Pimephales promelas (fathead minnow)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Natural Food Sources
- Latitude/Altitude Ranges
- Water Tolerances
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Threatened Species
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Pimephales promelas Rafinesque, 1820
Preferred Common Name
- fathead minnow
Other Scientific Names
- Cliola smithii Evermann & Cox, 1896
- Colistus parietalis Cope, 1871
- Hyborhynchus confertus Girard, 1856
- Hyborhynchus nigellus Cope, 1875
- Hyborhynchus puniceus Girard, 1856
- Pimephales agassizii Cope, 1867
- Pimephales anuli Kendall, 1903
- Pimephales fasciatus Girard, 1856
- Pimephales maculosus Girard, 1856
- Pimephales milesii Cope, 1865
- Pimephales promelas harveyensis Hubbs & Lagler, 1949
- Pimephales promelas promelas Rafinesque, 1820
- Pimephalus promelas Rafinesque, 1820
- Plargyrus melanocephalus Abbott, 1860
International Common Names
- English: black-head minnow; rosey reds
- Spanish: carpita cabezona; mino cabezón
- French: tête de boule
Local Common Names
- Canada: tête-de-boule
- Czech Republic: jelecek velkohlavý
- Denmark: tykhovedet elritse; tyrehoved
- Finland: paksupäämutu
- Germany: Fettköpfige Elritze
- Sweden: knölskallelöja
Summary of InvasivenessTop of page
P. promelas, commonly known as the fathead minnow, is a small-bodied, short-lived, nest guarding cyprinid fish. It is one of the most widely distributed fishes in North America and its native range covers large expanses from the Rocky Mountains in Canada south to Texas and northern Mexico. It has been accidently or intentionally introduced to many areas, where it has been stocked for its use as bait or as an aquarium pet. It is tolerant of a range of environmental conditions, has a high reproductive output and consumes a variety of food resources. It therefore has a high invasive potential and has been recorded as a pest in many countries. It is known to cause declines of native and endangered species including Colorado pikeminnow (Ptychocheilus lucius), the Lost River sucker (Deltistes luxatus) and shortnose sucker (Chasmistes brevirostris) in Oregon and California. It has also negatively impacted habitats, such as wetlands in the Prairie Pothole Region, USA. In Europe, its introduction has resulted in the spread of enteric redmouth disease through the introduction of a bacterial pathogen (Yersinia ruckeri) (Anseeuw et al., 2012).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Chordata
- Subphylum: Vertebrata
- Class: Actinopterygii
- Order: Cypriniformes
- Family: Cyprinidae
- Genus: Pimephales
- Species: Pimephales promelas
Notes on Taxonomy and NomenclatureTop of page
Three subspecies of Pimephales promelas were recognized up until the 1960s: P. promelas subsp. promelas, P. promelas subsp. confertus and P. promelas subsp. harveyensis. Although morphologically dissimilar, Vandermeer (1966) concluded from a study on the meristic and morphometric characters that the subspecies were unwarranted and unworkable (Danylchuk et al., 2012).
Two evolutionarily closely related species to the P. promelas are the bluntnose minnow (Pimephales notatus) and the slim minnow (Pimephales tenellus) (Ross et al., 2001). Hybridisation between P. promelas and P. notatus has been observed in their natural environment (Trautman, 1981; Danylchuk et al., 2012).
DescriptionTop of page
P. promelas is olive-yellow in colour with a yellow-silvery underside. Juveniles have a distinctive grey-black lateral band that runs from the gill cover to the tail. Breeding males are often much darker in colour with a black head and a large black spot at the anterior base of the dorsal fin. The body is slightly to moderately compressed, the lateral line tends to be incomplete (with 7-40 pored scales), and the pre-dorsal scales are crowded and significantly smaller than those on the sides of the body (Ross et al., 2001). The blunt, dorsally rounded shape of the head in breeding males (moderately so in others) gives the species its common name ‘fathead minnow’ (Nelson and Paetz, 1992).
The following describes the morphological changes for both sexes: ‘Breeding males develop a large, mucus-secreting, fleshy pad on the nape (Smith and Murphy 1974; Smith 1978) and 16 large tubercles in three rows on the snout. Females develop an enlarged urogenital papilla approximately one month prior to spawning (Flickinger 1969)’ (Hassan-Williams and Bonner, 2012).
Both the male and female of the colour morph commonly known as the ‘rosy red minnow’ are pinky-red in colour on the body and fins.
P. promelas has a moderately forked caudal fin, 8 dorsal fin rays, 7 anal fin rays and 44-50 lateral line scales. The maximum length recorded for the fish is 10 cm but length can range from 2.5 cm to 7.5 cm long. It has a pharyngeal teeth count of 0,4-4,0 (Nelson and Paetz, 1992).
DistributionTop of page
P. promelas is found throughout North America, ‘occurring naturally between the Rocky Mountains and the Appalachians in the Mississippi, Hudson Bay, and Great Lakes basins and from Louisiana, Texas, and Chihuahua, Mexico, in the south to the Alberta-Northwest Territories border and northern Ontario in the north’ (Danylchuk et al., 2012). In the Northeast, it is found in all states in the region (Werner, 2004). It occurs in high densities in the wetlands of the Prairie Pothole Region (Zimmer et al., 2001).
It has been widely introduced outside of its native range to many areas west of the Continental Divide and east of the Appalachians. Populations of P. promelas have been reported as established in Puerto Rico (USGS, 2012). It has also been introduced intentionally into Iran and Europe, where wild populations have become established in the UK, Belgium and France. It is uncertain as to whether wild populations are present in Germany as well (Danylchuk et al., 2012).
The exact native range of P. promelasis difficult to determine due to its widespread introductions from bait bucket releases. The literature therefore presents a confused distribution of this species for some areas; for example, its status as native to Tennessee is uncertain (USGS, 2012).
Distribution TableTop of page
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.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|United Kingdom||Present, Few occurrences||Introduced||2008||First wild population recorded in North Yorkshire|
|-Alberta||Present||Native||Widespread in central and southern Alberta|
|-Saskatchewan||Present||In the south|
|-Louisiana||Present||Native and Introduced||Native and introduced|
|-Mississippi||Present||Introduced||Not native but may have the possible exception of occurrences in the Tennessee drainage; Original citation: Ross et al. (2001)|
|-Nebraska||Present||Native and Introduced||Native and introduced|
|-Wisconsin||Present, Widespread||Present in all three major drainage basins – Mississippi River, Lake Superior and Lake Michigan)|
History of Introduction and SpreadTop of page
For many areas, P. promelas has been accidentally or intentionally introduced into the natural environment. For example, in Alabama it was introduced into streams and rivers accidentally through bait bucket releases or where it had escaped from nearby ponds. It was first recorded in California in 1950 and was probably released into the Colorado River from a bait tank. It was later introduced as forage for sport fishes and then became widespread as the bait industry grew. Other states where introductions from bait bucket releases are known are: Leon County in Florida, Meadow Creek in Kentucky, Youghiogheny River drainage in Maryland, and in Pennsylvania, Maryland, Massachusetts, Minnesota and Texas (USGS, 2012).
The species has been imported both locally and state wide; for example, the Minnesota bait industry sold P. promelas locally and as far as Florida and New Mexico (USGS, 2012). In some cases, it was introduced to areas that already had native populations, such as Nebraska and Ohio. In other areas, including Idaho, Louisiana, Utah and Virginia this species was introduced as a forage fish for bass and other game fish, and used in fish hatcheries. In the Twin Cities area of Minnesota, P. promelas was introduced to control mosquitoes (USGS, 2012). The species has also been introduced and stocked in sewerage treatment ponds in Wisconsin (Danylchuk et al., 2012).
It was introduced to Belgium for game and fish stocking and the first wild population to be become established was recorded in 1986 (Anseeuw et al., 2012). In the UK, the first population of P. promelas was observed in 2008 but few observations have been made of established populations (Godard et al., 2010).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Belgium||North America||1986||Hunting, angling, sport or racing (pathway cause)||Yes||Yes||Anseeuw et al. (2012)|
|California||North America||1950||Hunting, angling, sport or racing (pathway cause)||Yes||Yes||USGS (2012)||Initially introduced as bait but then stocked by the Department of Fish and Game|
|France||North America||Hunting, angling, sport or racing (pathway cause)||Yes||Sommer (2012)|
|Idaho||North America||1945||Forage (pathway cause)||USGS (2012)|
|Iran||North America||Sommer (2012)||unknown reason|
|Kentucky||North America||Hunting, angling, sport or racing (pathway cause)||USGS (2012)|
|Maryland||North America||Hunting, angling, sport or racing (pathway cause)||Yes||USGS (2012)|
|Massachusetts||North America||Hunting, angling, sport or racing (pathway cause)||Yes||USGS (2012)|
|Minnesota||North America||Hunting, angling, sport or racing (pathway cause)||Yes||USGS (2012)|
|UK||North America||2008||Godard et al. (2010)||unknown reason|
|Utah||North America||1969||Forage (pathway cause)||Yes||USGS (2012)|
|Virginia||North America||1950||Hunting, angling, sport or racing (pathway cause)||Yes||USGS (2012)|
Risk of IntroductionTop of page
The continued use of the P. promelas as bait and as an aquarium pet presents a continued risk of their release and introduction into non-native habitats. Their rapid maturation, high reproductive potential and ability to tolerate a wide range of habitats poses a high invasive risk and a substantial threat to native species.
In the UK, climatic conditions are thought to be unfavourable for yearlings, meaning that presently there is a low risk of populations becoming established. However, modelling predicts that the P. promelas could benefit from climatic warming by 2050 and so the risk of introduction could increase, and subsequently its invasiveness (Godard et al. 2010).
HabitatTop of page
P. promelas can be found in a variety of freshwater habitats, including creeks, headwaters, small rivers, lakes and ponds. Its ability to establish itself outside of its native range is facilitated by its tolerance of high temperatures, turbidity, high salinity and low oxygenated waters (NatureServe, 2012). It does not tend to be found in upland areas or at high altitudes and seems to prefer slow moving steams and ponds (Werner, 2004).
Habitat ListTop of page
|Freshwater||Irrigation channels||Secondary/tolerated habitat||Natural|
|Freshwater||Rivers / streams||Principal habitat||Natural|
Biology and EcologyTop of page
The haploid (n) and diploid (2n) chromosome numbers for P. promelas in the USA have been reported as n = 25 and 2n = 50-50 (Froese and Pauly, 2012). The genetics of the species have also been well studied by April et al. (2013).
P. promelas can spawn anytime between May and September. During this time, ‘a single female may produce 6803-10,164 eggs and may have participated in 16-26 spawning sequences’ (Ross et al., 2001). P. promelas can reach maturity and spawn within their first year.
Spawning sites are carefully selected for by the male and tend to be in shallow water, less than a meter deep, with a log, branch, rock or other object being used for the eggs to be deposited underneath. This species is unusual in that it shows some level of parental care. After spawning, the male chases the female away from the nest and then guards the eggs until they hatch. He cleans them with the thick spongy pad on his back and chases away any intruders (Nelson and Paetz, 1992). Eggs have a low hatching success rate if not tended to by the male. A male may accumulate as many as 12,000 eggs. Hatching times can vary from 13 days at lower temperatures (15°C) to 4 days at higher temperatures (25°C) (Ross et al., 2001).
Physiology and Phenology
In Canadian boreal lakes, populations of P. promelas show high phenotypic diversity. Although tolerant to a range of conditions, some environmental variables may affect the size of individuals. For example, in some lakes and ponds, which vary in oxygen levels from year to year, it may lead to a disproportionate number of smaller individuals within a population as the larger individuals cannot survive. Much of this variation may also be dependent on the social environment where the absence of larger, socially dominant males may cause smaller males to spawn earlier in the season, producing a larger number of young which are small in size (Danylchuk and Tonn, 2000).
There has been selective breeding of a red colour morph of P. promelas, which is commonly known as the rosy red minnow. This colour fish is frequently used in the aquarium trade.
This species is non-migratory.
Population Size and Structure
Size demography of individuals within a population may be affected by environmental variables, such as low oxygen levels. This can cause high mortality of larger individuals within a population (Danylchuk and Tonn, 2000). Populations of P. promelas in Ontario, Canada, have been shown to respond negatively to concentrations of oestrogen which have contaminated freshwater. Population size decreases as a result of the effects of oestrogen on reproductive health (Kidd et al., 2007).
Primarily a benthic feeder, most of its diet consists of algae and protozoans, but it will also eat zooplankton and aquatic insect larvae (Nelson and Paetz, 1992). Two types of feeding strategies, particle and filter-feeding, are used, resulting in the consumption of a wide range of aquatic invertebrates (Hanson et al., 2005). One study in Alberta observed P. promelas largely consuming invertebrates including crustaceans and chironomid pupae (Price et al., 1991).
P. promelas is generally found in schools. They can be found swimming with other small fish, including the brook stickleback (Culaea inconstans), southern redbelly dace (Chrosomus erythrogaster) and the bluntnose minnow (P. notatus) (Nelson and Paetz, 1992).
It is tolerant of a range of water quality conditions, with the exception of high acidity, and is frequently found in wetlands with high alkaline and saline conditions. It is also physiologically capable of surviving very low oxygen conditions in lakes.
Natural Food SourcesTop of page
|Food Source||Food Source Datasheet||Life Stage||Contribution to Total Food Intake (%)||Details|
ClimateTop of page
|B - Dry (arid and semi-arid)||Preferred||< 860mm precipitation annually|
|Cs - Warm temperate climate with dry summer||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Cw - Warm temperate climate with dry winter||Preferred||Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)|
|Ds - Continental climate with dry summer||Tolerated||Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Water TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|Dissolved oxygen (mg/l)||1||Harmful||Reproduction reduced at 2 mg/l and larval growth at 4 mg/l|
|Water pH (pH)||7.0||7.5||Optimum|
|Water temperature (ºC temperature)||10||29||Optimum||Tolerates higher temperatures of 33-40°C|
Natural enemiesTop of page
Notes on Natural EnemiesTop of page
P. promelas provides an important dietary component for many piscivorous fishes and birds. Numerous game fish species, such as lake trout (Salvelinus namaycush), walleye (Sander vitreus) and northern pike (Esox lucius) prey on P. promelas (Kidd et al., 2007). Yellow perch (Perca favescens) are also known to feed on P. promelas and a study demonstrated that they consistently feed on small individuals when given a choice of different sizes (Paszkowski and Tonn, 1994).
Means of Movement and DispersalTop of page
Natural Dispersal (Non-Biotic)
In some areas it is unclear as to whether native populations have dispersed or if new populations have been established due to bait or aquarium release.
The origin of many introduced populations of P. promelas has been attributed to their use as bait by anglers. Accidental escapes from stocked ponds may also be responsible for some introductions.
It has been intentionally introduced into new areas through the aquarium trade, where it is sold either as an aquarium pet or a feeder fish (Chang et al, 2009). Some introductions into waterways may be as a result of its release from aquariums and hatcheries (USGS, 2012).
Pathway CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
|Economic/livelihood||Positive and negative|
|Environment (generally)||Positive and negative|
Economic ImpactTop of page
Repeated introductions of P. promelas to wetland habitats in USA and Canada are causing economic repercussions for wetland managers responsible for water quality, flood protection and optimization of waterfowl production (Zimmer et al., 2006).
Environmental ImpactTop of page
Impact on Habitats
Introduced populations of P. promelas can grow rapidly because of their high reproductive output and their ability to utilize a wide range of food resources. High densities can have deleterious effects on the energy flow in wetlands, such as the Prairie Pothole Region of the US and Canada (Zimmer et al., 2006). Populations of native fish often flux as winterkill reduces or eliminates densities each year, but repeated introductions of fathead minnows can cause algal blooms, high turbidity and a loss of wetland plants, invertebrates and other vertebrate species. Therefore, introductions of P. promelas to wetlands can significantly reduce habitat quality (Butler and Hanson, 1996; Zimmer et al., 2001; Zimmer et al., 2006).
Impact on Biodiversity
This species is considered a pest in much of its introduced and native range. It may compete with other fish for food resources and habitat, especially during the spawning season when males are territorial at nest sites. It can indirectly affect aquatic insects, large cladocerans and other vertebrate and invertebrate species by reducing habitat quality (by increasing turbidity, phosphrous and chlorophyll a) (Zimmer et al., 2001).
Among other introduced fish, it is thought to predate on young suckers, including the Lost River sucker (Deltistes luxatus) and shortnose sucker (Chasmistes brevirostris) in Oregon and northern California. It has been implicated with the decline of other species, such as the endangered Colorado pikeminnow (Ptychocheilus lucius), wood frog (Lithobates sylvaticus) and, in southeastern Arizona, the Chiracahua leopard frog (Lithobates chiricahuensis) (USGS, 2012). Negative interactions between fathead minnows and endangered native fishes, such as woundfin (Plagopterus argentissimus) and Virgin River chub (Gila seminuda) within the Colorado River Baisin have also been documented (US Fish and Wildlife Service, 2008).
In Belgium it is not only considered a pest as it predates on the eggs and larvae of native fishes but it is also thought to be responsible for the introduction of Yersinia ruckeri, the agent of enteric redmouth disease (Anseeuw et al., 2012).
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Chasmistes brevirostris||EN (IUCN red list: Endangered)||USA; California; Oregon||Predation||USGS (2012)|
|Deltistes luxatus||EN (IUCN red list: Endangered)||USA; California; Oregon||Predation||USGS (2012)|
|Gila seminuda (Virgin River chub)||EN (IUCN red list: Endangered); National list(s); USA ESA listing as endangered species||USA||Competition||US Fish and Wildlife Service (2008)|
|Lithobates sylvaticus (wood frog)||LC (IUCN red list: Least concern)||USA||Competition; Predation||USGS (2012)|
|Plagopterus argentissimus (woundfin)||EN (IUCN red list: Endangered); USA ESA listing as endangered species||USA; Arizona; New Mexico||Competition||US Fish and Wildlife Service (2008)|
|Ptychocheilus lucius (Colorado pikeminnow)||No Details||USA||Competition; Predation||USGS (2012)|
|Rana chiricahuensis (Chiricahua leopard frog)||VU (IUCN red list: Vulnerable)||USA; Arizona||Competition; Predation||USGS (2012)|
Risk and Impact FactorsTop of page
- 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
- Capable of securing and ingesting a wide range of food
- Highly mobile locally
- Fast growing
- Has high reproductive potential
- Altered trophic level
- Damaged ecosystem services
- Ecosystem change/ habitat alteration
- Modification of natural benthic communities
- Modification of nutrient regime
- Reduced native biodiversity
- Threat to/ loss of endangered species
- Threat to/ loss of native species
- Competition (unspecified)
- Pest and disease transmission
- Highly likely to be transported internationally accidentally
- Highly likely to be transported internationally deliberately
- Difficult to identify/detect in the field
UsesTop of page
P. promelas is the most common species of minnow sold as bait for angling activities. It is commonly sold as an aquarium pet and the rosy red morph has been bred for this purpose as well as its use as bait (Robison and Buchanan, 1984).
They are also used as supplementary forage in bass-bream ponds. In Mississippi, the species is stocked annually in spring. They produce a number of offspring and so are useful forage for bass and larger bream.
They can also be used to make biomass from nutrient and plant material in sewage treatment ponds (Becker, 1983).
The species is an important bioassay animal due to the ease in which it can be cultured and maintained (Danylchuk et al., 2012). It is widely used in the USA in many laboratory toxicity studies (Ross et al., 2001).
It has been used for mosquito control in sloughs, ponds, ditches and storm water drainage channels. In Madison, USA, it was even found to decrease densities of mosquito that were associated with the spread of West Nile virus (Irwin and Paskewitz, 2009; Danylchuk et al., 2012).
Uses ListTop of page
Animal feed, fodder, forage
- Laboratory use
- Pet/aquarium trade
- Sport (hunting, shooting, fishing, racing)
Similarities to Other Species/ConditionsTop of page
P. promelas is similar to other minnows but can generally be differentiated by the high number of small pre-dorsal scales, which are noticeably smaller than those scales on the sides. The pre-dorsal scales on other minnows tend to be similar in size, number and compactness to those on the sides of the fish (Danylchuk et al., 2012). The creek chub (Semotilus atromaculatus) can be easily mistaken for P. promelas, but this species can be identified by its larger jaw, 8 anal fins, a complete lateral line and 11 or fewer gill rakers (Hassan-Williams and Bonner, 2012). Other species within the Pimephales genus, such as the bluntnose minnow (P. notatus) can be distinguished by their terminal and slightly oblique to subterminal mouth, in contrast to the distinctive strongly oblique to superior and almost vertical mouth of P. promelas (Werner, 2004; Danylchuk et al., 2012). Other differences, such as the slender body form or complete lateral line, can be used to distinguish the closely related bullhead minnow (P. vigilax) from P. promelas (Ross et al., 2001).
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
The aquarium trade is an important vector for the release of this species into local waterways. Therefore, educating both retailers and customers about the risks posed by both pet and bait bucket releases could reduce the further introduction of this species, as well as other aquatic species. Furthermore, improving the labelling and identification of fish species in stores and facilitating correct disposal of unwanted fish could help minimize new introductions (Chang et al., 2009).
Policies to prevent stocking of fish in certain vulnerable habitats could reduce the further spread of this species (Butler and Hanson, 1996).
Introducing or increasing the number of predators within systems, such as walleye or lake trout, could also control P. promelas populations. However, the effects of this on the rest of the ecosystem could be more harmful than the high abundance of P. promelas.
Gaps in Knowledge/Research NeedsTop of page
It has been suggested that P. promelas could become invasive in the UK within the next 40 years, so further research is needed to identify areas of elevated risk, and control and eradication plans would need to be proposed and implemented (Godard et al., 2010).
ReferencesTop of page
Anseeuw D; Branquart E; Lieffrig F; Micha JC; Parkinson D; Verreycken H, 2012. Pimephales promelas: fathead minnow. Invasive species in Belgium. Pimephales promelas: fathead minnow., Belgium: The Belgian Biodiversity Platform (online). [Invasive species in Belgium.] http://ias.biodiversity.be/
April J; Hunner RH; Dion-Cote AM; Bernatchez L, 2013. Glacial cycles as an allopatric speciation pump in north-eastern American freshwater fishes. Mol. Ecol, 22(2):409-422.
Butler MG; Hanson MA, 1996. Water Resources research grant proposal: Influences of fathead minnows on nutrient partitioning, water clarity, and ecosystem structure in prairie wetlands. Influences of fathead minnows on nutrient partitioning, water clarity, and ecosystem structure in prairie wetlands., USA: USGS Water Resources. http://water.usgs.gov/wrri/96grants/ncr12nd.htm
Chang AL; Grossman JD; Spezio TS; Weiskel HW; Blum JC; Burt JW; Muir AA; Piovia-Scott J; Veblen KE; Grosholz ED, 2009. Tackling aquatic invasions: risks and opportunities for the aquarium fish industry. Biological Invasions, 11(4):773-785. http://www.springerlink.com/content/b330w27880752rr4/fulltext.html
Danylchuk AJ; Tonn WM, 2000. Effects of social structure on reproductive activity in male fathead minnows (Pimephales promelas), 12(4):482-489.
Danylchuk Tonn WM; Paszkowski CA, 2012. Fishes of Wisconsin. Wisconsin, USA: Wisconsin Department of Natural Resources, US Geological Survey. http://www.fow-ebook.us
Flickenger SA, 1969. Determination of sexes in the fathead minnow, 98(3):526-527.
Froese R; Pauly D, 2004. FishBase DVD. Penang, Malaysia: Worldfish Center. Online at www.fishbase.org.
Froese R; Pauly D, 2012. Fishbase [ed. by Froese, R. \Pauly, D.]. Fishbase (online). http://www.fishbase.org
Godard MJ; Britton JR; Guillerault N; Zieba G; Copp GH, 2010. Fathead minnows in Europe: preliminary results. Fathead minnows in Europe., UK: Cefas-Lowestoft and Bournemouth University. http://icais.org/pdf/2010abstracts/Gordon%20Copp.pdf
Hanson MA; Zimmer KD; Butler MG; Tangen BA; Herwig BR; Euliss Jr NH, 2005. Biotic interactions as determinants of ecosystem structure in prairie wetlands: an example using fish. Biotic interactions as determinants of ecosystem structure in prairie wetlands, 25(3):764-775. http://digitalcommons.unl.edu/usgsnpwrc/270
Hassan-Williams C; Bonner TH, 2012. Texas Freshwater Fishes. Texas, USA: Texas State University (San Marcos Biology Department). http://www.bio.txstate.edu/
Irwin P; Paskewitz S, 2009. Investigation of fathead minnows (Pimephales promelas) as a biological control agent of Culex mosquitoes under laboratory and field conditions. Journal of the American Mosquito Control Association, 25(3):301-309. http://www.bioone.org/perlserv/?request=get-current-issue
ITIS, 2013. Integrated Taxonomic Information System (ITIS). Washington, DC, USA: Smithsonian Institution/NMNH. http://www.itis.gov/
Kidd KA; Blanchfield PJ; Mills KH; Palace VP; Evans RE; Lazorchak JM; Flick RW, 2007. Collapse of a fish population after exposure to a synthetic estrogen. Proceedings of the National Academy of Sciences of the United States of America, 104(21):8897-8901. http://www.pnas.org/
NatureServe, 2012. NatureServe Explorer: An online encyclopedia of life. Arlington, Virginia, USA: NatureServe. http://www.natureserve.org/
Nelson JS; Paetz MJ, 1992. The fishes of Alberta. Alberta, Canada: University of Alberta, 437 pp.
Paszkowski CA; Tonn WM, 1994. Effects of Prey Size, Abundance, and Population Structure on Piscivory by Yellow Perch, 123(6):855-865.
Price CJ; Tonn WM; Paszkowski CA, 1991. Intraspecific patterns of resource use by fathead minnows in a small boreal lake, 69:2109-2115.
Robison HW; Buchanan TM, 1984. Fishes of Arkansas. Arkansas, USA: University of Arkansas Press, 536 pp.
Smith RJF, 1978. Seasonal changes in the histology of the gonads and dorsal skin of the fathead minnow, Pimephales promelas, 56(10):2103-2109.
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31/12/12 Original text by:
Nicola Wakefield, CABI, UK
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