Invasive Species Compendium

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Datasheet

Phoxinus phoxinus
(European minnow)

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Datasheet

Phoxinus phoxinus (European minnow)

Summary

  • Last modified
  • 25 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Animal
  • Preferred Scientific Name
  • Phoxinus phoxinus
  • Preferred Common Name
  • European minnow
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Chordata
  •       Subphylum: Vertebrata
  •         Class: Actinopterygii
  • Summary of Invasiveness
  • P. phoxinus is mainly being introduced to new watercourses bordering on watercourses where it already is established, thereby slowly but steadily widening its area of distribution. The species is able to establ...

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Pictures

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PictureTitleCaptionCopyright
Eurasian minnows (Phoxinus phoxinus).
TitleMinnows swimming
CaptionEurasian minnows (Phoxinus phoxinus).
CopyrightBjørn Ove Johnsen
Eurasian minnows (Phoxinus phoxinus).
Minnows swimmingEurasian minnows (Phoxinus phoxinus).Bjørn Ove Johnsen

Identity

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Preferred Scientific Name

  • Phoxinus phoxinus (Linnaeus, 1758)

Preferred Common Name

  • European minnow

International Common Names

  • English: Eurasian minnow

Local Common Names

  • Denmark: elritse
  • France: vairon
  • Germany: elritze
  • Norway: ørekyt
  • Sweden: elritsa

Summary of Invasiveness

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P. phoxinus is mainly being introduced to new watercourses bordering on watercourses where it already is established, thereby slowly but steadily widening its area of distribution. The species is able to establish viable populations in most freshwater systems, from lowland to high alpine areas, in particular where few other fish species are present. However, successful establishment seems to require habitats which include some slow-flowing or lake-like areas. Rivers with only swift currents seem not to provide suitable habitats for P. phoxinus to complete its lifecycle.

Impacts on native ecosystems have not been well documented, except in the case of allopatric brown trout, where establishment of P. phoxinus leads to reduced brown trout densities.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Chordata
  •             Subphylum: Vertebrata
  •                 Class: Actinopterygii
  •                     Order: Cypriniformes
  •                         Family: Cyprinidae
  •                             Genus: Phoxinus
  •                                 Species: Phoxinus phoxinus

Notes on Taxonomy and Nomenclature

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This species retains the name given by Linnaeus in 1758. The common English name, minnow, needs a geographical qualifier (“European” or “Eurasian”) to be precise, as “minnow” may be used for a number of small cyprinids, both in Eurasia and America. Some authors indicate that several species may be included in the present Phoxinus phoxinus (cf. Kottelat and Freyhof, 2007).

Description

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P. phoxinus has a torpedo-shaped body, with 80-100 small cycloid scales along the lateral line.

P. phoxinus has variable colours, but are normally brownish green on the back, separated from the whitish belly by numerous brown and black blotches along the side, sometimes uniting to form a stripe. Males are brightly coloured during spawning, with white flashes at the fins, reddish pectoral and pelvic fins, a black throat, green along the sides and a scarlet belly (Maitland, 2004). 

Common size is 6-10 cm, with a maximum of 14-15 cm. The growth rates and age and size at maturation of P. phoxinus varies greatly with factors such as population density and numerous environmental factors (Lien, 1981; Myllylä et al., 1983; Mills and Eloranta, 1985; Mills, 1987; 1988; Museth et al., 2002). 

Distribution

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P. phoxinus is found in almost all of Europe (including the British Isles) and northern Asia. Notable exceptions concerning native distribution are north western Scotland and major parts of Norway. Native distribution in Norway is restricted to the south eastern low altitude areas and parts of Finnmark county in the far north. In Scandinavia, the minnow was also originally absent from most alpine areas. This is less known for other parts of the distribution area.

On the Red List (IUCN), minnows are stated as of least concern. However, their status differs highly in different European countries. In Denmark, P. phoxinus is considered rare (Frier, 1994). In Germany, P. phoxinus is listed as an endangered native species on the red list of all Federal states (Hesthagen and Sandlund, 2007), and in some areas the species is produced in hatcheries and released to sustain natural populations (e.g. in the river Treene in Schleswig-Holstein; LANU, 2002). In the Baltic countries, P. phoxinus is very common (Hesthagen and Sandlund, 2007). It is found in many Latvian rivers, also in brooks and ditches, but not in lakes and coastal waters.

In Norway, P. phoxinus is still being translocated and introduced into new areas, developing dense populations in most localities. At present there are no serious threats to minnow populations in Norway. However, during the past decades acidification affected this species in some areas of southern Norway, and more than 100 populations were either lost or damaged (Hesthagen et al., 1999). These populations are to a large extent located within the native distribution area for P. phoxinus. However, acidification is no longer a serious threat to minnows in this region. Only a few of the lost populations have been re-established (Hesthagen et al., 2007). Improved water quality in this region facilitates establishment of P. phoxinus in watercourses where it is non-native (cf. Larsen et al., 2007). A similar situation is seen in northwestern Scotland (Adams, 1994).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ArmeniaPresentNativeFroese and Pauly, 2008
AzerbaijanPresentNativeFroese and Pauly, 2008
ChinaPresentNativeFroese and Pauly, 2008
-HunanPresentNativeZipCodeZoo, 2008
-XinjiangPresentNativeZipCodeZoo, 2008
Georgia (Republic of)PresentNativeZipCodeZoo, 2008
KazakhstanPresentNativeZipCodeZoo, 2008
Korea, DPRPresentNativeFroese and Pauly, 2008
Korea, Republic ofPresentNativeZipCodeZoo, 2008
TurkeyPresentNativeFroese and Pauly, 2008; ZipCodeZoo, 2008
UzbekistanPresentNativeFroese and Pauly, 2008; ZipCodeZoo, 2008

Africa

EgyptLocalisedZipCodeZoo, 2008Sinai

Europe

AustriaWidespreadNativeFroese and Pauly, 2008
BelarusWidespreadNativeFroese and Pauly, 2008
BelgiumWidespreadNativeSysteme d'Informations sur la Biodiversite en Wallonie, 2008
Bosnia-HercegovinaPresentNativeFroese and Pauly, 2008
BulgariaPresentCAB ABSTRACTS Data Mining 2001; Froese and Pauly, 2008
CroatiaPresentNativeFroese and Pauly, 2008
Czech RepublicWidespreadNativeFroese and Pauly, 2008
DenmarkLocalisedNativeFrier, 1994Indigeneous, but present occurrence reduced
EstoniaWidespreadNativeFroese and Pauly, 2008
FinlandWidespreadNativeRask et al., 2000Indigeneous, widespread
FranceWidespreadNativeAllardi and Keith, 1991
GermanyWidespreadNativeFroese and Pauly, 2008
GreecePresentNativeFroese and Pauly, 2008
HungaryWidespreadNativeFroese and Pauly, 2008
IrelandWidespreadNativeFroese and Pauly, 2008Native according to Maitland 2004
ItalyPresentNativeFroese and Pauly, 2008
LatviaWidespreadNativePlikss, 2002
LiechtensteinWidespreadNativeFroese and Pauly, 2008
LithuaniaWidespreadNativeFroese and Pauly, 2008
LuxembourgWidespreadNativeFroese and Pauly, 2008
MoldovaPresentNativeFroese and Pauly, 2008
NetherlandsWidespreadNativeFroese and Pauly, 2008
NorwayWidespread Invasive Hesthagen and Sandlund, 1996; Hesthagen and Sandlund, 2006; Museth et al., 2007Originally restricted distribution in the country, introduced into numerous new watercourses over the last 100-140 yrs.
PolandWidespreadNativeFroese and Pauly, 2008
RomaniaPresentNativeFroese and Pauly, 2008
Russian FederationWidespreadNativeReshetnikov et al., 1997Indigeneous, widespread
SerbiaPresentNativeFroese and Pauly, 2008
SlovakiaWidespreadNativeFroese and Pauly, 2008
SloveniaPresentNativeFroese and Pauly, 2008
SpainLocalisedNativeFroese and Pauly, 2008
SwedenWidespreadFilipsson, 1994Common throughout Sweden, introduced to few localities.
SwitzerlandWidespreadNativeFroese and Pauly, 2008
UKPresentCAB ABSTRACTS Data Mining 2001; Maitland, 2004; Froese and Pauly, 2008
UkraineWidespreadNativeFroese and Pauly, 2008

History of Introduction and Spread

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Originally, minnows were spread because fishermen used them as live bait for catching species like brown trout (Salmo trutta), Arctic charr (Salvelinus alpinus), perch (Perca fluviatilis) and pike (Exos lucius) (Huitfeldt-Kaas, 1918). This practice is considered to be the main reason for most introductions throughout the 1900s. However, minnows have also been accidentally introduced in a large number of lakes together with stocked hatchery-reared brown trout (Borgstrøm, 1973; Lura and Kålås, 1994). Brown trout stocking has been routinely done especially in lakes modified as hydropower reservoirs, in order to compensate for reduced natural recruitment (Vøllestad and Hesthagen, 2001). These reservoirs are often located in the upper sections of watersheds. Whenever minnows were introduced, they were able to subsequently migrate downstream and become established in more lakes. This frequently occurred during the 1960s and 1970s. Minnows have also been spread through tunnels constructed for hydropower development between watersheds. In a few cases minnows have been intentionally introduced to provide forage fish for brown trout. In one case minnows were introduced as a control measure against the locally bothersome ‘Tune fly’ (Simuliidae) (Halleraker and Hesthagen, 1994).

Risk of Introduction

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As minnows are quite hardy animals, they may be kept and transported alive in very small bodies of water with high temperatures and low oxygen contents. This makes it easy for anyone to move the species between lakes or water courses.

P. phoxinus disperse easily downstream, but extended river stretches with continuous swift currents may appear to constitute a barrier to downstream migration. In the river Sanddøla, central Norway, a major tributary of the River Namsen, minnows were established in the headwater Lake Otersjøen around 1960. By 2005 they had still not spread downstream in Sanddøla, probably due to the continuous swift currents over a distance of more than 45 km (Thorstad et al., 2006). Based on the observation in several cases that downstream spread by minnows may cover 3-7 km per year, it may be speculated that the species require appropriate habitats for feeding, over wintering and possibly reproducing (i.e. lakes, pools or slow flowing river habitats) at suitable intervals. In such extreme lotic habitats, “resting habitats” at 5-10 km intervals may be necessary for individuals to survive the downstream migration.

Minnows are able to migrate against relatively strong currents for very short distances, but in small streams it is possible to construct barriers that stop minnows but allow the passage of larger brown trout (Holthe et al., 2005).

Habitat

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P. phoxinus are found in a variety of habitats over a wide geographical range throughout its native distributional area; in brackish water as well as in different types of freshwater; streams, rivers, ponds, and large lakes located from coastal areas to high mountains. P. phoxinus has been found at an altitude of 1,403 m above sea level. in a lake in the central mountain area in southern Norway (Jotunheimen), and even up 2000 m above sea level in other parts of the distribution area (Lelek, 1987). The species is less numerous in steep, fast flowing rivers. It occurs most abundantly in shallow lakes and slow flowing streams and rivers. P. phoxinus is also abundant in regulated lakes, even when the water level might vary by several metres throughout the year.

Laboratory studies of minnows revealed a significant preference for stony substratum (grain diameter 5-50 mm) over sand (grain diameter 0.5-1.0 mm) (Jacobsen, 1979). The preference for a stony substratum was strongest in old, schooling individuals, and significantly higher than in schools of juveniles aged 2-5 months. Substrate selection in minnows is probably associated with shelter against predator fish. In Lake Øvre Heimdalsvatn, located at 1,090 m above sea level. In southern Norway, where minnows were introduced in the late 1960s, brown trout preyed heavily on mature minnows shortly after ice break at the end of June, when minnows constituted 9 and 20% of the stomach volume of trout in length groups 16-30 and = 30 cm, respectively (Museth et al., 2005). Predation on minnows was only occasionally detected during July, August and September. Brown trout selectively preyed on minnows infected by Ligula intestinalis (Museth, 2001).

In Lake Øvre Heimdalsvatn, gillnet catches of minnows decreased significantly with increasing depth, being 32.1, 13.1 and 0.9 fish per 100 m2 net area at 1.5, 3.0 and 6.0 m depths, respectively (Museth et al., 2002). The highest densities of minnows were obtained at depths between 0.2 and 0.5 m (Museth et al., 2002). Furthermore, the minnows captured by gillnets were restricted to the net area close to the bottom, and less than 1% were captured more than 50 cm above the bottom.

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Freshwater
Irrigation channels Present, no further details Natural
Lakes Present, no further details Natural
Reservoirs Present, no further details Natural
Rivers / streams Present, no further details Natural
Ponds Present, no further details Natural
Brackish
Estuaries Present, no further details Natural

Biology and Ecology

Top of page Reproductive Biology

P. phoxinus displays considerable variability in life-history traits, i.e. in age and size at sexual maturity, growth rate and longevity (Mills, 1988). Age at maturity has been recorded over a gradient from 0+ to 6+, in fast and slow growing populations, respectively (Museth et al., 2002). Sexual maturity occurs at a smaller body size and at a lower age in lowland localities compared with those located at a higher altitude and latitude. In most cases, however, size at maturity deviates little from 50 mm. In the river Utsjoki in Finnish Lapland, maturity was strongly size-dependent and delayed until the fish reached 5, 6 or even 7 years of age, with a maximum age of 13 years at a length of only 75 mm (Mills, 1988). In Norway, sexual maturity in minnows has been recorded at between 2 and 15 years. In the alpine lake Øvre Heimdalsvatn, minnows of age 4 and 5 years made up about 67% of the spawning stock (Museth et al., 2002). All mature individuals were larger than 50 mm in length, and only a few specimens were smaller than 55 mm. Whereas no minnows older than 3 years were recorded in River Frome, UK (Mills, 1988), the oldest individual in the alpine lake Øvre Heimdalsvatn was 13 years (Museth et al., 2002).

In Norway, P. phoxinus spawns mainly in June and July, depending on altitude and latitude. The fish spawn in shoals over stones and gravel, either in running water or in shallow areas close to the shore line. The adhesive eggs stick to the substratum. In Øvre Heimdalsvatn, spawning activity was observed only 4-8 days after ice break in early June, with the spawning period lasting about 3 weeks (Museth et al., 2002). The adhesive yellow eggs of about 1.0-1.5 mm in diameter hatch after 5-10 days. Individual fecundity is between 200 and 1000 eggs.

It may appear that sexually mature minnows change behaviour towards spawning time, becoming more susceptible to fish predation (Museth et al., 2005).

Nutrition
P. phoxinus feeds on invertebrates (mainly crustaceans and insect larvae), and some plant material. They may also prey on salmonid alevins (Huusko and Sutela, 1997; J Museth, Agricultural University of Norway, personal communication, 2008).

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Water pH (pH) Optimum 6.5–7.5 tolerated
Water temperature (ºC temperature) Optimum 2–20 tolerated

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Salmo trutta Predator All Stages not specific

Means of Movement and Dispersal

Top of page Accidental Introduction

Originally, minnows were spread because fishermen used it as live bait for catching species like brown trout (Salmo trutta), Arctic charr (Salvelinus alpinus), perch (Perca fluviatilis) and pike (Exos lucius) (Huitfeldt-Kaas, 1918). This practice is considered to be the main reason for most introductions throughout the 1900s. However, minnows have also been accidentally introduced together with stocked hatchery-reared brown trout in a large number of lakes (Borgstrøm, 1973; Lura and Kålås, 1994). Brown trout stocking has been routinely done especially in lakes modified as hydropower reservoirs in order to compensate for reduced natural recruitment (Vøllestad and Hesthagen, 2001).

Natural Dispersal (Non-Biotic)

These reservoirs are often located in the upper sections of watersheds. Whenever minnows were introduced, they were in most cases able to subsequently migrate downstream and become established in more lakes. This frequently occurred during the 1960s and 1970s. Minnows have also been spread through tunnels between watersheds constructed for hydropower development.

Intentional Introduction
In a few cases minnows have been intentionally introduced to provide forage fish for brown trout. In one case minnows have been introduced as a control measure against the so-called tune fly (Simuliidae) (Halleraker and Hesthagen, 1994).

Pathway Causes

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Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Bait Yes Yes

Impact Summary

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CategoryImpact
Environment (generally) Negative

Environmental Impact

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P. phoxinus may introduce new parasites where they become established. In some sub-alpine lakes in southern Norway introduced minnows caused infection with new parasite species in snails, mussels and different insects, but not in brown trout (Hartvigsen, 1997).

In Norway, survey net catches of brown trout in lakes with and without introduced European minnows demonstrated a 35% reduction in catches in lakes where brown trout were sympatric with introduced minnows (Museth et al., 2007).

The abundance of important food items for brown trout may show a significant decline after the introduction of P. phoxinus. In Lake Øvre Heimdalsvatn, the introduction of minnows caused major changes in the benthic community (Brittain et al., 1988; 1995). Zoobenthos diversity declined, with a marked increase in numbers of oligochaetes and small forms, especially chironomids. There was also a marked decline in numbers of Gammarus lacustris, especially the proportion of larger individuals. However, total benthic densities remained similar to pre-introduction. G. lacustris formed a major component of the diet of minnows, while it’s occurrence in brown trout stomachs declined greatly. Lepidurus arcticus also virtually disappeared from the trout diet, probably due to minnow predation. In another Norwegian reservoir, introduced European minnows fed on the planktonic stages of L. arcticus, and after a few years adult specimens became an insignificant part of the diet of brown trout (Borgstrøm et al., 1985).

Introduction of European minnows may also cause reduced recruitment in brown trout. In Lake Øvre Heimdalsvatn, the cohort size of age-class 4 was reduced by approximately 50% during a period in sympatry with minnows compared to the situation before the introduction of minnows. There was no significant change in annual individual length increment (Borgstrøm et al., 1996). It is uncertain whether the reduction of trout recruitment was due to direct interactions with minnows in the nursery streams, or an indirect effect caused, for example, by increased brown trout cannibalism. Minnows may prey on salmonid larvae (Huusko and Sutela, 1997).

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Capable of securing and ingesting a wide range of food
  • Highly mobile locally
  • Long lived
  • Has high reproductive potential
  • Gregarious
Impact outcomes
  • Altered trophic level
  • Damaged ecosystem services
  • Modification of natural benthic communities
  • Modification of nutrient regime
  • Negatively impacts aquaculture/fisheries
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Pest and disease transmission
  • Predation
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Highly likely to be transported internationally deliberately
  • Highly likely to be transported internationally illegally
  • Difficult/costly to control

Uses List

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Animal feed, fodder, forage

  • Bait/attractant

Prevention and Control

Top of page Prevention

Raising public awareness appears to be the only measure with some potential to prevent or delay spreading of P. phoxinus to new localities. In some cases eradication from small water bodies may be feasible, in particular if the locality is expected to serve as a centre of further spreading into more localities. Control by habitat modification or bio manipulation may be possible but is rarely investigated.

Gaps in Knowledge/Research Needs

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Research is needed on the potential of applying habitat modification in order to reduce the impact of introduced P. phoxinus on salmonids and other fish species. This is particularly relevant in regulated rivers (Heavily Modified Water Bodies according to the EU Water Framework Directive). There may also be a potential for bio manipulation (management of predators or competitors) to reduce population densities and impacts of introduced minnows.

References

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Adams CE, 1994. The fish community of Loch Lomond, Scotland: Its history and rapidly changing status. Hydrobiologia, 290:91-102.

Allardi J; Keith P, 1991. Atlas préliminaire des poissons d'eau douce de France. Coll. Patrimoines Naturels. Paris, : Secrétariat Faune Flore, Muséum National d'Histoire Naturelle,.

Borgstrøm R, 1973. Spredning av ørekyt. Jakt - Fiske - Friluftsliv, 102:28-29.

Borgstrøm R; Brittain JE; Hasle K; Skjølas S; Dokk JG, 1996. Reduced recruitment in brown trout Salmo trutta, the role of interactions with the minnow Phoxinus phoxinus. Nordic Journal of Freshwater Reserach, 72:30-38.

Borgstrøm R; Garnas E; Saltveit SJ, 1985. Interactions between brown trout, Salmo trutta L. and minnow, Phoxinus phoxinus (L.) for their common prey, Lepidurus arcticus (PALLAS). Verh. Internat. Verein. Limnol, 22:2548-2552.

Brittain JE; Brabrand A; Saltveit SJ, 1995. Effekt pa fisk og næringsdyr ved introduksjon av ørekyt. Spredning av ferskvannsorganismer. Seminarreferat DN-Notat 1995-4., 146-148.

Brittain JE; Brabrand A; Saltveit SJ; Bremnes T; Røsten E, 1988. The biology and population dynamics of Gammarus lacustris in relation to the introduction of minnows, Phoxinus phoxinus, into Øvre Heimdalsvatn, a Norwegian subalpine lake. Rapp. Lab. Ferskv. Økol. Innlandsfiske, 109:1-30.

Filipsson O, 1994. [English title not available]. (Nya fiskbestand genom inplantering eller spridning av fisk) Information fran Sötvattenslaboratoriet Drottningholm, 2:1-65.

Frier J-O, 1994. [English title not available]. (Danske ferskvandsfisk og deres udbredelsesomrade) Truede ferskvandsfiskearter i Norden [ed. by Frier JO]., 83-99.

Froese R; Pauly D, 2008. FishBase. http://www.fishbase.org

Halleraker JH; Hesthagen T, 1994. [English title not available]. (Kategorisering av innlandsfiskesystemer i deler av Glommavassdraget) NINA Oppdragsmelding,, 302:1-18.

Hartvigsen R, 1997. [English title not available]. (Spredning av parasitter ved innvandring og/eller introduksjon av nye fiskearter: spredning av ørekyt (Phoxinus phoxinus) til ørretvassdrag) NINA Oppdragsmelding,, 466:1-14.

Hesthagen T; Sandlund OT, 1996. Changes in the distribution of minnow (Phoxinus phoxinus) in Norway: Causes and effects. NINA Forskningsrapport,, 013:1-16.

Hesthagen T; Sandlund OT, 2006. NOBANIS - Invasive Alien Species Fact Sheet - Phoxinus phoxinus. Online Database of the North European and Baltic Network on Invasive Alien Species - NOBANIS. http://www.nobanis.org

Hesthagen T; Sevaldrud IH; Berger HM, 1999. Assessment of damage to fish populations in Norwegian lakes due to acidification. Ambio, 28:12-17.

Hesthagen T; Walseng B; Karlsen LR; Langaker RM, 2007. Effects of liming on the aquatic fauna in a Norwegian watershed: Why do crustaceans and fish respond differently? Water and Soil Pollut. Focus, 7:339-345.

Holthe E; Lund E; Finstad B; Thorstad EB; Mckinley RS, 2005. A fish selective obstacle to prevent dispersion of an unwanted fish species, based on leaping capabilities. Fisheries Management and Ecology, 12:143-147.

Huitfeldt-Kaas H, 1918. Ferskvandsfiskenes utbredelse og indvandring i Norge med et tillæg om krebsen. Centraltrykkeriet, Kristiania., 106 pp.

Huusko A; Sutela T, 1997. Minnow predation on vendace larvae: intersection of alternative prey phenologies and size-based vulnerability. Journal of Fish Biology, 50:965-977.

Jacobsen OJ, 1979. Substrate preference in the minnow (Phoxinus phoxinus). Polskie Archiwum Hydrobiologie, 26:371-378.

Kottelat M; Freyhof J, 2007. Handbook of European Freshwater Fishes. Cornol, Switzerland: Publications Kottelat, 646 pp.

LANU, 2002. [English title not available]. (Die Süsswasserfische und Neunaugen Schleswig-Holsteins - Rote Liste) Landesamt für Natur und Umwelt des Landes Schleswig-Holstein., 1-58.

Larsen BM; Sandlund OT; Berger HM; Hesthagen T, 2007. Invasives, introductions and acidification: the dynamics of a stressed river fish community. Water, Air & Soil Pollution: Focus, 7:285-291.

Lelek A, 1987. The freshwater fishes of Europe. Threatened fishes of Europe. AULA-Verlag, Wiesbaden.

Lien L, 1981. Biology of the minnow Phoxinus phoxinus and its interactions with brown trout Salmo trutta in Øvre Heimdalsvatn, Norway. Holarctic Ecology, 4:191-200.

Lura H; Kalas S, 1994. [English title not available]. (Ferskvassfiskane si utbreiing i Sogn og Fjordane, Hordaland og Rogaland) Rapport Universitetet i Bergen, Zoologisk Museum.

Maitland PS, 2004. Keys to the freshwater fish of Britain and Ireland, with notes on their distribution and ecology. Ambleside, Cumbria, UK: UK Freshwater Biological Association (FBA), 248 pp.

Mills CA, 1988. Life history of the minnow Phoxinus phoxinus (L.) in a productive stream. Freshwater Biology, 17:53-67.

Mills CA, 1988. The effect of extreme northerly climatic conditions on the life history of the minnow, Phoxinus phoxinus (L.). Journal of Fish Biology, 33:545-561.

Mills CA; Eloranta A, 1985. The biology of Phoxinus phoxinus (L.) and other littoral zone fishes in Lake Konnevesi, central Finland. Annales Zoologici Fennici, 22:1-12.

Museth J, 2001. Effects of Ligula intestinalis on habitat use, predation risk and catchability in European minnows. Journal of Fish Biology, 59(4):1070-1080.

Museth J, 2002. Dynamics in European minnow Phoxinus phoxinus and brown trout Salmo trutta populations in mountain habitats: effects of climate and inter- and intraspecific interactions. Agricultural University of Norway, 29.

Museth J; Borgstrøm R; Brittian JE; Herberg I; Naalsund C, 2002. Introduction of the European minnow into a subalpine lake: habitat use and long-term changes in population dynamics. Journal of Fish Biology, 60:1308-1321.

Museth J; Borgstrøm R; Hame T; Holen LA, 2005. Predation by brown trout: a major mortality factor for sexually mature European minnows. Journal of Fish Biology, 62:692-705.

Museth J; Hesthagen T; Sandlund OT; Thorstad EB; Ugedal O, 2007. The history of the European minnow in Norway: from harmless species to pest. Journal of Fish Biology, 71(Supplement D):184-195.

Myllylä M; Torssonen M; Pulliainen E; Kuusela K, 1983. Biological studies on the minnow, Phoxinus phoxinus, in northern Finland. Aquilo: Series Zoologica., 149-156.

Plikss M, 2002. Fish of Latvia. http://latvijas.daba.lv/scripts/db/saraksti/saraksti.cgi?d=zivis&l=en

Rask M; Appelberg M; Hesthagen T; Tammi J; Beier U; Lappalainen A, 2000. Fish status survey of Nordic lakes - species composition, distribution, effects of environmental changes., 508.

Reshetnikov YS; Bogutskaya NG; Vasil'eva ED; Dorofeeva EA; Naseka AM; Popova OA; Savvaitova KA; Sideleva VG; Sokolov LI, 1997. An annotated check-list of the freshwater fishes of Russia. J. Ichthyol, 37(9):687-736.

Systeme d'Informations sur la Biodiversite en Wallonie, 2008. Entry for Phoxinus phoxinus. http://biodiversite.wallonie.be

Thorstad EB; Sandlund OT; Heggberget TG; Finstad A; Museth J; Berger HM; Hesthagen T; Berg OK, 2006. [English title not available]. (Ørekyt I Namsenvassdraget. Utbredelse, spredningsrisiko og tiltak) NINA Rapport, 155:1-69.

Vøllestad LA; Hesthagen T, 2001. Stocking of freshwater fish in Norway: management goals and effects. Nordic Journal of Freshwater Research, 75:143-152.

ZipCodeZoo, 2008. Entry for Phoxinus phoxinus. http://zipcodezoo.com/

Links to Websites

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WebsiteURLComment
European Research Network on Aquatic Invasive Species (ERNAIS)http://www.zin.ru/rbic/projects/ernais/
The North European and Baltic Network on Invasive Alien Specieshttp:// www.nobanis.org/
The Norwegian Biodiversity Information Centrehttp://www.biodiversity.no/

Contributors

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24/01/08 Original text by:

Odd Sandlund, Norwegian Institute for Nature Research, Tungasletta 2, NO-7485 Trondheim, Norway

Distribution Maps

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