Comparison of survival of winter-run steelhead based on stocking location 1 2 3 Jory L. Jonas1, James L. Dexter Jr.2, Neil Ledet3, Richard O’Neal4, Martha 4 Wolgamood5, and Jan VanAmberg6 5 6 7 1 Michigan Department of Natural Resources, Charlevoix Fisheries Research Station, 96 Grant Street, Charlevoix, Michigan 49720, USA. 8 9 2 10 Michigan Department of Natural Resources, Plainwell Operations Service 11 Center, 12 621 N. 10th Street, Plainwell, MI 49080, USA. 13 14 3 Indiana Department of Natural Resources, Fawn River State Fish Hatchery, 6889 N. State Road 327, Orland, IN 46776 USA. 15 16 4 17 Michigan Department of Natural Resources, Muskegon State Game Area, 7550 E. Messenger Road, Twin Lake, MI 49457, USA. 18 19 20 5 Michigan Department of Natural Resources, Wolf Lake State Fish Hatchery, 34270 County Road 652, Mattawan, MI 49071, USA. 21 22 23 24 6 Michigan Department of Natural Resources, Thompson State Fish Hatchery, 9445 South State Highway M149, Manistique, MI 498545, USA. 25 1 26 Abstract.— 27 The influences of stocking location on the survival of stocked fish are poorly 28 understood in the Great Lakes and only a few poorly replicated studies have 29 been carried out. To evaluate the survival of stocked Michigan strain steelhead 30 we investigated the relative survival and recreational fishery contributions of 31 coded-wire tagged fish stocked at up-, mid- and down-stream sites in four river 32 systems. Three are tributary to Lake Michigan (St. Joseph River, Big Manistee 33 River, and Muskegon River), and one tributary to Lake Huron (Au Sable River). 34 Tags were recovered through creel surveys, use of dedicated head collectors 35 and volunteer angler returns. Steelhead stocked in the Muskegon River on a 36 whole returned better followed by those planted in the Au Sable and St. Joseph 37 Rivers, which were similar. The Manistee River consistently produced lower 38 returns of stocked fish relative to the other study rivers. Contrary to reports from 39 previous investigations, up-stream stocking locations led to greater apparent 40 survival to open-lake and river fisheries. The pattern was observed for all four of 41 the study rivers and in all survey methodologies providing strong evidence for the 42 strength of the effect. To maximize the investment in hatchery fish, fisheries 43 managers and hatchery biologists would do well to consider upstream locations 44 when planning stocking programs for anadromous steelhead in the Great Lakes. 45 2 46 47 Introduction.— Anadromous rainbow trout or steelhead Oncorhynchus mykiss provide 48 important recreational fishing opportunities in the Great Lakes and tributary 49 streams. According to recent national surveys of hunting and fishing behavior, 50 from 2001 to 2006 nearly five percent of all angling nationwide occurred in Great 51 Lakes waters, representing US$1.5 billion dollars in expenditures per year (U.S. 52 Department of Interior 2006). These surveys further indicate that approximately 53 13% of Great Lakes anglers targeted steelhead when fishing. Over one-third of 54 all angling in the Great Lakes occurred on Lake Michigan, where steelhead 55 represented 5-15% of the recreational catch annually from 1996 to 2006 (Breidert 56 et al. 2007). 57 Both natural reproduction and supplemental stocking of hatchery-raised fish 58 maintain Great Lakes steelhead fisheries (Keller et al. 1990). Given the relatively 59 strong reliance on hatcheries to maintain fisheries, there is a keen interest in 60 maximizing post-stocking survival of hatchery fish. In Michigan, steelhead are 61 typically stocked as spring yearlings into tributary rivers of the Great Lakes. The 62 influences of stocking location on the survival of stocked fish are poorly 63 understood in the Great Lakes. Only a few poorly replicated studies have been 64 undertaken and reported. 65 In marine systems much thought has been given to the selection of stocking 66 locations for steelhead to optimize their survivability, spatial distributions, and 67 spawning runs (Ward and Slaney 1990; Wagner 1969). However, these marine 68 studies do not translate directly to the Great Lakes due to the substantially 69 different environmental conditions. 70 We investigate the relative survival and recreational fishery contributions of 71 steelhead stocked at up-, mid- and down-stream sites in four river systems. 72 Three are tributary to Lake Michigan (St. Joseph River, Big Manistee River, and 73 Muskegon River), and one is tributary to Lake Huron (Au Sable River). 74 75 Methods.— 76 Study site.—Three tributaries to Lake Michigan and one tributary to Lake Huron 3 77 were selected for this investigation, based on previous stocking histories, 78 suitability for survival of rainbow trout, and spatial location. The St. Joseph River, 79 located in southeast Michigan, the Muskegon River approximately 113 km to the 80 north and the Manistee River an additional 97 km north all flow into eastern Lake 81 Michigan (Figure 1). The Au Sable River flows into the west central Lake Huron 82 Basin. 83 The St. Joseph River drains approximately 12,134 km2 of southwest Michigan 84 and northern Indiana. The average annual discharge rate at the mouth is 130 85 m3/s. The main stem is 338 km long and there are 2,641 km of tributaries. The 86 river is predominately a warm-water system with some cold and cool water 87 habitats in tributaries (Wesley and Duffy 1999). 88 The Muskegon River drains approximately 6,086 km2, with an average annual 89 discharge rate of 56 m3/s. The river is classified as a coolwater stream with 90 moderate to low gradient flows below Croton Dam (O’Neal 1997). The main 91 stem is 341 km long and there are 94 named tributaries. 92 The Manistee River drains an area of approximately 4,557 km 2; the average 93 annual discharge rate is 57 m3/s. The river is considered a high quality coldwater 94 system. The main stem is 373 km long and there are 109 named tributaries 95 covering 3,694 km (Rozich 1998). 96 The Au Sable River drains approximately 5,000 km2, with an average annual 97 discharge rate of 43 m3/s. The river below Foote Dam is classified as a low 98 gradient cool water stream (Zorn and Sendek 2001). The main stem is 208 km 99 long and there are approximately 52 named tributaries covering 766 km (Zorn 100 101 and Sendek 2001). Impoundments influence runs of anadromous salmonids in all of the rivers 102 investigated. The St. Joseph River is the more heavily impounded system with 103 five fish ladders allowing upstream fish passage. A dam blocks upstream 104 movement 101 km upstream of the confluence near the Bodine State Fish 105 Hatchery (Twin Branch Dam) in Indiana (Figure 1). Tippy Dam (43 km upstream 106 of the confluence) blocks upstream movement of salmonids and other species on 107 the Manistee River, as do Croton Dam on the Muskegon and Foote Dam on the 4 108 Au Sable River (Figure 1). 109 110 Marking and stocking.—Gametes were obtained from weir collections of 111 naturalized adult runs of steelhead on the Little Manistee River during late March 112 and early April (Hay 2003). Fish were raised at the Thompson State Fish 113 Hatchery, Thompson, MI in 1996 and at Wolf Lake State Fish Hatchery, 114 Mattawan, MI from 1997 to 1999. 115 Prior to stocking, all study fish were marked as spring yearlings with a coded- 116 wire tag inserted in the snout (Wydoski and Emery 1983; Northwest Marine 117 Technology, Inc., Shaw Island, WA) and given an adipose fin clip. Uniquely 118 coded tags were used to identify each stocking site, and year-class (Table 1). 119 The actual number of marked fish counted as being released was adjusted for 120 tag loss, finclip quality and mortality experienced prior to stocking. 121 In each of four years (1996 to 1999), yearling smolts were stocked during late 122 April or early May. Stocking occurred prior to smolting, which typically takes 123 place in mid-May (Seelbach and Whelan 1988). Similar numbers were stocked 124 at multiple locations up- and down-stream in each of the river systems (Table 1; 125 Figure 1). During the first year (1996) fish were not stocked at the furthest 126 upstream site on the St. Joseph River due to logistical constraints (Table 1). 127 128 Tag recovery.—Tags were recovered through creel surveys, targeted collections 129 through use of dedicated head collectors and volunteer angler returns. In all, 130 nearly 7,500 tagged steelhead were recovered and the majority of these fish 131 (97%) were captured during five years from 1998 to 2002. Volunteer anglers 132 provided the greatest number of tag returns from Great Lakes and tributary 133 streams throughout the State of Michigan. Creel clerks collected more samples 134 from rivers, and dedicated head collectors more from Great Lakes open-water 135 fisheries (Table 2). 136 By collecting more detailed information, dedicated head collectors and creel 137 clerks provided data that enabled estimations of temporal and spatial scale 138 corrections for variations in collection efforts. Creel clerks collected marked fish, 5 139 but their primary responsibility was to obtain count and interview data used to 140 interpret overall recreational fishing effort and harvest. Tags were recovered 141 from Lake Michigan and Lake Huron fisheries in standardized lake-wide creel 142 surveys conducted from April through September each year from 1997 to 2004 in 143 State of Michigan waters (Lockwood 2000). Tags were recovered from the St. 144 Joseph River (1998 to 2004), Muskegon River (1999 to 2004), Manistee River 145 (1999 to 2004), and Au Sable River (1999 and 2000) fisheries through river 146 specific creel surveys conducted from March through December each year. 147 Targeted collection efforts (head-hunters) were not conducted for the Au Sable 148 River. 149 Dedicated collectors were seasonally employed personnel hired for the sole 150 purpose of collecting heads/snouts from fish with adipose fin clips which indicate 151 the likely presence of a coded wire tag in the snout. These individuals worked 152 the open-water fisheries of Lake Michigan from 1997 to 2006 and the St. Joseph 153 and Manistee River fisheries in 1998. These dedicated collectors recorded the 154 total number of fish evaluated, as well as the number with adipose fin clips for 155 each salmonid species per interview. 156 At the time of collection the total length, weight (if available), sex, location and 157 date were recorded and returned with the head/snout. Snouts were returned to 158 the MDNR Charlevoix Fisheries Research Station where tags were extracted, 159 read, and information entered into appropriate databases. Creel clerks collected 160 scale samples to facilitate age determination and definition of spawning history. 161 162 Analyses.— 163 Return rates.—The year-class and stocking site provided the base unit of 164 replication for comparisons within Great Lakes creel surveys, river specific creel 165 surveys, dedicated collections and volunteer angler returns. Returns of marked 166 fish were standardized (per 10,000 stocked) for each replicate to account for 167 variations actual numbers stocked. To compare return rates within a collection 168 methodology we used generalized linear model procedures. The dependent 169 variable was the standardized return rate and the independent factor was 6 170 stocking location within a river. 171 172 173 Results.— Comparisons among rivers indicate that the Muskegon River on a whole 174 produced the best steelhead returns followed by the Au Sable and St. Joseph 175 rivers which were similar (Wald 21,5>25.0, P<0.001). The Manistee River 176 consistently produced lower returns of stocked fish relative to the other study 177 rivers (Figures 2, 3, and 4). 178 For the St. Joseph River, fish stocked at the furthest upstream site at 179 Mishawaka (97 km from the mouth) produced the highest return rates in all three 180 open-lake survey methodologies. Fish stocked at the Arden rearing pond 181 location (23 km upstream from the mouth) generally ranked second in return 182 rates and all other sites were lower and similar to each other (Figures 2, 3, 4). In 183 creel surveys, the apparent survival (returns) of fish stocked at the up-stream 184 Mishawaka site was nearly twice that of fish stocked at the mid- and down- 185 stream sites on the St. Joseph River (Wald 21,4=8.91, P=0.06; Figure 2). 186 Whereas, results from targeted surveys and volunteer angler returns indicated 187 steelhead survival greater than 3.5 times that of the other sites (targeted returns, 188 Wald 21,4=15.7, P=0.003; volunteer returns, Wald 21,4=18.3, P=0.001; Figures 3 189 and 4). Returns from fish stocked at Arden Pond were generally 20 to 50 percent 190 greater than that those from fish stocked at other mid- and down-stream 191 locations. 192 For the Muskegon River, the three open-lake survey methodologies showed 193 similar patterns for returns of fish from different stocking locations (creel surveys, 194 Wald 21,2=7.2, P=0.028; targeted returns, Wald 21,2=7.5, P=0.023; volunteer 195 returns, Wald 21,2=8.6, P=0.013). Fish stocked at the upstream site experienced 196 the best survival, followed by those stocked at the mid- and down-stream 197 locations. Apparent survival of fish stocked at the upstream site on the 198 Muskegon River was over 3.5 times that of fish stocked at the down-stream 199 location and 20 to 30 percent greater than for fish stocked at the mid-stream 200 location; Figures 2, 3, and 4). 7 201 For the Manistee River, fish from the mid-stream stocking location produced 202 relatively poor returns in all open-lake survey methodologies. In all surveys, fish 203 from the upstream site produced the best returns by nearly 2 to 3 times those of 204 fish from the mid-stream location (creel surveys, Wald 21,2=5.2, P=0.074; 205 targeted returns, Wald 21,2=4.1, P=0.131; volunteer returns, Wald 21,2=8.8, 206 P=0.012). Targeted collections and volunteer angler returns also indicated poor 207 survival of fish stocked at the down-stream location. However, creel surveys 208 results showed near equal returns between fish stocked at up- and down-stream 209 locations (Figures 2, 3, and 4). 210 For the Au Sable River returns for fish stocked at the up-stream location were 211 always higher than those stocked at the downstream site. All three open-lake 212 survey methodologies indicated apparent survival that was 30 to 50 percent 213 greater for fish stocked at upstream versus downstream locations (creel surveys, 214 Wald 21,1=0.8, P=0.370; targeted returns, Wald 21,1=5.2, P=0.022; volunteer 215 returns, Wald 21,1=2.6, P=0.104). 216 Based on river specific creel surveys and volunteer angler returns it was clear 217 that steelhead had a strong propensity to return to the river in which they were 218 stocked (Table 3). Few fish strayed among systems as greater than 90% 219 returned to the same river system into which they had been stocked. Returns to 220 rivers represented patterns similar to those observed in Great Lakes surveys. 221 On the Au Sable River fish stocked at upstream sites produced the higher returns 222 in river fisheries relative to fish stocked at downstream sites. Fish stocked at the 223 upstream location on the Muskegon River contributed most to river fisheries 224 followed by fish stocked at the mid- and down-stream locations and similar 225 patterns were observed on the Manistee River. For the St. Joseph River fishery, 226 fish stocked at Mishawaka, the furthest upstream stocking location, again 227 outperformed fish stocked at all other locations. However, the returns of fish 228 stocked in Arden Pond were not much different from those stocked at the other 229 mid- or down-stream locations. The Muskegon River fishery had the best overall 230 return rates, followed by the St. Joseph River and the Au Sable River fisheries. 231 As we had observed in open-lake survey methodologies, the Manistee River 8 232 again produced the lowest returns of stocked fish in river fisheries. 233 234 235 Discussion.— Our results provide interesting insights into stocking strategies and survival of 236 stocked steelhead in tributaries of the Great Lakes. Contrary to reports from 237 previous marine investigations (Ward and Slaney 1990; Wagner 1969), up- 238 stream stocking locations in Great Lakes tributaries led to greater apparent 239 survival and contribution to open-lake and river fisheries. This pattern was 240 observed for all four of the study rivers and in all survey methodologies providing 241 strong evidence for the validity of the effect. It may be that predation pressures 242 on fish stocked in rivers are greater with increasing proximity to the Great Lakes. 243 It also may be that environmental conditions up-stream are somehow more 244 conducive to survival after stocking. One may also hypothesize that post- 245 stocking survival is somehow enhanced below impoundments, as the upstream 246 location on each of the rivers investigated is directly below an impoundment. 247 However, we would point out that there are four impoundments on the St. Joseph 248 River prior to reaching the furthest upstream location at Mishawaka. Three of our 249 study sites are located below the first impoundment and improved survival was 250 not evident at any of these locations. 251 There was a strong tendency for steelhead to return to the river in which they 252 were stocked and little straying was observed among river systems. Returns of 253 fish from the Arden Pond stocking location were somewhat higher than those of 254 fish from neighboring mid-stream stocking site locations. This may indicate some 255 improvement in survival for fish stocked at passive rearing facilities such as 256 holding ponds or net pens. Interestingly, the differences in returns of Arden 257 Pond-stocked fish to the river fishery were less pronounced for the open-lake 258 fishery. Often we assume that holding ponds or net pens improve imprinting as 259 well as survival thereby improving return rates to stocking locations. We also 260 note that observed improvements in survival were not substantial and that the 261 costs and benefits associated with rearing practices need to be considered 262 carefully before the practice is adopted wholesale. 9 263 It appears that fisheries managers and hatchery biologists would do well to 264 consider upstream locations when planning stocking programs. This is 265 particularly true for large cold to cool water rivers as investigated in our study. 266 267 Acknowledgments.—We would like to thank the many creel clerks, hatchery staff, 268 head-hunters and others with the Michigan and Indiana Department’s of Natural 269 Resources who have done an excellent job of raising and collecting steelhead for 270 this investigation. We also thank Tom Rozich (MDNR), Dr. Ed Rutherford of the 271 University of Michigan and Dr. Paul Seelbach of the University of Michigan and 272 MDNR, and Brian Briedert (INDNR) for their original ideas in initiating this project. 273 Randall Claramunt, John Clevenger, Eric Crissman, Pat O’neill and Patrick 274 Hanchin and other staff at the Charlevoix Fisheries Research Station who 275 provided invaluable input regarding the analysis and interpretation of results. 276 10 277 References.— 278 Hay, R. 2003. Little Manistee River weir steelhead egg-take report, 1967-92. 279 Michigan Department of Natural Resources, Technical Report 96-1, Ann 280 Arbor. 281 282 Lockwood, R. N. 2000. Conducting roving and access site angler surveys. 283 Chapter 14 in J. C. Schneider, editor. Manual of fisheries survey methods 284 II: with periodic updates. 285 Fisheries Special Report 25, Ann Arbor. Michigan Department of Natural Resources, 286 287 O’Neal, R. P. 1997. Muskegon River Watershed Assessment. Michigan 288 Department of Natural Resources, Fisheries Special Report 19, Ann 289 Arbor, MI. 290 291 292 Rozich, T. J. 1998. Manistee River Assessment. Michigan Department of Natural Resources, Fisheries Special Report 21, Ann Arbor, MI. 293 294 Seelbach, P. W., and G. E. Whelan. 1988. Identification and contribution of wild 295 and hatchery steelhead stocks in Lake Michigan tributaries. Michigan 296 Department of Natural Resources, Fisheries Research Report 1950, Ann 297 Arbor. 298 299 U.S. Department of the Interior, Fish and Wildlife Service, U.S. Department of 300 Commerce, U.S. Census Bureau. 2006. National Survey of Fishing, 301 Hunting, and Wildlife-Associated Recreation. U.S. Fish and Wildlife 302 Service. Available: 303 http://wsfrprograms.fws.gov/Subpages/NationalSurvey/2006_Survey.htm. 304 11 305 Wagner, H. H. 1969. Effect of stocking location of juvenile steelhead trout, Salmo 306 gairdneri, on adult catch. Transactions of the American Fisheries Society 307 98:27-34. 308 309 Ward, B. R., and P. A. Slaney. 1990. Returns of pen-reared steelhead from 310 riverine, estuarine, and marine releases. Transactions of the American 311 Fisheries Society 119:492-499. 312 313 Wesley, J. K., and J. E. Duffy. 1999. St. Joseph River Assessment. Michigan 314 Department of Natural Resources, Fisheries Special Report 24, Ann 315 Arbor. 316 317 Wydoski, R., and L. Emery. 1983. Tagging and marking. Pages 215-237 in L. 318 Nielsen and D. Johnson, editors. Fisheries Techniques. American 319 Fisheries Society, Bethesda, Maryland. 320 321 Zorn, T. G., and S. P. Sendek 2001. Au Sable River Assessment. Michigan 322 Department of Natural Resources, Fisheries Special Report 26, Ann 323 Arbor, MI. 324 12 Table 1.–Stocking locations, distance from the mouth (km) and numbers of marked (coded-wire tag and adipose fin clip) steelhead stocked into study rivers during 1996-1999. NS indicates sites not stocked. Distance River Stocking Location (km) 1996 1997 1998 1999 2 9,961 15,076 9,982 15,030 Sportsman’s Club-Arden 23 10,921 11,652 11,697 10,577 Shamrock Park-Berrien 35 9,847 14,923 10,173 10,049 Buchanan City Launch 50 9,801 14,780 10,107 9,987 Mishawaka-Lincoln Park 97 NS 19,819 20,317 20,054 Mouth <1 14,795 15,102 16,727 15,080 High Bridge 35 15,787 14,787 15,044 15,444 Tippy Dam 43 15,950 15,005 15,110 15,010 Mouth <1 10,163 10,056 10,180 10,095 Henning Park 34 21,489 19,965 20,218 20,022 Pine Street 44 22,072 20,198 20,180 20,058 Harbor <1 28,426 21,095 22,134 25,050 Rea Road 11 27,172 24,812 25,027 25,426 St. Joseph Pier 33 Manistee Muskegon Au Sable Total 196,384 217,270 206,896 211,882 13 1 Table 2.–Numbers of study fish returned in recreational fisheries by collection methodology and year. 1997 Creel survey collections Lake Michigan Manistee River Muskegon River St. Joseph River Lake Huron Au Sable River Targeted collections Lake Michigan Manistee River Muskegon River St. Joseph River Lake Huron Au Sable River Volunteer angler returns Lake Michigan Manistee River Muskegon River St. Joseph River Lake Huron Au Sable River Total Capture year 2001 2002 1998 1999 2000 2003 2004 2005 2006 Total 20 ------6 --- 59 ----273 15 --- 87 115 191 284 15 44 42 134 131 256 15 93 55 58 57 210 21 --- 20 42 43 89 4 --- 2 7 18 18 4 --- --5 2 9 1 --- ------------- ------------- 285 361 442 1,139 81 137 34 --5 --8 --- 100 117 49 54 15 --- 147 ------21 --- 102 ------31 --- 89 ------33 --- 49 ------6 --- 9 ----------- 2 ----------- 1 ----------- 1 ----------- 534 117 54 54 114 0 8 2 2 6 ----- 151 107 184 66 40 31 235 428 347 373 40 109 130 249 166 352 32 31 122 199 187 215 32 27 58 75 81 40 7 9 12 3 12 1 4 1 2 1 ----1 --- --1 --------- ------------- 718 1,065 979 1,053 156 208 91 1,261 2,436 1,764 1,305 523 91 23 2 1 7,497 2 14 3 Table 3.–Study fish (adjusted per 10,000 stocked) returned to study river watersheds during river creel surveys on the Big 4 Manistee, Muskegon and St. Joseph Rivers (1999 to 2004) and from volunteer angler returns (1997 to 2004). “--“ 5 indicates missing values for standard deviations which could not be estimated due to low sample size. Collection Method/Return Location (watershed) Creel Stocking Location Big Manistee Muskegon Volunteer St. Joseph Au Sable Big Manistee Muskegon St. Joseph Au Sable Big Manistee River Mouth 18.1±16.7 0.5±0.9 0.4±0.9 0 15.8±7.6 2.6±1.6 0 0 High Bridge 60.9±33.6 0.5±1.0 0.5±1.0 0 59.0±20.8 1.0±0.3 0.7±-- 0 Tippy Dam 91.1±36.2 1.6±2.1 0.5±1.0 0 84.0±30.8 0.9±0.4 0 0 Mouth 0 29.0±19.3 0 0 2.1±1.9 19.4±13.3 0 0 Henning Park 34.5±38.6 176.6±143.8 3.3±6.7 0 29.7±27.7 183.5±80.2 2.0±0.0 0 Pine Street 0 351.8±208.1 0 0 31.9±17.4 184.0±109.7 1.7±0.9 2.0±-- Pier 33 0 0 137.7±60.2 0 0.9±0.1 2.0±-- 34.7±7.7 0 Sportsman’s Club-Arden 0 1.2±2.3 115.4±18.7 0 1.3±-- 2.0±0.8 38.5±11.8 0 Shamrock Park-Berrien 0 0 154.6±70.3 0 1.0±0.3 2.0±-- 29.5±8.8 0 Buchanan City Launch 0 0.9±1.8 161.3±63.8 0 1.6±0.8 1.1±0.4 37.2±18.5 0 Mishawaka-Lincoln Park 0 4.3±3.8 531.9±262.6 0 4.0±2.8 3.0±1.6 193.8±157.0 0 Muskegon River St. Joseph River 15 Au Sable River Mouth 0 0 0 62.5±37.9 3.0±2.3 0 2.1±-- 30.9±31.3 Rea Road 0 0 0 83.2±44.3 5.6±2.8 0 2.1±-- 38.6±29.8 16 6 Figure captions.— 7 Figure 1. Map indicating key ports and locations of the Manistee, Muskegon and 8 St. Joseph rivers on Lake Michigan and the Au Sable River on Lake Huron. 9 Steelhead stocking locations are indicated by closed circles within expanded 10 detailed river sections. 11 12 Figure 2. Plot of steelhead returns to open-lake creel surveys per 10,000 fish 13 stocked for each river and site as indicated by distance upstream (kilometers). 14 15 Figure 3. Plot of open-lake steelhead returns from targeted collections per 16 10,000 fish stocked for each river and site as indicated by distance upstream 17 (kilometers). 18 19 Figure 4. Plot of open-lake volunteer angler steelhead returns per 10,000 fish 20 stocked for each river and site as indicated by distance upstream (kilometers). 21 17 22 Figure 1. Manistee River Tippy Dam # # # Au Sable River Leland Frankfort St. Joseph River # Foote Dam \ Manistee # # Ludington # Muskegon River # Croton Dam Berrien Springs Fish Ladder Grand Haven # Buchanan Fish Ladder Niles Fish Ladder \ # Holland South Haven St. Joseph New Buffalo 23 # # Legend: Stocking location Impoundment South Bend Fish Ladder # Mishawaka Fish Ladder 18 24 Figure 2. 25 26 11.4 Number per 10,000 stocked 10 8 8.8 6 4 2 9.2 9.0 5.9 7.0 6.2 4.5 5.4 4.0 4.8 1.7 3.5 Dis 0 tan 97 ce ups 50 44 trea 35 m ( 23 kilo 11 me ters 2 1 ) AuSable River Manistee Riv er Muskegon Riv er St. Joseph Riv er 19 Figure 3. 120 116.7 Number per 10,000 stocked 100 103.4 80 89.9 60 62.9 40 48.3 20 27.1 40.7 22.6 31.0 15.3 39.3 Dis 0 tan 97 ce ups 50 44 trea 35 m ( 23 kilo 11 me ters 2 1 ) 19.7 11.0 AuSable River Manistee Riv er Muskegon Riv er St. Joseph Riv er 20 Figure 4. 60 Number per 10,000 stocked 57.8 50 45.6 40 38.9 33.3 30 20 10 10.2 10.4 21.9 13.4 7.6 14.1 16.3 0 Dis tan 11.3 ce 97 50 ups 44 trea 35 m ( 23 kilo 11 me ters 2 1 ) 7.0 AuSable River Manistee Riv er Muskegon Riv er St. Joseph Riv er 21