mathematics

Article
Stability and Bifurcation Analysis in a Discrete Predator–Prey
System of Leslie Type with Radio-Dependent Simplified Holling
Type IV Functional Response
Luyao Lv and Xianyi Li *

Department of Big Data Science, School of Science, Zhejiang University of Science and Technology,
Hangzhou 310023, China; 212109701012@zust.edu.cn
* Correspondence: mathxyli@zust.edu.cn

Abstract: In this paper, we use a semi-discretization method to consider the predator–prey model of
Leslie type with ratio-dependent simplified Holling type IV functional response. First, we discuss the
existence and stability of the positive fixed point in total parameter space. Subsequently, through
using the central manifold theorem and bifurcation theory, we obtain sufficient conditions for the flip
bifurcation and Neimark–Sacker bifurcation of this system to occur. Finally, the numerical simulations
illustrate the existence of Neimark–Sacker bifurcation and obtain some new dynamical phenomena
of the system—the existence of a limit cycle. Corresponding biological meanings are also formulated.

Keywords: discrete predator–prey system of Leslie type; Holling type IV functional response;
semi-discretization method; flip bifurcation; Neimark–Sacker bifurcation

MSC: 39A28; 39A30

1. Introduction and Preliminaries

In the past few decades, mathematical models have played a crucial role in the study
Citation: Lv, L.; Li, X. Stability and
of biology. According to different application environments, the interaction between
Bifurcation Analysis in a Discrete
Predator–Prey System of Leslie Type
populations can be expressed as consumer–resource [1], plant–herbivore [2], parasite–
with Radio-Dependent Simplified
host [3], etc. Predator–prey models are the building blocks of ecosystems, as biomass grows
Holling Type IV Functional Response.
from resource masses. This topic plays an important role in ecology [4–16].
Mathematics 2024, 12, 1803. https:// Two of the most famous predator–prey dynamical models are the Lotka–Volterra
doi.org/10.3390/math12121803 model and the Leslie type model. Based on these models, subsequent scholars have
considered more influencing factors, such as the fear effect [9], the Allee effect [12], super-
Academic Editors: Lingju Kong and
predators [13], etc.
Min Wang
In [4], Leslie first proposed that the carrying capacity of the environment for the
Received: 13 May 2024 predator is proportional to the number of prey. So, this form of predator growth is also
Revised: 31 May 2024 called Leslie–Gower type and can be represented as
Accepted: 4 June 2024
Published: 10 June 2024 dy y
= sy(1 − )
dt hx
Therefore, the dynamical relationship between prey and predator may be expressed as
Copyright: © 2024 by the authors. (
dx
Licensee MDPI, Basel, Switzerland. dt = g ( x ) x − f ( x, y ) y (1)
dy y
This article is an open access article dt = sy (1 − hx )
distributed under the terms and
conditions of the Creative Commons where x and y represent prey and predator population sizes or densities, respectively; the
Attribution (CC BY) license (https:// function g( x ) characterizes the growth rate of prey in
the absence of a predator, and may be
creativecommons.org/licenses/by/ represented through logistic growth g( x ) = r 1 − xk ; f ( x, y) is a functional response curve
4.0/). and has many different forms, such as Holling types I–IV [17,18], Beddington–DeAngelis

Mathematics 2024, 12, 1803. https://doi.org/10.3390/math12121803 https://www.mdpi.com/journal/mathematics

Mathematics 2024, 12, 1803 2 of 16

type [19,20], Hassell–Varley type [15,21], etc.; the parameter s signifies the intrinsic growth
rate of the predator; and k and h denote the carrying capacity of the prey and predator
provided by the environment, respectively.
As is well-known, the generalized Holling-IV response function is f ( x, y) = ax2 +mx bx +1
.
There is far less research on the Holling-IV response function than on the Holling-I–III
response functions. Here, we assume that the prey growth follows logistic growth, and the
functional response f ( x, y) is taken as a Holling-IV response function. Then, the system (1)
can be reformulated as (
dx x mx
dt = rx (1 − k ) − a+ x2 y
dy y (2)
dt = sy (1 − hx )
Here, m and a are positive constants, the parameter m is the maximal predator per capita
consumption rate, the parameter a is the number of prey necessary to achieve half of the
maximum rate m, and the parameter r signifies the intrinsic growth rate of prey.
m yx
If one takes the functional response f (x, y) as the ratio-dependent type, i.e., f (x, y) = a+( yx )2
.,
then one has
mxy2
(
dx
dt = rx (1 − xk ) − ay2 + x2 (3)
dy y
dt = sy(1 − hx )
For the sake of the simplicity of mathematical analysis, we now non-dimensionalize
y 2
the system (3). To accomplish this, let xk → u, l → v, rt → τ, hk → l, kmr → α, ka2 → β, and
s
r → δ. Then, one can derive an equivalent to the system (3) as follows:

αuv2
(
du
dτ = u (1 − u ) − βv2 +u2 (4)
dv
dτ = δv(1 − uv )

This continuous system has been discussed in [18], whereas its discrete version has
not been found to be investigated yet. It is very difficult to solve a complicated continuous
equation or system without using a computer. Meanwhile, many models in nature look
more reasonable in their discrete forms. So, it is crucial to consider the discrete version
corresponding to a continuous model.
Now, we use a semi-discretization method to discretize the system (4). Let [t] denote
the greatest integer not exceeding t. Consider the average change rate of the system (4) at
integer number points, namely, the following model:

 1 du = 1 − u([τ ]) − αv2 ([τ ])
u(τ ) dτ βv2 ([τ ])+u2 ([τ ])
(5)
 1 dv = δ(1 − v([τ ]) )
v(τ ) dτ u([τ ])

One can see that a solution (u(τ ), v(τ )) of the system (5) for τ ∈ [0, +∞) has the
following characteristics:
1. On the interval [0, +∞), u(τ ) and v(τ ) are continuous;
du(τ ) dv(τ )
2. dτ and dτ exist everywhere when τ ∈ [0, +∞), possibly except at the points
{0, 1, 2, 3, · · · };
3. The system (5) is true on any interval [n, n + 1) with n = 0, 1, 2, · · · .
The following system can be obtained by integrating the system (5) over the interval
[n,τ] for any τ ∈ [n, n + 1) and n = 0, 1, 2, · · ·

α ( v n )2
 
 ln u(τ ) = 1 − un − ( τ − n ),
u(n) 2
β(vn ) +(un ) 2
(6)
 ln v(τ ) = δ(1 − vn )(τ − n)
v(n) un
Mathematics 2024, 12, 1803 3 of 16

Subsequently, we simultaneously take the exponent with e as the base for Equation (6):
  
α ( v n )2
1− u n − (τ −n)
β(vn )2 +(un )2

u(τ ) = un e (7)
vn
v(τ ) = vn eδ(1− un )(τ −n)


where un = u(n) and vn = v(n). Letting τ → (n + 1)− in the system (7) produces

α ( v n )2
 1− u n −
u n +1 = u n e β ( v n )2 +(un )2
v n
(8)
v n +1 = v n e δ (1− u n )


where the parameters α, β, δ > 0, and they are the same as in (4). The system (8) will be
considered in the sequel.
The rest of this paper is organized as follows. In Section 2, we discuss the existence
and stability of the positive fixed point because of the biological significance. In Section 3,
we provide the sufficient conditions for the existence of the flip bifurcation and Neimark–
Sacker bifurcation. In Section 4, we illustrate the theoretical results derived with numerical
simulations. Finally, some conclusions and discussions are stated in Section 5.

2. Existence and Stability of Fixed Points

We discuss the existence and stability of non-negative fixed points of the system (8) in
this section. The fixed point of the system (8) satisfies the following equation:

1− u − αv2 v
u − ue βv2 +u2 = 0, v − veδ(1− u ) = 0

It is easy to show that the system (8) has two non-negative fixed points E1 = (1, 0)
and E2 = (u∗ , v∗ ) for α < β + 1, where

1+β−α ∗
u∗ = , v = u∗
1+β

The Jacobian matrix of the system (8) at a fixed point E( x, y) is

 
  1−u− αv2 1−u− αv
2
2αu2 v2 2 + u2 2αu3 v 2 + u2
 1 − u + (u2 + βv2 )2 e βv − (u2 + βv2 )2 e βv
J ( E) = 

 
δv2 δ(1− uv ) v)

e 1− u e δv δ ( 1 − u
u2

and its characteristic equation is

F (λ) = λ2 − Tr ( J ( E))λ + Det( J ( E))= 0

where
2
2αu2 v2
   
1−u− αv 2 + u2 δv δ(1− v )
Tr ( J ( E)) = 1 − u + 2 e βv + 1− e u
(u + βv2 )2 u
2
2αu2 v2 2αuv3
  
v 1−u− αv +δ(1− uv )
Det( J ( E)) = 1−u+ 2 ( 1 − ) + e βv2 +u2
(u + βv2 )2 u (u2 + βv2 )2
Before analyzing the properties of the fixed points of the system (8), we provide the
following definition and Lemma [22–25].

Definition 1. Let E( x, y) be a fixed point of the system (8) with multipliers λ1 and λ2 .
(i ) If |λ1 | < 1 and |λ2 | < 1, E( x, y) is called sink, then a sink is locally asymptotically stable.
(ii ) If |λ1 | > 1 and |λ2 | > 1, E( x, y) is called source, then a source is locally asymptotically
unstable.
(iii ) If |λ1 | < 1 and |λ2 | > 1 (or |λ1 | > 1 and |λ2 | < 1), then E( x, y) is called saddle.
Mathematics 2024, 12, 1803 4 of 16

(iv) If either |λ1 | = 1 or |λ2 | = 1, then E( x, y) is called to be non-hyperbolic.

Lemma 1. Let F (λ) = λ2 + Bλ + C, where B and C are two real constants. Suppose λ1 and λ2
are two roots of F (λ) = 0. Then, the following statements hold.
(i ) If F (1) > 0, then
(i.1) |λ1 | < 1 and |λ2 | < 1 if and only if F (−1) > 0 and C < 1;
(i.2) λ1 = −1 and λ2 ̸= −1 if and only if F (−1) = 0 and B ̸= 2;
(i.3) |λ1 | < 1 and |λ2 | > 1 if and only if F (−1) < 0;
(i.4) |λ1 | > 1 and |λ2 | > 1 if and only if F (−1) > 0 and C > 1;
(i.5) λ1 and λ2 are a pair of conjugate complex roots and |λ1 | = |λ2 | = 1
if and only if −2 < B < 2 and C = 1;
(i.6) λ1 = λ2 = −1 if and only if F (−1) = 0 and B = 2.
(ii ) If F (1) = 0, namely, 1 is one root of F (λ) = 0, then another root
λ satisfies |λ| = (<, >)1 if and only if |C | = (<, >)1.
(iii ) If F (1) < 0, then F (λ) = 0 has one root lying in (1, ∞). Moreover
(iii.1) the other root λ satisfies λ < (=) − 1 if and only if F (−1) < (=)0;
(iii.2) the other root −1 < λ < 1 if and only if F (−1) > 0.

Due to biological significance, we only consider E2 . By using Definition 1 and Lemma 1,

the following result can be obtained.

1+ β − α 1+ β − α
Theorem 1. For α < 1 + β, E2 = (u∗ , v∗ ) = ( 1+ β , 1+ β ) is a positive fixed point of the
system (8).
3+ β 1+ β
Let δ1 = 2 + (1+α+4α β)(1+ β)
and δ2 = 1+ β − α . The following statements are true about
the positive fixed point E2 .
(1+ β )2
Case 1. When 0 < α ≤ 3+ β , δ2 ≤ 0 < δ1 :
(1) if 0 < δ < δ1 , then E2 is a sink;
(2) if δ = δ1 , then E2 is non-hyperbolic;
(3) if δ > δ1 , then E2 is a saddle.
(1+ β )2
Case 2. When 3+ β < α < 1 + β, 0 < δ2 < δ1 :
(1) if 0 < δ < δ2 , then E2 is a sink;
(2) if δ = δ2 , then E2 is non-hyperbolic;
(3) if δ2 < δ < δ1 , then E2 is a source;
(4)if δ = δ1 , then E2 is non-hyperbolic;
(5) if δ > δ1 , then E2 is a saddle.

Proof. The Jacobian matrix J ( E) of the system (8) at E2 is

!
α
1+ β + (1+2αβ)2 − (1+2αβ)2
J ( E2 ) =
δ 1−δ

whose characteristic polynomial can be written as

F (λ) = λ2 − Pλ + Q (9)

with
α 2α
P= + +1−δ
1 + β (1 + β )2

 
α 2α 2δα
Q= + (1 − δ ) +
1 + β (1 + β )2 (1 + β )2
 
α 3+β
= −δ
1+β 1+β
Mathematics 2024, 12, 1803 5 of 16

It is easy to see that F (1) = δ(1 − α

1+ β ) > 0 always holds for α < 1 + β. Simple
calculations display that

α 2α α
F (−1) = 2[1 + + ] − δ (1 + )
1 + β (1 + β )2 1+β
1+α+β
= (δ1 − δ)
1+β
α 2α δα
Q−1 = + 2
− −1
1 + β (1 + β ) 1+β
α
= (δ2 − δ)
1+β

We can see that when δ > (=, <)δ1 , then F (−1) < (=, >)0, and when δ > (=, <)δ2 ,
then Q < (=, >)1.
(1+ β )2
Case 1. If 0 < α ≤ 3+ β , then δ2 ≤ 0 < δ1 .
If 0 < δ < δ1 , then F (−1) > 0. But Q − 1 = 1+α β (δ2 − δ) < 0, namely, Q < 1. Lemma 1
(i.1) states that E2 is a stable node, i.e., a sink. If δ = δ1 , then F (−1) = 0; hence, E2 is
non-hyperbolic. If δ > δ1 , then F (−1) < 0, then Lemma 1 (i.3) says that E2 is a saddle.
(1+ β )2
Case 2. If 3+ β < α < 1 + β, then 0 < δ2 < δ1 .
When 0 < δ < δ2 , then F (−1) > 0 and Q > 1. By Lemma 1 (i.4), |λ1 | > 1 and |λ2 | > 1;
therefore, E2 is an unstable node, i.e., a source. When δ = δ2 , F (−1) = 0 and Q = 1. On
one hand, the following applies:
 
α 2α 3+β 1+β
P+2 = + +1− − +2
1 + β (1 + β )2 1+β α
α 2α 2β 1+β
= + + + >0
1 + β (1 + β )2 1+β α

So, P > −2. On the other hand, the following applies:

 
α 2α 3+β 1+β
P−2 = + − − −1
1 + β (1 + β )2 1+β α
α 2α 4 + 2β 1 + β
= + 2
− +
1 + β (1 + β ) 1+β α
 
1 3+β 2 2
= α − 2(1 + β ) α + (1 + β )
α (1 + β ) 1 + β
(1 + β )2
< 0 because < α < 1+β
3+β

Hence, P < 2. Accordingly, −2 < P < 2. By Lemma 1 (i.5), Equation (9) has a pair of
conjugate complex roots λ1 and λ1 with |λ1 | = |λ2 | = 1, implying that E2 is non-hyperbolic.
When δ2 < δ < δ1 , Q < 1. Lemma 1 (i.1) tells us that |λ1 | < 1 and |λ2 | < 1, so E2 is a stable
node, i.e., a sink.
When δ = δ1 , then F (−1) = 0 and E2 is non-hyperbolic.
When δ > δ1 , according to Lemma 1 (i.3), |λ1 | < 1 and |λ2 | > 1. Therefore, E2 is a
saddle. Summarizing the above discussions, we obtain the following Table 1.
Mathematics 2024, 12, 1803 6 of 16

Table 1. Type of the fixed point E2 .

Conditions Eigenvalues Properties

0 < δ < δ1 |λ1 | < 1, |λ2 | < 1 sink (stable node)
(1+ β )2 δ = δ1 λ1 = −1, λ2 ̸= −1 non-hyperbolic
0<α≤ 3+ β
δ > δ1 |λ1 | < 1, |λ2 | > 1 saddle
source (unstable
0 < δ < δ2 |λ1 | > 1, |λ2 | > 1
node)

(1+ β )2
δ = δ2 | λ1 | = | λ2 | = 1 non-hyperbolic
3+ β < α < 1+β
δ2 < δ < δ1 |λ1 | < 1, |λ2 | < 1 sink (stable node)
δ = δ1 λ1 = −1, λ2 ̸= −1 non-hyperbolic
δ > δ1 |λ1 | < 1, |λ2 | > 1 saddle

3. Bifurcation Analysis
In this section, we employ the center manifold theorem and bifurcation theory to
research the local bifurcation problems of the system (8) at the fixed point E2 .

3.1. Main Results

One can see from Theorem 1 that the fixed point E2 is a non-hyperbolic fixed point
when δ = δ1 . As soon as the parameter δ goes through the critical value δ1 , a flip bifurcation
probably occurs near the fixed point E2 . Namely, the bifurcation probably occurs in the
space of parameters
n o
(α, β) ∈ Ω1 = (α, β) ∈ R2+ | 0 < α < 1 + β, β > 0

In fact, a result is obtained as follows.

Theorem 2. Suppose the paramenters (α, β) ∈ Ω1 . Let δ1 = 2 + 4α

(1+α+ β)(1+ β)
. Assume
c2200+ c300 ̸= 0, where c200 and c300 will be defined in the sequel. If the parameter δ varies in a
small neighborhood of the critical value δ1 , then the system (8) experiences a flip bifurcation at the
fixed point E2 .

When δ = δ2 , there is a pair of conjugate imaginary roots with |λ1 | = |λ2 | = 1, which
ensures the necessary condition for a Neimark–Sacker bifurcation to occur. The following
result may be obtained.

Theorem 3. Suppose we have the following parameters:

(1 + β )2
 
(α, β) ∈ Ω2 = (α, β) ∈ R2+ | < α < 1 + β, β > 0
3+β
3+ β 1+ β
Let δ2 = 1+ β − α , and let L be defined in (18). Then, the system (8) undergoes a Neimark–
Sacker bifurcation at the fixed point E2 when the parament δ varies in a small neighborhood of the
critical value δ2 . Moreover, if L < (>)0, then a (an) stable (unstable) invariant closed orbit is
bifurcated out from the fixed point E2 of the system (8).
Mathematics 2024, 12, 1803 7 of 16

3.2. Proof of Main Results

Proof of Theorem 2. First, let Xn = un − u∗ and Yn = vn − u∗ , which transforms the fixed
point E2 to the origin. Then, the system (8) becomes

α(Yn +u ) ∗ 2
1− Xn − u ∗ −
+ u∗ )e β(Yn +u∗ )2 +( Xn +u∗ )2 − u∗

X n +1 = ( X n (10)
Yn +u∗
δ (1− X )
Yn+1 = (Yn + u∗ )e n +u∗ − u∗


Second, giving a small perturbation δ∗ of the parameter δ around δ1 , i.e., δ∗ = δ − δ1

with 0 < |δ∗ | ≪ 1, and letting δn∗+1 = δn∗ = δ∗ , the system (10) is perturbed into

α(Yn +u∗ )2

1− Xn − u ∗ −
 X n +1 = ( X n + u ∗ ) e β(Yn +u∗ )2 +( Xn +u∗ )2 − u ∗


∗
Yn +u
(δ1 +δn∗ )(1− X ) (11)
Y = (Yn + u∗ )e +u∗n − u∗
 ∗n+1


δn+1 = δn∗

Using the Taylor expansion of the system (11) at ( Xn , Yn , δn∗ ) = (0, 0, 0), one has

a010 Yn + a200 Xn2 + a020 Yn2 + a110 Xn Yn


 Xn+1 = a100 Xn +

+ a300 Xn + a030 Yn3 + a210 Xn2 Yn + a120 Xn Yn2 + o (ρ31 )
3





Yn+1 = b100 Xn + b010 Yn + b001 δn∗ + b200 Xn2 + b020 Yn2 + b002 (δn∗ )2


(12)

 +b110 Xn Yn + b101 Xn δn∗ + b011 Yn δn∗ + b300 Xn3 + b030 Yn3
+b003 (δn∗ )3 + b210 Xn2 Yn + b201 Xn2 δn∗ + b120 Xn Yn2 + b021 Yn2 δn∗




+b102 Xn (δn∗ )2 + b012 Yn (δn∗ )2 + b111 Xn Yn δn∗ + o (ρ31 )



p
where ρ1 = Xn2 + Yn2 + (δn∗ )2 .

α 2α 2α
a100 = + 2
, a010 = −
1 + β (1 + β ) (1 + β )2
u∗ α ( β − 3)
 
2α α
a200 = ∗ 2
1 + 2
− + ∗
u (1 + β ) (1 + β ) 2 u (1 + β )3
2α2 α(3β − 1)
a020 = + ∗
u ∗ (1 +
β )4 u (1 + β )3
4α(1 − β)
 
2α 2α α
a110 = − − + ∗
(1 + β )2 u ∗ (1 + β )2 1 + β u (1 + β )3
4u∗ 1 3 1 2
   
α α 2α
a300 = ∗ 2
− + 2 ∗ 2
− − ∗ 2
3 u (1 + β ) 2 u (1 + β ) 2 ( u ) (1 + β )3
2α( β − 3) α ( β − 3) 2( u ∗ )2
   
α 1 1
+ ∗ 2 − + −
( u ) (1 + β )3 u ∗ (1 + β )2 2 1+β (u∗ )2 (1 + β)2 β(1 − 3β)
α(3β − 1)
   
α 2 2α 2α
a030 = ∗ 2 + + α + − 1
( u ) (1 + β )3 (1 + β )2 1 + β ( u ∗ )2 (1 + β )3 3(1 + β )3
 ∗
u −β
  
4α 1 αβ 3β
a210 = ∗ + + 16 −
u (1 + β )2 u ∗ (1 + β ) 2 (1 + β )3 ( u ∗ )2
1 2 2α( β − 3)
   
4α α β α
− − + 1 − −
3(1 + β )2 u ∗ (1 + β )2 2 ( u ∗ )2 (1 + β )3 1 + β (1 + β )2
4(1 − β )
  
α α 1
+ ∗ 2
−2 ∗ 2
−
u (1 + β ) 1 + β u (1 + β ) 2

10α 2 (α + 1)(3β − 1)

α 1

a120 = + −
3u∗ (1 + β)4 u ∗ (1 + β )3 u ∗ (1 + β )2 2
α(3β − 1)
   
16αβ β 2β
+ ∗ 2 − 1 + + 1
( u ) (1 + β )3 ( u ∗ )2 (1 + β ) ( u ∗ )2 (1 + β )3 1 + β
Mathematics 2024, 12, 1803 8 of 16

 
2α α β
+ ∗ 2 − ∗ 2 +1
( u ) (1 + β )2(1 + β ) 2 ( u ) (1 + β )
α2 ( β − 3)
 
+ ∗ 2 − 8 ( 1 + β )
(u ) (1 + β)5 3u∗ (1 + β)2
δ1 (2 − δ1 )
b001 = b002 = b003 = b102 = b012 = 0, b100 = mδ1 , b010 = 1 − δ1 , b200 = −
2u∗

δ1 (2 − δ1 ) δ1 (2 − δ1 ) δ12 (3 − δ1 )
b020 = − , b110 = , b101 = 1, b011 = − 1, b030 =
2u∗ u∗ 6( u ∗ )2
!
δ (3 − δ1 ) δ2 ( δ − 1)
 
δ1 δ 5δ
b300 = ∗ 2
2 − δ1 − 1 , b210 = ∗1 2 − 1 + 1 − 2 , b201 = 1 ∗
2( u ) 3 (u ) 2 2 u
!
δ δ2 δ −1 2(1 − δ1 )
b120 = ∗2 2 1 − 2δ1 + 1 , b021 = 1 ∗ , b111 =
(u ) 2 u u∗

Therefore, we obtain the Jacobian matrix of the system (11) at the fixed point E2
 
2α
α
1+ β + (1+ β )2 − (1+2αβ)2 0
J ( E2 ) =  1 − δ1
 
δ1 0 
0 0 1

and its eigenvalues

λ1 = −1, λ2 = a100 + 2 − δ1 , λ3 = 1
with corresponding eigenvectors

    
1 1 0
ξ 1 =  M , ξ 2 =  N , ξ 3 =  0 
0 0 1

(1+ β)(1+α+ β) 2(1+ β )

where M = 1 + 2α and N = 1+ α + β .
Set T = (ξ 1 , ξ 2 , ξ 3 ), i.e.
 
1 1 0
T=  M N 0 
0 0 1
then
N
− N −1 M
 
N−M 0
T −1 =  M
− N− M
1
N−M 0 
0 0 1
Taking the transformation

( Xn , Yn , δn∗ )T = T (ln , mn , ωn )T

the system (11) is changed into the following form:

g1 (ln , mn , ωn ) + o (ρ32 )
      
l n +1 −1 0 0 ln
 m n +1  =  0 λ2 0mn  +  g2 (ln , mn , ωn ) + o (ρ32 ) (13)
ω n +1 0 0 1 ωn 0
p
where ρ2 = ln2 + m2n + ωn2 .

g1 (ln , mn , ωn ) = c200 ln2 + c020 m2n + c002 ωn2 + c110 ln mn + c101 ln ωn + c011 mn ωn
Mathematics 2024, 12, 1803 9 of 16

+ c300 ln3 + c030 m3n + c003 ωn3 + c210 ln2 mn + c201 ln2 ωn + c120 ln m2n
+ c102 ln ωn2 + c012 mn ωn2 + c021 m2n ωn + c111 ln mn ωn
g2 (ln , mn , ωn ) = d200 ln2 + d020 m2n + d002 ωn2 + d110 ln mn + d101 ln ωn + d011 mn ωn
+ d300 ln3 + d030 m3n + d003 ωn3 + d210 ln2 mn + d201 ln2 ωn + d120 ln m2n
+ d102 ln ωn2 + d012 mn ωn2 + d021 m2n ωn + d111 ln mn ωn

c102 = c012 = c002 = c003 = 0

c200 = γa200 + µb200 + M(γa110 + µb110 ) + M2 (γa020 + µb020 )
c110 = 2(γa200 + µb200 ) + ( M + N )(γa110 + µb110 ) + 2MN (γa020 + µb020 )
c020 = γa200 + µb200 + N (γa110 + µb110 ) + N 2 (γa020 + µb020 )
c101 = µMb011 + µb101 , c011 = µNb011 + µb101
c300 = γa300 + µb300 + M3 (γa030 + µb030 ) + M (γa210 + µb210 )
+ M2 (γa120 + µb120 ), c030 = γa300 + µb300 + N 3 (γa030 + µb030 )
+ N (γa210 + µb210 ) + N 2 (γa120 + µb120 )
c210 = 3(γa300 + µb300 ) + 3M2 N (γa030 + µb030 ) + (2M + N )(γa210 + µb210 )
+( M2 + 2MN )(γa120 + µb120 ), c120 = 3(γa300 + µb300 ) + 3MN 2 (γa030 + µb030 )
+( M + 2N )(γa210 + µb210 ) + ( N 2 + 2MN )(γa120 + µb120 )
c201 = µb201 + µM2 b021 + µMb111 , c021 = µb201 + µN 2 b021 + µNb111
µ( M + N )b111
c111 = µb201 + µMNb021 +
2
d102 = d012 = d002 = d003 =0
d200 = ϵa200 − µb200 + M (ϵa110 − µb110 ) + M2 (ϵa020 − µb020 )
d110 = 2(ϵa200 − µb200 ) + ( M + N )(ϵa110 − µb110 ) + 2MN (ϵa020 − µb020 )
d020 = ϵa200 − µb200 + N (ϵa110 − µb110 ) + N 2 (ϵa020 + µb020 )
d101 = −µMb011 − µb101 , c011 = −µNb011 − µb101
d300 = ϵa300 − µb300 + M3 (ϵa030 − µb030 ) + M (ϵa210 − µb210 )
+ M2 (ϵa120 − µb120 ), d030 = ϵa300 − µb300 + N 3 (ϵa030 − µb030 )
+ N (ϵa210 − µb210 ) + N 2 (ϵa120 − µb120 )
d210 = 3(ϵa300 − µb300 ) + 3M2 N (ϵa030 − µb030 ) + (2M + N )(ϵa210 − µb210 )
+( M2 + 2MN )(ϵa120 − µb120 ), d120 = 3(ϵa300 − µb300 ) + 3MN 2 (ϵa030 − µb030 )
+( M + 2N )(ϵa210 − µb210 ) + ( M2 + 2MN )(ϵa120 − µb120 )
d201 = −µb201 − µM2 b021 − µMb111 , d021 = −µb201 − µN 2 b021 − µNb111
µ( M + N )b111
d111 = −µb201 − µMNb021 −
2
N M 1
γ= , ϵ=− , µ=−
N−M N−M N−M
Assume on the center manifold that

mn = h(ln , ωn ) = h20 ln2 + h11 ln ωn + h02 ωn2 + o (ρ23 )

p
where ρ3 = ln2 + ωn2 . Then, according to (13), we obtain

mn+1 =h(ln+1 , ωn+1 ) = λ2 h(ln , ωn ) + g2 (ln , h(ln , ωn ), ωn ) + o (ρ33 )

h(ln+1 , ωn+1 ) = h20 (−ln + g1 (ln , h(ln , ωn ), ωn ))2
Mathematics 2024, 12, 1803 10 of 16

+ h11 (−ln + g1 (ln , h(ln , ωn ), ωn ))ωn + h02 ωn2 + o (ρ33 )

Comparing the corresponding coefficients of terms in the above center manifold

equation, it is easy to derive that

d200 d
h20 = , h = 101 , h02 = 0
1 − λ2 11 1 − λ2

So, the system (13) restricted to the center manifold is given by

ln+1 = f (ln , ωn ) =: −ln + g1 (ln , h(ln , ωn ), ωn ) + o (ρ33 )

= −ln + c20 ln2 + c11 ln ωn + c30 ln3 + c21 ln2 ωn + c12 ln ωn2 + o (ρ33 )

Accordingly, we have the following:

f 2 (ln , ωn ) = ln − 2c11 ln ωn − 2(c220 + c30 )ln3 + (c211 − 2c12 )ln ωn2 − c11 c20 ln2 ωn + o (ρ33 )

with c20 = c200 , c11 = c101 , c30 = c300 , c21 = c100 h11 + c011 h20 + c201 , and c12 = c011 .
It is not difficult to calculate

∂f ∂f2 ∂2 f 2
f (ln , ωn )|(0,0) = 0, |(0,0) = −1, |(0,0) = 0, | =0
∂ln ∂ωn ∂ln2 (0,0)
∂2 f 2
| = −2c11 = −2c101 = −2µ(1 − M)
∂ln ∂ωn (0,0)
2(1 + β)(1 + α + β)2
=
2α(1 + β − α) − (1 + β)(1 + α + β)2
2(1 + β)(1 + α + β)2
= < 0(̸= 0)
−2α( β2 + β + 1) − (1 + β)((1 + β)2 + α2 )
∂3 f 2
|(0,0) = −12(c220 + c30 ) = −12(c2200 + c300 ) ̸= 0
∂ln3

According to (21.1.43)–(21.1.46) in [26], p. 507, all conditions are valid for a flip
bifurcation to occur; hence, the system (8) undergoes a flip bifurcation at the fixed point E2 .
The proof is complete.

Next, we provide a proof for Theorem 3.

Proof of Theorem 3. First, give a small perturbation δ∗∗ of the parameter δ around δ2 in the
system (10), i.e., δ∗∗ = δ − δ2 with 0 < |δ∗∗ | ≪ 1. Under the perturbation, the system (10) is

α(Yn +u∗ )2

1− Xn − u ∗ −
X n +1 = ( X n + u ∗ ) e β(Yn +u∗ )2 +( Xn +u∗ )2 − u∗

Yn +u∗
(14)
(δ +δ∗∗ )(1− X )
Yn+1 = (Yn + u∗ ) e 2 n +u∗ − u∗


The characteristic equation of the linearized equation of the system (14) at the origin
(0,0) is
F (λ) = λ2 − p(δ∗∗ )λ + q(δ∗∗ ) = 0 (15)
where
α (3 + β )
p(δ∗∗ ) = + 1 − δ2
1+β
α (3 + β ) α(δ2 − δ∗∗ )
q(δ∗∗ ) = (1 − δ2 − δ∗∗ ) +
1+β (1 + β )2
Mathematics 2024, 12, 1803 11 of 16

 2
Notice that p2 (0) − 4q(0) = 2 + αδ2
1+ β − δ2 − 4 < 0; so, for 0 < |δ∗∗ | ≪ 1, the two
roots of F (λ) = 0 in (15) are

p(δ∗∗ ) ± i
p
∗∗ 4q(δ∗∗ ) − p2 (δ∗∗ )
λ1,2 (δ ) =
2
The occurrence of a Neimark–Sacker bifurcation requires the following two conditions
to be satisfied [26]:
d|λ1,2 (δ∗∗ )|
 
1. dδ ∗∗ ̸= 0;
δ∗∗ =0
2. i (0) ̸ = 1, i = 1, 2, 3, 4.
λ1,2
It is easy to observe that |λ1,2 (δ∗∗ )| = q(δ∗∗ ) and (|λ1,2 (δ∗∗ )|)|δ∗∗ =0 = q(0) = 1.
p p

Therefore
 d|λ (δ∗∗ )|  α (2 + β )2
1,2
= − < 0(̸= 0)
dδ∗∗ δ∗∗ =0 (1 + β )2
i (0) ̸ = 1 for i = 1, 2, 3, 4,; so, the two conditions are satisfied.
Obviously, λ1,2
Second, in order to derive the normal form of the system (14), one expands (14) in a
power series up to the third-order term around the origin to obtain

Xn+1 = s10 Xn + s01 Yn + s20 Xn2 + s11 Xn Yn + s02 Yn2




+s30 Xn3 + s21 Xn2 Yn + s12 Xn Yn2 + s03 Yn3 + o (ρ37 )



(16)

 Yn+1 = t10 Xn + t01 Yn + t20 Xn2 + t11 Xn Yn + t02 Yn2
+t30 Xn3 + t21 Xn2 Yn + t12 Xn Yn2 + t03 Yn3 + o (ρ37 )




p
where ρ7 = Xn2 + Yn2 .

s10 = a100 , s01 = a010 , s20 = a200 , s11 = a110

s02 = a020 , s30 = a300 , s21 = a210 , s12 = a120 , s03 = a030
δ2 (δ2 − 2) δ2 (δ2 − 2)
t10 = δ2 , t01 = 1 − δ2 , t20 = , t02 =
2u∗ 2u∗
2
mδ2 (δ2 − 6δ2 + 6)
δ2 (δ2 − 2)
t11 = − , t30 =
u∗ 6( u ∗ )2
δ22 (δ2 − 3) δ2 (δ22 − 2δ2 + 2)
t03 = − , t 12 =
6( u ∗ )2 2( u ∗ )2
δ2 (δ22 − 5δ2 + 4)
b21 = −
2( u ∗ )2

Take matrix
ζ −1 1
! !
0 s01 −1 η01 η
T= , then T = 1
η 1−ζ s01 0
Make a change in variables
( X, Y )T = T ( M, N )T
Then, the system (16) is changed to the following form:

g ( M, N ) + o (ρ48 )
      
M ζ −η M
→ + 3 (17)
N η ζ N g4 ( M, N ) + o (ρ48 )
√
where ρ8 = M2 + N 2 .

g3 ( M, N ) = j20 X 2 + j11 XY + j02 Y 2 + j30 X 3 + j21 X 2 v + j12 XY 2 + j03 Y 3

Mathematics 2024, 12, 1803 12 of 16

g4 ( M, N ) = k20 X 2 + k11 XY + k02 Y 2 + k30 X 3 + k21 X 2 Y + k12 XY 2 + k03 Y 3

X = s01 N, Y = η M + (1 − ζ ) N

s20 (ζ − 1) t20 s02 (ζ − 1) t02 s ( ζ − 1) t11

j20 = + , j02 = + , j = 11 +
ηs01 η ηa01 η 11 ηs01 η
s30 (ζ − 1) t s (ζ − 1) t03 s ( ζ − 1) t12
j30 = + 30 , j03 = 03 + , j = 12 +
ηs01 η ηs01 η 12 ηa01 η
s ( ζ − 1) t s s s s
j21 = 21 + 21 , k20 = 20 , k02 = 02 , k11 = 11 , k30 = 30
ηs01 η s01 s01 s01 s01
s03 s12 s
k03 = ,k = , k21 = 21
s01 12 s01 s01

Furthermore

FXX |(0,0) = 2j02 η 3 , FXY |(0,0) = j11 s01 η + 2j02 η (1 − ζ )

FYY |(0,0) = 2j02 s201 + 2j11 s01 (1 − ζ ), FXXX |(0,0) = 6j03 η 3
FXXY |(0,0) = 2j21 s01 η 2 + 6j03 η 2 (1 − ζ )
FXYY |(0,0) = 2j21 s201 η + 4j12 s01 η (1 − ζ ) + 6j03 η (1 − ζ )2
FYYY |(0,0) = 4(1 − ζ )3 + 6j30 s301 + 4j21 s201 (1 − ζ ) + 6j12 s01 (1 − ζ )2
GXX |(0,0) = 2k02 η 3 , GXY |(0,0) = k11 s01 η + 2k02 η (1 − ζ )
GYY |(0,0) = 2k02 s201 + 2k11 s01 (1 − ζ ), GXXX |(0,0) = 6j03 η 3
GXXY |(0,0) = 2k21 s01 η 2 + 6k03 η 2 (1 − ζ )
GXYY |(0,0) = 2k21 s201 η + 4k12 s01 η (1 − ζ ) + 6k03 η (1 − ζ )2
GYYY |(0,0) = 4(1 − ζ )3 + 6k30 s301 + 4k21 s201 (1 − ζ ) + 6k12 s01 (1 − ζ )2

To determine the stability and direction of the bifurcation curve (closed orbit) for the
system (8), the discriminating quantity L should be calculated and not be zero, where
 (1 − 2λ )λ2  1
1 2
L = − Re θ20 θ11 − |θ11 |2 − |θ02 |2 + Re(λ2 θ21 ) (18)
1 − λ1 2
1
θ20 = [ FXX − FYY + 2GXY + i ( GXX − GYY − 2FXY )]|(0,0)
8
1
θ11 = [ FXX + FYY + i ( GXX + GYY )]|(0,0)
4
1
θ02 = [ FXX − FYY − 2GXY + i ( GXX − GYY + 2FXY )]|(0,0)
8
1
θ21 = [ FXXX + FXYY + GXXY + GYYY
16
+ i ( GXXX + GXYY − FXXY − FYYY )]|(0,0)

Based on [26–28], we see that if L < (>)0, then an attracting (a repelling) invariant
closed curve bifurcates from the fixed point.
The proof is then complete.
Mathematics 2024, 12, 1803 13 of 16

4. Numerical Simulation
In this section, by using the software Matlab, we obtain the bifurcation diagrams and
phase portraits of the system (8) at the fixed point E2 , which illustrate our theoretical results
previously derived and reveal some new dynamical behaviors.
First, vary δ in the range (2.7, 3) and (0.35, 0.6), respectively, and fix α = 0.8, β = 0.5
with the initial value ( x0 , y0 ) = (0.4667, 0.4667). Figure 1a shows the existence of a flip
bifurcation at the fixed point E2 = (0.4667, 0.4667) when δ = δ1 = 2 + (1+α+4α β)(1+ β)
≈ 2.93
and indicates the periodic orbits and chaos in the system (8) as δ increases. Meanwhile, we
∂2 f 2 ∂3 f 2
can calculate that c2200 + c300 ≈ −10.92 < 0 and − ∂ln ∂ωn / ∂l 3 |(0,0) > 0, which means that
n
the direction of the flip bifurcation is on the right side of the critical value. Furthermore,
according to Case 2, we can clearly see that the nature of the system (8) changes from
unstable to stable near δ2 . This change is shown in Figure 1b, and the periodic orbit is
simulated in Figure 2b. This agrees with the conclusion in Theorem 2.
Then, we choose different values of the parameter δ. The corresponding phase portraits
are plotted in Figures 3 and 4, respectively. Figure 3 implies that the closed curve is stable
inside, while Figure 4 indicates that the closed curve is stable outside. That is to say, there
occurs a stable invariant closed curve around the fixed point E2 . This agrees with the
conclusion in Theorem 3.
Finally, take initial values ( x0 , y0 ) = (0.43, 0.43) in Figure 2a and (0.4667, 0.4667) in
Figure 2b. One finds a new dynamical phenomenon—the existence of a limit cycle. This
means that the system produces periodic oscillations here.

(a) δ ∈ (2.7, 3) (b) δ ∈ (0.35, 0.6)

Figure 1. Bifurcation of the system (8) in (δ, x )-plane with α = 0.8, β = 0.5, and the initial value
( x0 , y0 ) = (0.4667, 0.4667).

(a) δ = 0.67857 (b) δ = 0.45833

Figure 2. Phase portraits of the system (8) with different parameter values: (a) α = 0.8, β = 0.4,
and δ = 0.67857; and (b) α = 0.8, β = 0.5, and δ = 0.45833. Different initial values: (a) ( x0 , y0 ) =
(0.43, 0.43) and (b) ( x0 , y0 ) = (0.4667, 0.4667).
Mathematics 2024, 12, 1803 14 of 16

(a) δ = 0.3 (b) δ = 0.35

(c) δ = 0.39 (d) δ = 0.4

Figure 3. Phase portraits for the system (8) with α = 0.8, β = 0.5, and different δ with the initial value
( x0 , y0 ) = (0.4667, 0.4667) inside the closed orbit.

(a) δ = 0.45 (b) δ = 0.49

(c) δ = 0.5 (d) δ = 0.52

Figure 4. Phase portraits for the system (8) with α = 0.8, β = 0.5, and different δ with the initial value
( x0 , y0 ) = (0.4667, 0.4667) outside the closed orbit.

5. Conclusions and Discussion

In this paper, we analyze a predator–prey model of Leslie type with ratio-dependent
simplified Holling type IV functional response. By using the semi-discretization method,
Mathematics 2024, 12, 1803 15 of 16

the system (4) is transformed to the discrete system (8). At first, one considers the existence
1+ β − α 1+ β − α
and stability of the positive fixed point E2 = ( 1+ β , 1+ β ). Subsequently, one studies
the existence conditions of the flip bifurcation and Neimark–Sacker bifurcation of the
system (8) at the fixed point E2 by using the center manifold theorem and bifurcation
theory. In the end, we confirm the correctness of the theoretical results previously derived
through numerical simulations. In the process of simulation, the existence of a limit cycle is
also found.
As for the biological significance, our results indicate that a limit cycle will occur
when the parameter δ is small. This means that the interaction between prey and predator
leads to periodic oscillations, indicating the rich dynamic properties of the system. When
appropriately adding the value of the parameter δ, the prey and predator populations will
coexist and the limit cycle will be eliminated. Our studies provide a theoretical basis for
the stable coexistence of predator and prey.
However, there still are some questions worth investigating. For example, we only know
of the existence and bifurcation of the system (3) at the fixed point E2 when 0 < α < 1 + β.
How about the case when α ≥ 1 + β? Are there more interesting dynamical properties if we
discuss the impact of seasonality on the system’s behavior? How about using discrete methods
other than the semi-discretization method that we use in this paper? We hope that interested
readers consider these questions.

Author Contributions: L.L. and X.L. contributed equally and significantly in writing this paper. All
authors have read and agreed to the published version of the manuscript.
Funding: This work is partly supported by the National Natural Science Foundation of China
(61473340) and the Distinguished Professor Foundation of Qianjiang Scholar in Zhejiang Province
(F708108P02).
Data Availability Statement: There are no applicable data associated with this manuscript.
Conflicts of Interest: The authors declare that they have no competing interests.

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