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A MODIFIED LESLIE-GOWER PREDATOR-PREY MODEL

WITH HYPERBOLIC FUNCTIONAL RESPONSE AND
ALLEE EFFECT ON PREY
CLAUDIO ARANCIBIA-IBARRA AND EDUARDO GONZLEZ-OLIVARES
Grupo de Ecologa Matemtica, Instituto de Matemticas,
Ponticia Universidad Catlica de Valparaso, Chile
E-mail: clanarib@gmail.com, ejgonzal@ucv.cl
This work deals with a modied Leslie-Gower type predator-prey model consider-
ing two important aspects for describe the interaction: the functional response is
Hollling type II and the Allee eect acting in the prey growth function.
With both assumptions, we have a modication of the known May-Holling-Tanner
model, and the model obtained has a signicative dierence with those model due
the existence of an equilibrium point over y-axis, which is an attractor for all pa-
rameter values.
We prove the existence of separatrix curves on the phase plane dividing the behav-
ior of the trajectories, which have dierent ! l{mit. System has solutions highly
sensitives to initial conditions To simplify the calculus we consider a topologically
equivalent system with a minor quantity of parameters. For this new model, we
prove that for certain subset of parameters, the model exhibits biestability phe-
nomenon, since there exists an stable limit cycle surrounding a singularities of
vector eld or an stable positive equilibrium point.
.
1. INTRODUCTION
This work deals with a continuous-time predator-prey considering that:
i) The functional response is the hyperbolic type
23
,
ii) the growth of the predator population is of logistic form
18,23
, and
iii) the growth of the prey population is aected by the Allee eect
7,22
.
This last assumption characterize the Leslie-Gower type models
14,19
in
which the environmental carrying capacity 1

is a function of prey size

r
18,23
, that is, it depends on the available resources. Usually it is assumed
that 1

= 1(r) = :r, i. e., the carrying capacity is proportional to

the prey abundance, just as it is considered in the May-Holling-Tanner
model
1,11,21
.
We will assume that the predators have an alternative food when the
1
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2
quantity of prey diminish, this is, 1(r) = :r + c, obtaining a modied
Leslie-Gower model
2,15
, being c the measure of their alternative food. For
r = 0, then 1(r) = c, concluding that the predator is generalist since it
has an alternative food
2
.
The predator functional response or consumption function refers to the
change in attacked prey density per unit of time per predator when the prey
density changes. In many predator-prey models considered in books, it is
assumed that the functional response grows monotonic, being the inherent
assumption the more prey in the environment, the better for the predator
23
.
In this work, the predator functional response is expressed by the hyper-
bolic function ,(r) =
j r
r + o
, corresponding to the Holling type II
23
. Here,
the parameter a is a measure of abruptness for the functional response
12
.
If a 0, the curve grows quickly, while if a 1, the curve grows slowly,
i. e., a bigger amount of prey is needed to obtain
j
2
.
On the other hand, any ecological mechanism that may lead to a positive
ratio between measurable component of individual tness and the popula-
tion number or density may be called a mechanism of Allee eect
6,22
.
This phenomenon has been also called depensation in Fisheries
Sciences
4,17
, or negative competition eect, inverse density dependence or
positive density dependence in Population Dynamics
17
.
Populations can exhibit Allee eect dynamics due to a wide range of
biological phenomena, such as reduced antipredator vigilance, social ther-
moregulation, genetic drift, mating diculty, reduced antipredator defense,
and decient feeding at low densities; however, several other causes may
lead to this phenomenon (see Table 2.1 in
7
).
Many algebraic forms have been used for describing the Allee eect
13,24
,
but we use the most common to describe this phenomenon. However it is
possible to prove the topological equivalence between them
13
.
In turn, the existence and uniqueness of limit cycles are interesting
subjects in predator-prey models, but the determination of the amount of
limit cycles that may arise by the bifurcation of a center focus singularity
5
is not a easy task; this matter is still without answer for the predator-prey
models
16
just as it happens with the Hilbert 16th Problem for polynomial
dierential equations systems
10,16
.
The main goal of this work is to describe the behaviour of model es-
tablishing its bifurcation diagram
3
based on the parameter values and to
study the number of limit cycles.
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3
2. THE MODEL
We analyze the modied Leslie-Gower type predator-prey model, described
by the autonomous bidimensional dierential equations system
A

:
_
_
_
Jr
J|
=
_
r
_
1
r
1
_
(r :)
j
r+o
j
_
r
J
J|
= :
_
1

nr+c
_
j
(1)
which is of Kolmogorov type system
9
, where r = r(t) and j = j(t) indicate
the prey and predator population sizes, respectively (measure in number
of individuals, density or biomass); j = (r, 1, , :, a, :, :, c) R
8
+
and
parameters have the following meanings:
r is the intrinsic prey growth rate,
1 is the environmental carrying capacity,
is the consuming maximum rate per capita of the predators ,
a is the amount of prey to reach half of ,
: is intrinsic predator growth rate,
: is the food quality and it indicates how the predators turn eaten prey
into new predator births, and
c is the amount of alternative food available for the predators.
The parameter : satises that 1 _ : << 1. When : 0, the
population growth rate decreases if the population size is below the level :
and the population goes to extinction, i.e., : the minimum of viable prey
population or extinction threshold. In this case, the prey population is
aected by a strong Allee eect
24
. If : _ 0, it is said that the population
is aected by a weak Allee eect
7
. In sheries the same phenomena are
called critical and pure depensation, respectively
4,17
.
The parameter c 0 indicates that the predator is generalist and that
if it does not exist available prey it has a source of alternative food.
As system (1) is of Kolmogorov type, the coordinates axis are invariable
sets of the model and it is dened in the rst quadrant:
= (r, j) R
2
, r _ 0, j _ 0 = R
+
0
R
+
0
The equilibrium points of the system (1) or singularities of vector
eld A

are Q
0
= (0, 0), Q
n
= (:, 0), Q
1
= (1, 0), Q
c
= (0, c)
and Q
t
= (r
t
, j
t
), the positive equilibrium points (in the interior of the
rst quadrant) satisfying the equations of the isoclines j = :r + c and
j =
:
jr
(1
r
1
) (r :) (r +a).
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To simplify the calculus we follow the methodology used in
?,19,21
, doing
the change of variable and the time rescaling, given by the function , :

R R; so that
,(n, , t) =
_
1n, 1:,
_
n +
o
1
_ _
n +
c
1n
_
r1
t
_
= (r, j, t)
and det 1,(n, , t) =
1n(u +
a
K
)(u+
c
Kn
)
:
0; hence, , is a
dieomorphism
3
and the vector eld A

, in the new coordinates, is topo-

logically equivalent to the vector eld 1
i
= , A

, having the form

1
i
= 1(n, )
J
Ju
+ Q(n, )
J
Ju
; the associated dierential equation system
is given by the polynomial system of fth degree:
1
i
:
_
Ju
Jr
(n +C) ( (1 n) (n ') (n + ) Q) n
Ju
Jr
o (n +) ( n +C )
(2)
with i = (, ', o, C, Q) =]0, 1[]0, 1[R
3
+
, where =
o
1
< 1, o =
s
:1
, ' =
n
1
< 1, Q =
nj
:1
, C =
c
1n
. System (2) is dened in

= (n, ) R
2
, n _ 0, _ 0.
The equilibrium points of system (2) or singularities of vector eld 1
i
are: 1
0
= (0, 0), 1
1
= (', 0), 1
1
= (1, 0), 1
c
= (0, C) and the points lie
in the intersection of the isoclines curves =
1
Q
(1 n) (n ') (n + )
and = n +C, obtaining:
n
3
(' + 1 ) n
2
((' + 1) Q') n + (' +CQ) = 0 (c)
Applying the Descartes signs rule, equation (c) can have two positives
roots, one of multiplicity two, or none, for any sign of (' + 1) Q',
since the factors ' + (1 ) 0 ( < 1) and ' +CQ 0. Replacing
n in (c) we have:
n
3
(' + 1 ) n
2
+ ((' + 1) Q') n + (' +CQ) = 0,
which has a unique change of sign; then, it exists a unique negative real
root that we will denote it by H.
Due to the dicult to determine the exact solutions of equation (c),
we divide it by n + H. Denoting by
1
= 1 + H + ' and
0
=
Q+'H (H +' + 1) '+1, the following equation is obtained: