Range relationships of mule deer, elk and cattle in a rest rotation grazing system during summer and
fall
by Craig James Knowles
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 Craig James Knowles (1975)
Abstract:
A study was conducted in the timbered breaks adjacent to the Missouri River, northcentral Montana,
during the summers of 1973 and 1974 and the fall of 1974 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. Canopy coverages
and frequencies of occurrence were determined for low growing plant taxa in each of the plant
communities. Numbers, productivity, distribution, and range use of mule deer and elk were determined
from two early winter helicopter surveys and regular ground observations. Fawn:doe ratios were 50.0
and 54.8 for mule deer and calf:cow ratios were 65 and 70 for elk, respectively, during the two years of
the study. Numbers and distribution of mule deer within and between years showed no consistent
trends in relation to grazing by cattle and pasture treatments. Numbers and distribution of elk using the
study area within and between years were greatly influenced by grazing of cattle and previous pasture
treatments. Elk avoided areas of heavy use by cattle. Home ranges of three radio marked mule deer
were largest for the two males and smallest for the female. Home ranges of four radio marked elk were
substantially larger than those of mule deer, with home ranges of males larger than females.
Observations of marked elk indicated that previous and current grazing treatments of a pasture were
influential in determining use of a home range within a pasture by elk. The Pinus-Juniperus habitat type
was the most important for use by mule deer and elk during both summer and fall, while the
Artemisia-Agropyron habitat type the most important for use by cattle during both summer and fall.
Mule deer and elk made intensive use of steeper slopes and side ridges while cattle used main ridgetops
and major coulee bottoms most intensively. Food habits were determined from feeding site
examination and supplemented by rumen analysis. Forbs, browse, grass and browse, forbs, grass was
the order of importance of forage classes used by mule deer in summer and fall, respectively. Forbs,
grass, browse was the order of importance of forage classes used by elk in both summer and fall. Grass,
forbs, browse was the order of importance of forage classes used by cattle in both summer and fall.
Yellow sweetclover was the most important forb in the diet of each of the three ungulates. Use by mule
deer was shifted to browse and use by elk was shifted to grass as forbs were desiccated. Differences in
use of forage classes by mule deer between pastures in July 1974 was a result of previous grazing
treatments affecting reproductive success and subsequent abundance of yellow sweetclover.
Interspecific relationships, effect of rest rotation grazing and management implications 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
be allowed without my written permission.
Signature
Date
RANGE RELATIONSHIPS OF MULE DEER, ELK AND CATTLE
IN A REST ROTATION GRAZING SYSTEM
DURING SUMMER AND FALL
by
CRAIG" JAMES KNOWLES
A thesis submitted in partial fulfillment
of the requirements for the .degree'
of
MASTER OF SCIENCE
in
Fish and Wildlife Management
Approved:
Chairman, Examining Committee
Graduate yDean
MONTANA STATE UNIVERSITY
Bozeman, Montana
June
1975
ill
- ACKNOWLEDGMENT
. To the following, among others, I wish to express my sincere
appreciation for their contributions to this study:
Dr. Don C. Quimby.
and Dr. Richard J. Mackie, Montana State University, for project
planning, technical advice and guidance in preparation of the manu­
script; Mr. R. Bruce Campbell, and other personnel of Region Six,
Montana Fish and Game Department, for field assistance and cooperation;
Mr. Kenneth R. Greer, Montana Fish and Game Department, for lab assist­
ance and use of facilities; personnel of the Bureau of Land Management,
Malta District Office, for their cooperation; and to personnel of the
Lazy J D Cattle Company for their cooperation.
During the study, the
author was supported by the Montana Fish and Game Department under
Federal Aid Project W-I2O-R-5 and W-120-R-6.
iv
TABLE OF CONTENTS
Page
V I T A ..................................................................ii
ACKNOWLEDGMENT .......................................
ill
TABLE OF C O N T E N T S . .............................
iv
LIST OF T A B L E S ............................................
LIST OF FIGURES.
................. ..
vii
. . . ....................... xii
ABSTRACT ...................................................
xiv
INTRODUCTION . ........................................
DESCRIPTION OF STUDY AREA.
. , .......................
I
. . . . . .
2
METHODS..................... .. .....................................
9
Habitat Analysis ................... . . . . . . . . . . . . .
9
Populations and Range Use..............
9
Food Habits....................................................... 11
RESULTS.,........................................................
.
12
Habitat Types................... ' ..................... ..
Artemisia-Agropyron Habitat Type ............
. . . . . .
Aptemisiatridentata-AgropyTonspioatim community. . .
Artemisia tridentata-Agropyron smithii community . . . 1
Artemisia tridentata-Agropyron smithiiBouteloua graailis Community ..............
Pinus-JuniperusHabitat Type ..................
Pinus'ponderosa-Agropyron spioatum Community . . . . .
Pinus ponderosa-Juniperus soopulorum Community . . . .
Pinus ponderosa-Artemisia longifolia Community . . . .
. Pseudotsuga-Juniperus Habitat Type
;
Sarcobatus-Agropyron Habitat Type.........................
. Artemisia longifolia Habitat Type................
Agropyron-Symphoricarpos Habitat T y p e . ...................
■ Xanthiwn strumarium Habitat Type . . . . . . . . . . . . .
Miscellaneous Habitat Types
........................ . . .
12
12
12
6
18
20
20
21
21
2.3
25
26
26
28
28
V
TABLE OF CONTENTS
(Continued)
Page
Populations and Range U s e .....................................
Mule D e e r .........................................
'Numbers, Productivity and Distribution................
Home Range, Movements, Use Patterns and
Relationships.................................
Home Range S i z e ................................ .
M o v e m e n t s .............................
Use Patterns and Relationships................
Group Characteristics..........................
Activity....................... .. . . ...............
Use of Habitat T y p e ................. ' ...............
■ Summer.......................................
Fall................................................
Use of Slope and E x p o s u r e .............................
E l k ......................................
Numbers, Productivity andDistribution. . . . . . . .
Home Range, Movements, Use Patterns and
.
Relationships ......................... . . . . . .
Home Range S i z e ...................................
M o v e m e n t s ..............................
Use Patterns and Relationships....................
Group Characteristics.......... .. . ...............
Activity......................................
Use of Habitat T y p e ...............................
Summer......................................
Fall........................................
Use of Slope and E x p o s u r e ......................
Cattle. .'..........................................
Numbers and Distribution...............................
M o v e m e n t s .............................................
Group Characteristics .............................
Activity........................................
Use of Habitat T y p e .......................
S u m m e r ..............................
F a l l . ............................
Use of Slope and Exposure ......................
Food Habits ..............................................
Mule D e e r .......................................
Summer. ............................
Fall. . . . . . . . . . . . .
........ . . . . .
30
30
30
33
33
35
36
40
42
44
44
46
47
48
,
56
56
56
59
64
66
67
67
70
7.1
72
72
76
77
78
80
80
83
85
85
90
vi
TABLE OF CONTENTS
(Continued)
Page
Elk . . .
Summer
• Fall.
Cattle. .
Summer
' Fall.
DISCUSSION.
91
91
95
98
98
100
. ....................................................... 102
Interspecific. Range Relationships........................... '
Mule Deer and Cattle..................................... .
Elk and Cattle..................
Mule Deer and E l k .................................
Comparison of Rest Rotation Grazing to Season Long Grazing. .
Management Implications . ..........................
LITERATURE CITED.
. .............................
102
102
104
106
107
109
HO
vii
LIST OF TABLES
Table
Page
1.
Land status and grazing capacity of Nichols Coulee.RCA . .
5
2.
Mean temperature and precipitation for 1973 and 1974
together'with the 11-year means. U. S . Department of
Commerce Weather Station Roy 24 NE (Mobridge), Montana . .
8
3.
4.
5.
6.
7.
Mean percentage canopy coverage (C) and frequency of
occurrence (0) of plant taxa which obtained a mean
coverage of I percent or more in one or more of the
eleven plant communities. 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.......... ..
13
Mule deer numbers by sex," age class and locality during
January 1974 and 1975-helicopter surveys of Nichols
Coulee R C A .......................................
31
Summary of home ranges and movements during late
summer and early fall for three radio marked mule deer . .
34
Differential use of areas of home range by three radio
marked mule deer .................................
38
Mean monthly and seasonal mule deer group sizes observed
during the study by sex and for all mule deer combined . .
41
8 . A summary of characteristics of groups associated with
9.
10.
three radio marked mule d e e r .......... ..................
42
Seasonal percentages of all activities of mule deer
observed during the study within each habitat type and
total seasonal averages. ..................................
43
Monthly and seasonal percentages for all use of habitat
type by mule deer observed during the study for each ,
habitat type and percentages of use by mule deer for .
each community within a habitat type. Trace (T) amounts
are percentages less than I p e r c e n t . .......... ..
45
viii
LIST OF TABLES
(Continued)
Table
11.
12.
13.
14.
15.
16.
17.
18.
19.
Page
Seasonal percentages of all use of slope by mule deer
observed during the study within each habitat type and
total seasonal averages...........................
48
Seasonal percentages of all use of exposure by mule
deer observed during the study within each habitat
type and total seasonal a v e r a g e s .......... .. ...........
49
Summary of elk observations and classifications and on
and adjacent to Nichols Coulee RCA - helicopter survey,
early January 1974 and 1975...............................
51
Ratios between July and August 1973 and July and August
1974 for numbers of mule deer and elk observed within
Nichols Coulee RCA during these periods. . . . . . . . . .
53
Percentage distribution by pastures, month, and season
of all elk groups observed in 1973 and 1974, together
with percentage distribution of five marked elk when
observed inside Nichols Coulee RCA. Number of groups
of elk appear in parenthesis. Data includes elk groups
that were not classified.
. ................
54
Home range size and use of home range by locality for
five of six marked elk in Nichols Coulee RCA. Radio
channel number appears in parenthesis........ ..
57
Average distances between successive relocations of six
marked elk using Nichols Coulee RCA for 1974, for season
and two periods in fall, together with the maximum
distances between any two observations. Number of
relocations are in parenthesis ............. .. ...........
58
Mean monthly and seasonal elk group sizes observed "during
the study by sex and for all elk combined.
^
.
66
Coefficients of associations for six marked elk using
Nichols Coulee RCA during 1974 ........ . . . . . . . . .
67
XX
LIST OF TABLES
(Continued)
Table
20.
Page
Seasonal percentages of all activities observed during
this study within each habitat type and total seasonal
averages...................................... ; .......... ..
68
21.
Monthly and seasonal percentages for all use of habitat
type by elk observed during the study for each habitat
type and percentages of use by elk for each community
within a habitat type. Trace (T) amounts are percentages
less than I p e r c e n t ........ ............ .................. 69
22.
Seasonal percentages of all use of slope by elk observed
during the study within.each habitat type and total
seasonal averages ........................... .. . . . . . .
72
Seasonal percentages of all use of exposure by elk
observed during the study within each habitat type and
total seasonal averages ................... . . . . . . . .
73
Summary.of the grazing, seasons from 1972 to 1974 for
each pasture and totals for, Nichols Coulee RCA. ., ....
74
23.
24.
. .
25.
Summary of movements for four recognizable cattle ........
77
26.
Mean monthly cattle group sizes observed during the
study by pasture and year and mean monthly: arid seasonal
group sizes, for all cattle c o m b i n e d ........ ■. '. L . . . .
78
Seasonal percentages of all activities of cattle observed
during, the study within each habitat type and total
seasonal averages . . . . . . .
. . .... . .". .
79
Monthly and seasonal percentages for all use of habitat .
type by, cattle observed during the study for each habitat
type and percentages of use by cattle for each community
within.a. habitht type. Trace (T) amounts aria percentages
less than I percent ..........................
81
Seasonal percentages of use of slope by cattle observed
during the study within each habitat type and total
seasonal averages. . . . . . . . . . . . . . . . . . . . . . . .
83
27.
28.
29.
X
LIST OF TABLES
(Continued)
Table
30.
31.
32.
33.
34.
35.
Page
Seasonal percentages of all use of exposure by cattle
observed during the study within each habitat type
and total seasonal averages...............................
84
Seasonal frequencies of occurrence (0), aggregate mean
percentages of use (U) and percent canopy coverage
(C) by habitat type and by all types combined for each
plant species used by mule deer which accounted for I
percent- or more of the use recorded at feeding sites in
one or more habitat types. Trace (T) amounts are less
than I percent. Numbers of feeding sites examined are
in pa r e n t h e s i s ......................... ..................
86
Frequencies of occurrence (0), aggregate mean percentages
of use (U) and canopy coverage (C) by pasture for each
important plant taxa used by mule deer at feeding sites
examined during July 1974............... . ...............
89
Frequency of occurrence (0) and mean aggregate volume
percentage (V) for each plant taxa and forage class which
occurred in three or more of 25 mule deer rumen samples
collected in October and November 1974. Trace (T) amounts
are less than I percent'. . ............. .. ................
92
Seasonal frequencies of occurrence (0), aggregate mean
percentages of use (U) and percent canopy coverage (C)
by habitat type and by all types combined for each
plant species used by elk which accounted for I percent
or -more of the use recorded at feeding sites in one or
more habitat types. Trace (T) amounts are less than I
percent. Numbers of feeding sites examined are in
. parenthesis.......... ........................... ..
Frequency of occurrence (0) and mean aggregate volume
percentage (V) for each plant taxa and forage class
which accounted for I percent or more of the volume in
one o r :more of four elk rumen samples collected in
September 1973 and December 1974. Trace (T) amounts
are less than I percent. . . . . . . . . ..........
93
97
xi
LIST OF TABLES
. (Continued)
Table
36.
37.
Page
Seasonal frequencies of occurrence (0), aggregate mean
percentages of use (U) and percent canopy coverage (C)
by habitat type and by all types combined for each
plant species used by cattle which accounted for I
percent or more of the use recorded at feeding sites in
one or more habitat types. Trace (T) amounts are less
than I percent ........................... . . . . . . .
Frequency of occurrence (0) and mean aggregate volume
percentage (V) for each plant taxa and forage "class
which accounted for I percent or more of the volume in
one or more of four cattle rumen samples collected
August through November 1974. Trace (T) amounts are
less than I percent....................... ..
99
101
xii
LIST OF FIGURES
Figure
Page
1.
Map of the study area showing boundaries and drainages . .
3
2.
Grazing formula for Nichols Coulee RCA . . ...............
6
3.
Order of treatments for pastures in Nichols Coulee RCA
within and between years .................................
7
4.
Artemisia tridentata-Agropyron spioatum community....... 17
5.
Artemisia tridentata-Agropyron smithii community ........
17
6 . Artemisia tridentata-Agropyron smithii-Bouttelou
graeilis community ........................................
7 . Finns ponderosa-Agropyron spioatum community
19
19
8.
Finns ponderosa-Juniperus sooputornm c o m m u n i t y ......... .
22
9.
Finns ponderosa-Artemisia longifolia community . . . . . .
22
10.
Pseudotsugg menziesii-Juniperus sooputornm community . . .
24
11.
Saroobatus vermioulatus-Agropyrgn smithii community,
...
24
12.
Artemisia longifolia community ...........................
27
13.
Agropyrdn smithii-Syrrrphorioarpos oooidentalis community.
14.
Xanthium strumarium community.
15.
Distribution of mule deer January 1975-helicopter survey .
1.6.
Home ranges of three radio marked mule deer showing
specific areas and burned areas within their home ranges
(see Table 6, and Text)................................... 37
17.
Distribution of elk during specific periods of the study . 52
18.
. . . . . . . . . .
.
27
........ 29
Home range and movements of the radio marked 2.5-year-old
■male e l k .......... .................. ..
32
61
xiii
LIST OF FIGURES
(Continued)
Figure
19.
20.
21.
22.
. Page
Home range and movements of the radio marked 8-10-yearold female e l k ..............
62
Home range and movements of the radio marked 4-6-yearold female elk . . .......................................
63
Home range and movements of the neck-banded 2.5-yearold male e l k .................................
65
Generalized distribution of cattle density for each
year of the study.................................
75
ABSTRACT
A study was conducted in the timbered breaks adjacent to the
Missouri River, northcentral Montana, during the summers of 1973 and
1974 and the fall of 1974 to obtain quantitative data on populations,
range use and .food habits of mule deer, e.lk and cattle within an area
managed by rest rotation grazing.
Seven major habitat types consisting
of eleven plant communities were recognized. Canopy coverages and
frequencies of occurrence were determined for low growing plant taxa in
each of the plant communities. Numbers, productivity, distribution, and
range use of mule deer and elk were determined from two early winter
helicopter surveys and regular ground observations. Fawn:doe ratios
were 50.0 and 54.8 for mule deer and calf:cow ratios were 65 and 70 for
elk, respectively, during the two years of the study.. Numbers and dis­
tribution of mule deer within arid between years showed no consistent
trends in relation to grazing by cattle and pasture treatments. Numbers
and distribution of elk using the study area within and between years
were greatly influenced by grazing of cattle and previous pasture treat­
ments. Elk avoided areas of heavy use by cattle. Home ranges of three
radio marked mule deer were largest for the two males and smallest for
the female. Home ranges of four radio marked elk were substantially
larger than those of mule deer, with home ranges of males larger than
females. Observations of marked elk indicated that previous and current
grazing treatments of a pasture were influential in determining use of
a home range within a pasture by elk. The Pinus-Juniperus habitat type
was the most important for use by mule deer and elk duririg both summer
and fall, while the Artemisia-Agropyron habitat type the most important
for use by cattle during both summer and fall. Mule deer and elk made
intensive use of steeper slopes and side ridges while cattle used main
ridgetops and major coulee bottoms most intensively. Food habits were
determined from feeding site examination and supplemented by rumen
analysis. Forbs, browse, grass and browse, forbs, grass was the order
of importance of forage classes used by mule deer in summer and fall,
respectively. Forbs, grass, browse was the order of importance of
forage classes used by elk in both summer and fall. Grass, forbs,
browse was the order of importance of forage classes used by cattle in
both summer and fall. Yellow sweetclover was the most important forb
in the diet of each of the three ungulates. Use by mule deer was
shifted to browse and use by elk was shifted to grass as forbs were
desiccated. Differences in use of forage classes by mule deer between
pastures in July 1974 was a result of previous grazing treatments
affecting reproductive success and subsequent abundance of yellow
sweetclover. ' Interspecific relationships, effect of rest rotation
grazing and management implications of mule deer, elk and cattle on a
rest rotation grazing system were discussed.
INTRODUCTION
Rest-rotation grazing systems (Hormay and Talbot, 1961) have
become increasingly popular in recent years as a means of managing
.
rangelands grazed by livestock.
These systems, by incorporating periods
of rest into the grazing formula, provide conditions which favor the
development of desirable plant composition and abundance in spite of
intensive grazing during certain periods within or between years.
Personnel of the Bureau of Land Management, in 1965, initiated a restrotation grazing system for cattle (Bos taurus) in the Nichols Coulee
Resource Area (RCA) located bn and adjacent to the Charles M. Russell
Game Range in North-Central Montana.
Both mule deer (Odooovleus
hemionus) and elk (Cemus canadensis) also use parts, of this area dur'i
■
'
'
■
'
ing at least some seasons. Mackie (1970), reported on mule deer, elk
and cattle; range relationships for the Missouri River Breaks; the area
that includes part .of the RCA.
At the time of his study no rest-
rotation grazing systems were in operation.
Consequently he was
unable to observe the effects of rest-rotation systems for cattle bn
the range ecology of mule deer and elk.
This study, conducted on a full time basis during the.summer of
1973 and the.summer and fall of 1974, considered the range ecology of
mule deer, elk.and cattle within the pastures included in the system.
DESCRIPTION OF THE STUDY AREA
The study area of approximately 88,810 acres is located in South
Phillips County 55 miles southwest of Malta, Montana.
shown in Figure I.
Boundaries are
The area is part of a highly dissected plateau
which slopes gradually to the southeast.
Elevations range from 3,000
feet along the north boundary to 2,246 feet at the Missouri River.
Breaks topography extends from the river, north, to the Charles M.
Russell Game Range boundary which roughly delineates the end of the
timbered breaks.
From here, rolling plains develop.
Soils, mostly heavy clay loams of. the Lismas, Pierre and Phillips
series, are derived from the underlying Bear Paw Shale Formation
(Gieseker, 1926).
Shale outcrops are commorr, especially in the breaks
portion of the study area.
Alluvium from the Little Rocky Mountains
to the north occurs in isolated areas.
evidence of feeble glaciation.
Higher, level ridgetops show
Runoff from such ridges, is. high.
Major coulees, characterized by much meandering in a well develop­
ed flood plain, cut through shale outcrops and alluvial benches.
These coulees contain.intermittent streams and are subject to flash
floods during periods of heavy rains.
In addition to natural springs
and seepagesv nine wells and 83 stockponds have been developed on the
study area.
NicKols Coulee R C A .is a four pasture system.
Grazing is adminis­
tered by personnel of the Bureau of Land Management.
Land ownership.
-3-
Pe «t. I
hemp Creek
C M 1RusseII H
Oeme Ronge
Pest. IV
Post III
Missouri River
Legend
Nichols Coulee RCA
Creek or Coulee ---Holding posture ----------- hp
Figure I.
Map
of the study area showing boundaries and drainages.
-4acreages and AUMs for the study area are shown in Table I.
ing formula is shown in Figure 2.
counter-clockwise (Fig. 3).
clockwise (Fig. 3).
The graz­
Order of treatments within a year is
Rotation of treatments between years is
Pasture II lacks adequate natural shelter for
calving; therefore the years pasture II is grazed early, cows heavywith-calf are often turned into pasture I April I.
Pastures I and II are mostly rolling uplands dissected by a
dendtritic pattern of side coulees which flow into a few major drain-t­
ag es.
For the mdst part, relief is low.
Pastures III and IV are
strikingly different from pastures I and II.
Relief increases to as
much as 400 feet immediately north of the river; ridges become narrower
and break sharply into deeply entrenched drainages.
..
Climatological data were obtained from the United States Depart­
ment of Commerce Weather Station Roy 24 NE (Mobridge), located 17 miles
southwest of the study area center.
Mean monthly and yearly tempera­
tures and precipitation for the two years of the study,.as well as
the eleven year means, are shown in Table 2.
Climate of this area.is
considered semi-arid with short hot summers and long cold winters
(Gieseker, 1926).
Eighty percent of the precipitation comes in the
half year period of April through September.
Despite above average
precipitation in 1973, little rain fell between mid-June and late
August.
Mean number of frost free days is 118.
this figure was 117 and 102 days respectively.
For 1973 and 1974
Table I.
Land status and grazing capacity of Nichols Coulee RCA.
Total foir Area
AUMs
Z
Pasture I
Acres
AUMs
Pasture II
AUMs
Acres
Pasture III
AUMs
Acres
100.0
17,352
3,015
20,516
3,431
28,913
2,984
21,109
2,607
920
234
9,795
79.8
14,352
2,466
13,726
2,144
27,040
2,728
19,809
2,453
20
4
9.3
1,534
12.5
1,720
326
4,996
873
593
99
40
6
900
230
6.3
942
7.7
1,280
223
1,794
414
1,280
157
1,260
148
0
0
Ownership of
Control
Acres
Total
88,810
100.0
12,271
Public Domain
74,947
84.4
Private
8,249
Lease
5,614
Z
Pasture IV
Acres
AUMs
Holding Past.
Acres
AUMs
Month
TREATMENT
REST FOR VIGOR
GRAZE
; GRAZE
RESI FOR SEEDRII E (STVI)
RESI FOR I;STABLI SHMENT OF REPRODUCTION
Figure 2.
Grazing formula for Nichols Coulee RCA
—7—
Years combination occurred
1965
1969
1973
PASTURE I
PASTURE II
TREATMENT A
TREATMENT D
PASTURE IV
TREATMENT B
PASTURE III
TREATMENT C
Years combination occurred
1967
1971
Years combination occurred
1966
1970
1974
PASTURE I
%
TREATMENT A
PASTURE IV
TREATMENT C
PASTURE II
TREATMENT A
PASTURE III
TREATMENT D
Years combination occurred
1968
1972
PASTURE I
PASTURE II
PASTURE I
PASTURE II
TREATMENT C
TREATMENT B
TREATMENT D
TREATMENT C
PASTURE IV
TREATMENT D
Figure 3.
PASTURE III
TREATMENT A
PASTURE IV
TREATMENT A
PASTURE III
TREATMENT B
Order of treatments for pastures in Nichols Coulee RCA
within and between years.
— 8—
Table 2.
Mean temperature and precipitation for 1973 and 1974 together
with the 11-year means. U. S . Department of Commerce Weather
Station Roy 24 NE (Mobridge), Montana.
Temperature (degrees Fahrenheit)
Month
Year
Jan.
Feb.
Mar.
Apr.
May
June
July
Aug.
Sept.
Oct.
Nov.
Dec.
1973
20.5
27.2
38.2
42.7
55.8
64.9
70.9
72.4
58.2
49.2
23.0
23.0
1974
17.5
29.7
33.1
46.9
50.4
68.0
73.7
64.8
55.6
48.2
34.5
26.9
Totals
45.5
45.8
1No data Jan. and Feb. 1964.
11-Year1
Mean
11.7
22.1
31.8
44.1
55.1
64.3
71.5
70.7
57.6
47.1
31.2
17.3
43.8
Precipitation (inches)
1973
T
.11
.16
2.55
1.33
3.97
.52
4.12
2.27
.63
.53
.30
Year
1974
.12
.10
.83
.81
4.44
2.08
4.82
4.09
.61
.65
.41
.04
ll-Ye;
Mean
.58
.21
.57
1.36
2.02
2.80
1.69
1.90
1.06
.58
.32
.43
16.49
19.00
13.52
METHODS
Habitat Analysis
Habitat types of the study area were generally similar to those
described by Mackie (1970).
To verify similarities, 23 transects were
established in six major habitat types.
The method of analysis follow­
ed the canopy method.of Daubenmire (1959).
Twenty 2x5 dm plots for
grasses, forbs and shrubs and twenty 4x10 dm plots (Pyrah 1973) for
shrubs only were placed at 10 foot intervals along each of two 100 foot
lines at each sample site.
The sample units were placed in a relative­
ly homogenous and undisturbed portion of each habitat type.
In
addition, plant frequency and coverage recorded at mule deer, elk and
cattle feeding sites were combined with transect data to provide a
better representation of each habitat type.
Most work was done in
pastures III and IV.
Plant nomenclature follows Booth (1950) and Booth and Wright
(1966).
.
' •
Populations and Range Use
Early winter surveys of pastures III and IV, made from a heli­
copter, were conducted each year to obtain accurate Counts and
classification of mule deer and elk.
Productivity, population
structure and numbers are based on these data.
-10Radio telemetry of mule deer and elk was used in 1974 to more
precisely determine range relationships and movements of these
ungulates.
This phase of the study was conducted in conjunction with
Region Six personnel of the Montana Fish and Game Department.
some cases elk were.marked with neck-bands only.
made both from the ground and air.
Relocations were
Generally, attempts at relocating
all animals were made at least once a week.
sidered a relocation between days.
In
An observation was con­
To more fully understand daily
activity and daily movements, each fadio marked animal was observed a
minimum of twice on at least one day.
Home range data were calculated
by the minimum area method (Mohr, 1947) and modified (Harvey, 1965)
whenever data were insufficient or otherwise.failed to justify connect­
ing the outlying points.
Nine vehicle routes were established.
Eight of these were in
pastures III and IV, giving almost complete coverage of these pastures.
Emphasis of this study was placed on these pastures where the major
mule deer and elk concentrations occurred.
All routes in pastures III.
and IV were covered systematically at about one week intervals during
morning and evening observation periods.
Observations for all animals
were recorded with respect to time, group size, activity,habitat type,
slope, exposure and location when first seen.
In addition, for mule
deer and elk, sex and age class were recorded whenever possible.
—11—
Animals within 100 yd. of each other and/or displaying definite group
relationships were recorded collectively.
Food Habits
Food habits data were obtained by examining and recording frequency
of use of various plant taxa at recently vacated feeding sites.
Utili­
zation of one rooted stem for grasses and grass like plants, one stem
or leaf for forbs, and one twig or leaf for shrubs constituted one
instance of use.
Minimas of 50 instances of use for mule deer and elk
and 100 for cattle on one habitat type at one site were the requisite
for a feeding site.
Frequencies of occurrence and coverages of plant
taxa present were recorded at each feeding site to provide a basis for
evaluating food preferences.
The method used was similar to that pre-_
v IousIy described for habitat analysis except that ten 4x10-dm plots
for shrubs were used and plots were placed randomly in the area of the
feeding site.
Rumen samples to supplement feeding site data were
collected whenever possible.
Hunter killed animals in the fall
provided the bulk of the samples.
The aggregate percentage method
(Martin et al., 1946) was used to tablulate data for feeding site
examinations by season and habitat type and for rumen samples.
RESULTS
Habitat Types
Seven of, eight habitat types described by Mackie (1970) for the
Missouri River breaks occurred on the study area.
The results of the
vegetational measurements for the habitats, arranged in a manner
similar to that used by Mackie, are presented in Table 3.
,
•
•■
Artemisia-Agropyron Habitat Type
This type was most characteristic of level ridge tops
but it was
also present on more level portions of the sides of ridges and in the
major coulee, bottoms located at distances of seven miles or more from ,
the Missouri River.
It occupied about 57 percent of pastures III. and
IV and almost the entire areas of pastures I and II.
Mackie described
three plant communities within this type as follows:
Artemisia tridentata-Agropyron spioatum Community
Well developed stands of this community were generally found along
edges of the main ridges, on the tops of side ridges (Fig. 4) and on
the more level portions of the sides of ridges.
Bluebundhwheatgrass
(Agropyron spioatum) and big sagebrush (Artemisia tridentaia) was the
most common grass and shrub, respectively.
Forbs of the Tragopogon '
union were, the most common, especially yellow sweetclover (Melilotus
BARE GROUND
LITTER
GRASS
FORBS
SHRUBS
(23)
C 0
62/99
25/99
41/93
19/74
14/49
(23)
C 0
58/99
27/99
49/99
20/83
12/52
Ii
If
Il
Artemisia
longifolia
III
1,
PinusArtemiaia
I
1 ill
III PI
PinusJuniperus
Mean percentage canopy coverage (C) and frequency of occurrence (0) of plant taxa which obtained a mean coverage of I percent or more
In one or more of the eleven plant communities. 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.
PinusAgropyron
Table 3.
Bi Il
Jl
(17)
C 0
47/99
24/99
49/98
14/71
5/29
(27)
C 0
51/98
37/100
51/97
15/67
12/49
(9)
C 0
40/89
44/100
42/93
24/67
29/68
(13)
C 0
70/99
23/100
27/77
23/58
8/34
(I)
C 0
7/60
89/100
5/70
1/20
17/55
(24)
C 0
67/99
21/100
46/94
17/68
10/38
(14)
C 0
83/100
13/98
14/56
14/61
8/38
14/49
11/35
8/34
T/6
2/19
3/17
4/55
2/10
3/10
2/9
1/5
T/2
T/3
1/5
T/4
2/7
T/6
1/4
3/9
T/3
1/5
T/2
9/37
T/8
4/20
T/3
T/4
14/34
2/7
28/74
1/4
1/14
T/3
5/22
(16)
C 0
52/97
32/99
60/99
46/85
21/57
(2)
C 0
76/100
7/98
35/100
46/98
1/10
UNION:TAXA
AGROPYROti SPICA TUM
A. Spioatum
Muhlenbergia cuspidata
ARTEMISIA TRIDEIiTATA
A . tridentata
A. frigida
13/53
6/27
2/11
5/22
T/4
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
AGROPYROti SMITHII
A. cristatum
A. smithii
Bouteloua gracilis
Koleria cristata
Schedonardus paniculatus
Stipa viridula
Carex fH i folia
7/31
T/6
POA SECUHDA
P. secunda
T/4
T/9
7/38
T/3
T/3
1/24
T/8
2/31
T/l
T/4
TRAGOPOGOti DUBIUS
Artemisia ludovicinana
Aster cormutatus
Comandra umbellatum
Lactuca pulchelW
T/T
4/31
2/7
4/33
6/25
2/7
5/30
3/6
40/88
20/40
1/5
T/4
T/4
10/33
2/3
T/4
T/4
T/8
T/5
3/28
2/8
1/6
T/5
1/22
T/4
T/l
T/4
T/5
T/2
1/5
T/14
T/3
T/T
T/3
T/12
T/3
T/5
T/2
Table 3.
Continued.
£
"S E 5
a K
-g g ..
a -5 q
-S ^ #
P
E- C-. Q.
Q
^
(23)
C O
(23)
C 0
10/26
T/3
T/9
T/8
10/28
T/6
1/1 3
1/8
JUNIPERUS SCOPULORUM
J. scopulorum
Rhus trilobata
Carex geyeri
1 /2
T/2
2/9
T/ T
T /l
T/2
PRUNUS VIRGINIANA
Rosa nutkana
T /l
T/5
TRAGOPOGON DUBTUS (cont)
Melilotus official
Opuntia polycantha
Psoralea argophylla
Vicia americana
SYMPHORICARPOS OCCIDENTALIS
Artemisia cana
Symphoricapos occidentalis
Agropyron trachyeaulum
Poa eompressa
ARTEMISIA LONGIFOLIA
A. Iongifolia
Calamovilfa longifolia
SARCOBATUS VERMICULATUS
S. Vermiculatus
Distiehlis stricta
UNTHIUM STRUMARIUM
Grindelia squarrosa
Iva Axillaria
Rumex mexicanus
Xanthium strumarium
Hordeum jubatum
l
$ 6 1
(17)
C 0
•2
E
H
SI
(27)
C 0
C
(9)
0
si
25
C
(13)
0
16/32
T/2
| |
It
C
(I)
0
T/ 5
T/7
11/30
T/ 5
T/5
T/ T
T/ T
T /l
7/6
1/6
10/39
13/27
T/5
9/32
T /l
1/1 0
3/21
1/5
2/10
T /l
T/ 3
T /l
2/1 5
10/47
T/7
T/10
T/ 5
T/ 2
T/4
3/14
(24)
C 0
9/25
T/7
20/33
T/ 2
T/ 5
E
EE
T/5
7/23
T/2
2/15
1/11
5/19
2/19
T/2
I Il
T/2
T/l
T/T
T /l
1/4
T/ 4
T/ 3
T/ 5
T/ 5
T /l
T /l
2/5
T/l
T/T
T/ 5
2/5
T/ 3
1/7
10/35
T/2
2/4
6/20
T/4
E&8
(14)
C 0
(16)
C 0
C__0
11/33
T/l
41/69
27/83
T/2
T/l
2/3
T/2
T/3
T/5
5/22
T/l
T/4
1/6
9/31
2/6
T/4
3/15
4/19
T/2
T/8
8/55
T/5
T/T
T/3
2/10
T/2
T/3
T/2
(2)
T/l
T/2
T /l
T/l
I
3g
55
2/8
T/3
Cl
T/l
T/l
2/18
3/23
2/18
3/20
5/49
5/33
Table 3.
Continued
Other important plants with lesser values were:
ARTEMISIA TRIDENTATA-VRION
Astragalus bisulcatus
Atriplex nuttallii
Chrysotharmus nauseosus
Gutierrazia sarothrae
Phlox hoodii
JUNIPERUS SCOPELORUM-VI!ICU
Rosa Arkansana
Solidago missouriensis
Thermovsis rhombir?lia
PRUIWS VIRGINIARA-UUIOll
P. virginiana
AGROPYRON SMITHII-URIOH
SYMPHORICARPOS OCCIDENTALIS-UIIICN
A. Cristatum
Stippa comata
Glyoyrrizha lepidota
Ratibida colurmifera
POA SECUNDA-UNION
ARTEMISIA LONGIFOLIA-UNIOIl
Allium textile
Miaroseris nutans
Plantago spinulosa
Eriogonum multiaeps
Oryzopsis hymenoides
Yucca glauca
TRAGOPOGON DUBIUS-UHION
SARCOBATUS VERMICVLATVS-UUIOH
Achillea millefolium
Bahia oppositifolia
Gaura cocainea
Laatuoa serriola
Petalostemum candidun
Sphaeralaea coccinae
Stephanomeria runcinata
Tragopogon dubius
Conringia orientalis
Helianthos petiolaris
XAHTHIUM STRUMARIUM-UNI0I1
Elymus canadensis
Spartina pectinata
-16officinalis) and bastard toadflax (Comandra vmbellata).
Greasewood
(Savcobatus vevmiculatus) end Rocky Mountain juniper (Juniperus
scopulovum) occurred on level portions of the sides of ridges where
community structure was often poorly defined, thus explaining their
occurrence in more than trace amounts.
Sampling on sides of ridges,
as well as on the tops of ridges, resulted in values for bare ground
that exceeded values for other communities of the Artemisia-Agropyron
habitat type, opposite of Mackie's findings.
Artemisia tridentata-Agropyron smithii Community
This community was found on sites similar to those occupied by the
Artemisia tridentata-Agropyron spioatum community.
best represented on.the main ridges (Fig. 5).
(Agropyron smithii) was the dominant grass.
However, it was
Westerhwheatgrass
Its relatively low value
for cover despite high percent occurrence probably resulted from the
sampling of many grazed areas.
The Poa union was better represented
here than i n .the Artemisia tridentata-Agropyron spioatum community.
The early spring phenology of the Poa union probably has resulted in
an underestimation o f ,its. importance in this community and in others.
Taxa of this union were rarely recorded in fall.
obtained its maximum abundance in this community.
Bastard toadflax
Big sagebrush
showed little.change from the Artemisia tridentata-Agropyron spioatum
community.
Transect data for both these communities suggested that
.
—17—
Figure 4
Figure 5.
Artemisia tridentata-Agropyron smithii community.
—18differences in coverage and frequency of big sagebrush between that
reported here and that reported by Mackie are related to sampling bias.
Artemisia tvidentata-Aqvoyyvon smitht-i-Boutetoua.avaoilis Community
The occurrence of this community in the breaks portion of the
study area was restricted to level ridge tops (Fig. 6).
In pastures I
and Il it was the dominant community, even occurring in coulee bottoms.
It was especially well developed on the Phillips clay loam.
These clay-
pan soils were covered with dense mats of blue grama (Boutetoua
graaitis)'.
Despite the appearance of severe sheet erosion this commu­
nity had the lowest value for bare ground for any community in the
Artemisia-Agropyron habitat type.
Local areas of much bare ground did
exist.. Blue grama, tumblegrass .(Sahedonngrdus paniculatus)\,_ fringed
sagewort (Artemisia frigida)3 prickly pear (Opuntia polyaantha)3. spindle
plantain (Plant'ego spinulosa) all obtained maximum abundance in this
-
community.
■
•
-
'
Big sagebrush, the dominant shrub, provided the least cover­
age for any of the communities of the Artemisia-Agrdpyron habitat type,
an opposite trend from Mackie's results.
was found to exist in this community.
No evidence of sampling bias
Therefore, the large amounts of
blue grama.and.low. abundance of big sagebrush suggested that, this
community on the north side of the Missouri River varies in species,
abundance from that described by.Mackie.,
-19-
Figure 6.
Artemisia tridentata-Agropyron smithii-Bouteloua gracilis
community.
Figure 7.
Pinus ponderosa-Agropyron spicatum community.
-20Pinus-Juniperus Habitat Type
This habitat type was characteristic of sloping ridge sides.
Exposure and degree of slope determined its development.
This habitat
type was almost entirely within pastures III and IV where it occupied
26 percent of the area.
Somewhat greater coverage was afforded in
pasture III where it extended to the game range boundary than in
pasture IV where it was only poorly developed in the upper one fourth
of this pasture.
Mackie described three plant communities within this
type as follows:
1
Pinus vonderosa-Aqroyyvorr sy-Leatim Community
This community occurred along borders of the Pinus-Juniperus
community where it meets Artemisia-Agropyron dominated ridgetops
(Fig. 7), in shallow basin-heads, and on narrow side ridgetops where
scattered growths of ponderosa pine (Pinus ponderosa) occurred.
The
open nature of the pine, canopy allowed much herbaceous growth to
develop.
The
Agropyron-spieatum union was well developed.
Plains
muhly (Muhlenb'ergia ouspidata) obtained maximum coverage and frequency
in this community.
Prairie sandgrass (Cdlamovitfa lorigifolio.) was
present, on occasional patches of unstable soil.
In the Pinus-
Juniperus habitat type, grass coverage and frequency were greatest
here.
' ..v
Rocky Mountain juniper was the dominant shrub..
■
. •'
.'
■
Skunkbrush
(Rhus triiobdta)s. sroA., elk sedge (Carexgeyeri) obtained maximum
-21abundance in.this community.
Ponderosa pine occurred as scattered
individuals.
Pinus Ponderosa-Jun-Cpevus soopuloTum Community
.
This community was best developed on slopes with at least some
north exposure (Fig. 8).
It did, however, occur on all exposures but
with decreasing frequency toward south exposures.
Ponderosa pine and
Rocky Mountain juniper were the dominant species.
Often dense thickets
of juniper formed almost to the exclusion of pine from the canopy.
This community had the least amount of bare ground and the most litter
of the Piniis-Juniperus'habitat type.
Bluebunch wheatgrass, western
wheatgrass and green needlegrass (Stipa viridula) were all equally
present in moderate amounts.
Yellow sweetclover grew exceptionally
well along the edges of this community.
In old burns, hootka rose
(Rosa nutkana) and western snowberry (Symphovioanpos oooidentalis) were
the common shrubs.
Sampling in burned areas and along edges of this
community resulted in greater than normal values of grasses, forbs,
and bare ground and greater than trace amounts of big sagebrush and
greasewood.
Pinus pondevosa-Aytemisid longifblia Community
This community occurred on steep, unstable shale slopes usually,
with a south aspect (Fig. 9).
It contained the most bare ground and
-22-
Figure 8
Pinus ponderosa-Juniperus soopulorum community.
Figure 9.
Pinus ponderosa-Artemisia longifolia community.
-23the least amount of litter of all communities in the Pinus-Juniperus
habitat type.
Such unstable conditions allowed the development of the
Artemisia longifolia union; sandgrass, soapweed (Yueoa glauaa), and
longleaf sagebrush (Artemisia longifolia) were its principal grass, forb,
and shrub, respectively.
community.
The former was the dominant grass in this
Yellow sweetclover appeared to do well on these unstable
slopes when proper environmental conditions existed. . The Shrub layer
in this community was poorly developed.
Ponderosa pine occurred as.
scattered individuals.
No line intercepts were, conducted in any of the Pinus communities.
However, of these three communities it was. felt that the Pinus
ponderosa-Juniperus seopulorum community probably varied most from
Mackie1s description.
Due to the limited distribution of the
Pseudotsuga-Juniperus habitat type on this study, area, the Finns
ponderosa-Juniperus seopulorum community had largely taken over its
place on north slopes with, the role of pine increasing in the canopy.
PseudotAuga-Juhiperus Habitat Type
This community almost without exception occurred on relatively
steep northerly exposures (Fig. 10)*
This minor habitat type was
restricted to pastures III and IV on the study area where it occupied
one percent of the terrain.
Maximum abundance was along the west.bound­
ary of the study area, while along the east boundary it was absent.
-24-
Figure 10.
Pseudotsuga menziesii-Juniperus scopulorum community.
Figure 11.
Sareobatus vermiaulatus-Agropyron smithii community.
-25Observations indicated that its importance increased immediately west
of the study area.
Results are based on only one transect.
age was recorded in this habitat type.
Tree cover
Accumulation of Douglas fir
CPseudotsuga menziesCi) and ponderosa pine needles resulted in the
least amount of bare ground and the most litter,
habitat type.
recorded in any
Grass coverage, which was low, consisted, almost entirely
of bluebunch wheatgrass.
Line intercept coverage was 17.8 percent,
21.5 percent and 54.5 percent for Rocky Mountain juniper, ponderosa
pine and Douglas fir, respectively.
type and in others where it occurred,
The Prunus union, in this habitat
. was usually poorly developed.
Nootka rose appeared to be the only abundant and wide spread member.
Chokecherry (Pvunus Vivg-LnCana) /was abundant on the study area only in
a few locations.
Sarcobatus-Agropyron Habitat Type
This habitat type occurred along bottoms of the Missouri River,
coulee bottoms (Fig. 11), benches and footslopes.
pastures III and IV were occupied by this type.
western wheatgrass.
About 6 percent of
The major grass was
Limited amounts of bluebunch wheatgrass and plains
muhly occurred on footslopes.
again been underestimated.
The importance of the Poa union has
Greasewood was the dominant shrub only .
along the river and for short distances north in the major drainages.
Big sagebrush gradually increased in importance with distance from the
-26river. and at about seven miles, the Artemisia-Agropyron habitat type
replaced the Sarcobatus-Agropyron habitat type.
Artemisi-d IongifoUa Habitat Type •
This habitat type occurred on unstable, steep shale slopes
(Fig. 12).
It was most common on south and south-west slopes; being
extremely conspicuous in the lower section of the breaks.
Approxi­
mately 4 percent of pastures III and IV were occupied by this type.
Of all habitat types examined, this type had the most bare ground.
Correspondingly, litter, grass, forb and shrub coverages were all
equally low.
Western wheatgrass and sandgrass were the major grasses.
Yellow sweetclover was the only forb present in more than trace amounts.
Eriogonum (Eriogonwn muttioepts)3 of the Artemisia Iongifolia union,
obtained its. maximum development on this habitat type.
brush was the major shrub.
Longleaf sage­
Greasewood, big sagebrush and skunkbrush
were also commonly found.
Agropyron-Symphoriearpos Habitat Type.
This habitat type occurred in the flood plains of major couless
(Fig. 13) and in minor drainage ways.
Combined with the Xanthivm
strvmarivm habitat type it occupied about 6 percent of pastures III
and IV.
Frequent flooding and subsequent siltation in the summer of
1974 resulted in bdreground and litter being over and underestimated,
respectively.
Grass and forb coverage were the highest of all habitat
-27-
-28types. Western snowberry was the dominant shrub.
In drainages through
burns, nootka rose co-dominated; in major coulee bottoms, stands of
green rabbitbrush (Ckpysothamus V-Lsoidiftovus) and silver sagebrush
(Artemisia oand) were occasionally dominant.
Rocky Mountain juniper
was recorded in minor drainages bordering the Pinus-Juniperus community.
Xanthivm strumarium Habitat Type
This was a minor habitat type, occurring in the actual cuts of
intermittent streams (Fig. 14).
Vegetation was sparse and generally
restricted to silty banks and areas not severely flooded.
Because only
two feeding, site examinations were conducted in this type sampling was
biased to areas of vegetation.
Though western wheatgrass and slender
wheatgrass (Agropyron traohyoalum) were the dominant grasses, Canada
wildrye (Elymus canadensis) and prairie cordgrass (Spartina peotinata)
were also common grasses.
Dominant forbs were of the Xanthium union.•
The shrub layer- consisted of an occasional green rabbitbrush.
Miscellaneous Habitat Types
Backwaters of the Fort Peck Reservoir extended onto bottomlands of the
Missouri River along the south boundary of the study, area.
Because of
this and plus the fact that water levels vary greatly within and
between years, normal river bottom vegetation, as described by Allen
(1968), was substantially altered.
During low water periods vegetation
-29-
Figure 14.
Xanthium strumarium community.
—30—
resembled that of the Xanthium union.
Foxtail barley (Hovdeum jubatum)
dominated on higher sites while forbs were prevalent on lower sites.
About 700 acres were affected.
In a few locations in major coulee bottoms, plains cottonwood
(Populus sargentii) was found in limited numbers.
was the prominent grass.
Western wheatgrass
No quantitative work was done in either of
these two minor habitat types.
Populations and Range Use
Mule Deer
Results are based on 660 mule deer groups totaling 960 mule deer
observed during the summers (June-August) of 1973 and 1974; figures for
the first half of September 1973 and the fall (September-December) of
1974 were 478 and 880, respectively.
A total of 385 mule deer was
counted during two aerial surveys, one each in January of 1974 and 1975.
Numbers, Productivity and. Distribution
v
•
Data from the January helicopter surveys of pastures III and IV
(Table 4) suggested that mule deer number's increased 9.2 percent from
1974 to 1975.
All this increase was accounted for in pasture III where
numbers increased 13.1 percent, partly as a result of increased numbers
of fawns.
Densities varied between areas (Fig. 15 and Table 4).
Density was 1.5 and 1.9 times greater in pasture III than in pasture
Table 4.
Mule deer numbers by sex, age class and locality during January 1974 and 1975 helicopter surveys of Nichols Coulee RCA.
Area
Males: Fawns: Fawns:
%
Year
Total Males Females Fawns 100
100
100
%
Yr l .
__________________________________________________ Females Females Adults Fawns Males
Pastures III & IV
1974
184
34
100
50
34.0
50.0
37.3
27.2 52.9
201
40
104
57
54.8
1975
33.5
39.6
28.4 43.6
Pasture III
1974
1975
Deer/
Sg. Mi.
3.1
3.4
145
164
29
32
76
82
40
50
38.2
39.0
52.6
61.0
38.1
43.9
27.6
30.5
62.1
51.6
3.4
3.9
39
37
5
8
24
22
10
7
20.8
36.4
41.7
31.8
34.5
23.3
25.6
18.9
___
12.5
2.2
2.1
HO
147
23
31
57
73
30
43
40.4
42.4
52.6
58.9
37.5
41.3
27.3
29.3
73.9
50.0
3.6
4.9
Nichols Coulee
1974
1975
35
17
6
I
19
9
10
7
31.6
11.1
52.6
77.8
40.0
70.0
28.6 16.7
41.2 100.0
3.0
1.4
Seven Mile Coulee
1974
1975
39
37
5
8
24
22
10
7
20.8
36.4
41.7
31.8
34.5
23.3
25.6
18.9
Pasture IV
1974
1975
CK Creek
1974
1975
___
12.5
2.2
2.1
Post. IV
• •••••
• •••••••
•••••••••
•••••••%•
• • • • • • »v
» ••••••
lySi/'f-
Legend
X __/
^ > 5 D eer/sq. mi.
Missouri
River
I - 5 D eey sq. mi.
<^1 Deei/sq. mi.
Not Surveyed
Figure 15.
Distribution of mule deer January 1975-helicopter survey.
-33IV in 1974 and 1975, respectively.
The areas of the highest and lowest
densities occurred the same year in the CK and Nichols Coulee drainages,
both of which are within pasture III.
In January 1975, CK Creek aver­
aged 4.9 deer/sq. mi. while Nichols Coulee averaged only 1.4 deer/sq.mi.
Numbers of fawns per 100 females, as an indicator of productivity,
increased 9.6 percent from 1974 to 1975.
All this increase was account­
ed for in pasture III, where the number of fawns per 100 females
increased 16.0 percent.
Pasture IV showed a marked decline of 23.7 per­
cent in the number of fawns per 100 females.
The fawn:doe ratio
obtained during the fall of 1974 from ground observations in both
pastures was 50.3.
Home Range, Movements, Use Patterns and Relationships
Data on home ranges and movements of three radio marked mule deer,
two males and one female, were obtained from 134 observations from
August 21 to December 21, 1974. . That part of the home range which
encompassed the areas of intensive use was considered as the normal
home range.
Total home range includes all points of deer observations.
including extended.short term or seasonal movements, as during the
rut.
Home Range Size
Normal and total home range were the largest for the two-year-old
male (Table 5), intermediate for the yearling male and smallest for
Table 5.
Summary of home ranges and movements during late summer and fall for three radio
marked mule deer.
Animals
Sex
Age
Times
Observed
/Days
Relocated
Dates of
first-last
observation
Areas (acres)
Normal Total
Home
Home
Range Range
Distances (Miles)
Ma x . beBetween Successive Relocations
Aug.tween anyMean Oct. Nov. Dec. Max Min two obs.
Male
2.5
58/53
8/21-12/18
1,863
4,955
1.16
1.03
1.57
1.08
3.5
0.0
5.4
Male
1.5
30/27
8/30-12/21
1,498
2,485
1.15
1.05
1.55
.96
2.1
0.5
2.8
Female 1.5
46/43
8/23-12/20
767
1,322
.74
.74
.94
.47
2.8
0.1
3.3
I
w
-O
I
-35the yearling female.
During the rut, total home range of the two-year-
old male increased 2.66 times and for the yearling male 1.66 times.
The total home range of the yearling female increased only slightly
during this period.
The effect of extended movements during the rut
on subsequent home range size and shape was not determined; but there
was some indication that the normal home range of the yearling male
expanded following the rut.
Larger home range and greater mobility of
males has been reported on by Robinette (1966) and Dasmann and Taber
(1956).
Movements
The longest and shortest over night movements recorded were made
by the two-year-old male and were 2.25 mi. and 3 yds., respectively.
Despite the fact all deer appeared to be able to cross their home
range in an over night period it was rarely done.
The mean distances
traveled between successive relocations (Table 5) were almost identical
for both males, except in December.
recorded for the yearling female.
Considerably smaller values were
However, when home range sizes are
considered, this represents a more intensive utilization of home range
by the female.
The effect of the rut during November is well illustrated by
increased distances traveled between successive relocations (Table 5).
Movements of the two-year-old male during this period are best
—36—
summarized as incessant travel.
At least five excursions beginning
from and ending in the normal home range were made between November 12
and December 11.
Durations from one to twelve days and distances up
to two miles from the closest boundary of the normal home range were
recorded for such excursions.
Infrequent relocations of the yearling
male make comparisons impractical.
No departures from the normal home
range were recorded for either male until the beginning of the rut.
Significant departures of the yearling female.were recorded on October
25 and November 20 when on. these dates this deer was observed in the
same small drainage 0.75 mi. and 0.25 m i . , respectively, from the
Missouri River.
The latter date was the only time the yearling female
was observed with other deer.
Activities of a yearling male in the
group suggested that the marked female was in estrus.
Dasmann and
Taber (19.56) reported that female Columbian black-tailed deer were
more active near or during the time of estrus and sometimes traveled
beyond home range.boundaries.
The small distances recorded between
successive relocations in December for the yearling female were associ­
ated with movements restricted to about 150 acres in the northeast
section of her,normal home range.
U se Patterns and Relationships
Use patterns were characterized by a few specific areas in the
. ■
normal home range receiving differential use (Fig. 16 and Table 6).
IN
l- S - y e e r - e ld mole
VS -y e w -e ld me|e
-yeer-eW femele
Tefel Heme tenge
Burned Aree
•eserwelr A Well
Missouri
Figure 16.
Bluer
Home ranges of three radio marked mule deer showing specific areas and burned
areas within their home ranges (see Table 6, and Text).
-38Table 6.
Differential use of areas of home range by three radio
marked mule deer.
Specific Area
% of reloca­
tions in an
area1
Area
No.
Area % of
normal
home
range
Burned Area
% of obs.
in or
% of normal
adjacent
home range
to burns
burned
Sex
Age
Male
2.5
I
2
•3
. 4
28.6
26.2
16.7
14.3
7.0
11.8
5.7
7.7
72.9
39.9
Male
1.5
I
2
3
4
16.7
16.7
16.7
16.7
5.0
4.9
1.0
3.2
40.0
15.1
Female 1.5
I
2
3
4
39.0
24.4
14.6
9.8
9.3
5.1
3.7
1.0
44.2
10.8
.
'
1Based on observations for yearling male.
Movements, except during the rut, were associated with travel between
these few areas.
Use of these areas was
to forage and cover.
believed to be in relation
One recognizable relationship of this was the
proximity of one or more of these areas, of heavy use to an area that
showed evidence of having been burned.
This resulted in a dispropor­
tionate percentage of observations being in or adjacent to burned
areas (Table 6).
No definite relationships to water were observed.
The normal home range of the two-year-old male was well interspersed
—39wit h sources of water.
Two reservoirs and intermittent pools of water
in CK Creek and four other minor drainages were the known sources of
water.
One well and one reservoir in Seven Mile Coulee were the only
known sources of water in the normal home range of the yearling male.
No sources of water were found within the normal home range of the
yearling female.
However, a well and a reservoir were located on
either side of her normal home range.
Use of area four may have been
related to its proximity to the well in Nichols Coulee.
If use of
either of these water sources occurred, it was at night.
The yearling male was the only radio marked deer whose normal
home range encompassed areas of cattle use.
Forty-three percent of
the yearling male's normal home range was off the study area in a
pasture which was grazed continuously by cattle from April I to Novem­
ber 30.
Thirty-six percent of the observations of this deer were in
that area.
Three observations were made when cattle were at distances
'■
■
■
'-
less than 1/8 mi., eight.when cattle were between 1/8 mi. and 1/4 mi.
and 15 other observations when cattle were at distances greater than
1/4 mi. and still present in the grazing systems.
As compared to the
two-year-old male, the yearling male made more intensive use of his
home range, possibly as a result of his home range having been grazed
by cattle during most of the period he was followed.
Normal h ome.ranges of all three deer were within the timbered
breaks.
The two-year-old male during the rut expanded his total home
-40range north and west, as well as south, to the limits of conifer growth
in the CK drainage system.
The yearling male during the rut made use
of range lands immediately north of the timber in Seven Mile Coulee.
This area comprised about 1/4 of his total home range..
Bedding was the activity most often observed.
juniper was the cover most often used for bedding.
Rocky Mountain
Most feeding and
travel took place during periods too dark for visual observations.
j
Group Characteristics
Mean group siize progressively increased from early summer to late
fall (Table 7).
Summer group sizes were consistently higher in 1973,
when 1.62 was the average group size, as compared to 1974 when the
average group size was 1.36.
Better forage conditions throughout the
summer of 1974 were believed to be responsible for this difference.
During summer and again during November and December, an obvious diff­
erence was recorded in average group sizes observed for males and
females (Table 7).
During summer and early fall, males were often
observed in groups of two to four.
Areas of mature male concentrations,
similar to that described by Mackie (1970), were observed.
In November
and. December, during the rut, males dispersed and 95.7. percent of the
observations of males were either as single individuals or with females.
The low average group size of females during June and July reflected
the intolerance of adult females, towards other deer during the fawning
-41Table 7.
Mean monthly and seasonal mule deer group sizes observed
during the study by sex and for all mule deer combined.
June
Males .
Females
Ave.
period.
July
1.33 .1.68
1.15 1.13
1.25 1.38
Aug.
1.58.
1.26
1.61
Month , ■
_____ '
.
. Season
Summer Fall
.Sept. Oct. Nov. Dec.
1.37
1.36
1.70
1.40
1.37
1.89
1.04
1.37
1.98
1.13
1.51
2,09
1.66
i.19
1.45
1.25
1.39
1.88
This was probably related to the high nutritional require­
ments of a female with a fawn during this period.
Dasmahn and Taber
(1956) reported mutual intolerance of adult females and spacing of
centers of activity by adult females when fawns were young..
...
Characteristics of the groups with which the three radio marked
deer associated changed often (Table 8) .
The association, of the
marked two-year-old male with the same recognizable mature male on 11
Successive relocations and 14 observations from September 21 to
October 10 was the only example of group consistency observed.
-42Table 8.
A summary of characteristics of groups associated with three
radio marked mule deer.
Animal
Sex
Age
Average
group size
Male
2.5
Male
1.5
Female 1.5
1.48
1.63
1.10
1YM
MM
D
D&F
= Yearling male's.
= Mature males.
= Females
=.Females and fawns.
% of obs.
deer occurred
alone
62.1
63.3
95.0
■ Associations
period*
Aug.^ Oct.
Noyi
. . YM 5= MM
Y M 5M M 5D 5 D&F
NONE
■ ■
■
.
.
Dec.
D 5 D&F, & MM
Y M 5M M 5D 5 D&F
Y M 5D1
■*
Activity
Over-all, feeding was the dominant activity observed during summer
(Table 9).
The.period of most intensive feeding observed during the .
study was in June and July when 31 percent and 34 percent, respectively,
of all.mule deer observed were feeding.
The decreased.percentages of
..
.
•
mule deer observed in feeding and travel and the increased percentages
in alert and running during the fall represented a trend which began
in September and peaked in November, and probably was associated with
increased wariness of the deer, due to disturbance by hunters. ,
Activity periods during summer lasted about three hours after sunrise
and began again about one hour before sunset.,
In fall the morning
activity period was shortened considerablyi approximating that
V /
.
;;
. ■■: .
■ ■■
, ■-
- I
Table 9.
Seasonal percentages of all activities of mule deer observed during the study
within each habitat type and total seasonal averages.
Habitat Type
Feeding
Activity - Summer/Fall
Bedding Alert Travel Running
Drinking
Artevisia-Agropyron
31/21
4/2
25/31
22/22
16/24
2/-
Pinus-Juniperus
27/20
8/16
31/26
25/17
9/21
T/-
Pseudotsuga-Juniperus
25/-
25/33
25/50
25/-
-/17
Saraobatus-Agropyron
34/39
-/5
21/22
29/6
16/28
Artemisia longifolia
23/49
12/6
19/15
31/15
15/15
Agropyron-Symphorioarpos
30/29
-/14
18/14
25/24
27/19
Xanthium strumarium
50/-
-/-
25/-
25/-
-/-
Average
29/22
5/12
26/18
13/21
T/-
-/26/27
-/-
—44—
bserved in the evening.
This resulted in a greater percentage of mule
deer activity occurring.in periods too dark to make observations.
Use of Habitat Types
Summer
The highest over-all use recorded during summer was on the Pinus
Juniperus habitat type (Table 10).
During the summer period, the
Artemisia-Agropyfon and Sarcobatus-Agropyron habitat types showed an
inverse relationship in mule deer use to the Pinus-Juniperus habitat
type.
During June, the former two habitat types received greater use
than for any other period during the study; apparently in relation to
the early maturation arid/or abundance of forbs of the Poa and Tragopogbn unions on' these types.
Increased use on the Pinus-Juniperus
type during the latter half of summer reflected later development and
desiccation of forbs and the relative abundance of skunkbrush on this
type.
Mule deer usage of all other types was relatively minor.
Use
of the Pseudotsuga-Juniperus habitat type was never great because of .
its limited distribution on the. study area; almost all recorded use on
this type was on the west side of Seven Mile Coulee.
Use of the
Artemisia longifpli'a habitat type in July was associated with the
variety, of forbs and summer browse plants on this type." Use by deer
of the Agropyrori-Symphoricarpos.habitat type was generally restricted
to side drainages. . The importance of this type to. mule deer, has .
■■
-
i;. - -
'.. v
Y ' . :■ ;■
' .. .■
. ■■■■'■ ■■■ :■
.
Table 10.
Monthly and seasonal percentages for all use of habitat type by mule deer
observed during the study for each habitat type and percentages of use by
mule deer for each community within a habitat type. Trace (T) amounts are
percentages less than I percent.
Habitat Type
Community
June
July
Aug.
Summer
Sept.
Oct.
Nov.
Dec.
Fall
Art&nisia-Agropyx-on
A . tridentata-A. spioation
A. tvidentata-A. smithii
A. tridentata-A. smithiiBouteloua gracilis
65
67
29
4
39
44
36
20
27
50
28
22
40
51
31
18
27
48
32
20
23
44
32
24
24
38
38
24
26
50
14
36
25
45
31
24
Pinus-Juniperus
Pinus-Agropyron
Pinus-Juniperus
Pinus-Artemisia
19
35
41
24
44
58
34
8
63
41
46
13
46
46
41
13
61
34
52
14
63
39
40
21
58
35
47
18
64
42
45
13
61
36
47
17
T
T
T
T
I
Sarcobatus-Agropyron
10
6
3
6
3
2
5
I
3
Artemisia longifolia
T
5
2
3
3
10
8
6
6
Agropyron-Symphoricarpos
4
4
4
4
5
2
3
3
4
T
T
Pseudotsuga-Juniperus
Xanthivan s tm m a riio n
I
—
2
—
I
T
—46—
probably been under-estimated due to.poor observability into bottoms
of side drainages.
times was minor.
Use of the Xanthivm stvvmavivm habitat type at all
The observed, preference" of mule deer for the. Avtemisia
tvidentata-Agvo’
p yvon spieatvm community over the Artemisia tridentataAgropyron smithii community was probably a result of the distribution
of the former coinciding more closely to those topographic areas and
cover types preferred by, mule deer.
A breakdown of observed deer
activity within each habitat type (Table 9). showed the importance of
the Artemisia-Agropyron and Sarcobatus-Agrdpyron habitat types to mule
deer as feeding areas during summer.
Fall
No dominant; trends were recorded in fall.
In general, trends in
the use of habitat types established in late summer continued through­
out fall (Table 10).
The only exception was the sharp increase in
amounts of use recorded for the Pinus-Artemisia and Artemisia lohgifolia
communities during October and November.
This was associated with
mule deer digging.out roots of first year stems of yellow.sweetclover
from unstable shale slopes, an activity which began concomitant with
the desiccation of this plant in these and other communities.
The
increased use of the Pinus-Juniperus community during August continued .
on through the fall period.
The increase in use of the Artemisia .
tridentata-Agrop’
y ron spieatvm and Pinus-Agrppyron communities during
•
-47December was related to utilization of rubber rabbitbrush (Chrysotharmus
naueosus) and the relative abundance of this plant on these communities.
The two fold increase in both total use and feeding usage observed on
the Artemisia longifolia habitat type indicated increased importance
to mule deer during fall.
Degree of usage of the Agropyron-Symphori-
carpos habitat type by mule deer for feeding and for all activities
while relatively minor for both summer and fall; were more consistent
between seasons than for any other habitat type.
As in summer, the
relatively minor use of the Sarcobatus-Agropyron habitat type for feed­
ing was on footslopes, and benches.
Use of Slope and Exposure
'
'
.
During summer and fall, morfe than one half of all mule deer
observed were on slopes greater than 10 degrees (Table 11) and slightly
less than one half of the observations occurred on northerly exposures
(Table 12).
This largely reflected heavy use by deer of the Pinus-
Juniperus habitat type.
A 17 percent increase in the percentage of
observations occurring on slopes greater than 10 degrees and a 62 per­
cent, increase in the percentage of observations noted for northerly
exposure from June to August corresponded closely with the increase
in use of the. Pinus-Juniperus habitat type during summer.
The apparent
greater .use of east exposures as compared to west exposures ^zas
■
1
..
■. ■ "■
■i
.
.
-48r
Table 11.
Seasonal percentages of all use of slope by mule deer
observed during the study within each habitat type and
total seasonal averages.
Slope in degrees-summer/fall
11-25
1-10
26-35 .36-45
Habitat type
Level
Artemisia-Agropyron
Pinus-Juniperus
Pseudotsuga-Juniperus
Saroobatus-Agropyron
Artemisia tongifolia
Agropyron-Symphorioarpos
Xanthium strumarium.
26/28 40/43
4/2
26/33
-/- . -/25/12 29/38
6/8
7/17
22/47 50/37
33/67/-
26/25
40/42
25/40
15/25
7/17
25/11
-/-
8/4
: -/25/18
4/5
50/60
.25/12/6
19/19
33/33
40/25
3/5 .
-/.-/-/- •
-/I/-/-/7/-/-/-
Average .
15/10
30/33
20/16
T1/-
30/36
5/5
45+
1 Less than I percent.
probably a result of a greater number.of mule deer observed in the
mornings when observations were predominantly to, the west,.-
, y.
'
'' [Eik/. ' . " ' T '
-
/
Results•are based on 145. elk groups totaling 588' elk observed dur­
ing the summers of 1973.and 1974; figures for the first.half of Septem­
ber 1973 atid fall 1974 were 203 and 775, respectively.
A total of 89
elk was counted during the two winter surveys.
Numbers, Productivity and Distribution
Data from the two winter surveys (Table 13) showed a substantially
greater number of ■elk, tlsing:-.the study area, in early .winter of 1975 as
compared to 1974.. Pasture III, the only, pasture where elk were counted
Table 12.
Seasonal percentages of all use of exposure by mule deer observed during
the study within each habitat type and total seasonal averages.
Habitat Type
Exposure - Summer/Fall
E
SW
SE
S
17/24 16/13
9/9
7/13
W
13/13
WW
8/9
ArterrrLsia-AgrtOpyrOK
N
11/8
NE
19/11
Pinus-Juniperus
19/19
25/24
15/10
11/12
11/10
5/6
6/4
8/15
Pseudotsuga-Juniperus
25/50
75/50
-/-
-/-
-/-
-/-
-/-
-/-
Saroobatus-Agropyron
5/8
14/17
28/17
19/25
22/33
3/-
6/-
3/-
Artemisia longifolia
8/6
4/-
4/7
25/25
33/56
9/-
17/-
-/6
Agropyron-Symphorioarpos
8/30
17/10
21/10
17/-
13/10
8/20
4/10
Xanthium strumarium
-/-
-/-
-/-
-/-
-/-
-/-
-/-
-/-
16/17
21/20
16/13
13/13
12/12
6/7
9/6
7/12
Average
12/-
-50in 1974, accounted for 97 percent of this increase.
To increase size
of sample and establish a more valid indication of elk productivity in
1974, data were also collected from the next two major drainages east
of the study area, Garden Coulee and Beauchamp Creek.
part of the 1974 combined totals,
These data were
Although reproduction was excellent
both years (Table 13), this factor could not account for all of the
increase in elk counted from 1974 to 1975.
Elk distribution, as determined from regular observations, was
wide spread within pasture III.during June and July 1973 and within
pastures III and IV. during September 1974.
During most other months
and seasons of the study elk distribution was confined to.a few .
specific areas (Fig. 17).
Of special significance was the concentra­
tion of elk during August and September 1973 in areas of Nichols
Coulee and CK Creek that had, received little or no previous grazing .by
cattle that year. . In June and July. 1974 elk appeared tc avoid those
areas of. the heaviest use by cattle.during the previous year.
Only
19 percent of the observations of, elk during this period occurred on
sections (sq* ml.) that accounted for 2.5 percent dr,more of the
total number of.cattle observed in each pasture during 1973.
This
area constituted about 31 percent of the area regularly surveyed during
1973.
Areas of. elk concentrations during the summer of. 1974 were not
'V .
areas of elk concentrations during the fall of 1974.
^ V
''
-51Table 13.
Summary of elk observations and classifications on and
adjacent to the Nichols Coulee RCA - helicopter survey,
early January 1974 and 1975.
Area
Total
Pasture III
1974
1975
Pasture IV
1974
1975
Bull
Cow
Calf
8
58
12
5
27
3
19
O
2
O
2
O
O
O
O
Garden CouleeBeauchamp Cr.
1974
1975
21
—
I
—
12
8
DID NOT
COUNT
Combined Totals
1974
1975
29
60
I
14
17
27
11
19
Bulls:
100 Cows
Calves:
100 Cows
Bulls
44
60
70
— —
— —
100.00
8
—
67
—
100.0
—
6
52
65
70
100.0
35.7
25.0
To make valid comparisons of elk use of the study area between the
summers of 1973 and 1974 only data from July and August, two months of
equal observations each year, were used.
Mule deer, whose numbers as
indicated from the winter surveys increased only slightly between
years, were used as a standard of observability.
While total numbers
of mule deer observed between summers remained the same (Table 14),
total numbers of elk observed in 1974 increase 1.66 times over that
observed during the same period in 1973.
In addition, the number of
-52Pamt IV
Pamt III
1973
Missouri Rivor
Past IV
P f i ** III
Legend
Ju n e A ju ly 1973
AUO ASEP 1973©
SUMMER
1974 O
SEP
1974 <2>
O CT- DEC 1974
Figure 17.
s areas w hich accounted for 5%
of the observations
Distribution of elk during specific peiiods of the study.
-53Table 14.
Ratios between July and August 1973 and July and August 1974
for numbers of mule deer and elk. observed within Nichols
Coulee RCA during these periods. . . .
Species compared .
Mule deer 1974/
.Mule deer
Elk 1974/
Elk
Elk 197.3/M u l e d e e r
Elk 1974/
Mule deer
Elk/Mule deer 1974/Elk/Mule deer
1973
1973
1973
1974
1973
Ratio
1.00
1.66
.45
.76
1.69*
V
I
elk observed per mule deer in 1974 was 1.69 times greater than in 1973.
All indications were that use by elk of pastures III and IV increased
from 1973 to 1974.
This increase in elk observed was probably related
to pasture III receiving the rest treatment in 1974 and cattle not
entering the timbered portion of pasture.IV until September.
"
Distribu-
tion of cattle within pasture IV was not complete until mid-September.
During 1973, elk were observed on only four occasions within
pasture IV (Table 15).
The percent of elk observations within pasture
IV increased 3.5 times from the summer of 1973 to the summer of 1974.
Despite this increase, over-all use of pasture IV by elk was low.
The
preference shown by elk to. pasture III over pasture IV: during the
summer of 19.74 could have been related, to previous treatments and.
amounts of use each pasture received in 1972 and 1973 (Table 24).
Both years, pasture III received less use by cattle per fated AUM and
received the use later in the grazing season than did pasture IV.
Table 15.
Percentage distribution by pastures, month, and season of all elk groups observed
in 1973 and 1974, together with the percentage distribution by pasture of five
marked elk when observed inside Nichols Coulee RCA. Number of groups of elk
appear in parenthesis. Data includes elk groups that were not classified.
Year______
Pasture
1973
Pasture III
Pasture IV
1974
Pasture III
Pasture IV
I
Pasture
1974
Pasture III
Pasture IV
Pasture
I
_________________________ Month____________________Season
Aug.
June
July
Sept.
Oct.
Nov.
Dec.
—
92(24)
8(2 )
100(21)
00 (- )
71(7)
29(2)
-
84(27) 84(27)
16(5) 16(5)
-
87(27)
13(4)
50(22)
50(22)
88(43)
12(6)
-
-
-
-
92(49)
6(3)
2(1)
-
-
Summer
96(45)
4(2 )
86(42) 85(81)
12(6) 15(14)
2(1)
Combined percentage distribution of five marked elk
82(179)
16(36)
2(4)
Fall
-
-
80(156)
19(36)
T (2)
Totals for all
elk
observed
82(23/)
17(50)
T(2)
-55During September of both years, there was a marked increase in the
percentage of elk observations that occurred within pasture IV as com­
pared to summer.
Fifty percent of the elk observations in 1974 were
in pasture IV as compared to 29 percent in 1973.
Probable causes of
this shift in use in pastures were the onset of the rut and later, the
occurrence of bow hunters on the study area from early September to
early October each year.
During both years of the study, elk were
observed in pasture IV in September before the opening of the hunting
season.
Though the percentage of observations of elk in pasture IV
during fall 1974 was greater than in summer 1974, percentages for all
months in fall except September were all below those for any summer
month (Table 15).
The increase in the percentage of observations in
pasture IV during December 1974 was a result of three observations of
the Channel (Ch.) 9 radio marked elk, two of which could be related
to disturbance by hunters, and three observations of a small group of
elk.on the west side of Seven Mile Coulee near the Missouri River.
Two of these observations were on the bottomlands of the river within
an area of about 160 acres that was fenced to exclude cattle that year.
The observed differences of elk distribution and numbers between
years, between pastures within a,year, and within a pasture within a
year, for the most part, was related to quantity acd quality of forage
available.
Grazing by cattle appeared to be a major influence bn
these factors
-56Home Range, Movements, Use Patterns and Relationships
Home ranges and movements of four radio marked elk, two males and
two females, and two neck-banded elk, one male and one female, were
obtained from 234 observations made from January 18, 1974 to December
26, 1974.
.
Home Range Size
Home range was the largest for 'the two radio marked males and
smallest for the Ch. 7 female (Table 16) who was accompanied by a calf
during summer and fall.
The C h . 10 female, who did not have a calf,
had a home range comparable in size to the two radio marked males.
Data were too limited to make any conclusive statements concerning the
•
•
'
. . .
expansion of home range by male elk during the rut as was characteris­
tic of mule deer.
Movements
Elk appeared to be extremely mobile within their home range.
Oyer-all, movements appeared to increase slightly from summer to fall
(Table 17).
Most of the increase occurred during the September and
October period.
The.C h . 10 female was the only elk with sufficient
relocations to show a decline from summer to fall.
Average distances
traveled between successive relocations were similar for both radio
marked females but they showed opposite trends within fail.
Movements
recorded for the neck-banded male during summer were similar to those
Table 16.
Home range size and use of home range by locality for five of the sex marked
elk in Nichols Coulee RCA. Radio channel number appears in parenthesis.
Animal
Sex
Age
Type1S1
2
of tag
Date of
first-last
observations
Area of
home
N o . of
range
obs./
reloc. (sq. mi .)
% of home range/observations
within a given area
Past. IV Past. ITI Past. I
Study
Area
10.5
R4 (H)
8/20-11/26
18/14
42.9
19/6
63/72
-
82/78
Male
2.5
R ( 9)
8/20-12/26
52/47
42.6
37/23
62/73
T/4
100/100
Male
2.5
1/18-12/16
26/23
25.9
8/28
73/57
T/8
82/93
Female
8-10
R 5 ( 7)
1/23-12/26
46/43
20.7
4/11
96/89
-
Female
4— 6
R (10)
1/23-12/26
86/75
38.0
37/13
56/85
-
Female
4— 6
Male3
NB
NB
1/18-12/4
6/6
—
—
—
—
1Elk radio tracked from ground during fall only.
2R = radio; NB = neckband only.
3Killed by a hunter 12/8/74.
ttRadio malfunctioned early November.
5Radio malfunctioned 1/24-9/11 located as neck-banded only elk between these dates.
100/100 wj
93/98
—
Table 17.
Animal
Sex
Male
Average distances recorded between successive relocations of six marked elk using
Nichols Coulee RCA for 1974, for season and two periods in fall, together with
the maximum distance between any two observations. Number of relocations are
in parenthesis.
Age
10.5
Tag1
type
R 11
Average distances recorded between successive relocations in miles
Winter
Max. length
Spring
Summer
Fall
Sept.-Oct. Nov.-Dec. of home range
Mean
6.6(2)z 5.4(11)
5.5(10)
5.3(13)
4.5(1)
15.0
3.7(2)1
2
2.5(44)
3.0(25)
2.0(19)
5.5(2)
1.4(7)
1.7(13)
1.9(4)
1.7(9)
1.8(42)
1.9(3)
1.4(9)
1.9(30)
1.7(16)
2.1(14)
6.8
R 10
2.3(74)
4.1(11)
2.0(19)
1.9(44)
2.2(22)
1.6(22)
9.8
NB
3.2(5)
4.9(2)
-
2.2(3)
2.5(2)
1.5(1)
8.0
Av e . for males
2.8(81)
5.5(2)
2.8(11)
2.9(68)
3.5(39)
2.0(29)
11.8
Av e . for females
2.2(121)
4.0(16)
1.8(28)
1.9(77)
2.0(40)
1.8(37)
8.2
Total averages
2.6(202)
4.2(18)
2.1(39)
2.3(145)
2.8(79)
1.9(66)
10.0
Male
2.5
R 9
2.5(46)
Male
2.5
NB
1.9(22)
Female
8-10 R 7
Female
4-6
Female
4—6
-
1R = Radio and Channel Number; NB = Neck Band Only.
2Late summer movements.
9.3
11.0
recorded for the Ch. 7 female.
During fall, the average distances
traveled between successive relocation were larger for males than
females, especially during September and October.
The high values
recorded for the Ch. 11 male during September and October may be indica­
tive of the role older males play during the rut.
On seven of the eight
instances of harassment by hunters, radio marked elk moved to another
.part of their home range.
Average distance traveled from the point of
original disturbance to the next relocation was 6 . mi. for the two
males and 2.7 mi. for the two females.
Use Patterns and Relationships
Differential use of home range, except by males during the rut,
was noted for the matked elk during summer and fall.
relation to forage and cover.
This was in
Five of six marked elk centered their
activity within pasture III (Table 16).
Two general trends were noted;
either the percentage of observations in pasture III was greater than
percent of home range in that pasture or if the reverse, that percent
of the home range outside pasture III was a minor part of the total
home range.
The former was the case for the three elk with larger
home ranges and the latter,was the case for the t#o with the smaller
home ranges.
Evidences of shifting patterns of home range use were seen for
each radio marked elk during December.
The. Ch. 11 male who was last
-60observed in CK Creek on November 26, was killed by a hunter on December
8, seventeen miles west of the point last observed.
moved to the west side of Seven Mile Coulee twice.
such movement was a result of hunter disturbance.
The C h .9 male
However,the.first
The C h . 7 female
expanded her home range north in CK Creek, close to
the area where she
was observed most often in July.
moved to the south
boundary of her home range.
The Ch. 10 female
During the periods covered, the most
southerly and easterly points of observations of the C h , 10 female
were in association with the Ch. 7 female.
From November I to December 8, the C h ..9 male was observed 17
consecutive times within an area of 6 sq. mi. in the upper part of
pasture III and.a small portion of pasture I (Fig. 18).
General elk
observations were also concentrated in this area during this period
(Fig. 17).
Abundant amounts of first year stems of yellow sweetclover
and in addition, in December, sources of open water were the two major
factors responsible for this concentration of elk.
From August 8 until
December 9, twenty-mine of 31 observations of the C h . 7 female were
within 23% mi. of the Missouri River (Fig. 19).
The C h . 10 female was
observed out of pasture III on only 13 occasions:six times in winter
and spring when she tended to wander about her home range, one time
in June, two times at the end of August and four times in fall as a
result of hunter disturbance.
By the end of June, home range size and
shape changed little for the Ch. 10 female (Fig. 20).
W-'
The neck-banded
Fest I
Fdst IV
*"
I
O
\
\
/
I —
- J
\ »
,Im ile ,
<
X
tL
X 1x?
«1V\k «» wI *22
37^
38° ,, S 3
a
U
f
29"
a
on
V
Past III
Legend
Observe*:#
AUG 1A2
S IF 3 - 1 7
OCT 18-30
NO V 31-40
DEC 41-52
Home Range _
Figure 18.
Missouri River
Home range and movements of the radio marked 2.5-year-old male elk.
AUG
9-12
DEC. 3 5 -3 6
SEP. 13 - 19
Home R a n g e __________ ______
Figure 19.
Home range and movements of the radio marked 8-10-year-old female elk.
Post I
HO
du
» f O 60
V o o Se
*i
.d„
m o
2 Jf
Past. Ill
Missouri River
J A N -M A Y 1-11
JUNE 12-17
OCT 45—59'
JULY 18-26
NOV 60-71
AUG 2 7-32
DEC 7 2-86
SEP 3 3 -4 4
Horae R a n g e -------------------------- --
Figure 20.
Home range and movements of the radio marked 4-6-year-old female elk.
—64—
male apparently wintered off the study area in the Garden Coulee area
(Fig. 21).
The summer range of this elk was shifted to an area of 3.8
sq. ml. about equally divided between pasture III and IV (Fig. 21).
Seven of ten observations from June 17 to September 8 were within
pasture IV.
The observation of this elk in pasture IV on September 8
was the first time cattle were recorded within his home range; in this
case they were less than 1/8 mi. away.
The next observation of this
elk was on September 24 when he was observed just northeast of his
summer home .range.
From September 24 to December 16, all 14 observa­
tions of this elk were within an area of 7.7 sq. ml. in the upper part
of pasture III and a small portion of pasture I.
The neck-banded
female apparently wintered .in the same area as the neck-banded male.
This elk was not observed on the study area until October 21* one day
after, the opening of the deer hunting season.
Three more observations
of this elk up to December 4 indicated she had shifted her use of home
range to parts of pasture III..
Group Characteristics
Group sizes increased progressively through summer,, dropped sharply
in September and remained low until December (Table 18).
size recorded was.22 in late August 1973.
observed in summer.
groups.
Maximum group,
Mature males were rarely
From late November on, males were banded in small
Average group sizes during July and August, were larger in 1974
Post. I
Legend
Observations
J A N -M A Y 1 * 2
JUNE 3
OCT 14
JULY 4 - 6
NO V 15-21
AUC 7 -1 0
DEC 2 2 - 2 6
SEF 11-13
Home R a n g e _______ __________
Figure 21.
Home range and movements of the neck-banded 2.5-year-old male elk.
-66Table 18.
Males
Females
Ave.
Mean monthly and seasonal elk .group sizes observed during
the study by sex and for all elk combined.
June
July
Aug.
Month
____
Sept. Oct. Nov.
1.40
2.33
2.50
1.59
3.09
4.43
1.87
3.11
4.70
1.04
2.44
3.44
1.32
2.13
3.61
1.96
2.58
3.48
Dec.
2.60
3.12
4.86
Season
Summer Fall
1.68
. 2.92
4.06
1.70
2.55
3.82
when 4.9 elk per group were recorded as compared to 4.3 elk per group
recorded in 1973. . Observations of marked elk showed group character­
istics constantly changed.
Knight (1970) reported similar results.
All coefficients of associations (Cole, 1949) for marked elk were low
(Table 19).
Activity ■
'
•
'
.
Feeding was the dominant activity observed both summer and fall
(Table 20).
Morning activity periods ended about one hour after sun­
rise and began again about one hour before sunset in the evening for
both summer and fall.
The increase in bedding during fall was largely
due to observations of radio marked elk during this period.
in feeding were observed within or between seasons.
.
No trends
December was the
.
month of most intensive feeding when 33 percent of the elk observed
were feeding.
Table 19.
Coefficients of association for six marked elk using Nichols
Coulee RCA during 1974.
Animal
Sex
Associations
Tag or
Animal
Age- .
Ch.
Sex
Male
2.5
Male
2.5
Female
8-10 •
‘
Age
Tag or
Ch.
Coefficient of
association
9
Male
Female
Female
2.5
8-10
4— 6
NB
7
10
•24
.07
.02
NB
Male
Female
Male
10
4-6
2.5
11
NB
9
,06
.06
.24
7
Female
Female
Male
4—6
4— 6
2.5
10
NB
9
.11
. . -05
. .07
' ..’
Use of Habitat Type
V- -' '
^
Summer
'
The highest over-all use recorded during summer was on the PinusJuniperus habitat type (Table 21).
During summer, the Artemisia-
Agropyron habitat type showed an inverse relationship in use by elk to
the. Pinus-Jun-iperus habitat type.
This trend was related to .the later
development and desiccation.of forbs on the Pinus-Juniperus habitat
type.
Use of all other types was relatively minor.
When used,
especially in June, the Sarcobatus-Agropyron habitat type was an
important area for feeding (Table 20).
Use of this type in early
summer was generally on footslopes and benches and was related to
Table 20.
Seasonal percentages of all activities of elk observed during the study within
each habitat type and total seasonal averages.
Habitat Type
Feeding
Activity - Summer/Fall
Bedding Alert Travel Running
Drinking
Artemisia-Agropyron
32/33
1/4
12/26
39/10
13/27
3/-
Pinus-Juniperus
22/24
13/23
32/28
18/14
15/11
-/-
Pseudotsuga-Juniperus
-/-
-/67
-/33
-/-
-/-
Saroobatus-Agropyron
42/28
-/-
8/18
17/27
33/27
-/-
Artemisia longifolia
29/50
14/3
14/31
29/10
14/6
-/-
Agropyron-Symphoricarpos
45/28
11/9
22/27
22/18
-/-
-/-
100/100
-/-
23/28
27/13
Xanthium strumarium
Average
28/29
7/15
-/18
-/-/-
14/15
T/-
Table 21.
Monthly and seasonal percentages for all use of habitat type by elk observed
during the study for each habitat type and percentages of use by elk for each
community within a habitat type. Trace (T) amounts are percentages less than
I percent.
Habitat Type
Community
June
July
Aug.
Summer
Sept.
Oct.
Nov.
Dec.
Fall
Artemisia-Agropyron
A . tridentata-A. spioatum
A. tridentata-A. smithii
A. tridentata-A smithiiBouteloua gracilis
50
40
50
10
42
67
33
-
25
13
37
50
39
47
39
14
20
33
56
11
22
60
30
10
43
35
39
26
17
50
25
25
26
43
36
21
Pinus-Juniperus
Pinus-Agropyron
Pinus-Juniperus
Pinus-Artemisia
38
42
47
11
47
51
42
7
59
31
56
13
48
41
49
10
53
7
72
21
52
12
70
18
51
31
60
9
57
18
65
17
53
17
67
16
Pseudotsuga-Juniperus
-
-
-
-
3
-
-
-
T
Saroobatus-Agropyron
8
4
5
6
10
11
10
8
Artemisia longifolia
2
3
4
3
11
8
6
8
8
Agropyron-Symphorioarpos
2
4
5
4
3
6
-
6
4
Xanthium strumarium
—
—
2
T
—
I
—
2
T
-
—70ear Iy development of forbs.
Use of the Agropyron-Symphoricarpos
habitat type by elk increased through summer.
The importance of this
type to elk was underestimated by observational bias.
Where it occur­
red in small side drainages next to the Pinus-Juniperus community, elk
made use of the Agropyron-Symphoricarpos habitat type for feeding dur­
ing periods between, as well as within, the morning and evening
activity periods.
Use of this type in major coulee bottoms was during
twilight hours and at night.
Within the Artemisia-Agropyroh habitat
type the Artemisia tridentata-Agropyron spiaatum community appeared to
be preferred for the same reason as described for mule deer;
The
Pinus-Juniperus community received the most use within the Pinus
habitat type and was extremely important for bedding cover.
Fall
The Pinus-Juniperus and Artemisia-Agropyron habitat types again
rated first and second in over-all importance (Table 21) and their use
for feeding changed little from summer (Table 20).
An abrupt increase
in the use of the Artemisia-Agropyron habitat type in November was
related to use of Junegrass (Koeleria oristata) and dried first year
stems of yellow sweetclover, both.of which figured prominently in
feeding site examinations during this period.
Late August and early
September rains in 1973 resulted in.regrowth of Sandberg bluegrass
(Poa seounda) and western wheatgrass.
Use by elk of the Sarcobatus-
-71Agropyron habitat type in September 1973 was related to these two
plants and Junegrass.
In 1974, use of this habitat type; was related to
use of yellow sweetclover.
A sharp increase in use was recorded during
September and October on the Pinus-Artemisia and AvterrrLsia IongifoLia
communities, where yellow sweetclover, longleaf sagebrush and in
December soapweed were sought out by elk.
During September and October
elk made intensive use of the cover provided by the Pinus-Juniperus
community.
Also, in September, the dnly use of the Pseudotsuga-
Juniperus habitat type was recorded.
Use of Slope and Exposure
■
.
■■ :
■' '
■ .
'
-
.
An increased use by elk of steeper slopes was recorded from summer
to fall (Table 22).
This was largely a result of increased use of the
Pinus-Juniperus,. Pinus-Artemisia and Artemisia LongifoLia communities
during fall.
Use of exposure (Table 23) tended to favor those slopes
noted for a northerly aspect, where the Pinus-Juniperus habitat type
was used most intensively.
-72Table 22.
Seasonal percentages of all use of slope by elk observed
during the study within each habitat type and.total
seasonal averages.
Habitat type
Level
Slope in degrees-summer/fall
1-10
11-25
26-35 ..
36t 45 . 45+
36/20
Artemisia-Agropypon
5/1
Finus-Juntpepus
Fseudotsuga-Junipepus
-/45/48
Sapoobatus-Agpopypon
11/3
Artemisia longifolia .
Agpopypon-Symphopioappos 50/55
Xanthium stpumgpium
100/100
41/38
29/28
-/55/19
IlA
50/45
-/-
20/34
46/39
-/67
-/29
33/12
-/-/-
Average
35/27
32/33 . 12/20
21/11
3/8 .
-/■-/-.
-/6
20/26
-/33
-/-/-/4
-/34/35
' 11/41 -/9
-/-/-/- •
-A, -/... I/8
-/I
1Less than I percent..
.
■
Cattle '
Results are based on 641 groups of cattle totaling 4,240 individ­
uals observed during the summers of 1973 and 1974:
figures for the
first half of September 1973 and fall 1974 were 598 and 3,251*
respectively.
...
.
-
Numbers and Distribution
During the two years of the study an average of about 1,700 herefprd cattle were grazed in Nichols Coulee RCA during the period April I
to November 30.
The exact number and dates varied each'year.
Animal
Unit Months used (Table 24) in Nichols Coulee RCA and within each
Table 23.
Seasonal percentages of all use of exposure by elk observed during the study
within each habitat type and total seasonal averages.
Habitat Type
Artemisia-Agropyrcm
N
22/9
NE
11/15
Pinus-Juniperus
22/19
21/24
Pseudotsuga-Juniperus
-/-
Saroobatus-Agropyron
-/100
17/8
Exposure - Summer/Fall
E
SE
S
SW
27/23 11/10 9/13
7/13
NW
9/18/14
11/12
6/6
9/7
5/8
8/10
-/-
-/-
-/-
-/-
-/-
-/17
50/25
-/42
-/8
33/-
60/10
-/9
Artemisia Zongifolia
-/-
Agropyron-Symphorioarpos
-I-
-/-
50/-
25/-
Xanthium strumarium
-/-
-/-
-/-
-/-
20/14
16/19
17/13
10/10
Average
W
4/17
20/5
-/28 20/43
-/50
-/25
-/25
-/-
-/-
-/-
9/14
7/9
7/11
-/-
-/5
25/-/14/10
—74Table 24.
Summary of the grazing seasons from 1972 to 1974 for each
pasture and totals for Nichols Coulee RCA.
Year
Past. I
Past. II
Past. Ill
Past. IV
Total
1972
Treatment
Dates used
AUMs used
AUMs per
rated AUM
D
rest
rest
rest
C
8/11-12/5
2,555
.74
B
5/17-11/151
2,785
.93
A
4/1-11/15
6,479
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
2.20
D
rest
rest
rest
C
7/20-11/30
3,357
1.12
B
5/28-11/5
3,942
1.50
4/1-11/30
13,931
1.13
1974
Treatment
Dates used
AUMs used
AUMs per
rated AUM
B
6/2-11/10
3,054
1.01
A
4/1-10/24
5,841
1.70
D
rest
rest
rest
C
7/31-11/25
3,326
1.27
4/1-11/25
12,221
.99
1Gates left open between pastures III and IV after 8/11.
individual pasture varied between years and between pastures within
years.
Of particular importance was the heavy use of pasture IV in
1973 and 1974.
In 1972, when pasture IV received the "A" treatment,
cattle were present from April I to November 15.
Cattle distribution within a pasture was greatly influenced by
habitat types, forage conditions and sources of water.
Bottoms of major
coulees and tops of level ridges generally were areas of concentrations
of cattle (Fig. 22).
In August and September 1973, cattle were con-
-75-
Figure 22.
Generalized distribution of cattle density for each
year of the study.
centrated on the bottomlands of the Missouri River.' .In pastures III
and IV during this period up to 16 percent and 63 percent, respectively
of the cattle present within each pasture were within one mile of the •
Missouri River.
In pasture IV, the concentration, was so pronounced
that during this period the upper one-fourth of the pasture received
relatively little use by cattle.
Bottomlands were flooded in 1974
and cattle were more evenly distributed within pasture IV (Fig. 22).
All areas accounting for 2.5 percent or more of the total number of
cattle observed within each pasture were within one mile of a source
of water.
However; not all areas within one mile of a source of water
were.areas of heavy use.
Distribution of cattle was modified by
fences and herding in relation to pasture treatments.
Time required
for complete distribution of cattle within a pasture depended on the
number and location of entry points, herding and possibly forage,
conditions. ’ Distribution appeared to be less rapid in 1974 when
forage conditions were better than;in 1973.
Generally,; it appeared
that 30.to 45 days were needed for cattle to become completely distri­
buted throughout, a pasture following entrance. '
Movements
Movements of .males were determined from 15 observations of four
recognizable individuals over a period of 10 to 53 day s . . Two of these
-77traveled as a pair In pasture IV during July 1974 and data for these
were analyzed collectively.
All recognizable cattle appeared to be fairly mobile (Table 25).
Activities appeared to be centered around sources of water with neriodic
movements between sources of water.
Movement data more closely reflect
these periodic movements than movements associated with daily activities.
Table 25.
Summary of movements for four recognizable cattle.
,AnimflL_
Age1
Sex
Male
IM
Males1
2
Male
IM
M
Dates of
first-last
observations
Breed
Hereford7/25/73-9/9/73
• Angus Steer
Hereford
7/13/74-7/22/74
Hereford
7/22/73-9/12/73
No. of
reloca­
tions
Distances in miles
Ave.
between
Max.
reloca.
7
. 2.00
6.5
4
4
1.91
2.25
3.8
4.5
2.04
4.9
Average
1 IM = Immature; M = m a t u r e . '
'
'
2 Two - traveled as a pair, data analyzed collectively.
Group Characteristics
.
_
'
•
.
Data for June were combined with those for July because cattle .
observations were not begun until late June 1973 in pasture IV.
Aver­
age group size was largest in June and July and declined progressively
to November (Table 26).
During mid-day, groups of cattle up to 75 in
—78“
Table 26.
Mean monthly cattle groups sizes observed during the study
by pasture and year and mean monthly and seasonal group
' sizes for all cattle combined.
Pasture III 1973
Pasture IV 1973
Pasture IV 1974
A v e . group size
June &
Julv
8.4
7.2
8.0
Aug.
6;8
6.3
6.1
6.4
Sent.
6.4
6.3
5.2
Oct.
Nov.
5.1
5.0
5.8
5.1
5.0
Summer
6.6
Fall
5.4
number, but more commonly 15 to 30, were observed around reservoirs and
wells.
During morning and evening activity periods, when most, of the
observations of cattle were made, cattle were dispersed, and. average
group sizes were considerably smaller.
Average group size within a
pasture was largest during the month-of entrance into the pasture.
The
higher average group sizes recorded in 1973 as compared to 1974 were
probably a result of the concentration of cattle along the Missouri
River in 1973.
'
-
.
.
Activity ■
Feeding, was the dominant activity observed both summer and fall
(Table 27).
Cattle were observed to feed throughout the day but feed-
Ing was most intensive during a three'to four hour period after sunrise'
and before "sunset.
Differences in percentage of cdttle observed feed­
ing were, recorded between pastures III and IV during .1973; these figures
Table 27.
Seasonal percentages of all activities of cattle observed during the study within
each habitat type and total seasonal averages.
Habitat Type
Feeding
Activity - Summer/Fall
Bedding Alert Travel Running
T/T
Drinking
Artemisia-Agropyron
59/64
21/18
8/6
12/11
Pinus-Juniperus
54/81
27/9
8/10
11/-
Pseudotsuga-Juniperus
-/-
-/-
-/-
-/-
-/-
Saroobatus-Agropyron
68/76
14/9
2/2
13/12
-/I
Artemisia longifolia
50/82
-/-
—/18
50/-
Agropyron-Symphorioarpos
74/74
20/14
-/2
6/8
-/I
-/I
Xanthium stmanarium
73/89
18/6
2/-
2/2
2/1
3/2
Average
63/73
19/13
5/4
11/9
T/T
1/T
T/-
3/-
.—80—
were 57 percent and 78 percent, respectively.
Each year and within each
pasture there was.an increase in feeding a s .time progressed.
Months of
the most intensive feeding were September 1973 and November, 1974 when
76 percent and 80 percent, respectively, of all castle observed were
feeding.
The highest values for travel recorded within a pasture Occur­
red both years during the initial month of occupancy.
Use of Habitat Type
Summer
The highest over-rail use recorded during summer was on the
Artemisia-Agropyroh habitat type,(Table 28).
However, in relation to
area, the most intensive use recorded was on the Sarcobatus-Agropyroh
habitat type.
A possible exception to this was in August and September
1973 when 21 percent and 47 percent,.respectively, of all cattle observ­
ed within pasture IV were on the Xanthiym strimaviim habitat type along
bottomlands of the Missouri River.
At other periods and other locations
use of this latter type was minor.
The Agropyron-Symphoricarpos
habitat type was of moderate importance because of its importance as an
.area for feeding (Table 27);
The Pinus-Juniperus habitat type appeared
to be used moderately for bedding in addition to feeding, during summer.
The Artemisia trideritata-Agropyron spioatum community received, the
least use. within the Artemisia-Agropyron habitat type.
This was related
to cattle using areas where this community was only poorly developed.
Table 28.
Monthly and seasonal percentages for all use of habitat type by cattle
observed during the study for each habitat type and percentages of use
by cattle for each community within a habitat type. Trace (T) amounts
are percentages less than I percent.
Habitat Type
Community
June &
Aug.
July
Summer
Sept.
Oct.
Nov.
Fall
Artemisia-Agropyron
A. Tridentata-A. spioatum
A. tridentata-A. smithii
A. tridentata-A. smithiiBouteloua gracilis
59
10
48
42
49
24
43
33
53
22
44
34
27
23
55
22
56
14
31
55
42
13
47
40
43
18
43
39
Pinus-Juniperus
Pinus-Agropyron
Pinus-Juniperus
Pinus-Artemisia
10
57
43
-
8
78
20
2
9
75
23
2
7
48
38
14
10
57
30
13
15
53
12
35
9
52
28
20
-
-
-
-
-
-
-
Sarcobatus-Agrcpyron
22
24
23
32
17
27
25
Artemisia longifolia
I
T
T
I
3
12
3
Agropyron-Symphoricarpos
3
6
5
16
12
4
12
Xanthium stnonarium
5
13
10
17
2
_
8
Pseudotsuga-Juniperus
-82Principal use of the Pinus-Juniper.us habitat type was in the Pirius-,
Agropyron community. ■ Use of habitat types and areas by cattle after
entering a pasture generally followed a pattern.
Mainridgetops and
major coulee bottoms were first used; later, use of level tops of side
ridges and bottoms of side coulees gradually increased;" finally,
cattle were observed on level portions of sides o f .ridges and steeper
slopes.
Fall
The Artemisia-Agropyron and Sarcobatus-Agropyron habitat types
were again first and second in order of usage (Table 28).
In September
of both years, use of the Artemisia-Agropyron habitat type declined
considerably as compared to summer. . This was associated with cattle
predominantly being in the bottoms of coulees and along the bottomlands
of the Missouri River when an increase in use of. the three habitat
types characteristic of these areas was noted.
trend was observed.
In October, a reverse
These trends were probably related to the early
desiccation of grasses on xeric upland areap and later, the exhaustion
of forage supplies oh the more restricted, niesic lowland areas.
Use .
of the Pinus-Juniperus habitat type increased progressively through the
fall period.
The importance of this type for feeding in fall increased
greatly as compared to summer (Table 27).
The Pinus-Artemisia,
Artemisia longifblia and Sarcobatus-Agropyron communities, character- ;
—83—
istically found on steeper sides of ridges and footslopes, showed a
marked increase.in use by cattle during November.
Use of Slope.and Exposure
During both summer and fall over three-fourths of all cattle
observed were on slopes of less than eleven degrees (Table 29).
Very
few cattle were observed on slopes of greater than 25 degrees.
During
fall cattle appeared to use somewhat steeper slopes.
Especially, in
November, when 39 percent of the cattle observed were oh slopes greater
than ten.degrees. . Use of exposure during summer.was fairly evenly
distributed among the various exposures (Table 30). • In fall a slight
decrease in use of slopes noted for. a northerly aspect was recorded.
Table 29.
Seasonal percentages of use of slope by cattle observed
during the study within each habitat, type and total seasonal
averages.
Habitat type
Level
/Irtemisia-Agropyron 42/43
.15/2
Pinus-Juniperus '
Pseudotsuga-Juniperus- -/Harcobatus-Agropuron 65/67
A rLarnisid longifo Iia. -/5.
68/83
Agropyron. Symphoriaarpps
Xanthium strurnarivm 99/94
Average.
52/46
'
■ ■'
1-10
36/43
49/39
-/20/21
-/13 ;
27/14
11-25
16/17
25/39
-/12/8
-/23
5/3
26-35
36-45
6/7
-/10/16
1/4
-I- . ' rZ- .
3/4
-Z t
.100/32.
-/27
-/-/-
1/6
-/-
-/-
-/•“ •
29/30
14/15
5/7
T/2
45+
-/-/•-/-/-/-/•
-/-/-
Table 30.
Seasonal percentages of all use of exposure by cattle observed during the study
within each habitat and total seasonal averages.
Habitat Type
Arterrrisia-Agropyron
N
11/8
Pinus-Juniperus
21/15
NE
12/12
Exposure - Summer/Fall
S
E
SE
16/13
12/18
13/19
8/15
4/12
SW
13/10
W
11/10
NW
12/10
13/9
14/10
13/14
10/16
17/9
Pseudo tsuga-Juniperus
-/-
-/-
-/-
-/-
-/-
-/-
-/-
-/-
Saraobatus-Agropyron
6/3
9/2
16/25
17/16
15/11
15/17
15/23
7/3
Artemisis longifolia
-/-
-/15
-H
50/11
50/33
-/22
-/8
-/4
Agropyron-Symphoriaarpos
6/7
V-
27/-
13/20
13/27
7/40
20/6
7/-
Xanthium strumarium
-/-
-/33
50/33
-/-
-/-
-/33
50/-
—/-
11/8
11/11
12/16
13/16
16/14
13/14
Average
12/13
12/8
-85Food Habits
Mule Deer
Results are based on examinations of 34 mule deer feeding sites
totaling 5,616 instances of use during the summers of 1973 and 1974:
figures for the first half of September 1973 and fall 1974 were 23 and
5,561, respectively.
Twenty-five mule deer rumen samples collected
in October and November 1974 were also analyzed.
Summer
. Forbs accounted for about two-thirds of the total instances of use
recorded during summer (Table 31).
Use of grass was miixor.
Yellow
sweetclover and skunkbrush were the most important forb and shrub,
respectively.
feeding sites.
Together they accounted for 76 percent of the use at.
In 1973, yellow sweetclover accounted for 43 percent
of the instances of use while skunkbrush accounted for 34. percent;
figures for 1974 were 59 percent and 17 percent, respectively.
This
difference was related to the greater abundance of first year stems of
yellow sweetclover in 1974.
In the summer of 1973, after the desic-
,
cation of yellow sweetclover in late July, use was shifted to skunkbrush .
This suggested that, use of skunkbrush during summer by mule
*
deer was inversely related to use of yellow sweetclover and that mule
.
■
deer preferred forbs when succulent.
■■■
._
:* ■■
■
'
:
Mackie (1970) also found similar
Table 31.
Seasonal frequencies of occurrence (0), aggregate mean percentages of use (U) and percent canopy coverage (C) by habitat type and by all types
combined for each plant species used by mule deer which accounted for I percent cr more of the use recorded at feeding sites in one or more
habitat types. Trace (T) amounts are less than I percent. Numbers of feeding sites examined are in parenthesis.
Sc&ZCbCLZUSFORAGE CLASS
PLANT SPECIES
USED2
GRASS It SEDGES
AGCR
Total Grass
FORBS
ASCO
COUM
ERMU
GLLE
LASE
MEAL
MEOF
MINU
PECA
PEPU
PHLI
POER
TRDU
YUGL
Total Forbs
BROWSE
CHNA
CHVI
JUSC
PRVT
RHTR
RICE
RONU
SYOC
Total Browse
1
All Types Combined
Summer
Fall
Arterisia-Asropuron
Pinus-Juniz>er-*s
Summer
Summer
Fall
(34)
(23)
O U C
O U C
4/ T/ I
3/ I/ I
29/ T/43 17/ 1/39
(9)
O U C
(4)
O U C
(13)
O U C
(14)
O U C
(7)
O U C
22/ T/47
25/ 1/42
28/ T/50
43/ T/40
43/ T/43
6/ T/ T
27/ 2/ I
3/ T/ T
3/ I/ T
15/ I/ T
6/ I/ T
68/54/11
3/ T/ T
9/ T/ I
3/ 2/ T
3/ T/ T
3/ T/ T
35/ 4/ T
88/68/21
3/ T/ T
6/ T/
56/22/
6/ I/
21/ 6/
26/ 2/
65/32/
T
I
T
I
3
9
18/
9/
9/
9/
I/
T/
T/
T/
T
T
T
T
63/36/ 9
Fall
25/ 5/ T
22/ 6/ 3
14/ 2/ I
25/ 2/ T
11/ T/ T
11/ I/ T
89/72/10
21/
7/
7/
7/
Summer
T/
T
T/
I/
I
T
T
T
7/ T/ T
50/28/ 3
50/34/ 6
57/33/ 7
11/ I/ I
18/ T/ I
5/ T/ T
87/40/17
36/ 6/ T
33/ 6/ T
100/86/19
75/35/11
I
T
4
I
T
11/ I/ T
50/50/ 4
33/11/ I
25/ 4/ T
46/12/ 4
50/11/ 5
87/60/21
22/ I/ I
25/ 2/ 2
50/ 8/ 2
100/65/a
36/25/
9/ 2/
5/ I/
9/ T/
23/ 9/
56/13/12
57/41/17
14/ 2/ T
86/45/ 3
7/ 2/ T
28/ 7/ 2
28/ 2/ 3
93/58/12
Fall
Aaropuron-Snr^hcriaaroos
Summer
Summer
Fall
(2)
O U C
50/ 1/19
100/ 2/62
(2)
O U C
50/ T/ 6
100/ 1/49
100/76/26
50/50/30
50/10/15
100/77/28
50/50/36
50/10/15
50/42/ 6
50/ 7/17
50/49/2(
100/41/31
100/47/22
100/89/57
Fall
(3)
O U C
29/ T/ T
43/ I/
14/ T/
86/83/
14/ 2/
14/ T/
Arterrisia loncifolia
T
T
7
T
T
67/
67/
33/
33/
33/
I/
T/
7/
I/
T/
(3)
O U C
T
T
T
T
T
67/29/ I
33/ T/ T
33/ I/ T
33/ 4/ T
33/24/ T
14/ 3/ T
14/ 3/ T
57/ I/ T
24/ I/ T
7/ I/ T
93/37/17
100/99/31
43/27/ I
14/ 2/ T
7/ 2/16
14/ I/ T
29/10/ I
14/ T/ T
43/12/ 2
50/10/ 5
86/62/18
14/ T/ 9
100/69/ 9
100/19/
33/ 2/
33/ 2/
67/ 2/
100/32/
T
T
T
T
T
33A9/ T
33/ 3/ T
67/23/ 5
Table 31.
Continued
*No feeding sites examined in fall.
2Genus species names listed below:
AGCR » Agropyron cristatum
ASCO
COUM
ERMU
GLLE
LASE
MEAL
MEOF
MINU
PECA
PEPU
PHLI
POER
TRDU
YUGL
=
=
=
*
■
=
=
=
=
Aster Cormutatus
Cotmandra umbellata
Eriogonun multceps
Glycyrrhiza lepidota
Laetuca serriola
Melilotus alba
M. officinalis
Mieroseris nutans
Petalostemun aandidun
P. purpareum
Phaaellia linearis
Polygonum ereatum
Tragopogon dubius
Yueaa glauea
CHNA
CHYI
JUSC
PRVI
RHTR
RICE
RONU
SYOC
■
»
=
=
Chrysotharmus nauseosus
C. vtiseidiflorus
Juniperus seopulorun
Prunus -jirginiana
P.hus trilobate
P.ibes cereun
Posa KUtkana
Symphoriearpos oaeidentalis
I
00
I
-88trends between summers which he related to abundance of forbs.
Martin
(1973) cited Eustace as reporting skunkbrush to be excellent summer
browse with protein levels of about 12 percent.
Use of yellow sweet-
clover began about mid-June in 1974, concomitant with flowering of
second year stems, but not until early August 1974 when most mature
second year stems had died, did mule deer use the first year stems of
yellow sweetclover.
Qualitative observations in 1974 indicated that
fewer second year stems of yellow sweetclover were present in pasture
IV than pasture III; and canopy coverage of yellow sweetclover was
considerably greater at feeding sites in pasture III as compared to
values obtained at feeding sites in pasture IV.
This difference may
have resulted from heavier cattle use of pasture IV in 1972 and 1973
(Table 24).
Early, heavy and continued use associated with the "A"
treatment of pasture IV in 1972 may have resulted in a high percentage
of utilization of yellow sweetclover before
reproduction in 1973.
flowering and lower
The difference in abundance of yellow sweet­
clover between pastures III and IV was reflected in the food habits of
mule deer in the two pastures during July 1974 (Table 32).
Use of
,
yellow sweetclover was greater than use of skunkbrush in pasture III,
while in pasture IV use of the latter was greater than the former.
Again the inverse relationship of use of skunkbrush and yellow sweet­
clover was evident.
Total use of forbs and browse did not differ as .
greatly between pastures as compared to the difference in use of
-89yellow sweetclover and skunkbrush.
This was largely a result of greater
use of rose in pasture III and a greater use of common salsify
(Tvagopogon dubius) in pasture IV.
Table 32.
Frequencies of occurrence (0), aggregate mean percentages of
use. (U) and canopy coverage (C) by pasture for each important
plant taxa used by mule deer at feeding sites examined during
July 1974.
Forbs ■
Pasture III1
O U C .
- Pasture IV2
O U C
24/ 4/ I
71/56/12
57/ 6/ T
86/66/24
25/ 5/ 2.
75/26/ 3
50/18/ T
100/50/14
Browse
Rhus tvilobata.
Rosa- sp.
Symphovicavpos occidentalis
Total browse .
57/21/ I
42/12/ 2
28/ T/ 3
86/33/10
100/48/ 2
25/ T/ I
25/ I/ T
100/50/11
Grass
Total grass
28/ T/55
Cornandva umbellata
Melilotus officinalis
■Tvagopogon'dubius
, Total forbs
,
-/ -/43
'•
1Seven feeding site examinations - two in the Artemisia-Agropyron and
five in the Plnus-Juniperus habitat types.
2Four feeding site examinations - one in the Artemisia-Agropyron and
three in the.Pinus-Juniperus habitat types.
Yellow sweetclover was the most important forage plant in summer
on all habitat types, except Pinus-juniperus where browse, especially
skunkbrush, appeared to be extremely important (Table 31).
Forbs were
the principle Items used by mule deer feeding on the ArtemisiaAgropyron and Sarcobatus-Agropyron habitat types.
A variety of forbs
of the Tragopogon and Poa unions were used on these two habitat types
in June.
Western snowberry was the only plant species accounting for
more than I percent of the total over-all use by mule deer during summer
where its percent utilization was below percent canopy coverage at
feeding site examinations.
Nootka rose was sought out in old burns by
mule deer, especially in the Agropyron-Symphoricarpos dominated drain­
ages in these areas.
In August and early September, chckechefry was
generally used whenever it occurred within the area of. a feeding site.
Fall
Over-all, use of browse in fall exceeded the use of forbs by mule
deer (Table 31).
Use of grass was incidental.. Yellow sweet clover,
though greatly reduced in importance, was again the single most import­
ant forage plant.
Use of skunkbrush ceased after its leaves were
drop­
ped in mid-September. Consistent use of rubber rabbitbrush, the most
important browse plant in fall, did not begin until early October,
when this plant appeared to be highly selected for on the Artemisiatridentata-Agropyron spioatwn and Pihus ponderosa-Agropyron'spicatum .
communities.
Preference by mule deer for western snowberry appeared to
increase in fall, when it became an important forage plant on both the
Pinus-Juniperus and Agropyron-Symphoricarpos habitat types.
Nootka
-91rose was used mainly in September though mule deer were observed to
eat rose hips in October and November.
Use of yellow sweetelover on
the Pinus-Juniperus habitat type, almost unchanged from summer, was
greater than that recorded for any other plant species on this type.
Increased usage of yellow sweetelover on the Artemisia Zongifolia
habitat type was associated with mule.deer digging out roots of first
year stems of yellow sweetelover.
Roots of yellow sweetelover com­
prised as much as 84 percent of the content of rumen samples collected
in October.
Analysis of rumen samples (Table 33) showed a greater
usage of eriogonum than indicated by feeding site examinations.
Rocky
Mountain juniper was of minor importance in rumen samples collected in
November.
Elk
Results are based on examination of 28 elk feeding sites totaling
6,082 instances of use during the summers of 1973 and 1974:
figures
for the first half of September 1973 and fall 1974 were 22 and 7,193,
respectively.
Four elk rumen samples collected in September 1973 and
December 1974 were also analyzed.
Summer
Forbs, specifically yellow sweetelover and bastard toadflax, were
the main taxa taken at.summer feeding sites (Table 34).
Over-all,
I
—92Table 33.
Frequency of occurrence (0)'and mean aggregate volume per­
centage (V) for each plant taxa and forage class which
occurred in three or more of 25 mule deer rumen samples
collected in October and November 1974. Trace (T) amounts
.are less than I percent.
O
Grass
Total Grass
Forbs
Eviogonum 'multieeps
Melilotus officinalis
Tragopogaon dubius
Yucca glauca"
Total Forbs
Browse .
Artemisia tongifolia ■
A Lrdplex nuttallii
Ckvysothamnus sp.
tiuniperus,scopulovum
Rosa sp.
Syrnphoricarpds occidentalis
Total Browse
V
36/ T
56/ 7
100/38
39/ T
36/ T
100/47
. 32/ T
12/ T
96/18
56/ 2
44/ 2
76/11
100/50
Mushrooms
16/ T
Unidentified.material
28/ 2
Litter
32/ T
'
.
■'
-'
Seasonal frequencies of occurrence (0), aggregate mean percentages of use (U) and percent canopy coverage (C) by habitat type and by all types
combined for each plant species used by elk which accounted for I percent or more of the use recorded at feeding sites in one or more habitat types. Trace (T) amounts are less than I percent. Numbers of feeding sites examined are in parenthesis.
FORAGE CLASS
PLANT SPECIES
USED2
GRASS <& SEDGES
AGSM
CAGE
KOCR
MUCU
POSE
STVI
Total Grass
FORBS
ARFR
COUM
ERMU
GLLE
LAPU
LASE
MEAL
MEOF
MEDE
SPCO
TRDU
YUGL
Total Forbs
BROWSE
ARLO
CHNA
CHVI
PRVI
RHTR
ROAR
RONU
SAVE
SYOC
Total Browse
All Types Combined
Summer
Fall
(28)
O U C
57/ 6/16
18/ I/ 2
39/ 9/ 2
36/ I/ 7
25/ I/ 4
78/18/42
43/11/ I
4/ T/ T
21/ 2/ I
14/ T/ T
14/ I/ T
4/ T/ I
79/54/23
4/ T/ T
7/ T/ T
32/ T/ T
100/73/34
4/ T/ T
7/
21/
11/
11/
4/
7/
64/
4/ I
T/ T
2/ T
T/ T
T/ T
I/ 2
8/10
Arterrisia-.-.^vcvpron
Summer
Fall
(22)
(8)
(i d
O U C
O U C
O U C
70/11/18
81/ 7/16 88/ 7/16
9/ T/ T
18/ I/ 2
36/ 8/ 2
64/20/ 4 25/ 6/ I
5/ T/ 3
36/ I/ 6 13/ T/ 2
9/ 2/ T
41/ T/ 6
45/ 2/ 6 38/ T/ 6
81/22/42
81/31/45 100/15/48
5/ T/ T
36/ T/ T
64/14/ 2
18/ 4/ T
9/ I/ T
86/59/18
P i n u s er-^.3
Summer
Fall
Scrocba tus-Aarovyr'CK
AgrotyrcnArterisia ZenaifeZia Survkcrioarvoe
Summer
Summer
Fall
(9)
O U C
22/ T/ 3
33/ 2/ 2
33/ 3/ 3
67/ 3/13
(3)
0 IJ C
33/ T/13
33/ I/ 2
(2)
(5)
O U C
O U C
50/42/25 100/32/18
40/ T/ T
50/ T/ T
80/25/ 3
11/ T/ 3
89/ 8/38
33/ T/ 5
67/ 1/37
50/43/28
13/ 2/ T
50/ I/ T
56/17/ 2
33/ T/ I
25/ T/ T
33/ 3/ T
33/ T/ T
18/ I/ T
9/ T/ T
11/ I/ T
64/43/13 100/75/20
89/55/29
40/ 7/ I
80/ 3/ 7
100/67/44
(2)
O V C
50/ T/ 6
Fall
Summer
(3)
(3)
O U C
100/ 4/48
Fall
(3)
O U C
100/ 4/43
(I)
O U C
100/ 5/35
33/ T/ T
50/ 1/14
33/ I/ 8
100/ 5/65
33/ T/ 7
100/ 5/55
32/ 2/ I
33/ T/ I
33/ 2/ T
100/ 5/48
60/ 3/ T
50/ 9/ T
50/ 2/ T
33/35/ I
50/14/ 3
67/58/19
Xanthium
strumarium1
ee-
Table 34.
50/35/ I
100/50/35
100/ 6/15
100/ 3/ 4
100/ T/ I
100/ 9/ T
100/73/38
100/58/40
100/93/60
33/ 6/ 2
60/29/ 6
100/71/ 9
50/ I/ T
100/58/14
100/89/61
50/ 5/ T
45/ I/ T
5/ 4/ T
95/69/22
18/ I/ T
9/ T/ T
9/ 3/ T
22/ I/ T
100/63/26 100/80/24
100/79/37
33/ T/ T
33/27/ 2
100/86/21
100/55/17
80/33/11
100/83/12
67/ 7/ 4
9/ T/ T
100/93/15
100/91/72
100/ 2/ T
67/ I/ T
25/ 4/ T
33/ 6/ T
11/
44/
11/
22/
18/ T/ T
18/ 4/ T
5/ I/ T
8/
2/
T/
T/
33/10/ T
T
T
T
T
50/14/ 2
50/ I/ I
30/ 1 / 2
33/ T/ 2
50/ 2/ 5
23/ 4/ 2
59/ 9/11
27/ T/ T
55/ 5/12
25/ I/ T
63/ 5/11
33/ I/ 3
67/13/11
Ino/13/ll
50/ 2/ R
50/ I/ T
100/17/ 8
33/ 6/ T
67/ 7/ V
33/ 4/ 5
33/ 4/10
67/26/15
100/37/24
100/ 2/ T
100/ 2/ I
Table 34.
Continued
1No feeding sites examined in fall.
2Genus species names listed below:
AGSM
CAGE
KOCR
MUCU
POSE
STVI
=
=
=
=
”
Agropyvon smithii smithii
Carex geyeri
Koeteria cristata
Muhlenbergia ouspidata
Poa eecunda
Stipa virioula
AFRF
COUM
ERMU
GLLE
LAPU
LASE
MEAL
MEDF
MEDE
SPCO
TRDU
YUGL
Artemisia frigida
Comandra umbellata
Eriogonum multoeps
Clyoyrrhiza lepidota
Lactuaa pulehella
L. serriola
Melilotutus alba
'4. officinalis
Mentzellia deoapetala
Sphaeralcea aoeeinae
Tragopogon dubius
Yueaa glauaa
ARLO CHNA =
CHVl »
PRVI RHTR"ROAR RONU =
SAVE SYOC -
Artemisia longifolia
Chrysothamnu nauseosus
C. viseidiflorus
Prunus virginiana
Rhus trilobate
Rosa arkansana
Rosa nutkana
Saraobatus vermiaulatus
Symphoriaarpos oaaidentalis
\o
**
-95grasses and grass-like plants were second in importance.
Junegrass
appeared to be the only grass consistently selected for at feeding sites.
Western wheatgrass was also taken in considerable quantities in late
August 1973.'
In the summer of 1973, grasses accounted for 54 percent
of the total instances of use as compared to 5 percent in the summer of
1974.
The difference was due largely to the sharply increased use of
grasses in August 1973.
Forbs accounted for 32 percent of the total
instances of use in 1973 while in 1974 forbs accounted for 90 percent
of the use recorded, largely as a result of use of yellow sweetclover
throughout the summer of 1974.
Browse, though of moderate importance
in 1973, was of relatively minor importance over-rail.
Yellow sweetclover was highly selected for on all types and was
the dominant forage
plant for elk on all but the Sarcobatus-Agropyron
habitat type during summer (Table 34).
Junegrass and bastard toadflax
were often taken in conjunction with each other when elk fed in the
Artemisia tvidentata-Agvopyvon spioatum and Pinus-Agropyron communities.
Chokecherry, in August and early September, was of moderate importance
at elk feeding sites on the Pinus-Juniperus and Avtemisia tongifolia
habitat types.
’
Fall
Use of forage classes changed little from summer to fall (Table 34)
because of the persistent abundance of first year stems of yellow sweet­
clover throughout the fall of 1974.
Yellow sweetclover was again the
—96principal plant used in all habitat types except for the SarcobatusAgropyron type.
Wild licorice (Glycyvrhiza lepidota) was frequently
taken in small amounts early in fall, while use of soapweed was. record­
ed only in December.
During the fall of 1974, Junegrass appeared to be
the only grass selected for by elk.
However, in September 1973, grass
accounted for 91 percent of the total instances of use recorded at
three feeding site examinations conducted in the Sarcobatus-Agropyron
habitat type; principal among these were western wheatgrass, Junegrass
and Sandberg bluegrass.
It appeared the normal, late September shift
from forbs to grasses, as reported by Mackie (1970), occurred during
August in 1973 but not in the fall of 1974.
browse plants shifted from summer to fall.
Species importance among
Preference for western,
snowberry appeared to increase and green rabbitbrush was important on
the Pinus-Juniperus dominated northerly slopes and Agropyron-Symphorcarpos dominated coulee bottoms.
The rumen sample collected in September 1973 indicated a greater
importance of chokecherry than feeding site data for that period.
Aside from this, difference, rumen sample date (Table 35). tended to
support feeding site data for fall.
-97Table 35..
Frequency of occurrence (0) and mean aggregate volume per­
centage (V) for each plant taxa and forage class which
accounted for I percent or more of the.volume in one or more
of four elk rumen samples collected in September 1973 and
December 1974. Trace (T) amounts are less than I percent.
0
Grass
Total Grass
V
100/18
. Forbs
Melilotus officinalis
Opuntia polycantha
Yucca glaucd
Total Forbs
75/43
25/ T
50/ 4
100/48
Browse
Artemisia. Iongifolia
Gutierrezia sarothrae
Populus sargeyitii
Prunus virginiana
Symphoricarpos sp.
Chrysothamnus sp.
Total Browse
50/ T
25/ T
25/ T
25/5
100/ 2
100/ 4
100/13
Unidentified material
:
50/21
—98~
Cattle
Results are based on examination of 23 cattle feeding sites total­
ing 7,249 instances of use during the summers of 1973 and 1974:
figures
for the first half of September 1973 and fall 1974 were 18 and 5,631,
respectively.
Four cattle rumens collected from late August through
November 1974 were also analyzed.
Summer
Grasses and grass-like plants ranked as the most important forage
class during both summers with western wheatgrass and plains muhly
accounting for over one-half of the total instances of use (Table 36).
Forbs were second in importance over-all but accounted for only 8 per­
cent of the total use recorded at cattle feeding sites examined in 1973,
when browse was slightly more important.
percent of the recorded use.
In 1974, forbs comprised 39
Yellow sweetclover accounted for over 90
percent of the use of forbs by cattle in summer.
Browse consisted of
greasewood and was of. minor importance at all times.
In the Artemisia-Agropyron habitat type, plains muhly appeared to
be selected oyer western wheatgrass when both occurred on a feeding
site (Table 36).
The most intensive.use of grass and grass-like plants
occurred on the Pinus-Juniperus habitat type.
Yellow sweetclover was .
the dominant taxa used on the Agropyron-Symphorcarpos and Xanthiim .
StTimaviwn habitat types and was. of considerable importance on the
Table 36.
Seasonal frequencies of occurrence (0), aggregate mean percentages of use (I") and percent canopy coverage (C) by habitat type and by all types
combined for each plant species used by cattle which accounted for I percent or more of the use recorded at feeding sites in one or more
habitat types. Trace (T) amounts are less than I percent. Number of feeding sites examined are in parenthesis.
AntKiaia
FORAGE CLASS
PLANT SPECIES
USED2
GRASS 6, SEDGES
AGSM
AGSP
AGTR
BOGR
CALO
CAGE
HOJU
KOCR
MUCU
STVI
Total Grass
BROWSE
ARLO
ATNU
CHNA
RHTR
ROAR
SAVE
SYOC
Total Browse
(23)
O U C
96/29/19
9/ T/ 3
4/ I/ T
35/ T/ 2
9/ I/ T
9/ 2/ T
4/ T/ T
39/ 2/ 2
52/22/ 6
61/ 6/ 6
100/65/49
26/ T/ T
61/26/11
17/ T/ T
4/ T/ T
9/ T/ T
78/28/19
4/ T/ T
9/ I/ T
17/ I/ T
26/ 4/ 2
56/ 7/ 8
Summer
(18)
(13)
O U C
O U C
94/45/23 100/20/14
22/ 4/ 5
Fall
Pinus-Jur.iperus
Summer
Fall
Agropyron-
Saroobatus-Apropyron
Lcngifclia
Su^rphorioarpos
Summer
Summer
Summer
(4)
(3)
(8)
(3)
O U C
O U C
O U C
O U C
100/50/28 100/29/ 7 75/ 4/ 6 100/72/28
12/ I/ 5 67/ I/ 6 75/14/ 9
Fall
(4)
O U C
100/87/35
Fall
(i)
O U C
100/ 2/ I
(3)
O U C
100/38/48
Fall
(i)
O U C
100/48/42
=
=
=
=
=
=
Summer
a)
O U C
100/26/10
17/ I/ 5
11/ I/ I
54/ 2/ 4
15/ 2/ T
33/ I/ T
37/ 2/12
100/ 3/13
25/ 4/ I
67/12/17
6/ T/ T
61/12/ 2
33/ 4/ 4
67/ 5/ 5
100/72/46
46/ 3/ 3
77/29/ 8
46/ 2/ 5
100/60/45
100/11/ 9
100/84/55
25/ T/ T
25/ T/ T
25/ T/ T
100/ 4/ 7
100/93/49
100/ 5/15
100/18/16
100/56/64
100/ 4/ 2
100/52/46
100/31/21
25/ 6/ T
75/34/ 7
33/ 6/ T
50/ I/ T
100/60/ 5
67/42/38
100/48/50
100/65/17
33/ T/ T 75/ I/ T
67/ 8/ 9 100/42/ 8
33/ 6/ 6
50/ 2/ 3
100/60/ 7
67/43/43
100/48/50
87/24/ 3 100/ 4/ 3 75/ 6/ 4
25/ 4/ I 67/36/10 75/12/ 6
62/ 3/ 2 67/ 8/ 5 75/ 8/ 9
100/84/51 100/91/52 100/48/40
28/ T/ T
6/ I/ T
72/19/ 5
46/ I/ T
25/ I/ T
69/28/ 9
75/11/ T
28/ T/ T
89/22/ 7
8/ T/ T
92/31/18
100/14/ 4
8/ I/ T
25/ T/ T
33/ 5/ I
33/ T/ 2
100/ 5/10
100/ 2/ 7
6/
17/
6/
6/
T/
2/
T/
T/
T
T
T
T
39/ 2/ 2
17/ I/ 2
68/ 6/13
25/ I/ T
8/ 2/ T
23/ 2/ T
8/ 4/ T
S3/ 9/ 7
100/31/ T
33/ T/ T
33/ T/ T
25/ 3/ T
25/ T/ T
12/ T/ T
37/ 2/12
50/ 4/ 9
67/ I/ 6 100/10/20
100/10/11
100/ 5/ 7
100/10/11
100/ 5/12
100/ 2/ T
100/ 2/ T
100/35/12
33/ I/ T
33/ I/ 9
1No feeding sites examined in summer.
zGenus species names listed below:
AGSM
AGSP
AGTR
BOGR
CALO
CAGE
HOJU
KOCR
MUCU
STVI
Xar.-thiur"
Agropyron smithii
A. spicatum
A. trachycalum
Bouteloua gracilis
Calamovilea longifolia
Carex geyeri
Hordeum jubatum
Koleria aristata
Muhlenbergia cuspidata
Stipa Viridula
COUM
GLLE
MEOF
PEPU
RUME
TRDU
«
=
=
»
«
-
Conmandra umbellate
Glycyrrhiza lepidota
Melilotus officinalis
Petalosteum purpareum
Rumex mexicanus
Tragopogon dubius
ARLO
ATNU
CHNA
RHTR
ROAR
SAVE
SYOC
=
=
■
Artemisia longifolia
Atriplex nuttallii
Chrysothamnus nauseosus
Rhus trilobate
Rosa Arkansa
Sarcobatus vermiculatus
Symphoricarpos occidentalis
100/69/32
-99-
FORBS
COUM
GLLE
MEOF
PEPU
RUME
TRDU
Total Forbs
Arterrlsic:-Agropyron
All Types Combined
Summer
Fall
-XOOArtemisia-Agropyron habitat type.
Intensive use of skunkbrush was
noted in early August 1974 when cattle first entered pasture IV.
Fall
Although the relative importance of forage classes did not change
from summer to fall (Table 36), the use of grass increased from 56 per­
cent in the summer of 1974 to 70 percent of the total instances, of use
in fall.
Use of plains muhly diminished sharply after September and
use of Junegrass became increasingly important for the remainder of the
fall period.
Bluebunch wheatgrass received some use in November when
cattle made increased use of side ridges and the Pinus-Juniperus
habitat type.
Within the Pinus-Juniperus habitat type, however, use of
yellow sweetclover greatly exceeded that of other plants.
clover was again the most important forb.
Yellow sweet-
Greasewood, taken frequently
in low amounts, was the major browse plant; Npttall saltbrush (AtvvpZex'
nuttalZZi) and western, snowberry were also of minor importance to
cattle.
Minor use by cattle of green rabbitbrush, silver sagebrush and
western snowberry was noted in areas of heavy use in the AgropyronSymphoricarpos habitat type but was not recorded in any feeding site
examination.
Most feeding site examinations were conducted after an area was
initially grazed.
Observations indicated that cattle were less select­
ive with subsequent grazing of an area.
The discrepancy between data
-101from rumen samples (Table 37) and feeding site examinations was probably
a result of this sampling bias.
Table 37.
Frequency of occurrence (0) and mean aggregate volume per­
centage (V) for each plant taxa and forage class which
accounted for I percent or more of the volume in one or more
of four cattle rumen samples collected August through
November 1974. Trace (T) amounts are less than I percent.
0
Grass
Total Grass
Forbs
Melilotus' offioinalis
Total Forbs .
Browse
Saroobatus vermiculatus
Symphorioarpos sp.
Total Browse.
V
100/97
50/ I
50/ I
75/ I
50/ T
100/ 2
DISCUSSION
Interspecific Range Relationships
Inlander (1958) considered the prime factors controlling inter­
specific competition between two species of ungulates to be the extent
to which the two species use the same area and prefer the same forage
plants.
Smith and Julander (1953) differentiated interspecific compe­
tition into two broad categories; forage competition and land use
competition.
Forage competition occurs whenever the supply of a forage
plant is inadequate for the requirements of all individuals of one or
both species.
Land use competition can occur even though forage
supplies are adequate for all individuals of both species because
reducing of numbers of one would allow greater numbers of the other.
These criteria were used as guidelines to evaluate interspecific
competition.
Mule Deer and Cattle
Broad differences recorded for use of range between mule deer and
cattle indicated that under conditions existing during this study,
opportunities for forage competition were limited during summer and
fall, except.for June when 75 percent of the mule deer and 81 percent
of the cattle observed were on the Artemisia-Agropyron and SarcobatusAgropyrpn habitat types.
From August to December, well over one-half
-103of the mule deer observed were in the Pinus-Juniperus habitat type.
During the same period not more than 15 percent of the cattle observed
in any one month were in this habitat type.
During both summer and
fall, more than one-half of the mule deer and less than one-?fourth of
the cattle observed were on slopes greater than ten degrees.
Prolonged
usage of a pasture by cattle resulted in increased usage by cattle of
areas and habitat types preferred by mule deer but only to a minor
degree.
Grass, which was of incidental use to mule deer, was the major
forage class used by cattle both summer and fall.
The only forage
plant used in common and important in the diet of both ungulates was
yellow sweetclover.
When this plant was in low supply in summer,. mule
deer shifted their use to skunkbrush, a plant normally little used by
cattle.
No shifts in mule deer distribution within and between years
related to grazing by cattle were documented.
Skolvin (1968) reported
no significant preferences by migratory mule deer for areas not grazed
by cattle or any significant preferences for any particular level of
stocking of cattle.
The data suggested that grazing by cattle affected both forage
■
■
'
class usage and movements within normal home range of mule deer.
The
striking differences recorded between food habits of mule deer feeding
in pastures III and pasture IV. in July 1974 was a result of pasture
treatments the previous two years.
During this month forbs were the
dominant forage class used in pasture III while in pasture IV
-104use of browse equaled use of forbs as a result of a lower abundance of
second year stems of yellow sweetclover.
The radio marked'yearling male,
whose home range was grazed by cattle, exhibited abnormally long move­
ments which were comparable to the two-year-old male.
This suggested
a more intensive use of home range by the yearling male.
Dasman and
Taber (1956) found yearling males to be less mobile than older males
and to have home ranges similar to females.
this study.
This was not the case in
It would be expected that the above two factors would
ultimately be manifested in the reproductive success of mule deer.
The
more intensive use by cattle of pasture IV as compared to pasture. Ill
from 1972 through 1974 was viewed as a partial explanation for the
lower reproductive success of mule deer in pasture IV.
Failure of mule
deer in pasture IV to respond to the decline in use of the pasture by
cattle during this period through increased reproduction, as.was
observed for mule deer in pasture III and as would be expected if heavy
grazing by cattle did in fact affect mule deer reproduction, was
probably related to the reduced amounts of a crop of yellow sweetclover
for two years in pasture IV.,
Elk and Cattle
Distribution of elk within and between years suggested that land
use competition occurred between elk and cattle.
The large increase in
numbers of elk observed on the study area in 1974 ever 1973 was
-105attributed to elk concentrating In areas not grazed,by cattle,
especially the rest pasture.
The preference displayed by elk for
pasture III over pasture IV during the summer of 1974., when cattle were
not present in either pasture, was related to both the greater abundance
of yellow sweetclover in pasture III resulting from pasture treatments
the two previous years and the absence of cattle grazing.
This differ­
ence in distribution precluded any possibility of severe forage com­
petition between elk arid cattle occurring during the study.
Elk made
use of steeper slopes and more intensive use of the
Pinus-Junlperus habitat type than did cattle in a manner similar to
that described for mule deer.
Elk, however, made greater use of the
main ridge tops and major coulee bottoms, areas of primary range for ,
cattle, than did mule deer.
Grass, which was the most important forage
class, used by cattle, was of. secondary importance to elk during most
periods of the study.
Elk preferred forbs when available.
However,,
the shift to grasses by elk in August 1973 resulted in food habits
'
which closely paralleled those of cattle.
.
Grass comprised 62 percent
of the content of the elk rumen sample collected in September 1973.
Western wheatgrass and Junegrass, both of which were used intensively
by cattle, collectively accounted for 76 percent of the total iristances
of use at elk feeding sites in September 1973.
This shift to grasses
in 1973 represented opportunity for direct forage competition to occur
between elk and cattle.
Home range and movement data iridicated this
“106situation would be. important only when cattle grazed contiguous, large
blocks of land.
The mobility of elk allowed them to seek out areas
where forage conditions were optimum.
.
Skolvin (1968) found use of an area by elk decreased significantly
as levels of grazing by cattle increased and use of an area by elk
increased significantly when it was not grazed by cattle.
Similar
results were found in this study. Where grazing by cattle affected elk
largely through distribution.
.
Mule Deer and Elk
.Range use of ,mule deer and elk during summer overlapped at m a n y
points.. Use of slope and exposure was quite similar though mule deer
used.slightly steeper slopes than elk.
Not only was over-all use of
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habitat types comparable, but also trends in use of habitat types, with­
in summer corresponded, quite closely.
those areas preferred by mule deer.
Elk also, made intensive use of
As long as forbs were available in
summer, food habits of mule deer and elk overlapped.
both showed marked preferences for yellow sweetclovef.
Elk and mule deer
Divergence was.
noted in summer food habits as forbs were desiccated; mule deer
increased their use of skunkbrush and other browse while elk increased
their use of grasses.
Mackie (1970) pointed out that any factdr
causing mule deer to shift their use to browse earlier than normal
caused a sequential advancement in use of the less palatable browse
—107species as supplies of more palatable browse species were exhausted.
Competition for forbs between mule deer and elk can be one of these
factors;
In fall, range use of mule deer and elk continued to be
similar but food habits were substantially changed.
Browse was the
most important forage class to mule deer and forbs continued to be the
over-all most, important forage class to elk.
However, grasses, such as
in September 1973, would normally be the dominant forage class used by
elk in fall as reported by Mackie (1970), Rouse (1957). and Knight (1970).
Elk did make minor use of browse preferred by mule deer in fall.
Real possibilities of forage competition between mule deer .and elk
existed, especially if numbers of elk were allowed,to increase.
The
low to moderate fawn:doe ratios for mule deer obtained during the two
winter surveys compared to the high calf:cow ratios for elk suggested
that land use,, competition between mule deer and elk and/or cattle had
occurred.
„
Comparison of Rest Rotation Grazing to Season Long Grazing
Only a few major differences were found in range use and food
habits of mule deer, elk.and cattle between those reported here and by
Mackie(1970) on an area where season long grazing by cattle, was in,
effect.
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Home ranges of mule deer,in this study were considerably larger
than those described by Mackie.
This difference might be explained by
—108—
the greater diversity of habitat types and a population density about
..
three times greater on Mackie1s study area.
Smaller standard diameters
of home range at higher population densities was reported by Knight
(.1970) for elk and by White (1964) for deer mice CPevomysaus leucopus).
The smaller, group sizes of elk and more^intensive use of the PinusJuniperus habitat type by elk in fall observed in this study as compared
to that repotted by Mackie was probably a result of persistence of first
year stems of yellow sweetclover in the fall of 1974 and disturbance
by hunters throughout most of the fall period.
The greater use of. the Sarcobatus-Agropyron habitat type by cattle
reported, in this study as compared to that reported by Mackie was
probably related to the well developed system of reservoirs and wells
in the major, doulees in Nichols. Coulee RCA.
Food habits of mule deer, elk and cattle between study areas were
quite comparable, especially when seasonal trends and minor vegetational
differences are., considered.
Except for the difference of food habits
of mule deer between pastures III and IV noted in.July 1974 and the
heavy use by. elk of the rest pasture, it would appear that the over-all
effect of rest rotation grazing has not affected any major changes in
the range use and food habits of mule deer, elk and cattle in summer
and fall.
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Management Implications
The benefits derived from a rest rotation grazing system to key
forage plants and to cattle have been reported by Hormay and Talbot
(1961), Johnson (1965) and Ratcliff and Rader (1962).
The mobility of
elk, found in this study, enabled them to concentrate, in pastures and
areas previously not grazed by Cattle; this appeared to make rest
rotation grazing equally beneficial to elk.
However, the limited
mobility of mule deer made them vulnerable to any situation which
resulted in intensive use of their home, range by other ungulates.
The
heavy use of a pasture by cattle associated with the "A" and "B" treat­
ments and by elk concentrating in pastures and areas not grazed by
cattle during early summer under the "C" treatment and throughout the
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year of "D" treatment resulted in conditions where mule deer were .
frequently exposed to heavy use of their home range by other ungulates.
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
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—H O Allen, E. 0. 1968. Range use, foods, condition, and productivity of
white-tailed deer in. Montana." J. Wildl. Mgmt. 32(1) :130-141..
Booth, W. E. 1950. Flora of Montana, Part I, Conifers and Moncots.
Res. Found; Montana St. Coll., Bozeman.
232 p.
, and J..C. Wright.
1959. Flora of Montana, Part II, Dicotyledons.
Montana St. Coll., Bozeman.
280 p.
Cole, L. C. 1949. The Measurement of interspecific association.
Ecology 30(4):411-424.
Dasmann, R. F, and R. D. Taber.
1956. Behavior of Columbian black- ■
tailed deer with reference to population ecology. J. Mammal.
37(2):143-164.
Daubenmire, R; 1959. A canopy-coverage method of vegetatidnal analysis.
Northwest Sci. 33:43-64.
Gieseker, L . .F . ■ 1926. Soils.of Phillips County.
. Sta. Bull. No. 199. 61 p.
Montana A g r . Expt;
Harvey, M . J . and R. W. Barbour. 1965. Home range.of Microtus
ochogaster as determined by a modified minimum area method.
J. Mammal. 4.6(3) :398-402.
J
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Hormay, A. L: and M. W. Talbot.
1961. Rest-Rotation Grazing.... A New
Management for perinial bunch grass ranges. .USDA forest Service,
Prod. Res. Rpt. No. 51, 43 p.
Johnson, W. M.. 1965. Rotation, Rest-Rotation, and Season-Long grazing
on a Mountain range in Wyoming. USDA, Forest Service Rky. Mtn.
F&R Expf. Sta., ReA. Paper EM-14.
16 p.
Julander, 0. 1958.. Techniques.in studying competition between big
game and livestock. J. Range Mgmt. 11(1):18-21. .
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Knight,.R ..H. 1970. The Sun River.Elk Herd.
'66 p.;
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Wildlt- Monograph No. 23,
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Mackie, R. J. 1970. Range ecology and relations of mule deer, elk,
and cattle in the Missouri River breaks. "Wild!. Monograph No. 20,
77 pi- / \
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-111Martin, A. C., R. H. Gensch, and C. P. Brown.
1946. Alternative
methods in upland game bird food analysis. J. Wildl. Mgmt.
10(1):8-12.
Martin, P. R. 1973. Ecology of skunkbrush sumac (Rhus tvitibata Nutt.)
in Montana, with special reference to use by mule deer. MS thesis,
Monta. State Univ. 97 p.
Mohr, C. 0. 1947. Table of equivalent populations of North American
small mammals. Am. Midi. Nat. 3 7 (I):223-249.
Pyrah, D. B. 1973. Measuring Sage brush canopy coverage and cover
index. Montana Fish and Game Dept. Job Compl. Rept., Project
W-105-R-7, 8. 127-131.
Ratcliff, R. D. and L. Rader. 1962. Drought hurts less with RestRotation management. USDA, Forest Service, Pacific Southwest
Forest & Range Expt., Res. Note #196, 4 p.
Robinette, W. L. 1966. Mule deer home ranges and dispersal in Utah.
J. Wildl,. Mgmt. 30.(2):335-349.
Rouse, R. A.
1957. Elk food habits, range use, and movements,
Gravelly Mountains Montana. MS thesis, Mont. St. Coll., 29 pp.
Skovlin, J. M. et al. 1968. The influence of cattle’management on
deer and elk. Trans. N..Am. Wildl. Nat. Resour.•Conf. 33:169-181.
Smith, J. G., and 0. Inlander.
1953. Deer and sheep competition in
Utah. J. ,Wildl. Mgmt. 17(2):101-112.
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Climatological data- Montana.
Monthly Summaries. U. S . Govt. Print. Off., Washington, D. C.
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White, J. E; 1964. An index of the range of activity.
Nat. .71(2):369-373.
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MONTANA STATE UNIVERSITY LIBRARIES
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