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Analysis of long transients and detection of early warning signals of extinction in a class of predator–prey models exhibiting bistable behavior

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Abstract

In this paper, we develop a method of analyzing long transient dynamics in a class of predator–prey models with two species of predators competing explicitly for their common prey, where the prey evolves on a faster timescale than the predators. In a parameter regime near a singular zero-Hopf bifurcation of the coexistence equilibrium state, we assume that the system under study exhibits bistability between a periodic attractor that bifurcates from the singular Hopf point and another attractor, which could be a periodic attractor or a point attractor, such that the invariant manifolds of the coexistence equilibrium point play central roles in organizing the dynamics. To find whether a solution that starts in a vicinity of the coexistence equilibrium approaches the periodic attractor or the other attractor, we reduce the equations to a suitable normal form, and examine the basin boundary near the singular Hopf point. A key component of our study includes an analysis of the long transient dynamics, characterized by their rapid oscillations with a slow variation in amplitude, by applying a moving average technique. We obtain a set of necessary and sufficient conditions on the initial values of a solution near the coexistence equilibrium to determine whether it lies in the basin of attraction of the periodic attractor. As a result of our analysis, we devise a method of identifying early warning signals, significantly in advance, of a future crisis that could lead to extinction of one of the predators. The analysis is applied to the predator–prey model considered in Sadhu (Discrete Contin Dyn Syst B 26:5251–5279, 2021) and we find that our theory is in good agreement with the numerical simulations carried out for this model.

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Acknowledgements

The authors thank the anonymous reviewers for their suggestions that led to a significant improvement in the presentation of the article. S.S acknowledges the Provost’s Summer Research Fellowship program in Georgia College for supporting this research and is grateful for having fruitful discussions at the workshops on “Advances in Mathematical Ecology" at the Fields Institute in December 2022 and at the University of Pittsburgh in June 2023.

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Appendix

Appendix

Expressions for the coefficients in system (6): The expressions for \(\omega \), \(F_{13}\), \(F_{111}\), \(H_3\), \(H_{11}\), and \(\alpha (h)\) in Theorem 3.1 are given below:

$$\begin{aligned} \left\{ \begin{array}{lllll} \omega &{}=\sqrt{-\bar{x} (\bar{y}\bar{\phi _y}\bar{\chi _x}+\bar{z} \bar{\phi _z}\bar{\psi _x})},\\ F_{13}&{}= \bar{x} \bar{\phi _z}(\bar{z}\bar{\psi _z}-\bar{y}\bar{\chi _y})+\frac{\bar{x}}{\bar{\phi _y}}(\bar{y}\bar{\phi ^2_y}\bar{\chi _z} -\bar{z}\bar{\phi ^2_z}\bar{\psi _y}) +\frac{\omega ^2}{\bar{\phi _{xx}}\bar{\phi _y}}(\bar{\phi _{xz}}\bar{\phi _y} - \bar{\phi _{xy}}\bar{\phi _z}),\\ F_{111} &{}= \frac{\omega ^2}{\bar{x}^2\bar{\phi ^2_{xx}}} (3\bar{\phi }_{xx}+\bar{x}\bar{\phi }_{xxx}) +\frac{\bar{x}}{\omega ^2}[\bar{y}\bar{\chi _x}(\bar{y} \bar{\phi _y}\bar{\chi _y} + \bar{z}\bar{\phi _z}\bar{\psi _y})+ \bar{z}\bar{\psi _x} (\bar{y}\bar{\phi _y}\bar{\chi _z}+\bar{z} \bar{\phi _z}\bar{\psi _z} )] \\ &{}+\frac{1}{\bar{x}\bar{\phi }_{xx}}[\bar{y}\bar{\chi _x}(\bar{\phi _y}+\bar{x}\bar{\phi }_{xy}) +\bar{z}\bar{\psi _x}(\bar{\phi _z} +\bar{x}\bar{\phi }_{xz})+ \bar{x}(\bar{y}\bar{\phi _y}\bar{\chi }_{xx}+ \bar{z}\bar{\phi _z}\bar{\psi }_{xx})], \\ H_3 &{}= \frac{\bar{x}\bar{y} \bar{z}}{\omega ^2}[\bar{\psi _x}(\bar{\phi _y} \bar{\chi _z} -\bar{\phi _z}\bar{\chi _y} ) - \bar{\chi _x}(\bar{\phi _y} \bar{\psi _z} -\bar{\phi _z}\bar{\psi _y} ) ],\\ H_{11} &{}= \frac{\bar{x}\bar{y}\bar{z}}{\omega ^4}\bar{\phi _y}(\bar{\psi _x}-\bar{\chi _x}) \left( \bar{y} \bar{\chi _x}\bar{\chi _y}+\bar{z}\bar{\psi _x}\bar{\psi _z}\right) + \frac{\bar{y}\bar{z}}{\omega ^2 \bar{x}\bar{\phi }_{xx}}\bar{\phi _y} \left( \bar{z} \bar{\psi _x}\bar{\chi }_{xx}-\bar{x}\bar{\chi _x}\bar{\psi }_{xx}\right) , \\ \alpha (h)&{}= \left\{ -\begin{pmatrix} (\bar{f}_1)_{xx} &{} (\bar{f}_1)_{xy} &{} (\bar{f}_1)_{xz} \end{pmatrix} J^{-1} \begin{pmatrix} (\bar{f}_1)_h \\ (\bar{f}_2)_h\\ (\bar{f}_3)_h \end{pmatrix} +(\bar{f}_1)_{xh}\right\} \Big (\frac{h-\bar{h}}{\zeta }\Big ) \\ {} &{} -\frac{1}{\omega ^2}\{(\bar{f}_1)_y [(\bar{f}_2)_y (\bar{f}_2)_x + (\bar{f}_2)_z (\bar{f}_3)_x] + (\bar{f}_1)_z [(\bar{f}_3)_y (\bar{f}_2)_x + (\bar{f}_3)_z (\bar{f}_3)_x) ] \}. \end{array} \right. \end{aligned}$$

Hopf bifurcation of system (3): System (3) undergoes a Hopf bifurcation at \(h=\bar{h}+\zeta A+O(\zeta ^{3/2})\) for sufficiently small \(\zeta >0\), where A is the solution of the equation

$$\begin{aligned} \left\{ -\begin{pmatrix} (\bar{f}_1)_{xx}&(\bar{f}_1)_{xy}&(\bar{f}_1)_{xz} \end{pmatrix} J^{-1} \begin{pmatrix} (\bar{f}_1)_h \\ (\bar{f}_2)_h\\ (\bar{f}_3)_h \end{pmatrix} +(\bar{f}_1)_{xh}\right\} A= & {} \frac{1}{\omega ^2}\{(\bar{f}_1)_y [(\bar{f}_2)_y (\bar{f}_2)_x + (\bar{f}_2)_z (\bar{f}_3)_x] \nonumber \\+ & {} (\bar{f}_1)_z [(\bar{f}_3)_y (\bar{f}_2)_x + (\bar{f}_3)_z (\bar{f}_3)_x) ] \}.\nonumber \\ \end{aligned}$$
(27)

Transformation to normal coordinates: The transformations from the original coordinates (xyz) to the normal form coordinates (uvw) are given below:

$$\begin{aligned} \left\{ \begin{array}{ll} u &{}= \frac{\bar{f_{1xx}}}{\omega } X -\delta (\bar{f_{1y}} B_1(A_1, A_2, X, Y, Z)+\bar{f_{1z}}B_2(A_1, A_2, X, Y, Z) \\ &{}\quad + \frac{\delta }{3}C_{uv} \Big [ \Big ( \frac{\bar{f_{1xx}}}{\omega } X - \delta (\bar{f_{1y}} B_1(A_1, A_2, X,Y,Z) \\ &{} \quad +\bar{f_{1z}}B_2((A_1, A_2, X, Y, Z))\Big )^2 \Big (-1+\frac{\bar{f_{1xx}}}{2\omega ^2} (\bar{f_{1y}}Y+\bar{f_1}_zZ )\Big ) \\ &{}\quad +\frac{\bar{f^2_{1xx}}}{\omega ^4} (\bar{f_{1y}}Y+\bar{f_{1z}}Z)^2 \Big ],\\ v &{}= \frac{\bar{f_{1xx}}}{\omega ^2} (\bar{f_{1y}}Y+\bar{f_{1z}}Z),\\ w &{}=- \frac{\bar{f_{1xx}} \bar{f_{1y}}}{\omega ^2 \bar{f_{1z}}} \Big [ \Big (1+ \frac{\bar{f_{2x}}\bar{f_1}_{y}}{\omega ^2}\Big ) Y + {\bar{f_{2x}}\bar{f_{1z}}}{\omega ^2} Z + \frac{\delta }{\omega }\bar{f_1}_{xx} A_1 X\Big ], \end{array} \right. \end{aligned}$$
(28)

where

$$\begin{aligned} X = \frac{x-x_0}{\sqrt{\zeta }}, \ Y = \frac{y-y_0}{\zeta }, \ Z = \frac{z-z_0}{\zeta }, \end{aligned}$$

and

$$\begin{aligned} A_1= & {} \left( 1+ \frac{\bar{f_{2x}}\bar{f_{1y}}}{\omega ^2}\right) (\bar{f_{2y}}\bar{f_{2x}} + \bar{f_{2z}}\bar{f_{3x}}) + \frac{\bar{f_{2x}}\bar{f_{1z}}}{\omega ^2} (\bar{f_{3y}}\bar{f_{2x}} + \bar{f_{3z}}\bar{f_{2x}}), \\ A_2= & {} \frac{\bar{f_{3x}}\bar{f_{1y}}}{\omega ^2} (\bar{f_{2y}}\bar{f_{2x}} + \bar{f_{2z}}\bar{f_{3x}}) + \left( 1+\frac{\bar{f_{3x}}\bar{f_{1z}}}{\omega ^2}\right) (\bar{f_{3y}}\bar{f_{2x}} + \bar{f_{3z}}\bar{f_{2x}}),\\ C_{uv}= & {} -\frac{1}{\omega ^2} \Big [ \bar{f_{1y}}(\bar{f_{2y}}\bar{f_{2x}} + \bar{f_{2z}}\bar{f_{3x}}) + \bar{f_{1z}} (\bar{f_{3y}}\bar{f_{2x}} + \bar{f_{3z}}\bar{f_{2x}})\Big ] \\ {}- & {} \frac{1}{\bar{f_{1xx}}} \Big [\bar{f_{1xy}}\bar{f_{2x}} + \bar{f_{1xz}}\bar{f_{3x}} + \bar{f_{1y}}\bar{f_{2xx}} + \bar{f_{1z}}\bar{f_{3xz}}\Big ], \\ B_1= & {} -\frac{\bar{f_{1xx}}}{\omega ^4} (\bar{f_{1y}}Y+ \bar{f_{1z}}Z) (\bar{f_{2y}}\bar{f_{2x}} + \bar{f_{2z}}\bar{f_{3x}}) +\frac{\bar{f_{1xx}}\bar{f_{2xx}}}{2\omega ^2} + \frac{\bar{f_{2y}}\bar{f_{1xx}} }{\omega ^2}\Big [\left( 1+ \frac{\bar{f_{2x}}\bar{f_{1y}}}{\omega ^2}\right) Y \\ {}+ & {} \frac{\bar{f_{2x}}\bar{f_{1z}}}{\omega ^2} Z + \frac{\delta A_1 X}{\omega } \Big ] + \frac{\bar{f_{2z}}\bar{f_{1xx}} }{\omega ^2}\Big [ \frac{\bar{f_{3x}}\bar{f_{1y}}}{\omega ^2} Y + \Big (1+\frac{\bar{f_{3x}}\bar{f_{1z}}}{\omega ^2}\Big ) Z + \frac{\delta A_2 X}{\omega } \Big ],\\ B_2= & {} -\frac{\bar{f_{1xx}}}{\omega ^4} (\bar{f_{1y}}Y+ \bar{f_{1z}}Z) (\bar{f_{3y}}\bar{f_{2x}} + \bar{f_{3z}}\bar{f_{3x}}) + \frac{\bar{f_{1xx}}\bar{f_{3xx}}}{2\omega ^2} + \frac{\bar{f_{3y}}\bar{f_{1xx}} }{\omega ^2}\Big [\left( 1+ \frac{\bar{f_{2x}}\bar{f_{1y}}}{\omega ^2}\right) Y \\+ & {} \frac{\bar{f_{2x}}\bar{f_{1z}}}{\omega ^2} Z + \frac{\delta A_1 X}{\omega } \Big ] + \frac{\bar{f_{3z}}\bar{f_{1xx}} }{\omega ^2}\Big [ \frac{\bar{f_{3x}}\bar{f_{1y}}}{\omega ^2} Y + \Big (1+\frac{\bar{f_{3x}}\bar{f_{1z}}}{\omega ^2}\Big ) Z + \frac{\delta A_2 X}{\omega } \Big ]. \end{aligned}$$

Proof of Lemma 3.1:

Let \(M\ge N\) be the largest integer such that \(\{u(\tau _i)\}_{i=1}^M\) decreases and \(w(\tau )>0\) on \([0, \tau _M]\). By Remark 3.1, the expression for \(\bar{u^2}_{\text {est}}(\tau )\), defined by (15)–(16), holds up to \(O(\delta ^2)\) for all \(\tau \in [\tau _1, \tau _M]\). It is clear from (P2) that \(\bar{w}(\tau )\) also has a decreasing envelope for all \(\tau \in [\tau _1, \tau _N]\). Estimating \(\bar{w}(\tau )\) by \(\bar{w}_{\text {base}}(\tau )\), where \(\bar{w}_{\text {base}}(\tau )\) is defined by (17), it can be easily verified that \(\bar{w}_{\text {base}}(\tau )\) has a unique critical point at \(\tau =\bar{\tau }_m>0\), where

$$\begin{aligned} \bar{\tau }_m = \frac{1}{\delta H_3-b_2}\ln \Big (\frac{H_{11}b_1b_2e^{(\delta H_3-b_2) \tau _1}}{H_3(2\bar{w}(\tau _1)(\delta H_3-b_2)+\delta H_{11}b_1)} \Big ) \end{aligned}$$

if (18) holds. The critical point corresponds to a minimum of \(\bar{w}_{\text {base}}(\tau )\) with

$$\begin{aligned}\bar{w}_{\text {base}}(\bar{\tau }_m)= \frac{-H_{11}b_1}{2 H_3}e^{b_2(\bar{\tau }_m-\tau _1)}>0. \end{aligned}$$

Since \(\bar{w}(\tau )-\bar{w}_{\text {base}}(\tau )=O(\delta ^2)\) as long as \(w(\tau ) =O(\delta )\), it follows that \(\bar{w}(\tau )\) and the lower envelope of \(w(\tau )\) also attain their minima. Consequently, \(w(\tau )\) attains its global minimum at \(\tau = {\tau }_{\textrm{min}}\), where \({\tau }_{\textrm{min}}= {\bar{\tau }}_m +O(\delta ^2)\). Finally, by (P2), since \(w(\tau )\) has a decreasing envelope on \([\tau _1, \tau _N]\), it is clear that \({\tau }_{\textrm{min}}\ge \tau _N\). \(\square \)

Proof of Lemma 3.2:

We start by considering the relative position of \(w(\tau )\) with respect to \(-\frac{H_{11}}{2H_3} (u^2+v^2)(\tau )\) and show that there exists some \(\tau _c> {\tau }_{\textrm{min}}\) such that \(w(\tau )\) intersects with the upper envelope of \(-\frac{H_{11}}{2H_3} (u^2+v^2)(\tau )\) at \(\tau =\tau _c\). By (P2), \(w(\tau )<-\frac{H_{11}}{2H_3} (u^2+v^2)(\tau )\) for all \(\tau \in [0, \tau _{N}]\). Let \(\{t_k\}_{k=1}^{M}\) and \(\{s_k\}_{k=1}^{M-1}\) be an increasing sequence of locations of relative maxima and minima of \(v(\tau )\) in I respectively with \(t_k<\tau _k<s_k\). Since the trajectory is spiraling inwards, we have that \(\{v(t_k)\}_{k=1}^M\) and \(\{v(s_k)\}_{k=1}^{M-1}\) are decreasing and increasing respectively. Without loss of generality, we assume that \(|v(t_k)|\le |v(s_k)|\) for all \(1\le k \le M-1\). With the aid of (6), we note that the zeros of \(u(\tau )\) correspond to the critical points of \((u^2 +v^2)(\tau )\). Hence the relative maxima and minima of \((u^2 +v^2)(\tau )\) occur at \(s_k\) and \(t_k\) respectively with values \(v^2(s_k)\) and \(v^2(t_k)\) for all k. Denoting the local maxima and minima of \(-\frac{H_{11}}{2H_3} (u^2+v^2)(\tau )\) on the interval \([t_k, s_k]\) by \(M_k\) and \(m_k\) respectively, we then have that \(\{M_k\}_{k=1}^{M-1}\) and \(\{m_k\}_{k=1}^{M-1}\) are both decreasing. If \(u_{\text {env}}(\tau )\) and \(v_{\text {env}}(\tau )\) denote the upper envelopes of \(u(\tau )\) and \(v(\tau )\) respectively, we then have from (9) that up to \(O(\delta ^2)\), \(u_{\text {env}}(\tau )= Ae^{c_2(\tau -\tau _1)}\) and \(v_{\text {env}}(\tau )= Be^{c_2(\tau -\tau _1)}\) for \(\tau \in [\tau _1, \tau _M]\), where A is defined by (20) and

$$\begin{aligned} B= \frac{A}{c_2^2+\vartheta ^2},\ c_2=\frac{b_2}{2}=\frac{\alpha \delta }{2} +O(\delta ^k). \end{aligned}$$
(29)

Since \((u^2+v^2)(\tau )\le (u_{\text {env}}^2(\tau ) +v_{\text {env}}^2(\tau ))\) and \(|w(\tau )- \bar{w}_{\text {base}}(\tau )|=O(\delta ^2)\), it suffices to show that there exists some \(\tau _c \in ({\tau }_{\textrm{min}}, \tau _M)\) such that \(\bar{w}_{\text {base}}(\tau )\) intersects with \(\frac{-H_{11}}{2H_3} (u_{\text {env}}^2(\tau ) +v_{\text {env}}^2(\tau ))\) at \(\tau = \tau _c\). To this end, we first note from (17), (19) and (29) that \(\bar{w}_{\text {base}}(\tau )\) intersects with \(\frac{-H_{11}}{2H_3} (u_{\text {env}}^2(\tau ) +v_{\text {env}}^2(\tau ))\) if and only if \(\frac{-H_{11}(A^2+B^2)}{2H_3} > \frac{\delta H_{11}b_1}{2(b_2-\delta H_3)}\), i.e.

$$\begin{aligned} 1+\frac{1}{c_2^2+\vartheta ^2} > - \frac{1-e^{\frac{-4 \pi c_2}{\vartheta }}}{\frac{8\pi c_2}{\vartheta }\Big (\frac{2c_2}{\delta H_3}-1\Big )}. \end{aligned}$$
(30)

Since \(c_2 =O(\alpha \delta /2)\) and \(\vartheta \approx 1\), it then follows that the left hand side of (30) is approximately equal to 2, whereas the right hand side of (30) is less than 1/2 for \(\delta \) sufficiently small, and thus (30) holds. Next, we observe from (13) and (16) that as long as \(\bar{w}_{\text {base}}(\tau )\) is decreasing, \( \bar{w}_{\text {base}}(\tau )\le \frac{-H_{11}}{2H_3} b_1\). Since \(b_1 \approx A^2/2\) and \(c_2<0\), we then have from the definition of \(u_{\text {env}}\) and \(v_{\text {env}}\) that

$$\begin{aligned} \bar{w}_{\text {base}}(\tau ) \le \frac{-H_{11}}{2H_3} b_1< -\frac{H_{11}}{2H_3} (u_{\text {env}}^2(\tau ) +v_{\text {env}}^2(\tau )) \ \text {on} \ [\tau _1, \bar{\tau }_m]. \end{aligned}$$

Hence, for sufficiently small \(\delta >0\),

we must have \(\tau _c > \bar{\tau }_m = \tau _{\text {min}} +O(\delta ^2)\).

Finally, we note that the existence of \(\tau _c\) implies that \(w(\tau )\ge M_i\) on \([s_i, t_{i+1}]\) for some i between 1 and N, and hence the trajectory is in \({\mathbb {R}}^3 {\setminus } \Omega _{\alpha }\) over the interval \([s_i, t_{i+1}]\). The monotonic properties of \(\{M_k\}_{k=1}^{M-1}\) and \(\{m_k\}_{k=1}^{M-1}\), and the fact that \(\bar{w}'(\tau )>0\) for \(\tau >{\tau }_{\textrm{min}}\) will then imply that the trajectory is in \({\mathbb {R}}^3 {\setminus } \Omega _{\alpha }\) over the interval \([s_i, \tau _M]\). Choosing \(\tau _a=s_j\), where \(1<j<M\) is the smallest integer such that \(s_j\le \tau _c\) with \(w(s_j)>M_j\), and \(\tau _b=\tau _M\) then yields the result.

Proof of Lemma 3.3:

Let \(\{\tau _i\}_{i=1}^M\) be an increasing sequence of locations of relative maxima of \(u(\tau )\) such that \(\{u(\tau _i)\}_{i=1}^M\) is decreasing, where M is some integer greater than N to be chosen later. By Remark 3.1, we note that the expressions for \(\bar{u^2}_{\text {est}}(\tau )\) and \(\bar{w}_{\text {base}}(\tau ) \) defined by (15) and (17) respectively, hold up to \(O(\delta ^2)\) for all \(\tau \in [\tau _1, \tau _M]\). If \(\bar{w}(\tau _1)< \frac{\delta H_{11}b_1}{2(b_2-\delta H_3)}\), then it follows from (17) that \(\bar{w}_{\text {base}}(\tau )\) decreases and eventually becomes negative at \(\tau =\tau _e\), where

$$\begin{aligned} {\tau _e} = \frac{1}{\delta H_3-b_2}\ln \Big (\frac{\delta H_{11}b_1e^{(\delta H_3 -b_2)\tau _1}}{\delta H_{11}b_2 e^{b_2 \tau _1}+2\bar{w}(\tau _1)(\delta H_3-b_2)} \Big ). \end{aligned}$$

Since \( |w(\tau ) - \bar{w}_{\text {base}}(\tau )|=O(\delta ^2)\) and \(\bar{w}_{\text {base}}'(\tau )<0\) for all \(\tau \in [0, \tau _M]\), we must have that \(w(\tau )\) also changes its sign near \(\tau _e\). Let \(\tilde{\tau }=\tau _e+o(1)\) be such that \(w(\tilde{\tau })=0\). Choose M large enough such that \(\tilde{\tau } <\tau _{M}\). Since \(q_e\) is unstable along the w-direction and \(w'(\tilde{\tau }) = \frac{\delta }{2} H_{11}u^2 (\tilde{\tau })\le 0\) (follows from (6)), we cannot have \(w(\tau )=0\) for all \(\tau \ge \tilde{\tau }\), thereby leading to \(w(\tau )<0\le -\frac{H_{11}}{2H_3} (u^2+v^2)(\tau )\) for all \(\tau >\tilde{\tau }\). Thus \((u(\tau ), v(\tau ), w(\tau ))\in \Omega _{\alpha }\) for all \(\tau >\tilde{\tau }\). It is also evident from (6) that \(w'(\tau ) < \delta H_3 w\) and therefore if \(w(\bar{\tau })=-\tilde{k}\) for some \(\tilde{k}>0\) and \(\bar{\tau } > \tilde{\tau }\), then by Gronwall’s inequality, \(w(\tau ) < -\tilde{k} e^{\delta H_3 (\tau -\bar{\tau })} \rightarrow -\infty \) as \(\tau \rightarrow \infty \). Consequently, the fast variables \((u(\tau ), v(\tau ))\), governed by system (10), approach (0, 0) as \(\tau \rightarrow \infty \).

Next, we will prove that \(\Omega _{\alpha }\) is positively invariant with respect to \((u(\tau ),v(\tau ),w(\tau ))\) for all \(\tau \ge 0\). Denoting the sequences of local minima and maxima of \(-\frac{H_{11}}{2 H_3}(u^2+v^2)\) by \(\{m_k\}_{k=1}^{M-1}\) and \(\{M_k\}_{k=1}^{M-1}\) and their locations by \(\{t_k\}_{k=1}^{M-1}\) and \(\{s_k\}_{k=1}^{M-1}\) respectively with \(t_k<\tau _k<s_k\), and recalling that \((u(\tau ), v(\tau ), w(\tau )) \in \Omega _{\alpha }\) on \([0, \tau _N]\), we then have that \(w(\tau ) < m_k\) on the interval \([t_k, t_{k+1})\) for all \(1\le k\le N-1\). As in the proof of Lemma 3.2, we have that \(m_k =-\frac{H_{11}}{2H_3}v^2(t_k) =-\frac{H_{11}B^2}{2H_3} e^{b_2 (t_k-\tau _1)}\) up to \(O(\delta ^2)\), where B is defined by (29), and thus we have \(\bar{w}_{\text {base}}(\tau )\approx w(\tau ) <-\frac{H_{11}B^2}{2H_3} e^{b_2 (\tau -\tau _1)}\) on \([0, \tau _{N}]\), i.e.

$$\begin{aligned} \frac{\delta H_{11} b_1}{2(b_2-\delta H_3)} < -\frac{H_{11} B^2}{2H_3}. \end{aligned}$$
(31)

Since by our assumption \(\bar{w}(\tau _1)< \frac{\delta H_{11}b_1}{2(b_2-\delta H_3)}\), we have from (17) in combination with (31) that \(\bar{w}_{\text {base}}(\tau ) < m_k\) on \([t_k, t_{k+1})\) for all \( 1\le k\le M-1\), i.e. \(\bar{w}_{\text {base}}(\tau )< -\frac{H_{11}}{2 H_3}(u^2+v^2)(\tau )\) for all \(\tau \in [0,\tau _M]\). Finally, since \(w(\tau )=\bar{w}_{\text {base}}(\tau )+O(\delta ^2)\), we can conclude that the solution is in \(\Omega _{\alpha }\) for all \(\tau \in [0,\tau _M]\). Combining this with the fact that \((u(\tau ), v(\tau ), w(\tau ))\in \Omega _{\alpha }\) for all \(\tau >\tilde{\tau }\) proves the lemma. \(\square \)

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Sadhu, S., Chakraborty Thakur, S. Analysis of long transients and detection of early warning signals of extinction in a class of predator–prey models exhibiting bistable behavior. J. Math. Biol. 88, 70 (2024). https://doi.org/10.1007/s00285-024-02095-8

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