Range use, relationship to logging, and food habits of the elk in the Little Belt Mountains, Montana
by John Ballard Kirsch
A thesis submitted to the Graduate Faculty in partial fulfillment of the requirements for the degree of
MASTER OF SCIENCE in Fish and Wildlife Management
Montana State University
© Copyright by John Ballard Kirsch (1962)
Abstract:
An elk (Cervus canadensis) study was conducted in the Little Belt Mountains of central Montana
during the summer of 196.0 and the spring, summer and early fall of 1961 to investigate distribution,
movement, range use, relationships to block clear cut logging of lodgepole, pine (Pinus contorta) and
food habits. Elk movements and,range use were discussed in relation to broadly designated plant
communities and more specific vegetative types if heavily used by elk. Size, age and moisture
conditions of the lodgepole clear cuts were correlated with elk usage. A vegetational analysis of two
clear cuts and the adjoining uncut lodgepole was used to compare the changes resulting from logging.
Food habits during spring, summer and early fall were determined from 95 feeding sites examined.
Supplementing the feeding observations was the inclusion of data from 41 elk rumens, most of which
were collected during the late fall and winter of 1955 and 1956.
J .RANGE. USE, RELATIONSHIP TO LOGGING, AND FOOD.HABITS
OF THE ELK IN THE LITTLE BELT MOUNTAINS, MONTANA '
by
JOHN BALUARD KIRSCH
A thesis submitted to. the Graduate Faculty in partial
fulfillment of the requirements for the degree
of
MASTER OF SCIENCE
in
Fish and Wildlife Management
Approved s
MONTANA STATE COLLEGE
Bozemana Montana •
November, 1962
iii
ACKNOWLEDGEMENT
The author is indebted to the following among others, and wishes to .
acknowledge their aid during the studys
Dr= Don C= Quimbyg Montana State
College, who directed the study and aided in the preparation of the manu­
script; Glen F= Cole, Montana Fish and Game Department, for project plan­
ning, field assistance and permission to use the elk rumens included in .
this study; D r 0 John
Ho
Rumelys, Montana State College? for field assist-
ance and the photographs illustrating vegetative types; Dr= W= E= Booth,
Montana State College, for aid in verification of plant specimens; Robert
Varner, manager of the Judith Game Range, for hospitality and cooperation;
Kenneth Greer, Montana Fish and Game Department, for use of laboratory
t'
.
facilities and cooperation; the IL S= Forest Service for use of facilities
and information on lodgepole pine clear cut areas='
The writer was employ­
ed by the Montana Fish and Game Department under Project W-75-Rr-6 and
W-98-R.-2 during the study =
L
TAsPtE OF CONTENTS
Page,
Abstract
vii ;
. . . . . .
Introduction
•o*
***
• •••
Description of area . . . . . . . . . . . . . . . .
Ponderosa pine woodland - grassland community.
Ponderdsa pine pommunity . . , . . . . . » . .
Douglas fir community > . . . . . .
.
• p
Douglas fir forest type .
. .
Lodgepole pine type t . .
. ..
».
».
Open conifer - park type V , .. .
Burn type . © * .
• * . . ■ .v ?
Spruce ~ fir community . . .. . . .
• *
Spruce forest type . . . . . .
Lodgepole pine type . .
, . .
Open conifer
park type ... , o'* * * *
* ,• .t e •
•• • •• •
« • * •
f
. • . ..
.
« •"? '•
•
•
.3
■■.4 ■
5 '
- ?:■'
•
*
'
• < r'
• •
•
.
•.
• ...
.
8
8
.•...
•* 9 . »_ * . * .• ' 9
9 *■ • • o ’«'*
o e • • o • o ♦ f 9 • • . •■ • .
and movement's . . . . . . . . .
'I'
•
. 8,
‘ .9.
9
. ... .
13 y
0 * * * 9
Elk relationships to block clear cutting of lodgepole pine
Elk food habits . ........... .......................................
Spring (iApril ^ May 1961
Summer. (June, July, August 196J.).. . • • • • • • • •. * • • •-.
Summer (June 15-July 31, 1960 and 1961) . . . . . . . . .
Fall (.September) . # ... . . . . . . © . * .. * . * . . . .. ■. ®
Ponderosa pine woodland - grassland community . . . . . . . ..
Douglas fir community (open conifer -r park type) . . . . . . .
Spruce - fir community (open conifer - park type) . . . . . .
Lodgepole pipe block clear cuts
Rumen' analyses . . . . . ..... . . , . . . . . , . . ... . ...
Discussion . . . . . . . . . . .
........... . . . . . . . . . 3
29 .
32 •
.33
33
34
35
35.
35
35
36
8
Summary
40
Literature cited
»
42
■v
LIST OF TABLES
Table
I.
II.
III.
IV.
V.
VI.
VII.
VIII.
Page ,
Percentage distribution of observed elk on designated vege
tative types by periods, 1960 and 1961
14 ■
Physical characteristics of block clear cuts and .a. summary
of elk observations in the Beadhorse Sale Area . . . . . . .
22
Elk use in relation to size of clear cuts
Elk and elk groups observed during 1960 and 1961 in block
clear cuts of the Deadhorse Sale Area
25 .
Physical, quadrant tree and canopy-rCoverage data for two
clear cuts and three adjacent uncut lodgepole pine sites on
the north slope west of Spring Creek road, Deadhorse Sale
Area, September 15, 19 and 20, 196.1 . . . . . . . . . . .
.
26
Elk food habits as determined from 12,008 instances of plant
use at 95 elk feeding sites . . . . . . . . . . . . . . . . .
30
Analyses of 41 elk rumens collected in the Little Belt
Mountains, 1955 and 1956 . . . . . . . . . . . . . . . . . .
37
Per cent seasonal forage Blass use comparison of four
Montana elk studies . . . . . . . . . . . . . .
. . . . . .
38
vi
LIS-X OF FIGURES
Figure
I,
2
.
3.
Page
Elk study area, Little Belt Mountains, Montana » , . . , . , ,
Jhe ponderosa pine woodland-grassland community in the fore­
ground with the Douglas fir forest and burn type (right) in
background 0 0 0 0 0 0 0 0 0 0 -« 0 0 0 0 0 0 0 0 * 0 0 0 0 0
The open conifer - park type of the Douglas fir (background),
with restocking lodgepole pine clear cut (foreground)
4.
Recent block clear cutting of dense lodgepole pine , . .
5.
Block clear cutting within the lodgepole pine type of the •
spruce-tf ir community « * 0 * 0 0 0 0 0 0 . o * , . * . , 0 0
6
.
2
0
6
« » 10
0
10
An open ridge top park at 8,150 feet elevation in the sprucef ir comm,unity o * e o * * * o o * * o
* 0 . 0 0 * 0 0 0 ., . . l l
7,
The spruce-fir community with open conifer - park type in
background (southern exposure) and depse spruce forest in
foreground (northern exposure). * . . * * * « * . * . * « . . * I!
80
Clear cut blocks of lpdtfbpole pine in the Deadhprse Sale Area*
(Location of vegetative measurements discussed in the text
designated by X-symbols) , * . . o * * . * . . * . . « 0 . 0 * 21
vii
' ABSTRACT
An .elk (Cervus canadensis) study was conducted in the Little Belt
Mountains of central Montana during the summer of 196.0 and the spring,
sujmmer and early fall of 1961 to investigate distribution^ movement,
range use, relationships to block clear cut logging of lodgepole pine .
(Pinus contorts) and food habits. Elk movements and,grange use were dis­
cussed in relation to broadly designated plant communities and more
specific vegetative types if heavily used by elk. Size, age and.moisture
conditions of the lodgepole clear cuts were correlated with elk usage. A
vegetatibngl analysis of two clear cuts and the adjoining uncut lodgepole
was used to compare the changes resulting from logging, Food habits dur­
ing spring, summer and early fall were determined from 95 feeding sites .
examined. Supplementing the feeding observations was the inclusion of
data from 41 elk rumens, most of which were collected during the late fall
and winter of 1955 and 1956.
J
- 1-
INTRODUOTION
The gbundance of game in- central Montana before 1880 has been re­
corded by Worthen (1928).
By 1885 this..area had become one of the more
important stock raising regions of the state (Phillips 9 1925).
The Little
Belt Mountains 9 which are a prominent feature of the landscape, project
plainsward into this region from the Rocky Mountain cordillera.
\
Elk (Cervus canadensis) were present (Smith 9 1929; Rush 9 1930) but not
numerous in the Little Belts during the 1920'h.
86
According to West (1941),
elk were transplanted to this area from Yellowstone National Park in
1927-28.
.
He gives the 1941 elk population in the Judith River drainage as
300 animals.
Elk from.an unconfined private herd may also have augmented
the native animals in the region in 1929 (Rush, 1930).
The present elk.
population of the Little Belts roughly numbers more thah l><)00 animals.
A study in 1941-42 (McDowell, 1942) reported that 60 per cent of the
winter range within the forest boundary was of little forage value. . Elk
began to winder on the privately controlled foothills which had long been
over-used by sheep and cattle (Leiberg 9 1904).
To alleviate the resulting
game-livestock conflicts in the foothills, the Montana Fish and Game. .Com­
mission in the early 1940‘s began acquiring strategically located land for
elk winter range,
This relatively small tract, about 7,000 acres at present,
becomes a winter concentration, point for elk funneling out of the Judith
River drainage from distances up to 20 airline miles.
Varner (1962) re-
ported 550 elk wintering on the Judith Game Range (Fig. I) during the
winter of 1961-62.
A detailed study of mule deer (Odocoileus hemionus)
LEGEND
POND EROSA PINE WOOD LA NOGRASSLANO COMMUNITY
PONDEROSA PINE COMMUNITY
DOUGLAS FlR COMMUNITY
SPRUCE-FIR COMMUNITY
M AIN
DIVIDES
LOCATION OF MARKED ELK
6 MILES
IO
I
FIG.I.ELK
STUDY
A R EA ,
LITTLE
BELT MOUNTAINS, MONTANA.
range use and food habits on this game range and the Judith River drainage
was conducted by Lovaas (1958)„
Information of a fragmentary nature has been accumulating on this elk
herd since 1939.
herd.
Picton (1961) published a population study of the Judith
A detailed study of the elk was needed to provide basic information •
to manage the herd in relation to other important land uses. . The study,
reported here, was concerned with elk distribution, movements, range use.
and food habits during spying and summer.
harvest and elk range-use yvas investigated.
The relationship between timber
The expanded market, in recent
years, for lodgepole pine ('Rinus.contorta) has increased logging in the
area, resulting in marked changes in the vegetative cover.
DESCRIPTION OF AREA.
The Little Belt Mountains, extending northwest-southeast for about 70
miles, are 30 to 40 miles', wide in the western portion and draw to a point
in the eastern extremity.
The maximum elevation is 9,175 feet above mean
sea level, while the adjoining plains are about 5,200 feet.
Gieseker e.t o
a l . (1953) gives their area,as 1,648 square miles.
The study area of approximately 600,000 acres comprised all of Lewis
and Clark National Forest within the Little Belts east of U. S. Highway 89,
and certain foothills adjacent to the forest boundary (Fig 9 I).
The topography of the study area can be described as a low plateau­
like mountain range cut by deep narrow canyons.
However, local regions,
especially'in the north, have rugged rocky crests with prominent cliffs
ahd talus slopes.
Relatively broad, flat, open ridges separate water
4
c—4
courses.
,According to Weed (1899) the deep trenched canyons are open and
wide in soft shaly rock, and narrow in. the hard limestone strata.
Towards
the bases of the mountains, canyons are narrow and straight-walled while
the head waters flow in relatively open wide valleys.
i.
Two main crests delimit areal drainage (Fig. I).
Ypgo, Middle Fork
and South Fork, (najor tributaries of the Judith River, head in deep open
cuts in the divide and cross the study area west to east.
The Judith
system in its entirety carries all waters north and east of the main
crests.
Jhe southern slope is mostly drained by short deep canyons to the
Musselshell River.
Smith River and Belt Creek are minor drainages
the
study area.
The typing of vegetation used by elk was, influenced by Daubenmire’s
(1'943; 1946; 1952; 1953) work on zonatioh in the Rocky Mountains.
Four
areas designated as major plant communities (Fig. I) were recognized by
the presence or absence of certain conifer species, understory plants,
elevation, and exposure.
Within two of these communities, the Douglas' fir
and spruce-fir, four and three■"types", respectively, related to elk range
use, were recognized and described.
Scientific and common names of plants
used in this paper follow Booth (1950) and Booth and Wright (1959).
Pondbrosa pine woodland - grassland community,;
This community, which
occurs at elevations of about 5,300 feet, does not circle the mountain base
in a continuous belt, as does the adjacent grassland, but comprises local
tracts, the most extensive being in the eastern section of the study area
({Fig. I).
It has been -described by Leiberg (1904) and Lovaas (1958).
'
/
-5-
Jhis community is characterized by limber pine CPinus fle-xilis.) and
ponderosa pine (Pinus ponderosa) growing as scattered trees, thin lines
along ridge tops and patches of forest in a grassland matrix of bluebunch
fescue (Festuca idahoensis) and bluebunch wheatgrass (Aqroovron spicatum)
in various stages of disturbance (Fig. 2).
The trees are character­
istically freely-branched, of low stature, often gnarled and wind-trained.
other plants of this community include creeping juniper
zontalis), shrubby cinquefoil (Pdtentilla fruticosa), fringed sagewort
(Artemisia friaida). pussytoes (Antennaria spp.), Hood's phlox (Phlox
hoodii), draba (Draba
spp.
),
.
rough fescue (Festuca scabreI l a L needle-and- .
thread (Stipa comata)„ bluegrass (Poa spp.). Junegrass (Koeleria cristata).
smpoth brome (Bromus inermis).and crested wheatgrass (Aaropyron desertoruta).
The two latter species were characteristic of abandoned cultivated
fields.
The writings of Weed (1899) and Leiberg (1904) imply ah extensive
ground cover of creeping juniper in the area during an earlier period.
Ob­
servations of numerous dead branches of this plant substantiate their
statements.
Ponderosa pine communitys, This community is very limited and dis­
continuous in distributiono
The only tracts of any size are on the
Musselshell drainage and the lower South Fork of the Judith River (Fig. I).
It has an open, canopy and an abundant uhderstory including bluebunch wheatgrass, bluebunch fescue, big sagebrush (Artemisia tridentata) and arrowleaf balsamroot (BaIsamorrhiza saaittata).
TJie absence of limber pine,
the relatively high density of tall, clear-boled ponderosa pine, and the
different ground level species distinguishes this from the ponderosa pine
t
—6—
Fig. 2.
The ponderosa pine woodland-grassland community in the foreground
with the Douglas fir forest and burn type (right) in background=
Fig. 3.
The open conifer - park type of the Douglas fir (background)$,
with restocking lodgepole pine clear cut (foreground).
-7-
woodland-grassland community.
Douglas fir community*
coverage (Fig. I).
grassland.
TJiis community has, by far, the greatest areal
Along most of its lower elevation it abuts directly on
Its upper boundary is the spruce-fir community, usually at
about 7,200 feet, but the altitudinal range of this as well as the other
communities, fluctuates with changing topography and environmental factors
resulting in what has been described by Daubenmire (1943) as interfingering
or inversion of zones.
Four distinct types, were recognized in this
community.
Douglas fir forest type*
This type is characterized by the presence
of Douglas fir ('Pseudotsuaa mehziasii) (Fig. 2) often in dense stands on
north slopes with all age classes represented, while on xeric south slopes,
a more open scrubby type was observed,
Plants in the understory include
Oregon grape (Berberis reoens). kinnikinnick (Arctostaphvlos uva-ursi).
common juniper (Juniperus communis) and twin-flower (Linnaea borealis).
Lodaeoole pine type*
Much of the Douglas fir forest has been subject­
ed to fires (Leibergi 1904).
Within its range, lodgepole pine usually re­
stocks with dense, 'stands-JkJie
(Picea enqelmanni) forests.
arid.,Engelmaph spruce
Thus lodgepple pine now covers vast acreages.
Under the closed cariOpy formed by dense stands,.understory vegetation is
usually not abundant.
Open conifer - park types. This type includes all open non-timbered
parks and ridges, and the areas of scattered open conifer characteristic
of southern exposures (Fig. 3).
A vast array of grasses and forbs is
present because of the wide range o f .successional -stages in this type.
/ -I,
-8Usually forbs prevail on mesic sites while grasses dominate the drier
siteso
A few of the common plants are bluebunch fescueg yarrow (Achillea
lanulosa). blue bells (Mertehsia spp» )$ shrubby cinquefoilg big sagebrush,
copnon juhiper 9 k innIkinnick 9 limber pine and Douglas fir.
Burn types
In the eastern and southeastern sections of the study
area are extensive tracts of burned forest (Fig. 2).
to be restocking at. a very slow rate.
These areas appear
This typp is characterized by forbs
and browse with scattered scrubby conifers.
A few of the plant species
are arnica (Arnica cordifolia). Canadian buffalo-berry (Shepherdia cana­
densis^. spiraea (Spiraea.s o d . ). common juniper, shrubby cinquefoil, kinnik innick, limber pine and Douglas fir.
On certain sites, probably because
of adverse environmental factors, barren conditions have followed fire.
Spruce - fir community:
This community, characterized by Englemann
spruce and 1 subalpi.ne fir (Abies laslocarpa) ranges from approximately
Y 9200 feet elevation upward to the highest peaks (Fig. I).
states there is no definite upper timber line.
Leiberg (1904)
On north slopes Englemann
spruce, was found somewhat below 7,200 feet and it characteristically ex­
tends to lower elevations down narrow, cool, mesic canyons.
Three types
were recognized in this community.
Spruce forest type:
This type (Fig.. 7) is best represented at the
head of the Middle Fork of the Judith River where it forms dense stands,
choked with fallen timber, on north exposures.
ages of these stands as 300 to 350 years.
of the Englemann spruce forest includes.
Leiberg (1904) gave the
Plants found in the understory
Arnica, marsh valeriana (Varleriana-
d161 ca ). Paysoh's louseworl (Pedicularia pa vsoniana-) and low f ed huckle­
berry CVaccinium scoparium).
Lodaepole pine type;
Lodgepole pine which has restocked the firef
destroyed spruce forest forms an important type in this community and is
logged in certain areas by.the block clear cutting method (Figs. 4, 5). ;
The most conspicuous plant in the understory is low red huckleberry.’ Vege­
tation is discussed in detail under the section on block clear cutting of
..
Iodgepole pine.
Open cortifer - park types. Included in this type are all open parks
(Fig.
6
), most often on southern exposures, ranging from mesic sedge
meadows to xeric ridge tops.
Also in■Vthis type are open scattered conifer
groves (Fig. 7) represented.by limber.pine, subalpine fir and at higher
elevations, whitebark pine (Pinus albicaulis).
Grasslands in this type are
dominated by bluebunch fescue, but forbs predominate on extensive areas.
This type is the most important source of forbs for elk during the summer.
Some forbs and grasses include dandelion ('Taraxacum-s p p .
pale agoseris
(Aqoseris crlauca). geranium (Geranium spp *), northwest cinquefoil (Potentilla gracilis), daisy (Erioeron spp.), bedstraw (GaH u m spp.), brook saxi-'
fraga (Saxifraoa arouta). arrow!eaf groundsel (Senecio triangularis). bluegrasses, mountain brome (Bromus maroinatus). purple onion grass (Melica
spectabiIis). rough fescue and greenleaf fescue (Festuca viridula).
METHODS
Field work occupied full time from June 15 to August S i , .1960 and
April I to September 22, 1961.
The size of the study area made the use of
-10'1
Fig0 4.
Recent block clear cutting of dense lodgepole pine.
Fig, 5,
Block clear cutting within the lodgepole pine type of the
spruce-fir community.
-11-
Fig, 6.
An open ridge top park at 8,150 feet elevation in the spruce-fir
community.
Fig, 7,
The spruce-fir community with open conifer - park type in back- ground (southern exposure) and dense spruce forest in foreground
(northern exposure).
-12-
a rvejiicle necessary, but usually the best observations were made while
stalking on foot.
Elk observed with the aid of 7 x 35 wide angle bin­
oculars were classified, if possible, as calves, cows or bulls.
Other
pertinent data recorded included group size, location, use of vegetative
types, behavior, feeding habits and recognition of marked elk.
Two
hundred fifty*>£tiur elk have been tagged in this area since 1953 (Picton,
1961).
Metal ear tags and in some instances plastic markers and/or
streamers were used.
When feeding elk were observed an effort was made to record plant use.
Plants in the feeding area were carefully examined for use by elk.
Most
instances of plant use were recorded in areas known to have been recently
deserted by elk which were observed at close range.
A few feeding site
examinations were made on areas where elk were observed from great dis­
tances and old use was recorded.
Some pbservations were recorded in areas
where, the presence of elk was recognized by 'Vsign":.
These data were in­
cluded after careful consideration of possible deer or livestock use in
the vicinity.
Following the procedure of Cole and Wilkins (1958) the unit
constituting one instance of plant use was a rooted stem for grass-like
plants and forbs and an individual leader for browse.
A reference plant
collection facilitated identification of plants,
Instances of plant use on a given species were expressed as a per­
centage of total recorded plant use for each ,feeding site»
Percentages
were aggregated and averaged for the period involved ip computing food
habits in relation to seasons and\vegetative types.
The rumens of 41 elk collected in the Little Belt Mountains during
-13-
1955 and 1956 were analyzed by personnel of the Montana Fish and Game De­
partment.
Food habits were computed from the raw data by the author,
using the aggregate percentage method of.Martin, et, al. (1946).
The find­
ings were included ip this study.
Vegetational changes that have resulted from the block clear cutting
method of harvesting lodgepole pipe were considered in relation to elk
use.
For a quantitative expression of ground level vegetative changes>
130, two x five decimeter plots were established in two clear cuts and the
adjacent uncut lodgepole pipe,
Canopy-coverage and frequency were measured
and tabulated in accordance with the method proposed by Daubenmire (T959)>
The centers.of 45 canopy-coverage plots served as points in using the
quadrant method of tree spacing (Cottam and Curtis 9 1956).
considered as the center of four quarters.
The point was
At 25 points in the clear cut,
the distance from the point to the closest tree in each of the four
quarters was recorded by species, height and basal diameter*
At 20 points
in the uncut lodgepole pine forest the distance from the point to the
closest tree, four inches or over in diameter at breast height (dbh), in
each quarter was recorded by species and dbh.
Canopy cover and height
also were estimated in the uncut forest.
RANGE USE AND MOVEMENT?
A. total of 2,598 elk in 326 groups was recorded during
tional trips in the study area.
292
observer
Approximate counts of the largest observed
groups on winter and summer range were 90 and 56 elk, respectively,.
(I960) reported a maximum of 119 elk seen in one summering group,
Picton
The
-14-
number of elk observed during April almost equals all elk observed during
the remainder of the study (Table I).
Table I.
Fewer elk were seen in August (both
Percentage distribution of observed elk on designated vegetative
types by periods, 1960 and 1961.
Doualas fir community
CL
•a
A
re
i—l
<u
S
O
Mn
Mn
O -P
°!
® Q)
I S0i)
as O
SJ2
SB O
30
16
18
17
35
75
38
36
27
1205
252
175
50
268
409
Period
April
May
1-15
16-31
June 1-15
15-30
July
Aug.
1-15
16-31
Sent. 1-22
Totals
292
Q) "u)
C 0)
0)
<o T
a
S
O
p
C
C
Ij^ I
II I
99
74
53
•P
0)
MOh
M
Mh
Cfi
r-H
D
S
V
U
Q)
C
a
<u
O
Q>
-a
S
*1 /
2598
S
a
9
2
8
23
66
46
127
5
11
57
I
*
2
i!
O
19
30
46
17
2
*
Q)
•H
O
*
6
P
CD
t
Burr
U)
Spruce -fir community
U
a
C
CL
0)
O
O
Q)
-o
3
I
CD
•H
C
O
o
r*
§
s-
I
2
13
24
13
I
23
3
4
20
8
11
50
55
11
22
3
41
52
41
77
17
11
15
6
9
<D
3
2
l/ Less than one per cent.
1960 and 1961) than in any other month, although the greatest number of
observation trips were in August 1961.
Table I shows the percentage dis­
tribution of observed elk on designated vegetative types by periods.
About 99 per cent of the 1,205 elk observed during 30 observational
trips in April were confined to the ponderosa pine woodland - grassland
community.
All but three trips during the month were in this community in
-1 5 -
the foothills In the general vicinity of the Judith Game Range (fig. I),
On ,April 7 no sign of elk was evident in the vicinity of Russian Greek at
an elevation of about 6,500 feet in the Douglas fir community.
The tracks
of only three elk were seen along Ettien ridge east of Ettien Springs at an
elevation of about 6,000 feet in the Douglas fir community on April 15.
In the vicinity of High Spring at an elevation of about 6,000 feet in the
Douglas fir community, no fresh elk sign could be found on April 17=
From
this vantage point the lodgepole clear, cuts along the main divide east of
Hoover Mountain (Fig, I ) at about 7,200 feet elevation were observed to be
still deeply covered with snow.
. ' .
During April elk Used grassland openings as feeding sites.
areas were used for bedding, cover and escape.
the observed daily behavior pattern.
w
Timbered
This use is reflected in
At about 4:00 p.m., elk were usually
observed moving from the" timber cover to a particular grassland opening.
During early April a preference for southern exposures Was noted.
intensive feeding was just before darkness.
The most
During early morning elk were
usually observed gradually moving towards timber cover and by about 7:00
a.m. feeding activity had greatly decreased.
Although elk were concentrat­
ed in a relatively small area on their winter range, observations suggested
thaf. individual groups of varying sizes maintained autonomous behavior.
Range use and movements were somewhat modified between groups because of
this behavior.
Deer and elk were found together on feeding sites and while
elk
used
both native and cultivated grasslands, large numbers of deer were usually
found only in the cultivated (brome) meadows.
V
Throughout fhe study deer
=16"
were often foi^nd with elkb
In late April the winter range was still .receiving heavy elk Use=
On
April 24, 23.9 elk were pbserved in the ponderosa pine woodland - grassland
communityo
By the latter part of May elk use on the Judith .Same Range had greatly
decreased and a nptice@h.le-niovement "from,-, the-dry grassland ranges into'the
mountains had begun,
Picton (19.60). reported migrations ffdm the winter
range in Apfil and May=
Early in the month the open parks of the Douglas
fir community (fig = 3) were being used at. Russian Flats and by May 24, elk
were in the open parks along the South Fork of the Judith River in the
vicinity of their summer range above Russian Creek (.Fig= I) =
On May 25 no signs of elk or deer were evident in the vicinity of the
lodgepole clear cuts in the spruce-fir community along Spring Creek road
(Fig, I). . This may have been due to the retarded development of vegetation
at this elevation=
pf Bpring Greek
During May 9 elk were observed along the southern parts
road,
using open parks In the Douglas fir community,
A;
.
!
trip into Dry Pole Canypn (Fig= I) at the. end of the month revealed small
groups and single animals widely scattered over the upper part of the
canyon as high as the spruce,-fir community, with most df the animals high
on the past wall Using thp. Douglas fir community, burn type =
abundant in Dry Pole Canyon throughout the summer=
Water was
Some calves were ob­
served with the widely dispersed cows in the burn type in. June =
cates that the Douglas fir community was. used during calving=
(T951) gives thp peak pf births, in thp Gallatin herd as June. 1=
This-indi­
Jdhnspn
Pidtpn
(1960) found that the majority of .elk observed during the calving season,
-17-
iti June, were -using the grassland areas at lower elevations.
After the.
middle of June the grouping of cows with calves steadily increased.
The increased Use of forbs early in June (see food habits) is re­
flected in the use of types favoring these plants.
<At this season the
open conifer - park and burn types of the Douglas fir community contained
succulent forbs and received heavy elk use as shown in Table I.
Dry areas
which lacked succulent forbs in the. DoUglas fir community (see area de­
scription) were not used by elk during the summer season.
After the middle
of June a decreasing use of the Douglas fir ,community was matched by an in-f
creasing use of the spruce-fir community, particularly in the open conifer
- park type (Table I).
During the latter part of June the open conifer -
park type of the spruce-fir community at the drainage heads of the Judith
River became important elk concentration points (Fig. I),
A parallelism ■
was noted in Piston's (i960) observations during the same period.
He
stated that elk moved through the forest types in late June and early July
and by mid July were along the upper edges of the subalpine forest and in
the subalpine barrens where the forb subtype was favored.
The sites used by elk in 1960 were similarly located in 1961«
In the
evening elk moved up out of the drainsge bottoms to the open ridge tops
(Pig.
6
) w/here succulent forbs, were found at this season-
These herds
were often observed until after darkness when most activity and feeding
occurred.
Elk were again observed in these same localities at day break,
Bed^grounds were observed on top of ridges indicating that at least some
bedded down during the night-
Early in the morning the elk moved gradually
down the drainage towards the■timber=
This behavior appeared to be Of
common occurrence and was also observed in the drainages of the lodgepole
pine clear cuts (to be discussed later).
By the end of Juhe 9 elk were distributed to ail parts of their summer
range and were found at elevations up to approximately B 9OOO feet.
During July elk were concentrated in the spruce-fir community at
drainage heads located along the main divides ((Fig^xI), The dp'eh cdriifer I
■
,
park type was most heavily used (’T able 'l),» .The .largest -summering group 9
56 elk, was observed on July B 9 ,1961 in the spruce-fir commuhity.
The use of open ridge tops decreased during July and although there
was an upward movement of elk in the evening, the animals usually stayed
within the drainage rather than on the ridge tops.
The decreased use of
ridge tops may have been due to loss of succulence in the vegetation, or
to the use of these ridges by domestic sheep at that time.
During the summer months elk,appeared to favor habitats closely
associated with water,
Generaliy the most m.esic sites were located within
the spruce-fir community, and withift the study area the mesic drainage
heads along the north and east slopes of the main divides (:Eig^ hi) appear­
ed to be ..favored by elk.
Marked elk observed during JUne and July showed that movement from
the winter range was considerable-.
On June 13, 1961 a tagged cow elk was
observed on the southern slope of the main divide, approximately
miles from the Judith.Game Range headquarters,
20
airline
(This does not imply that
the cow spent the previous winter on the game range.)
Picton (1960) re­
ported a maximum movement of 28 airline miles from the winter range as
-1.9-
shown hy two tag returns 0
Five marked elk were, observed at 1 about 5,300
feet elevation in April, while late in July 9 1961 a tagged cow was seen at
8,300 feeto
The locations of all 16 marked elk observed during the study
are plotted (Tig. I ).
In August elk withdrew to their host secure habitat and many extensive:;.'.;
trips during the month revealed few animals.
There, is some suggestion from
limited observations of elk that drainage bottoms with water and spruce
forest or other forest cover on north exposures are used during this
period (Table I).
.Fresh elk signs observed at the sites of succulent
vegetation within the above1described general area substantiated these ob­
servations.
Picton (i960) reported that in late August there was a move­
ment downward into the forested areas.
fhe first rutting activity by bulls was noted September 5, 1961, a n d .
j
this greatly increased after the middle of September.
"Raghorns"? and
spikes which had summered with the cows and calves were driven out by
mature bulls.
All elk became increasingly observable during the month,
and older bulls seldom seep during the spring and summer were easily ob­
served, particularly in clear cuts.
Table I shows that most elk observed
during September were using the lodgepoie pipe type within the spruce-fir
community.
Thia usage included clear cuts dind is discussed under elk re­
lationships to clear cuts.
This is. npt at a great variance with Pictonls
(i960) observation of all sexually active males being seen in the forest
types.
All elk observed between June 15 and August 31, for both summers were
in either the DOuglas fir or spruce-fir communities, but usage, was not
/
—2 0 - r
equal between years,
In I960, forty-four per cent of the observed elk
were in the Douglas fir and 56 per cent in the spruce-fir communities,
while 'In 1-961 the figures were 13 and 87 per cent respectively.
The much
heavier use of the spruce-fir community in 1961 may have been due to
drought conditions and the resulting effects .on forhs,
Powever, in I9 6 0 .
the locations of elk were not well known and the whole study area was
generally covered.
community.
About an equal amount of time was spent in each
In 1961 I looked for elk where they would likely be found and
i
about 90 per cent of the observations were in the spruc.e^fir community,
Table I shows a lower percentage of elk observations in. timber cover
than open areas,-
This was probably due to the reduced visibility, in
timber, the fact that elk were not usually sought out in timber, and ob-.
nervation periods frequently paralleled feeding periods when open areas
were used by elk,
ELK RELATIONSHIPS TO BLOCK CLEAR. CUTTING OF LODGEPOLE PINE
Much of the study area along the divide in the upper portion of the
South Fork of the Judith River supports a fully stocked closed canopy
lodgepole pine forest.
Parts of this closed canopy have been broken by
the block clear cutting method of harvesting IodgepoLe pi-he (Bigj '
:8 )>
The status of block clear cuts in the. Deadhorse Sale Area (Fig, l)
where most elk observations in clear cuts were made, ,is summarized in
Table II which was compiled from the U. S„ Forest Service, Timber Survey
Map of the area.
Some lodgepole pine cuts were on south slopes in the
Douglas fir community, while others were on north slopes in the spruce-fir
-21-
Fig „
8
.
Clear cut blocks of lodgepole pine in the Deadhorse Sale Area 0
(Location of vegetative measurements discussed in the text
designated by X»symbols)«,
(U 0 S 0 For, Ser 0 Aerial Photo)
-22-
Table II.
Physical characteristics of block clear cuts and a summary of
elk observations in the Deadhorse Sale Area.
South slope
of divide
Block clear cuts
Number
36
Size range (acres)
10-55
Av g . size (acres)
23.65
851.4
Total acres cut
Cutting dates (range)
7/51-6/60
6
No. of cuts in each 1951
4
section by year i/ 1952
1953
1954
2 ( 1 )2 /
4
1955
1956 1 0 (2 )
4
1957
2
1958
0
1959
1960
I
1961
0
North slope of divide
East of Spr. West of Spr.
Cr. rd. less Cr. rd. more
mesic
mesic
39
5.1-44.3
23.52
917.3
6/54-11/60
0
24
10-38.5
2 2 .2 2
533.2
9/51-8/60
3 (9)
0
2
0
(1 2 )
6 (13)
I (4)
2
(I)
2
(I)
1 1 (I)
9 (I)
10
0
0
0
0
I
I
0
0
Elk observations (95 observation periods, 1960-61)
No. groups in clear cuts
3
3
40
No. groups not in clear
2
cuts
I
13
Total no. groups
4
5
53
No. elk in clear cuts
5
6
279
No. elk not in clear
2
cuts
I
96
Total elk
6
8
375
No. of clear cuts
used by elk
3
3
16
Year of cutting unknown for
6
99
5.1-55
23.25
2301.9
7/51-J1/60
9 (9)
6
8
2
Totals
(1 2 )
10 (14)
15 (5)
23 (4)
13 (I)
4
10
0
(I)
3
(I)
0
46
16
62
290
99
389
22
block clear cuts.
2/ Figure in parentheses indicates number of elk groups observed in clear
cuts of this age.
—2 3 —
community-
The clear cuts on north slopes located west of Spring Creek
road (Fig* I) were more often associated with the mesic conditions of
drainage heads than were those ioceted east of this road.
More elk were observed in clear cuts than in uncut portions of the
Deadhorse Sale area (Table II).
This was probably influenced by reduced
visibility in the close-set JrOdgepole pine timber (-Fig, 4), or by the time
of observation, which coincided with elk feeding activity-.
appear that clear cuts were used largely for feeding.
It would
About SOOper dent
of all elk groups using clear cuts in the Deadhorse Sale Area were feeding
when first observed as compared to 14 per cent in the uncut portion..
Ex­
cept for. September, elk were seldom observed in clear cuts during midday.
The groups of clear cuts associated with mesic drainage heads on
north slopes received, by far, the greatest usqge.
Table TI shows that 22
clear cuts were used by elk, but only in nine of these were two or more
groups observed.
All nine were in that group of clear cuts associated with
mesic conditions of drainage heads.
The clear cut in which the greatest
number of elk groups were observed is bisected by a mesic drainage.
Cutting dates varied from July 1951 to November 1960.
The 1953 and
1954 cuts, approximately eight and seven years old respectively, received
57 per cent of the total usage of all clear cuts by ellc groups, despite
the fact that they comprised only about 22 per cent of the total.
Age of a
cut does not fully explain elk use, however, as different clear cuts of the
same age did not receive equal use even in a local areg,
The cuts varied in size from 5.1 to 55 acres (Table Il)..
observed in cuts ranging in size from 12.3 to 38.5 acres.
Elk were
Seventy-four
-24-
p^x cent of aj.1 elk groups observed in clear cuts were using cuts in the
15.2 to 25.2 gpre size class (Table III),
of the total clear cuts.
These cuts comprised 57 per cent
Approximately 50 per cent of the clear cuts used .
by elk were duplicated in size by cu^s in which elk were not observed, •
Table III.
ElK use in relation tc3 size of clear cuts.
’•
'
Size ranae of clear cuts (acres)
5.1-15.1
No. clear
cuts
Per cent of
clear cuts
No. elk
groups
Per cent of
'61k groups
15,2-25.2
25.3-35.3
56
U
35.4-45.4
45.5-55.5
4
27
Total- '
I
99
4
I
. IOQ
'
'
57
ii
•I
2
27 ■
'
■
34
7
4-
0
.
74
T5 .
9
0,
46'
100
"
Seasonal use of the clear cuts in the Deadhorse Sale Area is shown inr
Table IV.■ No elk were observed, in this area during May, but 96 were ob-r.
served at lower elevations south and north of the area.
■
On- June 7 new ■'
'
.
•
green vegetation was noted in the clear cut areas and the first elk tracks,
but no elk or deer were observed.
horse Area. June 22.
The first elk was observed in the DeaJ-
Elk use on adjoining areas declined.
Elk numbers in .
clear cuts reached a peak, in July, dropped to a minimum in late August and
roee again in September.
Use in the later month is better compared to July
use if both elk numbers' and groups are considered.
Clear cuts in this area are commonly used by deer and cattle.
Whereas
elk used the mpre mesic sites, deer tended to heavily use %eric clear cuts
of the north and south slopes.
Cattle usage increased, especially at high-
-25-
Table IV.
Elk and elk groups observed during I960 and 1961 in block
clear cuts of the Deadhorse Sale Area.
• T T - V - ...
No. of
elk
observed
f. , t-,..
April
May
June
1-15
Jiine 16-30
July
1-15
July 16^31
Aug,
l-i5
Aug. .16-31
Sepi. 1-15
Sept.. 16-22.
Totals
0
' No. of
elk
groups
observed
.
0
■
0
Elk
observed
per
trip ■
Nd. of
obser­
vational
trips
'Elk
''
groups
observed.
, per trip.. .
0
0
0
0
0
0
0
0
3
3
0
0
7
8
2 .8 8
6
13
14
17
16
8.08
23
105
28
27
7
7
12
2
4
2 .0 0
12
31
- ...64
.
.. .13 ..
. 290
.
46
. 9 .... .
..
95 .
er elevations, as the summer progressed.
1.59
0.75
2.58
7,11.
.8 8
.46
.50
.41
.13
.33
.. . . •. li.4.4
3.05
...
.48
.
■
:
The clear cut in which the most,
elk groups were recorded was also used by cattle.
Ground vegetation in both clear $nd uncut lodgepole was analyzed by
methods previously described.
The results of these methods, employed in
clear cuts heavily used by elk and in adjacent uncut
lodgepole pine timber (Fig.
8
1 2 0 -2 0 0
), are tabulated in Table V.
year old
This table shows
en increase in canopy-coverage and number of species recorded for ground
level vegetation in seven and eight year old clear cuts as compared to ad­
joining uncut lodgepole pine sites.
It should' be noted that canopy cover­
age and frequency listed in Table V. are not necessarily related as the two
cap vary independently.
The table shows a great increase in forbs and
grasses in clear cut areas, as compared to uncut areas, but a sharp de­
crease in low red huckleberry (Vaccjnium scoparium). thin leaf huckleberry '
Table V.
Physical, quadrant tree and canopy-coverage data for two clear cuts and three adjacent
uncut lodgepole pine sites on the north slope west of Spring Creek Road, Deadhorse Sale
Area, September 15, 19 and 20, 1961.
Lodaeoole nine
clear
cut site
North
7200
22.7
6-53
11-53
50-75
North
7200
2W
5/40
11/80
5/15
—
2/
site _B
site C
Lodgepole pine
uncut
clear
B & C sites cut
combined
site A
25
8-54
11-54
80-95
East
7200
East
7200
East
7200
249
105.1
98
751
147.6
8 .8
6
60 ft.
50 ft.
2064
2.9
92
0.5
16.1 in.
75
80-90
10
20
20
20
12/50
20/50
18/50
6/55
17/60
12/40
4/40
3/5
1/5
1/5
2 /2 0
2 /2 0
6/35
100
1 /1 0
6/15
—
40
18/53
15/45
5/48
2/5
4 / 5 2/
3/8
50
9/36
9/46
8/46
6/32
10/28
4/18
■26-
Physical data:
Clear cuts
Size (acres)
Date cut
Date burned (slash)
Est. floor litter {%)
Exposure
Elevation (feet)
Quadrant tree data:
Density (trees per acre)
Basal stem area (sq.ft./acre)
Relative freq., lodgepole
Avg. tree dia. (inches)
Tree height (est, in uncut)
Canopy-coverage (est. % closed
for uncut sites)
Canopy-coverage data: I/
Numbers of plots
Vaccinium scoparium
Vaccinium membranaceum
Spiraea spp.
Linnaea borealis
Berberis repens
Pinus contorta
Astragalus spp.
uncut
site
Uncut lodgepole
pine
Table V,
continued.
Lodqeoole Dine
uncut
site
Aster spp.
Carex qeyeri
Lupinus spp.
Eoilobium anaustifolium
Trisetum spicatum
Poa. spp.
Festuca idahoensis
Calamaorostis rubescens
Elk droppings
— ~=—
—
—
—
—
—
clear
cut site
— —
26/75
19/55
11/95
5/35
6/45
— —
—
—
— —
/o
/15
Uncut lodgepole
oine
site B
+/15
15/25
—
site C
1/20
4/10
—
—
—
—
Lodoeoole oine
uncut
clear
B & C sites cut
combined
site A
1/18
9/18
«—
—
—
—
—
—
==«■»
8/14
—
—
/15
9/32
25/66
9/28
5/26
4/18
■■ nm
/10
/15
6/10
A
-I
Excluded from the table were six species that occurred in plots on uncut lodgepole pine and 15
on clear cut sites which had less than five per cent average canopy-coverage at all sites.
2/
The figure preceding each / is average per cent canopy-coverage for all plots on the site;
following figure is per cent frequency (coverage the basis of inclusion) in the same plots.
3/
The symbol-fp- indicates less than 0.5 per cent.
—2 8 —
(Vaccinlum membrahaceum) and twin flower .(Linnaea borealis).
(Spiraea
sop.
)
Spiraea
increased in one clear cut but decreased in the other. . The
increase in elk sedge (Carex qeveri) in clear cuts is noticeable.
The high
elk sedge coverage on uncut site .’IB” is explained by-the open nature, of
the tree canopy.
It was present in five of the last six plots .which were
under, a more open tree canopy.*
Table "V shows that the clear cuts had a
greater ground-vegetation canopy-coverage of the shade intolerant,lodgepole pine (Pinus contort a ) re.stocljage than did the uncut areas.
The two
'
most important forbs, as far as coverage and frequency are concerned,
lupine (lupinus spp,) and fireweed (Epilobium- anaustifolium). appeared in
clear cuts but not in uncut lodgepole plots.
The. average canopy-coverage
of aster (Aster spp.) was nine per cent in clear cut ”A ” .
section for elk usage of these forbs).
(See food habits
'Grass did not appear in any plot ;
-under,dense lodgepole pine timber but amounted to considerable coverage in.
clear cuts,
A comparison' of the. tree quadrant data is interesting as concerns t h e ’
,
number, of trees per acre,,basal stem area and canopyrcpverage.
lodgepole pine with an average height of 16.1 Inchps^ for.
measured, and a density of over
2 ,0 0 0
100
Restocking ••
trees,
trees per acre, has a canopy-coverage
of 10 per cent, as compared to thp estimated coverage pf .75 to 90 per cent
in mature uncut lodgepole pine*.
The heterogeneous nature of the vegetative cover of clear cuts is
illustrated as follows;,
only in seven consecutive plots falling along a
logging road in clear cut ”A" were bluebunch fescue (.Festuca idahoensls)
-29“
and redtop fAarostis alba) recorded, but the oanopyrcoverage and frequency
of these seven plots was 57/100 and 12/71 per cent for bluebunch fescue
and redtop respectively.
Combining these data in all 50 plots of !'Aw
masks the concentration that.did occur,
Daubenmire (1959) states that 40
plots may be satisfactory, but this should not be accepted everywhere with™
put tests.
The mnny varied disturbances within a clear cut make it any­
thing but homogeneous,
A 1 considerable number of plots appeared to be
necessary in most clear cuts to obtain an adequate sampling of the vegeta™
tion,
Fewer plots gave an adequate sample under the uncut lodgepole pine
timber.
Table V shows that coverage and frequency of low red huckleberry
at all three sites in the uncut lodgepole pine have rather close values,
As sites !-eB!';and wC". were similar, data from the forty plots were combined.
The resulting eanopy^coverage and frequency gf low red huckleberry was
ll/|3 per cent 9
if canopy-ogverage of the ground vegetation is used as a comparison
between yncut mature, lodgepole and clear cuts on similar sites, the super™
igfity of clear guts is evident, as shown in fable v.
This could well ex™
plain the. increased usage of clear cuts as feeding sites by elk,
y
.
•
ELK FOOD HABIfS
.Food habits
were
determined from 12,008 instances pf plant use
re=
corded during the examinations of 95 elk feeding sites involving approxi­
mately 969 elk.
Table VI shows both plant and forage-class usage by
seasons and range types.
Greater meaning can be attached to, total in=
stances of plant use if the number.of feeding sites contributing to that
T a b l e VI.
Elk
f o o d h a b i t s a s d e t e r m i n e d f r o m 1 2 , 0 0 8 i n s t a n c e s of p l a n t u s e a t 9 5 e l k f e e d i n g sites.
S e a s o n a l
S p r i n g
1 9 6 1
f e e d i n g
s i t e
e x a m s
P e r
N u m b e r
i n s t a n c e s
P l a n t
u s a g e
T a r a x a c u m
‘ L u p i n u s
A s t e r
I /
o f
.
s d d
s p p .
s p p .
A a o s e r i s
a l a u c a
A a o s e r i s
s d d
E o i l o b i u m
.
a n a u s t i f o l i u m
P o t e n t i l l a
a r a c i l i s
P o t e n t i l l a
s p p .
F a l l
1 9 6 1
1 9 6 1
R a n g e
S u m m e r
J u n e
c e n t
s i t e s
w h e r e
u s e d
S e p t .
3 5
1 3
1 6
-
P o n d e r o s a
J u l y
3 1
1 9 6 1
2 6
O p e n
p i n e
p a r k
D o u g l a s
S p r u c e -
g r a s s l a n d
f i r
f i r
2 6
c o m .
c o m .
1 3
3 2
s i t e s
s i t e s
s i t e s
%
%
%
%
%
%
%
%
%
u s e
—
1 5
I
u s e
u s e
4 4
2
i
2 9
5
4
—
u s e
u s e
u s e
u s e
4 0
4 0
2
I
—
2
I
7
—
T r .
5
—
5
5
—
2
—
4
—
I
—
1 7 3
8
—
I
—
3
—
3
—
T r .
— —
—
—
— —
3
—
—
—
3
4
—
— —
T r .
—
4
—
—
3
4
—
T r .
2
I
5 8
2
— —
2
1 0 4
7
— —
3
—
—
T r .
I
I
3
—
—
—
3
—
3
8 9
3
—
3
—
9 6
1 5
—
3
—
4 3
I
—
—
I
4
1 7 5
3 6
2
2
T r .
5
5 1 8
9
1 4
T r .
■ —
_
3 6 5
8
2
T r .
—
2 2
—
—
2
—
—
— —
— —
7
—
I
3
6
— —
F e s t u c a
i d a h o e n s i s
—
-
2
c r i s t a t u m
1 8 3
3
6
A a r o o v r o n
s o i c a t u m
1 6 2
3
3
—
—
A a r o o v r o n
d a s v s t a c h v u m
7 0
I
3
—
— —
—
—
—
—
—
3
—
—
T r .
6
P o a
o r a t e n s i s
4 8
I
P g a
s p p .
9 2
1 2
s o i c a t u m
U n i d e n t i f i e d
C a r e x
a e v e r i
S a l i x
s p p .
S p i r a e a
s d d
.
g r a s s
—
I
—
I
T r .
3
—
2
2
1 2
4
7 4
3
3 9 8 2
6 3
9 0
3
—
1 6 6
5
—
I
8
—
1 1 7
1 2
—
3
I
T r .
5 6
I
—
—
— —
—
A a r o o v r o n
T r i s e t u m
—
—
T r .
s o p .
i n e r m i s
i
i
v i s c o s s i s s i m u m
B r o m u s
—
5
—
2
i
4
—
G e r a n i u m
f o r b s
2 3
8
—
— —
3
1 0
—
1 2
—
1 7
—
T r .
——
5
2
9
3
1 0
—
3
—
—
—
—
4
—
i
—
2
—
4
T r .
-
I
2
— —
—
—
—
—
—
I
8
—
3
_
u s e
4 6
1 2 8
3
— —
—
u s e
2 0 3
2
2
3 3
G e r a n i u m
s u l o h u r e s c e n s
1 8
s i t e s
2 2
1 0 7
c u t s
s i t e s
2 9 1
s o l e n d e n s
b l o c k
c l e a r
s i t e s
2 3 1
C k v t r o o i s
L o d g e p o l e
p i n e
s i t e s
—
8
U n i d e n t i f i e d
t y p e
w o o d l a n d -
s i t e s
1 5
1 3
H e d v s a r u m
u s e
c o n i f e r -
s i t e s
4 4
6 8
s p p .
A u g .
3 6 8
1 1 1
A r n i c a
M a y
2 5 7 9
f r i o i d a
c o r d i f o l i a
J u l y
3 1
1 5
1 9 6 0
A p r i l
f e e d i n g
A r t e m i s i a
A r n i c a
( f o r b s )
u s e
o f
t r a c e - less than 0 . 5 p e r cent)
u s e
S u m m e r
J u n e
1 5
(Tr. =
7
—
I
—
T r .
4
—
I
— —
T r .
2
5 8
1 6
5
—
—
—
I
4
T r .
4
— —
— —
—
—
I
—
3
2
T r .
7
T a b l e VI.
(Cont i n u e d )
S e a s o n a l
S p r i n g
F a l l
1 9 6 1
1 9 6 1
1 9 6 1
R a n g e
u s e
S u m m e r
S u m m e r
J u n e
A p r i l
J u l y
M a y
A u g .
S e p t .
3 5
1 3
3 1
c l a s s
A g g r e g a t e
u s a g e
%
2 /
u s e
p l a n t s
i n s t a n c e s
o f
p l a n t s
P l a n t s
t h a t
c o n s t i t u t e d
l i s t e d .
G l a b r a .
P l a n t s
A s t r a g a l u s
l i n e a r i s .
t h a n
3
f r o m
t h e
s p p . ,
a r l s t a t a .
n u t a n s .
a c r i f o r m i s ,
T r a q o p o q o n
s p p . ,
A q r o s t i s
v e l u t i n u s .
i d a e u s ,
p e r c e n t a g e s
n u m b e r
s i t e s .
2 /
A l l
5 Z
N u m b e r s
p l a n t
i n
s p p . ,
a l b a ,
f i r
2 6
L o d g e p o l e
'
f i r
b l o c k
c l e a r
c o m .
c o m .
1 3
3 2
2 6
p i n e
c u t s
1 8
s i t e s
s i t e s
s i t e s
s i t e s
s i t e s
s i t e s
%
%
%
%
%
%
%
%
u s e
u s e
u s e
u s e
u s e
u s e
u s e
u s e
8 6
5 1
8 1
8 7
8
6 3
8 2
6 5
6
4 0
1 9
5
9 2
3 5
2 1
I
7
1 0
I
8
T r .
2
1 3
4
3 5 6 3
4 2 0
1 3 2 8
2 8 5 7
2 3 8
9 5 4
2 8 9 6
1 2 5 3
( 8 6 )
( 3 8 )
( 8 5 )
( 9 0 )
( 5 )
( 6 1 )
( 8 5 )
( 6 1 )
2 2 5
1 0 4
4 3 8 1
5 8 5
( 1 4 )
( 3 )
( 3 7 )
9
2 0 2
( 9 5 )
7
( I )
( 6 )
( T r . )
5 7 3
( 9 3 )
( 5 1 )
3 7
( 7 )
3 1 4
1 2 5
( I )
( 8 )
( H )
J u n i p e r u s
p e r c e n t a g e s
p a r e n t h e s e s
-
t h e
a n d
a r e
p e r
B r o m u s
V a c e i n i u m
i n s t a n c e s
o f
o f
p e r c e n t a g e s
u s e
u s e
P h l o x
V i c i a
S t l o a
F o t e n t i l l a
a r e
e a c h
1 4
f r o m
i n
P s e u d o t s u q a
i n
t o t a l
s p p . ,
r e l a t i o n
S e n e c i o
i n t e r m e d i a .
t o
s p p . ,
C a r e x
F a m . ,
m e n z i e s i i ,
Z y q a d e n u s
s p p . ,
R i b e s
5 1 1
( H )
1 4 5
( 2 5 )
( 4 )
( 1 4 )
1 5
s p p . ,
L i l i a c e a e
d o u g l a s i i .
2 9 7
t o t a l
s p p . ,
a r v e n s e ,
E r i g e r o n
P o l y g o n u m
g l a u c a .
t h a n
A n e m o n e
C i r s i u m
i n t e g e r r i m u s ,
E l y m u s
F a m . ,
s p p . ,
L e g u m i n o s a e
b i s t o r t o i d e s .
Z y g a d e n u s
C y p e r a c e a e
E q u i s e t u n
p l a n t ,
s p p . ,
o r ■ l e s s
u r s i n u s ,
3 6 9
( 2 )
m a r g a r i t a c e a .
C i r s i t m
E r i g e r o n
a l b i f l o r u m .
P o l y g o n u m
D a n t h o n i a
f r u t i c o s a ,
s p p . ,
3 3
c a t e g o r i e s ,
A n a c h a l i s
s p p . ,
i n d e c o r u s .
V i o l a
C o l u m b i a n a .
c o n s i d e r e d
c a l c u l a t e d
s p p . ,
S e n e c i o
s p p . ,
m e m b r a n a c e u m .
f o r
H i e r a c l u m
r a n g e
a n d
s p p . ,
C a s t l l l e i a
D o d e c a t h e o n
s p p . ,
c a n u s .
s e a s o n a l
A c h i l l e a
s a g i t t a t a .
i n a r g i n a t u s .
h v s t r i x .
h o r i z o n t a i l s ,
c e n t
F a m . ,
w e r e :
b i c o l o r .
H e u c h e r a
S e n e c i o
U n b e l l i f e r a e
s p p . ,
e l k
g a i r d n e r i .
s p p . ,
s p p . ,
i n d i v i d u a l
a l l
b y
D e l p h i n i u m
t r i f l o r u m .
S i t a n i o n
i n
u s e d
B a l s a m o r r h i z a
F a m .,
S e d u m
2 /
2 8 6
u s a g e
b u t
P t e r i d e r i d i a
B r o m u s
s p p . ,
S v m p h o r i c a r p o s
A g g r e g a t e
o f
s e r i c e a ,
P h l e u m
m i s e r ,
G e u m
r h o m b o i d e a ,
T r i f o l i u m
c r i s t a t a .
C e a n o t h u s
s p p . ,
c e n t
t a b l e
C r u c i f e r a e
G a l i u m
C x y t r o p i s
p e r
A s t r a g a l u s
F a m . ,
S a x i f r a q a
t r a c h y c a u l u m ,
K o e l e r i a
l e s s
e x c l u d e d
C o m p o s i t a e
G a i l l a r d i a
2 /
g r a s s l a n d
t y p e
S p r u c e -
s i t e s
4 8 1 5
B r o w s e
R u b u s
D o u g l a s
1 1
3 1 3
( 6 )
G r a s s - l i k e
s e r i s
1 6
p a r k
w o o d l a n d -
u s e
F o r b s
n o t
p i n e
3 1
1 9 6 1
8 8
B r o w s e
l /
J u l y
u s e
c o n i f e r -
p e r c e n t a g e
F o r b s
G r a s s - l i k e
T o t a l
2 /
■-
O p e n
s i t e s
s i t e s
F o r a g e
1 5
1 9 6 0
J u n e
P o n d e r o s a
i n s t a n c e s
A n e m o n e
C l a y t o n i a
E r i o g o n u m
F a m .,
R a n u n c u l u s
m i s s o u r i e n s i s .
e l e g a n s ,
A q r o p y r o n
F e s t u c a
s c a b r e l l a .
A r c t o s t a p h y l o s
v i s c o s s i s s i m u n .
s p p . ,
o f
s p p . ,
R o s a e e a e
R o s a
C o l l o m i a
F r a g a r i a
s p p . .
p l a n t s
u s e d
f o r
e a c h
s i t e ,
w e r e
a d d e d ,
t h e n
u s e
a t
a l l
f e e d i n g
s i t e s
f o r
t h e
p e r i o d
i n d i c a t e d .
d i v i d e d
s p p . ,
M i c r o -
R a n u n c u l u s
s p p . ,
h o o k e r ! ,
r e p e n s ,
s p p . ,
M u s h r o o m s .
a l l
a r e
A r a b i s
A q r o p y r o n
B e r b e r i s
F a m . ,
u s e ,
T h a l i c t r u m
H e l i c t o t r i c h o n
u v a - u r s i ,
c o m p u t i n g .
i n s t a n c e s
s p p . ,
s p p . ,
H e r t e n s i a
S o l i d a g o
o f
p a t e n s .
b y
t h e
total are known-
The second column of the tabla expresses this number as
a per cent of the 95 feeding sites.
Spring (April and May, 1961);
During April and May 5,165 instances.;
of plant use were recorded at 31 elk feeding sites.
plants constituted
88
per cent of the total.
Use of grass-like
Evaluation of use by species
had little meaning when 50 per cent or more of the usage was' recorded in
the unidentified grass category (Table VI).
New green grasses were often.
closely grazed making identification difficult.
About 70 pier cent of all
grass usage, recorded during the study for all seasons, was in the un­
identified grass category.
Field observations suggested the bluegras.ses
(Poa spp.), bluebunch fescue (Festuca, idahoensi-s). and rough fescue (Festuca scabrella) were heavily used.
It appeared that elk avoided the use
of dry grass during spring when green growth was available.
New growth
intermingled with dry grass usually was not as readily used as was the
green grass alone.
When new growth was scarce or covered by snow, usage .
of dry grass increased.
The limited availability of forbs during spring is reflected in the Y
low use percentage.
One of the earliest to Vgreen-Up", fringed sagewort
(Artemisia friaida). was. heavily used for only a short period in early
April.
False dandelion CAgoseris spp.) was the only other forb used in
appreciable amounts.
The first recorded Use of succulent forbs was May 16
with false dandelion receiving heavy use.
The first recorded use of
succulent forbs did not mark a. change, over, from grass-like plants to forbs
as grass continued to be the dominant plant in the diet during the remain­
=33”
der of the periods
Dandelion ft.Taraxa.cum spp0 ) was first observed in. bloom
during the middle of May 0
The first observed use of dandelion was on May
30 0
Summer (Junes July and August 9 1961).a
succulent forbs was apparent*
About June I increased use of
Dandelion was the most heavily used forb
during the summer months when forbs accounted for
plant usage.
86
per cent of the total
Use on this plant was recorded at 41 of 51 feeding sites ex=
amined during Juhe 9 July and August of both 1960 and 1961,
(Geranium spp,)„ aster (Aster spp ,
) 9
Geranium
willow-herb (Epilobium spp,
) 9
arnica
(Arnica spp,) 9 cinquefoil (Eotentilia spp,) and false dandelion (Aaoseris
spp,) were other important forbs,
Use on each of these genera was rather
uniform varying between four and six per cent.
was minor during the period.
Use of grass-like plants
Two species used but not listed in the table
were mountain brome (Bromus marginatus), recorded at ten pf 35 sites ex­
amined with one per cent usage, and timothy (PhleUm spp,) at five of 35
sites with two per cent usage.
Use on browse was about equal in percent­
age to use of grass-1ike plants with spiraea (Spiraea spp,).showing the
greatest use.
Summer (June 15-July S l 9 I960 and 196l)g
Elk feeding sites were ex­
amined in 1960 only between June 15 and July 31, and for an equal compari­
son only those sites examined between these dates in 1961 are used (Table
V l ),
Dandelion usage during both summers was 40 per cent,
exceeded seven per cent usage in either year.
No other forb
Total forbs used in 1961
slightly exceeded the I960 percentage, however, the table shows a marked
-3 4 -
differeiace in grass and Jbr.ows.e usage between years„x This fnay be due to'
climatic variances.
The total recorded precipitation at the Kings Hill
Weather Station on the western boundary of the study area s in the sprucefir Communitys, at an elevation of T 9SOQ feet* from April to July inclusive,
was 7.29 and
1 0 .1 0
inches for 1960 and 19^1 respectively.
Snow-fall re­
corded at phis station during April and May I960 was 46 Inches 9 but during
the same period in 1961 was 87 inches (U. S. Dept. Comm .
9
J.96Q and 1961).
Twenty-five per cent of the feeding sites examined in 1960 were in the
spruce-fir community in the general area of Kings Hill 9 while in, 19619
forty-two per cent of the sites from' June 15 to July 31 were in this area.
Differences in the location of feeding site ,examinations may account for
the observed differences in food habits for the two summers.
Fall (September)s A total of I9IlS instances of plant use was re­
corded at 13 feeding sites during September.
use of grass increased as compared to summer.
Use of fprbs decreased and
The first heavy recorded
use of grass after the spring period occurred September 4 and usage was
mainly confined to the green Ieavep of bluebunC'h fescue.
A, reliable com­
parison of species usage can he made of the grass-1 ike plants used during
this period since unidentified grass constituted only two per cents
bunch fescue received the greatest usage (22 per cent)?
Blue-
Rouse (1957),re­
corded 30 per cent usage on bluebunch fescue at feeding site examinations
during September.
had none.
Lupine (Lupinus spp.) received heavy Use ,while dandelion
Use of yarrow (Achillea spp,),, not listed ip the -table9 was re­
corded at 17 of 95 feeding sites in small amounts 9 and dyripg September
usage.was two per cent=
During this period browse was Used in larger
-3 5 -
ampUnts than either spring or summer.
Elk food habits on four range types are recorded in Jable VI.
Elk
food habits op specific range types are closely associated with seasonal
food habits because of elk movements.
Ppnderosa pine wpodland-grassIapd community?,
Cp this type^ which is
used as winter and early spring elk range,, grass-like plants received
per cent of the plant Usage.
92
Jhe balance was composed mainly of forbs?
with browse being insignificant.
Douglas fir community (open conifer ? park type)?.
like plants on this
type
Jhe use of grass­
was 35 per cent? while forb use was 63 per cent.
Bluebupch fescue and bluebupch wheatgra,ss (Adropyron spicatum) had recordfed usage of nine and six per cent respectively.
Dandelion and false dande­
lion were important forbs.
Spruce-fir community (open conifer - park type)B
the most Important forage plass.
Forbs constituted
Dandelion reached a peak of utilization
In this type while false dandelion? cinquefoil and geranium were import-,
ant? but at much, lower percentage levels.
Usage on grass-Iike plants was
at a lower level than for any other type.
,Ap abundance, of forbs is
characteristic of this type? especially on me sic and disturbed sites.
Lodgepole pine block clear cuts,?. Jpe rather high use Of grassflike
plants is a result of the Usage Of soft succulent grasses including bluebunch fescue? bluegrasses? spike trisetUm (Trisetum sbicat.um) and timothy
during September.
Lupine? abundant in. the clear cuts? was used heavily ip
September? especially before severe frosts which occurred after the middle
—3 6 —
pf the month.=
Use on lupine may be heavier than indicated in the table =
The plants were often 18 inches tall and over a foot in diameter 9 and
heavy use on. a single plant was recorded as a single instance=
servations suggested the ysage of lupine pods by elk=
Field ob­
This contention was
later substantiated by data from rumen samples collected in the area=Other fprbs used include fireweed (Epilobium angustifol-ium)= aster,?
arnica and dandelion=
Although over 50 per cent of the feeding sites in
the clear cuts were examined during the summer and over 80 per cent were
in the spruce-fir community, dandelion usage was surprisingly low as com­
pared to other range types and reflects the' pautity of this forb in the
clear cut area=
Browse use. was at a maximum as compared to all other
categories in,Table VI.
No use was recorded on low red huckleberry
(VacclniUm scobarlum) one of the most abundant plants in, the vicinity of
the clear cuts=
Thinleaved huckleberry ^Vaccinium Membranaceum) and.buck-
brush (SvmDhoricarpoS spp =).were used at the two per.cant level=
A.general
comparison between elk food habits in clear cuts and availability of plants
present can be made by comparing ,Tables V and VI=
Ruinen AnalysesiB. Table VII shows, elk food habits as determined from
41 rumens collected by Montana Fish and Game, personnel during 1955 and
1956 from the study area =
Analyses of samples were performed in the Wild­
life Laboratory under ,the supervision of Kenneth Gpeer=
,The author comput­
ed the raw data by the same aggregate, percentage method as used for feeding
sites=
Inclusion of these samples furnishes data on forage plass usage for
all seasonss, including late fall and winter which are lacking in the feed­
ing site examinations=
The stomach analyses verified the heavy use of
-
Table VIIo
37-
Analyses of 41 elk rumens collected in the Little Belt
Mountains, 1955 and 1956= I/
Plants 2/
W
Winter
Rumens
Oct.
Nov.
% vol.3/
Rumens
Dec.
Jan.
Feb.
% vol.
Rumens
Mar,
Apr.
May
% vol.
I
Tr.d/
Tr.
3
84
I
Aster spp.
Unidentified forbs
Unidentified grass
Arctostaohvlos uva-ursi
Linnaea spp.
Berberis repens
Unidentified browse
Junioerus communis
Pseudotsuaa menziesii
Unidentified conifer
Mosses and lichens
4
3
I
—
I
Tr.
Forage-classes
Forbs
Grass-like plants
Browse
13
74
13
11
6
74
88
2
2
Sorina
20
I
—
Tr.
2
I
I
2
——
7
88
6
2
4
2
—
— —
5
I
3
84
13
Summer
I
Rumen
June
% vol.
-
68
22
—
—
3
5
——
——
I
I
68
22
9
I/ Raw data supplied to author by Montana Fish and Game Department,
Wildlife Laboratory.
2/ The following plants constituting less than 0.5 per cent in all indivi­
dual seasonal categories have been omitted from the table: Achillea
spp., Antennaria spp., Artemisia frioida. Cirsium spp., Luoinus spp.,
Phlox spp., Compositae Fam., Cyperaceae Fam=, Chimaohila spp., Rosa
spp., Spiraea spp,, Junioerus spp., and Pinus oonderosa.
2/ The volume of each separated plant item was considered as a per cent of
the total identifiable material in each rumen. These percentages were
added for the periods indicated, then divided by the number of rumens
involved in the period.
i/ Tr. = less than 0.5 per cent.
~38 ~
grass in spring and forbs in summer as indicated by feeding sites.
Grass­
like plants constituted the major item in both fall and winter collected
stomachs.
Discussion*
Although elk utilized a wide variety of plants (Table
VI) a few species formed the bulk of the diet.
Table VIII,
Per cent seasonal forage class use comparison of four
Montana elk studies.
Study ^
Forbs
I
II
III
IV
Grass-like plants
I
II
III
IV
Browse
V
Table VIII, which compares
I
II
III
IV
Summer
June
July
Aug.
FaU
Sept.
Oct.
Nov.
11
86
6
91
76
15
13
14
19
Tr.
7
3
4
Tr.
90
91
jSorina
Mar.
Apr.
May
14
Tr.
Winter
Dec,
Jan.
Feb.
88
6
90
82
6
2
74
80
69
100
85
100
100
7
3
13
3
6
22
12
Tr.
Tr.
I
3
4
Tr.
88
7
5
Tr.
Study I (this paper). Spring and summer percentages determined from
feeding sites, fall and winter from rumen analysis.
Study II (Rouse, 1957). Volume percentage determined from 3 rumens
each, summer and winter, 4 rumens each, spring and fall.
Study III (Mackie, 1961).
elk feeding sites.
Percentages determined from examination of
Study IV (Morris & Schwartz, 1957).
analysis of stomach samples.
Percentages determined from
four Montana studies, shows the importance of grass-like plants and forbs,
-
and the light use of h^owse by elk*
39
”
Iri contrast to this^ referenda to the
heavy use oxf browse by elk is cited by Murie (1951) and West (1941)0
Gaffney (1941) records browse as an important elk winter food.
Evident in, the first three studies listed in Table VIII is the heavy
use of forbs during Juhe 9 ,July and ,August..
In study I dandelion received
the greatest Usage 9 in study II sticky geranium (Geranium viscossissimum).
and in study III yellow sweetclover (Melilotus officinalis).
Skinner
(1928) recorded the imporjtanbe of forbs in June 9 July and August ,with
herbaceous cinquefoil (Potentilla diversifolia) most heavily used.
According tb Pickford and Reid (1943)9 weed"herbage formed more than 80
per cent of the elk summer diet in eastern Oregon 9 with pokeweed fleeceflower (Polygonum Phytolaccaefolium) the most important forb used.
the statements of .Smith (1952) are applicable here.
Perhaps
He indicates that the
findings of a food habits study may have local application Only 9 so far as
species evaluation is concerned.
He further suggests that the one food
habit characteristic that may have general application to a big game
species is the one expressing the part played by the various forage classes
in the diet.
Consideration of Table VIII suggests a general pattern of forage class
usage.
Studies I 9 "II and III indicated that grass-like plants were im­
portant at all seasons except summer when forbs were most heavily used.
Browse use varied between seasons, but at a much lower level than either
grass-1ike plants or forbs.
However 9 all elk food habit investigations do
not follow this trend as large seasonal variations of forage classes are
evident.
Study IV (Table VIII) showed a high percentage of grass usage
—4 0 —
during the summer=
Gaffney (1941) gives figures of 60 and 90 per cent
browse in the winter«
A meaningful appraisal of elk food habits involves not only the plant
item consumeds, but also a knowledge of the vegetative type including suc­
cessions! status within which, the animal was feeding=
According to Mohlers
Dalke and Shaw (1958) the shrubby growth following large fires■in. Idaho
has provided an almost unlimited source of food and today elk are largely
found associated with these extensive shrub areas.
The lack of a definite
set pattern of seasonal forage class usage in the elk reflects the adapt­
ability of this animal's feeding habits.
SUMMARY
The elk of the Little Belt Mountains, Montana were studied during
spring and summer of 1961 and summer of 1960 to secure data on distriJS^r,■
tions movements, range Use 9 relationships to block clear cut logging and
food habits.
Four major plant communities were recognized.
These were
described as the ponderosa pine woodland-grassland, ponderosa pine,
Douglas fir and spruce-fir forest communities.
Approximately 2,600 ,elk
comprising 326 groups were classified as to range type used.
About 90 per
cent of elk observed during April and May were using the ponderosa pine
woodland-grassland community of the foothills.
In late May decreasing use
of this community corresponded with increased usage of the Douglas fir
community.
August.
The spruce-fir, community was most heavily used during July and
During late August all elk observed were in this community, usual­
ly in the most inaccessible areas.
Clear cuts were generally used only for
-4 1 -
feeding and those, seven and eight years of age, usually located in re­
lation to mesic conditions, received 57 per cent of all usage by elk
groups observed in clear cuts.
Seventy-four per cent of elk groups in
clear, cuts were using those of a size range of 15.2" to 25.2 acres.
)
Elk I
use, of the clear cuts studied, was not. observed Until the latter part of
June when 0.88 .elk group p e r .observation was recorded,
this figure was 0.13.
-J
hate in August
J
It increased sharply in September to. 1.44, the
highest for any period.■ Measurement of ground vegetation in cut and uncut
lddgepole pine showed an increase of plant species and canopy-coverage in
logged areas, although Vaccinum sop, coverage decreased in cut-over blocks.
Feeding site examinations showed that grass-1ike plants constituted
88
,
6
and 40 per cent of the plants used for spring, summer and fall respective­
ly, while the figures for forbs were 11,.
seasons.
86
, and 51 per cent for comparable
Dandelion constituted 44 per cent of the plant usage during June,
July and August.
Grass-like plants accounted for
88
per cept of the con­
tents of 20 rumens collected during December, January and February.
Forb
Usage was at a minimum, (eight per cent) in the pgnderosa pine woodlandx '•
grassland community, and at a maximum (82 per cent) in the spruce-fir com­
munity,
Figures for grass-like plants were 92 and 13 per cent QRt the two
communities respectively.
Elk food habits ,in ,clear cut areas reflected the
availability of certain plant species,
Comparing the food habits results
J
of this study with a few others revealed a close correlation, in seasonal
forage class usage, but a wide divergence in plant species usage,
-42-
LITERATURjE CITED
Booths, .W0 E 0 1950o Flora of Montana ^ Part I9 Conifers and Monocots0
Research Foundation at Montana State College, Bozeman9 Montana»
232 p p 0
; • '''i .and J 0 C 0 Wright0 1959° Flora of Montana, Part II9
Dicotyledons. Montana State College? Bozeman 9 Montana„ 280 pp 0
Cole 9 G 0 F 0 and B 0 T 0 Wilkins.
1958» The pronghorn antelope 9 its range
use and food habits in central Montana with special reference to
wheat, Tech 0 Bull, N o 0 2, Montana Fish and Game Department, 39 pp,
Cottam 9 G, and J 0 T, Curtis 0 1956° The use of distance measures in
phytpsociological sampling. Ecology, 37(3)$ 451-460,
Dapbenmire9 R 0 F 0 1943, VegetsTtIonal zonation in the Rocky Mountains,
The Botanical Review 9 9 (6 )s 325-393,
o 1946, The life zone problem in the northern intermountain
region. Northwest Science, 20(2) s. 28-38,
■"
’
. 1952, Forest vegetation of northern Idaho and adjacent
Washington, and its bearing on. concepts of vegetation classification,
Ecol, Monog,, 22(4)s 301-330,
■— ■ ... — >
1953° Classification of the conifer forests of eastern
Washington and northern Idaho, Northwest Science, 27(I )s 17-24,
,0
1 9 5 9 „ A canopy-coverage method of vegetations! analysis.
Northwest Science, 33(1)s 43-64,
Gaffney, W, S, 1941, The effects of winter elk browsing, South Fork of
the Flathead River, Montana, J, Wildl, Mgmt,, 5(4)5.427-453,
Gieseker, L, F,, C, B° Manifold, A, T, Strahorn 9 and 0 , F» Bartholomew,
1953, Soil survey, central Montana,
U, S 0 Dept, of Ag^, and Mont,
Agr, Expt, Sta, Series 1940, No, 9, 133 pp,
Johnson, D, E, 1951, Biology of the elk calf, Gervus canadensis nelsoni,
J, Wi.ldl, Mgmt0, 15(4)5 396-410,
' '
Leiberg 9 J, B, 1004, Forest conditions in the Little Belt Mountains for­
est reserve, Montana, and the Little Belt- Mountain quadrangle,
U, Sd
Geological Survey, Professional paper No, 30, Govt, Print, Off,
Washington, D, C 0 75 pp,
LoVaas , A., L 0 1958, Mule deer food habits and.range use, Little Belt
Mountains, Montana, J 0 Wildl, Mgmt,, 22(3)?. 275-283,
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Macki.e, R.
1961, DGer, elk and cattle food habits and range relation­
ships in the Missouri breaks area, P« R, Completion Report, Project
W^98-R.-l. Mont. Fish and Game Dept. Multilith, 28 pp.
Martin, A» C . , R, H. Gen.s.ch and C. p. Brown.
1946. Alternative methods
in Upland game bird food analysis, J. Wildl« Mgmt. 10(1)$ 8-12.
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1942, Judith River big game, investigation 1941-42.
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Mohler, L._L», P. D. Dalke and W« M. Shaw.
1958. Elk and elk hunting in
IdahoT Trans. N. Am. .Wildl. Conf. 23$ 491.-501.
Morris, M. S, and J. E» Schwartz.
1957. Mule deer and elk food habits on
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1951. Jhe elk of North America,
burg, Pa. 376 pp.
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1957. '.Elk food habits, range use and movements, Gravelly
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Unpubi. thesis (M.S. )s Montana State College,
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1928.
The elk situation.
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Smith, G. A.
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..
Uo S 0 Dep,to Commerce0 1960; 1961= ' Climatological
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Varner,
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k
*