Research Article

Spatial Ecology of Bobcats and Gray Foxes in Urban

and Rural Zones of a National Park
SETH P. D. RILEY,1,2 Department of Environmental Science and Policy, University of California, Davis, CA 95616, USA

Abstract
Urbanization threatens the persistence of many wildlife populations, particularly those of wide-ranging and low-density species
such as mammalian carnivores. Effective conservation of carnivore populations requires an understanding of the impacts of
adjacent urbanization on carnivores in reserves. I compared the spatial ecology of bobcats (Lynx rufus) and gray foxes (Urocyon
cinereoargenteus) between urban and rural zones of a national park in northern California, USA. In the urban zone, gray foxes used
the entire landscape from interior natural areas across the park edges and into the neighboring developed areas, although fox core
areas were always within the park. Bobcats never entered development, and radiocollared adult female bobcats maintained home
ranges in the interior of the park, far from the urban edge. Bobcats appeared to avoid crossing paved roads, while foxes crossed
roads regularly. For adult female bobcats, home ranges were smaller in the urban zone, and core areas were both smaller and
overlapped more. Home range size and overlap did not differ between zones for gray foxes. Bobcats seem to be more affected by
the proximity of urbanization than foxes, perhaps because of differences in diet and social structure. The more flexible use of the
landscape by foxes may give them access to increased resources and habitat, but also may expose them to more human-
associated risks. If female bobcats are more sensitive to urbanization, this sensitivity could affect the long-term viability of bobcat
populations in urban areas. Knowledge of how bobcats and gray foxes use the landscape in urban areas will allow more effective
conservation and improved coexistence with these widespread carnivores by helping to predict where and why conservation or
management issues may occur. (JOURNAL OF WILDLIFE MANAGEMENT 70(5):1425–1435; 2006)

Key words
bobcat, California, edge effects, gray fox, home range overlap, home range size, Lynx rufus, Marin County, roads,
urbanization, Urocyon cinereoargenteus.

Urbanization is a major cause of the loss and fragmentation
of wildlife habitat. To manage and conserve wildlife
populations, we must not only preserve habitat but also
understand the impact of adjacent development on remain-
ing natural areas. Many species are displaced by urban-
ization, but others survive and even thrive in proximity to
dense human populations. Although mammalian carnivores
are often the hardest species to conserve because of their low
densities, large home ranges, and specific dietary require-
ments (Noss et al. 1996, Gittleman et al. 2001), some
carnivores reach high densities in urban areas. This is
particularly true for more omnivorous species such as
raccoons (Procyon lotor; Riley et al. 1998, Prange et al.
2003), red foxes (Vulpes vulpes; Harris 1981), and striped
skunks (Mephitis mephitis; Rosatte et al. 1992). Even the
larger coyote (Canis latrans) coexists with humans in many
metropolitan areas (Gill and Bonnet 1973, Bounds and
Shaw 1997, Quinn 1997a, Fedriani et al. 2001), and
endangered San Joaquin kit foxes (Vulpes macrotis motica)
maintain better body condition in the city of Bakersfield
than outside of it (Cypher and Frost 1999). More strictly
carnivorous species, however, may be less able to persist in
urban areas. I examined the spatial ecology of 2 medium-
sized carnivores, the carnivorous bobcat (Lynx rufus) and the
more omnivorous gray fox (Urocyon cinereoargenteus), in
urban and rural zones of a national park to determine the
response of each species to adjacent urban development.
1
E-mail: seth_riley@nps.gov
2
Present address: Santa Monica Mountains National Rec-
reation Area, Thousand Oaks, CA 91360, USA
The bobcat may represent a carnivore that is intermediate
in its sensitivity to urbanization and fragmentation (Kam-
radt 1995, Crooks 2002). Although large carnivores such as
mountain lions (Puma concolor) and wolves (Canis lupus) may
tolerate some proximity to development (Fuller et al. 1992,
Beier 1995, Mech 1995), their extensive prey and spatial
requirements and their sensitivity to roads (Thiel 1985,
Mech et al. 1988, Mladenoff et al. 1995) and other
disturbance (Van Dyke et al. 1986) threaten their persis-
tence in urban areas. Bobcats may be able to coexist with
some development as long as a minimum amount of
functional natural habitat remains (see also Riley et al.
2003). Although some carnivores can take advantage of a
variety of food resources that may be more available in
developed areas, such as ornamental fruit, insects, garbage,
pet food, and even pets (e.g., McClure et al. 1995, Quinn
1997b, Fedriani et al. 2001), bobcats are strict carnivores and
generally eat only fresh meat (Ewer 1973, Anderson and
Lovallo 2003). Other species such as raccoons (Fritzell
1978, Gehrt and Fritzell 1998), red foxes (MacDonald
1981), and coyotes (Bowen 1981) also have more flexible
social systems than bobcats, in which the size and type of
social groups can vary depending on resources and
population density. Bobcats are generally solitary and
territorial, especially between adult females (Anderson and
Lovallo 2003; e.g., Bailey 1974 but see Zezulak 1998,
Nielsen and Woolf 2002).
My objectives were to determine the space use of bobcats
and gray foxes relative to human development and roads, to
compare this use between species, and to compare the spatial

Riley
Urban and Rural Bobcats and Gray Foxes 1425


Figure 1. Location of bobcat and gray fox study areas in urban and
rural zones of Golden Gate National Recreation Area, Marin County,
California, USA, 1992–1995.
ecology of both carnivores between the urban and rural
zones of the park. Because of their more restricted diet,
greater spatial requirements, and more restrictive social
system, I hypothesized that bobcats would be more affected
by urbanization than gray foxes. Specifically, I expected that
foxes would utilize developed areas more than bobcats. I
further expected that gray foxes would exhibit fewer
differences in home range size or intraspecific home range
overlap between the urban and rural zones of the park than
would bobcats, and that an increased utilization of develop-
ment by foxes would lead to them more readily cross roads
within and adjacent to the park.
Study Area
I conducted my study in Golden Gate National Recreation
Area (GGNRA) in Marin County, California, USA, a
30,000-ha national park in the San Francisco Bay Area that
received approximately 14 million visitors per year. Annual
grasslands, coastal sage scrub, riparian woodlands, and oak–
bay woodlands characterized coastal Marin County, with
occasional evergreen forests to the north.
I studied bobcats and foxes in 2 areas of GGNRA: the
urban zone in the southern part of the park, and the rural
zone to the north (Fig. 1). In the urban zone, the park was
adjacent to the cities of Sausalito, Marin City, and Mill
Valley. The rural zone, which began 15 km to the
northwest, was 7–17 km from any large areas of develop-
ment, although there were occasional dwellings and small
settlements within the park boundary.
1426
In the urban zone, Highway 101, a major 6–8-lane
freeway, was adjacent to the park (Fig. 1), and one paved
road into the park received regular use by visitors and
residents (Tennessee Valley Road). In the rural zone 2 roads
received sometimes heavy traffic from tourists and com-
muters (Fig. 1).
Methods
Live-Trapping and Handling
I captured animals in homemade box traps (bobcats and gray
foxes; Zezulak 1998) and Tomahawk live traps (gray foxes;
Tomahawk Live Trap Co., Tomahawk, Wisconsin). I
chemically immobilized both species with a 5:1 mixture of
ketamine hydrochloride and xylazine hydrochloride, which
was injected intramuscularly. I ear-tagged, measured, and
sexed animals, aged them as juvenile or adult based on tooth
eruption and wear and body size, and monitored vital rates
during anesthesia. I radiocollared animals of adult size
(Telonics Inc., Mesa, Arizona; model 315 radiocollar for
bobcats, model 225 radio with a nylon belting collar for gray
foxes). When the animal started to emerge from the effects
of the ketamine, I administered an intramuscular injection
of yohimbine hydrochloride to antagonize the xylazine.
When fully recovered, I released animals at the site of
capture. A protocol for all capture and handling procedures
was approved by the Animal Care and Use Committee at
the University of California at Davis (protocol # 5328; May
1992). For animals that died during the study, age was
determined by counting cementum annuli of canine teeth
(Matson Laboratories, Milltown, Montana).
Radiotracking
Animals in both the rural and urban zones were intensively
radiotracked for at least 15 months (Aug 1992–Mar 1994 in
the urban zone, Jan 1994–Mar 1995 in the rural zone),
during which period at least 2 day locations and 1 night
location were obtained each week for each animal. I also
conducted sequential radiotracking sessions each season
during which I tracked a group of 4–6 animals every hour
for the full 24-hour cycle, in 2 12-hour sessions. I did not
use the sequential locations to compute home ranges, since
they were not independent; however, the home ranges
computed from the independent locations encompassed all
of the sequential locations.
Locations were obtained by triangulation of 3 compass
bearings using an H-Adcock peak antenna. All the bearings
for any one location were taken within 15 minutes, and
when I could not obtain 3 reliable bearings within this
period, I used 2 bearings (approx. 20% of locations). I took
bearings from the same drainage as the radiocollared animal
to minimize the distance to the animal and the effects of
intervening topography. I determined receiver locations to
within 2–5 m using a Global Positioning System (GPS;
Pathfinder plus, Trimble Inc., Sunnyvale, California).
Animals were sometimes directly observed, and these visual
locations were determined with 2 bearings or 1 bearing and
the distance. I measured telemetry system accuracy using the
GPS locations of test collars. The mean distance between
The Journal of Wildlife Management
70(5)
Figure 2. Radio-locations and home ranges (95% adaptive kernels) of
bobcats relative to development in the urban zone of Golden Gate
National Recreation Area, Marin County, California, USA, 1992–1995.
triangulated locations and test collars was 76.6 m 6 SD ¼
53.3 m, range ¼ 17.7–287.1, n ¼ 37.
Home Range Size
I computed 100% and 95% minimum convex polygon
home ranges (MCPs; Hayne 1949) and 95% adaptive
kernel home ranges (AKs; Worton 1989) as representations
of the overall home range, and 50% adaptive kernel home
ranges as a representation of the core area of use (Kaufman
1962). I used 95% AKs for statistical analyses because
kernels are the most accurate estimators that incorporate
location density (Seaman and Powell 1996, Powell 2000).
Because kernel home ranges include areas outside the actual
locations of the animal, a number of home ranges included
open water of Bolinas lagoon in the rural zone and of the
Pacific Ocean in the urban zone. I removed these areas from
the home ranges using Arcview Geographic Information
System (GIS; ESRI, Redlands, California; Fig. 2). Often
ecologically important events such as the use of residential
areas were represented by a small number of locations, so I
used 100% MCPs to show these events graphically and to
reflect the use of outlying areas without exaggerating it (Fig.
3).
I compared home range sizes between the urban and rural
zones and between sexes with a 2-way analysis of variance,
using the program SYSTAT (SPSS, Inc., Chicago, Illinois).
Home ranges and core areas were log- or square-root (fox
core areas) transformed to meet the assumptions of
Riley
Urban and Rural Bobcats and Gray Foxes
Figure 3. Home ranges (100% minimum convex polygons) of gray
foxes relative to development in the urban zone of Golden Gate National
Recreation Area, Marin County, California, USA, 1992–1995. One
hundred percent minimum convex polygon home ranges are depicted
to show all foxes that used developed areas while not overestimating this
use, as can be the case with kernel home ranges (e.g., B001 in Fig. 2).
ANOVA. Because male bobcat home ranges are consistently
larger than those of females (Anderson and Lovallo 2003), I
also conducted contrasts between zones for males and
females separately.
Home Range Overlap
I computed the percentage of home range overlap for all
con-specific pairs whose ranges overlapped, and I classified
each percentage as female-on-female, male-on-male, male-
on-female, or female-on-male. For the same pairs whose
95% ranges overlapped, I calculated core area overlap. For
bobcats I tested for differences in overlap of 95% AKs
between the zones and between the sex combinations with
2-way analysis of variance, after a square-root trans-
formation. The 50% core area overlap data had a skewed
distribution because of the large number of zero values, and
it could not be effectively transformed, so I compared
overlap between zones within overlap type (e.g., female-on-
female between urban and rural zones) using Mann–
Whitney U tests.
For foxes, I also classified male-on-female and female-on-
male overlaps as belonging to a mated pair (Fritzell 1987,
Greenberg and Pelton 1994, Chamberlain and Leopold
2000) if an adult male and adult female were regularly
1427

Figure 4. Core areas (50% adaptive kernels) of gray foxes relative to
development in the urban zone of Golden Gate National Recreation
Area, Marin County, California, USA, 1992–1995.
located together and had home ranges and core areas that
overlapped by 50% or more. I compared 95% AK overlap
between the urban and rural zones, between sex combina-
tions, and between paired and unpaired animal pairs using a
3-way analysis of variance. Overlap data were square-root
transformed. Again, the core area overlaps could not be
transformed, so I tested for differences between zones with
Mann–Whitney U tests.
Use of Interior Park, Park Edge, and
Developed Areas
In the urban zone, to establish whether animals used all parts
of the park or crossed the park boundary into residential areas,
I determined the location of home ranges, core areas, and
individual radio-locations relative to development. First, I
outlined polygons of developed areas both within and outside
the park with Arcview using geographically rectified aerial
photographs of the park and surrounding areas. I included all
urban areas adjacent to the park and the 2 larger developed
areas within the park (2 southern polygons; Figs. 2–4). I then
determined whether each animal’s home range and core area
overlapped or abutted developed areas. Quantitatively, for
each animal I also measured the minimum distance between
the closest urban development and each animal’s nearest home
range boundary, and I calculated the average of the minimum
distance between each of its radio-locations and the nearest
development edge. I determined these distances using the
1428
Nearest Feature extension (Jenness 2000) for Arcview. These
data could not be effectively transformed, so I compared home
range and average radio-location distances to development
between sexes (within species) using Mann–Whitney U tests.
I made comparisons between sex or species groups using a
Kruskall-Wallis test and tested for pair-wise differences with
contrasts (Daniel 1990).
Home Range Use and Roads
Many studies that examine the effects of roads on wildlife
analyze the influence of road density on behavior and
ecology (e.g., Theil 1985, Mech et al. 1988, Lovallo and
Anderson 1996). In my study home ranges were too small
and road density too low for this kind of analysis. Instead, I
examined the home range placement and radio-locations of
carnivores relative to roads, specifically paved roads that
received regular vehicle use. I assumed that if an animal
inhabited an area near a road (i.e., its 100% MCP home
range overlapped or was within 500 m of the road) but
,5% of its locations were across that road, then the road
acted as a home range boundary, and the animal avoided
crossing it. I used a Fisher exact test to test for differences
between bobcats and foxes in whether roads represented
home range boundaries.
Results
Home Range Size
Bobcats.—On average, bobcat 95% AK home ranges
were 4.2 times (females) and 2.3 times (males) larger in the
rural zone, and male home ranges were 4.2 times (urban
zone) and 2.3 times (rural zone) larger than those of females
(Table 1; Zone: F1,15 ¼ 8.14, P ¼ 0.012; Sex: F1,15 ¼ 11.38,
P ¼ 0.004). Similarly, core areas were 4.7 times (females)
and 1.2 times (males) larger in the rural zone, and male core
areas were 6.8 times (urban zone) and 1.7 times (rural zone)
larger than female core areas (Table 1; Zone: F1,15 ¼ 5.40, P
¼ 0.035; Sex: F1,15 ¼ 17.45, P ¼ 0.001). The interaction
between zone and sex was not significant for home ranges
(F1,15 ¼ 0.527, P ¼ 0.479) and was marginally so for core
areas (F1,15 ¼ 3.26, P ¼ 0.091). However, despite these
larger average sizes for male bobcats in the rural zone, there
was significant variability (Table 1), and contrasts (total a ¼
0.05, Bonferroni correction) revealed no significant differ-
ence between the urban and rural zones for male bobcats for
either the 95% ranges (difference ¼ 0.685, critical difference
¼ 1.498, t ¼ 2.571, df ¼ 5) or the 50% core areas (difference
¼ 0.157, critical difference ¼ 1.233, t ¼ 2.571, df ¼ 5). In
contrast, for adult females, average home range size was
significantly larger in the rural zone for both 95% ranges
(total a ¼ 0.05, Bonferroni correction: difference ¼ 1.152,
critical difference ¼ 0.873, t ¼ 2.228, df ¼ 10) and 50% core
areas (Table 1; difference ¼ 1.248, critical difference ¼
0.819, t ¼ 2.228, df ¼ 10).
Gray foxes.—Gray fox 95% AK home range size and core
area size were similar between zones (95% AKs: F1,25 ¼0.076,
P ¼ 0.785; 50% AKs: F1,25 ¼ 0.720, P ¼ 0.405) and between
sexes (95% AKs: F1,25 ¼ 0.300, P ¼ 0.589; 50% AKs: F1,25 ¼
The Journal of Wildlife Management
70(5)
Table 1. Mean home range size estimates (ha) and standard deviations for bobcats and gray foxes in urban and rural zones of Golden Gate National
Recreation Area, Marin County, California, USA, 1992–1995.

Edited 95% AKa 95% AK 95% MCPb 50% AK

No. individuals Mean SD Mean SD Mean SD Mean SD

Female bobcats
Urban zone 5 150.9 31.0 151.4 31.7 132.1 30.4 20.8 4.3
Rural zone 7 633.1 656.4 634.9 655.1 534.8 496.8 98.6 102.6
Male bobcats
Urban zone 3 638.5 407.8 674.0 413.0 641.4 479.8 141.0 89.0
Rural zone 4 1,453.0 1,254.7 1,673.3 1,490.6 1,351.7 1,262.3 165.5 117.7
Female gray foxes
Urban zone 7 100.3 51.8 100.3 51.8 87.9 61.9 16.1 9.7
Rural zone 6 78.3 31.3 81.8 29.8 61.3 14.5 11.5 2.6
Male gray foxes
Urban zone 9 85.7 40.8 85.7 40.8 59.2 31.0 9.7 8.6
Rural zone 6 96.4 44.5 110.5 64.8 66.2 18.8 13.7 5.1
a
AK ¼ adaptive kernel home range. Edited 95% AK home ranges are the 95% adaptive kernel home ranges with the areas of open water
removed (see methods section in text).
b
MCP ¼ minimum convex polygon home range.

0.430, P ¼ 0.519; Table 1). There was no significant
interaction between sex and zone for home ranges (F1,25 ¼
1.395, P ¼ 0.249) and a marginally significant interaction for
core areas (F1,25 ¼ 3.52, P ¼ 0.072).
Home Range Overlap
Bobcats.—For the 95% AK home ranges, average
overlap was 2.3–5.6 times larger in the rural zone for 3 of
4 sex pairs, but not for female-on-female overlap (Table 2;
Zone: F1,50 ¼ 11.15, P ¼ 0.002). Male-on-male and male-
on-female overlap were 1.8–4.2 times larger than female-
on-male overlap, although differences based on sex combi-
nation were marginal (F3,56 ¼ 2.24, P ¼ 0.093), and there
was also a marginally significant interaction between zone
and sex combination (F3,56 ¼ 2.61, P ¼ 0.060) that was
evident in the female-on-female overlap (Table 2). Core
area overlap was ,6% on average for all zone and sex
combinations, except for female-on-female overlap, which
was 9 times higher in the urban zone (Table 2; U ¼ 42, n ¼
26, P ¼ 0.023).
Gray foxes.—For foxes, there was no consistent relation-
ship between overlap and either zone or sex combination for
95% AK home ranges (Table 3; Sex combination: F3,85 ¼
0.982, P ¼ 0.405; Zone: F1,89 ¼ 0.041, P ¼ 0.840), although
not surprisingly overlap was 1.7–4.0 times higher in paired
than unpaired animals (F1,85 ¼ 18.50, P , 0.001). There
was no significant interaction between sex combination and
zone (F3,85 ¼ 0.276, P ¼ 0.843) or between paired versus
unpaired animals and zone (F3,85 ¼ 0.456, P ¼ 0.501).
Similarly, core area overlap was ,12% for all unpaired foxes
and did not vary by zone (Table 3; male-on-male: U ¼ 52, n
¼ 22, P ¼ 0.502; male-on-female: U ¼ 66, n ¼ 24, P ¼ 0.872;
female-on-male: U ¼ 63, n ¼ 22, P ¼ 0.936). Sample sizes
were too small for core area overlap to make comparisons
with paired foxes or female-on-female overlap, although no
trends were apparent.

Table 2. Average home range overlap (%) and standard deviations for bobcats in urban and rural zone study areas of Golden Gate National
Recreation Area, Marin County, California, USA, 1992–1995.

Overlap (%) of 95% adaptive kernel home ranges

Rural zone Urban zone

Overlap type No. overlaps Mean SD No. overlaps Mean SD

Female-on-female 14 35.3 25.6 14 36.3 28.6

Male-on-male 4 54.2 34.9 6 23.6 8.7
Male-on-female 5 64.2 31.0 6 26.0 26.1
Female-on-male 5 34.8 31.0 6 6.2 7.7

Overlap (%) of 50% adaptive kernel home ranges

Rural zone Urban zone
Overlap type No. overlaps Mean SD No. overlaps Mean SD
Female-on-female 14 4.0 9.9 14 35.9 34.1
Male-on-male 4 0.1 0.2 6 0.0 0.0
Male-on-female 5 5.4 13.1 6 0.0 0.0
Female-on-male 5 2.4 5.8 6 0.0 0.0

Riley
Urban and Rural Bobcats and Gray Foxes 1429

Table 3. Average home range overlap (%) and standard deviations for gray foxes in urban and rural zone study areas of Golden Gate National
Recreation Area, Marin County, California, USA, 1992–1995.

Overlap (%) of 95% adaptive kernel home ranges

Rural zone Urban zone

Overlap type No. overlaps Mean SD No. overlaps Mean SD

Female-on-female 2 19.2 6.1 11 23.8 21.4

Male-on-male 10 29.2 30.2 12 32.0 29.7
Male-on-female, pairs 1 97.0 — 4 83.5 18.7
Female-on-male, pairs 1 82.7 — 4 56.6 24.9
Male-on-female, unpaired 8 36.9 33.9 17 34.5 25.4
Female-on-male, unpaired 8 20.9 21.7 17 33.8 26.6

Overlap (%) of 50% adaptive kernel home ranges

Rural zone Urban zone
Overlap type No. overlaps Mean SD No. overlaps Mean SD
Female-on-female 2 0.0 0.0 11 0.0 0.0
Male-on-male 10 3.9 8.3 12 9.6 17.9
Male-on-female, pairs 1 70.7 — 4 75.9 17.7
Female-on-male, pairs 1 75.0 — 4 61.7 34.2
Male-on-female, unpaired 8 10.8 27.0 17 6.1 14.9
Female-on-male, unpaired 8 7.5 18.0 17 11.4 28.8

Use of Interior Park, Park Edge, and
Developed Areas
Bobcats.—No bobcats were ever radio-located outside of
the park (Fig. 2). Among radiocollared animals, adult male
bobcats in the urban zone used the landscape differently
than adult females. All 3 adult male bobcats either used the
edges of the park out to Highway 101 and over a freeway
tunnel to the borders of the residential areas of Sausalito and
Marin City (B001 and B002), or they had a home range
centered around developed areas (office and residential
buildings) within the park (B005; Fig. 2).
The 5 adult female bobcats that I radiotracked in the
urban zone had home ranges far from the urban edge or
areas of intense human activity (Fig. 2). Adult female bobcat
home ranges centered on dense riparian vegetation
surrounded by scrub and grasslands. One yearling female
was an exception in that she utilized areas along the freeway
and the urban edge. She appeared to be a transient animal
(Litvaitis et al. 1987) searching for a territory, and she was
located throughout the urban zone before settling in a home
range in the interior of the park. Her range after settling also
centered on dense riparian vegetation and grasslands in an
area previously used by 2 older radiocollared females, 7 and
11 years old, that had recently died.
In the rural zone, no radiocollared bobcats ever entered the
town of Stinson Beach or utilized the other small
settlements within the park, although the home ranges of
2 male bobcats included or abutted these developed areas.
Gray foxes.—Of 18 foxes radiotracked in the urban zone,
10 were located in residential areas outside the park at least
once (Fig. 3). Two others were located near areas of intense
human use within the park including stables, offices and
residences. Of the remaining 6 animals, 3 had home ranges
that abutted the Marinview development (Fig. 3). Thus,
only 3 of 18 collared foxes in the urban zone had home
ranges that did not either include or border on developed
areas (Fig. 3), and these 3 animals were also monitored for
4 months each. Although the rural zone was 7–17 km
from any urban centers, foxes in the rural zone also utilized
areas of human use including the town of Stinson Beach,
another small settlement in the northern part of the rural
zone, and a feeding station stocked nightly by a park
resident (see also Riley et al. 2004).
Fox core areas were located within the park (Fig. 4).
Although 2 foxes had core areas centered on stables and
residences within the park, only 1 of 17 animals had a core
area that included any urban area outside the park (Fig. 4).
One fox in the urban zone, G010, died after only 10
locations, so I did not compute home ranges for her,
although 8 of her 10 locations were within the Marinview
community (Fig. 3).
Distance from development.—Distance to development
(Table 4) was not different between male and female foxes
for either home range edges (Mann–Whitney U ¼ 33, n ¼
17, P ¼ 0.761) or radio-locations (U ¼ 31, n ¼ 17, P ¼
0.630), so I pooled male and female foxes for comparisons
with bobcats. On average, the minimum distance between
home range edges and development was 30 times smaller,
and the average distance between radio-locations and
development was 2.2 times smaller for male bobcats than
for female bobcats (Table 4; Minimum home range distance:
U ¼ 15, P ¼ 0.024, n ¼ 8; Average radio-location distance: U
¼ 15, P ¼ 0.024, n ¼ 8). Therefore, I treated male and female
bobcats as separate groups for comparison with foxes.
On average, foxes, male bobcats, and female bobcats
differed for both home range distance (Table 4; H ¼ 11.07,
P ¼ 0.004, df ¼ 2) and average radio-location distance
(Table 4; H ¼ 9.23, P ¼ 0.010, df ¼ 2). Specifically, for home
range distance to closest development, there was no
significant difference between male bobcats and foxes

1430 The Journal of Wildlife Management
70(5)

Table 4. Mean, standard deviation, and range for minimum home range distance to development and average radio-location distance to
development for bobcats and gray foxes in the urban zone of Golden Gate National Recreation Area, Marin County, California, USA, 1992–1995.
Minimum distance from home-range edge
to development
No. individuals Mean SD
Female bobcats 5 893.8 216.3
Male bobcats 3 29.8 51.7
Gray foxes 17 248.6 318.8
(Contrasts with total a ¼ 0.05, Bonferroni correction: mean
rank difference ¼ 3.57, critical difference ¼ 9.03), but female
bobcat ranges were 3.6 times farther from development than
those of foxes (Contrast: mean rank difference ¼ 10.96,
critical difference ¼ 7.34). Similarly, for average radio-
location distance to closest development, male bobcats and
foxes exhibited no differences (Contrasts with total a ¼ 0.05,
Bonferroni correction: mean rank difference ¼ 2.53, critical
difference ¼ 9.03), while on average female bobcat locations
were 1.7 times farther from development than those of foxes
(mean rank difference ¼ 10.60, critical difference ¼ 7.34).
Home Range Use and Roads
In the urban zone, 2 males (B001 and B002) were the only
bobcats with home ranges near roads (Table 5). They were
both located adjacent to the 101 freeway but were never
located across it except in the vicinity of the ridge over the
freeway tunnel (Fig. 2) where they did not have to cross the
freeway surface itself. Seven bobcats in the rural zone and 5
foxes in both the rural and urban zones had home ranges
along roads (Table 5). Roads represented home range
boundaries for 7 of 9 bobcats but for only 2 of 10 foxes that
lived near roads (Fisher exact test, P ¼ 0.023). The 2 bobcats
for which .5% of locations were across roads were yearling
females, while 6 of the 7 bobcats for whom roads
represented home range boundaries were adults.
Discussion
Rarely has the spatial ecology of either bobcats or gray foxes
been investigated in urban landscapes, and it was interesting,
although perhaps not surprising, that both carnivores were
predominantly located in natural habitat within the park.
Both species were affected by development and roads, but as
I expected, bobcats were affected more than foxes. Although
the core areas of both species were within the natural habitat
of the park, no radiocollared bobcats were ever located in
developed areas outside the park, but .50% of collared gray
foxes were, particularly at night. For gray foxes, home range
size, core area size, home range overlap, and core area
overlap were not significantly different between the urban
and rural zones of the park, whereas for bobcats, home range
size, home range overlap, and core area size were all related
to zone, and female bobcats, in particular, had smaller home
ranges, smaller core areas, and greater core area overlap in
the urban zone. Roads also represented home range
boundaries for 75% of the bobcats that lived near them,
but for only 1 in 5 foxes. Carnivore spatial ecology can vary
Riley
Urban and Rural Bobcats and Gray Foxes
Average radio-location distance
to development
Range Mean SD Range
534.1–1080.1 1493.4 53.0 1424.2–1544.6
0.0–89.5 665.6 414.5 313.4–1122.4
0.0–894.8 842.8 487.4 227.2–1690.9
between regions and landscapes, and I hope that future
studies will continue to elucidate the relationship between
these common predators and expanding urbanization.
Although they are few, other studies of the relationship
between development and gray foxes and bobcats have
found similar results. Harrison (1997, 1998) compared gray
fox ecology and recorded bobcat sightings in rural residential
and undeveloped landscapes in New Mexico, USA. Foxes
also appeared to tolerate and even benefit from residential
areas as long as the human density was not too high (,125
residences/km2), and home range size was not significantly
different between foxes in the residential and undeveloped
landscapes (Harrison 1997). Harrison (1997) also found that
gray foxes could coexist with roads, especially if culverts
were present, and that foxes in the residential area had more
irregularly shaped home ranges that included satellite areas,
peninsulas, and intra-home range gaps. I found no
indication of these kinds of home range irregularities, but
this may have been because the New Mexico foxes lived
entirely within the residential areas. The urban zone foxes in
my study were all initially captured within the park and
maintained core areas outside the residential areas, even
though they moved into them. It is possible that there were
foxes that were more urban-associated than those that I
radiotracked (i.e., animals that maintained their entire
ranges within the developed landscape). Interestingly,
bobcats in New Mexico were sighted by homeowners often
,25 m from buildings (Harrison 1998). Nielsen and Woolf
(2001) also found bobcats near, and occasionally within,
human structures in rural Illinois, although overall bobcats
avoided the area around human dwellings, and there were
fewer dwellings within bobcat core areas than within home
ranges. In a highly fragmented landscape in southern
California, bobcats, particularly females, also utilized
developed areas less than a sympatric canid, in that case
coyotes (Riley et al. 2003), although bobcats did utilize
developed areas more than in the current study.
The different diets of foxes and bobcats likely contributed
to differences in the use of development. Although the more
omnivorous foxes may find ornamental fruits and nuts,
insects, and human foods (e.g., pet food or garbage) in
residential areas, none of these items are common food for
the more strictly carnivorous bobcats. Many successful urban
mammals, including raccoons, red foxes, opossums (Didel-
phis virginianus), and coyotes, are similarly omnivorous, and
I found that fruit and insects were important in the diet of
foxes (Riley 1999, 2001). Although residential or developed
1431
Table 5. Percentage of radio-locations across roads from core area, for bobcats and gray foxes in urban and rural zones of Golden Gate National
Recreation Area, Marin County, California, USA, 1992–1995.

No. locations % locations

Zone Age and sexa No. locations across roads across roads

Bobcats
B017 Rural Ad F 96 2 2.1
B020 Rural Ad F 130 2 1.5
B018 Rural Yg F 134 50 37.3
B024 Rural Ad M 120 3 2.5
B023 Rural Ad F 110 1 0.9
B022 Rural Yg F 102 24 23.5
B021 Rural Ad F 89 1 1.1
B001 Urban Ad M 214 0 0.0
B002 Urban Ad M 217 0 0.0
Gray foxes
G047 Rural Yg F 46 1 2.2
G045 Rural Ad M 57 23 40.4
G046 Rural Yg M 54 23 42.6
G041 Rural Ad F 138 66 47.8
G042 Rural Ad M 139 40 28.8
G019 Urban Ad M 152 35 23.0
G008 Urban Ad M 57 11 19.3
G002 Urban Ad F 157 16 10.2
G024 Urban Ad F 133 1 0.8
G004 Urban Ad M 206 83 40.3
a
Ad F ¼ Adult female, Yg F ¼ yearling female, Ad M ¼ Adult male, Yg M ¼ yearling male.

areas may have provided occasional bobcat prey in the form
of woodrats (Neotoma fuscipes), Norway (Rattus norvegicus)
or black rats (Rattus rattus), house mice (Mus musculus), or
songbirds, quality habitat for the common bobcat prey items
in this region (e.g., lagomorphs [Lepus californicus and
Sylvilagus bachmani], pocket gophers [Thomomys bottae], and
especially voles [Microtus californicus]; Riley 2001) were not
present. The residential areas of southern Marin County
were mostly built on steep forested hillsides and lacked the
grassland habitat (or even lawns) preferred by these species. I
do not have information on the abundance of prey species
from inside or outside of the park, so it is possible that even
though the diversity of prey decreased outside the park, the
overall abundance of suitable prey may not have.
Foxes actually appeared to reach their highest densities in
the areas near and including urbanization (Fig. 3), although
Figure 3 should not imply that foxes did not utilize all of the
natural zones of the park, since I was not able to radiotrack
every fox in the area or even every fox that I captured.
Increased food resources and perhaps a reduction in
interference competition from other carnivores, specifically
bobcats (Riley 2001), could contribute to increased fox
density along the urban edge. Coyotes, which can cause
significant gray fox mortality (Fedriani et al. 2000, Farias et
al. 2005), had been extirpated from the urban zone and were
essentially absent during my study.
If individual animals avoid areas outside a reserve or even
the edges of the reserve, then the reserve is smaller than its
administrative boundaries for that species (Schonewald-Cox
and Bayless 1986). However, if a species is able to utilize the
whole park and the matrix outside of it, then the reserve
essentially becomes larger than just the area within it. The
urban gray fox population I studied used the entire park out
to its borders and even across them, so for this species the
area of usable landscape was larger than the park. However,
foxes maintained core areas almost entirely within the
natural habitats of the park, so gray fox populations may still
require a certain amount of undeveloped area adjacent to
development to persist. Moreover, utilization of developed
areas could expose foxes to risks associated with develop-
ment (e.g., increased exposure to disease from domestic
animals; Riley et al. 2004). Radiocollared bobcats did not
cross park boundaries, so for this species, the park was at
most as large as its official size. If female bobcats are more
sensitive to human presence or disturbance than males and
avoid the urban edge (see below), then the acreage of the
urban zone of the park that effectively supports bobcat
reproduction could actually be smaller than its boundaries.
Both bobcats and gray foxes exhibited less landscape
flexibility, i.e., they utilized less of the landscape than, for
example, red foxes in England, which den in backyards and
maintain home ranges and core areas entirely within cities
(Saunders et al. 1997).
The fact that bobcats crossed hard landscape boundaries
such as roads and development less often than foxes may
have contributed to the greater differences between the
zones in home range size and overlap. In comparison to
other studies, the bobcat home ranges (for both sexes) from
the urban zone were among the smallest reported (Anderson
and Lovallo 2003), with a few other studies in the southeast
(Marshall and Jenkins 1966, Hall and Newsom 1976, Miller
and Speake 1979) and 2 in southern California (Lembeck
and Gould 1979, Riley et al. 2003) reporting bobcat home
ranges of a similarly small size. For gray foxes, there are
relatively few studies for comparison, particularly with
sample sizes .10; however, home ranges from the urban
and rural zones were similar to those in other parts of the
west (Cypher 2003). Small home ranges are likely related to

1432 The Journal of Wildlife Management
70(5)

high-quality habitat, and indeed the smaller home ranges
must obviously still provide the requisite resources to sustain
the individual bobcats. In urban landscapes such as my study
area, bobcats may also have smaller home ranges because of
the reduction of suitable habitat, and for the female bobcats
in my study, smaller home ranges and more intra-sexual
overlap were consistent with restricted use of the landscape.
Red foxes (Harris 1981) and an avian predator, Cooper’s
hawks (Mannan and Boal 2000), have exhibited high
population density and small home ranges in urban areas,
although this may be related to rich urban resources rather
than, or in addition to, a restriction of available space.
The reluctance of bobcats to cross paved roads also is
consistent with the results of other bobcat and lynx (Lynx
canadensis) studies. Bobcats in Wisconsin, USA, crossed
paved roads less than expected given the road density within
home ranges (Lovallo and Anderson 1996), and Canadian
lynx seemed to be indifferent to unpaved forest roads
(Mowat et al. 2000) but avoided crossing paved roads and
also treated roads as home range boundaries (Apps 2000).
Bobcats may actually prefer unpaved roads for travel and
hunting (e.g., Bradley and Fagre 1988), and bobcats in my
study often deposited scat or were seen on dirt roads in the
park, roads which were not open to vehicle access by the
public. In the rural zone, the only 2 bobcats for which .5%
of locations were across roads were yearling females, and a
yearling was the only female in the urban zone to move out
to the park edge and near the freeway. Younger female
bobcats may be more willing to cross roads and venture near
developed areas than adult females, since they are less likely
to have kittens and may be searching for an empty territory
(see also Riley et al. 2003).
The increased frequency of road crossings by foxes could
lead to more deaths from vehicles, although even animals
that rarely traversed roads were susceptible to this mortality
source. Only one radiocollared fox was killed by a vehicle
during my study, but I recovered uncollared foxes killed by
vehicles, usually young animals, on highways in the urban
and rural zones. Late in my study, 2 bobcats were killed by
vehicles: a juvenile male bobcat that dispersed from the
urban zone to the rural zone, and an adult female in the
rural zone that was located across paved roads only once, out
of .120 locations, before her death. In urban southern
California, roads were the major source of bobcat mortality
even for animals with little or no developed area within their
home range (Riley et al. 2003). As in the current study,
roads often formed home range boundaries in southern
California, and a large freeway represented a significant
barrier, not only to movement but also to gene flow (Riley et
al. 2003, 2006). Land managers and planners should be
aware of the potential effects of paved roads, both large and
small, on individuals and populations of carnivores such as
bobcats.
Adult female bobcats were located in or near urbanized
areas significantly less often than gray foxes or male bobcats,
and no female bobcat home ranges abutted urban areas.
There are a number of potential explanations for this result.
Riley
Urban and Rural Bobcats and Gray Foxes
One possible explanation is that I did not capture and
radiotrack the adult female bobcats using the urban edge. It
is my experience, as well as that of other bobcat researchers,
that female bobcats can be particularly difficult to recapture,
and generally male bobcat home ranges overlap those of
multiple females (Anderson and Lovallo 2003). Based on
my experience in the field, I do not believe that uncollared
female bobcats occupied the urban edges in my study area.
Because my primary goal was to evaluate the effects of
urbanization on both species, I trapped intensively along the
urban edge when I initially radiocollared animals and again
at the end of my study, when I attempted to remove
radiocollars, and I captured 2 male bobcats and many foxes
that used the edge. Also, in a heavily visited National Park
where bobcats were regularly visible, I never saw uncollared
bobcats near the residential areas nor received reports of
them, but I did receive reports from park employees and
visitors of collared bobcats in edge areas. However, it is
essentially impossible to reliably establish the absence of
uncollared animals, so the presence of uncollared female
bobcats remains a possibility. Another potential explanation
is that female bobcats are more wary of areas of intense
human use than males (or foxes). If so, it may be because
female bobcats raise kittens on their own and must,
therefore, maintain secure denning areas and fulfill the
resource needs of both themselves and their offspring within
their home range. Other studies in a highly fragmented
urban landscape in southern California have also docu-
mented sex differences in the sensitivity of bobcats to
urbanization (Tigas et al. 2002, Riley et al. 2003). If it
proves to be real and repeatable, the sexual difference in
space use in these urban studies represents another example
of sexual dimorphism in bobcats, a species with pronounced
dimorphism both morphologically and ecologically. Male
bobcats are larger, have consistently larger home ranges
(Anderson and Lovallo 2003), and often take larger prey
(Fritts and Sealander 1978, Toweill 1982, Litvaitis et al.
1984). Connor et al. (1999) also found that over time, the
home ranges of individual female bobcats decrease in size
while those of males become larger, presumably because
males are searching for more mating opportunities. In urban
areas, differences in landscape use may reflect not only an
increased sensitivity of females to disturbance, but also an
increased willingness by males to explore new areas, perhaps
again in search of new females.
Management Implications
Wildlife managers in urban areas should be aware that gray
foxes may cross roads and venture outside reserves, and they
will, therefore, be more visible to people and more likely to
be involved in conflicts, real or perceived. Bobcats, on the
other hand, may be more restricted to undeveloped natural
areas. Female bobcats, in particular, may be restricted to
higher-quality habitat in the interior of reserves, which is
important from a management perspective because females
drive population dynamics in this species. Conservation
plans or habitat suitability maps for bobcats may need to
1433
incorporate landscape restrictions in urban areas. However,
the reduced tendency for bobcats to cross roads does not
eliminate the potential for vehicle mortality (see also Riley et
al. 2003). Finally, smaller home ranges and higher home
range overlap in urban areas may result in increased bobcat
density and thereby a greater susceptibility to disease or
increased levels of interaction with other species (e.g.,
competitors; Riley 2001).
Acknowledgments
I thank Golden Gate National Recreation Area for logistical
and financial support throughout the field portion of this
work and J. Howell, C. Schonewald, and D. Van Vuren for
encouragement and assistance both in the field and after. I
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