Dynamical analysis in predator-prey-

scavenger model with harvesting

intervention on prey population
Cite as: AIP Conference Proceedings 2192, 060005 (2019); https://doi.org/10.1063/1.5139151
Published Online: 19 December 2019

Putri A. Dumbela and Dipo Aldila

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AIP Conference Proceedings 2192, 060005 (2019); https://doi.org/10.1063/1.5139151 2192, 060005

© 2019 Author(s).
Dynamical Analysis in Predator-Prey-Scavenger Model with
Harvesting Intervention on Prey Population
Putri A. Dumbela1,a) and Dipo Aldila1,b)
1
Department Mathematics, Faculty of Mathematics and Natural Science, Universitas Indonesia, Indonesia
a)
putri.angriani@sci.ui.ac.id
b)
Corresponding author: aldiladipo@sci.ui.ac.id

Abstract. This study deals with the interaction between predator, prey, and scavenger. In our model, we assume that the prey
population grows following a logistic model with predation from predator and scavenger. Harvesting intervention included in
the prey population to understand scenarios where the prey has competitive effects upon the other species. The analytical result
began with a non-dimensionalization process to the original model, which successfully reduced the number of parameters of the
model. Equilibrium points were shown analytically. Local stability of the fixed p oint w as a nalyzed u sing a r espected Jacobian
matrix of each fixed p oint. T he s ensitivity a nalysis o f t he m odel r egarding t he h arvesting r ate w as c onducted n umerically. We
find t hat w henever t he h arvesting i n t he p rey p opulation b ecomes u ncontrolled ( above t he u pper b ound), t hen t he p redator and
scavenger population will die out together with the prey population. We also find t hat t he t ime s cale s eparation f rom t he non-
dimensionalization process does not change the stability criteria of the fixed point.
Keywords: Predator-Prey-Scavenger Model, Local Stability, Harvesting.

INTRODUCTION
An ecosystem consists of all organisms that live in a community and also include all abiotic factors which interact
with these organisms. To interact with abiotic factors, a group of similar species or populations will also interact with
individuals in other populations. One kind of interaction is predation. The dynamic relationship between prey and
predators is very closely related. If one species in the food chain is lost, then the ecosystem will be disrupted, thus
causing the extinction of other species [14]. This phenomenon can be modeled mathematically into a predator-prey
model.
The simplest predator-prey model was introduced by Lotka [4, 5] and Volterra [6] in the early 1900s. The model
is given as follows:
dx
= ax − βxy,
dt
dy
= δxy − cy,
dt
where x(t) and y(t) present the prey and predator population in time t, respectively. a, β, δ , and c are the growth
rate of the prey, the effect of predation, rate of the predator in the presence of prey, and the natural date rate of
the predator population, respectively. However, the model has a weakness because, in Malthus’s growth model, the
population grows not limited along with the increased time. In fact, a population cannot increase continuously because
of the inhibiting factors such as living space, food sources, and environment. Therefore, the Lotka-Volterra model was
modified, assuming prey growth following the growth of logistics to better reflects reality [3].
Moreover, the Lotka Volterra model does not add the time needed by predators to digest their food and the fact
that food from prey is limited. Then in 1950, Holling introduced the response function, which is the amount of food
eaten by predators as a function of food density [9, 10]. In recent years, many researchers have studied the type of
functional response behaves; for example, see [12]. They compared a system with two predator-prey models using a
a Leslie and Laissez-faire-type model with type I functional responses.

Proceedings of The 8th SEAMS-UGM International Conference on Mathematics and its Applications 2019
AIP Conf. Proc. 2192, 060005-1–060005-8; https://doi.org/10.1063/1.5139151
Published by AIP Publishing. 978-0-7354-1943-8/$30.00

060005-1
 Furthermore, many studies have shown that scavenger is found in food chains. Authors in [1, 2, 3, 7, 8] success-
fully introduce how prey interact with predator and/or with scavenger in the same ecosystem. Authors in [1] introduce
a simple 3D model between prey, semipredator, and predator population. Their numerical result shows the existence
of chaotic behavior. The author in [2] introduces a model interaction between one prey and two competing predators,
and the existence of two limit cycle have been proved by authors. A similar model with the author in [2], but with
different functional responses have been introduced by the author in [3]. The analytical result regarding equilibrium
and the dynamical property also discussed. A fractional-order derivative model for interaction between prey, predator,
and scavenger introduced by the author in [7], but do not involve the Allee effect and fear factor in the prey growth
term. A mathematical model for interaction between prey-predator and scavenger population introduced by the author
in [8] as a three dimensional of an ordinary differential equation. A comprehensive analytical result regarding period-
doubling cascades has been conducted in their paper. Although the model in [8] does not involve the harvesting rate.
Besides, several kinds of literature have mentioned that harvesting [13, 15, 16] as human intervention can have a sig-
nificant effect on the behavior of the ecosystem. Thus, based on the above discussion, we consider a three-dimensional
system for predator- prey-scavenger (where scavenger has no inhibitory effect on predator) with harvesting on prey
population, based on the model in [8] which given by :
 
dx x
= ax 1 − − β1 xy − β2 xz − μ1 x − hx,
dt K
dy
= c1 β1 xy − μ2 y, (1)
dt
dz
= c3 β1 xy + c2 β2 xz − μ3 z,
dt
where x = x(t), y = y(t), z = z(t) represent the population size of prey, predator and scavenger species at time t,
respectively. The growth of the prey population is taken as a logistic type with a and K present the growth rate and
carrying capacity, respectively. The predator and scavenger consume the prey by a Hollings type I, with β1 and β2
present the contact rate between prey-predator and prey-scavenger, respectively. The term μ1 , μ2 and μ3 denote the
death rate of the prey, predator and scavenger, respectively. Note that μ1 < μ2 < μ3 . On the other hand c1 , c2 , and c3
are the energy conversion caused by predation on the prey population. All the parameters are assumed to be positive
except h̄ which is non-negative and all variables are non-negative.
To simplify the analytical calculation, we consider the following new variable and parameters.
x β1 y β2 z μ1 μ2 β 1 μ3 β2 h c3 ab1 ab2 t
x= ,y = ,z = , μ1 = , μ2 = , μ3 = , h = , c = , 1 = , 2 = ,t = .
K a a a c1 K c2 K a c2 c1 K c2 K a
Substitute these to system (1), we have a following non dimensional system of predator-prey-scavenger in the form
of :
dx
= x (1 − x) − xy − xz − μ1 x − hx,
dt
dy 1
= (xy − μ2 y), (2)
dt 1
dz 1
= (cxy + xz − μ3 z),
dt 2
where 1 and 2 is the time scale for predator and scavenger respect to prey population. Since our population are
nonnegative, then we have (x, y, z) ∈ Ω where Ω = {(x, y, z)|x ≥ 0, y ≥ 0, z ≥ 0}.
The paper is organized as follows: In Sect.1 we begin with review of the dynamics classic two-species Lotka
Voltera and introduce a scavenger into the model. In Sect. 2 we discuss the existence and stability of trajectories in
each of coordinate planes. In Sect. 3, we give some numerical simulations. Finally, some conclusions close the paper
in Sect. 4.

DYNAMICAL BEHAVIOR
In this section, we investigate the dynamics of the system (2) by analyzing the sign of the eigenvalues of the corre-
sponding characteristic equation. Taking the right-hand side of the system (2) equal to zero, and solve it respect to x, y

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and z, we have four fixed points of system (2) which given by

Ω1 = (0, 0, 0),
Ω2 = (1 − h − μ1 , 0, 0),
Ω3 = (μ3 , 0, 1 − h − μ1 − μ3 ), (3)
 
(μ3 − μ2 ) (1 − h − μ1 − μ2 ) cμ2 (1 − h − μ1 − μ2 )
Ω4 = μ2 , , .
cμ2 + μ3 − μ2 cμ2 + μ3 − μ2

Please note that all this equilibrium will have a biological meaning if Ωi ∈ Ω for i = 1, 2, 3, 4. Further analysis of the
existence of Ωi will be given later. Next, to analyze the local stability of each Ωi , let us define the Jacobian matrix of
system (2) based on standard linearization technique and using the Routh-Hurwitz criterion [11] as follows :
⎡ ⎤
⎢⎢⎢ 1 − 2 x − y − z − μ1 − h −x −x ⎥⎥
⎥⎥⎥
⎢⎢⎢ ⎥
J = ⎢⎢⎢⎢⎢ 0 ⎥⎥⎥⎥
y x−μ2
1 1 ⎥⎥ (4)
⎢⎢⎣
cy+z cx x−μ3 ⎥ ⎦
2 2 2

With the Jacobian matrix J and fixed point Ωi in hand, we are ready to step forward for analytical calculation
for the existence and local stability of Ωi .
1. The trivial fixed point Ω1 is always has a biological meanings (Ω1 ∈ Ω), which describe the extinction of all
populations. The Jacobian matrix at the interior fixed point Ω1 takes the form
⎡ ⎤
⎢⎢⎢ −h − μ1 + 1 0 0 ⎥⎥⎥
⎢⎢⎢ ⎥⎥⎥
J1 = ⎢⎢⎢⎢⎢ 0 − μ12 0 ⎥⎥⎥ ,
⎥⎥⎥ (5)
⎢⎢⎣ ⎥⎦
0 0 − μ23

where the eigenvalues of J1 are :

μ2 μ3
λ1 = 1 − h − μ1 , λ2 = − , λ3 = − .
1 2
It follows that the fixed point Ω1 is locally asymptotically stable if and only if h + μ1 > 1. This result tells us that
if the harvesting and natural date of prey are sufficiently large and more than 1, then the system cannot sustain
the populations of prey, which has an impact on the extinction of predator and scavenger. Based on the above
analysis, we obtained:
Theorem 1. System (2) has an extinction fixed point Ω1 = (0, 0, 0) and locally asymptotically stable when
h + μ1 > 1, and saddle node if h + μ1 < 1.

2. Coexistence of prey Ω2 has biological interpretation if only if h + μ1 < 1. This fixed point represents the limiting
population of prey in the absence of the predator and scavenger. Note that, from Theorem 1 we have h + μ1 > 1
where it contradicts the coexistence of Ω2 . Therefore, we conclude that the existence of Ω2 causing instability
of Ω1 and vice versa.
The Jacobian of Ω2 takes the form
⎡ ⎤
⎢⎢⎢ h + μ1 − 1 h + μ1 − 1 h + μ1 − 1 ⎥⎥⎥
⎢⎢⎢⎢ −h−μ1 +1−μ2
⎥⎥⎥⎥
J2 = ⎢⎢⎢⎢ 0 0 ⎥⎥⎥ , (6)
⎢⎢⎢ 1
⎥⎥⎥
⎣ c(−h−μ1 +1) −h−μ1 +1−μ3 ⎥ ⎦
0 2 2

where the eigenvalues of J2 are:

1 − (h + μ1 + μ2 ) 1 − (h + μ1 + μ3 )
λ1 = h + μ1 − 1, λ2 = , λ3 =
1 2

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 We see that λ1 < 0 is always hold due to coexistence of Ω2 , while λ2 and λ3 has negative sign if h + μ1 + μ2 > 1
and h + μ1 + μ3 > 1, respectively.
From the above analysis, we obtained :
Theorem 2. System (2) has fixed point Ω2 = (1 − h − μ1 , 0, 0) which has biological meanings if h + μ1 < 1 and
asymptotically stable if h + μ1 + μ2 > 1 and h + μ1 + μ3 > 1.
According to Theorem 1 and 2, we have the following corollary:
Corollary 1. Whenever Ω1 is stable, then Ω2 will not exist.

3. Extinction of predator fixed point Ω3 feasible if h + μ1 + μ3 < 1, i.e. harvesting with a natural date of prey and
scavenger less then 1. The Jacobian matrix related to this fixed point is
⎡ ⎤
⎢⎢⎢ −μ3 −μ3 −μ3 ⎥⎥
⎢⎢⎢ ⎥⎥⎥
J3 = ⎢⎢⎢⎢⎢ 0 μ3 −μ2
1 0 ⎥⎥⎥⎥⎥ (7)
⎢⎢⎣ ⎥⎥⎥
−h−μ1 +1−μ3 cμ3 ⎦
2 2 0

whose characteristic equation of J3 is given by


(λ 1 + μ2 − μ3 ) λ2 2 + λ 2 μ3 − hμ3 − μ1 μ3 − μ3 2 + μ3
= 0.
1 2
It follows that Ω3 stable if and only if all eigenvalues have a negative sign. Note that, one eigenvalue is given
by λ1 = μ3−μ
1
2
, which is always negative since μ3 < μ2 always hold true based on the previous assumption. The
other two eigenvalues are given by the solution of the quadratic equation:

λ2 2 + λ 2 μ3 + μ3 − hμ3 − μ1 μ3 − μ3 2 = 0.

From the above equation, we have a maximum of two eigenvalues that have a negative sign. To guarantee the
stability of J3 we are using the standard basic theory of quadratic equation,
(a) λ1 .λ2 = −hμ3 −μ1 μ23 −μ3 +μ3 > 0
2

↔ −hμ3 − μ1 μ3 − μ3 2 + μ3 > 0
↔ −h − μ1 − μ3 + 1 > 0
↔ h + μ1 + μ3 < 1.
we see that h + μ1 + μ3 < 1 is always satisfied since Ω3 exist.
(b) λ1 + λ2 = − 2μ2 3 < 0 always hold since all parameters are positive.
Since from existence criteria of Ω3 we have that h + μ1 + μ3 < 1, then it is always held that μ1 + μ3 < 1.Based
on the above analysis, we obtained the following theorem.
Theorem 3. System (2) has a fixed point Ω3 = (μ3 , 0, 1−h−μ1 −μ3 ) which has biological meaning if h+μ1 +μ3 < 1
and always locally stable if μμ32 < 1
According to Theorem 2 and 3, we have the following corollary:
Corollary 2. Whenever Ω2 is stable, then Ω3 will not exist.

4. Steady state of the coexistence, called, Ω4 is feasible if and only if μμ32 > 1 and h + μ1 + μ2 < 1. Now we turn to
analyze the stability of this fixed point by looking the sign of the eigenvalues of the following Jacobian matrix :
⎡ ⎤
⎢⎢⎢ 1 − 2 μ2 − hμ2 −hμ3 +μ1 μ2 −μ1 μ3 +μ2 −μ2 μ3 −μ2 +μ3 + cμ2 (h+μ1 +μ2 −1) − μ1 − h −μ2 ⎥⎥⎥
2
−μ2
⎢⎢⎢ cμ2 −μ2 +μ3 cμ2 −μ2 +μ3 ⎥⎥⎥
⎢⎢⎢ ⎥⎥⎥
J4 = ⎢⎢⎢⎢⎢ 0 ⎥⎥⎥⎥ ,
hμ2 −hμ 3 +μ μ
1 2 −μ μ
1 3 +μ 2
2
−μ μ
2 3 −μ2 +μ 3
1 (cμ2 −μ2 +μ3 ) 0 (8)
⎢⎢⎢ ⎥⎥⎥
⎢⎢⎣  ⎥⎥
1 c(hμ2 −hμ3 +μ1 μ2 −μ1 μ3 +μ2 −μ2 μ3 −μ2 +μ3 ) μ2 −μ3 ⎥ ⎦
2

2 cμ2 −μ2 +μ3 − cμ2 (h+μ1 +μ2 −1)

cμ2 −μ2 +μ3
cμ2
2 2

whose characteristic equation of J4 is given by

λ3 + Aλ2 + Bλ + C = 0,

060005-4
 where
2 μ2 − μ2 + μ3
A =


2 
μ2 μ3 1 (ch + cμ1 − c − μ3 ) − h2 (μ2 − μ3 ) − μ2 2 (−c1 + 1 + 2 ) + μ2 μ3 (2 1 + 2 ) − 2 μ1 (μ2 − μ3 ) + 2 (μ2 − μ3 )
B = −
1 (cμ2 − μ2 + μ3 ) 2
(μ2 − μ3 ) (h + μ1 + μ2 − 1) μ2
C = .
1 2
According to the Routh-Hurwitz criterion, to make the coexistence fixed point stable, we need A > 0, C > 0,
and AB − C > 0. It is easy to see that A > 0 and C > 0 always hold true due to coexistence of Ω4 . Therefore,
we only need to have that AB − C > 0, or equivalently AB C > 1. From the above analysis, we obtained :

Theorem 4. System (2) has fixed point Ω4 = μ2 , (μ3 −μcμ2 )(1−h−μ 1 −μ2 ) cμ2 (1−h−μ1 −μ2 )
2 +μ3 −μ2
, cμ2 +μ3 −μ2 which has biological meaning
μ3
if h + μ1 + μ2 < 1 and μ2 > 1. In the other hand, Ω4 will locally asymptotically stable if AB C > 1.
According to Theorem 2 and 4, we have the following corollary:
Corollary 3. Whenever Ω2 is stable, then Ω4 will not exist.

According to the above discussion, we conclude that 1 and 2 do not affect the existence and stability criteria of all
fixed points. We closed this section with the following table, which describes the existence and local stability criteria
of all fixed points of system (2).

TABLE 1. Summary of the existence and local stability of system (2)

Fixed Point Ω1 = (0, 0, 0) Ω2 = (x∗ , 0, 0) Ω3 = (x∗ , 0, z∗ ) Ω4 = (x∗ , y∗ , z∗ )
μ3
Existence - h + μ1 < 1 h + μ 1 + μ3 < 1 >1
μ2
h + μ1 + μ2 < 1
μ3
Stability h + μ1 > 1 h + μ 1 + μ2 > 1 μ2 <1 AB
C >1
h + μ1 + μ3 > 1

From Table 1, it can be seen that most of all, existence and local stability criteria for all equilibrium points depend
on the harvesting rate. For example, the local stability of Ω1 is given by condition h + μ1 > 1, which means that more
harvesting allowed in the environment to prey population, then more possible that the extinction equilibrium will
occur and stable. A similar interpretation might be used to the other parameter based on Table 1.

NUMERICAL SIMULATION
In this section, we conduct some numerical simulations to illustrate the theoretical result given in the previous section.

Harvesting Effect
Figure (1) show how the effect of harvesting against the dynamic of each population in the system (2). Using 1 =
1, 2 = 1, μ1 = 0.1, μ2 = 0.4, μ3 = 1, c = 0.2 with initial condition x(0) = 0.5, y(0) = 0.3, z(0) = 0.03, we have
all trajectory of figure 1 will goes to Ω4 (stable node). As h increase further, the population of prey will decrease
over time, resulting in reduced food resources for predator and scavenger. Therefore, the population of predator and
scavenger will also decrease whenever h increasing. Furthermore, based on table 1, using the above parameters, we
found that when h = 0.6 > 0.5, then Ω4 will not exist, and the trajectory of system (2) will goes to Ω2 where the
predator and scavenger are driven to extinct. Based on this simulation, controlling harvesting rate in prey population
play an important rule to the existence of all population (predator, prey, and scavenger) in the ecosystem.

060005-5
 FIGURE 1. Time series of the prey x(t), predator y(t), and scavenger z(t) with various h

Time Scale Separation Effect

In this simulation, we conduct a numerical simulation to determine the effect of 1 and 2 to the behavior of the
system (2), even though it does not change the existence or the local stability of fixed points (based on analytical
result). For an illustration, we set parameters value μ1 = 0.1, μ2 = 0.4, μ3 = 1, c = 0.2, and h = 0.2. The 1 and 2 will
be varying to understand how the time scale separtion affects the dynamics of the system (2).

1. 1 = 2
For the first simulation we set a special case 1 = 2 which the predator and scavenger have the same lifetime.
Two species that might meet the above conditions are hyenas and lions, for example. For each value of 1 = 2 ,
we found that the system will goes to Ω4 with eigenvalues for red(1 = 2 = 0.1), magenta(1 = 2 = 0.4),
blue(1 = 2 = 0.7) and green(1 = 2 = 1) line are (−0.218 + 1.076I, −0.218 − 1.018I, −5.963), (−0.220 +
0.509I, −0.220 − 0.509I, −1.458), (−0.224 + 0.362I, −0.224 − 0.362I, −0.808), (−0.229 + 0.283I, −0.229 −
0.283I, −0.541), respectively. We notice that the smaller 1 = 2 causes the acceleration in the coexistence of
fixed point to approach the stability. The reason is simply that larger 1 will reduce the value of the imaginer
component of the eigenvalues. Furthermore, this simulation proves our theoretical in the previous section that
1 and 2 do not change the stability of the coexistence. An example of these behaviors is shown in figure 2.

FIGURE 2. Time series of the prey x(t), predator y(t), and scavenger z(t) with various 1 and 2 .

2. Varying 1
In this simulation, we will show how the dynamic of the system (2) behaved when 1 varying, while 2 re-
mains constant. It is observed that the system (2) approaches asymptotically to the coexistence fixed point Ω4 .
When 1 increasing, x(t), y(t), and z(t) are plotted individually versus t, we found that the periodic variation
in predator y(t) and scavenger population z(t) slightly behind the prey population x(t). This result also based
on the eigenvalues of each scenario. We found that the eigenvalue of the system (2) in Ω4 when 1 = 0.1 are
−0.174+1.037I, −0.174−1.037I, −0.650, when 1 = 0.4 are −0.199+.0509I, −0.199−0.509I, −0.600, when 1 =

060005-6
 0.7 are −0.216+0.366I, −0.216−0.366I, −0.567 and when 1 = 1 are −0.229+0.283∗I, −0.229−0.283I, −0.541.
It can be seen that when 1 is getting bigger, then the real component is increasing, which will accelerate the
time to reach the Ω4 . On the other hand, increasing of 1 will reduce the imaginary part of the eigenvalues,
which will reduce the periodicity of the system to reach Ω4 . To be clear, the dynamic of system (2) with this
scenario is shown in Figure 3.

FIGURE 3. Time series of the prey x(t), predator y(t), and scavenger z(t) with various 1 .

3. Varying 2
Similar to the previous simulation, this simulation is given to understand how the time scale of scavenger affects
the dynamic of the system (2), while the time scale for predator remains constant. The numerical result respect
to the eigenvalues for this scenario behaved contrast with the previous scenario when 1 was varying while 2
constant. In this scenario, increasing the value of 2 will reduce the value of the real part of the eigenvalue, but
increase the value of the imaginary part. To be exact, the eigenvalues for this scenario are 1 = 0.1 are −0.230 +
0.260I, −0.230 − 0.260I, −5.939, when 1 = 0.4 are −0.233 + 0.266I, −0.233 − 0.266I, −1.433, when 1 = 0.7
are −0.233 + 0.275I, −0.233 − 0.275I, −0.790 and when 1 = 1 are −0.229 + 0.283I, −0.229 − 0.283I, −0.541. It
means that shorter the life span of scavenger (2 smaller) will accelerate the time to reach the stable equilibrium,
but increase the periodicity of the model. This result present in Figure (4).

FIGURE 4. Time series of the prey x(t), predator y(t), and scavenger z(t) with various 2 .

CONCLUSION
Harvesting is one of the populations among several populations in the same ecosystem is a very crucial factor in
guaranteeing the coexistence of all populations in the ecosystem. Here in this paper, we try to show how the harvest-
ing might affect the behavior of the dynamic among interaction between prey, predator, and scavenger. Our results
from the analytical and numerical approach support the previous hypothesis that harvesting in prey population un-
controlled can end up in extinction on the predator, scavenger, or even on all populations in the ecosystem. From the
non-dimensionalization approach, the time scale parameter which describes the life span differences between prey,

060005-7
predator, and scavenger appears. Although this time scale parameter does not change the existence or criteria of the
stability of all fixed points, they can accelerate/decelerate and increase/decrease the periodicity of the system to reach
a stable equilibrium.

ACKNOWLEDGMENTS
We thank all reviewers for their valuable comments. This research is funded by Universitas Indonesia with QQ re-
search project (ID:NKB-0268/UN2.R3.1/HKP.05.00/2019).

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