Models of Landscape Structure

Instructor: K. McGarigal

Assigned Reading: McGarigal (Lecture notes)

Objective: Provide a basic description of several alternative models of landscape structure,

including models based on point, categorical and continuous patterns. Highlight the importance
of selecting a meaningful model for the question under consideration given the constraints of
data availability and software tools available for analyzing pattern-process relationships.

Topics covered:
1. Models of landscape structure
2. Point pattern model
3. Patch mosaic model - island biogeographic and landscape mosaic models
4. Landscape gradient model
5. Graph matrix model
1. Models of Landscape Structure

There are many different ways to model or represent landscape structure corresponding to
different perspectives on landscape heterogeneity. Here we will review five common alternative
models: (1) point pattern model; (2) island biogeographic model based on categorical map
patterns; (3) landscape mosaic model based on categorical map patterns; (4) landscape gradient
model based on continuous surface patterns; and (5) graph-theoretic model.

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The choice of model in any particular application depends on several criteria:

• The ecological pattern-process under consideration and the objective of the analysis
• The spatial character of the landscape with respect to the relevant attributes
• Available spatial data (type, structure and quality)
• Available analytical methods (software tools)
• Available computational resources

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2. Point Pattern Model

2.1. Data Characteristics

Point pattern data comprise collections of the locations of entities of interest, wherein the data
consists of a list of entities referenced by their (x,y) locations. Familiar examples include:

• Map of all trees in a forest stand, perhaps by species

• Map of all occurrences of a focal ecosystem (e.g., seasonal wetland) in a study area
• Map of all detections of an individual of a species during a season or over a lifetime

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2.2. Data Structure

Point pattern data typically are represented using a vector data structure, wherein each point is
referenced by its (x,y) location and occupies no real space. Alternatively, it may be convenient in
some applications (see below) to represent point patterns using a raster data structure, wherein
each point is represented as a cell (or pixel) in a raster grid and thus occupies the space of one
cell. Lastly, it may be useful in some applications to address point patterns using a graph matrix
data structure, which is described later as a special model of landscape structure.

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2.3. Pattern Elements

Point pattern data consist of a single pattern element - points. The points are often
indistinguishable from each other (i.e., unweighted), wherein only the (x,y) location of the points
is of interest. Alternatively, the points may be distinguished from each other on the basis of one
or more attributes (e.g., weights), so that not all points are equal, and this information is then
taken into account in the analysis.

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2.4. Pattern Metrics

The goal of point pattern analysis is typically to quantify the intensity of points at multiple scales
and there are numerous methods for doing so. Here we will review only a few of the more
popular approaches.

(1) 1st-order point patterns

The primary data for point pattern analysis consist of n points tallied by location within an area of
size A (e.g., hundreds of individual trees in a 1-ha stand). The simplest of pattern metrics merely
quantify the number and density of points. However, a plethora of techniques have been
developed for analyzing more complex aspects of spatial point patterns, some based on sample
quadrats or plots and others based on nearest-neighbor distances. A typical distance-based
approach is to use the mean point-to-point distance to derive a mean area per point, and then to
invert this to get a mean point density (points per unit area), Lambda, from which test statistics
about expected point density are derived. There are nearly uncountable variations on this theme,
ranging from quite simple to less so (e.g., Clark and Evans 1954). Most of these techniques
provide a single global measure of point pattern aimed at distinguishing clumped and uniform
distributions from random distributions, but do not help to distinguish the characteristic scale or
scales of the point pattern.

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(2) 2nd -order point patterns – Ripley’s K-distribution

The most popular means of analyzing (i.e., scaling) point patterns is the use of second-order
statistics (statistics based on the co-occurrences of pairs of points). The most common technique
is Ripley's K-distribution or K-function (Ripley 1976, 1977), which we discussed previously as a
scaling technique for point pattern data. Recall that the K-distribution is the cumulative
frequency distribution of observations at a given point-to-point distance (or within a distance
class); that is, it is based on the number of points tallied within a given distance or distance class.
Because it preserves distances at multiple scales, Ripley's K can quantify the intensity of pattern
at multiple scales.

In the real-world example shown here for the distribution of vernal pools in a small region in
Massachusetts, the K function reveals that pools are more clumped than expected under a
spatially random distribution out to a distance of at least 3.5 km, and are perhaps most clumped
at a scale of about 400 m. The scale of clumping of pools is interesting given the dependence of
many vernal pool amphibians on metapopulation processes such as dispersal of individuals
among ponds. For most of these vernal pool-dependent species the pools are highly clumped at
scales corresponding to the range of dispersal distances.

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(3) Local pattern intensity – Kernel estimators

The previous methods provide a numerical summary of the global point pattern; that is, they
provide a quantitative description of the average pattern of points across the entire landscape.
Often times, however, it is more useful to assess the local point pattern and produce a pattern
intensity map. The kernel estimator (Silverman 1986; Worton 1989) is a density estimator, which
we discussed previously as a scaling technique for point pattern data. Recall that a kernel
estimator involves placing a “kernel” of any specified shape and width over each point and
summing the values to create a cumulative kernel surface that represents a distance-weighted
point density estimate. Because we can specify any bandwidth, the kernel estimator can be used
to depict the intensity of point pattern at multiple scales.

In the example shown here, a bivariate normal kernel was placed over each vernal pool in a small
study area from western Massachusetts, and the cumulative kernel surfaces are illustrated here in
three dimensions. The kernel surface on the left results from a bivariate normal kernel with a
bandwidth (standard deviation) of 200m. The one on the right has a bandwidth of 400m. Clearly,
the smaller the bandwidth, the rougher the surface. The peaks represent regions of high vernal
pool density, the troughs represent regions of low vernal pool density.

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2.5. Applications

(1) Home range estimation and mapping

One of the most common applications of point pattern analysis is in home range estimation and
mapping, wherein the data consist of a set of point locations for a single animal collected over
the course of a fixed time period. While there are a number of methods for estimating home
range size and distribution based on such data, the most popular technique is based on kernel
estimation. In this approach, a kernel of specified shape and width is centered on each point
location. The cumulative kernel surface represents the spatial utilization distribution for the
animal. Slicing the cumulative kernel surface at a particular height allows one to depict the area
(or areas) used during a specified percentage of the time. Thus, by slicing the kernel surface quite
low, for example, one can depict say the 95% utilization distribution; that is, the area containing
95% of the animal’s home range use. Conversely, by slicing the kernel surface quite high, one
can depict only core high use areas. In the example shown here, the kernel home range based on
several utilization thresholds is overlaid on the minimum convex polygon surrounding all
locations for a moose in central Massachusetts. It is clear that the kernel approach does a much
better job of estimating and depicting the spatial distribution of home range use than the
minimum convex polygon.

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(2) Modeling ecological processes

Another way in which kernel point pattern analysis is being applied is in the modeling of
ecological processes. Kernels are ideal for modeling point-based ecological processes because
the shape and size or width of the kernel can be adjusted to reflect the particular ecological
process under consideration. The flexibility in the shape of the kernel allows one to depict
nonlinear and even nonparametric ecological processes. In the example shown here, we applied
the concept of a dispersal kernel, which describes the scatter of offspring about the parent plant
in the form of a probability density function, to a landscape dispersal kernel weighted by percent
cover of ponderosa pine in areas adjacent to high-severity patches. Essentially, this involves
placing a kernel over every potential seed tree within the burn and surrounding area. We used a
Gaussian kernel and varied the smoothing parameter (h) to examine different potential seed
dispersal distance functions. Number of seedlings at distances less than 60 m can be high in early
post-fire periods, but remotely dispersed individuals may lead to variations at greater distances
through time. The Gaussian model approaches zero rapidly with distance, making migration a
coherent, stepwise process as compared to fat-tailed models which are expected for higher rates
of dispersal. As shown here, an h=150 m had the best fit and shown a strong relationship with
ponderosa pine regeneration in terms of both total percent cover and stem density. The resulting
kernel map depicts the expected spatial distribution in pine regeneration within the high severity
burn patches.

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In this example, Meador et al. (2009) used point pattern analysis to quantify the spatial
distribution of ponderosa pine trees in historical stands (pre-European land use) and compare
them to the contemporary stand structures (post-European land use) in northern Arizona in an
effort to discern the effects of human land use practices on forest structure. Specifically, they
surveyed a 2.59 ha stand of ponderosa pine forest characteristic of southwestern ponderosa pine
forests in northern Arizona and recorded the location (and size) of all trees existing prior to the
first selective harvest in 1894. Historical trees were located based on the presence of old live
trees and “evidences” of historical (pre-1894) trees, based primarily on stumps and logs, which is
possible because of the arid conditions and very slow decomposition rates. Based on this survey,
they were able to plot the location of every tree (with some confidence) in the pre-1894 stand and
every tree in the contemporary stand.

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As shown here, they used Ripley’s K distribution to quantify the scale of clumping in the tree
distribution. Specifically, they observed a moderate, but significant, clumping of trees at the 0-20
m scale in the pre-harvest (or pre-1894) stand, indicative of trees existing in small clumps and
groups of a few to several trees. Post-harvest, the trees were significantly more clumped and at a
much coarser scale, or even to some degree at all scales considered. However, in the
contemporary stand, the magnitude of clumping decreased dramatically even though it was still
significantly clumped at all scales. They interpreted the differences as indicative of contemporary
stands having lost the fine-scale “clumpy-groupy” structure that was characteristic of pre-
European land use stands.

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(3) Modeling species-environment relationships

Point pattern analysis offers myriad possibilities for modeling species-environment relationships
where either the species or the relevant environmental attributes are best represented as point
features of the landscape. In the example shown here, we used kernel estimators to depict the
spatial distribution of piping plover nests and productivity on Long Island, New York, in relation
to several environmental determinants. In this first slide, the distribution of plover nests along a
portion of the barrier beach are depicted as point locations.

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In this slide, we have used a Guassian kernel to estimate nest density as a continuous surface.
The width of the kernel was selected based on empirical data on territory size. The cumulative
kernel surface clearly distinguishes the areas of highest nest density. The following two slides
depict the distribution of nest productivity (i.e., number of young fledged per nest)(red) in
relation to nest density (blue). Note, the nest productivity distribution is based on weighted point
data, where each point location (nest) also contains a weight, in this case, productivity of the
nest. In addition, note that high nest density does not always equal high nest productivity – why?

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We also used point-based kernels to depict the distribution of certain environmental variables
deemed to be potentially important determinants of plover distribution, abundance and/or
productivity. Human activity on beeches is believed to be a major form of disturbance to nesting
plovers, and it comes in many forms as shown in this slide. The next two slides depict the
distribution of people on the beach recorded during regular aerial surveys and the cumulative
kernel surface derived from the point locations. The kernel map only shows the highest values
(288-4000 people/km2 ).

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This slide shows the plover nest density kernel overlaid on the people on beech kernel and there
is some indication of an inverse relationship between nest density and people density (i.e., nest
density increases where people density decreases). However, this relationship must be explored
statistically before any inferences about the relationship can be made.

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Lastly, we have used the point-based kernel approach to estimate the distribution of several
point-based environmental variables. Two others shown here are beach off-road vehicle (ORV)
activity (where each ORV is recorded as a separate point location) and gull intensity (where each
recorded gull is recorded as a separate point location). Gulls of several species are potential
predators of plover chicks. Currently, we are using statistical procedures to examine the
relationship between plover nest density and productivity and each of these and several other
environmental variables. The key point here is that point pattern analysis underpins most of our
analysis into the plover-environment relationship.

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2.6. Pros and Cons

Considering the information presented thus far, what are the strengths and limitations of the point
pattern model of landscape structure?

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3. Patch Mosaic Model

3.1. Data Characteristics

The patch mosaic model represents data in which the system property of interest is represented as
a mosaic of discrete patches that can be intuitively defined by the notion of "edge" (a patch is an
area with edges adjoining other patches). From an ecological perspective, patches represent
relatively discrete areas of relatively homogeneous environmental conditions at a particular scale.
The patch boundaries are distinguished by abrupt discontinuities (boundaries) in environmental
character states from their surroundings of magnitudes that are relevant to the ecological
phenomenon under. The patch mosaic model is the dominant model of landscape structure in use
today. Familiar examples include:

• Map of land cover types

• Map of ownership parcels

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3.2. Data Structure

In the patch mosaic model, the data consists of polygons (vector) or grid cells (raster) classified
into discrete classes. There are many methods for deriving a categorical map of patches. Patches
may be classified and delineated qualitatively through visual interpretation of the data (e.g.,
delineating vegetation polygons through interpretation of aerial photographs), as is typically the
case with vector maps constructed from digitized lines. Alternatively, with raster grids,
information at every location, typically obtained through remote sensing, may be used to classify
cells into discrete classes and then to delineate patches by outlining them, and there are a variety
of methods for doing this.

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3.3. Pattern Elements

In the patch mosaic model three major landscape elements are typically recognized: patches,
corridors and matrix, and the extent and configuration of these elements defines the pattern of
the landscape. The patch mosaic model is most powerful when meaningful patches can be clearly
defined and accurately mapped as discrete patches, and when the variation within a patch is
deemed relatively insignificant. The patch mosaic model assumes patches are homogeneous
within (or treats them so) and categorically different from one another. For example, breeding
habitat for many pond-breeding amphibians can be clearly defined and delineated with relatively
little uncertainty, and the variation in habitat quality within ponds is insignificant compared to
among-pond differences or pond-upland differences. Similarly, forested woodlots embedded
within a contrasting agricultural or urban landscape, fields in a forested landscape, stands of
deciduous trees within a coniferous forest, and many other examples can be represented easily
and meaningfully in a categorical map. In general, whenever disturbances (natural or
anthropogenic) either create discrete patches or leave behind discrete remnant patches, the patch
mosaic model is likely to be useful.

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(1) Patch

In the patch mosaic model, landscapes are composed of a mosaic of patches (Urban et al. 1987).
Landscape ecologists have used a variety of terms to refer to the basic elements or units that
make up a patch mosaic, including ecotope, biotope, landscape component, landscape element,
landscape unit, landscape cell, geotope, facies, habitat, and site (Forman and Godron 1986). Any
of these terms, when defined, are satisfactory according to the preference of the investigator. Like
the landscape, patches comprising the landscape are not self-evident; patches must be defined
relative to the phenomenon under consideration. For example, from a timber management
perspective a patch may correspond to the forest stand. However, the stand may not function as a
patch from a particular organism's perspective. From an ecological perspective, patches represent
relatively discrete areas (spatial domain) or periods (temporal domain) of relatively homogeneous
environmental conditions where the patch boundaries are distinguished by discontinuities in
environmental character states from their surroundings of magnitudes that are perceived by or
relevant to the organism or ecological phenomenon under consideration (Wiens 1976). From a
strictly organism-centered view, patches may be defined as environmental units between which
fitness prospects, or "quality", differ; although, in practice, patches may be more appropriately
defined by nonrandom distribution of activity or resource utilization among environmental units,
as recognized in the concept of "Grain Response".

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Patches are dynamic and occur on a variety of spatial and temporal scales that, from an organism-
centered perspective, vary as a function of each animal's perceptions (Wiens 1976 and 1989,
Wiens and Milne 1989). A patch at any given scale has an internal structure that is a reflection of
patchiness at finer scales, and the mosaic containing that patch has a structure that is determined
by patchiness at broader scales (Kotliar and Wiens 1990). Thus, regardless of the basis for
defining patches, a landscape does not contain a single patch mosaic, but contains a hierarchy of
patch mosaics across a range of scales.

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Patch boundaries are artificially imposed and are in fact meaningful only when referenced to a
particular scale (i.e., grain size and extent). For example, even a relatively discrete patch
boundary between an aquatic surface (e.g., lake) and terrestrial surface becomes more and more
like a continuous gradient as one progresses to a finer and finer resolution. However, most
environmental dimensions possess one or more "domains of scale" (Wiens 1989) at which the
individual spatial or temporal patches can be treated as functionally homogeneous; at
intermediate scales the environmental dimensions appear more as gradients of continuous
variation in character states. Thus, as one moves from a finer resolution to coarser resolution,
patches may be distinct at some scales (i.e., domains of scale) but not at others.

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KEY POINT: It is not my intent to argue for a particular definition of patch. Rather, I wish to
point out the following: (1) that patch must be defined relative to the phenomenon under
investigation or management; (2) that, regardless of the phenomenon under consideration (e.g.,
a species, geomorphological disturbances, etc), patches are dynamic and occur at multiple
scales; and (3) that patch boundaries are only meaningful when referenced to a particular scale.
It is incumbent upon the investigator or manager to establish the basis for delineating among
patches and at a scale appropriate to the phenomenon under consideration.

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(2) Corridor

Corridors are linear landscape elements that can be defined on the basis of structure or function.
Forman and Godron (1986) define corridors as “narrow strips of land which differ from the
matrix on either side. Corridors may be isolated strips, but are usually attached to a patch of
somewhat similar vegetation.” These authors focus on the structural aspects of the linear
landscape element and recognize three different types of structural corridors: (1) line corridors,
in which the width of the corridor is too narrow to allow for interior environmental conditions to
develop; (2) strip corridors, in which the width of the corridor is wide enough to allow for
interior conditions to develop; and (3) stream corridors, which are a special category.
Alternatively, these authors also classify corridors based on the agent of formation, including: (1)
disturbance corridors, in which the corridor is established by a disturbance event, usually
anthropogenic (e.g., roads, utility lines, fences); (2) remnant corridors, in which the corridor is
the result of disturbance around the corridor, leaving the corridor as an intact remnant of the
formally widespread cover type; and (3) environmental corridors, in which the corridor is the
result of a strong linear environmental gradient, such as a riparian corridor created by the land-
water interface along a stream.

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As a consequence of their form and context, structural corridors may function as habitat,
dispersal conduits, barriers, or as a source of abiotic and biotic effects on the surrounding matrix:

• Habitat Corridor.--Linear landscape element that provides for survivorship, natality, and
movement (i.e., habitat), and may provide either temporary or permanent habitat. Habitat
corridors passively increase landscape connectivity for the focal organism(s).

• Facilitated Movement Corridor.–Linear landscape element that provides for survivorship and
movement, but not necessarily natality, between other habitat patches. Facilitated movement
corridors actively increase landscape connectivity for the focal organism(s).

• Barrier or Filter Corridor.–Linear landscape element that prohibits (i.e., barrier) or

differentially impedes (i.e., filter) the flow of energy, mineral nutrients, and/or species across
(i.e., flows perpendicular to the length of the corridor). Barrier or filter corridors actively
decrease matrix connectivity for the focal process.

• Source of Abiotic and Biotic Effects on the Surrounding Matrix.–Linear landscape element
that modifies the inputs of energy, mineral nutrients, and/or species to the surrounding matrix
and thereby effects the functioning of the surrounding matrix.

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Most of the past attention and debate has focused on facilitated movement corridors. It has been
argued that this corridor function can only be demonstrated when the immigration rate to the
target patch is increased over what it would be if the linear element was not present (Rosenberg
et al. 1997). Unfortunately, as Rosenberg et al. point out, there have been few attempts to
experimentally demonstrate this. In addition, just because a corridor can be distinguished on the
basis of structure, it does not mean that it assumes any of the above functions. Moreover, the
function of the corridor will vary among organisms due to the differences in how organisms
perceive and scale the environment. More recently, the attention has shifted to the role of
corridors as barriers or impediments to ecological flows.

KEY POINT: Corridors are distinguished from patches by their linear nature and can be
defined on the basis of either structure or function or both. If a corridor is specified, it is
incumbent upon the investigator or manager to define the structure and implied function relative
to the phenomena (e.g., species) under consideration.

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(3) Matrix

In the patch mosaic model, the matrix is the most extensive and most connected element, and
therefore plays the dominant role in the functioning of the landscape. For example, in a large
contiguous area of mature forest embedded with numerous small disturbance patches, the mature
forest constitutes the matrix because it is greatest in extent, is mostly connected, and exerts a
dominant influence on the biota and ecological processes. In most landscapes the matrix is
obvious to the investigator or manager. However, in some landscapes, or at a certain point in
time, the matrix will not be obvious, and it may not be appropriate to consider any element as the
matrix. The designation of a matrix is largely dependent upon the phenomenon under
consideration. For example, in the study of geomorphological processes, the geological substrate
may serve to define the matrix and patches; whereas, in the study of vertebrate populations,
vegetation structure may serve to define the matrix and patches. In addition, what constitutes the
matrix is dependent on the scale of investigation or management. For example, at a particular
scale, mature forest may be the matrix with disturbance patches embedded within; whereas, at a
coarser scale, agricultural land may be the matrix with mature forest patches embedded within.

KEY POINT: A matrix element is not inherent to the landscape. It is incumbent upon the
investigator or manager to determine whether a matrix element exists and should be designated
given the scale and phenomenon under consideration.

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3.4. Perspectives on Categorical Landscapes

There are at least two different perspectives on the patch mosaic mode and categorical map
patterns that have profoundly influenced the development of landscape ecology and have
important implications for the analysis of landscape patterns.

(1) Island biogeographic model

In the island biogeographic model, the emphasis is on a single patch type; disjunct patches (e.g.,
habitat fragments) are viewed as analogues of oceanic islands embedded in an inhospitable or
ecologically neutral background (matrix). This perspective emerged from the theory of island
biogeography (MacArthur and Wilson 1967) and subsequent interest in habitat fragmentation
(Saunders et al. 1991). Under this perspective, there is a binary patch structure in which the focal
patches (fragments) are embedded in a neutral matrix. Here, the emphasis is on the extent, spatial
character, and distribution of the focal patch type without explicitly considering the role of the
matrix. Under this perspective, for example, connectivity may be assessed by the spatial
aggregation of the focal patch type without consideration of how intervening patches affect the
functional connectedness among patches of the focal class. The island biogeography perspective
has been the dominant perspective since inception of the theory. The major advantage of the
island model is its simplicity. Given a focal patch type, it is quite simple to represent the structure

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of the landscape in terms of focal patches contrasted sharply against a uniform matrix, and it is
relatively simple to devise metrics that quantify this structure. Moreover, by considering the
matrix as ecologically neutral, it invites ecologists to focus on those patch attributes, such as size
and isolation, that have the strongest effect on species persistence at the patch level. The major
disadvantage of the strict island model is that it assumes a uniform and neutral matrix, which in
most real-world cases is a drastic over-simplification of how organisms interact with landscape
patterns.

(2) Landscape mosaic model

In the landscape mosaic model, landscapes are viewed as spatially complex, heterogeneous
assemblages of patch types, which can not be simply categorized into discrete elements such as
patches, matrix, and corridors (With 2000). Rather, the landscape is viewed from the perspective
of the organism or process of interest. Patches are bounded by patches of other patch types that
may be more or less similar to the focal patch type, as opposed to highly contrasting and often
hostile habitats, as in the case of the island model. Connectivity, for example, may be assessed by
the extent to which movement is facilitated or impeded through different patch types across the
landscape. The landscape mosaic perspective derives from landscape ecology (Forman 1995) and
has only recently emerged as a viable alternative to the island biogeographic model. The major
advantage of the landscape mosaic model is its more realistic representation of how organisms
perceive and interact with landscape patterns. Few organisms, for example, exhibit a binary (all
or none) response to habitats (patch types), but rather use habitats proportionate to the fitness
they confer to the organism. Moreover, movement among suitable habitat patches usually is a
function of the character of the intervening habitats. The major disadvantage of the landscape
mosaic model is that it requires detailed understanding of how organisms interact with landscape
pattern, and this has delayed the development of additional quantitative methods that adopt this
perspective.

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3.5. Pattern Metrics

Regardless of data format (raster or vector) and method of classifying and delineating patches,
the goal of categorical map pattern analysis with such data is to characterize the composition and
spatial configuration of the patch mosaic, and a plethora of metrics has been developed for this
purpose. We will explore these pattern metrics in detail in a subsequent lecture (and lab), so for
now, suffice it to say that there are many different metrics available for quantifying the
composition and configuration of the patch mosaic. It is worth noting that in contrast to the
emphasis on identifying the scale of pattern with point data and continuous data, so-called
scaling techniques for categorical map data are less commonly employed in landscape ecology.
This may seem somewhat surprising given the predominant use of categorical data in landscape
ecological investigations – after all, the predominant patch mosaic model of landscape structure
is based on a categorical data format. However, there has been a plethora of landscape metrics
developed for quantifying various aspects of categorical map patterns and these have largely
taken the place of the more conventional scaling techniques. In addition, in applications
involving categorical map patterns, the relevant scale of the mosaic is often defined a priori
based on the phenomenon under consideration. In such cases, it is usually assumed that it would
be meaningless to determine the so-called characteristic scale of the mosaic after its construction.

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3.6. Applications

(1) Landscape ecological assessment

Landscape ecological assessment based on the patch mosaic model of landscape structure is
increasingly common. Given the inherent complexity of ecological systems and the daunting
challenges confronting managers seeking to sustain ecosystems in the face of increasing human
pressures, it is not too surprising that managers are increasingly seeking effective ecological
indicators. Landscape metrics that quantify the composition and configuration of the patch
mosaic are increasingly being used as course-scale ecological indicators of change. In the
example shown here, several different pattern metrics were evaluated for their sensitivity to land
use change between 1974 and 1999 at Fort Benning, Georgia. In the realm of wetland ecological
assessment there has been a similar explosion in the use of landscape pattern metrics to evaluate
the condition of wetlands as part of a regional wetlands monitoring and assessment program.
While the current methods almost exclusively adopt pattern metrics based on the patch mosaic
model of landscape structure, it is important to note that this is only by convention and that
metrics derived from other conceptual models of landscape structure apply equally well.

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(2) Landscape disturbance-succession modeling

One of the more common applications of the patch mosaic model in land management has been
in landscape disturbance-succession modeling (LDSM). In most LDSMs the landscape is
represented as patch mosaic, where the patches represent discrete land cover types (e.g.,
vegetation communities) and disturbance and succession processes operate in various ways to
alter the structure of the mosaic over time, such as shown in the animation on this slide. Pattern
metrics based on the patch mosaic are used to quantify the structure of the landscape at each
point in time during the simulation. Statistical summaries of the resulting trajectory of change
can then be used to quantify the range of variation in landscape structure under the particular
scenario simulated. This is a popular basis for characterizing the historic range of variability
(HRV) in landscapes and in comparing alternative future land management scenarios, two
subjects that we will explore in greater depth in subsequent lectures and labs.

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(3) Modeling species-environment relationships

Perhaps the most common application of pattern analysis based on the patch mosaic model is in
modeling species-environment relationships, and there are myriad examples of such applications.
An increasingly common example involves quantifying the landscape structure around sample
locations for the purpose of building statistical models to explain and/or predict the distribution,
abundance or performance of a species. In the example shown here, we adopted the patch mosaic
model to map the distribution of several major cover types deemed potentially relevant to a
variety of moth species in the pine barrens of southeastern Massachusetts. Conventional
landscape pattern analysis involves quantifying the structure of the entire landscape mosaic and
returning a single computed value for each metric.

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However, in this study we were more interested in quantifying the local neighborhood structure
around each point location (represented as a cell in the raster grid). To do this, we passed a
moving window of specified shape and size across the landscape one cell at time. Within each
window, we compute the desired landscape metric and returned the value to the focal (center)
cell. By doing this for every cell in the input landscape, the result was a new grid depicting a
continuous surface representing the local landscape structure as measured by the particular
metric.

As shown in the next two slides, we repeated this process for several different landscape metrics
at several different scales (i.e., window sizes). These local landscape structure surfaces
represented independent variables in a logistic regression, where the dependent variable was
presence/absence of a particular moth species. Using statistical procedures, we identified the
combination of metrics and scales that best predicted each species presence/absence. By
overlaying the predicted distribution of each species, we were able to identify areas of high moth
richness, which might serve to inform conservation decisions regarding where to focus land
conservation efforts.

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3.7. Pros and Cons

Considering the information presented thus far, what are the strengths and limitations of the
patch mosaic model of landscape structure?

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4. Landscape Gradient Model

4.1. Data Characteristics

An alternative approach to the conventional patch mosaic model involves representing

heterogeneity continuously – as a gradient. In this model, heterogeneity does not exist in discrete
patches, but rather exists as a continuously varying property of the local environment and
landscape. Here, the data can be conceptualized as representing a three-dimensional surface,
where the measured value at each geographic location is represented by the height of the surface.
For example, instead of representing habitat as discrete patches, it is represented as a suitability
or capability index, where the value at each location represents the quality of habitat and can take
on continuous values. In practice, habitat suitability or capability is often classified into
categories representing, say, high-, moderate-, and low-quality habitat, but this is more often
done for convenience to facilitate further analysis with conventional categorical-based
procedures or to simplify the presentation of results. Other familiar examples of inherently
continuous gradients include:

• Map of elevation
• Map of burn severity
• Map of leaf area index

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4.2. Data Structure

Continuous surface pattern data typically are represented using a raster data structure, wherein
each cell (or pixel) takes on a continuous value. Alternatively, it may be convenient in some
applications to represent continuous surface data using a vector data structure, wherein the
surface is represented as a series of contours (lines) as in the familiar topographic map.

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4.3. Pattern Elements

Interestingly, there are no pattern elements in the landscape gradient model, making it the most
parsimonious of the landscape structure models.

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4.4. Pattern Metrics

A wide variety of methods have been developed for quantifying the intensity and scale of pattern
in regionalized quantitative variables. Recall that a “regionalized” variable is one that takes on
values based on its spatial location and that the analysis of the spatial dependencies (or
autocorrelation – the ability to predict values of a variable from known values at other locations,
as described by a ‘structure function’) in the measured characteristic is the purview of
geostatistics. Recall that a variety of techniques exist for measuring the intensity and scale of this
spatial autocorrelation, and while the location of the data points (or quadrats) is known and of
interest, it is the values of the measurement taken at each point that are of primary concern.

(1) Autocorrelation Structure Functions

The most basic and common measures of pattern in regionalized quantitative variables (i.e.,
landscape gradients) are based on autocorrelation structure functions, including for example
Moran’s I autocorrelation coefficient and semivariance, which we described previously as scaling
techniques for continuous gradient data. Both measures are typically used to describe the
magnitude of autocorrelation as a function of distance between locations, as expressed by the
correlogram and variogram (or semi-variogram), respectively, and illustrated here for a gradient
in the topographic moisture index along a transect at the Coweeta Experiment Station.

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(2) Other Structure Functions

There are many other methods for analyzing the intensity and scale of pattern with continuous
data, especially when the data is collected along continuous transects or two-dimensional
surfaces, which we described previously as scaling techniques for continuous gradient data. Like
the autocorrelation structure functions, these techniques are typically used to quantify spatial
dependencies in a quantitative variable in relation to scale (distance). Recall that no one
technique has been found to be superior for all applications.

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(3) Surface metrology metrics

The autocorrelation and other related structure functions described above can provide useful
indices to quantitatively compare the intensity and extent of autocorrelation in quantitative
variables among landscapes. However, while they can provide information on the distance at
which the measured variable becomes statistically independent, and reveal the scales of repeated
patterns in the variable, if they exist, they do little to describe other interesting aspects of the
surface. For example, the degree of relief, density of troughs or ridges, and steepness of slopes
are not measured. Fortunately, a number of gradient-based metrics that summarize these and
other interesting properties of continuous surfaces have been developed in the physical sciences
for analyzing three-dimensional surface structures (Barbato et al. 1996, Sout et al. 1994,
Villarrubia 1997). In the past ten years, researchers involved in microscopy and molecular
physics have made tremendous progress in this area, creating the field of surface metrology
(Barbato et al. 1996).

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In surface metrology, several families of surface pattern metrics have become widely utilized.
One so-called family of metrics quantify intuitive measures of surface amplitude in terms of its
overall roughness, skewness and kurtosis, and total and relative amplitude. Another family
records attributes of surfaces that combine amplitude and spatial characteristics such as the
curvature of local peaks. Together these metrics quantify important aspects of the texture and
complexity of a surface. A third family measures certain spatial attributes of the surface
associated with the orientation of the dominant texture. A final family of metrics is based on the
surface bearing area ratio curve (or Abbott curve). The Abbott curve is computed by inversion of
the cumulative height distribution histogram. The curve describes the distribution of mass in the
surface across the height profile. A number of indices have been developed from the proportions
of this cumulative height-volume curve which describe structural attributes of the surface.

Many of the patch-based metrics for analyzing categorical landscapes have analogs in surface
metrology. For example, compositional metrics such as patch density, percent of landscape and
largest patch index are matched with peak density, surface volume, and maximum peak height.
Configuration metrics such as edge density, nearest neighbor index and fractal dimension index
are matched with mean slope, mean nearest maximum index and surface fractal dimension. Many
of the surface metrology metrics, however, measure attributes that are conceptually quite foreign
to conventional landscape pattern analysis. Landscape ecologists have not yet explored the
behavior and meaning of these new metrics; it remains for them to demonstrate the utility of
these metrics, or develop new surface metrics better suited for landscape ecological questions.

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McGarigal et al. (2009) examined landscapes in Turkey defined using both the landscape
gradient and patch mosaic models according to a variety of landscape definition schemes and
conducted multivariate statistical analyses to identify the universal, consistent and important
components of surface patterns and their relationship to patch-based metrics. They observed four
relatively distinct components of landscape structure based on empirical relationships among 17
surface metrics across 18 landscape gradient models:

1. Surface roughness

The dominant structural component of the surfaces was actually a combination of two distinct
sub-components: (1) the overall variability in surface height and (2) the local variability in slope.
The first sub-component refers to the nonspatial (composition) aspect of the vertical height
profile; that is, the overall variation in the height of the surface without reference to the
horizontal variability in the surface, and is represented by three surface amplitude metrics:
average roughness (Sa), root mean square roughness (Sq), and ten-point height
(S10z)(Appendix). These metrics are analogous to the patch type diversity measures (e.g.,
Simpson's diversity index) in the patch mosaic paradigm, whereby greater variation in surface
height equates to greater landscape diversity. Importantly, while these metrics reflect overall
variability in surface height, they say nothing about the spatial heterogeneity in the surface.

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The second sub-component refers to the spatial (configuration) aspect of surface roughness with
respect to local variability in height (or steepness of slope), and includes two surface metrics:
surface area ratio (Sdr) and root mean square slope (Sdq)(Appendix). These metrics are
analogous to the edge density and contrast metrics (e.g., contrast-weighted edge density, total
edge contrast index) in the patch mosaic paradigm, whereby greater local slope variation equates
to greater density and contrast of edges. Interestingly, while these surface metrics reflect
something akin to edge contrast, they do so without the need to supply edge contrast weights
because they are structural metrics. These two metrics appear to have the greatest overall analogy
to the patch-based measures of spatial heterogeneity and overall patchiness. A fine-grained patch
mosaic (as represented by any number of common patch metrics, such as mean patch size or
density) is conceptually equivalent to a rough surface with high local variability.

On conceptual and theoretical grounds, these spatial and nonspatial aspects of surface roughness
are independent components of landscape structure; however, in the landscape gradients we
examined these two aspects were highly correlated empirically. This distinction between
conceptually and/or theoretically related metrics and groupings based on their empirical behavior
has also been demonstrated for patch metrics.

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2. Shape of the surface height distribution

Another important nonspatial (composition) component of the surfaces we examined was the
shape of the surface height distribution. This component was comprised of five metrics:
skewness (Ssk), kurtosis (Sku), surface bearing index (Sbi), valley fluid retention index (Svi),
and core fluid retention index (Sci). All of these metrics measure departure from a Gaussian
distribution of surface heights, but emphasize different aspects of departure from normality
(Appendix). Ssk and Sku measure the familiar skewness and kurtosis of the surface height
distribution, while the surface bearing metrics, Sbi, Sci and Svi, measure different aspects of the
surface height distribution in its cumulative form. This component was universally present across
landscape models, but the composition of metrics varied somewhat among models reflecting the
complexities inherent in measuring non-parametric shape distributions. There were no strong
patch mosaic analogs to these surface metrics; however, departure from a Gaussian distribution
of surface heights was weakly correlated with, and conceptually most closely related to,
patch-based measures of landscape dominance (or its compliment, evenness) such as Simpson's
evenness index (SIEI) and largest patch index (LPI). Importantly, these five surface metrics
measure the 'shape' of the surface height distribution and are not affected by the surface
roughness (as defined above) per se.

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3. Angular texture

A third prominent component of the surfaces we examined was the angular orientation
(direction) of the surface texture and its magnitude. This component is inherently spatial, since
the arrangement of surface peaks and valleys determines whether the surface has a particular
orientation or not, and is represented by four spatial metrics: dominant texture direction (Std),
texture direction index (Stdi), and two texture aspect ratios (Str20 and Str37). The computational
methods behind these metrics are too complex to describe here (but see Appendix), but are based
on common geostatistical methods (Fourier spectral analysis and autocorrelation functions) that
determine the degree of anisotropy (orientation) in the surface. Not surprisingly given our
knowledge of the study landscape, we did not observe sample landscapes with a strong texture
orientation. We did observe mild levels of texture orientation in some landscapes, but many were
without apparent orientation. Importantly, the measurement of texture direction has no obvious
analog in the patch mosaic paradigm; indeed, we observed no pairwise correlation greater than
±0.22 between any of these four surface metrics and any of the 28 patch metrics.

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4. Radial texture

The fourth prominent component of the surfaces we examined was the radial texture of the
surface and its magnitude. Radial texture refers to repeated patterns of variation in surface height
radiating outward in concentric circles from any location. Like angular texture, this component is
inherently spatial, since the arrangement of surface peaks and valleys determines whether the
surface has any radial texture or not, and is represent by three spatial metrics: dominant radial
wavelength (Srw), radial wave index (Srwi), and fractal dimension (Sfd). Again, the
computational methods behind these metrics are based on common geostatistical methods. A
limitation of these and other metrics based on Fourier spectral analysis and autocorrelation
functions is that they are only sensitive to repeated, regular patterns. We observed that in the
absence of a prominent radial texture, the dominant radial wavelength (Srw) ends up being equal
to the diameter of the sample landscape. As a result, in some of our landscape gradient models
we observed too little variation in this metric and were forced to drop it from the final analyses.
Despite these limitations, we observed sample landscapes with varying degrees of radial texture
based on the other two metrics. In contrast to angular texture, the measurement of radial texture
has at least one conceptual analog in the patch mosaic paradigm - mean and variability in nearest
neighbor distance. On conceptual grounds, Srw should equate to mean nearest neighbor distance,
and Srwi and Sfd should equate to the coefficient of variation in nearest neighbor distance.
However, in our study the corresponding pairwise correlations did not exceed ±0.22, nor were
there any pairwise correlations greater than ±0.40 between either of these surface metrics and any
of the 28 patch metrics.

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4.5. Applications

(1) Scaling gradient patterns

By far the most common application of surface pattern metrics in landscape ecology has been to
identify the characteristic scale or scales of the ecological phenomena or to elucidate the nature
of the scaling relationship. One thing that is true of pattern, as we have already seen, is that its
expression varies with scale. Thus, a careful evaluation of the relationship between scale and
pattern is often considered an important first step in the characterization of pattern. In addition,
we noted previously, quantifying the scaling relationship of the ecological phenomenon of
interest can provide unique insights into potential pattern-process relationships. For example,
multi-scaled patterns (i.e., those with two or more distinctive characteristic scales) revealed
through scaling techniques can indicate the existence of multiple agents of pattern formation
operating at different scales. Conversely, scaling techniques can be used to assess a priori
hypotheses concerning agents of pattern formation; e.g., is the expected scale or scales of pattern
expressed in the data?

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In the example shown here, semivariance was used to examine the scale of pattern in the
abundance of coastal cutthroat trout in several Pacific Northwest streams. In this case, the
regionalized quantitative variable of interest is fish abundance and the spatial lag is stream
distance; i.e., the distance between two points on the stream along the flow path. The
semivariograms reveal different patterns of variation among streams. For example, the pattern of
variation for Glenn Creek in the Coast Range shows a distinct sill with a range of approximately
1100 m. In contrast, the pattern for Miller Creek in the Cascade Range reveals a nested patchy
pattern with a gradient; that is, a distinct scale of patchiness at roughly 300-400 m superimposed
on a coarse-scale gradient of increasing dissimilarity in fish abundance with increasing stream
distance. These patterns may reveal something about the differences between these stream
systems in the underlying distribution of habitat (e.g., the distribution of channel unit types).

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In the example shown here (from Meador et al. 2009), continuing with the example shown earlier
in the point pattern analysis section, correlograms were used to discern the pattern of spatial
autocorrelation in tree diameter (dbh) as a proxy fo establishment age in historical forest stands
(pre-European land use) versus contemporary forest stands (post-European land use) in
ponderosa pine forests of northern Arizona. In all cases, there was strong autocorrelation out
to15-30 m in the pre-1894 and contemporary stands, which the authors interpreted as evidence of
the “clumpy-groupy” nature of tree establishment, whereby trees establish in small groups largely
in the openings between the patches of canopy.

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They also estimated tree establishment age for all the historical trees and contemporary trees and
fit a semivariogram to the data (as shown) and then used the process of krigging to create an
interpolated surface of tree establishment age. In the figure shown here, the colored areas
represent contours of tree establishment age, with the immediate post-harvest 1909 residual trees
shown as points (with the symbol sized proportionate to dbh). The pattern reveals that trees
largely established in cohorts up until the last regeneration event in 1953 and established first in
the openings in clumps/groups and then gradually established closer to the existing canopies and
lastly underneath the existing canopy.

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(2) Landscape disturbance-succession modeling

Another application of landscape gradient model in land management has been in landscape
disturbance-succession modeling (LDSM). As noted previously, most LDSMs adopt the patch
mosaic model of landscape structure; however, some models utilize a gradient-based approach or
a hybrid approach in which some aspects of the landscape are represented as a continuous
surface. For example, the LANDIS model represents vegetation as a continuous surface where
each cell can take on continuous values for the abundance of each tree species and age cohort.
Disturbance and succession processes operate in various ways to alter the distribution of
individual species and age cohorts over time, essentially modifying the continuous surface
structure in species abundance and age. Simple landscape composition metrics that quantify the
total area occupied by each species and age class can then be computed for each timestep of the
model and summarized for the simulation as a whole. And while surface metrology metrics have
not yet been employed to describe the changing landscape structure in connection with this
simulation model, the potential exists to do so.

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(3) Modeling pattern-process relationships

There are myriad possibilities for using the landscape gradient model to explore pattern-process
relationships, which we will illustrate here with a couple of examples.

In the first example, Moody et al. (2007) used the gradient model to examine the relationship
between burn severity and runoff after a large wildfire. Extreme floods often follow wildfire in
mountainous watersheds. However, a quantitative relation between the runoff response and burn
severity at the watershed scale has not been established. Runoff response was measured as the
runoff coefficient C, which is equal to the peak discharge per unit drainage area divided by the
average maximum 30 min rainfall intensity during each rain storm. The magnitude of the burn
severity was expressed as the change in the normalized burn ratio, a continuous surface derived
from LandsatTM imagery. A new burn severity variable, hydraulic functional connectivity Çi was
developed and incorporates both the magnitude of the burn severity and the spatial sequence of
the burn severity along hillslope flow paths. The runoff response and the burn severity were
measured in seven subwatersheds in the upper part of Rendija Canyon burned by the 2000 Cerro
Grande Fire near Los Alamos, New Mexico, USA. The runoff coefficient was a linear function of
the mean hydraulic functional connectivity of the subwatersheds. Moreover, the variability of the
mean hydraulic functional connectivity was related to the variability of the mean runoff
coefficient, and this relation provides physical insight into why the runoff response from the
same subwatershed can vary for different rainstorms with the same rainfall intensity.

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In the second example, Cushman et al. (2006) used the gradient model to examine the
relationship between gene flow and landscape structure in black bears in northern Idaho. They
tested several hypotheses about environmental factors affecting gene flow, including the
hypothesis that observed gene flow was best explained by landscape resistance represented as a
continuous surface based on a factorial combination of elevation, slope, vegetative cover, and
roads. Here, landscape resistance derived from multiple environmental attributes was portrayed
as a continuous surface using the landscape gradient model of landscape structure. As will be
explained in more detail in a subsequent lecture, they used statistical procedures to compare the
genetic distance among the individual bears sampled across the landscape to the ecological
distance among sample locations based on least cost distances derived from the resistant
landscape surface.

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4.6. Pros and Cons

Considering the information presented thus far, what are the strengths and limitations of the
landscape gradient model of landscape structure?

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5. Graph matrix model

5.1. Data Characteristics

An altogether different approach for representing landscape structure that in many ways adopts
aspects of all the preceding models is based on graph theory. In the graph-theoretic model, the
data consists of a collection of nodes (patches) and linkages (connections) represented as data
matrices. In this model, the landscape is simplified into a graph depicting the focal points or
patches as the nodes and the linkages between nodes as connections. In the example shown here,
the grass openings, representing habitat patches, are the nodes and the shortest pathway between
patches are the links. Graph theory is widely used in computer science and engineering, and
became popular in food-web theory and landscape ecology in the early 90's. In landscape
ecology, graph theory has been described as a tool that bridges the gap between structural
measures and functional measures of a landscape, where it has been proven useful in examining
connectivity and ecological flows.

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5.2. Data Structure

The graph-matrix model is distinctive in representing the landscape structure as a set of data
matrices, wherein the spatial heterogeneity is summarized in two non-spatial data matrices:

1. Node data matrix – contains the x,y coordinates of the nodes and may also contain additional
information about the nodes such as the size and/or quality of the patch.
2. Link data matrix – square matrix containing information pertaining to the links between
every pair of nodes. There are at least three different sorts of edge matrices possible:
a. Distance matrix – containing ecological distances between every pair of nodes, based on
simple Euclidean distance or functional distance, for example using least cost paths
derived from resistance surfaces. In addition, distances can be from nearest patch edge to
patch edge or from patch centroid to patch centroid.
b. Flux/Dispersal probability matrix – containing rates of ecological flow (e.g., dispersal)
between every pair of nodes, derived from the distance matrix by applying some function
to the distance, from some attribute of the nodes (e.g., patch size, population size), or
from empirical data collected in the field.
c. Adjacency matrix – binary indicator matrix containing 0's for disconnected links and 1's
for connected links. The adjacency matrix could be created from either of above matrices
using a threshold distance to define connectivity.

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5.3. Pattern Elements

Graph matrices contain two basic elements: nodes and links.

1. Nodes – nodes (also referred to as vertices) can represent any point or patch features of the
landscape relevant to the phenomenon under consideration. It is important to recognize that in the
graph nodes are depicted as points, but in reality they may represent patches of widely varying
sizes. In most cases, the patch centroid is used to represent the patch location in the graph.

2. Links – links (also referred to as edges in the graph theoretic framework) represent the
connections between nodes. Again, it is important to recognize that in the graph links are
depicted as straight lines between nodes, but in reality they may represent nonlinear pathways
between nodes or flux rates between nodes that don’t have an explicit spatial representation.
Moreover, the links can have directionality; that is, the distance or flux rate between two nodes
may not be the same in both directions. For example, if flux rate is influenced by gravitational
fields, then the “distance” between node A downslope and node B upslope may depend
dramatically on whether the link as from A to B or the reverse.

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In addition to the two basic elements of a graph (nodes and links), there are a number of other
attributes of the graph that warrant brief description because that are often utilized in the pattern
metrics:

• Path (or Walk) = any series of nodes in a graph.

• Cycle = closed Path of at least 3 nodes.
• Tree = Path that has no Cycles; note, there can be many different trees in a graph.
• Spanning Tree = Tree that connects all nodes in a graph.
• Minimum Spanning Tree = shortest Path of all Spanning Trees in a graph; in other words, the
ree that connects all nodes in a graph using the shortest possible total linkage distance.
• Component (or Cluster) = Nodes that are “connected” to each other.

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• Cut-node/Cut-link = a Node/Link that singularly disconnects a connected graph.

5.4. Pattern Metrics

Not surprisingly, there are many ways to measure the differences between graphs. Here, we
review some of the more common ones:

• Number = number of components (disjunct clusters) in a graph.

• Order = number of nodes in the largest component.
• Diameter = diameter of the largest component, where diameter is defined as the “distance”
between the two nodes furthest apart and the path taken is always the shortest (also called the
Maximum Eccentricity).
• Expected cluster size = area-weighted mean component (cluster) size, where the component
size is defined as the total area contained in the corresponding nodes.
• Correlation length = area-weighted mean component (cluster) radius of gyration, where the
radius of gyration is defined as the mean distance between each point or cell in a cluster and
the centroid of the cluster. Note, this measure is conceptually very similar to expected cluster
size, but uses the average distance traversed within a cluster as opposed to the absolute area
of the cluster. Thus, the unit of measurement is a distance instead of an area.

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• Connected graph = logical or binary indicator of whether all nodes are connected to all other
nodes or not.
• Node-connectivity = number of nodes that need to be removed to disconnect a connected
graph.
• Edge-connectivity = number of links that need to be removed to disconnect a connected
graph.

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It is important to remember that graph links can be defined on the basis of any measure of
ecological distance, univariate or multivariate, and it is this feature that provides great flexibility
to the graph matrix model.

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5.5. Applications

(1) Examining connectivity thresholds

The vast majority of applications have involved examining connectivity thresholds using a
mixture of both theoretical and empirical approaches. Let’s begin with a brief theoretical
example fro Urban and Keitt (2001). One way to explore connectivity thresholds is to
sequentially remove links by progressively decreasing the threshold distance for determining
whether nodes are connected or not – the so called “edge thinning” approach. In the hypothetical
example shown here, the graph is nearly connected at a threshold of 1500 m – there are but two
components (clusters). As the threshold distance is progressively decreased, the graph is
increasingly fragmented into a larger number of smaller components.

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The structural changes in the graph occurring during this edge thinning experiment are shown
here. Specifically, the figure shown here depicts the changes in the number of components
(number), number of nodes in the largest remaining component (order), and diameter of the
largest remaining component (diameter). As edges are progressively loss, moving from the right
to the left on the graph, at first there is no change in the metrics. This is because the edge-
connectivity of the graph is greater than one, which means that each node is connected by
multiple links to other nodes so that the loss of any one link does not effect the overall
connectivity of the graph. However, at some point the diameter of the largest component goes up
before eventually going down, which seems counter-intuitive at first. This happens because the
direct paths between nodes are lost at first- then stepping stones are lost. Thus, the cluster order
remains the same (i.e., no nodes are lost), but the shortest pathway between the two furthest apart
nodes (diameter) increases. As links continue to be lost, at some point there is a rather abrupt
change in all of the metrics. This threshold-like behavior reflects sudden and dramatic changes in
the connectivity of the graph as the connected graph begins to get fragmented into many smaller
components. Ecologically, the implications are obvious – there may be critical ecological
distances (e.g., dispersal distances) that once crossed dramatically reduce connectivity of the
landscape.

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An alternative way to explore connectivity thresholds is to sequentially remove nodes – the so
called “node thinning” approach. In the hypothetical example shown here, nodes were
progressively removed using three different approaches: (1) randomly, (2) the smallest nodes (in
terms of area), and (3) the end-node with the smallest area. The right-hand figure shows the
relationship between the number of nodes removed under the three removal methods and the
graph diameter. There were a couple of important findings from this study. First, the exact shape
of the resulting curves was landscape dependent, preventing generally applicable conclusions.
Second, in this particular landscape, end-node removal showed an advantage over the other
removal methods and exhibited threshold behavior, suggesting that node position in the
landscape can be important and that connectivity may change abruptly with the loss of critical
nodes.

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Now let’s consider an empirical example. In the example shown here, the graph matrix model
was used to examine connectivity thresholds in winter habitat of woodland Caribou in Manitoba,
Canada (O’Brien et al. 2006). Caribou, a threatened species in Canada, require lichen-rich,
mature conifer habitat, especially in winter. Past forestry practices have favored white-tailed deer
and moose over caribou, though caribou can adapt to some habitat fragmentation as long as
enough connected habitat is provided.

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In this study, the graph matrix model was used to depict the distribution of high-quality winter
habitat patches and the potential linkages between those patches using data from 20 radio-
collared animals. More specifically, nodes were defined as high-quality (lichen-rich jack pine and
sparsely treed rock) winter habitat patches >5 ha. It is worth noting that this approach essentially
treats all other land cover as non-habitat that is only traversed to get between habitat patches.
This may or may not be a reasonable assumption. Links were defined as the least-cost pathways
between habitat patches, where cost was based on a resource selection function derived using
logistic regression.

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The figure shown here depicts the relationship between expected cluster size (defined earlier),
interpreted as a measure of overall landscape connectivity, and the cost distance based on least
cost pathways. Several things are evident in this figure. First, habitat patches connect gradually
over small scales, with large increases in ECS occurring at cost distances of 800, 1600 and 2250.
These correspond to Euclidean distances of approximately 650 m, 1250 m, and 1750 m,
respectively. These thresholds represent scales where the graph includes sub-clusters connected
by only a few links into one much larger cluster. Second, at a cost distance of roughly 2500 m
(approximately 2000 m Euclidean distance), the graph is completely connected (i.e., all habitat
patches are connected into a single large cluster). Third, the curve for the late winter habitat use
is greater than the curve for all winter and very close to the curve for the maximum cluster size,
indicating that late winter animals utilize the largest available habitat clusters.

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The maps shown here depict the graph structure for thresholds corresponding to the important
transitions in ECS. Generally, clusters of well-connected high-quality habitat exist in the north,
central and southeast regions of the winter home range. For cost distances of up to 800, clusters
of high-quality habitat in these areas form gradually where patches are aggregated. Above a cost
distance of 800, two large clusters in the central region of the winter home range link together,
corresponding to the large increase in ECS at this threshold, while clusters in the north and south
remain isolated. Including links up to a cost distance of 1600 joins the northern cluster with the
central cluster resulting in another large and rapid increase in ECS. At a cost distance of 2250 the
large southern cluster coalesces with the main central cluster. At this threshold the ECS
approaches the maximum, but small clusters in the northwest and central portions of the winter
home range remain isolated.

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Lastly, O’Brien et al. (2006) compared the graph structure for the current winter range with the
historical winter range. In both the Owl Lake and Kississing ranges, expected cluster size values
increased more rapidly at short cost distances (0–2000 units) in the winter home range than in the
historical home range. High-quality habitat patches within the winter home range are spaced
more closely together and coalesce into clusters more rapidly than patches in the historical home
range.

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(2) Identifying critical linkages

A closely related application of graph theory involves not only examining connectivity thresholds
but identifying the critical linkages and/or nodes associated with threshold changes in
connectivity. In the classic study by Keitt et al. (1997), the authors used the graph matrix model
to represent habitat patches of the Mexican spotted owl in Colorado, Utah, Arizona and New
Mexico. The links were simply Euclidean distances between patches.

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In the following series of figures, habitat patches are depicted in yellow and the links are in
purple. In the first figure, all links < 10 km are highlighted. The graph is highly disconnected.
The second figure depicts all links <30 km. The graph is more connected, but still largely
comprised of several separate clusters. The next three figures depict the graph structure for
threshold distances of 50, 70 and 90 km. By definition, the correlation length of the graph
(defined above) increases monotonically with increasing threshold distance. However, the graph
shows a distinct threshold in the vicinity of 40 km, where correlation length almost doubles as
the threshold distance increases slightly. Interestingly, 40 km corresponds roughly to the
dispersal distance of juvenile spotted owls.

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7.86
The figure shown here depicts the graph sensitivity at different threshold distances. Patches are
ordered by size. Sensitivity is the change in correlation length when a patch was removed. Not
surprisingly, at pretty much all threshold distances the largest patches are the most sensitive. That
is because they constitute a disproportionately large area and their removal results in a significant
change in correlation length. However, at intermediate threshold distances (40-50 km), several
smaller patches showed significant sensitivity, indicating that their position in the landscape
makes them particular important as stepping stones between large clusters and that their removal
has a disproportionate impact on graph connectivity as measured by correlation length.

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This figure portrays the results of the sensitivity analysis in map form. The left-hand figure
depicts the importance index of each patch averaged over all threshold distances between 0-100
km. Not surprisingly, the few largest patches have the highest overall absolute importance; that
is, they are essential to the connectivity of the graph. However, the right-hand figure depicts the
importance index on a per unit area basis, revealing that a few relatively small patches have a
very high importance due largely to their position in the landscape. By connecting two large
clusters to the north and south, they serve a critical role as stepping stones at threshold distances
of 40-50 km. These particular habitat patches might be the logical focus of conservation efforts
aimed at maintaining habitat connectivity for spotted owls.

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5.6. Pros and Cons

Considering the information presented thus far, what are the strengths and limitations of the
graph matrix model of landscape structure?

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