Ecological Modelling 220 (2009) 867–878
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Ecological Modelling
journal homepage: www.elsevier.com/locate/ecolmodel
Simulating population variation and movement within fragmented landscapes:
An application to the gopher tortoise (Gopherus polyphemus)
Todd BenDor
a,∗
, James Westervelt
b
, J.P. Aurambout
c
, William Meyer
b
a
Department of City and Regional Planning, University of North Carolina at Chapel-Hill, Chapel Hill, NC 27599, USA
b
U.S. Army Engineer Research and Development Center, Construction Engineering Research Laboratory (CERL), Champaign, IL 61826, USA
c
Australian Department of Primary Industries, Parkville, Victoria 3554, Australia
article info
Article history:
Received 27 February 2008
Received in revised form 17 December 2008
Accepted 5 January 2009
Keywords:
Habitat fragmentation
Spatial dynamic model
Gopher tortoise (Gopherus polyphemus)
Conservation biology
Landscape ecology
Land-use change
abstract
As the human activity footprint grows, land-use decisions play an increasing role in determining the future
of plant and animal species. Studies have shown that urban and agricultural development cannot only
harm species populations directly through habitat destruction, but also by destroying the corridors that
connect habitat patches and populations within a metapopulation. Without these pathways, populations
can encounter inbreeding depression and degeneration, which can increase death rates and lower rates
of reproduction. This article describes the development and application of the FRAGGLE model, a spatial
system dynamics model designed to calculate connectivity indices among populations. FRAGGLE can help
planners and managers identify the relative contribution of populations associated with habitat patches
to future populations in those patches, taking into account the importance of interstitial land to migration
success. The model is applied to the gopher tortoise (Gopherus polyphemus), a threatened species whose
southeastern U.S. distribution has diminished significantly within its native range due to agricultural
and urban development over the last several decades. This model is parameterized with life history
and movement traits of the gopher tortoise in order to simulate population demographics and spatial
distribution within an area in west-central Georgia that supports a significant tortoise population. The
implications of this simulation modeling effort are demonstrated using simple landscape representations
and a hypothetical on land-use management scenario. Our findings show that development resulting in
even limited habitat losses (10%) may lead to significant increases in fragmentation as measured by a loss
in the rate of dispersions (31%) among area subpopulations.
© 2009 Elsevier B.V. All rights reserved.
1. Introduction
Habitat loss and associated habitat fragmentation is a growing
problem worldwide and is considered to be one of the greatest
threats to biodiversity as well as a primary cause of the current
high rate of species extinction (Wu et al., 2003). Habitat fragmen-
tation is defined as the process through which a natural habitat
becomes divided into isolated small patches of complex geometri-
cal form, within a sea of generally inhospitable land uses (Bunnell,
1999; McComb, 1999).
Fragmentation negatively affects many species, since smaller
habitat patches support disproportionately smaller populations
that are more prone to local extinction than larger ones (Hanski,
1997; McComb, 1999; Berec, 2002). By increasing patch isola-
∗
Corresponding author. Department of City and Regional Planning, CB# 3140,
UNC-Chapel Hill, New East Building, Chapel Hill, NC 27599-3140, USA.
Tel.: +1 919 962 4760; fax: +1 919 962 5206.
E-mail address: bendor@unc.edu (T. BenDor).
tion, fragmentation effectively decreases the gene pool of local
populations, thereby favoring inbreeding depression and making
the population more prone to demographic stochasticity (Berec,
2002). It also modifies the quality of the remaining habitat by
increasing edges and reducing core habitat, which can further
increase biodiversity loss among core-dwelling species. FRAGGLE
is designed to generate connectivity indices among populations
within a metapopulation to help understand the relative mixing
of the populations.
However, the consequences of fragmentation depend on the par-
ticular species considered, and both positive and negative effects
have been reported (Kremsater and Bunnell, 1999). Fragmenta-
tion tends to negatively impact species that prefer interior habitat,
while benefiting edge-adapted species (Fahrig, 1999; With, 1999).
The extent to which fragmentation impedes movements within a
metapopulation (set of local populations connected by migrating
individuals) has also been linked to the vagility and life history
characteristics of each species (McComb, 1999). Although fragmen-
tation is strongly connected to habitat loss, in this article we focus
on the connectivity issues that result from fragmentation.
0304-3800/$ – see front matter © 2009 Elsevier B.V. All rights reserved.
doi:10.1016/j.ecolmodel.2009.01.005