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ECOSYSTEMS

Ecosystems (2002) 5: 329 338


DOI: 10.1007/s10021-001-0077-1

2002 Springer-Verlag

Contagious Disturbance, Ecological


Memory, and the Emergence of
Landscape Pattern
Garry D. Peterson
Center for Limnology, University of Wisconsin, Madison, Wisconsin, 53706, USA

ABSTRACT
Landscapes are strongly shaped by the degree of
interaction between pattern and process. This paper
examines how ecological memory, the degree to
which an ecological process is shaped by its past
modifications of a landscape, influences landscape
dynamics. I use a simulation model to examine how
ecological memory shapes the landscape dynamics
produced by the interaction of vegetative regrowth
and fire. The model illustrated that increased ecological memory increased the strength and spatial
extent of landscape pattern. The extent of these
changes depended upon the relative rates of vegetative recovery and fire initiation. When ecological
memory is strong, landscape pattern is persistent;

pattern tends to be maintained rather than destroyed by fire. The generality of the simulation
model suggests that these results may also apply to
disturbance processes other than fire. The existence
of ecological memory in ecosystems may allow processes to produce ecological pattern that can entrain
other ecosystem variables. The methods presented
in this paper to analyze pattern in model ecosystems could be used to detect such pattern in actual
ecosystems.

INTRODUCTION

previous fires, then memory is shaping their dynamics. The presence of memory allows ecological
processes to interact with one another; its absence
means that pattern is generated by a single process
imposing some type of template of organization on
a system. However, in most ecological systems, ecological processes cannot be neatly divided between
those that exhibit memory and those that do not.
Even apparently one-way relationships, such as the
effects of climate on vegetation, contain interactions (for example, the effects of vegetation upon
humidity). How much does memory influence
landscape dynamics? In this paper, I use a simulation model of fire and vegetation dynamics to examine the role of ecological memory in producing
landscape pattern. Specifically, I examine how ecological memory influences the landscape pattern
that is produced by the interaction between fire
frequency and vegetative regrowth.
Fire is a key structuring processes in many eco-

Key words: landscape ecology; self-organization;


spatial dynamics; patch dynamics; keystone processes; scale; autocorrelation.

Landscape pattern is a key attribute of ecosystems,


organizing and regulating the flow of ecological
goods and services (Ludwig and others 2000).
Landscape pattern is itself shaped by ecological processes. The extent of the interaction between landscape pattern and the processes that shape it has a
fundamental influence on landscape dynamics.
Does a process impose a pattern upon a landscape
unilaterally, or is there a two-way dynamic interaction between pattern and process? The degree to
which an ecological process is shaped by its history
can be thought of as the strength of the ecological
memory of that process. For example, if the location of tree-fall gaps is influenced by where past
tree falls occurred or if fire spread is influenced by

Received 14 November 2000; accepted 21 September 2001.


e-mail: gdpeterson@facstaff.wisc.edu

329

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G. D. Peterson

systems (Bond and van Wilgen 1996). Although the


duration of a fire is much shorter than the life span
of the vegetation it consumes, fire produces landscape patterns that persist for long periods (Baker
1993). At small spatial scales, fire homogenizes the
landscape by killing aboveground growth of trees,
producing patches of even-aged vegetation. At
larger spatial scales, fires produce heterogeneity by
creating a mosaic of burned and unburned patches.
Although fire produces landscape pattern, landscape pattern (for example, the presence of fire
breaks) can also influence fire dynamics (Baker
1995). Researchers working in different locations
have variously proposed that the probability of fire
spread is either independent of time since fire
(Bessie and Johnson 1995), or that it increases with
time since fire (Minnich 1983). These differences
suggest that some forests have little ecological
memory, whereas others have a significant
amount.
Fire is representative of a larger group of contagious disturbance processes. Contagious disturbance processeswhich include fires, insect outbreaks, and grazing herbivoresspread themselves
across a landscape. Unlike noncontagious disturbances, such as ice storms, hurricanes, or clearcutting, the extent and duration of a contagious
disturbance event are dynamically determined by
the interaction of the disturbance with a landscape.
The size of a contagious disturbance depends, at
least partially, on the spatial configuration of landscape being disturbed. Changes in landscape pattern
will alter the nature of a contagious disturbance
regime, but will not alter a noncontagious disturbance regime. For example, fragmentation of a forest by roads will impede the spread of a wildfire, but
not determine the path of a hurricane. A consequence of this interactivity is that the same driving
forces will produce different contagious disturbance
behaviors in landscapes with different spatial patterns.
I used a simulation model to investigate this
question because of the difficulty of taking an empirical approach. Disentangling the cross-scale interactions in heterogeneous real ecosystems is a
difficult task that is compounded by the fact that
spatially explicit data that could be used to assess
ecological pattern have only recently become available. Assessing the dynamics of landscape pattern
requires spatially explicit data that span at least
several cycles of structuring ecological processes.
Fire return times, as well as return times of other
contagious disturbance processes such as insect outbreaks, usually range from decades to centuries.
Long-term, spatially explicit records that span hun-

dreds of years do not, to the best of my knowledge,


currently exist. A modeling approach allows ecological memory to be analyzed in the absence of
confounding factors. Although such analyses cannot reveal what happens in real ecosystems, they
can suggest where and when theory predicts events
should be observable and unobservable in real ecosystems. It is in this sprit that I use simulation
models to identify ecological situations in which
ecological memory can be expected to have a large
effect.

FOREST FIRE MODEL


Ecologists have developed a variety of models of
forest fire dynamics. These models range from fire
risk assessment models that predict the detailed
dynamics of fire intensity and spread (Andrews
1986; Finney 1998) to spatially implicit models of
the interaction between fire frequency and vegetation dynamics (Casagrandi and Rinaldi 1999). Detailed spatial models such as the EMBYR model of
Yellowstone (Hargrove and others 2000) include
details of vegetation type, wind, and long-distance
fire dispersal via spotting. Spatial models that do not
model fire spread explicitly have also been used to
examine the distribution of forest age classes (Boychuk and others 1997).
Because I am interested in understanding how
ecological memory shapes landscape pattern, I created a simple spatial model, based on a minimal
model of forest fire dynamics developed and analyzed in statistical physics literature (Clar and others
1996; Drossel and Schwabl 1992). This model represents a forest as a rectangular matrix of sites. Each
site is either empty, occupied by trees, or occupied
by burning trees. In this model, a forest fire regime
is defined by three rates: the rate at which trees
regrow into empty sites, the rate at which burning
trees ignite neighboring nonburning trees, and the
rate/area at which fires are ignited across the landscape. To produce a fire regime in which fire disturbs rather than interacts with forest regrowth,
these rates must operate at different scales (Drossel
1997).
I developed a modified version of this model that
allowed me to examine the effect of ecological
memory. My model assumes that a site does not
always burn when fire burns neighboring sites;
rather, each site has a probability of burning that
depends on the time since the site was last burned.
This assumption supposes that after a fire there is a
process of fuel accumulation at a site. For example,
if fuel accumulates slowly, then a fire will be un-

Disturbance, Landscape Pattern, and Memory

331

Figure 1. An illustration of
the functioning of the fire
model. During each year, a
number of fires are initiated
at random locations. After a
fire is initiated at a site, it
can spread to any of its eight
neighboring sites. Fire spread
into a site is a probabilistic
function of the time since a
site last burned. Fire spreads
until there are no burning
sites. When a fire is extinguished, either another fire is
initiated or the landscape
ages a year and another
round of fires are initiated.

likely to spread into recently burned sites due to the


absence of fuel.
In my model, a fire regime is defined by the rate
of fire ignition, the rate of vegetation recovery following a fire, and the rate of fire spread. These
relationships can be thought of as the interactions
of different scale processes, such as climate and
vegetation growth, interacting to define a fire regime. Ecological memory is captured in the relationship between vegetation recovery and the probability of fire spread. Rather than explicitly
representing the speed of fire spread, my model
assumes that fire spreads much faster than vegetation recovers from fire. Drossel (1997) has shown
that in the statistical physics forest fire model, if fire
spreads much faster than a forest recovers from fire,
the average fire size is proportional to the rate of
recovery divided by the rate of ignition. This relationship means that as fire ignition events become
less frequent relative to the rate of recovery, fires
will tend to become larger and vice versa. Climate
and topographic variation influence the probability
of fire spread across time and space. Because this
investigation focuses upon the relationship between regeneration and fire, I hold climate and
topography constant.

Model Structure
My model is spatially explicit. The basic functioning
of the model is illustrated in Figure 1. The model
divides a landscape into a matrix of sites. The be-

havior produced by this model is general; its applicability to a specific type of disturbance depends on
the degree to which the grain and extent of that
disturbance can be captured by this model. The
modeled landscape is 440 440 sites. The model is
roughly parameterized so that each site in the matrix represents an area of vegetation 50 m on edge
(0.25 ha). Each site is described by the time since
that site was last burned, which affects the probability of fire spread.
The dynamics of the model consist of a withinyear process of fire initiation and spread, and succession. The entire process of fire initiation and
spread occurs within a simulated year. Randomly
selected sites in the landscape are ignited by fires at
the fixed rate. I used three fire initiation rates: 1
fire/100 cells/y, 4 fires/100 cells/y, and 16 fires/100
cells/y. The probability of fire spreading from a
burning site into a neighboring nonburning site is
modeled as a monotonically increasing function of
the nonburning sites time since fire (TSF). A fire
may spread from a site to any of its eight adjacent
sites, provided they have not already burned that
year. A burning site has only one chance to ignite
each of its neighboring sites. A fire will continue to
spread across the landscape until it fails to spread to
any unburned cells. Following the extinction of a
years fires, vegetation regrows at sites across the
landscape. The combination of fire and vegetation
regrowth produces a landscape composed of a mosaic of differently aged patches (Figure 2).

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G. D. Peterson

Figure 2. A pattern of firegenerated patches emerges


from a random landscape
after a history of fires. (A)
Forest after 1 year of fires.
(B) Forest after 300 years of
fires.

To simplify the comparison of systems exhibiting


different amounts of ecological memory, I assumed
that the probability of fire spread plateaus at a high
value (Pmax) following a maximum time since fire
(TPmax). To allow the strength of ecological memory
to be varied using a single parameter (), I designed
a mathematical function:
PrFireSpreadTSF 1 P max TPmax 1, TSF
TSF

TP max
PrFireSpreadTSF P max

,TSF TP max
(1)

The ecological memory parameter, , varies the


shape of the relationship between time since fire
and the probability of fire spread. I found that variation in ecological memory, ranging from no memory to strong memory, was well captured by using
values of 1/100, 1/4, 1/2, 1, 4, 100.
The relationships between time since fire of a site
and the probability of fire spreading into such a site
are shown in Figure 3. The figure illustrates that
when 1/100, fire spread is independent of time
since fire; as increases, the effect of fire becomes
more pronounced. When 1, there is an approximately linear relationship between time since fire
and the probability of fire spread; when 100,
there is approximately a step function between not
burning and Pmax when the time since fire equals
TPmax.
The time period over which vegetation reaches
maximum ability to carry fire (TPmax) varies greatly
in different ecosystems. Because my model works at
a yearly time step, I had to choose a value of TPmax
that permitted significant differences between the
fire spread functions produced by different values of
. I examine fire recovery periods (TPmax) of 12, 25,
and 50 years. These values span a reasonable eco-

Figure 3. Relationships between time since fire and the


probability of fire spread from Eq. (1). Vegetations memory of past fires is controlled by . If is low, the vegetation has no memory of past fires, and fire spreads
independently of vegetation structure. At high values of
, vegetation has a strong influence on fire spread, and
fire spread becomes dependent on vegetative pattern.

logical range, yet are brief enough in duration to


allow the tractable analysis of multiple fire periods.
The maximum probability of fire spreading into
an unburned neighboring site is Pmax. I choose a
value of Pmax located in the center of the percolation transition in this model. The percolation
threshold is the value of the probability of fire
spread at which a fire will percolate (Stauffer and
Aharony 1994); that is, fire will be able to spread
from one side of an arbitrarily large landscape to
another. A fire that percolates across a landscape
will not burn the entire landscape; some patches
will escape fire. In this model, percolation depends
on the details of how fire spreads (Stauffer and
Aharony 1994). Fire can spread to any of the eight
neighbors of a burning site, provided they have not
previously burned. I calculated the probability of
percolation, in homogenous landscapes of Pmax, for
different values of Pmax. I chose Pmax 0.24, a value
at which fire will frequently, but not always, percolate across the landscape. This value is ecologically reasonable because it probabilistically allows

Disturbance, Landscape Pattern, and Memory


some fires to spread far while limiting the spread of
others. Higher or lower values would assume that
fire behavior tends to be exclusively either large
rapidly spreading fires or small gradually spreading
fires, respectively. Similar values of Pmax (0.22 0.25)
do not change the general behavior of the model.

METHODS
To assess the relationship between contagious disturbance and ecological memory, I conducted multiple runs of the forest fire model for each of a range
of different amounts of ecological memory (), different rates of recovery following fire (TPmax), and
different fire frequencies. For each set of model
runs, I compared the cross-scale vegetation pattern
using correlograms.
Correlograms measure autocorrelation among
points on a landscape by calculating correlation
between pairs of points that are separated by different lags (Legendre and Fortin 1989; Radeloff and
others 2000; Rossi and others 1992). Specifically,
correlograms measure the autocorrelation between
two points separated by a lag for a series of n points.
Correlograms are similar to semivariograms, but
they are more robust to local changes in the means
and variances across a data set. I used a two-dimensional version of the correlogram to measure autocorrelation across both space and time.
I did this by sampling 6000 randomly selected
points (the 0.25-ha sites) that are separated by a
combination of temporal lags ranging from 0 to 84
years and spatial lags ranging from 0 to 200 cells.
These dimensions were chosen to include several
fire cycles while still remaining analytically tractable. In this case lag (h) is a vector that has both a
spatial and temporal component. Lag autocorrelation is estimated by:

z x m

Nh

1
p h
Nh

z x i h m h

i1

S hS h
(2)

where p(h) is an estimate of the autocorrelation at


lag h. z (xi) and z (xi h) are two data points separated by the lag h. N(h) is the number of data
points that are separated by lag h. Data point z (xi) is
the tail and z (xi h) is the head of a vector, and
mh and mh correspond to the mean values of the
points at the head and tail end of all the vectors of
lag h. sh and sh represent the standard deviations
of the tail and head values of the vectors at lag h.

333

Plotting lag autocorrelation against lag (h) produces


a correlogram.
Correlograms were constructed from the dynamics of a landscape 20 km on edge over 180 years.
The dimensions of this cube of data are such that
the maximum lags in space and time were less than
half the span of the landscape and less than half the
duration period during which landscapes were recorded.
No external disturbance enters the simulated
landscape from beyond the models edges, and fire
cannot spread beyond the edges of the simulated
area. These constraints produce an edge effect,
which results in the cells near the edges of the
simulated area burning less frequently than the
cells near the center of the matrix. The extent of the
area experiencing edge effects depends on the relative size of the simulated fires compared to the
total extent of the simulated area. The strength of
edge effects is inversely proportional to the amount
of memory in the model; however, edge effects are
typically limited to 10 20 cells. Consequently, correlograms were only calculated between points that
were at least 20 cells from the edge of the modeled
area.
In the temporal equivalent of an edge effect, initial landscape conditions influence landscape pattern for some period, but their influence declines
over time. To remove the effects of initial conditions, data for the correlograms were only collected
after allowing the landscape to organize for 600
years (at least 12 fire cycles). Based on test model
runs, this duration was roughly twice as long as
needed for a dynamic equilibrium to be reached.
I analyzed my model with different amounts of
ecological memory ( 1/100, 1/4, 1, 4, and 100),
with different rates of vegetation recovery following fire (TPmax 12, 25, and 50 years) and different
fire frequencies (1 fire/100 cells/y, 4 fires/100
cells/y, and 16 fires/100 cells/y). For each set of
model parameters, the model was run for each
value 26 times. The average autocorrelation at each
lag distance and time was calculated. Because the
standard error around a mean is a function of the
square root of the number of samples taken, the
benefit of increasing the number of model runs
diminishes rapidly as the number of samples increases. From these multiple model runs, standard
deviations of the average autocorrelation were estimated, and a two-tailed t-test was used to determine whether the autocorrelation at each lag was
significantly different from zero with a P 0.01
(Zar 1984). Autocorrelations at a lag that are not
significantly different from zero are plotted as zero.
From these calculations, autocorrelation surfaces

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G. D. Peterson

Figure 4. Covariance at spatial and temporal lags for 1/100, 1/4, 1, 4, and 100 at fire frequencies of 1, 4, and 16
fires/100 cells/y. Horizontal and vertical scales are log axes. Plus signs indicate positive autocorrelation; negative signs
indicate negative zones of autocorrelation. At long temporal lags and high values of , there are many cycles of positive
and negative autocorrelation, which are unlabeled.

were constructed. These surfaces show the autocorrelation of vegetation pattern (represented as time
since fire) at different spatial and temporal lags.
These figures show positive autocorrelation (that is,
sites are similar) at some temporal and spatial lags
and negative autocorrelation (that is, sites are consistently dissimilar) at other temporal and spatial
lags. The figures illustrate the spatial and temporal
lags at which pattern exists from the viewpoint of a
model cell.

RESULTS
The sets of correlograms produced by the models
are shown in Figures 4 and 5. In each figure, memory increases from left to right. Autocorrelation indicates that pattern exists at a given lag. Positive
autocorrelation indicates that sites at a given lag are
similar; negative autocorrelation indicates that sites
at a given lag are paired with sites that are different
(for example, young sites are paired with older sites

and vice versa). Figure 4 compares landscape pattern produced by models experiencing different fire
frequencies. Figure 5 compares landscape pattern
produced by models with different vegetation recovery periods. As suggested by the inverse relationship between forest recovery and fire initiation
(Drossel 1997), the patterns produced by changes in
fire frequency and forest recovery period are similar. Decreases in fire frequency have an effect similar, but not identical, to increases in the period of
forest recovery.
When there is no ecological memory (
1/100), spatial pattern is produced by fire, but that
pattern does not persist over time. There is a significant change in spatial and temporal pattern with
the transition from no memory to memory (
1/100 to 1/4). Spatial pattern spans less area
and persists longer. As ecological memory increases,
pattern begins to emerge at longer temporal lags,
which match the vegetation recovery period
(TPmax). When memory is strong, positive autocor-

Disturbance, Landscape Pattern, and Memory

335

Figure 5. Covariance at spatial and temporal lags for 1/100, 1/4, 1, 4, and 100 at vegetation recovery times (TPmax)
of 12, 25, and 50 years. Horizontal and vertical scales are log axes. Plus signs indicate positive autocorrelation; negative
signs indicate zones of negative autocorrelation. At long temporal lags and high values of , there are many cycles of
positive and negative autocorrelation, which are unlabeled.

relation appears at multiples of the vegetation recovery period. This increase in temporal structure is
also matched by an increase in spatial extent and
strength of spatial structure.
As ecological memory () increases, the spatial
and temporal organization of the landscape
changes. At all values of , the landscape is highly
correlated at short distances and over brief periods.
As increases, spatial correlation increases, and this
correlation decreases faster over time. Along with
this pattern, negative autocorrelation emerges at
intermediate temporal lags and positive autocorrelation at a frequency about that of the recovery
rate. With further increases of , the strength of
autocorrelation increases and extends across a
greater range of spatial lags. Additionally, the temporal lag over which there is correlation declines.
Furthermore, as increases, discrete ranges of negative and positive correlation emerge at temporal
lags related to TPmax at short spatial lags. At very
high values of , weak autocorrelation emerges at

intermediate frequencies at very large spatial lags.


This pattern is repeated at different fire frequencies
and values of TPmax; however, variation in fire frequency and TPmax changes the spatial and temporal
lags at which these zones of autocorrelation appear.
As frequency of fire decreases, the scale of spatial
correlation increases, and correlation over time increases. For example, at an intermediate value of
( 1), there is minimal organization at larger and
longer lags at a higher fire frequency (16 fires/100
cells/y); but as fire frequency decreases, the range
of spatial lags at which the landscape is correlated
and the intensity of correlation increases. Furthermore, at a lower fire frequency (1 fire/100 cells/y),
a new weak negative correlation emerges at about a
40-year lag.
At low , there is no difference between the
scales of correlation for different values of TPmax;
however, as increases, differences emerge. The
clearest change is that the temporal lags at which
positive and negative autocorrelation occur track

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G. D. Peterson

the changes in TPmax. When recovery time is brief


(12 years), autocorrelation decays quickly and then
reemerges rapidly; whereas if recovery is longer,
autocorrelation decreases gradually with temporal
lag and a new zone of autocorrelation emerges at
longer periods. As recovery time increases, the spatial scale over which autocorrelation occurs decreases and the strength of autocorrelation also decreases. This pattern is similar to what is observed as
fire frequency decreases.

DISCUSSION
The correlograms reveal that fires generate spatial
pattern and that the strength and spatial span of this
pattern depend on fire frequency and rates of vegetation recovery. They also reveal that the persistence of the patterns depends on ecological memorythe degree to which previous fires influence
the spread of fire. The amount of memory () necessary to produce landscape pattern depends on the
rate of vegetation recovery and the frequency of fire
initiation. As fire initiation rates decrease, or the
period of vegetation recovery decreases, the
amount of memory necessary to produce persistent
landscape pattern also decreases.
Ecological memory produces persistent pattern
because it establishes a feedback loop between fire
spread and landscape pattern. This feedback loop
allows ecological pattern and ecological process to
regulate one another (Watt 1947). In the absence of
memory, there is a one-way relationship between
fire and landscape pattern. Fire shapes landscape
pattern but is not shaped by it. Memory causes
landscape pattern to influence fire spread and vice
versa. Mutual influence between fire patterns and
landscape patterns encourages the formation of
mutually reinforcing patterns.
The feedback between fire spread and forest pattern produces a patch mosaic on the landscape.
Relatively homogenous patches of forest are produced due to the spatially contagious nature of fire
spread. Sites within these patches have the same
probability of fire spread, because they were previously burned at the same time. This homogeneity
means that fires ignited within these patches will
tend to either fail to spread or spread across the
entire patch. Fire spread tends to stop at the edge of
patches. Patches neighboring a burning patch are
more likely to have a lower probability of fire
spread, because patches with a higher probability of
fire spread are unlikely to have remained unburned. The mosaic of patches with different probabilities of fire spread reduces the ability of fire to
spread across the entire landscape. As memory in-

creases, these tendencies are strengthened, increasing the spatial extent of autocorrelation and the
persistence of spatial pattern over time.
The memory of past fires is the degree to which a
landscapes vegetative pattern influences the spread
of fire. The memory of past fires fades over time,
because differences in the probability of fire spread
that were produced by different fire histories diminish as the time since fire increases. For example,
when there is ecological memory, the probability of
fire spread varies between different-aged sites if
those sites are younger than the vegetation recovery period (TPmax), whereas there is no difference
between the probability of fire spread into sites
whose age is greater than TPmax. When forest pattern strongly influences the spread of fires (for example, when is greater than 1), forest pattern will
tend to be renewed by fire. When memory is weak
(for example, when is less than 1), future fires
will produce a new pattern that erases past patterns.

Implications of Ecological Memory


The consequences of manipulating landscape pattern or the processes that control the fire regime
will be more complex when an ecosystem has
memory than when it does not. Ecological memory
is encoded in the pattern of vegetation across the
landscape. This pattern constrains and channels the
spread of fire. Consequently, changes in vegetative
pattern, via abiotic, biotic, or anthropogenic processes, have the potential to alter a fire regime, even
if the drivers of that disturbance regime, such as the
climate and vegetation of the region, remain constant. Homogenizing the landscape may remove
barriers to fire spread, leading to a period of large
fires. Alternatively, fragmenting the landscape will
lead to a period of smaller fires. Despite these generalities, the degree to which a fire regime changes
in response to landscape modification will depend
on the specific details of landscape pattern and how
it is changed.
Similarly, due to the constraining effects of landscape pattern, shifts in either vegetation regrowth
or climate can also alter a fire regime in complex
ways. For example, a change in climate could increase fire frequency, which would produce smaller
and more frequent fires. Alternatively, a region invaded by pyrogenic grass will more rapidly become
combustible following a fire, decreasing the vegetation recovery time (DAntonio and Vitousek 1992).
The behavior of ecosystems with little memory will
simply track changes in vegetation recovery and fire
frequency. However, ecosystems with strong memory can be expected to respond to changes in driving processes in complex ways. Strong memory

Disturbance, Landscape Pattern, and Memory


should initially inhibit ecological response to
change. However, if change is rapid and intense,
extreme transient behavior can be expected. For
example, if fire frequency decreases, the existing
landscape pattern will constrain fires, but eventually the lack of maintenance of this pattern, due to
the reduced fire frequency, will result in very large
fires, before a new landscape structure is established. This suggests that ecosystems with large
amounts of memory will respond to changes in
disturbance regimes in more complicated ways than
ecosystems with little memory of the past.

Ecological Memory in Ecosystems


The general nature of the fire model suggests that
the interaction of ecological memory and contagious disturbances may influence landscape dynamics in a number of situations. In California
chaparral, fire intensity depends on vegetation density over small spatial scales (approximately 1 m),
while regeneration occurs primarily in areas that
experience less intense fire (Odion and Davis 2000).
In this ecosystem, the connection between vegetation pattern and fire introduces the memory of past
disturbances into the system and thus produces persistent spatial pattern. Ecosystems that are dominated by externally driven fluctuations in climate,
water levels, or biota are unlikely to be shaped by
the memory of past disturbances. For example, fire
in the western boreal forest of North America appears to be driven by climate variation (Johnson
1992), which suggests that ecological memory does
not influence the fire regime.
Animal behavior, like fire, has the potential to
encode memory into landscape pattern. Selective
grazing by herbivores produces disturbance patterns that are determined by existing landscape pattern. If the vegetative response to disturbance influences future grazing decisions by herbivores,
then the ecosystem exhibits memory and the interaction of selective herbivores can produce persistent
spatial pattern. For example, in the boreal forest of
eastern North America, moose selectively consume
early successional deciduous tree species. This
browsing facilitates the replacement of deciduous
species by coniferous species, producing a change in
vegetation pattern that alters future browsing decisions. This foraging behavior, in conjunction with
other contagious disturbance processes, produces
persistent spatial pattern in the forest (Pastor and
others 1998, 1999). In general, I expect that ecological memory is likely to arise in ecosystems in
which biotic variation has a strong influence on
ecological dynamics.

337

Extended Keystone Hypothesis


The results of my fire model show that ecological
memory encourages the emergence of persistent
spatial pattern, which provides some support for
the extended keystone hypothesis (Holling 1992)
and also suggests some limits. Holling (1992) proposed that landscape pattern is generated by the
interaction of a few keystone processes that operate at separate and distinct spatial and temporal
scales. He argues that these keystone processes entrain other ecological processes and variables to the
characteristic frequencies of these processes; consequently, the properties of ecosystems should exhibit
discrete rather than continuous structure. Although
Holling (1992) and others (Allen and others 1999;
Havlicek and Carpenter 2001; Manly 1996; Raffaelli
and others 2000) have examined the question of
whether ecosystem attributes exhibit a discontinuous structure, there has been little consideration of
how feasible it is for the interactions of keystone
ecological processes to produce discrete ecological
pattern.
Holling and others (1996) suggested that contagious disturbance processes, such as fire, are examples of keystone processes. My results show that
models of fire have the potential to produce discrete
landscape pattern, thus supporting the idea that fire
is indeed a keystone process. Persistent patterns
occur at temporal frequencies related to the recovery time of the vegetation from fire (that is, at 12-,
25-, and 50-year frequencies in the model). However, my model suggests that such patterns do not
emerge in the absence of ecological memory. These
results suggest that the extended keystone hypothesis is most likely to be demonstrated in ecosystems
that exhibit strong ecological memory ecosystems
in which the spread of a key disturbance process is
strongly influenced by the legacies of past disturbance.
The spatial and temporal correlograms that I used
to search for persistent pattern could also be used to
test the extended keystone hypothesis in actual
ecosystems. It is difficult to find data that span a
time period several times the frequency of a disturbance over a spatial scale several times the extent of
the disturbance. However, historical and paleoecological research is making such data sets increasingly available (Niklasson and Granstrom 2000).

CONCLUSIONS
Ecological memory shapes landscape pattern by increasing the strength of the interaction between
ecological processes and landscape pattern. When

338

G. D. Peterson

ecological memory is strong, landscape pattern is


persistent; pattern tends to be maintained rather
than destroyed by fire. The existence of ecological
memory in ecosystems may allow processes to produce ecological pattern that can entrain other ecosystem variables. Changes in landscape pattern or
key process drivers, such as fire frequency or vegetation recovery rates following disturbance, are
more likely to have nonlinear and surprising effects
in ecosystems with strong memory. There is some
evidence that ecological memory exists in real ecosystems. The methods presented in this paper to
analyze pattern in model ecosystems could be used
to detect evidence of persistent landscape pattern in
actual ecosystems.

ACKNOWLEDGMENTS
A part of this work was conducted while I was a
postdoctoral fellow at the National Center for Ecological Analysis and Synthesis at the University of
California Santa Barbara. It uses models that I developed during my doctoral research, which was
funded by a NASA Earth Science Fellowship. The
paper benefited from conversations with Bruce
Milne, and comments from Craig Allen, Elena Bennett, Lisa Dent, Tim Essington, and two anonymous
reviewers.

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