United States
Environmental Protection
Agency
Environmental Research
Laboratory
Duluth MN 55804
Research and Development
EPA/600/S3-85/049 Aug. 1985
f/EPA Project Summary
Reproduction and Distribution of
Fishes in a Cooling Lake:
Wisconsin Power Plant Impact
Study
Dennis W. Rondorf and James F. Kitchell
Spatial and temporal patterns during
reproduction and early life history of
fishes were studied in a manmade cool-
ing lake. Lake Columbia, impounded in
1974, near Portage, Wisconsin, has an
area of 190 ha, a mean depth of 2.1 m,
and a 15°C temperature gradient de-
rived from the thermal effluent of a
527-MW fossil-fueled generating sta-
tion which began operation in 1975.
The lake was initially colonized by
fishes when filled with Wisconsin River
water. Observations suggested a de-
cline of species diversity of the fish
community due to: (1) direct action of
upper lethal temperatures, (2) absence
of colonization by warm-water, lake-
dwelling species, and (3) lack of recruit-
ment for certain species.
Spatial and temporal patterns of
spawning of black crappie (Pomoxis
nigromaculatus) were altered by a rapid
rise in water temperatures following
plant startup after a 3-week shutdown.
Water temperatures above expected
spawning temperatures reduced avail-
able spawning area and induced aggre-
gation of sexually mature black crappie
at coolest available temperatures. Ele-
vated temperatures subsequently short-
ened the spawning season, induced
resorption of ova, and caused loss of
secondary sexual characteristics.
A second generating unit began opera-
tion in February 1978. Spawning of
black crappie and white bass (Morone
chrysops) occurred 1 month earlier dur-
ing the spring of 1978 than in 1977.
Species abundance of larval fish
catches was greater in 1978 when the
spawning season was not unusually
abbreviated, as in 1977. After initially
drifting with water current, juvenile
stages of sunfish (Lepomis sp.) and giz-
zard shad (Dorosoma cepedianum) re-
sponded to changes in the thermal gra-
dient by horizontal and vertical shifts in
abundance.
This Project Summary was developed
by EPA's Environmental Research Lab-
oratory, Duluth, MN, to announce key
findings of the research project that is
fully documented in a separate report of
the same title (see Project Report order-
ing information at back).
Introduction
Impoundment of a man-made cooling
lake provides an opportunity for develop-
ment of a recreational fishery. Thermally
elevated areas of a cooling lake can
increase angler use in temperate climates
(McNurney era/. 1977). Because the cool-
ing lake is a relatively small man-made
system, it has greater potential and flexi-
bility for recreational fishery management
than many systems.
Fish management strategies of stock-
ing or fish harvest for temperate lakes
based on surface area of morphoedaphic
characteristics are not valid for lakes sub-
stantially altered by thermal input. The
elevated thermal conditions of cooling
lakes create temporal and spatial limita-
tions of organisms native to nearby lakes.
Growth (Bennett and Gibbons 1974), dis-
tribution (Merriman and Thorpe 1976),
and reproduction (Bennett and Gibbons
1975, Kaya 1977) are modified in ther-
-------
4
mally altered areas. Lake Columbia, lo-
cated at the Columbia Generating Station
near Portage, Wisconsin, provided an
opportunity for research and development
of management strategies specifically for
cooling lakes in the Great Lakes region.
As a subproject of an assessment of a
developing cooling lake ecosystem (Lo-
zano et al. 1978), this study of fishes con-
cerned the reproductive responses of
adults and distributional patterns of adult,
larval, and early juvenile forms in Lake
Columbia. The objectives were (1) deter-
mine changes in species composition of
the fish population; (2) delineate temporal
and spatial limits of fish reproduction;
and (3) determine distributional patterns
of larval and early juvenile fishes.
Relative Abundance of Fishes
New impoundments are known for the
dramatic changes they cause in fish
communities after filling. Studies sug-
gest these changes can be attributed to
changes of biotic and abiotic characteris-
tics of the impoundment (Jenkins and
Morals 1971). In the cooling lake envir-
onment, the effects of the unique thermal
regime are imposed on these expected
changes in the fish community. There-
fore, the first objective was to determine
if changes in species composition of the
fish population occurred with time.
Methods
Composition of the fish community and
the relative abundance of adults at loca-
tions in Lake Columbia were assessed
using fyke nets. Netted fish were identi-
fied, measured for total length, and
weighed. Water temperatures were mea-
sured at 0.5, 1.0, 1.5, and 2.0-m depths
when nets were set and raised. Die!
temperature changes were continually
monitored with recording thermographs.
The number of fish per fyke net set—
catch per effort (CPE)—was used as a
measu re of the relative abundance of fish
species over time. Species diversity of the
fish population in quarterly fyke net
catches was calculated using the Shan-
non and Weaver (1963) general index of
diversity (H).
Results
The Shannon-Weaver diversity index
suggested a decline in diversity of quar-
terly catches that began in the winter of
1975-76 and continued through the fall
of 1976 (Figure 1). I n fall 1976 only 34%
of the catch was similar to that of the
summer quarter 1975. Species diversity
did not return to 1975 levels (Figure 2).
2.0-1
I
I
<3
1.0-
Percent similar to first sample
J
S F
1975
V
W
S
1976
l
W
rlOO
-50
to
I
Q.
S
1977
Figure 1. Shannon-Weaver diversity index (H) and percent similarity to first sample of
quarterly fyke net catches in Lake Columbia.
Changes in the diversity of fyke net
catches were attributed to a decline of
CPE for some species and numerical dom-
inance by others. The mean quarterly CPE
of centrarchids in Lake Columbia (Figure
2A) indicated that the abundance of
purnpkinseedsunfish(/.epo/r7/s<7/6A0si/s)
declined somewhat between July 1975
and August 1977. The mean CPE for
white bass(Morone chrysops)(Figure 2B)
was high in the winter quarter samples.
Bluegill (Lepomis macrochirus) catches
increased dramatically, while gizzard
shad (Dorosoma cepedianum) CPE in-
creased from none caught during the first
quarter to a CEP >5 in 1977 (Figure 2B).
Bluegill and gizzard shad length frequen-
cies indicated successful growth and
reproduciton in the new lake environ-
ment, as evidenced by increased CPE.
Other fish species common to the Wis-
consin River were collected in Lake
Columbia. The relationship between
changes in relative abundance, final
temperature preferenda, and upper incip-
ient lethal temperatures of the most
abundant species in fyke net catches is
shown in Figure 3. Yellow perch (Perca
flavescens) and walleye (Stizostedion
vitreum) made up <0.5% of all catches.
Species at low abundance that decreased
in CPE included quillback (Carpoides
cyprinus), smallmouth buffalo (Ictiobus
bubalus). and carp (Cyprinus carpio) (Fig-
ures 2C, 3). Minnow trap catches indi-
cated fathead minnows (Pimephales
promelas] and mud minnows (Umbra limi)
were abundant when the plant began
60-i
40-
20-
o
15-
1 /0"
<<
c
***
o
7.5-
1-
0.5-
, Bluegill
'--_/ /\' f-B. crappie
-Pumpkinseed
®
V-.,,---.. /._/V. pfte
© '"' Buffalo sp.
I '
"\-Carp
.Quillback
SFWSSFWSS
1975 1976 1977
Figure 2. Mean catch per effort in fyke
nets during quarterly sampling
periods in Lake Columbia.
-------
operation in March 1975, but abundance
declined by July. The mean quarterly CPE
of northern pike declined 99% between
the first and third year of the study and
disappeared from catches during the
summer quarter of 1976 (Figure 2B).
Bluegill abundance increased more than
eight-fold between the first and third
years (Figure 3). This species was well
adapted for the cooling lake environment
because of its relatively high thermal
preference, abilityto forage in open water,
and the presence of suitable spawning
habitat.
Reproduction of Fishes
Thermal requirements for reproduction
of a species are one of the primary limits
to distribution and abundance (Alderdice
and Forrester 1968, Hokanson 1977).
Temperature and photoperiod are envi-
ronmental factors important in the recru-
desence of gametogensis of many t°ie-
osts (De Vlaming 1972, 1974, Schreck
1974, Hokanson 1977). The unique ther-
mal regimes in the cooling lakes create
temperatures during winter and early
spring at or above those associated with
the initiation of spring spawning. The
thermal gradient was expected to deter-
mine the spatial and temporal location of
spawning within limits of photoperiod
control. Thus, the second objective was to
study the temporal and spatial limits of
reproduction within Lake Columbia.
Methods
White bass and black crappie were
selected for study because adults were
growing well but did not appear to be pro-
ducing young.'
Temporal patterns of reproduction were
determined by monitoring the maturity of
white bass and black crappie from the
initiation to the completion of spawning
during the springs of 1977 and 1978. The
percent gonad weight of total body weight
or gonadosomatic index was calculated
as a measure of maturity at 2-week inter-
vals starting in February 1977.
Spatial limitations of spawning relative
to the thermal gradient were examined
during the springs of 1977 and 1978.
Relative abundance of sexually mature
fish at varous locations in Lake Columbia
was determined by fyke net sampling and
regression analysis methods(Rondorf and
Kitchell, 1983). Thermal exposure of adult
black crappie was estimated from equa-
tions describing distribution of adults and
the thermal gradient.
Preferred Temperature ± S.D.
Incipient Lethal Temperature
Percent
Decrease/Increase
Gizzard shad
Bluegill
Black Bullhead
Black crappie
Yellow perch
White bass
Pumpkinseed
Carp
Carpsucker sp.
Buffalo sp.
Northern pike
20 30 40
Temperature (CJ
100
800
Results
A rapid rise in water temperatures fol-
lowing a 3-week power plant shutdown
during 1977 stimulated spawning, re-
duced available spawning area, and in-
duced aggregation of black crappie at
coolest available water temperatures.
Elevated water temperatures subsequent-
ly induced resorption of black crappie ova,
loss of secondary sexual characteristics,
and were probably near upper lethal
temperatures of embryo and larval stages.
A temporally shortened spawning season
was associated with a rapid rise in water
temperatures, while additional thermal
input by the Columbia II generating unit
caused spawning to occur 30 days earlier
in the spring.
Distribution of Larval Fishes
Larvae of many spring spawning fishes
initially disperse into limnetic waters
(Faber 1967, Werner 1969, Netsch et at.
1971, Amundrud et al. 1974, Kelso and
Ward 1977) and later aggregate as juve-
niles in littoral areas that have higher
water temperatures. Juvenile fishes ex-
hibit thermoregulatory behavior in the
laboratory (Cherry et al. 1977) and after
reviewing temperature preference data
Reproductive
Guild
N. Litho-pelagophil
G. Lithophil
G. Lithophil
G. Phytophil
N. Phyto-Lithophil
N. Phyto-Lithophil
G. Polyphil
N. Phytophil
N. Psammophil
N. Phytophil
N. Phytophil
Reference
Preferred. Lethal
a.b
a,c
a.d.e
a,b
a,c
a.e
a.b
a.b
a.e
f,e
N = nonguarding spawners
G = guarding spawners
Figure 3. Percent increase or decrease in abundance of selected fishes in Lake Columbia. Preferred and upper incipient lethal temperatures of fish
are from other sources fReutter and Herdendorf 1974.bBrungs and Jones 1977;c Cherry eta\. 1977;" Hart J 952;'Cvancaraeta\. 1977;
'Coutant 1977).
-------
of fish, Coutant (1977) concluded that
laboratory and field results were reason-
ably consistent. During the development
of thermoregulatory behavior, increasing
mobility probably facilitates the ability of
individual juveniles to thermoregulate.
However, little is known about the thermal
responsiveness of pelagic larvae and early
juveniles of fish when distributed in the
limnetic zone. Therefore, an objective of
this study was to describe distributional
responses of pelagic larvae and juvenile
fish to changes in the thermal gradient.
Certain abiotic factors—i.e., temperature—
are correlated with year-class strength of
some fish populations (Kramer and Smith
1962, Koonce et al. 1977).
Methods
Gizzard shad and bluegill, two species
whose populations increased in the cool-
ing lake, were studied to determine how
the larval forms responded to the dynamic
heterothermal environment of the cool-
ing lake. Additional observations were
designed to investigate factors affecting
the relative abundance of larval fish. Time
of capture, depth of capture, station loca-
tion, and die! movement patterns were
examined.
Tows to assess horizontal distribution
patterns were made at a depth of 0.5 m
between 1900 and 2400 h central stand-
ard time (CST). Clear plexiglass traps
equipped with lights were used to collect
larval fish. Larval and juvenile fish were
preserved in the field then counted and
identified in the laboratory.
Data were analyzed using mathemati-
cal methods to describe horizontal distri-
bution patterns.
Results
Species diversity of larval fish catches
was low in 1977 when water tempera-
tures increased rapidly. The median tem-
perature of capture of larval Lepomis sp.
and gizzard shad was near 30°C. Tem-
peratures > 31 °C during thermal stratifi-
cation reduced the abundance of gizzard
shad completing die! vertical movements.
After initially drifing with the current,
Lepomis sp. and gizzard shad responded
to water temperature changes by hori-
zontal shifts in abundance with a mode at
28° to 31 °C.
Conclusions
Observations suggest that the species
diversity of fish in Lake Columbia, Wis-
consin, declined during the first year of
thermal input by the plant. Habitat modi-
fication, such as reduced vegetation as a
direct result of thermal input, may be the
reason for the decrease in abundance of
some species. The decline in species
diversity was accentuated by fish mortal-
ity from temperatures exceeding upper
lethal limits, an absence of colonization of
warm-water lake-dwelling species, and
limited reproductive success.
Thermal inputs by the power plant mod-
ified temporal and spatial characteristics
of spawning white bass and black crap-
pie. Resumption of plant operation follow-
ing a 3-week shutdown resulted in a rapid
increase in water temperatures that
stimulated spawning, reduced available
spawning area, and induced aggregation
of sexually mature black crappie at cool-
est available water temperatures. Water
temperatures above expected spawning
temperatures induced partial resorption
of ovaries, loss of secondary sexual char-
acteristics, and abbreviation of spawning
duration. The combined operation of the
Columbia I and II generating units in-
duced spawning about 1 month earlier
than when only Columbia I was operating.
The rapid increase in water tempera-
tures that induced spawning and subse-
quent gonadal resorption in 1977 was
associated with a lower number of spe-
cies of larval fishes. If a number of species
were stimulated to spawn, then a greater
number of ichthyoplankton species would
be expected at that time, but this was not
observed. Therefore, factors associated
with the increase in temperatures must
have been responsible for reducing ich-
thyoplankton species abundance. The
number of species of larval fishes was
lower during 1977 when water tempera-
tures increased rapidly and reproductive
responses were aberrant. This may ex-
plain the limited reproductive success
observed for some species successful as
adults in the cooling lake environment.
Larval and early juvenile stages of blue-
gill and gizzard shad were responsive to
temperature changes within the thermal
gradient. Larval forms initially drifted
downstream. Pelagic larval and juvenile
stages of bluegill and gizzard shad were
most abundant at 28° to 31 °C and were
capable of responding to changes in the
thermal gradient induced by changing air
temperatures. Larval gizzard shad exhib-
ited reduced diel vertical movements
when temperatures were > 31 °C. Pelagic
larval and early juvenile forms responded
to thermal dynamics of the cooling lake by
modifying horizontal and vertical distribu-
tion patterns.
Recommendations
1. The disappearance of aquatic vege-
tation should be expected in cooling
lakes with heavy thermal loading.
Management to enhance fish popu-
lations of species that require vege-
tation to spawn should not be
attempted.
2. Initiation of spawning was approx-
imately 1 month earlier with the
thermal input of two 527-MW gen-
erating units. If fishery manage-
ment agencies protect spawning
adult fish during spring by closed
season, the regulations may not be
applicable to cooling lake fisheries.
The accelerating of spawning dur-
ing spring can provide the oppor-
tunity to open a fishery while other
temperate lake fisheries are closed
during normal reproductive season.
3. Power plant shutdowns are often
scheduled during spring but rapid
temperature increases when plants
resume operation can cause aber-
rant reproductive responses in
spring-spawning fish. The opera-
tion and more than one generating
unit may be beneficial because
additional units buffer the effects of
rapid changes in temperature of a
single unit.
4. Successful reproduction of fish in a
cooling lake is spatially limited to
water temperatures within the ther-
mal tolerance limits of reproduc-
tion. Cooling lake water tempera-
tures and corresponding areas can
be estimated during planning. De-
sign should provide adequate area
with water temperatures within the
thermal tolerance limits of oogene-
sis, spawning, and incubation dur-
ing spring.
5. Pelagic larval stages drift down-
stream in a recirculating cooling
lake and will subsequently be en-
trained by the power plant if cooling
lake turnover time is not adequate
to permit metamorphosis into the
more mobile early juvenile stages.
Entrainment should be accepted as
a part of the cooling lake environ-
ment, however, careful design can
minimize entrainment of larval and
early juvenile stages. Additional
generating units should be aug-
mented by increasing cooling lake
area and volume or by providing
cooling towers to increase lake turn-
over time.
-------
6. Early juvenile fish in pelagic stages
are mobile and respond to thermal
gradients and water currents by
congregating in preferred habitats.
Cooling lake design can increase
juvenile nursery areas and species
diversity by providing heterogene-
ous habitats with a diversity of
water depths, substrates, and shore-
line configurations. These design
modifications would not inhibit the
cooling capacity of the lake because
cooling is most dependent on sur-
face area.
7. After power plant operation began,
species diversity declined as a few
eurythermal species incresed in
abundance and other species de-
clined. Sport fish should be stocked
early in the operation of the lake so
that juveniles can utilize forage
species likely to become abundant
after the power plant begins opera-
tion. Thermally tolerant nest-guard-
ing Centrarchids and Ictalurids,
such as largemouth bass and chan-
nel catfish, are likely to be the most
successful native species in cooling
lakes and should be preferred for
initial stocking.
8. After the power plant begins opera-
tion, water temperatures of the
cooling lake may exceed the upper
lethal limits of popular cool water
sport fish resulting in fish kills.
Design can create thermal refugia
using water depth and circulation
patterns. Fishery managers andthe
public should anticipate fish kills as
a part of starting the long-term
management of a cooling lake, but
recurrent fish kills can be avoided
through careful design.
9. Largemouth bass, a species sus-
ceptible to over-harvest, was caught
at the highest rate by angling near
the outfall in winter (0.28 bass/
cast) and in cool water near the
intake in late spring (0.27 bass/
cast: Lozano et at. 1978). Thus, the
area near the intake nad outfall
should be permanently closed to
fishing to protect summer and win-
ter aggregations of fish under sea-
sonally extreme thermal conditions.
10. Small reservoirs are often subject
to over-exploitation of sport fish
populations, particularly when fish
are spatially limited by temperature
or lake morphometry. Thus, when
power plant security is designed,
consideration should be given to
minimize over-harvest of fish in
outfall and intake areas and reduce
potential conflicts between power
plant security and fisherman.
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Dennis W. Rondorf and James F. Kitchell are with Water Resources Center,
University of Wisconsin-Madison, Madison, Wl 53706.
Gary E. Glass is the EPA Project Officer (see below).
The complete report, entitled "Reproduction and Distribution of Fishes in a
Cooling Lake: Wisconsin Power Plant Impact Study," (Order No. PB 85-217
669/AS; Cost: $10.00, subject to change) will be available only from:
National Technical Information Service
5285 Port Royal Road
Springfield, VA 22161
Telephone: 703-487-4650
The EPA Project Officer can be contacted at:
Environmental Research Laboratory
U.S. Environmental Protection Agency
Duluth, MN 55804
•fr U. S. GOVERNMENT PRINTING OFFICE: 1985/559-111/20636
-------
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Environmental Protection
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Center for Environmental Research
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