism, especially during agonistic and distress behaviors. Catfish are able to produce a variety of sounds for communication that can be classified into two groups: drumming sounds and stridulation sounds. The variability in catfish sound signals differs due to a few factors: the mechanism by which the sound is produced, the function of the resulting sound, and physiological differences such as size, sex, and age. To create a drumming sound, catfish use an indirect vibration mechanism using a swimbladder. In these fishes, sonic muscles insert on the ramus Mulleri, also known as the elastic spring. The sonic muscles pull the elastic spring forward and extend the swimbladder. When the muscles relax, the tension in the spring quickly returns the swimbladder to its original position, which produces the sound.
Catfish also have a sound-generating mechanism in their pectoral fins. Many species in the catfish family possess an enhanced first pectoral fin ray, called the spine, which can be moved by large abductor and adductor muscles. The base of the catfishes' spines has a sequence of ridges, and the spine normally slides within a groove on the fish's pelvic girdle during routine movement; but, pressing the ridges on the spine against the pelvic girdle groove creates a series of short pulses. The movement is analogous to a finger moving down the teeth of a comb, and consequently a series of sharp taps is produced.
Sound-generating mechanisms are often different between the sexes. In some catfish, pectoral fins are longer in males than in females of similar length, and differences in the characteristic of the sounds produced were also observed. Comparison between families of the same order of catfish demonstrated family and species-specific patterns of vocalization, according to a study by Maria Clara Amorim. During courtship behavior in three species of Corydoras catfish, all males actively produced stridulation sounds before egg fertilization, and the species' songs were different in pulse number and sound duration.
Sound production in catfish may also be correlated with fighting and alarm calls. According to a study by Kaatz, sounds for disturbance (e.g. alarm) and agonistic behavior were not significantly different, which suggests distress sounds can be used to sample variation in agonistic sound production. However, in a comparison of a few different species of tropical catfish, some fish put under distress conditions produced a higher intensity of stridulatory sounds than drumming sounds. Differences in the proportion of drumming versus stridulation sounds depend on morphological constraints, such as different sizes of drumming muscles and pectoral spines. Due to these constraints, some fish may not even be able to produce a specific sound. In several different species of catfish, aggressive sound production occurs during cover site defense or during threats from other fish. More specifically, in long-whiskered catfish, drumming sounds are used as a threatening signal and stridulations are used as a defense signal. Kaatz investigated 83 species from 14 families of catfish, and determined that catfish produce more stridulatory sounds in disturbance situations and more swimbladder sounds in intraspecific conflicts.
Economic importance
Aquaculture
Main article: Aquaculture of catfishCatfish are easy to farm in warm climates, leading to inexpensive and safe food at local grocers. About 60% of U.S. farm-raised catfish are grown within a 65-mile (100-km) radius of Belzoni, Mississippi. Channel catfish (Ictalurus punctatus) supports a $450 million/yr aquaculture industry. The largest producers are located in the Southern United States, including Mississippi, Alabama, and Arkansas.
Catfish raised in inland tanks or channels are usually considered safe for the environment, since their waste and disease should be contained and not spread to the wild.
In Asia, many catfish species are important as food. Several airbreathing catfish (Clariidae) and shark catfish (Pangasiidae) species are heavily cultured in Africa and Asia. Exports of one particular shark catfish species from Vietnam, Pangasius bocourti, have met with pressures from the U.S. catfish industry. In 2003, The United States Congress passed a law preventing the imported fish from being labeled as catfish. As a result, the Vietnamese exporters of this fish now label their products sold in the U.S. as "basa fish." Trader Joe's has labeled frozen fillets of Vietnamese Pangasius hypophthalmus as "striper."
There is a large and growing ornamental fish trade, with hundreds of species of catfish, such as Corydoras and armored suckermouth catfish (often called plecos), being a popular component of many aquaria. Other catfish commonly found in the aquarium trade are banjo catfish, talking catfish, and long-whiskered catfish.
Catfish as food
Catfish have widely been caught and farmed for food for thousands of years in Africa, Asia, Europe, and North America. Judgments as to the quality and flavor vary, with some food critics considering catfish excellent to eat, while others dismiss them as watery and lacking in flavor. Catfish is high in vitamin D. Farm-raised catfish contains low levels of omega-3 fatty acids and a much higher proportion of omega-6 fatty acids.
In Central Europe, catfish were often viewed as a delicacy to be enjoyed on feast days and holidays. Migrants from Europe and Africa to the United States brought along this tradition, and in the Southern United States, catfish is an extremely popular food.
The most commonly eaten species in the United States are the channel catfish and the blue catfish, both of which are common in the wild and increasingly widely farmed. Farm-raised catfish became such a staple of the U.S. diet that President Ronald Reagan proclaimed National Catfish Day on June 25, 1987, to recognize "the value of farm-raised catfish."
Catfish is eaten in a variety of ways. In Europe, it is often cooked in similar ways to carp, but in the United States it is popularly crumbed with cornmeal and fried.
In Indonesia, catfish is usually served fried or grilled in street stalls called warung and eaten with vegetables, sambal (a spicy relish or sauce), and usually nasi uduk (traditional coconut rice). The dish is called pecel lele or pecak lele. Lele is the Indonesian word for catfish. The same dish can also be called as lele penyet (squashed catfish) if the fish is lightly squashed along with sambal with a stone mortar-and-pestle. The pecel or pecak version presents the fish in a separate plate while the mortar is solely for sambal.
In Malaysia, catfish is called ikan keli and is fried with spices or grilled and eaten with tamarind and Thai chili gravy and is also often eaten with steamed rice.
In Bangladesh and the Indian states of Odisha, West Bengal and Assam, catfish (locally known as magur) is eaten as a favored delicacy during the monsoons. In the Indian state of Kerala, the local catfish, known as thedu' or etta in Malayalam, is also popular.
In Hungary, catfish is often cooked in paprika sauce (Harcsapaprikás) typical of Hungarian cuisine. It is traditionally served with pasta smothered with curd cheese (túrós csusza).
In Myanmar (formally Burma), catfish is usually used in mohinga, a traditional noodle fish soup cooked with lemon grass, ginger, garlic, pepper, banana stem, onions, and other local ingredients.
Vietnamese catfish, of the genus Pangasius, cannot be legally marketed as catfish in the United States, and so is referred to as swai or basa. Only fish of the family Ictaluridae may be marketed as catfish in the United States. In the UK, Vietnamese catfish is sometimes sold as "Vietnamese river cobbler", although more commonly as Basa.
In Nigeria, catfish is often cooked in a variety of stews. It is particularly cooked in a delicacy popularly known as "catfish pepper soup" which is enjoyed throughout the nation.
In Jewish dietary law, known as kashrut, fish must have fins and scales to be kosher. Since catfish lack scales, they are not kosher.
Mythology
In the mythology of the Japanese Shinto religion natural phenomenon are caused by kami. Earthquakes are caused by a giant catfish called Namazu. There are other kami associated with earthquakes. In Kyoto it's usually an eel, but after the 1855 Edo earthquake Namazu-e (鯰絵, "catfish prints") were printed giving more popularity to the catfish kami that has been known since the 16th century Otsu-e. In one catfish print the divine white horse of Amaterasu is depicted knocking down the earthquake-causing catfish.
Dangers to humans
While the vast majority of catfish are harmless to humans, a few species are known to present some risk. Many catfish species have "stings" (actually non-venomous in most cases) embedded behind their fins; thus precautions must be taken when handling them. Stings by the venomous striped eel catfish have killed people in rare cases.
Catfish fishing records
By information from International Game Fish Association IGFA the most outstanding record:
- The biggest flathead catfish caught was by Ken Paulie in the Elk City Reservoir in Kansas, US on 19 May 1998 that weighed 55.79 kg (123 lb 0 oz)
See also
References
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External links
- All catfish species inventory
- "Giant Baghair caught in Jamuna" Archived 29 November 2014 at the Wayback Machine in The Daily Star (Bangladesh), 12 May 2009
- Skelton, Paul H. and Teugels, Guy G. 1992. Ichthyological Bulletin; No. 56: Neotype description for the African catfish Clarias Gariepinus (Burchell, 1822) (Pisces: Siluroidei: Clariidae). J.L.B. Smith Institute of Ichthyology, Rhodes University, Grahamstown, South Africa
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