Analysis of The Rosenzweig-MacArthur Predator-Prey Model with Anti-Predator Behavior CAUCHY –Jurnal Matematika Murni dan Aplikasi Volume 6(4) (2021), Pages 260-269 p-ISSN: 2086-0382; e-ISSN: 2477-3344 Submitted: January 22, 2021 Reviewed: March 17, 2021 Accepted: April 14, 2021 DOI: http://dx.doi.org/10.18860/ca.v6i4.11472 Analysis of The Rosenzweig-MacArthur Predator-Prey Model with Anti-Predator Behavior Ismail Djakaria1, Muhammad Bachtiar Gaib2 , Resmawan3 1,2,3Department of Mathematics, Universitas Negeri Gorontalo, Indonesia Email: iskar@ung.ac.id, m.tiargaib@gmail.com, resmawan@ung.ac.id ABSTRACT This paper discusses the analysis of the Rosenzweig-MacArthur predator-prey model with anti- predator behavior. The analysis is started by determining the equilibrium points, existence, and conditions of the stability. Identifying the type of Hopf bifurcation by using the divergence criterion. It has shown that the model has three equilibrium points, i.e., the extinction of population equilibrium point (𝐸0), the non-predatory equilibrium point (𝐸1), and the co-existence equilibrium point (𝐸2). The existence and stability of each equilibrium point can be shown by satisfying several conditions of parameters. The divergence criterion indicates the existence of the supercritical Hopf-bifurcation around the equilibrium point 𝐸2. Finally, our model's dynamics population is confirmed by our numerical simulations by using the 4th-order Runge-Kutta methods. Keywords: Rosenzweig-MacArthur; predator-prey model; anti-predator behaviour; Hopf Bifurcation; divergence criterion; equilibrium point. INTRODUCTION Population dynamics are the most interesting research in mathematical biology which discusses the interactions that occur between prey and predator in a particular ecosystem [1]. This interaction has implemented to a simple mathematical model known as the Lotka-Volterra predator-prey model [2]. In a mathematical model, the predation process (interaction between prey and predator) is expressed in some form that is known as a functional response. This functional response has classified three functions, i.e. Holling-Type I, Holling-Type II, and Holling-Type III where each type determine the characteristic of the predator [3]. On the progress, Rosenzweig and MacArthur modifying the Lotka-Volterra predator-prey model with the assumption the attack rate of predator increases at a decreasing rate with prey density until it becomes constant due to satiation which is affected by Holling-Type II functional response [4]. Further, some modified of Lotka-Volterra predator-prey model by considering the infectious disease [5]-[7]. Several research has discussed the modification of the Rosenzweig-MacArthur predator-prey model [8][9] is introduced predator foraging facilitation into Holling-Type II functional response. Furthermore, the Rosenzweig-MacArthur model has modified with various factors, e.g. the stage-structure [10][11], the refuge effect [12][13], the harvesting to one or more population [14][15]. From several studies described above, no one http://dx.doi.org/10.18860/ca.v6i4.11472 mailto:iskar@ung.ac.id mailto:m.tiargaib@gmail.com mailto:resmawan@ung.ac.id Analysis of The Rosenzweig-MacArthur Predator-Prey Model with Anti-Predator Behaviour Ismail Djakaria 261 (1) considering anti-predator behavior factors. In this article, the Rosenzweig-MacArthur predator-prey model by [6] modified considering anti-predator behavior factors [16]. These factors can be considered in the model because the dynamics of the model will be very complex when the prey population prefers to defending and provide resistance when the predation process is occurring. The structure of this paper is as follows. In the next section, the methods in our work are described. Then, the analysis of the model has been discussed. Finally, a brief conclusion of our work is given. METHODS The dynamics of the model is analyzed by carrying out the following steps: 1. Modifying the Rosenzweig-MacArthur predator-prey model considering anti- predator behavior factors. 2. Simplifying the model by using non-dimensional to reduce the number of parameters and solving the equilibrium points of the model. 3. Identifying the existence, local stability, and global stability of the equilibrium points. 4. Identifying the Hopf-bifurcation type by using the divergence criterion. 5. Demonstrated the numerical simulations of the model to describe the analysis results by using the 4th-order Runge-Kutta method. RESULTS AND DISCUSSION Mathematical Model In this article, the mathematical model is formulated based on the following assumptions: 1. The prey population is assumed to grow logistically with an intrinsic growth rate of π‘Ÿ and carrying capacity of the environment of 𝐾 and reduced due to the predation process. 2. The predator population is assumed to grow due to the predation process. 𝑐 is the conversion rate of the consumed prey into predator births. 3. The predation process follows Holling-Type II functional response which is affected by the encounter rate function where there is foraging facilitation of predator (𝑀 = 0), π‘Ž is the saturated rate of the predator, 𝑏 is coefficient interaction on both population and β„Ž is the predator time handling. 4. π‘š is the mortality of predators. 5. πœ‚ is the anti-predator behavior. From the following assumptions above, the dynamics of the model can be represented by the following set of differential equations: 𝑑π‘₯ 𝑑𝑑 = π‘Ÿπ‘₯(1βˆ’ π‘₯ 𝐾 )βˆ’ (π‘Ž βˆ’π‘)π‘₯𝑦 𝑦 +β„Ž(π‘Ž βˆ’π‘)π‘₯ 𝑑𝑦 𝑑𝑑 = 𝑐(π‘Ž βˆ’π‘)π‘₯𝑦 𝑦 +β„Ž(π‘Ž βˆ’π‘)π‘₯ βˆ’π‘šπ‘¦ βˆ’πœ‚π‘₯𝑦 Where π‘₯ and 𝑦 are respectively the densities of prey and predator population at time 𝑑 and π‘₯(0),𝑦(0) > 0. Analysis of The Rosenzweig-MacArthur Predator-Prey Model with Anti-Predator Behaviour Ismail Djakaria 262 (2) (3) (4) To simplify our analysis, we reduce the number of parameters in system (1) by using the following parameter scales [17]: π‘₯ β†’ π‘₯𝐾, 𝑦 β†’ 𝑦(π‘Ž βˆ’π‘)πΎβ„Ž, 𝑑 β†’ 𝑑 π‘Ÿ We obtain the following non-dimensional model 𝑑π‘₯ 𝑑𝑑 = π‘₯(1βˆ’π‘₯)βˆ’ 𝛼π‘₯𝑦 π‘₯ +𝑦 𝑑𝑦 𝑑𝑑 = 𝛽π‘₯𝑦 π‘₯ +𝑦 βˆ’π›Ύπ‘¦ βˆ’π›Ώπ‘₯𝑦 where 𝛼 = (π‘Ž βˆ’π‘) π‘Ÿ , 𝛽 = 𝑐 β„Žπ‘Ÿ , 𝛾 = π‘š π‘Ÿ , 𝛿 = πœ‚πΎ π‘Ÿ Existence and Stability Analysis of Equilibrium Points In this section, the equilibrium point of model (2) is obtained by solving [18]: π‘₯(1βˆ’π‘₯)βˆ’ 𝛼π‘₯𝑦 π‘₯ +𝑦 = 0 𝛽π‘₯𝑦 π‘₯ +𝑦 βˆ’π›Ύπ‘¦ βˆ’π›Ώπ‘₯𝑦 = 0 Thus, from the system (3), we obtain the following equilibrium points, i.e.: 1. A trivial equilibrium point 𝐸0 = (0,0), always exists. 2. A non-predator equilibrium point 𝐸1 = (1,0), always exists too. 3. A co-existence equilibrium point 𝐸2 = (π‘₯ βˆ—,π‘¦βˆ—), where π‘₯βˆ— = 𝛽 βˆ’π›Όπ›½ +𝛼𝛾 𝛽 βˆ’π›Όπ›Ώ , π‘¦βˆ— = (𝛽 βˆ’π›Όπ›½ +𝛼𝛾)(𝛽 βˆ’π›Ύ βˆ’π›Ώ) (𝛽 βˆ’π›Όπ›Ώ)(𝛾 +𝛿 βˆ’π›Όπ›Ώ) which exists if 𝛽 > 𝛼(𝛽 βˆ’π›Ύ), 𝛾 +𝛿 < 𝛼𝛿 < 𝛽 Now, study the local stability of the dynamics of the system (3) around each of equilibrium point. The Jacobian matrix from the system (3) is determined as [19]: 𝐽(π‘₯,𝑦) = ( 1βˆ’2π‘₯ βˆ’ 𝛼𝑦 π‘₯ +𝑦 + 𝛼π‘₯𝑦 (π‘₯ +𝑦)2 βˆ’ 𝛼π‘₯ π‘₯ +𝑦 + 𝛼π‘₯𝑦 (π‘₯ +𝑦)2 𝛽𝑦 π‘₯ +𝑦 βˆ’ 𝛽π‘₯𝑦 (π‘₯ +𝑦)2 βˆ’π›Ώπ‘¦ 𝛽π‘₯ π‘₯ +𝑦 βˆ’ 𝛽π‘₯𝑦 (π‘₯ +𝑦)2 βˆ’π›Ύ βˆ’π›Ώπ‘₯ ) By evaluating this Jacobian matrix (4) at each equilibrium point, we obtain the local stability properties of 𝐸0, 𝐸1, and 𝐸2 as follows. Analysis of The Rosenzweig-MacArthur Predator-Prey Model with Anti-Predator Behaviour Ismail Djakaria 263 Theorem 1. The trivial equilibrium point 𝐸0 always unstable (saddle). Proof: The Jacobian matrix (4) evaluated in equilibrium point 𝐸0 is given by 𝐽(𝐸0) = ( 1 0 0 βˆ’π›Ύ ) So, by solving the characteristic equation, we obtained the eigenvalues of 𝐽(𝐸0) is πœ†1 = 1 and πœ†2 = βˆ’π›Ύ. It means πœ†1 > 0 and πœ†2 < 0. Therefore, stability of equilibrium point 𝐸0 is unstable (saddle).∎ Theorem 2. If 𝛿 > 𝛽 βˆ’π›Ύ, then the non-predatory equilibrium point 𝐸1 of system (2) is locally asymptotically stable. Proof: The Jacobian matrix (4) evaluated in equilibrium point 𝐸1 is given by 𝐽(𝐸1) = ( βˆ’1 βˆ’π›Ό 0 𝛽 βˆ’π›Ύ βˆ’π›Ώ ) So, by solving the characteristic equation, we obtained the eigenvalues of 𝐽(𝐸1) is πœ†1 = βˆ’1 and πœ†2 = 𝛽 βˆ’π›Ύ βˆ’π›Ώ. It means πœ†1 < 0. Therefore, if 𝛿 > 𝛽 βˆ’π›Ύ then each the eigenvalues of 𝐽(𝐸1) are negatif, and 𝐸1 is locally asymptotically stable.∎ Theorem 3. The co-existence equilibrium point 𝐸2 is locally asymptotically stable if the conditions below are satisfied 𝛿2 < Θ+Ξ₯ Ξ– Proof: The Jacobian matrix (4) evaluated in equilibrium point 𝐸1 is given by 𝐽(𝐸2) = ( 𝑀11 𝑀12 𝑀21 𝑀22 ) Where 𝑀11 = βˆ’π›½2 +𝛼𝛽2 βˆ’π›Όπ›Ύ2 βˆ’2𝛼𝛿(𝛼 βˆ’1)(𝛽 βˆ’π›Ύ)βˆ’π›Όπ›Ώ2 +𝛼2𝛿2 (𝛽 βˆ’π›Όπ›Ώ)2 𝑀12 = βˆ’ 𝛼(𝛾 +𝛿 βˆ’π›Όπ›Ώ)2 (𝛽 βˆ’π›Όπ›Ώ)2 𝑀21 = (𝛽 βˆ’π›Ύ βˆ’π›Ώ)(𝛽2𝛾 +𝛼2𝛾𝛿2 βˆ’π›½(𝛾2 +2𝛾𝛿 +𝛿2(𝛼 βˆ’1)2)) (𝛽 βˆ’π›Όπ›Ώ)2 𝑀22 = βˆ’ 𝛽(𝛽 βˆ’π›Ύ βˆ’π›Ώ)(𝛾 +𝛿 βˆ’π›Όπ›Ώ) (𝛽 βˆ’π›Όπ›Ώ)2 By solving the characteristic equation, we obtained the eigenvalues of 𝐽(𝐸2) is πœ†1,2 = 1 2 . 1 (𝛽 βˆ’π›Όπ›Ώ)2 (𝐴±𝐡) Analysis of The Rosenzweig-MacArthur Predator-Prey Model with Anti-Predator Behaviour Ismail Djakaria 264 Where 𝐴 = Ζ𝛿2 βˆ’Ξ˜βˆ’Ξ₯ and 𝐡 = Ξ¨2 βˆ’π›ΌΞ© With Ξ– = (𝛼2 βˆ’π›Ό +𝛽 βˆ’π›Όπ›½) Θ = 𝛿(𝛽(𝛽 βˆ’2𝛾)+2𝛼2(𝛽 βˆ’π›Ύ)βˆ’π›Ό(𝛽2 +2(𝛽 βˆ’π›Ύ)βˆ’π›½π›Ύ)) Ξ₯ = 𝛽2(𝛾 βˆ’π›Ό +1)+𝛾2(𝛼 +𝛽) Ξ¨ = (𝛽2 βˆ’π›Όπ›½2 +𝛼𝛾2 +2𝛼𝛿(𝛼 βˆ’1)(𝛽 βˆ’π›Ύ)βˆ’π›Όπ›Ώ2(𝛼 βˆ’1)βˆ’π›½(𝛽 βˆ’π›Ύ βˆ’π›Ώ)(𝛾 +𝛿 βˆ’π›Όπ›Ώ)) Ξ© = 4(𝛽 βˆ’π›Ύ βˆ’π›Ώ)(𝛾 +𝛿 βˆ’π›Όπ›Ώ)(𝛽2𝛾 +𝛼2𝛾𝛿2 βˆ’π›½((𝛼 βˆ’1)2𝛿2 +𝛾2 +2𝛾𝛿)) According to (), the stability of equilibrium point 𝐸2 depending on the value of 𝐴. If 𝐴 < 0, we obtained: Ζ𝛿2 βˆ’Ξ˜π›Ώ βˆ’Ξ₯ < 0 Ζ𝛿2 < Ξ˜π›Ώ +Ξ₯ 𝛿2 < Θ+Ξ₯ Ξ– By the conditions above, the stability of equilibrium point 𝐸2 is locally asymptotically stable.∎ Next, study the global stability of the dynamics of the system (3) around equilibrium point 𝐸2. We obtain the global stability properties of 𝐸2 by using the Lyapunov function [20] as follows. Theorem 4. The co-existence equilibrium 𝐸2 is globally asymptotically stable if the conditions below are satisfied: π‘₯βˆ— < (𝛼 βˆ’π›½ +𝛾 +𝛿)(𝛾 +𝛿 βˆ’π›Όπ›Ώ) 𝛼(𝛾 +𝛿 βˆ’π›Όπ›Ώ)βˆ’(𝛽 βˆ’π›Ύ βˆ’π›Ώ)2 Proof: Define a Lyapunov function as follows 𝑉(π‘₯,𝑦) = [π‘₯ βˆ’π‘₯βˆ— βˆ’π‘₯βˆ— ln( π‘₯ π‘₯βˆ— )]+[𝑦 βˆ’π‘¦βˆ— βˆ’π‘¦βˆ— ln( 𝑦 π‘¦βˆ— )] By using the function οΏ½Μ‡οΏ½ < 0,βˆ€ (π‘₯,𝑦) ∈ ℝ2 +, we obtain: πœ•π‘‰ πœ•π‘₯ . πœ•π‘₯ πœ•π‘‘ + πœ•π‘‰ πœ•π‘¦ . πœ•π‘¦ πœ•π‘‘ ≀ 0 (1βˆ’ π‘₯βˆ— π‘₯ )(π‘₯(1βˆ’π‘₯)βˆ’ 𝛼π‘₯𝑦 π‘₯ +𝑦 )+(1βˆ’ π‘¦βˆ— 𝑦 )( 𝛽π‘₯𝑦 π‘₯ +𝑦 βˆ’π›Ύπ‘¦ βˆ’π›Ώπ‘₯𝑦) ≀ 0 ( (1βˆ’π‘₯)(π‘₯ +𝑦)βˆ’π›Όπ‘¦ π‘₯ +𝑦 )(π‘₯ βˆ’π‘₯βˆ—)+( 𝛽π‘₯ βˆ’π›Ύ(π‘₯ +𝑦)βˆ’π›Ώπ‘₯(π‘₯ +𝑦) π‘₯ +𝑦 )(𝑦 βˆ’π‘¦βˆ—) ≀ 0 For (π‘₯,𝑦) ∈ ℝ2 +, we obtain: Analysis of The Rosenzweig-MacArthur Predator-Prey Model with Anti-Predator Behaviour Ismail Djakaria 265 (5) βˆ’π›Ό +𝛼π‘₯βˆ— +𝛽 βˆ’π›Ύ βˆ’π›Ώ βˆ’(𝛽 βˆ’π›Ύ βˆ’π›Ώ)π‘¦βˆ— < 0 βˆ’π›Ό +𝛼π‘₯βˆ— +𝛽 βˆ’π›Ύ βˆ’π›Ώ βˆ’π‘₯βˆ— (𝛽 βˆ’π›Ύ βˆ’π›Ώ)2 (𝛾 +𝛿 βˆ’π›Όπ›Ώ) < 0 π‘₯βˆ— ( 𝛼(𝛾 +𝛿 βˆ’π›Όπ›Ώ)βˆ’((𝛽 βˆ’π›Ύ βˆ’π›Ώ)2) (𝛾 +𝛿 βˆ’π›Όπ›Ώ) ) < 𝛼 βˆ’π›½ +𝛾 +𝛿 π‘₯βˆ— < (𝛾 +𝛿 βˆ’π›Όπ›Ώ)(𝛼 βˆ’π›½ +𝛾 +𝛿) 𝛼(𝛾 +𝛿 βˆ’π›Όπ›Ώ)βˆ’((𝛽 βˆ’π›Ύ βˆ’π›Ώ)2) By the conditions above, the stability of equilibrium point 𝐸2 is globally asymptotically stable.∎ Analysis of Hopf Bifurcation Type In this section, we’ll define the Hopf-bifurcation type by using the divergence criterion [21]. System (3) underwent a Hopf-bifurcation when it satisfies the following conditions: 𝛿2 < Θ+Ξ₯ Ξ– and 𝛼 > Ξ¨2 Ξ© To determine the Hopf-bifurcation type of system (3) on equilibrium point 𝐸2, then we formed a new system. Let πœ™(π‘₯,𝑦) is a divergence of (π‘Žπ‘“,π‘Žπ‘”). We obtain the coefficient value of π‘Ž(π‘₯,𝑦) of the system (3) when the parameter value 𝛼 = 2, 𝛽 = 0.79, 𝛾 = 0.5, and 𝛿 = 0.0186 with equilibrium point 𝐸2 βˆ— = (0.279;0.157) as follows: π‘Ž(π‘₯,𝑦) = 1+6.956π‘₯ +13,386𝑦 βˆ’6.77π‘₯2 +32.968π‘₯𝑦 +55.507𝑦2 So that a new system is obtained: 𝑧(π‘₯,𝑦) = (1+6.956π‘₯ +13,386𝑦 βˆ’6.77π‘₯2 +32.968π‘₯𝑦 +55.507𝑦2) (π‘₯(1βˆ’π‘₯)βˆ’ 𝛼π‘₯𝑦 π‘₯ +𝑦 ) 𝑀(π‘₯,𝑦) = (1+6.956π‘₯ +13,386𝑦 βˆ’6.77π‘₯2 +32.968π‘₯𝑦 +55.507𝑦2) ( 𝛽π‘₯𝑦 π‘₯ +𝑦 βˆ’π›Ύπ‘¦ βˆ’π›Ώπ‘₯𝑦) By linearizing system (4), we obtained: 𝐽(𝐸2 βˆ—) = ( 1.337 βˆ’6.002 0.732 βˆ’1.337 ) By solving the characteristic equation, we obtained the eigenvalues of 𝐽(𝐸2 βˆ—) is πœ†1,2 = Β±1.615𝑖 For a system (5) to obtain the eigenvalues of conjugate complex numbers, then we can analyze the Hopf-bifurcation of system (3) type by looking at the divergence value of system (3). We obtained: πœ™π‘₯π‘₯(𝐸2 βˆ—) = βˆ’21.109 Analysis of The Rosenzweig-MacArthur Predator-Prey Model with Anti-Predator Behaviour Ismail Djakaria 266 Based on the divergence value above, a stable limit cycle appears in the system (3). Therefore, system (3) underwent a Supercritical Hopf-bifurcation. Numerical Simulations In this section, the numerical simulation is solved using the 4th-order Runge-Kutta method [22] with initial conditions and some values of the parameters. We choose the following set of parameter values: 𝛼 = 2, 𝛽 = 0.79, 𝛾 = 0.5 With different parameter control values as follows 𝛿1 = 0.011, 𝛿2 = 0.0186 and 𝛿3 = 0.026. We using the initial condition is π‘₯(0) = 0.3 and 𝑦(0) = 0.3. (a) (b) Figure 1. (a) Phase Portrait of Case 1 and (b) Time-Series Portrait (a) (b) Figure 2. (a) Phase Portrait of Case 2 and (b) Time-Series Portrait In case 1, we obtained the dynamics of the solution on the system (3) with parameter control values 𝛿1 = 0.011. Based on figure 1(a), the trivial equilibrium point 𝐸0 = (0,0) is unstable (saddle) with eigenvalues πœ†1 = 1 and πœ†2 = βˆ’0.5. This coincides with Theorem 1. The non-predator equilibrium point 𝐸1 = (1,0) is unstable (saddle) with eigenvalues πœ†1 = βˆ’1 and πœ†2 = 0.279. This coincides with Theorem 2 on condition 𝛿 < 𝛽 βˆ’π›Ύ. The co-existence equilibrium point 𝐸2 = (0.273;0.156) is unstable (spiral) with eigenvalues πœ†1,2 = 0.003Β±0.220𝑖. This Analysis of The Rosenzweig-MacArthur Predator-Prey Model with Anti-Predator Behaviour Ismail Djakaria 267 coincides with Theorem 3 on condition 𝛿2 < Θ+Ξ₯ Ξ– . Based on figure 1(b), the prey population and predator population have increased and decreased of total populations. The case continuously oscillates with a greater deviation value. Hence, both population is unstable to a specific point. (a) (b) Figure 3. (a) Phase Portrait of Case 3 and (b) Time-Series Portrait In case 2, we obtained the dynamics of the solution on the system (3) with parameter control values 𝛿1 = 0.0186. Based on figure 2(a), the trivial equilibrium point 𝐸0 = (0,0) is unstable (saddle) with eigenvalues πœ†1 = 1 and πœ†2 = βˆ’0.5. This coincides with Theorem 1. The non-predator equilibrium point 𝐸1 = (1,0) is unstable (saddle) with eigenvalues πœ†1 = βˆ’1 and πœ†2 = 0.271. This coincides with Theorem 2 on condition 𝛿 < 𝛽 βˆ’π›Ύ. The co- existence equilibrium point 𝐸2 = (0.279;0.157) is center (spiral) with eigenvalues πœ†1,2 = Β±0.220𝑖. This coincides with Theorem 3 on condition 𝛿2 = Θ+Ξ₯ Ξ– . Based on figure 2(b), the oscillations that occur have a smaller deviation value. This condition explains that there is a stability transition from unstable to stable to a specific point. This stability transition has led to the appearance of Hopf-bifurcation. In case 3, we obtained the dynamics of the solution on the system (3) with parameter control values 𝛿1 = 0.026. Based on figure 3(a), the trivial equilibrium point 𝐸0 = (0,0) is unstable (saddle) with eigenvalues πœ†1 = 1 and πœ†2 = βˆ’0.5. This coincides with Theorem 1. The non-predator equilibrium point 𝐸1 = (1,0) is unstable (saddle) with eigenvalues πœ†1 = βˆ’1 and πœ†2 = 0.264. This coincides with Theorem 2 on condition 𝛿 < 𝛽 βˆ’π›Ύ. The co- existence equilibrium point 𝐸2 = (0.285;0.159) is stable (spiral) with eigenvalues πœ†1,2 = βˆ’0.003Β±0.220𝑖. This coincides with Theorem 3 on condition 𝛿2 > Θ+Ξ₯ Ξ– . Based on figure 3(b), the dynamics between prey and predator begin to stabilize at 1500 days to a specific point. CONCLUSIONS The Rosenzweig-MacArthur predator-prey model with anti-predator behavior has been studied. 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