Content uploaded by Heather A. Stewart
Author content
All content in this area was uploaded by Heather A. Stewart on Aug 18, 2014
Content may be subject to copyright.
This article was downloaded by: [Heather Stewart]
On: 04 June 2014, At: 20:28
Publisher: Taylor & Francis
Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House,
37-41 Mortimer Street, London W1T 3JH, UK
North American Journal of Fisheries Management
Publication details, including instructions for authors and subscription information:
http://www.tandfonline.com/loi/ujfm20
Laboratory Investigations on the Use of Strobe Lights
and Bubble Curtains to Deter Dam Escapes of Age-0
Muskellunge
Heather A. Stewart
a
, Max H. Wolter
b
& David H. Wahl
b
a
Department of Natural Resources and Environmental Sciences, University of Illinois at
Urbana–Champaign, 1102 South Goodwin Avenue, Mail Code 047, Urbana, Illinois 61801, USA
b
Illinois Natural History Survey, Kaskaskia Biological Station, 1235 County Road 1000N,
Sullivan, Illinois 61951, USA
Published online: 22 May 2014.
To cite this article: Heather A. Stewart, Max H. Wolter & David H. Wahl (2014) Laboratory Investigations on the Use of Strobe
Lights and Bubble Curtains to Deter Dam Escapes of Age-0 Muskellunge, North American Journal of Fisheries Management,
34:3, 571-575
To link to this article: http://dx.doi.org/10.1080/02755947.2014.892549
PLEASE SCROLL DOWN FOR ARTICLE
Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained
in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no
representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the
Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and
are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and
should be independently verified with primary sources of information. Taylor and Francis shall not be liable for
any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever
or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of
the Content.
This article may be used for research, teaching, and private study purposes. Any substantial or systematic
reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any
form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://
www.tandfonline.com/page/terms-and-conditions
North American Journal of Fisheries Management 34:571–575, 2014
C
American Fisheries Society 2014
ISSN: 0275-5947 print / 1548-8675 online
DOI: 10.1080/02755947.2014.892549
ARTICLE
Laboratory Investigations on the Use of Strobe
Lights and Bubble Curtains to Deter Dam Escapes
of Age-0 Muskellunge
Heather A. Stewart*
Department of Natural Resources and Environmental Sciences,
University of Illinois at Urbana–Champaign, 1102 South Goodwin Avenue, Mail Code 047,
Urbana, Illinois 61801, USA
Max H. Wolter and David H. Wahl
Illinois Natural History Survey, Kaskaskia Biological Station, 1235 County Road 1000N,
Sullivan, Illinois 61951, USA
Abstract
The movement of Muskellunge Esox masquinongy over a dam to leave a reservoir is known as dam escape. It is
common in Midwestern U.S. reservoirs and has been as high as 25% of the adult population. A variety of barrier
and guidance systems have been used to control fish movement, but the success of such barriers has been mixed and
appears to be very species dependent. We examined the effectiveness of a simple, relatively low-power and low-cost
bubble curtain, strobe light, and bubble curtain with strobe light barriers to deter Muskellunge from escaping over
spillways. In 15 replicate trials of each treatment type conducted in a simulated spillway, age-0 Muskellunge were
more likely to escape during daytime trials (P < 0.01), but the three barrier combinations did not reduce rates of
escape. Light and bubble curtain barriers will likely not be effective in reducing spillway escapes by Muskellunge.
Muskellunge Esox masquinongy are important sport fish in
the United States (Simonson and Hewett 1999) and stocking
is an important tool for enhancing populations in reservoirs
(Margenau 1992; Wahl 1999). Following introduction, Muskel-
lunge are frequently observed escaping over spillways after
high-water events. About 25% of a Muskellunge population in
an Illinois reservoir escaped in 1 year (Wolter et al. 2013). Es-
cape of Muskellunge appeared to be an active response as most
happened during peak activity times and in proximity to spawn-
ing. Escaped Muskellunge are s ubject to an uncertain fate and,
unless rescued, are lost to the reservoir fishery. Additionally, the
influx of these large piscivores into downstream systems could
contribute to a decline of native fishes that use the streams for
nurseries (Stanford et al. 1986). Reports of dam escapes have
generated interest in installing barrier nets to prevent fish es-
*Corresponding author: hstewart@cfr.msstate.edu
Received September 18, 2013; accepted January 22, 2014
capes, but they are not always possible or effective due to debris
accumulation (Plosila and White 1970).
Electrical barriers, illumination, air bubble curtains, and
sonic devices are alternatives to barrier nets that deserve consid-
eration (McIninch and Hocutt 1987). Diverting fish with the use
of air bubbles has been suggested for decades (Von Brandt 1967;
Stewart 1982) and has the advantage of technical simplicity and
lower power consumption per unit area than electric barriers
(Stewart 1982). It has also been hypothesized that bubble cur-
tains have increased effectiveness if used with a light source
(Patrick et al. 1985). Previous studies using light and bubble
curtain barriers have proved successful in redirecting a num-
ber of different fishes. Strobe lights alone deterred Largemouth
Bass Micropterus salmoides, Chinook Salmon Oncorhynchus
tshawytscha, and Yellow Perch Perca flavescens (Richards and
571
Downloaded by [Heather Stewart] at 20:28 04 June 2014
572 STEWART ET AL.
Chipps 2007). The highest deterrence occurred when a strobe
light and a bubble curtain were used in combination, as seen with
Atlantic Menhaden Brevoortia tyrannus, White Perch Morone
americana, and Spot Leiostomus xanthurus (Patrick et al. 1985;
McIninch and Hocutt 1987). However, other studies reported
that some fishes were attracted to such barriers (Patrick et al.
1985; Sager et al. 1987). Age-0 Rainbow Smelt Osmerus mor-
dax were repelled by strobe lights, whereas juveniles and adults
were not (Stafford-Glase and Homa 1997). Our objective was to
examine whether a bubble curtain, strobe light, or a combination
of these two would be effective in reducing dam escapes of age-
0 Muskellunge. We examined the potential of these approaches
using a scaled replica of a spillway in a controlled laboratory
environment.
METHODS
Simulated dam and spillway.—In the laboratory, using flow
over a simulated barrier we examined a variety of variables
for their effect on age-0 Muskellunge movement. We created a
spillwayina460 × 100 × 50-cm fiberglass tank by blocking
one end with a notched (2 × 18 cm) board, simulating a dam
following the design of Wolter et al. (2013). Pumps (
1
/
4
hp,
0.76 L/s) were used to move water from below the spillway back
to the other end of the tank creating a closed loop (Figure 1).
The upper meter of the tank was partitioned with a net so that
fish in the trial arena could not encounter the area where pumps
discharged water. Spillway overflow height has been identified
as an important determinant of escape (Lewis et al. 1968; Paller
et al. 2006), so overflow height in this system was greater than
the body depth (1.25–1.75 cm) of an age-0 Muskellunge, and
water velocity at the face of the simulated dam (6 cm/s) was
comparable to velocities observed at the mouth of the spillway
at Lake Sam Dale, Illinois (4 cm/s) at similar levels of overflow.
Water depth was 34 cm in the trial arena when pumps were
running, whereas the water level in the catch basin was 24 cm,
which prevented fish from moving back into the test arena after
escaping. Flow was low enough that a refuge was not needed
as some fish did not pass over the dam. The trial arena was
378 L in volume and 198 cm long, an approximate distance of
15 body lengths for an average fish used in the trials. Three
identical tanks were used for replication and housed indoors.
Tanks were surrounded by opaque curtains to prevent fish from
being disturbed during the trials. A single age-0 Muskellunge
(120–165 mm TL) was randomly selected from one of four
pools of fish (∼200 individuals) and was placed into the arena
and allowed to acclimate for 1 h under trial conditions with
no flow over the barrier. After acclimation, the pumps were
activated to begin the 2-h trial. Treatments included the presence
and absence of a strobe light, presence and absence of a bubble
curtain, both in combination, and a control with no barriers.
Muskellunge escapes over the 2-h trial period was recorded.
Because diel period is an important factor of escape (Wolter
et al. 2013), we conducted day and night trials. Each of the
three barrier treatments, as well as day and night trials, were
replicated 15 times in a fully factorial design. Controls were run
with no strobe light or bubble curtain both during the day and
at night. After a trial, individuals were placed in a separate tank
so they would not be reused within 48 h.
Bubble curtain and strobe light.—The bubble curtain, which
was placed 110 mm from the spillway was created with a sec-
tion of PVC tubing, 500 mm long and 20 mm diameter, that
had 0.39-mm holes spaced 16 mm apart. The curtain was con-
nected by tubing to an air compressor delivering 984.3 kg/m
2
of pressure. The strobe light used was a Zeagle Underwater
Strobe with a xenon flash (60 flashes per minute) and visibility
of 3.2 km. The strobe light was angled to point into the bubble
curtain 390 mm from the spillway ( Figure 1), similar in scale to
FIGURE 1. Schematic of the simulated spillway system used to examine the effects of bubble curtain, strobe light, and their combination for reducing escapes
of Muskellunge. Dimensions of the system were 4.6 m long × 1.0 m wide × 0.5 m deep. The bubble curtain was 110 mm from the spillway and 500 mm across,
and had a width of 20 mm. The strobe was placed 390 mm from the spillway. Escape was determined when a fish passed over the dam into the catch basin as
monitored by a closed-circuit television camera, except for bubble curtain trials.
Downloaded by [Heather Stewart] at 20:28 04 June 2014
STROBE LIGHTS AND BUBBLE CURTAINS TO DETER DAM ESCAPES 573
laboratory studies by Sager et al. (1987, 250 mm) and McIninch
and Hocutt (1987, 300 and 850 mm). A single-plane wall of
bubbles was created that was illuminated by the strobe light.
Behavior trials were recorded using video cameras (AUC-75;
Atlantis Camera, Englewood, New Jersey) positioned 2 m above
the water surface of the simulated dam. Cameras were connected
to camera junction boxes, which sent video feed to video cas-
sette recorders (VCRs) to assess behavior during movement
of Muskellunge. The bubble curtain disrupted the surface of the
water so only control and strobe light trials were video-recorded
during the day. Trials were assessed using frame-by-frame anal-
ysis (30 frames/s) (Wagner et al. 2009). The behaviors assessed
were number of approaches before escape (defined as an indi-
vidual coming within one body length of the spillway), time
to escape, and activity prior to escape. A fish was classified as
making multiple approaches if it was observed to retreat before
approaching again. For activity prior to escape, both distance
from the spillway and body orientation of each individual ap-
proach was evaluated. Body orientation was recorded as degrees
from spillway, and as either head first or sideways. Head first
was defined as an angle of 45–135
◦
to the spillway and sideways
was defined as a 0–45
◦
or 135–180
◦
angle. Time to escape and
number of approaches were both quantified with two-sample
t-tests between escaping and nonescaping fish assuming equal
variances. For night and bubble curtain trials, visual observa-
tions were recorded to determine whether the fish went over the
spillway and, if so, at what time. Approaching behavior was not
recorded for night and bubble curtain trials.
Logistic multiple regression analysis (PROC LOGISTIC
SAS 9.2) was used to determine the variables that had an effect
on rates of escape. Categorical variables—diel period, strobe
light, and bubble curtain—were coded as either 1 or 0 based on
presence or absence, respectively. Length was used as a contin-
uous covariate. Fish were classified as having either “escaped”
or “not escaped” during the 2-h trials resulting in a binomial
response variable, 1 or 0 to code escape or no escape, respec-
tively, and significance was determined at α = 0.05. Percent
concordance was used as a measure of goodness of fit.
RESULTS
Fish length varied over a relatively narrow range and did
not differ between treatments (chi-square = 0.31, df = 1, P =
0.58). Diel period had an effect on escape and rates of escape
were higher during the day than at night (chi-square = 29.43,
df = 1, P < 0.01; Table 1). An explanatory model determining
the probability of escape under various conditions demonstrated
how diel period affected the probability of escape, with escapes
occurring more than twice as frequently during the day than at
night (Figure 2). There was also an effect of strobe on escape
with presence causing increased probability of escape compared
with a control (chi-square = 4.21, df = 1, P = 0.04; Table 1; Fig-
ure 2). The presence of bubbles did not reduce escapes compared
with controls (chi-square = 0.29, df = 1, P = 0.60; Table 1) nor
TABLE 1. Results of logistic regression analysis on Muskellunge escapes
during trials in a simulated spillway. The equation predicts a logit with a prob-
ability (p) that can be computed as p = 1/[1 + exp(logit)]. Presence–absence
of ambient light, strobe light, and bubbles were examined as factors influenc-
ing escape. Statistical significance is indicated by p-values in bold text with an
asterisk (*); no interactions were statistically significant (α = 0.05).
Wald chi p > chi
Parameter df Estimate SE square square
Intercept 1 1.20 0.45 7.28 <0.01*
Diel period 1 −2.73 0.50 29.43 <0.01*
Strobe light 1 −0.96 0.47 4.21 0.04*
Bubble curtain 1 −0.25 0.46 0.29 0.60
did the strobe light–bubble curtain combination (chi-square =
0.08, df = 1, P = 0.78). Escape was not affected by any interac-
tion of variables, including between diel period and strobe light
presence (chi-square = 0.62, df = 1, P = 0.43). Although not
statistically different, night time rates of escape were 47% with
strobe light (SD = 18.0) and 25% without strobe (SD = 12.5).
Fish Behavior
All escapes that were observed occurred within the first
20 min of the 2-h trials. Type and number of approaches did
not differ between strobe trials and control. The time to escape
after a trial began was similar for both strobe (mean, 8.9 min)
and no strobe (mean, 5.1 min; t-test: P = 0.15). Number of ap-
proaches to the spillway prior to escape did not differ between
strobe (mean, 1.3) and no strobe (mean, 2.0) trials (t -test: P =
0.15). Most (90%) escaping fish approached head first with the
body perpendicular to the spillway. The majority of the individ-
uals that escaped swam parallel to the spillway, then stopped
FIGURE 2. Probability of Muskellunge escape from the simulated spillway
during day and night with and without the strobe barrier present. Diel period
had an effect on escape (P < 0.01), and there was also an effect of strobe on
escape (P = 0.04); however, there was no significant interaction (P = 0.43).
Downloaded by [Heather Stewart] at 20:28 04 June 2014
574 STEWART ET AL.
and hovered before turning their head towards the spillway and
swimming over.
DISCUSSION
Strobe lights, bubble curtains, or their combination in a sim-
ulated spillway did not serve as a successful barrier to age-0
Muskellunge. In fact, the strobe light increased the number of
age-0 Muskellunge escapes. Positioning of the strobe light and
bubble curtain left a portion of the trial arena opposite of the
spillway as a refuge for fish that wanted to avoid these stimuli,
suggesting that the Muskellunge approaching were not merely
trying to escape the stimulation. Fishes at lower trophic lev-
els such as Alewife Alosa pseudoharengus, Rainbow Smelt,
Gizzard Shad Dorosoma cepedianum, White Perch, Spot, and
Atlantic Menhaden can be deterred by strobe light–bubble bar-
riers (Patrick et al. 1985). Muskellunge may react differently to
these stimuli than do other fishes as a result of species-specific
behavioral patterns (Bibko et al. 1974; Patrick et al. 1985;
McIninch and Hocutt 1987; Sager et al. 1987). Striped Bass
M. saxatilis have been observed to have only a temporary de-
terrence from strobe light (Bibko et al. 1974). Several other
studies have documented patterns of strobe light attraction in
select fishes, indicating that light may increase foraging effi-
ciency for visually foraging fishes (Brett and McKinnon 1953;
Alveras 1974; Hocutt 1980; Fiest and Anderson 1991; John-
son et al. 2003; Richards and Chipps 2007). Muskellunge are
visual predators (New and Kang 2000), which could explain
the observed attraction response to strobe light. There are many
factors that could account for the disparity in escapes among
fishes. Responses to light barriers may be species-specific due
to differences among feeding strategies, swimming activity and
ability, differences in visual systems, diel activity patterns, and
habitat selection (Patrick et al. 1985; McIninch and Hocutt 1987;
Sager et al. 1987). Variation in response to artificial illumination
can depend on light intensity, color, order of color presentation,
length of exposure, and mode of intensity change (Fields and
Finger 1956; Patrick 1982; Patrick et al. 1985; Sager et al. 1987;
Marchesan et al. 2004). Strobe lights have proven to be the most
successful of flashing lights, but the flash frequency is important
to be effective (Sager et al. 1987). Strobe light efficacy in causing
a startle response depends on a flash frequency distinguishable
from any natural light fluctuations observed underwater due to
waves or cloud movement (McFarland and Loew 1983; Schech-
ner and Karpel 2004). Because no previous studies using strobe
lights had been performed on Muskellunge, we used a flash rate
of 60 per minute to assess avoidance response. The Muskellunge
were not deterred by the low frequency strobe light, so future
studies should assess avoidance of higher flash frequencies.
Muskellunge escaped more often during daylight hours than
at night. Similar daylight escape patterns have been observed
with Largemouth Bass, whereas Black Bullhead Ameiurus
melas in the same study showed increased escapes at night
(Lewis et al. 1968). These escape patterns may be related to
foraging behavior and activity levels of the different fishes. The
escaping bullheads were primarily age-0 fish, which exhibit two
feeding periods, one at dawn and one at dusk, and the majority
of escapes were at these times (Darnell and Meierotto 1965;
Lewis et al. 1968). Largemouth Bass have higher foraging suc-
cess during daylight hours (McMahon and Holanov 1995), thus
the higher rate of escape during the day may be related to for-
aging activity. Since Muskellunge are primarily visual foragers,
it might be expected that they would exhibit similar behavior in
terms of daytime escape as other visual hunters such as Large-
mouth Bass (New et al. 2001).
The level of avoidance Muskellunge would exhibit towards
bubble barriers was uncertain given the limited research on Es-
ocidae with nonphysical barriers. Previous literature also pro-
vided little insight to possible responses due to the few trials
with predatory fishes. Piscivorous fishes such as White Perch
and Striped Bass were observed to have limited avoidance of
bubble barriers alone (Sager et al. 1987), which suggests that
these responses could be based on foraging behavior. However,
pelagic fishes such as Alewife, Rainbow Smelt, and Gizzard
Shad were found to be repelled by bubble barriers, whereas some
demersal fishes such as White Sucker Catostomus commersonii
and Spot were attracted (Patrick et al. 1985). The combination
of strobe light with bubble curtain barriers has demonstrated an
improved avoidance for some pelagic fishes (Sager et al. 1987).
We used age-0 Muskellunge that may not exhibit the same es-
cape behaviors as older adult fish. However, Wolter et al. (2013)
found a correlation between escapes in laboratory studies of
immature Muskellunge and field trials of adult Muskellunge,
suggesting our results on effects of nonphysical barriers could
also be applied to adult fish.
Muskellunge in these trials behaved similarly as they did
during tank acclimation, where fish were suspended in the wa-
ter column and moved slowly about the tank. Darting or other
types of burst swimming behavior were very rare. Observations
of Muskellunge behavior suggest that active swimming was in-
volved in escapes and it was not just the flow pulling the fish
over the spillway (Wolter et al. 2013). The majority of the indi-
viduals that escaped swam parallel to the spillway, before they
turned and swam over the barrier. Similar approach behavior
has been observed in flatfishes (Lemon Sole Microstomus kitt,
European Plaice Pleuronectes platessa, and Common Dab Li-
manda limanda) when electrified barriers were used (Stewart
1982). Frequently, the fish would swim parallel to the barrier,
turn away to retreat, and then approach the barrier again. These
behaviors were thought to be testing the barrier to find out if
it was harmless, and when no harm appeared to come from
the barrier, the fish quickly escaped. A similar response was
observed in age-0 Muskellunge.
Before installing light and bubble curtain barriers, response
to a range of stimuli and intensity for targeted fishes should
be understood to maximize barrier effectiveness. Future studies
should also assess sound barriers for Muskellunge, as some
fishes are more sensitive than others to particular wavelengths
Downloaded by [Heather Stewart] at 20:28 04 June 2014
STROBE LIGHTS AND BUBBLE CURTAINS TO DETER DAM ESCAPES 575
and frequencies (Sager et al. 1987; Mann et al. 2007). Strobe
light and bubble curtain barriers have attracted interest from
fish managers because of their simplicity and low costs, but our
study suggests these barriers will not be effective at reducing
spillway encounters for age-0 Muskellunge.
ACKNOWLEDGMENTS
Funding for this project was provided by the Hugh C.
Becker Memorial Foundation and from Federal Aid in Sportfish
Restoration Act Project F-151-R. We thank L. Dunham and S.
Pallo who coordinated activities with the Illinois Department of
Natural Resources. We thank Steve Miranda for his statistical
help. We also thank L. Einfalt for the care of fish and laboratory
assistance, as well as C. DeBoom for assistance with experimen-
tal design and three anonymous reviewers for providing helpful
comments on this manuscript.
REFERENCES
Alveras, R. A. 1974. Status of air bubble fish protection system at Indian Point
Station on the Hudson River. Pages 289–291 in L. D. Jensen, editor. Entrain-
ment and intake screening. Proceedings of the second entrainment and intake
screening workshop. Electric Power Research Institute, Baltimore, Maryland.
Bibko, P. N., L. Witrenan, and P. E. Kuester. 1974. Preliminary studies on the
effects of air bubbles and intense illumination on the swimming behavior
of the Striped Bass (Morone saxatilis) and the Gizzard Shad (Dorosoma
cepedianum). Pages 293–304 in L. D. Jensen, editor. Entrainment and in-
take screening. Proceedings of the second entrainment and intake screening
workshop. Electric Power Research Institute, Baltimore, Maryland.
Brett, J. R., and K. D. McKinnon. 1953. Preliminary experiments using lights
and bubbles to deflect migrating young spring salmon. Journal of the Fisheries
Research Board of Canada 10:548–559.
Darnell, R. M., and R. R. Meierotto. 1965. Diurnal periodicity in the Black Bull-
head, Ictalurus melas (Rafinesque). Transactions of the American Fisheries
Society 94:1–8.
Fields, P. E., and G. L. Finger. 1956. The effectiveness of constant and intermit-
tently flashing light barriers in guiding young Silver Salmon. University of
Washington, School of Fisheries, Technical Report 22, Seattle.
Fiest, B. E., and J. J. Anderson. 1991. Collected bibliography for review and
design criteria of behavioral fish guidance systems. University of Washington,
Seattle.
Hocutt, C. H. 1980. Behavioral barriers and guidance systems. Pages 183–205 in
C. H. Hocutt, J. R. Stauffer Jr., J. E. Edinger, L. W. Hall Jr., and R. P. Morgan
II, editors. Power plants: effects on fish and shellfish behavior. Academic
Press, New York.
Johnson, R. L., M. Simmons, C. S. Simmons, C. A. McKinstry, C. B . Cook, S. L.
Thorsen, R. LeCaire, and S. Francis. 2003. Strobe light deterrent efficacy and
fish behavior determination at Grand Coulee Dam third powerplant forebay.
Pacific Northwest National Laboratory, PNNL-14177, Richland, Washington.
Lewis, W. M., R. Heidinger, and M. Konikoff. 1968. Loss of fishes over the
drop box spillway of a lake. Transactions of the American Fisheries Society
97:492–494.
Mann, D. A., P. A. Cott, B. W. Hanna, and A. N. Popper. 2007. Hearing in
eight species of northern Canadian freshwater fishes. Journal of Fish Biology
70:109–120.
Marchesan, M., M. Spoto, L. Verginella, and E. A. Ferrero. 2004. Behavioural
effects of artificial light on fish species of commercial interest. Fisheries
Research 73(1-2):171–185.
Margenau, T. L. 1992. Survival and cost-effectiveness of stocked fall fingerling
and spring yearling Muskellunge in Wisconsin. North American Journal of
Fisheries Management 12:484–493.
McFarland, W. N., and E. R. Loew. 1983. Wave produced changes in underwater
light and their relations to vision. Environmental Biology of Fishes 8:173–
184.
McIninch, S. P., and C. H. Hocutt. 1987. Effects of turbidity on estuarine fish
response to strobe lights. Journal of Applied Ichthyology 3:97–144.
McMahon, T. E., and S. H. Holanov. 1995. Foraging success of Largemouth
Bass at different light intensities: implications for time and depth of feeding.
Journal of Fish Biology 46:759–767.
New, J. G., L. A. Fewkes, and A. N. Khan. 2001. Strike feeding behavior in the
Muskellunge, Esox masquinongy: contributions of the lateral line and visual
sensory systems. Journal of Experimental Biology 204:1207–1221.
New, J. G., and P. Y. Kang. 2000. Multimodal sensory integration in the strike-
feeding behaviour of predatory fishes. Philosophical Transactions of the Royal
Society of London Series B Biological Sciences 355:1321–1324.
Paller, M. H., D. E. Fletcher, M. M. Standora, T. B. Grabowski, T. A. Jones, S. A.
Dyer, and J. J. Isely. 2006. Emigration of fish from two South Carolina cooling
reservoirs. North American Journal of Fisheries Management 26:976–982.
Patrick, P. H. 1982. Responses of Alewife to flashing light. Ontario Hydro
Research Division, Report 82-305-K, Toronto.
Patrick, P. H., A. E. Christie, D. Sager, C. Hocutt, and J. R. Stauffer Jr. 1985.
Responses of fish to a strobe light/air bubble barrier. Fisheries Research
3:157–172.
Plosila, D. S., and B. D. White. 1970. A swinging vertical screen for fish barrier
dams. Progressive Fish-Culturist 32:178–179.
Richards, N. S., and S. R. Chipps. 2007. Stress response and avoidance behav-
ior of fishes as influenced by high-frequency strobe lights. North American
Journal of Fisheries Management 27:1310–1315.
Sager, D. R., C. H. Hocutt, and J. R. Stauffer Jr. 1987. Estuarine fish responses
to strobe light, bubble curtains and strobe light/bubble-curtain combinations
as influenced by water flow rate and flash frequencies. Fisheries Research
5:383–399.
Schechner, Y. Y., and N. Karpel. 2004. Attenuating natural flicker patterns. Pages
1261–1268 in Oceans 2004 MTS/IEEE (Marine Technology Society/Institute
of Electrical and Electronic Engineers) Oceans ‘04 conference 3. MTS/IEEE,
New York.
Simonson, T. D., and S. W. Hewett. 1999. Trends in Wisconsin’s Muskellunge
fishery. North American Journal of Fisheries Management 19:291–299.
Stafford-Glase, M., and J. Homa. 1997. An evaluation of fish entrainment and
the effectiveness of the strobe light deterrent system at Milliken Station on
Cayuga Lake, Thompkins County, and New York. Prepared for New York
State Electric and Gas Corporation, Binghampton.
Stanford, J. A., J. V. Ward, W. J. Liss, C. A. Frissell, R. N. Williams, J. A.
Lichatowich, and C. C. Coutant. 1986. A general protocol for restoration
of regulated rivers. Regulated Rivers Research and Management 12:391–
413.
Stewart, P. A. 1982. An investigation into the reactions of fish to electrified
barriers and bubble curtains. Fisheries Research 1:3–22.
Von Brandt, A. 1967. Application of observations on fish behavior for fishing
methods and gear construction. FAO (Food and Agriculture Organization of
the United Nations) Fisheries Report 62:169–191.
Wagner, C. P., L. M. Einfalt, A. B. Scimone, and D. H. Wahl. 2009. Effects
of fin-clipping on the foraging behavior and growth of age-0 Muskellunge.
North American Journal of Fisheries Management 29:1644–1652.
Wahl, D. H. 1999. An ecological context for evaluating the factors influencing
Muskellunge stocking success. North American Journal of Fisheries Man-
agement 19:238–248.
Wolter, M. H., C. S. DeBoom, and D. H. Wahl. 2013. Field and laboratory
evaluation of dam escapement of Muskellunge. North American Journal of
Fisheries Management 33:829–838.
Downloaded by [Heather Stewart] at 20:28 04 June 2014