You are on page 1of 8

Advances in Water Resources 28 (2005) 956963

www.elsevier.com/locate/advwatres

The inuence of soil properties on the formation of


unstable vegetation patterns on hillsides of semiarid catchments
Nadia Ursino

Universita di Padova, Dept. IMAGe, via Loredan 20, I-35131 Padova, Italy
Received 17 June 2004; received in revised form 14 February 2005; accepted 22 February 2005
Available online 10 May 2005

Abstract
On hillsides of semiarid catchments regular bands of vegetation have been observed to form under low rainfall conditions. Many
authors have observed the existence of a slope gradient threshold below which no banded patterns were observed; this increases with
the mean annual rainfall. A simple model for soil moisture balance and vegetation growth, explicitly accounting for basic soil physics, is demonstrated and discussed. The inuence of relevant soil characteristics on vegetation patterns and their patchiness is
addressed by linear stability analysis. The results conrm the experimental evidence of a threshold slope gradient depending on
the mean annual net water supply, and demonstrate the inuence of the soil properties (saturated conductivity and capillary rise)
on the stability condition and on the threshold slope. Nevertheless, the concluding remark concerns the oversimplied models for
vegetation patterns, such as the one discussed here, that require properly dened eective soil parameters in order to become predictive tools.
 2005 Elsevier Ltd. All rights reserved.
Keywords: Ecohydrology; Semiarid catchments; Vegetation banding; Stability analysis; Patterns initiation

1. Introduction
Regular bands of vegetation have been observed to
form on hillsides of semiarid catchments under low rainfall conditions. Numerous experimental studies have
been conducted in dierent parts of the world, mainly
in Africa and in Australia, in order to collect data and
classify patterns of vegetation organized in a two-phase
mosaic as well as to dene their connection with geomorphic and climatic conditions. Valentin et al. [24]
present a summary of the experimental evidence collected in the last 30 years circa. There is a continuous
interest in trying to understand and model vegetation
patterns formation and evolution. Lefever and Lejeune
[8] explain the formation of tiger bush through a mechanism of resources competition. Gilad et al. [5] attribute
*

Fax: +39 49 8275446.


E-mail address: nadia@idra.unipd.it

0309-1708/$ - see front matter  2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.advwatres.2005.02.009

the appearance of mosaic vegetation to the cooperation


of plant and microorganism. Ares et al. [1] recognize
that grazing and thus anthropic use may modify the spatial patterns of vegetation. The formation of vegetation
bands alternating with bare soil has been linked by
many authors to a low scale process of redistribution
of water by runo [22,8,9,7,24,3,6,25,14,2]. The redistribution of surface water is considered as an important
factor explaining the display of mosaic vegetation patterns, according to the following self-organizing mechanism. Rain falling on bare patches of soil fails to
inltrate and runs o [19]. This run-o water accumulates in the vegetated patches, where it can inltrate
more easily; the spatial vegetation distribution then
inuences the spatial distribution of soil particles, organic matter and nutrients [14]. HilleRisLambers et al.
[6] demonstrate the eect of bare soil inltration
capacity on patterns formation. Rietkerk et al. [15] recently summarized model results and perspectives on

N. Ursino / Advances in Water Resources 28 (2005) 956963

echosystems bistability and catastrophic shifts that may


be linked to vegetation patchiness. Despite the many diverse recent contributions to the comprehension of the
genesis and evolution of vegetation mosaic, there remain
questions concerning the origin of these vegetation and
soil moisture paths that still remain unanswered, such
as: do the averaged soil properties have an impact on
vegetation patchiness? And, eventually, to what extent?
Notwithstanding the complexity of the interplay between vegetation, soil, moisture and nutrients, a simple
model for soil moisture and plant biomass [7] shows that
local interactions (such as the feedback between water
and biomass), coupled with dispersion, can cause regular vegetation patterns to develop in the absence of soil
heterogeneity as the result of Turing-like instability
[23,10]. The results previously obtained with this model
demonstrate that the emergent behavior of large, complex landscapes in a prolonged period of disequilibrium
may simplify into a form that can be observed without
an explicit consideration of soil eective parameters. In
other words, although soil may have a crucial role in
water and nutrient transport and thus in vegetation
growth, soil properties are usually not explicitly considered within this model. The following questions arise:
how may relevant soil properties be taken into account
in hydro-biological models? Are the parameters and
concepts commonly adopted in soil physics adequate
to describe the role that the soils play in creating vegetation patterns?
The model of Klausmeier [7] (revised in Section 2.1),
is reinterpreted here in order to account for relevant soil
properties. The new modied version of this model is
demonstrated in Section 2.2. Factors such as surface
water, nutrient transport as well as animals and microorganism action are neglected, in the belief that the impact of soil properties on the stability condition could be
captured anyway. Many authors have observed that a
slope gradient threshold, below which no banded patterns were observed, exists and that it tends to increase
with mean annual rainfall (e.g. [26,21]). The new model
allows derivation of the marginal stability condition as a
function of the hill slope, the saturated conductivity,
and the capillary rise. The results presented here were
obtained analytically and confronted with experimental
data taken from literature (Section 3). In Section 4, the
dierent processes that appear to have an inuence on
the interaction between water, soil and vegetation are
discussed. It has been demonstrated that the relationship
between critical annual rainfall and critical slope is
strongly aected by the averaged over depth saturated
conductivity and by the mean capillary rise. Nevertheless, the parameterization of the model results is particularly delicate. It may also be said that properly dened
eective parameters (which may resemble the subscale
soil processes that determine the macroscopic evolution
of the unstable ecosystem) are needed.

957

2. Theory
Turing [23] provided a hypothesis to explain the generation of patterns through mechanisms of reaction and
diusion. By superimposing an initial pattern of given
wave-number upon the steady state homogeneous solution and allowing reaction and diusion to act over
time, the initial disturbance is expected to either grow
or decline. Obviously, the patterns which grow faster
in time will be most visible. The mathematical framework for this stability analysis is the Fourier analysis.
The relations between the attributes of the unstable patterns (growth rate and characteristic length) and those
of the environment (mean rainfall, hill slope and soil
properties) are investigated below.
The characteristic length of the bands of vegetation
considered here is much larger in the direction perpendicular to the slope than in the parallel one. The soil
moisture follows a spatial distribution similar to that
of the vegetation. Furthermore, the water content variability in the direction perpendicular to the soil surface
may be neglected since the soil depth that inuences vegetation growth is shallow as compared to the surface
extension of the domain (a similar hypothesis is commonly adopted in remote sensing studies, see e.g. [20]).
Thus, based on the above simplied hypothesis, the
theoretical analysis of the soil moisture and of the vegetation balance equation is limited here to the one-dimensional case only. On at ground the extension of the
model and the related results to the two-dimensional
case is straightforward, by replacing the squared mode
considered here with the summation of the squared
unstable modes in two directions.
2.1. Linear stability analysis of soil moisture
and biomass balances
The model of Klausmeier [7] is a pair of partial dierential equations for soil moisture (W) and plant biomass
(N) that are dened on an innite one dimensional domain indexed by X as a function of time T.
oW
oW
A  LW  RWN 2 V
oT
oX
2
oN
o
N
RJWN 2  MN D 2
oT
oX

where A is the uniform water supply (according to


Klausmeier [7] A does not account for the feedback between inltration and biomass [6] even if this mechanism
has been proved to induce instability); LW is the water
loss due to evaporation and leakage (this may be
approximately considered to be a linear function of
the soil moisture under stress conditions, e.g. [16,18]);
MN is the plant biomass lost due to mortality; V is
the downhill water speed; J is the yield of plant biomass
per unit water consumed; D is the diusion coecient of

958

N. Ursino / Advances in Water Resources 28 (2005) 956963

plant dispersal, and RWN2 is the plants uptake. According to Klausmeier [7] plants take up water at rate
RG(W)F(N)Nwhere G(W) is the response of plants to
water and F(N) is a function that describes how plants
increase inltration. Klausmeier [7] linearizes the problem assuming G(W) = W and F(N) = N, and arguing
that the results are not sensitive to the exact form of
these functions. The lateral loss V oW
, that is responsible
oX
for the instability of the model, will be disclosed in the
next section.
Equation (1) can be reduced in dimensionless form
assuming T = L1t, X = D1/2L1/2x, V = (LD)1/2v, M =
Lm, W = Wsw, N = WsJn and A = LWsa, and obtaining:
ow
ow
a  w  rwn2 v
ot
ox
2
on
o
n
rwn2  mn 2
ot
ox

where r W 2s J 2 RL1 .
The spatially uniform steady state solution (w0, n0) of
the coupled soil moisture and vegetation problem (1) is
the solution of
lw; n a  w  rwn2 0

3
hw; n rwn2  mn 0
p
1
namely: w0 0.5a  a2  4m2 r1 and n0 mw1
0 r .
Adding a small perturbation to the steady state
(w0, n0),


w  w0
u
4
n  n0
in the form
X
ck ekt U k x;
u

where the growth rate k is the eigenvalue, the constants


ck are determined by a Fourier expansion of the perturbation in terms of Uk(x), and k is the wave number.
Substituting (4) into (2) and linearizing yields to the following relation
ut Au V ru  Dr2 u;
where

lw
A
hw






hn
v 0
0 0
;V
;D
0 0
0 1
ln

and lw, hw, hn and ln are the derivative of l and h with respect to w or n, evaluated in (w0, n0). Substituting (5) into
(6), the following expression for the eigenvalue k may be
found:
k A jkV  k 2 D;

where j is the imaginary unit. The steady state (w0, n0) is


stable in the absence of any spatial eect (when k = 0)
and unstable for some k50. The rst condition gives

jAj > 0 and lw + hn < 0. The second condition is fullled


when the real part of k is negative [10].
2.2. A linear stability model for subsurface soil
moisture dynamics on slopes accounting for eective
soil parameters
The model of Klausmeier [7] includes within the
water balance equation the term V oW
, that accounts
oX
for the ow parallel to the soil surface and induces instability [10]. It may be interpreted as the ow of soil moisture within the root depth and may be expressed as a
function of the slope gradient and of the soil properties
as will be shown below. Further, by analyzing the subsurface ow by the continuum approach, an additional
diusive term appears. The diusive term neglected by
Klausmeier [7] is found in other instability models (e.g.
[14]) and will also be demonstrated to lead to instability.
According to the Richards equation [13] that governs
the vadose-zone water ow, the net variation of soil
moisture due to the horizontal soil water ux q (x-direction) may be expressed as follows



dq
d
dW


Kh
i
8
dx dx
dx
where W 6 0 is the negative pressure head in the unsaturated soil, h is the soil moisture content, K(h) is a mean
value of the soil moisture dependent conductivity in the
dz
unsaturated root zone i  dx
is the slope gradient (the
vertical coordinate z is taken as positive upward). Set
h = hs eaW and K = Ks eaW = h Ks/hs [4], where a is
the inverse of the mean capillary rise and depends on
the soil characteristics [11,12]. a represents the ratio between the gravity and the capillary forces. Substituting
K = K(W) and h = h(W) into (8) yields to


dq
d2 h
dh
Dw 2 V w ;
dx
dx
dx

where Dw = Ks/(hs a) and Vw = i Ks/hs.


By averaging q and h we obtain the horizontal
ux Q
RH
through the root zone depth H : Q 0 q dz, and the
RH
averaged soil moisture content W 0 h dz the upper
limit of which Ws is the extractable soil moisture and
may be estimated from the dierence between the wettest and the dryest prole in a time series of prole soil
moisture data. Ws depends on the soil type and on the
vegetation type since deeper rooted vegetation has access to a larger depth H. Here the extractable soil moisture is set approximately Ws = hsH. Taking into account
the vertical ux of water through a plane element, due to
the net rainfall supply, evapotranspiration and leakage,
we nd an expression similar to the rst of Eq. (1),
2
where Vw = V, and the additional term Dw ddxW2 appears
on the right side accounting for soil moisture diusion
due to tension gradients.

N. Ursino / Advances in Water Resources 28 (2005) 956963

The destabilizing advective term that appears in the


model of [7] Eq. (2) has been demonstrated to be the
component of the subsurface ow in the direction parallel to the hill slope. It depends on the nature of the lower
boundary of the root zone which is investigated here. An
impermeable boundary deviates the whole inltrating
soil moisture, whereas, if the rain inltrates into a sloped
soil of innite extension, the subsurface ow parallel to
the slope reduces to the projection of the vertical ux in
the direction of the slope. In order to properly account
for the transverse loss of moisture, the leakage must
be carefully estimated and expressed as a function of
the soil properties.
The dimensionless form of the new pair of water and
vegetation balance equations is the following:
ow
ow
o2 w
a  w  rwn2 v
d 2
ot
ox
ox
2
on
on
rwn2  mn 2
ot
ox
1

10

where d = DwD . In the new water balance equation


(the rst of (10)) both dimensionless numbers v and d
are linked to the soil properties Ks and a, since
1=2
d = Ks(hs a)1D1 and v i
K s h1
. When the
s LD
capillary forces are negligible and a becomes very
large, d tends to 0 and the proposed model (10) formally reduces to the model of Klausmeier [7] (2). The
diusive term allows instability to appear also on at
ground.
The eigenvalues k as a function of the wave number k
are still given by (6), where now


d 0
D
;
11
0 1

959

whereas A and V (Eq. (3)) and the steady state solution


(w0, n0) remain unchanged.
The critical stability condition upon the real part of
the growth rate Rek = 0, identies the boundary between stable and unstable conditions. It may be expressed in terms of the correspondence between two or
more soil and/or vegetation parameters, such as i and
A in the cases presented here. It allows the evaluation
of the critical wave number k that has maximum growth
rate Rek = 0, and is derived below (according to Rovinsky and Menzinger [17]).
i1=2
1=2
1 h
Rek lw hn  d 1k 2 p q2 p2
q
2
12
where Rek is the real part of the growth rate k,


p 2kv lw  hn  d  1k 2 ;
and

2
q lw  hn  d  1k 2 4ln hw  k 2 v2 .
Based on (3): lw 1 rn20 , ln = 2m, hw rn20 and
hn = m.
Fig. 1 illustrates the signicance of the critical stability condition Rek = 0. The linear stability analysis demonstrated in this section distinguishes between stable
and unstable scenarios on the base of the growth rate
of a very small initial disturbance, but says nothing
about the resulting nite size pattern. Fig. 1 shows the
growth rate curves (12) at the critical stability condition
(A = 374 mm, and i = 0.02), below (A > 374 mm, and
i < 0.02), and beyond (A < 374 mm, and i > 0.02). Below
the critical stability condition all the perturbation modes
are stable since Rek is always negative. Beyond, a

Fig. 1. Growth coecient Rek as a function of wavelength k. Depending on A and i no wavelength may lead to instability (Rek < 0), or a continuous
range of unstable modes may do this. At the critical stability condition only one k has Rek = 0; the other modes are stable. In the case shown here, the
parameters that characterize the marginal stability condition are A = 374 mm, and i = 0.02. The other relevant parameters are M = 1.8 y1,
J = 0.003 kg m2 mm1 y1; R = 100 m4 kg2 y1, D = 1 m2 y1, L = 4 y1, hs = 0.4, H = 0.1 m, Ks = 3000 m y1 and a = 100 m1.

960

N. Ursino / Advances in Water Resources 28 (2005) 956963

continuous range of unstable modes may arise if stimulated. In the next section, the focus is on the pair Ai
leading to the marginal stability condition for dierent
biomass and soil parameters.

3. Results
A sensitivity analysis of the critical stability condition
(12) for soil parameters was developed. All variables
were set according to literature data or reasonable
assumptions.
Rainfall in semi-arid regions is between 50 and 750 mm
y1 [24]. A is set in this range. Klausmeier [7] assumes for
grass M = 1.8 y1; J = 0.003 kg m2 mm1 y1; R =
100 m4 kg2 y1 (obtained from the expression for equilibrium plant biomass); D = 1 m2 y1, and L = 4 y1.
Thus, m = 0.45 and r = 0.36. The maximum stored soil
moisture Ws hsH depends on the active soil depth H.
If hs = 0.4, and H = 0.1 m, then Ws = 40 mm. In the depth
averaged water balance only the transverse conductivity
comes into play. It was set Ks = 3 102 and 3
103 m y1 (for more clayey and more sandy soils respectively). Rather low wettability of the bare zones and lateral diusion due to capillary forces are expected to
support the so called runo runon mechanism of water
redistribution, which is responsible for the formation of
vegetation patterns. Thus, very high values of the ratio between gravity and capillary forces a have been considered:
a = 10, 100, 1000 m1. Klausmeier [7] does not take into
account soil properties and assumes v = 182.5 and d = 0.
Here, for i = 4%, L = 4 y1, and a = 100 m1,
1=2
v iK s h1
16, 160 and d = Ks(hsaD)1 = 7.5,
s LD
75 depending on Ks (case 1). Setting L = 2 y1, v = 22,
220 and d = 11, 110 (case 2). Assuming that a = 1000,

which indicates negligible capillary forces acting on soil


moisture, the proposed model reproduces the results previously obtained by Klausmeier [7] neglecting soil moisture diusion.
The critical slope i is plotted in Fig. 2 as a function of
the net precipitation A for dierent Ks and a. It is worthwhile repeating here that on sites characterized by a hillslopes below the critical i no banded vegetation patterns
are expected to appear, and that the threshold gradient
slope has been observed to increase with the mean rainfall intensity. Fig. 2 demonstrates that the results of the
proposed model are in agreement with this experimental
evidence. Furthermore, the range of precipitation that
induces the formation of banded vegetation extends towards the higher A with increasing saturated conductivity Ks (curves on the left side of Fig. 2 versus
corresponding curves on the right side). This result
could be expected since Ks inuences both: the advective
and the diusive terms in the water balance Eq. (10). Indeed advection and diusion contribute toward the
reduction of the already scarce soil moisture.
The inuence of the newly introduced diusive term
may be inferred confronting case a = 1000 m1 (circles)
with the other two: a = 10 and 100 m1 (squares and triangles, respectively). For gentle hill slopes, soils characterized by lower a values reach the critical stability
condition at higher A (e.g. triangles and squares for
i < 0.04). Conversely, for steeper slopes, the critical stability condition is reached at higher net precipitation by
the soils characterized by larger a. It may otherwise be
stated that for very low A the critical stability is reached
at gentle slope i when a is low; conversely, for higher A
the soil with larger a shows an unstable behavior at steep
slopes. Fig. 2 demonstrates that the lower a values
(a = 10) are associated with unstable behavior also for

Fig. 2. Case 1. Grass critical stability condition for dierent slope gradients i and net inow A. ( ) Experimental data taken from literature. Open
symbols: analytical solutions of the marginal stability condition problem Rek = 0 (12). Water loss due to evaporation and leakage per unit soil
moisture L = 4 y1 and coecient of plant uptake R = 100 m4 kg2 year 1. Soil parameters: saturated conductivity Ks = 3000 m y1 (left) and
Ks = 300 m y1 (right); a = 1000 m1 (n); a = 100 m1 ( ) a = 10 m1 (h).

N. Ursino / Advances in Water Resources 28 (2005) 956963

i = 0 in the lower range of precipitation (between a minimum Amin and the intersection of the marginal stability
condition with the axis i = 0). In this case, instability is
impelled by capillary diusion only (d is maximum when
a is minimum). In general, the inuence of Ks, a and i on
the critical stability condition is more evident for large
Ks (left side of Fig. 2).
In Fig. 2 the analytical model results are plotted with
the critical stability conditions observed by dierent
authors at dierent sites in Africa and Australia (bold
circles in Fig. 2). All the experimental data are referred
by Valentin et al. [24] in a review paper. Unfortunately
the sparse details concerning the soil characteristics reported in the literature references do not allow us to
accurately parameterize our model. At any rate, the
hypothesis of homogeneous soil adopted here for the
sake of simplicity sounds rather unrealistic, and thus it
does not justify a very accurate description of the subsoil, other than in terms of homogeneous, eective
parameters. The eective parameters are meant here to
be those values that somehow match the experimental
evidence and thus make the model soil behavior similar
to the observed one. Indeed, only the processes in action
and the relevant parameters are discussed, whereas the
exact comparison of the experimental data with the theoretical prediction is retained to be a complementary
target of this study. Further, the data shown are taken
from diverse experimental sites, requiring, in principle,
separate interpolations. Many of the experimental data
shown in Fig. 2 match poorly with the theoretical prediction, since the experimental precipitation rate A is
below the theoretical minimum meaningful precipitation: Amin = 2 mr1/2 (see the constraint for w0 and n0
to be real). In order to overcome this limitation, a second case was tackled. R and L have been adjusted maintaining the same ratio LR1 in order to leave the plant

961

biomass unchanged [see Klausmeier [7] for more details].


The critical condition, obtained for L = 2 y1 and
R = 50 m4 kg2 y1 is shown in Fig. 3. A better correspondence between model results and eld data is
achieved. The inuence of Ks and a on the stability condition remains the same as in the rst case (Fig. 2) and
similar comments apply to Figs. 2 and 3.
On slopes steeper than 0.2% in arid regions, typical
vegetation bands with a width in the range of a few dozens of meters are observed (see [24] and [15] for a review). Lefever and Lejeune [8] report on the evidence
of an inverse correlation between the band wavelength
and the ground slope, and of a direct correlation between band length and aridity. Figs. 4 and 5 show the
unstable vegetation mode k versus A corresponding to
the critical stability condition plotted in Figs. 2 and 3.
k in the range 0.320.48 corresponds to bands of width
2pk1D1/2L1/2 in the range 2639 m for L = 4 y1 (Fig.
4); and 2028 m for L = 2 y1 (Fig. 5). The evaluated
critical mode appears to be reasonably in agreement
with the observed band characteristic length. There is
no unique trend in the vegetation critical mode k as a
function of the net rainfall amount A. Indeed, depending on Ks and a, k increases with A, decreases, presents
a maximum or a minimum. Conversely, R has minor
inuence on k, in fact Figs. 4 and 5 show substantially
similar results.

4. Discussion and conclusions


A new model for linear stability analysis of vegetation
patterns was derived and solved analytically in order to
obtain the critical stability condition of the soil moisture
and biomass balance equations. The model explicitly
takes into account two commonly referred soil proper-

Fig. 3. Case 2. Grass critical stability condition for dierent slope gradients i and net inow A. ( ) Experimental data taken from literature. Open
symbols: analytical solutions of the marginal stability condition problem Rek = 0 (12). L = 2 y1; R = 50 m4 kg2 year1. Ks = 3000 m y1 (left) and
Ks = 300 m y1 (right); a = 1000 m1 (n); a = 100 m1 ( ) a = 10 m1 (h).

962

N. Ursino / Advances in Water Resources 28 (2005) 956963

Fig. 4. Case 1. Grass critical wave length for dierent slope gradients i and net inow A. L = 4 y1 and R = 100 m4 kg2 year1. Soil parameters:
saturated conductivity Ks = 3000 m y1 (left) and Ks = 300 m y1 (right); a = 1000 m1 (n); a = 100 m1 ( ) a = 10 m1 (h).

Fig. 5. Case 2. Grass critical wave length for dierent slope gradients i and net inow A. L = 2 y1; R = 50 m4 kg2 year1. Soil parameters:
saturated conductivity Ks = 3000 m y1 (left) and Ks = 300 m y1 (right); a = 1000 m1 (n); a = 100 m1 ( ) a = 10 m1 (h).

ties: the soil saturated conductivity and the ratio between the gravity and the capillary forces. The model
is based on the Klausmeiers [7]. All terms and parameters were set accordingly, except those related to subsurface ow. The model results were compared with
literature data. In order to t the experimental data,
the soil parameters had to be stretched beyond the usual
range. The result suggests that the subsurface processes
that have a relevance for banded vegetation formation
may not have been captured and other relevant processes must be taken into account. At least three terms
within the soil moisture balance should be parameterized with care: net vertical inow, leaching, and lateral
destabilizing ow. These depend on the upper and lower
boundary conditions that do not come into play within
the horizontal soil moisture balance. In this case, the
eective parameters are the upscale keys to the scale of
the soil moisture balances relevant processes taking
place at smaller scales.

Despite the limitations discussed above, the results


support some relevant phenomenological evidence: (i)
the threshold slope increases with eective rainfall; (ii)
lateral soil moisture diusion (that may be impeded by
the low wettability of the bare zones) has a destabilizing
eect (leading to a lower threshold gradient) in case of
low precipitation, whereas, conversely, the eect is stabilizing in the case of higher precipitation (leading to a larger threshold gradient); (iii) the soils characterized by
larger saturated conductivity are more prone to the
appearance of unstable plant development (the instability eld is broader); (iv) dierent ratios a between the
gravity and the capillary forces have a minor impact
on the behavior of the less conductive soils. Only point
(i) has already been proven experimentally.
Due to the many simplifying hypotheses that have
been postulated, it seems rather ambitious to pretend
that the model prediction fully matches the manifold
experimental data taken from literature. Indeed they

N. Ursino / Advances in Water Resources 28 (2005) 956963

relate to dierent heterogeneous scenarios. Nevertheless,


the comparison between theoretical and experimental
data suggests that eective soil parameters may be found
resulting in a good t. A major question concerning the
eective parameters is whether they could be physically
supported. The early results presented here induce the
belief that such a physical basis may exist.
The model introduced and discussed here was used to
speculate on the impact of soil properties on unstable
environments. Thus, it has been kept as simple as possible. Nevertheless, the impact that soil parameters may
have on the stability condition, has been clearly demonstrated. The model may be led closer to reality by adding
new degrees of complexity, such as the consideration of
soil heterogeneity or non-averaged over depth soil
parameters. Nevertheless, in its present form, it evidences the crucial role of soil physics in plant development. It may seem rather obvious, but this point is
often neglected. The lack of relevant elements makes
the predictive power of the model less quantitative than
it might be. In order to achieve a predictive power, it
should be coupled, at least, with the surface water and
nutrients balance equations.

References
[1] Ares J, Del Valle H, Bisigato A. Detection of process-related
changes in plant patterns at extended spatial scales during early
dryland desertication. Glob Change Biol 2003;9:164359.
[2] Fernando TM, Cortina J. Spatial patterns of surface soil
properties and vegetation in a Mediterranean semi-arid steppe.
Plant Soil 2002;241:27991.
[3] Galle S, Ehrmann M, Peugeot C. Water balance in a banded
vegetation pattern. A case study of tiger bush in western Niger.
Catena 1999;37:197216.
[4] Gardner WR. Some steady state solutions of the unsaturated soil
moisture ow equation with application to evaporation from a
water table. Soil Sci 1958;4(85):22832.
[5] Gilad E, von Hardenberg J, Provenzale A, Shachak M, Meron E.
Ecosystem engineers: From pattern formation to habitat creation.
Phys Rev Lett 2004;93:098105.
[6] HilleRisLambers R, Rietkerk M, van den Bosch F, Prins HHT, de
Kroon H. Vegetation patterns formation in semi-arid grazing
systems. Ecology 2001;82(1):5061.

963

[7] Klausmeier CA. Regular and irregular patterns in semiarid


vegetation. Science 1999;284:18268.
[8] Lefever R, Lejeune O. On the origin of tiger bush. Bull Math Biol
1997;59:26394.
[9] Ludwig JA, Tongway DJ, Marsden SG. Stripes, strands or
stipples: Modelling the inuence of three landscape banding
patterns on resource capture and productivity in semiarid
woodlands, Australia. Catena 1999;37:25773.
[10] Murray JD. Mathematical biology. Berlin: Springer-Verlag;
1989.
[11] Philip JR. Theory of inltration. Adv Hydrosci 1969;5:21596.
[12] Pullan A. The quasilinear approximation for unsaturated porous
media. Water Resour Res 1990;26:121934.
[13] Richards LA. Capillary conduction of liquids through porous
medium. Physics 1931;1:31833.
[14] Rietkerk M, Boerlijst MC, van langevelde F, HilleRisLambers R,
vandeKoppel J, Kumar L, et al. Self-organization of vegetation in
arid ecosystems. Am Nat 2002;160:52430.
[15] Rietkerk M, Dekker S, de Ruiter PC, van de Koppel J. Selforganized patchiness and catastrophic shifts in ecosystem. Science
2004;305:19269.
[16] Rodriguez-Iturbe I, Porporato A, Ridol L, Isham V, Cox DR.
Probabilistic modelling of water balance at a point: the role of
climate soil, and vegetation. Proc R Soc Lond A 1999;455:
3789805.
[17] Rovinsky AB, Menzinger M. Chemical instability induced by a
dierential ow. Phys Rev Lett 1992;69:11936.
[18] Salvucci GD. Estimating the moisture dependence of root zone
water loss using conditional averaging precipitation. Water
Resour Res 2001;37(5):135765.
[19] Schlesinger WH, Reynolds JF, Cunningham GL, Huenneke LF,
Jarrell WM, Virginia RA, et al. Biological feedbacks in global
desertication. Science 1990;247:10438.
[20] Schmugge TJ, Jackson TL, McKim HL. Survey of methods for
soil moisture determination. Water Resour Res 1980;16:96179.
[21] Slatyer RO. Methodology of a water balance study conducted on
a desert woodland (Acacia aneura F. Muell) community in central
Australia. In: Plantwater relationships in arid and semi-arid
conditions. Proceedings of the Madrid symposium. UNESCO
Arid Zone Research, vol. 16. 1961. p. 1525.
[22] Thiery J, dHerbes JM, valentin CA. A model simulating the
genesis of banded vegetation patterns in Niger. J Ecol 1995;83:
497507.
[23] Turing AM. Philos Trans R Soc London Ser B 1952;237:37.
[24] Valentin C, dHerbes JM, Poesen J. Soil and water components of
banded vegetation patterns. Catena 1999;37:124.
[25] Von Hardenberg J, Meron E, Shachak M, Zarmi Y. Diversity of
vegetation patterns and desertication. Phys Rev Lett 2001;87:
198101.
[26] Worral GA. Patchiness in vegetation in the northern Sudan. J
Ecol 1960;48:10717.

You might also like