PB81-198954
Natural Variation  in  Abundance of Salmonid
Populations  in  Streams  and Its Implications for
Design of  Impact Studies.  A Review
Oregon State Univ.
Corvallis
Prepared  for

Corvallis Environmental  Research Lab.,  OR
Feb 81
                      U.S. DEPARTMENT OF COMMERCE
                   National Technical Information Service

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                                            EPA 600/3-81-021
                                            Febraury 1981
NATURAL VARIATION IN ABUNDANCE OF SALMONID POPULATIONS IN STREAMS
        AND ITS.IMPLICATIONS FOR DESIGN OF IMPACT STUDIES
                            A Review
                              by
               James D. Hall and Ned J. Knight
             Department of Fisheries and Wildlife
                    Oregon State University
                     Corvallis, OR  97331
                        Project Officer
                      Jack H. Gakstatter
             U.S. Environmental Protection Agency
                     200 S.W. 35th Street
                     Corvallis, OR  97330

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. FRO M THE BEST COP Y F URN ISH ED USB Y
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. .
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. IN THE INTEREST OF MAKING AVAILABLE
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BIBLIOGRAPHIC INFORMATION
PB81-198954
Natural Variation in Abundance of Salmonid Populations in
,Streams and Its Implication's for Design of Impact Studies. A
Review,
, Fe b 81
JamesD. Hall, and Ned J.Knight.,
PERFORMER:
Oregon State Univ., Corvallis. Dept. of
Fisheries and wildlife.
EPA~600/3-81-021 .
SPONSOR:
Corvallis Environmental Research Lab., .OR.
," ------ .----. -- -- - - -'------.. --~
Literature on stock size and production of salmonid
populations in streams has been reviewed. The objective is
to bring together data on the magnitude of natural variation
in population size and to relate this variability to
environmental conditions where possible. Recommendations are
presented for the use of this information in designing
studies to ~stimate the impact of non-point source
pollutiion. A partially annotated bibliography of 260
relevant reference is included.
KE YWO RDS:
*Ecology, *Salmon, *Water pollution.
Available from the National Technical Information Service,
Springfield, Va. 22161
PR ICE CODE:
PC A05/MF AO 1
i

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DISCLAIMER
This report has been reviewed by the Corvallis Environmental Research
Laboratory, U~S. Environmental Protection Agency, and approved for pub-
lication~ Approval does not signify that the contents necessarily reflect
the views and policies of the U.S. Environmental Protection Agency, nor does
mention of trade names or commercial products constitute endorsement or
recommendation for use.
ii

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ABSTRACT
, '
Literature on stock size and, production of salmonid populations in
streams has been reviewed. The objective is to bring together data on the
magnitude of natural variation in population size and to relate this vari-
ability to environmental conditions where possible. Recommendations are
presented for the use of this information in,designing studies to estimate
the impact of non-point source pollution. A partially annotated bibliography
of 260 ,relevant references is included. '
A number of long-term studies, some up to 15-20 years, have provided
useful data on temporal variation in population abundance. Other studies
have examined spatial variation. Data from the best examples of both kinds
of variation are presented in, Appendix Tables. Temporal and spatial varia-
tion may be as high as several orders of magnitude in the extreme, and even
at the least are sufficient to mask very significant perturbations caused by
non-point source pollutants. Environmental variables most closely associated
with spatial variation are those relating to the quality of salmonid habitat,
particularly physical characteristics such as cover in its many forms. '
Streamflow and food abundance have been associated with both temporal and
~patial variation. In general, physical characteristics of, habitat seem
most promising as descriptors of variability.
Systems of rating habitat quality should receive considerable emphasis
in attempts to minimize the effects of natural variation in the evaluation
of impacts of non-point source pollutants. First priority should be placed
on assessment of physical features. This approach has been used so far
mainly to explain spatial variation,' but has promise of explaining temporal
variation as well, particularly in reference to fluctuation in streamflow.
. The other major emphasis should be in further development of systems of
stream and watershed classification. The most useful of these devised to
date take a perspective from geomorphology and focus on the potential of a
stream system for oiological production. More emphasis on study of basic
processes in stream ecosystems and more extensive use of paired comparisons
in design of impact studies are also suggested as means of more clearly
separating natural variation from damage caused by non-point source
pollutants.
. ., ~
III

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CONTENTS
Abstract. . . . . . . . . . . . . . . . . . . . .

Tables. . . . -- . . . . . . . . . . . . . . . . .

Appendix Tables.
Acknowledgment. .
. . . . .
. . . . .
. . . .
. . . .
. ; . .
. . . . .
. . . . . . . .
. . . . . .
. . . . . . . . .
. . . . .
1.
2.
3.
4.
Introduction. . . . . . . . . '. ; . . '. . . . . . . . . . . .

Conclusions and Recommendations. . . . . . . . . . . . . . .
Studies of Variability. . . . . . . . . . . . . .
Factors Affecting Natural Variability. . . . . . . . . . . .
Physical factors. . . . . . . . . . . . . .
Biological factors. . . . . . . . . . . . . . . . . . .
Other factors~ . . . . . . . ," . . . . . . . . .
Minimizing the Effects of Variability in Impact Studies
Habitat quality rating systems. . . . . . . . . .
Process studies. .'. .'. . . . . . . . . . . . . . . . .
Stream class{fication. . . . . . . . . . . . . . .
Improved study design. . . . . . . . . . . .
5.
Bibliography. . . . . .
Appendix. . . . . . . .
. . .8 . .
. . . . . . . . .
. . . . . . . . .
. . . . . .
. . . .
. . . . . . . . . .
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Number
TABLES
1
Long-term Studies of Stream Salmonid Populations. . .
. . . . .
2
Correlations Between Mean Monthly Discharge and Annual
Smolt Count and Mean June-April Biomass for Coho
Salmon, Alsea Watershed Study, June 1960-May 1969 .
. . . . .
3
Correlations Between Mean Monthly Discharge and Mean
September Biomass in G/M2 for Cutthroat Trout, Alsea
Watershed Study, October 1961-September 1972. . . . .
. . . .
4
Correlations Between Mean Monthly Discharge and Mean
Annual Biomass of Brown, Brook, and Rainbow Trout,
Sagehen Creek, California, 1954-1961. . . . . . . .
. . . . .
5
Summary of Advantages and Disadvantages of the Four
Maj or Approaches to. Watershed Stream Analysis. .
. . . . . .
vi
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4
6
7
9
27

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Number
A-I
A-2
A-3
A-4
A-5
A-6
A-7
A-8
A-9
A-10
A-ll
A-12
APPENDIX TABLES
, ,
Potential Egg Depos1tion and Freshwater Survival of Pink
Saimon, Sash in Creek, A!aska, 1940-1959. . . .
. . . .
Weir Counts of Coho Salmon Fry and Smolts, Sashin Creek
Alaska, 1956-1968. . . . . . . . . . . . . .' .
. . . .
Weir Counts of Downstream Migrating Pink and Chum Salmon
Fry, Hooknose Creek, British COlumbia, 1947-1956 . . .
Biomass of Coho Salmon and Rainbow and Cutthroat Trout in
Streams in the Vicinity of the Carnation Creek
Watershed, British Columbia, 1970-1977 . . . . . . . : . . .
Biomass of Cutthroat Trout, Coho Salmon, and Sculpin in
, Different Habitats of Six British,Columbia Streams,

1973-1976. . . . . . . . . . . . ',' . . . . . . . .
. . . .
Escapement, Potential Egg Deposition, and Freshwater
Survival of Wild Coho Salmon, Minter Creek, Washington,

1938-1953. . . . . . . . . . . . . . . . . : . . . . . . . .
Counts of Spawning Coho Salmon and Smolts at Downstream
Weir on Gnat Creek, Oregon, 1954-1959. . . . . . . .
. . . .
Escapement, Potential Egg Deposition, and Freshwater
Survi val of Coho Salmon, Deer Creek, Oregon, 1959-1971
Estimated Biomass of Juvenile Coho' Salmon, Deer Creek,
Oregon, 1959-1968. . . . . . . . . . . . . . .
. . . .
Escapement, Potential Egg Deposition, and Freshwater
Survival of Coho Salmon, Flynn Creek, Oregon, 1959-1971. . .
Estimated Biomass of Juvenile Coho Salmon, Flynn Creek,
Oregon, 1959-1968. . . . . -'-~ ,- -' . . . . . . . . . . . . .
Escapement, Potential Egg Deposition, and Freshwater
Survival of Coho Salmon, Needle Branch, Oregon,
1959-1971. . . . . . . . . . . . . . . . . . . . .
. . . . .
vii
Page
61
62
62
63
64
65
65
66
66
67
67
68

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'Number
A-i3
A-14
A-15
A-16
A-17
. A-18
A-19
A-20
A-21
A-22
A-23
A-24
A-25
A-26
A-27
APPENDIX TABLES
(continued)
Estimated Biomass of Juvenile Coho Salmon, Needle Branch,
Oregon, 1959-1968. . . . . . . . . . . . .
Biomass of Cutthroat Trout, Alsea Watershed Study,
1962-1973. . . . . . . . . . . . . .
Biomass of SalmonidSpecies in Three Northern California
Streams, 1967-1969 . . : . . . . . . . . . . . . . . .
.Biomass of Brook Trout in 10 Sections of Sagehen Creek,
California, 1952-1961. . . . . . . . . . . . . . . .
. . . .
Biomass of Brown Trout in 10 Sections of Sagehen Creek,
California, 1952-1961. . . . . . . . . . . . . . . .
. . . .
Biomass of Rainbow Trout in 10 Sections of Sagehen Creek,
California, 1952-1961. . . . . . . . . . . . . . . . .
Escapement, Potential Egg Deposition, and Freshwater
. Survival of Coho Salmon, Waddell Creek, California,

1933-1940. . . . . . . . . . . . . . . . . . .
. . . .
Downstream Trap Counts of Steelhead Trout by Age Group,
Waddell Creek, California, 1933-1942 .. .
. . . .
Biomass of Brook, Rainbow, and Brown.Trout, Trout Creek,
Montana, 1950-1951 . ... . . . . . . . . . . . . . . .
Biomass of Brown, Rainbow, and Brook Trout in 11 Sections
of Little Prickly Pear Creek, Montana, Summer 1966 . .
Number of Cutthroat Trout. Trapped, and Number Remaining
in Stream After Trap was Removed, Arnica Creek,
Yellowstone Park, Wyoming, 1950-1958 . . . . . . . .
. . . "
Biomass of Brook Trout, Lawrence Creek, Wisconsin,

1953-1957. . . . . . . . . . . . . . . . . . . .
. . . . . .
September Population Estimates of Ages 0 and I Brook
Trout in Lawrence Creek and Big Roche-A-Cri Creek,
Wisconsin, 1953-1964 . . . . . . . . . . . .
. . . . .
Biomass of Brook'Trout by Age Group in April and September,
Lawrence Creek, Wisconsin, 1960-1970 . . . . . . . . . . . .
Annual Production of Brook Trout by Section and Age Group,
Lawrence Creek, Wisconsin, 1960-1970 . " . . . . . . . . . .
viii
Page
69
69
70
70
71.
71
72
72
73
73
74
75
75
76
77

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'Number
A-28
A-29
A-30
A-31
A-32
A-33
A-34
A-35
A-36
A-37
APPENDIX TABLES
( continued)
Physical Characteristics and Biomass of Brook, Brown,'and
Rainbow Trout in Sections of Three Michigan Streams,

1937 . . . . . . . . . . . . . . . . ." . . . .
Number of Brook Trout Present in September in Hunt Creek,
Michigan by Age-Group. ',' . . . . . . . . . . . . . .
Mean Annual Biomass of Brook Trout in Streams in Matamek
Watershed" Quebec, 1971-1973 . . . . . . . . . . . . .
Numbers of Brook Trout in a 4l1~m Section of Hayes Brook,
Prince Edward Island, 1947-l~60. . . . . . . . . . . .
Counts of, Atlantic Salmon Smo1ts and Seaward Migrating
Brook Trout, Little Codroy River, Newfoundland,

1954-1963. . . . . . . . . . . . .,. . . . . . . . .
. . . .
Biomass of Brown Trout in Tributaries and the Main Stem
of the Upper River Tees System, England, ,1967-1970 .
. . . .
Mean Biomass of Brown Trout in Five Tributaries of the
River Tees System, England, in May, August, and

October. . . '. . . . . . . . . . . . . . . . .
. . e. .
Production of At1antic Salmon and Brown Trout in Three
Sections of Shelligan Burn, Scotland, 1966-1968. . .
. . . .
Biomass of Atlantic Salmon and Brown Trout at the End of
the Growing Season, She11igan Burn, Scotland, 1966-1975.
Biomass of Brown Trout in Six Sections of Horokiwi Stream,
New Zealand, 1940-1941 . . . .- . . . . . . . . . . . . . . .
ix
Page
78
79
80
81
81
82
83
84
85
-
85

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ACKNOWLEDGMENT
We appreciate the advice of Mr. Michael Crouse, who provided the
original impetus for the review, and that of Dr. Jack Gakstatter, Project
Officer. Drs. Hiram Li and Terry Finger reviewed an early version of the
manuscript. Dr. Scott Overton a4ded useful comments to the section on '
study design. Dr. Richard Gard graciously provided unpublished data from
Sagehen Creek, California. The p~rmission of copyright holders and publish-
ers to reprint data tables is gratefully acknowledged. One of us (J.D.H.)
is especially appreciative of space and facilities made available by Dr.
'W. C. Clark, Zoology Department, University of Canterbury, and Dr. R. M.
McDowall, Fisheries Research Laboratory, Ministry of Agriculture and
Fisheries, Christchurch, New Zealand, where final work on the manuscript'
was completed. We would also like to acknowledge the secretarial staff of
the Department of Fisheries and Wildlife for exceptional service under
difficult conditions, and Marthanne Norgren for skillfully coordinating
final production of the manuscript. This is Technical Paper No. 5608, ,
Oregon Agricultural Experiment Station. ' '
x

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SECTION I
INTRODUCTION
Assessment of impacts on streams caused by non-point source pollutants
is now receiving increasing atte~tion. Salmonids are the principal fish
species of economic importance affected in the western United States.Assess-
ment of damage to these populations cannot be undertaken without some under-
standing of natural variation in abundance within and between populations.
Strategies of analysis' must be devised. that will separate natural variation
from effects due to disturbance~ It is the purpose of this review to bring'
together literature and unpublished data on the natural variation in abundance
of salmonid populations in streams and to attempt to relate this variation to
environmental variables--physical, chemical, and biological.
There are two kinds of variability to be considered, spatial and temporal.
Spatial variability can be studied at several levels of resolution, ranging
from microhabitat preferences to variability within and between streams. .
Tempor~l fluctuations in abundance can occur on a diel,. seasonal, or .annual
scale. . .' .
This paper will concentrate on studies ofsalmonid spec1es during that
part of their lives spent in the stream environment. The species include the
coho salmon (Oncorhynchus kisutch), chinook salmon (0. tshawytscha), pink
salmon (0. gorbuscha), chum salmon (0. keta), brown trout (Salmo trutta),
rainbow trout(S. gairdneri), steelhead trout (S. gairdneri-gaLrdneri), cut-
throat trout (S~ clarki), Atlantic salmon (S. salar), brook.trout (Salvelinus
fontinalis), and Dolly Varden (S~ malma). We began this review with the
intention of emphasizing studies on the west coast of North America. However,
we found that most of the quantitative data on variability in resident
salmonid populations came from other areas, and much of that information has
been included. .
Much less information is available on population levels of the fish
species associated with salmonids. Though not included here, the importance
of this element of the aquatic system should be emphasized and steps taken to
fill this gap in our knowledge of stream fish communities.
"-- .~-
I

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SECTION 2
. .
CONCLUSIONS AND RECOMMENDATIONS
The standing stock biomass of salmonid fishes in streams shows great
natural variation, both in time ~d space. 2Reported levels of biomass vary
from zero or just above to just over 60 g/m. This variation is sufficient
to mask large-scale perturbations caused by non-point source pollutants, such
as result from logging and agricultural practices. Among the most important
,causes of variation are differences in physical characteristics of streams,
inCluding streamflow and habitat quality, particularly cover. Biological
factors, such as food abUndance and predation, may sometimes influence abund-
ance, but their mode of action is less clear and the case for their involve-
, ment more equivocal than that of the physical elements of the habitat.
We recommend several courses of action that will help to minimize the
effects of this natural variation when attempts are made to evaluate impacts
of a particular non-point source pollutant. Habitat quality rating systems
are being developed that show promise of explaining much of the spatial varia-
tion in salmonid populations in streams. These rating systems are based pri-
marily on assessment of physical features. They may also help to explain
temporal variation caused by changes in streamflow, but other influences on
temporal variation need further study. The other major approach that may aid
impact assessment is development of schemes of , stream and watershed classifi-
cation, such as those of Platts (1974) and Warren (1979). The latter is
particularly promising in that it focuses' on the potential of a system for
biological production, rather than a particular value of the moment, and takes
a biogeoclimatic perspective. Continuing emphasis on study of the basic
physical and biological processes that lead to growth, mortality, and produc-
tion of. stream salmonids is another promising approach to understanding
natural variation in abundance. Finally, new approaches to the design of
impact studies are suggestd that may aid in more clearly separating natural
variation from that c~used by non-point source pollutants and in monitoring
the time required for biological systems to recover from perturbation.
2

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SECTION 3
STUDIES OF VARIABiLITY
Natural variability of salmonid populations in streams has been measured
by two principal methods. In some streams, weirs or traps have been con-
structed to get reliable counts of migrating fish. Other studies have
examined standing crops in the stream by electro shocking, netting, or angling.
There have beert a number of important long~term studies on natural
variation in abundance ofanadromous and resident species, which are briefly
described in Table 1. As an aid to. further analysis, data from these studies
and others of shorter duration that deal with spatial variation have been
compiled from original sources and are included in tables in the Appendix.
Further description of many studies is included in the annotated bibliography.
We performed some preliminary analyses on the data in the Appendix Tables
and in other publications, in search of general patterns in variation ..-
over the species and geographical areas included. We used the range in
abundance as a fraction of mean abundance for a measure of relative vari-
ability, rather than the coefficient of variation, owing to small sample
sizes. Not surprisingly, the extremes of temporal variation occur in pink
and chum salmon fry; their numbers may vary over several orders of magnitude.
The most stable populations are those of brook trout in Wisconsin and .
Michigan, where the range is in the order of only one-half the mean abundance. .
Notably, two of the most useful analyses of variation and its causes were from
these two populations (McFadden et al. 1967; Hunt 1974). Where good compari-
sons of both temporal and spatial variation could be made in the same stream
system (Sagehen Creek, Calfiornia and Lawrence Creek, Wisconsin), spatial
variation was the greater, by a significant margin. This, again, may not be
a surprising result, but is one with important implications for impact
studies. .
It appears that inferences about natural fluctuation in abundance and its
causes may best be found in detailed analyses of individual research studies,
including information on as many relevant environmental variables as possible.
Thus the bulk of this review is concerned with attempts to relate variation
in abundance to the environmental factors with which it may be associated.
3

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'TABLE 1.
/.
i.
LONG-TERM STUDIES 9F STREAM SALMONID POPULATIONS.

J
Location
Sashin Creek,
Alaska
Hooknose Creek,
British Columbia
Carnation Creek,
British Columbia
Minter Creek,
Washington
A1se.a River,
Oregon
Waddell Creek,
California
Sagehen Creek,
California
Lawrence Creek,
Wisconsin '.
Hunt Creek,
Michigan
Au Sable River,
Michigan
Hayes Brook,
Prince Edward
Island
Little Codroy
Ri ver ,
Newfoundland
Shelligan Burn,
Scotland
Species
pink salmon
coho salmon
pink salmon
chum salmon
coho salmon
cutthroat trout
stee1head trout
coho salmon
coho. salmon
cutthroat trout
coho salmon
stee1head trout
rainbow trout
brown trout
brook trout
brook trout
brook trout
. brook trout
brown trout
brook trout
Atlantic salmon
brook trout
Atlantic salmon
brown trout
I.i"
InclUSive Dates
II.
for Data Presented
. 1940-1959
1956-1968
1947-1956
1970-1977
(con tinuing)
1938-1953
1959-1973
1933-1944
1952-1961
1953-1970
1949-1962
1957-1967
1947-1960
1954-1963
1966-1975
Principal Reference
Merrell (1962)
Crone and Bond (1976)
Hunter (1959)
"Narv-er.and':Arraersen
(1974)
Sa10 and Bayliff (1958)
Moring and Lantz (1975)
Knight (1980)
Shapova1ov and Taft
(1954)
Gard and F1ittner
(1974)
Hunt (1974)
McFadden et al. (1967)
Alexander (1979)
Saunders and Smith
(1962)
Murray (1968)
Egg1ishaw and Shackley
(1977)
4

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SECTION 4
FACTORS AFFECTING NATURAL VARIABILITY
One approach to listing the important factors or variables in the stream
environment that can affect abundance of sa1monid populations is the following:
A.
Physical factors
1.
2.
Streamflow
Habitat quality
'B.
Biological factors
1.
2.
3.
Food abundance
Predation
Movement and migration
In most instances these variables may interact to influence a popula-
tion, and the classification is inevitably artificial. For example, habitat
preferences are often related to food availability. Under natural conditions,
it is often difficult to measure the effect of one factor independently.
However, the variables will be considered separately in this discussion, with
an attempt to show how interactions may be involved.
PHYSICAL FACTORS
Streamflow
, '
One of the earliest studies that attempted to relate streamflow to
sa1monid abundance was conducted by McKernan et al. (1950). They found that
low summer flows correlated with subsequent low returns of adult coho salmon
in the Si1etz River, Oregon from 1924 to 1945. No relation was apparent in
the Coquille River from 1923 to 1948. Scarnecchia (1978) found a significant
correlation (r = 0.68) between total streamflow in the 17-month period of
stream residence of juvenile coho and the commercial troll catch of adult
salmon 2 years later. These data came from five Oregon rivers from 1942 to
1962. In addition, there was a significant correlation (r = 0.56) between
total annual flow and catch 2 years later. Smoker (1955) obtained an even
higher correlation (r = 0.91) in the same analysis (total annual flow vs.
catch of adult coho 2 years later) for Puget Sound streams from 1935 to 1954.
In Cowichan Bay, B. C., a lower availability of coho to the sport fishery was
noted for year classes that experienced low summer streamf10ws in their
juvenile stages (Neave 1949). In Nile Creek, B. ~., from 1946 ,through 1949
5

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the output of coho smolts varied directly with the minimum mon/thly rai.nfall
during the previous summer (Wickett 1951). These studies sho~that stream-
flow ~uring some part of the. freshwater phase of coho life hi~tory can influ-
. ence J.ts level of abundance J.n the catch. ! '..;"
, . \~ ~ :t..\ .
We carried' out a s~milar analysis for juvenile coho salmon in two of the
streams that were part of the Alsea Watershed Study in Oregon.. Mean monthly
and seasonal discharge were correlated with mean June-April biomass and also
with the smolt count in the same period, from June 1960 through May 1968. In .
both streams the few significant correlations were mostly in the spring (Table
2) .
TABLE 2.
CORRELATIONS BETWEEN MEAN MONTHLY DISCHARGE AND ANNUAL SMOLT COUNT
AND MEAN. JUNE-APRIL BIOMASS FOR COHO SALMON, ALSEA WATERSHED STUDY,
JUNE 1960-MAY 1969. .
 Deer Cr. . Flynn Cr.
Period Smo1t Biomass Smo1t Biomass
June -0.'080 -0.080 -0.307 -0.321
July 0.095 -0.442 -0.004 -0.126
August 0.050 -0.199 -0.155. -0.267
September 0.098 -0.045 -0.206 -0.221
October 0.431 0.661 -0.122 0.145
November -0.076 -0.461 0.132 0.032
December 0.153 0.003 0.291 -0.373
January -0.398 -0.218 0.531 -0.654
February -0.099 . -0.006 0.042 0.088
March -0.687* -0.426 0.099 -0.055
April . 0.630 0.076 0.931** 0.936**
May 0.569 -0 . 162 0.714 * 0.694.*
June-May -0 . 350 -0 . 507 0.150 -0.364
Nov-Apr -0.482 -0.544 -0.209 -0 . 446
Jan-Apr -0.691* -0.448 -0.097 -0.279
Mar-Apr -0.344 -0.352 0.518 0.385
June-Sept -0.021 -0.125 -0.262 -0.230
* P <0.05    
** P <0.01    
  6  

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Knight (1980) used a longer series of data on smo1t abundance alone' and found
significant negative corre~ations between. mean January discharge and total
November-May smo1t count for Deer Creek and Flynn Creek (r = -0.64 and -0.65
. respectively) for the 1959-1960 through 1972-1973 seas~:ms" We performed a
similar analysis for cutthroat trout from September biomass data and mean
monthly discharge data (October 1961-September 1972) for all three streams
in the A1sea Watershed Study. 'Generally, 'corre1ations were' negative, but
nonsignificant, in the winter months 'in all three streams (Table 3).
TABLE 3. CORRELATIONS ~1WEEN MEAN MONTIfLY DISCHARGE AND MEAl'l SEPTEMBER 
  BIOMASS IN G/ FOR CUTTHROAT TROUT, ALSEA WATERSHED STUDY, 
  OCTOBER 1961-SEPTEMBER 1972.     
  . .      
  Deer Cr. Flynn Cr.   Needle Br.  
Month(s) All ye ars All years All years Pre-logging a Post-logging b
October -0.119 -0.014  -0.221 0.829  0.586 
November -0.133 0.085  -0.060 -0.393  0.150 
December 0.222 -0.174  -0.384 -0.382  0.292 
January -0.061 -0.139  -0.538 -0 . 804  -0.309 
February -0.136 -0.311  -0.269 0.645  0.686 
March 0.209 0.304  -0.302 0.646  -0.144 
April .0.571 0.399  -0.071 0.335  -0.256 
May 0.415 0.423  0.589 0.558  0.277 . 
June -0.212 -0.210  -0.095 0.993**  0.720 
July -0.092 0.094  -0.158' 0.335  0.465 
August -0.367 0.013  0.063 0.002  0 . 650 
September O. 097 0.117>  -0.262 0.264  0.718 
Oct-Sept 0.214 0.006  -0.565 -0 . 906  0.336 
Nov-Apr 0.195 -0.027  -0.574 -0.867  0.109 
Jan-Apr 0.186 0.058'  -0.711* -0.654  -0.085 
Mar-Apr 0.419 0.417  -0.249 0.669  -0 . 202 
June-Sept -00173 -0.100  -0.135 0.551  0.755 
 a 1962-1965       
 b 1967-1972 .       
 * P <0.05        
** P <0.01        
     7     

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In all three of 'these analyses the lack of consistency in the correla-
tions was notable. Although one can attach plausible explanations to the
statistically significant correlations, there were hardly more of them than
. might be expected due to chance in a series of that many analyses. Our
conclusion is that there is no solid basis for a relationship between stream-
flow and abundance of coho salmon and cutthroat trout in these streams, a
surprising result in the face of so much other evidence for such a relation.
The small size of the streams involved and the resultant low numbers of
juvenile fish may have reduced the power of the analysis, however.
In several western Oregon streams, Pearson et ale (1970) did a prelim-
inary short-term study of the effects of streamflow on juvenile coho salmon
during the summer low-flow season: From 1962 to 1965, thZy found a signifi-
cant positive relationship between coho density (number/m) and minimum
streamflow in McKay Creek, a tributary to the Tualatin River. They also
found a significant positive relationship between mean water velocity in
pools and coho density for 50 pools in fi ve ,streams of the Nehalem River
system. Preliminary data also indicated that streams with higher flows sup-
ported coho of larger sizes.
In streams in Maine, Havey and Davis (1970) found through multiple
regression analysis of several environmental variables that rainfall in July
and August, presumed to be an index of streamflow during the dry season, was
the single most important factor influencing survival of Atlantic salmon from
age 0+ to age 1+. Their multiple regression analysis was weakened, however,
by a small sample size.
Wickett (1958) reviewed the effects of low water levels on adult migra-
tion and egg deposition by pink and chum salmon in British Columbia streams.
Low flows result in excessive spawning density, leading to. superimposition
of redds and crowding of eggs. Adult migration is inhibited by low stream-
flow; other consequences include failure of egg deposition and increased
predation on spawning fish crowded in shallow water.
The effects of streamflow on survival of pink and chum salmon in spawning
beds were studied by McNeil (1966;1968) in streams in southeastern Alaska.
Below normal streamflow, both in summer and winter, caused significant
mortality of eggs and alevins in the gravel. In summer,. low streamflow acted
by causing low levels of dissolved oxygen inintragravel water. In winter,
low streamflow led to freezing of eggs and alevins, especially in streams
subject to greatly fluctuating flows. High streamflow during winter caused
mortality by displacement of eggs and alevins from spawning gravel.
Studies have. also been undertaken on the influence of streamflow on
resident populations of salmonids. In Big Roche-a-Cri Creek, Wisconsin, brook
trout biomass fluctuated greatly with streamflow. White (1975) found that
from 1958 through 1966, biomass was significantly correlated with mean
January-February discharge (r = 0.867). -
Using data from Sagehen Creek, California, kindly provided by Dr. Richard
Gard, we correlated mean monthly and seasonal discharge with mean annual bio-
mass of brook, brown, and rainbow trout (Table 4).: Brown trout biomass was
8

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best correlated with December flows ~dbrook trout biomass showed the best
correlation with February discharge. Neither of these was statistically
significant, however. Rainbow trout biomass showed significant negative
. correlations with discharge in January, 'April,. and June. Again, the total
number of significant correlations among the 51 comparisons is very close to
the number that would be expected by chance. However, the predominance of
negative correlations for brook and rainbow trout and positive correlations
for brown trout is in itself a significant result that deserves further
analysis.' .
TABLE 4.
CORRELATIONS BETWEEN MEAN M)NTHLY DISCHARGE AND MEAN ANNUAL
BIOMASS OF BROWN, BROOK-, AND RAINBOW TROUT. SAGE HEN CREEK,
CALIFORNIA, 1954-1961. DATA FROM DR. RICHARD GARD (PERSONAL
COMMUNICATION). .
Month(s) Brown Brook Rainbow
January 0.049 -0.142 -0.806*
February -0.174 -0.632 -0.495
March -0.168 0.072 -0.449
April 0.067 0.023 -0.746*
May 0.183 -0.488 -0.595
June 0.123 -0.547 -0. 727*
July 0.109 -0.494 -0. 649
Augus t 0.259 -0.408 -0. 592
September 0.354 -0.275 -0.468
October -0.416 -0.199 0.373
November -0.050 -0.209 -0.127
  , ,
December 0.514 0.146 0.147
Jan-Dec 0.208 -0.461 -0.294
Jan-June 0.137 -0.448 -0.305
March-June 0.151 -0.449 -0.339
March-April 0.164 -0.510 -0.454
July-Dee 0.426 -0.184 -0.080
* P < 0.05   
Floods can have a very severe impact on salmonid and other fish popula-
tions. Wickett (1958) reported that floods are a ',major cause of mortality in
9

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pink and chum salmon streams in British Columbia and have often reduced the
size of succeeding runs. The principal cause of mortality is scouring of
eggs and alevins from the gravel. In Nile Creek, B.C., chum salmon survival
, was considerably reduced in years of severe floods. In 1945-46, there were
no floods and the fry had a 3% survival rate. There were several severe
floods in 1946-47 and 1947-48, with survival rates dropping to 0.44% and
0.38%, respectively. ' There was high water but no severe flooding in 1948-49
and fry survival increased to 6.0% (Neave and Wickett 1953)~
In the Horokiwi stream, New Zealand, severe flooding occurred between
May and October 1941. Based on studies over the previous year, Allen (1951)
estimated the effects of these floods on the streambed, the benthic fauna,
and the brown trout population. The bottom fauna was reduced to 40-50% of '
levels of the previous year. The estimated number of most age classes present
in October 1941 was only 25-50% the number present of the same age in October
1940. Destruction of eggs by flooding represented 80-90%, compared to a
negligible loss the year before. The reduction in bottom fauna resulted 'in a
higher percentage of this food resource being required by the remaining
reduced trout population just for maintenance. This reduction, left a lower
proportion of the food for growth. Thus the floods caused a reduction in the
bottom fauna that limited the trout stock to a lower biomass and production.
This effect occurred independently of the direct reduction in numbers of
trout caused by the floods. Although the study terminated at that time, the
limitation was presumed to be only temporary, with both benthos and trout
populations returning to original levels in periods of normal rainfall.'
In Valley Creek, Minnesota, four severe floods were recorded in 1965
and 1966. Two year classes of brook trout were nearly eliminated from the
population. The older age groups were reduced as a result of changes in
habitat caused by flooding (Elwood and Waters 1969). A later study showed
that the brook trout population made a substantial recovery in 4-5 years.
S~anding.crop increased from 418.fish/ha in 1966 t02l?,882 fish/ha in 1969.
B~omass ~ncreasedfrom 2.5 g/m ~n 1966 to 14.8 g/m ~n 1970 (Hans02 and
Waters 1974),.still somewhat lower than the average of about 25 g/m from
1961 to 1965.
In Sagehen Creek, California, survival of spring-spawned rainbow trout
fry increased in years following winter floods (Seegrist and Gard 1972).
This increased survival of age-O rainbow trout was presumed to be caused by
reduced competition from young brook trout, a consequence of brook trout eggs
being destroyed by flooding. When floods occurred in May, rainbow trout eggs
were destroyed and survival of young brook trout was improved. Adult trout
were less affected by flooding than were the young.
These studies illustrate the impacts that floods can have on sa1monid
populations. Generally, they affect the eggs and young, older fish being
somewhat more resistant. The magnitude of the impact, however, can vary
according to the severity of the storm, the particular species, the time of
year, and the physical characteristics of the stream. '
10.

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Habitat Quality
Salmonids are not uniformly distributed within a stream reach. If
, habitat preference or use can be defined for a species, the potential exists
for prediction of spatial variation in abundance based on measurement o'f
habitat qualityo It may also be possibie to relate temporal variability in
. abundance to seasonal changes in habitat caused by changing streamflow or
other variables. A number of studies have attempted qualitative or quantita-
tive des~ription of habitat use by stream salmonids.
Juvenile coho salmon in their first summer prefer a pool environment.
Emerging fry in Waddell Creek,California, initially utilized shallow gravel
areas, particularly those near the stream margin (Shapovalov and Taft 1954).
'The youngest fry tended to school, but as the fish grew larger, these schools
broke up and individuals took up territories, which they defended. The larger
fry moved into deeper water and by July and August were mainly found in the
,deeper pools. Chapman (1962) further defined this territorial and aggressive
behavior and related it to habitat utilization. Ruggles (1966) found that
over twice as many fry remained in a pool-like environment than a riffle-like
condition in stream channels in British Columbia. In Oregon streams, Nickelson
and ~eisenbichler (1977) described characteristics of prime habitat for juve-
nile. coho salmon as having water depth of at least 30 cm, velocity of less
than 30 cm/sec, a cobble substrate, and cover consisting of undercut banks and
submerged roots.
In the South Fork system of the Salmon River, Idaho, juvenile summer
chinook salmon rear primarily in the main stem. Platts and Partridge (1978)
recently reported significant use of many tributaries as well. In these
tributaries the juvenile salmon preferred high quality pools iri the larger
streams that had lower channel gradients and grassy streambanks. Yet 59% of
all the salmon were found in stream reaches where less than 20% of the
channel consisted of pools. This distribution was presumably the result of .
the fact that most of the juvenile chinook in the tributaries occupied stream
reaches within 400 m of the main river, where there was naturally a low pool/
riffle ratio. . , '
Ina north central Colorado stream, Stewart (1970) sampled 41 sections
four times from June through September. He found mean depth and underwater,
overhanging rock cover to be the most important variables determining the
density of brook and rainbow trout larger than 18 em. Undercut banks and
areas of deep turbulent water seemed to be related to brook trout density,
but not that of rainbow trout. He also presents,useful data on spatial and
temporal variation in biomass of the 41 sections, along with the physical
data. Biomass of brook trout >18 em varied from 0 to 63.9 g/m, rainbow trout
o to 8103 g/m, and combined trout 0 to 117.5 g/meter of stream (data on area
not presented).
In Little PrickJy Pear Creek, Montana, Lewis (1969) conducted a similar
study involving 19 sections. He found that cover was the most important
factor determining the density of brown trout. Increased stream velocity was
associated with increases in density of both brown and rainbow trout per unit
area of pool surface and per unit area of cover. :The most stable trout
11

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populations occurred in deep. slow pools with extensive cover; brown trout
showed greater stability than rainbow trout. Current velocity was the most
important factor determining density of rainbow trout. Useful data on spatial
, variation in density are presented. but there is no information on biomass.
Use of habitat by steelhead trout was studied by Shapovalov and Taft
(1954) in Waddell Creek. California. Young fry showed similar tendencies to
coho fry. initially congregating in schools and later setting up territories.
However. unlike coho. steelhead fry inhabited riffles in late summer rather
than deep pools.
Dolly Varden fry in Hood Bay Creek. Alaska. were found in quiet water
near stream banks and in small pools. The fry were usually inactive and
found in or on the substrate. in contrast to the more aggressive coho fry.
often found in the same habitat. The coho were actively swimming and feeding
from the water surface (Blackett 1968).'
Species interaction can have a strong influence on habitat utilization.
Apparent preferences shown in the presence of a competing species may change
if that species is absent. or if another is present. so care must be used in
interpreting results from field studies of species interaction. Careful field
observation coupled with experimental analysis is needed to define these
interrelations. . .
Seasonal habitat preferences and behavior of juvenile coho salmon and
steelhead trout were studied by Hartman (1965) in British Columbia. In spring
and summer coho occupied pools and steelhead occupied riffles. Both were
aggressive in defending their respective habitats. This behavior is similar
to that observed in Waddell Creek, California. discussed earlier. In winter,
however, both species inhabited pools. Low population numbers, low aggres~
. siveness. and different microhabitat preferences were thought to be responsi-
ble for this coenstence.
Glova (1978) examined sympatric and allopatric populations of coho salmon
and cutthroat trout in six British Columbia streams. In each. three habitats
were defined in terms of stream veloci ty--pools « 8 cm/ sec), glides (8-20
cm/sec), and riffles (> 20 cm/sec). . In sampling during 1973 in Bush and
Holland creeks, where both species occurred, coho salmon dominated the salmonid
biomass in pools. composing 53-91% of the combined biomass. compared to 9-47%'
made up by trout. In riffles trout were dominant, making up 63-88% of the
combined total biomass. Glides were areas of intermediate biomass for both
species. although coho also tended to dominate here, with 52-81% of total
biomass. Above barrier falls. where they were found alone. cutthroat trout
utilized pools more so than riffles, possibly due to the absence of coho.
His analysis of diets suggested that coho were more specialized feed~rs.
relying mainly on drifting foods. whereas cutthroat were more generalized.
utilizing both drift and benthos. Glova (1978) noted that cutthroat emerged
much later in the ye~r than did coho salmon. into an environment that may
already be saturated by coho fry.: As a result of aggressive interaction with
young coho. the trout would be largely restricted. to riffle areas during
summer and early fall. and this habitat type is usually less abundant than
pools at this time. He concluded that production:of sympatric trout may be
12

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limited by interspecific interaction, although total fish production may be
greater..in multi-species streams.
Glova (1978) also found that pools were more extensively utilzed by the,
total fish species complex than were riffles. There was a strong negative
correlation (r = -0.92) between the biomass of all fish species combined
(coho salmon, cutthroat trout, and Coastrange sculpin, Cottus aleuticus) and
mean stream velocity in Holland Creek during September. Based on behavioral
studies he postulated that large pools would be less densely populated by ,
salmonids than small ones, owing to competition near the heads of pools for
incoming food and resultant low densities of fish in the downstream ends of
the larger pools. In support of this hypothesis, he found a significant
negative correlation (r = -0.40) between logarithms of salmonid biomass and
pool surface area, based on,data from a total of 37 pools in three streams.
In British Columbia, Bustard and Narver (1975b) found in experiments
that overwintering coho s.almon and cutthroat trout strongly preferred side-
pools with overhanging bank cover to those without such cover. Given a choice
between clean rubble substrate and silted rubble, they preferred the side-
pools with clean rubble. In a natural stream studied'during winter, age 1+
coho and steelhead were found mainly at greater depths and in deeper water'
than age 0 fish of either spec~es. As stream temperature dropped below 90 C,
coho and age 1+ steelhead occupied progressively de'eper water 'and both'
species moved closer to cover (gustard and Narver 1975a). Logs and upturned
roots were the most commonly used cover. Steelhead were more closely asso-
ciated with the substrate than were coho.
Habitat utilization by sympatric populations of coho and chinook salmon
fry was studied by Lister and Genoe (1970) in the Big Qualicum River, British
Columbia~ At emergence, fry of both species were found along stream margins
in association with streambank cover. As the young fish grew they moved into'
areas of, faster velocity. Spatial segregation soon occurred between the two
species because chinook fry emerged about one month earlier than coho fry and
grew at a faster rate. As a result chinook prefe.rred higher current veloc-
ities than did coho fry at a given date. .' Somewhat different results, invol v-
ing more overlap of distribution and more interspecific interaction, were
noted in an Oregon river where the two species emerged more nearly at the
same time (Stein et al. 1972). .
Diel variability in habitat preferences of juvenile chinook salmon and
steelhead trout in Idaho streams was shown by Edmundson et al. (1968). Both
species tended to move inshore at night to areas of quieter and shallower water
than those occupied during the day. Steelhead used areas of faster velocity
during the day than did chinook. Everest and Chapman (1972) found that most
age 0 steelhead trout and chinook salmon in two Idaho streams lived in water
velocities of less than 0.15 m/sec during summer. However, chinook occupied
areas of finer substrate and deeper water than did steelhead. There is little
interaction for living space between the two species because they spawn and
emerge at different times; steelhead spawn in spring and chinook spawn in
early fall. The larger juveniles tend to occupy deeper water, and the size
differences resulting from these different spawning periods thus reduce compe-
tition for food and space between the two species (Chapman and Bjornn 1969).
13

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Platts (1974) conducted an extensive study of fish habitats in 291 ,
sites in 38 streams within the upper South Fork of the Salmon River system,
Idaho. Geomorphic characteristics were an important determinant of popula-
, tion abundance. He found the highest fish population densities in channels
having 30-50% pools. Total density of the fish populations was positively
correlated with width and depth of the sampled streams. Rainbow trout and
chinook salmon dominated the populations. Rainbow trout were predominantly
found in riffles that were combined with shallow pools. Juvenile chinook
were found most abundant in high quality pools. '
In the Miramichi River, New Brunswick" Keenleyside (1962) studied
habitats and feeding behavior of Atlantic salmon and brook trout. Salmon
, fry were most abtmdant in the upper reaches, where rapids and riffles were
conunon. The fry were most abundant in fast water over substrate composed
of small gravel and stones. Salmon parr (1-4 years old) were also more
abundant in the upper sections of the river than the lower, but were found
in de~per water and over larger substrate. Brook trout were found only in
the upstream areas. Fry were most common in shallow slow-moving water
, along the margin. Older fish were found in deeper water that was often
swift or turbulent. Keenleyside (1962) noticed feeding segregation between
the species. Salmon fry and parr fed on benthic fauna and surface'organ-
isms, whereas trout fed almost exclusively on surface foods, possibly
because they held positions further above the substrate than the salmon.
In the Indalsalven River, Sweden, brown trout and Atlantic salmon
were found together (Lindroth 1955). The young trout (age 0+) occupied
shallow water near the stream margin. The trout were territorial and
aggressive, actively chasing salmon fry away from these areas. In Scottish
streams, Mills (1969) fotmd varying degrees of dominance between Atlantic
salmon and brown trout. In some streams he found salmon fry and parr and
juvenile trout living together in the same pools and riffles. All possible
combinations were noted, from predominance of trout in some streams through
to predominance of salmon in others. ' ,
Additional evidence that habitat quality is an important determinant
of salmonid biomass comes from efforts to improve the quality of existing
stream habitat. Although much of this work has gone unevaluated, a number
of careful studies have shown population response to habitat development.
Among the best documented is the work of Hunt (1971) at Lawrence Creek,
Wisconsin. Habitat development in one 0.7 km section of the stream in
1964 increased permanent bank cover by 416% and pool area by 289%. As a
result, total brook trout biomass increased from a mean of 59 kg in
1961-63 to 110 kg in 1965-67. In a follow-up study, Hunt (1976) fotmd the
mean tota~ biomass in 1968-70 to have increased even further, to 165 kg
(21.9 glm ).
One of the earliest studies on habitat development in the West was
conducted by Tarzwell. (1938) in two Arizona streams. In Horton Creek,
small log dams, deflectors, and artificial bank cover were added to one
section. A section of nearby Upper Tonto Creek was left unimproved as a
control. From 1932 to 1937, 25,150 brook, brown, and rainbow trout were
stocked in Horton Creek and 46,190 trout were stoc~ed in Upper Tonto Creek.
14

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A complete creel 'census was conducted in both streams. In 1936 and 1937,
following improvement in Horton Creek, that stream yielded more trout to the
angler ,and a greater weight of trout per hectare than did Upper Tonto Creek"
,in spite of the much heavier stocking of the latter stream.
The effects of cover manipulation on trout abundance were studied by
Boussu (1954) in Trout Creek, Montana. Four inventories were carried out
before alteration of habitat (June, December, March, and June); three inven-
tories were made after the alterations (September, December, and March).
Rainbow and brook trout comprised about 98% of the sa2monids, the remainder
being a few brown trout. 'Brush cover totaling 14.42m was added to four
sections of the stream having a total area of 263 m. Following the cover
addition, t2tal trout bi2mass in those sections increased from 1.13 kg to 4.04
kg (4.3 g/m to 15.4 g/m). Trout biomas2 in three un2ltered control 2 '
sections increased only 22% (from 8.5 g/m to 10.4g/m). When 11.9 m 2
of natu:al brush cover were removed from two sectionswit2 an area of 208 m ,
trout bl.Omass decreased from 3.83 kg to 2.28 kg (35.5 g/m to 21.1 g/m). 2
At the same2time trout biomass in a controL section increased26% (38.1 g/m
to 40.5 g/m ). The third treatment~nvo1ved removal of 1.4 m ' of undercut.
bank from two sectionstota2ing 80 m . 2In this case biomass decreased from'
0.68 kg to 0.45 kg (8.5 g/m 2to 5.6 g/m )2 while biomass in a control area
increased 20%, from 14.4 g/m to 17.3 g/m. In each of'the three treatments
the response by legal-sized fish (>18 cm) to change in cover was greater than
that of smaller fish. Another result of his work not explicitly presented was
the finding of a very significant spatial variation in trout biomass. In t2e
13 sections used for the study t2e pre-alteration biomass averaged 16.4 g/m ,
but ranged from 0.11 to 46.7 g/m. Because the data are reported as
averages for four sampling dates, actual variability was undoubtedly greater.
It should also be noted that these data resulted from a single pass with an
electroshocker through each section blocked with stop nets, rather than from a
formal population estimate.
Thirteen dams , 12 deflectors, and several covers were constructed in a
4ll-meter section of Hayes Brook, Prince Edward Island, in 1959. In the, .
following year, the number of age ° brook trout increased to 526 ,compared, .
with a 13-year pre-treatment mean of 482 (Saunders and Smith 1962). Numbers
of older trout increased from a mean of 348 (1947-1959) to 611 in 1960. .
Many of these studies have shown great variability in habitat preferences
between species, at different times of the year, for different ages of fish,
and in association with other species present. Knowledge of these preferences
is an important concern in the design of a sampling program. '
BIOLOGICAL FACTORS
Food Abundance
There has been an enormous amount of work done on food habits and feeding'
behavior of stream salmonids. However, very few of these studies bear directly
on the matter at issue here: can differences in abundance or availability of
food account for spatial or temporal variation in salmonid biomass in streams?
15

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The question is complicated by difficulty in defining an appropriate measure
of food availability--benthos, drift, or some combination. Very few studies
have focused on these important issues. A reorientation of feeding studies
. is required before a definitive answer to the question of food limitation is
possible. Our review will concentrat~ on the few studies relating food abun-
dance to variation in salmonid abundance. '
A starting point is to examine the significance of invertebrate drift.
Drifting invertebrates represent a potential food source of considerable magni-
tude, but of variable availability. Of particular importance is a strong diel '
periodicity, most drift occurring during darkness. Other factors that may
affect the rate of drift include water temperature, current velocity, stage of
the life cycle, and population density (Waters 1969). Some studies (Mason and
Chapman 1965; Elliot 1973; Gibson and Galbraith 1975) have shoWn greater fish
standing crops in stream sections with greater incoming drift. Yet other
studies have shown a significant part of the diet to be made up of non-drift
benthic forms. For example, Warren et ale (1964) reported the greatest food
consumption in stream sections with the least drift, possibly because of a
much greater abundance of benthic fauna in these sections. Other work has
shown little correlation between drift and diet. One such study was conducted
by Mundie (1969) on coho salmon fry in ,British Columbia. In seeking an explan-
ation for the lack of correlation he postulated diel and spatial variation in
drift composition, and variation in fry behavior. It is clear that there is
considerable variation in the degree to which drift is utilized as food by
, stream dwelling salmonids. "
There is evidence that food can be a limiting factor for some popula-
tions of stream salmonids. One of the strongest cases was brought forward by
Mason (1976). He found that food limited the stream production of juvenile
coho salmon during the summer in Sandy Creek, B.C. Through supplemental
feeding, the summer biomass was increased 6-7 fold compared with previous,
levels. However, there was no significant increase in the number of smolts
the following spring. The estimate of smolt yield under natural conditions,
was 212 fish, and the February population estimate was 257+71 fish surviving
from supplemental feeding the previous summer. Thus in thIs stream the winter
carrying capacity appeared to be the ultimate limit to smolt production.

, In the Horokiwi stream, New Zealand, Allen (1951) found evidence suggest-
ing that the food supply of brown trout, primarily the benthic fauna, could
play an important role in regulating the trout population. He found that an
increase in trout abundance increased pressure on the food supply, decreasing
the density of that supply. This resulted in a reduction in surplus food (the
amount that could be used for growth and production). Consequently, there was
a decrease ,in mean individual growth rate. This resulted in a feedback system
that would tend to keep the population biomass relatively constant by changing
growth rate in response to changes in population size. '
In a later review, Allen (1969) discussed the role of the benthic fauna
in regulating production of stream salmonids as a group. He suggests that
fish production can be limited by the density of the bottom fauna, which in
turn may be controlled by consumption by fish. This interaction provides a
mechanism for stabilizing the salmonid production rate.
16

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Ellis and Gowing (1957) examined bottom fauna and brown trout popula-
tions above and below a domestic sewage outfall into Houghton Creek, Michigan.
Although the biomasses of trout were similar above and below, there were'
. significant increases in the benthic fauna and condition of trout below the
point of sewage input., They also noted that trout below the outfall relied
less heavily on terrestrial foods, and concluded that trout growth was
strongly influenced by the quantity and kinds of food consumed.
Symons (1971) experimented with effects of fluctuating food quantities
on behavior and abundance of Atlantic salmon parr in a stream tank. He
found that such fluctuations had little effect on'the abundance of socially
dominant parr. Socially subdominant fish, however, seemed more abundant
where food was plentiful than where it was scarce. Thus total fish' abundance
was 'higher in channels where food was more abundant. Mason and Chapman (1965)
studied behavior and abundance of juvenile coho salmon in two experimental
stream channels. They found that one channel received about a third more.
volume in potential food organisms, 'and'this was associated with about a two-
thirds increase in total fish weight in that channel. However, there was no
replication, and other causes may also have been involved. .
Variation in food abundance was associated with spatial variation in
abundance of cutthroat trout populations in the Oregon Cascades. One pair
of open and shaded ,stream reaches was studied intensively for 4 years.
Primary production and insect emergence were significantly 'greater in the
open area compared to the forested section (Triska et ale 1980). Production,
growth rate, and biomass of cutthroat trout were about twice as great in the'
open area (Hall et al. 1978). Murphy (1979) expanded the study to include
nine pairs of open and forested sites, the openings being the results of
earlier clearcuts. He found the same general relations to hold, including
increased abundance of primary producers, predatory insects, and cutthroat
trout in the open areas. .
Predation
Although predation has been shown to cause some significant mortality
in stream salmonids (Hunter 1959; Mills 1964; Tagmaz'yan 1971), there have
been very few studies to support the position that variation in level of
predation leads to ultimate variation in size of the salmonid population.
One of the few studies to combine stream population studies with preda-
tor manipulation was carried out over a number of years in New Brunswick.
Elson (1962) reports investigations of predation on juvenile Atlantic salmon
by mergansers and kingfishers from 1942 to 1953. , In a sample of 117 mergan-
ser stomachs analyzed, an average of 42.1% of the number of items were salmon.
These salmon comprised an average of 10.3% of the total fish numbers in the
river, yielding a forage ratio of 4.1. Kingfishers also selectively fed on
salmon and had a forage ratio of 3.1. Predator control was practiced from
1947 through 1950 and. the abundanc~ of mergansers and kingfishers was signifi-
cantly reduced. Consumption of salmon by these two species of predators was
estimated to have been reduced to about 10% of pre~control levels. Before
control, smalt output ranged from approximately 1,000 to 5,000 each year.
During predator control, output ranged from 14,000 :to 24,000 smolts. Elson
17

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(1962) concluded that predation by mergansers waS a limiting factor on
Atlantic salmon smo1t production. Unforttmate1y, the study design was some-
what flawed by differing levels of stocking in the pre-control and control
'years, and by lack of data on adult returns.
One of the most detailed long-ferm studies on trout populations and
predation has-been carried out in Michigan on the North Branch of ' the Au Sable
River (Alexander 1979). Estimates of population size of brook and brown trout
were made in spring and fall each year from 1957 to 1967. Catch by anglers
was determined from a statistically designed creel census in two sections of
the river, one in which normal angling regulations prevailed and another in
w~ich angling was significantly restricted. Predators were collected for
stomach analysis from 1960 to 197~. From these analyses Alexander concluded
that the annual rate of mortality of both brook and brown trout was very high
(average rates calculated by cnapman-Robson method for age groups O-IV from
his data in Tables 2-5: brook - 0.84 and 0.82; bro\~ - 0.64 and 0.74, normal
and special regulations respectively). Consumption by known predators (prin-
cipally the American merganser, great blue heron, be 1 ted kingfisher, mink,
otter, and large brown trout) accotmted for a large fraction of this mortality;
their consumption was estimated between 43 and 46% of annual production.
Anglers'took another 37 and 8% in the normal and restricted water respectively.
Notwithstanding the sizeable mortality caused by predators, Alexander is of
the opinion that reduction of their abundance, short of complete removal of all
predators, would not, have a significant impact on salmonid abundance, owing to
, a compensatory kill rate that would be demonstrated by the remaining predators.
The fact that total annual mortality rates are similar for each age group in
the two sections, in the face' of much less angling "predation" in the special
water, supports this view. More effort must be put into well-designed stream
studies such as this one before a definitive conclusion on the significance of
predation to population abundance of stream dwelling salmonids can be provided.
Movement and Migration
Nearly all salmonid species undergo varying degrees of movement in their
lifetime. Some non-anadromous species undergo annual migrations within the
same stream system for the purpose of spawning. Others remain in the same
general area, undergoing local movements motivated by food, temperature, ,
streamflow, or other factors. Movement and migration can be considered a form
of temporal variability. The timing and magnitude of these movements need to
be understood in order to know what age and size range of fish to expect from
sampling ata particular time of the year. A comprehensive review of migra-
tory strategies of freshwater fishes and their significance to fish production
is provided by Northcote (1978).
Migrations of anadromous species are so conspicuous and generally well
known that it seems unnecessary to include them in this review. One caution-
ary example is perhaps in order, however. Conventional wisdom for many years
held that juvenile faU chinook salmon migrated to saltwater shortly after
emergence from the gravel, whereas juvenile spring chinook resided in fresh
water for a full year before migrating to the ocean. More recent studies
have indicated considerable variation from this pattern, both within and
between stocks of fall and spring chinook (Reimers:and Loeffel 1967; Reimers
18

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1973; Schlueter and Lichatowich 1977). These results indicate the impor-
tance of careful studies of the migratory pattern in each stock of fish.
Most studies of resident salmonids have found their movement to be
quite restricted, with the exception of some activity associated with.
spawning. In Kettle Creek, Pennsylvania, Watts et al. (1942) observed an
upstream migration of brook trout into. colder tributaries in late May and
early June. Spawning .took place in the fall, after which the trout moved
downstream once again. Resident trout in this watershed, however, moved
little between tributaries.
There have been several studies of the movements of resident brown
trout. Solomon and Templeton (1976) studied a population in a 7.5 km
section of a chalk stream in. England, from which they recognized five life
history stages with respect to movements and migration. The first was a
downstream movement from hatching to nursery areas. Fish stayed in these
areas for about 6.months. Then came a second movement further downstream
to areas of adult growth, where the trout remained until they were about 15
months old. Following this was .a period of very limited adult movement
. until maturation..Then came an upstream spawning migration followed by
downstream movements after spawning. .

In the Pine River, Michigan, Mense (1975} studied effects of varying
brown trout densities on movement. Among fish >15 em, he found no change in
movement patterns in a comparison of densities of 209 and 87 trout/hao He
does not przsent data on biomass, but we have made a rough estimate of 3.8
and 3.3 gfm , based on his data for the two respective years. Both
values for biomass are rather low, and the fact that the average size of
fish was much larger in the year of lower density reduced the power of his
test of the hypothesis.
In Convict Creek, California, Needham and Cramer (1943) found extensive
downstream movement of brown trout during spring. The peak coincided with
rising, but not maximum, streamflows, although flow was not felt to be a
causative factor. Most migrants were sexually immature. The downstream
migration may have been initiated by lack of adequate food and shelter in
the upper reaches of the stream. Little migration of rainbow or cutthroat
trout was noted.
Movement into and out of an intermittent tributary was shown to be "an
important feature in the life history of rainbow trout in Sagehen Creek,
California. From 39 to 47% of the spawning adults used this tributary from
1972 to 1975. Two possible reasons were given for this high use of an inter-
mittent stream while permanently flowing tributaries were. used by only a
small percentage of the spawning fish. Peak runoff from snow melt is much
greater and occurs earlier in the year in the intermittent tributary. In
addition, there is no competition from brook trout.because they cannot spawn
there in the fallowing to insufficient flow (Erman and Hawthorne 1976).
The rainbow trout fry from this tributary showed a .diel periodicity in down-
stream movement that differed between a dry and wet year. In 1973, the dry
year, fry moved downstream mainly during the day. In 1974, when the tribu-
tary retained permanent flow, fry migrated d~wnstream mainly at night. In
19

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that year many fry remained in the tributary throughout the summer (Erman
and Leidy,1975).
In another population of rainbow trout, movement was not extensive. In
Elder Creek, Oregon, Osborn (1967) made 755 observations of rainbow trout
movement, based on recaptures of marked fish larger than 75 mID. Less than 4%
of the fish had moved more than 91 m. '
Several studies have indicated that resident cutthroat trout undergo
relatively limited movements. In Gorge Creek, Alberta, Miller (1957) found
that of 58 tagged fish recaptured, 32 (55%) were recovered in the same pool
in which they were tagged. He concluded that most cutthroat in this stream
had a home territory less than 18 m long. In Lookout Creek, Oregon,
restricted'home ranges were also found for cutthroat trout. Wyatt (1959)
noticed no general downstream movement, but he did obs~rve two periods of
limited ,upstream movement. From October through January some trout made
$cattered vislts to tributaries. Then from the end of March to early June
there was a spawning migration, with a peak in April. .
OTHER FACTORS
There are a number of other factors that may affect natural variation
in abundance of salmonids. This section includes consideration of those
factors that are worthy of mention but have not been studied in enough detail
to warrant discussion in separate sections.
In the Pigeon River, Michigan, Benson (1953b) studied the effects of
ground water on brook and brown trout populations. Spawning of brook trout
occurred only in sections, with considerable ground water seepage. Brown
trout spawned in more widely scattered areas, but the greatest concentration
of redds was located where ground. water was abundant. In turn, these areas
of greater spawning produced higher population estimates. In a later study
in'the same river system, Latta (1969) found that numbers of brook trout fry
were directly correlated with ground water levels. He suggested that the
relation would be' stronger in lower reaches of streams than in headwaters.
Ice formation can have substantial effects on overwintering salmonid
populations in high mountain streams or high latitudes. In Sagehen Creek,
California, Needham and Jones (1959) noted that anchor ice, which forms under-
water in riffle areas, is an important ecological factor in that it can raise
the water level in pools and reduce streamflow over riffles. The breakup or
melting of anchor ice can dislodge the benthic fauna, making more food avail-
able to trout. In British Columbia, Bustard (1974) found collapsing snow and
subsurface ice to be two major causes of winter mortality in salmonids.
Beaver dams can significantly alter physical characteristics and carry-
ing capacity of salmonid streams. .In Sagehen Creek, California, the balance
of abundance of brook, brown, and rainbow trout was shifted by the presence
or absence of dams (Garq 1961).
20

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Chemical properties of stream water may influence salmonid abundance'and
growth rate. In New South Wales, Lake (1957) examined brown and rainbow trout
populations in 1~0 streams. He found a strong correlation between water
, chemistry and growth rate. Streams with the hardest water and highest pH had
the most abundant bottom fauna and produced trout with the greatest length at
a given age. Kennedy and Fitzmaurice (1971) examined over 40 streams arid
rivers in Ireland. They found the largest and fastest growing ,brown trout in
streams having a high calcium content. The smallest and slowest growing ones
were in lime deficient waters draining acid rocks. These results are supported
by Thomas (1964) from rivers in west Wales. He found that the growth rate of
brown trout in waters having a pH of 7 or more with a high ion and calcium
content was greater than that in more acid waters. Brown trout populations
in six streams of varying hardness- in Pennsylvania were sampled by McFadden
. (196la). There was no consistent difference in trout density between hard
and soft water streams, yet brown trout growth rate was consistently
greater in hard water streams. Fish of similar size had greater fecundity
in hard water. . . ,
Stream gradient usually operates to limit distribution of salmonids,
rather than abundance. However, in transitional areas" where two species
are involved, consideration of gradient may help to explain variation in
abundance. In the ClearWater Rive.r system" Idaho, Griffith (1972) f<:>und
evidence suggesting that stream gradient may influence the relative abundance
of brook and cutthroat trout in streams inhabited, by both species. In some
parts of Idaho cutthroat live in slow water. « 6 em/see) when not associated
with brook trout~ but they did not occupy this habitat in association with
brook trout in Crystal Creek. Brook trout were found in the low gradient
sections of Crystal Creek, whereas cutthroat were more abundant upstream iri
areas of higher gradient. The same distribution of the two species was also
found in a tributary of the St. Joe River. .
21

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SECTION 5
MINIMIZING THE EFFECTS OF VARIABILITY IN IMPACT STUDIES
The temporal and spatial variability in populations of stream salmonids
are clearly sufficient to mask very significant man-caused changes in these
populations. This is especially true for damage done by non-point source
pollutants. If we are to effectively monitor impacts of such perturbations,
means must be found to minimize the effects of natural variability in detect-
ing these effects.. It now seems. clear that the traditional watershed study
design, with its long-term pre-treatment calibration and post-treatment evalu-
ation, is not adequate for such analysis (Hall et al. 1978). After reviewing.
existing approaches to the problem, we present several interrelated ideas
that may improve sensitivity of future studies.
HABITAT QUALITY RATING SYSTEMS
. Models that quantitatively describe the qu~lity of salmonid habitat
promise to significantly reduce the amouht of unexplained va,riabili ty in
population abundance. The principal stimulus for the development of many of
these models has been concern about loss of water from streams caused by
irrigation or other appropriation. Hence the focus has been on determination
of minimum streamflow requirements and on ch~ges in habitat quality and .
quantity with changing streamflow. A good review of the historical basis for
this work is provided in proceedings of the Symposium on Instream Flow Needs
(Orsborn and Allman 1976).'. .
One of the early attempts to develop such a model was made by Wesche.
(1973), who combined hydrologic parameters, surface area, and available trout
cover to define available habitat for brown trout. Continuation of this
work extended the analysis to a cover rating system that provided a signifi-
cant linear predictor of brown trout biomass in several stream systems (Wesche,
1976) .
Another early study was initiated by the Oregon Department of Fish and
Wildlife, mentioned earlier. They began with an attempt to relate habitat
quantity and quality to streamflow by manipulating flow in a natural stream
channel through a diversion (Keeley and Nickelson 1974; Nickelson 1975).
Though initial work was marred by technical difficulties in establishing the
diversion, recent results have been quite promising. Two types of models are
presently being developed. One describes the relation between stream habitat
and rearing potential of salmonids during the low flow period. Another is
designed to predict the amount of habitat for an/ value of streamflow (Nickel-
son and Reisenbichler 1977; Nickelson and Hafele 1~78). Pool volume alone
22

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explained 93.5% of the variation in summer standing crop of juvenile coho
salmon in 12 sections of three coastal streams. For cutthroat trout a habitat'
quality rating (HQR) is computed as 'a product of a cover value, velocity
.preference factor, and wetted area. The cover value is a combination of
depth, escape cover, overhanging cover, turbulence, and velocity shelter.
Two alternative formulations of the HQR explained 91 and 87% of the variation
in cutthroat trout standing crop in 31 sections. of six streams' (Nickelson
and Hafele 1978). A related HQR for steelhead trout, involving cover, depth
and velocity, . and wetted area, explained 79% of the variation in standing.
crop of juveniles in 23 sections of four streams. Further work is underway
to validate these models. .
A related approach has been taken in a follow-up of work done in Wyoming
streams by Wesche (1976). Binns and Eiserman (1979) developed a habitat.
quality index for trout from analysis of 22 physical, chemical, and biological
at.tributes in a sample of 36 streams. Using a multiple regression approach
for'selection2of model attributes, they constructed an index (Model I) that
produced an R value of 0.955 for the .initial 20 streams sampled. When 2
this model was used to predict trout standing crop at 16 new stations, R
dropped to 0.594. A new model was developed for 'all 36 sites, based. on only.
nine habitat attributes, all physical and chemical (late summer flow, annual
flow variation, maximum stream temperature, and a food index and, a cover index
that are combinations of nitrate nitrogen, cover, eroding stream banks, sub-
state, water velocity, and stream width). This new model (Mo~l II) explained
97% of the variation in trout standing crop at the 36 sites (R = 0.~66).
However,. this analysis highlights a frequent misinterpretation of R as a
measure of reliability of the model (W.S. Overton, Department of Statistics,
Oregon State Univ., personal communication). One vrlue of standi~g crop is .
more than twice as large as the next largzst (63.4'V vs 28.4 g/m ). This
one point tends to inflate the value of R by its large contribution to the
sum of squares for standing crop. A more valid measure of the goodness of fit
is the relative predict~on error. The authors noted ~hat no prediction was in
error more than 5.5 g/m and that an error of 5.4 g/m at Sand Creek (the
highest trout population) was within 9% of the measured value. However, the
percent error at many stations with lower biomass was substantially higher
than that, and averaged 32.4% for the 36 stations with Model I (range 0-179%)
and 26.2% (range 0-157%) for Model II: Nonetheless, this approach is a very
useful one that promises to increase the precision of impact evaluation.
The most extensive development of indices to habitat quality has been
undertaken by the Cooperative Instream Flow Service Group of the U.S. Fish
and Wildlife Service (Bovee and Cochnauer 1977, Bovee 1978). Their general
approach has been to couple information on the state of several hydraulic
parameters of the stream environment with a "probability of use" for a com-
bination of these parameters. A weighted usable area is then calculated for
each level of discharge for the various life history stages of each species
1 It is noteworthy that this estimate appears to be one of the
ever reported for sa1monid biomass in streams, especially in
resulted from only a single pass through the study section.
largest
that it
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of interest. This effort, focused on the effects of,incremental losses of
streamflow on reduction in quality and quantity of. fish habitat, has been
substantially influenced by the thinking of physical scient:l.sts, primarily
. hydrologists. The input from physical scientists has been a significant
feature of the program and one that should be encouraged. Addition of the
perspective of geomorphology (Platts 1974; Swanson and Lienkaemper1978)
could significantly improve the generality of the approach.
PROCESS STUDIES
Unde~standing of the basic physical and biological processes that lead
to biological production and eventually to fish production will provide a
much sounder basis for assessment than has been available through the case
history approach. One particularly relevant example is found in the analysis
of temperature changes following logging in the Alsea Watershed Study (Brown
1967; Brown and Krygier 1970). By developing a model of the heating and cool-
ing process in an undisturbed stream and quantifying each element in the
energy budget, Brown was able to identify direct solar radiation as the pri-
mary source of warming in streams. This procedure allowed a prediction to be
made of the potential impact before timber was cut, and thereby provided a
basis for planning necessary buffer strips to minimize adverse effects caused
by warming of stream water. The process study provided an energy budget
approach that is general enough to be applied in most watersheds.
It is probably more feasible to carry out such studies of the physical
processes in streams than those of the biological components. Additional
work on physical process is now underway, for example, in suspended sediment
and bedload transport (Beschta 1978; Beschta and Jackson 1979). Nonetheless,
studies of biological process are essential to an understanding of variability
in stream salmonid populations, and further emphasis must be placed there.
Though far from complete, the work in Mack Creek carried out under the
Coniferous .Forest Biome Study and mentioned earlier (Triska et al. 1980)
provided some evidence of the validity of this approach. Knowledge of pri-
mary production, insect abundance, and trout production provided evidence that
the higher trout biomass in streams flowing through clearcut areas was a real
phenomenon rather than simply the result of movement of trout in response to
preference for open areas. It also provided some evidence of at least one
pathway through which the increase in trout production might have been
achieved. Much more work will be necessary in many more systems, however,
before models of biological processes will achieve the same level of under-
standing and predictability now enjoyed by models of physical processes in
streams. '.
. _._~-
STREAM CLASSIFICATION
It seems clear that some sort of classification of streams and their
watersheds will be an essential element of future impact assessment (W.S.
Overton, pers. corom.). Classification has had a long history, especially'in
Europe, where it has been incorporated in management schemes (cf. Huet 1959).
24

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However, the perspective of those involved in classification has often seemed
to focus on differences rather than similarities of stream ecosystems, thus
leading to unmanageable complexity in the system of classification (cf. Pennak
. 1971) . .
One of the approaches most adaptable to the present problem is that of
Platts (1974). His classification is based on stream order and a ,small 'number
of geomorphic characteristics and provides a manageable and quantifiable
system. Application to a stream ecosystem encompassing 220 km of the South
Fork of the Salmon River in Idaho provided significant explanation of vari-
ability in distribution and abundance of nine fish species.
A recent synthesis by Warren -(1979) forwards a more inclusive classifi-
cation scheme, based on a biogeoclimatic perspective. It takes the promising
approach of classification based.on capacity or potential of a system rather
than its present state. This potential would be indexed solely by geomorphic
characteristics of the stream habitat and the watershed system within which it
is imbedded. The scheme thus avoids much of the complexity inherent in meas-
uring both taxonomic and quantitative variability in biological components
within and between stream ecosystems. Further development of this concept
should provide a much more solid basis for impact assessment in the future.
IMPROVED STUDY DESIGN
Another source of improvement in efficiency of detection of impacts
appears available through modifications in the way in which observational
data are gathered. Field observations will probably always be the major
basis for impact assessment. As a consequence, 'much of the body of experi-
ence and theory in the field of experimental design will not be directly
applicable to such analysis. A sampling perspective is more appropriate,
and Overton (1978) provides a useful discussion of three levels at which
sampling questions can be addressed, along with general guidelines on study
design.
Eberhardt (1978) provides a valuable review of the problems of appraising
variability in population studies, one that should be required reading for
anyone beginning a study to assess impacts of non-point source pollution. A
related article (Eberhardt 1976) provides further detail, particularly on his
suggestions for handling the "single-site problem" that is often a character-
istic of impact assessment. He proposes substitution of repeated observa-
tions in time or space for true replication. The ratio of population density
in the affected area to that in the "control" site(s) would be the measure of
impact. He is cautious, indicating potential problems and suggesting the
whole approach as a "pseudodesign." Nonetheless, these two papers are a very
significant contribution to the topic under review here.
A number of different approaches to field observation are possible, and
appropriate combinations may lead to more fruitful results than will a single
approach. These possible approaches have been cl~ssified in two ways by Hall
et al. (1978). In a review of effects of watershed perturbations on streams,
they grouped studies according to whether they bracketed (before-after) or
2S

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followed (post-) treatment. The other level of classification was based on
whether detailed studies were made on one or very few streams (intensive)
compared to less detailed work on many streams, including a wide range of
. habitat types (extensive}. This two-level classification results in four
categories, which are evaluated for efficiency and sensitivity of impact
detection. An expanded listing.of advantages and disadvantages of each type
(Table 5) reveals that no one design is optimum. The extensive post- ,
treatment approach does have a number of advantages over the cl~ssical water-
shed study (intensive before~after). The best approach appears to be a .
combination of ,extensive post-treatment analysis with carefully designed
process studies carried out at one or more locations. '
Pairing of treatment and control is proposed to improve sensit~vity of
detection (Hall et al. 1978). This procedure places an upstream control
very close to a treatment area on each stream. It proved to be a sensitive
design to investigate changes in both predator populations and their habitat
in small clearcutsin the western Cascades in Oregon (Murphy 1979; Murphy and
Hall MS.)~ By inclusion of watersheds that had been harvested up to 35 years'
earlier, it also provided some insight into the rate of change of physical
and biological characteristics following treatment. This approach does have
the limitation that it can detect only those effects that occur in the immedi-
ate stream reach affected by the treatment. It is relatively insensitive to
downstream effects or those that accumulate over the larger watershed. '
A modification that would provide some in~ight into effects on that
scale would pair watersheds, treated and untreated. However, it would often
be difficult to find untreated watersheds adjacent to treated areas, and such
pairs would undoubtedly be more unlike than adjacent reaches of the same
stream. Nonetheless Welch et al. (1977) used a variation on this approach to
document effects of forestry and agriculture on streams in New Brunswick,
examining a total of 34 watersheds, all smaller than about 1000 ha.
~
Erman et al. (1977) used an innovative' form of this approach in a study
of effects of clearcutting on invertebrate populations in Northern California
streams. They sampled a total of 62 streams, all in small watersheds «800
ha). There were two objecives: to test effects of various widths of buffer
strips in preventing changes in invertebrate populations, and to examine
localized effects of point disturbances such as road-related landslides. For
the latter purpose their design was to sample upstream of the landslide as a
control, at the disturbance point, and downstream where no visual evidence of
the disturbance remained.
To evaluate the role of bufferstrips, they used a design that employed
two controls for each logged section, one upstream from the treatment and
another in an adjacent untreated watershed. The hypothesis tested was that
if effects occurred, the two control streams should be more similar than
either control and the treated section. Various measures of similarity were
compared and nonparam~tric ranking tests were used in the statistical anal-
ysis. They found significant effects on community composition in unbuffered
streams and found no significant differences between controls and streams
with wide bufferstrips (Newbold et al. in press).
26

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TABLE 5.
SUMMARY OF ADVANTAGES AND DISADVANTAGES OF THE FOUR MAJOR
APPROACHES TO WATERSHED STREAM ANALYSIS.
A.
Advantages
Intensive Before-After (10-15 years; 5-7 years before ,and after treatment).
Disadvantages
1) Possible to assess year-to-year
variation and place size of impact
in context of that variation.

2) Can assess short-term rate of -
recovery (ca. 5 years).
3) No assumptions required about
initial conditions.

4) Possible to monitor whole water-
shed impacts (provided substan-
tial investment in facilities
such, as flow and sediment
sampling wiers, fish traps).

, ' ,
5) Long time frame provides format
for extensive process s4udies.
1) No replication; results must be
viewed as a case study.

'2) Results not necessarily applicable'
elsewhere (areas of different soils,
geology, fish species, etc.)

3) Results vulnerable, to unusual
. climatic events (e.g. high or low
rainfall season(s) immediately
following treatment). '

4) Final results and management recom-
mendations require exceptionally
long time to formulate - up to 15
yrs after initial planning stage.

5) Difficult to maintain intensity of
investigation and continuity of
investigators over such a long
period..

6) Must rely on outside agencies or
firms to complete treatments as
scheduled - considerable coordina-
tion required. .
B.
'Advantages
Extensive Before-After (2-4 years; 1 year before treatment, 1 year after).
Disadvantages
1) Provides broader perspective
across geographical area than (A).

2) Larger number of streams examined
lessens danger of extreme case.

3) Increased generality of results
allows some extrapolation to
other areas.'
4) Relatively short time to achieve
results (3-4 years from planning
stage) .
1) Lack of long-term perspective--
little opportunity to observe
year-to-year variation.

2) Able to assess only immediate
results, which may not be repre-
sentative of longer time sequence.
3) Treatment vulnerable to unusual
weather (if all treatments in same
year) . .

4) Must rely on outside agency (see
(A) above).
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TABLE 5.
(Continued. )
C.
Intensive Post-Treatment
following treatment).

Advantages
(One Watershed--Paired Sites) (4~5 years,
Disadvantages
1) Shorter tlme for results than
(A) .

2) Moderate ability to assess
year-to~year variation.
3) Provides opportunity for
moderate level of effort on
process studies.
1) Provides no strict control--
requires assumption that upstream
control was identical to treated
area prior to. treatment.

2) "Control" most logically must be.
located upstream of treatment.
Strong downstream trend in any
feature would confound analysis.

3) Provides no spatial perspective--
results of limited' application
elsewhere.
(or more); all observations in
D. Extensive Post-Treatment, 10-30 Watersheds
1~2 years (vaFiable time' after treatment).

Advantages
Disadvantages
1) Wide spatial perspective allows
extrapolation to, other areas.

2) Long temporal perspective is
possible--can assess recovery
for as many years as past treat-
ments have occurred.

3) Provides ability to assess
interaction of physical setting
and treatment effects (e.g.
effects of sediment input at
different stream gradients).

4) Requires least time of all
four designs to get results--
as little as 2 years.

5) Probably most economical of
all four approaches per unit
of information.
1} No data available on pre-treatment
conditions--forces assumption that
control and treatment were identi~
cal (on average).

2) Control predominately upstream.

3) Total cost concentrated in very
short period--requires extensive
planning.

4) Not as effective as (A) in
assessing whole watershed effects.

5) Methods used in early treatments
may. not be comparable to later
ones.
---~~._- ~.- -
28

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Although much variability will undoubtedly remain in any study of
natural populations in field situations, the ideas discussed above should
help to resolve some of the uncertainty that has been present in past
. analyses. A good deal of ingenuity and insight will be needed in making
the right choices of habitat parameters and in devising methods of quanti-
fying them. Choosing the appropriate variables for watershed classification
will likewise be a formidable task. Hopefully, however, some judicious'
com9ination of these approaches should make the task of assessing and con-
trolling ,non-point source pollution a more effective and rational one.
29

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BIBLIOGRAPHY
As part of a development of strategy for analysis of impacts of non-point
source pollutants on streams, we have searched the literature on' population
dynamics of stream salmonids. The following list is not exhaustive, but we
have put emphasis on studies that ~eport quantitative estimates of population
size, and those that deal with spatial and temporal variability in such data.
In addition we have included studies that may be useful in interpreting this
variability. Some emphasis is placed on work on the west coast of North
America, but many valuable contributions from other areas have also been
included. The more important contributions in the following list are
annotated; other reports of interest are listed only by title.
Aho,
R.S. 1976. A population study of the cutthroat trout in
and shaded section of stream. M.S. thesis. Oregon State
Corvallis. 87 pp.
an unshaded
Univ. ,
Alexander, G.R. 1979. Predators of fish in coldwater streams. Pp. 153-170
in Henry Clepper Ced.), Predator-prey systems in fisheries management.
Int. Symp. on Predator-Prey Systems in Fish Communities and their Role
in Fisheries Management. Sport Fishing Institute, Washington, D.C. '
Allen, K.R. 1940. Studies on the biology of the early stages of the salmon,
(Salmo salar). 1. Growth in the River Eden. J. Anim. Ecol. 9:1-23. '
An investigation of the juvenile Atlantic Salmon population in the River'
Eden, England. Growth is considered in terms of length and weight. In
addition, data are presented on seasonal variability of the major'
benthic organisms, and a study is also made of those consumed by the
salmon.
Allen, K.R. 1951. The Horokiwi stream: a study of a trout population.
New Zealand Mar. Dept. Fish. Bull. No. 10. 231 pp.
A classic study of the brown trout and benthic fauna in a New Zealand
stream. Spatial variability in mean estimated trout biomass in six
sections of the stream is presented for 1940 and 1941~ Movements,
mortality, production, growth, length-weight relationships, and angling
effects are also analyzed. The bottom fauna is sampled in each section
and total numbers and weights are estimated. The availability of these
organisms in the-stream is compared to the consumption by the trout
and is expressed as forage ratios. The'role of the food supply as a pos-
sible limiting factor in trout production is discussed. The effects of
floods on the invertebrate fauna arid trout population are also examined.
30

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Allen, K.R. 1969. Limitations on production.m salmonid populations in
streams. pp. 3-18 in T.G. Northcote (ed.), Symposium on salmon and
trout in st~eams'; H:"'R. MacMillan Lectures in Fisheries.' Uni v.
British Columbia, Vancouver. .
A review of factors that can limit salmonid production in streams.
In. the egg stage in gravel, stream discharge and gravel permeability
are critical. Predation, territoriality, and food availability are
other stream factors of importance. The extent to which these factors
can be limiting depends on the species, the amount of.time spent in
fresh water, the stream's physical characteristics, and climatic
condi tions.
Andersen, B.C. 1978. Fish
Barkley So~d streams,
118 pp. .
populations of Carnation Creek and other
1975-1977. Fish~ Mar. Servo Data Rep. 89.
Andersen, B.C., and D.W. Narver. 1975. Fish populations of Carnation Creek
and other Barkley Sound streams 1974: data record and progress report.
Fish. Res. Bd. Can. MS Rep. 1351. 73 pp. .
~trong, R.H. 1970. Age, food, and migration of Dolly Varden smolts in
southeastern Alaska. . J. Fish. Res. Bd. Can. 27:991-1004.
Armstrong, R.H. 1974. Migration .of anadromous Dolly Varden (Salvelinus
malma) in southeastern Alaska.. J. Fish. Res. Bd. Can. 31:435-444.

Au, D.W.K. 1972. Population dynamics of the coho salmon and. its response
to logging in three coastal streams. Ph.D. thesis. Oregon State Univ.,
Corvallis. 245 pp.
An encompassing study of coho salmon in the Alsea Watershed Study~ in
which orie watershed was clearcut down to the streambank in 1966, another
watershed was clearcut but buffer strips were left, .and a third was left
unlogged as a control. Fish traps enabled total counts to be made of
upstream migrating adults and downstream migrating juveniles. Data on
mean monthly biomass of juveniles are presented for the 1963-1968 year
classes in each stream. Spawning migrations are analyzed in relation
to streamflow. Sex ratios, redd location, fecundity, redd survival,
and fry emergence are also discussed. Fry dispersal, colonization,
behavior, and migration are also investigated. Population estimates,
survival, growth, net production, and smolt yields are determined. No
significant long-term effects of logging were noted in these populations.
See also Chapman (1961; 1965) and Moring and Lantz (1975).
Becker, C.D. 1973. Food and growth parameters of juvenile chinook salmon,
Oncorhynchus tshawytscha, in central Columbia River. Fish. .Bull.
n : 387 -400.
Benson, N.G. 1953a. Seasonal fluctuations in the feeding of brook trout
in the Pigeon River, ~lichigan. Trans. Amer. Fish. Soc. 83:76-83.
31

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Benson, N.G. 1953b. The importance of ground water to, trout populations in
the Pigeon River, Michigan. Trans. N. Amer. Wildl. Conf. l8:26~-28l.
. Benson, N.G. 1960. Factors influencing production of immature cutthroat
trout in Arnica Creek, Yellowstone Park. Trans. Amer. Fish. Soc.
89:168-175.
Presents data on the number of cutthroat trout in age groups 0, I,
and II from 1950 through 1958. Production of these year classes
is also investigated. Factors that may be related to production,
include migration down into Yellowstone Lake, streamflow, photoperiod,
predation, number of spawners, and timing of the spawning run. '
Beschta, R.L. 1978. Long-term patterns of sediment production following
road construction and logging in the Oregon Coast Range. Water
Resource Res. 14:1011-1016.
Beschta, R.L., and W.L; Jackson.. 1979. The intrusion of fine sediments
into a stable gravel bed. J. Fish. Res. Bd. Can; 36:204-210.
Binns, N.A., and F.M. Eiserman. 1979. Quantification of fluvial trout
, habitat in Wyoming. Trans. Am. Fish. Soc. 108:215-228.
Bjornn, T.C. ,1965. The production of juvenile stee1head trout in an
Idaho stream. ' Proc. 45th Ann. Conf. West. Assoc. State Game Fish
Commrs. 45:210-216.
Bjornn, T.C. 1968. Survival and emergence of trout and salmon frY in
various gravel-sand mixtures. pp. 80-88 in Proc. Forum on the relation
between logging and salmon. Feb. 8-9, 1968. Alaska Distr. Amer. Inst.
Fish. Res. BioI., Juneau.
Bjornn, T.C. 1971. Trout and salmon movements in two Idaho streams as
related to temperature, food, streamflow, cover and population
density. Trans. Amer. Fish. Soc. 100:423-438.
Bjornn, T.C. 1978. Survival, production, and yield of trout and chinook
salmon in the Lemhi River, Idaho. Univ. Idaho ColI. For., Wild1.,
Range Sci. Bull. No. 27. 57 pp.
A long-term study of chinook salmon, stee1head trout, and resident
trout from 1962 through 1975. Stocking the river with different
densities of stee1head fry and subsequent yields of subyear1ings
are discussed. Survival, growth, mortality, and predation are
examined. The effects of introducing anadromous fish on resident
trout and interactions between stee1head trout and chinook salmon
are also investigated.
Blackett, R.F. 1968. Spawning behavior, fecundity, and early life
history of anadrornous Dolly Varden Sa1velinus'ma1ma (Walbaum) in
southeastern Alaska. Alaska Dept. Fish Game Res. Rep. No.6. 85 pp.
32

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Bohlin, T. 1977. Habitat selection and intercohort competition of
juvenile sea-trout Salmo trutta. Oikos 29:112-117.
A study of juvenile brown trout in a small Swedish stream from 1971
through 1975. Seven study sections were established to determine
densities of ages 0+ and I+ trout in relation to water depth and
substrate type. Differences in trout lengths in the various habitats
are also investigated. Competition between year classes is shown in
stream tank experiments. Additional.field studies show biennial
fluctuations in densities of the two groups. The role of intercohort
competition as a possible cause of these variations is discussed.
Boussu, M.F. 1954.
small stream.
- .
Relationship between trout populations and cover on a
J. Wildl. Mgmt. 18:229-239.
Bovee, K.D. 1978. Probability-of-use criteria for the family Salmonidae.
Instream Flow Info. Pap. 4. FWSjOBS-78j07. 80 pp. .
. Bovee, K.D., and T. Cochnauer. ~977. Development and evaluation of
weight~d criteria; probabi1ity-of-use curves for instream flow
assessments: fisheries. Instream Flow Info. Pap. 3. FWSjOBS-
77j63. 39 pp.
Brown, G.W. 1967. Temperature prediction using energy budget techniques
on small mountain streams. Ph.D. thesis. Oregon State Univ.,
Corvallis. 120 pp.
Brown, G.W., and J.T. Krygier. 1970.
temperature. Water Resource Res.
Effect of clearcutting on stream
6: 1133-1139.
Burnet, A.M.R. 1959. Some observations on natural fluctuations of trout
populations numbers. New Zealand J. Sci. 2 :410-42l.

Burns, J.W. 1970. Spawning bed sedimentation studies in northern California
streams. Calif. Fish Game 56:253-270.
1 )
Burns, J.W. 1971. The carrying capacity for juvenile salmonids in some
northern California streams. Calif. Fish Game 57:44-57.
Presents 3 years of data on salmonid biomass (1967-196Q) from seven
northern California streams. An attempt is made to define the
natural carrying capacity of these streams. Biomass per unit of
surface area seems to be the best method of expressing this
capacity. The use of streambed sediments, total dissolved solids,
alkalinity, and total phosphate in predicting carrying capacity
is also discussed. Changes in annual carrying capacity up to 50%
are attributed to natural variation.
Bustard, D. 1974. Some possible effects of logging on wintering juvenile
salmonids. In Stream ecology: a symposium, Parksvi11e, B.C. Assoc.
British Columbia Prof. Foresters. 19 pp. .
33

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Bustard, D.R., and DoW. Narver. 1975a.. Aspects of the winter ecology of
juvenile. coho salmon (Oncorhynchus kisutch) and steelhead trout
(Salmo gairdneri). J. Fish. Res. Bd. Can. 32:667-680.
Bustard, D.Ro, and D.W. Narver. 1975bo Preferences of juvenile coho
. salmon (Oncorhynchus kisutch) and cutthroat trout (Salmo clarki)
relative to simulated alteration of winter habitat. J. Fish. Res.
Bd. Can. 32:681-687.
Chapman, DoW. 1957. Studies on age, growth, and migration of steelhead
trout, Salmo gairdneri gairdneri, in the Alsea River, Oregon. M.S.
thesis. Oregon State Univ., Corvallis. 96 pp.
Chapman, D.W. 1961. Factors determining production of coho salmon,
Oncorhynchus kisutch, in three Oregon streams. Ph.D. thesis.
Oregon State Univ., Corvallis. 214 pp.
A comprehensive study of coho salmon production and ecology in the
three streams of the AlseaWatershed Study from 1958 to 1961. Physical
factors investigated include streamflow, rainfall, water temperature,
suspended sediment loads) light incidence, water chemistry, and stream
area measurements. Biomass, yield, and growth rates of juvenile coho
salmon are examined in determining production estimates. Behavior
patterns of fry are also studied, including dominance, hierarchy
formation, hiding, territorial defense, and aggression. Food habits
of the fry are also analyzed. In addition, the aquatic insect food
habits are examined to determine th~ proportion of coho food that
comes either directly or indirectly from allochthonous sources.
Chapman, D.W,. 1962. Aggressive behavior in juvenile coho salmon as
a cause of emigration. J. Fish. Res. Bd. Can. 19:1047-1080.
Chapman, D. IV. 1965.
Oregon streams. 0
Net production of juvenile coho salmon in three
Trans. Amer. Fish. Soc. 94:40-52.
,i Presents data on mean monthly biomass, production, and instantaneous
growth rate of the 1959-1962 year classes in all three streams of
the Alsea Watershed Study. The smolt yield for these years is also
presented. Relatively large freshets seem to increase downstream
movements of juveniles. Downstream drift of fry and smoltmigrations
are discussed as possible sources of bias in estimates of growth and
production in the residual populations.
Chapman, D.W. 1966. Food and space as regulators of salmonid populations
. in streams. Amer. Nat. 100:345-357.
An important synthesis emphasizing the importance of social behavior
in population regulation of stream salmonids. No new data are
included, but many earlier studies are reviewed. The author makes
the case for a food-space interaction mediated through social behavior
as the important mechanism controlling population size in streams.
34

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Chapman, D.W., and T.C. Bjornn. 1969. Distribution of salmonids in
streams, with special reference to food and feeding. pp. 153-176
in T.G. Northcote (ed.), Symposium on salmon and trout in streams;
H.R. MacMillan Lectures in Fisheries. Univ. British Columbia,
Vancouver.
Chapman, WoM., and E. Quistorff. 1938. The food of certain fishes of
north central Columbia River drainage, in particular, young chinook
salmon and steelhead trout. Wash. Dept. Fish. BioI. Rep. No.. 37A.
14 pp.
Clary, JoR. 1970. Predation on the brown trout (Salmo trutta) by the
slimy sculpin (Cottus cognatus). Ph.D. thesis. Pennsylvania State
Univ., University Park. 50 pp.
Cordone, A.J., and. D.W. Kelley. 1961. .The influences of i~organic
sediment on the aquatic life of streams. Calif. Fish Game 47:189-228.
Creaser, C.W. 1930. Relative importance of hydrogen-ion concentration,
. temperature~ dissolved oxygen, and carbon-dioxide tension, on
habitat selection by brook-trOut~ Ecology' 11:246-262.

Crisp, D.T., R.HoK~ Mann, and J.C. McCormack; 1974. The populations of
fish at Cow Green, Upper Teesdale, before impoundment. J. Appl. Ecol.
11:969-996. . . .
A study of fish populations in several tributaries and the main stem
of the upper River Tees, England from August 1967 through May 1970.
Species studied are brown trout, bullhead (Cottus), and minnow. Data
are presented on physical and chemical characteristics of each stream
study site. Biological parameters examined for each species include
distribution, length, weight, density, biomass, growth, and
production. These parameters are also considered by age group for
trout and bullheads. Initial effects of dam construction on these
parameters are also investigated.
Crisp, D~T., R~H.K. Mann, and J. C. McCormack. 1975. The populations
of fish in the River Tees system on. the Moor House National Nature
Reserve, Westmorland. J. Fish BioI. 7:573-593.
A study of brown trout and bullhead (Cottus) populations in tributaries
and the main stem of the River Tees system, England. Mean density and
biomass from 1968 through 1972 are presented for May, August, and
October for five tributary sites. Growth rates, survival, production,
and recruitment are also investigated.
Crone, R.A. 1968. Behavior and survival of coho salmon, Oncorhynchus
kisutch (Walba~), in Sashin Creek, southeastern Alaska. M.S. thesis.
Oregon State Univ., Corvallis. 79 pp.
35

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A study of 'coho salmon from 1963 to 1966. Spawning adult
investigations include number, timing, and age of returning migrants,
distribution on the spawning grounds, effects on pink salmon~ redd
life, fecundities, and egg retention. Juvenile investigations
include population size, distribution in the stream, age composition,
food habits, mortality, and timing and size of smolt migrations.

Crone, R.A., and C.E. Bond. 1976. Life history of coho salmon, Oncorhynchus
, kisutch, in Sashin Creek, southeastern Alaska. Fish. Bull. 74:897-923.
Presents data on coho fry and smolt weir counts from 1956 through 1968.
Number of adults, ti~ng of migration, age composition, fecundity, and
spawner distribution and density are examined. Interspecific
competition with pink salmon is also investigated. Freshwater survival,
growth, mortality, and size and age of smolts are analyzed. In
addition, the significance of fry migration and early salinity
tolerance is discussed. '
Cummins, K.W. 1974. Structure and function of stream ecosystems.
Bioscience 24:631-641.
, ,
Davidson, F.A., and S.J. Hutchinson. 1942. Natural reproduction of
pink salmon studied at Little Port Walter, Alaska. Ecology
23:234-235.
Dodge, D.P., and H.R. MacCrimmon. 1971. Environmental influences on
extended spawning of rainbow trout (Salmo gairdneri). Trans. '
Amer.Fish. Soc. 100:312-318.
Drucker, B. 1972., Some life history characteristics of coho salmon of
the Karluk River System, Kodiak Island, Alaska. Fish. Bull.
70: 79-94.
Eberhardt, L.L. 1976. Quantitative ecology and impact assessment.
Environ. Mgmt. 4:27-70.
J.
Eberhardt, L.L. 1978. Appraising variability in population studies.
Wildl. Mgmt. 42:207-238.
J.
Edmundson, E., F.E. Everest, and D.W. Chapman. 1968. Permanence of
station in juvenile chinook salmon and steelhead trout. J. Fish.
Res. Bd. Can. 25:1453-1464.
Egglishaw, H.J.
Scotland.
1970. Production of salmon and trout in a stream in
J. Fish BioI. 2:117-136.
A study of Atlantic salmon and brown trout populations in Shelligan
Burn, Scotland from 1966 through 1968. Population size, growth
rates, biomass, production, and yield data are presented. Variability
in production is ~hown in three stream sections for the 1963-1968 year
classes. Contributions of each year class to the total production for
both species are discussed. .
36

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Egglishaw, H.J.,' and P.E. Shackley. 1977.. Growth, ~urvival, and production
of juvenile salmon and trout in a Scottish stream, 1966-1975. J. Fish
BioI. 11:647~672.
A long-term study (1966-1975) of Atlantic salmon and brown trout.
populations in Shelligan Burn, ~cotland. Mean biomass of each age
group and total biomass of each species every year for the 10-year
period are presented. Seasonal variability in growth rates, mean
size and weight, and population density are investigated. Variability
within each cohort through its lifetime is also discussed.
Elliott, J.M. 1973. The food of brown and rainbow trout (Salmo trutta
and S. gairdneri) in relation to the abundance of drifting invertebrates
in a-mountain stream. Oecologia 12:329-347.
Elliott, J.M. 1976.
Salmo trutta L.
The downstream drifti~g of eggs of brown trout,
J. Fish BioI. 9:45~50.
Ellis, R.J., andH. Gowing. 1957. Relationship betweert food supply
and condition of wild brown trout, Salmo trutta Linnaeus, in a
Michigan stream. Lirnnol. Oceanogr.---Z:Z99-308. .
Elser, A.A. 1968. Fish populations of a trout stream in relation to.
o major habitat zones and channel alterations. Trans. Amer. Fish.
Soc. 97:389-397.
A study of brown, rainbow, and brook trout populations in Little
Prickly Pear Creek, Montana, during the summers of 1965 and 1966.
Spatial variability is shown through" data on biomass in 11 sections
of the stream. Four of these sections have been altered due to
adjacent railroad and highway construction. Major changes include
straightening 'of the channel, loss of streambank cover, and changes
in pool-riffle ratios. Comparisons are made between altered and
unaltered sections in the same habitat zones in terms of physical
stream characteristics and trout abundance.
Elson, P.F. 1957. The importance of size in the change from parr to
smolt in Atlantic salmon. Can. Fish Cult. 21:1-6.
Elson, P.F. 1962. Predator-prey relationships between fish-eating
birds and Atlantic salmon. Fish. Res. Bd. Can. Bull. 133. 87 pp.
A study of relationships between mergansers, kingfishers, and
juvenile Atlantic salmon in the Pollett River, New Brunswick, from
1942 to 1953. Control of the predators was applied .from 1947 to 1950,
reducing pressure on the salmon by approximately 90%. Survival of
planted underyearlings in years before and during control is examined.
Data are presented on number of fish planted, number of parr one year
later, and smolt output for each year of the ,study. The role of these
predators as limiting factors for smolt production is discussed.
Methods of predator control are also investigated. The optimum
levels of each predator species for this river. section are determined.
37

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Elson, P.F. 1975. Atlantic salmon rivers, smolt production and optimal
spawning: an overview of natural production. pp. 96-119 in J.R.
Bohne ,and L. Sochasky (eds.), New England Atlantic salmon-restoration
conference. Int. Atlantic Salmon Spec. Publ. Sere No.6.
Elwood, J.W., and T.F. Waters. 1969. Effects of floods on food consUmption
and production rates of a stream brook trout population. Trans. Amer.
Fish. Soc. 98:253-262.
Erman, D.C., and V.M. Hawthorne. 1976. The quantitative importance of
an intermittent stream in the spawning of rainbow trout. Trans.
Amer. Fish. Soc. 105:675-681.
Erman, D.C., and G.R. Leidy. 1975. Downstream movement of rainbow trout
fry in a tributary of Sagehen Creek, under permanent and intermittent
flow. Trans. Amer. Fish. Soc. 104:467-473. '
Erman, D.C., J.D. Newbold, and K.B. Roby. 1977.' Evaluation of
streamside buffers trips for protecting aquatic organisms.
165. Calif. Water'Resource Center, Davis. 48 pp.
Contr. No.
, ,
Everest, F.H. 1973. Ecology and management of summer steelhead in the
Rogue River. Ore. State Game Comm. Fish. Res. Rep. No.7. 48 pp.
Everest, F.H., and D.W. Chapman. 1972. Habitat selection and spatial
interaction by juvenile chinook salmon and steelhead trout in two
Idaho streams. J. Fish. Res. Bd. Can. 29:91-100.' ,
Foerster, R.E., and W.E. Ricker. 1941. The effect of reduction of
, predaceous fish on survival of young sockeye salmon at Cultus
Lake. J. Fish. Res. Bd. Can. 5:315-336.
Foerster, R.E., and W.E. Ricker. 1953. The coho salmon of Cultus Lake
and SweItzer Creek.J. Fish. Res. Bd. Can. 10:293-319.
Gard, R. 1961. Effects of beaver on trout in Sagehen Creek, California.
J. Wildl. Mgmt. 25:221-242.
Gard, R.,and G.A. Flittner. 1974.
in Sagehen Creek, California.
Distribution and abundance of fishes
J. Wildl. Mgmt. 38:347-358.
A 10-year study of salmonid and other fish populations, from 1952
through 1961. Data on average biomass over the entire period in
11 sections of the stream for brook, brown, and rainbow trout show
spatial variability and relative distributions of the species.
Temporal variability in abundance is also shown. The effects of
stream gradient, water temperature, elevation, floods, and beavers
on these two kinds of variability are examined.
38

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Gard, R., and D.W. Seegrist. 1972. Abtn1dance' and harvest of trout in
Sagehen Creek, California. 'Trans. Amer. Fish Soc. 101:463-477.,
'Gibbons, D.R., and E.O. Salo. 1973. An annotated bibliography of the
effects of logging on fish of the western United States and Canada.
U.S.D.A. For. Servo Gen. Tech. Rep. PNW-10. 145 pp.
Gibson, R.J. 1973. The interrelationships of brook trout, Salvelinus
fontinalis (Mi tchill), and juvenile Atlantic salmon, Salmo salar
L. Ph.D. thesis. Univ. Waterloo, Ontario.
Gibson, R.J., and D. Galbraith. 1975. The relationships between invertebrate
drift and salmonid populations in the Matamek River, Quebec, below a
lake. Trans. Amer. Fish. Soc. 104:529-535.
Gibson, R.J., and G. Power.' 1975. Selection by brook trout (Salvelinus
fontinalis) and juvenile Atlantic salmon (Salmo salar) of shade'
related to water depth. J. Fish. Res ~ Bd. Can. 32:1652-1656.
Giger, R.D.
Oregon.
1972. Ecology and management of coastal cutthroat trout in
Ore. State Game Comm. Fish. Res., Rep. No.6. 61 pp.
Glova, G.J. 1978~ Pattern and mechanism of resource p~rtitioning
between stream populations of juvenile coho salmon (Oncorhynchus
'kisutch) and coastal cutthroat trout (Salmo clarki clarki). Ph.D.
thesis. Univ. British Columbia; Vancouver. 185 pp.
An investigation of sympatric and allopatric populations 'of coho
salmon and cutthroat trout in six British Columbia streams from 1973
to 1976. Data are presented on biomass and fish length in pool, glide,
and riffle habitats in each stream; Relationships between biomass and
streamflow,pool area, and volume are examined. Spatial variability in
salmonid density and biomass in three study sections in each of two
streams is shown. Interactions between the two species are discussed
in terms of abundances in different habitats. The importance of drift
organisms in the diet of each species is investigated. Density and
biomass of the sculpin (Cottus aleuticus) are estimated, and. its'
relationships to the salmonids are studied.
Griffith, J.S., Jr. 1972. Comparative behavior and habitat utilization
of brook trout (Salvelinus fontinalis) and cutthroat trout (Salmo
clarki) in small streams in northern Idaho. J. Fish. Res. Bd. Can.
29: 265-273.
Griffith, J.S., Jr. 1974. Utilization of invertebrate drift by brook
trout (Salvelinus fontinalis) and cutthroat trout (Salmo clarki) in
small streams in Idaho. Trans. Amer. Fish. Soc. 103:440-447.
Hall, J.D., M.L. Murphy, and R.S. Aho. 1978. An improved design for
assessing impacts of watershed practices on small streams. Verh.
Int. Ver. Limnol. 20:1359-1365.
39
. I

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Four approaches to studying impacts of watershed management practices
on streams are discussed. The long-term study is an intensive
approach involving pre-treatment and post-treatment work on sites
in the same stream. The extensive before-after approach is a short-
term study involving sites in several streams. The intensive post-
treatment analysis compares two adjacent sections of the same stream,
one in a previously treated area and the other just upstream, over
several years. The last technique is extensive post-treatment
analysis, a short-term study involving comparisons between adjacent
logged and forested stream sections in many watersheds. Advantages
and disadvantages of each approach are discussed, as well as
recommendations from statisti~al, practical, and economic viewpoints.
Hallam, J.C. 1959. Habitat and associated fauna of four species of
fish in ontario streams. J. Fish. Res. Bd. Can. 16:147-173.
Hanson, D.L., and T.F. Waters. 1974. Recovery of standing crop and
. production rate of a brook trout population in a flood-damaged
stream. Trans. Amer. Fish. Soc. 103:431-439.
Hanzel, D.A. 1960. The distribution of the cutthroat trout (Salmo
. clarki) in Montana. Proc.Montana Acad. Sci. 19:32-71.-----
Hartman, G.F. 1965. The role of behavior in the ecology and interaction
of underyearling coho salmon (Oncorhynchus kisutch) and steelhead
trout (Salmo gairdneri). J. Fish. Res. Bd. Can. 22: 1035-1081.

Hartman, G.F., and C.A. Gill. 1968. Distributions of juvenile steelhead
and cutthroat trout (Salmo gairdneri and~. clarki clarki) within
streams in southwestern British Columbia. J. Fish. Res. Bd. Can.
25:33-48.' .
Havey, K.A., and R.M. Davis. 1970. Factors influencing standing crops
and survival of juvenile salmon at Barrows Stream, Maine. Trans.
Amer. Fish. Soc. 99:297-311.
Presents data on biomass of Atlantic salmon in this stream from 1960
through 1965. Biological and physical variables that could influence
biomass and survival are studied. These factors include rainfall,
streamflow, air temperature, water temperature, numerical standing
crop, and biomass of other fish species in the stream. Regression
analyses between these parameters and survival and biomass of age 0+
and 1+ salmon are performed to determine those of greatest significance.
Hazzard, A.S., and M.J. Madsen. 1933. Studies of the food of the cutthroat
trout. Trans. Amer. Fish. Soc. 63:198-207.
Heiser, D.W. 1966. Age and growth of anadromous Dolly Varden char
Salvelinus malrna (Walbaurn) in Eva Creek, Baranof Island, southeastern
Alaska. Alaska Dept. Fish Game. Res. Rep. No.5. 29 pp.
40

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Hildebrand, S.G. 1971. The effect of coho spawning on the benthic
invertebrates of the Platte River, Benzie County, Michigan. Trans.
Amer. Fish. Soc. 100:61-68.
Hoar, W.S. 1951. The behavior of chum, pink and coho salmon in relation
to ~heir seaward migration~ J. Fish. Res. Bd. Can. 8:24l-263~
Hobbs, D.P. 1937. Natural reproduction of quinnat salmon, brown and
rainbow trout in certain New Zealand waters. New Zealand Mar. Dept.
Fish. Bull. No.6. 104 pp.
Holton, G.D. 1953. A trout population study on a small creek in Gallatin
County, Montana. J. Wildl. Mgmt. 17:62-82.
Spatial variability in biomass of brook, brown, and rainbow trout is
shown for four sections of Trout Creek, Montana in 1950 and 1951.
This variability may be related tothe relative distribution of the
species, stream section width and depth, and pool-riffle ratiQs.
Survival, growth, reproduction, arid movements of these species are also
examined. In addition, survival of planted brook trout is investigated.
Hoopes, R.L. 1975. Flooding as the result of Hurricane Agnes, and its
effect on a native brook trout population in an infertile headwater
stream in central Pennsylvania. Trans. Amer. Fish. Soc. 104:96-99.
Hopkins, C.L. 1970. Some aspects of the bionomics of fish in a brown.
trOut nursery stream. New Zealand Mar. Dept. Fish. Res. Bull. No.4. .
38 pp.
A study of brown trout and benthic invertebrate populations in two
New Zealand streams. Seasonal trout population estimates from October
1964 through June 1966 are presented.. Population density, growth, and
food of the trout are analyzed. The availability of food and living
space are discussed as potential limiting factors for the trout. The
extent to ~hich these factors are important with respect to size and
age of the fish is also examined. In addition, data on biomass of the
dominant benthic organisms in three study sections from 1964 through
1966 are presented.
Horton, P.A. 1961. The bionomics of brown trout in a Dartmoor stream.
Anim. Ecol. 30:311-338.
J.
Horton, P.A., R.G. Baily, and S.I. Wilsdon. 1968. A comparative study of
the bionomics of the salmonids of three Devon streams. Arch. Hydrobiol.
65:187-204.
Huet, M. 1959. Profiles and biology of Western European streams as related
to fish management. Trans. Amer. Fish. Soc. . 88:155-163.
Hunt, R.L. 1971. Responses ofa brook trout population to habitat
development in Lawrence Creek. Wise. Dept. Nat. Resour. Tech. Bull.
No. 48. 35 pp.
41

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HW1t, R.L. 1974. Annual production by brook trout in Lawrence Creek
during eleven successive years. Wise. Dept. Nat. Resour. Tech. Bull.
No. 82. 28 pp.
Extensive data are presented on a long-term study of the brook
trout'population in this Wisconsin stream from 1960 through 1970.
Production is given for the entire population and by age group for'
o four stream study sections. Temporal variability in biomass is also
shown by age group for April and September.' Comparison is made
between production in one section that W1derwent habitat development
and production in the other sections. A discussion of compensatory
mechanisms in growth and survival of each age group that keep the
total production at a relatively stable level is also included. '
HW1t, R.L. '1976. A long-term evaluation of trout habitat development
and its relation to improving management-related research. Trans.
, Amer~ Fish Soc. 105:361-364. '
HW1ter, J.G. 1959. Survival and production of pink and chum salmon in
a coastal stream. J. Fish. Res. Bd. Can. 16:835-886.
'A comprehensive study of factors affecting survival and production of
pink and chum salmon in Hooknose Creek, British Columbia, from 1947
through 1956. Effects of temperature, stream discharge, sex ratio,
population density, and predation on these populations are analyzed.
Data on the number of predators (coho salmon and sculpin) as well as
weir COW1tS of pink and chum salmon fry and percent predation are
given for each year of the study. 0
Hynes, H.B.N.
19:1-15.
1975.
The stream and its valley.
Verh. Int. Ver. Limnol.
Idyll, C.1942. Food of rainbow, cutthroat, and brown trout in the
Cowichan River system, B.C. J. Fish. Res~ Bd. Can. 5:448-458.
Jones, A~N. 1975. A preliminary study of fish segregation in salmon
spawning streams. J. Fish BioI. 7:95-104.
Keeley, P.L., and T.E. Nickelson.
salmonids. Ore. Wildl. Comm.
18 pp.
1974. Streamflow requirements of
Ann. Prog. Rep. AFS-62-3.
Keenleyside, M.H.A. 1962. Skin-diving observations of Atlantic salmon and
brook trout in the Miramichi River, New BrW1swick. J. Fish. Res. Bd.
Can. 19:625-634.
Kennedy, M~, and P. Fitzmaurice. 197I.
Salmo-trutta (L.) in Irish waters.
B. (18):269-352.
Growth and food of brown trout
Proc. Roy~ Irish Acad. 71, Sect.
42

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Knight. N.J. 1980. Factors affecting the smo1t yield of coho salmon
(Oncorhynchus kisutch) in three Oregon streams. M.S. thesis. Oregon
State Un'iv.. Corvallis. 101 pp.
Koski. K.V. 1966. The survival of coho salmon (Oncorhynchus kisutch) from
egg deposition to emergence in three Oregon coastal streams. M.S.
thesis. Oregon State Univ.. Corvallis. 84 pp. '
Koski. K V. 1975. The survival and fitness of two stocks of chum salmon
(Oncorhynchus keta) from egg deposition to emergence in a contro11ed-
stream environment at Big Beef Creek. Ph.D. thesis. Univ. of
Washington. Seattle. 211 pp.. .
Krohn. D.C. 1967. Production of the reticulate sculpin (Cottus perp1exus)
and its predation on salmon fry in three Oregon streams. M.S. thesis.
Oregon State Uni v.. Corvall is. 78 pp.
, 'Lake. J.S.
Austr.
1957. Trout populations and habitats in ,New South ,Wales.
J~ Mar. Freshw. Res. 8:414-450.
A comprehensive survey of 100 streams containing brown'and/or rainbow
trout. These are divided into three general categories with'
corresponding differences in mean pH. mean temperatures. bottom fauna
production. and trout growth. Analysis of angling effects on the
trout populations is also unde~taken. ' '
Larkin. P.A. 1971. Simulation studies of the Adams River sockeye
salmon (Oncorhynchus nerka). J. Fish. Res. Bd. Can. 28:1493-1502.

Latta. W.C. 1969. Some factors affecting survival of young-of-the-
year brook trout. Sa1velinus fontinalis (Mitchill). in streams.
pp. 229-242 in T.G. Northcote (ed.). Symposium on salmon and trout in
streams; H.R:-MacMil1an Lectures in Fisheries. Univ. British
Columbia. Vancouver.
A study of age 0 brook trout from the Pigeon River. Michigan, in
1962. 1964. and 1965. Magnitude and periodicity of mortality are
discussed in a brief review of other studies. Mortality from
starvation, in relation to water temperature. is investigated in
stream aquaria. Relationships between levels of groundwater and
abundance of fry are also examined. The importance of grOtmdwater
levels in regulating the carrying capacity of the stream is discussed.
LeCren. E.D. 1969. Estimates of fish populations and production in small
streams in England. Pp. 269-280 in T.G. Northcote (ed.). Symposium
on salmon and trout in streams; ~R. MacMillan Lectures in Fisheries.
Univ. British Columbia, Vancouver. .
Leonard, J.W. 1941. Some observations on the winter feeding habits of
brook trout fingerlings in relation to natural food organisms present.
Trans. Amer. Fish. Soc. 71:219-235. '
43

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Lewis, S.L. 1969. Physical factors influencing fish populations in
pools of a trout. stream. ,Trans. Amer. Fish. Soc. 98:14-19.
A study of brown and rainbow trout populations in pools of Little Prickly
Pear Creek, Montana, in the sunnners of 1965 and 1966. The importance
of surface area, volume, depth, streamflow~ and cover in accounting
for the variation in population numbers, is. examined. Combinations of
these' factors are also considered as well as differential responses of
,.the two species. Cover is discussed, in terms of protection and .
photonegative response; streamflow is analyzed in terms of space-
food relationships.
Lindroth, A. '1955. Distribution," territorial behaviour and movements
of sea trout fry in the River Indalsa1ven. Inst. Freshw. Res.
Drottningholm Rep. 36:l04-l19~ '
Lindsay, R.B. 1974., Distribution and survival of coho salmon fry after
emigration from natal streams. M.S. thesis. Oregon State Univ.,
Corvallis. 41 pp.
, ,
Lister, D.B. and H.S. Genoe. 1970. Stream habitat utilization by
cohabiting underyearlings of chinook (Oncorhynchus tshawytscha)
and coho (0. kisutch) salmon in the Big Qua1icum River, British
Columbia. -J. Fish. Res. Bd.Can. 27: 1215-1224.
Lorz, H.W. 1974. Ecology and management of brown trout in Little
Deschutes River. Ore. Wildl. Connn. Fish. Res. Rep. No.8. '49 pp.
Lowry, G.R. 1964. Net production, movement, and food of cutthroat
trout (Salmo clarki Richardson) in three Oregon coastal streams.
M.S. thesis. Oregon State Univ., Corvallis. 72 pp.
A study of cutthroat trout populations in the streams of the Alsea
Watershed Study in 1962 and 1963. Population size, growth rate, ,
and net production estimates are made. Data on monthly biomass for
the 1959-1962 year classes are presented. Magnitude and timing of
upstream adult and downstream kelt and smol t migrations are
investigated. Trout food habits are studied during the time of
emergence and early growth of coho salmon fry to determine the extent
of predation. Seasonal variation in kind and quantity of food
consumed is also examined. Much of the information is published in
Lowry (1965 and 1966).
Lowry, G.R. 1965. Movement of cutthroat trout, Salmo clarki clarki
(Richardson) in three Oregon coastal streams. Trans. Amer. Fish. Soc.
94:334-338.
Lowry, G.R. 1966. Production and. food of cutthroat trout in three
Oregon coastal streams. J. Wild1. Mgmt. 30:754-767.
44

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Maciolek, J.A;" and P.R. Needham. 1952. Ecological effects of winter
conditions on trout and trout foods in Convict Creek, California,
1951. Trans. Amer. Fish. Soc. 81:202-217.
Maher, F.P., and P.A. Larkin. 1954. Life history of the steelhead trout
'of the Chilliwack River, British Columbia. Trans. Amer. Fish. Soc.
84:27-38.
Maitland, 'P.S. 1965. The feeding relationships of salmon, trout,
minnows, stone loach and three~spined sticklebacks in the River
Endrick, Scotland. ' J. Anim. Ecol. 34:109-133.
Major, R.L., andJ.L. Mighell. 1969. Egg-to-migrant survival of spring
chinook salmon (Oncorhynchus tshawytscha) in the Yakima River,
. Washington. Fish. Bull. 67:347-359.
'Manion, P.J. 1977. Downstream movement of fish in a tributary of
. southern Lake Superior. Progr. Fish-Cult. 39:14-16. .

Mann,R.H.K. 1971. The populations, growth 'and production of fish in
four small streams in southern England. : J. Anim. Ecol. 40:155-190.
Mason, J.C. 1974. Movements of fish populations in
Vancouver Island: A' summary of weir operations
including comments on specific life histories.
'Serv~ Tech. Rep. 483. 35 pp. '
Lynm Creek,
during 1971 and 1972
Can. Fish. Mar.
Mason, J.C. 1975. Seaward movement of juvenile fishes, including lunar
periodicity in the movement of coho salmon (Oncorhynchus kisutch) .
fry. J. Fish. Res. Bd. Can. 32:2542-2547. ' .
Mason;J.C. 1976. Response of underyearling coho salmon to supplemental
feeding in a natural stream. J. Wildl. Mgmt. 40:775-788.
An, important study on the role of food as a limiting factor in juvenile
coho salmon production in a small British Columbia stream.
Supplemental feeding resulted in a six-fold increase in the stream
carrying capacity. Effects of this feeding on survival, growth, and
pre-winter lipid reserve are also examined. However, there were no
effects on the smo1t yield the following spring. Attempts to increase
winter carrying capacity by installing artificial refuges were
ineffective. Complex winter behavior patterns may be responsible in
limiting the population size.
Mason, J.C., and D.W. Chapman. 1965. Significance of early emergence,
environmental rearing capacity, and behavioral ecology of juvenile
coho salmon in stream channels. J. Fish. Res. Bd. Can. 22:173-190.
A study of juvenile coho salmon behavior in artificial stream .
channels. Aggressive behavior is observed within 1 week of emergence
and is investigated for 5 months. Territorial defense, threat, and
45

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chasing areexaminedo Distribution in pool. and riffle areas is
studied. The effects of different times of emergence on fish size
and territoriality are also examined~ Environmental rearing
capacity is discussed in terms of food and stream area.

McDonald,.J. 1960. The behaviour of Pacific salmon fry during their
downstream migration to freshwater and saltwater nursery areas.
J. Fish. Res. Bd. Can. 17:655,..676.,
McFadden, J.T. 1961a. An ecological comparison of six brown trout
CSalmo trutta) populations. Ph.D. thesis. Pennsylvania State Univ.
,91 pp. '
McFadden, J.T. 1961bo A population study of the brook trout,
Salvelinus fontinalis. Wildl. MonOgr. No.7. 73 pp. '
A comprehensive study of the brook trout population in Lawrence
Creek, Wisconsin, from 1953 through 1957. Data are presented on
population, estimates, lengths, age structure, density, biomass,
and natural mortality in each of four study sections for each age
group of trout. Length-fecundity relationships are also examined.
Angling intensity is investigated by means of a complete creel
census each year. Data obtained include number of fishing trips,
number, sex, size, and weight of trout caught and exploitation
rates. Angler mortality is compared to natural mortality of the
trout, and management options are discussed as means of keeping an
adequate recruitment to the population.
McFadden, JoT. 1969~ DYnamics and regulation of sa1monid populations
in streams. pp. 313-329 in T.G. Northcote Ced.), Symposium on salmon
and trout in streams; H.~ MacMillan Lectures in Fisheries.Univ.
British Columbia, Vancouver.
McFadden, J,. T., G.R. Alexander, and D.S. Shetter. 1967. Numerical changes
and population regulation in brook trout Salvelinus fontinalis. J.
Fish. Res. Bd. Can. 24:1425-1459.
McKernan, D.L., D.R. Johnson, and J.I. Hodges. 1950. Some factors
influencing the trends of salmon populations in Oregon. Trans.
N. Amer. Wildl. Conf. 15:427-449.
One of the earlier studies attempting to explain fluctuations in
salmon population levels in Oregon. Columbia River chinook
salmon catch data are presented from 1866 through 1948 and coho salmon
catch data from coastal rivers are presented from 1923 to 1948.
-P61li.ltiOn, changes in angling regulations, hatcheries, logging,
. streamflow, and salinity are analyzed as potential factors
responsible for the observed fluctuations. Ih addition, the,
effects of the fishery, indicated by marking experiments, fishing
intensity, and economic trends, are examined.-
46

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McNeil, W.J. 1966. Effect of the spawning bed environment on reproduction
of pink and chum salmon. Fish. Bull. 65 :495-523. .
'McNeil, W.J. 1968. Effect of streamflow on survival of pink and chum
salmon in spawning beds. ?p. 96-114 in Proc. Forum on the relation
between logging and salmon. Feb. 8-9:-1968. Alaska Distr., Amer. Inst.
Fish. Res. BioI., Jtmeau. .
McNeil, W.J. 1969. Survival of pink and chum salmon eggs and alevins.
?po 101-117 in T.G. Northcote (ed.), Symposium on salmon and trout in
streams; H.R:-MacMillan Lectures in Fisheries. Univ. British
Columbia.
Reviews a long-term study of freshwater survival from 1940 through
1965. Also discussed are a dome-shaped production curve, density-
independent mortaiity from droughts~ floods, and freezing
temperatures, artd' density-dependent mortality from redd superimposition.
Mense, J.B. 19700 Relation of density to brown trout movement'in a
Michigan stream. Ph.D. thesis. Mich. State Univ. 82 pp.
Mense, J.B. 1975. Relation 'of density to brown trout movement in a
Michigan stream. Trans. Amero Fish. Soco 104:688-695.
, Merrell, T.R. 1962. freshwater survival of pink salmon at Sashin
Alaska. Pp. 59-72 in NoJ~ Wilimovsky (ed.), Symposium on pink
H.R. MacMillan Lectures in Fisheries. Univ. British Columbia,
Vancouver.
Creek,
salmon;
Presents data on number of spawners, egg deposition, number of
seaward migrating fry, and freshwater survival from 1940 through
1959. Differences in magnitude and timing between odd-year runs
and even-year rtmS are examined. Preliminary data in the last year
of the study seem to indicate survival is related to stream
gradient 0 There also seems to be a relationship between size of
the rtm and section of the stream used for spawningo
Miller, R.B. 1957. Permanence and size of home territory in stream-dwelling
cutthroat trout. J. Fish. Res. Bd. Can. 14:687-691.
Miller, W.H. 1971. Factors influencing migration of chinook salmon fry
(Oncorhynchus tshawytscha) in the Salmon River, Idaho. Ph.D. thesis.
Univ. Idaho, Moscow.
Mills, D.H. 1964. The ecology of the young stages of the Atlantic
salmon in the River Bran, Ross-Shireo Freshw. Salmon Fish. Res.
Edinburgh 0 No. 32. . 58 ppo
Mills, D.H. 1969. The survival of juvenile Atlantic salmon and brown
trout in some Scottish streams. pp. 217-228 in T.G. Northcote (ed.),
47

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Symposium on salmon and trout in streams~ HoR. MacMillan Lectures
in Fisheries. Univ. British Columbia, Vancouver.
'Moring, J.R. 1975. The Alsea Watershed Study: Effects of logging on
the aquatic resources of thr~e headwater streams of the Alsea River,
Oregon. Part II -.Changes in environmental conditions. Ore. Dept.
Fish Wildl., Fish. Res. Rep. No. 9~ 39 pp.
Moring, J.R., and R.L. Lantz~ 1975. The Alsea Waters~ed Study: Effects'
of logging on the aquatic resources of three headwater streams of the
Alsea River, Oregon. Part I - Biological studies. Ore. Dept. Fish
Wildl., Fish. Res. Rep. No.9. 66 pp.
A summary' report, incorporating data from a number of earlier
publications from the study. Three small tributaries were,studied,
one clearcut down to the streambank, one clearcut with buffer strips
left, and the third left uncut as a control, for 15 years. The
pre-logging period was 1959 to 1965 and the post-logging period
was 1967 to 1973. Biological studies concentrate mainly on coho
salmon and cutthroat trout, the dominant species in the streams. '
For adults, data include numbers, timing of migration, size, sex
ratio, and fecundity. Data on juveniles include emergence, growth,
biomass, production, mortality, and downstream migration. Effects of
logging are reported.' ,
Mortensen, Eo 1977. Pop~lation, survival, growth and production of
trout Salmo trutta in a small Danish stream. Oikos 28:9~15.
Population dynamics of brown trout in a small Danish stream'were
'examined from 1973 through 1975. Biomass of the 1971-1975 year
classes is investigated. Population size, production, mortality,
length-weight relationships, and smolt yield are also discussed.
Mortensen, E. 1978. The population dynamics and production of trout
(Salmo trutta L.) in a small Danish stream. Pp. 151-160 in J.R. '
Moring (ed.), Proc. wild trout - catchable trout symp. Ore. Dept.
Fish Wildlo, Res~ Devel. Sect., Portland.
Population size, survival, growth, biomass, and production of brown
trout were studied in three sections of a small Danish stream from
1974 through 1976. Variability in age group composition and biomass
between the three sections is shown. Density-dependent fry mortality,
density-independent mortality of older fish, and variability in
production-biomass ratios between age groups are also analyzed.
Mundie, JoH. 1969. Ecological-J.IDplications of the diet of juvenile coho
in streams. Pp. 135-152 in T.G. Northcote (ed.), Symposium on salmon
and trout in streams; Ho~ MacMillan Lectures in Fisheries. Univ.
British Columbia, Vancouver.
Mundie, J.H. 1974.
Res. Bd. Can.
Optimization of the salmonid nursery stream.
31:1827-1837.
J. Fish.
48

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Murphy, M.L.1979. Predator assemblages in old-growth and logged sections
of small Cascade streams. M.S. thesis. Oregon State Univ., Corvallis.
72 pp.
. Murphy, M.L., and J.D. Hall. MS. Effects of clearcutting on predators and
their habitat in small streams of the Cascade mountains, Oregon.
Can. J. Fish. Aquat. S'ci. (in press).
. Murray, A.R. J968. Numbers of Atlantic salmon and brook .trout captured
and marked at the Litle Codroy River, Newfoundland, counting fence and
auxiliary traps, 1954-1963. Fish. Res. Bd. Can. Tech. Rep. No. 84. .
135 pp.
Magnitude and timing of upstream adult and downstream kelt and
smolt migrations in this river are examined for both species.
Marking experiments are used to determine movements,' utilization,
and survival. Marine survival is determined from commercial
catch and escapement of marked fish.
Murray, A.R., and.T.J Harmon. 1969. A preliminary consideration of the
factors affecting the productivity of Newfoundland streams. Fish.
Res. Board Can. Tech. Rep. No. 130. 405 pp. .
An extensive survey of the major streams in Newfoundland,. including.
description, location, and certain morphometric, edaphic, climatic, and .
biotic. characteristics. Morphometric factors examined include watershed
shape, elevation, gradient, vegetational cover, runoff, and discharge.
Edaphic factors include surface geology, total dissolved solids,
micronutrients, acidity, conductivity, and hardness. Climatic factors
examined are air temperature and precipitation. Competition, predation,
fecundity, emigration, survival, and disease are the biotic factors
investigated. Both fish and aquatic invertebrate populations are
studied. Native salmonid species are Atlantic salmon, brook trout,
and Arctic char. Pink salmon, rainbow trout, and brown trout have
been introduced. In addition, the sport fishery for Atlantic salmon
is examined. .
Muttkowski, R.A. 1929. The ecology of trout streams in Yellowstone
National Park. Roosevelt Wildl. Ann. 2:155-240.
Narver, D.W. 1971. Effects of logging debris on fish production.
pp. 100-111 in J.T. Krygier and J.D. Hall (eds.), A symposium: Forest
land uses an~stream environment. Oregon State Univ., Corvallis.
Narver, D.W., and B.C. Andersen. 1974. Fish populations of Carnation
Creek and other Barkley Sound streams - 1970-1973: Data record
and progress report. Fish. Res. Bd. Can.. MS Rep. 1303. 115 pp.
The initial data from a long-term study of the effects of logging on
the aquatic resources of a British Columbia watershed. The study'
is divided into pre-logging (1970-1974), logging (1975-1979), and
post-logging (after 1979) periods. In partic~lar sampling is
49

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conducted on sections of upper and lower Carnation Creek, "C".
tributary, "1600" tributary, and Useless, Frederick, Ritherdon,
and South Pachena creeks., This report contains data on population
estimates, density, late summer biomass, ,growth, length-weight
relationships, and condition of both resident and anadromous .
salmonids. The primary species include coho and chwn .salmon, and
rainbow,steelhead, and cutthroat trout. Data are updated for
every year through 1977 by Andersen and Narver (1975) and
Andersen (1978).
Neave, F. 1947. Natural propagation of chum salmon in a coastal stream..
Fish. Res. Bd.Can. Progr. Rep., Pac~ Coast Sta. No. 70:20-21.
Neave, F. 1949. Game fish populations of the Cowichan River.
Res. Bd. Can. Bull. 84. 32 pp.
Fish.
Neave, F. 1953. Principles affecting the size of pink and chum salmon
populations in British Columbia. J. Fish. Res. Bd. Can. 9:450-491.
Neave, F~, and W.P. Wickett. 1953.. Factors affecting the freshwater
development of Pacific salmon in British Columbia. Proc. Seventh
Pac., Sci. Congr. (1949). 4:548-556.
A review of some chemical, physical, and biological factors that
.affect salmon populations in British Columbia streams. Chemic~l
factors include dissolved oxygen and pH. Important physical factors
are bottom type, particularly on spawning grounds, obstructions,
streamflow, including floods, and water temperature. Predation,
interspecific and intraspecific competition, and disease are important
biological factors discussed. Relationships between precipitation
. and adult returns two years later are also analyzed.
Needham; P.R., and F.K. Cramer. 1943. Movement of trout in Convict Creek,
California. J. Wild1. Mgmt. 7:142-148.
Needham, P.R., and A.C. Jones. 1959. Flow, temperature, solar radiation,
and ice in relation to activities of fishes in Sagehen Creek, California.
Ecology 40:465-474.
Needham, P.R., J.W. Moffett, and D.W. Slater. 1945. Fluctuations in
wild brown trout populations in Convict Creek, California. J. Wi1dl.
Mgmt. 9:9-25.
A study of the brown trout populations in two sections of this stream
from 1939 to 1944, one closed and the other open to angling.
Reproduction, survival, seasonal weight changes, and- density are
analyzed in both sections. The effects of angling on these.
parameters are s~own. The possibility of stocking the stream with
more trout is discussed as a method to offset angling pressure.
50

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Neill, R.M. 1938. The food and feeding of the brown trout (Salmo trutta
L.) in relation to the organic environment. Trans. Roy. Soc. Edinburgh
59(pt. 11):481-520.
Newbold, J.D., D.C. Erman, and K.B. Roby. 1980.. Logging effects on
stream macroinvertebrates and the role of bufferstrips. J. Fish.
Res. Bd~ Can. (In press.) .
.Nickelson, T.E. 1975. Streamflow requirements of salmonids.
Wildl. Comma Ann. Prog. R~p. AFS-62-4. 19 pp.
Ore.
Nickelson, T.E., and R.E. Hafele. 1978. Streamflow requirements
of salmonids. Ore. Dept. Fish Wildl. Proj. AFS-62. 26 pp.
Nickelson, T .E. ,. and R. R. Reisenbichler. 1977. Streamflow requirements
of salmonids. Ore. Dept. Fish Wildl. Proj. AFS-62. 24 pp.

Northcote, T.G. 1978. Migratory strategies and production in freshwater
fishes. Pp. 326-~59 in S.D. Gerking (ed.), Ecology of freshwater fish
production. BlackweIT Scientific Publ., Oxford.
NylIian, O.L.1970. Ecological interaction of brown trout, Salmo trutta
L., and brook trout, Salvelinus fontinalis (Mitchil1), in a stream.
Can. Field-Nat. 84:343-350.
O'Connor, J"F., and G. Power. 1976.. Production by brook trout (Salvelinus
fOhtinalis) in four streams in the Matamek Watershed, Quebec. J.
Fish. Res. Bd. Can. 33:6-18.
Four streams were examined for brook trout populations from 1971
through 1973. Biomass of each age group each year is presented
as well as total biomass in the study sections. Production and
production-biomass ratios in each stream are also examinedo Differences
in these parameters between streams are related to variations in
stream cover, recruitment, and total biomass 0 The roles of food
and cover as possible limiting factors on adult production are
discussed.
Orsborn, JoF., and C.H. Allman (edso). 19760 Instream flow needs:
a symposium. Vol. 2. Amer. Fish. Soc. 657 pp.
Osborn, C.E. 1967. A population study of the rainbow trout (Salmo
gairdneri) in a central Oregon stream. M.S. thesis. Oregon State Univ.,
Corvallis. 65 pp. .
Osterdahl, L. 1969. Thesmo1t run of a small Swedish river. pp. 205-215
in T.G. Northcote (ed.) , Symposium on salmon and trout in streams;
~R. MacMillan Lectures in Fisherieso Univ. British Columbia, Vancouver.
Otto, C. 1976. Size, growth, population density and food of brown trout
Sa1mo trutta L. in two sections of a south Swedish stream. J. Fish
BioI. 8:477-488..
51

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Overton, W.S. 1978. Study design considerations for impact assessments.
pp. 24-25 in W.T. Mason, Jr. (ed.),Methods for the assessment and
prediction-of mineral mining impacts on aquatic communities.
U.S. Fish Wildl. Servo FWSjOBS-78j30.
Pearson, L.S., K.R. Conover, and R.E. Sams. 1970. Factors affecting the
natural rearing of juvenile coho salmon during the summer low flow'
season. Ore. ,Fish Comm., Portland. 64 pp. (Processed.)
A study of summer juvenile coho salmon populations in 19 streams
in four Oregon river systems from 1962 through 1966. Relationships
between minimum streamflow and coho density are examined. Mean
pool velocity and fish density in pools also seem to be related.
Effects of streamflow on f~sh size and incidence of disease are
investigated. Fish numbers and'size are related to the rearing
capacity of a stream. Shade, cover, temperature, bottom composition,
stream size, comPetition, and food production can also affect
coho rearing capacity.
Pennak, R.W. 1971.
Hydrobiologia
Toward a classification of lotic habitats.
38:321-334.
Phillips, R.W. 1970. Effects of sediment on the gravel environment
and fish production. pp. 64-74 in J.T. Krygier and J.D. Hall (eds.),
A symposium: Forest land uses and stream environment. Oregon State
Uni V. , Corvallis.' .
Platts, W.S. 1974. Geomorphic and aquatic conditions influencing salmonids
and stream classification with application to ecosystem classification.
Surface Environ. Mining, U.S. For. Serv., Billings, Montana. xiv +
199 pp.
An extensive study from 1970 to 1972 on conditions influencing
salmonid populations in 291 stream sites in the upper South Fork
of the Salmon River watershed, Idaho. Physical and hydrologic
parameters examined include climate, geological processes and types,
stream channel width, depth, gradient, and order, elevation, pool-
riffle ratios, and streambank vegetation and stability. Effects of
these factors on the salmonid populations are analyzed. The
salmonid species present include rainbow, steelhead, and brook
trout, Dolly Varden, chinook salmon, and mountain whitefish.
Platts, W.S., and F.E. Partridge. 1978. Rearing of chinook salmon in
. tributaries of the South Fork Salmon River, Idaho.. U.S.D.A. For.
Servo Res. Pap. INT-205. 12 pp.
Power, G. 1973. Estimates of age, growth, standing crop and production
of salmonids in ~ome north Norwegian rivers and streams. Inst.
Freshw. Res. Drottningholm Rep. 53:78-111.
Pritchard, A.L. 1944. Physical characteristics and behavior of pink
salmon fry at McClinton Creek, B.C. J. Fish. ,Res. Bd. Can. 6:217-227.
52

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Raleigh, R.F. . 1978. Habitat evaluation procedure for aquatic assessments.
pp. 131-137 in W.T. Mason, Jr. (ed.), Methods for the assessment
and prediction of mineral mining impacts on aquatic. communities.
U.S. Fish Wildl. Servo A~S/OBS-18/30.
Rayner, H.J. 1937. Notes on the food of trout of.Yosemite National
Park. Calif. Fish Game 23:149-156.
Redmond, M.A. 1975. Natural production. Pp. 134-135 in J.R. Bohne and
L. Sochasky (eds~), New England Atlantic salmon restoration conference.
Int. Atlantic Salmon Spec. P~bl. Sere No.6.'
Reed, R.J. 1967..0bservat:Lon of -fishes associated with spawning salmon.
Trans. Amer. Fish. Soc. 96:62-67.
Reimers, N. '1957. Some aspects of the relation between stream foods
and trout survival. Calif. Fish Game 43:43-69.
Provides data on the percentage composit~on of the major benthic
fauna groups, organisms in the drift, and numbers and volume of food
consumed by brown trout in Convict Creek, California. The speed
and effectiveness of trout digestion at various water temperatures
are also investigated. In addition, .the effects of prolonged
starvation on the. condition and viability of trout in different
seasons are examined. .
Reimers, P.E. 1973. ,The length of residence of juvenile fall chinook
salmon in Sixes River, Oregon. Ore. Fish Comm. Res. Rep. 4(2) :1-43.
Reimers, P.E., and RoE. Loeffel. 1967. The length of residence of
juvenile fall chinook salmon in selected Columbia River tributaries.
Fish Comm. Ore. Res. Briefs 13(1):5-19.
Ringstad, N.R. 1974. Food competition between freshwater sculpins
(genus Cottus) and juvenile coho salmon (Oncorhynchus kisutch):
experimental and ecological study in a British Columbia coastal
stream. Can. Fish. Mar. Servo Tech. Rep. 457. 88 pp.
An
Ruggles, C.P. 1959. Salmon populations and bottom fauna in the Wenatchee
River, Washington. Trans. Amer. Fish. Soc. 88:186-190.
Ruggles, C.P. 1966. Depth and velocity as a factor in stream rearing
and production of juvenile coho salmon. Can. Fish Cult. 38:37-53.
Salo, E.O.,and W.H. Bayliff. 1958. Artificial and natural production
of silver salmon, Oncorhynchus kisutch, at Minter Creek, Washington.
Wash. Dept. Fish. Res. Bull. 4. 76 pp. + appendix.
A comprehensive long-term study of wild and hatchery coho salmon
populations from 1938 to 1953. Magnitude and' timing of upstream
adult and jack migrations are investigated. Numbers, lengths, and
timing of both hatchery and wild smolt migrations are studied.
53

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For the hatchery-reared fish, fecundity, freshwater survival, marine
survival, and total survival are evaluated for various rearing period
durations. For wild fish, fecundity and survival are also .
investigated. Comparisons of these data, as well as contributions to
the commercial fishery, are made. In addition, estimates for
optimum wild escapement' are presented.
Salt, G.W. 1969. The role of laboratory experimentation in ecological
. research. pp. 87-100 in J. Cairns, Jr. (ed.), The structure and
function of fresh~water microbial communities. Virginia Polytechnic
Inst. State Univ., Res. Div. Monograph 3. .
Saunders, J.N., and M.W. Smith. 1~62. Physical alterat~on of stream
, habitat to improve brook trout production. Trans. Amer. Fish Soc.
91:185-188. "
A study of initial effects of habitat development on the brook
trout population in one ,section of Hays Brook, Prince Edward
Islando Data are presented on numbers of trout from 1947 through
1959 before development and in 1960 after developmento Artificial
changes to the stream include 13 dams, 12 deflectors, and several
covers in a 450-yd (4ll-m) study section. Changes in survival and.
growth are also examined.
Saunders, J.W., and M.W. Smith. 1965. Changes in a stream population
of trout assoCiated with increased silt. J. Fish. Res.' Bd. Can.
22:395-404.
Saunders, RoL., and J.H. Gee. 1964. Movements of young Atlantic salmon
in a small stream. J. Fish. Res. Bd. Can. 21:27-36.
".
Scarnecchia, DoL. 1978. Factors affecting coho salmon production in
Oregon. M.S. thesis. Oregon State Univ., Corvallis. 100 pp.
Schaffer, W.M., and P.F. Elson. 1975. The adaptive significance of
variations in life history among local populations of Atlantic
salmon in North America. Ecology 56:577-590.
Schlucter, M.D., and J.A. Lichatowich. 1977. Juvenile life histories
of Rogue River spring chinook salmon Oncorhynchus tshawytscha
(Walbaum), as determined by scale analysis. Ore. Dept. Fish
Wildl. Info. Rep. Fish. 77-5. 24 pp.
Schuck, H.A. 1943. Survival, population density, growth, and movement of
the wild brown trout in Crystal Creek. Trans. Amer. Fish. Soc.
73:209-230.
Seegrist, D.W., and R~ Gard. 1972~ Effects of floods on trout in Sagehen
Creek, California. Trans. Amer. Fisho Soc. 101:478-482.
Shapovalov, L., and A.C. Taft 0 1954. The life histories of the steelhead
rainbow trout (Salmo gairdneri gairdneri) and:silver salmon
54

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(Oncorhynchus'kisutch) with special'reference to Waddell Creek,'
California, and recommendations regarding their management. Calif.
Dept. Fish Game Bull. No. 98. 3?5 pp'.
A comprehensive treatment of spawning migration, egg production,
freshwater life, smolt migration, marine life, and survival of both
species in this stream" with data from 1933 to 1944. Predators in
both fresh and sal t water are als'o examined.. A review of other studies
within each'topic is included as well. '
Shetter, D.S. 1961. Survival of brook trout from egg to ,fingerling
stage in two Michigan trout 'streams.' Trans. Amer. Fish. Soc. 90:
252-258. '
Brook,trout populations were examined in Hunt Creek from 1943 through
1948 and in the Pigeon River from +949 through 1958. Number of
spawners, fecundity, egg deposition, number of fingerlings, and
survival rates are investigated. Variability in these parameters is
shown through data for the entire period.
Shetter, D.So, and A.So Hazzard. 1938. Species compositon
and stability of fish populations in sections of three
trout streams during the summer of 1937. Trans. Amer.
28l-~02. ' ' ,
by age groups
Michigan
Fish Soc. 68:
, ,
One of the earliest studies to quantitatively show spatial variability
between sections of the' same stream for brook, brown, and rainbow
trout in 1935. A total of eight sections in three streams were
examined for physical 'characteristics~ relative amounts of shade, trout
biomass, and age group composition.
Skeesick, DoG. ,1970. The fall immigration of juvenile coho salmon into
a small tributary. are. Fish Comm. Res. Rep. 2(1):90-95.
Slack, H.D. 1934. The winter food of brown trout (Salmo trutta L.).
J. Anim. Ecol. 3 :105-108.
Smith, M.W., and J.W. Saunders. 1958. Movements of brook trout,
Salvelinus fontinalis (Mitchill), between and within fresh and salt
water. J. Fisho Res. Bd. Can. 15:1403-1449.
Smoker, W.A. 1953. Stream flow and silver salmon production in
western Washington. \'lash. Dept. Fish., Fish. Res. Pap. 1(1) :5-12.
Smoker, W.A. 1955. Effects of streamflow on silver salmon production
in western Washington. Ph.D. thesis. Univ. Washington, Seattle.
198 pp.
Solomon, D.J., and R.G. Templeton.
trutta L. in a chalk stream.
1976. Movements of brown trout Salmo
J. Fish BioI. 9:411-423.
55

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Stauffer, T.M. 1972. Age, growth, 'and downstream migration of juvenile
rainbow trout in a Lake Michigan tributary. Trans. Amer. Fish. Soc.
101:18-28. '
Stein, R.A., P.E. Reimers, and J.D. Hall.
juvenile coho (Oncorhynchus kisutch)
,tshaWytscha) in Sixes River, Oregon.
1737-1748.
1972. Social interaction between
and fall chinook salmon (0.
J. Fish. Res. Bd. Can. 29:
Stewart, P.A.
stream.
1970. Physical factors influencing trout density in a small
Ph.D. thesis. Colorado State Univ., Ft. Collins. 78 pp.
An investigation of physical variables affecting the density of brook
and rainbow trout in a small Colorado stream. These variables,
include mean depth, underwater overhanging rock cover, undercut banks,
streamflow, and turbulence. Multiple regression analysis is, performed
to determine which factors are statistically important to the density
of each species. '
Straskraba,M. 1966. On the distribution of the macrofauna and fish in
two streams, Lucina and Moravka. Arch. Hydrobiol. 61:515-536.
Sumner, F.H. 1952. Migrations of salmonids in Sand Creek, Oregon.
Amer. Fish. Soc. 82:139-150.
Trans.
Surber, E.W. 1951. Bottom fauna and temperature conditions in relation
to trout management in St. Mary's River, Augusta County, Virginia.
Virginia J. Sci. 2:190-202.
Swanson, F.J., and G.W. Lienkaemper. 1978. Physical consequences of large
organic debris in Pacific Northwest streams. USDA Forest Servo Gen.
Tech. Rep. PNW-69. 12 pp.
Symons, P.E.K. 1971. Behavioral adjustment of population density to
available food by juvenile Atlantic salmon Salmo salar. J. Anim.
Ecol. 40:569-587.
Symons, P.E.K. 1974. Territorial behavior of juvenile Atlantic salmon
reduces predation by brook trout. Can. J. Zool. 52:677-679.
Symons, P.E.K., and M. Heland. 1978. Stream habitats and behavioral
interactions of underyearling and yearling Atlantic salmon (Salmo
salar). J. Fish. Res. Bd. Can. 35:175-183.
Tagmaz'yan, Z.1. 1971. Relationship between the density of the downstream
migr~tion and predation of young pink salmon [Oncorhynchus gorbuscha
(Walb.)]. J. 1chthyol. 11:984-987.
Tarzwell, C.M. 1938. An evaluation of the methods and results of stream
improvement in the Southwest. Trans. N. Amer; Wildl. Conf. 3:
339-364.
56

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Tebo, L.B., Jr. 1957.
Amer. For. Proc.
Effects of sil tation on trout streams'.
1956:198-202.
Soc.
Tebo, L.B., Jr., and 1'1.1'1. Hassler. 1963. ,Food of brook, brown, and
rainbow trout from streams in western North'Carolina. J. Elisha
Mitchell Soc. 79:44-53.
Thomas, J.D. 1962. The food and growth of brown trout (Salmo 'trutta
, L.) and its feeding relationships with the salmon parr (Salmo
salar L~) and the eel (Anguilla anguilla L.)in the river Teify,
West Wales. J. Anim. Ecol. 31:175-205. '
Thomas, J.D. 1964. Studies on the growth of trout, Salmo trutta; from
four contrasting habitats. Proc. Zool. Soc. London 142:459-509.
Tomasson, T. 1978. Age and growth of cutthroat trout, Salmo clarki
clarki ,(Richardson), in the Rogue River, Oregon. M~hesis.
Oregon State Univ., Corvallis. 75 pp.
Triska, F.J., J.R.Sedell, and S.V. Gregory. 1980. The coniferous forest
stream: physical, chemical and biological interactions through ,
'space and time. In R.L. Edmonds (ed.), The natural behavior and response
to stress of western coniferous forests. ' Dowden, Hutchinson; and '
Ross, Inc., Stroudsburg, Pa. (In press.) ,
Vincent" R.E., and \'I.H. Miller. 1969. Altitudinal distribution of
brown trout and other fishes in a headwater tributary of the
South Platte River, Colorado. Ecology 50:464-467.
Warren, C.E. 1979. Toward classification and rationale for watershed
management and stream protection. U.S. Environ. Proto Agency, EPA~
600/3-79-059. 143 pp. '
Warren, C.E., J.H. Wales, G.E. Davis, and P. Doudoroff. 1964. Trout
production in an experimental stream enriched with sucrose.
J. Wildl. Mgmt. 28:617-660.
Waters, T.F. 1969. Invertebrate drift--ecology and significance to
stream fishes. pp. 121-134 in T.G. Northcote, (ed.), Symposium on
salmon and trout in streams:- H.R. MacMillan Lectures in Fisheries.
Univ. British Columbia, Vancouver.
Watts, R.L., G.L. Trembley, and G.W. Harvey. 1942. Brook trout in
Kettle Creek and tributaries. Bull. Penn. Agr. Exp. Stat. No. 437.
41 pp.
Welch, H.E., P.E.K. Symons, and D.W. Narver. 1977. Some effects of
potato farming and forest clearcutting on New Brunswick streams.
Can. Fish. Mar. Serve Tech. Rep. 745. 13 pp. .
57

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. .'
W~sche,:T.A. 1973. Parametric determination of
trout. Univ~ Wyoming. Water Resource Res.
Ser. No. ~7. 102 pp.
minimum streamflow for
Inst., Water Resource
, Wesche, T .A., 1976. Development and application of a trout cover rating
system for IFN determinations. Pp. 224-234 in J.F. Orsborn and
C.H. Allman (eds.), Instream flow needs: a-Symposium. Amer. Fish.
Soc. '
White, H.C. ,1930. Some observations on the eastern brook trout (S.
fontinalis) of Prince Edward Island. Trans. Amer. Fish. Soc.-
60:101-'108.
White, R.J. 1975. Trout population responses to streamflow fluctuation
. and habitat management in Big Roche-a-Cri Creek, Wisconsin. Verh.
Int. Ver. Limnol. 19:2469-2477.
White, R.J., and R.L. Hunt. 1969. Regularly occurring fluctuations in
year-class strength of two brook trout populations. Trans. Wisc.
Acad. Sci., Arts, and Letters. 57:135-153.
Brook trout populations in Lawrence Creek and Big Roche-a-Cri Creek.
were studied from 1953 through 1964. Fluctuations in population
numbers of age 0 and I trout were observed in both streams, with peaks
every other year. These cycles may be related to competition for food
and/or space in the streams.' ' ,
Wickett, W.P. 1951. The coho salmon population of Nile Creek.
Res. Bd. Can. Progr. Rep. (Pac.). 89:88-89.
Fish.
Wickett, W.P. 1958. Review of certain environmental factors affecting
the production of pink and chum salmon. J. Fish. Res. Bd. Can.
15: 1103-11260
Streamflow is the primary factor investigated in relation to pink
and chum salmon pr~quction in British Columbia streams. Effects
of low flow on adult migration and egg deposition are discussed.
Negative impacts of floods on populations are also examined. Effects
of streamflow on incubation stages and the survival of deposited eggs
are analyzed. Another factor discussed is spawner density, in
relation to egg density and gravel permeability.
Wickett, W.P. 1962. Environmental variability and reproduction potentials
of pink salmon in British Columbia. Pp. 73-86 in N.J. Wilimovsky (ed.),
Symposium on pink salmon; H.R. MacMillan Lectures in Fisheries. Univ.
British Columbia, Vancouver.
Willis, R.A. 1962. Gnat Creek weir studies.
Ore. 71 pp. (Mimeo.)
Final report, Fish Corom.
A weir was set up on this Oregon tributary to the lower Columbia River.
Counts of migrating coho and chinook salmon we~e made from 1955
58

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th~ough 1961. Timing of migration, adult-jack relationships" sex
r~tios, fecundity, and freshwater survival of wild fish are examined.
S~rvival of hatchery-released fish in this stream is also analyzed.
ptel~minary data indicate that commercial gill-net catches in the
l4w~~ Columbia River are related to discharge in Gnat Creek two years
earlier.
Wiseman, J'.S. 1951. A quantitative analysis of foods eaten' by eastern
brook ,trout.' Wyoming Wild1. 15(10) :12-17.'
Withler, I.L. 1966. Variability in 'life history characteristics of
steelhead trout (Salmo gairdneri) along the Pacific coast of
North America. J. Fish. Res; Bd. Can. 23:365-393.
Wong, D.M. 1975. Aspects of the life history of 'the Paiute cutthroat
trout, Salmo clarki seleniris Snyder, in North Fork Cottonwood Creek,
Mono County, California, with notes on behaviour in a stream
aquarium. M.A. thesis. California State Univ., Long Beach. 190 pp.
Wyatt, B. 1959. Observations on the movements and reproduction of the
Cascade form of cutthroat ,trout. M.S. thesis. Oregon State Univ.,
Corvallis. 60 pp.
59

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APPENDIX
This appendix is a compilation of examples of the best data available
on temporal and spatial variation in populations of stream salmonids
. (reprinted here with permission of the copyright owners and publishers).
The tables are arranged geographically--north to south, west to east.
A dash in lieu of data indicates "not sampled."
The tables are reprinted with permission of the following organizations
and individuals:
American Fisheries Society.: Tables A-9, A-11, A-13, A-22, A-23, A-28, A-31.

Blackwell Scientific Publications: Table A-33.

California Department of Fish and Game: Tables A-IS, A-19, A-20.

Scientific Information and Publication Branch, Canada Department of
Fisheries and Oceans: Tables A-3, A-29, A-30.
Pacific Biological Station, Canada Department of Fisheries and Oceans,
Nanaimo, British Columbia: Table A-4. . .

Research and Resource Services, Canada Department of Fisheries and
Oceans, St. Johns ,Newfoundland: Table A- 32.
The Fisheries Society of the British Isles:
Tables A-34, A-35, A-36.
Fisheries Research Division, New Zealand ~Iinistry of Agricul ture and
Fisheries: Table A-37.
Oregon

United
Table A-2.

Institute of Animal Resource Ecology, University of British Columbia:
Table A-I.
Department of Fish and Wildlife: Table A-7.
'~ I
States Department of Commerce, National Marine Fisheries Service:
Washington State Department of Fisheries: Table A-6.
The Wildlife Society: Tables A-2l, A-24.
Wisconsin Academy of Science, Arts, and Letters: Table A-25.
Wisconsin Department of Natural Resources:
Dr. David Au: Tables A-9, A-II, A-13.
Dr. Richard Gard: Tables A-16, A-17, A-18.
Dr. Gordon G1ova: Table A~5.
Tables A-26, A-27.
Mr. Gerald Lowry:
Table A-14.
60

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TABLE A-I. POTENTIAL EGG DEPO~:iTIONAND FRESHWATER SURVIVAL OF PINK SALMON,
 SASHIN CREEK, ALAS ,A, 1940-1959 (FROM MERRELL 1962).  
  ,I' .,'.   
  ,   
  !..,/   
Brood Potential Egg .. Number of . Freshwater
I' .,:.
Year .Depositi.ona  Mi grating Fry Survival (%)
1940 52,858,000  3,402~830 6.4 
1941 88,678,000  1,024,364 1.2 
1942 81,502,000  674,672 0.8 
1943 14,980,000  227.,673 1.5 
1944 3,904,000'  104,113 2.7 
1945 5,062,000  41,900 0.8 
1946 736,000  1, 168 0.2 
1947 .1,330,000  26,454 2.0 
1948 516,000  9,016 1.7 
1949 4,800,000  176,025 3.7 
1950 86,000  (50 killed) 0.1 
. 1951 4,062,000  379,585 9.3 
1952 run destroyed  0  
1953 1, 284,000  90,219 7.0 
1954 12,000  576 4.8 
1955 10,286,000  1,232,872 12.2 
1956 1,018,000  5,043 0.5 
1957 2,587,758  588,976 22.8 
1958 174,OQO  10,577 6.1 
1959 40,379,327  5,332,468 13.2 
a Based on 2,000 eggs per female except when actual fecundity was calculated
in 1957 (1,986 eggs) and 1959 (2,021 eggs).
61

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T~BLE A-2. WEIR COUNTS OF COHO SALMON FRY AND SMOLTS, SASH IN CREEK, ALASKA,
1956-1968 (FROM CRONE AND BOND 1976).. . . .
Year
Total Count
Fry
Smolts
t956
1957
1958
1959
1960
1961
1962
1963
1964b
1965b .
1967
1968
373
.2 ,854
218
9,923
2,699
1,209
1,236
44,023
12,000
10,000
1,665'
928
1,961
1)015
1,587
1,258
2,489
2,865
1,599a
334

1,400
1,440
. a Partial count.

b Weir not functional.
damaged in 1966 - no
Counts are estimates "from fyke net sampl ing. Heir
sampling conducted. .
TABLE A-3. HEIR COUNTS OF DOWNSTREAM rUGRATING PINK AND CHUM SAU10N FRY,
HOOKNOSE CREEK, BRITISH COLUMBIA, 1947-1956 (FROM HUNTER 1959).
Brood Year Pink Chum Total
   , ,
1947 33,349 108,746 142,095
1948 64,312 77 ,539 141,851
1949 54,061 44,463 98,524
1950 234,396 431,399 665,795
1951 242,993 .269,701 512,694
1952 1,227,025 182,200 1,409,225
1953 204,250 984,504 1,188,754
1954 907,458 353,761 1,261,219
1955 86,256 49,443 135,699
1956 454,148 69,830 523,978
62

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TIIBLE 11-4. BIOMIISS (~/i IN LATE SUI~MER) OF COIIO SilL MOil IINO RIIINBOH IItID CUTTIIROIIT TROUT III smEM1S III TliE VICINITY OF TIlE
CIIIWIITrON CREEK WIlTERSII£[) , BRITISII COllJMB III , 1970-1977 (FROM NIIRVER /\NO IItlDmSON 1974; IINDERSOtl IIND NIIRVER 1975;
IIND IlII0ERSON 1978). . .
. ..- -.---.-------- "---_____.0____._.-- ---_.- ----.--...-- - - . ---.-- -.-- --'-'--.----"--,--" -_._-..... ---------.-.------------ -----------.----------- ------_...----- -.--.-..., ,
---.-- ---"-.---.-----.-----..--.-----.-- .-. ...._---.-- ----
------'----'-- - -. ---- ...---- 0.__..---.-- .-.-.-- .-- . ---.. .--.-- .---.---.-- .-- -_.- - --..- '4'__--"--"-.'---' - . .. ...- ---- .....-
Year
Lower
Carnation Cr.
Coho --ii" iiliiow
Upper
Carnation Cr.
'--Cutthr-oa}---
Trib "C"
CuTthroaT
Trib
"1600"
colio"ClifiJiroai:
Useless
Cr.
COlIn -cilTIII-I'oaT
Ft"eded ck
(I". .
Colw" AiiTrlbow
RiLherdon
Cr.
C"1,tTI1'rc.iiil
. S. Pachena
Cr.
Colio- - fiii i ilho~i
.-.. ..n__-.,-._-_.... _.~ ---- -'"~'---'-'_.--'-- - -.. .------...--- _.. -_._-~ --.- -.-.-.- - -.-". .~----._.__.. -- -_'____h_--____--.-.-.-.-.. .--.-..---- - -.- -- .--"'-'--.---,.--
..- n - . _.. - -. -..-.-
 1970 2.72 1. 21            
 1971 \. 89 0.92 3.46       1.06 0.0 5.36 1.,87 1.24
 1972 1. 47 0.43 2.90 4.79 4.88 \. 93 0.84 1.f>4 1. 54 0.33 4.50 1. 31 1.11
 1973 1.46 0.59 3.97 5.61\ 2.45 1. 97 0.19 1.66   3.07 0.74 0.4<'
 1974 \. 59 0.49 \. 94 3.45 2.84 0.95 0.2B 2,39 0.44 0.01\ 3.31 0.59 O. HI
0\               
VI 1975 \.61\ 0.44            
 1.66 3.71 4.19 0.39 0,70 2.01 1. 36 0.07 2.84 \.85 .0.23
 1976 1. 23 0.30 1. 94 2.77 2.75 0.25 0.38 0.67 0.66 0.02 1.1\0 0.82 0.17
 1977 1.62 0.32 1. 08 2.47   0.53 0.85 0.95 0.09 1. 33 1.1\0 0.37
 ----.----.---. .._---._----_..~.__._._--- -- ------..-.----.---------- .-----------.-.----------------.-.- ...:.. ---..- .---. ---.-. -----.---. -.. -.- .._"- -_._~-".-..__.
 '---'-'---'--.--- ._----------_._~. ----.-.- ..---..----.-- ---.-.-.--------- --.--- .--- ..--.-.-.- _._---- --..----..--. --.---.--. ..-....--.- ---..--.- -- --. -----...._--

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T/\BLE /\-5. BIOM/\SS (20 clII/sec).  
  -  
 b Mainly Cottus aleuticus.              
    -...,.--.--. ---,-              

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TABLE A-6., ESCAPE~,1ENT, POTENTIAL EGG DEPOSITION, AND FRESHHATER SURVIVAL OF
WILD COHO SALMON, MINTER CREEK, WASHINGTON, 1938-1953 (FROM SALO
AND BAYLIFF 1958) .
Brood Females Released Egg  Freshwater
Year Upstream Potential Smolt Count Survival (%)
1938 967 2,657,316 35,452 1.33
1940 1,393 4,577,398 32,085 , 0.70
1942 786 ~,873,038 31,893 1. 70
1943 906 2,092,860 23,177 1.11
1944 500 1,376,500 30,408 2.21
1946 500 '1,097,000 41 ,848 3.81
1948 98 186,200. 17,839 . 9.58
1949 114 287,964 27,.781 9;65 .
1951 411 1,086,684 22,545 2.07
1952 753 . 1,929,186 31,363 L63
1953 491 1,150,413 18,620 . 1.62
TABLE A'-7.. COUNTS OF SPAWNI.NG COHO SALMON AND SMOLTS AT DOWNSTREAM I~EIR ON
GNAT CREEK, OREGON, 1954-1959 (FROM WILLIS 1962). .
Brood Year Female Spawners Smolt Count
1955 26 2,996
1956 29 1 ,847
1957 67 . 1,013
1958 40 1,061
1959 45 3,226
65

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TABLE A-8. ESCAPEMENT, POTENTIAL EGG DEPOSITION, AND FRESHWATER SURVIVAL OF
 COHO SALMON. DEER CREEK, OREGON, 1959~1971 (FROM KNIGHT 1980).
Brood. Female Egg Smolt . Freshwater
Year Escapement Potentiala Count Survival (%)
1959 21 43,197 ' 1,917 4.44
1960 19 44,156 2,210 5.00
1961 28 67,620 2,775 4.10
1962 18 ,'42,030 2,082 4.95
1963 27 62,964, 2,368 3.76
1964 44 104,940 1,836 1. 75
1965 24 55,176 2,245 4.07
196,6 56 141,798 2,461 1. 74
1967 23 52,815 2,160 4.09
.1968 39 . 80,301 1,484 1.85
1969 8 15,484 738 ' 4.77
1970 10 22,119 1,072 4.85
1971 36 73,134 1,923 2.63
a Calculated from reg~ession equation (Koski 1966), Y = -3,184 + 7.81 X,
where X = average length in mm (from unpublished data) and Y = individual
fecundity. Total fecundity equals Y times the number of female spawners.
TABLE A-9. '
ESTIMATED BIOMASS (g/m2) OF, JUVENILE COHO SALMON, DEER CREEK,
OREGON, 1959-1968 (FROM CHAPMAN 1965 AND AU 1972). DATA ARE
INTERPOLATED FOR THE BEGINNING OF EACH MONTH INDICATED, FROM
POPULATION ESTIMATES MADE LESS FREQUENTLY THROUGHOUT THE YEAR.
 1959 1960 1961 1962 1963 1964 1965 1966 1967 1968
June 4.8 2.7 2.1 5.9' 4.2 2.1 4.8 5.0 8.7 2.3
July 4.0 2.5 2.5 3.6 3.7 2.1 5.9 4.3 7.2 3.0
Aug 3.1 2.3 3.0 3.0 3.4 2.1 4.3 3.8 6.2 3.7
Sept 2.9 2.3 3.2 4.0 3.1 2.6 3.2 3.6 5.9 4.4
Oct 3.1 2.5 3.4 4.7 2.5 3.1 2.9 3.8 6.1 5.0
. Nov 3.6 2.7 3.4 5.1 2.4 3.5 2.8 4.0 6.3 5.3
Dec 3.6 2.0 3.2 4.0 2.4 3.7 2.7 4.1 6.1 5.1
Jan 3.8 2.0 3.8 4.5 2.5 3.6 3.0 4.2 4.4 4.1
Feb 4.0 2.0 1.7 4.5 2.5 2.8 2.9 4.2 3..5 2.9
Mar 4.2 2.0' 1.7 3.6 2.4 1.9 2.4 3.7 3.2 2.2
Apr 1.8 1.3 1.3 2.1 1.6 1.5 0.8 1.9 1.9 1.4
May 0.5 0.4 0.6 0.5      
66

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TABLE A-10. ESCAPEMENT, POTENTIAL EGG DEPOSITION, AND FRESHWATER SURVIVAL OF
 COHO SALMON, FLYNN CREEK, OREGON, 1959-1971 (FROM KNIGHT 1980).
Brood Female Egg Smolt Freshwater
Year Escapement Potentiala Count Survival (%)
1959 8 17 ,368 875 5.04
1960 26 66,742 776 1.16
1961 51 131,427 " 1,354 1.03
1962 2 4,644 565 12.17
1963 20 44,220 736 1.66
1964 . 10 . 24,020 663 2.76
. 1965. 11 26,565 968 3.64
1966 55 138,050 616 . 0.45
1967 10 23,130 430 1.86
1968 19 38,931 207 0.53
1969 5 9,625 140' 1.45
1970 "5 13,745 330 2.40.
1971 18 37,404 404 1.08
a Calculated from regression equation (Koski 1966), Y = -3,184 + 7.81 X, 
where X = average length in mm (from unpublished data) and Y = average 
individual fecundity. Total fecundity equalsY times the number of female
spawners.         
TABLE A-II. ESTIMATED BIOMASS (g/m2) OF JUVENILE. COHO SALMON, FLYNN CREEK,
  OREGON, 1959-1968 (FROM CHAPMAN 1965 AND AU 1972). DATA ARE 
  INTERPOLATED FOR THE BEGINNING OF EACH MONTH INDICATED, FROM 
  POPULATION ESTIMATES MADE LESS FREQUENTLY THROUGHOUT THE~YEAR.
 1959 1960 1961 1962 1963 1964 1965 1966 1967 1968
June 4.1 2.9 2.1 8.3 1.3 2.2 4.1 4.0 6.0 1.1
July 3.3 2.8 2.0 3.0 1.7 1.7 1.7 2.8 3.9 1.3
Aug 3.0 2.7 2.1 2.3 2.0 1.5 1.4 2.6 2.9 1.8
Sept 3.1 2.6 2.3 2.6 2.2 1.7 1.6 2.9 2.1 2.3
Oct 2.9 2.5 2.5 3.4 2.4 1.8 1.7 2.8 1.8 2.5
Nov 2.9 2.5 2.5 3.8 '2.2 2.0 1.7 2.6 1.7 2.3
Dec 2.4 2.0 3.5 4.5 2.0 1.9 1.7 2.4 1.7 1.8
Jan 2.2 1.9 2.1 4.1 1.8 1.9 1.7 2.5 1.8 1.1
Feb 2.2 1. 7- 2.0 4.1 1.6 1.7 1.6 2.5 1.7 0.9
Mar 2.6 1.8 1.8 3.8 1.4 1.7 1.1 2.1 1.6 . 0.8
Apr 1.5 1.7 1.6 2.6 1.1 1.3 0.4 0.9 1.4 0.5
May 0.5 1.3 0.7 1.4      
     67     

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TABLE A-12.
ESCAPEMENT, POTENTIAL EGG DEPOS ITION, AND FRESHWATER SURVIVAL OF
COHO SALMON, NEEDLE BRANCH, OREGON, .1959-1971 (FROM KNIGHT 1980).
Brood Female Egg a Smolt Freshwater
Year Escapement Potential Count Survival. (%)
1959 2b 4,471 b 462 10.3
1960 2 4,192 . 223 5.32
1961 15 33,135 470 1.42
1962 4 . 9,632d 314 3.26
1963 15c 33,530d 160 0.477
1964 25c 55,884d 286 0.512
1965 28c 62,590 333 0.532
1966 19 46,664 277 0~594
1967 15 40,460 421 1.04
1968 17 35,088 194 0.55
1969 1 2,666 76 2.85
1970 2 5,386 113 2.10
1971 18 35,604 369 1.04
"
a Calculated from regression equation (Koski 1966)~ Y = -3,184 + 7.81 X,.
. where X = average length in mm (from unpublished data) and Y = average
individual fecundity. Total fecundity equals Y times the number of female
sp~wners. . .

b Estimated equivalents from 1,627 planted fry.
c Estimated from redd surveys.

d Estimated from mean female length (693.9 mm) from the other years of the
study. ~
68

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TABLE A-13..
ESTIMATED BIOMASS (g/m2) OF .JUVENILE COHO SALMON, NEEDLE BRANCH,
OREGON, 1959-1968 (FROM CHAPMAN 1965 AND AU 1972). DATA ARE
INTERPOLATED FOR THE BEGINNING .OF EACH MONTH INDICATED, FROM
POPULATION ESTIMATES MADE LESS FREQUENTLY THROUGHOUT THE YEAR.
 1959 1960 1961 1962 1963 1964 1965 1966 1967 1968
June 2.3 1.4 2.1 7.2 3.5 4.0 5.0 7.6 6.9  3.1
July 1.6 1.7 1.9 3.9 3.0 2.4 3.9 4.0 9.0  3.4
Aug 1.8 2.0 1.8 3..2 .2.9 1.5 3.5 3.1 7.8  3.8
Sept 1.9 2.0 1.8 2.8 2.7 1.3 3.4 3.7 6.5  4.2
Oet 2.2 2.2 1.7 2.8 2.8 1.2 3.3 '4.3 6.2  4..6
Nov 2.4 2.4. 1.7 2.9 2.7 1.0 2.9 4.1 3.2  4.4
Dee 2.0 2.2 1.8 2.8 2.6 0.7 2.2 4.4 3.1. 3.7
Jan 1.9 2.2 1.8 3.3 2.3 0.8 1.9 4.0 3.2  2.4
Feb 1.9 2.4 1.7 3.1 2.3 0.9 1.3 3.1 3.9  1.7
Mar 2.4 1.5 1.4 3.1 1.5 1.0 0.8 1.5 2.6  0.4
Apr 2.4' 1.1 1.8 2.1. 1.0 0.8 0.4 0.4 0.2  0.1
May . 0.9 0.3 0.7 0.9       
TABLE A-14. BIOMASS (g/m2 IN SEPTEMBER) OF CUTTHROAT TROUT, ALSEA l,/ATERSHED .
  STUDY, 1962-1973 (~ROM LOWRY 1964 AND UNPUBLISHED DATA). .. 
Year  Deer Cr.   Flynn Cr.   Needle Br.
1962   5.07 .   5.82.    3.89 
1963   .2.93   3.54    3.41 
1964  . 1.90   4.04    3.16 
1965   2.93   2.72    2.97 
1966 .   2.05   2.73    1.09 
.1967   3.29   4.26    0.68 
1968   2.15   2.71    1.65 
1969   2.80   3.70    1.46 
1970   3.83   4.01    1.14 
1971   4.20   4.27    1.32 
1972   4.03   4.13    1.39 
1973       3.79    1.53 
69

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TABLE A-15. BIOMASS (g/m2) OF SALt10NID SPECIES IN THREE NORTHERN CALIFORNIA
STREAMS; 1967-1969 (FROM BURNS 1971). .
No. Fork,
Casper Cr.

Coho Steel head
S. Fork
Yager Cr.

Steel head
Godwood Cr.
a
Coho Trout
June 1967 0.18 1.09
July 1967  
Aug 1967  
Oct 1967 0.15 1.46
June 1968 0.13 1.16
July 1968  
Aug 1968  
Oct 1968 0.19 1.44
June 1969 0.61 0.98 .
Jul y 1969.  
Aug 1969  
Oct 1969 0.81 1.13
a 1  
Stee head and cutthroat. 
 1.09 0.57
3.22  
 0.76 0.49
4.21  
 0.34 0.51
2.94  
TABLE A-16. BIOMASS OF BROOK TROUT (g/m2 IN MID-AUGUST) IN 10 SECTIONS OF
SAGEHEN CREEK, CALIFORNIA, 1952-1961 (FROM R. GARD PERS. COMM.).
SECTION I IS UPSTREAM.
 I I! II! IV V VI VII VIII IX X
1952 5.15 13.90 3.75 1.48 0.39 0.80 2.01 1.06 0.24 0.01
1953 4.71 14.24 4.74 1. 38 0.50 3.25 1.46 0.67 0.07 0
1954 4.85 10.12 2.17 1.28 0 1.29 1.01 1.60 0 0
1955 4.48 6.87 2.45 0.41 0.01 0.74 0.06 1.31 0 0
1956 2.47 6.65 1.80 0.96 0.18 0.55 0.41 0.13 0.01 0
1957 4.56 3.67 1.88 1.49 0.25 0.80 0.18 0.19 0 0
1958 2.24 2.91 1.23 0.25 0.53 1.14 0.96 1.84 0 0
1959  7.40 2.31 0.85 0.35 1. 78 0.24 0.19 0.01 0
1960 4.63 2.32 1.95 0.19 0.54 1.64 0.86 0.75 0 0
1961 3.36 2.50 0.86 1.12 0.55 2.85 1.56 0.40 0 0
70

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I
TABLE A-17. BIOMASS OF BRO~1N TROUT (g/m2 IN MID-AUGUST) IN' 10 SE~TIOt-lS OF '
SAGEHEN CREEK, CALIFORNIA, 1952-1961 (FROM R. GARD PERS. COMM.).
SECTION I IS UPSTREAM.
 I II' III .IV V VI VII VIII IX X
1952 0' 0' 0 0 0 0 0.63 ' 8.56 1.99 0.40
1953 0 0 0 0 0 0.08 6.34 1.18 1.18 0.78'
,1954 0 0 0 0, 0 0 1.69 '1.47 2.03 0.67
1955. 0 0 0 0 0 0.01 3.96 1.66 4.18 0.45
1956 0 0 0 0 a 0 1.48 1.06 2.54 0.11
1957 0 0 0 0 0 0.44 0.91 0 2.95 0.01
1958 0 0 0.54 0 0 0.08 0 0.13' 4.11 0.16
1959  0 0 0 0 0.02 1.30 1. 97 2.82 0
,1960 0 0 0 0 0.06 0 1.30 0.37 2.93 0
1961 0 0 0 0 0 0.02 1. 51 0.16 1. 74 0.01
TABLE A-18. . BIOMASS OF RAINBOW TROUT (g/m2 IN MID-'AUGUST) IN 10 SECTIONS OF .
SAGEHEN CREEK, CALIFORNIA, 1952-1961 (FROM R. GARD PERS. COMM.).
SECTION I IS UPSTREAM'.
 I II III IV V VI ' VII VIII IX X
1952 0 1.12 1.45 4.69 0.75 .1. 01 0.99 0.54 0.21 0.07
1953 0 1.27 4.30 4.78 0.77 1.20 ,1.46 0.99 0.28 0.03
1954 0 1.42 2.31 3.89 0.37 1. 73 2.42 0.45 0.01 (j
1955 0 1.04 2.82 4.07 0.31 2.26 1.43 0.77 0 0.12
1956 0 0.41 1. 30 2.21 0.45 1. 75 0.41 0.06 0 0
1957 0 1.09 1.23 3.45 0.44 2.93 1.82 0.64 0.15 0
1958 0 1.12 0.91 2.35 1.43 3.55 1. 79 0.75 0.59 0.07
1959'  1.04 0.54 4.02 0.37 5.30 2.50 0.91 0 0
1960 0 0.37 0.83 4.60, 0.83 4.29 1.54 0.11 0.49 0
1961 0 0.41 0.36 5.91 0.89 6.67 1.64 0.96 0 0
71

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                                                                                         ikd
TABLE A-19.  ESCAPEMENT, POTENTIAL EGG DEPOSITION,  AND FRESHWATER  SURVIVAL  OF
             COHO SALMON, WADDELL CREEK, CALIFORNIA,  1933-1940 (FROM
             SHAPOVALOV AMD TAFT 1954)

Brood
Year
1933
1934
1935
1936
1937
1938
1939
1940
Femal e
Escapement
222
309
59
157
37
56
150
115
Egg
Potential
560,690
725,014
141,233
377,352
91 ,728
130,074
396,321
257,886
Smolt
Count
3,573
4,911
1,067
1,926
852
1,740
152
711
Freshwater
Survival (%)
0.64
0.68
0.76
0.51
0.93
1.34
0.038
0.28

TABLE A-20.  DOWNSTREAM TRAP COUNTS OF STEELHEAD TROUT BY AGE GROUP,  WADDELL
             CREEK, CALIFORNIA, 1933-1942 (FROM SHAPOVALOV AND TAFT 1954).
                 0
                        I
II
III
 Year
Number  Percent  Number  Percent  Number  Percent   Number   Percent
1933-34
1934-35
1935-36
1936-37
-1937-38
1938-39
1939-40
1940-41
1941-42
604
699
1,365
1,875
1,946
691
2,239
3,306
2,009
19
39
35
53
57
11
64
59
35
741
578
1,655
1,191
1,015
3,699
945
2,049 .
2,834
24
32
42
34
30
60
27
36
50
1,657
484
830
451
410
1,720
. 292
251
843
53
27
21
13
12
28
8
4
15
112
28
90
11
19
77
7
9
33
4
2
2
a
1
1
a
a
1

a < 1 percent.
                                     72

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TIIOLE A-21. BIOHASS (g/m2) OF BROOK, RAINBOII, AND BROI-IN TROUT, TROllr CREEK, MONTAflA, 1950-1951 (FR0I1110LTON 1953). SECTION I
IS UPSTREAM.
Sec ti on
2
-----------.----- ------ --------------- -'--'--'----'------------'.-_n______...--- --.--..-.--.----.-----------.------..
4
---..-- -----.-.-. _._- . -----.--.,.- -'.'--'-'-----'----'--'-----'- -.----.+---- ----...------..-- -- ..-_.._--_.-..-. ._-. ---- "..-- ----- ,.--. ----------- --_.__..---
---- -'--
- ---.- "-"--'----,-,-----
-_._----~-- ..-.-......-- .'-__'_h. ---_..
Date
Auy. Nov. I-lay Aug.
24 18 20 12
1950 1950 1951 1951
July Sept. flov. !Iay Aug.'
25 14 19 30 4
1950 1950 1950 1951 1951
_..----..-.
--" ----------.-'-----"'---
---- --... .'-.;---.-.-.---. _._---_._.~-
Br'ook
13.0 34.9 12.4 10.B
3.B B.O 3.7 2.1 5.3
3.6, 6.5 5.0 6.6 11. 1
0.4 0.0 4.6 6.7 13.4
RainlJow
2.0
3.0
0.9
D.9
o
Brown
o
o
o
~---------- -...-.-----------.------------.---.----------.-.--------------.---.------------.----
3
---------- ------------.--.-..-------
-_-4____..- '.'--- ..
..__.--_.~-..-
Aug. Sept. !Iov. June I\ug.
9 14 4 3 11
1950 1950 1950 1951 1951
Aug. Oct. May Aug.
1 20 17' 5
1950 1950 1951 1951
----- --------.-------- --------- .
-------.----.-.- --'.-. --- ---
2.2 2.1 :3.4 1.3 2.1
4.9 5.0 5.0 6.9 5.7
0.5 0.5 10.6 0.1 0.3
2.1 1.4 1.1 0.2
3.9 6.4 3.0 6.2
0.6 9.7 4.1 0.7
.----.---..-.-----.--- ---.------.------------.------..----.---.----------------- --------- -.-- -~-_._--- ----.--_.- --- -- -.-------..-.-- -. ~n
TABLE A-2? BIOMASS (y/l) OF BROHII, RAIUBOH. AND BROOK TI!OUT HI 11 SECTIONS or LITTLE PRICKLY PEAI{ CREEK; MONTANA, SUMMER 1966
(FIWH ELSER 1960). SECTION 1 IS UPSTREAM .
See ti on
3
4<1
-------------.---------.---.------.-----.-..---- - ---.---------.-.--.---------- ---_._.~----_. --- --.-.-------.--. -.-- .---.---
9
lOa
.....
VI
-.--.---..----------.--.-------------------------.-----------------.----.----..-----------..--.-------------------.-..---..- ..--
lA
2
5
6<1
Oa
7
--~--_."._"-------_._-_._._.._---------_._-,--_._---------------------.---------.------------------.--.-.----------.-----'-.-..----
 2 2308  1862 2266 29% 2630 3157   1255 3213
Area (III ) 1093 21'1S 17'10
Mean width (III) 5.5 2.7 5.5 7.3 7.6 8.2 B.O 7.3 6.'1 7.6 13.4
,.Brown  l. 57 3.14 12.9 16.7 3.36 17.6 4.37 17. 1 10.'1 11. l' 1. 57.
Rainbow 4.70 5.0'1 5.60 5.72 0.90 6.39 2.02 7.85 1. 91 5.9'1 '1.04
Brook  1. 01 10.8 3.81 2.02 0.22 0.'15 0.11 0.3'1 0.67 0.22 0.11
Total trout 7.23 19.0 22.3 24.4 4.'18 2'1.'1 6.50 25.3 13.0 17.3 5.72
a Altered sections.
.- ---.--.- ---.- - --.-- -.-.-------.- -.-----..- --~- ---.---- ----.---'--.-----.--.-.-.-----. .---' -------- -- -----..---.--------.-----.--. --..._~ --- --r---------...---
----._------- ------.-.-.. --..-- ----..
--,---- _.~_._.- -- -~.. ..- .-.- .--....----- ---.--.-- ~ ---.-.--..---.- -.----.----..--.-- ---.._-----.~_.- -'-----'--------.-. .--.. --_u- ..----.------.--.-.--.

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TABLE A-23. NUMBER OF CUTTIIROAT TROUT TRAPPED, AND IIUMBER REMAINING IN STREAM AFTER TRAP WAS REMOVED,a ARNICA CREEK, YELlOWSTOtIE
PARK, WYOMING, 1950-1958 (FROM BENSON 1960),
--~_._------ -------..-.------.-----..-----------.----------------------'-----'-------'-----.--'-----------.-----'-
.._-----_._-------_._---------------------~-------------------.-.------------.----------.--------.---------.-
, .
Year.
Oil te trap
removed
_~~9~~.!.!r.~!lJI- Q.----
Trapped In Stream
.._.A9~.:.9EP.!!P.-l.__--
TI'apped, In Stream
fi~ap~~a---9!'.9-f!H{ ream
Tota 1
frappe-a---'n Stream
Grand
To ta 1
. --'-'-_._-----'--'--'-'--'-'----~--------------------'-------'---------'--'-----'-------'----.---------.-..---.
 1950 Sept. 28 9,556  300  95  9,951  9,951
 1951 Oct. 6 5,240  365  39  5,641  5,644
 1952 Sept. 21 502  792  242  1.536  1,536
 1953 Sept. 28 1,332 708 763 407 82 67 2,177 1,182 3,359
 1954 Sept. 14 4,151  943  47  5,141  5,141
 1955 Sept. 24 4,182 244 340 15 7 0 4,529 259 4,788
 1956 Sept. 13 4,268 612 386 56 132 5 4,786 673 5,459
 1957 Sept. 25 2,850 405 121 270 5 3 . 2,976 678 3,654
--..j 1958 Aug. 31 36 1,950 9 30  15 46 1,995 2,041
.j:>. 
 --------------- ..--------.------......-..----.------------------------'--'----'-------------'--.-'---'-'--'-.--.---.--'--'---.--.-----
 . .   
 ---'--.---------'------'--'-'--------------,---------,-----.".-'------'--"-------'--.-------,.-----,,-
a Data not available on number of fish in stream after dismantling of trap for 1950, 1951, 1952, and 1954.

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TABLE A-24. BIOMASS (g/m2) OF BROOK TROUT, LAWRENCE CREEK, \.:IISCONSIN, 1953-
  1957 (FROM McFADDEN 1961b). SECTION A is UPSTREAM 
   Section    
  A B C.. D Total Mean.
Sept. 1953 10.30 9.81 8.40 5.52 34.03 8.51
Sept. 1954 14.53 12.36 8.81 6.31 42.01 10.50
April 1955 11. 18 9.64 6.82 6.60 34.24 8.56 .
Sept. 1955 7.11 8.25 6..10 3.02 24.48 6.12
April 1956 4.41 4.2"7 3.89 2.45 15.02 3.75
Sept. 1956 10.76. 6.01 5.50 1. 30 23.57 5.89
April 1957 14.48. 6.93 5.35 3.37 30.13 7.53
Sept. 1957 . 26.18 10.74 6.37 4.20 47.49 11.87
TABLE A-25. SEPTEMBER POPULATION ESTIMATES OF AGES 0 AND I BROOK TROUT IN
LAWRENCE CREEK AND ~IG ROCHE-A-CRI CREEK, WiSCONSIN, 1953-1964
(FROM WHITE AND HUNT 1969). . .
 Lawrence Creek
Year. 0 I
1953 10,113 2,040
1954 13,523 2,749
1955 5,720 . 2,754
1956 10,853 816
1957 13 ,258 3,370
1958 4,166 4,393
1959 22,646 1,044
1960 8,507 3,324
1961 14,313 2,360
1962 7,611 4,523
1963 10,367 2,388
1964 9,680 4,382
Big Roche-a-Cri Creek
o ..1
2,012
6,229
2,637
9,915
4.,361
5,632
4,964
7,420
1,135
. 474.
1,817
1,257
2,630
1,609
1,623
1,072
75

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ll\(}l[ 11-26. BIOr1J\SS (g/n?)a OF BROOK TROUT BY IIGE GROUP III IIPRll 11110 SEPHMBER, llll/REtICE CHEEK, IIISCONSIN, 1960-1970 (FHOM ItUNT
1974).. .
-.----. -.- -'-'------.-------.---..-- ----,------ '--___"0 --- ---- ----- .-- ------ --...-..--.-.-- --.--.-- -"---.,----, - .-..--.--. ----....----. ------- -..----....-.-.----..-- -"----'-'-"--' . .--..-.-...--.--...
- --~-----_. -.__'__.0_._- ----'--- -- ------"----------".----'----- ---------"- -- -..---.. ---- -.. --'''-------'----'---.--. ---.-.---."---.---.. -.------... - ------- ----- -.-.."0_....- .-.---..--
Year
.-.----..--.-.---- .____~I_i.l. ..---- ... .--..---....-.-....-..- .---
I II III IVt Total
"---.--..-.-----.-.- .S_e.P. ~e.!!!.~E___.__---_._-- .:..----.-- ___h_-
o [ II III IV+ Total
 -- - -------------...- ----------.-----------..---- --------'------'-'----- ---_.,-~---_.__._---.-_._._---_._._..._--_._-----_._----..-.-------.
 1960 4.00 0.375  .0.542 0.0245  4.94  2.01 4.75 0.105 0.0417 0 6.91
 1961 l. 97 1.48  0.0245 0.0294  3.50  3.83 4.39 0.922 0.0172 0 9.16
 1962 5.48 2.30  0.412 0.0049  8.20  1. 70 6.0B 0.507 0.0930 0 8.38
 1963 2.71 4.62  0.333 0.035B  7.75  2.80 3.39 1. 78 0.137 0.0392 11.15
 1964 5.20 2.72  1.20 0.108  9.23  2.51 5.56 0.995 0.387 0.0319 9.48
 1965 2.43 3.30  0.517 0.174  6.47  2.13 4.30 1.13 0.167 0.0294 . 7.81
 1966 3.96 3.38  0.934 0.130  8.40  2.54 6.19. 1.64 0.299 0.0392 10.7
 1967 3.15 5.65  1.04 0.427 10.3  2.14 4.03 2.02 0.332 0.0613 8.63
" 1968 3.35 4.43  l. 62 0.287  9.69  2.85 3.40 0.BG3 0.280 0.0686 7.46
0\   
               .    
 1969 5.25 3.75  0.699 0.194  9.B9  3.10 4.51 1. 02 0.177 0.0809 8.89
 1970 3.88 5.17  0.701 0.142  9.89  3.32 3.78 0.9110 0.145 0;0466 8.27
 r1ean 3.77 3.38  0.729 0.146  8.02  2.63 4.58 1. 09 U.194 0.0361 8.53
 . --.- -.. '.-.----."--'---------'------"-'----'-'--'- .--------.------.----- --.-----..-.--.-------.-.-.---------.-----....-------..-----.--
 ._- --- _. .------------------.------.-.----- ...- -.- .....__.-- -.---.-.- --------_.._---_. ------------.-- _._----- ....---.-- ---- --------.--.-.-.----.-.
 a Biomass f.'om IIppendix Table 1 has been divided by stream  of 4.0fi ha.    
  area    

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TIIRlE 11-27. ANNUlll pHODUCTI orl (9/nl /yr) OF OIWOK THOUT OY SECT ION Arm AGE GHOUl', lllHHENCE CHEEK, IH SCONS IN, 1960-1970 (FROM
HUNT 1974). SECTION II IS UPSTREN1. "
-----------------.---------.----------- --------------.-.---.---..
---.---------.-----------.-----------.-.--.---. -_._----._-_._--------~----_.__._---_. ---
" .
-.-----
Year
Section"
-A----Ir------c~n_____----o-
----_._------~~J1.r:.()~-~._- --.---..----.
o I' II . III IVI
Stream
Total
 _._----~-------------._----------- .,-----_.._---------_._-_._---_.__.__._--_._-----_.~_...--'---
 1960 13.0 10.1 13.4 14.0 4.1 7.7 0.3 0.4 <0.1 12.5
 1961 17.2 13.5 10.2 8.0 6.8 3.9 1.1 <0.1 0.1 11.9
 1962 14.0 10.0 11. 1 IO.IJ 3.8 6.2 0.9 0.2 <0.1 1l.2
 1963 16.5 12.0 12.9 11. 5 6.4 4.0 2.2 0.2 <0.1 12.9
 1964 19.8 12.8 9.8 1J.6 5.2 5.2 1.2 0.5 <0. I 12.2
 1965 19.5 9.8 11.0 5.4 4.3 4.2 1.9 0.2 <0.1 10.6
 1966 15.2 12.6 9.8 6.3 3.1 5.7 1.4 0.4 <0.1 10.6.
 1967 21.7 9.4 10.9 6.7 3.8 4.5 2.4 0.5 <0. I 11.2
-...j 19GIJ 21.3 12.0 1J.9 5.1 4.5 "4.0 1.9 0.6 O. I II. 1
-...j
 1969 25.8 12.0 7.9 6.6 4.7 5.3 1.6 0.3 0.1 12.0
 1970 20.5 13.2 ro.1 7.8 5.3 4.6 2.2 0.2 <,), 1 12.3
 Mean 18.8 11.6 10.6 8.2 4.3 5.0 1.6 .0.3 <0. I 11. 7
 -----------------------.---..--------.-------.-.---------.--.---.---.----
      --------------------------.--......---.-------------..-.

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TADLE A-28.
PIIYSICAL CllIIRIlCTERIST/CS ANIIIIIOMASS (g/II?) OF BROOK, DROWN, AND RAINOOW TROUT III SECT/OtiS OF TlmEE MICIIIGAN
STREAMS, 1937 (FROt1 SIiElTER AND IIAZZIIRD 1931:).
-----.-.- "..-- -------.---------..----.-.--..--.---------.------.-----.--------. ----.--'------------.-------------
-----.----.---.-----------------------..------ .--- ------ -~ -------.-.- -.-.---- - ---1"'----------'-' .---. .--. -'--'----.----.---.-...-----.. ----._--.
Stream
Section
Length
(m)
Mean
Hidth (m)
Mean
Depth (elll)
Velocity
(em/see)
Re 1 a t i ve
Shade
Oiomass
ifi:Ook'------ ifrown---lfiilnljpw-'--T 0 taT
.
-----_._~-_.._-----------~---_._----_._---_.- ---.---------------.-.----..--------..---.-----
 South 8raneh, Upper 29.5 7.6 24.9 22.9 Pa rt 1 y 0.95 0 0.35 1. 30
 Pine River          
  Middle 29.0 5.4 27.7 26.2 Densely. 0.87 0 0.53 1.40
  Lower 31.5 6.7 34.8 20.7 Partly 2.40 U.27 1. 82 4.49
 . Little Manistee Upper 32.5 8.5 43.7 4t.!! Partly 0.21 1. 92 1. 75 3.88
 River          
  Middle 46.9 11.2 44.2  Exposed 0.12 0.73 2.54 3.39
 North IIraneh, Upper 42.6 7.5 25.7 94.5. Partly 0.028 0.84 0 0.87
 Ooardman River          
  Middle 29.0 8.5 29.0 46.3 Exposed 0.041 0.13 0 0.17
"-I  Lower 37.0 9.1 23.1 51.5 Partly 0.048 0.10 0 0.15
'JO 
 --.---.------------------------      
 --------    ---.--------.---------    .------,..---

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TABLE A-29. NUMBER OF BROOK TROUT PRESENT IN SEPTEMBER IN HUNT CREEK,
MICHIGAN BY AGE~GROUP (FROM MCFADDEN ET AL. 1967).
Year 0 I II III IV Tota 1
1949 4,471 2,036 287 14 0 " 6,808
1950 3,941 2,013 304 13 0 6,271
1951 4,287 1,851 265 16 1 6,820
1952 5,033 1,763 261 16 0 7;073
1953 5,387 1,637 175 13 0 7,212
1954 6,325 2,035 234 13 0 8,607
1955 4,235 2 ,325 383 " 24 0 6,947
1956" 4,949 1,612 392 51 1 7,005
1957 6,703 1,796 309 . 33 1 " 8,842
1958 5,097 2,653 355 26 2 8 , 133
1959 4,038 2,395 685 68 0 7,186
1960 5,057 2,217 473 47 1 7,795
1961 2,809 2,017 409 23 0 5,258
1962 . 5,052 1,589 448 52 2 7,143
79

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TABLE 11-30. MEAN ANNUAL BIOMASS (g/m2) OF BROOK THOUT IN STREAMS IN MATAMEK ~'ATERSItEO. QU[(JEC. 1971-1973 (FROM O'CONNOR 111m
POWER 1976).
-.-.- .---.- ------~. ---"------'------------------ -'-~'- _....--_....._----- --_.__._-~------_._---_.~...~,._-_._-_._._----- '-------'--'--"--'-'---.------'-
--- -'---'----- _. ----------_. ----'.'--'-"-'.----..' ---....---.- ---. ---........---.------ -------.-.--.----..-..-------- ...---.--------- --_.-.----_._--,- ..+
Stream
See ti 0/1
Length (m)
Average
Hidth (m)
Year
. Biomass
u Dt"u --------l+--------~-----------T~------ - --- -;n--- ------5+------ ---6:j-- - ----To-taT
Kaikhosru
6.1
-- -- ._----------- --.'-'-"---'.----' .,-- ---'-'-'--' --.--- -.._--- -----'--'- ----- ---- '-----------'--'--~- -.- ,...- -."-.. .'--'--'- ..-.. -- .--- --.--.. -------'------'--.---..-.-.-.'--
355
Ga 11 i enne
330
6.6
Tehillie3man
620
15. 1
00
Q
Sherry
215
4.2
1971
1972
1971
1972
1971
1977
1971
1972
1973
0.35
0.35
0.08
0.08
0.23
0.28
0.28
1. 93 0.33 0.54 0.05 0.17 0 4.17
2.23 0.66 0.44 0.22 0.06 0 3.96
2.40 2.12 0.57 0.14 0.10 0 5.33
2.83 1. 74 0.49 0.00 0.14 0 5.28
0.51 0.26 0.25 0.05 0.01 0.06 1. 21
0.42 0.48 0.32 0.07 0.01 0.03 1.42
1. 25 0.87 1.13 0.25 0 0 3.78
0.79 0.51 0.30 n.15 0 0 2.03
0.57 .0.2B 0.16 0.17 0 0 1. 4 7
.._-~---_._------_._---------------_._--------- -_.~-----_._._------_.._---_._.__._---_._---_._--
----.-.---.-----.---------.------------..--------.------.-.-.---...--------------.------.--.------- ---

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TABLE A-31. NUMBERS OF BROOK TROUT HI A 411-m SECTION OF HAYES BROOK, PRINCE
Em~ARD ISLAND, 1947-1960 (FROM,SAUNDERS,.lXND Sf1ITH 1962).
Year, Age 0 Age I + Total
1947 588 351 939
1948 729 342 1,071
1949 539 279 818
1950 321 223 544
1951 166 418 584
1952 611 372 983
1953 308 362 670
1954 468 294 762
1955 758 383 1,141
1956 580 467 1,047
1957 350 363 713
1958 4$1 314 795
.1959a 371 352 723
1960 526 611 1,137
a Afte; habitat,development.  
TABLE A.,.32. COUNTS OF ATLANTIC SAU10N SMOL TS AND SEAl-lARD MIGRATING BROOK
TROUT, LITTLE CODROY RIVER, NE~/FOUNDLAND, 1954-1963 (FROM ~1URRAY
1968)., '
Year Sa 1 man Trout
1954 12,210 
1955 11 ,248 
1956 14,772 706
1957 8,900 1,067
1958 9 , 341 889
1959 12,099 1,074
1960 7,829 457
1961 8,058 312
1962 8,193 698
1963 7,326 485
81

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TABLE A-33. BIOI-lASS (g/m2) OF BROUN TROUT IN TRIBUTARIES AND TilE MAIN STEM OF TIlE UPPEH RIVER lEES SYSTEM, EIiGlAND, ]967-]970
(FROM CRISP [T Al. ]974). .
"---'---'--------"'----------'- ..---- ----.--..-.--------.-.. "-'."-- --'----'.'----- - .---.- --- -- -------- ---._u_-.---.-----.---- -... ".. -..--.----.---.- -".'.------.--..-.--..-...
--"------ - --------- --------"---_. .--..--.- -.---.-. ..-..------------ .----.- --.-.-. - - ...-----.- - ---- -_.._..~-_...__._--...._.._._. -.-- .-. .--- .-.- ..----.---------.. .-- -..-.--.-. -.- - -'.
Mai ze
Beck
River Tees
below Cauldron
Snout
River Tees
above the
Weel
Wee1head
SHe
Dubby
Sike
Mattergi11
Sike
lod!1eg i 11
Sike
-.-------------- ---------------.---------.---.--------.----------.----"--- ----'--------.-'."--.'---------.-.------. --.--.-
 Section length (lit) 70.9 30.5 23,4 .97.5 40.8 47.2 45.7
 Mean Width (Ill)  9.94 6.91 ]1.20 1. ]6 1. 54 3.26 3.0!i
 Section Area  2 705 211 262 ]13 63 ]54 139
 (m )
 -'--'-"-_._~-"-'--'-------_.,--,-,-------,----,-,-------.-------.----.-- -_._._._--_.._-~ -----------.'--------.'-' -_u_.---------- --._--- -'------'-
 August 1967   3.63 3.65  3.2]  5. ]4 
 Uc tober 1967   1,43 2.16 0.08 ]6.63 6.85 5.84 3.34
 May ]968    0.81 1. 28  2.81 2.55 .1.03 0.32
 July ]968   0.30 1.28 0.37 3,43 4.69 1.72 0.60
00       2.24 0.01 4.73 3.20 ] 1. 76 5.25
N October 1968   0.92
 May ]969    0.48 1.11  1. 96 2,45 1.18 1. 87
 August 1969   0.42 1.2]  4.44 4.09 2.68 1. 21
 October 1969   1.18 0.96  5.45 3.]4 4.87 1. 26
 May 1970    1.05 2.05  1. 56 1.66 1.63 0.57
"."..-..._- -'--"--'------------------------------ '-------'----..
-'---~---'-'-'--'-'-'.'-----'----------,-----,-,--
_....~-------_._._-- ---.---------- ---~--"-'-------'----'-'---'-----. ------ --.-------------.---.-----.--..----- --'-.---"---------'-'--'- - ---. ....-

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TABLE A-34.
MEAN (1963-1972) BIOMIISS (gll) OF BROWN TROUT IN FivE TlUBIITI\RIES OF TIlE RIVER TEES SYSTEM, ENGLAND, IN MAY,
AUGUST. Atm OCTOBER (FROM CRISP ET AI.. 1975), . .
'~----'----"- -.----.. -.-.-.----------. --_.- --- --------.---...---. _..~------_.. "---,.-. -." -----.------....--------------------.-.- ---'---'--'--'---
-.'-.-- .- --.--.--.-.---.. -.-- '-'--"-----'---'-" - '--------- - - --- _no _.. ------------ --- --. ----- -.--. ------ - ------ --.---.--..-.--..-----.-.-.---.. ..- . -- .--.---- .-...----..
~loss
Burn
Nether
lIearth Sike
Trout
Bed
Great Dodgen
Pot Sike 'A'
Great Dodgen
Pot Sike 'B'
-- -- ------------- -..---".-- ------ "--- .--.- - --_.... -...-- ..-.. .'--- -------- ------.----" --- -'---'--'-'-'- -.-----... .-... "--- . - -...,.---..-- .--- .-.---- --.--.-..-- ----
 Meall IHdth (m)a 1.9
  2 a 254
 Surface Area (m )
  Minimum 0.8
 May t1ean 1. 19
  Maximum 1.6
  Minill1U111 1.4
 August Mean 2.00
00  Max imlun 2.9
VI.   
  Minimum 1.0
 Oc toher tlean 1. 37
  Maximum 2.2
4.1 5.6 0.9 1.6
209 205 34 115
0.7 1.0 3.0 1.1
1. 14 1. 45 4.55 1.85
1.5 1.8 6.8 2.7
1.3 2.4 3.5 1.6
2.62 3.97 5.54 3.77
3.3 6.2 10.1 7. I
1.2 1.0 5.3 2.9
2.02 1.47 5.41 3.8n
2.8 1.9 8.1 5.1
. a Based 011 measurements made in tlay 1968.
..-..-------.--.---.------.-------..------------.---------.--------. -----.__.__._----------~._-----_._,-_._-_._._-------_.-
-------------------.------------.----.----------------.---.----.--.--.-.-------.--------------- --- --'------.--.----.-.--...---

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TABLE A-35. PRODUCTION (g/nh OF ATLANTIC SALMON MID BROWN mOUT IN TltREE SECTIONS OF SIIELlIGlI1I BURN, SCOTlAND, 1966-196f1
(FROM EGGLISIIAW 1970).
---- .-.- -----.-.---...---.----.-- --.---- --_.- -'-------'-".'-'--- .-..-.--.---- -- ..--- "- .-- -----------..----.--------- -----'--'-----'--------'---,-
. ,
. .
---- -.- '-----.-,--..---------------.----.---.------ --". ---.---..------ --'-'~----'- -.------.-.----- --------------....----..--------- --'---_.,----_._~_._-
, . . Year,Class
1963----1964-----1965-- ---1%6------1 967 -------T~68
Annual
, To ta 1 s
Sec ti on
Length (III)
Mean
Width (III)
Section
Area (1112)
 -,. .-------------.----.---- -.-----------------.------------.. --------- ---'------------------------'--------'-_'_"_0. --..----..-.----.-.---. - -,------ ..-.
          - .
 Sec ti on I-Ups trealll        30.8 4.10 126.0
 1966 Salmon 0.23 1. 57 1. 25 4.47   7.52   
  Trout 0.65 1. 47 3.42 4.10   9.64   
 1967 Salilion 0 0.11 0.63 2.22 7.38  10.84   
  T rou t 0 0.52 0.77 5.05 5:82  12.16   
 1968 Salilion 0 0 0.16 0.64 2.53 8.54 11.87   
  Trout 0 0 0 0.94 3. JO 2.67 6.71   
 Sec ti on 2        27.9 3.07 86.0
 1966 Salmon 0 1. 12 1.07 3.52   5.71   
  Trou t 0.51 2.49 4.69 2.80   1O.62a   
'00 1967 Salmon 0 0.03 O. HJ 1. 92 6.59  87.2   
-too  frout 0 0 2.07 5.95 3.98  12.00   
 1968 Salmon 0 0 0 0.43 2.51 7.28 10.22   
  Trout 0 0 0 1. 91 4.40 2.39 8.70   
 Sect ion 3-Downs tream        27.5 3.43 9'1.0
 1966 Salmon 0 0.51 1.03 4.69   6.23   
  Trout 0.41 1. 96 '1.44 3.95   10..76   
 1967 Salmon 0 0.30 0.52 2.87 8.59  12.28   
  Trout 0 0.46 2.02 5.68 4.54  12.70   
 1968 Sa lilion 0 0 0 0.72 2.56 7.89 11.17   
  Tt'out 0 0 0 1.45 3.92 2.114 7.81   
--.-- ----- .._--_._----------_.~--------,-.- ------.-.---..-...--- ".---'-------'- -+-----'-----'_.-__h.'__,.,. -..,.+.-. -----"---'-~---'-'----'-- ----'- -- ---. - -. ---.-..- -----'--~'---'- ---
. ."-".-'.' '-----'.-. --_. - _.__.m___.__.----- .. --------.--... -',-----,-,_u.,--".,. - - "-- "---.. __n___.,..--..----.---. - - ---'----'----'-'- -'---'--'--.- - - -'----____-.n.- ----------- .-..., -.-- .-..--------.-
a hlcludes 0.13 9/1112 production of the 1962 yea,. class.

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TABLE A-36.
BIOMASS (g/m2) OF ATLANTIC SAU1or: AND BRO\'m TROUT AT THE END OF
THE GROWING SEASON, SHELLIGAN BURN, SCOTLAND, 1966-1975 (FROM
EGGLISHAt'J Arm SHACKLEY 1977). TOTAL BIOMASS INCLUDES :1: STANDARD
ERROR.
   Salmon    Trout  Total
. Year 0+ 1+ '2+ All 0+ 1+ 2+ All Biomass
1966 3.5 1.0 1.5 6 . 0:1: 1. 9 2.5 4.0 3.2 9.8:t1.1 15.8:1:0.9
1967 2.7 1.7 0.2 4.6:1: 1. 2 3.6 6.1 1.8 11.4:1:1.5 16.1:1:2.6
1968 2.5 1.7 0.7 4.9:1:0.9 1.9 3.2 0.9 6 . 0:1: 1. 1 10.9:1:2.1
1969 4.4 2.2 0.7 7 . 3:1: 1. 9 5.0 3.8 2.9 11.6:1:2.1 18.9:1:2.8
1970 1.9 . 2.5 0.3 4.7:1:2.3 1.9 6.0 2.1 10.1:1:2.5 14. .8:1:3.9
1971 4.7 2.4 0.5 7.6:1: 1. 9 3.8 4.9 1.4 10.1:t2.9 17~7:t4.2
1972 2.6 4.6 0.3 7.5:1:2.8 3.2 6.9 0.8 10.9:1:3.0 18.4:1:4.2
1973 3.7 3.8 0.2' 7.6:1:1.4 4.0 5.4 1.2 10.7:1:2.5 18.3:1:3.2
1974 3.2 3.4 .0.0 6.6:t1.2 2.5 4.5 0.4 7.4:t2.2 14.0:1:3.2
1975 6.1 2.9 0.2 9.2:1:2.6 1.9 4.3 0.8 7. l:t2 . 1 16.3:1:4.0
Me.an . 3.5 2.6 0.5 6.6:1:1.5 3.0 4.9 1.6 . 9.5:1:2.0 16.l:t2.4
TABLE A-37. BIOMASS (g/m2) OF BROWN TROUT IN SIX SECTIONS OF HOROKIHISTREAM,
NEW ZEALAND, 1940-1941 (FROM ALLEN 1951). SECTION I IS
DOWNSTREAM. .
Zone  I IIM IIR III IV V 
Length (m) 3,167 2,035 1 ,918. 2,719 1,602 527 Total
July 1940 24.1 31.5 11. 7 24.9 41.0 18.8 25.6
Oct. 1940 28.0 36.1 11. 7 25.7 46.4 24.7 28.6
Jan. 1941 34.1 32.7 11.0 16.4 29.8 21.9 26.5
May 1941 22.8 42.8 14.9 21. 3 26.0 24.4 25.8
Oct. 1941 1.8 13.7 2.5 2.7 18.7 18.5 6.5
8'S

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