Abstract
Coral reefs provide refuge for prey and are important for the preservation of an oceanic ecosystem. However, they have been experiencing severe destruction by environmental changes and human activities. In this paper, we propose and analyze a tri-trophic food chain model consisting of coral, Crown-of-thorns starfish (CoTS), and triton in deterministic and stochastic environments. We investigate the effects of harvesting in the deterministic system and environmental noises in the stochastic system, respectively. The existence of possible steady states along with their stability is rigorously discussed. From the economic perspective, we examine the existence of the bionomic equilibrium and establish the optimal harvesting policy. Subsequently, the deterministic system is extended to a stochastic system through nonlinear perturbation. The stochastic system admits a unique positive global solution initiating from the interior of the positive quadrant. The long-time behaviors of the stochastic system are explored. Numerical simulations are provided to validate and complement our theoretical results. We show that over-harvesting of triton is not beneficial to coral reefs and modest harvesting of CoTS may promote sustainable growth in coral reefs. In addition, the presence of strong noises can lead to population extinction.
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Acknowledgements
Zhao, Liu and Fan are supported by NSFC (No.12071068); Wang is supported by NSERC Discovery Grant RGPIN-2020-03911 and NSERC Accelerator Grant RGPAS-2020-00090.
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Appendices
Appendix A: Proofs for Deterministic System
1.1 Proof of Lemma 1
Proof
According to the first equation of (1), we have
Similarly, we obtain
and
It is clear that \(x(t)>0\), \(y(t)>0\), and \(z(t)>0\) whenever \(x(0)>0\), \(y(0)>0\), and \(z(0)>0\). Then any solution from the first quadrant of xyz-plane gives a positive solution. This completes the proof. \(\square \)
1.2 Proof of Theorem 1
Proof
Define \(W=e_{1}e_{2}x+e_{2}y+z\). Taking time derivative of W leads to
Due to the positivity of parameters and the nonnegativity of solutions, we obtain
Taking the limit of the above inequality as t tends to \(\infty \), we obtain
Therefore, all solutions of (1) with initial values \((x(0),y(0),z(0))\in \mathbb {R}^{3}_{+}Z\) are eventually confined to the region
\(\square \)
1.3 Proof of Lemma 2
Proof
When \(n=1\), it is obvious that the claim is valid. Assume that the claim holds for \(n=2k-1\). Then, when \(n=2k+1\), one has
If \(\alpha (x)=0\) has one positive root, then the claim is valid. If all the roots are negative, then there is a contradiction with \(p_0>0\). Thus, \(\alpha (x)=0\) has at least a pair of conjugate complex roots denoted by \(x_0=a+bi, \bar{x}_0=a-bi\) with \(a\not =0\). Then,
where \(\tilde{\alpha }(x)=(-1)^{2k-1}x^{2k-1}+q_{2k-2}x^{2k-2}+\cdots +q_{1}x+q_{0}\) and \(q_{0}(a^{2}+b^{2})=p_{0}>0\). By the method of mathematical induction, \(\alpha (x)\) has at least one positive root if \(p_{0}>0\). The proof is complete. \(\square \)
1.4 Proof of Theorem 2
Proof
The local stability of equilibria is determined by computing the eigenvalues of the Jacobian matrix about each equilibrium. The Jacobian matrix of (1) at \(E_{0}\) is
The three eigenvalues of this matrix are
Obviously, \(E_{0}=(0,0,0)\) is unstable. The proof is complete. \(\square \)
1.5 Proof of Theorem 3
Proof
The Jacobian matrix of (1) at \(E_{x}\) reads
The three eigenvalues of this matrix are
The equilibrium \(E_{x}=(K_{1},0,0)\) is locally asymptotically stable if \(h_{1}>r_{2}+{e_{1}m_{1}K_{1}}/({a_{1}+K_{1}})\), otherwise it is a saddle point with stable manifold in \(x-z\) plane and unstable manifold in y direction. The proof is complete. \(\square \)
1.6 Proof of Theorem 4
Proof
The Jacobian matrix of (1) at \(E_{y}\) takes the form
Its eigenvalues are
Then it is not difficult to show that all the eigenvalues of \(J\mid _{E_{y}}\) have negative real parts when (6) is satisfied, then \(E_{y}\) is locally asymptotically stable. The proof is complete. \(\square \)
1.7 Proof of Theorem 5
Proof
By (3), one has
Substituting the above equation into the second equation of (3) shows that \(x_{3}\) is a positive solution of the following equation:
where \(\hat{\nu }=[{m_{1}K_{1}K_{2}(h_{1}-r_{2}-e_{1}m_{1})+a_{1}r_{1}r_{2}(2K_{1}-a_{1})}]/(r_{1}r_{2})\). From Lemma 2, it follows that \(x_{3}\) is positive if (7) is satisfied.
The Jacobian matrix of (1) at \(E_{xy}\) writes
The characteristic equation of \(J\mid _{E_{xy}}\) of (1) is
Then \(\lambda _{1}<0,~\lambda _{2}<0\), and \(\lambda _{3}<0\) under the conditions of Theorem 5. Thus, \(E_{xy}=(x_{3},y_{3},0)\) is locally asymptotically stable. The proof is complete. \(\square \)
1.8 Proof of Theorem 6
Proof
By (4), one has
It is trivial to show that \(E_{yz}=\left( 0,y_{4},z_{4}\right) \) exists when (8) is satisfied.
The Jacobian matrix of (1) evaluated at \(E_{yz}\) reads
The characteristic equation of \(J\mid _{E_{yz}}\) is
Then, \(\lambda _{1}<0\), \(\lambda _{2}<0\), and \(\lambda _{3}<0\) when (9) holds. Therefore, \(E_{yz}=(0,y_{4},z_{4})\) is locally asymptotically stable. The proof is complete. \(\square \)
1.9 Proof of Theorem 7
Proof
It is not difficult to show that \(y^*>0\) and \(z^*>0\) when (11) is satisfied. The existence of \(E_{xyz}(x^{*},y^{*},z^{*})\) follows from direct calculation. The Jacobian matrix of (1) evaluated at \(E_{xyz}\) is
The characteristic equation of \(J\mid _{E_{xyz}}\) of (1) is
where
It is obvious that \(A_{1}>0\) and \(A_{3}>0\) when the conditions in Theorem 7 are satisfied. By the Routh-Hurwitz criterion, it follows that \(E_{xyz}\) is locally asymptotically stable if \(A_{1}A_{2}>A_{3}\).
For the global asymptotic stability of \(E_{xyz}\), consider the positive definite Lyapunov function
where \(l_{1}\) and \(l_{2}\) are positive constants to be determined later.
By calculating the derivative of V(x, y, z) along the solutions of (1), we obtain
After simplification,
Set
such that the right hand side of (A2) can be written as \(-X^{\textrm{T}}DX\), where \(X^{\textrm{T}}=(x-x^{*},y-y^{*},z-z^{*})\) and the symmetric matrix D is given by
Then \({\textrm{d}V}/{\textrm{d}t}<0\) if and only if the symmetric matrix D is positive definite, which is equivalent to (12). Therefore, \(E_{xyz}\) is globally asymptotically stable. The proof is complete. \(\square \)
Appendix B: Proofs for Optimal Harvesting
1.1 Proof of Theorem 8
Proof
The bionomic equilibrium solves the system of algebraic equations
Define
Obviously, \(\bar{\Omega }\) is a positive invariant set of (B4). From the first equation, one has
Substituting \(\bar{y}\) into the third equation produces
where \(\bar{E}>0\) since \(d<\frac{e_{2}m_{2}\bar{y}}{a_{2}+\bar{y}}\). Similarly, we have
where \(\bar{z}>0\) since \(c>p_1q_1\bar{y}\). According to the second equation of (B4), we can see \(\bar{x}\) is a positive root of the following equation
which can be simplified by
where
must be a positive constant under the condition \(r_{1}<\frac{m_{1}K_{2}}{a_{1}}\) and \(\frac{p_1}{p_2}>e_{2}+\frac{m_{1}c}{a_{1}r_{1}q_1p_2}\). From Lemma 2, it follows that there exists at least one positive root \(\bar{x}\). Therefore, the positive bionomic equilibrium exists if (13) is satisfied. The proof is complete. \(\square \)
1.2 Proof of Theorem 9
Proof
By Pontryagin’s maximum principle (1987), the adjoint equations read
Consider the equilibrium of (B4), then the adjoint equations reduce to
Then
Combining (17) with (B9), one has
where
This leads to the solution
where \(\Xi _{1}\) and \(\Xi _{2}\) are constants of integration, \(\triangle _{1}\) and \(\triangle _{2}\) are the roots of the auxiliary equation \(\triangle ^{2}+\mathcal {Q}_{1}\triangle +\mathcal {Q}_{2}=0\). Imposing the transverse condition \(\chi _{2}(t)\rightarrow 0\) as \(t\rightarrow \infty \) leads to \(\Xi _{1}=\Xi _{2}=0\). Then,
Substituting \(\chi _2\) into (17) reaches
Similarly,
where
\(\chi _{1}\) can be simplified as
It is not difficult to find that the shadow prices \(\chi _{1}(t)\textrm{e}^{\delta _{u}t}\), \(\chi _{2}(t)\textrm{e}^{\delta _{u}t}\), and \(\chi _{3}(t)\textrm{e}^{\delta _{u}t}\) are bounded and constant over time in optimal equilibrium when they satisfy the transversality condition at \(\infty \) (Sadhukhan et al 2010).
Substituting the values of \(\chi _{2},\chi _{3}\) into (B8), one obtains
The optimal equilibrium solution \((x^{**},y^{**},z^{**},E^{**})\) of the control problem can be obtained by solving steady state equations together with (B12). Moreover, the maximum net present revenue is
The proof is complete. \(\square \)
Appendix C: Proofs for Stochastic System
1.1 Proof of Theorem 10
Proof
Since (2) has a unique local solution \((x(t),y(t),z(t))\in \mathbb {R}^{3}_{+}\) on \([0,\tau _{e})\) a.s., where \(\tau _{e}\) is the explosion time. In order to show that this solution is global, i.e., \(\tau _{e}=\infty \) a.s., let \(n_{0}\ge 0\) be sufficiently large such that \(x_{0}\), \(y_{0}\), and \(z_{0}\) are lying within the interval \([{1}/{n_{0}},n_{0}]\). For each integer \(n\ge n_{0}\), define the stopping time as
Obviously, \(\tau _{n}\) is increasing as \(n\rightarrow \infty \). Set \(\tau _{\infty }=\lim _{n\rightarrow \infty } \tau _{n}\), whence \(\tau _{\infty }\le \tau _{e}\) a.s. Hence, it only needs to prove that \(\tau _{\infty }=\infty \) a.s. It can be proved by contradiction arguments. Assume that the statement is not true, then there exist a pair of constants \(T>0\) and \(\kappa \in (0,1)\) such that \(\mathbb {P}\left\{ \tau _{\infty }\le T\right\} >\kappa .\) Whence there exists an integer \(n_{1}\ge n_{0}\) such that \(\mathbb {P}\left\{ \tau _{n}\le T\right\} \ge \kappa ,~n\ge n_{1}.\) Define a \(C^{2}\)-function V: \(\mathbb {R}^{3}_{+}\rightarrow \mathbb {R}_{+}\) by
with \(0<\theta <1\). The nonnegativity of V follows from \( u^{\theta }-\ln u-(1-\ln (1/\theta ))/\theta \ge 0,~u\ge 0. \) By applying Itô’s formula, one has
Then, by Lemma 3,
where
and \(\rho \) is a positive constant being independent of x, y, z, and t. Consequently,
Integrating it from 0 to \(\tau _{n}\wedge T\) and taking expectation produces
Set \(\Omega _{n}=\left\{ \omega \in \Omega : \tau _{n}=\tau _{n}(\omega )\le T\right\} \), then \(\mathbb {P}(\Omega _{n})\ge \kappa \), and, for any \(\omega \in \Omega _{n}\), \(x(\tau _{n},\omega )\), \(y(\tau _{n},\omega )\) or \(z(\tau _{n},\omega )\), equals either n or 1/n. Hence,
where \(I_{\Omega _{n}}\) denotes the indicator function of \(\Omega _{n}\). Let \(n\rightarrow \infty \), then there is a contradiction
Therefore, \(\tau _{\infty }=\infty \). The proof is complete. \(\square \)
1.2 Proof of Theorem 11
Proof
Define
By the generalized Itô’s formula, one has
where
For any \(0<\beta <d+h_2\), applying Itô’s formula to \(\textrm{e}^{\beta t}\tilde{V}\) leads to
where
On the one hand,
on the other side,
where the following H\(\mathrm {\ddot{o}}\)lder inequality has been used
Hence,
which is bounded, say by \(\mathcal {G}^{*}(\tilde{\theta })\). Then
Letting \(n\rightarrow \infty \) and then \(t\rightarrow \infty \) yields
It follows that
Set
then
The proof is complete. \(\square \)
1.3 Proof of Theorem 12
Proof
Note that, for any \((x_{0},y_{0},z_{0})\in \mathbb {R}^{3}_{+}\), the solution of (2) is positive, then
Consider the following auxiliary one-dimensional stochastic differential equation
with an initial value \(X(0)=x_{0}>0\). Set
Direct calculation shows that
Then
where
Letting
then
which implies that \(\displaystyle \int \limits _{0}^{1}\tilde{\phi }(x)\textrm{d}x\) has the same convergence as \(\displaystyle \int \limits _{0}^{1}\tilde{\phi }_{1}(x)\textrm{d}x\). Since
and \(r_{1}>\sigma _{11}^{2}/2\), one has \(\displaystyle \int \limits _{0}^{1}x^{2(r_{1}/\sigma _{11}^{2}-1)}\textrm{d}x<+\infty .\) On th
then
which implies that \(\displaystyle \int \limits _{1}^{\infty }\tilde{\phi }(x)\textrm{d}x\) has the same convergence as \(\displaystyle \int \limits _{1}^{\infty }\tilde{\phi }_{2}(x)\textrm{d}x\). Then
Whence
Due to the ergodic property (Kutoyants and Kutojanc 2004), the invariant density of (C13) is
where Q is a constant such that \(\int _{0}^{\infty }\varphi (x)\textrm{d}x=1\). From the ergodic theorem, it follows that
Let X(t) be the solution of (C13) with \(X(0)=x_{0}\), then, by the comparison theorem of one-dimensional stochastic differential equation (Peng and Zhu 2006), one has \(x(t)\le X(t)\) for \(t\ge 0\) a.s.
Applying the Itô’s formula yields
Integrating both sides of (C14) from 0 to t and then dividing it by t leads to
Let \(\tilde{M}(t)=\sigma _{22}\int _{0}^{t}y(s)\textrm{d}B_{2}(s)\), then \(\tilde{M}(t)\) is a local martingale whose quadratic variations is \(\langle \tilde{M},\tilde{M}\rangle _{t}=\sigma _{22}^{2}\int _{0}^{t}y^{2}(s)\textrm{d}s\). By employing the exponential martingales inequality (Mao 2007), for any positive constant \(\delta \), one obtains
From Borel-Cantelli Lemma (Mao 2007), it follows that, for almost all \(\omega \in \Omega \), there exists a \(\delta _{0}=\delta _{0}(\omega )\) such that
Then
Hence, for \(0\le \delta -1\le t<\delta \), the following inequality holds:
Taking the superior limit on both sides of (C16) and using \(\lim _{t\rightarrow \infty }{B_{2}(t)}/{t}=0\) a.s., by the strong law of large numbers for local martingale (Mao 2007), one gets
that is, \(\lim _{t\rightarrow \infty }y(t)=0\), and then \(\lim _{t\rightarrow \infty }z(t)=0\) a.s. As a result, for any \(\epsilon >0\), there exists an \(\Omega _{\epsilon }\subset \Omega \) such that
Note that
and
the distribution of the process x(t) converges to the measure with the density \(\varphi \). The proof is complete. \(\square \)
1.4 Proof of Theorem 13
Proof
Applying the Itô’s formula to \(\ln x\) produces
Integrating (C17) from 0 to t and dividing it by t on both sides leads to
Let \(M(t)=\sigma _{12}\int _{0}^{t}x(s)\textrm{d}B_{1}(s)\), then M(t) is a local martingale whose quadratic variations is \(\langle M,M\rangle _{t}=\sigma _{12}^{2}\int _{0}^{t}x^{2}(s)\textrm{d}s\). By employing the exponential martingales inequality (Mao 2007), for any positive constant \(\delta \), one obtains the following inequality
From Borel-Cantelli Lemma (Mao 2007), it follows that, for almost all \(\omega \in \Omega \), there exists \(\delta _{0}=\delta _{0}(\omega )\) such that
It is trivial to show that
Hence, for \(0\le \delta -1\le t<\delta \), the following inequality holds
Taking the superior limit on the both sides of (C19) and noting that \(\lim _{t\rightarrow \infty }{B_{1}(t)}/{t}=0\) a.s. the strong law of large numbers for local martingale (Mao 2007) implies that
then \(\lim _{t\rightarrow \infty }x(t)=0\) a.s. Therefore, there exists a set \(\Omega _{\epsilon }\subset \Omega \) such that
Applying the Itô’s formula to \(\ln y\), integrating it from 0 to t, and dividing it by t on both sides, one has
Taking the superior limit on both sides of (C20) and using the arbitrariness of \(\epsilon \) yield
which implies that \(\lim _{t\rightarrow \infty }y(t)=0\) and \(\lim _{t\rightarrow \infty }z(t)=0\) a.s. The proof is complete.
\(\square \)
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Zhao, X., Liu, L., Wang, H. et al. Ecological Effects of Predator Harvesting and Environmental Noises on Oceanic Coral Reefs. Bull Math Biol 85, 59 (2023). https://doi.org/10.1007/s11538-023-01166-z
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DOI: https://doi.org/10.1007/s11538-023-01166-z