Harvesting in a Competitive

Environment
Eugene Wu
Math 360-2 Applied Analysis
February 2, 2015
In this project, we will extend our model of competition between two species. We will introduce harvesting of the two species into this system, where both species will lose members of their populations to this
harvesting effect. We will explore both the constant-effort model and the constand-yield model of
harvesting.

Constant Effort Harvesting I

(a) Let us consider a constant-effort model applied to only species x, with a positive constant E measuring harvest rate, given by
dx

= x(1 - E - x - y)
dt
dy
= y(0.75 - y - 0.5 x)
dt
For E = 0, we find the critical points where

dx
dt

dy
dt

(1)

= 0. These critical points occur at (0,0), (0, 0.75),

(1,0), and (0.5, 0.5).

For E 0, the critical points in terms of E become (0,0), (0, 0.75), (1 - E, 0), (0.5 - 2 E, 0.5 + E) . Thus,
as E increases, the critical points (0,0) and (0,0.75) stay the same, (1,0) shifts in the negative x direction
and (0.5,0.5) shifts in the negative x and positive y directions.
(b) We want to find the value of E = E0 such that the critical point at (0.5 - 2 E, 0.5 + E), coincides with
(0,0.75). Setting 0.5 - 2E = 0, we find that E0 = 0.25. This is consistent with the y-values, because 0.5 +
E = 0.5 + 0.25 = 0.75.
(c) Now, we will draw a direction field for the system when E = E 0 , and when E is slightly less than and
when E is slightly greater than E0 . The system at E = E0 is:
dx

= x(0.75 - x - y)
dt
dy
= y(0.75 - y - 0.5 x)
dt

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(2)

Harvesting in a Competitive Environment.nb

E>E0
1.0

0.8

0.6

0.4

0.2

0.0
0.0

0.2

0.4

0.6

0.8

1.0

E<E0

E= E0
1.0

1.0

0.8

0.8

0.6

0.6

0.4

0.4

0.2

0.2

0.0

0.0
0.0

0.2

0.4

0.6

(d) The Jacobian J of the system is

J(0, 0.75) =

0.8

1.0

0.0

0.2

0.4

0.6

0.8

1.0

1-E -2x-y
-x
. At (0, 0.75),
-0.5 y
0.75 - 2 y - 0.5 x

0.25 - E
0
. SInce this matrix is lower triangular, its eigenvalues are 1 = 0.25 - E
-0.375 -0.75

and 2 = -0.75. When E is slightly less than E0 = 0.25, 1 is positive and 2 is negative, making (0,0.75)
an unstable saddle point. When E > E0 , 1 and 2 are both negative. Then (0,0.75) changes from an
unstable saddle point to an asymptotically stable node as E passes through E0 .
(e) For E > E0 , species x goes extinct, unless the initial condition includes none of species y. Trajectories will all head toward (0,0.75) unless they begin on the x-axis, in which case they head toward
(1 - E,0). At E 1, x will certainly go extinct no matter what the initial conditions.

Constant Effort Harvesting II

Now the constant-effort model of harvesting only species x will be applied to a new system, given by

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Harvesting in a Competitive Environment.nb

dx
dt
dy
dt

= x(1 - E - x - y)
(3)

= y(0.8 - 0.6 y - x)

(a) For E = 0, we find the critical points where

dx
dt

dy
dt

= 0. These critical points occur at (0,0), (0,1.33),

(1,0), and (0.5,0.5).

For E 0, the critical points in terms of E become (0,0), (0, 1.33), (1 - E, 0), (0.5 + 1.5 E, 0.5 - 2.5 E).
Thus, as E increases, the critical points (0,0) and (0,1.33) stay the same, (1,0) shifts in the negative x
direction and (0.5,0.5) shifts in the positive x and negative y directions.
(b) We want to find the value of E = E0 such that the critical point at (0.5 + 1.5 E, 0.5 - 2.5 E), coincides
with (1 - E, 0). Setting 0.5 + 1.5E = 1 - E, we find that E0 = 0.2. This is consistent with the y-values,
because 0.5 - 2.5E = 0.5 - 0.5 = 0.
(c) Now, we will draw a direction field for the system when E = E 0 , and when E is slightly less than and
when E is slightly greater than E0 . The system at E = E0 is:
dx

= x(0.8 - x - y)
dt
dy
= y(0.8 - 0.6 y - x)
dt

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(4)

Harvesting in a Competitive Environment.nb

E=E0
1.5

1.0

0.5

0.0
0.0

0.2

0.4

0.6

0.8

1.0

E>E0

E<E0
1.5

1.5

1.0

1.0

0.5

0.5

0.0

0.0
0.0

0.2

0.4

0.6

0.8

1.0

0.0

0.2

0.4

0.6

0.8

1.0

The basin of attraction of the point (0,1.33) is shown above in blue, while the basin of attraction for the
point (1 - E, 0) is shown in green. The point (1 - E, 0) does not appear to be asymptotically stable
unless E < E0 . There are no other asymptotically stable points.
(d) The Jacobian J of the system is

1-E -2x-y
-x
. At (1 - E, 0), the critical point
-y
0.8 - 1.2 y - x

originally located at (1,0), J(1 - E, 0) =

E -1 E -1
. Since the matrix is upper triangular, its eigen0
E - 0.2

values are 1 = E - 1 and 2 = E - 0.2. When E < E0 = 0.2, 1 and 2 are negative. When E is slightly
greater than E0 , 1 is negative while 2 is positive. Thus, (1 - E, 0) changes from an asymptotically
stable nodal sink to an unstable saddle point as E passes through E 0 .

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Harvesting in a Competitive Environment.nb

(e) Species x will go extinct for E > E0 , because (1 - E, 0) is an unstable saddle point, and all trajectories will head towards (0,1.33) unless the initial conditions include only species x, in which case species
x will head toward the critical point (1 - E, 0). if E 1, then species x will go extinct no matter what the
initial condition is.

Constant Effort Harvesting III

Now consider a system where both species x and species y are harvested, given by:
dx
dt
dy
dt

= x(1 - E1 - x - y)
(5)

= y(0.75 - E2 - y - 0.5 x)

(a) When E1 = E2 = 0, the point (0.5,0.5) is an asymptotically stable node. We want to find the conditions
on E1 and E2 that permit the long-term survival of both species. By setting

dx
dt

dy
dt

= 0, we find the

critical points in terms of E to be (0,0), (0, 0.75 - E 2 ), (1 - E1 , 0), (0.5 - 2 E1 + 2 E2 , 0.5 + E1 - 2 E2 ).

Because all other critical points lie on the axes where only one population exists, we want to find conditions on E1 and E2 such that (0.5 - 2 E1 + 2 E2 , 0.5 + E1 - 2 E2 ) is asymptotically stable.

Critical Point: Both Species x and y

The Jacobian J of the system is given by

1 - E1 - 2 x - y
-x
. Then the Jacobian
-0.5 y
0.75 - E2 - 2 y - 0.5 x

at the critical point in question is

J(0.5 - 2 E1 + 2 E2 , 0.5 + E1 - 2 E2 ) =

-0.5 + 2 E1 - 2 E2 -0.5 + 2 E1 - 2 E2
. The locally linear system
-0.25 - 0.5 E1 + E2 -0.5 - E1 + 2 E2

at the critical point (0.5 - 2 E1 + 2 E2 , 0.5 + E1 - 2 E2 ) is given by u=Ju. The characteristic equation is:
(-0.5 + 2 E1 - 2 E2 - ) (-0.5 - E1 + 2 E2 - ) - (-0.5 + 2 E1 - 2 E2 ) (-0.25 - 0.5 E1 + E2 ) = 0
0.25 + 0.5 E1 - E2 + 0.5 - E1 - 2 E1 2 + 4 E1 E2 - 2 E1 + E2 + 2 E1 E2 - 4 E2 2 + 2 E2 + 0.5 + E1 2 E2 + 2 - 0.125 + 0.25 E1 - 0.5 E2 - 0.5 E1 - E1 2 + 2 E1 E2 + 0.5 E2 + E1 E2 - 2 E2 2 = 0
0.125 - 0.25 E1 + - E1 2 + 3 E1 E2 - E1 + 2 - 2 E2 2 = 0
2 + (1 - E1 ) + 0.125 - 0.25 E1 - E1 2 - 2 E2 2 + 3 E1 E2 = 0
=

(1 - E1 )2 - 4 0.125 - 0.25 E1 - E1 2 - 2 E2 2 + 3 E1 E2

-1 + E1

2
-1 + E1

E1 2 - 2 E1 + 1 - 0.5 + E1 + 4 E1 2 + 8 E2 2 - 12 E1 E2
2

-1 + E1

5 E1 2 - E1 + 0.5 - 12 E1 E2 + 8 E2 2
2

In order for both species to survive, the critical point must be asympotically stable, so both eigenvalues
must be negative. We can ensure that this is satisfied by setting
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Harvesting in a Competitive Environment.nb

-1 + E1 + 5 E1 2 - E1 + 0.5 - 12 E1 E2 + 8 E2 2 < 0. To check for the case that the eigenvalues are
complex, we enter the expression under the radical into a solver and find that it is never negative. Thus,
the eigenvalues are not complex.
Solve5 x2 - x + 0.5 - 12 x * y + 8 y2 < 0, {x, y}, Reals
Solve::ratnz : Solve was unable to solve the system with inexact coefficients.
The answer was obtained by solving a corresponding exact system and numericizing the result.

{}

(b) Next, we want to more analyze other possible scenarios where only one species survives, or where
no species survive. This will be made easier by finding an expression for the eigenvalues correspnding
to the locally linear system at each critical point. The work for this is shown below.

Critical Point: Species x

The critical point (1 - E1 , 0) corresponds to the existence of only species x. The Jacobian J of the
E1 - 1
E1 - 1
. Because this matrix is upper triangular,
system is given by J(1 - E1 , 0) =
0
0.25 + 0.5 E1 - E2
its eigenvalues are 1 = E1 - 1 and 2 = 0.25 + 0.5 E1 - E2 .

Critical Point: Species y

The critical point (0, 0.75 - E2 ) corresponds to the existence of only species y. The Jacobian J of the
0.25 - E1 + E2
0
. Because the matrix is upper triangusystem is given by J(0, 0.75 - E2 ) =
-0.375 + 0.5 E2 -0.75 + E2
lar, its eigenvalues are 1 = 0.25 - E1 + E2 and 2 = -0.75 + E2 .

Critical Point: No Species

The critical point (0,0) corresponds to the absence of both species x and species y. The Jacobian J of
1 - E1
0
. Because the matrix is triangular, its eigenvalues
the system is given by J(0,0) =
0
0.75 - E2
are 1 = 1 - E1 and 2 = 0.75 - E2 .

Constant-Effort Harvesting Plot

As found in part (a), -1 + E1 + 5 E1 2 - E1 + 0.5 - 12 E1 E2 + 8 E2 2 < 0 is the condition that must be
satisfied in order for both species to survive.
Now, let us first find the condition on E1 and E2 that allows for the survival of either one or the other
species. For species x to survive, the critical point (0, 0.75 - E 2 ) must be asymptotically stable, so both
of the corresponding linear systems eigenvalues must be negative. For species y to survive, the critical
point (0, 0.75 - E2 ) must be asymptotically stable, so both of the corresponding linear systems eigenval-

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Harvesting in a Competitive Environment.nb

ues must be negative.

In order for both species to perish, the critical point at (0,0) must be asymptotically stable, so both of the
corresponding linear systems eigenvalues must be negative.
The region in the E1 E2 -plane that corresponds to the long-term effect on both of the species is plotted
below.
1.4

1.2

1.0

x and y survive

0.8
E2

x survives
y survives

0.6

NO SURVIVORS!!!

0.4

0.2

0.0
0.0

0.2

0.4

0.6

0.8

1.0

1.2

1.4

E1

Nothing can be said of the point (E1 , E2 ) = (1, 0.75), because this is the point at which all critical points
converge on the origin and take on zero eigenvalues. Locally linear systems cannot give us any insight
into the behavior of the non-linear system in this case. However, a direction field of this scenario is
provided below, and it suggests stablity at the origin, if not asymptotic stability.

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Harvesting in a Competitive Environment.nb

E= (1,0.75)
1.0

0.8

0.6

0.4

0.2

0.0

0.0

0.2

0.4

0.6

0.8

1.0

Constant Yield Harvesting

A constant-yield model of harvesting will reduce the rate of population change by some positive constant that is not dependent on the current population size. A system where constant-yield harvesting is
used only on species x is given by:
dx

= x(1 - x - y) - H
dt
dy
= y(0.75 - y - 0.5 x)
dt
(a) If H = 0, the nullclines (where

dx
dt

(6)

= 0) are x = 0 and y = 1 - x. If H 0, then the nullclines occur where

x(1 - x - y) - H = 0. Then,
x - x2 - xy - H = 0
This equation takes the polynomial conic form of Ax2 + Bxy + Cy2 + Dx + Ey + F = 0. We can identify
what type of conic it is by looking at B2 - 4 AC. Because B2 - 4 AC = (-1)2 - 4 * -1 * 0 = 1 > 0 in this case,
the conic is a hyperbola. This is also shown below by plotting the equation, where we arbitrarily set H =
0.1. We also plot the lines x = 0 and x + y = 1, and see that these are indeed the asymptotes of the
hyperbola.

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Harvesting in a Competitive Environment.nb

x-nullcline
x=0
x+y=1
y-nullcline

-2

-2

(b) To find the critical points, we set

dx
dt

dy
dt

= 0. When H = 0, the critical points are (0,0), (0,0.75), (1,0),

and (0.5,0.5). Now, let H 0. The y-nullclines of y = 0 and y = 0.75 - 0.5 x remain the same. When the xnullcline intersects y = 0, x2 - x + H = 0. By the quadratic formula, the critical point(s) is

1-4 H
2

,0 ,

corresponding with (0,0) and (1,0). When the x-nullcline intersects y = 0.75 - 0.5 x,
0.5 x2 - 0.25 x + H = 0. By the quadratic formula, the critical point(s) is
0.25

0.0625 - 2 H , 0.625 0.5

0.0625 - 2 H , corresponding with (0.5,0.5) and (0,0.75). As H

increases, the critical points originally at (0,0) and (1,0) move towards each other, and eventually meet
at (0.5,0) when H = 0.25. Both critical points disappear when H exceeds 0.25. As H increases, the
critical points
originally at (0,0.75) and (0.5,0.5) both move toward (0.25,0.625), which they reach when H = 0.03125.
Both critical points disappear when H exceeds 0.03125.
(c) As stated above, when H = Hc = 0.03125, the critical points originally at (0,0.75) and (0.5,0.5) coincide at the point (0.25,0.625).
(d) As stated in part (b), when H > Hc , both critical points originally at (0,0.75) and (0.5,0.5) disappear.
Now consider the other two critical points,
system, J, is

1-

1-4 H
2

1-

1-4 H
2

, 0 and

1+

1-4 H
2

1-2x-y
-x
. Then, the Jacobian at
-0.5 y
0.75 - 2 y - 0.5 x

,0 =

1-4H
0

-1+

1-4 H
2

, 0 is

1-4 H
2

2+

1-

, 0 . The Jacobian of the

. Because this matrix is upper-triangular, its eigenvalues are

1-4 H
4

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10

Harvesting in a Competitive Environment.nb

1 - 4 H and 2 =

1 =
1-

1-4 H
2

1+

1-4 H
4

. For Hc < H < 0.25, these eigenvalues are positive, and thus

, 0 is an unstable node. The Jacobian at the other critical point is

1-4 H
2

2+

,0 =

- 1-4H

-1-

2-

1 = - 1 - 4 H and 2 =

1-4 H
2

. Because the matrix is upper-triangular, its eigenvalues are

1-4 H
4

2-

1-4 H
4

. For Hc < H < 0.25, these 1 < 0 and 2 > 0, so

1+

1-4 H
2

, 0 is an

unstable saddle point.

(e) Species x, unless the initial conditions include no members of species y, will go extinct. Both critical
points on the x-axis are unstable. There are no other critical points, so we will examine the systems
behavior on the y-axis to determine what happens to species y. On the y-axis,

dy
dt

= y(0.75 - y). Thus,

dy
dt

is positive under y = 0.75 and negative above it. Thus, the population of y will tend towards 0.75 unless
it starts with 0 members.
(f) Now, we will draw a direction field for the system when H = Hc , and when H is slightly less than and
when H is slightly greater than Hc . The system at H = Hc is:
dx

= x(1 - x - y) - 0.03125
dt
dy
= y(0.75 - y - 0.5 x)
dt

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(7)

Harvesting in a Competitive Environment.nb

H>Hc
1.5

1.0

0.5

0.0
0.0

0.2

0.4

0.6

0.8

1.0

H=Hc

H<Hc
1.5

1.5

1.0

1.0

0.5

0.5

0.0

0.0
0.0

0.2

0.4

0.6

0.8

1.0

0.0

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0.2

0.4

0.6

0.8

1.0

11