Progress reports

Landscape ecology and

biogeography: Rethinking
landscape metrics in a
post-FRAGSTATS landscape

Progress in Physical Geography

36(3) 400420
The Author(s) 2012
Reprints and permission:
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DOI: 10.1177/0309133312439594
ppg.sagepub.com

John A. Kupfer
University of South Carolina, USA

Abstract
Landscape pattern indicators or metrics provide simple measures of landscape structure that can be easily
calculated with readily available data and software. Unfortunately, the ecological relevance of many metrics
(i.e. the relationship between metric values and the real-world ecological processes that they are meant to
serve as proxies for) is often unproven and questionable, and concerns are regularly voiced that such metrics
fail to capture important aspects of landscape function. In this paper, I provide a review of landscape measures
that may better link landscape pattern and function, ranging from approaches that extend existing metrics by
incorporating a more functional component (e.g. core area measures, least cost distances) to those rooted in
graph, network, and electrical circuit theory. While more functional approaches are becoming increasingly
popular, the selection of appropriate landscape metrics in many applications involves tradeoffs regarding data
requirements, ease of calculation, functional basis, and simplicity of interpretation by a range of specialist and
non-specialist stakeholders. Regardless, there continues to be a need for landscape metrics because they are
seen by many land managers and stakeholders as simple, intuitive tools for assessing and monitoring changes
in landscape pattern and, by extension, the effects on underlying ecological processes. Future needs include:
(1) the development of more user-friendly landscape analysis software that can simplify graph-based analyses
and visualization; and (2) studies that clarify the strengths and weaknesses of different approaches, including
the potential limitations and biases in graph and network-based measures.
Keywords
circuit theory, graph theory, habitat network, landscape pattern indicator, least cost distance, scale

The ability to quantify landscape structure is prerequisite to the study of landscape function and
change. For this reason, much emphasis has been
placed on developing methods to quantify landscape structure . . . This report describes a program
called FRAGSTATS that we developed to quantify
landscape structure. FRAGSTATS offers a comprehensive choice of landscape metrics and was
designed to be as versatile as possible. The program
is almost completely automated and thus requires
little technical training. (McGarigal and Marks,
1995: 2)

(E)cological relevance of landscape indices is more

often presumed than established, with inadequate
supporting empirical evidence in the literature. In the
absence of such evidence, landscape indices are
mathematical constructs that have no inherent ecological meanings. The validity of landscape analysis

Corresponding author:
Department of Geography, 709 Bull Street, Columbia, SC
29208, USA
Email: kupfer@sc.edu

Kupfer
will increase if indices are selected according to
their ecological relevance rather than the convenience of computer programs. (Li and Wu, 2004:
391)

I Introduction
A central tenet of landscape ecological theory is
that the spatial pattern of organisms, populations, and ecosystems across a landscape
reflects the influence of underlying gradients
and processes but in turn acts to shape ecological processes such as dispersal, competition,
disturbance, and fluxes of energy and matter
across space (Kupfer, 2011). This reciprocal
relationship between landscape pattern and process has made the quantification of landscape
structure (the composition and spatial configuration of ecological entities such as habitat
types) one of the most fundamental pursuits in
landscape ecology. Technological advances in
remote sensing and geographic information systems have played a significant role in advancing
the availability and analysis of geospatial data
over the last three decades, but it could be
argued that it was the release of the FRAGSTATS software analysis package nearly 20
years ago that helped to revolutionize the analysis of landscape structure and firmly entrench
landscape pattern indices or landscape metrics
in the minds and statistical tool boxes of many
landscape ecologists and biogeographers.
The two primary releases of FRAGSTATS
(McGarigal and Marks, 1995; McGarigal et
al., 2002) have now been cited thousands of
times, and many of the original FRAGSTATS
metrics have been incorporated into other
widely used stand-alone and GIS-integrated
landscape analysis packages.
Landscape metrics are quantitative indices
that describe compositional and spatial aspects
of landscapes based on data from maps, remotely
sensed images and GIS coverages. Typically,
landscape elements are defined as discrete entities or patches, and landscape pattern is described

401

using metrics developed to quantify patch- (e.g.

size, shape, isolation) and mosaic- (patch richness and diversity, connectivity, contagion) level
characteristics. Ideally, the goal of landscape
analyses should then be to link measures of landscape structure to specific effects on ecological
processes, rather than treating the quantitative
description of spatial pattern as an end unto itself
(e.g. ONeill et al., 1997). Haines-Young and
Chopping (1996) and McGarigal et al. (2002)
provide comprehensive reviews of common
metrics that are included in landscape analysis
packages, including FRAGSTATS (see also
Table 1).
While FRAGSTATS-style metrics continue
to be widely used, they have well-documented
limitations that have led some to question
whether many can be related to real-world ecological processes in a predictable manner (Cardille
et al., 2005). Metric values are sensitive to data
resolution (Moody and Woodcock, 1995), study
area extent (Turner et al., 1989), and thematic
resolution of the input data (Huang et al.,
2006), and even what are regarded as acceptably
low misclassification rates in the source data can
be magnified into substantial errors in metric values (Langford et al., 2006; Shao and Wu, 2008).
Such limitations can often be addressed, or put in
perspective, through careful data manipulation,
analysis and interpretation. More fundamental,
however, are limitations related to bridging the
disconnect between spatial pattern and ecological process, and thereby establishing the ecological relevancy of landscape metrics. Correlation
analysis involving landscape metrics is hampered by the difficulty of replicating large-scale
experiments and the complicated responses of
metrics to changes in scale and spatial pattern
(Li and Wu, 2004). Relationships between some
metrics and ecological patterns or processes may
be confounded by interactions with other attributes of the landscape (Hargis et al., 1999).
Quantifying the unique effects of habitat configuration on biotic responses, for example, is difficult because many widely used configuration

402

Perimeter-Area Ratio
Shape Index/Fractal Dimension
Index
Linearity Index
Related Circumscribing Circle
Contiguity Index
Core Area
Number of Core Areas
Core Area Index
Average Depth Index
Maximum Depth Index
Proximity Index
Similarity Index
Euclidean Nearest Neighbor
Distance
Functional Nearest Neighbor
Distance
Edge Contrast Index

Shape

Contrast

Isolation/Proximity

Core Area

Patch Area
Patch Perimeter
Radius of Gyration

Patch

Area/Density/Edge

FRAGSTATS
METRICS

Total Core Area

Number/Density of Disjunct Core Areas
Core Area Distribution
Disjunct Core Area Distribution
Core Area Index Distribution
Proximity Index Distribution
Similarity Index Distribution
Euclidean Nearest Neighbor Distance
Distribution
Functional Nearest Neighbor Distance
Distribution
Contrast-Weighted Edge Density
Total Edge Contrast Index
Edge Contrast Index Distribution

Percentage of Landscape
Number of Patches/Patch Density
Total Edge/Edge Density
Landscape Shape Index
Largest Patch Index
Patch Area Distribution
Radius of Gyration Distribution
Perimeter-Area Fractal Dimension
Perimeter-Area Ratio Distribution
Shape Index/Fractal Index Distribution
Linearity Index Distribution
Contiguity Index Distribution

Class

Table 1. Representative landscape metrics calculated by FRAGSTATS v3.3 (McGarigal et al., 2002)

(continued)

Total Core Area

Number/Density of Disjunct Core Areas
Core Area Distribution
Disjunct Core Area Distribution
Core Area Index Distribution
Proximity Index Distribution
Similarity Index Distribution
Euclidean Nearest Neighbor Distance
Distribution
Functional Nearest Neighbor Distance
Distribution
Contrast-Weighted Edge Density
Total Edge Contrast Index
Edge Contrast Index Distribution

Perimeter-Area Fractal Dimension

Perimeter-Area Ratio Distribution
Shape Index/Fractal Index Distribution
Linearity Index Distribution
Contiguity Index Distribution

Number of Patches/Patch Density

Total Edge/Edge Density
Landscape Shape Index
Largest Patch Index
Patch Area Distribution
Radius of Gyration Distribution

Landscape

403

Connectivity

Diversity

Patch

Contagion/
Interspersion

FRAGSTATS
METRICS

Table 1. (continued)

Percentage of Like Adjacencies

Aggregation Index
Interspersion and Juxtaposition Index
Mass Fractal Dimension
Landscape Division Index
Splitting Index
Effective Mesh Size
Patch Cohesion Index
Connectance Index
Traversability Index

Class

Percentage of Like Adjacencies

Contagion
Aggregation Index
Interspersion and Juxtaposition Index
Landscape Division Index
Splitting Index
Effective Mesh Size
Patch Cohesion Index
Connectance Index
Traversability Index
Patch Richness/Patch Richness Density
Relative Patch Richness
Shannons and Simpsons Diversity Indices
Shannons and Simpsons Evenness Indices

Landscape

404

metrics are correlated with habitat extent (Smith

et al., 2009; Wang and Cumming, 2011).
In spite of these problems, landscape metrics
continue to be widely used because they are
seen as simple, intuitive tools for assessing
and monitoring changes in landscape pattern
and, by extension, the effects on underlying
ecological processes. Landscape metrics that
quantify spatial patterns of deforestation and
fragmentation may be linked with measures of
species persistence to elucidate fragmentation
effects and develop tools and indices that can
be used to aid forest management (Baskent and
Jordan, 1996; Ripple et al., 1991). Because the
data needed to calculate metrics are easy to
obtain from satellite imagery, they provide a
means for monitoring landscape change and
serve as perhaps the simplest proxies for estimating aspects of species and community health
in heterogeneous settings.
Li and Wu (2004: 395) nonetheless noted
eight years ago that: After two decades of
extensive research, interpreting indices
remains difficult because the merits and
caveats of landscape metrics remain poorly
understood. What an index really measures
is uncertain even when the analytical aspects
of most indices are quite clear. Not much has
changed in the last decade, as consistent generalizations regarding pattern-process relationships for many metrics remain elusive
(Turner, 2005). The good news is that advances
in spatial analysis, modeling and metric development have facilitated the creation of
new landscape measures and approaches to
linking landscape pattern and function. A number of these indices are meant to more adequately or flexibly address one of the most
significant sources of problems with the first
generation of landscape metrics: their inherent
focus on landscape structure, rather than landscape function.
The purpose of this paper is to summarize
some of these advances. While landscape
metrics have been developed to describe aspects

Progress in Physical Geography 36(3)

of patch edge, shape, diversity and evenness,

contagion and interspersion, and contrast
(McGarigal et al., 2002), I will focus primarily
on metrics associated with patch area, isolation,
and connectivity, because: (1) the linkages
between such measures and ecological processes are often perceived to be relatively clear,
and (2) such metrics are most commonly used to
quantify the structural changes associated with
forest loss and fragmentation (Kupfer et al.,
2006), one of the most typical applications of
landscape metrics. Further, while metrics have
been used to quantify spatial patterns at a broad
range of spatial scales (Kupfer, 2006), I will
focus on their usage at the scale at which
pattern-process linkages are often of greatest
management interest, that of human-observed
landscapes (e.g. 10s1000s of km2: Forman and
Godron, 1986).

II Putting the fun(ction) in

landscape metrics
At the end of a landscape ecology seminar that I
taught in 2001, a doctoral student gave me a cartoon that humorously depicted a pitfall of poorly
designed landscape analyses, that is, a failure to
question the ecological meaning of metric
values (Figure 1). By lowering technological
barriers, FRAGSTATS and related packages
allow users to calculate dozens of measures
of landscape structure, but the generation of
such information has outpaced our understanding of how such measures relate to actual ecological processes (e.g. Herzog and Lausch, 2001).
In particular, the assumption that metrics which
quantify landscape structure capture functional
landscape properties is perhaps too uncritically
accepted, and the ecological relevance of many
landscape indices is often unproven and
questionable.
The most basic measures of landscape structure, those addressing patch area and isolation,
stem from island biogeography theory (IBT),
which states that the number of species on

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405

Figure 1. Technological advances have facilitated the generation of dozens of measures of landscape structure, but the amount of information produced has often outpaced our understanding of how such measures
relate to ecological processes.
Source: T. Crimmins (personal communication, 2001).

oceanic islands is a function of island size

(because of its effects on extinction rates and
habitat heterogeneity) and distance from the
mainland (which influences the arrival of potential colonists) (MacArthur and Wilson, 1967).
Principles from IBT have since been applied
to the design of nature reserves, and the conceptual importance of patch area, connectivity, and
isolation remains central to contemporary ecological theories, including those associated with
metapopulations, source-sink dynamics, and
metacommunities (Kupfer, 1995).
Efforts to link landscape pattern and its
change through time to biotic responses have
most commonly used metrics such as: (1) patch
area, edge density and nearest neighbor distance, at the individual patch level; or (2) mean
patch size, largest patch index, mean nearest

neighbor distance, or cohesion/juxtaposition,

at the class level (e.g. Magness et al., 2006; Patterson and Malcolm, 2010; Scharine et al.,
2009). Unfortunately, while field and simulation studies have documented that habitat area
and landscape connectivity influence the presence and dispersal of species and the persistence
of metapopulations, it is less clear whether
many landscape metrics capture such effects
and can be unambiguously linked to functional
properties of patches or the landscape mosaic
(Lindenmayer et al., 2002; McAlpine and Eyre,
2002). Alternative approaches have thus been
advocated for incorporating greater functionality into landscape metrics.
The simplest method for doing so is to add a
more functional component to existing metrics.
Core area, for example, quantifies the extent of

406

patch area remaining after eliminating a specified edge buffer (Laurance, 1991), the width
of which is defined on the basis of a functional
property. The depth of edge influence following
forest removal and edge creation has been
defined for a range of primary responses (e.g.
changes in microclimate, litterfall, nutrient
cycling, decomposition; damage to vegetation;
increased seed and pollen dispersal) and secondary responses (e.g. alterations in sapling
density, understory cover, shrub height, community composition, and non-native species
abundance) (Harper et al., 2005; Laurance et
al., 2002). Core area has been shown to be a useful predictor of the presence and abundance of
area-sensitive species (Austen et al., 2001),
although the specific nature of ecological
responses may be complicated by the broaderscale landscape context of a patch (Mazerolle
and Villard, 1999; Smith et al., 2011). Further,
core area is sensitive to errors in the specified
depth of edge influence, ignores heterogeneity
inherent in the edge to interior transition, and
does not account for the dynamic aspects of
some edge effects (Kupfer and Runkle, 2003).
Nearest neighbor distance is perhaps the most
commonly used isolation metric, yet it is often
poorly related to population properties (e.g.
size, persistence) and ecological processes
(e.g. dispersal) and is sensitive to sample sizes
and variations in patch size and shape (Bender
et al., 2003; Moilanen and Nieminen, 2002).
There are several reasons for these shortcomings (Calabrese and Fagan, 2004), one of which
is that the movement of organisms is determined
by a landscapes functional connectivity for
each species, which is a factor of more than distance. The most widely used alternative for
assessing inter-patch distances is least cost distance (Adriaensen et al., 2003). A landscape is
represented as a grid in which each cell is
assigned a resistance value based on the cost
that it imposes on species movement. Cost
values reflect variables that are relevant for a
particular species such as vegetation cover

Progress in Physical Geography 36(3)

type and structure, human infrastructure, and

topography. Estimates of resistance values are
typically based on expert opinion, but more
recent efforts have used telemetry or trapping
data, measures of relative gene flow, or behavioral data (Epps et al., 2007; Rayfield et al.,
2010; Sawyer et al., 2011). Once a resistance
surface has been calculated, the values are used
to identify a path that minimizes cumulative
costs between locations. A number of studies
have demonstrated that least cost distance measures may be better predictors of dispersal and
patch occupancy (Chardon et al., 2003; Sutcliffe
et al., 2003; Verbeylen et al., 2003) and genetic
patterns (Coulon et al., 2004; Hokit et al., 2010)
than Euclidean distances, although such relationships may be stronger for habitat specialists
than generalists (Shanahan et al., 2011).
A second area of advancement in making
existing landscape metrics more functional
involves the treatment of spatial scale. Ecological patterns and processes show variability
on a range of spatial, temporal, and organizational scales, and there is no single scale at
which ecological phenomena should be studied (Levin, 1992). Spatial scale thus poses a
problem for many landscape metrics because
their values are based on static classifications
of land-cover data that are provided at scales
that may be inappropriate for the organisms
or processes in question. For example,
attempts at linking landscape metrics to wildlife responses to fragmentation have often
been equivocal (Tischendorf, 2001), in part
because metrics may capture landscape features that are irrelevant to how species perceive and utilize the landscape (Cale and
Hobbs, 1994; Lindenmayer et al., 2002). Consequently, scale-oriented approaches to landscape metric calculation and analysis that
have potential relevance for linking the usage
of metrics and indicator species in evaluations
of environmental change and biodiversity loss
(e.g. Banks-Leite et al., 2011; Lindenmayer
and Likens, 2011) have great potential value.

Kupfer

One regularly used approach to addressing

scale involves examining how metric values
change with changes in resolution and extent.
Recognizing that organism-specific behavioral
and perceptual responses to landscape structure
should guide the delineation of habitat patches
during landscape analysis, Girvetz and Greco
(2007, 2009) developed and implemented an
algorithm that delineated patches across a range
of spatial scales based on organism-specific
thresholds in land-cover density, gaps, and habitat patch thickness. Their results confirmed that
the spatial scale at which habitat characteristics
were measured influenced estimates of patch
suitability.
Taking a more functional approach to linking
spatial patterns with population dynamics and
viability at scales relevant to organisms, Vos
et al. (2001) proposed the use of ecologically
scaled landscape indices (ESLIs) such as average patch carrying capacity and average patch
connectivity, which were defined on the basis
of species traits. A trait-based approach to
developing pattern-process relationships related
to area and connectivity implicitly recognizes
functional aspects of how organisms perceive
and respond to their environment, and has since
been shown to be useful for a range of organism
types (Gehring and Swihart, 2003; Goheen et
al., 2003; Meyer et al., 2008; Schleicher et al.,
2011).

III From landscape mosaics to

landscape graphs
Driven by a desire for a more functional
approach to landscape quantification, landscape
ecologists have turned to the use of graph theory. In general, graphs conceptualize an entity
as a set of points or nodes connected by edges
or links; when node locations and connections
are tied to specific spatial locations, the resulting feature is termed a spatial graph (Fall et
al., 2007). In landscape ecological applications,
a landscape is represented as a set of nodes

407

(habitat patches) that are connected by links

(e.g. via dispersal), resulting in a landscape
graph (Cantwell and Forman, 1993). The use
of landscape graphs to examine patterns of connectivity, identify potential dispersal pathways,
and prioritize habitat patches for conservation
has increased rapidly over the last decade
because graphs have flexible data requirements
and are efficient for characterizing landscape
connectivity at broad spatial scales with many
habitat patches (Rayfield et al., 2011; Treml et
al., 2008). Graphs also provide a useful tool for
examining the dynamics of metapopulations
and metacommunities by linking patterns of
individuals, populations, and habitats with
behavior and dispersal (Dale and Fortin, 2010;
Economo and Keitt, 2008).
Metrics used to quantify graph properties can
be classified by the structural level of the graph
to which they apply and the aspect of the element that they quantify (Baranyi et al., 2011;
Rayfield et al., 2011). Many commonly used
measures relate to characteristics of nodes and
links and their role in network pattern, including: (1) the number of linkages connected to a
given node (node degree), which serves as an
indicator of node accessibility and how important the node is to overall connectivity (e.g.
Benedek et al., 2011; Cantwell and Forman,
1993; Jordan et al., 2007); and (2) node centrality (e.g. betweenness centrality, closeness centrality, subgraph centrality), which quantifies
the importance of a nodes position in paths
between other pairs of nodes (Bodin and Norberg, 2007; Economo and Keitt, 2010; Estrada
and Bodin, 2008; Newman, 2005). Weights may
be assigned to nodes or links on the basis of qualitative or quantitative characteristics (e.g. for
nodes: measures of habitat area or quality; for
links: the flow of organisms or probability of
movement) to better represent their relative
contributions to habitat provision and maintenance of connectivity (Jordan et al., 2007; Lookingbill et al., 2010). Such measures can be used
to approximate node or dispersal flux,

408

movements into or out of a patch, or movement

between pairs of habitat patches (Minor and
Urban, 2007; Urban and Keitt, 2001).
A second set of graph metrics quantifies
aspects of the entire graph or subgraph components (sets of nodes connected to each
other but disconnected from parts of the
larger graph). Examples include measures
of: (1) the number or density of nodes, links
or components (Brooks et al., 2008; Bunn et
al., 2000; Jordan et al., 2003), which provides
information on the number of habitat patches
and their degree of linkage; (2) how component or network properties change with node
or link removal (Bodin and Norberg, 2007;
Cantwell and Forman, 1993; Treml et al.,
2008; Urban and Keitt, 2001), an indicator
of vulnerability to patch or link removal
through habitat loss; and (3) component/
graph diameter and path lengths (Economo
and Keitt, 2008; Ferrari et al., 2007; Minor
and Urban, 2008), which describe the density, compactness, and potential ease of
movement. Indices developed by Saura and
colleagues assess habitat availability by integrating habitat area and connectivity into a
single measure (class coincidence probability
and integral index of connectivity: PascualHortal and Saura, 2006; probability of connectivity: Saura and Pascual-Hortal, 2007;
equivalent connected area: Saura et al.,
2011a). Subsequent work has coupled such
indices with centrality measures (Bodin and
Saura, 2010) and partitioned values into separate components to quantify the ways in
which a specific patch may contribute to
habitat connectivity and availability (Saura
and Rubio, 2010). This latter approach, in
particular, may be valuable for identifying
the best and highest-quality habitat areas
regardless of their position within a landscape network.
The use of landscape graphs and their associated metrics offers an opportunity to move
beyond the emphasis on structural properties

Progress in Physical Geography 36(3)

inherent in most traditional landscape metrics,

but the degree to which graphs do so is dictated
by the way in which nodes and links are defined
(Rayfield et al., 2011). Structural connectivity
can be assessed simply by analyzing information
about the physical adjacency of habitat patches
(topology) or the Euclidean distances among
them. However, the relationship of such measures
to ecological pattern and processes (e.g. interpatch movement) still needs to be established,
and whether graph measures provide a better
means for doing so than traditional landscape
metrics should be tested. By incorporating estimates of mobility derived from species traits or
mark-recapture studies, or through the use of
species-specific least cost surfaces, it becomes
possible to quantify potential connectivity, a
proxy for how connected a given landscape or
patch will be for a focal species (Calabrese and
Fagan, 2004). For example, data on the maximum
dispersal distance threshold of a species or modeled dispersal patterns may be used to eliminate
links that fail to meet certain criteria and produce a representation of potential connectivity
for a focal species (Brooks et al., 2008; OBrien
et al., 2006) (Figure 2). Observed movement
patterns of individuals among habitat patches
or through a landscape (e.g. from radiotelemetry) can be used to quantify actual connectivity.
These more direct measures explicitly account
for the availability of potential paths among
nodes in a network (Matisziw and Murray,
2009), but they require data that is often prohibitively time-consuming or expensive to collect for a large number of individuals.

IV Oceans, islands and all things in

between
In our 2006 paper, Not seeing the ocean for
the islands: The mediating influence of
matrix-based processes on forest fragmentation
effects, my colleagues and I highlighted a
growing awareness of the need to consider not
only the characteristics of habitat patches in

Kupfer

409

Figure 2. Graph theoretic representation of 138 potential habitat patches for the Delmarva fox squirrel
(Sciurus niger cinereus) and their connections based on simulated dispersal events. For each patch, 100,000
dispersers were released and successful transfers from source to destination patches were tracked. Figures
represent different thresholds of connectedness as determined by inter-patch flux rates; two patches are
assumed to be connected if the flux rates are above these thresholds. The flux rates are (A) > 0; (B) >
1000; (C) > 2000.
Source: Data courtesy of T. Lookingbill.

assessing forest loss and fragmentation effects

(islands, in the island biogeography metaphor), but also characteristics of the intervening
human-dominated matrix (the ocean) (Kupfer
et al., 2006). The underlying message was that
the matrix may neither be uniformly unsuitable
nor serve as a fully absorbing barrier to the dispersal of certain forest taxa. This point has been
made elsewhere (e.g. Ewers and Didham, 2006;
Fischer and Lindenmayer, 2007; Ricketts, 2001;
Vandermeer and Carvajal, 2001), and the growing number of papers confirming the significance of matrix-related processes attests to a
recognized need for quantifying matrix patterns
and properties (Bender and Fahrig, 2005;

Prevedello and Vieira, 2010; Umetsu et al.,

2008; Watling et al., 2011).
Efforts to extend or improve landscape
metrics aimed specifically at quantifying spatial
patterns across the entire landscape, including
the matrix, have generally addressed one of two
issues. The first involves the influence of the
matrix on the length, relative quality, and redundancy of dispersal routes connecting habitat
patches (e.g. Pinto and Keitt, 2009). The movement of organisms across a landscape is determined by factors that are not easily captured
by traditional landscape metrics, including
matrix permeability, the presence of stepping
stones, the width and quality of corridors, and

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the willingness of a species to cross gaps in

habitat (e.g. Belisle, 2005; Uezu et al., 2007;
Vos et al., 2007). The most common method for
incorporating the effects of the matrix, or, more
broadly, non-habitat patches, on movement
involves the use of resistance surfaces to calculate cost paths (as described earlier). Once calculated, least cost distances can be directly
incorporated into landscape graphs as an alternative to Euclidean distance for defining link
weights and analyzing habitat connectivity
(Chetkiewicz et al., 2006). The subsequent
graph measures may thus reflect the effects of
matrix traversability if the shortest path passes
through the matrix.
An inherent risk of this approach is that simply replacing nearest neighbor Euclidean distances with least cost distances may still fail to
adequately capture landscape connectivity. This
may occur if the between-patch movement
potential is equated with the distance of a single
path between nodes, even though individuals
rarely use a single optimum route (Driezen et
al., 2007). Recent attempts at representing
habitat networks have therefore involved treating links and pathways as multiple routes or corridors, rather than single, least cost routes. Two
examples of such methods are the conditional
minimum transit cost method (Pinto and Keitt,
2009; Theobald, 2006) and the minimum shortest paths method (Pinto and Keitt, 2009),
which identify a set of routes between pairs
of habitat patches and retain all paths shorter
than a user-specified threshold, for example,
all paths within 10% of the least cost route path
(Urban et al., 2009).
An alternative approach for analyzing crossmatrix connectivity involves the use of concepts
borrowed from electrical circuit theory (McRae
et al., 2008). Circuits are graphs in which relative flow rates between nodes are quantified and
visualized based on resistance, in this case, the
opposition of a habitat type to an organisms
movement, or conductance, the facilitation of
movement through a cell or along a graph edge

Progress in Physical Geography 36(3)

relative to others available to it (McRae et al.,

2008). Ease of movement between nodes can
be quantified using measures such as effective
resistance (a measure of isolation between pairs
of nodes that incorporates multiple pathways),
which can then be related to individual movement patterns and gene flow (e.g. McRae and
Beier, 2007). Measures of cumulative flux
between habitat patches distributed over multiple routes or link redundancy, which indicates
the number of parallel pathways between a pair
of nodes, can also be calculated (Rayfield
et al., 2011).
Beyond its role as movement conduit, filter
or barrier, the matrix may influence populations
in forest remnants if it provides an increased
resource base by enlarging foraging or breeding
area (Kupfer et al., 2006). Because most traditional landscape metrics are calculated after
categorizing landscape elements into discrete,
homogeneous units, broader neighborhood
effects are typically ignored. A number of
researchers have advocated reframing how
landscapes are conceptualized and pushed
for raster-based analyses that better capture gradients in environmental resources and habitat
quality (Kent, 2009; Murphy and Lovett-Doust,
2004). Rather than being represented as discrete
patches within a large mosaic, landscapes may,
for example, be represented as a continuous
surface of a habitat variable (e.g. forest cover)
(McIntyre and Hobbs, 1999). A grid-based
landscape representation recognizes that areas
of habitat also exhibit variability in conditions
and suitability.
One approach to quantifying landscape patterns as continuous surfaces has involved the
use of moving windows, in which each cell in
the landscape is assigned a value or a category
based on values of all cells within a kernel centered on the cell of interest. This approach can
be computationally efficient for large areas,
effective at capturing the context of a point relative to larger landscape neighborhood effects,
and useful for examining the effects of scale

Kupfer

on forest patterns. FRAGSTATS calculates a

range of class and landscape metrics within
moving windows, and Riitters and colleagues
have used moving window analyses to quantify
several aspects of landscape pattern (e.g. Riitters and Coulston, 2005; Riitters et al., 2009).
While such measures are useful for summarizing the extent or pattern of focal cover types
within the windows (e.g. forests or natural
covers), efforts to create landscape metrics
that more broadly quantify characteristics of
the matrix and the landscape as a whole, and
particularly those that can be related to landscape function, require further improvements.
The development of such metrics would be
particularly useful given advances in species
distribution modeling that facilitate the creation of continuous grids of habitat suitability
(Miller, 2010).
The other primary approaches to analyzing
patterns of continuous variations in landscape
characteristics involve geostatistical techniques.
Measures of spatial autocorrelation and pattern
are now regularly used in landscape ecology
and biogeography (e.g. Vaclavk et al., 2012),
and McGarigal et al. (2009) introduced a variety
of metrics for quantifying landscape gradients
from the field of surface metrology, the analysis of three-dimensional surfaces. These surface metrics, which captured structural
aspects rather than functional components of
the landscape, described aspects of surface
roughness, height and texture. While some
surface metrics had strong analogs among
more traditional patch metrics, others provided
unique measures of landscape pattern. As with
moving window approaches, though, more
research is needed to determine the ecological
relevance of these measures.

V Are metrics just fun tools for

landscape ecologists?
In her recent paper, Spatial metrics useful indicators for society or mainly fun tools for

411

landscape ecologists?, Dramstad (2009) examined the extent to which traditional landscape
metrics have been linked to ecological processes
and questioned their potential as indicators for
managers and policy-makers. She argued that
certain issues still need to be better resolved
before many spatial metrics can provide reliable
guidance for spatial planning and land management, including solidifying the links between the
ecological patterns revealed by metrics and the
processes we want them to indicate. Sawyer et
al. (2011) give a comparable example regarding
the use of least cost distance models, stating:
Conservation planners are faced with a critical
question: will such models improve placement of
linkages/corridors by explicitly incorporating habitat effects on movement, or will they result in misleading and potentially costly recommendations
for conservation by concealing invalidated assumptions? (Sawyer et al., 2011: 669)

Beyond questions about the ability of structural

metrics to capture functional landscape attributes, an additional concern is how easily landscape measures can be interpreted by those
responsible for conservation planning and land
management. Most stakeholders and decision
makers want measures that are transparent,
easy to understand, and inexpensive to measure
(Gustavson et al., 1999). Landscape pattern
indicators thus need to be not only ecologically
relevant but also easily interpretable by a range
of non-scientists, including politicians, land
managers and, in some cases, the public. In reality, the selection of appropriate landscape
metrics to facilitate effective land management
and site prioritization in conservation plans
often involves tradeoffs regarding three factors:
(1) data requirements and ease of metric calculation; (2) their basis in structural versus functional properties of a landscape; and (3) their
ease of interpretation by a range of specialist
and non-specialist stakeholders.
Simple structural metrics that quantify patch
(e.g. patch area, nearest neighbor distance) or

412

landscape (e.g. patch cohesion) properties independent of any specific process have been popular because they can be easily calculated with
readily available land-cover data and software,
require little or no additional parameterization,
and often have intuitive interpretations (Figure 3).
However, given the limitations of such measures
and questions regarding their ecological relevance, these metrics may be best suited for
exploratory and descriptive analysis. At the other
extreme, approaches utilizing spatially explicit
population models consider population dynamics
resulting from birth, mortality, emigration, and
immigration processes in individual patches and
are appropriate when analyses require an assessment of spatiotemporal population trends and
persistence. However, such approaches are computationally intensive and require extensive
knowledge of species biology and behavior, limiting their application in many cases to small
study areas (Minor and Urban, 2007).
Node, link, component, and network measures rooted in graph, network, and circuit theory represent a middle ground in that
they typically require more information and
user-based parameterization than structural
landscape metrics, but they can better bridge the
gap between structure and function. Further,
graph-based applications may make similar
predictions to and offer insights not available
from spatially explicit population models while
having less stringent data requirements (Minor
and Urban, 2007). The tradeoff is that many
element-based (e.g. node degree, centrality,
resistance distance) or network-based (e.g.
area-weighted flux, graph diameter, characteristic path length, probability of connectivity)
graph measures are more difficult to calculate
and interpret than commonly used structural
metrics such as patch area or nearest neighbor
distance (Figure 3).
The nature of tradeoffs among data requirements, ease of calculation, functional basis,
and simplicity of interpretation changes with
theoretical and technological advances.

Progress in Physical Geography 36(3)

Figure 3. Conceptual diagram of landscape metric

tradeoffs with respect to: (1) ease of implementation, which reflects data requirements and the
availability and simplicity of software to calculate
the metric; (2) structural versus functional basis;
and (3) ease of interpretability by scientists and
non-scientist stakeholders. Measures include (A)
patch/element metrics and (B) landscape/network
metrics. (Metric abbreviations: NND Euclidean
nearest neighbor distance; LCD least cost distance; Betw. Centrality betweenness centrality;
Resist. Dist. resistance distance; AWF areaweighted flux; Graph Diam. graph diameter;
Char. Path Len. characteristic path length;
PC probability of connectivity; Surf. Metrol.
surface metrology.)

Kupfer

413

Table 2. Software packages that have been used for landscape graph analysis. A number of other software
packages and tools (e.g. social network analysis packages, R libraries) have been developed to quantify aspects
of graph properties but are not included here because they lack the ability to directly analyze landscape
graphs. All websites were current as of 28 November 2011.
Software

Description

Download available

Citation

Conefor
Calculates node-, link- and graph-based
http://
Sensinode
metrics, including: number of links, number www.conefor.org
of components, Harary Index, class and
landscape coincidence probability, integral
index of connectivity, flux and areaweighted flux, probability of connectivity

Saura and Torne

(2009)

JMatrixNet

Identifies a network of habitat patches within http://www.ecology.

a landscape that can then be analyzed using su.se/jmatrixnet
network software (e.g. Pajek) to calculate a
limited number of patch and network
measures

Bodin and Norberg

(2007)

FunConn

A modeling toolbox for ArcGIS v9.3 that

http://warnercnr.
calculates minimum spanning trees and
colostate.edu/
shortest paths, and provides a range of link,
*davet
node, and network-based operations

Theobald et al. (2011)

SELES

A structured language for modeling

http://seles.info
landscape dynamics that includes methods
for calculating minimum planar graphs and
spanning trees

Fall and Fall (2001);

Fall et al. (2007)

LQGraph

Optimizes the connectivity of sites

http://uts.cc.utexas.edu/ Fuller and Sarkar
administered to protect biodiversity; it
*consbio/
(2006)
calculates minimum spanning trees and
Cons/Labframeset.html
performs a limited number of node and link
operations

Circuitscape Calculates and maps measures of resistance, http://www.circuit

conductance, current flows, and voltage
scape.org

Landscape metric calculations can be simplified, for example, by expanding well-known

software packages to include a broader range
of landscape metrics or introducing new software. A forthcoming update to FRAGSTATS
will include new cell-level, kernel-based
metrics as well as surface metrics for landscape
gradient data (K. McGarigal, personal communication, 2011). In contrast, the software needed
to construct and analyze landscape graphs and

McRae and Shah

(2009)

make such information readily available to

researchers and practitioners is still largely lacking. The few publicly accessible packages for
conducting landscape graph analyses collectively perform only a subset of the analyses discussed here (Table 2), and the calculation of
many landscape graph metrics requires exporting data from GIS or other software packages
for analysis in social network analysis software
(e.g. Pajek: de Nooy et al., 2005; see also http://

414

pajek.imfm.si/doku.php) or R (e.g. the igraph

library: http://igraph.sourceforge.net).
In terms of approaches, there is a general
move toward utilizing measures that include a
greater functional basis, in part because much
has been written about the shortcomings of
structural landscape metrics. However, utilizing
more functional approaches without understanding their statistical and ecological limitations and the implications for error does not
make them inherently more valid. The use of
least cost distances may be hampered, for example, by uncertainties about an organisms dispersal ability through different landscape
elements, sensitivity of analyses to the selection
of resistance values, and the basic assumption
that a disperser has complete knowledge of the
landscape and uses such knowledge to choose
dispersal routes (Jacobson and Peres-Neto,
2010; Rayfield et al., 2010; Sawyer et al., 2011).
Potential limitations and biases in graph
measures are perhaps even less well recognized (Dunn, 2010). In their study of reserve
networks in Central Finland, Laita et al.
(2011) demonstrated that: (1) conceptual differences in graph-theoretic connectivity measures could cause inconsistent connectivity
evaluations at both the patch and landscape
scales; (2) network characteristics may affect
the performance of connectivity measures;
and (3) patch prioritizations based on a node
removal analysis were sensitive to the measures used to assess changes in network connectivity and the methods used to weight
patch size and patch location. More research
on how well ecological processes are incorporated into graph structures and analyses is
clearly necessary. Particularly needed are studies
that: (1) test the behavior and responses of graph
measures to changing landscape structures at a
range of spatial scales; and (2) compare the abilities of graph measures and structural landscape
metrics to capture fundamental ecological patterns and processes (e.g. patch occupancy, dispersal) in a range of environments.

Progress in Physical Geography 36(3)

Finally, one approach to making landscape

analyses more interpretable has been an evolution away from solely indicator-based measures
to methods that incorporate visualization
techniques. Vogt et al. (2007, 2009) used Morphological Spatial Pattern Analysis (MSPA), a
pixel-level analysis of cover maps using image
segmentation, to classify and map individual pixels into core, patch, connector, perforation, and
edge categories. Goetz et al. (2009) used this classification to evaluate the location and connectivity
of core forest areas in the Northeastern and midAtlantic USA and to identify parks that play a key
role as connecting links between core areas. These
analyses can be conducted using the freely available Guidos software package (http://forest.jrc.ec.
europa.eu/download/software/guidos).
Graph-based approaches that produce maps
identifying key sites and linkages on the basis
of habitat availability metrics (Gurrutxaga
et al., 2011; Pascual-Hortal and Saura, 2006)
or that use circuit theory to depict spatial patterns of landscape resistance or conductance
(Schwartz et al., 2009; Shirk et al., 2010) provide an easily interpretable means for conveying more complicated metric values and can
be used to portray uncertainty of modeled linkages under different assumptions (Beier et
al., 2008). Saura et al. (2011b) have combined
the MSPA approach to classifying forest spatial
patterns with graph-based metrics of habitat
availability provided by the Conefor Sensinode
package to quantify properties of core habitat
areas and the structural connectors between
them for two case studies in Spain.

VI Summary: rethinking landscape

metrics in a post-FRAGSTATS
landscape
Despite concerns that landscape metrics based
on simple topological relationships often fail
to adequately capture ecologically meaningful
information about landscape structure, they

Kupfer

remain popular. They can be easily calculated

with readily available land-cover data and software such as FRAGSTATS needs little or no
parameterization, and often has intuitive interpretations. Such metrics have served to focus
efforts on the quantification of landscape structure and will continue to have a role in exploratory and descriptive landscape analysis and in
landscape monitoring for the foreseeable future.
Further, recent advances in analytical approaches
(Cardille and Lambois, 2010; Cushman et al.,
2008) and ecoregionalization techniques (Long
et al., 2010) have facilitated the calculation,
interpretation, and visualization of information
content in large sets of traditional landscape
metrics at broad spatial scales.
On the other hand, metrics rooted in graph,
network, and circuit theory, including alternative methods for quantifying patch isolation and
landscape connectivity such as least cost distances and resistance surfaces, offer the promise
of a more ecologically oriented approach to
quantifying landscape pattern and process.
Graph-based approaches may more effectively
bridge the gap between structure and function
that has often been cited as a shortcoming of
many landscape metrics, but are much less data
demanding than spatially explicit population
models. The primary limitations to the effective implementation of least cost measures and
graph-based approaches are, in many respects,
similar to those that faced structural metrics
1520 years ago when FRAGSTATS was
first introduced. Specifically there are needs
for: (1) a comprehensive software package that
allows users to easily extract one- and twodimensional node and link information from
landscape rasters, incorporates a wide selection of graph construction variants and metrics,
and performs basic graph-based analyses such
as link thresholding and node removal experiments (Galpern et al., 2011); and (2) focused
studies that provide a more thorough understanding of the caveats and justifications of
graph-based measures.

415

Acknowledgements
Tim Warner, Jeff Cardille, Paul Galpern, Josh
Leisen, Kimberly Meitzen, and two anonymous
reviewers provided constructive comments that
improved the quality of this paper. Peng Gao, Todd
Lookingbill, and O.E. Jakes provided additional
assistance and data.

Funding
This research received no specific grant from any
funding agency in the public, commercial, or notfor-profit sectors.

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