A Delayed Predator-Prey Model with Fear Effect and Cannibalism ()
1. Introduction
Predation serves as the primary determinant of prey mortality. During the course of evolution, prey has developed a variety of sophisticated strategies, such as detecting, eluding, or combating predators, while actively searching for new food sources [1]-[3]. A growing body of research has demonstrated that the mere presence of predators can exert a substantial influence on the physiological traits and behavioral patterns of prey. In other words, the presence of a predator has a greater effect on the prey than the predator directly hunting. Zanette et al. [4] studied the effects of predators on the reproductive habits of songbirds and found that the presence of fear factors influenced the birth and survival rates of songbirds. Wang et al. [5] proposed a specific mathematical model to characterize the role of the fear effect on predator-prey systems:
where prey and predator density are denoted by
and
,
reflects the fear level of the prey due to perceiving the risk of predation,
is the natural birth rate of prey in the absence of predators,
is the cost of the antipredation defense of the prey due to fear and it is monotonically decreasing in
and
,
is the natural death rate of prey,
is the density-dependent death rate of the prey due to intraspecies competition,
is the functional response function which is independent of
, and
is the predator death rate. It obtained some results on how the fear effect affects the dynamic behavior of predator-prey models through mathematical analysis and numerical simulations. For more related research, see [6]-[10].
Certain predator populations are capable of consuming their own kind to gain energy, enabling them to survive in the absence of prey. And we study such models by considering cannibalism. In some primates [11], fish [12], carnivorous mammals [13] and spiders [14], cannibalism [15], also known as intraspecific predation, involves eating offspring or siblings. This phenomenon is sometimes referred to as the “lifeboat mechanism” because it prevents the extinction of predator communities. Depending on the rate of cannibalism, cannibalism can have a positive or negative impact on the population. Many scholars have been inspired to study and produce many remarkable research results. Therefore, for some predators, it makes more sense to include predators in a stage structure model as well. Deng et al. [16] proposed a Lotka-Volterra prey-predator with predator cannibalism:
where parameter
represents the intrinsic growth rate of the prey,
represents the carrying capacity of the prey in the environment,
represents the rate of predation,
represents the rate at which prey biomass is converted into predator birth,
represents the rate at which cannibalism is converted into predator birth,
represents the rate at which predators die,
represents the rate at which cannibalism occurs within predator individuals, and
represents the cannibalism half-saturation constant. The final term and the second term in the second equation of the system represent the phenomena of cannibalism.
The existence of delay is common in ecosystem. In ecology, physics, biology and other fields, delayed differential equations are more useful than conventional equations. In ecosystem, changes in growth and development, reproduction processes, and environmental factors can produce time lags, and population conditions are affected by time lags. Considering the time delay factor in the predator model can better reflect the actual situation of the ecosystem. In many current studies, the energy obtained by predators through food does not immediately affect the reproduction of predator groups, and this effect can be described by the Holling-type functional response function, which often displays time delay. In the delayed predator-captor model, delay may affect the stability or instability of prey density due to predation (see [17]-[20]). Due to new ecological evidence and theoretical advances, researchers are making new advances in modeling various aspects of biological interactions. Holling [21] proposed a more accurate point of view that has become one of the most widely used ecologies. Hussien et al. [22] considered the existence of cannibalism in the predator population, considered the pregnancy delay of the predator population, and added the predator refuge constant to obtain the following model:
(1)
where parameter
represents the cannibalism coefficient of the predator population,
represents the half-satiation constant of cannibalism, and
represents the predator refuge constant. The results show that reducing the cannibalism rate will destroy the coexistence equilibrium point and make the system close to periodic dynamics. On this basis, the Holling Type II functional response, based on the traditional Lotka-Volterra model, is a function of increases, concave, smooths out, and saturation at high prey numbers. Many authors have studied predator-prey models that use this functional response function, both with and without time delay, and even with spatial dependence (see [23] [24]).
In natural ecosystems, long-term monitoring of predator populations can reveal patterns related to cannibalism and density. Rudolf [25] studied dragonfly predation models, providing the first experimental evidence for the indirect effects of cannibalism behavior and density constraints on prey. The study of biological populations and the analysis of related ecological phenomena have certain relevance to the study of the relationship between cannibalism and density in fish populations. To some extent, it shows the importance and significance of long-term monitoring of predator populations in natural ecosystems. Meanwhile, prey animals are not only affected by the immediate presence of predators (fear effect), but predators also sometimes resort to cannibalism when resources are scarce. Moreover, biological processes such as reproduction and development take time, which is captured by the time-delay factor. The previous study did not couple these parts in the same model to accurately describe how these factors interact over time and affect the stability and dynamics of the predator-prey system. Therefore, assuming that young individuals have density limitations due to their dependence on nutritional or biological resources. We propose the following model:
(2)
with the initial conditions
(3)
The rest of this paper is organized as follows. In Section 2, we establish the well-posedness of solutions of system (2). In Section 3, we investigate the existence and stability of equilibrium and the existence of Hopf bifurcation are investigated. In Section 4, we analyze the stability of Hopf bifurcation. In Section 5, we conducted some numerical simulations. Finally, a brief conclusion is given in Section 6.
2. Well-Posedness
In this section, we will prove the well-posedness of model (2) and obtain the following theorem.
Theorem 1. For any initial value
, system (2) has a unique solution satisfying (3), which exists globally in
, is nonnegative, and remains bounded. Moreover, the region
is both positively invariant and attractive for (3), where
Proof. According to [26] (Lemma 4), we derive that every solution of (2) with the initial condition (3) is positive, that is, any solutions remain positive when
in the region
. The following is a proof of well-posedness.
Firstly, similar to the Lyapunov function as in [22], let
where
are positive constants. After calculation, the
Set
, then
Choose a constant
such that
take
, then
. Since
, so that
has a maximum value of
.
Applying differential inequality theory results, we obtain
As
, we have
. Therefore, all the solutions of system (2) are confined in the region
This completes the proof.
3. Equilibria and Its Stability
In this section, we will show all equilibrium of the model (2) and analyze its stability.
First, we can easily obtain the following results:
1) The trivial equilibrium point
;
2) The predator-free point
, where
always exist in
;
3) The prey-free point
, where
, exists if
;
4) The coexistence equilibrium point
satisfying
By solving the above equations, we get
The equilibrium point exists only if
and
satisfies the equation
where
According to the Descartes rule of sign and
, we find that it exists at least one positive root.
Next, we will discuss the stability of the equilibrium point. To study its local stability, we linearize the system at equilibrium point
to obtain the Jacobian matrix.
Let
, and for simplicity, we still denote them as
. The linearized system is
where
(4)
Then, the characteristic equation of the above system at
can be determined by
(5)
Theorem 2. The trivial equilibrium point
is unstable for all
.
Proof. Substitute
into the characteristic Equation (5) to obtain
and further derive the eigenvalues
. Then, if
then
is a source and if
then
is a saddle point. So,
is always unstable for all
.
Theorem 3. Assume that
and
. Then, the predator-free equilibrium point
is asymptotically stable.
Proof. Substitute
into the characteristic Equation (5) to obtain
where
, after calculation, one eigenvalue is obtained as
, and the other eigenvalue is obtained from the following equation
For
, it is obtained
, if and only if
, the eigenvalue is negative, but when the previous assumption is not met, the eigenvalue
is positive and the equilibrium point
is the saddle point.
Now, when
, by [19] proposition 1, let
, the eigenvalue
has a negative real part when
and
is satisfied. Otherwise, the eigenvalue
has a positive real part.
Therefore, for any
, if
, the predator-free equilibrium point is locally asymptotically stable, while it is an unstable point for
when the assumption doesn’t hold.
This proves the theorem.
Theorem 4. Assume that
and
, where
. Then, the prey-free equilibrium point
is locally asymptotically stable for all
.
Proof. Substitute
into the characteristic Equation (5) to obtain
(6)
where
. The characteristic Equation (6) becomes
(7)
after calculation, we get
In order to obtain the presence of negative real parts in the eigenvalues, it holds when
and
, that is
direct computation gives the following two expressions
where
.
Therefore, the prey-free equilibrium point
is locally asymptotically stable if the above assumptions hold.
This proves the theorem.
Theorem 5. The following results hold:
1) If
and
, or
holds, then the equilibrium point
is locally asymptotically stable for any
;
2) If
and
holds, then the equilibrium point
is locally asymptotically stable for
,
is unstable when
;
3) If
or
and
holds, then the equilibrium point
is locally asymptotically stable for
,
is unstable when
.
Proof. Substitute
into the characteristic Equation (5) to obtain
The above characteristic equation becomes
(8)
where
For
, the characteristic equation is
(9)
According to the Routh-Hurwitz criteria, if and only if
, Equation (9) has two roots and both of its real parts are negative. Hence,
is locally asymptotically stable.
Now, when
, assuming
is a root of Equation (8), substituting
into it yields
then, by separating the real and imaginary parts of the above equation, it is obtained that:
(10)
By calculation, we obtain:
(11)
where
and
.
In the following, we will analyze the roots of Equation (11).
1) If
and
, or
holds, then Equation (11) has no positive roots.
2) If
and
holds, then Equation (11) has two positive roots, which are
and
, respectively, where
3) If
or
and
holds, then Equation (11) has a positive root about, which is
;
In summary, it can be assumed that Equation (11) has two positive roots, denoted as
, which can be substituted into the equation to obtain
Then,
can be expressed as
Denote
This proves the theorem.
4. Hopf Bifurcation Analysis
In this section, the possibility of occurrence of Hopf bifurcation at the coexistence equilibrium point is discissed. Through the analyses in the preceding sections, we have known that Hopf bifurcation will occur under the appropriate conditions. According to Theorem (5), Equation (8) has a pair of purely imaginary roots
at
, and all other roots have non-zero real parts. Therefore, let
be the pair of complex conjugate roots of Equation (8) at
, where
.
According to the stability theory of time-delay differential equations, it can be concluded that when
, the system is stable and when
, the real part of the eigenvalues of the linearized system is discussed.
Theorem 6. If
, a Hopf bifurcation occur at the positive equilibrium point
.
Proof. Firstly, we will verify the transversality conditions. Take the derivative of both sides of characteristic Equation (8) with respect to
, we get
(12)
Next,
Hence,
Clearly,
.
Therefore, the transversality conditions are satisfied, that is, the system has Hopf bifurcation at the positive equilibrium point
.
This proves the theorem.
Next, we will explore the properties of the Hopf bifurcation, including the direction and stability. The direction of the periodic trajectory of the bifurcation of the positive equilibrium point
at the critical of delay
and the stable periodic solution will be given through the center manifold and normal form theory.
Theorem 7. The following statements hold:
1) If
, then a supercritical (subcritical) Hopf bifurcation occurs at
;
2) If
, then the bifurcating periodic solution is asymptotically stable (unstable);
3) If
, then the period of the bifurcating periodic solution increases (decreases).
Proof. Linearizing the system (2) in
, yields the following time-delay differential equation:
(13)
where
and
,
, with
where
are given in Equation (4),
According to the Riesz representation theorem, there exists a bounded variation function
in
such that
In fact, it can be chosen
where
is Dirac delta function and is defined as
For
, define
and
Then, system (13) is equivalent to the following abstract operator equation:
(14)
For
, the adjoint operator
of
is defined as
and define the bilinear inner product as
(15)
where
and
,
and
are adjoint operators. The eigenvalue corresponding to
is
. It is easy to know that
is also the eigenvalue of the conjugate operator
.
Let
where
,
.
and
are the eigenvectors of
and
to the eigenvalues
and
, respectively. Moreover, determine the parameter value of
, such that:
(16)
According to Equation (15), we have
Therefore, due to (16), it is obtained that
According to the theory in [27], we can obtain the property of Hopf bifurcation. Calculating the direction of the Hopf bifurcation and the stability coefficient of the periodic solution at the positive equilibrium point
:
where
with
and
Based on the above analysis and the expressions for
and
, we get the following values:
The above parameters determine the Hopf bifurcation.
5. Numerical Simulation
In this section, we will show some numerical simulation results to support the above research.
For this purpose, we choose the initial value condition
and refer to some parameters in [22] to obtain the following parameters
(17)
In this case, there is a unique coexistence equilibrium point
, and
.
Firstly, fixed parameter (17), we obtained the solution and phase diagram of the positive equilibrium point, as well as the Hopf bifurcation diagram, as shown below.
Figure 1. The Hopf bifurcation diagram.
Figure 2. Solution and phase diagram of system.
From Figure 1, we can find that the system has a Hopf bifurcation as the parameter
passing the bifurcation point
(see Figure 1). From Figure 2, the solution of the system is approached to the coexistence equilibrium point
starting from the initial point (see Figure 2).
Next, we analyze the impact of the parameters
and
on the system.
1) Taking
, other parameters are the same as in (17), the numerical simulations are as follows.
Figure 3. The system solution and phase diagrams of cannibalism rate in small-parameter predator individuals.
We can see from Figure 3 that the cannibalism rate
of the rate at which cannibalism occurs within predator individuals below a specific value destabilizes the coexistence equilibrium point and the system has periodic solution (see Figure 3).
2) Taking
, other parameters are the same as in (17), the numerical simulations are as follows.
Figure 4. The system solution and phase diagrams of cannibalism rate in big-parameter predator individuals.
From Figure 4, we know that increase the conversion rate causes destabilizing the coexistence equilibrium point and the system has a stable limit cycle (see Figure 4).
Finally, we analyze the impact of the time delay
on the model, fixed parameter in (17), we get the following results:
1) Taking
, the numerical simulations are as follows.
Figure 5. Solution and phase diagram of model when
.
2) Taking
, the numerical simulations are as follows.
Figure 6. Solution and phase diagram of model when
.
We can see from Figure 6 that when the pregnancy delay parameter is greater than
, the system has a Hopf bifurcation (see Figure 5 and Figure 6).
6. Conclusions
This paper proposes a predator-prey model that considers the existence of the fear effect and the gestation delay, as well as cannibalism. Firstly, we prove the positivity and boundedness of the system (2). Secondly, it is obtained that the system (2) has four possible non-negative equilibrium points. The local stability of them is studied for
, and the stability conditions are determined. The existence of Hopf bifurcation as a function of
is proved. The stability and direction of the bifurcated periodic dynamics are investigated using the center manifold and normal form theory. Finally, numerical simulation results present the influence of different model parameters on the dynamics of system.
The research results indicate that cannibalism and delay pregnancy in predator populations have significant impacts on population dynamics. The final state and stability of a population mainly depend on predation behavior and parameter configuration. The research findings provide a new perspective for understanding population interactions in ecosystems, emphasizing the crucial role of cannibalism and delayed pregnancy of predator populations in ecosystem stability and population dynamics. Future research can further explore the dynamic characteristics of population models under different predation relationships, which provide insight into species interactions and stability in ecosystems.
Acknowledgements
The authors are grateful to Dr. Danfeng Pang for her valuable suggestion leading to a substantial improvement of the manuscript.