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A Graph-Theory Framework For Evaluating Landscape
A Graph-Theory Framework For Evaluating Landscape
Abstract: Connectivity of habitat patches is thought to be important for movement of genes, individuals,
populations, and species over multiple temporal and spatial scales. We used graph theory to characterize
multiple aspects of landscape connectivity in a habitat network in the North Carolina Piedmont (U.S.A).. We
compared this landscape with simulated networks with known topology, resistance to disturbance, and rate
of movement. We introduced graph measures such as compartmentalization and clustering, which can be
used to identify locations on the landscape that may be especially resilient to human development or areas
that may be most suitable for conservation. Our analyses indicated that for songbirds the Piedmont habitat
network was well connected. Furthermore, the habitat network had commonalities with planar networks,
which exhibit slow movement, and scale-free networks, which are resistant to random disturbances. These
results suggest that connectivity in the habitat network was high enough to prevent the negative consequences
of isolation but not so high as to allow rapid spread of disease. Our graph-theory framework provided insight
into regional and emergent global network properties in an intuitive and visual way and allowed us to make
inferences about rates and paths of species movements and vulnerability to disturbance. This approach can
be applied easily to assessing habitat connectivity in any fragmented or patchy landscape.
Keywords: dispersal, fragmented landscapes, graph theory, habitat connectivity, habitat network, network
theory, spread of disturbance.
Un Marco de Referencia Teórico-Gráfico para Evaluar la Conectividad del Paisaje y Planificar Conservación
Resumen: Se piensa que la conectividad de los parches de hábitat es importante para el movimiento de
genes, individuos, poblaciones y especies en múltiples escalas temporales y espaciales. Utilizamos la teorı́a
de gráficos para caracterizar múltiples aspectos de la conectividad del paisaje en una red de hábitats en el
Pie de Monte en Carolina del Norte (E.U.A.). Comparamos este paisaje con redes simuladas con topologı́a,
resistencia a la perturbación y tasa de desplazamiento conocidas. Introdujimos medidas gráficas como la
compartimentación y el agrupamiento, que pueden ser utilizados para identificar localidades en el paisaje
que pueden ser especialmente resilientes al desarrollo humano o áreas que pueden ser más adecuadas para
la conservación. Nuestros análisis indicaron que la red de hábitats en el Pie de Monte estaba bien conectada
para las aves. Más aun, la red de hábitats tenı́a caracterı́sticas en común con las redes en planicies, que
exhiben desplazamiento lento y con redes sin escalas, que son resistentes a las perturbaciones aleatorias. Estos
resultados sugieren que la conectividad en la red de hábitats fue suficiente para prevenir las consecuencias
negativas del aislamiento pero no para permitir la rápida dispersión de enfermedades. Nuestro marco de
referencia teórico-gráfico proporcionó entendimiento de las propiedades regionales y globales de las redes de
manera intuitiva y visual y nos permitió inferir las tasas y direcciones de los movimientos de las especies y su
vulnerabilidad a la perturbación. Este método se puede aplicar fácilmente a la evaluación de la conectividad
del hábitat en cualquier hábitat fragmentado.
∗ Current address: University of Maryland Center for Environmental Science, Appalachian Laboratory, Frostburg, MD 21532, U.S.A., email
eminor@al.umces.edu
Paper submitted March 22, 2007; revised manuscript accepted August 22, 2007.
297
Conservation Biology, Volume 22, No. 2, 297–307
C 2008 Society for Conservation Biology
DOI: 10.1111/j.1523-1739.2007.00871.x
298 Graph Theory, Connectivity, and Conservation
Palabras Clave: conectividad de hábitat, dispersión, dispersión de la perturbación, paisajes fragmentados, red
de hábitat, teorı́a de gráfic0s, teorı́a de redes
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Table 1. Definitions, ecological relevance, and examples of graph terminology used in text.
Characteristic path A network attribute measuring the If CPL is short, all patches tend to
length (CPL) average shortest path length be easily reachable. This implies
over the network a patchy population rather than
a metapopulation or
subpopulations.
Clustering coefficient A node attribute measuring the Highly clustered nodes facilitate Nodes 1–5 (Fig. 1) are highly
average fraction of the node’s dispersal and spread of clustered, whereas nodes on
neighbors that are also disturbances. They may be the right side of the graph are
neighbors with each other more resilient to patch removal not clustered.
due to many redundant
pathways.
Compartmentalization A network attribute measuring the High compartmentalization slows
or connectivity relationship between node movement through a network
correlation degree and average node and may isolate the potentially
degree of its neighbors cascading effects of
disturbance.
Component A set of nodes that are connected Patches in the same component Figure 1 shows a single graph
to each other are mutually reachable. There is component that contains all the
no movement between nodes in the network.
different components, implying
eventual genetic divergence.
Degree A node attribute measuring the Low-degree patches may be Node 2 has a node degree of 4,
number of edges (or neighbors) vulnerable to extinction if whereas node 6 has a node
adjoining a node neighboring patches are degree of 2 (Fig. 1).
developed. High-degree patches
may be population sources or
sinks, depending on size and
quality of patch.
Diameter A network attribute measuring the Short diameter implies fast Ignoring the shortcut, one
longest shortest path joining movement through the diameter in Fig. 1 is from node
any two nodes in the network; network. This could be 3 to 11 (along the path
there may be more than one beneficial (dispersal is easy) or 3→5→6→7→12→11). There
detrimental (spread of disease) are multiple diameters in this
for the focal organism. network.
Path A sequence of consecutive edges Represents the possible routes an In Fig. 1, there are multiple paths
in a network joining any two individual may take while between nodes 3 and 1: some
nodes traveling across the landscape. alternatives include 3→2→1
and 3→4→2→1. The shortest
path is 3→1.
Random networks (Fig. 2b) consist of nodes with ran- all complex networks as random, although it has been
domly placed connections. In these networks, a plot of recognized recently that most self-forming networks are
node-degree distribution is often bell shaped, with most more complex (e.g., scale free or small world) (Albert &
nodes having approximately the same number of edges Barabasi 2002; Proulx et al. 2005).
(i.e., there are no hubs). They typically do not display A scale-free network (Fig. 2c) is characterized by pref-
clustering and may or may not be planar. In fact, the U.S. erential attachment to certain nodes, so there are a few
highway system has been described as random (Barabasi high-degree nodes (i.e., hubs), whereas the majority of
& Bonabeau 2003). In the past science theory treated nodes are low-degree nodes (Barabasi & Bonabeau 2003).
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The result is that the node degree distribution follows a 2002). Network robustness depends strongly on node-
continuously decreasing function. This is evident in the degree distribution; thus, networks with significant vari-
World Wide Web, in which a few highly connected pages ance in node connectivity are most robust to random
(e.g., google.com) have millions of connections and are removal of nodes (Albert et al. 2000). A compartmental-
responsible for holding the entire Web together. In highly ized pattern of nodes may also increase overall network
fragmented landscapes, conservation efforts may result robustness by isolating deleterious effects of disturbances
in the formation of one or more landscape hubs such as (Maslov & Sneppen 2002; Melian & Bascompte 2002). In
national or state parks connected to many smaller and addition, a highly compartmentalized network may con-
scattered habitat patches such as undeveloped lots and fer a higher resistance to fragmentation if a fraction of the
city parks. Scale-free networks are highly resistant to ran- nodes is removed (Melian & Bascompte 2002), offering
dom disturbances but vulnerable to deliberate attacks an alternative form of robustness. Compartmentalization
on the hubs (Barabasi & Bonabeau 2003). In a scale-free is not a distinguishing feature of any of the types of net-
habitat network, if a disease or invasive species were in- works described above, but it does require that node
troduced into a random habitat patch it would probably degree be somewhat heterogeneous. The scale-free net-
not spread far or quickly because most patches have few work in Fig. 2 displays a fairly high level of compartmenta-
connections. Nevertheless, if the introduction was made lization.
into a hub, the invasion would quickly spread through- From a conservation standpoint, the ideal habitat net-
out the network. Similarly, network connectivity would work might resemble a scale-free network with several
show little change if most of the smaller patches were large hubs connected to multiple smaller patches. This
removed but would quickly break apart if hubs were re- would create a landscape with heterogeneous node de-
moved. Consequently, conservation and monitoring ef- gree and resilience to patch removal. The hubs would
forts would be best spent on hub patches. be protected areas, such as parks or reserves, so that
A small-world network (Fig. 2d) is characterized by there would be no threat of removal (i.e., develop-
shortcuts that allow rapid and direct movement between ment). The hubs could be managed and/or monitored
distant nodes (Fig. 1), which results in a small diameter to prevent spread of invasive species, disease, or other
relative to the number of nodes (Watts & Strogatz 1998). disturbances. Widely scattering the hubs across space
In habitat networks, these shortcuts are likely to be the would create a compartmentalized landscape, isolat-
result of natural disturbances or human intervention. For ing disturbances while allowing dispersal across the
example, hurricane-force winds may carry a bird much landscape.
farther than it would fly on its own. Alternatively, peo- Alternatively, clustering might also be desirable in habi-
ple often intentionally or unknowingly transport organ- tat networks because highly clustered areas have many re-
isms over long distances. Small-world networks are much dundant connections and can probably lose more nodes
more vulnerable to random disturbance than scale-free without losing connectivity. Clustering is common in
networks because the shortcuts make spreading through small-world networks but not in scale-free networks.
the network relatively quick and easy. Small-world net- Clustering may also confer stability to populations (Mi-
works also tend to have a high clustering coefficient com- nor & Urban 2007)—yet another desirable attribute in a
pared with random graphs, so that a node’s neighbors are habitat network.
often connected to each other (Watts & Strogatz 1998). It is not generally known whether habitat networks
tend to display the qualities described above or even
what the topology of a typical habitat network might be.
Habitat Networks
Recently, a network of bat-roosting trees was shown to
Important features in a habitat network include connec- have scale-free topology (Rhodes et al. 2006). Neverthe-
tivity and resilience to disturbance (Peterson 2002; Op- less, a single landscape can have very different connecti-
dam et al. 2003), both of which are affected by network vity characteristics when examined from the perspective
topology. An intermediate level of connectivity is most of different organisms (Bunn et al. 2000). We exam-
desirable—too little and patches will be isolated from ined landscape connectivity from the perspective of
each other, too much and disease or other disturbance songbirds—a relatively mobile taxon. We measured the
will spread rapidly (Jules et al. 2002). We use resilience network characteristics described above and determined
to refer to the number of patches that can be removed whether our study area resembled other naturally oc-
without altering network connectivity. In the network curring networks by displaying scale-free or small-world
literature, these 2 features are sometimes referred to as topology or whether it fit the simpler null model of a pla-
network robustness (i.e., robustness to the spread of nar network. Analyzing landscapes within this framework
a deleterious mutation and to the fragmentation of the allows assessment of multiple aspects of connectivity and
network as an increasing number of nodes are deleted) subsequently can lead to more informed conservation
(Albert et al. 2000; Sole & Montoya 2001; Dunne et al. plans.
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Table 2. Graph metrics used to assess connectivity for each network. the right (skewness = 1.0; Fig. 4). By design, mean node
degree was similar in the random and the scale-free net-
Triangle Random Scale-free
networka networkb networkb works (3.6 and 3.7 respectively), but the random node-
degree distribution was close to normal (skewness =
Number of components 55 43 306 0.5), whereas the scale-free node-degree distribution had
Size of largest component 1148 1337 1054 a skewness of 2.0.
Diameter 60 14 12
Diameter/size of 0.05 0.01 0.01 The clustering coefficient was higher in the Triangle
largest component network than in either of the simulated networks by at
Characteristic path length 22.1 5.8 4.6 least one order of magnitude (Table 2). The clustering
Clustering coefficient 0.08 0.0004 0.002 coefficient was 0.08 in the Triangle network, 0.0004 in
Connectivity correlation 0.47 0.00 0.48 the random network, and 0.002 in the scale-free network.
a Forest-habitat network in the Triangle region of North Carolina Finally, connectivity correlation (i.e., compartmentaliza-
(U.S.A.). tion) values were positive for both the Triangle network
b Simulated networks.
and the scale-free network and zero for the random net-
work (Table 2, Fig. 5).
Figure 4. Node-degree
distribution for the Triangle
network (a forest habitat
network) and the random and
scale-free networks (simulated
networks). The higher the node
degree, the more neighbors
connected to each node. An
asterisk (∗) indicates maximum
node degree for each network.
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topology, but it was not clear whether it resembled a park is a large (2201 ha) protected area and appears to
planar or scale-free network. be more compartmentalized than clustered, potentially
The connectivity correlation in the Triangle network conferring some protection from disturbances in the out-
was positive, indicating the inverse of compartmenta- side landscape. Finally, Jordan Lake State Recreation Area
lization—hubs tend to be clustered next to each other on is a large (1585 ha) protected area contained mostly in
the landscape. High-degree nodes (larger circles) tended Chatham County that is composed of several smaller for-
to be grouped together and separated from groups of est patches, two of which appear to be large hubs. The
low-degree nodes (smaller circles) (Fig. 6). These groups southern hub is much more clustered than the north-
also tended to be in highly clustered areas (darker colors), ern one, and this variation may provide an interesting
resulting in many darkly colored large circles and pale- opportunity to test some of the concepts discussed in
colored small circles. This occurred because there were this paper, such as the effect of clustering on population
large regions on the landscape with high forest cover stability or spread of disease.
and many patches (e.g., Chatham county—the south- For forest birds, the Triangle network appears to be
western portion of the map) and other regions with low somewhere between a planar network and a scale-free
forest cover and few patches (e.g., Wake and Johnston network, although network topology might change com-
counties—the southeastern portion of the map). Conser- pletely if a different organism were under consideration.
vation strategy in the Triangle might focus on these highly For example, if the focal organism was Norway Maple
clustered regions. On one hand, they may be desirable ar- (Acer platanoides), an exotic tree commonly used in res-
eas for many forest interior species because of their high idential landscaping, the average edge might be much
forest cover and densely connected patches, which facil- shorter than for birds because seed dispersal is more
itate dispersal. On the other hand, however, their highly limited. Nevertheless, the network would also contain
interconnected nature makes them vulnerable to spread many shortcuts radiating from plant nurseries to locations
of disease. Furthermore, some might argue that because across the Triangle. This network would more closely
clustering confers resilience to patch removal, these re- resemble a small-world network and would display the
gions can withstand development more than other re- small diameter and fast spread of disturbance character-
gions in the Triangle and development would be bet- istic of this topology.
ter concentrated in these areas than in more vulnerable,
less-clustered areas, where connectivity may be easily
Applications and Future Work with Graph Theory
severed.
Figure 6 highlights some areas of conservation inter- Graph theory may be well suited for selecting habitat re-
est in the Triangle network. Research Triangle Park, a serves (Opdam et al. 2006; Pascual-Hortal & Saura 2006;
research park in southeastern Durham County, is a heav- Minor & Urban 2007). Our results, however, do not sug-
ily forested area that is undergoing rapid development. gest a formula for reserve design because each landscape
It currently contains many hubs that are also highly clus- and conservation problem is different. Despite these dif-
tered, perhaps making this a safe choice for additional ferences, the importance of considering landscape con-
well-planned development. Umstead State Park is located nectivity when designing reserves is becoming widely
in Wake County adjacent to the international airport. The acknowledged (e.g., Briers 2002; Nikolakaki 2004;
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306 Graph Theory, Connectivity, and Conservation
Moilanen & Wintle 2007). Here, we presented some new Bunn, A. G., D. L. Urban, and T. H. Keitt. 2000. Landscape connectivity: a
tools for measuring aspects of connectivity that might conservation application of graph theory. Journal of Environmental
Management 59:265–278.
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Calabrese, J. M., and W. F. Fagan. 2004. A comparison-shopper’s guide
times a high level of connectivity is desirable (Jordan et to connectivity metrics. Frontiers in Ecology and The Environment
al. 2003), whereas at others it can be detrimental (Con- 2:529–536.
deso & Meentemeyer 2007). Difficult choices must some- Clergeau, P., and F. Burel. 1997. The role of spatio-temporal patch
times be made, such as whether it is more important to connectivity at the landscape level: an example in a bird distribution.
have a network that is robust to spread of disturbance or Landscape and Urban Planning 38:37.
Colizza, V., A. Flammini, A. Maritan, and A. Vespignani. 2005. Char-
one that maximizes population stability. If the former is acterization and modeling of protein-protein interaction networks.
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reserve might be the better design. Graph theory cannot heterogeneity on the emerging forest disease sudden oak death.
Journal of Ecology 95:364–375.
make these decisions, but it does provide tools that make
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these decisions easier. ture and biodiversity loss in food webs: robustness increases with
The next step in developing graph-based conservation connectance. Ecological Letters 5: 558–567.
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Acknowledgments Jules, E. S., M. J. Kauffman, W. D. Ritts, and A. L. Carroll. 2002. Spread
of an invasive pathogen over a variable landscape: a nonnative root
We thank the members of the Duke University Nicholas rot on Port Orford cedar. Ecology 83:3167–3181.
Keitt, T. H., D. L. Urban, and B. T. Milne. 1997. Detecting critical scales
School graph group, especially E. Treml, for discussion
in fragmented landscapes. Conservation Ecology 1: http://www.
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Moilanen, and the anonymous reviewers for their com- King, A. W., and K. A. With. 2002. Dispersal success on spatially struc-
ments on this manuscript. tured landscapes: when do spatial pattern and dispersal behavior
really matter? Ecological Modelling 147:23–39.
Kossinets, G., and D. J. Watts. 2006. Empirical analysis of an evolving
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