MULTISPECIES MODELS

All of the models that we have examined so far have been based on the notion that we have a
fishery harvesting from a single stock of fish. This is a gross oversimplification of the real situation
in most fisheries. Most types of fishing gear catch a variety of fish species and most fishing vessels
operate in a variety of fisheries, with the consequence that few fisheries can fairly be classified as
being exclusively single species. Furthermore, there are likely to be ecological linkages between
the various species harvested, either because the organisms feed on each other or because they
share a common food source. As you might imagine, the models for multispecies fisheries can be
very complicated. We will examine a few examples of simple multispecies models to illustrate the
general principles involved in analyzing a multispecies fishery.

Technical Interactions in Multispecies Fisheries

Technical interactions between fish stocks occur when several stocks share the same
geographic region and are caught together by the fishing gear, but the stocks do not exert any
strong biological or ecological influence over each other. That is, the biological interactions among
the stocks are sufficiently small that they can be ignored. In salmon fisheries this is often described
as the problem of a mixed stock fishery.

To illustrate the problem consider the following hypothetical example, from Ricker (1958), on the
Supplemental Reading list. We have three salmon stocks having different spawner-recruit
relationships, but all with the same unexploited equilibrium stock size.

Seq
R=S

B
Recruits

Spawners

To proceed further we must express the equilibrium catch as a function of the exploitation rate. For
a Ricker SR model the equilibrium stock size Seq is related to the exploitation rate µ according to
the following

ln a⋅ ( 1 − µ )
Seq =
b

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and the equilibrium catch Ceq is related to µ according to

ln a⋅ ( 1 − µ )  ln a⋅ ( 1 − µ )  − ln a⋅ ( 1 − µ )

Ceq = R ( Seq) − Seq = a⋅ ⋅ exp −b⋅ 
b  b  b

ln a⋅ ( 1 − µ ) ln a⋅ ( 1 − µ ) ln a⋅ ( 1 − µ )  µ 

Ceq = − = ⋅ 
b⋅ ( 1 − µ ) b b 1 − µ

µ MSY
For our three salmon stocks we
get the following picture of
equilibrium catch versus
exploitation rate.
A+B+C

Equilibrium Catch
Notice that the stocks have
different maximum equilibrium
catches (MSY) and that they
C
occur at different exploitation
rates. If we set our exploitation
rate so as to maximize the
B
combined catches, we would end
up eliminating the least
productive stock. To guard
A
against this problem many of our
salmon stocks are managed to
protect the weakest stock.
Exploitation Rate

The Supplemental Reading list contains some other references on the topic of mixed-stock
fisheries. Paulik, Hourston and Larkin (1967) derive a solution for the exploitation rate that gives
MSY in a mixed stock salmon fishery. Ricker (1973) discusses the general problems associated
with mixed stock fisheries and draws attention to the so-called fishing up period experienced in
many fisheries. Initial harvests from an unexploited stock often are much higher than sustainable
catches. When this occurs it can give the fishers and the managers false expectations about how
productive the fishery will be in the future.

The problems associated with fishing on mixed-stocks are not limited to semelparous species like
Pacific salmon. Clark (1985) in chapter 5 illustrates the problem with a simple two-species
differential equation model,

dX dY
= GX ( X) − qX⋅ f⋅ X and = GY ( Y) − qY⋅ f⋅ Y
dt dt

Species X and Y are ecologically separate. The growth function of X is independent of Y and vice
versa. However, the two species are caught together.

At equilibrium we have the following relationships.

dX
=0 ==> Gx ( X) = qx⋅ f⋅ X
dt

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 dY
=0 ==> Gy ( Y) = qy⋅ f⋅ Y
dt

Gy ( Y) Gx ( X)
so the following condition must hold, =f= .
q y⋅ Y q x⋅ X

If both species grow according to the logistic growth model, then we have


ry⋅ Y⋅ 1 −
Y 
rx⋅ X⋅  1 −
X
 
 Ky 
=f=
 Kx 
q y⋅ Y q x⋅ X

ry  Y rx  X
⋅ 1 −  =f= ⋅ 1 − 
qy  Ky  qx  Kx 

Y qy qy r x  X
1− = ⋅f = ⋅ ⋅ 1 − 
Ky ry r y qx  Kx 

Y qy qy r x  X
=1− ⋅f = 1 − ⋅ ⋅ 1 − 
Ky ry r y qx  Kx 

 qy   qy r x  X 
Y = Ky⋅  1 − ⋅ f = Ky⋅ 1 − ⋅ ⋅ 1 − 
 ry   r y qx  Kx 

ry
Let f approach and observe that Y goes to zero and that the expression
qy

 qy r x  X   r y qx 
1 − ⋅ ⋅  1 −  also goes to zero, which means that X goes to Kx⋅  1 − ⋅  .
 r y qx  Kx   r x qy 

Kx

Ky
Species Y

  Species X

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 rx ry
In the graph I have assumed that > so that X remains greater than zero.
qx qy

What would cause fishing effort to increase to the level ry/qy and beyond? Open access fishing
effort for species X will result in X being less than

 r y qx   r y qx 
Kx⋅ 1 − ⋅  provided c < px⋅ qx⋅ Kx⋅  1 − ⋅ 
 r x qy   r x qy 

Costs per unit fishing effort must be less

than revenue per unit fishing effort.

Brander (1988), on the Recommended Reading list, gives some case histories for trawl fisheries in
the Irish Sea. One species of ray has apparently been eliminated from the region even though
they are only caught incidentally. Another example is a fishery for flatfish that targets sole but also
catches plaice. The price for sole is about five times the price for plaice. There are quotas on
plaice landings,but they are ineffective. Catches beyond the quota are simply discarded at sea. In
the trawl fisheries here in the Pacific Northwest we have similar problems with discarding of our
managed species.

Murawski (1984), on the Supplemental Reading list, contains a yield-per-recruit analysis for a
mixed species trawl fishery. The idea is that mesh size and the rate of fishing mortality can in
theory be adjusted to obtain a better species mix. The analysis ignores the problem of different
prices for different species, however.

Biological Interactions in Multispecies Fisheries

For most fish populations the factors contributing to population regulation (somatic growth,
mortality, reproduction) depend to some degree on the abundance of other species. In general, a
population's growth rate will depend on the abundance of prey, competitors, and predators.
Single-species models, such as the Schaefer surplus-production model, ignore these
complications. Krebs (1985), on the Recommended Reading list, in the chapters entitled "Species
interactions - competition" and "Species interactions - predation", provides a good introduction to
the basic concepts and mathematical equations associated with modeling a collection of interacting
species. We now will examine a fisheries example from May et al. (1979), on the Recommended
Reading list. The "fish" species involved are Antarctic krill, which is a small euphausid that forms
vast and dense schools in the waters surrounding Antarctica, and Antarctic baleen whales that
feed almost exclusively on Antarctic krill.

May et al. model the unexploited krill population dynamics using the following modified logistic
growth model.

dNk  Nk 
= rk⋅ Nk⋅  1 −  − a⋅ Nk⋅ Nw
dt  Kk 

where Nk denotes the abundance of krill and Nw denotes the abundance of whales. Consumption
of krill per whale occurs at rate a·Nk, proportional to krill abundance.

In the graph on the next page the diagonal line represents the consumption of krill by whales. It
has a slope of a·Nw. If there were no whales present, the krill population would be at equilibrium at
Kk, the carrying capacity when there are no whales.

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 Change in Krill Abundance

Krill Abundance Kk

The whale dynamics, in the absence of any harvesting, are assumed to be governed by the
following differential equation.

dNw  Nw 
= rw⋅ Nw⋅  1 − 
dt  α ⋅ Nk 

The carrying capacity for the whales is the quantity α·Nk. It is proportional to the abundance of krill.
The intrinsic growth rate for the whales is unaffected by the abundance of krill.

Rather than working with the equations as they are given above, May et al. put the equations into
an equivalent, but dimensionless form. This same technique can be applied to advantage with
most differential equation models. May et al. define the following two new variables:

Nk Nw
Xk = and Xw =
Kk α ⋅ Kk

dXk 1 dNk
The new differential equation for the krill is = ⋅ .
dt Kk dt

dXk  Nk    Nk   Nk 
= r k⋅   ⋅ 1 −   − a⋅   ⋅ Nw = rk⋅ Xk⋅ ( 1 − Xk) − a⋅ Xk⋅ Nw
dt  Kk    Kk   Kk 

We can reformulate the last term using a new parameter (ν) to transform the NW

dXk  1 ⋅ a⋅ α ⋅ K  ⋅  Nw 
= rk⋅ Xk⋅ ( 1 − Xk) − rk⋅ Xk⋅  k  
dt  rk   α ⋅ Kk 
ν Xw

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 dXk
and simplify the differential equation to = rk⋅ Xk⋅ ( 1 − Xk − ν ⋅ Xw) .
dt

dXw 1 dNw
For the whales the new differential equation is = ⋅ .
dt α ⋅ Kk dt

dXw Nw  Nw   Nw  Kk   Xw 
= r w⋅ ⋅ 1 −  = rw⋅ Xw⋅ 1 − ⋅   = rw⋅ Xw⋅ 1 − 
dt α ⋅ Kk  α ⋅ Nk   α ⋅ Kk  Nk   Xk 

Now we have simplified both differential equations and reduced the number of parameters from five
(rk, rw, Kk, a, and α) to three (rk, rw, and ν). We were able to do this because in the original
equations some of the parameters were redundant. For example, the parameters α and K k always
occurred together as a product.

Now let's add harvesting to the models for both krill and whales. These two species are always
harvested independently; they are never caught together, as was the case with the species in the
earlier models of technical interactions.

We will use scaled fishing mortality coefficients, which are dimensionless.

Fk Fw
Fishing for krill: F'k = Fishing for whales: F'w =
rk rw

F k and Fw are the usual instantaneous rates of fishing mortality. The only realistic values for the
new scaled fishing mortality coefficients occur over the range (0,1).

F' = 1 ==> F=r ==> Extinction

When we add the scaled fishing mortalities to the model we get

dXk
for the krill = rk⋅ Xk⋅ ( 1 − F'k − Xk − ν ⋅ Xw)
dt

dXw  Xw 
and for the whales = r w⋅ Xw⋅  1 − F'w − 
dt  Xk 

By deriving isoclines for the krill (combinations of Xk and Xw for which dXk/dt = 0) and for the whales
(combinations of Xk and Xw for which dXw/dt = 0) we can generate the phase plane diagram for this
system and explore how the system will change over time.

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The isocline for the krill is

Xk = 0 This does not tell us

dXk
 anything very useful.

dt
=0 ==>  or

 Xw = ( 1 − F'k − Xk) ⋅
1
This is an equation
ν for a straight line.

The isocline for the whales is

Xw = 0
dXw

dt
=0 ==>  or

 Xw = ( 1 − F'w) ⋅ Xk Another straight line.

On the phase plane diagram the isoclines look like the following

(1-F' k)/ν
Relative Whale Abundance

The krill isocline intersects

←
the whale axis at (1-F'k)/ν
↓
and has slope equal to
(-1/ν). The whale isocline
passes through the origin
and has slope equal to →
(1-F'w). The point where ↓ ↑
the two isoclines intersect is ←
the only equilibrium point
for this system. ↑
→

Relative Krill Abundance (1-F'k)

dXk dXw
At the equilibrium point neither population is changing = = 0.
dt dt

Xw = ( 1 − F'k − Xk) ⋅ Xw = ( 1 − F'w) ⋅ Xk

1
==> and
ν

To get equations for the equilibrium values for Xk and Xw we can solve for Xk in the second equation
(which comes from the whale isocline) and substitute into the result the right-hand side of the first
equation (which comes from the krill isocline).

− −

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 Xew 1 − F'k − Xek
Xek = =
1 − F'w (1 − F'w)⋅ ν

Xek⋅ ( 1 − F'w) ⋅ ν = ( 1 − F'k − Xek) Xek⋅ 1 + ( 1 − F'w) ⋅ ν = 1 − F'k

1 − F'k (1 − F'w)⋅ (1 − F'k)

Xek = Xew = ( 1 − F'w) ⋅ Xek =
1 + ( 1 − F'w) ⋅ ν 1 + ( 1 − F'w) ⋅ ν

The graphs below indicate what happens to the system as either of the fishing mortality coefficients
change.

Increased Fishing on the Krill. Increased Fishing on the Whales.

Relative Whale Abundance

Relative Whale Abundance

Relative Krill Abundance Relative Krill Abundance

With increased fishing on the krill both the krill and the whale populations decline. Krill harvests
reduce krill abundance and the whale carrying capacity. With increased fishing on the whales,
however, the whale population declines but the krill population increases. Whale harvests reduce
whale abundance, which lessens the amount of predation by the whales on the krill.

The equilibrium yields are

rk⋅ Kk⋅ F'k⋅ ( 1 − F'k)

Y_k = F k⋅ N_k = ( rk⋅ F'k) ⋅ ( Kk⋅ X_k) =
1 + ( 1 − F'w) ⋅ ν

α ⋅ rw⋅ Kk⋅ ( 1 − F'k) ⋅ F'w⋅ ( 1 − F'w)

Y_w = F w⋅ N_w = ( rw⋅ F'w) ⋅ ( α ⋅ Kk⋅ X_w) =
1 + ( 1 − F'w) ⋅ ν

The diagram on the next page shows krill yield as a function of the scaled krill fishing mortality
(increasing to the right) and the scaled whale fishing mortality (increasing to the left).

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 Relative Yield of Krill.

In addition to this simple two-species model May et al. examine two models with three interacting
species: a model with one prey (krill) and two competing predators (whales and seals) and a model
with three trophic levels (krill that are eaten by squid, that are eaten by sperm whales).

There is no end to how complicated these multi- species models can become, but there is a limit to
our ability to thoroughly anayze and understand why a model behaves as it does. There is little
point to building a model that is as complicated as the real system that it is supposed to mimic.

Models that use coupled differential equations, such as the above model for whales and krill and
the earlier bioeconomic model for a fish stock and fishing fleet, are often described as dynamical
systems. Clark (1976), on the Supplemental Reading list, provides additional mathematical
background for dynamical systems and methods for their analysis.

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