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2002 Springer-Verlag
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-
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G. D. Peterson
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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.
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
TP max
PrFireSpreadTSF P max
,TSF TP max
(1)
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)
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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-
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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
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G. D. Peterson
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.
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CONCLUSIONS
Ecological memory shapes landscape pattern by increasing the strength of the interaction between
ecological processes and landscape pattern. When
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G. D. Peterson
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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