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"The problem of pattern and scale is the central problem in ecology,

unifying population biology and ecosystems science, and marrying
Scale and Hierarchy basic and applied ecology. Applied challenges ... require the
interfacing of phenomena that occur on very different scales of space,

Theory time, and ecological organization. Furthermore, there is no single

natural scale at which ecological phenomena should be studied;
systems generally show characteristic variability on a range of spatial,
temporal, and organizational scales."

Simon Levin 1992

Temporal and Spatial Scales in Ecology

Theories of Scale
Different patterns may emerge at differing scales of
investigation of almost every aspect of every ecological system.
Value of Scale Theories:
Early example were species-area
curves, which showed that nonlinear patterns existed, and that
Heuristic value
study area size must be accounted for when interpreting results
of studies of species richness.
Focus measurement

Model Parameterization

Management - Range of Natural Variation

Source: Rahbek Ecology Letters 2005

Temporal and Spatial Scales in Ecology Ecological scaling: definitions

Different patterns may emerge at differing scales Grain = minimum resolution

of investigation of almost every aspect of every of the data, defined by the
ecological system. cell size (for raster data) or
minimum polygon size
(vector data).

Extent = the scope or

domain of the data, defined
as the size of the landscape
or study area under
Local scale: red pine is declining Landscape scale: red pine is
and may disappear from the thriving in Superior Ntl Forest due
Boundary Water Canoe Area to active management.

Ecological scaling: definitions
Ecological scaling: notes and rules

Grain & Vector Data Grain and extent are correlated: Reality
As the study area increases, we
Grain = minimum resolution generally lose resolution; high

of the data = minimum resolution usually requires a smaller Scale of
mapping unit. study area. investigation

The smaller the grain, generally

Is Grain a relevant concept
Upper limit
the more information or data

of resolution
for vector data? collected.

Yes, because: * The upper limit of data resolution

a) Grain influences both area is set by the extent
and length. * The lower limit by the grain.
* We cannot detect patterns finer Lower limit

of resolution
b) You must be aware of the than the grain or coarser than the
Fine Coarse
grain when combining or
using multiple data sets.

Ecological scaling: notes and rules Effects of scale: Abiotic/biotic

To an Organism: Biological interactions may Biota may introduce indirect
separate or reduce the relationships effects, feedbacks, and spatial or
Grain is the smallest component between the abiotic template and temporal lags that are not
of the environment that is relevant landscape pattern. observable at larger scales.
to the organism.

Extent is the maximum distance

at which an organism interacts with
an external object.
To Humans:

Grain might be the finest unit of

management or study.

Extent is the total area under

investigation or management.

May be determined by the scale

of the data or other technological

Effects of scale: Openness Effects of scale: Detail and Mechanism

Openness is defined by the When systems are open, processes
strength of the interactions between at broader scales will affect those at
scales. the scale of observation. Reveals generalizations
A. Fine-scale studies & system dynamics
Island population may reveal more detail
closed to
about the underlying
mechanisms of a pattern Coarse
Temporal scale

Weak Scale`
Scale of study
B. Mechanisms will be
Patch open
to dispersal generalized at broader

Open C. The scale(s) of the

mechanism determines
Sc ine

Strong Reveals mechanisms &


the patterns that can be


ixns Island population underlying patterns

open to
detected. Spatial scale

Theories of Scale: Characteristic scale Theories of Scale: Hierarchy theory
Ecological phenomena
have characteristic spatial Higher-level Scale
and temporal scales, or

spatiotemporal domains,
Dynamics too slow to
Species be a variable
and should be addressed at Extinctions Provides Appears as a constant
their characteristic scales. mechanisms Sets constraints or

Temporal scale Species boundaries

Temporal scale
As the spatial or temporal Reference Scale
scale changes, the Secondary Phenomena or
scale of interest
phenomena of interest
Windthrow Lower-level Scale
Fire context
Treefalls Dynamics fast are
Short-term changes often experienced as averages

affect small areas while Recruitment Appears as background

noise or variability that is
long-term changes affect averaged
Spatial scale
larger areas. Small Broad
Spatial scale

Hierarchy Theory: Examples

Hierarchy theory: examples
Objective: predict the increase in biomass of a forest stand over 100 years
Ecoregion: defined by
climate and geology; 100,000s
ha; 1,000s-10,000 yrs. Higher-level Scale

Dynamics too slow to
Landscape: defined by Soil
be parent materal
a variable
physiographic province,
Landscape as aposition
Mechanisms Ecoregion mechanisms Sets constraints or
changes in land use, boundaries
Temporal scale

disturbance regimes: 10,000s

Temporal scale

Leaf area Scale
Landscape ha, 100-1,000 yrs. Phenomena or
Avg. tree diameter
scale of interest
Stand: defined by Lower-level Scale
Stand topographic position, Tree energy budgets:
disturbance patches: 1-10s ha,

Dynamics fast are

tree physiology
Constraints/ experienced as averages
Gap boundaries 10-100s yrs. root respiration
Appears as background
noise productivity
or variability that is
Gap: defined by the
Spatial scale influence of a single large tree: Spatial scale
0.01-0.1 ha, 1-10s yrs.

Hierarchy theory: examples

Objective: predict forest stand dynamics over 100 years
Dealing with ecological scale: Scaling up
Mechanisms boundaries
Temporal scale

Landscape soil parent materal

landscape position Scaling up involves extrapolating
data to scales of higher levels;
Phenomena of interest

values are inferred for higher scales

often beyond the range of the data
collected for estimation.
Stand Dynamics:
Spatial scale

Stand Stand Stand Tree diameters

Spp Composition
Not all data may be scaled up, since
it assumes constant processes
Gap Gap across scales, linearities, and a
Gap Gap lack of thresholds.
Gap Gap Gap
Gap tree physiology
Gap root respiration
Gap annual productivity
Spatial scale

Theories of Scale: Hierarchy theory Ecological scale: Summary

Summary 1. The scale of an observation has very strong influence over what
you observe, which is important because scales of study are often
Ecosystems can be divided into smaller components arbitrary and based on human perception.
that operate at finer-scales than the phenomena of
interest. 2. Conclusions, as well as data, documented at one scale may not
be applicable or transferable to another scale.
Ecosystems are nested within increasingly larger
ecosystems that influence processes occurring in the 3. Finding the appropriate scale is
systems. sometimes difficult, and there are few
shortcuts in doing so; one must consider
Lower level generate behaviors for levels above; higher the hierarchy and processes
levels constrain levels below. for the object in question.