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Predator-prey interactions under fear effect and multiple foraging strategies

  • * Corresponding author: Joydeb Bhattacharyya

    * Corresponding author: Joydeb Bhattacharyya 

SH is supported by CSIR, Govt. of India grant (09/106(0161)/2017-EMR-I). JB is supported by SERB, Govt. of India grant (TAR/2018/000283). SP is supported by WBSCST, Govt. of India grant (ST/P/S & T/16G-22/2018)

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  • We propose and analyze the effects of a generalist predator-driven fear effect on a prey population by considering a modified Leslie-Gower predator-prey model. We assume that the prey population suffers from reduced fecundity due to the fear of predators. We investigate the predator-prey dynamics by incorporating linear, Holling type Ⅱ and Holling type Ⅲ foraging strategies of the generalist predator. As a control strategy, we have considered density-dependent harvesting of the organisms in the system. We show that the systems with linear and Holling type Ⅲ foraging exhibit transcritical bifurcation, whereas the system with Holling type Ⅱ foraging has a much more complex dynamics with transcritical, saddle-node, and Hopf bifurcations. It is observed that the prey population in the system with Holling type Ⅲ foraging of the predator gets severely affected by the predation-driven fear effect in comparison with the same with linear and Holling type Ⅱ foraging rates of the predator. Our model simulation results show that an increase in the harvesting rate of the predator is a viable strategy in recovering the prey population.

    Mathematics Subject Classification: Primary: 92B05, 92D40; Secondary: 92D25.

    Citation:

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  • Figure 1.  Mutual position of prey-nullclines (red) and predator-nullclines (blue) of the system (3) due to the changes in $ h_1 $ and $ h_2 $, other parameters are taken from Table 2. The system is LAS at $ (a) $ $ E^* $ ($ E_0 $, $ E_1 $ and $ E_2 $ are unstable), $ (b) $ $ E_2 $ ($ E_0 $ and $ E_1 $ are unstable; $ E^* $ does not exist), $ (c) $ $ E_1 $ ($ E_0 $ is unstable; $ E^* $ and $ E_2 $ do not exist) and $ (d) $ $ E_0 $ ($ E^* $, $ E_1 $ and $ E_2 $ do not exist)

    Figure 2.  One-parameter bifurcation plots of the system $ (4) $ due to the changes in $ (a) $ $ h_1 $ where $ h_2<r $, $ (b) $ $ h_1 $ where $ h_2>r $, $ (c) $ $ h_2 $ where $ h_1<h_1^* $, and $ (d) $ $ h_2 $ where $ h_1>h_1^* $

    Figure 3.  Two-parameter bifurcation plots with the bifurcation parameters $ (a) $ $ h_1 $ and $ h_2 $, $ (b) $ $ h_1 $ and $ \beta $, $ (c) $ $ h_2 $ and $ \beta $, $ (d) $ $ \beta $ and $ \eta_1 $, $ (e) $ $ h_1 $ and $ \eta_1 $, $ (f) $ $ h_2 $ and $ \eta_1 $. All other parameters are taken from Table 2. The coloured bars represent the prey population density

    Figure 4.  Mutual position of prey-nullclines (red) and predator-nullclines (blue) of the system (4) due to the changes in $ h_1 $ and $ h_2 $, all other parameters are taken from Table 2. $ (a) $ The system is LAS at the unique interior equilibrium $ E^*_1 $ ($ E_0 $, $ E_1 $ and $ E_2 $ are unstable). $ (b) $ The system has bistability at $ E_2 $ and $ E^*_1 $ ($ E_0 $, $ E_1 $ and $ E^*_2 $ are unstable). The system is LAS at $ (c) $ $ E_2 $ ($ E_0 $ and $ E_1 $ are unstable; $ E^*_i $ does not exist), $ (d) $ $ E_2 $ ($ E_0 $ is unstable; $ E^*_i $ and $ E_1 $ do not exist), $ (e) $ $ E_1 $ ($ E_0 $ is unstable; $ E^*_i $ and $ E_2 $ do not exist) and $ (f) $ $ E_0 $ ($ E^*_i $, $ E_1 $ and $ E_2 $ do not exist) $ (i = 1,2) $

    Figure 5.  One-parameter bifurcation plots of the system (4) due to the changes in $ h_1 $ $ (a) $ for $ h_2<r $ and $ \eta_1 = 0.05 $, where a transcritical bifurcation occurs at $ h_1^{**} = 0.6585 $; $ (b) $ for $ h_2<r $ and $ \eta_1 = 0.125 $, a transcritical and a saddle-node bifurcation occur at $ h_1^{**} = 0.1475 $ and $ h_{1cr}^- = 0.3685 $ respectively. One-parameter bifurcation plots of the system (4) due to the changes in $ h_2 $ $ (c) $ for $ h_1<h_1^* $ and $ \eta_1 = 0.05 $, where a transcritical bifurcation occurs at $ h_2^{**} = 0.3495 $; $ (d) $ for $ h_1<h_1^* $ and $ \eta_1 = 0.125 $, a transcritical and a saddle-node bifurcation occur at $ h_2^{**} = 0.74 $ and $ h_{2sn} = 0.64 $ respectively

    Figure 6.  Two-parameter bifurcation plots with $ h_1 $ and $ h_2 $ as bifurcation parameters where $ (a) $ $ \eta_1 = 0.05 $ and $ (c) $ $ \eta_1 = 0.25 $, where $ f_{SN} = 0 $ is a saddle-node bifurcation curve, $ h_i = h_i^{**} $ $ (i = 1,2) $ and $ h_2 = r $ are transcritical bifurcation curves

    Figure 7.  Two-parameter bifurcation plots with the bifurcation parameters $ (a) $ $ h_1 $ and $ \beta $, $ (b) $ $ h_2 $ and $ \beta $, $ (c) $ $ h_1 $, and $ \eta_1 $, $ (d) $ $ h_2 $ and $ \eta_1 $; other parameter values are taken from Table 2

    Figure 8.  $ (a) $ For $ \alpha = 0.4993 $, the phase space shows the existence of stable (in green) and unstable (in red) manifolds of the system (4). The unstable limit cycle around $ E^*_1 $ is represented in blue. The curves representing the changes in $ (b) $ $ {\rm{Tr}}(J^*_1) $, $ {\rm{Det}}(J^*_1) $ and $ (c) $ $ \frac{d}{d\alpha}{\rm{Tr}}(J^*_1) $ due to the changes in $ \alpha $ verify the occurrence of a Hopf bifurcation of the system (4) at $ \alpha = 0.4993 $

    Figure 9.  Mutual position of prey-nullclines (red) and predator-nullclines (blue) of the system (5) due to the changes in $ h_1 $ and $ h_2 $, other parameters are taken from Table $ 1 $. The system is LAS at $ (a) $ $ E_* $ ($ E_0 $, $ E_1 $ and $ E_2 $ are unstable), $ (b) $ $ E_2 $ ($ E_0 $ and $ E_1 $ are unstable; $ E_* $ does not exist), $ (c) $ $ E_1 $ ($ E_0 $ is unstable; $ E_* $ and $ E_2 $ do not exist) and $ (d) $ $ E_0 $ ($ E_* $, $ E_1 $ and $ E_2 $ do not exist)

    Figure 10.  One-parameter bifurcation plots of the system (5) due to the changes in $ (a) $ $ h_1 $ where $ h_2<r $, $ (b) $ $ h_2 $ where $ h_1<h_1^{\#} $. $ (c) $ A two-parameter bifurcation plot with $ h_1 $ and $ h_2 $ as bifurcation parameters, where $ h_1 = 1 $, $ h_1 = h_1^{\#} $ and $ h_2 = r $ are transcritical bifurcation curves. All other parameters are taken from Table 2

    Figure 11.  Two-parameter bifurcation plots of the system (5) with the bifurcation parameters $ (a) $ $ h_1 $ and $ \beta $, $ (b) $ $ h_2 $ and $ \beta $, $ (c) $ $ h_1 $ and $ \eta_1 $, $ (d) $ $ h_2 $ and $ \eta_1 $, where $ h_1 = h_1^{\#} $, $ h_2 = h_2^{\#} $, and $ h_2 = r $ are transcritical bifurcation curves

    Figure 12.  Local sensitivity of the prey response for $ (a) $ linear, $ (b) $ Holling type Ⅱ, and $ (c) $ Holling type Ⅲ foraging of the predator. For comparison, model simulations before parameter manipulations, are shown in black line. The prey density from simulations where particular parameter values were increased by $ 10\% $ are shown in red lines, as are the prey density from simulations where particular parameter values were decreased by $ 10\% $ in blue lines

    Figure 13.  One-parameter bifurcation plots due to the changes in $ \beta $, where $ h_1<1 $ and $ h_2<r $ for $ (a) $ linear, $ (b) $ Holling type Ⅱ, and $ (c) $ Holling type Ⅲ foraging rates

    Figure 14.  One-parameter bifurcation plots due to the changes in $ \eta_1 $, where $ h_1<1 $ and $ h_2<r $ for $ (a) $ linear, $ (b) $ Holling type Ⅱ, and $ (c) $ Holling type Ⅲ foraging rates

    Figure 15.  Local sensitivity of the predator response for $ (a) $ linear, $ (b) $ Holling type Ⅱ, and $ (c) $ Holling type Ⅲ foraging of the predator. For comparison, model simulations before parameter manipulations, are shown in black line. The predator density from simulations where particular parameter values were increased by $ 10\% $ are shown in red lines, as are the predator density from simulations where particular parameter values were decreased by $ 10\% $ in blue lines

    Table 1.  Non-dimensionalized system

    $ \delta $ $ \delta=1 $ (Linear) $ \delta=2 $ (Holling type-Ⅱ) $ \delta=3 $ (Holling type-Ⅲ)
    Transformation $X=Kx$, $Y= \frac{R_{1}}{A_{1}}y$, $T= \frac{1}{R_{1}}t$,
    $r=\frac{R_{2}}{R_{1}}$, $\beta=\frac{BR_{1}}{A_{1}}$, $h_{1}=\frac{H_{1}}{R_{1}}$,
    $h_{2}=\frac{H_{2}}{R_{1}}$, $\alpha=\frac{A_{2}}{A_{1}K}$, $\eta_{1}=\frac{\eta}{K}$
    $X=Kx$, $Y= \frac{R_{1}K}{A_{1}}y$, $T= \frac{1}{R_{1}}t$,
    $r=\frac{R_{2}}{R_{1}}$, $\beta=\frac{BR_{1}K}{A_{1}}$, $h_{1}=\frac{H_{1}}{R_{1}}$, $h_{2}=\frac{H_{2}}{R_{1}}$,
    $\alpha=\frac{A_{2}}{A_{1}}$, $\eta_{1}=\frac{\eta}{K}$, $b=\frac{B_{1}}{K}.$
    $X=Kx$, $Y= \frac{R_{1}K}{A_{1}}y$, $T= \frac{1}{R_{1}}t$,
    $r=\frac{R_{2}}{R_{1}}$, $\beta=\frac{BR_{1}K}{A_{1}}$, $h_{1}=\frac{H_{1}}{R_{1}}$, $h_{2}=\frac{H_{2}}{R_{1}}$,
    $\alpha=\frac{A_{2}R_{1}}{A_{1}}$, $\eta_{1}=\frac{\eta}{K}$, $b_{1}=\frac{B_{1}}{K^2}.$
    Non-dimensional system $ \begin{array}{ll}\frac{dx}{dt}=\frac{x(1-x)}{1+\beta y}- xyc_\delta(x)-h_{1}x\equiv f^\delta_1 \; \;\;\;\;\;\;\;\;\;(2)\end{array}$
    $ \frac{dy}{dt}=y\left(r-\frac{\alpha y}{x+\eta_{1}}\right)-h_{2}y\equiv f^\delta_2,$
    where $c_{\delta}(x)=\left\{\begin{array}{ll} 1, &{ \textrm{if }}\;\delta=1\\ \frac{1}{b+x}, &{ \textrm{if }}\;\delta=2\\ \frac{x}{b_1+x^2}, &{ \textrm{if }}\;\delta=3, \end{array}\right.$
    $x(0)\geq 0$ and $y(0)\geq 0$.
     | Show Table
    DownLoad: CSV

    Table 2.  Tables of parameter values

    (a)
    Original parameters
    Parameter Description Value
    $R_1$ Intrinsic growth rate of prey 0.03
    $R_2$ Intrinsic growth rate of predator 0.03
    $B$ The level of fear 4
    $A_1$ Consumption rate of predator 0.5
    $A_2$ Intraspecific competition of predator 0.5
    $K$ Carrying capacity of prey 2
    $\eta$ Alternative prey density 0.25
    $B_1$ Half saturation coefficient 0.1
    $H_1$ Harvesting rate of prey 0.01
    $H_2$ Harvesting rate of predator 0.02
    (b)
    Non-dimensional parameters
    Parameter Value
    $r$ 1
    $\alpha$ 0.03
    $\beta$ 0.48
    $\eta_1$ 0.125
    $b$ 0.025
    $h_1$ 0.333
    $h_2$ 0.667
     | Show Table
    DownLoad: CSV

    Table 3.  Existence and local stability of equilibria of system (3)

    Equilibria Sufficient condition for existence Local asymptotic stability
    $ E_0 $ Always $ h_1>1 $ and $ h_2>r $
    $ E_1 $ $ h_1<1 $ $ h_{1}<1 $ and $ h_{2}>r $
    $ E_2 $ $ h_2<r $ $ h_{1}>h_{1}^* $ and $ h_{2}<r $
    $ E^* $ $ h_1<\min\{1,h_{1}^*\} $ and $ h_2<r $ $ h_1<\min\{1,h_{1}^*\} $ and $ h_2<r $
     | Show Table
    DownLoad: CSV

    Table 4.  Existence and local stability of equilibria of system (4)

    Equilibria Sufficient condition for existence Local asymptotic stability
    $ E_0 $ Always $ h_1>1 $ and $ h_2>r $
    $ E_1 $ $ h_1<1 $ $ h_{1}<1 $ and $ h_{2}>r $
    $ E_2 $ $ h_2<r $ $ h_{1}>h_{1}^{**} $ and $ h_{2}<r $
    $ E^*_i $ $ h_1<\min\{1,h_{1}^{**}\} $ and $ h_2<r $ $ \mbox{Tr}(J^*_i)<0 $ and $ \mbox{Det}(J^*_i)>0 $
     | Show Table
    DownLoad: CSV

    Table 5.  Existence and local stability of equilibria of system (5)

    Equilibria Sufficient condition for existence Local asymptotic stability
    $ E_0 $ Always $ h_1>1 $ and $ h_2>r $
    $ E_1 $ $ h_1<1 $ $ h_{1}<1 $ and $ h_{2}>r $
    $ E_2 $ $ h_2<r $ $ h_{1}>h_1^{\#} $ and $ h_{2}<r $
    $ E_* $ $ h_1<\min\{1,h_1^{\#}\} $ and $ h_2<r $ $ \mbox{Tr}(J_*)<0 $ and $ \mbox{Det}(J_*)>0 $
     | Show Table
    DownLoad: CSV

    Table 6.  Comparison of the critical threshold values for transcritical bifurcation (TB) and saddle-node bifurcation (SNB) of the three systems

    Bifurcation parameter Linear Holling type Ⅱ Holling type Ⅲ
    Threshold Bifurcation Threshold Bifurcation Threshold Bifurcation
    $h_1$
    ($h_2 < r$)
    $h_1^*=0.8971$ TB $h_1^{**}=0.1475$
    $h_{1sn}^-=0.3685$
    TB
    SNB
    $h_1^{\#}=0.79$ TB
    $h_1$
    ($h_2>r$)
    $h_1^*=1$ TB $h_1^{**}=1$ TB $h_1^{\#}=1$ TB
    $h_2$
    ($h_1 < 1$)
    $h_2^*=1$ TB $h_2^{**}=0.74$
    $h_{2sn}=0.64$
    TB
    SNB
    $h_2^{\#}=1$ TB
    $\beta$
    ($h_1 < 1$ & $h_2 < r$)
    $\beta^*=16.8$ TB $\beta^{**}=16$
    $\beta_{sn}=18.51$
    TB
    SNB
    $\beta^{\#}=1.5$ TB
    $\eta_1$
    ($h_1 < 1$ & $h_2 < r$)
    $\eta_1^*=0.825$ TB $\eta_1^{**}=0.2$
    $\eta_{1sn}=0.441$
    TB
    SNB
    $\eta_1^{\#}=0.3721$ TB
     | Show Table
    DownLoad: CSV

    Table 7.  Bifurcation parameters with different foraging types and corresponding basins of attraction at $ E^* $

    Parameters Largest basin of recovery Smallest basin of recovery
    $ h_1 $ & $ h_2 $ Linear Holling type Ⅱ
    $ h_1 $ & $ \beta $ Linear Holling type Ⅲ
    $ h_2 $ & $ \beta $ Holling type Ⅲ Holling type Ⅱ
    $ h_1 $ & $ \eta_1 $ Linear Holling type Ⅱ
    $ h_2 $ & $ \eta_1 $ Linear Holling type Ⅱ
     | Show Table
    DownLoad: CSV
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