A review of the life history and biology
of Northern Pike Esox lucius Linnaeus1
Northern pike belong to the family Esocidae and are regarded as a top predator in most aquatic systems. They are considered a coolwater species (Casselman 1978) and are a popular freshwater game fish in most of their native range.
Northern pike have an attenuated fusiform body with the front of the head shaped much like a duck's bill; stout sharp teeth; prominent eyes; dorsal fin and anal fin set at posterior portion of body; mildly forked caudal fin; narrow caudal peduncle; lateral and dorsal coloration generally greenish with yellowish markings, darker on dorsal surface; whitish colored ventral surface, fins slightly marked but less so in juveniles. In California, anglers have confused northern pike with brown trout, Salmo trutta, lake trout, Salvelinus namaycush, and pike minnow Ptychocheilus grandis.
Northern pike distribution is holartic, ranging from northwestern Europe, across northern Asia, to northern North America. In North America they range from Alaska to Labrador, south through New England and much of New York, the northern part of the Ohio Valley, the Great Lakes region, and southward to Missouri and Nebraska. According to Dill and Cordone (1997), northern pike were supposedly brought to California in December 1891 by the U.S. Fish Commission. However, there is confusion regarding the identification and the fish may not have been northern pike but rather grass pickerel (Esox americanus vermiculatus). Neither species became established in California as a result of that introduction.
Northern pike are listed in the California Code of Regulations Title 14 Section 671 as a "detrimental animal" and it is unlawful to import, transport or possess northern pike alive except under permit. However, in 1988 an angler reported catching a 7-lb northern pike from Frenchman Lake, Plumas County. In 1991, Frenchman Lake was successfully chemically treated to remove northern pike. Subsequently, the species was collected from the Feather River and Lake Davis, Plumas County in 1992 and 1994, respectively. Both waters were chemically treated to removed northern pike but the species reappeared in Lake Davis in the summer of 1999, two years after treatment.
In it's native range, northern pike abundance varies and in northern Wisconsin lakes northern pike >35 cm (14 in.) was reported to averaged 16.1 fish/ha (40 fish/acre) and ranged from 2.8-38.0 fish/ha (7-94 fish/acre) (Margenau et al 1998).
Northern pike are adapted for shallow, moderately productive mesotrophic-eutrophic environments (Casselman 1996); usually select water less than 4 m deep, with vegetation, and within 300 m of shore (Diana et al 1977, Cook and Bergersen 1988); and aquatic vegetation has been considered crucial to northern pike success (Grimm and Klinge 1996) although some investigators have reported that there may be no relationship with biomass or number and littoral area (Margenau et al.1998). Juvenile pike were reported to occupy nearshore areas with abundant aquatic vegetation and emergent plants (McClane 1998); age-0 muskellunge, Esox masquinongy, a species similar to northern pike, were reported to occupy shallow habitats <1.5 m (5 ft) (Farrell and Werner (1999), adult northern pike were reported in or close to weed beds during summer in about 4 feet of water; in the fall move to precipitous shores, and in winter return to summer habitats or deep water (McClane 1998).
Northern pike location within a water has been suggested to be influenced by prey availability. In Alaska, northern pike were reported to live in the side sloughs and channels of moderate river systems (D. Rutz, Alaska Department of Fish and Game, transcripts of Lake Davis Steering Committee meeting, August 19, 1999). In Minnesota, northern pike were reported to prefer lake areas that have sedges, grasses, and wild rice (R. Pierce, Minnesota Department of Natural Resources, transcripts of Lake Davis Steering Committee meeting, August 25, 1999).
Pike have been reported to coexist perfectly in Alaskan rainbow trout lakes that have deep areas (mean depth 40-50 ft) and moderate to little amount of littoral area (D. Rutz, Alaska Department of Fish and Game, transcripts of Lake Davis Steering Committee meeting, August 19, 1999).
In Alaskan streams, northern pike were reported to fare poorly in clear, fast waters - but where the water is darker and slower, as in the Deshka or Alexander rivers, or in the side sloughs of the Sustina River itself, the species proliferates (Medred 1998).
Temperature
Upper lethal temperature for northern pike was reported to be 29o C (84o F), optimal growth occurs from 19-25o C (66-77o F) (Casselman 1978, Bevelhimer et al. 1985, Jacobson 1992), and upper lethal temperature based from measurements in different parts of U.S. was reported to be 30o C (86o F) (R. Pierce, Minnesota Department of Natural Resources, transcripts of Lake Davis Steering Committee meeting, August 25, 1999). Northern pike were reported to be susceptible to winter kill in small shallow lakes (Margenau et al. 1998).
Salinity
The species is salinity tolerant and occurs regularly in the Baltic Sea (Crossman 1979). In Alaska, northern pike were reported to have migrated downstream from the Sustina River, survived the brackish waters along the Sustina Flats, and moved into the Little Susitna River (Medred 1998). Northern pike were report caught by commercial set netters from the Cook Inlet, Alaska (salt water) (D. Rutz, Alaska Department of Fish and Game, transcripts of Lake Davis Steering Committee meeting, August 19, 1999).
Dissolved oxygen
Dissolved oxygen lower threshold for growth was reported to be 3 mg DO/l (Headrick and Carline 1993).
Ph
Ph typically ranges from 6.1 to 8.6 in northern pike lakes (Margenau et al. 1998)
Spawning occurs in the spring soon after ice out (Clark 1950, Franklin and Smith 1963). In the spring, individual northern pike may migrate from lakes into tributary streams (Miller 1948). Northern pike are reported to spawn in shallow marshes if available, typically with abundant flooded and aquatic vegetation, and spawning activity occurs mostly during afternoon in marshy areas of lakes or connected sloughs (Franklin and Smith 1963). Temperature, daily light intensity, and suitable vegetation was reported to stimulate spawning (Fabricius and Gustafson 1958). Spawning duration was reported to range from a few days to a month or more (Franklin and Smith 1963). Northern pike were reported to not use cattails for spawning, but rather select firm vegetative mats as good egg substrate; grasses, sedges, and rushes with fine leaves make best substrates (Franklin and Smith 1963).
Sexual maturity
Northern pike in the southern parts of its range spawn occasionally at one year of age, while the majority spawn initially at two years (McClane 1998). In Minnesota 1% were one year old, 84% were 2-5 years, and 15% were over 5 years of age at spawning. Spawning migrations are variable but pike have been reported to migrate up to 25 miles in streams.
Temperature
Water temperatures immediately following ice-out range from 32-40o F. Northern pike spawning and egg incubation were reported to occur at water temperatures from 45 to 60o F. (Franklin and Smith 1963).
Fecundity
The number of eggs produced by individual northern pike were expressed as Y = 4401.4X - 66,245 (Y = number of eggs and X = total length in inches) (Fanklin and Smith 1963). Female pike weighing 0.75 pound may produce 9-10,000 eggs, while a 10 pound female may produce 100,000 eggs (Lagler 1956). Highest densities of egg deposition related to vegetation (Fabricus and Gustafson 1958).
Embryo development
Embryo period was reported to range from 1½ to 2 weeks (Dorier 1938, Franklin and Smith 1963).
Growth at age varies with latitude (Figure 1). Northern pike are reported susceptible to stunting due to lack of prey (Diana 1996) and are typically associated with a high population densities (Casselman 1996). Slow growing high density populations are more common in Minnesota lakes <200 ha (494 acres) (Margenau et al 1998). When habitat is not limited and with removal (angling, netting and trapping), a compensatory response resulting in small-size individuals may be triggered (Colby et al 1987)
Northern pike length weight relationship is expressed in Figure 2. Maximum size was reported to several hundred pounds in Europe, while the Northern America all-tackle gamefish world record weighed 46 pounds 2 ounces and was caught in New York, USA in 1940. Typical state and line class records range from 20 to 40 pounds.
Northern pike demonstrate opportunistic feeding strategies and reflect species availability (Margenau et al 1998). Fusiform shaped prey are preferred, juvenile northern pike 10 - 12.5 mm (4-5 in.) reported to feed on microcrustacea and aquatic insects; juvenile pike feed on aquatic insects with increasing piscivorous as size increases; adults are very piscivorous but will include a variety of items in their diet including frogs, crayfish, ducklings (Lawler 1965, Frost 1954, Allen 1939, Hunt and Carbine 1950, Lux and Smith 1960, Lagler 1956).
Daily movement was reported to be variable in direction and magnitude of displacement (Diana et al 1977). Although not proven, Diana et al. (1977) suggested that pike populations may include both resident and nomadic individuals as in other fish populations (Funk 1957). Northern pike were most active during April and May and least active during October in a Colorado reservoir (Cook and Bergersen 1988).
Growth is reported to be related to northern pike density, water transparency, and abundance of small bluegills, while water temperatures >21oC limit growth and size structure (Margenau et al 1998).
In small Michigan lakes northern pike were ineffective in controlling an overabundance of bluegills (Beyerle 1971).
In southern Wisconsin centrarchid dominated waters, sunfish provide food for pike, but are not an optimal prey species, and northern pike have a tendency to stunt (T. Margenau, Wisconsin Department of Natural Resources, transcripts of Lake Davis Steering Committee meeting, August 19, 1999).
Northern pike fry are reported to be very adaptable, and "can live in a whole host of conditions" (R. Pierce, Minnesota Department of Natural Resources, transcripts of Lake Davis Steering Committee meeting, August 25, 1999).
Minnesota experienced "absolutely no success in angler control of northern pike populations" (R. Pierce, Minnesota Department of Natural Resources, transcripts of Lake Davis Steering Committee meeting, August 25, 1999).
Beyerle and Williams (1968) suggested that the results of observations on the food selectivity by northern pike increased doubts as to the effectiveness of pike for controlling pan fish populations in lakes, and suggested that soft-rayed food fishes favor good growth of northern pike.
"In those systems (Alaska) with shallow lakes and ponds, some of them have been completely devastated to the point where there are none left" (D. Rutz, Alaska Department of Fish and Game, transcripts of Lake Davis Steering Committee meeting, August 19, 1999, referring to northern pike predation on salmonids).
Goeman et al (1993) reported that northern pike removal in small Minnesota lakes though trap-netting and angler harvest was ineffective in altering population size structure.
REFERENCES
Allen, K.R. 1939. A note on the food of pike (Esox lucius L.) In Windermere, J. Animal Ecol. 8:72-75.
Beyerle, G.B. 1971. A study of two northern pike-bluegill populations. Trans. Am. Fish. Soc. 100:69-73.
Beyerle, G.B. and J.E. Williams. 1968. Some observations of food selectivity by northern pike in aquaria. Trans. Am. Fish Soc. 97:28-31.
Bevelhimer, M.S., R.A. Stein, and R.F. Carline. 1985. Assessing significance of physiological differences among three esocids with a bioenergetics model. Canadian Journal of Fisheries and Aquatic Sciences 42:57-69.
Casselman, J.M. 1978. Effects of environmental factors on growth, survival, activity, and exploitation of northern pike. Pages 114-128 in R.L. Kendall, editor. Selected coolwater fishes of North America. American Fisheries Society, Spec. Pub. 11, Bethesda, MD.
Casselman, J.M. 1996. Age, growth, and environmental requirements of pike. Pages 69-101 in J.F. Craig, editor, Pike: biology and exploitation. Chapman and Hall, London.
Clark, C.F. 1950. Observations on the early life history of the northern pike, Esox lucius L., in Houghton Lake, Michigan. Trans. Am. Fish. Soc. 71:149-164.
Colby, P.J., P.A. Ryan, D.H. Schupp, and S.L. Serns. 1987. Interactions in north-temperature lake fish communities. Canadian Journal of Fisheries and Aquatic Sciences 44(Supplement 2):104-128.
Cook, M.F. and E.P. Bergersen. 1988. Movements, habitat selection, and activity periods of northern pike in Eleven Mile Reservoir, Colorado. Trans. Am. Fish Soc. 117:495-502.
Crossman, E.J. 1979. Atlas of North American Freshwater Fishes.
Diana, J.S., W.C. Mackay, and M. Ehrman. 1977. Movements and habitat preferences of northern pike (Esox lucius) in Lac Ste. Anne, Alberta, Trans. Am. Fish. Soc. 106:560-565.
Dill, W.A. and A.J. Cordone. 1997. History and status of introduced fishes in California, 1871- 1996. Ca Dept Fish and Game, Fish Bull 178, 414 p.
Fabricius, E. and K.J. Gustafson. 1963. Some new observations on the spawning behavior of the pike, Esox lucius L. Fish Bd. Sweden, Inst. Freshwater Res. Drottningholm, 39:23-54.
Farrell, J.M. and R.G. Werner. 1999. Distribution, abundance, and survival of age-o muskellunge in Upper St. Lawrence River nursery bays. North American Journal of Fisheries Management 19:309-320.
Franklin, D.R. and L.L. Smith Jr. 1963. Early life history of Northern Pike, Esox lucius L., with special reference to the factors influencing the numerical strength of year classes. Trans. Am. Fish. Soc. 92:91-110.
Frost, W.E. 1954. The food of pike, Esox lucius, L., in Windermere. Jour. Animal Ecol. 23:339-360.
Funk, J.L. 1957. Movement of stream fishes in Missouri. Trans. Am. Fish. Soc. 85:39-57.
Goeman, T.J., P.D. Spencer, and R.B. Pierce. 1993. Effectiveness of liberalized bag limits as management tools for altering northern pike population size structure. NAJFM 13:621-624.
Grimm, M.P. and M. Klinge. 1996. Pike and some aspects of its dependence on vegetation. Pages 125-156, in J.F. Craig, editor, Pike: biology and exploitation. Chapman and Hall, London.
Headrick, M.R. and R.F. Carline. 1993. Restricted summer habitat and growth of northern pike in two southern Ohio impoundments. Trans. Am. Fish. Soc. 122:228-236.
Hickley, P. And A. Sutton. 1984. A standard growth curve for pike. Fisheries management 15:29-30.
Hunt, B.P. and W.F. Carbine. 1950. Food of young pike, Esox lucius, L., and associated fishes in Peterson's ditches, Houghton Lake, Michigan. Trans. Am. Fish. Soc. 80:67-83.
Jacobson, L.D. 1992. Analysis of factors affecting growth of northern pike in Minnesota, Minnesota, Dept. of Nat. Res. Section of Fisheries Investigational Report 424.
Lagler, K.F. 1956. Freshwater Fishery Biology. 2nd Edition, Wm. C. Brown Co. Publishers, Dubuque, Iowa.
Lawler, G.H. 1965. The food of pike Esox lucius, in Heming Lake Manitoba. J. Fish. Res. Bd. Canada 22:1357-1377.
Lux, F.E. and L.L. Smith. 1960. Some factors influencing seasonal changes in angler catches in a Minnesota lake. Trans. Am. Fish. Soc. 89:67-79.
Margenau, T.L., P.W. Rasmussen, and J.M. Kampa. 1998. Factors affecting growth of northern pike in small northern Wisconsin lakes. North American Journal of Fisheries Management 18:625-639.
McClane, A. J. (editor). 1998. McClane's New Standard Fishing Encyclopedia, Random House Value Publishing.
Medred, C. 1998. Killer Pike. Anchorage Daily News, May 31, 1998.
Miller, R.B. 1948. A note on the movement of the pike, Esox lucius. Copeia, 1948:62.
Figure 1. Length at age for northern pike (from Lagler 1956).
Figure 2. Length weight regression for northern pike.
1/ Prepared by Dennis P. Lee, Senior Fisheries Biologist, Fisheries Programs Branch, September 1999.