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    The Annals of Applied Probability2001, Vol. 11, No. 4, 12631291

    ASYMPTOTIC ANALYSIS AND EXTINCTION INA STOCHASTIC LOTKAVOLTERRA MODEL

    By F. C. Klebaner1 and R. Liptser1

    University of Melbourne and Tel Aviv University

    A stochastic LotkaVolterra model is formulated by using the semi-

    martingale approach. The large deviation principle is established, and is

    used to obtain a bound for the asymptotics of the time to extinction of prey

    population. The bound is given in terms of past-dependent ODEs closely

    related to the dynamics of the deterministic LotkaVolterra model.

    1. Introduction. Main result.

    1.1. Deterministic LotkaVolterra system. The LotkaVolterra system of

    ordinary differential equations [Lotka (1925) and Volterra (1926)],

    xt = xt xtytyt = xtyt yt

    (1.1)

    with positive x0 y0 and positive parameters describes a behavior ofa predatorprey system in terms of the prey and predator intensities xt andyt. Here, is the rate of increase of prey in the absence of predator, is a rateof decrease of predator in the absence of prey while the rate of decrease in

    prey is proportional to the number of predators yt, and similarly the rate ofincrease in predator is proportional to the number of prey xt [see, e.g., May(1976)]. The system (1.1) is one of the simplest nonlinear systems.

    Since the population numbers are discrete, a description of the predator

    prey model in terms of continuous intensities xt yt is based implicitly ona natural assumption that the numbers of both populations are large, andthe intensities are obtained by a normalization of population numbers by a

    large parameter K. Thus (1.1) is an approximation, an asymptotic descriptionof the interaction between the predator and prey. Although this model may

    capture some essential elements in that interaction, it is not suitable to answer

    questions of extinction of populations, as the extinction never occurs in the

    deterministic model; see Figure 1 for the pair xt yt in the phase plane.We introduce here a probabilistic model which has as its limit the determin-

    istic LotkaVolterra model, evolves in continuous time according to the same

    local interactions and allows evaluating asymptotically the time for extinction

    of prey species.

    There is a vast amount of literature on the LotkaVolterra model, and a his-

    tory of research on stochastic perturbations of this system both exact, approx-

    Received June 2000; revised January 2001.1Supported in part by Australian Research Council.

    AMS 2000 subject classifications. Primary 60I27, 60F10.

    Key words and phrases. Predatorprey models, large deviations, semimartingales, extinction

    1263

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    1264 F. C. KLEBANER AND R. LIPTSER

    Fig. 1. rx y when = 5, = 1, = 5, = 1.

    imate and numerical; see, for example, Goel, Maitra and Montroll (1971),

    Turelli (1977), Kesten and Ogura (1981), Hitchcock (1986), Watson (1987),

    Roozen (1989) and references therein. We approach the problem of extinc-

    tion via the theory of large deviations, thus obtaining new theoretical results,

    which previously were studied numerically.

    The system (1.1) possesses the first integral which is a closed orbit in the

    first quadrant of phase plane x y. It is given by

    rx y = cx logx + y logy + r0(1.2)where r0 is an arbitrary constant. It depends only on the initial points x0 y0(see Figure 1).

    1.2. Stochastic LotkaVolterra system. In this paper, we introduce and

    analyze a probabilistic model of preypredator population related to the clas-

    sical LotkaVolterra equations.

    Let Xt, and Yt be numbers of prey and predators at time t. We start withsimple balance equations for preypredator populations

    Xt = X0 + t t Yt = Y0 + t t

    (1.3)

    where:t is a number of prey born up to time t.t is a number of prey killed up to time t.t is a number of predators born up to time t.t is a number of predators died up to time t.

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1265

    We assume that t t

    t

    t , are double Poisson processes with the following

    random rates: Xt,

    KXtYt,

    KXtYt, Yt, respectively and disjoint jumps [the

    latter assumption reflects the fact that in a short time interval t t + tonly one prey might be born and only one might be killed, only one predator

    might be born and only one might die, with the above-mentioned intensities;

    moreover all these events are disjoint in time].

    The existence of such a model is not obvious; therefore in Section 2 we give

    its detailed probabilistic derivation.

    Assume X0 = Kx0 and Y0 = Kx0 for some fixed positive x0 y0 and alarge integer parameter K. Introduce the normalized by K prey and predatorpopulations

    xKt =XtK

    and yKt =YtK

    In terms ofxKt and yKt the introduced intensities for double Poisson processes

    can be written as

    KxKt x

    Kt y

    Kt x

    Kt y

    Kt y

    Kt

    We justify the choice of the probabilistic model given in (1.3) by Theorem 2

    which states that the solution of the LotkaVolterra equations is the limit (in

    probability) for xKt yKt ,xKt y

    Kt

    xt yt K Such an approximation is known as the fluid approximation. Results on the

    fluid approximation for Markov discontinuous processes can be found in Kurtz

    (1981) and are adapted to the case considered here, despite that in our case

    the two intensities do not satisfy the linear growth condition in xKt yKt .

    1.3. Fomulation of the problem. Here, we are interested in evaluation of

    the prey extinction, namely, the asymptotics in K of

    P

    TKext T inftT

    yKt > 0

    = 0

    where TKext = inft > 0 xKt = 0 is the prey extinction time. Unfortunately,the fluid approximation does not provide much information on the extinction

    time TKext. Since for x0 > 0, y0 > 0 the fluid limit xt yt remains positivefor any t > 0 (see Figure l), we have

    limK

    P

    TKext T inftT

    yKt > 0

    = 0(1.4)

    A historical comment onT

    K

    extof evaluation is due. There is a large amount of

    literature on the subject of extinction mostly using some simplification of the

    original problem, such as linearization, and numerical studies. For a somewhat

    different model with a state-dependent noise, Hitchcock (1986) showed that

    ultimate extinction is certain, and derived exact probabilities for the predators

    to become eventually extinct when the prey birth rate is zero. A power series

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    1266 F. C. KLEBANER AND R. LIPTSER

    approximation for the extinction probabilities as well as the number of steps

    to extinction in special cases were given. Numerical studies of probabilities of

    extinction were also done in Smith and Mead (1979, 1980), and Watson (1987)where a rough approximation (based on the normal approximation) was also

    given.

    Due to (1.4), the rate of convergence in (1.4) is of interest, and is the subject

    of this paper. Neither fluid nor even diffusion approximations for the stretched

    differences

    K

    xKt xt

    K

    yKt yt

    are effective for such analysis.

    Freidlin (1998), Freidlin and Weber (1998) carried out an effective asymp-

    totic analysis for randomly perturbed oscillators and other Hamiltonian

    systems. Their approach is based on the approximation of the first integral

    process [in our case rxKt yKt ] by a scalar diffusion. However, in our case thisapproach does not seem to be of use, since at the time of extinction the first

    integral process rxKt yKt explodes; see (1.2).A large deviation (LD) type evaluation yields results in our case. The ran-

    dom process xKt yKt is a vector semimartingale, so that it appears that onecan have the large deviation principle (LDP) by using a general results from

    Pukhaskii (1999). However, the method from Pukhaskii does not serve the

    model studied here, since the intensities of two double Poisson processes are

    of quadratic form in xKt yKt . For the same reason we could not find adequate

    methods for proving the (LDP) in the literature, for example, Wentzell (1989),

    Dupuis and Ellis (1997), Freidlin and Wentzel (1984), Dembo and Zeitouni

    (1993).

    It is well known that the main helpful tool in verification of the LDP with

    unbounded intensities, satisfying a linear growth condition, is the exponen-

    tial negligibility for sets suptT xKt C, suptT yKt C, for large K, Cand every T > 0. In our case two from four intensities do not satisfy the lin-ear growth condition. Nevertheless, specifics of the model and the fact that

    xKt yKt are nonnegative processes with bounded jumps allow establishing the

    above-mentioned exponential negligibility (Lemma 2) and deriving the LDP

    similarly to Liptser and Pukhalskii (1992).

    Although in principle the LDP allows finding the logarithmic rate with

    norming 1K

    for

    P

    TKext T inftT

    yKt > 0

    a realistic procedure for determining the required rate depends heavily on the

    structure of the LDP rate function. In our case we deal with purely discontin-

    uous process and the rate function is extremely inconvenient for this purpose;

    therefore, we restrict ourselves to finding only a lower bound for the required

    rate.

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1267

    Fig. 2. Trajectory that attains the lower bound. Parameters are the same as for Figure 1.

    1.4. Main result. To formulate the main result, we need to introduce a

    system of past-dependent differential equations parameterized by q > 0,

    qt = qt

    qt

    qe t0 qt dsqt qt

    qt = qt

    qt

    (1.5)

    subject to the initial condition q0 = y0, q0 = x0, and denote

    Tq = inft > 0 qt = 0(1.6)

    Theorem 1. For every T > 0,

    liminfK

    1

    Klog P

    TKext T inf

    tTyKt > 0

    x

    20

    2Tq

    0 e2t0 qt dsqt qt dt

    where Tq is associated with the smallest q = q for which Tq T.1.5. Example. We give here an example with = 5, = 1, = 5, = 1 and

    T = 15. The trajectories for qt qt , on which the lower bound is attainable, aregiven in the phase plane (see Figure 2). Parameter q 00023 and Tq 15,while the value of the rate function defined later in (4.2) is Jq q 00018.

    1.6. Remark. We can show for a stochastic LotkaVolterra model with a

    different noise structure (much simpler and somewhat artificial) the exact

    asymptotic relation in Theorem 1 (rather than a lower bound), as well as that

    the minimum of the rate function in the LDP occurs on the solution similar

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    1268 F. C. KLEBANER AND R. LIPTSER

    to (1.5). Therefore, loosely speaking, (1.5) (see Figure 2) gives a likely path to

    extinction in the stochastic LotkaVolterra model.

    The article is organized as follows. In Section 2 we give description of thestochastic model, in Section 3 we show the fluid approximation, in Section 4

    we formulate the LDP and prove the main result. The verification of the LDP,

    which is quite technical, is done in the Appendix.

    2. The model: description of stochastic dynamics.

    2.1. Existence. In this section, we show that the random process Xt Ytis well defined by (1.3). To this end, let us introduce four independent

    sequences of processes,

    t =

    t 1 t 2

    /K

    t = /Kt 1 /Kt 2 /Kt =

    /Kt 1 /Kt 2

    t =

    t 1 t 2

    Each of them is a sequence of i.i.d. Poisson processes characterized by rates

    , K

    , K

    , , respectively. Define the processes Xt Yt by the system of Itoequations

    Xt = X0 +t

    0

    n1

    IXs n dsn t

    0

    n1

    IXsYs n d/Ks n

    Yt = Y0 + t

    0 n1IXsYs n d/Ks n

    t

    0 n1IYs n dsn

    (2.1)

    governed by these Poisson processes, which obviously has a unique solution

    on the time interval 0 T, whereT = inft > 0 Xt Yt =

    The double Poisson processes involved in (1.3) are obtained then in the follow-

    ing way:

    t =t

    0

    n1

    IXs n dsn

    t =

    t

    0 n1IXsYs n d/Ks n

    t =t

    0

    n1

    IXsYs n d/Ks n

    t =t

    0

    n1

    IYs n dsn

    (2.2)

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1269

    Let us show that t, t ,

    t,

    t defined in 22 satisfy the required prop-

    erties. Since all Poisson processes are independent, their jumps are disjoint,

    so that the jumps of t, t , t, t are disjoint as well. To describe structureof intensities for t,

    t ,

    t,

    t , let us introduce a stochastic basis F =

    tt0 P supplied by the filtration F generated by all Poisson processes andsatisfying the general conditions. Then, obviously, the random process

    At =t

    0

    n1

    IXs nds

    is the compensator of t. On the other hand, since Xs is an integer-valuedrandom variable, we have

    n1 IXs n = Xs; that is, At =

    t0 Xs ds and

    the intensity of t is Xt. Analogously, others compensators are seen to be

    At =t

    0

    KXsYs ds

    At =

    t0

    KXsYs ds

    At =

    t0Ys ds

    and thus all other intensities have the required form.

    We now show that the process Xt Ytt0 does not explode.Lemma 1.

    PT = = 1Proof. Set TXn = inft > 0 Xt n, n 1 and denote by TX =

    limn TXn . Due to (2.1) it holds that

    EXtTXn X0 +t

    0EXsTXn ds

    and so, by the GronwallBellman inequality EXTXn T X0eT for every T > 0.Hence by the Fatou lemma EXTXT X0eT. Consequently,

    P

    TX

    T =

    0

    T > 0

    Set TY = inft Yt , 1 and denote by TY = lim TY . Due to (2.1)it holds that

    EYtTXn TYt Y0 +t

    0

    KEXsTXn TY YsTXn TY ds

    Y0 +t

    0

    KnEYsTXn TY ds

    Hence, by the GronwallBellman inequality for every T > 0 we haveEYTTxnTy Y0enT and by the Fatou lemma,

    EYTTXn TY Y0e/KnT n 1Consequently, P

    TY

    TXn

    T

    =0

    T > 0, n

    1 and, since TXn

    ,

    n , we obtainPTY T = 0 T > 0

    Since T = TX TY, PT T = PTX T + PTY T, and wehave PT T = 0 for any T > 0.

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    1270 F. C. KLEBANER AND R. LIPTSER

    Corollary 1. For Tn = inft Xt Yt n,

    limn PTn T = 0 T > 0

    The above description of the model allows claiming that Xt Yt is acontinuous-time pure jump Markov process with jumps of two possible sizes

    in both coordinates: 1 and 1 and infinitesimal transition probabilities (ast 0,

    PXt+t = Xt + 1Xt Yt = Xtt + ot

    PXt+t = Xt 1Xt Yt =

    KXtYtt + ot

    PYt+t = Yt + 1Xt Yt =

    KXtYtt + ot

    PYt+t = Yt 1Xt Yt = Ytt + ot

    2.2. Semimartingale description for xKt yKt . Let At, At , At, At be thecompensators of t,

    t ,

    t,

    t defined above. Introduce martingales

    Mt = t At Mt = t At Mt = t At Mt = t At and also normalized martingales

    mKt =Mt Mt

    Kand mKt =

    Mt MtK

    (2.3)

    Then, from (1.3) it follows that the process xKt yKt admits the semimartin-gale decomposition

    xKt = x0 +t

    0

    xKs Ks xKs yKs

    ds + mKt (2.4)

    yKt = y0 +t

    0

    xKs y

    Ks yKs

    ds + mKt (2.5)

    which is a stochastic analogue (in integral form) of (1.1)

    In the sequel we need quadratic variations of the martingales in (2.4) and

    (2.5). It is well known [see, e.g., Liptser and Shiryaev (1998), Chapter 18 or

    Klebaner (1998), Theorem 9.3] that all martingales are locally square inte-

    grable and possess the predictable quadratic variations

    Mt = At Mt = At and Mt = At Mt = At (2.6)and zero mutual predictable quadratic variations M Mt 0 M Mt 0, implied by the disjointness of jumps for t, t , t, t .

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1271

    Hence

    mK

    m

    K

    t 0mKt =

    1

    K

    t0

    xKs + xKs yKs

    ds

    mKt =1

    K

    t0

    xKs y

    Ks + yKs

    ds

    (2.7)

    2.3. Stochastic exponential and cumulant function. In the large deviation

    theory we use stochastic exponential, and to this end the following represen-

    tation is more convenient:

    xKt = x0 =tK

    t

    K

    yKt = y0 = tK tK (2.8)

    tK

    ,tK

    ,tK

    ,tK

    are counting process with jumps of the unit size K1 and com-

    pensatorsAtK

    ,AtK

    ,AtK

    ,AtK

    , respectively. With every pair tK

    ,AtK

    , , tK

    ,AtK

    and a predictable process t such that for any T > 0 and K large enough,T

    0et/K

    xKt + yKt + xKt yKt

    dt < P-a.s.

    we associate nonnegative processes

    zt

    K = expt

    0

    sK

    ds

    es/K

    1

    dAs

    (2.9)

    zt

    K

    = exp

    t0

    sK

    ds es/K 1 d AsApplying the Ito formula to zt K, we find

    dzt

    K

    = zt

    K

    et/K 1 dt Atthat is, zt K is a local martingale [analogously zt K, zt K, zt K are localmartingales as well]. All these martingales are nonnegatives, so that they

    are supermartingales [see Problem 1.4.4 in Liptser and Shiryayev (1989) or

    Theorem 7.20 in Klebaner (1998)]. Hence for any Markov time we haveEz K 1 Ezt K 1.

    Introduce also (for predictable processes and )

    zKt = zt

    K

    zt

    K

    zt

    K

    zt

    K

    (2.10)

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    1272 F. C. KLEBANER AND R. LIPTSER

    Applying the Ito formula to zKt and taking into account that jumps oft,

    t ,

    t,

    t are disjoint we find

    dZKt = zKt et/K 1 dt At + et/K 1 dt At + et/K 1 d t At + et/K 1 d t At

    so that ZKt is a positive local martingale as well as a supermartingalewith

    EZK 1for any Markov time .

    Set

    G u v= vu + e 1 u + e 1 + uv

    G u v= u v + e

    1 uv + e

    1 + v(2.11)

    and define the so-called cumulant function,

    G u v = G u v + G u v(2.12)Using the cumulant function, (2.10) can be written as

    ZKt = expt

    0

    s dxKs +

    t0

    s dyKs

    expt

    0KG

    s

    K

    sK

    xKs yKs

    ds

    (2.13)

    Hence an equivalent multiplicative decomposition holds for the exponential

    semimartingale

    expt

    0

    s dxKs + s dyKs = ZKt VKt

    with the positive local martingale ZKt and a positive predictable process

    VKt = expt

    0KG

    s

    K

    sK

    xKs yKs

    ds

    3. Fluid approximation. In this section we justify the choice of thestochastic dynamics by showing that the LotkaVolterra equations describe

    a limit (fluid approximation) for the family

    xKt y

    Kt

    K

    In what follows xt yt is a solution of the LotkaVolterra equation (1.1).Theorem 2. For any T > 0 and > 0,

    limK

    P

    suptT

    xKt xt + yKt yt

    >

    = 0

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1273

    Proof. Set

    TKn =

    inft xKt yKt n(3.1)Since TKn = TnK (see Corollary 1 to Lemma 1),

    limn

    limsupK

    P

    TKn T = 0(3.2)

    Hence, it suffices to show that for every n 1,

    limK

    P

    sup

    tTKn T

    xKt xt + yKt yt

    >

    = 0(3.3)

    Since suptTKn TxKt yKt n+ 1, there is a constant Ln, depending on n andT, such that for t TKn T,

    xKt xKt yKt xt xtyt LnxKt xt+ yKt ytxKt yKt yKt xtyt yt LnxKt xt + yKt ytThese inequalities and (1.1), (2.4) implyxKTKn T xTKn T+ yKTKn T yTKn T

    2Lnt

    0

    xKsTKn xsTKn + yKsTKn ysTKn ds+ sup

    tTKn T

    mKt + suptTKn T

    mKt Now, by the GronwallBellman inequality we find

    suptTKn T

    xKt xt + yKt yt e2LnT suptTKn T

    mKt + suptTKn T

    mKt

    Therefore (3.3) holds, if both suptTKn T mKt and suptTKn T mKt convergein probability to zero as K . By the Doob inequality for martingales (see,e.g., Theorem 1.9.1(3) and Problem 1.9.2 in Liptser and Shiryayev (1989) the

    required convergence takes place provided that both mKTKn T and mKTKn Tconverge to zero in probability as K . But by (2.7) mKTKn T constK , mKTKn T constK , and the proof is complete.

    4. Formulation of the LDP and the proof of Theorem 1.

    4.1. LDP. The random process xKt yKt t0 has nonnegative paths fromthe Skorokhod space 2 = 20 . Since we are going to apply the LDPfor asymptotic analysis of the extinction time on a finite interval 0 T, weexamine the LDP in 20 T. Because of the fluid approximation, the limit forxKt yKt tT is xt yttT with continuous differentiable functions xt and yt.

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    1274 F. C. KLEBANER AND R. LIPTSER

    This fact allows examining the LDP in the metric space 20 T suppliedby the uniform metric

    x x

    2

    0 T

    and

    x x =2

    j=1suptT

    xtj xt j(4.1)

    More exactly, since xKt 0, yKt 0 andKx = inft > 0 xKt = 0 Ky = inft > 0 yKt = 0

    are also absorption points for processes xKt and yKt , respectively, let us intro-

    duce a subspace 2 +0 T of nonnegative function from

    20 T. It is obvious that

    2 +0 T is a closed subset of

    20 T in the metric .

    Therefore we formulate the LDP in metric space 2 +0 T .

    Theorem 3. For every T > 0, the family xK

    t yK

    t tT, K obeys theLDP in the metric space 2 +0 T with the rate of speed 1K and the (good)rate function

    JT=

    T

    0sup

    t +t Gtt

    dt dtdtdtdt

    0=x00=y0 ,

    otherwise.(4.2)

    The proof of the LDP is given in the Appendix.

    4.2. Proof of Theorem 1. Denote by o the interior of the set

    2 +0 T inf

    t

    T

    t = 0 inft

    T

    t > 0Due to the LDP,

    lim infK

    1

    KlogP

    inftT

    xKt = 0 inftT

    yKt > 0

    inf

    JT (4.3)

    It is clear that

    inf o

    JT inf o JT

    where is a subset of o of absolutely continuous functions t, t with0 = x0, 0 = y0 and for any = ttT,

    t = tt (4.4)To emphasize the fact that t is a solution of (4.4) we use the notation

    t .

    The function t remains positive for any finite time interval and moreover,

    sup

    t + t G t t

    = sup

    t G t t

    = sup

    t t t e 1 t e 1 + tt

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1275

    Taking into account that functions ts from have to be absorbed on 0 T,we choose a subset

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    1276 F. C. KLEBANER AND R. LIPTSER

    that is,

    t

    x0 pt. Now, for any T > 0, the choice p =

    x0T

    guarantees T T.If some ut is chosen and associated with T < T, it is obvious that ut 0

    for t T T. Then, particularly, in (4.6) the integral T0

    u2t dt is replacedby

    T0

    u2t dt. On the other hand, due to T = 0 we have

    0 = eT

    0 s ds

    x0

    T0

    et

    0 s dsut

    t

    t dt

    that is, x0 =T

    0e

    t0

    s dsut

    t

    t dt. Further, the CauchySchwarz

    inequality implies

    T

    0 u

    2

    t dt x20T

    0e2 t0 s dstt dt (4.7)

    Take any positive q and ut = qet

    0 s ds

    t

    t then the inequality given

    in (4.7) becomes an equality, and only for

    q = x0T0

    e2t

    0 s dst

    t dt

    (4.8)

    we have T = 0.Let q Tq be a pair such that (1.6) holds with q and T replaced by q and

    Tq respectively and Tq

    T. Denote by

    qt and

    qt the solution of (1.5) on

    0 Tq corresponding to the pair q Tq. Then we get the lower bound

    lim infK

    1

    Klog P

    TKext T inf

    tTyKt > 0

    inf x

    20

    2Tq

    0 e2 t0 s dsutqt qt dt

    where inf is taken over all Tq T .The final step of the proof uses the property

    q < q x2

    0Tq0 e

    2 t0 qs dsqt q dt C

    = (A.1)

    lim0

    lim supK

    1

    Klog sup P

    supt

    xKt+ xK + supt

    yKt+ yK > =

    (A.2)

    for any > 0, where sup is taken over all stopping times T. C local LDPis valid, if for any 20 T

    lim sup0 limsupK

    1

    K logPxK yK JT (A.3)

    lim inf0

    lim infK

    1

    KlogPxK yK JT (A.4)

    Since the linear growth condition for intensities of counting processes is

    lost, a verification of (A.1) requires a different technique from the one used in

    Liptser and Pukhalskii (1992). We verify (A.1) in the next subsection.

    A.2. Exponential negligibility of suptT xKt , suptT y

    Kt .

    Lemma 2. For every T > 0,

    limL

    lim supK

    1

    KlogP

    suptT

    xKt L =

    limC

    limsupK

    1

    KlogP

    suptT

    yKt C

    =

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    1278 F. C. KLEBANER AND R. LIPTSER

    Proof. Recall that xKt = x0 + ttK

    and Mt = t At; that is, dxKt dtK

    =xKt dt

    +dMt

    Kand so xKt

    et

    x0

    +1

    K t

    0es dMs

    . Consequently, the

    first statement of the lemma is valid provided that

    limL

    lim supK

    1

    KlogP

    suptT

    t0

    es dMs KL

    =

    It is clear that with KL = inftt

    0es dMs KL it suffices to establish

    limL

    lim supK

    1

    KlogP

    KL T

    = (A.5)

    To this end, with r > 0 and s = res we define zt by the first formula in(2.9) with

    sK

    replaced by s and use an obvious modification of this formula,

    zt = expt0 s dMs t0 es 1 s dAs(A.6)Recall that zt is a supermartingale with EzKL T 1. We use thisinequality for the next one,

    1 EzKL T

    IKL T(A.7)We sharpen that inequality, by evaluating below z

    KL T on the set KL T,

    logzKL T

    = rKL T

    0es dMs

    KL T0

    ere

    s 1 resKxKs ds

    rKL

    KL T

    0 er

    1

    r

    KxKs ds

    rKL T

    0er 1 rKxK

    sKLds

    To continue this evaluation we find an upper bound for xKsKL

    . Since

    KL T0

    esdMs KL + 1

    (recall that jumps oft

    0esdMs is bounded by 1), we claim

    xKKL t

    etx0 + 1 + L

    Hence, with p =eT

    x0

    +1

    +L

    L , we arrive at the lower boundlogz

    KL T KLr er 1 rTp

    It is clear that r > 0 can be chosen so that r er 1 rTp = > 0 andtherefore logz

    KL T KL > 0.

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1279

    Thus, (A.7) with s = res replaced by sres, implies1

    K logPKL T L L

    that is, (A.5) holds.

    The proof of the second statement of the lemma uses (A.5) heavily. SincesuptT

    yKt C

    suptKL T

    yKt C

    KL Tand thereby

    P

    suptT

    yKt C

    2

    P

    sup

    tKL TyKt C

    P

    KL T

    by virtue of (A.5) it suffices to check that for every fixed L,

    limC

    lim supK

    1K

    logP suptKL T

    yKt + C = (A.8)The verification of (A.8) is similar to the proof of the first statement of the

    lemma. It is clear that there is a positive constant RL so that IKL txKt RL, t 0. Further, due to dyKt xKt yKt dt + 1K dMt we have

    dyKtKL

    IKL txKt yKt dt +1

    KdM

    tKL

    RLyKt dt +1

    KdM

    tKL

    and hence yKt

    KL

    eRLt

    y0

    + tKL

    0 eRLd

    Ms

    Now, introduce

    KL C = inf

    ttKL

    0eRL dMs C

    and note that (A.8) holds provided that

    limC

    lim supK

    1

    KlogP

    KL C T

    = L > 0(A.9)The proof of (A.9) is similar to the proof of (A.5), so here we give only a sketch

    of it. Set = RL and for positive r take s = res. Introduce a super-martingale

    zt = expt

    0s d

    Ms

    t

    0 es 1 s

    d

    As

    Due to EzKL TKL CT

    1 write 1 EzKL TKL CT

    IKL C T and evaluatefrom below z

    KL TKL CT on the set KL C T as

    log zKL TKL CT

    KCr er 1 rTp

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    1280 F. C. KLEBANER AND R. LIPTSER

    With r such that r er 1rTp = > 0 similarly to the proof of thefirst statement of the lemma we obtain 1

    KlogPKL C T C and (A.9)

    holds.

    A.3. C-Exponential tightness. (A.1) is implied by Lemma 2.To verify (A.2), let us note that

    xKt+ xK =1

    K

    t+

    + 1K

    t+

    yKt+ yK =1

    K

    t+

    + 1K

    t+

    Evidently, it suffices to establish the validity of (A.2) for every

    1

    Kt+

    1

    Kt+

    separately.Recall that the Markov time TKn is defined in (3.1). By virtue of Lemma 2

    we claim that limn lim supK1

    KlogPTKn T = . Hence, with any

    n 1 we have to verify (A.2) only for

    limK

    1

    Klog sup

    TP

    +TKn TKn > K

    = 0

    limK

    1

    Klog sup

    TP

    +TKn TKn > K

    = 0

    The proofs for the above relations are similar. So, we give below only one of

    them for t . With r > 0 set

    s

    =rI

    TKn

    +

    TKn

    s

    . The random process

    s is bounded and left continuous (and thereby predictable). Consequently[compare (2.9)], the random process

    zt = expt

    0s ds

    t0

    es 1dAs

    is a supermartingale with EzT+ 1. Therefore,

    1 EzT+I

    +TKn TKn > K

    (A.10)

    Since A+TKn ATKn

    Kn + 12, on the set +TKn

    TKn > K,

    logzT+ = r

    +TKn

    TKn

    er 1

    A+TKn A

    TKn

    Kr er 1n + 12Now, with small enough and r = log

    n+12 ,

    zT+ exp

    K

    log

    n + 12

    + n + 12

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1281

    Hence

    1

    K log supT P+TKn TKn > K log

    n + 12

    + n + 12 0

    A.4. C-Local-LDP. Upper bound. Obviously, for 0 = x0 or 0 = y0 theupper bound is equal to . The proof that for 0 = x0 and 0 = y0 and 20T\20T the upper bound is equal to under the C-exponentialtightness can be found in Theorem 1.3 in Liptser and Pukhalskii (1992).

    Thus, the proof is concentrated on the case 20 T with 0 = x0,0 = y0. Let ZKt K K be defined in (2.13) with piecewise constant (deter-ministic) functions

    t

    and

    t

    . Since EZK

    T K K

    1, write

    1 EZKT

    K

    K

    I

    xK yK

    (A.11)

    On the set xK yK , we evaluate ZKt K K from below. Since are piecewise constant functions notations,

    T0

    t dtT

    0t dt will be

    used for tjT

    tj1tj tj1 and

    tiTti1ti ti1

    respectively. Obviously, a positive constant c can be chosen such that

    logZKT K K K

    c +

    T0

    t dt + t dt Gt t t t dt

    Therefore, this lower bound jointly with (A.11) imply

    lim sup0

    lim supK

    1

    KlogPxK yK

    T

    0t dt + t dt Gt t t t dt

    Since the left side of this inequality is independent oft t

    the required

    upper bound is obtained by minimization of the right side in t t. As inLiptser and Pukhalskii [(1992), Theorem 6.1], we find that for not absolutely

    continuous t or t the minimal value of the right-hand side is equal to ,while if both t t are absolutely continuous functions the minimal valuecoincides with JT .

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    1282 F. C. KLEBANER AND R. LIPTSER

    A.5. C-Local-LDP. Lower bound. It suffices to verify (A.4) only for JT, < . That supposes the verification of (A.4) for absolutely continuousfunctions t ttT from

    2+0 T with 0 = x0 0 = y0. Moreover, to satisfy

    JT < functions t t have to be absorbed on axis x = 0; y = 0(recall that xKt y

    Kt are absorbed on these axes). For definiteness we denote

    the class of the above-mentioned functions t t by .Set T = inft > 0 t = 0 and T = inft > 0 t = 0 inf = and

    introduce

    JT =

    TT0

    sup

    t G t t dt(A.12)

    JT =

    TT0

    sup

    t G t t dt(A.13)

    Then, obviously, we have JT = JT + JT . Below we givedetailed description of t = argmaxt G t t and t =argmaxt G t t for T T = ,

    t= log

    t2t

    +

    2t4t2

    + t

    t= log

    t2tt

    +

    2t4tt2

    + t

    (A.14)

    A.5.1. Main Lemma.

    Lemma 3. Assume

    (i) .(ii) T T = .

    (iii) t const t const

    Then there is a positive constant L , depending on L , such thatfor > 0 small enough,

    lim infK

    1

    K

    logP

    xK yK

    JT

    L

    Proof. Set K = inft xKt t and K = inft yKt t .Since jumps of xKt y

    Kt are bounded by

    1K

    , by the triangular inequality we

    have xKt t xKt t + 1K and yKt t yKt t + 1K . Consequently

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1283

    with K 4

    the inclusion holds:

    suptT xKt t + suptT yKt t

    suptT

    xKt t + suptT

    yKt t

    2

    =

    suptK T

    xKt t + suptK T

    yKt t

    2

    Hence, the first lower bound we use is the following:

    PxK yK

    P

    sup

    tK

    T

    xKt t + supt

    K

    T

    yKt t

    2

    (A.15)

    For K large enough set

    Kt= log

    t

    2xKt+

    2t

    xKt2+ x

    Kty

    Kt

    xKt

    t K T

    Kt= log

    t

    2xKtyKt

    +

    2t

    2xKtyKt2yKt

    xKtyKt

    t K T

    (A.16)

    [Kt Kt are defined similarly with an obvious change]. It is clear thatfor K large enough Kt Kt are bounded and strictly positive.

    We define now the positive supermartingale [compare (2.13)]

    ZKt K K= exp

    KtK T

    0

    Kt dxKt GKt xKt yKt dt

    exp

    K

    tK T0

    Kt dyKt GKt xKt yKt dt

    (A.17)

    We show that ZKt K KtT is the square integrable martingale withEZKT K K = 1

    To prove this property we have to check that EZKT K K2 < . (A.17)implies

    ZKT

    K K

    2

    =ZKT

    2K 2K

    exp

    K

    K T0

    G

    2Kt xKt yKt

    2GKt xKt ykt dt exp

    K

    K T0

    G

    2Kt xKt yKt

    2GKt xKt ykt dt

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    1284 F. C. KLEBANER AND R. LIPTSER

    The process ZKT 2K 2KtT is a supermartingale as well; that is

    EZ

    K

    T 2K

    2

    K

    1and at the same time under assumptions of the lemma G2Kt xKt yKt ,GKt xKt yKt and G2Kt xKt yKt GKt xKt yKt are boundedfunctions on the time intervals 0 K T and 0 K T, respectively. Con-sequently, ZKT K K2 has a finite expectation.

    Now, F = tt0 QK with dQK = ZKT K K dP is the stochasticbasis. Due to the positiveness of ZKT K K not only QK P but also P QK with dP = ZKT K K1 dQK. We apply this formula for the right sideof (A.15). With the notation

    K =

    sup

    tK TxKt t + sup

    tK TyKt t

    2

    we have

    PK =

    K

    ZKT K K1 dQK(A.18)

    The random process ZKtTK K is the martingale with respect to P andsince P QK it is a semimartingale with respect to QK [see, e.g., Liptser andShiryayev (1989), Chapter 4, Section 5]. For further analysis QK semimartin-gale description of ZKtTK K is required. To this end,we use the fact thatP-semimartingales xKt y

    KT are Q

    K semimartingales as well [see Liptser and

    Shiryayev (1989), Chapter 4, Section 5] and find their semimartingale decom-

    positions. Let us note that t t

    t

    t are counting processes with respect to

    both measures P and QK. Recall that At

    At

    At At are their compensatorswith respect to P and denote by AQt AQt AQt AQt their compensators withrespect to QK. Then the P-martingale mKt = 1Kt At t At obeysthe semimartingale decomposition (P-and QK-a.s.)

    mKt =1

    K

    t AQt

    t AQt 1

    K

    At AQt

    At AQt = mKQt

    1

    K

    At AQt

    At AQt (A.19)

    with QK- square integrable martingales mK Qt with the predicatable quadratic

    variation mK Qt =1

    KAQ

    t + AQ

    t . Analogously, the QK-semimartingaledecomposition mKt = mK Qt 1KAt AQt At AQt holds with thesquare integrable martingale m

    K Qt with mK Qt = 1KAQt + AQt and mK Q,

    mK Qt 0 (recall that disjointness for jumps of counting processes remainsvalid with respect to QK).

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1285

    We give now formulas for AQt A

    Qt

    A

    Qt

    A

    Qt . With

    t = t t1 ,

    t = t t we have

    ZKt K K= XKtK K expKtt T+Ktt t

    (A.20)

    AQt : It is well known that AQt is defined by integral equation: for anybounded and predictable function ut ,

    T0

    ut dt dQK =

    T0

    ut dAQt dQK

    Taking into account that dQK = ZKT K KP and ZKtTK K is the mar-tingale with respect to P we get

    T0

    ut dt dQK =

    T0

    ut ZKT K K dt dP

    =

    T0

    ut ZKt K K dt dP

    Since jumps of counting processes are disjoint the right side of the above equal-

    ity is equal to [see (A.20)]

    T0

    ut ZKtK KeKtt dtdP or, what is

    the same,

    T0

    ut ZKtK KeKt dt dP. Now, since Z

    KtK K is

    the predictable process, the latter integral coincides with

    T0 ut ZKtK KeKt dAt dPwhich, due to the martingale property of ZKt K K, is equal to

    T0

    ut ZKT K KeKt dAt dP =

    T0

    ut eKt dAt dQK

    Consequently,

    dAQt = e

    Kt dAt

    and dAQt = eKt dAt d

    AQt = eKt d

    At d A

    Qt = eKtd

    At are

    obtained in the same way.Hence

    dmKt = dmKQt +

    eKt 1xKt eKt 1xKt yKt dt

    dmKt = dmKQt +

    eKt 1xKt yKt eKt 1yKt dt

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    1286 F. C. KLEBANER AND R. LIPTSER

    and thereby we find the semimartingale decomposition with respect to QK:

    dxKt= xKt yKt +eKt 1xKt eKt 1xKt yKt dt + dmKQt=

    G

    Kt xKt yKt

    dt + dmKQt = t dt + dmKQt

    dyKt =

    yKtxKt

    +eKt 1xKt yKt eKt 1yKt dt + dmKQt

    =

    G

    Kt xKt yKt

    dt + dmKQt = t dt + dmKQt

    (A.21)

    The use of this decomposition allows to obtain a description for 1K

    log ZKT K,K with respect to QK:

    1K

    logZKT K K

    =K T

    0Kt dmKQt +

    K T0

    Ktt GKt xKt yKt

    dt

    +K T

    0Kt dmKQt +

    K T0

    Ktt GKt xKt yKt

    dt

    Let us estimate above the right side of this equality on the set K . There are

    positive constants c and c so that

    suptK T

    Kt t + suptK T

    GKt xKt yKt Gt t t c

    suptK T

    K

    t

    t +

    suptK T

    G

    K

    t

    xK

    t yK

    t G

    t

    t

    t c

    Further, since

    tt Gt t t = sup

    t G t t

    tt Gt t t = sup

    t G t t

    and

    JT =K T

    0sup

    t G t t dt

    +

    K T

    0sup

    t G t t dt

    we find

    1

    KlogZKT K K

    K T0

    Kt dmKQt+ K T

    0Kt dmKQt

    +c + cT + JT

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1287

    This upper bound implies the lower one,

    ZK

    T KK

    exp

    K

    J

    T

    + c

    +c

    T

    exp

    K

    K T0

    Kt dmKQtKK T

    0Kt dmKQt

    which, being applied in (A.18), gives

    PK expKJ + c + cT

    K

    exp

    K K

    0Kt dmKQt

    K

    K T0

    Kt mKQt

    dQK

    (A.22)

    (A.22) can be sharpened as

    PK eKJT+c+cT+2QK

    K K K

    where

    K =

    K T0

    Kt dmKQt < and K = K T

    0Kt dmKQt

    < Hence

    liminfK

    1

    Klog PK JT c + cT 2

    + lim infK

    1

    Klog QK

    K K K

    The latter inequality jointly with (A.15) imply the statement of the lemma

    provided that for fixed and it holds lim infK QKK K K = 1.We derive that property from

    limK

    QK

    \K = 0 lim

    KQK

    \K

    = 0 limK

    QK

    \K = 0(A.23)

    The first part of (A.23) follows from the Chebyshev inequality (EK is theexpectation with respect to QK),

    QK

    \K = QK K T

    0Kt dmKQt

    1

    2EK

    K T

    0 K

    t

    2 d

    mKQ

    t

    1K22

    EKK T

    0Kt2eKt dAt + e

    Kt dAt

    constK2

    0 K

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    1288 F. C. KLEBANER AND R. LIPTSER

    The validity of the second part of (A.23) is verified similarly. The third part

    in (A.23) is verified similarly to the first and second ones since by virtue of

    (A.21),

    \K =

    suptK T

    mKQt + suptK T

    mKQt 2

    A.5.2. The lower bound under T T = .

    Lemma 4. Assume

    (i) .(ii) T T = .Then

    lim inf0 lim infK1

    K logPxK yK JT Proof. Set nt = x0 +

    r0

    sIs n ds. SinceT

    0t dt < , it holds

    limn

    suptT

    t nt T

    0It > nt dt = 0

    Similarly, for nt = y0+r

    0 sIs n ds we have limn suptT tnt = 0. Letus choose n0 so that for n n0 it holds suptT t nt + suptT t nt 2 .Since by the triangular inequality,

    supt

    T

    xKt t+ sup

    t

    T

    yKt t+ sup

    t

    T

    xKt nt+ sup

    t

    T

    yKt nt+

    2

    for any n n0 we get

    PxK yK P

    xK yK n n 2

    Therefore, by Lemma 3,

    lim infK

    1

    KlogP

    xK yK n n

    2

    JTn n Ln n

    Since inftT t > 0 and inftT t > 0 and t t and approximated by nt

    nt

    uniformly in t T, for n large enough a majorant L for Ln n can bechosen, so that

    lim infK

    1K

    logPxK yK 2 JTn n L

    We show that

    lim supn

    JTn n JT (A.24)

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    ANALYSIS AND EXTINCTION IN LOTKAVOLTERRA MODEL 1289

    Since for n large enough inftT nt > 0, inftT

    nt > 0, introduce

    nt= log nt2nt + n2t4nt 2 + nt t= log

    nt

    2nt nt

    +

    n2t4nt nt 2

    + nt

    (A.25)

    and note that

    JTn n =T

    0It n

    ntt G

    nt nt nt

    dt

    +T

    0It > n

    G

    nt nt nt

    dt

    + T0

    It nntt Gnt nt nt dt+

    T0

    It > nGnt n ndt

    JT +T

    0

    Gt t t Gnt nt nt dt+

    T0

    Gt t t Gnt nt nt dtThe second and third terms in the right side of the above inequality converge

    to zero, as n , by virtue of the uniform (in t T) convergence n n

    and T0 nt nt tt dt 0 and T0 nt nt tt dt 0;that is, (A.24) holds.

    Thus

    lim infK

    1

    KlogP

    xK yK

    2

    JT L (A.26)

    and the statement of the lemma holds.

    A.5.3. The lower bound under T T < . For and h smallenough we have

    lim inf

    0

    lim infK

    1

    KlogPxK yK

    = lim inf0

    liminfK

    P

    suptTT

    xKt t+ suptTT

    yKt t lim inf

    0liminf

    KP

    sup

    tThT

    xKt t+ suptThT

    yKt t

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    1290 F. C. KLEBANER AND R. LIPTSER

    On the other side, for ht t, t T h, ht t, t T h, and fort > T h, t > T h,

    ht = ht ht ht = ht ht respectively, we have

    liminf0

    lim infK

    P

    sup

    tThT

    xKt t + suptThT

    yKt t = lim inf

    0liminf

    KPxK yK h h JTh h

    where the latter inequality follows from Lemma 4.

    Finally,

    JYh h =

    ThT

    0sup

    t G t t dt

    + ThT0

    sup

    t G t t dt

    TT

    0sup

    t G t t dt

    +TT

    0sup

    t G t t dt

    = JT Now combining the obtained results above we arrive at the required lower

    bound,

    lim inf0 lim infK 1K logPxK yK JT

    REFERENCES

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    Department of Mathematics

    and Statistics

    University of Melbourne

    Parkville, Victoria 3052

    AustraliaE-mail: [email protected]

    Department of Electrical

    Engineering Systems

    Tel Aviv Univeristy

    69978 Tel Aviv

    IsraelE-mail: [email protected]


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