Transactions of the American Fisheries Society 118:284-289, 1989 Rearing of Juvenile Chinook Salmon in Nonnatal Tributaries of the Lower Fraser River, British Columbia C. B. MURRAY AND M. L. ROSENAU! Department of Fisheries and Oceans, Biological Sciences Branch Pacific Biological Station. Nanaimo, British Columbia V9R 5K6 Canada Abstract. —Juvenile chinook salmon Oncorhynchus tshawytscha were collected from May to June 1980 in seven nonnatal tributaries of the lower Fraser River in British Columbia, Canada. The juveniles were collected predominantly in pools and had migrated 0.4-6.5 km up the nonnatal tributaries to reach the rearing areas. In 1981, two of the tributaries, Nathan Creek and the Brunette River, were sampled every 2 weeks from late February to late June. During this period, the mean fork lengths of juvenile chinook salmon increased from 45 to 80 mm in Nathan Creek and from 44 to 68 mm in the Brunette River. The use of nonnatal rearing areas is of importance to the survival and productivity of juvenile chinook salmon in the Fraser River. Freshwater residence by juvenile chinook salmon Oncorhynchus tshawytscha is highly variable. Some fish have "ocean-type" behavior in that they migrate seaward after a period of freshwater residence ranging from a few days to a few months. Others exhibit "stream-type" behavior in that they remain in fresh water for a year or longer before migrating seaward (Reimers 1973; Major et al. 1978; Healey 1983). Chinook salmon in the Fraser River exhibit both stream-type and ocean-type life history patterns (Fraser et al. 1982). Stocks of chinook salmon that spawn and rear in upper Fraser River tributaries tend to produce more streamtype juveniles than those in the lower Fraser River (Tutty and Yole 1978). The Harrison River, a large tributary of the lower Fraser River with about 25% of the mean annual runs of returning chinook salmon to the Fraser River, is probably the largest contributor of ocean-type juveniles in the whole system (Fraser et al. 1982). The ocean type of juvenile Fraser River chinook salmon varies in rearing and migratory behavior. The juveniles either migrate directly to the ocean with no freshwater growth, rear in the estuary for up to 60 d before moving offshore, or reside for 60-150 d in their natal streams before migrating to the estuary or ocean (Fraser et al. 1982; Levy and Northcote 1982). In the Nechako and Chilcotin rivers of the upper Fraser River system, migrating juvenile chinook salmon have been marked in the main stem and then recovered upstream from the release site and in nonnatal tributaries 1 Present address: Department of Biological Sciences, University of Waikato, Hamilton, New Zealand. of these rivers (Delaney et al. 1982; Russell et al. 1983). However, there has been no documentation of upstream migrations into nonnatal tributaries of the lower Fraser River. In this study, we describe and document the use of nonnatal tributaries of the lower Fraser River by juvenile chinook salmon for rearing, and we document their residence time, growth, and habitat distributions. Methods In 1980, 21 tributaries of the lower Fraser River in the Fraser Valley were sampled in May and June to determine the distribution and abundance of juvenile salmonids (Figure 1). In 1981, the Brunette River and Nathan Creek were sampled every 2 weeks from early February until late June; these two tributaries were selected for further investigation because they contained large numbers of juvenile chinook salmon in 1980. In both years, collections were made with an electroshocker (Smith-Root Type VII) and a stop net (2 m x 10 m with 0.75-cm stretch mesh). The collection sites were chosen according to juvenile salmonid distribution and segregated according to habitat type (pool, run, and riffle as defined by Helm 1985). Sampling was conducted in a representative reach (100 m in length) of each stream. The stop net was located downstream from each sampling habitat, and collecting continued until no more fish were captured. Juvenile chinook salmon from each habitat were identified, counted, anesthetized, and preserved in 10% formalin. All juveniles from each collection were blotted dry and then weighed to the nearest 0.01 g; fork length was measured to the nearest 0.1 mm. Differences in fork length and weight offish among 285 JUVENILE CHINOOK SALMON IN BRITISH COLUMBIA 10 20km FIGURE I.—Streams in the Fraser River (British Columbia, Canada) drainage sampled in 1980 and 1981 for the presence of juvenile salmonids. Circled numbers indicate streams that contained juvenile Chinook salmon. (1) Tincan Creek (near Vancouver), (2) Brunette River, (3) Coquitlam River, (4) Scott Creek, (5) Alouette River, (6) Kanaka Creek, (7) Whonnock Creek, (8) Silverdale Creek, (9) Lagace (Bouchier) Creek, (10) Chilqua (Thompson) Creek, (11) Norrish (Suicide) Creek, (12) Nicomen Slough, (13) Squakum Creek, (14) Hicks Creek, (15) Salmon River, (16) West Creek, (17) Nathan (Beaver) Creek, (18) Clayburn (Kelly) Creek, (19) Wade Creek, (20) Hope Slough, and (21) Silverhope Creek. rivers in 1980 and between rivers and among time periods in 1981 were tested by either one-way (river) or two-way (river, time) analysis of variance. stream surveys revealed additional juveniles, whereas upstream surveys did not. These observations indicated that juvenile chinook salmon used these seven nonnatal tributaries for rearing and that the collection sites were close to the most upstream distribution of the fish. Results In 1980, more juvenile chinook salmon were Juvenile chinook salmon were found in 7 of the collected from pools than from runs or riffles (Ta21 tributaries of the lower Fraser River sampled ble 1). Differences in fork length (F = 11.01, df = in 1980 (Figure 1). No young chinook salmon with 6, 203, P < 0.01) and weight (F = 5.67, P < 0.01) yolk sacs were collected, and there was no infor- were observed among tributaries. Juveniles colmation available from yearly (1947-1981) spawn- lected from the Nathan, West, and Brunette tribing ground surveys during probable spawning pe- utaries in early May were larger than those colriods to suggest that chinook salmon have ever lected from the other tributaries in late May and spawned in these seven tributaries. Collection sites early June. Higher indices of total dissolved solids with juvenile chinook salmon were located 0.4- were obtained from the Brunette, Nathan, and 6.5 km upstream from the confluence of the trib- West tributaries (Table 1), and indices of total utaries with the Fraser River (Table 1). These sev- dissolved solids were significantly correlated with en tributaries probably were accessible to juve- juvenile length (r = 0.87, df = 5, P < 0.05) and niles because of their low gradients and the absence weight (r = 0.83, P < 0.05). In addition to variof barriers, such as flood control gates, bridge foot- ation in size, there was considerable variation in ings, culverts, and waterfalls. At each site, down- fish density among tributaries (Table 1); higher 286 MURRAY AND ROSENAU TABLE 1.—Numbers of juvenile chinook salmon collected from different habitats, stream reach characteristics, and mean sizes of juveniles measured. Fish were collected during 1980 from sites on seven tributaries of the Fraser River, British Columbia. Location is number of kilometers upstream from the confluence with the Fraser River; TDS is the index of total dissolved solids; SEs are in parentheses. Reach characteristics Number offish collected from Tributary Date Pool Run Riffle Nathan West Brunette Scott Whonnock Squakum Wade Total % of total May8 May 9 May 12 May 21 May 29 JunS Jun 6 245 32 226 161 26 58 25 773 66 86 18 39 10 15 55 10 233 20 73 12 30 5 6 30 2 158 14 Water MaxitemperWetted mum area density TDS aturc Loca(m2) (fish/m2) 0«s/cm) (°Q tion 594 490 1,354 540 767 990 620 5,355 0.680 0.126 0.217 0.326 0.061 0.144 0.060 199.3 149.0 161.7 15.2 13.6 19.2 18.4 10.5 10.1 14.9 12.4 11.1 12.2 9.2 6.5 2.1 4.1 6.3 0.5 4.5 0.4 Fish measurements N Fork length (mm) 40 25 40 40 25 25 15 52.8(0.49) 54.4(1.72) 54.2(0.97) 47.7(1.00) 47.1(1.67) 49.9(0.92) 45.2(0.90) Wet weight (g) 1.88(0.06) 2.16(0.20) 2.14(0.14) 1.51(0.14) 1.24(0.16) 1.71(0.12) 1.30(0.09) densities occurred in Nathan and Scott creeks and migrations (upstream, downstream, or both) that in the Brunette River than in the other tributaries. are genetically and environmentally controlled In 1981, sampling was limited to the Brunette (Kelso et al. 1981). Most newly emerged chinook River and Nathan Creek. Juvenile chinook salm- salmon are carried downstream for several days on were collected from early April to late June in and then either develop residence behavior, mithe Brunette River and from late March to early grate back upstream, or actively continue the June in Nathan Creek (Table 2). The first juveniles downstream migration to the estuary (Lister and collected in each stream measured 43.6-44.7 mm Genoe 1970; Reimers 1973; Fraser et al. 1982; fork length and had a streamlined body shape with Anonymous 1987). Healey (1980) found that many no visible or residual yolk. This suggests that they young chinook salmon collected in the Nanaimo were not newly emerged. Sampling continued in River estuary between March and April were new Nathan Creek from early June to late September emergents with visible yolk. The first juveniles and produced no juvenile or adult chinook salm- collected in Nathan Creek and Brunette River were on. In each sampling period, water temperature past the yolk-sac stage. A survey of the lower Frawas higher in the Brunette River than in Nathan ser River (Schubert 1982) and extensive yearly Creek. In both tributaries, juvenile chinook showed surveys of spawning grounds conducted on foot significant increases in fork length (F = 1,011.6, by fisheries officers during probable chinook salmdf = 6, 231, P < 0.01) and weight (F = 240.8, P on spawning times (Hancock and Marshall 1985a, < 0.01) over the sampling period, but the juve- 1985b, 1985c) revealed no chinook salmon niles from Nathan Creek were consistently longer spawning in the seven tributaries where we found (F = 986.3, P < 0.01) and heavier (F = 130.2, P juveniles. This suggests that habitat requirements < 0.01) than those collected from the Brunette do not restrict juveniles to their natal streams and River. In both tributaries, the juveniles were found that the absence of spawners does not necessarily predominantly in pools. In 1981, densities of ju- indicate an absence of appropriate habitat. venile chinook salmon were usually higher in the Our findings and those of others suggest that the Brunette River than in Nathan Creek. However, dispersal and migratory patterns of young chinook densities within each stream were dissimilar be- salmon increase the use of available rearing areas tween years. In early May 1980, the numbers of (Chapman 1966; Reimers 1973; Fraser etal. 1982; juveniles collected from Nathan Creek and the Levy and Northcote 1982). The method of coloBrunette River were 404 and 295, respectively, nization is not known, but in a large system like and in early May 1981, they were 16 and 210, the Fraser River and its tributaries, a passive respectively. downstream migration over several days during a spring freshet could result in considerable downDiscussion stream displacement. Juvenile chinook salmon are Movements of young salmonids from spawning territorial (Chapman and Bjornn 1969), and terareas to rearing areas may consist of complex local ritorial interactions among and within salmonid 287 JUVENILE CHINOOK SALMON IN BRITISH COLUMBIA TABLE 2.—Habitat characteristics, number, and mean (SE) measurements of juvenile Chinook salmon collected during 1981 in the Brunette River and Nathan Creek, British Columbia, Canada. Sampling locations are the same as in Table 1. W is the number of juvenile Chinook salmon measured from each stream. Number offish collected from Tributary Date Pool Run Riffle Wetted area (m 2 ) Maximum density (fish/m2) Water temperature (°C) Fish measurements N Fork length (mm) Wet weight (g) Nathan Brunette Feb22 Fcb23 0 0 0 0 0 0 502 1,315 6.5 7.0 Nathan Brunette Mar 10 Mar 11 0 0 0 0 0 0 545 1,340 6.5 7.5 Nathan Brunette Mar 23 Mar 24 9 0 2 0 1 0 554 1.345 0.022 7.5 8.5 12 44.7(0.99) 1.10(0.08) Nathan Brunette Apr 6 Apr? 9 1 0 0 2 0 607 L375 0.018 8.0 9.5 11 1 51.6(0.65) 43.6 1.75(0.07) 0.75 Nathan Brunette Apr 21 Apr 22 15 382 16 160 34 54 557 1,340 0.117 0.445 9.0 10.0 40 40 52.2(0.45) 41.6(0.29) 1.82(0.05) 0.80(0.02) Nathan Brunette May 3 May 4 1 154 8 48 7 8 564 1,352 0.028 0.155 9.0 13.0 16 40 58.8(1.24) 43.4(0.68) 2.63(0.18) 1.03(0.07) Nathan Brunette May 22 May 23 11 22 3 2 3 1 600 1,365 0.028 0.018 11.0 14.0 17 25 70.0(1.18) 44.9(0.96) 4.79(0.24) 1.26(0.10) Nathan Brunette Jun 8 Jun9 8 26 1 0 0 0 582 1.345 0.015 0.019 12.0 13.5 9 26 80.5(1.55) 68.1(1.13) 6.65(0.72) 4.39(0.19) Nathan Brunette Total number offish % of total Jun 29 Jun 30 0 0 0 1 0 0 589 1.360 12.0 13.5 1 66.3 4.28 638 64 241 24 110 11 species cause downstream displacement of subordinate fish (Stein et al. 1972). However, dispersal to other rearing areas implies that the juveniles must imprint on their natal stream as alevins or emergents, not as smolts (Cooper and Hirsch 1982; Rehnberg et al. 1985). High tides coupled with high water flows in the main-stem Fraser River during spring freshets cause flows to stop or reverse in those lower tributaries that lack flood control systems. Juvenile chinook salmon displaced downstream and carried into the lower reaches of these tributaries could then migrate upstream and develop residency behavior. This is similar to colonization of tidal channels by young chinook salmon in the Fraser River estuary (Levy and Northcote 1982). Upstream migrations of these fish after initial downstream displacement have been observed in the Nechako River (Russell et al. 1983). Tributaries of the lower Fraser River tend to be less turbid than the main stem (Cooper 1965; Servizi and Martens 1987). Feeding success (reaction distance, capture rate, and percentage of prey ingested) of juvenile coho salmon O. kisutch decreases with increased turbidity (Berg and Northcote 1985), and young coho salmon reared in turbid water have reduced growth rates (Grouse et al. 1981). Therefore, those young chinook salmon that enter nonnatal tributaries of the lower Fraser River may be able to maximize their growth and survival. Small tributaries are generally more productive than large rivers (Mundie 1969), and the upstream migrations of up to 6.5 km suggest that these areas have productive rearing habitats. Levels of productivity also vary among streams, and differences in productivity could account for some of the differences in juvenile size we observed among streams. The larger size in spite of lower rearing temperatures and the stable abundance of juvenile chinook salmon in Nathan Creek suggest that juveniles rear there for a long period; conversely, the smaller size at higher rearing temperatures and the rapid increases and decreases in abundance over time suggest a short rearing period for juveniles in the Brunette River. Colonization of tributaries by chinook salmon stocks with differences in initial egg size (Beacham and Murray 1989) could also produce differences in juvenile size among streams. Growth of young chinook salmon rearing in 288 MURRAY AND ROSENAU various habitats and from many different stocks in the Pacific Northwest is remarkably consistent. Chinook salmon rearing in estuaries in southern British Columbia from early April to late June increase in fork length from 33 to 80 mm (Kask and Parker 1972; Healey 1980; Levy and Northcote 1982). Fish that remain in their natal streams increase from 37 to 83 mm fork length before migrating to the ocean in late July (Lister and Genoe 1970; Reimers 1973; Delaney et al. 1982; Russell et al. 1983; Anonymous 1987). Our finding that young chinook salmon increased in fork length from 42 to 80 mm suggests that growth in nonnatal tributaries is similiar to growth in natal streams and estuaries. Levy and Northcote (1982) reported an average density of 0.18 juveniles/m 2 in the Fraser River estuary. Rearing densities in our study were within this range and were considerably higher for the Brunette River. This suggests that the nonnatal tributaries of the Fraser River are important rearing areas for juvenile chinook salmon and that further research is needed to assess methods of colonization, survival, carrying capacity, and the origin of juveniles that use these tributaries. Acknowledgments The extensive field sampling necessary for this project was accomplished with the help of Sam Bowman, Jim Kristmanson, Wally Osterman, and Lynn Yamanaka. Wally Barner assisted in the data analysis. T. D. Beacham, J. H. Mundie, and W. R. Meehan provided helpful suggestions for improvements to the manuscript. We thank J. D. McPhail for supporting the study through his grant from the Natural Sciences and Engineering Research Council of Canada (grant A3451). References Anonymous. 1987. Nechako River project. Studies of juvenile chinook salmon in the Nechako River, British Columbia—1985 and 1986. Canadian Manuscript Report of Fisheries and Aquatic Sciences 1954. Beacham. T. D., and C. B. Murray. 1989. Variation in developmental biology of sockeye salmon (Oncorhynchus nerkd) and chinook salmon (O. tshawytscha) in British Columbia. Canadian Journal of Zoology 67:2081-2089. Berg, L., and T. G. Northcote. 1985. 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