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1. Introduction
Traditionally, the researchers usually connect the predator and their prey only through direct predations [1, 2]. In the recent decades, lots of experiments show that the relationship between the predator and the prey is far more complex than hunting. For one aspect, due to the fear effect of the predation [3–5], the prey will choose to adopt the antipredator behaviours, such as moving to new habitats, searching for safer foods, and other physiological changes, aiming at avoiding the predation. This will definitely help to decrease the death rate of the prey. However, the growth rate of the preys will also be decreased because of the extra costs of adopting antipredator behaviours. Therefore, there could be the trade-off of the fear effect on the growth of the prey. Also, it will be important to study the impacts of the fear effect of prey on the dynamic behaviours between the prey and their predators.
Recently, some scholars have also considered the influence of fear effect on the dynamic behaviours of predator-prey models. For example, Wang et al. [6] first proposed a predator-prey model with the fear effect. In their paper, they have demonstrated a correlation between the fear effect and the direct effect of predators on prey and gave three specific expressions for the fear effect. They also demonstrated that the fear effect does not affect the dynamic behaviours of the system with the Holling-I response function. However, when the functional response is Holling-II, higher levels of fear can stabilize the system by eliminating the presence of oscillatory behaviours. When the appropriate birth rate and fear effect are selected, the system will have a limit cycle. The authors in [7] established a Beddington–DeAngelis predator-prey model with fear effect, refuge, and harvest. They found that there is a critical value of the fear effect, and the two species can continue to exist for a long time through positive density levels. There are also other critical values so that both species can exist at a positive density, but their density levels swing periodically over time. Zhang et al. [8] proved that the system exhibits multiple spatiotemporal patterns due to spatial memory delay and nonlocal fear effect delay. Researchers explored the impact of the fear effect and the Leslie–Gower function on the dynamical behaviours of the predator-prey model and found that as the fear effect increases, the dynamical behaviours of the system switches multiple times until the prey eventually becomes extinct, while the predator survives due to the presence of alternative prey in [9]. For more relevant research on the fear effect, refer to the literature [10–14].
When the number of predators reaches a certain level, the prey will have a fear effect and show antipredator behaviours, resulting in a change in the functional response between the predator and the prey. Therefore, we need to establish a model with threshold conditions. But most predator-prey models are described by ordinary differential equations with continuous right-hand sides, but these models cannot reflect the influence of refuge, group defense, and other factors on population dynamics. Therefore, a discontinuous right-hand nonsmooth dynamic system widely used in mechanics has attracted the attention of ecologists [15–19]. This kind of system is called the Filippov system or switched system, which provides a basic framework for the establishment of many mathematical models with practical significance [20–22]. Therefore, based on the above research, this study proposes a Filippov predator-prey model, which depends on the number of predators using the threshold strategy. Our model extends the existing predator-prey model with the fear effect by introducing a threshold strategy to describe the impact of the fear effect when the number of predators exceeds the threshold. When the number of predators is below the threshold, the functional response is Holling-I. When the number of predators is above the threshold, the functional response changes to Holling-II, and the fear effect is produced.
This study is organized as follows: In Section 2, a Filippov predator-prey model with fear effect is established, and the basic theory and related definitions of the Filippov system will be provided. In Section 3, the dynamic behaviours of the two subsystems are analyzed. In Section 4, the sliding region, sliding dynamics, and the existence of various equilibria of the model are analyzed theoretically. In Section 5, the regular/virtual equilibrium bifurcation, boundary equilibrium bifurcation, and global sliding bifurcation are numerically verified. Finally, the conclusions are presented in Section 6.
2. Model Formulation and Preliminaries
2.1. Model Formulation
We use the classical Lotka–Volterra model to describe the interaction between the predator and prey [23], i.e.,
We assume that the fear effect of the prey to their predator is triggered on by a threshold value of the density of predators
(1)
(2)
(3)
(4)
A simple form of
So, systems (2) and (3) can be integrated into the following nonsmooth dynamical system:
2.2. Preliminaries
To further analyze the Filippov system, here are some symbols [24–26]. Let
For convenience, we denote
Then, system (4) with (6) can be rewritten as the following Filippov system:
Furthermore,
Let
Accordingly, the boundary will be classified as follows:
(i)
(ii)
(iii)
For the convenience of the later study, it is useful to list the definitions of the various equilibria of the Filippov system [16, 17] for this paper.
Definition 1.
If a point
Definition 2.
If a point
Definition 3.
If
Definition 4.
If
3. Dynamics Analysis of Subsystems
The Filippov system (8) consists of three parts: two smooth subsystems and a discontinuity boundary, so it is necessary to study the dynamical behaviours of two subsystems before analyzing the complete Filippov system. In this section, we analyze the existence and stability of the equilibria of the two subsystems separately.
3.1. Dynamics of the Subsystem
When
Theorem 5.
(I) The trivial equilibrium
(II) The boundary equilibrium
(III) The interior equilibrium
Proof.
We analyze the stability of the equilibria by analyzing the corresponding Jacobian matrix for each equilibrium. Now, the Jacobian matrix at
Since
The trace and determinant corresponding to the matrix
Theorem 6.
When
Proof.
The two functions on the right-hand side of system (1) are denoted by
For
When
3.2. Dynamics of the Subsystem
For the subsystem
Notice that the equilibria
Theorem 7.
The trivial equilibrium
Proof.
The Jacobian matrix at
Obviously, we can get
The following lemmas [6] give the stability analysis regarding the boundary equilibrium
Lemma 8.
The boundary equilibrium
Lemma 9.
The boundary equilibrium
Lemma 10.
The internal equilibrium
It is unstable if
Lemma 11.
The internal equilibrium
4. Sliding Region and Equilibria of the Filippov System (8)
In this section, we calculate the sliding region and various equilibria of the Filippov system (8) according to the definitions.
4.1. Sliding Segment and Region
Through simple calculation, the following Lie derivatives can be obtained:
Theorem 12.
(I) If
(II) If
Proof.
(I) If
Moreover, there are two possible scenarios that can lead to the presence of transversal intersection and crossing region:
or
From (28), we calculate that
(II) If
Similarly, (28) and (29) can be obtained for the crossing region. Simplifying (28) can yield
We next examine the existence of four types of equilibria of the Filippov system (8) according to the definitions in Section 2.2.
4.2. Pseudoequilibrium
Based on Definition 2, we first calculate
Substituting the expressions for
We know from [27] that we only need to discuss the case of the molecular roots of
Let
It can be seen that the intersection of the function with the vertical axis is
From this, it follows that the discriminant of the roots of the derivative equation and the axis of symmetry of the image are
Since all parameters are non-negative numbers,
The next step is to determine the positive roots of the original function based on the derivative.
(A)
In this case,
(B)
In this case, the sign of
(I)
In this case, since
(II)
Combining all the above conditions shows that
(i) When
(ii) When
(iii) When
(iv) If
(v) If
[figure(s) omitted; refer to PDF]
Next, we analyze the stability of the pseudoequilibria. We can see from Figure 1 that
We summarize the above in Table 1.
Table 1
Summary of pseudoequilibrium.
Condition | The case of pseudoequilibrium |
4.3. Regular Equilibria
For the subsystem
Moreover, about regular equilibria for the subsystem
The expressions of
Similarly, when assuming that
Based on the above analysis, when both
We summarize the regular and virtual states of the internal equilibria under different conditions in the following Table 2.
Table 2
Summary of regular equilibria.
Prerequisite | Size relation between | Regular and virtual state of equilibria |
4.4. Boundary Equilibrium
The boundary equilibrium of the Filippov system (8) satisfies the following equation:
We denote
So, the coordinates of the boundary equilibria are given in the following equation:
Therefore, the boundary equilibria
4.5. Tangent Points
According to Definition 4, a point
The coordinates of the two tangent points can be obtained by solving the above equations (43) and (44):
5. Sliding Bifurcation Analysis of the Filippov System (8)
5.1. Regular/Virtual Equilibrium Bifurcation
Based on the above analysis of the different equilibria it is known that
The four curves divide the parameter space into seven regions and mark whether the equilibrium in each region is regular equilibrium or virtual equilibrium. Similarly, the boundary equilibria
[figure(s) omitted; refer to PDF]
5.2. Boundary Equilibrium Bifurcation
Boundary equilibrium bifurcation may occur in the Filippov system (8) once
[figure(s) omitted; refer to PDF]
5.2.1. Boundary Node Bifurcation
A simple calculation gives the critical value of
The regular equilibrium
For
5.2.2. Boundary Focus Bifurcation
The critical value
For
The boundary equilibrium
5.3. Global Sliding Bifurcation
It can be seen from [6] that there may be standard periodic solutions that lie entirely in the region
It is known from [29] that standard periodic solutions can collide with sliding segments, and this bifurcation is called a grazing (touching) bifurcation. When
[figure(s) omitted; refer to PDF]
Especially when the bifurcation parameter
5.4. Impact of
In this section, we analyze the effect of the fear effect
It can be seen from Figures 6(a) and 7(a) that the number of predator and prey in the two systems is in a state of periodic fluctuation (i.e. the solution is a periodic solution), but the peak value of fluctuation in Figure 6(a) is higher than that in Figure 7(a). The low-level fear effect ultimately reduce the range of fluctuations in the solutions of the subsystem
[figure(s) omitted; refer to PDF]
6. Discussion
In recent years, the Filippov system has been widely used in integrated pest management, epidemic control, and predator-prey research [15–18, 21, 30]. The Filippov system demonstrates complex dynamic behaviors on sliding segments based on the inheritance of the dynamic behaviors of atomic systems. New equilibria such as pseudoequilibrium, boundary equilibrium and tangent points can appear on the sliding segment. In addition, new bifurcations such as boundary bifurcations and global sliding bifurcations will also appear on the sliding segment.
In this study, we establish and analyze a Filippov predator-prey model with the fear effect. The threshold of the model is the number of predators, and when the number of predators is below the threshold, the prey does not have a fear effect; when the number of predators exceeds the threshold, the prey has fear effect. Also, the functional responses that respond to the predator-prey relationship are different in the two subsystems.
First, we analyze the existence and stability of the equilibrium of the two subsystems, and discuss whether the equilibria of the two subsystems are regular equilibria or virtual equilibria. The analysis results are summarized in Table 2. In addition, the numerical simulation of the regular/virtual equilibrium bifurcation is also carried out. It is found that with the change of threshold and birth rate, the regular and virtuality of the equilibria also change, and there will be two regular equilibria or virtual equilibria, as shown in Figure 2. Second, by using the theory of the Filippov system, we give the conditions for the existence of sliding segments and various equilibria. We focus on the existence and stability of the pseudoequilibrium and summarize the results in Table 1.
Finally, we study the boundary equilibrium bifurcation and global sliding bifurcation. From the above numerical simulation, we find that in the case of
This paper has obtained some meaningful results, of course, there are some shortcomings. For example, we only analyze the possible existence of pseudoequilibrium using a combination of theoretical and graphical methods and discuss the local stability of the pseudoequilibrium, without examining their global stability, which will provide a direction for our next study.
Acknowledgments
This study was partially supported by the National Natural Science Foundation of China (Grant nos. 11961024, 12261104, and 12201196).
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Abstract
Evidences of the fear effect of the prey are well documented, which can greatly affect the dynamics of the predator-prey system. In this study, considering that the fear effect of the prey is triggered on as the density of the predator reaches and exceeds a threshold value, we develop a Filippov system of predator-prey model with the fear effect. In addition, we also include a modify factor of the growth rate of the prey when they adopt the antipredator behaviours due to the fear effect. We initially analyze the dynamics of the two subsystems, including the existence and stability of the equilibria. Utilizing the theory of the Filippov system, we discuss the sliding dynamics, i.e., the existence of sliding region and sliding equilibria. By choosing the threshold as the bifurcation parameter, we investigate the bifurcations near the regular equilibria. The solution curve has three cases: crossing the threshold curve, sliding on the threshold curve, and approaching the pseudoequilibrium. Finally, we numerically verified the existence of the global sliding bifurcation near the regular equilibrium and also the touching bifurcation.
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