European Journal of Wildlife Research (2020) 66: 69

https://doi.org/10.1007/s10344-020-01385-y

ORIGINAL ARTICLE

Effective corridor width: linking the spatial ecology of wildlife

with land use policy
Adam T. Ford 1 & Emily J. Sunter 1 & Catherine Fauvelle 2 & Jacob L. Bradshaw 3 & Brett Ford 1 & Jenna Hutchen 1 &
Nicholas Phillipow 1 & Kristine J. Teichman 1

Received: 22 August 2019 / Revised: 15 April 2020 / Accepted: 29 May 2020 / Published online: 29 July 2020
# Springer-Verlag GmbH Germany, part of Springer Nature 2020

Abstract
Wildlife corridors are a policy tool to maintain or restore connectivity through human-modified landscapes. While corridors have
been an important concept in wildlife conservation for decades, there is little evidence to guide their design. Specifically, width is
one of the most contentious dimensions of corridors because it often limits industrial, residential, or recreational expansion.
Building on concepts that describe how wildlife avoid infrastructure or people—zone of influence and the flight initiation
distance—we present a conceptual approach to design and assess the management effectiveness of wildlife corridors through
a concept we define as effective corridor width. The effective corridor width is the minimum spatial dimension needed to abate
human influence on animal movement through the corridor. We searched for case studies documenting the zone of influence and
the flight initiation distance from recreational trails and residential development for four species of carnivore: black bears, grizzly
(brown) bears, gray wolves, and cougars. From these studies, we synthesized information on the effective corridor width, which
varied from 3000 to 6000 m for residential areas and 400 to 1000 m for trails, depending on the species. We examined the
effective corridor width in a case study from Alberta, Canada, where debate over the appropriate width of designated wildlife
corridors has been ongoing for decades. Our analysis is a step towards developing robust, empirically derived guidelines for
designing and managing wildlife corridors.

Keywords Carnivore . Connectivity conservation . Flight initiation distance . Wildlife . Zone of influence

Introduction 2014; Lamb et al. 2017). To help counter these negative ef-
fects, jurisdictions have designated “wildlife corridors” as a
Wildlife populations are negatively affected by human activ- part of their land use planning. Designated wildlife corridors
ities such as agriculture, urbanization, and recreation refer to the policy or regulatory tools that are intended to
(Woodroffe 2000; Polfus and Krausman 2012; Ripple et al. maintain the flow of animals, genes, or ecological processes
through human-dominated areas. This policy-related concept
This article is part of the Topical Collection on Road Ecology of wildlife corridors is distinct from that of “animal movement
Guest Editor: Marcello D’Amico corridors,” which are areas that have a high concentration of
Electronic supplementary material The online version of this article
animal movement pathways (LaPoint et al. 2013; Keeley et al.
(https://doi.org/10.1007/s10344-020-01385-y) contains supplementary 2017; Scharf et al. 2018).
material, which is available to authorized users. Designated wildlife corridors and animal movement corri-
dors may not always co-occur. For example, corridors may be
* Adam T. Ford designated in areas that have lower potential for economic
[email protected] development, e.g., the valley sides with steep slopes or
flood-prone areas, but these areas may be less preferred habitat
1
Department of Biology, University of British Columbia, Okanagan for some wildlife species (Hilty et al. 2019). Moreover, and
Campus Kelowna BC Canada unlike designated wildlife corridors, the location of animal
2
Department of Biology, University of Calgary, Calgary AB Canada movement corridors may not be known to managers when
3
Ecosystem Science and Management, University of Northern British land use decisions are being made. High-resolution tracking
Columbia, Prince George BC Canada of individual animals has been suggested as one of the only
69 Page 2 of 10 Eur J Wildl Res (2020) 66: 69

means to quantify animal-defined corridors (LaPoint et al. to design and assess the management effectiveness of wildlife
2013). Such fine-scale data can be impractical to collect be- corridors through a concept we define as effective corridor
cause of the cost of acquiring and deploying GPS collars and width. The effective corridor width is the minimum spatial
the time required to collect and analyze GPS collar data. dimension needed to abate human influence on animal move-
Despite these challenges, an understanding of how animal ment through the corridor. Second, we briefly synthesize re-
movements are affected by corridor design (i.e., by land use search on how residential areas and non-motorized recreation-
decisions within and adjacent to policy corridors) is a critical al trails influence the spatial ecology of four species of large
knowledge gap affecting conservation and economic objec- carnivore: cougar (Puma concolor), black bear (Ursus
tives (Kennett 2005). americanus), grizzly bear (Ursus arctos spp.), and gray wolf
Width appears to be one of the most uncertain and conten- (Canis lupus). We focus on these two sources of disturbance
tious considerations in designing wildlife corridors (Beier because they commonly occur near corridors (i.e., for residen-
et al. 2008; Gregory and Beier 2014; Beier 2018). Corridor tial development) or within it (i.e., for recreational trails). We
width influences restrictions on land use—the wider the cor- focus on these four species because they are widespread and
ridor, the more restrictive land use becomes, potentially threat- often are the focus of conservation and management planning
ening economic opportunities (Boisjolie et al. 2017; Wilhere in human-occupied landscapes (Berier 1995, Carroll et al.
and Quinn 2018). In spite of its impact on policy and econom- 2001; Gutzwiller et al. 2017; McDonald et al. 2018). Third,
ic interests, there are few evidence-based guidelines for how we integrate the effective corridor width concept with data
wide a wildlife corridor should be in order to be effective. from the synthesis to create recommendations for corridor
Intuitively, a wider corridor ought to convey more move- design, linking to a case study in Alberta, Canada, where
ment than a narrower one, yet an animal only needs a space discussions on corridor design have been ongoing for decades.
wider than its body for passage to occur (Ford et al. 2017). We also highlight where the effective corridor width concept
Beier (1995) suggested that a 100-m-wide corridor is needed requires further investigation.
for cougars over short distances (800 m), but wider corridors
(400 m) are needed for longer corridors (> 1 km in length). The effective corridor width concept
Harrison (2010) suggested using the species home range to
design the width of corridors. For a number of species, and We draw upon the concepts such as the “zone of influence”
based on considerations of home range dimensions and (ZOI) (Boulanger et al. 2012) and “flight initiation distance”
vagility, (Beier (2018) suggests a “rule of thumb” of a 2-km (FID) to inform wildlife corridor width. In this case, the ZOI
width to accommodate gene flow and recolonization. For cor- refers to an effect of human activity or infrastructure that is
ridors created over or under roads, width is an important attri- projected over space and onto ecological processes (Fig. 1).
bute that positively predicts passage rates (Clevenger and FID refers to the distance at which animals flee from an ap-
Waltho 2000), but even small (~ 2 m wide) passages are used proaching person. Concepts related to the ZOI/FID are vari-
often by large carnivores like grizzly bears (Ford et al. 2017). ously referred to as the “core-area model” (Laurance and
In some systems, width has been shown to have no effect (La Yensen 1991), the “edge effect” (Lidicker 1999), “the road-
Polla and Barrett 1993; Lindenmayer and Nix 1993), a non- effect zone” (Forman and Alexander 1998), and the “human
linear effect (Andreassen et al. 1996), or a positive effect on shield” (Berger 2007). ZOI typically refers to a response to
biodiversity (Haddad 1999; Damschen et al. 2008). Ideally, anthropogenic disturbance or infrastructure (e.g., houses,
rigorously derived evidence (e.g., experimental titration of roads, trails), whereas FID usually refers to the response to
corridor dimensions) on corridor width effects would be avail- people per se.
able for focal taxa and used to guide corridor planning. For The effective corridor width is the residual space that oc-
some species—like large terrestrial mammals—the applica- curs beyond the ZOI/FID of human activity within a designat-
tion of such controlled or experimental approaches is not fea- ed wildlife (i.e., policy) corridor. If, for example, a 400 m wide
sible, yet the demand for evidence in decisions regarding cor- designated wildlife corridor is adjacent to a residential devel-
ridor designation and regulation persists. opment, and the ZOI from that development is 100 m, then the
Creating and managing corridors through land use plan- effective corridor width is 300 m (Fig. 1). In this case, the
ning, development bylaws, and other regulatory approaches adjacent development causes a 25% loss in the effective cor-
is aided by empirical guidance (Krizek et al. 2009). Such ridor width. The ZOI/FID is species and context dependent;
empirical guidance is needed for corridor policy in many for example, the FID of habituated or foraging grizzly bears
areas. Here, we present one approach that can help fill the may be a few meters while foraging (Albert and Bowyer
corridor design knowledge gap, particularly in cases where 1992), yet this species is known to avoid denning near human
data on animal-defined corridors are unavailable and where settlements by upwards of 4 km (Whiteman et al. 2017).
land use changes are proposed within or adjacent to existing Likewise, the habitat needs and sensitivity to human activity
designated corridors. First, we present a conceptual approach for adults with young may differ from that of dispersing
Eur J Wildl Res (2020) 66: 69 Page 3 of 10 69

Whittington 2006) and camera trapping (Muhly et al. 2011),

whereas animal-based methods include backtracking
(Whittington et al. 2005) and telemetry (Chetkiewicz and
Boyce 2009; Rogala et al. 2011). Response variables, such
as habitat selection, occupancy, kill rates, or other behavioral
outcomes, have been recorded as part of ZOI (Table S1 and
S2) and used in connectivity research globally (Lyra-Jorge
et al. 2010; Almasieh et al. 2019).
There are three ways through which the ZOI and FID in-
form effective corridor width: direction, magnitude, and dis-
tance. The directional effect can be positive (e.g., animals
select for areas near towns), negative (e.g., animals avoid
towns), or neutral (e.g., animals use areas near towns in pro-
portion to their availability). The magnitude can be strong,
weak, or neutral referring to the effect that distance has on
the response variable. The distance effect is the spatial extent
within which significant changes in the response variable are
observed. The distance, magnitude, and direction of the ZOI
can be nonlinear, with peak positive selection occurring at
intermediate distances from the disturbance (Dyer et al. 2001).
The effective corridor width provides an evidence-based
approach to design corridors, to guide corridor management,
and to estimate how different land use scenarios may affect
wildlife. For example, the effective corridor width can inform
setbacks or mitigations to reduce the impacts of urban devel-
opment adjacent to a designated wildlife corridor. However,
the effective corridor width does not, on its own, prescribe if
such land use is “too much” or “within acceptable limits.”
Instead, the effective corridor width provides a measure of
potential changes in corridor function. To make use of the
effective corridor width in policy, we suggest that decision-
makers establish a priori what the acceptable limits of change
are before planning land use adjacent to, or within, a desig-
nated wildlife corridor.

Linking human activity to corridor design: a case

study for large carnivores

Fig. 1 The effective corridor width concept. In these examples, the To demonstrate how the effective corridor width can be esti-
yellow area is the effective corridor width that lies outside of the zone mated, we performed a literature review to find examples
of influence (orange shading) of a residential areas occurring on either where the spatial response of four large carnivores has been
side of the designated wildlife corridor; b residential areas occurring on quantified. We used the Institute of Science’s Web of Science
one side of a designated wildlife corridor and a low-use (e.g., steep
slopes) natural feature on the other side; c a recreational trail occurring as well as Google Scholar to survey published literature. We
on one side of a designated wildlife corridor and a low-use natural feature searched for these papers during 2016 and 2017 and included
on the other side; and d residential areas occurring on one side of a papers published from 1985 to 2017. We used the following
designated wildlife corridor, recreation trails occurring within the corri- terms to structure our query: carnivore, predator, cougar,
dor, and a low-use natural feature on the other side of the corridor
wolf, bear, residen* (yielding both residential and residence),
recreation, trail, bik* (yielding biking, biker, and bike), and
individuals, giving rise to corridor prescriptions optimized for anthropogenic disturbance. We recorded whether studies re-
life history requirements (Ford et al. 2017). ported a positive, negative, or neutral ZOI/FID and, if avail-
For many wildlife species, the ZOI can be estimated from a able, determined the distances at which ZOI and FID values
number of place-based and animal-based methods. Place- were detected. Methods used to quantify the ZOI included
based approaches include snow transects (Shepherd and resource selection functions (41% of studies), occurrence
69 Page 4 of 10 Eur J Wildl Res (2020) 66: 69

(23%), utilization (18%), mortality (8%), and occupancy (5%) (n = 13) reported a distance over which the negative response
(Table S3). Further details of this search are available in the occurs, with a range of 1–7 studies per species.
Supporting Information. Using the reported negative (avoidance) values from these
We found 39 studies examining the ZOI for residential studies, we estimated the effective corridor width for the four
development (range: 4–16 studies per species, with a mean carnivore species. ZOI distances for residential development
of 44 individuals per study) (Supporting Information; ranged from a low of 50 m (cougars) to a maximum of 8000 m
Table S1, S2). These studies ranged in sample size from 5 for grizzly bears (Table 1). The species-specific median ZOI
(black bears) to 362 (grizzly bears) individuals per study. for residential development ranged from a low of 3105 m
Across studies on all species, 74% found a negative direction- (cougar) to 6000 m (grizzly bear), for a median, cross-
al response of residential development, 30% positive, and species ZOI of 4413 m. This means that wildlife corridors
21% neutral (Fig. 2). Of the 39 studies reporting a ZOI for need to be at least 4400 m wide to have an effective width
residential development, 28% (n = 11) reported a distance that is greater than 0 m when adjacent to urban development.
over which the negative response occurs (Table S2). For trails, negative ZOI distances ranged from 21 m (grizzly
For recreational trails, we found 19 studies reporting a ZOI bears) to 2000 m (black bears). The species-specific median
(range: 3–9 studies per species) with a mean of 11 individuals ZOI for trails ranged from a low of 100 m (cougar) to a max-
per study. Grizzly bears (47% of studies) were the most com- imum of 732 m (black bear), for a median, cross-species ZOI
monly studied species, followed by wolves (4 studies), and of 465 m. This means, on average, that wildlife corridors need
black bears and cougars at 3 studies each. Across species, to be at least 465 m wide to have an effective width that is
there was a negative ZOI for recreational trails in 68% of greater than 0 m when adjacent to recreational trails.
studies, 42% positive, and 42% neutral. These values exceed In addition to responses to residential development and trails,
100% because more than one response can be detected within our search revealed fine-scale responses of wildlife to an ap-
a given study. Of the 19 studies reporting a ZOI for trails, 68% proaching human (FID). The distance at which animals flee an
approaching human ranged from 21 to 157 m for grizzly bears,
17–488 m for wolves, and 10–25 m for black bears (Fig. 3). We
found no studies reporting FID for cougars. Like the ZOI, the
FID provides an estimate of how much space wildlife require in
order to move naturally through the landscape.

A test case for the effective corridor width: the Bow

Valley, Alberta, Canada

In the Bow Valley (Alberta, Canada), the effect of urban expan-

sion on the movement and persistence of large mammals has
been a key concern for governments, the public, and developers
for decades. In the 1990s, a ruling by a judicial review (the
National Resources Conservation Board) stated that wildlife cor-
ridors must be used to mitigate the impacts of a proposed ~
15,000 person development in a town that, at the time, had a
population of 6000 (Kennett 2005). The ruling included some
corridor design guidelines, including a minimum width of
350 m. The 350-m width guideline has been supported by a local,
multi-stakeholder working group that formed in 1995 to help
guide development in the area (BCEAG 2016). However, ongo-
ing deliberation between the community, government, and de-
velopers is testing the effectiveness and underlying evidence of
the 350 m width. For example, in a recent decision (July 2018)
Fig. 2 The spatial response of large carnivores to a residential areas and b
recreation trails, derived from published studies. Positive responses are
made by the local land regulator, an urban development proposal
taken from the zone of influence and indicate selection for the feature or in this area was rejected because it changed the width of one of
lower mortality/conflict, as distance to the feature decreases. Negative the designated wildlife corridors in the area to less than 350 m in
responses indicate avoidance of the feature and higher mortality/ some places. The decision letter indicated that if the width (i.e., in
conflict, as distance to the feature decreases. Neutral responses indicate
that no effect of the feature was observed. A mixed result occurred when
a revised proposal) could be widened to between 400 and 450 m,
there were at least two different directional responses, such as age, sex, or there would be “…enough room for wildlife to move through
diel period-specific effects this area unimpeded” (see Supporting Information).
Eur J Wildl Res (2020) 66: 69 Page 5 of 10 69

Table 1 The zone of influence

distance associated with Disturbance Species Range (m) Median (m) Number of studies
avoidance of residential areas and
recreational trails Residential areas Cougar 50–9990 3105 3
Wolves 1000–6100 3546 3
Grizzly bear 4000–8000 6000 4
Black bear 5000 5000 1
Trails Cougar 100 100 1
Wolves 400 400 1
Grizzly bear 21–8000 628 7
Black bear 122–2000 732 2

Details of the studies found in this review are described in the Supporting Information

Based on our review of ZOI and FIDs, a 350-m-wide cor- infrastructure and human activity on the movement of wild-
ridor has no effective corridor width for the four species of life. Here, we integrated ZOI and FID into the concept of
carnivore if urban development and recreational trails are lo- effective corridor width, which can be used to evaluate man-
cated adjacent to the corridor (Fig. 4; Table 1). In addition, the agement effectiveness of land use policies inside and adjacent
FID of large carnivores was as low as 25 m in some cases but to designated wildlife corridors.
extended up to 490 m, suggesting that the influence of direct We envision at least two applications of the effective corridor
encounters with people in or adjacent to the corridor can in- width in wildlife conservation and land use policy. First, we see
fluence significant portions of its 350-m width. This finding the predominately negative (Fig. 2) response of large carnivores
does not mean that the animals will be absent from the Bow to residential development and recreational trails (but see wolves)
Valley corridors (e.g., see Chetkiewicz and Boyce 2009), but as justification for a precautionary approach to land use planning
that people are very likely exerting a negative influence on in areas where the movement of wildlife is a priority for man-
wildlife behavior within the areas specifically designated to agement. Specifically, corridors less than 4 km wide will likely
facilitate animal movement. Mitigations targeted at reducing be negatively influenced by urban development along their
residential development and human activity within and adja- boundaries for cougars, wolves, and black and grizzly bears
cent to the corridor may be needed to maximize the effective- (Fig. 4). The strength of this negative influence is not well-
ness of this space for animal movement. known and moving forward, a key knowledge gap to quantify
just how much, and under what context is, wildlife movement
affected by urban development, trails, and other disturbances.
Discussion Though we found many cases of negative effects of residen-
tial development, we also found ~ 55% of cases where the ZOI
Concepts such as the zone of influence (ZOI), flight initiation was positive or mixed. For example, wolves tended to select for
distance (FID), and related approaches to understanding edge recreational trails in many cases (Fig. 2). In situations where
effects can help inform policies that minimize the impact of residential developments create an ecological trap by attracting

Fig. 3 The flight initiation distances (FID) of North American carnivores. was no flight initiation distances identified for cougars. The figure repre-
FID can be used to assess the effective corridor width, especially for sents the median value for two studies (McLellan and Shackleton 1989;
recreational trails or other areas where people are active in the outdoors. Reimchen 1998), while the others show the mean reported by each study
References for studies are provided in the Supporting Information. There
69 Page 6 of 10 Eur J Wildl Res (2020) 66: 69

Fig. 4 The effective corridor

width for large carnivores based
on the median, negative zone of
influence for a residential
development and b recreational
trails (see Table 1 for values). To
illustrate this concept, we simu-
lated a designated wildlife corri-
dor with a width of 7000 m for
residential areas and 3500 m for
recreational trails. The species-
species effective corridor width is
shown as the distance from
boundary of the zone of influence
to the corridor edge farthest from
human activity

wildlife and facilitating human-wildlife interactions (Lamb et al. riparian areas to regional habitat linkages (Hilty et al. 2019).
2017), corridors will need to be much wider than this “zone of Measuring success across these scales must match appropriate
attraction.” Wide corridors may therefore help facilitate animal levels of ecological process, from foraging behavior in small-
movement by providing space for people and wildlife to avoid scale corridors to gene flow and population persistence through
one another. Corridors less than 465 m wide are likely going to regional linkages (Beier and Loe 1992; Beier et al. 2008). The
be negatively influenced by recreational trails along their bound- effective corridor width can be applied to these scale-dependent
aries for cougars, wolves, and black and grizzly bears. Finally, process to help guide policy. For example, urban area ZOI dis-
and based on the FID review, most species’ movement in corri- tances can be applied for large-scale (100 s of km) linkages and
dors less than 400 m wide will be negatively influenced by the regional plans (Chester 2015), while recreational trail-based ZOI
presence of people along the edge of the corridor. Corridor and FID could be used to justify regulation of the timing and
widths need to be even greater if residential development, trails, location of trail closures occurring inside or adjacent to corridors
or human activity are permitted within the boundaries of the (10 s of meters). In the Bow Valley, AB, case study described
designated wildlife corridor. here, the “urban development” ZOI influences the entire valley
Second, the effective corridor width can be informed by scale- floor, where all four carnivores species we reviewed occur
specific land use decisions. Broadly speaking, land use planning (Barrueto et al. 2014). In this case, we suggest that measuring
occurs at a variety of scales, from where to prioritize biodiversity the impacts of potential urban developments on corridor effec-
conservation in a region (e.g., national parks) (Coristine et al. tiveness needs to consider finer-scale spatial processes and indi-
2018), demarcating the boundaries of development (Fleury and cators, such as the FID.
Brown 1997), regulating human activity via bylaws, and plan- The effective corridor width builds on existing and well-
ning the route of a new recreational trail (Shepherd and established concepts in spatial ecology to help guide
Whittington 2006). These scales invoke different policy and legal evidenced-based land use policy aimed at preserving or restoring
mechanisms that regulate or guide development (Boisjolie et al. animal movement. The effective corridor width is therefore an
2017). Similarly, wildlife corridors can range from small-scale important advance for systems where there is demand for
Eur J Wildl Res (2020) 66: 69 Page 7 of 10 69

evidence or where expert opinion has heretofore driven policy. which selection or avoidance occurs (in some cases even when
Moving forward, there are at least five areas of further investiga- including distance to housing or trails as a covariate,(e.g., Hiller
tion that will enhance the utility of effective corridor width con- et al. 2015), making it difficult to calculate a ZOI from these
cept. First, the effective corridor width does not answer the ques- studies. In the future, studies seeking to translate knowledge
tion: what does the minimum width of a wildlife corridor need to from movement ecology into management and policy (Fraser
be? Instead, the effective corridor width rather answers: to what et al. 2018) should provide model outcomes that can be more
extent will this corridor remain effective under different develop- clearly interpreted in management plans, for example, the pro-
ment scenarios? However, Beier (2018) suggested that the ques- portional reduction in space use by animals as a function of
tion of how narrow a corridor can be while retaining ecological distance to human disturbance.
functions is “….analogous to asking an engineer, what are the Fourth, our review did not address the combined impacts of
fewest number of rivets that might keep this wing on the air- multiple sources of disturbance or myriad other forms of human
plane?” Nonetheless, the tone of many land use discussions, such activity that influence wildlife. Housing density and amount of
as the Bow Valley case study we described (Kennett 2005), trail use affect wildlife movements (Rogala et al. 2011; Ladle
reflects the very real tension imposed by economic, social, and et al. 2018a), as do roads, mines, and motorized recreation
conservation objectives whose manifestation is incompatible at (Tucker et al. 2018). These sources of human activity likely
the same location. Until this tension is resolved, there will be an would have zones of influence that are distinct individually and
ongoing need to know if corridors are too narrow for their cumulatively, from the two sources of disturbance we examined
intended purpose. in this study. Regional or global patterns in the cumulative effects
Second, our review focused on the spatial responses of from multiple sources of disturbance are associated with range
large carnivores, but did not address specific ecological pro- contractions in some large mammals (Shackelford et al. 2018),
cesses or flows. For example, a corridor designed for foraging diel activity (Gaynor et al. 2018), and changes in movement
may have different attributes, including width, than one de- propensity (Tucker et al. 2018). Given that these cumulative
signed for dispersal (Abrahms et al. 2017). Corridors and ad- changes to the landscape are increasing globally (Venter et al.
jacent human activity may also influence predator-prey or 2016), further research is needed to understand how co-occurring
other biotic interactions (Ladle et al. 2018b). For example, disturbance types influences fine-scale animal movement in the
areas avoided by predators may attract prey, creating localized context of corridor design.
trophic cascades that negatively impact non-focal species like Fifth, we recognize that animals may adjust their behavior
songbirds and beavers (Hebblewhite et al. 2005). Ford et al. in such a manner that would make them resilient to land use
(2017) found that grizzly bear use of wildlife corridors built cha nges an d human activity near c orridors. For
across roads varied by reproductive status and sex, with family example,(Gaynor et al. (2018) found that wildlife have be-
groups preferring wider corridors than singletons. In come more nocturnal as human activity increases. This sug-
carnivore-rich areas such as Alberta’s Bow Valley, we may gests that animals may continue to use corridors frequented by
also anticipate that intraguild (between carnivore species) in- people by shifting activity to periods when people are less
teractions could influence the effectiveness of corridors. For active. Wildlife vary in their tolerance or capacity to habituate
example, wolves are known to displace cougars (Kortello to people, and this may influence the ZOI/FID over time. For
et al. 2007) and coyotes (Merkle et al. 2009) from kills, while example,Barrueto et al. (2014) found that black bears and
grizzly bears likely are dominant in direct interactions with wolves avoided wildlife crossing structures used frequently
black bears (Mattson et al. 1992). Thus, for corridor designs by people, while cougars and grizzly bears showed only weak
targeted at multiple species, it will be important to not only responses to human activity. Further work is needed to under-
account for the influence of a human-caused ZOI and FID but stand which species are likely to accommodate changes in
also allow enough room so that species interactions do not land use and human activity in or near corridors.
interfere with the objectives of the corridor (e.g., the flow of
animals). These process- and demographic-specific design
criteria will continue to emerge as researchers gain better un- Conclusion
derstanding of movement as an ecological process (Nathan
et al. 2008) and the need to integrate trophic interactions in The effective corridor width provides a starting place to over-
landscape restoration practices (Fraser et al. 2015). lay evidence of species biology with the “landscape of policy”
Third, we did not systematically address the strength of re- that is used to guide the management of designated wildlife
sponses to residential development or trails in our review. We corridors. Data on how species respond to edges has been
relied on reported measures of statistical significance, some of published widely, including for ungulates and urban areas
which reported the odds ratio to quantify the per unit change in (Polfus and Krausman 2012), various species and roads
use as a function of distance to infrastructure. Most of the stud- (Forman and Alexander 1998), and other species (reviewed
ies in our review did not explicitly provide a distance over in Beier 2018). Applying these broad streams of evidence to
69 Page 8 of 10 Eur J Wildl Res (2020) 66: 69

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