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''Coregonus'' is a diverse
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
in the salmon
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
(
Salmonidae Salmonidae (, ) is a family (biology), family of ray-finned fish, the only extant member of the suborder Salmonoidei, consisting of 11 extant genera and over 200 species collectively known as "salmonids" or "salmonoids". The family includes salmo ...
). The ''Coregonus'' species are known as whitefishes. The genus contains at least 68 described
extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
taxa, but the true number of species is a matter of debate. The
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of the genus is ''
Coregonus lavaretus ''Coregonus lavaretus'' is a species of freshwater whitefish, in the family Salmonidae. It is the type species of its genus '' Coregonus''. There are widely different concepts about the delimitation of the species ''Coregonus lavaretus'' and abo ...
''. Most ''Coregonus'' species inhabit lakes and rivers, and several species, including the
Arctic cisco The Arctic cisco (''Coregonus autumnalis''), also known as Arctic omul (), is an anadromy, anadromous species of freshwater whitefish that inhabits the Arctic parts of Siberia especially Yenisey Gulf. It can also be found in Alaska and Canada. I ...
(''C. autumnalis''), the Bering cisco (''C. laurettae''), and the least cisco (''C. sardinella'') are
anadromous Fish migration is mass relocation by fish from one area or body of water to another. Many types of fish migrate on a regular basis, on time scales ranging from daily to annually or longer, and over distances ranging from a few metres to thousa ...
, moving between
salt water Saline water (more commonly known as salt water) is water that contains a high concentration of dissolved salts (mainly sodium chloride). On the United States Geological Survey (USGS) salinity scale, saline water is saltier than brackish wate ...
and
fresh water Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salt (chemistry), salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include ...
. Many whitefish species or
ecotype Ecotypes are organisms which belong to the same species but possess different phenotypical features as a result of environmental factors such as elevation, climate and predation. Ecotypes can be seen in wide geographical distributions and may event ...
s, especially from the
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes spanning the Canada–United States border. The five lakes are Lake Superior, Superior, Lake Michigan, Michigan, Lake Huron, H ...
and the Alpine lakes of
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
, have gone
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
over the past century or are
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching, inv ...
. Among 12 freshwater fish considered extinct in Europe, 6 are ''Coregonus''. All ''Coregonus'' species are protected under appendix III of the Bern Convention,Council of Europe, 1979. Convention on the Conservation of European Wildlife and Natural Heritage. Bern, Switzerland. Available at: http://conventions.coe.int/Treaty/EN/Treaties/Html/104.htm and Annex IV of the EC Habitats Directive (92/43/EEC)


Taxonomy

Phylogenetic evidence indicates that the most basal member of the genus is the highly endangered Atlantic whitefish (''C. huntsmani''), which is
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
to a single river basin in
Nova Scotia Nova Scotia is a Provinces and territories of Canada, province of Canada, located on its east coast. It is one of the three Maritime Canada, Maritime provinces and Population of Canada by province and territory, most populous province in Atlan ...
,
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
. The Atlantic whitefish is thought to have diverged from the rest of the genus during the mid-
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
, about 15 million years ago. The genus was previously subdivided into two subgenera ''Coregonus'' ("true whitefishes") and ''Leucichthys'' ("
ciscoes The ciscoes (or ''ciscos'') are salmonid fish that differ from other members of the genus ''Coregonus'' in having upper and lower jaws of approximately equal length and high gill raker counts. These species have been the focus of much study rece ...
"), ''Coregonus'' comprising taxa with sub-terminal mouth and usually a benthic feeding habit, ''Leucichthys'' those with terminal or supra-terminal mouth and usually a pelagic plankton-feeding habit. This classification is not natural however: based on molecular data, ciscoes comprise two distinct lineages within the genus. Moreover, the genus ''
Stenodus ''Stenodus'' is a genus of large-sized whitefish in the family Salmonidae. It consists of two species; one of them ( beloribitsa) is extinct in the wild. The two species have alternatively been considered subspecies of the single species ''Steno ...
'' is not phylogenetically distinct from ''Coregonus''; although ''Stenodus'' occupies a basal position within the genus, phylogenetic evidence indicates that ''C. huntsmani'' is even more basal than it. The scientific name given to this genus of fish comes from the Greek
κόρη Κόρη may refer to: * Kore (sculpture), a type of ancient Greek sculpture depicting a young female * Persephone, a goddess in Greek mythology * Kore (moon) Kore , also known as , is the outermost named natural satellite of Jupiter. It was d ...
(''kórē'') "daughter ; eye pupil" and γωνία (''gōnía'') "angle", because their
pupil The pupil is a hole located in the center of the iris of the eye that allows light to strike the retina.Cassin, B. and Solomon, S. (1990) ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing Company. It appears black becau ...
makes an angle, even though they share this feature with a large number of other fish.


Species diversity

There is much uncertainty and confusion in the classification of the many species of this genus. Particularly, one extreme view of diversity recognises just two main species in Northern and Central Europe, the
common whitefish ''Coregonus lavaretus'' is a species of freshwater whitefish, in the family Salmonidae. It is the type species of its genus ''Coregonus''. There are widely different concepts about the delimitation of the species ''Coregonus lavaretus'' and abou ...
''C. lavaretus'' and the vendace ''C. albula'', whereas others would divide these into numerous, often narrowly distributed species. A drastic increase in number of recognized species occurred in 2007, when a review advocated that more than 50 local European populations should be considered as distinct based on morphological differences. It has been estimated that several of them are very young, having separated from each other less than 15,000 years ago. Many of these were primarily defined based on number of
gill raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of th ...
s. Although this largely is
hereditary Heredity, also called inheritance or biological inheritance, is the passing on of traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cells or organisms acquire the genetic inform ...
, the number is highly variable (even within single populations and species), can change relatively fast in response to changes and genetic studies have shown that they often are of limited use in predicting relationships among populations (a large difference in gill raker number does not necessarily equal a distant relationship). Genetic differences between several of the recently proposed species, even ones that are relatively distinct morphologically, are very limited and sometimes they are not
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
. Various ''Coregonus'', whether regarded as separate species or not, readily interbreed with each other. A review of whitefish in the United Kingdom found that the
identification key In biology, an identification key, taxonomic key, or frequently just key, is a printed or computer-aided device that aids in the identification of biological organisms. Historically, the most common type of identification key is the dichotomous k ...
provided in 2007 did not match most individuals and that solid evidence for more than one species in that region is lacking. Many European lakes have more than one ''Coregonus'' morph differing in ecology and morphology (especially gill rakers). Such morphs are sometimes partially
reproductively isolated The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offsprin ...
from each other, leading to suggestions of recognizing them as separate but clinal species. The morphs or clinal species may rapidly disappear (in 15 years or less, equalling three ''Coregonus'' generations) by merging into a single in response to changes in the habitat. A similar pattern can be seen in North America where the
ciscoes The ciscoes (or ''ciscos'') are salmonid fish that differ from other members of the genus ''Coregonus'' in having upper and lower jaws of approximately equal length and high gill raker counts. These species have been the focus of much study rece ...
of the ''
Coregonus artedi ''Coregonus artedi'', commonly known as the cisco, is a North American species of freshwater whitefish in the family (biology), family Salmonidae. The number of species and definition of species limits in North American cisco (fish), ciscoes is a ...
''
complex Complex commonly refers to: * Complexity, the behaviour of a system whose components interact in multiple ways so possible interactions are difficult to describe ** Complex system, a system composed of many components which may interact with each ...
in the
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes spanning the Canada–United States border. The five lakes are Lake Superior, Superior, Lake Michigan, Michigan, Lake Huron, H ...
and elsewhere comprise several, often co-occurring morphs or ecotypes, whose taxonomic status remains controversial.


Species

In 2017,
FishBase FishBase is a global species database of fish species (specifically finfish). It is the largest and most extensively accessed online database on adult finfish on the web.
listed 78 species, including the more than 50 proposed for Europe in 2007. Some of these are recently extinct (marked with a dagger, "†") and ''C. reighardi'' is likely extinct. * '' Coregonus acrinasus'' Oliver M. Selz, Carmela J. Dönz, Pascal Vonlanthen, Ole Seehausen, 2020 * '' Coregonus albellus'' Fatio, 1890 (autumn brienzlig) * ''
Coregonus albula ''Coregonus albula'', known as the vendace or as the European cisco, is a species of freshwater whitefish in the family Salmonidae. It is found in lakes in northern Europe, especially Finland, Latvia, Lithuania, Sweden, Russia and Estonia, and in ...
''
Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
,
1758 Events January–March * January 1 – Swedish biologist Carl Linnaeus (Carl von Linné) publishes in Stockholm the first volume (''Animalia'') of the 10th edition of ''Systema Naturae'', the starting point of modern zoologic ...
(vendace) *
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Coregonus alpenae'' ( Koelz, 1924) (longjaw cisco) * '' Coregonus alpinus'' Fatio, 1885 (kropfer) * '' Coregonus anaulorum'' Chereshnev, 1996 * '' Coregonus arenicolus'' Kottelat, 1997 * ''
Coregonus artedi ''Coregonus artedi'', commonly known as the cisco, is a North American species of freshwater whitefish in the family (biology), family Salmonidae. The number of species and definition of species limits in North American cisco (fish), ciscoes is a ...
'' Lesueur, 1818 (northern cisco or lake herring) * '' Coregonus atterensis'' Kottelat, 1997 * '' Coregonus austriacus'' C. C. Vogt, 1909 * '' Coregonus autumnalis'' (
Pallas Pallas may refer to: Astronomy * 2 Pallas asteroid ** Pallas family, a group of asteroids that includes 2 Pallas * Pallas (crater), a crater on Earth's moon Mythology * Pallas (Giant), a son of Uranus and Gaia, killed and flayed by Athena * Pa ...
, 1776)
(Arctic cisco) * '' Coregonus baerii'' Kessler, 1864 * '' Coregonus baicalensis'' Dybowski, 1874 * '' Coregonus baunti'' Mukhomediyarov, 1948 * ''
Coregonus bavaricus ''Coregonus bavaricus'', the Ammersee kilch, is a species of freshwater Coregonus, whitefish endemic to Ammersee, Lake Ammersee in the Germany, German state of Upper Bavaria. A small, silver-colored fish, it typically lives between deep, though ...
'' Hofer, 1909 *
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Coregonus bezola'' Fatio, 1888 (bezoule) * '' Coregonus brienzii'' Oliver M. Selz, Carmela J. Dönz, Pascal Vonlanthen, Ole Seehausen, 2020 * '' Coregonus candidus'' Goll, 1883 * '' Coregonus chadary'' Dybowski, 1869 (Khadary whitefish) * '' Coregonus clupeaformis'' ( Mitchill, 1818) (lake whitefish) * '' Coregonus clupeoides'' Lacépède, 1803 (powan) * '' Coregonus confusus'' Fatio, 1885 * '' Coregonus danneri'' C. C. Vogt, 1908 * '' Coregonus duplex'' Fatio, 1890 * '' Coregonus fatioi'' Kottelat, 1997 *
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Coregonus fera'' Jurine, 1825 (fera) * '' Coregonus fontanae'' M. Schulz & Freyhof, 2003 (Stechlin cisco) *
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Coregonus gutturosus'' ( C. C. Gmelin ( de), 1818) * '' Coregonus heglingus'' Schinz, 1822 *
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Coregonus hiemalis'' Jurine, 1825 (gravenche) * '' Coregonus hoferi'' L. S. Berg, 1932 * '' Coregonus holsata'' Thienemann, 1916 * '' Coregonus hoyi'' ( Milner, 1874) (bloater) * '' Coregonus huntsmani'' W. B. Scott, 1987 (Atlantic whitefish) *
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Coregonus johannae'' ( G. Wagner, 1910) (deepwater cisco) * '' Coregonus kiletz'' Michailovsky, 1903 * '' Coregonus kiyi'' ( Koelz, 1921) (kiyi) * '' Coregonus ladogae'' Pravdin, Golubev & Belyaeva, 1938 * '' Coregonus laurettae'' T. H. Bean, 1881 (Bering cisco) * ''
Coregonus lavaretus ''Coregonus lavaretus'' is a species of freshwater whitefish, in the family Salmonidae. It is the type species of its genus '' Coregonus''. There are widely different concepts about the delimitation of the species ''Coregonus lavaretus'' and abo ...
''
Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
,
1758 Events January–March * January 1 – Swedish biologist Carl Linnaeus (Carl von Linné) publishes in Stockholm the first volume (''Animalia'') of the 10th edition of ''Systema Naturae'', the starting point of modern zoologic ...
(common whitefish, European whitefish; lavaret) * '' Coregonus lucinensis'' Thienemann, 1933 * '' Coregonus lutokka'' Kottelat, Bogutskaya & Freyhof, 2005 * '' Coregonus macrophthalmus'' Nüsslin, 1882 * '' Coregonus maraena'' (
Bloch Bloch is a surname of German origin. Notable people with this surname include: A *Adele Bloch-Bauer (1881–1925), Austrian entrepreneur *Albert Bloch (1882–1961), American painter *Alexandre Bloch (1857–1919), French painter *Alfred Bloch ( ...
, 1779)
(maraena whitefish) * '' Coregonus maraenoides'' L. S. Berg, 1916 * '' Coregonus maxillaris'' Günther, 1866 * '' Coregonus megalops'' Widegren, 1863 (lacustrine fluvial whitefish) * '' Coregonus migratorius'' ( Georgi, 1775) (omul) * '' Coregonus muksun'' (
Pallas Pallas may refer to: Astronomy * 2 Pallas asteroid ** Pallas family, a group of asteroids that includes 2 Pallas * Pallas (crater), a crater on Earth's moon Mythology * Pallas (Giant), a son of Uranus and Gaia, killed and flayed by Athena * Pa ...
, 1814)
(muksun) * '' Coregonus nasus'' (
Pallas Pallas may refer to: Astronomy * 2 Pallas asteroid ** Pallas family, a group of asteroids that includes 2 Pallas * Pallas (crater), a crater on Earth's moon Mythology * Pallas (Giant), a son of Uranus and Gaia, killed and flayed by Athena * Pa ...
, 1776)
(broad whitefish) * '' Coregonus nelsonii'' T. H. Bean, 1884 (Alaska whitefish) *
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Coregonus nigripinnis'' ( Milner, 1874) (blackfin cisco) * '' Coregonus nilssoni''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1848
* '' Coregonus nipigon'' ( Koelz, 1925) * '' Coregonus nobilis'' Haack, 1882 *
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Coregonus oxyrinchus''
Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
,
1758 Events January–March * January 1 – Swedish biologist Carl Linnaeus (Carl von Linné) publishes in Stockholm the first volume (''Animalia'') of the 10th edition of ''Systema Naturae'', the starting point of modern zoologic ...
(houting) * '' Coregonus palaea'' G. Cuvier, 1829 * '' Coregonus pallasii''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1848
* '' Coregonus peled'' ( J. F. Gmelin, 1789) (peled) * '' Coregonus pennantii''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1848
(gwyniad) * ''
Coregonus pidschian The humpback whitefish (''Coregonus pidschian''), also referred to as the bottom whitefish, the Arctic whitefishKottelat, M. and Freyhof, J. 2007. Handbook of European Freshwater Fishes. Publications Kottelat, Cornol, Switzerland. or the pidschian ...
'' ( J. F. Gmelin, 1789) (humpback whitefish) * '' Coregonus pollan'' W. Thompson, 1835 (Irish pollan) * '' Coregonus pravdinellus'' Dulkeit, 1949 * ''
Coregonus profundus ''Coregonus'' is a diverse genus of fish in the salmon family (Salmonidae). The ''Coregonus'' species are known as whitefishes. The genus contains at least 68 described extant taxa, but the true number of species is a matter of debate. The type ...
'' Oliver M. Selz, Carmela J. Dönz, Pascal Vonlanthen, Ole Seehausen, 2020 * ''
Coregonus reighardi The shortnose cisco (''Coregonus reighardi'') is a North American freshwater whitefish in the salmon family (biology), family Salmonidae. One of the members of the broader Coregonus artedi#The Coregonus artedi species complex, ''Coregonus artedi' ...
'' ( Koelz, 1924) (shortnose cisco) * '' Coregonus renke'' ( Schrank, 1783) *
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
'' Coregonus restrictus'' Fatio, 1885 * '' Coregonus sardinella''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1848
(Sardine cisco) * '' Coregonus steinmanni'' Oliver M. Selz, Carmela J. Dönz, Pascal Vonlanthen, Ole Seehausen, 2020 * '' Coregonus stigmaticus'' Regan, 1908 (schelly) * '' Coregonus subautumnalis'' Kaganowsky, 1932 * '' Coregonus suidteri'' Fatio, 1885 * '' Coregonus trybomi'' Svärdson ( sv), 1979 * '' Coregonus tugun'' (
Pallas Pallas may refer to: Astronomy * 2 Pallas asteroid ** Pallas family, a group of asteroids that includes 2 Pallas * Pallas (crater), a crater on Earth's moon Mythology * Pallas (Giant), a son of Uranus and Gaia, killed and flayed by Athena * Pa ...
, 1814)
* '' Coregonus ussuriensis'' L. S. Berg, 1906 (Amur whitefish) * '' Coregonus vandesius'' J. Richardson, 1836 (vendace) * '' Coregonus vessicus'' Dryagin, 1932 * ''
Coregonus wartmanni ''Coregonus wartmanni'' is a freshwater fish of the family Salmonidae found in Lake Constance Lake Constance (, ) refers to three bodies of water on the Rhine at the northern foot of the Alps: Upper Lake Constance (''Obersee''), Lower Lake ...
'' (
Bloch Bloch is a surname of German origin. Notable people with this surname include: A *Adele Bloch-Bauer (1881–1925), Austrian entrepreneur *Albert Bloch (1882–1961), American painter *Alexandre Bloch (1857–1919), French painter *Alfred Bloch ( ...
, 1784)
* '' Coregonus widegreni'' Malmgren, 1863 (Valaam whitefish) * '' Coregonus zenithicus'' ( D. S. Jordan & Evermann, 1909) (shortjaw cisco) * '' Coregonus zuerichensis'' Nüsslin, 1882 * '' Coregonus zugensis'' Nüsslin, 1882


References


External links

* * * * {{Authority control Ray-finned fish genera Taxa named by Carl Linnaeus