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Journal of

Ecology 2000,
Fire, resprouting and variability: a recipe for grasstree
88, 213229 coexistence in savanna
*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

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

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

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 mean (Rx; mm) and standard deviation (Rsd; mm) of
do not model individual grass tufts but model grass annual rainfall; s is the eect of the long-term peri-
patches. Because we are interested in capturing the odicity of rainfall (mm); l is the frequency of peri-
heterogeneity introduced by tree neighbourhoods on odicity; and y is the simulation year.
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 Most authors use linear regression to describe the
postulate that interannual variation in rainfall is the relationship between rainfall and grass production,
key source of variation, and because most of the as this produces the best t to the data from
data we had access to was annual data. However, as savanna regions (O'Connor 1985; Scholes & Walker
discussed above, fuel properties vary considerably 1993). Grass production can be written as:
within a year; we deal with this problem by allowing
the day of re ignition to be a random variable of Gp gg R eqn 2
dened moments. Below we discuss the assumptions
where Gp is the predicted above-ground production
and the functions used to simulate the ecological
(kg ha 1), R is the annual rainfall (mm), and gg is
processes described above; we then describe how we
the growth coecient. Using data from southern
combine these functions to dene a dynamic model.
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
Mean annual rainfall is variable in savanna systems; & Coughenour 1990). Grass production beneath
this variability can be divided into two components. tree canopies can be boosted by almost 300% or
The rst component is stochastic variation; the sec- suppressed by over 50% (Mordelet & Menaut
ond component is long-term periodicity. We use a 1995). To account for the eect of stem neighbour-
sine wave function, which captures both these com- hoods on grass production we can write:
ponents of variability, to generate rainfall:

gg R ; if Ci;j 0
 y Gpi;j eqn 3
R xRx ; Rsd sin 2p s; Re0 eqn 1 gg Rc ; if Ci;j 1
# 2000 British l
Ecological Society
Journal of Ecology, Here R is the annual rainfall (mm); x is a nor- Here c is a coecient that describes the eect of
88, 213229 mally distributed random number dened by the the stem neighbourhood on grass production at site
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 amount of grass carried over from one year to the
site (i,j) is in a stem neighbourhood (C(i,j) 1) or next (Gy 1(i,j)), we could write:
not (C(i,j) 0); stem neighbourhoods are dened

below (equation 12). The grass standing crop at the Gi;j atd utd ; if Bi;j 0
end of the growing season (G(i,j)) is therefore: Gy1i;j
Gi;j atd utd bGi;j ; if Bi;j 1

Gi;j Gpi;j Gy1i;j eqn 4 eqn 6

where Gy 1(i,j) is the amount of grass material that is Here u is the decomposition rate (kg ha 1 day 1);
carried over from the previous year (cf. equation 6). td is the length of the dry season in days; b is the
The levels of herbivory, the rate of grass decomposi- completeness of the burn; and B(i,j) is an array that
tion and whether a re has occurred will determine describes whether site (i,j) is burnt or not.
how much grass is carried over from one year to the


HERBIVORY AND DECOMPOSITION The moisture content of the fuel inuences re

While herbivores are a ubiquitous feature of intensity. Grass growing in a tree's neighbourhood
savanna ecosystems and inuence savanna dynamics may be moister and retain moisture levels for longer
in many ways (Cumming 1982; Pellew 1983; Dublin into the dry season (Weltzin & Coughenour 1990;
et al. 1990; Prins & van der Jeugd 1993; Scholes & Vetaas 1992; Webber 1997). If we assume that the
Walker 1993), we are primarily interested in herbi- moisture content of grass decays exponentially into
vores' ability to manipulate fuel loads; for this rea- the dry season (Cheney & Sullivan 1997), we could
son we do not consider browsing. Following write the moisture of grass on the day of ignition as:
Danckwerts (1982) we assume that herbivores can 
reduce the grass standing crop as a linear function Mo expdo ti ; if Ci;j 0
Mf i;j eqn 7
of time since production: Mc expdc ti ; if Ci;j 1

Gfi;j Gi;j ati eqn 5

Here Mf(i,j) is the moisture content (%) at location
where Gf(i,j) is the grass standing crop on the day of (i,j) ti days after the start of the dry season; Mo and
ignition, ti is the ignition day (days since the start of Mc are the moisture contents of grass (outside and
the dry season), a is the grazing rate (kg ha 1 inside the tree neighbourhoods) at the end of the
day 1), and G(i,j) is the grass standing crop at the growing season; and do and dc are the drying rates
end of the growing season. Note that this function for outside and inside the tree neighbourhood. The
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implies that growth and consumption are treated as array C(i,j) records if a cell is inside a tree neighbour-
Journal of Ecology, discrete events in the model, and ignores spatial and hood (see equation 12 for the denition of tree
88, 213229 temporal heterogeneity in grazing. To estimate the neighbourhood).
We use empirically derived relationships to predict Stems that have been topkilled usually resprout
coexistence in
re intensity, as these statistical models provide, from rootstocks. Savanna species have very high
given the information available, a better prediction probabilities of resprouting (Lacey et al. 1982;
of re intensity than physical re models (Trollope Trollope 1982, 1984; Boo et al. 1997; Gignoux et al.
1998). The statistical model was developed using 1997). Resprouting ability is generally thought to
200 monitored res in South African savannas (P increase with stem size (Wright et al. 1976; Moreno
< 0.01, d.f. 196, R2 0.60; Trollope 1998). The & Oechel 1993) but, in some tree species, decreases
model was tested against independent re behaviour again in the larger size classes (Trollope 1974;
data and accounted for 56% of the variation in re Hodgkinson 1998; K. Maze and W.J. Bond, unpub-
intensity (Trollope 1998). The multiple regression lished data). The eect of stem size on the probabil-
model is: ity of resprouting ( pr) can be written as:

p 8
> 0 ; if h<hr
Qi;j 2729 0:8684Gfi;j 530 Mfi;j >
< pmax
pr   eqn 10
596 >
> h h0:5 ; if hehr
0:907H 2 eqn 8 : 1 exp
W vr

where pmax is the maximum probability of resprout-

Here Q(i,j) is the re intensity (kJ s 1 m 1) at
ing, h0.5 is the stem height (h) at which there is a
site (i,j); Gf(i,j) is grass standing crop (kg ha 1) at
50% chance of resprouting, and vr is a constant that
site (i,j); Mf(i,j) is fuel moisture (%) at site (i,j); H is
describes how rapidly the probability of resprouting
the relative humidity (%); and W is wind speed (m
changes with stem height. Recent germinants do not
s 1). The empirical model therefore proposes that
have the root reserves to resprout (Moreno &
the realized re intensity is dependent on the grass
Oechel 1993); we therefore assume that stems less
standing crop, the grass moisture content, the rela-
than the resprouting height (hr) cannot resprout.
tive humidity, and the wind speed on the day of the
The model only allows one stem per site; it therefore
re. For the simulations we assume that humidity
does not consider the resprouting of multiple stems.
and wind speed are normally distributed random
Because the probability of re survival is inuence
numbers dened by the site's mean and standard
by tree height and not stem number, we do not con-
deviation of humidity (Hx,Hsd ;H r 0) and wind
sider this an important limitation.
speed (Wx,Wsd ;W r 0).

Recent carbon dating evidence suggests that
The probability of stem mortality (or `topkill') in a savanna trees can be more than 1000 years old
re is a function of stem height and re intensity (CSIR, personal communication). Rates of mortality
(Trollope 1984). We used data on the survival rates due to stress are therefore expected to be low.
of 7400 stems of 76 species in 40 res of known Andersen et al. (1998) reported annual mortality
intensities (W.S.W. Trollope, A.L.F. Potgieter and rates of 0.01 for Australian savannas; Trapnell
N. Zambatis, unpublished data) to estimate a logis- (1959) reported mortality rates of 0.04 in
tic regression model of the probability of stem mor- Zimbabwean miombo woodlands; and Shackleton
tality (P < 0.01, R2 0.48, d.f. 7397): (1997) reported mortality rates of 0.05 in South
African savannas. In the model trees of maximum
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 The stem mortality and stem resprouting functions
m 1; equation 8). Interestingly Williams et al. all use stem height information. Very little data exist
(1999), using data from a single intense re in a tro- on height or diameter growth of savanna stems. We
pical savanna in northern Australia, found that pt know that stems initially grow rapidly in height but
was a quadratic function of tree size, with larger subsequently growth slows (K. Maze and W.J.
# 2000 British
Ecological Society
and small trees suering highest pt. The susceptibil- Bond, unpublished data); using this information we
Journal of Ecology, ity of large trees to topkill was attributed to termite can describe stem growth using a dierence equa-
88, 213229 damage to large trees (Williams et al. 1999). tion:
219 hy1 long-distance dispersal. The importance of local vs.
h hy1 1 gs eqn 11
S.I. Higgins, hmax long-distance dispersal motivated us to model dis-
W.J. Bond & where gs is the growth rate of stems (cm year 1), persal as a stratied process that explicitly considers
W.S.W. Trollope hmax is the maximum stem height (m), and hy1 is both local and long-distance dispersal (Higgins &
the stem height in the previous year. Stem growth Richardson 1999). We can use a mixture of two
rates and maximum tree heights in savannas are exponential distributions to describe a probability
known to be inuenced by moisture and nutrient density function of dispersal distances d(x):
availability (Shackleton 1997), although we do not
dx pl exp bl x 1 pl exp bf x eqn 14
explicitly consider these eects here.
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-
A stem's neighbourhood has two components. First, meter bf).
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- Seed banks of savanna tree species are not very
theless, that the diameter of the neighbourhood (n; long-lived. Those that do not suer predation by
m) increases as a linear function of tree size: insects and rodents either decay rapidly, germinate
or lose viability (Tybirk et al. 1993). We summarize
n k gn h eqn 12 all these processes by assuming that a constant pro-
where gn is a growth coecient, h is stem height portion of seeds decay each year (sd). We could not
(m), and k is a constant. We do not have data on nd any published estimates of sd, although anecdo-
below-ground neighbourhoods and we therefore use tal evidence suggests that the decay rate is relatively
data on the relationships between stem height and high (Skoglund 1992; Tybirk et al. 1993; Miller
canopy diameter (W.S.W. Trollope and A.L.F. 1994; Chidumayo & Frost 1996).
Potgieter, unpublished data) to estimate k and gn.


SEED PRODUCTION Little is known about the regeneration niches of

savanna tree species, although we know that many
Following Ribbens et al. (1994), we dene seed pro-
seedlings are shade-intolerant and that high grass
duction as a function of tree size, such that:
biomass can suppress their recruitment, while seed-
8 lings of other species are shade-tolerant and little
< 0 ; if h<hf
 2 inuenced by grass biomass. What is clear is that
F h eqn 13
:f ; if hehf one of the major factors limiting establishment is
hstd the availability of moisture: droughts during the wet
season of more than 30 days can lead to seedling
Here F is tree fecundity (seeds year 1) and f is the mortality (Medina & Silva 1990; Chidumayo &
number of seeds dispersed by a tree of a reference Frost 1996; Homan 1996). If we assume that the
height (hstd); stems smaller than the height of repro- number of rainfall events during the wet season is
ductive maturity (hf) do not produce seeds. Data on positively correlated with annual rainfall, then the
the seed production of savanna trees (Tybirk et al. probability of wet season drought should decrease
1993) is used to estimate hf and f. with annual rainfall. We can express the probability
of wet season drought ( pd) as:

pd   eqn 15
1 exp
Many savanna tree species are dispersed passively
and by animals (bird, ungulates, rodents, termites Here R0.5 is the annual rainfall at which there is a
and ants; Brown & Archer 1989; Tybirk et al. 1993; 0.5 chance of a wet season drought; and vd is a con-
Miller 1994). Previous demographic models of stant that describes the rate at which the probability
# 2000 British
Ecological Society
savanna (Menaut et al. 1990; Hochberg et al. 1994; of wet season drought changes with rainfall (R). If
Journal of Ecology, Jeltsch et al. 1996, 1998) have emphasized the impor- there is no wet season drought then the probability
88, 213229 tance of tree clumps and hence the role of local vs. of establishment ( pe) is:
220 >
> 0 ; if fCi;j 1 & ct 1g eqn 16.1
Grasstree 1
pei;j   eqn 16.2
coexistence in >
> G i;j G0:5 ; if fCi;j 0 & ct 1g
: 1 exp
savanna ve or {ct = 0}

Here we account for the observations that some (1994). We assume that res can spread if a thresh-
species require light for establishment (ct 1), while old re intensity is exceeded; this threshold has been
others are shade tolerant (ct 0). If the light condi- estimated as 150 kJ s 1 m 1 in savanna systems
tions are suitable (equation 16.2) then the probabil- (van Wilgen & Scholes 1997). The re spread algo-
ity of establishment is a function of the grass rithm allows a re to spread to neighbouring cells if
standing crop. In equation 16.2 G0.5 is the grass a neighbouring cell's potential re intensity exceeds
standing crop at which the probability of establish- the threshold. The re spread algorithm is not inu-
ment ( pe) is 0.5, G(i,j) is the grass standing crop and enced by wind or topography. The way we model
ve is the rate at which the probability of establish- re ensures that fuel properties rather than ignition
ment changes with grass standing crop. 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-
IMPLEMENTATION tion 9). The likelihood of a dead stem resprouting is
Each year the model sequentially simulates the fol- estimated as a function of stem height (equation 10).
lowing ecological processes: rainfall, tree growth, Adult trees face a pm probability of death each year.
seed dispersal, grass production, potential re inten- Seeds can only germinate and establish if there is no
sity and re spread, stem mortality due to res, wet season drought (equation 15). If there is no
resprouting, adult tree mortality, seedling establish- drought then a seed can establish, provided the
ment, and seed bank decay. Rainfall is generated grass biomass is low enough, light conditions are
using equation 1. Tree heights are incremented each suitable (equation 16) and the site is unoccupied by
year (equation 11) and tree neighbourhoods are sub- a tree stem. The seed bank is decayed by a constant
sequently calculated from the tree height (equation proportion (sd) each year.
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 Model behaviour
the distance each seed moves is a random number
dened by the mixture distribution (equation 14). Our analysis of the model's behaviour is divided
This distance and a randomly selected direction are into two sections. We rst examine whether the
used to calculate the location of each seed. Grass model successfully predicts coexistence, and attempt
production is calculated as a function of the rainfall to understand the behaviour of the model in the
and neighbourhood state (equation 3). Estimating context of theoretical coexistence models. The sec-
potential re intensity (equation 8) requires the esti- ond section analyses the sensitivity of the coexis-
mation of grass standing crop, grass moisture con- tence state to key parameters and hence investigates
tent, relative humidity and wind speed on the day of the environmental conditions for which we would
the re. We assume that ignition can occur on any expect grasstree coexistence.
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 For this section we initialized the model with the
of the re can be estimated by using equations 5 best parameter estimates available. Sources of these,
and 7 and by assuming that ignition occurs ti days largely southern African, parameter estimates are
into the dry season. In the current version of the discussed under operational denition, and the para-
model one re ignition occurs per year. We assume meter values used are listed in Table 1. We then var-
that ti is a normally distributed random number ied the base parameterization (Table 1) to simulate
with a mean (Ix) and standard deviation (Isd) char- four sites representing a rainfall gradient from arid
acteristic of the temporal distribution of ignition to mesic savanna (Table 2). In southern Africa this
events. This method of generating ignition events gradient is associated with a change from palatable
emphasizes the temporal distribution of ignition to relatively unpalatable grass; and we simulate this
# 2000 British
Ecological Society
events, i.e. there is an emphasis on re intensity by decreasing the grazing rate with increasing rain-
Journal of Ecology, rather than re frequency. The re spread algorithm fall. We also assume that (i) stem growth rates will
88, 213229 is analogous to that proposed by Turner & Romme 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
coexistence in

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

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