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Pacific salmon (Oncorhynchus spp.) early marine feeding patterns based on 15N/14N and 13C/12C in Prince William Sound, Alaska
Thomas C. Kline, Jr., and T. Mark Willette

Abstract: Nitrogen and carbon mass and stable isotope composition among cohorts of Pacific salmon (Oncorhynchus spp.) released from Prince William Sound, Alaska, hatcheries in 1994 varied widely, suggesting a range in early marine feeding patterns. Analyses consisted of whole-body stable carbon and nitrogen mass and stable isotope composition of selected release-date cohorts that had been identified by implanted coded wire tags (CWT). Nitrogen isotopic and mass shifts suggested that the initial protein pool within individual fish was replaced at different rates among cohorts. There was a notable difference in carbon source dependency among hatcheries. Salmon from the hatchery closest to the Gulf of Alaska had a 13C-depleted carbon signature consistent with Gulf carbon, whereas salmon from the other hatcheries had Sound signatures. Differences in early marine feeding histories among 1994 hatchery-release-date cohorts reconstructed from the stable isotope composition of fry bore no relationship to marine survival pattern. Varied survival rates of 1994 Prince William Sound hatchery salmon were more likely related to the fry size at time of release, the observed differences in growth rate among release cohorts, and predation refuge effects of pen-rearing. Rsum : Les masses dazote et de carbone et la composition en isotopes stables varient considrablement dune cohorte lautre chez les saumons du Pacifique (Oncorhynchus spp.) provenant de piscicultures du dtroit du Prince William, Alaska, ce qui laisse croire quils possdent une gamme de patterns alimentaires diffrents au dbut de leur vie en mer. Nous avons dtermin les masses dazote et de carbone stables dans le corps entier, ainsi que la composition en isotopes stables, chez des cohortes choisies en fonction de la date de leur mise en eau et dont les individus avaient t marqus par limplantation de fils de fer cods par couleur. Les variations des isotopes et de la masse dazote indiquent que le pool initial dazote est remplac des taux divers chez les poissons des diffrentes cohortes. Il y a des diffrences apprciables entre les sources de carbone utilises par les poissons des diffrentes piscicultures. Les saumons provenant de la pisciculture la plus proche du golfe de lAlaska ont une signature de carbone appauvrie en 13C, ce qui correspond au carbone prsent dans le golfe, alors que les saumons des autres piscicultures ont une signature de carbone correspondant au carbone du dtroit. Les variations de lalimentation durant les premiers temps en mer chez les cohortes de pisciculture de dates de libration diffrentes en 1994, telles que rvles par ltude de la composition en isotopes stables des alevins, ne sont pas relies aux patterns de survie en mer. Les taux variables de survie des saumons provenant des piscicultures du dtroit du Prince William en 1994 sont plus probablement relis la taille des alevins au moment de leur libration, aux diffrences observes dans les taux de croissance chez les cohortes libres et aux effets de refuge dus llevage en enclos. [Traduit par la Rdaction] Kline and Willette 1638

Introduction
Approximately 500 million Pacific salmon (Oncorhynchus spp.) fry are released into the waters of Prince William Sound (PWS), Alaska, each year from five private nonprofit hatcheries (Cooney 1993). These salmon initially feed within PWS, then migrate into the Gulf of Alaska (GOA), returning to PWS as maturing fish one or more (depending upon the species) years later. Survivorship of maturing salmon returning to PWS hatcheries has declined in recent years. Presently, survival rates, which are based on comparing population size

at date of release with return, generally range between 3 and 5% (Willette et al. 1999b). Survival of salmon from juvenile to adult stages is thought to be determined by mortality occurring during early marine life history (Parker 1968; Hartt 1980; Bax 1983). Manipulating time of release by PWS hatchery managers was explored as a means for increasing marine survival rate (Willette et al. 1995, 1996). In 1994, pink salmon (Oncorhynchus gorbuscha, the numerically dominant Pacific salmon species in the study area) fry that were released in June at two PWS hatcheries, at a larger size, had survival rates of 7 and 22% compared with 1% survival for

Received 29 March 2001. Accepted 17 July 2002. Published on the NRC Research Press Web site at http://cjfas.nrc.ca on 11 October 2002. J16292 T.C. Kline, Jr. Prince William Sound Science Center, 300 Breakwater Ave., P.O. Box 705, Cordova, AK 99574, U.S.A. T.M. Willette. Alaska Department of Fish and Game, 43961 Kalifornsky Beach Road Suite B, Soldotna, AK 99835-7563, U.S.A.
1

Corresponding author (e-mail: tkline@grizzly.pwssc.gen.ak.us).


DOI: 10.1139/F02-126 2002 NRC Canada

Can. J. Fish. Aquat. Sci. 59: 16261638 (2002)

Kline and Willette

1627 48 39 43 6 41 13 46 15 July July July 8 May to 16 8 May to 10 26 Jun to 12 7 and 8 July 7 and 8 July 4 June to 16 5 June to 16 4 June to 12 Pink Pink Pink Pink Pink Chum Chum Sockeye
2002 NRC Canada Note: The survival rate of chum salmon was not tracked. WHN, W.H. Noerenberg hatchery; AFK, A.F. Koernig hatchery; MB, Main Bay hatchery; ND, not determined. a Data from Willette et al. (1996), Joyce and Riffe (1995), and provided by the Prince William Sound Aquaculture Corp. b Among pink salmon releases. c Detailed listings showing sample size for each species, hatchery, release date, and capture date combination are given in Table A1 among pink salmon releases.

Table 1. Survival rates and sampling information of select Prince William Sound hatchery salmon release cohorts.

Survival to adult (%)a

Materials and methods


Study area Salmon naturally migrate into PWS from adjacent natal (mostly intertidal and short streams) and juvenile rearing (lakes and streams) habitats or are released into PWS from hatcheries during AprilMay. This time period in PWS is marked by energy-rich diatom and Neocalanus blooms, which are food sources for juvenile salmon production (Cooney et al. 2001). Salmon then rear in PWS during the transition from winter to summer conditions (Cooney et al. 2001). Accordingly, seasonal transitioning, including a concomitant complex of oceanographic processes, in part due to the fjord and estuarine nature of PWS, alters the distribution of potential food sources for salmon and salmon predators (Gay and Vaughan 2001; Vaughan et al. 2001; Wang et al. 2001). This early marine life-history phase when salmon are rearing within PWS, which is thought to be important for determining year-class strength and has been the subject of recent intensive investigation (Cooney et al. 2001), was the focus of this study. Following PWS rearing, salmon emigrate to the greater Gulf of Alaska, only returning to spawn as a maturing fish one or more years later (Cooney et al. 2001).

Wet mass at releasea

Number released (millions)a

Hatchery releases

Hatchery

Salmon species

WHN WHN WHN AFK AFK WHN WHN MB

Late April Early May Early June Early May Early June Late May Early June All

Release period

48.0 154.7 7.7 84.8 7.0 12.0 10.0 3.8

0.4 0.4 1.4 0.2 1.1 1.1 6.0 7.9

1.0 0.4 22.1 0.4 7.2 ND ND 14.4

those released during the conventional period of April to May (Table 1). Processes leading to fish mortality are frequently dichotomized into bottom up versus top-down control, which in this case, is the relative importance of salmon forage versus salmon-as-forage, respectively. The availability of zooplankton forage, especially Neocalanus copepods, is important also because it may cause a shift in the behavior of salmon predators, in addition to being important as salmon prey (Cooney et al. 2001; Willette et al. 2001). Neocalanus and other zooplankton from PWS were found to be distinguishable by their stable carbon isotope ratio content from those from the adjacent GOA during 1994 (Kline 1999). Juvenile salmon sampled off Vancouver Island appeared to have fed on isotopically distinctive food sources (Perry et al. 1996). We determined whether natural stable isotope abundance based trophic differences in early marine stage salmon, suggested by a pilot study (Kline 1996), were important for marine survival. There is a predictable relationship between the isotopic composition of a consumer and that of its diet (reviewed by Michener and Schell 1994). A secondary consumers isotopic signature reflects that of its diet, but with a consistent increase in the heavy isotopes 15N and 13C relative to diet (Vander Zanden and Rasmussen 2001). Isotopic composition shifts occurring at low trophic levels can thus propagate to higher trophic levels. A change in diet to one of differing isotopic composition can thus be measured in a secondary consumer like salmon. The approach taken here was to determine whether carbon or nitrogen isotopic composition patterns paralleled observed survival patterns for certain identifiable (by tagging) hatchery-produced salmon cohorts from 1994. If it could be shown that isotopic and survival patterns were concordant, then it could be argued that prey was a factor affecting survival, since isotopic signatures derive from diet.

Survival to adult (%)a

Capture datesc

Isotope sample-specific releases

Wet mass at releasea

Cohort code

A B C D E

Relative survivalb

Low Low High Low High High

25 April 7 May 11 June 10 and 17 May 13 June 30 May 3 June 28 to 31 May

Release dates

0.4 0.4 1.4 0.2 1.1 1.1 6.0 7.9

1.0 0.2 22.3 1.0 6.9 ND ND 14.4

July July July

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Can. J. Fish. Aquat. Sci. Vol. 59, 2002 Fig. 1. Map of the Prince William Sound study area showing locations of salmon hatcheries and its geographic location within Alaska, U.S.A. (insert).

Sampling The sampling of CWT-marked salmon by Willette et al. (2001) enabled us to perform this study. Approximately one in every 600 salmon released from PWS hatcheries in 1994 was labeled with a CWT indicating release date and hatchery of origin, thus enabling the selection of specific cohorts (Willette et al. 1999a). Fine-mesh purse seines were used to sample juvenile salmon during their migration from western PWS hatchery-release sites from early May through mid-July 1994 (Willette et al. 1999a, 2001). Schools of fish were targeted for seining after being located visually in near-shore nursery habitats. CWT fish were immediately identified using a portable CWT detector. Samples were logged into a database that included release cohort information for each fish. Samples were stored frozen (20C) organized by capture date. The CWT samples selected for this study were based on release cohorts that yielded samples over the May to July sampling period, as well as samples analyzed in a pilot study (Table 1, Table A1; Kline 1996). Salmon selected for this study originated primarily from the Wallace H. Noerenberg (WHN) hatchery (Fig. 1) because the migration path of these salmon within PWS was known (Willette et al. 1999a, 2001). Furthermore, their route along Knight Island Passage within PWS enabled multiple recapture of the individual release cohorts over the spring to summer out-migration period (Fig. 1, Willette et al. 1999a; see Table A1 for sampling dates). However, sampling was destructive (fish caught were removed from the pool) and characterized by decreasing probability of resampling over time owing to dispersion (after a few days from release, frequently only one individual from a given cohort was found in a day of sampling, Table A1), which is a function of zooplankton density (Willette 2001). This study focused on pink salmon because they dominate both natural and artificial salmon production in PWS (Cooney 1993). A total of 1192 pink salmon fry with CWT were recovered from the 1 199 348 individuals that were sampled from May to July 1994 (Willette et al. 1995). Marine survival (total number returning as maturing fish minus number of fry released) rates of pink salmon hatchery releases were dichotomized into high (>5% survival) or low (1% survival) relative survival from provided data (Table 1). To also provide a comparison by species, a majority of the recovered CWT chum salmon fry (Oncorhynchus keta; 93 CWT recovered, all from WHN hatchery) and sockeye salmon fry (Oncorhynchus nerka; 23 CWT recovered, all from Main Bay (MB) hatchery; Fig. 1) were also analyzed. Data from a pilot study (Kline 1996) that included pink salmon fry from the Armin F. Koernig (AFK) hatchery (Fig. 1) are included here for comparison with pink salmon fry from the WHN hatchery. Laboratory analysis CWT fry samples were stored frozen (20C) in their sampling vials until they were selected for analysis. Selected samples were thawed, fork length was measured to nearest 0.5 mm, and then samples were refrozen and freeze-dried. Freeze-dried weight was determined and then samples were ground to a fine powder with a dental amalgamator. Pilot study (Kline 1996) samples followed the same protocol omitting the length and weight measurements (these samples

are indicated in Table A1). Powdered samples were sent to the University of Alaska Fairbanks Stable Isotope Facility for analysis. A mass spectrometric analysis generated the following data: 13C/12C and 15N/14N ratios expressed in standard delta units, 13C and 15N, respectively; and %C and %N. The delta notation used to express stable isotope ratios is reported as the per mil () deviation relative to an international standard, air for nitrogen, and Vienna Pee Dee belemnite (VPDB) for carbon. By definition, the isotope standards have delta values of zero, i.e., 15N = 0 for atmospheric N2. Mass spectrometric analysis quality assurance protocols consisted of running of laboratory standards before and after groups of five unknowns and duplicate analyses of each sample. Samples were run again when duplicate analyses differed by more than 0.6. Data analysis Data used for analysis consisted of mean 13C, 15N, and N and C mass, and C/N of the duplicate determinations made for each fish. The %C and %N data were used to calculate C/N atom ratios. Nitrogen and carbon mass was calculated by multiplying the freeze-dried weight by the %N and %C, respectively. Stable carbon isotope ratios were normalized for lipid content following the methods of McConnaughey and McRoy (1979) and expressed as 13C following Kline (1999). Release date was subtracted from capture date to calculate days since release for use as an independent variable for each fish. Data were computer fitted using Graphpad Prism 3.0 (GraphPad Software, Inc., San Diego, Calif.) to simple mathematical models (Table A2) according to fit determined by R2, and when appropriate, P value (significance level was 0.05) and the ratio of predicted half-time (half-life or doubling time) confidence intervals (the smaller the ratio the better the fit). Considering only mass-balance constraints, growth in N and C mass was expected to dilute the original isotopic
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composition in proportion to mass increase. For example, if a salmon with initial isotopic composition A grew on a diet that resulted in isotopic composition B, the isotopic composition would be halfway between A and B when the initial mass was doubled, i.e., the fish would consist of 50% initial material and 50% new material. Once a fish increased in mass by an order of magnitude, the isotopic composition would equal 10% initial material plus 90% new material. Accordingly, isotopic estimates were made for an initial 15N of 13.0 and a 15N of 10.0 for a salmon virtually rid of its initial N mass, i.e., 100% new material, as B, above. The variable initial condition was N mass. Starting mass values were 5, 7.5, 10, 15, and 20 mg N. The terminology isotope dilution model is used herewith to refer to the results of this predictable shift. Salmon isotopic patterns were characterized and compared qualitatively among high and low survival cohorts. Mean cohort 13C values were characterized as high or low, depending on whether they were greater than or less than 13C = 22.0 after showing evidence of growth, i.e., having turned over their initial natal or hatchery signature (Kline et al. 1993). A cohorts nitrogen isotope turnover was characterized as rapid when the regression line of 15N vs. time crossed 15N = 11.5 before 30 days, and slow when the cohort crossed it 30 days after release and later. Similar isotopic patterns would be conferred if all cohorts within a survival category (low vs. high marine survival) had either slow or rapid nitrogen turnover or all had either high or low 13C.

Results
Following release into Prince William Sound from hatcheries, carbon and nitrogen mass increased exponentially (Fig. 2). Salmon released on later dates, between 30 May and 11 June (all chum and sockeye fry and June WHN pink fry), attained ~200 mg N and ~1000 mg C in 3040 days compared with 6070 days for the April and May releases. An overall decline in C/N atom ratio with time suggested a difference in C and N mass turnover (Fig. 2c). The C/N ratios of WHN pink salmon fry released in April and May declined more slowly than the chum salmon fry released from WHN from late May to June (Fig. 2c). Data points for June WHN pink and MB sockeye fry (Fig. 2c) occur near the WHN chum regression, whereas May AFK pink fry fell in with the April and May WHN regressions and June AFK pink fry fell in between conferring a speedier C/N decline for late spring releases. Fry 13C content depended upon location (AFK is located ~80 km south of the WHN hatchery), whereas 15N content depended upon growth (change of the fry mass with time). From initial 15N values of ~+13, 15N content declined rapidly during the first two to three weeks following release, then appeared to become asymptotic at values ranging from approximately +10 (e.g., May WHN pink salmon) to about +11 (e.g., April WHN pink salmon; Fig. 3a). While the rapid increase in mass was reflected by a concomitant change in 15N composition, 13C composition remained approximately constant (regression lines ranged up to ~1; Fig. 3b). Although C/N declined, the C/N-based lipid normalization (only normalized data are shown) changed pink, chum, and sockeye salmon 13C values by just 0.4 (stan-

dard deviation, SD = 0.5), 0.5 (SD = 0.3), and 0.2 (SD = 0.2), respectively, compared with the raw data. The 13C composition of early and late releases from a given hatchery ranged about 2. Nevertheless, there was a significant correlation for 13C vs. time (Fig. 3b, Table A2). AFK salmon fry were 13C depleted by 34 compared with those from the other hatcheries (Fig. 3b). Ontogenetic shifts in 15N content were suggested from most xy plots of 15N and N mass (Fig. 4a). The very rapid decline in 15N (Fig. 3a) corresponded to when fry N mass was <~50 mg. Salmon fry appear to have reached a 15N minimum when N mass was between ~50 and 150 mg. May WHN fry were 15N depleted by about 0.5 compared with April and June releases when in the 15N-minimum size range. Fry >~150 mg N (corresponding to length >~90 mm) appear to have increased in 15N. Many of the larger fish, particularly those with 15N values >~11.5, were sockeye salmon. Data aggregated into 10-day averages enabled simultaneous elucidation of the effects of calendar date, number of days since release, and cohort, suggesting that N mass and 15N decreased more slowly for salmon released before day 130 compared with those released later (Table 2). For example, April salmon exceeded 200 mg N and had 15N values <11.0 after about two months compared with about one month for those released on day 150 and later. The isotope dilution model results fitted a hyperbolic model with R2 > 0.99. The effect of increased starting mass on the isotope dilution model was to shift the curves upwardly (Fig. 4a). The May WHN pink cohort fitted well to the isotope dilution model, whereas the others did not. However, their asymptote was closer to 10.4, rather than 10.0 as used in the model calculations. Shift in 15N values among cohorts was correlated to C/N decline (Fig. 4b). When salmon fry approached asymptotic 15N values (Fig. 3a), C/N ratios were ~4.5 to 4.0 (Fig. 2c), suggesting that C/N could be used as an independent variable to ascertain turnover of initial material (Fig. 4b). A range in asymptotic 15N values can be seen in the right portion of Fig. 4b above the abscissa for the C/N range <4.5; the slopes for April and May WHN pink and sockeye salmon were similar, ranging from ~1 to 1.3 (Fig. 4b, Table A2). The apparent lesser slope and lower R2 for chum salmon may, in part, have reflected their lower initial C/N (cf. Fig. 2c). The ratio C/N was used as an independent variable to distinguish salmon that had turned over their natal C (Fig. 5). Species other than chum salmon with C/N <4.5 ranged in 13C value from about 19.0 to 24.0 (Fig. 5). The 13C dichotomy of ~3 between AFK salmon and the others occurred at C/N values consistent with initial material turnover. Accordingly, only AFK salmon were categorized as having low 13C (Table 3). Qualitative assessment suggested that there were both high and low 13C and both rapid and slow 15N turnover rates within each survival category (Table 3). Of the high marine survival cohorts (Table 1), only MB sockeye salmon had sufficient data to ascertain slow turnover from the regression of 15N with days since release, crossing the +11.5 line at the 30-day mark. Because the 15N June WHN pink salmon was already +11 before 20 days, they must have
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Fig. 2. (a) Coded wire tagged (CWT) fry nitrogen mass in relation to time, in days since release from hatcheries, for three W.H. Norenberg (WHN) hatchery pink salmon (Oncorhynchus gorbuscha) release dates (25 April, 7 May, and 11 June 1994; indicated by Apr WHN Pink, May WHN Pink, and June WHN Pink, respectively), WHN hatchery chum salmon (Oncorhynchus keta) released from 30 May to 3 June (indicated by WHN Chum), and Main Bay (MB) hatchery sockeye salmon (Oncorhynchus nerka) released from 28 to 31 May 1994 (indicated by MB Sockeye). (b) CWT salmon fry carbon mass in relation to time, in days since release from hatcheries, for the same cohorts as shown in a. (c) CWT salmon fry C/N atom ratio in relation to time, in days since release from hatcheries, for the same cohorts as shown in a with the addition of data for the A.F. Koernig (AFK) hatchery pink salmon releases of 10 and 17 May 1994, which are indicated in the legend as May AFK Pink, and the AFK pink salmon hatchery release of 13 June, which is indicated as June AFK Pink.

crossed +11.5 earlier, assuming they had starting 15N values similar to the other cohorts. Their turnover was thus categorized as rapid. Because the 15N values of June AFK pink

salmon were about +11.5 before 30 days, their turnover rate was categorized as rapid. It is assumed that their starting 15N was similar to WHN salmon. April and May WHN
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Fig. 3. (a) Coded wire tagged (CWT) salmon fry 15N in relation to time, in days since release from hatcheries, for three W.H. Norenberg (WHN) hatchery pink salmon (Oncorhynchus gorbuscha) release dates (25 April, 7 May, and 11 June 1994; indicated by Apr WHN Pink, May WHN Pink, and June WHN Pink, respectively), WHN hatchery chum salmon (Oncorhynchus keta) released from 30 May to 3 June (indicated by WHN Chum in the legend), Main Bay (MB) hatchery sockeye salmon (Oncorhynchus nerka) released from 28 to 31 May 1994 (indicated by MB Sockeye), A.F. Koernig (AFK) hatchery pink salmon releases of 10 and 17 May 1994, which are indicated in the legend as May AFK Pink, and the AFK pink salmon hatchery release of 13 June, which is indicated in the legend as June AFK Pink. The reference line is at 15N = 11.5, which was used as the criterion to differentiate rapid and slow nitrogen isotopic turnover. (b) Only a slight change of WHN and MB CWT salmon fry 13C in relation to time, in days since release from hatcheries, for the same cohorts as shown in a. AFK cohorts, however, were significantly 13C depleted, suggesting that the Gulf of Alaska was the carbon source.

pink salmon had low marine survival; however, the former had slow 15N turnover, since the regression crossed at 30 days, whereas the latter had rapid turnover because the regression crossed before day 20. May AFK pink salmon, a low marine survival cohort, were categorized as having slow 15N turnover because their values were relatively high (+11.0 to 11.5) at 50 days from release, especially when compared with May WHN pink salmon at 50 days.

Discussion
Isotopic shifts concomitant with salmon fry growth Given a difference between the isotopic composition of salmon diet in PWS and that of their natal or hatchery food source, rapidly growing fry were expected to shift their isotopic composition in proportion to their increase in body mass (Hesslein et al. 1993). This was the case for 15N; the

high 15N content natal-hatchery signature was replaced 34 weeks after release (more quickly for earlier releases), suggesting that after ranging free for a month, their bodies consisted principally of protein acquired from feeding in the natural environment. Our observations were similar to those for sockeye salmon fry feeding in a lake, which shifted from a parental-derived natal isotopic signature until they were ~4045 mm in length (Kline et al. 1993). In contrast to 15N, salmon fry from the WHN and MB hatcheries did not change appreciably in 13C content because 13C signatures derived from the natal-hatchery period and those resulting from consuming potential food sources in PWS appear not to have differed appreciably. The salmon fry 15N shift of ~2.5 reflected incorporation of 15N-depleted (relative to their natal-hatchery signature) N mass during growth. A 15N change model based solely upon dilution by N mass increase and given assumed
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Fig. 4. (a) The relationship of coded wire tagged (CWT) salmon fry 15N with N mass for three W.H. Norenberg (WHN) hatchery pink salmon (Oncorhynchus gorbuscha) release dates (25 April, 7 May, and 11 June 1994; indicated by Apr WHN Pink, May WHN Pink, and June WHN Pink, respectively), WHN hatchery chum salmon (Oncorhynchus keta) released from 30 May to 3 June (indicated by WHN Chum in the legend), and Main Bay (MB) hatchery sockeye salmon (Oncorhynchus nerka) released from 28 to 31 May 1994 (indicated by MB Sockeye). The thin lines indicate change in 15N predicted by isotopic dilution model for initial N masses ranging from 5 to 20 mg indicated by inset legend. (b) Linear (abscissa reversed so that data are approximately chronological, from left to right, along each line) relationship of CWT salmon fry 15N released from indicated PWS hatcheries in 1994 in relation to C/N, with the addition of data for the A.F. Koernig (AFK) hatchery pink salmon releases of 10 and 17 May 1994, which are indicated in the legend as May AFK Pink, and the AFK pink salmon hatchery release of 13 June, which is indicated as June AFK Pink.

initial and assumed final isotopic values produced results that were in the range of our observations when N mass was small (<~50 mg). Observed variability in the 15N data among the cohorts could thus, in part, be explained by differences in initial mass. Because the 15N content of PWS biota is proportional to trophic level (e.g., Kline and Pauly 1998), shifts to increased trophic level with size could explain how larger salmon fry had elevated 15N values. Herbivorous zooplankton species like the copepod Neocalanus had 15N values of ~+8.0 (Kline 1999). Omnivorous and carnivorous zooplankton had correspondingly higher 15N values (Kline and Pauly 1998; Kline 1999). Based on trophic 15 N enrichment, salmon fry consuming only herbivores were expected to have 15N values of ~+11.4. This estimate, however, must be examined the context of the isotopic variability of herbivores. For example, the Neocalanus 15N SD value from PWS is ~1.0 (Kline 2001b). Furthermore, PWS zooplankton 15N values increased during early May of 1994

when phytoplankton exhausted the nutrient supply (Kline 1999). The slower decrease in 15N of fry released in April may thus have reflected feeding on this more positive food source, i.e., a food source less different from their natal-hatchery signature than those released in May, since these latter fry fed when the 15N values of production had rebounded to less positive values. Nevertheless, May WHN fry attained their least positive 15N values of +10.4 at ~125 mg N on approximately day (of the year) 180 when April releases were larger, but slightly more positive (>200 mg and 15N = +10.8). Larger fry had 15N values up to +12.5, suggesting a shift to a diet averaging approximately one third of a trophic level higher than that reflected by a 15N value of +11.4 (trophic level estimated using formula in Kline and Pauly 1998). Salmon with >150 mg N may therefore have consumed a greater proportion of higher (than herbivore) trophic level prey than smaller fry. This conjecture is sup 2002 NRC Canada

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Table 2. Mean N mass and isotopic ratio for 1994 salmon fry in relation to release date (cohort) and time since release with standard deviation (in parentheses) and N (when N > 1).
Days since release from hatchery Release cohort (day of year, species)

110

1120 18.7 (11.2) 4 12.5 (0.4) 22 15.6 (4.0) 8 11.5 (0.2) 5 39.8 11.8 96.5 (38.0) 26 10.9 (0.2) 27

2130

3140

4150 77.9 (18.0) 14 11.1 (0.4) 14

5160

6182 236.5 (102.5) 8 10.8 (0.3) 11

Day 115, pink salmon mg N 27.8 12.5 15N Day 127, pink salmon mg N 6.0 (2.5) 6 12.8 (0.6) 5 15N Days 150 and 154, chum salmon mg N 20.1 (6.8) 47 12.5 (0.3) 47 15N Day 162, pink salmon mg N 15N

29.6 (2.8) 5 11.0 (0.2) 5 227.5 (45.6) 3 10.8 (0.3) 3 166.8 (63.1) 11 10.9 (0.2) 11 227.0 (63.0) 5 10.9 (0.2) 5 321.7 (54.1) 8 11.2 (0.3) 8

127.8 (33.2) 18 10.4 (0.2) 19

175.8 10.6

ported by stomach content analyses made by M. Sturdevant (given in appendix 4 of Oakey and Pauly 1998) of pink and chum salmon fry (size range not specified) during 1994 that suggest a diet consisting, in part, of a combination of carnivorous and herbivorous zooplankton. Stomach contents for both species also included fish, presumably larvae, in approximately the same proportion as carnivorous and herbivorous zooplankton combined, when averaged over time. In May, large calanoids and euphausiids (life stages not specified) were the dominant constituents in pink salmon diets, whereas fish and large calanoids were the dominant constituents in chum salmon diets. Large calanoids were less important in June and July for pink salmon, while small calanoids were more important. Gastropods (presumably the common pteropod Limacina) became the second most important diet component, after fish, for both pink and chum salmon fry during July. Because zooplankton taxa other than the large calanoid copepod Neocalanus had higher 15N, and therefore had a higher trophic level compared to Neocalanus (Kline and Pauly 1998; Kline 1999), the reduced proportion of large calanoids in the stomach content analysis could explain the 15N increases measured in salmon during July. Carbon isotopic composition It was possible to assess dietary information from salmon fry 13C content for those fry with C derived from the open (nonnatal nonhatchery) environment because of C and N mass increases, direct evidence of turnover from 15N data, and the C/N decline from ~6 to ~4.04.5, which took place during early marine feeding. Furthermore, the C/N decline suggested that the natal-hatchery C pool was lost more rapidly than N. The declining C/N of hatchery-produced salmon from time of release also suggested a concomitant decrease in lipid content and therefore whole-body energetic content (McConnaughey and McRoy 1979). This conjecture, which depends upon the relationship between C/N and fish whole-body energetic content (Paul et al. 2001), was supported by the observation of Paul and Willette (1997) that whole-body energetic content of hatchery pink salmon sampled near the WHN hatchery was lower at the end of May compared to early May, while body length increased.

The distinctive 13C values of AFK salmon compared with those from the more northern hatcheries in PWS reflected feeding on a separate, 13C-depleted food source. Such a difference in food source can be taxonomically independent because a single copepod species, Neocalanus cristatus, in the study area exhibited the same 13C range as observed here for salmon fry (Kline 1999). The species composition of diet can thus be invariant, e.g., 100% Neocalanus, when at the same time the isotopic composition of diet can be highly variable. Therefore, isotopically based diets must be referred to in nontaxonomic terms like food sources. Organic carbon (food sources) originating in the GOA in 19941995 was characterized as 13C depleted compared to food sources originating within PWS (Kline 1999). The proximity (~15 km) of the AFK hatchery to the GOA (Fig. 1), and the direct connection to it via one passage is consistent with the notion that AFK hatchery fry primarily consumed Gulf carbon. In contrast, the other hatcheries are >50 km from the Gulf and connect to it via several passages. Isotopic patterns versus survival A far greater proportion of the pink salmon released in June 1994 from both the AFK and WHN hatcheries survived to maturity than did earlier 1994 releases. Did early marine feeding history differences detected with stable isotopes confer any survival advantage? Large differences in food source were apparent from 13C values that, however, appeared to have a hatchery-derived geographic dependency that was not related to survival, since salmon from the AFK and WHN hatcheries consisted of releases with both high and low survival. The dependency on PWS vs. GOA carbon during early marine feeding appeared to confer no survival advantage to 1994 salmon fry because such a dependency difference was not observed within a hatchery where survival differences occurred. Salmon that were released from hatcheries from the end of May (~day 150) and later were larger, either from faster growth (in terms of change in either C or N mass) or from being released at a larger size, than those released earlier. The fry released at a large size in June exhibited much higher survival rates. These differences could have resulted
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Fig. 5. 13C in relation to C/N ratio for three W.H. Norenberg (WHN) hatchery pink salmon (Oncorhynchus gorbuscha) release dates (25 April, 7 May, and 11 June 1994; indicated by Apr WHN Pink, May WHN Pink, and June WHN Pink, respectively), WHN hatchery chum salmon (Oncorhynchus keta) released from 30 May to 3 June (indicated by WHN Chum), Main Bay (MB) hatchery sockeye salmon (Oncorhynchus nerka) released from 28 to 31 May 1994 (indicated by MB Sockeye), A.F. Koernig (AFK) hatchery pink salmon releases of 10 and 17 May 1994, which are indicated in the legend as May AFK Pink, and the AFK pink salmon hatchery release of 13 June, which is indicated in the legend as June AFK Pink.

Table 3. Qualitative assessment of high- and low-mortality cohorts to isotopic pattern. Cohort code from Table 1 High survival C E F Low survival A B D Relative N isotope turnover Rapid Rapid Slow Slow Rapid Slow Relative C isotope signature High Low High High High Low

because (i) the larger fry were less vulnerable to predation due to their size, or (ii) the large fry group was being held in net pens while the earlier release groups were subjected to a period of high predation losses. Willette et al. (1999b) found no significant differences in the mean wet weight, length, and condition factor between the early and large late release groups when they were mixed together in late June. Therefore, the second alternative was more likely the cause of the different survival rates between the two groups. The effects of temperature and prey density on growth were possible factors determining salmon fry mortality (Willette et al. 1999b). Water temperature may have been an important factor affecting material turnover, since seasonal warming of the water column (Cooney et al. 2001) accelerated growth and protein turnover rates in later-released fry. Warming water temperatures may also explain the accelerated C/N change of late May and June releases. The more rapid C/N ratio decline of later releases suggested an accelerated consumption of initial lipid stores (assuming C/N as a lipid proxy; McConnaughey and McRoy 1979) that was probably due to greater metabolic demand from seasonal warming and that should have been an additional factor af-

fecting these fish. From their N mass change, protein turnover was also suggested to be very rapid. These observations imply that foraging rates for the later releases were greater than for earlier releases, but this is is a contraindication of a hypothetical inverse relationship between foraging stress and survival rate (Cooney 1993). Year-to-year differences in zooplankton density, particularly if they are due to variation in the GOA carbon subsidy like those evidenced here for AFK fry, could be suggested by isotopic analysis of fishes such as salmon fry or their predators (Kline 2001a). Willette et al. (1999b) described results from a statistical model that predicted survival to adult from size, number, and timing of hatchery salmon releases and subsequent duration of the macrozooplankton bloom. Recent modeling studies indicate that year-to-year variability in PWS pelagic productivity patterns may reflect weather conditions occurring during the spring phytoplankton bloom period (Eslinger et al. 2001). Productivity-affected salmon growth rates would, in turn, affect vulnerability to predation, because slower-growing fish are susceptible to predation for a longer time (Parker 1971; West and Larkin 1987). It may be possible to elucidate pelagic productivity patterns, including long-term change, from stable isotope data. For example, Schell (2001) conjectured from systematic decrease in both 13C and 15N content of bowhead whale baleen that primary productivity in the Bering Sea has declined over multidecadal time periods. It is thus tempting to speculate that the carbon isotope differences ascribed to GOA and PWS carbon observed in juvenile salmon may also reflect differences in primary productivity within the region. This speculation is supported by productivity modeling that suggested greater zooplankton productivity during cold and stormy spring conditions than warm and calm springs (Eslinger et al. 2001). Eslinger et al. (2001) suggested that the latter pattern leads to a rapid but short-lived PWS phytoplankton blooms, increasing contributions to the benthos because the phytoplankton cannot be fully utilized by the zoo 2002 NRC Canada

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plankton. Fast algal growth rate and concomitant [CO2]aq and nutrient depletion occurring during warm springs is a nonexclusive cause for increased phytoplankton 13C values (e.g., Cullen et al. 2001). These higher 13C values when propagated up the food chain may thus be an indicator of this type of phytoplankton bloom. The higher 13C values found in PWS may therefore reflect the greater water stability characteristic of this type of bloom and of PWS relative to GOA (Vaughan et al. 2001). Consequently, the WHN salmon may have been more dependent on zooplankton that had fed on phytoplankton that had undergone more rapid growth than those from AFK, thus reflecting a bottom-up effect (i.e., a physicsnutrientphytoplankton interaction effect propagated to the fish). Nonetheless, this difference between the hatcheries was not manifested by a parallel difference in survival. Simulation models suggested that survival rate patterns could be explained by differences in effects of predation among release cohorts (Willette et al. 2001). If predation on salmon fry was the principal factor controlling marine survival, isotopically evidenced differences in food source and trophic shifts occurring in 1994 appeared not to predispose fry to this loss. Alternative hypotheses therefore need to be considered. For example, the later releases may have been less vulnerable to predation because of predator behavior (e.g., prey-switching; Willette et al. 1999a). Such differences in vulnerability would not be reflected by fry isotopic signature. Thus, the highly variable survival rates of 1994 PWS hatchery salmon were more likely due to causes other than those that can be inferred from differences in observed isotope abundance. Other applications The turnover of a natal or hatchery isotopic signature is important for interpretation of field data (Kline et al. 1993). For example, Perry et al. (1996) conjectured that the observed variability in stable isotope abundance among fry sampled near Vancouver Island reflected isotopic differences in diet of hatchery and wild (nonhatchery) salmon. This study suggested protocols for selecting fry so that a data set will reflect only postnatal or posthatchery feeding based on N mass gain or C/N decline. The empirical criteria found here may be useful in another setting until local criteria can be established. In this study, fry with C/N < 4.5 and N mass > ~50 mg reflected postnatal or hatchery feeding. Salmon fry selected on these criteria exhibited a range in pelagic organic carbon source dependency based on 13C content. Although these criteria could be applied to other studies, caution should be used because temperature appeared to be an important factor in regulating turnover in PWS. The variability in turnover rate in PWS suggested that time per se (e.g., a specified period since release) may not be an effective criterion for selecting fish not containing significant proportions of previously acquired material. Elucidation of material turnover from initial (isotopic and mass) values, which was aided by knowing capture and release dates of individual hatchery salmon that were sampled after free-ranging for up to three months, would not have been possible without CWTs and a large sampling effort. CWTs are being supplanted by otolith thermal marking (OTM) technology. OTM technology makes it possible to determine very effectively which salmon in a sample from

PWS (and other locations where the technology is applied) are from hatcheries, and from an absence of markings, which are from natural propagation. Whereas OTM technology enables the mass marking of virtually all of a hatcherys production (all PWS hatchery salmon are presently being OTM marked), there will be a trade-off, since OTM fish may not be marked specifically for a date of release. In summary, we have shown that salmon fry exhibited a variety of early marine feeding histories based on their having a large range in both carbon and nitrogen stable isotope abundance. These patterns of isotopic variability could not be related directly to marine survival rates. Additionally, our data provided direct evidence for rapid turnover of material in early stage juvenile fishes through a noncaptive feeding experiment complementing previous captive studies (Hesslein et al. 1993).

Acknowledgments
The research described in this paper was funded by the Exxon Valdez Oil Spill Trustee Council. The findings presented by the authors are their own and not necessarily the Trustee Councils position. We were assisted by a large crew from the Alaska Department of Fish and Game that collected fish specimens. M. Powell and K. Antonucci facilitated the retrieval of archived salmon samples. K. Antonucci prepared samples for mass spectrometry. Bruce Barnett at the University of Alaska Fairbanks Stable Isotope Facility performed the stable isotope analysis. V. Patrick provided suggestions for the data analysis. Tim Joyce of the Alaska Department of Fish and Game provided hatchery data. S. Baumann prepared the map shown in Fig. 1. This paper benefitted from suggestions made by Ole A. Mathisen, three anonymous reviewers, and an anonymous associate editor.

References
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1636 Hartt, A.C. 1980. Juvenile salmonids in the oceanic ecosystemthe critical first summer. In Salmonid ecosystems of the North Pacific. Edited by W.J. McNeil and D.C. Himsworth. Oregon State University Press, Corvallis, Oreg. pp. 2558. Hesslein, R.H., Hallard, K.A., and Ramlal, P. 1993. Replacement of sulfur, carbon, and nitrogen in tissue of growing broad whitefish (Coregonus nasus) in response to a change in diet traced by 34S, 13C, and 15N. Can. J. Fish. Aquat. Sci. 50: 20712076. Joyce, T., and Riffe, R. 1995. Summary of Pacific salmon coded-wire tag application and recovery, Prince William, 1995. Regional Information Report No. 2A95-51. Alaska Dept. of Fish and Game. Anchorage, Alaska. Kline, T.C., Jr. 1996. Sound Ecosystem Assessment: confirming food web dependencies in the Prince William Sound Ecosystem using stable isotope tracers (SEA-FOOD). Exxon Valdez Restoration Project Annual Report (Restoration Project 95320I), Alaska Dept. of Fish and Game, Anchorage, Alaska. Kline, T.C., Jr. 1999. Temporal and spatial variability of 13C/12C and 15N/14N in pelagic biota of Prince William Sound, Alaska. Can. J. Fish. Aquat. Sci. 56(Suppl. 1): 94117. Kline, T.C., Jr. 2001a. Evidence of biophysical coupling from shifts in natural stable carbon and nitrogen isotopes in Prince William Sound, Alaska. In Spatial processes and management of marine populations. Edited by G.H Kruse,. N. Bez, A. Booth, M.W. Dorn, S. Hills, R.N. Lipcius, D. Pelletier, C. Roy, S.J. Smith, and D. Witherell. University of Alaska Sea Grant, AK-SG-01-02, Fairbanks. pp. 363375. Kline, T.C., Jr. 2001b. The trophic position of Pacific herring in Prince William Sound, Alaska based on their stable isotope abundance. In Herring: expectations for a new millennium. Edited by F. Funk, J. Blackburn, D. Hay, A.J. Paul, R. Stephenson, R. Toresen, and D. Witherell. University of Alaska Sea Grant, AK-SG-01-04, Fairbanks. pp. 6980. Kline, T.C., Jr., and Pauly, D. 1998. Cross-validation of trophic level estimates from a mass-balance model of Prince William Sound using 15N/14N data. In Fishery stock assessment models. Edited by F. Funk, T.J. Quinn, II, J. Heifetz, J.N. Ianelli, J.E. Powers, J.F. Schweigert, P.J. Sullivan, and C.-I. Zhang. Alaska Sea Grant College Program Report No. AK-SG-98-01, University of Alaska Fairbanks, 1998. pp. 693702. Kline, T.C., Jr., Goering, J.J., Mathisen, O.A., Poe, P.H., Parker, P.L., and Scalan, R.S. 1993. Recycling of elements transported upstream by runs of Pacific salmon: II. 15N and 13C evidence in the Kvichak River Watershed, Bristol Bay, southwestern Alaska. Can. J. Fish. Aquat. Sci. 50: 23502365. McConnaughey, T., and McRoy, C.P. 1979. Food-web structure and the fractionation of carbon isotopes in the Bering Sea. Mar. Biol. 53: 257262. Michener, R.H., and Schell, D.M. 1994. Stable isotope ratios as tracers in marine food webs. In Stable isotopes in ecology and environmental science. Edited by K. Lajtha and R.H. Michener. Blackwell Scientific Publications, Oxford, U.K. pp. 138157. Oakey, T.A., and Pauly, D. (Editors). 1998. Trophic mass-balance model of Alaskas Prince William Sound Ecosystem, for the post-spill period 19941996. Fisheries Centre Research Reports 6(4). University of British Columbia, Vancouver. Paul, A.J., and Willette, M. 1997. Geographical variation in somatic energy content of migrating pink salmon from prince William Sound: a tool to measure nutritional status. In Forage fishes

Can. J. Fish. Aquat. Sci. Vol. 59, 2002 in marine ecosystems. Proceedings of the international symposium on the role of forage fishes in marine ecosystems, 1316 Nov. 1996, Anchorage, Alaska. Alaska Sea Grant College Program Report No. 9701. University of Alaska Fairbanks. pp. 707720. Paul, A.J., Paul, J.M., and Kline, T.C. 2001. Estimating whole body energy content for juvenile Pacific herring from condition factor, dry weight, and carbon/nitrogen ratio. In Herring: expectations for a new millennium. Edited by F. Funk, J. Blackburn, D. Hay, A.J. Paul, R. Stephenson, R. Toresen, and D. Witherell. University of Alaska Sea Grant, AK-SG-01-04, Fairbanks. pp. 121133. Parker, R.R. 1968. Marine mortality schedules of pink salmon of the Bella Coola River, central British Columbia. J. Fish. Res. Board Can. 25: 757794. Parker, R.R. 1971. Size selective predation among juvenile salmonid fishes in a British Columbia inlet. J. Fish. Res. Board Can. 28: 15031510. Perry, R.I., Hargreaves, N.B., Waddell, B.J., and Mackas, D.L. 1996. Spatial variations in feeding and condition of juvenile pink and chum salmon off Vancouver Island, British Columbia. Fish. Oceanogr. 5: 7388. Schell, D.M. 2001. Carbon isotope ratio variations in Bering Sea biota: The role of anthropogenic carbon dioxide. Limnol. Oceanogr. 46: 9991000. Vander Zanden, M.J., and Rasmussen, J.B. 2001. Variation in 15N and 13C trophic fractionation: implications for aquatic food web studies. Limnol. Oceanogr. 46: 20612066. Vaughan, S.L., Mooers, C.N.K., and Gay, S.M., III. 2001. Physical variability in Prince William Sound during the SEA study (19941998). Fish. Oceanogr. 10(Suppl. 1): 1441. Wang, J., Jin, M., Patrick, E.V., Allen, J.R., Eslinger, D.L., Mooers, C.N.K., and Cooney, R.T. 2001. Numerical simulations of the seasonal circulation patterns and thermohaline structures of Prince William Sound, Alaska. Fish. Oceanogr. 10(Suppl. 1): 1441. West, C.J., and Larkin, P.A. 1987. Evidence of size-selective mortality of juvenile sockeye salmon (Oncorhynchus nerka) in Babine Lake, British Columbia. Can. J. Fish. Aquat. Sci. 44: 712721. Willette, T.M. 2001. Foraging behaviour of juvenile pink salmon (Oncorhynchus gorbuscha) and size-dependent predation risk. Fish. Oceanogr. 10(Suppl. 1): 110131. Willette, T.M., Carpenter, G., and Debevec, E. 1995. Sound ecosystem assessment: salmon growth and mortality. Restoration Project 94320A Final Report, Alaska Dept. Fish and Game, Cordova, Alaska. Willette, T.M., Carpenter, G., Saddler, P., and Powell, M. 1996. Sound ecosystem assessment: Salmon growth and mortality. Exxon Valdez Restoration Project Annual Report (Restoration Project 95320A), Alaska Dept. Fish and Game, Cordova, Alaska. Willette, T.M., Cooney, R.T., and Hyer, K. 1999a. Predator foraging mode shifts affecting mortality of juvenile fishes during the subarctic spring bloom. Can. J. Fish. Aquat. Sci. 56: 364376. Willette, T.M., Cooney, T., and Hyer, K. 1999b. Some processes affecting mortality of juvenile fishes during the spring bloom in Prince William Sound, Alaska. In Ecosystem approaches in fisheries management. University of Alaska Sea Grant, AK-SG-99-01, Fairbanks. pp. 137142. Willette, T.M., Cooney, R.T., Patrick, V., Mason, D.M., Thomas, G.L., and Scheel, D. 2001. Ecological processes influencing mortality of juvenile pink salmon (Oncorhynchus gorbuscha) in Prince William Sound, Alaska. Fish. Oceanogr. 10(Suppl. 1): 1441.

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Appendix A
Table A1. Sample inventory, ordered by hatchery release date, indicating total number (N) of samples from each releasecapture date combination for each species and hatchery; same information from pilot study in right-most column. Hatchery WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN WHN AFK AFK AFK AFK MB MB MB MB MB MB MB MB WHN WHN WHN WHN WHN WHN MB MB MB MB WHN WHN WHN WHN WHN WHN WHN WHN WHN Salmon species Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Sockeye Sockeye Sockeye Sockeye Sockeye Sockeye Sockeye Sockeye Chum Chum Chum Chum Chum Chum Sockeye Sockeye Sockeye Sockeye Chum Chum Chum Chum Chum Chum Chum Pink Pink Release date 25 April 25 April 25 April 25 April 25 April 25 April 25 April 25 April 25 April 25 April 25 April 25 April 25 April 25 April 25 April 7 May 7 May 7 May 7 May 7 May 7 May 7 May 7 May 10 May 10 May 17 May 17 May 28 May 28 May 28 May 28 May 28 May 28 May 29 May 29 May 30 May 30 May 30 May 30 May 30 May 30 May 30 May 30 May 31 May 31 May 3 June 3 June 3 June 3 June 3 June 3 June 3 June 11 June 11 June Capture date 8 May 9 May 8 June 9 June 10 June 11 June 12 June 13 June 14 June 27 June 28 June 1 July 10 Jul 11 July 16 July 8 May 11 May 25 May 2 June 27 June 28 June 29 June 10 July 7 July 8 July 7 July 8 July 24 June 25 June 26 June 1 July 9 July 12 July 25 June 9 July 4 June 5 June 13 June 14 July 15 July 16 July 26 June 12 July 4 June 25 June 4 June 5 June 11 July 13 July 14 July 15 July 16 July 25 June 26 June Days since release 13 14 44 45 46 47 48 49 50 63 64 67 76 77 82 1 4 18 26 51 52 53 64 58 59 51 52 27 28 29 34 42 45 27 41 5 6 14 45 46 47 27 43 4 25 1 2 38 40 41 42 43 14 15 N total 22 1 1 2 2 6 1 1 1 2 4 1 1 2 1 4 2 8 5 13 5 1 1 1 1 2 2 1 1 3 1 2 1 1 1 4 4 1 1 2 1 1 1 1 1 23 16 1 2 1 2 1 1 3 N pilot study 5

1 2

1 1 2 2

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1638 Table A1 (concluded). Hatchery WHN WHN WHN WHN WHN WHN WHN WHN AFK AFK Salmon species Pink Pink Pink Pink Pink Pink Pink Pink Pink Pink Release date 11 June 11 June 11 June 11 June 11 June 11 June 11 June 11 June 13 June 13 June Capture date 27 June 28 June 29 June 30 June 1 July 10 July 11 July 12 July 7 July 8 July

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Days since release 16 17 18 19 20 29 30 31 24 25

N total 5 3 11 3 1 6 5 5 21 20

N pilot study

21 20

Note: Hatchery abbreviations are as follows: WHN, W.H. Norenberg; MB, Main Bay; AFK, A.F. Koernig. Dates are from 1994.

Table A2. Regression parameters for Figs. 2 to 5.


Fig. No. 2a 2a 2a 2a 2a 2b 2b 2b 2b 2b 2c 2c 2c 2c 2c 3a 3a 3a 3a 3a 3b 3b 3b 3b 3b 4a 4a 4a 4a 4a 4b 4b 4b 4b 4b 5 5 5 5 5 Cohort April WHN pink May WHN pink June WHN pink WHN chum MB sockeye April WHN pink May WHN pink June WHN pink WHN chum MB sockeye April WHN pink May WHN pink June WHN pink Chum WHN Sockeye MB April WHN pink May WHN pink June WHN pink WHN chum MB sockeye April WHN pink May WHN pink June WHN pink WHN chum MB sockeye April WHN pink May WHN pink June WHN pink WHN chum MB sockeye April WHN pink May WHN pink June WHN pink WHN chum MB sockeye April WHN pink May WHN pink June WHN pink WHN chum MB sockeye Model N (mg) = Ae N (mg) = AeB(d) N (mg) = AeB(d) N (mg) = AeB(d) N (mg) = AeB(d) C (mg) = AeB(d) C (mg) = AeB(d) C (mg) = AeB(d) C (mg) = AeB(d) C (mg) = AeB(d) C/N (atom ratio) = AeB(d) C/N (atom ratio) = AeB(d) C/N (atom ratio) = AeB(d) C/N (atom ratio) = AeB(d) C/N (atom ratio) = AeB(d) 15N = AeB(d) + C 15N = AeB(d) + C 15N = AeB(d) + C 15N = AeB(d) + C 15N = AeB(d) + C 13C = A(d) B 13C = A(d) B 13C = A(d) B 13C = A(d) B 13C = A(d) B 15N = A M/(B + M) 15N = A M/(B + M) 15N = A M/(B + M) 15N = A M/(B + M) 15N = A M/(B + M) 15N = A + B(C/N) 15N = A + B(C/N) 15N = A + B(C/N) 15N = A + B(C/N) 15N = A + B(C/N) 13C = A + B(C/N) 13C = A + B(C/N) 13C = A + B(C/N) 13C = A + B(C/N) 13C = A + B(C/N)
B(d)

Independent variable d d d d d d d d d d d d d d d d d d d d d d d d d M M M M M C/N C/N C/N C/N C/N C/N C/N C/N C/N C/N

A 10.76 10.50 37.18 19.74 318 42.85 42.89 127.8 74.54 1126 2.27 1.26 6.31E+31 0.80 1.76 3.348 2.99 72.05 1.669 2.195 0.009 0.027 0.038 0.016 0.025 0.1412 0.05961 0.07961 0.1223 0.1947 5.50 6.49 9.84 9.10 6.76 21.08 23.77 16.41 18.24 24.96

B 0.043 0.048 0.054 0.063 0.014 0.041 0.044 0.054 0.060 0.014 0.057 0.029 5.438 0.083 0.353 0.473 0.040 6.10E-06 0.069 0.055 19.38 19.28 21.2 19.31 18.66 0.095 0.066 0.702 0.183 5.127 1.350 0.998 0.263 0.703 1.259 0.301 0.7748 0.9927 0.2738 1.352

R2 0.83 0.86 0.47 0.96 0.20 0.81 0.85 0.46 0.96 0.18 0.88 0.25 0.06 0.40 0.79 0.79 0.86 0.00 0.76 0.55 0.18 0.48 0.43 0.12 0.31 0.20 0.90 0.03 0.39 0.35 0.71 0.63 0.01 0.23 0.40 0.11 0.49 0.02 0.70 0.32

T2 16.3 14.5 12.8 11.0 49.6 17.1 15.7 12.9 11.5 51.3 12.2 23.7 0.1 3.9 4.2 14.7 17.2 113561 10.0 12.6

CI ratio 1.6 1.6 2.1 1.2 undefined 1.6 1.6 2.1 1.2 undefined 1.9 9.8 1.1 26.6 undefined 73.6 2.7 undefined 6.3 undefined

+ + + + +

4 4 4 4 4

10.7 10.1 82.9 11.1 11.4

Note: Relative goodness-of-fit for nonlinear models is indicated by bold for high correlation coefficient while having a half-time (T2) confidence interval ratio (CI ratio) <7. Significant (P < 0.05) linear models indicated by bold. Hatchery abbreviations: WHN, W.H. Norenberg; MB, Main Bay; AFK, A.F. Koernig. d, days since release; M, mass of nitrogen in milligrams; C/N, atom ratio. The models come out of the datavia computer fit, they were not a priori.

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