Feeding Habits and Diet of the Muskellunge

(Esox masquinongy): A Review of Potential

Impacts on Resident Biota

January 2016

Report prepared by

Steven J. Kerr

`for

Muskies Canada Inc.

and

Ontario Ministry of Natural Resources and Forestry

Suggested Citation: Kerr, S. J. 2016. Feeding Habits and Diet of the Muskellunge (Esox masquinongy): A

Review of Potential Impacts on Resident Biota. Report prepared for Muskies Canada Inc. and the Ontario

Ministry of Natural Resources and Forestry. Peterborough, Ontario. 24 p. + appendices.

Executive Summary

The Muskellunge (Esox masquinongy) is known as a voracious apex predator. In instances

where muskellunge are extending their range, either through intentional or inadvertent

introduction and natural range extension, concerns have been identified about the potential

negative impacts on resident fishes and aquatic biota. This review has been conducted to

assemble information on muskellunge predatory habits and diet as well as interspecific

competition with other species.

Muskellunge prey on a wide variety of organisms but prefer other fishes. Predation is based

largely on whatever species is available at the preferred size. There is a considerable amount

of evidence to indicate that Muskellunge prefer soft-rayed fishes and the availability of soft-

rayed prey could determine the degree of predation on other species.

Generally, there are few definitive studies to quantify impacts (if any) of Muskellunge on other

fish species. There is very little evidence to indicate that Muskellunge have a significant

negative impact on populations of other popular sport fish species including Walleye,

Largemouth Bass and Smallmouth Bass. In fact, there are numerous instances where these fish

species successfully co-habit the same waterbody. Since Muskellunge seldom occupy

coldwater habitats, their interactions with coldwater fishes (i.e., salmonids and coregonids) are

poorly understood. This is an area which requires future study.

Potential negative impacts of Muskellunge on other fish species are probably related to the size

of waterbody and the composition of the resident fish community. Larger waterbodies and

those waters having a diverse forage fish community seem to be relatively unaffected by the

presence of Muskellunge. The presence/abundance of soft-rayed fish species likely reduces the

predation on other resident fish species.

Other fish species can have negative impacts on the Muskellunge. Northern Pike are known to

have a competitive advantage over Muskellunge where they coexist. Young Muskellunge are

also subject to predation by other fishes including Largemouth Bass, Yellow Perch, Rock Bass

and Walleye.

Based on this literature review several recommendations are offered. These are related to

initiating more quantified studies to document impacts (if any) when Muskellunge are introduced

or become established in new waters, utilizing new state-of-the-art techniques to determine

diets and predatory-prey relationships amongst a broader range of fish community types

(including salmonids and species at risk), and developing efforts to improve the public

perception of Muskellunge.

(i)

Table of Contents

Executive Summary ..................................................................................................................... (i)

Table of Contents ......................................................................................................................... (ii)

Common and Scientific Names of Fishes Cited in the Text ........................................................ (iii)

Introduction..................................................................................................................................... 1

Distribution of Muskellunge in Canada ............................................................................. 1

Brief Summary of Muskellunge Life History ...................................................................... 1

Description of Issue ........................................................................................................... 2

Muskellunge Prey and Feeding Activity ......................................................................................... 2

General Diet ...................................................................................................................... 2

Diet of Young Muskellunge ............................................................................................... 3

Diet of Older Muskellunge ................................................................................................. 3

Prey Selection ................................................................................................................... 3

Bass ..................................................................................................................... 3

Bluegill and Crappies ........................................................................................... 4

Gizzard Shad ....................................................................................................... 4

Muskellunge ......................................................................................................... 4

Salmonids ............................................................................................................ 6

Walleye ................................................................................................................ 6

Yellow Perch ........................................................................................................ 7

Species at Risk .................................................................................................... 7

Seasonal Variation in Feeding Activity............................................................................. 7

Size of Prey ...................................................................................................................... 8

Interspecific Competition ................................................................................................................ 9

Information Gaps and Future Research Needs ........................................................................... 11

Acknowledgements ...................................................................................................................... 13

Personal Communications ........................................................................................................... 13

References and Literature Cited .................................................................................................. 13

General References ........................................................................................................ 13

Additional General References ....................................................................................... 16

Diet References .............................................................................................................. 16

Additional Diet References ............................................................................................. 19

Interspecific Competition References ............................................................................. 20

Additional Interspecific References ................................................................................. 23

Glossary ....................................................................................................................................... 24

Appendix 1. Fish Species Predated by Muskellunge.

(ii)

Common and Scientific Names of Fishes Cited in the Text

Alewife (Alosa pseudoharengus) Atlantic Salmon (Salmo salar) Banded Killifish (Fundulus diaphanous) Bigmouth Shiner (Notropis dorsalis) Black Bullhead (Ameiurus melas)

Redbreast Sunfish (Lepomis auritus) Red Shiner (Cyprinelia lutrensis) River Redhorse (Moxostoma carinatum) Rock Bass (Ambloplites rupestris) Shorthead Redhorse (Moxostoma macrolepidotum)

Black Crappie (Pomoxis nigromaculatus) Blacknose Dace (Rhinichthys atratulus) Blacknose Shiner (Notropis heterolepis) Bluegill (Lepomis macrochirus)

Silver Shiner (Notropis photogenis) Smallmouth Bass (Micropterus dolomieu) Spotfin Shiner (Cyprinella spiloptera) Spottail Shiner (Notropis hudsonius)

Bluntnose Minnow (Pimephales promelas) Telescope Shiner (Notropis telescopus) Brook Silverside (Labidesthes sicculus) Brook Trout (Salvelinus fontinalis) Brown Bullhead (Ameiurus nebulosis) Bull Trout (Salvelinus confluentus) Central Mudminnow (Umbra limi) Central Stoneroller (Campostoma anomalum) Chain Pickerel (Esox niger) Channel Catfish (Ictalurus punctatus) Common Carp (Cyprinus carpio)

Tessellated Darter (Etheostoma olmstedi) Tiger Muskellunge (Esox masquinongy x E. lucius) Trout-Perch (Percopsis omiscomaycus) Walleye (Sander vitreus) White Bass (Morone chrysops) White Crappie (Pomoxis annularis) White Sucker (Catostomus commersoni) Yellow Bullhead (Ameiurus natalis) Yellow Perch (Perca flavescens)

Creek Chub (Semotilus atromaculatus) Fathead Minnow (Pimephales promelas) Gizzard Shad (Dorosoma cepedianum) Golden Redhorse (Moxostoma erythrurum)

Golden Shiner (Notemigonus crysoleucas) Goldfish (Carassius auratus) Hornyhead Chub (Nocomis biguttatus) Iowa Darter (Etheostoma exile) Johnny Darter (Etheostoma nigrum) Lake Herring (Coregonus artedii) Lake Whitefish (Coregonus clupeaformis) Largemouth Bass (Micropterus salmoides) Logperch (Percina caprodes) Longnose Dace (Rhinichthys cataractae) Mooneye (Hidon tergisus) Mottled Sculpin (Cottus bairdi) Muskellunge (Esox masquinongy) Ninespine Stickleback (Pungitius pungitius) Northern Hogsucker (Hypentelium nigricans) Northern Pike (Esox lucius) Northern Pikeminnow (Ptychocheilus oregonensis)

Orangespotted Sunfish (Lepomis humilis) Pumpkinseed (Lepomis gibbosus) Rainbow Trout (Oncorhynchus mykiss) Redband Trout (Oncorhynchus mykiss gibbsi)

(iii)

Introduction

Distribution of Muskellunge in Canada

In Canada, the Muskellunge’s original range was limited to the provinces of Ontario and Québec (Scott and Crossman 1973). In Québec, Muskellunge are found in the Saint Lawrence River and southern Québec (north and south of the St. Lawrence River). In Ontario, Muskellunge are found in the Great Lakes as well as the southern and northwestern portions of the province (OMNR 1987, Kerr 2011). There have been some occurrences of Muskellunge in the Winnipeg River in the extreme eastern part of Manitoba (Stewart and Watkinson 2004) and, more recently, reports of muskellunge in Lake Winnipeg (Owen 2014).

Brief Summary of Life History

The Muskellunge spawns during the spring at water temperatures which usually range from 10-

15°C. Eggs are broadcast over submerged woody debris, aquatic vegetation and detritus

substrate. There is no nesting behaviour or parental care but there is strong evidence of

reproductive homing to the same spawning site (Crossman 1990, Kerr and Grant 2000).

Nursery habitats are usually characterized as shallow, vegetated areas which warm quickly.

Shortly after hatching, Muskellunge fry feed upon zooplankton. After reaching a size of

approximately 40 mm young Muskellunge switch to a piscivorous diet (Cook and Solomon

1987).

Muskellunge are sit-and-wait ambush predators which are sight oriented (Wisconsin DNR

2008). The predatory response of Muskellunge has two basic stages: (i) slow stalk with minimal

body movement, and (ii) explosive lunge at the prey item. They usually attack from the side and

impale their prey near their center mass – a point that moves least during escape attempts

(Webb 1984, Webb and Skadsen 1980). The prey item is subsequently rotated and swallowed

head first. Muskellunge rarely pursue their prey after a miss (Engstrom-Heg et al.1986).

Feeding activity generally peaks at water temperatures in the 18-20°C range and declines as

temperatures approach 30°C (Minnesota DNR undated). In the Rideau River, Ontario,

Muskellunge activity was lowest at dawn, increased throughout the day, peaked at dusk and

declined at night (Landsman et al. 2015). Since they rely heavily on sight to capture prey,

Muskellunge feeding activity is believed to be reduced at night and may be impaired in turbid

waters.

Adult Muskellunge are often closely associated with aquatic vegetation. Although they can

withstand temperatures as high as 32.2°C, they prefer temperatures in the 24-26°C range (Scott

1.

and Crossman 1973). Muskellunge can be sedentary or migratory. Migrations are usually

associated with spawning, searching for food or establishing summer and winter home ranges.

The maximum lifespan of a Muskellunge can approach thirty years. Muskellunge attain sexual

maturity at ages ranging from 3-4 years (males) and 4-5 years (females) (Parsons 1959).

Muskellunge display sexual dimorphism in that females grow faster, attain larger sizes and live

longer than males.

Description of Issue – Traditionally, there have been concerns that the introduction or

expansion of Muskellunge would have detrimental impacts on resident biota, particularly other

sport fish species such as bass, Walleye, trout and salmon (Seelbach 1988, Knapp et al. 2012,

Wolter et al. 2012, Midwood et al. 2015, Thomas et al. undated).

In many instances, proposed introductions of Muskellunge have been controversial because of

the belief that Muskellunge would compete with or prey upon other resident sport fishes

particularly those under rehabilitation (Bacigalupi et al. 2011, Wolter et al. 2012, Dan Taillon

personal communication). Planned introductions to new waters in MInnesota have encountered

some public resistance (www.dnr.state.mn.us/muskie/index.html) over concerns on impacts on

resident fish species. In Maine, there are concerns about the presence of Muskellunge having a

deleterious impact on resident Brook Trout and landlocked Atlantic Salmon (Brautigan and

Lucas 2008). The inadvertent introduction and expansion of Muskellunge in the Saint John

River, New Brunswick, is believed to potentially represent a threat to severely depleted stocks of

Atlantic Salmon (Curry et al. 2007, Kidd et al. 2011). In Manitoba, muskellunge appear to be

expanding their range westward moving down the Winnipeg River and into Lake Winnipeg

(Owen 2014).

This review was initiated in order to assemble available information on Muskellunge diet and

feeding activity in order to evaluate potential impacts as well as identify knowledge gaps

requiring further research.

Muskellunge Prey and Feeding Activity

Muskellunge predation can be evaluated based on a number of factors including prey selection,

size of prey, and seasonal variation in feeding activity.

General Diet –Generally, Muskellunge are considered to be solitary opportunistic predators

whose diverse diet is known to include crayfish, frogs, insects, tadpoles, snakes, fish,

2.

mudpuppies, ducklings and small mammals (muskrats, mice, shrews, etc.) (Hunter and Rankin

1939, Anderson 1948, Scott and Crossman 1973, Bozek et al. 1999). The bulk of their diet

consists of fish however (Scott and Crossman 1973) (see Appendix 1). In a northern Wisconsin

study involving 34 different lakes, Bozek et al. (1999) reported that fish comprised almost 98%

of the Muskellunge’s diet.

Muskellunge diet information from Canadian waters is very sparse. Many angled fish that don’t

recover when released have empty stomachs (John Casselman personal communication).

Diet of Young Muskellunge - The abundance and structure of the prey fish community

can strongly influence survival of young-of-the-year Muskellunge (Kapusincski and Farrell

2013). Young Muskellunge initially feed upon large zooplankton (usually cladocerans).

Applegate (1981) reported that, of five genera of invertebrates, Muskellunge fry preferred the

crustacean Moina brachiata over other species. They soon switch to larger prey when they

reach lengths of approximately 35-40 mm (MacCrimmon and Skobe 1970, Brautigan and Lucas

2008). Foraging success and growth of young Muskellunge are correlated with prey density

(Szendrey and Wahl 1996). Muskellunge grow most quickly during the first three years of life

(Wisconsin Department of Natural Resources 2008).

In the St. Lawrence and upper Niagara rivers, Banded Killifish, cyprinids and darters were the

most important prey for young-of-the-year Muskellunge (Kapuscinski 2011, Kapuscinski et al.

2012). In that study, young-of-the-year Muskellunge consumed more longer, fusiform-shaped

prey than laterally compressed prey. Krska and Applegate (1982) reported that young

Muskellunge, stocked in the Big Stone Power Plant in northeastern South Dakota, preyed

predominantly on Johnny Darter, Fathead Minnow, Bluegill and Orangespotted Sunfish.

Diet of Older Muskellunge - At a certain size, Muskellunge exhibit an ontogenetic shift in

diet. In the New River, Virginia, Brenden et al. (2004) found that at lengths from 800-900 mm,

Muskellunge changed from a diet consisting primarily of cyprinids to one comprised largely of

catostomids. Diet studies have shown that muskellunge growth rates were positively related

with abundance of minnows and suckers (Harrison and Hadley 1979, Hanson 1986). In Lake

Simcoe, MacCrimmon and Skobe (1970) reported that, although Muskellunge may be found at

depths of 10-15 meters, principal food items were species found in warm littoral or shoal waters.

It is widely believed that Muskellunge prefer cylindrical, soft-rayed prey over larger spiny- rayed

fishes (Oehmcke et al. 1958, Kerr and Grant 2000, Brown and Yokum 2012).

Prey Selection - The potential predatory impacts of Muskellunge on other fish species may

be summarized as follows:

Bass - Woomer et al. (2012) concluded that game fish, such as bass and Walleye, did not

comprise a large proportion of the Muskellunge diet. In Big Crooked Lake, Wisconsin,

Frey (2003) reported that Muskellunge predation on Smallmouth Bass was low. In a five

3.

year study involving seven Illinois lakes, Wolter et al. (2012) concluded that Muskellunge

stocking did not have any significant impacts on Largemouth Bass, Redear Sunfish

Gizzard Shad, or Common Carp populations. In another Illinois study, Storck and

Newman (1986) found that there was no evidence either in their study or in published

accounts to indicate that Tiger Muskellunge consumed substantial quantities of

Largemouth Bass. Brenden et al. (2004) found that Smallmouth Bass accounted for only

4% (by weight) of the Muskellunge diet and concluded that Muskellunge predation likely

had little overall impact on Smallmouth Bass in the New River, Virginia. A subsequent

study on the New River revealed that the most common diet items consisted of cyprinids,

sunfishes and suckers (Brittle 2014). Seelbach (1988) also concluded that Muskellunge

predation was unlikely to have a noticeable impact on Smallmouth Bass populations in

Michigan. Wolter et al. (2012) concluded that there were no negative effects of a

Muskellunge introduction on resident Largemouth Bass. Storck and Newman (1986)

found no evidence that Tiger Muskellunge consumed substantial quantities of Largemouth

Bass. Conversely, during a period of low Yellow Perch abundance and some strong year

classes of Largemouth Bass, Gammon and Hasler (1965) reported that bass fry and

fingerlings were prominent in the diet of Muskellunge.

Bluegill and Crappies – Bluegills and crappies were found to be the principal prey items

of Tiger Muskellunge stocked into Allerton Lake, Illinois (Storck and Newman 1986).

Bluegill were found to be less susceptible to Muskellunge predation than Fathead

Minnows (Moody et al. 1983). Wolter et al. (2012) reported that, when soft-rayed prey was

not available, Bluegill became the primary (>80%) prey item in the Muskellunge diet. They

also believed that some Black and White Crappie populations were sensitive to the

impacts of Muskellunge predation.

Gizzard Shad - Where present, Gizzard Shad seem to be a preferred prey for

Muskellunge perhaps because they are more easily captured than centrarchid prey (Wahl

and Stein 1991) (Table 1). Wahl and Stein (1988) concluded that growth and survival were

maximized when Muskellunge were stocked in systems with soft-rayed or fusiform prey

including cyprinids or Gizzard Shad. In the Susquehanna River, Pennsylvania, Deutsch

(1986) reported that the Common Carp and a variety of sucker species comprised the bulk

of the Muskellunge diet. Non-game fish species, such as Gizzard Shad, have been found

to be more vulnerable to esocid predation than game fishes (Weithman and Anderson

1977, Wahl and Stein 1991). They found that vulnerability to esocid predation was highest

for Gizzard Shad, Goldfish and Golden Shiners, intermediate for Bluegills and Largemouth

Bass and lowest for Channel Catfish. After a literature review, Hess and Heartwell (1978)

concluded that Gizzard Shad was probably the best forage for large esocids. In seven

Illinois lakes, Gizzard Shad overwhelmingly dominated in the diet of Muskellunge (Wolter

et al. 2012).

Muskellunge - Muskellunge are known to be cannabilistic (Kerr and Grant 2000). In

4.

Nogies Creek, Ontario, McNeil (1979) found that predation by two-year-old Muskellunge

on young-of-the year and yearling Muskellunge was significant and probably a factor

limiting Muskellunge production in that waterbody. Predation on stocked Muskellunge

fingerlings by older Muskellunge has been attributed as a cause of stocking failure

(Margenau 1992, Serns and Andrews 1986). Overall, however, it is generally not believed

that other esocids are an important dietary item of Muskellunge (Kerr and Grant 2000).

Table 1. Proportion of diet items of the Muskellunge and Tiger Muskellunge as determined from various

North American studies.

Location

Sample Size

(# Fish)

Diet Items

Reference

Corrine Lake (Wisconsin)

53 Yellow Perch – 62.3% Largemouth Bass - 3.8% Unidentified Fish - 32.1% Insects – 1.9%

Gammon and Hasler (1965)

Five Ohio reservoirs

Unknown Gizzard Shad (77-97% by weight)

Wahl and Stein (1991)

George Lake (Wisconsin)

24 Yellow Perch – 50.0% Largemouth Bass – 29.2% Smallmouth Bass – 4.2% Unidentified Fish 16.7%

Gammon and Hasler (1965)

Mingo Lake (Illinois)

Unknown Gizzard Shad - >95% Wolter et al. (2012)

Otter Lake (Illinois) Unknown

Gizzard Shad – ~90% White Bass – ~10%

Wolter et al. (2012)

Pierce Lake (IIlinois) Unknown Gizzard Shad – >85% Bluegill - <10% Yellow Perch <10%

Wolter et al. (2012)

Ridge Lake (Illinois) Unknown Bluegill – ~ 80-95% Largemouth Bass – ~15%

Wolter et al. (2012)

Sam Dale Lake (Illinois)

Unknown Gizzard Shad - >95% Wolter et al. (2012)

Shelbyville Lake (Illinois)

Unknown Gizzard Shad – 60-80% Bluegill - <10% Walleye - < 5%

Wolter et al. (2012)

St. John River (New Brunswick)

36 Insects – 11% Alewife – 3% White Perch – 3% White Sucker – 3% Unidentified Fish – 19% Empty Stomach – 61%

Stocek et al. (1999)

5.

Salmonids – Since the ranges of Muskellunge and salmonids seldom overlap there is

relatively limited information on predation by Muskellunge on salmonids. Baker et al.

(2015) documented evidence of some predation by Tiger Muskellunge on stocked

Rainbow Trout in Curlew Lake, Washington. When exposed to predation by introduced

Tiger Muskellunge, Lepak et al (2012) found that White Suckers had greater survival than

stocked Rainbow Trout. In that study, Tiger Muskellunge consumed greater proportions

(53-84% by mass) of stocked salmonids as compared to White Suckers (Lepak et al.

2014).

Tiger Muskellunge have been stocked to eradicate non-native Brook Trout in some high

elevation alpine lakes in North America. They have been found most successful in small,

lower elevation lakes not having an inlet or outlet (Koenig et al. 2015).

In the Mayfield Reservoir, Washington, Tipping (2001) reported that differences in habitat

selection by Tiger Muskellunge, Bull Trout and Redband Trout reduced their interaction

and the potential for predation.

Muskellunge coexist with Lake Trout and Lake Whitefish in Lake of the Woods and

several other northwestern Ontario lakes (Barry Corbett personal communication).

Similarly, Muskellunge formerly cohabitated in Lake Simcoe with Lake Trout, Lake

Herring and Lake Whitefish (MacCrimmon and Skobe 1970). Current Muskellunge

restoration efforts are not believed to conflict with management objectives for this

coldwater fish community (Buchanan and LeBeau 2000).

Relatively few studies have been conducted on predation of Atlantic Salmon. A study of

fish predation of newly stocked Atlantic Salmon in Maine (Warner et al. 1968) revealed

that Chain Pickerel was the worst predator. Thirty-one percent (N = 173) of the Chain

Pickerel sampled had consumed one or more of the recently stocked salmon. The study

did not include Muskellunge. Several researchers have, however, concluded that

Muskellunge represented only a minimal increase in predation for Atlantic Salmon in New

Brunswick (Curry et al. 2007, Landsman 2015). Anadromous salmon had probabilities of

less than 7% of being in the diet of Saint John River Muskellunge and less than 5% of

Muskellunge’s annual food intake occurred during the smolt migration (Curry et al. 2007).

It is worthy of note that a small population of landlocked Atlantic Salmon coexist with

Muskellunge in Trout Lake, Ontario (Fitchko et al. 1996, Bowman et al. 1997).

Walleye - Muskellunge are known to predate Walleye. In a Wisconsin study, a single

Muskellunge was believed to consume as many as twenty age 2-3 Walleye per year (2-

12 age 2-3 Walleye ha-1 year-1) (Gaeta et al. 2015, Jerme Gaeta, personal

communication). It was concluded that the ultimate effect of this rate of predation could

have series implications to the Walleye fishery.

There is, however, a considerable amount of evidence to indicate that Walleye and bass

are not preferred diet items of Muskellunge. During a four year Wisconsin study which

6.

involved examination of stomach contents from 1,092 Muskellunge, only five Walleye

were documented (Bozek et al. 1999). In a study to evaluate competition and predation

involving Walleye and four other sport fishes in a set of northern Wisconsin lakes, Fayram

et al. (2005) reported that direct competition or predation between Walleye and

Muskellunge was unlikely to be occurring. Waldera (undated) concluded that, although

Muskellunge may occasionally feed on Walleye, Walleye ranked extremely low in the diet

of a Muskellunge and their impact on Walleye populations was minimal. In Lake

Vermilion, Minnesota, Williams (undated) found that the introduction of Muskellunge had

not resulted in any major change to the resident Walleye population. Seelbach (1988)

concluded that the introduction of Muskellunge into several southern Michigan rivers was

unlikely to have a noticeable impact on valued game fishes such as Walleye and

Smallmouth Bass. In an Illinois study there was little predation on either Walleye or bass

(Wolter et al. 2012). Similarly, in a study involving 41 Minnesota lakes, there was no

significant decrease in mean abundance for any species, including Walleye, after

Muskellunge stocking (Knapp et al. 2008). They concluded that Walleye generally

coexisted well with Muskellunge in those lakes.

Yellow Perch – Yellow Perch are a common species which inhabit most Muskellunge

waters. Yellow Perch and catostomids were consistently found to be important in the diet

of Wisconsin Muskellunge (Bozek et al. 1999). Hourston (1952) reported that Yellow

Perch was the species more frequently eaten by Muskellunge in three different regions of

Ontario (Lake of the Woods, Kawartha Lakes and the St. Lawrence River). In the New

River, Virginia, the major species in the diet of Muskellunge was Yellow Perch and

Gizzard Shad (Brenden et al. 2004).

Species at Risk – In some instances there are concerns about the predatory impacts of

Muskellunge on other organisms which have been designated as species at risk (SAR) or

at low levels of abundance. Seelbach (1988) concluded that, since predator and prey

would be relatively rare (i.e., low densities), Muskellunge should not pose major threat.

There have been few, if any, other studies in this area.

Seasonal Variation in Feeding Activity – Food consumption by Muskellunge is known

to vary on a seasonal basis. Feeding occurs at temperatures ranging from 10 - 27° C (50 - 80°

F) but it usually peaks at temperatures near 20° C (70° F). Consumption rates decrease greatly

at temperatures below 14.5° C (40° F) and greater than 29.5° C (85° F) (Biesinger et al. 1979).

Bozek et al. (1999) reported that Muskellunge with food items in their stomach varied from

25.4% in the spring, 53.5% in the summer and 69.0% in the autumn. Wahl and Stein (1991)

found that the diet of young-of-the-year Muskellunge was dominated by Gizzard Shad in the

summer-early fall switching to sunfishes (Lepomis spp.) and Brook Silverside from late fall to

spring. In two Washington lakes, Rainbow Trout were more heavily predated in the spring than

at other times of year. Dombeck (1979) concluded that increased Muskellunge movements

during the autumn were related to increased feeding activity.

7.

Size of Prey - Muskellunge are ambush predators which seem to favour one large meal over

several smaller meals (Scott and Crossman 1973, Seelbach 1988, Sternburg 1996).

Muskellunge often remain inactive for long periods of time before resuming feeding activity

(Woomer et al. 2012). This is evidenced by the fact that, in many studies, the majority of

Muskellunge have empty stomachs (Table 2). Similar observations have been reported for

Northern Pike (Johnson 1969). In three Ohio reservoirs, Wahl and Stein (1993) found that

empty stomachs were common across all age groups of esocids.

Table 2. Muskellunge and Tiger Muskellunge feeding activity reported from selected North American

studies.

Waterbody

No. Fish Examined

Number of Fish (%) With Empty Stomachs

Reference

Allerton Lake 40 22 (55.0%) Storck and Newman (1986)

Big Stone Power Plant Reservoir (South Dakota)

107 60 (56.1%) Krska (1980), Krska and Applegate (1982)

Corrine Lake (Wisconsin)

106 56 (52.8%) Gammon and Hasler (1965)

Curlew Lake (Washington)

277 169 (61.0%) Baker et al. (2015)

George Lake (Wisconsin)

40 16 (40.0%) Gammon and Hasler (1965)

Lake of the Woods 40 33 (82.5%) Storck and Newman (1986)

New River (Virginia)

171 75 (43.9%) Brenden et al. (2004)

Silver Lake (Washington)

201 96 (48.0%) Baker et al. (2015)

Saint John River (New Brunswick)

36 22 (61.1%) Stocek et al. (1999)

Three regions in Ontario

202 121 (59.9%) Hourston (1952)

7 Illinois lakes

1,728 1,176 (68.1%) Wolter et al. (2012)

34 Wisconsin lakes 1,092 7170 (65.7%) Bozek et al. (1999)

Generally, the size of Muskellunge prey items increases as their size increases (Crossman

1962, Schmitz and Hetfield 1965, Gillen et al. 1981, Wahl and Stein 1993). Muskellunge can

consume prey fish ranging in size up to 47% of their own length (Bozek et al. 1999). Krska

8.

(1980) reported that there was a positive correlation of Muskellunge jaw width (closed) and prey

size (length and body depth). He also reported a correlation between Muskellunge body depth

and prey size. Kapuscinski et al. (2012) found that, as Muskellunge length increased, lengths of

laterally compressed spiny-rayed fish increased at a slower rate than did the lengths of fusiform

prey. Crossman (1962) concluded that, based on the wide variety of fish and other vertebrates

consumed, prey selection was largely size specific.

The growth of large Muskellunge may be impaired if food of an adequate size is not available in

spite of large numbers of smaller fish (Seelbach 1988, Wisconsin DNR 2008).

Interspecific Competition

Interspecific competition and impacts of Muskellunge on the forage base have been identified

as concerns amongst fisheries managers (Kerr 2011). A review of interspecific competition,

involving Muskellunge, was conducted by Kerr and Grant in 2000. They concluded that any

potential impacts of Muskellunge were largely based on predation of resident biota and

competition with Northern Pike.

It is important to realize that Muskellunge coexist with many other popular sport fishes. Based

on a review of 373 waterbodies from Ontario’s Aquatic Resource Area (ARA) database, several

different sport fish species commonly coexist in the same waterbodies as Muskellunge. For

example, three-quarters of all inland muskellunge waters also contain self-sustaining

populations of Smallmouth Bass (Table 3).

Table 3. Ontario Muskellunge waters (excluding the Great Lakes and connecting channels) which are

also inhabited by other sport fishes (Source: Ontario ARA database)..

Number of Waterbodies (% of Total)

Sport Fish Species Lakes Rivers/Streams Total

Black Crappie 49 15 64 (17.2%) Largemouth Bass 139 18 157 (42.1%) Northern Pike 169 28 197 (52.8%) Smallmouth Bass 253 31 284 (76.1%) Walleye 205 30 235 (63.0%) Coldwater species* 81 3 84 (22.5%)

* Includes salmonids and coregonids

Young Muskellunge are susceptible to predation by a variety of other fish species including

Largemouth Bass (Stein et al. 1981, John 1982, Carline et al. 1986), Yellow Perch (Murry and

Farrell 2007), Northern Pike (Serns and Andrews 1983, Margenau and Snow 1984) and Rock

Bass (Murry and Farrell 2007). Stocked Tiger Muskellunge have been found to be heavily

9.

predated by Largemouth Bass (Stein et al. 1981). In Missouri, the poor survival of stocked

Muskellunge has been attributed to predation on young fish by Largemouth Bass and other

predators (http://mdc.mo.gov/discover-nature/field-guide/muskellunge-0). Wahl (1995) reported

that Muskellunge were more susceptible to Largemouth bass predation than were Walleye.

There is some evidence for negative impacts, through predation, of Muskellunge on resident

fishes. In two small Wisconsin lakes, Gammon and Hasler (1965) reported that the introduction

of Muskellunge caused a dramatic decline in the abundance of Yellow Perch. The introduction

of Muskellunge to Iron Lake, Michigan, resulted in the decline of prey fishes such as White

Suckers and Black Crappie (Siler and Beyerle 1986). In two Wisconsin lakes, interactions

between Muskellunge and Largemouth Bass (which occupy similar habitats) were believed

responsible for a decline in Largemouth Bass (Becker 1983).

Conversely, in a study involving forty-one Minnesota lakes which included populations of

Northern Pike, Walleye, Yellow Perch, Lake Herring, Black Crappie, White Sucker and Bluegill,

Knapp et al. (2012) found no significant decreases among seven different fish species after the

introduction of Muskellunge. They concluded that the various fish species coexisted well with

Muskellunge at the densities that resulted from stocking. Inskip and Magnuson (1983) reported

on the stability of the Big Pine Lake, Wisconsin, fish community, which included Muskellunge,

Walleye and bass, over a forty year period. Weithman and Anderson (1977) found that non-

game fishes were more vulnerable than game species to Muskellunge predation in holding

tanks. The Minnesota Department of Natural Resources concluded that Muskellunge coexisted

with healthy populations of Walleye and other game fishes (Minnesota Department of Natural

Resources 2015). Colby et al. (1987) speculated that, in northern coolwater fish communities,

the abundance of percids, such as Walleye, depended on how well apex predators (such as

Muskellunge and Northern Pike) controlled the abundance of Common White Suckers and

Bluegill.

There is considerable interest regarding Muskellunge and Walleye interactions. In a project

involving a set of Wisconsin lakes, Fayram et al. (2005) studied species which had the greatest

interaction with Walleyes and concluded that Muskellunge-Walleye interactions (competition

and predation) were not significant. In some instances, there is evidence to suggest that high

Walleye densities can influence Muskellunge abundance. In a study involving 17 Wisconsin

lakes, Klingbiel (1966) concluded that there was an inverse relationship between the size of the

resident Walleye population and the proportion of stocked Muskellunge in the harvest.

Probably one of the most debated interactions involves Muskellunge and Northern Pike (Inskip

1986). Oehmcke (1969) considered Northern Pike to be a direct competitor with Muskellunge

and was ultimately the dominant of the two species. After the introduction of Northern Pike to

Big Pine Lake, Wisconsin, the abundance of Muskellunge declined (Inskip and Magnuson

1983). Since Northern Pike spawn and hatch earlier than Muskellunge, it is widely believed that

pike fry predate the younger and smaller Muskellunge (Threinen and Oehmcke 1950). These

10.

two species have many similar behavioural characteristics (Wahl and Stein 1993). They are

known to compete for food, especially in their first year of life, as well as cover. The two species

hybridize naturally (Eddy 1944). Although their ranges overlap, Muskellunge seem to prefer

warmer water temperatures and more lotic conditions than Northern Pike.

At older life stages, Muskellunge and Northern Pike display evidence of spatial and/or temporal

segregation. In the Niagara River, Harrison and Hadley (1978, 1979) speculated that

Muskellunge and Northern Pike were spatially segregated during the spawning period with

Northern Pike preferring lentic waters and Muskellunge seeking lotic conditions. Similar

observations were reported by Osterberg (1981) and Cooper et al. (2008) on the St. Lawrence

River and in Leech Lake, Minnesota (Strand 1986).

Muskellunge and Tiger Muskellunge have been used as a management tool to improve

predator-prey balance and enhance the growth rate of prey species. In some Illinois waters,

stocking of Tiger Muskellunge to reduce Bluegill density was considered unsuccessful (Storck

and Newman 1986). Similar failures have been reported in Wisconsin (Snow 1988). Failures of

stocked Muskellunge to control overabundant panfish populations has been related to the

relatively low density of Muskellunge, their preference for larger prey fish, and their habit of

being inactive for long periods of time before resuming feeding (Woomer et al. 2012). Stocked

Muskellunge have been successful in reducing stunted perch populations, however (Gammon

and Hasler 1965, Oehmcke 1969).

Information Gaps and Future Research Needs

Based on this review, several information gaps were identified:

Unpublished Diet Information - Despite the fact that this survey revealed a relative shortage

of published literature pertaining to muskellunge diet, it is believed that there is additional

information which has been accumulated on some of the larger, more heavily utilized

muskellunge fisheries. For example, in Ontario this could include Lake of the Woods (Tom

Mosindy, personal communication), Georgian Bay (Arunas Liskauskas personal

communication) and the Kawartha lakes. The collection of this information was beyond the

scope of this project.

Efforts are required to identify sources of unpublished data regarding muskellunge diet

in order to summarize and consolidate this information.

Impacts of Muskellunge on Salmonids - Most of the diet information available comes from

waters having coolwater and warmwater fish communities. Since their natural ranges are

almost mutually exclusive, relatively few studies have been conducted of Muskellunge predation

11.

on Atlantic Salmon. Curry et al. (2007) concluded that it was highly improbable that newly

established Muskellunge would have a significant impact on migratory Atlantic Salmon in the

Saint John River, New Brunswick. Bozek et al. (1999) recommended that future studies needed

to evaluate Muskellunge diet across a suite of waterbodies having different fish communities.

Future studies are required to evaluate feeding habits and diet in coldwater habitats,

involving both lakes and tributaries, to ascertain interspecific relationships with species

including trout, salmon and coregonids.

Impacts of Muskellunge on Species at Risk – It is possible that increased predation from an

introduced Muskellunge population could further impair the status of species at risk such as

River Redhorse or American Eel. There are few, if any, definitive studies on this topic. Similarly,

there seems to be little quantification of the effects of Muskellunge on suppressed populations

of fish species such as Walleye.

The examination of potential impacts of Muskellunge on species at risk or species at low

levels of abundance in waters where they coexist needs further study.

New Tools for Determining Prey Consumption – Traditional techniques for determining diet

of fishes, including Muskellunge, have involved physical removal of stomach contents or

examination of fecal content (see reviews by Kamler and Pope, 2001 and Kerr 2015). These

techniques have many shortcomings largely due to variation in digestive rates (Sheppard and

Hardwood 2005).

Over the past number of years, new techniques such as DNA analysis and the use of stable

isotopes have been developed to investigate predator-prey food webs (Carreon-Martinez and

Heath 2010).

Future diet studies should utilize new and alternative techniques, such as stable

isotopes and DNA analysis, for diet analysis and determination of trophic relationships.

Public Perception of Muskellunge – For many anglers, Muskellunge are a highly prized

species, yet the public perception about the impacts of Muskellunge on other fish species has

been identified as a management challenge (Midwood et al. 2015). In addition, generalist

anglers are more likely to harvest a Muskellunge (Margenau and Petchenik 2001).

More efforts are required to disseminate results of new diet and ecological impact

studies which may be conducted in the future.

12.

Acknowledgements

Kevin Collins, Ontario Ministry of Natural Resources and Forestry, provided access to the

province’s ARA database. I am also grateful for constructive editorial comments received from

Davin Heinbuck, Brenda Koenig, Peter Levic and Dan Taillon. This project was sponsored

cooperatively by Muskies Canada Incorporated (MCI) and the Ontario Ministry of Natural

Resources and Forestry (MNRF).

Personal Communications

Casselman, John, Queens University, Kingston, Ontario.

Collins, Kevin, Ontario Ministry of Natural Resources and Forestry, Peterborough, Ontario.

Corbett, Barry, Ontario Ministry of Natural Resources and Forestry (retired). Kenora, Ontario.

Gaeta, Jerme. Utah State University. Logan, Utah.

Liskauskas, Arunas. Ontario Ministry of Natural Resources and Forestry. Owen Sound, Ontario.

Mosindy, Tom, Ministry of Natural Resources and Forestry (retired). Kenora, Ontario.

Taillon, Dan. Ministry of Natural Resources and Forestry, Peterborough, Ontario.

Zelman, Kaleb. University of New Brunswick. Fredericton, New Brunswick.

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changes on the segregation of northern pike (Esox lucius) and muskellunge (Esox masquinongy)

in the upper St. Lawrence River. Hydrobiologia 601:41-53.

Crossman, E. J. 1962. Predator-prey relationships in pikes (Esocidae). Journal of the Fisheries Research Board of Canada 19:979-980.

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Eddy, S. 1944. Hybridization between northern pike (Esox lucius) and muskellunge (Esox masquinongy).

Proceedings of the Minnesota Academy of Science 12:28-43.

Fayram, A. H., M. J. Hansen and T. J. Ehlinger. 2005. Interactions between walleyes and four fish species with implications for walleye stocking. North American Journal of Fisheries Management 25:1321-1330. Frey, A. P. 2003. Trophic interactions between walleye and smallmouth bass in a north temperate lake. M.Sc. Thesis. University of Wisconsin. Stevens Point, Wisconsin. Gaeta, J. W., M. J. Vander Zanden, G. Hansen and S. R. Carptenter. 2015. Muskellunge (Esox masquinongy) predation on walleye (Sander vitreus) affects population size structure and harvest: A modelling approach. Presentation at the 145

th Annual Meeting of the American

Fisheries Society. August 16-20, 2015. Portland, Oregon. Gammon, J. R. and A. D. Hasler. 1965. Predation by introduced muskellunge on perch and bass, I:Years 1-5. Wisconsin Academy of Sciences, Arts and Letter 54:249-272. Harrison, E. J. and W. F. Hadley. 1978. Ecologic separation of sympatric muskellunge and northern pike.

American Fisheries Special Publication 11:129-134.

Inskip, P. D. 1986. Negative associations between abundances of muskellunge and northern pike:

Evidence and possible explanations. American Fisheries Society Special Publication15:135-150.

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Lake, Wisconsin. Transactions of the American Fisheries Society 112:378-389.

John, L. D. 1982. Factors affecting short term survival of stocked muskellunge fingerlings in Wisconsin.

Research Report 117. Wisconsin Department of Natural Resources. Madison, Wisconsin. 24 p.

Kapuscinski, K. L. 2011. Comparative ecology of muskellunge and nearshore fish assemblages in the Great Lakes. Ph.D. Dissertation. State University of New York. Syracuse, New York. 191 p. Kerr, S. J. and R. E. Grant. 2000. Ecological impacts of fish introductions: Evaluating the risk. Fish and Wildlife Branch. Ontario Ministry of Natural Resources. Peterborough, Ontario. 473 p. Klingbiel, J. 1966. An evaluation of stocking large muskellunge fingerlings. Fisheries Management

Report No. 3. Wisconsin Department of Natural Resources. Madison, Wisconsin.11 p.

Knapp, M. L., S. W. Mero, D. J. Bohlander and D. F. Staples. 2008. Fish community response to the introduction of muskellunge in Minnesota lakes. Special Publication 166. Minnesota Department of Natural Resources. Brainerd, Minnesota. 22 p. Knapp, M. L., S. W. Mero, D. J. Bohlander, D. F. Staples and J. A. Younk. 2012. Fish community responses to the introduction of muskellunge into Minnesota lakes. North American Journal of Fisheries Management 32:191-201. Margenau, T. L. and H. Snow. 1984. An evaluation of muskellunge stocking in Murphy Flowage.

Research Report 128. Wisconsin Department of Natural Resources. Madison, Wisconsin. 10 p.

Margenau, T. L. and J. B. Petchenik. 2004. Social aspects of muskellunge management in Wisconsin.

North American Journal of Fisheries Management 24:82-93.

21.

Minnesota Department of Natural Resources. 2015. Muskellunge management: New lakes proposed http://www.dnr.state.mn.us/muskie/index.html

Murry, B. A. and J. M. Farrell. 2007. Quantification of native muskellunge nursery habitat: Influence of

body size, fish community composition and vegetation structure. Environmental Biology of Fishes

79:37-47.

Oehmcke, A. A. 1969. Muskellunge management in Wisconsin. Fisheries Report No. 19. Wisconsin

Department of Natural Resources. Madison, Wisconsin.

Osterberg, D. M. 1981. Spawning and early life history of the Great Lakes muskellunge (Esox

masquinongy) in the international section of the St. Lawrence River. Project Progress Report.

State University of New York. Potsdam, New York.

Seelbach, P. W. 1988. Considerations regarding the introduction of muskellunge into southern Michigan

rivers. Fisheries Division Technical Report 88-5. Michigan Department of Natural Resources.

Lansing, Michigan. 14 p.

Serns, S. L. and L. M. Andrews. 1983. Survival and growth of muskellunge fingerlings stocked in four

Vilas County, Wisconsin, lakes. Fisheries Management Report 116. Wisconsin Department of

Natural Resources. Madison, Wisconsin. 15 p.

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Michigan. American Fisheries Society Special Publication 15:229-237.

Snow, H. E. 1988. Effects of the introduction of muskellunge and walleye on bluegill and other species in

Clear Lake, Sawyer County, Wisconsin, 1959-1984.Research Report 147. Wisconsin Department

of Natural Resources. Madison, Wisconsin. 17 p.

Stein, R. A., R. F. Carline and R. S. Hayward. 1981. Largemouth bass predation on stocked tiger

muskellunge. Transactions of the American Fisheries Society 110:604-612.

Storck, T. W. and D. L. Newman. 1986. Evaluation of the introduction of tiger muskellunge into

impoundments dominated by the bass-bluegill combination, evaluation of a partial creel and size-

specific survival of stocked channel catfish. Technical Report 1986-4. Aquatic Biology Section,

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Wolter, M. H., C. S. DeBoom, C. P. Wagner, M. J. Diana and D. H. Wahl. 2012. Evaluation of growth and

survival of different genetic stocks of muskellunge: Implications for stocking programs in Illinois

and the Midwest. INHS Technical Report 2012-33. Illinois Natural History Survey, University of

Illinois. Champaign, Illinois.71 p.

22.

Additional Interspecific Competition References

Caplan, D. L. 1982. An experimental study of interactions between young of the year pike (Esox lucius) and muskellunge (Esox masquinongy). M.Sc. Thesis. University of Wisconsin. Madison, Wisconsin. 57 p.

Carey, M. P. and D. H. Wahl. 2010. Interactions of multiple predators with different foraging modes in an aquatic food web. Oecologia 162:443-452.

Gammon, J. R. and A. D. Hasler. 1961. A predator-prey interaction: The effect of muskellunge on the

resident fish populations of two big lakes. Paper 354. Proceedings of the American Society of

Limnology and Oceanography. Washington, D. C.

Graff, D. R. 1972. Observations on interactions between juveniles of Esox lucius and Esox masquinongy.

Pennsylvania Cooperative Fishery Research Unit. State College, Pennsylvania. 79 p.

23.

Glossary

Ambush Predator – An organism whose foraging strategy is based on a “sit-and-wait”

approach. They capture or trap prey by stealth rather than by speed or by strength.

Biota - The total collection of organisms of a geographic region or a time period. The biota, or

biotic component of the Earth make up the biosphere.

Coolwater Fish Community – Ecosystems which are mesothermal (intermediate between cold

and relatively unproductive coldwater and warmer, more nutrient-rich warmwater ecosystems).

Coldwater fish communities are usually characterized by species such as Walleye, Northern

Pike and Muskellunge.

Cyprinids - Any of numerous often small freshwater fishes of the family Cyprinidae, which

includes the minnows, carps, and shiners.

Fusiform - Having a spindle-like body shape that is wide in the middle and tapers at both ends.

Lentic – Still or standing water such as lakes and ponds.

Lotic - Flowing water. Lotic waters can range from springs only a few centimeters wide to

major rivers kilometers in width.

Ontogenetic – History of the development of an organism.

Piscivorous – Feeding on fish.

Sexual Dimorphism - Differences in appearance between males and females of the same

species, such as in colour, shape, size, and structure.

Warmwater Fish Community – Ecosystems characterized by warm year-round temperatures,

nutrient rich (mesotrophic to eutrophic) and characterized by fish species having relatively high

thermal preferences.

Zooplankton - Plankton are living organisms drifting in bodies of water. Plankton collectively

refers to both phytoplankton (plants) and zooplankton (animals). Individual zooplankton are

usually microscopic, but some are larger and visible with the naked eye. Many forms of

zooplankton are important prey for young fish.

24.

Appendix 1. Fish species predated by Muskellunge.

Fish Species Reference

Alewife

Stocek et al. (1999), Brautigan and Lucas (2008), (Zelman personal communication).

Atlantic Salmon Curry et al. (2007)

Banded Killifish

Kerr and Grant (2000), Kapuscinski et al. (2012)

Bass (Micropterus spp.)

Hourston (1952)

Bigmouth Shiner

Bozek et al. (1999)

Black Bullhead

Bozek et al. (1999)

Black Crappie

Wolter et al. (2012)

Blacknose Dace

Bozek et al. (1999)

Blacknose Shiner

Bozek et al. (1999)

Bluegill

Weithman and Anderson (1977), Krska (1980), Gillen et al. (1981), Krska and Applegate (1982), Moody et al. (1983), Tomcko et al. (1984), Storck and Newman (1986), Wahl and Stein (1991, 1993), Bozek et al. (1999), Kerr and Grant (2000), Brenden et al. (2004). Wolter et al. (2012)

Bluntnose Minnow

Gammon (1963), Wahl and Stein (1991), Kerr and Grant (2000)

Brook Silverside

Wahl and Stein (1991,1993), Kerr and Grant (2000), Wolter et al. (2012)

Brook Trout

Koenig et al. (2015)

Brown Bullhead

Kerr and Grant (2000)

Bullheads and Catfishes (Ictalurus spp. and Ameiurus spp.)

Hourston (1952), MacCrimmon and Skobe (1970), Scott and Crossman (1973), Brautigan and Lucas (2008)

Central mudminnow

Bozek et al. (1999)

Central Stoneroller

Brenden et al. (2004)

Channel Catfish

Weithman and Anderson (1977), Kerr and Grant (2000)

Common Carp Weed (1925), Axon (1978), Deutsch (1986), Cook and Solomon (1987), Bozek et al. (1999), Kerr and Grant (2000), Woomer et al. (2012)

Coregonids (Coregonidae)

Oehmcke (1969)

Fish Species

Reference

Creek Chub Bozek et al. (1999)

Cyprinids (Cyprinidae) Anderson (1948), Hourston (1952), Brewer (1960), MacCrimmon and Skobe (1970), Scott and Crossman (1973), Krska (1980), Webb and Skadsen (1980), Gillen et al. (1981), Cook and Solomon (1987), Seelbach (1988), Wahl and Stein (1993), Brautigan and Lucas 2008), Kapuscinski et al. (2012), Brittle (2014)

Fathead Minnows Krska (1980), Webb and Skadsen (1980), Krska and Applegate (1982), Tomcko et al. (1984), Moody et al. (1983), Kerr and Grant (2000), New et al. (2001)

Gizzard Shad

Anderson (1948), Hourston (1952), Weithman and Anderson (1977), Axon (1978), Hess and Heartwell (1978), Storck and Newman (1986), Cook and Solomon (1987), Wahl and Stein (1991, 1993), Kerr and Grant (2000), Brenden et al. (2004), Brautigan and Lucas (2008), Wolter et al. (2012)

Golden Redhorse

Seelbach (1988), Kerr and Grant (2000)

Golden Shiner Weithman and Anderson (1977), Bozek et al. (1999), Kerr and Grant (2000), Wagner (2012)

Goldfish

Weithman and Anderson (1977)

Hornyhead Chub

Bozek et al. (1999)

Iowa Darter

Bozek et al. (1999)

Johnny Darter

Krska (1980), Krska and Applegate (1982), Bozek et al. (1999), Kerr and Grant (2000)

Lake Herring Bozek et al. (1999), Kerr and Grant (2000), Simonson (2012), Williams (undated)

Lake Whitefish

Kerr and Grant (2000), Brautigan and Lucas (2008)

Largemouth Bass Gammon and Hasler (1965), Weithman and Anderson (1977), Storck and Newman (1986), Wolter et al. (2012), Baker et al. (2015)

Logperch

Bozek et al. (1999)

Longnose Dace Bozek et al. (1999)

Mooneye Hourston (1952), Scott and Crossman (1973)

Mottled Sculpin

Bozek et al. (1999)

Fish Species Reference

Muskellunge

Serns and Andrews (1986), Margenau (1992), Bozek et al. (1999)

Ninespine Stickleback

Bozek et al. (1999)

Northern Hog Sucker Bozek et al. (1999), Brenden et al. (2004)

Northern Pike

Bozek et al. (1999)

Northern Pikeminnow

Baker et al. (2015)

Orangespotted Sunfish

Krska (1980), Krska and Applegate (1982), Kerr and Grant (2000)

Pumpkinseed

Bozek et al. (1999), Kerr and Grant (2000)

Rainbow Trout

Baker et al. (2015)

Redbreast Sunfish

Brenden et al. (2004)

Redhorse (Moxostoma spp.) Hourston (1952)

Red Shiner

Wolter et al. (2012)

Rock Bass

Hourston (1952), Kerr and Grant (2000), Brenden et al. (2004)

Shorthead Redhorse

Bozek et al. (1999), Casselman (2011)

Silver Shiner

Brenden et al. (2004)

Smallmouth Bass

Hourston (1952), Gammon and Hasler (1965)

Spotfin Shiner

Wahl and Stein (1991)

Spottail Shiner

Brenden et al. (2004)

Suckers (Catastomus spp.) Anderson (1948), Hourston (1952), MacKay (1963), Brewer (1969), Oehcke (1969), MacCrimmon and Skobe (1970), Scott and Crossman (1973), Deutsch (1986), Seelbach (1988), Cook and Solomon (1987), Brenden et al. (2004), Brautigan and Lucas (2008), Wisconsin DNR (2008), Simonson (2012), Woomer et al. (2012), Brittle (2014)

Sunfishes (Lepomis spp.)

Scott and Crossman (1973), Wahl and Stein (1991), Brautigan and Lucas (2008), Brittle (2014)

Telescope Shiner Brenden et al. (2004)

Tessellated Darter

Kapuscinski et al. (2012)

Fish Species Reference

Trout-Perch

Bozek et al. (1999)

Walleye

Anderson (1948), Hourston (1952), Bozek et al. (1999), Wolter et al. (2012)

White Bass

Wolter et al. (2012)

White Crappie Wahl and Stein (1991)

White Perch

Stocek et al. (1999)

White Sucker

Hourston (1952), Crossman (1962), Bozek et al. (1999), Stocek et al. (1999), Kerr and Grant (2000), Williams (undated)

Yellow Bullhead

Kerr and Grant (2000), Wolter et al. (2012)

Yellow Perch Anderson (1948), Hourston (1952), Crossman (1962), MacKay (1963), Gammon and Hasler (1965), Schmitz, W. R. and Hatfield (1965), MacCrimmon and Skobe (1970), Scott and Crossman (1973), Seelbach (1988), Bozek et al. (1999), Kerr and Grant (2000), Brautigan and Lucas 2008), Wisconsin DNR (2008), Simonson (2012), Williams (undated)