The E.ffects of Hydroelectric Develdprne,nt on .. the Biology of No.rthern Fishes (Reproduction and . · Population·Dynarnics} ln. Yellow Walleye · Stizostedion vitreum vitreum (Mit chill).
A L.iterafl.lre Review and Bibliography.
by Kazimierz Machniak
FISHERIES AND MARINE SERVICE SERVICE DES PECHES ET DES SCIENCES DE LA MER
1975 £ Environment Environnement "l!ij'F Canada Canada
Fisheries Service des peches and Marine et des sciences Service de Ia mer Technical Reports
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Fisheries and Marine Service Service des Peches et des Sciences de la mer
Research and Development Directorate Direction de la Recherche et Developpement
TECHNICAL REPORT NO. 529 RAPPORT TECHNIQUE N°. 529
(Numbers 1-456 in this series were issued (Les numeros 1-456 dans cette serie furent as Technical Reports of the Fisheries utilises comme Rapports Techniques de l'Office Research Board of Canada. The Series des recherches sur les pecheries du Canada. name was changed with report number 457) Le nom de la serie fut change avec le rapport numero 457)
THE EFFECTS OF HYDROELECTRIC DEVELOPMENT ON THE
BIOLOGY OF NORTHERN ~ISHES (REPRODUCTION AND POPl~ATION DYNAMICS)
III. YELLOW WALLEYE STIZOSTEDION VITREUM VITREUM (MIT CHILL) .
A LITERATURE REVIEW AND BIBLIOGRAPHY
by
KAZIMIERZ MACHNIAK
This is the sixtieth Ceci est le soixantieme
Technical Report from the Rapport Technique de la Direction de la
Research and Development Directorate Recherche et Developpement
Freshwater Institute Institut des eaux douces
Winnipeg, Manitoba Winnipeg, Manitoba
1975 i
Machniak, Kazimierz. 1975. The Effects of Hydroelectric Development on the Biology of Northern Fishes (Reproduction and Population Dynamics) III. Yellow Walleye Stizostedion vitreum vitreum (Mitchill). A Literature Review and Bibliography. Fish. Mar. Serv. Res. Dev. Tech. Rep. 529, 68 pp.
The reproduction and early life of the yellow walleye, Stizostedion vitreum vitreum, is reviewed. Walleye commonly spawn in riffles of streams or along shorelines of lakes. Although they have been reported to spawn on a wide variety of substrata it appears they prefer clean gravel bottoms at depths less than 1.5 metres. In impoundments, walleye are apparently less influenced by water levels during the spawning period than are other "shallow water" spawners. Nonetheless, fairly stable or slightly rising levels during spawning and incubation are recommended if spawning is to be successful. Among the many hazards to reproduction, the silting over of spawning beds due to erosion along lakeshores and flooded streams is probably the major cause of spawning failure in impoundments. Growth and numbers could increase, but will be dependent upon the availability of forage species and spawning habitat.
Cet ouvrage etudie la reproduction et le debut de la vie du dare jaune, Stizostedion vitreum vitreum. Le dare jaune fraye generalement dans les bas-fonds des rivieres ou pres du bard des lacs. Bien que les etudes montrent qu'il fraye a des profondeurs tres variables, il prefere fonds de gravier propres a des profondeurs inferieures a 1.5 metre. Dans les frayeres artificielles, le niveau de l'eau semble mains influencer le dare jaune pendant la saison du frai que les autres especes qui frayent dans les "hauts fonds." Neanmoins, pour que le frai soit bon, il est preferable que le niveau de l'eau soit assez stable ou monte legerement pendant le frai et la periode d'incubation. Parmi les nombreux facteurs qui peuvent gener la reproduction dans les frayeres artificielles, la principale est probablement ie recouvrement des frayeres par suite de !'erosion qui se produit le long des rives des lacs et des rivieres en crue. La croissance et la multiplication du dare jaune depend de la quantite des poissons dont il se nourrit et du nombre d'endroits qui peuvent servir de frayeres. ii
TABLE OF CONTENTS
ABSTRACT i TABLE OF CONTENTS ii LIST OF TABLES iv ACKNOWLEDGEMENTS iv INTRODUCTION 1 REPRODUCTION AND EARLY LIFE HISTORY OF THE YELLOW WALLEYE Stizostedion vit~eum vit~eum (Mitchill) Place of spawning 2 Time of spawning 2 Spawning grounds 2 Sexual maturity 5 Fecundity 8 Sex ratios 8 Sexual dimorphism 11 Migration 11 Homing 11 Spawning behaviour 12 DEVELOPMENT OF EGGS AND.LARVAE Description 13 Fertilization 14 Incubation period and hatching 14 Early life 15 Mortality of eggs and larvae 16 FACTORS AFFECTING REPRODUCTION Temperature 16 Water levels and discharge 17 Quality of spawning sites 17 Water quality 18 Siltation 18 Pollution 18 Predation 19 Summary 19 EFFECTS OF IMPOUNDMENT ON WALLEYE REPRODUCTION Sexual maturity 20 Fecundity 20 Migration 20 Water temperature 21 Egg properties 21 Spawning grounds 21 Water level regulation and discharge rate 22 Siltation 24 Predation 24 Summary 24 WALLEYE POPULATIONS AND IMPOUNDMENT Population size 25 Food supply 26 Growth 27 iii
Water level regulation 29 Distribution 29 Summary 30 BIBLIOGRAPHY OF YELLOW WALLEYE - REPRODUCTION AND EARLY LIFE 31 BIBLIOGRAPHY OF YELLOW WALLEYE - IMPOUNDMENT LITERATURE 50 ADDITIONAL CITATIONS 68 iv
LIST OF TABLES
TABLE
1. Time and Temperature of Walleye Spawning 3
2. Depth and Bottom Types of Walleye Spawning 6
3. Fecundity of Walleye in Various Localities 9
ACKNOWLEDGEMENTS
Numerous persons have assisted me in providing information for this report. Their assistance is gratefully acknowledged. In particular, I wish to thank Dr. A. L. Hamilton, Dr. R. E. Reeky, and H. A. Ayles who have critically reviewed this manuscript. 1
INTRODUCTION
This is the third in a series of technical reports concerning the impact of hydroelectric development on the biology of certain fishes. The report summarizes the available information on the repro duction and success of yellow walleye populations in reservoirs. Spawning in natural waters is also emphasized in assessing environmental requirements. 2
REPRODUCTION AND EARLY LIFE
HISTORY OF THE YELLOW WALLEYE,
STIZOSTEDION VITREUM VITREUM (MITCHILL)
The walleye has a very extensive distribution, occurring as far north as the Mackenzie Delta and south down to Texas. However, most of the published materials dealing with reservoir impoundment originate from southern regions where the majority of the walleye populations are introductions and/or lake spawners. Nonetheless, in Canada, the basic biology of this species has been well studied, although there exists little data in relation to hydroelectric develop ment and reproduction.
Place of Spawning
Walleye commonly spawn in the rapids of streams, shallow offshore reefs, or along shorelines of lakes. Some populations are also known to spawn in marshes (Priegel, 1970).
Time of Spawning
Spawning occurs in the spring or early summer, depending on latitude and water temperature. Spawning throughout the walleye range may begin in early March or as late as June (see Table 1). Normally, walleye begin spawning at about the time of ice break-up in a lake, at water temperatures of 6.7 to 8.90 C, but have been known to spawn over a temperature range of 5.6 to 11.10 C (Scott and Crossman, 1973).
Spawning Grounds
The types of spawning areas utilized by walleye have been described by numerous .workers, among them; Bean (1903), Dymond (1926), Derback (194 7), Eschmeyer (1950), Rawson (1957), Niemuth et al. (1959), Johnson (1961), Ellis and Giles (1965), Regier et al. (1969) and Priegel (1970).
Walleye have been reported to spawn on a wide variety of substrata, although it appears that walleye select gravel bottom for spawning when it is available (see Table 2). For example, Eschmeyer Table 1. Time and temperature of walleye spawning.
Location Time of Temperature at Authority Spawning Spawning (oC)
Alberta May Bidgood (1971)
Saskatchewan April 30-May 18 3. 3-11.1 Rawson (1957) (Lac la Range) *(7.7-10.0)
Manitoba May 6.1 Derback (1947)
April-May 6.7-13.0 Ellis and Giles (1965)
Ontario w (L. Erie) Mid-April 7.2. Parsons (1972)
(L. of the Woods) Late April-early 3.9-7.2 Macins (1972) June
(Talbot R.) April 7.2-15.6 MacCrimmon and ~kobe (1970)
Northwest Territories June 5-June 11 Jessop et aZ. (1973) (Rabbitskin R.)
Canada Early April-end June 5.6-11.1 Scott and Crossman (1973)
Iowa 7.2-10.0 Harlan and Speaker (1969)
Michigan May 4.4-6.7 Eschmeyer (1950) *(7.8-8.9) Table 1. (Cont'd)
Location Time of Temperature at Authority Spawning Spawning (OC)
New York (Oneida L.) March-April Raney and Lachner (1942) Forney (1966) April Houde (1969)
Minnesota
(L. Winnibigoshish) April-May 5.0-17.8 Johnson (1961)
Wisconsin Mid-April- 6.1-17.2 Niemuth et al. (1959) early May *(8.9-10.0) ~
(Spoehr' s Marsh April 3-April 25 2.2-12.2 Priegel (1970) Wolf R.)
(Fox R.) March 31-April 28 3.3-15.6 Priegel (1970)
(L. Winnebago) April 17-May 4 3.9-11.1 Priegel (1970)
Reservoirs
Oklahoma March 10-March 20 3.9-14.4 Grinstead. (1971) (Canton Reservoir) March 4.4-15.6 Gennings (1967)
Tennessee-Kentucky (Dale Hollow Reservoir) March 15-April 28 5.6-14.4 Libbey (MS 1969) . *(10.0)
* Optimum Temperature 5
(1950) observed that sand was avoided while gravel areas only a few feet in diameter were utilized for spawning.
He also noticed that unused areas generally had steeper depth gradients and were less often wave-washed. Johnson (1961) made similar observations. Walleye eggs were found frequently on small isolated patches of firm gravel and rubble along extensive shorelines of pure sand where there was little or no spawning. Spawning also takes place on less favourable bottom types when gravel is not present. Smith (1892), Fish (1932), Priegel (1970), Bidgood (1971), and others have noted walleye spawning on sand bottoms in lakes and rivers. Some populations even spawn over vegetation in flooded areas (Nevin, 1900; Niemuth et al., 1959; Priegel, 1963; 1970; and Schumann, 1964). Suspected spawning and nursery areas of yellow walleye, whitefish, and northern pike in the Mackenzie River System were summarize-d by Stein et al. (1973 a, b) and Jessop et al. (1973). However, all walleye spawning sites have in common a supply of moving water for aeration and clean substrate on which to deposit the eggs.
Walleye spawn in relative shallow water (see Table 2). According to various observers '(Breder and Rosen, 1966), "the spawning of walleye occurs in water from 0. 9 to 3 m in depth." Derksen (MS 1967), however, stated that spawning occurs generally in water less than one metre deep. Similarly, Niemuth et al. (1959) reported that eggs are mostly deposited in water less than 0.6 m deep and no more than 1.2 m deep. Johnson (1961) found walleye eggs spawned on gravel in water as shallow as 5.1 em, with the deepest water from which walleye eggs were collected being approximately 1.2 m. Ellis and Giles (1965) also observed walleye spawning in a stream-compound in water less than 0.6 m deep.
Ellis and Giles (MS 1964) state "that of those species whose spawning has been investigated from the ethological point of view, the whitefish Coregonus lavar?tus (Fabricius and Lindroth, 1954) most closely resembles the walleye." The published literature indicates that walleye and whitefish (e.g., "Storsik", Fabricius, 1950) are both species which can spawn in a variety of shallow-water habitats, including lake shallows and rivers (even marshes for walleye). In fact, Bidgood (1972) noted that Buck Lake (Alberta) walleyes were documented as spawning on the same surficial lake sediments of boulders, sand and gravel, as lake whitefish utilize for spawning. The walleye-whitefish pattern appears to be adapted for spawning in a variety of habitats and hence walleye are less specialized and less restricted in their spawning.
Sexual Maturity
Accor~ing to Niemuth et al. (1959), males from most populations become mature when two to three years old and 30.5 to 33.3 em long. Females mature when four to five years old, and at lengths of 38.1 to 43.2 em. Also, Scott and Crossman (1973) state that male walleyes generally mature at 2 to 4 years of age, over 11 in. (27.9 em) in length, Table 2. Depth and bottom types of walleye spawning.
Location Depth of Substrate Authority Spa'>ming (metres)
Alberta (Maybelle R.) Shallow water Sand Bidgood (1971)
Manitoba (Northwest Creek) Less than 0.9 Hard stony bottom Derback (1947)
Less than 0,6 Gravel Ellis and Giles (1965)
Less than 1~0 Gravel Derksen (MS 1967)
0\ Ontario Shallow water Bouldery riffles, sand, MacKay (1963) gravelly or stony shoals in lakes
(Lake of the Woods) Gravel Evermann and Latimer (1910)
(Georgian Bay) Sticks and stones in running water. Bensley (1915)
(L. Erie) Sandy bars Fish (1932)
(Talbot R.) 0.2 to over 0.9 Coarse gravel and rubble MacCrimmon and Skobe (1970)
Canada Shallow water Rocky areas in rapids Scott and Crossman (1973) (rivers) boulder, coarse gravel shoals in lakes Table 2. (Cant' d)
Location Depth of Substrate Authority Spawning (metres)
Michigan Shallow water Gravel, rubble, boulders Eschmeyer (1950) and sand
Minnesota Broken rock Cobb (1923)
(L. Winnibigoshish) Less than 1.2 Gravel-rubble, sand Johnson (1961) and muck
Wisconsin 0.6-1.2 Gravel Niemuth et al. (1959) '-J (L. Winnebago region) Less than 0.6 Marsh vegetation and Priegel (1970) gravel
New York Shallow water Sandy bars Bean (1903)
Gravel Kingsbury (1948)
Iowa 0.3-1.5 Rock reefs, sand bars Harlan and Speaker (1969) or gravel
Oklahoma (Canton Reservoir) Less than 1. 5 Rip-rap Grinstead (1971) Gennings (1967)
South Dakota (Lake Frances Case Less than 1. 2 Gravel-rubble-boulder North Central Reservoir (reservoir) ) Investigation 1968 (1969) 8 and females at 3 to 6 years of age or 14 to 17 in. in length (35.6- 43.2 em).
Rawson (1957) reported that the dominant age group for spawning walleyes in Lac la Range, Saskatchewan, were age groups VIII to X, and that few walleye of age group V were present on the spawning run. In Wisconsin, Priegel (1970) noted that the dominant age groups in the spawning populations for males and females were VI through VII. The length distributions of spawning run walleye have been reported by several authors and these show considerable variation (Rawson, 1957; Smith and Pycha, 1960; Priegel, 1969a; Johnson, 1971; and others).
Wolfert (1969) reported a decrease in age at maturity of walleye since 1927-28 in Lake Erie accompanied by a large increase in growth rate. Here some females are mature at age III and all were mature by age V, while almost all age II males are mature at present growth rates.
Fecundity
"A few estimates have been published on the egg production of the walleye, but most of these estimates have been based on a small number of fish and the size range has been limited" (Priegel, 1970). Nevertheless, walleye have a high fecundity; various studies have shown between 30,000 and 600,000 or more eggs per female. Data on fecundity in various waters are given in Table 3. Hood (1969) reported egg size varied with the size of the female. A female of 38.1 em had eggs with a diameter of 1.83 mm while a female of 71.1 em had eggs of 2.29 mm. However, Wolfert (1969) found no relation between egg diameter and length or age of walleye in Lake Erie.
Sex Ratios
Males usually predominate throughout the spawning run. They arrive first on the spawning ground and remain longer than the females (Rawson, 1957; Eschmeyer, 1950; Niemuth et al.~ 1959; Libbey, MS 1969; Harlan and Speaker, 1969; Priegel, 1970; Scott and Crossman, 1973; and others). The male to female ratio can range from a low of 0.8:1 to a high of 14:1 (Johnson, 1971). During the late stage of the run, females usually outnumber males.
However, accor~ing to Priegel (1970), "sex rattos are difficult to obtain for 3 reasons: (1) On the one hand, males arrive on spawning grounds before the females do and they remain th~re throughout most of the spawning season. On the other hand, females move on to the spawning area, spawn, then leave immediately. On the basis of different spawning behavior for the two sexes, there will always be more males than females on natural spawning areas. (2) Males also begin, to reach sexual maturity at the end of their third year of life, and are completely mature by the Table 3. Fecundity of walleye in various localities.
Location Range/kg Av. no. /kg Authority
Ontario (Lake of the Woods) 50,054 Carlander (1945)
(Lake Erie) 99,225 Leach (1927)
((Eastern Basin)) 41,284-97,135 61,288
((Western Basin)) 56,314-123,249 82,703 Wolfert (1969)
Minnesota (Little Cut Foot 65,383 Johnson (1971) Sioux Lake) 1.0
Michigan (L. Gogebic) 58,053-67,952
(Muskegon R.) 65,927-96,173 Eschmeyer (1950)
(Saginaw Bay) 91,904
Iowa 50,715-110,250 Harlan & Speaker (1969)
Pennsylvania 55,125-88,200 Hood (1969)
Wisconsin 28,665-99,225 Niemuth et aZ. (1959)
(L. Winnebago) 48,089-63,586 Priegel (1969a) Table 3. (Cont'd)
Location Range/kg Av. no./kg Authority
Reservoirs
Tennessee (Norris Reservoir) 28,478-32,802 29,768 Smith (1941)
(Central Hill More than 63,945 Muench (MS 1966) Reservoir)
(Dale Hollow 26,460-50,715 45,571 Libbey (MS 1969) Reservoir)
1-' 0 11 end of their sixth year. Females, however, begin to reach maturity at the end of their fourth year of life and all are mature at the end of their eighthyear (Priegel, 1969a). Because they mature earlier than females, more male walleyes will be found on the spawning run. (3) Most spawning areas are too large to permit a statistically large enough sample of spawning walleyes to be captured." It appears that sex ratios can be quite variable for the same run over a period of years and also that sex ratio depends upon what stage of the run is monitored.
Sexual Dimorphism
Females are larger in length and weight than males of the same age. "The divergence in growth rates between the sexes occurs before the fish mature for the first time and is likely a secondary sex characteristic under genetic and/or hormonal control" (Bidgood, 1971). According to Adams and Hankinson (1928) and McPhail and Lindsey (1970), the lower lobe of the male caudal fin is distinctly striped with white as are the anterior borders of the paired fins. Ellis and Giles (MS 1964), however, noticed no secondary sexual differences amongst colour patterns in walleye and the only distinguishing feature between the sexes was the distended abdomen of ripe females.
Migration
Water temperature appears to be the controlling influence for the spawning run. Rawson (1957) found that walleye runs from Lac la Range, Saskatchewan, occurred long before the break-up of ice on the lake and postulated that the warmer river water stimulated the spawning migration of these fish. Little movement took place at 4.4 to 5.6°C but most active spawning was observed between 7.7 to 10°C. He also demonstrated that in cold years with late lake ice breakup the spawning run was protracted to 3 or 4 weeks. Grinstead's (1971) data also seems to support the theory that water temperature is the primary controlling influence in determining the time walleye spawn. He showed that in Canton Reservoir, Oklahoma, they spawn considerably earlier (March 10 - March 20), but within temperature ranges similar to walleye in other regions.
Homing
Walleyes exhibit a homing behaviour, returning annually to particular spawning sites, rather than seeking any suitable spawning area. The tendency of walleye to return to specific stream spawning grounds has been noted by many authors. Stoudt (1939), Stoudt and Eddy (1939), Eddy and Surber (1947), Eschmeyer (1950), Smith et al. (1952), Eschmeyer and Crowe (1955), Rawson (1957), Forney (1961, 1963), 12
Priegel (1970), and others; all observed that stream spawning walleye tagged on specific spawning grounds tended to return to .them. A similar phenomenon has been observed in lake spawning populations (Forney, 1971; Olson and Scidmore, 1962; Crowe, 1962; Crowe et aZ. 1963; Payne, 1964; Anonymous, 1966; Johnson and Johnson, 1971; and others). "As a consequence of the tendency to return to chosen spawning sites, stocks of walleye are probably composed of discrete sub-populations identified with particular spawning areas" (Crowe, 1962). If, however, spawning grounds are within a few miles of each other, then extensive intermingling of lake-spawning populations of walleyes occurs as demonstrated by Whitney (1958) in Clear Lake, Iowa.
Stoudt and Eddy (1939) and Smith et aZ. (1952) suggested that the homing instinct might be weak. Olson and Scidmore (1962) demonstrated that the tendency of walleyes to return to a particular spawning site is not evidenced to the same degree by all fish. Their results show a greater return of fish which had a previous history of appearing in the run during years of low flow. This suggested that these fish were either better able to locate the areas when flow was reduced or were more strongly motivated to enter the stream under the less favorable conditions. The mechanism of homing in walleyes is as yet unknown.
Spawning Behaviour
The spawning behaviour of walleye has been described by a number of investigators (Bean, 1903; Adams and Hankinson, 1928; Eschmeyer, 1950; Schumann, 1964; Ellis and Giles, 1965; Baker and Manz, 1967; Priegel, 1970; and others). The literature on walleye suggests that it is not a territorial fish at spawning time (Eschmeyer, 1950), but that some slight form of courtship behavior occurs among grouped fish at night over shallow spawning grounds. In addition, the rather drab coloration and lack of specific color patterns which might serve as social releasers (Baerends, 1957) support the implication in the literature that a complex courtship ritual does not take place.
Walleye are known to be essentially nocturnal spawners, although Harlan and Speaker (1969), MacCrimmon and Skobe (1970), and Priegel (1970) observed fish spawning in broad daylight. Ellis and Giles (1965) found that spawning occurred most frequently in the early evening.
The females are usually larger and spawn with one or more males. "The fact that a number of males escort one female may be a behavioral adaptation to compensate for a relatively low likelihood of fertilization due to a short-lived viability of either the egg or sperm" (Regier et aZ. 3 1969). Ellis and Giles (1965) described the spawning act as follows: "Overt courtship began by either males or females approaching another of either sex from behind or laterally and pushing sideways against it, or drifting back and circling around, pushing the approached fish backwards. The first dorsal fin was alternately erected and flattened during these approaches. The approached fish would either hold position or withdraw. 13
Approaches and contact of this sort appeared to be the preliminary essentials of courtship and were promiscuous, i.e., there was not continued relationship between any particular pair of fish. Activity increased in frequency and intensity and individuals began to make preliminary darts forward and upward. Finally, one or more females and one or more males came closely together and the compact group rushed upwards. At the surface the group swam vigorously around the compound unt'il the moment of orgasm.when swimming stopped and the females frequently turned or were pushed violently onto their sides. This sideways movement by the females was taken as an indicator of spawning even when no eggs or milt were seen. In summary, based on clear sightings of simultaneously released eggs and milt, spawning consisted of a series of synchronized acts by promiscuous groups of fish. Each act was preceded by simple short courtship consisting of approaches and bodily contacts between individuals. There was no indication of terri torial defense, even though some fish would maintain position for hours on end. Females can spawn out completely in one night, whereas males have the potential for spawning over a longer period."
Walleye exhibit a diel behavioral cycle on natural spawning grounds. The cycle consits of low activity in daytime, expressed mainly by position-holding. According to Derback (1947), walleye rest during daylight hours in protected bays or behind rocks in rapids. Eschmeyer (1950) noted that walleyes are very sensitive to light. However, Priegel (1970) found no indication that they left the spawning area during the day to retreat to deeper water. During the evening, walleye show increased activity, expressed by courtship.
Walleye spawn annually after reaching sexual maturity, although there is some indication that they may miss spawning seasons (Carlander, 1945; Deason, 1933). Also, during the spawning season walleye feed little (Parsons, 1953; Muench, MS 1966; Libbey, MS 1969).
Development of Eggs and Larvae
Description
The mature eggs are spherical in shape, with a large yolk enclosing an oil drop which has a diameter more than a third that of the egg itself. The eggs are relatively transparent with a yellow-brownish colouration (Reighard, 1890).
Walleye eggs are adhesive for some hours after spawning and then lose their adhesiveness (Reighard, 1890; Leach, 1927; Nelson et al.~ 1965). Egg diameter is 1.5- 2.0 mm (Scott and Crossman, 1973). "Walleye eggs are hyaline and turgid early in development but often are flaccid during the eyed stage, especially just before hatching" (Johnson, 1961). 14
Fertilization
Hatchery workers have found that the range of fertilization among walleye eggs is very wide. Klienert and Degurse (1968) noted a range of 20.1 to 95.3 percent and an average of 72.1 percent. Hurley (1972) found that the percentage of fertilized eggs ranged from 65.0 to 97.5 percent, averaging 85.3 percent.
The percentage fertilization of eggs deposited naturally is also quite variable. Baker and Manz (1967) found that over a 7-year period between 19 and 49 percent of large numbers of eggs taken from reefs in western Lake Erie had live, developing embryos. Johnson (1961) noted that shortly after spawning, between 28.3 and 90.8 percent of the eggs were alive and presumably represented fertilized eggs. However, by late incubation the percentage of live eggs ranged from 4.7 to 51~9 percent on the various spawning grounds. He also noted, that since egg fertility was consistently high in the areas studied, it seems unlikely that inadequate fertilization was a significant factor in egg survival.
The "group" spawning behavior of walleye probably facilitates the initial high fertility of walleye eggs. Stranahan (1900) stated that walleye milt dies in two minutes and eggs cannot be impregnated after 6 minutes exposure to water. "If this is so, simultaneous spawning by a larger number of walleye over a small area should increase the probability of fertilization" (Regier et al.~ 1969).
Incubation Period and Hatching
The incubation period usually runs from 12 to 18 days, depending upon the water temperature. Niemuth et al. (1959) stated that the eggs hatch in 26 days when the water temperature is 40°F (4.4°C), 21 days when 50 to 58°F (10-14.4°C) and 7 days at a mean temperature of 57°F (13.9°C).
Temperatures on the spawning grounds can be quite variable. Priegel (1970) found water temperatures during egg development ranged between 36 to 66 0 F (2.2-18.9 0 C). Johnson (1961) reported "that most rapid embryo develop- ment and short incubation periods were associated with high daytime water temperatures and high minimum water temperatures." He also found once that survival was highest in the year with the shorter incubation period.
Other factors beside water temperature can influence the incubation period. Oseid and Smith (1971) demonstrated that eggs hatched under lower oxygen levels required one to four days longer incubation. According to Priegel (1970), "four factors influenced egg development; (1) Water levels and flow, (2) Substrate type, (3) Predator activity and (4) Dissolved oxygen." 15
Early Life
Reighard (1890), Fish (1932), Norden (1961), Olson (1966), Nelson (1968), and others, gave details on the early development of the young. Fry, upon hatching, will absorb their yolk sacs within a 3 to 5 day period (Priegel, 1970). Feeding takes plac~ prior to disappearance of yolk, in fact, Hurley (1972), noted cannibalism near the end of yolk absorption. According to Priegel (1970), newly-hatched larvae have no developed paired fins and the movement is restricted to vertical swimming. Only after a 10 to 12 day period do their paired fins develop sufficiently to allow horizontal movements.
Newly-hatched fry range in total length from 6.0 to 8.6 mm. Small fry have been observed in schools at the spawning grounds, but they soon disperse. Cheney (1897) believed that after hatching, the brood remains together for the first season if not destroyed, making a solid compact mass during the first two weeks. Bajkov (1930) said that the fry usually school in comparatively shallow places. Houde (1967) showed that walleye larvae lead a pelagic existence for their first 4 to 6 weeks and were concentrated near the surface in protected bays. However, the movements of walleye immediately after hatching and for a period thereafter are not well known.
Smith and Moyle (1943), studying rearing ponds, reported that walleye fry began feeding on rotifers and nauplii and that as the fish increased in size, entomostraca, insects and fish successively became important items in the diet. Hahn (1966) found that diatoms were the first food of pelagic fry in western Lake Erie. On the other hand, Houde (1967) found that large zooplankters, particularly copepods, were the initial food source of walleye in Oneida Lake. Priegel (1970) also found that walleye in the 10-50 mm size class feed principally on copepods and cladocerans. "Apparently some food selectivity occurs during this stage of development" (Houde, 1967). In fact, Priegel (1970) reported that "Leptodora which was the least abundanct zooplankton in Lake Winnebago, was positively sought and selected by young walleye. The preference shown by young walleye for Leptodora was probably due to its relatively large size which made it a more attractive food item than other species of zooplankton."
Dobie (1966) reported that when young walleye had reached a length of about 30 mm, they shifted to feeding on fish (yellow perch). Maloney and Johnson (1955) showed that most of the stomach contents of young walleye from Mille Lacs Lake was composed of fish (98.9 percent by volume), with young yellow perch accounting for 77 percent of this volume. Priegel (1970), however, found that "trout-perch and freshwater drum fry were the most important species of forage fish utilized" in the Lake Winnebago region by young walleye. In years when species of forage fish were absent, consumption of chironomids, copepods and cladocerans increased. According to Forney (1966), growth rate of walleye in Oneida Lake in late summer tended to be rapid in years when walleye fed on fish and slower when invertebrates were common in the diet. 16
Mortality of Eggs and Larvae
"Weather conditions may be the critical factor affecting survival at this early life-stage" (Regier et aZ.~ 1969). In a natural situation, fluctuating water temperatures, washing ashore, siltation, and other factors cause a high mortality of eggs with only 5 to 20 percent survival to hatching. Even then, the odds of survival from hatching through the first year are about one in ten thousand (Shields, 1965).
Factors Affectjn g Beproducti on
Temperature
Spawning success could depend on favorable water temperatures during the spawning period of the walleye. Payne (MS 1964) found some evidence that stronger year classes of walleyes tend to arise during warmer than average spawning seasons. Furthermore, Tait (1973) stated that the density of eggs on the spawning grounds in Lake Erie and the calculated average daily temperature rise are strongly correlated (r = 0.81). Strong year classes of walleye developed in years when water temperatures rose steadily and rapidly during the spawning and incubation periods, and thus reduced the period of exposure of the walleye eggs to predation, low o~gen tensions, disease, sedimentation and wave action.
Unfavorable temperatures, on the other hand, have been known to disrupt spawning. Fluctuating temperatures would tend to disrupt spawning in the sense that the fish might spawn intermittently over a period of weeks instead of over a much shorter interval if temperatures rose steadily. Derback (1947) noted that walleyes were spawning normally in a stream in northern Manitoba at a'water temperature of 43°F (6.1°C), when a sudden and prolonged cold spell caused a drop in tempera ture resulting in the spawners leaving the stream and not returning for the season. Walleye taken later in June were resorbing their eggs. Similarly, Schumann (1964) reported that during a cold spring, following a warm initial burst, many walleye failed to spawn and many of the eggs spawned late in the season were sterile, apparently because they were physiologically too old. Although cold weather prolonged spawning activity on Spoehr's Marsh, Wisconsin, it never inhibited it over an extended period of time (Priegel, 1970).
The fact that the walleye is predominately a river-spawning species tends to indicate that the eggs and fry are tolerant of rapid temperature fluctuations. Indeed, Allbaugh and Manz (1964) demonstrated that walleye eggs were fairly tolerant of water temperature fluctuations. However, Johnson (1961) stated that "egg mortality, especially as associated with unusually cold water during the egg incubation period, may be an important factor in the establishment of year classes." He 17 noted that survival was best in years of warmer and shorter incubation periods. Doan (1942) and Carlander (1945) observed no relation between spring air temperature and subsequent size of walleye age classes. "Whether temperature is, in fact, the immediate factor, or whether it is some combination of correlated factors is not known" (Regier et al.,, 1969).
Water Levels and Discharge
Carlander (1945) in the Lake of the Woods and Tait (1973) in Lake Erie found no correlation between spawning success and water levels. Johnson (1961) and Priegel (1970) felt that fluctuations of water levels during the spawning period were not an important factor. Lake-spawning walleye generally spawn in depths of water great enough that subsequent changes in water level appear to have little effect. However, where spawning sites are in shallow water sudden drops in water level may have an adverse effect. According to Priegel (1970), "on Spoehr's Marsh and all other marshes along the Wolf and Fox rivers, water level on the marshes are a major factor in spawning success." For example, in 1963, no eggs hatched when the marsh dried up completely after all spawning had occurred. Derksen (MS 1967) reported that variations in year class abundance of walleye in Cedar and Moose Lakes (Manitoba) were positively correlated with variations in discharges of the Saskatchewan River. A walleye investigation in Minnesota showed more fish in a spawning run in years of intermediate to large stream flow, and fewest when stream flow was low (Olson and Scidmore, 1962). High discharges can also increase the success of spawning by providing optimum conditions during incubation or promoting first-year growth.
Quality of Spawning Sites
Areas with clean bottom types and .a good flow of water seem to provide the best spawning sites for walleye reproduction. Johnson (1961) examined egg survival on five bottom types utilized by walleye; soft muck, sand, gravel, rubble and boulders. He found that egg survival was poorest on the soft muck - detritus bottom, intermediate on firm, clean sand bottom and was best on clean gravel - rubble bottom. "Walleye eggs deposited on the soft mud are probably experiencing a high mortality rate as a result of insufficient circulation of water through the mud and a subsequent lack of available oxygen" (Dickson, MS 1966). The superiority of gravel-rubble bottom over sand was aptly demonstrated by the artificial addition of such a substrate - resulting in percentage survival of eggs increasing 5 times, and fry production increasing more than 100 times. "It appears that eggs deposited on clean, firm gravel-rubble substrate are less subject to entanglement in debris and scouring from waves compared to those on other bottom types" (Johnson, 1961). 18
Water Quality
Colby and Smith (1967) noted low survival of walleye eggs in field situations with low oxygen levels. Priegel (1970) reported that "dissolved oxygen was an important factor in spawning success over most of the Fox River marshes. Low dissolved oxygen concentrations were directly related to low water levels, dense vegetation, carp activity and excessive algae growth." Van Horn and Balch (1956) and Oseid and Smith (1971) tested the survival of walleye eggs at various oxygen levels. They found that at lower levels of oxygen, hatching time is substantially extended and mean length of fry is smaller. "If shorter incubation periods and larger size at hatching time are considered to be advantageous to placing larger year classes in a natural system, optimum oxygen levels for the incubation of walleye eggs appear to be not lower than 5 to 6 p. p.m." (Oseid and Smith, 1971) •
Stein et al. (1973b) noted differences in water quality between spawning areas of different species. Yellow walleye, Arctic grayling, Arctic char and burbot spawning and nursery areas were predominant in clear streams. Spawning and nursery areas of whitefish, cisco, longnose sucker and northern pike were found in waters that ranged from clear to turbid.
Siltation
Sedimentation as a result of erosion or decreased water flows can cause egg mortality during the incubation period. Hood (1969) noted that experiments have been conducted in hatcheries on the effect of siltation on eggs in hatching jars. In the experimental units in which siltation was allowed to occur, all eggs were lost; in the control units where preventive measures were taken, an 82 percent hatch was obtained. Whether this is a serious factor affecting hatching success in nature is unknown, although from the walleye's preference of spawning sites it must be deduced that silt deposition would be deleterious to egg survival as it affects oxygen availability.
Pollution
Pollution in some streams or lakes may possibly inhibit repro duction or result in a poor hatch success. Some studies have been conducted on the effects of pulp-mill effluents on survival of walleye eggs and larvae (Longtin, 1953; Smith and Kramer, 1963; Smith et al. 1966). Their results show that conifer groundwood at 50 to 150 ppm acts as a loading and limiting stress and reduces the scope for activity of the walleye fingerling. Present studies also indicate that the deleterious effect of suspended wood fibers on walleye egg survival is small, and that mortality in water carrying mill effluents at levels observed in the Rainy River is due primarily to Spaherotilus growths, and, in some lesser degree, to toxic or oxygen-demanding bottom 19
deposits of fiber. Colby and Smith (1967), however, show that sulfide levels of 0.3 ppm approximating river conditions are acutely lethal to walleye eggs and fry.
Predation
Heavy predation on eggs and larvae could be a serious factor in hatching success. Some of the following fishes have been observed consuming walleye eggs; yellow perch, spottail shiner, white sucker, stonecat, and various minnows. "No detailed studies have been reported on the predation of larger fish on walleye fry" (Regier et al.~ 1969), although adult perch, white bass, walleye, pike, alewife, American smelt, and sauger prey on the young as probably do a wide variety of other predatory fishes. The behaviour of the fry after the yolk has been absorbed adds to the problems of survival. Cuff (MS 1973) states that in the early stages, "cannibalistic behaviour occurs regardless of the amount of food present; walleye, not recognizing one another as conspecifics, teat each other as potential food." "Cannibalism is most likely to affect year class size in a population where predator density is high and where there is intense competition for prey" (Chevalier, 1973). Forney (1974) found that intensity of cannibalism was partly a function of young perch density.
Goode (1903) mentioned the destructive inroads of sturgeon, catfish and suckers upon walleye spawning beds. In contradiction Eschmeyer (1950) and Priegel (1970), noted there was no indication of predation on walleye eggs by any fish species on spawning sites, although some fish were associated with walleye during the spawning season. Priegel (1970) noted that the presence of carp was detrimental to walleye eggs. The spawning carp roil up the bottom and muddy the water.
Summary
Walleye commonly spawn in riffles of streams or along shorelines of lakes. It appears that walleyes selct gravel bottoms, generally in water less than 1.5 m deep. Water temperature, discharge (flow) rates, water levels, dissolved oxygen levels, sedimentation and pollution are probably the main factors affecting spawning success. 20
Effects of Impoundment on Walleye Reproduction
Sexual Maturity
Forney (1965) found that faster-growing walleye matured at an earlier age than slower-growing individuals. Grinstead (1971) demon strated that walleye reached sexual maturity at an earlier age in Canton Reservoir than walleye in other localities. Some individuals were found to be sexually mature after only one year of growth. Apparently, sexual maturity in the walleye is primarily a function of size, and the fast growth of Canton Reservoir walleye is a partial explanation for their early development. Similarly, Libbey (MS 1969) found that walleye in Dale Hollow Reservoir, Tennessee, matured at an early age (age I-III).
Fecundity
The fecundity could respond adaptively to a change in the food supply, since growth rate effects the time of' sexual maturity, which in turn affects fecundity. Smith (1941) found that the very fast-growing walleye of Norris Reservoir, Tennessee, produced about 13,500 eggs per pound (29,768 per kg) of fish. This differs considerably with the average per pound of fish of 23,000 to 50,000 (50,715 - 110,250 per kg) eggs produced by walleye in other areas. On the other hand, Muench (MS 1966) found that the rapidly growing walleye in Center Hill Reservoir, Tennessee, produced an average of over 29,000 eggs per pound of fish weight (63,945 per kg).
Migration
Construction of dams can reduce the abundance of walleye by preventing the highly-migratory species from ascending the streams (Trautman, 1957). This problem can be particularly acute if walleye exhibit any homing tendencies. However, the flooding of tributary streams which serve as spawning sites should not seriously affect reproduction provided that discharge rates and sedimentation levels are not altered significantly. Also, the walleye is a far-ranging species and could possibly migrate to the next set of rapids if prevailing conditions are unsuitable. The diversion of river flows, however, might seriously affect walleye reproduction in some circumstances by altering natural river flows which could affect the spawning behaviour and proper incubation of eggs, migration of fry, as well as homing to spawning grounds.
The importance of olfaction in certain salmonids in locating their spawning tributaries and individual spawning grounds is well documented (Hasler, 1966). "If olfaction is also necessary for walleyes 21 to locate their spawning grounds, then any dissolved foreign matter introduced irito the stream below the spawning grounds would result in disorientation of the fish" (Ryder, 1968). High concentrations of total dissolved solids have been known to block the spawning migration of some fishes, for example, the striped bass, Roccus saxatiUs (Radtke and Turner, 1967).
As stated above, walleye can undergo quite extensive movements. Of 505 tagged walleyes captured within about 7 months after transfer from northern Lake Champlain to Chambly Reservoir, in the Richelieu River, Quebec, 20 had travelled distances of 100 to 175 mi (Desrochers, 1953). Similarly, Eschmeyer and Crowe (1955) studied the movements of walleyes transferred to upstream impoundments. They found the walleyes exhibited a "marked proclivity to move downstream past the power dams toward their original habitat."
Water Temperature
According to Johnson (1961), egg mortality, especially as associated with unusually cold water during the egg incubation period, may be an important factor in the establishment of year classes of walleyes. This was illustrated in Lake Francis Case, where a decrease in water temperature appeared to be an important factor in fish embryo or larval mortality for walleye (North Central Reservoir Investigations 1970, 1971). Many eggs were killed when the water temperature dropped 6°C during one 24-hour period in mid-May 1969. Downstream from a reservoir, deep-water discharges from a dam, might delay or even prevent reproduction (Pfitzer, 1967).
Egg Properties
Differences in adhesion of the eggs constitute an adaption that increases the effectiveness of breeding in certain fish. Most (1965) felt that for this reason sauger would do better in reservoirs than its close relative, the walleye. "Apparently, each of these fish lay eggs in moving water at the upper end of the reservoirs. Because sauger eggs are more adhesive and stick to the bottom better than those of walleyes, they hold better in the current, particularly during power releases from upstream dams."
Spawning Grounds
Suitability and availability of spawning sites will determine the success ,of walleye reproduction in impoundments. Generally, reproduction of walleye in impoundments can follow apattern that is the 22 reverse of that observed in northern pike. Initially, flooding of a reservoir results in poor spawning success of walleye due to the lack of suitable spawning substrate and the increased siltation. However, as more coarse materials in a reservoir bottom become exposed, walleye reproduction dramatically improves. Such is the pattern that has been observed in many Missouri River mainstem reservoirs. Erickson (1972) stated that it apparently requires five or more years of wave washing action to produce clean gravel qars in on-stream reservoirs. This probably explains why Jenkins (l970b) found a significant correlation (0.02 level) that with increase in reservoir age, walleye numbers increase.
Erickson (1972) stated that the presence of gravel bars appears to be essential for walleye survival. Although it appears that walleye select gravel bottom for spawning when it is available, spawning can also take place on less favourable bottom types when gravel is not present. However, "egg survival on bottom types other than gravel rubble, both as percentage survival and number of eggs surviving the eyed stage, is quite low" (Johnson, 1961).
Artificial spawning beds have been constructed in a number of impoundments. Generally, these consist of coarse gravel and rubble deposits in shallow waters (Yeager and Weber, 1972). Some have proved unsuccessful, apparently because they were placed too deep (Klingbiel, 1971). According to Grinstead (1971), "the two criteria- rock substrate and wind-swept shoreline - were not sufficient to define totally the requirements for a walleye spawning site. It appears that depth may be important, as Eschmeyer (1950) reported that areas not used as spawning sites usually had a steeper depth gradient." However, orien tation of sites with respect to winds and currents should not be over looked, as walleye commonly spawn in riffles or where wave action keeps the water in motion. Also, the homing instinct that walleyes exhibit for specific sites could be quite important.
Water Level Regulation and Discharge Rate
The manipulation of water levels, particularly during the walleye spawning season, is most important. Walleye spawn in waters less than two metres in depth in most reservoirs, and it has been shown that water level fluctuations have a negative effect on walleye (Jenkins, 1970b). According to Klingbiel (1971), most States attempt to maintain fairly stable or slightly rising levels during spawning and incubation. Also, Erickson (1972) stated that on-stream impoundments which have slow water level fluctuations produce the best walleye populations.
However, fluctuations of water levels during the spawning period may not necessarily be an important factor in the spawning success of some walleye populations. Apparently, walleye spawning succiess in Lake Francis Case (Missouri River reservoir) shows little relationship to water levels since vast gravel-rubble spawning grounds are available at a wide range of depths (North Central Reservoir 23
Investigations 1970, 1971). Benson (pers. comm.) states that walleye abundance in all Missouri River reservoirs is increasing because they are not littoral spawners~ that is, "they spawn at depths below the effects of water level fluctuation" (Benson, 1973b).
Discharge flow rates rather than water level fluctuations may be more important to walleye spawning, particularly in river impoundments. For, as Nilsson (1973) stated, "from a biological point of view the essential feature of river reservoirs is not the water level fluctuation, but the change in water flow as compared with the stabilized river system of falls, rapids, streams and calmer reaches; in reservoirs the velocity of the stream often 'Changes overnight." Actual spawning may be interrupted or ended prematurely by the operation of a control dam. Insufficient flows can also affect walleye spawning runs, spawning runs, spawning itself, egg incubation, post-spawning mortality and fry transport. For example, Bidgood (1971) found that a decrease in daily discharge of water in the Peace River as a result of the operative Peace River Dam lead to lowered water levels in the Athabasca River delta. This, he felt, was "detrimental to the survival of post-spawning walleye and could reduce the size of subsequent spawning population and recruitment to the population." MacCrimmon and Skobe (1970) noted that when water flow was reduced by the control dam on the Talbot River, walleye left the riffle areas. Control structures can also result in poor water quality on walleye spawning grounds as noted by Stone (MS 1963) and Crowe (MS 1969).
The manner in which water levels are regulated in impoundments might have serious consequences, particularly for age 0 walleye survival. Current velocities in re'servoirs are generally related to discharge or flushing rate. Houde (1969) reported that walleye larvae less than 9.5 mm in length could not sustain currents greater than 3.0 em/sec., and 50 percent only of larvae 10 to 16 mm long could maintain their position in currents in excess of 3 to 5 em/sec. Reservoirs having a high flushing rate would therefore be detrimental to larvae survival.
Indeed, Walburg, (1971) has demonstrated that many age 0 fish are lost in the discharge from Lewis and Clark Lake each summer. He found that sauger-walleye 1osses reached a peak of 700,000 on June 10, 1969 and 110,000 on June 3, 1970. Similarly, Armbruster (1962) estimated that a total of 19,102 walleyes were lost during the period from December through April when rapid reductions in water levels were made. The fish lost were chiefly fingerlings, of which 42 percent were dead or would soon die of injuries sustained while escaping from the reservoir. However, in spite of losses of walleye from Berlin Reservoir he found that there has been a rather consistent production - this may be due to the periodic reduction in population pressure.
Benson (1973a), however, found that the lowering of the water level in Lake Frances Case each fall by 11.6 m for hydroelectric production did not have a major effect on survival of age 0 fish. This was probably due to the manner in which the water was withdrawn. In the case of Lewis and Clark Lake, where fish losses were extremely high, the 24
water was drawn from approximately 3 metres below the surface to the reservoir bottom, whereas, in Lake Francis Case, few, if any, small fish were lost because water was drawn from near the reservoir bottom at a depth of 40 metres.
-5iltation
Sedimentation on lakeshore and stream spawning sites could be particularly harmful to embryo survival in newly-flooded impoundments characterized by bank instability. According to Trautman (1957), . construction of dams in Ohio leadsto increased turbidity and silting over of the hard bottoms with soft clayey silts - one of the major factors in the demise of walleye populations.
Predation
Initial population increases of such species as yellow perch and northern pike could threaten the success of,walleye reproduction in some impoundments. However, Scott and Crossman (1973) state that more important in controlling populations are "water temperature, stream flow and wind at spawning time, and other species which spawn over the walleye eggs or roil up the silt."
Summary
Walleye are apparently less influenced by water levels during the spawning period than are other "shallow-water" spawners. "Walleye usually spawn at depths below the effects of water level fluctuation in impoundments (Missouri River ) (Benson, 1973b) and will, therefore, have more consistent reproduction than near-shore spawners such as northern pike." Accessibility to spawning grounds, water temperature, siltation, water level and flow will be prime factors in determining walleye spawning success in impoundments. 25
Walleye Populations and Impoundment
Although the walleye is an extremely important sport fish. It is also harvested commercially in some of the larger waters. "The walleye is probably the most economically valuable species of fish in Canada's inland waters" (Scott and Crossman, 1973). With the establish ment of impoundments on major rivers and the control of lakes, knowledge of factors that produce good year classes of walleye are imperative if management of this species is to be successful.
Walleye have been introduced into numerous reservoirs, particu larly in the United States. Surprising success has been attained in many shallow, sand and silt-bottomed reservoirs in Nebraska, Kansas, Oklahoma and Texas. "Standing crops and catch rates of walleye are, however, lower in most reservoirs than in natural lakes" (Jenkins, 1970a). Unfortunately, very few studies document changes occurring within natural walleye populations following impoundment. Those studies that are available come mainly from research conducted on Missouri and Tennessee River reservoirs. To the author's knowledge, very little information exists on lake impoundment and its impact on natural walleye populations.
Population Size
In some cases where a river has been impounded, walleye popula tions have not adapted well to conditions found in reservoirs and have declined in numbers (Trautman, 1957; Patriarche and Campbell, 1958; Miller and Paetz, 1959; Walburg, 1964; Carter, 1969; SNBB, 1972; and others). The decline in numbers is attributable to a number of factors; (1) unsuitable temperature regime (2) lack of spawning sites (3) increased turbidity and siltation (4) fluctuating water levels (5) increase in predators, etc. In other instances, walleye populations are quite successful (Stroud, 1949; Muench, MS 1966; Libbey, MS 1969; Wahtola et al., 1972; and others). Attempts to support or maintain walleye by stocking have been made, but usually prove unsuccessful. Houser (pers. comm.) states that stocking to supplement natural populations in Arkansas reservoirs has not been very successful; after 10 years of stocking they remain a minor species in the catch.
Jenkins (1970b) examined the effects of selected reservoir environmental variables on fish standing crop through partial correlation and multiple regression analysis. He showed that the age of a reservoir is positively correlated with walleye standing crop. Similarly, Walburg et al. (1971) found that the proportion of walleye to sauger in Lewis and Clark Lake increased from 1 in 10 in the early 1960s to 3 in 10 in 1968. Erickson (1972) found that reservoirs which support white bass (Morone chrysops) and smallmouth bass (Micropterus doZomieu dolomieu) will also support walleyes. This is probably because of the 26 similar environmental requirements of the three species. Benson (pers. comm.) observed that walleye abundance in all the Missouri River impoundments is increasing mainly because their spawning habitat is increasing.
Food Supply
Hohn (1966) found that diatoms were the first food of pelagic walleye fry in western Lake Erie. Similarly, Paulus (1969) reported that fry under 9.0 mm fed on diatoms. This food source usually increases in abundance after impoundment. For e~ample, in Missouri reservoirs, as in most rivers, diatoms dominate the phytoplankton. Comparison of phytoplankton data from pre- and post-impoundment studies show ten-fold increases after dam closure and flooding (SNBB, 1972). Zooplankters take over in importance in walleye diet with an increase in size. Houde (1967) and Priegel (1963) reported that walleye larvae lead a pelagic existence for their first 4 to 6 weeks, feeding on copepods, Cladocera and fish. Such foods also increase in abundance after impound ment. Therefore, it does not appear that the larval stage of the walleye will be a "critical period" in terms of survival in impoundments. The only forseeable factor is the constantly changing plankton community structure in reservoirs and its effect on the availability to walleye fry. However, according to Regier et al. (1969) "whether walleyes at any life stage have a pronounced preference scale of pr~y species is debatable." Apparently, no one has examined the problem critically, taking into account among other factors, the relative size of predator and prey, although, Houde (1967) stated .that it is probable that selection does occur for the larger food organisms by walleye fry. In short, walleye would be likely to starve only in those waters where few mobile organisms of intermediate size are available.
Adult walleye are primarily piscivorous and hence the abundance of forage fishes will determine their success in impoundments. Most authors consider the yellow perch (Perea flavescens) the primary food of walleye, and feel this is because yellow perch are available in sufficient numbers in most walleye waters (Galligan, 1960). However, "walleye are generalized predators and feed on a wide variety of fishes; - rainbow smelt, alewife, ciscoes, ninespine stickleback, white sucker, longnose sucker, lake whitefish, sauger, spottail shiner, darters, white perch, freshwater drum, trout perch, emerald shiner, common shiner, silver chub, gizzard shad, rock bass, pumpkinseed, black crappie, smallmouth bass, brown bullhead, goldeye, mooneye, cottids, burbot and others" (Scott and Crossman, 1973).
Although other forage species may play an important role in the diet of walleye (e.g. gizzard shad; Henderson, 1967) it appears that' yellow perch may be the most important food item to walleye. Maloney and Johnson (1957) reported that the failuare of the walleye fingerling crop was found to be coincident with the failure of the perch crop which the young walleye used for forage. Similarly, Smith and Krefting 27
(1953) noted a remarkably close correlation between the six year classes of walleye and perch in the Red Lakes, Minnesota. Forney (1974) states that "production of large year classes of young yellow perch should enhance survival of young walleye in the same year." It thus appears that the size of the perch crop may be an important factor in determining the year class strength of the walleye and so it might be in some impounded waters.
Yellow perch appear to do quite well in most reservoirs (Walburg, 1964; Gasaway, 1970; Beckman and Elrod, 1971; and others). The importance of the perch-walleye relationship is illustrated by the North Central Reservoir (1973) investigations. They found that yellow perch increased markedly in abundance from 1958 to 1965, to become the primary forage species, but then declined in Oahe Reservoir. The decline was due to the fact that brushy areas which provided good spawning habitat deteriorated beginning in the lower end of the reservoir. This caused the perch to move upstream. The shift in perch distribution produced a decline in abundance and growth rate of walleye in the lower areas of the reservoir. It also caused walleye to become more abundant in the upper parts of the reservoir as a result of the increase in perch abundance in that area.
Scott and Crossman (1973) state that walleye are highly cannibalistic if small yellow perch or another forage fish are not more readily available. However, a high abundance of perch in impoundments does not necessarily mean a high abundance of walleye, as predation by very numerous small perch upon walleye fry could limit the abundance of walleyes.
The walleye apparently relies largely on sight to find its prey. "Thus, if turbidity is excessive, we would expect the walleye to be an ineffective predator" (Regier et aZ._, 1969). However, its noctural habits (Niemuth et al. _, 1959) and other evidence implies "a partial reliance on some, or all, of the senses of sound, taste or smell for feeding" (Regier et aZ._, 1969). Increased turbidity as the result of impoundment should, therefore, not adversely affect the feeding behaviour of walleye, although it may affect its food organisms.
Growth
Availability of food is probably the limiting factor regarding the growth of walleye in impoundments. In the natural state, Forney (1965) has found that slow growth of the walleye was related to low abundance of prey of suitable size. Also, Morsell (1970) found that the highest consumption of yellow perch occurred in those years when walleye growth was fastest and the lowest consumption occurred in those years when walleye growth was slowest. Years of slow walleye growth corresponded to years when invertebrates predominated in the diet. 28
Stroud, 1949a; b; Netsch and Turner, 1964; Muench, MS 1966; Libbey, MS 1969; Elrod and Hassler, 1969; Lewis, 1970; Tucker and Taub, 1970; Grinstread, 1971; Wahtola et aZ.~ 1972; and others, have studied the growth rate of walleyes in impoundments. Generally, walleyes show a superior growth rate, at least initially, as a result of abundant food supplies. According to Carlander (1948), the richness of the water, population densities and length of growing season are factors that modify growth of the walleye.
Eschmeyer (1940) reported a growth of 41.4 em in two years in impounded T.V.A. waters. Stroud (1949b) found that "the growth rate increased through the first 7 to 9 years of impoundment (in Norris Reservoir) and was probably correlated with the generally increasing food supply. However, growth was somewhat less rapid through the twelfth year of impoundment due to significant increases in population density during this period. Present growth rate is fully as rapid as during the first five years of impoundment." Roseberry (1951) also found that walleye in Clayton Lake, a reservoir in Virginia, had growth rates greater than Stroud's (1949b) after three years of age. Libbey (MS 1969) found growth rates of walleye in Dale Hollow Reservoir were excellent when compared to that of other waters. In general, the growth rates of walleye were substantially better than the national average reported by Vidal (1965). The comparatively long growing season and the abundant forage available undoubtedly contributed to the rapid growth of the walleye in this case. Similarly, Lewis (1970) found that walleye in Canton Reservoir, Oklahoma, had a higher growth rate and an earlier sexual maturity than walleye in most other states. Wahtola et aZ.~ (1972) also found that walleye from Lake Sakakawea (Garrison Reservoir) had good linear growth when compared with walleyes from waters in north central United States. Good growth rates were attributed to an abundance of small goldeye (Hiodon aZosoides) and yellow perch.
The superior growth rate of walleyes in impoundments, cannot be expected to be maintained in most cases. Benson (pers. comm.) stated that forage fish abundance has decreased rapidly in Missouri River reservoirs because of deteriorating spawning and nursery areas (erosion and lack of vegetation). In fact, North Central Reservoir (1973) investigations showed that the mean length of female walleyes in Lake Sharpe declined for the third consecutive year. This was attributed to the lack of reservoir fish foods which have decreased over the years.
As in the natural state, there is a difference in the rate of growth of females over males in impounded waters (Stroud, 1949b; Lewis, 1970; Tucker and Taub, 1970; and others). Libbey (MS 1969) found that females in Dale Hollow Reservoir showed greater annual growth increments in age classes I through III. Females invariably grow faster than males, although Elrod and Hassler (1969) reported males grew at a faster rate than females from age 0 to I and from I to II in Lake Sharpe. However, beginning at age III, mean length of females was greater than that of males in all other years. 29
Water Level Regulation
Although fairly stable or slightly r1s1ng levels during spawning and the incubation period are desirable, stable levels have not proved to be necessary at other times of the year. According to Klingbiel (1971), some of Wisconsin's best walleye populations exist in water storage reservoirs which are lowered as much as 5.4 m each winter, reducing their surface acreage to 25% of normal. Similarly, North Central Reservoir 1970 (1971) investigations reported that fall drawdowns of 12 m had no effect on feeding intensity of walleye in Lake Francis Case, i. e. it did not cause a significant increase in the utilization of forage fish. The results also suggested that fall drawdowns did not have a major effect on survival of young fish.
Distribution
Dendy (1945, 1946a, 1948) examined the relationship of tempera ture and dissolved o~gen to the depth distribution of walleye in Norris, Douglas and Cherokee Reservoirs. The nettings showed two things of importance to depth distribution; "(1) that in the presence of an adequate supply of dissolved o~gen the depth distribution of fish was related to thermal stratification and (2) that the oxygen requirements of fishes were different, with walleye being intermediate between largemouth bass and sauger." Carter and Eley (1968) looked at vertical distribution of fishes (among them walleye) and physicochemical parameters before and after flood waters entered Keystone Reservoir, Oklahoma. They found that flood waters were cooler and carried a heavy load of suspended solids and organic matter. This produced a decrease in temperature, specific conductance and dissolved oxygen and also, an increase in turbidity. The fish responded to these physico-chemical changes by moving closer to the surface. O~gen and turbidity were probably the critical factors in causing a decrease in mean depth.
Grinstread (1971) followed the seasonal distribution of walleye fry in Canton Reservoir, Oklahoma. "In Canton Reservoir, larval walleye moved from the spawning site at the dam soon after hatching and were pelagic throughout the reservoir until May." This is similar to reports by Eschmeyer (1950), Forney (1966), Faber (1967), Johnson (1969) and Priegel (1970) which indicate that young walleye in the large northern lakes leave shoreward areas soon after hatching and are pelagic until they reach a total length of approximately 25 mm. "The young then concentrated in the shallow water near the shoreline, with the highest concentrations within coves near the deeper areas of the reservoir. During August, they returned to open water, concentrating near the bottom. During November and December, they moved to the deepest portion of the reservoir, where they remained throughout January and February." The seasonal distribution described by Grinstead is similar to that observed by Forney (1966) and Johnson (1969) for young walleye in Lakes Oneida and Little Cutfoot Sioux respectively. Beckman and Elrod (1971) also examined the abundance and distribution of young-of-the year fishes 30
in Oahe Reservoir, South Dakota. They found that embayments with permanent tributaries contained more species and had higher levels of abundance than embayments of intermittent streams. In the case of the walleye, abundance was highest in the upper end of the major tributary embayments and decreased toward the lower portions. However, distribution appeared to be random within the smaller bays of lower Oahe.
Summary
Impoundment during the first year tends to produce good growth and excellent survival in most fish. Growth rates and population size of yellow walleye could increase, but will be dependent upon the availability of forage species and spawning success. For example, walleye had a rich year class the year when Lake Sharpe, South Dakota, was impounded. Elrod and Hassler (1969) observed that when a new lake is created by a dam in a river, pike and walleye numbers tend to change in reversed directions; pike decline and walleye increase. Similarly, Jenkins (1970) found that with increase in reservoir age, an increase in walleye occurs. 31
WALLEYE REPRODUCTION
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WALLEYE IMPOUNDMENT LITERATURE
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1959. History of fish and fishing in Norris - a TVA tributary reservoir. Proc. 12th Ann. Conf. Southeastern Ass. Game Fish Comm. (195R):ll6-127.
CLARK, C. F. 1965. Walleyes in Ohio and its management. Ohio Div. Wildl . 25 pp .
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1946a. Further studies of depth distribution of fish, ------~--~--~Norris Reservoir, Tennesse, J. Tenn. Acad. Sci. 21(1):94-104.
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1948. Predicting dep.th distribution of fish in ------~--~~~-three TVA storage-type reservoirs. Trans. Amer. Fish. Soc. 75:65-71.
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------1969. An evaluation of Ohio's walleye stocking programs. Ohio Dept. Natur. Resour., Fed. Aid Fish Wildl. Restoration Proj. F-29-R-8, Jobs 9-1 to 9-6, 44 pp.
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------:------1967b. Management consultation and experimental design. Ohio Dept. Natur. Res our., Fed. Aid Fish Wildl. Restoration Proj. F-29-R-6, Job 12, 7 pp.
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------196lb. Report of fisheries investigations during the second year of impoundment of Oahe Reservoir, South Dakota, 1959. S. Dak. Dept. Game, Fish Parks, Fed. Aid Fish Wildl. Restoration Proj. F-1-R-9, Jobs 12, 13, 14, 55 pp.
1963. Report of fisheries investigations during the ------~----~-fourth year of impoundment of Oahe Reservoir, South Dakota, 1961. S. Dak. Dept. Game, Fish Parks, Fed. Aid Fish Wildl. Restora tion Proj. F-1-R-11, Jobs 10, 11, 12, 36 pp.
1964a. Summation of four years of creel census on ------~--Oahe tailwaters, July 1959 through June 1963. S. Dak. Dept. Game, Fish Parks, Fed. Aid Fish Wildl. Restoration Proj. F-1-R-3, Job 12-A, 20 pp.
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1966. Factors affecting first year growth of walleye in Oneida Lake. New York. N.Y. Fish and Game J. 13(2):146-167.
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------1967. The sport fishery of Tenkiller Ferry Reservoir, Oklahoma. Okla. Fish. Res. Lab. Bull. 7:21 pp.
1970. Changes in the fish population in Lake ------~------~--Francis Case in South Dakota in the fist 16 years of impoundment. U. S. Bur. Sport Fish. Wildl., Tech. Paper 56:30 pp.
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______1962a. Report of fisheries investigations during the seventh year of impoundment of Cavins Point Reservoir, South Dakota, 1961. S. Dak. Dept. Game, Fish Parks, Fed. Aid Fish Wildl. Restoration Proj. F-1-R-11, Job 1, 2, 3-A, 7, 40 pp.
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------~~~~---1932b. The depth distribution of certain species of fish in some of the lakes of New York. Trans. Amer. Fish. Soc. 62:331-335.
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1961c. Observations on the limnological dynamics of ------~~~~Tenkiller Ferry Reservoir. Survey of Walleye literature. Okla. Dept. Wildl. Conserv. Fed. Aid Div. Job Compl. Rept. Sept. 1961:43-52.
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ADDITIONAL CITATIONS
BAERENDS, r,. P. 1957. The ethological analysis of fish behavior, pp. 229-269. M. E. Brown (Ed.), The Physiology of Fishes. Vol 2. Academic Press, N. Y.
CALLIGAN, J. P. 1960. Winter food habits of pike perch in Oneida Lake. N.Y. Fish and Game 7(2):156-157.
HASLER, A. D. 1966. Underwater guidposts. Univ. Wise. Press, Madison, Wise. 155 pp.
HOliDE, E. D. 1969. Sustained swimming ability of larvae of walleye (Stizostedion vitreum vitreum) and yellow perch (Perea flavescens). J. Fish. Res. Bd. Canada 26(6): 1647-1659.
MALONEY, J. E., and F. H. JOHNSON. 1955. Life histories and inter relationships of walleye and yellow perch, especially during their first summer, in two Minnesota lakes. Trans. Amer. Fish. Soc. 85:191-202.
RADTKE, L. D., and J. L. TURNER. 1967. High concentrations of total dissolved solids block spawning migration of striped bass, Roccus saxatilis~ in the San Joaquin River, California. Trans. Amer. Fish. Soc. 96(4):405-407.
VIDAL, P. 1965. Halleye and sauger facts. Ill. Dept. of Conserv., Div. Fish., Fish Mgmt. Mimeo. No. 31:3 pp.