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The Evolution of Stability in a Stochastic Predator-Prey System

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This master's thesis aim to see if fluctuations in the environment can make a stable predator-prey system naturally evolve into an unstable system. In [1] Abrams and Matsuda uses numerical methods to investigate the stability properties of a predator-prey system and looks at the effect that the unstable population dynamics have on the mean population sizes and the mean predation pressure. After re-deriving some of their main results, I introduce white noise in the prey equation to model the fluctuations which occur in e.g. the weather. Using both algebraic and numerical methods I present how different sets of parameters will affect the evolution of the stability of the system. The evolution of the prey population is assumed to select through the trait that codes for the prey's vulnerability. The parameter describing the prey vulnerability is therefore used in the evaluation of the direction of the evolution, which is done using techniques from Adaptive Dynamics.Unlike [1], I choose to separate the time scales of which the population dynamics and evolution occur on and evaluate the direction of the evolution by looking at the selection gradient. It is found out that the system can change its stability properties if the fluctuations reach a certain level. Based on numerical simulations, this level of fluctuations seems possible to exist in real ecological systems. The unstable evolution is possible since cycling dynamics have a negative effect on the mean predator population and positive on the prey dito.

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  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

    http:///reader/full/the-evolution-of-stability-in-a-stochastic-predator-prey-sys

  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

    http:///reader/full/the-evolution-of-stability-in-a-stochastic-predator-prey-sys

  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

    http:///reader/full/the-evolution-of-stability-in-a-stochastic-predator-prey-sys

  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

    http:///reader/full/the-evolution-of-stability-in-a-stochastic-predator-prey-sys

  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

    http:///reader/full/the-evolution-of-stability-in-a-stochastic-predator-prey-sys

  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

    http:///reader/full/the-evolution-of-stability-in-a-stochastic-predator-prey-sys

  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

    http:///reader/full/the-evolution-of-stability-in-a-stochastic-predator-prey-sys

  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

    http:///reader/full/the-evolution-of-stability-in-a-stochastic-predator-prey-sys

  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

    http:///reader/full/the-evolution-of-stability-in-a-stochastic-predator-prey-sys

    n

    n + 1

    n

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  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

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    G

    N

    G = N /N

  • 5/19/2018 The Evolution of Stability in a Stochastic Predator-Prey System

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    N=fN(N, P)

    P =fP(N, P)

    N

    P

    100

    2

    1

    2

    fN fP

    fN fP

    fN =N[r(C) kN CP (C, N)]

    fP =P[BCN(C, N) d].

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    C

    C >0

    r(C) =R + qC

    R,q >0

    (C, N) = 1/(1+hCN)

    h

    CN (C, N)

    CP (C, N)

    B

    r(C) N

    d P

    N P

    N =N

    R+ qC kN

    CP

    1 + hCN

    P =P

    BC N

    1 + hCN d

    ,

    (C, N) = 1

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    C

    C

    d

    h

    q

    B

    k

    R

    C C

    d d

    h h

    q q

    C =B

    kC, d =

    1

    Rd, h =

    R

    Bh,

    q =

    k

    BRq

    N

    P

    N =N R/k

    P = PBR/k

    t = t/R

    N=N

    1 + qC N

    CP

    1 + hCN

    P =P CN

    1 + hCN d ,

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    G

    C

    G(C, C) =N(C)

    N(C)= 1 + qC N(C)

    CP(C)

    1 + hCN(C).

    C

    C

    G(C, C) = 1 + qC N(C) CP(C)

    1 + hCN(C).

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    G(C, C)

    G(C, C)

    C = C

    C > C

    C < C

    G(C=C, C)

    G(C, C)

    C

    C

    G/C

    g

    g(C) =d C(1 dh)

    C2(1 dh) .

    C

    g(C)

    x

    C = d

    1 dh,

    G(C, C)

    C

    2G

    C2

    C=C

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    dg

    dC

    C=C

    =

    1 dh

    d

    2,

    C

    C

    C

    C

    C

    C

    C

    C < C

    C =C

    C > C

    g(C) > 0

    C

    C

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    0

    The invasion fitness evaluated at a specific C

    Vulnerability, C

    G

    (C,

    C

    )

    C

    G

    C < C

    g(C) > 0

    0

    The invasion fitness evaluated at an ESS

    Vulnerability, C

    G(C,

    C

    )

    ESS

    C

    ESS

    G C =C g(C) = 0

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    G

    g

    G= 1 + qC N( )

    CP( )

    1 + hCN( )

    g= q P(

    )

    1 + hCN( )

    .

    N( ) P( )

    P(t)

    1/(1 +hCN(t))

    C

    C

    =

    P( )

    1 + hCN( )

    .

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    (Ni, P

    i)

    (N1 , P1 ) =

    d

    C(1 dh),1 + hCN

    1

    C (1 + qC N1 )

    ,

    (N2 , P2 ) = (1 + qC, 0)

    (N3 , P3 ) = (0, 0).

    dh < 1

    1 + qC > N1

    d

    d < C(1 + qC)1 + hC+ hqC2 ,

    N

    P

    (N, P)

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    (N, P) =

    1 + qC 2N

    CP

    (1 + hCN)2

    CN

    1 + hCNCP

    (1 + hCN)2CN

    1 + hCN d

    .

    (N, P)

    J22

    C

    Ci

    Ci = 1

    2q1 +

    4q

    h

    1 + dh

    1 dh 1 .

    C Ci (N, P)

    1 2

    C

    Cd

    Ch

    Cq

    (N, P)

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    0 5 10 15 2010

    5

    0

    5

    10

    Eigenvalues of the Jacobian as a function of the prey vulnerability

    Vulnerability, C

    Re{1}Im{1}

    Re{2}Im{2}

    C

    1.2 1.4 1.6 1.8 2

    2

    1.5

    1

    0.5

    0

    0.5

    1

    genva ues o t e aco an as a unct on o t e prey vu nera ty

    Vulnerability, C

    Re{1}Im{1}Re{2}Im{2}

    C

    {2}

    1.05

    C

    C = Ci

    C > 1.05

    1.05 < C 1.35

    1.35 < C Ci

    Ci C 9.45

    C >9.45 {1}

    d= 1.0

    h= 0.5

    q= 0.85

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    w(t)

    w= w(t + t) w(t)

    w

    t

    w

    2t

    w(t)

    t 0

    dw

    2dt

    dt

    t

    dw

    dw= AN,

    0< A 0.2

    dt

    N

    dw

    dN=fNdt + dw

    dP =fPdt.

    x1 x2

    x1= N N

    x2= P P.

    d =

    dt + d

    = [x1 x2]

    T

    d

    = [dw 0]T

    g= g0+ gNx1+ g

    Px2+

    1

    2gN,Nx

    21+ g

    N,Px1x2+

    1

    2gP,Px

    22.

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    g

    g= g0+ gNx1+ g

    Px2+

    1

    2gN,NV(x1)

    + gN,PC(x1, x2) +1

    2gP,PV(x2).

    dx1 dx2

    d

    +

    T

    + = 0.

    V11 = V(x1)V22 = V(x2) V12 = V21 = C(x1, x2)

    211 dw

    V(x1) = 2

    11

    2J11,

    V(x2) = J21

    J12V(x1)

    C(x1, x2) = 0.

    V(x1) J11

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    0 =J11x1+ J12x2+1

    2(fN)

    N,NV(x1)+

    + (fN)N,PC(x1, x2) +

    1

    2(fN)

    P,PV(x2)

    0 =J21x1+ J22x2+1

    2(fP)

    N,NV(x1)+

    + (fP)N,PC(x1, x2) +12(fP)P,PV(x2).

    J22 = (fN)

    P,P = (fP)

    P,P = 0

    J11x1+ J12x2+

    hC2P

    (1 + hCN)3 1

    V(x1) = 0

    J21x1 hC2P

    (1 + hCN)3V(x1) = 0.

    x1= C2hP

    (1 + hCN)3V(x1)

    J21

    x2= 1

    J12

    1

    hC2P

    (1 + hCN)3

    V(x1) J11x1

    .

    g= q P

    1 + hCN+

    hCP

    (1 + hCN)2x1

    1

    1 + hCNx2

    2h2C2P

    (1 + hCN)3V(x1).

    x21 x

    22 x1x2

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    Ni+1= Ni+ Ni

    1 + qC Ni

    CPi1 + hCNi

    dt + dwi

    Pi+1= Pi+ Pi

    CNi

    1 + hCNi d

    dt.

    N(

    )

    P(

    )

    gj Cj

    gj =q P( )

    1 + hCjN( ).

    C

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    d= 1.0

    h= 0.5

    q= 0.85

    dt= 103

    C

    C

    Ci = 2.1329

    C Ci

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    0 20 40 60 80 1000.5

    1

    1.5

    2

    Population densities as functions of time

    Time, t

    N(t)P(t)

    1.3 1.4 1.5 1.6 1.70

    0.5

    1

    1.5

    Phase portrait

    N(t)

    P(t)

    C= 1.2

    0 10 20 30 400.5

    1

    1.5

    2

    Population densities as functions of time

    Time, t

    N(t)P(t)

    0.8 1 1.2 1.4 1.6

    1.4

    1.6

    1.8

    2

    Phase portrait

    N(t)

    P(t)

    C= 1.9

    0 20 40 600

    1

    2

    3

    4

    Population densities as functions of time

    Time, t

    N(t)P(t)

    0 1 2 30

    1

    2

    3

    4

    Phase portrait

    N(t)

    P(t)

    C= 2.4

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    3 4 5 60

    0.5

    1

    1.5

    2

    2.5

    3

    Mean and equilibrium densities as functions of the vulnerability

    Vulnerability, C

    N(t)N

    P(t)P

    Ci

    C

    [1.055, 6.4]

    N

    P

    N(

    )

    P(

    )

    2 3 4 5 60

    0.2

    0.4

    0.6

    0.8

    1

    The MPP as a function of prey vulnerability

    Vulnerability, C

    MPP(N(t), P(t))MPP(N, P)

    C Ci

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    C

    C >5.92

    C= 2.70

    C

    C

    = 2.3

    C

    N

    P

    C

    C

    C

    C < C

    C < C

    C

    C

    C =C

    C

    G(C, C)

    C

    CC

    C

    C = C

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    1.2 1.4 1.6 1.8 2

    0.1

    0.05

    0

    0.05

    0.1

    0.15

    0.2

    0.25 The invasionfi

    tness evaluated atfi

    xed C

    s

    Vulnerability, C

    G(C,

    C

    =

    constant)

    C =1.05

    C =

    1.10

    C =

    1.15

    C =

    1.20

    C =

    1.25

    C =

    1.30

    C =

    1.35

    C =1

    .45

    C =1

    .50

    C =1

    .40

    C =1.5

    5

    C =1.6

    0

    C =1.65

    C

    = 1.70

    C

    = 1.75

    C

    = 1.80

    C

    = 1.85

    C = 1.90

    C

    = 1.95

    C

    = 2.00

    C

    = 2.05

    C

    = 2.10

    C

    C

    C

    1.5 2 2.5 3

    1.5

    2

    2.5

    3

    C =C

    C

    =

    CESS

    g(C

    )=0

    g(C

    )=0

    The invasion fitness in the CC-plane

    Resident vulnerability, C

    Mu

    tantvulnerability,C

    + - +

    - + -

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    2 3 4 5 60.4

    0.2

    0

    0.2

    0.4

    0.6

    0.8

    1

    The selection gradient as a function of the prey vulnerability

    Vulnerability, C

    g(N(t), P(t))g(N, P)

    x

    C= C

    C

    C Ci

    g(C)

    C < C

    C > Ci

    g(C)

    x

    C

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    0 1 2 3 4 5 60

    0.5

    0.85

    1

    1.5

    2

    2.5

    The stability areas of the different parameter spaces

    Vulnerability, C

    Cq-borderCd-borderCh-border

    Cd

    Ch

    Cq

    y

    q= 0.85

    d= 1.0

    h= 0.5

    C= 2

    C = 2

    C

    2.1329

    {2} = 0 C = 1.0546 C

    C > 1.0546

    d

    h

    q

    d

    C

    h

    q

    Cd

    Ch

    Cq

    q

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    C

    q

    Ci

    C

    C < Ci

    q

    C

    qi

    C =Ci

    qi =1 dh

    d2h ,

    d h C

    Ci

    d

    h

    C

    Ci

    d

    h

    C

    Ci

    C

    C

    Ci

    g

    C =Ci

    q

    A

    A

    A

    A

    A

    C

    A

    A

    C

    C

    A

    0.0 A 0.12

    0 t 103

    t= 103

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    Ci

    0.0 A 0.10

    A 0.16 A 0.17 A= 0.12

    Ci

    A

    Ci

    C

    Ci

    C

    A = 0.12

    C

    0.555 105 Ci

    2.257 105

    C

    Ci

    g

    C

    C Ci

    gP g

    N,N g

    N

    gN,P

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    A C < C i C Ci C Ci

    1.8 C 2.3

    0 t 103

    t = 103

    Ci

    A

    Ci 105

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    1.

    8

    1.

    9

    2

    2.

    1

    2

    .2

    2.

    3

    0.

    04

    0.

    020

    0.

    02

    0.

    04

    0.

    06

    0.

    08

    0.

    1

    Theselec

    tiongradientwithd

    ifferentamountofw

    hitenoise

    Vulnera

    bility,

    C

    Theselectiongradient

    A

    =

    0.22

    A

    =

    0.20

    A

    =

    0.18

    A

    =

    0.16

    A

    =

    0.14

    A

    =

    0.12

    A

    =

    0.10

    A

    =

    0.08

    A

    =

    0.06

    A

    =

    0.04

    A

    =

    0.02

    A

    =

    0

    A

    A

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    C

    Ci

    C

    Ch

    h

    h

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    h

    h

    Ch

    C

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    A

    C

    0.01 A 0.06

    0.07 A 0.10 0.11 A 0.14

    0.15 A 0.17

    0.18 A 0.22

    A

    A Ci

    g(C)< 0

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    C

    C = 2.3

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