Behavior and environmental selection by elk (Cervus canadensis nelsoni) during summer and fall in the
first and second Yellow Mule drainages, Madison County, Montana
by Gayle Lynne Joslin
A thesis submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE
in Zoology
Montana State University
© Copyright by Gayle Lynne Joslin (1975)
Abstract:
A study was conducted in the First and Second Yellow Mule drainages of southcentral Montana during
the summer and early fall of 1973 and 1974 to investigate ecological aspects of elk (Cervus canadensis
nelsoni). The study area was divided into park, broken park, and timber cover types covering 31, 13,
and 56 percent of the area, respectively. Six habitat types were delineated on the timbered regions.
Large mixed sex bull-cow groups observed in late July dispersed to form small single sex bull groups
and cow groups which increased in size to late August. Environmental features recorded to compare
areas selected by each elk group type were cover type, habitat type, timber type, distance to water,
distance to the next closest cover type, downfall, sight distance, location, topography, elevation, aspect,
and slope. Significant differences existed between preferences of group types for topography and
aspect (P<.05) and location.(P<.01). Selection of other environmental features were not statistically
different. Bulls abandoned their environmental preferences in late summer to join cow groups in their
preferred habitat.
Canopy cover of tree and shrub species, general ground cover, and the three dominant species in each
of nine 2X5 decimeter frames were recorded at 21 feeding and 28 bedding sites. Ground cover at both
types of sites was similar. Each group was observed in one of five basic activity patterns and in one of
three response states. Elk in the restless response state were significantly more common during weather
phase one (Landsberg, 1969). The "head jerk" and "humped back" postures and other dominant and
subordinate gestures were observed. Rut-related activities began with velvet shedding, followed by
herding, wallowing, bugling, and sparring.
Dry weight standing crops and percentage moisture contents were determined for forbs and grasses
which were clipped every two weeks from elk feeding sites and from established timber and meadow
plots at three elevations. Forbs at elk feeding sites had significantly greater moisture contents than
forbs in all meadow and most timber sites; standing crops of forbs at feeding sites were also
significantly greater than forbs at all timber and most meadow sites. Percentage moisture content of
forbs fell below that of grasses during October, which was also the only time that moisture content of
grasses at established plots exceeded that of grasses at feeding sites. However, elk moved into the
timber in September, coincident with the rut, and reappeared in high open parks in October. Based on
relative percentage moisture content, movement into the timber solely for succulent vegetation would
not be founded; therefore, during this study it appeared that security requirements of elk were involved
in this fall movement into the timber. STATEMENT OF PERMISSION TO COPY
In presenting this thesis in partial fulfillment of the
•requirements for an advanced degree at Montana State University,
I agree that the Library shall make it freely available for
inspection.
I further agree that permission for extensive copying
of this thesis for scholarly purposes may be granted by my major
professor, or, in his absence, by the Director of Libraries.
It
is understood that any copying or publication of this thesis for
financial gain shall not be allowed without my written permission.
Signature
Date
BEHAVIOR AND ENVIRONMENTAL SELECTION BY ELK (CERVUS CANADENSIS
NELSONI) DURING SUMMER M D FALL IN THE FIRST M D SECOND
YELLOW MULE DRAINAGES, MADISON COUNTY, M O N T M A
by
GAYLE LYNNE JOSLIN
A thesis submitted in partial fulfillment
of the requirements for the degree
of
• MASTER OF SCIENCE
in
Zoology.
Approved:
Chairman, Examining Committee
ad. Major Department
Graduate Aean
MONTANA STATE UNIVERSITY
Bozeman,. Montana
August, 1975■
-iiiACKNOWLEDGMENT
The author wishes to express her appreciation to the following
people for their contributions to this study:
Dr. Robert E. Moore,
Montana State University, who directed this study and aided in
preparation of the manuscript; Dr. Harold D. Picton and Dr. Theodore
W. Weaver, Montana State University, for critical review of the
manuscript and for assistance in project planning; Mr. and Mrs.
John Cada, Porcupine Game Range, Montana Fish and Game Department,
for hospitality and cooperation; Mr. Terry N. Lonner, Game Research
Biologist, Montana Fish and Game Department, for helpful suggestions
and assistance; personnel of the Squaw Creek Ranger District,
Gallatin National Forest, for use of the Yellow Mule Ranger Station;
Dr. John H. Rumely, curator of the Montana State University herbarium
for aid in verification of plant specimens; Mr. and Mrs. Harry L.
Joslin for encouragement and support.
During this study the author
was employed as a Graduate Teaching Assistant at Montana State
University.
iv
TABLE OF CONTENTS
Page
VITA
.............................. .. . . . . . . . .
.
ACKNOWLEDGMENT..........................................
TABLE OF CONTENTS
LIST OF FIGURES
iii
.............................-...............
LIST OF T A B L E S .......... ...............................
ii
....
iv
vi
'. . ...........■ . . ......................... .. viii
ABSTRACT .......................................................... ix
INTRODUCTION.................................. ................
I
METHODS
3
......................................................
THE STUDY AREA ........................................
8
Abies lasiocarpa(T?inus aVbiaauiis)/Vaeainium
sooipaviym habitatt y p e .......................
Abies Zasiooavpa/Vacoinivm soopavium
12
.habitat t y p e ................................
15
Abies Zasiooavpa/Linnaea borealis
habitat t y p e ................................
16
Abies lasioodvpa/Calamagvostis vubesoens
habitat type
..............
18
Abies Zmiooavpa/Caldmagvpstis.canadensis
habitat t y p e ................
20
Abies Zasiocavpa/GaZivm tviflovim
habitat t y p e ............
RESULTS
20
.......................................................
Elk Group Characteristics.......................... .. . .
Group Size and Composition..................
Sites Used by E l k ............................
Vegetation Analysis ofElk Feeding and Bedding Sites .
Elk Activities
. . . • ...................................
General Daily Activities ........................
. .
23
23
24
26
32
36
36
V
TABLE OF CONTENTS
(Continued)
Pege
Social Behavior . . . . . . . . . . ................
' 37
Rut—Related Activities . . . . / . . . .. . . . .. . .. .' • 40
Vegetation Phenology . . . . . . . . . . . . . . . . .
42
DISCUSSION............ . .......................................
49
APPENDIX . ............. ..................................... .
55
LITERATURE CITED
63
vi
LIST OF TABLES
Table
1.
2.
3.
4.
5.
Page
PERCENT OF ELK GROUPS AND MEAN NUMBER OF ELK PER.
GROUP OBSERVED' IN SEVEN TWO WEEK PERIODS DURING
1973 AND 1974 . . . ................. . . . . . . . . . .
25
PERCENT OF ELK GROUPS.AND MEAN NUMBER OF ELK OBSERVED PER GROUP DURING SIX WEATHER PHASES
. . ...............
26
CHI SQUARE VALUES CALCULATED FROM COMPARISONS OF
THREE ELK GROUP TYPES FOR EACH OF TWELVE '
ENVIRONMENTAL. FEATURES .......... . . . . . . . . . . .
27
PERCENT OF ELK GROUPS AND MEAN NUMBER OF ELK PER
GROUP IN THREE GROUP TYPES OBSERVED ON SIX
TOPOGRAPHICAL AREAS ................................... .
28
PERCENT OF ELK GROUPS AND MEAN NUMBER OF ELK PER GROUP ■
IN THREE GROUP TYPES OBSERVED ON EIGHT ASPECT
CLASSES
............ .. . . . . . .
28
6.
PERCENT OF ELK GROUPS AND MEAN NUMBER OF ELK PER GROUP
IN THREE GROUP TYPES OBSERVED IN TWO LOCATIONS .... . ... 28
7.
PERCENTAGES OF 170 ELK GROUPS RELATED TO VARIOUS ,
ENVIRONMENTAL FEATURES, MEASURED DURING 1973 AND 1974 • •
29
PERCENTAGES OF 83 ELK BEDDING SITES AS RELATED TO
VARIOUS ENVIRONMENTAL FEATURES, MEASURED DURING 1974
31
8.
9.
,. .
MEAN PERCENT CANOPY COVERAGE OF GENERAL VEGETATION
GROUPS AT TWENTY-ONE FEEDING AND TWENTY-EIGHT BEDDING
SITES AS DETERMINED BY EXAMINATION OF NlNE 2 X 5
DECIMETER FRAMES AT. EACH S I T E ................ ..
10. VEGETATION ASSOCIATED WITH TWENTY-ONE FEEDING SITES
■ EXAMINED OVER A FOURTEEN WEEK PERIOD WITH REGARD TO
CONSTANCY, OVERALL DOMINANCE VALUES, AND. PRESENCE, .
MEASURED IN NINE 2 X 5 DECIMETER FRAMES WITHIN A,
CIRCULAR PLOT AT EACH S I T E ........ ....................
32
33
vii
LIST OF TABLES
(Continued)
Table
11.
12.
13.
14..
15.
Page
VEGETATION ASSOCIATED WITH'TWENTY-EIGHT BEDDING
SITES EXAMINED OVER A FOURTEEN WEEK PERIOD WITH
REGARD TO CONSTANCY, OVERALL DOMINANCE VALUES, AND
PRESENCE, MEASURED IN NINE 2 X 5 DECIMETER FRAMES
WITHIN A CIRCULAR PLOT AT EACH S I T E .......... ..............
34
COMBINATIONS OF PRESSURE, CLOUD COVER, TEMPERATURE
AND WIND TO PRODUCE SIX WEATHER PHASES .....................
56
CONSTANCY AND OVERALL DOMINANCE
ASSOCIATED WITH 189 AND 253 2 X
MEASURED AT FEEDING AND BEDDING
OVER A FOURTEEN WEEK PERIOD
.
VALUES OF SPECIES
5 DECIMETER FRAMES
SITES, RESPECTIVELY,
. . . . . . . . . . . . ........
57
STANDING CROP AND PERCENTAGE MOISTURE CONTENT OF
VEGETATION ON TWENTY-ONE ELK FEEDING SITES ................
61
STANDING CROP AND PERCENTAGE MOISTURE CONTENT OF
VEGETATION ON SIX ESTABLISHED PLOTS AT THREE
ELEVATIONS . . . . . ......................................
62
viii
LIST OF FIGURES
Figure
Page
1.
Map of study area showing cover types ............
2.
Terraced basin of the upper Second Yellow
Mule Creek ................................
9
11
3.
Broken park cover t y p e .................................... 11
4.
Map of study area showing habitat types and
timber types ............................................
13
Abiea laaioearpatPinua albioaulis)/Vacoinivm
aaoparium habitat type ....................
14
5.
6.
7.
. . . . . . .
Abies lasioearpa/Vaeeinivm aeopariim habitat type,
the Vaecinivm seopaTium p h a s e ........................ ■ . .
19
Abies lasioearpa/Catamagrostis rubeseens
habitat t y p e .............................................. 19
8.
9.
Abies lasioearpa/Calamagrostis canadensis "stringer"
surrounded by the Abies lasioearpa(Finns albieautis)/
Vaeeininm seoparium habitat type ........................
21
Duration of rut-related activities
41
. . . . .
............
10.
Grams dry weight standing crop of forbs (left) and
grasses (right) at twenty-one elk feeding sites arid six
established plots clipped every two weeks from mid July
to mid O c t o b e r ............................................ 46
11.
Percentage moisture content of forbs (left) andgrasses
(right) at 21 elk feeding sites, and six established
plots clipped every two weeks from mid July to mid
O c t o b e r ......................................
47
ix
ABSTRACT
A study was conducted in the First and Second Yellow Mule drain­
ages of southcentral Montana during the summer and early fall of 1973
and 1974 to investigate ecological aspects of elk (Cervus canadensis
nelsoni)* The study area was divided into park, broken park, and
timber cover types covering 31, 13, and 56 percent of the area, re­
spectively. Six habitat types were delineated on.the timbered regions.
Large mixed sex bull-cow groups observed in late July dispersed to form
small single sex bull groups and cow groups which increased in size to
late August. Environmental features recorded to compare areas selected
by each elk group type were cover type, habitat type, timber type,
distance to water, distance to the next closest cover type, downfall,
sight distance, location, topography, elevation, aspect, and slope.
Significant differences existed between preferences of group types for
topography and aspect (P<.05) and location (P<.01). Selection of other
environmental features were not statistically different. Bulls
abandoned their environmental preferences in late summer to.join cow
groups in their preferred habitat.
Canopy cover of tree and shrub species, general ground cover, and
the three dominant species in each of nine 2 X 5 decimeter frames were
recorded at 21 feeding and 28 bedding sites. Ground cover at both
types of sites was similar. Each group was observed in one of five
basic activity patterns and in one of three response states. Elk in
the restless response state were significantly more common during
weather phase one (Landsberg, 1969). The "head jerk" and "humped back"
postures and other dominant and subordinate gestures were observed.
Rut-related activities began with velvet shedding, followed by herding,
wallowing, bugling, and sparring.
Dry weight standing crops and percentage moisture contents were
determined for forbs and grasses which were clipped every two weeks
from elk feeding sites and from established timber and meadow plots at
three elevations. Forbs at elk feeding sites had significantly greater
moisture contents than forbs in all meadow and most timber sites;
standing crops of forbs at feeding sites were also significantly greater
than forbs at all timber and most meadow sites. Percentage moisture
content of forbs fell below that of grasses during October, which was
also the only time that moisture content of grasses at established
plots exceeded that of grasses at feeding sites. However, elk moved
into the timber in September, coincident with the rut, and reappeared
in high open parks in October. Based on relative percentage moisture
content, movement into the timber solely for succulent vegetation would
not be founded; therefore, during this study it appeared that security
requirements of elk were involved in this fall movement into the timber.
INTRODUCTION
The Rocky Mountain elk
(Cevvus canadensis netsoni) is an integral
part of many Montana ecosystems, and because elk are valuable animals
from the standpoint of economics, recreation and aesthetics, knowledge
of the relationship of the species to the environment is essential.
Elk consistently select certain aspects of various environmental
features (Lonner, 1974) such as slope, aspect, topography, and cover
type.
.
The size, composition, and habitat preferences of elk groups
change with time and environmental conditions (Knight, 1970).
Large
early summer migratory groups become smaller dispersal groups of mid
summer which become larger bull-cow groups once again with the onset
of rut (Martinka, 1965).
Behavioral patterns and hierarchical systems
change with the season and hormonal development (Struhsaker, 1967).
Matriarchal group control is the rule throughout the summer (Cole,
1969), but aggressive mature males join already formed large cow
groups during the rut and assert their dominance in the new associ­
ation (Altmahn, 1952).
According to Darling (1937), red deer stags
move from their summering areas to join hind groups in their
preferred habitat in the late summer, but similar movements have not
been noted for elk.
Seasonal movements of elk have been attributed;
to the phenologies! development of vegetation (Brazda, 1953; Picton,
-2—
1959; Kirsch, 1962; Martinka, 1965; Stevens, 1966; Knight, 1970;
Coop, 1971; Day, 1973), but though this may be a cause of such
seasonal movements, it may not be the only cause (Cole, 1969; Ream
et at., 1972).
During the summer and fall of 1973 and 1974, elk were studied .
in the Yellow Mule drainages of southcentral Montana.
The purpose of
the study was to describe some of the environmental features of the
elk's summer habitat and various daily and seasonal social inter­
actions and group changes.
This information may help to shed
additional light on the ecological requirements of elk and provide
some baseline data for future monitoring of the elk in the Yellow
Mule area.
The National Science Foundation RANN Program provided support
for the Gallatin Canyon Case Study team to assess the impact of
the Big Sky development on the Gallatin Canyon region.
My project
was an offshoot of this endeavor and was funded in part by the NSF
RANN Program grant numbers GI-29908x and GI-29908xl.
METHODS
Field work was conducted from July 24 to October 5, 1973, and
from July 6 to October 15, 1974.
Travel within the area was
unrestricted in 1973, but in 1974 the area was closed to all but two
wheeled motorized vehicles which were restricted to authorized trails.
Travel to and from the area was by motorcycle, and travel within the
area was on foot.
Elk observed with the aid of a 32X.spotting scope and 7 X 35
binoculars were classified into one of three elk group types.
Bull
groups consisted of bulls and spikes, alone or in association with
each other.
Cow.groups included cows, calves and spikes, alone or in
association with each other, but lone spikes were classified as bulls.
Bull-cow groups always included at least a bull and cow, but might
also include calves and spikes.
The size, composition, activity,
response state and behavior of groups were recorded.
Aerial obser­
vations in 1973 and 1974 were made over the area.during 12 flights
in a light plane.
Environmental characteristics recorded for each group were
cover type, habitat type, timber type, distance to water, distance
to next closest cover type, downfall, sight distance, location,
topography, elevation, aspect, slope and weather conditions.
Relative
tree densities and distribution ascertained from aerial photos and
-Iir-
topographic maps were used to determine the park, broken park and
timber cover types.
Forest areas were classified and mapped by
habitat type, according to Pfister
et al. (1974).
Points along
systematically traveled routes were habitat-typed and plotted on
topographic maps.. Habitat type boundaries were mapped by
delineating areas of similarly habitat-typed points.
Notes were made
at each point on understory and overstory vegetation composition.
Associations of five tree species comprise four timber types present
in the overstory.• Identification and distribution of each timber
type was determined from ground reconnaissance, aerial photos, and
U.S.D.A. Forest Service timber inventory maps.
Measurements of
distance to free surface water and next closest cover type were
grouped in categories of .less than 100 meters, less than 500 meters,
and more than 500 meters.
Downfall in the nearest timber stand was
recorded as none, light (scattered fallen trees which do not .appear
to deter elk movement), moderate (fallen trees which retard movement),
and heavy (thick downfall in which movement is extremely difficult).
Sight distance, in the timber stand where elk were observed, or with
which elk were most closely associated, was recorded as less than 20
meters, less than 50 meters, and more than 50 meters.
noted by drainage.
Location was
The topography of the immediate area where a
group was observed was recorded as ridge, upper slope, mid slope,
:
'
lower slope, bench or flat, or stream bottom. Elevation was recorded
.
to the nearest 30.5 meter interval using a U.S. Geological Survey
. ' 1 i'1
•.
15 minute topographic map.
Aspect of ea;ch site was categorized as
north, northeast, east, southeast, south, southwest, west, and north­
west.
Slope categories included 0-15 percent, 16-25 percent, 26-35
percent, 36-45 percent, and over 45 percent.. ■Aspect and slope were
'
.
measured with a Bruntdn transet.
•
Wind direction and velocity, cloud
cover,.and temperature were estimated in the. field for each obser­
vation.
Barometric pressure for each day was later derived from
records kept in Bozeman by Harold D. Picton.. Various combinations of
wind velocity, cloud cover, temperature and barometric pressure
comprise six weather phases as described by Landsberg (1969) .
(Appendix Table 12).
One of the six wehther phases was ascribed to
each observation of elk.
Use of the terms significant and highly significant indicate
statistical significance at the five and one percent levels,
'
respectively.
,
•
"
‘
Selected feeding and bedding sites from each two week period in
1974 were sampled with a 141.5 square meter circular plot centered
on the spot of most intense elk activity;
In it a species list was
compiled; scientific and common names follow Booth (1950) and Booth
and Wright (1959).
General ground cover•including grass, forbs,
litter, rock and bare ground was recorded by the canopy coverage .
method (Daubehmire, 1959) in nine 2 X 5
decimeter.frames;
Frames .
—6—
were arranged in three concentric circles within the circular plot.
The outer circle of four frames circumscribed the perimeter of the
plot:
the inner, circle of four frames located half way between the
edge and center alternate with those in the outer circle; a single
frame was placed in the center.
The three low growing species with
the greatest aerial coverage per frame were recorded as dominant
species.
Overall dominance values were assigned to each dominant
species to indicate the percentage of frames it occurred as a
dominant species in 189 and 252, 2 X 5
decimeter frames analyzed
at feeding and at bedding sites, respectively.
Data recorded for each
site indicated whether a species was present, and if it was a dominant,
in what percentage of nine frames it occurred as a dominant.
Shrub
cover was measured with the line intercept method of Canfield (1950),
along two perpendicular lines each bisecting the plot.
Tree species,
size classes in four centimeter increments (dbh), and canopy coverage
classes as described by Daubenmire (1959) were recorded for each
site plot.
In 1974, six 10.5 square meter permanent plots were established
east of the study area boundaries, near the Doe Creek road, to monitor
the phenoldgical development of vegetation.
located at.each of three elevations:
Pairs of plots were
2200, 2500, and 2800 meters.
One of each pair of plots was located in a meadow and the other was
located in the timber.
Ten 2 X 5
decimeter frames were clipped in
-7each plot every two weeks from mid July to mid October, 1974.
Vegetation from each plot was separated into grasses and forbs, put
in plastic bags, labeled, weighed in the laboratory on a Mettler
balance scale to the nearest 0.25 gram, transferred to paper bags,
labeled, dried for at least 24 hours at 90 C, and reweighed.
Per­
centage moisture content was calculated for grasses and forbs from
each plot according to the formula, Wet
Vegetation from five of nine 2 X 5
X 100.
decimeter frames at each of
21 elk feeding sites was clipped and separated into grasses and forbs
and similarly weighed, dried and reweighed.
sites was clipped each two week period.
A maximum of four feeding
THE STUDY AREA
The Yellow Mule study area (Figure I) is located about 4.8 kilo­
meters south of Meadow Village of Big Sky, Montana, in the Madison
Mountain Range of the Gallatin National Forest.
The 24.1 square
kilometer area is bisected from north to south by the Madison-Gallatin
County line and encompasses the First and Second Yellow Mule Creeks
and the bordering ridges from.the Buck Creek ridge at 2865 meters to
the 2195 meter level near the South Fork of the West Fork of the
Gallatin River.
In 1973 the area boundaries were not well defined,
and observation trips were also made to Buck Creek, Beaver Creek,
McAtee Basin, Muddy Creek.and Third Yellow Mule Creek.
The large amphitheater basins of. Muddy Creek, the First, Second,
and Third Yellow Mule Creeks are indicative of possible glacial action
during the Bull Lake glacial stade.
There are no interlocking spur
creeks between drainages, and the valleys are wide straight troughs
providing evidence of glacial activity, although mass gravity action
in the region has largely obscured glacial evidence.
Extensive
alluvial deposits occur in the basins at the heads of these drainages.
They are derived from sedimentary parent material of sandstone, shale
and limestone (Walsh, 1971).
The Gallatin Gateway 26 SSW weather station is located 4.8 kilo­
meters northeast of the study area in Beaver Creek at 2012 meters and
provides the nearest record of climatological data.
The seven year
PARK
' I BROKEN PARK
T I MBER
Flattop ^Atrv
I Ki l omet e r
Figure I
Map of study area showing cover types.
-10record (U.S. Weather-Bureau, 1973 and 1974) indicates an.average annual
temperature of 2.6 C and an average annual precipitation of 56.95
centimeters.
The study area is from 240 meters to 850 meters above
the weather station and receives substantially more precipitation
and experiences colder temperatures.
According to the U.S.D.A. - S .C ;S
(1974) precipitation map, from 76.2 to 127.0 centimeters of moisture
falls annually, and Caprio (1965). indicates that less than thirty
frost free days occur per year on the study area.
The area is part of a cattle grazing allotment leased by the
Forest Service from July I to October 15.
Cattle did not arrive on •
the area until September 9, 1973, but they were present at high
elevations within the area from July 11 until after October 15, 1974.
The study area was divided into three general cover types as seen
in Figure I.
The park cover type covers 31.0 percent (7.5 square
kilometers) of the area.
It consists of terraced benches with long
bands and island-like clumps of subalpine fir
Engelmann spruce
(Abies 'lasioeavpa),
(TPicea engelmannii), and whitebark pine (Pinus
albicaulis) (Figure 2).
The large basins of both the First and,Second
Yellow Mule Creeks contain 67 percent of the park cover type and
have a highly heterogeneous nature with interspersed creeks, willow
flats, tree islands, meadows, rock piles and bare ground.
The broken park type is composed of small meadows separated by
interconnecting tree stands (Figure 3).
It covers 13.3 percent (3.2
-11-
Figure 3.
Broken park cover type.
-12square kilometers) of the study area and is located primarily on the
southeast border, but broken park areas heavily used by elk are
centered below Flattop Mountain (Figure I).
These cover 0.8 square
kilometers, or less than one percent of the study area.
The timber cover type covers 55.7 percent (13.4 square kilo­
meters) of the area.
Six of Pfister1s et at. (1974) forest habitat
types, along with seven phases, were recognized within the timber
cover type.
The area was mapped on a gross scale according to the
following four major habitat types; Abtes Zasiooappa (Pi-nus
atbieaulis)/Vaecinivm Seopaviim3 Abies lasioeavpa/Vaeeinium Seopaviim3
Abies Zasioedvpa/Linnaea boveaZis3 Abies Zasioeavpa/CaZamagvostis
vubeseens (Figure 4).
All habitat types fell within the Abies
Zasioeavpa series which is composed of three elevational categories:
temperate, subalpine and timberline.
There were no timberline
habitat types on the area.
Abies Zasioeappa(Pinus aZbieauZis)/Vaceinivm seopavivm habitat
type (AF(WBP)/Vase) : This is the highest and most extensive subalpine
habitat type covering 48.5 percent of the forested area (6.5 square
kilometers) from approximately 2560 meters to 2865 meters (Figure 5).
It is the only subalpine type to occur on the area.
Subalpine fir
is the climax tree species, but whitebark pine is a long lived serai
dominant which is a conspicuous feature along the dry windy rims of
HABI TAT
TYPES
A iries Ia s io e G T ia (Jrvr-^s a Z&i&guZi a J/ Vcicciniien scoparium
IDI
1
2
3
4
A bies laeicecz^cz/Vaccir.ii<r. s c c z a riu r.
A bies IasicQGTeaZLirjrjeaecL borea l i e
A bies I a s i a c a r f ^ CalanagToet i e Tubeseer.s
timber
types
subalpine fir, vhirebark pine, Engelmann spruce
subalpine fir, Engelcann spruce, lodgepole pine
lodgepole pine
subalpine fir, Engelmann spruce, lodgepole pine,
Douglas-fir
I
H
W
I
A
I K i l o m ete r
Figure 4.
Map of study area showing habitat types and timber types.
-14-
Figure 5.
Abies IaeioocatPa(Pinus albioaulis)/Vaccinium soopariwn
habitat type.
-15the amphitheater basins.
Varying, proportions of subalpine fir;
whitebark pine and Engelmann spruce comprise the single timber type
which occurs on this habitat type (Figure 4).
The understory has
a paucity of species but consistently includes grouse whortleberry
(Vaceiriiim seopariurn)'which often forms a thick mat in the more open
stands where whitebark pine is highly represented.
Elk sedge (Capex
geyeri) may replace grouse whortleberry on the dry upper ridges.
Where the canopy is closed, bare ground, litter and scattered Avriica
oovdifolia are most common.
Abies Zasioeavpa/Vaoeiniwi seopaviim habitat type (AF/Vaso):
This temperate habitat type forms the lower border of the Abies .
lasiooavpa(Finns albicaulis)/Vapeinium seopaviim type, generally,
ranging from 2316 meters to 2560 meters and covers about 32.9 percept
of the forested area (4.4 square kilometers).
of two timber types as shown on Figure 4.
It includes portions
Subalpine fir is the
climax species, and Engelmann spruce is a major component on one of
the three phases.
Lodgepole pine (Finns eontovta) is widely
distributed in stands of varying size classes and densities.
The three phases of this habitat type occur on distinct areas
but often intermix to form complicated mosaics.
The Vaeeiniwn
seopaviim phase apparently occupies cold dry sites on relatively.
flat ground, usually in the upper regions of the habitat type
(Figure 6), however, it does occur along the northern edge of the
I
—16type to 2316 meters under lodgepole pine stands or under heavy down­
fall.
Other understory species are sparse but include elk sedge and
Arnica cordifolia. .
Shady, cool, moist sites appear to support the ThaZ-ictrum
occZdentate phase, characteristic of inclines and swales, whereas the
VaccZnZim eeoparZum phase occupies flat areas.
typical of this phase.
Engelmann spruce is
Understory species include western meadow rue
(ThalZctrwn oeeZdentale) ,• grouse whortleberry, pine reedgrass
(CaZamagrostZs rubescens) , elk sedge, ArnZaa cordZfoZZa and ArnZea'
.
ZatZfoZZa.
Probably the least extensive of the three phases is the
CatamagrostZs rubescens phase which occurs on warmer,,more open areas.
These sites are characterized by an association of pine reedgrass and
Oregon grape (BerberZs repens) or kinikinnick (AretostaphyZos uvd-ursZ)
and are most common along the lower edge of the type around 2260 .
meters.
The CaZamagrostZs rubescens phase becomes increasingly
interspersed with the ThaZZetrum oceZdentaZe phase to 2350 meters.
Only this phase occurs in association with Douglas-fir (Tseudotsuga
menztesZZ).
Pine reedgrass occurs at higher elevations but without
the other necessary indicator species.
AbZes .ZasZooarpa/Ltnnaea boreatts habitat type (AF/Ltbo) :
This
habitat type covers 17;I percent (2.3 square kilometers) of the
forested regions, and forms the northern boundary of the study area.
-17It extends beyond the lower boundary where logging operations are in
progress.
The lower elevational limits were not determined.
Gn the
west side of the First Yellow Mule Creek this type extends up to 2320
meters where it merges with the Abies Zasioear^a/Vaocvniim sooparium
habitat type Calamagrostis rubescens phase.
Small isolated patches
occur at 2440 meters on the west side of the area, but east of the
First Yellow Mule Creek, along the Beaver Creek-First Yellow Mule
ridge, the type is continuous to 2440 meters.
Lodgepole pine and
Douglas-fir are the most common tree species on this type, but subalpine fir and Engelmann spruce occur on wet sites, and subalpine fir
is the indicated climax.
Two phases are present in this habitat type.
The Abies- ldsioearpa/
Linnaea borealis habitat type Vacoinivm seoparivm-phase often occurs
in conjunction with the Calamogrostis rubeseens phase of the Abies
lasioearpa/Vaeeinivm seopariim habitat type.
Where stands of Douglas- •
fir and lodgepole pine form a mosaic pattern it appears as though the
Abies lasioaarpa/Linnaea borealis habitat type Vaeeinivm sooparium
phase is associated with the Douglas-fir, while the Calamagrostis
rubeseens
pine.
phase
of this
habitat
type
occurs under the lodgepole
The Vaeeinivm seopariim phase of the Abies lasioaarpa/Linnaea,
borealis habitat type is more common than the Linnaea borealis phase
because it occupies the more abundant dry areas.
Along with twin-
flower (Linnaea borealis) and grouse whortleberry, western baneberry
-18(Aatea rubra), Oregon grape, klnikinnick, Arnica aordifolia, Arnica
IatifoZia , wlntergreen (Pyrola asarifolia) and western meadow rue
occur as some of the major understory species.
The Linnaea borealis phase is minor and usually is limited to the
cool wet buffer zone between the dry Vaccinium saoparium'phase of the
timber and the wet Abies ldsiocarpa/Galium triflorum habitat type of
creeks.
Pfister et al. (1974) indicated that the Vaccinium saoparium
phase was commonly found on the Beaverhead and Deerlodge National
Forests and in the Little Belt Mountains between 1920 and 2225 meters
elevation.
1
Abies lasioeccrpa/Calamagrostis rubesaens habitat type (AF/Caru):
This uncommon open canopy habitat type occurs on dry, warm, steep
slopes below 2380 meters.
It covers 1.5 percent (0.2 square kilo­
meters) of the forested area.
The most extensive sites are on the
west slope of the Beaver Creek-First Yellow Mule ridge..
Its presence
coincides with the southern limb of the timber type in which Douglasfir occurs (Figure 4).
Douglas-fir and lodgepole pine dominate the
stand although subalpine fir and Engelmann spruce are present.
Heavy
litter is common in the understory and grouse whortleberry is con­
spicuously absent.
Elk sedge, western meadow rue and Oregon grape
are some of the major understory species which occur along with pine
reedgrass (Figure 7).
i
H
VO
I
Figure 6.
Abies lasiocarpa/Vacoiniicm
sooparinm habitat type, the
Vaooinium sooparium phase.
Figure 7.
Abies lasiocarpa/Calamagrostis rubesoens
habitat type.
-20-
Abi-es lasiocavpa/Calamagrost1Ls canadensis habitat type (AF/Cacg):
Occupying cool, poorly drained seep areas or creek banks, this local
habitat type is the most extensive "wet" type occurring at all ele­
vations within the Abies lasiocavpa(Finns albicaulis)/Vaeeininm
soopavium and Abies lasiocavpa/Vaeciniwn saopavium habitat types.
It
is replaced on the eastern ridge of the study area by phases of the
Abies lasioeavpa/Galium tviflovum type.
Engelmann spruce is always
present, often as the dominant tree because of the wet edaphic
conditions.
Associated species include bluejoint reedgrass
magvostis eanadensis) , arrowleaf groundsel
(Senecio
(Cala-
tviangulavis ),
American globeflower (Tvollius laxus), horsetail (Equisetum avvense)
and occasionally Labrador tea (Ledum glandulosum) and American
mannagrass
(Glyeevia
gvandis).
This type was often identified in the
broken park cover type as "stringers" (Figure 8).
Abies lasioeavpa/Galium tviflovum habitat type (AF/Gatv):
Because Abies lasioeavpa/Galium tviflovum is the warmest of the wetsite Abies lasioeavpa habitat types, it is confined to lower ele­
vations of the study area.
Both of the Abies lasioeavpa/Galium
tviflovum habitat type phases were present, but a distributional
dimorphism exists.
Because the Galium tviflovum phase is the warmer
of the two phases (Pfister, et al. , 1974), it was located exclusively
on the Beaver Creek-First Yellow Mule ridge, which
seems
to possess
Figure 8.
Abies lasiooarpa/Calcamgvostis canadensis "stringer"
surrounded by the Abies lasiocavpa(Finns albicaulis)/
Vaccinium seopaviian habitat type.
-22a somewhat warmer climate than the rest of the study area as. expressed
by the other warm climate habitat types found there.
associated with Engelmann spruce and Douglas-fir.
It was always
The Calamagrostis
canadensis phase is transitional to the cooler Abies lasioccorpa/.
'Calamagrostis canadensis habitat type and was located west of the
First Yellow Mule Creek, often in areas surrounded by lodgepole pine,
but. Engelmann.spruce always occupied the immediate site.
In contrast
to the "stringer" nature of Abies lasiocanpa/Cglamagrostis
canadensis habitat type, Abies lasiocorpa/Galium triflorum habitat
type is a restricted condition characteristic of swales.
The
indicator species, western baneberry, is found in association With
sweetscehted bedstraw (Galium triflonm).
Other species common to
both phases in this type are twinflower, horsetail, wintergreen,
western meadow rue and swamp currant (Ribes lacustre) . Arrowleaf
groundsel and bluejoint reedgrass are found only in the Calamdgrostis
canadensis phase.
.
RESULTS
Elk Group Characteristics
During the course of this study,, a total of 976 elk were ob­
served from the ground.
Of these, 281 were observed in 1974.
Eighty-eight percent of all elk were observed in parks.
If the total
number of elk using the area did not vary from year to year, it is
Vassumed that.a large portion of elk in 1974 were utilizing the
broken park and.timber cover types in which.observation was difficult.
Although fewer elk were observed from the ground in 1974, more were
observed from the air.
Ninety-two elk were observed in 1973 and 162
elk were observed in 1974 from the air.
The study area boundaries were redefined in 1974 to.make the area
smaller,, but when all elk observed outside the 1974 boundaries were
excluded, more than twice as many elk were still observed in 1973 as
in 1974.
More time was also spent looking for elk in 1974, so. the
area covered and time spent in the field were not factors in the
numerical discrepancy;
Although the observed number of elk varied from one year to the
next, those animals observed appeared to exhibit similar activities
both years and provided a basis for. analysis of interactions of elk .
with other elk and with the environment.
-24Group Size and Composition
The sizes of mixed sex bull-cow groups were greatest during .the
second half of July. As the size of these groups decreased in August
:
1
.
"
.■
•
bull and cow groups increased -In size; bull groups in 1974 were an
exception (Table, I).
Large bull-cow groups of early summer dispersed
in mid summer to form single sex cow and bull groups.
Bull-cow
groups reformed in late August and early September, but moved into
the timber during September and became unobservable.
in October after rut.
They reappeared
In 1974, more bull-cow groups were observed in
October than in any other month, and although no bull-cow groups were
seen from the ground in October, 1973, several groups were observed
in high parks from the air.
A total of 181 elk were observed in
bull-cow groups on parks above 2560 meters during flights made in .
October 1973 and 1974 in drainages of the South and Middle Forks of
the West Fork of the Gallatin River.
The number, and size of .elk groups were compared under six
possible weather phases to determine whether either of these group
characteristics were influenced by weather.
Elk were equally as
likely to-be observed during each phase since an equal number of
observation trips made during each phase were successful and unsuc­
cessful.
The number of groups seen during each phase was pro­
portionate to the time spent in search of elk during each phase.
TABLE I.
PERCENT OF ELK GROUPS AND MEAN NUMBER OF ELK PER GROUP OBSERVED IN SEVEN TWO WEEK
PERIODS DURING 1973 AND 1974.
All Elk
1974
1973
96/6951
74/281
July 1-15
—
11/4.O2
Bull Groups
1974
1973
42/102
42/81
—
12/3.2
Cow Groups
1973
1974
32/248
21/95
—
14/5.3
Bull-Cow Groups
1973
1974
22/345
11/105
—
—
July 16-31
16/13.4
35/4.6
17/2.4
36/2.0
16/11.2
43/4.6
August 1-15
31/5.4
12/3.1
24/1.4
17/2.3
41/6.8
5/2.0
32/8.6
August 16-31
23/7.1
10/5.0
24/4.6
5/1.5
31/9.1
14/6.3
9/9.5
September 1-15
24/7.3
11/3.0
21/1.9
10/1.0
13/4.3
14/4.3
45/13.8
September 16-30
6/1.3
9/2.1
14/1.3
13/1.5
—
—
—
10/6.0
12/3.1
—
7/1.0
—
10/1.5
—
36/5.5
October 1-15
—
1Total number of elk groups/total number of elk.
2Percent of elk groups/mean number of elk per group.
14/43.0
18/23.5
9/10.0
18/6.5
9/7.0
Largest group sizes were observed during phase three (Table 2),
the period prior to a storm, yet the differences in group sizes.were
not significantly different (t-test).
Large group size cannot be:
attributed to time of year because groups observed during phase three
appeared to be nearly equally distributed throughout the summer.
■TABLE 2.
PERCENT OF ELK GROUPS AND MEAN NUMBER OF. ELK OBSERVED PER
GROUP DURING SIX WEATHER PHASES.
(Weather phases follow
those of Landsberg, 1969).1 .
Weather Phase
Elk2
I
14/5.O3
2
3
4
5
6
22/4.8
' 10/9.2
11/5.6
22/6.5
21/4.9
.
1See Appendix Table 12.
2Elk observed in 1973 and 1974 combined.
3Percerit of 170 groups observed/mean number of elk seen per group. -
Sites Used b y .Elk
Twelve environmental features of elk activity sites were tested
with a Chi Square test against cow, bull.and bull-cow group categories
to assess.differences amorig group preferences, with regard to each
feature. (Table 3).
Observations for 1973 and 1974 were combined.
Groups exhibited, differential selection with regard to topography
aspect and location.
topography.
Table 4 shows group differences with regard to
All three group types utilized the bench or flat areas
heavily, but bull groups were observed often on lower slopes while
cow groups arid bull-cow groups were observed on upper, and mid slopes
-27TABLE 3.
CHI SQUARE VALUES CALCULATED FROM COMPARISONS OF THREE ELK
GROUP TYPES FOR EACH OF TWELVE ENVIRONMENTAL FEATURES. .
Environmental Feature
Cover type
Habitat type
Timber type
Distance to water
Distance to next cover type
Downfall
Sight distance
Location
Topography
Elevation
Aspect
Slope
Calculated Chi
Square Value
3.824 .
2.529
5.792
4.479
4.906
4.307
1.719
35.103*
20.743**
6.978
28.091**
8.254
Degrees of
,Freedom
4
8
6
4
4
6
4
2
10
6
14
8
*Signifleant at the .01 probability level.
**Signifleant at the .05 probability level.
and only rarely on lower slopes.
All groups were commonly observed on
the north aspect (Table 5); however, cow and bull-cow groups were most
often observed on the northeast aspect, and bull groups were most
often observed on the west aspect.
Bull groups selected the Second
Yellow Mule while cow and bull-cow groups selected the First Yellow
Mule (Table 6).
The differences in distribution are highly significant
Only observations made in the upper basins of the First and Second
Yellow Mule were used in the Chi Square test because sample sizes were
too small in other areas for testing.
Table 7 is a summary of all environmental features in which the
preferences of different elk group types did not significantly vary;
-28TABLE 4.
PERCENT OF ELK GROUPS AND MEAN NUMBER OF ELK PER GROUP IN
THREE GROUP TYPES OBSERVED ON SIX TOPOGRAPHICAL AREAS.
Bull Groups
Cow Groups
Bull-Cow Groups
Topography
33/450
84/1831
53/343
10/2.Iz
Ridge
3/7.0
11/11.3
Upper Slope
19/6.0
12/1.4
. 24/14.0
Mid Slope
17/12.1
26/6.6
37/14.0 .
Lower Slope
24/2.1
3/7.0
5/1.7
28/6.6
Bench or Flat
31/3.3
27/16.6
Stream Bottom
6/1.2
6/4.0
9/3.6
1Total number of elk groups/total number of elk.
2Percent of elk groups/mean number of elk per group.
TABLE 5.
Aspect1
North
Northeast
East
Southeast
South
Southwest
West
Northwest
PERCENT OF ELK GROUPS AND MEAN NUMBER OF ELK PER GROUP IN
THREE GROUP TYPES OBSERVED ON EIGHT ASPECT CLASSES.
Bull-Cow Groups
Cow Groups
Bull Groups
33/450
84/1832
53/343
27/2.93
24/14.0
26/6.4
28/17.2 "
13/1.7
21/8.9:
8/2.1.
13/2.7
15/7.4
13/6.6
4/1.0
9/13.3
—
4/7.0
6/2.8
9/11.2
3/26.0
30/1.7
18/10.0
10/2.4
12/1.5
3/22.0
4/1.5
—
1Each class spans an arc of 45 degrees, with 22.5 degrees on either
side of center.
2Total number of elk groups/total number of elk.
3Percent of elk groups/mean number of elk per group.
TABLE 6.
PERCENT OF ELK GROUPS AND MEAN NUMBER OF ELK PER GROUP IN
THREE GROUP.TYPES OBSERVED IN TWO LOCATIONS.
Bull-Cow Groups
Cow Groups
Bull Groups
32/183
Location
65/1521
29/354
Upper First
28/1.52
84/5.6
76/12.1
Yellow. Mule
Upper Second
72/2.6
24/12,6
Yellow Mule
16/6.3
1Total number of elk groups/total number of elk.
2Percent of elk groups/mean number of elk per group.
-29TABLE 7., PERCENTAGES OF 170 ELK GROUPS RELATED TO VARIOUS ENVIRONMENTAL FEATURES, MEASURED DURING 1973 AND 1974.
Cover Type1
P
BP
T
80
11
Habitat Types"2
AF(WBP)/Vaso
9
AF/Vaso
81
10
Timber Types 3
AF-WBP--ES .AF-ES-LPP AF-ES!-LPP-DF
12
81
21
I
2 .
Distance to Water in Meters
500 or
100 or
500 or
Less
Less
More
I
39
N
Downfall4
L M H
4
25
50
AF/Caru ■ .
7
LPP
6
Distance to Next Closest
Cover Type, in Meters
500 or
500 or
100 or
Less
Less
More
78
AF/Caad
33
28 '
Sight Distance
in Meters
.20 or
.50 or
50 or
Less
Less
More
21
52.
38
10
Elevation in Meters
Less
Than
2530
12
Slope in Percent
2530-2650 . 2651-2772
:
40
23
2773-2865
1-15
25
16-25
54
19
26-35
9
Over
36-45 ■ 45
14
1P - park, BP = broken park, T = timber,
2See Study Area description, page 8.
^AF = subalpine fir, WBP = whitebark pine, ES = Engelmann spruce*
LPP = lodgepole pine, DF = Douglas-fir.
■•
4N = none, L = light, M = moderate, H = heavy.
4 •
-30data from all groups were pooled.
Although not significantly
different, environmental preferences of bull groups were consistently
different from the similar preferences shown by cow and bull-cow
groups.
Elk were most often observed in the park cover type
associated with the
',
Abies Zasiocarpa(Pinus: aZbioaulis)/Vaooinivm
saoparium habitat type.
Elk were not observed in close association
with water, but they were usually close to an adjacent cover type. '
Nearby timber stands had moderate downfall arid sight distances of
less than 20 meters.
They utilized slopes of less than 15 percent
and were often, seen between 2530 and 2650 meters elevation.
Because
an environmental feature may have been distributed in unequal pro­
portions in the landscape, elk may have used it. in proportion to its
occurrence, and the percentages listed in Table 7 may be a reflection
of this;
The objective was not to determine the type of habitat elk
prefer, but to assess differences in site characteristics selected
by different elk group types.
Elk bedding sites were analyzed separately from elk activity
sites.
A bedding site had from one to 106 beds.
were collected during the 1974 season (Table 8).
Data for bed sites
Beds were most
often found in the broken park cover type% north of Flattop Mountain,
less than 100 meters from water and cover, in areas associated with
light to moderate downfall.
Sight distance in adjacent timber
stands was usually less than 20 meters.
Sites were usually located
-31TABLE 8.
PERCENTAGES OF 83 ELK BEDDING SITES AS"RELATED TO VARIOUS
ENVIRONMENTAL FEATURES, MEASURED DURING 1974.
,•- •
Cover Type1
P
BP
T
31
45
Habitat Types2
AFCf/BPj/Vasa
24
AF-WBP-ES
30 .
LPP
5
Distance to Next Closest
Cover Type in Meters
500 or
100 or
Less
• Less
02
N
8
73
16
,
11
Sight Distance
in Meters
20 or
50 or
50 or
Less
Less
, More
Downfall4
L .M
H
51
I ■
4
Distance to Water in Meters
500 or . 500 or
100 or
Less
Less
More .
2
28
AF/Gatv
44
Timber Type3
AF-ES-LPP AF-ES-LPP-DF
'
AF/Cacq
16
35
63
98
AF/Vdsa
13
55
34
11
■
Slope in Percent
R
12
Topography5
US MS LS B/F SB
7 29
2 25 .25
Aspect6
N NE E SE S SW
48 21 7
11
W NW
5 12
5
1-15 16-25 26-35 36-45
75
19
4
I
1P = park, BP = broken park, T = timber.
2See Study Area description, page 8.
3AF = subalpine fir, WBP = whitebark pine, ES = Engelmann spruce,
LPP = lodgepole pine, DF = Douglas-fir.'
4N = none, L = light, M = moderate, H = heavy. '
5R = ridge, US = upper slope, MS = mid slope, LS = lower slope,
B/F = bench or flat, SB = stream bottom,
6Each aspect class encompasses a 45 degree arc.
.
Over
45
I
-32in moist- areas characterized by the
Abies lasioaarpa/Calamagvostis
canadensis habitat type, Engelmann spruce trees, the stream bottom
or flat topographies, on gentle north slopes.
Vegetation Analysis of Elk Feeding and Bedding Sites
General characteristics of ground cover on 21 feeding and 28
bedding sites were analyzed by the canopy coverage method of
Daubenmire (1959) as shown in Table 9.
While grass and forb coverage,
of feeding and bedding sites are nearly equal, bedding sites have
more litter coverage and feeding sites have more rock and bare ground
coverage.
TABLE 9. . MEAN PERCENT CANOPY COVERAGE OF GENERAL VEGETATION GROUPS AT
TWENTY-ONE FEEDING AND TWENTY-EIGHT BEDDING SITES AS DETER­
MINED BY EXAMINATION OF NINE 2 X 5 DECIMETER FRAMES AT ..EACH
SITE.
■
Feeding Sites
Bedding Sites
Grass
Forbs
Litter
. Rock
Bare Ground
27.8 .
28.7 .
46.7
42.9
19.1
29.3
2.2
0.7
13.7
.6.7
'
Table 10 and 11 list species with greater than 20 percent
constancy along with the dominance value of each and a presence list,
of all species at each feeding and bedding site.
Eighty percent of
the forb species over 20 percent constancy occurred in common on
feeding and bedding sites, which had 44 and 43 forb species,
respectively.
Eighteen grass species over'20 percent constancy
33
TABLE 10.
VEGETATION ASSOCIATED WITH TWENTY-ONE FEEDING SITES EXAMINED OVER A FOURTEEN WEEK PERIOD WITH REGARD TO CONSTANCY,
OVERALL DOMINANCE VALUES, AND PRESENCE, MEASURED IN NINE 2X5 DECIMETER FRAMES WITHIN A CIRCULAR PLOT AT EACH SITE.
Only species with twenty percent or greater constancy are listed.
C1
D2
100
91
91
86
81
71
67
67
67
67
62
62
52
52
52
52
52
48
43
43
38
38
38
38
38
38
38
33
33
33
33
33
33
29
29
29
29
29
24
24
24
24
24
24
20
8
13
17
I
I
24
5
10
6
7
2
I
91
86
71
67
62
62
52
43
43
29
13
18
25
15
2
2
I
9
3
_________________________________Individual Bedding Sites_______________________________
Mid July
Late July
Mid August
Late August
Mid September
Mid October
FORBS
Aohillea millifolium
Agoeerie glauoa
Potentilla graoilie
Taraxicum offioianale
Solidago multiradiata
Erithronium grandiflorum
Aeter fo Iiaoeue
Seneoio oraeeulue
Geranium viBooeeieeimum
Lupinue argenteue
Helianthella quinquinervie
Thaliotrum, oocidentale
Arabie drurmondi
Epilobium anguetifolium
Eragaria virginiana
Merteneia oiliata
Pedicularie bracteoea
Polygonum bietortoidee
Allium eohoenopraeum
Aetragalue alpinue
Arnica cordifolia
Collomia linearie
Dodeoatheon pauoiflorum
Epilobium glanduloeum
Thlaepi fendleri
Valeriana oocidentale
Viola nuttallii
Delphinium bicolor
Delphinium oocidentale
Geranium riohai'deonii
Lupinue eerioeue
Myoeotie eylvatioue
Hanunoulue eeoholtnii
Rumex pauoifoliue
Equieetum arvenee
Gaum triflorum
Heraoleum lanatum
Polygonum douglaeeii
E H o g o n u m umbellatum
Lyohnie druimondii
Phlox multi flora
Plantago tueedyi
Senecio triangularie
Trolliue laxue
+3 56- 22
_5 22 33
11
+
56 22
22
11
+
+
+
11
I
I
2
3
+
11 11 11 33
11
11 22
11 33 +
67 11 + 44
11
I
+ 11 11 +
11
11 11
22 33 + 33
22
33
22
11
33 11 22 44
+ 11
+ 11
22 11
22
11
11
22 11 44 67
11
11
11
22
11
89 89
33 11
11 11 22
11
89
44
56
- 11
56
44
33
11
33
11
22
22
22
11
+
11
22
56
22
67
22
+
11
33
44
56
44
11
33
11
56
11
+
22
11
7
6
2
I
4
3
11
33
11
44
11
44
-
33
11
+
+
—
11
+
+
—
-
11
-
+
11 + 33
11 11 11
33
11
22
33
11
11
11 11 22
33
11
11
44
33
11
11
+
I
I
5
-
11
22
44
+
-
3
I
8
3
I
I
44
11
11 11
+
-
33
11
56
33
56
44
+ 22
-
22
11
I
I
3
5
—
—
+
+
—
—
+
+
-
+
+
11
-
-
11
11
22
33
22
33
56
GRASS AND GRASS-LIKE PLANTS
Phleum alpina
Agropyron oaninum
Peetuoa idahoeneie
Bromue oarinatue
Melioa epeotabilie
T H e e t u m epioatum
Poa reflexa
Carex spp.
Deeotuwrpeia oaeepitoea
Poa alpina
11
22
22 33
89
89
22 44
44 11
44
56 11
33
11
11
11
56 11
11
11
33
-
22 22
33 11 67 22
44
11
11
11
33
11
+
11 22
67
- 78
44
I
11
56 11
-
22
+
44
33
+
22
11
11
33
22
11
33
11
11
33
67
11
11
11
1C - Constancy, percent occurrence among all sites.
2D - Dominance Value, percent occurrence of the species as a dominant within 189 2X5 decimeter frames analyzed on bedding sites.
*+ - Proaunt on the site but not occurring as a dominant.
'•percent occurrence as a dominant in nine 2X5 decimeter frames analyzed on each site.
,J- - Not present on the site.
34
TABLE 11.
VEGETATION ASSOCIATED WITH TWENTY-EIGHT BEDDING SITES EXAMINED OVER A FOURTEEN WEEK PERIOD WITH REGARD TO CONSTANCY,
OVERALL DOMINANCE VALUES, AND PRESENCE, MEASURED IN NINE 2X5 DECIMETER FRAMES WITHIN A CIRCULAR PLOT AT EACH SITE.
Only species with twenty percent or greater constancy are listed.
C1
D23
4
86
86
82
68
64
61
61
61
57
54
50
50
50
46
46
46
43
43
39
36
36
36
36
36
32
32
32
32
32
32
32
32
29
29
25
25
25
25
25
21
21
21
21
10
5
15
7
8
15
4
5
16
7
10
2
12
I
3
I
I
75
75
68
68
61
54
54
39
39
29
29
25
25
21
21
21
21
21
18
2
19
2
3
5
I
I
I
32
6
____________________________________ Individual Bedding Sites____________________________________
Mid July
Late July
Mid August Late August Mid September Late September
Mid October
FORBS
Achillea millifoliwn
Erithroniwn grandiflorwn
Potentilla graoilia
Agoeerie glauoa
Taraxiown offioianale
Aetor fo liaoeue
Atitragalue alpinue
Pragaria virginiana
Eenoaio trianyuLaria
Thaliotrwn oooidentale
A m i o a oordifolia
Geranium vieaoeeieeimwn
Trolliue laxue
Epilobium anguetifolium
Lupinue argenteue
Pedioularie braoteoea
Epilobiwn g Ianuloeum
Polygonum bietortoidee
Equieetum arvenee
Caetillija rhexifolia
Helianthella quinquinervie
Ranunculue unainatue
Merteneia ciliata
Viola nuttallii
Dodecatheon pauciflorun
Haeaklia floribunda
Lupinue eericeue
Plantago tveedyi
Senecio oraeeulue
Solidago multiradiata
Unidentified Forbs
Veronioa uormekjoldii
Alliwn echoenopraeum
Arabia drwmondi
Epilobiwn alpinum
Pamaeeia fimbriata
Polygonum douglaeeii
Saxifraga arguta
Veronica eerpyllifolia
Delphinium biaolor
Myoeotie ayIvatiaue
Hanunaulue eeoholtzii
Valeriana ooaidentalie
+
11
+ 22
+
11 33
11 44 33 11
22 33
33 11 11
11
22 11 11 44
22 33
44
11
22
44
56
33
11
33
11
+
+ 22
+
33
11
11
33 44 11 44
44
33 22
44
67 33 44
22
11
22
22
33 33 56
11
11 11
56 22
22
11
11
11
11
11
11
33
44
33
33
22
- 11
11
11
11
11
11 22
1 1 + + - + + + + +
33 22 + - +
22 22 56
11 33 11
+
+
+
22
-
11
-
+ 22
-
11
11 89
11 33 + +
- - 44 +
11 22 22 11
11 - 44 67
22 + - +
- - 33 44
+
22 +
- 33 56 11
+ 22
22 44 11
- 33
3 3 + - - +
22
11
22
11
9
I
I
I
3
I
89
67 56
11
11 44
11
+
+
-
-
-
11
- - - 2 2 +
33 22
22 11
I
6
I
2
4
6
I
I
I
11 11
1 1 - - - 22
11
22 67
22
11
22
22
11
33
22
22
11
22 33
11
22
11
33
11
- 22
-
- 44 11
- 11
+ 11
11
I
22
4
I
I
I
I
GRASSES AND GRASS-LIKE PLANTS
Carex epp.
Phleum alpina
Peetuoa idahoeneie
Trieetum epioatum
Agropyron aaninum
Bromue oarinatue
Melioa epeotabilie
Poa reflexa
Deeohampeia oaeepitoea
Calamagroetie rubeeaene
Stipa oooidentalie
Calamagroetie oanadeneie
Poa alpina
Bromue anomalue
Danthonia intermedia
Elymue glauaue
Junoue spp.
Poa junoifolia
-3 22' 22 + s
67
- 76
22 44 44
44
+
_
+
22
22
. .
- 11
89 22
44 11
11
-
. 33
+
+
22 67
11
- 22 11
+ -
11 11
+ 44
33 11 22
+
33 44 22 11
89
44
11
67 22
33
11 22
33 11
11 11
22 11 11 33
22
11
44
+
-
1 1 - - - -
22
22 56
U l l
- - +
+ +
11 22
-
*
11
22
-
- 22 + - - 3 3
+ - -
+ 11
- 11 11 56
22
- 1 1 + - -
11
33
2 2 + - - 22
11
6
33
11
I
7
- 11 89
+ 11
-
+67
22
56
11
-
-
22
+
I
I
I
I
+ 11
11
11
+
2 2 + + - 11
11
11
33
*
++
+
+
SHRUBS
Vaooinium eooparium
+
+
+
+
+
+
1C - Constancy, percent occurrence among all aitea.
2D - Dominance Value, percent occurrence of the species as a dominant within 252 2X5 decimeter frames analyzed on bedding sites.
3- ■ Not present on the alts.
4Percent occurrence as a dominant in nine 2X5 decimeter frames analyzed on each site.
s+ - Present on the site but not occurring as a dominant.
- 44 33
-35occurred on bedding sites while feeding sites had ten species; all .
grasses occurring on feeding sites also occurred on bedding sites.
Species having constancies of less than 20 percent, along with their
overall dominance values, are listed in Appendix Table 13.
Shrub species intercepted on less than five percent of the
combined line length were not recorded as dominants, but when inter­
cepted on five to 25 percent of the line, they were recorded as trace
dominants.
line.
No shrub was intercepted on more than 25 percent of the
Shrubs were uncommon and scattered but occurred more often and
with a greater diversity on bedding sites than oh feeding sites.
On
bedding sites only grouse whortleberry appeared with over 20'percent
constancy.
No shrub of 20 percent constancy occurred on feeding sites'
The number, size (dbh classes), species and canopy coverage of
trees occurring in 141.5 square meter circular plots were.noted.
Nineteen of the 28 bedding sites were under trees while only three of
21 feeding sites were under trees.
Bed sites occurred under all
classes of canopy coverage except the 95 to 100 percent cover class.
Most beds were located under 25 to 50 percent cover, while feeding
sites always occurred under less than 25 percent cover.
,Subalpine
fir occurred on 19 of the areas sampled and was the most abundant tree
over all, while Engelmann spruce, whitebark pine, lodgepole pine and
Douglas-fir .occurred on 12, 12, 5, and I of the sites, respectively,
and decreased in abundance in the same order.
The number of trees
-36per site ranged from I to 99.
Due to the small sample size this is
a poor indicator of local tree density..
Trees ranged in size from
0 to 52 centimeters in diameter (dbh).
Eighty-seven percent of the
sites which had trees had at least one large tree (20 centimeters dbh
or larger).
Elk Activities
General Daily Activities
Each elk group observed was assigned one of the following five
basic activities:
bugling.
feeding, bedding, feeding and bedding, moving and
Groups were observed in these activities during 60, 10, 9;
16, and 5 percent of the observations, respectively.
Bugling, which
was not observed in conjunction with some other activity, was listed
as a basic activity.
The following minor activities, moving, bugling,
playing, herding, and sparring, occurred in conjunction with the
basic activities during 8, 6, 5, 5, and 4 percent of the observations,
respectively.
■ A diurnal, movement connected with feeding activity was noted
through the summer, but decreased with the onset of the rut in late
August.
In the evenings elk moved from timber to parks at high
elevations, where they fed. After feeding again in the mornings,
they returned to the timber.
-37Days were divided into three periods:
morning, from dawn to
10:00 a.m., mountain day-light.time; midday, from 10:00 a.m. to 5:00
p.m.; and evening from 5:00 p.m. to dark.
Ninety-three percent of
the groups were seen during the morning and evening periods, with 58
percent of the groups; and 63 percent of the elk being observed in the
morning.
The response state of elk was recorded with regard to weather,
to determine if weather was an influence on elk attitudes.
Each elk.
group was. categorized into one of three.response state, classes:
calm, alert, or restless.
Results of a Chi Square test indicated that
groups in the calm and alert classes, were as likely to be seen during
any of the weather phases, but restless groups were significantly more
likely to be seen during weather phase one, a period of high stable
pressure, clear skies, cool rising temperatures, and calm wind.
When
weather phase one and the associated elk groups were removed from the
.test, a Chi Square test on the five remaining weather phases and their
associated groups was not significant.
Social Behavior
The social structure of elk involves communication signals which
convey meaning to members of the group.
Individual conflicts are
minimized and group stability is maintained through agonistic and sub­
missive gestures of group members. ,
-38Three categories of agonistic behavior as described by deVos
et'at.
(1967) include weapon threats, scare threats, and present threats.
I
■
■
Weapon threats involve presentation of weapons which are to be used
during an interaction such as hooves, teeth, and antlers. Weapon
threats'were first observed in males on.July 8, and were last observed
on October 4., while in females they were observed only in July.
Compared to weapon threats, scare threats and present threats are
lower intensity intimidation gestures.
Scare threats, first observed
in males July 14, were last observed September 30, while in females
they were first observed July 8 and last observed September 7.
Present,
threats, seen only in males, extended from August 14 to September 10.
Gestures may differ between males and females. Weapon threats
exhibited by females included the single foreleg kick, rising onto
hind legs and lashing out with front feet, and elevated muzzle, which
is.a low intensity version of rising onto the hind legs (Struhsaker,
.1967).
Jawing was observed as a dominance threat with the apparent
intent of. biting rather than a submissive gesture as indicated by
Struhsaker (1967).
The only male weapon threat was lowering of the
antlers as in preparation for combat (Struhsaker, 1967).
Both males and females performed low stretch and the direct stare
during scare threats (Geist, 1971).
A scare threat observed twice
during this study, and exhibited by males, was labeled the head jerk.
During this display, the head and neck were extended parallel to the
—39ground and the antlers were located on either side of the back.
The
bull would quickly rotate his head on the horizontal axis causing the
antlers to swing over the back.
at the full extent of the arc.
The surroyal tines touched each flank
This gesture elicited immediate flight
in the individual being displayed to.
Present threats of bulls emphasized physical features of the
displaying individual.
An elevated
head
fully exposed the dark mane,
while a broadside present emphasized body size, and swinging the head
to and fro gave full view of the antlers to the displayee.
Submissive postures were elicited from subordinate elk by
dominant elk who displayed some type of agonistic behavior.
Lowering
the head and looking away, the nose down retreat, and the muzzle up
retreat, were submissive postures observed during this study, which
have been described by Struhsaker (1967) and Geist (1971).
Both
retreat postures resulted in presentation of the rump patch when the
individual walked, trotted or stotted away.
Another submissive
posture, which was observed twice during this study and has not been
previously described, was the humped back posture.
A small or
subordinate individual would stand or hop with all four feet close
together, causing a rounded back and lowered head.
Small or young animals were subordinate to larger or older
animals.
Bulls were dominant over cows, cows were dominant over
spikes and calves, and spikes were dominant over yearling cows.
!
)
-40Calves assumed the rank of their mothers, but were lowest in the
system when disassociated from the parent.
Rut - Related Activities
Physical and behavioral changes were evident in male elk during
mid to late summer.
The occurrence of rut-related activities such as
velvet shedding, herding, wallowing, bugling and sparring are pre­
sented in reference to time in Figure 9.
First indications of velvet
shedding were noted July 28, and most bulls had completed the process
by August 24. 'Herding of cows and calves by a single bull was first
observed on July 25, but did not become common until mid to late
August. . The first observation of wallowing was noted August 13 when
a conflict over a wallow occurred between two bulls.
sites were noted up to October 14.
Active wallowing
Spikes were observed to wallow in
both dirt and water during mid July, but the activity was related to
insect annoyance and not- to rut.
The initiation' of bugling and
wallowing coincided, however, only two bugles were noted before the
first of September.
On August 8, two bulls responded to imitation
bugles by running toward the sound,, but they did not bugle. ' Peak '
bugling occurred during the last ten days of September.
sham fighting were first observed August 28.
Sparring and
This activity peaked
during late August and quickly diminished, but was last noted October
4.
High pitched squealing sometimes accompanied this activity.
An
apparent modification of sparring was noted on July 14 when two bulls
24
28
V e l vet
C
S hedding
3
25
H e rd in g
I___________
13
14
W a llo w in g
13
- 1.‘iv
4
B u g lin g
28
4
S parring
JU L Y
Figure 9.
A U G U S T
Duration of rut-related activities.
S E P T E M B E R
OCTOBER.
(Data from 1973 and 1974 have been pooled.)
in velvet approached one another, placed their noses in contact, and
commenced pushing.■
Other rut-related activities were noted.
Bulls holding- their
mouths open as described by Struhsaker (1967)' were recorded three
times, and a single observation of the lip curl was recorded August
29.
In early. August, harassment by mature bulls began with calves; by
late August it had extended to spikes, and by mid September conflicts
between adult bulls were observed.
A spike expressing interest in a
cow on September 10, repeatedly smelled her perineal region, and was
soon expelled from the group by the harem bull.
approached a bull as in no manner seen before,
On September 10 a cow
She approached him
closely, circled, and stared at him for several minutes.
her with his antlers and herded her into the timber.
He threatened
A declining
interest.in harems was apparent during the first half of
October when
.herd bulls became tolerant of the presence of smaller bulls and herding
activities dwindled.
-
'
'
•
Vegetation Phenology
.
'
'
-
The standing crop and percentage moisture of vegetation at twelve
cow and nine bull elk feeding sites, and at six permanent plots were
monitored during the 1974 summer (Appendix Tables 14 and 15).
Data,
from bull and cow feeding sites were pooled because they were not
significantly different in standing crop or moisture content.when
tested with an analysis of variance test.
The dry weight standing
-■
-
43
-
crop and percentage moisture of forbs at elk feeding sites were
significantly greater than grasses (P<.01).
Data on standing crop and percentage moisture from established
plots were tested with an analysis of variance test to determine
whether differences existed between meadow forbs in each of three
elevational plots, between timber forbs in each of three elevational
plots, between meadow forbs and timber forbs at each elevation, between
forbs and grasses in meadows at each elevation, and between forbs and
grasses in timber at each elevation.
The first three tests were also
calculated with data from grasses.
The standing crop of forbs in meadow plots decreased with in­
creasing elevation.
The differences between plots at each successive
elevation was highly significant.
Grasses in the 2500 meter meadow
plot had the highest standing crop while those in the 2800 meter plot
had the lowest, but the standing crop of all plots were significantly
different (P<.01).
The standing crop of forbs in timber was highly
significantly greater in the 2500 meter plot than in the 2200 meter
plot and was significantly greater in the 2800 meter plot than in the .
2200 meter plot.
The standing crop of grasses in timber was signifi­
cantly greater in the 2200 meter plot than in the 2500 meter, plot arid
was highly significantly greater in the 2200 meter plot than in the
2800 meter plot.
Standing crops of forbs in meadow were significantly
larger, at the one percent level, than forbs in timber at each
-44elevation.
This was also true.for grasses.
When forbs were compared
with grasses in meadows' and in timber plots at each elevation, grasses
in the 2200 meter timber plot had a significantly larger standing crop
than forbs (P<.01), while forbs had a significantly larger standing
crop than grasses in the 2500 meter, timber and 2800 timber and meadow
plots (P<.01).
Moisture content did not significantly differ between grasses and
■forbs within each plot, or when comparing either grasses or forbs be­
tween meadow and timber.
Only in the 2500 meter timber plot was the
percentage moisture o f 'forbs significantly higher than grasses..
All elk feeding sites were in.meadows, so they were compared with
established meadow plots.
Forbs at elk feeding sites had a signifi­
cantly larger, standing crop than forbs in the 2500 and 2800 meter
plots (P<.01).
The standing crops of grasses at elk feeding sites
were significantly higher than grasses at 2800 meters.
The percentage
moisture content of both forbs and grasses was significantly larger at
elk feeding sites than forbs and grasses at all meadow plots (Pc.01).
Data collected from elk feeding sites during the mid September and mid
October collection periods indicate that forbs had a significantly
higher moisture content than forbs from established plots clipped
during the same periods.
Grasses from feeding sites clipped during
the mid August and mid September periods also had a significantly
higher moisture content than grasses at established plots.
The
-45percentage moisture content .of both forks and grasses from feeding ,
sites were also significantly higher than forbs and grasses from
timber plots.
The standing crops of vegetation on both elk feeding sites and
.established plots varied during the summer, and exhibited an overall,
decrease from July to October (Figure 10).
Grasses exhibited less
fluctuation and less overall standing.crop.decrease than forbs.
At
established plots, standing crops of forbs in the timber varied IesS
and decreased less than in meadows.
Standing crops of grasses also
varied less in timber than in meadows, but remained about the same, or
increased slightly in timber sites from July to October.
The percentage moisture content of clipped vegetation at
established plots decreased as the summer progressed (Figure 11).
Percentage moisture content of vegetation decreased faster in meadow
plots than in timber plots, and it decreased faster in forbs than it
did in grasses.
The percentage moisture content of forbs in meadows
was slightly greater, than that of grasses until late August in the
2200 and 2500 meter plots and until late September in the.2800 meter
plot.
In the timber, percentage moisture content of forbs was
slightly greater than grasses until mid October.
At elk feeding sites percentage moisture content also decreased
.faster in forbs than it did in grasses, but at all times, forbs had a
higher percentage moisture content than grasses.
The rate of
GRAMS
ELK
PARK
TIMBER
VELK
Zlll
AUGUST
Figure 10.
SEPTEMBER
OCTOBER
JULY
AUGUST
SEPTEMBER
OCTOBER
Grams dry weight standing crop of forbs (left) and grasses (right) at twenty-one
elk feeding sites and six established plots clipped every two weeks from mid July
to mid October.
(Refer to Tables 14 and 15 for data from individual sites. Data
from individual elk feeding sites were averaged for each clipping period.)
\ «71)
47-
PERCENTAGE MOISTURE CONTENT
---PARK
TIMBER
JULY
Figure 11.
AUGUST
SEPTEMBER
OCTOBER
AUGUST
SEPTEMBER
OCTOBER
Percentage moisture content of forbs (left) and grasses (right) at 21 elk feeding
sites and six established plots clipped every two weeks from mid July to mid
October.
(Refer to Tables 14 and 15 for data from individual sites. Data from
individual elk feeding sites were averaged for each clipping period.)
“48desiccation of forbs at elk feeding sites was less than fofbs at
established.plots in both meadow and timber, while the rate of
desiccation of grasses at feeding sites was actually faster than.,
grasses at all established plots except the 2800 meter meadow plot.
In comparing percentage moisture content of forbs at elk feeding sites
with those of established plots for each sampling period, established
meadow plots surpassed elk feeding sites only during the mid July
period.
Percentage moisture content of forbs at timber plots.sur­
passed that of feeding sites during the mid July and mid August
periods.
Percentage moisture content of grasses in meadow plots never
surpassed grasses at feeding sites, but moisture content of grasses in
timber plots exceeded that of grasses at feeding sites during mid
October.
DISCUSSION
Elk group size, composition, and use of the landscape changed
from July to October.
Mixed sex migratory groups arrived on the
summer range during July and began to disperse into small single’ sex
cow and bull groups which reached peak sizes by late August.
At that
time members of bull groups dispersed, to either join an already
assembled harem, as indicated by Altmann (1952), or wander as
unattached singles.
During September, bull-cow groups moved from the high open basins
of the Yellow Mule Creeks into lower timbered areas.
They stayed in
the timber until October, when they reappeared in high parks above
2560 meters.
Ream et
at. (1972), also found that "following the peak
of the rut, the animals moved back to the upper slopes and ridges,
using mostly open exposures".
Areas used by elk reflected a certain need for security.
If not
actually in cover, elk and elk bedding sites were nearly always near
the edges of timber having moderate downfall and short sight distances
Altmann (1952) states that "elk prefer to rest in locations surrounded
by fallen trees which seemingly provide protection" and Allen (1972)
indicates that timber with short sight distances is preferred.
Janson
(1974) and Lonner (1974) found that heavy timber was preferred during
the rutting season, as was inferred during.this study because of
-infrequent observations made during this time.. The significant
-50differences in selection of topography, location, and aspect between
cow, bull-cow, and bull groups may be related to different security
level needs.
As indicated earlier,the consistent selection of
similar areas by cow and bull-cow groups, in contrast to areas
selected by bulls, suggests that female members of harems were
responsible for selecting the habitat used by the group.
Although
cows within the harem appeared to determine group movements, the
master bull would assert his dominance by keeping the harem herded
into a group and defending it from other bulls.
.Since bulls left the
Second Yellow Mule, where they had spent the majority of the summer,
.
to.join cow groups in the First Yellow Mule, it.appears, that they
abandoned their environmental preferences to assume the habits of the
cow groups which they joined.
Darling (1937) indicates that compared
to stags, red deer hinds have a higher requirement for security related
to maternal instincts; this might well be the case for elk.
Along with secure habitat, elk appeared to select areas which
provided cool, comfortable sites for daily activities. .Typical bed- '
ding sites were associated with moist areas characterized by gentle
slopes, the stream bottom and flat topographies, the
Abies lasiodarpa/
-Calamagrostis canadensis habitat type and Engelmann spruce trees,,
but the wetness of an area appeared to be the important factor in
selection of a site.
Diurnal movements of elk to open areas in the
evening and to timbered areas in the day indicate that elk may have
T - S l-
been selecting the coolest areas on a 24 hour basis where different
rates of heat loss caused' variation in microclimate.. Kirsch .(1962)
noted the same daily movements and related them to selection of
succulent vegetation in high open parks and ridges; this is in agree­
ment with the findings of this study which indicate that elk selected
'
meadow feeding sites which had higher percentage moisture content and
standing crop than vegetation at established meadow and timber plots.
Since most elk were observed feeding during the cool hours of dawn
and dusk and were only occasionally seen in .the open during the day,
it is likely that both vegetation moisture and climatic factors are
involved in this movement.
First signs of the rutting season occurred in late July of 1973
v#ien bulls began to shed antler velvet.
Cole (1969) indicates that
bulls started to shed velvet as early, as August 13 and Murie. (1951)
indicates that infrequently some bulls started shedding during, the
first week of August. A rising interest in rut-related activities
was noted with the response of two bulls which were "bugled in" on
August 8, 1973.
Murie (1951) indicates that the earliest observation
of herding ever noted was on August 15, while during this study it
was observed once in late July but became common during mid August.
Wallowing and bugling began in mid August.
Struhsaker (1967) noted
that overt response by other elk to the wallowing of a bull never
occurred, but during this study, the wallowing activities of a fivfe
-52point. bull appeared to aggravate a six point bull who displaced.the
smaller one from the .wallow with antler threats and pursued him into '
the timber.
was sparring.
The last activity observed in this chronological sequence
Struhsaker (1967) observed that squeals, during sparring
encounters, were only made by two and- one-half year old bulls.
During
this study, bulls two and one-half years of age and older.made
squealing noises.
The rising antagonism of adult bulls was first
expressed toward calves; Espmark
(1964) notes that in. caribou, calves
often take the brunt of bull antagonisms.. Spikes were next to receive
bullying, but as Struhsaker'(1967) notes, they were tolerated in the
harem group longest, compared to other males.
Spikes do not normally.
take part in rut activities (Murie, 1951), but this is possibly because
such activities are precluded by the vigilance of the harem bull.
Elk moved into the timber.in late summer and stayed for the
duration of the’rut.
Craighead
et'al. (1973) monitored cow. elk
fitted with radios, and noted that they were "difficult to observe"
because they "remained relatively sedentary ;.. in dense timber until
the break up of harems .
With shifts in the hormonal balances of elk
during the rutting season (Altmann, 1952), physiological changes may
bring about behavioral modifications'.
Preoccupation with, rutting
activities may reduce the usual wariness of elk, and consequently the
use of more secure habitats during the rut would have a selective
advantage.■
-53Several authors indicate that desiccation of vegetation in high
elevation parks may be responsible for this altitudinal migration to
timbered areas containing "succulent vegetation".
Yet., data collected
from elk feeding sites indicate that during the sampling period from
mid July to mid October, elk were able to select meadow sites which
were superior in moisture content to both established meadow and
timber plots at all three elevations.
During this study, it appeared that elk were selecting sites with
regard to forbs.
Throughout the summer, forbs at elk feeding sites
appeared to be superior to grasses based on desiccation rates, relative
standing crop weights and percentage moisture contents.
Rouse (1957)
and Stevens (1966) indicate that elk food habits undergo a transition
from forbs to grasses in September, however, Knowles (1975) indicates
that under conditions of a moist year, this transition is delayed.
If it was necessary for the elk in the Yellow Mule area to make the
conversion from forbs to grasses, it is logical that they would move
to timbered areas in October when the percentage moisture content of
grasses exceeded that of forbs, but.elk moved into the timber in
September when grasses were still inferior in percentage moisture .
content and. standing crop to forbs.
At elk feeding sites., the
standing crop.of forbs was generally greater than that of timber
grasses, and the percentage moisture content was consistently
significantly greater than that of timber grasses through Septembei.
-54Where elk have the ability to select high quality meadow areas
in which to feed, it would not be advantageous to move into, the timber
unless, such a move would benefit them in some other way.
This move­
ment coincident with the time of rut is evidence to support the
hypothesis of an increased security requirement of elk during this
time of year.
appendix
TABLE 12.
COMBINATIONS OF PRESSURE, CLOUD COVER, TEMPERATURE AND WIND TO PRODUCE SIX
WEATHER PHASES.1
Weather Code
Pressure
Cloud Cover
Temperature
Wind
1From:
I
2
3
4
5
6
High and
Stable
High and
Falling
Slightly
High,
Falling
Fast
Low,
Falling
Low,
Rising
High,
Rising
High
Fluffy
to Clear
Clear
Clouding,
Storm
Coming
Cloudy,
Stormy
Cloudy,
and
Breaking Up
Partly
Cloudy
and
Clearing
Cool and
Rising
Warm,
Fluctuating
and Rising
Hot and
Falling
Warm,
Rising
Slowly
Falling
to Cool
Cool
Rising
and
Fluctu­
ating
None to
Breezy
Breezy
Breezy to
Windy
Windy
Windy
Breezy
Landsberg, 1969, Weather and Health, An Introduction to Biometerology.
-57TABLE 13.
CONSTANCY AND OVERALL DOMINANCE VALUES OF SPECIES ASSOCI­
ATED WITH 189 AND 253 2 X 5 DECIMETER FRAMES MEASURED AT
FEEDING AND BEDDING SITES, RESPECTIVELY, OVER A FOURTEEN
WEEK PERIOD. Only species with less than twenty percent
constancy are listed.
Species
Feeding Sites
C1
D2
Bedding Sites
C
D3
19
5
10
7
7
18
7
7
4
7
7
14
4
11
11
4
—
tr4
—
—
tr
—
—
I
2
—
tr
I
—
7
11
7
4
4
4
14
18
4
11
—
11
4
7
4
—
—
tr
—
4
4
18
7
18
—
FORBS
Aaquilegia flaveecene
Agoserie aurantioa
Androsaoe ocoidentalis
Anemone multifida
Angelica pinnata
Antennaria alpina
Antennaria raaemosa
Antennaria rosea
Antennaria spp.
Arenaria congesta
Arnica diversifolia
Arnica latifolia
Arnica longifolia
Arnica mollius
Aster spp.
Aster conspicuus
Astragalus dasyglottis
Astragalus spp.
Besseya wyomingensis
Caltha leptosepala
Campanula rotundifolia
Castilliga miniata
Castillija pulchella
Cerastium beeringiana
Collinsia parviflora
Collomia linearis
I
I
—
10
5
14
14
14
—
—
—
14
10
10
10
—
—
10
5
—
I
2
I
I
3
10
—
5
—
Cruciferae spp.
Delphinium occidentale
Descurainia pinnata
Epilobium alpinum
Erigeron compositus
Erigeron formosissimus
Erigeron peregrinus
Erigeron speciosus
Erigeron ursinus
19
10
10
19
I
—
—
—
I
—
—
—
—
—
tr
—
—
—
I
—
3
-58TABLE 13.
Continued.
Species
Feeding Sites
C
D
Bedding Sites
C
D
19
14
19
tr
—
—
5
14
—
14
4
14
4
4
18
14
4
11
tr
—
tr
—
—
2
tr
—
tr
18
7
7
4
4
4
4
I
—
—
tr
tr
—
—
FORBS (Continued)
Frasera speciosa
Galium aparine
Galium boreale
Galium triflorum
Gentiana amarella
Geranium riohardsonii
Geum triflorum
Goodyera oblongifolia
Habenaria dilitata
Haeoklia floribunda
Heraoleum lanatum
Hieraoium graoile
Hieraoium oynoglossaides
Hydrophyllum oapitatum
Hymenoxys gradiflora
Lewisia pygmaea
Lingustiaum angustifolium
Lingustiaum filioinim
Linnaea borealis
Linum perenne
Lithophragma parviflora
Lomatium aous
Lupinus sulphurous
Lychnis drummondii
Mertensia oblongifolia
Miorosteris nigresoens
Miorosteris gracilis
5
5
——
5
—
5
5
—
—
5
—
5
10
—
—
4
Mushrooms
Osmorhiza depauperata
Oxytropis besseyi
Pedioularis oystopteridifolia
Pedioularis groenlandioa
Penstemon spp.
Perideridia gairdneri
Phaoelia heterophylla
Phaoelia serioea
Phlox multiflora
5
14
—
14
5
14
10
5
—
I
I
—
—
—
—
4
4
7
11
4
—
—
—
—
tr
7
4
18
4
—
—
tr
—
7
18
14
4
——
—
tr
—
7
7
tr
I
-59TABLE 13.
Continued.
Species
Feeding Sites
C
D
Bedding Sites
C
D
FORBS (Continued)
Potentilla divevsifolia
Pyrola asarifolia
Rorippa obtuea
Rudbeokia oooidentalie
Rumex pauoifolius
Saxifraga arguta
Saxifraga montanensis
Sedum lanoeolatum
Seneoio integerrimus
Seneoio 8erra
Sibbaldia prooumbene
Smilaoena stellata
Stellaria crispa
Thlaspi fendleri
Toimsendia parryi
Unidentified Forbs
Urtioa dioioa
Valeriana dioioa
Valeriana edulis
Veronica amerioana
Veronica serpyIlifolia
Veronica wormskjoldii
Zygadenus elegans
5
—
5
5
—
—
10
5
10
14
5
14
5
19
5
5
5
19
—
—
—
tr
—
4
—
—
—
—
I
.
7
18
I
2
18
tr
4
11
—
—
4
4
4
14
4
—
tr
—
—
—
4
14
4
14
—
tr
—
tr
14
tr
7
11
7
tr
—
tr
7
4
tr
—
GRASS AND GRASS-LIKE PLANTS
Agrostis exarata
Agrostis soabra
Bromus seoalinus
Calamagrostis canadensis
Danthonia intermedia
Desohampsia atropurpurea
Elymus glauous
Festuoa ovina
Glyoeria grandis
Hordeum braohyantherum
Junous spp.
Luzula wahlenbergii
Poa ousiokii
10
—
5
14
5
5
10
I
tr
—
I
—
14
19
tr
—
14
5
tr
—
-60TABLE 13.
Continued.
Species
Feeding Sites
D
C
Bedding Sites
C
D
10
19
19
19
19
10
14
19
tr
—
tr
tr
tr
tr
tr
—
14
—
14
—
7
11
—
tr
10
5
—
7
11
7
4
tr
—
—
—
10
—
5
—
4
4
7
11
7
—
—
—
——
—
5
—
10
5
—
—
14
—
GRASS AND GRASS-LIKE PLANTS
(Continued)
Poa glauoifolia
Poa junoifolia
Poa leptoooma
Poa nemoralis
Poa pratensis
Poa spp.
Poa stenantha
Stipa oocidentalis
SHRUBS
Artemesia tridentata
Berberis repens
Juniperus oommunis
Lonioera utdhensis
Prunus virginiana
Ribes amerioanum
Ribes laoustre
Rosa spp.
Salix montioola
Sambuous raoemosa
Spiaraea betulifolia
Symphorioarpos oooidentalis
Vacoinium sooparium
tr
1C = Constancy, percent occurrence among all sites.
2D = Dominance Value, percent occurrence of the species as a dominant
within 189 2 X 5 decimeter frames analyzed on feeding sites.
3D = Dominance Value, percent occurrence of the species as a dominant
within 252 2 X 5 decimeter frames analyzed on bed sites.
^tr = trace, species occurs as a dominant on less than one percent
of the plots.
-61TABLE 14.
STANDING CROP AND PERCENTAGE MOISTURE CONTENT OF VEGETATION
ON TWENTY-ONE ELK FEEDING SITES.
Standing Crop1
Forbs
Grass
Percentage Moisture Content2
Forbs
Grass
Mid July
82.0
172.0
119.0
194.0
35.0
91.5
67.5
96.0
379.9
263.4
328.6
332.2
207.1
210.4
205.2
196.4
Late July
197.0
170.0
164.0
213.0
44.0
90.0
66.0
150.0
376.1
368.2
261.0
422.1
200.0
261.1
139.4
250.7
Mid August
237.5
149.0
150.0
150.5
57.5
246.0
82.0
84.5
323.4
341.3
299.0
243.5
157.4
199.4
161.6
165.7
Late August
156.0
136.0
140.5
90.0
2.5
95.0
73.0
200.0
410.9
329.0
140.0
134.4
280.0
182.1
102.1
70.8
180.0
169.0
114.0
31.0
15.5
171.0
298.3
139.1
214.5
156.5
87.1
123.1
61.5
40.5
63.5
53.5
71.5
137.0
40.2
31.4
Mid September
Mid October
1Dry weight standing crop per square meter.
2Percent moisture content = (Wet weight-Dry weight)/Dry weight X 100
-62TABLE 15.
STANDING CROP AND PERCENTAGE MOISTURE CONTENT OF VEGETATION
ON SIX ESTABLISHED PLOTS AT THREE ELEVATIONS.
Standing, Crop1
Meadow
Timber
Forbs Grass Forbs Grass
Percentage Moisture Content2
Meadow
Timber
Forbs Grass Forbs Grass
2200 Meters
Mid July
Late July
Mid August
Late August
Mid September
Late September
Mid October
310.0
178.0
125.3
139.8
188.5
81.0
77.8
97.0
50.5
19.5
40.0
59.8
76.3
39.3
25.3
23.0
20.0
18.5
15.0
10.0
8.0
24.5
21.3
24.5
23.8
41.0
37.3
25.5
230.8 106.7
153.4 77.2
120.0 92.9
69.4 80.6
13.8 28.0
7.4 17.0
9.3 13.4
387.1 134.7
241.3 107.1
213.8 171.4
171.6 78.9
88.3 66.5
67.5 55.0
25.0 45.1
321.0 179.6
173.3 123.9
164.7 144.2
86.6 93.0
44.2 41.8
11.9 30.2
2.2
1.8
268.3 150.0
278.8 88.9
366.1 65.1
245.5 95.0
98.2 70.0
53.1 37.5
47.5 51.9
201.1 202.4
336.1 156.7
245.3 121.7
178.8 97.6
57.9 36.7
27.1 33.3
18.2 25.0
471.2
265.3
318.4
231.6
77.6
64.1
20.5
2500 Meters
Mid July
Late July
Mid August
Late August
Mid September
Late September
Mid October
124.0
138.5
94.3
58.0
47.5
102.8
42.3
111.5
113.0
110.8
104.0
94.5
95.3
78.5
67.0
49.5
47.3
33.5
27.5
24.0
30.0
2.5
9.0
10.8
10.0
2.5
2.0
6.8
2800 Meters
Mid July
Late July
Mid August
Late August
Mid September
Late September
Mid October
87.5
41.5
47.5
47.3
35.0
38.8
22.0
21.0
30.0
30.0
31.5
39.5
30.8
20.0
26.0
50.5
43.5
24.5
21.3
19.5
18.3
4.8
4.0
4.0
6.5
7.0
6.0
5.8
194.7
175.0
131.3
115.4
64.3
54.2
39.1
1Dry weight standing crop in grams per square meter.
^Percentage Moisture Content = (Wet weight-Dry weight)/Dry weight X 100.
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MONTANA STATE UNIVERSITY LIBRARIES
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