<<

Journal of Great Lakes Research 36 (2010) 490–496

Contents lists available at ScienceDirect

Journal of Great Lakes Research

journal homepage: www.elsevier.com/locate/jglr

Spawning by (Sander vitreus) and white sucker ( commersoni)in the Detroit River: Implications for spawning habitat enhancement

Bruce A. Manny a,⁎, Gregory W. Kennedy a, James C. Boase b, Jeffrey D. Allen a, Edward F. Roseman a a US Geological Survey, Great Lakes Science Center, 1451 Green Rd, Ann Arbor, MI 48105, USA b U.S. Fish and Wildlife Service, Alpena National Fish and Wildlife Conservation Office, Waterford Fisheries Station, 7806 Gale Road, Waterford, MI 48327, USA article info abstract

Article history: Few active fish spawning grounds have been found in channels connecting the Great Lakes. Here, we describe one Received 5 August 2009 near Belle Isle in the Detroit River, part of the channel connecting lakes Huron and Erie. There, in 2005, we Accepted 9 May 2010 collected 1,573 fish eggs, cultured them, and identified the hatched larvae as walleye (Sander vitreus)andwhite sucker (Catostomus commersoni). Walleye spawning peaked during the week of April 12–19; white sucker Communicated by Geoffrey Steinhart spawning peaked on May 10. Average areal rate of egg deposition by walleye and white sucker at this spawning ground in 2005 was 346 and 25 eggs/m2, respectively. Our environmental measurements showed that bottom Index words: fi Walleye substrates on this spawning ground were largely sand, not optimal for sh reproduction. We hypothesize that White sucker reproduction of these fish at this spawning ground could be enhanced by adding rock and gravel substrates for Spawning ground protection of deposited fish eggs and suggest that reproduction by walleye in the Detroit River may add resilience Detroit River to production of walleye in western Lake Erie. Published by Elsevier B.V.

Introduction were collected in the river in 1977 (Goodyear et al., 1982; Hatcher and Nester, 1983). Historically, many fish species were reputed to in channels The walleye population in western Lake Erie is presently thought connecting the Laurentian Great Lakes, including the Detroit River, to be a mixture of fish that spawn in the Sandusky and Maumee rivers part of the channel connecting lakes Huron and Erie (Goodyear et al., (Mion et al., 1998) and on shoals in Lake Erie (Roseman et al., 1996). The 1982; Manny et al., 1988). Recent evidence for fish spawning in the Detroit River is not recognized as a walleye spawning area by current Detroit River is sparse, and their contribution to fishery productivity fishery management paradigms (Jones et al., 2003; MacLennan et al., in lakes Huron and Erie has been largely overlooked (Mackey and 2003; Strange and Stepien, 2007). Recent evidence of egg deposition by Goforth, 2005; Minns and Wichert, 2005). Of fish community goals fish in the Detroit River is limited to lake sturgeon, lake whitefish, and and objectives for all of the Great Lakes, only those for lakes Erie and walleye (Caswell et al., 2004; Roseman et al., 2007; Manny et al., 2007). St. Clair recognize the potential contribution of stream spawning These latter authors postulated that walleye eggs they collected in fishes to fishery productivity in those lakes (MacLennan et al., 2003; 2004 drifted from a spawning site upstream of Belle Isle. The purpose Ryan et al., 2003). of this study was to identify a potential source of walleye eggs collected In the early 1900s, the Detroit River was a reputed spawning ground by Manny et al. (2007) near Belle Isle, determine how suitable the for cisco (Coregonus artedii), lake sturgeon (Acipenser fulvescens), shoal at the head of Belle Isle was for spawning by fish, and put lake whitefish (Coregonus clupeaformis), and walleye (Sander vitreus; spawning by walleye in the Detroit River in perspective with the socially Goodyear et al., 1982). Walleye spawning was reputed at four areas in and economically valuable metapopulation of walleye in lakes Erie and the lower Detroit River, and the head of Belle Isle was reputed to be a St. Clair. nursery area for walleye larvae (Goodyear et al., 1982). White sucker were not reputed to spawn in the Detroit River, but white sucker larvae Materials and methods

Study area

⁎ Corresponding author. Our study area was a shoal at the head of Belle Isle beside the Fleming E-mail addresses: [email protected] (B.A. Manny), [email protected] (G.W. Kennedy), [email protected] (J.C. Boase), [email protected] (J.D. Allen), Channel in the Detroit River (Fig. 1), about 1 and 50 river km from lakes [email protected] (E.F. Roseman). St. Clair and Erie, respectively. Water depth on the spawning ground in

0380-1330/$ – see front matter. Published by Elsevier B.V. doi:10.1016/j.jglr.2010.05.008 B.A. Manny et al. / Journal of Great Lakes Research 36 (2010) 490–496 491

Fig. 1. Study area at the head of Belle Isle in the Detroit River showing nine locations where 45 egg mats were deployed in 2005.

our study area ranged from 2 to 6 m; water depth in the adjacent and reset at the same location. All eggs visible on the interior and channels exceeds 10 m. Width of the river near the study area is 1.6 km exterior mat surfaces were removed with forceps, placed in Detroit (Derecki, 1984). Mean annual discharge of the Detroit River near Belle River water in a sealed 100-ml glass jar, and transported on ice to the Isle is 5,210 m3/s (Edwards et al., 1989). Great Lakes Science Center in Ann Arbor, MI. Because the eggs had been exposed to many pathogens in the Detroit River that could potentially Egg sampling reduce egg survival, eggs were then cultured in McDonald hatching jars in temperature-controlled (6.3–12.5 °C), filtered (10-μmporosity),UV- During April–May 2005, we sampled fish eggs by setting mats on the sterilized (Emperor Aquatics, Model EU40, Pottstown, PA), river water river bottom following the methods of Nichols et al. (2003).Eachmat that was recirculated in an apparatus of our own design to remove or kill consisted of a rectangle (38×50×2.5 cm) of latex-coated, natural-fiber, potential pathogens (USGS Great Lakes Science Center, unpublished furnace-filter material (product no. 383-716-601; American Air Filter procedure). Fish larvae hatched from the eggs were identified following International, Louisville, KY) wrapped around a metal frame Auer (1982). The density of eggs on each gang of egg mats was derived (38×24×0.5 cm) and secured with four 5×2.5 cm steel-spring binder by summing the number of eggs removed from each of the five mats clips (final mat dimensions: 38×25×4.5 cm). Mats were designed to be each week and dividing by the combined upper surface area of all five thin and flat, allowing close contact with the river bottom. Because only mats (0.46 m2/gang). Walleye eggs were visually distinguished from one side of each mat was exposed to the water column, we assumed that larger sucker eggs and smaller eggs of other species on the basis of egg eggs collected by the mats settled out of the water column onto the diameter. The range in diameter of walleye, sucker, and other eggs was upper surface area of the mat. We calculated the surface area of each mat 2.0–2.1 mm, 3.0–3.1 mm, and 1.4–1.5 mm, respectively. Weekly catch as a 2-dimensional surface area representing only one side of the mat of walleye, white sucker, and other eggs in 2005 was derived by (0.092 m2). Five such mats linked together with braided nylon line summing all of the walleye, sucker, and other eggs, respectively, that we formed a gang of egg mats. Each gang was anchored with a small collected each week on all egg mats combined. Cumulative density of (3.6 kg) Danforth anchor on each end. The downstream anchor was walleye and sucker eggs deposited per sampling location and at all connected by line to a surface buoy for gear retrieval (cf., Manny et al., sampling locations combined during our study was derived by adding 2007). We deployed mats in waters 2–6 m deep, based on water depths the walleye and sucker eggs collected weekly per square meter on each where walleye were reported to spawn in large streams (Geiling et al., gang during the study, respectively, and calculating an overall egg 1996; Corbett and Powles, 1986; Pitlo, 1989). Sampling locations were density for walleye and sucker eggs, respectively, on the spawning area, equally spaced across the study area to sample three locations each on as a mean of those cumulative egg densities per gang. The areal extent of the 2-, 4-, and 6-m contours. Beyond the 6-m contour, water depth the spawning ground at the head of Belle Isle was defined by creating a increased rapidly on either side of the shoal to 11 m in the shipping polygon of the area surrounded by the 6-m depth contour, using channels, where passing vessels made it unsafe to sample. On April 5, geographic information system software (ArcGIS v.9.x, 2008, ESRI, 2005, a week after ice-out, nine gangs (45 mats total) were deployed on Redlands, CA). The area of the spawning ground, calculated with the river bottom in the study area (Fig. 1). Weekly during each of the geographic information system software was 202,378 m2 or 20 ha. Total next eight weeks, each gang of egg mats was lifted, examined for eggs, eggs deposited by walleye on the spawning ground was calculated as 492 B.A. Manny et al. / Journal of Great Lakes Research 36 (2010) 490–496 the product of the spawning ground area (202,378 m2)andthemean tified) eggs were collected during this study. Walleye eggs were cumulative density of walleye eggs deposited over the spawning ground collected at all nine sampling locations from April 12 to May 17 and for in 2005 (346 walleye eggs/m2). six consecutive weeks at sampling locations 2, 4, 5, and 7 (Figs. 2 and 3). Cumulative, mean, areal rate of egg deposition by walleye at each Environmental measurements sampling location was 346 walleye eggs/m2 (range: 27–747 eggs/m2). White sucker eggs were collected at 7 of the 9 sampling locations and To understand environmental conditions on the spawning ground, for five consecutive weeks from April 26 to May 24 at only locations we characterized physical aspects of our study area in May 2005 and 8 and 9 (Figs. 2 and 4). Cumulative, mean, areal rate of egg deposition July 2007. On May 9, 2005, at each of the nine sampling locations, by white sucker at each sampling location was 25 eggs/m2 (range: 0– water depth and geospatial coordinates were determined with a boat- 127 eggs/m2). No white sucker eggs were collected at sampling mounted, combination global positioning system and depth sounder locations 3 and 6 in shallow, slow-moving water near the island. (Garmin International, Olathe, KS). Water temperature was measured at Greatest egg deposition by walleye and white suckers was concen- 0.5-m water depth with a mercury stem thermometer (Model 14-983- trated along the southern (Fleming Channel) margin of the spawning 10B; Fisher Scientific, Chicago, IL). To characterize the composition and ground (Figs. 3 and 4). arrangement of river bottom substrates and assess interstitial void space Walleye spawning peaked during the week of April 12–19, as among the substrates at the sampling locations, bottom substrates were water temperature increased from 6.5 to 9.0 °C (Fig. 2). That week, examined and photographed in 2005 under ambient light conditions 958 walleye eggs were collected on 42 of the 45 mats deployed. On with a high-resolution underwater video camera (Multi-Sea Cam 1050; April 19, the deposition of walleye eggs was highest at locations DeepSea Power and Light, San Diego, CA) attached to a 14-kg depressor 8 (563/m2), 6 (387/m2), and 7 (290/m2) in deeper (3.2–6.0 m) water plate, as the boat drifted over each sampling location. The video along the southern edge of the study area (Fig. 3). That same week, camera was not used to judge or estimate the particle size of bottom the next highest density of walleye eggs was found at locations 2 substrates. During video transects, water depth was measured with a (221/m2), 5 (219/m2), and 1 (161/m2) in deeper water (4.9–6.0 m) boat-mounted depth sounder and boat movements were recorded in along the northern edge of the study area. Also that week, lowest egg time and space using global positioning system software (Garmin densities were found at locations in shallow waters (1.9–2.9 m deep) International, Olathe, KS) following Edsall et al. (1997). On July 20, 2007, closer to the island at locations 4 (120/m2), 3 (92/m2), and 9 (4/m2) to describe water depth, water velocity at the water–substrate interface, (Fig. 3). Thereafter, walleye egg deposition per week decreased at all and bottom substrates at each sampling location where egg mats were locations, as water temperature decreased to 5–6 °C between April 26 deployed in 2005, three measurements of water depth were made with and May 3 and then increased to 9–11 °C between May 10 and 17 a boat-mounted depth sounder and averaged; three measurements of (Fig. 2). Averaged over the sampling period, cumulative walleye egg water velocity were made at the water–substrate interface with an deposition rate was greatest at location 8 (747/m2), next greatest at electronic current meter (Model 201; Marsh McBirney, Fredrick, MD) locations 6 and 7 (474–479/m2), intermediate at locations 1–5 (129– and averaged; and one sample of bottom substrates was collected with a 366/m2), and least at location 9 (27/m2). All differences in egg standard PONAR grab (Model 1725-F10; Wildlife Supply Co., Buffalo, deposition rates herein are numerical differences but not statistically NY), placed in a pan and examined to assess particle size of the significant differences. substrates by reference to Wentworth (1922).Wedidnotmeasure White sucker spawning peaked on May 10, 3 weeks after peak dissolved oxygen concentration because dissolved oxygen concentra- walleye spawning (Fig. 2). That week, a total of 43 white sucker eggs tions throughout the Detroit River are high enough to sustain fish all the were collected. White sucker eggs were collected for five consecutive time (6.5–10.5 mg/L; Davis, 1979; Manny et al., 1988; McClain and weeks from April 26 to May 24 at locations 8 and 9 as water temperature Manny, 2000). We also did not measure sedimentation rate in the study increased from 6 to 12 °C (Fig 2). On May 10, total mean deposition of area. white sucker eggs was greatest at location 9 (54 eggs/m2). That week, To check if there was a significant difference in physical conditions lesser rates of white sucker egg deposition were found at locations 7 and at our study site between the two sampling periods, we evaluated 8(11eggs/m2) along the southern edge of the study area. Averaged over whether there was any difference in water levels in our study area the sampling period, cumulative white sucker egg deposition rate between May 2005, when we collected fish eggs, and July 2007, when wasgreatestatlocation9(127eggs/m2), next greatest at location 8 (43 we measured physical environmental properties at the spawning eggs/m2), and lower at locations 1, 2, 4, 5, and 7 (range: 4–19 eggs/m2). ground. We compared historic, published water levels measured at No white sucker eggs were collected at locations 3 and 6 in shallow those times at the Belle Isle gauging station about 1200-m down- waters (1.9–2.9 m deep) closer to the island (Fig. 4). stream from our study area (cf., www.lre.usace.army.mil/greatlakes/ hh/greatlakeswaterlevels/currentconditions/connectingchannels- waterlevels/index.cfm). Mean monthly water levels at the Belle Isle gaging station in May 2005 and July 2007 were 175.01 m and 174.82 m, International Great Lakes Datum 1985, respectively. Likewise, the within-month variation in water level at the Belle Isle gaging station in May 2005 and July 2007 was 0.20 m and 0.26 m, respectively, and the difference between mean water level in May 2005 and July 2007 was 0.19 m. Hence, the difference in mean water level between May 2005 and July 2007 at Belle Isle was less than the within-month variation there in water level in either May 2005 or July 2007 and of no environ- mental consequence between our two survey sampling periods.

Results

Egg deposition

Fig. 2. Total eggs of walleye and white sucker collected per week and water temperature at fi Egg deposition by sh was widespread in our study area. A total of Belle Isle in the Detroit River during April–May 2005. The total of eight other eggs collected 1455 walleye eggs, 110 white sucker eggs, and eight other (uniden- on May 3 and 10 was too few to depict here at this scale. B.A. Manny et al. / Journal of Great Lakes Research 36 (2010) 490–496 493

Fig. 3. Map of cumulative walleye egg deposition (eggs/m2) on egg mats at nine locations on a shoal at the head of Belle Isle in the Detroit River in 2005.

A total of eight other eggs were collected at locations 5 and 6 on substrates on the shoal. Fine silts were found at locations 3 and 6 near May 3 and 10, as water temperature increased from 6 to 12 °C (Fig. 2). the island where we found the least water depth, lowest water These eggs were smaller than walleye eggs but were not taxonom- velocity, and lowest walleye egg deposition rates. Location 7 along the ically identified because they died in culture. From the total of 1455 southern margin of the shoal was the only place where rocks and walleye eggs cultured, 77 larvae hatched and were identified as walleye. gravel were found (Table 1). In July 2007, at locations 3, 6, and 9, the From the total of 110 white sucker eggs cultured, 18 larvae hatched and river bottom was covered with several centimeters of fine silts and were identified as white sucker. colonized by shoots of submersed macrophytic and patches of filamentous algae. Environment measurements Discussion During the 7-week sampling period for walleye and white sucker, water temperature increased (Fig. 2). Walleye spawned during a Our discovery of viable, fertilized walleye and white sucker eggs 3-week increase in water temperature from 3.5 to 9 °C. Suckers spawned deposited over a wide area on the river bottom at the head of Belle Isle during a 4-week increase in water temperature from 6 to 10 °C between indicates that walleye and white sucker spawned in the Detroit River April 26 and May 17 (Fig. 2). Mean water depth varied from 1.9 m near at this location in April 2005. This evidence supports our hypothesis the island at location 3 to 6.0 m at locations 2, 5, and 8 near the upstream that an upstream spawning ground somewhere in the Detroit River, channel margins of the shoal area (Table 1). Mean water velocity at possibly this spawning ground, was the source of 136 viable, fertilized the water substrate interface varied from 3 cm/s near the island at walleye eggs collected approximately 600 m downstream of this locations 3 and 6 to 34 cm/s at locations 2 and 8 near the channel margins spawning ground in April and May 2004 by Manny et al. (2007). Peak of the shoal area (Table 1). From April 5 to June 2, 2005, water clarity spawning by walleye in this study corresponded closely in time and (Secchi disc depth) was high (mean: 1.91 m) and increased steadily from water temperature with peak spawning by walleye 600 m down- aminimumof0.3monApril13ontoamaximumof3.6monJune1. stream on April 15–22, 2004, at a water temperature of 5.6–9.7 °C Images of eleven underwater video camera transects (duration: (Manny et al., 2007) and peak spawning by walleye on shoals in 165 minutes) recorded over the shoal on May 9, 2005, showed that western Lake Erie during April 10–20, 1994, at water temperatures of sand was the predominate bottom type, interspersed with large, thin 5–7°C(Roseman et al., 1996). patches of vacant dreissenid shells, within our study area. Data from To our knowledge, this report is the first scientific evidence of PONAR grabs at each egg mat location on July 20, 2007, showed that spawning by white sucker in the Detroit River. Deposition of walleye bottom substrates varied from coarse sand in areas of higher water eggs, then white sucker eggs at this spawning ground in the Detroit velocity to silt and clay near the island in areas of low water velocity River closely resembled that described for these two species in an (Table 1). Interstitial (void) space in the bottom substrates within Ontario stream (Corbett and Powles, 1986) and elsewhere (Scott and our study area was low, owing to a general lack of rock and gravel Crossman, 1973; Johnston, 1997). 494 B.A. Manny et al. / Journal of Great Lakes Research 36 (2010) 490–496

Fig. 4. Map of cumulative white sucker egg deposition (eggs/m2) on egg at nine locations on a shoal at the head of Belle Isle in the Detroit River in 2005.

Walleye and white sucker deposit higher densities of eggs on may be the relatively low year class strength of 2005 walleye (10.3) gravel and cobble than on sand (Balon, 1975; Jones et al., 2003). The within the western Lake Erie summer bottom trawl index values maximum, cumulative density of walleye eggs deposited during the for age-0 walleye, 2001–2007 (range: 0.1–155.5; Ohio Division of spawning period in our study area (747 eggs/m2) is low compared Wildlife, 2008). with the range of walleye eggs found elsewhere on sand in rivers and Our estimated size of the walleye spawning ground at the head of lakes (1001–1302 eggs/m2; Jones et al., 2003) or on rubble, cobble, Belle Ile (20 ha) falls near the low end of the size range of fluvial and gravel substrates (2537–7050 eggs/m2) by other investigators walleye spawning grounds reported in the literature (6–240 ha; (Jones et al., 2003; Johnson, 1961; Corbett and Powles, 1986). Likewise, Cheng et al., 2006; MacDougall et al., 2007). Extrapolating from our the maximum, cumulative egg deposition by white sucker that we average cumulative density of 346 walleye eggs/m2 measured in measured at this Belle Isle spawning ground (127 eggs/m2; Fig. 4)was 2005 within the 6-m contour area that we sampled on the Belle Isle in the low end of the range of white sucker egg density measured with a spawning ground (202,378 m2), we estimate that 70,022,788 total Surber sampler in an Ontario stream (0.5–710 eggs/m2; Corbett eggs were deposited on this 20-ha spawning ground in 2005. If we and Powles, 1986). An explanation for the relatively low density of assume that the average fecundity of a walleye spawning on this deposited walleye eggs on the Belle Isle spawning ground in our study ground at Belle Isle was equal to that of walleye from western Lake Erie in 1990-91 (228,875 eggs/female; Muth and Ickes, 1993), we estimate that 306 females spawned at Belle Isle in 2005. Using an Table 1 estimated spawning substrate requirement of 20 m2/female (Furlong Water depth (m) and velocity (cm/s) at the water substrate interface, and river bottom et al., 2006 in MacDougall et al., 2007), the Belle Isle spawning ground substrates at nine locations on the spawning ground at the head of Belle Isle on July 20, 2007. could theoretically accommodate 10,119 females, if optimal spawning Location Depth (m) Velocity (cm/s) River bottom substrates substrates were available. Because walleye spawn on shoals in Lake 1 4.9 20 Coarse sand and shells Erie at water depths of 10 m (Roseman et al., 1996), and by design we 2 6.0 34 Fine sand restricted our egg sampling to water depths of 2–6 m, we may 3 1.9 3 Sand and silt have missed egg deposition by walleye around this shoal at depths 4 2.9 6 Clay covered by sand greater than 6 m and underestimated the size of this spawning 5 6.0 27 Coarse sand and shells 6 3.2 3 Silt and clay ground at Belle Isle. 7 4.4 23 Rocks, gravel, and shells The preference of both walleye and white sucker to spawn on rock 8 6.0 34 Clay covered by coarse sand substrates may be illustrated by greatest egg deposition by both 9 2.2 9 Silt, sand, and shells species along the southern margin of this spawning ground, where we Bottom substrates were collected with a PONAR grab. found the only rocks and gravel (Table 1). Millions of tons of rock and B.A. Manny et al. / Journal of Great Lakes Research 36 (2010) 490–496 495 gravel have been excavated from the river bottom to create 96 km of spawning stocks of walleye contribute to the mixed-stock walleye deep-draft navigation channels in the 51-km Detroit River (Larson, population in eastern Lake Erie, and walleye that spawned in the Black 1981). Because much of the rock and gravel once present in this river Sturgeon River were genetically related to walleye that were was removed, walleye and white sucker may spawn on less than historically harvested in Black Bay, (Wilson et al., 2007). optimal substrates where other environmental conditions are suitable Reproduction and larval recruitment of walleye are largely a function for reproduction, such as this sandy shoal at the head of Belle Isle. of spawner abundance (Cheng et al., 2006) but can be enhanced by Water depth at the head of Belle Isle resembled that on productive creation of new spawning habitat (Colby et al., 1994; Geiling et al., walleye spawning grounds described by others (Johnson, 1961; Corbett 1996). Once established as a spawning stock, walleye home to fluvial and Powles, 1986; Geiling et al., 1996; Paragamian, 1989; Pitlo, 1989; spawning grounds (Crowe, 1962; Jennings et al., 1996). Protection Jones et al., 2003). Sand, the predominant bottom substrate type on and rehabilitation of fish spawning grounds in connecting channels of this Belle Isle spawning ground, was characterized by Jones et al. (2003) the Great Lakes could be desirable because management of migratory as average habitat quality for spawning by walleye, on which 23% fish species in the Great Lakes can be complicated when adults are of deposited eggs could be expected to hatch. Higher rates of egg harvested in one jurisdiction but reproduce in another where degra- deposition, hatch, and survival of both walleye and white sucker dation of their spawning habitat may be unregulated (Welsh, 2004). might be expected on this shoal at Belle Isle, if spawning habitat there Rehabilitation of fish spawning habitat in the Detroit River has been were enhanced. Adding rock rubble, cobble, or gravel that are preferred proposed as a means of delisting loss of fish and wildlife habitat as an by spawning walleye (Jones et al., 2003)andpossessmoreinterstitial impaired beneficial water use in that river (Manny, 2003). A draft fish- space for protection of fish eggs and larvae from predation and dis- community goal and objective of maintaining stable, self-sustaining lodgement may increase survival of walleye eggs at this Belle Isle stocks of fish in the Detroit River Area of Concern, lists walleye as the spawning ground. first management priority (Hartig, 1993). Protection and restoration Our study area is a historic, reputed nursery area of walleye larvae of self-sustaining stream-spawning stocks of walleye is also a fish (Goodyear et al., 1982). The area may be suitable for survival of both community objective for Lake Erie (Ryan et al., 2003). Therefore, walleye and white sucker larvae because high dissolved oxygen this shoal at the head of Belle Isle would be a good site to explore concentrations needed by walleye larvae (McElman and Balon, 1979) rehabilitation of spawning habitat for walleye. are present in the water year round (Manny et al., 1988; McClain and – Manny, 2000). From April to June when water temperature is low (3 Acknowledgments 10 °C) and fish larvae are present in our study area, dissolved oxygen concentrations would be near saturation (Dobson, 1967). During our We thank D. Wilcox, M. Bur, and R. Drouin for manuscript review; survey of the shoal on July 20, 2007, beds of aquatic plants were seen D. Bennion, M.G. Black, J. Craig, R. Quintal, S. Riley, and E. Stockwell for beneath the water surface, and PONAR samples at several of the egg field assistance and/or data analysis; and the US Coast Guard, Belle mat locations (locations 1, 3, 6, and 7) contained roots and leaves of Isle Station for technical support. Financial support was provided, in submersed aquatic plants (USGS Great lakes Science Center, unpub- part, by grant no. 02CR-1.19 from the (NOAA) Coastal Restoration fi lished data) that could provide shelter for young sh. Grant Program and grant no. 2002.232 from the Great Lakes Fishery Survival of walleye larvae produced on spawning habitat at Belle Trust to the Michigan Sea Grant College Program at the University of Isle may, in part, supplement recruitment of the walleye metapopula- Michigan for the Belle Isle/Detroit River Sturgeon Habitat Restoration, tion in Lake Erie (Manny et al., 2007), provided that transport of Monitoring, and Education Project, and by the USGS Science Support walleye larvae is rapid to zooplankton-rich, nursery habitat down- Program for Projects 03-R3-02 and 06-R3-04. Reference to trade stream (Jones et al., 2003; Zhao et al., 2009). During out-migration names does not imply endorsement by the US Government. This article from Ohio rivers, survival of walleye larvae was reduced by high river is contribution no. 1590 of the Great Lakes Science Center. discharge, suspended sediment concentration, and water velocity (Mion et al., 1998; Jones et al., 2003). In May 2005, walleye larvae References produced at Belle Isle would have been transported by river currents 50 km down the Detroit River to Lake Erie. Then, the average water Auer, N.A., 1982. Identification of larval fishes of the Great Lakes basin, with emphasis velocity (0.3–0.7 m/s; Derecki, 1984; Manny et al., 1988) and water on the Lake Michigan Drainage. Great Lakes Fishery Commission, Ann Arbor, MI: temperature (6–10 °C; Fig. 2) in the Detroit River would have Spec. Publ., vol. 82-3. Balon, E.K., 1975. Reproductive guilds of fishes: a proposal and definition. J. Fish. Res. theoretically predicted a moderate- to low-transport survival rate of Board Can. 32, 821–864. walleye larvae (0.05–0.1; Jones et al., 2003; Fig. 8). In 2006, from mid- Caswell, N.M., Peterson, D.L., Manny, B.A., Kennedy, G.W., 2004. Spawning by lake March through mid-June, 29 walleye larvae were collected at sturgeon (Acipenser fulvescens) in the Detroit River. J. Appl. Ichthyol. 20, 1–6. Cheng, F., Zika, U., Banachowski, K., Gillenwater, D., Granata, T., 2006. Modelling the effects midchannel sampling stations in the Detroit River and over 90% of dam removal on migratory walleye (Sander vitreus) early life-history stages. River of these larvae were collected from sites in the lower half of the river Res. Appl. 22 (8), 837–851. (E.F. Roseman, USGS Great Lakes Science Center, unpublished data). Colby, P.J., Lewis, C.A., Eshenroder, R.L., Haas, R.C., Hushak, L.J., 1994. Walleye- rehabilitation guidelines for the Great Lakes area. Great Lakes Fishery Commission, Fluvial spawning grounds in the Detroit River may be more Ann Arbor, MI. 112 pp. protected than shoals in western Lake Erie from wind-generated Corbett, B.W., Powles, P.M., 1986. Spawning and larval drift of sympatric and water currents that dislodge walleye eggs (Roseman et al., 2001). white suckers in an Ontario stream. Trans. Am. Fish. Soc. 115, 41–46. fl Crowe, W., 1962. Homing behavior of walleyes. Trans. Am. Fish. Soc. 91, 350–354. Such uvial spawning grounds may add resilience to year-class Davis, J.C., 1979. Dissolved oxygen. In: Thurston, R.V., Russo, R.C., Fetterolf Jr., C.M., Edsall, T.A., strength of walleye in Lake Erie, when reproduction by shoal- Barber Jr., Y.M. (Eds.), A review of the EPA red book: quality criteria for water: Water spawning walleye in Lake Erie is reduced by strong water currents Quality Section, American Fisheries Society, Bethesda, MD, pp. 169–174. (Roseman et al., 2001, 2005; Zhao et al., 2009). Fluvial spawning Derecki, J.A., 1984. Detroit River, physical and hydraulic characteristics. Natl. Oceanogr. Atmos. Adm., Great Lakes Environ. Res. Lab., GLERL Contrib. No. 417. Ann Arbor, MI. 11 pp. grounds in the Detroit River may also be less susceptible than Dobson, H.H., 1967. Principle ions and dissolved oxygen in Lake Ontario. Proc. 10th Conf. spawning grounds on shoals in Lake Erie to climate-induced increases Great Lakes Res., Internat. Assoc. Great Lakes Res., Ann Arbor, MI, pp. 337–356. fi in water temperature that could theoretically reduce use of spawning Doka, S., Bakelaar, C., Bouvier, L., 2006. Chapter 6. Coastal wetland sh community fi assessment of climate change in the lower Great Lakes. In: Mortesh, L., Ingram, J., and nursery habitats by sh in the Great Lakes (Eshenroder, 2003; Hebb, A., Doka, S. (Eds.), Great Lakes coastal wetland communities: vulnerability to Jones et al., 2006; Doka et al., 2006). Walleye that spawn in tributaries climate change and response to adaptation strategies. Environment Canada and the smaller than this connecting channel contribute to mixed-stock Department of Fisheries and Oceans, Toronto, Ontario, pp. 101–128. Edsall, T.A., Behrendt, T., Cholwek, G., Frey, J., Kennedy, G., Smith, S., 1997. Use of walleye populations elsewhere in the Great Lakes. For example, remote-sensing techniques to survey the physical habitat of large rivers. U.S. Geol. Gatt et al. (2003) and MacDougall et al. (2007) showed that fluvial Survey, Great Lakes Sci. Cent., Ann Arbor, MI, Contrib. No. 983. 20 pp. 496 B.A. Manny et al. / Journal of Great Lakes Research 36 (2010) 490–496

Edwards, C.J., Hudson, P.L., Duffy, W.G., Nepszy, S.J., McNabb, C.D., Haas, R.C., Liston, C.R., Manny, B.A., Kennedy, G.W., Allen, J.D., French, J.R.P.I.I.I., 2007. First evidence of egg Manny, B.A., Busch, W.D.N., 1989. Hydrological, morphometrical, and biological deposition by walleye (Sander vitreus) in the Detroit River. J. Great Lakes Res. 33 characteristics of the connecting rivers of the international Great Lakes: a review. (2), 512–516. In: Dodge, D.P. (Ed.), Proceedings of the International Large Rivers Symposium: McClain, J., Manny, B.A., 2000. [online] Evaluation of lake sturgeon habitat in the Detroit Can. Spec. Publ. Fish. Aquat. Sci., vol. 106, pp. 240–264. River. Final report to the U.S. Environ. Prot. Agency for Award #GL98105. Great Lakes Eshenroder, R.L., 2003. Persistence of upper Great Lakes walleye (Sander vitreus) Natl. Prog. Off., Region 5, 77 West Jackson Blvd., Chicago, IL. 14pp. + appendices. populations in relation to river flow. In: Barry, Terence P., Malison, Jeffrey A. (Eds.), Available from http://www.fws.gov/midwest/alpena/documents/rpt-detrstn- Proceedings of PERCIS III, the Third International Percid Fish Symposium, hab99.pdf. Accessed Feb 9, 2010. University of Wisconsin, Madison, Wisconsin, U.S.A., July 20-24, pp. 69–70. McElman, J.F., Balon, E.K., 1979. Early ontogeny of walleye, Stizostedion vitreum, with Furlong, P., Foster, R.F., Colby, P.J., Friday, M., 2006. Black Sturgeon River dam: a barrier steps of saltatory development. Environ. Biol. Fish. 4, 309–348. to the rehabilitation of Black Bay walleye. Technical Report 06-03, Ontario Ministry Minns, C.K., Wichert, G.A., 2005. A framework for defining fish habitat domains in Lake of Natural Resources, Upper Great Lakes Management Unit-Lake Superior. Ontario and its drainage. J. Great Lakes Res. 31 (Suppl. 1), 6–27. Gatt, M.H., McParland, T., Halyk, L.C., Ferguson, M.M., 2003. Mitochondrial DNA Mion, J.B., Stein, R.A., Marschall, E.A., 1998. River discharge drives survival of larval variation and mixed-stock analysis of recreational and commercial walleye walleye. Ecol. Appl. 8, 88–103. fisheries in eastern Lake Erie. N. Am. J. Fish. Manage. 23, 431–440. Muth, K.M., Ickes, B.S., 1993. Fecundity of walleyes in western Lake Erie, 1966 and Geiling, W.D., Kelso, J.R.M., Iwachewski, E., 1996. Benefits from incremental additions 1990–91. J. Great Lakes Res. 19 (4), 715–719. to walleye spawning habitat in the Current River, with reference to habitat Nichols, S.J., Kennedy, G., Crawford, E., Allen, J., French III, J., Black, G., Blouin, M., Hickey, modification as a walleye management tool in Ontario. Can. J. Fish. Aquat. Sci. 53 J., Chernyak, S., Haas, R., Thomas, M., 2003. Assessment of lake sturgeon (Acipenser (Suppl. 1), 79–87. fulvescens) spawning efforts in the lower St. Clair River. J. Great Lakes Res. 29 (3), Goodyear, C.D., Edsall, T.A., Demsey, D.M., Moss, G.D., Polanski, P.E., 1982. Atlas of the 383–391. spawning and nursery areas of Great Lakes fishes. U.S. Fish and Wildlife Service, Ohio Division of Wildlife, 2008. Ohio's Lake Erie Fisheries, 2007. Annual status report. FWS/OBS-82/52, Vol. I-XIV. Federal Aid in Fish Restoration Project F-69-P. Ohio Department of Natural Hartig, J.H., 1993. A survey of fish-community and habitat goals/objectives/targets and Resources, Division of Wildlife, Lake Erie Fisheries Units, Fairport and Sandusky. status in Great Lakes areas of concern. Great Lakes Fishery Commission, Ann Arbor, 110 pp. MI. 95 pp. Paragamian, V.L., 1989. Seasonal habitat use by walleye in a warmwater river system, as Hatcher, C.O., Nester, R.T., 1983. Distribution and abundance of fish larvae in the St. Clair determined by radiotelemetry. N. Am. J. Fish. Manage. 9 (4), 392–401. and Detroit Rivers. U.S. Fish and Wildlife Service, Great Lakes Fishery Laboratory, Pitlo Jr., J., 1989. Walleye spawning habitat in Pool 13 of the upper Mississippi River. Ann Arbor, MI: Admin. Rpt., vol. 83-5. 41 pp. N. Am. J. Fish. Manage. 9 (3), 303–308. Jennings, M., Claussen, J., David, P., 1996. Evidence for heritable preferences for Roseman, E.F., Taylor, W.W., Hayes, D.B., Haas, R.C., Knight, R.L., Paxton, K.O., 1996. spawning habitat between two walleye populations. Trans. Am. Fish. Soc. 125, Walleye egg deposition and survival on reefs in western Lake Erie. Proceedings 978–982. of the Second International Percid Fishes Symposium: Annales Zoologici Fennici., Johnson, F.H., 1961. Walleye egg survival during incubation on several types of bottom vol. 33, pp. 341–351. in Lake Winnibigoshish, Minnesota, and connecting waters. Trans. Am. Fish. Soc. 90, Roseman, E.F., Taylor, W.W., Hayes, D.B., Knight, R.L., Haas, R.C., 2001. Removal of 312–322. walleye eggs from reefs in Western Lake Erie by a catastrophic storm. Trans. Amer. Johnston, T.A., 1997. Within-population variability in egg characteristics of walleye Fish. Soc. 130, 341–346. (Stizostedion vitreum)andwhitesucker(Catostomus commersoni). Can. J. Fish. Aquat. Roseman, E.F., Taylor, W.W., Hayes, D.B., Tyson, J.T., Haas, R.C., 2005. Spatial patterns Sci. 54, 1006–1014. emphasize the importance of coastal zones as nursery areas for larval walleye in Jones, M.L., Netto, J.K., Stockwell, J.D., Mion, J.B., 2003. Does the value of newly western Lake Erie. J. Great Lakes Res. 31 (Suppl. 1), 28–44. accessible spawning habitat for walleye (Stizostedion vitreum) depend on its Roseman, E.F., Kennedy, G.W., Boase, J.C., Manny, B.A., Todd, T.N., Stott, W., 2007. location relative to nursery habitats? Can. J. Fish. Aquat. Sci. 60, 1527–1538. Evidence of lake whitefish spawning in the Detroit River: implications for habitat Jones, M., Shuter, B.J., Zhao, Y., Stockwell, J.D., 2006. Forcasting effects of climate change and population recovery. J. Great Lakes Res. 33, 397–406. on Great Lakes fisheries: models that link habitat supply to population dynamics Ryan, P.A., Knight, R., MacGregor, R., Towns, G., Hoopes, R., Culligan, W., 2003. Fish- can help. Can. J. Fish. Aquat. Sci. 63, 457–468. community goals and objectives for Lake Erie. online Great Lakes Fishery Commission, Larson, J.W., 1981. Essayons. A history of the Detroit District U.S. Army Corps of Ann Arbor, MI: Spec. Publication, vol. 03-02. Available at: http://www.glfc.org/pubs/ Engineers. U.S. Government Printing Office, 1981-0-750-976. 215 pp. SpecialPubs/Sp03_2.pdf. Accessed Feb 9, 2010. MacDougall, T.M., Wilson, C.C., Richardson, L.M., Lavender, M., Ryan, P.A., 2007. Walleye in Scott, W.B., Crossman, E.J., 1973. Freshwater fishes of Canada. Bull. Fish. Res. Board the Grand River, Ontario: an overview of rehabilitation efforts, their effectiveness, and Canada 184 966 pp. implications for eastern Lake Erie fisheries. J. Great Lakes Res. 33 (Suppl. 1), 103–117. Strange, R.M., Stepien, C.A., 2007. Genetic divergence and connectivity among river and MacLennan, D.S. and Haas, R.C., Towns, G., Thomas, M.V., Roseman, E.F., Francis, J., reef spawning groups of walleye (Sander vitreus vitreus) in Lake Erie. Can. J. Fish. Braunscheidel, J., Halyk, L., Hector, D., Locke, B., McGregor, R., Morencie, M., Murray, A., Aquat. Sci. 64, 437–448. 2003. [online] Draft fish-community goal and objectives for Lake St. Clair, St. Clair Welsh, A.B., 2004. Factors influencing the effectiveness of local versus national River, and Detroit River (St. Clair System). Great Lakes Fishery Commission, Ann Arbor, protection of migratory species: a case study of lake sturgeon in the Great Lakes, MI. Available at http://www.glfc.org/lakecom/lec/LEC_docs/other_docs/scsfcgo. North America. Environ. Sci. Pol. 7, 315–328. pdf. [Accessed Feb 9, 2010] Wentworth, C.K., 1922. A scale of grade and class terms for clastic sediments. J. Ecol. 30, Mackey, S.D., Goforth, R.R., 2005. Great Lakes nearshore habitat science. J. Great Lakes 377–392. Res. 31 (Suppl. 1), 1–5. Wilson, C.C., Lavender, M., Black, J., 2007. Genetic assessment of walleye (Sander Manny, B.A., 2003. Setting priorities for conserving and rehabilitating Detroit River vitreus) restoration efforts and options in Nipigon Bay and Black Bay, Lake Superior. habitats. In: Hartig, J.H. (Ed.), Honoring our Detroit River, caring for our home. J. Great Lakes Res. 33 (Suppl. 1), 133–144. Cranbrook Institute of Science, Bloomfield Hills, MI, pp. 121–140. Zhao, Y., Jones, M.L., Shuter, B.J., Roseman, E.F., 2009. A biophysical model of Lake Erie Manny, B.A., Edsall, T.A., Jaworski, E., 1988. The Detroit River, Michigan: an ecological walleye (Sander vitreus) explains interannual variations in recruitment. Can. J. Fish. profile. U.S. Fish and Wildlife Service. Biol. Rep. 85 (7.17). Aquat. Sci. 66, 114–125.