Range relationships of mule deer, elk and cattle on a rest-rotation grazing system during winter and
spring
by Thomas James Komberec
A thesis submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE
in Fish and Wildlife Management
Montana State University
© Copyright by Thomas James Komberec (1976)
Abstract:
A study was conducted in the timbered breaks adjacent to the Missouri River, northcentral Montana,
during the summer of 1974 and the winter and spring of 1975 to obtain quantitative data on
populations, range use and food habits of mule deer, elk and cattle within an area managed by
rest-rotation grazing. Seven major habitat types consisting of eleven plant communities were
recognized. Numbers, productivity, distribution, and range use of mule deer and elk were determined
from one early winter helicopter purvey, two fixed-wing airplane surveys and regular ground
observations. The fawn:doe ratio was 54.8 for mule deer and the cow:calf ratio was 70.0 for elk in late
January 1975. Numbers and distribution of mule deer during the study showed no consistent trends in
relation to grazing by cattle and pasture treatments. Numbers and distribution of elk during this study
were greatly influenced by grazing of cattle. Elk moved from areas of previous use when cattle began
using the area. Home ranges of four marked mule deer were largest for the two adult males and
smallest for the female and male fawn. Three radio-collared elk had home ranges much larger than
those of the mule deer, with the male having a home range more than twice as large as the home ranges
of two female elk.
The Artemisia-Agpopyron habitat type was used most often by both mule deer and elk during winter
and spring and by cattle during spring.
Marked mule deer and elk used the Pinus-Juniperus habitat type most often during both winter and
spring. Mule deer, elk and cattle all used 0-10 degree slopes most often during winter and spring. Mule
deer and elk preferred southerly exposures during both seasons while cattle used ridge tops and coulee
bottoms most often. Food habits were determined from feeding site examinations, supplemented with
one cow rumen sample. Browse, forbs, grasses and forbs, browse, grasses was the order of importance
of forage classes used by mule deer in winter and spring, respectively. Forbs, grasses, browse was the
order of importance of forage classes used by elk in winter. Grasses, forbs, browse was the order of
importance of forage classes used by cattle during spring. Yellow sweetclover was the most important
forb in the diet of each of the three ungulates. Western wheatgrass was the most important grass in the
diet of both elk and cattle. Interspecific relationships, effect of rest rotation grazing and management
recommendations of mule deer, elk and cattle on a rest rotation grazing system were discussed. STATEMENT OF PERMISSION TO COPY
/
In presenting this thesis in partial fulfillment of the require­
ments 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
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.
Signature
Date.
RANGE RELATIONSHIPS.OF MULE DEER,. ELK AND CATTLE ON A RESTROTATION GRAZING SYSTEM DURING WINTER AND SPRING
by
THOMAS JAMES KOMBEREC
.A thesis submitted in partial fulfillment
of the requirements for the degree
of
MASTER OF SCIENCE
in
Fish and Wildlife Management
.MONTANA STATE UNIVERSITY
Bozeman, Montana
March, 1976
iii
ACKNOWLEDGMENT
The author wishes to express his sincere appreciation to the
following, among others, for their aid and contributions to this study:
Dr. D. C. Quimby and Dr. R. J. Mackie, Montana State University, for
project planning, technical advice and guidance in preparation of the
manuscript.; Dr. W. R. Gould and Dr. R. L. Eng, Montana State University,
for critical review of the manuscript; Dr. J. H. Rumely, Montana State
University, for.verification of plant specimens and for critical review
of the manuscript; Mr. R. Trueblood, Mr. R. B. Campbell, and other
personnel of Region Six, Montana Department of Fish and Game, for field
assistance and cooperation; Mr. J. Jones, and other personnel of the
Bureau of Land Management, Malta District Office, for their cooperation;
and to the Robinson brothers of the Lazy J. D. Cattle Company for their
assistance and cooperation.
Last, but not least, to my parents, family and friends, for their
unstinting support through the long years of academic pursuit, I wish to
express my heartfelt thanks.
During the study, the author was supported by the Montana
Department of Fish and Game under Federal Aid Project W-120-R-6 and
W-120-R-7.
iv
TABLE OF CONTENTS
Page
V I T A ...........................................................
ii
ACKNOWLEDGMENT ................................................
iii
LIST OF T A B L E S ................................................
vi
LIST OF F I G U R E S ..............................................
x
A B S T R A C T .......................................................
xi
INTRODUCTION ..................................................
I
DESCRIPTION OF THE STUDY A R E A ...............................
3
6
7
OO OO
Artemisia-Agropyvon Habitat Type .......................
Pinus-Juniperus Habitat Type
...........................
Pseudotsuga-Juniperus Habitat Type
Saroobatus-Agropyron Habitat Type
Artemisia longifolia Habitat Type .......................
Agropyron-Symphorioarpos Habitat Type ...................
Xanthium strumarium Habitat Type
.......................
9
9
9
M E T H O D S .......................................................
11
R E S U L T S .......................................................
14
Mule Deer Populations and Range U s e .....................
14
Population Characteristics .........................
Home Range and M o v e m e n t s ...........................
Group Size and Characteristics.....................
Activity Habits and Patterns .......................
Use of Habitat Types, Slopes, and Exposures
. . . .
14
18
24
25
25
Elk Populations and Range U s e ...........................
Population Characteristics .........................
Home Range and M o v e m e n t s ...........................
Group size and Characteristics.....................
Activity Habits and Patterns .......................
Use of Habitat Types, Slopes, and Exposures
.. . .
30
30
33
39
40
42
V
TABLE OF CONTENTS
(Continued)
Page
Cattle Populations and Range Use ........
. . . . ....
.
Numbers and Distribution ............................
Group Characteristics............
Activity Habits and Patterns
. ......................
Use of Habitat Type, Slope, and Exposure . . . . . . .
Food H a b i t s ............
46
49
50
50
.53
Mule D e e r ............ .. . . . . . ............... ..
Elk . . ............... ' ................. ............
C a t t l e ...................
D I S C U S S I O N ..............................................
Interspecific Range Relationships
46
53
55
57
61
.......................
61
Mule Deer and Cattle ................................
Elk and Cattle
. .....................................
Mule Deer and E l k ....................................
63
65
Rest-Rotation Grazing Versus Season-Long Grazing . . . . .
66
67
MANAGEMENT R E C O M M E N D A T I O N S .............' ........... ...........
70
A P P E N D I X ........................................
71
LITERATURE CITED
77
vi
LIST OF TABLES
Table
1.
2.
3.
Page
MEAN TEMPERATURE AND PRECIPITATION FOR 1974 AND JANUARYJUNE 1975., SNOWFALL FOR 1975, AND. 11-YEAR MEANS. U. S.
DEPARTMENT OF COMMERCE WEATHER STATION ROY 24 NE '■
(MOBRIDGE), MONTANA .................................
MULE DEER NUMBERS BY SEX, AGE CLASS AND LOCALITY
DURING.JANUARY 1975 HELICOPTER SURVEYS OF
NICHOL'S COULEE RCA . ............
16
SUMMARY OF HOME RANGES AND MOVEMENTS DURING WINTER AND
SPRING FOR THREE RADIO-COLLARED AND ONE EAR-TAGGED
MULE D E E R ............
20
4.. MONTHLY AND SEASONAL FREQUENCIES OF MULE DEER GROUP
SIZES AND MEAN GROUP SIZE DURING WINTER AND SPRING . . .
5.
6.
7.
5
24
SEASONAL PERCENTAGES OF ALL MULE DEER ACTIVITIES
OBSERVED WITHIN EACH HABITAT TYPE AND TOTAL SEASONAL
AVERAGES; AND MONTHLY PERCENTAGES OF ACTIVITIES FOR
ALL MULE DEER ON ALL HABITAT TYPES
......................
26
PERCENTAGES OF OBSERVATIONS OF 4 MARKED MULE DEER BY
ACTIVITY CLASS, HABITAT TYPE, EXPOSURE, AND SLOPE
DURING WINTER AND SPRING ................. . . . . . . .
27
MONTHLY AND SEASONAL PERCENTAGES FOR ALL USE OF HABITAT
TYPE BY MULE DEER OBSERVED, FOR EACH HABITAT TYPE AND
FOR EACH COMMUNITY WITHIN A HABITAT TYPE. TRACE (T)
AMOUNTS < 1 % ..................... .........................
28
8 . MONTHLY AND SEASONAL PERCENTAGES OF ALL USE OF SLOPES
BY MULE DEER OBSERVED DURING WINTER AND SPRING
' 9.
10.
. . . . .
29
MONTHLY AND SEASONAL PERCENTAGES OF ALL USE OF
EXPOSURES BY ALL MULE DEER OBSERVED DURING WINTER
AND S P R I N G ..............................................
31
NUMBERS AND SEX AND AGE COMPOSITION OF ELK OBSERVED ON
THE NICHOL'S COULEE RCA DURING HELICOPTER SURVEYS
■ ■JANUARY 1975
32
vii
LIST OF TABLES
(Continued)
Table
11.
12.
13.
14.
15.
16.
17.
Page
NUMBERS OF RELOCATIONS, HOME RANGE SIZES, AND
PERCENTAGES OF RELOCATING WITHIN A GIVEN AREA DURING
WINTER AND SPRING FOR THREE RADIO-COLLARED ELK ON
THE NCRCA . . '........................... ................
37
MONTHLY AND SEASONAL FREQUENCIES OF ELK GROUP SIZES AND
MEAN GROUP SIZE FOR ALL ELK OBSERVED DURING WINTER AND
SPRING; AND SEASONAL FREQUENCIES AND MEAN GROUP SIZE
OBSERVED FOR THREE RADIO-COLLARED ELK DURING
WINTER/SPRING ........... ■............................ .
40
SEASONAL FREQUENCIES OF ALL ELK ACTIVITIES OBSERVED ■
WITHIN EACH HABITAT TYPE AND TOTAL SEASONAL AVERAGES;
AND MONTHLY PERCENTAGES OF ACTIVITIES FOR ALL ELK ON
ALL HABITAT T Y P E S ........................... ............
41
PERCENTAGES OF OBSERVATIONS OF 3 RADIO-COLLARED ELK BY
ACTIVITY CLASS, HABITAT TYPE, EXPOSURE AND SLOPE
» DURING WINTER AND SPRING ' ..................... ' .........
43
MONTHLY AND' SEASONAL PERCENTAGES FOR ALL USE OF HABITAT
TYPES BY ELK OBSERVED DURING WINTER AND SPRING FOR EACH
HABITAT TYPE AND FOR EACH COMMUNITY WITHIN A
HABITAT TYPE . ■..........................................
44
FREQUENCIES OF ALL USE OF SLOPES BY ELK OBSERVED DURING
WINTER AND SPRING BY. MONTH AND S E A S O N ................. ..
46
FREQUENCIES OF ALL USE OF EXPOSURE BY ELK DURING WINTER
AND SPRING BY MONTH AND S E A S O N .........................
47
18.
FREQUENCIES OF CATTLE GROUP SIZE CLASSES; AND MEAN GROUP •
SIZES DURING SPRING BY MONTH AND SEASON
...............
49
19.
PERCENTAGES OF ALL ACTIVITIES RECORDED FOR CATTLE
OBSERVED'DURING SPRING BY MONTH AND SEASON .............
50
viii
LIST OF TABLES
(Continued)
Table
20.
-
Page
FREQUENCIES OF ALL.CATTLE ACTIVITIES OBSERVED DURING ■
SPRING FOR EACH HABITAT TYPE AND TOTAL CATTLE USE OF
EACH HABITAT TYPE. TRACE (T) AMOUNTS < 1 % .......... .. .
51
-PERCENTAGES OF.USE OF SLOPES BY ALL CATTLE OBSERVED
DURING; SPRING BY MONTH AND SEASON.. TRACE (T) '
AMOUNTS < 1 % ..................... '............... .
51
FREQUENCIES OF USE OF EXPOSURE BY CATTLE OBSERVED DURING
SPRING BY MONTH AND SEASON. TRACE (T) AMOUNTS <1%
. .
52
AGGREGATE MEAN
FOR EACH PLANT
SPRING. TRACE
SITES.
SPRING
PERCENTAGES AND FREQUENCY OF OCCURRENCE
SPECIES USED BY MULE DEER, WINTER AND
(T) AMOUNTS ARE <1%. WINTER - 6 FEEDING •
- 3 FEEDING SITES
...............
54
24. . AGGREGATE MEAN PERCENTAGES AND FREQUENCY OF OCCURRENCE
FOR EACH PLANT SPECIES USED BY ELK DURING WINTER AT
NINE FEEDING SITES EXAMINED. TRACE (T) AMOUNTS <1% . . .
56
21.
22.
23.
25.
'
AGGREGATE MEAN PERCENTAGES AND FREQUENCY OF OCCURRENCE
FOR EACH PLANT SPECIES USED BY CATTLE DURING SPRING AT
..SIX FEEDING SITES EXAMINED . .'.........................
58
26. PERCENTAGE OF RUMEN CONTENTS OBTAINED FROM ONE CATTLE •
: RUMEN DURING JUNE 1975 ‘.......... ............ ..
27.
28.
LAND STATUS AND GRAZING CAPACITY OF NICHOL'S
COULEE R C A .......... ............................... ..
60
.
MEAN PERCENTAGE CANOPY COVERAGE (C) AND FREQUENCY OF
OCCURRENCE (O) OF PLANT TAXA WHICH OBTAINED A MEAN
COVERAGE OF I PERCENT OR MORE IN ONE OR MORE OF THE
ELEVEN PLANT COMMUNITIES DURING SUMMER AND.FALL.
VALUES FOR BARE GROUND AND LITTER ARE ALSO INCLUDED.
TRACE .(T). AMOUNTS .ARE FOR VALUES BETWEEN•I PERCENT .AND
0.1 PERCENT.
NUMBERS OF SITES EXAMINED ARE IN
PARENTHESIS
................ .............. • . .
72
75
ix
LIST OF TABLES
(Continued)
Table
29.
■
SUMMARY OF THE GRAZING SEASONS FROM 1972 TO .1974 FOR .
EACH PASTURE AND TOTALS FOR NICHOL'S COULEE RCA
Page
76
X
LIST OF FIGURES
Figure
1.
2.
3.
4.
Page
Map of the study area showing boundaries and
drainages ........................ .................. ..
4
Distribution of mule deer January 1975-helicopter
s u r v e y ....................... .. . . . ............ .
15
Distribution of mule deer on Nichol's Coulee RCA
during winter 1975 ......................................
17
Distribution of mule deer on Nichol's Coulee RCA
during spring 1975 ............... .................. ; .
19
5.
Home ranges of three radio-collared and one ear-tagged
mule deer on Nichol's CoUlee RCA during winter and
spring 1975 . ............... 22
.6 .
Distribution of elk on Nichol's Coulee RCA during
winter 1975 .'..................... .. . . ............. '.
7.
8.
Distribution of elk on Nichol's Coulee RCA during
spring 1975 . . . . . . . .
.......................... ..
34
.
35
Home ranges of three radio-collared elk on Nichol's
Coulee RCA during winter and spring 1975 ...............
38
Distribution of cattle on Nichol's Coulee RCA during
spring 1975 ............................................ ..
48
10.
Grazing formula for Nichol's Coulee RCA
...............
73
11.
Order of treatments for pastures in Nichol's Coulee
RCA within and between years
.. ............. .......... ..
74
9.
ABSTRACT '
A study was conducted in the timbered breaks adjacent to the
Missouri River, northcentral Montana, during the summer of 1974 and
the winter and spring of 1975' to obtain quantitative data on popu­
lations, range use and food habits of mule deer, elk and cattle within
an area managed by rest-rotation grazing. Seven major habitat types
consisting of eleven, plant communities were recognized. Numbers,
productivity, distribution, and range use of mule deer and elk were
determined from one early winter helicopter purvey, two fixed-wing
airplane surveys and regular ground observations. The fawn:doe ratio
was 54.8 for mule deer and the cow:calf ratio was 70.0 for elk in late
January 1975. Numbers and distribution of mule deer during the study
showed no consistent trends in relation to grazing by cattle and
pasture treatments.
Numbers and distribution of elk during this study
were greatly influenced by grazing of cattle. Elk moved from areas of
previous use when cattle began using the area. Home ranges of four
marked mule deer were largest for the two adult males and smallest for
the female and male fawn. Three radio-collared elk had home ranges
much larger than those of the mule deer, with the male having a home
range more than twice as large as the home ranges of two female elk.
The Artem-is-ia-Agrppyvon habitat type was used most often by both mule
deer and elk during winter and spring and by cattle during spring.
Marked mule deer and elk used the P-inus-Juniperus habitat type most
often during both winter and spring. Mule deer, elk and cattle all
used 0-10 degree slopes most often during winter and spring. Mule deer
and.elk preferred southerly exposures during both seasons while cattle
used ridge tops and coulee bottoms, most often.
Food habits were deter­
mined from feeding site examinations, supplemented with one cow rumen
sample. Browse, forbs, grasses and forbs, browse, grasses was the order
of importance of forage classes used by mule deer in winter and spring,
respectively.
Forbs, grasses, browse was the order of importance of
forage classes used by elk in winter.
Grasses, forbs, browse was the
order of importance of forage classes used by cattle during spring.
Yellow sweetclover was the most important forb in the diet of each of
the three ungulates.. Western wheatgrass was the most important grass
in. the diet of both elk and cattle.
Interspecific relationships, effect
of rest rotation grazing and management recommendations of mule deer,
elk and cattle on a rest rotation grazing system were discussed. .
INTRODUCTION
Rest-rotation grazing (Hormay and .Talbot 1961) has been
increasingly employed by range,and other land managers' to relieve
overgrazing and improve vegetation while maintaining maximum production
of livestock on rangelands.
The benefits of rest-rotation grazing for
livestock and rangeland vegetation have been well documented (Hickey
1966).
Little is known as yet. of the possible effects or influences
\
of rest-rotation grazing of livestock on habitat values for wildlife
or on wildlife-livestock relationships.
During the summer of 1973, the Montana Department of Fish and
Game initiated a study of the range relationships of mule deer
(Odocoileus hemionus), elk (C e m u s .canadensis), and cattle (Bos
taupus) on the. Nichol's Coulee Resource Conservation Area, a restrotation grazing system for cattle in northcentral Montana (Knowles
1975).
The objectives of.the study were to obtain quantitative data
on mule deer, elk, and cattle distribution, movements,. range use, and
food habits within the Nichol's Coulee Area, and to establish basic
criteria to assist range and wildlife managers, in planning and
conducting sound multiple-use management on a rest-rotation grazing
■'
.
. ' ,
system. . Knowles (1975) described range relationships of mule deer,
elk, and cattle on the area during summer and fall.
My study,
conducted full-time during the summer of 1974 arid.from January.through
•I .
* 4 « ■’
-2June, 1975, considered these relationships during winter and spring.
DESCRIPTION OF THE STUDY AREA
The 88,810 acre Nichol's .Coulee RCA study area (Fig. I) is
located in Phillips County, about 55 miles southwest of Malta,
Montana.
Knowles (1975) described the general physiographic and
climatic characteristics of this area as well as the primary features
of the rest-rotation grazing system.
Ownership of land and grazing
capacities are listed in Appendix Table 27.
The grazing formula for
the study area is shown in Appendix Figure 10, and order of treatments
within and between years is given in Appendix Figure 11.
Climatological data for the period January through June, 1975
(Table I) were obtained from the U. S . Department of Commerce Weather
Station Roy 24 NE (Mobridge), located 17 miles southwest of the study
area.
Except for January, the mean monthly temperatures were an
average of 3.62° F less than the. corresponding 11 year monthly means;
the range of deviation of monthly temperatures from normals was minus
1.0 for May to minus 6.8 for February.
Northwest winds averaging 5
to 10 miles-per-hour with gusts to 30 miles-per-hour were common during
winter and early spring.
Eighty-four percent of the 41.5 inches of
snowfall from January through June 1975 occurred during March and
April, while 85.5 percent of total precipitation for the same period
fell during April, M a y , and June.
—4—
P ast. Il
P a st. I
C M .R ussall H
G a m e R an g e
P ast. Ill
M isso u ri R iver
L egend
N ichols C o u le e RCA
C ree k o r C o u lee -----H o lding p a s t u r e ------------ hp
Figure I.
Map
of the study area showing boundaries and drainages.
-5TABLE I.
MEAN TEMPERATURE AND PRECIPITATION FOR 1974 AND JANUARY-JUNE
1975, SNOWFALL FOR 1975, AND Il-YEAR MEANS. U. S. DEPARTMENT
OF COMMERCE WEATHER STATION ROY 24 NE (MOBRIDGE), MONTANA.
Month
Temperature (° F)
Year
11-Year1
1974
1975
Mean
Precipitation (inches)
Year
Snowfall
Il-Year
1974
1975
Mean
1975
Jan.
Feb.
March
April
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
17.5
29.7
33.1
46.9
50.4
68.0
73.7
64.8
55.6
48.2
34.5
26.9
21.4
15.3
29.1
38.4
54.1
62.4
—
—
—
—
—
—
11.7
27.1
31.8
44.1
55.1
64.3
71.5
70.7
57.6
47.1
31.2
17.3
.12
.10
.83
.81
4.44
2.08
4.82
4.09
.61
.65
.41
.04
.26
.19
.98
2.18
3.21
3.07
—
—
—
—
—
—
1.0
5.5
19.5
15.5
0
0
—
—
—
—
—
—
.58
.21
.57
1.36
2.02
2.80
1.69
1.90
1.06
.58
.32
.43
Totals
45.8
36.78
43.8
19.00
9.89
41.5
13.52
-No data January and February 1964.
Knowles (1975) reported that seven of the eight habitat types
described by Mackie (1970) for the Missouri River breaks, occur on the
Nichol's Coulee RCA study area.
A summary of the general occurrence
and vegetational characteristics of these habitat types and the 11
plant communities which currently occur on each as described in detail
by Knowles (1975) is presented below.
Mean percentage canopy coverage
and frequency of occurrence of plant taxa found in each of the 11
communities during summer and fall are shown in Appendix Table 28.
-6ATtemisia-Agvopyron Habitat Type
This habitat type occupied approximately 57 percent of Pastures
III and IV and almost all of Pastures I and II.
It was comprised of
three distinct plant communities as follows:
The Artemisia tridentata-Agropyron spicatum community was well
developed along edges of main ridges, on the tops of side ridges and
oh the more level portions of ridge sides.
Big sagebrush .(Artemisia
tridentata), bluebunch wheatgrass (Agropyron spicatum), and yellow
sweetclover (Melitotus officinalis) was the most common shrub, grass,
and forb, respectively.
These taxa were also dominant species recorded
at ungulate feeding sites during winter and spring.
While the Artemisia tridentata-Agropyron smithii community was
best represented on the main ridges, it was also found on sites similar
to the Artemisia tridentata-Agropyron spicatum community.
Western
wheatgrass (Agropyron smithii), big sagebrush, and pale bastard toad­
flax (Comandra umbellata) was the dominant grass, shrub, and forb,
respectively.
Big sagebrush and western wheatgrass were feeding site
dominants on this community during winter.
These taxa arid pale bastard
toadflax were dominant on ungulate feeding sites recorded during spring.
The Artemisia tridentata-Agropyron smithii-Bouteloua gracilis
community was the dominant community of Pastures I and II, but was
restricted to level ridge tops in Pastures III and IV.
Blue grama
('Bouteloua gracilis) and western wheatgrass were the dominant grasses.
-7
Big sagebrush and' fringed sagewort (Artemisia fvigida) was the most
frequent shrub and forb, respectively.
The above taxa were the most
often recorded as dominants at feeding sites during winter and spring.
However,. forks of the Tragopogon dubius union (Mackie 1970) increased
in importance as spring progressed.
Finus-Juniperus Habitat Type
Characteristic of sloping ridge sides, with development determined
by degree of slope and exposure, this habitat type occupied 26 percent
of the area.
Distribution was confined mostly to Pastures III and IV.
Three plant communities were characteristic:
The Pinus ponderosa-Agropyron spioatum community occurred at the
borders between the Pinus-Juniperus type and Avtemisia-Agvopyron
dominated ridge tops, in shallow basins at the heads of drainages, and
on narrow side-ridge tops with scattered ponderosa pine (Pinus
pondevosa).
Rocky Mountain juniper (Junipevus soopulovum), bluebunch
wheatgrass, and yellow sweetclover was the dominant shrub, grass, and
forb, respectively.
Winter feeding site examinations showed these
plant taxa to be dominant then.
The Pinus pondevosa-Juhipevus'scopuldrum community was best
developed on slopes with some north exposure, with decreasing frequency
of occurrence toward south exposures.
juniper, were characteristic.
Dense thickets of Rocky Mountain
Green needlegrass (Stipa vividula),
yellow sweetclover, and juniper had the greatest coverage for grasses.
-8forbs, and shrubs, respectively.
The Pinus pondevosa-Artemisia longifolia community occurred on
steep, unstable shale slopes usually, with a south aspect.
Prairie
sand reedgrass ,(Calamovilfa longifolia) was the dominant grass.
Long-
leaf sagebrush (Artemisia longifolia) was the most frequent shrub.
Yellow sweetclover was the dominant forb, although soapweed '(Yueea
glauea) was quite characteristic of this community.
Along with the
above taxa, western wheatgrass was recorded as a feeding site dominant
on this community during winter.•
Pseudotsuga-Juniperus Habitat Type
This minor habitat type, characteristic of steep northerly
exposures was restricted to Pastures III and IV where it covered one
percent of the area.
Maximum abundance was along the west edge of the
Study area, while, this type was absent on the ea!st boundary.
'Sarcobatus-Agropyron Habitat Type
About six percent of Pastures III and IV along the Missouri River
bottoms, coulee bottoms, and footslopes was covered by.this habitat
type.
Western wheatgrass was the dominant grass, especially in coulee
bottoms.
Greasewood (Sareobatus vermieulatus) was the dominant shrub
along the river and a short distance north in major coulees.
Big
sagebrush gradually increased in abundance with distance from the
river.
Yellow sweetclover was the most important forb.
These plant
—9taxa were also most often recorded in feeding sites during winter.
Artemisia longifolia Habitat Type
This habitat type occurred on unstable, steep south and southwest
shale slopes, being most conspicuous in the southern part of Pastures
III and IV, where it covered four percent of the area.
Longleaf sage­
brush and yellow sweetclover were dominant shrubs, and forbs, respectively
Western wheatgrass and prairie sand reedgrass were the dominant grasses.
Soapweed was also recorded as a feeding site dominant during winter.
Agropyron-Symphorioarpos Habitat•Type
Found in the flood plains of major coulees and.in minor drainages,
this habitat type combined with the Xanthium strumarium type occupied
approximately six percent of Pastures III and IV.
Western wheatgrass,
western snowberry (Symphorioarpos oooidentalis) and yellow sweetclover
was the predominant grass, shrub, and forb, respectively.
Green
rabbitbrush (Chrysothamnus viscidiflorus), silver sagebrush (Artemisia
oana), and wild licorice .(Glyoyrrhiza iepidota) yere. also recorded as
dominant taxa on ungulate feeding sites examined during winter and
spring.
-
'
.
Xanthium strumarium Habitat Type
This minor habitat type occurred in the actual cuts of inter­
mittent streams.
Vegetation was sparse and generally restricted to
silty banks, arid areas not severely flooded.
Cocklebur (Xanthium
-10
stmmariurri) was the characterizing species.
During winter, shrubs, dry grasses, and stems of persistent forbs,
such as yellow sweetclover, were the dominant vegetative forms in all
habitat types on the study area.
With the progression of spring, forbs
of the Tvagopogon and Poa unions, especially, contributed greatly to
the floral diversity of Nichol's Coulee RCA. .
METHODS
Habitat analyses were similar to those described by Knowles (1975).
Twenty-three permanent transects were established in stands repre­
sentative of six major habitat types during the summer of 1974.
Canopy
coverage (Daubenmire 1959) and frequency of occurrence of herbaceous
taxa were recorded among twenty 2 X 5 dm. plots on each transect; these
data for shrubs were recorded in superimposed 4 X 10 dm. plots (Pyrah
1973).
Supplementary data on plant composition in habitat types were
obtained by measuring coverage and frequency using ten 2 X 5 dm. and
ten 4 X 10 dm. plots for forbs and shrubs, respectively, at each
ungulate feeding site examined (Knowles 1975). . Plant nomenclature
followed Booth (1950) and Booth and Wright (1966).
Range usage was determined by recording observations of mule deer,
elk, and cattle along nine vehicle observation routes similar to those
established by Knowles (1975).
Eight of these routes were located
within Pastures III and IV of the Nichol's Coulee RCA.
Routes, were
covered systematically, at approximately weekly intervals, during
morning and evening.
Observations were recorded as to species, time,
group size, activity, habitat type, slope, exposure, and location.
Sex and age-class of mule deer and elk were recorded when possible.
The occurrence of any two or more of the three ungulate species within
a distance of one-eighth mile of each other was also noted.
—12—
Additional data on range usage were obtained by two aerial
surveys over the study area using a fixed-wing aircraft during January
and April, 1975, and an intensive helicopter survey of Pastures III
and IV during mid-January, 1975.
The latter also provided basic popu­
lation data for mule deer and elk on the study area.
Movements and home ranges of mule deer, and elk were determined by
relocations of five individually marked mule deer, including three
equipped with radio-transmitters, and three radio-transmitter equipped
elk.
Radio transmitters were applied by personnel of the Montana Fish
and Game Department during August, 1974 (Knowles 1975); two additional
mule deer were ear tagged and marked by Knowles during June, 1974.
Relocations of radio-equipped animals were usually made during mid-day;
those of ear-marked mule deer were made during normal observation
periods.
Home range areas of marked mule deer and elk were determined
by the modified minimum area method of Harvey and Barbour (1965);
Food habits were determined by examination of recently vacated
ungulate feeding sites to record frequencies of plant use by species
(Cole 1956).
One instance of use was defined as usage of an individual
leaf or rooted stem for grass and grass-like plants, an individual leaf
or twig for shrubs and trees, and individual leaves or stems for forbs.
A minimum of 50 instances of use were recorded at each feeding site.
Dominant plant, species were recorded at all feeding sites examined..
Food habits data were compiled using the aggregate percentage method
-13(Martin et at. 1946) by season and habitat type.
Supplementary data
on the food habits of cattle were obtained by examination of contents
of the rumen.of one cow found dead on,,the study area d.uring June,.
1975.
RESULTS
Mule Deer Populations and Range Use
Results are based on observations of 612 mule deer, comprising
119 separate groups during winter (January-March), 1975, and 225
individuals in 81 groups during spring (April I-June 15).
An
additional 201 mule deer were observed during helicopter surveys of
Pastures III and IV in January, 1975.
Population Characteristics
Knowles (1975) described mule deer population characteristics and
trends on the study area during early January of 1974 and 1975.
These
data indicated an increase of 9.2 percent in numbers of mule deer on the
area from 1974 to 1975. .The increase occurred entirely within Pasture
III, which had not been grazed by cattle during the previous grazing
season.
Mule deer densities varied between pastures and between drainages
within each pasture (Fig. 2, Table 2).
In January, 1975, the deer
density was 1.9 times greater in Pasture III than in Pasture IV.
Within Pasture III, the density of mule deer in the CK Creek drainage
was 3.5 times greater than in the Nichol's Coulee drainage.
Of the 612 mule deer observed from the ground during the winter of
1975, 82 percent occurred in Pasture III.
Most of these were found in
four extensive concentration areas (Fig. 3).
Three areas of concentrated
Past. IV
Iv S c /''
L eg en d
\
^
^ > 5 D e e r / s q . m i.
M isso u ri
R iv er
I - 5 D e e y sq . mi.
< ^1 D e e r/s q . m i.
N o t S u rv e y e d
Figure 2.
Distribution of mule deer January 1975-helicopter survey.
TABLE 2.
MULE DEER NUMBERS BY SEX, AGE CLASS AND LOCALITY DURING JANUARY 1975
HELICOPTER SURVEYS OF NICHOL'S COULEE RCA.
Males:
Fawns:
Fawns:
100
100
100
Females
Adults
Percent
Fawns
Deer Per
Sq. Mile
Total
Males
Females
Fawns
Females
Pastures III & IV
201
40
104
57
38.5
54.8
39.6
28.4
3.4
Pasture III
164
32
82
50
39.0
61.0
43.9
30.5
3.9
37
8
22
7
36.4
31.8
23.3
18.9
2.1
147
31
73
43
42.5
58.9
41.4
29.3
4.9
Nichol's Coulee
17
I
9
7
11.1
77.8
70.0
41.2
1.4
Seven Mile Coulee
37
8
22
7
36.4
31.8
23.3
18.9
2.1
Area
Pasture IV
CK Coulee
Post. I I I
Missouri
Figure 3.
Distribution of
River
mule deer on NlchoVs Coulee RCA during winter 1975
-18use occurred in Pasture IV, though these areas were much smaller, more
widely separated, and contained fewer mule deer than those in Pasture
III.
Ground observations during the spring showed much the same
distribution as winter (Fig. 4), with 83 percent of 235 mule deer
observed being in Pasture III.
The January, 1975, helicopter survey showed 54.8 fawns per 100
females in Pastures III and IV, an increase of 9.6 percent from 1974
to 1975 (Knowles 1975).
The number of fawns per 100 does was 1.9
times greater in Pasture III than in Pasture IV.
The fawn:doe ratio
for ground observations during January, 1975, over the whole study area
was 67.5:100.
The difference in fawn:doe ratios between the helicopter
survey and ground observations during January, 1975, may have been due
to more frequent observations in Pasture III where higher than average
numbers of fawns occurred.
Home Range and Movements
Knowles (1975) differentiated between normal and total home range,
in describing mule deer movements during summer and fall, with normal
including areas of intensive use, and total, all points of relocation.
I compiled home ranges for mule deer relocated during the winter and
spring in a manner comparable to the normal classification.
Three radio-collared and one ear-tagged mule deer were relocated
a total of 30 times during winter and spring (Table 3). •The 2.5
Post. IV
19-
TABLE 3.
Sex
SUMMARY OF HOME RANGES AND MOVEMENTS DURING WINTER AND SPRING FOR THREE RADIOCOLLARED AND ONE EAR-TAGGED MULE DEER.
Animals
Age
Times
Relocated
Dates of FirstLast Observations
Home Range
Area (Acres)
Distance Moved (Miles)
Max.
Min.
Mean
Male
2.5+
14
1/20 -- 6/24
1703
1.7
0.22
1.1
Male
1.5+
8
2/10 -- 6/24
1138
2.6
0.16
1.1
Female
1.5+
3
1/18 -- 1/31
506
1.3
0.34
0.9
Male
Fawn
5
1/19 -- 6/2
370
3.8
0.50
1.6
I
N)
0
1
-21year-old male had the largest home range, 1,703 acres, followed In
descending order by the yearling male, 1,138 acres, the yearling
female, 506 acres, and the male fawn, 370 acres (Fig. 5, Table 3).
Home ranges of the three radio-collared deer during winter and spring
were much smaller than their normal home ranges during summer and fall.
Differences ranged from 91 percent for the 2.5 year-old male to 66
percent for the yearling female.
Radid-triangulations of the radio-
collared deer, without visual contact, suggested that the home range ■
of the 2.5 year-old male may have been about 50 percent larger than
determined only by recorded visual relocations.
Addition of relo­
cations by triangulation did not affect the.determined home range
sizes of the two yearlings.
The yearling female was last seen on
January 31 and apparently died sometime.prior to recovery of the
collar in May.
During summer and fall, 1974, the yearling male was the only
marked mule deer which experienced cattle grazing within its home
range (Knowles 1975).
spring, 1975.
No cattle grazed that area during winter and
Large numbers of cattle grazed the home range of the
2.5 year-old male and some cattle grazed the male fawn's home range ■
from April I, 1975, until observations ceased on June 15.
Knowles (1975) noted an extension during late fall of the 2.5
year-old male's home range to the edge of timber in CK Creek drainage
and of the yearling buck's use of rangeland to the north of the timber
_______ 2 . 5 Y e e r - o l d
_____ 1 . 5
______ 1.5
Figure 5.
M o le
Y e o r - o ld M ole
Y e o r - old
----f
Fem ole
Home ranges of three radio-collared and one ear-tagged mule deer on Nichol1s
Coulee RCA during winter and spring 1975.
-23in Seven Mile Coulee.
These areas continued to be used extensively
during winter and early spring.
I did not observe the heavy use of burned areas within home ranges
of radio-collared deer described by KndWles (1975).
Also, the radio-
collared animals did not appear to be attracted by or associated with
water sources, although some deer were observed eating snow and gnawing
ice on frozen streams during the winter.
Heavy precipitation during
spring provided free water throughout the study area.
Although the marked mule deer appeared to have smaller home ranges
during winter and spring than those recorded by Knowles (1975),
distances moved between successive relocations were quite similar.
The maximum, minimum, and mean distances moved by the marked deer
between successive relocations from early January through late June
are listed in Table 3.
The longest recorded movement was by the male
fawn on June 2 when he was observed nearly four miles from his normal
home range.
The yearling male moved the least distance between
successive relocations.
adult males.
Mean movements were about equal for the two
Mean distances traveled by the yearling female and male
fawn were lower than those of adult males through the winter until the
fawn moved abruptly in early June.
The 2.5 year-old male also seemed
to move greater distances in early June, possibly because of intensive
human activity within its home range.
-
24
-
Group Size and Characteristics
Mean group size increased from January through March, then
decreased through June.
During winter the most commonly observed
group size was 2-5 (Table 4).
TABLE 4.
Single mule deer and groups of over 11
MONTHLY AND SEASONAL FREQUENCIES OF MULE DEER GROUP SIZES
An d m e a n g r o u p s i z e d u r i n g w i n t e r a n d s p r i n g .
Month
Season
January
February
March
Winter
April
May
June (1-15)
Spring■
I
9.1
9.7
4:5
7.6
8.3
41.5
68.8
37.0
2-5
70.5
51.6
45.5
56.3
50.0
51.2
31.2
46.9
Group Size
6-10
13.6
29.0
45.5
29.4
41.7
7.3
16.1
Mean
Group Size
11+
6.8
9.7
4.5
6.7
—
.
4.5
5.4 .
5.6
5.1 .
4.8
2.4
1.4
2.9
animals were observed with about equal frequencies during winter.
groups larger than ten mule deer were observed after March.
No
The
tendency toward larger group sizes during late winter appeared to be
related to use of open and level areas during that period.
Groups of
2-5 deer continued to be observed most commonly during spring, except
in June when the frequency of single animals increased sharply. . This
change has been related to intolerance of pregnant females to other
deer during fawning (Dasmann and Taber 1956, Einarsen 1956, Geist 1971,
and Knowles 1975).
-25Activity Habits and Patterns
Running was the most frequently recorded activity among all deer
observed during winter (Table 5).
Feeding was the dominant activity
over-all during spring and ranked second for observations during the
winter.
The dominant activity for the marked mule deer was feeding
during both seasons (Table 6).
Knowles (1975) reported an increase in
the occurrence of alert and running deer during fall, which he related
to increased wariness due to hunter disturbance.
The greater occur­
rence of observations of alert and running animals during winter may
reflect increased usage of open and level areas affording less security
and greater sight distances than the timbered breaks.
Activity periods were somewhat longer during winter and early
spring than reported by Knowles for summer and fall; some deer were
observable during most daylight hours.
Similar activity periods were
described by Mackie (1970).
Use of Habitat Types, Slopes, and Exposures
Mule deer used the Artemisia-Agropyron habitat type most fre­
quently during both winter and spring (Table 7).
The Pinus-Juniperus
habitat type was second most important during the winter and during
June.
It was the single most important type used by marked mule deer
during both winter and spring (Table 6).
The Sareobatus-Agropyron
habitat type was more frequently used over-all during spring, except
TABLE 5.
SEASONAL PERCENTAGES OF ALL MULE DEER ACTIVITIES OBSERVED WITHIN EACH HABITAT
TYPE AND TOTAL SEASONAL AVERAGES; AND MONTHLY PERCENTAGES OF ACTIVITIES FOR ALL
MULE DEER ON ALL HABITAT TYPES.
Habitat Type
Month
Traveling
Feeding
Activity Winter/Spring
Feeding and
Traveling
Bedded
Running
Alert
Artemisia-Agropyron
70/47
55/75
90/-
6/5
62/74
44/41
Pinus-Juniperus
21/27
7/4
10/-
71/62
31/13
37/29
Artemisia longifolia
-/7
23/4
-/-
12/5
5/2
17/2
Saroobatus-Agropyron
9/20
15/17
-/-
12/30
3/11
3/29
16/6
25/44
5/-
4/9
32/20
19/21
40
17
-
5
12
26
February
5
17
8
5
48
16
March
3
36
6
I
38
15
April
5
44
-
12
20
18
May
9
52
-
3
18
17
June
-
9
-
18
23
50
Average
January
-
TABLE 6.
27
-
PERCENTAGES OF OBSERVATIONS OF 4 MARKED MULE DEER BY
ACTIVITY CLASS, HABITAT TYPE, EXPOSURE, AND SLOPE
DURING WINTER AND SPRING.
Male
2.5
Marked Mule Deer
Male
Female
Male
Fawn
1.5
1.5
Mean
42.9
21.4
14.3
14.3
7.1
—
37.5
25.0
37.5
—
33.3
33.3
33.3
—
—
20.0
40.0
40.0
—
24.1
23.1
28.2
23.0
1.8
28.5
64.3
7.1
50.0
12.5
—
25.0
12.5
33.3
66.7
—
—
—
40.0
60.0
—
—
—
38.0
50.9
1.8
6.3
3.1
Activity
Alert
Bedded
Feeding
Running
Traveling
—
Habitat Type
Artemisia-Agropyron
Pinus-Juniperus
Artemisia longifolia
Saroobatus-Agropyron
Agropyron-Symphorioarpos
—
—
Exposure
N
NW
NE
S
SW
SE
W
E
Ridge Top
21.4
21.4
21.4
7.1
7.1
7.1
14.3
—
—
—
12.5
—
—
12.5
37.5
12.5
12.5
33.3
33.3
—
—
—
33.3
—
—
—
60.0
20.0
—
—
—
—
—
20.0
—
28.7
18.7
8.5
4.9
1.8
13.2
13.0
8.1
3.1
35.7
50.0
14.3
87.5
12.5
—
66.7
33.3
—
40.0
40.0
20.0
57.5
25.6
8.6
—
Slope
0-10°
10-25°
25-35°
TABLE 7.
MONTHLY AND SEASONAL PERCENTAGES FOR ALL USE OF HABITAT TYPE BY MULE DEER OBSERVED,
FOR EACH HABITAT TYPE AND FOR EACH COMMUNITY WITHIN A HABITAT TYPE.
TRACE (T) AMOUNTS <1%.
Habitat Type
Community
January
February
March
Winter
April
May
Artemisia-Agropyron
Artr-Agsp
Artr-Agsm
Artr-Agsm-Bogr
48
4
7
—
63
7
16
—
62
7
10
—
58
6
11
—
66
—
—
—
54
—
—
—
Pinus -Juniperus
Pipo-Agsp
Pipo-Jusc
Pipo-Arlo
30
4
14
8
16
7
5
4
26
12
8
5
24
8
9
5
19
9
—
4
17
7
4
5
Artemisia longifolia
11
13
9
11
—
Sareobatus-Agropyron
11
8
2
7
Artr
Agsp
Agsm
Arlo
Bogr
Jusc
Pipo
=
=
=
=
=
=
=
Artemisia tridentata
Agropyron spioatwn
Agropyron smithii
Artemisia longifolia
Bouteloua gracilis
Juniperus scopulorum
Pinus ponderosa
22
8
20
June (1-15)
50
Spring
5
60
—
—
T
46
9
5
32
18
8
2
7
—
—
—
3
5
20
-
29
-
in J u n e H e a v y usage of the Artemisia-Agropyron and SaroobatusAgropyron types in spring probably reflects the abundance of forbs of
the Poa and Tragopogon unions on these types (Knowles 1975).
Increased
use of the Pinus-Juniperus type during June appeared to reflect usage
of the open Pinus ponderosa-Artemisia longifolia community in which
extensive stands of yellow sweetclover occurred.
The Artemisia
longifolia habitat type was of some importance during winter.
Although
observations indicated that other habitat types received only minor use,
the Agropyron-Symphorioarpos type may have been used extensively during
late evening or at night as indicated by the frequent occurrence of
tracks.
No use was recorded on either the Xanthium strumarium of the
Pseudotsuga-Juniperus habitat types.
Deer, including the marked animals, were most commonly observed
on slopes of 0-10 degrees during winter and spring (Tables 6 and 8).
TABLE 8.
MONTHLY.AND SEASONAL PERCENTAGES OF ALL USE OF SLOPES BY
MULE DEER OBSERVED DURING WINTER AND SPRING.
Month
Season
January
February
March
Winter
April
May
June (1-15)
Spring.
0-10°
32
69
63
56.
64
57
55
60
Slope in Degrees
10-25°
57
22
26
35
35
32
32
33.
25-35°
11 •
9
11
10
2
11
14
7
-
30
-
Southerly exposures were most frequently used by all deer throughout
winter and spring; the marked deer preferred northerly exposures (Tables
6 and 9).
Knowles (1975) reported dominant usage of northerly exposures
by all mule deer during summer and fall..
Frequent use of open, level habitat types during winter and spring
y
corresponds to the high use of gentle slopes.
The spring of 1975 was
characterized by heavy snowfall during April and May, which may account
for extensive use of southerly exposures during that period.
Elk Populations and Range Use
Observations were made of 413 elk in 87 groups during winter,
1975, and 139 individuals in 54 groups during spring.
In addition,
60 elk were counted during helicopter surveys of Pastures. Ill and IV
in January, 1975.
Population Characteristics
Knowles (1975) described elk population trends on the study area
and adjacent areas during early January of 1974 and 1975.
A two-fold
increase in numbers of elk using these areas was concurrent with.a 97
percent increase of elk use in Pasture III from 1974 to 1975.
Age and
sex ratios from the January, 1975, helicopter survey of the study area
are listed by pasture and within Pasture III by drainages (Table 10).
These data show an over-all calf:cow ratio of 70:100.
Ground classi­
fications during the same month indicated a ratio of 56 calves:100 cows.
TABLE 9.
MONTHLY AND SEASONAL PERCENTAGES OF ALL USE OF EXPOSURES BY ALL MULE DEER OBSERVED
DURING WINTER AND SPRING.
Month
Season
_________________________________ Exposure
SW
SE
E
W
NW
NE
S
N
Coulee Bottom*
Ridge Top*
17
9
10
-
14
24
8
7
2
February
5
-
4
19
20
23
14
5
-
March
8
2
I
6
24
7
5
32
7
9
10
4
4
8
20
16
9
17
3
9
12
2
I
13
24
13
3
12
7
14
8
13
3
16
29
10
6
14
-
-
June (1-15)
14
23
18
9
14
9
9
5
-
-
Spring
11
9
3
14
25
12
5
12
3
7
January
Winter
April
May
Level areas where no exposure could be determined.
8
10
TABLE 10.
NUMBERS AND SEX AND AGE COMPOSITION OF ELK OBSERVED ON THE NICHOL'S COULEE
RCA DURING HELICOPTER SURVEYS JANUARY 1975 •
Area
Pasture III
Pasture IV
CK Coulee
Nichol1s Coulee
Total
Calves
Males:100
Females
Calves:100
Females
Calves:100
Adults
Total
Males
Females
58
12
27
19
44
70
49
2
2
0
0
0
0
0
50
6
26
18
23
69
56
8
6
I
I
—
—
—
60
14
27
19
52
70
46
-
33
-
Of 413 elk observed from the ground during winter, 98 percent
were in Pasture III.
CK Creek drainage.
Nearly 80 percent of these were observed in the
Three major winter concentration areas were
located in Pasture III (Fig. 6).
Elk were observed in one small area
of Pasture IV only in January, 1975.
Ground observations of elk
during spring showed a shift in use to the southern part of Pasture
III and some movement to Pasture IV (Fig. 7).
Of 139 elk observed
during spring, 86 percent were in Pasture ill; approximately 61 per­
cent of these were in the Nichol's Coulee drainage.
This shift in
distribution coincided with the onset of cattle grazing in Pasture
III in early April.
Initially the cattle were concentrated along
the northern boundary fence of Pasture III.
From here, they slowly
worked south along the CK Creek bottom and along ridges to the head of
Nichol's Coulee.
Observations during this period indicated that elk
avoided areas which they had used extensively, after cattle moved into
them.
Knowles (1975) reported 85 and 80 percent of all elk observed
during summer and fall 1974, respectively, were in Pasture III.
He
related this distributional pattern to quantity and quality of forage
available in the absence of cattle.
Knowles also reported that elk
avoided areas of cattle use during summer and fall.
Home Range and .Movements
Winter-spring home ranges and movements were determined from 90
relocations of three radio-collared elk between January I and June 24,
Past. I V
Pa s t . I l l
Missouri
Figure 6.
River
Distribution of elk on Nichol's Coulee RCA during winter 1975.
Past.
Figure 7.
IV
Past. III
Distribution of elk on Nichol1s Coulee RCA during spring 1975
—
1975 (Table 11).
square miles.
36
—
The adult male had the largest home range, 57.6
Home ranges of the two cows (24.9 and 22.4 square miles),
were less than half as large as that of the male (Fig. 8, Table 11).
Home ranges of these animals during the previous summer and fall were
42.6 square miles for the bull, and 38.0 and 20.7 square miles for the
younger cow and older female, respectively.
Seventy-one percent of relocations of the radioed bull, during
winter and spring, were in Pasture III, while comparable data for the
older and younger female elk were 87 and 78 percent, respectively
(Table 11).
Knowles reported 73, 89, and 85 percent of relocations
of the bull, older cow, and younger cow, respectively, were in Pasture
III during late summer and fall, 1974.
Cattle were present on portions
of the home ranges of all three radio-collared elk from April I to June
15, 1975, when the study was concluded.
All showed movements
characteristic of the general shift of elk to areas not used by cattle
as spring progressed.
The elk were extremely mobile within their comparatively large
home ranges.
Mean distances between successive relocations of radio-
collared elk range from 2.1 miles for the older female to 3.5 miles
for the male (Table 11).
The maximum distance between successive
relocations was recorded for the male (8.8 miles).
this animal traveled more than 4 miles overnight.
On one occasion
The minimum
distance between successive relocations was 0.1 miles, recorded for
TABLE 'll.
NUMBERS OF RELOCATIONS, HOME RANGE SIZES, AND PERCENTAGES OF- RELOCATING WITHIN A
GIVEN AREA DURING WINTER AND SPRING FOR THREE RADIO-COLLARED ELK ON THE NCRCA.
Distances
Home
Moved
Range
Between .
Dates of First Area
Successive
Relocations
Animals
Times
and Last .
(Sq.
Sex ■ Age -Relocated Observations Miles) Max. Min. Mean
1/9
6/17
57.6 . 8.8
0.6
3.5'
30
1/7
6/24
22.4
6.4
0.1
2.1
—
32
1/13
6/24
24.9
7.0
0.1
2.5
_
Male
2.5+.
28
'Female
8-10 .
4-6
Female
Percentages of Relocations
^ ____ . Within a Given Area
West of
East of
Pasture Pasture Pasture Pasture
IV.
III
' IV.
III
.
3.6
14.3
—
18.8
71.4
10.7
86.7
13.3
78.1
3.1
--O
O
Figure 8.
»-10 Yeo
4-6
Y e o r -old
Fe
Home ranges of three radio-collared elk on Nichol's Coulee RCA during
winter and spring 1975.
-
both cow elk.
39
-
All three radioed elk showed greater mean movements
between relocations in winter and spring, 1975, than during the fall
of 1974.
Numbers of relocations were not sufficient to quantify
differences in the extent of travel between winter and spring; however
trends suggested greater movement for the male, and lesser movement
for the two females during spring.
Group Size and Characteristics
Mean group sizes for elk were relatively constant during the
winter months, but decreased generally from winter to spring as well
as monthly from April through June.
Groups with which radio-collared
elk were associated showed similar decreases in mean numbers from
winter to spring (Table 12).
During winter observed groups most
commonly numbered 2 to 5 elk; single elk and groups of over 11 animals
being observed with about equal frequency, except in February, when
single animals were seen more than twice as frequently as large groups
Single animals were also observed more frequently than the larger
groups during spring.
No groups larger than 10 elk were observed
after March and none larger than 5 were seen during June.
Among the
radio-collared elk, the bull and older cow associated with groups of
2-5 most frequently during winter; the younger female associated with
groups of 6-10 most often.
Both females were observed most frequently
alone during spring, while the male continued to associate with groups
-
TABLE 12.
Female 8-10
Female 4-6
Male 2.5+
-
MONTHLY AND SEASONAL FREQUENCIES OF ELK GROUP SIZES AND
MEAN GROUP SIZE FOR ALL ELK OBSERVED DURING WINTER AND
SPRING; AND SEASONAL FREQUENCIES AND MEAN GROUP SIZE
OBSERVED F O R .THREE RADIO-COLLARED ELK DURING WINTER/SPRING.
Month
Season
January
February
March
Winter
April
May
June
Spring
40
I
2-5
10.3
24.1
15.8
16.1
30.8
45.5
75.0
.46.3
,
Group Size
6-10 .
53.9
58.6
47.4
54.0
38.5
45.5
25.0 ■
40.7
50/13
15/22
67/80
28/73
23/61
11/10
of 2-5 most commonly (Table 12).
during February.
23.1
.6.9
26.3
18.4
30.8
9.1
—— '
13.0
17/13
" 46/11
17/-
Mean
Group.Size
11+
12.8
10.3
10.5
11.6
—
—
5.0
4.4
4.8
4.8
3.5
2.5
1.5
2.6
—
-6/16/6
6/10
4.5/2.1
6.972.2
4.8/3.6
The largest group was 19, observed
The two radio-collared females were observed to­
gether or in the same groups four times during winter and spring; the
bull was observed in the same group as the younger cow once.
Knowles
(1975) reported constantly changing group characteristics among elk
on the study area.
Knight (1970) reported similar findings in the Sun
River area of Montana.
• I
Activity Habits and Patterns
During winter, feeding was the most frequently recorded activity
among all elk observed; alert and bedded animals ranked second (Table
13).
Alert animals dominated in observations during spring when feeding
TABLE 13.
SEASONAL FREQUENCIES OF ALL ELK ACTIVITIES OBSERVED WITHIN EACH HABITAT TYPE AND
TOTAL SEASONAL AVERAGES; AND MONTHLY PERCENTAGES OF ACTIVITIES FOR ALL ELK ON ALL
HABITAT TYPES.
Habitat Type
Month
__________________ Activity - Winter/Spring
Feeding and
Feeding
Traveling
Traveling
Bedded
Artemisia-Agroipijron
42/-
51/51
Pinus-Juniperus
13/54
12/3
Artemisia longifolia
15/8
11/3
Saraobatus-Agropyron
31/39
14/43
-/-
12/-
17/9
27/27
January
25
26
February
18
March
Running
Alert
32/58
7/7
42/31
39/41
-/42
77/67
47/46
41/43
-/-
-/23
-/-
16/20
11/-
21/10
-/7
-/-
-/6
20/11
9/9
20/35
—
19
6
24
17
18
18
8
21
—
44
6
24
15
11
April
—
22
—
11
9
59
May
16
31
15
5
9
25
June (1-15)
—
17
—
50
17
17
Agropyron-Symphoricarpos
Average
-/68/7/9
—
ranked second.
42
—
Radio-collared elk were most frequently alert when
first observed; bedded was second and feeding ranked third (Table 14).
The increase in alert elk during spring probably was related to
predominant usage of the open, generally level, Aptemisia-Agvopyron
habitat type during April.
As with mule deer, elk activity periods were longer during winter
and early spring than during summer and fall.
during most daylight hours.
Elk were observable
Between 10:00 AM and 2:00 PM most elk were
bedded when first observed.
Use of Habitat Types, Slopes, and Exposures
Over-all, elk usage was greatest on the Artemisia-Agropyvon
habitat type during winter and spring.
The same was also true for
all months except January and June,, when the Pinus-Juniperus type,
received 41 and 83 percent, respectively, of the total observed elk
use (Table 15).
The Sarcobatus-Agropyron habitat type also received
important use throughout the period.
During winter, elk observed on
this type were most commonly traveling, while feeding was recorded
most frequently during spring.
Elk were also observed frequently on
the Agropyron-Symphoricarpos type in major coulee bottoms during
winter, especially in February.
minor usage during both seasons.
Other habitat types received only
Increased usage on the Artemisia-
Agropyron and Sareobatus-Agropyron habitat types during spring
appeared related to the vernal succulence and availability of forbs
/
-
TABLE 14.
43
-
PERCENTAGES OF OBSERVATIONS OF 3 RADIO-COLLARED ELK BY
ACTIVITY CLASS, HABITAT TYPE, EXPOSURE AND SLOPE DURING
WINTER AND SPRING.
Female 8-10
Radio-Collared Elk
Female 4-6 Male 2.5+
Mean
Activity
Alert
Bedded
Feeding
Running
Traveling
Feeding & Traveling
34.4
31.3
18.8
9.4
3.1
3.1
30.3
33.3
15.2
9.1
9.1
3.0
50.0
28.6
14.3
7.1
—
38.2
31.1
16.1
6.2
6.4
2.0
Habitat Type
Artemisia-Agvopyron
Pinus-Juniperus
Artemisia longifolia
Sarcobatus-Agropyron
Agropyron-Symphoriearpos
9.4
56.3
—
31.3
3.1
14.7
58.8
—
20.6
5.9
10.7
67.9
3.6
10.7
7.1
11.6
61.0
1.2
20.9
5.4
Exposure
N
NW
NE
S
SW
SE
W
E
Ridge Top
Coulee Bottom
16.1
12.9
9.7
16.1
16.1
9.7
6.5
6.5
—
6.5
11.8
8.8
11.8
20.6
17.7
8.8
—
11.8
2.9
5.9
21.4
14.3
3.6
3.6
17.9
3.6
10.7
7.1
3.6
14.3
16.4
12.0
8.4
13.4
17.2
7.4
5.7
8.5
2.2
8.9
Slope
0-10°
10-25°
25-35°
58.1
38.7
3.2
52.9
38.2
8.8
64.3
35.7
58.4
37.5
4.0
—
—
TABLE 15.
MONTHLY AND SEASONAL PERCENTAGES FOR ALL USE OF HABITAT TYPES BY ELK OBSERVED
DURING WINTER AND SPRING FOR EACH HABITAT TYPE AND FOR EACH COMMUNITY WITHIN
A HABITAT TYPE.
Habitat Type
Community
January
February
March
Winter
April
May
June (1-15)
Spring
Artemisia-Agropyron
Artr-Agsp
Artr-Agsm
Artr-Agsm-Bogr
26
37
16
4
43
32
—
—
50
27
—
—
36
32
5
I
57
—
—
—
31
—
—
—
——
—
—
—
37
—
—
—
Pinus-Juniperus
Pipo-Agsp
Pipo-Jusc
Pipo-Arlo
41
16
53
20
25
—
9
66
26
42
13
46
33
17
35
35
37
—
59
—
28
—
65
22
83
—
50
50
36
—
60
20
Artemisia longifolia
11
I
Saroobatus-Agropyron
19
17
17
17
3
15
8
8
Agropyron-Symphoricarpos
Artr
Agsp
Agsm
Bogr
Pipo
Jusc
Arlo
=
=
=
=
=
=
=
Artemisia tridentata
Agropyron spicatim
Agropyron smithii
Bouteloua gracilis
Pinus ponderosa
Juniperus scopulorum
Artemisia longifolia
—
6
—
6
7
—
30
5
—
17
—
4
21
3
-
on these types.
45
-
Repeated relocations of radio-collared cows during
the calving period accounted for most of the 83 percent use of the
Pinus-Juni-perus habitat type during June.
Elk were observed feeding
on the loose shale slopes of the Pinus ponderosa-Avtemisia longifolia
community throughout winter,
Extensive stands of yellow sweetclover
occurred on this community during spring, and were used quite heavily
by elk.
Radio-collared elk used the Pinus-Juniperus habitat type most
frequently during winter and spring, while the Sareobatus-Agropyron
type ranked second (Table 14).
Slopes of 0-10 degrees were most commonly used during winter and
spring by all elk, including the three radio-collared animals (Tables
14 and 16).
Use of steeper slopes (10 to 35 degrees) steadily
increased from April through June, and was related to greater usage
of the Pinus-Juniperus and Pinus-Artemisia communities during spring.
Southerly exposures were used most frequently by all elk observed,
including radio-collared animals, during winter and spring (Tables
14 and 17).
Elk were observed on coulee bottoms most often during
February, reflecting greater usage of the Agropyron-Symphoriearpos
habitat type during that month.
Knowles (1975) reported increased
usage of steeper slopes from summer to fall, and predominant use of
northerly slopes.
He also related this to greater use of Pinus-
Juniperus communities during late summer and fall.
-46TABLE 16.
FREQUENCIES OF ALL USE OF SLOPES BY ELK OBSERVED DURING
WINTER AND SPRING BY MONTH AND SEASON.
Month
Season
0-10
January
February
March
Winter
April
May
June
Spring
65
85
54
69
74
52
25
56
Slope in Degrees
10-25
25-35
6
8
5
5
14
33
13
26
7
' 46
25
21
35
42
31
35+
3
- •
.I
—
Cattle Populations and Range Use
Results are based on 116 groups of cattle totaling 1,221
individuals observed during spring, 1975.
Numbers and Distribution
Cattle grazed the study area beginning April I, when .approxi­
mately 800 Hereford cows without calves were placed in Pasture III.
Calving occurred in the pasture during April and May..
About one-half
of these cattle were moved from Pasture III to Pasture II in early
June.
Knowles (1975) summarized grazing usage from 1972 to 1974 for
each pasture and totals for the study area (Appendix Table 29).
During spring, cattle were observed primarily in major coulee
bottoms and on level upland ridge tops.
Greatest concentrations
(Fig. 9) occurred along the bottoms of CK Coulee and major side
(
TABLE 17.
Month
Season
FREQUENCIES OF ALL USE OF EXPOSURE BY ELK DURING WINTER AND SPRING BY MONTH
AND SEASON.
________________________________ Exposure
NW
NE
S
SW
SE
W
E
N
January
26
I
10
February
10
3
5
March
—
Winter
11
—
15
4
6
6
21
21
6
4
April
May
—
June (1-15)
17
25
Spring
3
12
—
8
10
Coulee Bottom*
Ridge Top*
3
17
I
7
17
9
—
29
2
2
17
28
3
17
19
26
11
9
8
8
15
15
4
11
19
5
—
30
6
6
—
—
9
11
17
22
6
20
17
8
33
9
20
19
Level areas where no exposure could be determined.
—
—
6
13
—
5
—
9
—
3
8
Missouri
Figure 9.
River
Distribution of cattle on Nichol's Coulee RCA during spring 1975.
-
49
-
coulees, as well as on uplands and ridge tops along the north and east
boundaries of Pasture III and at the head of Nichol's Coulee.
grazed along the Missouri River only in CK Coulee.
Cattle
This distributional
pattern was reached by the end of April and Was essentially maintained
through mid-June when observations ended.
As reported by Knowles
(1975), all areas of heavy cattle use were within a mile of a water
source, though not all areas within this distance were heavily used.
Group Characteristics
Group sizes for cattle were lowest during. April and increased
through May to June.
Groups numbering from one to ten cattle were
most frequent during all months (Table 18).
TABLE 18.
The largest group
FREQUENCIES OF CATTLE GROUP SIZE CLASSES; AND MEAN GROUP
. SIZES DURING SPRING BY MONTH AND SEASON.
Month
Season
1-10
April
May
June (1-15)
Spring
82.4
68.8
71.7
. 72.1 .
Group Size
11-20
21-30
31+
11.8
20.8
■8.7
14.4
5.9
6.3
6.7
7.2
— —
4.2
10.9
6;3
Mean
Group Size
6.1
11.0
12.2
10.5
observed was 82, at a well on the CK' Coulee bottom in late May, 'The
trend toward larger group sizes during May and June reflected increased
aggregation around water sources during mid-day as temperatures in­
creased.
The mean group size for spring was nearly twice as large as
-
50
-
those reported by Knowles (1975) for summer and fall.
Activity Habits and Patterns
Most cattle observed during spring were feeding (Table 19).
TABLE 19.
PERCENTAGES OF ALL ACTIVITIES RECORDED FOR CATTLE OBSERVED
DURING SPRING BY MONTH AND SEASON.
Activity
April
Loafing
Feeding
Bedded
Alert
Traveling
' 1.5
63.4
26.1
2.2
6.7
May
June (1-15)
Spring
30.7
20.4
28.4
5.9
13.8
29.4
30.8
26.2
3.6
10.0
34.7
33.4
23.5
1.5
6:8
However, feeding decreased in frequency among observations from April
to June 15, as greater percentages of cattle were observed bedded and
loafing around water sources.
Although cattle were observed,feeding
during all daylight hours, most intensive feeding occurred during
early morning and late evening.
Use of Habitat Type, Slope, and Exposure
The Artemisia-Agropyron habitat type was most frequently used by
cattle.
The Sareobatus-Agropyron type ranked second (Table 20).
Only
minor use was made of the Pinus-Juniperus and Agropyron-Symphoricarpos
habitat types.
-51TABLE 20.
FREQUENCIES OF ALL CATTLE ACTIVITIES OBSERVED DURING SPRING
FOR EACH HABITAT TYPE AND TOTAL CATTLE USE OF EACH HABITAT
TYPE. TRACE (T) AMOUNTS <1%.
■
Habitat Type.
. ■
Loafing
Avtem-LsrLa-Agvoyyron
. 58
P-Lnus-Junipevus
7
2
Artemisia ZongifoZia
Saveobatus-Agvopyvon
25
AgvopyvonSymphovicavpos9
.
Activity
Bedded 'Alert
Feeding
34 .
T .
—
43
11'
100
35
40 ■
—
31
6
Traveling
Total
37
.63 ■
46
5
T
35
•--
14
.
Slopes of 0-10 i
degrees were most frequently used by cattle overall during spring and for all months.
Use of slopes over ten degrees
increased during May but decreased in June (Table 21) .
TABLE 21.
Level areas
PERCENTAGES OF USE OF SLOPES BY ALL CATTLE OBSERVED DURING
SPRING BY MONTH AND SEASON. TRACE (T) AMOUNTS <1%.
Month
Season
0-10
April ;
May.
June (1-15)
Spring
92.5
84.4
95.2
90.3
,
Slope in Degrees
10-25
■7.5
14.8
4.8
. 9.4
25-35
—
T '
—
. T
along coulee bottoms and ridge tops were most often used over-all,
while southerly and easterly exposures were also used frequently by
cattle (Table 22).
TABLE 22.
Month
Season
April
May
June (1-15)
Spring
FREQUENCIES OF USE OF EXPOSURE BY CATTLE OBSERVED DURING SPRING BY MONTH AND
SEASON. TRACE (T) AMOUNTS <1%.
Ridge Top*
Coulee Bottom*
Exposure
N
NW
NE
S
SW
SE
W
E
—
—
55.2
T
2.2
—
5.2
12.7
6.7
T
42.5
6.5
5.9
1.3 13.1
4.0
12.3
5.7
29.6
4.6
T
8.2
5.0
9.6
1.4
3.0 32.3
4.8
38.0
5.0
3.0
. 4.3
8.5
7.5
6.7
7.0 15.1
17.2
Level areas where no exposure could be determined.
13.1
T
-
53
-
Food Habits
Mule Deer
Results are based on examination.of six mule deer feeding sites
totaling 2,071 instances of use during winter, and three feeding sites
totaling 752 instance^ of use during spring.
During winter, browse was the most important forage class, while
forbs ranked second (Table 23).
negligible.
Utilization of grasses and sedges was
Rubber rabbitbrush (Chvysothamnus nauseosus) and yellow
sweetclover was the most important shrub and forb, respectively.
Deer
were frequently observed to dig for roots of yellow sweetclover on
the loose shale slopes of the Avtemistd longifotia habitat type and
the Pinus pondevosa-Avtemisia longifotia community.
Knowles (1975)
noted deer digging for yellow sweetclover roots on the Avtemisia
longifotia type during fall and found roots of sweetclover to comprise
as much as 84 percent of the contents of rumen samples collected in
October.
Mule deer actively sought the roots of yellow sweetclover
from mid-fall to early spring; however, dried stems of first-year
sweetclover and some second-year stalks were also eaten frequently.
Utilization of big sagebrush was restricted to the mid-winter period.
Dusek (1971) reported rubber rabbitbrush to be the most important item
in the winter diet of mule deer on a prairie habitat in Northcentral
Montana; Mackie (1970) found big sagebrush and rubber rabbitbrush to
-
TABLE 23.
54
-
AGGREGATE MEAN PERCENTAGES AND FREQUENCY OF OCCURRENCE FOR
EACH PLANT SPECIES USED BY MULE DEER, WINTER AND SPRING.
TRACE (T) AMOUNTS ARE <1%. WINTER - 6 FEEDING SITES.
SPRING - 3 FEEDING SITES.
Forage Class
Plant Species Used
Aggregate Mean
Percentage W/S
Frequency of
Occurrence W/S
Browse
Artemisia longifolia
Artemisia tridentata
Chrysothamnus nauseosus
Juniperus scopulorum
Pinus ponderosa
Rhus trilobata
Rosa spp.
Saroobatus vemrlculatus
Symphoricarpos spp.
6.7/19.3/22.9/8.4/1.4/10.1
-/23.3
T/1.6/T/-
33/33/50/33/17/33
-/33
17/17/17/-
61.5/33.3
83/33
-/7.4
-/32.1
3.7/33.3/-/6.4
1.0/T/-
-/33
-/33
17/67/-/67
33/17/-
38.5/45.8
67/67
Agropyron smithii
Bouteloua gracilis
Carex geyeri
Stipa viridula
-/T
-/11.5
T/-/8.5
-/33
-/33
17/-/33
Total Grasses
T/20.8
17/33
Total Browse
Forbs
Allium textile
Comandra umbellatum
Eriogonum multioeps
Melilotus officinalis
Musineon divarication
Tragopogon dubius
Yucca glauca
Total Forbs
Grasses and Sedges
-
55
-
be the most and second most important forage species, respectively,
during four winters.
During spring forbs and grasses increased in importance and
utilization of browse declined (Table 23).
Pale bastard toadflax,
blue gramma, and skunkbush sumac (Rhus tvitobata) was the most
important forb* grass, and shrub, respectively.
Mackie (1970)
reported an increase in use of forbs and grasses by mule deer during
spring, although browse remained the most important forage class.
He
found Sandberg's bluegrass (Roa seounda) to be the most important
grass utilized by mule deer during spring.
Elk
Results are based on examination on nine feeding sites totaling
4,020 instances of.use during winter.
No quantitative data on the
food habits of elk were obtained during spring.
Forbs were the most important forage class during winter (Table
24).
Yellow sweetclover was the most important forb; soapweed ranked
second.
Grasses comprised 38 percent of recorded instances of use,
with western wheatgrass the most important grass.
Green rabbitbrush
was the major browse species used by elk.
Mackie (1970) reported predominant use of grasses and minor use
of forbs by elk during four winter periods.
He also found western
wheatgrass to be the single most important grass.
-
TABLE 24.
56
-
AGGREGATE MEAN PERCENTAGES AND FREQUENCY OF OCCURRENCE
FOR EACH PLANT SPECIES USED BY ELK DURING WINTER AT NINE
FEEDING SITES EXAMINED. TRACE (T) AMOUNTS <1%.
Forage Class
Plant Species Used
Aggregate Mean
Percentage
Frequency of
Occurrence
Browse
Artemisia longifolia
Chrysothamnus visoidiflorus
Rhus trilobata
Total Browse
1.8
9.7
T
33
22
11
11.5
56
2.0
T
T
5.8
29.2
T
12.9
22
11
11
11
77
11
33
50.5
100
20.1
3.6
4.5
1.4
5.3
3.1
66
11
11
11
11
11
38.0
89
Forbs
Artemisia ludoviciana
Aster spp.
Cirsium spp.
Eriogonum multioeps
Melilotus officinalis
Tragopogon dubius
Yucca glauca
Total Forbs
Grasses
Agropyron smithii
Ca lamoviIfa longifo Iia
Koeleria cristata
Muhlenbergia cuspidata
Oryzopsis hymenoides
Stipa comata
Total Grasses
-
57
-
Like mule deer, elk were observed to dig for yellow sweetclovef
roots on the loose shale slopes of the Artemisia longifolia habitat
type and the Pinus ponderosa-Artemisia longifolia community.
In late March, elk were observed feeding on new green growth of
grasses, especially needle-and-thread (Siipa oomata) and western
wheatgrass.
As spring progressed, elk appeared to feed increasingly
on early forbs such as pale bastard toadflax, wild parsley (Musineon
divarioatum) ,■and American vetch (Vioia am'erioana).
As yellow sweet-
clover flowered during June, elk were observed to feed almost
exclusively on that species.
Mackie (1970) reported grasses to be the most important forage
class for elk during spring.
Western wheatgrass individually com­
prised about 50 percent of the total grass use.
He also found that
as forbs become more abundant, elk made increasing use of this forage
class.
Cattle
Results are based on examination of six cattle feeding sites,
totaling 3,771 instances of use, during spring and examination of
contents of one rumen from a cow found dead on the area in June.
Grasses and sedges constituted the most important forage class,
contributing 50 percent of total instances of use at feeding sites
(Table 25). ■ Western wheatgrass, green needlegrass, and blue grama
-58TABLE 25.
AGGREGATE MEAN PERCENTAGES AND FREQUENCY OF OCCURRENCE FOR
EACH PLANT SPECIES USED BY CATTLE DURING SPRING AT SIX
FEEDING SITES EXAMINED.
Forage Class
Plant Species Used
Aggregate Mean
Percentage
Frequency of
Occurrence
Browse
Artemisia aana
Chrysothamnus viscidif torus
Eurotia lanata
Total Browse
.
1.4
. 1.4
1.7
17
17
17
4.5
33
Forbs
Altium textile
Lomatium spp.
Hymenoxys riahardsonii
Melilotus officinalis
Musineon divarication
Taraxacum officinalis
Vicia americana
Total Forbs
6.8
1.9
1.7
25.1
4.0 .
1.0
4.9
45.2
33
17
17
67
50
17
33
100
Grasses and Sedges
Agropyron smithii
Agropyron spicatum
Bouteloua gracilis
Carex fiIifoIia
■Koeleria cristata
Stipa viridula
Total Grasses and Sedges
14.0
5.4
10.4
5.4
2.8
12.4
83
17
50
17
17
50.
50.3
10.0
-
59
-
accounted.for 37 percent of total Instances of use.
Forbs, largely
yellow sweetclover, constituted 45 percent of total instances of use.
Utilization of browse was minor, comprising only five percent of total
use at feeding sites.
Mackie (1970) reported grasses constituted 82
percent of total cattle use during four spring periods, with western
wheatgrass being the most important grass species.
Yellow sweetclover comprised approximately 97 percent of the
contents from a cattle rumen obtained during June, 1975 (Table 26).
By the second week of June, second-year stems of yellow sweetclover
were flowering profusely, and cattle appeared to be feeding almost
exclusively on that species.
The extensive use of yellow sweetclover
may account for lesser use of grasses than reported by Mackie (1970).
-
TABLE 26.
60
-
PERCENTAGE OF RUMEN CONTENTS OBTAINED FROM ONE CATTLE
RUMEN DURING JUNE 1975.
Forage Class
Percentage
Forbs
Metitotus offioinatis
Unknown forb
Total Forbs
96.8
0.6
97.4
1.0
Grass
Browse
Artemisia spp.
Rhus tritobata
Total Browse
0.3
1.3
1.6
DISCUSSION
Interspecific Range Relationships
Interspecific conflict between large wild and domestic ungulates
may involve either forage or land use competition (Smith and Julander
1953).
Forage competition occurs if the supply of a forage species
used in common is inadequate for the requirements of one or both
species of animals.
Land use competition exists when a reduction in
numbers of one animal species would result in increased numbers of
another on a given area.
Cole (1958) deemed four conditions necessary
for competition in ungulates to exist:
(I) both species use the same
range areas, (2) both use the same forage plants, (3) the forage plants
are important for either or both species,
(4) the forage plants are in
limited supply, or deteriorating in production as a result of combined
use.
Both animals need not use the same area or forage plants at the
same time, since use during one season may affect the amount or quality
of forage available during another season.
Competition may involve
elements of "interference" as well as "exploitation" (Miller 1967).
Interference includes any activity which either directly or indirectly
limits a competitor's access to a necessary resource; while exploi­
tation relates to the utilization of a resource once access has been
achieved.
An element of interference is implicit in the "disturbance
competition" concept of Deriniston (1956).
Disturbance competition
—62—
usually involves movement of one animal from and/or avoidance of areas
used by another, due perhaps to "social intolerance" (Lonner 1975).
This type of interaction has been observed between moose (Aloes aloes)
and cattle (Denniston 1956, Stone 1971, Schladweiler 1974), elk and
cattle (Jeffery 1963, Stevens 1966, Skovlin 1968, Mackie 1970, Knowles
1975, Lonner 1975, Pictpn 1976), between elk and domestic sheep
(Kirsch 1962, Coop 1971), and between deer and cattle (McMahan 1966,
Firebaugh 1969, Dusek 1971).
The extent to which social intolerance (or disturbance competition)
occurs or is observed may be determined by the availability of suitable
undisturbed habitat for the intolerant species on a particular range.
Thus, Schladweiler (1974) stated "... The reaction of moose to the
presence of cattle may be related to the availability of suitable
adjacent moose habitat.
areas.
Where it is ample, moose may avoid cattle-use
However, if cattle occupy most, or all, of the suitable moose
habitat, the moose would be forced to use the area along with the
cattle."
Knowles (1975) discussed interspecific relationships on the
Nichol's Coulee study area on the basis of possible forage and land
use competition during summer and fall.
His observations of dis­
turbance movements of mule deer and, especially, elk were related to
land use rather than distinct disturbance competition.
My evaluation
of range relationships and possible competition between mule deer, elk.
-
63
-
and cattle during winter and spring follows similar criteria; however,
additional consideration is given to social intolerance and possible
disturbance competition as distinct from land use competition in the
sense of Smith and Julahder (1953).
Mule Deer and Cattle
The study area was not grazed by cattle during winter, and forage
plants or plant parts utilized by deer during the winter were generally
different than those used by cattle during the previous summer and fall.
During spring, the Avtemis-Ia-AgropyTon and Sareobatus-Agropyron
habitat types were used extensively by both deer and cattle.
Deer were
observed on the Pinus-Juniperus habitat type 3.6 times as often as were
cattle.
Slopes of less than ten degrees were used almost exclusively
by cattle and 60 percent of the time by deer.
Southerly exposures
were most important for deer, while cattle were most frequently ob­
served on level coulee bottoms.
Forbs comprised approximately 45 per­
cent of the diet of both species, but different forb species appeared
to be utilized.
Grass, the major forage class used by cattle, ranked
third in importance for mule deer.
Forbs and browse ranked first and
second, respectively, in the diet of deer.
These data indicated little
opportunity for forage competition between cattle and mule deer during
the period.
However, very, few data were obtained during early spring,
when greater overlap in food habits might have occurred as both mule
-
64
-
deer and cattle utilized new growth of early grasses and forbs.
Because of food habit differences, land use competition was also
believed to be of little consequence on the study area, during spring.
The general distribution of mule deer during spring, when cattle
were present, differed very little from winter.
On one occasion eight
mule deer were observed feeding approximately 100 yards from cattle.
In three other instances deer were observed less than 1/8 mile from
cattle.
Avoidance of cattle use areas by.deer has been reported by
McMahan (1966), Firebaugh (1969), and Dusek (1971).
Skovlin (1968)
reported that cattle grazing had less influence on the distribution
of deer than on that of elk.
Knowles (1975) reported that the radio-
collared yearling male, whose home range was grazed by cattle,
exhibited abnormally long movements comparable to those of a two-yearold male mule deer.
The only unusual movement of a radioed deer during
spring occurred in June when the 2.5-year-old. male reacted to the
presence of cattle by crossing a fence to an ungrazed area (Knowles
pers. comm.).
Yellow sweetclover, the preferred forage at the time,
was very abundant on both sides of the fence and forage availability
could not be considered a probable cause of the movement.
Although these data suggest that some social intolerance of mule
deer for cattle may occur, there was no evidence that disturbance
competition was widespread and/or important, compared with forage and
—
65
-
land use competition.
Elk and Cattle
Both elk and cattle used the. -Artem-IsrIa-Agvopyron and SarcobatusAgropyron habitat types extensively; elk also used, the 'Pinus-Juniperus
type heavily.
■
Only five percent of all cattle observed were on the
■■
i
Pinus-Juniperus type.
less, than cattle.
Elk used steep slopes, more, and. coulee bottoms
Food habits of elk and cattle were not compared
quantitatively during spring, but forage competition between the
two species was precluded by the movement of elk from areas of cattle
concentration.
Soon after cattle entered
Pasture III, elk moved to portions of
the pasture where few, if any, cattle occurred.
As spring progressed,
increasing numbers of elk also occurred in the ungrazed Pasture IV.
Knowles (1975) suggested the occurrence of land use competition
between elk and cattle on the basis of distributional differences
during summer and fall.
He attributed a preference of elk for Pasture
III over Pasture IV, when both pastures were ungrazed, to greater
abundance of yellow sweetclover in Pasture III.
My observations
indicated that changes in the distribution of elk were more closely
related to the presence of cattle and a "social intolerance" of elk
for cattle than to the kind and amount of forage available on the area
Because of this, the interaction involved might be more appropriately
—66—
considered as disturbance competition rather than land use competition,
though the consequences might be similar.
Social intolerance of elk
to livestock has been reported or implied in several studies where elk
were observed to move from areas, of cattle concentration before pre­
ferred forage supplies became depleted (Jeffery 1963, Stevens 1966,
Sk'ovlin 1968, Mackie 1970, Lonner 1975, and Picton 1976).
In the
Little Belt Mountains of central Montana, elk used many areas which
were also grazed by cattle, but avoided areas used by sheep (Kirsch
1962, Coop 1971).
During my study, elk were observed within 1/8 mile
of cattle on only one occasion. .
Because adequate adjacent habitat was available to elk during
spring, they readily moved to areas not grazed by cattle, and the
opportunities for competition, especially for forage, did not occur.
Changes in the distribution of elk from winter to spring as cattle
became distributed throughout Pasture III appeared to indicate a
social intolerance of elk towards cattle and possible disturbance
competition.
Although the effects of this disturbance on the elk
population are not known, they did not appear important at the present
time.
Mule Deer and Elk
Range use and food habits of mule deer and elk on the study area
overlapped extensively during summer (Knowles 1975).
During fall,
-67range use continued to be similar, but food habits diverged.
Data
for winter indicated generally similar range use habits, although
mule deer used steeper slopes somewhat more and coulee bottoms less
than did the elk.
Loose shale slopes of southerly aspect received
considerable use by both species.
Browse comprised about 62 percent
of the mule deer diet during winter, and forbs made up 51 percent of
the elk diet.
species.
elk.
Yellow sweetclover was extensively utilized by both
Range use changed little during spring for either deer or
Use of grasses by deer increased considerably during early
spring, but forbs remained important to both ungulate species.
Knowles (1975) indicated that forage competition between mule
deer and elk could occur, especially if elk numbers were allowed to
increase.
fall.
Land use competition may have occurred during summer and
Data obtained during this study indicated that both forage and
land use competition could have occurred during winter and early spring
as both deer and elk heavily utilized sweetclover on southerly shale
slopes of the Artemisia longifolia and Pinus-Juniipevus habitat types.
Mule deer and elk appeared tolerant of each other on the area.
On six occasions mule deer were observed less than 1/8 mile from elk.
Rest-Rotation Grazing Versus Season-Long Grazing
Knowles (1975) reported few major differences in range use and
food habits of mule deer, elk and cattle on the Nichol's Coulee area
during summer and fall as compared with earlier studies on an adjacent
-68area of season-long cattle grazing (Mackie 1970).
were also indicated for winter and spring.
Few differences
As during summer and fall
(Knowles 1975), home ranges of mule deer during winter and spring were
considerably larger than those described by Mackie (1970).
Knowles
(1975) suggested that the difference may have reflected greater
habitat diversity and population density on Mackie's study area.
Habitat usage by mule deer was similar to that reported by
Mackie (1970), except that greater use of the'Artemisia longifolia
habitat type was recorded in winter and greater use of the
Sarcobatus-Agropyron type occurred during spring.
also generally similar.
Food habits were
Browse ranked first in the winter diet of
mule deer in both studies.
My findings indicated that forbs were of
greater importance during winter and spring, but this probably was
related to the exceptional abundance of yellow sweetclover on the
Nichol's Coulee area in 1974 and 1975.
Mackie found considerable
differences in usage of forbs and browse between seasons and years
in relation to the abundance of forbs on his study area.
The average
size of mule deer groups was similar to that reported by Mackie (1970)
in winter but only about half as large during spring.
Habitat usage by elk was generally similar to that found by
Mackie (1970) during both seasons, with only the Saroobatus-Agropyron
habitat type receiving more use during this study.
Mackie (1970)
reported grasses to be the most important item in the diet of elk
-
69
-
dur'ing winter and spring, with only minor use of forbs.
Forbs
accounted for 50 percent of the elk winter diet on the Nicholrs Coulee
area, with yellow sweetclover and soapweed being the most important.
Elk group sizes averaged about half as large as those reported by
Mackie (1970) for both seasons.
Cattle used coulee bottom habitat types more extensively during
spring on the Nicholrs Coulee area than on Mackiers (1970) study area.
Knowles (1975) found greater use of the Saroobatus-AgvopyTon habitat
type during summer and fall and attributed it to the well developed
system of reservoirs and wells in the major coulees of NicholrS Coulee
RCA.
Grasses and forbs ranked first and second, respectively, in the
diet of cattle during spring.
Heavy fo r b 'usage reflected the almost
exclusive feeding on yellow sweetclover in early June.
Mackie (1970)
found 82 percent use of grasses by cattle in spring on the continuously
grazed area where sweetclover was abundant during only one of three
years.
Knowles (1975) concluded that rest-rotation grazing had not
affected any major changes in range use and food habits of mule deer,
elk, and cattle during summer and fall.
Similarly, the differences
in range use and food habits between those reported by Mackie (1970)
for a season-long grazing system and those found during this study
cannot be ascribed to any effects of a rest-rotation, grazing system.
MANAGEMENT RECOMMENDATIONS
Results obtained during winter and spring of 1975 support the
conclusion of Knowles (1975) that:
"Proper stocking rates of cattle
and a conscientious effort to control elk numbers in relation to the
forage needs of mule deer can result in an efficient use of a forage
resource in this rest-rotation grazing system."
APPENDIX
Ta b l e 27.
LAND s t a t u s a n d g r a z i n g c a p a c i t y o f NICHOL'S COULEE RCA.
AUMs
100.0
Total
Public Domain
Private
Lease
Pasture I
AUMs
Total for Area
Ownership of
Control
12,271
74,947
9,795
8,249
1,534
1 0 0 .0
17,352
1,720
1,280
Pasture II
AUMs
3,015
20,516
2,466
13,726
3,431
Pasture III
AUMs
28,913
2,984
Pasture IV
AUMs
21,109
2,607
19,809
2,453
4,996
1,280
1,260
Holding Pasture
Acres
Month
TREATMENT
J
F
M
A
M
J
J
S
A
0
N
*vX
GRAZE
II I
§§i
W
A
B
REST FOR Vl GOR
C
REST FOR SEEDRII E (ST\ D
D
gri.ZE
H I
REST FOR ISTABLI SHMENT OF RE PRODUC TION
Figure 10. Grazing formula for Nichols Coulee RCA
SR:GRi
S
ll
..
Il
I
D
-
74
-
Years combination occurred
1965
1969
1973
PASTURE I
PASTURE II
Years combination occurred
1966
1970
1974
PASTURE I
PASTURE II
TREATMENT A
TREATMENT D
TREATMENT B
TREATMENT A
PASTURE IV
PASTURE III
PASTURE IV
PASTURE III
TREATMENT B
TREATMENT C
TREATMENT C
TREATMENT D
Years combination occurred
1967
1971
1975
PASTURE I
PASTURE II
Years combination occurred
1968
1972
PASTURE I
PASTURE II
TREATMENT C
TREATMENT B
TREATMENT D
TREATMENT C
PASTURE IV
PASTURE III
PASTURE IV
PASTURE III
TREATMENT D
TREATMENT A
TREATMENT A
TREATMENT B
res in Nichol's Coulee RCA
within and between years.
75
MEAN PERCENTAGE CANOPY COVERAGE (C) AND FREQUENCY OF OCCURRENCE (O) OF PLANT TAXA WHICH OBTAINED A MEAN COVERAGE OF I PERCENT OR MOKE IN ONE
OR MORE OF THE ELEVEN PLAN! COMMUNITIES DURING SUMMER AND FALL. VALUES FOR BARE GROUND AND LITTER ARE ALSO INCLUDED. TRACE (T) AMOUNTS ARE
FOR VALUES BETWEEN I PERCENT AND 0.1 PERCENT. NUMBERS OF SITES EXAMINED ARE IN PARENTHESIS.
----------------------- — — — — :— :
c .: • : : : : : Lrr ::;T r r r r r r : ™ : :.r.r* :r::::::rrrr---------------- -------------------A r te m ia ia - A r t t m i e i a - HrtemTai <iA gropyron A g r o p y n n /I. ami t h i t H outeloua
a m ith ii
ep in a tu m
(17)
(23)
(23)
C 0
C 0
C 0
AGROPYRON SPICATUM
A. e p io a tu m
M uhlen b erg ia o u e p id a ta
ARTEMISIA TRIDENTATA
A. t r i d e n t a t a
A. f r i g i d a
AGROPYHON SMITHII
A. a r ie ta tu m
A. e m i t h i i
B o u te lo u a g r a o i l i e
K o e le r ia o r i e t a t a
S a h edonnardue p a n io u la tu e
S tip a v ir id u la
C arex f i l i f o l i a
POA SECUNDA
P. eeaunda
A jn p y n n
C
0
47/99
24/99
49/98
14/71
5/29
51/98
17/100
51/97
15/67
12/49
2/11
5/22
TM
8/26
1/9
8/30
T/6
5/19
4/30
6/36
2/9
2/20
27/86
3/12
3/28
T/3
6/32
2/11
26/74
25/76
3/25
2/12
2/10
2/5
62/99
25/99
41/93
19/74
14/49
58/99
27/99
49/99
20/8)
12/52
13/53
6/27
7/31
T/6
Tj1
fIf
23/100
89/100
5/70
1/20
17/55
83/100
13/98
14/56
14/61
8/38
52/97
32/99
60/99
46/85
21/57
2/10
3/10
2/9
1/5
T/2
T/3
T/3
1/5
T/2
2/19
3/17
1/5
TM
2/7
T/6
1/4
3/9
9/37
28/74
1/4
1/14
T/T
4/31
2/7
4/3)
T/8
4/20
T/3
TM
6/25
2/7
14/34
2/7
5/22
1/6
T/5
1/22
T/l
TM
%
7/21
T/2
20/3)
T/2
T/3
T/12
T/3
9/23
T/7
I/ll
T/5
7/6
1/6
10/39
13/27
T/5
9/32
T/8
2/31
T/l
TM
10/26
T/3
T/9
T/8
10/28
T/6
1/13
1/8
JUNIPERUS SC0PUL0RUM
J . eeopulorum
Rhue t r i lo b a ta
C arex g e y e r i
1/2
T/2
2/9
T/T
T/l
T/2
PRUNUS VIRGINIANA
Roea n u tka n a
T/l
T/l
1/10
SYMPHiRICARPOS OCCIUfNTALIS
A r te m ie ia ea rn
Sym ph o ria a rp o e o e o i d e n ta lia
A g n p y r o n tra e h y e a u lu m
Poa eom preeea
T/5
T/l
T/3
2/15
5/19
2/19
T/2
T/5
4/55
T/14
16/32
T/2
T/5
T/5
T/7
T/2
TM
3/14
11/30
T/5
T/5
TZT
T/T
T/l
3/21
1/5
2/10
10/47
T/7
T/10
5/30
T/T
T/l
1/4
TM
T/l
T/l
2/5
T/5
1/7
10/35
T/T
2/5
T/2
10/33
2/3
T/T
%
T/8
T/5
41/69
3/28
27/83
T/5
T/2
11/33
T/l
T/l
T/2
2/3
T/2
T/3
T/5
5/22
TM
1/6
9/31
2/6
TM
2M
3/15
4/19
T/T
6/20
T/2
T/3
2/10
T/l
T/2
T/l
T/l
T/l
T/2
T/8
8/55
T/5
T/3
T/l
20/40
1/5
T/2
T/l
T/l
T/5
3/6
40/88
76/100
7/98
35/100
46/98
1/10
TM
2/8
T/3
A g ro p yro n S ym p h o ri- Xanthium
narpoa
et-ruim rii.
(16)
(2)
C O
C 0
67/99
21/100
46/94
17/68
10/38
8/34
T/6
7/18
XANTHIUM STRUMARIUM
G r in d e lia e q u a rro ea
Iua a x i l l a r i e
Rumex mexicanuB
X anthium etru m a riu m
Hordeum ju b a tu m
/.Il/BU
14/49
11/35
T/9
SAfiCOBAWS VERMICULAWS
S . v e r m ic u la tu e
D ie tie h lie s tr ia ta
Par ,-] ‘tan i-i- C xiroabatua- A r te m ie ia
A gropynn
lo n jifo lia
J u td p rv u a
(24)
(I)
C 0
C 0
21/58
8/34
T/3
T/3
1/24
ARTEMISIA LOHCIFOLIA
A. Ia n g if o l i a
C a la m o v ilfa l o n g i f o l i a
A r te m ie iu
(13)
C 0
29/68
TM
TRAGOPOGON DUBIUS
A r te m ie ia lu d o u ic ia n a
A e te r cotm u t a Cue
Comandra um bel l a t a
L a o tu ca p u I e h e I l a
M e lilo tu a o f f i e i n a l i e
O p u n tia p o ly a e a n th a
P e o ra le a Orgophd I la
V ie ia am eriaana
J u n ip r r u a
(9)
C 0
2/18
3/23
2/18
3/20
5/49
5/33
OTHER IMPORTANT PLANTS WITH LESSER VALUES WERE:
AFTEHISIA TRIDENTATA-UMlOH
T M : HVMN OVfiJUGr UNI ON
A rh i I l e i mi I I r ] '. Zturn
Ikih iu of![XieLtif o l i a
A a tr a g a lu e b i e u la a tu e
A tr ip le x n u t t a l l i i
C hryaothaim ue nauaeoaue
G u tie r r e s iu e o r o th r o e
P h lo x h o o d ii
rStmm poorimo
L a rtu a a e e r r i u l a
PetuloBtxm um aanariulum
S p h a e ra ln e a m a n in e u
Stephanom ori I r u n r iu n ta
Tnjopo-JO n JuOiun
AGROPYRON SHITHlI-VHlOH
A. a r i a to turn
S t i p a coma to
G ly n y r r is h a le p id o ta
R u tib id n n o lurnnif e n
AHnW S I A LUNGlPOL lA - W ION
E rio j onum mu I t ini
Orynope in hymenoi-laa
Yunna j launn
mmm.
Hoaa urkam uina
S o Iid u j o m ie iu -u r iin n ia
THeimtopaiit rhom hif o l i a
POA SECUNDA-WIOH
A lliu m t e x t i l e
M ic r o e e r ie nuU ine
P la n ta g o e p in u lo e a
YMI’IIOHI IiAHPOS OCCJPfJtTJMISr MJON
W
W
V M lJ J A M -U N lO H
V. iHnilniima
fo n rin y ia o r ie n ta lia
H e liu n th u a p a t i o l a r i a
XMTHiyM JTfMAhJGM-UU ION
Elymun irm n/lehttir.
S fx ir tim p e n tim ta
-
TABLE 29.
Year
76
-
SUMMARY OF THE GRAZING SEASONS FROM 1972 TO 1974 FOR EACH
PASTURE AND TOTALS FOR NICHOL'S COULEE RCA.
Pasture I
Pasture II
Pasture III
Pasture IV
D
rest
rest
C
8/11-12/5
2,555
B
5/17-11/15
2,785
A
4/1-11/15
6,479
Total
1972
Treatment
Dates used
AUMs used
AUMs per
rated AUM
rest
.74
.93
2.48
4/1-12/5
11,819
.96
1973
Treatment
Dates used
AUMs used
AUMs per
rated AUM
A
4/1-11/20
6,632
D
rest
rest
C
7/20-11/30
3,357
B
5/28-11/5
3,942
2.20
rest
1.12
1.50
B
6/2-11/10
3,054
A
4/1-10/24
5,841
D
rest
rest
C
7/31-11/25
3,326
1.01
1.70
rest
1.27
4/1-11/30
13,931
1.13
1974
Treatment
Dates used
AUMs used
AUMs per
rated AUM
4/1-11/25
12,221
.99
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