Journal of

Ecology 2000,
Fire, resprouting and variability: a recipe for grasstree
88, 213229 coexistence in savanna
STEVEN I. HIGGINS*, WILLIAM J. BOND { and WINSTON S.
W. TROLLOPE{
*Centre for Water in the Environment, Department of Animal, Plant and Environmental Sciences, University of
the Witwatersrand, PO WITS 2050, South Africa; {Department of Botany, University of Cape Town, Private
Bag Rondebosch 7701, South Africa; and {Department of Livestock and Pasture Science, Faculty of
Agriculture, University of Fort Hare, Alice 5700, South Africa

Summary
1 Savanna ecosystems are characterized by the codominance of two dierent life
forms: grasses and trees. An operational understanding of how these two dierent
life forms coexist is essential for understanding savanna function and for predicting
its response to future environmental change.
2 The existing model, which proposes that grasses and trees coexist by a separation
of rooting niches, is not supported by recent empirical investigations. Our aim was
to dene an alternative mechanism of grasstree coexistence in savanna ecosystems.
The model we have built concentrates on life historydisturbance interactions
between grasses and trees.
3 The model demonstrates coexistence for a wide range of environmental condi-
tions, and exhibits long periods of slow decline in adult tree numbers interspersed
with relatively infrequent recruitment events. Recruitment is controlled by rainfall,
which limits seedling establishment, and re, which prevents recruitment into adult
size classes. Decline in adult tree numbers is the result of continuing, but low levels,
of adult mortality. Both aspects of the dynamics are consistent with an established
non-equilibrium mechanism of coexistence (the storage eect).
4 A sensitivity analysis indicated that data on tree resprouting ability, stem growth
rates and the relationship between seedling establishment and wet season drought
are essential for predicting both the range of conditions for which coexistence is
possible and the response of savanna ecosystems to environmental change.
5 Our analysis suggests that understanding grasstree interactions in savanna
requires consideration of the long-term eects of life historydisturbance interac-
tions on demography, rather than the ne-scale eects of resource competition on
physiological performance.

Key-words: bush encroachment, coexistence, environmental change, re, non-equili-

brium dynamics, spatially explicit individual-based model, storage eect
Journal of Ecology (2000) 88, 213229

Introduction with savanna type, the physiognomy of savanna

Savanna is a curious vegetation state characterized
by the coexistence of grasses and trees. Although
the exact ratio of grass to tree varies considerably
Correspondence: Steven I. Higgins, National Botanical
# 2000 British Institute, Private Bag X7, Claremont 7735, South Africa
Ecological Society (fax 27 21 7976903; e-mail higgins@nbict.nbi.ac.za).
remains clearly distinct from that of grassland and
forest. Most authors would agree that a complex
web of factors, notably water, herbivory, re, soil
texture and nutrients, inuences the balance between
grass and trees in savanna (Cole 1986; Skarpe 1992;
Scholes & Walker 1993; Frost 1996). Given this
complexity, the question of how grasses and trees
coexist over such a wide range of climatic, edaphic,
214 biogeographic and historical conditions is intriguing:
Grasstree so intriguing that it has been referred to as the
coexistence in `savanna problem' (Sarmiento 1984).
savanna Initially it was felt that grasstree coexistence
could be explained by equilibrium theories of coexis-
tence. The LotkaVolterra model is the classic equi-
librium model of coexistence; it predicts that stable
coexistence can occur if the eects of intraspecic
competition are greater than the eects of interspeci-
c competition. In essence, in the LotkaVolterra
model the mechanism of coexistence is through
niche dierentiation. It is therefore not surprising
that a niche dierentiation model has been invoked
to explain grasstree coexistence in savanna. The
Walter hypothesis (Walter 1971) proposes that
grasstree coexistence is made possible by separation
of the rooting niche, with trees having sole access to
water in deeper soil horizons and grasses having pre-
ferential access to, and being superior competitors
for, water in the surface soil horizons. The Walter
hypothesis was articulated in an analytical model by
Walker & Noy-Meir (1982), and they demonstrated
that rooting niche dierentiation could allow the
stable coexistence of grasses and trees. Although
some data on root distributions and water uptake
support the Walter hypothesis (Helsa et al. 1985;
Knoop & Walker 1985; Weltzin & McPherson
1997), enough dissenting evidence exists (Johns
1984; Richards & Caldwell 1987; Belsky 1990;
Belsky 1994; Le Roux et al. 1995; Seghieri 1995;
Mordelet et al. 1997) to question its validity as the
ubiquitous mechanism of grasstree coexistence
(reviewed recently in Scholes & Archer 1997).
Evidence against the rooting niche separation
mechanism does not, however, necessarily preclude
the possibility of another equilibrium explanation,
although experimental evidence suggests that inter-
specic competition between grass and trees is often
stronger than intraspecic competition (Scholes &
Archer 1997) and this violates the equilibrium mod-
el's conditions for stable coexistence.
It is clear that alternative theories of coexistence
are needed to explain grasstree coexistence in
savanna. Several mechanisms by which strongly
competing organisms can coexist have been pro-
posed (Shmida & Ellner 1984), although these the-
ories have not been applied to the grasstree
coexistence problem. A promising non-equilibrium
model of coexistence was developed by Chesson &
Warner (1981); their model shows how recruitment
uctuations can promote coexistence between
strongly competing, long-lived organisms in lottery
systems. Later they generalized their ndings
beyond lottery systems and called this mechanism
the storage eect (Warner & Chesson 1985; Chesson
& Huntly 1989). The storage eect depends on the
# 2000 British
occurrence of overlapping generations and uctuat-
Ecological Society
Journal of Ecology, ing recruitment rates; under these conditions the
88, 213229 reproductive potential is `stored' between genera-
tions, allowing the population to recruit strongly
when conditions are favourable. The average popu-
lation growth rate is thus more strongly inuenced
by the benets of the favourable periods than the
costs of the unfavourable periods (Warner &
Chesson 1985). The longevity of savanna trees and
the highly variable climates (which lead to variable
recruitment rates) in savanna ecosystems suggest
that the storage eect could be a signicant contri-
butor to the coexistence of grasses and trees in
savanna. In essence, the promise of the storage eect
suggests that there may be a demographic explana-
tion for the coexistence of grasses and trees; this
represents a departure from existing dynamic models
of savanna ecosystem function (Walker & Noy-Meir
1982; Eagleson 1989; Jeltsch et al. 1996; Jeltsch et al.
1998) which emphasize physiological mechanisms.
In this paper we develop a demographic model of
the interactions between grasses and trees in
savanna. The aim of the model is to (i) integrate our
existing understanding and empirical data on the
demography of savanna ecosystems and (ii) explore
whether a demographic mechanism of grasstree
coexistence can be found. Theoretical models of the
storage eect have already shown the theoretical
possibilities for coexistence. What is needed is to see
whether existing empirical data sets from savanna
systems can be used to build and parameterize a
more realistic model that is consistent with the sto-
rage eect. Coexistence between grasses and trees in
savanna is also an unusual coexistence problem
because the competing organisms belong to unlike
growth forms, yet a ubiquitous niche separation
does not seem to exist. We hope that the model will
help us understand savanna ecosystem dynamics
and the sensitivity of savanna to climate, re, her-
bivory and wood harvesting, or at least help identify
the demographic (proximate) and physiological (ulti-
mate) information needed to predict how savannas
will respond to environmental change.
Model denition
CONCEPTUAL DEFINITION
The model rephrases the grasstree coexistence ques-
tion as: why do grasses not eliminate trees, and why
do trees not thicken up to form forests that would
exclude grasses? We propose that the storage eect
promotes the persistence of trees at low densities
through variations in seedling establishment and
adult recruitment against a background of low adult
mortality. We believe that the storage eect operates
in savanna because (i) seedling establishment rates
depend on rainfall, which is highly variable in
savanna; (ii) grass res, which vary considerably in
intensity in savanna, can prevent tree recruitment;
and (iii) savanna trees are long-lived. It follows that
215 understanding grasstree coexistence requires an
S.I. Higgins, understanding of grass re behaviour, re-induced
W.J. Bond & tree damage, tree recruitment and seedling establish-
W.S.W. Trollope ment (Fig. 1). In this paper we use the term estab-
lishment to refer to the seed to seedling transition,
and the term recruitment to refer to the seedling to
adult transition. The model is based largely on data
and assumptions from southern African studies of
savannas that burn relatively frequently, but similar
savannas that also burn relatively frequently are
characteristic of large areas of Africa, South
America, Asia and Australia. We do not consider
savannas heavily impacted by herbivores, although
we believe that the demographic problems of trees
escaping from re resemble the problems of trees
escaping from browsing (Pellew 1983; Dublin et al.
1990; Prins & van der Jeugd 1993).
Grass res occur in savanna ecosystems because
grass production in the wet season is followed by an
extended dry season leading to a continuous cover
of fuel, and there is a ready source of ignitions
(lightning and human). We hypothesize that savan-
nas exist under conditions where res are intense
enough to limit the recruitment rates of trees, but
not so intense to prevent recruitment (as in grass-
lands) or so mild to not limit recruitment (as in sur-
face res in forests). Variation in re intensity can
be attributed to variations in grass standing crop,
grass moisture content (which varies with species
and season), air temperature, humidity and wind
speed (Trollope 1982; Cheney et al. 1993; Cheney &
Sullivan 1997; Trollope 1998). Spatial variation in
re intensity may therefore be due to patchy grass
production (Chidumayo 1997), patchy herbivory
(Coughenour 1991), the eects of tree neighbour-
hoods on grass production (Mordelet & Menaut
1995) and grass moisture contents (Vetaas 1992;
Webber 1997). The temporal variation in re inten-
sity may be due to both interannual variation of
rainfall and variation in the timing of ignition events
and hence fuel conditions (Trollope 1982; Cheney &
Sullivan 1997). In our model (Fig. 1) rainfall is the
primary determinant of grass production. Local site
variables, notably soil characteristics and nutrient
availability, obviously also inuence grass produc-
tion (Scholes & Walker 1993), as does grass species
composition (Trollope et al. 1989), but such subtlety
is not our concern. Grass standing crop increases
during the wet growing season and decreases as the
dry season progresses, due to herbivory and decom-
position; the moisture content also decreases as the
dry season progresses. The realized re intensity is
therefore dependent on the grass standing crop and
grass moisture content on the day of the re, the
species of grass (as dierent species have dierent
# 2000 British
moisture contents), as well as the temperature,
Ecological Society
Journal of Ecology, humidity and wind speed on the day of the re
88, 213229 (Fig. 1). It follows that the intensity of a re regime
is strongly dependent on the seasonal distribution of
ignition events.
Understanding the variation in tree recruitment
needs not only an understanding of variation in re
intensity but also an understanding of the life his-
tory of savanna trees. Savanna trees only recruit
into the adult population once they escape the zone
of inuence of grass res. The ability of stems that
are killed in a re to resprout is a key life-history
trait that promotes the persistence of trees in
savanna (Walter 1971; Bond & van Wilgen 1996;
Gignoux et al. 1997; Trollope 1998). Tree seedlings
may persist as suppressed juveniles (called `gullivers'
by Bond & van Wilgen 1996) for many years
because such stems continue to resprout repeatedly
after being burnt back by res. We model the fre-
quency of escape of gullivers from the ame zone
into the adult population by simulating how re
intensity and tree size inuence the likelihood of
stem mortality (Fig. 1). Taller, thicker stems and
stems with thicker bark have a higher chance of sur-
viving a re of a given intensity (Wright et al. 1976;
Moreno & Oechel 1993; Gignoux et al. 1997;
Trollope 1998; Williams et al. 1999). Hence the fre-
quency of gulliver escape depends strongly on stem
growth rates and the frequency and intensity of re
(Trollope 1984).
Gulliver banks are maintained by both resprout-
ing and seedling establishment. Little is known
about the regeneration niches of savanna tree spe-
cies. It is generally believed that the seedlings of
many savanna species are shade intolerant (Smith &
Shackleton 1988), and high grass biomass can sup-
press tree seedlings (Brown & Booysen 1967; Walker
et al. 1981; Knoop & Walker 1985; Harrington
1991). Other evidence suggests that establishment is
facilitated by the presence of grasses (Brown &
Archer 1989; Holmgren et al. 1997; Davis et al.
1998), and that some savanna species are shade tol-
erant (Smith & Walker 1983; O'Connor 1995;
Homann 1996). What is clear is that most savanna
germinants cannot tolerate droughts during the wet
growing season (du Toit 1965; Medina & Silva 1990;
Harrington 1991; Hodgkinson 1991; O'Connor
1995; Homann 1996). It seems, therefore, that the
likelihood of wet season droughts will strongly inu-
ence seedling establishment patterns and hence
grasstree coexistence (Fig. 1).
OPERATIONAL DEFINITION
We developed an individual-based, spatially explicit
simulation model of grass and tree dynamics
because this class of model allows exible simulation
of a wide range of ecological processes. In particular
the individual-based approach allows us to keep
track of the size and fate of individual tree stems as
inuenced by their neighbourhoods. Because of the
216
Grasstree
coexistence in
savanna
Fig. 1 Conceptual model of grasstree interaction. The model shows the factors that inuence seedling establishment, re
intensity and the probability of stem mortality.
large dierences in the size of grasses and trees, we
do not model individual grass tufts but model grass
patches. Because we are interested in capturing the
heterogeneity introduced by tree neighbourhoods on
grass patches, we chose a spatial grain of 1 m2 and
assumed that only one stem can occupy each 1-m2
site. The area the model simulates can be varied, but
for this study we use a 1-ha area (100  100 cells).
We chose to use an annual time step because we
postulate that interannual variation in rainfall is the
key source of variation, and because most of the
data we had access to was annual data. However, as
discussed above, fuel properties vary considerably
within a year; we deal with this problem by allowing
the day of re ignition to be a random variable of
dened moments. Below we discuss the assumptions
and the functions used to simulate the ecological
processes described above; we then describe how we
combine these functions to dene a dynamic model.
RAINFALL
Mean annual rainfall is variable in savanna systems;
this variability can be divided into two components.
The rst component is stochastic variation; the sec-
ond component is long-term periodicity. We use a
sine wave function, which captures both these com-
ponents of variability, to generate rainfall:
 y
R xRx ; Rsd sin 2p s; Re0 eqn 1
# 2000 British l
Ecological Society
Journal of Ecology, Here R is the annual rainfall (mm); x is a nor-
88, 213229 mally distributed random number dened by the
mean (Rx; mm) and standard deviation (Rsd; mm) of
annual rainfall; s is the eect of the long-term peri-
odicity of rainfall (mm); l is the frequency of peri-
odicity; and y is the simulation year.
GRASS PRODUCTION
Most authors use linear regression to describe the
relationship between rainfall and grass production,
as this produces the best t to the data from
savanna regions (O'Connor 1985; Scholes & Walker
1993). Grass production can be written as:
Gp gg R eqn 2
where Gp is the predicted above-ground production
(kg ha 1), R is the annual rainfall (mm), and gg is
the growth coecient. Using data from southern
Africa we estimated Gp 3.369  R (P < 0.0001,
d.f. 71; Fig. 2). Grass production can be nega-
tively (Grunow et al. 1980; O'Connor 1985;
Mordelet & Menaut 1995) or positively inuenced
by tree neighbourhoods (Belsky et al. 1989; Weltzin
& Coughenour 1990). Grass production beneath
tree canopies can be boosted by almost 300% or
suppressed by over 50% (Mordelet & Menaut
1995). To account for the eect of stem neighbour-
hoods on grass production we can write:

gg R ; if Ci;j 0
Gpi;j eqn 3
gg Rc ; if Ci;j 1
Here c is a coecient that describes the eect of
the stem neighbourhood on grass production at site
217
S.I. Higgins,
W.J. Bond &
W.S.W. Trollope
Fig. 2 Grass productionrainfall relationship from the savanna regions of southern Africa. Data are from O'Connor (1985),
Scholes & Walker (1993) and O'Connor & Bredenkamp (1997). These data are used to dene the rainfallgrass production
relationship used in the model (see operational denition).
(i,j) and C(i,j) is an array that describes whether the
site (i,j) is in a stem neighbourhood (C(i,j) 1) or
not (C(i,j) 0); stem neighbourhoods are dened
below (equation 12). The grass standing crop at the
end of the growing season (G(i,j)) is therefore:
Gi;j Gpi;j Gy1i;j eqn 4
where Gy 1(i,j) is the amount of grass material that is
carried over from the previous year (cf. equation 6).
The levels of herbivory, the rate of grass decomposi-
tion and whether a re has occurred will determine
how much grass is carried over from one year to the
next.
HERBIVORY AND DECOMPOSITION
While herbivores are a ubiquitous feature of
savanna ecosystems and inuence savanna dynamics
in many ways (Cumming 1982; Pellew 1983; Dublin
et al. 1990; Prins & van der Jeugd 1993; Scholes &
Walker 1993), we are primarily interested in herbi-
vores' ability to manipulate fuel loads; for this rea-
son we do not consider browsing. Following
Danckwerts (1982) we assume that herbivores can
reduce the grass standing crop as a linear function
of time since production:
Gfi;j Gi;j ati eqn 5
where Gf(i,j) is the grass standing crop on the day of
ignition, ti is the ignition day (days since the start of
the dry season), a is the grazing rate (kg ha 1
day 1), and G(i,j) is the grass standing crop at the
end of the growing season. Note that this function
# 2000 British
Ecological Society
implies that growth and consumption are treated as
Journal of Ecology, discrete events in the model, and ignores spatial and
88, 213229 temporal heterogeneity in grazing. To estimate the
amount of grass carried over from one year to the
next (Gy 1(i,j)), we could write:

Gi;j atd utd ; if Bi;j 0
Gy1i;j
Gi;j atd utd bGi;j ; if Bi;j 1
eqn 6
Here u is the decomposition rate (kg ha 1 day 1);
td is the length of the dry season in days; b is the
completeness of the burn; and B(i,j) is an array that
describes whether site (i,j) is burnt or not.
GRASS MOISTURE CONTENT
The moisture content of the fuel inuences re
intensity. Grass growing in a tree's neighbourhood
may be moister and retain moisture levels for longer
into the dry season (Weltzin & Coughenour 1990;
Vetaas 1992; Webber 1997). If we assume that the
moisture content of grass decays exponentially into
the dry season (Cheney & Sullivan 1997), we could
write the moisture of grass on the day of ignition as:

Mo expdo ti ; if Ci;j 0
Mf i;j eqn 7
Mc expdc ti ; if Ci;j 1
Here Mf(i,j) is the moisture content (%) at location
(i,j) ti days after the start of the dry season; Mo and
Mc are the moisture contents of grass (outside and
inside the tree neighbourhoods) at the end of the
growing season; and do and dc are the drying rates
for outside and inside the tree neighbourhood. The
array C(i,j) records if a cell is inside a tree neighbour-
hood (see equation 12 for the denition of tree
neighbourhood).
218 FIRE INTENSITY
Grasstree
We use empirically derived relationships to predict
coexistence in
re intensity, as these statistical models provide,
savanna
given the information available, a better prediction
of re intensity than physical re models (Trollope
1998). The statistical model was developed using
200 monitored res in South African savannas (P
< 0.01, d.f. 196, R2 0.60; Trollope 1998). The
model was tested against independent re behaviour
data and accounted for 56% of the variation in re
intensity (Trollope 1998). The multiple regression
model is:
p 8
Qi;j 2729 0:8684Gfi;j 530 Mfi;j
596
0:907H 2 eqn 8
W vr
Here Q(i,j) is the re intensity (kJ s 1 m 1) at
site (i,j); Gf(i,j) is grass standing crop (kg ha 1) at
site (i,j); Mf(i,j) is fuel moisture (%) at site (i,j); H is
the relative humidity (%); and W is wind speed (m
s 1). The empirical model therefore proposes that
the realized re intensity is dependent on the grass
standing crop, the grass moisture content, the rela-
tive humidity, and the wind speed on the day of the
re. For the simulations we assume that humidity
and wind speed are normally distributed random
numbers dened by the site's mean and standard
deviation of humidity (Hx,Hsd ;H r 0) and wind
speed (Wx,Wsd ;W r 0).
STEM MORTALITY
The probability of stem mortality (or `topkill') in a
re is a function of stem height and re intensity
(Trollope 1984). We used data on the survival rates
of 7400 stems of 76 species in 40 res of known
intensities (W.S.W. Trollope, A.L.F. Potgieter and
N. Zambatis, unpublished data) to estimate a logis-
tic regression model of the probability of stem mor-
tality (P < 0.01, R2 0.48, d.f. 7397):
p height (hmax) face a pm chance of mortality each
exp4:3 5:003lnh 0:004408 Q
pt p year.
1 exp4:3 5:003lnh 0:004408 Q
eqn 9
Here pt is the probability of stem mortality; h is
stem height (m); and Q is the re intensity (kJ s 1
m 1; equation 8). Interestingly Williams et al.
(1999), using data from a single intense re in a tro-
pical savanna in northern Australia, found that pt
was a quadratic function of tree size, with larger
# 2000 British
Ecological Society
and small trees suering highest pt. The susceptibil-
Journal of Ecology, ity of large trees to topkill was attributed to termite
88, 213229 damage to large trees (Williams et al. 1999).
STEM RESPROUTING
Stems that have been topkilled usually resprout
from rootstocks. Savanna species have very high
probabilities of resprouting (Lacey et al. 1982;
Trollope 1982, 1984; Boo et al. 1997; Gignoux et al.
1997). Resprouting ability is generally thought to
increase with stem size (Wright et al. 1976; Moreno
& Oechel 1993) but, in some tree species, decreases
again in the larger size classes (Trollope 1974;
Hodgkinson 1998; K. Maze and W.J. Bond, unpub-
lished data). The eect of stem size on the probabil-
ity of resprouting ( pr) can be written as:
> 0 ; if h<hr
>
< pmax
pr   eqn 10
>
> h h0:5 ; if hehr
: 1 exp
where pmax is the maximum probability of resprout-
ing, h0.5 is the stem height (h) at which there is a
50% chance of resprouting, and vr is a constant that
describes how rapidly the probability of resprouting
changes with stem height. Recent germinants do not
have the root reserves to resprout (Moreno &
Oechel 1993); we therefore assume that stems less
than the resprouting height (hr) cannot resprout.
The model only allows one stem per site; it therefore
does not consider the resprouting of multiple stems.
Because the probability of re survival is inuence
by tree height and not stem number, we do not con-
sider this an important limitation.
TREE MORTALITY
Recent carbon dating evidence suggests that
savanna trees can be more than 1000 years old
(CSIR, personal communication). Rates of mortality
due to stress are therefore expected to be low.
Andersen et al. (1998) reported annual mortality
rates of 0.01 for Australian savannas; Trapnell
(1959) reported mortality rates of 0.04 in
Zimbabwean miombo woodlands; and Shackleton
(1997) reported mortality rates of 0.05 in South
African savannas. In the model trees of maximum
STEM GROWTH RATES
The stem mortality and stem resprouting functions
all use stem height information. Very little data exist
on height or diameter growth of savanna stems. We
know that stems initially grow rapidly in height but
subsequently growth slows (K. Maze and W.J.
Bond, unpublished data); using this information we
can describe stem growth using a dierence equa-
tion:
  
219 hy1
h hy1 1 gs eqn 11
S.I. Higgins, hmax
W.J. Bond & where gs is the growth rate of stems (cm year 1),
W.S.W. Trollope hmax is the maximum stem height (m), and hy1 is
the stem height in the previous year. Stem growth
rates and maximum tree heights in savannas are
known to be inuenced by moisture and nutrient
availability (Shackleton 1997), although we do not
explicitly consider these eects here.
STEM NEIGHBOURHOODS
A stem's neighbourhood has two components. First,
a stem's canopy creates a moister and shadier envir-
onment; secondly, the laterally spreading roots inu-
ence the soil moisture and soil nutrient status. The
root- and canopy-dened neighbourhoods do not
always overlap (Vetaas 1992), but we assume, never-
theless, that the diameter of the neighbourhood (n;
m) increases as a linear function of tree size:
n k gn h eqn 12
where gn is a growth coecient, h is stem height
(m), and k is a constant. We do not have data on
below-ground neighbourhoods and we therefore use
data on the relationships between stem height and
canopy diameter (W.S.W. Trollope and A.L.F.
Potgieter, unpublished data) to estimate k and gn.
SEED PRODUCTION
Following Ribbens et al. (1994), we dene seed pro-
duction as a function of tree size, such that:
8 lings of other species are shade-tolerant and little
>
< 0 ; if h<hf
 2
F h eqn 13
>
:f ; if hehf
hstd
Here F is tree fecundity (seeds year 1) and f is the
number of seeds dispersed by a tree of a reference
height (hstd); stems smaller than the height of repro-
ductive maturity (hf) do not produce seeds. Data on
the seed production of savanna trees (Tybirk et al.
1993) is used to estimate hf and f.
SEED DISPERSAL
Many savanna tree species are dispersed passively
and by animals (bird, ungulates, rodents, termites
and ants; Brown & Archer 1989; Tybirk et al. 1993;
Miller 1994). Previous demographic models of
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Ecological Society
savanna (Menaut et al. 1990; Hochberg et al. 1994;
Journal of Ecology, Jeltsch et al. 1996, 1998) have emphasized the impor-
88, 213229 tance of tree clumps and hence the role of local vs.
long-distance dispersal. The importance of local vs.
long-distance dispersal motivated us to model dis-
persal as a stratied process that explicitly considers
both local and long-distance dispersal (Higgins &
Richardson 1999). We can use a mixture of two
exponential distributions to describe a probability
density function of dispersal distances d(x):
dx pl exp bl x 1 pl exp bf x eqn 14
Here pl is the proportion of seeds that are dis-
persed short distances (described by the parameter
bl) and (1pl) is the proportion of seeds that are
dispersed longer distances (described by the para-
meter bf).
SEED BANK DECAY
Seed banks of savanna tree species are not very
long-lived. Those that do not suer predation by
insects and rodents either decay rapidly, germinate
or lose viability (Tybirk et al. 1993). We summarize
all these processes by assuming that a constant pro-
portion of seeds decay each year (sd). We could not
nd any published estimates of sd, although anecdo-
tal evidence suggests that the decay rate is relatively
high (Skoglund 1992; Tybirk et al. 1993; Miller
1994; Chidumayo & Frost 1996).
SEEDLING ESTABLISHMENT
Little is known about the regeneration niches of
savanna tree species, although we know that many
seedlings are shade-intolerant and that high grass
biomass can suppress their recruitment, while seed-
inuenced by grass biomass. What is clear is that
one of the major factors limiting establishment is
the availability of moisture: droughts during the wet
season of more than 30 days can lead to seedling
mortality (Medina & Silva 1990; Chidumayo &
Frost 1996; Homan 1996). If we assume that the
number of rainfall events during the wet season is
positively correlated with annual rainfall, then the
probability of wet season drought should decrease
with annual rainfall. We can express the probability
of wet season drought ( pd) as:
1
pd   eqn 15
R R0:5
1 exp
vd
Here R0.5 is the annual rainfall at which there is a
0.5 chance of a wet season drought; and vd is a con-
stant that describes the rate at which the probability
of wet season drought changes with rainfall (R). If
there is no wet season drought then the probability
of establishment ( pe) is:
 8
220 >
> 0 ; if fCi;j 1 & ct 1g
>
<
Grasstree 1
pei;j  
coexistence in >
> G i;j G0:5 ; if fCi;j 0 & ct 1g
>
: 1 exp
savanna ve
Here we account for the observations that some
species require light for establishment (ct 1), while
others are shade tolerant (ct 0). If the light condi-
tions are suitable (equation 16.2) then the probabil-
ity of establishment is a function of the grass
standing crop. In equation 16.2 G0.5 is the grass
standing crop at which the probability of establish-
ment ( pe) is 0.5, G(i,j) is the grass standing crop and
ve is the rate at which the probability of establish-
ment changes with grass standing crop.
IMPLEMENTATION
Each year the model sequentially simulates the fol-
lowing ecological processes: rainfall, tree growth,
seed dispersal, grass production, potential re inten-
sity and re spread, stem mortality due to res,
resprouting, adult tree mortality, seedling establish-
ment, and seed bank decay. Rainfall is generated
using equation 1. Tree heights are incremented each
year (equation 11) and tree neighbourhoods are sub-
sequently calculated from the tree height (equation
12). The number of seeds produced by each stem is
calculated (equation 13) and these are available for
dispersal. Each seed is dispersed individually and
the distance each seed moves is a random number
dened by the mixture distribution (equation 14).
This distance and a randomly selected direction are
used to calculate the location of each seed. Grass
production is calculated as a function of the rainfall
and neighbourhood state (equation 3). Estimating
potential re intensity (equation 8) requires the esti-
mation of grass standing crop, grass moisture con-
tent, relative humidity and wind speed on the day of
the re. We assume that ignition can occur on any
day during the dry season. The relative humidity
and wind speed on the day of the re are generated
by assuming that daily humidity and wind speed are
normally distributed random numbers. The grass
standing crop and grass moisture content on the day
of the re can be estimated by using equations 5
and 7 and by assuming that ignition occurs ti days
into the dry season. In the current version of the
model one re ignition occurs per year. We assume
that ti is a normally distributed random number
with a mean (Ix) and standard deviation (Isd) char-
acteristic of the temporal distribution of ignition
events. This method of generating ignition events
emphasizes the temporal distribution of ignition
# 2000 British
Ecological Society
events, i.e. there is an emphasis on re intensity
Journal of Ecology, rather than re frequency. The re spread algorithm
88, 213229 is analogous to that proposed by Turner & Romme
eqn 16.1
eqn 16.2
or {ct = 0}
(1994). We assume that res can spread if a thresh-
old re intensity is exceeded; this threshold has been
estimated as 150 kJ s 1 m 1 in savanna systems
(van Wilgen & Scholes 1997). The re spread algo-
rithm allows a re to spread to neighbouring cells if
a neighbouring cell's potential re intensity exceeds
the threshold. The re spread algorithm is not inu-
enced by wind or topography. The way we model
re ensures that fuel properties rather than ignition
frequency determines the modelled re frequency.
The average re intensity in the nine 1-m2 cells in a
tree's neighbourhood and the tree's height are used
to estimate the probability of stem mortality (equa-
tion 9). The likelihood of a dead stem resprouting is
estimated as a function of stem height (equation 10).
Adult trees face a pm probability of death each year.
Seeds can only germinate and establish if there is no
wet season drought (equation 15). If there is no
drought then a seed can establish, provided the
grass biomass is low enough, light conditions are
suitable (equation 16) and the site is unoccupied by
a tree stem. The seed bank is decayed by a constant
proportion (sd) each year.
Model behaviour
Our analysis of the model's behaviour is divided
into two sections. We rst examine whether the
model successfully predicts coexistence, and attempt
to understand the behaviour of the model in the
context of theoretical coexistence models. The sec-
ond section analyses the sensitivity of the coexis-
tence state to key parameters and hence investigates
the environmental conditions for which we would
expect grasstree coexistence.
THE NATURE OF GRASSTREE
COEXISTENCE
For this section we initialized the model with the
best parameter estimates available. Sources of these,
largely southern African, parameter estimates are
discussed under operational denition, and the para-
meter values used are listed in Table 1. We then var-
ied the base parameterization (Table 1) to simulate
four sites representing a rainfall gradient from arid
to mesic savanna (Table 2). In southern Africa this
gradient is associated with a change from palatable
to relatively unpalatable grass; and we simulate this
by decreasing the grazing rate with increasing rain-
fall. We also assume that (i) stem growth rates will
increase with rainfall and (ii) that the periodicity
221 Table 1 Parameter symbols, names and default values used for the simulation runs. Sources of parameter estimates are dis-
cussed under operational denition
S.I. Higgins,
W.J. Bond & Symbol Parameter name Default value
W.S.W. Trollope
Rx Mean annual rainfall 1000 mm
Rsd Standard deviation of mean annual rainfall 62 mm
s Strength of periodicity in rainfall 188 mm
l Period length of periodicity in rainfall 20 years
gg Grass growth coecient 3.369 kg ha 1mm 1
c Eect of tree neighbourhood on grass production 1
a Grazing rate 7 kg ha 1day 1
u Decomposition rate 1 kg ha 1day 1
b Completeness of burn 0.9
Mo Moisture content of between canopy grass 30%
Mc Moisture content of beneath canopy grass 50%
do Drying rate of between canopy grass 0.01
dc Drying rate of beneath canopy grass 0.001
Hx Mean daily humidity 20%
Hsd Standard deviation of mean daily humidity 20%
Wx Mean daily wind speed 5 m s 1
Wsd Standard deviation of mean daily wind speed 5 m s 1
Ix Mean day of re ignition (days after growing season) 150 days
Isd Standard deviation of day of re ignition (days after growing season) 50 days
pmax Maximum probability of resprouting 0.9
h0.5 Stem height for 50% chance of resprouting 800 cm
vr Rate of change of resprouting probability with stem height 100
hr Height at which resprouting ability is attained 30 cm
pm Probability of stem mortality due to age 0.001
gs Growth rate of stems 60 cm year 1
hmax Maximum stem height 600 cm
gn Growth coecient of stem neighbourhood 0.5
k Constant describing change in stem neighbourhood 0.3
f Seeds produced by a stem of reference height (hstd) 4 seeds year 1
hstd Reference stem height 400 cm
hf Height of reproductive maturity 300 cm
pl Proportion of seeds dispersed locally 0.9
bl Scale parameter for local dispersal 0.5
bf Scale parameter for long-distance dispersal 0.02
sd Rate of seed decay 0.7
R0.5 Annual rainfall for 50% chance of wet season drought 700 mm
vd Rate of change of wet season drought probability with annual rainfall 50
G0.5 Grass biomass for 50% chance of seedling establishment 2500 g m 2
ve Rate of change of seedling establishment probability with grass biomass 400
ct Shade tolerance (binary factor) 0

and stochasticity in rainfall (equation 1) changes chasticity and strength of periodicity of rainfall. The
from arid to mesic savanna. The fact that the coe- relative contribution of the periodicity vs. stochasti-
cient of variation of rainfall tends to decrease with city in rainfall to the coecient of variation of rain-
increasing rainfall is simulated by changing the sto- fall is varied to simulate situations where rainfall

Table 2 Parameter symbols, names and parameter settings used for the simulation runs to describe four savanna sites across
a rainfall gradient. Other parameters are set to the values listed in Table 1

Site name

Symbol Parameter name Arid Semi-arid Semi-mesic Mesic

Rx Mean annual rainfall (mm) 300 600 1000 1400

Rsd Standard deviation of mean annual rainfall (mm) 120 38 62 140
# 2000 British S Strength of periodicity in rainfall (mm) 0 112 188 0
Ecological Society A Grazing rate (kg ha 1 day 1) 12 10 7 2
Journal of Ecology, gs Growth rate of stems (cm year 1) 35 45 60 80
88, 213229
222
Grasstree
coexistence in
savanna

Fig. 3 Four-thousand year trajectory of adult and gulliver (non-reproductive) stem numbers for four hypothetical sites
spanning arid to mesic savanna. The four parameterizations are variations on the default parameterization of the model
(see Table 1 for default parameter settings and Table 2 for the variations used in these runs). The model was initiated with a
0.1 tree density; we show only data from year 1000 to year 5000 to remove the eect of initial conditions.

cycles are not present. While we do not claim that
these parameterizations are full representations of
the dierences between arid and mesic savannas,
they do illustrate the dierent kinds of dynamics the
model can produce.
Running the model with the parameter settings
listed in Table 2 generates coexistence between
grasses and trees at all four sites (Fig. 3), in that
trees persisted but did not reach 100% cover. The
tree dynamics at all sites were characterized by long
periods of slow decline in adult stems punctuated by
occasional recruitment events. The frequency of
recruitment events and the ratio of gulliver (non-
reproductive) to adult stems and the stem densities
vary across the rainfall gradient; these dierences
are best explained by examining the mean and var-
iance in rates of establishment, recruitment and
mortality (Fig. 4). First, low mortality rates at all
# 2000 British
sites explains the slow rate of decline of adult num-
Ecological Society
Journal of Ecology, bers in the absence of recruitment. The low gulliver
88, 213229 relative to adult stem numbers at the arid site is due
to low seedling establishment rates (due to the high
frequency of wet season droughts, cf. equation 15)
and the fact that when establishment does occur at
the arid site it often leads to tree recruitment (due to
the low frequency of high-intensity res caused by
low fuel loads). At the semi-arid and semi-mesic
sites establishment rates are relatively high (due to
less frequent drought and low grass standing crop)
but recruitment into the adult stage is lower and
more variable (due to relatively intense res). The
combination of high establishment rates and low
recruitment explains the accumulation of large num-
bers of gullivers at both these sites. The high var-
iance in gulliver stem numbers at the semi-arid site
is due to relatively high establishment rates, coupled
with a slow growth rate that prevents many of the
smaller gulliver stems from resprouting after topkill.
At the semi-mesic site gulliver stem numbers accu-
mulate because they are large enough to resprout
but too small to recruit frequently. At the mesic site
establishment rates are lower due to the negative
223
S.I. Higgins,
W.J. Bond &
W.S.W. Trollope
Fig. 4 Mean and coecient of variation of seedling estab-
lishment, tree recruitment and mortality rates generated
from low density ( < 0.01) model runs of 5000 years using
the default parameterization of the model for four
hypothetical sites spanning arid to mesic savanna. The four
parameterizations are variations on the default parameteri-
zation of the model (see Table 1 for default parameter set-
tings and Table 2 for the variations used in these runs).
eect of high grass standing crop (produced by the
higher rainfall) on seedling establishment, but the
higher growth rates of stems means that recruitment
rates are maintained. The lower coecient of varia-
tion of rainfall at the mesic site accounts for the
lower variance in establishment and recruitment
rates and hence the more constant adult population
size.
While the patterns in establishment, recruitment
and mortality are instructive, the challenge is to
understand in more general terms the factors
responsible for generating coexistence. It is estab-
lished in the coexistence literature that varying
environments are themselves not recipes for coexis-
tence: some interaction between environmental
variability and species behaviour is needed for varia-
bility to promote coexistence (Turelli & Gillespie
1980; Chesson & Warner 1981; Chesson & Huntly
1989). It follows that understanding coexistence in
heterogeneous systems requires understanding the
interaction between environmental variability and
life history. One such interaction occurs when adult
survival is high and recruitment rates are variable;
# 2000 British
and it is this combination of factors that constitutes
Ecological Society
Journal of Ecology, the storage eect (Warner & Chesson 1985). The
88, 213229 low adult mortality and variable seedling establish-
ment and recruitment rates produced by our model
(Fig. 4) are consistent with the storage mechanism of
coexistence (Warner & Chesson 1985). Moreover,
partitioning out the contribution of the storage
eect to the growth rate of the population (follow-
ing Warner & Chesson 1985) shows that the popula-
tion growth rate for trees was negative or zero when
the storage eect was excluded ( 0.0019, 0.0019,
0.0020, 8.0E 05; for the arid to mesic sites,
respectively, cf. Table 2) and positive with the sto-
rage eect included (0.0024, 0.0050, 0.0071, 0.0055),
suggesting that the storage eect is essential for the
persistence of trees in the model system. The reasons
for the relatively high and constant adult survival
rates are clear: savanna trees are long lived and have
a low likelihood of suering re-induced stem mor-
tality (equation 9). The relatively high variance in
tree seedling establishment and recruitment rates
can be related, respectively, to the variations in rain-
fall and re intensity. In more arid systems variation
enters at the seedling establishment and recruitment
phase, whereas in more mesic systems variation
enters primarily at the tree recruitment phase (Fig.
4). In agreement with this result, high variance in
recruitment rates has been reported in Australian
savannas (Harrington 1991). Harrington (1991)
attributed the high variance in recruitment rates to
the rarity of synchronization between adequate
moisture conditions for seedling establishment and
res of intensities low enough to allow recruitment.
The eect of the rarity of such synchronization is
most easily detected in the model runs from the arid
site (Fig. 3).
THE SENSITIVITY OF GRASSTREE
COEXISTENCE TO ENVIRONMENTAL
CHANGE
Exploring the sensitivity of the coexistence state to a
range of potential inuencing variables can help us
understand the factors that inuence grasstree
coexistence as well as its susceptibility to environ-
mental change. The model we have constructed,
however, contains 48 parameters [40 are listed in
Table 1; the re intensity and the stem mortality
models (equations 8 and 9) contain an additional
eight parameters]: consequently an exhaustive sensi-
tivity analysis is not feasible here. We do, however,
explore the sensitivity by varying key parameters
from the base parameter estimates as dened in
Table 1. For the sensitivity simulation runs the
model was run for 2000 simulation years, a single
run was used for each parameter setting, and the
mean and standard deviation of tree density in the
last 500 simulation years is reported.
We rst vary the mean annual rainfall from 200
to 2000 mm (Fig. 5a; constant humidity); this shows
that the trees can coexist with grasses between 500
and 1600 mm mean annual rainfall. At low rainfall
224
Grasstree
coexistence in
savanna

Fig. 5 Sensitivity of the number of adult tree stems to variation in key model parameters; all other parameters are set to the
default parameter settings (Table 1). The points and bars are the mean and standard deviation of stem density for the last
500 years of a 2000-year simulation run. The model was initiated with a 0.1 tree density for these runs.

the model predicts that trees are limited by moisture shown that, in arid savanna, re cannot control tree
# 2000 British
conditions for establishment, whereas at high rain- densities but can keep trees in the browse zone. Our
Ecological Society
Journal of Ecology, fall trees are limited by re intensity (in agreement model predicted highest tree densities at lower rain-
88, 213229 with Trollope 1980). Trollope (1974, 1980) has fall, suggesting that an additional factor such as
225 browsing (which is excluded here) is needed to con-
S.I. Higgins, trol tree densities in more arid areas. Browsing was
W.J. Bond & also regarded as important in regulating tree densi-
W.S.W. Trollope ties in east African savannas (Pellew 1983; Prins &
van der Jeugd 1993). However, because other para-
meters (e.g. tree growth rates, humidity and wet sea-
son drought) co-vary with rainfall, the rainfall range
that allowed coexistence here can only be taken as a
rough guide to the rainfall limits of savanna. For
instance, if we increase humidity as we increase rain-
fall to simulate less ammable fuel conditions (Fig.
5a; increasing humidity) the model does not predict
tree exclusion at higher rainfall. None the less,
savanna ecosystems occur over a similar range of
mean annual precipitation to the range predicted
here (c. 3001800 mm; Scholes & Walker 1993;
O'Connor & Bredenkamp 1997; Scholes 1997).
Hence both our model and empirical data suggest
that rainfall is a key determinant of grasstree
ratios; although other factors must also inuence
this ratio.
We have established that variability in recruit-
ment rates and low adult mortality rates (Fig. 4)
allow trees to coexist with grasses, but the key to
understanding the coexistence mechanism is deter-
mining what generates this variability. The model
does not appear to be sensitive to the variability
(Fig. 5b; the coecient of variation in rainfall is
increased by increasing s and Rsd) or the stochasti-
city (Fig. 5c; the contribution of Rsd is increased and
the contribution of s to the coecient of variation
in rainfall is decreased) in rainfall. However, remov-
ing variance in re intensity (by changing variation
in rainfall, relative humidity, wind speed and re
ignition day) can lead to the exclusion of trees,
whereas increasing this variance favours trees (Fig.
5d). Hence variable re intensities provide opportu-
nities for tree stems to escape the ame zone, where
they are most susceptible to re, and recruit into the
more re-resistant size classes. In other words, var-
iance in re intensity produces the variance in
recruitment rates that is necessary for the storage
eect to operate. Our model therefore suggests that
a re-mediated recruitment bottleneck (Walter 1971;
Trollope 1974; Bond & van Wilgen 1996; Gignoux
et al. 1997; Andersen et al. 1998) is central to under-
standing how re mediates coexistence of grasses
and trees. Other factors do, however, inuence the
regeneration niche and hence the tightness of the
recruitment bottleneck. Very low seed production
can suppress tree numbers, but the eect of seed
production on tree density rapidly asymptotes ( f;
Fig. 5e). The eect of grass competition on seedling
establishment (G0.5; Fig. 5f) does not appear to inu-
ence tree density. The likelihood of wet season
drought (R0.5; Fig. 5g) does inuence tree density;
# 2000 British
this parameter is likely to be more important in arid
Ecological Society
Journal of Ecology, systems, where variation in recruitment is controlled
88, 213229 by variance in establishment conditions rather than
variance in recruitment conditions (Figs 3 and 4).
Interestingly, although the presence of moister sub-
canopy grass is likely to facilitate the coexistence of
grasses and trees by buering tree stems against re
intensity, increasing the moisture content of the sub-
canopy grasses does not lead to increased tree num-
bers (the subconopy moisture content is increased
and the subcanopy drying rate is decreased relative
to the between canopy values; Fig. 5h). This suggests
that the spatial component of heterogeneity in re
intensity is not as important as the temporal one
(Fig. 5d). However, the eects of tree canopies on
grass production could introduce additional spatial
heterogeneity; such eects were not considered here
(c was set to 1 for these simulation runs).
The rate of stem growth (Fig. 5i) and the maxi-
mum likelihood of a damaged stem resprouting
(Fig. 5j) strongly inuenced tree dominance. Stem
growth rates of more than 50 cm year 1 are needed
for trees to persist, while stem growth rates greater
than 70 cm year 1 lead to tree dominance. The pau-
city of existing data on stem growth rates and the
sensitivity of the model to this parameter provides
motivation for collecting stem growth rate data.
Similarly, a probability of resprouting in excess of
0.6 is needed for tree persistence, while a probability
of resprouting of 0.99 leads to tree dominance. Data
suggest that resprouting probabilities in savanna are
typically greater than 0.8 in savanna (Trollope 1974,
1998; Boo et al. 1997; Gignoux et al. 1997). The rate
of adult mortality due to factors other than re is
another factor that strongly inuences tree persis-
tence; the model suggested that low adult mortality
rates (< 0.05) are necessary for tree persistence
(Fig. 5k). In apparent contradiction to this model
prediction, annual mortality rates, which may
include the eects of re, of c. 0.040.05 have been
reported in southern African savannas (Trapnell
1959; Shackleton 1997). However, Andersen et al.
(1998) reported mortality rates of 0.01 for
Australian savannas, and Dublin et al. (1990) used
0.01 in a model based on eld data from East
Africa. Interestingly, elephants have been responsi-
ble for tree morality rates of 0.18 in Zimbabwean
savannas (Thomson 1975); suggesting that their role
as ecosystem modiers should not be disregarded.
Increasing grass production negatively inuences
tree density (Fig. 5l) by making it more dicult for
seedlings to establish, and by eectively increasing
the re intensity and therefore reducing escape
opportunities into re-resistant size classes. The
grazing rate has the opposite eect (Fig. 5m), in gen-
eral agreement with observations that high grazing
rates promote bush encroachment (Archer et al.
1988; Skarpe 1991; Archer 1995). Our model there-
fore suggests that bush encroachment occurs due to
increased tree recruitment caused by reductions in
standing crop and hence re intensity. This contra-
dicts the competitive release mechanism of bush
226 encroachment (Walker & Noy-Meir 1982; Stuart-
Grasstree Hill & Tainton 1989; Jeltsch et al. 1997), whereby
coexistence in the decreased grass standing crop as a result of graz-
savanna ing reduces competition between grasses and trees
and thus increases opportunities for tree recruit-
ment. Recent empirical studies also challenge the
competitive release mechanism by showing that, in
resource-limited systems, establishment and recruit-
ment are limited more by resource availability than
competition (Davis et al. 1998). In our model, grass
standing crop had only a weak direct eect on tree
recruitment (Fig. 5f), i.e. the eect of grazing on
trees is manifested through the eect of grazing on
grass standing crop and hence re intensities. Our
model's mechanism is consistent with the hypothesis
that bush encroachment is constrained by soil moist-
ure availability and re intensity rather than grass
competition for soil moisture (du Toit 1967;
Harrington 1991). In addition, heavy grazing can
favour less ammable and less productive grasses,
further decreasing re intensities (Trollope 1998).
Conclusions
The rooting niche separation (Walker & Noy-Meir
1982) model of grass tree interaction predicts an
equilibrium coexistence between grasses and trees.
Dissatisfaction with the assumptions of the Walker
Noy-Meir model (Scholes & Archer 1997) has moti-
vated the search for alternative mechanisms of
grasstree coexistence (Menaut et al. 1990;
Hochberg et al. 1994; Jeltsch et al. 1996, 1998).
Models presented by Menaut et al. (1990), Hochberg
et al. (1994) and Jeltsch et al. (1996) could not pre-
dict coexistence, although Jeltsch et al. (1996)
reported coexistence for a narrow range of condi-
tions. In a revised model, Jeltsch et al. (1998) intro-
duced safe sites for seedling establishment by
simulating the eects of a range of small-scale het-
erogeneities, and these revisions allowed grasstree
coexistence. Our model, and the data used to para-
meterize the model, also demonstrates grasstree
coexistence and shows that it can occur for a wide
range of conditions.
Although many of the post WalkerNoy-Meir
models included re, they tended to concentrate on
the eects of re frequency and re distribution
rather than re intensity and its variance. We there-
fore believe that the novel feature of our model that
promotes coexistence, is its simulation of the eects
of re intensity on tree recruitment; further, by
including tree resprouting in our model, the role of
re is to limit tree recruitment, allowing adult mor-
tality to remain low. Perhaps more importantly, the
model simulates the eects of variation in re inten-
sity as inuenced by variations in grass production,
# 2000 British
grazing and tree neighbourhoods. In addition to
Ecological Society
Journal of Ecology, variations in re intensity, the model also simulates
88, 213229 how variable rainfall could result in variations in
seedling establishment. We predict that rainfall-dri-
ven variation in recruitment is more important in
arid savannas, where res are less intense and more
infrequent. In summary, it is variations in rainfall
and re intensity that lead to variations in seedling
establishment and tree recruitment that, against a
background of low levels of adult mortality, allow
the storage eect (Warner & Chesson 1985) to pro-
mote coexistence. Hence our hypothesis is that
grasstree coexistence is driven by the limited
opportunities for tree seedlings to escape both
drought and the ame zone into the adult stage.
Our model emphasizes temporal variance in recruit-
ment opportunities, while Jeltsch et al. (1998)
emphasized spatial variation in opportunities for
recruitment. Hence we suspect that the storage eect
may also be mediating grasstree coexistence in the
model developed by Jeltsch et al. (1998), although
they did not interpret their results in the context of
the storage eect.
While we have demonstrated that coexistence
between grasses and trees can occur for a wide
range of parameter values, how much environmental
and geographical space this translates into needs to
be explored; this could be done by using the model
to guide the collection of data from a range of
savanna ecosystems. By parameterizing the model
for a range of sites we will then be able to test
whether the patterns produced by the model are
consistent with the patterns observed in the eld,
and the kinds of savanna for which the model is
appropriate. What is clear is that the answer to the
`savanna problem' (Sarmiento 1984) lies in stepping
back from the details of ne-scale interactions
between grasses and trees and observing the longer
term eects of disturbance, life history (Noble &
Slatyer 1980) and regeneration (Grubb 1977) on
demography.
Acknowledgements
Thanks to Harry Biggs, Mary Cadenasso, Neil
Eccles, Jessica Kemper, Henri Laurie, Jeremy
Midgely, Norman Owen-Smith, Kevin Rogers and
Ed Witkowski for stimulating discussions on the
ideas presented here. Thanks to Andre Potgieter and
Nick Zambatis of the National Parks Board for
allowing us to use unpublished data. This work is a
contribution to the riparian corridors in savanna
landscapes programme. The support of the Andrew
Mellon foundation is gratefully acknowledged.
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