Bifurcations, chaos, and multistability in a

nonautonomous predator–prey model with

fear
Cite as: Chaos 31, 123134 (2021); https://doi.org/10.1063/5.0067046
Submitted: 13 August 2021 • Accepted: 06 December 2021 • Published Online: 30 December 2021

Mainul Hossain, Saheb Pal, Pankaj Kumar Tiwari, et al.

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Chaos 31, 123134 (2021); https://doi.org/10.1063/5.0067046 31, 123134

© 2021 Author(s).
 Chaos ARTICLE scitation.org/journal/cha

Bifurcations, chaos, and multistability in a

nonautonomous predator–prey model with fear
Cite as: Chaos 31, 123134 (2021); doi: 10.1063/5.0067046
Submitted: 13 August 2021 · Accepted: 6 December 2021 ·
Published Online: 30 December 2021 View Online Export Citation CrossMark

Mainul Hossain,1 Saheb Pal,1 Pankaj Kumar Tiwari,2 and Nikhil Pal1,a)

AFFILIATIONS
1
Department of Mathematics, Visva-Bharati, Santiniketan 731235, India
2
Department of Basic Science and Humanities, Indian Institute of Information Technology, Bhagalpur 813210, India

a)
Author to whom correspondence should be addressed: nikhil.pal@visva-bharati.ac.in

ABSTRACT
Classical predator–prey models usually emphasize direct predation as the primary means of interaction between predators and prey.
However, several field studies and experiments suggest that the mere presence of predators nearby can reduce prey density by forcing them to
adopt costly defensive strategies. Adoption of such kind would cause a substantial change in prey demography. The present paper investigates
a predator–prey model in which the predator’s consumption rate (described by a functional response) is affected by both prey and predator
densities. Perceived fear of predators leads to a drop in prey’s birth rate. We also consider both constant and time-varying (seasonal) forms
of prey’s birth rate and investigate the model system’s respective autonomous and nonautonomous implementations. Our analytical studies
include finding conditions for the local stability of equilibrium points, the existence, direction of Hopf bifurcation, etc. Numerical illustra-
tions include bifurcation diagrams assisted by phase portraits, construction of isospike and Lyapunov exponent diagrams in bi-parametric
space that reveal the rich and complex dynamics embedded in the system. We observe different organized periodic structures within the
chaotic regime, multistability between multiple pairs of coexisting attractors with intriguing basins of attractions. Our results show that even
relatively slight changes in system parameters, perturbations, or environmental fluctuations may have drastic consequences on population
oscillations. Our observations indicate that the fear effect alters the system dynamics significantly and drives an otherwise irregular system
toward regularity.
Published under an exclusive license by AIP Publishing. https://doi.org/10.1063/5.0067046

Classical predator–prey models have revolved primarily around
direct predation. However, in recent years, the indirect effects
of predators on their prey have gained considerable attention
among ecological researchers. Several field studies have been
carried out to assess the impact of prey’s perceived fear of
being killed by its predators as opposed to just direct preda-
tion. Furthermore, theoretical studies have revealed many aspects
of the role of fear effect on the stability and coexistence across
a wide range of ecological contexts. Inspired by recent works
on ecological systems incorporating fear effect, we investigate
a nonautonomous predator–prey model with a time-varying
birth rate and direct predation being governed by Cosner func-
tional response. Here, fear is introduced as a decreasing factor
to prey’s birth rate. We explore both autonomous and nonau-
tonomous implementations of our model system. This includes
finding the conditions for the local stability of equilibrium points,
examining the existence and direction of Hopf bifurcation, etc.,
analytically. With the help of advanced numerical tools and tech-
niques, we explore the role of different system parameters by
using bifurcation diagrams assisted by phase portraits. We also
construct several isospike and Lyapunov exponent diagrams in
the bi-parametric space of fear and birth rate parameters. In this
way, we illustrate the intricate and rich dynamics woven into our
system. Our observations include the presence of different orga-
nized periodic structures (shrimp-shaped periodic islands and
Arnold tongues) within the chaotic regime. We also detect multi-
stability between several pairs of coexisting attractors. It appears
from their basins of attractions that they are of fractal nature. Our
results suggest that the parameter values and the initial popula-
tion sizes of prey and predator play a vital role in the dynamics
of the system; a slight alteration in either of these can lead to a
drastic change in the outcome. We observe that the fear effect
has a stabilizing role on the autonomous system, whereas in the
nonautonomous system, the fear effect drives the system toward

Chaos 31, 123134 (2021); doi: 10.1063/5.0067046 31, 123134-1

Published under an exclusive license by AIP Publishing
 Chaos
regularity. In summary, we conclude that the nonautonomous
ecological systems are inherently complex and exhibit intricate
dynamical behaviors.
I. BIOLOGICAL BACKGROUND AND MODEL
FORMULATION
Evolution in organisms is shaped by interspecies interactions.
Among them, predator–prey interactions are a widely studied phe-
nomena in the scientific community. At its core, the interaction
is between an organism and its natural enemies. Plant–herbivore,
host–parasitoid, herbivore–carnivore, and host–pathogen interac-
tions are a few examples of prey–predator interactions. Recent
trends in population biology are to develop several mathemati-
cal models and study the long-term behavior of the species, their
coexistence, and how they impact each other’s densities.1,2 Two-
species mathematical models look simple; however, they exhibit rich
and complex dynamics. A prototypical two-species predator–prey
model introducing x(t) and y(t) as the prey and predator population
densities at time t, respectively, can be written as
( dx
dt
= bx − dx − ax2 − 8(x, y)y,
dy
(1.1)
dt
= µ8(x, y)y − my,
where b and d are the birth and death rates of prey, respectively,
a denotes the intraspecific competition rate of prey, µ represents
the conversion factor of prey biomass into predator biomass, 8(x, y)
describes the prey consumption rate of individual predator depend-
ing on both the predator and prey densities, and m is the predator
mortality rate. Functional response is a basic modeling unit in
a predator–prey system. It describes the intake rate at which a
predator consumes prey. The behavioral characteristic of predator
species reflects through the term 8(x, y), and predation is typically
described by the functional response.
The most widely used functional responses in describing preda-
tor–prey interactions are of Holling types,3 which depend on prey
density only. However, such prey-dependent functional responses
do not consider the interference among predators.4,5 Some biolo-
gists have claimed that in many situations, especially when predators
have to search for food (and therefore, have to share or compete
for food), the functional response in a predator–prey model should
be predator-dependent.6–9 However, the appropriate form of func-
tional response has been debated. For a better understanding of
the proper representation of predation processes, detailed observa-
tions of environmental factors that influence functional response
are required. Cosner et al.10 proposed a functional response that is
predator-dependent and describes the schooling behavior of both
the predator and prey populations. A biological example regarding
this type of behavior is that a school of tuna searches, contacts, and
then hunts a herd of prey.11 The functional form of such response
function is given by
Ce0 xy
8(x, y) = , (1.2)
1 + hCe0 xy
where C is the amount of prey killed by a predator per encounter,
e0 is the total encounter coefficient between the predator and the
Chaos 31, 123134 (2021); doi: 10.1063/5.0067046
Published under an exclusive license by AIP Publishing
ARTICLE scitation.org/journal/cha
prey, and h is the handling time per prey. Unlike the other predator-
dependent functional responses (such as ratio-dependent or Bed-
dington–DeAngelis), the functional response (1.2) is increasing in
predator density y since it presumed that predators were foraging in
a group. For the convenience of mathematical analysis, we consider
α as the product of C and e0 in (1.2). Then, the model system (1.1)
can be written in the form

αxy2
 dx

dt
= bx − dx − ax2 − ,
 1 + hαxy
(1.3)
µαxy2
 dy =


dt
− my.
1 + hαxy
Recent empirical and theoretical studies suggested that preda-
tor has both direct and indirect impacts (consumptive and non-
consumptive) upon their prey. In a direct way, the predator kills and
consumes prey, but in an indirect way, the predator induces fear in
the prey population and changes the prey’s behavior. Due to fear
of predation, the prey population enhances vigilance, limits forag-
ing activity, sacrifices higher intake zone and feeds in a safer place,
adjust reproductive cycle, etc.12–16 These adjustments create trade-
offs such that prey will enhance their anti-predator behavior and
limit their foraging intensity or reproductive behavior. Therefore,
prey compensates for mortality risk, which ultimately affects the
growth rate of prey species. Several researchers empirically observed
such types of phenomena in different predator–prey systems.17,18
Here, we assume that due to fear of predation risk, the birth rate
of the prey population is reduced. To incorporate the fear phe-
nomenon, we multiply the birth rate (b) of prey population with a
decreasing function of the predator density (y) and the intensity of
fear (k). k measures the level of fear exerted by a particular predator
on a particular prey. It may be understood this way: if the predators
are very aggressive and fearsome, k is higher, and if the predators are
less fearsome, k assumes a lesser value. Although it may vary from
species to species depending on many biological factors (age, terrain,
etc.), for simplicity, we assume it to remain constant over time for all
age groups of the prey and predator. Now, from a biological point of
view, the fear function 2(k, y) must have the following properties:
1. 2(0, y) = 1 : if there is no fear of predator, then the birth rate of
prey remains unchanged.
2. 2(k, 0) = 1 : if there is no predator in the system, then there is
no reduction in prey reproduction due to fear.
3. limk→∞ 2(k, y) = 0 : when strength of fear is very large and
predator density y > 0, then prey reproduction decreases and
ultimately becomes zero.
4. limy→∞ 2(k, y) = 0 : when predator density is very large and
strength of fear k > 0, then prey reproduction decreases and
ultimately becomes zero due to large strength of fear.
5. ∂2(k,y)
∂k
< 0 : the reproduction of prey decreases with an increase
in the strength of fear.
6. ∂2(k,y)
∂y
< 0 : the reproduction of prey decreases with an increase
in predator density.
The simplest form of 2(k, y) for which the above properties
1 19–23
hold well is given by 1+ky .
31, 123134-2
 Chaos ARTICLE scitation.org/journal/cha

Thus, the model (1.3) with a fear phenomenon becomes

bx αxy2

dx 2
 dt = 1 + ky − dx − ax − 1 + hαxy ,


(1.4)
µαxy2
 dy =


dt
− my.
1 + hαxy
All the parameters involved in the system (1.4) are non-
negative. We investigate the behaviors of the above model both
analytically and numerically.

II. MODEL ANALYSIS

Mathematical analysis of the model gives important informa-
tion about the steady state (where the system does not change with
time), coexistence, and extinction condition of each species. The
predator–prey model (1.4) always exhibits the trivial equilibrium
E0 = (0, 0), where both the species extinct

from the system. Also, the
predator-free equilibrium Ek = b−d a
, 0 exists for the model when
b > d.
For the coexistence equilibrium, where both the prey and
predator populations survive, we have to obtain the positive solu- FIG. 1. Figure shows non-trivial nullclines of the system (1.4) for k = 0.2,
tion(s) of the following system of algebraic equations: k = 0.7132, and k = 1.2. Other parameter values are b = 1.4, d = 0.3,
a = 0.4, α = 0.2, h = 2, µ = 0.7, and m = 0.1. Here, prey and predator
(
b αy2 nullclines are represented by red and green curves, respectively.
1+ky
− d − ax − 1+hαxy = 0,
µαxy
(2.1)
1+hαxy
− m = 0.
value of k. It is to be noted that the predator nullcline is independent
From the second equation of (2.1), we have
of the fear level.
(µ − mh)αxy − m = 0, Now, to determine the stability properties of coexistence equi-
librium Ē = (x̄, ȳ), we have to check the signs of trace and deter-
which demonstrates that model (1.4) has no coexistence equilibrium minant values of the corresponding variational matrix LĒ . The
for µ ≤ mh. From the ecological point of view, both the populations algebraic expression of LĒ is given by
can persist in the system if the conversion factor from prey biomass  
to predator biomass is sufficiently large compared to the product L L12
LĒ = 11 ,
of handling time and mortality rate of predators. Thus, we consider L21 L22
µ > mh, and the prey density of coexistence equilibrium is given by
m
x̄ = (µ−mh)αȳ , where the predator density ȳ is a positive root of the where
following biquadratic equation: hα 2 x̄ȳ3 kbx̄ α x̄ȳ(2 + hα x̄ȳ)
L11 = −ax̄ + , L12 = − − ,
ρ0 y4 + ρ1 y3 + ρ2 y2 + ρ3 y + ρ4 = 0, (1 + hα x̄ȳ)2 (1 + kȳ)2 (1 + hα x̄ȳ)2
µαȳ2 µα x̄ȳ
with L21 = 2
, L22 = .
(1 + hα x̄ȳ) (1 + hα x̄ȳ)2
2 2 2 2
ρ0 = kα (µ − mh) , ρ1 = α (µ − mh) ,
The trace (T) and determinant (D) values of the above variational
ρ2 = µdkα(µ − mh), matrix are given by
ρ3 = µamk + µα(µ − mh)(d − b), ρ4 = µam. T(x̄, ȳ) = L11 (x̄, ȳ) + L22 (x̄, ȳ), (2.2)
Note that all the coefficients are always positive for µ > mh
except ρ3 . A necessary (but not sufficient) condition for ρ3 to be neg- D(x̄, ȳ) = L11 (x̄, ȳ)L22 (x̄, ȳ) − L12 (x̄, ȳ)L21 (x̄, ȳ). (2.3)
ative is that b > d. The above biquadratic polynomial equation in y
has at most two positive roots according to Descartes’ rule of signs. Thus, the coexistence equilibrium Ē = (x̄, ȳ) is locally asymptotically
Thus, the model (1.4) exhibits at most two coexistence equilibria. In stable (LAS) if T(x̄, ȳ) < 0 and D(x̄, ȳ) > 0.
Fig. 1, we draw non-trivial prey and predator nullclines for different The local stability of extinction and predator-free equilibria is
values of k to visualize the existence of interior equilibrium points, straightforward, as the eigenvalues of the corresponding variational
where all other parameter values are fixed. We observe that below matrix are located along the diagonal. In Table I, we summarize all
a threshold value of k, both prey and predator nullclines intersect the steady states, their feasibility, and local stability of the model
at exactly two points; the nullclines do not cross above a threshold system (1.4).

Chaos 31, 123134 (2021); doi: 10.1063/5.0067046 31, 123134-3

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 Chaos ARTICLE scitation.org/journal/cha

TABLE I. Existence conditions and stability criteria of different steady states of the For the existence of a pair of purely imaginary roots of the
model (1.4). above equation, we must have D(x̄(k),
p ȳ(k)) > 0 at k = kh . Thus, we
consider λ1,2 = ±iν0 , where ν0 = D(x̄(k), ȳ(k))|k=kh .
Steady state Feasibility Stability Now, we check the transversality condition that confirms that
the eigenvalues cross the imaginary axis of the complex plane with
E0 = (0, 0)
Always exists LAS if b < d
non-zero speed. For this, let at any neighboring point k of kh , the
Ek = b−d ,0 b>d Always LAS if it
a eigenvalues of the variational matrix are λ1,2 = ξ(k) ± iν(k), where
exists
Coexistence state µ > mh and ρ 3 < 0 LAS if T(x̄, ȳ) < 0 T(x̄(k), ȳ(k))
Ē = (x̄, ȳ) and D(x̄, ȳ) > 0 ξ(k) =
2
and
r
In Fig. 2, we plot the densities of the prey population at equi- T2 (x̄(k), ȳ(k))
librium by changing the parameter k with two different values of ν(k) = D(x̄(k), ȳ(k)) − .
4
b(= 0.9/1.4), where other parameter values are fixed at d = 0.3,
a = 0.4, α = 0.2, h = 2, µ = 0.7, and m = 0.1. Note that the Now,
system has at most two coexisting (positive) equilibrium points.   
d  1 d
The equilibrium point that moves along the magenta curve always ξ(k)
 = T(x̄(k), ȳ(k)) .
exhibits saddle behavior. In the present work, the dynamics of the dk k=kh 2 dk k=kh
system (1.4) are explored with the help of numerical simulations
d
around the equilibrium point, where the equilibrium point moves Thus, the transversality condition is satisfied if dk T(x̄(k), ȳ(k))
along the green curve. A nice animation is also made available on the 6= 0 at k = kh . Therefore, the system (1.4) undergoes Hopf bifurca-
electronic version of this article (see the additional high-resolution tion at k = kh . 
movies in the supplementary material), which explains the existence
and stability of the equilibrium points. B. Direction of the Hopf bifurcation
At this point, we analyze the stability properties of the bifurcat-
A. Hopf bifurcation ing periodic solutions originating from the coexisting equilibrium
Nonlinear mathematical models of interacting populations point Ē = (x̄, ȳ) via the Hopf bifurcation. For this, we calculate the
show rich and complex dynamical behaviors even when system 1st Lyapunov coefficient by obtaining the normal form of the Hopf
complexity is low (two or three species). Oscillating behavior is the bifurcation. As mentioned earlier, at k = kh , the variational matrix
most frequent dynamical property in population dynamics. Oscil- LĒ has eigenvalues λ1,2 = ±iν0 .
lating behavior or the existence of a limit cycle leads to the Hopf Now, to obtain the normal form, first, we perform a linear
bifurcation of the system. The Hopf bifurcation is defined as the coordinate shift by introducing two new variables w1 and w2 ,
appearance or disappearance of a periodic orbit through a local
change in the stability properties of an equilibrium point. Here, we w1 = x − x̄, w2 = y − ȳ.
explore the possibility of occurrence of the Hopf bifurcation and
Then, the equilibrium point Ē = (x̄, ȳ) of the system (1.4)
its direction around an arbitrary coexistence equilibrium Ē = (x̄, ȳ)
transforms into the origin, and the system (1.4) becomes
with respect to bifurcating parameter k.
The following theorem gives the necessary and sufficient con- 
b(w1 + x̄)
ditions for Hopf bifurcation of the system (1.4).  dw

dt
1
= − d(w1 + x̄) − a(w1 + x̄)2



 1 + k(w2 + ȳ)
Theorem II.1. The necessary and sufficient conditions for which α(w1 + x̄)(w2 + ȳ)2


the system (1.4) undergoes the Hopf bifurcation at k = kh around − ,
 1 + hα(w1 + x̄)(w2 + ȳ)
the coexistence equilibrium point Ē are that D(x̄(k), ȳ(k)) > 0 and

 2
 dw2 = µα(w1 + x̄)(w2 + ȳ) − m(w + ȳ).

d

T(x̄(k), ȳ(k)) 6= 0 at k = kh .

dk  dt 2
1 + hα(w1 + x̄)(w2 + ȳ)
Proof. The characteristic equation of the variational matrix LĒ
is given by Expanding Taylor’s series of the above system at (w1 , w2 )
= (0, 0) up to terms of order 3 produces the following system:
λ2 − T(x̄(k), ȳ(k))λ + D(x̄(k), ȳ(k)) = 0,
ẇ1 = e10 w1 + e01 w2 + e20 w21 + e11 w1 w2

where T and D are trace and determinant values as given in (2.2)


+e02 w22 + e30 w31 + e21 w21 w2 + e12 w1 w22


and (2.3), respectively. For the Hopf bifurcation, the matrix LĒ must



+e03 w32 + O(|z|4 ),

have a pair of purely imaginary eigenvalues. At a critical value

(2.4)
k = kh , T(x̄(k), ȳ(k)) = 0. Therefore, the above characteristic 2
ẇ2 = f10 w1 + f01 w2 + f20 w1 + f11 w1 w2

equation becomes 


 +f02 w22 + f30 w31 + f21 w21 w2 + f12 w1 w22


λ2 + D(x̄(k), ȳ(k))|k=kh = 0. +f03 w32 + O(|z|4 ),


Chaos 31, 123134 (2021); doi: 10.1063/5.0067046 31, 123134-4

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 Chaos ARTICLE scitation.org/journal/cha

FIG. 2. Figure shows the variations of

coexistence equilibrium densities of the
prey population of the system (1.4) with
respect to the fear level k. In the left figure,
b = 0.9, and in the right figure, b = 1.4,
and the remaining parameter values
are d = 0.3, a = 0.4, α = 0.2, h = 2,
µ = 0.7, and m = 0.1.

where the upper dot denotes the derivative with respect to time t, where W = (w1 , w2 )T and
and

e20 w21 + e11 w1 w2 + e02 w22 + e30 w31

 
hα 2 ȳ3
 
e10 e01  
 +e21 w21 w2 + e12 w1 w22 + e03 w32 
= LĒ , e20 = −a + , M1
f10 f01 (1 + hα x̄ȳ)3 M=  f20 w2 + f11 w1 w2 + f02 w2 + f30 w3  .
= 
M2 1 2 1
bk 2αȳ +f21 w21 w2 + f12 w1 w22 + f03 w32
e11 = − 2
− ,
(1 + kȳ) (1 + hα x̄ȳ)3
bk2 x̄ α x̄ The eigenvector v̄ of variational matrix LĒ corresponding to the
e02 = − ,
(1 + kȳ)3 (1 + hα x̄ȳ)3 eigenvalue iν0 at k = kh is v̄ = (e01 , iν0 − e10 )T . Now, define
h2 α 3 ȳ4 3hα 2 ȳ2
e30 = − 4
, e21 = ,  
(1 + hα x̄ȳ) (1 + hα x̄ȳ)4 R = (Re(v̄), −Im(v̄)) =
e01 0
.
−e10 −ν0
bk2 α(1 − 2hα x̄ȳ)
e12 = 3
− ,
(1 + kȳ) (1 + hα x̄ȳ)4
Let W = RV or V = R−1 W, where V = (v1 , v2 )T . Under this
bk3 x̄ hα 2 x̄2
e03 = − + , transformation, system (2.5) becomes
(1 + kȳ)4 (1 + hα x̄ȳ)4
µhα 2 ȳ3 2µαȳ V̇ = (R−1 LĒ R)V + R−1 M(RV).
f20 = − , f11 = ,
(1 + hα x̄ȳ)3 (1 + hα x̄ȳ)3
µα x̄ µh2 α 3 ȳ4 This can be written as
f02 = 3
, f30 = ,
(1 + hα x̄ȳ) (1 + hα x̄ȳ)4
3µhα 2 ȳ2
      1 
µα(1 − 2hα x̄ȳ) v˙1 0 −ν0 v1 F (v1 , v2 ; k = kh )
f21 = − , f12 = , = + 2 , (2.6)
(1 + hα x̄ȳ)4 (1 + hα x̄ȳ)4 v˙2 ν0 0 v2 F (v1 , v2 ; k = kh )
µhα 2 x̄2
f03 = − .
(1 + hα x̄ȳ)4 where F1 and F2 are nonlinear in v1 and v2 and are given by

Neglecting the higher-order terms of degree 4 and above, the 1

F1 (v1 , v2 ; k = kh ) = M1 ,
system (2.4) can be written in the vector form as e01
1
F2 (v1 , v2 ; k = kh ) = − (e10 M1 + e01 M2 ),
Ẇ = LĒ W + M(W), (2.5) ν0 e01

Chaos 31, 123134 (2021); doi: 10.1063/5.0067046 31, 123134-5

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 Chaos ARTICLE scitation.org/journal/cha

with
M1 = (e20 e201 − e11 e01 e10 + e02 e210 )v21 + ν0 (2e02 e10 − e11 e01 )v1 v2
+ ν02 e02 v22 + (e30 e301 − e21 e201 e10 + e12 e01 e210 − e03 e310 )v31
+ ν0 (2e12 e10 e01 − e21 e201 − 3e03 e210 )v21 v2
+ ν02 (e12 e01 − 3e03 e10 )v1 v22 − ν03 e03 v32 ,
M2 = (f20 e201 − f11 e01 e10 + f02 e210 )v21 + ν0 (2f02 e10 − f11 e01 )v1 v2
+ ν02 f02 v22 + (f30 e301 − f21 e201 e10 + f12 e01 e210 − f03 e310 )v31
+ ν0 (2f12 e01 e10 − f21 e201 − 3f03 e210 )v21 v2
+ ν02 (f12 e01 − 3f03 e10 )v1 v22 − ν03 f03 v32 .
Now, we calculate the 1st Lyapunov coefficient, based on the
normal form (2.6), which is given by [Wiggins,24 Eq. (20.2.14), p.
385]
1  1
Fv1 v1 v1 + F1v1 v2 v2 + F2v1 v1 v2 + F2v2 v2 v2

l1 =
16
1  1
Fv1 v2 F1v1 v1 + F1v2 v2 − F2v1 v2 F2v1 v1 + F2v2 v2



+
16ν0
−F1v1 v1 F2v1 v1 + F1v2 v2 F2v2 v2 ,


where all the partial derivatives are calculated at the bifurcation

point; i.e., (v1 , v2 ; k) = (0, 0; kh ).
By applying the result given in Wiggins’s book,24 the Hopf
bifurcation is supercritical if l1 < 0 and subcritical if l1 > 0. Remark
that, when l1 = 0, the system (1.4) exhibits a generalized Hopf
(GH) bifurcation. The GH bifurcation point separates branches

FIG. 4. Figure shows bifurcation diagrams for both populations [prey in (a) and
predator in (b)] of the system (1.4) with respect to prey’s birth rate (b). Depending
on the nature of stability, we break it into two groups: stable equilibrium (gray area)
and limit cycle (blue area). In the first portion of the gray region, predators cannot
survive moving the system into a predator-free equilibrium. At a sufficiently higher
birth rate, both species coexist (last part of the gray region) and in the blue region
populations show periodic oscillations. Here, other parameter values are chosen
from (3.1), and the initial condition is fixed at (0.5, 1.0).

of subcritical and supercritical Hopf bifurcations in the parameter

plane.
FIG. 3. Figure shows the phase portrait of the system (1.4) in the absence of fear
effect, i.e., at k = 0. Other parameter values are chosen from (3.1), and the initial
condition is fixed at (0.5, 1.0). Here, both prey and predator species coexist in III. NUMERICAL SIMULATIONS
an oscillatory manner forming a stable limit cycle around the unstable equilibrium
point (0.3889, 2.5712). In this section, we analyze the system (1.4) with the help of
various numerical tools and techniques. We use the renowned RK4

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method for solving the differential equations. We solve the system

for at least T = 20 000 (enough for discarding initial fluctuations) for
plotting our results. The programs are written and run on MAT-
LAB. Our primary goal here is to extract more insight into the
system dynamics, which is almost impossible to get from an ana-
lytical approach. This also serves as a way to verify some of our
analytical results.
To start with, we first choose a biologically reasonable set of
parameter values given by the following:
b = 1.4, d = 0.3, a = 0.4, α = 0.2,
(3.1)
h = 2, µ = 0.7, m = 0.1.
We also fix the initial condition at (0.5, 1.0), unless otherwise
stated.
With parameter values chosen from (3.1) and in the absence of
fear effect (i.e., with k = 0), a phase-portrait diagram (see Fig. 3) of
the autonomous system (1.4) shows that the two interacting species
oscillate periodically forming a stable limit cycle around the unsta-
ble equilibrium point (0.3889, 2.5712). The birth rate (b) of the prey
(x) plays an important role in its survivability as well as its coexis-
tence with predator species. A bifurcation study of the system (1.4)
with respect to b, keeping the initial condition fixed at (0.5, 1.0),
reveals that the system shows three types of behaviors: prey-only
state, stable coexistence, and oscillatory coexistence. From Fig. 4,
we can draw a conclusion that the predator’s survivability is largely
dependent on prey’s birth rate (b), and only a sufficiently higher
value of b (namely, b > 0.87) makes the coexistence possible. For
0.87 < b < 1.011, the populations show stable coexistence and then
switch to limit cycle behavior via a Hopf bifurcation at b = 1.011.
We draw another phase portrait of the system (1.4) for b = 0.9
to reaffirm the stable coexistence of the two interacting species as

FIG. 6. Figure shows bifurcation diagrams of system (1.4) with respect to k, at

parameter values given in (3.1). (a) shows the change in the prey population, and
(b) shows the change in the predator population. Here, we observe that with a
gradual increase in the value of k, the amplitudes of periodic oscillations shrink
down and ultimately form a stable equilibrium via a Hopf bifurcation occurring at
k = 0.206 and thereafter remains stable. Further increase in k leads to predator’s
extinction.

indicated by the bifurcation diagrams (Fig. 4). Figure 5 shows that

FIG. 5. Figure shows the phase portrait of the system (1.4) reaffirming the stable
coexistence of the two interacting species at b = 0.9, keeping other parameters
fixed at those given in (3.1).
among the two coexisting equilibrium states, (1.2831, 0.7794) and
(0.6471, 1.5454), the former one is saddle and the latter is spirally
stable (the corresponding variational matrix has a pair of complex
conjugate eigenvalues with negative real part).
Now, we move our focus to the effect of fear (k) in the sys-
tem (1.4). Fear could have both stabilizing and destabilizing effects.25
At first, we explore the situation when initially the system shows a

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FIG. 8. Figure shows the bifurcation diagram of system (3.2) with respect to θ.
We have plotted the maximum peaks in blue and minimums in red. The system
shows chaotic dynamics along with some periodic and higher periodic windows.
Here, b = 1.4, k = 0, w = 1, and other parameters are chosen from (3.1).

A. The nonautonomous case

A large volume of literature exists in the study of a
mathematical model that describes the predator–prey relationship,
and in the last few decades, much progress has been seen in this
direction. Essentially, all these models consider within the frame-
work of an unvarying, deterministic environment. However, such

FIG. 7. Figure shows bifurcation diagrams of system (1.4) with respect to k, at

b = 0.9 and other parameter values chosen from (3.1). (a) shows the change
in prey population and (b) shows the change in predator population. Here, we
observe that with a gradual increase in the value of k, the stable equilibrium
remains stable and a further increase in k leads to predator’s extinction.

periodic oscillation. In Fig. 6, we have plotted two bifurcation dia-

grams of system (1.4) with respect to k, at parameter values given
in (3.1). It is observed that as we gradually increase the intensity of
fear, the amplitudes of periodic oscillations shrink down and ulti-
mately form a stable equilibrium via a Hopf bifurcation occurring at
k = 0.206 and thereafter remains stable. Further increase in k leads
FIG. 9. Figure shows the bifurcation diagram of system (3.2) with respect to θ.
to predator’s extinction. A similar study with b = 0.9 shows that the We have plotted the maximum peaks in blue and minimums in red. Here, b = 0.9,
stable equilibrium remains stable until further increase in k leads to k = 0, w = 1, and other parameter values are chosen from (3.1).
predator’s extinction (see Fig. 7).

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 Chaos
FIG. 10. Figure shows the bifurcation diagram of system (3.2) with respect to
k. We have plotted the maximum peaks in blue and minimums in red. Here,
the system shows chaotic dynamics along with some periodic and higher peri-
odic windows. Here, θ = 0.5, w = 1, and other parameter values are chosen
from (3.1).
an unvarying or constant environment cannot reflect the reality of
nature. Most natural environments are physically highly variable,
and in response, the population’s birth and death rates, carrying
capacities, competition coefficient, and many other parameters that
characterize the deterministic model vary greatly in time.26 For this
FIG. 12. Figure shows the largest (a) and the second-largest (b) Lyapunov exponents (LE) of system (3.2) in θ-k biparametric space. Other parameter values are chosen
from (3.1).
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FIG. 11. Figure shows isoperiodic regions in θ-k biparametric space of system
(3.2). Here, period means the periodicity of the peaks of respective time-series
evaluations discarding initial fluctuations. EP denotes the equilibrium state. Apart
from equilibrium, we have presented regions of period 1 to period 18 in distinct
colors and the rest in black color, which includes chaotic attractors. A sidebar rep-
resents the periods corresponding to each color. Here, we can also see some
shrimp-shaped structures and another structure that seems to be like Arnold-
tongue. During this simulation, we fixed our initial condition at (0.5, 1.0), and
other parameter values are chosen from (3.1).
reason, time-dependent predator–prey systems have been widely
investigated.
It is well accepted that temporal fluctuations in the phys-
ical environment are major drivers of population fluctuations.
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FIG. 13. Figure shows different versions of attractor-following isospike regions in θ-k biparametric space of system (3.2). All four subfigures are drawn in an attractor-following
manner. (a) is drawn by first solving system (3.2) for θ = 0 and for all k values (vertical line θ = 0). The last point of the solution is used as the initial condition for the next
value of θ. This process is repeated until we reach θ = 0.8 (horizontal forward). (b) is drawn changing θ backward from 0.8 to 0 (horizontal backward) following the same
process. Similarly, in (c), we first take the horizontal line k = 0 for all θ and then increase k until we reach k = 0.25 (vertical forward). In (d), we just change k backward from
0.25 to 0 (vertical backward). For the first iteration, we used (0.5, 1.0) as the initial state of system (3.2). Here, the last points of the solutions of each iteration are used as
the initial states of the next iteration. Other parameter values are chosen from (3.1).

Theoretical evidence to date indicates that many populations and
community patterns represent intricate interactions between biol-
ogy and variation in the physical environment (for a brief review,
see Chesson27 and related papers in the same issue). When envi-
ronmental fluctuation is taken under consideration, a model must
be nonautonomous and thus more difficult to analyze in general.
However, one can take advantage of the properties of those varying
parameters by assuming that the parameters are periodic or almost
periodic for seasonal reasons.
Seasonal variation in birth rates is common among many
species. There could be many factors contributing to such fluc-
tuations, for example, temperature, humidity, rainfall, change in
daylight period, an abundance of food, etc.28–30 It may happen due to
some seasons being evolutionarily beneficial and favorable for giving
birth and raising the young. According to most research, seasonal
breeding is driven by dynamics related to resource availability. Many
wild mammals31–33 and birds29,34,35 show seasonal breeding. In North
America, during the spring and early summer months, when food is

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 Chaos
FIG. 14. Figure shows the zoomed-in view of two different modes of the attractor following the isospike diagram in θ − k biparametric space. (a) is drawn in by taking the
vertical line θ = 0.65 and moving it backward (horizontal backward). (b) is drawn in by taking the horizontal line k = 0.06 and moving it backward (vertical backward). Other
parameter values are chosen from (3.1).
plentiful and it is not too cold, mule deer give birth.36,37 In savanna
ecosystems, the variation of resource availability is greatly depen-
dent on rainfall. Ryan et al.38 observed that African buffalos are more
likely to give birth during the wet season of every year. In Canada,
the song sparrow lays its eggs and hatches them between April and
June every year.17 There is ample evidence of seasonal breeding in
marine and coastal ecosystems from several studies.39–41
We, therefore, take another step by introducing the periodic
birth rate into the system (1.4). Thus, we replace the birth rate b with
b + θ cos(wt) making system (1.4) into a nonautonomous system
and get
(b + θ cos wt)x αxy2

dx
− dx − ax2 − , Although bifurcation diagrams are very useful in revealing
 dt =


1 + ky 1 + hαxy
(3.2)
µαxy 2
 dy =


dt
− my.
1 + hαxy
Here, θ is the amplitude of variation and w represents the corre-
sponding frequency. We also set w = 1 for all subsequent simula-
tions of system (3.2).
Now, we simulate the nonautonomous system (3.2) by first
changing θ and keeping k = 0 and choosing other parameter values
from (3.1). Figure 8 shows the corresponding bifurcation diagram
of system (3.2). Here, the blue dots represent the local maximums
of the time-series data, and the red ones represent the local mini-
mums. We observe that the system moves from the initial periodic
state to chaotic oscillations along with some periodic windows. We
have added a colorbar below along with a sidebar on the right to
represent the periodicity of the peaks corresponding to each color.
Here, black dots are associated with the periodicity higher than 18,
which also includes the chaotic regimes.
Chaos 31, 123134 (2021); doi: 10.1063/5.0067046
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For b = 0.9 and k = 0, we draw another bifurcation diagram
of system (3.2) with respect to θ (see Fig. 9). Here, we see that
the system moves from a stable to periodic state at the very first
instant where θ becomes nonzero and remains the same except for
an increase in the amplitude of oscillation (due to an increase in θ ).
In Fig. 10, we draw the bifurcation diagram of the system (3.2)
with respect to the fear parameter k with θ = 0.5, w = 1, and other
parameter values are chosen from (3.1). We see that with an ade-
quate increase in the strength of fear, the system (3.2) becomes
periodic from a chaotic state and remains there. One can also check
that beyond a certain level of fear, the predator population (y)
becomes extinct.
the rich dynamics of a system such as (3.2), it is quite limited to
only showing changes for a single parameter value. Variation of
one more parameter gives us extra advantages and often unfolds
many organized periodic structures in the chaotic regime that can-
not be seen otherwise.42 More parameters can be varied at once,
but it is very hard to visualize on the plane of a paper. For this,
we draw an isospike diagram of the system (3.2) in θ − k bipara-
metric space keeping other parameter values fixed at those given in
(3.1). (Fig. 11). We also fix our initial condition at (0.5, 1.0) through-
out the simulation. Here, period means the periodicity of the peaks
(spikes) of the time-series data for each pair of parameter values.
We have discarded the first T = 20 000 time-series data to remove
initial fluctuations. In this rectangular region ([0, 0.8] × [0, 0.25]),
filled with 600 × 600 colored dots, each color is associated with
unique periodicity mentioned in the sidebar on the right. Here, EP
represents the equilibrium state. Chaotic and periodic regions with
periodicity higher than 18 are represented in black. One can now
easily correlate Figs. 8 and 10 with Fig. 11 and verify the results.
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FIG. 15. Figure shows sections of time-series evaluations and corresponding basins of attraction for three different pairs of (θ, k) values for which system (3.2)
shows multistability between multiple coexisting attractors. (a) and (b) corresponding to (θ, k) = (0.4, 0.053) show the multistability between attractors of peri-
odicities 1, 2, and 3. Similarly, (c) and (d) corresponding to (θ, k) = (0.4, 0.048) show the multistability between attractors of periodicities 1, 2, and 6, and (e)
and (f) corresponding to (θ, k) = (0.452, 0.048) show the multistability between attractors of periodicities 1, 2, and 12. Here, other parameter values are chosen
from (3.1).

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 Chaos
FIG. 16. Figure shows the largest Lyapunov diagram of two different segments from θ − k biparametric space (Fig. 12) containing different organized periodic structures of
system (3.2). (a) shows shrimp-shaped structures, and (b) shows Arnold tongues. Here, other parameter values are chosen from (3.1).
Here, we see many organized periodic structures including shrimp-
shaped periodic islands inside the chaotic regime. We also see the
overlapping of different periodic zones indicating the possible pres-
ence of multistability between multiple coexisting attractors.43 We
discuss more about these phenomena in Subsection III A 1.
Lyapunov exponents are very well-known measures of expo-
nential convergence or divergence of infinitesimally close orbits.
In general, with very few exceptions, a positive value of the largest
Lyapunov exponent (LLE) indicates the chaotic nature of a system.
Similarly, a negative value of LLE represents a stable equilibrium
state. Regular oscillations give zero LLE. Although very useful, the
computational cost of calculating the Lyapunov spectrum of a sys-
tem such as (3.2) for a large set of parameter values is huge. Here,
we draw the largest [Fig. 12(a)] and second-largest [Fig. 12(b)]
Lyapunov exponents (LE) of system (3.2) in θ − k biparametric
space ([0, 0.8] × [0, 0.25]). We used Wolf’s algorithm involving
variational equations. This involves discarding of first T = 20 000
time-series data to make sure the absence of any initial fluctuations
and averaging relative exponential divergence rates of nearby tra-
jectories for another T = 10 000. A side-by-side comparison with
Fig. 13 and some easy time-series evaluations reveal that the regions
where the system has positive LLE shows chaotic nature.
1. Multistability
To confirm the multistable nature of our system (3.2) in some
certain zones in the abovesaid biparametric space, we draw four dif-
ferent versions of Fig. 11 using an attractor-following technique. In
this technique, we first choose a vertical or horizontal line segment
from the four boundary lines of the rectangular area and solve the
system for each pair of parameter values on the line segment. In
this step, we use a fixed initial state (here [0.5, 1.0]) for each point
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of the line segment. We then use the last points of the solutions
as the initial states of the neighboring line segment and continue
this process until we reach the other boundary. Depending on the
choice of starting line, we have four different variations presented in
Fig. 13. In Fig. 13(a), we choose the vertical line θ = 0 and proceed
in a forward direction (θ increasing), and in Fig. 13(b), we use the
vertical line θ = 0.8 and move backward to 0 (θ decreasing). Simi-
larly, in Figs. 13(c) and 13(d), we, respectively, choose the horizontal
lines k = 0 and k = 0.25 and proceed in forward (k increasing) and
backward directions (k decreasing), correspondingly.
From the four subfigures in Fig. 13, one can easily point
out their color differences and distinguish some of the prominent
multistability zones. To take a closer look, we redraw the region
enclosed by the white dotted rectangle ([0.35, 0.65] × [0.016, 0.06]
in θ − k biparametric space) in Fig. 13(b) (horizontal backward
version) and Fig. 13(d) (vertical backward version) and display in
Fig. 14. Here, we can clearly observe the overlapping of multiple
pairs of coexisting attractors of different periodicities. These differ-
ences strengthen our claim for the presence of multistability in our
system (3.2).
To be accurate, we further pick three (θ , k) pairs from three
different multistable regions displayed in Fig. 14 and explore their
corresponding time-series evaluations for three different sets of
initial conditions. We also draw their basins of attraction. Those
three (θ , k) pairs are (0.4, 0.053) from a red-green overlapping
zone, (0.4, 0.048) from a deep green-green overlapping zone, and
(0.452, 0.048) from a violet-green overlapping zone. In Fig. 15, we
plot the time-series evolutions on the first column, and correspond-
ing basins of attraction are placed on their right. Figures 15(a)
and 15(b) corresponding to (θ , k) = (0.4, 0.053) show the multi-
stability between attractors of periodicities 1, 2, and 3. Similarly,
Figs. 15(c) and 15(d) corresponding to (θ , k) = (0.4, 0.048) show
31, 123134-13
 Chaos
FIG. 17. Figure shows self-similar patterns in different zoomed-in versions of the fractal basin presented in Fig. 15(b) corresponding to (θ, k) = (0.4, 0.0453). (a) represents
the area inside the white dotted rectangle in Fig. 15(b). (b) is the zoomed-in view of (a). (c) represents the area inside a smaller rectangle (at the left), and (b) and (d) represent
the larger one (at the right). Here, other parameter values are chosen from (3.1).
the multistability between attractors of periodicities 1, 2, and 6, and
Figs. 15(e) and 15(f) corresponding to (θ , k) = (0.452, 0.048) show
the multistability between attractors of periodicities 1, 2, and 12.
Since the birth rate of the prey varies periodically with time, the sys-
tem has no stable equilibrium point. All the subfigures presented in
Fig. 15 are drawn in terms of the prey population (x). In the blue
region, predators cannot survive and prey oscillates due to the peri-
odic nature of its birth rate. The readers may notice the self-similar
pattern of the basin of attraction in all three cases. We explore it
further in Sec. III A 3.
2. Organized periodic structures
In Fig. 12, we observe different organized periodic
structures. Among them, we found shrimp-shaped structures and
Arnold tongues particularly noteworthy to report. We selected two
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different rectangular sections (violet-bordered boxes) of the (θ , k)
plane presented in Fig. 12. In Fig. 16(a), we scatter plot the LLE for
a grid of 500 × 500 equispaced pairs of parameter values (θ , k) ∈
[0.37, 0.4] × [0.03, 0.04]. Here, we observe multiple shrimp-shaped
organized periodic structures inside the chaotic regime. It is inter-
esting to note that one can explore the period-bubbling transition
to chaos near the inner boundaries of the shrimp.42,44 Similarly,
for (θ , k) ∈ [0.095, 0.225] × [0.065, 0.095], we see the presence of
Arnold tongues in Fig. 16(b). The colorbars represent the LLE values
for colors attached to them.
3. Fractal basin
In all three basins of attraction presented in Fig. 15, we see the
presence of self-similar (fractal) patterns. To demonstrate this, we
31, 123134-14
 Chaos
pick a rectangular region from Fig. 15(b) (indicated by the white
dotted area) and draw it again on Fig. 17(a). Figure 17(b) shows
a zoomed-in view of the white rectangular area at the center of
Fig. 17(a). It displays a similar pattern. We again plot two dif-
ferent rectangular sections of the basin of attraction of Fig. 17(b)
in Figs. 17(c) and 17(d) (in the left to right order). Both exhibit
self-similar patterns. This kind of phenomenon is quite rare in
continuous-time models. Here, each area is divided into 600 × 600
equispaced grid points, and then the system (3.2) is solved for all
360 000 pairs of initial conditions and for time T = 30 000. We ana-
lyzed the system behavior for the last T = 5000 (i.e., discarding first
T = 25 000 time-series data) to detect the periodicity of the attractors
and plot them in their respective colors.
IV. CONCLUSION
In the present study, we have investigated the impact of the fear
effect in a predator–prey model. We considered that due to the fear
of predation risk, the birth rate of the prey population reduces in
both autonomous and nonautonomous versions of the model. The
whole investigation of the system is mainly centered on two impor-
tant factors, namely, the birth rate of prey and the fear effect. Using
phase-portrait, bifurcation, isospike, and Lyapunov exponent dia-
grams, we explored the rich and complex dynamical behaviors of
the system. First, we explored how changes in seasonal fluctuations
in the birth rate influenced the population cycles. In contrast to the
autonomous system, which shows periodic behaviors, the nonau-
tonomous system exhibits chaotic oscillations. Seasonal variations
in the level of birth rate lead to higher periodic and chaotic oscilla-
tions and produce dynamical complexities in the ecosystem. It was
also noticed that increased fear of predation risk potentially sup-
presses the irregular oscillations in the system. We also explored
the global behavior of the systems in the biparameter space (birth
rate and fear effect) by varying the control parameters. We found
various types of several periodic islands in the chaotic regimes.
Therefore, unexpected shifts in the dynamics of the system can hap-
pen in a small variation of the control parameters. The well-known
organized structures such as shrimp-shaped structures and Arnold’s
tongues are detected in the biparameter space. It is to be noted
that period-bubbling transition to chaos can be found due to the
existence of shrimp-shaped structures. Furthermore, in the isospike
diagram, overlapping of multiple periodic windows is found, and
multistability between topologically different coexisting attractors is
detected. The complexities of the multistable attractors are exam-
ined by drawing the basin of attractions. We drew basins of several
coexisting attractors and found fractal basin boundaries. Here, the
parameter values and the initial population sizes of prey and preda-
tor play a vital role in the dynamics of the system, and the solution
trajectories can converge to different non-equivalent attractors. It is
observed that the persistence of the predator population also highly
depends on the initial population densities. Therefore, small changes
in the parameters, perturbations, or environmental fluctuations can
have drastic outcomes in the oscillations of the populations. In the
present model system, the fear effect has a stabilizing role on the
autonomous system, whereas in the nonautonomous system, the
fear effect drives the system toward regularity. In summary, we
Chaos 31, 123134 (2021); doi: 10.1063/5.0067046
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conclude that the nonautonomous ecological systems are inherently
complex and exhibit intricate dynamical behaviors.
SUPPLEMENTARY MATERIAL
In the supplementary material, we include two movies show-
ing the qualitative change in the dynamics of the model (1.4) when
the strength of fear (k) is varied. We plot the nullclines of both
species in the first quadrant to show the coexisting equilibrium
points. Throughout the animations, the red curve represents the
prey nullcline, and the green one represents the predator nullcline.
We added the vector fields corresponding to each value of k by using
the “quiver” command in MATLAB. We also plotted solution curves
for three arbitrarily chosen initial conditions to show where the solu-
tions converge. This gives a clear understanding of the stability of the
coexisting equilibrium points.
The movie named nullcline1 shows nullclines of the system
(1.4) for different values of k, ranging from 0 to 1 where b = 1.4
and other parameter values are chosen from (3.1). The other movie,
named nullcline2, does the same except for b = 0.9 and varying k in
the range of 0–0.1. In both cases, we see that there are at most two
coexisting equilibrium states in the system (1.4), which are repre-
sented by the intersection of the nullclines. Among these two, one
is always saddle. In the present work, the dynamics of the system
(1.4) are explored with the help of numerical simulations around
the non-saddle equilibrium point.
ACKNOWLEDGMENTS
The authors are thankful to the anonymous reviewers and the
handling editor for their valuable comments and suggestions, which
helped us improve the quality of the manuscript. M. Hossain is
grateful to the Department of Science and Technology (DST), India,
for providing financial support under the INSPIRE Fellowship
(No. IF-170522) program.
AUTHOR DECLARATIONS
Conflict of Interest
The authors have no conflicts to disclose.
Author Contributions
All authors contributed equally to this work.
DATA AVAILABILITY
The data that support the findings of this study are available
within the article.
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