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J. Differential Equations 258 (2015) 1058–1105
www.elsevier.com/locate/jde
Global attractivity, spreading speeds and traveling

waves of delayed nonlocal reaction–diffusion systems
Shi-Liang Wu a,∗,1 , Cheng-Hsiung Hsu b,2 , Yanyu Xiao c
a School of Mathematics and Statistics, Xidian University, Xi’an, Shaanxi 710071, People’s Republic of China
b Department of Mathematics, National Central University, Chungli 32001, Taiwan
c Department of Mathematics, University of Miami, P.O. Box 249085, Coral Gables, FL 33124-4250, USA
Received 22 December 2013; revised 17 September 2014

Available online 22 November 2014
Abstract
The purpose of this work is to study the spatial dynamics of some delayed nonlocal reaction–diffusion
systems in whole space. We first establish a series of comparison theorems to investigate the global attractiv-
ity of the equilibria for a delayed nonlocal reaction–diffusion system with and without quasi-monotonicity.
Then we show that the spreading speed of a general system without quasi-monotone conditions is coin-
cident with the minimal wave speed. Applying a fluctuation method, we further provide some sufficient
conditions to ensure the upward convergence of the spreading speed and traveling wave solutions. Finally,
we point out the effects of the delay and nonlocality on the spreading speed of the non-quasi-monotone
systems.
© 2014 Elsevier Inc. All rights reserved.
MSC: 35K57; 35R10; 35B40
Keywords: Global attractivity; Spreading speed; Traveling wave solution; Minimal wave speed; Time delay and
nonlocality
* Corresponding author.
E-mail addresses: slwu@xidian.edu.cn (S.-L. Wu), chhsu@math.ncu.edu.tw (C.-H. Hsu), yanyuxiao@gmail.com
(Y. Xiao).
1 Research supported in part by the NSF of China (11301407), NSF of Shaanxi Province (2013JQ1012) and
Fundamental Research Funds for the Central Universities (K5051370002).
2 Research supported in part by MST and NCTS of Taiwan.
http://dx.doi.org/10.1016/j.jde.2014.10.009
0022-0396/© 2014 Elsevier Inc. All rights reserved.
S.-L. Wu et al. / J. Differential Equations 258 (2015) 1058–1105 1059
1. Introduction
To model the spatial spread of epidemics via the environmental pollution produced by the in-
fective human population, Capasso and Maddalena [5] proposed the following reaction–diffusion
system:

ut (x, t) = d1 uxx − a11 u(x, t) + a12 v(x, t),
(1.1)
vt (x, t) = d2 vxx − a22 v(x, t) + g u(x, t) ,
where u(x, t) and v(x, t), respectively, represent the spatial densities of bacteria and infective
population at a point x in the habitat Ω ⊆ R and time t ≥ 0; d1 > 0 and d2 ≥ 0 are diffusion
coefficients; a11 is the natural death rate of bacteria and a12 v is the contribution of infectious
population to the density of bacteria; a22 is the natural diminishing rate of infected individuals.
The nonlinearity g(u) gives the “force of infection” on human due to the concentration of bac-
teria. In the past decades, dynamics of the model (1.1) with specific reaction terms have been
widely discussed and analyzed, see [3,4,6,11,12,37,43] and references cited therein.
Later, in view of the latent period of a virus, Thieme and Zhao [25] modified the system (1.1)
by studying the following reaction–diffusion model with distributed time delay:
⎧
⎪ ut (x, t) = d1 uxx − a11 u(x, t) + a12 v(x, t),
⎪
⎪
⎨
∞
(1.2)
⎪ vt (x, t) = d2 vxx − a22 v(x, t) + g u(x, t − s) P (ds),
⎪
⎪
⎩
0
where P is a probability measure on R+ which describes the distribution of the latent period. In
particular, if the latent period has a fixed length τ , system (1.2) takes the form:

ut (x, t) = d1 uxx − a11 u(x, t) + a12 v(x, t),
(1.3)
vt (x, t) = d2 vxx − a22 v(x, t) + g u(x, t − τ ) .
However, for some infection agents, such as indirect transmission diseases (typhoid fever, schis-
tosomiasis, malaria, etc.), u depends on not only the infective humans v at the spatial point x but
also at the spatial neighbor points of x (even points in the whole region). Following this aspect,
Capasso [2] further proposed and studied the following nonlocal model (see also [3,38]):
⎧
⎪
⎨ ut (x, t) = d1 uxx − a11 u(x, t) + J (x − y)v(y, t)dy,
(1.4)
⎪
⎩
Ω

vt (x, t) = d2 vxx − a22 v(x, t) + g u(x, t) ,
where the kernel function J (x) describes the transfer kernel of the infective agents produced by
the infective humans.
Recently, many researchers pointed out that the interaction effect of spatial movement, non-
local interaction and time delay such as latent period on the spread of the disease should be
taken into account for more realistic models. We refer to the survey papers Gourley et al. [10]
1060 S.-L. Wu et al. / J. Differential Equations 258 (2015) 1058–1105
and Ruan [22]. Therefore, in this work, we will examine the spatial dynamics of the following
delayed nonlocal reaction–diffusion system:

ut = d1 uxx − αu(x, t) + h (J1 ∗ v)(x, t) ,
(1.5)
vt = d2 vxx − βv(x, t) + g (J2 ∗ u)(x, t) ,
where d1 > 0, d2 ≥ 0; the convolution terms in (1.5) are defined by

(J1 ∗ v)(x, t) = J1 (x − y)v(y, t − τ1 )dy,
Ω

(J2 ∗ u)(x, t) = J2 (x − y)u(y, t − τ2 )dy, (1.6)
Ω
here τ1 , τ2 ≥ 0 are given constants and Ω = R. Noting that for the derivation of some specific
forms of the kernel functions Ji (·), i = 1, 2, we refer to the survey paper [10].
In recent years, in order to study the interaction effects of spatial diffusion and time delay
on the evolutionary behavior of biological and epidemic systems, some incisive results for the
dynamics of delayed nonlocal reaction–diffusion models have subsequently been done. Here we
only refer to the survey papers Gourley et al. [10] and Ruan [22]. Note that the main research
topics include coexistence and global stability of steady states, existence of spreading speed
and traveling wave solutions, etc. There have been many significant results on the stability of
equilibria for reaction–diffusion equations defined on bounded domains, see [4,6,32,39,41] and
the references therein. However, it seems that only little result has been established for those
systems defined on whole space. For examples, Kyrychko et al. [14] derived a delayed stage-
structured population model on an isolated lattice and studied the stability of the unique positive
equilibrium. Recently, Wang and Li [27] extended the method in [14] to a delayed nonlocal
reaction–diffusion equation in Rn . For other related results, we refer to [40,42].
Inspired by the works of Kyrychko et al. [14] and Wang and Li [27], the first subject of this
paper is to study the stability of equilibria of the delayed nonlocal system (1.5) in R2 with and
without quasi-monotone conditions. Throughout this article, we assume that the kernel functions
Ji (·) and nonlinearities g(·), h(·) satisfy the following assumptions:

+∞
(A1 ) Ji ∈ L1 (R), Ji (−x) = Ji (x) ≥ 0 for x ∈ R, −∞ Ji (y)dy = 1 and there exists a λ0 > 0
(λ0 may be +∞) such that
+∞ +∞
e−λy Ji (y)dy < +∞ for λ ∈ [0, λ0 ) and lim e−λy Ji (y)dy = +∞.
λ→λ0 −0
−∞ −∞
(A2 ) g, h ∈ C 2 (R+ , R), h(0) = g(0) = 0,
g(u), h(u) > 0 for u > 0 and h (v) > 0 for v ∈ [0, +∞).
By the above assumptions and some restrictions, we first prove the positivity and boundedness
for the solutions and establish a series of comparison theorems by applying the theory of abstract
functional differential equations developed in the work [21]. Next, we show that if there has no
positive equilibrium, the zero equilibrium is globally attractive. We also give some sufficient con-
ditions to guarantee the global attractivity of the unique positive equilibrium. If these conditions
do not hold, we obtain a bounded and positive attractor of positive solutions. For example, if we
take h(v) = v and g(u) = pue−δu , where p, δ > 0 are given parameters, then our results show
that
(i) if 0 < p ≤ αβ, then the zero equilibrium (0, 0) is global attractive;
(ii) if αβ < p ≤ αβe2 , then the unique positive equilibrium (K1 , K2 ) = (δ −1 ln αβ p
, αK1 ) is
global attractive; and
(iii) if p > αβe2 , then C[(k − ,k − ),(k + ,k + )] is a global attractor. See Remark 3.7 for details and the
1 2 1 2
definitions of C[(k − ,k − ),(k + ,k + )] , (k1− , k2− ) and (k1+ , k2+ ).
1 2 1 2
To the best of our knowledge, we give the first work to study the stability of the equilibria for
delayed nonlocal reaction–diffusion systems in whole space, especially for non-quasi-monotone
systems.
Next, we consider the spreading speeds and traveling wave solutions of (1.5). Specifically,
when g (u) ≥ 0 and h (v) ≥ 0 for (u, v) ∈ [0, K] where K is the unique positive equilibrium of
(1.5), (1.5) is a quasi-monotone system and the comparison principle holds. The spreading speed
and minimal wave speed for such monotone systems can be easily established by applying the
powerful theory developed by Liang and Zhao [18] for monotone semiflow. On the other hand,
if the above monotone conditions on h and g do not hold, then (1.5) is a non-quasi-monotone
system. In this case, the solution semiflow for such systems may not be monotone and Liang and
Zhao’s theory for monotone semiflow can not be applied directly to obtain the existence results
of traveling wave solutions and spreading speeds. Moreover, for the case d1, d2 > 0, it is also
difficult to transform system (1.5) into integral equations as those discussed in the work [25].
Due to these reasons, it is worthwhile to study the traveling wave solutions and spreading speeds
for such non-quasi-monotone delayed nonlocal reaction–diffusion systems. This constitutes the
second subject of this paper.
More precisely, we shall consider the spreading speeds and traveling wave solutions for the
following general delayed nonlocal reaction–diffusion system without quasi-monotone assump-
tion:

ut = d1 uxx + f1 u(x, t), (J1 ∗ v)(x, t) ,
(1.7)
vt = d2 vxx + f2 (J2 ∗ u)(x, t), v(x, t) ,
where d1 > 0, d2 , τ1 , τ2 ≥ 0; the convolutions are given by (1.6), and the nonlinearity F :=
(f1 , f2 ) ∈ C 2 (R2+ , R2 ) satisfies the following conditions:
(B1 ) there exists a K := (K1 , K2 )

0 := (0, 0) such that F (0) = F (K) = 0, and there is no
other positive zero of F between 0 and K;
(B2 ) ∂1 f1 (0, 0)∂2 f2 (0, 0) < ∂2 f1 (0, 0)∂1 f2 (0, 0) and ∂i fi (0, 0) < 0, i = 1, 2.
To overcome the difficulty arising from the non-qausi-monotone nonlinearity, we introduce two
auxiliary quasi-monotone systems to “trap” system (1.7). Note that the method has been used by
many researchers for various scalar non-monotone delayed equations [7,8,13,15,20,35] and for
non-cooperative systems without delay [26]. Here, we extend the method to non-quasi-monotone
systems with delay and nonlocal interaction. Based on a comparison argument for solutions of
Cauchy problems of the auxiliary and original systems, we establish the existence of spreading
speed c∗ and obtain the non-existence result of traveling wave solutions when the wave speed c
is less than c∗ . Moreover, applying Schauder’s fixed-point theorem and a limiting argument, we
establish the existence of traveling wave solutions when the wave speed c ≥ c∗ . The asymptotic
behavior of the traveling wave solutions with speed c > c∗ at −∞ and the downward convergence
of the minimal wave are also obtained.
We would like to mention that it is not easy to obtain the upward convergence of the spreading
speed and traveling wave solutions due to the non-quasi-monotone nonlinearity. In fact, the trav-
eling wave solutions for such non-quasi-monotone delayed systems may not be monotone (see
e.g. [9]). In this article, we establish some sufficient conditions to ensure the upward convergence
of the spreading speed by using a fluctuation method [7,8,13,24]. This method was developed by
Thieme and Zhao [24] for a non-local delayed and diffusive predator-prey model. Authors of [7,
8,13] also used the method to prove the upward convergence of the spreading speed for various
non-monotone evolution equations. The upward convergence of the traveling wave solutions is
then obtained. In addition, we point out the effects of the delay and nonlocality on the spread-
ing speeds of the non-quasi-monotone system. According to our analyzes, one can see that the
delay slows the spreading speed and the nonlocality increases the spreading speed. Similar phe-
nomenons have also been founded by Li et al. [17], Zou [44] and Wang et al. [28,29] for scalar
delayed nonlocal reaction–diffusion equations.
The rest of the paper is organized as follows. In Section 2, we obtain the positivity and bound-
edness of solutions for (1.5) and establish a series of comparison theorems. Section 3 is devoted
to the global attractivity of the equilibria of (1.5). A specific example is also given to illustrate
the main results (see Remark 3.7). In Section 4, we first recall some known results on spreading
speeds and traveling wave solutions for system (1.7) satisfying the quasi-monotone conditions.
Then, using squeeze method combining with two auxiliary systems, we obtain the minimal wave
speed and spreading speed of system (1.7) without satisfying the quasi-monotone conditions. In
particular, we give some sufficient conditions to ensure the upward convergence of the spread-
ing speed and traveling wave solutions. The downward convergence of the minimal wave is also
obtained. Finally, we investigate the effects of delay and nonlocality on the spreading speed and
apply our results to a specific delayed nonlocal epidemic model.
2. Solutions of Cauchy problem and comparison theorems
In this section, we first prove the positivity and boundedness of solutions for the Cauchy
problem of (1.5) when initial values are positive and bounded. Then we establish a series of
comparison theorems which can help us to investigate the global attractivity of the equilibria in
the next section. To this end, two classes of the nonlinearity g are considered as following:
(A3 ) g(·) is increasing on [0, +∞) and limu→+∞ g(u) := g∞ > 0;

(A3 ) there exists a number u∗ > 0 such that g(u) is increasing for 0 < u ≤ u∗ and decreasing
for u ≥ u∗ .
For convenience, some notations and definitions are introduced in the sequel.
Notation 2.1.
(1) Let X := BUC(R, R2 ) be the Banach space of all bounded and uniformly continuous func-
tions from R into R2 with the supremum norm · X . The space BUC(R, R) is defined
similarly. Moreover, we denote

X + := ϕ = (ϕ1 , ϕ2 ) ∈ X : ϕi (x) ≥ 0, x ∈ R, i = 1, 2 .
(2) Let τ := max{τ1 , τ2 } and C := C([−τ, 0], X) be the Banach space of continuous func-
tions from [−τ, 0] into X with the supremum norm. Moreover, we denote C + := {φ ∈ C :
φ(s) ∈ X + , s ∈ [−τ, 0]}.
(3) Given L
0, let’s denote the spaces

X[0,L] := ϕ ∈ X : ϕ(x) ∈ [0, L], x ∈ R , (2.1)

C[0,L] := φ ∈ C : φ(x, s) ∈ [0, L], x ∈ R, s ∈ [−τ, 0] . (2.2)
(4) As usual, we identify an element φ ∈ C as a function from R × [−τ, 0] into R2 defined by

φ(x, s) = φ(s)(x). For any continuous function w : [−τ, b) → X, b > 0, we define w t ∈ C,
t ∈ [0, b) by w t (s) = w(t + s), s ∈ [−τ, 0]. Then t → w t is a continuous function from [0, b)
to C.
According to Notation 2.1, it is easy to see that X + and C + are closed cones of X and C,
respectively.
Notation 2.2.
(1) Let T (t) = diag(T1 (t), T2 (t)) be a family of linear operators defined for t ≥ 0 by Ti (0) = I
and

Ti (t)[ϕ](x) := Γi (x − y, t)ϕ(y)dy, for ϕ ∈ BUC(R, R), x ∈ R, t > 0,
R
where

x2
Γ1 (x, t) := (4d1 πt)−1/2 exp − − αt , (2.3)
4d1 t
2
(4d2 πt)−1/2 exp{− 4dx 2 t − βt}, if d2 > 0,
Γ2 (x, t) := (2.4)
e−βt δ(x), if d2 = 0.
Here δ(·) is the Dirac function. It is clear that T (t) is a linear semigroup on X and
T (t)X + ⊂ X + .
(2) Let’s define F = (F1 , F2 ) : C → X by

F1 (φ1 , φ2 )(x) := h J1 (x − y)φ2 (y, −τ1 )dy ,
R

F2 (φ1 , φ2 )(x) := g J2 (x − y)φ1 (y, −τ2 )dy .
R
2.1. Positivity and boundedness of solutions
Following the above notations and definitions, we have the following results.
Theorem 2.3. Assume that (A1 ), (A2 ) and (A3 ) or (A3 ) hold. Then, for any ϕ ∈ C + , there exists
a unique solution w(x, t; ϕ) = (u(x, t; ϕ), v(x, t; ϕ)) of (1.5) on [0, +∞) such that

e−αt δ1 , e−βt δ2 ≤ w(x, t; ϕ) ≤ K̄ := (K̄1 , K̄2 ) for any (x, t) ∈ R × [0, +∞), (2.5)
where δ1 := infx∈R ϕ1 (x, 0), δ2 := infx∈R ϕ2 (x, 0), Mi := maxs∈[−τ,,0] supx∈R ϕi (x, s), i = 1, 2
and

(max{α −1 h(K̄2 ), M1 }, max{β −1 g∞ , M2 }), if (A3 ) hold;
K̄ :=
(max{α h(K̄2 ), M1 }, max{β g(u∗ ), M2 }), if (A3 ) hold.
−1 −1
Furthermore, w(x, t; ϕ) is classical on (τ, +∞) and if ϕ(0) ∈ Int(X + ), then w(t) ∈ Int(X + ) for
t ≥ 0 and w t ∈ Int(C + ) for t > τ .
Proof. According to the abstract setting of [21], one can see that a mild solution of (1.5) is a
continuous solution to its associated integral equation
⎧
⎪
⎪ t
⎪
⎨
w(t) = T (t)ϕ(0) + T (t − s)F w s ds, t > 0,
⎪
⎪ 0
⎪
⎩
w(s) = ϕ(s), s ∈ [−τ, 0].
First, we assume g(·) satisfies the condition (A3 ). In this case, it is easy to verify that
w− (x, t) := (0, 0) and w+ (x, t) := K̄ constitute a pair of sub-solution and super-solution of
(1.5). Moreover, one can easily see that F : C[0,K̄] → X is globally Lipschitz continuous and
quasi-monotonic on C[0,K̄] in the sense that
1
lim dist ψ(0) − φ(0) + η F (ψ) − F (φ) , X+ = 0
η→0+ η
for all φ, ψ ∈ C[0,K̄] with ψ ≥ φ. Therefore, the existence and uniqueness of w(x, t; ϕ) follows
from [21, Corollary 5] with S(t, s) = T (t, s) = T (t − s) for t ≥ s ≥ 0 and B(t, φ) = F (ϕ).
Moreover, it follows from [31, Corollary 2.2.5] that w(x, t; ϕ) is classical for t > τ .
Next, we consider the case that (A3 ) holds. For any φ = (φ1 , φ2 ) ∈ C + , let’s define

B + (φ) := B1+ (φ), B2+ (φ) := h(φ2 ), g(u∗ ) ,

B(φ) := B1 (φ), B2 (φ) := h(φ2 ), g(φ1 ) ,

B − (φ) := B1− (φ), B2− (φ) := h(φ2 ), 0 .
Then B(φ) is Lipschitz continuous on C[0,K̄] and it is easy to verify that w− (x, t) := 0 and
w+ (x, t) := K̄ satisfy
t
r
w+ (t) ≥ T (t − s)w+ (s) + T (t − r)B + w+ dr,
s
t
r
w− (t) = T (t − s)w− (s) + T (t − r)B − w− dr
s
for any 0 ≤ s < t < +∞. Moreover, for any φ ∈ C[0,K̄] and η > 0, we have
t
w+ (t) − φ(0) + η B + w+ − B(φ) ≥ K̄ − φ(0) ≥ 0 in X,
which implies that
1 t
lim dist w+ (t) − φ(0) + η B + w+ − B(φ) , X + = 0, for any t ≥ 0.
η→0+ η
Similarly, we can show that
1 t +
lim dist φ(0) − w− (t) + η B(φ) − B − w− , X = 0, for any t ≥ 0.
η→0+ η
Take S ± (t, s) = S(t, s) = T (t, s) = T (t − s). Then [21, Proposition 3] implies that there exists
a unique solution w(x, t; ϕ) of (1.5) on [0, +∞) such that 0 ≤ w(x, t; ϕ) ≤ K̄ for any (x, t) ∈
R × [0, +∞).
Finally, since g(u), h(u) > 0 for u > 0, we have

w(x, t; ϕ) ≥ T (t)ϕ(0)(x) ≥ e−αt inf ϕ1 (x, 0), e−βt inf ϕ2 (x, 0) = e−αt δ1 , e−βt δ2 ,
x∈R x∈R
for (x, t) ∈ R × [0, +∞). It follows that if ϕ(0) ∈ Int(X + ), then w(t) ∈ Int(X + ) for t ≥ 0 and
w t ∈ Int(C + ) for t > τ . This completes the proof. 2
2.2. Comparison theorems
Now, we establish two comparison theorems which will be used to prove the global attractivity
of the equilibria.
Lemma 2.4. Assume (A1 )–(A3 ). Given any a ∈ R. Let w+ (x, t) and w− (x, t) be a pair of
bounded functions defined on [a, ∞) which satisfy
t
r
w+ (t) ≥ T (t − s)w+ (s) + T (t − r)F w+ dr, for any a ≤ s < t < ∞,
s
t
r
w− (t) ≤ T (t − s)w− (s) + T (t − r)F w− dr, for any a ≤ s < t < ∞,
s
respectively. If w+ (x, s) ≥ w− (x, s) ≥ 0 for x ∈ R and s ∈ [a − τ, a], then w+ (x, t) ≥

w− (x, t) ≥ 0 for (x, t) ∈ R × [a, ∞).
Proof. Since g(·) and h(·) are non-decreasing on [0, ∞), the assertion of this lemma follows
from [21, Corollary 5]. We omit its proof. 2
For non-monotone function g, we first introduce the definitions of super-solution and sub-
solution of (1.5).
Definition 2.5. Two functions w + = (u+ , v + ) and w − = (u− , v − ) are called a pair of super-
solution and sub-solution of (1.5) if
(i) there exists M̄ := (M̄1 , M̄2 ) > 0 such that 0 ≤ w − (x, t) ≤ w + (x, t) ≤ M̄ for (x, t) ∈ R ×
[−τ, +∞);
(ii) for any function φ = (φ1 , φ2 ) with w − (x, t) ≤ φ(x, t) ≤ w + (x, t) for (x, t) ∈ R ×
[−τ, +∞), there holds

u+ + +
t ≥ d1 uxx − αu (x, t) + h (J1 ∗ φ2 )(x, t) ,
(2.6)
vt+ ≥ d2 vxx
+
− βv + (x, t) + g (J2 ∗ φ1 )(x, t) ,
and

u− − −
t ≤ d1 uxx − αu (x, t) + h (J1 ∗ φ2 )(x, t) ,
(2.7)
vt− ≤ d2 vxx
−
− βv − (x, t) + g (J2 ∗ φ1 )(x, t) ,
for (x, t) ∈ R × (0, +∞).
Then we establish the following comparison theorem.
Lemma 2.6. Assume (A1 ), (A2 ) and (A3 ) . Let w + = (u+ , v + ) and w − = (u− , v − ) be a pair of
super-solution and sub-solution of (1.5). If
w − (x, s) ≤ ϕ(x, s) ≤ w + (x, s) for (x, s) ∈ R × [−τ, 0],
then the unique solution w(x, t; ϕ) = (u(x, t; ϕ), v(x, t; ϕ)) of (1.5) with initial value ϕ ∈ C +
satisfies
w − (x, t) ≤ w(x, t; ϕ) ≤ w + (x, t), for all x ∈ R and t > 0.
Proof. By Definition 2.5 and Theorem 2.3, we see that 0 ≤ w(x, t; ϕ), w + (x, t) ≤ K̄ for any
(x, t) ∈ R × [0, +∞), where K̄ := (K̄1 , K̄2 ) =(max{α −1 h(K̄2 ), M̄1 }, max{β −1 g(u∗ ), M̄2 }). Set

Z(x, t) = Z1 (x, t), Z2 (x, t) := w(x, t; ϕ) − w + (x, t), for (x, t) ∈ R × [−τ, +∞).
For any φ = (φ1 , φ2 ) satisfies the condition of (ii) in Definition 2.5, it is easily seen that

(Z1 )t ≤ d1 (Z1 )xx − αZ1 + h (J1 ∗ v)(x, t) − h (J1 ∗ φ2 )(x, t) ,
(2.8)
(Z2 )t ≤ d2 (Z2 )xx − βZ2 + g (J2 ∗ u)(x, t) − g (J2 ∗ φ1 )(x, t) ,
for (x, t) ∈ R × (0, +∞). Four cases are considered in the sequel.
Case (i). τ1 , τ2 > 0.
Note that τm := min{τ1 , τ2 } > 0 and τ := max{τ1 , τ2 }. Let’s choose φ(x, s) = ϕ(x, s) for
(x, s) ∈ R × [−τ, 0]. Then it follows from (2.8) that

(Z1 )t ≤ d1 (Z1 )xx − αZ1 ,
for (x, t) ∈ R × (0, τm ], (2.9)
(Z2 )t ≤ d2 (Z2 )xx − βZ2 ,
which implies that
Zi (x, t) ≤ Ti (t)Zi (x, 0) ≤ 0, for (x, t) ∈ R × (0, τm ], i = 1, 2.
Hence, Z(x, t) ≤ 0 for (x, t) ∈ R × [0, τm ], that is, w(x, t; ϕ) ≤ w + (x, t) for (x, t) ∈ R × [0, τm ].
Similarly, we also obtain that w − (x, t) ≤ w(x, t; ϕ) for (x, t) ∈ R × [0, τm ].
Now we choose φ(x, s) = w(x, s; ϕ) for (x, s) ∈ R × [τm − τ, τm ]. From (2.8), we also have

(Z1 )t ≤ d1 (Z1 )xx − αZ1 ,
for (x, t) ∈ R × (τm , 2τm ],
(Z2 )t ≤ d2 (Z2 )xx − βZ2 ,
which also implies that Z(x, t) ≤ 0 for (x, t) ∈ R × [τm , 2τm ], i.e. w(x, t; ϕ) ≤ w + (x, t) for
(x, t) ∈ R × [τm , 2τm ]. And similarly that w − (x, t) ≤ w(x, t; ϕ) for (x, t) ∈ R × [τm , 2τm ]. Fol-
lowing the same procures, we conclude that w − (x, t) ≤ w(x, t; ϕ) ≤ w + (x, t) for all x ∈ R and
t > 0.
Case (ii). τ1 = 0 and τ2 > 0.
Let’s choose φ1 (x, s) = ϕ1 (x, s) for (x, s) ∈ R × [−τ2 , 0]. By (2.8), we have
(Z2 )t ≤ d2 (Z2 )xx − βZ2 , for (x, t) ∈ R × (0, τ2 ].

Thus, v(x, t; ϕ) ≤ v + (x, t) for (x, t) ∈ R × [0, τ2 ]. Similarly, we have v − (x, t) ≤ v(x, t; ϕ) for
(x, t) ∈ R × [0, τ2 ]. Now we choose φ2 (x, t) = v(x, t; ϕ) for (x, t) ∈ R × [0, τ2 ]. It follows from
the first inequality of (2.8) that
(Z1 )t ≤ d1 (Z1 )xx − αZ1 for (x, t) ∈ R × (0, τ2 ].
Thus, u(x, t; ϕ) ≤ u+ (x, t) for (x, t) ∈ R × [0, τ2 ]. Similarly, we have u− (x, t) ≤ u(x, t; ϕ) for
(x, t) ∈ R ×[0, τ2 ]. Hence, w − (x, t) ≤ w(x, t; ϕ) ≤ w+ (x, t) for (x, t) ∈ R ×[0, τ2 ]. Inductively,
we obtain that w − (x, t) ≤ w(x, t; ϕ) ≤ w+ (x, t) for (x, t) ∈ R × [0, +∞).
Case (iii). τ2 = 0 and τ1 > 0.
The proof is similar to that of the case (ii) and omitted.
Case (iv). τ1 = τ2 = 0.
Let’s choose φ = w + . By the first equation of (2.8), we have
∞

(Z1 )t ≤ d1 (Z1 )xx − αZ1 + h η(x, t) J1 (x − y)Z2 (y, t)dy (2.10)
−∞
for (x, t) ∈ R × (0, +∞), where
∞

η(x, t) := J1 (x − y) v + (y, t) + θ Z2 (y, t) dy ∈ [0, K̄2 ] and θ ∈ (0, 1).
−∞
Denote Lh := maxv∈[0,K̄2 ] h (v) and [B]+ := max{B, 0} for any B ∈ R. Note that Z1 (x, 0) ≤ 0
for all x ∈ R and h (u) > 0 for all u ≥ 0. It follows from (2.10) that
∞
Z1 (x, t) ≤ Γ1 (y, t)Z1 (x − y, 0)dy
−∞
t +∞ ∞

+ Γ1 (y, s)h η(x − y, t − s) J1 (z)Z2 (x − y − z, t − s)dzdyds
0 −∞ −∞
t +∞ ∞

≤ Γ1 (y, s)h η(x − y, t − s) J1 (z) Z2 (x − y − z, t − s) + dzdyds
0 −∞ −∞
t +∞

≤ Lh Γ1 (y, s) J1 ∗ [Z2 ]+ (x − y, t − s)dyds,
0 −∞
where Γ1 is defined in (2.3). Hence, for (x, t) ∈ R × (0, +∞),

t +∞

Z1 (x, t) + ≤ Lh Γ1 (y, s) J1 ∗ [Z2 ]+ (x − y, t − s)dyds.
0 −∞
Moreover, we set

Z̃i,λ (t) := sup Zi (x, t) + e−λt and Z̃i,λ := sup Z̃i,λ (t), for i = 1, 2, and λ > 0.
x∈R t∈[0,∞)
Note that 0 ≤ (Z̃1,λ , Z̃2,λ ) ≤ K̄. Then, we have
t +∞
Z̃1,λ (t) ≤ Lh Γ1 (y, s)Z̃2,λ (t − s)e−λs dyds
0 −∞
t
≤ Lh K̄2 e−(λ+α)s ds ≤ Lh K̄2 /(λ + α).
0
Thus, Z̃1,λ ≤ Lh K̄2 /(λ + α) → 0 as λ → +∞ which implies that u(x, t; ϕ) ≤ u+ (x, t) for
(x, t) ∈ R × [0, +∞). Similarly, we obtain u− (x, t) ≤ u(x, t; ϕ) for (x, t) ∈ R × [0, +∞).
Now, let φ1 (x, t) = u(x, t; ϕ) for (x, t) ∈ R × [0, +∞). It then follows from the second in-
equality of (2.8) that
(Z2 )t ≤ d2 (Z2 )xx − βZ2 , for (x, t) ∈ R × (0, +∞).
Since Z2 (x, 0) ≤ 0 for all x ∈ R, we get v(x, t; ϕ) ≤ v + (x, t) for (x, t) ∈ R × [0, +∞). Sim-
ilarly, we also obtain v − (x, t) ≤ v(x, t; ϕ) for (x, t) ∈ R × [0, +∞). Therefore, w − (x, t) ≤
w(x, t; ϕ) ≤ w + (x, t) for (x, t) ∈ R × [0, +∞). This completes the proof. 2
3. Global attractivity of equilibria
In this section, we will consider the global attractivity of the equilibria of (1.5) when g satisfies
the assumptions (A3 ) and (A3 ) respectively. To guarantee the existence and uniqueness of a
positive equilibrium, we impose the following condition for the nonlinearities g(·) and h(·):
(A4 ) αβ < h (0)g (0) and there exists a constant K1 > 0 such that h(β −1 g(u)) > αu for u ∈
(0, K1 ) and h(β −1 g(u)) < αu for u > K1 .
One can easily verify that (A4 ) implies the existence of a unique positive equilibrium K :=
(K1 , K2 ) with K2 = β −1 g(K1 ), even if αβ = h (0)g (0). However, the strict inequality αβ <
h (0)g (0) is required in investigating the attractivity of K, see the following lemmas. Note also
that if (A3 ) holds, then system (1.5) satisfies the quasi-monotonicity on R2 and if (A3 ) and (A4 )
with K1 ≤ u∗ hold, then system (1.5) also satisfies the quasi-monotonicity on [0, K]; otherwise,
it is a non-quasi-monotone system.
We first establish the following lemma which plays an important role in the proof of the
global attractivity of the unique positive equilibrium. Recall τ = max{τ1 , τ2 }. For convenience,
let Cτ := C([−τ, 0], R2 ) and Cτ+ := {φ ∈ Cτ : φ(s) ≥ 0, s ∈ [−τ, 0]}. Given function z(t) ∈
C([−τ, +∞), R2 ), then zt ∈ Cτ means zt (θ ) = z(t + θ ), for θ ∈ [−τ, 0]. We also denote the
symbol “ ˆ ” as the inclusion R2 → Cτ by z → ẑ, ẑ(θ ) = z, θ ∈ [−τ, 0].
Lemma 3.1. Assume (A2 ). Given any r ∈ R. Let w(t, ϕ) = (u(t, ϕ), v(t, ϕ)) be a solution of the
following system

u (t) = −αu(t) + h v(t − τ1 ) , t > r,
(3.1)
v (t) = −βv(t) + g u(t − τ2 ) , t >r
with initial value
w r = ϕ = (ϕ1 , ϕ2 ) ∈ Cτ+ . (3.2)
Then the following statements hold:
(1) if (A3 ) holds and αu > h(β −1 g(u)) for u > 0, then limt→∞ w(t, ϕ) = 0;
(2) if (A3 ) and (A4 ) hold and ϕ(r) > 0, then limt→∞ w(t, ϕ) − K = 0;
(3) if (A3 ) and (A4 ) hold, K1 < u∗ and ϕ(r) > 0, then limt→∞ w(t, ϕ) − K = 0.
Proof. Since (3.1) is autonomous, it suffices to consider the case where r = 0. If we replace
X = BUC(R, R2 ) with R2 , it follows from Theorem 2.3 that for any ϕ ∈ Cτ+ , (3.1) has a unique
solution w(t, ϕ) = (u(t, ϕ), v(t, ϕ)) on [0, ∞). Moreover, if (A3 ) holds, one can easily see that
(3.1) is a cooperative system on Cτ+ . Let’s denote f (·) = (f1 (·), f2 (·)) : Cτ+ → R2 and b =
(b1 , b2 ) by

f1 (φ) := −αφ1 (0) + h φ2 (−τ1 ) , f2 (φ) := −βφ2 (0) + g φ1 (−τ2 ) ,

b1 := max α −1 h(b2 ), sup ϕ1 (s) , b2 := max β −1 g∞ , sup ϕ2 (s) .
s∈[−τ,0] s∈[−τ,0]
The assertions are proved in the sequel.

(1) We first show that Σ0 := [0̂, b̂] is positively invariant for (3.1). Let φ ∈ Σ0 . It is easy to
see that if φi (0) = 0, then fi (φ) ≥ 0, i = 1, 2. Further, we have

f1 (φ) = −αb1 + h φ2 (−τ1 ) ≤ −αb1 + h(b2 ) ≤ 0, if φ1 (0) = b1 , and

f2 (φ) = −βb2 + g φ1 (−τ2 ) ≤ −βb2 + g∞ ≤ 0, if φ2 (0) = b2 .
Since (3.1) is cooperative on Σ0 , it then follows from [23, Proposition 5.2.3 and Corollary 5.2.4]
that
lim w(t, 0̂) = c and lim w(t, b̂) = d,

t→∞ t→∞
where 0 ≤ c ≤ d ≤ b and f (ĉ) = f (d̂) = 0. By αu > h(β −1 g(u)) for u > 0, we see that c =
d = 0. Moreover, since ϕ ∈ Σ0 , we have
w(t, 0̂) ≤ w(t, ϕ) ≤ w(t, b̂), for t > 0,
(see e.g. [23, Theorem 5.1.1]). Therefore, limt→∞ w(t, ϕ) = 0.

(2) Since ϕ(0) > 0, it is easy to verify that w(t, ϕ)
0 for t > τ . By renewing the initial time,
we can assume that infs∈[−τ,0] ϕ(s)
0. Set a := (a1 , a2 ) and Σ := [â, b̂], where

a1 ∈ 0, min K1 , inf ϕ1 (s), g −1 β inf ϕ2 (s) and a2 := β −1 g(a1 ).
s∈[−τ,0] s∈[−τ,0]
Clearly, a
0 and ϕ ∈ [â, b̂]. By similar arguments in part (1), we can show that Σ is positively
invariant for (3.1). Again, by [23, Proposition 5.2.3 and Corollary 5.2.4], we obtain that
lim w(t, â) = c and lim w(t, b̂) = d,

t→∞ t→∞
where a ≤ c ≤ d ≤ b and f (ĉ) = f (d̂) = 0. Since a

0 and K is the unique positive equilibrium
of (3.1), it must be c = d = K. Noting that
w(t, â) ≤ w(t, ϕ) ≤ w(t, b̂), for t > 0,
we conclude that limt→∞ w(t, ϕ) − K = 0.

(3) Let W1 (t, ϕ) = (U1 (t, ϕ), V1 (t, ϕ)) be the solution of the following problem:
⎧
⎨ U1 (t) = −αU1 (t) + h V1 (t − τ1 ) ,
⎪ t > 0,
V1 (t) = −βV1 (t) + g(u∗ ), t > 0,
⎪
⎩ 0
W1 = ϕ.
According to [23, Theorem 5.1.1], we have w(t, ϕ) ≤ W1 (t, ϕ) for t > 0. Moreover, it is easy to
see that

lim U1 (t, ϕ), V1 (t, ϕ) = α −1 h β −1 g(u∗ ) , β −1 g(u∗ ) .
t→∞
By K1 < u∗ , we have

K1 < α −1 h β −1 g(u∗ ) < u∗ and β −1 g(u∗ ) < h−1 (αu∗ ).
Then, there exists a T1 > τ such that

0 w(t, ϕ) ≤ W1 (t, ϕ) ≤ u∗ , h−1 (αu∗ ) , for t ≥ T1 .
Now we consider the initial value problem of (3.1) starting at time T1 + τ . Set

L := u∗ , h−1 (αu∗ ) and l := (l1 , l2 )
0
where

l1 ∈ 0, min K1 , inf u(s, ϕ), g −1 β inf v(s, ϕ) and l2 := β −1 g(l1 ).
s∈[T1 ,T1 +τ ] s∈[T1 ,T1 +τ ]
Since g(u) and h(v) are increasing on [0, u∗ ] and [0, +∞), respectively, (3.1) is cooperative on
Σ1 := [l̂, L̂]. Similar to the proof of part (2), we can show that Σ1 is positively invariant for (3.1)
and that limt→∞ w(t, ϕ) − K = 0. This completes the proof. 2
Now, we state the main result on the attractivity of the equilibria for the monotone case. For
convenience, for given constant M > 0 and vector δ := (δ1 , δ2 ) ≥ 0, we set the space

+
Cδ,M := ϕ ∈ C + : ϕC ≤ M and inf ϕ(x, 0) ≥ δ .
x∈R
In the remainder of this section, we always denote the solution of (1.5) with the initial value
ϕ ∈ C + by w(x, t; ϕ).
Theorem 3.2. Assume that (A1 )–(A3 ) hold. Then the following results hold:
(1) If αu > h(β −1 g(u)) for u > 0, then for any δ = (δ1 , δ2 ) ≥ 0 and M > 0,

lim w(·, t; ϕ) = 0 +
uniformly for ϕ ∈ Cδ,M .
t→∞
(2) If (A4 ) holds, then for any δ = (δ1 , δ2 ) > 0 and M > 0,

lim w(·, t; ϕ) − K = 0 +
t→∞
Proof. Denote K̄ := (max{α −1 h(K̄2 ), M}, max{β −1 g∞ , M}). From Theorem 2.3, we have
+
δ1 e−αt , δ2 e−βt ≤ w(x, t; ϕ) ≤ K̄ for any (x, t) ∈ R × [0, +∞) and ϕ ∈ Cδ,M .
Let w̄(t) and w(t) be solutions of the following problems:

⎧
⎨ ū (t) = −α ū(t) + hv̄(t − τ1 ),
⎪ t > τ,
v̄ (t) = −β v̄(t) + g ū(t − τ2 ) , t > τ, and
⎪
⎩
w̄(s) = K̄, s ∈ [0, τ ],
⎧
⎪
⎨ u (t) = −αu(t) + h v(t − τ 1 ) , t > τ,

v (t) = −βv(t) + g u(t − τ2 ) , t > τ,
⎪
⎩
w(s) = δ1 e−ατ , δ2 e−βτ , s ∈ [0, τ ],
respectively. Then it follows from Lemma 2.4 that
u(t) ≤ u(x, t; ϕ) ≤ ū(t), for x ∈ R and t ≥ τ. (3.3)
Hence, the assertions of this theorem follow from Lemma 3.1. The proof is complete. 2
Next, we consider the case that (A3 ) holds. The global attractivity of the trivial equilibrium
is stated as follows.
Theorem 3.3. Assume that (A1 ), (A2 ) and (A3 ) hold. If αu > h(β −1 g(u)) for u > 0, then for
any δ = (δ1 , δ2 ) ≥ 0 and M > 0, we have

lim w(·, t; ϕ) = 0 +
t→∞
Proof. We first construct an upper auxiliary system:

u+ (t) = −αu+ (t) + h v + (t − τ1 ) , t > 0,
+ (3.4)
v (t) = −βv + (t) + g + u+ (t − τ2 ) , t > 0,
with initial value

u+ (s), v + (s) = (M, M), s ∈ [−τ, 0],
where

g(u), u ∈ [0, u∗ ],
g + (u) := (3.5)
g(u∗ ), u > u∗ .
Since βh−1 (αu) > g(u) for u > 0 and h(·) is non-decreasing on [0, ∞), we have βh−1 (αu) >
g + (u) ≥ g(u) for u > 0. Moreover, g + (·) is non-decreasing on [0, ∞). It then follows from
part (1) of Lemma 3.1 that limt→∞ (u+ (t), v + (t)) = 0. Moreover, by [21, Proposition 1] (see
also Lemma 4.7), we have

0 ≤ w(x, t; ϕ) ≤ u+ (t), v + (t) for all x ∈ R, t > 0.
Therefore, limt→∞ w(·, t; ϕ) = 0. This completes the proof. 2
To study the global attractivity of K, we first introduce the function:

− minv∈[u,K0 ] g(v), u ∈ [0, K0 ],
g (u) := (3.6)
g(u), u > K0 ,
where K0 := α −1 h(β −1 g(u∗ )) + 1. It is clear that if K1 ≥ u∗ , then there exists k1− ∈ (0, K1 ) such
that β −1 g − (u) > h−1 (αu) for u ∈ (0, k1− ) and β −1 g − (u) < h−1 (αu) for u > k1− . Moreover,
g(u) ≥ g − (u) for all u ≥ 0. For the graph of g − (u), see Fig. 1.
Then, the global attractivity of K for two cases: (i) K1 ≤ u∗ and (ii) K1 > u∗ , are stated in the
sequel.
Fig. 1. Graph of β −1 g − (u) with K1 = u∗ . Note that K̄0 < K1 with g − (K̄0 ) = g(K̄0 ) and g − (u) = g(u) for u ∈
[0, K̄0 ] ∪ [K0 , ∞).
Fig. 2. The graphs of g and h for the case K1 ≤ u∗ .
3.1. (i) K1 ≤ u∗
Theorem 3.4. Assume (A1 )–(A2 ), (A3 ) and (A4 ). If K1 ≤ u∗ (cf. Fig. 2), then for any δ =
(δ1 , δ2 ) > 0 and M > 0, we have

lim w(·, t; ϕ) − K = 0 +
t→∞
Proof. Let K̄ = (K̄1 , K̄2 ) := (max{α −1 h(K̄2 ), M}, max{β −1 g(u∗ ), M}). Then, Theorem 2.3
yields
+
e−αt δ1 , e−βt δ2 ≤ w(x, t; ϕ) ≤ K̄ for any x ∈ R, t ≥ 0 and ϕ ∈ Cδ,M .
It is easy to see that the functions w − (x, t) = 0 and w + (x, t) = (U1 (t), V1 (t)) are a pair of
sub-solution and super-solution of (1.5), where w + (x, t) satisfies
⎧
⎨ U1 (t) = −αU1 (t) + h V1 (t − τ1 ) , t > τ,
⎪
V1 (t) = −βV1 (t) + g(u∗ ), t > τ, (3.7)
⎪
⎩
U1 (s), V1 (s) = K̄, s ∈ [0, τ ].
It then follows from Lemma 2.6 that

lim sup sup w(x, t; ϕ) ≤ lim w + (x, t) = α −1 h β −1 g(u∗ ) , β −1 g(u∗ ) .
t→∞ x∈R t→∞
If K1 < u∗ , we have K1 < α −1 h(β −1 g(u∗ )) < u∗ and β −1 g(u∗ ) < h−1 (αu∗ ). Thus, there
exists a T > 0 such that

w(x, t; ϕ) ≤ w + (x, t) ≤ u∗ , h−1 (αu∗ ) , for x ∈ R and t ≥ T .
Let w̄(t) = (ū(t), v̄(t)) and w(t) = (u(t), v(t)) be solutions of the following problems:
⎧ ⎧
⎪ ū (t) = −α ū(t) + h v̄(t − τ1 ) , ⎪ u (t) = −αu(t) + h v(t − τ1 ) ,
⎨ ⎨
v̄ (t) = −β v̄(t) + g ū(t − τ2 ) , and v (t) = −βv(t) + g u(t − τ2 ) ,
⎪
⎩ ⎪
⎩
w̄(s) = u∗ , h−1 (αu∗ ) , w(s) = δ1 e−α(T +τ ) , δ2 e−β(T +τ ) ,
respectively, where t > T + τ and s ∈ [T , T + τ ]. By the assertion (3) of Lemma 3.1, we have
limt→∞ w(t) = limt→∞ w̄(t) = K. Note that w(t) and w̄(t) are also solutions of (1.5) with
initial values ϕ(·) = w(·) and ϕ(·) = w̄(·), respectively. Since g(u) is increasing for u ∈ [0, u∗ ]
and h(v) is increasing for v ∈ [0, +∞), applying [21, Corollary 5], we obtain
w(t) ≤ w(x, t; ϕ) ≤ w̄(t), for x ∈ R and t > T + τ.

+
Therefore, limt→∞ w(·, t; ϕ) − K = 0 uniformly for ϕ ∈ Cδ,M .
If K1 = u∗ , we first construct another sub-solution of (1.5) by considering the following prob-
lem:
⎧
⎨ u1 (t) = −αu1 (t) + h v1 (t− τ1 ) ,
⎪

t > τ,

v1 (t) = −βv1 (t) + min g u1 (t − τ2 ) , g U1 (t − τ2 ) , t > τ, (3.8)
⎪
⎩ −ατ
−βτ
u1 (s), v1 (s) = e δ1 , e δ2 , s ∈ [0, τ ].
Since g(u) ≤ g(u∗ ) for all u ≥ 0, it is easy to show that

0 ≤ u1 (t), v1 (t) ≤ U1 (t), V1 (t) for t ≥ τ.
Moreover, it is not difficult to verify that (u1 (t), v1 (t)) and (U1 (t), V1 (t)) are a pair of sub- and
super-solutions of (1.5). From Lemma 2.6, we have

u1 (t), v1 (t) ≤ w(x, t; ϕ) ≤ U1 (t), V1 (t) , for x ∈ R and t ≥ τ. (3.9)
Note that when K1 = u∗ ,

lim U1 (t), V1 (t) = α −1 h β −1 g(u∗ ) , β −1 g(u∗ ) = K. (3.10)
t→∞
Hence, to prove this assertion, it is sufficient to show that limt→∞ (u1 (t), v1 (t)) = K. To this end,
we further consider the following problem:
⎧
⎪ (t) = −αu1 (t) + h v 1 (t − τ1 ) ,
⎨ u1
t > τ,
−

v 1 (t) = −βv 1 (t) + g u1 (t − τ2 ) , t > τ,
⎪
⎩
u1 (s), v 1 (s) = e−ατ δ1 , e−βτ δ2 , s ∈ [0, τ ],
where g − (·) is defined in (3.6). Since g(u) ≥ g − (u) for all u ≥ 0, we have (u1 (t), v 1 (t)) ≤
(u1 (t), v1 (t)) for t > τ . From (3.9) and (3.10), there exists T > τ such that for t > T ,

u1 (t), v 1 (t) ≤ u1 (t), v1 (t) ≤ U1 (t), V1 (t) ≤ K0 , β −1 g(u∗ ) + 1 . (3.11)
Noting that g − (u) is decreasing on [0, K0 ] and k1− < K1 = u∗ , it follows from the assertion (3)
of Lemma 3.1 that limt→∞ (u1 (t), v 1 (t)) = (k1− , k2− ), where k2− := β −1 g(k1− ). Moreover,

0 k1− , k2− ≤ lim inf u1 (t), v1 (t) ≤ lim sup u1 (t), v1 (t) ≤ K.
t→∞ t→∞
It is easy to see that limt→∞ u1 (t) exists if and only if limt→∞ v1 (t) exists.
We first prove that limt→∞ (u1 (t), v1 (t)) exists. If this fact is false, then
− −
k1 , k2 ≤ (u− , v− ) := lim inf u1 (t), v1 (t) lim sup u1 (t), v1 (t) ≤ K.
t→∞ t→∞
Let {tj }j ∈N be a sequence with tj → ∞ as j → ∞ such that limj →∞ (u1 (tj ), v1 (tj )) = (u− , v− )
and limj →∞ (u1 (tj ), v1 (tj )) = 0. We can choose a subsequence of {tj }j ∈N , still denote by
{tj }j ∈N , such that limj →∞ u1 (tj − τ2 ) and limj →∞ v1 (tj − τ1 ) exists, say ũ− and ṽ− , respec-
tively. It is clear that (u− , v− ) ≤ (ũ− , ṽ− ) ≤ K, (u− , v− ) K and

αu− = h(ṽ− ) ≥ h(v− ) and βv− = max g(ũ− ), g(K1 ) = g(ũ− ).
Then we consider two cases: (i) ũ− = K1 and (ii) ũ− < K1 . For case (i), we have
h−1 (αK1 ) = β −1 g(K1 ) = v− ≤ h−1 (αu− ).
Hence u− ≥ K1 which leads to a contradiction. For case (ii), noting that β −1 g(u) > h−1 (αu) for
u ∈ (0, K1 ), we have
h−1 (αu− ) ≥ v− = β −1 g(ũ− ) > h−1 (α ũ− ).
Thus, u− > ũ− which also gives a contradiction. Then, the limit (u+ , v+ ) := limt→∞ (u1 (t),
v1 (t)) exists and (3.8) implies that
αu+ = h(v+ ) and βv+ = g(u+ ); or h−1 (αu+ ) = β −1 g(u+ ).

Fig. 3. The graphs of g and h with K1 > u∗ := α −1 h(β −1 g(α −1 h(β −1 g(u∗ )))) > u∗ .
Since 0 (k1− , k2− ) ≤ (u+ , v+ ) ≤ K, it must be (u+ , v+ ) = K. Therefore, limt→∞ (u1 (t),
v1 (t)) = K. The proof is complete. 2
3.2. (ii) K1 > u∗
To obtain global attractivity of K in this case, we further impose the following assumptions.
(Q) ub(u) is strictly increasing in u ∈ (0, α −1 h(β −1 g(u∗ ))], where b(u) := α −1 h(β −1 g(u)).

< 2K1 − u, if u ∈ [u∗ , K1 ),
(Q) b(u)
≥ 2K1 − u, if u ∈ [K1 , 2K1 ].
Theorem 3.5. Assume (A1 )–(A2 ), (A3 ) and (A4 ). If K1 > u∗ (cf. Fig. 3),

u∗ := α −1 h β −1 g α −1 h β −1 g(u∗ ) > u∗ and (Q) or (Q) holds, (3.12)
then, for any δ = (δ1 , δ2 ) > 0 and M > 0, we have

lim w(·, t; ϕ) − K = 0 +
t→∞
Proof. Let (U1 (t), V1 (t)) and (u1 (t), v1 (t)) be the solutions of problems (3.7) and (3.8), respec-
tively. We first prove the following claims.
Claim 1: limt→∞ (u1 (t), v1 (t)) = (k 1 , k 2 ), where

(k 1 , k 2 ) := α −1 h(k 2 ), β −1 g α −1 h β −1 g(u∗ ) .
Let (ũ(t), ṽ(t)) be the solution of the following problem:

⎧
⎨ ũ (t) = −α ũ(t) + hṽ(t − τ1 ) ,
⎪ t > τ,
ũ (t) = −β ũ(t) + g U1 (t − τ2 ) , t > τ,
⎪
⎩
ũ(s), ṽ(s) = e−ατ δ1 , e−βτ δ2 , s ∈ [0, τ ].
One can easily verify that limt→∞ (ũ(t), ṽ(t)) = (k 1 , k 2 ) and (u1 (t), v1 (t)) ≤ (ũ(t), ṽ(t)) for
t > τ . Hence,

(u− , v− ) := lim inf u1 (t), v1 (t) ≤ lim sup u1 (t), v1 (t) ≤ (k 1 , k 2 ).
t→∞ t→∞
If u− > u∗ , then there exists T0 > 0 such that U1 (t) ≥ u1 (t) > u∗ for t > T0 . Since g is
non-increasing on [u∗ , ∞), the function (u1 (t), v1 (t)) satisfies

u1 (t) = −αu1 (t) + h v1 (t − τ1 ) , if t > T0 + τ,

v1 (t) = −βv1 (t) + g U1 (t − τ2 ) , if t > T0 + τ.
Since limt→∞ U1 (t) = α −1 h(β −1 g(u∗ )), we have limt→∞ (u1 (t), v1 (t)) = (k 1 , k 2 ).
Thus, to prove this claim, it suffices to show that u− > u∗ . Suppose u− ≤ u∗ , let {tj }j ∈N be a
sequence with tj → ∞ as j → ∞ such that

lim u1 (tj ), v1 (tj ) = (u− , v− ) and lim u1 (tj ), v1 (tj ) = 0.
j →∞ j →∞
Then we can choose a subsequence of {tj }j ∈N , still denote by {tj }j ∈N , such that (ũ− , ṽ− ) :=
limj →∞ (u1 (tj − τ2 ), v1 (tj − τ1 )) exists. It is clear that (u− , v− ) ≤ (ũ− , ṽ− ) ≤ (k 1 , k 2 ). From
(3.8), we have

αu− = h(ṽ− ) ≥ h(v− ),

βv− = min g(ũ− ), g α −1 h β −1 g(u∗ ) ,
which implies that

β −1 min g(ũ− ), g α −1 h β −1 g(u∗ ) ≤ h−1 (αu− ). (3.13)
By the assumption u∗ > u∗ , it is easy to see that

β −1 g α −1 h β −1 g(u∗ ) > h−1 (αu∗ ) ≥ h−1 (αu− ).
Note that ũ− ≤ k 1 and k 1 < K1 when K1 > u∗ . Thus, β −1 g(ũ− ) > h−1 (α ũ− ) ≥ h−1 (αu− ).
Hence,

β −1 min g(ũ− ), g α −1 h β −1 g(u∗ ) > h−1 (αu− ),
which contradicts to (3.13). Therefore, u− > u∗ and limt→∞ (u1 (t), v1 (t)) = (k 1 , k 2 ).
Moreover, one can see that

(k 1 , k 2 ) K α −1 h β −1 g(u∗ ) , β −1 g(u∗ ) when K1 > u∗ .
By the assumption α −1 h(k 2 ) = u∗ > u∗ , there exits a T1 > 0 such that

(ω1 , ω2 ) ≤ u1 (t), v1 (t) ≤ w(x, t; ϕ), K ≤ U1 (t), V1 (t) , for t > T1 , (3.14)
where (ω1 , ω2 ) := ( 12 (α −1 h(k 2 ) + u∗ ), 12 β −1 g(α −1 h(β −1 g(u∗ ))))

(u∗ , 0).
Next, we construct a sequence of pairs of sub-solution and super-solution of (1.5). Let
(Un (t), Vn (t))(n ≥ 2) and (un (t), vn (t))(n ≥ 2) be the solutions of the following problems:
⎧ ⎧
⎨ Un (t) = −αUn (t) + h Vn−1 (t − τ1 ) ,
⎪ ⎨ un (t) = −αun (t) + h vn−1 (t − τ1 ) ,
⎪
Vn (t) = −βVn (t) + g un−1 (t − τ2 ) , and vn (t) = −βvn (t) + g Un−1 (t − τ2 ) ,
⎪
⎩ ⎪
⎩
Un (s), Vn (s) = K̄, un (s), vn (s) = (ω1 , ω2 ),
for t > T1 + τ and s ∈ [T1 , T1 + τ ], respectively.

Claim 2: We claim that

u1 (t), v1 (t) ≤ · · · ≤ un−1 (t), vn−1 (t) ≤ un (t), vn (t) ≤ K

≤ Un (t), Vn (t) ≤ Un−1 (t), Vn−1 (t) ≤ · · · ≤ U1 (t), V1 (t) , for t > T1 + τ. (3.15)
We prove this claim by induction. By (3.14), one can see that (3.15) holds for n = 1. For n = 2,
we see that

u2 (t) + αu2 (t) = h v1 (t − τ1 ) = u1 (t) + αu1 (t),
for t > T1 + τ,
v2 (t) + βv2 (t) = g U1 (t − τ2 ) ≥ v1 (t) + βv1 (t),
with (u2 (T1 + τ ), v2 (T1 + τ )) = (u1 (T1 + τ ), v1 (T1 + τ )). Hence,

u1 (t), v1 (t) ≤ u2 (t), v2 (t) , for t > T1 + τ.
Note that g is non-increasing on [u∗ , ∞) and h is non-decreasing on [0, ∞). Similarly, we obtain
that

u2 (t), v2 (t) ≤ K and K ≤ U2 (t), V2 (t) ≤ U1 (t), V1 (t) , for t > T1 + τ.
Now, assume that (3.15) holds, let’s show that

un (t), vn (t) ≤ un+1 (t), vn+1 (t) ≤ K ≤ Un+1 (t), Vn+1 (t) ≤ Un (t), Vn (t) ,
for t > T1 + τ . Since (Un+1 (s), Vn+1 (s)) = (Un (s), Vn (s)) for s ∈ [T1 , T1 + τ ], by (3.15), we
obtain

Un+1 (t) + αUn+1 (t) = h Vn (t − τ1 ) ≤ h Vn−1 (t − τ1 ) = Un (t) + αUn (t),

Vn+1 (t) + βVn+1 (t) = g un (t − τ2 ) ≤ g un−1 (t − τ2 ) = Vn (t) + βVn (t)
for t > T1 + τ , which implies that (Un+1 (t), Vn+1 (t)) ≤ (Un (t), Vn (t)) for t > T1 + τ . Similarly,
we have

K ≤ Un+1 (t), Vn+1 (t) and un (t), vn (t) ≤ un+1 (t), vn+1 (t) ≤ K for t > T1 + τ.
Hence the claim follows by using the inductive argument.

Let’s denote (u+ + + +
n , vn ) := limt→∞ (un (t), vn (t)) and (Un , Vn ) := limt→∞ (Un (t), Vn (t)).
Then
+ + + + + + +
u+
1 , v1 ≤ · · · ≤ un , vn ≤ K ≤ Un , Vn ≤ · · · ≤ U1 , V1 ,
+ + +
αUn+ = h Vn−1 , αun = h vn−1 ,
+
+ and +
+ (3.16)
βVn = g un−1 , βvn = g Un−1 .
Set (u+ , v + ) := limn→∞ (u+ + + + + +

n , vn ) and (U , V ) := limn→∞ (Un , Vn ). It then follows from
(3.16) that

αU + = h V + , αu+ = h v + ,
and (3.17)
βV + = g u+ , βv + = g U + ,
which yields that

U + = α −1 h β −1 g u+ = b u+ and u+ = α −1 h β −1 g U + = b U + .
And, we have

U + b U + = b u+ b U + = b u+ u+ .
Suppose that (Q) holds. Since limt→∞ U1 (t) = α −1 h(β −1 g(u∗ )), we have u+ ≤ K1 ≤
U + ≤ α −1 h(β −1 g(u∗ )). Then it must be u+ = K1 = U + , which implies that (u+ , v + ) =
(U + , V + ) = K.
On the other hand, if (Q) holds, by considering two curves U = b(u) and u = b(U ) in the
(u, U )-plane and the similar discussions in Wang and Li [27], we can show that u = b(U ) and
U = b(u) have only one positive intersection point (K1 , K1 ). Hence, (u+ , v + ) = (U + , V + ) = K.
Therefore, we have

lim w(·, t; ϕ) − K = 0 uniformly for ϕ ∈ Cδ,M
+
.
t→∞
This completes the proof. 2
Theorem 3.6. Assume (A1 )–(A2 ), (A3 ) and (A4 ). If K1 > u∗ , then, for any ϕ ∈ C + with
infx∈R ϕ(x, 0) > 0, there holds

k1− , k2− ≤ lim inf inf w(x, t; ϕ) ≤ lim sup sup w(x, t; ϕ) ≤ α −1 h β −1 g(u∗ ) , β −1 g(u∗ ) ,
t→∞ x∈R t→∞ x∈R
where k1− ∈ (0, K1 ) is the unique positive root of the equation β −1 g − (u) = h−1 (αu), g − (·) is
defined in (3.6) and k2− := β −1 g(k1− ).
Proof. Let W0 (t) = (U0 (t), V0 (t)) and w0 (t) = (u0 (t), v0 (t)) be the solutions of the following
problems:
⎧
⎨ U0 (t) = −αU0 (t) + h V0 (t − τ1 ) ,
⎪ t > τ,
V0 (t) = −βV0 (t) + g(u∗ ), t > τ,
⎪
⎩
W0 (s) = K̄, s ∈ [0, τ ],
and
⎧
⎪ u0 (t) = −αu0 (t) + h v0 (t − τ1 ) , t > τ,
⎪
⎨
v0 (t) = −βv0 (t) + min g u0 (t − τ2 ) , g U0 (t − τ2 ) , t > τ,
⎪
⎪
⎩ w0 (s) = e−ατ inf ϕ1 (x, 0), e−βτ inf ϕ2 (x, 0) , s ∈ [0, τ ],
x∈R x∈R
respectively, where li =: sup(x,r)∈R×[−τ,0] ϕi (x, r), i = 1, 2, and

K̄ = (K̄1 , K̄2 ) := max α −1 h(K̄2 ), l1 , max β −1 g(u∗ ), l2 .
It is easy to verify that w0 and W0 are a pair of sub-solution and super-solution of (1.5). From
Theorem 2.3, we have w0 (s) ≤ w(x, s; ϕ) ≤ W0 (s) for (x, s) ∈ R × [0, τ ]. It then follows from
Lemma 2.6 that
w0 (t) ≤ w(x, t; ϕ) ≤ W0 (t) for (x, t) ∈ R × [τ, ∞).
It is clear that

lim W0 (t) = α −1 h β −1 g(u∗ ) , β −1 g(u∗ ) .
t→∞
By constructing a lower auxiliary system via the function g − (·), using a similar method as in the
proof of Theorem 3.4, we can show that lim inft→∞ w0 (t) ≥ (k1− , k2− ), and the assertion of this
theorem follows. 2
Remark 3.7. (i) Note that it’s possible that u∗ ≤ u∗ for some nonlinearities, see Fig. 4. Here we
remark that for general functions g and h, the condition u∗ > u∗ holds when K1 (> u∗ ) is close
to u∗ . Let

f (u) := α −1 h β −1 g α −1 h β −1 g(u) − u.
Direct computations show that f (K1 ) = 0 and

f (K1 ) = α −1 β −1 h β −1 g(K1 ) g (K1 ) + αβ h β −1 g(K1 ) g (K1 ) − αβ .
Since g (u∗ ) = 0, g (K1 ) < 0 and h (u) > 0 for u > 0, we have f (K1 ) < 0 for 0 < K1 − u∗ 1
which implies that f (u∗ ) > 0 for 0 < K1 − u∗ 1. Therefore, the condition u∗ > u∗ holds for
0 < K1 − u∗ 1.
Fig. 4. The graphs of g and h with K1 > u∗ > u∗ := α −1 h(β −1 g(α −1 h(β −1 g(u∗ )))).
In the following, we give a specific example to support the condition (3.12) and main results
for the non-monotone function g.
Assume h(v) = v and g(u) = pue−δu , where p, δ > 0. It’s obvious that u∗ = δ −1 . If p >
αβ, it’s easy to verify that system (1.5) has a unique positive equilibrium K = (K1 , K2 ) :=
(δ −1 ln αβ
p
, αK1 ). Moreover, we have
(i) K1 ≤ u∗ , if αβ < p ≤ αβe, and (ii) K1 > u∗ , if p > αβe.
Direct computation shows that the condition u∗ > u∗ yields to
p p
2 ln − − 1 > 0.
αβ αβe
It is easy to see that the above inequality holds for αβe < p ≤ αβe2 . Moreover, it is not difficult
to show that (Q) holds when αβe < p ≤ αβe2 . According to Theorems 3.3–3.6, the following
statements hold:
(1) If 0 < p ≤ αβ, then the zero equilibrium 0 is global attractive.

(2) If αβ < p ≤ αβe, then K1 ≤ u∗ and the positive equilibrium K is global attractive.
(3) If αβe < p ≤ αβe2 , then K1 > u∗ and the positive equilibrium K is global attractive.
(4) If p > αβe2 , then C[(k − ,k − ),(k + ,k + )] is a global attractor, where
1 2 1 2

p p p 2 + pαβδe p + αβδe
k1+ , k2+ := , , k1− = exp − and k2− = αk1− .
αβδe βδe α 2 β 2 δe αβe
In addition, if we replace X = BUC(R, R2 ) with X = R2 , then our main results, i.e. Theo-
rems 3.3, 3.4, 3.5 and 3.6 hold for the following delay system:
⎧
⎨ u (t) = −αu(t) + hv(t − τ1 ), t > 0,
⎪
v (t) = −βv(t) + g u(t − τ2 ) , t > 0, (3.18)
⎪
⎩
u(s), v(s) = ϕ(s), s ∈ [−τ, 0].
In fact, our result can be easily extended to the delayed nonlocal system (1.5) on a bounded
domain with Neumann boundary conditions. We leave the details to the readers.
4. Spreading speed and traveling wave solutions
In this section, we first state some results on monotone traveling wave solutions and spreading
speed of (1.7) in quasi-monotone case. Then by squeezing method combining with two auxiliary
systems, we obtain the minimal wave speed and spreading speed of (1.7) without satisfying
the quasi-monotone condition. By applying a fluctuation method, we also provide some suffi-
cient conditions to ensure the upward convergence of the spreading speed and traveling wave
solutions. Our results reveal that the minimal wave speed coincides with the spreading speed.
Finally, we point out the effects of the delay and nonlocality on the spreading speed of the non-
quasi-monotone system.
We first give the definition of spreading speed, see e.g. [1,8,18,25,30].
Definition 4.1. A function w : R × R+ → R2+ is said to have a spreading speed c∗ > 0 if there
exists some vector ε
0 such that
lim w(x, t) = 0 for all c > c∗ and lim inf w(x, t) ≥ ε for all c ∈ (0, c∗ ).
t→∞,|x|≥ct t→∞,|x|≤ct
According to Definition 4.1, we see that c∗ and ε depend on the function w. However, if all
solutions w(x, t) of a system with initial functions having compact supports share the same c∗
and ε, then we call such c∗ the spreading speed of the system.
4.1. Results for monotone delayed systems
In this subsection, we give the following additional assumption:
(B3 ) ∂j fi (w) ≥ 0 and F (w) ≤ DF(0)w for all 1 ≤ i = j ≤ 2 and w ∈ [0, K], i = 1, 2.
For convenience, we denote
αi = ∂i fi (0, 0) < 0 and βi = ∂j fi (0, 0) > 0 for 1 ≤ i = j ≤ 2.
It’s easy to see that the characteristic function of system (1.7) with respect to the trivial equilib-
rium 0 has the form:
1 (c, λ) := P1 (c, λ) − P2 (c, λ), for c ≥ 0 and λ ∈ C, (4.1)
where

P1 (c, λ) := d1 λ2 − cλ + α1 d2 λ2 − cλ + α2
+∞ +∞
−cλ(τ1 +τ2 ) −λy
P2 (c, λ) := β1 β2 e J1 (y)e dy J2 (y)e−λy dy.
−∞ −∞
Let’s denote

c+ c2 − 4d1 α1 c+ c2 − 4d2 α2
λc1 := , λc2 := , if d2 > 0,
2d1 2d2

min{λc1 , λc2 }, if d2 > 0,
λcm :=
λc1 , if d2 = 0.
Note that P1 (c, λcm ) = 0 and α1 α2 < β1 β2 . Similar to [33, Lemma 2.1], we can obtain the fol-
lowing result.
Lemma 4.2. There exists a positive number c∗ such that
(1) if c ≥ c∗ , then the equation 1 (c, λ) = 0 has two positive real roots Λ1 := Λ1 (c) and Λ2 :=
Λ2 (c) with 0 < Λ1 (c) ≤ Λ2 (c);
(2) if c = c∗ , then Λ1 (c∗ ) = Λ2 (c∗ ) := Λ∗ , and if c > c∗ , then Λ1 (c) < Λ∗ < Λ2 (c), 1 (c, ·) <
0 in (0, λcm ) \ (Λ1 (c), Λ2 (c)) and 1 (c, ·) > 0 in (Λ1 (c), Λ2 (c)).
In the quasi-monotone case, by the theory for monotone semiflow developed in Liang and
Zhao [18], as applied to the solution semiflow associated with (1.7), we can prove that c∗ is the
spreading speed of system (1.7). Here, we omit the proof of this result since it is essentially the
same as that of [18, Theorem 5.1].
Theorem 4.3. Assume (A1 ) and (B1 )–(B3 ). For any ϕ ∈ C[0,K] , system (1.7) has a unique solution
w(x, t) with w(x, s) = ϕ(x, s) for x ∈ R and s ∈ [−τ, 0] and 0 ≤ w(x, t) ≤ K for x ∈ R and
t ≥ 0. Furthermore, the following statements hold:
(1) For any c > c∗ , if 0 ≤ ϕ K and ϕ(x, ·) = 0 for x outside a bounded interval, then
limt→∞,|x|≥ct w(x, t) = 0,
(2) For any c < c∗ and any vector σ ∈ R2 with 0 σ ≤ K, there is a positive number
rσ such that if ϕ ∈ C[0,K] and ϕ(x, ·)
σ for x on an interval of length 2rσ , then
limt→∞,|x|≤ct w(x, t) = K.
Similar to Theorem 2.3, the existence and uniqueness of solutions for the Cauchy problem of
(1.7) can be obtained by using the theory of abstract functional differential equations developed
by Martin and Smith [21], see also [16].
Now, we consider the traveling wave solutions of (1.7). Let’s substitute

u(x, t), v(x, t) = φc (ξ ), ψc (ξ ) , ξ = x + ct,
for some functions φc , ψc into (1.7), then we obtain the corresponding wave system

cφc (ξ ) = d1 φc (ξ ) + f1 φc (ξ ), (J1 ∗ ψc )(ξ ) ,
(4.2)
cψc (ξ ) = d2 ψc (ξ ) + f2 (J2 ∗ φc )(ξ ), ψc (ξ ) ,
where
+∞
(Ji ∗ ϕ)(ξ ) = Ji (y)ϕ(ξ − y − cτi )dy, for i = 1, 2 and ϕ ∈ BC(R, R).
−∞
We point out that the existence of monotone traveling wave solutions with c > c∗ can be
obtained by applying the Liang and Zhao’s theory. However, we still provide a sketch of the proof
for the existence of the monotone traveling wave solutions with c > c∗ by using the method of
sub-solutions and super-solutions (cf. [19]), because we need some more specific information on
the asymptotic behavior of the traveling wave solutions, which will be used in later sections for
proving the existence and asymptotic behavior of the traveling wave solutions for non-monotone
delayed systems.
According to [19, Theorem 2.2], the existence of solutions of (4.2) can be reduced to the exis-
tence of an appropriate pair of super-solution and sub-solution of (4.2). See [19, Definition 2.1].
By Lemma 4.2, we are ready to construct a pair of super-solution and sub-solution of (4.2).
Lemma 4.4. Let (A1 ) and (B1 )–(B3 ) hold, c > c∗ , μ ∈ (1, min{2, Λ2 /Λ1 }) and q > 0. Define

Φ̄c (ξ ) = φ̄c (ξ ), ψ̄c (ξ ) := min K1 , eΛ1 ξ , min K2 , b(c)eΛ1 ξ ,

Φ̂c (ξ ) = φ̂c (ξ ), ψ̂c (ξ )

:= max 0, eΛ1 ξ − qeμΛ1 ξ , max 0, b(c)eΛ1 ξ − qB(c, μ)eμΛ1 ξ ,

+∞
where b(c) := β2 e−cΛ1 τ2 −∞ J2 (y)e−Λ1 y dy/(cΛ1 − d2 Λ21 − α2 ) and

+∞
β2 e−cΛ1 μτ2 −∞ J2 (y)e−Λ1 μy dy c(Λ1 μ) − d1 (Λ1 μ)2 − α1
< B(c, μ) <
+∞ .
c(Λ1 μ) − d2 (Λ1 μ)2 − α2 β1 e−cΛ1 μτ1 −∞ J1 (y)e−Λ1 μy dy
Then Φ̄c (ξ ) is a super-solution of (4.2), and Φ̂c (ξ ) is a sub-solution of (4.2) provided

q > max{1, b(c)/B(c, μ)} is large enough.
Proof. Using the assumptions F ∈ C 2 (R2+ , R2 ) and (B3 ), the proof is direct and easy. We omit
it here. 2
Based on Lemma 4.2, Theorem 4.3 and the above construction of super-solution and sub-
solution of (4.2), we can prove the following existence and non-existence of monotone traveling
wave solutions of (1.7).
Theorem 4.5. Assume (A1 ) and (B1 )–(B3 ). Then, the following result holds.
(1) For each c ≥ c∗ , system (1.7) has a monotone traveling wave Φc (ξ ) := (φc (ξ ), ψc (ξ )), ξ =
x + ct, which satisfies
Φc (−∞) = 0, Φc (+∞) = K and Φc (ξ )

0 for all ξ ∈ R.
Moreover, if c > c∗ , then
lim Φc (ξ )e−Λ1 (c)ξ = νc and Φc (ξ ) ≤ eΛ1 (c)ξ νc (4.3)

ξ →−∞
for all ξ ∈ R, where νc = (1, b(c)).

(2) For 0 < c < c∗ , (1.7) has no traveling wave solutions connecting 0 and K.
Proof. The existence of monotone traveling wave solutions with speed c > c∗ follows from
Lemma 4.4 and [19, Theorem 2.2]. From the constructions of the super-solution and sub-solution,
we have the asymptotic behavior (4.3).
For the case c = c∗ , the existence of the monotone traveling wave could be obtained by using
a limiting argument, see e.g. [25, Theorem 3.4].
The non-existence of traveling wave solutions is a direct consequence of Theorem 4.3. This
proof is complete. 2
From Theorems 4.3 and 4.5, we see that the spreading speed of (1.7) coincides with the
minimal wave speed when (1.7) is quasi-monotone. In what follows, we shall extend the result
to the non-quasi-monotone case. To overcome the difficulty for the non-quasi-monotone delayed
system, we need the following additional assumptions:
(B3 ) There exist K± = (K1± , K2± )

0 with 0 K− ≤ K ≤ K+ and two continuous and twice
piecewise continuous differentiable functions F ± := (f1± , f2± ) : [0, K+ ] → R2 such that
(i) F ± (0) = F + (K+ ) = F − (K− ) = 0 and there is no other positive zeros of F ± between
0 and K± ;
(ii) ∂1 f1± , ∂2 f2± ∈ C([0, K+ ], R) and there exists a constant L1 > 0 such that

f1± (u, v1 ) − f1± (u, v2 ) ≤ L1 max{0, v1 − v2 }, for all (u, v1 ), (u, v2 ) ∈ 0, K+ ,

f2± (u1 , v) − f2± (u2 , v) ≤ L1 max{0, u1 − u2 }, for all (u1 , v), (u2 , v) ∈ 0, K+ ;
(iii) fi± (w) are non-decreasing with respect to wj for 1 ≤ i = j ≤ 2 and w ∈ [0, K+ ];
(iv) F (w) and F ± (w) have the same Jacobian matrix DF(0) at w = 0 and

F − (w) ≤ F (w) ≤ F + (w) ≤ DF(0)w for all w ∈ 0, K+ .
Remark 4.6. Based on (B3 ) , we can construct two quasi-monotone systems whose reaction
terms are F + and F − , respectively, to “trap” system (1.7). If ∂j fi (w) ≥ 0 for 1 ≤ i = j ≤ 2
and w ∈ [0, K], i.e. (1.7) is quasi-monotone on [0, K], then it is natural to choose F ± (·) =
F (·). In Subsection 4.5, we shall give the constructions of the auxiliary functions F ± for a
specific epidemic model. We also note that the assumption (B3 ) (ii) is only used to establish the
comparison theorem (see Lemma 4.7).
4.2. Spreading speeds
In this subsection, we establish the spreading speed and the non-existence of traveling wave
solutions for (1.7) in non-quasi-monotone case.
We remark that the well-posedness result for the initial value problem of (1.7) follows from the
theory of abstract functional differential equations developed by Martin and Smith [21], see also
[16]. More precisely, for any ϕ ∈ C[0,K+ ] , system (1.7) has a unique global solution w(x, t; ϕ)
with w(x, s; ϕ) = ϕ(x, s) and 0 ≤ w(x, t; ϕ) ≤ K+ for x ∈ R, s ∈ [−τ, 0], t ≥ 0. Hereafter, we
define C[0,K± ] by replacing K with K+ and K− in the definition of C[0,K] , respectively.
According to (B3 ) , we consider the following two auxiliary monotone delayed systems:

ut = d1 uxx + f1+ u(x, t), (J1 ∗ v)(x, t) ,
(4.4)
vt = d2 vxx + f2+ (J2 ∗ u)(x, t), v(x, t) ,
and

ut = d1 uxx + f1− u(x, t), (J1 ∗ v)(x, t) ,
(4.5)
vt = d2 vxx + f2− (J2 ∗ u)(x, t), v(x, t) ,
where x ∈ R, t ∈ R. To obtain the spreading speed of (1.7) in the non-quasi-monotone case, we

need to establish a comparison theorem for solutions to systems (1.7), (4.4) and (4.5). Note that
∂1 f1± , ∂2 f2± ∈ C([0, K+ ], R). We first introduce some notations and definitions. Let’s denote

L̄ := max max ∂i fi+ (w), ∂i fi− (w) (4.6)
w∈[0,K+ ] i=1,2
and Q(φ) = (Q1 (φ), Q2 (φ)) by

∞
Q1 (φ)(x) := f1 φ1 (x, 0), J1 (x − y)φ2 (y, −τ1 )dy + L̄φ1 (x, 0),
−∞
∞
Q2 (φ)(x) := f2 J2 (x − y)φ1 (y, −τ2 )dy, φ2 (x, 0) + L̄φ2 (x, 0),
−∞
for φ = (φ1 , φ2 ) ∈ C[0,K+ ] . Similarly, we define Q± (φ) := (Q± ±

1 (φ), Q2 (φ)) by replacing
F (φ) = (f1 (φ), f2 (φ)) with F (φ) = (f1 (φ), f2 (φ)) and F (φ) = (f1 (φ), f2− (φ)), respec-
+ + + − −
tively. It is clear that Q± (·) is non-decreasing in C[0,K+ ] and
Q− (φ) ≤ Q(φ) ≤ Q+ (φ), for φ ∈ C[0,K+ ] .
Furthermore, we denote the linear operator T̃ (t) = diag(T̃1 (t), T̃2 (t)) by T̃i (0) = I and

T̃i (t)[ϕ](x) := Γ˜i (x − y, t)ϕ(y)dy, for x ∈ R, t > 0, ϕ ∈ BUC(R, R),
R
where Γ˜1 (x, t) := e(α−L̄)t Γ1 (x, t) and Γ˜2 (x, t) := e(β−L̄)t Γ2 (x, t).
Then we have the following comparison results.
Lemma 4.7. Assume (A1 ), (B1 ), (B2 ) and (B3 ) . For any ϕ, ϕ + ∈ C[0,K+ ] and ϕ − ∈ C[0,K− ] with
ϕ − (x, s) ≤ ϕ(x, s) ≤ ϕ + (x, s) for x ∈ R and s ∈ [−τ, 0],
let w − (x, t; ϕ − ), w(x, t; ϕ) and w + (x, t; ϕ + ) be the solutions of systems (4.5), (1.7) and (4.4)
through ϕ − , ϕ and ϕ + , respectively. Then,

w − x, t; ϕ − ≤ w(x, t; ϕ) ≤ w + x, t; ϕ + , for x ∈ R and t ≥ 0.
Proof. For convenience, we denote w − (x, t; ϕ − ), w(x, t; ϕ) and w + (x, t; ϕ + ) by w − (x, t),
w(x, t) and w + (x, t), respectively. It is clear that 0 ≤ w(x, t), w ± (x, t) ≤ K+ for all x ∈ R,
t ≥ −τ . We only prove that w(x, t) ≤ w + (x, t) for x ∈ R and t ≥ 0, since the other case can be
proved similarly. Let

z(x, t) = z1 (x, t), z2 (x, t) := w(x, t) − w + (x, t), x ∈ R, t ≥ −τ.
Note that w(x, s) ≤ w + (x, s) for x ∈ R and s ∈ [−τ, 0], and
t

w(t)(x) = T̃ (t) ϕ(0) (x) + T̃ (t − s) Q w s (x)ds, (4.7)
0
t

w (t)(x) = T̃ (t) ϕ + (0) (x) +
+
T̃ (t − s) Q+ w +s (x)ds. (4.8)
0
There are four cases considered in the sequel.

Case 1: τ1 > 0 and τ2 > 0.
Since fi+ (w) are non-decreasing with respect to wj for 1 ≤ i = j ≤ 2 and w ∈ [0, K+ ] and
L̄ + ∂1 f1+ (w) ≥ 0 for any w ∈ [0, K+ ], for any x ∈ R and t ∈ [0, τ1 ], we have
+t
Q1 w t (x) − Q+ 1 w (x)

≤ Q+ 1 w (x) − Q1 w
t + +t
(x)
+
= L̄z1 (x, t) + f1 u(x, t), (J1 ∗ v)(x, t) − f1+ u+ (x, t), J1 ∗ v + (x, t)

≤ L̄z1 (x, t) + f1+ u(x, t), J1 ∗ v + (x, t) − f1+ u+ (x, t), J1 ∗ v + (x, t)

= L̄ + ∂1 f1+ θ u(x, t) + (1 − θ )u+ (x, t), J1 ∗ v + (x, t) z1 (x, t)

≤ 2L̄ max 0, z1 (x, t) , (4.9)
where θ ∈ (0, 1). Let [B]+ := max{B, 0} for B ∈ R. It then follows from (4.7)–(4.9) that
t
+s
z1 (t)(x) = T̃1 (t) z1 (0) (x) + T̃1 (t − s) Q1 w s − Q+
1 w (x)ds
0
t +∞

≤ 2L̄ Γ˜1 (x − y, t − s) z1 (y, s) + dyds, for any x ∈ R and t ∈ [0, τ1 ],
0 −∞
which implies that
t +∞

z1 (x, t) + ≤ 2L̄ Γ˜1 (y, s) z1 (x − y, t − s) + dyds. (4.10)
0 −∞
Moreover, for any λ > 0, we set

z̃1,λ (t) := sup z1 (x, t) + e−λt and z̃1,λ := sup z̃1,λ (t).
x∈R t∈[0,τ1 ]
From (4.10), for any t ∈ [0, τ1 ], we can obtain
t +∞ τ1
˜ −λs +
z̃1,λ (t) ≤ 2L̄ Γ1 (y, s)z̃1,λ (t − s)e dyds ≤ 2L̄K1 e−(λ+L̄)s ds.
0 −∞ 0
Then we have

z̃1,λ ≤ 2L̄K1+ 1 − e−(λ+L̄)τ1 /(λ + L̄) → 0 as λ → +∞.
Hence, z̃1,λ ≤ 0 for sufficiently large λ, which implies that z1 (x, t) ≤ 0 for x ∈ R and t ∈ [0, τ1 ].
Similarly, we can show that z2 (x, t) ≤ 0 for x ∈ R and t ∈ [0, τ2 ]. Thus, z(x, t) ≤ 0 for x ∈ R and
t ∈ [0, τm ]. Recall that τm = min{τ1 , τ2 }. By the same argument, we can obtain z(x, t) ≤ 0 for
x ∈ R and t ∈ [τm , 2τm ]. Inductively, we obtain that z(x, t) ≤ 0 for x ∈ R and t ≥ 0, and hence
w(x, t) ≤ w + (x, t) for x ∈ R and t ≥ 0.
Case 2: τ1 > 0 and τ2 = 0.
Similar to Case 1, we can show that z1 (x, t) ≤ 0 for x ∈ R and t ∈ [0, τ1 ], that is u(x, t) ≤
u+ (x, t) for x ∈ R and t ∈ [0, τ1 ]. Note that L̄ + ∂2 f2+ (w) ≥ 0 for any w ∈ [0, K+ ] and when
τ2 = 0,

(J2 ∗ q)(x, t) = J2 (x − y)q(y, t)dy for q = u or u+ .
R
Thus, for x ∈ R and t ∈ [0, τ1 ],

+t
Q2 w t (x) − Q+ 2 w (x)

≤ L̄z2 (x, t) + f2+ J2 ∗ u+ (x, t), v(x, t) − f2+ J2 ∗ u+ (x, t), v + (x, t)

= L̄ + ∂2 f2+ J2 ∗ u+ (x, t), θ v(x, t) + (1 − θ )v + (x, t) z2 (x, t)

≤ 2L̄ max 0, z2 (x, t) , where θ ∈ (0, 1).
By the same arguments as above, we can obtain z2 (x, t) ≤ 0 for x ∈ R and t ∈ [0, τ1 ]. Hence,
z(x, t) ≤ 0 for x ∈ R and t ∈ [0, τ1 ]. Inductively, we obtain that z(x, t) ≤ 0 for x ∈ R and t ≥ 0.
Therefore, w(x, t) ≤ w+ (x, t) for x ∈ R and t ≥ 0.
Case 3: τ2 > 0 and τ1 = 0.
The proof is similar to that for the Case 2 and omitted.
Case 4: τ1 = τ2 = 0.
By part (ii) of the assumption (B3 ) , for any x ∈ R and t ∈ [0, ∞), we have
+t
Q1 w t (x) − Q+ 1 w (x)

≤ L̄z1 (x, t) + f1+ u(x, t), (J1 ∗ v)(x, t) − f1+ u+ (x, t), (J1 ∗ v)(x, t)

+ f1+ u+ (x, t), (J1 ∗ v)(x, t) − f1+ u+ (x, t), J1 ∗ v + (x, t)

≤ 2L̄ max 0, z1 (x, t) + L1 max 0, (J1 ∗ z2 )(x, t) . (4.11)
Then it follows from (4.7), (4.8) and (4.11) that
t +∞

z1 (x, t) + ≤ L Γ˜1 (y, s) z1 (x − y, t − s) + + J1 ∗ [z2 ]+ (x − y, t − s) dyds,
0 −∞
where L := L1 + 2L̄. Similarly, we obtain
t +∞

z2 (x, t) + ≤ L Γ˜2 (y, s) z2 (x − y, t − s) + + J2 ∗ [z1 ]+ (x − y, t − s) dyds.
0 −∞
Using the similar method as in the proof of Lemma 2.6, we can show that w(x, t) ≤ w+ (x, t) for
x ∈ R and t ≥ 0. The proof is complete. 2
Applying the comparison Lemma 4.7, we have the following results on the spreading speed
for (1.7) in the non-quasi-monotone case. Recall that b(u) = α −1 h(β −1 g(u)).
Theorem 4.8. Assume (A1 ), (B1 ), (B2 ) and (B3 ) . For any ϕ ∈ C[0,K+ ] , system (1.7) has a unique
solution w(x, t; ϕ) = (u(x, t; ϕ), v(x, t; ϕ)) with w(x, s; ϕ) = ϕ(x, s) for x ∈ R and s ∈ [−τ, 0]
and 0 ≤ w(x, t; ϕ) ≤ K+ for x ∈ R and t ≥ 0. Furthermore, the following statements hold:
(1) For any c > c∗ , if 0 ≤ ϕ K+ and ϕ(x, ·) = 0 for x outside a bounded interval, then
limt→∞,|x|≥ct w(x, t; ϕ) = 0,
(2) For any c < c∗ and any vector σ ∈ R2 with 0 σ ≤ K+ , there is a positive number rσ such
that if ϕ ∈ C[0,K+ ] and ϕ(x, ·)
σ for x on an interval of length 2rσ , then
K− ≤ lim inf w(x, t; ϕ) ≤ lim sup w(x, t; ϕ) ≤ K+ .

t→∞,|x|≤ct t→∞,|x|≤ct
Moreover, if f1 (u, v) = −αu + h(v), f2 (u, v) = −βv + g(u), (A2 ), (A4 ), and one of the
following assumptions hold:
(i) (A3 ) holds and K1 ≤ u∗ ;
(ii) b(u)/u is strictly decreasing for u ∈ [K1− , K1+ ] and b(u) satisfies the property:
(P) ∀u1 , u2 ∈ [K1− , K1+ ] satisfying u2 ≤ K1 ≤ u1 , u2 ≥ b(u1 ) and u1 ≤ b(u2 ), there
holds u1 = u2 ,
then limt→∞,|x|≤ct w(x, t; ϕ) = K.
Proof. The proof is similar to those of [13, Theorem 2.3] and [8, Theorem 3.3]. For any ϕ ∈
C[0,K+ ] , define ϕ̃ ∈ C[0,K− ] by ϕ̃(x, t) = min{ϕ(x, t), K− }. From Lemma 4.7, we have
w − (x, t; ϕ̃) ≤ w(x, t; ϕ) ≤ w + (x, t; ϕ) for x ∈ R and t ≥ 0.
Further, by Theorem 4.5, c∗ is the spreading speed of solutions for both systems (4.4) and (4.5),
which together with the above inequalities implies that c∗ satisfies the statements (1) and the first
part of (2). That is, c∗ is the spreading speed of system (1.7).
Now, we prove the upward convergence in the property of spreading speeds. Suppose the con-
dition (i) holds, we can choose F ± (·, ·) = F (·, ·) and K± = K. Hence, the upward convergence
result follows.
Next, assume the condition (ii), we shall use the fluctuation method (see [7,8,13,24]) to prove
the upward convergence of the spreading speed. First, we simplify the notation w(x, t; ϕ) =
(u(x, t; ϕ), v(x, t; ϕ)) by w(x, t) = (u(x, t), v(x, t)) and define G(·, ·) ∈ C(R2+ , R) by

minw∈[u,v] g(w), if u ≤ v,
G(u, v) :=
maxw∈[v,u] g(w), if v ≤ u.
Clearly, G(u, v) is non-decreasing in u and non-increasing in v, and G(u, u) = g(u). Moreover,

for any ν ∈ (0, c∗ ), we set
U∗ (ν) := lim inf u(x, t) and U ∗ (ν) := lim sup u(x, t),
t→∞,|x|≤νt t→∞,|x|≤νt
∗
V∗ (ν) := lim inf v(x, t) and V (ν) := lim sup v(x, t).
t→∞,|x|≤νt t→∞,|x|≤νt
Note that system (1.5) can be rewritten in the following way:
∞
u(x, t) = e−αt Γ¯1 (x − y, t)u(y, 0)dy
−∞
t +∞

+ e−α(t−s) Γ¯1 (y, t − s)h (J1 ∗ v)(x − y, s) dyds
0 −∞
∞
= e−αt Γ¯1 (x − y, t)u(y, 0)dy
−∞
0 +∞

+ eαs Γ¯1 (y, −s)h (J1 ∗ v)(x − y, t + s) dyds, (4.12)
−t −∞
and
∞
v(x, t) = e−βt Γ¯2 (x − y, t)v(y, 0)dy
−∞
t +∞

+ e−β(t−s) Γ¯2 (y, t − s)g (J2 ∗ u)(x − y, s) dyds
0 −∞
∞ 0 +∞
= e −βt
Γ¯2 (x − y, t)v(y, 0)dy + eβs Γ¯2 (y, −s)
−∞ −t −∞

× G (J2 ∗ u)(x − y, t + s), (J2 ∗ u)(x − y, t + s) dyds, (4.13)
where Γ¯1 (x, t) := Γ1 (x, t)eαt , Γ¯2 (x, t) := Γ2 (x, t)eβt , and Γ1 and Γ2 are defined in (2.3) and
(2.4), respectively.
Let c ∈ (0, c∗ ) be given. We fix a number γ ∈ (c, c∗ ) and define
U∗ (c, γ ) := inf U∗ (ω) and U ∗ (c, γ ) := sup U ∗ (ω),

c<ω<γ c<ω<γ
V∗ (c, γ ) := inf V∗ (ω) and V ∗ (c, γ ) := sup V ∗ (ω).

c<ω<γ c<ω<γ
For any ν ∈ (c, γ ), we can choose sequences {tj } in (0, ∞) and {xj } in R such that |xj | ≤ νtj ,
tj → ∞ as j → ∞ and limj →∞ u(xj , tj ) = U∗ (ν). Moreover, for any ι ∈ R and ω ∈ R, we see
that |xj − ω| ≤ γ (tj + ι) when j large enough. Then, we have
lim inf v(xj − y − z, tj + s − τ1 ) ≥ V∗ (γ ).

j →∞
Note that h (v) > 0 for v ∈ [0, +∞). Using Fatou’s lemma, it then follows from (4.12) that
0 +∞

U∗ (ν) ≥ lim inf eαs Γ¯1 (y, −s)h (J1 ∗ v)(xj − y, tj + s) dyds
j →∞
−tj −∞
0 +∞
≥ eαs Γ¯1 (y, −s)h lim inf(J1 ∗ v)(xj − y, tj + s) dyds
j →∞
−∞ −∞
0 +∞ +∞
αs ¯
≥ e Γ1 (y, −s)h J1 (z) lim inf v(xj − y − z, tj + s − τ1 )dz dyds
j →∞
−∞ −∞ −∞
0 +∞

≥ eαs Γ¯1 (y, −s)h V∗ (γ ) dyds = h V∗ (γ ) /α,
−∞ −∞
which implies that

αU∗ (ν) ≥ h V∗ (γ ) .
Similarly, from (4.12) and (4.13), we can show that

αU ∗ (ν) ≤ h V ∗ (γ ) , βV∗ (ν) ≥ G U∗ (γ ), U ∗ (γ ) and βV ∗ (ν) ≤ G U ∗ (γ ), U∗ (γ ) .
Then it follows that

αU∗ (c, γ ) ≥ h V∗ (c, γ ) , αU ∗ (c, γ ) ≤ h V ∗ (c, γ ) ,

βV∗ (c, γ ) ≥ G U∗ (c, γ ), U ∗ (c, γ ) and βV ∗ (c, γ ) ≤ G U ∗ (c, γ ), U∗ (c, γ ) . (4.14)
By the definition of function G(·, ·), we can find u1 , u2 ∈ [U∗ (c, γ ), U ∗ (c, γ )] ⊂ [K1− , K1+ ] such
that

G U∗ (c, γ ), U ∗ (c, γ ) = g(u1 ) and G U ∗ (c, γ ), U∗ (c, γ ) = g(u2 ).
Hence, (4.14) implies

b(u1 ) = α −1 h g(u1 )/β ≤ α −1 h V∗ (c, γ ) ≤ U∗ (c, γ )

≤ u1 , u2 ≤ U ∗ (c, γ ) ≤ α −1 h V ∗ (c, γ ) ≤ α −1 h g(u2 )/β = b(u2 ). (4.15)
Thus, we have
b(u1 ) b(K1 ) b(u2 )

≤1= ≤ .
u1 K1 u2
Combing the above inequality and the strict monotonicity of b(u)/u on [K1− , K1+ ], we conclude
that u2 ≤ K1 ≤ u1 . Moreover, by (4.15) and the property (P), we can obtain u1 = u2 . Hence,
u1 = u2 = K1 . It then follows from (4.15) that
U∗ (c, γ ) = U ∗ (c, γ ) = u1 = u2 = K1 and V∗ (c, γ ) = V ∗ (c, γ ) = K2 .
Consequently, we have
K1 = U∗ (c, γ ) ≤ U∗ (c) ≤ U ∗ (c) ≤ U ∗ (c, γ ) = K1 , and

K2 = V∗ (c, γ ) ≤ V∗ (c) ≤ V ∗ (c) ≤ V ∗ (c, γ ) = K2 ,
which imply that limt→∞,|x|≤ct w(x, t; ϕ) = K. This completes the proof. 2
Remark 4.9. As pointed out by Fang et al. [8], Fang and Zhao [7], and Hsu and Zhao [13], either
of the following two conditions is sufficient for (P) to hold:
(P1 ) ub(u) is strictly increasing for u ∈ [K1− , K1+ ], or

(P2 ) b(u) is non-increasing for u ∈ (K1 , K1+ ] and b(b(u))/u is strictly decreasing for u ∈
(0, K1 ].
The non-existence of traveling wave solutions is a consequence of the result on the spreading
speed. Since its proof is similar to that of [25, Theorem 3.5], see also [26, Theorem 2.1]. Thus,
we only state the non-existence result and skip the proof.
Theorem 4.10. Assume (A1 ), (B1 ), (B2 ) and (B3 ) . For 0 < c < c∗ , (1.7) does not admit a trav-
eling wave solution Φ(ξ ) with lim infξ →∞ Φ(ξ )
0 and Φ(−∞) = 0.
4.3. Existence of minimal wave speed
In this subsection, we consider the existence of traveling wave solutions of (1.7) in the non-
quasi-monotone case. Let c∗ be the spreading speed of (1.7) established in Theorem 4.8. We shall
prove that for any c ≥ c∗ , (1.7) has a traveling wave solution (Φ, c) with Φ(−∞) = 0. This result
together with Theorems 4.8 and 4.10 imply that the spreading speed of (1.7) coincides with the
minimal wave speed in the non-quasi-monotone case.
To find the traveling wave solutions of (1.7) with speed c > c∗ , we will employ Schauder’s
fixed theorem and construct a profile set in a suitable Banach space by using the monotone
traveling wave solutions of the lower auxiliary system (4.5) and its asymptotic behavior at −∞.
Such construction of the profile set provides us a merit to obtain the asymptotic behavior of the
traveling wave solutions at −∞ (see (4.17) below).
Since F (w) and F ± (w) have the same Jacobian matrix DF(0) at w = 0, it is clear that
1 (c, λ) = 0 is also the characteristic equation of (4.4) and (4.5) with respect to the trivial equi-
librium 0. Therefore, by Theorem 4.5, we have the following result on the monotone traveling
wave solutions for the lower auxiliary system (4.5).
Proposition 4.11. Assume (A1 ), (B1 ), (B2 ) and (B3 ) . For any c > c∗ , (4.5) has a non-decreasing
traveling wave solution Φc− (ξ ) = (φc− (ξ ), ψc− (ξ )), ξ = x + ct, which satisfies
Φc− (·)
0, Φc− (−∞) = 0, Φc− (+∞) = K− ,
lim Φc− (ξ )e−Λ1 (c)ξ = νc and Φc− (ξ ) ≤ νc eΛ1 (c)ξ for all ξ ∈ R.
ξ →−∞
Now, we state the main result in this subsection.
Theorem 4.12. Assume (A1 ), (B1 ), (B2 ) and (B3 ) . For each c > c∗ , (1.7) admits a traveling
wave solution Φc (ξ ) such that
Φc (−∞) = 0, 0 Φc (ξ ) ≤ K+ , for ξ ∈ R, (4.16)

K− ≤ lim inf Φc (ξ ) ≤ lim sup Φc (ξ ) ≤ K+ and lim Φc (ξ )e−Λ1 (c)ξ = νc . (4.17)
ξ →∞ ξ →∞ ξ →−∞
Moreover, if f1 (u, v) = −αu +h(v), f2 (u, v) = −βv +g(u), (A2 ), (A4 ), and one of the following
assumptions hold:

(ii) b(u)/u is strictly decreasing for u ∈ [K1− , K1+ ] and b(u) satisfies the property (P),
then limξ →∞ Φc (ξ ) = K.
Proof. The proof of the first assertion is similar to those of [19, Theorem 1.1] and [13, Theo-
rem 3.1]. So, we only sketch the outline in the following three steps.
Step 1. Let’s define the operator H = (H1 , H2 ) : C(R, [0, K+ ]) → C(R, R2 ) by

H1 (φ, ψ)(ξ ) := L̃φ(ξ ) + f1 φ(ξ ), (J1 ∗ ψ)(ξ ) ,

H2 (φ, ψ)(ξ ) := L̃ψ(ξ ) + f2 (J2 ∗ φ)(ξ ), ψ(ξ ) , (4.18)
where L̄ is given as (4.6) and L̃ := max{L̄, maxw∈[0,K+ ] maxi,j =1,2 |∂j fi (w)|}. We also de-
fine H + and H − by replacing F with F + and F − in (4.18), respectively. Since F ± (w1 , w2 )
are non-decreasing with respect to the second variable w2 for all w = (w1 , w2 ) ∈ [0, K+ ], it
is clear that H ± (·) are non-decreasing in C(R, [0, K+ ]). Furthermore, we define the operator
T = (T1 , T2 ) : C(R, [0, K+ ]) → C(R, R2 ) by
ξ +∞
1 λi1 (ξ −s) λi2 (ξ −s)
Ti (Ψ )(ξ ) = e Hi (Ψ )(s)ds + e Hi (Ψ )(s)ds (4.19)
c2 + 4di L̃ −∞ ξ
with
⎧ ⎧
⎨ c− c2 +4di L̃ ⎨ c+ c2 +4di L̃
λi1 = 2di , di > 0, and λi2 = 2di , di > 0,
⎩ ⎩
−L̃/c, d2 = 0, +∞, d2 = 0.
Similarly, we define T + and T − by replacing H with H + and H − in (4.19), respectively. Clearly,

T ± (Ψ ) are non-decreasing in C(R, [0, K+ ]), and

T − (Ψ ) ≤ T (Ψ ) ≤ T + (Ψ ), ∀Ψ ∈ C R, 0, K+ .
Step 2. For any c > c∗ , let Λ1 (c) be defined as in Lemma 4.2 and

Φ̃ + (ξ ) := min K1+ , eΛ1 (c)ξ , min K2+ , b(c)eΛ1 (c)ξ ,
where b(c) is defined as in Lemma 4.4. Since F + (w) ≤ DF(0)w for all w ∈ [0, K+ ], it is easy to
verify that T + (Φ̃ + )(ξ ) ≤ Φ̃ + (ξ ) for ξ ∈ R. Moreover, for a given λ ∈ (0, min{Λ1 (c), λ12 , λ22 }),
denote

Xλ := Ψ ∈ C R, R2 : supΨ (ξ )e−λξ < +∞
ξ ∈R
with Ψ λ = supξ ∈R Ψ (ξ )e−λξ . Then (Xλ , · λ ) is a Banach space. It is easy to see that
Φc− , Φ̃ + ∈ Xλ . Thus the set Y ⊂ Xλ defined by Y := {Ψ ∈ Xλ : Φc− ≤ Ψ ≤ Φ̃ + } is a convex and
closed subset of Xλ . In addition, for any Ψ ∈ Y, one can see that

Φc− = T − Φc− ≤ T − (Ψ ) ≤ T (Ψ ) ≤ T + (Ψ ) ≤ T + Φ̃ + ≤ Φ̃ + ,
which implies that T : Y → Y.

Step 3. We claim that T is compact on Y. We first show that T is continuous on Y. For any
Ψ1 = (φ1 , ψ1 ), Ψ2 = (φ2 , ψ2 ) ∈ Y, we have

H2 (Ψ1 )(ξ ) − H2 (Ψ2 )(ξ )e−λξ

≤ L̃ψ1 (ξ ) − ψ2 (ξ )e−λξ + ∂2 f2 (J2 ∗ φ1 )(ξ ), η1 (ξ ) ψ1 (ξ ) − ψ2 (ξ )e−λξ
+∞

+ ∂1 f2 η2 (ξ ), ψ2 (ξ ) J2 (y)φ1 (ξ − y − cτ2 ) − φ2 (ξ − y − cτ2 )e−λξ dy
−∞
≤ L̂1 Ψ1 − Ψ2 λ ,

+∞
where L̂1 := L̃(2 + −∞ J2 (y)e−λ(y+cτ2 ) dy).
η1 (ξ ) := θ1 ψ1 (ξ ) + (1 − θ1 )ψ2 (ξ ) and η2 (ξ ) := θ2 (J2 ∗ φ1 )(ξ ) + (1 − θ2 )(J2 ∗ φ2 )(ξ )
with θ1 , θ2 ∈ (0, 1). If d2 > 0, then we have
L̂1
T2 (Ψ1 )(ξ ) − T2 (Ψ2 )(ξ )e−λξ ≤ Ψ1 − Ψ2 λ ,
d2 (λ − λ21 )(λ22 − λ)
and if d2 = 0, there holds
ξ

T2 (Ψ1 )(ξ ) − T2 (Ψ2 )(ξ )e−λξ ≤ 1 e− c (ξ −s) e−λξ H2 (Ψ1 )(s) − H2 (Ψ2 )(s)ds
L̃
c
−∞
L̂1
≤ Ψ1 − Ψ2 λ .
cλ + L̃
Similarly, we can also obtain that

L̂2 Ψ1 − Ψ2 λ
T1 (Ψ1 )(ξ ) − T1 (Ψ2 )(ξ )e−λξ ≤ ,
d1 (λ − λ11 )(λ12 − λ)

+∞
where L̂2 := L̃(2 + −∞ J1 (y)e−λ(y+cτ1 ) dy). Therefore, we obtain
(L̂1 + L̂2 )Ψ1 − Ψ2 λ
T (Ψ1 ) − T (Ψ2 ) ≤ ,
λ
min{d1 (λ − λ11 )(λ12 − λ), d2 (λ − λ21 )(λ22 − λ), cλ + L̃}
which implies that T is continuous on Y.

Next, we show that T (Y) is compact in Xλ . For any Ψ ∈ Y and ξ ∈ R, direct computations
show that

Ti (Ψ )(ξ ) ≤ K + + 1 maxw∈[0,K+ ] fi (w),
i L̃
for i = 1, 2.
Ti (Ψ ) (ξ ) ≤ 2 L̃K + maxw∈[0,K+ ] fi (w) / c2 + 4di L̃,
+
1
Therefore, T (Y) is a family of uniformly bounded and equi-continuous functions on R. Thus,

using the method as in [13], we can show that T (Y) is compact in Xλ .
Therefore, Schauder’s fixed point theorem implies that the operator T has a fixed point Φc
in Y, which is a traveling wave solution of (1.7) for c > c∗ . Since
0 Φc− (ξ ) ≤ Φc (ξ ) ≤ Φ̃ + (ξ ), for ξ ∈ R,
then the assertions of (4.16) and (4.17) hold obviously.

Finally, when the condition (i) holds, we can choose F ± (·, ·) = F (·, ·) and K± = K. Hence
limξ →∞ Φc (ξ ) = K. In the case where (ii) holds, by using the upward convergence in the prop-
erty of spreading speeds, the proof of the limit limξ →∞ Φc (ξ ) = K is very similar to that of [13,
Theorem 2.3], see also [8, Theorem 4.1]. We omit it here. This completes the proof. 2
Next, using the limiting argument, we can establish the existence of traveling wave solution
Φ∗ with the minimal wave speed c∗ . In particular, under some additional assumptions, we can
further prove that Φ∗ (−∞) = 0. The results are stated as following.
Theorem 4.13. Assume (A1 ), (B1 ), (B2 ), (B3 ) and c = c∗ . For any vector σ
0 with σ 1,
(1.7) admits a non-constant traveling wave solution Φ∗ (ξ ) such that
0 ≤ Φ∗ (ξ ) ≤ K+ for ξ ∈ R, Φ∗ (ξ ) ≤ σ for ξ ≤ 0, and (4.20)

− +
K ≤ lim inf Φ∗ (ξ ) ≤ lim sup Φ∗ (ξ ) ≤ K . (4.21)
ξ →∞ ξ →∞
In addition, if α1 + β2 > 0 and α2 + β1 > 0, then Φ∗ (−∞) = 0. Note that αi = ∂i fi (0, 0) and
βi = ∂j fi (0, 0) for 1 ≤ i = j ≤ 2.
Moreover, if f1 (u, v) = −αu + h(v), f2 (u, v) = −βv + g(u), (A2 ), (A4 ), and one of the
following assumptions hold:

(ii) b(u)/u is strictly decreasing for u ∈ [K1− , K1+ ] and b(u) satisfies the property (P),
then limξ →∞ Φ∗ (ξ ) = K.
Proof. By using the limiting argument, we choose a sequence {cj } ⊂ (c∗ , +∞) such that
limj →∞ cj = c∗ . According to Theorem 4.12, there exists a traveling wave (Φj , cj ) of (1.7)
for each j such that
K− ≤ lim inf Φj (ξ ) ≤ lim sup Φj (ξ ) ≤ K+ .

ξ →∞ ξ →∞
Given any σ
0 with σ 1. Since Φj (ξ + h), h ∈ R, is also a solution satisfying
Φj (−∞) = 0, we can assume that Φj (ξ ) ≤ σ for ξ ≤ 0. Similar to the proof of Theorem 4.12,
we can prove that {Φj (ξ )}∞ j =1 is an equi-continuous and uniformly bounded sequence of func-
tions on R. Then, by Ascoli’s theorem and a nested subsequence argument, we obtain that there
exists a subsequence of {cj }, still denoted by {cj }, such that Φj (ξ ) converges uniformly on ev-
ery bounded interval, and hence pointwise on R to a function Φ∗ (ξ ) := (φ∗ (ξ ), ψ∗ (ξ )). Note that
Φj (ξ ) = T (Φj )(ξ ), ξ ∈ R, where
ξ +∞
1 λi1 (ξ −s) λi2 (ξ −s)
Ti (Φj )(ξ ) = e Hi (Φj )(s)ds + e Hi (Φj )(s)ds ,
c2 + 4di L̃ −∞ ξ
for i = 1, 2. Letting j → ∞ in the above equation and using the dominated convergence theorem,
we get Φ∗ (ξ ) = T (Φ∗ )(ξ ) for ξ ∈ R and the assertions of (4.20) and (4.21).
Next, we show that Φ∗ (−∞) = 0 when α1 + β2 > 0 and α2 + β1 > 0. We first show that
0 0
φ∗ (ξ )dξ < +∞ and ψ∗ (ξ )dξ < +∞. (4.22)
−∞ −∞
Note that Φ∗ (ξ ) ≤ σ for any ξ ≤ 0. Choose σ = (σ1 , σ2 )

0 with

1 α1 + β2 α2 + β1
σ ≤ min , ,
5M 4 4

+∞
where M := maxw∈[0,K+ ] max{∂ij F (w), i, j = 1, 2}. Since −∞ Ji (y)dy = 1, there exists a
constant T0 < 0 such that
T0 −cτ1 T0 −cτ2
σ2 σ1
J1 (y)dy < + and J2 (y)dy < .
K2 K1+
−∞ −∞
Hence, for ξ ≤ T0 , we have

+∞
(J2 ∗ φ∗ )(ξ ) = J2 (y)φ∗ (ξ − y − cτ2 )dy
−∞
−cτ2
ξ +∞
= J2 (y)φ∗ (ξ − y − cτ2 )dy + J2 (y)φ∗ (ξ − y − cτ2 )dy
−∞ ξ −cτ2
T0 −cτ2
≤ K1+ J2 (y)dy + σ1 ≤ 2σ1 .

−∞
Similarly, we can show that (J1 ∗ ψ∗ )(ξ ) ≤ 2σ2 for ξ ≤ T0 . Therefore, for ξ ≤ T0 ,

M φ∗2 (ξ ) + ψ∗2 (ξ ) + 2φ∗ (ξ )(J1 ∗ ψ∗ )(ξ )

+ 2(J2 ∗ φ∗ )(ξ )ψ∗ (ξ ) + (J1 ∗ ψ∗ )2 (ξ ) + (J2 ∗ φ∗ )2 (ξ )
α1 + β2 α2 + β1
≤ φ∗ (ξ ) + (J2 ∗ φ∗ )(ξ ) + ψ∗ (ξ ) + (J1 ∗ ψ∗ )(ξ ) .
4 4
By Taylor’s expansion, it then follows from (4.2) that
cφ∗ (ξ ) + cψ∗ (ξ ) − d1 φ∗ (ξ ) − d2 ψ∗ (ξ )

= f1 φ∗ (ξ ), (J1 ∗ ψ∗ )(ξ ) + f2 (J2 ∗ φ∗ )(ξ ), ψ∗ (ξ )
≥ α1 φ∗ (ξ ) + β1 (J1 ∗ ψ∗ )(ξ ) + α2 ψ∗ (ξ ) + β2 (J2 ∗ φ∗ )(ξ )

− M φ∗2 (ξ ) + 2φ∗ (ξ )(J1 ∗ ψ∗ )(ξ ) + (J1 ∗ ψ∗ )2 (ξ )

− M ψ∗2 (ξ ) + 2(J2 ∗ φ∗ )(ξ )ψ∗ (ξ ) + (J2 ∗ φ∗ )2 (ξ )
α1 + β2 β2 − α1
≥ φ∗ (ξ ) + (J2 ∗ φ∗ )(ξ ) + (J2 ∗ φ∗ )(ξ ) − φ∗ (ξ )
4 2
α2 + β1 β1 − α2
+ ψ∗ (ξ ) + (J1 ∗ ψ∗ )(ξ ) + (J1 ∗ ψ∗ )(ξ ) − ψ∗ (ξ ) (4.23)
4 2
for ξ ≤ T0 . Integrating (4.23) over [y, T0 ] with y < T0 , we can obtain

c φ∗ (T0 ) − φ∗ (y) + c ψ∗ (T0 ) − ψ∗ (y) − d1 φ∗ (T0 ) − φ∗ (y) − d2 ψ∗ (T0 ) − ψ∗ (y)
T0 T0
β2 − α1 β1 − α2
− (J2 ∗ φ∗ )(ξ ) − φ∗ (ξ ) dξ − (J1 ∗ ψ∗ )(ξ ) − ψ∗ (ξ ) dξ
2 2
y y
T0 T0
α1 + β2 α2 + β1
≥ φ∗ (ξ ) + (J2 ∗ φ∗ )(ξ ) dξ + ψ∗ (ξ ) + (J1 ∗ ψ∗ )(ξ ) dξ. (4.24)
4 4
y y
It is easy to see that Φ∗ (ξ ) is uniformly bounded in R. Moreover, for any y ≤ T0 , we have
T0

(J2 ∗ φ∗ )(ξ ) − φ∗ (ξ ) dξ

y
T0 +∞ 1

= (x + cτ2 )J2 (x) φ∗ ξ − θ (x + cτ2 ) dθ dxdξ

y −∞ 0
+∞ 1

= (x + cτ2 )J2 (x) φ∗ T0 − θ (x + cτ2 ) − φ∗ y − θ (x + cτ2 ) dθ dx

−∞ 0
+∞
≤ 2K1+ cτ2 + |x|J2 (x)dx < +∞
−∞
and
T0 +∞

+
ψ∗ (s − cτ1 ) − ψ∗ (s) ds ≤ 2K2 cτ1 + |x|J1 (x)dx < +∞.

y −∞

T0
T
Thus y [(J1 ∗ φ∗ )(ξ ) − φ∗ (ξ )]dξ and y 0 [(J2 ∗ ψ∗ )(ξ ) − ψ∗ (ξ )]dξ are bounded on (−∞, T0 ].

T
T
Therefore, from (4.24), we see that y 0 φ∗ (s)ds and y 0 ψ∗ (s)ds are bounded on (−∞, T0 ].
Hence (4.22) follows.
Now, we are able to show that Φ∗ (−∞) = (φ∗ (−∞), ψ∗ (−∞)) = 0. Suppose for the contrary
that for some 0 > 0, there exists a sequence ξn → −∞ such that φ∗ (ξn ) > 0 . By the uniform
boundedness of φ∗ , we see that φ∗ (ξ ) is uniformly continuous on R. Then, there exists a δ > 0
(independent of ξn ) such that
0
φ∗ (ξn ) > , ∀ξ ∈ (ξn − δ, ξn + δ).
2
Without loss of generality, we may assume that (ξm − δ, ξm + δ) ∩ (ξk − δ, ξk + δ) = ∅, ∀m = k.

Then,
0 ∞ ξ n +δ
∞
0
φ∗ (s)ds ≥ φ∗ (s)ds ≥ 2δ = ∞,
2
−∞ n=1 ξ −δ n=1
n
which leads to a contradiction. Hence, φ∗ (−∞) = 0. Similarly, we can show that ψ∗ (−∞) = 0.
Therefore, Φ∗ (−∞) = 0.
The proof of the upward convergence Φ∗ (+∞) = K is similar to that of Theorem 4.12 and
omitted. This completes the proof. 2
4.4. The effects of delay and nonlocality on the spreading speed
In this subsection, we shall consider the effects of the delays and nonlocality in (1.7) on
the spreading speed c∗ . In the past, the similar problem for scalar delayed nonlocal reaction–
diffusion equations have been considered in the works Li et al. [17], Zou [44] and Wang
et al. [29]. In [34], the authors also studied the problem for a special epidemic model. Here
we remark that system (1.7) is more general than those of the mentioned works.
From Theorems 4.8, 4.10, 4.12 and 4.13, we know that the c∗ defined by Lemma 4.2 is the
minimal wave speed and coincides with the spreading speed. First, we consider the effect of the
delays on the spreading speed. For simplicity, let’s take Ji (y) = δ(y). Then

1 (c, λ) := d1 λ2 − cλ + α1 d2 λ2 − cλ + α2 − β1 β2 e−cλ(τ1 +τ2 ) .
By 1 (c∗ , λ∗ ) = 0 and ∂λ ∂
1 (c∗ , λ)|λ=λ∗ = 0, it is easy to see that c∗ = c∗ (τ1 , τ2 ) and λ∗ =
λ∗ (τ1 , τ2 ) are differentiable functions with respect to τ1 , τ2 . Since di λ2∗ − c∗ λ∗ + αi < 0, i = 1, 2,
direct computations show that
∂c∗ (τ1 , τ2 ) β1 β2 ce−c∗ λ∗ (τ1 +τ2 )

= <0
∂τi (d1 + d2 )λ2∗ − 2c∗ λ∗ + α1 + α2 − β1 β2 (τ1 + τ2 )e−c∗ λ∗ (τ1 +τ2 )
for i = 1, 2. Thus, the delays τ1 and τ2 will slow the spreading speed.
Next, we consider the effect of the nonlocality on the spreading speed. Fixing τ1 and τ2 and
take
2 2
1 −x 1 −x
J1 (x) = √ e 41 and J2 (x) = √ e 42 .
4π1 4π2
Obviously, the larger 1 and 2 , the stronger the effect of nonlocality; vice versa. By direct
computations. We have

1 (c, λ) := d1 λ2 − cλ + α1 d2 λ2 − cλ + α2 − β1 β2 e(1 +2 )λ −cλ(τ1 +τ2 ) .
2
Similarly, c∗ = c∗ (1 , 2 ) and λ∗ = λ∗ (1 , 2 ) are differentiable functions with respect to 1 ,

2 , and
−β1 β2 λ∗ e(1 +2 )λ∗ −c∗ λ∗ (τ1 +τ2 )

2
∂c∗ (1 , 2 )
= > 0,
(d1 + d2 )λ2∗ − 2c∗ λ∗ + α1 + α2 − β1 β2 (τ1 + τ2 )e(1 +2 )λ∗ −c∗ λ∗ (τ1 +τ2 )
2
∂i
i = 1, 2. Hence, the nonlocality will increase the spreading speed.
4.5. An example
Consider the following delayed epidemic model:

ut = d1 uxx − u(x, t) + γ (J1 ∗ v)(x, t),
(4.25)
vt = d2 vxx − βv(x, t) + g (J2 ∗ u)(x, t) ,
where d1 , β, γ > 0, d2 , τ1 , τ2 ≥ 0, (J1 ∗ v)(x, t) and (J2 ∗ u)(x, t) are defined as (1.6).
The spreading speeds and traveling wave solutions for some special cases of (4.25) when g
is increasing has been extensively studied, see e.g. [25,37,43]. For example, in the case where
d2 = τ1 = τ2 = 0 and J2 (·) = J1 (·) = δ(·), Xu and Zhao [37] proved the existence, unique-
ness and stability of bistable traveling wave fronts of (4.25). Zhao and Wang [43] also estab-
lished the existence and non-existence of monostable traveling wave fronts. For the case where
d2 = τ1 = τ2 = 0 and J2 (·) = δ(·), Xu and Zhao [38] considered the spreading speed and monos-
table traveling wave fronts. When d2 = τ1 = 0 and J2 (·) = J1 (·) = δ(·), Thieme and Zhao [25]
obtained the existence of spreading speed and minimal wave speed by applying their theory de-
veloped for integral equations. These results on traveling wave solutions were then extended
by Wu and Liu [35] to the non-quasi-monotone case by transforming the system into an inte-
gral equation and constructing two auxiliary monotone integral equations. However, for the case
where d1 , d2 > 0, (4.25) can not be transform into an integral equation. In fact, to the best of
our knowledge, there has been no results on the traveling wave solutions and spreading speed for
(4.25) in the case where d1 , d2 > 0 and g is non-monotone.
We assume that J1 (·) and J2 (·) satisfy (A1 ) and the following conditions hold:
(H1 ) g ∈ C 2 ([0, +∞), [0, +∞)), g(0) = g(k) − βγ −1 k = 0, g(u) > βγ −1 u for u ∈ (0, k), and
g(u) ≤ g (0)u for u ∈ [0, k], where k > 0 is a constant.
(H2 ) One of the following holds:
(a) g(u) is increasing for u > 0;
(b) there exists a number u∗ > 0 such that g(u) is increasing for 0 < u ≤ u∗ and decreasing
for u > u∗ .
Let’s denote K := (k, β −1 g(k)) and F (u, v) := (f1 (u, v), f2 (u, v)) = (−u + γ v, −βv + g(u)).
Clearly, F (0) = F (K) = 0 and

−1 γ
DF(0) = .
g (0) −β
From the assumption (H1 ), we see that
F (w) ≤ DF(0)w for w ∈ [0, K] and g (0) > βγ −1 > 0,
i.e. ∂1 f1 (0, 0)∂2 f2 (0, 0) < ∂2 f1 (0, 0)∂1 f2 (0, 0). Hence, (B1 ) and (B2 ) hold for (4.25).
In addition, if (H1 ) and (H2 )(a) or (H2 )(b) hold with k ≤ u∗ , it is easy to see that (B3 ) holds
for (4.25). We also claim that (B3 ) holds for system (4.25) provided that (H1 ) and (H2 )(b) hold
with k > u∗ . In fact, let’s set

umin := inf u ∈ (0, u∗ ] | g(u) = g γβ −1 g(u∗ ) .
It’s obvious that umin > 0. Similar to [36], we can define two functions F ± (u, v) as follows:

F ± (u, v) = −u + γ v, −βv + g ± (u) ,
where

+ g(u), u ∈ [0, u∗ ], − g(u), u ∈ [0, umin ],
g (u) = and g (u) =
g(u∗ ), u ∈ [u∗ , γβ −1 g(u∗ )] g(umin ), u ∈ [umin , γβ −1 g(u∗ )].
Clearly, g + (u) ≤ g (0)u for u ∈ [0, γβ −1 g(u∗ )]. Hence, F + (w) ≤ DF(0)w for w ∈ [0, K+ ].
Thus, one can easily check the condition (B3 ) with K = (k, β −1 g(k)),

K+ = k1+ , k2+ = γβ −1 g(u∗ ), g(u∗ ) and K− = k1− , k2− = γβ −1 g(umin ), g(umin ) .
Moreover, it’s clear that the characteristic function for (4.25) with respect to the trivial equi-
librium 0 can be represented by

2 (c, λ) := d1 λ2 − cλ − 1 d2 λ2 − cλ − β
+∞ +∞
−cλ(τ1 +τ2 ) −λy
− g (0)γ e J1 (y)e dy J2 (y)e−λy dy
−∞ −∞
for c ≥ 0 and λ ∈ C. By Lemma 4.2, there exists a unique (c∗ , λ∗ ) satisfies the following system:
∂
2 (c, λ) = 0 and 2 (c, λ) = 0.
∂λ
Therefore, following the previous theorems, we have the following results.
Theorem 4.14. Assume (A1 ) and (H1 ). The following statements hold:
(1) If (H2 )(a) or (H2 )(b) holds with k ≤ u∗ , then the conclusions of Theorems 4.3 and 4.5 hold
for (4.25);
(2) If (H2 )(b) holds with k > u∗ , then the conclusions of Theorems 4.8, 4.10, 4.12 and 4.13 hold
for (4.25). Additionally, if b(u)/u is strictly decreasing for u ∈ [k1− , k1+ ] and b(u) satisfies
the property (P), where b(u) = γβ −1 g(u), then the upward convergence of the spreading
speed and traveling wave solutions holds.
In particular, the spreading speed c∗ for each of the above two cases is linearly determinate
and coincides with the minimal wave speed of traveling wave solutions of (4.25).
Acknowledgment
The authors are grateful to the referee for his/her valuable comments and suggestions which
have led to an improvement of the presentation.
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Global attractivity, spreading speeds and traveling

Received 22 December 2013; revised 17 September 2014

MSC: 35K57; 35R10; 35B40

where d1 > 0, d2 ≥ 0; the convolution terms in (1.5) are defined by

(A2 ) g, h ∈ C 2 (R+ , R), h(0) = g(0) = 0,

(B1 ) there exists a K := (K1 , K2 )

2. Solutions of Cauchy problem and comparison theorems

(A3 ) g(·) is increasing on [0, +∞) and limu→+∞ g(u) := g∞ > 0;

(4) As usual, we identify an element φ ∈ C as a function from R × [−τ, 0] into R2 defined by

(2) Let’s define F = (F1 , F2 ) : C → X by

2.1. Positivity and boundedness of solutions

which implies that

Similarly, we can show that

2.2. Comparison theorems

respectively. If w+ (x, s) ≥ w− (x, s) ≥ 0 for x ∈ R and s ∈ [a − τ, a], then w+ (x, t) ≥

for (x, t) ∈ R × (0, +∞).

Then we establish the following comparison theorem.

w − (x, s) ≤ ϕ(x, s) ≤ w + (x, s) for (x, s) ∈ R × [−τ, 0],

w − (x, t) ≤ w(x, t; ϕ) ≤ w + (x, t), for all x ∈ R and t > 0.

which implies that

Zi (x, t) ≤ Ti (t)Zi (x, 0) ≤ 0, for (x, t) ∈ R × (0, τm ], i = 1, 2.

(Z2 )t ≤ d2 (Z2 )xx − βZ2 , for (x, t) ∈ R × (0, τ2 ].

(Z1 )t ≤ d1 (Z1 )xx − αZ1 for (x, t) ∈ R × (0, τ2 ].

for (x, t) ∈ R × (0, +∞), where

where Γ1 is defined in (2.3). Hence, for (x, t) ∈ R × (0, +∞),

Note that 0 ≤ (Z̃1,λ , Z̃2,λ ) ≤ K̄. Then, we have

(Z2 )t ≤ d2 (Z2 )xx − βZ2 , for (x, t) ∈ R × (0, +∞).

3. Global attractivity of equilibria

with initial value

w r = ϕ = (ϕ1 , ϕ2 ) ∈ Cτ+ . (3.2)

Then the following statements hold:

The assertions are proved in the sequel.

lim w(t, 0̂) = c and lim w(t, b̂) = d,

w(t, 0̂) ≤ w(t, ϕ) ≤ w(t, b̂), for t > 0,

(see e.g. [23, Theorem 5.1.1]). Therefore, limt→∞ w(t, ϕ) = 0.

lim w(t, â) = c and lim w(t, b̂) = d,

where a ≤ c ≤ d ≤ b and f (ĉ) = f (d̂) = 0. Since a

w(t, â) ≤ w(t, ϕ) ≤ w(t, b̂), for t > 0,

we conclude that limt→∞ w(t, ϕ) − K = 0.

Then, there exists a T1 > τ such that

Let w̄(t) and w(t) be solutions of the following problems:

respectively. Then it follows from Lemma 2.4 that

u(t) ≤ u(x, t; ϕ) ≤ ū(t), for x ∈ R and t ≥ τ. (3.3)

Proof. We first construct an upper auxiliary system:

with initial value

Therefore, limt→∞ w(·, t; ϕ) = 0. This completes the proof. 2

To study the global attractivity of K, we first introduce the function:

Fig. 2. The graphs of g and h for the case K1 ≤ u∗ .

It then follows from Lemma 2.6 that

w(t) ≤ w(x, t; ϕ) ≤ w̄(t), for x ∈ R and t > T + τ.

Since g(u) ≤ g(u∗ ) for all u ≥ 0, it is easy to show that

Note that when K1 = u∗ ,

h−1 (αK1 ) = β −1 g(K1 ) = v− ≤ h−1 (αu− ).

h−1 (αu− ) ≥ v− = β −1 g(ũ− ) > h−1 (α ũ− ).

αu+ = h(v+ ) and βv+ = g(u+ ); or h−1 (αu+ ) = β −1 g(u+ ).

3.2. (ii) K1 > u∗

then, for any δ = (δ1 , δ2 ) > 0 and M > 0, we have

αi = ∂i fi (0, 0) < 0 and βi = ∂j fi (0, 0) > 0 for 1 ≤ i = j ≤ 2.

1 (c, λ) := P1 (c, λ) − P2 (c, λ), for c ≥ 0 and λ ∈ C, (4.1)

(B3 ) There exist K± = (K1± , K2± )

(i) (A3 ) holds and K1 ≤ u∗ ;