1. Introduction
The study of discrete time models defined by difference equations has become increasingly popular in biological mathematics research, particularly for populations with non-overlapping generations [
1,
2,
3,
4,
5,
6,
7,
8,
9]. Among the various mathematical models used in population dynamics, the Moran–Ricker model is considered one of the most important for characterizing density dependence in single-species population dynamics. The Moran–Ricker model is often employed to analyze the population dynamics of various species, such as the red king crab off Kodiak Island [
10]. In previous works [
10,
11], the Moran–Ricker model with delay was explored in depth to study its dynamical modes, including the appearance and stability criteria for the two- and three-year cycles. More recently, the Moran–Ricker equation has been used to explain local population models with a fundamental age structure and density-dependent regulation, leading to the discovery of various dynamic modes (stable, periodic, and chaotic) that can coexist under the same conditions [
11,
12,
13,
14,
15]. Interestingly, these models suggest that a change in the observed dynamic mode could result from a random variation in the current population size. In addition to the red king crab, the Moran–Ricker model was also applied to investigating the dynamics of the Larch bud moth [
12,
13,
14,
15,
16], revealing its chaotic dynamics behavior with time lag. In this study, we use the inner product approach to compute the critical normal form of two types of one-parameter bifurcations and the generating cases, including the 1:2, 1:3, and 1:4 resonances at fixed points of the model. This work provides a deeper understanding of the dynamics of the Moran–Ricker model, and sheds light on the various dynamic modes that can be observed under different conditions.
Time delay is an important feature in ecological models because it can affect the dynamics of populations and communities over time. In many ecological systems, there are time lags between the occurrence of events and their effects on other variables, such as the response of a population to changes in environmental conditions or the feedback between predator and prey populations. Time delays can arise due to a variety of factors, such as biological processes (e.g., growth and reproduction rates), physical transport processes (e.g., the dispersal of organisms or nutrients), and behavioral interactions (e.g., predator avoidance or prey switching).
The aim of this article is to investigate all of the codim-1 bifurcations of the Moran-Ricker model on the basis of such a time lag as transcritical, period-doubling, and Neimark–Sacker bifurcations, considering different values of model parameters and codim-2 bifurcations, such as resonance 1:2, 1:3, and 1:4, assuming the combination of two parameters. The non-degeneracy of codim-1 and codim-2 bifurcations for the delayed Moran-Ricker model is verified using normal form coefficients in this work. This approach has the advantage of avoiding the direct computation of the central manifold and the conversion of the linear component to the Jordan form, and the complexity of the computations is substantially lower than that of the way of computing the central manifold and converting the linear part into the Jordan form. More information can be found in [
17,
18,
19,
20]. The rest of this paper is organized as follows.
Section 2 presents the necessary conditions for flip bifurcation and Neimark–Sacker bifurcation at the fixed points of a discrete-time Moran-Ricker model with delay. In
Section 2, the two parameters in the model were chosen as free parameters to investigate the local dynamics generated by 1:2, 1:3, and 1:4 resonance, and we provide the required conditions for 1:2, 1:3, and 1:4 resonance at the model’s fixed points.
Section 3 provides numerical analysis to demonstrate the theoretical results, and complicated dynamics are shown. Lastly, in
Section 4, the conclusions are outlined.
2. Moran-Ricker Model Based on a Time Delay
In Nedorezov [
11], the author proposed the following Moran–Ricker model based on a time delay:
To eliminate the delay, we could convert the original one-dimensional delay Model (
1) into a two-dimensional model as follows:
Model (
2) are represented with the map depicted below:
To obtain the fixed points of Model (
3), we solve the following equations:
The solutions of (
4) are:
The Jacobian matrix of Map (
3) is as follows:
Map
can be written as follows:
where
and
2.1. Codimension 1 Bifurcation of
Theorem 1. undergoes a transcritical (pitchfork) bifurcation at ,
Proof. It is clear that
has eigenvalues
and
. This has a simple eigenvalue
on the provided unit circle
. The central manifold corresponding to
is one-dimensional and can be considered as follows:
where
and
The restriction of
to (
7) has the following form:
where
The dynamics of the Moran–Ricker model at the fixed point
depend on the
parameter. If this parameter is non-zero, the fixed point undergoes a transcritical bifurcation. Otherwise, a pitchfork bifurcation occurs. Notably,
remains a fixed point and is stable throughout the bifurcation. For a more comprehensive explanation, we refer the reader to [
21,
22]. □
Remark 1. In discrete dynamical systems, a fold bifurcation occurs when the Jacobian matrix evaluated at a fixed point has only a simple eigenvalue of and no other eigenvalues on the unit circle. In this case, the central manifold of the system corresponding to the map has dimension 1, and the map restricted to this manifold has the following form:where is the critical normal form coefficient of the fold bifurcation determined by non-degeneracy and the scenario of the bifurcation. 2.2. Codimension 1 Bifurcation of
Theorem 2. undergoes a flip bifurcation at .
Proof. It is clear that
has eigenvalues
and
. It has a simple eigenvalue
on the provided unit circle
. The central manifold corresponding to
is one-dimensional and can be considered as follows:
where
and
The restriction of
to (
7) has the following form:
where
The bifurcation is generic, provided
. If
(
) that the bifurcation is supercritical (subcritical); for more details, see [
21,
22]. □
Remark 2. In discrete dynamical systems, a flip bifurcation occurs when the Jacobian matrix evaluated at a fixed point has only a simple eigenvalue of and no other eigenvalues on the unit circle. In this case, the central manifold of the system corresponding to the map has dimension 1, and the map restricted to this manifold has the following form:where is the critical normal form coefficient of the flip bifurcation determined by non-degeneracy and the scenario of the bifurcation. This remark provides a brief summary of the key characteristics of flip bifurcation in discrete dynamical systems, including the conditions for its occurrence, the dimensionality of the central manifold, and the form of the map restricted to this manifold. It also introduces the concept of the critical normal form coefficient, which is an important parameter for characterizing the behavior of the system near the bifurcation point.
Theorem 3. undergoes a Neimark–Sacker bifurcation at , provided that .
Proof. Clearly,
has eigenvalues
that
. As
, the pair of complex conjugate
lies on the unit circle. This implies that the condition for Neimark–Sacker bifurcation is fulfilled.
The central manifold corresponding to
is two-dimensional and can be considered as follows:
where
and
The restriction of
to (
8) has the form
where
with
The bifurcation is generic, provided
, where
If
(
) the bifurcation is subcritical (supercritical); for more details, see [
23]. □
Remark 3. In discrete dynamical systems, a Neimark–Sacker bifurcation occurs when the Jacobian matrix evaluated at a fixed point has a simple complex conjugate pair of eigenvalues on the unit circle and no other eigenvalues on the unit circle. In this case, the center manifold of the system corresponding to the map has dimension 2, and the map restricted to this manifold has the form:where is the critical normal form coefficient of the Neimark–Sacker bifurcation. The value of determines the non-degeneracy and the scenario of the bifurcation. This remark provides a concise description of the key features of the Neimark–Sacker bifurcation in discrete dynamical systems, including the conditions for its occurrence, the dimensionality of the central manifold, and the form of the map restricted to this manifold. It also introduces the concept of the first Lyapunov coefficient and the normal form coefficient, which are important parameters for characterizing the behavior of the system near the bifurcation point.
2.3. Codimension 2 Bifurcation of
Theorem 4. undergoes a 1:2 resonance bifurcation at Proof. Clearly,
has eigenvalues
on the unit circle.
The central manifold corresponding to
is two-dimensional and can be considered as follows:
where
and
The restriction of
to (
9) has the following form:
where
The bifurcation is generic because of and . □
Remark 4. In discrete dynamical systems, a 1:2 resonance bifurcation occurs when the Jacobian matrix evaluated at a fixed point has two eigenvalues of −1 on the unit circle and no other eigenvalues on the unit circle. In this case, the central manifold of the system corresponding to the map has dimension 2, and the map restricted to this manifold has the following form: Here, and are the critical normal form coefficients of the 1:2 resonance bifurcation that determine the non-degeneracy and the scenario of the bifurcation.
Theorem 5. undergoes a 1:3 resonance bifurcation at Proof. Clearly,
has eigenvalues
on the unit circle.
The central manifold corresponding to
is two-dimensional and can be considered as follows:
where
and
The restriction of
to (
10) has the following form:
where
The bifurcation is generic because of . □
Remark 5. In discrete dynamical systems, a 1:3 resonance bifurcation occurs when the Jacobian matrix evaluated at a fixed point has two eigenvalues of on the unit circle and no other eigenvalues on the unit circle. In this case, the central manifold of the system corresponding to the map has dimension 2, and the map restricted to this manifold has the following form: Here, and are the critical normal form coefficients of the 1:3 resonance bifurcation that determine the non-degeneracy and the scenario of the bifurcation:
Theorem 6. undergoes a 1:4 resonance bifurcation at Proof. Clearly,
has eigenvalues
on the unit circle.
The central manifold corresponding to
is two-dimensional and can be considered as follows:
where
and
The restriction of
to (
11) has the form
where
The bifurcation scenario is determined by
Since there are two limit-point curves of the fourth iteration of . □
Remark 6. In discrete dynamical systems, a 1:4 resonance bifurcation occurs when the Jacobian matrix evaluated at a fixed point has two eigenvalues of on the unit circle, and no other eigenvalues on the unit circle. In this case, the central manifold of the system corresponding to the map has dimension 2, and the map restricted to this manifold has the following form: Here, and are the critical normal form coefficients of the 1:4 resonance bifurcation that determine the non-degeneracy and the scenario of the bifurcation:
3. Numerical Bifurcation Analysis of
MatContM is a version of MatCont that is specifically designed for the continuation and bifurcation analysis of maps or discrete dynamical systems. It extends the capabilities of MatCont to handle systems that are defined by iterated maps rather than differential equations.
Like MatCont, MatContM is an open-source software package that can be used to analyze a wide range of nonlinear systems. It provides a user-friendly interface for specifying the map, selecting the continuation algorithm, and visualizing the results.
MatContM implements several continuation algorithms that are tailored to the needs of discrete dynamical systems. These include the parameter-homotopy, pseudo-arclength, and tangent-bifurcation methods. It also provides tools for detecting and analyzing various types of bifurcations, including period-doubling, Neimark–Sacker, and saddle-node bifurcations.
One of the strengths of MatContM is its ability to handle systems with multiple parameters and discontinuities. It can also handle systems with higher-dimensional maps, such as those arising in spatially extended systems.
In summary, MatContM is a powerful tool for the continuation and bifurcation analysis of maps or discrete dynamical systems. It is a valuable addition to a toolkit if one is working with discrete dynamical systems, and wants to provide a rigorous and quantitative analysis of their behavior. To confirm the analytical results, we used
MatcontM, which is a toolbox of
Matlab that works on the basis of the numerical continuation method; for more details, see [
23,
24]. Here,
and
are considered a free parameter and a fixed parameter, respectively.
We consider two cases for fixed parameter .
- (i)
Case 1: If
, we consider
. With this assumption, a flip bifurcation occurs for
at
with critical coefficient
. Because this coefficient is positive, we conclude that the flip bifurcation is supercritical, and the double cycle bifurcated from
is stable. The period doubles when a curve emanates from a PD bifurcation is shown in
Figure 1a. A 1:2 resonance bifurcation occurs on the curve of flip bifurcation for
and
at
, see
Figure 1b.
- (ii)
Case 2: If
, we consider
. With this assumption, a Neimark–Sacker bifurcation occurs for
at
with the first Lyapunov coefficient
. Since
, we conclude that the Neimark–Sacker bifurcation is supercritical. This phenomenon is shown in
Figure 2.
On the curve, Neimark–Sacker bifurcation caused a 1:4 resonance fifurcation for
at
and a 1:3 resonance fifurcation for
at
, see
Figure 3a. The neutral saddle curve of the third iterate
is presented in
Figure 3b.
In the following, the structure of periodic orbits and transitions between them is considered. The 1:4 resonance was set as the initial point, and the corresponding period was computed on a mesh of
parameters (alpha, sigma). Different colors are associated with different periods of the trajectories. It is clear that the stable regions were becoming increasingly smaller as the parameters increased. Almost periodic regions coexist with period-doubling regions. For example, the period-18 and -9 regions. Meanwhile, when compared with
Figure 1b and
Figure 3a, the Neimark–Sacker bifurcation curve and period-doubling bifurcation curve were also consistent with the border between different types of bifurcations in
Figure 4. It is interesting that there existed a series of a period-adding and bar-shaped region with a period higher than four.
4. Conclusions
In conclusion, this study highlights the importance of considering time delay factors in ecological models, particularly for local population dynamics. The findings demonstrated that the incorporation of delay factors can produce complex and diverse dynamic modes, including equilibrium, periodic oscillations, and chaotic fluctuations. Moreover, the Moran–Ricker model with time lag effectively described the dynamics of several species, indicating the relevance of delay models in understanding the population dynamics of real ecosystems. The study also emphasizes the potential for shifts in dynamic modes due to changes in population size or external factors, which has significant implications for the management and conservation of natural populations. Overall, the findings of this study have important implications for ecological research, and highlight the need for the continued development of modeling techniques that can capture the complexity of real ecosystems.
In addition to providing insights into the dynamics of the Moran–Ricker model with time delay, our study has important implications for understanding and managing real-world populations. For example, the model has been used to describe the dynamics of several species, including Zeiraphera griseana and Epinotia tedella. Our findings highlight the significance of estimating population parameters, and the potential for shifts in dynamic modes due to changes in population size or external factors. Understanding these shifts and their underlying mechanisms is crucial for predicting the responses of populations to environmental change, such as global climate change or habitat loss.