Ecological Archives M073-002-A1

Kung-Sik Chan, Nils Chr. Stenseth, Kyrre Lekve, and Jakob Gjøsæter. 2003. Modeling pulse disturbance impact on cod population dynamics: the 1988 algal bloom of Skagerrak, Norway. Ecological Monographs 73:151–171.

Appendix A. Derivation of a fjord-based, age-structured model for cod (Gadus morhua L.) dynamics along the Norwegian Skagerrak coast. Click here for a pdf version.

Derivation of Eqs. 1 through 4.

We now derive Eqs. 2a–2c from 1a–1c. First, we generalize Eqs. 1a–1c by replacing Zf,t by (Zf,t + xf,t(0)) and incorporating the covariate species in Eq. 1b to obtain

Xs,t = (Zf,t+ x(0)f,t) exp(s + t + s,t)

(A.1a)

Ys,t = Xs,t-1exp(-xs,t-1-ys,t-1+ ws,t-1)

(A.1b)

Zf,t = mtYs,t-1+ tZ f,t-1.

(A.1c)

In the log-scale, the preceding equations become

xs,t = log(exp(zf,t)+ x(0)f,t) + s+ t+ s,t

(A.2a)

ys,t = (1-)xs,t-1-ys,t-1+ws,t-1

(A.2b)

zf,t = log(m exp(-F1,t-1+ys,t-1)+ exp(-F2,t-1+ z f,t-1)),

(A.2c)

where xs,t = log(Xs,t ), ys,t = log(Ys,t), and zf,t = log(zf,t); all summations sums over all sites s in a fjord f. Note that mt = m exp(-F1,t-1) and t = exp(-F2,t-1), where the F’s denote fishing mortality. Consider the case that = 0, F1,t-10, F2,t-10, and assume the simplified model is stationarity, with Y,s and mZ,f being the stationary means of Ys,t and Zf,t, respectively. Also, write y,s = log(Y,s ) and z,f = log(Z,f ). For the simplified model, taking expectation on both sides of Eq. A.1c yields

Z,f = s mY,s + Z,f.

Hence, (1-)Z,f = s mY,s, which will be useful below. The basic idea for deriving Eqs. 2a–2c is to approximate the right side of Eqs. A.2a–A.2c by a first order Taylor expansion around = 0, F1,t-1 = 0, F2,t-1 = 0, ys,t-1 = y,s, z f,t-1 = z,f and z f,t = z,f . Equations 2a–2c are obtained by noting that for Eq. A.2a, xs,t /zf,t = 1 and xs,t /l = x(0)f,t /z,f, where all derivatives are evaluated at = 0, F1,t-1 = 0, F2,t-1 = 0, ys,t-1 = y,s, z f,t-1 = z,f and z f,t = z,f. For Eq. A.2c,

zf,t /F1,t-1 = -( mY,s)/(s mY,s + Z,f ) = -(1- ), zf,t /F2,t-1 = -Z,f /(s mY,s+Z,f )

= -, zf,t / ys,t-1 = mY,s /(s mY,s + Z,f ), and zf,t / z f,t-1 = Z,f /(s mY,s + Z,f ) = .

Define cs = mY,s /[(s mY,s + Z,f )(1- )]. Then it can be readily checked that s cs = 1. Hence, Eqs. A.2a–A.2c approximately equal

xs,t = zf,t + fxf,t(0) + s + t + s,t

(2a)

ys,t = (1-)xs,t-1-ys,t-1 + ws,t-1

(2b)

zf,t = df + (1-)csys,t-1 + z f,t-1- [(1-)F1,t-1 + F2,t-1];

(2c)

where df =z,f -(1-)csy,s -z,f is a constant, and f = /Z,f. Consequently, we have

xf,t = zf,t + fx(0)f,t + f + t + f,t

(3a)

yf,t = (1-)xf,t-1-yf,t-1+wf,t-1

(3b)

zf,t = df + (1-q)yf,t-1+ z f,t-1- [(1-)F1,t-1 + F2,t-1],

(3c)

where the last equation [Eq. 3c] is obtained by re-labeling some of the variables in Eqs. 2 so as to make all the model-variables fjord-specific (i.e., xf,t = csxs,t and similarly we can define yf,t).

Next, we outline the derivation of Eq. 4. Adding to the left side of Eq. 3c the product of times the lag 1 of the left side of Eq. 3c and doing likewise to the right side of Eq. 3c eliminates the y’s from the equation, because yf,t + yf,t-1 = (1-)xf,t-1 +
wf,t-
1, owing to Eq. 3b. Specifically,

zf,t + zf,t-1

= (1+ )df + (1-)(yf,t-1+yf,t-2)+ (z f,t-1 + zf,t-2)- [(1-)(F1,t-1 +F1,t-2)

+ (F2,t-1 + F2,t-2)]

= (1 + )df + (1-)[(1-)xf,t-2 + wf,t-2]+ (z f,t-1+zf,t-2)

[(1-)(F1,t-1 + F1,t-2) + (F2,t-1 + F2,t-2)].

(A.3)

Equation 3a implies that zf,t = xf,t –( fx(0)f,t + f + t + f,t ). Upon substituting this expression into Eq. A.3, we obtain Eq. A.4 after some algebra and noting that t is modeled as a linear combination of water temperature and the NAO:

xf,t = (-)xf,t-1 + ( + (1-)(1-))xf,t-2 + (1-)wf,t-2

+ fx(0)f,t + f(-)x(0)f,t-1 - fx(0)f,t-2

+ constf + f,t + (-q)f,t-1 - f,t-2

+ {-[(1-)F1,t + F2,t] - [(1-)F1,t-1 + F2,t-1]}

+ weathert

(A.4)

where weathert = 0Tt + 1Tt-1 + 2Tt-2 + k0naot + k1naot-1 + k2naot-2 (where further 0 = , 1 = (-) and
2 = -, and k0 = k, k1 = (-)k and k2 = -gk) and constf = (1 + )df + (1-)(1+ )f.

Table A1a. A complete list of variables and their description.

Variables

Symbol

Description

s,t

stochastic effect (white noise) on cod reproduction at site s in year t

f,t

(weighted) stochastic effect (white noise) on cod reproduction in fjord f in year t

It

dummy variable of the 1988 bloom; equals 1 for t = 1988 and 0 otherwise

naot

Northern Atlantic Oscillation (NAO) index in year t

Tt

spring water temperature in year t

ws,t

log abundance of covariate species at site s in year t

wf,t

(weighted) log abundance of covariate species in fjord f in year t

xf,t(0)

amount (in million) of larvae released in fjord f in year t

Xs,t

0-group cod abundance at site s (within fjord f) in year t

xs,t

log 0-group cod abundance at site s (within fjord f) in year t

xf,t

(weighted) log 0-group cod abundance in fjord f and in year t

Ys,t

1-group cod abundance at site s (within fjord f) in year t

ys,t

log 1-group cod abundance at site s (within fjord f) in year t

yf,t

(weighted) log 1-group cod abundance in fjord f and in year t

Zf,t

abundance of adult and mature cod in fjord f in year t

zf,t

log abundance of adult and mature cod in fjord f in year t

TableA1b. A complete list of variables and their description.

Parameters

Symbol

Description

a0, a1, a2

common direct (lag1, lag2) bloom effect on the (weighted) log 0-group abundance of any given fjord

Af

direct bloom effect on the (weighted) log 0 -group abundance in fjord f

s

site effect in cod reproduction

t

year effect in cod reproduction

ar1, ar2

Lag 1 and lag 2 autoregressive coefficients in Eq. 5

within-cohort intraspecific effect

Bf

lag-1 bloom effect on the (weighted) log 0 -group abundance in fjord f

cs

relative 1-group cod abundance at site s within fjord f

Bf

lag-2 bloom effect on the (weighted) log 0 -group abundance in fjord f

dt

intervention effect function of the 1988 bloom

covariate species effects

f

fraction of the natural spawning population size that gives rise to 1 million larvae in fjord f

equals 0

0, 1, 2

water temperature effects from current year, last year and two years ago.

F1,t

fishing mortality rate of the 1-group cod

F2,t

fishing mortality rate of the mature cod

Between-cohort intraspecific effect

equals 0

0, 1, 2

NAO effects from current year, last year and two years ago.

average number of mature cod needed to spawn 1 million larvae

ma1, ma2

lag 1 and lag 2 moving-average coefficients in Eq. 5

Y,s

stationary mean 1-group abundance at site s under the simplified (1a-1b) with no trends nor covariates

y,s

equals log(Y,s)

Z,s

stationary mean mature cod abundance in fjord f under the simplified (1a-1b) with no trends nor covariates

z,s

equals log(µZ,f)

mt

survival rate from the 1-group cod to 2-group

annual rate of change in fishing mortality of mature cod

common direct bloom effect on the 1-group cod

common indirect bloom effect on the 1-group cod

baseline survival rate of the mature cod (in year 1970)

t

survival rate of adult and mature cod

common bloom effect on the 0-group cod

Table A2a. Parameter estimates obtained through the model fitting assuming, for reference, no effect of the algae bloom (see Chan et al. 2002b); f = /Z,f. Bold P values represent significance at the 5% level. Larvae releases are only performed in some fjords.

Parameter
Estimate
SE
Ratio
P value
Fjord name
ma1 -0.13 0.105 -1.23 0.226  
ar2 0.45 0.090 5.02 0.000
ma2 -0.36 0.098 -3.68 0.001
1 -0.0073 0.0019 -3.90 0.000
k4 -0.12 0.048 -2.53 0.015
0 -0.073 0.054 -1.37 0.177
1 0.098 0.056 1.75 0.086
2 0.13 0.058 2.21 0.032
0 0.043 0.031 1.41 0.164
1 -0.068 0.033 -2.02 0.049
2 -0.055 0.037 -1.49 0.143
1 0.13 0.054 2.33 0.024 Torvefjord
2 -0.041 0.030 -1.39 0.171 Topdalsfjord
3 0.036 0.017 2.13 0.038 Høvåg
4 0.045 0.014 3.30 0.002 Bufjord
5 -0.0033 0.0097 -0.34 0.735 Flødevigen
7 -0.0047 0.0086 -0.54 0.590 Sandnesfjord
8 0.021 0.018 1.21 0.234 Søndeledfjord
11 0.016 0.27 0.57 0.569 Kilsfjord
16 -0.011 0.010 -1.11 0.272 Nøtterø
17 0.018 0.0061 2.99 0.004 Holmestrand area
20 -0.0086 0.0046 -1.88 0.067 Hvaler

 

Table A2b. Parameter estimates of the ecological parameters obtained from the equating the coefficients in Eqs. 4 and 5. Bold P values represent significance at the 5% level.

Term Estimate SE Ratio P value
0.67 0.071 9.49 Term
0.54 0.11 4.77 Term
0.73 0.094 7.80 Term
Pollack old -0.37 0.16 -2.35 Term

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