Wako

ARBAMINCH UNIVERSITY
SCHOOL OF POST GRADUATE STUDIES

COLLAGE OF NATURAL SCIENCE
DEPARTMENT OF MATHEMATICS(OPTIMIZATION)
A THESIS ON:
OPTIMAL HARVESTING OF TWO PREYS AND A
PREDATOR SPECIES
By
WAKO KURA ADI
ADVISOR: SIMON DARKEE (PHD)
SEPTEMBER, 2021
ARBAMINCH, ETHIOPIA
ARBA MINCH UNIVERSITY
SCHOOL OF GRADUATE STUDIES
DEPARTMENT OF MATHEMATICS
Certificate
This is to certify that the thesis entitled “Optimal harvesting of two preys
and a predator species” in partial fulfillment of requirements for the award of
the masters of science (MSc.) in Mathematics(Optimization) submitted to the
department of Mathematics, Arbaminch university carried out by WAKO KURA
(I.D PSNS/1689/09) under my supervision. Therefore I recommend that the
student’s Thesis can be presented for review and open oral presentation.
ARBA MINCH UNIVERSITY SIMON D. ZAWKA(PHD)

Advisor,
ARBA MINCH
Department of Mathematics
September, 2021 Arba Minch University.
i
Declaration
I here by declare that this MSc. Specialty or equivalent thesis dissertation is my
original work and has not been presented for a degree in any other university,
and all sources of material used for this thesis / dissertation have been duly
acknowledged.
Name: WAKO KURA ADI
Signature: ———————————————-
Date: —————————————————-
ii
APPROVAL
As member of Board of Examiners of the final MSc. Graduate thesis open defense
Examination, we certify that we have read and evaluate this graduate thesis
prepared by WAKO KURA entitled by “ Optimal harvesting of two preys and
a predator species” and recommended that it can be accepted as fulfilling the
thesis requirement for the degree of MSc. In Mathematics (Optimization).
APPROVED BY
Advisor
Name :Dr.Simon Darkee Signature ————————– Date—————-
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iii
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iv
Acknowledgements
First and for most, I am indebted to my advisor Dr.SIMON DARKEE from the
department of mathematics, for his inspiration, encouragement, humble guidance
and patience by constructive comments and great contributions in training me
how to use Latex software. His tireless efforts and patience in advising me are
greatly appreciated.
Moral support from my parents is also acknowledged. The presence of my family

throughout the period of this effort has made the hard time of my work less
stressful. Special thanks to my wife Kabale Galgalo and my sons, Kura Wako and
Guyo Wako, for their love, support, and understanding.
Finally, I acknowledge the AMU Department of Mathematics who gave me the

chance to do this thesis.
WAKO KURA
v
Abstract
This thesis deals with optimal harvesting of two preys and a predator species
where three species are subject to combined harvesting. The model is based on
Lotka -Volterra dynamics with two competing species which are affected not only
by harvesting but also by the presence of a predator, the third species. Preys
populations are competing each other (eg. competition for space and food).
Therefore, the problem of harvesting two competing species in the presence of
a predator species which feeds on both the competing species where studied.
We are consider constant harvesting and proportional harvesting for the species.
The possibility of existence of equilibrium is discussed, we verify positivity and
boundedness of the solution for the considered model. Finally, some numerical
examples are discussed.
vi
Contents
Certificate i
Declaration ii
Acknowledgements v
Abstract vi
Contents vii
1 Introduction 1
1.1 Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2 Statement of the Problem . . . . . . . . . . . . . . . . . . . . . . . 4
1.3 Objective of the Study . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.3.1 General Objective of the Study . . . . . . . . . . . . . . . . 4
1.3.2 Specific Objectives of the Study . . . . . . . . . . . . . . . . 5
1.4 Significance of the Study . . . . . . . . . . . . . . . . . . . . . . . . 5
1.5 Delimitation of the Study . . . . . . . . . . . . . . . . . . . . . . . 6
2 Literature Review 7
2.1 Logistic growth Equation . . . . . . . . . . . . . . . . . . . . . . . . 9
2.2 Basic Lotka -Volterra Model . . . . . . . . . . . . . . . . . . . . . . 10
2.3 prey-predator model . . . . . . . . . . . . . . . . . . . . . . . . . . 16
2.3.1 Constant Harvesting . . . . . . . . . . . . . . . . . . . . . . 22
2.3.2 Proportional Harvesting . . . . . . . . . . . . . . . . . . . . 24
2.3.3 Maximum sustainable yield . . . . . . . . . . . . . . . . . . 28
2.4 Bioeconomic Models . . . . . . . . . . . . . . . . . . . . . . . . . . 29
2.5 Optimal harvesting . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
3 Research Methodology 34
3.1 Description of the study . . . . . . . . . . . . . . . . . . . . . . . . 34
3.2 Research design . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
vii
4 Mathematical and Ecological Preliminaries 36
4.1 Nonlinear Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
4.2 Equilibrium Points and their Stability . . . . . . . . . . . . . . . . . 37
4.3 Basic Growth Models . . . . . . . . . . . . . . . . . . . . . . . . . . 38
4.3.1 Logistic growth Equation . . . . . . . . . . . . . . . . . . . . 38
4.3.2 Basic Lotka -Volterra Model . . . . . . . . . . . . . . . . . . 39
4.3.3 prey-predator model . . . . . . . . . . . . . . . . . . . . . . 45
4.4 Harvest Models . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
4.4.1 Constant Harvesting . . . . . . . . . . . . . . . . . . . . . . 50
4.4.2 Proportional Harvesting . . . . . . . . . . . . . . . . . . . . 51
4.5 Maximum sustainable yield . . . . . . . . . . . . . . . . . . . . . . 56
4.6 Bionomic Equilibrium . . . . . . . . . . . . . . . . . . . . . . . . . 57
4.7 Optimal harvesting . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
5 Results and Discussion 62

5.1 Assumptions in the Model . . . . . . . . . . . . . . . . . . . . . . . 62
5.2 The existing Model . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
5.3 Model analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
5.3.1 Existence of equilibrium . . . . . . . . . . . . . . . . . . . 64
5.3.2 Stability Analysis . . . . . . . . . . . . . . . . . . . . . . . . 67
5.4 Optimal harvesting Policy . . . . . . . . . . . . . . . . . . . . . . . 71
6 Numerical Simulations 74
7 Conclusion and Recommendations 77

7.1 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
7.2 Recommendations . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
References 79
viii
CHAPTER 1
Introduction
1.1 Background
The exploitation of natural resources such as fisheries and forestries

has gained importance in recent years. The techniques and issues
associated with the bioeconomic exploitation of these resources have
been discussed in detail by Clark [16]. Since most marine fisheries
are essentially multi species in nature, exploitation of mixed-species
fisheries has started to draw attention from researchers. The
population of human being has grown enormously in the past two
centuries. Billions of people use up resources quickly as they eat
food, build houses, produce goods, and burn fuel for transportation
and electricity. The continuation of life as we know it depends on
the careful use of natural resources.
On the other hand, People often misuse the natural resources.

Species are over harvested. Marine resources are depleted. Forests
are cleared, exposing land to wind and water damage. Fertile soil
is exhausted and lost to erosion because of poor farming practices.
If resources are carelessly managed, many will be used up. If used
1
Chapter 1. Introduction
wisely and efficiently, however, renewable resources will last much

longer. Through conservation, people can reduce waste and manage
natural resources wisely.
The need to conserve resources often conflicts with other needs. A

timber company may want to harvest the area’s trees for construction
materials. A business may want to build a factory or shopping hall
on the land. All these needs are valid, but sometimes the plants
and animals that live in the area are forgotten. When we use the
environment in ways that ensure we have resources for the future, it is
called sustainable development. There are many different resources
we need to conserve in order to live sustainable[12][13].
Exploitation of a renewable resource such as a fish has attracted

our attention. Exploitation of natural resource, though it hardly
encompasses protection is also natural. Mathematical population
models have been used to study the dynamics of prey predator
systems since Lotka (1925) and Volterra (1927) proposed the simple
model of prey-predator interactions now called the Lotka -Volterra
as result, the study of optimal harvesting of prey-predator.
Modeling is frequently evolving process, to gain a deep understanding

of the mathematical aspects of the problem and to yield non
trivial biological insights; one must carefully construct biologically
meaningful and mathematically tractable population models (Kiang,
2002). Some of the aspects that need to be critically considered in a
realistic and plausible mathematical model include; carrying capacity
which is the maximum number of prey that the ecosystem can sustain
in absence of predator, competition among prey and predators which
can be intra specific or inter specific, harvesting of prey or predators
and functional responses of predators.
2
Chaudhuri [7] formulated an optimal control problem for the combine

harvesting of two competing species. Most of the mathematical
models on the harvesting of a multi species have so far assumed
that the species are affected by harvesting only. To the researcher
knowledge, no fully adequate attempt has been made to study a
harvesting model of two competing species in the presence of a
predator, the third species.
Therefore, the problem of harvesting two competing species in the

presence of a predator species which feeds on both the competing
species where studied. A combined harvesting effort is devoted to the
exploitation of the first two (prey) species and the third (predator)
species. We analyzed the existence and stability of the equilibria
of the system. Harvesting models such as; constant harvesting,
proportional harvesting and so on is a few well known in literature
that deals with the optimal harvesting of two preys and a predator
(such as a fish)[8].
In this work, the researcher gives more attention to the model

associated with the Lotka -Volterra dynamics with two competing
species which are affected not only by harvesting but also by the
presence of a predator, the third species.
3
1.2 Statement of the Problem
The thesis seeks to address the problem of determine the optimal

harvesting of two preys and a predator. The prey populations
competing for resource. There is a predator which survives on this
preys.
We assume that the predator is low commercial value. The

harvesting activity does not affect the predator population directly
unless some special effort is explicitly put in to bring about a change
in the predator population. But the harvesting activity does reduce
the predator population indirectly by reducing the availability of the
preys to the predator.
Similarly the presence of a predator in the system indirectly reduces

the catch to the harvesting agency (here after stated as agency)
resulting in lower returns to the agency. Hence our problem is
harvesting fishery model with competition to analyze the effects of
the predator species in an ecosystem.
1.3 Objective of the Study
1.3.1 General Objective of the Study
The main objective of this study is to analysis mathematical model

on optimal harvesting of two preys and a predator species with
diametrical goals to harvest as much as possible while allowing
optimal population growth.
4
1.3.2 Specific Objectives of the Study
The specific objectives of this study will be:
✓ To verify boundedness and positivity of a modified model.
✓ perform the existence and stability analysis of the equilibrium

points.
✓ To discuss maximum sustainable yield.
✓ To discuss optimal harvesting strategy and bioeconomic

equilibrium.
✓ To give numerical simulation to demonstrate main findings of

the study.
1.4 Significance of the Study
The following are the importance of doing this thesis/ research/
✓ To provide potentially relevant information for agency whose

income is based on resource harvesting.
✓ To create the decision support tools which should help us to

understand human behavior within complex system in which
the prices of harvested object play an important role on one
hand the availability of the object is important on the other side.
5
✓ To provide preliminary information for other researcher.
1.5 Delimitation of the Study
The delimitation of this study is to investigating the optimal

harvesting of two preys in the presence of a predator in content.
We are not consider a primary data due to some constraints(i.e time,

budget and so on.)
6
CHAPTER 2
Literature Review
Ra o et al.[21] investigated analytically the dynamical behavior of the

model for three species with one prey and two competing predators.
they also discussed the policy of optimal harvesting. They used the
functional response of predator that it is limited by density of prey.
Kar (2003) studied a prey-predator system with delay, Hollings

Type II functional response and harvesting of the prey. The study
showed that as harvesting effort increased, the predator’s population
decreased as expected. This showed that harvesting of the prey alone
indirectly affected the population density of the predators and also
played a crucial role in stabilizing the dynamics of the prey-predator
systems[26].
Chaudhuri and Kar (2004) proposed and analyzed a fishery model

of a two prey-one predator system in which the prey were being
harvested and the feeding rate of the predator increases linearly
with prey density. They derived conditions for global stability of
the system using a Lyapunov function. Using Pontryagin’s maximal
principal, they established the conditions for optimal harvest. They
7
Chapter 2. Literature Review
also analyzed bioeconomic equilibrium solution and the policy of

optimal harvesting[8].
Green (2004) studied a model on two prey-one predator system in

which the prey were non competing and predation followed the
density gradient of the prey. It was discovered that when the
predator divides its time between the two preys depending on their
comparative density, the predator stabilized the system. The model
did not consider competition among prey and prey harvesting[9].
Das et al.[23] and Chaudhuri K . L [7] proposed the model one

prey-one predator with combined harvesting. They analyzed the
dynamical behavior of the model and the optimal harvesting policy.
They assumed that the growth of both prey and predator are
formulated in the logistic terms.
Kafue (1995) worked on a model on the interactions within a

four species ecosystem. The model was based on his research on
the interactions among the four main species in Queen Elizabeth
National Park (Uganda) namely; lions, hyenas, Uganda Cobs and
water bucks. The lions and hyenas were the predators while the
Uganda Cobs and the water bucks were the prey. Using the
Rout-Hurwitz criteria, he established conditions for stable existence
of all four species. He also investigated how mutual cooperation,
intraspecific and/or inter specific competition among species affected
the equilibrium of the ecosystem. However, the system was based
on the general linear prey-predator competition model and excluded
terms such as logistic growth with carrying capacity and functional
responses.
Optimal control theory deals with the problem of finding a control

law for a given system such that a certain optimality criteria is
8
achieved. It is an extension of the calculus of variations, and is a

mathematical optimization method for deriving control policies. The
method is largely due to the work of Lev. Pontryagin and Richard
Bellman in the 1950s.
A control problem includes a cost functional that is a function of

state and control variable. An optimal control is a set of differential
equations describing the paths of the control variables that minimize
the cost function.
In this literature review, focus is on optimal harvesting of two preys

and a predator species.
2.1 Logistic growth Equation
Exponential growth increases without bound. This is reasonable for

some situations; however, for populations there is usually some type
of upper bound. This can be caused by limitations on food, space
or other scarce resources. The effect of this limiting upper bound
is a curve that grows exponentially at first and then slows down
and hardly grows at all (ref. Figure 4.1). This is characteristic of a
logistic growth model. The Logistic growth equation is defined by
dx
= F (x), (2.1.1)
dt
where the function F is given by
x
F (x) = rx(1 − ). (2.1.2)
K
9
x
F (x) = rx(1 − K)
Growth function F (x)
0
0 Resource x K K
2
Figure 2.1: The figure highlights the solution curve for Logistic growth
equation. For the given initial state x0 > 0, the stock approaches the
environmental carrying capacity K asymptotically
2.2 Basic Lotka -Volterra Model
Mathematical modeling and analysis of multiple species ecological

problems was first done by Volterra (1927). Volterra had been
introduced to an ecological problem that in the years after the first
World War, the proportion of the predatory fishes caught in the
Upper Adriatic Sea was found to be considerably higher than in the
years before the war, where as the proportion of prey fishes was
down.
The Prey-Predator model with linear per capita growth rates is
x· = (b − py)x
y · = (rx − d)y
This system is referred to as the Lotka -Volterra model: it represents

one of the earliest models in mathematical ecology. The parameter b
10
is the growth rate of species x (the prey) in the absence of interaction

with species y (the predators).
Prey numbers are diminished by these interactions: The per capita

growth rate decreases (here linearly) with increasing y , possibly
becoming negative. The parameter p measures the impact of
x.
predation on . The parameter d is the death (or emigration) rate
x
of species y in the absence of interaction with species x . The term
rx denotes the net rate of growth (or immigration) of the predator
population in response to the size of the prey population.
Lotka -Volterra proposed the competition equations as follows:

dN1 N1
= r1 N1 (1 − )
dt K1
dN2 N2
= r2 N2 (1 − )
dt K2
The competition in these equations intra-specific. When the
species compete with each other (for nesting sites, food, etc.) the
intra-specific competition is detrimental to both specie’s per capita
growth rates. Then, the Lotka -Volterra competition model assumes
logistic growth for the population of each species to account for the
intra-species competitions, and that the growth rates are reduced
due to inter-specific competition.
The mathematical model for two competing species is:

dN1 N1
= r1 N1 (1 − ) − a12 N1 N2
dt k1
(2.2.1)
dN2 N2
= r2 N2 (1 − ) − a21 N1 N2
dt k2
11
The complete characterization of equation(2.2.1)revolves around the

orientation of the zero-growth isocline’s.
Figure 2.2: The figure highlights phase potrait for a12 > kk21 , a21 < kk12 . N2
rapidly excludes N1 . Species 2 approaches its carrying capacity.
12
Figure 2.3: The figure highlights phase potrait for a12 < kk21 , a21 > kk12 . N1
13
Figure 2.4: The figure highlights phase portrait for a12 > kk21 , a21 > kk21 . N2
zero growth isocline cross the N1 zero growth from above. Inter Specific effects
are large for both species.
14
Figure 2.5: The figure highlights phase portrait for a12 < kk21 , a21 < kk21 . Inter
specific effects were weak relative to intra specific effects. The two competing
species coexist.
15
2.3 prey-predator model
Predator-prey models are arguably the building blocks of the bio

and ecosystems as biomasses are grown out of their resource masses.
Species compete, evolve and disperse simply for the purpose of
seeking resources to sustain their struggle for their very existence.
Depending on their specific settings of applications, they can take the
forms of resource-consumer, plant-herbivore, parasite-host, tumor
cells (virus)-immune system, susceptible-infectious interactions, etc.
They deal with the general loss-win interactions and hence may have
applications outside of ecosystems. When seemingly competitive
interactions are carefully examined, they are often in fact some forms
of predator-prey interaction in disguise.
Kar,T.K and Chaudhuri, K.S propose harvesting in a two-prey

one-predator fishery. They make two prey species have the same
growth functional response of the predator to draw a precise
comparison based on the model.
dx1 x1
= r1 (1 − ) − β1 y − q1 E1 − σx2 (1 − q2 E2 )
dt k1
dx2 (1 − q2 E2 x2 )
= r2 [1 − ] − β2 y − σx1 (2.3.1)
dt k2
dy
= η1 β1 x1 + η2 β2 (1 − q2 E2 )x2 − m
dt
The three species are subject to the positive initial conditions. The
feeding rate of the predator species is assumed to increase linearly
with prey density. In their paper in which the harvesting function for
prey x1 adopts the traditional form and the other harvesting function
for prey x2 takes the new form.
16
They find that the new form of harvesting function refines the effects
of human intervention, which shows that harvesting prey affects not
only the growth of prey population but also the growth of predator
population[8].
Figure 2.6: The periodic activity generated by the predator-prey model.
17
Figure 2.7: Prey-Predator dynamics as described by the level curves of a

conserved quantity. The arrows describe the velocity and direction of solutions.
In this simulation, the data are d = r = b = d = 1 . There are equilibria at
x = 1, y = 1 and at x = 0, y = 0.
18
Liu(1994) propose the dynamical behaviors of a two prey one

predator system. In the absence of the predator the prey population
density grows according to logistic law of growth. Two prey species
are competitive in nature and there is reaction time for predator.
The model is given by;
dX X
= r(1 − ) − a1 XY − ω1 XZ
dt k
dY Y ω2 Y Z
= s(1 − ) − a2 XY − (2.3.2)
dt L (m + Y )
dZ Y (t − τ )Z
= b1 ω1 X(t − τ )Z + b2 ω2 − cZ
dt m + Y (t − τ )
Where r and s the intrinsic growth rate of two preys, k and L are
their carrying capacities, c is the mortality rate coefficient of the
predator, a1 , a2 are inter-species interference of two prey species,
ω1 and ω2 are first and second prey species searching efficiency of
predator, b1 and b2 are conversion factors denoting the number of
newly born predator for each captured of first and second prey
m is half saturation co-efficient. A discrete time delay(τ ≥ 0) is
introduced to the functional response term involved with the growth
equation of predator to allow for a reaction time.
In this paper Hollings type I response function is taken to represent

the interaction between one of the prey and predator. The
interaction between the other prey and the predator is assumed to
be governed by a Hollings type II response function. Such difference
of functional response may be particularly useful when handling
time for one prey is negligible, whereas the predator needs sufficient
handling time for other prey.
19
Figure 2.8: Kermack-McKendrick model of propagation of infectious disease.

The tipping point is at x = 1.0 . If x(0) is above this value, an epidemic will
ensue. The severity can be estimated by tracing the curve emanating from
x(0) until it converges to the horizontal axis. This indicates the size of the
susceptible population that avoids the infection. Note that the horizontal axis
as drawn here begins at x = 0.1 to avoid the logarithmic singularity.
20
Figure 2.9: Crocodiles are some of the evolutionarily oldest and dangerous
predators
21
2.3.1 Constant Harvesting
Constant harvesting is where a fixed number of resource stock was

removed each year [20]. Let h (t) = h represents the constant
rate of harvesting. Now, using the Logistic growth equation for a
single-species we have the following harvesting model.
dx x
= rx(1 − ) − h, x(0) > 0. (2.3.3)
dt K
The equilibrium stock levels associated with constant harvesting can
be seen. we observe that, for a given constant yield Y = h, the
system admits two equilibria, which are x1 and x2 . Where x1 is
unstable and x2 is stable in a sense that x(t) → x2 as t → ∞, in the
immediate presence of x2 . From these expressions we conclude that
constant harvest rate h is sustainable when,
h < maxx G(x) and x0 > x1 .
But if h > maxG(x) then population will reduce to extinction with in

dx
finite time because is always negative. Therefore our prediction in
dt
this harvest model is sufficiently low harvest rate can be sustained
for ever, while harvest rates that are too high will, if maintained,
gradually exterminate population.
22
Yield
Y =h
Unstable
equilibrium Natural
growth rate
Stable
equilibrium
0 x1 x2
0 K x
Figure 2.10: The graph of a constant harvesting
23
2.3.2 Proportional Harvesting
B. Dubeya, Peeyush Chandrab, Prawal Sinhab conducted a research

fishery in different zones, one reserved and the other is not (free
fishing zone). In modeling, they considered no fishing is permitted
in the reserved zone while the unreserved area is an open-access
fishery zone. Let N(t) and M(t) be the respective biomass densities
of the same fish population inside the unreserved and reserved
areas, respectively, at a time t. Let the fish subpopulation of the
unreserved area migrate into reserved area at a rate p1 and the fish
subpopulation of the reserved area migrate into unreserved area at
a p2 rate. Let E be the total effort applied for harvesting the fish
population in the unreserved area.
They assume that in each zone growth of fish population follows

logistic model. Keeping these in view, the dynamics of fish
subpopulations in unreserved and reserved areas may be governed
by the following autonomous system of differential equations[15]
dN N
= rN (1 − ) − p1 N + p2 M − qEN
dt K (2.3.4)
dM M
= sM (1 − ) − p2 M + p1 N
dt L
r and s are constants and known as intrinsic rate of
growth. Where K and L are the respective carrying capacity and N
and M are population size of unreserved and reserved zone of fish
species respectively, q is the catch ability coefficient of fish species
in the unreserved area.
Consider a population of fish that is growing logistically and that is

being harvested. Let us assume that the catch of fish per unit effort
24
is proportional to the stock level x,

dx x
= rx(1 − ) − qEx, x(0) > 0, (2.3.5)
dt K
Where x(t) is population size of species and q is the catch ability
coefficient of species. The parameters r, q and k are assumed to be
positive constants. q represents the proportion of the current stock
N caught by one standard vessel in one time unit.
The harvest rate is the product of three terms: the fishing effort E, a
proportionality constant q that measures catch ability, and the stock
level x.
h = qEx (2.3.6)
The product of the catch ability and of the effort, qE, is the fishing
mortality; it has the same dimensions as r and will play an important
role in what follows. h is the harvest rate both being measured in
tonnes per unit time.
The ideas behind this assumed catch relation are, first that catch
rates should be higher if the fish population is larger, and second
that the total catch rate should increase in proportion to fleet size.
It has been found that the total user’s cost of harvest per unit of
effort equals to the discounted value of the future marginal profit of
the effort at the steady-state level. It has also been noted that if the
discount rate increases, then the economic rent decreases and even
may tend to zero if the discount rate tend to infinity. Thus, it has
been concluded that high interest rate will cause high inflation rate.
Fishery management is the balance between harvesting and its

ecological implications[1] .
25
In fishery management, it is important to fish in such a way that a

species is sustainable and not in danger of becoming endangered
or going extinct. They do not want to over harvest small fish
populations. The key idea in their study was implementing different
fishing strategies to maximize catch in a sustainable manner. The
idea that supports the importance of optimal harvesting model
is raised by[14, 15]. Species throughout the oceans currently
have different population sizes; there are some populations above
equilibrium and some below.
These factors should be taken into consideration when constructing

a harvest strategy. They have equilibria, x∗ , whenever the growth
rate of the fish population equals the harvest rate:
∗ ∗ x∗
qEx = rx (1 − ) (2.3.7)
K
There are two equilibrium points.
qE
x∗ = K(1 − ) and x∗ = 0 (2.3.8)
r
that gives
qE
Y = qEx∗ = qEK(1 − ) (2.3.9)
r
26
Natural Y = qEx
growth rate
Yield
Stable
equilibrium F (x)
0 x x2
0 K
Figure 2.11: Graph of the proportional harvesting
27
2.3.3 Maximum sustainable yield
The foundation for the concept of MSY was laid down in the 1930s
by Russel (1931). Since then, there have been many comprehensive
accounts written of its use and misuse [12, 13]. MSY as defined
by Colin Clark is the largest harvest rate that can be sustained
indefinitely. It is based on differential equation that relate growth
rate with harvest rate assuming that harvest rate is always less
than maximum growth rate of the population and the initial
population size before harvesting begin is greater than the stock
size at equilibrium point. This strategy can results in sufficiently
low harvest rate sustained for ever, and if the harvest rate become
too high, and maintained for long time gradually the population will
be extinct.
(MSY) in case of constant harvest rate is hM SY = maxx G(x) on

x(t). Harvesting rate at any point of population size different from
the point at which maximum growth rate occur yields rapid decrease
in population size. It is also really difficult to make census on
population size at a given time interval because of different biological
and environmental changes like mortality, migration etc. Rather,
what the management will do is estimating population size which
can not be certain.
Because of presence of nuisance species in ecosystem, optimal

harvesting strategies are still in difficulty to management. That is
why currently, the predominant biological reference point for U.S.
fisheries management is the maximum sustainable yield (MSY) of
each individual species in an ecosystem however it mitigates [11].
In 1977, Peter Larkin published his now-famous paper,’An epitaph
for the concept of maximum sustained yield’. Larkin criticized the
28
concept of single maximum sustained yield (MSY) for many reasons.

First, there is a possibility that it may not guard against recruitment
failure. Second, it is impossible to maximize sustainable yield for all
species simultaneously.
For constant harvesting model MSY is obtained at

E
x∗ = (1 − )K (2.3.10)
r
and for proportional harvesting, MSY calculated will occur when
dY 2qE
= qK(1 − )=0 (2.3.11)
dE r
and it is obtained at
qE
x∗ = (1 −
)K (2.3.12)
r
where q is the constant of proportionality. Therefore
rK r
M SY = and EM SY = (2.3.13)
4 2q
2.4 Bioeconomic Models
Bioeconomic Models are integrated economic-ecological models

with all the advantages and disadvantages of such models, Most
bioeconomic modeling seeks appropriate levels of stock and catch
to assist resource managers, normally with environmental conditions
assumed constant[16].
Let us discuss the Gordon-Schaefer model which is based on logistic

growth equation:
X
F (X) = rX(1 − ), (2.4.1)
k
29
where F(X) is surplus biomass growth per unit of time : X is stock

biomass. The harvest rate(H) is assumed by the simple relation of
Schaefer catch function,
H(E, X) = qEX (2.4.2)
where E is fishing effort and q is a constant catch ability coefficient.

Sustainable yield occurs when harvest equals the surplus growth.
dX
= F (X) − H(E, X) = 0. (2.4.3)
dt
This implies
X
qEX = rX(1 − ).
k
Therefore, biomass at equilibrium is
qE
X = k(1 − ) (2.4.4)
r
Inserting (2.4.4) into (2.4.2) gives the long-term catch equation,
qE q 2 kE 2
H(E) = qkE(1 − ) = qkE − (2.4.5)
r r
Dividing both sides of(2.4.5) by effort (E) gives the linear relationship
between catch per unit of effort(CPUE) and fishing effort:
H q 2 kE
CP U E = = qk = (2.4.6)
E r
Assuming constant price, equation (2.4.5) can be used to define total
revenue (TR) in equilibrium as a function of standardized effort:
T R(E) = p.H(E) (2.4.7)
30
Where p denotes a constant price per unit of harvest. Total cost of

fishing effort (TC) is given by
T C(E) = c.E (2.4.8)
where c denotes unit cost of effort including opportunity cost of labor

and capital. From equations (2.4.7) and (2.4.8), the equilibrium
resource rent (⨿) can be derived as a function of fishing effort(E).
⨿ (E) = T R(E) − T C(E) (2.4.9)
Figure 2.12: The-standard-bio-economic-model
31
2.5 Optimal harvesting
In optimal harvesting policy for constant effort harvesting, a

population size x(t) which obeys the generalized logistic model is
stated by means of a constant effort, by the equation
dx x
= r(t)x[1 − ] − Ex (2.5.1)
dt K(t)
where r(t) and k(t) are both continuous period 1 functions and K(t)
R1
is positive and 0 rtdt > E > 0 for E is constant. In this case
h(x) = Ex is constant and named constant harvest model. But if
harvesting effort become a function of time t, it will be periodic and
E(t) > 0, piecewise continuous first order differentiable function,
h=E(t)x, thus in turn gives proportional harvest model. Hence
optimal harvesting become a result of optimal harvesting effort and
the optimal time-spectrum. In many papers, it is shown that optimal
policy is driven by applying time dependent logistic equation taking
in account of the effect of harvesting effort on the yield of single
species exploitation.
Meng Fan and ke wang has shown that the time-dependent

periodic Logistic equation has a unique positive periodic solution,
which is globally asymptotically stable for positive solutions, and
they obtain its explicit representation. Further, they choose the
maximum annual-sustainable yield as the management objective,
and investigated the optimal harvesting policies for constant harvest
and periodic harvest.
Now I am supposed to introduce the optimal dynamic optimization

model. If the control variable E(t) is used for 0 ≤ E(t) ≤ Emax
then the present value J of a continuous time-stream of revenues is
32
denoted by
Z ∞
J= e−δt [p1 q1 N1 + p2 q2 N2 + p3 q3 P − C]Edt (2.5.2)
0
where δ denotes the instantaneous annual rate of discount[8] is

wished to maximize subject to
dN1 N1
= r1 N1 (1 − ) − a12 N1 N2 − a13 N1 p − q1 EN1
dt k1
dN2 N2
= r2 N2 (1 − ) − a21 N1 N2 − a23 N2 p − q2 EN2 (2.5.3)
dt k2
dp p
= r3 p(1 − ) + a31 N1 N2 + a32 N2 p − q3 Ep
dt k3
for all t. Thus this is a dynamic optimization problem to be solved
in this study[2].
33
CHAPTER 3
Research Methodology
3.1 Description of the study
Here we studied the optimal harvesting of two preys and a predator

model. The mathematical model consists objective function that we
went to maximize and the constrained equation representing preys
and predator growth under harvesting of three species.
3.2 Research design
Once we formulate mathematical problem, next we perform stability

analysis of the equilibrium points and discuss optimal policy on
harvesting of two preys and a predator. It is also both interesting
and important to study the nature of various equilibrium solutions
associated with growth equation.
Finally, the problem is solved as an optimal control problem

and the optimal harvesting policy is discussed. For complicated
mathematical computation, we use computer algebra package like
mat lab or Mathematica. Furthermore, we use books, published
34
Chapter 3. Research Methodology
articles and related studies from Internet services as a source of

information for this study.
35
CHAPTER 4
Mathematical and Ecological Preliminaries
4.1 Nonlinear Systems
Consider the initial value problem
χ(t) = f (χ(t))
(4.1.1)
χ(t0 ) = χ0 .
where the set ψ ⊂ Rn is open, f : ψ −→ Rn be a map, and χ0 ∈ ψ.

Definition: An equilibrium point of the system (4.1.1) is a point χ∗
solution of the equation f (χ∗ ) = 0.
Definition: An equilibrium point χ∗ is said to be locally stable, if
any trajectory starting near to χ∗ it converges to χ∗ .
The local stability of χ∗ is determined by the sign of the eigenvalue
of the Jacobian matrix J at the equilibrium points.
36
Chapter 4. Mathematical and Ecological Preliminaries
4.2 Equilibrium Points and their

Stability
Let χ∗ be the equilibrium of system (4.1.1). The Jacobian matrix of

the system at χ∗ is given by
∂fi (χ∗ )

J(χ∗ ) = (4.2.1)
∂χi
Let λ1 , ..., λn be the eigenvalue of J. Which are the roots of the
characteristic polynomial:
p(λ) = λn + an−1 λn−1 + ... + a2 λ2 + a1 λ + a0 .
We have the following prepositions:
Proposition 1.
If the real parts of all eigenvalues are negative, then the

equilibrium point χ∗ is locally asymptotically stable.
If the real parts of all eigenvalues are positive, then the

equilibrium point χ∗ is unstable.
The limitation of the above proposition is that eigenvalues cannot

always be determined depending on the dimension of the system. In
this case we use the Routh-Hurwitz Criteria which informs on the
sign of the eigenvalues without providing them explicitly.
Theorem 1 (Routh-Hurwitz Criteria for Stability [reference]).
Consider the characteristic polynomial form:
p(λ) = λn + an−1 λn−1 + ... + a2 λ2 + a1 λ + a0 .
I. For n=1, the eigenvalue of the matrix J has negative real part
if and only if, a0 > 0.
37
II. For n=2, the eigenvalue of the matrix J has negative real part
if and only if; a0 , a1 > 0.
III. For n=3, the The eigenvalue of the matrix J has negative real
part if and only if; a0 , a1 , a2 > 0 and a1 a2 > a0 .
IV. For n=4, the eigenvalue of the matrix J has negative real part
if and only if; a0 , a1 , a2 , a3 > 0, a1 a2 > a0 a3 and
(a1 a2 − a0 a3 )a3 − a21 a4 > 0.
where J is the Jacobian matrix of the given system at χ∗ .
4.3 Basic Growth Models
4.3.1 Logistic growth Equation
Exponential growth increases without bound. This is reasonable for

some situations; however, for populations there is usually some type
of upper bound. This can be caused by limitations on food, space
or other scarce resources. The effect of this limiting upper bound
is a curve that grows exponentially at first and then slows down
and hardly grows at all (ref. Figure 4.1). This is characteristic of a
logistic growth model. The Logistic growth equation is defined by
dx
= F (x), (4.3.1)
dt
where the function F is given by
x
F (x) = rx(1 − ). (4.3.2)
K
38
x
F (x) = rx(1 − K)
Growth function F (x)
0
0 Resource x K K
2
Figure 4.1: The figure highlights the solution curve for Logistic growth
equation. For the given initial state x0 > 0, the stock approaches the
environmental carrying capacity K asymptotically
4.3.2 Basic Lotka -Volterra Model
Mathematical modeling and analysis of multiple species ecological

problems was first done by Volterra (1927). Volterra had been
introduced to an ecological problem that in the years after the first
World War, the proportion of the predatory fishes caught in the
Upper Adriatic Sea was found to be considerably higher than in the
years before the war, where as the proportion of prey fishes was
down.
The Prey-Predator model with linear per capita growth rates is
x· = (b − py)x
y · = (rx − d)y
This system is referred to as the Lotka-Volterra model: it represents

one of the earliest models in mathematical ecology. The parameter b
is the growth rate of species x (the prey) in the absence of interaction
with species y (the predators).
39
Prey numbers are diminished by these interactions: The per capita

growth rate decreases (here linearly) with increasing y , possibly
becoming negative. The parameter p measures the impact of
x·
predation on . The parameter d is the death (or emigration) rate
x
of species y in the absence of interaction with species x . The term
rx denotes the net rate of growth (or immigration) of the predator
population in response to the size of the prey population.
Lotka -Volterra proposed the competition equations as follows:

dN1 N1
= r1 N1 (1 − )
dt K1
dN2 N2
= r2 N2 (1 − )
dt K2
The competition in these equations intra-specific. When the
species compete with each other (for nesting sites, food, etc.) the
intra-specific competition is detrimental to both specie’s per capita
growth rates. Then, the Lotka-Volterra competition model assumes
logistic growth for the population of each species to account for the
intra-species competitions, and that the growth rates are reduced
due to inter-specific competition.
The mathematical model for two competing species is:

dN1 N1
= r1 N1 (1 − ) − a12 N1 N2
dt k1
(4.3.3)
dN2 N2
= r2 N2 (1 − ) − a21 N1 N2
dt k2
The complete characterization of equation(4.3.3)revolves around the

orientation of the zero-growth isocline’s.
40
Figure 4.2: The figure highlights phase potrait for a12 > kk21 , a21 < kk12 . N2
41
Figure 4.3: The figure highlights phase potrait for a12 < kk21 , a21 > kk12 . N1
42
Figure 4.4: The figure highlights phase potrait for a12 > kk21 , a21 > kk12 . N2
zero growth isocline cross the N1 zero growth from above. Inter Specific effects
are large for both species.
43
Figure 4.5: The figure highlights phase potrait for a12 < kk12 , a21 < kk21 . Inter
specific effects were weak relative to intra specific effects. The two competing
species coexist.
44
4.3.3 prey-predator model
Predator-prey models are arguably the building blocks of the bio-

and ecosystems as biomasses are grown out of their resource masses.
Species compete, evolve and disperse simply for the purpose of
seeking resources to sustain their struggle for their very existence.
Depending on their specific settings of applications, they can take the
forms of resource-consumer, plant-herbivore, parasite-host, tumor
cells (virus)-immune system, susceptible-infectious interactions, etc.
They deal with the general loss-win interactions and hence may have
applications outside of ecosystems. When seemingly competitive
interactions are carefully examined, they are often in fact some forms
of predator-prey interaction in disguise.
Liu(1994)propose the dynamical behaviors of a two prey one predator

system. In the absence of the predator the prey population density
grows according to logistic law of growth. Two prey species are
competitive in nature and there is reaction time for predator. The
model is
dX X
= r(1 − ) − a1 XY − ω1 XZ
dt k
dY Y ω2 Y Z
= s(1 − ) − a2 XY − (4.3.4)
dt L (m + Y )
dZ Y (t − τ )Z
= b1 ω1 X(t − τ )Z + b2 ω2 − cZ
dt m + Y (t − τ )
Where r and s the intrinsic growth rate of two preys, k and L are
their carrying capacities, c is the mortality rate coefficient of the
predator, a1 , a2 are inter-species interference of two prey species,
ω1 and ω2 are first and second prey species searching efficiency of
predator, b1 and b2 are conversion factors denoting the number of
45
newly born predator for each captured of first and second prey
m is half saturation co-efficient. A discrete time delay(τ ≥ 0) is
introduced to the functional response term involved with the growth
equation of predator to allow for a reaction time.
In this paper Hollings type I response function is taken to represent

the interaction between one of the prey and predator. The
interaction between the other prey and the predator is assumed to
be governed by a Hollings type II response function. Such difference
of functional response may be particularly useful when handling
time for one prey is negligible, whereas the predator needs sufficient
handling time for other prey.
Figure 4.6: The periodic activity generated by the predator-prey model.
46
Figure 4.7: Prey-Predator dynamics as described by the level curves of a

conserved quantity. The arrows describe the velocity and direction of solutions.
In this simulation, the data are d = r = b = d = 1 . There are equilibria at
x = 1, y = 1 and at x = 0, y = 0.
47
Figure 4.8: Kermack-McKendrick model of propagation of infectious disease.

The tipping point is at x = 1.0 . If x(0) is above this value, an epidemic will
ensue. The severity can be estimated by tracing the curve emanating from
x(0) until it converges to the horizontal axis. This indicates the size of the
susceptible population that avoids the infection. Note that the horizontal axis
as drawn here begins at x = 0.1 to avoid the logarithmic singularity.
48
Figure 4.9: Crocodiles are some of the evolutionarily oldest and dangerous
predators
49
4.4 Harvest Models
Prey-predator theory is traced from Malthus-verhust logistic

equation through the I - IV equation. Harvesting in predator-prey
systems has been of interest to economists, ecologists and natural
resource managers for some time now.
4.4.1 Constant Harvesting
Constant harvesting is where a fixed number of resource stock was

removed each year[20]. Let h (t) = h represents the constant
rate of harvesting. Now, using the Logistic growth equation for a
single-species we have the following harvesting model.
dx x
= rx(1 − ) − h, x(0) = x0 > 0. (4.4.1)
dt K
The equilibrium stock levels associated with constant harvesting can
be seen. we observe that, for a given constant yield Y = h, the
system admits two equilibria, which are x1 and x2 . Where x1 is
unstable and x2 is stable in a sense that x(t) → x2 as t → ∞, in the
immediate presence of x2 . From these expressions we conclude that
constant harvest rate h is sustainable when,
h < maxx G(x) and x0 > x1 .
But if h > maxG(x) then population will reduce to extinction with in

dx
finite time because is always negative. Therefore our prediction in
dt
this harvest model is sufficiently low harvest rate can be sustained
for ever, while harvest rates that are too high will, if maintained,
gradually exterminate population.
50
Yield
Y =h
Unstable
equilibrium Natural
growth rate
Stable
equilibrium
0 x1 x2
0 K x
Figure 4.10: The graph of a constant harvesting
4.4.2 Proportional Harvesting
Kar,T.K and Chaudhuri, K.S propose harvesting in a two-prey

one-predator fishery. They make two prey species have the same
growth functional response of the predator to draw a precise
comparison based on the model.
dx1 x1
= r1 (1 − ) − β1 y − q1 E1 − σx2 (1 − q2 E2 )
dt k1
dx2 (1 − q2 E2 x2 )
= r2 [1 − ] − β2 y − σx1 (4.4.2)
dt k2
dy
= η1 β1 x1 + η2 β2 (1 − q2 E2 )x2 − m
dt
The three species are subject to the positive initial conditions. The
feeding rate of the predator species is assumed to increase linearly
with prey density. In their paper in which the harvesting function for
prey x1 adopts the traditional form and the other harvesting function
for prey x2 takes the new form.
51
They find that the new form of harvesting function refines the effects
of human intervention, which shows that harvesting prey affects not
only the growth of prey population but also the growth of predator
population[8].
B. Dubeya, Peeyush Chandrab, Prawal Sinhab conducted a research

fishery in different zones, one reserved and the other is not (free
fishing zone). In modeling,they considered no fishing is permitted
in the reserved zone while the unreserved area is an open-access
fishery zone. Let N(t) and M(t) be the respective biomass densities
of the same fish population inside the unreserved and reserved
areas, respectively, at a time t. Let the fish subpopulation of the
unreserved area migrate into reserved area at a rate p1 and the fish
subpopulation of the reserved area migrate into unreserved area at
a p2 rate. Let E be the total effort applied for harvesting the fish
population in the unreserved area.
They assume that in each zone growth of fish population follows

logistic model. Keeping these in view, the dynamics of fish
subpopulations in unreserved and reserved areas may be governed
by the following autonomous system of differential equations[15].
dN N
= rN (1 − ) − p1 N + p2 M − qEN
dt K (4.4.3)
dM M
= sM (1 − ) − p2 M + p1 N
dt L
r and s are constants and known as intrinsic rate of
growth. Where K and L are the respective carrying capacity and N
and M are population size of unreserved and reserved zone of fish
species respectively, q is the catch ability coefficient of fish species
in the unreserved area.
52
Consider a population of fish that is growing logistically and that is

being harvested. Let us assume that the catch of fish per unit effort
is proportional to the stock level x,
dx x
= rx(1 − ) − qEx, x(0) > 0, (4.4.4)
dt K
Where x(t) is population size of species and q is the catch ability
coefficient of species. The parameters r, q, and k are assumed to be
positive constants. q represents the proportion of the current stock
N caught by one standard vessel in one time unit.
The harvest rate is the product of three terms: the fishing effort E, a
proportionality constant q that measures catchability, and the stock
level x.
h = qEx (4.4.5)
The product of the catchability and of the effort, qE, is the fishing
mortality; it has the same dimensions as r and will play an important
role in what follows. h is the harvest rate both being measured in
tonnes per unit time.
The ideas behind this assumed catch relation are, first that catch
rates should be higher if the fish population is larger, and second
that the total catch rate should increase in proportion to fleet size.
It has been found that the total user’s cost of harvest per unit of
effort equals to the discounted value of the future marginal profit of
the effort at the steady-state level. It has also been noted that if the
discount rate increases, then the economic rent decreases and even
may tend to zero if the discount rate tend to infinity. Thus, it has
been concluded that high interest rate will cause high inflation rate.
53
Fishery management is the balance between harvesting and its

ecological implications[1] .
In fishery management, it is important to fish in such a way that a

species is sustainable and not in danger of becoming endangered
or going extinct. They do not want to over harvest small fish
populations. The key idea in their study was implementing different
fishing strategies to maximize catch in a sustainable manner. The
idea that supports the importance of optimal harvesting model
is raised by[14, 15]. Species throughout the oceans currently
have different population sizes; there are some populations above
equilibrium and some below.
These factors should be taken into consideration when constructing

a harvest strategy. We have equilibria, x∗ , whenever the growth rate
of the fish population equals the harvest rate:
x∗
qEx∗ = rx∗ (1 − ) (4.4.6)
K
There are two equilibrium points.
qE
x∗ = K(1 − ) and x∗ = 0 (4.4.7)
r
that gives
qE
Y = qEx∗ = qEK(1 − ) (4.4.8)
r
54
Natural Y = qEx
growth rate
Yield
Stable
equilibrium F (x)
0 x x2
0 K
Figure 4.11: Graph of the proportional harvesting
55
4.5 Maximum sustainable yield
The foundation for the concept of MSY was laid down in the 1930s
by Russel (1931). Since then, there have been many comprehensive
accounts written of its use and misuse [12, 13]. MSY as defined
by Colin Clark is the largest harvest rate that can be sustained
indefinitely. It is based on differential equation that relate growth
rate with harvest rate assuming that harvest rate is always less
than maximum growth rate of the population and the initial
population size before harvesting begin is greater than the stock
size at equilibrium point. This strategy can results in sufficiently
low harvest rate sustained for ever, and if the harvest rate become
too high, and maintained for long time gradually the population will
be extinct.
(MSY) in case of constant harvest rate is hM SY = maxx G(x) on

x(t). Harvesting rate at any point of population size different from
the point at which maximum growth rate occur yields rapid decrease
in population size. It is also really difficult to make census on
population size at a given time interval because of different biological
and environmental changes like mortality, migration etc. Rather,
what the management will do is estimating population size which
can not be certain.
Because of presence of nuisance species in ecosystem, optimal

harvesting strategies are still in difficulty to management. That is
why currently, the predominant biological reference point for U.S.
fisheries management is the maximum sustainable yield (MSY) of
each individual species in an ecosystem however it mitigates [11].
In 1977, Peter Larkin published his now-famous paper,’An epitaph
56
for the concept of maximum sustained yield’. Larkin criticized the

concept of single maximum sustained yield (MSY) for many reasons.
First, there is a possibility that it may not guard against recruitment
failure. Second, it is impossible to maximize sustainable yield for all
species simultaneously.
For constant harvesting model MSY is obtained at

E
x∗ = (1 − )K (4.5.1)
r
and for proportional harvesting, MSY calculated will occur when
dY 2qE
= qK(1 − )=0 (4.5.2)
dE r
and it is obtained at
qE
x∗ = (1 − )K (4.5.3)
r
where q is the constant of proportionality.Therefore
rK r
M SY = and EM SY = (4.5.4)
4 2q
4.6 Bionomic Equilibrium
Bioeconomic Models are integrated economic-ecological models

with all the advantages and disadvantages of such models, Most
bioeconomic modeling seeks appropriate levels of stock and catch
to assist resource managers, normally with environmental conditions
assumed constant[16].
Let us discuss the Gordon-Schaefer model which is based on logistic

growth equation:
X
F (X) = rX(1 − ), (4.6.1)
k
57
where F(X) is surplus biomass growth per unit of time : X is stock

biomass. The harvest rate(H) is assumed by the simple relation of
Schaefer catch function,
H(E, X) = qEX (4.6.2)
where E is fishing effort and q is a constant catch ability coefficient.

Sustainable yield occurs when harvest equals the surplus growth.
dX
= F (X) − H(E, X) = 0. (4.6.3)
dt
This implies
X
qEX = rX(1 − ).
k
Therefore, biomass at equilibrium is
qE
X = k(1 − ) (4.6.4)
r
Inserting (2.4.4) into (2.4.2) gives the long-term catch equation,
qE q 2 kE 2
H(E) = qkE(1 − ) = qkE − (4.6.5)
r r
Dividing both sides of(2.4.5) by effort (E) gives the linear relationship
between catch per unit of effort(CPUE) and fishing effort:
H q 2 kE
CP U E = = qk = (4.6.6)
E r
Assuming constant price, equation(2.4.5)can be used to define total
revenue(TR)in equilibrium as a function of standardized effort:
T R(E) = p.H(E) (4.6.7)
58
Where p denotes a constant price per unit of harvest. Total cost of

fishing effort(TC) is given by
T C(E) = c.E (4.6.8)
where c denotes unit cost of effort including opportunity cost of labor

and capital. From equations (2.4.7) and (2.4.8), the equilibrium
resource rent (⨿)can be derived as a function of fishing effort(E).
⨿ (E) = T R(E) − T C(E) (4.6.9)
Figure 4.12: The-standard-bio-economic-model
59
4.7 Optimal harvesting
In optimal harvesting policy for constant effort harvesting, a

population size x(t) which obeys the generalized logistic model is
stated by means of a constant effort, by the equation
dx x
= r(t)x[1 − ] − Ex (4.7.1)
dt K(t)
where r(t) and k(t) are both continuous period 1 functions and K(t)
R1
is positive and 0 rtdt > E > 0 for E is constant. In this case
h(x) = Ex is constant and named constant harvest model. But if
harvesting effort become a function of time t, it will be periodic and
E(t) > 0, piecewise continuous first order differentiable function,
h = E(t)x, thus in turn gives proportional harvest model. Hence
optimal harvesting become a result of optimal harvesting effort and
the optimal time-spectrum. In many papers, it is shown that optimal
policy is driven by applying time dependent logistic equation taking
in account of the effect of harvesting effort on the yield of single
species exploitation.
Meng Fan and ke wang has shown that the time-dependent

periodic Logistic equation has a unique positive periodic solution,
which is globally asymptotically stable for positive solutions, and
they obtain its explicit representation. Further, they choose the
maximum annual-sustainable yield as the management objective,
and investigated the optimal harvesting policies for constant harvest
and periodic harvest.
Now I am supposed to introduce the optimal dynamic optimization

model. If the control variable E(t) is used for 0 ≤ E(t) ≤ Emax
then the present value J of a continuous time-stream of revenues is
60
denoted by
Z ∞
0
where δ denotes the instantaneous annual rate of discount[8] is

wished to maximize subject to
dN1 N1
= r1 N1 (1 − ) − a12 N1 N2 − a13 N1 p − q1 EN1
dt k1
dN2 N2
= r2 N2 (1 − ) − a21 N1 N2 − a23 N2 p − q2 EN2 (4.7.3)
dt k2
dp p
= r3 p(1 − ) + a31 N1 N2 + a32 N2 p − q3 Ep
dt k3
for all t. Thus this is a dynamic optimization problem to be solved
in this study[2].
61
CHAPTER 5
Results and Discussion
5.1 Assumptions in the Model
Let N1 (t) represent the population sizes of the first species at

time t, N2 (t) represent the population sizes of the second species
at time t and p(t) represent the population sizes of the third
species at time t.
Assume prey species follows the logistic growth model for

interactions within the species.
Model has a term for prey-predator interaction.
The differential equation for each species has a loss term for
inter-species competition.
Assume interaction between the species are represented by the

product of the species.
If two species compete for a single resource, then
1. Competitive exclusion - one species out competes the other

and becomes the only survivor.
62
Chapter 5. Results and Discussion
2. Coexistence - both species coexist around a stable

equilibrium.
We consider that all the species are harvested with harvesting

efforts E.
We assume that ri − qi E > 0 for i = 1, 2, 3.
All the parameters are assumed to be positive constant.
We assume here that the interior equilibrium point(N1∗ , N2∗ , p∗ )

exists.
Table 5.1: Model parameters and their description
Parameters Descriptions
r1 The natural growth rate of the first species
r2 The natural growth rate of the second species
k1 The carrying capacities of the first species
k2 The carrying capacities of the second species
a12 is rate of decrease of the first prey due to the competition with the second prey
a21 is rate of decrease of the second prey due to the competition with the first prey
a31 is rate of increase of the predator due to successful attacks on the first prey
a13 is rate of decrease of the first prey due to inhibition by the predator
a32 is rate of increase of the predator due to successful attacks on the second prey
a23 is rate of decrease of the second prey due to inhibition by the predator
q1 The catch-ability coefficient of the first species
q3 The catch-ability coefficient of the third species
q2 The catch-ability coefficient of the second species
E The harvesting effort
q1 E The harvesting rates of the first species at time
q2 E The harvesting rates of the second species at time
q3 E The harvesting rates of the third species at time
63
5.2 The existing Model
The governing equations of the system can be written as

dN1 N1
= r1 N1 (1 − ) − a12 N1 N2 − a13 N1 p − q1 EN1
dt k1
dN2 N2
= r2 N2 (1 − ) − a21 N1 N2 − a23 N2 p − q2 EN2 (5.2.1)
dt k2
dp p
= r3 p(1 − ) + a31 N1 N2 + a32 N2 p − q3 Ep
dt k3
Where q1 EN1 ,q2 EN2 and q3 Ep, are the catch-rate functions based
on the catch-per-unit-effort hypothesis.
5.3 Model analysis
In the section, we analyze the existence and the stability of equilibria.
5.3.1 Existence of equilibrium
We determine the conditions for the existence of the equilibrium

points of the system (6.0.1).
By equating the right hand side to zero, we obtain the equilibrium

states.
The possible equilibrium states are given as follows,
P0 (0, 0, 0) : The state of all washed out.
P1 (0, N2∗ , p∗ ) : The state in which the second prey and the predator
species survive and the first prey species extinct out.
64
Where
r3 r2
a23(−r3 +q3 E) + (r3 − q3 E) a32(r3 −q2 E) + (r3 − q3 E)
k3 k2
N2∗ = r2 r3 , p∗ = r2 r3
a32 a23 + a32 a23 +
k2 k3 k2 k3
r3
The equilibrium point is exists when: a23 (r3 − q3 E) <
k3 (r2 − q2 E).
P2 (N1∗ , 0, 0) : The state in which only first prey species survive.
qE
Where N1∗ = k1 (1 − 1 ) With r1 > q1 E
r1
P3 (0, N2 , 0) : The state in which only second prey species survive.
qE
Where N2∗ = k2 (1 − 2 ) With r2 > q2 E
r2
P4 (N1∗ , 0, p∗ ) : The state in which the first prey and the predator
species survive and the second prey species extinct out.
Where.
r3 r1
a13(−r3 +q3 E) + (r1 − q1 E) a31(r1 −q1 E) + (r3 − q3 E)
k3 k1
N1∗ = r1 r3 , p∗ = r1 r3
a31 a13 + a31 a13 +
k1 k3 k1 k3
r3
The equilibrium point is exists when: a13 (r3 − q3 E) <
k3 (r1 − q1 E).
P5 (N1∗ , N2∗ , 0) : The state in which the first prey and the second prey
species survive and the predator species extinct out.
We have the system:

r1
r1 − N1 − a12 N2 − q1 E = 0
k1
r2
r2 − N2 − a21 N1 − q2 E = 0
k2
Solve this system, Thus the non zero equilibrium point is
65
k1 [a12 k2 (r2 −q2 E)−r2 (r1 −q1 E)] k2 [a21 k1 (r1 −q1 E)−r1 (r2 −q2 E)]
N1∗ = a12 a21 k1 k2 −r1 r2 , N2∗ = a12 a21 k1 k2 −r1 r2
The equilibrium point is exists if and only if it satisfies the following

conditions.
i. a12 k2 (r2 − q2 E) > r2 (r1 − q1 E),

a21 k1 (r1 − q1 E) > r1 (r2 − q2 E) and a12 a21 k1 k2 > r1 r2 .
ii. a12 k2 (r2 − q2 E) < r2 (r1 − q1 E),

a21 k1 (r1 − q1 E) < r1 (r2 − q2 E) and a12 a21 k1 k2 < r1 r2 .
P6 (0, 0, p∗ ) : The state in which only second the predator survives,

two prey species are washed out.
k3 (r3 − q3 E)
p∗ =
r3
P7 (N1∗ , N2∗ , p∗ ) : The co-existence state, the state in which two prey
and predator species exists. The equilibrium points are the solution
of the equations
r1 N1 (1 − Nk11 ) − a12 N1 N2 − a13 N1 p − q1 EN1 = 0
r2 N2 (1 − Nk22 ) − a21 N1 N2 − a23 N2 p − q2 EN2 = 0
r3 p(1 − kp3 ) + a31 N1 N2 + a32 N2 p − q3 Ep = 0
This system of equations must be solved simultaneously.
1 r2 r3 1 r2 r3
N1∗ = [(−r1 +q1 E)(a32 a23 + )]+ [(−r1 +q1 E)(a32 a23 + )]+
L k2 k3 L k2 k3
1 r2 r3
[(−r1 + q1 E)(a32 a23 + )]
L k2 k3
1 r3 1 r1
N2∗ = [(r1 −q1 E)(a31 a23 + a21 )]+ [(r3 −q3 E)(a23 a23 −a21 a13 )]+
L k3 L k1
1 r1 r3
[−(r2 − q2 E)(a31 a13 + )]
L k1 k3
66
1 r2 a31 1 r1 a32
p∗ = [(r1 − q1 E)(a21 a32 − )] + [(r2 − q2 E)(a12 a31 − )] +
L k2 L k1
1 r1 r2
[(r3 − q3 E)(a21 a12 − )],
L k1 k2
where,
r2 a13 r1 a23 r1 a23
L = (a12 a23 − )a31 + (a21 a13 − )a32 + (a21 a13 − )a32
k2 k1 k1
5.3.2 Stability Analysis
Local stability
We study the local stability of each equilibrium point by finding the

eigenvalues of the variational matrix at the equilibrium point.
The variational matrix of the system at the equilibrium point

(N1∗ , N2∗ , p∗ ) is given by
∂f1 (N1∗ , N2∗ , p∗ ) ∂f1 (N1∗ , N2∗ , p∗ ) ∂f1 (N1∗ , N2∗ , p∗ )

 
 ∂N1 ∂N2 ∂p 
∗ ∗ ∗
 ∂f2 (N1∗ , N2∗ , p∗ ) ∂f2 (N1∗ , N2∗ , p∗ ) ∂f2 (N1∗ , N2∗ , p∗ ) 
V (N1 , N2 , p ) = 
 

 ∂N1 ∂N2 ∂p 
 ∂f3 (N1 , N2 , p ) ∂f3 (N1 , N2 , p ) ∂f3 (N1 , N2∗ , p∗ )
∗ ∗ ∗ ∗ ∗ ∗ ∗ 
∂N1 ∂N2 ∂p
 
V11 −a12 N1 −a13 P
= −a21 N2 V22 −a23 N2
 

a31 P a32 P (a31 N1 + a32 N2 − 2P )
Where
2r1 N1
V11 = (r1 − k1 − a12 N2 − a13 P − q1 E),
2r2 N1
V22 = (r2 − k2 − a21 N1 − a33 P − q2 E).
67
Stability at the origin (0,0,0)
The Variational matrix at a trivial equilibrium point (0, 0, 0) is given

by  
r1 − q 1 E 0 0
V0 (0, 0, 0) =  0 r2 − q2 E 0 
 
0 0 0
It is a diagonal matrix. Its eigenvalues are elements on the diagonal.
λ1 =r1 − q1 E, λ2 = r2 − q2 E and, λ3 = 0 are eigenvalues of V at the
trivial equilibrium point.
Hence the integral curves terminate in a plane corresponding to the
steady state.
Theorem 2. The equilibrium state p7 (N1∗ , N2∗ , p∗ ) is globally
asymptotically stable if,
r1 r2
(i) a13 = a31 , a23 = a32 , (ii) 4 > (a12 + a21 )2 .
k1 k2
Proof. To examine the globally stability of the system(6.1), we
define a lyapunov function
N1 N2 P
V (N1 , N2 , P ) = N1 − N1∗ ln( ) + N 2 − N ∗
2 ln( ) + P − P ∗
ln( )
N1∗ N2∗ P∗
We see that V is definite positive. The time derivative of V along
the solutions of the system (6.1),is given by,
dV N1 − N1∗ dN1 N2 − N2∗ dN2 P − P ∗ dP
= + + .
dt N1 dt N2 dt P dt
dV
After simplifying, can be rewritten as,
dt
dV
= −X T AX (5.3.1)
dt
where , X T = [N1 − N1∗ , N2 − N2∗ , P − P ∗ ], and A is
68
 
r1 1 1
− (a12 + a21 ) (a31 − a13 )
 k1 2 2 
 1 r2 1 
− (a12 + a21 ) (a32 − a23 )
 
2 k2 2
 
 
 1 1 r3 
(a31 − a13 ) (a32 − a23 )
2 2 k3
dV
We observe that < 0 if the matrix A is definite positive. The
dt
matrix A is definite positive if,
1 1
(a31 − a13 ) = 0, (a32 − a23 ) = 0, and
2 2
r1 r2
4 − (a12 + a21 )2 > 0.
k1 k2
So, we conclude that the co-existence equilibrium is globally
asymptotically stable, if a13 = a31 , a23 = a32 and
r1 r2
4 > (a12 + a21 )2 > 0.
k1 k2
Stability at (N1 ,N2 ,0)
When the predator is absent the interaction becomes a two-species

competition model.
where
k1 [a12 k2 (r2 − q2 E) − r2 (r1 − q1 E)]
N1 = , and
a12 a21 k1 k2 − r1 r2
k2 [a21 k1 (r1 − q1 E) − r1 (r2 − q2 E)]
N2 =
a12 a21 k1 k2 − r1 r2
The variation matrix at equilibrium point (N1 , N2 ) is given by:
" #
c1 −a12 N1
V3 =
−a21 N2 c2
69
2r1 N1
Where c1 = r1 − k1 − c12 N2 − q1 E
r1 N1 r1 N1
=r1 (1 − k1 ) − k1 − a12 N2 − q1 E
=- r1kN1 1 c2
2r2 N2
=r2 − k2 − a21 N1 − q2 E
r2 N2 r2 N2
=r2 (1 − k2 ) − k2 − a21 N1 − q2 E
=- r2kN2 2
The trace τ and the determinant ∆ of the matrix V3 are
τ = e1 + e2 < 0,
r1 r2 − k1 k2 a12 a21
∆ = λ1 .λ2 = ( )N1 N2
k1 k2
case 1: If r1 r2 < k1 k2 c12 c21 ,we have ∆ < 0, then (N1 , N2 ) is
unstable.
case 2: If r1 r2 > k1 k2 c12 c21 ,we have ∆ > 0, then (x∗ , y∗ ) is locally
asymptotically stable.
In this case the competition low , and both species can coexist.
Stability at (N1∗ ,N2∗ ,P ∗ )
∗
− r1kN1 1 −12 N1∗ −a13 N1∗
 
∗
V4 (N1 , N2∗ , P ∗ ) =  −a21 N2∗ − r2kN2 2 −a23 N2∗ 
 
a31 p∗ a32 p∗ −p∗

The characteristic equation for the V4 is b3 β 3 + b2 β 2 + b1 β + b0 =0.
Using the Routh-Hurwitz criteria, the stability of the system is given
by
r1 a12 a31 r2 a21 a32
> and >
k1 a32 k2 a31
70
5.4 Optimal harvesting Policy
Once the process of harvesting the resource is stated the problem

of management of the resources can be viewed in terms of rent
maximization. We have already discussed the dynamics of the system
governed by the equation (1)-(3) and now we are in a position to
apply maximum principle[8] to solve the optimization problem[2] and
obtain optimal harvesting policy.
We restate our problem to maximize the objective function

Z ∞
0
subject to the state equations (1)-(3) and the control constraints

0 ≤ E(t) ≤ Emax
The Hamiltonian for the problem is given by
H = e−δt [p1 q1 N1 + p2 q2 N2 + p3 q3 P − C]Edt

N1
+β1 (r1 N1 (1 − k1 ) − a12 N1 N2 − a13 N1 p − q1 EN1 )
N2
+β2 (r2 N2 (1 − k2 ) − a21 N1 N2 − a23 N2 p − q2 EN2 )
p
+β3 (r3 p(1 − k3 ) + a31 N1 N2 + a32 N2 p − q3 Ep),
Where βi , i = 1, 2, 3, are the adjoint variables.
When the shadow price equals the net economic revenues on a

unit harvest, that is µ(t) = 0, then the Hamiltonian H becomes
independent of the control variable E(t),
∂H
i.e = 0.
∂E
71
This is a necessary condition for the singular control E ∗ (t) to be

optimal over control set.
Thus ,the optimal harvesting policy is:


 E , µ(t) > 0
 max


E(t) = 0 , µ(t) < 0



E
∗ , µ(t) = 0
Since µ(t) causes E(t) to switch between the levels 0 and Emax , µ(t)
are called switching function.
We seek to find the optimal equilibrium solution of the problem,

hence N1 , N2 , P and E are to be treated as constants in the
subsequent steps.
The adjoint equations are

dβ1 ∂H
=-
dt ∂N1
β1 r1 N1
= - e−δt p1 q1 E + + β2 a21 N2 − β3 a31 P
k1
dβ2 β2 r2 N2
= - e−δt p2 q2 E + β1 a12 N1 + − β3 a32 P
dt k2
dβ3 ∂H
=-
dt ∂P
β3 r3 P
= - e−δt p3 q3 E + β1 a13 N1 + β2 a23 N2 +
k3
Now, by eliminating β1 and β3 from above equation, we get a reduced
differential equation for β2 as
(D3 + c2 D2 + c1 D + c0 )β2 = A2 e−δt , (5.4.2)
where
72
r1 r2 r3
c2 = -( N1 + N2 + N3 ),
k1 k2 k3
r1 r2 r1 r3
c1 = ( - a21 a12 )N1 N2 + ( + a31 a13 )N1 N3 +
k1 k2 k1 k3
r2 r3
( + a21 a12 )N2 N3 ,
k2 k3
r1 a32
c0 = (a32 a13 a21 - a23 + a31 a13 a23 )N1 N2 P +
k1
r3 r2 r1 r2 r3
( a21 - a31 a13 - )N1 N2 P
k3 k2 k1 k2 k3
r3
A2 = [(a32 a13 + a12 )N1 P + a12 δN1 ]p1 q1 E +
k3
r1 r1 r3
[(a31 a12 − a32 )N1 P − a32 δP ]p3 q3 E - ( + a31 a13 )N1 P p2 q2 E -
k1 k1 k3
r1 r3
( δN1 + δP + δ 2 )
k1 k3
We find the shadow prices βi (t)eδt , i = 1 , 2 , 3, of the three
fish species remain bounded as t −→ ∞ and hence satisfy the
transversality condition at ∞.
73
CHAPTER 6
Numerical Simulations
In this section, we present the simulation results to illustrate the

dynamical of the model. In our optimal problem, we have an
equilibrium solution that satisfies the necessary conditions of the
maximum principle[8]. Let us consider a set of values of parameters
as follows in appropriate units:
r1 =2.09, r2 =2.07, k1 =200, k2 =300, q1 =0.04, q2 =0.01, r3 =2.09,

a12 =0.001, a21 =0.001, a13 =0.01, a31 =0.3, a23 =0.02, a32 =0.3, p1 =6,
p2 =8, E=10, C=50 and δ=0.05.
74
Chapter 6. Numerical Simulations
Figure 6.1: Variation of the populations against time,beginning with

N1 =150,N2 =250 and p=130.
75
Chapter 6. Numerical Simulations
Figure 6.2: Phase-space trajectories corresponding to the optimal effort E=

11.4 units,with reference to differential initial levels. The trajectories indicate
that the optimal equilibrium p(91,102,58) is asymptotically stable
76
CHAPTER 7
Conclusion and Recommendations
7.1 Conclusions
In this thesis, we have considered the effects of harvesting in a

two-species competitive system in the presence of a predator species.
It has been assumed that the two species compete with each other
for common resources such as food and each species grow logistically
in the absence of the other species.
We have first studied the existence and stability (local) and discussed
the effect of the predator on two competing species in which
three species are harvested.The globally stability of the system in
co-existence state is examined by using Lyapunov function.
Next, the optimal harvest problem was discussed to obtain the

optimal equilibrium solution. The present value of a continuous
time-stream of revenues is maximized by invoking Pontryagin’s
maximum principle. It is found that the shadow prices remain
constant over time, so the boundedness and positivity condition are
satisfied. Also the total user cost of harvest per unit effort equals
77
Chapter 7. Conclusion and Recommendations
the discounted value of the future profit at the steady state effort
level.
7.2 Recommendations
Based on the results of model analysis; I recommended that, the

species should not be harvested at a rate higher than their intrinsic
growth rate. However optimal harvesting of the preys at a rate much
lower than their intrinsic growth rate is permissible, since this would
not lead to collapse of the system in the long term.
I believe that these thesis makes an important contribution to the

technical side of renewable resource harvesting models that look at
a time and also has policy implication. In the next research, will be
developed for interaction of predator and prey species based on the
limited density of prey population, in which it is more reasonable in
an ecological system.
78
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ARBAMINCH UNIVERSITY

SCHOOL OF POST GRADUATE STUDIES

ADVISOR: SIMON DARKEE (PHD)

and a predator species” in partial fulfillment of requirements for the award of

the masters of science (MSc.) in Mathematics(Optimization) submitted to the

department of Mathematics, Arbaminch university carried out by WAKO KURA

(I.D PSNS/1689/09) under my supervision. Therefore I recommend that the

ARBA MINCH UNIVERSITY SIMON D. ZAWKA(PHD)

Name: WAKO KURA ADI

Name of the candidate —————————————————————————-

Moral support from my parents is also acknowledged. The presence of my family

Finally, I acknowledge the AMU Department of Mathematics who gave me the

5 Results and Discussion 62

7 Conclusion and Recommendations 77

The exploitation of natural resources such as fisheries and forestries

On the other hand, People often misuse the natural resources.

wisely and efficiently, however, renewable resources will last much

The need to conserve resources often conflicts with other needs. A

Exploitation of a renewable resource such as a fish has attracted

Modeling is frequently evolving process, to gain a deep understanding

Chaudhuri [7] formulated an optimal control problem for the combine

Therefore, the problem of harvesting two competing species in the

In this work, the researcher gives more attention to the model

1.2 Statement of the Problem

The thesis seeks to address the problem of determine the optimal

We assume that the predator is low commercial value. The

Similarly the presence of a predator in the system indirectly reduces

1.3 Objective of the Study

1.3.1 General Objective of the Study

The main objective of this study is to analysis mathematical model

1.3.2 Specific Objectives of the Study

The specific objectives of this study will be:

✓ To verify boundedness and positivity of a modified model.

✓ perform the existence and stability analysis of the equilibrium

✓ To discuss maximum sustainable yield.

✓ To discuss optimal harvesting strategy and bioeconomic

✓ To give numerical simulation to demonstrate main findings of

1.4 Significance of the Study

The following are the importance of doing this thesis/ research/

✓ To provide potentially relevant information for agency whose

✓ To create the decision support tools which should help us to

✓ To provide preliminary information for other researcher.

1.5 Delimitation of the Study

The delimitation of this study is to investigating the optimal

We are not consider a primary data due to some constraints(i.e time,

Ra o et al.[21] investigated analytically the dynamical behavior of the

Kar (2003) studied a prey-predator system with delay, Hollings

Chaudhuri and Kar (2004) proposed and analyzed a fishery model

also analyzed bioeconomic equilibrium solution and the policy of

Green (2004) studied a model on two prey-one predator system in

Das et al.[23] and Chaudhuri K . L [7] proposed the model one

Kafue (1995) worked on a model on the interactions within a

Optimal control theory deals with the problem of finding a control

achieved. It is an extension of the calculus of variations, and is a

A control problem includes a cost functional that is a function of

In this literature review, focus is on optimal harvesting of two preys

2.1 Logistic growth Equation

Exponential growth increases without bound. This is reasonable for

Growth function F (x)

2.2 Basic Lotka -Volterra Model

Mathematical modeling and analysis of multiple species ecological

The Prey-Predator model with linear per capita growth rates is

This system is referred to as the Lotka -Volterra model: it represents

If the real parts of all eigenvalues are negative, then the

If the real parts of all eigenvalues are positive, then the