Stability Problem For Coupled Systems of

Differential Equations
(Lotka-Volterra Predator-Prey Model)

Keoagile Balekane Raborife (16528301)

Supervisor: Prof JMW. Munganga

Honours Research in Applied Mathematics

Department of Mathematical Science

University of South Africa

5 January 2023

1
Declaration
I Keoagile Balekane Raborife, hereby declare that all the information submitted is the product
of my efforts, both in conception and execution, and that all the sources that I have used or
quoted have been indicated and properly acknowledged through complete references.

2
 Abstract

Based on observational data collected in the Kruger National Park game reserve, we
construct the Lotka-Volterra (predator-prey equations) model to predict the interactions
between lions and wildebeest. The coefficients (parameters) for both species’ intrinsic
growth rates and the effectiveness of predators in converting food into fertility were de-
termined from the field data.
We alter the model by considering three lion cropping strategies: constant rate crop-
ping, cropping proportional to the population present, and cropping proportional to the
square of the population present. We see how the populations change from the phase
potraits in our figures from the alteration of the model, and we also discover intriguing
interpretations of how the populations of both species might develop.

3
Acknowledgment
I want to thank God for always being there with me, the Lord knows this was not an easy
journey to start and accomplish. To Mrs. E. Sibanyoni, thank you very much for helping me
to register on time to do my honors and offering to assist me with paying for my fees, am
forever grateful. To my beautiful mother, the best I could have ever asked for, thank you for
your prayers, your love which is the sweetest I’ve ever known, and your support throughout
my academic journey and life as a whole. To Ms. Dipuo Tikane, thank you for being the best
mentor and sister, your love for mathematics triggered me to also pursue mathematics. Also, to
prof Munganga my supervisor thank you for your feedback, and for directing me on the correct
path as to complete this project in time.

4
Contents
1 Introduction 7

2 Preliminaries 8
2.1 Systems of differential equations . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2.2 Locally Lipschitz functions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2.3 Well-posedness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.4 Equilibrium points . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2.5 Stability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2.6 Linearization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

3 The Mathematical Model 12

4 Species interaction 13

5 Stability analysis 15

6 Estimation of the parameters 16

7 Management of population through cropping 17

7.1 Cropping at a constant rate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
7.2 Cropping at a rate proportional to the present population . . . . . . . . . . . . . 18
7.3 Cropping at a rate proportional to the square of the population present . . . . . 19

8 Concluding remarks 20

Bibliography 21

Index 22

A Appendix 23

B Numerical solution code 25

5
List of Tables
1 Wildebeest, zebra and lion population densities since 1972 . . . . . . . . . . . . 23
2 Number of lions to be cropped to maintain a stable population using strategy
(12) over a year . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

List of Figures
1 Phase plot of a 2D predator-prey system for 3 initial conditions [4] . . . . . . . . 14
2 Numerical simulation of the Lotka-Volterra model [4] . . . . . . . . . . . . . . . 14
3 Numerical simulation of the Lotka-Volterra model [5] . . . . . . . . . . . . . . . 17
4 Trajectories for cropping at a constant rate R of 0.1 and 0.2 respectively [5] . . . 18
5 Trajectories for cropping proportional to the population R of 0.4 and 0.8 [5] . . 19
6 Trajectories for cropping proportional to the square population R of 0.4 and 0.8
[5] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

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K.B Raborife (16528301) HRAPM82 Assignment 10

1 Introduction
The Lotka-Volterra model was the first to explain interactions between two species, a predator
and a prey, living in the same ecosystem and it can be used to predict what the populations
of the two species over time would be like. A basic understanding of ecology is required to
comprehend the lotka-volterra model. When two species coexist in the same habitat, they
share the same resources. In ecology, two terms should be defined: habitat and ecological
niche. A habitat is a place where an organism lives, whereas an ecological niche is made up of
biotic factors, food sources, and the habits of the organisms as a whole [2]. As a result, the
niche is much more precise and specific than the habitat, which consists solely of biotic factors
and the biotic elements that live there, and as a result, the competition’s outcome will be either
a win or loss or partitioning [7].
In an ideal world, no two species can coexist in the same niche without becoming competitive.
As a result, if two species share the same niche, they should compete. That concept is known
in ecology as the competitive exclusion principle or Gause’s Law. According to Gause’s Law,
if the niche between two species is indifferent, the system will result in a competition. If
competition arises between these two species, one will win and the other will lose. Furthermore,
the successful species will live in that habitat, while the unsuccessful species will become locally
extinct. Partitioning, on the other hand, occurs when two species coexist in the same habitat
and share the same niche, but have limited resources [7].
The goal is to study the relationship between the prey and the predator by applying the Lotka-
Volterra model and differential equations. To characterize the relationship between the two
species, we investigate the interactions that occur between predator and their prey and try to
provide an alternative viewpoint on how these interactions should be described. The species’
connection will be outlined and figures illustrating the relationship between the species will be
provided.
We explain how to estimate the coefficients of the components in this system of nonlinear
first-order differential equations using information received from the field.
We discuss how the data obtained can be utilized to estimate the parameters of the terms in this
system of nonlinear first-order differential equations. The lion (Panthera leo) and wildebeest
(Connochaetes taurinus) population projections, kill rates, cropping rates, and calf survival
rates are all contained in the given dataset [8] that was gathered over six years in the Kruger
National Park (South Africa).

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2 Preliminaries
We introduce the basic concepts of stability and derive some results concerning the existence
and uniqueness of solutions.

2.1 Systems of differential equations

Definition 1 (Autonomous systems of differential equations) Let f¯ be a function de-
fined on a subset Rn with values in Rn . An autonomous system of a differential equation is an
equation of the form [9].
′
x (t) = f¯(x(t)) for all t in some interval. (1)
′
We will omit the t in the above equation and write the system as, x = f¯(x)
Definition 2 (Solution) A vector-valued function x(t), is a solution of a system of differential
equations if it is differentiable on some interval and satisfies the system of differential equations
on that interval [9].
Definition 3 (Initial Value Problem) Let D be a subset of Rn , b ∈ D, f¯ a function defined
on D with values in Rn and s is any real number. An initial value problem involves finding a
function x satisfying the differential equation [9].
′
x = f¯(x)
on some interval containing s ∈ R, with the condition x(s) = b.
This function x is called a solution to the initial value problem [9].

2.2 Locally Lipschitz functions

Definition 4 (Lipschitz function) Let f¯ : Rn → Rn be defined on a set D. f¯ is said to be
Lischiptz on D, if there exists a positive number K Lipschitz constant for f¯ such that
∥f¯(x) − f¯(y)∥ ≤ K||x − y∥
for any x and y in D[9].
Definition 5 (Open and closed ball) For every x ∈ Rn and very real number r > 0 the set
Br (p) = x ∈ Rn : ||x − p∥ < r}
is called an open ball with center p and radius r [9].
The set
B̄r (p) = x ∈ Rn : ||x − p∥ ≤ r}
is called a closed ball with center p̄ and radius r.
Definition 6 (Locally Lipschitz function) Consider a function f¯ : Rn → Rn defined on an
open set D. If, for each p ∈ D, there exists a ball B̄r (p) such that f¯ is Lipschitz on B̄r (p), then
f¯ is called locally Lipschitz on D [9].
Proposition 1 A function f¯ : Rn → Rn is locally Lipschitz on an open set D provided the
partial derivatives of its components are continuous on D.

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2.3 Well-posedness
Theorem 1 (Uniqueness) Consider the autonomous system x′ = f¯(x) and suppose the func-
tion f¯ is locally Lipschitz on its domain. If x and y are solutions of the system defined on the
interval (α, β) and x(s) = y(s) for some s ∈ (α, β), then x = y on (α, β) [9].
Remark 1 A function f¯ : Rn → Rn is locally Lipschitz on its domain D if the partial deriva-
tives of its components are continuous on its domain D.
Theorem 2 (Local existence) Consider a function f¯ : Rn → Rn defined on an open D in
Rn and consider the initial value problem

x′ (t) = f¯(x(t)) (2)

with x(s) = b [9].

If f¯ is locally Lipschitz on D and b ∈ D, then there exists a positive number δ and a function
x such that x is the solution of the initial value problem on the interval (s − δ, s + δ).
In the example below we look at our model 5 and we look to answer two questions; do solutions
exist and are the solutions unique? We make use of theorems 1 and 2 to investigate the problem.
Example 1 Consider the initial value problem x̄ = f¯(x, y), x̄(s) = b̄ with b̄ ∈ R2 . Suppose that
f¯ is locally Lipschitz on an open set D ∈ R2 .
Do solutions to the system exist?
The functions αx − βxy and δxy − γy are both continuous on R2 . By theorem 2, with the abuse
of notation, there exists δ1 > 0 and δ2 > 0 and solutions x and y for the IVP that is defined on
(s − δ1 , s + δ1 ) and (s − δ2 , s + δ2 ) respectively.
Are the solutions to the system unique?
Since f¯ is locally Lipschitz on R2 , thus by remark 1 the partial derivatives of f¯ are continuous on
R2 . Hence, by theorem 1, the solutions x and y defined above on (s−δ1 , s+δ1 ) and (s−δ2 , s+δ2 )
respectively are unique.
From the example above we observe that we expect solutions to the Lotka-Volterra model to
exist and also should be unique.

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2.4 Equilibrium points

Definition 7 (Equilibrium point) A point p is an equilibrium point of (1) if f¯(x) = 0 for
all t ≥ 0.
To find the equilibrium points of (5) we set the left-hand side of the equations to 0, that is,
x′ = 0 and y ′ = 0.

x′ = αx − βxy = 0,
y ′ = δxy − γy = 0

Thus we get that;

α
x(α − βy) = 0 ⇐⇒ x = 0 or α − βy = 0 ⇒ y = β
γ
y(δx − γ) = 0 ⇐⇒ y = 0 or δx − γy = 0 ⇒ x = δ

Hence the two possible equilibrium points for all values of t, are (x, y) = (0, 0) and (x∗ , y ∗ ) =
( γδ , αβ ).
The equilibrium point (x, y) = (0, 0) suggests that both of our species (lions and wildebeest)
are being extinct, and the point (x∗ , y ∗ ) = ( γδ , αβ ), corresponds to the population density at
which the size of the population for both species will remain.

2.5 Stability
Types of stability
Definition 8 Point p is stable if, for each ϵ > 0 and t0 ∈ R, there exists δ > 0 such that if
x(t) is a solution of (1) and ∥x(t0 ) − p∥ < δ then ∥x(t) − p∥ < ϵ for all t ≥ t0 .
The point p, if it is not stable then it is called unstable.
Definition 9 The point p is Asymptotically stable if it is stable and if for every t0 ∈ R, there
exists δ > 0 such that if x(t) is a solution of (1) and ∥x(t0 ) − p∥ < δ then

lim x(t) = p
t→∞

Definition 10 The point p is a Globally asymptotically stable equilibrium point of (1) if it is

stable and
lim x(t) = p for all x0 ∈ Rn
t→∞

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2.6 Linearization
In some cases, the behavior of the solutions of a non-linear system like the Lotka-Volterra model
(5) near an equilibrium point p can be resolved by studying a linear system.
Definition 11 (Jacobian matrix) Suppose the function f¯ defined on Rn → Rn is a vector
field.
The matrix is given by
 
∂1 f1 (p) ∂2 f1 (p) . . . ∂n f1 (p)
 ∂1 f2 (p) ∂1 f2 (p) . . . ∂n f2 (p) 
Jf¯(p) =  ..
 
.. .. .. 
 . . . . 
∂1 fn (p) ∂2 fn (p) . . . ∂n fn (p)

is called the Jacobian matrix of the vector field f¯ at the equilibrium point p.
Remark 2 If A = Jf¯(p), then the behavior of the solutions of nonlinear systems near the
equilibrium point p is approximately the same behavior as that of solutions of linear systems.
Theorem 3 Suppose the point p is an equilibrium point of the system x̄′ = f¯(x̄) .
(a) If all the eigenvalues of Jf¯(p) have negative real parts, then p is asymptotically stable.
(b) If any eigenvalue of Jf¯(p) has positive real parts, then p is unstable.
Theorem 4 Suppose the point p is an equilibrium point of the system x′ = f¯(x). If there is no
eigenvalue of the matrix Jf¯(p) that has a positive real part but there exists a zero eigenvalue
or eigenvalue with a zero real part, then the equilibrium point p can either be
i. asymptotically stable,
ii. stable but not asymptotically stable,
iii. unstable.
.

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3 The Mathematical Model

In this section, we construct a mathematical model of how two species, a predator and prey
may interact with each other. We make the following claims to develop the model [1]:
1. The food supply for the prey population is never-ending at all times.
2. In the exclusion of predators, the population of the prey x increases in proportion to its
size x′ = αx, α > 0. This coefficient α, was termed the coefficient of auto-increase by
Vito Volterra [1]. Integrating the equation x′ = αx, α > 0, the population of prey would
increase exponentially given by the equation x(t) = x0 exp(αt) with x0 being the initial
population size of the prey.
3. In absence of prey, the predator population y would decline in proportion to its size
y ′ = −γx, γ > 0. By integration, the predator population would decline exponentially
and eventually become extinct given by the equation y(t) = y0 exp(−γt) with y0 the initial
population size of the predator.
4. When both species are present, a decline in the population of the prey and an increase in
the population of the predator would be observed. This decline and growth of both these
species would occur at rates proportional to the number of times they come into contact
with each other. The rates of encounter for the prey and predators are −βxy and δxy
respectively, with both β, δ > 0.
The x and y variables represent the populations of wildebeest and lions respectively.
Putting these claims into place, our prey equation is

x′ = αx − βxy. (3)

This equation tells us that the change in the prey’s population depends on its growth minus
the rate at which it encounters the predator. And similarly, the predator equation is given by

y ′ = −γy + δxy. (4)

The equation shows that the change in the predator population is influenced mainly by the
food supply (the prey).
Thus, the Lotka-Volterra predator-prey model [6, 10] is given by the following system of differ-
ential equations:

x′ = αx − βxy,
y ′ = δxy − γy. (5)

with α, β, δ, γ > 0.

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4 Species interaction
The analysis of the relationship between the two species may be achieved by analyzing the
integrals of the individual populations using the separation of the variable method. Consider
the prey population equation:

dx
= x(α − βy).
dt
intergrating and simplifying the equation yield x(t) = A exp(α − βy)t, where A = exp(C) ∈ R.
Similarly the predator population equation

dy
= y(δx − γ).
dt
simplifies to y(t) = B exp(δx − γ)t where A = exp(C) ∈ R. From this, we see that the
population of the predator depends on the prey population.
dy
If x′ = dx
dt
and y ′ = dt
, then x′ = αx − βxy, y ′ = δxy − γy.
Thus
dy δxy − γy
= . (6)
dx αx − βxy
dy y(δx − γ)
= .
dx x(α − βy)
Cross multiply then divide by xy into both sides we get
α − βy δx − γ
dy = dx. (7)
y x

Integrating both sides of (7), we get the solution

α − βy δx − γ
Z Z
dy = dx
y x
α ln(y) − βy = δx − γ ln(x) + C, (8)
where C is the constant that depends on some given initial condition.
Equation (8) cannot be written in an explicit form. Thus, this gives us phase curves determined
by the value of C.
The curve formed by (8) with different values of C would give us both solutions x(t) and y(t).
And by choosing how the solution say x(t) behaves, then (8) would be used to determine how
the solution y(t) would in turn behave as seen in figure 1, and since our population size is
in 1000’s thus, to avoid using large quantities on our figures we use point decimals in our
code, where 0.1 represents 100 lions/wildebeests. With the initial conditions (x(0), y(0) =
(0.3, 0.3), (0.45, 0.45), (0.67, 0.67) for α = β = γ = δ = 3.
The connection between the two species is addressed in figure 2 underneath. The Lotka-Volterra
model was designed using Matlab according to the following parameters [3]: h = 0.1, tf =
20, (x(0), y(0)) = (20, 5), α = 10, β = 1, δ = 0.1, γ = 1.

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Figure 1: Phase plot of a 2D predator-prey system for 3 initial conditions [4]

.

Figure 2: Numerical simulation of the Lotka-Volterra model [4]

.

.
The phase portraits in Figure 1 clearly show the oscillation’s periodicity. The curves move
counterclockwise: when predator numbers are low, prey numbers increase, and when predator
numbers are high, prey numbers decrease. From Figure 2, we take note of how the predator
population lags behind the prey population: an increase in prey numbers results in a delayed
increase in predator numbers as the predators consume more prey.

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5 Stability analysis
We aim to determine and classify the stability of the equilibrium points of the Lotka-Volterra
population model, by making of linearization theory discussed above matrix. The equations of
our model are given by

x′ = f (x, y) = αx − βxy,
y ′ = g(x, y) = δxy − γy.

The Jacobian of our system is

 
∂ f (x, y) ∂y f (x, y)
Jf¯ = x
∂x g(x, y) ∂y g(x, y)
 
α − βy −βx
Jf¯ =
δy δx − γ
Under the equilibria section (2.4) we found that the equilibrium points of our predator-prey
model were (x, y) = (0, 0) and (x∗ , y ∗ ) = ( γδ , αβ )
Thus, at the point (x, y) = (0, 0):  
α 0
Jf¯ =
0 −γ

To find the eigenvalue(s) λ we consider the characteristic equation:

α−λ 0
det(Jf − λI) =
0 −γ − λ
(9)
= (α − λ)(−γ − λ)
=0

which for the two-by-two Jacobian matrix results in a quadratic equation for λ. Our eigenvalues
are α and -γ respectively.
Hence, By theorem 3, we can conclude that the equilibrium point (x, y) = (0, 0) is unstable.
Consider the equilibrium point (x∗ , y ∗ ) = ( γδ , αβ ):

0 −βx∗
 
Jf∗¯ =
δy ∗ 0

0 − λ − βγ
det(Jf∗ − λI) = δα δ
β
0
(10)
= λ2 + αγ
=0
√
the solutions are λ = ± αγ, and they are purely imaginary. When the eigenvalues of the
two-by-two Jacobian are purely imaginary, the equilibrium point is referred to as a center, and

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the perturbation oscillates and does not grow or decays. In this case, the angular frequency of
√
oscillation is ω = αγ and the period of the oscillation is 2π
ω
.
And from theorem 4 our equilibrium point (x∗ , y ∗ ) = ( γδ , αβ )can be either asymptotically stable,
or stable but not asymptotically stable or unstable.

6 Estimation of the parameters

In this section, we look at how the coefficients of the Lotka-Volterra model (5) are determined
from the field data gathered. The population of wildebeest and lions are denoted by x(t) and
y(t) respectively (measured in thousands) at time t.
The model’s parameters were determined as follows:
The average calf survival rate of α = 0.405 is the wildebeest’s intrinsic growth rate. This is
the mean of the calf survival rates recorded over six years. The likelihood of a lion-wildebeest
interaction resulting in a kill has been estimated to be around 80%.To make the statistics in
our model as simple as possible, we picked β = 81% = 0.81 likelihood since the critical value
yielded is exactly 500 lions, whereas 80% produces 506 lions [8].
The lions’ intrinsic growth rate is estimated to be δ = 1.5 cubs per year per adult female [8] p.7.
In the model, this value is taken to be negative since cubs die in the absence of prey. Again to
keep our model (11) as simple as possible, the age groups and the sex composition of the lion
population are not taken into consideration.
The predator’s efficiency to convert food into fertility is the ratio of the number of cubs, which
is 1.5, to the number of kills per adult lion, which is 4.5, multiplied by the ratio of adult females
1.5
to the pride, which is 0,375, so γ = 4.5 × 0.375 = 0.125. This is not the only model that can be
used to describe the interactions between wildebeest and lions, but when the solution curves of
this model are plotted in the population quadrant of the phase plane, they fit the data well.
From the information in [8], the possible Lotka-Volterra model is,

x′ = 0, 405x − 0, 81xy,
y ′ = 0, 125xy − 1, 5y. (11)

From the system, we get the equilibrium points (0, 0) and (12, 0.5). The equilibrium point (0, 0)
is the saddle point and it is not much of interest, and the point (12, 0.5) is the center and we
can see from Figure 3 that the trajectories revolve around the center point. The population of
the wildebeest is on the x-axis and the population of the lions on the y-axis both measured in
thousands.
.

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Figure 3: Numerical simulation of the Lotka-Volterra model [5]

.

7 Management of population through cropping

Between 1976 and 1986, a lion cropping program was implemented in the Kruger National
Park, as detailed by Smuts [?]. According to the report, the goal of cropping was to reduce
lion populations to the point where wildebeest and zebra populations could recover.
In this section, we look at three strategies that can be used to crop lions in a way that would
not lead to extinction.

7.1 Cropping at a constant rate

We make use of the model we developed (11) with the addition of constant cropping rate R to
the equation that relates the population of lions.

x′ = 0, 405x − 0, 81xy,
y ′ = 0, 125xy − 1, 5y − R.

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Figure 4: Trajectories for cropping at a constant rate R of 0.1 and 0.2 respectively [5]
.

The consequence of continual lion removal over time is catastrophic, as seen in figure 4. The
trajectories illustrated in this figure are for cropping constant rates of 0.1 and 0.2, respectively.
These trajectories spiral out and finally cross the x-axis, signaling lion extinction. This suggests
that employing this method is ineffective.

7.2 Cropping at a rate proportional to the present population

We consider our cropping rate R proportional to the population present:

x′ = 0, 405x − 0, 81xy,
y ′ = 0, 125xy − 1, 5y − Ry.

This model’s phase plane trajectories with R = 0.4 and R = 0.8 are illustrated in figure 5. The
trajectories reveal that the model’s equilibrium points are in the center, and the populations
move in a cyclic pattern. The steady-state population of lions remains constant at 500, but
the steady-state population of wildebeest grows as the R-value increases. However, Starfield
[8] suggests that it is impossible to accurately regulate the value of R over time, making this
method difficult to adopt in reality, and figure 5 shows that the model is particularly sensitive
to slight changes in R.

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Figure 5: Trajectories for cropping proportional to the population R of 0.4 and 0.8 [5]
.

7.3 Cropping at a rate proportional to the square of the

population present
We consider the model;

x′ = 0, 405x − 0, 81xy,
y ′ = 0, 125xy − 1, 5y − Ry 2 .

We can observe that the phase plane trajectories as shown in figure 6 for this model spiral inward
toward a fixed point when we plot the trajectories with the same R-values as in section 7.2.
This implies that, over time, both populations will tend towards stable population groupings,
and the drastic shift in nature of the critical points, from center when R = 0 to an attractive
spiral when R > 0 but not too big, indicates that bifurcation is occurring about this parameter.

Figure 6: Trajectories for cropping proportional to the square population R of 0.4 and 0.8 [5]
.

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8 Concluding remarks
Our Lotka-Volterra predator-prey model is a model that can be utilized to predict the behavior
of the two species but it is not 100% accurate, because the parameter values are expected to
change, thus the model would not fit the census data in Table 1 perfectly or over a long period.
As a result, our parameter estimates are only educated guesses at best. Starfield and Bleloch
[8] summarize this ‘we can never really validate the sort of model that we have built, we can
only hope to gain confidence in it.’
From the three strategies of cropping introduced, the first cropping strategy was unproductive
because, for example, no one who sets out to implement this strategy would continue to crop
a population once it was evident that the population was declining. As shown in Figure 4, if
this strategy was implemented, the lions would become extinct. Figure 5 shows that the second
strategy was also ineffective because of the difficulty of maintaining a consistent crop rate. The
third strategy was effective, and as shown in Figure 6, the populations of both species spiral
toward a stable population.
More questions arise with regards to our model and the alterations made to accommodate
cropping, questions such as:
ˆ How accurate are the parameter estimates in the model?
ˆ What level of certainty can be implied for model predictions?
ˆ What is the model’s sensitivity to small changes in parameter values?
For our model to be more realistic and best predict how the two species would best interact and
how their populations would vary over time, factors such as rainfall, temperature and humidity,
diseases and other factors must be considered.

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K.B Raborife (16528301) HRAPM82 Assignment 10

References
[1] Mira-Cristiana Anisiu. Lotka, volterra and their model. Didáctica mathematica, 32:9–17,
2014.
[2] Carly Dodd. Habitat and ecological niche. https://www.worldatlas.com/articles/
habitat-and-niche.html, 2021. Accessed January 10, 2023.
[3] Daniel M Dubois. A model of patchiness for prey—predator plankton populations. Eco-
logical Modelling, 1(1):67–80, 1975.
[4] Inasse El Arabi, Anas Chafi, and S Kammouri Alami. Numerical simulation of the sir
and lotka-volterra models used in biology. In 2019 International Conference on Intelligent
Systems and Advanced Computing Sciences (ISACS), pages 1–4. IEEE, 2019.
[5] Temple H Fay and Johanna C Greeff. Lyons and wildebeest: A predator-prey model.
Mathematics and Computer Education, 33:106–119, 1999.
[6] Alfred J Lotka. Analytical note on certain rhythmic relations in organic systems. Proceed-
ings of the National Academy of Sciences, 6(7):410–415, 1920.
[7] Sarah P Otto and Troy Day. A biologist’s guide to mathematical modeling in ecology
and evolution. In A Biologist’s Guide to Mathematical Modeling in Ecology and Evolution.
Princeton University Press, 2011.
[8] Anthony M Starfield and Andrew L Bleloch. Building models for conservation and wildlife
management. Macmillan, 1986.
[9] N.F.J van Rensburg. Differential equations and dynamical systems, 2021.
[10] Vito Volterra. The general equations of biological strife in the case of historical actions.
Proceedings of the Edinburgh Mathematical Society, 6(1):4–10, 1939.

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Index
Asymptotically stable point, 10
Autonomous systems, 8

Cropping, 17, 18

Differential equation, 8

Ecological niche, 7
Ecology, 7
Eigenvalues, 11
Equilibrium points, 10
Extinction, 10

Globally asymptotically stable point, 10

Habitation, 7

Initial value problem, 8

Integration of differential equations, 13

Jacobian, 11, 15

Linear system, 11
Lipschitz function, 8
Locally lipschitz, 9
locally lipschitz, 8
Lotka-Volterra model, 7, 12

Managing population, 17

Partial derivatives, 8
Phase potrait, 14
Predator population in terms of differential equa-
tions, 13
Prey population in terms of differential equa-
tions, 13

Separation of variables, 13
Solution of a differential equation, 8
Species competition, 7
Stability of a point, 10
Stable point, 10

Unstable point, 10

22
K.B Raborife (16528301) HRAPM82 Assignment 10

A Appendix

Table 1: Wildebeest, zebra and lion population densities since 1972

.
Year Wildebeest Zebra
1972 10600 10500
1974 7931 7523
1975 6745 7850
1976 5783 7616
1977 5066 7649
1978 5141 8316
1979 5502 8511
1980 5816 8877
1981 6512 10834
1982 8127 11603
1983 7584 9807
1984 8062 12830
1985 8634 11822
1986 9406 12520
1987 9915 13097
1988 9650 12431
1989 9547 113008
1990 9807 12176
1991 9788 13240
1992 10574 12060
1993 10658 13577
1994 11243 13004

Table 1 shows population census numbers for wildebeest and zebra from 1972 to 1994, whereas
the lion population has been maintained at around 300-500 by migrating nomads [?].

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K.B Raborife (16528301) HRAPM82 Assignment 10

Table 2: Number of lions to be cropped to maintain a stable population using strategy (12)
over a year
.
Month Population of Lions Number of lions to be cropped
0 500 -
1 476 24
2 454 22
3 434 20
4 415 19
5 398 17
6 383 15
7 368 15
8 355 13
9 342 13
10 331 11
11 321 10
12 311 10

Table 2 above gives us an indication as to how many lions should be culled so that the population
of wildebeest would recover [5].

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K.B Raborife (16528301) HRAPM82 Assignment 10

B Numerical solution code

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