Density and composition of macroinvertebrate populations were examined prior to and during the migration and spawning of coho and chinook salmon in a small stream in Michigan. Data gathered from bottom samples indicated that disruption of substrate in the experimental area resulted in a significant decrease in the numbers and species of the macroinvertebrate community. This reduction of invertebrates can create a severe stress on native trout populations

Denison, Douglas L

Meier, Peter G

English

ValpoScholar

The Great Lakes Entomologist 
Volume 12 
Number 1 - Spring 1979 Number 1 - Spring 1979 Article 10 
April 1979 
Effects of Salmon Spawning Activity on Macroinvertebrates in a 
Small Michigan Stream 
Douglas L. Denison 
University of Michigan 
Peter G. Meier 
University of Michigan 
Follow this and additional works at: https://scholar.valpo.edu/tgle 
 Part of the Entomology Commons 
Recommended Citation 
Denison, Douglas L. and Meier, Peter G. 1979. "Effects of Salmon Spawning Activity on 
Macroinvertebrates in a Small Michigan Stream," The Great Lakes Entomologist, vol 12 (1) 
Available at: https://scholar.valpo.edu/tgle/vol12/iss1/10 
This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. 
It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. 
For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 
THE GREAT LAKES ENTOMOLOGIST 
EFFECTS OF SALMON SPAWNING ACTIVITY OIU 
MACROINVERTEBRATES IN A SMALL MICHIGAN STREAM 
Douglas L. Denison and Peter G. ~ e i e r '  
ABSTRACT 
Density and composition of macroinvertebrate populations were examined prior to  
and during the migration and spawning of coho and chinook salmon in a small stream in 
Michigan. Data gathered from bottom samples indicated that disruption of substrate in 
the experimental area resulted in a significant decrease in the numbers and species of the 
macroinvertebrate community. This reduction of invertebrates can create a severe stress 
on native trout populations. 
As salmon migrate upstream, severe disturbances in the stream substrate occur due to 
vigorous swimming movement and redd-digging activity. The benthic population is 
reduced by this disruption, which destroys the habitats provided by the deposited silt and 
sand and the existing macrophytic growth. A mean salmon redd area of 2.8 m2 was 
reported by Burner (1951) for the Toutle River, a tributary of the Columbia River, and 
Hildebrand (1971) estimated that as much as 20,773 m2 of substrate area was disrupted 
by 7,000 spawning female salmon in Michigan's Platte River. The loss of benthic 
macroinvertebrates in such large areas may be temporarily catastrophic and results in a 
substantial decrease in the food supply available to native salmonids. 
Pacific coho salmon (Onchorynchus kisutch Walbaum) were planted by the Michigan 
Department of Natural Resources in three Michigan streams during 1966. The stocking 
project was adopted in an effort to reduce extremely large populations of the alewife 
(Alosa pseudoharengus Wilson) in the Great Lakes, while providing a valuable sport fish 
resource. Two years after the release, strays of returning adult salmon spawners were 
entering most trout streams of northern Lake Michigan and southern Lake Superior, 
causing disruption of the benthic macroinvertebrate communities in these streams. 
One such stream, Cedar Creek, was chosen for study in order that the actual effects 
of the spawning disturbance might be quantified. The benthic population was sampled 
and examined before and after the spawning period and changes have been evaluated to 
determine the severity of the damage. In addition, community diversity and similarity 
within the study area have been calculated. 
STUDY AREA 
Cedar Creek is located 6.4 km north of Traverse City in Elmwood Township, Leelanau 
County, Michigan (T.28N,R.llW,S.28). It is a relatively short stream, flowing easterly 
from Cedar Lake into the west arm of Grand Traverse Bay. The substrate of 40% gravel, 
40% sand, and 20% silt is reported as being optimal for salmon spawning (Shapovalov and 
Taft, 1954). The stream averages 5.3 m in width, 0.39 m in depth and maintains a stable 
flow throughout the year. I t  supports a diverse population of macroinvertebrates and 
trout (brown, rainbow and brook trout). Vegetation surrounding the stream is heavy, 
with cedar and lowland hardwoods dominant. 
The study area begins 30 m from the stream mouth and extends 121 m upstream to a 
small dam. Within this area, four sub-sections, each 15 m in length, were designated for 
study purposes. The downstream sub-sections, 3 and 4, displayed an average current of 
- - 
l ~ e ~ a r t m e n t  of Environmental and Industrial Health, School of Public Health, The 
University of Michigan, Ann Arbor, MI 48109. 
1
Denison and Meier: Effects of Salmon Spawning Activity on Macroinvertebrates in a Sm
Published by ValpoScholar, 1979
58 THE GREAT LAKES ENTOMOLOGIST Vol. 12, No. 1 
15 cm sec-', a velocity which allows for deposition of silt and sand which support 
macrophytes Potarnogeton, Myriophllum, and Ceratophyllum cover 65% of the substrate 
in these sub-sections. Upstream, in subsections 1 and 2, velocity increases to over 60 cm 
sec-I. The substrate in this area is composed of large gravel and some sand with very 
little macrophytic growth. 
METHODS 
The entire study area was divided lengthwise in order to create control (C) and 
experimental (E) sections. The experimental section was easily accessible to adult 
migrating salmon. The control section, however, was enclosed with 3.5 cm wire mesh. 
This fence allowed native fish species to pass freely between the experimental and control 
sections while preventing anadromous fish from entering the control section. 
Two sampling periods were chosen: the f i s t  in August, 1976, prior to the salmon run, 
and the second in November, 1976, five weeks after the run had started. Sampling sites 
were determined by a coordinate system within each sub-section, selected from a table of 
random numbers. The modified Hess sampler of 0.052 m2 was used to take triplicate 
bottom samples from each sub-section (Waters and Knapp, 1961). 
AU aquatic macroinvertebrates from the samples were hand picked, enumerated and 
identified to the lowest practical levels (Hilsenhoff, 1975). The Shannon-Weaver function 
(Lloyd, et  al., 1968), community similarity coefficients and the single-tail F-test 
(Campbell, 1967) were determined for each sample. These tests were used to compare 
control and experimental populations before and after the salmon run. The control 
populations before and after the run were also compared in order to verify the absence 
of change in this section during the sampling periods. 
RESULTS 
The first mature female salmon entered the stream on 12 October, 1976 and large 
concentrations remained throughout November. An informal survey conducted on 22 
October indicated approximately 420 salmon in the study area. It was estimated that one 
half of these were spawning females which could potentially disrupt 600 mZ of substrate. 
Since the total study area was only 557 mZ,  virtually all of the substrate was overturned, 
causing complete elimination of periphytic growth. Redds were observed throughout the 
stream bed in the experimental section, with depths as great as 40 cm and areas as large 
as 2.6 ma. The disruption was most noticeable in the upstream sub-sections, E-1 and E-2. 
The macroinvertebrate data compiled in Table 1 are expressed as the mean value of 
three samples. During the study, a total of 55 invertebrate genera representing eight 
orders were identified The major portion of these belonged to the family Chironomidae 
(Diptera). Sub-section 4, which had a sand-silt substrate, produced the greatest number of 
species and individuals. 
The spawning activity affected the entire benthic community, resulting in an 
overall reduction of individuals, particularly in sub-sections E-1 and E-2 (Table I) .  The 
mean number of organisms in E-1 and E-2 fell below two individuals in November. The 
comparable control sub-sections (C-1 and C-2) showed a mean of 207 individuals during 
the same sampling period. A similar decrease in the mean number of taxa was also found, 
with the control section averaging 16.5 taxa and the experimental section 1.5. Since the 
greatest spawning activity occurred in sub-sections E-1 and E-2, the reduction in mean 
number and taxa was most severe. 
The Shannon-Weaver Index was applied to samples taken in both sampling periods and 
the calculated index values appear in Table 2. August samples showed relatively constant 
diversity values throughout both the control and experimental sections, ranging from a 
low value of 2.10 (C-4) to  a high value of 2.95 (E-2). In November, diversity index values 
dropped in sub-sections E-1 and E-2, the area of' greatest spawning activity. Diversity in 
the control section showed little variance between August and November. Sub-sections 
E-3 and E-4 showed a slight increase in diversity between August and November. These 
sub-sections tended to support benthos composed primarily of midges. Since the 
2
The Great Lakes Entomologist, Vol. 12, No. 1 [1979], Art. 10
https://scholar.valpo.edu/tgle/vol12/iss1/10
1979 THE GREAT LAKES ENTOMOLOGIST 5 9 
TABLE 1. Macroinvertebrate Composition of Study Area. 
Section Sub-section Month No. Taxa No. Organisms No. Diptera 
Control 
Control 
Experimental 1 
2 
Experimental 1 
2 
August 17 
19 
16 
18 
November 17 
20 
13 
20 
August 15 
18 
17 
21 
November 1 
2 
12 
18 
TABLE 2. Shannon-Weaver Index of Diversity. 
- 
Section 
Mean Diversity Mean Diversity 
Sub-section (Aug.1 (Nov.) 
Control 
Experimental 
Chironomidae are active within the sediment, the spawning and swimming activities had 
little effect. 
The community similarity (Jaccard, 1928) showed that, before spawning, the benthos 
in the control and experimental sections displayed similarity coefficients near unity 
(Table 3). This coefficient dropped after spawning occurred, indicating a significant 
change in the communities. Furthermore, testing the similarity of each sub-section before 
and after the spawning period showed a distinct difference in the experimental popula- 
tions, while similarity remained near unity in the control section. 
In order to verify the significance of this change, the single-tailed F-test was applied. 
For both collection periods, the F-test indicated no significant difference between the 
total number of organisms and taxa for all subsections, except for E-1 and E-2. 
These areas showed a statistically significant loss of macroinvertebrates coincident with a 
very high number of spawning salmon. 
DISCUSSION 
As a result of the salmon spawning, some areas of the experimental section underwent 
a significant decrease in the total number of benthic macroinvertebrates, total number of 
3
Denison and Meier: Effects of Salmon Spawning Activity on Macroinvertebrates in a Sm
Published by ValpoScholar, 1979
6 0 THE GREAT LAKES ENTOMOLOGIST Vol. 12, No. 1 
TABLE 3. Community Similarity 
Sub-sections Coefficient of Community Similarity 
August, 1976 (before run) 
C-1 vs. E-1 
C-2 vs. E-2 
C-3 vs. E-3 
C-4 vs. E-4 
November, 1976 (after run) 
C-1 vs. E.l 
C-2 VS. E-2 
C-3 VS. E-3 
C-4 vs. E-4 
Before Run vs. After Run 
C-1 vs. C-1 
C-2 vs. C-2 
C-3 vs. C-3 
C-4 vs. C-4 
macroinvertebrate taxa, and community diversity. This decrease is d i r e c e  attributable to 
the vigorous swimming behavior and redd-digging activity of the spauning salmon. The 
importance of this decrease in terms of the entire stream cannot be ascertained from this 
phase of study. Further work is required to determine the long-term effects on the 
stream ecosystem. Continuation of the spawning activity will cause further reduction and 
loss of representative organisms can be experienced over time, resulting in an unstable 
community. This loss of important food chain members would have its peatest impact 
on the next trophic level, the native trout, since they depend on the benthic community 
as a major food source. 
The rate of recolonization by drift and adult reproduction must be sondered  when 
assessing the stream once it has become denuded of macroinvertebrater Hildebrand 
(1971) obse~ved that five months was insufficient to completely recolonize a portion of 
the Platte River during the Fall of 1967. If indeed this process requires a loWr time 
period, the food supply will remain below normal levels during the peak growing and 
feeding periods of trout. Taube (1975) reports that the reduction of benthic o-s is 
responsible for reduced reproduction in brown trout although no noticeable effect on 
older trout can be seen. If reproduction is slowed and the trend is for survival of older 
trout, a change in the population structure is imminent. 
LITERATURE CITED 
Burner, C. J. 1951. Characteristics of spawning nests of Columbia River salmon. U.S. Fish 
Wildl. Serv. Fish Bull. 61:97-110. 
Campbell, R. C. 1967. Statistics for biologists. Cambridge Univ. Press. 
Hildebrand, S. G. 1971. The effect of coho salmon on the benthic imertebrates of the 
Platte River, Benzie County, Michigan. Trans. Amer. Fish. Soc. 100:6168. 
Iiilsenhoff, W. L. 1975. Aquatic insects of Wisconsin. Tech. BulL No. 89, Dept  Sat. 
Res., Madison, Wis. 
Jaccard, P. 1928. Die statistische-floristische methode als _erundlage der pflanzensozie 
logie. Handb. Biol. Arbeitsmeth. Abderhalden. 11:165-202. 
4
The Great Lakes Entomologist, Vol. 12, No. 1 [1979], Art. 10
https://scholar.valpo.edu/tgle/vol12/iss1/10
1979 THE GREAT LAKES ENTOMOLOGIST 6 1 
Lloyd, M., J. H. Zar and J. R. Karr. 1968. On the calculation of information-theoretical 
measures of diversity. Amer. Midl. Nat. 79: 257-272. 
Shapovalov, L. and A. C. Taft. 1954. The life histories of the steelhead trout and silver 
salmon with special reference to Waddell Creek, California, and recommendations 
regarding their management. Calif. Fish and Game Fish Bull. 98:l-375.  
Taube, C. M. 1975. Abundance and growth, biomass and interrelationship of trout and 
coho salmon in the Platte River. Fish Rep. No. 1830, Mich. Dept. Nat. Res. Fish. Div. 
Waters, T. F. and R. J. Knapp. 1961. An improved stream bottom sampler. Trans. Amer. 
Fish. Sac. 93:225-227. 
5
Denison and Meier: Effects of Salmon Spawning Activity on Macroinvertebrates in a Sm
Published by ValpoScholar, 1979


Effects of Salmon Spawning Activity on Macroinvertebrates in a Small Michigan Stream

Valparaiso University

The Great Lakes Entomologist Volume 12 Number 1 - Spring 1979 Number 1 - Spring 1979 Article 10 April 1979 Effects of Salmon Spawning Activity on Macroinvertebrates in a Small Michigan Stream Douglas L. Denison University of Michigan Peter G. Meier University of Michigan Follow this and additional works at: https://scholar.valpo.edu/tgle  Part of the Entomology Commons Recommended Citation Denison, Douglas L. and Meier, Peter G. 1979. "Effects of Salmon Spawning Activity on Macroinvertebrates in a Small Michigan Stream," The Great Lakes Entomologist, vol 12 (1) Available at: https://scholar.valpo.edu/tgle/vol12/iss1/10 This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. THE GREAT LAKES ENTOMOLOGIST EFFECTS OF SALMON SPAWNING ACTIVITY OIU MACROINVERTEBRATES IN A SMALL MICHIGAN STREAM Douglas L. Denison and Peter G. ~ e i e r '  ABSTRACT Density and composition of macroinvertebrate populations were examined prior to  and during the migration and spawning of coho and chinook salmon in a small stream in Michigan. Data gathered from bottom samples indicated that disruption of substrate in the experimental area resulted in a significant decrease in the numbers and species of the macroinvertebrate community. This reduction of invertebrates can create a severe stress on native trout populations. As salmon migrate upstream, severe disturbances in the stream substrate occur due to vigorous swimming movement and redd-digging activity. The benthic population is reduced by this disruption, which destroys the habitats provided by the deposited silt and sand and the existing macrophytic growth. A mean salmon redd area of 2.8 m2 was reported by Burner (1951) for the Toutle River, a tributary of the Columbia River, and Hildebrand (1971) estimated that as much as 20,773 m2 of substrate area was disrupted by 7,000 spawning female salmon in Michigan's Platte River. The loss of benthic macroinvertebrates in such large areas may be temporarily catastrophic and results in a substantial decrease in the food supply available to native salmonids. Pacific coho salmon (Onchorynchus kisutch Walbaum) were planted by the Michigan Department of Natural Resources in three Michigan streams during 1966. The stocking project was adopted in an effort to reduce extremely large populations of the alewife (Alosa pseudoharengus Wilson) in the Great Lakes, while providing a valuable sport fish resource. Two years after the release, strays of returning adult salmon spawners were entering most trout streams of northern Lake Michigan and southern Lake Superior, causing disruption of the benthic macroinvertebrate communities in these streams. One such stream, Cedar Creek, was chosen for study in order that the actual effects of the spawning disturbance might be quantified. The benthic population was sampled and examined before and after the spawning period and changes have been evaluated to determine the severity of the damage. In addition, community diversity and similarity within the study area have been calculated. STUDY AREA Cedar Creek is located 6.4 km north of Traverse City in Elmwood Township, Leelanau County, Michigan (T.28N,R.llW,S.28). It is a relatively short stream, flowing easterly from Cedar Lake into the west arm of Grand Traverse Bay. The substrate of 40% gravel, 40% sand, and 20% silt is reported as being optimal for salmon spawning (Shapovalov and Taft, 1954). The stream averages 5.3 m in width, 0.39 m in depth and maintains a stable flow throughout the year. I t  supports a diverse population of macroinvertebrates and trout (brown, rainbow and brook trout). Vegetation surrounding the stream is heavy, with cedar and lowland hardwoods dominant. The study area begins 30 m from the stream mouth and extends 121 m upstream to a small dam. Within this area, four sub-sections, each 15 m in length, were designated for study purposes. The downstream sub-sections, 3 and 4, displayed an average current of - - l ~ e ~ a r t m e n t  of Environmental and Industrial Health, School of Public Health, The University of Michigan, Ann Arbor, MI 48109. 1Denison and Meier: Effects of Salmon Spawning Activity on Macroinvertebrates in a SmPublished by ValpoScholar, 197958 THE GREAT LAKES ENTOMOLOGIST Vol. 12, No. 1 15 cm sec-', a velocity which allows for deposition of silt and sand which support macrophytes Potarnogeton, Myriophllum, and Ceratophyllum cover 65% of the substrate in these sub-sections. Upstream, in subsections 1 and 2, velocity increases to over 60 cm sec-I. The substrate in this area is composed of large gravel and some sand with very little macrophytic growth. METHODS The entire study area was divided lengthwise in order to create control (C) and experimental (E) sections. The experimental section was easily accessible to adult migrating salmon. The control section, however, was enclosed with 3.5 cm wire mesh. This fence allowed native fish species to pass freely between the experimental and control sections while preventing anadromous fish from entering the control section. Two sampling periods were chosen: the f i s t  in August, 1976, prior to the salmon run, and the second in November, 1976, five weeks after the run had started. Sampling sites were determined by a coordinate system within each sub-section, selected from a table of random numbers. The modified Hess sampler of 0.052 m2 was used to take triplicate bottom samples from each sub-section (Waters and Knapp, 1961). AU aquatic macroinvertebrates from the samples were hand picked, enumerated and identified to the lowest practical levels (Hilsenhoff, 1975). The Shannon-Weaver function (Lloyd, et  al., 1968), community similarity coefficients and the single-tail F-test (Campbell, 1967) were determined for each sample. These tests were used to compare control and experimental populations before and after the salmon run. The control populations before and after the run were also compared in order to verify the absence of change in this section during the sampling periods. RESULTS The first mature female salmon entered the stream on 12 October, 1976 and large concentrations remained throughout November. An informal survey conducted on 22 October indicated approximately 420 salmon in the study area. It was estimated that one half of these were spawning females which could potentially disrupt 600 mZ of substrate. Since the total study area was only 557 mZ,  virtually all of the substrate was overturned, causing complete elimination of periphytic growth. Redds were observed throughout the stream bed in the experimental section, with depths as great as 40 cm and areas as large as 2.6 ma. The disruption was most noticeable in the upstream sub-sections, E-1 and E-2. The macroinvertebrate data compiled in Table 1 are expressed as the mean value of three samples. During the study, a total of 55 invertebrate genera representing eight orders were identified The major portion of these belonged to the family Chironomidae (Diptera). Sub-section 4, which had a sand-silt substrate, produced the greatest number of species and individuals. The spawning activity affected the entire benthic community, resulting in an overall reduction of individuals, particularly in sub-sections E-1 and E-2 (Table I) .  The mean number of organisms in E-1 and E-2 fell below two individuals in November. The comparable control sub-sections (C-1 and C-2) showed a mean of 207 individuals during the same sampling period. A similar decrease in the mean number of taxa was also found, with the control section averaging 16.5 taxa and the experimental section 1.5. Since the greatest spawning activity occurred in sub-sections E-1 and E-2, the reduction in mean number and taxa was most severe. The Shannon-Weaver Index was applied to samples taken in both sampling periods and the calculated index values appear in Table 2. August samples showed relatively constant diversity values throughout both the control and experimental sections, ranging from a low value of 2.10 (C-4) to  a high value of 2.95 (E-2). In November, diversity index values dropped in sub-sections E-1 and E-2, the area of' greatest spawning activity. Diversity in the control section showed little variance between August and November. Sub-sections E-3 and E-4 showed a slight increase in diversity between August and November. These sub-sections tended to support benthos composed primarily of midges. Since the 2The Great Lakes Entomologist, Vol. 12, No. 1 [1979], Art. 10https://scholar.valpo.edu/tgle/vol12/iss1/101979 THE GREAT LAKES ENTOMOLOGIST 5 9 TABLE 1. Macroinvertebrate Composition of Study Area. Section Sub-section Month No. Taxa No. Organisms No. Diptera Control Control Experimental 1 2 Experimental 1 2 August 17 19 16 18 November 17 20 13 20 August 15 18 17 21 November 1 2 12 18 TABLE 2. Shannon-Weaver Index of Diversity. - Section Mean Diversity Mean Diversity Sub-section (Aug.1 (Nov.) Control Experimental Chironomidae are active within the sediment, the spawning and swimming activities had little effect. The community similarity (Jaccard, 1928) showed that, before spawning, the benthos in the control and experimental sections displayed similarity coefficients near unity (Table 3). This coefficient dropped after spawning occurred, indicating a significant change in the communities. Furthermore, testing the similarity of each sub-section before and after the spawning period showed a distinct difference in the experimental popula- tions, while similarity remained near unity in the control section. In order to verify the significance of this change, the single-tailed F-test was applied. For both collection periods, the F-test indicated no significant difference between the total number of organisms and taxa for all subsections, except for E-1 and E-2. These areas showed a statistically significant loss of macroinvertebrates coincident with a very high number of spawning salmon. DISCUSSION As a result of the salmon spawning, some areas of the experimental section underwent a significant decrease in the total number of benthic macroinvertebrates, total number of 3Denison and Meier: Effects of Salmon Spawning Activity on Macroinvertebrates in a SmPublished by ValpoScholar, 19796 0 THE GREAT LAKES ENTOMOLOGIST Vol. 12, No. 1 TABLE 3. Community Similarity Sub-sections Coefficient of Community Similarity August, 1976 (before run) C-1 vs. E-1 C-2 vs. E-2 C-3 vs. E-3 C-4 vs. E-4 November, 1976 (after run) C-1 vs. E.l C-2 VS. E-2 C-3 VS. E-3 C-4 vs. E-4 Before Run vs. After Run C-1 vs. C-1 C-2 vs. C-2 C-3 vs. C-3 C-4 vs. C-4 macroinvertebrate taxa, and community diversity. This decrease is d i r e c e  attributable to the vigorous swimming behavior and redd-digging activity of the spauning salmon. The importance of this decrease in terms of the entire stream cannot be ascertained from this phase of study. Further work is required to determine the long-term effects on the stream ecosystem. Continuation of the spawning activity will cause further reduction and loss of representative organisms can be experienced over time, resulting in an unstable community. This loss of important food chain members would have its peatest impact on the next trophic level, the native trout, since they depend on the benthic community as a major food source. The rate of recolonization by drift and adult reproduction must be sondered  when assessing the stream once it has become denuded of macroinvertebrater Hildebrand (1971) obse~ved that five months was insufficient to completely recolonize a portion of the Platte River during the Fall of 1967. If indeed this process requires a loWr time period, the food supply will remain below normal levels during the peak growing and feeding periods of trout. Taube (1975) reports that the reduction of benthic o-s is responsible for reduced reproduction in brown trout although no noticeable effect on older trout can be seen. If reproduction is slowed and the trend is for survival of older trout, a change in the population structure is imminent. LITERATURE CITED Burner, C. J. 1951. Characteristics of spawning nests of Columbia River salmon. U.S. Fish Wildl. Serv. Fish Bull. 61:97-110. Campbell, R. C. 1967. Statistics for biologists. Cambridge Univ. Press. Hildebrand, S. G. 1971. The effect of coho salmon on the benthic imertebrates of the Platte River, Benzie County, Michigan. Trans. Amer. Fish. Soc. 100:6168. Iiilsenhoff, W. L. 1975. Aquatic insects of Wisconsin. Tech. BulL No. 89, Dept  Sat. Res., Madison, Wis. Jaccard, P. 1928. Die statistische-floristische methode als _erundlage der pflanzensozie logie. Handb. Biol. Arbeitsmeth. Abderhalden. 11:165-202. 4The Great Lakes Entomologist, Vol. 12, No. 1 [1979], Art. 10https://scholar.valpo.edu/tgle/vol12/iss1/101979 THE GREAT LAKES ENTOMOLOGIST 6 1 Lloyd, M., J. H. Zar and J. R. Karr. 1968. On the calculation of information-theoretical measures of diversity. Amer. Midl. Nat. 79: 257-272. Shapovalov, L. and A. C. Taft. 1954. The life histories of the steelhead trout and silver salmon with special reference to Waddell Creek, California, and recommendations regarding their management. Calif. Fish and Game Fish Bull. 98:l-375.  Taube, C. M. 1975. Abundance and growth, biomass and interrelationship of trout and coho salmon in the Platte River. Fish Rep. No. 1830, Mich. Dept. Nat. Res. Fish. Div. Waters, T. F. and R. J. Knapp. 1961. An improved stream bottom sampler. Trans. Amer. Fish. Sac. 93:225-227. 5Denison and Meier: Effects of Salmon Spawning Activity on Macroinvertebrates in a SmPublished by ValpoScholar, 1979

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Effects of Salmon Spawning Activity on Macroinvertebrates in a Small Michigan Stream

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