Annual and seasonal home range characteristics of female Asiatic
black bears in Karuizawa, Nagano Prefecture, Japan
Toshiaki Yamamoto1,4, Hiroo Tamatani2, Junpei Tanaka2, Shotaro Yokoyama2, Kumiko Kamiike2,
Masaru Koyama3, Kanako Seki1, Sayoko Kakefuda1, Yuko Kato1, and Natsumi Izawa1
1
Department of Veterinary Nursing and Technology, Nippon Veterinary and Life Science University, Musashino,
Tokyo 180-8602, Japan
2
Department of Bear Management, Picchio Wildlife Research Center, Hoshino Karuizawa, Kitasaku-gun,
Nagano 389-0194, Japan
3
Department of Tourism and Economy, Karuizawa Town Office, Karuizawa, Kitasaku-gun, Nagano, Japan
Abstract: We characterized annual and seasonal home ranges (HR) of 27 female Asiatic black
bears (Ursus thibetanus) during 2003–11 in Karuizawa, Nagano Prefecture, Japan. Mean annual
HR size for bears calculated by the 95% fixed kernel method was 10.0 km2 (SD 5 8.5) and
differed depending on the year. However, annual HR sizes were not affected by reproductive
status (with or without cubs) or age (subadult or adult). Home ranges tended to be larger in
summer than in autumn. We speculate that a contributing factor to differences in HR size was
annual or seasonal fluctuation of food resources. Annual shifts in HR centers and HR overlaps
were estimated to be 1.2 km (SD 5 0.9) and 0.55 km (SD 5 0.13), respectively. Seasonal shifts in
HR centers and seasonal overlaps were 1.1 km (SD 5 1.1) and 0.54 km (SD 5 0.12),
respectively. Together, these results suggest that female Asiatic black bears in our study area do
not migrate long distances seasonally or annually.
Key words: Asiatic black bear, GIS, home range, Japan, overlap ratio, shift distance, telemetry, Ursus
thibetanus
Ursus 23(2):218–225 (2012)
the Environment 2007). Hence, appropriate conservation and management plans are needed for each
population (Mano et al. 2008). Documenting black
bear behavior, including home range sizes and shifts,
is useful for the development of sound conservation
and management plans to reduce depredation levels
while minimizing the impact on bear populations
and habitats (Sutherland 1998, Caro 2007).
In Japan, past studies have showed that male and
female black bears had very different home range
(HR) sizes (Hazumi and Maruyama 1986), that there
were clear seasonal shifts in the HR sizes (Hazumi
et al. 1985), and that, in the case of black bears with
access to an alpine zone, there was a distinct seasonal
shift in HR elevation (Izumiyama and Shiraishi 2004).
Furthermore, little is known about the influence of
age and reproductive status of females on HR
characteristics (Dahle et al. 2006, Moyer et al. 2007).
To our knowledge, no study has quantified annual
shifts or overlaps of HRs of Asiatic black bears.
We conducted GIS analyses using radio telemetry
data of 27 female Asiatic black bears collected
between 2003 and 2011 in Karuizawa, Nagano
The Asiatic black bear (Ursus thibetanus), an
umbrella species positioned at the top of the
ecosystems it occupies, requires vast and rich natural
habitats. However, in Japan, the impacts of human
activities and encroachment on bear habitat and
resulting habitat deterioration and fragmentation
have become concerns in recent years (Japan
Wildlife Research Center 1993). In fact, the Asiatic black bear populations of 6 regions in Japan
(Kyusyu, Shikoku, Western Chugoku, Eastern Chugoku, Kii Peninsula, and Shimokita Peninsula) have
been designated as ‘‘Locally Threatened Population
(LP)’’ on the Red List published by the Ministry of
the Environment (Japanese Environment Agency
2002).
However, conflicts with bears, including those
involving in agriculture, forestry, and attacks on
humans, remain prevalent in most regions where
black bears occur in Japan. Consequently, 1,000–
2,000 bears on average are destroyed as pests
annually, with peaks as high as 4,700 (Ministry of
4
[email protected]
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HOME RANGE CHARACTERISTICS OF ASIATIC BLACK BEARS N Yamamoto et al.
219
Fig. 1. Study area in Karuizawa, Nagano Prefecture, Japan, located in a matrix of forest (dark gray), private
land (striped gray), agricultural and grassland (white), and urban land (black).
Prefecture, central Japan. We documented HR
characteristics (sizes, shifts, and overlap ratios) and
assessed the influence of age and reproductive status.
Materials and methods
Study area
The study area was centered on Karuizawa Town,
Nagano Prefecture, central Japan. Karuizawa Town
consists of forests, private lands, agricultural lands,
grasslands, and urban areas (Fig. 1). The forestlands
(approximately 50%) have been partially protected
Ursus 23(2):218–225 (2012)
as a wildlife sanctuary since the 1940s and are
dominated by Japanese chestnut trees (Castanea
crenata), Mongolian oak (Quercus crispula), and
Japanese white oak (Quercus serrata). Karuizawa is
a popular summer resort in Japan, and second
homes occupy a substantial part of the study area.
Elevations range between 798 m and 2,568 m, and
mean daily temperatures ranged from 23.8uC in
January to 20.7uC in July in 2008. Maximum snow
depth in winter is approximately 50 cm.
The Picchio Wildlife Research Center, which is
located in the study area, has been commissioned by
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HOME RANGE CHARACTERISTICS OF ASIATIC BLACK BEARS N Yamamoto et al.
Table 1. Numbers of locations and tracking times using radio-telemetry for female Asian black bears in
Karuizawa, Nagano Prefecture, Japan. Tracking time was separated into day (0600–1800) and night (1800–
0600). Age of individuals was estimated in 2011. Parentheses indicate the reproductive status: 0 = without cub;
1 = with cub; underbar = unknown.
the Karuizawa Town office to conserve and manage
Asiatic black bears and has been enforcing various
non-lethal depredation control measures since 2000,
including garbage management, hazing and, since
2005, patrols using trained Karelian bear dogs
provided by the Wind River Bear Institute (Hunt
2003). Bears in our study area have been hazed using
fireworks and dogs, although the average number of
actions/bear/yr has been ,5. This may have affected
their behavior, but because these bears were usually
distributed in the forested areas of Karuizawa, we
believe the influence of this hazing on their home
ranges was minimal (J. Tanaka et al., unpublished
data). To further ensure that our results were not
affected by these countermeasures, we excluded from
our analysis all 3 bears that consumed anthropogenic food (e.g., garbage).
Data collection
From 2001 to 2011, 63 bears (27 males and 36
females) were captured in drum traps with honey
bait and 27 female bears were used for the analysis of
HR characteristics. They were anesthetized with
tiletamine and zolazepam (1:1, 10 mg/kg). The
tranquilized bears were sexed, measured, weighed,
and fitted with radiocollars (ATS Inc., Isanti,
Minnesota, USA).
From 2003 to 2011, radiocollared bears were
located by triangulation (White and Garrott 1990)
with handheld receivers and a 3-element Yagi
Ursus 23(2):218–225 (2012)
HOME RANGE CHARACTERISTICS OF ASIATIC BLACK BEARS N Yamamoto et al.
antenna from known positions along roads. A total
of 5,634 locations were recorded (Table 1). We
conducted tracking once a week on average and
recorded only one location per bear per day using
locations obtained from .2 bearings collected within
30 minutes. Tracking occurred during both daytime
(0600–1800) and nighttime (1800–0600) periods.
Several reports indicate that Asiatic black bears are
primarily diurnal (Hwang and Garshelis 2007,
Yamazaki et al. 2008). However, bears that lived in
undeveloped woodland near populated areas were
found to be more nocturnal (Yamada and Ueuma
2009). In our study area, the number of cottages in
forested areas may have influenced bear behavior,
particularly during the summer. Thus, we recorded
bear locations both during the day and night (1,454
daytime locations, 26%; 4,180 nighttime locations,
74%; Table 1). To assess telemetry error, we
estimated the locations of 8 collars placed at
locations unknown to the researchers. The mean
telemetry error was 84.3 m (n 5 8, range: 20–161 m).
We estimated bear ages from an extracted
premolar (Willey 1974). Subadults were defined as
individuals younger than 4 years, because females
become reproductively mature at that age (Katayama et al. 1996). The reproductive status of adult
females was determined by direct observation during
the active period (May–Nov) or by winter den visits
between February and April, during which we could
hear the vocalizations of cubs of the year. We could
not confirm reproductive status during 16 bearyears.
GIS analysis
We plotted location estimates on 1:25,000 scale
maps using ArcView GIS. Bears were radiolocated
an average of 81.7 times/yr (SD 5 36.6; Table 1). We
estimated home range (HR) size with the fixed kernel
method (95% and 50%) using Home Range Extensions (HRT; Rodgers et al. 2007) and bandwidth
selected by least squares cross-validation (LSCV;
Seaman and Powell 1996). We used only female
bears with a minimum of 30 radiolocations between
June and November to estimate the annual HR of
each individual (Seaman et al. 1999). We excluded
locations from December to May to avoid denning
chronology interference.
To estimate seasonal HRs, we separated the active
period into 2 seasons (Benson and Chamberlain
2007): summer (Jun–Aug) and autumn (Sep–Nov).
To ensure the accuracy of seasonal HR estimates,
Ursus 23(2):218–225 (2012)
221
only bears with a minimum of 20 radiolocations per
season were included in the calculations, (mean
number of locations 5 47.3, Appendix). We
compared range shift distances and temporal overlap
ratio between the HRs of each individual in
successive years. We calculated the shift distance
between successive HRs by calculating the centers of
the 95% and 50% isopleths. We measured the shift
distance between paired centers of each individual
using Hawth’s tools for ArcView 9.2 software. The
overlap area of annual and seasonal HRs was
calculated as follows (Minta 1992):
Mean overlap of annual HR~
sffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
Overlap area
Overlap area
|
home range of(i) year home range of (i{1) year
Mean overlap of seasonal HR~
sffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
Overlap area
Overlap area
|
home range of summer home range of autumn
where i 5 year.
Statistical analysis
We tested for differences among years in HR size,
shift distance, and overlap ratio using Kruskal-Wallis
tests, and for differences in seasonal HR size (summer
versus autumn) using Wilcoxon signed-rank tests. All
statistical analyses were performed using SPSS 16.0
for Windows.
Results
Annual and seasonal home range sizes
Mean annual HR sizes of the 27 female bears were
10.0 km2 (SD 5 8.5) and 2.0 km2 (SD 5 1.6) as
estimated with the 95% and 50% kernel ispoleths
(n 5 69). Seasonal 95% HR sizes averaged 9.5 km2
(SD 5 7.6) in summer and 8.5 km2 (SD 5 8.1) in
autumn, and 50% HR sizes were 2.2 km2 (SD 5 1.8)
in summer and 1.8 km2 (SE 5 1.9) in autumn (n 5
50, Table 2). Annual and autumn 95% HR sizes
differed among years (Kruskal-Wallis test: annual
HR, x2 5 15.94, P 5 0.04; autumn HR, x2 5 15.12,
P 5 0.05), but summer 95% HR size did not (x2 5
9.02, P 5 0.34). Summer 50% HR size differed from
autumn 50% HR size, but 95% HR size did not
differ among seasons (Wilcoxon signed-rank test;
222
HOME RANGE CHARACTERISTICS OF ASIATIC BLACK BEARS N Yamamoto et al.
Table 2. Annual and seasonal home range size (SD) for subadult (,4 years old) and adult (.4 years old) female
Asiatic black bears in Karuizawa, Nagano Prefecture, Japan estimated by the 50% and 95% methods. Sample
size (N) is the number of home ranges used to estimate average home range size.
95% HR, Z 5 21.55, P 5 0.12; 50% HR, Z 5
22.23, P 5 0.03, n 5 50). We were unable to
document significant differences in HR size among
any pairs of bear categories (subadults, adults with
cubs, and adults without cubs; P . 0.05 in all cases)
regardless of time period considered or isopleths
used.
Overlap ratio and shift distance of annual and
seasonal home ranges
Bears shifted their range centers only approximately 1.0 km yearly, and HR displayed approximately 50% overlap (Table 3). Specifically, the
annual 95% HR shift distance was 1.2 km and the
overlap ratio was 0.55 km. Seasonal 95% HR shift
distance was 1.1 km and the overlap ratio was
0.54 km. Neither shift distances nor overlap ratios
differed among seasons or years (Kruskal-Wallis
test; P . 0.05 in all cases).
Discussion
Annual and seasonal HR sizes of female bears are
influenced by population density (Dahle and Swenson 2003, Dahle et al. 2006), the quantity of food
resources (Koehler and Pierce 2003), and the
presence of cubs (Dahle and Swenson 2003). Our
study revealed that the average annual HR size of
female Asiatic black bears inhabiting the forest
around Karuizawa was 10 km2. The HR size of
female Asiatic black bears has been investigated
previously in several different regions (Koike and
Hazumi 2008, Ueuma and Yamada 2008, Izumiyama et al. 2009). According to those studies,
HR size of female Asiatic black bears was 10–20 km2
(using the 95% method), which was slightly larger
than those we report here. Regional difference in HR
size may be related to the density of bears. Because
forest land in Karuizawa has been protected as a
wildlife sanctuary for 70 years, the density of Asiatic
black bears is higher than other areas; this may have
resulted in decreased HR size.
HR size of bears in our study differed among years
and seasons, suggesting that the annual and seasonal
fluctuation in resource availability influences HR
size of Asiatic black bears in our region. In
Karuizawa, the summer diets of Asiatic black bears
consist predominantly of herbaceous plants (e.g.
Angelica pubescens), insects (e.g. ants), and soft mast
(e.g. Chinese mulberry [Morus bombycis] and wild
Table 3. Overlap ratios and shift distances of home ranges estimated by the 95% and 5% method between
years and between summer and autumn. Shift distances between years and seasons were calculated from the
centers of annual home ranges for female Asiatic black bears in Karuizawa, Nagano Prefecture, Japan
determined by the 95% and 50% methods.
Ursus 23(2):218–225 (2012)
HOME RANGE CHARACTERISTICS OF ASIATIC BLACK BEARS N Yamamoto et al.
cherries [Prunus spp.]), whereas in autumn diets
consist predominantly of Fagaceae nuts (oak acorns
and chestnuts). Unfortunately, the distribution and
density of those food resources have not been
thoroughly investigated, particularly summer food
resources. However, in northern Japan, the abundance of Fagaceae nuts fluctuates between years
(Oka et al. 2004). Thus, the fluctuation in resource
availability in our study area could help explain our
observation that HR sizes varied markedly between
years and between autumns.
We found no relationships between age (subadult
and adult) and HR size. This is in line with findings
for American black bears (U. americanus; Smith and
Pelton 1990) and brown bears (U. arctos; Glenn and
Miller 1980). We also detected no significant effects
of female reproductive status on HR size. Previous
studies have indicated that in American black bears,
HR sizes may differ between females with cubs and
without cubs, because females with cubs may have
different resource and safety requirements than those
without cubs (Alt et al. 1980). Moyer et al. (2007)
showed that for Florida black bears, differences in
HR size between females with and without cubs was
significant only in years with limited resource
availability. In our study, data for an entire year
were pooled for each bear. Thus, in our study, the
influence of resource availability might mask differences in HR size between females with cubs and
those without cubs.
Our findings on HR shifts and overlap ratios
indicated only slight annual and seasonal movements, similar to other studies (Hashimoto et al.
2003, Ueuma and Yamada 2008). A strong association between HR shifts and overlap ratios and the
distribution and density of food resources has been
suggested (McNab 1963, Clutton-Brock and Harvey
1978). For instance, seasonal shifts may occur when
the distributions of food resources during summer
and autumn vary in elevation (Hashimoto et al. 2003,
Izumiyama and Shiraishi 2004). In Karuizawa, the
small seasonal shifts and large overlap ratios we
observed may be explained by the mosaic-like habitat
throughout the study area, resulting in a relatively
similar food distribution in summer and autumn.
We believe that a better understanding of the
distribution and quantity of food resources in our
study area would enable us to predict HR sizes and
shift patterns more accurately, and this would allow
us to develop more appropriate management and
conservation plans for Asiatic black bears.
Ursus 23(2):218–225 (2012)
223
Acknowledgments
This study was performed as a part of the
Karuizawa town bear management and damage
control project. We thank the staff of the Karuizawa
town office for their assistance and advice in the field.
We also thank A. Konose for her help with our
geographic and information system and the Nagano
Environmental Conservation Research Institute for
estimating the age of the bears we caught. This study
was supported in part by the PRO NATURA FUND
and the Mitsui & Co., Ltd. Environment Fund.
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Associate Editor: R. Harris
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225
Appendix. Annual (Y), summer (S), and autumn (A) home range sizes of each individual estimated by the 95%
method. Values estimated by the 50% method are shown in parentheses. n indicates the number of locations
used in the estimation.
Ursus 23(2):218–225 (2012)