Distribution of polar bears (Ursus maritimus) during the ice-free period in

western Hudson Bay

Department of Zoology, University of Alberta, Edmonton, Alta., Canada T6G 2E9

AND

IANSTIRLING
Canadian Wildlife Service, 5320 122 Street, Edmonton, Alta., Canada T6H 3S5 and Department of Zoology,
University of Alberta, Edmonton, Alta., Canada T6H 2E9
Received May 5, 1989

DEROCHER, A. E., and STIRLING, I. 1990. Distribution of polar bears (Ursus maritimus) during the ice-free period in western
Hudson Bay. Can. J. Zool. 68: 1395- 1403.
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The distribution and movements of polar bears (Ursus maritimus) during the ice-free period in western Hudson Bay were
studied using mark and recapture and radiotelemetry locating of animals of known age and sex collected between 1966 and 1987.
Polar bears segregated themselves by age, sex, and reproductive status. Adult males occupied coastal areas. Family groups and
pregnant females occupied areas farther inland. Subadult females and males occupied similar habitats which overlapped with
those of adult males. Avoidance of conspecifics, energy conservation, philopatry, denning habitat, and habitat selection
appeared to influence observed patterns. A southward shift in the population was evident early in the ice-free period and was
followed by a return movement northward during October and November. This may be due to ice-formation patterns which
permit earlier access to the sea ice along the north coast and hunting habitat to the northeast. All bears moved less than bears on
the sea ice, and movements were consistent with a strategy of energy conservation. Most bears appear to have travelled directly
from the sea ice to the study area and remained until the sea ice reformed. Little exchange with adjacent populations and a high
degree of philopatry were evident for all age and sex groups; this may be a function of the distribution of denning habitat, the
winter distribution of sea ice habitat and seals, and the noncompetitive conditions that prevail during the ice-free period which
make dispersal of limited benefit.

DEROCHER, A. E., et STIRLING, I. 1990. Distribution of polar bears (Ursus maritimus) during the ice-free period in western
For personal use only.

Hudson Bay. Can. J. Zool. 68 : 1395- 1403.

Des techniques de marquage-recapture et de radiotklkmktrie ont servi a ktudier la rkpartition et les dkplacements d'Ours
polaires (Ursus maritimus) au cours de la pkriode sans glace dans l'ouest de la Baie d'Hudson en permettant de repkrer les
animaux d'ige et de sexe connus entre 1966 et 1987. Les ours se rkpartissaient selon leur ige, leur sexe et leur statut reproducteur.
Les miles adultes occupaient les rkgions cbtieres. Les farnilles et les femelles enceintes se tenaient dans les zones plus
continentales. Les femelles et les miles subadultes occupaient des habitats semblables qui chevauchaient ceux des miles adultes.
La fuite des autres ours, la conservation de l'knergie, la philopatrie, le lieu du repaire et le choix de l'habitat semblent les facteurs
dkterminants des rkpartitions observkes. La population s'est dkplacke vers le sud au dkbut de la pkriode sans glace et il s'est
effectuk un mouvement de retour vers le nord en octobre et novembre. Ces dkplacements sont probablement dus aux mouvements
de formation des glaces qui permettent un acces plus rapide aux glaces marines le long de la c6te nord et aux zones de chasse vers
le nord-est. Tous les ours se dkplasaient moins durant la pkriode sans glace que sur les glaces marines et ces dkplacements
semblaient rkpondre a une stratkgie de conservation d'knergie. Les ours semblent tous s'Ctre rendus directement des glaces
marines a la rkgion d'ktude et y sont restks jusqu'au retour des glaces. Chez les ours des deux sexes et de tous les Ages, il y avait
peu d'interactions avec les populations adjacentes et une nette tendance a la philopatrie, phknomhnes probablement reliks a la
rkpartition des repaires, a la rkpartition des glaces marines et des phoques en hiver, et aux conditions qui suppriment la nkcessitk
de compktition durant la pkriode sans glace et qui font que la dispersion comporte peu d'avantages.
[Traduit par la revue]

Introduction 1977; Ramsay and Andriashek 1986). 'The breeding season

Polar bears (Ursus maritimus) have a circumpolar distribu-
tion closely tied to the arctic pack ice (DeMaster and Stirling
1981). They are solitary carnivores, preying predominantly
upon ringed seals (Phoca hispida) and secondarily on bearded
seals (Erignathus barbatus) (Stirling and Archibald 1977;
Smith 1980). In some polar bear populations, such as the one in
western Hudson Bay, the bears migrate onto land when the sea
ice melts during the summer (Stirling et al. 1977). While on
land, polar bears spend 70-90% of their time resting (Latour
1981; Lunn and Stirling 1985) and feed little (Russell 1975;
Knudsen 1978; Lunn and Stirling 1985). Fat deposited during
the previous spring is their predominant energy source. All
bears, except for pregnant females, return to the sea ice in
November Or early December. Pregnant femiles remain inland
in dens where they give birth to cubs in late December or
January (Harington 1968). They return to the sea ice between
mid-February and early April (Harington 1968; Stirling et al.
occurs while polar bears are on the sea ice from lace March
through May (L@n@1970).
Polar bears on land in western Hudson Bay are segregated by
age and sex class (Stirling et al. 1977). In general, adult males
predominate along the coastal areas, whereas adult females with
and without cubs are found farther inland (Stirling et al. 1977;
Latour 1981). The purpose of this study was to quantify the
spatial and temporal distribution and movements of polar bears
during the summer and autumn and examine possible reasons
for the observed patterns. In addition, we also examine patterns
of philopatry and site-specific fidelity within the study area.
Studv area
~ h ,study area is the western Hudson Bay coast between the
Churchill and Nelson rivers, inland to94O15'W. The western boundary
corresponds to the maximum westward movements observed (Fig. . " 1).
~ ~ ~ r o . x i m a700-900
t e l ~ bears are found in the study area during the
 1396 CAN. J. ZOOL. VOL. 68, 1990
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For personal use only.
FIG. 1.. Location of the study area and locations mentioned in the
text.
ice-free period (Derocher 1987). The habitat is a flat peatland along the
ecotone between the subarctic open boreal forest and the arctic tundra.
Lichen tundra, numerous small lakes, and bogs characterize the inland
area. Open scrub spruce (Picea spp.) forest is present along rivers,
streams, some lakes, and throughout the southern quarter and along
the western edge of the study area. The coast is characterized by sedge
and grass meadows, extensive salt marshes, and patches of willow
(Salix spp.). Mean temperatures are between 12°C in July and - 12°C
in November. During the summer, daytime temperatures can exceed
25°C.
The annual ice in Hudson Bay begins to break up in May and is gone
by mid-August. The last of the major ice floes occur off the coast of
Manitoba and Ontario. Freezeup begins along the coast north of Rankin
Inlet in late October and moves southward, forming a rim of ice several
kilometres wide. By the first or second week of November, the ice
extends south to Cape Churchill, and most of the bay freezes within the
next few weeks.
Methods
Data were collected between 1966 and 1987, although the objec-
tives, methods, and intensity of tagging operations have varied. Most
animals were immobilized from a Bell 206B helicopter, between
August and October, using remote injection equipment. Techniques
were modified from those outlined by Lentfer (1968) and Larsen
(1971). Some bears were captured in Aldrich leg snares or culvert
traps, and others were darted from motor vehicles. The sex, reproduc-
tive status, condition, and other selected parameters were recorded
from immobilized bears. Ages were determined from an extracted pre-
molar, and weights were estimated from axillary girth (Stirling et al.
1977). A plastic tag was placed in each ear and tattoos were applied to
both upper lips with a unique number to facilitate long-term identifica-
tion. Temporary letters or numbers were painted on the back of each
bear to permit identification from aircraft. Eartag, glue-on, or neck-
collar radios were placed on 166 bears. Relocations of marked and
radio-tagged bears were obtained during capture sessions, aerial sur-
veys, and from reliable observers.
Bears were classified into one of five groups: family groups
(females with cubs of the year or yearlings), solitary females (5 years
and older), subadult females (independent females 1-4 years old),
adult males (5 years and older), and subadult males (independent males
1-4 years old). Because of limited variation between adult females
with cubs of the year and those with yearlings (Derocher 1987), these
two groups were pooled. There was little variation in distribution and
movement patterns between years (Derocher 1987), so the data were
pooled.
Locations of known animals were described according to their
distance from the coast. North-south distributional aspects were not
considered in detail because sampling constraints in some years may
have introduced biases. Only relocations of animals made within 30
days were used for the analyses of daily movement distances. Direction
of movement was calculated using movements of 2 km or more, as
shorter movements were within the range of possible mapping error in
some habitats.
Data were analyzed using BMDP (Department of Biomathematics,
The University of California, Los Angeles) and SAS (SAS Institute
Inc., Cary , NC) statistical software. Kruskal- Wallis (K- W) and
Mann- Whitney (M-W) tests were used for most comparisons, as data
distributions were highly skewed. Multiple-comparison tests were
performed by analyzing the ranks of the distance from the coast using
the GT2 method (Sokal and Rohlf 1981). For all tests, probabilities
greater than 0.05 were considered not significant. Directional data
were analyzed using circular statistical methods, including the Rayleigh
test for nonrandom orientation (Batschelet 1981). Where visual inspec-
tion of plotted data indicated movement in opposite directions, an axis
of orientation was determined.
Population definition was examined using mark and recapture data
and returns from tagged bears killed by Inuit. Seasonal fidelity was
evaluated from locations where an individual bear was recaptured
during the same month, in different years. This was done for adults,
subadults relocated as adults, subadults relocated as subadults, and
cubs relocated as independent bears. Emigration from the study area
was also investigated by examining longer distance movements of
bears caught in the study area.
Results
Between 1966 and 1987, 1528 captures and 1073 resightings
of known independent animals were recorded. During the
autumn of most years, there is an influx of females with cubs
and subadults into the area near the Churchill townsite, and
these animals constituted a large portion of the sample prior to
1977 (Stirling et al. 1977; Lunn and Stirling 1985). After 1977,
the region sampled increased, and by 1980, animals were
captured throughout the study area. Over 65% (1704/2601) of
the locations of known polar bears have occurred since 1980.
The overall mean resighting or recaptured interval was 20 days
(SE = 1, n = 1201) or 7 days (SE = 0.3, n = 9 13) for bears
relocated within 30 days.
Distribution
The monthly distributions of bears located between 1966 and
1987 are described by the mean and range of distances from the
coast (Table I). Individuals of all five groups could be found
along the coast throughout the study period, and therefore the
range is equal to the maximum distance from the coast.
Significant differences were found between groups in their
mean distance from the coast (August, K-W: H = 92.3, df = 4,
P < 0.001; September, K-W: H = 171.2, df = 4, P < 0.001;
 DEROCHER AND STIRLING: I1

TABLE1 . Distance from the coast (km) of polar bears of different age, sex, and reproduc-
tive status for July-November

Lone
adult Family Subadult Subadult Adult
females groups females males males

July
n
Mean (SD)
Median
Range
August
n
Mean (SD)
Median
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Range
September
n
Mean (SD)
Median
Range
October
n
Mean (SD)
Median
Range
November
n
For personal use only.

Mean (SD)
Median
Range
NOTE:Within each month (except July, when there were insufficient data for analysis), groups whose range values
are followed by the same letter are not significantly different (GT2 test, P > 0.05).

October, K-W: H = 276.2, df = 4, P < 0.001; and November,
K-W: H = 49.0, df = 4, P < 0.001). Adult males were
significantly separated from all other groups during August
(Table I). There were no significant differences in distribution
between subadult females, subadult males, and adult males
during September, October, and November, although these
groups occupied different areas than family groups and solitary
females during this period (Table 1). Although family groups
and solitary females were not significantly separated in Novem-
ber, solitary. females occupied the areas farthest inland. Sub-
adults of both sexes and adult males used areas closer to the
coast. There was a wide degree of variation within each group
and individuals of each group could be found throughout the
study area. Adult males tended to aggregate at prominent points
and small islands along the coast.
Monthly variation in observed distances from the coast was
found in all groups (family groups, K-W: H = 61.9, df = 4,
P < 0.001; solitary females, K-W: H = 58.3, df = 4, P <
0.001; subadult females, K-W: H = 20.8, df = 4, P < 0.001;
subadult males, K-W: H = 27.8, df = 4, P < 0.001; adult
males, K-W: H = 52.4, df = 4, P < 0.001). Family groups,
adult males, and subadults were nearest the coast in July, as they
arrived from the sea ice, and movement inland reached a
maximum in August or September. Changes in observed distance
from the coast varied considerably between the different
segments of the population. Family groups were the last to
leave inland areas. Solitary females differed from all other
groups because pregnant females remained inland to give birth
to young.
Solitary females that remained along the coast or returned
there during October and November after being inland repre-
sented 21% (221106) of the group. Solitary females found in
October within 10 krn of the coast weighed significantly less (t =
-2.77, df = 62, P = 0.007) than those found farther inland
(coast: i = 173 kg, SE = 9, n = 13; inland: i = 224 kg,
SE = 9, n = 5 1). There were no differences (t = - 1.61, df = 62,
P = 0.113) in their mean ages. During November, these
differences persisted (t = -3.76, df = 33, P = 0.007; coast:
i = 156 kg, SE = 14, n = 16; in1and:i = 223 kg, SE = 11,
n = 19), although there were still no significant differences (t =
0.62, df = 33, P = 0.542) in their ages.
Movement patterns
There were no significant differences between groups (K-W
test) in mean distance moved per day. However, within groups,
significant variation in distance moved per day was found
between months for family groups (K-W: H = 8.2, df = 3, P =
0.043), subadult females (K-W: H = 20.2, df = 3, P < 0.001),
arid adult males (K-W: H = 22.9, df = 3, P < 0.00 1). These
three groups moved less during September than either before or
afterwards (Table 2). When nonpregnant females were removed
from the solitary female group, there was significant variation
between months (K-W: H = 8.73, df = 3, P = 0.033) in
distance moved per day. The shortest daily movements were
made during November (Table 2).
The movements of several groups showed a significant
degree of orientation. Most bears moved south along the coast
or inland after arriving on land and then returned northward
 CAN. J . ZOOL. VOL. 68, 1990

TABLE2. Distance (krn)moved per day for polar bears of different age, sex, and repro-
ductive class

Family Solitary Subadult Adult Subadult

groups females females males males

August
n
Mean (SE)
Median
Range
September
n
Mean (SE)
Median
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Range
October
n
Mean (SE)
Median
Range
November
n
Mean (SE)
Median
Range

TABLE3. Proportion of tagged polar bears in capture and harvest ( r = 0.45, n = 111, P < 0.001), 352"(r = 0.41, n = 103,
For personal use only.

samples from locations about Hudson Bay (locations are listed clock- P < 0.001), and 329" (r = 0.41, n = 22, P = 0.027),
wise, starting from the study area) respectively. An axis of orientation was found during September
with a bearing of 348"-168" ( r = 0.48, n = 63, P < 0.001).
Capture Bears tagged Subadult males as a group had a significant nonrandom orienta-
and in study area Proportion tion during October, with a mean heading of 328" (r = 0.29,
Location harvest sample in sample tagged
n = 40, P = 0.034).
Study areaa
Eskimo Point Population definition
Whale Cove Bears tagged in the study area are recaptured or killed
Rankin Inlet throughout the Hudson Bay region (Table 3). The harvest of
Chesterfield Inlet polar bears along the western coast of Hudson Bay from Eskimo
Coral Harbour Point to Rankin Inlet is composed of 44% subadult males, 27%
Southampton 1slandb adult males, 20% subadult females, and 9% adult females (n =
Quebec coast 509). In comparison, the approximate composition of the
Belcher Islands population of independent bears from the capture sample is 28%
Ontario
Southeast ~ a n i t o b a ~ adult females, 28% adult males, 19% subadult females, and
25% subadult males (n = 1528). While information from
"Includes spring captures. eastern Hudson Bay is limited and the total harvest unrecorded,
bGovernment of N.W.T. study. few bears tagged in the study area were reported there. The
'Harvest was not recorded in Quebec.
d ~ r e from
a the mouth of the Nelson River east to the Manitoba-Ontario border. proportion of bears tagged in the study area, relative to the total
number that are captured or killed in the study area or elsewhere,
provides an index of population distribution and fidelity
prior to sea ice formation. In August and October, the (Table 3).
movements of family groups had a mean heading of 196" (r = Of 122 bears originally tagged in the study area and harvested
0.44, n = 21, P = 0.015) and 30" (r = 0.42, n = 48, P < between Eskimo Point and Rankin Inlet, 46 (38%) were known
0.001). Significant orientation was not found during September to have been within the study area in the previous 6 months. Of
and November. A significant ( P < 0.001) orientation of 169" the eight tag returns from Quebec, six were from males with a
(r = 0.70, n = 15) was found for solitary females during August. mean age of 4 years (SE = 0.3, n = 6). All eight bears were
A significant (P < 0.001) axis of orientation of 339"- 159" (r = known to have been in the study area within a mean of 13
0.92, n = 12) was found during September. Subadult females months (SE = 6, n = 8), although four were killed within 5
showed no significant orientation during August and Septem- months. The single tagged bear killed in the Belcher Islands was
ber, but showed a significant mean bearing of 324" (r = 0.30, a 5-year-old male that had moved over 800 km in the 4 months
n = 41, P = 0.026) and 340" ( r = 0.32, n = 28, P = 0.046) since it was last handled near Churchill. There were 28
during October and November, respectively. Adult males recaptures in Ontario of bears that had been handled in Manitoba
showed significant orientation in their movements during and all were within 100 krn of the Manitoba-Ontario border
August, 0itober , and November, with mean bearings of 1660 (G. Kolenosky , personal communication).
 DEROCHER AND STIRLING: I1
Legend
a3 ADULT-ADULT
I SUBADULT-ADULT
IZ9 SUBADULT-SUBADULT
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RELOCATION DISTANCE (km)
FIG. 2. Frequency distribution of relocation distances for adult
females recaptured as adults (adult-adult), subadult females recap-
tured as adults (subadult-adult), and subadult females recaptured as
subadults (subadult-subadult).
For personal use only.
Legend
a3 ADULT-ADULT
I SUBADULT-ADULT
IZ9 SUBADULT-SUBADULT
RELOCATION DISTANCE (km)
FIG. 3. ~ r e ~ u e ndistribution
c~ of relocation distances for adult
males recaptured as adults (adult-adult), subadult males recaptured as
adults (subadult-adult), and subadult males recaptured as subadults
(subadult-subadult) .
From 1967 to 1975, 73 bears were caught along the Ontario
coast and in James Bay (Jonkel et al. 1976). Only one bear, a
2-year-old male when captured in 1968, was recaptured in our
study area, in 1984 and 1985. In contrast, 23% (17173) have
been harvested in the Belcher Islands, Ontario, or along the
Quebec coast. Less than 1% (11270) of the bears harvested in the
Belcher Islands were handled previously in our study area. A
recent and ongoing study by the Northwest Territories Wildlife
Service in the area of Southampton Island and Wager Bay (west
of Southampton Island) has resulted in over 200 captures
(M. Taylor, personal communication). To date, no exchange of
tagged animals has been documented between this population
and our study area. None of the 23 1 polar bears captured in the
southeast Baffin Island area during 1974- 1979 (Stirling et al.
1980) has been recaptured in western Hudson Bay.
'i'
RELOCATION DISTANCE (km)
FIG.4. Frequency distribution of relocation distances for cubs of the
year and yearlings with their mothers and subsequently located as
independent animals.
Fidelity
The distances between where bears were first captured and
where they were recaptured in subsequent years were similar for
all groups. These distances were also substantially shorter than
the distance between a random sample of points within the study
area which had a mean distance of 91 km (SE = 4, n = 150)
between points. Adult females were relocated a mean of 33 km
(SE = 3, n = 129) between positions (Fig. 2). Females handled
as subadults and later as adults were found a mean of 32 km (SE
= 4, n = 103) between positions (Fig. 2). Subadult female
bears, relocated as subadults, had a mean of 30 km (SE = 6, n =
47) between positions (Fig. 2). The mean distance between
relocations of adult males in different years was 32 km (SE = 3,
n = 273) (Fig. 3) and that between the locations of subadult
males and their locations as adults was 23 km (SE = 3, n = 125)
(Fig. 3). Subadult males relocated as subadults were found a
mean of 16 km (SE = 2, n = 96) between positions (Fig. 3).
The mean distance between locations of female and male cubs
of the year and yearlings with their mothers and locations as
subadults and adults was 35 km (SE = 4, n = 150) (Fig. 4).
During 1984 and 1985, areas outside the study area were
searched to assess the degree of emigration during a single
ice-free period. Of 200 independent bears captured during the
autumn of 1984 and 1985, only two were known to have left
prior to freezeup. One 3-year-old male moved 94 km to the
northwest, and the second, a 19-year-old male, moved 156 km
to the southeast. Of the independent bears handled during 1984
and 1985, 86% (1721200) were resighted within the study area.
Twenty-eight bears (14%) were not relocated. Using radio-
tracking data, at least 96% (85189) of the bears handled
remained within the study area until freezeup. Four bears with
radios were not relocated after release. There appears to be little
immigration to the study area after the ice breaks up and the
bears are ashore. Only 3 of over 400 bears handled in Ontario
and eastern Manitoba during 1984 and 1985 by the Ontario
Ministry of Natural Resources were sighted in the study area.
Winter distribution
After freezeup, polar bears from the study area were relocated
on the sea ice in western Hudson Bay (Fig. 5). In 1985 and
1986, the mean distance between the last known position of
 1400 CAN. J. ZOOL. VOL. 68, 1990
I I I I
620-
- at which body reserves are used will influence the condition of
a 60"-
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4
- 58"-
1985-86 Spring
For personal use only.
FIG.5. Locations of radio-collared polar bears on the sea ice.
bears on land in the study area in the autumn and their location
on the sea ice during the winter was 195 krn (SE = 23, n = 12).
A significant (P < 0.01) orientation of 53" (r = 0.68, n = 12)
was found in the movements of bears from the study area to the
sea ice. The late autumn locations demonstrate northward
movement along the newly formed sea ice as it rims the coast
before moving farther offshore (Fig. 5). .
Discussion
. From Tables 1 and 2 it is clear there are significant differences
in the distribution of different age and sex groups of polar bears
on land in western Hudson Bay during the ice-free period.
While the dispersion of resources is usually fundamental to the
spacing and structure of carnivore society (Macdonald 1983),
our study population during the late summer and fall is of
particular interest because of the absence of competition for
either mates or food. Nevertheless, individuals need to survive
for protracted periods on stored resources, and aspects of social
organization and environmental factors appear important.
Seals are unavailable to polar bears during the ice-free period
in Hudson Bay, and it is essential that they conserve body fat
reserves until freezeup when they can return to the sea ice to
feed. Most bears lose weight while onshore (Lunn and Stirling
1985), although the extent of the fat stores and energy expended
may differ for different segments of the population. The greatest
nutritional stress is likely experienced by females accompanied
by nursing cubs and by subadults, which are not as experienced
at catching seals and are less able to defend their kills from
dominant animals. Conversely, adult males and adult females
without cubs are usually in better condition than other bears of
other groups because of their greater ability to defend their own
seal kills and to usurp those of subordinates.
Polar bears are locomotively inefficient (Best 1982; Hurst et
al. 1982a, 1982b) and are easily affected by hyperthermia,
especially when travelling under warm ambient temperatures
(Oritsland 1970; Best 1982). Therefore, reduction of unneces-
sary movement is important for conservation of energy. The rate
an animal when it returns to the sea ice and may affect
subsequent survival and reproductive success. Through popula-
tion segregation, individuals can reduce avoidance behaviour
and confrontations and thereby reduce energy expenditure.
Distribution
Most adult males remain in the coastal areas where they are
able to minimize movements and conserve energy. We suggest
that because wind alters convective heat loss (Best 1982), cool
breezes from Hudson Bay make coastal areas preferred habitat
for adult males. Fidelity to specific coastal or inland sites could
be a function of individuals finding suitable habitat for
summering when young. Although only a small portion of the
adult males are found in the inland areas during the ice-free
period, they are similar in age and weight to males along the
coast (Derocher 1987). Use of pits and dens in the inland area,
dug down to the permafrost, could represent a viable thermoreg-
ulatory option to remaining along the coast.
Females leaving the denning area in the spring return during
the ice-free period (A. E. Derocher and I. Stirling, unpublished
data). Philopatry to inland areas by females with cubs appears to
familiarize female cubs with habitats suitable for denning,
which they will use themselves in later years. Young female
black bears ( Ursus americanus) and brown bears ( Ursus arctos)
often remain on the maternal home range and subsequently
include it, or portions of it, in their own adult home range
(Pearson 1975; Jonkel and Cowan 1971;Glenn and Miller 1980;
Rogers 1987). Although polar bears in western Hudson Bay
move much greater distances between denning areas and feeding
and breeding habitat than do either black or brown bears, females
born in the study area return there to den (A. E. Derocher and
I. Stirling, unpublished data). Hansson and Thomassen (1983)
found that polar bear cubs only spend 2 weeks near their
maternal den, after breaking out in the spring, before leaving for
the sea ice. This is probably not long enough for a cub to learn
the location of the denning area. Thus, the return of the female
to the denning area with her litter probably serves to reinforce
travel routes taken between the breeding and hunting areas and
the denning area. Family groups in higher latitude populations
have also been observed during the summer in areas used for
maternal denning (Schweinsburg 1979; Schweinsburg et al.
1982; A. E. Derocher and I. Stirling, unpublished data).
Adult male polar bears dominate subordinate conspecifics
and the threat of cannibalism can influence their distribution
(Stirling et al. 1981; Latour 1981; Kolenosky and Prevett
1983; Lunn and Stenhouse 1985; Taylor et al. 1985). Similar
behaviour is found in black and brown bears where social
interactions and the dominance of adult males also influence
distribution and movements of subordinates (Beecham 1980;
Young and Ruff 1982; McCullough 1981; Stringham 1983;
Tietje et al. 1986; Rogers 1987). On the sea ice, females with
young cubs of the year avoid habitats used by adult males
probably because of the threat of infanticide (Stirling et al.
1981). Females with yearling and 2-year-old cubs, which are as
mobile as adults, use the same habitats as adult males because of
the higher availability of ringed seals (Stirling et al. 1981). As
 DEROCHER AND STIRLING: I1
older cubs can outrun adult males (Stirling 1974), intraspecific
predation is probably unusual during the ice-free period.
Infanticide as an adaptive strategy for males may function
when a male can terminate a female's investment in offspring
fathered by other males and stimulate a rapid resumption of
receptivity and subsequent breeding opportunity (Hardy 1974;
Bertram 1975; Packer and Pusey 1984; Taylor et al. 1985). The
substantial temporal spacing between the ice-free period and the
breeding season preclude infanticide as a reproductive strategy
during the summer and autumn. Given the high energetic cost to
an adult male moving inland to pursue the more mobile family
groups, the risk of iqjury , and the small size of cubs, the benefits
of infanticide to procure food appear limited. However, females
with cubs may perceive adult males as a threat to their offspring
Can. J. Zool. Downloaded from www.nrcresearchpress.com by UNIV CALGARY on 10/09/12
even after cubs are large enough to evade adult males. Adult
males have been recorded chasing family groups during the
ice-free period (A. E. Derocher and I. Stirling, unpublished
data), and cubs are extemely nervous in meetings with unrelated
bears (Stirling 1974; Latour 1981). These observations suggest
that the avoidance of conspecifics could have a strong adaptive
value. Adult females with cubs may use the inland area partly to
avoid adult males. Given the low reproductive potential of
female polar bears and the substantial investment represented by
each cub, the loss of offspring during the autumn may seriously
affect net reproductive output.
However, intraspecific aggression does not adequately ex-
plain all of the dynamics of the spatial and temporal distribution
of family groups. If the distribution of females with cubs was
For personal use only.
only a function of the threat of cannibalism, then all females
should move to areas where the probability of encountering an
adult male is minimal. This could easily be achieved by moving
inland or by using areas where bear densities are low, such as the
region north of Churchill. The absence of family groups and
other bears in such areas indicates that factors other than
intraspecific aggression alone are involved.
Segregation of the population may also be facilitated by
greater tolerance of conspecifics of similar age and sex. Latour
(1981) reported that most of an individual's social behaviour is
directed at members of the same age and sex group. Similarly,
adult female polar bears, with and without cubs, have been
observed to interact nonaggressively (Lutzyuk 1978; Hansson
and Thomassen 1983; Lunn 1986).
Pregnant females in dens were found 10-80 km from the
coast. This .area seems preferred because of the extensive peat
banks in which dens can be dug. Most dens are located on the
banks of lakes and creeks, under clumps of scrub spruce, the
roots of which help consolidate the soil and cause additional
drifting snow to accumulate over the dens. Peat dens are cool
inside, and bears resting in them during the summer escape
harassment by insects and probably find it easier to thermoreg-
ulate than if they were exposed to direct sunlight. Pregnant
females remain in peat dens after the rest of the population has
returned to the sea ice. When sufficient snow has accumulated,
the tunnel out into a snow den which they excavate from the
snow drift that forms over the bank. The inland movement of
pregnant females is probably constrained by the presence of
continuous coniferous forest because the deep unconsolidated
snow that accumulates there is unsuitable for dens and difficult
to travel through during the spring. There is also an energetic
cost to moving far inland, both for denning in the autumn and
again in the spring, when the families return to the sea ice. At
higher latitudes, most pregnant females den within a few
kilometres of the coast (Harington 1968; Stirling et al. 1975:
Larsen 1985). Pregnant females in Hudson Bay may den inland
1401
to minimize interactions with other bears and because that is
where the best denning habitat is. Pregnant females show little
fidelity to specific sites, but do return to the same general area
(Ramsay and Stirling 1990). Repeated visits may increase an
individual bear's familiarity with the distribution of denning
habitat.
The mean weight of pregnant females in the autumn is 234 kg
(Ramsay and Stirling 1988). In comparison, the mean weight of
solitary females near the coast was approximately 50-70 kg
lighter, or approximately the same weight as females with cubs
of the year when they emerge from dens in the spring.
Implantation in ursids is delayed (Wimsatt 1963; L@n@1972)
and female black bears with insufficient body reserves may fail
to implant the blastocyst and forgo pregnancy that year (Rogers
1976, 1987). Similarly, it is likely that solitary female polar
bears with lower body weight may also fail to implant and not
receive hormonal cues associated with denning behaviour.
Using serum progesterone levels, Ramsay and Stirling (1988)
reported 13% of solitary females were not pregnant. Based on
movement information alone, it appears that up to 21% of the
solitary females may not be pregnant or fail to implant.
The distribution of subadults and the factors involved are not
as apparent as for older bears. The low position of subadults in
the dominance hierarchy and their displacement by higher
ranking bears may influence distribution and movements. In
contrast with subadult black and brown bears, which avoid adult
males and may disperse when densities of adult males are high
(Jonkel and Cowan 1971;Pearson 1975; Glenn and Miller 1980;
Young and Ruff 1982; Rogers 1987), the areas used by subadult
polar bears overlapped with adult males. The habitat along the
coast of western Hudson Bay is very open and bears can usually
see each other from some distance. Consequently, since
subadults can outrun adult males, they may perceive them as
less of a threat. Also, since there are no resources to compete
for, adult males may simply ignore subadults. Segregation of
subadults and adult females is probably due to a general
avoidance of conspecifics by adult females, whether or not they
are accompanied by cubs.
Although not fully explained, the distribution patterns de-
scribed here indicate that the selection of specific habitats is
important. There are large areas in which bears were not found,
despite extensive surveying. In general, wet areas are not used.
For example, very few bears have been seen or captured in the
fen area 5-30 km south of Churchill. Bears in the inland areas
utilize dry patches of lichen or habitat raised above the sodden
tundra. Along the coast, the drier sandy or gravel beaches are
used more frequently than the wetter areas. This may partially
account for the higher density of bears along the northern coast
of the study area, which tends to be drier than the southern
region.
Polar bears have been reported to feed sparingly while on land
and the energetic returns are thought to be low (Russell 1975;
Knudsen 1978; Lunn and Stirling 1985). However, Derocher
(1987) reported extensive foraging on Vaccinium uliginosum
and Empetrum nigrum by females and subadults in the inland
areas. The importance of this feeding and its influence on
distribution is unknown, but it may provide a high quality food
source for a brief period for some bears when alternate resources
are limited.
Movement patterns
All groups of polar bears on land during the ice-free period
moved much less than bears on the sea ice. Daily movements of
17 km (SE = 2, n = 70) were found for seven different adult
 1402 CAN. J. ZOOL. VOL. 68, 1990
females on the sea ice in the Beaufort Sea (calculated from
Amstrup 1986), which is eight to nine times the distance we
recorded during the ice-free period. The analysis of distance
moved per day by family groups, subadult females, and adult
males showed decreased movement during September. Several
bears spent many weeks at the same location. Moving to
preferred areas during July and August followed by reduced
activity until the sea ice starts to form probably functions to
conserve energy.
Solitary females move inland soon after they arrive off the sea
ice. The median distance moved per day decreased throughout
the ice-free period and is indicative of pregnant females finding
suitable denning sites and remaining there. By late September,
most pregnant bears have moved inland and occupied dens. Few
Can. J. Zool. Downloaded from www.nrcresearchpress.com by UNIV CALGARY on 10/09/12
pregnant females were seen outside their dens after the middle
of October.
Many bears arrive from the sea ice in the northeast comer of
the study area and subsequently move south along the coast or
inland. In October and November, the direction of movement is
reversed as bears move north along the coast and begin to leave
the inland area. By November, many bears have reached the
north coast of the study area where they await the formation of
the sea ice. The heading that family groups took as they moved
inland or back to the coast was approximately the same as, or
180" to, the orientation of 39" that Ramsay and Andriashek
(1986) reported for bears leaving the denning area in the spring.
They speculated that the observed orientation may have been a
function of the bears moving to an area of active sea ice with
For personal use only.
good hunting identified by Stirling et al. (1977). The use of the
area to the northeast of Cape Churchill was supported by the
radio locations obtained from bears on the sea ice.
Fidelity
Distribution and movement patterns are influenced by fidelity
to the study area and specific sites within the study area. Fidelity
patterns were similar for all groups of bears. Polar bears born in
the study area are philopatric, and adults and subadults show
long-term fidelity to the study area. Findings of site fidelity in
polar bears are not new. Polar bears on the sea ice in the spring
return to the same feeding areas over many years (Stirling et al.
1980, 1984; Lentfer 1983; Schweinsburg et al. 1982). Immigra-
tion into the study area is limited, and the exchange between the
Ontario and Manitoba populations results from bears making
periodic visits outside of their normal summering areas. Prevett
and Kolenosky (1982) suggested that there was an annual
exchange of bears between Manitoba and Ontario. However,
they only considered the portion of Manitoba near the Ontario
border. Our study clearly indicates that extensive exchange of
bears between our study area in Manitoba and Ontario does not
occur within or between annual ice-free periods.
Recovery of tags from bears captured in the study area
decreased as the distance from it increased. The limits of the
western Hudson Bay population appear to lie between Rankin
Inlet and Chesterfield Inlet to the north and between the mouth
of the Nelson River and the Manitoba-Ontario border to the
south. Given that harvest along the Keewatin coast is centred on
subadults, which have a lower tag density than adults in the
population, the decrease in tag returns moving north probably
underestimates the proportion of the study population being
harvested. The large proportion of bears known to have recently
been in the study area and being harvested farther north is a
function of the northward migration along the newly formed sea
ice. The analysis of hunter kill information is confounded
because dispersal and regular seasonal movements cannot be
differentiated. However, it is clear that while many of the bears
from our study area spend the winter and spring in western
Hudson Bay, some bears, predominantly young males, do range
throughout Hudson Bay.
The conditions during the ice-free period are such that the
benefits of dispersal by subadults may be limited and any
mechanisms that induce dispersal may not function at this time.
The benefits of philopatry and site-specific fidelity are probably
similar to those derived by other animals from home ranges and
territories: familiarity with conspecifics and the distribution of
resources. Philopatry may also relate to navigational patterns
learned from mothers. The manner in which polar bears
navigate is unknown, but it is essential that they return to land as
the sea ice melts.
The sea ice near the study area may be productive for hunting
seals and the study area represents the energetically most
efficient place to wait until the sea ice reforms. There is little
known about the distribution of seals in the Hudson Bay area,
but the density of ringed seals is variable (Smith 1975) and
probably determines the distribution of polar bears. The separa-
tion of the subpopulations that spend the ice-free period in
Ontario and Manitoba may be influenced by hunting in different
areas in the winter and spring.
In summary, it appears that the distribution and movement
patterns of polar bears during the ice-free period are influenced
by social and energetic factors interacting with learned beha-
viour and site fidelity. The spatial and temporal segregation of
polar bears during the ice-free period is markedly different from
that found in other ursids and permits this normally solitary
species to pass a noncompetitive season, at high densities, while
conserving energy stores.
Acknowledgements
Dennis Andriashek provided technical support and field
assistance that was crucial to the success of the field studies.
Wendy Calvert provided invaluable assistance in the laboratory
overseeing data management. We wish to thank M. Cattet, M.
Gillespie, R. Hansson, D. Jacobs, S. Kearney, L. Knutsen,
S. Miller, J. Miller, K. Pontus, M. Ramsay M. Shoesmith,
C. Spencer, I. Thorleifson, R. Tease, and L. Voissey for their
assistance in this study. Two anonymous reviewers provided
constructive criticism of an earlier draft of this paper. Funding
was provided by the Boreal Institute for Northern Studies,
the Canadian Wildlife Service, the Manitoba Department of
Natural Resources, the Natural Sciences and Engineering
Research Council of Canada, the Northwest Territories Depart-
ment of Renewable Resources, the Folar Continental Shelf
Project, the University of Alberta, and World Wildlife Fund
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