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Estimating landscape resistance from habitat suitability: effects of data

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Landscape Ecol
DOI 10.1007/s10980-016-0387-5
RESEARCH ARTICLE
Estimating landscape resistance from habitat suitability:
effects of data source and nonlinearities
Annika T. H. Keeley . Paul Beier .
Jeffrey W. Gagnon
Received: 31 August 2015 / Accepted: 18 April 2016
Ó Springer Science+Business Media Dordrecht 2016
Abstract
Context Conservation corridors must facilitate long-
distance dispersal movements to promote gene flow,
prevent inbreeding, and allow animals to shift ranges
with climate change. Least-cost models are used to
identify areas that support long-distance movement.
These models rely on estimates of landscape resistance,
which are typically derived from habitat suitability.
Objectives We examine two key steps in estimating
resistance from habitat suitability: choosing a procedure to
estimate habitat suitability, and choosing a transformation
function to translate habitat suitability into resistance.
Methods We used linear and nonlinear functions to
convert three types of habitat suitability estimates
(from expert opinion, resource selection functions, and
step selection functions) into resistances for elk
(Cervus canadensis) and desert bighorn sheep (Ovis
canadensis nelsoni). We evaluated the resulting
Electronic supplementary material The online version of
this article (doi:10.1007/s10980-016-0387-5) contains supple-
mentary material, which is available to authorized users.
A. T. H. Keeley (&)  P. Beier
School of Forestry, Northern Arizona University, 200 E
Pine Knoll Drive, Flagstaff, AZ 86011, USA
e-mail: annika.keeley@yahoo.com
J. W. Gagnon
Arizona Game and Fish Department, 5000 W Carefree
Highway, Phoenix, AZ 85023, USA
resistance maps on an independent set of observed
long-distance, prospecting movements.
Results A negative exponential function best
described the relationship between resistance values
and habitat suitability for desert bighorn sheep indi-
cating long-distance movers readily travel through
moderately-suitable areas and avoid only the least
suitable habitat. For desert bighorn sheep, all three
suitability estimates performed better than chance, and
resource and step selection functions outperformed
expert opinion. For elk, all three suitability estimates
performed the same as chance.
Conclusions When designing corridors to facilitate
long-distance movements of mobile animals, we
recommend transforming habitat suitability into resis-
tance with a negative exponential function. Use of an
exponential transformation means that larger fractions
of the landscape offer low resistance, allowing greater
flexibility in where a corridor is located.
Keywords Cervus canadensis  Desert bighorn
sheep  Elk  Expert opinion  Exploratory movements 
Least-cost models  Ovis canadensis  Resource
selection function  Step selection function  Wildlife
corridors
Introduction
Wildlife corridors are intended to allow animals to
move through the landscape, maintain genetic diver-
sity, and shift ranges with climate change (Hilty et al.
123

2012). Corridors must meet all life-history needs for
relatively sedentary species (corridor dwellers—Beier
et al. 2008a). But for relatively mobile species,
including most large mammals, corridors are intended
to support migration and dispersal movements. Dis-
persal is defined as movement of an animal from the
area where it is born (or, rarely, from a breeding home
range) to another area where it joins, or attempts to
join, the local breeding population (Baguette et al.
2013). Dispersal movements, although rare, are the
most important movements underlying gene flow,
demographic viability of metapopulations, recolo-
nization, and range shifts (Trakhtenbrot et al. 2005).
Prospecting movements, during which animals
become familiar with the habitat surrounding the
home range, frequently precede dispersal movements
(Stamps and Krishnan 1999; Stamps 2001). Here, we
assume that habitat use during long-distance,
prospecting movements is similar to habitat use during
long-distance, dispersal movements.
Least-cost models (Adriaensen et al. 2003) are
commonly applied to identify wildlife corridors in a
conservation context (Beier et al. 2008a, 2011; Sawyer
et al. 2011). These models use estimates of landscape
resistance (the degree to which a grid cell facilitates or
impedes movement of the study organism; Spear et al.
2010) to identify the path with the least accumulated
resistance between two locations. Thus resistance
estimates are critical to least cost corridor models
(Beier et al. 2008a; Sawyer et al. 2011; Zeller et al.
2012). If estimates are wrong, then land-use planners,
transportation agencies, and other decision-makers
may be implementing poor corridor designs. Resis-
tance estimates are commonly derived from habitat
suitability values because habitat suitability is easier to
study than landscape use during rare prospecting and
dispersal movements. Here, we examine two key steps
in estimating resistance from habitat suitability:
choosing a procedure to estimate habitat suitability,
and choosing a transformation function to transform
habitat suitability into resistance.
Habitat suitability is typically modeled as a func-
tion of land cover, elevation, and other attributes. To
estimate habitat suitability, expert opinion models are
popular because they are less expensive and do not
require intensive field data collection (Walker and
Craighead 1997; Hoctor et al. 2000; Singleton et al.
2002; Hunter et al. 2003; Larkin et al. 2004; Wikra-
manayake et al. 2004; Beier and Majka 2006;
123
Landscape Ecol
Adriaensen et al. 2007; Beier et al. 2008a). Alterna-
tively, landscape variables at sites used by an animal
are compared to equivalent sites that are either unused
or potentially available (Manly et al. 2002) to estimate
habitat suitability. When the used sites are animal
point locations, resource selection functions (RSF) are
employed to model habitat suitability (e.g., Johnson
et al. 2000). However, when used sites are movement
steps (straight lines connecting consecutive locations
separated by no more than a few hours), step selection
functions (SSF) are used to model habitat suitability
(e.g., Fortin et al. 2005). We hypothesize that all three
procedures yield habitat suitability estimates that can
inform useful least-cost models. We predict that
resistance derived from expert opinion is least reliable
because expert opinion is subjective, and the resis-
tance derived from SSF is most reliable because SSF
are based on the data set that contains the most
information about the animals’ habitat use.
When resistance is estimated from habitat suitabil-
ity, it is usually assumed to be a negative linear
function of suitability (Walker and Craighead 1997;
Hoctor et al. 2000; Ferreras 2001; Singleton et al.
2002; Hunter et al. 2003; Larkin et al. 2004; Wikra-
manayake et al. 2004; Richards-Zawacki 2009; Pull-
inger and Johnson 2010) such that resistance decreases
at a constant rate as suitability increases, as in the
upper line in Fig. 1. Beier et al. (2008a) called
attention to the lack of empirical support for this
crucial assumption, labelled it the ‘‘subjective trans-
lation problem,’’ and argued that it was a crucial
uncertainty in all corridor modeling. It seems plausible
that during long-distance dispersal or pre-dispersal
prospecting movements, animals might readily move
through moderately suitable areas such that resistance
increases only modestly as suitability decreases from
its maximum value, and then increases dramatically at
lower suitability values (lower curves in Fig. 1). Such
a relationship was reported by Trainor et al. (2013) for
red-cockaded woodpeckers (Picoides borealis) and by
Mateo-Sánchez et al. (2015) for brown bears (Ursus
arctos). We hypothesized that this negative exponen-
tial relationship applies to most animals.
In this study, we test whether estimates of habitat
suitability can predict habitat use during long-dis-
tance, prospecting movements. We compare resis-
tance estimates from linear and nonlinear
transformations of three types of suitability estimates
(from expert opinion, RSF, and SSF) in terms of their
Landscape Ecol
Fig. 1 Eight curves used to transform habitat suitability values
into resistance values. The curves are based on the transforma-
tion function R ¼ 100  99  ðð1  expðc  HÞÞ =
ð1  expðcÞÞÞ, where R is resistance, H is suitability, and
the factor c determines the shape of the curves
consistency with observed long-distance prospecting
movements for two species of large mobile mammals.
Our method of evaluating each resistance model (a
combination of a particular suitability estimate and
transformation function) assumes that during
prospecting animals choose paths of low cumulative
resistance (Adriaensen et al. 2003). If this assumption
is true, long-distance paths will have a relatively low
cost compared to random paths of identical topology
in the same geographic area.
Methods
Study area
We studied elk on and near the Tonto, Coconino and
South Kaibab National Forests in northern Arizona.
Elevations range from 1100 to 3800 m, and vegetation
types include pinyon (Pinus edulis)–juniper (Junipe-
rus spp.) woodlands and chaparral (Arctostaphylos
spp.) at lower elevations, ponderosa pine (P. pon-
derosa) forest at medium elevations and mixed conifer
forest at upper elevations.
We studied desert bighorn sheep in the Black
Mountains in northwestern Arizona on land managed
by the Bureau of Land Management. Elevations range
from 150 m at the Colorado River to 1663 m. Mojave
Desert scrub and creosote (Larrea tridentata) shrublands
are dominant vegetation types. Topography includes
rugged mountainous terrain with steep talus slopes and
cliffs as well as dry washes among rolling hills.
Habitat data
To estimate habitat suitability, we first identified land
cover and geographic variables that are known to affect the
distribution of elk (Wallace and Krausman 1997; Boyce
et al. 2003; Wait and McNally 2004; Creel et al. 2005;
Mao et al. 2005; Coe et al. 2011; Beck et al. 2013) and
desert bighorn sheep (Turner et al. 2004; Sappington et al.
2007; Hoglander 2012). We reclassified vegetation types
from the Landfire Program (Landfire 2008) into broad
land-cover variables (Tables S1, S2) to limit the number of
variables. We obtained elevation and slope from 30-m
resolution digital elevation models (DEM, http://
nationalmap.gov/elevation.html, last accessed August
21, 2015). We applied the DEM Surface Tools extension
(Jenness 2013) to generate aspect and slope rasters in
ArcGIS 10.2 (ESRI 2011). We transformed aspect, a cir-
cular variable, into two independent variables, northness
(cos(aspect)) and eastness (sin(aspect)). We obtained road
and highway maps from the topologically integrated
geographic encoding and referencing (TIGER) project of
the U.S. Census Bureau (http://www.census.gov/geo/
maps-data/data/tiger.html, last accessed August 21,
2015). We calculated distances to roads or highways in
ArcGIS 10.2 (ESRI 2011). All variables were resampled
to the North American geographic coordinate system from
1983 with a cell size of 30 9 30 m.
Following the commonly used threshold of r = 0.7
(Booth et al. 1994; Dormann et al. 2013), correlations
among habitat covariates for elk were relatively low
(maximum r = -0.57, Table S3), allowing us to
retain all predictor variables in our models. For
bighorn sheep, we excluded the topographic position
index because it was strongly correlated with slope
(r = 0.72, Table S4).
Animal locations
The Arizona Game and Fish Department (AZGFD)
radio collared elk from 2004 through 2012 with global
positioning system (GPS) collars (models TGW-3600,
TGW-3690, TGW 4600; Telonics, Inc., Mesa, AZ;
and NSG-LD2 Northstar Science and Technology,
LLC, King George Virginia; Dodd et al. 2012a, b;
Gagnon et al. 2012, 2013). Each collar recorded the
animal’s location every 2 h between 17:00 and 7:00 or
123

between 16:00 and 10:00, or, for a few individuals,
every 1.5 or 4 h throughout the day. GPS error was
±10 m (Dodd et al. 2006). To minimize heterogeneity
within models, we restricted our analyses to data of
yearling bull elk in April, May, and June, because
yearling bulls made more prospecting movements
than any other age-sex class, and because elk disperse
at about this age (Petersburg et al. 2000). To avoid
locations with large errors, we removed two-dimen-
sional fixes with a PDOP [5 (Lewis et al. 2007).
AZGFD equipped desert bighorn sheep in the Black
Mountains from 2004 through 2012 with remote-
download, self-dropping, store-on-board GPS collars
(model TTW-3590; Telonics, Inc., Mesa, AZ; McKin-
ney and Smith 2007; Bristow and Crabb 2008) and
satellite uplink collars (NSG-D2 Northstar Science
and Technology, LLC, King George Virginia; Gagnon
et al. 2011). Most collars recorded the animal’s
location every 5 h for 6–24 months, some collars
recorded the location every 2 h (Gagnon et al. 2011).
Because precipitation amounts can affect desert big-
horn sheep space and resource use (Hoglander 2012),
we selected bighorn sheep location data from July to
March, but omitted data from April, May and June
when mean precipitation over the last 30 years was
below 1 cm per month (http://www.usclimatedata.
com/climate/henderson/nevada/united-states/usnv0122).
We removed two-dimensional fixes with a PDOP [ 5.
For the RSF, we reduced the GPS collar data sets
to resemble VHF telemetry study data sets by
randomly selecting one location between 17:00 and
19:00 h per day for each elk, and one location
between 9:00 and 11:00 h per day for each bighorn
sheep. The time windows reflected times of peak
activity for elk (Green and Bear 1990) and bighorn
sheep (Alderman et al. 1989). Short time intervals
would likely have greater dependence between
successive locations. We only used the data for
animals that maintained coherent home ranges
throughout the 12- to 18-month life of their collars
(i.e., ‘‘stay-at-home’’ individuals).
For the SSF, we limited the data sets to include only
consecutive GPS fixes with the minimum GPS fix
acquisition intervals (elk: 2 h; bighorn sheep: 5 h). We
defined a movement step as the straight-line path
between two consecutive GPS fixes. To focus on
movement behavior, we excluded steps \200 m for
elk and\100 m for bighorn sheep. We only used data
of stay-at-home individuals.
123
Landscape Ecol
For testing resistance estimates from expert opin-
ion, RSF, and SSF in terms of their consistency with
observed long-distance prospecting movements, we
visually identified long-distance movements by indi-
vidual elk and bighorn sheep outside their established
home range (Fig. 2). The beginning and ending points
were located at the edge of the home ranges; all the
other points of the prospecting movements were
located outside the home ranges. To avoid circularity,
we used data for stay-at-home individuals to estimate
resistance and data for animals that undertook these
prospecting movements to evaluate the models.
Estimating habitat suitability
To build habitat suitability models based on expert
opinion, we used previously published expert opinions
of resistance for elk (Table S5; Beier et al. 2008b) and
bighorn sheep (Table S6; Beier and Majka 2006). To
generate these opinions, biologists familiar with the
region reviewed the literature on habitat selection by
the focal species. They then assigned a score between
1 and 10 (1 being most suitable) to vegetation classes,
topographic positions, elevation classes, and distance-
to-road classes, and weighted these four factors
according to their estimated importance.
To estimate habitat suitability with a resource
selection function (RSF), we conducted a two-step
conditional logistic regression to quantify selection
for each habitat attribute (Compton et al. 2002;
Manly et al. 2002; Zeller et al. 2014). First, we
determined the optimum spatial scale of each habitat
covariate to consider as available. Second, we created
a multi-scale model by using the best spatial scale for
each habitat covariate. In a conditional logistic
regression, used habitat is compared to available
habitat. We characterized used habitat by values of
habitat variables in grid cells with GPS fixes. To
characterize available habitat, we first fit a general-
ized Pareto distribution to the straight-line distances
between GPS points separated by a fixed time interval
(Zeller et al. 2014) using the POT package (Ribatet
2012). At each location, we then placed a Pareto
kernel and truncated this kernel at the smaller of the
97.5 percentile or the maximum observed movement
distance. We calculated the mean value of each
habitat variable in the kernel, inversely weighted by
the distance to the observed location (Zeller et al.
2014). We calculated the difference between used
Landscape Ecol
Fig. 2 Example of a long-
distance path (diamonds) of
a yearling male elk in
relation to the home range
(circles)
and available for each variable, and set the response
variable equal to one for each GPS location.
To determine the optimum spatial scale of each
habitat covariate to consider as available in the first
step, we calculated four Pareto kernels for elk,
corresponding to 2-, 4-, 8-, and 16-h intervals between
consecutive fixes, and 3 Pareto kernels for bighorn
sheep, corresponding to 5-, 10-, and 20-h acquisition
intervals. We then used a conditional mixed-effects
logistic regression (lmer function in the lme4 package,
v.0.999999-2, Bates et al. 2014), with individual
animals as a random effect, and identified the scale
with the lowest AICc ranking (AIC corrected for small
sample size; Burnham and Anderson 2002) as the best
scale at which to assess availability.
Our top models were generally clearly better than
alternative models. Therefore, to create the multi-scale
model we used the best scale for each habitat
covariate, as identified in step one. Because many of
our paired mixed-effects models using individual
animals as the random effect failed to converge, we
used generalized linear models with a binomial error
distribution (glm function, R Core Team 2014). We
conducted an automated model selection using the
glmulti package in R (Calcagno and de Mazancourt
2010). This package uses a genetic algorithm to find
the best models for large candidate sets. We selected
the best model based on the AICc value.
To estimate habitat suitability with a step selection
function (SSF), we characterized used habitat as the
proportion of each land-cover type, and the mean of the
continuous variables, within a 30-m buffer on each side
of the segment to account for GPS error (Rettie and
McLoughlin 1999) and to include the immediate
environment around each step. We characterized avail-
able habitat as these same proportions and means within
a Pareto kernel centered on the starting point of each
step. We generated the Pareto kernel the same way as in
the RSF calculations, but only for one scale (Zeller et al.
2014, 2015). We used the differences between used and
available habitat values as predictor variables. In this
specification, the response variable is 1 and there is no
model intercept (Agresti 2002; Zeller et al. 2015). We
created a full model and conducted an automated model
selection with the glm function in the glmulti package
(Calcagno and de Mazancourt 2010). We selected the
best model based on the AICc value.
Generating resistance surfaces
We used the expert habitat suitability estimates and
the coefficients of the best RSF and SSF models to
create habitat suitability surfaces. We parameterized
resistances per cell, rx, across the landscape as:
r x ¼  1  z 1 þ 2  z 2 þ    þ i  z i ; ð1Þ
123

where ß1-i are the habitat suitability beta coefficients,
and z1-i are the environmental covariates.
We rescaled the suitability values to a range of 0–1
with Eq. 2.
f ðxÞ ¼ ðx  minÞ=ðmax  minÞ; ð2Þ
where x is the habitat suitability value of a grid cell,
and min and max are the minimum and maximum
habitat suitability values of the habitat suitability
surface, respectively. Values near one indicate the
most suitable habitat conditions and values near zero
indicate the least suitable habitat.
Transforming habitat suitability into resistance
To transform suitability values into resistance values,
we used eight curves based on the transformation
function (Eq. 3) to define the relationship between
suitability and resistance (Trainor et al. 2013):
100  99  ðð1  expðc  hÞÞ = ð1  expðcÞÞÞ;
ð3Þ
where h is the suitability matrix, and c = 0.25, 0.5, 1,
2, 4, 8, 16 or 32.
For each of the eight transformations, resis-
tance = 1 when suitability = 1 and resistance =100
when suitability = 0. At c = 0.25, the relationship is
nearly linear; as c increases, resistance values become
an increasingly nonlinear negative exponential func-
tion of suitability (Fig. 1). For each group of animals
(yearling bull elk, bighorn sheep ewes, bighorn sheep
rams), we generated 24 resistance surfaces: eight
transformations each of expert habitat suitability
estimates (Tables S5, S6), RSF suitability values,
and SSF habitat suitability values.
Evaluating resistance surfaces
To evaluate whether the observed long-distance
movement paths were low-cost paths as predicted by
one or more resistance models, we generated 49
‘‘random’’ paths per observed path by rotating the
observed path around the center point (see Cushman
et al. 2010, 2011) using the R-package adehabitatLT
(Calenge 2006; Fig. S1). For each resistance model,
we calculated the cost of each of the 50 paths (1
observed, 49 random) by summing the least cost
distances between consecutive GPS fixes of paths
123
Landscape Ecol
using the R-package gdistance (van Etten 2014). We
then arrayed the 50 paths by cost, and calculated the
cost rank of the observed path. The mean cost rank
(MCR) of several observed paths is a measure of
aptness of the underlying resistance model. An MCR
of 1 is best, MCR values near the median (25.5)
indicate a model that performs about the same as
chance, and MCR of 50 is worst. Therefore, MCR
evaluates how well the model predicts long-distance
paths. If actual paths consistently have low costs under
a model, then the model can be expected to identify
corridors that would support exploratory and dispersal
movements by that species.
We tested for differences between the eight trans-
formations of each suitability model (expert opinion,
RSF, SSF) with linear mixed effects models (R-
packages lme4: Bates et al. 2014; and lmerTest:
Kuznetsova et al. 2015):
cost difference  suitability model  transformation
þ ð1jIDÞ þ ð1jID : rotated pathÞ;
ð4Þ
where cost_difference is the difference in cost
between the observed path and the rotated paths and
ID refers to an observed path and its rotated versions.
The last two terms specify random effects attributed to
the effects of the ID, and to the interaction between the
ID and the individuals rotated paths, respectively.
P-values based on Satterthwaite’s approximations
indicate significantly different models and transfor-
mations. We also tested whether a resistance model
was significantly better than random (rank of 25.5)
with 1-tailed Student’s t-tests.
Results
Animal locations
We analyzed 35,836 fixes across 33 stay-at-home
yearling bulls (range = 68–3092, median = 987);
44,390 GPS fixes of 29 stay-at-home ewes (range =
359–2418; median = 1711), and 22,308 fixes of 17
stay-at-home rams (range = 627–2266, median =
1481). We identified nine long-distance movements
made by six yearling bull elk, 11 long-distance
movements by seven ewes, and 15 long-distance
movements by 13 rams.
Landscape Ecol
Habitat suitability estimates
The Pareto kernel distances for elk in the RSF varied
from 1430 to 4705 m for elk and from 1000 to 2529 m
for bighorn sheep (Table 1). The characteristic scale
varied among habitat variables. For yearling bull elk,
the strongest response occurred at the finest scale (2-h
and 1430-m) for elevation, slope, northness, distance
to highways, and shrub; but at a coarse scale (16-h and
4705-m) for eastness, distance to road, open, riparian
and developed areas of medium to high intensity
(Table S7). Bighorn sheep ewes and rams responded
most strongly to most habitat variables at the 20-h and
[2400-m scale (Tables S8, S9). Rams selected
elevation and sparse vegetation at the 5-h and
1366-m scale and grassland at the 10-h and 1743-m
scale. Ewes also responded to elevation and sparse
vegetation, but also to solar insolation and open
developed areas at the finest scale.
In both the best multivariate RSF and SSF
models, yearling bull elk strongly avoided devel-
oped areas of medium–high intensity, but strongly
preferred developed open areas such as golf courses
(Tables S10, S11a) and north-facing areas. Although
elk avoided roads and highways in the RSF model,
elk were indifferent to small paved roads and tended
to occur close to highways in the step selection
model. The expert opinion model differed most from
the RSF and SSF models in suitability of open
developed areas, which were moderately avoided in
the expert opinion model but strongly favored in
RSF and SSF models.
As expected, bighorn sheep preferred medium
levels of elevation and ruggedness and preferred to
be near permanent water (Table S11b, c). Ewes
Table 1 GPS fix Species and sex
acquisition intervals and the
associated Pareto kernel Elk yearling bulls
distances
Bighorn sheep rams
Bighorn sheep ewes
preferred desert scrub and riparian areas in the RSF
model, but avoided these landcover types in the SSF
model. Rams showed a slight preference for developed
areas in the RSF model, but strongly avoided them in
the SSF model. The expert model matches the RSF
and SSF models closely: all three models show that
bighorn sheep prefer medium elevations and steep
slope but avoid roads.
Transforming habitat suitability into resistance
The relationship between resistance and suitability
was better defined by negative exponential functions
for the resource and step selection models of bighorn
sheep rams and ewes (Fig. 3; Tables S12, S13). For the
expert models of bighorn sheep and for all elk models,
there were no significant differences among transfor-
mation functions, but there was a trend towards better
performance of moderately exponential functions
(Tables S12, S13; Fig. 3).
Evaluating resistance surfaces
At the transformations that yielded the lowest MCR,
none of the elk resistance models performed better
than chance (1-tailed t-tests: p [ 0.07; Fig. 3). For
bighorn sheep ewes and rams, all models performed
better than chance (1-tailed t-tests: p \ 0.015), includ-
ing models in which suitability was linearly trans-
formed to estimate resistance (Fig. 3).
For bighorn sheep ewes and rams, the resource and
step selection models performed significantly better
than the expert model, and there was no difference in
performance between the resource and step selection
models (Fig. 3; Table S13). For elk, we did not detect
GPS acquisition interval Pareto kernel distance (m)
2-h steps 1430
4-h steps 2173
8-h steps 2891
16-h steps 4705
5-h steps 1366
10-h steps 1743
20-h steps 2529
5-h steps 1003
10-h steps 1679
20-h steps 2396
123

(a)
(b)
(c)
Fig. 3 Effect of the function used to transform habitat
suitability (see Fig. 1) into resistance estimates on the mean
(±1 SE) cost rank of observed paths in comparison to 49 random
paths. Lower mean ranks indicate a better transformation or
model (expert opinion, resource selection functions, step
selection functions). A rank of 25.5 (dashed line) indicates
performance of a random resistance model. a elk, b bighorn
sheep ewes, c bighorn sheep rams
123
Landscape Ecol
a difference in performance among the three models
(Fig. 3; Table S13).
Discussion
Our results indicate habitat suitability estimates can
yield resistance estimates that reliably identify long-
distance movement paths for desert bighorn sheep, a
habitat specialist, but that do not identify long-
distance paths of elk, a habitat generalist. For bighorn
sheep ewes and rams, the expert opinion model did not
perform as well as the data-intensive RSF and SSF
models, but still performed better than chance. Con-
sidering the cost of obtaining data for RSF or SSF
models, the combination of an expert opinion model
generated for the study area and a negative exponential
transformation of habitat suitability into resistance
may be, as in the case of the bighorn sheep, a reliable
foundation for corridor design. However, for little
studied species it may be safer to spend the time and
money needed to estimate habitat suitability to ensure
a sound underlying model for generating resistance
values.
Contrary to our hypothesis, resistance derived from
RSF (i.e., from telemetry locations obtained once per
day during a period of peak activity) was as reliable as
resistance derived from SSF (movement steps). We
observed a few differences between habitat suitability
models that might be due to incorporating different
spatial scales in the RSF models but only working with
the 2-h scale in the SSF models. We are not aware of
any studies comparing these two approaches, but if
this pattern holds for other species, resistance can be
estimated using data from relatively inexpensive
radio-transmitters.
For desert bighorn sheep, the resource and step
selection models had the lowest MCR when the
relationship between resistance values and habitat
suitability followed a strongly negative exponential
curve. This indicates that bighorn sheep, when under-
taking long-distance movements, move through habi-
tat of moderate and moderately low suitability as
readily as through highly suitable habitat. Only the
least suitable habitat poses high resistance to long-
distance movements. Similarly, landscape resistance
to exploratory movements of red-cockaded
Landscape Ecol
woodpeckers was a strongly negative exponential
function of habitat suitability (Trainor et al. 2013),
and landscape genetic patterns of brown bears and
kinkajous (Potos flavus) were consistent with a negative
exponential relationship between resistance and habitat
suitability (Mateo-Sánchez et al. 2015; Keeley 2015).
Similarly, habitat conditions that were avoided within
the home range were readily traversed by dispersing
individuals in many species, including the butterflies
Junonia coenia and Euptoieta claudia (Haddad and
Tewksbury 2005), hispid cotton rats (Sigmodon
hispidus, Bowne et al. 1999), Siberian flying squirrels
(Pteromys volans, Selonen and Hanski 2006), Iberian
lynx (Lynx pardinus, Gastón et al. 2016), and lions
(Panthera leo, Elliot et al. 2014). Similar results are
evident in landscape genetic studies that—instead of
transforming combined habitat suitability values into
resistance with negative exponential functions—first
conducted univariate scaling to determine the best scale
and functional form of each landscape variables. In
these studies, initial parameter values were selected
based on literature review and expert opinion. In the
optimized multivariate models that best explained how
landscape structure influences gene flow, only 1–2 of
the initial 4–5 landscape variables were retained,
indicating that mountain goats (Oreamnos americanus,
Shirk et al. 2010), American marten (Martes ameri-
cana, Wasserman et al. 2010), and American pikas
(Ochotona princeps, Castillo et al. 2014) are con-
strained by fewer landscape variables during dispersal
and mating movements than during daily movements.
Given this evidence, we propose that when
researchers and managers design corridors to facilitate
dispersal, migration, and other long-distance move-
ments of mobile animals, a negative exponential
relationship should be the default assumption when
transforming habitat suitability into resistance. Until
now, the negative linear relationship has been the
default, despite the lack of any empirical studies
supporting a linear transformation (Beier et al. 2008a).
Use of an exponential transformation has important
implications for corridor design because it means that
larger fractions of the landscape offer low resistance,
allowing greater flexibility in where a corridor is
located. We caution, however, that all species studied
were relatively mobile animals that can pass through a
long corridor designed to connect adjacent popula-
tions in a single movement lasting a few hours or days.
In contrast, corridor dwellers require generations to
move their genes through these long corridors. For
these species, a long corridor can be conceptualized as
stepping stones of suitable habitat, separated by gaps
crossed in discrete dispersal events (Beier et al.
2008a). Research is needed on the minimum size of
stepping stones, and on how resistance and habitat use
are related for these less-mobile species.
For elk, none of the three models performed better
than chance. We think the main reason for this
outcome is that elk are generalist herbivores (Van
Deelen et al. 1997), whereas bighorn sheep are habitat
specialists (Turner et al. 2004). The habitat niche of
elk was strikingly broad during the long distance
movements that we observed. Similarly, McClure-
Rainey (2012) found that least-cost paths informed by
habitat suitability models using observed migration
paths of elk predicted an independent set of migration
paths no better than null models. In contrast, Poor et al.
(2012) examined how well least-cost models matched
migratory movements of pronghorn (Antilocapra
americana), a habitat specialist (Wood 1989). They
created least-cost paths based on Maxent habitat
suitability models and buffered them by 10 % of the
most traversable habitat. Because 71 % of migrating
pronghorn GPS locations occurred in these least-cost
corridors, Poor et al. (2012) concluded the least-cost
method is a valid approach for conservation planning.
Collectively, these results support the conclusion of
Beier et al. (2009) that habitat specialists are the most
important focal species for design of multi-species
corridors because habitat generalists can easily move
through landscapes conserved for habitat specialists,
but habitat specialists may not readily move through a
corridor designed for a generalist.
This paper introduces MCR as a tool to evaluate
aptness of resistance models. MCR can determine if a
model is statistically better than random with only a
small number of observed paths (as few as nine paths
in our analyses). MCR can also be used to statistically
test for differences between alternative resistance
models. Finally, this tool considers habitat use along
the entire path, not only at the actual telemetry fix
locations, and thus takes advantage of the rich
information yielded by GPS radio-tags.
Some studies that compared resistance estimates
from expert opinion and empirical methods found
expert opinion is not reliable (e.g., Seoane et al. 2005;
Shirk et al. 2010; Charney 2012). However, our study
shows expert opinion can predict long-distance
123

movements of desert bighorn sheep. Because many
corridor designs are based on resistances derived from
expert opinion (e.g., Arizona Missing Linkages: http://
corridordesign.org/linkages/arizona, South Coast
Missing Linkages in California: Beier et al. 2006), it is
useful to know that expert opinion paired with litera-
ture review of empirical studies can be a sound source
for resistance estimates for some focal species. We
expect expert opinion models of well-studied species
to be reliable, but recommend habitat suitability
modeling for little studied species to ensure a good
underlying model for generating resistance values.
Studies of additional habitat specialists are needed to
verify our findings and to quantify the tradeoff between
cost and reliability of different suitability estimates.
Acknowledgments We thank Ester Rubin, AZGFD, for
arranging use of data, Christopher Coffey for computer cluster
support, Jeff Jenness for ArcGIS support, and Kathy Zeller for
advice on the analyses. NAU School of Forestry, McIntire-
Stennis Cooperative Forestry Research Program, Arizona
Board of Forestry, and the Hafen, Krimminger, Prather, Czak,
Berry, and Forestry Faculty scholarships at Northern Arizona
University supported A.T.H.K. during this work. Sam Cushman,
Carol Chambers, and two anonymous reviewers helped to
significantly improve the manuscript. Ungulate data were
collected by AZGFD with funding from Wildlife Restoration
Act, Special Big Game License Tag Funds raised by the Arizona
Desert Bighorn Sheep Society, Arizona Elk Society, Rocky
Mountain Elk Foundation, Arizona Antelope Foundation, and
Arizona Big Game Super Raffle.
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