Relations of moose, cattle, and willows in southwestern Montana
by Robert Donald Dorn
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 Robert Donald Dorn (1969)
Abstract:
The Shiras moose {Aloes aloes shirasi) was studied on Red Rock Lakes National Wildlife Refuge in
southwest Montana from June 10, 1968 to March 17, 1969 to determine use of vegetation types, food
habits, sex and age ratios, and the extent of moose-cattle competition. A means for checking the
adequacy of a sample obtained by the feeding site method of food habits analysis was presented.
Vegetation of the study area was analyzed quantitatively. The average height of a willow (Salix)
species was related to its most common habitat. Observations of moose in the willow type accounted
for 84 percent of the total in summer and 93 percent in winter. Food habits primarily determined the
use of vegetation types. Browse accounted for 98.3 percent of all forage used by moose in summer with
Salix myvtillifolia, Betula glandulosa, Salix geyeriana3 and Salix planifolia accounting for 58.1, 11.8,
9.6, and 6,7 percent, respectively. In winter browse accounted for 99.8 percent of the forage used by
moose with S. myvtillifolia, S. planifolia, S. bebbiana, and S. geyeriana accounting for 25.0, 24.2, 15.4,
and 10.5 percent, respectively. The factors determining the amount of various plant species in the diet
of moose were availability, adaptation, palatability, and habit. It appeared that all species of willow
present in this study were potentially important as forage for moose at some time during the year,
especially in winter. Salix Wolfii, S, myrtillifolia, S. geyeriana, and S. bebbiana accounted for 50.2,
15.9, 11.2, and 10.9 percent, respectively, of all browse used by cattle in the willow and sedge types in
summer. Forage competition between moose and cattle was not significant under conditions prevailing
during this study. Competition would be expected under certain conditions. Forage competition
between moose and beaver (Castor canadensis) was greater than competition between moose and cattle
but was not significant. Observation data show that in summer bulls were more readily identified from
the ground than cows and that many calves were not observed from the ground. The data also show that
observed sex and age ratios were influenced by home range and migration patterns and differential use
of vegetation types between sexes. September ratios obtained from aerial observations were probably
most reliable for the composition of the breeding population. Only a small proportion of adult bulls
wintered on the study area. Resident, summer, winter, and transient population segments appeared to
be using the area. The importance to management of movements and twinning rates was discussed. The
range appeared to be in satisfactory condition during the study except for one local area. Management
recommendations were given. I n
RELATIONS OF MOOSE, CATTLE, AND WILLOWS
IN SOUTHWESTERN MONTANA
by
ROBERT DONALD DORN
A th e s is subm itted to
fu lfillm e n t o f the
th e G raduate F acu lty in p a r t i a l
requirem ents fo r th e degree
of
MASTER OF SCIENCE
in
Fish
and W i l d l i f e
Management
Approved:
MONTANA STATE UNIVERSITY
Bozeman, M ontana •
‘
June,
1969
iii
ACKNOWLEDGMENT
A p p r e c i a t i o n i s e x te n d e d t o t h e f o ll o w i n g f o r t h e i r a s s i s t a n c e o r
c o o p e r a t i o n d u r in g t h e s t u d y :
ty , fo r s e ttin g
D r. D. C. Q uim by, M ontana S t a t e U n i v e r s i ­
up t h e s tu d y and f o r g u id a n c e and s u g g e s t i o n s i n p re -.
p a r i n g t h e m a n u s c r ip t ; D r. R. J . Graham and D r. R. L . E ng, M ontana S t a t e
U n i v e r s i t y , f o r r e a d i n g t h e m a n u s c r ip t and o f f e r i n g s u g g e s t i o n s , f o r i t s
im p ro v e m e n t; D r. W. E . B o o th , M ontana S t a t e U n i v e r s i t y , f o r many h o u rs
a t v e r i f i c a t i o n o f p l a n t s p e c im e n s ; D r. G. W. A rg u s , U n i v e r s i t y o f
S a s k a tc h e w a n , S a s k a to o n , f o r v e r i f i c a t i o n o f w illo w s p e c im e n s ; M r. Thomas
M u sse h l and M r, P h i l i p S c h l a d w e i l e r , M o n tan a F i s h and Game D e p a rtm e n t,
f o r a s s i s t a n c e i n v a r i o u s p h a s e s o f t h e s t u d y ; M r. K e n n e th G r e e r and
s t a f f , M ontana F i s h and Game D e p a rtm e n t, f o r g r o s s s e p a r a t i o n o f rumen
s a m p le s ; o t h e r p e r s o n n e l o f M ontana F i s h and Game D e p a rtm e n t f o r a s s i s ­
t a n c e and c o o p e r a t i o n ; U. S ., F i s h and W i l d l i f e S e r v ic e p e r s o n n e l and
r e s i d e n t s o f L a k e v ie w , M o n id a , and t h e C e n t e n n i a l V a lle y f o r a s s i s t a n c e ,
c o o p e r a t i o n , and h o s p i t a l i t y ; and M r, G e o rg e Kam ps, M ontana S t a t e U ni­
v e r s i t y , f o r m aking a v a i l a b l e f e e d in g s i t e d a t a f o r m u le d e e r and w h i t e t a i le d d e e r.
The s tu d y w as f in a n c e d b y t h e M ontana F i s h an d Game
D e p a rtm e n t u n d e r F e d e r a l A id P r o j e c t s W-9 S -R -9 and W -73-R -14,
iv
TABLE OF CONTENTS
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ACKNOWLEDGMENT . . . .
. iii
TABLE OF CONTENTS .
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INTRODUCTION
METHODS
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TABLE OF CONTENTS
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S e x a n d Age R a t i o s 9 P o p u l a t i o n T r e n d s 9 a n d M o v e m e n t s
Range S u r v ey
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39
APPENDIX . . . ©©, ©
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LITERATURE CITED
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75
Management Recommendations
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1
VX
L I ST OF TABLES
T a b le
H
H
I.
in.
Page
CLIMATOLOGICAL DATA FOR LAKEVTEW, MONTANA ..........................
SAMPLING DATES, SAMPLE SIZE, AND" METHODS USED
FOR VEGETATION TYPE ■ANALYSES .......................................... ... ='. ...
2
.
£
INTERCEPT, CANOPY-COVERAGE, AND FREQUENCY OF IMPORTANT
PLANTS ON THE STUDY AREA BY VEGETATION T Y P E ............... ..
13
HEIGHTS AND FREQUENCY INDICES OF WILLOWS ON THE
STUDY ^VREA.
16
USE OF VEGETATION TYPES BY.MOOSE FOR FEEDING
AND BEDDING
19
• o o o o o o o o o e e e e o e e e o c o e e o e o c e o e e . e e e e e e ' t e c e - e e e e e
V.
e e o o e o o e o o e
VIo
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6, 0
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o
o
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e o e e e e o e o e e e ' e e e e e e e e e e e e o e
FOOD HABITS OF CATTLE AND MOOSE .
.
21
PROBABLE MOST IMPORTANT FACTORS DETERMINING THE AMOUNT
OF USE OF IMPORTANT FORAGE SPECIES OF MOOSE IN THIS
STUDY
26
OBSERVED SEX AND AGE RATIOS OF MOOSE ON THE
STUDY ^VREA
29
x ■
PERCENT VOLUME AND FREQUENCY OF AQUATIC TYPE TAXA
42
Xo
PERCENT CANOPY-COVERAGE, FREQUENCY, AND INTERCEPT
OF SEDGE TYFE TAXA
43
PERCENT CANOPY-COVERAGE, FREQUENCY, AND INTERCEPT
OF WILLOW TYPE *TAXA
45
PERCENT CANOPY-COVERAGE, FREQUENCY, AND INTERCEPT
OF SPRUCE TYPE TAXA ©b©©©'©©©©©©©©©©©©©©©©©©©©©©©©©©,©©©*®©©"
48
VIIo
e e e o o o e e o e o s o o ' e e e e o o e o e o e e ' e e o p o e o e ' e o e o e e e o o e o e e o e g e
VTIIo
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o « e e * « o o * o o c o o © ‘e o o o ' o e o o © o o o ‘ © « ® o o o e « < s © o *
XI.
X II.
X IIIo
PERCENT CANOPY-COVERAGE AND FREQUENCY OF GRASSLAND
50
XIVo
PERCENT' CANOPY-COVERAGE, FREQUENCY, AND, INTERCEPT
OF ASPEN TYPE TAXA
52
PERCENT CANOPY-COVERAGE, .FREQUENCY, AND INTERCEPT OF
DOUGLAS—FIR TYPE TAXA ©o*©©©.©©"©©,©©©©©©©©©©#*©©*©©**©©.©©©©
55
© © © © © © ' © © e © © © © © © © © © © © © ' © © © © © © © © © © © © © © ' © © ©
XVo
v ii ,
T a b le
XVI.
X V II.
X V III.
XIX.
L I ST OF T AB L E S .
(continued)
Page
PERCENT CANOPY-COVERAGE a FREQUENCY, AND INTERCEPT
OF SPRUCE-FIR TYPE TAXA . . . . . ...................................................... . . . . . .
57
RESULTS OF POINT-CENTERED QUARTER METHOD ANALYSES
OF TREES ................. ..................................... ........................................................... ..
58
PLANT DISTRIBUTION BY VEGETATION TYPE ON THE STUDY
AREA . . . . . . . . o o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
59
FORAGE PLANTS WITH LESS THAN 2 PERCENT USE (EXCLUDING
THE GENUS SALIX) BY MOOSE AND CATTLE IN ALL CATEGORIES
EXCEPT FREQUENCY FOR RUMEN SAMPLES ...........................................
73
v iii
L I S T OF FIGURES
F ig u re
I o
2,
3,
4©
P ag e
L o c a ti o n o f t h e s tu d y a r e a , v e g e t a t i o n t y p e s , and
v e g e t a t i o n t r a n s e c t s o«© o o©©oo©©©©oo©oo©oo©©c©©9o©o© ©©
3
I n s t a n c e s - o f - u s e c u r v e s f o r d e te r m in in g t h e a d e q u a c y
o f a s a m p le o b t a i n e d by t h e f e e d i n g s i t e m ethod o f
fo o d h a b i t s a n a l y s i s : A, m oose i n sum m er; B , c a t t l e
i n sum m er; C, m oose i n w i n t e r ©ocooeoooocoooo©©©' ©©©©' ©©© * ©©©©
9
M ost common h a b i t a t o f w illo w s p e c i e s w i t h r e s p e c t t o
s o i l m o i s t u r e and m i c r o c l i m a t e ©©©©©©©©©©©©© ©©©©©©©'©©©©'.©©©©
17
Maximum d a i l y o b s e r v a t i o n s o f i n d i v i d u a l m o o se .b y
s e m im o n th ly p e r i o d s ©©©©©©©©©©©©©©©©©©©©©©©©©©©©,©c
D i s t r i b u t i o n o f w illo w h e i g h t s on t h e s tu d y a r e a
by s p e c i e s
72
G round o b s e r v a t i o n s o f m oose i n summer and w i n t e r by
q u a r t e r - s e c t i o n and o b s e r v a t i o n d a t e s and l o c a t i o n s
o f t^zo m arked b u l l s ©©©©©©©o©©©©©©©©©*©©©©©©©©©©©©©©©©©©©©
74 .
© o © © © © © © © © © © © © © © © © © © © © © © © © © © © © © © © © © . © o ©
6©
ix
ABSTRACT
The S h i r a s m oose (AZ-ces a lo es s h ir a s i) was s t u d i e d on Red Rock L akes
N a t i o n a l W i l d l i f e R e fu g e i n s o u th w e s t M o n tan a from J u n e 1 0 , 1968 to M arch
1 7 , 1969 t o d e te r m in e u s e o f v e g e t a t i o n t y p e s , f o o d . h a b i t s , s e x and ag e
r a t i o s , an d t h e e x t e n t o f n io o s e -c a t t l e c o m p e t it io n .
A m eans f o r ch e ck ­
in g t h e a d e q u a c y o f a s a m p le o b t a i n e d b y t h e f e e d in g s i t e m eth o d o f fo o d
h a b i t s a n a l y s i s was p r e s e n t e d . • V e g e t a t i o n o f t h e s tu d y a r e a was a n a ly z e d
q u a n tita tiv e ly .
The a v e r a g e h e i g h t o f a w illo w (S a lix ) s p e c i e s was r e ­
l a t e d t o i t s m o st common h a b i t a t .
O b s e r v a tio n s o f m oose i n t h e w illo w
t y p e a c c o u n te d f o r 84 p e r c e n t o f t h e t o t a l i n summer and 93 p e r c e n t i n
w in te r.
F o o d .h a b i t s p r i m a r i l y d e te r m in e d t h e u s e o f v e g e t a t i o n ty p e s ..
B row se a c c o u n te d f o r 9 8 .3 p e r c e n t o f a l l f o r a g e u se d by m oose i n summer
w i t h S a lix m y v t i l l i f o l i a 3 B etu la Qlandulosa3 S a lix Qeyeriana3 and S a lix
Q la n ifo lia a c c o u n tin g f o r 5 8 .1 , 1 1 .8 , 9 . 6 , and 6 ,7 p e r c e n t , r e s p e c t i v e l y .
I n w i n t e r b ro w s e a c c o u n te d f o r 9 9 .8 p e r c e n t o f t h e f o r a g e u s e d by m oose
w i t h S. myV t i l l i f o l i a 3 S . Q la n ifo lia 3 5 . Bebbiana3 and S. geyeviana a c ­
c o u n tin g f o r 2 5 .0 , 2 4 .2 , 1 5 .4 , and 1 0 .5 p e r c e n t , r e s p e c t i v e l y .
The
f a c t o r s d e te r m in in g t h e am ount o f v a r i o u s p l a n t s p e c i e s i n t h e d i e t o f
m oose w e re a v a i l a b i l i t y , a d a p t a t i o n , p a l a t a b i l i t y , and h a b i t .
I t ap­
p e a r e d t h a t a l l s p e c i e s o f w illo w p r e s e n t i n t h i s s tu d y w e re p o t e n t i a l l y
i m p o r ta n t a s f o r a g e f o r m oose a t some tim e d u r in g t h e y e a r , e s p e c i a l l y i n
w i n t e r . S a lix W o lfii3 S. myV t i l l i f o l i a 3 S. Qeyeviana3 and S . bebbiana
a c c o u n te d f o r 5 0 .2 , 1 5 .9 , 1 1 .2 , and 1 0 .9 p e r c e n t , r e s p e c t i v e l y , o f a l l
b ro w s e u s e d b y c a t t l e i n t h e w illo w and s e d g e ty p e s i n sum m er. F o ra g e
c o m p e t it io n b e tw e e n m oose an d c a t t l e was n o t s i g n i f i c a n t u n d e r c o n d i­
t i o n s p r e v a i l i n g d u r in g t h i s s tu d y .
C o m p e titio n w o u ld b e e x p e c te d
u n d e r c e r t a i n c o n d i t i o n s * F o ra g e c o m p e t it io n b e tw e e n m oose and b e a v e r
(Castor canadensis) was g r e a t e r t h a n c o m p e t it io n b e tw e e n m oose and
c a t t l e b u t was n o t s i g n i f i c a n t .
O b s e r v a t i o n d a t a show t h a t i n summer
b u l l s w e re m ore r e a d i l y i d e n t i f i e d from t h e g ro u n d t h a n cows and t h a t
many c a lv e s w e re n o t o b s e rv e d from t h e g r o u n d . The d a t a a l s o show
t h a t o b s e rv e d s e x and a g e r a t i o s w e re i n f l u e n c e d by home r a n g e and m i­
g r a t i o n p a t t e r n s and d i f f e r e n t i a l u s e o f v e g e t a t i o n t y p e s b e tw e e n s e x e s .
S e p te m b e r r a t i o s o b t a i n e d from a e r i a l o b s e r v a t i o n s w e re p r o b a b l y m o st
r e l i a b l e f o r t h e c o m p o s itio n o f t h e b r e e d i n g p o p u l a t i o n .
O nly a s m a ll
p r o p o r t i o n o f a d u l t b u l l s w i n t e r e d on t h e s tu d y a r e a .
R e s i d e n t , summ er,
w i n t e r , and t r a n s i e n t p o p u l a t i o n s e g m e n ts a p p e a r e d t o b e u s in g t h e a r e a .
The im p o r ta n c e t o m anagem ent b f m ovem ents an d tw in n in g r a t e s was d i s ­
c u s s e d . The r a n g e a p p e a r e d t o b e i n s a t i s f a c t o r y c o n d i t i o n d u r in g th e
s t u d y e x c e p t f o r one l o c a l a r e a . M anagem ent re c o m m e n d a tio n s w e re g iv e n .
INTRODUCTION
T h is i n v e s t i g a t i o n w as a n o t h e r se g m e n t o f t h e e c o l o g i c a l s t u d i e s ■
o f t h e S h i r a s m oose i n s o u th w e s t e r n M o n ta n a , - F i e l d w ork w as c o n d u c te d
f u l l - t i m e i n summer a n d w i n t e r and p a r t - t i m e i n f a l l fro m J u n e 1 0 , 1968
t o M arch 1 7 , 1969„
The m a in o b j e c t i v e s w e fe t o d e te r m in e u s e o f v e g e ­
t a t i o n t y p e s , fo o d h a b i t s ; s e x and a g e r a t i o s , and t h e e x t e n t o f m o o sec a t t l e c o m p e t it io n i n a p r e d o m in a n tly w i l lo w - s e d g e a r e a . - P r e v io u s
s t u d i e s i n s o u th w e s t e r n M o n tan a w e re by K n o w lto n .( 1 9 6 0 ) , P e e k ( 1 9 6 2 ),
S te v e n s ( 1 9 6 5 ) , and S te v e n s . ( 1 9 6 7 ).
M c M illa n ( 1 9 5 3 ) , i n Y e l lo w s to n e ,
N a t i o n a l P a r k , and D e n n is to n ( 1 9 5 6 ) , H a r ry ( 1 9 5 7 ), and H o u s to n ( 1 9 6 8 ),
i n J a c k s o n H o le , Wyoming, a l s o s t u d i e d t h e S h i r a s m oose.
AREA DESCRIPTION
The s tu d y a r e a , a b o u t 22 s q u a r e m i l e s , was m a in ly on t h e Red Rock
L ak es N a t i o n a l W i l d l i f e R e fu g e i n t h e C e n t e n n ia l V a l l e y , B e a v e rh e a d
C o u n ty , M o ntana ( F ig u r e I ) „
The Red Rock L ak es l i e i n a d o w n -d ro p p ed
b a s i n a t 6 ,6 0 0 f e e t e l e v a t i o n (Banko 1 9 6 0 :4 0 -4 1 )=
The C e n t e n n ia l
M o u n ta in s r i s e a b r u p t l y t o n e a r l y 1 0 ,0 0 0 f e e t im m e d ia te ly t o t h e s o u th
and t h e f o o t h i l l s o f t h e G r a v e ll y R ange l i e
to th e n o r th .
A ld e n (1953)
d i s c u s s e d a r e a g e o lo g y a n d N i e ls o n and F a r n s w o rth (1 9 6 5 ) c l a s s i f i e d
a re a s o i l s .
The a r e a h a s lo n g , c o ld .w i n t e r s and s h o r t , c o o l sum m ers.
C lim a to ­
l o g i c a l d a t a f o r L a k e v ie w , on t h e s tu d y a r e a , a r e p r e s e n t e d i n T a b le I ,
Snow c o v e r i s v a r i a b l e from e x t e n s i v e d r i f t i n g .
TABLE I .
CLIMATOLOGICAL DATA FOR LAKEVIEW, MONTANA (U. S. DEPARTMENT
OF COMMERCE, WEATHER BUREAU 1 9 6 5 , FARNES 1 9 6 9 ).
A v e ra g e a n n u a l p r e c i p i t a t i o n
A v e ra g e a n n u a l s n o w f a ll
Mean t e m p e r a t u r e
A nnual
J a n u a ry
J u ly
T e m p e r a tu r e e x tre m e s
Maximum
Minimum
Snow c o v e r ^
J a n u a r y 2 , 1969
J a n u a r y 3 1 , 1969
M arch 3 , 1969
19=96 i n c h e s
1 4 2 .1 i n c h e s
34.6°?
1 0 .3 ° F
58.9°F
91° • F
-4 4 ° F
26 i n c h e s
71 i n c h e s
78 i n c h e s
I / D a ta from F a m e s (1 9 6 9 ) f o r snow s u r v e y c o u r s e i n D o u g l a s - f I r — s p r u c e f i r e c o to n e a t 6 j9 3 0 f e e t e l e v a t i o n .
LEGEND
V egetation
Types
Roads
Vegetation
Transects
Gravelly
Moun tains
Aspen
Streams
A
D
Dougl as-f i r
Lakeview
Montana
O
Gr as s l a n d
Aquatic
-
Sedge
□
Willow
8
A
Spruce
Sedge
Wi l l o w
Aspen
G rassland
WlW
Douglas-fi
Spruce-fir
I
^ U p p . ,
Centennial
Red
lake
R ockiS gg
Mountains
Scale
F ig u re I .
L o c a tio n o f t h e s tu d y a r e a , v e g e t a t i o n t y p e s , and v e g e t a t i o n t r a n s e c t s
UJ
I
—4—
Red Rock L ak es R e fu g e , now a b o u t 4 0 ,0 0 0 a c r e s , was e s t a b l i s h e d i n
1935 t o p r e s e r v e t h e t r u m p e te r swan- (Olov buccinator)',
u s e s i n n o n -sw a n h a b i t a t a r e c a t t l e g r a z i n g a n d h a y i n g „
The p r im a r y la n d
R e c re a tio n a l
u s e i s h e a v y i n some a r e a s .
Moose w e re o b s e r v e d i n t h e C e n t e n n ia l V a lle y s h o r t l y a f t e r 1900 b u t
w e re n o t num erous u n t i l a f t e r 1940,,
1950 ( Banko 1 9 5 1 : 1 ) .
A c o n s p ic u o u s i n c r e a s e o c c u r r e d a b o u t
H o u s to n ( 1 9 6 8 :1 6 -1 7 ) d e s c r i b e d t h i s sam e p a t t e r n
f o r J a c k s o n H o le , Wyoming.
H o u sto n ( 1 9 6 8 :1 3 , 35) a l s o r e p o r t e d a p r o ­
g r e s s i v e i n c r e a s e i n m ean a n n u a l s n o w f a l l i n J a c k s o n H o le from 122 in c h e s
i n t h e 1 9 1 2 -1 9 3 0 p e r i o d t o 179 i n c h e s i n t h e 1 951-1966 p e r i o d .
D a ta f o r
L ak ev iew (U.- S . D e p a rtm e n t o f Commerce, W e a th e r B u re a u 1 9 5 5 :4 0 , 1 9 6 5 :4 2 )
show a s i m i l a r i n c r e a s e from 132 i n c h e s i n t h e 194 3 -1 9 5 2 p e r i o d t o 159
i n c h e s i n t h e 1951 -1 9 6 0 p e r i o d .
T h is may h a v e r e s u l t e d i n c o n d i t i o n s
w h ic h i n c r e a s e d f a v o r a b l e h a b i t a t f o r m oose a t t h e sam e tim e t h a t t h e
g r i z z l y b e a r {Ursus h o r r i b i t i s )
and t im b e r w o lf (Canis lupus ) , t h e
p r im a r y p r e d a t o r s o f m oose (H o u sto n 1 9 6 8 :7 9 , Mech 1 9 6 6 :1 1 4 ) , w e re b e in g
red u ced .
C hanges i n la n d u s e may a l s o h a v e i n f l u e n c e d t h e i n c r e a s e .
O v e r - u t i l i z a t i o n o f b ro w se r e s u l t e d from t h e p o p u l a t i o n i n c r e a s e on
t h e r e f u g e a ro u n d 1950 (Banko 1 9 5 1 :2 ) .
A d i e - o f f ■o f w illo w s o c c u r r e d
.
s h o r t l y a f t e r from unknown c a u s e s .
The r e f u g e was o p e n e d t o m oose h u n t ­
in g b y p e r m it i n 1952 w hen t h e m tiose p o p u l a t i o n a t p e a k u s e was a b o u t 60
,
A n im a ls.
A t o t a l o f 128 m oose was k i l l e d o v e r e i g h t s u c c e s s i v e s e a s o n s .
A c l o s u r e o f 5 y e a r s ' f o ll o w e d .
F iv e p e r m i t s w e re i s s u e d a n n u a ll y from
1965 t o 1968 a n d 19 m oose w e re k i l l e d
(R e fu g e r e c o r d s ) .
a t p e a k u s e i n 1 968-69 was a b o u t 55 a n im a ls .
The p o p u l a t i o n
METHODS
V e g e t a t i on
The c a n o p y -c o v e ra g e m eth o d (B a u b e n m ire 19.59) , l i n e - i n t e r c e p t m eth o d
(G a te s 1 9 4 9 :3 6 - 3 8 ) , p o i n t - c e n t e r e d q u a r t e r m eth o d ( C o ttam an d C u r t is
1 9 5 6 ) , and h a b i t a t i n v e n t o r y t e c h n i q u e (W e b s te r 1965) w e re u s e d t o sa m p le
v e g e t a t i o n ty p e s =
S a m p lin g d a t e s , s a m p le s i z e , and m e th o d s f o r e a ch v e g e ­
t a t i o n ty p e a r e p r e s e n t e d i n T a b le I I .
S am ple t r a n s e c t s w e re m o s tly s e ­
l e c t e d i n a r e a s f r e q u e n t e d b y m oose and n e a r s e c t i o n l i n e s o r la n d m a rk s
( F ig u r e I ) .
T r a n s e c t s w e re 300 f e e t lo n g w i t h 33 f e e t 4 i n c h e s b e tw e e n
p l o t s o r p o in ts =
U pper Red Rock L a k e , ra n d o m ly sa m p le d b y t h e r e f u g e
b i o l o g i s t , r e p r e s e n t e d t h e a q u a t i c ty p e=
V e g e t a t io n ty p e s w e re mapped
fro m g ro u n d o b s e r v a t i o n s and a e r i a l p h o to s =
W illo w h e i g h t s w e re m e a s u re d w i t h an 1 1 - f o o t p o l e m ark e d i n f e e t .
H e ig h ts o v e r 20 f e e t w e re e s t i m a t e d .
The c l o s e s t p l a n t s a m p lin g
t e c h n i q u e (C o le 1 9 6 3 :7 ) was u s e d f o r common s p e c ie s =
L e s s common s p e c i e s
w e re m e r e ly s o u g h t o u t . ■ H e ig h ts w e re f i r s t m e a s u re d w i t h i n a b o u t 50
y a r d s o f t h e t r a n s e c t s i n t h e w illo w ty p e=
I n s u f f i c i e n t s a m p le s w e re
t h e n c o m p le te d i n a r e a s a t l e a s t o n e - q u a r t e r m i l e from any t r a n s e c t = .
The d e g r e e o f d i f f i c u l t y i n l o c a t i n g p l a n t s o f a . s p e c i e s t o m e a s u re
was p r o p o r t i o n a l t o t h e r e l a t i v e a b u n d a n c e o f t h a t s p e c ie s =
i n d i c e s f o r e a c h s p e c i e s c o u ld t h e n b e d e te r m in e d .
F re q u e n c y
I f a t l e a s t te n
p l a n t s o f a s p e c i e s w e re fo u n d w i t h i n a b o u t 50 y a r d s o f e a c h o f th e t e n
t r a n s e c t s i n t h e w illo w t y p e , t h e f r e q u e n c y in d e x was 10 0 .
a b l e sa m p le w as a l s o c o n s id e r e d f o r r a r e s p e c ie s =
The o b t a i n ­
O n ly . t e n p l a n t s o f
o n e s p e c i e s w e re f o u n d , b u t a l l w e re n e a r a t r a n s e c t f o r a f re q u e n c y
TABLE I I .
SAMPLING DATES, SAMPLE SIZ E , AND METHODS USED FOR VEGETATION TYPE ANALYSES.
V e g e t a t io n
Type
S am p lin g
D a te s
A q u a tic
8/24
Sedge
8 /2 7 -2 8 -
W illow
7 /3 - 7
S p ru e e
No... o f
T r a n s e c ts
No. -o f P l o t s
o r P o in ts
T r e e Group
METHOD^/
Shrub Group
50
5
50
L i n e - i n t e r cep t
10
100
L in e -in te rc e p t L in e -in te rc e p t
7 /2 3 -2 5
3
’ 30
G r a s s la n d
6 /2 5 -7 /2
6
60
A spen
7 /1 5 -1 8
5
50
D o u g la s -fi r
7 /1 9 n 2 0 -
4
40
S p ru c e -fir
7 /1 9
I
10
I / S e e .t e x t , page 5 , f o r c i t a t i o n .
L in e -in te rc e p t L in e -in te rc e p t
P o in t-q u a rte r
C a n o p y -c o v e ra g e
L in e -in te r c e p t L in e -in te rc e p t
P o in t-q u a rte r
L in e -in te rc e p t L in e -in te rc e p t
P o in t-q u a rte r
L in e -in te rc e p t L in e -in te rc e p t
P o in t-q u a rte r
H erb Group
H a b ita t
in v e n to ry
Canopy-r
c o v e ra g e
C anopyc o v e ra g e
C anopyc o v e ra g e
C anopyc o v e ra g e
C anopyc o v e ra g e
C anopyc o v e ra g e
C anopyc o v e ra g e
-7 -
in d a x o f 1 0 . . F o r a n o t h e r s p e c i e s 35 p l a n t s w e re f o u n d , b u t o n ly 5 w e re
n e a r a t r a n s e c t f o r a f re q u e n c y in d e x o f 5 .
The l a r g e r num ber o f p l a n t s
fo u n d o f t h e l a t t e r s p e c i e s i n d i c a t e d a h i g h e r a b u n d a n c e t h a n f o r t h e
fo rm e r d e s p i t e t h e f r e q u e n c y in d e x f i g u r e s .
R e p r e s e n t a t i v e s o f m o st p l a n t s p e c i e s W ere c o l l e c t e d a n d a r e f i l e d
i n my p e r s o n a l c o l l e c t i o n , i n t h e h e r b a r iu m a t M ontana S t a t e U n i v e r s i t y ,
a n d / o r a t Red Rock L a k e s R e fu g e .
P l a n t n o m e n c la tu r e f o ll o w s P o r t e r
(1 9 6 7 ) f o r t h e g e n u s Sdl-Lx a n d B o o th (1 9 5 0 ) and B o o th and W rig h t (1966) ,
w i t h some u n p u b l is h e d c o r r e c t i o n s , f o r o t h e r p l a n t s u n l e s s o t h e r w i s e
n o te d .
N in e ty - o n e w illo w s i d e n t i f i e d i n summer w e re ta g g e d t o f a c i l i t a t e
i d e n t i f i c a t i o n o f w illo w s i n w i n t e r .
G e n e r a l d i s t r i b u t i o n o f W illow
s p e c i e s w as d e te r m in e d from f i e l d o b s e r v a t i o n s .
V e g e t a t i o n T y p e Us e
O b s e r v a t i o n s o f m oose i n sum m er, J u n e 10 to. S e p te m b e r 3 0 , w e re
m o s tly from r o a d s an d w e re a id e d by a 7 x 50 b i n o c u l a r and a 25X s p o t t i n g
scope.
ris e .
A r e g u l a r r o u t e was t r a v e r s e d n e a r l y e v e ry day s h o r t l y a f t e r s u n ­
O b se rv e d m oose w e re r e c o r d e d b y v e g e t a t i o n ty p e an d l o c a t i o n .
e v e n in g o b s e r v a t i o n s w e re m ade.
Few
O b s e r v a tio n s i n w i n t e r , D ecem ber 20 t o
M arch 1 7 , w e re i n c i d e n t a l t o t h e s tu d y o f fo o d h a b i t s e x c e p t f o r an i n ­
t e n s i v e s u r v e y tw i c e e a c h m onth by sn o w m o b ile .
N e a r ly a l l a n im a ls p r e ­
s e n t w e re p r o b a b l y o b s e rv e d d u r in g t h e s e s u r v e y s e x c e p t i n D ecem ber arid
Ja n u ary .
A f e e d i n g s i t e ex am in ed w h i l e snow t r a c k i n g w i t h o u t o b s e r v in g
t h e a n im a l w as - c o n s id e r e d a n o b s e r v a t i o n .
—8—
Food H a b i t s
Food h a b i t s o f m oose and c a t t l e i n summer and o f m o o s e .in w i n t e r
w e re d e te r m in e d from f e e d i n g s i t e o b s e r v a t i o n s (K now Iton 1 9 6 0 :1 6 2 -1 6 3 )„
An i n s t a n c e o f u s e on b ro w s e was u t i l i z a t i o n o f one l e a f o r one l e a d e r
i n summer and one l e a d e r i n w i n t e r and on f o r b s , g r a s s , a n d g r a s s - l i k e
p l a n t s was u t i l i z a t i o n o f o n e . l e a f . o r o n e .s te m .
An i n s t a n c e o f u s e on
b a r k was u t i l i z a t i o n o f a volum e e s t i m a t e d t o b e e q u i v a l e n t t o a n a v e r - .
age l e a d e r .
Food h a b i t s d a t a o f c a t t l e i n c l u d e d o n ly b ro w s e u s e i n t h e
w illo w and s e d g e t y p e s .
Food h a b i t s o f m oose i n f a l l w e re d e te r m in e d
from rum en s a m p le s from h u n t e r - k i l l e d m o o se.
Rumen a n a l y s i s was as
d e s c r i b e d by K n o w lto n (1 9 6 0 :1 6 3 ) e x c e p t m e a s u re m e n ts w e re t o t h e n e a r e s t
Oo5 Ce.
I n w i n t e r e a c h v e g e t a t i o n t y p e w as s u rv e y e d f o r f e e d i n g s i t e s
i n a p p r o x im a te p r o p o r t i o n t o i t s
o c c u r r e n c e on t h e s tu d y a r e a .
d a t a from t h e s p r u c e - f i r a n d D o u g l a s - f i r ty p e s w e re p a r t l y
im m e d ia te ly a d j a c e n t t o t h e s tu d y a r e a .
W in te r
from a r e a s
D a ta from f e e d i n g s i t e s and
rum en s a m p le s w e re c o m p ile d by t h e a g g r e g a t e volum e m eth o d ( M a r ti n , e t
at.
1946).
T he a d e q u a c y o f a sa m p le o b t a i n e d by t h e f e e d in g s i t e m eth o d can
b e d e te r m in e d w i t h a p l o t t e d c u rv e s i m i l a r t o t h e s p e c i e s : a r e a c u rv e
u s e d by p l a n t .e c o l o g i s t s
( C o s tin g 1 9 5 6 :4 4 - 4 7 ) .
The a c c u m u la te d t o t a l
o f i n s t a n c e s o f u s e on a l l p l a n t s p e c i e s i s p l o t t e d on t h e x a x i s .
The
p e r c e n t o f t h e a c c u m u la te d t o t a l o f i n s t a n c e s o f u s e t h a t i s a c c o u n te d
f o r by a n i m p o r ta n t s p e c i e s i s p l o t t e d on t h e y a x is ( F i g u r e 2 ) .
The
t h r e e p l a n t s p e c i e s w i t h t h e m o st u s e s h o u ld e a c h b e u s e d f o r p l o t t i n g
-9 -
Salix my rtillifolia
Salix g e y e r ia n a
B e tu la g l a n d u l e s a
22
26
30
34
26
30
34
38
S alix wolfii
Salix m y r ti ll if oli a
Salix g e y e r i a n a
'
Satix p la n ifo lia
Salix my r ti ll if oli a
Salix b e b b i a n a
18
Total
A ccumulated
22
Instances
of
Us e
(x 1 0 0 0 )
F i g u r e 2„
I n s t a n c e s - o f - u s e c u rv e s f o r d e te r m in in g t h e a d e q u a c y o f a
sa m p le o b t a i n e d by t h e f e e d i n g s i t e m ethod o f fo o d h a b i t s
a n a l y s i s : A, m oose i n summ er; B , c a t t l e i n sum m er; C,
m oose i n w i n t e r .
—1 0 —
a s e p a r a t e curve=
T h is m in im iz e s m i s i n t e r p r e t a t i o n s w h ic h a r e m ore
p r o b a b l e when o n ly one c u r v e i s u s e d .
c u rv e s l e v e l o f f .
The s a m p le . i s a d e q u a te when t h e
A r b i t r a r y v a l u e s , d e p e n d in g on t h e c o n f i d e n c e d e s i r e d ,
c a n b e s e l e c t e d f o r d e te r m in in g when t h e c u rv e s l e v e l o f f .
In g e n e ra l,
t h e s m a l l e r t h e a v e r a g e num ber o f i n s t a n c e s o f u s e p e r f e e d i n g s i t e , t h e
m ore dem anding s h o u ld b e t h e c r i t e r i a .
A n .e x a m p le i s :
no m ore t h a n a
3 p e r c e n t c h a n g e w i t h a n ad d e d 5 ,0 0 0 i n s t a n c e s o f u s e f o r f e e d i n g s i t e s
a v e r a g in g a b o u t 1 ,0 0 0 i n s t a n c e s o f u s e . ■ T h is p o i n t c a n b e d e te r m in e d
i n t h e c a l c u l a t i o n s w i t h o u t p l o t t i n g a c u rv e (,K orschgen 1 9 6 3 :3 2 0 ) , b u t
a c u r v e h e l p s t o m in im iz e m i s i n t e r p r e t a t i o n s o f c a l c u l a t e d r e s u l t s by
p r o v id i n g a c o n ti n u o u s , d e v e lo p in g p a t t e r n w h ic h c a n b e e x a m in ed v i s u a l ­
l y , and i t a l s o f a c i l i t a t e s
d a ta t h a t a re n eed ed .
t h e e s t i m a t i o n o f t h e am ount o f a d d i t i o n a l
I f .f o o d h a b its d a ta a r e s e g re g a te d by v e g e ta tio n
ty p e , c u rv e s a re n eeded f o r each ty p e .
The b a s i c
u n it o f in s ta n c e s o f.
u s e on t h e x a x i s s h o u ld b e a b b u t t w i c e t h e a v e ra g e num ber o f i n s t a n c e s
o f u se p e r fe e d in g s i t e .
I n . t h i s s tu d y t h e a v e r a g e s , f o r t o t a l u s e ,
w e re 920 f o r m oose i n sum m er, 594 f o r c a t t l e , and 928 f o r m oose i n
w in te r.
The b a s i c u n i t s s e l e c t e d w e re 2 ,0 0 0 , 1 ,0 0 0 , a n d 2 ,0 0 0 i n s t a n c e s
o f u s e , r e s p e c t i v e l y . , S m a ll i n t e r v a l s b e tw e e n t h e u n i t d i v i s i o n s on t h e
x a x is and a l o n g ■y a x i s , w i t h i n l i m i t s , g iv e t h e b e s t c u r v e .
S e x a n d Age R a t i o s ,
P o p u l a t i o n T r e n d s , a n d Movements
D a ta on s e x and a g e r a t i o s , p o p u l a t i o n t r e n d s , and m ovem ents w e re
r e c o r d e d w i t h d a t a on t h e u s e o f v e g e t a t i o n t y p e s .
F our a e r i a l su rv e y s
w e re flo w n i n f ix e d - w in g a i r c r a f t t o s u p p le m e n t t h e s e d a t a .
N e a r ly a l l
- 1 1 -
m oose t h a t w e re p r e s e n t w e re p r o b a b ly o b s e rv e d d u r in g t h e a e r i a l s u r ­
v e y s e x c e p t t h o s e i n t h e c o n i f e r o u s t im b e r ty p e s =
S te v e n s '
(1 9 6 5 :3 1 ) o b s e r v a t i o n s »
T h is a g r e e s w ith
Y e a r l i n g s w e re c o n s id e r e d a d u l t s
and i n d i v i d u a l m o o se ,w e re n o t k n o w in g ly r e c o r d e d m o r e .t h a n o n c e d a i l y .
M ost cows' w i t h o u t c a lv e s w e re s e x e d by t h e p r e s e n c e o f a . l i g h t c o lo r e d p a t c h o f h a i r a ro u n d t h e v u l v a . - T h is m e th o d , p r e v i o u s l y u se d
by r e f u g e p e r s o n n e l , H o u s to n ( 1 9 6 8 :7 ) , an d o t h e r s , app’e a r s t o b e r e ­
lia b le in th is a re a .
I n a l l g ro u n d o b s e r v a t i o n s , o n ly one cow was n o te d
w i t h a n o b s c u r e p a t c h and no b u l l s w e re o b s e r v e d w i t h a p a t c h . ,
RESULTS AND CONCLUSIONS
V egetation
Q u a n tita tiv e r e s u l t s o f th e v e g e ta tio n a n a ly s is a re p re s e n te d in
T a b le s I I I an d IV and i n T a b le s IX -X V III i n t h e a p p e n d ix .
v e g e t a t i o n t y p e s w e re d i f f e r e n t i a t e d
E ig h t
(F ig u re I ) .
A quatic T ype.
T h is t y p e w as m a in ly i n U pper Red Rock L a k e .
I t was d o m in a te d by
Potamogeton T ich a rd so n ii3 Chara s p . ■( f o l lo w s H o tc h k is s 1 9 6 7 :5 2 ) , and
MyriophyVlvm spioatmrio
R e s p e c ti v e v o lu m es w e re 3 6 , 3 3 , and 15 p e r c e n t
o f t h e t o t a l an d r e s p e c t i v e f r e q u e n c i e s w e re 7 0 , 4 6 , a nd 28 p e r c e n t
(U= S= D e p a rtm e n t o f I n t e r i o r 1968)=
T h is ty p e c o m p rise d a b o u t 15
p e r c e n t o f t h e s tu d y a rea=
Sedge Type
T h is ty p e was i n w e t lo w la n d s a n d w as d o m in a te d by Carex sp p =
( m o s tly C=
T ostrata3 CL Simulata3 and CL nebraskensis) , C la s s M usci
( f o l lo w s J a q u e s 1 9 4 9 :1 6 0 ) , and
Juncus balticus*
I t su cceed ed th e
a q u a t i c t y p e and c o m p ris e d a b o u t 35 p e r c e n t o f t h e s tu d y a r e a .
Willow Type
T h is t y p e was i n w e t lo w la n d s and a lo n g s tr e a m s .
I t w as d o m in a te d
by S a lix myT t i l l i f o l i a 3 S 0 p l a n i f o lia 3 CL W o lfii3 SL geyeriana3 and
B etu la glandulosa i n t h e s h ru b g ro u p and b y Carex s p p = , C la s s M u s c i,
and Junous b a ltie u s i n t h e h e rb g ro u p =
T h is ty p e s u c c e e d e d t h e s e d g e
ty p e and c o m p ris e d a b o u t 15 p e r c e n t o f t h e s tu d y a r e a .
Spruce Type
T h is ty p e w as i n w e t lo w la n d s and w as d o m in a te d by Pieea engelmanni3
Linnaea b o r e a lis 3 and Cornus canadensis i n t h e t r e e and s h ru b g ro u p s and
-1 3 -
TABLE I I I .
INTERCEPT, CANOPY-COVERAGE, AND FREQUENCY OF IMPORTANT PLANTS
ON THE STUDY AREA BY VEGETATION TYPE. A SINGLE VALUE INDI­
CATES PERCENT INTERCEPT AND TWO VALUES SEPARATED BY A SLASH
INDICATE PERCENT CANOPY-COVERAGE AND FREQUENCY, RESPECTIVELY.
T DENOTES LESS THAN 0„5 PERCENT AND + DENOTES PRESENT BUT
NOT OCCURRING IN PLOTS.
T axon
SE
—
WI
T.
+
Abies lasiooarpa
Pioea engelmanni
Pinus o o n to rta
Pseudotsuga m e n z ie s ii
Popuius trem uioides
+
15
S a tix m y r t i t t i f o l i a
S a lix p l a n ifo lia
I
14
I
9'
S a lix w o l f i i
T
S a lix geyeriana
5
B etu la glandulosa
I
5
T
Linnaea b o r e a lis
Cornus oanadensis
Symphorioarpos o reo p h ilu s Spiraea b e t u l i f o l i a
Clem atis Columbiana
B erb eris repens
S a lix bebbiana
T
I
+
T
S a lix drummondiana
6 6 /1 0 0 5 2 /9 8
Garex s p p . 2 /
C la s s Musc l —
3 5 /6 2
1 5 /4 6
Junous' b a ltio u s
A ste r fo lia o e u s
6/34
Equisetum arvensesJ
Fragaria v irg in ia n a
Geranium rio h a rd so n ii
Aotaed rubra
Sfnilaoina s t e l l a t a
G lyo etia e la ta
T /4
Taraxacum s p p ,
1/20
Thaliotrum venulosum
1 /1 0
T rifo liu m longipes
Calamagrostis rubescens f —
Lupinus afrgenteus .
v a r . p a r tiflo r u s — .
Geranium viscosissim um
Arnica o o r d ifo lia
2 0 /3 9
5 /2 1
1 /1 4
1/26
1 /6
T /7 ■
+
2 /1 3
+
1 /1 7
1 /6
1 /6
—
-
VEGETATION TYPE l /
AS
SP
DF
+
. 2
I
I
49
—
12
7"
71
+
+
75
T
I
I
T
+
+
+ .
+
13
4
-
+
3
T
I
2
40/86
27/66
T /3
+
1 8 /5 6
1 0 /5 0
2
I
1 /2 2 .
1 /6
4*
5 /1 2
5 /3 2
9/43
8/20
7 /3 0
7 /4 0
6/43
6/20
3/32
3 /2 0
1 /7
-
-
“
1 3 /7 6
1 1 /4 4
9/26
8/38
7/40
7/34
6/24
2
6
3
T
+
1 /5
2/25
1 /8
+
6 /4 5
+
T /3
6 /3 5
2 5 /8 0
' SF
60
28
—
19
T
T
3
GR
—
+
+
T /2
-
+
T /9
T /2
T /2
2/20
4 /3 0
6/60
T/1 0 .
—
—
T/1 0
1 1 /8 7
1 2 /8 0
1 /1 7 •
1 0 /6 0
6
3 /2 0
-
1 /1 0
2 /1 3
T /1 0
-
22/85
4/60
—
1 /2
-
—1 4 -
TABLE I I I .
(CONTINUED)
SE
WI
T /1 2
+
T /l
1 /1 2
T axon
Phleum p v a te n s e ■
A stev oonspiouus
Ligustioum filio in u m
Festuoa ovina
Lupinus s p p „
M iovosevis nigvesoens
Besseya wyomingensis
Galium boveale
T /2
VEGETATION TYPE
SP
AS
DF
+
T /?
5/32
2/6
1 /8
+
2 /3 0
—
7 /1 8
+
1 /1 0
SF
-
1 1 /7 0
7 /3 0
1 /4 0
GR
+
“
1 6 /7 3
9/68
6 /5 4
5/56
5 /3 9
JL/ SE = , s e d g e , WI = w i l lo w , SP = s p r u c e , AS = a s p e n , DF = D o u g l a s - f i r ,
SF = s p r u c e - f i r , and GR = g r a s s l a n d .
_2/ F o llo w s J a q u e s ( 1 9 4 9 :1 6 0 ) .
3 / F o llo w s D a v is ( 1 9 5 2 :5 1 ) .
4 / F o llo w s H itc h c o c k and C r o n q u is t ( 1 9 6 1 :3 0 2 ) .
by Cavex s p p . , C la s s M u s c i, and Equisetum avvense ( f o l lo w s D a v is 1 9 5 2 :5 1 )
i n t h e h e rb g r o u p .
I t s u c c e e d e d t h e w illo w and a s p e n ty p e s a n d c o m p rise d
l e s s t h a n I p e r c e n t o f t h e s tu d y a r e a .
Gvass land Type
T h is ty p e was on d ry t o m o d e r a te ly m o is t u p la n d s and w as d o m in a te d
by Festuoa Ovina3 Taraxacum s p p . , and Lupinus s p p .
An o v e r s t o r y o f
A vtem isia tv id e n ta ta a n d /o r Chvysothamnus s p p . was p r e s e n t i n some a r e a s .
T h is ty p e s u c c e e d e d t h e s e d g e and w illo w t y p e s and c o m p ris e d a b o u t 20
p e r c e n t o f t h e s tu d y a r e a .
Aspen Type
T h is ty p e was on m o d e r a te ly m o is t u p la n d s and i n w e t .l o w l a n d s .
It
-1 5 -
was d o m in a te d b y Poputvs ^ e m u lo id e s 3 Symphoricarpos O reophilus3 and
B erb eris repens i n t h e t r e e and s h ru b g ro u p s and by Taraxaovm s p p 0 ,
T halictrum Venulosvm3 and T rifo liu m longipes i n t h e h e rb g r o u p „
T h is
t y p e s u c c e e d e d t h e w illo w t y p e i n t h e lo w la n d s and t h e g r a s s l a n d ty p e
i n t h e u p la n d s and c o m p ris e d l e s s t h a n I p e r c e n t o f t h e s tu d y a r e a .
D o u g la s-fir Type ■
T h is t y p e was on m o d e r a te ly m o is t u p la n d s m o s tly b e lo w 7 ,5 0 0 f e e t „
I t was d o m in a te d by Pseudotsuga m e n z ie s ii3 Spiraea b e tu li: fo liq 3 and
■.;
Clem atis Columbiana i n t h e t r e e and s h ru b g ro u p s and by Calamagrostis
riibeseens3 Arnica O o rd ifo lia 3 and A s te r eonspieuus i n t h e h e r b .g ro u p .
T h is ty p e s u c c e e d e d t h e a s p e n and g r a s s l a n d t y p e s and i s ; c o n s id e r e d
t h e c lim a x on d r i e r s i t e s m o s tly b e lo w 7 ,5 0 0 f e e t ( P a t t e n 1 9 6 3 :3 7 8 ,
386, 3 9 8 )o
I t c o m p ris e d a b o u t 10 p e r c e n t o f t h e s tu d y a r e a .
S p r u c e - fir .Type
T h is , type, w as on w e t u p la n d s and w as d o m in a te d on t h e s tu d y a r e a
by Abies Ia sioearpa3 B erb eris repens3 an d Clematis columbiana i n th e
t r e e an d s h ru b g ro u p s and by T halietrvm Venulosvm3 A ste r eonspicuus3
and Calamagrostis rvbeseens i n t h e h e r b g r o u p .
T h is t y p e i s c o n s id ­
e r e d t h e c lim a x on m o is t s i t e s m o s tly a b o v e .7 ,5 0 0 f e e t (D au b en m ire
1 9 4 3 , P a t t e n 1 9 6 3 :3 7 8 , 3 9 7 - 3 9 8 ) .
I t c o m p ris e d l e s s t h a n I p e r c e n t o f
t h e s tu d y a r e a .
,
•
. . .
.
-
.
•••
D a ta on w illo w h e i g h t s a r e p r e s e n t e d i n 'T a b l e TV and i n F i g u r e 5
i n t h e a p p e n d ix .
The a v e r a g e h e i g h t o f a w illo w s p e c i e s w as r e l a t e d
t o i t s m o st common h a b i t a t w i t h r e s p e c t t o s o i l m o i s t u r e , m i c r o c l i m a t e j
, ~
-1 6 -
TABLE IVo
HEIGHTS AND FREQUENCY INDICES OF WILLOWS ON. THE STUDY .AREA.
Sam ple
S iz e
S p e c ie s
S.
S.
Sc
Sc
Sc
Sc
Sc
Sc
Sc
Sc
Sc
Sc
Candida
w o lfi i
e x ig u a ■
v ig id a
p la n ifo lia
drurmondiana
m onticola
m y r tillifo lia
lasiandra
bebbiana
geyeriana
sco u leria n a
50
100
50 •
10
100
100
50 •
100
35
100
100
20
A v e ra g e
2 .2
2 .9
3 .4
3 .5
3 .7 '
4 .9
5 .3 '
5 .6
5 .7
6 .2
7 .8
1 9 .6
HEIGHT ( f t . )
Maximum
Minimum
F re q u e n c y
In d e x i/
5 .0
5 .5
7 .0
7 .0
1 3 .5
9 .0
1 4 .0
1 4 .0
1 9 .5
1 8 .0
1 .0
1 .0
1 .0
2 .0
1 .0
2 .0
1 .5
1 .5
2 .0
1 .5
14
80
10
10
100
22
14
90
5
23
1 8 .5
4 0 .0
2 .0
3 .0
90
-
I / S ee t e x t , p a g e 5 , f o r e x p l a n a t i o n .
o r b o t h . ( F ig u r e 3 ) .
G e n e r a l l y , t h e s h o r t e s t s p e c i e s w e re i n a r e a s w i t h
t h e m o st m o i s t u r e and t h e g r e a t e s t t e m p e r a t u r e e x tre m e s w h i l e t h e t a l l ­
e s t s p e c i e s w e re i n a r e a s w i t h t h e l e a s t m o i s t u r e and t h e l e a s t te m p e ra ­
t u r e e x tre m e s d u r in g t h e g ro w in g s e a s o n .
re la tio n s h ip .
fe e t.
S tre a m s a p p a r e n t l y u p s e t t h i s
A l l s p e c i e s w e re w i t h i n an e l e v a t i o n a l r a n g e o f 150
Sc rigi-da was to o r a r e f o r e v a l u a t i o n .
T h is h e i g h t - h a b i t a t r e l a t i o n s h i p p a r a l l e l s th e d w a r fin g o f w illo w s
w i t h i n c r e a s i n g e l e v a t i o n and l a t i t u d e .
T he w e t , m ossy b o g s ro u g h ly
c o r r e s p o n d t o s u b a l p i n e s l o p e s i n t h e m o u n ta in s and t h e lo w e r t u n d r a
in th e n o r th .
Raup ( 1 9 5 9 :2 0 ) , r e f e r r i n g
t o w illo w s i n b o r e a l w e s t e r n
A m e ric a , s t a t e d t h a t m o st p r o s t r a t e w illo w s grow i n t h e a r c t i c o r
G r e a t e s t T e m p e r a t u r e E x t r e m e s — „ — — -----------------------------------------— _
We t M o s s y Bog
M o d e r a t e l y We t L o w l a n d
S. oandida • SL w o lfi-i
2=2
2 .9
S= p ta n ifo lia
3=7
L e a s t T e m p e ra tu re Extrem es
D r i e r Lowland Edges
Dr y U p l a n d F o r e s t
S= m onticola
S= bebbiana
-S= scouleviana
5 .3
6 .2
1 9 .6
S. m y r t it l i f o l i a
S=. geyeriana
5 .6
'
7=8
S t r e a m Bank G roup
F ig u re 3.
S= exigya
3.4-
S= dvimmondiana
4 .9
S= Iasiandva
5 .7
M ost common h a b i t a t o f w illo w s p e c i e s w i t h r e s p e c t t o s o i l m o i s t u r e and m ic r o c l im a te .
The num bers a r e a v e r a g e h e i g h t s o f t h e s p e c i e s i n f e e t .
-1 8 -
a lp in e
t u n d r a , m o st m e d iu m -s iz e d s p e c i e s a r e i n s w a le s o r a lo n g s tr e a m s
i n t h e t u n d r a o r on o p e n l a n d o f f o r e s t a r e a s , and m o st t a l l s p e c i e s a r e
i n f o r e s t s o r on t h e m a r g in s o f f o r e s t s .
T h is r e l a t i o n s h i p i s a p p a r e n t l y
o n ly e v i d e n t u n d e r n e a r l y i d e a l c o n d i t i o n s b e c a u s e o f t h e many f a c t o r s
in flu e n c in g th e h e ig h ts .
P a t t e n (1 9 6 8 :1 1 1 2 ) d i s c u s s e d t h e e f f e c t s o f
b ro w s in g o n . s e v e r a l s p e c i e s o f w illo w .
Many o f t h e f a c t o r s . l i s t e d by
H anson (1 9.53:139) a s i n f l u e n c i n g t h e d e v e lo p m e n t, m a in te n a n c e , o r
c h a n g e s o f . t h e s p e c i e s c o m p o s itio n w i t h i n v e g e t a t i o n t y p e s a l s o i n ­
f l u e n c e w illo w h e i g h t s o . U n d er l e s s t h a n i d e a l c o n d i t i o n s , d w a rfin g , i s
m o st a p p a r e n t w i t h i n c r e a s i n g l a t i t u d e , som ew hat l e s s a p p a r e n t w i t h i n ­
c r e a s i n g e l e v a t i o n , and l e a s t a p p a r e n t on a s m a ll a r e a .
The s m a l l e r t h e
a r e a , th e . m ore p ro n o u n c e d a r e t h e e f f e c t s o f ; i n f l u e n c i n g f a c t o r s i n
m a sk in g t h e h e i g h t r e l a t i o n s h i p .
The h i g h m o u n ta in v a l l e y w i t h s i g n i f i ­
c a n t te m p e ra tu re changes o v er a s m a l l.a r e a , a g ra d u a l change i n s o i l
m o i s t u r e , a v a r i e t y o f w illo w s p e c i e s , m o d e r a te b r o w s in g , and o t h e r
f a c t o r s p r o v id e d n e a r l y i d e a l c o n d i t i o n s f o r e x p r e s s i o n o f t h i s h e i g h t h a b i t a t r e l a t i o n s h i p on t h e s tu d y a r e a .
The m o st common h a b i t a t ( F ig u r e 3) d o e s n o t r e f l e c t t h e r a n g e o f
d is trib u tio n .
In g e n e ra l, 5 .
Oandida3 S . - Scoiileviana3 and t h e s tr e a m
b a n k g ro u p w e re m o s tly r e s t r i c t e d t o m ossy b o g s , u p la n d f o r e s t s , and
s tr e a m b a n k s , r e s p e c t i v e l y . .
tw e e n t h e p o s i t i o n s o f So
b a n k p o s i t i o n ( F ig u r e 3 ) .
The r e m a in in g s p e c i e s m o s tly r a n g e d b e ­
w o lfii and S. geyeviana3 i n c l u d i n g t h e s tr e a m
-1 9
V e g e t a t i o n T y p e U se
O b s e r v a tio n s o f m oose f e e d i n g o r b e d d e d i n summer an d w i n t e r t o t a l e d
581 and 4 1 3 , r e s p e c t i v e l y , • O b s e r v a tio n s i n t h e w illo w t y p e a c c o u n te d f o r
84 p e r c e n t o f t h e t o t a l i n summer a n d .93 p e r c e n t i n w i n t e r ( T a b le V )„
.
U se o f t h e t im b e r t y p e s w as p r o b a b ly m o re i n summer - t h a n t h e d a t a i n d i ­
c a te .
S ev en p e r c e n t o f a l l o b s e r v a t i o n s w e re o f m oose m oving t o and
from t h e D o u g l a s - f i r t y p e i n t h e m o rn in g and e v e n in g , r e s p e c t i v e l y , s u g ­
g e s t i n g m ore u s e o f t h i s t y p e t h a n o b s e r v e d .
Use o f t h e s e d g e and g r a s s ­
lan d . t y p e s was p r o b a b ly l e s s i n s u m m e r-th a n t h e d a t a i n d i c a t e due t o
e x c e lle n t v i s i b i l i t y
in th e s e ty p e s . .
The d i f f e r e n c e s i n t h e u s e o f t y p e s b e tw e e n summer an d w i n t e r w e re
m a in ly due t o snow c o v e r i n f l u e n c i n g a v a i l a b i l i t y o f f o r a g e an d v i s i b i l i ­
ty .
Food h a b i t s p r i m a r i l y d e te r m in e d t h e u s e o f v e g e t a t i o n t y p e s w h ich
e x p l a i n s t h e im p o r ta n c e o f t h e w illo w t y p e .
D iffe re n t a re a s have
d i f f e r e n t p a t t e r n s o f u s e b e c a u s e fo o d h a b i t s u s u a l l y d i f f e r by a r e a .
TABLE V.
USE OF VEGETATION TYPES BY MOOSE FOR FEEDING AND BEDDING.
NUMBER OF OBSERVATIONS IS IN PARENTHESES.. T DENOTES LESS
THAN 0 .5 PERCENT.
V e g e t a t io n
Type
W illow
A spen
Sedge
G r a s s la n d
A q u a tic
D o u g la s -fir
S p ru c e
S p ru c e -fir
Total
S tm m er(S S l)
PERCENT USE
'
W in te r( 4 1 3 )
84
6
6
3
■T
T
93
3
I
0
T
T
T
3
T
T
1 00
99
-2 0 -
The d i s t r i b u t i o n o f m oose w i t h i n t h e w illo w ty p e d i f f e r e d b e tw e e n
summer and w i n t e r ( F ig u r e 6 i n t h e a p p e n d i x ) „- T h is may h a v e b e e n due
t o d i f f e r e n c e s i n a v a i l a b i l i t y o f f o r a g e , c o v e r r e q u i r e m e n t s , re m a in in g
i n t h e f i r s t s t a n d o f s u i t a b l e w illo w s e n c o u n te r e d a f t e r l e a v i n g tim ­
b e r e d a r e a s i n w i n t e r , o r a c o m b in a tio n o f t h e s e .
T h is d i f f e r e n c e -i n
d i s t r i b u t i o n h e lp e d r e d u c e t h e b ro w s in g p r e s s u r e on t h e w i n t e r f o r a g e
s u p p ly .
Moose w e re so m e tim e s o b s e rv e d b e d d e d on t h e i c e o f U pper Red Rock
L ake p o s s i b l y t o t a k e a d v a n ta g e o f s o l a r r a d i a t i o n .
T h is was con­
s i d e r e d u s e o f t h e a d j a c e n t w illo w t y p e .
Food H a b i t s
M oose-sum er
B row se a c c o u n te d f o r 9 8 .3 p e r c e n t o f a l l f o r a g e u s e d b y m oose i n
summer ( T a b le V I ) .
K h o w lto n (1 9 6 0 :1 6 6 ) and H o u sto n ( 1 9 6 8 :2 6 ) r e p o r t e d
p e r c e n t a g e s o f 29 and a b o u t 75 p e r c e n t , r e s p e c t i v e l y .
T he a b u n d a n c e i n
q u a n t i t y and v a r i e t y o f w illo w s i n t h i s s tu d y l a r g e l y a c c o u n te d f o r t h e
h e a v y u s e o f b ro w s e .
S a lix m y v t i l l i f o l i a 3 B etu la glandulosa3 S a lix Qeyeriana3 and
S a lix 'p la n ifo lia a c c o u n te d f o r 5 8 .1 , 1 1 .8 , 9 . 6 , and 6 .7 p e r c e n t , r e s p e c ­
t i v e l y , o f a l l f o r a g e u s e d ( T a b le V I ) .
O th e r i n v e s t i g a t o r s fo u n d v a r y ­
in g d e g r e e s o f u s e o f o n e o r m ore o f t h e s e s p e c i e s e x c e p t 51. p la n ifo lia
(M cM illan 1 9 5 3 :1 0 3 , D e n n is to n 1 9 5 6 :1 0 6 , P e e k 1 9 6 3 :2 2 8 -2 2 9 , H o u sto n
1 9 6 8 :2 7 ,1 0 1 )„
•TABLE V I.
FOOD HABITS OF CATTLE AND MOOSE. VALUES ARE PERCENTAGES. T DENOTES LESS THAN 0 .0 5 PERCENT AND NA DENOTES NOT AVAILABLE.
PLANTS WITH LESS THAN 2 PERCENT USE (EXCLUDING THE GENUS SALIX) IN ALL CATEGORIES EXCEPT FREQUENCY FOR RUMEN SAMPLES
APPEAR IN TABLE XIX IN THE APPENDIX.
SUMMER
Moose
C a ttle
A re a o f U s e ^ :
A l 30
Ca 19
A l 40
As 8
I n s t a n c e s o f U se :
1 7 ,8 1 3
NA
NA
2 .7
0 .0
0 .0
NA
NA
0 .0
1 0 .9
0 .5
1 2 ,2 7 1 NA
NA
5 .1
2 .0
T
NA
NA
0 .0
2 .0
0 .0
3 6 ,8 1 2
0 .0
0 .0
1 1 .8
3 .3
T
0 .4
0 .0
0 .1
2 .6
0 .0
2 ,2 2 1 - '
0 .0
0 .0
0 .0
4 2 .3
0 .0
6 .7
0 .0
1 .5
1 7 .9
NA
A s t e r e o n sp ie u u s
L upinus a r g e n te u s .
v a r . a r g e n te u s—
T o ta l F o ro s - ^
2 .8
0 .1
9 .6
0 .2
0 .0
0 .8
5 8 .1
0 .0
6 .7
0 .1
T
0 .1
~
0 .1
NA
NA
NA
NA
NA
NA
1 5 .4
NA
1 2 .5
0 .0
0 .0
0 .0
0 .9
1 0 0 .0 1 /
9 5 .4
9 8 .3
-
NA
0 .0
4 .5
“3/
G ram in eae and Carex s p p .
T o ta l G r a s s , G r a s s - I i k e ^
-J l
-
Wi
-
Ti I^
A l 35
Wi 24
V o l. F r e q . V o l.
7 .1
0 .2
14
NA
~
0 .0
0 .0
0
NA
0 .0
29
1 .1
0 3 4 .6
0 .0
NA
0 .0
NA
0 .0
0 .0
0
0 .0
-
3 2 ,4 8 0
2 .5
2 .2
0 .7
3 .1
1 .5
0 .3
0 .6
1 .7
1 5 .4
T
2 6 ,7 4 8
NA
NA
0 .9
0 .0
0 .0
NA
NA
T
1 7 .5
T
3 ,4 8 7 —
1 7 .1
2 0 .2
NA
1 1 .9
1 4 .3
0 .5
4 .8
1 .5
8 .7
NA
0 .9
0 .1
1 0 .5
2 .0
0 .7
2 5 .0
1 .8
2 4 .2
0 .0
1 .9
2 .5
1 .1
0 .8
1 .0
0 .1
1 2 .8
2 .4
0 .8
3 0 .4
NA
2 9 .4
0 .0
NA
3 .1
1 .3
NA
NA
NA
NA
NA
0 .0
0 .0
NA
NA
NA
1 7 .1
NA
0 .3
3 .4
NA
NA
NA
NA
NA
0 .0
NA
0 .0
0 .0
1 .5
0 .0
8 .2
-
7-
-
-
_
Df 9
As 5
1 ,6 2 7 - '
1 0 .9
0 .6
0 .0
3 6 .2
0 .0
4 .7
1 .7
3 0 .1
2 .2
NA
9 9 .0
8 5 .2
0 .1
1 .9
9 5 .5
:
100
13
38
100
9 6 .6
NA
0 .8
9 9 .6
100
29
100
:
2 2 .2
0 .4
7 .5
7 2 .2
9 9 .8
9 9 .8
io o T o
1 0 0 .0
0 .0
3 .8
13
NA
_
2 4 .8
0 .0
NA
0 .0
0 .0
1.5
0 .0
0 .0
0
0 .0
0
0 .0
0 .0
0 .0
NA
0 .0 .
4 .5
1 .5
1 .0
3 .8
50
0 .1
43
2 4 .8
T
T
0 .1
_
0 .7
75
0 .3
71
3 .0
0 .2
T
0 .7
75
0 .3
71
3 .0
0 .1
0 .2
0 .0
IO
O
U n i d e n t i f i e d b ro w se
T o ta l B ro w se* /
6 .3
0 .2
2 1 .0
NA
NA
2 .4
4 5 .6
NA
1 0 .3
NA
NA
0 .0
~
NA
2 .9
0 .3
1 1 .2
NA
NA
0 .3
1 5 .9
NA
4 .8
NA
NA
5 0 .2
NA
^
O
O
A b ie s la s io c a r p a
A eer g l a b r m
B e tu la g la n d u lo s a
P opulus tre m u lo id e s
P opulus tr e m u lo id e s ( b a r k )
Prunus v ir g in ia n a
P seu d o tsu g a m e n z ie s ii
Rosa w o o d sii
S a l i x b e bbiana
S a l i x oandida
S a l i x dm cm ondiana
S a l i x e x ig u a
S a l i x g e y e r ia n a
S a l i x g la u o a l'
S a trIx VasrIandva
S a l i x m o n tie o la
S a lix m y v t i l li f o li a
S a l i x pentandva=-'
S a lix p la n ifo lia
S a lix r ig id a
S a l i x s e o u le r ia n a
S a lix w o lfi i
S a lix sp p .
Sym phorioarpos o r e o p h iln s
Al
V o l. F r e q ..
1 .3
25
0 .0
0
0
0 .0
0 .9
25
13
5 .3
0 .0
0
0 .0
0
0 .0
0
-
WINTER
Moose
O I
T axon
FALL
M oose _ #
I / A l = a l l a r e a s , Ca - c a t t l e a r e a s , As = a s p e n t y p e , Wi = w illo w t y p e , T i = u p la n d t i m b e r t y p e s , and Df = D o u g l a s - f i r t y p e .
Numbers a r e t o t a l f e e d i n g s i t e s f o r summer an d w i n t e r and t o t a l rum en sa m p le s f o r f a l l .
2 / I n s u f f i c i e n t sa m p le .
3 / R e co rd ed i n a n a r e a a d j a c e n t t o t h e s t u d y a r e a .
4 / I n c l u d e s p l a n t s i n T a b le XIX i n t h e a p p e n d ix .
5 / C a t t l e u s e on non-w oody p l a n t s w as n o t r e c o r d e d .
J i/ F o llo w s H itc h c o c k and C r o n q u is t ( 1 9 6 1 :3 0 6 ) .
I
No
V
-2 2 -
F o r a g e u s e i n t h e w illo w ty p e was s i m i l a r t o o v e r a l l u s e .
The
s a m p le i n t h e a s p e n ty p e was n o t s u f f i c i e n t f o r r e l i a b l e p e r c e n t a g e s ,
b u t th e d a ta s u g g e st p o s s ib le p ro p o rtio n s o f u se .
Populus tvem uloides
was m o st im p o r ta n t f o llo w e d by t h r e e s p e c i e s o f w illo w
( T a b le V I ) .
M o o se-fa ll
T h re e rum en s a m p le s from h u n t e r - k i l l e d m oose w e re c o l l e c t e d O c to b e r
1 9 , 1958 (K n o w lto n 1 9 6 0 :1 6 7 ) , one was c o l l e c t e d November 2 1 , 1959 by
M ontana F i s h an d Game D e p a rtm e n t p e r s o n n e l , and I c o l l e c t e d f o u r on
O c to b e r 2 0 , 1968„
T h is s a m p le was n o t s u f f i c i e n t (H anson a n d G r a y b i l l
1956) , b u t i t d o e s s u g g e s t p o s s i b l e fo o d h a b i t s i n f a l l .
The u s e o f
b ro w s e i n a l l v e g e t a t i o n t y p e s was 9 5 .5 p e r c e n t o f t h e t o t a l u s e (T a b le
V I) com pared t o o v e r 85 p e r c e n t i n o t h e r s t u d i e s
S te v e n s 1 9 6 7 :1 3 , H o u s to n 1 9 6 8 :2 6 ) .
(K no w lto n 1 9 6 0 :1 6 6 ,
S p e c ie s o f S a lix a c c o u n te d f o r 9 6 .6
p e r c e n t o f t h e u s e i n t h e w illo w t y p e .
One ru m e n :sa m p le r e p r e s e n t i n g
t h e u p la n d t im b e r t y p e s c o n ta i n e d o n ly 7 2 .2 p e r c e n t b r o w s e , m o s tly P .
tv e m u lo id e s b a r k and s p e c i e s o f S a l i x a
One f o r b , A s te r OonspiouiLSj
a c c o u n te d f o r 2 4 .8 p e r c e n t o f t h e t o t a l o f t h i s s a m p le .
M oose-winter
B row se a c c o u n te d f o r 9 9 .8 p e r c e n t o f a l l f o r a g e u s e d b y m oose i n
w i n t e r ( T a b le V I ) .
K n o w lto n ( 1 9 6 0 :1 6 7 ) , S te v e n s ( 1 9 6 7 :1 5 ) , and H o u sto n
(1 9 6 8 :2 5 ) r e p o r t e d s i m i l a r p e r c e n t a g e s .
S . myr t i l l i f o lia 3 S a p l a n i f o lia 3 S a bebbiana3 and S a geyeriana a c ­
c o u n te d f o r 2 5 .0 , 2 4 .2 , 1 5 .4 , and 1 0 .5 p e r c e n t , r e s p e c t i v e l y , o f a l l u s e .
-2 3 -
T h e s e p e r c e n t a g e s w e re 2 8 .4 , 1 7 .2 , 2 2 .6 , a n d 1 5 .0 , r e s p e c t i v e l y , d u r in g
t h e p e r i o d o f deep snow i n F e b r u a r y a n d .M a rc h .
The im p o r ta n c e o f t h e
t a l l s p e c i e s i n c r e a s e d w h i l e t h a t o f t h e s h o r t e s t s p e c i e s , S . PlccnifoZrIa,
d e c re a se d .
S', w o lfH w as i m p o r ta n t e a r l y i n t h e w i n t e r w h i l e i t was
a v a ila b le .
H o u sto n ' ( 1 9 6 8 :2 8 , 103) r e p o r t e d v a r y in g d e g r e e s o f u s e o f
a l l t h e s e s p e c i e s e x c e p t S c p la w ifO lia c
U se i n t h e w illo w ty p e w as s i m i l a r t o o v e r a l l u s e .
U se i n th e
D o u g l a s - f i r and a s p e n t y p e s was o n ly s u g g e s t i v e b e c a u s e o f t h e s m a ll
s a m p le s .
P . tvem uloides. an d Rosa w oodsii w e re im p o r ta n t i n t h e a s p e n
ty p e and P . fcvemuloides, Aoev gldbopvm, Abies lasiocarpcc, an d S a lix
sco u levia n a w e re i m p o r ta n t i n t h e D o u g l a s - f i r t y p e .
C a ttle-sim n er
So w o l f i i . So m y r t i l l i f o l i a , So geyeriana, and S c bebbiana a c ­
c o u n te d f o r 5 0 .2 , 1 5 .9 , 1 1 .2 , and 1 0 .9 p e r c e n t , r e s p e c t i v e l y , o f a l l
b ro w s e u s e d by c a t t l e i n summer ( T a b le V I ) .
r e p o r t e d a s u s e d by c a t t l e o r o t h e r l i v e s t o c k
A ll th e s e s p e c ie s have been
(Van D e r s a l 1 9 3 8 :2 4 5 -2 5 4 ,
H o u s to n 1 9 6 8 :4 5 ) .
Conolusions
'
;
A v a i l a b i l i t y w as o n e f a c t o r d e te r m i n in g t h e am ount o f v a r i o u s p l a n t
s p e c i e s i n t h e d i e t o f m o o se .
I n t h i s s t u d y S c p la n ifo lia was i m p o r ta n t .
I n J a c k s o n H o le i t a p p a r e n t l y w as n o t p r e s e n t (H o u sto n 1 9 6 8 ) .
Cornus
s to lo n ife r a w as i m p o r ta n t i n t h e d i e t o f m oose i n w i n t e r i n J a c k s o n H o le
(H o u s to n 1 9 6 8 :2 9 , 32) b u t w as n o t i m p o r ta n t i n t h i s s tu d y d u e t o i t s
lim ite d a v a i l a b i l i t y .
M oose w e re o b s e r v e d u s in g a q u a t i c s o n ly tw ic e i n
—2 4 —
t h i s s tu d y .
B o th p e r i o d s o f f e e d in g w e re b r i e f .
P e t e r s o n (1 9 5 5 :1 4 0 )
fo u n d t h a t Potamogeton r-iohavdsonii was t h e m o st im p o r ta n t a q u a t i c
fo o d i n a s tu d y i n O n t a r i o .
on t h i s s tu d y a r e a .
T h is s p e c i e s w as t h e m o s t ; a b u n d a n t a q u a t i c
Moose a p p a r e n t l y u s e few a q u a t i c s i f w illo w s a r e
a b u n d a n t.
Many f a c t o r s i n f l u e n c e a v a i l a b i l i t y .
s tu d y was s n o w c o v e r .
T he m o st i m p o r ta n t i n t h i s
E x t e n s i v e d r i f t i n g r e s u l t e d i n co m p lex i n f l u e n c e s .
D r i f t s c o v e re d some p l a n t s 10 f e e t h i g h , b u t b e c a u s e t h e s e d r i f t s
c o u ld
s u p p o r t a m o o se , th e y m ade a v a i l a b l e some p l a n t s o v e r 15 f e e t h i g h . .
M ost p l a n t s l e s s t h a n 4 f e e t h i g h w e re u n a v a i l a b l e by m i d - w i n t e r .
Win­
t e r u s e o f S . g ta n d u to s a and S'. W o t f t i 3 b o t h lo w -g ro w in g s h r u b s , was
lim ite d to e a rly w in te r.
S* b'ebbiana3 o f m in o r im p o r ta n c e i n summer
and e a r l y w i n t e r , becam e i m p o r ta n t i n l a t e w i n t e r p a r t l y th r o u g h an
in c r e a s e i n r e l a t i v e a v a i l a b i l i t y due to i t s
t a l l h e i g h t , ( F i g u r e 2 C ).
A d a p t a ti o n w as a s e c o n d f a c t o r d e te r m i n in g t h e am ount o f v a r i o u s
p la n t s p e c ie s in th e d i e t .
F l e r o v (1 9 6 0 :1 7 4 ) and M cM illan (.1953:109)
d i s c u s s e d t h e a d a p t a t i o n s o f m oose f o r b r o w s in g .
w h ic h m oose s t r i p
ta tio n .
The e f f i c i e n c y w ith
t h e l e a v e s from w illo w tw ig s i l l u s t r a t e s
o n e .a d a p ­
M ost u s e o f b ro w s e by moose, i n summer was by l e a f s t r i p p i n g .
T h is a d a p t a t i o n l a r g e l y a c c o u n te d f o r t h e l i g h t u s e o f S. w o t f i i by
m oose i n sum m er.
M ost l e a f s t r i p p i n g w as o v e r 3 f e e t a b o v e g ro u n d
l e v e l j a p p a r e n t l y t h e a r e a f o r m o st e f f i c i e n c y .
S'. W o tfii p l a n t s .
T h is was a b o v e m o s t.
S in c e u s e i n w i n t e r w as by n ip p in g l e a d e r s , o f t e n
c l o s e t o t h e snow l e v e l , /S', w o t f i i becam e i m p o r ta n t t o m oose w h i l e i t
-2 5 -
w as a v a i l a b l e b e c a u s e i t
s p e c ie s =
c o u ld b e a s . e f f i c i e n t l y u t i l i z e d
as o th e r
U se o f S c WoZfi-yL by c a t t l e i n summer was heavy=
C a ttle ,
b e in g a d a p te d f o r g r a z i n g , a r e i n e f f i c i e n t b r o w s e r s »
g ro w th o f So w o l f i i p u t i t i n a g r a z i n g p o s i t i o n =
T he low form o f
M c M illa n (1 9 5 3 :1 0 4 )
fo u n d t h a t m oose u s e d S 0 W oZfii l e s s t h a n So geyeviana i n sum m er'an d
s u g g e s t e d i t was p a r t l y due t o i t s
d i f f i c u l t t o brow se=
low fo rm o f g ro w th m ak in g i t m ore
He a l s o n o te d m ore u s e o f .So W oZfii by e l k
(.Cei1Vus canadensis n e tso n i) w h ic h a r e m o re a d a p te d f o r g r a z i n g th a n
moose=
P a l a t a b i l i t y w as a t h i r d f a c t o r d e te r m i n in g t h e am ount o f v a r i o u s
p l a n t s p e c i e s i n t h e d ie t=
I n t h e D o u g l a s - f i r ty p e i n . w i n t e r
Abies
Zasiocaxr^ a w as u s e d much m ore t h a n Pseudotsuga m e n zie s ii . w h ic h was -much ■
m ore
abundant
( T a b le s I I I and VI) =
U se o f A=
Zasiocavga w as l i m i t e d
t o c e r t a i n t r e e s w h ic h w e re o f t e n s e v e r e l y h e d g e d .
a l s o n o t e d t h i s p a t t e r n o f u s e o f A=
d e c ad e n t w illo w s , e s p e c ia lly
H o u s to n (1 9 6 8 :3 0 )
Zasiocavpao . C o n s i s t e n t u s e o f
So pZanifoZia and So bebbiana3 w i t h u se
o f g ro w th a s o ld a s 7 y e a r s , r a t h e r t h a n t h e p r e v i o u s y e a r 's g ro w th ,
was o b s e rv e d i n t h i s stu d y =
A f o u r t h f a c t o r d e te r m i n in g t h e am ount o f v a r i o u s .p l a n t s p e c i e s
i n t h e d i e t w as h a b i t (L e o p o ld 1 9 3 3 :2 5 5 -2 5 8 )=
A s u ita b le s p e c ie s w i l l
n o t b e u s e d u n l e s s an a n im a l h a s l e a r n e d t o u s e i t th r o u g h t r a i n i n g o r
e x p e r i e n c e (L e o p o ld 1 9 3 3 :2 5 8 , K en d eig h 1 9 6 1 :1 8 8 -1 8 9 ) .
a t i o n i s i n c l u d e d h e re =
fa c to r.
In d iv id u a l v a r i­
My d a t a w e re i n s u f f i c i e n t t o d e m o n s tr a te t h i s
—2 6 -
A v a i l a b i l i t y , was m o st i m p o r ta n t i n d e te r m in in g t h e c o m p o s itio n o f
th e d i e t f o llo w e d by a d a p t a t i o n and p a l a t a b i l i t y „
E v a lu a tio n o f th e
a b s o l u t e im p o r ta n c e o f i n d i v i d u a l p l a n t s p e c i e s was d i f f i c u l t b e c a u s e
t h e s e f a c t o r s w e re a l l i n t e r r e l a t e d .
I t a p p e are d t h a t a l l s p e c ie s of
w illo w t h a t w e re p r e s e n t i n t h i s s tu d y w e re p o t e n t i a l l y im p o r ta n t as
f o r a g e f o r m oose a t some tim e d u r in g t h e y e a r , e s p e c i a l l y i n w i n t e r .
R e l a t i v e l y low a v a i l a b i l i t i e s k e p t m o st s p e c i e s from b e in g i m p o r t a n t .
The p r o b a b l e m o st im p o r ta n t f a c t o r d e te r m in in g t h e am ount o f u s e f o r
f i v e i m p o r ta n t f o r a g e s p e c i e s o f m oose i n t h i s s tu d y i s p r e s e n t e d i n
T a b le V I I .
The g ro w th form o f S , T n y v tillifo lia p r o b a b ly a llo w e d i t t o
b e u s e d m ore e f f i c i e n t l y t h a n o t h e r s p e c i e s i n sum m er, t h e r e f o r e a c ­
c o u n tin g f o r i t s h e a v y u s e .
B e s id e s w i l l o w s , two o t h e r s p e c i e s w e re
p o t e n t i a l l y im p o r ta n t i n w i n t e r .
T h e se w e re P . tvem uloides and A.
Iasiooavpa^ They w e re i m p o r ta n t i n J a c k s o n H o le (H a rry 1 9 5 7 :5 5 ,
H o u sto n 1 9 6 8 :2 8 , 3 3 , 44) and P 0 tvem uloides was i m p o r ta n t i n p a r t s o f
A la s k a (S p e n c e r and C h a t e l a i n 1 9 5 3 :5 4 2 ).
TABLE V I I .
PROBABLE MOST IMPORTANT FACTORS DETERMINING THE AMOUNT OF
USE OF IMPORTANT FORAGE SPECIES OF MOOSE IN THIS STUDY.
FACTOR
T axon
S a lix m y v t i l l i f o l i a
S a lix p l a n i f o l i a
S a lix bebbiana
SarI ix geyeviana
B etu la glandulosa
Summer
A d a p ta tio n
A v a ila b ility
A v a ila b ility
A v a ila b ility
P a la ta b ility •
W in te r
A v a ila b ility
P a la ta b ility
A v a ila b ility
A v a ila b ility
A v a ila b ility
-2 7 -
M oose-C attle
C om petition
F o r a g e c o m p e t i t i o n (S m ith a n d i n l a n d e r 1 9 5 3 :1 0 1 ) b e tw e e n m oose a n d
c a t t l e was n o t s i g n i f i c a n t u n d e r t h e c o n d i t i o n s p r e v a i l i n g d u r in g t h i s
s tu d y .
S. Wotfi-Vi a c c o u n t in g f o r 5 0 .2 p e r c e n t o f a l l b r o w s e u s e d by
c a t t l e , was r a r e l y u s e d b y m oose e x c e p t i n e a r l y w i n t e r ( T a b le V I ) .
T he
a b u n d a n c e o f t h i s s p e c i e s a c t e d a s a b u f f e r t o r e d u c e t h e u s e by c a t t l e
o f S. myTt-Ltt-Vfotiai S. b e b b i a n a j
an d S. g e y e r i q n a .
C o m p e titio n w o u ld
b e e x p e c te d t o i n c r e a s e i n a r e a s w i t h a low a v a i l a b i l i t y
o r absence o f
S. w o t f i i o r w h e re m oose m u st' d e p e n d o n S. W o t f i i d u e t o a b s e n c e o f
o th e r w illo w s .
H o u s to n ( 1 9 6 8 :4 4 - 4 5 ) r e p o r t e d m o o s e - c a t t l e c o m p e t i t i o n
on S, m y r t i t t i f o t i a .
I n t h i s s tu d y c a t t l e u sed l i t t l e
b ro w s e b e f o r e t h e
m ore p a l a t a b l e f o r b s , g r a s s e s , a n d g r a s s - l i k e p l a n t s w e re l a r g e l y u t i ­
liz e d .
A le s s e r a v a ila b ility
o f t h e s e w o u ld i n c r e a s e t h e u s e o f b r o w s e .
M ost o f t h e b ro w s e u s e d by c a t t l e was l e s s t h a n 5 f e e t a b o v e g ro u n d
le v e l.
M ost o f t h i s w as c o v e r e d w i t h snow i n w i n t e r s o w as u n a v a i l a b l e
t o m o o se .
E l i m i n a t i o n o f c a t t l e u s e w o u ld p r o b a b l y h a v e l i t t l e
e ffe c t
on a ll o w i n g m o re w illo w s t o r e a c h a b o v e t h e snow i n w i n t e r o n t h e s tu d y
a re a ,
S. w o t f i i r a r e l y g ro w s o v e r 4 f e e t t a l l a n d c a t t l e u s e on o t h e r
w illo w s w as m o s t l y on lo w e r l a t e r a l s h o o t s .
F o ra g e c o m p e t i t i o n c a u s e d
b y t r a m p li n g a n d r u b b in g c o u ld becom e s i g n i f i c a n t w i t h h e a v i e r s t o c k i n g .
T he a p p r o x im a te s t o c k i n g r a t e w as o n e a n im a l p e r 5 a c r e s f o r a p e r i o d o f
82 d a y s i n a n a r e a w i t h t h e f o ll o w i n g p e r c e n t a g e s o f v e g e t a t i o n :
w illo w
ty p e 5 0 , se d g e ty p e 4 5 , g r a s s la n d ty p e 3, a sp e n ty p e I , and a q u a tic
ty p e I .
" D i s t u r b a n c e 'c o m p e t i t i o n , " a s h o t e d b y D e n n is to n ( 1 9 5 6 :1 1 4 ) i
-2 8 -
w h e re b y m oose move o u t when c a t t l e move i n , was n o t o b s e r v e d .
Moose
and c a t t l e w e re o b s e rv e d f e e d in g w i t h i n 10 f e e t o f e a c h o t h e r w i t h no
a p p a re n t c o n c ern e x c e p t c u r i o s it y .
O ther Com petition
F o ra g e c o m p e t it io n b e tw e e n m oose and b e a v e r ,w a s g r e a t e r th a n
c o m p e t it io n b e tw e e n m oose and c a t t l e .
Damage t o w illo w s by c u t t i n g and
f l o o d i n g was a p p a r e n t i n s e v e r a l a r e a s b u t w as n o t s i g n i f i c a n t .
It
c o u ld becom e s i g n i f i c a n t w i t h a s m a l l i n c r e a s e o f t h e b e a v e r p o p u l a t i o n .
M ost a s p e n c l o s e t o s u i t a b l e w a t e r a r e a s h a s b e e n e l i m i n a t e d so w illo w
was t h e p r im a r y woody fo o d and c o n s t r u c t i o n m a t e r i a l f o r b e a v e r .
P a c k a rd (1 9 4 7 :2 2 7 ) and R u d e r s d o r f ( 1 9 5 2 :3 7 , 51) r e p o r t e d t h a t b e a v e r
t u r n t o w illo w a f t e r a s p e n h a s b e e n e l i m i n a t e d and a p p a r e n t l y a c c e p t
i t as a .s a t i s f a c t o r y s u b s ti tu t e .
S e x a n d Age R a t i o s ,
P o p u l a t i o n T r e n d s , and M ovem ents■
Sex and a g e r a t i o s o f m oose i n summer d e te r m in e d fro m a e r i a l ob­
s e r v a t i o n s w e re m o re a c c u r a t e t h a n from g ro u n d o b s e r v a t i o n s . ■ C om pari­
s o n o f d a t a from g ro u n d o b s e r v a t i o n s from J u n e th ro u g h S e p te m b e r and
com bined a e r i a l o b s e r v a t i o n s from J u l y and S e p tem b e r ( T a b le V I I I )
i n d i c a t e s t h a t b u l l s w e re m ore r e a d i l y i d e n t i f i e d from t h e g ro u n d t h a n
cow s.
Know.lton ( 1 9 6 0 :1 6 8 -1 6 9 ) o b s e rv e d t h i s
a ls o .
The s e x o f 16 p e r ­
c e n t o f a l l m oose o b s e r v e d from t h e g ro u n d in .su m m e r w as n o t , d e te r m in e d .
M ost w e re p r o b a b l y cow s.
A dding t h e s e t o t h e cow g ro u p g i v e s a r a t i o
o f 73 b u l l s t o 100 cows w h ic h i s n e a r l y t h e sam e a s t h e r a t i o
from
-2 9 -
a e r i a l o b s e r v a t i o n s i n summer o f 7 4 : IOO0
The same c o m p a ris o n i n d i c a t e s
t h a t many c a l v e s w e re n o t o b s e rv e d from t h e g ro u n d . . P a r t o f t h e d if-r
f e r e n c e was d u e t o t r a n s i e n t cow s, w h ic h r a r e l y h ad c a l v e s , a p p e a r in g
i n t h e g ro u n d d a t a .
The m a jo r d i f f e r e n c e r e s u l t e d from d i f f i c u l t y i n
s e e in g th e s m a lle r c a lv e s 0
K no w lto n ( 1 9 6 0 :1 6 9 ) , E dw ards an d R itc e y
(1 9 5 8 :2 6 .1 ), P i m l o t t (1 9 5 9 a : 3 9 2 ) , a n d H o u s to n (1 9 6 8 :2 1 ) n o t e d t h a t o b - .
s e r v e d c a l f t o cow r a t i o s w e re n o t r e l i a b l e a t l e a s t i n sum m er.
TABLE V I I I 0
OBSERVED SEX AND AGE RATIOS OF MOOSE ON THE STUDY AREA.
NUMBER OF MOOSE OBSERVED IS IN PARENTHESES.
A p ril
AERIAL
OBSERVATIONS
O c t.
J u ly
S e p t.
J u ly -S e p t.
(50)1/
(23)
(31)
(54)
(16)
B u l l s : Cows
6 0 :1 0 0 8 5 :1 0 0 5 7 :1 0 0
C a lv e s:C o w s
6 0 :1 0 0 5 4 :1 0 0 7 1 :1 0 0
C a l v e s : A d u l t s 3 5 :1 0 0 3 5 :1 0 0 2 9 :1 0 0 4 5 :1 0 0
—
-
7 4 :1 0 0 ■
5 7 :1 0 0
3 2 :1 0 0
GROUND
OBSERVATIONS
Ju n e -S e p t.
M ar.
(674)
(46)2/
1 0 5 :1 0 0
3 4 :1 0 0
1 3 :1 0 0
15 :1 0 0
5 6 :1 0 0
4 8 :1 0 0 2 /
I / Two m o n th s b e f o r e s t a r t o f s tu d y .
Tj Snow m obile s u r v e y .
3 / N in e m a le and s i x fe m a le c a lv e s w e re o b s e r v e d .
The d a t a on s e x an d a g e r a t i o s
tio n s
from a e r i a l and sn o w m o b ile o b s e rv a ­
( T a b le V I I I ) and d a t a on t h e num ber o f m oose o b s e rv e d by p e r i o d s ,
th r o u g h o u t t h e s tu d y ( F ig u r e 4) i n d i c a t e t h a t o b s e rv e d s e x and ag e
r a t i o s w e re i n f l u e n c e d by home r a n g e and m i g r a t i o n p a t t e r n s a n d d i f f e r ­
e n t i a l u s e o f v e g e t a t i o n t y p e s b e tw e e n s e x e s .
P e e k .( 1 9 6 2 :3 6 1 - 3 6 2 ) n o te d
t h a t d i f f e r e n t i a l u s e o f v e g e t a t i o n t y p e s b e tw e e n s e x e s an d d i f f e r e n t i a l
Observations
3 30
-o 20-
Maximum
Daily
Aerial
O b s e r vat ions
Ground
O bservations
Apr
May
Aug
Sep
Ti me
F ig u r e 4 .
Oc t
Nov
Dec
F e b lM a
Period
Maximum d a i l y o b s e r v a t io n s o f i n d i v i d u a l m oose by se m im o n th ly p e r i o d s .
-3 1 -
m ovem ents from y e a r t o y e a r i n f l u e n c e d s e x a n d .a g e r a t i o s and P i m l o t t
(1 9 5 9 b :4 4 2 ) n o te d t h a t s e x r a t i o s w e re i n f l u e n c e d by d i f f e r e n t i a l u s e
b e tw e e n s e x e s o f o p e n h a b i t a t s .
S e p te m b e r r a t i o s
from a e r i a l o b s e r v a ­
t i o n s w e re p r o b a b l y m o st r e l i a b l e f o r t h e c o m p o s itio n o f t h e b r e e d in g
p o p u la tio n =
The m o st b u l l s and t h e m o st m o o se , e x c e p t f o r w i n t e r ,
w e re o b s e r v e d on t h e s tu d y a r e a i n S eptem ber=
w i t h t h e e a r l y ru t=
T h is p e r i o d c o in c i d e d
M ost b u l l s a p p a r e n t l y r a n g e d t h e tim b e r e d u p la n d s
d u r in g m o s t .o f t h e summer and moved down t o t h e lo w la n d s i n l a t e A u g u st
and Septem ber=
Cowan (1 9 5 0 :5 8 3 ) n o te d t h a t many b u l l s ra n g e d a t h i g h e r
e l e v a t i o n s t h a n t h e m a in h e r d s =
A ltm ann (1 9 5 9 :4 2 1 -4 2 2 ) n o te d u s e o f
w illo w f l a t s by l a r g e r b u l l s i n summer u n t i l A u g u st w hen t h e y moved t o ,
t h e tim b e r e d s l o p e s =
f o r t h e ru t=
I n S e p te m b e r th e y r e t u r n e d t o t h e w illo w f l a t s
P e e k (1 9 6 2 :3 6 3 ) o b s e rv e d a n i n c r e a s e d u s e o f w illo w b o t ­
tom s by b u l l s i n l a t e A u g u st and e a r l y S e p tem b e r= . R a t io s o b s e rv e d a t ,
o t h e r tim e s o f t h e y e a r g a v e d i f f e r e n t r e s u l t s =
I n w i n t e r w hen m oose
a r e on w i n t e r i n g a r e a s a n d r e a d i l y o b s e r v a b l e , t h e c o m p o s itio n o f t h e
p o p u l a t i o n on a g iv e n a r e a m a y b e d i f f e r e n t . from t h e b r e e d i n g p o p u la ­
tio n =
An i n t e n s i v e g ro u n d s u r v e y by sn o w m o b ile i n M arch i n d i c a t e d t h a t
o n ly a s m a l l p r o p o r t i o n o f a d u l t b u l l s w i n t e r e d on t h e s tu d y a r e a
( T a b le V I I I ) =
C o m p ariso n o f r a t i o s from d i f f e r e n t a r e a s i s p r o b a b ly
m e a n in g le s s b e c a u s e o f d i f f e r e n c e s i n t h e m ovem ent p a t t e r n s
o f each
p o p u la tio n = . U n t i l m o re i s known a b o u t m ovem ents o f m o o se , s e x and ag e
r a t i o s a r e b e s t u s e d a s i n d i c e s f o r d e te r m i n in g t r e n d s i n a g iv e n a r e a .
-3 2 -
The v a l i d i t y o f t h e S e p te m b e r r a t i o s i s q u e s t i o n a b l e .
s a m p le g i v e s a w id e c o n f i d e n c e i n t e r v a l .
The s m a ll
I f i t i s assum ed t h a t t h e
s a m p lin g was random and t h a t t h e p o p u l a t i o n was s t a b l e , t h e 95 p e r c e n t
c o n f id e n c e i n t e r v a l f o r t h e b u l l t o cow r a t i o w o u ld b e 6 6 :1 0 0 t o 94:
100 ( S p i e g e l 1 9 6 1 :1 5 8 -1 6 3 ) ,
The same a p p l i e s f o r t h e o t h e r r a t i o s . .
Moose n o t o b s e rv e d i n t h e c o n if e r o u s t im b e r ty p e s c o u ld c a u s e t h i s
la rg e v a r i a t i o n ,'
Cows w ith c a lv e s w e re r e p e a t e d l y o b s e rv e d i n a r e a s o f o n e - h a l f
t o two s q u a r e m i l e s th r o u g h o u t t h e summer and i n t o f a l l .
D u rin g t h i s
p e r i o d t h e c a l f t o cow r a t i o was s m a l l w hen t h e t o t a l num ber o f a n i ­
m a ls o b s e rv e d was l a r g e and v i c e v e r s a ( T a b le V I I I a n d F i g u r e 4) i n d i ­
c a t i n g a r e l a t i v e c o n s ta n c y o f c o w - c a lf g r o u p s .
On t h e J u l y , S e p te m b e r,
and O c to b e r f l i g h t s , s i x , s e v e n , and f i v e c a lv e s w e re o b s e r v e d , r e s p e o r
tiv e ly .
A cow w i t h tw in c a lv e s was o b s e r v e d i n th e sam e a r e a i n J u l y
and S e p te m b e r b u t was n o t o b s e rv e d i n O c to b e r ,
Some s o l i t a r y
cows
w e re r e p e a t e d l y o b s e rv e d i n t h e sam e a r e a s i n summer b u t o t h e r s w e re
o b s e rv e d i n a n a r e a f o r o n ly s h o r t p e r i o d s .
T h is may h a v e b e e n due
t o a g e d i f f e r e n c e s w i t h t h e y o u n g e r a n im a ls m oving t h e m o s t,
No b u l l s w e re o b s e rv e d f o r any e x te n d e d p e r i o d i n , any one a r e a
d u r in g sum m er.
They u s u a l l y w e re on t h e s tu d y a r e a f o r a , s h o r t p e r i o d
and th e n l e f t ,
a p p a r e n t l y i n t o t h e u p la n d t i m b e r , f o r a . l o n g e r p e r i o d , 1
A b u l l w i t h a r e d n e c k b a n d , p u t on b y Id a h o F is h and Game D e p a rtm e n t
p e r s o n n e l ( N i e l s o n and Shaw 1 9 6 7 ), w as o b s e rv e d on t h e s tu d y a r e a i r ­
r e g u l a r l y t h r o u g h o u t t h e summer ( F ig u r e 6 i n t h e a p p e n d ix ) ,
He was
-3 3 -
n e v e r o b s e r v e d on m ore t h a n t h r e e c o n s e c u t i v e d ay s and t h e i n t e r v a l s
w h e n -h e w as n o t o b s e rv e d ra n g e d from I t o 3 w e e k s „
The b u l l t o cow
r a t i o was l a r g e s t w hen t h e t o t a l num ber o f a n im a ls o b s e r v e d w as l a r g ­
e s t . a n d v i c e v e r s a ( T a b le V I I I a n d F i g u r e 4) i n d i c a t i n g t h a t b u l l s
'
I
ra n g e d o v e r g r e a t e r a r e a s t h a n cows d u r in g summer=
M ovem ents r e p o r t e d i n o t h e r s t u d i e s v a r y ( D e n n is to n 1 9 5 6 :1 1 0 ,
K now lton 1 9 6 0 :1 6 6 , P e e k 1 9 6 2 :3 6 2 , H o u s to n 1 9 6 8 :5 1 -5 3 )=
Home ra n g e s
a p p a r e n t l y v a r y w i t h t h e a r e a a s s u g g e s t e d by P e t e r s o n (1 9 5 5 :1 1 3 )
and w i t h c h a n g in g e n v ir o n m e n ta l c o n d i t i o n s , a s s u g g e s t e d b y P e e k
(1 9 6 2 :3 6 2 )=
The fe w e r o b s e r v a t i o n s o f b u l l s a p p a r e n t l y was n o t d u e t o u se
o f t im b e r t y p e s d u r in g th e day o n ly b u t r a t h e r t o c o n t i n u o u s , u s e o f
th e s e ty p e s =
C o w -c a lf g ro u p s a c c o u n t e d ‘f o r 62 p e r c e n t o f a l l o b s e r ­
v a t i o n s o f m o o s e . c r o s s i n g b e tw e e n t h e lo w la n d s and u p la n d t i m b e r .
G roups i n c l u d i n g b u l l s a c c o u n te d f o r 19 p e r c e n t o f .t h e t o t a l .
Some
cows w i t h c a lv e s u s e d t h e tim b e r f o r c o v e r ,d u r in g t h e day an d r e t u r n e d
t o t h e lo w la n d s t o f e e d i n t h e e v e n in g and e a r l y m orning=
T h is h a b i t
o c c a s i o n a l l y c a u s e s c a l f m o r t a l i t y on f e n c e s =
A d u lt m oose e a s i l y jump
m o st f e n c e s b u t c a lv e s u s u a l l y h a v e d i f f i c u l t y
and s o m e tim e s becom e •
e n ta n g le d =
M ig r a ti o n p a t t e r n s a p p e a r e d t o b e complex=
The s h a r p d e c l i n e i n
t h e num ber o f a n im a ls o b s e rv e d i n e a r l y O c to b e r ( F ig u r e 4) r e s u l t e d
from m i g r a t i o n o r a s h i f t b a c k t o t h e tim b e r e d a r e a s o f home r a n g e s .
The d e c l i n e c o r r e s p o n d e d w i t h two n a t u r a l phenom ena w h ic h c o u ld c a u s e
-34-
a te m p o ra ry s h i f t .
W illo w l e a v e s w e re k i l l e d a t t h i s tim e w h ic h may
h a v e m ade them u n p a l a t a b l e .
H o s le y ( 1 9 4 9 :1 6 ) c i t e d Cooney a s s t a t i n g
t h a t m oose r e t u r n e d t o tim b e r e d s l o p e s a t t h e end o f S e p te m b e r a f t e r
w illo w l e a v e s , w h ic h a r e p r e f e r r e d t o t h e b r o w s e , w e re k i l l e d .
i c e was a l s o fo rm e d a t t h i s
T h in
tim e w h ic h may h a v e made t r a v e l i n t h e low -
l a n d s d i f f i c u l t o r e v e n p a i n f u l f o r m o o se.
T h e s e phenom ena c o u ld a l s o
s t i m u l a t e m i g r a t i o n . ' The s m a l l c h a n g e o f t h e c a l f t o cow r a t i o b e tw e e n
J u l y an d S e p te m b e r and t h e l a r g e c h a n g e from S e p te m b e r t o O c to b e r
( T a b le V I I I ) i n d i c a t e d t h a t b o t h s o l i t a r y
O c to b e r .
cows and b u l l s m oved o u t i n
The p r e s e n c e on t h e s tu d y a r e a o f t h e m arked b u l l f o r s e v e r a l
summ ers b u t n o t i n w i n t e r ( 0 . H. V i y i o n , p e r s o n a l c o m m u n ic a tio n ) , t h e
s e a s o n w hen h e w as h a n d e d i n I d a h o , i n d i c a t e s t h a t some o f t h e m oose
p r e s e n t i n summer w i n t e r e ls e w h e r e , p r o b a b ly i n I d a h o .
The low p r o p o r ­
t i o n o f b u l l s on t h e s tu d y a r e a i n w i n t e r ( T a b le V I I I ) i n d i c a t e d t h a t
m o st o f t h e b u l l s w i n t e r e ls e w h e r e .
Cowan (1 9 5 0 :5 8 3 ) n o t e d t h a t many
b u l l s r a n g e d a t h i g h e r e l e v a t i o n s i n w i n t e r t h a n t h e m ain h e r d s and
S te v e n s (1 9 6 7 :1 1 ) o b s e rv e d o v e r 20 p e r c e n t o f t h e m oose i n o n e a r e a
w i n t e r i n g i n t h e s p r u c e - f i r com m unity w h ic h i s m o s tly a t h i g h e r e l e v a ­
tio n s .
I f a summer p o p u l a t i o n seg m en t e x i s t s , t h e d e f i n i t i o n o f m ig ra ­
t i o n s p r e s e n t e d by E dw ards and R itc e y ( 1 9 5 6 :4 8 7 ) , w h e re b y d i f f e r e n t
e n v ir o n m e n ts a r e u t i l i z e d t o m in im iz e d e t r i m e n t a l c o n d i t i o n s , may n e e d
re -e x a m in a tio n .
C o n d itio n s i n p r o b a b l e w i n t e r i n g e n v ir o n m e n ts o f f t h e
s tu d y a r e a a r e p o s s i b l y m ore d e t r i m e n t a l .
The c o n s ta n c y o f c o w - c a l f g ro u p s i n t o O c to b e r and t h e i r p re d o m i-
-3 5 -
n a tin g p re s e n c e i n e a r ly w in te r s u g g e s te d t h a t a r e s i d e n t p o p u la tio n s e g ­
m ent was p r e s e n t m ade up m o s tly o f c o w - c a l f g r o u p s .
P e t e r s o n (1 9 5 5 :1 1 0 )
i n d i c a t e d t h a t some m o o se , e s p e c i a l l y co w s, c a l v e s , and y e a r l i n g s ,
re m a in e d i n t h e lo w la n d s a l l y e a r .
A w i n t e r p o p u l a t i o n se g m e n t was p r e s e n t i n t h i s s tu d y a s i n d i c a t e d
by t h e many m o o s e ■o b s e r v e d b y l a t e w i n t e r a n d t h e s h a r p d e c l i n e i n s p r i n g
(F ig u re 4 ) .
The s h a r p i n c r e a s e i n t h e num ber o f m oose o b s e r v e d o n .t h e
s tu d y a r e a i n e a r l y F e b r u a r y ( F ig u r e 4) o c c u r r e d a f t e r a s e r i e s o f s to rm s
w h ic h i n c r e a s e d t h e snow c o v e r i n t h e lo w e r tim b e r e d a r e a s from a b o u t 3
to 6 f e e t o
M ovements t o w i n t e r r a n g e s i n r e s p o n s e t o i n c r e a s i n g sn o w .
d e p th s w e re o b s e rv e d by Edw ards and R i t c e y ( 1 9 5 6 :4 9 3 ) , P e e k ( 1 9 6 3 :2 2 9 ) ,
and H o u s to n ( 1 9 6 8 :4 7 ) .
Snow i n t h e lo w la n d s d i d n o t r e s t r i c t m ovem ents
e x c e p t f o r a few d a y s a f t e r h e a v y s n o w f a l l s „
S e t t l i n g and w in d a c t i o n
co m p a c te d t h e snow s u f f i c i e n t l y t o s u p p o r t , t h e m oose.
R e c o g n i ti o n o f i n d i v i d u a l m o o se, m o s t l y y e a r l i n g s , . o v e r s h o r t p e r i ­
ods i n d i c a t e d t h a t a t r a n s i e n t p o p u l a t i o n se g m e n t was a l s o p r e s e n t i n
sum m er.
H o u s to n ( 1 9 6 8 :1 7 , 21) o b s e rv e d r e s i d e n t , w i n t e r , and t r a n s i e n t
s e g m e n ts i n J a c k s o n H o l e . ■ He d id n o t c o n s i d e r a summer p o p u l a t i o n s e g ­
m e n t, i f p r e s e n t , t o b e s i g n i f i c a n t ( p e r s o n a l c o m m u n ic a tio n ).
S in c e t h e c o n d i t i o n o f t h e w i n t e r r a n g e i s im p o r ta n t i n t h e m anage­
m en t o f m o o se , i t i s d e s i r a b l e t o know w h e re t h e w i n t e r p o p u l a t i o n s e g ­
m en t i s d u r in g t h e h u n t i n g s e a s o n , a b o u t O c to b e r 20 t o D ecem ber 1 5 , and
w h a t p a r t o f th e p o p u l a t i o n se g m e n t p r e s e n t d u r in g t h e h u n t i n g s e a s o n
i s a ls o p re s e n t in w in te r.
The a p p e a r a n c e o f m oose on b o t h t h e n o r th
-3 6 -
and s o u th s i d e s o f t h e s tu d y a r e a i n w i n t e r s u g g e s te d t h a t p a r t o f t h e
w i n t e r p o p u l a t i o n se g m e n t sum m ered i n t h e G r a v e ll y M o u n ta in s and p a r t ,
i n t h e C e n t e n n ia l M o u n ta in s „
The a n im a ls p r e s e n t d u r in g t h e h u n tin g
s e a s o n w e re p r o b a b l y m o s tly r e s i d e n t s „
w o u ld h a v e l i t t l e
E x c e s s iv e h a r v e s t o f t h i s g ro u p
e f f e c t on t h e w i n t e r r a n g e u n l e s s a n im a ls from th e
o t h e r p o p u l a t i o n se g m e n ts m oved i n t o t a k e t h e i r p l a c e s a n d w e re s u b ­
j e c t e d to s u b se q u e n t h a r v e s t s .
The a p p a r e n t . r e d u c t i o n o f a l l p o p u la ­
t i o n se g m e n ts a f t e r i n t e n s i v e h u n t in g i n t h e 1 9 5 0 's (R e fu g e r e c o r d s )
s u g g e s t s t h a t t h i s r e p la c e m e n t d i d t a k e p l a d e .
H o u s to n ’ s (1 9 6 8 ;5 6 )
o b s e r v a t i o n s s u g g e st, t h a t v a c a n t a r e a s w o u ld q u i c k l y b e . f i l l e d by
y e a rlin g s .
The i n c r e a s i n g e v id e n c e t h a t o rp h a n e d c a lv e s r a r e l y s u r v i v e t h e
w i n t e r ( D e n n is to n 1 9 5 6 :1 1 2 , A ltm ann 1 9 5 8 :1 5 8 , H o u sto n 1 9 6 8 :7 7 ) and
t h a t tw in n in g r a t e s and t h e o c c u r r e n c e o f t r i p l e t s may b e g e n e t i c a l l y
c o n t r o l l e d ( P i m l o t t 1 9 5 9 a :3 8 8 , H o u s to n 1 9 6 8 :6 9 - 7 1 ) , a lt h o u g h n u t r i t i o n
a p p a r e n t l y i s a l s o i n v o lv e d ( P i m l o t t 1 9 5 9 a :3 9 7 - 3 9 9 ), s u g g e s t s t h a t a
r e g u l a t i o n o r r e q u e s t t o p r o t e c t cows f o llo w e d by a c a l f o r c a lv e s may
b e n e f i t p r o d u c t i v i t y an d y i e l d .
P i m l o t t (1 9 5 9 b :4 2 8 ) c i t e d M aki as i n d i ­
c a t i n g t h a t t h i s r e g u l a t i o n was i n e f f e c t i n F i n l a n d .
T he a p p a r e n t r e -
■
d u c t i o n i n t h e tw in n in g r a t e i n t h e s u r r o u n d in g a r e a i n r e c e n t y e a r s
( B a il e y 1 9 3 0 :4 1 , D e n n is to n 1 9 5 6 :1 0 8 , P e e k 1 9 6 2 :3 6 2 , K n o w lto n 1 9 6 0 :1 6 9 ,
S te v e n s 1 9 6 5 :3 1 , H o u s to n 1 9 6 8 :6 9 ) may b e p a r t l y due t o s e l e c t i o n a g a i n s t
p r o d u c t i v e cows i n " a n y m oose" h u n t in g s e a s o n s . ■ The o c c u r r e n c e o f cows
w i t h c a lv e s i n a c c e s s i b l e lo w la n d a r e a s w o u ld f a c i l i t a t e s u c h s e l e c t i o n .
-3 7 -
D a ta o f E dw ards and R i t c e y (1 9 5 8 :2 6 3 -2 6 4 ) s u g g e s t a p o s s i b l e s e l e c t i o n
a g a i n s t lo w la n d moose* . M oose sum m ering a t h i g h e r e l e v a t i o n s h a d a
h i g h e r tw in n in g r a t e an d d i d n o t a lw a y s a r r i v e on w i n t e r a r e a s so o n
enough t o b e h a r v e s t e d .
They th o u g h t some e n v ir o n m e n ta l f a c t o r a t
h i g h e r e l e v a t i o n s m ig h t b e r e s p o n s i b l e f o r t h e h i g h e r tw in n in g r a t e
th e r e *
D e t e r i o r a t i o n o f t h e r a n g e p r o b a b ly c a u se d t h e d e c l i n e i n t h e
tw in n in g r a t e i n a r e a s w h e re h u n t in g h a s b e e n p r o h i b i t e d .
B a i le y
( 1 9 3 0 :3 9 , 41) o b s e r v e d a h i g h tw in n in g r a t e i n Y e llo w s to n e N a t i o n a l
P a r k and a n a b u n d a n c e o f w illo w s a lth o u g h i n some a r e a s t h e y w e re
s e v e r e l y b ro w s e d . . P o o r , r e p r o d u c t i o n was o b s e rv e d b y de Vos (1 9 5 6 :5 2 0 )
on a r e f u g e w h e re m oose w e re p r o t e c t e d and h e th o u g h t i t m ig h t b e d u e.
t o p o o r r a n g e c o n d i t i o n s . • H o u sto n (1 9 6 8 :7 0 ) s u g g e s te d a p o s s i b l e
m echanism f o r c h a n g e s i n t h e tw in n in g r a t e i n r e s p o n s e t o .c h a n g e s i n
e n v ir o n m e n ta l s t r e s s e s .
T h e re was l i t t l e
e v id e n c e o f e n v ir o n m e n ta l
s t r e s s i n t h i s s tu d y .
The tw in n in g r a t e fo u n d i n t h i s s tu d y was h i g h e r t h a n i n s u r r o u n d ­
in g a re a s i n r e c e n t y e a r s .
The r a t e v a r i e d from 18 p e r c e n t i n A p r i l ,
1968 t o 17 p e r c e n t . i n S e p te m b e r, 1968 t o 13 p e r c e n t i n M a rc h , 1969;
t h e s e p e r c e n t a g e s i n v o l v e o n ly tw o , o n e , an d two s e t s o f t w i n s , r e s ­
p e c tiv e ly .
S te v e n s ( 1 9 6 5 :2 1 ) o b s e rv e d two s e t s o f tw in s on t h e s tu d y
a re a i n 1965.
H o u s to n (1 9 6 8 :6 9 ) o b s e rv e d a r a t e o f 4 .5 p e r c e n t i n
J a c k s o n H o le .
K n o w lto n (1 9 6 0 :1 6 9 ) o b s e r v e d no tw in s an d P e e k (196,2:
362) o b s e r v e d o n ly two s-fets i n one y e a r and no n e i n two o t h e r y e a r s .
N u t r i t i o n a p p a r e n t l y i s as i m p o r ta n t a s a g e n e t i c c o n t r o l s i n c e
-3 8 t h e h i g h e r tw in n in g r a t e s w e re r e p o r t e d from a r e a s w i t h a h i g h e r a v a i l ­
a b ility
o f w i l l o w s . ■ P i m l o t t ( 1 9 5 9 a :386) r e p o r t e d N o rth A m e ric an tw in n in g
r a t e s r a n g in g from 2 t o 28 p e r c e n t .
A lth o u g h t h e r e i s some c h a n c e f o r
e r r o r i n c o m p a rin g r a t e s from d i f f e r e n t a r e a s , t h e v a r i a b l e s b e tw e e n
a r e a s a s d i s c u s s e d e a r l i e r a r e c o n s i d e r a b l y l e s s b e c a u s e o n l y . c o w s ,w ith
c a lv e s a r e b e in g c o n s i d e r e d .
Range S u rv e y
F o u r . t r a n s e c t s f o r t h e C o le (1 9 6 3 ) r a n g e s u rv e y w e re e s t a b l i s h e d
on t h e s tu d y a r e a i n 1 9 6 5 .
c o n d itio n o f th e ra n g e .
They a r e o f l i t t l e
v a lu e i n d e te r m in in g t h e
A l l ta g g e d p l a n t s w e re n o t k e y s p e c i e s and o n ly
one t r a n s e c t was i n a r e p r e s e n t a t i v e l o c a t i o n i n a k e y a r e a .
l i f o l i a a p p e a r e d t o b e t h e k e y s p e c i e s on t h e s tu d y a r e a .
87) c o n s id e r e d t h i s t h e k ey s p e c i e s i n J a c k s o n H o le .
p la n ifo li-a was u n c e r t a i n .
S. m y r til-
H o u sto n (1 9 6 8 :
The s t a t u s o f S,
I t s g ro w th was g e n e r a l l y lo w e r, t h a n S, m y r til-
U f o l i a so w as l e s s a v a i l a b l e i n w i n t e r , i t p ro d u c e d l e s s f o r a g e b e c a u s e
of i t s
g ro w th fo rm , and i t was g e n e r a l l y i n p o o r c o n d i t i o n .
S. m y r t i l l i -
f o l i a was t h e p re d o m in a n t s p e c i e s ta g g e d i n o n ly two t r a n s e c t s .
A ll
t r a n s e c t s h a d m ore t h a n o n e s p e c i e s ta g g e d . . S te v e n s ( 1 9 6 5 :3 1 - 3 2 ) i n d i ­
c a te d t h a t o n ly one s p e c i e s o f w illo w s h o u ld b e sa m p le d i n a t r a n s e c t .
The a v a i l a b i l i t y
z o n e s on w illo w s w e re d i f f i c u l t t o e s t a b l i s h b e ­
c a u s e o f d i f f e r e n t i a l snow d e p t h s .
The v a r i a t i o n I n d a t a r e c o r d e d by
d i f f e r e n t i n d i v i d u a l s i s o f t e n due t o t h i s .
A f i x e d zo n e f o r e a ch
t r a n s e c t w o u ld l a r g e l y e l i m i n a t e t h i s v a r i a t i o n .
-3 9 -
C o le (1 9 6 3 :8 ) s u g g e s t e d a n a ll o w a b l e u s e o f l e a d e r s f o r w illo w o f
5 0 , 7 5 , and 95 p e r c e n t i n m i l d , a v e r a g e , and s e v e r e w i n t e r s , r e s p e c ­
tiv e ly .
H o u sto n (1 9 6 8 :8 5 ) s u g g e s t e d a ll o w a b l e u s e o f 5 0 , 7 0 , .and 90
p e r c e n t i n m i l d , a v e r a g e , a n d s e v e r e w i n t e r s , r e s p e c t i v e l y , f o r S.
Irtyrti-IlifotyCa.
T hese p e rc e n ta g e s a p p e a r s a t i s f a c t o r y .
Q u a l i t a t i v e o b s e r v a t i o n s on t h e c o n d i t i o n o f t h e r a n g e s h o u ld
a l s o b e a p a r t o f t h e s u r v e y a s s u g g e s t e d b y S te v e n s ( 1 9 6 5 :3 2 ) .
ra n g e a p p e are d to b e i n s a t i s f a c t o r y
The
c o n d i t i o n d u r in g t h e s tu d y .e x c e p t
f o r t h e a r e a o f w illo w s s o u th o f U pper Red Rock Lake w h ic h was o v e r ­
used.
The w i n t e r p o p u l a t i o n a p p e a r e d t o b e a p p ro a c h in g t h e c a r r y i n g
c a p a c ity o f th e ra n g e .
Management Recommendations
S to c k in g r a t e s o f c a t t l e s h o u ld t a k e i n t o a c c o u n t t h e a v a i l a b i l i t y
o f p a l a t a b l e f o r b s , g r a s s e s , and g r a s s - l i k e p l a n t s and o f S a lix W o lfii
on any g i v e n a r e a t o p r e v e n t s i g n i f i c a n t m o o s e - c a t t i e c o m p e t i t i o n . • The
p o s s ib ility
o f tr a m p li n g damage s h o u ld a l s o b e c o n s i d e r e d .
Fences
s h o u ld b e c o n s t r u c t e d a n d l o c a t e d t o m in im iz e h a z a r d s t o m o o se.
B eaver
s h o u ld b e h a r v e s t e d r e g u l a r l y t o m in im iz e c o m p e t it io n w ith .m o o s e .
U n t i l r e l i a b l e i n f o r m a t i o n i s o b t a i n e d on m o v em e n ts, a ,h u n t i n g
season s ta r tin g
a b o u t . S e p te m b e r 20 seem s p r e f e r a b l e t o t h e p r e s e n t
s e a s o n b e c a u s e o f t h e c o m p o s itio n and s i z e o f . t h e p o p u l a t i o n a t t h a t
tim e on t h e s tu d y a r e a .
Cows f o llo w e d by c a lv e s s h o u ld b e p r o t e c t e d
and t h e a n n u a l q u o ta , s h o u ld b e r a i s e d fro m f i v e t o e i g h t . ■
-4 0 -
New r a n g e s u r v e y t r a n s e c t s s h o u ld b e e s t a b l i s h e d t o r e p l a c e t h r e e
o f th e fo u r e x is tin g t r a n s e c t s .
C re e k i s s a t i s f a c t o r y .
The e x i s t i n g t r a n s e c t on Red Rock
An a d d i t i o n a l o n e s h o u ld b e e s t a b l i s h e d on t h i s
c r e e k t o t h e e a s t and two s h o u ld b e e s t a b l i s h e d s o u th o f U p p e r.R e d Rock
L ake b e tw e e n t h e m a in V-Ioea ervgetmann-l s t a n d and Shambow P o n d . ;
APPENDIX
-4 2 - .
TABLE IX .
PERCENT VOLUME AND FREQUENCY OF AQUATIC TYPE TAXA (U. S.
DEPARTMENT OF INTERIOR, 1 9 6 8 ) „ '
T axon
Potamogeton r tc h a r d s o n ti
Chara s p „ 2 /
M yrtophytlim sp ioatun
Potamogeton praelongus
Potamogeton p e o tin a tu s
S a g itta r ta ouneata
Etodea canadensis
Najas f l e x i l i s
Lemna tr i s u tc a
Total
V F o llo w s H o tc h k is s ( 1 9 6 7 :5 2 ) .
_2/ T d e n o te s l e s s t h a n 0 .5 p e r c e n t .
Volume
F re q u e n c y
36
33
15
5
5
70
46
28
12
10
4
I
14
12
10
2
T—/
100
-4 3 TABLE X .
PERCENT CANOPY-COVERAGE, FREQUENCY, AND INTERCEPT OF SEDGE TYPE
TAXA.
T . DENOTES LESS THAN 0 . 5 PERCENT.
T axon
S a lix -p la n ifo lia
S a lix w o l f i i
S a lix Candida
B e tu la glandulosa
S a lix geyeviana
S a lix bebbianaP o te n tilla fv u tic o s a
A rcto sta p h ylo s u va -u rsi
Carex s p p .
.
C la s s Musc i —
Juncus b a ltie u s
A ste r fo lia e e u s
Polemoniim o eeid en ta le
S w ertia perennis
P e d ie u lg ris groenlandiea
U n i d e n t i f i e d F o rb s
Gglium tr ifid u m
Paraxaeim s p p .
Po,a s p p .
Muhlenbergia racemosa
P o te n tilla g r a c ilis
T r ifo liim longipes .
A renarig l a t e r i f l o r a
Fhleum p ra ten se
V iola nephrophylla
Desehampsig ea esp ito sa
P arhdssia p a r v iflo r a t ,
Hordeum braehyantherum
A e h ille a m ille fo liu m
S t e l l a r i a lo n g ifo lia
Festued rubra] ' 1
Agropijron b a k e ri
C laytonia e h a m isso i■
Veronica amerieana
G lyceria e la ta
M uhlenbergia■f i l i f o r m i s ■
C anopyC o v e ra g e
F re q u e n c y
In te rc e p t
I
I
I
I
T
T
T
T
TABLE X .
(CO N T IN U E D )o
T axon
C anopyC o v e ra g e
Calamagrostis n eg le e ta
A llium sohoenoprasum
Solidago canadensis
Galium bo rea le
Alopecurus a lp in u s
Equisetum laevigatum^Phleum alpinum.
Antennaria anaphaioides
Eabenaria hyperborea
L itte r
W a te r
B a re
I / F o llo w s J a q u e s ( 1 9 4 9 :1 6 0 ) .
2 / F o llo w s D a v is (1 9 5 2 :5 3 )„
F re q u e n c y
T
T
T
T
T
2
2
2
2
2
T
T
T
T
47
3
I
2
2
.2
2
94
4
8
.
In te rc e p t
-4 5 TABLE X Io
PERCENT CANOPY-COVERAGE, . FREQUENCY, AND INTERCEPT OF WILLOW
TYPE TANA.
T .DENOTES LESS THAN 0 . 5 PERCENT.
C anopyC o v e ra g e
T axon
F re q u e n c y
In te rc e p t
S d lix m y v t i l l i f o U a
S d lix f lccnif o l i a
S d lix w o l f i i
S d lix geyeriana
B etu la glandulosa
F o te n tilla fv u tic o s a
S d lix Toebbiana
Eibes inevme
Lonioera in vo lu o ra ta
Rharmus a ln if o l i a
S d lix drurmondiana
S a lix m ontioola
S a l i x ' Candida
ArotofS taphy Ios uva- urs i
Linndea b o r e a lis
Corntis s to lo n ife r a
Pioeg engelmanni
Rosa w obdsii
Car'ex sp p „
,
C la s s Mpsc i —
Junous b a ltio u s
Swertig. perennis
P e d io u la ris groenlandioa
Sm ilaoina s t e l l a t a ■
Equisetum arvense=-'
Taraxacum s p p .
A s te r fo lia o e u s
Polemonium o co id en ta le
Galium bqreale
Geum macrophyllum
C d s tille ja m in ia ta
Fragaria virg in ia n a T rifo liu m longiges
Thaliotrum venulosum
15
14
9
5
5
2
I
I
I
I •
T
T
T
T
T
T
T
T
52
20
5
3
3
2
I
I
I
I
I
I
I
I
I
I
98
39
21
22
17
13
26
17
14
13
12
12
9
6
6
6
•
—4 6 TABLE X Io
(C O N T IN U E D )o
T axon
Poa p r a te n s is
.
C la s s H e p a t i c a e i '
Eubus aogu lis
Arenajpia l a t e r i f l o r a
Poa sp p o'
G ra m in e ae
S t e l l a r i a lo n g ifo lia
Pyrola a s a r if o l i a
Geranium ric h a rd so n ii
Viola nephrophylla
Galium tr ifid u m
Equisetum laevigatum —'
Veronica amerioana
P arnassia fim b ria ta
U n i d e n t i f i e d F o rb s
Hdbenaria hyperborea
Agrppyron repens
P arndssia p a r v iflo r a
Thaliotrum sp a rsiflo ru m
A ngelica arguta
Ejpilobium glandulosum
Mimulus' g u tta tu s
A c h ille a m ille fo liu m
P o te n tilla anserina
P o te n tilla g r a c ilis
Cirsium fo lio su m
AVlium schoenoprdsum
C o m p o sita e
Juncus saximontanus
Glayioriia cham issoi
GVyceria s t r i a t a
A ngeIica p in n a ta
Alopecurus a lp in u s
Lupinks a rg en teus 2 /
v a r o argenteus— 'Polygonum amphibium
Epilobium OnglU stifo liu m
C anopyC o v e ra g e
F re q u e n c y
I
I
I
T
T
T
T.
T
T
T
5
3
2
17
12
9
8
8
7
7
TT
T
T
TT
T
T
T
T
T
T
T
T
T
T
T
T
T
T
T
T
T
5
4
4
4
4
3
3
2
2
2
2
2
2
2
2
I
I
I
I
I
I
I
I
T
T
T
I
I
I
In te rc e p t
-4 7 TABLE XIo
(CONTINUED).
T axon
C anopyC o v erag e
Sot-Ldago m uttiLradiata
S te tla v L a s p ,
'T
T
L itte r
W a te r
B a re
46 •
7
I
F re q u e n c y
I / F o llo w s J a q u e s ( 1 9 4 9 :1 6 0 ) ,
2J F o llo w s D a v is ( 1 9 5 2 :5 1 - 5 3 ) ,
3 / F o llo w s H itc h c o c k and C r o n q u is t ( 1 9 6 1 :3 0 6 )„ .
I
I
84
16
6
In te rc e p t
-4 8 TABLE X I I .
PERCENT CANOPY-COVERAGE, FREQUENCY, AND INTERCEPT OF SPRUCE
TYPE TAXA.
T DENOTES LESS THAN 0 . 5 PERCENT.-
T axon
Canopy-r
C o v e ra g e
F re q u e n c y
51
49
13
4
Open O v e r s to r y
P i c e a engeXmanni
LXnnaea bovealXs
Comns canadensis
S d iix bebbiana
Lonicera in v o tu c ra ta
Cornus s to to n ife r a
S a iix -p ia n ifo tia
Ribes hudsonianim
S d lix w o i f i i
Rhamnus d l n i f o i i a
Ribes inerme
Junigerus communis
B etiiia gianduiosa
A rctb std p h yio s u va -u rsi
' S d tix W jj r t i i i if o i i a
C d re x S ^ p o 1 ,
C lk s s Miis c i —
.
Equisetum arvense—
Fragdria v irg in ia n a
Geranium ridhccrdsonii
U n i d e n t i f i e d F o rb s
Actdea rubra
Sm iideina s t e i i a t a
G iyeerid e ia ta
P d m d ssia fim b ria ta
Gdiium s p p ♦
Taraxaeum s p p .
Poiemonium o eeid en ta ie
Pyroia vire n s
Egiiobium angustifoU um
A ngeiiea arguta
A iiium h revistyiu m
Egiiobium gianduiosum
Geum maeroghyiium
Veronica amerieand
Thdiietrum venuiosum
In te rc e p t
2
2
2
I
I
I
I
T
T
T
T
T
J
10
9
7
7
6
6
3
3
3
2
2
2
2
86
66
56
50
43
59
30
43
20
53
40
20
13
13
13
13
I
I
I
I
I
13
10
10
7
7
;40
27
18
-4 9 TABLE X I I o
(C O N T IN U E D ).
T axon
C an o p y -.
C o v e ra g e .
F re q u e n c y
I
I
I
I
I
7
3
3
3
3
I
T
T
TT
T
3
7
7
7
3
3
T
T
T
T
T
3
3
3
3
3
Eabenaria hyperborea
T
T
T
'T
3
3
3
3
L itte r
B a re
64
I
92
3
A ntennaria anaphaioides
Pyrola a s a r ifo lia
C laytonia cham issoi
Seneoio tr ia n g u la r is
AngeIioa p in n a ta
Thaliotrum sp a rsiflo ru m
Calamagrostis canadensis •
P e d io u la ris groenlandioa
S t e l l a r i a lo n g if o l i a
Bromus o ilia tu s
Cinna l a t i f o l i a
C la s s H e p a t i c a e i ^
Barbarea orthooeras
Z iz ia a pt era
Jimous b a ltio u s .
Gentidria a f f i n i s Poa s p p .
L is te r a b o r e a lis
G ram in eae
I / F o llo w s J a q u e s ( 1 9 4 9 :1 6 0 ) .
2 / F o llo w s D a v is ( 1 9 5 2 :5 1 ) .
In te rc e p t
-5 0 TABLE X I I I .
PERCENT CANOPY-COVERAGE AND FREQUENCY OF GRASSLAND TYPE
TAXA.
T DENOTES LESS THAN 0 . 5 PERCENT.
T axon
Festuoa ovina
Taraxaoum s p p . •
Luptnus s p p .
U n i d e n t i f i e d F o rb s
M ioroseris nigresoens
Besseya wyomingensis
Galium b oreale
Cerastium arvense
Poa seounda
Agropyron s p p .
A m io a so ro ria
A o h ille a m ille fo liu m
Viola n u t t a l l i i
A hfennaria rosea
Fhtox- lo n g ifo lia
Anemorie patens
Agrppyron dasystaohyumA rth m isia tr id e n ta ta
P o te n tilla g r a o ilis
Ahenaria oongesta
G ra m in e ae
Arabis n u t t a l l i i
Zyg'ademls venenosus
S h xifra g a rhomboidea
T h ifo liu m longipes
Poa s p p .
Poa o u s io k ii ■
A stragalus m iserPhlox h o o d ii
Antennaria anaphaloides
Linum perenne
L eg u m in o sae
Geum tr iflo r u m
Chrysothamnus v is o i d i f to ru s
Frasera speoiosa
C anopyC o v e ra g e
F re q u e n c y
16
11
9
6
6
5
5
4
4
73
87
68
61
54
56
39
48
41
3
3
2
2
2
2
2
2
2
2
I
53
37
48
39
26
20
15
14
12
12
36
I
I
I
I
I
31
29
20
19
17
I
I
I
I
I
I
I
I
I
I
,
15 •
15
12
12
10
9
9
7
7
7
-5 1 TABLE X I I I o
(C O N T IN U E D )o
Taxon
U m b e ll if e r a e
Eriogonum umbellatum
K o eleria o r is t a t a
Agropyron b a keri
Geranium viseosissim um
A llium oernuum
Ranunoulus gldberrim us
Carex spp=
Solidago m u ltir a d ia ta
Muhleribergia s p .
Carex f i l i f o l i a
Androsaoe s e p te n tr io n a lis
Danthonia in term ed ia
C a s tille y a sulphurea
Senecio lugens
S o lid a g o nem oralis
Drdba nemorosa
Haplopappus i n t e g r i f o Iiu s
Peristemon radioosus
A s te r o o o id e n ta lis
I r i s m isso u rie n sis
Symphorioarpos oreo p h ilu s
Frayaria v irg in ia n a
Anemone m u ltifid a
Erigeron g ld b e llu s
Erigeron com positus
M ertensia o b lo n g if o l i a
Dodeoatheon sp„
Arabis s p .
Solidago m isso u rie n sis
G a illa rd ia a r is ta ta
A goseris glauea
A ste r fo lia o e u s
R osaceae
B a re
-
C anopyC o v e ra g e
F re q u e n c y
I
I
I
I
I
T
T
T
T
T
T
T
T
T
T
T
T
T .
T
T
T
T
T
T
T
7
7
5
2
2
17
12
9
9
9
7
7
5
5
3
3
3
3
3
3
2
2
2
2
2
T
T.
T
T
T
2
2
2
2
2
2
2
2
2
99
T,
T
T
T
12
-5 2 TABLE X IV .
PERCENT CANOPY-COVERAGE, FREQUENCY, AND INTERCEPT OF ASPEN
TYPE TAXAe
T DENOTES LESS TH AN'0 . 5 PERCENT.
T axon
C an o p y -.
C o v e ra g e
F re q u e n c y
Bcnpulus trem uloides
• 75
24
7
1
3
2
I
I .
I
I
T
T
Open i n O v e r s to r y
Bseudotsuga m e n s ie s ii■
Bioea engelmanni
SymphorioaJtPos o reophilus
B erb eris repens
Rosa w oodsii
S a lix bebbiana
Clematis Columbiana
Shepherdia oanadensis
Spiraea b e t u l i f o l i a
Lonioera in v o lu c ra ta
Taraxacum s p p .
U n i d e n t i f i e d F o rb s
Thaliotrum venulosum '
T rifo liu m longipes
Calamagrostis mibesoens
Geranium rio h a rd so n ii
Lupinus argenteus
v a r„ p a r v iflo r u s —
Aotaea rubra
Geranium visoosissim um
Arnica o o r d ifo lia
Frdgdria v irg in ia n a
Bhleum p ra ten se
Equisetum a rven seiJ
Sfniladina s t e l l a t a
A stragalus miserGalium b o r e a le '
A c h ille a m ille fo liu m
Elym us■glauous
Boa p a lu s tr is
Fragaria vesoa
C d stilleQ a m in ia ta
A ste r oonspiouus
In te rc e p t
13
11
11
9
8
8
76
50
44
26
38 20
7
7
7
6
5
5
40
40
34
24
32
32
5
3
3
2
2
2
2
2
2
2
12
32
26
30
24
16
16
12
6
6
-5 3 TABLE XIVo
(C O N T IN U E D ).
T axon
C anopyC o v e ra g e
A q u ileg ia fla v e so e n s
Ccccex s p p .
TPoa sp p i
A llium b v evistylvm
Arenavia l a t e r i f l o r a
2
I
I
I
I
G ra m in e ae
I
I
I
I
I
I
T
T
T
T
T
T
T
T
Agrogyron traehyoaulum
Ligustiow n filic in u m
A ntennaria rosea
C la s s M usci-L/
Solidago oanadensis
Bromus m arginatus
Pyrola seounda
AhdroSaoe s e g te n tr io n a lis
Collomia lin e a r is
S tip a eolumbiana
Bromus s p .
P o te n tilla spp=
Osmorhisa o o o id e n ta lis
C ru c ife ra e
A ste r o o o id e n ta lis
Geum s p .
Geum tr iflo r u m
Agropyron s p .
Poa p r a te n s is
S ile n e s p .
Arnioa oham issonis
Phleum alpinum
Veronioa amerioana
Barbavea orthooeras
Arabis s P=
A g ro stis sodbra
Seneoio se rra
Hordeum s p .
T
T
T
T
T.
T
T
T
T
T
T
T
T
.T
T
F re q u e n c y
4
22
20
16
16
12
10
8
6
6
4
14
6
6
6
6
4
4
4
4
2
2
2
2
2
2
2
2
2
2
2
2
2
2
—5 4 TABLE XIVo
(CO N T IN U E D )O
T axon
Agrepyron b a keri
V-iola adunoa
PediGutari-s paysoniana
Poa nervosa
Anemone m ultif-ida
Carex r o s s iv
Clem atis h ir s u tis s im a
K o eleria c r is t a t a
L itte r
B a re
Rock
C anopyC o v e ra g e
F re q u e n c y
T
T
T
T
T
2
2
2
2
2
T
T
T
75
5
2
2
2
2
96
20
4
I / F o llo w s H itc h c o c k and C r o n q u is t ( 1 9 6 1 :3 0 2 ) .
Tj F o llo w s D a v is (1 9 5 2 :5 1 )«
Zj F o llo w s J a q u e s (1 9 4 9 :1 6 0 )=
In te rc e p t
-5 5 TABLE XV= . PERCENT CANOPY-rCOVERAGE, FREQUENCY, AND INTERCEPT OF DOUGLASF IR TYPE TAXA.
T DENOTES LESS THAN 0 . 5 PERCENT.
T axon
C anopyC o v e ra g e
F re q u e n c y
" T
Pseudetsuga menz'les'i'L
Open i n O v e r s to r y
Pinus co n to vta
Abies tasioearipa
Pioea engetmanni
Spiraea b e t u l i f e l i a
Clem atis oolumbiana
Symphorioarpos o reo p h ilu s
S a lix so o u leria n a
Aoer gldbrum .
Juniperus oormunis
B erb eris repens
Ribes' Visoosissimum
Lofiioeira u ta h en sis
Rubiis p g r u ifio r u s
Calamagrostis rubesoens
Arniod o o r d ifo lid
A steridonspiouus
Aotaea rubra
ThdPiotrum venulosum
U n i d e n t i f i e d F o rb s
Antennaria raoemosa
Fraggria vesoa.
Aquilegi'a f la v e soens
C la s s M bsci—^
M ite lla sta u ro p e ta la
A stragalus m iser
Geraniwn v is o o s is simum
Poa p d lu s tr is
Pyrola seounda
Goodyera o b lo n g ifo lia
Viola adunoa
Galium boreale
Lupinus ccrgenteus
.
y d r . p a r v i f Iorus-P
In te rc e p t
6
3
2
I
I
T
T
T
T
T
25
22
7
6
6
4
4
4
3
2
2
2
2
I
80
85
18
45
35
35
28
25
8
25
23
13
13
18
I
I
I
I
15
13
10
10
I
10
"
l
19
12
2
1
.
“
TABLE XV.
56
—
(CO N T IN U E D )„ •
Taxon
Fragavia v iv g in ia n a
Gevm s p p .
Carex r o s s i i
Valeriana d io ie a
Calypso bulbosa
Antennaria anaphaloides
Disporim traohyoarpumArenaria l a t e r i f l o r a
A c h ille a m ille fo liu m
Taraxacum s p p .
A ntennaria rosea
G ram in eae
Collomia lin e a r is
F o te n tilla g r a c ilis
L itte r
B a re
Rock
C anopyC o v e ra g e
F re q u e n c y
I
I
I
I
T
8
5
5
3
5
T
T
T
T
T
T
T
T
T
92
T
T
5
5
5
5
3
3
3
3
3
95
3
3
I / F o llo w s J a q u e s ( 1 9 4 9 :1 6 0 ) .
2,/ F o llo w s H itc h c o c k and C r o n q u i s t (1 9 6 1 :3 0 2 ) .
In te rc e p t
-5 7 TABLE X V I .
PERCENT CANOPY-COVERAGE, FREQUENCY, AND IN TE RC E PT OF SPR U C E F I R TYPE TAXA„
T' DENOTES L E S S THAN 0 . 5 P ER C EN T.
Canopy-r
C o v e ra g e
Taxon
Frequency
Ab-Ies Ias-Loaarpa
P-Lcea engetmann-L
Open i n O v e r s t o r y
",
.
I/
C l a s s Musc i —
Fragdria v irg in ia n a
U n i d e n ti f ie d F orbs
Gdliim boreale
Viola adunaa
Allium b revistylu m
Pyrola seaunda
A c h ille a m ille fo liu m
Taraxacum spp„
A stragalus 'm iser
Elymus glaucus
Sm ilaaina s t e l l a t a
Lupihus argenteus
,
v a r . p a r v iflo r u s —
L itte r
60
28
23
19
6
\
Poputus tremitlo-Ldes
Berber-Ls repens
Clemat-Ls oolurrib-Lana
Spiraea b e t u l i f o l i a
Symphoriaarpos o reo p h ilu s
Thaliatrum venulosum
A ste r oonspicuus ,
Calamagrostis rubescens
L ig u stio im filic in u m
Actaea rubra
Arnica c o rd if o l i a
Geranium rio h a rd so n ii
In te rc e p t
3
T
T
6
4
4
80
70
-60
30
60
60
30
3
2•
2
I
I
I
T
T
20
20
10
40
30
20
10
10
T
T
T
T
10
10
10
10
T
88
10
90
12
11
10
.
I
_!/ F o llo w s Ja q u e s , ( 1 9 4 9 : 1 6 0 ) .
2 / F o llo w s H i t c h c o c k . and C r o n q u i s t ( 1 9 6 1 : 3 0 2 ) .
.
'
-5 8 -
TABLE XVII.
RESULTS .OF POINT-CENTERED QUARTER METHOD ANALYSES OF TREES.
V e g e t a t i o n Type
and Taxon
R e la tiv e
Frequency
R e la tiv e
D e n sity
R e la tiv e
Dominance
I m p o r ta n c e
V a lu e
S pruce
Vioea engeimanni
100
100
100
300
A spen
Voipulus tvem uloides
Pseudotsuga m e n z ie s ii,
Pioea engeimanni
T o tal
7 9 .4
1 5 .9
4 .8
9 0 .5
7 .0
2 .5
9 6 .5
3 .0
0 .6
2 6 6 .4 .
2 5.9
7 .9
100.1
1 0 0 .0
100.1
3 0 0 .2
7 8 .0
1 6 .0
6 .0
8 5 .0
1 1 .3
3 .8
9 4 .0
4 .9
1 .1
2 5 7 .0
3 2 .2
1 0.9
1 0 0 .0
100.1
1 0 0 .0
3 0 0 .1
6 6 .7
2 0 .0
1 3 .3
7 0 .0
2 0 .0
1 0 .0
7 7 .2
1 4 .9
7 .9
2 13.9
5 4 .9
3 1 .2
1 0 0 .0
1 0 0 .0
1 0 0 .0
3 0 0 .0
D ougl a s - f i r
Pseudotsuga m e n zie siiPinus co n to rta
Abies lasiooavga
T o tal
S p ru c e -fir
Abies lasiooarpa
Populus tvem uloides
Pioea engeimanni
T o ta l
-59-
TABLE X V III.
PLANT 'DISTRIBUTION BY VEGETATION' TYPE ON THE STUDY AREA
(AQ = A Q U A T i c i ', SE = SEDGE, WI = WILLOW, SP = SPRUCE,
AS = ASPEN, DF = DOUGLAS-FIR, SF = SPRUCE-FIR, GR =
GRASSLAND, MI = MISCELLANEOUS!'). X DENOTES PRESENCE.
Taxon
SE
WI
SP
AS
X
X
X
X
X
X
X
X
X
X
X
DF
SF
GR
EQUISETACEAE3/
Eqwisetum arvens& - 3/
laevigatvong O lustre^J
X
X
X
PINACEAE
Abies lasioeavpa
Picea engelmanni
Pinus -eontovta
fte x ilis
Pseudotsvga m e n z ie sii
X
X
X
X
X
X
X
X
X
X
X
CUPRESSACEAE
Junipevus communis
seopulovum
X
TYPHACEAE
Typhq l a t i f o l i a
SPARGANIACEAE
Spavganium m ultipeduneulatum
X
POT AMOGET ONACEAE
Potamogeton f i l i f o v m i s
p e e tin a tu s
pvaelongus
p u s illu s
v ieh a vd so n ii
so ste v ifo v m is
Z a n n ie h e llia p a lu s tv is
X
X
X
X
X
X
X
NAJADACEAE
Najas f l e x i l i s
X
JUNCAGINACEAE
Tvigl'oehin mavitima
X
ALISMACEAE
S a g itta v ia euneata ■
X
HYDROCHARITACEAE
Elodea canadensis
X
X
X
X
X
-6 0 TABLE ' X V I I I .
( C O N T IN U E D ).
Taxon
AQ
SE
WI
SP
AS
X
X
X
DF
SF
GR
MI
GRAMINEAE
T rib e F e stu c e a e
Bromus anomalus
o i Ziatus
inerm is
m arginatus
teotorum
Catabrosa aquatioa
DaetyZis gZomerata
Festuaa ovina
rubra
GZyceria b'oreaZis
eZata
grandis
s tr ia ta
MeZiea sp e e ta b iZ is
Poa amp Za
eompressa
'eu sia kii
ep iZ is
nervosa
p a Z u stris
p r a te n s i s ■
s'eeunda
PueaineZZia d ista n s
• X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
T r i b e H o rd e a e
Agropyron b a keri
dasystaehyum
desertorum
pseudorepens
repens
traehyeauZum
EZymus ein ereu s
gZaucus
maeounii
Hordeum braehyantherum
■jubqtum
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X >' X
X
X
X
■■
X x
X
X
X
X
T r i b e A veneae
Danthonia interm edia
Desehampsia e a esp ito sa
KoeZeria a r is ta ta
T ris e tu n spieatw n
X
X
X
X
X
-6 1 TABLE X V I I I „
(C O N T IN U E D ).
Taxon
AQ
SE
WI
SP
AS
DF
SF
CR
MI
T rib e A g ro stid e a e
A g ro stis alba
d ieg o en sis
exargta
sodbra
Alopeourus a eq u a lis
a lp in u s
Calamagrostis oanadensis
inexpansa
n eg leo ta
riibesoens
Cinna l a t i f o l i a
Muhlenbergia. f i l i f o r m i s ■
....
raoemosa.
Phleim alpinum
p ra ten se
S tip q eolumbiana
: tA illia m s ii ’
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
T r ib e .'C h io r id e a e
Bedhnannia syzigaohne
SpaPtina g ra o i H s
X
X
X
T r i b e .P h a l a r i d e a e
E ieroohloe odorata
P hg ld ris arundinaoea
X
X
X
CYPERACEAE
Carex
S e c tio n D iv isae
CaPex doug la s i i .'
sim u la ta
X
X
S e c tio n B r a c te o s a e .
X
Carelc v a llio o la
S e c tio n H eleo n astes
X
Car'ex disperma
X
S e c tio n S t e l l u l a t a e
X
Carex in te r i o r
X
X
X
S e c t i o n O v a le s
Carex athrostaohya
fe s tiv e lla
m ioroptera
. xera n tio a
X
X
X
X
S e c tio n F i l i f o l i a e
Carex f H i f o l i a
X
X
-62TABLE X V I I I . . (C O N T IN U E D ).
T axon
AQ
SE
WI
SP
AS • DF
SF
GR
S e c t i o n M ontanae
Carex r o s s i i
X
X
S e c tio n F irm icu lm es
X
Carex g eyeri
S e c tio n B ic o lo re s
Carex awrea
X
S e c tio n H irta e
X
Carex lanuginosa
S e c tio n A tra ta e
X
Carex r a y n o ld s ii
X
S e c tio n A cutae
Carex n ebraskensis ■.
X
S e c tio n V e s ic a ria e
Carex r o s tr a ta
E leoaharis a c io u la ris
m aorostaohya''
Eriophorum a n g u stifb liu n i
SoirpUs aoutus
X
X
X
X
X
X
X
LEMNACEAE'
Lemna minima—
minor
.t r i s u l o a ’
X
X
' ' X
JUNCACEAE
Junous b a ltio u s
bufonius
confu su s
e ffu su s
Io n g istylis^
sajximontanus
Luzula 'p a r v iflo r a
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X . X
X
LILIACEAE
A lliion b re v isty iu m
oermXun.
sohoenograsum
Cdloohortvis n itid u s
n u tta llii
Camas'sia ’quamash Disponon traohyearpurrt •
F r i t i l l a r i a atropurpurea
pudioa
X
- X
X
X
X
X
X
X
X
X
MI ■
-6 3 TABLE X V I I I .
(C O N T IN U E D ).
Taxon
AQ
SmilaQ-Lna raoemosa
s te lla ta
Zygadenus elegans
ven en esu s .
I RIDACEAE
SE
WI
SP
AS
DF
SF
X
X
X
X
X
X
X
X
CR
MI
X
X
,
I v i s mis s oicviensis
Sisyvinahium a n g u stifo liu m
X
X
X
X
X
X
ORCHIDACEAE
X
X
Calypso bulbosa
Goodyeva o b lo n g ifo lia
Eabenavia hypevbovea
u n a lasoensis
L is te v a b o v e a lis
X
X
X
-
X
X
X
SALI CACEAE
X
X
Populus acuminata
tvem uloides
tviohooavpa
S a lix bebbiana
Candida
dvummondiana
exigua
fv a g ilis
geyeviana
lasiandva
m onticola
m y v tiI I i f o I i a
p la n ifo I ia
v ig id a
sco u levia n a
w o lfii
BETULACEAE
X
X
X
X
X
X
X
X
X
X
X
X
X
X.
X
X
X ■
X
X
X
X
X
X '
X
X
X
X
X
X
X
X
X
X
X
X
X
? :
X
X
X
X . X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
!
B etu la glandulosa
URTI CACEAE
X
Uvtica d io ic a
X
SANTALACEAE
X
Comandva um bellata
POLYGONACEAE
Eviogonum umbellatum
Polygonum amphibium
a vicu la ve
X
'
X•
X
X
X
—6 4 TABLE X V I I I .
(C O N T IN U E D ).
T axon
AQ
WI
SP
AS
DF
SF
GR
MI
X!ix!
b is to r to id e s
d o u g la s ii■
la p a th ifo Uwn
Rheim rhapontieim
Rimex ■e risp u s
m aritim us■
oceidentaU s
paueifo Uus
sa U e ifo U u s
SE
X
X
XJ
CHENOPODIACEAE
Chenopodium album
rubrum
' Monolepis n u tta llia n a
X
PORTULACACEAE
C laytonia ~ehamissoiCARYOPHYLLACEAE
X
X
X
X lX l
Avenavia congest a
la te v iflo v a
Cevastium avvense
vulgatum
Sagina o c e id e n ta U s—'
S ile h e pavvyi
S t e lla v ia lo n g ifo lia
X
X
X
X
X
RANUNCULACEAE
X
X
X
X
X
X
Xl Xl X!
X
X
X
X
Xl XJ Xl
Actaea viibva
Anemone m u ltifid a
patens
A q u ileg ia fla v e sc e n s
Clematis Columbiana
h iv s u tis s im d
Delphinium glaucescens
n e ls o n iS f
Ranunculus acvifovrhis
a q u a ti H s
cymbalavia
glabevvimus
g m e lin ii
inamoeniis
natans
sc e le v a tu s
Thalictvum spavsiflovum
venulosum
X
X
X
X
X
X
X
X
X
X ■
X
X
X
X
X
X X
X
X
-65TAKLE X V I I I .o--
(C O N T IN U E D ) .
Taxon.
AQ
SE -
WI
SP
AS
DF-
SF
CR
BERBERIDACEAE ~
B erb eris repens
FUMARIACEAE.
C orydatis aurea
CRUCI FERAE
CRASSULACEAE
Sedim laneeolatum
SAXIFRAGACEAE
Eeueliera p a r v ifo liq
M ite lla .staicropetala
P a m d ssia fin ib ria ta
p a r v iflo r a
S a x ifra g a arguta <
rhomboidea
GROSS UI^ARI ACEAE
Ribes hudsonianum
inerme
setosum,
viseosissim um ■
ROSACEAE
Am elanehier a l n i f o l i a
Fragaria vesea
v irg in ia n a
Geum eanadense
maerophyllum
ir iflo r u m
X M
Ardbis Tiirsuta
n u tta ilii
Barbarea orthooeras
C apsella .b u rsa -p a sto ris
Cardamine brew eri
D eseurainia r ie h a r d s o n ii
Draba nemorosa
Erysimum ch eira n th o id es
Lepidium densiflo ru m
Rorippa isla n d ie a
ly r a ta
na£tiwtium raqudticum
Sisymbrium l o e s e l i i
TKe lypodium ■sa g iith tu m
T h ld sp i arvense
MI
-66TABLE X V I I I o
(CO NTIN UED )o
T axon
AQ
P o te n tilla .anserina
fr u tic o s a
grad Hs
norveg.ica
p la tte n s is .
Prunus v irg in ia n a
Rosa.w oodsii
Rvibus a c a u lis
-Ldaens
-parv i f Lotus
Spiraea b e t u l i f o l i a
SE
X
X
WI
SP
AS
X
X X
X
X ■
X
DF-
SF
CR
MI
X
X
X
■ X
X
X
X
X
X
X
X
X
X
X
LEGUMINOSAE
X X
X
X
X
X
X
X
X
X
X
x
X
X
L I NACEAE
Linum perenne'
CALLlTRICHACEAE
C a llitr ic h e herm aphroditica
verna
X
X
X
X
-XX
X
X
XXX
X
X
X
GERANIACEAE
Geranium r ic h a r d s o n ii
viscosissim um
X
A stra g a lu s canadensis
d a s y g lo ttis
eucosmiis
in fle x u s .
m iser
te rm in a lis
Caragana arborescOns
Hkdysarum boreale
' '
su lp h u resc‘e n s ,
i, ,
Lupinus 'a rg en teus v a r =. drgenteus— ^ /
p a r v iflo r u s —
s e ric e u s .
Medicago lu p u lin a ,
s a tiv a
M elilo tu s alba
o ffic in a lis
O xytropis d e fle x a
se ric e a
T rifo liu m longipes
p ra ten se
repens
—6 7 —
T A B L E •X V I I I .
(C O N T IN U E D ).
Taxon
AQ
SE
WI
SP
AS
DF
SE
GR' MI
ACERACEAE
A c e r gtdbmm
RHAMiNACEAE
-
Ehamnus a l'n ifo lia
MALVACEAE
Itiam na v iv u ta r is
V I OLACEAE
Viola adunoa
nephrophyIla
n u tta llii
ELAEAGNACEAE
Shepherdia canadensis ,
ONAGRACEAE
Epilqbium a n g u stifo liu m
glandulosum
p a lu s tr e
Oenothera fla v a
HALORAGIDACEAE
Muriophy Hum spicatumHIPPURIDACEAE
H ippuris v u lg a ris
UMBELLIFERAE
Xl
Cornus canadensis.
s to lo n ife r a
X
CORNACEAE.
X
A n g elica arguta
;
'•
p in n a ta .
Bupleurwm americdhum
Heracleum lanatum
LigUsticum filic in u m
Loipdtium cous
tr ite r n a tu m ■
• Musineon divarication
Osmorhiza o c c id e n ta lis
P e rid e r id ia g a ird n eri
Siim suave
Z iz ia apt era
—6 8 TABLE X V I I I ,.
(C O N T IN U E D ),.
Taxon
AQ
SE
WI
SP
AS
X
X X X
X
X
DF
SE
CR
MI
ERICACEAE
A ro to sta p h ylo s u va -u rsi
Pyrola a s a r if o l i a
secunda
u n iflo r a
vire n s
X
X
X
X
X
X
X
PRIMULACEAE
X
X
Xi X!
X
Xi
Androsace s e p te n tr io n a lis .
Dodeoatheon oonjugens .
pauoiflorum
Primula inoana
GENTIANACEAE
X
X
X
X
X
X
X
X
Xi Xi Xi
Frasera speoiosa
Gentiana a ff i n i s
p r o s tr a ta
S w ertia p erennis
POLEMONIACEAE
X
X
X
X
X
Xi Xi Xi Xi
Collomia lin e a r is
Phlox ho o d ii
ke ls e y i
I q n g ifo lia
Polemoniim ooo id en ta le
X
HYDROPHYLLACEAE
Hydrophyllum oapitatum
Phaoelia h a st a t a
se rio e a
X
X
BORAGI NACEAE
X
X
Xi
X
X
Xi Xi
H aokelia flo rib u n d a
Lappula eohinata
M erteAsia o i l i a t a
oblongi f o l i a
M yosotis s y Iv a tio a
P lagiobothrys so o u le ri
X
X
LABI ATAE
Agdstaohe u r t i o i f o l i a
Mentha a rven sis
P ru n ella v u lg a ris
S o u te lla r ia g a le rio u la ta
Staohys p a lu s tr is
X
X
X
X
X
X
X
X
X
SOLANACEAE
Hyoscyamus■n ig e r
X
-6 9 TABLE X V I I I o
(CO NTIN UED ).
Taxon
AQ
SE. WI
SP
.AS
DF
X
X
X
X
X
X
X
X
X
X
X
X
I
X
SF
GR
MI
SCROPHULARIACEAE
X
X
X
X
X
XX
X
X
X
X
X
X! X X
Besseya wyomingensis
C a s tille g a m in iata
sulghuvea
L im o sella aquatioa .
Mimulus g u tta tu s
Ovthooavpus- ■lu teu s
P ed io u la vis gvoenlandioa
paysoniana
Penstemon vadicosus
vyd b evg ii
Vevonioa amevioana
a vven sis
s e v p y llifo lia
X
X
X
X
LENTIBULARIACEAE
U tvicu la via v u lg a ris
X
PLANTAGINACEAE
Plantago magov
X
X
RUB IACEAE
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Galium boveale
tv ifid u n
tv iflo v u m
CAP RIFOLI ACEAE
X
X
X
X
X
X
X
X
X
.X
X
X
X X X X
X
X
X
X
Linnaea b o v e a lis
Lgnio'eva in vo lu cva ta
■
u ta h en sis
Sambuous pubens
Syrnphovioavpos oveophi Ius
Vibuvnum edule
VALERIANACEAE
Valeviana d io io a
e d u lis
X
X
CAMPANULACEAE
Campanula v o tu n d ifo lia .
X
COMPOSITAE
T rib e H e lia n th e a e -
Balsamovhiza s a g itta ta
Bidens oem na
Evioghylbam lanatioh '
G a illa vd ia o v i s t a t a
■E e lia n th e lla u n if Iova
H elianthus annuus
Iv a x a n th ifo lia
X
X
X
X
X
X
—7 0 TABLE X V I I I t,
(C O N T IN U E D ) o
AQ
T axon
SE
WI
SP
A S ' DF
SF
GR
XX
Madia gtom erata
Rudbeckia o o e id e n ta lis
W yethia h e lia n th o id e s
MI
X
X
X
X
X
M X X ixi >4
X
AahiZZea miZZefoZium
A rte m isia b ie n n is
dracuncuZus
f r ig id a
Zudovioiana
tr id e n ta ta
Chrysanthemum■Zeuoanthemum ■
M a trio a ria m a tr io a r io id e s
Tanacetum vuZgarej ■
XX
T r i b e A n th em ideaerv
T r ib e S e n e c io n e a e ^ /
A rniea oham issonis
o o r d if o Zia
s o r o r ia
Seneoio oanus
d e b iZ is
hydrophiZus
Zugens
pseudaureus
s erra
sphaerocephaZus
tria n g u Z a ris
w erneriae'fo Zius
Tetradym ia eanescens
X
X
X
X
X
X
X
X
X
. X
X
X
X
X
X
X
X
X
X
X
T rib e A stereae4 /
A s te r oonspiouus
engeZmanni
faZoatus
fo Ziaoeus
h e sp eriu s
o e e id e n ta Z is
pereZegans
Chrysothamnus nauseosus
v i s e i d i f ZorusE rigeron oom positus
ooryrribosus
gZdbeZZus
ZonehophyZZus.
p h i ZadeZphious
sp e e io su s
X
X
X
X
X
X
X
X
'
X
X
X
X
X
X
X
X
X
X
X
■
X
X
X
X
—7 1 —
TABLE X V I I I .
(C O N T IN U E D ).
Taxon
AQ
GT-Lndelia squarrosa
Haplopappus a o a u lis
in te g r ifo liu s
u n iflo T u s
Solidago oanadensis
m isso u rie n sis
m u ltiT a d ia ta
nem ovalis
Toionsendia vaTTyi
SE
WI
SP
AS
DF
SF
GR
MI
X
X
X
X
X
X
X
•X
X
X
X
X
X
X
X
T rib e In u le a e ^ /
A ntennaria anaphaloides
raeemosa
rosea
Gnapjiglium p g lu s tr e
X
X
X
X
X
X
X
X
X
X
T rib e Q ynareae-'
Cirsium a rien se
fo lio su m
vulgare
X
X
X
X
X
X
T rib e C ic h o rie a e -'
A goseris glauca
Crepis acuminata
Hierqcium a lb iflo ru m
cynoglossoides
' Lactuea p u lc h e lla
M ieroseris n ig reseen s
Sonehus u lig in o su s
Taraxacum laevigatum
o ffic in a le
Tragopogon dubius
p r a te n s is
X
X
X
X
X
X
I / I n c l u d e s w e t . b a n k s , s h o r e s , and d i t c h e s .
F o l lo w s
F o llo w s
F o llo w s
F o l lo w s
X
X
X
2j I n c l u d e s r o a d s i d e s and o t h e r d i s t u r b e d a r e a s „
_3/
4/
_5/
6/
X
X
D a v is ( 1 9 5 2 ) .
F a s s e t t (1 9 6 6 :3 7 6 ).
Saw yer ( 1 9 6 7 ) .
H i t c h c o c k ' and C r o n q u i s t ( 1 9 6 1 : 3 0 2 - 3 0 6 ) .
X
X
X
X
X
X
X
X
-7 2 -
40-
40
40
30
30-
30
20-
20
20
10
I
-
10 -
10______
S.
M4 0 -
40-
Iaaiandia
40
L
3020-
°
H'
:
b e b b ia n a
io
5. d r u m m o n d i a n a
30-
30
30
20-
20-
20
III
S.
20
40-
10-
m y rtillifolia
30
20
: i o -
S.
10-
I Q-
I
l
5
Illll .
i
10
15
Height (feet)
40
S.
gay ar ia n a
30
20
10
I,
20-
10-
5
10
15
20
5
Height (feet)
10
Height
201
o °
15
20
(feet)
S a c oula r i a n a
10-
Z
-I-----1
-----1
-----r—r
3 - 4 7-8
15-16
Height
F i g u r e 5„
T----r
T -T
25
35
30
40
(feet)
D i s t r i b u t i o n o f w i l l o w h e i g h t s on t h e s t u d y a r e a by s p e c i e s ,
S am ples o f l e s s t h a n 100 p l a n t s a r e p r o j e c t e d t o 100„
TABLE XIX.
FORAGE PLANTS WITH LESS THAN 2 PERCENT USE (EXCLUDING THE GENUS SALIX) BY MOOSE AND CATTLE IN ALL CATEGORIES EXCEPT FREQUENCY
FOR RUMEN SAMPLES. VALUES ARE PERCENTAGES. T DENOTES LESS THAN 0 .0 5 PERCENT AND NA DENOTES NOT AVAILABLE.
________________SUMMER_______________ ____________ FALL_______________________________WINTER_________________
,,
C a ttle
M oose
_,
M oose _ #
,
Moose
A re a o f U se^ :
A l 30
Ca 19 , A l 40
As 8
A l &■'
U I
IT i I - ' A l 35 Wi 24
Df 9 _ , As 5
T ax on
I n s t a n c e s o f U se: 1 7 ,8 1 3
1 2 , 2 7 3 6 , 8 1 2 2 , 2 2 1 ^ V o l. F re q . V b l. F r e q . V o l. 3 2 .4 8 0
2 6 ,7 4 8 3 , 4 8 7 - '
1 ,6 2 7 ^ /
NA
0 .0
0 .1
NA
0 .2
0 .0
0 .0
0
1 .4
NA
0 .0
NA
A m ela n o h ier a l n i f o l i a
NA
0 .4
0 .0
NA
13
0 .0
0 .0
0 .0
0 .1
0 .0
NA
NA
B e r b e r is re p e n s
T
0 .0
0
NA
NA
0 .0
0 .0
0 .0
0 .0
0 .4
NA
NA
C lem a tis o o lim b ia n a
T
0 .0
T
14
0 .1
0 .1
0 .0
NA
0 .1
0 .0
13
0 .2
NA
C om us s t o l o n i f e r a
T
0
0 .0
0 .0
0 .0
0
0 .0
0 .0
1 .1
0 .4
0 .1
1 .1
0 .3
L o n io e ra in v o lu o r a ta
0
T
0 .0
NA
0 .0
0 .0
NA
0 .0
0
NA
0 .0
NA
0 .0
P o te n tilla fr u tio o s a
T
T
0 .7
~
NA
0 .0
T
0 .4
0 .3
B ib e s inerm e
T
T
NA
NA
NA
0 .0
NA
NA
R ib e s se to su m
14
0 .0
0 .1
T
0 .7
13
0 .9
0 .0
0 .9
R ib e s s p p .
T
0
0 .0
0 .0
0 .0
0 .0
0
0 .0
0 .0
0 .4
NA
0 .0
NA
S h e p h e r d ia oan a d en sis
0 .0
T
NA
0 .0
0
NA
0 .0
0 .0
0 .0
0 .2
NA
NA
S p ir a e a b e t u l i f o l i a
T
0
0 .0
T
0 .0
0 .0
0 .0
0
0 .0
NA
0 .0
0 .0
A s t e r f o li a o e u s
0 .0
0 .0
0
0 .0
0
0 .0
0 .0
T
0 .1
NA
0 .0
0 .0
E p ilo b iu m a n g u s tifo liu m
0
0 .0
T
0 .0
0
0 .0
0 .0
0 .0
0 .2
0 .0
0 .0
0 .0
F ra g a ria v ir g in ia n a
0 .0
0 .0
0
0 .0
0
0 .0
0 .0
T
0 .5
0 .0
0 .0
0 .0
Geranium r i a h a r d s o n ii
0 .0
0
NA
0 .0
0 .0
NA
T
0 .1
0 .0
0 .0
0 .0
Geranium v i s c o s i s s i m m
T
T
T
14
0 .0
0 .0
0 .0
T
0 .0
13
0 .0
0 .0
Geun m aorophyllun
L u p in u s a r g e n te u s
v a r. p a rvi flo r u s lJ
R udbeokia o o o i d e n t a l is
S m ila c in a s t e l l a t a
Taraxacum o f f i c i n a l e
U n id e n tif ie d fo rb s
Bromus m a rg in a tu s
C a la m a g ro stis oan a d en sis
C arex s p p .
Junaus b a l t i c u s
Phleum p r a te n s e
U n i d e n t i f i e d G ram in eae
-
-
-
NA
NA
0 .0
0 .0
-
T
T
T
T
0 .0
0 .0
0 .0
0 .0
0 .0
T
T
0 .1
0 .1
T
T
0 .1
0 .0
0 .0
0 .0
T
NA
0 .0
NA
T
0 .0
0 .0
0 .0
0 .0
T
-
0 .0
-
0
0
0
0
25
NA
0 .0
0 .0
0 .0
0 .1
0
0
0
29
0
0 .0
-
-
0
NA
-
-
-
-
0 .0
0 .0
0 .0
0 .0
0 .0
0 .0
0 .0
0 .0
0 .0
T
NA
0 .0
0 .0
0 .0
-
0 .0
NA
0 .0
0 .0
-
0 .0
0 .0
0 .0
0 .0
0 .1
0 .0
0 .0
0 .1
0 .0
0 .0
T
NA
0 .0
0 .2
0 .0
0 .0
T
0 .0
NA
0 .0
NA
NA
“
0 .0
NA
0 .0
NA
0 .0
-
V A l = a l l a r e a s , Ca = c a t t l e a r e a s , As = a s p e n t y p e , Wi = w illo w t y p e , T i = u p la n d tim b e r t y p e s , and Df = D o u g l a s - f i r t y p e .
Numbers a r e t o t a l f e e d i n g s i t e s f o r summer an d w i n t e r and t o t a l ru n e n sa m p le s f o r f a l l .
2 / I n s u f f i c i e n t sa m p le .
F o llo w s H itc h c o c k and C r o n q u is t ( 1 9 6 1 :3 0 2 ) .
LEGEND
M o o se O b s e rv a tio n s
S um m er
S
W in te r
5
M a r k e d A d u lt B u lls
A 6 /1 4
X 9 /ig
B 7/ 9
Y 9 /2 3
C 7 /3 1
Z 9 /2 6
D 8/ 9
E 8 /1 8
F 9 /1 1
G 9 /1 2
H 9 /1 3
76'
Road
S tream
I
■^1
■P>
I
F ig u re 6 .
Ground o b s e r v a t i o n s o f m oose i n summer and w i n t e r by q u a r t e r - s e c t i o n and
o b s e r v a t i o n d a t e s and l o c a t i o n s o f two m arked b u l l s .
See F i g u r e I f o r a
d e t a i l e d d i s t r i b u t i o n o f v e g e t a t i o n t y p e s and o t h e r f e a t u r e s .
LITERATURE CITED
A l d e n , W„ C.
1953» P h y s i o g r a p h y and g l a c i a l g e o lo g y o f w e s t e r n M ontana
and a d j a c e n t a r e a s ,
U„ .S„ D e p t 0 I n t e r i o r , G e o l . S u r v e y P r o f .
P a p e r 2 31.
200 p p .
A ltm ann, M a rg a r e t.
1 958.
S o c ia l i n te g r a t io n of th e m o o se .c a lf.
B eh av io r 6 (3 -4 ):1 5 5 -1 5 9 .
A nim al
A ltm a n n , M a r g a r e t .
1 9 5 9 . Group d y n a m ic s i n Wyoming m oose d u r i n g t h e
ru ttin g season,
J . Mammal.
4 0 (3 ):4 2 0 -4 2 4 .
B a i l e y , V.
1930. A nim al l i f e o f Y e l l o w s t o n e ' N a t i o n a l P a r k .
Thom as, S p r i n g f i e l d , , 1 1 1 .
241 p p .
C h a r l e s C.
B a n k o , W. E.
1 9 5 1 . . Moose b r o w s e i n v e s t i g a t i o n s .
Red Rock Lake's N a t l .
W i l d l . R e f u g e , M o n id a , M ont. T y p e w r i t t e n .
17 p p .
B a n k o , W» E.
1 9 6 0 . The t r u m p e t e r sw an. U. S. D e p t. I n t e r i o r , F i s h and
W i l d l . S e r v . ,N. Am. F a u n a No. 63 .
214 pp ,
B o o t h , W» E.
1950. F l o r a o f M ontana. P a r t . I .
a t M ont. S t a t e C o l l . , Bozeman,.
232 p p .
The R e s e a r c h F o u n d a t i o n
B o o t h , W. E . , and J . C9 W r i g h t .
196 6 . F l o r a o f M o n ta n a .
M ont. S t a t e U n i v . , Bozeman,
305 p p .
C o l e , G. F ,
1963. ' Range s u r v e y g u i d e .
M oose, Wyo . 22 p p .
P art II.
G rand T e t o n N a t . H i s t . A s s o c . ,
Cottarn, G . , and J . T. C u r t i s . ,1 9 5 6 . The u s e o f d i s t a n c e m e a s u r e s i n
p h y t o s o c i o l o g i c a l s a m p l i n g . ■ E c o lo g y 3 7 ( 3 ) : 4 5 1 - 4 6 0 .
Cowan, I . McT. 1 950.
Some v i t a l s t a t i s t i c s o f b i g game on o v e r s t o c k e d
m o u n ta in r a n g e .
T r a n s . N. Am. W i l d l . C o n f . 1 5 : 5 8 1 - 5 8 8 .
D a u b e n m ire , R. F . • 1943. V e g e t a t i o n a l z o n a t i o n i n t h e Rocky M o u n ta i n s .
The B o t a n . Rev.
9 ( 6 ) : 3 2 5 r3 9 3 .
D a u b e n m ire , R.
1959. A c a n o p y - c o v e r a g e m eth o d o f v e g e t a t i o n a l a n a l y s i s
N o rth w est S c i.
3 3 (l):4 3 -6 4 .
D a v i s , R. J .
836 p p .
1952.
F lo ra o f Idaho.
Wm. C. Brown C o ., D ubuque, Iow a.
D e n n i s t o n , R. H . , I I .
1956. E c o l o g y , b e h a v i o r and p o p u l a t i o n dynam ics
o f , t h e Wyoming o r Rocky M o u n ta in m o o se , Aloes aloes s h lv a s l^
Z o o lo g ic a 4 1 :1 0 5 -1 1 8 .
—7 6 —
De V o s , A. 1956„
W ildl= Conf=
Summer s t u d i e s o f m oose i n O n t a r i o *
2 1 :5 1 0 -5 2 5 =
E d w a rd s , R= Y . , and R= W= R itc e y =
J= Mammal= • 3 7 ( 4 ) : 4 8 6 - 4 9 4 = •
1956.
T rans= N= Am.
The m i g r a t i o n s o f a m oose h e r d
E d w a rd s , R= Y=, a n d . R= W= R itc e y =
1958= R e p r o d u c t i o n i n a m oose,
p o p u latio n =
J= W ild l= Mgmt=- 2 2 ( 3 ) : 2 6 1 - 2 6 8 =
F a m e s , P= E= 1969= W a te r s u p p l y o u t l o o k f o r M on ta n a . ■ U= S . D e p t.
A g r= , S o i l , Conserv= Serv= J a n = , F e b = , M ar.
F a s s e t t , N= C= 1966= A m a n u a l o f a q u a t i c p l a n t s .
P r e s s , M a d is o n = . 405 p p .
The Univ= o f W i s e . ■
F l e r o v , K= K= 1960= F a u n a o f USSR, Mammals = V o l. I , No, 2.
(T ra n sl.
from R u s s i a n ) I s r a e l Program, f o r S c i e n t i f i c T r a n s l a t i o n s , O f f i c e
o f Tech= S e r v = , U= S. Dept= Commerce, W a s h in g to n ,
257 p p .
G a t e s , F= C.
1949= . F i e l d m a n u a l o f p l a n t e cology=
C o ., New Y o rk ,
137 pp=
M c G ra w -H ill Book.
H a n s o n , H= C= 1953= V e g e t a t i o n t y p e s i n n o r t h w e s t e r n A l a s k a and
c o m p a r is o n s w i t h c o m m u n itie s i n o t h e r a r c t i c r e g i o n s =
E c o lo g y
34(1 ):1 1 1 -1 4 0 =
H a n s o n , W= R=, and F= G r a y b i l l =
a n a ly se s=
J= W ild l= Mgmt=
1956= Sam ple s i z e i n f o o d - h a b i t s
2 0 (l):6 4 -6 8 .
H a r r y , G= B.
1957= W i n t e r fo o d h a b i t s o f m oose i n J a c k s o n H o le ;
Wyoming = J i W i l d l . Mgmt= 2 1 ( 1 ) : 5 3 - 5 7 =
H i t c h c o c k , C= L=, and A= C r o n q u is t=
1961= . V a s c u l a r p l a n t s o f . t h e
P a c i f i c N o rth w e st=
P a r t 3. U n i v . o f Wash. P r e s s i S e a t t l e .
614 pp
H o s l e y , N= W= 1949= The m oose and i t s e c o lo g y = U= S. D e p t . I n t e r i o r ,
F i s h and W ild l= Serv= W ild l= L e a f l e t 312.
51 pp=
H o t c h k i s s , N= 1967= U n d e r w a t e r and f l o a t i n g - l e a v e d p l a n t s o f t h e
U n i t e d S t a t e s and Canada= U= S= Dept= I n t e r i o r , F i s h a n d . W i l d l .
Serv= R e s o u r c e Publ= 44= 124 pp=
H o u s t o n , D= B= 1968= The S h i r a s m o o s e . i n J a c k s o n H o l e , Wyoming.
T e t o n N a t . H is t= Assoc= Tech= B u l l . No= I ,
HG p p .
Grand
-7 7 J a q u e s , Hi E 0
1949„
P la n t fa m ilie s -h o w
B r o w n C o ., , D u b u q u e , I o w a . .
1 7 7 p p '.
to
k n o w . th e m .
2nd
e d . . Wm.
C.
K e n d e i g h , S „ C0 .1961. A n im al e c o l o g y .
C l i f f s , N. J . . 468 p p .
P r e n t i c e - H a l l , I n c . , Englewood
K n o w lto n , F . F . I 9 6 0 . Food , h a b i t s , m ovem ents and p o p u l a t i o n s o f moose
i n t h e G r a v e l l y M o u n t a i n s , M o n ta n a .
J . W i l d l . Mgmt.■ 2 4 ( 2 ) : 1 6 2 - 1 7 0 .
K o r s c h g e n , L. J .
1 963. R evisio n o / M a r t i n , A. C . , F o o d - h a b i t s p r o ­
c e d u r e s , C h a p t . 1 3 . In M osby, H0 S-. ( e d i t o r ) , W i l d l i f e i n v e s t i g a ­
t i o n a l t e c h n i q u e s . 2nd e d . The W i l d l i f e S o c i e t y , W a s h in g to n .
419 p p .
L e o p o l d , A.
1 933.
481 p p .
Game m anagem ent.
C h a r l e s S c r i b n e r ' s S o n s , New Y ork.
M a r t i n , A. C . , R. H. G e n s c h , and C. P . Brown.
1946. A l t e r n a t i v e m eth o d s
i n u p l a n d . g a m e b ird fo o d a n a l y s i s .
J : W i l d l . Mgmt. - 1 0 ( 1 ) : 8 - 1 2 .
M c M illa n , J . F .
1953.
Some f e e d i n g h a b i t s o f moose i n Y e l l o w s t o n e P a r k .
E c o lo g y 3 4 ( 1 ) : 1 0 2 - 1 1 0 .
M ech, L. D.
1 966. The w o l v e s o f I s l e Royale.o f t h e U. S. F a u n a S e t . 7.
210 p p .
Fauna o f t h e N a t l . P a rk s
N i e l s o n , A. E . , and W0 M. Shaw,. , 1967. A h e l i c o p t e r - d a r t g u n t e c h n i q u e
f o r c a p t u r i n g m oose. P r o c . A n n u a l C o n f. W e s t e r n A s s o c . S t a t e
Game and F i s h C o m m is s io n e rs 4 7 : 1 8 3 - 1 9 9 .
N i e l s o n , E. C . , and D. N. F a r n s w o r t h .
1965.
S o il su rv e y handbook f o r
Red Rock L a k e s M i g r a t o r y W a te r f o w l R e fu g e i n s u r v e y a r e a 0 6 1 . U.
S. D e p t. A g r.,, S o i l . C o n s e r v . S e r v . T y p e w r i t t e n .
178 p p .
C o s t i n g , H0 J .
1 956. The s t u d y o f p l a n t c o m m u n itie s . ■ 2nd e d .
F reem an and C o . , San F r a n c i s c o .
440 p p .
W. H.
P a c k a r d , F . M. 1 9 4 7 . A s u r v e y o f t h e b e a v e r p o p u l a t i o n o f Rocky
M o u n ta in N a t i o n a l P a r k , C o l o r a d o . J . Mammal.
2 8 (3 ):2 1 9 -2 2 7 .
P a t t e n , D. T.
1 963. V e g e t a t i o n a l p a t t e r n i n r e l a t i o n t o ,e n v ir o n m e n t s
i n t h e M a d iso n R a n g e , M o n ta n a . • E c o l . M onographs 3 3 : 3 7 5 - 4 0 6 .
P a t t e n , D. T .
1 968. Dynamics o f t h e s h r u b c o n tin u u m a l o n g t h e G a l l a t i n
R iv e r i n Y e llo w sto n e N a tio n a l P a rk .
E c o l o g y ,4 9 ( 6 ) : 1 1 0 7 - 1 1 1 2 .
-
78
-
Peek, J „ Mo
196 2 ,
S t u d i e s o f m o o s e . i n t h e G r a v e l l y and S n o w e r e s t
M o u n t a i n s , Montana= J . W ild l= Mgmt= 2 6 ( 4 ) :3 6 0 - 3 6 5 ,
P e e k , J= Mo 1963= A p p r a i s a l o f a .m o o s e r a n g e i n s o u t h w e s t e r n M ontana. ■
J . Range Mgmt= 1 6 ( 5 ) :227-231=
P e t e r s o n , R= L= 1955=
T o ro n to =
280 pp =
N o r th A m e ric a n moose=
Univ= o f T o r o n t o P r e s s ,
P i m l o t t , D= Ho . 1959a= R e p r o d u c t i o n a n d p r o d u c t i v i t y o f N e w foundland
m oose. J= W ildl= ,Mgmt= 2 3 ( 4 ) : 3 8 1 - 4 0 1 ;
P i m l o t t i D. H= 1 9 5 9 b = Moose h a r v e s t s i n N ew foundland and F e n n o s c a n d i a n
c o u n t r i e s = T rans= N= Am. W i l d l 0 C o n f . . 2 4 : 4 2 2 - 4 4 8 .
P o r t e r , C. L= 1967. A f l o r a o f Wyoming. P a r t V=
E x p t . Sta= R e s e a rc h , j o u r n a l 14.
37 p p .
U n i v . o f Wyo. Agr=
R a u p , Ho M= .1959= ■ The w i l l o w s . o f b o r e a l w e s t e r n A m e ric a .
G ray Herb = H a r v a r d Univ= No. •CLXXXV. 95 p p .
C o n trib =
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