Home Range and Landscape Use of Coyotes in a Metropolitan Landscape: Conflict or Coexistence?

Abstract

An understanding of how top mammalian carnivores respond to urbanization is important for conservation and management of human-wildlife conflicts. Coyotes (Canis latrans) have recently become more prevalent in many metropolitan areas; however, their apparent success is poorly understood. We estimated home-range size and selection of land-use types for coyotes in a heavily urbanized landscape, with a particular focus on responses of coyotes to those parts of the urban landscape with high levels of human development or activity. Mean (6 SE) annual home ranges of transient coyotes (X ¯ 5 26.80 6 2.95 km2) were larger than those of resident coyotes (X ¯ 5 4.95 6 0.34 km2), and home-range size for resident coyotes did not vary among seasons or between age and sex classes. Although most home ranges were associated with natural patches of habitat, there was considerable variation among coyotes, with some home ranges entirely lacking patches of natural habitat. Within home ranges, coyotes typically avoided land-use types associated with human activity (i.e., Residential, Urban Grass, and Urban Land) regardless of coyote characteristics, seasons, and activity periods. Few coyotes were nuisances, and conflicts occurred when coyotes were sick or exposed to wildlife feeding by humans. We found little evidence that coyotes were attracted to areas associated with human activity, despite at times having home ranges located in heavily developed areas.

Full text

HOME RANGE AND LANDSCAPE USE OF COYOTES IN A
METROPOLITAN LANDSCAPE: CONFLICT OR COEXISTENCE?
STANLEY D. GEHRT,* CHRIS ANCHOR, AND LYNSEY A. WHITE
School of Environment and Natural Resources, Ohio State University, 2021 Coffey Road, Columbus, OH 43210, USA
(SDG, LAW)
Forest Preserve District of Cook County, Route 4, Box 178, 28W040 State Route 58, Elgin, IL 60120, USA (CA)
An understanding of how top mammalian carnivores respond to urbanization is important for conservation and
management of human–wildlife conflicts. Coyotes (Canis latrans) have recently become more prevalent in
many metropolitan areas; however, their apparent success is poorly understood. We estimated home-range size
and selection of land-use types for coyotes in a heavily urbanized landscape, with a particular focus on
responses of coyotes to those parts of the urban landscape with high levels of human development or activity.
Mean (6 SE) annual home ranges of transient coyotes (X
¯
5 26.80 6 2.95 km
2
) were larger than those of
resident coyotes (X
¯
5 4.95 6 0.34 km
2
), and home-range size for resident coyotes did not vary among seasons
or between age and sex classes. Although most home ranges were associated with natural patches of habitat,
there was considerable variation among coyotes, with some home ranges entirely lacking patches of natural
habitat. Within home ranges, coyotes typically avoided land-use types associated with human activity (i.e.,
Residential, Urban Grass, and Urban Land) regardless of coyote characteristics, seasons, and activity periods.
Few coyotes were nuisances, and conflicts occurred when coyotes were sick or exposed to wildlife feeding by
humans. We found little evidence that coyotes were attracted to areas associated with human activity, despite at
times having home ranges located in heavily developed areas.
Key words: Canis latrans, Chicago, coyote, Illinois, resource selection, urbanization
A fundamental question regarding the presence of wildlife
in urban landscapes is whether they are attracted to human
activities and somehow benefit from urban areas (i.e.,
synanthropic species—Johnston 2001), or alternatively occur
in urban areas despite possible negative effects of human-
dominated areas, and thereby require habitat fragments
protected from development. Large carnivores tend to
disappear from areas dominated by humans, either through
direct persecution, competition for resources, or diminution of
resources (Cardillo et al. 2004; Woodroffe 2000; but see
Linnell et al. 2001). This trend would be expected to be
particularly apparent in urban landscapes, where habitat
fragmentation is extensive and human presence is the greatest,
and conflicts between carnivores and humans would seem to
be acute. Indeed, human intolerance or carnivore conflict may
prohibit larger carnivore species from becoming synanthropic.
Coyotes (Canis latrans) have dramatically expanded their
range across North America, in contrast to other, larger
Carnivora (Gompper 2002; Laliberte and Ripple 2004), and
are now found in an increasing number of cities in the United
States and Canada (Gehrt 2004). The emergence of coyotes in
urban systems can have important ecological implications
(Gehrt and Riley, in press; Gompper 2002), such as through
their role as an apex carnivore and subsequent effects on prey
or the creation of trophic cascades (Crooks and Soule´ 1999).
However, the emergence of coyotes in urban landscapes also
has increasingly resulted in conflicts with people (Gehrt 2004;
Gehrt and Riley, in press). Coyotes have the potential to attack
pets and, at times, people (Bounds and Shaw 1994; Carbyn
1989). Thus, the existence of coyotes in proximity to people
can create concern among the public and present dilemmas for
management agencies. An understanding of how coyotes
respond to urban areas, especially landscapes dominated by
high levels of human activities, is important for conservation
and management efforts.
The appearance of coyotes in urban areas would suggest
that they are relatively flexible in their use of the landscape,
while maintaining a reliance on natural habitats. However,
research to date has provided mixed results as to whether
coyotes are a true synanthropic species. Monitoring of track
stations has suggested that coyotes may have restrictions in
* Correspondent: gehrt.1@osu.edu
E
2009 American Society of Mammalogists
www.mammalogy.org
Journal of Mammalogy, 90(5):1045–1057, 2009
1045

their use of natural habitat patches or developed patches
within the urban matrix (Crooks 2002; Randa and Yunger
2006). But monitoring track stations provides only limited
information regarding use of developed areas, and provides no
information on the types of individuals that are likely to occur
in different parts of the landscape. Radiotelemetry studies of
coyotes in urban areas have produced mixed results, with
some studies reporting coyote avoidance of residential areas or
other types of development (Quinn 1997; Riley et al. 2003)
and others reporting use only at levels of availability (Gibeau
1998; Grinder and Krausman 2001; Way et al. 2004).
However, these studies were limited in sample size and
conducted near the perimeter of developed areas, in areas with
large tracts of rural or undeveloped use. Coyote behavior may
differ in areas with limited natural habitat and intense human
presence. Additionally, coyote use of the landscape may vary
in association with characteristics related to sex, age, social
status, or season, any of which may be difficult to identify
with small sample sizes.
Herein, we report on coyote use of the landscape within the
Chicago metropolitan area, one of the largest urban centers in
North America. Although common in the surrounding rural
landscape, coyotes were rare in the metropolitan area until the
1990s (Gehrt 2004). An apparent consequence of their success
was a dramatic increase in the number of coyotes captured for
nuisance control. Before 1990, .20 individuals were typically
removed from the area annually, and this increased to .300
annually within 10 years (Gehrt 2004). Given the level of
development, size of the human population, and an apparently
increasing coyote population, the Chicago area represents an
excellent opportunity to investigate how carnivores exploit an
urban landscape, and the implications for the humans sharing
that space.
We used patterns of home-range size and landscape use
within a heavily urbanized landscape to test predictions related
to synanthropy in coyotes. Given the close relationship
between home-range size and resource quality and distribution
for carnivores (Gittleman and Harvey 1982), we predicted
relatively smaller home ranges and no avoidance of developed
areas if coyotes thrive in urban landscapes. Other species
considered to be synanthropic, such as kit foxes (Vulpes
macrotis), red foxes (V. vulpes), and raccoons (Procyon lotor),
have smaller home ranges in urban landscapes (Cavallini
1996; Cypher, in press; Prange et al. 2004), and some fox
species readily use developed parts of the landscape (Cypher,
in press; Newman et al. 2003). Likewise, we predicted no
relationship between home-range size and the proportion of
the home range comprising developed land. However, if
coyotes avoid developed areas and concentrate activities in
natural habitat patches, we would predict a positive relation-
ship between home-range size and the proportion of
development in the home range. For both home-range analyses
and patterns of landscape use, we construct a series of tests
layered by potentially important covariates to the use of urban
landscapes, including season; activity period; and age, sex,
and social class of the coyote. We also provide the frequency
and characteristics of nuisance coyotes, or those that come into
apparent conflict with people. If urban coyotes develop an
affinity for developed areas, we predict that a relatively high
proportion of coyotes will be removed as nuisances because of
human fear of coyotes (Gehrt and Riley, in press). Finally, we
discuss the summary of these results with respect to the
following question: Do coyotes reside in urban areas because
of an attraction to human activities, or in spite of them?
MATERIALS AND METHODS
Study area.—The Chicago metropolitan area spans all or
parts of 6 counties in northeastern Illinois (Cook, DuPage,
Kane, Lake, McHenry, and Will), and extends into parts of
Wisconsin and Indiana. The 6 counties include .260
municipalities and a cumulative human population exceeding
9 million, making this one of the most heavily urbanized areas
in North America. General land cover in 1997 for this area was
estimated to be 33% agriculture, 30% urban, 16% natural
areas, and 21% unassociated vegetation (Wang and Moskovits
2001). Natural areas (including savannas, woodlands, grass-
lands, and wetlands) were fragmented, 1st by agriculture in the
early 1800s, and more recently through urbanization. The
extensive process of urbanization has produced a dynamic
landscape in these counties, especially recently. Between 1972
and 1997, urban land increased 49%, natural areas decreased
21%, and agricultural lands decreased 37% (Wang and
Moskovits 2001). An important feature of this landscape is
the patchwork of habitat fragments protected from develop-
ment, most of which are county forest preserves (Gehrt and
Chelsvig 2003). For example, forest preserves make up 11%
of the land area of Cook County, Illinois, providing an
important component of the landscape mosaic in addition to
the .5 million human inhabitants in that county.
Our fieldwork was largely focused in the northwestern
portion of the metropolitan area, including O’Hare Interna-
tional Airport (Fig. 1). The scope of the study area was
determined by the cumulative area of locations of radio-
collared, resident coyotes, which spanned approximately
1,173 km
2
. It is important to note that this study area occurred
within the urban matrix, in contrast to previous studies of
coyotes conducted at the periphery of urban areas. Our study
area had a paved road density of 6.11 km/km
2
, with traffic
volumes exceeding 100,000 vehicles daily for some roadways
(Illinois Department of Transportation, Springfield, Illinois),
and was composed of the following land-use types: agriculture
(14%), natural habitat (13%), residential (20%), urban land
(including commercial–industrial use, 43%), and other (10%).
Live captures.—Because of the constraints associated with
working in public areas, our trapping was largely opportunis-
tic. It was necessary to focus our trapping in areas that
afforded some seclusion from the public. In most cases these
were secure areas within large forest preserves, or private
properties. Trapping was conducted opportunistically through-
out the year excluding summer months when pups were
emerging from dens. Coyotes were livetrapped with padded
1046 JOURNAL OF MAMMALOGY Vol. 90, No. 5

foothold traps and cable restraint devices. Upon capture of an
unmarked individual, the coyote was usually transported to a
laboratory area and immobilized with an injection of Telazol
(Fort Dodge Animal Health, Fort Dodge, Iowa). Coyotes were
removed from the field to avoid people and pets while we
were processing animals. Coyotes were marked with uniquely
numbered plastic ear tags (NASCO Farm & Ranch, Fort
Atkinson, Wisconsin) and fitted with very-high-frequency
radiocollars (Advanced Telemetry Systems, Isanti, Minne-
sota). We weighed each coyote, and determined sex, age (via
tooth wear and reproductive condition), and physical condi-
tion. Once coyotes had recovered from immobilization, they
were released at the capture site. Our trapping and handling
protocols were approved by Ohio State University’s Institu-
tional Animal Care and Use Committee (IACUC 2003R0061),
and followed guidelines approved by the American Society of
Mammalogists (Gannon et al. 2007).
Radiotelemetry.—We obtained radiolocations for coyotes
using triangulation (with program LOCATE II; Pacer, Truro,
Nova Scotia, Canada) with a truck-mounted antenna or by
visual observations. We located coyotes once during the day,
typically 2 or 3 times per week, and at night during tracking
shifts in which we focused on a group of coyotes and obtained
sequential locations at 60- to 120-min intervals for 5–6 h
during the night. Early in the study, we determined that
coyotes confined most of their activity to nocturnal hours
(Morey 2004), which has been observed by virtually all
studies of urban coyotes (Atkinson and Shackleton 1991;
Gibeau 1998; Grinder and Krausman 2001; Quinn 1997; Riley
et al. 2003; Tigas et al. 2002). Mean (6 SD) error for test
transmitters was 108 6 87 m via triangulation (Morey 2004).
However, many of the coyotes were located within the urban
matrix, and because of the extensive road system it was
frequently possible to drive within a few meters of coyotes and
record their location directly with a global positioning system
unit. Coyote locations were recorded to the nearest meter
using the Universal Transverse Mercator grid system.
Home-range estimates.—We used the Home Range Exten-
sion (Hooge and Eichenlaub 1997) for ArcView 3.2
geographical information system software (Environmental
Systems Research Institute, Redlands, California) to plot
95% minimum convex polygon home-range estimates. We
found that kernel models produced disjunct home-range
boundaries for some coyotes that could not be smoothed
without expanding home-range boundaries to an unrealistic
size. This was especially true for coyotes with fragmented
home ranges in the urban matrix, a situation also encountered
by Riley et al. (2003). It was important to have continuous
home ranges to correctly estimate areas available for use by
coyotes within home ranges, and the minimum convex
polygon yielded the most conservative areas without disjunct
home-range boundaries.
We calculated annual home ranges for each coyote that had
a minimum of 47 radiolocations recorded during an annual
period (the minimum number of locations that spanned .1
season within an annual period). However, for transient
FIG.1.—Study area and land use within the Chicago, Illinois, metropolitan area.
October 2009 GEHRT ET AL.—URBAN COYOTE LANDSCAPE USE 1047

coyotes we used a lower minimum number of 30 locations
because of the difficulties associated with monitoring coyotes
with large home ranges in the metropolitan area, such as
locating telemetry signals, and because solitary individuals
sometimes dispersed and truncated our time to acquire
locations. Area-observation curves were not useful for
identifying minimum location numbers because of the fluid
nature of home ranges for transients. We classified a coyote as
a resident if it used 1 unique area for 1 biological season and
was observed with another coyote, and a transient if it
maintained a home range that overlapped multiple territories
of residents or was not observed associating with other coyotes
for .1 season (adapted from Gese et al. 1988). Home ranges
of residents were exclusive, whereas home ranges of transients
overlapped each other and those of residents (Gese et al. 1988;
Kamler and Gipson 2000). Coyotes that dispersed from the
study area were censored from data analysis.
For seasonal analyses, we partitioned the data into 3 periods
that corresponded to biological events: breeding (1 January–30
April), pup-rearing (1 May–31 August), and dispersal (1
September–31 December). We estimated seasonal home
ranges using a criterion of 30 locations recorded in a season,
providing that approximately equal numbers of locations were
obtained during daytime and nighttime periods.
Because of high survival and site fidelity, multiple annual
home ranges were calculated for some individuals. Thus, we
reduced the resident data set to only 1 estimate per coyote by
selecting the year with the greatest number of locations or
complete monitoring throughout the year. We used 2-way
fixed analysis of variance (ANOVA) with an interaction term
to assess whether sex or age explained variation in annual
home-range size. We subsequently pooled estimates across
sex–age groups to compare resident and solitary home ranges
with a t-test. Similarly, we determined if home-range sizes
vary by season among sex–age groups with a 3-way ANOVA
with sex, age, and season as main effects with interaction
terms. For each of these tests, home-range estimates were log-
transformed to conform to normal distributions.
We created a land-use–type coverage with 28.5-m resolu-
tion from 1997 Chicago Wilderness/NASA Landsat Thematic
Mapper images for use in ArcView geographical information
system software (Wang and Moskovits 2001). We reclassified
the original 164 Landsat categories into 8 broad land-cover
types: Agricultural (usually small fragments of row-crop land
use, but may also include small produce such as pumpkin
farms or vegetable gardens), Natural (fragments of natural
habitat typically protected from development, but often
exposed to extensive human use), Other (typically small areas
with a mix of developed and undeveloped properties, such as
golf courses or cemeteries), Residential (developed areas for
human residents), Urban Grass (managed lawns or parks,
including corporate campuses, mowed parks, or recreational
areas), Urban Land (industrial or commercial development,
often including a high degree of impervious surfaces),
Undeveloped (usually small fragments not managed for
wildlife, and either too small for development or in many
cases a buffer between developments, such as easements along
major thoroughfares), and Water (impoundments or streams,
often retention ponds resulting from development).
We used simple linear regression to determine the
relationship between the level of urbanization within a home
range and home-range size by determining the composite
proportion of Residential, Urban Grass, and Urban Land
categories within an annual home range, and regressing this
metric with home-range size. To satisfy assumptions of
normality, we log-transformed home-range size and used the
arcsine-transformed proportion of cumulative urban-associat-
ed land use. This analysis was restricted to the annual home
ranges of resident coyotes.
Resource use and selection.—We defined coyote use of
land-cover types as the observed proportion of locations
observed in a land-cover type, whereas selection of land-cover
types was defined as differences in observed use compared to
expected based on the null model that use would equal
availability if no selection occurred. Resource selection can
occur at multiple spatial scales (Johnson 1980); however, we
focused our analysis on 3rd-order selection (within the home
range). We did not assess 2nd-order selection (home range
within the study area) for 2 reasons. First, it was necessary to
restrict our trapping to areas with sufficient cover to avoid
conflicts with the public, which was either in private areas or,
more typically, public nature areas. Trapping in the most
heavily urbanized portions of the landscape was not possible.
Thus, our distribution of radiocollars was necessarily nonran-
dom among coyotes across the landscape. Second, the
territorial social system of coyotes likely prevented coyotes
from using some portions of the landscape at the home-range
scale, thereby altering our perception of the areas truly
available to individuals in ways that we could not measure
because we did not have all resident coyotes radiocollared
across the study area. However, we provide the composition of
home ranges to illustrate the range of use across the landscape
exhibited by coyotes and the availability of land-use types
within home ranges for 3rd-order selection. Use of landscape
types by coyotes was calculated for each animal annually and
by season by overlaying telemetry points on geographical
information system layers.
We assessed coyote selection of land-cover types using
Johnson’s (1980) rank method with the program PREFER 5.1
(Northern Prairie Science Center, United States Geological
Survey, Jamestown, North Dakota). This approach compares
rankings of use versus availability for resource components
(Johnson 1980), using the individual as the unit, and tests for a
significant deviation from an equal distribution with a multiple
comparison procedure (Waller and Duncan 1969). This
method also provides a test to identify which habitat
components differed in their rankings in the event of a
significant F-test. However, we were more interested in
general patterns of rankings among land-use types, and the
direction of their selection scores, rather than specific
statistical tests comparing 1 land-cover type to another. More
specifically, our attention was focused on land-cover types
1048 JOURNAL OF MAMMALOGY Vol. 90, No. 5

associated with people (Residential, Urban Grass, and Urban
Land) and whether their rankings varied with season or sex
class. In this paper, we use the term ‘‘selection score’’ to refer
to T-bar scores in Johnson (1980), which represent the mean
use ranking minus the mean availability ranking. In this case, a
large negative value would indicate selection for a land-use
category, whereas a large positive value results from a type of
patch with low use relative to availability and indicates
possible avoidance. All analyses required a minimum of 9
individuals, so for some groups (i.e., transients) it was not
possible to compare smaller subsets such as age or sex. As
with the home-range analysis, we selected the year with the
best data set for resident adults monitored in multiple years
and excluded the other years for those individuals to reduce
the possible effects of nonindependence. However, individuals
that changed status (i.e., age or social status) or moved to new
locations between years were retained in the analysis despite
appearing more than once.
We assessed selection of land-cover types within annual
home ranges separately for resident animals and transients. To
determine if selection of land-cover types varied by season, we
calculated mean selection scores for each year and used a 2-
way ANOVA (season and land-use type as main effects) on
these ratios with a focus on the interaction term between
season and land-cover type. Because of small sample sizes, we
did not assess seasonal variation in selection of land-cover
types for transients. We used a similar analytical approach to
compare land-cover selection by resident males and females,
in which selection scores were determined for each year and
compared with a 2-way ANOVA (sex and land-cover type as
main effects) with a focus on the interaction term.
Because coyotes shift their activity toward nocturnal hours
in more urban areas and rest in cover during the day (Gehrt
and Riley, in press), it is possible that resource selection may
differ between activity periods, especially regarding land-
cover types associated with human activities. Therefore, we
determined patterns of land-cover selection between activity
periods for resident and transient coyotes, with the hypothesis
that coyote selection for Residential and Urban Lands may
increase during nocturnal hours when human activity declines.
Because coyotes with home ranges composed largely of
human development may be assumed to be attracted to
human-associated land-cover types (given limited habitat
patches), we further focused this analysis on coyotes with
home ranges with a mix of natural and urban land use, and
coyotes with highly urbanized home ranges (.50% combi-
nation of Residential and Urban Land types). Resident coyotes
with home ranges located largely within natural fragments
were censored from this analysis.
Conservation efforts are often focused on minimum levels
of natural habitat required to maintain home ranges in urban
landscapes. Therefore, we report the size of home ranges
relative to the cumulative composition of natural habitat
within the home range. We report the smallest fragment of
natural habitat in which a coyote associated with a pack
maintained a territory for at least a full year. Likewise, we
report the cumulative proportion of natural habitat within
annual home ranges with a particular focus on the minimal
amount used by resident coyotes.
Nuisance coyotes.—We identified marked coyotes as
nuisances if .1 human resident registered complaints
(typically phone calls to an animal control agency) concerning
an individual animal, or if the complaint of an individual
resident resulted in control action against a coyote. In essence,
the public identified a nuisance coyote rather than us.
Unfortunately, our sample size of nuisance coyotes was not
sufficient for analyses of resource selection.
RESULTS
We captured and radiocollared 181 coyotes (including 17
adult females, 41 subadult females, 28 female pups, 28 adult
males, 40 subadult males, and 27 male pups). We recorded
25,509 locations, yielding sufficient numbers of locations to
estimate 182 annual home ranges, including multiple estimates
for some individuals. We initially estimated 118 annual home
ranges for residents and 40 home ranges for solitary transients.
Because some individuals were monitored for multiple years,
we reduced the number of home ranges to 84 residents (22
adult females, 11 subadult females, 29 adult males, and 22
subadult males). Individual coyotes appear in the analyses
more than once if they graduated from one age group to
another, or if they shifted from transient to resident or vice
versa. Twenty-four annual home ranges were calculated for
pups, which are presented for reference but are not included in
analyses because we assumed they were still associated with
adults.
Home-range size.—Annual home-range estimates for resi-
dent coyotes (Table 1) were similar by sex (F 5 0.03, d.f. 5 1,
80, P 5 0.86) and age (F 5 0.61, d.f. 5 1, 80, P 5 0.44)
classes with no interaction (F 5 0.89, d.f. 5 1, 80, P 5 0.35).
Similarly, differences in annual home-range estimates for
transient coyotes among sex and age classes were not
significant (all P . 0.12; Table 1). However, mean (6 SE)
annual home ranges of transient coyotes (X
¯
5 26.80 6
2.95 km
2
) were larger (t 5 12.6, d.f. 5 122, P , 0.001) than
TABLE 1.—Annual home-range estimates (km
2
) for radiocollared
coyotes (Canis latrans) in the Chicago, Illinois, metropolitan area
during 2000–2006, estimated with 95% minimum convex polygon.
Home-range estimates are partitioned by age–sex class within social
categories, except that males and females are pooled in the
juvenile class.
Category Age–sex group nX
¯
SE
Resident Adult female 22 4.80 0.66
Subadult female 11 5.17 0.98
Adult male 29 5.46 0.58
Subadult male 22 4.32 0.68
Juvenile 24 2.53 3.08
Transient Adult female 9 18.92 4.82
Subadult female 14 34.67 5.91
Adult male 15 23.70 3.92
Subadult male 2 30.47 18.64
October 2009 GEHRT ET AL.—URBAN COYOTE LANDSCAPE USE 1049

those of resident coyotes (X
¯
5 4.95 6 0.34 km
2
), with home
ranges of transients ranging up to 98 km
2
. Home-range size
for resident coyotes did not vary among seasons (F 5 1.02, d.f.
5 2, 210, P 5 0.36), or between age (F 5 2.63, d.f. 5 2, 210,
P 5 0.11) and sex (F 5 2.17, d.f. 5 2, 210, P 5 0.14) classes
(Table 2).
Home-range composition and land use.—Home ranges of
resident coyotes were typically associated with natural
habitats, and in many cases these were home ranges that were
almost completely encompassed within large habitat frag-
ments (Fig. 2). In these cases, boundaries of coyote home
ranges followed the borders between parks and surrounding
development. However, some coyote home ranges varied
considerably in the composition of natural habitat, with
concomitant variability in the composition of Urban Land
and Residential land use (Fig. 3). Unlike home ranges of
residents, home ranges of transients were not composed
exclusively within natural fragments, although Natural land
cover still dominated the other land-cover categories (Fig. 3).
Home-range size of residents was positively (r 5 0.38, n 5
84, P , 0.001) related to the amount of human-related
development within the home range (Fig. 4). However, home-
range size also varied substantially among coyotes with home
ranges composed almost exclusively of Natural or Undevel-
oped areas. Thirty-seven annual home ranges were located
nearly exclusively (.95%) within single fragments of Natural
land use, and ranged in size from 0.92 km
2
to 11.1 km
2
. The
smallest contiguous Natural fragment to exclusively occupied
a coyote pack was 247 ha. Larger fragments of Natural habitat
typically had multiple pack territories within them.
In contrast to those home ranges associated with Natural
land-use patches, there also were 24 annual home ranges of
residents that were composed of little (,10%) Natural land
use. Eight percent of annual home ranges had no (0%)
measurable patches of Natural land use within them.
Third-order selection.—Resident and transient coyotes
showed similar patterns of land use within annual home
ranges (Fig. 5), with both classes of coyotes using Natural
areas heavily. Similarly, selection of land-use types within
home ranges was nearly identical between coyote classes
(Fig. 5). In both cases, coyotes selected Undeveloped and
Other categories most (the 2 highest-ranked categories), and
avoided Urban Grass, Residential, and Urban Land as the
lowest-ranked categories (residents: F 5 10.49, d.f. 5 7, 89, P
, 0.001; transients: F 5 8.18, d.f. 5 7, 19, P , 0.001).
Coyotes avoided urbanized areas either by restricting their
movements to boundaries of Natural habitat fragments or by
focusing their activities within series of smaller patches of
undeveloped areas within home ranges (Fig. 6).
Selection by season.—Resident coyotes also exhibited
selection of land-use types within seasonal home ranges (all
P , 0.001), and there was no difference in the pattern of
selection among seasons (2-way ANOVA interaction term
season
3 land-use: F 5 1.21, d.f. 5 14, 96, P 5 0.28).
Undeveloped, Other, and Water were the most-selected
categories in each season, and the 3 land-use categories
associated with humans, Urban Grass, Urban Land, and
Residential, were avoided (Table 3).
Selection by sex.—Male (n 5 56) and female (n 5 40)
residents exhibited selection of land-use types within annual
home ranges (males: F 5 5.68, d.f. 5 7, 50, P , 0.0001;
females: F 5 6.37, d.f. 5 7, 33, P , 0.0001), and their
patterns of selection were similar across years (2-way
ANOVA interaction term sex
3 land-use class: F 5 0.36,
d.f. 5 7, 63, P 5 0.92). In both cases, Undeveloped and Other
were the highest-ranked categories (used greater than
expected), and Residential, Urban Grass, and Urban Land
were the 3 lowest-ranked categories, in each case with
selection scores indicating avoidance.
Selection by activity period.—Rankings of land-use selec-
tion by coyotes remained generally consistent between diurnal
and nocturnal periods (Fig. 7). Transient coyotes (n 5 14)
selectively used land-use categories during both periods
(diurnal: F 5 6.12, d.f. 5 7, 7, P 5 0.014; nocturnal: F 5
4.51, d.f. 5 7, 7, P 5 0.032). For both activity periods, the 3
top-ranked land-use types were Water, Undeveloped, and
Other. These were relatively strongly selected for during the
diurnal periods, with Water the most selected (Fig. 7). For
both periods, Residential was strongly avoided, along with
Urban Grass during diurnal periods, and Urban Land during
nocturnal periods.
We partitioned resident coyotes into those with home
ranges composed of .50% urban matrix (urban home ranges)
and those with highly urbanized home ranges (high-urban
home ranges). Coyotes with urban home ranges (n 5 41) had
significant selection during diurnal (F 5 7.64, d.f. 5 7, 34, P
, 0.001) and nocturnal (F 5 14.12, d.f. 5 7, 34, P , 0.001)
periods, and the ranking order was identical for both periods.
Undeveloped, Other, and Water were the 3 top-ranked
categories, in each case reflecting positive selection, and
Urban Grass, Residential, and Urban Land the lowest-ranked
categories, in each case indicating avoidance (Fig. 7).
Coyotes with high-urban home ranges (n 5 9) did not
exhibit significant selection by activity period (diurnal: F 5
7.86, d.f. 5 7, 2, P 5 0.12; nocturnal: F 5 3.83, d.f. 5 7, 2, P
TABLE 2.—Seasonal home-range estimates (km
2
) for resident,
radiocollared coyotes (Canis latrans) in the Chicago, Illinois,
metropolitan area, 2000–2006. Home ranges were estimated with
95% minimum convex polygon.
Season Age Sex nX
¯
SE
Breeding Adult Female 22 4.40 0.44
Male 31 3.89 0.39
Subadult Female 4 2.72 0.94
Male 10 3.79 0.74
Pup-rearing Adult Female 27 4.17 0.60
Male 36 4.00 0.39
Subadult Female 7 5.70 2.01
Male 12 2.80 0.58
Dispersal Adult Female 20 4.47 0.54
Male 27 4.73 0.40
Subadult Female 5 6.04 1.43
Male 10 3.00 0.50
1050 JOURNAL OF MAMMALOGY Vol. 90, No. 5

5 0.22). However, the lack of significance may be the result
of small sample size, because there were relatively strong
selection scores, with Undeveloped and Water as the 2 top-
ranked categories in both periods and Urban Grass, Residen-
tial, and Urban Land the lowest-ranked categories for both
periods (Fig. 7).
Nuisance coyotes.—We identified 7 radiocollared coyotes
that generated complaints from the public or were killed
during control efforts (Table 4), which comprised 4% of the
radiocollared sample. Nuisance coyotes were represented by a
range of sex–age classes. Four coyotes had dispersed from
their original territories, with 1 becoming a resident and the
rest transients at the time of removal. Overall, 3 coyotes were
residents and 4 were transients at the time of conflict
(Table 4). Two coyotes (adult males 2 and 3) were nuisances
immediately upon being radiocollared. Only coyote 3 was
known to possibly attack domestic animals and also was the
only suspected alpha male as a nuisance (based on visual
observations with a reproductive female); coyote 3 was in
good health, rested during the day in a large patch of natural
habitat, and moved into developed areas at night. Four coyotes
were in poor health at the time of conflicts, including 3
afflicted with mange (they were not known to create a conflict
before mange infections). In all cases they were observed near
houses during the day, which they apparently used for food or
shelter (2 were known to attempt to use dens under decks).
One of these (coyote 76) was monitored for nearly 4 years
without incident, until she developed a severe mange
infection. Two coyotes in good health at the time of conflict
were an adult (coyote 78) that was shot while located near
runways at O’Hare International Airport and a juvenile
(coyote 154) that was increasing his use of developed areas
in response to feeding by people before a collision with a
vehicle (other members of the pack continued to avoid
developed areas).
DISCUSSION
Home-range size can be an important indicator of resource
distribution and abundance (Gittleman and Harvey 1982), and
also may correlate with population density. Thus, comparisons
of home-range size between urban and rural landscapes can
provide important insights into the ecology of carnivores in
urban areas. At the landscape level, small home ranges can be
an indicator of high population densities (Andelt 1985;
Fedriani et al. 2001) in either urban or rural areas.
Are urban home ranges smaller than rural home ranges?—
Our estimates of annual and seasonal home-range sizes for
resident coyotes are smaller than winter (median 27.0 km
2
)or
summer (median 16.8 km
2
) home ranges reported for residents
in the agriculturally dominated landscape of rural Illinois
(Gosselink et al. 2003). Similarly, Atwood et al. (2004)
observed smaller home ranges for coyotes in suburban areas.
Likewise, there is a trend for home ranges of red foxes, a
FIG.2.—Distribution of annual home ranges of resident (yellow lines) and transient (white lines) coyotes (Canis latrans) in the Chicago,
Illinois, metropolitan area during 2004.
October 2009 GEHRT ET AL.—URBAN COYOTE LANDSCAPE USE 1051

synanthropic species, to decrease with urbanization (Cavillini
1996; Goszczyn
´
ski 2002).
Although comparisons across studies suggest a general
trend for smaller home ranges to occur in urban landscapes, at
the local scale habitat fragmentation resulting from urbaniza-
tion may cause coyotes to increase home-range size to meet
daily needs (Riley et al. 2003). The positive relationship
between urban land use and home-range size in our study is
consistent with this argument, but contradicts previous studies.
Within the Los Angeles, California, area, Tigas et al. (2002)
did not observe an increase in home-range size with increased
urban fragmentation, but their study had a small sample size,
and Riley et al. (2003) found a positive, but nonsignificant,
relationship between home-range size and urban development
from the same area.
Although we documented a positive relationship between
urban composition in the home range and home-range size, we
also documented considerable variation in home-range size
regardless of the composition of home ranges. Likewise,
coyotes in Tucson, Arizona (Grinder and Krausman 2001),
and urban red foxes in Great Britain (Doncaster and
Macdonald 1991; Soulsbury et al. 2007) also exhibit
considerable variation in home-range size. It appeared that
the size and shape of resident home ranges in our Chicago-
area study were simultaneously affected by the juxtaposition
of resources, fragmentation, and human activities. For
example, the home-range boundaries of many resident coyotes
conformed to the boundaries of natural areas, probably
reflecting the dramatic small-scale gradients typical of
urbanization. However, other resident coyotes maintained
home ranges that spanned such gradients, and individual
variation among coyotes in response to development and
human activities was apparent.
The upper range of home-range sizes of transients in our
study was similar to that from urban and rural studies,
although our mean size was relatively smaller than those
reported for other areas (Cape Cod, Massachusetts—115 km
2
[Way et al. 2002], and Tucson—105 km
2
[Grinder and
Krausman 2001]). Our relatively smaller mean may reflect
the considerable individual variation in the way coyotes leave
territories and their movements across the heavily developed
landscape in our study area. Also, our long-term monitoring of
individuals allowed us to identify when individuals separated
from their original territories and entered a transient phase,
regardless of the size of the area of their movements. In any
event, it is clear transients are capable of maintaining large
home ranges in highly urbanized landscapes.
Does home-range size vary with sex, age, or season?— Our
findings that home ranges were similar in size among different
sex and age groups and seasons are generally consistent with
previous studies. Urban and rural studies of coyotes have been
mixed when assessing sex differences in home-range size (see
Bekoff and Gese 2003; Laundre` and Keller 1984). For
example, in the same Los Angeles study, Tigas et al. (2002)
FIG.3.—Box plots of composition of annual home ranges by land-
use category for resident and transient coyotes (Canis latrans) in the
Chicago, Illinois, metropolitan area, 2000–2006. Horizontal lines
represents the mean, the box represents the standard deviation, 95%
confidence intervals, and points outside confidence intervals.
FIG.4.—Relationship between annual home-range size (95%
minimum convex polygon) and proportion of the home range
composed of urban land use (combined Residential, Urban Grass,
and Urban Land) for radiocollared coyotes (Canis latrans) in the
Chicago, Illinois, metropolitan area.
1052 JOURNAL OF MAMMALOGY Vol. 90, No. 5

reported females with larger home ranges than males, whereas
Riley et al. (2003) found that home ranges of males were
larger than those of females. As in our study, home-range size
was similar between sexes for coyotes in Tucson (Grinder and
Krausman 2001).
Few studies of urban coyotes have assessed seasonal
variation in home-range size, but coyotes in Tucson also
exhibited consistency in home-range size across seasons
(Grinder and Krausman 2001). Some studies from rural areas
have reported seasonal differences (Bekoff 1982; Gese et al.
1988), including in Illinois (Gosselink et al. 2003), whereas
others have not (Andelt 1985).
Are coyotes attracted to human-associated areas?— We
found no evidence that resident or transient coyotes, in
general, were attracted to human-associated areas within home
ranges, regardless of age, sex, season, or activity period,
despite considerable individual variation in home-range
compositions. Developed land-use types were consistently
the lowest-ranked habitats and selection scores indicated
avoidance. Even coyotes with home ranges primarily com-
posed of developed areas maintained a consistent avoidance of
areas associated with humans. Previous studies of urban
coyotes have either documented use of residential areas in
proportion to availability (Gibeau 1998; Grinder and Kraus-
man 2001; Way et al. 2004), or avoidance of residential or
other developed land-use types (Bogan 2004; Quinn 1997).
Foxes in urban areas have been observed to restrict their use
of residential areas to nocturnal hours (Harrison 1997;
Saunders et al. 1997), and some have suggested that coyotes
do the same (Quinn 1997). In contrast, we did not observe a
temporal shift in selection of developed areas in our study,
even for coyotes with highly urbanized home ranges.
However, avoidance does not imply that coyotes did not use
developed areas. We were able to visually observe resident
and transient coyotes with highly urbanized home ranges
passing through residential and commercial areas at night to
reach isolated habitat patches with no natural corridors, but
they did so quickly and these movements did not result in high
levels of use in our monitoring data. Indeed, it is likely these
coyotes would be removed by control efforts if they did not
move quickly and covertly through these areas during the
night. In contrast to most coyotes, 1 of the nuisance coyotes in
our study resided in an urban natural area during the day and
foraged in an adjacent residential area at night.
What kinds of land cover are selected by coyotes?— We did
not assess 2nd-order selection in our study, but it was notable
that many coyote home ranges largely comprised protected
areas in the form of Natural land use or areas protected from
development, which also was typical of coyotes in other urban
areas (Grinder and Krausman 2001; Riley et al. 2003).
However, as with coyotes in Tucson (Grinder and Krausman
2001), there was considerable variation in home-range
composition among coyotes, ranging from home ranges that
were composed exclusively of single large patches of Natural
land use to home ranges that did not include any patches of
Natural land use.
Although Natural land cover was consistently in the middle
of selection rankings with use by coyotes near that of
availability, this was undoubtedly affected by apparent 2nd-
order selection by coyotes that resulted in many home ranges
with large proportions of this land-use type. The importance of
natural habitat patches was reflected by the high level of use
exhibited by most coyotes and diets dominated by food items
associated with natural areas (Morey et al. 2007). Although
natural patches appear to serve as cover and provide food
items for coyotes, it is important to note that the vast majority
of properties associated with natural habitats in our study area
were public, with resulting heavy use by humans. For
example, the Ned Brown Forest Preserve in our study area
is typical of most Cook County forest preserves, in which the
approximately 15-km
2
area receives between 1 and 3 million
human visitors annually (Gehrt 2004), and is used for multiple
purposes including hiking, biking, picnicking, and as exercise
areas for pets. Thus, natural areas that might represent refuge
from people in other systems provide only limited respite from
FIG.5.—Use and selection of land-use types by radiocollared
coyotes (Canis latrans) in the Chicago, Illinois, metropolitan area,
2000–2006. Selection score represents mean difference in ranks
between use and availability within the home range (Johnson’s [1980]
T-bar), with negative scores representing selection for, and positive
selection scores representing avoidance.
October 2009 GEHRT ET AL.—URBAN COYOTE LANDSCAPE USE 1053

people in the Chicago metropolitan area. Indeed, it could be
argued that some of the public natural areas have human
activity on as high a level as traditional developed areas,
especially during warmer months. Coyotes in these areas are
still exposed to people and pets, and vice versa, on a consistent
basis, but nevertheless have more escape cover than in
developed areas. Even within large urban natural parks,
coyotes typically avoid areas or trails with high human activity
(George and Crooks 2006).
The land-cover types with consistently high selection
rankings and significant selection scores within home ranges
were Open, Undeveloped, and Water. Undeveloped, in
particular, was used for cover and foraging habitat, and was
particularly important for those coyotes with little or no
natural habitat in their home ranges. This land-cover type is
most easily characterized as small patches that are typically
too small to develop into buildings, and often serve as buffers
between developments or roads. Open areas also serve
multiple purposes for coyotes, especially when golf courses
or cemeteries maintain small patches of habitat that provide
cover. The Water category was most typically represented by
small water-retention ponds near developments. When coyotes
used these types of areas, they usually had emergent
vegetation such as cattails, rushes, or Phragmites that provided
cover for coyotes, especially during the day. We also observed
coyotes frequently using these areas during winter, possibly
for insulation from wind and low ambient temperatures.
Unfortunately, we are unsure as to their value for prey species.
FIG.6.—Variability of landscape use among urban coyotes (Canis latrans), as illustrated by patterns of use within annual home-range
boundaries for 3 resident coyotes during 2004 in the Chicago, Illinois, metropolitan area. Each color represents the locations and 95% minimum
convex polygon of a resident coyote from 3 territories. Each coyote exhibits avoidance of developed areas, despite considerable differences in
territory composition.
TABLE 3.—Seasonal land-use selection within home ranges by
radiocollared coyotes (Canis latrans) in the Chicago, Illinois,
metropolitan area (2000–2006). Land-use categories are ranked in
order of selection, with 1 representing the most-used category relative
to availability, and 8 the least-used relative to availability. Selection
scores represent the mean difference in ranks between use and
availability among land-use types (Johnson 1980). A negative
selection score reflects preferred categories, whereas positive values
reflect avoidance.
Land use
Breeding Pup-rearing Dispersal
Ranking Score Ranking Score Ranking Score
Undeveloped 1 21.07 1 21.00 1 21.11
Other 2 20.94 2 20.99 2 21.10
Water 3 20.50 3 20.64 3 20.44
Agriculture 4 20.19 4 20.44 4 20.40
Natural 5 20.14 5 20.13 5 20.17
Urban Grass 6 0.50 6 0.50 7 0.32
Residential 7 0.83 7 0.81 6 0.16
Urban Land 8 1.51 8 1.89 8 1.85
1054 JOURNAL OF MAMMALOGY Vol. 90, No. 5

However, we frequently observed coyotes hunting small
mammals in Undeveloped patches, especially easements along
major roadways.
The patterns of landscape use and selection we observed for
radiocollared coyotes were consistent with diet studies in our
study area, in that food items associated with natural areas,
rather than anthropogenic foods, dominated the diet across the
landscape (Morey et al. 2007). Further, coyotes did not use
developed habitat associated with human food within large
natural fragments. The high levels of human use within the
natural areas in our study area typically resulted in readily
available refuse near garbage cans and dumpsters, which were
used heavily by raccoons (Bozek et al. 2007; Prange et al.
2004). Yet, coyotes residing in these areas did not exhibit an
attraction to these areas within their territories (Gehrt 2004)
and consequently human-related foods rarely occurred in their
diets (Morey et al. 2007). Thus, the relatively heavy use of
natural areas by coyotes was not a result of the availability of
rich clumps of anthropogenic foods that occur in those areas.
Given that some studies using track stations have indicated
that coyotes have a threshold of fragmentation beyond which
they are absent (Crooks 2002; Randa and Yunger 2006), we
were surprised to observe coyotes maintaining home ranges in
areas with limited or no natural habitat. In each case these
were stable home ranges that were maintained for 1 or more
years, and in each case these were apparently associated with
packs because other coyotes were observed with radiocollared
individuals. Some of these packs were located in downtown or
industrial areas, including some of the most heavily developed
portions of our study area, in addition to occasional transients.
It is important to note that we were unable to trap for coyotes
in completely urbanized areas, and could only document that
use once radiocollared coyotes moved into those areas.
However, we frequently observed unmarked coyotes traveling
with radiocollared individuals and litter-rearing in these areas.
Indeed, we were aware of coyotes residing in portions of the
inner core of the city of Chicago. To some degree our results
contradict predictions, derived from track stations, that
coyotes in the Chicago area will decline in occurrence as
urbanization continues to convert rural lands to urban
development (Randa and Yunger 2006). Attempts to deter-
mine coyote presence with track stations and artificial
substrates may underestimate the presence of coyotes (Harris
and Knowlton 2001).
Nuisance coyotes.—We are unaware of previous descrip-
tions of the frequency and characteristics of nuisance coyotes
in an urban area. Coyotes of various sex and age classes
became nuisances, and in nearly all cases either disease or
feeding by residents was involved. The relatively low
FIG.7.—Selection of land-use classes within home ranges by
coyotes (Canis latrans) during diurnal and nocturnal hours: A)
transients (n 5 14), B) residents with mixed land-use types (n 5 41),
and C) residents with highly urbanized home ranges (n 5 9).
Selection scores represent differences between mean rankings of use
and availability (Johnson 1980). Negative scores indicate a higher use
ranking than available ranking for a land-use type, and vice versa for
positive scores. Coyotes were monitored during 2000–2006 in the
Chicago, Illinois, metropolitan area.
TABLE 4.—Characteristics of nuisance coyotes (Canis latrans) in the Chicago, Illinois, region, 2000–2006. Monitoring period refers to the
length of time between initial capture and last location or terminal capture. Status includes individuals that dispersed from group territories and
were subsequently transients (Disp. Tran.) or residents (Disp. Res.) at the time of capture.
Coyote no. Sex Age Monitoring period Status Condition Fate
2 Male Adult 25 March 2000–24 April 2001 Transient Fair–poor Disappeared
3 Male Adult 30 March 2000–31 May 2000 Resident Good Road-killed
76 Female Adult 25 October 2002–8 October 2006 Disp. Res. Poor, mange Euthanized
78 Male Adult 30 October 2002–10 January 2005 Disp. Tran. Good Shot at airport
93 Female Adult 3 April 2003–16 January 2004 Disp. Tran. Poor, mange Euthanized
154 Male Pup 12 August 2004–20 October 2004 Resident Good Road-killed
156 Male Pup 24 August 2004–28 December 2004 Disp. Tran. Poor, mange Shot by homeowner
October 2009 GEHRT ET AL.—URBAN COYOTE LANDSCAPE USE 1055

proportion of radiocollared coyotes that subsequently became
nuisances is consistent with the general pattern of avoidance
of human-related areas. Indeed, if those with health issues and
encroaching on airports are excluded, the proportion of
radiomarked coyotes that became nuisances was low (2 of
181) during our study. Only 1 of the nuisance coyotes, a
presumed alpha male, was reported attacking and killing
domestic animals.
Conclusion.—Our results can be summarized into 3 primary
observations. First, in a landscape dominated by human
development, natural habitat was used heavily by coyotes.
Second, despite the importance of natural habitat for coyotes,
some individuals are capable of maintaining territories in
portions of the landscape with minimal or no natural areas and
elevated human activity. Third, coyotes consistently demon-
strated avoidance of areas associated with humans, regardless
of their sex, social status (resident or transient), the activity
period, or the amount of urban development within their home
ranges. Our interpretation of coyotes avoiding human-related
areas also is supported by a shift in activity to nocturnal
periods and a lack of human-related foods in the diet.
It was clear that coyotes were using a variety of strategies to
exploit the landscape while avoiding people, with some
limiting their use exclusively to natural patches, and others
with a mixture of land-use types. However, coyotes were
consistent in apparently avoiding human activities despite
home ranges located in areas with nearly complete develop-
ment, which created a paradox of use and avoidance of
developed land-use types. Future research should further
explore the limitations of urban landscapes for coyotes,
including the influence on survival and reproductive rates in
different parts of the urban landscape, but our results
demonstrate that coyotes represent a medium-to-large carni-
vore capable of exploiting areas of intense development while
largely managing to avoid people and conflicts.
ACKNOWLEDGMENTS
Funding was provided by Cook County Animal and Rabies
Control, the Max McGraw Wildlife Foundation, and the Forest
Preserve District of Cook County. We thank C. Rizzo, M. Neri, J.
Brown, P. Morey, S. Prange, and many additional technicians and
graduate students that assisted with fieldwork. R. Erickson assisted
with trapping and provided snares. S. Riley provided comments on an
earlier draft, and the manuscript further benefited from comments
from P. Krausman and 1 anonymous reviewer. We especially
recognize the support of Dr. Dan Parmer (deceased).
LITERATURE CITED
ANDELT, W. F. 1985. Behavioral ecology of coyotes in south Texas.
Wildlife Monographs 94:1–45.
A
TKINSON, K. T., AND D. M. SHACKLETON. 1991. Coyote, Canis latrans,
ecology in a rural–urban environment. Canadian Field-Naturalist
105:49–54.
A
TWOOD, T. C., H. P. WEEKS, AND T. M. GEHRING. 2004. Spatial
ecology of coyotes along a suburban-to-rural gradient. Journal of
Wildlife Management 68:1000–1009.
B
EKOFF, M. 1982. Coyote. Pp. 447–459 in Wild mammals of North
America (J. A. Chapman and G. A. Feldhamer, eds.). Johns
Hopkins University Press, Baltimore, Maryland.
B
EKOFF, M., AND E. M. GESE. 2003. Coyote (Canis latrans). Pp. 467–
481 in Wild mammals of North America (G. A. Feldhamer, J. A.
Chapman, and B. C. Thompson, eds.). 2nd ed. Johns Hopkins
University Press, Baltimore, Maryland.
B
OGAN, D. A. 2004. Eastern coyote home range, habitat selection and
survival in the Albany Pine Bush landscape. M.S. thesis, State
University of New York, Albany.
B
OUNDS, D. L., AND W. W. SHAW. 1994. Managing coyotes in U.S.
national parks: human–coyote interactions. Natural Areas Journal
14:280–284.
B
OZEK, C. K., S. PRANGE, AND S. D. GEHRT. 2007. The influence of
anthropogenic resources on multi-scale habitat selection by
raccoons. Urban Ecosystems 10:413–425.
C
ARBYN, L. N. 1989. Coyote attacks on children in western North
America. Wildlife Society Bulletin 17:444–446.
C
ARDILLO, M., A. PURVIS,W.SECHREST,J.L.GITTLEMAN,J.BIELBY,
AND G. M. MACe. 2004. Human population density and extinction
risk in the world’s carnivores. PLoS Biology 2:909–914.
C
AVALLINI, P. 1996. Variation in the social system of the red fox.
Ethology Ecology & Evolution 8:323–342.
C
ROOKS, K. R. 2002. Relative sensitivities of mammalian carnivores
to habitat fragmentation. Conservation Biology 16:488–502.
C
ROOKS, K. R., AND M. E. SOULE
´
. 1999. Mesopredator release and
avifaunal extinctions in a fragmented system. Nature 400:
563–566.
C
YPHER, B. L. In press. Kit foxes (Vulpes macrotis). In Urban
carnivores: ecology, conflict, and conservation (S. D. Gehrt, S. P.
D. Riley, and B. L. Cypher, eds.). Johns Hopkins University Press,
Baltimore, Maryland.
D
ONCASTER, C. P., AND D. W. MACDONALD. 1991. Drifting territoriality
in the red fox Vulpes vulpes. Journal of Animal Ecology 60:423–
439.
F
EDRIANI, J. M., T. K. FULLER, AND R. M. SAUVAJOT. 2001. Does
availability of anthropogenic food enhance densities of omnivorous
mammals? An example with coyotes in southern California.
Ecography 24:325–331.
G
ANNON, W. L., R. S. SIKES, AND W. L. GANNON,R.S.SIKES, AND THE
ANIMAL CARE AND USE COMMITTEE OF THE AMERICAN SOCIETY OF
MAMMALOGISTS. 2007. Guidelines of the American Society of
Mammalogists for the use of wild mammals in research. Journal of
Mammalogy 88:809–823.
G
EHRT, S. D. 2004. Ecology and management of striped skunks,
raccoons, and coyotes in urban landscapes. Pp. 81–104 in Predators
and people: from conflict to conservation (N. Fascione, A. Delach,
and M. Smith, eds.). Island Press, Washington, D.C.
G
EHRT, S. D., AND J. E. CHELSVIG. 2003. Bat activity in an urban
landscape: patterns at the landscape and microhabitat scale.
Ecological Applications 13:939–950.
G
EHRT, S. D., AND S. P. D. RILEY. In press. Coyotes (Canis latrans). In
Urban carnivores: ecology, conflict, and conservation (S. D. Gehrt,
S. P. D. Riley, and B. L. Cypher, eds.). Johns Hopkins University
Press, Baltimore, Maryland.
G
EORGE, S. L., AND K. R. CROOKS. 2006. Recreation and large
mammal activity in an urban nature reserve. Biological Conser-
vation 133:107–117.
G
ESE, E. M., O. J. RONGSTAD, AND W. R. MYTTON. 1988. Home range
and habitat use of coyotes in southeastern Colorado. Journal of
Wildlife Management 52:640–646.
1056 JOURNAL OF MAMMALOGY Vol. 90, No. 5

GIBEAU, M. L. 1998. Use of urban habitats by coyotes in the vicinity
of Banff Alberta. Urban Ecosystems 2:129–139.
G
ITTLEMAN, J. L., AND P. H. HARVEY 1982. Carnivore home range size,
metabolic needs and ecology. Behavioral Ecology and Sociobiol-
ogy 10:57–63.
G
OMPPER, M. E. 2002. Top carnivores in the suburbs? Ecological and
conservation issues raised by colonization of northeastern North
America by coyotes. BioScience 52:185–190.
G
OSSELINK, T. E., T. R. VAN DEELEN,R.E.WARNER, AND M. G.
J
OSELYN. 2003. Temporal habitat partitioning and spatial use of
coyotes and red foxes in east central Illinois. Journal of Wildlife
Management 67:90–103.
G
OSZCZYN
´
SKI, J. 2002. Home ranges in red fox: territoriality diminishes
with increasing area. Acta Theriologica Supplement 1:103–114.
G
RINDER, M., AND P. R. KRAUSMAN. 2001. Home range, habitat use,
and nocturnal activity of coyotes in an urban environment. Journal
of Wildlife Management 65:887–898.
H
ARRIS, C. E., AND F. F. KNOWLTON. 2001. Differential responses of
coyotes to novel stimuli in familiar and unfamiliar settings.
Canadian Journal of Zoology 79:2005–2013.
H
ARRISON, R. L. 1997. A comparison of gray fox ecology between
residential and undeveloped rural landscapes. Journal of Wildlife
Management 61:112–122.
H
OOGE, P. N., AND B. EICHENLAUB. 1997. Animal Movement Extension
to ArcView. Version 1.1. United States Geological Survey, Alaska
Biological Science Center, Anchorage.
J
OHNSON, D. H. 1980. The comparison of usage and availability
measurements for evaluation of resource preference. Ecology
61:65–71.
J
OHNSTON, R. F. 2001. Synanthropic birds of North America. Pp. 49–
67 in Avian ecology and conservation in an urbanizing world (J. M.
Marzluff, R. Bowman, and R. Donnelly, eds.). Kluwer, Boston,
Massachusetts.
K
AMLER, J. F., AND P. S. GIPSON. 2000. Space and habitat use by
resident and transient coyotes. Canadian Journal of Zoology
78:2106–2111.
L
ALIBERTE, A. S., AND W. J. RIPPLE. 2004. Range contractions of North
American carnivores and ungulates. BioScience 54:123–138.
L
AUNDRE
`
, J. W., AND B. L. KELLER. 1984. Home range size of coyotes:
a critical review. Journal of Wildlife Management 48:127–139.
L
INNELL, J. D. C., J. E. SWENSON, AND R. ANDERSON. 2001. Predators
and people: conservation of large carnivores is possible at high
human densities if management policy is favourable. Animal
Conservation 4:345–349.
M
OREY, P. S. 2004. Landscape use and diet of coyotes, Canis latrans,
in the Chicago metropolitan area. M.S. thesis, Utah State
University, Logan.
M
OREY, P. S., E. M. GESE, AND S. GEHRT. 2007. Spatial and temporal
variation in the diet of coyotes in the Chicago metropolitan area.
American Midland Naturalist 158:147–161.
N
EWMAN, T. J., P. J. BAKER,E.SIMCOCK,G.SAUNDERS,P.C.L.WHITE,
AND S. HARRIS. 2003. Changes in red fox habitat preference and rest
site fidelity following a disease-induced population decline. Acta
Theriologica 48:79–91.
P
RANGE, S., S. D. GEHRT, AND E. P. WIGGERS. 2004. Influences of
anthropogenic resources on raccoon (Procyon lotor)move-
ments and spatial distribution. Journal of Mammalogy 85:483–
490.
Q
UINN, T. 1997. Coyote (Canis latrans) habitat selection in urban
areas of western Washington via analysis of routine movements.
Northwest Science 71:289–297.
R
ANDA, L. A., AND J. A. YUNGER. 2006. Carnivore occurrence along an
urban–rural gradient: a landscape-level analysis. Journal of
Mammalogy 87:1154–1164.
R
ILEY, S. P. D., ET AL. 2003. Effects of urbanization and habitat
fragmentation on bobcats and coyotes in southern California.
Conservation Biology 17:566–576.
S
AUNDERS, B., P. C. L. WHITE, AND S. HARRIS. 1997. Habitat utilization
by urban foxes (Vulpes vulpes) and the implications for rabies
control. Mammalia 61:497–510.
S
OULSBURY, C. D., G. IOSSA,P.J.BAKER,N.C.COLE,S.M.FUNK, AND
S. HARRIS. 2007. The impact of sarcoptic mange Sarcoptes scabiei
on the British fox Vulpes vulpes population. Mammal Review
37:278–296.
T
IGAS, L. A., D. H. VAN VUREN, AND R. M. SAUVAJOT. 2002.
Behavioral responses of bobcats and coyotes to habitat fragmen-
tation and corridors in an urban environment. Biological
Conservation 108:299–306.
W
ALLER, R. A., AND D. B. DUNCAN. 1969. A Bayes rule for the
symmetric multiple comparisons problem. Journal of the American
Statisticians Association 64:1484–1503.
W
ANG, Y., AND D. K. MOSKOVITS. 2001. Tracking fragmentation of
natural communities and changes in land cover: applications of
Landsat data for conservation in an urban landscape (Chicago
wilderness). Conservation Biology 15:835–843.
W
AY, J. G., I. M. ORTEGA, AND E. G. STRAUSS. 2002. Eastern coyote
home range, territoriality, and sociality on urbanized Cape Cod.
Northeast Wildlife 57:1–18.
W
AY, J. G., I. M. ORTEGA, AND E. G. STRAUSS. 2004. Movement and
activity patterns of eastern coyotes in a coastal, suburban
environment. Northeastern Naturalist 11:237–254.
W
OODROFFE, R. 2000. Predators and people: using human densities to
interpret declines of large carnivores. Animal Conservation 3:
165–173.
Submitted 29 August 2008. Accepted 16 April 2009.
Associate Editor was John A. Yunger.
October 2009 GEHRT ET AL.—URBAN COYOTE LANDSCAPE USE 1057