North Amsncwi J o u r ~ olf Firhcrlar MuHQ&WWU
19:793-803. 1999
0 Copyrighl hy tho Amodcnn Fisheries Society 1999
Attraction and Passage Efflciency of White Suckers
and Smallmouth Bass by Two Denil FIshways
Wnrcrloo Riotelemetry Institute, Deparrmenr of Biology, IJniver,rity of Waterloo,
Waterloo, Ontario N2L 3G1, C a d
Depurtn~enrof F i s h e r i ~ sand Oceans, Central and Arctic Region.
Winnipeg, Munitoha R3T 2N6, Conaah
Waterloo Biot~lemetryInstitute, Depnrlmenf r$ Biology. University of Wulerloo.
Waterloo, Onlario N21, 3G1, Cunada
Ahslrurbt.-We compared two Uenil fishways, locatcd on the west (low velocity. 10% slope) and
east (high velocity, 20% slope) sides of the Mannhcim weir. Grand River, OnLurio, for use by
upstream-migrating white suckcrs Catostomus commersoni and smnllmouth bnfis Micropterus doIornieu. Mark-recapture and rndiotclcmctry were used to assess attraction and fish passagc. Moverncnt of 85 radio-lugged fish was monitored continuously during s p r i n ~and early sulnmcr of 1995
and 1996. Attrnction and passage efficiencies of white suckers at thc west hshway were upproxinlately 50% and 5 5 % , respectively. Attraction efficiency of whitc suckers at the east fishway was
approximately 59%. and passage efficicncy was 38%. The altrnction and passage efficiencies of
smnllmouth huss at the west fishway were approximately 82% and 36%. respectively. At thc cnst
fishway, attraction efficiency of snldlmouth bass was npproximalely 5 5 % , and passagc efficiency
was 33%. Thcrc was an exponential decline in the numbers of both species Lhat used each fishway
relative to watcr velocity. The maximum water velocity used by white suckers was 0.96 mls and
that used by smallmouth bass was 0.99 m/s. Distracting flows near thc wcst fishwny nppeurtd to
aflect attraction. Both fishways passed equal numbers uC smallmouth bass per year, nnd ~lnntlrnouth
hnss that used the east fivhway were significantly larger thnn individtlals l h ~ used
l
the wcst fishway.
In contrast, more lhan twice as numy white suckers U R C the
~
west flshway, and these fish were
significnntly larger than those lhat used the east fishway. Differcncce in passage were related to
burst and critical swimming speeds and the use of velncily rerugiu within the fishways.
Fishways allow upstream-migrating fish to bypass natural and artificial rivcr barricrs (Beach
1984; Clay 1995). Biologically oriented fishway
research has focused on anadromous fishes, such
as salmonids and clupeids. As a result, fishways
do not usually include defiign fcaturcs that are relevant to the behavior and swimming pcrformance
of freshwater fish (Lucas and Frear 1997). This
information is vital if fishways designed to accommodate heshwatcr spccies are to be successful.
Dams m d othcr river barricrs increasingly compromise fish movements in temperate and k o p i c ~ l
streams and rivers. Many families of warmwater
fishes including ccntrarchids, catoslomids, csucids. percids, cyprinids and ictalurids migrate up
streams and rivers, cspccially during spawning pc~ o d s(Raney and Wcbster 1942; Schultz 1955;
* Corresponding author:
smckinle@scihorg.uwaterIoo.ca
Rweived June 24, 1998; accepted February 27. 1999
Rawson 1957). Some members of thcsc families
are known to use Rshways (Nclson 1983; Derlrsen
1988; Langhurst and Schocnikc 1990; Harris and
Mallen-Cooperl994; Lucas and Frear 1997). Fishway use varies, huwevcr, according to fishway design, water velocities within the fishways, water
temperature, time of day, fish size, and swimming
ability (Femet 1984; Schwalme ct al. 1985; Derksen 1988; Pavlov 1989; Katopodis et al. 1991),
Passagc efficicncy of a limited number of coolwater, coldwater, and warmwater fishes has been
investigated using mark-recapture (Linlokken
1993) and videography (Dexter and Ledet 1996;
Haro and Kynard 1997). Radiotelemetry has not
been used to track individual movements and hchavior of nonsalmonid fishes in Cshways to dcterrninc attraction and passage efficicncy.
Thc objectives of this study wcrc to describe
conditions during iishway usc und to quantify attraction efficiency and passagc cfficicncy of white
suckers Catostomus commersoni and smallmoulh
794
RUNT ET A L
ONTARIO
Lake Ontarlo
NORTH
FICX~WE
I.-The
location of the Mnnnhelm weir on the Grand River, nenr Kitchcner. Ontnrio. Approximntely
6,700 km2 nrc draincd into Lakc Eric by thc Grnnd River watershed.
bass Micropterus dolomieu downstream from a rcccntly constructed weir equipped with two Dcnil
fishways. We chose to study whitc suckers and
smallmuuth bass hecause they are morphologically
dislinct and differ in swimming ability. The primary null hypothcscv for cach species wcrc ( I )
east iishway attraction efficiency equals west fishway attraction efficiency, and (2) east fishway passage efficiency equals west fishway passage efficiency. We also measured the critical swimming
speeds of adult smallmouth bass and used similar
data for white suckers from the litcrature to hclp
cxplain successful passagc relative to hydraulic
conditions within the two fishways.
Methods
Study area.-This
utudy was conductcd at the
Mannheirn weir on the Grand River, near Kitchcncr, Ontario. The Orand River is a mid-order
stream that flows 297 k n ~from its source in Dundalk, Ontario, to the eastern basin of Lake Erie
(Smith 1994). The Mannheim weir is located approximately midway along the river (Figure 1) and
creates an impoundment for the extraction of regional drinking water. Mean depth downstream
from the weir is approximately 0.5 m, mean annual
discharge is approximately 33 mVs, and primary
subslrales consist of cobble and broken rock (Bunt
ct al. 1998). The construction of the 2.2-m high
weir and fishways was completed in 1990. Characteristic of Dcnll fishways. those at the Mannheim
weir use baffles to turn the flow of watcr back on
itself to reduce the velocity of a primary flow,
through which fish must swim. Prior to 1990, fish
movements were not restricted at this site.
To allow upstream passage of fish, n 27-m Denil
fishway that doubled back on itself twice was constructed from reinforced concrete along the west
bank of the river (Figure 2). Each of three parallel
flumes (slope 10%) were fitted with metal baffles
spaced approximately 25 cm apart. On the east
bank US the river, a much simpler and less expensive Denil fishway was cunfitructcd. It consistcd
of a single 11-m reinforced concrete flumc with
baffles along a 20% slopc (Figure 2 ) . All flumes
were 0.6 m wide and 2.15 m dccp and werc complctcly covcrcd with removable steel plates.
Telemetry.-Digitally coded radio tags (1.9 g in
watcr; 10.6 Inm X 28 mm X 7 mm; 2 . 5 s pulse
rates) were externally attached to 32 white suckers
and 53 smallmouth bass exceeding 300 rnm total
length. External tagging was chosen because most
fish wcre ready to spawn and therefore considered
unfit for surgical implantation without the risk of
affecting normal behavior. We tagged equal proportions of malts and femalcs of each species. Thc
FIGLIRE
2.-Schcmntjc
weir. Shaded arcns in Ihr
Rclntive velocity isoplet
Sigma PVM velocity me
ratio between tag wcii
than 2%, as recornmen
a bricf recovery period
and fish were releaser
approximately 150 m
Movemcnt of radiowhite suckers was de
seven stationary submr
and within thc fishwa
tioned within cnch of r
fishway and another w:
the east fishway. One 1
at the entrance of cacl
tenna was positioned b
e6 to record back and
antennas wcre scanned
signals. A receiver anc
(SRX 400 and DSP 5
Engineering Limited) ,
antenna switching cap:
minc transmitter loca
point of the nearest ar
ination was used to id,
strength calibrations a
terrnine the distance b
cciving antennas lo w
ccption areas at the u
Sigma PVM vclocily meter.
ratio between tag weight and fish weight was less
than 2%, as recommended by Winter (1983). After
a bricf rccovcry period, each transmitter was tested
and fish were released at randomly chosen sites,
approximately 150 m downstream from the weir.
Movement of radio-tagged smallmouth bass and
white suckers was determined using an array of
scvcn stationary submerged antennas near the weir
and within the fishways. OIK antenna wa8 positioned within each of thc tlucc flumes of thc west
fishwny m d another was positioned midway within
the east fishway. One antenna was also positioned
at the entrance of each fishway. and the final antcnna was positioned bctwccn the fishway cntrancos to record back nnd forth movcmcnt. All seven
antcnnas were scanned cvcry 7.5 ms for telemetry
signals, A receiver and digital spectrum processor
(SRX 400 and DSP 500., respectively, of Lotek
Engineering Limited) equipped with fast multiple
antenna switching capabilities were used to determine transmitter locations relative to the focal
point o f the ncarcst antenna. Pulse-code discrimination was used lo identity valid signals. Signalstrength calibrations allowed us to accurately determine the distance between transmitters and r t ceiving antcnnas to within 0.3 m. The signal reception areas at the weir were verified regulnrly
796
HUNT ET A L
ging a large number of both species and comparing
their recapturc patterns with radio-lagged fish. We
were not able to collect all fish for anchor-tagging
and radio attachmenl from the fishway traps.
Thcrcfore, by a ~ g l i n gwe captured 11 smallniouth
bass ncar the fishway cntrances and statistically
compared their use of the fishways with smallmouth bass that were caught in the fishway traps.
If no differences wcrc dctcctcd bctwcen thc numbcr of anglcd versus trapped smallmouth bass that
used the fishways, data for smallmouth bass were
pooled.
Mark-recapture and data anu1ysis.-Fish that
succcsvfully asccnded each of the two fishways
were trapped just before the upstream exit in fishway traps that consisted of aluminum blocking
screens (mesh size 1.5 ctn) and removable aluminum funnels made from 2-cm wire mesh (Figurc
2). Inslallation and monitoring of fishway traps
began in mid-April before m y fish werc obsorvcd
near the weir. Water ternpcraturcs werc recorded
each timc fisliway traps werc checked and at midday using a hand-held themlometer at the west
fishway cntrancc. The Grand Rivcr is wcll mixed,
and there is no difference in water tcnlpcrature on
either side of the weir. When the watcr ternperaturc
was warm enough, while suckers were trapped in
the Iishways and smallmouth bass were either
(rapped in the fishways or angled near the fishway
entrances. These fish were either radio-tagged, or
externally marked (Floy FD-94 anchor tag), and
released at randomly chosen release sites, downstream. Fishway traps wcrc chcckcd and cleared
daily between 0900 and 1400 hours and bctwccn
1600 and 0200 hours. All trapped fish wcrc removed from the fishway traps with a dipnet, enumerated, and measured for total length (TL, mm).
Most were tagged and released downstream. In all
situations, fish that returned to either fishway trap
more than once were released upstream from the
weir. The mean total lengths of white suckers and
smallmouth bass, including recaptured fish collected from the fishway traps but excluding juveniles (<200 rnm for both species [Scott and
Crossman 1973]), wcrc coniparcd with one-way
analysis of variance (ANOVA). Nurnhers of
anchor-taggcd and radio-tagged fish that were subsequently recaptured in the fishway traps wcrc
cotnpnred with chi-square contingency analysis.
Radio-tagged smallmouth bass and white suckers
that were recaptured or detected in the fishway
traps were considered to have mndc a successful
attempt at using a fishway. Attempts to use a fishway occurrcd wlicn radio-taggcd fish wcrc de-
tected more than 2 m inside either fishway entrance. Attraction and passage efficiencies of both
species were compared at each fishway using 2 X
2 contingcncy analysis for proportions wilh corrections for continuity, and the power of the pcrformed test was detcrmincd according to the mcthods in Zar (1984). All statistical tests were considered significant at an alpha level of 0.05, and
all means are reported 1 SE.
Fishway conditions.-Denil
fishway water
depths, discharge, velocities, and water surface
profilcs arc interdcpcndcnt and relationships bctween thcm have been dcvclopcd through hydraulic model studies (Katopodis and Rajaratnam
1983; Rajaratnam and Katopodis 1984). Several
extcnsive laboratory investigations with scale and
prototype models of Dcnil fishways have been conductcd (Rajaratnam and Katopodis 1984; Katopodis et al. 19971, Tn thcsc studies, two modcl
scales (1:6 and 1:3) were examined and compared
with a prototype scale (1: 1) for the standard Denil
design, similar to those at the Mannheim weir. Thc
similarity of the results from the threc scales was
demonstrated, and the three data sets fit the same
curve. Results were also confirmed in a field study
(Rajaralnam et al. 1992), which proved the reliability of these rclationships and the usc of dcpth
rneasurcments to estimate fishway discharges and
velocitics. In addition, concordance betwccn estitnated velocities and actual velocities at the
Mannhcim weir was verified with dircct measurcmcnts of watcr vclocity within the fishways with
a Sigma PVM ultrasonic velocity meter. Water
dcpths wcre therefore measured directly a1 the
Mnnnhcim weir and used to calculnte vclocitics in
the two fishways.
Watcr depths downstream from each fishway
trap were recorded to produce a flow profile and
to estimate water velocitics from a velocity-rating
curve. The minimum velocities yielded by the rating curves wcrc 0.24 and 0.33 m/s for thc wefit
and east fishways. rcspectivcly. Using a calibratcd
rod, we measured the vertical distance (nearest cm)
to the water surface at a series of fixed points along
the rtshway. Water depths for the various measurement location^ wcrc cnlculatcd by subtracting
these field measurements horn the previously measured vertical distance to thc baffle crests. Turbulence at the surface of the primary flow compromises true estimates of vclocity (Katopodis and
Rajarainam 1983). Calculations of maximum water vclocity that any fish would expcricncc during
fishway usc wcrc thcrcforc dcrivcd for n region
corresponding lo 0.8 x water dcpth.
+
Swirnrnin~abilities
swimming speeds (U
mouth bass that wcre
for radio-tagging. CI
measure of prolonge
the maximum veloci
prescribed pcriod of
held without food fo
tion. All swimming I
ducted in ambient riv,
L Blazka-Fry respiro
to the raspirometer fo
approximated 0.5 bc
swimming speeds (m,
rnin increments betwt
ity increases of appro
ing to the procedure:
werc corrected (Smit
fecls caused by the flc
of largc fish within th
perature for all trial:
20°C, and all fish wcr
dark cycles to minirni
and temperature on cc
mance (Kolok 1991).
and total length were
analysis. lnformntion
suckers was derived
(Joncs et al. 1974).
Hydra~rlicConditinns
The mean watcr v,
(0.89 0.02 d s . rang
was significantly less
the east fishway (0.95
2.05 mls, N = 162, P
occasions when large
laled on the east fishw
accumulations of dcl
screens altered water
watcr velocitics was
throughout the study F
ways varied on a daily
and was cleared from
ations in flow and velc
after mid-June, when
lished upstream. Veloc
way traps and blockin
locilics were Iowcst a,
had accurnulatcd on bl
Maximum fishway us
velocities when some
the blocking screens.
*
USE OF 'TWO D E N l L F I S H W A Y S
Swimming abi1iries.-We determined the critical
swimming speeds (U crit) for Grand River smallmouth bass that were similar in size to fish chosen
for radio-tagging. Critical swimming speed is a
measure of proloqged swimming and represents
the maximum vclocity a fish can maintain for a
prcscribcd pcriod of time (Brett 1964). Fish wcrc
hcld without food for 48 h prior to cxpcrimentation. All swimming speed experimcntv wcrc conducted in ambient rivcr watcr using a modified 70L Blazkn-Fry respirorneter. Fish wcrc acclimated
to the respirometer for 24 h at water velocities that
approximated 0.5 body lengths (BL)/s. Critical
swimming speeds (m/s) were detcrmincd using 10
min increments between constant step-wisc vclocity increases of approximately 1-1.5 BL/s according to the procedures of Brett (1964). Velocities
were corrected (Smit et al. 1971) for blocking effects caused by the flow of watcr around thc bodies
of large fish within the swim tubc. The water ternperature for all lrialr; rangcd bctwccn 15°C and
20°C. nnd all fish wcrc acclimated to natural light:
dark cycles to minimize the cffccts of photoperiod
and temperature on centratchid swimnung perfornlancc (Kolok 1991). Relationships hetwcen U crit
and lotal length were determined using regresaion
analysis. Information of swinlming ability of white
suckers wns derived from thc rivailablc literature
(Jones et al. 1974).
Results
Hydruulic Conditions in Fishways
The mean water velocity in the wcst fishway
(0.89 +. 0.02 rnls, range c0.24-1.57 m/s,N =- 207)
was significantly less than the water vclocity in
thc cast fishway (0.99 -C 0.03 m/s, range <0.332.05 m/s. N = 162, P < 0.001), except during rare
occasions when large arnounts of debris accumulated on the east fishway blucking screen. Variable
accumulations of debris on upstrcam blocking
scrccns altered water depths so that a rangc of
water velocities was available in both fishways
throughout the study period. Velocitics in the fishways varied on a daily basis as debris accurnulatcd
md was cleared from the blocking screens. Variations in flow and velocity were most pronounced
after mid-June, when macrophytcs bccamc established upstream. Vclocitics were greatest after fishway traps and blocking screens were cleaned. Velocitics were lowest after large amounts of debris
had accurnulatcd on blocking screens after storms.
Maximum fishway use occurred at intermediate
velocities when some debris had accumulated on
the blocking screens.
White Suckers
Whitc suckcrs were first observed at the fishway
entrances and began to use both fishways on 3 May
1995 after which, midday water ternpernturcs were
consistently above 9°C. Maximum passage occurred on 3 May 1995 when 92 white suckers wcrc
captured in the west fishway trap, The last recorded
passage was on 15 July 1999 whcn two white suckers were caught in the east fishway and the watcr
tcrnpcrature was 23°C. In 1996, whitc suckcrs begnn to use the west fishway on 6 May, after which
water lemperaturcs wcrc consistcntly above 8°C.
Maximum passagc at the west fishway occumcd
on 17 May when 101 white suckers passed upstream and the watcr tcnipernture was 11°C. Whitc
~ u c k c r sbegan to use the east fishway on 9 May
1996 when the watcr tempcraturc was consistently
above 10°C. Maximum passage at the east fishway
occurrcd on 11 May 1996 when 76 whitc suckers
used thc fishway qnd the water tempcraturc was
10°C. The last recorded passage of white suckers
at the west and east fishways was on 9 July (18°C)
m d 15 July (21°C), respectively. White suckers
were caught most frcqucntly in the fishway traps
from 3 May lo 12 May 1995 and 6 May to 18 May
1996. Aggrcgutions of white suckers werc observed near the weir und fish often appeared to
follow one anothcr.
Whitc suckers were periodically within the vicinity of the weir (within 20 m) for 12 ? 2 d.
During this period 50% (95% CI = 33-67%) and
59% (95% CI = 42-76%) of radio-tagged white
suckers were attracted to the wcst and east fishways, respectively. Thc attraction efficiency o r
while suckers at both fishways was not significantly different ( Z = 0.78, P > 0.5). Radio-lagged
white suckers were detected near ihc weir and fishway entrances within 1 d of release (mean 0.75 &
0.25 d). Twenty-two fish approached the weir, and
45% wcrc first dctcctcd in the high velocity zonc
upstream from the west fishway entrance (Figure
2). Highly variahlc cxploratory movements (Pigurc 3a) as well as long periods with no movcmcnt
near the weir and fishwnys werc common.
Approximately half of all radio-tagged white
suckers that entered the wost fishway (N = 11)
swam to the exit. Passage efficiency of whitc suckers by the west fishwdy was estimated to be 55%
(95% CI = 25-84%). There were eight attempts
to use the east fishway, where passage efficiency
of whitc suckers was estimated at 38% (95% C1
= 4-71%). Passage efficiency at each fishway did
not diffcr statistically (Z = 0.17, P > 0.5). Over
BUNT ET AL
TABLEI .-Menn totnl
and July 1995 and 1996.
traps are shown. For tact
oC fish caught wcre simil
Species
TL
Srnallmouih b u ~ s
White suckers
321.
326.
fishways ~ c c u r r c dor
smallmouth bass wcrc
and the watcr temper;
corded passagc of srr
on 27 Junc (water te
June (18°C) for the w,
lively. In 1996, small
east fishway on 18 Ma
occurrcd on 18 May,
used the east fishway.
began on 20 May (16?
for smallmouth bass i:
TABLE
?.-Tracking
su
and pattcrns of use by wh
Variable
Nocturnal, west fishwsy
Daytime, waat tifihway
Nociurnul, enst fishway
Daytime, east fishway
1-4
1-1-
27-Jun-95
I~-Ju~.Q~
1
2
3
Position
4
1
2
3
4
Position
FI~UK
3.-Examples
E
of bnck and forth movements by (a) white suckcrs and (b) slnallmouth bass immediately
downslrcnrn from thc weir during the spring (IT 1995. Positions along the ubvcissa urc as follows: 1 = west f l ~ h w a y
entrance. 2 = high-velocity area in spillwny, 3 = rcgion between fishway entrances, and 4 = enst fishwny entrance.
69% o f total white sucker passage occurred at the
wcst fishway (Table 1). These white suckers were
significantly larger than whitc suckcrs that used
the east fishway (F = 25.75, df = 1328, P < 0.001;
Table 1). Approximately 27% of white suckcrs
caught in the west fishway usad velocities less than
0.24 mls. Similarly, approximately 70% of while
suckcrs used the east fishway when water velocities were less than 0.33 d s . Numbers of while
suckers using each fishway declined exponentially
relative to watcr velocity. Thc maximum water velocities used by white suckers was 0.96 d s . White
suckers wcrc located ncar both fishways at night,
but twice as many attempts to usc either fishway
occurred during daylight hours (Tablc 2).
Smallmouth Bass
Smallmouth bass began to use both fishways on
9 May 1995 when the water temperature was consistently above 10°C. Maximum passage at both
The!
In pro1 1 , west fishwey
In pool 2, wovl fishway
For full asccnl, west fishway
Fur full ascenL rasl fiuhway
Overall and succmr
Overull, wcst fishway
Succarr~ful.west finhwny
Overall, emt Rahwny
Succe~uful,east fishway
Passuge (P) and attm
95% con
A, waat tishwey
P. wmt tishwny
A, e m fishwuy
P, east fishway
a
Nocturnul attempts wcrc bet
Overall auampt~mny not eq
attempts hecnuse sumc fish
the west fishway.
799
USE OF TWO DENIL FISHWAYS
TABLE1.-Mean total length (ZSE)of srnnllrnouth buss and while ~ u c k e r scaught in the fishwny tmps bctween May
and July 1995 and 1996. Numbers of anchor-tagged nnd rudio-tagged f i ~ hand subsequent recaptures from the Ashway
traps rue shown. Fur each species, total lengths (TLR)wcrc significantly differen1 ( P < 0.05) between fishways, numbers
of fish cuught were similar for smallmouth b u s but significnntly diffcrcnt (P r' 0.05) for white suckers.
Nurnher of fish
Wost fi~hwny
East tishway
Species
TL (mmj
N
TL ( m m )
N
Anchortagged
Smnllmouth bnss
White suckcfi
326.8 Z 0.3
321.9 -C 2.2
37
919
748.8 2 7.2
302.3 2 3.2
48
410
51
818
fishwnys occurred on 5 June 1'195, when cight
h
trap
smallmouth bass were caught in e ~ c fishway
and the water temperature was 16°C. Thc last rccorded passage of smallmouth bass in 1995 was
on 27 June (water temperature = 20°C) and 17
June (18°C) for the west and east fishway, rcapcctively. In 1996. smallmouth bass began using the
cast fishway on 18 May (14°C). Maximum passage
occurred on 18 May, when nine smallmouth bass
uvod the cast fishway. Passage at the west Gshway
bagan on 20 May (16°C). The last recorded passage
for smallmouth bass in the west and east fishways
TABLE
?.-Trucking
summary, d i d use of the fishways,
md pnttclns of use by white suckers nnd smallmouih has.
While
wckeru
Snidl~uouth
bnvv
N (%) tracked nenr the flshwaya
Attempts lu
Ntrclurnul, WUHL fi~hwuy
Daytime, wevt flshway
Nocturnal, anst tivhwny
Daytime, ewl Ashw~~y
HshwuyR
In p o l 1 , weclt fishwuy
In pool 2, west R~hway
For full nuccnt, west tishway
For full a6canl. east hhway
1-20
5-1 76
12-85
4-6
2-80
3-1 140
43-1240
I
O v e d h snd s u c m h i sttcmpb to urn Ashwuy
Overall, west fishwny
II
11
Succos~ful,wa6t tishwny
6
4
Overnll, eual fiuhwuy
8
3
Successful, m t finhwuy
3
1
Psenage (P) and stbnctlon ( A ) elRclency (%) sud
95% cunfidcnce lntcrvnl
A. wevt tishwny
wcsl fihhwny
A, east fishway
P, cast fishwnv
'
' Nocturnd
attempts were helwwn 2000 nntl 0600 hours.
Ovtrull uclernpts may nut equnl the yuni of nucturnul nnd daytime
Itttemp tmcaure wmc finh wailed overnight in renting puuls ul'
the w-t fluhway.
Rndiotuggad
53
32
Number of
recaptures (%)
Anchor tng
Radio tug
3 (5.9)
42 (5.1)
2 (3.8)
1 (3.1)
was on 14 July and 15 July, respectively, when thc
water temperature was 21°C. Most snlallmouth
bass were caplured in the fishway traps from 15
May to 5 Junc 1995 and 18 May to 13 June 1996.
Frcqucnt freshets and abnormally cool June tempcraturcs interrupted smallmouth bass migrations
in 1996. Tagged and untagged smallmouth bass
werc oftcn observed in lhe high- velocity zone near
the west fishway entrance (Figure 2). Smallmouth
bass were first detected ncar the weir and fishway
entrances 3.0 ? 0.5 d after release. There was no
difference in time lo detection or in attempts to
use the fishways among lrapped and angled fish,
so data for smallmouth bass werc poolcd.
During the study, 82% (95% CI = 69-95%) of
smallmouth bass were attracted to the west fishway, and 55% (95% CI = 38-72%) were attracted
to the cast fishway. The attraction efficiency of
smallmouth bass by each fishway was not signiLicantly different (Z = 1.82. 0.1 < P < 0.05, power
= 0.43). As with white suckers, exploratory movemcnts were common (Figure 3b) and lasted for 16
-' 2 d for fish that did not successfully pass upstream from the weir. The number uC attempts to
use the fishways during the day exceeded nocturnal
use by a factor of five (Table 2).
Passage efficiency of mallmouth bass at the
west fishway was estimated to bc 36% (95% CI =
X -65%). Only thrcc radio-tagged s~nallmouthbass
entered the east fishway, and one swam upstream
to the exit. Therefore, east fishway passage cfficicncy of smallmouth bass was roughly cstimatcd
to hc 33% (95% CI = 0-87%). Passage efficiency
of smallmouth bass at the west fishway did not
appear to differ from passage efficiency at the east
fishway (2 = 0.08. P > 0.5). Eighty-five untagged
and tagged smallmouth bass were recovered from
the fishway traps, and 48 ( 5 6 % ) were caught in
the easl fishway (Table I). Equal number8 of smallmouth bass used each fishway ( x 2 = 1.42, P <
0.25), and equal proportions of radio-tagged and
anchor-tagged smallmouth bass wcrc rccapturcd in
800
BUNT ET AL.
the fishway traps (Table 1). More than 71% of
smallmouth bass used the wcst fishway when water
velocities werc less than 0.24 mls. At thc cast fishway a similar pattern emerged: 60% of smallmouth
bass uscd watcr velocities less than 0.33 m/s. Numbcrs of smallmouth bass using tach fishway declined exponentially relative lo water velocity. The
maximum water velocities used by smallmouth
bass in the wcst and cast fishway wtrc 0.72 and
0.99 m/s, respectively. Smallmouth bass that uscd
the east fishway were significantly larger than
smallmouth bass that used the west fishway (F =
4.93, df = 84, P = 0.03; Table I).
Critical Swimming Speeds
The range of critical swimming speeds recorded
for smallmouth bass 262-378 mm TL was 0.501.18 d s . Thcre was a highly significant positivc
relationship between critical swimming spccd and
total length of fish tested (N = 11, r = 0.43, P <
0.001). The relationship was best described as U
crit = 0.009534 X TL0.7mZ".
Discussion
Studying fishway use and effectiveness from a
biological perspective is a notoriously difficult
challenge (Beach 1984), and this study was no
exception. Various misfortunes confronted us during this project. For examplc, antennas werc tcmporarily severed twice from debris accumulation
during storms, which may have resulted in undetected attempts at rht east fishway. We were unable
to collect sufficient numbers of fish that were potentially naive of the weir, and this may have affected our estimates of attraction and passage.
Learning m o n g fish has not been proven to exist.
and we therefore feel that these effects were negligible. Our observations suggest that some fish
migratc upstrcam to barriers. Others find habitat
that suits them downstream from river barriers and
may never interact with the barriers. Clearly, some
fish within a spatially distinct area will naturally
relocate in the springtime while others will not. 11
is thcrefore impossible to know, a priori, which
individuals have a propensity for physical displaccmcnt. Even if fish for this study had bccn
collected from areas away from the weir, thcre
would bc no evidence they had not intcractcd with
the lishways or the weir prior to this work. As
such, it is extremely difficult to gauge the importance of fishways with random samples of fish collected downstream from rivcr barriers. Other problems that we encountcrcd whilc comparing thc
Mannheim fishways included the high velocity re-
gion upstream from the west fishway that distractcd some fish during the study. Also, the m o d e ~ t
degree of complete fishway use by radio-tagged
fish was insufficient for the application of powerful
statistical analyses. It is likely, based on limitcd
numbers of white suckers and smallmouth bass
that used each fishway annually, that an economically excessive number of fish would have to be
tagged to yield statistically adequate data. Noncthclcss, this was the first biological asscssmcnt and
evaluation of fishway use by warmwater species
rind some intriguing patterns emerged.
A complete assessment of fishway performance
should address entrance attraction efficiency, difficulty or physical output associated with upstream
passage, and finally. passage efficiency. Simple
observation ur mark-recapture cxperimcnts at existing fishways (c.g., Fcrnet 1984; Schwalmc et al.
1985; Monk et al. 1989; Dcxter and Lcdet 1996)
have not cffcctivcly provided information that relates overall attempts to successful attempts, the
timing of fishway use, and other factors necessary
for a clear understanding of tishway efficiency.
One recent study used two receiver check-points
or gate-keepers, spaced 50 m apart to monitor the
movements and ascent of barbel Barbus barbus
over a flow-gauging weir in England (Lucas and
Frear 1997). However, until now, no studies have
uscd radiotclcmctry to continuously monitor the
detailed behavior of warmwater fish in fishways.
Although the literature contains numerous quantitative and anecdotal reports of anadromous fish
passage, we are not aware of any study relating
the success and failures of individual attempts to
use fishways by warmwater fish.
The behavioral and physiological impacts of river barriers on white suckers and smallmouth bass
arc largely unknown. It appears that upstream
movcmcnts of both species are interrupted and delayed at dams despite the presence or absence of
fishways. In a recently published report, Kanehl el
al. (1997) demonstrated the positivc effects of lowhead dam removal on smallmouth bas6 abundance
and biomass. Numerous other species experience
significant delays passing river barriers. For examplo, dclays of several weeks have been reported
among northern pikc Esox luclus in Canada (Fernet
1984), barbel in England (Lucas and Frear 1997),
Atlantic salmon Salnio salar in Scotland (Webb
1990), and several percid spccies in Australia
(Harris and Mallcn-Cooper 1994). At the Mannhcim weir, dclays wcrc often indefinite, and most
radio-tnggcd fish did not pass upstream. Delays
among salmonid and cyprinid species have been
attributed to a relucti
(Priede and Holliday
to swim at speeds ner
(Lucas and Frear 195
numerous spccies of
tentially catastrophic
after extensive dclays
1971).
In our study, watel
wcrc usually less thar
ming speeds, which a
bass 378 mm TL anc
370 mm fork length (
ditions should have I
use. It is interesting
began using the low-1
season and at lower
high-velocity fishway
fish rely heavily on a
to support intense sw
1992). Cooler watcr
swimming capacity (1
that the onset of bur:
at lower water vcloci
low a species-spocific
achieving and mainta
essary to negotiate hi
lengths of some fish.
swimming performar
Denil fishways may
ative behavioral respc
air bubbles, and a hi1
the surface of the flo,
gested that fish must t
opposing flows to prr
velocities and turbulc
have prohibited succt
ticularly small small
suckers. It should be
the east fishway cxt
mended maximum.
The tracking methc
study permitted direc,
tial delays associatet
trances, time spent in
of passage. Both wh
bass rested much lon
pools in the west fis
dication of fatigue. 1
servations (sonior ac
smallmouth bass migl
with water velocities
0.5 rnls indicate that
fluenced by turbulent
801
USE OF TWO DENIL PISHWAYS
attributed to a reluctance to swim at high spceds
(Priede and Holliday 1980) or to physical inability
to swim at speeds necessary for successful asccnt
(Lucas and Frear 1997). Regardless of the cause,
numerous speciesof fish havc demonstrated a potentially catastrophic tendency to resorb gametes
after extensive delays below dams (Shikhshahekov
1971).
In our study, water velocities in both fishwayfi
were usually less than maximum prolonged swimming spceds, which were 1.18 rn/s for smallmouth
bass 378 mrn TL and 0.73 m/s for white suckers
370 mm fork length (Jones et al. 1974). Such conditions should havc permitted succcssful fishway
use. It is intcrcsting to note that whitc suckers
began using the low-velocity fishway earlier in the
season and at lowcr water temperatures than the
high-velocity fishway. At low water temperatures,
Ash rely heavily on anaerobic whitc muscle tissue
to support intense swimming activity (Rome et al.
1992). Coolcr water temperatures reduce aerobic
swimming capacity (Jayne and Lauder 1994) such
that the onset of burst activity occurs sooner and
at lower watcr velocities. Water ternpcra(ures helow a species-specific minimum prcvcnt fish from
achieving and maintaining swimming spceds ncccssary to negotiate high water velocities d o n g the
lcngihs of some fishways. Comparisons between
swimming performclnce and passage success in
Denil fishways may he funher obscured hy negative behavioral responses to turbulence, entrained
air bubbles, and a high-velocity lnyer of water at
the surface of the flow profile. Beach (1984) suggested that fish must swim at lcast 30% faster than
opposing flows to progress upstream. High watcr
velocities and turbulcnce in the east fishway may
have prohibited successlul use by some fish, particularly small smallmouth bass and large white
suckers. It should bc mentioned that the slope of
the cast fishway exceeds Clay's (199.5) recommcndcd maximum.
Thc trucking method incorporated in the present
study permitted direct measurements of thc potential delays associatcd with locating fishway entrances, time spent in resting pools, and total time
of pasgage. Both whitc suckers and smallmouth
bass rested much longcr in the second of thc two
pools in the west fishway. which may bc an indication of fatigue. Visual and videographic observations (senior author's unpublished data) of
smallmouth bass migrating through Denil fishwnys
with water velocities in excess of approximately
0.5 m/s indicate that fish positions are greatly influenced by turbulcnce. Nonbenthic species, such
as smallmouth bass, mufit withstand a great deal
of turbulcnce while negotiating maximum water
velocities near the surface of Denil fishway flows.
These maximum velocitics and the behavioral effects of turbulence on fish pnssagc uhould be addrcvsed to refine design criteria for Dcnil fishways.
Fish that were able to fit in the spaces between
baffles without injury sometimes passcd upstream
when high writer velocities should have prohibited
succcssful fishway use. This was accomplished by
burst swimming upstream from refuge to refuge,
as observed among cyprinids in other Denil fishways (Scbwalme et al. 1985). Fish that were able
to maintain position near the bottom of the flow
profile, such as benthic suckers, were able to usc
the fishways whcn surface water velocitics appeared exccssivc. The relative importancc of burst
swimming, position-holding abilities within thc
boundary layers of flows, prolongcd swimming
abilitics, and injury rates caused by impacts with
baffles also require further investigation lo maximize passage rates of warmwater species through
Denil fishways. Experiments using electromyogram tclemtlry may help to determine relative
physiological costs (McKinley and Power 1992;
Demers et al. 1996; Hinch et al. 1996) associated
with the use of different fishways under a variety
of conditions.
In summary, the relatively simplc and inexpensive fishway design on the east side of the Mannhcim weir permitted largc smallmouth bass as well
as small white suckers. a species with lesser swimming abilities, to paw upstream. Watcr velocities
in hoth fishways wcrc generally within thc range
of critical swimming spceds of smallmouth bass
and white suckers, and equal numbers of smallmouth bass were collected from cnch of the fishway traps. However, the lower-vclocity fishway
with resting pools, on the west side of the Mannhcim weir permitted morc white suckers to pass
upstream. The numbcr of white suckers and smallmouth bass that used each fishway declined exponentially as water velocities increased. Maximum velocities used by white suckcrs and smallmouth bass were 0.96 and 0.99 mls, respectively.
Passage efficiencies of each species at hoth fishways wcrc low to moderate. Futurc work should
address the physiological and behavioral characteristics of fish that affect passage through fishways.
Acknowledgmenb
We thank Oabrielle Held. Scan Clancy, and
Stcven Cooke for thcir assistance in the ficld;
802
BUNT ET AL.
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