Abstract
In this article, an eco-epidemiological model with strong Allee effect in prey population growth is presented by a system of delay differential equations. The time lag in terms of the delay parameter corresponds to the predator gestation period. We inspect elementary mathematical characteristic of the proposed model such as uniform persistence, stability and Hopf bifurcation at the interior equilibrium point of the system. We execute several numerical simulations to illustrate the proposed mathematical model and our analytical findings. We use basic tools of nonlinear dynamic analysis as first return maps, Poincare sections and Lyapunov exponents to identify chaotic behavior of the system. We observe that the system exhibits chaotic oscillation due to the increase of the delay parameter. Such chaotic behavior can be suppressed by the strength of Allee effect.
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Acknowledgments
SB’s and MD’s research is supported by the junior research fellowship from the University Grants Commission, Government of India. SS’s research work is supported by NBHM postdoctoral fellowship. JC’s research is partially supported by a DAE project (Ref No. 2/48(4)/2010-R & D II/8870).
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Appendix
Appendix
1.1 Proof of the Proposition 1:
Proof
The Jacobian matrix of the model (2) at its any equilibrium point \((S_{*}, I_{*}, P_{*})\) is described as follows
After substituting \(E_i = \left( S^*,I^*,P^*\right) ,~ i~=~0,~\theta ,~1,~2\) into (23), we obtain the eigenvalues for each equilibrium:
-
1.
\(E_0 = \left( 0, 0, 0\right) \) is always locally asymptotically stable since eigenvalues associated with (23) at \(E_0\) can be presented as follows:
$$\begin{aligned} \lambda _1= & {} -\theta ~~ \left( <0\right) ,\quad \lambda _2 = -\mu ~~ (<0) \quad \hbox {and}\\ \lambda _3= & {} -d~~ (<0). \end{aligned}$$ -
2.
\(E_\theta = \left( \theta , 0, 0\right) \) is always unstable since eigenvalues associated with (23) at \(E_1\) are given by,
$$\begin{aligned} \lambda _1= & {} \theta (1-\theta ) ~~(>0),\\ \lambda _2= & {} \beta \theta -\mu \biggl \{^{<0\quad \text{ if } \beta \theta <\mu }_{>0\quad \text{ if } \beta \theta >\mu }\quad \text{ and } \lambda _3 = -d \left( <0\right) . \end{aligned}$$ -
3.
\(E_1 = \left( 1, 0, 0\right) \) is locally asymptotically stable if \(\beta < \mu \) since eigenvalues associated with (23) at \(E_1\) are given by,
$$\begin{aligned} \lambda _1= & {} \theta -1 ~~(<0),\\ \lambda _2= & {} \beta -\mu \biggl \{^{<0\quad \text{ if } \beta <\mu }_{>0\quad \text{ if } \beta >\mu }~~~~ \text{ and } \lambda _3 = -d \left( <0\right) . \end{aligned}$$ -
4.
\(E_{2} = \left( \frac{\mu }{\beta }, \frac{\left( \frac{\mu }{\beta }-\theta \right) \left( 1-\frac{\mu }{\beta }\right) }{\frac{\mu }{\beta }+\beta -\theta },0\right) \) is locally asymptotically stable if \(\alpha \frac{\left( \frac{\mu }{\beta }-\theta \right) \left( 1-\frac{\mu }{\beta }\right) }{\frac{\mu }{\beta }+\beta -\theta } -d<0\) and \(1<\frac{\beta }{\mu }<\min \Bigg \{\frac{1}{\theta },\frac{\beta -\theta +\sqrt{\beta ^2-\beta \theta +\beta }}{\beta +\beta \theta -\theta ^2}\Bigg \}\). Since the eigenvalues associated with (23) at \(E_{2}\) are given by
$$\begin{aligned} \lambda _3= & {} \alpha \frac{\left( \frac{\mu }{\beta }-\theta \right) \left( 1-\frac{\mu }{\beta }\right) }{\frac{\mu }{\beta }+\beta -\theta }\\&-d\biggl \{^{<0\quad \text{ if } \alpha \frac{\left( \frac{\mu }{\beta }-\theta \right) \left( 1-\frac{\mu }{\beta }\right) }{\frac{\mu }{\beta }+\beta -\theta } -d<0}_{>0\quad \text{ if } \alpha \frac{\left( \frac{\mu }{\beta }-\theta \right) \left( 1-\frac{\mu }{\beta }\right) }{\frac{\mu }{\beta }+\beta -\theta } -d>0} \end{aligned}$$and the other two eigenvalues are the roots of the equation
$$\begin{aligned} \lambda ^2 - A\lambda +B = 0. \end{aligned}$$Where
$$\begin{aligned} \begin{array}{lll} A &{}=&{} \frac{\mu }{\beta }\left( 1-2\frac{\mu }{\beta }-\frac{\left( \frac{\mu }{\beta }-\theta \right) \left( 1-\frac{\mu }{\beta }\right) }{\frac{\mu }{\beta }+\beta -\theta }+\theta \right) \\ &{}=&{} \frac{\left( \beta +\beta \theta -\theta ^2\right) \left( \frac{\beta }{\mu }\right) ^2-2\frac{\beta }{\mu }(\beta -\theta )-1}{\left( \frac{\beta }{\mu }\right) ^2\left( 1+\frac{\beta }{\mu }(\beta -\theta )\right) }\\ B &{}=&{} \mu \left( \frac{\mu }{\beta }-\theta \right) \left( 1-\frac{\mu }{\beta }\right) >0 \end{array} \end{aligned}$$(24)Thus, we have
$$\begin{aligned}&A>0\quad \text{ if } \frac{\beta }{\mu }>\frac{\beta -\theta +\sqrt{\beta ^2-\beta \theta +\beta }}{\beta +\beta \theta -\theta ^2} \text{ while } \\&A<0\quad \text{ if } \frac{\beta }{\mu }<\frac{\beta -\theta +\sqrt{\beta ^2-\beta \theta +\beta }}{\beta +\beta \theta -\theta ^2}. \end{aligned}$$Therefore, \(E_2\) exists and is locally asymptotically stable if
$$\begin{aligned} 1<\frac{\beta }{\mu }<\min \Bigg \{\frac{1}{\theta },\frac{\beta -\theta +\sqrt{\beta ^2-\beta \theta +\beta }}{\beta +\beta \theta -\theta ^2}\Bigg \} \end{aligned}$$
Again, the Jacobian matrix of the model (2) at its interior equilibrium point \(\widetilde{E} = (\widetilde{S}, \widetilde{I}, \widetilde{P})\) can be written as follows
where its characteristic equation reads as follows:
with \(\lambda _i, i = 1,2,3\) being roots of (26). If all the real parts of \(\lambda _i, i=1,2,3\) are negative, then we have
Thus, interior equilibrium is locally asymptotically stable if the following conditions hold
So, from (27), we see that \(E_2^*\) is stable and \(E_1^*\) is unstable.
1.2 Proof of the permanence of the model (1)
Proof
In this section, we shall prove that the boundary planes of \({\mathbb {R}}^{3}_{+}\) repel the positive solutions of system (1) uniformly. Let us define
where \(C([-\tau , 0],~{\mathbb {R}}^{3}_{+})\) denote the space of continuous function mapping \([-\tau , 0]\) in to \({\mathbb {R}}^{3}_{+}\).
If \(C_{0} = C_{1}\cup C_{2}\cup C_{3}\) and \(C^{0} = int C\left( [-\tau , 0], {\mathbb {R}}^{3}_{+}\right) \), it suffices to show that there exists an \(\epsilon _{0} >0 \) such that for any solution \(u_{t}\) of system (1) initiating from \(C_{0}\), \(\displaystyle {\lim _{t\rightarrow \infty }\inf \ d(u_{t}, C_{0} )\ge \epsilon _{0}}\).
Now, we verify below that the conditions of Lemma 1 are satisfied. By definition of \(C_{0}\) and \(C^{0}\) and system (1), it is easy to see that \(C_{0}\) and \(C^{0}\) are positively invariant. Moreover, it is clear that conditions (i) and (ii) of Lemma 1 are satisfied. Thus, we need to confirm conditions (iii) and (iv).
Three constant solutions in \(C_{0}\) corresponding to \(\left( S(t) = 0, I(t) = 0, P(t) = 0\right) \), \(\left( S(t)=1, I(t)=0, P(t)=0\right) \) and \(\left( S(t) = S_{2}, I(t) = I_{2}, P(t) = 0\right) \) are respectively \(E_{0}\), \(E_{1}\) and \(E_{2}\).
If \(\left( S(t), I(t), P(t)\right) \) is any solution of system (1) initiating from \(C_{1}\) with \(\psi _{1}(0) = 0\) then \(S(t)\rightarrow 0\), \(I(t)\rightarrow 0\), \(P(t)\rightarrow 0\) as \(t\rightarrow \infty \). If \(\left( S(t), I(t), P(t)\right) \) is a solution of system (1) initiating from \(C_{2}\) with \(\psi _{1}(0) > 0\), it follows that \(S(t)\rightarrow 1\), \(I(t)\rightarrow 0\), \(P(t)\rightarrow 0\) as \(t\rightarrow \infty \). If \(\left( S(t), I(t), P(t)\right) \) is a solution of system (1) initiating from \(C_{3}\) with \(\psi _{1}(0) \psi _{2}(0) > 0\), it follows that \(S(t)\rightarrow S_{2}\), \(I(t)\rightarrow I_{2}\), \(P(t)\rightarrow 0\) as \(t\rightarrow \infty \).
This shows that invariant sets \(E_{0}\), \(E_{1}\) and \(E_{2}\) are isolated invariant, and then, \(E_{0}\), \(E_{1}\) and \(E_{2}\) are an isolated as well as an acyclic covering, satisfying condition (iii) of Lemma 1.
We now show that \(W^{s}(E_{0}) \cap C^{0} = {\varPhi }\), \(W^{s}(E_{1}) \cap C^{0} = {\varPhi }\) and \(W^{s}(E_{2}) \cap C^{0} = {\varPhi }\). The proof for the first part is simple, so we ignore it. We shall prove the second part through contradiction. Let us assume that \(W^{s}(E_{1}) \cap C^{0} \ne {\varPhi }\), then there exists a positive solution \(\left( S(t), I(t), P(t)\right) \) of system (1) such that \(\left( S(t), I(t), P(t)\right) \rightarrow (1, 0, 0)\) as \(t\rightarrow +\infty \). Let us choose \(\epsilon _{1} > 0 \) small enough such that
for some \(t>t_{1}\), where \(t_{1}\) be sufficiently large. Then, from first and second equations of the system (1), we have for \(t>t_{1}\)
Now let us consider
Let \(V = (v_1,v_2)\) and \(\zeta >0\) be small enough such that \(\zeta v_1<S(t_1),\zeta v_2<I(t_1)\). If \(\left( y_1(t),y_2(t)\right) \) is a solution of system (29) satisfying \(y_i(t_1) = \zeta v_i, i = 1, 2\). We know from comparison theorem that \(S(t) > y_1(t),~ I(t) > y_2(t)\) for all \(t>t_1\). We can check easily that the system (29) has a unique positive equilibrium
Now \(S(t) > y_1(t), I(t) > y_2(t)\) for all \(t>t_1\) and \(\displaystyle {\lim _{t\rightarrow \infty } y_2(t)}=y^*_2\). This is a contradiction. Hence, \(W^{s}(E_{2} ) \cap C^{0} = {\varPhi }\).
Let \(W^{s}(E_{2}) \cap C^{0} \ne {\varPhi }\). Then, there exists a positive solution \(\left( S(t), I(t), P(t)\right) \) of the system such that \(\left( S(t), I(t), P(t)\right) \rightarrow (S_{2}, I_{2}, 0)\) as \(t\rightarrow \infty \). Let us choose \(\epsilon _2>0\) small enough such that \(I_2-\epsilon _2 < I(t) < I_2+\epsilon _2\) for \(t > t_2-\tau \).
Then, from third equation of the system (1), we have for \(t > t_{2}-\tau \)
Now, let us consider
Let \(u_{1}\) and \(v>0\) be small enough such that \(vu_{1}<P(t_{2})\). If \(z_{1}\) is a solution of system (31) satisfying \(z_{1}(t_{2})=wu_{1}\), we know from comparison theorem, \(P(t)\ge z_{1}(t)\) for all \(t >t_{2}-\tau \). We also observe that the solution \(z_{1}\) of Eq. (31) satisfies \(\displaystyle {\lim _{t\rightarrow \infty } z_{1}(t)} \rightarrow +\infty \) (from condition (ii)).
Since \(P(t)\ge z_{1}(t)\) for all \( t >t_{2} \), so \(\displaystyle {\lim _{t\rightarrow \infty } P(t)}\nrightarrow 0\). This contradicts that \(W^{s}(E_{2} ) \cap C^{0} = {\varPhi }\). From Lemma 1, we conclude that \(C_{0}\) repels the positive solutions of (1) uniformly. Hence, the system (1) is permanent. This proves the theorem.
1.3 Direction and stability of Hopf bifurcation of model (1)
We consider the transformation \(z_{1}(t) = S(\tau t)-S_{*}\), \(z_{2}(t) = I(\tau t)-I_{*}\), \(z_{3}(t) = P(\tau t)-P_{*}\).
Let \(\tau =\tau ^{*}+\mu \), \(\mu \in \mathbf {R}\). Then, \(\mu =0\) is the Hopf bifurcation value of the system (1). The Eq. (1) can be written in the form
where \(z(t)=(z_{1}(t), z_{2}(t), z_{3}(t))^{T}\in \mathbf {R^{3}}\). For \(\psi =(\psi _{1}, \psi _{2}, \psi _{3})^{T}\in \mathbf {C}([-1, 0], \mathbf {R^{3}_{+}})\); \(L_{\mu }: \mathbf {C}\rightarrow \mathbf {R}\) and \(F: \mathbf {R}\times \mathbf {C}\rightarrow \mathbf {R}\) are given by
and
where
By the Riesz representation theorem [87], there exists a function \(\eta (\theta , \mu )\) of bounded variation for \(\theta \in [-1, 0]\), such that
In fact, we can choose
where \(\delta \) is defined by \(\delta (\theta )=\Big \{^{1, \ \ \theta =0,}_{0, \ \ \theta \ne 0.} \)
For \(\psi \in \mathbf {C}^{1}\left( [-1, 0], \mathbf {R^{3}_{+}}\right) \), define
and
Then, the system (32) is of the form
where \(z_{t}(\theta )=z_{t}(t+\theta )\) for \(\theta \in [-1, 0]\).
For \(\phi \in \mathbf {C}^{1}([0, 1], (\mathbf {R^{3}_{+}})^{*})\), define
and a bilinear inner product
where \(\eta (\theta )=\eta (\theta , 0)\). Clearly, A(0) and \(A^{*}\) are adjoint operators. We know that \(\pm i\rho _{0}\tau ^{*}\) are eigenvalues of A(0). So, they are also eigenvalues of \(A^{*}\). Now we search for the eigenvector of A(0) and \(A^{*}\) corresponding to \(i\rho _{0}\tau ^{*}\) and \(-i\rho _{0}\tau ^{*}\) respectively.
We assume that \(q(\theta )=(1, u, w)^{T} e^{i\rho _{0}\tau ^{*}\theta }\) and \(q^{*}(s)\) are the eigenvectors of A(0) and \(A^{*}\) corresponding to \(i\rho _{0}\tau ^{*}\) and \(-i\rho _{0}\tau ^{*}\), respectively. Then, we have \(A(0)q(\theta )=i\rho _{0}\tau ^{*}q(\theta )\). By the definition of A(0) and from (35) and (36), it follows that
Then, we can get \(q(0)=(1, u, w)^{T}\),
where
Similarly, we can obtain
We choose D in such a way that \(\langle q^{*}(s), q(\theta ) \rangle =1\), \(\langle q^{*}(s), \overline{q}(\theta ) \rangle =0\).
Hence
Thus, we can choose D as \(D=\frac{1}{1+\overline{u}u^{*}+\overline{w}w^{*}+ \tau ^{*} \overline{w} \alpha (u^{*}P_*+ w^{*}I_*) e^{i\rho _{0}\tau ^{*}}}\).
To describe the center manifold \(\mathbf {C}_{0} \) at \(\mu =0 \), we compute the coordinates by using the same notations and procedures as proposed by Hassard et al. [36].
Let \( z_{t}\) be the solution of of Eq. (32) when \(\mu =0\).
Define
On the center manifold \(\mathbf {C}_{0} \), we have
where
\(\text {z}\) and \(\overline{\text {z}}\) are local coordinates for center manifold \(\mathbf {C}_{0}\) in the direction of \(q^{*}\) and \(\overline{q}^{*}\). Here W is real when \(z_{t}\) is real. Now, we consider only real solutions. For solution \(z_{t} \in \mathbf {C}_{0}\) of Eq. (32), since \(\mu =0\), we can obtain
we rewrite this equation as \(\dot{\text {z}}=i\rho _{0}\tau ^{*}\text {z}+g(\text {z}, \overline{\text {z}})\) with
Then, from Eq. (40), we have
Thus, from Eq. (41), we can get
Comparing with the coefficients with (41), we can obtain
To calculate the value of \(g_{21}\), we need to compute the values of \(W_{20}(\theta )\) and \(W_{11}(\theta )\). From Eqs. (37) and (40), we have
where
Expanding the above series and comparing the corresponding coefficients, we obtain
From Eq. (45), we know that for \(\theta \in [-1, 0),\)
Comparing the coefficients with (46) gives that
and
Since \(q(\theta )=(1, u, w)^{T} e^{i\rho _{0}\tau ^{*}\theta }\), hence
where \(E_{1}=(E_{1}^{(1)}, E_{1}^{(2)}, E_{1}^{(3)}) \in \mathbf {R}^{3}\) is a constant vector.
Similarly, from Eqs. (47) and (50), we get
where \(E_{2}=(E_{2}^{(1)}, E_{2}^{(2)}, E_{2}^{(3)}) \in \mathbf {R}^{3}\) is a constant vector.
In what follows, we shall seek appropriate \(E_{1}\) and \(E_{2}\) in (51) and (52) respectively. From the definition of A and (47), we obtain
and
where \(\eta (\theta )=\eta (0, \theta )\). From (47), we have
and
Noting that
and
and putting (51) and (55) into (53), we get
which implies that
It follows that
where
Similarly putting (52) and (56) into (54), we have
which implies that
and hence,
where
From the above analysis, we can compute the following quantities:
The above three quantities \(\mu _2\), \(\beta _2\) and \(\tau _2\) will determine the direction, stability and the periods of the bifurcating periodic solutions.
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Biswas, S., Saifuddin, M., Sasmal, S.K. et al. A delayed prey–predator system with prey subject to the strong Allee effect and disease. Nonlinear Dyn 84, 1569–1594 (2016). https://doi.org/10.1007/s11071-015-2589-9
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DOI: https://doi.org/10.1007/s11071-015-2589-9