Age-Structured Stock Assessments

K. Limburg lecture notes, Fisheries Science

Outline:
1. What are age-structured assessments? 2. Virtual population assessments (VPA), cohort analysis, and the modern-day descendants 3. Yield-per-recruit models 4. Eggs-per-recruit models (just a mention)

Figure: King (1995)

What are age-structured assessments?

These are tools that keep explicit track of age* classes as part of the assessment procedure. Empirical methods that do not assume any underlying stock/recruit relationships. Typically used by fisheries managers to help set levels of fishing mortality (F) Can also be used to help diagnose overfishing or other causes of stock declines

History of catch-at-age analysis

Virtual Population Analysis Virtual Population Analysis Cohort Analysis CAGEAN (not covered here)
(Pope 1972) (Deriso, Quinn, and Neal 1985) (Gulland 1965) (Fry 1949)

* Or length classes in cases where age cannot be determined

P. Sullivan, Cornell University

The basic theory behind agestructured analysis

Cohort Numbers at Age

800

N t N 0 exp( Zt )
can break this up, to look at year-to-year changes (here, t is used for Age)

200

400

600

5
National Marine Fisheries Service

10 Age

15

20

P. Sullivan, Cornell University

Virtual Population Analysis (VPA)

Virtual Population Analysis (VPA)

Nt1 Nt e
Nt
natural mortality

(F M) t

The idea:
If you know how many fish in a given cohort were caught one year, thats a minimum estimate of how many were alive. If we add in natural mortality, then weve got total mortality and so at least that many fish were alive the year before. Work backwards, year by year, to get annual estimates of cohort abundances and mortality rates. If we assume that M is constant, then we obtain estimates of F for all years in the past, for that cohort.

The abundance of fish of age (t+1), the survivors from the year before, equals Nt reduced by fishing and natural mortality.

Nt+1
Fishing mortality

Virtual Population Analysis (VPA)

For the last year of life:

80

All fish in the last year of a cohorts existence die, either from natural mortality M or fishing mortality F. That number of fish is called the terminal abundance and can be calculated, if you have estimates of M and F, and you know the catch, C

Q: why look backwards?

Answer:
Catch (Number) 40 60 0 20

With a good picture of past and present population dynamics, one can then make the forecasts needed to assess management options.
(Jennings et al. 2001, p. 138)

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 Age
P. Sullivan, Cornell University

VPA, Step 1: Calculating the terminal abundance

Once you know the terminal abundance N in the previous year can be estimated retrospectively, if you know the catch for the previous year and you assume that M is constant. VPA is a retrospective analysis of the cohorts past abundance, based on catch statistics.

Nt

Ct Ft (1 exp( Z t )) Zt

Nt

Ct Ft (1 exp( Z t )) Zt
Ct is the catch that year Ft is your estimate of F that year Zt = Ft + M = total mortality rate (note: assume that M is constant)

Where

Jennings et al. (2001)

VPA, Step 2: Calculating the previous years F Now that we know the terminal abundance, we can employ Gullands version of the catch equation to estimate the previous years instantaneous fishing mortality (Ft-1).

Age Class 1 2 3

Catches 70 90 80

Estimated Assumed C_t M 70.0 0.2 90.1 0.2 0.2

F Z 0.1725 0.3725 0.349 0.549 0.6 0.8

Stock Size 486 335 194

Ct 1

Ft 1 N t ( e Ft 1 M 1) Ft 1 M

Step 3. Once weve solved for Ct-1, we can substitute the Ft-1 and Ct-1 into the equation:

In Step 2, we have to solve for F by iteration. We do this by making a column (I called it the Estimated C_t for estimated catch), typing in the catch equation, and then trying out values for F that make the estimated C_t come out as closely as possible to the actual catch that year. Once you have F for that year, you can calculate N (the Stock Size) for that year, too (Step 3). Repeat Steps 2 & 3 and work backwards through time.

N t 1

Ct 1 Ft 1 (1 exp( Z t 1 )) Z t 1

Same equation as in Step 1, only at t-1

Age-based cohort analysis. alternative to VPA developed by Pope (1972) approximates the results of VPA, without having to iterate to determine the values of F. we still use catches and initial guesstimates of F and M to get the terminal year abundance we still work backwards through time to reconstruct previous values of F and stock structure.

N t 0. 5 N t e M / 2 N t e M / 2 Ct
N t 1 ( N t e M / 2 Ct ) eM / 2

Jennings et al. (2001)

N t 1 ( N t e M / 2 Ct ) e M / 2
We want to have an expression for Nt, so we have to re-arrange this equation:

Cohort analysis, step-by-step. 1. Same as Step 1 in VPA: calculate the terminal abundance of the most recent year, t.

N t ( N t 1e M / 2 Ct ) e M / 2
Nt is the # of fish alive at the start of the year, based on the catch throughout the year and the number surviving until next year. Now were ready for step-by-step cohort analysis.

Nt

Ct Ft (1 exp( Z t )) Zt
Ct is the catch that year Ft is your estimate of F that year Zt = Ft + M = total mortality rate (note: assume that M is constant)

Where

Cohort analysis, step-by-step. 2. Use the new equation to calculate the # of fish alive at the start of the previous year:

Cohort analysis, step-by-step. 3. Now determine F for age-2 fish from the usual exponential decay model:

N t ( N t 1e M / 2 Ct ) e M / 2
Example: say we are following the same three years of a cohorts existence, and that N3 was calculated to be 194 by Step 1 in the VPA. M is 0.2, and 90 fish were caught (Ct = 90). Then,

N t 1 N t e ( Ft M )
N t 1 e ( Ft M ) , Nt ln(

Re-arrange to get Ft:

N2 = (194 e(0.2/2) + 90) e(0.2/2) = 336 fish

N t 1 ) Ft M , Nt

(Step 3)

N Ft ln( t ) M N t 1
F2 = ln(336/194) 0.2 = ln(1.732) 0.2 = 0.549 0.2 = 0.349
Same as VPA result

You keep on repeating Steps 2 and 3 to work back and build the historical pattern of agebased stock sizes and fishing mortalities. -----------------With either VPA or cohort analysis, you do these calculations for each cohort in the fishery, and in this way, track each ones history of fishing mortalities. Caveat: assumption of constant M can be tough to estimate! (how would you do it?) and is it really constant?

Since we now know that N3 = 194, that N2 = 336, and M = 0.2, were ready to compute F2:

Yield-per-Recruit Models
120000 100000 80000

Change in population size

70.0 60.0

Change in Length (von Bertalanffy growth)

centimeters
0 5 10 15

50.0 40.0 30.0 20.0 10.0 0.0 0 5 10 15

Another class of age-based assessment is called the dynamic pool approach. In addition to population size, natural mortality, and fishing mortality, these keep track of fish growth and hence total biomass.

60000 40000 20000 0

Age (years)

Age (years)

X
Change in W eight (von Bertalanffy growth)
6.0 60000 50000

Change in Population Biomass

kilograms per fish

kilograms

Thus, a manager tries to select a fishing mortality rate that maximizes the yield (biomass) per recruit. Here is the general idea: consider a single cohort

5.0 4.0 3.0 2.0 1.0 0.0 0 5 10 15

=
Age (years)

40000 30000 20000 10000 0 0 5 10 15

Age (years)

kilograms

If you set fishing levels too high, too many fish will be taken before they have a chance to grow (growth overfishing). On the other hand, if you set F too low, you will have larger fish, but low biomass yields.

Change in Population Biomass

60000 50000 40000 30000 20000 10000 0 0 5 10 15

Fisheries managers use yield per recruit models to guide: 1. The size at which fish enter a fishery (are recruited) and 2. The amount of fishing effort, geared to produce the maximum harvests possible, subject to the constraints of conserving the stocks.

Age (years)

The optimal time to harvest is here.

Beverton and Holt, of stock-recruitment fame, came up with the idea that by regulating a parameter called

Beverton and Holts model expresses yields as the product of fishing mortality (Ft), stock abundance (Nt) and weight of fish (Wt) integrated over all fish-able ages (tc to the maximum age of the cohort, tmax)

tc = the age at first capture in a fishery,

and F, the fishing mortality rate, you can manipulate the biomass development in populations of fish. Thus, a fishery can potentially tune the population to produce large amounts of yield. (Beverton and Holt, 1957)

tmax

F N W dt
t t t

tc

The solution to this is nasty looking, but it can be used in a spreadsheet without too much difficulty:
3 Y U e nK ( tc t0 ) FW e M ( tc tr ) n (1 e( Z nK )( tmax tc ) ) R n 0 ( Z nK )

The models main advantage is that it can be used to estimate biomass yields by varying things that fisheries can be managed for, that is, F and tc (Q: how do you do this?) Disadvantages:

The values of Un are as follows: U0 = 1 U1 = -3 U2 = 3 U3 = -1

Assumption of steady state (all cohorts behave identically) No warnings of recruitment overfishing. Works best when M is low (M < 0.5, which is < 40%/yr).

Study this table!

A practical way to work through a yield per recruit example is given by Jennings et al. (2001). They create a spreadsheet of age, weight-at-age, the number of fish remaining after mortality, catch in numbers and also in terms of biomass, and finally, the remaining biomass in the living population.

Age (yrs) 1 2 3 4 5 6 7 8 9 10

W (kg) 0.6 0.9 2.1 4.1 6.3 8.4 10 11.2 12.6 13.5

N alive 100 44.9 20.2 9.1 4.1 1.8 0.8 0.4 0.2 0.1

Catch (#) Catch (kg) Pop Biom. 41.3 24.8 60.0 18.6 16.7 40.4 8.3 17.5 42.4 3.7 15.4 37.2 1.7 10.6 25.7 0.8 6.4 15.4 0.3 3.4 8.2 0.2 1.7 4.1 0.1 0.9 2.1 0.0 0.4 1.0 97.7 0.98 237 2.37

M 0.2 F 0.6 Z 0.8

Sum (kg) Sum per recruit

Catch is determined by the catch equation (F/ZN(1 e-Z) N alive is after fishing and natural mortality occur (i.e., Nt = Nt-1 e-(F+M) )

Once you create a table like the one in the last slide, you can run different scenarios by simply varying the values of F. You can then build up a picture of how F affects the yields, and how it affects what is left alive, for example:
Biomass Values of remaining F YPR (kg) per recr. 0 0.00 21.46 0.1 1.12 12.93 0.2 1.36 8.26 0.3 1.32 5.59 0.4 1.20 4.00 0.5 1.08 3.01 0.6 0.98 2.37 0.7 0.89 1.93 0.8 0.83 1.63 0.9 0.77 1.42 1 0.73 1.26
Yield per recruit model
25.00 20.00 15.00 10.00 5.00 0.00 0 0.2 0.4 0.6 0.8 1 Values of F
Biomass remaining per recr. YPR (kg)

You can even create a 3-D set of possible yield-per-recruit outcomes, by varying the size at recruitment (here indexed as Lc/L)

1.60 1.40 1.20 1.00 0.80 0.60 0.40 0.20 0.00

Yields per recruit

Biomass remaining

http://www.fishbase.org/manual/English/fishbaseyieldperrecruit_analyses.htm

Eggs per Recruit (EPR) method: Literature cited and further reading: developed by John Boreman (ESF grad!) very similar to Yield per Recruit, only instead of looking at potential yields, it estimates the total, potential, lifetime egg production of females this is a way of estimating the sustainability of a population when it is fished can compare EPR values among different stocks, different species, etc.
Beverton, R.J.H., and S.J.Holt. 1957. On the Dynamics of Exploited Fish Populations. Ministry of Agriculture, Fisheries and Food, London. Fry (1949) Gulland, J.A. 1965. Estimation of mortality rates. Annex to Arctic Fisheries Working Group Report (ICES, C.M. Doc #3). Jennings, S., M.J. Kaiser, J.D. Reynolds. 2001. Marine Fisheries Ecology. Blackwell Science, London. Pope, J.G. 1972. An investigation of the accuracy of virtual population analysis. ICNAF Research Bulletin 9:65-74.