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This is to certify that the
thesis entitled
POPULATION PARAMETERS AND ABUNDANCE OF PINK SALMON

IN THE UPPER GREAT LAKES

presented by

John Francis Kocik

has been accepted towards fulfillment
of the requirements for

Master of Science degree in Fisheries and Wildlife

 

 

Major professor

flaw/M74

Date AUES‘: 23, 1988

 

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POPULATION PARAMETERS AND ABUNDANCE OF PINK SALMON
IN THE UPPER GREAT LAKES

BY
John Francis Kocik

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE

Department of Fisheries and Wildlife

1988

ABSTRACT

POPULATION PARAMETERS AND ABUNDANCE OF PINK SALMON
IN THE UPPER GREAT LAKES

BY
John Francis Kocik

Pink salmon (Oncorhynchus gorbuscha) spawner and fry
abundance, length and weight, and spawning habitat were
studied in selected tributaries of Lakes Huron, Michigan,
and Superior from 1984 to 1987. Pink salmon stocks in Lake
Superior have declined substantially since 1980, Lake Huron
stocks are steady to slightly increasing, and those in Lake
Michigan are increasing. Adult pink salmon from Lake
Michigan (males: 482 mm, 1045 9: females 449 mm, 840 g)
were significantly larger than fish from Lakes Huron
(males: 418 mm, 574 g; females 389 mm, 450 g) or Superior
(males: 410 mm, 567 g; females 388 mm, 466 g). There was
no significant size difference between Lakes Huron or
Superior fish. Spawning habitat utilized by pink salmon
was composed of gravel to cobble substrate in 0.50 to 1.50
feet of water with velocities from 0.50 to 1.5 feet per
second. There was no significant difference in the size of
outmigrating fry from Lake Huron (31.94 mm, 0.138 9), Lake
Michigan (32.64 mm, 0.132 g), and Lake Superior (31.96 mm,
0.117 g). Fry abundance was variable between years and
dependent upon spawner abundance and environmental factors.
Low estimates of survival to outmigration (0.12 %-1.15 %)
indicate that the stream phase is critical in determining

pink salmon recruitment in Great Lakes ecosystems.

To my parents.

ii

ACKNOWLEDGEMENTS

I would like to thank my major professor, Dr. William
Taylor for his unending encouragement, faith and support
throughout all phases of this project. Appreciation is
also extended to my committee members Dr. Thomas Burton and
Dr. Howard Tanner for their assistance and advice.

A special thanks to Wilbert Wagner of the Marquette
Fisheries Research Station, Michigan Department of Natural
Resources for sharing his experience, knowledge, and
insights. His cooperation and coordination of our research
efforts has enhanced this research substantially. I would
also like to thank Richard Schorfhaar, Paul Hannuksela,
Richard Jamsen, and James Peck of the Marquette Fisheries
Research Station for their help and friendship.

I extend special thanks to my wife Linda for her
constant love, encouragement, and support as well as her
invaluable assistance as a technician, data entry
specialist, and editor.

Thanks are extended to the following for their help in
the field and in the laboratory; Russell Brown, David
Dowling, Jeff Eibler, Robert Elliot, Jill Dufour, Mark

Freeberg, Katherine Grimm, Daniel Hayes, Russell Hanshue,

iii

Scott Hanshue, Andrew Loftus, John Lott, Paul Padding,
Martin Smale, Paul Thomas, Mark Turner, Gary Whelan, and
Robin Ziegler.

This publication is a result of work sponsored by the
Michigan Sea Grant College Program, project R/GLF-lS, under
grants NA85AA-8G045 and NA86AA-D-SG043 from the office of
Sea Grant, U. S. Department of Commerce, and funds from the
State of Michigan. Additional financial support for this
research was provided by the Michigan Agricultural
Experiment station, the Great Lakes Council of the
Federation of Fly Fishers, the Michigan Salmon and
Steelhead Fishing Association, and the Oakland County

Sportfishing Association.

iv

TABLE OF CONTENTS
Page
List of Tables .........................................vii
List of Figures .........................................ix
List of Appendices .......................................x
Introduction .............................................1
Study Streams ............................................8
Methods .................................................15
Fall Sampling .........................................15
Visual Survey of Spawning Adults ....................15
Population Estimates ................................16
Vital Statistics: Adults ............................17
Spawning Habitat Utilization ........................18
Spring Sampling ............................... ...... ..19
Relative Fry Abundance ..............................19
Vital Statistics: Fry ...............................21
Results .................................................22
Fall ...................................................22
Visual Survey of Spawning Adults ....................22
Population Estimates ................................25
Vital Statistics: Adults ............................28
Spawning Habitat Utilization ........................43

Spring 0......0.000000000000000000000000000.0.0.00000049

Relative Fry Abundance ..................... ...... ...49
Vital Statistics: Fry ...............................51
Discussion ............. ..... .................... ....... .55
Fall ..................................................55
Spawner Abundance ..................... ..... . ...... ..55
Vital Statistics: Adults ................. ......... ..63
Spawning Habitat ............................ ...... ..69
Spring ..................................... ......... ..72
Relative Fry Abundance ..............................72
Vital Statistics: Fry ...............................76
Ecological Role of Pink Salmon ........................77
Summary .................................................78
References ..............................................80

AppendiceBOOOOOOOOOOOOOOOOOOOO..OOOOOOOOOOOOOOOOOOOOOO0.087

vi

Table

Table

Table

Table

Table

Table

Table

Table

Table

LIST OF TABLES

Page
General trends in pink salmon abundance in
selected Michigan tributaries of the Great

Lakes. Abundance designations after Wagner

and Stauffer 1982 OOOOOOOOOOOOOOOOOOIOOO00.0.0023

Population estimates of pink salmon abundance.
in the Laughing Whitefish River ...............26

Population estimates of pink salmon abundance

in Albany Creek (* - Michigan Department of
Natural Resources Sample: - Removed all

fish in area) .................................27

Comparison of mark-recapture population estimates
of pink salmon abundance compared to visual
surveys of abundance in Albany Creek and the
Laughing Whitefish River ......................29

Average total length, weight, and numbers of
adult pink salmon by sex from selected
tributaries of the Great Lakes, 1984

and 1986. No confidence limits are reported

for sample sizes less than 4. .................31

Average total length, weight, and numbers
of adult pink salmon by sex from selected
tributaries of the Great Lakes 1985. ..........33

Average total length, weight, and numbers of
adult pink salmon by sex from selected
tributaries of the Great Lakes for 1987. ......35

Average total length, weight, and numbers of
adult pink salmon by sex from Lakes Huron,
Michigan and Superior from 1984 to 1987. ......37

Weight-length regression values for pink salmon
from selected tributaries of the Great Lakes.
Regression is in the form:

log weight - log a + b(log total length) ......39

vii

Table

Table

Table

Table

Table

10.

11.

12.

13.

14.

Sex ratio observed in pink salmon from
selected tributaries of the Great Lakes
during even year runs. A * denotes sex ratio
is significantly different from H°:sex ratio
is 1:1 using a chi square test at 0.95. ......

Sex ratio observed in pink salmon from
selected tributaries of the Great Lakes
during odd year runs. A * denotes sex ratio
is significantly different from H°:sex ratio
is 1:1 using a chi square test at 0.95. ......

Sex ratio observed in pink salmon from weekly
samples from Albany Creek. A * denotes sex
ratio is significantly different from Ho:sex
ratio is 50:50 using a Chi square test at

0.95. O...I...OOOCOOOOOOOOOOOOOOOOO0.0.0.0...O

Average total length, weight, and numbers of
pink salmon fry from selected tributaries of

the Great Lakes.

Average total length, weight, and number of
pink salmon fry from selected tributaries of

the Great Lakes.

viii

.40

.41

.42

.45

.53

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

9.

10.

12.

LIST OF FIGURES

Page
Location of all primary and secondary study
streams located in the Upper Peninsula of
Michigan and the Northern Lower Peninsula. .....9
Location of study areas on the Laughing
Whitefish River, Alger County, Michigan. ......11
Location of study areas on Albany Creek,
Chippewa County, Michigan. ............... ..... 13

Frequency distribution of spawning depth
utilized by adult pink salmon. ................46

Frequency distribution of spawning velocity
utilized by adult pink salmon. ................47

Frequency distribution of spawning substrates
utilized by adult pink salmon. Substrate

indices are standard designations (See

Appendix 3). ..................................48

Number of pink salmon fry outmigrating from
the Laughing Whitefish River in the spring of

1985 OOOOOOOOOOOOOOOOOOOOO...OOOOOOOOOOOOOOOOOOSO

Number of pink salmon fry outmigrating from
Albany Creek in the Spring of 1986. ...........52

Changes in the average size of pink salmon.
Data from Wagner (1985) and this study. .......67

Habitat utilization curves for spawning pink
salmon in the Great Lakes and Pacific Ocean:
Depth. OOOOOOOOIOOOOI00.000.000.00...00.0.000070

Habitat utilization curves for spawning pink
salmon in the Great Lakes and Pacific Ocean:
VQIOCitYo ...OOOIOOOOOOOOOOOO0.0.00.000000000071

Habitat utilization curves for spawning pink

salmon in the Great Lakes and Pacific Ocean:
Substrate. Index numbers from Appendix 3. ...73

ix

Appendix

Appendix

Appendix

Appendix

Appendix

Appendix

Appendix

Appendix

1.

LIST OF APPENDICES
Page

Non-target fish species collected during
electrofishing in Albany Creek (AC) and

the Laughing Whitefish River (LWR) during
spring and fall sampling. An * denotes

fish present in a stream. ..................87

Non-target fish species collected during
electrofishing in secondary study streams. .88

Generalized substrate classes for use in
studies to determine habitat utilization
and preference. ............................89

Descriptions of substrate materials by
percentages of embeddedness to the nearest
quartile. From Bovee (1986). ...............90

Results of Tukey's means test for total

length of pink salmon from separate streams
within each year. Streams with the same
letter have mean total lengths that are

not significantly different. ...............91

Results of Tukey's means test for weight of
pink salmon from separate streams within

each year. Streams with the same letter

have mean total lengths that are not
significantly different. ...................92

Multiple comparison analysis of pink salmon
total length by Lake and Year. .............93

List of streams surveyed over duration of
study during fall spawning runs.

Abundance designations after Wagner and
Stauffer (1982): N-None: P-Present (1-99):
C-Common (100-999): A-Abundant (1000-9999):
V-Very Abundant(>9999). ....................94

INTRODUCTION

Pink salmon (Oncorhynchus gorbuscha) are the smallest
of the five Oncorhynchid species indigenous to the Pacific
Coast of North America (Scott and Crossman 1973). They
average 340 to 630 mm fork length and about 1400 to 3600 g
in weight at maturity (Kaganovskii 1949: Ricker 1981). The
life cycle of the pink salmon is similar to that of other
Oncorhynchids and is described in detail by several authors
(Arc and Shepard 1967; Scott and Crossman 1973). Pink
salmon spawn predominately in gravel areas in freshwater
streams in the late summer to late fall. Eggs are
deposited in a redd dug by the female. The eggs incubate
in the redd until mid to late winter when the alevins hatch
from their egg. The alevins continue to live in the
interstitial spaces in the gravel feeding almost
exclusively on their yolk sac reserves until the late
spring when the fish, now called fry, emerge from the
gravel and migrate to the ocean. Upon entering the marine
environment they move directly to open waters where they
undertake extensive oceanic migrations before returning to
their native stream to spawn. In their native range, pink
salmon adhere to a stringent two year life cycle. The two

year life cycle results in reproductively isolated stocks

2
of pink salmon in even and odd years since the two stocks
do not interbreed. Considerable differences in genetics,
abundance, and biological characteristics occur between
even and odd year runs (Ricker 1962: Aspinwall 1974:
Beacham 1984). In their native range, a few exceptions to
the two year life cycle have occurred, but they are
exceedingly rare (Anas 1959: Turner and Bilton 1968: Foster
et al. 1981).

Pink salmon were indigenous from the Sacramento River
in California northward to the Aleutian Islands of Alaska
in North America (Scott and Crossman 1973) and from the
Yurappu River in Japan to the Kamchatka Peninsula of the
Soviet Union in Eurasia (Ishida 1967). Although the pink
salmon still occupies most of its native range, its
abundance has declined in some locations due to habitat
degradation and over-fishing (Ricker 1981).

Several attempts have been made to extend the marine
distribution of pink salmon within the Pacific Ocean
watershed and to introduce it to the Atlantic Ocean for a
variety of management objectives ranging from
reintroductions to the establishment of additional
fisheries. While some projects were successful (Yu and
Ustyugov 1977), the majority of them have met with little
success (Ishida 1967). One of these introduction attempts
inadvertently led to the establishment of pink salmon in
the Great Lakes. In the mid 1950's, the Ontario Department

of Lands and Forests was attempting to establish a Native

3
American subsistence fishery in the waters of Hudson Bay

utilizing pink salmon and chum salmon (Q. keta). The

 

fertilized pink salmon eggs for this project came from the
1955 year class in the Lakelse River, a tributary to the
Skeena River, in British Columbia (Arc 1979). These eggs
were raised to the fry stage at a hatchery facility located
on Thunder Bay, Lake Superior for transport to Hudson Bay
stocking locations. In the spring of 1956 an estimated
21,000 excess pink salmon fry remained at the end of the
stocking program. These pink salmon were discarded into a
sewer that led into the Current River, a tributary of Lake
Superior (Nunan 1967). The survival of these fish in
freshwater was thought implausible since pink salmon were
considered to be an obligatory anadromous species
(Rounsefell 1958: Hurley and Woodall 1968). Ironically,
the planting in Hudson Bay was unsuccessful, but the pink
salmon released into Lake Superior did survive,
establishing a unique freshwater population of pink salmon.
The first recorded occurrence of adult pink salmon in
the Great Lakes was in the fall of 1959, when anglers
reported catching pink salmon in two Minnesota tributaries
of Lake Superior (Schumacher and Eddy 1960). Pink salmon
populations remained at low but increasing levels in Lake
Superior through the 1960's (Schumacher and Hale 1962:
Kwain and Lawrie 1981: Wagner and Stauffer 1982). As this
population growth occurred, pink salmon colonized many

tributaries in Lake Superior and expanded their range to

4
the lower Great Lakes. Pink salmon were considered to be
well established in Lake Superior by 1971 (Wagner and
Stauffer 1982) and had extended their range to Lake Huron
by 1969, Lake Michigan by 1973, and to Lakes Erie and
Ontario by 1979 (Collins 1975: Emery 1981).

In addition to original odd year runs, Lake Superior
pink salmon have established even year runs that were first
observed in 1976 (Kwain and Chappel 1978). The origin of
these runs is thought to be 3-year-old pink salmon, which
are theorized to have arisen due to the cold, oligotrophic
nature of Lake Superior delaying their maturity (Wagner and
Stauffer 1980). Even year runs of pink salmon have also
been noted in Lakes Huron and Michigan. To date, only odd-
year runs have been documented in Lakes Erie and Ontario
(Emery 1981).

Since their introduction, pink salmon in Lake Superior
have expanded their range and increased in abundance
(Wagner and Stauffer 1982). By 1979, the pink salmon
population was estimated to be 10 to 20 times larger than
in past years (Wagner and Stauffer 1982). This phenomenal
growth did not continue into the 1980's. The 1981 run of
pink salmon was expected to be quite large due to high
spawner abundance in 1979. However, the runs were nearly
complete failures in most United States tributaries of Lake
Superior (Bagdovitz et a1. 1986). On the Canadian side, in
the northeast portion of Lake Superior, pink salmon

abundance has remained relatively high (Kwain 1987). The

5
abundance of pink salmon in the lower Great Lakes appears
to be expanding (Emery 1981). The trends in population
abundance in Great Lakes pink salmon can best be
categorized as unstable. Dynamic and highly variable
populations of pink salmon are common in the Pacific
Northwest (Beacham 1984).

Prior to 1981, little was known about Great Lakes pink
salmon. Research done by Michigan State University
(Bagdovitz 1985: Bagdovitz et a1. 1986), the Michigan
Department of Natural Resources (Wagner and Stauffer 1982:
Wagner 1985), the Ontario Ministry of Natural Resources
(Kwain 1982, Kwain and Kerr 1984: Kwain 1987), and the
University of Minnesota (Nicolette 1983: 1984) has
increased our knowledge of pink salmon populations in Lake
Superior. However, there is still much to be learned about
the ecology of pink salmon, not only in Lake Superior but
in all the Great Lakes.

Since pink salmon are a well established and expanding
species, information documenting their population dynamics
is needed to assess their ecological role in Great Lakes
ecosystems and to implement a well informed management plan
for these stocks. Except for the Lake Superior studies
noted above, little is known about the population
parameters and abundance of these fish in other Great Lakes
watersheds. In addition, the dynamic nature of pink salmon
population abundance poses a unique problem in assessing

their impact upon Great Lakes ecosystems. Insights into

6
their population dynamics will provide a clearer
understanding of the ecological role of Great Lake pink
salmon and the factors that influence their abundance.

This research was undertaken to continue the
monitoring of Lake Superior populations while expanding the
scope of the study to Lake Huron and Lake Michigan
populations. Specifically, my goal was to examine pink
salmon populations in two primary study rivers that
contained similar populations of pink salmon. These data
would be supplemented with information regarding the
relative abundance and biological parameters of pink salmon
in additional tributaries of Lakes Superior, Michigan, and
Huron. The incorporation of multiple study streams would
allow for the integration of data and observation to
determine if population and vital statistics shifts were
stream specific or lake wide.

In addition, the habitat utilized by spawning pink
salmon in the Great Lakes has not been adequately
documented. In order to better understand the population
dynamics of pink salmon a detailed description of their
spawning requirements is needed. This information will
allow a clearer understanding of the relationships between
the stream environments in the Great Lakes watershed and
pink salmon spawner escapements to these streams. This
information can also be used in habitat analysis techniques
such as the Instream Flow Incremental Methodology to

determine the amount of habitat available to these fish.

7

To meet the objectives outline above the following

components were examined:

1. Relative abundance of pink salmon adults during
fall spawning runs in selected tributaries of
Lakes Huron, Michigan, and Superior.

2. The relative abundance of pink salmon fry during
spring outmigration in selected tributaries.

3. Biological Parameters of:

a. adults: length, weight, sex,
b: fry: length, weight

4. The habitat types required for pink salmon
spawning and the development and refinement of a
Habitat Suitability Index (HSI) for Great Lakes

pink salmon.

STUDY STREAMS

Two primary and seven secondary study streams were
chosen to assess the pink salmon populations in the Upper
Great Lakes (Figure 1). The two primary study streams were
Albany Creek (450 58'N, 84° 05'W), located on the northern
shore of Lake Huron in Chippewa County and the Laughing
Whitefish River (47° 38'N, 87° 02'W), located on the south
shore of Lake Superior in Alger County. These streams were
chosen through consultation with Wilbert Wagner, Fisheries
Division, Michigan Department of Natural Resources (MDNR),
an expert on Great Lakes pink salmon. Their selection was
based on: similarities in size and habitat composition: the
occurrence of abundant spawning populations of pink salmon:
and their utilization as MDNR study streams since the mid
1970's.

Within each study stream, specific study areas were
chosen. Selection of study areas within the streams was
based upon availability of pink salmon spawning habitat,
visibility and clarity of the water, and their history as
areas that contained a majority of spawners within the
stream (Wilbert C. Wagner, Personal Communication,
Fisheries Division MDNR). The length of the study areas

varied according to the amount of available spawning

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habitat but none were under 100 m in length.

The Laughing Whitefish River was approximately 18 km
long and the predominate substrates were cobble, gravel,
and bedrock throughout most of the river. A marsh area,
spanning 400 m upstream, near the mouth of the stream was
characterized by aquatic vegetation and woody debris on
sandy substrate. Pink salmon spawning occurred upstream of
the marsh area in riffle and runs in areas of gravel and
cobble substrate. This region of the river contains
resident populations of cyprinids and juvenile steelhead
(Salmg gairdneri) as well as a few coho salmon juveniles
(Q. kisutch): anadromous runs of steelhead, coho salmon,
and chinook salmon (Q. tshagytggha) were also present in
the stream (Appendix 1). Two study areas were established
in the Laughing Whitefish River (Figure 2). An upstream
study area was located approximately 4 km from Lake
Superior and was 225 m in length. This study area was
predominately riffle and run habitat with an abundance of
gravel and cobble, ranging from 2 to 300 mm in diameter.
Historically, approximately 80 % or more of adult pink
salmon present in the stream utilized this stream segment
for spawning (Bagdovitz 1985). This stretch was to be the
only study area on the stream. However, when pink salmon
populations on the Laughing Whitefish River declined,
spawning pink salmon failed to return to this upstream
spawning area. It was necessary, therefore, to select a

study area in a stream segment that was being utilized by

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pink salmon. The area was located 2.5 km upstream from the
river mouth, downstream from the historical spawning area,
and was 100 m in length. The downstream area was composed
predominately of bedrock substrate but contained pockets of
spawning gravel that are utilized by spawning pink salmon.

Albany Creek is approximately 9 km in length. At the
mouth, a marsh spanning 350 m upstream is characterized by
sandy substrate, woody debris, detritus and sparse aquatic
vegetation. Upstream from the marsh, gradient and velocity
increase and the predominate substrates are cobble, gravel,
and sand throughout in the mid-section of the river.
Approximately 4 km from the mouth, the upstream regions of
the creek are slowed by a complex of beaver dams and
contain a silt and detritus bottom. It was in the
midstream area of Albany Creek where the majority of pink
salmon spawning occurred. In this section the fish
community contains numerous cyprinids, brook trout
(Salvelinus fontinalis), juvenile steelhead (Salmg
gairdneri), coho salmon juveniles (Q. kisutgh) as well as
anadromous runs of coaster (lake run) brook trout,
steelhead, coho salmon, and chinook salmon (Appendix 1).
The study area on Albany Creek (Figure 3), was located
approximately 0.5 km upstream from the river mouth and was
700 m in length. The Albany Creek study area consisted
primarily of gravel riffles and runs and several log jam

complexes that form plunge pools.

13

 

 

 

\,

Albany Creek

 

Route M48

§E~Studyhrea
Y»
Q60
0° - Route M134

~2\
1 Kilometer Lo ke Huron

*1

 

 

 

Figure 3. Location of study areas on Alban Creek Chi ewa
County, Michigan. ' y ' pp

14

Secondary study streams were used to compare vital
statistics and abundance of pink salmon populations from
Lakes Superior, Huron, and Michigan. The secondary study
streams for Lake Superior were Harlow Creek (46° 38'N, 87°
28'W), the Little Garlic River (46° 38'N, 87° 30'W), and
the Rock River (46° 27'N, 86° 55'W)(Figure 1). Lake
Michigan secondary streams were the Black River (46° lO'N,
85° 25'W) and Thompson Creek (45° 54'N, 86° 30'W)(Figure
l). The secondary streams on Lake Huron were the Black
Mallard River (45° 35'N, 84° lO'W) and the Carp River (46°
05'N, 84° 40'W)(Figure 1).

Study areas within each secondary study stream were
selected by the criteria outlined for primary study
streams. All secondary study streams contained an
abundance of gravel and cobble substrates and an adequate
amount of riffles and runs, providing an abundance of
adequate spawning habitat. Most secondary study streams
were comparable in size to the primary study streams with
the exception of the Carp River which was substantially
larger than all other study streams. Secondary study
streams contained resident cyprinid populations and most
harbored runs of anadromous steelhead, coho salmon and
chinook salmon as well as resident juvenile steelhead and

coho salmon (Appendix 2).

METHODS

The field portion of this research was conducted
during two seasons: a fall assessment of spawning adults:

and a spring assessment of outmigrating fry.

Fall Sampling

Visual Surveys of Spawning Adults

Estimates of the relative abundance of spawning pink
salmon were made by visual surveys conducted on all primary
and secondary study streams. Visual surveys of the study
streams started on 25 August each fall, which is typically
one week prior to the stream entry of pink salmon
(Bagdovitz 1985). Primary and secondary study streams were
surveyed a minimum of once each week with primary streams
typically being surveyed two or more times per week.
Surveys were conducted by slowly walking in an upstream
direction and counting the number of pink salmon present in
the study area of each stream (Sheridan 1962: Wagner and
Stauffer 1982). Non-study areas of each stream were also
visually checked to insure that pinks were still using the
study area as their predominate spawning area. These areas

were surveyed at the beginning, at the peak, and at the end

15

16
of the spawning run. These surveys permitted the detection
of any large-scale changes that may have occurred in pink
salmon abundance between years, streams, and Lakes and
allow broader inferences to be drawn from observations made

from the primary study streams.

Population Estimates

Population estimates on the primary study sites were
made using modified Petersen mark-recapture population
estimates (Ricker 1975). Within each study area, pink
salmon were collected with electrofishing equipment and
marked with a fin clip. Marked fish were released back
into the streams slightly downstream from the point of
their initial capture. Recapture runs were started as soon
as the last group of fish marked was redistributed on
spawning areas, typically within 30 minutes of release.
Fish were collected and held on the recapture run in order
to measure length and weight, determine sex, and examine
the fish for any signs of injury. Upon completion of
measurements, the fish were released.

Electrofishing runs were conducted at the peak of the
run except in 1987 when weekly estimates were made in
Albany Creek. The estimation of the peak of the run was
made by examining records of past runs, the date of first
fish entry, and numeric increases observed in the visual
counts. In the Laughing Whitefish River, population

estimates were conducted at the upstream area in 1984 and

17
in both areas for the remainder of the study. In Albany
Creek, population estimates were made for the entire study
area in 1984 and 1986. However, in 1985 and 1987,
extremely high abundance of spawners made it necessary to
subsample the study area. In 1985, 100 m (14%) of the
Albany Creek Study Area were selected to represent the
entire study area. In 1987, two smaller study areas (10%)
were selected to represent the predominant habitat types in
the larger study area. One of
the areas was chosen as representative of the riffle areas
and measured 47 m in length and the other, chosen as
representative of the log jam complexes, measured 20 m in

length.

Vital Statistics: Adults

Vital statistics collected were total length (mm),
weight (g), and sex. Fish were sexed by external
inspection (presence of a hump), gonadal inspection, or
extrusion of gametes. During handling, pink salmon were
inspected for lamprey scars and other visible
abnormalities.

Pink salmon collected during population estimates in
primary study streams were measured for vital statistics.
Pink salmon were collected for vital statistics data by
single run electrofishing in the study areas of secondary
study streams. If only a few (<10) pink salmon were

collected in the study area, areas adjacent to the study

18
area were surveyed to collect additional fish for vital

statistics analysis.

Spawning Habitat Utilization

Surveys were conducted to determine the type of
habitat utilized by spawning female pink salmon. These
habitat utilization measurements were conducted in the
fall, primarily in the early portion of the spawning run
when pink salmon were spawning on substrates undisturbed by
previous spawners. When female pink salmon were observed
actively constructing redds during abundance surveys,
efforts to collect habitat measurements on these females
were initiated. Pink salmon redds were located again by a
second visual survey. Only those redds that where being
actively excavated or defended by the female were analyzed.
At these redd locations depth, velocity, and substrate were
measured in an undisturbed area immediately upstream from
redd construction. Habitat utilization variables were
collected using the methodology outlined in the Instream
Flow Incremental Methodology (IFIM) since these data will
be used for future modeling efforts (Bovee 1982). Depth
and velocity measurements were made using a top set wading
rod and Marsh-McBirney current meter. All depth and
velocity values were measured in english units since these
are the standard for the IFIM (Bovee 1982). Substrates
were visually categorized using standard Habitat

Suitability Index (HSI) methodology (Bovee 1986). The

19
predominate substrate was visually classified as belonging
to one of the designated substrate classes (ie. sand,
medium gravel, large cobble) standard for HSI studies
(Appendix 3). In addition, the percent embeddedness of
fine materials was visually classified using the
description (Appendix 4) and photographs of Bovee (1986).
Due to the relative abundance of spawning pink salmon,
habitat utilization measurements were made predominately in
Albany Creek with additional samples from the Laughing

Whitefish, Thompson Creek, and the Black River.

S Sa

Relative Fry Abundance

Data on outmigrating pink salmon fry were collected
primarily from Albany Creek and the Laughing Whitefish
River. Drift nets were fished immediately below the study
areas to intercept the fry on their outmigration. In order
to determine when the run started and to estimate abundance
throughout the run, sampling was started in mid to late
April. Fry sampling was terminated after catch rates fell
to 0 fish per day for five consecutive days.

Pink salmon fry were collected using drift nets that
were fished for 24 hours a day. These drift nets were made
of 0.48 cm regent nylon netting. The mouth of the net was
secured to a 61 cm by 122 cm frame made of stainless steel

conduit. The nets were 4.88 m deep and the cod and was

20
folded over and tied shut to close it. The net and frame
were placed in the stream and held in position by two 31 mm
cables that were anchored on both shores parallel to the
flow of the stream. One cable was run along the stream
bottom and the other cable was held in place approximately
1 m from the bottom of the stream. By securing the net to
both cables with rope, the net was fixed securely to the
bottom of the stream and mouth was held perpendicular to
the flow of the stream. The discharge was measured
immediately upstream from the nets in order to determine
the volume of water passing through the nets. This
measurement was made twice daily at each net check. From
the established relationship between surface area,
discharge, and the number of fry collected in each net, the
abundance of outmigrating fry was determined using the
methods of Acara and Smith (1971). Total abundance (A) was
calculated using the following equation:
A - S/Sn x C

where S is the cross-section area in square cm, Sn is the
area of the net opening in square cm, and C is the catch.

The nets were emptied twice daily, once an hour after
sunrise and once an hour before sunset. Before emptying
the net, all detritus and other debris were washed to the
cod end of the net by working the net in the current and
shaking debris from the side of the net. After working the
contents to the cod end, the net was untied, and the

contents of the net were emptied into a 190 liter gallon

21
plastic holding box. All fish sorted from the debris were
identified to species and enumerated. Pink salmon fry were
placed in a holding tank until all debris was sorted
through. All dead pink salmon fry were kept, and a random
sample of 50 % of live pink salmon fry were sacrificed for
measurement of vital statistics. The remaining fish were
released.

In an effort to compare pink salmon from different
tributaries and lakes, pink salmon fry from the Lake
Michigan tributary, the Black River were also sampled. The
Black River was sampled weekly during the 1985 fry
outmigration. These collections were made primarily for
comparison of vital statistics of the fry from different

lakes.

Vital Statistics: Fry

All pink salmon fry collected during relative
abundance estimates were measured for standard, fork, and
total length, to the nearest 0.001 mm using a
microdigitizing pad interfaced with a microcomputer. Wet
weights to the nearest 0.001 g were measured using an

electronic balance.

RESULTS

Fall

 

Visual Surveys of Spawning Adults

Visual surveys indicated a decline in Lake Superior
pink salmon populations from the onset of this study (Table
1). In the fall of 1984 an estimated 50 to 100 pink salmon
were counted in several primary and secondary study streams
in Lake Superior. In 1986, only the Laughing Whitefish
River and the Rock River contained pink salmon, and pink
salmon abundance was less than 25 fish. Spawning runs in
1985 and 1987 were also extremely poor. Fish were scarce
and difficult to detect, and pink salmon abundance was less
than 25 individuals in the Laughing Whitefish River, Harlow
Creek, and the Rock River. No pink salmon were seen in the
Little Garlic River after 1984.

Surveys on Lake Huron tributaries revealed increases
in pink salmon abundance in these streams from the onset of
the study (Table 1). In 1985, Albany Creek and the Carp
River had an estimated 10,000 spawners. The Black Mallard
River had a run of pink salmon numbering from between 500
to 1000 individuals, and this was the first run of pink
salmon recorded in this stream. In 1987, overall spawner

abundance was comparable to 1985 levels with increasing

22

23

Table 1. General trends in pink salmon abundance in
selected Michigan tributaries of the Great
Lakes. Abundance designations after Wagner and
Stauffer (l982):N-None: P-Present (1-99): C-
Common (100-999): A-Abundant (1000-9999): V—Very

Abundant (>9999).

 

 

Lake and Tributary
12§Z_

Lake Superior

Laughing Whitefish River
Harlow Creek

Little Garlic

Rock River

Lake Huron
Albany Creek

Carp River

Black Mallard River

£152.14.me
Black River

Thompson Creek

1984 1985

'U
'6sz

1986

’UZZ'U

'UZ'U'U

 

24
numbers of pink salmon in the Carp River and St. Mary's
River, while Albany Creek and the Black Mallard River
exhibited declines. It should be noted that water levels
were low in the Black Mallard River in the fall of 1987.
Many upstream areas were totally dewatered and the mouth of
the river was blocked off from the lake by sand from 18
September to 25 September probably forcing pink salmon to
migrate to another tributary for spawning. Lake Huron
tributaries surveyed in even years, 1984 and 1986, had 0 to
6 spawners present.

Both Lake Michigan tributaries exhibited increases in
spawner abundance in 1985 and the Black River showed
increases in 1987. These increases did not appear to be as
great as in Lake Huron tributaries, but populations in Lake
Michigan are continuing to increase. Spawners in the Black
River numbered approximately 5000 individuals in 1985. In
1987, the population size doubled with approximately 10,000
pink salmon spawners. Thompson Creek supported
approximately 1000 spawners in 1985 but less than 100 in
1987. The 1987 decline in Thompson Creek may have been a
true decline in the population or a result of a MDNR salmon
blocking weir placed in the stream prior to pink salmon
migration. The only pink salmon observed in any even year
runs on Lake Michigan tributaries was a single female on

Thompson Creek in 1986.

25
Population Estimates

Pink salmon abundance, as estimated by Petersen
estimates and electrofishing surveys, decreased
substantially from 1984 to 1987 in the Laughing Whitefish
River (Table 2). The spawning run of 1984 was the last
significant pink salmon run in the Laughing Whitefish
River. Even this run was relatively small with an
estimated 53 pink salmon in the upstream spawning area. A
few fish (< 10) were also observed in the downstream
spawning area, but no collections were made in this area in
1984. Numbers of pink salmon in the Laughing Whitefish
River study areas fell drastically after the 1984 survey.
No pink salmon were observed in upstream sections of the
stream and only very low numbers (< 10) were observed in
downstream sections. No pink salmon were collected in the
upstream or downstream study areas in 1985, 1986, or 1987.
Electrofishing collections made in non study areas below
the downstream study area did yield a total of 10 pink
salmon over the entire three year period.

In Albany Creek, pink salmon spawner abundance was
exceptionally high in 1985 (Table 3). A 100 meter reach of
the study section contained an estimated 298 pink salmon on
17 September 1985. Two years later, 17 September 1987, the
entire 700 m study area contained approximately 412 fish.
Despite the fact that the Albany Creek pink salmon were
more abundant in 1985, the 1987 pink salmon run was still a

large one.

26

 

 

 

Table 2. Population estimates of pink salmon abundance in
the Laughing Whitefish River.
Estimate
Year and Date Sitgf (95 % C. L.) # / 100 m
1984
17 September upstream 39 ( 16- 98) 20
30 September upstream l4 ( 7- 32) 7
1985
20 September upstream 0 0
20 September downstream 0 0
1986
26 September upstream 0 0
26 September downstream 0 0
1987
23 September upstream 0 0
23 September downstream 0 0

 

27

Table 3. Population estimates of pink salmon abundance in
Albany Creek (* - Michigan Department of Natural
Resources Sample: - Removed all fish in area).

 

 

 

Estimate
Year and Date Site (95 % C. L.) # / 100 m
1984
22 September study area 2 ( l- 4) 0.003
1985
18 September 100 m of
study area 298 (250-357) 298
1986
22 September study area 6* 0.009
1987
10 September study area 7 ( 3- 18) 0.01
17 September study area 412 (361-470) 0.59
24 September run area 148 (123-179) 314.89
24 September log jam 80 ( 61-104) 400
1 October run area 145 (121-174) 308.51
1 October log jam 60 ( 44- 81) 300
8 October run area 106 ( 86-132) 225.53
8 October log jam 33 ( 21- 51) 165
15 October run area 51 ( 38- 69) 108.51
15 October log jam 7 t 35
22 October run area 17 t 36
22 October log jam 0 t 0
29 October run area 0 t 0

29 October log jam 0 t 0

 

28

Petersen estimates of the entire study area in 1984
estimated only 2 pink salmon in the stream. In the fall of
1986 the Michigan Department of Natural Resources removed
all pink salmon in the stream prior to our collection.
Their collection efforts yielded 6 female pink salmon.

Mark-recapture data also illustrated the efficiency of
the visual surveys. When compared to mark-recapture
electrofishing, visual surveys underestimate the abundance
of pink salmon spawners in a tributary. However, the
visual estimates were typically within the range of the
lower limits of the 95 % confidence interval around the

population estimates (Table 4).

Vital Statistics: Adults

Measurements of mean total length and weight were
compared using a t-test to determine if the means were
significantly different between sexes (Steel and Torrie
1980). For most samples, differences in mean total length
and weight were found to be significant (P 5 0.05) with the
males being larger. Only in those samples with less than
10 fish per sex were differences non-significant. Thus,
males and females were treated as separate, distinct groups
for further statistical analysis.

To determine if mean total length and weight were
significantly different between sampling dates in the same
stream, a one-way analysis of variance was calculated in

conjunction with Tukey's multiple comparison test. The

29

Table 4. Comparison of mark-recapture population estimates
of pink salmon abundance compared to visual
surveys of abundance in Albany Creek and the
Laughing Whitefish River.

 

 

Tributary and Date 95 % C. L. Estimate
Count
Lowe U r
Laughing Whitefish River
17 Sept 84 16 98 39 12

30 Sept 84 7 32 14 12

Albany Creek

22 Sept 84 1 4 2 l
18 Sept 85 250 357 298 258
10 Sept 87 3 18 7 6
17 Sept 87 361 470 412 295
24 Sept 87 123 179 148 134
24 Sept 87 61 104 80 7O
8 Oct 87 86 132 106 88

8 Oct 87 21 51 33 28

 

30
only dates upon which mean total lengths and weights were
significantly different were for Albany Creek, the Black
Mallard River, and the Carp River for 1985. Pink salmon
collected in these streams during the early part of the run
were significantly larger (P < 0.05) than those collected
later in the run. The largest difference in mean size was
observed in the Carp River where male pink salmon averaged
460 mm and 802 g on 11 September and 429 mm and 613 g on 19
September. Length and weight differences observed between
dates in Albany Creek and the Black Mallard River were
smaller but still significant. The stream with the most
collections over the period of one spawning run was Albany
Creek in the fall of 1987 where pink salmon were collected
over a 7 week period. No significant differences were
observed between dates for Albany Creek in the fall of
1987.

The average size of pink salmon was compared between
streams in the same year using Tukey's multiple comparison
test (Appendix 5 and 6). Differences were observed between
streams within a lake and between streams flowing into
different lakes, but no consistent pattern was observed
over the course of the study.

In 1984 and 1986, the largest fish came from Albany
Creek and the Carp River, tributaries of Lake Huron (Table
5). In 1984, the difference in total length was
significant only in females with Albany Creek females

averaging 446 mm. However, in 1986, the difference was

31

Table 5. Average total length, weight, and number of adult
pink salmon by sex from selected tributaries of
the Great Lakes, 1984 and 1986.
limits are reported for sample sizes less than 4.

No confidence

 

 

 

1984
Lake and Tributary Sex
Lake Huron
Albany Creek M
F
Lake Superior
Laughing Whitefish River M
F
Harlow Creek M
F
1986
Lake and Tributary Sex
Lake Huron
Albany Creek
Carp River M
F
Lake Superior
Laughing Whitefish River M
F
Rock River M
F.

l2

17
19

PM

0%

Means i 95% C. L.
Lengthlmml Heightigi
480 1168
446 1 49 818 i 294
417 i 12 640 1 64
392 i 12 485 i 45
397 536
346 i 26 438 i 56

Means : 95% C. L.
MW
471 i 38 978 i 273
510 1032
515 1044
403 495
385 551
389 i 29 437 i 118
394 i 16 421 i 39

 

32
apparent in both males and females. The single male pink
salmon from the Carp River measured 510 mm and was
significantly larger than fish from Lake Superior
tributaries. The sole female pink salmon from the Carp
River was 515 mm in length while the average female pink
salmon from Albany Creek was only 471 mm in length. This
difference was not significant, but female pink salmon from
the Carp River were significantly longer than their Lake
Superior counterparts: the same relationship held true for
the average weights of males in 1984 and 1986 and for the
average weights of females in 1984. In 1986, the sole
female pink salmon from the Carp River weighed 1044 g but
was not significantly larger than females from Albany
Creek, the Laughing Whitefish River, and Thompson Creek.
The fish was larger than pink salmon from the Rock River
that averaged only 421 g.

Pink salmon in odd year runs from the Lake Michigan
tributaries of the Black River and Thompson Creek were
typically larger than fish from other tributaries. This
difference was more pronounced in males than in females
(Table 6). In 1985, male pink salmon from the Black River
averaged 488 mm and 1178 g and males from Thompson Creek
averaged 495 mm and 1170 g. No significant difference
existed between fish from these two Lake Michigan
tributaries, but the males were significantly longer and
heavier than pink salmon from tributaries of Lake Huron or

Lake Michigan. In the same year, female pink salmon from

33

Table 6. Average total length, weight, and number of adult
pink salmon by sex from selected tributaries of
the Great Lakes 1985.

 

 

Lake and Means i 95 % C. L.
Tributary Date Se_; N Length(mm) Weight(g)

 

Lake Huron

Albany Creek 9-11 M 46 433 i 10 708 i 96
F 25 418 i 13 590 i 86
9-18 M 138 426 i 5 -------
F 117 396 i 4 -------
Black Mallard River 9-16 M 21 427 i 16 601 i 59
F 15 415 i 17 590 i 98
9-26 M 58 422 i 12 557 i 55
F 54 382 i 5 414 i 20
Carp River 9-11 M 39 460 i 13 802 i 75
F 27 421 i 16 631 i 79
9-19 M 92 429 i 8 613 i 31
F 48 399 i 9 508 i 27
Lake Michigan
Black River M 47 488 i 12 1178 i 98
F 46 462 1 9 995 i 67
Thompson Creek M 57 495 i 8 1170 i 65
F 39 467 i 11 1015 i 76
Lake Superior
Harlow Creek M 16 408 i 19 598 i 67
F 4 429 i 57 630 i 188
Laughing Whitefish River F l 400 650
Rock River M 7 430 i 24 640 i 165
F 10 395 i 17 513 i 88

 

34

the Black River averaged 462 mm and 995 g and females from
Thompson Creek averaged 467 mm and 1015 g. No significant
difference in length or weight existed between females from
these Lake Michigan tributaries, but they were also larger
than any of their counterparts from Lake Huron or Lake
Michigan. No significant differences in length or weight
were noted between male pink salmon from tributaries of
Lake Huron and Lake Superior in 1985. Female pink salmon
from these tributaries did exhibit some significant
differences in average length but no trend was apparent and
no significant differences in average weight were noted.

Pink salmon collected in 1987 showed similar trends to
fish collected in 1985 but the lack of fish in some rivers
limits the number of comparison that can be made (Table 7).
The largest male pink salmon collected in 1987 came from
the Black River and averaged 467 mm and 880 g. No
significant difference was found between Black River males
and those from the Carp River that were 433 mm and 668 g.
The male Black River pink salmon were significantly longer
and heavier than their counterparts from Albany Creek and
the Laughing Whitefish River but no significant difference
was noted between Carp River males and their counterparts
from Albany Creek and the Laughing Whitefish River. Female
pink salmon from the Black River averaged 437 mm and 727 g.
Like the males, the female pink salmon were not
significantly larger or heavier than females from the Carp

River but they were significantly larger than all other

35

Table 7. Average total length, weight, and number of adult
pink salmon by sex from selected tributaries of
the Great Lakes for 1987.

 

 

Means and 95 % C. L.

 

Lake and Tributarv Sex N Length(mm) Weight(g)
Lake Huron

Albany Creek M 291 393 i 5 452 i 18
F 266 370 i 3 356 i 11
Carp River M 126 433 i 9 668 i 54
F 46 406 i 13 580 i 79

Lake Michigag
Black River M 83 467 i 12 880 i 75
137 437 i 9 727 i 53

Lake Superior
Harlow Creek F 4 377 i 12 318 i 46
Laughing Whitefish River M 2 377 i 70 350 i 381
F 4 373 i 16 358 1 198

 

36
females collected in 1987. All male and female pink salmon
from tributaries of Lake Huron and Lake Superior were not
significantly different in total length. In regards to
weight, some differences were noted but no trends were
consistent between both tributaries and sexes.

Samples within Lakes were pooled in order to summarize
length and weight data for pink salmon returning to
tributaries of the same lake (Table 8). Tukey's multiple
range test was used to compare the total lengths and
weights of male and female pink salmon between Lakes and
years (Appendix 7). This analysis revealed that female
pink salmon from Lake Michigan and even year fish from Lake
Huron were consistently the largest fish. However, males
of the same lake-year combinations were not significantly
different from most other lake-year combinations.

Data for each lake was additionally pooled by years
and compared utilizing a one-way analysis of variance and
Tukey's multiple comparison test. This pooling allowed
comparisons to be made between the lakes for mean lengths
and weights over the entire period of this study, 1984-
1987. Intotal length and weight, Lake Michigan pink
salmon were significantly larger than either Lake Huron or
Lake Superior fish (P < 0.05). Lake Huron pink salmon were
not significantly different than Lake Superior pink salmon
in length or weight.

The weight-length relationship for pink salmon was

calculated for individual streams by sex using the methods

37

Table 8. Average total length, weight, and number of adult
pink salmon by sex from Lakes Huron, Michigan and
Superior from 1984 to 1987.

 

 

 

 

 

Males Females

Means and 95 3 C. L. Meeee and 95 % C. L.
Lake
leaf. N Length(mm) Weigh§(g) N Leggth(mm) Weight(g)
Huron
1984 2 480 1168 7 446 i 49 818 1294
1985 395 431 i 4 652 i 30 286 400 i 3 517 i 24
1986 1 510 1032 6 471 i 38 978 i273
1987 454 405 i 4 520 i 22 437 376 i 3 391 i 15
Total 852 418 i 3 574 i 17 736 389 i 3 450 i 17
Michigan
1985 104 493 i 7 1174 + 56 85 464 i 7 1005 i 49
1986 - --- - ---- 1 441 733
1987 83 467 i 12 880 i 75 137 437 i 9 727 i 53
Total 186 482 i 7 1045 i 49 223 449 i 6 840 i 41
Superior
1984 18 416 i 12 627 i 60 26 397 i 60 468 i 34
1985 23 415 i 3 610 i 61 15 405 i 16 553 i 71
1986 9 392 i 21 450 i 99 10 393 i 14 434 i 46
1987 2 376 350 8 355 i 7 338 i 72
Total 52 410 + 8 567 i 43 59 388 i 8 466 i 31

 

38
of Ricker (1975). All regressions were tested using a F-
test to determine if a significant amount of the variance
had been explained by the regression and a t-test to
determine if the slope was significantly different from 0
(Sokal and Rohlf 1973). If both tests were significant (p
< 0.05), the regression was deemed valid. Nonsignificant
regressions were not reported. Regression formulas showed
a great amount of variance and the coefficient of variation
ranged from 0.41 to 0.96 for males and from 0.10 to 0.90
for females (Table 9). Data was pooled for each of the
Lakes in an attempt to summarize weight-length regressions
for each Lake in each year. These regressions were also
tested for significance as in the individual stream
samples. Only 2 of 7 regressions for Lake Superior were
significant: therefore, these data were not used for
comparisons or summary.

A chi square analysis was conducted to determine if
the sex ratios deviated significantly from a 1:1 ratio
(Table 10 and 11). Results from these statistical analyses
varied, with most samples not significantly different from
a 1:1 sex ratio. However, in 1985, the majority of samples
deviated significantly from a 1:1 sex ratio in favor of
males. In 1987, enough samples were taken weekly from
Albany Creek allowing a chi square analysis to be done for
each week throughout the spawning period (Table 12). The
sex ratio was skewed towards males in the early part of the

run and towards females in the later part of the run.

39

 

 

 

Table 9. Weight-length regression values for pink salmon
from selected tributaries of the Great Lakes.
Regression is in the form:
log weight - log a + b(1og total length).

Year / River a b 8:2 a gemalea

1984

Albany Creek n.s. -11.68 3.01 0.84

Laughing Whitefish R. - 9.60 2.66 0.90 n.s.

1985

Albany Creek -11.22 2.92 0.41 -16.86 3.84 0.67

Carp River 9-11 -10.06 2.73 0.85 -10.30 2.77 0.81

9-19 - 5.77 2.01 0.57 1.91 0.72 0.10

Black Mallard River I

9-16 - 7.93 2.36 0.72 -10.19 2.74 0.62
9-26 - 6.53 2.12 0.68 - 9.88 2.68 0.61

Black River -15.24 3.60 0.85 -14.47 3.48 0.83

Thompson Creek -12.47 3.14 0.77 -10.83 2.76 0.70

Harlow Creek - 8.01 2.39 0.74 n.s.

Rock River -l9.08 4.21 0.96 -14.27 3.43 0.88

1986

Albany Creek ---- 113.08 3.24 0.83

1987

Albany Creek - 4.27 1.73 0.41 - 6.27 2.05 0.35

Black River -l3.39 3.27 0.79 -12.81 3.19 0.66

Carp River -l3.23 3.24 0.89 -l4.06 3.39 0.90

 

40

Table 10. Sex ratio observed in pink salmon from selected
tributaries of the Great Lakes during even year

runs. A * denotes sex ratio is significantly

different from H :sex ratio is 1:1 using a chi

square test at 0995.

 

 

 

 

ea Stream Melee Females M:F Ratio
1984
Albany Creek 2 7 0.29 : 1
Laughing Whitefish River 17 19 0.88 : 1
Harlow Creek 1 7 0.14 : 1 *
1986
Albany Creek 0 6 -----
Carp River 1 l 1.00 : 1
Laughing Whitefish River 2 l 2.00 : 1
Rock River 7 9 0.78 : 1
Thompson Creek 0 1 -----

 

41

Table 11. Sex ratio observed in pink salmon from selected
tributaries of the Great Lakes during odd year
runs. A * denotes sex ratio is significantly
different from H°:sex ratio is 1:1 using a chi
square test at 0.95.

 

 

 

 

Year [ Stream Melee F ma 8 M:F Ratio
1985
Albany Creek 185 142 1.30:1*
Black Mallard 79 69 1.14:1
Black River 47 46 1.02:1
Carp River 131 75 1.75:1*
Harlow Creek 16 4 4.00:1*
Laughing Whitefish River 0 l -----
Rock River 7 10 0.70:1
Thompson Creek 57 39 1.46:1*
1987
Albany Creek 328 391 0.84:1
Black River 83 137 0.61:l*
Carp River 126 46 2.74:1*
Harlow Creek 0 4 -----

Laughing Whitefish River 2 4 0.50:1

 

42

Table 12. Sex ratio observed in pink salmon from weekly

samples from Albany Creek. An * denotes sex

ratio is significantly different from H°:sex
ratio is 1:1 using a chi square test at 0.95.

 

 

 

 

Date Melee Females M:F Ratio
10 September 3 2 1.50 : 1
17 September 179 126 1.42 : 1*!
24 September 71 84 0.84 : 1

1 October 31 54 0.57 : l

7 October 32 74 0.43 : 1*
15 October 10 36 0.27 : 1*
22 October 2 15 0.13 : 1*

 

43
During the mid point of the run, the sex ratio of males to
females was approximately 1:1.

All pink salmon collected were examined for any
external signs of parasitism or disease. Lamprey scars
were not seen on any pink salmon collected in 1984 or in
1986. Pink salmon did exhibit scarring in 1985 and 1987 in
tributaries to Lake Michigan and Lake Huron. Five pink
salmon were found with scars in 1987, 1 each from Albany
Creek (0.31 %), Black River (1.07 %), Black Mallard River
(0.68 %) and 2 from the Carp River (0.97 %). In 1987,
seven pink salmon with scars were found in Albany Creek
(1.26 %), five in the Black River (2.50 %), and none in the
Carp River.

In 1987 pink salmon were found in Albany Creek and the
Black River that exhibited an inflammation of the gills in
the isthmus area and frequently a mucosal buildup in the
gill arches. This condition is thought to be a form of
goiter (Sonstegard and Leatherland 1976: Leatherland and
Sonstegard 1987), although no biopsy was conducted. The
percentages of goiter occurrence were 2.69 % in Albany
Creek and 1.82 % in the Black River. Goiter was not

observed in any other pink salmon stocks analyzed.

Spawning Habitat Utilization
Habitat suitability data for spawning females were
collected primarily in Albany Creek with additional data

from Thompson Creek, the Black River, and the Laughing

44

Whitefish River. Mean values for depth, velocity, and
substrate were tested to determine if any differences in
habitat selection existed between streams. No difference
in mean depth or substrate chosen were found in the three
streams (Table 13). The mean velocity utilized at Thompson
Creek was significantly higher than was observed at Albany
Creek or the Black River (P < 0.05). This difference was
most likely due to the differences in gradient between the
three streams in the main spawning areas (Kocik 1988).

Frequency distributions for the three parameters were
constructed to illustrate habitat choice. Depths utilized
ranged from 0.20 feet to 3.90 feet, and over 80 % of redds
analyzed were in the water depth range of 0.50 to 1.50 feet
(Figure 4). Pink salmon utilized water velocities ranging
from 0.11 to 3.97 feet per second and over 70 % of redds
were constructed in water velocities ranging from 0.5 to
1.5 feet per second (Figure 5). Substrate was categorized
according to Bovee (1982, see Appendix 3 and 4). Substrate
utilized by pink salmon females for redd construction
ranged from sand (6) to large cobbles (13) (Figure 6).
Over 75 % of the redds were constructed in medium gravel
(9) to small cobble (12). The mean substrate value used
was 10.64 indicating substrate selection was skewed towards
gravel with relatively low percent embeddedness. Gravel
with a quartile 1 rating was utilized for 37.2 % of redds.
The percentage of redds then declined as the quartile

embeddedness ratings increased: quartile 2 (33.8 %),

45

Table 13. Average depth, velocity, and, substrate selected

by spawning pink salmon in selected tributaries
of the Great Lakes.

 

 

 

Parameter Albany Black Thompson
195 % C. L.) Creek River Creek
Depth (Feet) 0.89 i 0.04 1.05 i 0.14 0.89 i 0.14
Velocity

(Feet / Sec.) 1.02 i 0.06 0.89 i 0.98 1.28 i 0.18
Substrate

(Index Value:
Appendix 3) 10.72 i 0.12 10.08 i 0.27 10.47 i 0.48

 

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49
quartile 3 (24.2 %), and quartile 4 (4.5 %). The mean

percent embeddedness was 1.95.

Spring
Relative Fry Abundance

Pink salmon fry outmigrating from spawning tributaries
were enumerated in the spring of 1985 in Albany Creek and
the Laughing Whitefish River. No pink salmon fry were
collected in Albany Creek in the spring of 1985. In the
Laughing Whitefish River in 1985, deployment of drift nets
was delayed until 6 May due to extremely high spring flood
waters. Since 26 pink salmon were estimated to be
migrating downstream on this date, the onset of
outmigration is uncertain. Numbers outmigrating were
highest between 11 May and 15 May with the peak of
migration being on 13 May (Figure 7) The last pink salmon
outmigrating were collected on 16 May. A total of 324 pink
salmon were estimated to be outmigrating from the Laughing
Whitefish River during this time period.

In the spring of 1986, collection efforts were
concentrated on Albany Creek since the Laughing Whitefish
River had very few adult spawners the preceding fall.
Collection efforts were initiated in late April, a week
prior to any known pink salmon outmigration in the Great
Lakes in an effort to determine when outmigration began.
Despite early deployment, 49 pink salmon were estimated to

be outmigrating on the first day of fishing on 23 April.

50

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Pink salmon outmigration continued until 23 May. The peak
of pink salmon outmigration occurred between 27 April and 1
May with the most fish observed outmigrating on 29
April(Figure 8). A total of 5,380 pink salmon were
observed outmigrating from Albany Creek in the spring of
1986.

The Laughing Whitefish River was checked for fry
outmigration on 2 nights during the usual peak of pink
salmon outmigration. These samples yielded no pink salmon
migrating from this river in 1986. No population estimates
were made in the spring of 1987 due to the low abundance of
spawners in the Laughing Whitefish River and Albany Creek
in the fall of 1986. The streams were checked on two
nights during the normal peak outmigration periods in both
streams. Chinook salmon (Qe geneeygeehe) fry were
collected from both streams, but no pink salmon fry were

collected from either stream.

Vital Statistics: Fry

Length and weight data were collected from pink salmon
captured in the Laughing Whitefish River in 1985 and Albany
Creek and the Black River in 1986. Vital statistics data
were summarized for all pink salmon collected (Table 14).
Both length and weight data were compared by date using a
one-way ANOVA and Tukey's multiple comparison test.
Significant differences in both length and weight of pink

salmon on various dates were observed in the Laughing

52

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53

Table 14. Average total length, weight, and number of pink
salmon fry from selected tributaries of the Great

 

 

 

Lakes.
give; and Yea;

Laughing Albany Black
Vital Statistics Whitefish Creek River
(95 % C. L.) River 1985 1986 1986
Length (mm):
Total 31.96 1 0.29 31.94 1 0.11 32.64 1 0.25
Fork 30.99 1 0.28 31.06 1 0.11 31.66 1 0.24
Standard 28.02 1 0.27 28.48 1 0.10 28.74 1 0.24
Weight (g) 0.117 1 0.003 0.138 1 0.002 0.132 1 0.004

Weight-Length Regression:
log weight - log a + b(log total length)

a -8.8908 -7.9259 -7.8145
b 1.9453 1.7141 1.6557
r2 0.39 0.20 0.2557

 

54
Whitefish River in 1985. These differences in size were
erratic, and no overall trend of changing size was
observed. Multiple comparison tests for Albany Creek and
the Black River in 1986 showed no significant differences
(P < 0.05)in length or weight of pink salmon by date. The
weight-length relationship for pink salmon fry was not
strong since the coefficient of determination was low in
all cases, ranging from 0.20 in Albany Creek and the Black
River to 0.39 in the Laughing Whitefish River. All pink
salmon fry collected from the Laughing Whitefish River had
completely absorbed their egg sacs. Pink salmon fry
without complete egg sac absorption constituted 9.1 8 of
fry collected in Albany Creek and 8.8 % of all fry
collected in the Black River.

Pink salmon fry from the three rivers were compared to
determine if there were any significant differences in
total length and weight using a one-way ANOVA and Tukey's
multiple comparison test. No significant differences in
total length were observed between pink salmon fry from the
Black River, Albany Creek or the Laughing Whitefish River.
Although not significantly different, pink salmon fry from
Albany Creek were slightly heavier than Black River fry.
Pink salmon fry from the Laughing Whitefish River were
significantly smaller than pink salmon from Albany Creek

but not significantly different from Black River fry.

DISCUSSION

Fall

 

Spawner Abundance

Pink salmon spawning runs from the southwest basin of
Lake Superior reached a pinnacle in 1979, twenty years
after their initial detection in the watershed. In 1979,
several United States tributaries had 10,000 or more
spawners and 46 tributaries had documented populations of
spawning pink salmon (Wagner and Stauffer 1982). Since
1979, pink salmon spawning populations have been declining,
reaching levels comparable to the early 1970's by 1984
(Bagdovitz et a1. 1986). Bagdovitz et a1. (1986)
hypothesized that the even year-class of pink salmon would
begin to increase and become the predominant run of fish
since recruitment appeared to be better in those stocks.
However, my visual observations and mark-recapture
population estimates indicated that the decline in
abundance continued in both even and odd pink salmon year
classes in the southwestern basin of Lake Superior through
1987. Since 1984, populations have declined to critically
low levels of less than 20 spawners per tributary in all

primary and secondary study streams surveyed for both even

55

56
and odd year runs. However, pink salmon populations in the
northeastern basin of Lake Superior, predominantly in the
Canadian waters of the lake, have not gone through the
large scale decline in populations observed in the
southwest portion of the lake. Pink salmon populations in
this portion of the lake have leveled off from high levels
of abundance observed in the late 1970's, but pink salmon
continue to be abundant in the Canadian waters of Lake
Superior (Kwain 1987).

Pink salmon reached detectable levels in Lake Huron in
1969, approximately twelve years after they were documented
in Lake Superior (Wagner and Stauffer 1982). Their
populations started to expand rapidly in 1985, sixteen
years after initial detection with levels of abundance 10
to 20 time higher than any previous runs (Wilbert C.
Wagner, Personal Communication, Fisheries Division MDNR).
In 1987, a leveling off in abundance occurred. Some
streams had more spawners and others had less than in 1985,
but the general trend appears to be a stable to slightly
increasing pink salmon population in the United States
Waters of Lake Huron. Lacustrian abundance of these fish
in 1985 and 1987 was sufficiently large to create an
important sport fishery in the United States waters of the
lake (Kocik and Taylor 1987b: Rakoczy and Rogers 1988).
Creel surveys in 1987 estimated that nearly 29,500 pink
salmon were caught in the Lake Huron sport fishery, making
pink salmon the third most abundant salmonid in the Lake

57
Huron sport catch (Rakoczy and Rogers 1988). An additional
6,000 pink salmon were estimated to be caught in the St.
Marys River (John Schrouder, Personal Communication,
Fisheries Division, MDNR). Even year runs of pink salmon
in Lake Huron, although present, are not common and have
not shown any trends towards increasing and show signs of
decline. At present, the pink salmon runs in Lake Huron
appear to consist of strong odd year class and very weak
even year class.

Lake Huron pink salmon have not undergone the lake-
wide expansion that Lake Superior fish have. These fish
have only been documented in 5 streams in the United States
waters of the lake, and are generally limited to the
northern portion of the lake (Wagner and Stauffer 1982).
Populations in the lower peninsula are relatively low while
upper peninsula populations make up the majority of the
United States pink salmon population in Lake Huron. On the
Canadian side of Lake Huron, pink salmon are limited
predominately to the North Channel and Manitoulin Island
area (Collins 1975: Kwain 1987). Although expansion
appears much more limited, increases in abundance appear to
be following the pattern of relatively rapid and explosive
population growth exhibited by Lake Superior populations.
However, unlike pink salmon in the United States waters of
Lake Superior, populations in Lake Huron have not yet shown

signs of decline.

58

Lake Michigan pink salmon populations entered a period
of rapid population growth in the mid to late 1980's, only
12 years after their initial detection (Wagner and Stauffer
1982). Lake Michigan pink salmon populations increased in
1985 and again in 1987. Unlike pink salmon populations in
Lake Huron, these populations appear to still be in an
active phase of population growth. Even year pink salmon
runs have not yet reached detectable levels in most Lake
Michigan streams. Expansion to new spawning streams is
even more limited in Lake Michigan populations than in Lake
Huron fish. Lake Michigan pink salmon are only documented
in 3 streams in the Michigan waters of the lake, and are
generally limited to the northern portion of the lake
(Wagner and Stauffer 1982). Populations in the lower
peninsula are extremely low and distribution is spotty at
best.

Visual surveys on non-study streams indicated only a
very limited colonization of tributaries by pink salmon
since 1984. The most intensive surveying of tributaries
occurred in the fall of 1985. Lake Huron tributaries were
observed from the Thunder Bay River at Alpena northward to
the Straits of Mackinac. Lake Michigan tributaries were
checked from Thompson Creek eastward to the Mackinac
Straits and in some selected Lower Peninsula streams.
Colonization of new tributaries was limited considering the
relative abundance of fish in the Lakes and in major

spawning streams of relatively close proximity to

59
tributaries checked. Only one new spawning stream was
reported in Lake Huron, the Black Mallard River, and two in
Lake Michigan, Paquin Creek and the Cut River (Appendix 8).
No additional new spawning streams were noted over the
duration of this study.

An important aside to these surveys was the
observation that pink salmon were spawning along the
shoreline of Lake Huron. Shoreline spawning was observed
in 1985 and 1987 in the Les Cheneaux Island area of Lake
Huron. Pink salmon were seen excavating redds, guarding
them, and in the act of spawning. Some of the shoreline
spawning took place near culverts where temporal streams
and swamp drainages enter the lake, but substantial
spawning was also observed on shoals not adjacent to any
flowing waters. The utilization of these areas could be
important, even if spawning is only slightly successful,
since successful spawning in the lacustrian environment
could allow for further increases in abundance and range
expansion.

The unstable and changing abundances of pink salmon in
the Great Lakes present a complicated puzzle for
investigators and a difficult management situation for
biologists. The expansion of pink salmon populations is
probably the simplest parameter to explain, despite the
fact that the logistics of the expansion are somewhat
unclear. Pink salmon were introduced in a period of time

when Lake Superior's indigenous piscivore and planktivore

6O
populations were in a state of decline or already severely
depressed (Christie 1974). Under these conditions, pink
salmon were placed into an environment that was relatively
low in potential predators and competitors. Like many
exotic species, the pink salmon established itself slowly
and then its abundance began to increase rapidly in regions
with available habitat (Elton 1958). The increase in pink
salmon populations occurred in the late 1970's in Lake
Superior, the early 1980's in Lake Huron, and the mid to
late 1980's in Lake Michigan. Although the rate of
colonization has varied, an apparent trend towards
increases in abundance has been observed in all the lakes
(Emery 1981).

The declines observed in the Lake Superior populations
are more difficult to explain. One theory might be that
pink salmon exhibited a boom-bust population growth common
with many introduced animals (Elton 1958) and common to
many Great Lakes exotics (Christie 1974). This explanation
fails to account for the decline of pink salmon only in the
southwest basin of Lake Superior and not in the
northeastern portion of Lake Superior, Lake Huron, or Lake
Michigan. Bagdovitz (1985) proposed two hypotheses to
explain the decline in pink salmon populations in Lake
Superior: first, a decline in rainbow smelt (erezpe
moedeg), an important prey item of pink salmon, and,
secondly, high egg and fry mortality caused by low stream
flow during spawning. Although, little research has been

61
done in support of the declining forage base hypothesis:
it should be noted that a 90 % decrease in rainbow smelt
occurred in Lake Superior during the time period in which
Lake Superior pink salmon populations declined (Selgeby
1985). Kocik and Taylor (1987a) also proposed the latter
hypothesis to explain the decline in pink salmon
populations. By synthesizing information describing the
factors that control oncorhynchid recruitment in the
Pacific Northwest and integrating these factors with pink
salmon population trends and environmental variability in
the Great Lakes region, they determined that stream flow
could have a great impact upon pink salmon recruitment.
Particularly important is the hydrological stability of
tributaries that pink salmon utilize for spawning and
incubation. Tributaries that are predominantly controlled
by ground water seepage and have relatively high base flows
remain relatively constant environmentally, and, as such,
provide stable spawning and incubation conditions for pink
salmon. Tributaries that are controlled directly by runoff
tend to be subject to dramatic climatically induced changes
in flow and other environmental variables. These unstable
streams would have highly variable spawning conditions
dependent upon climatic conditions. Since the majority of
spawning tributaries in the southwest basin of Lake
Superior drain shallow soils that lie on top of bedrock
(Farrand and Bell 1984), the majority of these streams are

hydrologically unstable in nature. As such, Kocik and

62
Taylor (1987a) attributed the decline of the pink salmon
stocks in the United States waters of Lake Superior during
the early 1980's to the unstable stream environments in
this region combined with below normal precipitation events
during the 1979 spawning run. Since the initial decline,
pink salmon populations have failed to recover to levels of
abundance of the early 1970's, and some populations have
declined even further. Although the reasons for the
continued decline in pink salmon abundance and failure to
reestablish populations that have declined are unclear, the
following hypothesis has been suggested. At present, pink
salmon populations have been reduced to levels reminiscent
of the early to middle 1960's. These populations remained
relatively low and showed very little signs of increase
until the early 1970's. It is possible that pink salmon
populations have to build up to a critical level of
abundance before populations can begin to recover at a more
rapid rate. In addition, rainfall levels during the
spawning period were quite high in the early 1970's, and
the combined effects of a critical mass of spawners and
adequate rainfall may be needed for the Lake Superior
populations to recover and expand (Kocik and Taylor 1987a).
Tributaries utilized in northeastern Lake Superior, Lake
Huron, and Lake Michigan are typically either larger in
size or predominantly controlled by ground water inputs
(Kocik and Taylor 1987a). As such, tributaries in these

regions appear to provide a more stable environment for

63
pink salmon spawning and incubation under a variety of
environmental conditions.

An additional factor that may explain the boom-bust
population dynamics in Lake Superior and apparent stability
in Lakes Huron and Michigan is the genetic makeup of the
brood stock introduced to the Great Lakes. Pink salmon
introduced into the Great Lakes were from an odd year brood
stock of the Lakelse River, a tributary to the Skeena
River, in British Columbia (Arc 1979). It appears that
odd-year stocks of pink salmon from this region are better
adapted to mild southern environmental conditions than
harsher northern conditions (Beacham 1984). The adaptation
to a warmer environment may be related to the separation of
even-year and odd-year line during the last glaciation
(Aspinwall 1974). As a result, the genetic makeup of these
fish may be poorly suited to the harsh climates prevalent
in southwestern Lake Superior (Ward 1925).

Vital Statistics: Adults

The mean total length and weight of male pink salmon
was typically significantly larger than that of females.
This observation is in agreement with data from the Pacific
Ocean (Ricker 1962: Beacham and Murray 1985) and other
studies in the Great Lakes Region (Wagner 1985: Bagdovitz
et a1. 1986: Kocik and Taylor 1987b). Male and female pink
salmon from the Great Lakes are substantially smaller than

those from the Pacific Ocean (Bams 1974: Beacham and Murray

64
1985). Average lengths and weights were comparable to
other studies in the Great Lakes region (Wagner and
Stauffer 1975: Kwain 1982: Nicolette 1983: Wagner 1985).
Although differences did occur between streams and years,
no trends were apparent.

Significant differences in total length and weight
were noted between pink salmon collected from the same
river on different dates in only three cases. All three
cases are from tributaries of Lake Huron in 1985: Albany
Creek, the Black Mallard River, and the Carp River. In
each case, those pink salmon that were collected earlier
were significantly larger than those collected at a later
date. A detailed investigation of pink salmon in Albany
Creek was conducted in 1987 when weekly samples were taken
throughout the duration of the spawning run. In this
instance no significant differences were found in total
length or weight between the dates. Pink salmon from
different streams were also tested for significant
differences in total length and weight. Significant
differences were observed but no overall trend emerged from
the analysis. In the Pacific Ocean, stocks of pink salmon
from different rivers often exhibit significant differences
in length and weight (Beacham 1985). This phenomenon has
also been observed in the Great Lakes (Bagdovitz 1985:
Bagdovitz et al. 1986). Observed differences in these
vital statistics between streams imply that pink salmon

stocks from a single river may be acting as a cohesive

65
unit, feeding together and migrating together until
returning to a tributary for spawning. This theory is
supported by research of the genetic structure of Great
Lakes pink salmon stocks. Although the pink salmon in the
Great Lakes come from a common genetic stock, there appears
to be some preliminary signs of genetic divergence
occurring in the Great Lakes watershed (Gharrett and
Thomason 1987). Conversely, the significant differences in
vital statistics observed in fish collected from Lake Huron
tributaries on different dates would seen to preclude this
argument. It is, however, possible that pink salmon
collected were either part of a cohesive unit that strayed
from another tributary or part of a group from the same
tributary that acted as a separate group.

Comparisons of total length and weight were also made
of pink salmon from each of the three lakes between years.
No major trends were apparent from this analysis. When
analyzing the data pooled over the four year period, pink
salmon from Lake Michigan were significantly larger than
pink salmon from Lake Huron or Lake Superior. Although
pink salmon from Lake Huron were consistently larger than
those from Lake Superior, the difference was only
significant in 1986. These differences in size may be
attributable to the amount of forage available in each lake
(Christie 1974). This could explain why Lake Michigan fish
are larger than those from the other lakes. However, this

argument would suggest that pink salmon from Lake Huron

66

would be larger than those from Lake Superior, which is not
true. This fact may be explained by the relative abundance
of pink salmon in these Lakes. When pink salmon were
increasing in population size in Lake Huron but were still
at low levels, from 1979 to 1983, they were significantly
larger than pink salmon from Lake Superior (Wagner 1985).
However, when pink salmon densities increased in Lake
Huron, competition for food resources appears to have
precluded the advantage of having more available forage, as
a noticeable downward trend in the size of pink salmon
corresponding to increased populations was observed. The
same downward trend in the size of pink salmon has occurred
in each lake as populations increase (Figure 9).

Weight-length regressions for adult spawners showed a
great deal of variation both between streams and between
years. The coefficient of determination of many of the
regressions was relatively low. Bagdovitz (1985) also
documented a poor relationship between these two
parameters. The reason for the observed discrepancies is
probably a result of two factors. One factor that may be
responsible for the poor relationship is the relatively
narrow size spectra of pink salmon collected. When only a
small range of lengths is regressed by weights a poor
regression can result due to the variations in weights of
fish of similar sizes. Another related factor is the
varying stages of sexual maturity at the time of capture.

Since pink salmon may be green, carrying all of their

67

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o«m(\

68
gametes, or partially or totally spent, great variations in
weight would occur for the same fish over its stream
residency. This factor could greatly influence the
observed relationship between length and weight. In an
effort to sort out the variation that is related to sexual
condition, I attempted to categorize pink salmon by their
developmental stage in the field and then separate the
various stages for analysis. I tried this for five streams
and the results were the same for all of them. For
example, in the Black River in 1987 the overall average
coefficient of determination for females was 0.79: when
analyzed by developmental stage, coefficient of
determinations ranged from 0.29 to 0.95. Results were the
same for other samples: some developmental stages showed
improved coefficients of determination but the majority
showed no improvement. As such, no further work was done
with separating the fish for weight-length regressions.

The sex ratios observed in different tributaries were
typically not significantly different from a 1:1 male to
female ratio. In cases where the difference was
significant this difference is probably attributable to the
timing of the collection rather than the actual sex ratio
in the run, especially in cases where large samples were
taken. Weekly analysis of the sex ratio of pink salmon
entering Albany Creek in 1987 revealed that male pink
salmon were the first to enter the spawning stream, the sex

ratio was approximately 1:1 near the peak of the run, and

69
females dominated the run near the end of the run. This
corresponds to observations about pink salmon in the

Pacific Ocean (Scott and Crossman 1973).

Spawning Habitat

Habitat selection by spawning pink salmon was closely
related to that observed in other studies in the Pacific
Ocean watersheds (Kurko 1977: Wilson et al. 1981: Nadeeu
1984: Raleigh and Nelson 1985). Redd depth for Great Lakes
pink salmon ranged from 0.50 to 1.50 feet, which is
comparable to the most commonly utilized depths observed in
Pacific coast pink salmon redds (Figure 10). Pink salmon
were found to utilize water as deep as 4 feet for spawning
in west coast studies. The maximum redd depth observed in
this study was 3.6 feet, and redds were uncommon in water
deeper than 1.2 feet. Lack of utilization of these deeper
waters was probably not a function of habitat choice but
habitat availability: the streams where Great Lakes pink
salmon were observed spawning contain only limited water in
excess of 1.2 feet in depth. A similar observation was
made with the velocity utilized for spawning (Figure 11).
Pink salmon in the Great Lakes were seldom observed
spawning in water with a velocity greater than 1.8 feet per
second. Pink salmon in Pacific studies were found spawning
in water up to 5 feet per second but utilization dropped
off dramatically at 2.5 feet per second (Kurko 1977: Wilson

et a1. 1981: Nadeeu 1984). This discrepancy is probably

70

—— Kurko (1977)

........ wilson et 81. (1981)
----- Nadeeu (1984)
——-Kocik

 

UTILIZATION

 

 

DEPTH (FEET)

Figure 10. Habitat utilization curves for spawning pink
salmon in the Great Lakes and Pacific Ocean:
Depth.

71

— Kurko (1977)

-------- Wilson et 81. (1981)
Nandeau (1934)

—— Kocik

CD

UTILIZATION
CD

CD

 

O i 2 3 4 5
VELOCITY (FEET/SECOND)

Figure 11. Habitat utilization curves for spawning pink
salmon in the Great Lakes and Pacific Ocean:
Velocity.

72
due to the fact that velocities in this range were uncommon
in study streams during fall. Substrate selection was
similar in both Great Lakes and Pacific Ocean pink salmon
(Figure 12). Very coarse gravel (32-64 mm) was the
predominate spawning habitat in all studies. Pink salmon
in the Pacific Ocean utilized the substrate range from fine
gravel (2-8 mm) to small cobble (64-130) (Kurko 1977:
Wilson et al. 1981: Nadeeu 1984). In the Great Lakes the
selection range was much wider, ranging from sand to large
cobble (130-250 mm). In summary, Great Lakes spawning
tributaries appear to have comparable substrate conditions
to west coast streams, and, despite the fact that the
entire range of depth and velocity parameters utilized by
pink salmon are not available in the Great Lakes, adequate

depth and velocity conditions are available.

em
Relative Fry Abundance

In the Laughing Whitefish River only an estimated 324
pink salmon fry were outmigrating during the 1985 census.
This was a relatively low number considering the abundance
of spawners the previous year was an estimated 53 adults.
High spring flows that prevented early sampling of the run
may have washed some of the fry out of their redds and the
stream prematurely. Once in the lake, their chances for
survival would not be good since fry that exit spawning

tributaries prematurely typically have poor growth and low

73

-------- Wilson et al.(1981l
Nadeeu (1984)
— K08 1k

UTILIZATION

 

0o ' 2 4
SUBSTRATE INDEX

Figure 12. Habitat utilization curves for spawning pink

salmon in the Great Lakes and Pacific Ocean:
Index Numbers from Appendix 3.

Substrate.

74

survival (Foerster 1968: Bailey et a1. 1976: Simenstad et
al. 1976). Pink salmon outmigration from the Laughing
Whitefish River peaked on 13 May and was highest between 11
and 15 May. The peak period of migration corresponds with
outmigration rates in 1983 and 1984 (Bagdovitz 1985:
Bagdovitz et a1. 1986). The pink salmon outmigration in
1985 occurred over a shorter duration than in 1983 or 1984.
The 1985 outmigration lasted from 6 May to 16 May. In 1983
pink salmon outmigration started on 6 May and ended on 22
May and in 1984 it commenced on 1 May and did not end until
21 May (Bagdovitz 1985). Surveys during the peak migration
period on the Laughing Whitefish River in 1986 and 1987
failed to produce any outmigrating pink salmon fry.

Estimates of the relative abundance of pink salmon fry
outmigrating from Albany Creek in the spring of 1985
revealed no pink salmon outmigrating from the stream. Low
abundance was expected as only 2 pink salmon were estimated
to be in the 1984 spawning run. In the spring of 1986, an
estimated 5,380 pink salmon were observed outmigrating from
the stream. Although much higher than population estimates
in the Laughing Whitefish River, these estimates are
relatively low considering that the abundance of spawners
in 1985 was greater than 10,000 fish. Evaluation of
outmigrating pink salmon in the spring of 1987 revealed no
pink salmon fry outmigrating. Since no pink salmon were
collected in the main spawning area in 1986 no fry were

expected.

75

Utilizing my adult population estimates and sex ratios
and the fecundity data of Wagner (1985), I calculated the
potential egg deposition for spawning pink salmon in the
Laughing Whitefish River in 1984 and in Albany Creek in
1985. These calCulations reveal some insights into the
population dynamics of Great Lakes pink salmon. Potential
egg deposition for the 1984 Laughing Whitefish River was
28,056 eggs. Outmigrating fry numbered 324 or 1.15 % of
the potential egg deposition. In Albany Creek in 1985,
potential egg deposition was 4,648,012 eggs and the
resultant fry outmigration of 5,380 pink salmon fry
represented only 0.12 t of the potential egg deposition.
High mortality is common for pink salmon in the early
stages of their life history (Kocik and Taylor 1986a). The
differences in the survival rates in the two streams was
probably a result of low spawner densities in the Laughing
Whitefish River and extremely high densities in Albany
Creek. High spawner densities can have a detrimental
effect upon egg deposition and survival through a myriad of
factors such as spawner mortality (Wickett l958),increased
egg retention (Foerster 1968), displacement into
unfavorable spawning substrate (Kocik and Taylor 1986a),
redd superimposition (McNeil 1964), and streambed
overseeding (Heard 1978). A combination of these factors
probably led to the poor survival rate exhibited in the
pink salmon year-class from Albany Creek. It should be

noted that despite having a poor survival rate, Albany

76
Creek produced substantially more fry than the Laughing
Whitefish River. This fact is reinforced by the strong
return of adult pink salmon in Albany Creek in 1987,
compared to the weak return of pink salmon to the Laughing

Whitefish River in 1986.

Vital Statistics Fry

Great Lakes pink salmon fry are small when compared to most
pink salmon stocks in the Pacific Northwest that typically
range in size from 30 to 45mm fork length (Scott and
Crossman 1973: Healey 1980). Since the mean diameter of
eggs from the Great Lakes is slightly smaller than the
diameter of eggs from Pacific Ocean fish (Wagner 1985), and
egg size is frequently related to fry size in Pacific
salmon (Beacham et al. 1984), it would be expected that
Great Lake pink salmon fry would be smaller. No
significant differences in size were observed between fry
collected in this study and those collected by Bagdovitz
(1985) in three tributaries to Lake Superior.

Pink salmon fry outmigrating from Albany Creek, the
Black River, and the Laughing Whitefish River were not
significantly different in total length. Fry from the
Laughing Whitefish River weighed significantly less than
fry from Albany Creek but were not significantly different
from Black River fry. Thus, size differences in pink
salmon fry are minimal between the three lakes. Since all

pink salmon stocks are derived from a single stock of fish,

77
it is possible that either environmental or developmental-
stage differences are the cause of any observed deviations

in length or weight.

Ecological Role of Pink Salmop

The pink salmon was introduced to a Great Lakes
ecosystem that is constantly changing as a result of
natural and anthropogenic forces in the ecosystem (Scavia
et al. 1986). It has been successful in becoming an
established part of the Great Lakes ichthyofauna (Kocik and
Taylor 1987a). At present, the primary piscivores in these
ecosystems are salmonids, dominated by exotic species such
as the coho salmon, chinook salmon, brown trout eelpe
gppppe, and steelhead, and a re-established native fish,
the lake trout fielgelippe nemeyepeh. These salmonids have
become an established part of the ecosystem and a primary
part of the Great Lakes sports fishery (Scavia et a1.
1986). Since pink salmon feed upon rainbow smelt and
alewife (glee; pepegepegengee), forage fish that are
extensively utilized by the salmonid complex, the potential
for competitive and predatory interactions with these other
salmonid species exists (Kocik and Taylor 1987b). The
overlapping diets of pink salmon and the other salmonids
may result in direct competition for younger forage fish
and may reduce the abundance of these populations in future
years, thus adversely affecting the growth and survival of

various species in the Great Lakes salmonid complex.

SUMMARY

Since 1984, pink salmon populations have declined
substantially in the United States waters of Lake Superior.
This downward trend is a continuation of a decline first
noted in 1981. The reasons for this decline are unclear
but seem linked to the riverine phase of the pink salmon's
life history. In Lakes Huron and Michigan during the same
time period, pink salmon populations have increased. At
present, Lake Huron populations appear to have leveled off
at relatively high levels and Lake Michigan populations
appear to be on the increase.

Adult pink salmon spawned from late August to early
October. Utilization of spawning habitat is similar to
that chosen in Pacific Ocean populations. Slightly higher
water velocities and water depths were utilized by the West
Coast stocks due to the greater availability of these
habitats in those systems. Analysis of the size structure
of spawning pink salmon in the Great Lakes indicated pink
salmon males (average: 429 mm, 654 g) were consistently
larger than females (average: 407 mm, 536 9): similar
sexual dimorphism in size is common in the pink salmon's
native range. Size differences occur between lakes and

between streams but these differences appear to be linked

78

79
to environmental factors such as prey densities and lake
productivity. The average size of Great Lakes pink salmon
spawners is substantially smaller than their Pacific Ocean
counterparts.

The Great Lakes pink salmon's ecological utilization
of the stream environment is limited to spawning and
incubation of eggs and alevins since the fry migrate
directly to the Great Lakes environment in a manner similar
their Pacific Ocean counterparts oceanic migration. Fry
outmigration patterns followed those on the West Coast with
outmigration beginning in mid-April and ending in late May.
Fry sizes (average: 32.07 mm, 0.135 g) were slightly
smaller than pink salmon observed in West Coast runs.

In general, Great Lakes pink salmon are
morphologically smaller versions of their Pacific Ocean
counterparts exhibiting a similar migratory-pelagic ecology
and riverine spawning-incubation ecology. Pink salmon, the
first naturally reproducing Oncorhynchid in the Great
Lakes, appear to be a permanent component of the Upper
Great Lakes ecosystem. Since they feed upon the same foods
as other salmonids in the Great Lakes ecosystem the
potential for competition for food and space with these

species exists.

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APPENDICES

87

Appendix 1. Non-target fish species collected during
electrofishing in Albany Creek (AC) and the
Laughing Whitefish River (LWR) during spring
and fall sampling. An * denotes fish present
in a stream.

 

 

 

Streams

Spesies AC LWR
Sea Lamprey (Eetzomyzop peeippe) * *
Coho Salmon (Oncorhynchus kieppgh) * *
Chinook Salmon (Qpepppypepee pepepyeeepe) * *
Steelhead (ee1pg gelegpepi) * *
Brook Trout (mum 1283.18.11.15) *
Central Mudminnow (Umbze 11ml) * *
Rosyface Chub (Hypgpeie rubrifzgngl *

N. Redbelly Dace (Phgxiege e99) * *
Finescale Dace (Engxipge neggegng) * *
Longnose Dace (Bhiniethxs satarastee) * *
White Sucker (W W) * *
Northern Hog Sucker (Hypengelium nigrigenal * *
Burbot (Leta lots) *
Pumpkinseed (LQEQELE 91922525) *
Largemouth Bass (Mieggppexpe eelegigee) *
Johnny Darter (Ephegetgme nigrum) * *

Mottled Sculpin (Qgegge hairdi) * *

 

88

Appendix 2. Non-target fish species collected during
electrofishing in secondary study streams.

 

 

Black Mallard R.
Thompson Creek
Harlow Creek
Little Garlic R.

Carp River
Black River

Species
Sea Lamprey

Petromyzop marinus

Coho Salmon
Oncorhynchus kisutch * * t * 8

Mr
I-
I-
I»

Chinook Salmon
Oncorhynchus ts a scha * * * * *

Brown Trout
Salmo prutta * * *

 

Steelhead
Salmo geernee1 * e e t *

Brook Trout

Mavelnsmtmelis * * *

Central Mudminnow
932;; limi * e e

N. Redbelly Dace
Phoxipus eee * *

Finescale Dace
Phoeinue neogeeue *

Longnose Dace
s gategagtee * e e e e

White Sucker
__tQ§$_Ca omus W * * *

Northern Hog Sucker

Hypenteliue 1 ca 8 * *
Johnny Darter
Ethegspoma pigeep * * * *

Mottled Sculpin
Cottps beigdg * e e *

Rock River

89

 

 

 

 

Appendix 3. Generalized substrate classes for use in
studies to determine habitat utilization and
preference.

Li C1§§§,NQE§. Size Range (mm)

01 Organic Detritus

02 Vascular Plants

03 Attached Algae

04 Clay 0.00024 0.004

05 Silt 0.004 0.062

06 Sand 0.062 2.0

07 Very Fine Gravel 2 4

08 Fine Gravel 4 8

09 Medium Gravel 8 16

10 Coarse Gravel 16 32

11 Very Coarse Gravel 32 64

12 Small Cobbles 64 128

13 Large Cobbles 128 256

14 Small Boulders 256 512

15 Medium Boulders 512 1024

16 Large Boulders 1024 2048

17 Very Large Boulders 2048

18 Bedrock Plain, unfractured

19 Bedrock Plain, jointed

20 Bedrock Tilted, perpendicular

21 Bedrock Tilted, parallel

 

90

Appendix 4. Descriptions of substrate materials by
percentages of embeddedness to the nearest
quartile. From Bovee (1986).

 

 

 

Description % Fines Quartile
Openings between dominate sized particles 0-25 1

appear dark and are 1/4 to 1/2 the size of
the particles. Material seen through
openings are about the same size as the
dominate particles. Particle edges are
clearly discernable.

At least half of the openings between 25-50 2
dominate sized appear dark. Opening are

apparent, but less than 1/4 the size of

particles. Most particle edges are clearly

discernible, but up to half the edges are

obscured by fine materials.

Openings between dominant sized materials 50-75 3
appear to be completely filled with finer

materials. Less than half the edges of

dominate particles are clearly discernable,

but the size of the larger materials can

be determined without removing them from

the bed.

All openings between larger materials are 75-100 4
obscured. Bed appears to consist of fine

materials, but is solid to the touch. Only

one or two edges of the dominate particles

may be visible. Size of dominate particles

cannot be determined without removing them

from the bed.

 

91

Appendix 5. Results of Tukey's means test for total
length of pink salmon from separate streams
within each year. Streams with the same
letter have mean total lengths that are not
significantly different.

 

 

 

 

gear and Streams Melee Females
1984
Albany Creek A A
Laughing Whitefish River A B
Harlow Creek A ' B
1985
Thompson Creek A A
Black River A AB
Carp River B Harlow Creek ABC
Albany Creek B ABC
Rock River 8 Carp River BC
Black Mallard River B Laughing Whitefish River C
Harlow Creek B Rock River C
Black Mallard River C
198p
Carp River A Carp River A
Laughing Whitefish River B Albany Creek AB
Rock River B B
Laughing Whitefish River 8
1291
Black River A A
Carp River AB AB
Albany Creek 8 Harlow Creek B
Laughing Whitefish River B Rock River B
Albany Creek B

 

92

Appendix 6. Results of Tukey's means test for weight of
pink salmon from separate streams within
each year. Streams with the same letter
have mean total lengths that are not
significantly different.

 

 

 

 

 

_ear and Streams Males Femalee_
1984

Albany Creek A A

Laughing Whitefish River B

Harlow Creek A B
1985

Black River A Thompson Creek A

Thompson Creek A Black River A

Albany Creek 8 Laughing Whitefish River 8
Carp River B Harlow Creek B
Rock River B Albany Creek 8
Harlow Creek B Carp River B
Black Mallard River B Rock River B
Black Mallard River 8
12§6
Carp River A Carp River A
Laughing Whitefish River 8 Albany Creek AB
Rock River B Thompson Creek AB
Laughing Whitefish River AB
Rock River B
1287
Black River A A
Carp River AB AB
Albany Creek BC Laughing Whitefish River BC
Laughing Whitefish River C Albany Creek BC

Harlow Creek C

 

93

 

 

 

 

Appendix 7. Multiple comparison analysis of pink salmon
total length by Lake and Year.
Meles Total Length Females
Lakee Year Grouping, Lake_, Year
Grouping
Huron 86 A A
Michigan 85 AB AB
Huron 84 ABC ABC
Michigan 87 ABCD Michigan 86 ABC
Huron 85 ABCD Michigan 87 ABCD
Superior 84 ABCD Superior 85 BCDE
Superior 85 ABCD Huron 85 BCDE
Huron 87 BCD Superior 86 CDE
Superior 86 CD Huron 87 DE
Superior 87 D Superior 87 DE
Superior 84 E
Males Weight Females
Lek—_e Y__rea 9.1922189 LQISL_ Year. won 1
Michigan 85 A A
Huron 84 A Huron 86 A
Huron 86 AB Huron 84 AB
Michigan 87 ABC Michigan 87 ABC
Huron 85 ABC Michigan 86 ABCD
Superior 84 ABC Superior 85 BCDE
Superior 85 BC Huron 85 BCDE
Huron 87 BC Superior 84 BCDE
Superior 86 C CDE
Superior 87 C Huron 87 DE
Superior 87 E

 

Appendix 8.

94

study during fall spawning runs.
designations after Wagner and Stauffer

(l982):N-None: P-Present (1-99): C-Common
(loo-999): A-Abundant (1000-9999): V-Very

Abundant (>9999).

List of streams surveyed over duration of

Abundance

 

 

Lake 1 Stream

Lake Huron

Lake

Lake

Elliot Creek
Grand Lake Outlet
Greene River
Little Trout River
Mill Creek

North Devil River
Ocqueoc River
Saint Mary's River
South Devil River
Swan River
Thunderbay River
Trout River

Michigan

Cut River
Davenport Creek
Manistique River
Ogontz River
Paquin Creek
Rapid River
Stony Creek
Tacoosh.River
White River

Superior

Anna River

Dead River
Huron River

Two Heart River

1984

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