Abstract
We consider a periodic time-dependent predator-prey system with stage structure and impulsive harvesting, in which the prey has a life history that takes them through two stages, immature and mature. A set of sufficient and necessary conditions which guarantee the permanence of the system are obtained. Finally, we give a brief discussion of our results.
1. Introduction
In the natural world, there are many species whose individual members have a life history that takes them through two stages, immature and mature. In particular, we have mammalian populations and some amphibious animals in mind, which exhibit these two stages. From the view point of mathematics, the description of the stage structure of the population in the life history is also an interesting problem in population dynamics. The permanence and extinction of species are significant concepts for those stage-structured population dynamical systems. Recently, stage structure models have been studied by many authors [1–3]. This is not only because they are much more simple than the models governed by partial differential equations but also because they can exhibit phenomena similar to those of partial differential models [4], and many important physiological parameters can be incorporated. Much research has been devoted to the models concerning single-species population growth with the stage structure of immature and mature [5, 6]. Two species models with stage structure were investigated by Wang and Chen (1997), Magnusson (1999), Xiao and Chen (2003), as well as Cui and Song (2004). Also, Zhang, Chen, and Neumann (2000) proposed the following autonomous stage structure predator-prey system:where , , , , , , , , and are all positive constants, is a digesting constant.
On the other hand, since biological and environmental parameters are naturally subject to fluctuation in time, the effects of a periodically varying environment are considered as important selective forces on systems in a fluctuating environment. So more realistic and interesting models should take into account the seasonality of the changing environment [7, 8]. This motivated Cui and Y. Takeuchi (2006) to consider the following periodically nonautonomous predator-prey model with stage structure for prey:in which , , , , , , , , , and are all continuous -periodic functions; , , , , , , , are all positive; , are nonnegative; and denote the density of immature and mature population (prey), respectively; and is the density of the predator that only prey on (immature prey). They provided a set of sufficient and necessary conditions to guarantee the permanence of the above system.
Systems with impulsive effects describing evolution processes are characterized by the fact that at certain moments of time, they experience a change of state abruptly. Processes of such type are studied in almost every domain of applied science. Impulsive equations [9, 10] have been recently used in population dynamics in relation to impulsive vaccination [11, 12], population ecology [13, 14], the chemotherapeutic treatment of disease [15], birth pulses [16], as well as the theory of the chemostat [17].
Let us assume that the predator population is affected by harvesting (e.g., fishing or hunting). Further, as we all know that the harvesting does not occur continuously, that is, the harvesting occurs in regular pulses, then let us assume that at some fixed moments, the predator population in system (1.2) is subject to a perturbation which incorporates the proportional decrease. After a perturbation at step , the size of the population becomeswhere is the size of the predator population at step before the impulsive perturbation, and represents the rate at which the predator is harvested.
In this article, we extend the model (1.2) to the case when the predator population is omnivorous and affected by impulsive effects, which is governed by the following system:with the initial value conditionsin which and are the densities of immature and mature prey, respectively; and is the density of the predator that can prey on . When its favorite food is severely scarce, population can eat other resources: . Also, there exists a positive integer such that Here, , the number of the prey consumed per predator in unit time, is called the predator functional response. We assume that there exists a positive constant such thatThe last condition in (1.7) implies that as the prey population increases, the consumption rate of prey consumed per predator increases. The birth rate of the immature prey population is proportional to the existing mature prey population with a proportionality function . For the immature prey population, the death rate is proportional to the existing immature prey population with a proportionality function . The variable parameter represents that the immature prey population is density restriction. The transition rate from immature individuals to the mature individuals is assumed to be proportional to the existing immature population with proportionality coefficient . The death rate of the mature population is of a logistic nature with proportionality coefficient . Also, and are the coefficients that relate to conversion rates of the immature and, respectively, mature prey biomass into predator biomass. The coefficients in (1.4) are all continuous -periodic for . In fact, , , , , , , , and are all strictly positive, and is nonnegative .
The organization of this paper is as follows. In Section 2, we provide some preliminary results which will be useful. In Section 3, we investigate the permanence and extinction of system (1.4) by using analysis technique. In the last section, we give a biological example and a brief discussion of our result.
2. Preliminary Results
Before stating and proving our main results, we give the following definitions, notations, and lemmas which will be useful in the following section.
Let , and be a continuous -periodic function defined on , then we setLemma 2.1. System (1.4) is dissipative.Proof. Define a function such thatin which , . After a simple computations, we havein which is a positive constant. Obviously, the right-hand side of the above inequality is bounded above for all . Hence,where is a positive constant. When , we getAccording to Lemma 2.2 in [9, page 23 ], we derive thatwhich implies that system (1.4) is dissipative. This completes the proof.Next, we consider the following two subsystems of system (1.4):Lemma 2.2 ([18]). The system (2.7) has a positive -periodic solution which is globally asymptotically stable with respect to .Lemma 2.3. If the following conditions hold, then system (2.8) has a unique -periodic solution: and for every solution of system (2.8), Proof. Let and obtain the linear nonhomogeneous impulsive equationLet be the Cauchy matrix for the relevant homogeneous equation, then the solution of (2.12) has the formThe solution will be -periodic if , or ifIn view of conditions (2.9), (2.14) has a unique solutionThen, (2.13) has a unique -periodic solutionHence, (2.8) has a unique -periodic solutionFrom (2.12), we derive thatLetSince conditions (2.9) hold, we note that Then, from Lemma 2.1, the solution of (2.8) is governed byin which and are defined in Lemma 2.1. This completes the proof.
3. Permanence and Extinction of System
Theorem 3.1. System
(1.4) is permanent if and only if in which is the positive -periodic solution of system (2.7).We need the following
lemmas to prove Theorem 3.1.Lemma 3.2. There exist positive constants and such that Proof. In fact, from Lemmas 2.1 and 2.3, the proof of the lemma is obvious. This completes the proof.Lemma 3.3. There exists a positive constant such that Proof. From
Lemma 3.2, one notes that there exists a positive constant such thatwe then havefor According to Lemma 2.2, the following
auxiliary equations:has a globally asymptotically
stable -periodic solution .
Let be the solution of (3.6) with .
By the vector comparison theorem [19], we obtainAccording
to the global asymptotic stability of ,
for any positive constant ,
there exists a such that for all ,Hence, for all ,
we derive thatLetthenConsequently,This completes the proof.Lemma 3.4. Suppose that (3.1) holds, then there
exists a positive constant such that Proof. In view of (3.1), we can choose a
positive constant such thatin whichConsider the following system
with a positive parameter : By Lemma 2.2, system (3.16) has a
positive -periodic solution ,
which is globally asymptotically stable. Let be the solution of (3.16) with initial condition , ,
where is the positive periodic solution of (2.7).
Hence, for the above ,
there exists such thatfor , According to the continuity of the solution in
the parameter ,
we have uniformly in as .
Hence, for ,
there exists such that, Thus, from (3.17) and (3.18), we
get, Since and are all -periodic, we have
Choose a constant ,
from (3.20), we deriveSuppose that (3.13) is not true,
then for the above ,
there exists a such thatwhere is the solution of (1.4) with the initial
condition .
So there exists a constant such thatthen we derive
thatfor .
Let be the solution of (3.16) with and ,
then by the vector comparison theorem, we obtain By the global asymptotic stability of ,
for the given ,
there exists such that and hence, by (3.21), we getSince together with (1.4) and (1.7), we
haveHence, it follows from Lemma 2.2 in [9, page 23] thatthat is,By (3.14), we know that as ,
which leads to a contradiction. This completes the proof.Lemma 3.5. Assume that (3.1) holds, then there
exists a positive constant such that any solution of system (1.4) with initial conditions
satisfies Proof. Suppose that (3.31) is not true, there
must exists a time sequence , such thatand by Lemma 3.4, we have , .
Hence, for each ,
we choose two time sequences and ,
satisfying and as ,
as well asBy Lemma 3.2, for a given
positive integer ,
there exists such that and for all .
In view of as ,
there exists a positive integer such that as .
Hence, for any ,
we haveIntegrating the above inequality
from to ,
for any ,
then we haveObviously, it follows from (3.33)
thatHence, in view of the
periodicity of and ,
we getBy (3.14), (3.33), and (3.38), there
are positive constants and such thatfor for ,
and for all .
However, (3.39) implies thatfor Then, we getLet be the solution of (3.16) with and ,
then by the vector comparison theorem, we obtainFrom and Lemmas 3.2 and 3.3, we obtain that
for any ,
there is a such that for any ,
For , (3.16) has a globally asymptotically stable positive -periodic solution .
By the global asymptotic stability of ,
for the given , there exists ,
and is dependent of any and such thatand hence, by (3.21), we
getBy (3.44), there is
an such that for all and ,
where .
Hence, from (3.44) and (3.47), we obtainfor all , ,
and .
Since, for any , and ,
by (1.4) and (3.42), we have Integrating from to for any and ,
we obtainHence, by (3.34), (3.40), and (3.41),
we finally derive thatThis leads to a contradiction.
This completes the proof.According to Lemmas 3.2–3.5, we can
directly prove the sufficient part of the theorem. Next, we are ready to show
the necessity of this theorem.
Consider the case ofit is then easy to derive that the predator population must be extinct because of sterilization and impulsive harvesting. Suppose thatWe will show thatIn fact, from (1.7) and (3.53), we know that for any given , there exists such thatNote that for . Sincefor the given , it is easy to show that there exists such thatwe then haveWe now show that there must exist such that . Otherwise, by the last two equations in system (1.4), we haveThis implies that , which is a contradiction.
Let . Note that is bounded. We then show thatOtherwise, there exists such that in which , and By (3.58), we havewhich leads to a contradiction. This implies that (3.60) holds. In view of the arbitrariness of , we get that as . This completes the proof.
Remark 3.6. From the proof of Theorem 3.1 above, we note that the predator population in system (1.4) will be extinct provided thator excessive harvesting, that is,
4. Discussion
Our result could provide a useful insight into the conservation of beneficial animals, especially rare animals. As an example, we depict the case of Oreolalax omeimontis tadpole, Oreolalax omeimontis, and red-eared slider (T. scripta elegans). Oreolalax omeimontis is a rare species of frog found near Mt. Omei in Sichuan (China). The red-eared slider is a native of the Mississippi Valley area of the United States [8, 20]. Since the 1970s, large numbers of red-eared sliders have been produced on turtle farms in the USA for the international pet trade. Red-eared Turtles are traded as pet animals and have been introduced to many countries. They are omnivorous and will eat insects, tadpoles, crayfish, shrimp, worms, snails, amphibians and small fish, as well as aquatic plants, and they hardly may be controlled by a natural enemy. In our model, the variables and represent the density of Oreolalax omeimontis tadpole and Oreolalax omeimontis at time , respectively. The variable describes the density of the red-eared slider at time . It is well known that Oreolalax omeimontis is beneficial to humans because they eat so many insect pests. Ironically, although red-eared sliders have been widely introduced throughout the world, its detrimental effects have been reported by many researchers [21, 22]. The red-eared slider is one of main enemies of Oreolalax omeimontis. To protect these beneficial and rare toads, we must control the amount of red-eared sliders in the habitat of Oreolalax omeimontis. From a control point of view, our aim is to keep red-eared sliders at an acceptably low level with a minimum use of artificial control measures, not to eradicate all red-eared sliders. Hence, in the above example, the problem of nonextinction of populations becomes a description of reasonable harvesting rates .
According to our main result, system (1.4) is permanent if and only if the growth of the predator by foraging prey populations and plus its intrinsic rate of increase is positive on average during the period , and harvesting rates of red-eared sliders during the period are small enough to satisfy thatThese seem to be reasonable from a biological point of view; but it should be noted that condition (3.1) allows the predator to be for some time intervals among That is, under reasonable harvesting rates, the predator can survive together with prey populations even if the growth rate of the former is negative for some seasons during a period. We hope that our result can be used to help protect beneficial animals found in their habitats.
Acknowledgment
This work is supported by the National Natural Science Foundation of China (10671001).