Stillaguamish Summer Chinook:

Productivity Estimates from Coded-Wire Tag Recoveries and

A Simple Model for Setting Interim Exploitation Rate Objectives

 

 

 

Kit Rawson

Tulalip Fisheries

7615 Totem Beach Rd.

Marysville, WA 98271

Krawson@tulalip.nsn.us

 

May 17, 2000

 

Introduction

 

The joint state tribal Comprehensive Chinook working group (COOKS[1] draft plan, November 28, 1999) has established the following objective for management of the harvest of Puget Sound chinook salmon stocks:

 

Manage fisheries to ensure that harvest related mortality will not impede recovery of the productivity, abundance, and diversity of Puget Sound chinook salmon stocks to levels consistent with treaty-reserved fishing rights and salmon –related cultural and ecological values.

 

The Stillaguamish chinook management unit is comprised of two stocks (WDF et al. 1993): Stillaguamish summer chinook and Stillaguamish fall chinook.  Consistent with the COOKS draft plan, the management guidelines include an interim exploitation rate objective and a low abundance threshold.  The interim exploitation rate objective is the maximum fraction of the production from any brood year that is allowed to be removed by all sources of fishery-related mortality, including direct take, incidental take, and non-landed mortality.  The exploitation rate is expressed as an adult equivalent rate (see below for definition).  The low abundance threshold is the minimum natural spawning escapement allowable in any return year.  Whenever spawning escapement is projected to be below this level, fisheries will be managed to a very low exploitation rate (to be described in the COOKS plan in terms of specific fishery restrictions), lower than the interim exploitation rate objective.

 

In this report I will document analysis of available data pertaining to the Stillaguamish summer chinook stock and a simple model that can be used to project the consequences of employing different maximum exploitation rates.  Using this model, managers can select an interim rate to use in management.  The exploitation rate objective is termed “interim” because the comanagers expect and intend to collect new information pertaining to the current productivity, capacity, and diversity of the chinook salmon stocks.  The ultimate management plan will be based on analysis of this new data.

 

The interim exploitation rate objective is developed from the most recently available information concerning productivity and capacity.  For Stillaguamish chinook, there are 8 complete brood years of coded-wire tag (CWT) recoveries from the Stillaguamish Tribe’s indicator stock project.  From recoveries of these tags, it is possible to estimate the total numbers of recruits produced   from each brood year 1986-1993. Using spawner escapement estimates for those years, it is possible to estimate recruit per spawner values.  I used the 8 estimates of recruits per spawner to simulate future patterns of production and harvest under adult equivalent exploitation rates, varying from zero to .80 in intervals of .05.

 

Methods

 

Coded-wire tag codes used.  I used summarized and expanded recoveries of coded-wire tags from the Stillaguamish indicator stock project for brood years 1986-1993 to estimate actual exploitation rates.  Because this project involved taking broodstock from the summer run fish in the North Fork Stillaguamish, this analysis is specific to the summer chinook stock.  All escapement numbers, reconstructed run sizes, and estimates of recruits per spawner, are for the Stillaguamish Summer chinook stock only.  However, because of a current lack of data for the Stillaguamish fall chinook stock, the maximum exploitation rate objective recommended for the summer chinook stock may be applied to the fall chinook stock as well. Tag codes and release numbers are in Table 1.

 

Table 1. Tag codes used in reconstruction of Stillaguamish chinook exploitation rates.

Brood Yr.

Tag Code

Tags released

AD-Only

Unmarked

1986

21-22-21

  23,904

996

0

1987

21-25-55

127,910

9,333

7,923

1988

21-31-47

36,599

4,524

0

1989

21-18-26

44,964

1,873

0

1990

21-20-26

63,019

5,852

219

1991

21-22-05

165,620

5,303

5,833

1992

21-22-40

24,091

771

848

1992

21-22-51

89,207

5,708

5,206

1993

21-23-30

200,664

6,912

8,424

1993

21-20-45

3,855

133

162

1993

21-18-56

5,048

174

212

 

 

I obtained estimated CWTs by age for each tag code from the CRAS database management system[2].  I used the estimated contribution of CWTs to the fisheries directly from the CRAS reports and estimated the escapement of tagged fish to spawning escapement based on information on spawning escapement estimates and sampling rates assembled by Will Beattie (personal communication, 1999).  Estimated CWTs for each brood year are shown in Table 2.  Estimated CWTs were summed over all codes for the brood year for brood years with more than one tag code.

 

Table 2. Estimated contribution of Stillaguamish CWTs to fisheries and escapement for each brood year.  The Nominal exploitation rate (ER) in the righthand column is simply the sum of the estimated fishery CWTs divided by the sum of fishery and escapement CWTs.

Brood

Est. Fishery CWTsa

Est. Escapement CWTsb

Nominal

Year

Age 2

Age 3

Age 4

Age 5

Age 2

Age 3

Age 4

Age 5

Total

ER

1986

79.5

157.7

32.1

 

 

 

82.9

22.2

374.5

0.719

1987

131.7

256.9

77.8

1.0

 

40.2

395.8

 

903.4

0.517

1988

71.7

145.7

83.8

 

 

104.0

37.4

2.3

444.9

0.677

1989

75.7

259.9

97.2

 

 

28.9

50.0

 

511.7

0.846

1990

286.2

481.8

116.0

4.0

 

160.0

196.4

3.2

1247.7

0.712

1991

3.1

42.3

44.3

 

 

23.9

43.7

3.6

160.9

0.558

1992

11.2

72.1

35.1

 

7.8

67.9

87.3

10.7

292.2

0.405

1993

140.8

284.8

94.1

 

62.9

172.0

141.5

43.6

939.7

0.553

 

 

 

 

 

 

 

 

 

 

 

aFrom CRAS summary output.

bFrom information supplied by W. Beattie, see Appendix A.

 

Exploitation Rates.  The total of the estimated fishery CWTs divided by the total fishery CWTs plus the escapement CWTs is the nominal exploitation rate for the brood year.  To make exploitation rates comparable among brood years when the distribution of fishing mortality among ages may have been diffrent, it is necessary to adjust fishery recoveries by adult equivalency (AEQ) factors[3].  I obtained AEQ factors for these tag codes form the Pacific Salmon Commission’s Chinook Technical Committee (M. Alexandersdottir, Northwest Indian Fisheries Commission, personal communication).  Applying these to the fishery recoveries, I computed AEQ exploitation rates as the sum of the AEQ-adjusted fishery CWT contribution, divided by this sum plus the sum of the estimated escapement CWT contribution.  These values are given in Table 3.

 

Table 3. Computation of AEQ exploitation rates from recoveries of Stillaguamish coded-wire tags.

Brood

AEQ Fishery CWTsa

Fshry

Escp.

AEQ

 

Year

Age 2

Age 3

Age 4

Age 5

Total

Total

ER

 

1986

48.1

129.1

31.3

 

208.5

105.2

0.665

 

1987

77.7

209.6

77.6

1.0

365.8

436

0.456

 

1988

45.9

129.4

83.3

 

258.7

143.7

0.643

 

1989

44.9

216.9

97.0

 

358.9

78.9

0.820

 

1990

179.5

415.6

115.2

4.0

714.4

359.7

0.665

 

1991

1.8

35.4

44.0

 

81.2

71.2

0.533

 

1992

7.0

64.3

34.7

 

106.0

173.8

0.379

 

1993

84.2

238.8

93.4

 

416.3

420.0

0.498

 

 

 

 

 

 

 

 

 

 

aComputed by applying AEQ factors from the PSC chinook technical committee to the estimated tag recoveries by age (Table 2).

 

Reconstruction of Total Recruits and Recruits per Spawner.  Based on the above, I reconstructed the total brood year run size by expanding the spawning escapement by the estimated AEQ exploitation rate.  Because the Stillaguamish Tribe’s indicator stock project includes acquisition of broodstock from the escapement, there are four categories of spawning escapement that must be considered: 1) natural origin fish spawning in natural areas, 2) natural origin fish taken for broodstock, 3) fish originating from the indicator stock program spawning in natural areas, and 4) fish from the indicator stock program taken as broodstock.  I separated the escapement into these categories based on age and mark data from the Stillaguamish Tribe (J. Drotts, personal communication, 1999).  The spawning escapement that was expanded to reconstruct total recruitment included all natural origin fish (categories 1) and 2) ).  In order to estimate recruits per spawner I divided this total recruitment by the naturally spawning fish (categories 1) and 3) ) in the brood year.  These computations are summarized in Table 4.

 

Table 4. Reconstruction of total recruitment and recruits per spawner.

Brood

Year

Original

Escapementa

Subsequent

Escapementb

Brood Yr

Exp. Rate

 

Recruitment

Recruits

Per Spawner

 

1986

980

505

0.66

1,505

1.54

 

1987

1,065

695

0.46

1,278

1.20

 

1988

516

654

0.64

1,832

3.55

 

1989

510

458

0.82

2,544

4.99

 

1990

575

488

0.67

1,457

2.53

 

1991

1,331

486

0.53

1,040

0.78

 

1992

466

596

0.38

959

2.06

 

1993

563

585

0.50

1,165

2.07

 

 

 

 

 

 

 

aIncludes all fish spawning naturally in the brood year.

bIncludes all fish produced by the naturally spawners from the brood year which returned as escapement to the river (natural origin recruits).

 

Marine and Freshwater survival indices.  The recruits per spawner values in Table 4 are the product of a component representing survival in freshwater and marine survival.  Jim Scott (Washington Department of Fish and Wildlife, personal communication, November 1999) has developed a set of marine survival indices for Puget Sound chinook salmon from commonalties in recovery patterns of coded-wire tagged chinook from hatcheries throughout Puget Sound.  These are shown in Table 5, along with a Stillaguamish freshwater index, which is simply the recruit per spawner value from Table 4 divided by the marine survival index value for the particular brood year.

 


Table 5. Puget Sound marine survival indices and Stillaguamish freshwater indices derived from recruits per spawner and marine survival indices.  For each brood year, the recruits per spawner is the product of these indices.

Brood

Year

Mar. Surv.

Indexa 

Freshwater

Indexb

1986

2.35

.655

1987

.74

1.622

1988

.78

4.551

1989

.80

6.238

1990

1.66

1.524

1991

.29

2.690

1992

.37

5.568

1993

.68

3.044

 

 

 

aFrom Jim Scott, Wash. Dept. Fish and Wildlife.

bRecruits per spawner (Table 4) divided by marine survival index.

 

The Simulation Model and Results

 

In order to compare the effects of different exploitation rate strategies, I developed a simple simulation model using the above information. For each year simulated, the model selects a freshwater index and marine survival index at random from the values in Table 5.  The first stage in computing total recruitment for the brood year is to multiply brood year escapement times the chosen freshwater index.  If the resulting product exceeds a maximum value (5,000 in these simulations[4]) then the value of recruitment at this stage is reduced to that maximum .  This is a simple spawner-recruit relationship that does not include any density dependence, other than the maximum value of total recruitment[5].  The first stage recruits are then multiplied by a marine survival index to obtain the total brood year recruitment.  Each year’s marine survival index is the average of the randomly selected index and the index used for the previous year.  This averaging mimics some degree of autocorrelation in marine survivals.

 

The brood year recruitment is then subjected to fishing mortality, based on the target exploitation rate chosen for the run. For each year of each run, the exploitation rate was adjusted by a management error rate randomly chosen from 36 values computed for the 1988 through 1993 fishing years by comparing expected with actual exploitation rates for six Puget Sound chinook management units[6].  Surviving fish return to spawning escapement according to the average escapement age distribution observed for the 1986-1993 brood years[7].

 

Each model run projects the return and the escapement for 25 years. I replicated 1000 model runs at each exploitation rate ranging from 0 to .80 in increments of .05 (increments of 0.1 above 0.50).  All model runs were seeded with the observed North Fork Stillaguamish natural chinook escapement values for the first four years of the run (years 0 through 3). 

 

Examples of individual runs of this model at four different exploitation rates are shown in Appendix C.

 

For each target exploitation rate I computed the average percentage of years that the escapement was below a critical threshold (500 spawners in these runs)[8].  I also averaged the spawning escapements in years 0-3 and 22-25.  To compare rebuilding rates, I computed the ratio of years 22-25 escapement to years 0-3 escapement.  These quantities are tabulated (Table 6) below.

 

Table 6. Summary of results of 1000 runs at each exploitation rate.

Exploitation

Percent

Percent

Median

Median

Rate

< crit.

Escap. ratio>1

Escap. ratio

Escapement

0.00

1%

96%

        2.75

          3,597

0.05

1%

96%

        2.81

          3,377

0.10

1%

96%

        2.76

          3,165

0.15

2%

95%

        2.66

          2,964

0.20

2%

95%

        2.56

          2,758

0.25

3%

93%

        2.57

          2,418

0.30

4%

92%

        2.48

          2,210

0.35

6%

92%

        2.46

          1,920

0.40

7%

91%

        2.29

          1,686

0.45

11%

87%

        2.14

          1,444

0.50

17%

80%

        1.92

          1,180

0.60

41%

52%

        1.04

             648

0.70

73%

12%

        0.27

             259

0.80

94%

0%

        0.02

               55

 

 

Conclusion

 

The results from these simulations can be compared with criteria for evaluating fishery management plans under consideration by NMFS.  The first test is whether the proposed action increases the probability of the spawning escapement dropping below a threshold level by more than 5 percentage points, when compared with no action.  The comanagers have chosen a threshold of 500 spawners for the Stillaguamish summer chinook stock based on historical spawning escapement data showing that in 6 of 30 years (1969-1998) the estimated escapement for this stock was below this value.  Because of reduced fishing mortality, escapements below 500 have only occurred twice in the last 15 years (1984 and 1992).  Until detailed information on capacity and productivity is assembled, the comanagers have chosen to use 500 as the critical threshold in the interim management plan and I used the same value as the critical escapement level to test in these simulations.  Using the NMFS criterion and the comanagers’ critical escapement value of 500, exploitation rates up to 0.35 would be acceptable based on the results of these simulations (Table 6, comparing the percentages of runs with escapements < critical for exploitation rates of 0 and 0.35).

 

A second criterion proposed by NMFS is to test whether the probability of rebuilding under a proposed action is 80% or greater.  For these simulations, I defined rebuilding as occurring when the escapement ratio (years 22-25 escapement divided by years 0-3 escapement) is greater than 1.  Using this definition and the NMFS criterion, exploitation rates of up to 0.50 would be acceptable (Table 6).

 

Using the most restrictive of the two criteria, target exploitation rates of up to 0.35 would be acceptable for the Stillaguamish summer chinook stock.  Other factors, not included in the simulation analysis, might dictate a maximum rate lower than 0.35.  The most important of these is that the exploitation rate chosen will be applied to the entire Stillaguamish management unit, which includes both the Stillaguamish summer and Stillaguamish fall stocks.  Therefore, the chosen rate must be appropriate for both stocks. 

 

At the present time, there is very little information concerning the productivity of the Stillaguamish fall stock other than the fact that the average abundance of this stock is approximately 50% of the Stillaguamish summer stock (Appendix A, Table A.1, comparing percent North Fork escapement with estimated total escapement).  I ran the model described above, with initial escapements and maximum recruitment reduced to 50% of the values used for the Stillaguamish summer stock and the critical escapement threshold set at 250 spawners.  These resulted in an exploitation rate of 0.35 showing a probability of 4% of escapements below critical, as compared with a 1% probability for an exploitation rate of 0.  The probability of rebuilding at this exploitation rate was 96%.  This rough analysis indicates that a target exploitation rate of 0.35 would also be appropriate for the Stillaguamish fall stock.

 

The comanagers have selected an exploitation rate guideline of 0.30 for the Stillaguamish chinook management unit and a critical threshold of 500 natural origin spawners for the Stillaguamish summer chinook stock.  According to the above analysis, these guidelines should meet the NMFS criteria for an acceptable harvest management strategy for bot the Stillaguamish summer and Stillaguamish fall stocks.

 

References Cited

 

Morishima, Gary. 1999. When is a fish not a fish? Metrics of exploitation for chinook salmon.  Briefing paper prepared for WDFW/NWIFC modeling workshop. Quinault Management Center, Mercer Island, WA, October 1999.

 

Washington Department of Fisheries (WDF), Washington Department of Wildlife (WDW), and Western Washington Treaty Indian Tribes. 1993.  1992 Washington State salmon and steelhead stock inventory.  212 p. (Available from the Washington Department of Fish and Wildlife, Olympia, WA.)


Appendix A.  In-river tag recovery information (assembled by W. Beattie, NWIFC)

 

 

A1.  Escapement estimates and spawning ground CWT sampling summary

 

 

 

 

 

 

 

 

total

Est.

CWT

 

Est. CWT

 

 

Natural esc.

# Carcass

# CWT's

Pct.

exp.

NF % of

in

 

 

Year

N. Fork

Total

sampled

Recov'd

Natrl.

factor

total esc

Escp.

 

 

1988

516

717

0

 

 

 

72.0%

 

 

 

1989

510

811

72

 

 

7.08

62.9%

 

 

 

1990

575

842

65

 

 

8.85

68.3%

 

 

 

1991

1331

1632

187

 

 

7.12

81.6%

 

 

 

1992

466

780

84

11

86.9%

5.55

59.7%

61.0

 

 

1993

563

928

270

94

65.2%

2.09

60.7%

196.0

 

 

1994

659

954

206

65

68.4%

3.20

69.1%

207.9

 

 

1995

561

822

192

54

71.9%

2.92

68.2%

157.8

 

 

1996

918

1384

270

84

68.9%

3.40

66.3%

285.6

 

 

1997

771

1156

88

30

65.9%

8.76

66.7%

262.8

 

 

1998

955

1540

212

111

47.6%

4.50

62.0%

500.0

 

 

Average

 

 

 

 

 

 

67.0%   

 

 

 

 

 

 

 

 

 

 

 

 

 

 

A.2  Detail of observed and estimated Stillaguamish indicator stock CWT recoveries from stream surveys

 

 

 

 

 

 

 

 

 

 

 

 

Recovery Year

Brood Yr

1989

1990

1991

1992

1993

1994

1995

1996

1997

1998

1985

 

 

 

 

 

 

 

 

 

 

1986

0

9

3

 

 

 

 

 

 

 

1987

0

4

49

 

 

 

 

 

 

 

1988

 

 

13

6

1

 

 

 

 

 

1989

 

 

 

5

23

 

 

 

 

 

1990

 

 

 

 

70

56

1

 

 

 

1991

 

 

 

 

 

7

14

1

 

 

1992

 

 

 

 

 

2

19

21

1

 

1993

 

 

 

 

 

 

20

47

15

9

1994

 

 

 

 

 

 

 

15

11

83

1995

 

 

 

 

 

 

 

 

 

8

1996

 

 

 

 

 

 

 

 

 

11

total

0

13

65

11

94

65

54

84

27

111

 

 

 

 

 

 

 

 

 

 

 


 

A.3 Estimated recoveries expanded for sampling rate  (total carcasses sampled / estimate of escapement)

 

 

Recovery Year

 

Brood Yr

1990

1991

1992

1993

1994

1995

1996

1997

1998

 

 

 

 

 

 

 

 

 

 

 

 

1986

79.6

21.4

 

 

 

 

 

 

 

 

1987

35.4

348.8

 

 

 

 

 

 

 

 

1988

 

92.5

33.3

2.1

 

 

 

 

 

 

1989

 

 

27.7

48.0

 

 

 

 

 

 

1990

 

 

 

146.0

179.1

2.9

 

 

 

 

1991

 

 

 

 

22.4

40.9

3.4

 

 

 

1992

 

 

 

 

6.4

55.5

71.4

8.8

 

 

1993

 

 

 

 

 

58.4

159.8

131.4

40.5

 

1994

 

 

 

 

 

 

51.0

96.4

373.9

 

1995

 

 

 

 

 

 

 

 

36.0

 

1996

 

 

 

 

 

 

 

 

49.6

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

A.4. Escapement recoveries expanded for tagging rate at release

 

 

 

 

 

 

Recovery Year

 

Brood Yr

1990

1991

1992

1993

1994

1995

1996

1997

1998

 

 

 

 

 

 

 

 

 

 

 

 

1986

82.9

22.2

 

 

 

 

 

 

 

 

1987

40.2

395.8

 

 

 

 

 

 

 

 

1988

 

104.0

37.4

2.3

 

 

 

 

 

 

1989

 

 

28.9

50.0

 

 

 

 

 

 

1990

 

 

 

160.0

196.4

3.2

 

 

 

 

1991

 

 

 

 

23.9

43.7

3.6

 

 

 

1992

 

 

 

 

7.8

67.9

87.3

10.7

 

 

1993

 

 

 

 

 

62.9

172.0

141.5

43.6

 

1994

 

 

 

 

 

 

53.0

100.2

388.7

 

1995

 

 

 

 

 

 

 

 

43.6

 

1996

 

 

 

 

 

 

 

 

106.9

 

 

 

 

 

 

 

 

 

 

 

 

Total

123.1

522.0

66.3

212.3

228.2

177.7

316.0

252.4

582.9

 

% of NF

0.214

0.392

0.142

0.377

0.346

0.317

0.344

0.327

0.610

 

 


 

Appendix B. Management errors based on comparison of post-season exploitation rates with preseason predictions.  All values, except those for White River spring chinook from J. Gutmann’s memo (24 Feb 1998) were used in the simulations.  White River values were rejected because they were significantly different from the remaining values in the array (1-tailed t-test, see p-values on right).

 

 

 

 

 

 

Management

Percent

 

 

Management Unit

Type

Year

Error

 

 

LAKE WASHINGTON

S/F

1988

-0.116

t-test p-values

LAKE WASHINGTON

S/F

1989

-0.033

0.000426

WR vs S/F

LAKE WASHINGTON

S/F

1990

0.012

5.41E-05

WR vs rest

LAKE WASHINGTON

S/F

1991

-0.144

0.078335

SP vs S/F

LAKE WASHINGTON

S/F

1992

-0.085

 

 

LAKE WASHINGTON

S/F

1993

0.213

Averages

 

SKAGIT

S/F

1988

0.226

Lk WA

-0.02550

SKAGIT

S/F

1989

-0.101

Skagit S/F

-0.02217

SKAGIT

S/F

1990

0.126

Sno S/F

-0.00750

SKAGIT

S/F

1991

-0.071

Stil S/F

0.03900

SKAGIT

S/F

1992

-0.127

Nook SP

-0.02467

SKAGIT

S/F

1993

-0.186

Skagit SP

-0.02083

SNOHOMISH

S/F

1988

0.052

WR SP

0.26283

SNOHOMISH

S/F

1989

0.130

 

 

SNOHOMISH

S/F

1990

0.162

S/F

-0.004042

SNOHOMISH

S/F

1991

-0.134

non-WR

-0.010278

SNOHOMISH

S/F

1992

-0.182

SP

0.0724444

SNOHOMISH

S/F

1993

-0.073

 

 

STILLAGUAMISH

S/F

1988

-0.009

1988

0.06286

STILLAGUAMISH

S/F

1989

0.117

1989

0.04714

STILLAGUAMISH

S/F

1990

-0.252

1990

0.07257

STILLAGUAMISH

S/F

1991

0.067

1991

-0.04557

STILLAGUAMISH

S/F

1992

0.008

1992

-0.07986

STILLAGUAMISH

S/F

1993

0.303

1993

0.11529

NOOKSACK

SP

1988

0.009

1994

0.09267

NOOKSACK

SP

1989

0.049

 

 

NOOKSACK

SP

1990

-0.044

All

0.02874

NOOKSACK

SP

1991

-0.173

 

 

NOOKSACK

SP

1992

-0.028

 

 

NOOKSACK

SP

1993

0.039

 

 

SKAGIT

SP

1988

0.081

 

 

SKAGIT

SP

1989

-0.177

 

 

SKAGIT

SP

1990

-0.006

 

 

SKAGIT

SP

1991

-0.162

 

 

SKAGIT

SP

1992

-0.069

 

 

SKAGIT

SP

1993

0.208

 

 

WHITE RIVER

SP

1988

0.197

 

 

WHITE RIVER

SP

1989

0.345

 

 

WHITE RIVER

SP

1990

0.51

 

 

WHITE RIVER

SP

1991

0.298

 

 

WHITE RIVER

SP

1992

-0.076

 

 

WHITE RIVER

SP

1993

0.303

 

 

 


 

 

 

 

 

 

 


Appendix C.  Examples of individual simulations at exploitation rates of 0.0, 0.05, 0.30, and 0.50.

 


Exploitation Rate = 0

 

 

 

 



[1] The Committee to Organize Options for King Salmon (COOKS) is developing a harvest management plan for Puget Sound chinook salmon to be completed in early 2000.

[2] The Coded-wire tag Retrieval and Analysis System is maintained by the Northwest Indian Fisheries Commission in Olympia, WA (www.nwifc.wa.gov).  The tag recovery data are compiled and stored by the Pacific States Marine Fisheries Commission in Portland, OR.

[3] “The adult equivalence factor is an age- and stock-specific scalar that converts nominal fish into potential adults.” (Morishima 1999).

[4] Based on the upper quartile of total recruitment values inferred from observed escapements 1965-1993 and adjusted by the geometric mean of the marine survival indices so that the maximum total recruitment will average to approximately 4000.

[5] Because of the shape of the graph of recruits on spawners this has been called the “hockey stick” model.

[6] J. Gutmann, memorandum to Puget Sound management biologists, 24 February 1998.  See Appendix B for further detail.

[7] 30% age 3, 60% age 4, 10% age 5

[8] Only years 4-25 were counted, since years 0-3 were always seeded with the same values.