Two-prey one predator model

Submitted by
Ausmita Barman

Supervisor: Dr. Behzad Djafari-Rouhani

University of Texas at El Paso

Monday 29th June, 2015

Abstract

In this paper we propose a new multi-team preypredator model, in which the prey teams help each other.
We study its local stability. In the absence of predator, there is no help between the prey teams. So, we
study the global stability and persistence of the model without help.

Introduction
The growth and decline of populations in nature and the interaction between species have been a subject
of interest for many years. Among the founders of mathematical population models were Malthus [1],
who proposed a model of population dynamics for single species. The solution of this problem is an
exponential function in the form:
N (t) = N0 ekt

(1)

where N(t) is the size (number) of the population at time t, N0 is the initial population at time t = 0 and k
is the rate of reproduction per unit time. This model was applied to human populations, bacterial growth
(k > 0) and radioactive decay (k < 0).
Realistically, this model lose its accuracy with the increasing of time. Since, the growth of biological populations in the presence of a limited nutrient supply always decelerates and eventually stops. This
would tend to imply that k is not a constant but changes with time.
Later on 1838 Verhulst [2] proposed a more realistic model for the study of population dynamics,
this model is known as the logistic model, which takes the following non-dimensional form:
dN (t)
= N (t)F (N (t)) = rN (t)(1 N (t)
dt

(2)

where F(N(t)) is the rate of reproduction. The solution of equation (2) is

N (t) =

N0
N0 + (1 N0 )ert

(3)

where N0 = N(0) is the initial population and r is the intrinsic growth parameter. So, the Verhulst model
improves the Malthus model by including competition for resources.
When species interact, the population dynamics of each species is affected. There are three main
types of interaction [3]:
1. If the growth rate of one population is decreased while the growth rate of the other population
increased, then the populations are in a predatorprey situation.

2. If the growth rate of each population is decreased, then the two populations are in a competition
case.
3. If the growth rate of each population is enhanced, then it is a mutualism situation.
A system of two equations was proposed independently by Lotka [4] and Volterra [5] for studying
the preypredator interaction. LotkaVolterra model is the simplest model of predatorprey interactions.
They are among the first to study this phenomena by making a number of simplifying assumptions that
led to nontrivial but tractable mathematical problem. The model proposed to explain the oscillatory levels
of certain fish catches in the Adriatic sea during the first world war. Suppose that N(t) is the size of prey
population and P(t) be the size of the predator population at time t, then the LotkaVolterra model is given
by the following equations:

dN (t)
dt
dP (t)
dt

= N (t)F (N (t), P (t)) = N (t)(a bP (t))

= N (t)G(N (t), P (t)) = P (t)(cN (t) d)

(4)

where F(N,P) and G(N,P) are the reproduction rates of prey and predator, respectively. Also, the parameters a, b, c and d are positive constants.
The assumptions of this model are as follows:
(i) In the absence of any predation, the prey grows unboundedly in a Malthusian way; this is the (aN) term
in the first equation.
(ii) The effect of the predation is to reduce the prey growth rate by a term proportional to the prey
and predator populations;this is the (-bNP) term.
(iii) In the absence of any prey for sustenance the predators death rate results in exponential decay,
that is, (-dP) term in the second equation.
(vi) The preys contribution to the predator growth rate is (cNP) term; that is proportional to the available
prey as well as the size of the predator population.
Since their pioneering work, many other notable contributions were made [6,7]. One of the unrealistic assumptions in the LotkaVolterra model is that the prey growth is unbounded in the absence of the
predator. So, the prey growth have to changed to satisfy the logistic growth in the absence of any predator.
Then, the LotkaVolterra model can be written in the form:
dN (t)
N (t)
dt = rN (t)(1 K ) N (t)P (t)R(N (t)),
dP (t)
dt = P (t)G(N (t), P (t)) = P (t)(cN (t) d)

(5)

where R(N(t)) is the predation term. This term my be constant or variable (like Holling types) [812].
Also, there are a lot of studied on the delay predatorprey systems [1315].

Many creatures form teams. This has at least two main advantages, the first is the improvement in
foraging since looking for food in a team is more efficient than doing it alone. The second is that living in
a team reduces predation risk due to early spotting of predators and that existing in a team gives a higher
probability that the predator will attack another member of the team.
Here a model is given where one predator interacts with two teams of preys. The teams of preys
group help each other. We will study the equilibrium solutions, their local stability. Also, global stability
of the interior solutions and persistence of the system without help are studied.
A biological realization of our study is the herds of zebras and gazelles living side by side and attacked by one type of predator. The problem of multi-team game is very recent. It has been studied in
[16,17].

The model
In this section, we propose a system consists of two teams of preys with densities x(t), y(t), respectively,
interacting with one team of predator with densities z(t). The assumptions of this model are as follows:
In the absence of any predation, each team of preys grows logistically; this is ax(t)(1-x(t)) and
ay(t)(1-y(t)).
The effect of the predation is to reduce the prey growth rate by a term proportional to the prey and
predator populations; this is the -x(t)z(t) and -y(t)z(t) terms.
The teams of preys help each other against the predator, that is a x(t)y(t)z(t) term exist.
In the absence of any prey for sustenance, the predators death rate results in inverse decay, that is
the term cz 2 (t).
The preys contribution to the predator growth rate are dx(t)z(t), ey(t)z(t); that is proportional to the
available prey as well as the size of the predator population.
Using the above assumptions, the following model is proposed:
dx(t)
dt = ax(t)(1 x(t)) x(t)z(t) + x(t)y(t)z(t)
dy(t)
dt = by(t)(1 y(t)) y(t)z(t) + x(t)y(t)z(t)
dz(t)
2
dt = cz (t) + dx(t)z(t) + ey(t)z(t)

(6)

where the coefficients a, b, c, d and e are positive constants and x(0),y(0), z(0) > 0. It is clear that the two
teams of preys help each other e.g. in foraging and in early warning against predation. Note that this help
occurs only in the presence of predator. This is presented by the term x(t)y(t)z(t) in the preys equations.

The analysis of the model

By Solving the equations,
dx(t)
= ax(t)(1 x(t)) x(t)z(t) + x(t)y(t)z(t) = 0
dt

(7)

dy(t)
= by(t)(1 y(t)) y(t)z(t) + x(t)y(t)z(t) = 0
dt

(8)

dz(t)
= cz 2 (t) + dx(t)z(t) + ey(t)z(t) = 0
dt

(9)

From (7) we get,

x = 0 or a(1 x) z + yz = 0
For x = 0 (8) becomes,
y = 0 or b(1) z = 0
For x = 0, y = 0 (9) becomes,
z=0
E0 (0, 0, 0)
For x = 0 (8) becomes,
b(1 y) z = 0 y =
Now (9) becomes,
z = 0 or cz +

bz
b

eb ez
=0
b

E1 (0, 1, 0)
For x = 0, y =

bz
b

(9) becomes,
cz +

Now y =

eb ez
eb
=0z=
b
e + bc

cb
e+bc

E2 (0,

cb
eb
,
)
e + bc e + bc

From (7) we get,

a(1 x) z + yz = 0 x =
Now (8)becomes,
y = 0 or (1 y) z +

a z + yz
a

a z + yz
z=0
a
9

From (9) we get,

z = 0 or cz +

da dz
=0
a

E3 (1, 0, 0)
az
a ,y

For x =

= 0 Now (9) becomes,

cz +

da
da dz
=0z=
a
ca + d

E4 (

ca
da
, 0,
)
ca + d
ca + d

Now (8)becomes,
b(1 y) z +

a z + yz
z2
z2
z = 0 y( b) = ( b)
a
a
a

If ( za b) 6= 0 then y = 1
q
z2
If ( a b) = 0 then z = ab
For y = 1, x = 1 From (9) we get,
z = 0 or cz + d + e = 0
E5 (1, 1, 0)
For x = 1, y = 1 (9) becomes,
z=

d+e
c

E6 (1, 1,
For z =

b
a

d+e
)
c

we get infinitely many values of y.

Let y = where is a positive parameter.

From (7)

x=1+

E7 (1 +

If we assume,y = =
under the conditions

c abd(1 ab )
q
e+d ab

ab( 1)
a

ab( 1)
, , ab)
a

then x =

q
cb+e(1 ab )
q
e+d ab

r
b
b
cb + e(1
) > 0 cb + e > e
a
a
r
r

b
b
c ab d(1
) > 0 d c ab < d
a
a
Definitions: First, we define some types of stability for solutions of the equation:
dx(t)
= f (x, t)...................()
dt

Definition 1: A solution x
(t) of (*) is (Lyapunov) stable if for each  > 0 and t0 R there exists
= (, t0 ) > 0 such that if x(t) is a solution of (*) and |x(t0 ) x
(t0 )| <  then |x(t) x
(t)| < for
all t t0 .
Definition 2: A solution x
(t) of (*) is asymptotically stable if it is (Lyapunov) and if for every t0 R there
exists = (t0 ) > 0 such that if x(t) is a solution of (*) and |x(t0 ) x
(t0 )| < then |x(t) x
(t)| 0
for all t .
Definition 3: A solution x
(t) of (*) is uniformly (Lyapunov) stable if for each  > 0 and t0 R there
exists = () > 0 such that if x(t) is a solution of (*) and |x(t0 ) x
(t0 )| <  then |x(t) x
(t)| <
for all t t0 .
Proposition 1: Local stability analysis [18] shows that the system (6) has four unstable equilibrium
solutions E0 , E1 , E2 and E3 .
Proof: The Jaconian matrix of the above system (6) is given by:

a(1 2x) z(1 y)

xz
x(1 y)

J =
yz
b(1 2y) z(1 x)
y(1 x)

dz
ez
2cz + dx + ey
Substituting by the point E0 (0, 0, 0) in the above matrix,

a 0 0

JE0 = 0 b 0
0 0 0
det(JE0 I) = 0 = 0, b, a
which has two positive eigenvalues. So, it is unstable equilibrium point.
Similarly,

a 0 1

JE1 = 0 b 0
0
0
d
det(JE1 I) = 0 = d, b, a
which has two positive eigenvalues. So, it is unstable equilibrium point.
Similarly,

a 0
0

JE2 = 0 b 1
det(JE2

0 0
e
I) = 0 = a, b, e

E2 (0, 1, 0) has two positive eigenvalues. So, it is unstable equilibrium point.

By the same way,

a 0

JE3 = 0 b
det(JE3

0
0
0 d+e
I) = 0 = a, b, d + e

E3 (0, 1, 0) has two positive eigenvalues. So, it is unstable equilibrium point.

So, all of them are unstable equilibrium points.
Now,
a

JE4 =

cb
eb
bc+e (1 cb+e
2
ceb
(cb+e)2
ebd
cb+e

0
b(1

2cb
cb+e

be2
cb+e

cb
cb+e

2ceb
cb+e
+

ceb
cb+e

det(JE4 I) = 0
=a

be2
(cb+e)2

or 2 + ( bce+cb
bc+e ) +

ceb2
(bc+e)

=0

The equilibrium solution E4 is locally asymptotically stable under the condition,

a<

be2
(cb + e)2

Numerical simulations agreed with these results. Let a = 1.0, b = 2.0, c = 0.1, d = 1.0, e = 1.4, we get the
stable solution (0, 0.125, 1.75) as shown in Fig. 1.

Fig. 1. In this figure we used the values a = 1.0, b = 2.0, c = 0.1, d = 1.0, and e = 1.4. So, we get the stable
solution (x = 0, y = 0.125, z = 1.75).
Similarly

a(1

JE5 =

2ca
ca+d

ad
ca+d

0
ad2
ca+d

cda2
(ca+d)2
ad
ca
ca+d (1 ca+d
ead
ca+d

ca
ca+d

2cad
ca+d
+

cad
ca+d

det(JE5 I) = 0
=b

ad2
(ca+d)2

or 2 + ( cad+ca
ca+d ) +

dca2
(ca+d)

=0

The equilibrium solution E5 is locally asymptotically stable under the condition,

b<

ad2
(ca + d)2

Numerical simulations agreed with these results. Let a = 1.0, b = 0.4, c = 0.3, d = 1.0, e = 1.4, we get the
stable solution (0.231, 0, 0.769) as shown in Fig. 2.

Fig. 2. In this figure we used the values a = 1.0, b = 0.4, c = 0.3, d = 1.0, and e = 1.4. So, we get the stable
solution (x = 0.231, y = 0, z = 0.769).

d+e
0
a
c

Now JE6 = d+e

b
0
c
d2 +ed
c

ed+e2
c

d e

det(JE6 I) = 0
2
The first internal equilibrium solution E6 is locally asymptotically stable if ab > ( d+e
c ) . Numerical

simulations agreed with these results. Let a = 1.0, b = 2.0, c = 2.0, d = 1.0, e = 1.6, we get the stable
solution (1, 1, 1.3) as shown in Fig. 3.

Fig. 3. In this figure we used the values a = 1.0, b = 2.0, c = 2.0, d = 1.0, and e = 1.6. So, we get the stable

solution (x = 1, y = 1, z = 1.3).
The second internal equilibrium solution

ab( 1)
, , ab)
a

a + 3 ab( 1)
ab + b b
a + ab + ab2 2 ab

ab ab2
JE7 =

ab
b
a

d ab
e ab
2ac ab + ad + d ab d ab
det(JE7 I) = 0
E7 (1 +

We get the characteristic equation,

3 + a1 2 + a2 + a3 = 0

where

a1 = a 3 ab + 3 ab + b + 2ac ab ad b ab d ab

a2 = ab 3 abb2 + 3b ab + 2ca2 ab a2 d ab ab abad 6a2 cb 3ab ab + 32 b2 a +

3dab + 6a2 cb 3ad ab 3b2 a 3abd ab 2 b ab + b ab + ad ab dab 2 abd +

2abd + 2abc ab abd bd ab bd ab 2b + 2b + 2b2

a3 = b(a + 3 ab 3 ab)(2ac ab ad d ab d ab) + 2b a + 3 ab( 1) ( ab +

b b)db(b b2 ) (a + ab + ab2 2 ab)eab (a + ab + ab2 2 ab)bd ab +

( ab + b b)(2ac ab + ad + d ab d ab) ab
E7 is locally asymptotically stable under the Routh-Hurwitz conditions: a1 > 0, a3 > 0anda1 a2 >
a3 .Numerical simulations agreed with these results. Let a = 1.2, b = 1.4, c = 1.0, d = 1.0, e = 2.0, we
get the stable solution (0.40, 0.45, 1.29). Also, let a = 1.0, b = 1.44, c = 1.0, d = 1.0, e = 1.2, we get the
internal solution (0.51, 0.59, 1.23) as shown in Figs. 4 and 5.

Fig. 4. In this figure we used the values a = 1.2, b = 1.4, c = 1.0, d = 1.0, and e = 2.0. So, we get the

internal stable solution (x = 0.40, y = 0.45, z = 1.29).

Fig. 5. In this figure we used the values a = 1.0, b = 1.44, c = 1.0, d = 1.0, and e = 1.2. So, we get the
internal stable solution (x = 0.51, y = 0.59, z = 1.23).
Now, we consider the above system but without team interaction help term:
dx(t)
dt = ax(t)(1 x(t)) x(t)z(t)
dy(t)
dt = by(t)(1 y(t)) y(t)z(t)
dz(t)
dt

(10)

2
= cz (t) + dx(t)z(t) + ey(t)z(t)

By Solving the equations,

dx(t)
= ax(t)(1 x(t)) x(t)z(t) = 0
dt

(11)

dy(t)
= by(t)(1 y(t)) y(t)z(t) = 0
dt

(12)

dz(t)
= cz 2 (t) + dx(t)z(t) + ey(t)z(t) = 0
dt

(13)

bc
eb
We get these equilibrium points E10 (0, 0, 0), E20 (0, 1, 0), E30 (0, bc+e
, bc+e
), E40 (1, 0, 0), E50 (1, 1, 0),
ad(d+e)
abcbe+ae abc+bdad
E60 ( abc+bd+ae
, abc+bd+ae , abc+bd+ae
) To prove the global stability of the internal solution of the system

(10), we use the following propositions.

Proposition 2: The system
du(t)
= ru(t)(1 u(t))
dt

has two equilibrium solutions, u(t) = 0 and u(t) = 1, where the constant r > 0. The non-zero equilibrium
solution u(t) = 1 is globally stable for all solutions u(t) with the initial condition u(0) > 0.
Proof: This equation is a Bernoulli differential equation whose solution is
Let,
u = v 1

du
= ru ru2
dt
Now,
du
dv
= v 2
dt
dt
Then,
v 2

dv
= rv 1 rv 2
dt
dv
= rv + r
dt

Let,
(t) = e

rdt

= ert

Now,
ert

dv
+ rvert = rert
dt

dv rt
[ve ] = rert
dt

vert = rert + C

v = 1 + Cert

So
u=

1
1 + Cert

Then
u0 =

1
1 u0
C=
1+C
u0

u=

u0
u0 + (1 u0 )ert

which is non-zero, non-oscillatory solution that tends to 1 when t tends to .

Now consider two cases,
Case 1: Let
u0 < 1
u(t) = increasing < 1

If for some t0 , (1 u(t0 )) < ,then t > t0 , u(t) > u(t0 ).So 0 < (1 u(t)) < (1 u(t0 )) <
Case 2: Let
u0 > 1
u(t) = decreasing > 1

If for some t0 , (u(t0 ) 1) < ,then t > t0 , u(t0 ) > u(t).So 0 < (u(t) 1) < (u(t0 ) 1) < 
So, u(t) is uniformly stable.
Proposition 3: Consider the system (without help)
dx(t)
dt = ax(t)(1 x(t)) x(t)z(t)
dy(t)
dt = by(t)(1 y(t)) y(t)z(t)
dz(t)
dt

(14)

= cz 2 (t) + dx(t)z(t) + ey(t)z(t)

where a, b, c, d and e are positive constants. Then, the internal solutions of (14) are globally asymptotically
stable for all solutions with the initial conditions x(0) > 0, y(0) > 0 and z(0) > 0.
Proof: We use a method similar to the one used in [19]. The internal solution of the system (14)
is,x =

abcbe+ae
abc+bd+ae , y

abc+bdad
abc+bd+ae , z

ad(d+e)
abc+bd+ae

The first equation of the system (14) implies

dx(t)
ax(t)(1 x(t))
dt
using Proposition 2 then
x(t) U1x , U1x = 1

The second equation of the system (14) implies

dy(t)
by(t)(1 y(t))
dt
using Proposition 2 then
y(t) U1y , U1y = 1

Now, from the third equation of the system (14), we have

z(t)
z(t)
dU x + eU1y
cz 2 (t) + (dU1x + eU1y )z(t)
cz(t)( 1
z(t))
dt
dt
c
using Proposition 2 then
z(t)

U1z , U1z

dU1x + eU1y
=
c

On the other hand

dx(t)
Uz
dx(t)
ax(t)(1 x(t)) x(t))U1z
ax(t)((1 1 ) x(t))
t
t
a
using Proposition 2 then
x(t) Lx1 , Lx1 = 1

U1z
a

y(t) Ly1 , Ly1 = 1

U1z
b

By the same way,

Similarly
z(t)
z(t)
dLx + eLy1
cz 2 (t) + (dLx1 + eLy1 )z(t)
cz(t)( 1
z(t))
dt
dt
c
using Proposition 2 then
z(t) Lz1 , Lz1 =

dLx1 + eLy1
c

Since, U1z = supz(t), then z(t) Lz1

The first equation of the system (14) implies
dx(t)
dx(t)
Lz
ax(t)(1 x(t)) Lz1 z(t)
ax(t)((1 1 ) x(t))
dt
t
a

using Proposition 2 then

x(t) U1x , U1x = 1

Lz1
a

y(t) U1y , U1y = 1

Lz1
b

Similarly

Continuing the above process one gets a sequence of upper limits Uix , Uiy , Uiz and lower limits Lxi , Lyi , Lzi , i =
2, 3..... related by the relations:
Uix = 1

dUix + eUiy
Lzi
Lz
, Uiy = 1 i , Uiz =
a
b
c

Lxi = 1

dLxi + eLyi
Uiz y
Uz
, Li = 1 i , Lzi =
a
b
c

Now solving these equations,

Lzi

d(1
dLxi + eLyi
Lzi =
=
c

Uiz

Uiz
a )

+ e(1
c

d(1
dUix + eUiy
=
Lzi =
c

Lzi
a )

Uiz
b )

Lzi =

+ e(1
c

Lzi
b )

abd dbUiz + aeb aeUiz

abc

Uiz =

ab(d + e)
abc + bd + ae

Lxi = 1

ab(d + e)
abc be + ae
Lxi =
a(abc + bd + ae)
abc + bd + ae

Lyi = 1

ab(d + e)
abc + bd ad
Lyi =
b(abc + bd + ae)
abc + bd + ae

Lzi =

ab(d + e)
abc + bd + ae

Uix = 1

ab(d + e)
abc be + ae
Uix =
a(abc + bd + ae)
abc + bd + ae

Uiy = 1

ab(d + e)
abc + bd ad
Uiy =
b(abc + bd + ae)
abc + bd + ae

Definition 4: A system is persistent if there exist a compact region V subset of the interior of the state
space such that all solutions with positive initial conditions are attracted to V [19].
Propositiion 4: The system (14) is persistent.
Proof: From Proposition 3, we have proved that for all t > 0, all solutions with positive initial conditions
satisfy:
0 < Lxi x(t) Uix

0 < Lyi y(t) Uiy

0 < Lzi z(t) Uiz

Thus the system (14) is persistent.

Conclusion
Many animals live in groups. Different groups share one habitat hence these groups may cooperate,
compete with each other or form predatorprey system.
In this work we present a new model for predatorprey teams. Equilibrium solutions are derived, their
local stability, persistence and global stability are studied. A biological realization of our model is two
cooperating teams of gazelles and zebras attacked by one predator. We get a global stable internal solution
for the proposed system without preys help.

21

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