A DISCRETE-TIME HOST-PARASITOID MODEL

SOPHIA R.-J. JANG AND JUI-LING YU

We study a discrete-time host-parasitoid model proposed by May et al. In this model,

the parasitoid attacks the host first then followed by density dependence, where density
dependence depends only on those host populations that escaped from being parasitized.
Asymptotic dynamics of the resulting system are derived. There exist thresholds for which
both populations can coexist indefinitely.
Copyright 2006 S. R.-J. Jang and J.-L. Yu. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
1. Introduction
It is well known that the sequence of density dependence and parasitism in the host life
cycle can have a significant eect on the population dynamics of the host-parasitoid interaction. Consequently, the eect can have important implications for biological control. In
[10], May et al. proposed and numerically simulated three host-parasitoid models based
on the timing of parasitism and density dependence. In this work, we will study a model
proposed by May et al. [10] in which parasitism occurs first then followed by density dependence. However, density dependence only depends on the remaining host population
that escaped being parasitized.
2. The model
Let Nt be the host population at time t. The parasitoid population at time t is denoted
by Pt . An individual parasitoid must find a host to deposit its eggs so that the parasitoid
can reproduce. It is assumed that parasitism occurs first then followed by density dependence. Let be the average number of osprings that a parasitized host can reproduce
for a parasitoid individual. It is assumed that the number of encounters between host
and parasitoid populations at any time t 0 follows that of simple mass action, bNt Pt ,
where the searching eciency b is a constant. We assume that the number of encounters
is distributed randomly with a Poisson distribution. Consequently, the probability that
an individual host will escape from being parasitized when the parasitoid population is
Hindawi Publishing Corporation
Proceedings of the Conference on Dierential & Dierence Equations and Applications, pp. 451455

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A discrete-time host-parasitoid model

of size P is ebp . For simplicity, the host population in the absence of the parasitoid is
modeled by a simple Beverton-Holt equation N/(1 + kN), where parameters and k are
positive. Since density dependence occurs after parasitism, the interaction between the
host and the parasitoid is governed by the following system of dierence equations:
Nt+1 =

Nt
ebPt ,
1 + kNt ebPt


(2.1)

Pt+1 = Nt 1 ebPt ,

(2.2)

N0 ,P0 0.
Steady state E0 = (0,0) always exists. The Jacobian matrix can be given by


J=

J11

1e

J12


bP

bNebP

(2.3)

where
ebP
J11 = 
2 ,
1 + kNebP
bNebP
J12 = 
2 .

(2.4)

1 + kNebP

Note that


J(0,0) =

0 0

(2.5)

Thus it can be easily seen that E0 is the only steady state of system (2.1) if < 1 and it is
globally asymptotically stable. Indeed,
Nt+1 =

Nt
Nt
ebPt =

bP
t
1 + kNt e
kNt + ebPt

Nt

< Nt ,
1 + kNt

(2.6)

for t 0 implies limt Nt = 0 as < 1. As a result, we can show that limt Pt = 0 and
hence E0 = (0,0) is globally asymptotically stable.
Suppose now > 1. Then (2.1) has another boundary steady state E1 = (( 1)/k,0) =

(N,0)
and the Jacobian matrix of the system associated with E1 is

=
J(N,0)

J12 E1

1.
b
k

(2.7)

S. R.-J. Jang and J.-L. Yu 453

Thus E1 is locally asymptotically stable if b( 1)/k = bN < 1. We show that (2.1) has
no interior steady state if bN < 1. Notice that the P-component of an interior steady
state (N ,P ) must satisfy
= ebP + kh(P),

(2.8)

where h(P) = P/(1 ebP ) for P > 0. Since limP0+ h(P) = 1/b, h (P) > 0 for P > 0 and
limt h() = , we see that (2.8) has a positive solution P if and only if
b + k
< i bN > 1.
b

(2.9)

In this case P > 0 is unique and there is a unique interior steady state E1 = (N ,P ) if
bN > 1. We conclude that if > 1 and bN < 1, then E1 is locally asymptotically stable and there is no interior steady state. We show that solutions of (2.1) with N0 > 0 all
converge to E1 .
To this end,
Nt+1 =

Nt
Nt

,
ebPt + kNt 1 + kNt

(2.10)

for t 0 implies limsupt Nt ( 1)/k by a simple comparison argument. Then for

any  > 0 there exists t0 > 0 such that Nt < ( 1)/k +  for t t0 . Since bN < 1, we
choose  > 0 such that
b(N + ) < 1.

(2.11)

But then


Pt+1 = Nt 1 ebPt < (N + ) 1 ebPt b(N + )Pt ,

(2.12)

for t t0 implies limt Pt = 0. Consequently, we can prove that liminf t Nt (

1)/k if N0 > 0. Therefore, limt Nt = N and E1 is globally asymptotically stable.
Suppose now > 1 and bN > 1. Notice E0 and E1 are unstable and (2.1) has a unique
interior steady state. We prove that the system is uniformly persistent by using a result of
Hofbaur and So [6]. Clearly, system (2.1) has a global attractor X. Let Y = {(N,P) R2+ :
N = 0 or P = 0}, that is, Y is the union of nonnegative coordinate axes, and let M be the
maximal invariant set in Y . Then M = {E0 ,E1 }, where {E0 } and {E1 } are isolated in X.
We claim that the stable set W + (E0 ) = {(N,P) R2+ : Nt 0,Pt 0 as t } lies in Y .
For suppose there exists a solution (Nt ,Pt ) of (2.1) with N0 > 0, P0 > 0 such that
limt (Nt ,Pt ) = E0 , then since > 1, we can choose  > 0 such that eb > 0. For this
 > 0 there exists t1 > 0 such that Pt <  for t t1 , and consequently
Nt+1 =

Nt
Nt
>
,
ebPt + kNt eb + kNt

(2.13)

for t t1 . Hence liminf t Nt > ( eb )/k > 0 and we obtain a contradiction. Therefore
W + (E0 ) lies on Y . Similarly, if there exists a solution (Nt ,Pt ) of (2.1) with N0 ,P0 > 0 such

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A discrete-time host-parasitoid model

that limt (Nt ,Pt ) = E1 = (( 1)/k,0), then for any  > 0 there exists t2 > 0 such that
Nt > ( 1)/k  if t t2 . Since bN > 1, we choose  > 0 such that b(( 1)/k ) >
1. But then
Pt+1 >



1
 1 ebPt ,
k

(2.14)

for t t2 implies liminf t Pt > 0 and we obtain a contradiction. Therefore W + (E1 ) lies
on Y and system (2.1) is uniformly persistent by Hofbauer and So [6, Theorem 4.1].
We summarize the above discussion in the following theorem.
Theorem 2.1. Dynamics of system (2.1) can be summarized below.
(a) If < 1, then solutions of (2.1) all converge to E0 = (0,0).

In addi(b) If > 1, then system (2.1) has another boundary steady state E1 = (N,0).

tion if bN < 1, then solutions of (2.1) with N0 > 0 all converge to E1 . If bN > 1,
then system (2.1) has a unique interior steady state E2 = (N ,P ) and (2.1) is
uniformly persistent, that is, there exists M > 0 such that liminf t Nt M and
liminf t Pt M for all solutions (Nt ,Pt ) of (2.1) with N0 > 0 and P0 > 0.
3. Discussion
In this short chapter we investigated a model proposed by May et al. [10], where parasitism occurs before density dependence and density dependence depends only on the
remaining population that escaped from being parasitized. The model exhibits simple
asymptotic dynamics. Both populations go to extinction if the intrinsic growth rate of
the host is less than 1. When the host intrinsic growth rate is greater than 1, then the host
can stabilize in a positive steady state N in the absence of the parasitoid. Therefore the
parasitoid population becomes extinct if bN < 1, where bN can be interpreted as the
Both populations
growth rate of the parasitoid when the host is stabilized at the level N.

can coexist indefinitely if > 1 and bN > 1.

Notice the per capita population growth rate of the host in the absence of the parasitoid population is a decreasing function of the host population. Allee eects occur
when the per capita growth rate of a species is initially an increasing function of the
population size [1]. Allee eects may due to a variety of causes ranging from mating limitation, predator saturation, and antipredator defense and so forth. Among these is the
uncertainty of finding mates to reproduce or lack of cooperative individuals to exploit
resources eciently in spars populations. We refer the reader to [1, 2, 4, 5] for more biological discussion about Allee eects. See also [3, 79, 1114] and references cited therein
for models of Alee eects. We will next incorporate Allee eects into the host population and examine the Allee eects upon the dynamics of the host-parasitoid interaction
studied in this manuscript.
References
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[2] M. Begon, J. Harper, and C. Townsend, Ecology: Individuals, Populations and Communities,
Blackwell Science, New York, 1996.

S. R.-J. Jang and J.-L. Yu 455

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Sophia R.-J. Jang: Department of Mathematics, University of Louisiana at Lafayette, Lafayette,
LA 70504-1010, USA
E-mail address: jang@louisiana.edu
Jui-Ling Yu: Department of Applied Mathematics, Providence University, Taichung 43301, Taiwan
E-mail address: jlyu@pu.edu.tw