dcsimg

Behavior

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Perception Channels: tactile ; chemical

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Robin Street, University of Michigan-Ann Arbor
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Associations

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Adult walleye are top predators, which means that they do not have any natural predators in their habitat except humans. Humans do catch and eat adult walleye. The eggs and young fish are susceptible to predation by other fish such as white bass, muskellunge, white perch, largemouth bass, northern pike, and catfish. Young walleye avoid predation by staying near cover.

Known Predators:

  • white bass (Morone chrysops)
  • muskellunge (Esox masquinongy)
  • white perch (Morone americana)
  • largemouth bass (Micropterus salmoides)
  • northern pike (Esox lucius)
  • channel catfish (Ictalurus punctatus)
  • humans (Homo sapiens)
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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Robin Street, University of Michigan-Ann Arbor
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Morphology

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The name walleye refers to the glassy, large pupils of this fish; their white stare is a result of light reflected back through the pupil by crystalline matter in the retina. This allows the walleye to see extraordinarily well in darker waters. Walleyes are long and slim; brownish- green or silver above to creamy white below with dark stripes. The ventral lobe of the tail fin has a prominent white margin. Walleyes have large canine teeth. They have a large, visible, black spot at the base of the last three spines in the first of their two dorsal fins. They can reach 107 cm in length and can grow to 11 kg. Average weight is around 5 kg (Phillip, et al., 1982; Froese and Pauly, 2002)

Range mass: 2.0 to 10.0 kg.

Range length: 107.0 (high) cm.

Other Physical Features: bilateral symmetry

Sexual Dimorphism: sexes alike

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Robin Street, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Life Expectancy

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In southern areas, walleye may live 10 to 12 years but in northern waters they may live to be more than 20 years old

Range lifespan
Status: wild:
10.0 to 20.0 years.

Average lifespan
Status: wild:
18.0 years.

Average lifespan
Status: wild:
26.0 years.

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Robin Street, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Habitat

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Walleyes prefer deep lake and river water but will move into shallow flats to feed during early evening and night. (Tomelleri, 1990)

Range depth: 27.0 (high) m.

Habitat Regions: temperate ; freshwater

Aquatic Biomes: lakes and ponds; rivers and streams

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Robin Street, University of Michigan-Ann Arbor
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Distribution

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Walleye are native to the Nearctic Region. Walleye are abundant in many lakes and larger rivers over much of North America, from the Northwest Territories across Canada east of the Rocky Mountains to Labrador, southward along the Atlantic Coast to North Carolina, west to Arkansas, and north along the Missouri River. Their original range involved the Mississippi, Great Lakes, and St. Lawrence drainages, but they have been widely introduced into Atlantic, Pacific, and Gulf drainages.

Biogeographic Regions: nearctic (Native )

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Trophic Strategy

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Walleye are strictly carnivorous. Young walleye eat plankton. As they get older, they mostly eat fishes such as yellow perch and freshwater drum. They also eat insects, crayfish, snails, and mudpuppies (a kind of salamander). They even eat small mammals when fish and insects are not available. Feeding occurs at night. (Tomelleri, 1990; Froese and Pauly, 2002; Ontario Fishing Network, date unknown)

Primary Diet: carnivore (Eats terrestrial vertebrates, Eats non-insect arthropods)

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Associations

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Walleye are top predators. Once they reach adulthood, they primarily eat other animals and are not themselves eaten (except by people). They compete for food with other fish that are predators, including smallmouth bass and white perch.

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Benefits

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The walleye is perhaps the most sought after warm-water game fish and it supports a large fishing industry, particularly in the Central U.S. and Great Lakes area. (Phillips, et al., 1982)

Positive Impacts: food ; controls pest population

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Conservation Status

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Overall, walleye are not threatened or endangered. Populations of walleye are managed by humans as a game fish. One subspecies, Sander vitreus (blue pike) is believed to have gone extinct recently.

US Federal List: no special status

CITES: no special status

State of Michigan List: probably extirpated; no special status

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Untitled

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The walleye is the state fish of Minnesota and by far the most popular fish in that state. In southern areas, walleye may live 10 to 12 years but in northern waters they may live to be more than 20 years old (Phillips et al. 1982; Ontario Fishing Network, date unknown)

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Robin Street, University of Michigan-Ann Arbor
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Reproduction

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In the spring, Walleye migrate as far as 100 miles in search of prime spawning sites; they seem to prefer the rocky shores of lakes and dam ripraps. Males are not territorial and do not build nests. Spawning occurs at night. It involves one or two females and up to six males who chase, swim in circles, and erect their fins. They move to shallow water, where each female rolls on her side to release eggs while the males release sperms. A female deposits 20,000- 50,000 eggs per pound of body weight in a single night! Eggs typically hatch in twenty days. (Tomelleri, 1990; Phillips et al., 1982; Froese and Pauly, 2002; Ontario Fishing Network, date unknown)

Breeding interval: Breeding occurs once per year.

Breeding season: April through June

Range number of offspring: 16000.0 to 143000.0.

Average gestation period: 20.0 days.

Range age at sexual or reproductive maturity (female): 3.0 to 5.0 years.

Range age at sexual or reproductive maturity (male): 3.0 to 5.0 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; sexual

Average number of offspring: 150000.

The eggs are not placed in a nest or guarded, instead they drift in the water and become stuck among the rocks. Many of the eggs do not survive. When the baby fish (called fry)hatch, they are about 1 cm long and very hard to see in the water because they are so thin.

Parental Investment: no parental involvement

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Street, R. 2002. "Sander vitreus" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sander_vitreus.html
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Diagnostic Description

provided by Fishbase
Nuptial tubercles absent. Differentiation of sexes difficult. Branchiostegal rays 7,7 or 7,8 (Ref. 1998).
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Recorder
Estelita Emily Capuli
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Life Cycle

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Spawning occurs in small groups (a larger female and two smaller males or two females and up to six males) that engage in chasing, circular swimming, and fin erection. The group then ascends to shallow water, females roll on their side, and eggs and sperm are released. Deposition of eggs usually occurs in a single night (Ref. 1998). Larvae pelagic (Ref. 7471).
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Armi G. Torres
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Migration

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Potamodromous. Migrating within streams, migratory in rivers, e.g. Saliminus, Moxostoma, Labeo. Migrations should be cyclical and predictable and cover more than 100 km.
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Rainer Froese
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Morphology

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Dorsal spines (total): 13 - 17; Dorsal soft rays (total): 18 - 22; Analspines: 2; Analsoft rays: 11 - 14; Vertebrae: 44 - 48
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Estelita Emily Capuli
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Trophic Strategy

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Occurs in lakes, pools, backwaters, and runs of medium to large rivers. Prefers large, shallow lakes with high turbidity (Ref. 9988). Feeds at night, mainly on insects and fishes (prefers yellow perch and freshwater drum but will take any fish available) but feeds on crayfish, snails, frogs, mudpuppies, and small mammals when fish and insects are scarce (Ref. 1998). Although not widely farmed commercially for consumption, large numbers are hatched and raised for stocking lakes for game fishing (Ref. 9988).
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Pascualita Sa-a
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Biology

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Occurs in lakes, pools, backwaters, and runs of medium to large rivers. Frequently found in clear water, usually near brush (Ref. 86798). Prefers large, shallow lakes with high turbidity (Ref. 9988, 10294). Rarely found in brackish waters (Ref. 1998). Feeds at night, mainly on insects and fishes (prefers yellow perch and freshwater drum but will take any fish available) but feeds on crayfish, snails, frogs, mudpuppies, and small mammals when fish and insects are scarce (Ref. 1998). Although not widely farmed commercially for consumption, large numbers are hatched and raised for stocking lakes for game fishing (Ref. 9988). Utilized fresh or frozen; eaten pan-fried, broiled, microwaved and baked (Ref. 9988).
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Importance

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fisheries: commercial; aquaculture: experimental; gamefish: yes; aquarium: public aquariums
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Walleye

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The walleye (Sander vitreus, synonym Stizostedion vitreum), also called the yellow pike or yellow pikeperch or yellow pickerel,[3] is a freshwater perciform fish native to most of Canada and to the Northern United States. It is a North American close relative of the European zander, also known as the pikeperch. The walleye is sometimes called the yellow walleye to distinguish it from the blue walleye, which is a color morph that was once found in the southern Ontario and Quebec regions, but is now presumed extinct.[4] However, recent genetic analysis of a preserved (frozen) 'blue walleye' sample suggests that the blue and yellow walleye were simply phenotypes within the same species and do not merit separate taxonomic classification.[5]

In parts of its range in English-speaking Canada, the walleye is known as a pickerel, though the fish is not related to the true pickerels, which are members of the family Esocidae.[6]

Walleyes show a fair amount of variation across watersheds. In general, fish within a watershed are quite similar and are genetically distinct from those of nearby watersheds. The species has been artificially propagated for over a century and has been planted on top of existing populations or introduced into waters naturally devoid of the species, sometimes reducing the overall genetic distinctiveness of populations.

Etymology

Walleye, Sander vitreus

The name "walleye" comes from its pearlescent eyes caused by the reflective tapetum lucidum which, in addition to allowing the fish to see well in low-light conditions, gives its eyes an opaque appearance. Their vision affects their behavior. They avoid bright light and feed in low light on fish that cannot see as well as they do.[7] Many anglers look for walleyes at night since this is when major feeding efforts occur. The fish's eyes also allow them to see well in turbid waters (stained or rough, breaking waters), which gives them an advantage over their prey. Thus, walleye anglers commonly look for locations where a good "walleye chop" (i.e., rough water) occurs. Their vision also allows the fish to populate the deeper regions in a lake, and they can often be found in deeper water, particularly during the warmest part of the summer and at night.[8]

Description

Walleyes are largely olive and gold in color (hence the French common name: doré—golden). The dorsal side of a walleye is olive, grading into a golden hue on the flanks. The olive/gold pattern is broken up by five darker saddles that extend to the upper sides. The color shades to white on the belly. The mouth of a walleye is large and is armed with many sharp teeth. The first dorsal and anal fins are spinous, as is the operculum. Walleyes are distinguished from their close relative the sauger by the white coloration on the lower lobe of the caudal fin, which is absent on the sauger. In addition, the two dorsals and the caudal fin of the sauger are marked with distinctive rows of black dots which are absent from or indistinct on the same fins of walleyes.[9]

Length and weight

Weight and length of walleyes

Walleyes grow to about 80 cm (31 in) in length, and weigh up to about 9 kg (20 lb). The maximum recorded size for the fish is 107 cm (42 in) in length and 13 kilograms (29 lb) in weight. The rate depends partly on where in their range they occur, with southern populations often growing faster and larger. In general, females grow larger than males. Walleyes may live for decades; the maximum recorded age is 29 years. In heavily fished populations, however, few walleye older than five or six years of age are encountered. In North America, where they are highly prized, their typical size when caught is on the order of 30 to 50 cm (12 to 20 in), substantially below their potential size.

As walleye grow longer, they increase in weight. The relationship between total length (L) and total weight (W) for nearly all species of fish can be expressed by an equation of the form

W = c L b {displaystyle W=cL^{b},} W=cL^{b},

Invariably, b is close to 3.0 for all species, and c is a constant that varies among species. For walleye, b = 3.180 and c = 0.000228 (with units in inches and pounds) or b = 3.180 and c = 0.000005337 (with units in cm and kg).[10]

This relationship suggests a 50 cm (20 in) walleye will weigh about 1.5 kg (3.3 lb), while a 60 cm (24 in) walleye will likely weigh about 2.5 kg (5.5 lb).

Population dynamics

The Garrison Dam National Fish Hatchery at Garrison Dam, North Dakota, is the largest walleye hatchery in the world. Although they are in high demand for fishing and consumption in North Dakota, elsewhere they are considered a nuisance. For that reason GDNFH is also researching hormonal population control to provide control options to other areas.[11]

Reproduction

Walleye larva

In most of the species' range, male walleyes mature sexually between three and four years of age. Females normally mature about a year later. Adults migrate to tributary streams in late winter or early spring to lay eggs over gravel and rock, although open-water reef or shoal-spawning strains are seen, as well. Some populations are known to spawn on sand or vegetation. Spawning occurs at water temperatures of 6 to 10 °C (43 to 50 °F). A large female can lay up to 500,000 eggs, and no care is given by the parents to the eggs or fry. The eggs are slightly adhesive and fall into spaces between rocks. The incubation period for the embryos is temperature-dependent, but generally lasts from 12 to 30 days. After hatching, the free-swimming embryos spend about a week absorbing a relatively small amount of yolk. Once the yolk has been fully absorbed, the young walleyes begin to feed on invertebrates, such as fly larvæ and zooplankton. After 40 to 60 days, juvenile walleyes become piscivorous. Thenceforth, both juvenile and adult walleyes eat fish almost exclusively, frequently yellow perch or ciscoes, moving onto bars and shoals at night to feed. Walleye also feed heavily on crayfish, minnows, and leeches.

Taxonomy

The walleye is part of the North American clade within the genus Sander, alongside the sauger (S. canadensis). Hubbs described a taxon called the blue walleye (S. glaucus) from the Great Lakes but subsequent taxonomic work showed no consistent differences between this form and the "yellow" walleye and the blue walleye is now considered to be a synonym and color variant of the walleye.[12] The walleye was first formally described by the American naturalist Samuel Latham Mitchill (1764-1831) with the type locality given as Cayuga Lake near Ithaca, New York.[13]

As food

Fresh walleye being cooked over a fire

The walleye is considered to be a quite palatable freshwater fish, and consequently, is fished recreationally and commercially for food.[14] Because of its nocturnal feeding habits, it is most easily caught at night using live minnows or lures that mimic small fish. Most commercial fisheries for walleye are situated in the Canadian waters of the Great Lakes,[15] and fried walleye is considered a staple of Canadian cuisine.[16][17] In Minnesota, the walleye is often fished for in the late afternoon on windy days (known as a "walleye chop") or at night. Often served as a sandwich in Minnesota's pubs where the fish is very popular, deep fried walleye on a stick is a Minnesota State Fair food.[18]

Fishing

Because walleyes are popular with anglers, fishing for walleyes is regulated by most natural resource agencies. Management may include the use of quotas and length limits to ensure that populations are not overexploited. For example, in Michigan, walleyes shorter than 15 in (38 cm) may not be legally kept.

Since walleyes have excellent visual acuity under low illumination levels, they tend to feed more extensively at dawn and dusk, on cloudy or overcast days, and under choppy conditions when light penetration into the water column is disrupted. Although anglers interpret this as light avoidance, it is merely an expression of the walleyes' competitive advantage over their prey under those conditions. Similarly, in darkly stained or turbid waters, walleyes tend to feed throughout the day. In the spring and fall, walleyes are located near the shallower areas due to the spawning grounds, and they are most often located in shallower areas during higher winds due to the murkier, higher oxygenated water at around six feet deep.[19] On calm spring days, walleyes are more often located at the deep side of the shoreline drop-off and around shore slopes around or deeper than 10 feet.[20]

As a result of their widespread presence in Canada and the northern United States, walleyes are frequently caught while ice fishing, a popular winter pastime throughout those regions.

"Walleye chop" is a term used by walleye anglers for rough water typically with winds of 10 to 25 km/h (6 to 16 mph), and is one of the indicators for good walleye fishing due to the walleyes' increased feeding activity during such conditions. In addition to fishing this chop, night fishing with live bait can be very effective.

The current all-tackle world record for a walleye is held by Mabry Harper, who caught an 11.34-kg (25-lb) walleye in Old Hickory Lake in Tennessee on 2 August 1960.[21]

Cultural aspects

Large walleye statue at Lake Mille Lacs in Garrison, Minnesota

Walleye is a culturally significant food in the Upper Midwest.[22] Walleye is popular in Minnesota; the Minnesota Legislature declared walleye the official state fish in 1965. Three towns—Garrison, Minnesota, Baudette, Minnesota, and Garrison, North Dakota—each claim to be the "Walleye Capital of the World" and a large statue of the fish is erected in each town.[23] Walleye pike was declared the official "state warm water fish" of Vermont in 2012. (Vermont's official "state cold water fish" is the brook trout, salvelinus fontinalis.)[24]

South Dakota designated the walleye as its official state fish in 1982.[22] Although the fish is native to South Dakota, living in Missouri River reservoirs and eastern glacial lakes of the state, it only became a popular food in South Dakota in the 1970s and 1980s, when the fishing tournament circuit promoted the fish and operated walleye fishing contests in the state.[22]

The walleye is the official provincial fish of Manitoba.[25] Winnipeg, Manitoba, considers the walleye (referred to locally as "pickerel") its most important local fish.[26]: 76  Icelandic fishermen in Lake Winnipeg traditionally supplied the Winnipeg market.[26]: 23–26  The walleye is also the provincial fish of Saskatchewan, which declared the species its official fish in 2015 after it won a fish emblem contest.[27] Walleye is the most popular fish for sport fishing in Saskatchewan, and can be caught in many rivers, reservoirs, and lakes.[28] The International Underwater Spearfishing Association record for largest spearfishing-caught walleye is held by a 13.3-pound walleye caught in 2014 on the South Saskatchewan River north of Lake Diefenbaker.[27]

See also

References

  1. ^ NatureServe (2013). "Sander vitreus". IUCN Red List of Threatened Species. 2013: e.T202605A18229159. doi:10.2305/IUCN.UK.2013-1.RLTS.T202605A18229159.en. Retrieved 19 November 2021.
  2. ^ Froese, Rainer; Pauly, Daniel (eds.) (2019). Sander &speciesname= vitreum" Sander vitreum " in FishBase. December 2019 version.
  3. ^ "Ontario Freshwater Fishes Life History Database Species Detail".
  4. ^ "Le doré bleu existe!". lapresse.ca. 16 August 2008. Retrieved 24 March 2018.
  5. ^ Haponski, Amanda E.; Stepien, Carol A. (2014). "A population genetic window into the past and future of the walleye Sander vitreus: relation to historic walleye and the extinct "blue pike" S. v. "glaucus"". Giornale della Libreria. 14 (1): 133. doi:10.1186/1471-2148-14-133. PMC 4229939. PMID 24941945. Retrieved 10 July 2015.
  6. ^ Crossman, E.J. "Walleye - The Canadian Encyclopedia". Retrieved 29 April 2017.
  7. ^ "Walleye biology and identification". Minnesota Department of Natural Resources. Retrieved 29 May 2021.
  8. ^ Northern Wisconsin All-Outdoors Atlas & Field Guide. Sportsman's Connection. 2012. p. 5.
  9. ^ "In-Fisherman - The World's Foremost Authority On Freshwater Fishing". In-Fisherman. Retrieved 24 March 2018.
  10. ^ Anderson, R. O.; Neumann, R. M. (1996). "Length, Weight, and Associated Structural Indices". In Murphy, B. E.; Willis, D. W. (eds.). Fisheries Techniques (Second ed.). Bethesda, MD: American Fisheries Society. ISBN 1-888569-00-X.
  11. ^ Wilson, Malik (12 February 2021). "Garrison Dam National Fish Hatchery spearheading walleye population control project". KX NEWS. Retrieved 14 February 2021.
  12. ^ Carol A. Stepien & Amanda Haponski (2015). "Taxonomy, Distribution, and Evolution of the Percidae". In Patrick Kestemont; Konrad Dabrowski & Robert C. Summerfelt (eds.). Biology and Culture of Percid Fishes. Springer, Dordrecht. pp. 3–60. doi:10.1007/978-94-017-7227-3_1. ISBN 978-94-017-7227-3.
  13. ^ Eschmeyer, William N.; Fricke, Ron & van der Laan, Richard (eds.). "Perca vitrea". Catalog of Fishes. California Academy of Sciences. Retrieved 16 September 2020.
  14. ^ "Walleye, Sander vitreus". Department of Natural Resources (DNR), State of Michigan. Retrieved 15 March 2013.
  15. ^ "Walleye". Retrieved 25 April 2022. Seafood Source
  16. ^ "Best Fried Walleye". Retrieved 25 April 2022. Cook Me
  17. ^ "Best Fried Walleye". Retrieved 25 April 2022. All Recipes
  18. ^ "Field and Stream July 2005". July 2005.
  19. ^ Joe Fellegy, Jr., Walleyes and Walleye Fishing (Dillon Press, 1974), 57, 58
  20. ^ Fellegy, 60
  21. ^ International Game and Fish Association (1960). "IGFA All-Tackle World Record". IGFA. Retrieved 20 April 2014.
  22. ^ a b c Makenzie Huber, Why walleye? How this flaky fish became a South Dakota food favorite, Argus Leader (August 28, 2019).
  23. ^ "Walleyed War of the Walleye Capitals". RoadsideAmerica.com. Doug Kirby, Ken Smith, Mike Wilkins.
  24. ^ Vermont No. 82 of 2012.
  25. ^ "Official Emblems of Manibota" (PDF). Legislative Assembly of Manitoba. Archived (PDF) from the original on 20 April 2017.
  26. ^ a b Nicholson, Karen (May 2007). "A History of Manitoba's Commercial Fishery 1872-2005" (PDF). Manitoba Historic Resources Branch. Archived (PDF) from the original on 22 August 2017. Retrieved 8 May 2017.
  27. ^ a b "Saskatchewan's provincial symbols". Canadian Heritage. 15 August 2017. Retrieved 20 June 2022.
  28. ^ Michael Snook, Fishing Saskatchewan: An Angler's Guide to Provincial Waters (University of Regina Press: 2004), pp. 101-03.
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Walleye: Brief Summary

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The walleye (Sander vitreus, synonym Stizostedion vitreum), also called the yellow pike or yellow pikeperch or yellow pickerel, is a freshwater perciform fish native to most of Canada and to the Northern United States. It is a North American close relative of the European zander, also known as the pikeperch. The walleye is sometimes called the yellow walleye to distinguish it from the blue walleye, which is a color morph that was once found in the southern Ontario and Quebec regions, but is now presumed extinct. However, recent genetic analysis of a preserved (frozen) 'blue walleye' sample suggests that the blue and yellow walleye were simply phenotypes within the same species and do not merit separate taxonomic classification.

In parts of its range in English-speaking Canada, the walleye is known as a pickerel, though the fish is not related to the true pickerels, which are members of the family Esocidae.

Walleyes show a fair amount of variation across watersheds. In general, fish within a watershed are quite similar and are genetically distinct from those of nearby watersheds. The species has been artificially propagated for over a century and has been planted on top of existing populations or introduced into waters naturally devoid of the species, sometimes reducing the overall genetic distinctiveness of populations.

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