Glacier
Habitat Relationships of Grizzly Bears in Glacier National Park, Montana
C. J. Martinka
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Glacier National Park
West Glacier, Montana 59936
HABITAT RELATIONSHIPS OF GRIZZLY
BEARS IN GLACIER NATIONAL PARK, MONTANA
National Park Service Progress Report, 1972
For use by National Park Service and cooperating agencies
Prepared by
C. J. Martinka, Research Biologist
Office of Natural Science Studies
Glacier National Park
March 1, 1972
HABITAT RELATIONSHIPS OF GRIZZLY
BEARS IN GLACIER NATIONAL PARK, MONTANA
C. J. Martinka, Glacier National Park, Montana
Abstract: Habitat use and food preferences of
grizzly bears (Ursus arctos) were studied in Glacier National
Park fron 1967 through 1971. Park physiolography and vegetation were
described. Analysis of 353 sightings showed that grizzlies frequented
low elevation coniferous forest, grassland, and snowslide habitat types
in spring; higher elevation coniferous forest, shrub, and alpine types
in summer and early fall; lower elevation coniferous forests in late
fall. Analysis of 340 scats showed that the diet was predominated by
ungulates and unidentified grasses in early spring; horsetail
(Equisetem app.) and cow parsnip (Heracleum lanatum) in
late spring and early summer; huckleberry (Vaccineum spp.),
serviceberry (Amelanchier spp.), hawthorne (Crataegus
spp.) and mountain ash (Sorbus scopulina) in late summer and
early fall; grasses, roots and ungulates in late fall. Columbian ground
squirrel (Citellus columbianus) supplemented the diet during all
seasons. Relationships between grizzlies and Park ecosystems were
discussed.
TABLE OF CONTENTS
Abstract
Study Area
Methods
Results
Food Habits
Discussion
Literature Cited
Grizzly bears originally inhabited an extensive area
of western North America (Storer and Trevis 1955: 12-14). Historical
accounts generally characterized the grizzly as an omnivore which
utilized a wide variety of habitats and food items. This evaluation has
more recently been supported by studies in Canada and Alaska (Mundy
1963, Murie 1944, Murie 1961, and others). Extensive population declines
in the late 1800s, precluded similar studies for most historic ranges
south of Canada.
This study was conducted in Glacier National Park
from 1967 through 1971 to obtain quantitative data on status, dynamics,
habitat-relationships and management of the resident grizzly bear
population. Results relating to population characteristics and
management to protect park visitors have been reported elsewhere
(Martinka 1971a, 1971b). Data and evaluations concerning habitat use and
food preferences of the grizzly population are presented in this
report.
The author expresses appreciation to R. Gildart and
D. Shea for field assistance. Technical assistance in analyzing scats
was given by K. R. Greer, Montana Fish and Game Department. Cooperation
of Glacier National Park personnel in reporting grizzly bear sightings
made the study possible. The report is a contribution from National Park
Service Natural Science Project GLAC-N-16.
STUDY AREA
Glacier National Park includes 1583 square miles of
mountainous terrain in northwestern Montana. Topography is characterized
by a central axis of precipitous peaks and ridges with streams radiating
from the park as headwaters of the Saskatchewan, Missouri, and Columbia
River systems. Prominent geologic features include remnant glaciers,
cirques, moraines and lakes. Extensive talus slopes are common east of
the Continental Divide. Elevations vary from 3,100 to 10,500 feet.
Park climate is classified as continental with
Pacific maritime modifications on western slopes (Dightman 1967).
Weather records for Summit (5,213 feet) showed a mean annual
precipitation of 38.29 inches from 1937-66; about 60 percent fell as
snow from November through April. Annual snowfall of 251 inches
frequently resulted in accumulations exceeding 60 inches. Mean monthly
temperature was 35.9F with extremes 15.3F (January) and 59.9F (July).
Records of measurements at other elevations show substantial variations
in precipitation and temperatures when compared to these means.
Park vegetation is predominated by extensive zones of
alpine and coniferous forest communities. A discontinuous zone of
grassland communities occurs in peripheral areas. Considerations of
floral composition, physiographic sites, and successional status
permitted identification of five major habitat types within these zones.
Descriptions were aided by discussions in Habeck (1970).
Alpine: This type occurred generally above
6,000 feet. Meadow, fellfield and ledge communities were characterized
by low-growing plants including varileaf cinquefoil (Potentilla
diversifoila), snow willow (Salix nivalis), and sedge
(Carex tolmieri). Alpine fir (Abies lasiocarpa) krummholz
occupied a variety of physiographic sites within this type. Broad
ecotones frequently formed with coniferous forest and snowslide types of
lower elevations.
Coniferous Forest: This type occupied most
sites below 6,000 feet. Climax forests were dominated by alpine fir,
Engelman spruce (Picea engelmannii) and Douglas fir
(Pseudotsuga menziesii). Seral forests were composed of Douglas
fir, lodgepole pine (Pinus contorta) and western larch (Larix
occidentalis). A variety of shrubs and herbs occurred in field
strata of these types.
Snowslide: This type occurred frequently on
more precipitous slopes within the coniferous forest zone. Shrubs were
characteristic dominants with alder (Alnus sinuata) and willow
(Salix spp.) abundant on most sites. Mountain maple (Acer
glabrum), mountain ash and serviceberry were common in drier
situations.
Shrub: This type commonly occupied sites
within the coniferous forest zone for variable periods following
wildfire. Willow, western serviceberry, mountain maple, redstem
ceanothus (Ceanothus sanguineus) were dominant on drier sitess.
Huckleberry and smooth memziesia (Menziesia ferruginea)
characterized higher elevations.
Grasslands: This type occurred on alluvial
fans, colluvial slopes, windblown ridges, or as a seral community
following wildfire in dry coniferous forest types. Rough fescue
(Festuca scabrella), bluebunch fescue (F. idahoensis),
bluebunch wheatgrass (Agropyron spicatum) and sedges (Carex
spp.) were characteristic dominants. Bluegresses (Poa spp.)
and timothy (Phleum pratense) dominated moist depressions as well
as occasional meadows. A number of minor seral types was also present,
but not generally utilized by grizzlies.
METHODS
Data on habitat use were obtained from sightings of
grizzlies by the author, park personnel, and other qualified observers.
Bears were observed while traveling park trails on foot or horseback,
from a vehicle along roads, or during aerial flights. Date, location,
numbers, descriptions, habitat type and elevation were recorded for each
sighting. Descriptions included size, color and distinguishing features
and were used to exclude possible duplicate sightings within 30-day
periods. Each sighting was considered as a unit irregardless of number
or group composition.
Seasonal food habits were studied from analyses of
grizzly scats. These were collected along roads and trails as
opportunity permitted. The presence of tracks or diggings, a minimum
diameter of two inches, or both, were used as criteria to distinguish
grizzly from black bear scats. Each scat was placed in a plastic bag,
identified by number, and frozen upon return from the field. Laboratory
analysis for food content was facilitated by emersion in water and
separation in a pan. The percentage of individual food items or
categories present was visually estimated. Frequency of occurrence and
means of the aggregated volumes were computed for each season.
Scat analyses were complemented by evaluation of
sites where grizzlies had recently secured food by digging. Habitat type
was recorded for each site and food items determined from excavation
characteristics. Diggings for herbaceous items were relatively shallow
and showed evidence of bitten or chewed roots. In contrast, diggings for
rodents commonly exposed portions of burrow systems.
RESULTS
Use of Habitat Types
Analysis of 353 sightings revealed that each of the
major habitat types in Glacier National Park was utilized by grizzlies
(Table 1). Trends which occurred in seasonal use corresponded, in
general, with shifts in the altitudinal distribution of bears (Table 2).
These data, when considered with sighting locations, reflected the
general distribution of grizzlies throughout the park.
Table 1. Seasonal use of the different habitats by
grizzly bears in >Glacier National Park as determined from 353
sightings, 1967 through 1971.
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Season |
No. of Sightings |
Percent Observed
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Alpine |
Coniferous Forest |
Snowslide |
Shrub |
Grassland |
|
Spring (April June) |
165 |
7 |
46 |
21 |
10 |
16 |
Sumner (July Aug.) |
130 |
41 |
36 |
9 |
11 |
3 |
Fall (Sept. Nov.) |
58 |
29 |
35 |
12 |
19 |
5 |
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Yearlong |
353 |
23 |
41 |
15 |
12 |
9 |
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Table 2. Seasonal altitudinal distribution of grizzly
bears as determined from sightings in Glacier National Park, 1967
through 1971. Sample sizes as shown in Table 1.
|
Season |
Percent Observed
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3100 4000' |
4100 5000' |
5100 6000' |
Above 6000' |
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Spring (Apr. June) |
53 |
24 |
17 |
6 |
Sumner (July Aug.) |
20 |
25 |
14 |
41 |
Fall (Sept. Oct.) |
27 |
14 |
26 |
33 |
|
Yearlong |
37 |
23 |
17 |
23 |
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Spring: Grizzlies were most frequently
observed on lowland habitats. Coniferous forests were the most important
type with maximum use occurring along stream bottoms or in the vicinity
of grasslands. Extensive tracts of upland forests were utilized less
frequently. Snowslide and grassland types were used to a greater extent
during spring then in other seasons. Occurrence of grizzlies in shrub
types was most often recorded at low elevations.
Summer: A segment of the grizzly population
moved to higher elevation habitats. Increased use of the alpine type
coincided with decreased use of other types, particularly snowslide and
grassland. Alpine meadow and krummholz communities were most important.
Summer use of coniferous forests occurred at all elevations but a
preference for higher elevations was indicated for the shrub type.
Fall: Grizzlies continued to frequent high
elevation habitats in September but moved lower following frosts and
early snows. Decreased use of alpine types correspond with greater use
of shrub and snowslide types. Coniferous forests remained important but
grasslands continued to be of minor significance. The relatively few
observations made in late October and November suggested nearly
exclusive use of the coniferous forest type.
Winter: Bears were generally inactive although
occasional sightings of individual bears or tracks were recorded in
December and March. Limited data suggested that denning and hibernation
occurred in coniferous forests or higher elevation shrub types.
FOOD HABITS
Analysis of 340 scats showed that the grizzly diet
included a variety of plant and animal foods (Table 3). Records from 74
sites revealed that digging supplemented the diet with roots, bulbs and
small mammals (Table 4). Relatively few foods contributed significantly
to the diet during each seasonal period with the exception of Columbian
ground squirrel. Trends in the seasonal use of certain foods appeared
related, in part, to changes in habitat use.
Table 3. Seasonal food habits of grizzlies in Glacier
National Park as determined from analyses of 340 scats collected from
1967 through 1971. Values are expressed as percentages.
|
Food Item1/ |
Spring (Apr.June)
|
Summer (JulyAug.)
|
Fall (Sept.Oct.)
|
Vol.2/ | Freq.3/ |
Vol. | Freq. |
Vol. | Freq. |
|
Herbs and Roots |
Carex spp. | Tr | 3 |
6 | 23 | 3 | 10 |
Equisetem spp. | 9 | 35 |
4 | 17 | Tr. | 3 |
Gramineae | 43 | 78 |
16 | 52 | 16 | 46 |
Heracleum lanatum | 30 | 52 |
42 | 61 | 1 | 9 |
Misc. Herbs | 7 | 22 |
5 | 25 | 14 | 26 |
Roots and Bulbs | Tr | 3 |
4 | 20 | 6 | 10 |
Total | 89 | 96 |
77 | 82 | 40 | 60 |
Fruits |
Amelanchier spp. | | |
4 | 1 | 7 | 21 |
Crataegus spp. | | |
Tr | 1 | 6 | 8 |
Sorbus scopulina | | |
| | 5 | 11 |
Vaccineun spp. | 1 | 1 |
14 | 26 | 35 | 63 |
Misc. Fruits | Tr | 11 |
2 | 8 | 4 | 16 |
Total | 1 | 12 |
20 | 32 | 57 | 78 |
Mammals & Insects |
Cervidae | 9 | 15 |
1 | 2 | 2 | 3 |
Citellus columbianus | 1 | 8 |
Tr | 7 | 1 | 9 |
Misc, mammals | Tr | 11 |
Tr | 2 | Tr | 6 |
Insecta | Tr | 29 |
2 | 26 | Tr | 4 |
Total | 10 | 50 |
3 | 36 | 3 | 32 |
Sample Size | 139 |
99 |
102 |
1/ Items comprising less than 5 percent volume
and 25 percent frequency during each season are included in the
miscellaneous category.
2/ Average of the aggregated percentages for
that season. Tr denotes less than 0.5 percent.
3/ Percentage of scats in which item occurred
during that season.
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Table 4. Habitat types and food items at 74 grizzly
bear digging sites inspected from 1967 through 1970 in Glacier National
Park.
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Habitat Type |
Sites
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Food Item Frequency (%)
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No. | % |
Roots | Mammals | Both |
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Alpine |
33 |
45 |
40 |
36 |
24 |
Snowslide |
25 |
34 |
76 |
12 |
12 |
Grassland |
9 |
12 |
33 |
67 |
0 |
Shrub |
4 |
5 |
25 |
75 |
0 |
Coniferous Forest |
3 |
4 |
100 |
0 |
0 |
|
Totals |
4 |
100 |
53 |
32 |
15 |
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Spring: Herbaceous foods predominated the
diet. Unidentified grasses were the most important category with maximum
use occurring during early spring. Decreased use of grasses during late
May and June coincided with more frequent use of horsetail and cow
parsnip.
Ungulates were eaten more frequently in spring than
during other seasons. Greatest use was recorded in April when elk
(Cervus canadensis) and deer (Odoccileus spp.) bulked 54
percent of 9 scats examined. Additional species recorded in May and June
scats included moose (Alces alces) and mountain goat (Oreamnos
americanus). Most of the insects recorded in spring scats were
Coleoptera; a large proportion appeared to occur as incidental adjuncts
to decomposition.
Summer: Herbaceous items were supplemented
with fruits, roots and insects. Decreased use of grasses and horsetail
coincided with more frequent use of sedges and cow parsnip as compared
to spring. In turn, huckleberry and serviceberry progressively replaced
herbs as fruits ripened. Roots and bulbs were common diet constituents;
glacier lily (Erythronium grandiflorum) was the most important
species. Insects, primarily Formicidae, provided more bulk to the diet
than during spring.
Fall: Fruits occurred more frequently than
herbaceous items. Huckleberry was the principle fruit eaten;
serviceberry, hawthorne and mountain ash were important supplemental
species. Use of fruits appeared to decline sharply in late October.
Coincidently, unidentified roots, grasses and ungulates were the
principle components of the few scats examined.
DISCUSSION
Results of this study showed that grizzly bears
functioned as ommivores within Glacier National Park ecosystems. A
primary herbivorous role was supplemented with scavenging or predation
on certain mammals. These interactions resulted in occupancy of a
relatively distinct niche by grizzlies.
Ungulate carrion was an important food source for
grizzlies during early spring. Most use appeared to be of carcasses
which were not utilized by other secondary consumers following natural
winter deaths. Accidental deaths of wintering ungulates in snowslides
provided an additional source which became progresively available during
May and June. Spring distribution of grizzlies seemed closely related to
these food sources.
Direct predation on ungulates supplemented scavenging
activities of grizzlies during spring. Scat contents suggested that
newborn young were occasionally killed following birth. Data for other
age classes were unconclusive although one death of an adult female
moose was recorded. Cole (1972) reported that grizzlies frequently
preyed upon predisposed elk in Yellowstone National Park following
winter conditions which extended into May.
Grizzlies and ungulates interacted within
ecologically complete habitats (Cole 1971) in most areas of Glacier.
Natural regulation and relative numerical stability were characteristic
of these ungulate populations. Grizzlies benefited from both
stress-related and accidental deaths which field observations suggested
were largely independent of ungulate density. Accordingly, grizzlies did
not appear to be important influents on ungulate population levels. Cole
(1972) concluded that grizzly predation, scavenging, or both, was a
nonessential but assisting adjunct to natural processes that regulated
an elk population.
Spring distribution of grizzlies was also influenced
by the location of certain herbs which were characteristically obtained
from moist seres and bottomlands within the coniferous forest zone. Wet
meadows containing bluegrasses were utilized more frequently for grazing
than the drier bunchgrass types. Horsetail and cow parsnip were commonly
found in moist seeps and bottomlands within the coniferous forest type.
As snow melted, snowslides were an abundant source of cow parsnip,
particularly on north and east exposures. Use of cow parsnip during
early summer occurred at progressively higher elevations, but declined
sharply following maturation and the availability of fruits.
Coniferous forests were an inportant source of
huckleberry and other fruits. Early summer use by grizzlies occurred at
all elevations but use progressed toward the higher elevations in
August. High elevation were frequented until frosts, early snow, or
both, reduced availability of palatable fruits. Dispersal of bears to
lower elevation coniferous forests usually followed.
Wildfire was an important natural factor which
affected the abundance and productivity of favored fruit producing
plants, within the coniferous forest zone. Field observations showed
both greater and more consistant production of huckleberries on burned
sites than in mature forests. These relationships were most important at
high elevations where a slow rate of secondary succession following
wildfire permitted high production of huckleberries to continue for a
relatively long interval. Grizzlies appeared responsive and reached
densities as high as 1 per square mile on these sites during late
summer. Jonkel (1971) reported similar observations for black bears
(Ursus americanus).
Annual variations in spring weather apparently caused
year to year fluctuations in fruit production by huckleberry. During
this study, crop failures varied from local to parkwide in four of five
years. Fluctuations of similar magnitude were not observed among other
fruit-producing shrubs. Limited data suggested that these provided
important alternative foods for grizzlies during years or in location of
poor huckleberry production.
A distinct relationship occurred between grizzlies
and alpine habitats during the snow-free period from July through
September. Digging for foods was a characteristic foraging activity
which resulted in extensive disturbance to alpine soils in many areas.
Similar relationships have been described for populations in Canada
(Mundy 1963) and Alaska (Murie 1944).
Black bears were sympatric with grizzlies in Glacier
National Park. Maximum association occurred within forest and shrub
habitat types of the coniferous forest zone. Food preferences of black
bears (Tisch 1961) are similar to those of grizzlies in forest habitats
but interspecific relationships were not apparent. In contrast,
observations on the relatively open shrub types indicated that wide
spacing occurred between the species or that black bears were not
present. Barnes and Bray (1964) described dominance interactions between
the species which were quite complex in Yellowstone National Park.
Important differences occurred between black and
grizzly bears in the use of certain foods and habitats. Black bears were
seldom seen in association with wintering ungulates or as predators on
rodents. In addition, few black bears were observed on grassland or
alpine habitat types. Widespread occurrence of ungulate populations as
well as extensive alpine habitats possibly permitted relatively high
densities of grizzlies to inhabit the park, as compared to other areas
(Martinka 1971a, b). Jonkel (1971) reported that grizzlies seemed to
prefer more open habitats than black bears.
Most terrestrial ecosystems within Glacier are
essentially complete and intact. As an integral component, the grizzly
population appears to be at habitat carrying capacity, relatively
stable, and regulated by social interaction (Martinka 1971a, b).
Developmental intrusions by humans have occurred but human activity and
unnatural foods seemed more important than limited physical features in
affecting grizzly habitat relationships. Management to reduce
interactions between grizzlies and humans resulting from these factors
has been implemented (Martinka 1971a). Potential responses to further
human intrusions are not known.
LITERATURE CITED
Barnes, V. G. and O.E. Bray, 1967. Population
Characteristics and Activities of Black Bears in Yellowstone National
Park. Colorado Coop. Wildl. Res. Unit, Colorado State Univ. 199pp.
Clark, W. F. 1957. Seasonal Food Habits of the Kodiak
Bear. Trans. N. Am. Wildl. Conf. 22: 145-151.
Cole, G. F. 1971. An Ecological Rationale for the
Natural or Artificial Regulation of Native Ungulates in Parks. Trans. N.
A. Wildl. Conf. 36: 417-425.
______. 1972. Grizzly-elk Relationships in
Yellowstone National Park. J. Wildl. Mgmt. 36: (In press).
Dightman, R. A. 1967. Climate of Glacier National
Park, Montana. Glacier Natural History Association. Bull. No. 7.
9pp.
Habeck, J. R. 1970. The Vegetation of Glacier
National Park, Montana. Special Report. 132pp.
Jonkel, C. J. and I. M. Cowan. 1971. The Black Bear
in the Spruce-fir Forest. Wildl. Monog. No. 27. 57pp.
Martinka, C. J. 1971a. Status and Management of
Grizzly Bears in Glacier National Park, Montana. Trans. N. A. Wildl.
Conf. 36: 312-322.
______. 1971b. Population Characteristics of Grizzly
Bears in Glacier National Park. Symposium on Research in the National
Parks. (In press).
Mundy, K. R. D. 1963. Ecology of the Grizzly Bear
(Ursus arctos) in Glacier National Park, British Columbia.
Unpublished M.S. Thesis, University of Alberta, Edmonton. 103pp.
Murie, A. 1944. The Wolves of Mount McKinley. Fauna
of National Parks. No. 5. 238pp.
______. 1961. A Naturalist in Alaska. The Devin-Adair
Co., New York. 302pp.
Stover, T. I. and L. P. Trevis, Jr., 1955. California
Grizzly. University of California Press, Berkeley. 355pp.
Tisch, E. L. 1961. Seasonal Food Habits of the Black
Bear in the Whitefish Range of Northwestern Montana. Unpublished M. S.
Thesis, Univ. of Montana, Missoula. 108pp.
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