ALLEE EFFECT, EMIGRATION AND IMMIGRATION IN A CLASS OF PREDATOR-PREY MODELS

EDUARDO GONZALEZ-OLIVARES , JAIME MENA-LORCA HECTOR MENESES-ALCAY, BETSABE GONZALEZ-YANEZ Grupo Ecolog Matemtica, Instituto de Matemticas, a a a Ponticia Universidad Catlica de Valpara o so. Casilla 4950, Valpara so, Chile. E-mails: ejgonzal@ucv.cl, jmena@ucv.cl, hmeneses@ucv.cl, betsabe.gonzalez@ucv.cl JOSE D. FLORES Department of Mathematical Sciences The University of South Dakota, 314 East Clark Street, Vermillion, SD 57069, USA E-mail: jflores@usd.edu

In this work we analyze a predator-prey model proposed by Kent et. al. in16 , in which two aspect of the model are considered: an eect of emigration or immigration on prey population to constant rate and a prey threshold level for predators. We prove that the system when the immigration eect is introduced in the model has a dynamics that is similar to the Rosenzweig-MacArthur model. Also, when emigration is considered in the model, we show that the behavior of the system is strongly dependent on this phenomenon, this due to the fact that trajectories are highly sensitive to the initial conditions, in similar way as when Allee eect is assumed on prey. Furthermore, we determine constraints in the parameters space for which two stable attractor exist, indicating that the extinction of both population is possible in addition with the coexistence of oscillating of populations size in a unique stable limit cycle. We also show that the consideration of a threshold level of prey population for the predator is not essential in the dynamics of the model.

This work is partially supported by DI-PUCV project 124.711/2007, and by the Department of Mathematics at USD.

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1. Introduction In this paper we analyze the predation model proposed by Kent et al.16 where they model the case of resource subsidy supposing that the resource is a limitation prey to predators and assuming inux (immigration) or outux (emigration) of prey. They arm that the system is: (i) stabilized by an inux of prey in the form of a rescue eect, and (ii) destabilized by an outux of prey in the form of an Allee eect. In Population Dynamics, any mechanism that can lead to a positive relationship between a component of individual tness and either the number or density of conspecic can be termed a mechanism of Allee eect 5,25 , or depensation 4 , or negative competition eect 29 . The outux of prey to constant rate can be considered as Allee eect because a change on interaction dynamics is provoked, for instance, due to dicult of encountering mate. Other implicit assumptions for the proposed continuous time model model are; the populations are uniformly distributed in the environment, neither sex and nor age structure are considered, and abiotic phenomenon is not inuencing on the growth of both populations. In this paper we show that when immigration or emigration are not assumed, the model proposed in16 is topological equivalent to well known Rosenzweig-MacArthur predation model22,28 , for which the eects of entrance or exit of individual prey will be considered by modication to this last model. In order to make an exhaustive study of the proposed system we consider separately the immigration and emigration eects. The inux (immigration)or outux (emigration) of prey in the interaction is of constant rate, for which the new model is not of Kolmogorov type system6 . The lack of these mathematical properties of Kolmogorov type models provoke significant changes in the dynamics of new system with respect to the original Rosenzweig-MacArthur model when the emigration is assumed dierent to zero. Moreover, we show that if immigration (the inux) is considered, the new system has a similar behavior to the Rosenzweig-MacArthur model19,23 ; that is, there exists a parameter set for which the unique equilibrium point at interior of the rst quadrant is globally asymptotically stable or else the existence of a unique limit cycle is assured. We prove that there is a separatrix curve determined by an attractor point in the second quadrant, which divides the behavior of the trajectories. This result implies that the proposed model when prey emigrates at con-

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stant rate, is highly sensitive to initial conditions, since it is possible to nd points close enough in the phase plane whose trajectories can have dierent -limit, an equilibrium point at the second quadrant and the unique positive equilibrium point in the rst quadrant, moreover this later singularity could be surrounded by a unique limit cycle. Then, in this sense, a unique limit cycle can coexist with the extinction of both species depending only on the initial conditions. We consider that the extinction of both species is a result inherent to the Allee eect, and in no way is consequence of the combine presence of both, an Allee eect on prey and a predator functional response type II. Similar conclusion has been shown in13 by using another type of functional response or another predator growth equation7,10,12 . 2. The Model In this section we analyze the consequences of the emigration (outux) and the immigration (inux) eect of prey in a predation model. In the analysis of the model it is assumed that prey are consumed by predators at rate per capita that depends on prey abundance according to the classic disk Qx equation h(x) = Qhx+1 proposed by Holling18 . The deterministic continuous time model is described by the autonomous bi-dimensional dierential equations system
X : dx dt dy dt

= r (x + ) 1 Q(x) = P1+Qhx y

x K

Qxy Qhx+1

(1)

where = (r, K, Q, h, P, ) R6 and R, and the parameters have the + following meanings: r is the intrinsic rate, that is the average rate of prey births per capita in a pristine environment, is the average size of prey immigrants for positive values or emigrants for negative values, or intrinsic prey ux into or out of a prey population of given carrying capacity, K is the carrying capacity of prey at which births and migration reduce to zero, Q is a the predators average searching rate for prey, h is the handling time for each encountered prey, P is a conversion ratio of consumed prey into viable predator ospring, and is the size of prey that sustains one predator, and replaces it with a single ospring when it dies or the marginal subsistence demand for prey;

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also, it is considered as the threshold prey density below which predator density declines. We observed that in the system (1) which is dened in the rst quadrant, = {(x, y) R2 : x 0, y 0} = R+ R+ 0 0 the y-axis is not an invariant set, that is, the system is not of Kolmogorov type6 , except for the case = 0. The equilibrium points of system (1) (or singularities of vector eld X ) 1 are P = (, 0), PQh = ( Qh , 0), PK = (K, 0) and Pe = (xe , ye ), which is the positive(interior) equilibrium point with xe = and ye = r Q 1 K ( + )(1 + Qh).

Then, ye > 0, if and only if < K. In the next two subsections we will analyze separately the two cases; (a) emigration of the prey when R+ , and (b) immigration of prey when R both cases in the framework of the Rosenzweig-MacArthur predator-prey model22,28 . 2.1. Immigration Here we consider the eect of immigration (inux) of prey in model (1). That is, assuming the case R+ we have the following result. Lemma 2.1. The system (1) is topologically equivalent to the following system
X : dx dt dy dt

= r (x + ) 1 =
px x+a

x K

qxy x+a

c y

(2)

with = (r, K, q, a, p, c, ) R7 . + Proof. Qx First we reparametrize the function h(x) = Qhx+1 by using the substiqx 1 1 tutions a = Qh and q = h , hence h(x) has the form h(x) = a+x . a) We can see clearly that in the second equation of system (2),
ac (p c) x pc px c= , x+a x+a and using the second equation from system (1) we have

P q(x ) P Q(x ) = . 1 + Qhx a+x