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''Coregonus lavaretus'' is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
freshwater whitefish The freshwater whitefish are fishes of the subfamily Coregoninae, which contains whitefishes (both freshwater and anadromous) and ciscoes, and is one of three subfamilies in the salmon family Salmonidae. Apart from the subfamily Coregoninae, ...
, in the
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 ...
. It is 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 its genus '' Coregonus''. There are widely different concepts about the delimitation of the species ''Coregonus lavaretus'' and about the number of species in the genus ''Coregonus'' in general.


Lavaret

In a narrow sense, ''Coregonus lavaretus'', or the lavaret, is considered to be endemic to Lake Bourget and Lake d'Aiguebelette in the
Rhône The Rhône ( , ; Occitan language, Occitan: ''Ròse''; Franco-Provençal, Arpitan: ''Rôno'') is a major river in France and Switzerland, rising in the Alps and flowing west and south through Lake Geneva and Southeastern France before dischargi ...
river basin in
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
, whereas it formerly also occurred in Lake Geneva. According to this view there is a great number of distinct whitefish species in lakes, rivers and brackish waters of Central and Northern Europe.


European whitefish (common whitefish)

In the broad sense, ''Coregonus lavaretus'', in English referred to as the common whitefish or European whitefish, is widespread from central and northwest Europe to Siberia. Often called the ''C. lavaretus'' complex and considered as a superspecies, it encompasses many of the whitefish populations suggested by others to be locally restricted species (such as the British
powan The powan (''Coregonus clupeoides'') is a kind of freshwater whitefish endemic to two lochs in Scotland, Loch Lomond and Loch Eck. It has been successfully introduced in two other sites, Loch Sloy Hydro-Electric Scheme, Loch Sloy and the Carron V ...
and the
gwyniad The gwyniad (''Coregonus pennantii''), or Welsh whitefish, is a species of freshwater Actinopterygii, ray-finned fish belonging to the Family (biology), family Salmonidae, which includes the salmon, trouts and related fishes. This species is End ...
or the Alpine gravenche, as well as distinct intralacustrine morphs and populations characterized by different feeding habits, gill raker numbers, growth patterns and migration behaviour. Genetic studies suggest that the whitefish diversity within this complex is mostly of post-glacial origin. The resource polymorphism represented by the feeding morphs has evolved repeatedly and independently within individual lakes, and similar morphs in different lakes are not closely related to each other.


Description

There is much variation among the European whitefish forms, but in general they have a tapered body, a slightly protruding upper jaw and a fleshy dorsal fin that is typical of the salmon family. The snout is short and tapered, a fact that distinguishes this species from the two other North European ''Coregonus'' species, vendace (''Coregonus albula'') and the introduced peled (''Coregonus peled''). The former has a protuberant lower jaw while in the latter, the jaws are equal in length. The back is bluish green or brownish, the flanks silvery and the belly white. The fins are dark grey. This fish seldom grows more than long or exceeds in weight. (This description refers to the broad concept of European whitefish, from a
Fennoscandia __NOTOC__ Fennoscandia (Finnish language, Finnish, Swedish language, Swedish and ; ), or the Fennoscandian Peninsula, is a peninsula in Europe which includes the Scandinavian Peninsula, Scandinavian and Kola Peninsula, Kola peninsulas, mainland ...
n perspective)


Biology

The European whitefish mostly feed on bottom-dwelling invertebrates or zooplankton. Larger fish also take insects off the surface of the water and eat fish fry. Breeding takes place in the autumn between September and November, largely depending on the water temperature. Different populations in the same sections of water may spawn at different times. Many populations in seas and lakes tend to make their way up-river to spawn, but others populations remain in lakes or the sea even when breeding.


Houting

The houting (''Coregonus oxyrhinchus'') is a European
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of whitefish in the family
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 ...
that was long thought
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 ...
. It was native to the estuaries and rivers draining to the
North Sea The North Sea lies between Great Britain, Denmark, Norway, Germany, the Netherlands, Belgium, and France. A sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the Norwegian Se ...
. The houting is distinguishable from other '' Coregonus'' taxa by having a long, pointed snout, an inferior mouth and a different number of gill rakers.Freyhof, J. and C. Schöter. 2005. The houting Coregonus oxyrinchus (L.)(Salmoniformes: Coregonidae), a globally extinct species from the North Sea basin. Journal of Fish Biology 67:3, 713–729.Schöter C., 2002. Revision der Schnäpel und Großen Maränen des Nordseeund südwestlichen Ostseeraumes (Teleostei: Coregonidae). Diplomarbeit, Rheinischen Friedrich-Wilhelms-Universität, Bonn. The houting once occurred in
Belgium Belgium, officially the Kingdom of Belgium, is a country in Northwestern Europe. Situated in a coastal lowland region known as the Low Countries, it is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeas ...
,
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
,
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
, the
Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
and
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
. In 2023, DNA
analysis Analysis (: analyses) is the process of breaking a complex topic or substance into smaller parts in order to gain a better understanding of it. The technique has been applied in the study of mathematics and logic since before Aristotle (38 ...
found the houting to be genetically indistinguishable from ''Coregonus lavaretus'', and therefore not extinct.


Controversial status

There is controversy whether whitefish surviving in the southeastern North Sea sector of
Denmark Denmark is a Nordic countries, Nordic country in Northern Europe. It is the metropole and most populous constituent of the Kingdom of Denmark,, . also known as the Danish Realm, a constitutionally unitary state that includes the Autonomous a ...
(
Wadden Sea The Wadden Sea ( ; ; or ; ; ; ) is an intertidal zone in the southeastern part of the North Sea. It lies between the coast of northwestern continental Europe and the range of low-lying Frisian Islands, forming a shallow body of water with tida ...
) and considered there as houting (Danish: ''snæbel'') represent the same species as the houting that was
extirpated Local extinction, also extirpation, is the termination of a species (or other taxon) in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with global extinctions. Local extinctions mark a chan ...
from the more southwestern parts of the North Sea.Hansen M.M., Fraser D.J., Als T.D. & Mensberg K.L.D.(2008)
Reproductive isolation, evolutionary distinctiveness and setting conservation priorities: The case of a European lake whitefish and the endangered North Sea houting (''Coregonus'' spp.)
BMC Evolutionary Biology 8:137 doi:10.1186/14716-2.
Like the more southwestern population, the Danish North Sea population has a long, pointed snout and an inferior mouth, and it is anadromous, spending most of its adult life in coastal waters, but migrating into rivers to breed (some other European ''Coregonus'' occur in brackish water, but the ability to live long-term in full salt water is unique to the houting).Møller, P.R.; and H. Carl, editors (2012). Atlas over danske ferskvandsfisk tlas of Danish Freshwater Fish Natural History Museum of Denmark . The Danish houting is genetically part of the widespread ''Coregonus lavaretus''
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 ...
(including '' Coregonus maraena'' of the
Baltic Sea The Baltic Sea is an arm of the Atlantic Ocean that is enclosed by the countries of Denmark, Estonia, Finland, Germany, Latvia, Lithuania, Poland, Russia, Sweden, and the North European Plain, North and Central European Plain regions. It is the ...
basin; some prefer to include the Danish houting in this species), while its genetic relationship to the extinct population cannot be confirmed due to a lack of sufficient samples of the latter.Jacobsen M.W.; Hansen M.M.; Orlando L.; Bekkevold D.; Bernatchez L.; Willerslev E.; and Gilbert M.T. (2012). Mitogenome sequencing reveals shallow evolutionary histories and recent divergence time between morphologically and ecologically distinct European whitefish (Coregonus spp.). Mol Ecol. 21(11): 2727–2742. Nevertheless, there are some minor differences in the genetics of the Danish houting compared to other living members in the ''C. lavaretus'' complex, as well as the differences in morphology and ecology, making it an evolutionarily significant unit.Dierking, J.; L. Phelps; K. Præbel; G. Ramm; E. Prigge; J. Borcherding; M. Brunke; and C. Eizaguirre (2014). Anthropogenic hybridization between endangered migratory and commercially harvested stationary whitefish taxa (Coregonus spp.). Evol Appl. 7(9): 1068–1083. Hybridization and
introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Introg ...
between North Sea houting and its relatives is well-documented, and likely the result of translocations of ''Coregonus'' between different regions by humans. Some researchers argue that the morphological differences between different houting populations are not exceptional within the broader variation of the European whitefish, and probably no species-level extinction has taken place. The primary reason for treating the Danish houting and the extinct houting as separate are differences in the number of gill rakers (on average, the Danish has fewer than the extinct), but this number can vary extensively in ''Coregonus'', even within a single population and species,Christensen, G.H. (2010). Danmarks ferskvandsfisk enmark's Freshwater Fish pp. 47–48. and genetic studies of ''Coregonus'' have shown that gill rakers are of limited use in predicting relationship among populations. Some think that the morphological differences in number of gill rakers are sufficient for treating them as separate, and that the last true houting was caught in the lower
Rhine The Rhine ( ) is one of the List of rivers of Europe, major rivers in Europe. The river begins in the Swiss canton of Graubünden in the southeastern Swiss Alps. It forms part of the Swiss-Liechtenstein border, then part of the Austria–Swit ...
in 1940. Studies in the early 2000s (decade) indicated that there was no overlap in the possible number of gill rakers of the two (28–35 in the Danish; 38–46 in the extinct), but later reviews have shown that there is an overlap (up to 41 has been found in the
Elbe The Elbe ( ; ; or ''Elv''; Upper Sorbian, Upper and , ) is one of the major rivers of Central Europe. It rises in the Giant Mountains of the northern Czech Republic before traversing much of Bohemia (western half of the Czech Republic), then Ge ...
, a reintroduced population based on Danish houting).Ramm, G.; and J. Dierking (2014). . Geomar, Helmholtz Centre for Ocean Research Kiel. A €13 million restoration project of the Danish houting, partly funded by the European Union's LIFE programme and the Danish Natural Agency, was undertaken in 2005–2013, and there is ongoing monitoring of the species and regulation of the fish-eating great cormorant from important locations.Svendsen, J.C.; A.K.O. Alstrup; and L.F. Jensen (11 April 2018)
Save a North Sea fish from becoming museum piece.
Nature. Retrieved 12 April 2018.
As of 2019, a total of more than €20 million has been used on its conservation, with almost two-thirds funded by Denmark and the remaining by the European Union.Mandrup, L., and N.M. Jørgensen (26 April 2019)
Efter redningsaktioner for 150 mio. kr.: Danmarks mest sjældne fisk er i lille fremgang.
fter rescue projects for 150 million DKK: The population of Denmark's rarest fish is slowly increasing DR News. Retrieved 26 April 2019.
However, the only remaining fully natural and significant population of Danish houting is in the Vidå River, estimated in 2014 to consist of about 3,500 adults. Little is known about its exact spawning and juvenile requirements, and despite the earlier project it was still declining, leading to fears that it could become fully extinct unless more is done to preserve it. After years with a downward trend in its numbers, an increase to about 4,000 adult Danish houtings was registered in 2018–19, with most individuals in the Vidå and fewer in (both populations increasing). Individuals from the Danish population have been used as a basis for re-establishing houting in the Eider, Elbe (both indisputably a natural part of the range) and Rhine (arguably non-native, if the extinct is recognized as a separate species). On the 24th of October 2023 researchers of the Research Institute for Nature and Forest in Belgium captured a live specimen in the Scheldt river in Kruibeke, while the species was thought to be extinct in the country for over 100 years. Possibly the specimen migrated there from the Rhine population. In Annex II of the EC Habitats Directive (92/43/EEC), a species listed as ''Coregonus oxyrhynchus'' is protected for 'anadromous populations in certain sectors of the North Sea' In 2023, DNA analysis found the houting to be genetically indistinguishable from ''Coregonus lavaretus'', and therefore not extinct.


References

{{Authority control lavaretus Freshwater fish of Europe Fish described in 1758 Taxa named by Carl Linnaeus Habitats Directive species