Comparison of Survival of Winter-Run Steelhead Based on Stocking Location

1 Comparison of survival of winter-run steelhead based on stocking location 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 8 Station, 96 Grant Street, Charlevoix, Michigan 49720, USA. 9 10 2 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, 15 6889 N. State Road 327, Orland, IN 46776 USA. 16 17 4 Michigan Department of Natural Resources, Muskegon State Game Area, 18 7550 E. Messenger Road, Twin Lake, MI 49457, USA. 19 20 5 Michigan Department of Natural Resources, Wolf Lake State Fish Hatchery, 21 34270 County Road 652, Mattawan, MI 49071, USA. 22 23 6 Michigan Department of Natural Resources, Thompson State Fish Hatchery, 24 9445 South State Highway M149, Manistique, MI 498545, USA.

1 1 25Abstract.— 26The influences of stocking location on the survival of stocked fish are poorly 27understood in the Great Lakes and only a few poorly replicated studies have 28been carried out. To evaluate the survival of stocked Michigan strain steelhead 29we investigated the relative survival and recreational fishery contributions of 30coded-wire tagged fish stocked at up-, mid- and down-stream sites in four river 31systems. Three are tributary to Lake Michigan (St. Joseph River, Big Manistee 32River, and Muskegon River), and one tributary to Lake Huron (Au Sable River). 33Tags were recovered through creel surveys, use of dedicated head collectors 34and volunteer angler returns. Steelhead stocked in the Muskegon River on a 35whole returned better followed by those planted in the Au Sable and St. Joseph 36Rivers, which were similar. The Manistee River consistently produced lower 37returns of stocked fish relative to the other study rivers. Contrary to reports from 38previous investigations, up-stream stocking locations led to greater apparent 39survival to open-lake and river fisheries. The pattern was observed for all four of 40the study rivers and in all survey methodologies providing strong evidence for the 41strength of the effect. To maximize the investment in hatchery fish, fisheries 42managers and hatchery biologists would do well to consider upstream locations 43when planning stocking programs for anadromous steelhead in the Great Lakes.

2 2 44Introduction.— 45 Anadromous rainbow trout or steelhead Oncorhynchus mykiss provide 46important recreational fishing opportunities in the Great Lakes and tributary 47streams. According to recent national surveys of hunting and fishing behavior, 48from 2001 to 2006 nearly five percent of all angling nationwide occurred in Great 49Lakes waters, representing US$1.5 billion dollars in expenditures per year (U.S. 50Department of Interior 2006). These surveys further indicate that approximately 5113% of Great Lakes anglers targeted steelhead when fishing. Over one-third of 52all angling in the Great Lakes occurred on Lake Michigan, where steelhead 53represented 5-15% of the recreational catch annually from 1996 to 2006 (Breidert 54et al. 2007). 55 Both natural reproduction and supplemental stocking of hatchery-raised fish 56maintain Great Lakes steelhead fisheries (Keller et al. 1990). Given the relatively 57strong reliance on hatcheries to maintain fisheries, there is a keen interest in 58maximizing post-stocking survival of hatchery fish. In Michigan, steelhead are 59typically stocked as spring yearlings into tributary rivers of the Great Lakes. The 60influences of stocking location on the survival of stocked fish are poorly 61understood in the Great Lakes. Only a few poorly replicated studies have been 62undertaken and reported. 63 In marine systems much thought has been given to the selection of stocking 64locations for steelhead to optimize their survivability, spatial distributions, and 65spawning runs (Ward and Slaney 1990; Wagner 1969). However, these marine 66studies do not translate directly to the Great Lakes due to the substantially 67different environmental conditions. 68 We investigate the relative survival and recreational fishery contributions of 69steelhead stocked at up-, mid- and down-stream sites in four river systems. 70Three are tributary to Lake Michigan (St. Joseph River, Big Manistee River, and 71Muskegon River), and one is tributary to Lake Huron (Au Sable River). 72 73Methods.— 74Study site.—Three tributaries to Lake Michigan and one tributary to Lake Huron

3 3 75were selected for this investigation, based on previous stocking histories, 76suitability for survival of rainbow trout, and spatial location. The St. Joseph River, 77located in southeast Michigan, the Muskegon River approximately 113 km to the 78north and the Manistee River an additional 97 km north all flow into eastern Lake 79Michigan (Figure 1). The Au Sable River flows into the west central Lake Huron 80Basin. 81 The St. Joseph River drains approximately 12,134 km2 of southwest Michigan 82and northern Indiana. The average annual discharge rate at the mouth is 130 83m3/s. The main stem is 338 km long and there are 2,641 km of tributaries. The 84river is predominately a warm-water system with some cold and cool water 85habitats in tributaries (Wesley and Duffy 1999). 86 The Muskegon River drains approximately 6,086 km2, with an average annual 87discharge rate of 56 m3/s. The river is classified as a coolwater stream with 88moderate to low gradient flows below Croton Dam (O’Neal 1997). The main 89stem is 341 km long and there are 94 named tributaries. 90 The Manistee River drains an area of approximately 4,557 km2; the average 91annual discharge rate is 57 m3/s. The river is considered a high quality coldwater 92system. The main stem is 373 km long and there are 109 named tributaries 93covering 3,694 km (Rozich 1998). 94 The Au Sable River drains approximately 5,000 km2, with an average annual 95discharge rate of 43 m3/s. The river below Foote Dam is classified as a low 96gradient cool water stream (Zorn and Sendek 2001). The main stem is 208 km 97long and there are approximately 52 named tributaries covering 766 km (Zorn 98and Sendek 2001). 99 Impoundments influence runs of anadromous salmonids in all of the rivers 100investigated. The St. Joseph River is the more heavily impounded system with 101five fish ladders allowing upstream fish passage. A dam blocks upstream 102movement 101 km upstream of the confluence near the Bodine State Fish 103Hatchery (Twin Branch Dam) in Indiana (Figure 1). Tippy Dam (43 km upstream 104of the confluence) blocks upstream movement of salmonids and other species on 105the Manistee River, as do Croton Dam on the Muskegon and Foote Dam on the

4 4 106Au Sable River (Figure 1). 107 108Marking and stocking.—Gametes were obtained from weir collections of 109naturalized adult runs of steelhead on the Little Manistee River during late March 110and early April (Hay 2003). Fish were raised at the Thompson State Fish 111Hatchery, Thompson, MI in 1996 and at Wolf Lake State Fish Hatchery, 112Mattawan, MI from 1997 to 1999. 113 Prior to stocking, all study fish were marked as spring yearlings with a coded- 114wire tag inserted in the snout (Wydoski and Emery 1983; Northwest Marine 115Technology, Inc., Shaw Island, WA) and given an adipose fin clip. Uniquely 116coded tags were used to identify each stocking site, and year-class (Table 1). 117The actual number of marked fish counted as being released was adjusted for 118tag loss, finclip quality and mortality experienced prior to stocking. 119 In each of four years (1996 to 1999), yearling smolts were stocked during late 120April or early May. Stocking occurred prior to smolting, which typically takes 121place in mid-May (Seelbach and Whelan 1988). Similar numbers were stocked 122at multiple locations up- and down-stream in each of the river systems (Table 1; 123Figure 1). During the first year (1996) fish were not stocked at the furthest 124upstream site on the St. Joseph River due to logistical constraints (Table 1). 125 126Tag recovery.—Tags were recovered through creel surveys, targeted collections 127through use of dedicated head collectors and volunteer angler returns. In all, 128nearly 7,500 tagged steelhead were recovered and the majority of these fish 129(97%) were captured during five years from 1998 to 2002. Volunteer anglers 130provided the greatest number of tag returns from Great Lakes and tributary 131streams throughout the State of Michigan. Creel clerks collected more samples 132from rivers, and dedicated head collectors more from Great Lakes open-water 133fisheries (Table 2). 134 By collecting more detailed information, dedicated head collectors and creel 135clerks provided data that enabled estimations of temporal and spatial scale 136corrections for variations in collection efforts. Creel clerks collected marked fish,

5 5 137but their primary responsibility was to obtain count and interview data used to 138interpret overall recreational fishing effort and harvest. Tags were recovered 139from Lake Michigan and Lake Huron fisheries in standardized lake-wide creel 140surveys conducted from April through September each year from 1997 to 2004 in 141State of Michigan waters (Lockwood 2000). Tags were recovered from the St. 142Joseph River (1998 to 2004), Muskegon River (1999 to 2004), Manistee River 143(1999 to 2004), and Au Sable River (1999 and 2000) fisheries through river 144specific creel surveys conducted from March through December each year. 145Targeted collection efforts (head-hunters) were not conducted for the Au Sable 146River. 147 Dedicated collectors were seasonally employed personnel hired for the sole 148purpose of collecting heads/snouts from fish with adipose fin clips which indicate 149the likely presence of a coded wire tag in the snout. These individuals worked 150the open-water fisheries of Lake Michigan from 1997 to 2006 and the St. Joseph 151and Manistee River fisheries in 1998. These dedicated collectors recorded the 152total number of fish evaluated, as well as the number with adipose fin clips for 153each salmonid species per interview. 154 At the time of collection the total length, weight (if available), sex, location and 155date were recorded and returned with the head/snout. Snouts were returned to 156the MDNR Charlevoix Fisheries Research Station where tags were extracted, 157read, and information entered into appropriate databases. Creel clerks collected 158scale samples to facilitate age determination and definition of spawning history. 159 160Analyses.— 161Return rates.—The year-class and stocking site provided the base unit of 162replication for comparisons within Great Lakes creel surveys, river specific creel 163surveys, dedicated collections and volunteer angler returns. Returns of marked 164fish were standardized (per 10,000 stocked) for each replicate to account for 165variations actual numbers stocked. To compare return rates within a collection 166methodology we used generalized linear model procedures. The dependent 167variable was the standardized return rate and the independent factor was

6 6 168stocking location within a river. 169 170Results.— 171 Comparisons among rivers indicate that the Muskegon River on a whole 172produced the best steelhead returns followed by the Au Sable and St. Joseph

2 173rivers which were similar (Wald  1,5>25.0, P<0.001). The Manistee River 174consistently produced lower returns of stocked fish relative to the other study 175rivers (Figures 2, 3, and 4). 176 For the St. Joseph River, fish stocked at the furthest upstream site at 177Mishawaka (97 km from the mouth) produced the highest return rates in all three 178open-lake survey methodologies. Fish stocked at the Arden rearing pond 179location (23 km upstream from the mouth) generally ranked second in return 180rates and all other sites were lower and similar to each other (Figures 2, 3, 4). In 181creel surveys, the apparent survival (returns) of fish stocked at the up-stream 182Mishawaka site was nearly twice that of fish stocked at the mid- and down-

2 183stream sites on the St. Joseph River (Wald  1,4=8.91, P=0.06; Figure 2). 184Whereas, results from targeted surveys and volunteer angler returns indicated 185steelhead survival greater than 3.5 times that of the other sites (targeted returns,

2 2 186Wald  1,4=15.7, P=0.003; volunteer returns, Wald  1,4=18.3, P=0.001; Figures 3 187and 4). Returns from fish stocked at Arden Pond were generally 20 to 50 percent 188greater than that those from fish stocked at other mid- and down-stream 189locations. 190 For the Muskegon River, the three open-lake survey methodologies showed 191similar patterns for returns of fish from different stocking locations (creel surveys,

2 2 192Wald  1,2=7.2, P=0.028; targeted returns, Wald  1,2=7.5, P=0.023; volunteer

2 193returns, Wald  1,2=8.6, P=0.013). Fish stocked at the upstream site experienced 194the best survival, followed by those stocked at the mid- and down-stream 195locations. Apparent survival of fish stocked at the upstream site on the 196Muskegon River was over 3.5 times that of fish stocked at the down-stream 197location and 20 to 30 percent greater than for fish stocked at the mid-stream 198location; Figures 2, 3, and 4).

7 7 199 For the Manistee River, fish from the mid-stream stocking location produced 200relatively poor returns in all open-lake survey methodologies. In all surveys, fish 201from the upstream site produced the best returns by nearly 2 to 3 times those of

2 202fish from the mid-stream location (creel surveys, Wald  1,2=5.2, P=0.074;

2 2 203targeted returns, Wald  1,2=4.1, P=0.131; volunteer returns, Wald  1,2=8.8, 204P=0.012). Targeted collections and volunteer angler returns also indicated poor 205survival of fish stocked at the down-stream location. However, creel surveys 206results showed near equal returns between fish stocked at up- and down-stream 207locations (Figures 2, 3, and 4). 208 For the Au Sable River returns for fish stocked at the up-stream location were 209always higher than those stocked at the downstream site. All three open-lake 210survey methodologies indicated apparent survival that was 30 to 50 percent 211greater for fish stocked at upstream versus downstream locations (creel surveys,

2 2 212Wald  1,1=0.8, P=0.370; targeted returns, Wald  1,1=5.2, P=0.022; volunteer

2 213returns, Wald  1,1=2.6, P=0.104). 214 Based on river specific creel surveys and volunteer angler returns it was clear 215that steelhead had a strong propensity to return to the river in which they were 216stocked (Table 3). Few fish strayed among systems as greater than 90% 217returned to the same river system into which they had been stocked. Returns to 218rivers represented patterns similar to those observed in Great Lakes surveys. 219On the Au Sable River fish stocked at upstream sites produced the higher returns 220in river fisheries relative to fish stocked at downstream sites. Fish stocked at the 221upstream location on the Muskegon River contributed most to river fisheries 222followed by fish stocked at the mid- and down-stream locations and similar 223patterns were observed on the Manistee River. For the St. Joseph River fishery, 224fish stocked at Mishawaka, the furthest upstream stocking location, again 225outperformed fish stocked at all other locations. However, the returns of fish 226stocked in Arden Pond were not much different from those stocked at the other 227mid- or down-stream locations. The Muskegon River fishery had the best overall 228return rates, followed by the St. Joseph River and the Au Sable River fisheries. 229As we had observed in open-lake survey methodologies, the Manistee River

8 8 230again produced the lowest returns of stocked fish in river fisheries. 231 232Discussion.— 233 Our results provide interesting insights into stocking strategies and survival of 234stocked steelhead in tributaries of the Great Lakes. Contrary to reports from 235previous marine investigations (Ward and Slaney 1990; Wagner 1969), up- 236stream stocking locations in Great Lakes tributaries led to greater apparent 237survival and contribution to open-lake and river fisheries. This pattern was 238observed for all four of the study rivers and in all survey methodologies providing 239strong evidence for the validity of the effect. It may be that predation pressures 240on fish stocked in rivers are greater with increasing proximity to the Great Lakes. 241It also may be that environmental conditions up-stream are somehow more 242conducive to survival after stocking. One may also hypothesize that post- 243stocking survival is somehow enhanced below impoundments, as the upstream 244location on each of the rivers investigated is directly below an impoundment. 245However, we would point out that there are four impoundments on the St. Joseph 246River prior to reaching the furthest upstream location at Mishawaka. Three of our 247study sites are located below the first impoundment and improved survival was 248not evident at any of these locations. 249 There was a strong tendency for steelhead to return to the river in which they 250were stocked and little straying was observed among river systems. Returns of 251fish from the Arden Pond stocking location were somewhat higher than those of 252fish from neighboring mid-stream stocking site locations. This may indicate some 253improvement in survival for fish stocked at passive rearing facilities such as 254holding ponds or net pens. Interestingly, the differences in returns of Arden 255Pond-stocked fish to the river fishery were less pronounced for the open-lake 256fishery. Often we assume that holding ponds or net pens improve imprinting as 257well as survival thereby improving return rates to stocking locations. We also 258note that observed improvements in survival were not substantial and that the 259costs and benefits associated with rearing practices need to be considered 260carefully before the practice is adopted wholesale.

9 9 261 It appears that fisheries managers and hatchery biologists would do well to 262consider upstream locations when planning stocking programs. This is 263particularly true for large cold to cool water rivers as investigated in our study. 264 265Acknowledgments.—We would like to thank the many creel clerks, hatchery staff, 266head-hunters and others with the Michigan and Indiana Department’s of Natural 267Resources who have done an excellent job of raising and collecting steelhead for 268this investigation. We also thank Tom Rozich (MDNR), Dr. Ed Rutherford of the 269University of Michigan and Dr. Paul Seelbach of the University of Michigan and 270MDNR, and Brian Briedert (INDNR) for their original ideas in initiating this project. 271Randall Claramunt, John Clevenger, Eric Crissman, Pat O’neill and Patrick 272Hanchin and other staff at the Charlevoix Fisheries Research Station who 273provided invaluable input regarding the analysis and interpretation of results.

10 10 274References.— 275Hay, R. 2003. Little Manistee River weir steelhead egg-take report, 1967-92. 276 Michigan Department of Natural Resources, Technical Report 96-1, Ann 277 Arbor. 278 279Lockwood, R. N. 2000. Conducting roving and access site angler surveys. 280 Chapter 14 in J. C. Schneider, editor. Manual of fisheries survey methods 281 II: with periodic updates. Michigan Department of Natural Resources, 282 Fisheries Special Report 25, Ann Arbor. 283 284O’Neal, R. P. 1997. Muskegon River Watershed Assessment. Michigan 285 Department of Natural Resources, Fisheries Special Report 19, Ann 286 Arbor, MI. 287 288Rozich, T. J. 1998. Manistee River Assessment. Michigan Department of Natural 289 Resources, Fisheries Special Report 21, Ann Arbor, MI. 290 291Seelbach, P. W., and G. E. Whelan. 1988. Identification and contribution of wild 292 and hatchery steelhead stocks in Lake Michigan tributaries. Michigan 293 Department of Natural Resources, Fisheries Research Report 1950, Ann 294 Arbor. 295 296U.S. Department of the Interior, Fish and Wildlife Service, U.S. Department of

297 Commerce, U.S. Census Bureau. 2006. National Survey of Fishing,

298 Hunting, and Wildlife-Associated Recreation. U.S. Fish and Wildlife

299 Service. Available:

300 http://wsfrprograms.fws.gov/Subpages/NationalSurvey/2006_Survey.htm.

11 11 302Wagner, H. H. 1969. Effect of stocking location of juvenile steelhead trout, Salmo 303 gairdneri, on adult catch. Transactions of the American Fisheries Society 304 98:27-34.

306Ward, B. R., and P. A. Slaney. 1990. Returns of pen-reared steelhead from 307 riverine, estuarine, and marine releases. Transactions of the American 308 Fisheries Society 119:492-499.

309 310Wesley, J. K., and J. E. Duffy. 1999. St. Joseph River Assessment. Michigan 311 Department of Natural Resources, Fisheries Special Report 24, Ann 312 Arbor. 313 314Wydoski, R., and L. Emery. 1983. Tagging and marking. Pages 215-237 in L. 315 Nielsen and D. Johnson, editors. Fisheries Techniques. American 316 Fisheries Society, Bethesda, Maryland. 317 318Zorn, T. G., and S. P. Sendek 2001. Au Sable River Assessment. Michigan 319 Department of Natural Resources, Fisheries Special Report 26, Ann 320 Arbor, MI. 321

12 12 322Table 1.–Stocking locations, distance from the mouth (km) and numbers of 323marked (coded-wire tag and adipose fin clip) steelhead stocked into study rivers 324during 1996-1999. NS indicates sites not stocked. Distance River Stocking Location (km) 1996 1997 1998 1999 St. Joseph Pier 33 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 Manistee 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 Muskegon 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 Au Sable Harbor <1 28,426 21,095 22,134 25,050 Rea Road 11 27,172 24,812 25,027 25,426 Total 196,384 217,270 206,896 211,882

13 13 14 325Table 2.–Numbers of study fish returned in recreational fisheries by collection methodology and year.

Capture year 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 Total Creel survey collections Lake Michigan 20 59 87 42 55 20 2 ------285 Manistee River ------115 134 58 42 7 5 ------361 Muskegon River ------191 131 57 43 18 2 ------442 St. Joseph River --- 273 284 256 210 89 18 9 ------1,139 Lake Huron 6 15 15 15 21 4 4 1 ------81 Au Sable River ------44 93 ------137 Targeted collections Lake Michigan 34 100 147 102 89 49 9 2 1 1 534 Manistee River --- 117 ------117 Muskegon River 5 49 ------54 St. Joseph River --- 54 ------54 Lake Huron 8 15 21 31 33 6 ------114 Au Sable River ------0 Volunteer angler returns Lake Michigan 8 151 235 130 122 58 12 2 ------718 Manistee River 2 107 428 249 199 75 3 1 1 --- 1,065 Muskegon River 2 184 347 166 187 81 12 ------979 St. Joseph River 6 66 373 352 215 40 1 ------1,053 Lake Huron --- 40 40 32 32 7 4 1 ------156 Au Sable River --- 31 109 31 27 9 1 ------208 Total 91 1,261 2,436 1,764 1,305 523 91 23 2 1 7,497 326

15 14 16 327Table 3.–Study fish (adjusted per 10,000 stocked) returned to study river watersheds during river creel surveys on the Big 328Manistee, Muskegon and St. Joseph Rivers (1999 to 2004) and from volunteer angler returns (1997 to 2004). “--“ 329indicates missing values for standard deviations which could not be estimated due to low sample size. Collection Method/Return Location (watershed)

Creel Volunteer Stocking Location Big Manistee Muskegon 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

Muskegon River 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±--

St. Joseph River 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

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

17 15 18 330Figure captions.— 331Figure 1. Map indicating key ports and locations of the Manistee, Muskegon and 332St. Joseph rivers on Lake Michigan and the Au Sable River on Lake Huron. 333Steelhead stocking locations are indicated by closed circles within expanded 334detailed river sections. 335 336Figure 2. Plot of steelhead returns to open-lake creel surveys per 10,000 fish 337stocked for each river and site as indicated by distance upstream (kilometers). 338 339Figure 3. Plot of open-lake steelhead returns from targeted collections per 34010,000 fish stocked for each river and site as indicated by distance upstream 341(kilometers). 342 343Figure 4. Plot of open-lake volunteer angler steelhead returns per 10,000 fish 344stocked for each river and site as indicated by distance upstream (kilometers). 345

19 16 20 346Figure 1. Manistee River

Tippy Dam

Au Sable River Leland 

 Frankfort Foote Dam

St. Joseph River \ # #  Manistee #

 Ludington

# Muskegon River Croton Dam # Berrien Springs Fish Ladder #\  Grand Haven #

 Holland # Buchanan Fish Ladder Niles Fish Ladder  South Haven

 St. Joseph #

 New Buffalo 347 Legend: Stocking location South Bend Fish Ladder Impoundment # Mishawaka Fish Ladder

21 17 22 348Figure 2. 349 350

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