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  • 8/13/2019 Stock-Recruitment Analysis for Escapement Goal Development: a Case Study of Pacific Salmon in Alaska

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    743

    *Corresponding author: [email protected] email address: [email protected]

    Stock-Recruitment Analysis for Escapement Goal

    Development: a Case Study of Pacific Salmon in Alaska

    ROBERTA. CLARK*, DAVIDR. BERNARD1, ANDSTEVEJ. FLEISCHMAN

    Alaska Department of Fish and Game, Sport Fish Division

    Research and Technical Services

    333 Raspberry Road, Anchorage, Alaska 99518, USA

    American Fisheries Society Symposium 70:743757, 2009

    2009 by the American Fisheries Society

    Abstract.

    The constitutional mandate to sustain yields along with regulatory guidancefrom the Sustainable Salmon and Escapement Goal policies of the State of Alaska pro-

    vide the impetus for development and implementation of escapement goals for salmon.

    When appropriate, a stock-recruitment analysis can provide vital insight on the popula-

    tion dynamics of a salmon stock. This insight can greatly facilitate the development of

    a scientifically defensible escapement goal to sustain yields from the stock. However,

    the role of stock-recruitment analysis in escapement goal development is often misun-

    derstood and misapplied. Although many analysts focus on the quantity and quality of

    the data needed to conduct a stock-recruitment analysis, other factors such as the spe-

    cies of salmon, type and size of fishery, management constraints, social and economic

    constraints, and information content of the data are also important. Much of the confu-

    sion about stock-recruitment analysis arises because, while the analysis is primarily astatistical procedure, the actual development and implementation of an escapement goal

    is primarily a scientific and practical endeavor. Many see the ultimate goal of a stock-

    recruitment analysis as the identification of the escapement that produces maximum

    sustained yield, although in many cases it may identify much more than that, or less.

    Several case studies are used to illustrate the potential uses of stock-recruitment theory

    in the development of escapement goals.

    Introduction

    Article VIII, section 4 of the Alaska Con-

    stitution states that (from Harrison 1992):

    Fish, forests, wildlife, grasslands, and

    all other replenishable resources belonging

    to the State shall be utilized, developed, and

    maintained on the sustained yield principle,

    subject to preferences among benecial

    uses.

    This mandate for sustainable manage-ment of Pacic salmon Oncorhynchus spp.

    provided the impetus for development of a

    scientically defensible escapement goal

    policy in Alaska. Along with the statutory

    duties and powers of the Commissioner of

    the Alaska Department of Fish and Game

    (ADF&G) and relevant management plans

    for salmon stocks, the development of es-

    capement goals is regulated by the policy

    for the management of sustainable salmon

    sheries and the policy for statewide salmon

    escapement goals (Title 5 of the Alaska Ad-ministrative Code, Chapter 39).

    These two regulatory policies dene four

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    744 Clark et al.

    types of escapement goals, two of which are

    most important to sustained yield manage-

    ment of salmon stocks. The biological es-

    capement goal (BEG) is dened as: the es-

    capement that provides the greatest potentialfor maximum sustained yield (MSY). As an

    alternative to management for MSY, the sus-

    tainable escapement goal (SEG) is dened

    as: the escapement that is known to provide

    for sustained yield. Both of these escapement

    goals must be described as ranges that take

    into account our uncertainty in the data and

    variation in stock productivity. The two regu-

    latory policies also stipulate that BEGs and

    SEGs for Pacic salmon be developed from

    the best available data and be scientically

    defensible.

    The concept of scientic defensibility

    is embodied in the well-known production

    models and statistical formulations of stock-

    recruitment theory. Much of the scientic dis-

    course on salmon production and the mainte-

    nance of yields is focused on model selection

    (e.g., Schmidt et al. 1998), the effect of mea-surement error on model performance (e.g.,

    Walters and Ludwig 1981), and the incorpo-

    ration of accumulated knowledge about mod-

    el parameters into the statistical formulation

    (e.g., Hilborn and Liermann 1998) or what is

    herein referred to as stock-recruitment analy-

    sis. While these are important considerations

    in the methodological development of the

    theory, little of the peer-reviewed literature

    on stock-recruitment analysis focuses on im-provements in data collection and the inter-

    pretation of these data in relation to the sus-

    tained yield principle in Alaskan law.

    It is argued here that current salmon pro-

    duction theory is sufcient to address the

    question of sustaining yields of Pacic salm-

    on in Alaska via the development of BEGs or

    SEGs. When used judiciously, stock-recruit-

    ment analysis is an important tool in this de-

    velopment. It is also argued that factors such

    as the species of salmon, type and size of sh-

    ery, management constraints, social and eco-

    nomic constraints, and information content

    of the production data are as important as the

    availability of data and sophistication of the

    statistical machinery used for stock-recruit-

    ment analyses. Four case histories from sh-eries on salmon stocks throughout Alaska are

    presented to illustrate these often neglected

    considerations in escapement goal analysis.

    Methods

    Brood year tables were constructed from

    escapements (S) and subsequent production

    (R) from the following stocks that likely rep-resent the range of information content and

    shing power typical of salmon sheries in

    Alaska (Table 1; Figure 1): early-run Chignik

    River sockeye salmon O. nerka(Ruggerone

    et al. 1999; Witteveen et al. 2005; Table 2),

    Chilkat River Chinook salmon O. tshawyts-

    cha (Ericksen and McPherson 2004; Table

    3); Kenai River sockeye salmon, (Hasbrouck

    and Edmundson 2007; Table 4); and Good-

    news River Chinook salmon (Brannian et al.2006; Table 5).

    Simple stock-recruitment analyses were

    performed on each brood table using the lin-

    earized form of the Ricker relationship with

    multiplicative process error (Hilborn and

    Walters 1992) to estimate parameters (equa-

    tion (1) and reference points (equations (2)

    through (4)). Beginning with the familiar

    nonlinear form of the stochastic Ricker equa-

    tion,

    ( ) ( )= expexp SSR , (1a)

    and then dividing by S and taking natural logs

    to form the linear regression recipe

    ( )2,0~;lnln +=

    NSS

    R(1b)

    A linear regression of ln(R/S) on Swill

    estimate the parameters ln (y-intercept),

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    745Stock-Recruitment Analysis for Escapement Goal Development

    TABLE1.Salmon stocks used to illustrate the different techniques for developing an escapement goal

    based on the number of years of stock-recruit data, information content of these data, and the shing

    power of the sheries on these stocks.

    Stock n(years) Information Content Fishing Power

    Chignik sockeye 75 high high

    Chilkat Chinook 7 low very low

    Kenai sockeye 28 low very high

    Goodnews Chinook 17 high low

    Chilkat River

    Goodnews River

    Chignik River

    Kenai River

    Chilkat River

    Goodnews River

    Chignik River

    Kenai River

    FIGURE1.The state of Alaska with the location of major communities and four river systems discussed

    in this paper.

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    746 Clark et al.

    TABLE

    2.Escapementsandsubsequentproductionofearly-runChignikRiversockeyesalmonOncorhy

    nchusnerkaforthe1922-1996

    broodyears

    (fromRuggeroneetal.

    1999andWitteveenetal.

    2005).

    BroodYear

    Escapement

    Produ

    ction

    BroodYear

    Escapement

    Production

    Broo

    dYear

    Escapement

    Prod

    uction

    192

    2

    86,4

    21

    96

    3,8

    14

    1947

    2,3

    86,7

    33

    181,1

    12

    1

    972

    326,3

    20

    995,9

    71

    192

    3

    4,6

    42

    38

    0,3

    59

    1948

    3

    84,6

    37

    327,2

    95

    1

    973

    538,4

    62

    1,1

    72,4

    28

    192

    4

    121,9

    83

    1,32

    2,5

    57

    1949

    2

    13,2

    69

    308,5

    34

    1

    974

    364,6

    03

    607,5

    96

    192

    5

    386,3

    64

    11

    7,6

    62

    1950

    2

    06,2

    70

    392,6

    27

    1

    975

    319,8

    90

    655,8

    27

    192

    6

    289,0

    09

    53

    0,1

    94

    1951

    1

    25,1

    26

    625,6

    89

    1

    976

    548,9

    53

    982,3

    61

    192

    7

    857,8

    81

    2,67

    7,1

    84

    1952

    34,1

    55

    230,8

    20

    1

    977

    364,5

    57

    2,2

    33,7

    83

    192

    8

    507,3

    53

    82

    0,9

    81

    1953

    1

    68,3

    75

    357,6

    07

    1

    978

    419,7

    32

    968,2

    98

    192

    9

    995,8

    32

    1,05

    4,1

    67

    1954

    1

    84,9

    53

    142,4

    21

    1

    979

    491,4

    67

    3,6

    12,1

    07

    193

    0

    92,9

    55

    37

    7,4

    85

    1955

    2

    56,7

    57

    554,4

    95

    1

    980

    369,5

    80

    1,7

    32,7

    41

    193

    1

    96,2

    01

    1,12

    8,2

    31

    1956

    2

    89,0

    96

    208,1

    68

    1

    981

    570,2

    10

    1,8

    33,4

    32

    193

    2

    2,1

    51,7

    34

    34

    1,2

    98

    1957

    1

    92,4

    79

    350,5

    12

    1

    982

    616,1

    17

    2,3

    23,6

    43

    193

    3

    223,9

    13

    62

    1,4

    00

    1958

    1

    20,8

    62

    242,3

    70

    1

    983

    426,1

    78

    564,1

    74

    193

    4

    866,8

    90

    1,65

    8,4

    66

    1959

    1

    12,2

    26

    340,9

    46

    1

    984

    597,7

    13

    636,0

    40

    193

    5

    194,6

    36

    41

    9,7

    09

    1960

    2

    51,5

    67

    774,7

    56

    1

    985

    373,0

    40

    772,9

    20

    193

    6

    548,0

    39

    64

    5,9

    85

    1961

    1

    40,7

    14

    571,6

    45

    1

    986

    557,7

    72

    2,5

    23,2

    15

    193

    7

    205,6

    13

    80

    9,5

    50

    1962

    1

    67,6

    02

    693,4

    73

    1

    987

    589,2

    99

    1,4

    34,0

    36

    193

    8

    175,9

    72

    1,02

    5,5

    70

    1963

    3

    32,5

    36

    698,7

    03

    1

    988

    420,5

    80

    1,6

    58,4

    60

    193

    9

    1,1

    42,8

    52

    48

    9,2

    32

    1964

    1

    37,0

    73

    755,7

    26

    1

    989

    384,0

    01

    1,7

    06,4

    00

    194

    0

    176,3

    07

    50

    5,3

    79

    1965

    3

    07,1

    92

    1,9

    48,1

    44

    1

    990

    434,5

    50

    1,5

    26,8

    44

    194

    1

    374,4

    20

    1,58

    3,5

    79

    1966

    3

    83,5

    45

    1,3

    03,5

    67

    1

    991

    662,6

    60

    1,4

    95,5

    03

    194

    2

    442,9

    81

    3,05

    9,1

    05

    1967

    3

    28,0

    00

    240,7

    12

    1

    992

    360,6

    81

    929,7

    59

    194

    3

    701,8

    59

    70

    0,6

    58

    1968

    3

    42,3

    43

    1,2

    10,9

    27

    1

    993

    364,2

    61

    903,5

    37

    194

    4

    291,8

    44

    33

    4,0

    93

    1969

    3

    66,5

    89

    476,2

    82

    1

    994

    769,4

    65

    2,0

    38,9

    09

    194

    5

    217,8

    82

    24

    5,5

    34

    1970

    5

    36,2

    57

    441,6

    85

    1

    995

    366,4

    95

    2,1

    17,1

    30

    194

    6

    774,1

    30

    22

    4,1

    63

    1971

    6

    71,6

    68

    1,3

    83,4

    55

    1

    996

    464,7

    48

    1,5

    70,3

    75

    Average

    430,2

    54

    983,9

    14

    SD

    380,9

    63

    740,9

    58

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    747Stock-Recruitment Analysis for Escapement Goal Development

    TABLE 3.Escapements and subsequent production of Chilkat River Chinook salmon Oncorhynchus

    tshawytschafor the 19911997 brood years (from Ericksen and McPherson 2004).

    Brood Year Escapement Production

    1991 5,897 12,932

    1992 5,284 5,542

    1993 4,472 3,231

    1994 6,795 1,645

    1995 3,790 4,348

    1996 4,920 5,637

    1997 8,100 7,081

    Average 5,608 5,774

    SD 1,465 3,619

    TABLE4.Escapements and subsequent production (in thousands) of Kenai River sockeye salmon Onco-

    rhynchus nerkafor the 19681995 brood years (from Hasbrouck and Edmundson 2007).

    Brood year Escapement Production

    1968 82 916

    1969 52 409

    1970 72 520

    1971 289 863

    1972 302 2,1861973 358 1,995

    1974 144 665

    1975 129 895

    1976 353 1,187

    1977 664 2,811

    1978 350 3,451

    1979 246 1,111

    1980 398 2,346

    1981 359 2,268

    1982 566 8,9301983 557 8,697

    1984 310 3,252

    1985 396 2,246

    1986 400 1,741

    1987 1,333 9,531

    1988 839 2,120

    1989 1,334 3,898

    1990 439 1,334

    1991 376 3,926

    1992 752 3,469

    1993 670 1,2871994 895 2,511

    1995 521 1,421

    Average 471 2,714

    SD 328 2,455

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    748 Clark et al.

    (slope), and2 (residual error). Then ln is

    adjusted for lognormal process error (Hilborn

    1985),

    2

    2

    lnln

    += (1c)

    and to estimate the relevant reference points

    for salmon management from the regressionparameters:

    =

    lnEQS

    , (2)

    ln07.05.0EQMSY SS , and (3)

    ln07.05.0lnMSY . (4)

    Statistical uncertainty about the param-

    eters and reference points was assessed with

    a bootstrap technique (Efron and Tibshirani

    1993); resampling the residuals of the linear

    regression with replacement, calculating all

    parameter estimates and reference points for

    each bootstrap replicate, and using percentiles

    of the bootstrap values to obtain interval es-timates. Here, for comparison among stocks

    a nonparametric analog of the coefcient of

    variation (NPCV) is also calculated for each

    parameter and reference point (Prager and

    Mohr 1999):

    ; (5)

    where an NPCV of 25% or less was consid-ered precise.

    TABLE5.Escapements and subsequent production of Goodnews River Chinook salmon Oncorhynchus

    tshawytschafor the 19811997 brood years (from Brannian et al. 2006).

    Brood year Escapement Production

    1981 11,454 16,538

    1982 4,332 11,870

    1983 20,420 9,479

    1984 12,003 12,412

    1985 10,810 6,030

    1986 6,186 11,899

    1987 6,762 10,150

    1988 8,131 11,255

    1989 4,806 18,849

    1990 11,292 9,1341991 6,473 12,069

    1992 3,757 7,466

    1993 7,076 22,817

    1994 11,722 7,006

    1995 14,701 24,571

    1996 8,907 9,532

    1997 10,153 7,187

    Average 9,352 12,251

    SD 4,211 5,424

    ( )median

    percentilepercentileNPCV

    thth85.3015.69

    =

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    749Stock-Recruitment Analysis for Escapement Goal Development

    Apart from this analytical recipe for es-

    timating the reference points from the linear

    regression parameters, it is important to note

    that SEQ

    is an estimate of the carrying capaci-

    ty of a salmon stock, where carrying capacityis dened as the average escapement of the

    stock when it is not being shed (Figure 2).

    Moreover, ln is an estimate of the intrinsic

    rate of increase of the stock, where intrinsic

    rate of increase is dened as the natural loga-

    rithm of average recruits per spawner as Sap-

    proaches zero.

    These denitions also have relevance to

    the rate of harvest on a stock while stock-

    recruitment data are being collected. Walters

    and Hilborn (1976) showed that in the case

    of very low harvest rate most of the stock-

    recruitment data pairs are on the right-hand

    side of the stock-recruit curve, the carrying

    capacity of a stock is known, but not the in-

    trinsic rate of increase. Conversely, they also

    showed that in the case of a very high harvest

    rate when most of the stock-recruitment data

    pairs are on the left-hand side of the stock-re-

    cruitment curve, the intrinsic rate of increase

    is known but not the carrying capacity.

    Using information from each of the four

    salmon stocks as examples and the aforemen-tioned principles of stock-recruitment analy-

    sis, recommendations for establishing a BEG

    or SEG were developed for each stock that

    are scientically defensible based on infor-

    mation content of the stock-recruit data, the

    denitions of carrying capacity and intrinsic

    rate of increase, and the analytical relation-

    ship between the two.

    Case Studies

    Early run Chignik River sockeye

    salmon.The Chignik River is located on

    the Alaska Peninsula near the community of

    Chignik (Figure 1). Sockeye salmon are pri-

    marily harvested in a commercial seine sh-

    ery (Bouwens and Poetter 2006). Although

    a long history of exploitation exists on this

    Escapement

    Pro

    duction

    Escapement

    Pro

    duction

    FIGURE2.Graphical representation of reference points (SMSY

    , SEQ

    , MSY

    ) on a Ricker stock-recruitment

    curve. The diagonal line is the replacement line and the shaded region is the area of surplus produc-

    tion.

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    750 Clark et al.

    stock, recent rates of annual exploitation are

    moderate and range from 51 to 85%, aver-

    aging 69% during 19592003 (Witteveen et

    al. 2005). Recent stock assessment consists

    of escapements estimated by weir and har-vests estimated by allocation of age classes to

    stream of origin based on the relative magni-

    tude of escapements (Witteveen et al. 2005).

    The brood table consists of 75 years (1922

    1996) of escapement and subsequent produc-

    tion estimates (Table 2). Sufcient shing

    power occurs in the commercial seine shery

    to harvest all available surplus production so

    that a BEG was desired.

    A plot of subsequent production on

    brood-year escapement indicates that a broad

    range of escapements have occurred, and that

    production has not replaced escapement (i.e.,

    data points below the replacement line) in 11

    out of 75 years, most notably at the highest

    levels of escapement (Figure 3). The infor-

    mation content of these data are high because

    there is information about the intrinsic rate

    of increase at high exploitation rates and on

    carrying capacity at lower exploitation rates.

    The stockrecruitment analysis indicated that

    estimates of ln and were relatively precise

    with NPCV of 8% and 14%, respectively (Ta-

    ble 6). Estimates of SEQ, SMSY, and MSYwerealso precise (NPCV of 11%, 11% and 5%, re-

    spectively), indicating that a BEG based on

    SMSY

    could be developed from this analysis

    (Table 7). A BEG appears to be a good t to

    this shery as there is close correspondence

    between the observed average exploitation

    rate (69%) and MSY

    (69%; Table 6).

    Chilkat River Chinook salmon.The

    Chilkat River is located near the city of Haines

    in southeast Alaska (Figure 1). Chinook

    salmon are harvested in commercial and ma-

    rine sport sheries located in Lynn Canal as

    well as in the commercial troll shery (Erick-

    sen and McPherson 2004). Although a long

    history of exploitation exists on this stock,

    recent rates of annual exploitation are low

    and ranged from 8 to 19%, averaging 12%

    during 19881991. Recent stock assessment

    Escapement

    0

    500,000

    1,000,000

    1,500,000

    2,000,000

    2,500,000

    3,000,000

    3,500,000

    0 500,000 1,000,000 1,500,000 2,000,000 2,500,000 3,000,000 3,500,000

    Produ

    ction

    Escapement

    FIGURE3.Plot of subsequent production on brood year escapement and the estimated Ricker curve from

    the early-run of Chignik River sockeye salmon Oncorhynchus nerka, 19221996 brood years.

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    751Stock-Recruitment Analysis for Escapement Goal Development

    consists of escapements estimated by mark-

    recapture and harvests estimated by coded-

    wire tag recoveries. The brood table consistsof seven years (19911997) of escapement

    and subsequent production estimates (Table

    3). Prior to this analysis no escapement goal

    was available for this stock. Although la-

    tent shing power is available that could be

    brought to bear on this stock, the initial de-

    sire was to develop an escapement goal (SEG

    or BEG) for this stock so existing sheries

    could be liberalized when surplus production

    was realized.A plot of subsequent production on brood-

    year escapement indicates that a narrow range

    of escapements have occurred and that pro-

    duction has not replaced escapement in three

    out of seven years, indicating that this stock is

    at or near carrying capacity (Figure 4). Infor-

    mation content of these data are low because

    there is no information on the intrinsic rate

    of increase, but there is on carrying capacity.

    The stock-recruitment analysis indicated thatestimates of ln and were imprecise with

    NPCV of 67% and 70%, respectively (Table

    6). Estimates of SEQ

    and SMSY

    were notably

    more precise (NPCV of 19% and 25%, re-

    spectively) because there was good informa-tion about carrying capacity. The estimate of

    MSY

    was poor (NPCV of 53%) because of the

    lack of production data at low escapements

    and the resulting lack of information about

    intrinsic rate of increase. The estimate of SEQ

    is defensible; it differs little from the average

    of escapements observed while shing power

    was low (average S= 5,608; Table 3). From

    the estimate of SEQ

    , a conservative estimate of

    SMSYcould be developed by assuming ln was0 (assumes return-per-spawner is 1 at low es-

    capements) and solving equation (3) for SMSY

    .

    Alternatively, a less conservative estimate of

    SMSY

    could be calculated by using an estimate

    of average ln from Chinook salmon stocks

    in southeast Alaska (average ln = 1.92) as

    was done by Ericksen and McPherson (2004).

    Although these methods produced an estimate

    of SMSY

    , information was still missing on re-

    turns at escapements near SMSY

    for this stock,so that a SEG was recommended instead of a

    BEG (Table 7).

    TABLE6.Summary of parameter estimates, estimated reference points, and nonparametric CVs (in pa-

    rentheses) from linear regression and bootstrapping of stock-recruitment data from four salmon stocks

    in Alaska.

    Stock

    ln 2 SEQ SMSY MSY

    Chignik

    sockeye

    1.87

    (8%)

    1.68 10-6

    (14%)

    0.60

    (10%)

    1,114,168

    (11%)

    410,940

    (11%)

    69%

    (5%)

    Chilkat

    Chinook

    0.91

    (67%)

    1.38 10-4

    (70%)

    0.49

    (43%)

    6,590

    (19%)

    2,875

    (25%)

    40%

    (53%)

    Kenai

    sockeye

    2.11

    (8%)

    5.71 10-4

    (51%)

    0.29

    (13%)

    3,698a

    (48%)

    1,303a

    (50%)

    74%

    (5%)

    Goodnews

    Chinook

    1.39

    (19%)

    1.08 10-4

    (23%)

    0.19

    (16%)

    12,843

    (10%)

    5,169

    (12%)

    56%

    (14%)

    ain thousands.

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    752 Clark et al.

    TABLE

    7.Summaryoffourcasestudiesofsalmonescapementgoaldevel

    opment.

    Stock

    n

    Info

    rmation

    Fishing

    ln

    Goaltype

    Basisforgoal

    Con

    tent

    Power

    Chignik

    sockeye

    75

    high

    high

    kn

    own

    known

    BEG

    Rickerregression

    to

    estimateSMSY

    ChilkatChinook

    7

    low

    low

    un

    known

    unknown

    SEG

    Discountfromca

    rrying

    capacitybasedon

    averageescapement

    Kenaiso

    ckeye

    28

    low

    high

    kn

    own

    unknown

    SEG

    Exploitationrate

    fromln

    GoodnewsChinook

    17

    high

    low

    kn

    own

    known

    BEGorSEG

    Rickerregression

    to

    estimateSMSY

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    753Stock-Recruitment Analysis for Escapement Goal Development

    Kenai River sockeye salmon.The

    Kenai River is located on the Kenai Pen-insula near the cities of Soldotna and Ke-

    nai (Figure 1). Sockeye salmon are har-

    vested in marine commercial, freshwater

    sport, and personal-use fisheries (Shields

    2006). Escapement is estimated with so-

    nar near the mouth of the river, and ma-

    rine harvests are estimated by allocation

    of harvests by age-class to stream of ori-

    gin based on the relative magnitude of

    escapements (Hasbrouck and Edmundson2007). Stock assessments have occurred

    since 1968, with annual exploitation

    ranging from 48 to 94% and averaging

    79%. The brood table consists of 28 years

    (19681995) of escapement and subse-

    quent production estimates (Table 4).

    The current escapement goal is an SEG

    of 500800 thousand (Hasbrouck and Ed-

    mundon 2007). There is sufficient fishingpower in the currently configured fisher-

    ies to harvest all available surplus pro-

    duction so that a BEG was desired.

    A plot of subsequent production on

    brood year escapement indicates that a nar-row range of escapements have occurred

    and that production has always replaced es-

    capement, indicating a lack of information

    about this stocks carrying capacity (Figure

    5). Information content of these data are low

    because there is information on the intrinsic

    rate of increase, but little or none on carry-

    ing capacity. The stock-recruitment analysis

    indicated that the estimate of ln was pre-

    cise (NPCV of 8%) as was the estimate of

    MSY(NPCV of 5%; Table 6). Estimates of

    (NPCV of 51%), SEQ

    (NPCV of 48%), and

    SMSY

    (NPCV of 50%) were grossly impre-

    cise (Table 6), which prohibited the devel-

    opment of a defensible BEG. On the other

    hand, the estimate of MSY

    (74%) is lower

    than the observed average exploitation

    rate (81%), indicating that the current SEG

    should be increased somewhat. Althoughno defensible BEG could be developed, ad-

    vice to increase the current SEG is useful to

    shery managers (Table 7). While likely not

    Escapement

    0

    2,000

    4,000

    6,000

    8,000

    10,000

    12,000

    14,000

    0 2,000 4,000 6,000 8,000 10,000 12,000 14,000

    Spawning Abundance

    Production

    Escapement

    FIGURE4.Plot of subsequent production on brood year escapement from Chilkat River Chinook salmon

    Oncorhynchus tshawytscha, 19911997 brood years.

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    754 Clark et al.

    optimal in terms of producing MSY, the cur-

    rent SEG has been shown to be defensible

    and sustainable given that escapements ob-

    served since 1968 have always yielded sur-

    plus production.

    Goodnews River Chinook salmon.The

    Goodnews River is located near the commu-

    nity of Goodnews Bay in southwestern Alaska

    (Figure 1). Chinook salmon are harvested in

    commercial, sport, and subsistence sheries(Jones and Linderman 2006) that primarily

    target sockeye salmon. Escapement is esti-

    mated with a weir, and marine harvests are es-

    timated by allocation of harvests by age-class

    to stream of origin based on the relative mag-

    nitude of escapements (Brannian et al. 2006).

    Recent stock assessments have occurred since

    1981, with annual exploitation ranging from

    16 to 71% and averaging 33%. The brood

    table consists of 17 years (19811997) ofescapement and subsequent production es-

    timates (Table 5). The initial desire was to

    replace an SEG range based on escapements

    indexed with aerial surveys with one (SEG or

    BEG) developed from a brood table based on

    weir-based counts of escapement.

    A plot of subsequent production on

    brood-year escapement indicates that a broad

    range of escapements have occurred, and that

    production has not replaced escapement (i.e.,

    data points below the replacement line) in 5

    out of 17 years, most notably at the highest

    level of escapement (Figure 6). Informationcontent of these data are high because there

    is information on the intrinsic rate of increase

    at high exploitation rates and on carrying ca-

    pacity at lower exploitation rates. The stock

    recruitment analysis indicated that estimates

    of ln and were relatively precise with

    NPCV of 19% and 23%, respectively (Table

    6). Precise estimates of SEQ

    , SMSY

    , and MSY

    were estimated from the parameters (NPCV

    of 10%, 12% and 14% respectively), indicat-ing that a BEG based on S

    MSYcould be devel-

    oped from this analysis (Table 7). However,

    Escapement

    0

    2,000

    4,000

    6,000

    8,000

    10,000

    0 2,000 4,000 6,000 8,000 10,000

    Production

    Escapement

    FIGURE5.Plot of subsequent production on brood year escapement from Kenai River sockeye salmon

    Oncorhynchus nerka (in thousands), 19681995 brood years.

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    755Stock-Recruitment Analysis for Escapement Goal Development

    estimated SMSY

    (5,169 sh) was much lower

    than the observed average escapement for

    this stock (9,352 sh), so that the currently

    congured shery may be unable to harvest

    all available surplus production leading to achronic inability to achieve the BEG. A bet-

    ter option for this shery might be an SEG

    constructed around the range of escapements

    shown to produce sustained yields.

    Discussion

    As illustrated by the four case studies,

    the simple stock-recruitment analyses usedherein provide vital information regarding

    population dynamics and information content

    of the basic adult escapement and production

    data. This information, when combined with

    the practical considerations of the shery

    such as shing power and prior knowledge of

    a particular salmon species can help the ana-

    lyst and shery manager develop a scienti-

    cally defensible escapement goal that meets

    the mandates of Alaskan law. As seen in the

    example of Chignik sockeye salmon, shery

    objectives and the information content of thedata match and all that is needed is a simple

    stockrecruitment analysis to develop a BEG

    that has the potential for producing MSY in

    the long-term. More sophisticated models

    and analytical techniques of these stock and

    recruitment data are available to help rene

    our analysis, but the basic premise of a de-

    fensible BEG would remain. Our experience

    with salmon stocks and sheries in Alaska is

    that this situation is the exception rather than

    the norm.

    Situations similar to the Chilkat Chinook

    salmon and Kenai sockeye salmon case stud-

    ies are most often seen, where the results of

    simple stock-recruitment analyses are in-

    conclusive or partially conclusive, and the

    Escapement

    0

    5,000

    10,000

    15,000

    20,000

    25,000

    30,000

    0 5,000 10,000 15,000 20,000 25,000 30,000

    Pro

    duction

    Escapement

    FIGURE6.Plot of subsequent production on brood year escapement and the estimated Ricker curve from

    Goodnews River Chinook salmon Oncorhynchus tshawytscha, 19811997 brood years.

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    756 Clark et al.

    analyst must use theoretical considerations

    and past knowledge of similar stocks to de-

    velop a scientically defensible escapement

    goal. In these cases, an SEG is most often

    recommended to managers because escape-ment that produces MSY cannot be specied

    from the available information. In the case

    of Chilkat Chinook salmon this outcome has

    little consequence for a shery that has very

    low shing power which likely would not

    harvest all available surplus production even

    with generous shing time. If additional sh-

    ing power could be brought to bear on this

    stock, then information might be gained on

    production potential at levels of escapements

    lower than currently observed.

    The lack of a BEG is also self perpetu-

    ating in the Kenai sockeye salmon shery

    because current shing power is more than

    sufcient to harvest at rates higher than what

    might be specied at MSY. This results in

    production information that is conned to the

    existing SEG with little potential for gaining

    information on the carrying capacity of thisstock. In this case, it can be argued that the

    current SEG or one slightly higher has suc-

    cessfully sustained high levels of harvest and

    is therefore a defensible escapement goal.

    Situations similar to Goodnews Chinook

    salmon are also seen where knowledge of es-

    capement that produces MSY may not be the

    proper objective for the shery. In this example,

    stock assessment data are collected from a sh-

    ery that harvests multiple species where shingpower may be adequate to harvest surplus pro-

    duction of one species (sockeye salmon in this

    example) and not the other species (Chinook

    salmon). Sufcient information is possessed to

    develop a BEG from simple stock-recruitment

    analysis, but may not be elected to set an es-

    capement goal based on this analysis because

    there is insufcient shing power to harvest all

    available surplus production. In this case, the

    tradeoffs in harvest of two species in a single

    shery may best be handled by a BEG for one

    species and a SEG for the other.

    These case studies also illustrate the

    need for independent corroborative evidence

    of carrying capacity and intrinsic rate of in-

    crease in escapement goal development, es-

    pecially when one or both pieces of this in-formation are missing or inconclusive in the

    stock-recruitment analysis. Existing habitat-

    based and paleolimnological approaches for

    describing carrying capacity (e.g., Bradford

    et al. 1997 for coho salmon O. kisutch, Parken

    et al. 2004 for Chinook salmon, Schindler et

    al. 2005 for lake-rearing sockeye salmon) are

    valuable tools for the development of escape-

    ment goals. The development of these ap-

    proaches is encouraged for other salmon spe-

    cies such as river-rearing sockeye salmon and

    chum salmon O. keta. Similarly, meta analy-

    ses of intrinsic rates of increase for salmon

    species (e.g., Myers et al. 1999) can provide

    helpful information on production dynamics

    and scientic defensibility of an escapement

    goal, especially when shing power is low.

    While the continued statistical development

    of stock-recruitment analysis is laudable, it isargued that the science of setting escapement

    goals and understanding of production dy-

    namics of salmon would be better served by

    an increase in the collection of high quality

    stock assessment data across a broad range

    of salmon species, geographic settings, and

    shing power.

    Acknowledgments

    The authors would like to acknowledge

    the helpful comments and suggestions on

    the content and presentation of this mate-

    rial from the many Alaska Department of

    Fish and Game staff that participated in the

    Escapement Goal Analysis workshops held

    from March 2006 through February 2007

    in Anchorage and Juneau. We also greatly

    appreciate the helpful external peer reviewsprovided to us by the AYK SSI Editorial

    Board.

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    757Stock-Recruitment Analysis for Escapement Goal Development

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