Glacier
Homing Behavior of Transplanted Black Bears, Glacier National Park
Katherine L. McArthur
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HOMING BEHAVIOR OF TRANSPLANTED BLACK BEARS,
GLACIER NATIONAL PARK
National Park Service Progress Report
For use by National Park Service and cooperating agencies
Katherine L. McArthur
Research Division
Glacier National Park
West Glacier, Montana 59936
May 30, 1978
HOMING BEHAVIOR OF TRANSPLANTED BLACK BEARS, GLACIER NATIONAL PARK
KATHERINE L. McARTHUR, Glacier National Park, West
Glacier, Montana 59936
Abstract: Data on the transplanting of 101
black bears (Ursus americanus) on 155 occasions between 1967 and
1976 in Glacier National Park were evaluated to examine the homing
behavior of transplanted black bears and to identify factors that
contribute to transplant success. Females were more likely than males to
return to the capture area; males were more likely than females to
become nuisances in developed areas near the release sites. The more a
bear weighed, up to 100 kg, the more likely it was to return to the
capture area; two-thirds of the transplants of bears weighing 100 kg or
more were successful. Transplant success was very highly correlated with
distance of the transplant and with the estimated elevation gain that a
homing bear would encounter in returning; presence of a physiographic
barrier also contributed to transplant success. It is suggested that
male bears that repeatedly exhibit nuisance behavior may have difficulty
in obtaining adequate natural forage. Evidence suggested that a portion
of the adult population exists as transients, with very large home
ranges. Implications for management are discussed.
TABLE OF CONTENTS
Abstract
Study Area
Methods
Results
Discussion
Management Implications
Literature Cited
Where wilderness occupied by black bears interfaces
with human-use areas, bears are commonly attracted into the developed
areas by garbage, carelessly stored food, and other unnatural sources of
food (Barnes and Bray 1967, Sauer et al. 1969, Piekielek and Burton
1975, Beeman and Pelton 1976, Lindzey et al. 1976, Harms 1978). When a
bear enters a developed area for the first time, its actions may be
governed largely by chance, but if a behavior pattern is reinforced by a
food reward, the bear is likely to repeat it (Stokes 1970:1156). In an
effort to obtain food, bears may learn to threaten people or cause
property damage or personal injury.
In Glacier National Park, inhabited by an estimated
500 black bears (Glacier National Park 1978), management emphasizes
preservation of natural biotic relationships. However, annual visitation
increased dramatically from 67,200 in 1945 to 884,000 in 1967 and
1,660,000 in 1976. Associated with increased visitation were rising
numbers of bear/human conflicts. Management implemented to resolve these
conflicts has involved preventive as well as corrective measures.
Elimination of factors that lead to nuisance bear behavior is the most
effective and economical method for reducing the number of bear/human
conflicts (Beeman and Pelton 1976:87, Harms 1978). Between 1967 and
1970, a concerted effort was begun in Glacier National Park to inform
visitors and make unnatural sources of food unavailable to bears.
Despite this effort, some unnatural food sources continued to exist, as
when bearproof garbage cans were overfilled, when sightseers offered
food to bears grazing at the roadside, or when campers failed to
securely store their food or properly dispose of garbage.
Corrective management involves capture and
transplanting of nuisance bears, and destruction of bears that
repeatedly exhibit nuisance behavior. Erickson and Petrides (1964:60)
stated that troublesome bears can be transplanted without great
likelihood of their return. In contrast, Barnes and Bray (1967:116),
Harger (1967:6), Sauer et al. (1969:216-220), Payne (1975:812),
Piekielek and Burton (1975:23), and Beeman and Pelton (1976:90) observed
that bears have a definite tendency to return to the capture site and
that the likelihood that a bear would return was related to the distance
it was transplanted. Beeman and Pelton (1976:92) stated that rough
topography and dense vegetation may have hindered long movements of
bears in the Great Smoky Mountains. No other study has apparently
considered topographic features.
I am grateful to C. Martinka for providing the
opportunity to conduct this study and for offering valuable guidance and
encouragement. I also thank J. DeSanto, M. Meagher, C. Meslow, and J.
Peek for their thoughtful and constructive reviews of the manuscript.
STUDY AREA
Glacier National Park, an area of 4,100 km, is
situated adjacent to the Canadian border in northwestern Montana. The
park is bisected by the Continental Divide; elevations range from 961 m
to 3,190 m.
The terrain is very rugged. Physiographic features
reflect glacial and, to a lesser extent, fluvial erosion of the uplifted
sedimentary rock. Most precipitation falls west of the Continental
Divide. Up to 250 cm of precipitation falls at upper elevations, mostly
as snow (Glacier National Park 1976:23).
The west side of the park and portions of the east
side are blanketed by spruce (Picea engelmanni x glauca) /
fir (Abies lasiocarpa) forests and various successional stages of
lodgepole pine (Pinus contorta) communities. Bunchgrass
(Agropyron spicatum, Festuca spp.) communities are common on the
east side of the park and occur locally on the west side. One-third of
the park lies above timberline.
In addition to black bears, the park is inhabited by
approximately 200 grizzly bears (Ursus arctos) (Martinka
1974:23).
METHODS
Most bears were captured in or near developed areas
or along roads using baited culvert traps (Erickson 1957) and
subsequently immobilized. A limited number of free-ranging bears were
immobilized with dart guns employing succinylcholine chloride
(Sucostrin) or etorphine (M99). Each immobilized bear was marked with a
numbered metal tag in one or both ears. Sex and weight of the bear were
determined.
Bears were transported to remote sites within the
park and released. Release sites were selected by considering distance
from the capture site and proximity to other developed areas. During
transport, bears were not drugged and were able to see out of the
culvert trap.
Data analyzed included characteristics of the bear as
well as of the transplant. Characteristics of the bear included weight;
sex; whether cubs were present and, if present, whether they were also
transplanted; and the number of times the bear had been previously
transplanted. Since the age of the bear was unknown in most cases, the
bear's weight was used as a general indicator of its age. Whether the
bear obtained unnatural food at the capture area usually had not been
recorded; the influence of this factor therefore could not be evaluated.
Characteristics of the transplant included date, straight-line distance,
and the number of drainages away that the bear was transplanted. In
addition, the route that a homing bear might take was simulated by
plotting a straight-line route and adjusting it to consider topographic
features such as mountain passes and skirting ridges instead of going
directly over them; the total elevation gain that the bear would
encounter in returning to the capture area along this route was
estimated. Also noted was whether the homing bear would encounter a
physiographic barrier such as a sharp ridge at the head of a valley, or
be forced by the terrain to travel outside the eastern boundary of the
park, where there was severe hunting pressure. Data were not generally
available on the number of days until a bear returned, only on the
number of days until it was recaptured.
A transplant was considered successful if the
transplanted bear was not seen again through 1977. An unsuccessful
transplant was defined as one from which the bear returned to the
capture area. Other possible outcomes were that (1) the bear became a
nuisance in another area, or (2) the bear was killed outside the
park.
Data were analyzed using standard statistical
analyses including chi-square, t-test, and linear regression. For
analysis, transplants were separated into first and subsequent
transplants. An experienced bear was defined as a bear that had been
previously transplanted.
RESULTS
A total of 101 black bears were transplanted 155
times between 1967 and 1976 (Table 1). Thirty-five bears were
successfully transplanted the first time; 18 were successfully
transplanted on subsequent transplants.
Table 1. Transplant success associated with various
characteristics of transplanted black bears, Glacier National Park,
1967-76. Percentages are in parentheses.
|
Characteristic | Successful |
Returned to capture area |
Nuisance in different area |
Killed in different area |
Total |
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First-time transplants | 34 (35) | 50 (49) | 11 (11) | 5 (5) | 101 |
Subsequent transplants | 18 (33) | 27 (50) | 6 (11) | 3 (6) | 54 |
Total | 53 (34) | 77 (50) | 17 (11) | 8 (5) | 155 |
Sex |
Male | 29 (38) | 31 (40) | 14 (18) | 3 (4) | 77 |
Female | 17 (27) | 40 (63) | 3 (5) | 4 (6) | 64 |
Unknown | 7 (50) | 6 (43) | 0 (0) | 1 (7) | 14 |
Cubs present |
Entire litter transported | 2 (40) | 1 (20) | 2 (40) | 0 (0) | 5 |
Partial litter transported | 1 (50) | 1 (50) | 0 (0) | 0 (0) | 2 |
No cubs transported | 0 (0) | 3 (100) | 0 (0) | 0 (0) | 3 |
Weight (kg) |
019 | 1 (17) | 1 (17) | 3 (50) | 1 (17) | 6 |
2039 | 11 (42) | 9 (35) | 4 (15) | 2 (8) | 26 |
4059 | 14 (26) | 32 (60) | 6 (11) | 1 (2) | 53 |
6079 | 14 (45) | 16 (52) | 1 (3) | 0 (0) | 31 |
8099 | 1 (13) | 7 (87) | 0 (0) | 0 (0) | 8 |
>100 | 4 (67) | 1 (17) | 1 (17) | 0 (0) | 6 |
Unknown | 8 (32) | 11 (44) | 2 (8) | 4 (16) | 25 |
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Characteristics of the Bear
Erickson and Petrides (1964:50), Harger (1967:7),
Sauer et al. (1969:217), Piekielek and Burton (1975:22), and Beeman and
Pelton (1976:90) found that 78-91 percent of nuisance bears were males.
In Glacier National Park, sex of 91 percent of the nuisance bears was
recorded. Of these, 55 percent were males and 45 percent were females;
this ratio is significantly different from those of the previous studies
(P<0.05).
Transplants of female bears were somewhat less
successful than were transplants of male bears (Table 1). Females were
more likely than males to return to the capture area; transplanted males
were more likely than females to become nuisances in other areas. This
observation supports the statement of Bunnell and Tait (1978:48, 51)
that female bears defend specific territories while males do not.
Beeman and Pelton (1976:50) reported that only 1 (1.3
percent) of 76 nuisance bears was a female with cubs; in contrast, 64
(41.3 percent) of 155 transplants of nuisance bears in Glacier involved
females, and 10 (15.6 percent) of the 64 females had cubs with them.
When at least some of the cubs were transplanted with the mother, the
transplants met with varying degrees of success (Table 1). In each. of 3
instances in which a female was transplanted without any of her cubs,
the female returned to the capture site. One of these females returned
11 km in 8 hours.
Although Barnes and Bray (1967:118) observed that
homing was more prevalent among adults than subadults, Beeman and Pelton
(1976:90) found no significant difference between homing of adults and
that of subadults. In this study, regression analysis showed that weight
of the bear, up to 100 kg, was positively correlated with the percentage
of bears that returned to the capture area (Y = 0.785 X + 10.950,
r = 0.9405). In other words, the more a bear weighed, up to 100
kg, the more likely it was to return to the capture area. In direct
contrast, however, 67 percent of the transplants of bears weighing 100
kg or more were successful; in fact, this was the most successful weight
category. The reason for this relationship could not be ascertained.
Although the differences were not significant
(P>0.50), female bears successfully transplanted on their
first transplant tended to weigh less than those successfully
transplanted on a subsequent transplant (Table 2). This pattern also
held true for females and males that returned to the capture area: bears
transplanted for the first time tended to weigh less than those
transplanted a subsequent time. This tendency was expected because the
average bear transplanted for the first time is probably younger than
the average bear that has been transplanted previously. However, male
bears successfully transplanted on their first transplant tended to
weigh more than those whose subsequent transplants were successful.
Although bears tend to weigh less earlier in the year, the relationship
could not be adequately explained by transplant dates; the mean date of
successful first-time transplants of male bears was 17 July and the mean
date of successful subsequent transplants of male bears was 15 July
(Fig. 1). Male bears that weigh less may be more likely to return to the
capture area if they roam more widely and are familiar with a greater
area than heavier bears. Another possibility is that male bears that
weigh less have difficulty in obtaining adequate natural forage, are
more dependent on unnatural food, and therefore have greater incentive
to return to a known source of unnatural food. This hypothesis is
supported by the fact that the lightest-weight bears were the ones most
likely to become nuisances in other developed areas (Table 1).
Table 2. Mean weights (kg) and standard deviations
(SD) of females, males, and all bears that were transplanted
successfully and those that returned to the capture area, comparing
first-time transplants with subsequent transplants, Glacier National
Park, 1967-76.
|
| Female (±SD)
|
Male (±SD)
| Overalla (±SD)
|
Successful | Returned |
Successful | Returned |
Successful | Returned |
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First-time transplants |
47.2 ± 17.2 (N= 10) |
50.8 ± 18.4 (N= 22) |
63.2 ± 27.6 (N= 21) |
59.4 ± 21.1 (N= 22) |
58.7 ± 25.4 (N= 35) |
55.6 ± 19.8 (N= 50) |
Subsequent transplants |
53.8 ± 20.3 (N= 7) |
51.9 ± 15.0 (N= 18) |
50.0 ± 23.3 (N= 8) |
64.5 ± 15.1 (N= 9) |
51.9 ± 20.6 (N= 18) |
57.0 ± 15.9 (N= 27) |
Total |
49.6 ± 17.9 (N= 17) |
51.2 ± 17.1 (N= 40) |
59.6 ± 26.7 (N= 29) |
60.8 ± 19.6 (N= 31) |
56.6 ± 24.0 (N= 53) |
56.0 ± 18.6 (N= 77) |
aIncludes bears of unknown sex. |
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Fig. 1. percentages of successful
first-time ()(N=21) and successful subsequent (
)(N=8) transplants of male black bears that took
place during 20-day time periods, Glacier National Park, 1967-76.
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The percentage of transplants that were successful
was essentially the same between first and subsequent transplants (Table
1). Experienced bears that returned to the capture area had been
previously transplanted a mean of 1.7 times, whereas experienced bears
that were successfully transplanted had been previously transplanted a
mean of 1.4 times; the difference was not significant
(P<0.25).
Characteristics of the Transplant
Although increased movements by bears during autumn
have been noted (Erickson and Petrides 1964:56, Piekielek and Burton
1975:16), and Barnes and Bray (1967:118) reported greater success of
transplants in autumnboth associated with more abundant local food
sourceslittle correlation was found in this study between success
and time of year (Y = 0.096 X + 54.569, r = 0.3274), nor
was the percentage of bears that returned to the capture area correlated
with time of year (Y = 0.012 X + 53.339, r = 0.0724).
Bears transplanted a greater distance were
significantly less likely to return to the capture area (Fig. 2). In
first-time transplants, the mean distance of successful transplants was
29.9 km, whereas the mean distance of transplants where the bear
returned was 19.2 km; the difference was significant
(P<0.001). In subsequent transplants, the mean distance of
successful transplants was 35.3 km, whereas the mean distance of
transplants where the bear returned was 23.6 km; the difference was not
significant (P<0.25).
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Fig. 2. Percentages of transplants that
were successful () or from which the bear returned (
) in relation to distance of the transplant, Glacier National
Park, 1967-76. Sample sizes are in parentheses.
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The number of drainages across which a transplanted
bear had to return was also important (Fig. 3). In first-time
transplants, the mean number of drainages was 3.3 in successful
transplants and 1.6 in transplants where the bear returned; the
difference was significant (P<0.01). In subsequent
transplants, the mean number of drainages was 4.1 in successful
transplants and 2.4 in transplants where the bear returned; the
difference was not significant (P<0.25).
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Fig. 3. Percentages of transplants that
were successful () or from which the bear returned (
) in relation to the number of drainages across which the bear was
transplanted, Glacier National Park, 1967-76. Sample sizes are in
parentheses.
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The elevation gain a bear would encounter in homing
from the transplant site to the capture area has not been quantitatively
considered in previous studies. This factor was indicated to be an
important component of transplant success (Fig. 4). Data for elevations
of 1,500-1,999 m were not included in Fig. 4. The sample for those
elevations consisted of 3 transplants; none were successful. Two bears
returned to the capture area; one took 3 years and one took 7 years to
do so. The third bear became a nuisance in another area 2 years later.
If these data are included, the regression of percent success on
elevation becomes Y = 0.024 X + 17.292, r = 0.7532; and the
regression of percent returned to the capture area on elevation becomes
Y = 0.017 X + 57.839, r = 0.7386.
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Fig. 4. Percentages of transplants that
were successful () or from which the bear returned (
) in relation to the estimated elevation gain that the homing bear
would encounter, Glacier National Park, 1967-76. Sample sizes are in
parentheses.
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In first-time transplants, the mean elevation gain
involved was 834 m in successful transplants and 352 m in transplants
where the bear returned; the difference was significant
(P<0.01). In subsequent transplants, the mean elevation gain
involved was 1,214 m in successful transplants and 477 m in transplants
where the bear returned; again, the difference was significant
(P<0.01).
Presence of a physiographic barrier was another
apparently important contributor to transplant success. Of 23
transplants where the homing bear would encounter a barrier, 16 (70
percent) were successful; 3 (13 percent) bears returned to the capture
area. Three (13 percent) bears become nuisances in other developed
areas, indicating that a barrier did not simply shunt a problem bear to
another area.
DISCUSSION
Because home range fulfills physical needs as well as
psychic needs by allowing bears to move about familiar areas with
assurance and to efficiently utilize the resources there, transplanted
bears are highly motivated to return to their home ranges (Sauer et al.
1969:216; Beeman and Pelton 1976:93). To preclude bears from returning
to the capture area, they must be removed from the area with which they
are familiar (Sauer et al. 1969:222). Bears released in unfamiliar
surroundings evidently wander at random until they either find a
familiar area or can establish new home ranges (Erickson and Petrides
1964:60).
Previous studies have found mean summer home range
diameters of female black bears to be 0.9-11.9 km, and those of male
black bears to be 2.8-13.8 km (Erickson and Petrides 1964:56, Sauer et
al. 1969:212-216, Jonkel and Cowan 1971:19, Poelker and Hartwell
1973:62, Piekielek and Burtin 1975:16). Although these home range
studies have found that male bears are wider-ranging than females,
females in this study were more apt than males to return to the capture
site. Males, on the other hand, were more likely than females to become
nuisances in developed areas near the release sites. Evidence presented
in this paper indicates that males that repeatedly exhibit nuisance
behavior may have difficulty in obtaining adequate natural forage.
Poelker and Hartwell (1973:61-62) theorized that bear damage to conifers
was related to a shortage of preferred food and that wider ranging of
bears in tree-damage areas reflected more extensive searching for food.
The statement of Lindzey et al. (1976:219) that nuisance bears range
farther than non-nuisance bears is supported by the mean distances from
which nuisance bears returned in this study19.4 km for males and
21.7 km for femaleswhich are substantially greater than reported
normal home range sizes.
Several studies have noted unexplained long-distance
movements by some bears. Amstrup and Beecham (1976:345) observed a
female with a home range of 130.3 km2. Although Erickson and
Petrides (1964:57) maintained that bears could generally be transplanted
without returning, they reported one bear that returned from a
transplant of 103 km and another that returned 154 km. Erickson and
Petrides (1964:56) also reported that a female with a cub moved 31.2 km,
while summer movements of other females averaged 2.3 km. Jonkel and
Cowan (1971:19) noted unusually long movements by 5 adult bears. Such
movements have generally been designated as excursions outside of a
smaller home range, and are undoubtedly important in familiarizing a
bear with terrain and resources beyond its normal home range and assist
it in orienting itself after being transplanted from its home range
(Beeman and Pelton 1976:93). On the other hand, as suggested by Lindzey
and Meslow (1977:421), these may be transient adults, similar to
mountain lions (Felis concolor) described by Seidensticker et al.
(1973:55) and to adult "floaters" reported in several bird species
(Smith 1978). A mixture of resident and transient adults would promote
more complete utilization of resources and ensure the availability of a
breeding-age replacement should a resident bear be removed from the
population. Transient adults would not receive the benefits of intimate
knowledge of a relatively small home range, but would be familiar with a
large area and able to take advantage of widely dispersed sources of
locally abundant food. Such individuals would be difficult to transplant
successfully. They also may make up a large proportion of the nuisance
bear population.
MANAGEMENT IMPLICATIONS
Transplanting is an effective method of dealing with
nuisance bears, as indicated by the fact that 53 of 101 bears in this
study were eventually successfully transplanted. Selection of release
sites is an important consideration in the handling of nuisance bears
(Barnes and Bray 1967:118, Beeman and Pelton 1976:95). Results of this
and other studies show that bears transplanted greater distances are
less likely to return to the capture area. Bears should also be
transplanted across rough terrain, with large elevation changes and
physiographic barriers. Greater effort must be made in transplanting
transient bears, which are probably more difficult to successfully
transplant because they are familiar with a much larger area than are
resident bears.
An abundance of natural food at the release site may
be a major factor in determining the success of a transplant (Piekielek
and Burton 1975:23-24). Sauer et al. (1969:222) stated that where
sources of unnatural food are available near the release site,
transplanting a bear will simply transfer the problem. This study showed
that females and lighter-weight male bears were especially likely to
return to the capture area or to become nuisances in developed areas
near the release site. This may be due to the relative lack of easily
obtainable food at the release site. More research needs to be done
concerning the influence of the abundance and quality of food at release
sites.
Additional research should also be done to ascertain
the activities and fates of transplanted bears that are not seen
againbears that, in this study, were considered successfully
transplanted. Results of studies on the responses of bears released in
unfamiliar surroundings will be useful in programs to reestablish
populations of threatened or endangered species in formerly occupied
habitat.
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