YELLOW PERCH, PERCA FLAVESCENS, BEHAVIOR IN THE INDIANA WATERS
OF LAKE MICHIGAN IN 2009, 2011, AND 2012
A THESIS SUBMITTED TO THE GRADUATE SCHOOL IN PARTIAL
FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF
SCIENCE.
BY DAVID A. STARZYNSKI
ADVISOR THOMAS E. LAUER
BALL STATE UNIVERSITY
MUNICE, IN
JULY 2013
ii YELLOW PERCH, PERCA FLAVESCENS, BEHAVIOR IN THE INDIANA WATERS
OF LAKE MICHIGAN IN 2009, 2011, AND 2012
A THESIS SUBMITTED TO THE GRADUATE SCHOOL IN PARTIAL
FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE
MASTER OF SCIENCE
BY
DAVID A. STARZYNSKI
ADVISOR: THOMAS E. LAUER
Committee Approval:
______________________________________________________
Committee Chairperson
____________
Date
______________________________________________________
Committee Member
____________
Date
______________________________________________________
Committee Member
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Date
Departmental Approval:
______________________________________________________
Departmental Chairperson
Date
______________________________________________________
Dean of Graduate School
____________
Date
BALL STATE UNIVERSITY
MUNCIE, INDIANA
JULY 2013
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iii TABLE OF CONTENTS
TABLE OF CONTENTS.......................................................................................... iii
LIST OF FIGURES .................................................................................................. iv
LIST OF TABLES ......................................................................................................v
ACKNOWLEGEMENTS ......................................................................................... vi
CHAPTER 1
ABSTRACT ................................................................................................................1
INTRODUCTION ......................................................................................................2
METHODS .................................................................................................................4
RESULTS ...................................................................................................................6
DISCUSSION .............................................................................................................8
REFERENCES .........................................................................................................11
FIGURES AND TABLES ........................................................................................15
CHAPTER 2
ABSTRACT ..............................................................................................................21
INTRODUCTION ....................................................................................................22
METHODS ...............................................................................................................24
RESULTS .................................................................................................................27
DISCUSSION ...........................................................................................................29
REFERENCES .........................................................................................................32
FIGURES AND TABLES ........................................................................................36
iv LIST OF FIGURES
Figure
Page
CHAPTER 1
1.
Locations of yellow perch sampling zones near Michigan City (1), Burns
Harbor (2), and East Chicago (3) in the Indiana waters of Lake
Michigan. ...............................................................................................................15
2.
Logistic spawning curves for female and male yellow perch collected
from the Indiana waters of Lake Michigan in 2009, 2011, and 2012. The
dashed line marks the point of peak spawn (50% spent). The date
indicates the day at which peak spawn occurred ...................................................16
3.
Lake Michigan mean bottom temperatures by day for 2009, 2011, and
2012........................................................................................................................17
4.
Lake Michigan mean bottom temperature comparison for the sampling
periods in 2009, 2011, and 2012. The 2009 and 2011 temperatures were
statistically similar (a) while 2012 was higher (b).. ...............................................18
CHAPTER 2
1.
Locations of yellow perch sampling zones near Michigan City (1), Burns
Harbor (2), and East Chicago (3) in the Indiana waters of Lake
Michigan. ...............................................................................................................36
v LIST OF TABLES
Table
Page
CHAPTER 1
1.
Abundance of mature yellow perch captured in gill-nets fished in Lake
Michigan during 2009, 2011, and 2012 .................................................................19
CHAPTER 2
1.
Abundance of mature yellow perch captured in gill-nets fished in Lake
Michigan during 2009, 2011, and 2012 .................................................................37
2.
Results of multiple regression model for yellow perch collected within
the Indiana waters of Lake Michigan in 2009, 2011, and 2012 with
CPUE as the response variable ..............................................................................38
3.
Mean length ± SE of yellow perch collected in the Indiana waters of
Lake Michigan in 2009, 2011, and 2012 ...............................................................39
4.
Results of multiple regression model for yellow perch collected in the
Indiana waters of Lake Michigan in 2009, 2011, and 2012 with the
response variable (length) and corresponding P-values for each predictor ...........40
5.
Results of the logistic regression for yellow perch collected in the
Indiana waters of Lake Michigan in 2009, 2011, and 2012...................................41
vi ACKNOWELDGEMENTS
I would like to thank all those who helped with the completion of my thesis
project. I thank the many Ball State students who spent their time on Lake Michigan
collecting samples especially Kip Rounds and Nick Haunert. I also thank Dr. Thomas
Lauer for his guidance and recommendations over the last two years. I thank the Indiana
Department of Natural Resources for the use of their facilities and I thank Ball State
University and Indiana DNR for funding. I finally thank my family and friends for their
help and support throughout my career.
Chapter 1: Yellow perch Perca flavescens spawning behavior in the Indiana waters of
Lake Michigan in 2009, 2011, and 2012.
Abstract – Experimental gill nets were used to collect adult yellow perch from the
Indiana waters of Lake Michigan during 2009, 2011, and 2012 to estimate the length and
timing of the yellow perch spawning season. Sampling was conducted at three zones near
East Chicago, Burns Harbor, and Michigan City from May to August in 2009 and 2011;
however, sampling was conducted twice in April before continuing from May to August
in 2012. After collection, yellow perch were measured, counted, sexed, and maturity state
was determined. Logistic regression was then used to model the proportion of post-spawn
(spent) fish over time (days) for each year. The resulting models estimated the timing and
duration of yellow perch spawning and allowed me to determine the timing of peak
spawn (50% spent) for each year. I found that yellow perch spawning started when water
temperatures reached ~ 11⁰C and lasted 2-4 weeks. In 2009 and 2011, spawning started
mid-May and was finished early June, although in 2012 spawning started in early May
and finished before June. These differences are likely in response to unseasonably warm
lake temperatures caused by record high air temperatures in the Great Lakes region. My
findings suggest global climate change will alter yellow perch reproduction cycles;
however, the specific influences on survival are unknown.
Introduction
Yellow perch, Perca flavescens, are native to Lake Michigan and have been
important to the recreational and commercial angler since the late 1800s (Eshenroder et
al. 1995). Easy accessibility and angler preference made yellow perch the most
recreationally harvested fish in Lake Michigan in the 1980s and 1990s (Bence & Smith
1999). Historically, a large scale commercial fishery was in place for this fish, but major
declines in abundance prompted closure of the commercial fishery in the main body of
the lake by 1997 and reduced the daily bag limit for recreational anglers (Marsden &
Robillard 2004; Wilberg et al. 2005). Despite these changes, there has not been
resurgence in the yellow perch population abundance since that time (Headley & Lauer
2008). Commercial fishing was considered the primary cause of the yellow perch
population decline by selectively harvesting female fish, which skewed sex ratios and
reduced the spawning stock biomass (Lauer et al. 2008). Currently, a limited commercial
fishery for yellow perch exists, but only in Green Bay and is relatively small (generated
$160,000 in 2008; USGS 2008). Therefore, exploitation does not appear to be the only
factor contributing to the poor recruitment of yellow perch in Lake Michigan.
Abiotic and biotic influences on yellow perch recruitment have been a focal point
for research in Lake Michigan (Clapp & Dettmers 2004). Phosphorous control programs
decreased phosphorous loadings into Lake Michigan by the early 1990s and a
simultaneous decrease in primary production occurred in the near shore waters of Lake
Michigan (Madenjian et al. 2002) Decreases in primary production likely reduced critical
larval yellow perch prey (e.g., rotifers, copepods) which could lead to low year-class
2 strength (Fulford et al. 2005). Further, Dreissena mussels have altered the Lake
Michigan ecosystem and could negatively affect yellow perch recruitment through
changes in the structure of lower trophic levels, specifically, the phytoplankton and
zooplankton communities (Dettmers et al. 2003; Marsden & Robillard 2004; French III et
al. 2007). Yellow perch and alewife (Alsoa pseudoharengus) abundances were negatively
correlated (Shroyer & McComish 2000; Forsythe et al. 2013) likely due to predation of
larval yellow perch by adult alewife (Brandt et al. 1987). Thus, a variety of ecological
effects appear to be influencing yellow perch recruitment, illustrating the need to
understand more of yellow perch life history (e.g., migration, reproduction) in Lake
Michigan to better manage the fishery (Clapp & Dettmers 2004; Marsden & Robillard
2004).
Yellow perch spawning behavior, specifically duration, timing, and thermal
triggers have been minimally studied in the Great Lakes (Robillard & Marsden 2001;
Brown et al. 2009; Collingsworth & Marschall 2011) compared to smaller systems
(Forney 1971; Smith 1979; Brown et al. 2009). Yellow perch typically spawn in early
spring when water temperatures reach 7 to 13 ⁰C following an extended period of
temperatures < 10 ⁰C (Hokanson 1977; Kolkovski & Dabrowski 1998; Brown et al.
2009). Spawning duration for yellow perch can be from one to three weeks; however,
timing can change based on geographic location and local environmental factors
(Hokanson 1977; Thorpe 1977). Yellow perch in northern latitudes have a shorter
spawning duration that is later in spring compared to southern populations (Thorpe
1977). Populations in the western basin of Lake Erie spawned when mean lake bottom
3 temperatures were between 11 and 15 ⁰C (Collingsworth & Marschall 2011). Thus,
spawning season and temperature thresholds appear to vary for individual populations of
yellow perch.
Although overexploitation and exotic species interactions were identified as
recruitment limitations for yellow perch (Wilberg et al. 2005; Forsythe et al. 2012), other
factors may also influence recruitment. Unfortunately, our lack of understanding in this
area is exasperated by a limitation in basic yellow perch life history in southern Lake
Michigan. Spawning behavior in these waters is relatively unknown compared to smaller
lakes (Forney 1971; Smith 1979; Brown et al. 2009), and spawning habitat limitations
have not been thoroughly studied (Walters 2010). Further, temperature is intimately
related to yellow perch life history (Hokanson 1977; Kolkovski & Dabrowski 1998;
Brown et al. 2009), but research defining temperature influences in southern Lake
Michigan are scant (see Rydell et al. 2010). Thus, yellow perch life history must be
investigated to better understand potential limitations to recruitment and future impacts
on the fishery.
The goal of my study was to better understand spawning behavior of yellow perch
during the spawning season in southern Lake Michigan. My objective was to identify the
timing and duration of the yellow perch spawning season in southern Lake Michigan. For
this objective, I evaluated whether mature fish were ripe (gravid) or recently spent. By
identifying this period, I calculated the date of 50% spent fish which I used to establish
peak spawn. I hypothesized that yellow perch spawning would begin mid-May, run
several weeks, and be completed by the end of June. The second objective was to
4 determine whether timing and duration of the spawning season were consistent among
2009, 2011, and 2012. I was specifically interested whether differences existed among
years, and if so, were they related to variations in temperature. I hypothesized that
spawning timing and duration would be consistent among all years.
Methods
Study Site – My sampling sites were located in the Indiana waters of southern Lake
Michigan and were primarily comprised of shallow waters (< 20 m) with uniform sandy
substrates (Janssen et al. 2005). The Indiana waters were divided into three sampling
zones, one each near East Chicago, Burns Harbor, and Michigan City (Figure 1). Each
zone spanned a 12 km long section of lakeshore and was divided into 1 km long sampling
frames. On each day, one frame was randomly selected for sampling from the 12 possible
frames in each zone.
Field Methods – Yellow perch were sampled in 2009, 2011, and 2012. In 2009, samples
were collected daily from May 4th to June 4th and then once per week after June 4th until
the yellow perch population reached 100% spent or when spawning had ended (Walters
2010). In 2011, samples were collected twice a week from May 10th to June 20th and then
once per week from June 20th to August 1st. In 2012, sampling was initiated on April 13th
and 14th in response to a perceived early season warming trend and anecdotal
observations. Twice-weekly sampling resumed from May 1st to June 21st and then once
per week from June 21st to July 31st. Sample collection meant collecting a single sample
from each zone resulting in three samples per date. Sampling was occasionally altered
(typically delayed) based on existing weather conditions. Yellow perch were collected
5 using graded-mesh gill nets containing nine, 15.2 m panels of repeating 51, 64, and 76
mm stretch-mesh for a total length of 137 m and depth of 1.8 m. These mesh sizes
effectively collect mature yellow perch > 175 mm that were required for my study
(Thomas 2007). Gill nets were fished parallel to shore at a depth contour between 5 and
20 m, set before sunset and soaked overnight for ~ 12 hours. The following morning,
yellow perch were removed from the net, immediately placed on ice, and then transported
to the Indiana DNR field station in Michigan City, IN for processing. Onshore, yellow
perch were counted, measured (total length), sexed, and maturity state was determined
using a modified gonad stage index that included gravid, running, and spent categories
(Treasurer & Holliday 1981). Temperature profiles were taken when gill nets were
collected using a YSI ProDO Professional Series digital unit from the surface to the
bottom at 1 m increments.
Statistical Methods – Logistic regression was used to estimate peak spawning season by
modeling the proportion of fish in the spent maturity state (response variable) from time
in days (predictor variable; Sigma Plot 9.0). Using this analysis I was able to identify
when 50% of the population reached the spent maturity stage, which I labeled as peak
spawn. A logistic regression was chosen because it best models the nonlinear relationship
between my two binary variables (gravid and spent). Yellow perch sexes were separated
for all analysis, as the timing and duration of males and females differs (Kolkovski &
Dabrowski 1998). Spawning season was measured as the date I first collected spent
females or males (start) and the date I last collected gravid females or males (finish).
6 The temperature of peak spawning was identified using mean lake bottom values
on the date of peak spawn for each sex. I used multiple regression through a generalized
linear model function to identify differences in mean bottom temperature (response
variable) among day (Julian), year, and day* year interactions (predictor variables) (SPSS
19.0). A post hoc Fisher’s least significant difference pairwise comparison analysis was
used to determine whether temperature values among years varied significantly. For all
tests, α was set at 0.05.
Results
A total of 3083 yellow perch was collected from the Indiana waters of Lake
Michigan during the 2009, 2011, and 2012 sampling seasons (Table 1). Female fish (N =
2551) comprised 83% of the total with ranges varying in size from 166 to 355 mm
(2009), 132 to 369 mm (2011), and 146 to 360 mm (2012) total length. Male yellow
perch (N = 532) ranged in size from 160 to 325 mm (2009), 127 to 315 mm (2011), and
96 to 301 mm (2012) total length. Ninety percent of females and 30% of males were
collected post-spawn and considered spent.
Yellow perch spawning duration and timing were identified using the observed
maturity stage of collected fish. Yellow perch spawning ranged between two to four
weeks among all sampling years based on collection of gravid and spent fish. In 2009,
female spawning took place in a 17 day period with half the spawn occurring between
May 18th and May 23rd and peak spawn on May 20th (Figure 2; Table 2); male spawning
took place in a 21 day period with half the spawn occurring between May 18th and May
31st and peak spawn on May 23rd (Figure 2; Table 2). In 2011, female spawning was
7 extended to 27 days, with half the spawn occurring between May 23rd and May 29th and
peak spawn on May 26th (Figure 2; Table 2); male spawning took place in a 26 day
period with half the spawn occurring between June 1st and June 14th and peak spawn on
June 8th (Figure 2; Table 2). In 2012, female spawning took place in a 23 day period with
half the spawn occurring between May 4th and May 12th and peak spawn on May 8th
(Figure 2; Table 2); male spawning took place in a 22 day period with half the spawn
occurring between May 31st and June 8th and peak spawn on June 3rd. However, males
were ripe from May 2nd until completion of spawning in June indicating the potential to
spawn any time after May 2nd (Figure 2; Table 2).
Lake temperatures varied daily, however a general increase in temperature
occurred throughout the spawning season in all years (Figure 3). In 2009, the yellow
perch spawning season began at 10.9 ⁰C, peaked (50% females spent) at 11.6 ⁰C, and was
completed by 14 ⁰C. In 2011, the yellow perch spawning season began at 11.2 ⁰C,
peaked at 11.5 ⁰C, and was completed by 13.3 ⁰C. In 2012 the yellow perch spawning
season began at 11.6 ⁰C and peaked at 13.2 ⁰C, however during the last two days of the
spawning season temperature fell to 12.3 ⁰C due to an influx of cold bottom waters.
Although 2012 water temperatures were significantly warmer on a given date than 2009
or 2011, yellow perch still began spawning ~ 11 ⁰C. The average temperatures when
spawning began varied by < 1 ⁰C throughout all sampling years and suggests that yellow
perch spawning in the Indiana waters is triggered by water temperatures ~ 11⁰C. Mean
bottom temperatures varied significantly by year (P = 0.001, df = 2, Likelihood ratio =
13) and by day (P < 0.001, df = 1, Likelihood ratio = 68) and the post hoc pairwise
8 comparison suggested that temperatures in 2009 and 2011 (P = 0.39) were similar, while
in 2012, temperatures were significantly warmer (P < 0.001) (Figure 4).
Discussion
My results allowed me to estimate spawning timing and duration for yellow perch
in 2009, 2011, and 2012, and to determine whether timing and duration of the spawn
were consistent among sampling years. As hypothesized, yellow perch spawning in
southern Lake Michigan began in spring and lasted between two and four weeks each
year. However, my hypothesis of consistent spawn timing in 2009, 2011, and 2012 was
not supported. In 2009 and 2011, spawning began mid-May and was completed by midJune, but in 2012, spawning began at the start of May and was completed before June.
The 2012 spawn was earlier by approximately two weeks. The unexpected shift was
likely caused by warmer than normal temperatures in the region, as 2012 had the warmest
spring on record for Indiana, Illinois, Michigan, and Wisconsin (NOAA 2013). The
temperature range of spawning in this study was consistent with others (Forney 1971;
Tsai & Gibson 1971; Brazo et al. 1975; Hokanson 1977; Collingsworth & Marschall
2011). However, spawning timing appears to be later in southern Lake Michigan
compared to other areas throughout the yellow perch range, likely because the large water
volume of Lake Michigan warms relatively slowly (Tsai & Gibson 1971; Brazo et al.
1975; Hokanson 1977; Collingsworth & Marschall 2011).
Duration of yellow perch spawning differed for females and males within a given
year, with females having a shorter spawning period. This difference may be inherent in
the biology of each sex, as females spawn in a single event and males remain able to
9 spawn throughout the spawning season and in multiple events (Kolkovski & Dabrowski
1998). In my sample area, half of the sampled yellow perch females spawned in single 58 day period, while half of sampled yellow perch males spawned in 9-13 days. My
findings and others (Dorr 1982; Brown et al. 2009) imply that males arrive on spawning
grounds prior to females, and males remain for the duration of the spawn. For my study
the total duration of yellow perch spawning was similar among years and within the
expected 2-4 week period (Hokanson 1977). Similarly, Dorr (1982) found yellow perch
spawning in southeastern Lake Michigan ranged from 1-2 weeks in late May and early
June.
Temperature appears to be the main variable that influences spawning timing and
duration for yellow perch in southern Lake Michigan. Historically, the early 2012 spawn
was not expected based on the calendar, but could be explained by temperatures. Record
spring weather resulted in rapidly rising water temperatures in Lake Michigan in 2012
(NOAA 2013), with water temperatures in my sample area more than 2 ⁰C warmer in the
2012 spring compared to 2009 or 2011 (GLCW 2013). The accelerated warming period
aligns with the earlier spawning period of 2012 and likely demonstrates the effect of the
temperature threshold driving initiation of spawning, which appears to be ~ 11 ⁰C.
Yellow perch spawning is influenced by environmental temperatures throughout their
range (Hokanson 1977; Kayes & Calbert 1979; Brown et al. 2009; Collingsworth &
Marschall 2011), although differences in yellow perch spawning duration appear to
relate, in part, to an upper thermal limit between 13 and 14 ⁰C. In 2009, water
temperatures rose relatively quickly to 14 ⁰C resulting in the shortest yellow perch
10 spawning period; whereas 2011 water temperatures increased more gradually to 13.3 ⁰C
resulting in the longest spawning period. The 2012 water temperatures rose to 13.4 ⁰C
before declining to 12. 3 ⁰C resulting in an intermediate spawning period length. Despite
the 2012 yellow perch spawning period occurring two weeks earlier compared to 2009
and 2011, water temperatures at the start of spawning were within 1 ⁰C for all years,
suggesting that temperature is a major determinate on yellow perch spawning behavior.
My data suggested photoperiod did not influence yellow perch spawning activity
in southern Lake Michigan despite the contrary findings of others (Hokanson 1977;
Kolkovski & Dabrowski 1998; Hinshaw 2006). In hatcheries, reproduction of yellow
perch were found to occur when ~ 12 h of light per day was reached (Kolkovski &
Dabrowski 1998), while Kayes & Calbert (1979) found that wild yellow perch exposed to
rising water temperatures from 2.5 to 9.0 – 13 ⁰C spawned simultaneously regardless of
daylight that ranged between 10.5 and 13.5 h. Southern Lake Michigan has a photoperiod
in May that is approximately 14 h. If a shorter photoperiod was responsible for triggering
spawning, then, I might have expected spawning to occur as early as mid-March, which
was not the case. The absence of spawning in March at the 12 h photoperiod length
suggested that other factors must be met or are controlling spawning initiation.
My findings add another small piece in our understanding of global warming and
its influence on poikilothermic organisms, such as yellow perch. Should the temperature
regime in Lake Michigan shift to favor shorter winters and longer summers, the specific
influences on yellow perch are unknown. However, successful yellow perch reproduction
requires a minimum winter chill period of < 10 ⁰C for 160 days to ensure proper gonadal
11 development and maturation (Hokanson 1977; Dabrowski et al. 1996). After spawning,
yellow perch egg developmental period decreases with an increase in water temperature
(Huff et al. 2004). Therefore, in regions where winter temperatures promote gonad
maturation, warmer spring weather may extend yellow perch young-of-year growth
through earlier spawning, shorter hatching times, and fewer physical abnormalities
allowing for better survival to age 1 (Hokanson 1977; Huff et al. 2004). Conversely,
warmer than normal winter temperatures in the southern ranges of the yellow perch, such
as the Carolinas (Hinshaw 2006), could disrupt gonad development and potentially
extirpate yellow perch from these areas. My results also suggest species such as yellow
perch will have new environmental influences that must be recognized and monitored
with the advent of global climate change (National Research Council 2001). Ultimately,
global climate change may have a dual role in the ultimate fate of this fish,
simultaneously limiting yellow perch populations in some regions and fortifying or
promoting populations in others.
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Treasurer, J. W., and F. G. T. Holliday. 1981. Some aspects of the reproductive biology
of perch, Perca fluviatilis L. A histological description of the reproductive cycle.
Journal of Fish Biology. 18: 359-376.
Tsai, C. and G. R. Gibson. 1971. Fecundity of the yellow perch, Perca flavescens
Mitchill, in the Patuxent River, Maryland. Chesapeake Science. 12(4): 270-274.
United States Geological Survey (USGS). 2008. Commercial fish production – pounds
and value, 2008 Lake Michigan U.S. waters. Available:
http://www.glsc.usgs.gov/_files/cfreports/noaa08.txt. (July 2011).
Wilberg, M. J., J. R. Bence, B. T. Eggold, D. Makauskas, D. F. Clapp. 2005. Yellow
perch dynamics in southwestern Lake Michigan during 1986-2002. North
American Journal of Fisheries Management. 25: 1130-1152.
15 Figures
F
1
3 2 Figure 1. Loccations of yellow perch sampling
s
zonnes near Micchigan City (1), Burns
Harbor
H
(2), an
nd East Chiccago (3) in th
he Indiana w
waters of Lakke Michigann.
16 Females
1.0
Males
1.0
May 20
2009
May 23
0.8
0.8
0.6
0.6
0.4
0.4
0.2
0.2
0.0
0.0
1.0
1.0
June 8
May 26
2011
0.8
0.8
0.6
0.6
0.4
0.4
0.2
0.2
0.0
0.0
1.0
1.0
June 3
May 8
2012
0.8
0.8
0.6
0.6
0.4
0.4
0.2
0.2
0.0
0.0
April 1 April 20 M ay 10 M ay 30 June 19 June 29
July 19 April 1 April 20 M ay 10 M ay 30 June 19 June 29 July 19
Figure 2. Logistic spawning curves for female and male yellow perch collected from the
Indiana waters of Lake Michigan in 2009, 2011, and 2012. The dashed line marks the
point of peak spawn (50% spent). The date indicates the day at which peak spawn
occurred.
17 20
Water Temperature (C)
18
2009
2011
2012
16
14
12
10
8
6
April 1
April 20
May 10
May 30
June 19
July 9
July 29
Figure 3. Lake Michigan mean bottom temperatures by day for 2009, 2011, and 2012.
18 b
16
Water Temperature (C)
14
a
a
2009
2011
12
10
8
6
4
2
0
2012
Year
Figure 4. Lake Michigan mean bottom temperature comparisons for the sampling period
in 2009, 2011, and 2012. The 2009 and 2011 temperatures were statistically similar (a)
while 2012 was higher (b).
19 Tables
Table 1. Abundance of mature yellow perch captured in gill-nets fished in Lake Michigan
during 2009, 2011, and 2012.
Females
Males
Year
Pre-spawn
Post-spawn
Pre-spawn
Post-spawn
2012
12
638
34
24
2011
69
953
126
62
2009
178
785
208
85
Total
259
2292
368
164
Chapter 2: Pre- and post-spawn population demographics for yellow perch in the
Indiana waters of Lake Michigan.
Abstract – Yellow perch were collected from the Indiana waters of Lake Michigan in
2009, 2011, and 2012 to compare pre- and post-spawn fish demographics and determine
whether differences were nonrandom. To identify differences in demographics, I
compared pre- and post-spawn total abundance, CPUE, mean length, and sex ratios.
Yellow perch were collected weekly from May – August at zones near Michigan City,
Burns Harbor, and East Chicago. Fish were processed onshore where they were counted,
measured, sexed, and maturity state was determined. Mean length and CPUE increased
from pre-spawn to post-spawn for female yellow perch in all years while sex ratios
shifted from predominantly male groups pre-spawn to predominantly female groups postspawn in all years. Collectively, these results suggested large female yellow perch were
migrating into the Indiana waters after spawning. The increased number of yellow perch
post-spawn indicates that the warm, relatively shallow Indiana waters may not be
desirable spawning areas, but are attractive for other life history stages, and likely used
for feeding post spawn. My findings suggest that these fish act as a single population not
bound to political jurisdictions and that cooperative inter-state management of yellow
perch in the southern basin of Lake Michigan must occur to best manage the fishery.
Introduction
Recreational and commercial anglers have pursued yellow perch, Perca
flavescens, in Lake Michigan since the late 1800s (Eshenroder et al. 1995). Yellow perch
were caught by recreational anglers at the highest rates in Lake Michigan in the 1980s
and 1990s and continue to attract large numbers of anglers (Bence & Smith 1999). Prior
to 1997, an extensive commercial fishery harvested more than 2 million pounds of yellow
perch in some years from Lake Michigan (Baldwin et al. 2009). Unfortunately,
exploitation likely exceeded sustainable levels in the 1980s and 1990s (Wilberg et al.
2005), and when coupled with recruitment failure (Marsden and Robillard 2004),
declines in the existing population levels were observed.
By 1997, these declines in yellow perch abundance had not reversed which lead to
the closure of the commercial fishery and stricter harvest limits for the recreational
fishery throughout most of Lake Michigan (Marsden & Robillard 2004; Wilberg et al.
2005). Although these management actions taken by the bordering state agencies were
intended to enhance spawning stock biomass, increases failed to materialize and
remained at less than 10% of 1980s levels (Wilberg et al. 2005). These findings
intensified the need to investigate other possible causes of reproductive failure in Lake
Michigan, specifically those factors that are influencing early life history of yellow perch.
Three distinct stocks or populations of yellow perch likely occur in Lake
Michigan and include the southern basin of Lake Michigan, northern basin of Lake
Michigan, and Green Bay (Marsden et al. 1993; Miller 2003; Clapp & Dettmers 2004;
Glover et al. 2008). Yellow perch spawning groups in the southern basin of Lake
22 Michigan are a genetically distinct and homogenous population that is separate from
Green Bay (Miller 2003). This distinction was also shown using tagged yellow perch,
where movement was noted up to 101 km; however, no southern basin tagged fish were
recovered in the northern basin suggesting movement between northern and southern
regions of the lake was minimal (Glover et al. 2008). Finally, Beletsky et al. (2007) used
settlement models driven by currents to document larval fish movement away from natal
spawning areas in the southwestern portion of the lake to the entire southern basin. Adult
yellow perch have spawning site fidelity in southern Lake Michigan, with 35-80% of fish
returning to their location of capture (Glover et al. 2008). Similarly, Marsden et al.
(1993) documented yellow perch returning to tagging sites annually for spawning in
Illinois waters of Lake Michigan, but moving great distances throughout the southern
basin during summer and fall. Thus, yellow perch behavior along with the physical
attributes of the lake show that seasonal differences exist in habitat usage, yet multiple
stocks exist within the lake that are separate and distinct. This segregation of lake-wide
stocks must be acknowledged for successful evaluation and management of the resource
(Clapp & Dettmers 2004).
Spawning habits of yellow perch in Lake Michigan have been poorly studied in
contrast with smaller lakes (Smith 1979; Krieger et al. 1983; Robillard & Marsden 2001;
Mangan 2004; Brown et al. 2009). Typical spawning substrates, such as aquatic
macrophytes and woody debris are not present in Lake Michigan (Robillard & Marsden
2001), and heavy currents during storm events make shallow waters (< 5 m) unsuitable
for spawning activity (Smith 1979; Krieger et al. 1983). Clady and Hutchinson (1975)
23 documented large numbers of yellow perch eggs washed ashore in unprotected shorelines
in Oneida Lake, New York indicting the susceptibility of yellow perch eggs to rough
waters. These habitat limitations demand that successful yellow perch spawning must
occur in deeper waters (> 5 m) over solid substrates such as cobble, rock, and riprap
(Smith 1979; Dorr 1982; Robillard & Marsden 2001). Unfortunately, rocky substrates
are not present in all areas of the lake, and not in the shallow sandy shore of southern
Lake Michigan (Dorr 1982; Robillard & Marsden 2001; Janssen et al. 2005; Walters
2010). Suitable spawning habitat is essential to the recruitment of yellow perch (Mangan
2004), but the extent of this habitat in the Indiana waters is at best, limited (Glover et al.
2008).
My goal for this study was to determine which variables in the Indiana waters of
Lake Michigan influence yellow perch life history and spawning activity. My first
objective was to identify whether pre- and post-spawn yellow perch groups differed in
the Indiana waters of Lake Michigan using population demographics (abundance, sex
ratios, mean length, and catch-per-unit-effort). My second objective was to determine
whether demographic results were consistent among 2009, 2011, and 2012. I
hypothesized that pre- and post-spawn group demography would be non-random and that
differences would be consistent among all sampling years. If pre- and post-spawn
demography differed, then two distinct groups of the same population are present in
Indiana waters before and after spawning. Identifying these distinctions will aid
management efforts and facilitate our understanding of yellow perch in this large and
complex ecosystem.
24 Methods
Study site- Lake Michigan is the 5th largest lake in the world with a volume of 4,918 km3
and a maximum depth of 281 m and is bordered by Indiana, Illinois, Michigan, and
Wisconsin. My sampling occurred in the Indiana waters of southern Lake Michigan,
which are relatively shallow (< 20 m) and have a predominantly sand substrate (Janssen
et al. 2005; Walters 2010). Yellow perch sampling was located in zones within the
Indiana waters near East Chicago, Burns Harbor, and Michigan City (Figure 1).
Field methods- Yellow perch were collected in 2009, 2011, and 2012. In 2009, samples
were collected daily from May 4th to June 4th and then once per week after June 4th until
the yellow perch population reached 100% spent (Walters 2010). In 2011, samples were
collected twice a week from May 10th to June 20th and then once per week from June 20th
to August 1st. In 2012, samples were collected on April 13th and 14th in response to
perceived early season warming of lake waters and anecdotal observations. Twice weekly
sampling resumed from May 1st to June 21st, and then once per week until July 31st.
Each sampling zone was 12 km long and was divided into 12 equal sampling
frames. Daily sampling meant taking three samples, one from each zone. Within each
zone, one sampling frame was selected randomly on any given day using a random
number generator. Sampling was occasionally altered (typically delayed) due to
inclement weather.
Yellow perch were collected using gill nets that were experimental in design
containing nine, 15.2 m panels of repeating 51, 64, and 76 mm stretch-mesh for a total
25 length of 137 m and depth of 1.8 m. These mesh sizes effectively collect mature yellow
perch > 175 mm that were required for my study (Thomas 2007). Gill nets were fished
parallel to shore at a depth contour between 5 and 20 m based on previous catch rates and
water temperatures, set before sunset and soaked overnight for ~ 12 hours. The following
morning, yellow perch were removed from the net, immediately placed on ice, and then
transported to the Indiana DNR field station in Michigan City, IN. Onshore, yellow perch
were counted, measured (total length, mm), weighed (g), and sexed. Maturity state was
determined and classified using a modified gonad stage index that included gravid,
running, and spent categories (Treasurer & Holliday 1981).
Statistical methods- Data were collected from three zones in the Indiana waters of Lake
Michigan and pooled for all subsequent analyses. To determine whether differences
existed among sites before data combination I used a generalized linear model (SPSS
19.0) to create a multiple regression model with total abundance as the response variable
and zone, sex, maturity (pre- or post-spawn) and interactions as predictors. The model
allowed me to identify differences in zones based on total abundances of female and male
yellow perch during pre- or post-spawn seasons for each year.
I was interested in whether female and male CPUE differed between pre-spawn
and post-spawn periods across the three years of sampling. I used multiple regression
through a generalized linear model function with CPUE as the response variable and
maturity (pre- or post-spawn), year, sex, and interactions as predictors (SPSS 19.0). The
model allowed me to identify which predictors were significantly correlated with CPUE
and significance was determined using a Likelihood ratio test statistic. Because pre- and
26 post-spawn fish were collected simultaneously in gill nets, yellow perch CPUE had to be
calculated separately for these two classifications. This was accomplished by separating
pre-spawn and post-spawn fish into two groups and then separating females and males. I
was then left with CPUE values that represented the number of pre- or post-spawn female
and male fish collected in each net.
Mean differences in total length of pre- and post-spawn yellow perch were
compared to determine whether differences existed within and among sampling years. I
used a generalized linear model function (SPSS 19.0) to create a multiple regression
model using total length as the response variable and sex, maturity (pre- or post-spawn),
year, and the interactions as predictors. To account for yellow perch growth over the
sampling period, 7 mm was subtracted from the mean total length of post-spawn females
and 5 mm was subtracted from the mean total length of post-spawn males following
Walters (2010). This adjustment in total length was conservative (likely overestimating
growth) which attempted to statistically eliminate or minimize the confounding effect of
growth that occurred during the ~ 8 week period when most fish were collected. The
choice of length subtracted by sex was based on historic growth rate data for yellow
perch in Lake Michigan (see Headley and Lauer 2008). A post hoc Fisher’s least
significant difference pairwise comparison determined differences among factors. A
change in mean total length for pre- and post-spawn groups would indicate that different
groups of yellow perch were present before and after the spawn.
Lastly, I was interested in whether differences in sex ratios were present in the
pre- and post-spawn groups of yellow perch and whether differences were similar among
27 2009, 2011, and 2012. I used the generalized linear model function (SPSS 19.0) to create
a logistic regression model with a binomial response variable (sex) and maturity (pre- or
post-spawn), year, and maturity* year interactions as predictors (SPSS 19.0). Effects of
model predictors were evaluated using a Wald chi-square test statistic. Changes in sex
ratios would suggest that immigration, emigration, or differential mortality was occurring
between pre- and post-spawn periods each year. Alpha was set at 0.05 for all tests of
significance.
Results
A total of 3083 yellow perch was collected from the Indiana waters of Lake
Michigan during 2009, 2011, and 2012 (Table 1). Female fish (N = 2551) comprised 83%
of the total and ranged in total length (by year) from 166 to 355 mm (2009), 132 to 369
mm (2011), and 146 to 360 mm (2012). Male yellow perch (N = 532) ranged in total
length (by year) from 160 to 325 mm (2009), 127 to 315 mm (2011), and 96 to 301 mm
(2012). Ten percent of mature females and 70% of mature males were not spent and were
considered pre-spawn (Table 1).
Initial total abundance comparisons among zones using multiple regression found
no differences between the Michigan City zone and the Burns Harbor zone (P = 0.718);
however, the East Chicago zone had lower abundance and was ~ 13% of the total
abundance of Michigan City and Burns Harbor (P < 0.001). Despite the lower total
abundance of yellow perch at East Chicago, post-spawn increases in female abundance
and decreases in male abundance continued to occur, mimicking the trends in the other
28 two zones. These results and the goal of my study, to evaluate the Indiana waters as a
whole, indicated data from all zones could be pooled for analysis.
Yellow perch pre-spawn CPUE was about one third of post-spawn CPUE for all
years combined (Table 2) but total CPUE did not vary among years (Table 2). More
female yellow perch were collected than male yellow perch but the proportion of female
and male fish remained constant among all years and between pre- and post-spawn
seasons (Table 2). Female yellow perch CPUE was significantly lower pre-spawn
(reference) compared to post-spawn and post hoc comparisons revealed that CPUE (mean
± SE) for pre-spawn female yellow perch was an order of magnitude lower (3.0 ± 4.2) for
107 net sets compared to post-spawn female CPUE (29.0 ± 3.2) for 123 net sets. While
male CPUE appeared to be higher pre-spawn (6.7 ± 4.0) than post-spawn (4.1 ± 4.6), post
hoc comparisons revealed these differences were not significant (P = 0.662). Pre- and
post-spawn yellow perch were not present in all net sets, resulting in unequal numbers of
samples.
Yellow perch female lengths were greater than male lengths in all years (Table 3).
Female mean length for all years combined was 242.2 ± 1.7 mm; however, female
lengths in 2009 were ~ 10 mm smaller than female lengths in 2011 and 2012. Despite
differences in 2009 female lengths, an increase in length was seen for female yellow
perch from pre-spawn to post-spawn seasons in all years (Table 3). Post hoc comparisons
revealed length (mean ± SE) for female yellow perch increased from 231.5 ± 3.2 mm prespawn to 252.8 ± 0.8 mm post-spawn Table 3). Male yellow perch lengths were similar
29 among years and post hoc comparisons determined there were not significant changes
from 217.2 ± 2.4 mm pre-spawn to 221.1 ± 3.1 mm post spawn (Table 3).
Female yellow perch comprised 35% of the total pre-spawn abundance and 94%
of the total post-spawn abundance for all years combined. However, the total proportion
of female yellow perch collected in each year (~ 75%) did not vary among years (Table
5). Yellow perch sex ratios were skewed towards fewer females in the pre-spawn period
for all years (2009, 29%; 2011, 35%; 2012, 26%) when compared to the post-spawn
period (2009, 82%; 2011, 94%; 2012, 96%) and these differences were significant in all
years (Table 5).
Discussion
As hypothesized, yellow perch group demographics changed between pre- and
post-spawn periods in the Indiana waters of Lake Michigan all years sampled. The total
and relative (CPUE) number of female yellow perch and mean length increased from preto post-spawn in all years. Male abundance decreased from pre- to post-spawn in all
years, while no differences in total lengths were observed (Table 1). The change in
demographics suggests post-spawning immigration of female yellow perch into the
Indiana waters is occurring. Movement of yellow perch into the Indiana waters is not
unusual as indicated by tagging and genetic studies (Marsden et al. 1993; Miller 2003;
Glover et al. 2008). This movement throughout the southern basin allows for seasonal
usage of different regions (habitats) within the basin. Adult yellow perch will move
distances over 100 km and larval yellow perch have the potential to recruit anywhere in
the southern Lake Michigan basin from spawning grounds in southwestern Lake
30 Michigan (Marsden et al. 1993; Beletsky et al. 2007; Glover et al. 2008). Thus, suitable
spawning areas could be distant from locations where seasonal foraging occurs. My
results indicate distinctly different groups of the same population of yellow perch are
residing in the Indiana waters of Lake Michigan before and after the spawn.
A failure to identify few pre-spawn females in the Indiana portion of Lake
Michigan implies that suitable spawning habitat in these waters is limited at best. The
substrates in the Indiana waters of Lake Michigan are primarily sand and offer little
protection from strong currents and wave activity which can destroy eggs on unprotected
shorelines (Clady & Hutchinson 1975; Janssen et al. 2005; Walters 2010). In 2010, few
larval yellow perch were collected throughout the Indiana waters (Rounds 2011), while
simultaneous scuba and video observations failed to locate yellow perch egg skeins at
these same locations (Walters 2010). However, in nearby Illinois waters of Lake
Michigan where typical substrates include rock, cobble, and glacial boulders, yellow
perch spawning has been documented (Marsden et al. 1993; Janssen et al. 2005; Glover
et al. 2008). Further, Robillard and Marsden (2001) showed higher densities of yellow
perch prepared to or engaged in spawning in areas dominated by cobble substrates
compared to sand substrates in Lake Michigan near Lake Bluff, IL. Thus, limited
spawning habitat, few spawning adult yellow perch, absence of egg skeins, and few larval
yellow perch all support the same conclusion: spawning of yellow perch in the Indiana
portion of Lake Michigan is minimal or non-existent.
The Indiana waters of Lake Michigan appear to be well suited foraging grounds
for yellow perch because of the relatively warm, shallow waters preferred by yellow
31 perch in this area (Rydell et al. 2010). Growth for yellow perch begins after spawning
and requires high energy intake (Henderson et al. 2000; Collingsworth & Marschall
2011). Large amounts of forage fish are available in the Indiana waters and are
commonly eaten by adult yellow perch immediately following spawning (Truemper &
Lauer 2005; Forsythe et al. 2012). Additionally, Indiana commercial fishermen have
historically harvested a disproportionally high number of yellow perch based on area of
lake further supporting high use of Indiana waters by yellow perch (Baldwin et al. 2009).
Yellow perch grew larger in their first year of life in the Indiana waters of Lake Michigan
compared to other areas of the lake (Horns 2001), further suggesting that Indiana waters
are more productive feeding grounds during a time of high energy need.
Habitat use by pre-spawn male yellow perch does not appear to follow patterns
exhibited by female yellow perch. Although male pre- and post-spawn CPUE were
statistically similar, total abundance was greater pre-spawn compared to post-spawn
during each sampling year (Table 1) and corresponds with Dorr (1982) who showed that
male yellow perch congregate in shallow waters (6-12 m) before spawning and disperse
afterwards. While little spawning appears to be occurring in the Indiana waters, local
male yellow perch are likely grouping and moving into my sampling areas with the
perceived spawning season. Low male seasonal abundance when compared to females
indicates that proportionally few male yellow perch reside in Indiana waters and is
consistent with historic annual trends (Lauer & Doll 2012). The absence of large numbers
of male yellow perch further indicates spawning is limited or non-existent in the Indiana
waters.
32 Management implications - Yellow perch spawning behavior and movement in the
Indiana waters of Lake Michigan provides further evidence that the southern basin of
Lake Michigan must be managed cooperatively among bordering state agencies (Clapp &
Dettmers 2004; Glover et al. 2008). My results indicate females and males are behaving
separately in the spring and early summer, likely associated with spawning activity that
extends into the neighboring political jurisdictions. These yellow perch have varying
angler pressures among bordering states and combined fishing mortality for the yellow
perch population could have a greater effect than previously determined (Wilberg et al.
2005; Makauskas & Clapp 2009). Further, habitat limitations in southern Lake Michigan
should encourage protection of known spawning sites in southwestern Lake Michigan to
increase yellow perch spawning stock biomass (Marsden et al. 1993; Wilberg et al. 2005;
Glover et al. 2008). It appears that the Indiana waters of Lake Michigan serve as feeding
grounds after spawning for yellow perch; however, little reproduction occurs in Indiana
waters. Identification of other habitat limitations for yellow perch in the southern basin of
Lake Michigan is essential for improving management and recovery of this fish
population.
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Wilberg, M. J., J. R. Bence, B. T. Eggold, D. Makauskas, D. F. Clapp. 2005. Yellow
perch Dynamics in southwestern Lake Michigan during 1986-2002. North
American Journal of Fisheries Management. 25: 1130-1152.
36 Figures
F
1 3 2
s
zonnes Michigaan City (1), B
Burns Harboor (2),
Figure 1. Loccations of yellow perch sampling
nd East Chiccago (3) in th
he Indiana waters
w
of Lakke Michigann.
an
37 Tables
Table 1. Yellow perch gill net collection abundance from the Indiana waters of Lake
Michigan for 2009, 2011, and 2012.
Females
Males
Year
Pre-spawn
Post-spawn
Pre-spawn
Post-spawn
2009
178
785
208
85
2011
69
953
126
62
2012
12
638
34
24
Total
259
2292
368
164
38 Table 2. Results of the multiple regression model for yellow perch collected within the
Indiana waters of Lake Michigan in 2009, 2011, and 2012 with CPUE as the response
variable.
Source
df
Likelihood ratio
P - value
Intercept
1
26.312
< 0.001
Maturity
1
7.706
0.006
Sex
1
7.109
0.008
Year
2
1.96
0.375
Sex *Year
2
0.473
0.789
Maturity *Sex
1
11.32
0.001
Maturity * Year
2
0.759
0.684
Maturity *Year
*Sex
2
0.758
0.684
39 Table 3. Mean length ± SE of yellow perch collected in the Indiana waters of Lake
Michigan in 2009, 2011, and 2012.
Females
Males
Year
Pre-spawn
Post-spawn
Pre-spawn
Post-spawn
2009
225 ± 2.7
246 ± 1.3
220 ± 2.5
220 ± 4.0
2011
238 ± 4.4
254 ± 1.2
219 ± 3.3
220 ± 4.5
2012
239 ± 10.6
259 ± 1.4
210 ± 6.3
227 ± 7.5
40 Table 4. Results of the multiple regression model for yellow perch collected in the
Indiana waters of Lake Michigan in 2009, 2011, and 2012 with the response variable
(length) and corresponding P-values for each predictor.
Source
df
Likelihood ratio
P - value
Intercept
1
7067.129
< 0.001
Maturity
1
18.337
< 0.001
Sex
1
70.625
< 0.001
Year
2
6.051
0.049
Sex* Year
2
7.444
0.024
Maturity* Sex
1
5.448
0.020
Maturity* Year
2
1.457
0.478
41 Table 5. Results of the logistic regression for yellow perch collected in the Indiana waters
of Lake Michigan in 2009, 2011, and 2012.
Source
df
Wald chi-square
P - value
Intercept
1
189.122
< 0.001
Maturity
1
459.732
< 0.001
Year
2
0.191
0.909
Maturity* Year
2
29.246
< 0.001