Nutritional and developmental responses of three mountain bunchgrasses to summer cattle and winter
elk grazing
by William John Dragt
A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in
Range Science
Montana State University
© Copyright by William John Dragt (1985)
Abstract:
Elk-Livestock Studies in Montana was established to study elk-livestock interactions through
cooperative research programs of various state and federal agencies. The objectives of Elk-Livestock
Studies were: 1) to determine the influence of various cattle management practices on elk and their
habitats, 2) to evaluate alternative management practices, and 3) to develop beneficial guidelines for
dual use of the available resource. This research project fell within objective one, and its objectives
were: 1) to quantify the effects of summer cattle grazing at various phonological stages on the winter
nutritional values of three mountain bunchgrasses, and 2) to quantify the effects of winter elk grazing
on the growth and development in the following spring of the same three species. Rough fescue
(Festuca scabrella). Idaho fescue (Festuca idahoensis) and bluebunch wheatgrass (Agropyron spicatum)
were the species studied. The study was conducted on the South Crow elk winter range in the Elkhorn
Mountains of Montana.
For objective one a random experimental design of one factor with three levels and nine treatments was
used. The grass species were the factor levels and the treatments consisted of the phenological stage
when grazing had occurred plus an ungrazed stage. Winter chemical constituent values were the
dependent variables. For objective two a randomized block design having one factor with three levels
and two treatments in five blocks was used. Sites, species, grazing, and standing dead were the blocks,
factor and two treatments, respectively. The dependent variables were plant height and phenological
stage on six spring sampling dates.
None of the summer grazing treatments affected rough fescue or bluebunch wheatgrass winter
nutritional values, and grazed plants were similar to ungrazed plants. Some phenological stages of
summer grazing were reflected in Idaho fescue winter chemical constituent values, but no management
significance could be attached to the differences. Winter grazing did not affect spring leaf lengths in
rough fescue or bluebunch wheatgrass, and only occasionally affected Idaho fescue spring growth. The
presence of standing dead in the spring resulted in longer Idaho fescue leaves in the next spring.
Neither rough fescue nor bluebunch wheatgrass spring leaf lengths were affected by the presence of
overwintering residual material. These results were discussed from the perspective of the winter
physiological activity of each species of grass.. NUTRITIONAL AND DEVELOPMENTAL RESPONSES OF THREE MOUNTAIN BUNCHGRASSES
TO SUMMER CATTLE AND WINTER ELK GRAZING
W illiam John D ragt
A t h e s i s su b m itte d i n p a r t i a l f u l f i l l m e n t
of th e re q u ire m e n ts f o r the degree
of
M aster of S c ie n ce
in
Range S cience
MONTANA STATE UNIVERSITY
Bozeman, Montana
June 1985
/V 3 7 2
a 7 Cf
C .^
ii
APPROVAL
o f a t h e s i s su b m itted by
W illiam John Dragt
This t h e s i s has been re a d by each member of th e t h e s i s committee
and has been found to be s a t i s f a c t o r y re g a rd in g c o n te n t, E nglish
usage, form at, c i t a t i o n s , b ib lio g r a p h ic s t y l e , and c o n s is s te n c y and i s
ready f o r subm ission to the College o f G raduate S tu d ie s .
/& ,
C h airp erso n , Graduate Committee
Approved f o r th e Major Department
Head, Major Department
Approved f o r the College o f Graduate S tu d ie s
2. Q
Date
M f/'
Graduate Dean
iii
STATEMENT OF PERMISSION TO USE
In
p re se n tin g
th is
th e sis
in
p a rtia l
f u l f i l l m e n t o f th e
r e q u i r e m e n t s f o r a m a s t e r ’s d e g r e e a t M ontana S t a t e U n i v e r s i t y ,
I
a g r e e t h a t t h e L i b r a r y s h a l l make i t a v a i l a b l e t o b o r r o w e r s u n d e r
r u l e s of th e L ib ra ry .
B r i e f q u o t a t i o n s from ..this t h e s i s a r e a llo w a b le
w ith o u t s p e c i a l p e rm is sio n , provided t h a t a c c u r a te acknowledgement of
so u rce i s made.
P e rm iss io n f o r e x te n s iv e q u o t a t i o n from or r e p r o d u c tio n of t h i s
t h e s i s may be g ra n te d by my major p r o f e s s o r , o r i n h i s absence, by the
D ir e c to r of L i b r a r i e s when, i n th e o p in io n o f e i t h e r , th e proposed use
of th e m a t e r i a l i s f o r s c h o l a r l y purposes.
m a te ria l
Any copying or use of the
i n t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a llo w e d
w ith o u t my w r i t t e n p e rm issio n .
S ig n a tu r e / / > 7
iv
TABLE OF CONTENTS
Page
LIST OF TABLES......... ....................................................................................................
LIST OF FIGURES........... ....................... ............................................. ...................
vi
v iii
ABSTRACT........................................................'.............. ................... .............................
x
INTRODUCTION................................... .............................................................................
1
LITERATURE REVIEW......................... ................................................................... ..
3
m in \o
Elk d i e t s ........................................................................................................
C arbohydrate a l l o c a t i o n s t r a t e g i e s ......... ............................
Agroovron spioatum ..................................... .........................................
F e s tu c a s o a b r e l l a i .......................................................................................
F e s tu c a id a h o e n s is ............. ..................................................................... ..
13
18
STUDY AREA........... ............................................... ........................................................
23
S i t e d e s c r i p t i o n .....................................................................................
S i t e s e l e c t i o n ................................
23
28
METHODS........... ........................................................................................ .......................
29
T r a n s e c t s . ..................................... .............................. .....................................
Chemical a n a l y s i s . .............................................
Near->infrared s p e c tr o s c o p y ........................
S t a t i s t i c a l a n a l y s i s . ..................................................................................
Chemical C o n s titu e n t D a t a . ............................... ............ ............................
Transe c t D ata................................. ............................................. .....................
29
31
31
33
33
34
RESULTS AND DISCUSSION........................ .............................................................. ..
35
Chemical c o n s t i t u e n t s . .....................
Chemical c o n s t i t u e n t r e l a t i o n s . . . . . ........... ....................................
Summer g r a z in g e f f e c t s ..........................
S p e c ies c o m p o sitio n .................................................................................... ..
'—"Winter g r a z in g e f f e c t s . ...................... ....................... ................................
S ta n d in g dead e f f e c t s ........... .................................... ..................... ............
W inter l e a f l o s s ......... .................................. ............ ................... .................
—^ W inter n u t r i e n t a l l o c a t i o n p a t t e r n s . . . . . ......... ................................
35
39
41,
47
50
50
55
58
SUMMARY AND CONCLUSIONS............................................... ......................................
61
M eth o d s.........................
Summer g r a z in g e f f e c t s . . . . . . .....................
61
62
V
TABLE OF CONTENTS - Continued
Page
W inter g r a z in g e f f e c t s . . . . . . ........... ........................................... ..
P la n t w in te r p h y s io lo g ic a l a c t i v i t y ..........................................
Management i m p l i c a t i o n s ................................... ................... ............ ..
62
63
63
LITERATURE CITED.'................... ............ ................................ .............................. ..
65
APPENDICES.................................. ................................................... ..............................
72
Appendix A - R egression E q u atio n s f o r NIR P r e d ic te d
Chemical C o n s titu e n t V a lu e s ...................... ............ ..................................
Appendix B - Frequency of P h e n o lo g ical S t a g e s . . . . . ......... ..
73
79
vi
LIST OF TABLES
Table
1
2
3
4
5
Page
Rough fe s c u e ( F e s c ) , Idaho f e s c u e (F eid ) and b lu e bunch w h e a tg ra s s (Agsp) in s e a s o n a l e lk d i e t s i n
Montana, s e l e c t e d s t u d i e s ............................. : .......... ....................... ..
4
The p r o g r e s s io n o f bluebunch w h e a tg ra s s p h en o lo g ic a l
development i n th e N o rth ern R o c k i e s . .................................
7
N u t r i t i o n a l p r o f i l e s o f bluebunch w h e atg rass from f a l l
dormancy through s p r in g g r o w th ...........................................
9
The p r o g r e s s io n o f rough fe s c u e phenolo g ic a l develop­
ment i n A lb e rta and B r i t i s h Columbia..................................
15
N u t r i t i o n a l p r o f i l e s o f rough fe s c u e from f a l l dormancy
through s p r in g gro w th ...................................................................
19
6
F o u r -y e a r a v e ra g e s o f semimonthly a n a ly s e s o f Idaho
fe s c u e (dry m a tte r b a s i s ) (McCall 1939) ...................... ............... 20
7
N u t r i t i o n a l p r o f i l e s o f Idaho fe s c u e from f a l l dormancy
through s p r in g g r o w t h . . ............ ..........................................
8
9
22
Sampling s i t e d e s c r i p t i o n s on t h e South Crow w in te r
r a n g e ................................................................................
2
NIR p r e d i c t e d w in te r chemical c o n s t i t u e n t v a lu e s f o r
bluebunch w h e atg rass and the p h e n o lo g ic a l s ta g e when
g r a z in g had o c c u rre d d u rin g t h e prev io u s s u m m e r ............... 36
10
NIR p r e d ic te d w in te r chemical c o n s t i t u e n t v a lu e s f o r
rough fe scu e and th e p h e n o lo g ic al s ta g e when g r a z in g had
o c c u rre d d u rin g the p re v io u s s u m m e r . . . , ..................................... 37
11
NIR p r e d ic te d w in te r chemical c o n s t i t u e n t v a lu e s f o r
Idaho fe s c u e and the p h e n o lo g ic a l s ta g e when g r a z in g
o c c u rre d d u rin g th e p re v io u s summer............................................. 38
12
C lu s t e r a n a l y s i s o f w in te r bluebunch w h e atg rass chemical
c o n s t i t u e n t s ................................................ ............................................... 40
13
C lu s t e r a n a l y s i s o f w in te r rough fe s c u e chem ical
c o n stitu e n ts.
......... ................................ ...................
42
C lu s t e r a n a l y s i s o f w in te r Idaho fe s c u e chem ical
c o n s t i t u e n t s . ............................................................... .................
43
14
v ii
LIST OF TABLES - Continued
T a b le
15
P age
Comparisons o f NIR p r e d ic te d chem ical c o n s t i t u e n t mean
v a lu e s f o r rough fe sc u e ( F e s c ) , Idaho fe scu e (F e id )
and bluebunch w h e atg rass (A gsp)................................. ..
48
16
Number of rough fe sc u e ( F e s c ) , Idaho fe scu e (F e id ) and
bluebunch w h e a tg ra ss (Agsp) on each sampling s i t e i n
t h e spring* I 984 . . . . . . . . . . . e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 9
17
F req u e n c ies and number of p l a n t s grazed (G) and w ith
s ta n d in g dead p r e s e n t (SD) fo r rough fe scu e ( F e s c ) ,
Idaho fe sc u e (F e id ) and bluebunch w h e atg rass (Agsp)
p l a n t s on each s i t e . . . . ........... ...........................................
51
Average rough f e s c u e h e i g h t s on s i x sampling d a t e s f o r
p l a n t s w ith o u t (no SD) and w ith (SD) s ta n d in g dead a t
the b e g in n in g o f new growth i n 1 9 8 4 . . ......... .................
52
Average bluebunch w h e a tg ra ss h e i g h t s on s i x sam pling
d a t e s f o r p l a n t s w ith o u t (no SD) and w ith (SD) s ta n d ­
i n g dead a t th e b e g in n in g o f new growth i n 1984............. ..
53
18
19
20
Average Idaho f e s c u e h e i g h t s on s i x sampling d a t e s f o r
p l a n t s w ith o u t (no SD) and w ith (SD) s ta n d i n g dead a t
the b e g in n in g o f new growth i n 1984...................... ....................... 54
21
R eg ressio n e q u a tio n s used f o r th e NIR p r e d i c t i o n of
rough fe sc u e chemical c o n s t i t u e n t v a l u e s . . . . . . . . . . . . . . . . . 74
22
R eg ressio n e q u a tio n s used f o r th e NIR p r e d i c t i o n of
Idaho fe s c u e chemical c o n s t i t u e n t v a l u e s ........... ..
75
23
R egression e q u a tio n s used f o r th e NIR p r e d i c t i o n of
bluebunch w h e atg rass chemical c o n s t i t u e n t v a l u e s . . . . . . . . . 76
24
NIR p r e d ic te d rough fe scu e summer chemical c o n s t i t u e n t
v a lu e s and t r e a t m e n t s .......................................................
77
NIR p r e d ic te d Idaho fe scu e summer chemical c o n s t i t u e n t
v a lu e s and t r e a t m e n t s . . .
78
25
v iii
LIST OF FIGURES
F ig u re
Page
1
The l o c a t i o n o f th e Elkhorn Mountains i n M ontana.. . . . . . . . 24
2
The Elkhorn Mountains showing t h e Crow Creek w in te r
r a n g e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . o . . . . . . . 25
3
Drawing o f th e Muddy Lake and M iddle p a s tu r e s o f the
South Crow a l l o t m e n t showing the study s i t e l o c a t i o n s . . . . 26
4
Frequency of Agsp p h en o lo g ic a l s t a g e s a f t e r two t r e a t ments on th e JGI s i t e * s p rin g 1984@.. . . . . . . o . . . . . . . . . . . . .
80
Frequency o f Agsp p h en o lo g ic a l s ta g e s a f t e r two t r e a t ­
m e n ts on t h e J G I I s i t e , s p r i n g 1 9 8 4 . . . . . . . . . . . . . . . . . . . . . .
81
Frequency of Agsp p h en o lo g ic a l s t a g e s a f t e r two t r e a t ­
ments on the ML s i t e , s p rin g 1984 .............................
82
5 .
6
7
Frequency of Agsp p h en o lo g ic a l s ta g e s a f t e r two t r e a t ­
ments on th e SCI s i t e , s p rin g 1984................................................. 83
8
Frequency of Agsp p h en o lo g ic a l s ta g e s a f t e r two t r e a t me n t s on th e S CU s i t e, s p rin g 19 84 . . . . . . . . . . . . . . . . . O . . . .
9
10
11
12
13
14
15
84
Frequency of Feid p h en o lo g ic a l s ta g e s a f t e r two t r e a t ­
ments on th e JGI s i t e , s p rin g 1984 .......................................... - •. 85
Frequency of Feid p h en o lo g ic a l s t a g e s a f t e r two t r e a t ­
ments on th e J G II s i t e , s p rin g 1984..........................................
86
Frequency o f Feid p h e n o lo g ic a l s t a g e s a f t e r two t r e a t ­
ments on the ML s i t e , s p rin g 1984 ...................... ..
87
Frequency o f Feid p h e n o lo g ic a l s ta g e s a f t e r two t r e a t ­
ments on th e SCI s i t e , s p rin g 1984 ........................................
88
Frequency of Feid p h en o lo g ic a l s ta g e s a f t e r two t r e a t ­
ments on th e SCII s i t e , s p rin g 1984
......... ..
89
Frequency of Fesc p h en o lo g ic a l s ta g e s a f t e r two t r e a t ­
ments on th e JGI s i t e , s p rin g 1984................................................
90
Frequency of Fesc ph en o lo g ic a l s ta g e s a f t e r two t r e a t ­
ments on th e J G II s i t e , s p rin g 1 9 8 4 . . . ....................
91
ix
LIST OF FIGURES - C ontinued
F ig u re
16
Page
Frequency of Fesc p h e n o lo g ic a l s t a g e s a f t e r two t r e a t ­
ments on the ML s i t e , s p rin g 1984 . =
92
17
Frequency of Fesc p h e n o lo g ic a l s ta g e s a f t e r two t r e a t ­
ments on th e SCI s i t e , s p rin g 1 9 8 4 . . . . . . . . . . . . . . . . . . . . . . . 93
18
Frequency of Fesc p h e n o lo g ic a l s ta g e s a f t e r two t r e a t ­
ments on th e SCII s i t e , s p rin g 1984................
94
X
ABSTRACT
E lk -L iv e s to c k S tu d i e s i n Montana w as e s t a b l i s h e d t o s t u d y e l k l i v e s t o c k i n t e r a c t i o n s t h r o u g h c o o p e r a t i v e r e s e a r c h p ro g r a m s o f
v a r io u s s t a t e and f e d e r a l a g e n c ie s.
The o b j e c t i v e s o f E lk -L iv e sto ck
S tu d ie s w ere:
I) t o d e t e r m i n e t h e i n f l u e n c e o f v a r i o u s c a t t l e
m anagem ent p r a c t i c e s on e l k and t h e i r h a b i t a t s , 2 ) t o e v a l u a t e
a l t e r n a t i v e m anagem ent p r a c t i c e s , and 3 ) t o d e v e l o p b e n e f i c i a l
g u i d e l i n e s f o r dual use of th e a v a i l a b l e re s o u r c e .
T h is r e s e a r c h
p r o j e c t f e l l w i t h i n o b j e c t i v e one, and i t s o b j e c t i v e s w e r e :
I) t o
q u a n t i f y th e e f f e c t s of summer c a t t l e g r a z in g a t v a r io u s p h o n o lo g ical
s t a g e s on t h e w i n t e r n u t r i t i o n a l v a l u e s o f t h r e e m o u n t a i n
b u n chgrasses, and 2 ) to q u a n tif y th e e f f e c t s o f w i n t e r e lk g ra z in g on
t h e growth and developm ent i n t h e f o ll o w i n g s p r in g o f th e same th r e e
sp e c ie s.
Rough f e s c u e ( F e s t u o a s c a b r e l l a ). I d a h o f e s c u e ( F e s t u c a
i d a h o e n s i s ) and b lu e b u n c h w h e a t g r a s s (A g ro o v ro n s b i c a t u m ) w ere t h e
s p e c ie s s tu d ie d . The study was conducted on th e South Crow e lk w in te r
range i n t h e Elkhorn Mountains o f Montana.
For o b j e c t i v e one a random e x p e r im e n ta l d e sig n of one f a c t o r w ith
t h r e e l e v e l s and n in e t r e a t m e n t s was used. The g r a s s s p e c ie s were th e
f a c t o r l e v e l s and the t r e a tm e n ts c o n s is te d of the p h e n o lo g ie s! s ta g e
when g r a z i n g had o c c u r r e d p l u s an u n g r a z e d s t a g e . W in te r c h e m i c a l
c o n s t i t u e n t v a lu e s w ere the dependent v a r i a b l e s . For o b j e c t i v e two a
randomized block d e sig n h av in g one f a c t o r w ith th r e e l e v e l s and two
t r e a t m e n t s i n f i v e b l o c k s was u sed . S i t e s , s p e c i e s , g r a z i n g , and
s t a n d i n g d e a d w e re t h e b l o c k s , f a c t o r and tw o t r e a t m e n t s ,
r e s p e c t i v e l y . The d e p e n d e n t v a r i a b l e s w e r e p l a n t h e i g h t a n d
phenolo g ic a l s ta g e on s i x s p r i n g sam p lin g d a te s .
None of the summer g ra z in g t r e a t m e n t s a f f e c t e d rough fe sc u e o r
bluebunch w h e a tg ra s s w i n t e r n u t r i t i o n a l v a lu e s , and grazed p l a n t s were
s i m i l a r to u ngrazed p l a n t s .
Some p h e n o l o g i c a l s t a g e s o f summer
g r a z i n g w e re r e f l e c t e d i n Id a h o f e s c u e w i n t e r c h e m i c a l c o n s t i t u e n t
v a l u e s , b u t no m an ag em en t s i g n i f i c a n c e c o u ld be a t t a c h e d t o th e
d iffe re n c e s.
W in te r g r a z i n g d i d n o t a f f e c t s p r i n g l e a f l e n g t h s in-vrough fe s c u e or bluebunch w h e a tg ra s s, and only o c c a s i o n a l l y a ffe c te d ^
Id a h o f e s c u e s p r i n g g r o w th .
The p r e s e n c e o f s t a n d i n g d e a d i n t h e
s p r in g r e s u l t e d i n lo n g e r Idaho f e s c u e le a v e s i n th e n ex t s p rin g .
N e ith e r rough fe sc u e nor bluebunch w h e a tg r a s s s p rin g l e a f l e n g t h s were
a f f e c t e d by th e p resen ce of o v e r w i n t e r i n g r e s i d u a l m a t e r i a l .
These
r e s u l t s w e re d i s c u s s e d from th e p e r s p e c t i v e o f th e w i n t e r
p h y s io lo g ic a l a c t i v i t y of each s p e c ie s of g ra s s .
I
INTRODUCTION
M anagem ent and r e s e a r c h
g ro u p s
have
spent
m uch
effo rt
i n v e s t i g a t i n g th e v a r io u s a s p e c t s and r e l a t i o n s h i p s o f la n d , l i v e s t o c k
and e lk management.
One c u r r e n t program i s "E lk -L iv e sto c k S tu d ie s i n
M ontana", an i n t e r a g e n c y e f f o r t to stu d y e l k - l i v e s t o c k i n t e r a c t i o n s .
" E l k - L i v e s t o c k S t u d i e s " w as i n i t i a t e d i n I 982 t o :
I ) . d e te rm in e th e
i n f l u e n c e o f v a r i o u s c a t t l e m anagem ent p r a c t i c e s on e l k and t h e i r
h a b i t a t , 2 ) t o e v a l u a t e a l t e r n a t i v e m anagem ent p r a c t i c e s and 3 ) t o
develop b e n e f i c i a l g u i d e l i n e s f o r dual use of th e a v a i l a b l e re so u rce .
In a d d i t i o n to Montana S t a t e U n iv e r s ity ,
Range Experim ent S t a t i o n i n M issoula,
F ish ,
the In te rm o u n ta in F o r e s t and
MT,
th e Montana Department of
W i l d l i f e and P a r k s and t h e U n i v e r s i t y o f M ontana a r e a l s o
in v o lv e d i n t h e program.
T h is p r o j e c t f e l l w i t h i n t h e f i r s t o b j e c t i v e o f " E l k - L i v e s t o c k
S tu d ie s" . I t s o b j e c t i v e s were:
I j t o q u a n t i f y th e e f f e c t s of summer
c a t t l e g r a z in g a t v a r io u s p h en o lo g ic a l s ta g e s on th e r e s u l t i n g w in te r
n u t r i t i o n a l v a lu e s of th r e e m ountain bu n ch g rasses, and 2 ) t o q u a n tif y
t h e e f f e c t s o f e l k w i n t e r g r a z i n g on g r o w th and d e v e lo p m e n t o f t h e
same g r a s s e s i n th e f o ll o w i n g sp rin g .
Two d e c i s i o n s w e r e made w h ic h had i m p o r t a n t i n f l u e n c e s on t h e
p ro je c t.
F irst,
i t was d ecided t h a t t h i s p r o j e c t would d e s c r ib e one
a n n u a l c y c l e o f g r o w th and u t i l i z a t i o n fro m an i n d i v i d u a l p l a n t
p e rsp e c tiv e .
As a r e s u l t f i e l d w o r k w as d e s i g n e d t o m i n i m i z e t h e
im p a c t of th e p r o j e c t .
lin e
Some e f f e c t s o f t h i s d e c is io n w ere th e use of
tra n s e c ts in s te a d
o f c a g e s and a n im a ls f o r d e f o l i a t i o n
2
t r e a tm e n ts .
s a c rific e
This approach r e s u l t e d i n reduced t r e a t m e n t c o n tr o l and
of e x p e rim e n ta l
d e sig n i n t e g r i t y .
Some
co m b in a tio n s w ere m is s in g o r had s m a ll sample s iz e s .
tre a tm e n t
The s t a t i s t i c a l
a n a l y s i s was l e s s rig o r o u s than could have been ach ie v e d from a more
c o n t r o l l e d ex p erim en t.
The o t h e r i m p o r t a n t d e c i s i o n w as t h e s e l e c t i o n o f key s p e c i e s .
Rough f e s c u e ( F e s t u c a s c a b r e l l a
T o rr.),
Id ah o
fescu e
(F e stu c a
i d a h o e n s i s E lm e r .) and b lu e b u n c h w h e a t g r a s s (A g ro o v ro n s p i c a t u m
(P u r s h )
S c rib n .
and S m ith ) w e re
chosen,
because
th e y
are
b o th
i m p o r t a n t h a b i t a t c o m p o n e n ts and m a j o r c o n s t i t u e n t s o f w i n t e r e l k
d i e t s i n M ontana. Of th e t h r e e s p e c i e s I d a h o f e s c u e w a s t h e m o st
abundant and rough fe sc u e th e l e a s t .
S i t e s w ith both rough fescu e and
bluebunch w h e a tg r a s s appeared to be e c o to n e s between p a tc h e s of rough
fe s c u e and t h e g e n e r a l l y o c c u r rin g bluebunch w h e atg rass.
S e le c t io n of
th e se key s p e c ie s reduced the p o t e n t i a l sam pling s i t e s from the e n t i r e
a v a i l a b l e a r e a to s p e c i f i c s i t e s w here a l l t h r e e s p e c ie s o ccu rred .
3
LITERATURE REVIEW
E lk d i e t
se le c tio n
s t u d i e s fro m
t h e m o u n ta in o u s r e g i o n s o f
M ontana a r e p r e s e n t e d t o s u p p o r t t h e c h o i c e of ro u g h f e s c u e , Id a h o
fescue
and
b lu e b u n c h w h e a t g r a s s a s
th e
stu d ie d
sp e c ie s.
G rass
c a rb o h y d ra te a l l o c a t i o n s t r a t e g i e s and growth re s p o n se s t o g ra z in g a r e
summarized.
P h e n o lo g ic a l developm ent,
g r a z in g re s p o n s e s and n u t r i e n t
c o n c e n tr a tio n p a t t e r n s f o r rough fe s c u e and bluebunch w h e a tg ra s s a r e
p re s e n te d .
Only s ea so n al p ro g r e s s io n of n u t r i e n t c o n c e n tr a tio n s a r e
p r e s e n t e d f o r Id a h o f e s c u e .
The l i t e r a t u r e on bluebunch w h e a tg ra ss
was much more e x te n s iv e and com plete th a n f o r e i t h e r fe sc u e .
ELK DIETS
E lk fo o d h a b i t s t u d i e s i n t h e Rocky M o u n ta in r e g i o n h a v e been
e x te n s iv e ly review ed by Kufeld (1973).
From th e se s t u d i e s i t could be
concluded t h a t e lk s e l e c t e d t h e g r e e n e s t fo r a g e a v a i l a b l e .
During th e
seaso n s of p la n t dormancy t h e i r d i e t s were composed p re d o m in a n tly of
g ra m in o id s.
T h i s g e n e r a l p a t t e r n h a s a l s o b een o b s e r v e d i n M ontana
s t u d i e s ( t a b l e I) .
A d d i t i o n a l l y , i n M ontana, a l a r g e p r o p o r t i o n o f
t h e s p e c ie s s e l e c t e d w ere th e dominant a v a i l a b l e bunchgrasses.
In th e
E lk h o r n M o u n ta in s o f c e n t r a l M ontana up t o 62% o f e l k w i n t e r d i e t s
w ere composed o f
Idaho fe s c u e , rough fe s c u e and bluebunch w h e a tg ra ss,
t h e m o s t a b u n d a n t and a v a i l a b l e b u n c h g r a s s e s on t h e w i n t e r r a n g e
(DeSimone e t a l . 1984).
Other s t u d i e s from Montana a l s o r e p o r te d t h a t
rough fe s c u e , Idaho f e s c u e and bluebunch w h e a tg ra s s w ere im p o r ta n t e lk
w i n t e r d i e t components ( t a b l e I).
.;d
Table I
Rough fe sc u e ( F e s c ) , Idaho fe s c u e (F e ld ) and bluebunch w h e a tg ra s s (Agsp)
i n se a s o n a l e lk d i e t s i n Montana, s e l e c t e d s t u d i e s .
SPFClFS
FESC FEID AGSP
% OF DIET
48
10
6
I
27
21
48
49
8
5
55
5
39
24
I
6
"2
3
88% GRASS1
84% GRASS
100% GRASS-LIKE2
100% GRASS-LIKE
4
3
48
13
16
13
1}
5
37
90% GRASS1
90% GRASS
18 . 35
67% GRASS1
Ii3
43
IV
27
11
SEASON
LOCATION
WINTER
WINTER
SPRING
SPRING
April
May
Elkhorns MT
■Elkhorns MT
Ml CROHI STOLOGICAL DeS imone et AL. 1984
Feeding S ite Exam Gordon 1968
Elkhorns MT
Sun River MT
Feeding S ite Exam Stevens 1965
Feeding S ite Exam Knight 1961
Little Belts
MT
Feeding S ite Exam Kirsch 1963
Rumen Analysis
spring
WINTER
. SPRING
WINTER
SPRING
WINTER
WINTER
WINTER
WINTER
WINTER
SPRING
WINTER
March
WINTER
SPRING
SPRING
NAT'L Bison
Range MT
Sun River MT
Sun River
Canyon MT
Gravelly Mtns
MT
Gravelly Mtns
MT
Gallatin Canyon
MT
Rocky Mtn Front
MT
Sun River. MT
GRASSES ON THE STUDY'AREAT' ------- d
^
Z S K
rI
All Eestuca
^
TECHNIOllF ■■
Rumen Analysis
CITATION
Morris & Schwartz 1957
Feeding S ite Exam Casagranda s J anson 195
Feeding S ite Exam Schallenberger 1966
Feeding S ite Exam Eustace 1967
Clipping
Rumen Analysis
Rouse 195/
Feeding S ite Exam Constan 1967
Ml CROHISTOLOGICAL Kasworm et al. 1984
Feeding S ite Exam Knight 1970
SPECIES, BUT THE KEY SPECIES WERE THE DOMINANT
I s^
' ^ e W^ d H r^ V' DED■BY SPECIES' BUT THE KEY SPECIES WERE
in the study combined.
the
5
CARBOHYDRATE ALLOCATION STRATEGIES
C a r b o h y d r a te a l l o c a t i o n p a t t e r n s on a s e a s o n a l b a s i s and i n
re s p o h se to d e f o l i a t i o n and g ra z in g t r e a t m e n t s have been review ed by
T r l i c a (1977)» W h ite (1973) and Cook ( 1966).
Some g e n e r a l i z a t i o n s
s u p p o r t e d by a l l t h r e e a u t h o r s i n c l u d e d : I ) a l t h o u g h i t w as seld o m
emphasized t o t a l p la n t s o lu b le c a rb o h y d ra te s in c r e a s e d w ith m a tu r ity ,
2) carb o h y d rate a llo c a tio n
p a tte rn s
v a rie d
w ith
p h e n o lo g ic a l
developm ent and e n v iro n m e n ta l c o n d itio n s , and 3) th e l e v e l of s o lu b le
c a r b o h y d r a t e s l o c a t e d i n a p a r t i c u l a r t i s s u e a t any g i v e n t i m e w as
a l s o a f u n c t i o n o f c o n c u r r e n t p l a n t p h y s i o l o g i c a l a c t i v i t i e s and
b i o t i c and a b i o t i c e n v iro n m e n ta l in f l u e n c e s .
G razin g ,
a b i o t i c e n v iro n m e n ta l in flu e n c e ,
c arb o h y d rate a llo c a tio n
p a tte rn of a l l
sp e c ie s
has m o d ified th e
stu d ie d
to d a te .
S t u d i e s u s i n g m o re s o p h i s t i c a t e d t e c h n i q u e s have d e m o n stra te d t h a t
i n d i v i d u a l s p e c ie s responded d i f f e r e n t l y to g ra z in g (e.g, C aldw ell e t
al.
1981).
However some i n t e r s p e c i f i c g e n e r a l i z a t i o n s w ere ap p aren t.
I f g r a z in g ended b e fo re th e i n i t i a l grow th p e rio d ended,
d e f o l i a t i o n w as r e d u c e d .
i n j u r y from
F a l l r e grow t h w as much m ore i m p o r t a n t t o
p l a n t s g ra z ed through th e e n t i r e i n i t i a l growth p e rio d th a n to p l a n t s
g r a z e d e a r l y i n t h e i n i t i a l g ro w th p e r i o d o r u n g r a z e d (McLean and
Wikeem 19 8 5 a , b).
G ra m ^ n o id s t h a t w e re e n t i r e l y d o r m a n t d u r i n g th e
w i n t e r w ere more dependent on c a rb o h y d ra te r e s e r v e s f o r s p r in g growth
i n i t i a t i o n th a n th o s e w i t h p h o to s y n t h e t i c a l l y a c t i v e o v e r w i n t e r i n g
l e a v e s ( R o b e r t s o n and W oolhouse 1 9 8 4 b ). G ra m in o id s t h a t d ep en d e d on
o v e r w in te r in g p h o t o s y n t h e t i c a l l y a c t i v e l e a v e s t o f u e l i n i t i a l s p r in g
g r o w th
r e q u i r e d some i n t a c t l e a v e s i n t h e s p r i n g ( R o b e r ts o n and
6
Woolhouse 1984b).. There may a ls o be a d d itio n a l c arb o h y d ra te a ll o c a t i o n
p a tte r n s n o t y e t e lu c id a te d .
AGROPYROM SPICATUM
Quinton e t a l .
(1982) s tu d ie d bluebunch w h e a tg ra s s v e g e t a t i v e and
r e p r o d u c t i v e grow t h i n t h e a b s e n c e o f g r a z i n g fr o m I 9 7 1 t o I 973 i n
i n t e r i o r B r i t i s h Columbia. V e g e ta tiv e grow th i n i t i a t i o n was d eterm in ed
by e x t r a p o l a t i o n o f y i e l d and l e a f g r o w th c u r v e s t o z e r o . G row th
in itia tio n ,
e s ti m a te d from y i e l d c u rv e s,
o c cu rred from mid-March to
May. S o il te m p e r a tu r e s a t 10 cm w ere 6°C and average a i r te m p e ra tu re s
w i t h i n I 0 C.
S o i l m o i s t u r e a t 28 and 75 cm had b e en r e c h a r g e d by
s p r i n g snow m e l t .
e a rlie r.
le ss
L e a f g r o w th e s t i m a t e s o f g r o w th i n i t i a t i o n w e re
S o il te m p e ra tu re s w ere s t i l l n e a r O0C a t 10 cm and snow m elt
t h a n one w eek e a r l i e r .
A 6 °C s o i l
tem p eratu re
at
g ro w th
i n i t i a t i o n w as s u p p o r t e d by o t h e r s ( H a r r is . 1967 and A n d e rso n and
McNaughton I 973).
V e g e t a t i v e g r o w th i n B r i t i s h C o lu m b ia l a s t e d tw o m o n th s w i t h
culms r e a c h in g a maximum h e ig h t o f 19 cm.
Average culm le n g t h reached
38.2 cm i n W a s h in g to n ( R ic k a r d e t a l . I 975) and 16.4 cm a t D u b o is, ID
( B l a i s d e l l 1958).
so il
L e a f g ro w th. c e a s e d i n e a r l y summer.
m o is tu re l e v e l s w ere v a r i a b l e ,
A sso c ia ted
and a i r te m p e ra tu re s w ere s t i l l
c o o le r than optimum growth te m p e ra tu re s (DePuit and C ald w e ll 1975).
F a l l g r o w th i n B r i t i s h C o lu m b ia o c c u r r e d o n ly i n t h e y e a r w i t h
both th e w e t t e s t and w arm est September and October.
P r e c i p i t a t i o n and
te m p e r a tu r e w ere 135$ and 120$ o f av erag e, r e s p e c t i v e l y .
P h en o lo g ical
developm ent p a t t e r n s from s e v e ra l a r e a s a r e r e p o r te d i n t a b l e 2.
T a b le -2 .
The p ro g r e s s io n o f bluebunch w h e a tg ra s s phenolo g ic a l development i n t h e
N o rth ern R ockies.
PHENOLOGICAL
STAGE
snow f r e e
growth i n i t i a t i o n
boot
e a r l y head
f u l l head
e a r ly flow er
f u l l flow er
seed r i p e
seed s h a t t e r
growth c e s s a t i o n
f a l l growth
Years
L o c a tio n
C it a t i o n
DATE
mid-Ap
Jn I
e a r l y Jn
mid-J n
m id - la te J n
l a t e J n - mid-J l
la te Jl
l a t e J I - e a r l y Ag
e a r l y J n - mid-J I
l a s t h a l f Oc
1971-73
Kamloops, B.C.
Quinton e t a l .
1982
Ap
Ap
Ma
Ma
Jn
I
11
16
28
6
Jn
Jn
Jl
Jn
18
28
21
7
1937-40
Dubois, ID
B la isd e ll e t a l.
1952
.
Mr 30
Ap 4
Ma 22
'
l a t e Ap
Ma - e a r l y Jn
J n 14
Jn
. Jl
Jl
Jl
25
19
24
8
1941-47
Dubois, ID
B la isd e ll
1958
la te J l
la te J l
J l & Ag - Oc
M isso u la, MT
Daer and W illa rd
1981
8
C lim a tic p a t t e r n s m o d ified th e r a t e of phenolo g ic a l p r o g r e s s io n
(Quinton e t a l .
1982).
Where low s p r i n g te m p e r a tu r e s were l i m i t i n g ,
e a r l y p h e n o l o g i c a l s t a g e s w e re c o m p r e s s e d i n t i m e .
d ro u g h t was l i m i t i n g ,
la te r
p h o n o lo g ic a l
sta g e s
Where summer
w e re
sh o rte r.
B l a i s d e l l and P e c h a n e c (1949) r e p o r t e d t h a t c l i p p i n g d u r i n g s p r i n g
g r o w th d i d n o t a f f e c t t h e t i m e o r r a t e o f p h o n o l o g i c a l d e v e lo p m e n t.
S p ik e n u m b e rs w e r e n o t w e l l c o r r e l a t e d w i t h t i l l e r num ber o r b a s a l
a re a of p la n ts .
S ev eral
stu d ie s
c o n s t i t u e n t s h av e
a n a ly z in g
e sta b lish e d
b lu e b u n c h
seasonal v a ria tio n
a s s o c i a t e d w ith fo r a g e q u a l i t y .
in
c o m p o n e n ts
S to d d a rt (1946) r e p o r t e d t h a t t o t a l
c arb o h y d rates in c re a se d a s v e g e t a t i o n
c a rb o h y d ra te s ,
wh e a t g r a s s c h e m i c a l
m a tu re d .
N o n stru c tu ra l
presumably a v a i l a b l e f o r p la n t u t i l i z a t i o n or s to ra g e ,
i n c r e a s e d i n t h e s p r i n g ( C a l d w e l l e t a l . 1981).
D aer and W i l l a r d
(1 9 8 1 ) fo u n d t h a t minimum t o t a l n o n s t r u c t u r a l c a r b o h y d r a t e s (TNC)
l e v e l s corresponded t o i n i t i a l grow th and e a r ly boot s t a g e s and th a t
th e
m axim um
TNC o c c u r r e d
at
th e
la te
boot
sta g e .
"O th er
c a r b o h y d r a t e s " ( t o t a l c a r b o h y d r a t e s - c e l l u l o s e ) r e a c h e d maximum
c o n c e n tr a tio n s i n th e f a l l and minimum l e v e l s i n l a t e w i n t e r ( S to d d a rt
1946). C e l l u l o s e c o n t e n t r a n g e d f r o m 24% i n e a r l y s p r i n g t o 31% i n
la te
sp rin g ,
33% i n l a t e
summer and 43% i n l a t e w i n t e r .
L ig n in
i n c r e a s e d f r o m 4% i n e a r l y s p r i n g g r o w th t o 14% i n f a l l and 18% i n
l a t e w in te r .
P r o t e i n c o n c e n tr a tio n d e c lin e d from 26% i n new s p rin g
g r o w th t o 13% when h e a d s w e re e m e r g in g and 3% i n t h e f a l l .
th ro u g h s p r in g n u t r i e n t c o n te n ts a r e r e p o r t e d i n t a b l e 3„
F a ll
Table 3 .
Stage or
Season
N u t r i t i o n a l p r o f i l e s o f bluebunch w h e a tg ra s s from f a l l dormancy
through s p r in g growth.
To t a l 1 Crude
CH^O F iber
%
mature
FALL
Nov
J an
Apr
Oct
POST
SEED
SHATTER
NEW ,
GROWTH
MATURE
FOLIAGE
%
NFE
%
37.8
35.1
31.2
29.5
28.5
30.7
33.4
49.4
46.7
46.4
44.2
43.7
48.4
39.6
20.6
30.2
33.9
36.6
44.1
43.5
Crude
Cellulose Lignin Protein Ash
%
%
.
%
%
Citation
5.0
3.2
5.2 Knight et AL. 1908
12.5 McCreary 1927
7.9 McCreary 1931
f:6
10.9 .
it
4.2
13.4
"
3.0 , 8.2
2.9
10.1
McCall 1940
62.5
• 24,2
4.0
25.5
7.8
3.7
26.2
86.6
32.9
12.9
2.9
7.5
OLD
GROWTH
88.6
43.2
17.7
2.5
6.5
Ii
NEW
GROWTH
63.8
24.8
5.6
23.4
9.0
■I
DRY
MATURE
83.3
31.8
14.5
4.6-
Nbv
NEW
GROWTH
(4/20)
DRY
MATURE
(9/15)
(4/1)
(4/15)
(9/15)
1Total
carbohydrates 1
"
Blaisdell et al. 1952
"
"
8.6
8.9 .
Stoddart 1946
10
Herbage removal a f f e c t e d bluebunch w h e a tg ra s s n u t r i e n t c o n te n t i n
the same summer.
S to d d a r t (1946) r e p o r te d t h a t one m id s p rin g c lip p in g
r e s u l t e d i n t h e b e s t o v e r a l l f o r a g e q u a l i t y a s r e f l e c t e d by h i g h e r
p r o t e i n and lo w e r l i g n i n c o n t e n t s .
S u b s e q u e n t c l i p p i n g s a t o n e - or
two-week i n t e r v a l s r e s u l t e d i n s l i g h t f o r a g e v alu e d e c l i n e s ,
but a l l
c l i p p i n g t r e a t m e n t s had h ig h e r f o r a g e v a l u e s th a n u n d i p p e d c o n t r o l
p la n ts.
C aldw ell e t a l .
(1981) r e p o r t e d t h a t p r o t e i n c o n c e n tr a tio n s
i n re g ro w th l e a v e s w ere s i m i l a r to th o se o f u n d ip p e d l e a v e s one month
e a rlie r.
B o lto n and Brown (1980) a s s o c i a t e d h i g h e r n i t r o g e n (N)
c o n c e n tr a tio n s w ith h ig h e r p h o to s y n th e tic a c t i v i t y .
C ald w ell e t a l.
( 1981) m e a s u re d h i g h e r p h o t o s y n t h e t i c a c t i v i t y i n r e g r o w t h t i s s u e
c o m p ared t o u n c l i p p e d p h o t o s y n t h e t i c t i s s u e . N i t r o g e n i n c o r p o r a t e d
i n t o re g ro w th was new a s s i m i l a t e , because r o o t and crown N p o o ls did
no t d e c li n e (C ald w ell e t a l ,
1981).
T o t a l n o n s t r u c t u r a l c a r b o h y d r a t e p o o ls i n blue bunch w h e a tg ra ss,
w hich w ere c o n c e n tra te d i n stem s and s h e a th s i n i n t a c t p l a n t s ,
w ere
d i m i n i s h e d by c l i p p i n g and r e m a i n e d low i n r e g r o w t h co m p ared t o
un c lip p e d p l a n t s (C aldw ell e t a l . 1981).
" O th e r c a r b o h y d r a t e s " a l s o
d e c lin e d i n c lip p e d p l a n t s (S to d d a rt 1946). C arbohydrate p o o ls i n th e
r o o t s d e c l i n e d r a p i d l y w i t h g r o w th i n i t i a t i o n , i n c r e a s e d s l i g h t l y
d u r i n g t h e a c t i v e g r o w th p e r i o d , and w e r e c o m p l e t e l y r e p l e n i s h e d
f o l l o w i n g a c t i v e v e g e t a t i v e grow th (Daer and W illa rd 1981, McCarty and
P r i c e 1 942 and M c I l v a i n e 1942).
Removal o f p h o t o s y n t h e t i c t i s s u e
d i s r u p t e d t h i s p a t t e r n of a l l o c a t i o n t o th e r o o t s ,
re d u c in g r e s e r v e s
a v a i l a b l e f o r g r o w th i n i t i a t i o n t h e f o ll o w i n g s p r i n g (B l a i s d e l I and
Pechanec I 949).
11
B lu e b u n c h w h e a t g r a s s f o l i a g e r e m o v a l by b u r n i n g , g r a z i n g o r
c l i p p i n g i n any s e a s o n r e d u c e d p r o d u c t i o n i n a t l e a s t t h e f o l l o w i n g
y e a r ( S to d d a rt 1946, B l a i s d e l l and Pechanec 1949, Mueggler 1972, Sauer
1978, W ilm s e t a l . 1980 and McLean and Wikeem 1985a).
t i l l e r s r a r e l y produced new t i l l e r s (C ald w ell e t a l .
damage r e s u l t e d from both s p r i n g and f a l l
The c l i p p e d
1981).
S e v e re st
c lip p in g s (S to d d a r t 1946).
A s i n g l e c l i p p i n g t r e a t m e n t d u rin g t h e most m e t a b o l i c a l l y demanding
g r o w th p e r i o d s , a c t i v e r e p r o d u c t i v e s t a g e s o r c a r b o h y d r a t e r e s e r v e
re p le n ish m e n t
p e rio d s,
a ll
red u ced y i e l d
in th e fo llo w in g year
(McCarty and P r ic e 1942, M cIlv ain e 1942, B l a i s d e l l and Pechanec 1949
and W ils o n e t a l 1 9 6 6 ).
I f c lip p in g k i l l e d a t i l l e r i t u s u a lly d ie d
im m e d ia te ly o r w ith the o n s e t of d rought (C aldw ell e t a l . 1981).
When
c o m p e titio n from s u rro u n d in g v e g e t a t i o n was reduced, ex trem e c lip p in g
d u r i n g c r i t i c a l g r o w th s t a g e s w a s much l e s s d e t r i m e n t a l ( M u eg g le r
1972).
The key f a c t o r m i t i g a t i n g d e f o l i a t i o n damage appeared t o be th e
tim e and amount o f l e a f grow th re m a in in g f o ll o w i n g d e f o l i a t i o n (McLean
and Wikeem 1985a). P la n ts d e f o l i a t e d e a r l y i n th e s p r in g w ith s e v e ra l
weeks o f l e a f grow th f o l l o w i n g d e f o l i a t i o n had h ig h e r s u r v i v a b i l i t y
and v i g o r t h a n p l a n t s d e f o l i a t e d n e a r o r p a s t t h e end o f t h e l e a f
growth p e rio d (McLean and Wikeem 1985a,
S to d d a rt 1946 and B l a i s d e l l
and Pechanec 1949).
The e f f e c t s o f c l i p p i n g f a l l r e g r o w t h w e re m ore v a r i a b l e and
d i f f i c u l t to e x p la in .
B l a i s d e l l and P e c h an e c (1 9 4 9 ) r e p o r t e d t h a t
f a l l r e grow t h w as v e r y i m p o r t a n t t o p r o d u c t i o n i n t h e n e x t y e a r .
C lipped p l a n t s w ith re g ro w th y ie ld e d , i n the next grow ing season, 270$
12
o f p l a n t s w i t h no r e g r o w t h .
P l a n t s w i t h r e grow th a l s o p ro d u c e d 2.5
tim e s a s many flo w e r s t a l k s a s p l a n t s w ith o u t r e growth.
McLean and
Wikeem (1985a) r e p o r t e d t h a t c l i p p i n g p l a n t s w ith 5.1 g / p l a n t o f f a l l
re g ro w th
re su lte d
in
th a n
s ig n ific a n tly
c lip p in g
p la n ts
h ig h e r
w ith
p ercen t
2.7 o r
k ill
in
1.6 g o f
th e
fo llo w in g
year
fa ll
re g ro w th ,
s u g g e s t i n g t h a t some t h r e s h o l d l e v e l o f f a l l r e g r o w t h
p ro d u c tio n was r e q u i r e d t o have an im pact.
C lip p in g blue bun ch wheat g r a s s s t a n d i n g d e ad i n J a n u a r y r e d u c e d
p ro d u c tio n i n t h e f o l l o w i n g s p r in g ,
l e a f and culm y i e l d d e c lin e d 28%,
l e a f l e n g t h d e c r e a s e d 25% and l o s s o f , s t a n d i n g d ead d e c r e a s e d 21%.
F a c to r s a s s o c i a t e d w ith r e p r o d u c tio n w ere not s i g n i f i c a n t l y a f f e c t e d
( S a u e r 1978).
The c a e s p i t o s e g r o w th fo rm p r o v i d e s m i c r o s i t e m o d i f i c a t i o n s
(Sauer 1978, Chapin e t a l . 1979, C aldw ell e t a l . 1981 and Smith e t aL
1983).
By removing s ta n d i n g dead m a t e r i a l some b e n e f i c i a l e f f e c t s o f
the c a e s p ito s e grow th form w ere l o s t .
Sau er (1978) h y p o th e size d t h a t
f r e e r a i r c ir c u la tio n in c re a se d m o istu re s tr e s s ,
c a u s in g sto m a te
c lo s u r e , re d u c in g CO2 u p tak e and co n se q u e n tly p ro d u c tio n . Sauer (1978)
a l s o su g g este d p h o t o i n h i b i t i o n o f p h o to s y n th e s is .
However, blue bunch
w h e a t g r a s s l e a f d i s p l a y p a t t e r n s and culm d e n s i t i e s e x p o s e d m o s t
le a v e s to d i r e c t s o l a r r a d i a t i o n i n i n t a c t bunches (C a ld w e ll e t a l.
1 981).
B lu e b u n ch
w h e a tg ra ss
le a v e s
had
a
p h o to s y n th e tic r a t e p er a r e a (C aldw ell e t a l . 1981).
re la tiv e ly
h ig h
C o m p e titio n from
su rro u n d in g v e g e t a t i o n may have c o n t r i b u t e d t o d e c r e a s e d p r o d u c t i o n
( M u eg g le r 1 9 7 2 ).
O th e r r e s e a r c h e r s ( B l a i s d e l l and Pechanec 1949 and
13
W ils o n e t a l . I 966) h a v e r e p o r t e d t h a t o v e r w i n t e r i n g f a l l r e g r o w t h
becam e th e f i r s t
c a rb o h y d rate
so u rce in
th e s p r i n g ,
because
c a rb o h y d ra te r e s e r v e s d e c lin e d s h o r t l y a f t e r r a t h e r th a n c o n c u r re n tly
w ith
g r o w th i n i t i a t i o n
(C a ld w ell
e t a l.
1981).
A b sence o f f a l l
reg ro w th f o r e a r l i e s t s p r in g p h o t o s y n th e tic a c t i v i t y may a ls o i n h i b i t
s p rin g p ro d u c tio n .
F a l l r e g r o w t h h a s a l s o b e en r e p o r t e d a s a m a jo r
s o u rc e of new t i l l e r s (D aer-and W illa r d 1981).
All of th e se s t u d i e s used "treatm en ts which removed a l l t i l l e r s .
B lu e b u n c h w h e a tg r a .s s r e s p o n s e s t o
s e le c tiv e h e rb iv o ry
w ould be
s i m i l a r , bu t l e s s s e v e re , th a n those p re s e n te d above.Some l e a v e s would
re m a in ung ra z ed and could c o n tin u e p h o to s y n th e tic a c t i v i t y .
Stems and
s h e a th s com prised m ajor p o o l s o f n o n s t r u c t u r a l c a r b o h y d r a t e s w h ic h
c o u l d a p p a r e n t l y c o n t r i b u t e t o r e g r o w t h p ro d u c tio n (C ald w ell e t a l .
1981).
B lu e b u n c h w h e a t g r a s s s tem and s h e a t h p h o t o s y n t h e s i s was a s
p r o d u c t i v e and e f f i c i e n t a s l e a f p h o t o s y n t h e s i s ( C a l d w e l l e t a l .
1981).
T h e re fo re s e l e c t i v e g ra z in g o f l e a v e s would no t e l i m i n a t e a l l
p h o to sy n th e tica lly a c tiv e tissu e .
FESTUCA SCABRELLA
Rough fe s c u e grow th and p h en o lo g ic a l development w ere s tu d ie d a t
two s i t e s n e a r K am loops, B r i t i s h C o lu m b ia ( S t o u t e t a l . 1981) and a t
S ta v e ly and L e th b rid g e ,
A lb e rta (Jo h n sto n and McDonald 1967).
and p h e n o l o g i c a l d e v e l o p m e n t v a r i e d
b e tw e e n y e a r s
and
Growth
site s.
E n v i r o n m e n t a l f a c t o r s seem ed t o c o n t r o l i n i t i a t i o n o f g r o w th , b u t
p h e n o ty p ic a d a p ta tio n r e s u l t e d i n d i f f e r e n t r a t e s o f p h o n o lo g ic a l
pro g ress a t d if f e r e n t
site s.
For t h e t h r e e y e a r s s t u d i e d i n B.C.
14
( S t o u t e t a l . I 981) t h e r e w e re no m ore t h a n t h r e e w eek s v a r i a t i o n i n
the tim in g of any p h o n o l o g i c a l . sta g e .
S t o u t e t a l . (1 9 8 1 ) c a l c u l a t e d fr o m y i e l d c u r v e s t h a t g ro w th
i n i t i a t e d i n m i d - A p r i l w i t h s o i l t e m p e r a t u r e s o f 2 -3 ° C a t 10 cm.
J o h n sto n and McDonald (1967) r e p o r t e d new growth i n e a r l y May w ith 20
cm s o i l te m p e r a tu r e s o f 2°C and c a l c u l a t e d t h a t l e a f grow th had been
i n i t i a t e d two weeks e a r l i e r .
in itia tio n .
Culm growth began s i x weeks a f t e r growth
Growth i n i t i a t i o n was c o r r e l a t e d w i t h s o i l t e m p e r a t u r e
but no t a i r te m p e ra tu re .
C oncurrent s o i l m o is tu re had been co m p le tely
re c h arg e d by snow m e lt (B a ile y and Anderson 1978).
C e s s a tio n o f v e g e t a t i v e growth was keyed t o s o i l m o is tu re (Stout
e t a l . 1981) and m o d ified by a i r te m p e ra tu re i n f l u e n c e s on p l a n t w a te r
re la tio n s.
P la n t
w e ig h t
g a in
c o n tin u e d
fo llo w in g
le a f
g ro w th
c e s s a t i o n , b e c a u s e p h o t o s y n t h e s i s c o u ld c o n t i n u e a t v a p o r p r e s s u r e
\
l e v e l s which i n h i b i t e d c e l l expansion (Hsiao 1973).
C oncurrent s o i l
t e m p e r a t u r e s w e re n o t warm enough t o i n h i b i t g r o w th ( S m o lia k and
J o h n s t o n I 9 6 8 ).
I n A l b e r t a f o l i a g e had e n t e r e d w i n t e r d o rm ancy
e a r l y O ctober (Jo h n sto n and McDonald 1967).
by
P a t t e r n s of p h e n o lo g ie s!
development from B r i t i s h Columbia and A lb e rta a r e r e p o r t e d i n t a b l e 4.
J o h n s t o n a n d M cD o n a ld ( 1 9 6 7 ) r e p o r t e d
th a t rough fe s c u e
r e p r o d u c tiv e culm p ro d u c tio n ranged from 6.7 to 121.7 culm s p er p la n t.
S eed p r o d u c t i o n w as e r r a t i c
and d i f f i c u l t
to r e l a t e
to
sp e c ific
en v iro n m e n ta l cues (Jo h n sto n and McDonald 1967 and S to u t e t a l . 1981).
A b u n d an t f l o w e r p r o d u c t i o n y e a r s h a v e i n c l u d e d
1952,
1964, 1966
( J o h n s t o n and McDonald 1967) and 1972 ( S t o u t e t a l . 1 9 8 1 ).
F lo ra l
p r im o r d ia w ere i n i t i a t e d i n th e f a l l w ith seed p ro d u c tio n i n th e
Table 4 .
The p ro g r e s s io n o f rough fe sc u e phenolo g ic a l development i n A lb e rta
and B r i t i s h Columbia.
PHENOLOGICAL
STAGE
DATE
Ma I
Ma 18 Ma 8
Ma 25 Ma 30
J n I Ma 30
J n 15 Jn 6
J n 22 Jn 13
J n 29
- ■
J l 11
Jl 5
J l 24 J l 5
J l 7 -J n 13
none
Oc 16
boot
e a r l y head
f u l l head
e a r ly flo w er
f u l l flo w e r
end o f flo w er
seed i n milk
seed i n dough
seed r i p e
seed s h a t t e r
growth c e s s a t i o n
f a l l growth
Ma 27
Jn 3
J n 10
J n 17
J n 24
Jl I
-.
J l 22
J n 17
Se 3
Years
L o c a tio n
E le v a tio n
C ita tio n s
1972
1973
1971
Hamilton, B.C.
1158 m
S to u t e t a l . 1981
Ma 12
Ma 19
Ma 22
Jn 2
Jn 9
Jn 23
J n 30
Jl 7
J l 14
J n 23
Se 15
Ma 10
Ma 17
Ma 24
Ma 31
Jn 7
Jn 14
Jl 5
J l 17
J n 28
none
Ap
Ma
Ma
Ma
Jn
Jn
Jn
Jn
Jl
Jl
Ma
Oc
30
7
14
28
5
12
19
26
3
3
28
3
1972
1971
1973
East Mara, B.C.
854 m
S to u t e t a l . 1981
la te Jn
mid J l
'
e a r ly Au
1964 -1966
L e th b rig e , Al.
Johnston and McDonald
1967
16
f o l l o w i n g summer ( J o h n s t o n and McDonald 1967), a t r a i t s h a r e d w i t h
some o t h e r
n o rth e rn
g r a s s e s in c lu d in g Idaho fe s c u e
J o h n s t o n and McDonald 1967),
W ith in t h i s
(Hodgson i n
tim e span a v a r i e t y of
p o t e n t i a l l y i n f l u e n t i a l e n v ir o n m e n ta l e v e n t s c o u ld o c c u r . I n i t i a t e d
p r im o r d ia w ere cold t o l e r a r i t (Johnston and McDonald 1967).
Of 3,780
p rim o rd ia examined from 1963 through 1967, only one p e rc e n t s u f f e r e d
fro s t k illin g .
Minimum te m p e ra tu re s d u rin g t h i s p e rio d w ere -AO0C a i r
te m p e ra tu re and -11 and - 90 C; s o i l te m p e r a tu r e s a t 10 and 20 cm d ep th s,
re sp e c tiv e ly .
S t i l l o n ly a b o u t 35% o f i n i t i a t e d f l o r a l p r i m o r d i a
s u c c e s s f u l l y developed i n t o seedheads, and s p r in g management did not
ap p ea r to a f f e c t seedhead p ro d u c tio n .
i
t o 97$ (Jo h n sto n and McDonald 1967),
Seed g e r m in a tio n ran g ed from 86
i
Rough f e s c u e h ad c h a r a c t e r i s t i c s w hich w ere i n d i c a t i v e of both
g r a z i n g t o l e r a n c e and s u s c e p t i b i l i t y ( J o h n s t o n and McDonald 1967).
M e r i s t e m s o f v e g e t a t i v e cUlms w e r e n o t e l e v a t e d , and t h e r a n g e o f
r e p r o d u c tiv e , t o v e g e t a t i v e dulm r a t i o s , 0.299 to 0.001, was r e l a t i v e l y
low even i n y e a r s o f high seed p ro d u c tio n .
Grazing s u s c e p t i b l e t r a i t s
in c lu d e d th e e r e c t c a e s p i t o s e grow th form which made a high p ro p o r tio n
of p h o to s y n th e tic t i s s u e a v a i l a b l e to h e rb iv o ry .
T i l l e r i n g i n rough
f e s c u e was n e i t h e r v ig o ro u s nor s t im u la te d by th e removal o f culms or
e l e v a t e d m e r i s t e m s . Under g r a z i n g management, such a s r e s t - r o t a t i o n
and d e f e r r e d g r a z in g methods w ith m oderate u t i l i z a t i o n ,
rough fe s c u e
h a s re c o v e re d and m a in ta in e d i t s e l f (Anderson and Franzen 1983).
Rough f e s c u e w as dam aged by c o n s e c u t i v e y e a r s o f h e a v y summer
g r a z in g (Jo h n sto n and McDonald 1967), But p r o t e c t i o n from even l i g h t
g r a z in g ,
15$ to 25$ u t i l i z a t i o n f o r 12 y e a r s , in c r e a s e d th e p resen ce
J
17
of
ro u g h f e s c u e
i n an e x c l o s u r e ( J o h n s t o n 1961).
produced v a r y in g r e s u l t s (B a ile y and Anderson 1978).
cm o f new grow th r e s u l t e d i n reduced,
S p rin g b u rn in g
B urning w ith 10
but r e c o v e r in g canopy coverage
f o r a t l e a s t t h r e e y e a r s (39% of unburned i n y e a r one, 69% i n y e a r two
and 90% i n y e a r t h r e e ) .
When b u rn e d w i t h 4 cm o f new g r o w t h , canopy
c o v e r a g e d e c l i n e d l e s s and p r o d u c t i o n w as n o t a f f e c t e d .
F lo ra l
p r i m o r d i a w e r e u n a f f e c te d ^ by b u r n i n g u n t i l th e y had b e en e l e v a t e d
above th e r o o t crown i n the s p r in g , when seedhead d e n s i t y was g r e a t l y
r e d u c e d (3% o f u n b u rn e d one y e a r p o s t b u rn ) ( B a i l e y and A n d erso n
1978).
B ased on a n e c d o t a l e v i d e n c e some ro u g h fe s c u e p o p u la tio n s have
had a l o n g h i s t o r y o f h e a v y w i n t e r g r a z i n g ( J o h n s t o n and McDonald
1967).
is
In a r e a s o f th e n o rth e rn G re at P l a i n s where Fescue g r a s s la n d
t h e d o m in a n t v e g e t a t i o n
ty p e ,
such a s
th e
C ypress H i l l s
in
s o u t h e a s t e r n A lb e rta , b u f f a l o summered on t h e open p l a i n s and w in te re d
i n th e h i l l s .
I n s u c h a r e a s r o u g h f e s c u e was s u b j e c t e d t o s e v e r e
w i n t e r g r a z in g ev ery y e a r , but r e s t e d t h e re m a in d er of th e year.
C h e m ic a l a n a l y s e s
have found
th a t,
except fo r
p ro te in ,
n u t r i t i o n a l components o f rough fe s c u e w ere r e l a t i v e l y s t a b l e through
t h e y e a r ( J o h n s t o n .a n d B ezeau 1962 and B ezeau and J o h n s t o n 1962).
P r o t e i n d e c re a se d from 13.7% i n e a r l y v e g e t a t i v e growth t o 4.7% i n th e
f a l l and 4.2% by t h e end o f w i n t e r . . O v e r w i n t e r p r o t e i n d e c l i n e w as
m ore v a r i a b l e t h a n i n o t h e r s e a s o n s .
Crude f i b e r and c e l l u l o s e
i n c r e a s e d t h r o u g h t h e g r o w in g s e a s o n f r o m 29.9 t o 33.4% and 3 3 «2 t0
3 8.4%, r e s p e c t i v e l y .
O v e r w i n t e r c r u d e f i b e r i n c r e a s e d t o 34.8% and
c e l l u l o s e t o 39.5%. I n v i t r o d ry m a t t e r d i s a p p e a r a n c e d ro p p e d fro m
18
48.5? i n e a r l y s p r i n g t o 3 2 . 1? by f a l l and 24.0? a t t h e end o f w i n t e r .
Rough f e s c u e n u t r i e n t c o n t e n t s fro m d o rm an cy th r o u g h s p r i n g a r e
r e p o r t e d i n t a b l e 5.
FESTUCA IDAHOENSIS
For I d a h o f e s c u e o n ly t h e s e a s o n a l p r o g r e s s i o n o f d i e t q u a l i t y
w i l l be d is c u s s e d .
Growth and p h o n o l o g i c a l d e v e lo p m e n t and g r a z i n g
e f f e c t s a r e n o t p r e s e n t e d b e c a u s e t h i s i n f o r m a t i o n w a s n o t fo u n d .
M c C a ll (1 9 4 0 ) r e p o r t e d
th a t
th e
ty p ic a l
fa ll
d e c lin e
in
grass
n u t r i t i o n a l v a lu e s was l e s s pronounced i n Idaho fe s c u e th a n i n o th e r
b u nchgrasses s tu d ie d .
I n p a r t i c u l a r c r u d e p r o t e i n (CP) d ro p p e d t o
4.6? i n Id a h o f e s c u e and 2.9? i n b lu e b u n c h w h e a t g r a s s w h i l e c ru d e
f i b e r i n c r e a s e d t o 27*2? i n I d a h o f e s c u e and 33.4? i n
w h e a tg ra s s.
b lu e b u n c h
In d i g e s t i o n t r i a l s u s in g lam bs on m atu re f o r a g e s , Idaho
f e s c u e had h i g h e r t o t a l
d ig e stib le
n u t r i e n t s and p o s i t i v e
c ru d e
p r o t e i n d i g e s t i o n c o e f f i c i e n t s (? d i e t a r y CP - ? f e c a l CP). Bluebunch
w h e a t g r a s s h a d l o w e r t o t a l d i g e s t i b l e n u t r i e n t s and n e g a t i v e c r u d e
p r o t e i n d i g e s t i o n c o e f f i c i e n t s . Idaho f e s c u e was more n u t r i t i o u s th a n
bluebunch w h e a tg ra s s.
McCall (1939) d e te rm in e d biw eekly chem ical c o n s t i t u e n t s o f Idaho
fe s c u e over a f o u r y e a r p e rio d ( t a b l e 6). Crude f i b e r ro s e s l i g h t l y a s
(
Idaho fe s c u e m atured from "new grow th" t o "m aturing" s t a g e s , changed
l i t t l e i n t h e " f a l l " and r o s e a g a i n t h r o u g h th e " w i n t e r . "
Only "new
grow th" and " w in te r" c ru d e f i b e r l e v e l s w ere s i g n i f i c a n t l y d i f f e r e n t .
N itro g e n f r e e
e x tra c t
(NFE) i n
" w in te r"
and
"new
g r o w th " w e re
sig n ific a n tly
l o w e r t h a n i n " m a t u r i n g " and " f a l l " s t a g e s .
Crude
Table 5 .
N u t r i t i o n a l p r o f i l e s o f rough fe s c u e from f a l l dormancy through
s p r in g growth.
S tage or
Season
T o ta l1 Crude
CH2O" F ib e r
Ash
IVDMD
%
%
%
13.7
4 .7
4 .2
6.9
8 .5
8.6
48.5
32.1
24.0
4 .2
10.2
2.1
12.3
30.3
12.0
10.1
37.2
5.4
9.8
%
le af
cured
w eath ered
29.9
33.4
34.8
seed shed
(9/15)1
w eathered
(1 1 /2 5 )
36.0
82.6
le af
(6 /1 6 )
p a r t l y cured
( 8/ 10)
^T o tal c a rb o h y d ra te s
Crude
P r o te in
42.5
NFE
C e llu lo s e
%
%
33.2
38.4
39.5
46.6
C ita tio n
Jo h n sto n & Bezeau 1962
and
Bezeau & Jo h n sto n 1962
McLean & T is d a le I960
C larke & T is d a le 1945
20
Table 6 .
F o u r-y e a r a v e ra g e s o f semimonthly a n a ly s e s o f Idaho fe scu e
(dry m a tte r b a s i s ) (McCall 1939).
Period
M ature:
F i r s t p a rt of J a
Last p a rt o f J a
F i r s t p a r t of Fb
L a s t p a r t of Fb
F i r s t p a r t o f Mr
L a s t p a r t o f Mr
F i r s t p a r t o f Ap
New growth:
F i r s t p a r t o f Mr
L a s t p a r t o f Mr
F i r s t p a r t o f Ap
Middle p a r t of Ap
L a s t o f Ap-f i r s t of Ma
Middle p a r t of Ma
L a s t p a r t o f Ma
F ir s t part of Jn
L ast p a rt of Jn
F ir s t part of J l
L a s t p a r t of J l
F i r s t p a r t of Au
L a s t p a r t of Au
F i r s t o f Sp
Middle p a r t o f Sp
L a s t o f S p - f i r s t of Oc
Middle p a r t o f Oc
L a s t p a r t o f Oc
Middle p a r t o f Nv
L a s t o f N v - f i r s t o f Do
Middle p a r t o f Dc
L a s t p a r t of Do
Ash
Crude
P r o te i n
%
%
C arb o h y d rates
Crude N-f r e e
F ib e r e x t r a c t
%
$
16.2
16.4
15.3
16.6
18.8
18.5
18.6
4 .7
4 .7
4 .8
4 .9
5.1
5 .0
5.1
32.6
32.7
33.6
32.8
31.1
31.9
30.8
43.6
4 3 .2
44.1
43.0
42.3
42.4
4 2.5
10.0
9 .9
10.4
11.1
13.2
12.9
12.1
13.1
13.2
14.0
13.7
13.7
14.3
14.6
13.6
15.4
17.0
16.4
16.3
14.8
15.5
17.8
17.4
19.0
18.9
16.2
13.8
11.1
9.2
7 .8
6 .5
5 .0
4.6
4.1
4.1
3 .9
3 .7
4 .2
4.6
4 .7
4.4
4 .2
4.4
4 .6
26.0
24.1
25.1
24.0
28.3
28.2
30 .2
31.2
30.3
31.6
31.4
31.2
30.9
29.0
30.5
30.7
29.0
30.3
31.6
32.9
32.7
30.9
42.4
43.0
42.0
4 4 .8
41.2
4 4.3
45.2
44 .5
46.7
46.3
4 6.8
47.7
47.1
48 .7
48.9
46.2
45.9 .
45.0
44.4
45 .2
44.4
43 .8
21
p r o t e i n d e c lin e d from "new growth" t o "m aturing" s t a g e s th e n rem ained
s t a b l e th r o u g h t h e " f a l l " and " w i n t e r " .
"New g r o w th " c r u d e p r o t e i n
was s i g n i f i c a n t l y h ig h e r th a n t h e o th e r t h r e e s ta g e s w hich w ere not
s i g n i f i c a n t l y d i f f e r e n t from each o th e r .
g r o w th " t h r o u g h t h e y e a r .
Ash ro s e s t e a d i l y from "new
S e a s o n a l c h e m i c a l c o n s t i t u e n t s o f Id a h o
f e s c u e from s e v e r a l s t u d i e s a re summarized i n t a b l e 7. .
Table 7 .
N u t r i t i o n a l p r o f i l e s o f Idaho fe s c u e from f a l l dormancy through
s p r in g growth.
S tage or
Season
Crude
F ib e r
NFE
C e llu lo s e
%
%
%
Crude
P r o te i n ■Ash
%
%
IVDMD
%
C ita tio n
new growth
fa ll
w in te r
26.0
30.9
32.4
42.4
45.3
43.3
17.4
4 .4
4 .8
10.0
16.0
16.8
McCall 1939
p o st seed
sh a tte r
27.2
38.7
4 .6
14.6
McCall 1940
seed r i p e
cured
l a t e w in te r
32.7
31.9
33.4
7 .6
4 .9
4 .5
7.5
10.6
9.0
35.0
35.9
38.8
It.
n
27.0
18.8
14.6
Joh n sto n & Bezeau 1962
and
Bezeau & Jo h n sto n 1962
23
STUDY AREA
The f i e l d work f o r t h i s p r o j e c t was conducted i n Muddy Lake and
J e n k i n ’ s G ulch d r a i n a g e s ,
t r i b u t a r i e s o f Crow C re ek l o c a t e d on t h e
s o u t h e a s t e r n end of th e Elkhorn Mountains on th e Muddy Lake and Middle
p a s t u r e s o f t h e S o u th Crow c a t t l e a l l o t m e n t on t h e H e le n a N a t i o n a l
F o r e s t , M ontana ( f i g . I t o 3).
SITE DESCRIPTION
The c l i m a t e i n t h e r e g i o n i s m o d i f i e d c o n t i n e n t a l w i t h l a r g e
a n n u a l and d a i l y
te m p e ra tu re v a r ia tio n s
(DeSimone e t
a l.
1984).
E le v a tio n and p r e c i p i t a t i o n range from 1464 m to 2100 m and 51 cm to
76 cm, r e s p e c t i v e l y .
The g e n e r a l v e g e t a t i o n a s p e c t on th e study a re a
i s a r o l l i n g g r a s s la n d w ith s te e p c o n i f e r draws and b ro a d e r cottonwood
bottom s. Slo p es and rid g e to p s have s to n y , loamy s o i l s . The v e g e t a t i o n
on t h e u p la n d s i t e s i s d o m in a te d by b u n c h g r a s s e s .
u b iq u ito u s.
Idaho fe s c u e i s
Rough f e s c u e o c c u r s on t h e s l i g h t l y m o i s t e r s i t e s and
b lu e bunch w h e a tg r a s s on most o th e r s i t e s .
Each t r a n s e c t w a s l o c a t e d on a s i t e w h e re a l l
o ccu rred .
th re e sp e c ie s
T h e re w e r e f i v e s i t e s c a l l e d S o u th Crow I (SCI) and I I
( S C I I ) , J e n k i n ' s G ulch I (J G I) and I I ( J G I I ) and Muddy Lake (ML).
e le v a tio n ,
The
a s p e c t and s l o p e o f e ac h s i t e a r e r e p o r t e d i n t a b l e 8.
Based on o b s e r v a tio n s d u rin g f ie ld w o r k ,
th e J e n k in ’s Gulch s i t e s w ere
c o o le r and m o i s t e r and t h e South Crow s i t e s w ere warmer and d r i e r th an
the o t h e r s .
A pproxim ately 200 e lk have w in te r e d i n th e study a re a s in c e th e
1960’s (DeSimone e t a l . 1984).
G e n e r a l l y w i n t e r r a n g e i s u s e d fro m
■x
w ^
%
y
v iv x .
f1
IM iiii
-i^-Ic, -I
V.
4# &
%f,1%
dC ZL:
F ig u re I .
The l o c a t i o n o f the Elkhorn Mountains i n Montana.
; ./
m
■}
z
/'
•>?
;
25
Iaf" Ir
VJ
[2:
^
I ie n d a le * ,
„
U
'
d
"
:
F ig u r e 2 .
s r
-
I
I
M I
»
r
The Elkhorn Mountains showing t h e Crow Creek w in te r range
26
MUDDY LAKE
PASTURE
MIDDLE-PASTURE
SCII
South
LEGEND
A llotm ent
b o u n d r y a nd d i v i s i o n
S ite
Names
ML
JGI
JGII
SCI
SCII
Muddy L a k e
J e n k i n 1 s Gulc h I
J e n k i n 1s Gulch I I
S o u t h Crow I
S o u t h Crow I I
fences
/ --------- £
F ig u re 3. Drawing o f th e Muddy Lake and Middle p a s tu r e s o f th e South
Crow a llo tm e n t showing the study s i t e l o c a t i o n s .
27
Table 8.
Sampling s i t e d e s c r i p t i o n s on th e South Crow w in te r ra n g e .
SITE
TRANSECTS
number
ELEVATION
m eters
ASPECT
SLOPE
$
Muddy Lake
5
2048
. SSW
5
J e n k i n ’ s Gulch I
6
2073
W
6
J e n k i n 1s Gulch I I
3
1987
sw .
18
South Crow I
5
1926
SSE
24
South Crow I I
5
1939
SW
12
t h e c l o s e o f h u n t i n g s e a s o n u n t i l t h e summer r a n g e s a r e s n o w - f r e e ,
u s u a ll y December to June. The South Crow w i n t e r range i s no t a t y p i c a l
e lk w in te r range f o r s e v e ra l re a so n s.
The e l k p o p u l a t i o n i s one o f
th e
M ontana
densest
and m o st
c o m m u n ic a ti o n ) .
S e rv ic e la n d ,
p ro d u c tiv e in
(DeSimone
p ersonal
D u r in g a l l s e a s o n s 95$ o f t h e e l k a r e on F o r e s t
th u s a v o i d i n g c o n f l i c t s w i t h s u rro u n d in g la n d owners.
T y p i c a l l y 80$ o f t h e e l k i n t h e w e s t e r n U.S. w i n t e r on p r i v a t e l a n d
(DeSimone e t a l . I 984).
A lso t h e S o u th Crow w i n t e r r a n g e i s n o t i n
the F o r e s t S e rv ic e tim b e r base.
So d u rin g th e w i n t e r e lk w i n t e r range
i s th e p rim ary use of th e area.
S o u th Crow a l l o t m e n t h a s had a t h r e e p a s t u r e d e f e r r e d r o t a t i o n
g r a z in g system s in c e 1970.
w ith c a lv e s (USDA 1970).
t o O c t o b e r 15.
There a r e 2517 AUM’s a l l o c a t e d to 604 cows
In 1983 th e g r a z in g seaso n r a n from June 10
The M id d le p a s t u r e w as g r a z e d fr o m J u n e 10 i n t o
O c to b e r and t h e Muddy Lake p a s t u r e fro m A ugust 10 i n t o O c to b e r . The
c o m b in a tio n of h ig h q u a l i t y ra n g e ,
w e l l managed l i v e s t o c k and a
28
h e a lth y e lk p o p u la tio n made t h i s stu d y s i t e an example of a d e s i r a b l e
situ a tio n .
SITE SELECTION
The Crow C reek d r a i n a g e w as s e l e c t e d b e c a u s e tw o p r e v i o u s MSU
m a s t e r ’ s p r o j e c t s ( S t e v e n s 1-965-and G ordon 1968) h av e s t u d i e d e l k l i v e s t o c k i n t e r a c t i o n s i n t h i s a r e a and r e p o r t e d h a b i t a t d e s c r i p t i o n s ,
i
se a so n a l h a b i t a t use and d i e t a r y i n f o r m a t i o n . A d d i t i o n a l l y c u r r e n t
e l k w i n t e r l o c a t i o n d a ta w ere made a v a i l a b l e by th e Elkhorn Mountains
W i l d l i f e M o n ito rin g Program (EMWMP).
t u r e s w ere s e l e c t e d , p a r t i a l l y ,
s t u d i e s on the same a re a.
The Middle and Muddy Lake pas­
to f a c i l i t a t e c o o r d in a tio n w ith EMWMP
29
METHODS
TRANSECTS
When s e t t i n g up t h e f i e l d w o r k a p r i m a r y c o n s i d e r a t i o n w as t h e
i m p l e m e n t a t i o n o f p r o c e d u r e s w h ic h w o u ld m i n i m i z e t h e p r o j e c t ’ s
e f f e c t s on t h e p l a n t s , a n im a ls and s i t e s in v o lv e d .
A s tu d y of c r e s te d
w h e a t g r a s s d e f o l i a t i o n p a t t e r n s ( N o r to n and J o h n s o n 1981) r e p o r t e d
t h a t very
little
re g ra z in g of in d iv id u a l
e x te n siv e p a stu re s itu a tio n .
w ith in th is a r tic le ,
p l a n t s o c c u r r e d i n an
B ased on t h i s and s u p p o r t i n g e v id e n c e
i t w as d e c i d e d t h a t p r o t e c t i o n o f i n d i v i d u a l
p l a n t s from e x c e s s iv e u t i l i z a t i o n would not be a problem and permanent
t r a n s e c t s w ere s e le c te d .
Gammon and R o b erts (1978) u sed t r a n s e c t s t o
study d e f o l i a t i o n p a t t e r n s o f i n d i v i d u a l p l a n t s on s o u t h e r n A f r i c a n
v e ld ,
and r e p o r t e d t h a t two weeks betw een t r a n s e c t re a d in g s a llo w ed
i d e n t i f i c a t i o n of g r a z in g e f f e c t s and p h e n o lo g ic a l developm ent and so
tw o w eek s
p ro je c t.
b e tw e e n t r a n s e c t r e a d i n g s w a s a d o p t e d i n t h e c u r r e n t
The f i r s t s e t o f p l a n t h e i g h t r e a d i n g s w as s u b j e c t e d t o
S t e i n ' s t w o - s t a g e t e s t f o r s a m p le s i z e a d e q u a c y ( S t e e l and T o r r i e
1960), and i t was d e te rm in e d t h a t 15 t r a n s e c t s a d e q u a te ly sampled the
v a r i a t i o n i n l e a f le n g th f o r each s p e c ie s . T w en ty -fiv e t r a n s e c t s were
e s t a b l i s h e d t o allo w f o r u n fo resee n problem s.
The 25 t r a n s e c t s w e r e i n f i v e g r o u p s w i t h f i v e t r a n s e c t s p e r
g ro u p .
Each t r a n s e c t w as s u b j e c t i v e l y l o c a t e d
t o i n c l u d e ro u g h
fe s c u e ,
Idaho fe s c u e and bluebunch w h e a tg r a s s p l a n t s and ev id en ce of
w i n t e r e lk use and summer c a t t l e use. A t r a n s e c t was a perm anent 10 m
l i n e m a rk e d by f i x e d end p o i n t s .
When a t r a n s e c t w as r e a d a m e te r
30
t a p e was s t r e c h e d t i g h t l y
d ire c tio n .
b e tw e e n t h e s t a k e s a l w a y s i n t h e same
Canopy l i n e i n t e r c e p t f o r each bunch of each key s p e c ie s
was re c o rd ed .
Each bunch was assumed t o be an i n d i v i d u a l p la n t.
This
pro ced u re allo w ed i d e n t i f i c a t i o n o f i n d i v i d u a l p l a n t s th r o u g h tim e
w ith o u t d i s t u r b i n g p l a n t s o r m o d ify in g s i t e c h a r a c t e r i s t i c s .
T r a n s e c t s w e r e r e a d b iw e e k l y fro m A p r i l th r o u g h O c t o b e r , 1983.
H e ig h t
of
th e
lo n g e st
v e g e ta tiv e
u t i l i z a t i o n w ere re c o rd e d .
le a f,
p h e n o lo g ic a l
stag e
and
F e s c u e l e a f l e n g t h s w e r e m e a s u re d by
h o ld in g t h e le a v e s s t r a i g h t up and m e asu rin g th e lo n g e s t .
Bluebunch
w h e a t g r a s s had e l o n g a t e d v e g e t a t i v e c u lm s , so l e a f l e n g t h w as t h e
h e i g h t o f t h e h i g h e s t p o i n t o f t h e h i g h e s t l e a f on t h e culm .
p h e n o lo g i c a l
sta g e s
w ere
id e n tifie d ,
(I)
v e g e ta tiv e ,
E ight
(2)
boot
( s w e l l i n g v i s i b l e i n s h e a t h t o s e e d h e a d 50$ o u t o f t h e s h e a t h ) , (3)
e m e r g in g s e e d h e a d s ( s e e d h e a d s g r e a t e r t h a n 50$ o u t o f t h e s h e a t h t o
f u l l y e m e rg e d , b u t n o t f i l l i n g ) , (4) a n t h e s i s (any v i s i b l e a n t h e r s
r e g a r d l e s s o f c o n d itio n ) ,
(5) m ature f lo w e r s (from seedheads f i l l i n g
to
(6 )
fu lly
d e v e lo p e d ),
d isa rtic u la tio n ),
le av e s
d u lle r
(7) m a tu re f o l i a g e
green
re g re e n ih g of le a v e s).
At t h e
seed
o u tse t i t
th ro u g h
sh a tte r
(any
in d ic a tio n
of
(seedheads c o m p le te ly empty and
bro w n )
and
(8)
fa ll
r e g r o w th
(any
M a tu re f o l i a g e w a s th e o v e r w i n t e r i n g s t a g e .
w as h oped t h a t
some a c c e p t a b l e key t o a n d / o r
d e f i n i t i o n s o f p h e n o l o g i c a l s t a g e s w o u ld be f o u n d ; no n e w e re .
The
s t a g e s i d e n t i f i e d i n t h i s p r o j e c t w ere s e l e c t e d o u t o f ease of f i e l d
id e n tific a tio n .
F i e l d w o r k w as c o n t i n u e d fro m J a n u a r y th r o u g h e a r l y J u l y , I 984.
D uring t h i s w i n t e r , p r i m a r i l y i n March, p l a n t s w ith known u t i l i z a t i o n
31
h isto rie s
fro m
t h e p r e v i o u s summer w e r e c o l l e c t e d f o r
ch em ical
a n aly sis.
These sam p les were s to r e d a t ~50°C and low h u m id ity u n t i l
a n a l y z e d . C l i p p i n g r e m o v e d a p l a n t fro m t h e s a m p le p o p u l a t i o n . Also
d u r i n g March t h e t r a n s e c t s w e r e r e a d t o d e t e r m i n e w i n t e r l e a f l o s s
w h ic h w as m e a s u r e d i n tw o w ay s, w i n t e r g r a z i n g and t h e p r e s e n c e o f
s ta n d in g dead a t the b e g in n in g of new s p r i n g growth.
On some or a l l
o f t h e f o l l o w i n g d a t e s , d e p e n d in g on snow c o v e r , t h e t r a n s e c t s w e re
read:
A p r i l 19, May 10, May 18, J u n e 10, J u n e 18 and J u n e 25. H e ig h t
o f t h e l o n g e s t v e g e t a t i v e l e a f and p h e n o lo g ic a l s ta g e w ere re c o rd ed
u s in g the same te c h n iq u e s and d e l i n i a t i o n s p re v io u s ly d e sc rib e d .
CHEMICAL ANALYSIS
Twenty p e rc e n t o f th e c o l l e c t e d sam p les w ere an aly z e d u s in g th e
f o l l o w i n g p ro c e d u re s:
m acro -K jeld ah l p r o t e i n (PRO),
dry m a t t e r (DM)
and a s h (ASH) f o l l o w i n g AOAC (1 9 7 0 ) , n e u t r a l d e t e r g e n t f i b e r (NDF),
a c i d d e t e r g e n t f i b e r (ADF) and a c i d d e t e r g e n t l i g n i n
(ADL) f o ll o w i n g
G oering and VanSoest (1970) and i n v i t r o o rg a n ic m a t t e r d is a p p e a ra n c e
(IVOMD)
(H a rris
1 970).
A d d itio n a lly
c e llu lo se
(CELL)
and
h e m ic e llu lo s e (HG) were c a l c u l a t e d (CELL = ADF - ADL, HG = NDF - ADF)
(Goering and VanSoest 1970).
Twelve rough f e s c u e , 10 Idaho fe s c u e and
7 bluebunch w h e a tg r a s s v a lu e s f o r each c o n s t i t u e n t were, d eterm in ed .
NEAR-INFRARED SPECTROSCOPY
The chem ical a n a l y s i s d a ta w ere used t o c a l i b r a t e a TECHNICOR 400
INFRALYSER,
a n e a r-in fra re d
(NIR) s p e c t r o p h o t o m e t e r ,
th a t
th e n
p r e d i c t e d t h e c h e m i c a l c o n s t i t u e n t s f o r t h e r e m a i n i n g 80% o f t h e
32
sam ples.
T his NIR s p e c tro p h o to m e te r had 19 f i l t e r s w ith w av elen g th s
from 1445 t o 2348 nm, i n c l u s i v e .
I t w as d e c i d e d , d u r i n g t h e c o u r s e o f t h e p r o j e c t ,
p r e d i c t i o n o f chem ical c o n s t i t u e n t s f o r t h r e e re a s o n s :
t o u se NIR
I) to save on
t h e t i m e and e x p e n s e i n v o l v e d i n l a b o r a t o r y a n a l y s i s , 2) t o b e g in a
NIR d a ta base f o r ra n g e la n d f o r a g e s and 3) because sam p les w ere w in te r
c o l l e c t i o n s o f u s u a lly grazed p l a n ts ,
t h e am ount o f m a t e r i a l p e r
sample was to o s m a ll to p e rm it com plete chem ical a n a l y s i s .
Also f o r
t h e s e t h r e e r e a s o n s some o f t h e u s u a l NIR p r o c e d u r e s (AACC 1962,
N o r r i s e t a l . 1976 and Shenk e t a l .
1981) w ere m o d ified .
All sam ples w ere a n a ly z e d by NIR and t h e lo g v a lu e s re c o rd e d i n a
file .
The l a r g e s t s a m p l e s w e re c h o s e n f o r th e c h e m i c a l a n a l y s i s
d e s c r ib e d above.
Some sam ples w ere to o s m a ll to a llo w a n a l y s i s o f a l l
the chem ical c o n s t i t u e n t s .
fo r
o n ly
p art
of
th e
As a r e s u l t ,
c o n stitu e n ts,
some sam p les w ere analyzed
some w e re a n a l y z e d f o r a l l
c o n s t i t u e n t s b u t n o t r e r e a d by NIR, and some w e re a n a l y z e d f o r a l l
c o n s t i t u e n t s p l u s r e r e a d by NIR.
T h i s m eans t h a t s o u r c e s o f e r r o r
a s s o c i a t e d s i z e of sam p les and i n t r a s a m p l e c o n s t i t u e n t c o r r e l a t i o n s
w ere not a d e q u a te ly d e a l t w ith .
D e te rm in a tio n o f f i l t e r c o e f f i c i e n t s and an i n t e r c e p t v a lu e f o r
each c o n s t i t u e n t was done u s in g the "All P o s s ib le S u b s e ts R egression"
program (Dixon 1981).
a r e i n A p p e n d ix A.
The r e g r e s s i o n e q u a tio n s f o r each c o n s t i t u e n t
A ll s a m p l e s w h ic h w e re s t i l l l a r g e en o u g h w e re
r e r u n on t h e c a l i b r a t e d NIR m a c h in e t o p r e d i c t c o n s t i t u e n t v a l u e s .
Also th e l o g v a lu e s from th e i n i t i a l NIR re a d in g s f o r a l l sam p les w ere
e n t e r e d i n t o a r e g r e s s i o n m odel a l o n g w i t h t h e c o e f f i c i e n t s and
33
i n t e r c e p t s from th e c a l i b r a t i o n p ro ced u re.
The o u tp u t was p re d ic te d
chem ical c o n s ti tu e n t v a lu e s f o r a l l o f th e o r ig i n a l sam p les, in c lu d in g
th o s e to o s m a ll to re ru n by MIR.
STATISTICAL ANALYSIS
BMDP (D ixon I 981) and MSUSTAT (Lund 1983) s t a t i s t i c a l p a c k a g e s
w e re u s e d t o a n a l y z e
th e d a ta .
r e f e r r e d t o i n c o n te x t.
The s p e c i f i c
p r o g r a m s u sed a r e
For th e re m a in d e r of t h i s t h e s i s s i g n i f i c a n c e
means t h a t p<,05.
For t h e s t a t i s t i c a l
s e r i e s of s u b p r o je c ts .
f a c to r /tre a tm e n t
a n a l y s i s t h i s p r o j e c t w as b r o k e n i n t o a
T his was n e c e s sa ry f o r two re a s o n s :
c o m b in a tio n s
I) No two
h a d e q u a l s a m p le s i z e .
2) A ll
t r e a t m e n t s e x c e p t th o se a s s o c i a t e d w i t h l a b r e s u l t s in c lu d e d unequal
n u m b e rs o f m i s s i n g d a t a .
U sin g s u b p r o j e c t s a l l o w e d t h e u se o f
s t a t i s t i c a l packages f o r th e a n a l y s i s .
CHEMICAL CONSTITUENT DATA
For o b j e c t i v e o n e,
t r e a t m e n t s w as u s e d .
a random d e s i g n w i t h one f a c t o r and n in e
The f a c t o r w as s p e c i e s w i t h
th re e le v e ls.
T rea tm e n ts w ere th e phenolo g ic a l s t a g e s when g r a z in g had o c c u rre d i n
the p re v io u s summer, i n c l u d i n g no g r a z i n g .
w ere th e n in e c h e m ic a l c o n s t i t u e n t s .
n u m b e rs.
The d e p e n d e n t v a r i a b l e s
T h e re w e re u n e q u a l t r e a t m e n t
The c h e m i c a l c o n s t i t u e n t d a t a w as a n a l y z e d i n f o u r s t e p s .
The chem ical c o n s t i t u e n t d a ta f o r each s p e c ie s was d e s c r ib e d u s in g t h e
" C l u s t e r A n a l y s i s o f V a r i a b l e s " p ro g r a m (D ixon 1981).
"S te p w ise
D is c r im in a n t A n a ly sis" (Dixon 1981) was th e n perform ed on t h e chem ical
c o n s t i t u e n t r e s u l t s to d e te rm in e i f
th e p h e n o lo g ic a l
s t a g e when
34
g r a z in g had o c c u rre d was r e f l e c t e d i n w i n t e r n u t r i t i o n a l v a lu e s.
The
c a l c u l a t i o n s w i t h i n t h i s program w ere such t h a t f o r a v a r i a b l e to be
removed a s a d i s c r i m i n a t i n g v a r i a b l e r e q u i r e d an F -v a lu e o f 4.0 which
w as a b o u t e q u a l t o p=.05.
The v a l u e s f o r e ach c o n s t i t u e n t w e re
compared among s p e c i e s u s in g " M u l t i - f a c t o r A n aly sis o f V ariance" (Lund
1983). For c o n s t i t u e n t s w ith s i g n i f i c a n t F - v a l u e s t h e s p e c i e s means
were s e p a r a te d u s in g t h e L e a st S i g n i f i c a n t D iffe re n c e tech n iq u e.
TRANSECT DATA
The e x p e rim e n ta l d e sig n f o r th e second o b j e c t i v e was randomized
b lo ck h av in g one f a c t o r w ith th r e e l e v e l s and two t r e a t m e n t s i n f i v e
b lo c k s. S i t e s ,
s p e c i e s g r a z i n g and s t a n d i n g d e a d w e r e t h e b l o c k s ,
f a c t o r and two tr e a t m e n t s , r e s p e c t i v e l y . The dependent v a r i a b l e s w ere
p l a n t h e i g h t and p h e n o l o g i c a l s t a g e on s i x s a m p l i n g d a t e s .
In itia l
a n a l y s i s o f p l a n t h e ig h t u s in g a n a l y s i s o f v a ria n c e , program 2V (Dixon
1 9 8 1 ),
fo u n d
th a t
on e a c h
sig n ific a n tly d iffe re n t.
d a te
site s
and s p e c i e s
w e re a lw ay s
Each s i t e / s p e c i e s case was a l s o analyzed on
e a c h d a t e u s i n g a n a l y s i s o f v a r i a n c e , p ro g ra m 2 V (D ix o n 1 981).
T h is
e l i m i n a t e d much o f t h e m i s s i n g d a t a by c a u s i n g th e p ro g ra m t o n o t
r e c o g n iz e i t .
Grazing and s ta n d i n g dead w ere th e tr e a t m e n t s .
The p h e n o lo g ic a l s ta g e d a ta was d e term in e d to be u n s u i t a b l e f o r
s t a t i s t i c a l a n a l y s i s because two o f th e in d e x in g keys pred o m in ated and
t h e d i s t r i b u t i o n o f key v a l u e s d i d n o t a p p e a r t o be n o rm a l.
The
freq u e n cy of p h e n o lo g ic a l s ta g e s f o r each s i t e / s p e c i e s case and each
tr e a t m e n t w ere graphed (Appendix B.)
RESULTS AND DISCUSSION
CHEMICAL CONSTITUENTS
The NIR p r e d i c t e d
v a lu e s
of
p r e s e n t e d i n t a b l e s 9 th r o u g h 11.
th e
chem ical
c o n stitu e n ts are
A ll v a l u e s i n t h i s p r o j e c t w e re
p e r c e n t o f t o t a l s a m p le w e i g h t e x p r e s s e d on a d ry m a t t e r b a s i s .
l a r g e p o r t i o n o f w h a t f o l l o w s w i l l be b a s e d on t h e s e a n a l y s e s .
A
The
chem ical c o n s t i t u e n t v a lu e s w ere s u i t a b l e f o r w i t h i n - p r o j e c t a n a l y s i s
but must be used more c a u ti o u s ly i n com parisons w ith o th e r s tu d ie s .
The u se o f NIR f o r p r e d i c t i n g c h e m i c a l c o n s t i t u e n t s and f o r a g e
q u a l i t y p a r a m e t e r s i s i n c r e a s i n g l y a c c e p t e d , so t h e r e i s a g r o w in g
body o f l i t e r a t u r e
G eneral
c o n clu sio n s
(e .g . B rooks e t a l .
su p p o rte d
by
th is
1984 and P a r k e t a l . 1 983).
lite ra tu re
in c lu d e d
th e
f o l l o w i n g : NIR p r e d i c t i o n s o f c o n s t i t u e n t s t h a t could be a s s o c i a t e d
w i t h some l i m i t e d num ber o f s p e c i f i c c h e m i c a l bonds ( i . e . PRO) w e re
more a c c u r a te th a n o f c o n s t i t u e n t s which in c lu d e d b e h a v io r a l f a c t o r s
p l u s a w id e r a n g e o f c h e m i c a l bonds ( i . e .
IVOMD).
v a lu e s ten d ed t o be h ig h e r th a n w et c h e m is try v a lu es.
sample s i z e th e more a c c u r a t e was the p r e d ic tio n .
NIR p r e d i c t e d
The l a r g e r th e
I f the e n t i r e range
o f c o n s t i t u e n t v a lu e s was in c lu d e d i n t h e c a l i b r a t i o n ,
sam ple r e s u l t s
w e r e m ore a c c u r a t e t h a n i f o u t l y i n g c o n s t i t u e n t v a l u e s had t o be
p re d ic te d .
Ash h a s b e en p o o r l y p r e d i c t e d by t h e I n f r A l y s e r 4 0 0 ,
because i t does n o t re c o g n iz e phosphorus.
All of th e s e problem s and
c o n s t r a i n t s more or l e s s a f f e c t e d t h e NIR p r e d ic te d c o n s t i t u e n t v a lu e s
i n t h is p ro je c t.
36
T able 9 .
A ll NIR p re d ic te d w in te r chem ical c o n s titu e n t v a lu e s f o r
bluebunch w h e a tg ra ss and th e p h e n o lo g ic a l s ta g e when
g ra z in g had o c c u rre d d u rin g th e p re v io u s summer.
CHEMICAL CONSTITUENTS*1
NDF
STAGE2
$
I
I
I
2
2
3
3
3
4
6
6
7
7
7
7
8
9
9
81.43
80.5
76.2
79 .7
7 7 .8
79.2
89.0
82.0
7 4 .8
87.8
75.6
80.1
85.4
78.3
82.2
82.0
80.2
84.5
CELL
HC
PRO
DM
ASH
IVOMD
%
%
%
%
%
%
%
%
51.7
48.6
50.6
51.7
5 4.8
54.4
6 3.3
54.3
55.1
54.9
52.6
57.4
52.8
52.8
52.5
54.8
51.9
56.9
4 .8
3.7
4 .4
4 .3
6 .0
4 .9
6 .9
6.1
5 .8
7 .0
4 .6
5 .5
5.6
5 .9
5.1
6 .4
4 .8
6 .2
39.3
38.1
42.0
41.8
45.3
42.7
49.3
43.2
42.1
45.5
39.6
45.1
44.7
43.1
41.9
45.6
43.5
46.7
27.9
27.5
26.7
25.7
2 4 .8
24.6
26.8
28.3
24.9
32.9
24.4
24.0
31.5
28.4
31.8
29.5
26.9
30.3
8:5
19.0
31.6
58.1
69.5
41.2
59.5
105.5
27.9
19.1
43.8
16.0
69.0
70.7
38.4
59.3
50.5
67.5
75.7
ADF
ADL
5 .9 96.8
5.1 96.3
4 .4 97.2
4.1 95.9
3 .0 98.3
4.9 95.8
2 .3 95.3
3 .6 98.5
4 .3 97.1
1.9 ■ 99.9
6.1 96.2
4.4 95.4
1.7 98.0
3.5 98.7
4.1 97.2
2.5 98.9
2 .4 97.0
3 .0 97.2
8.9
7 .5
8.0
6 .0
8.0
7 .0
7 .2
7 .4
6.7
8 .3
7 .8
7 .3
7 .0
7 .9
6.5
7 .2
7 .4
1The chemical c o n s t i t u e n t s a r e n e u t r a l d e te r g e n t f i b e r (NDF)t acid
1 d e t e r g e n t f i b e r (ADF)t a c id d e t e r g e n t l i g n i n (ADL)„ c e l l u l o s e (CELL),
hemic e l l u l o s e (HG), p r o t e i n (PRO), dry m a tte r (DM), ash (ASH) and
i n v i t r o o rg a n ic m a tte r d is a p p e a ra n c e ( IVOMD).
^The ph en o lo g i c a l s ta g e s when g r a z in g o c c u rre d were v e g e t a t i v e ( I ) ,
boot ( 2 ) , seedheads emerging ( 3 ) , a n t h e s i s ( 4 ) , seed s h a t t e r ( 6 ) ,
m ature f o l i a g e (8) and ungrazed ( 9 ) .
^V alues a r e % of t o t a l sample w e ig h t on a dry m a tte r b a s i s .
37
Table 10.
A ll NIR p re d ic te d w in te r chem ical c o n s titu e n t v a lu e s f o r
rough fe s c u e and th e p h e n o lo g ic a l s ta g e when g ra z in g had
o c c u rre d d u rin g th e p re v io u s summer.
CHEMICAL CONSTITUENTS1
STAGE2
I
I
I
2
2
2
3
3
3
3
3
3
4
5
6
6
6
7
7
7
7
7
7
7
8
.. 8
9
NDF
ADF
ADL
CELL
HC
%
%
%
%
%
79.43
71.9
74.1
84.0
80.6
72.8
79.5
82.2
7 9.4
76.4
82.3
80.6
76.3
79.1
7 7 .8
79.1
7 9.4
78.4
7 9 .4
82.9
7 6 .8
79.9
80.2
77.6
78.4
85.7
77.5
49 .4
46.2
4 8 .2
49.4
47.5
43.6
49.1
48.9
50.4
50.3
51.3
47.8
49 .0
50.6
50.8
48.3
50.0
46.3
51.3
49.6
50.5
52.4
4 9 .2
47.9
4 8 .8
49.9
48.7
1.6
3 .0
3 .3
4 .8
2 .8
3.2
3 .2
2 .7
2 .9
4 .2
4 .2
3 .8
3 .2
3.4
3 .3
3 .9
3.1
3 .2
4.1
3 .8
4 .2
3 .9
3 .4
2 .6
3 .7
4.1
2 .7
41.7
34.5
37.5
49.0
44 .5
35.8
41.5
45.8
4 3.9
39.3
46.3
46.6
41.7
43.5
41.6
41.5
41.7
38.9
42.7
47.5
39.7
40.5
46.4
40.9
40.1
50.2
41.6
PRO
%
38.3 3 .2
18.6 6 .5
24.1 6 .6
23.5 3 .0
34.1 2 .4
19.0 6 .6
31.6 4 .8
30.5 3.1
31.0 3 .4
21.6 5.9
26.7 4.1
19.3 2.1
24.0 5 .2
29.5 3 .3
26.0 3 .9
20.5 . 6 .7
29.4 4 .3
30.2 5.1
29.7 4 .9
28.2 3 .3
22.3 5 .9
29.2 6.1
23.2 . 3 . 8
28.3 4 .8
30.1 5 .5
25.0 2.5
26.8 5 .0
DM
%
96.6
95.9
95.3
97.5
96.5
95.7
95.9
96.5
96.5
96.2
96.5
98.5
96.3
96.7
97.0
95.9
96.2
95.8
95.7
96.9
96.2
95.8
96.8
95.9
95.5
98.1
95.9
ASH
% ■
6 .7
8.1
10.4
6 .9
4 .3
7 .3
7 .3
5 .3
7 .4
11.6
7 .5
2 .7
9.1
7 .0
11.2
10.7
8 .0
8.5
8 .0
7 .0
11.3
9 .8
8 .4
8.4
8 .7
6.1
8 .2
IVOMD
%
60.6
44.8
47.1
59.6
6 0 .8
45.2
56.0
59.0
58.1
52.7
55 .0
60.8
4 9.9
58.3
57 .2
48.6
53 .8
55.8
52.5
59.2
52.5
50.5
53.5
5 2.7
52.6
62.6
50 .8
^The chemical c o n s t i t u e n t s a re n e u t r a l d e t e r g e n t f i b e r (NDF), a c id
d e t e r g e n t f i b e r (ADF), a c id d e te r g e n t l i g n i n (ADL), c e l l u l o s e (CELL),
h e m ic e llu lo s e (HG), p r o t e i n (PRO), dry m a t t e r (DM), a sh (ASH) and
i n v i t r o o rg a n ic m a tte r d is a p p e a ra n c e (IVOMD).
^The p h e n o lo g ic a l s ta g e s when g ra z in g o c c u rre d were v e g e t a t i v e ( I ) ,
boot ( 2 ) , seedheads emerging ( 3 ) , a n t h e s i s ( 4 ) , m ature f lo w e r s ( 5 ) ,
seed s h a t t e r ( 6 ) , m ature f o l i a g e ( 7 ) , f a l l reg ro w th (8 ) and ungrazed ( 9 ) .
^Values a r e % o f t o t a l sample w eig h t on a dry m a tte r b a s i s .
38
T able 11,
A ll NIR p r e d ic te d w in te r chem ical c o n s titu e n t v a lu e s f o r
Idaho fe s c u e and th e p h e n o lo g ic a l s ta g e when g ra z in g
o c c u rre d d u rin g th e p re v io u s summer.
CHEMICAL CONSTITUENTS1
NDF
STAGE2
I
I
I
I
I
I
2
2
2
2
2
3
3
3
3
3
6
6
6
7
7
7
7
8
8
9
9
%
7 1 . 33
86.7
82.1
82.4
71.7
72.9
77.0
71.8
80.8
75.8
7 4 .9
74.4
75.6
74.1
68.1
67.0
72.5
61.0
65 .4
67.9
81.1
69.4
74.6
7 1.7
73.0
76.4
77.1
ADF
ADL
%
%
46.2
49.9
4 5.2
47.3
46 .2
46.3
50.2
46.0
47.9
48.1
45.2
42.3
42 .8
52.8
46.6
45.3
42.0
39.2
33.9
41.6
52.9
43.4
4 8 .8
40.5
45 .8
44.3
48.0
4 .7
3 .0
3 .7
3 .6
3 .8
3 .5
3 .2
4 .2
2 .8
3.5
4 .2
3.6
3 .8
3 .7
3 .7
3 .5
4 .5
6.1
6.6
4.3
5.6
4 .2
4 .3
3 .8
3 .7
4.6
4 .2
CELL . HC
%
35.9
40.9
40.2
39.8
37.2
36.3
40.0
37.5
41.8
39.3
31 .8
35.1
35.7
41.9
36.3
36.0
37.6
25.7
26.6
35.1
35.2
35.6
37.0
33.5
36.2
36.2
36.7 .
%
25.3
27.6
31.7
31.7
25 .9
24.3
28.7
28.6
29.5
27.4
25.7
30.8
31.5
27.8
26.7
25.6
26.4
26.6
27.7
29.1
29.0
28.4
26.5
29.3
27.9
30.8
25.0
PRO
DM
ASH
IVOMD
%
%
%
%
93.1
90.3
91.1
90.9
94.1
93.7
95.6
96.7
95.6
95.9
97.0
96.8
96.9
96.4
96.2
97.2
96.6
99.6
97.3
96.8
94.4
97.3
96.0
95.5
96.7
96.2
95.6
15.4
10.6
10.7
12.3
5 .3
6.8
5 .4
3.9
7 .2
7 .4
8 .8
8.9
10.8
6.5
7.6
7 .3
9 .4
5.8
8.5
8,0
9.5
5.1
7 .8
8.1
7 .9
9.6
7.6
6 .6
5.1
6 .0
6 .2
6.1
6.1
4 .9
5.8
4 .9
5 .2
6 .3
6.1
6.1
5 .7
5.5
5.6
5 .9
6 .2
6.6
6 .5
9.4
5 .2
5 .0
5.9
6 .0
5 .5
4 .9
70.1
53.6
5 5 .2
55.4
52.7
51 .3
43.3
50.9
42.5
44.1
64.0
57.5
5 9.7
42.3
53.8
52.8
56.0
80.2
81.4
60 .8
44.5
59.0
61.1
60.1
53.8
64.0
55 .8
1The chemical c o n s t i t u e n t s a re n e u t r a l d e t e r g e n t f i b e r (NDF), a c id
d e te r g e n t f i b e r (ADF), a c id d e te r g e n t l i g n i n (ADL), c e l l u l o s e (CELL),
h e m ic e llu lo s e (HG), p r o t e i n (PRO), dry m a t t e r (DM), a s h (ASH) and
i n v i t r o o rg a n ic m a tte r d is a p p e a ra n c e (IVOMD),
2 The p h e n o lo g ic a l s ta g e s when g r a z in g o c c u rre d were v e g e t a t i v e ( I ) ,
boot ( 2 ) , seedheads emerging ( 3 ) , seed s h a t t e r ( 6 ) , m ature f o l i a g e ( 7 ) ,
f a l l re g ro w th (8 ) and ungrazed ( 9 ) .
^V alues a r e $ o f t o t a l sample w eig h t on a . d r y m a tte r b a s i s .
39
Bluebunch w h e a tg r a s s IVOMD r e s u l t s w ere v ery poor and ranged from
11% to 105%,
two u n lik e ly ex trem es.
The blue bunch w h e a tg r a s s IVOMD
mean w as n o t s i g n i f i c a n t l y d i f f e r e n t fro m t h a t o f e i t h e r f e s c u e .
H ow ever,
IVOMD1s f o r a l l
s p e c i e s w e r e h i g h e r (50% - 60%)
ex p ec te d (20% - 40%). Due to th e s e f a c t o r s ,
than
IVOMD w i l l no t be f u r t h e r
d is c u s s e d .
The ASH c o n s t i t u e n t was r e t a i n e d
because th e v a lu e s corresponded
w e l l w i t h th e c a l i b r a t i o n s a m p l e s .
T h i s may have b e en due t o low
phosphorus c o n te n t i n th e g r a s s e s ,
a t r a d i t i o n a l w e s te r n w i n t e r range
d efic ien c y .
CHEMICAL CONSTITUENT RELATIONS
C lu s t e r a n a l y s i s by s p e c ie s s e g r e g a te d w i n t e r c o n s t i t u e n t s i n t o
f i b e r and p r o t e i n c l u s t e r s .
Bluebunch w h e a tg ra s s c o n s t i t u e n t s ( ta b le
12) fo r m e d tw o l a r g e c l u s t e r s .
A c e l l u l o s e c l u s t e r o f ADF, CELL and
ADL and a h e m i c e l l u l o s e c l u s t e r w i t h NDF, HG and DM f o r m e d and com­
b in e d i n t o a f i b e r c l u s t e r .
a lso id e n tif ie d .
c o rre la ted .
A p r o t e i n c l u s t e r w i t h PRO and ASH w as
The f i b e r and p r o t e i n c l u s t e r s w e re n e g a t i v e l y
W inter bluebunch w h e a tg r a s s r e p r e s e n te d a " t y p i c a l " d o r­
m a n t w i n t e r g r a s s w i t h low PRO and h ig h ADL and ADF c o n c e n t r a t i o n s ,
and t h e c o n s t i t u e n t s c l u s t e r e d a s e x p e c t e d f o r s e v e r a l r e a s o n s : I )
C aldw ell e t a l . (1981) showed t h a t d u rin g th e grow ing season bluebunch
w h e a tg r a s s d id n o t respond t o d e f o l i a t i o n , and t h e r e was no i n d i c a t i o n
t h a t t h i s s i t u a t i o n c h an g e d d u r i n g t h e w i n t e r .
2) B lu e b u n c h w h e a t -
g r a s s a p p e a r e d , d o rm a n t i n A u g u st, e a r l i e r th a n e i t h e r f e s c u e . T h is
im p lie d t h a t once bluebunch w h e a tg ra s s began grow th i n the s p r in g , i t s
40
Table 12.
C lu s te r a n a ly s is o f w in te r bluebunch w h e a tg ra ss chem ical
c o n s ti tu e n ts .
-.4 0
30
-.20
.00
S 50
I
M
I
L
A 60
R
I
T
Y
70
.20
.40
.60
.80
100
1.00
,
NDF HC DM ADF CELL
ADL
IVOMD PRO
CHEMICAL CONSTITUENTS
ASH
C
0
R
R
E
L
A
T
I
0
N
41
g ro w th ,
d e v e lo p m e n t and s e n e s c e n c e w e r e m a i n l y f i x e d and l i t t l e
in f lu e n c e d by e x t e r n a l f a c t o r s such as g ra z in g . A fte r the c e s s a t i o n of
a c t i v e g r o w th d i f f e r e n t i a t i o n o f t h e c e l l u l o s e , h e m i c e l l u l o s e and
p r o t e i n c l u s t e r s co n tin u e d due t o le a c h i n g and r e s p i r a t i o n . . By March,
when th e p l a n t s w ere c o l l e c t e d ,
t h e r e w ere i d e n t i f i a b l e d i f f e r e n c e s
betw een th e th r e e c l u s t e r s .
For rough fe s c u e ( t a b l e 13) t h e r e was a lo o s e f i b e r c l u s t e r which
co m b in e d a NDF and CELL c l u s t e r w i t h DM, HG, ADF and ADL c l u s t e r s .
PRO and ASH form ed t h e p r o t e i n c l u s t e r .
fib e r
c lu ste r
d ig e stib ility
c o n ta in in g
ADF,
In Idaho f e s c u e ( t a b l e 14) a
CELL,
NDF,
HC a n d
ASH a n d
a
c l u s t e r c o n t a i n i n g ADL, PRO and DM w e r e i d e n t i f i e d .
SUMMER GRAZING EFFECTS
No rough fe s c u e or bluebunch w h e a tg r a s s w i n t e r chem ical c o n s t i ­
t u e n t v a l u e s c o u ld p r e d i c t any o f t h e s t a g e s d u r i n g w h ic h th e p l a n t
h ad b e e n g r a z e d i n t h e p r e v i o u s sum m er. A lso g r a z e d and u n g r a z e d
p la n ts
c o u ld
not
be d i s t i n g u i s h e d .
It
w as c o n c lu d e d
th a t
th e
ph en o lo g ic a l s ta g e when rough fe s c u e and bluebunch w h e a tg ra s s had been
g razed d id not s i g n i f i c a n t l y
a f f e c t w in te r c h em ical c o n s t i t u e n t
v a lu e s .
The f o l l o w i n g w as b a s e d on d i s c r i m i n a n t a n a l y s i s o f c h e m i c a l
c o n s t i t u e n t s fro m p o o le d w i n t e r and summer s a m p l e s .
d is c r im in a n t
In p a r t th e
a n a ly s is re in fo rc e d th e c lu s t e r a n a ly s is in th a t i t
tended to p ic k p rim ary c l u s t e r m em bers a s d i s c r i m i n a t i n g v a r i a b l e s .
For rough fe s c u e NDF was t h e d i s c r i m i n a t i n g v a r i a b l e ,
th e i n i t i a l
and i t was a l s o
c o n s ti tu e n t s e le c te d i n th e f i b e r c l u s t e r .
I n Id a h o
42
Table 13.
C lu s te r a n a ly s is o f w in te r rough fe sc u e chem ical
c o n s ti tu e n ts .
.20
.40
.60
.80
J .00
CHEMICAL CONSTITUENTS
C
0
R
R
E
L
A
T
I
0
N
43
T able 14.
C lu s te r a n a ly s is o f w in te r Idaho fe sc u e chem ical
c o n s ti tu e n ts .
60
.20
.40
.60
.80
,
CHEMICAL CONSTITUENTS
1.00
C
0
R
R
E
L
A
T
I
0
N
44
f e s c u e ADL, PRO, NDF and DM w e re t h e d i s c r i m i n a t i n g v a r i a b l e s .
ADL
and PRO w e re t h e tw o e x t r e m e m em bers o f t h e d i g e s t i b i l i t y c l u s t e r .
NDF was i n t h e f i b e r c l u s t e r .
DM form ed an ind ep en d en t c l u s t e r .
For rough f e s c u e th e a n t h e s i s , m ature f lo w e r s and ungrazed phenol o g i c a l s t a g e s o f g r a z in g w ere s u c c e s s f u l l y p r e d ic te d by th e chem ical
c o n s t i t u e n t s . F o r I d a h o f e s c u e v e g e t a t i v e , b o o t and u n g r a z e d p l a n t s
w ere
th e
sta g e s
p re d ic te d .
The s e e d s h a t t e r
s ta g e w as a ls o
s u c c e s s f u l l y p r e d ic te d 66.7% of th e tim e.
D is c rim in a n t
c o n stitu e n ts
a n a l y s i s w as p e r f o r m e d on t h e p o o le d c h e m i c a l
of b o th f e s c u e s .
T here w e re i n d i v i d u a l
d i f f e r e n c e s i n th e ch em ical c o n s ti tu e n ts ,
a c r o s s s p e c ie s ,
sp e c ie s
so l i t h e s a m p l e s p o o le d
p re v io u s ly d i s c r i m i n a t i n g c o n s t i t u e n t s w ere averaged
i n t o in sig n ific a n ce .
For w i n t e r c o l l e c t i o n s o f Id a h o f e s c u e ( n o t t h e p o o le d s a m p le )
DM, NDF and ADL w ere th e d i s c r i m i n a t i n g v a r i a b l e s ; however, only the
p l a n t s g ra z ed d u rin g t h e v e g e t a t i v e s ta g e w ere 100% p r e d i c t a b l e by th e
w in te r
c o n stitu e n ts.
p re d ic te d
c o rre c tly
The s e e d s h a t t e r and u n g r a z e d s t a g e s w e re
66.7%
and 50.0% o f
th e
tim e ,
re sp e c tiv e ly .
D is c r i m i n a n t a n a l y s i s i n d i c a t e d t h a t t h e p h e n o l o g i c a l s t a g e d u r i n g v
which g r a z in g o c c u rre d and g r a z in g i t s e l f did no t have a s i g n i f i c a n t ^
e f f e c t on r o u g h f e s c u e o r b lu e b u n c h w h e a t g r a s s w i n t e r n u t r i t i o n a l
v a lu e s .
By March any d i f f e r e n c e s i n chem ical c o n s t i t u e n t s t h a t may
have r e s u l t e d
fro m
th e
p h e n o lo g ic al
sta g e
at
g r a z i n g had
been
o b l i t e r a t e d by more i n f l u e n t i a l p r o c e s s e s , such a s t r a n s l o c a t i o n of
n u t r i e n t s a n d /o r w e a th e rin g . For Idaho fe s c u e g r a z in g a t some s ta g e s
o f growth d id have an a f f e c t on su b sequent w i n t e r n u t r i t i o n a l v a lu es.
45
T h e re w a s, h o w e v e r, no p a t t e r n o f im p ro v e m e n t o r damage r e l a t e d t o
g r a z in g a t any p h o n o lo g ic a l s ta g e , i n c l u d i n g ungrazed,
That g r a s s e s have s p e c ie s s p e c i f i c re s p o n se s t o g r a z in g h as been
w e l l d o c u m e n te d (W h ite 1973, Cook 1966 and C a l d w e l l e t a l . 1981).
T h a t t h e s e d i f f e r e n c e s i n g r a z i n g r e s p o n s e a r e p a r t o f an a r r a y o f
i n d i c a t o r s of s p e c ie s s p e c i f i c p h y sio lo g y i s a to p ic of c u r r e n t
d i s c u s s i o n and r e s e a r c h (C ald w ell e t a l . 1981, R obertson and Woolhouse
1984a, b a n d McLean and Wikeem 19 8 5 a , b). I n t h i s p r o j e c t b lu e b u n c h
w h e a tg ra s s r e p r e s e n t e d one d i s t i n c t p h y s io lo g ic a l type and th e f e s c u e s
a n o th e r type*
C l i m a t e i s one i m p o r t a n t g r a s s p h y s i o l o g y s e l e c t i o n f a c t o r .
B lu e b u n c h f h e a t g r a s s , a G r e a t B a s in s p e c i e s , h a s a d a p t e d t o t h e c o ld
d e s e r t c l i m a t e (C aldw ell e t a l . 1981). Conversely th e F e s tu c h genus i s
b e l i e v e d t o be o f n o r t h e r n o r i g i n s .
Rough f e s c u e d i s t r i b u t i o n i s
c u r r e n t l y c e n te r e d i n th e n o r th e r n G re at P la in s and Aspen p a rk la n d s i n
th e n o r t h e r n U n i t e d S t a t e s and Canada.
Idaho fe s c u e i s lo c a t e d i n
m ountains and f o o t h i l l s from th e C e n tra l Rockies i n t o s o u th e rn Canada.
Rough and I d a h o f e s c u e a r e l o c a t e d i n g e n e r a l l y m o i s t e r and c o o l e r
h a b i t a t s th a n bluebunch w h e a tg ra ss.
Due t o
t h e i r g e o g ra p h y and e l e v a t i o n ,
o f G re a t B asin ,
th e E lk h o rn s c o n ta in
flo ristic
e lem en ts
N o rth e rn
P la in s
h a b ita ts.
. By l o c a t i n g t r a n s e c t s t o i n c l u d e a l l
and
boreal
t h r e e key g r a s s
s p e c ie s unique m i c r o h a b i t a t s were sampled which a llo w ed th e m ixing of
t h r e e s p e c ie s from d i f f e r e n t g e o g ra p h ic re g io n s .
Rough fe s c u e was on
t h e warm and d ry end of_l_t s p o t e n t i a l d i s t r i b u t i o n Djo,t_e.ntl-al—w_h i l e
b lu e b u n c h w h e a t g r a s s w as i n
th e
c o o l e r and w e t t e r
p a rt of i t s
i»6
p o te n tia l
range.
Idaho
fescue
w as
in
ty p ic a l
h a b ita t.
As a
consequence, a s any of th e sam p lin g s i t e s improve i n ran g e c o n d itio n ,
im p ly in g b e t t e r m o is tu re r e l a t i o n s ,
rough fe sc u e would be exp ected t o
h a v e a c o m p e t i t i v e a d v a n t a g e o v e r b lu e b u n c h w h e a t g r a s s .
T h is w as
observed i n p la c e s w ith conserved m o is tu r e , d e p re s s io n s , snow d r i f t s ,
shaded a r e a s and a p p a r e n tly ungrazed a re a s .
S tu d i e s o f h e rb iv o ry re sp o n se s have su g g ested t h a t none of th e se
th r e e g r a s s e s a r e v e ry g r a z in g t o l e r a n t ,
but t h a t Idaho fe s c u e te n d s
to be th e more g r a z in g t o l e r a n t and rough fe s c u e th e l e a s t (McLean and
T is d a le 1966, Mueggler 1975
and McLean and Wikeem 1985a , b).
R e s u l t s fro m t h i s p r o j e c t s u p p o r t e d t h e concept t h a t bluebunch
w h e a tg r a s s and th e f e s c u e s r e p r e s e n t e d two d i f f e r e n t p h y s i o l o g i c a l
ty p es. The independence o f fe sc u e chem ical c o n s t i t u e n t v a lu e s was due
i n p a r t t o a l l p l a n t s o f b o th s p e c i e s h a v i n g g r e e n o v e r w i n t e r i n g
l e a v e s s u g g e s t i n g some l e v e l
of w in te r
p h o to sy n th e sis.
W in te r
p h o t o s y n t h e s i s by g r a m i n o i d s i n h a r s h w i n t e r e n v i r o n m e n t s h a s b e en
re p o rte d
( R o b e r t s a n d W oolhouse 1 9 8 4 a , b and C h a p in e t a l .
1980).
W inter p h o to s y n th e tic a c t i v i t y r e p r e s e n te d a s i g n i f i c a n t p h y s io lo g ic a l
d i f f e r e n c e betw een t h e f e s c u e s and bluebunch w h e a tg ra s s.
b e tw e e n rp u g h and I d a h o f e s c u e r e p r e s e n t e d
D iff e re n c e s
d iff e r e n c e s in w in te r
p h y s io lo g ic a l a c t i v i t y r a t h e r th a n i t s p re se n c e or a b se n c e.
The
g e n e r a l e x p la n a tio n f o r in c r e a s e d s t r u c t u r a l c a rb o h y d ra te s d u rin g th e
w in te r,
e sp e c ia lly
c e llu lo se
and l i g n i n ,
is
th a t
m ore
e a sily
m e ta b o liz e d c a r b o h y d ra te s a re used i n m aintenance l e v e l r e s p i r a t i o n .
W in ter
p h o to sy n th e tic
a c tiv ity
p ro v id e d
some h i g h l y
a v a ila b le
47
c a r b o h y d r a t e s , r e d u c i n g t i s s u e c a t a b o l i s m and p r e v e n tin g in c r e a s e d
c o n c e n tr a tio n s o f c e l l u l o s e and l i g n i n i n th e w i n t e r f i b e r component.
Idaho fe s c u e had h ig h e r w i n t e r PRO l e v e l s th a n t h e o th e r s p e c ie s
w ith l i t t l e
d e c lin e in
PRO fro m
sum m er i n t o
w in te r
(ta b le
15).
S i m i l a r f i n d i n g s have been r e p o r t e d by o t h e r s (McCall 1939 and 1940).
Since p r o t e i n c o n te n t was measured i n p e rc e n t, no d i s t i n c t i o n was made
between e i t h e r more p r o t e i n , l e s s d i l u t i o n by s t r u c t u r a l c a rb o h y d ra te s
or
b o th .
In
any
case,
it
m ust i n d i c a t e
a le v e l
of w in te r
p h y s i o l o g i c a l a c t i v i t y p r e s e n t i n few o t h e r g r a s s e s . I n " t y p i c a l "
g rasses,
p ro te in
c o n c e n tra tio n s
d e c lin e
by f a l l
c o n s ta n t th ro u g h w i n t e r (McCall 1939 and 1940).
and t e n d t o
In Idaho fe s c u e ,
be
th e
l a t e grow ing season draw down of p r o t e i n was l e s s th a n f o r most o th e r
g ra s s e s .
This could be due to red u ced f a l l t r a n s l o c a t i o n o f p r o t e i n
i n t o th e cro w n o r r o o t s a s h a s b e e n r e p o r t e d f o r e v e r g r e e n s h r u b s
(C h a p in e t a l . 1980).
SPECIES
COMPOSITION
The s p e c ie s c o m p o sitio n on th e f i v e s i t e s s tu d ie d a r e p re s e n te d
in ta b le
16. The J e n k i n ' s G u lch I I s i t e w as t h e m o s t d i s s i m i l a r ,
b e c a u s e one t r a n s e c t w as d e s t r o y e d by c a t t l e t r a m p l i n g e a r l y i n t h e
fa ll,
and t h e s i t e had snow c o v e r on m o st o f t h e s p r i n g s a m p l i n g
d a te s . The o th e r s i t e s w ere v ery s i m i l a r . One hundred f i f t y - f o u r rough
f e s c u e , 158 b l u e b u n c h w h e a t g r a s s and 575 I d a h o f e s c u e p l a n t s w e re
sampled.
48
Table 15.
Comparisons of NIR p r e d ic te d chem ical c o n s titu e n t mean
v a lu e s 2 f o r rough fe sc u e (P e s o ), Idaho fe s c u e (F e id ) and
bluebunch w h e a tg ra ss (A gsp).
SPECIES
CHEMICAL
CONSTITUENT
I
FESQ
W inter Summer
% ■
FEID
W inter Summer
%
%
%
NDF
81b
79be
67 d
74af
71 f
ADF
54 c
49be
45 d
46 a f
46 f
ADL
6c
3 ad
3 d
4bg
2 f
CELL
44 c
42be
34 d
36 a f
38 f
HC
28a
27 ae
23 d
28ag
21 f
PRO
4a
5bd
7 e
6ef
6 f
DM
97b
96 ad
96 d
96af
96 f
ASH
7a
Bad
11 e
8af
9 f
53 a
54 ae
43 d
57 ag
51 f
IVOMD
I
AQSp
W inter
.
Means w i t h i n t h e same row fo llo w ed by th e same l e t t e r a r e not
s i g n i f i c a n t l y d i f f e r e n t (p^.,05). Three comparisons w ere made f o r
each chem ical c o n s t i t u e n t : I) Agsp, Fesc and Feid w i n t e r means, 2)
w in te r and summer Fesc means and 3) w in te r and summer Feid means.
2Values a r e % o f t o t a l sample w eig h t on a dry m a tte r b a s i s .
49
Table 16.
Number o f rough fe sc u e ( F e s c ) l Idaho fe s c u e (F e ld ) and
bluebunch w h e atg rass (Agsp) on each sam pling s i t e in
th e s p rin g , 1984.
SPECIES
Feid
Fesc
SJTE
n
n
Agsp
n
T o tal
n
South Crow I
36 (15.32)1
South Crow I I
28 (1 5 .0 2 )
118 (6 3 .1 2 )
41 (2 1 .9 2 )
I87d
J e n k i n ' s Gulch I
33 (14.22)
168 (7 2 .4 2 )
31 (1 3 .4 2 )
23 2d
J e n k i n ' s Gulch I I
29 (4 1 .4 2 )
29 (41 .42)
12 (1 7 .1 2 )
70c
Muddy Lake
28 (1 7 .3 2 )
110 (6 7 .9 2 )
24 (1 4 .8 2 )
I62d
154 (1 7 .4 2 )
30.,8a1
575 (6 4 .8 2 )
115.0b 1
158 (1 7 .8 2 )
3 1 .6 a
887
T o ta l
Average
150 (63.62)
50 (2 1 .2 2 )
^ of key s p e c i e s t o t a l on s i t e (o r row t o t a l s ) .
2Values fo llo w e d by th e same l e t t e r a r e not s i g n i f i c a n t l y
d i f f e r e n t . There were two com parisons: I ) among s p e c ie s
(column a v e r a g e s ) and 2) among s i t e s (row t o t a l s ) .
236d
50
WINTER GRAZING EFFECTS
Rough f e s c u e u t i l i z a t i o n (86%) was g r e a t e r t h a n e i t h e r Id a h o
f e s c u e (49%) o r b lu e b u n c h w h e a t g r a s s (47%) w h ic h w e re n o t d i f f e r e n t
( t a b l e 1 7 ).
The num ber o f p l a n t s g r a z e d p e r s p e c i e s a r e r e p o r t e d i n
t a b l e 17. On t h i s w i n t e r r a n g e e l k s e l e c t e d r o u g h f e s c u e > Id a h o
fe s c u e > bluebunch w h e a tg ra s s.
When h e ig h t d a ta w ere pooled a c r o s s s i t e s a n d /o r s p e c ie s t h e r e was
no d a te when w i n t e r g r a z in g had a s i g n i f i c a n t e f f e c t on s p r i n g p la n t
h e i g h t s . F o r a l l s p e c i e s t h e r e w e r e 52 t o t a l i n d i v i d u a l d a t e / s i t e
cases.
One o f 17 b lu e b u n c h w h e a t g r a s s and s i x o f 19 I d a h o f e s c u e
c a s e s had s i g n i f i c a n t l y
ta lle r
p l a n t s a s s o c i a t e d w i t h no w i n t e r
I
g ra z in g ,
No i n d i v i d u a l c a s e s f o r rough f e s c u e had s i g n i f i c a n t g ra z in g
e ffe c ts.
F i v e o f t h e i n d i v i d u a l c a s e s o f Id a h o f e s c u e w i t h s h o r t e r
l e a v e s w ere on th e l a s t th r e e sam p lin g d a te s .
STANDING
DEAD
EFFECTS
More bluebunch w h e a tg ra s s (63%) and Idaho fe s c u e (60%) p l a n t s had
s t a n d i n g d e ad r e m a i n i n g a t t h e end o f w i n t e r t h a n d i d r o u g h f e s c u e
(20%).
When h e i g h t d a t a w e re p o o le d a c r o s s
site
a n d /o r
sp e c ie s
s t a n d i n g d e a d w as a s s o c i a t e d w i t h t a l l e r p l a n t s on 5 / 1 0 , 6 / 1 2 , 6 /1 8
and 6/24. I n d iv i d u a l s i t e / d a t e c a se s p e r s p e c ie s a r e shown i n t a b l e s
18 t o
20.
On t h e
S o u th
Crow
II
s ite
a ll
th re e
sp e c ie s
s i g n i f i c a n t l y t a l l e r p l a n t s a s s o c i a t e d w ith s ta n d in g dead.
had
For rough
f e s c u e and bluebunch w h e a tg r a s s t h i s was th e only s i t e where s ta n d in g
d e a d a p p e a r e d i m p o r t a n t . T h e re w e re tw o a d d i t i o n a l c a s e s f o r e ach
s p e c ie s when p l a n t s w ith s ta n d in g dead w ere s i g n i f i c a n t l y t a l l e r . For
51
Table 17.
F re q u e n c ie s and number of p l a n t s grazed (G) and w ith s ta n d ­
in g dead p r e s e n t (SD) f o r rough fe s c u e ( F e s c ) 9 Idaho fe s c u e
(F e id ) and bluebunch w h e a tg ra s s (Agsp) p l a n t s on each s i t e .
sm SPECIES
FESC
SCI
G %2
n
SD
%
n
FEID
G %
n
SD
%
n
AGSP
G %
n
SD
%
n
Mean
G %
SD %
SCII JGI
J G II
ML
T o tal
81
29
89
25
91
30
69
20
100
28
132
36
13
25
7
30
10
10
3
0
0
33
62
93
43
51
44
74
28
8
69
76
302
65
98
80
94
63
106
66
19
26
29
346
58
29
51
21
45
14
67
8
13
3
75
72
36
73
30
68
21
50
6
54
13
106
67e
58 f
6 le
59 f
60e
54 f
55e
42 f
6 le
27 f
Mean
SG
SSD
86b3
20c
49a
6 Od
47a
63d
IS i t e s a r e South Crow I (SCI) and I I ( S C I I ) , J e n k i n ' s Gulch I
(JG I) and I I (JG II) and Muddy Lake (ML).
^V alues a r e % o f s p e c i e s ' t o t a l on s i t e .
^Means fo llo w e d by th e same l e t t e r a r e not s i g n i f i c a n t l y d i f f e r e n t
( p £ .0 5 ) . Four com parisons were made: I ) g r a z in g among s p e c ie s (Mean
%G colum n), 2 ) s ta n d i n g dead among s p e c ie s (Mean J&SD colum n), 3)
g r a z in g among s i t e s (Mean G % row) and 4) s ta n d in g dead among s i t e s
(Mean SD % ro w ).
Table 18.
Average rough fe sc u e h e i g h t s (cm) on s i x sam pling d a te s f o r p l a n t s
w ith o u t (no SD) and w ith (SD) s ta n d in g dead a t the b eginning of
new growth i n 1984.
srrE 1
SAMPLING
DATES
SCI
no SD2 SD
4 /1 8
10.43 11.1
(ns)4
5/10
11.0
5/19
SCII
SD
no SD
NA
NA5
NA
JGI
no SD
SD
NA
NA
JG II
no SD
SD
NA
ML
no SD
SD
NA v
5.5
6
5.0
NA
NA
NA
12.4
(ns)
NA
21.7
NA
NA
NA
NA
NA
NA
NA
NA
NA
21.1
22.5
(ns)
NA
NA
22.7
18.9
8 .2
7 .7
6 .5
(ns)
(ns)
(O)
6/12
27.3
27.0
(ns)
6/18
NA
NA
27.5
33.8
(BO)
2 2 .7
24.8
(ns)
23.4
30.5
(ns)
NA
6/24
2 9 .2
31.5
(ns)
31.4
37.7
25.2
25.5
(ns)
26.8
35.7
26.3
(B)
(B)
1S i t e s a r e South Crow I (SCI) and I I ( S C I I ) , J e n k i n 1s Gulch I (JGI) and
I I (JG II) and Muddy Lake (ML).
2 Absent = 10% o f o v e rw in te rin g t i l l e r s w ith r e s i d u a l le a v e s . P re se n t > 10% o f
t i l l e r s w ith r e s i d u a l le a v e s .
3Mean h e ig h t (cm) o f lo n g e s t le a v e s .
4 AOV of p la n t h e i g h t s comparing no SD t o SD, (n s ) = p>.05, ( Q) = R l.0 5 ,
(BO ) = P i . 0 1 .
5D a t e / s i t e c a se s w ith more th a n 50% m issin g d a ta were not a n a ly se d .
6No p l a n t s on the ML s i t e had s ta n d in g dead.
6
NA
6
Vl
ro
Table 19»
SAMPLING
DATE
Average bluebunch w h e atg rass h e i g h t s (cm) on s i x sam pling d a te s f o r
p l a n t s w ith o u t (no Sb) and w ith (Sb) s ta n d in g dead a t the beg in n in g o f new growth i n 1984 •
SCI
no SD2 SD
4/18
NA3
5/10
7 .8
SCII
no SD
SD
SITE1
JGI
no SD
SD
NA
NA
NA
NA
8 .2
NA
NA
5 .5
.
6/12
14.4
19.8
NA
NA
5 .9
6 .3
(ns)
(n s)
5/19
JG II
no SD
SD
1.4.7
(n s)
8 .9
2 1.7
(ns)
19.1
8.4
NA
21.3
(ns)
17.0
ML
no SD SD
NA
2 . 04
2.5
(ns)5
6 .4
3 .7
(n s)
4 .3
(ns)
NA
NA
NA
NA
NA
21.1
NA
NA
16.9
19.5
(ns)
23.4
(ns)
2 5 .2
25.4
(n s)
NA
NA
27.4
31.0
32.5
(ns)
23.4
23.4
(ns)
(n s)
(0)
6/18
NA
NA
19.5
26.4
(OO)
19;8
6/24
23.8
24.6
(ns)
23.7
27.9
(oo)
20.8
(O)
^ S i t e s a r e South Crow I (SCI) and I I (S C II), J e n k i n 1s Gulch I (JGI) and
I I (JG II) and Muddy Lake (ML).
2 Absent = 10% o f o v e rw in te rin g t i l l e r s w ith r e s i d u a l le a v e s .
P re s e n t >10% of
t i l l e r s w ith r e s i d u a l l e a v e s .
3D a t e / s i t e c a se s w ith more th a n 50% m issin g d a ta were not a n a ly s e d .
4 Mean h e ig h t (cm) o f l o n g e s t le a v e s .
5 AOV of p la n t h e i g h t s comparing no SD to Sb, (n s ) = p>.05, ( e ) = P.1.05,
(OO) = pi* 01.
g
r0W
O
g H
SAMPLING
DATE
Average Idaho fe sc u e h e i g h t s (cm) on s i x sam pling d a te s f o r p l a n t s
w ith o u t (no SD) and w ith (SD) s ta n d in g dead a t the beg in n in g of
new growth i n 1984.
B
Table 2 0 .
4/18
NA3
NA
5/10
6.9* 8 .5
(°op
SCII
no SD
SD
NA
NA
5 .7
9 .2
(oo)
SITE1
JGI
no SD
SD
NA
NA
5 .0
6.3
(oo)
JG II
no SD
SD
NA
NA
4 .5
5.1
no SD
ML
SD
NA
NA
3 .8
4 .3
(O)
(ns)
5/19
10.1
12.9
(OO)
NA
NA
NA
NA
NA
NA
NA
NA
6/12
11.0
15.8
(oo)
10.1
14.5
(oo)
9.4
12.2
(oo)
NA
NA
10.4
10.9
(ns)
6/18
NA
NA
9 .7
14.8
(oo)
9 .7
12.7
(oo)
10.0
11.6
(ns)
NA
NA
6/24
11.4
15.8
(oo)
11.0
16.8
(oo)
10.2
13.2
(oo)
11.4
12.7
(ns)
11.0
12.0
(ns)
^ S i t e s a r e South Crow I: (SCI) and I I (S C II), J e n k i n ' s Gulch I (JGI) and
I I (JG II) and Muddy Lake (ML).
^Absent = 10% o f o v e rw in te rin g t i l l e r s w ith r e s i d u a l le a v e s . P rese n t >10% of
t i l l e r s w ith r e s i d u a l le a v e s .
^ D a t e / s i t e c a se s w ith more th a n 50% m issin g d a t a were not an aly se d .
llMean h e ig h t (cm) o f l o n g e s t le a v e s .
^AOV o f p la n t h e ig h ts comparing no SD t o SD, (n s ) = p>.05, ( ° ) = PJ&.05,
(oo) = p<.01.
55
3 o f 4 ro u g h f e s c u e c a s e s and 3 o f 4 b lu e b u n c h w h e a tg r a s s t h e c a s e s
w ith t a l l e r p la n ts w ere on th e l a s t two sam p lin g d a te s .
F or
Id a h o
fe scu e
in
13 o f 18 i n d i v i d u a l
s ta n d in g dead had s i g n i f i c a n t l y lo n g e r le a v e s .
c a s e s p l a n t s w ith
The J e n k in ’s Gulch I I
s i t e was th e only one w ith o u t a s ta n d in g dead a f f e c t .
WINTER LEAF LOSS
B lu eb u n ch w h e a tg r a s s i s c o n s id e r e d one o f th e m ore d e s i r a b l e
f o r a g e p l a n t s on w e s t e r n r a n g e la n d s .
w as d i f f e r e n t .
I n th e E lk h o rn s th e s i t u a t i o n
I n b o th summer (D eSim one e t a l . 1 984) and w i n t e r ,
bluebunch w h e a tg ra s s u t i l i z a t i o n w as l e s s th a n th a t of. e i t h e r fe sc u e .
In a l l
s e a so n s,
bluebunch w h e a tg ra s s had e lo n g a te d v e g e ta tiv e culm s.
N e ith e r fe s c u e e x h ib ite d t h i s re sp o n se .
D uring th e w in te r bluebunch
w h e a tg r a s s h ad t h e h i g h e s t v a l u e s f o r f i b e r c o n s t i t u e n t s and d ry
m a t t e r (ADF, ADL and DM w e re s i g n i f i c a n t maximum v a l u e s among th e
th r e e s p e c ie s ) and th e lo w e s t p r o te in v a lu e s . I t was a p p a re n tly l e s s
p a la ta b le .
The c o m b in a tio n o f c o a r s e
e re c t
c u lm s and c h e m ic a l
c o n s t i t u e n t s a s s o c i a t e d w ith lo w e r p a l a t a b i l i t y combined to red u ce
g ra z in g p re s s u re on bluebunch w h e a tg ra s s through th e e n t i r e y e a r.
S a u e r’s
(1 9 7 8 )
h y p o th e s iz e d
m e c h a n is m s f o r
s t a n d i n g d e ad
i n f l u e n c e in c l u d e d im p ro v e d m i d r o s i t e m o i s t u r e c o n d i t i o n s .
The
r e s u l t s from t h i s stu d y su p p o rte d such an h y p o th e sis.
The s ta n d in g dead and g ra z in g p a ra m e te rs appeared to be p o s itiv e
and n e g a t i v e i n d i c a t o r s o f l i t t e r ,
r e s p e c t i v e l y . The p r im a r y s i t e
im p a c t o f l i t t e r i s s h a d in g , c o o l i n g th e s o i l s u r f a c e an d r e d u c in g
e v a p o ra tio n .
On th e f i r s t
sam p lin g d a te th e r e w ere in d iv id u a l c a se s
56
o f p la n ts t h a t had been g razed and had no s ta n d in g dead h av in g lo n g e r
l e a v e s , b u t th e d i f f e r e n c e s w e re n o t s i g n i f i c a n t (NS). P l a n t s w ith
l e s s r e s id u a l m a te r ia l began grow th e a r l i e r ,
presum ably because th ey
c o u ld warm up s o o n e r . On a l l o t h e r d a t e s f o r a l l c a s e s , p l a n t s w i t h
s t a n d i n g d e a d a n d / o r no g r a z i n g w e re t a l l e r ,
s ig n ific a n tly .
th o u g h n o t a lw a y s
The p re sen c e o f s ta n d in g dead was more im p o rta n t than
th e method o f i t s rem oval.
The amount o f l i t t e r i n one y e a r 's s ta n d in g dead was n o t ad eq u ate
t o in d e p e n d e n tly i n f l u e n c e
w h e a tg ra s s,
by l i t t e r .
g ro w th o f ro u g h f e s c u e
or
b lu e b u n c h
but on th e South Crow I I s i t e s ta n d in g dead was augm ented
S o u th Crow I I w as th e o n ly s i t e
abundance o f l i t t e r ,
w ith
a n o b s e r v a b le
e s p e c ia lly Idaho fe sc u e (c a s u a lly m easured a t 6 -
10 cm deep). The p e rc e n t s ta n d in g dead f o r Idaho fe s c u e and bluebunch
w h e a tg r a s s w as h i g h e r on S o u th Crow I I th a n any o t h e r s i t e
(NS)
i n d i c a t i n g t h a t th e y e a r o f th e s tu d y w o u ld a l s o be c o n t r i b u t i n g
litte r.
The. l i t t e r
la y e r
m a in ta in e d
m o is tu re
and t e m p e r a t u r e
c o n d itio n s a d e q u a te f o r l e a f e x p an sio n lo n g e r i n t o th e a c tiv e grow th
p e rio d a llo w in g lo n g e r le a v e s .
le s s l i t t e r ,
On t h e o t h e r s i t e s , w h ic h had much
s o i l m o is tu re d e p le tio n d id n o t allo w th e e x p re s s io n of
s ta n d i n g .d e a d e f f e c t s . The s t r o n g a s s o c i a t i o n b e tw e e n Id a h o f e s c u e
l e a f g ro w th and s t a n d i n g d ead w as a f u n c t i o n o f i t s
much d e n s e r
s ta n d in g dead c r e a tin g a c o o le r m o is te r m ic r o s ite . In a d d itio n , Idaho
f e s c u e may e x p e r ie n c e optim um p h o t o s y n t h e s i s a t r a d i a t i o n l e v e l s
r e s u l t i n g from l e s s th a n d i r e c t s u n lig h t (C ald w ell e t a l. 1981).
There w ere some g e n e r a liz a tio n s a p p lic a b le to a l l
The in flu e n c e o f o v e rw in te r le a v e s was im p o rta n t
th re e s p e c ie s .
l a t e r i n th e s p rin g ,
57
fo llo w in g i n i t i a l new l e a f p ro d u c tio n .
The d a te s when s ta n d in g dead
was im p o rta n t corresponded to p la n t p h o n o lo g ic a l s ta g e s w ith h ig h e r
m e ta b o lic demands,
such a s seedhead em ergence and developm ent.
That
a l l s p e c i e s had s i g n i f i c a n t s t a n d i n g d e ad a f f e c t s on S o u th Crow I I
su g g e ste d t h a t m o is tu re a v a i l a b i l i t y
was th e more im p o rta n t f a c to r ,
perh ap s e x a c e rb a te d by th e s ta g e o f developm ent.
Rough fe s c u e and bluebunch w h e a tg ra s s s p rin g p la n t h e ig h ts w ere
not a ffe c te d
g r a z in g .
by th e
p re se n c e
of s p rin g
s t a n d i n g d ead
o r w in te r
Id a h o f e s c u e p l a n t s w ith s t a n d i n g d e a d w e re t a l l e r ,
w in te r g razed p la n ts w ere o n ly o c c a s i o n a l l y s h o r t e r .
but
T h ese r e s u l t s
help ed c o rro b o ra te th e f in d in g s from th e ch em ical c o n s titu e n t a n a ly s is
and f u r t h e r d i f f e r e n t i a t e p h y s io lo g ic a l d if f e r e n c e s betw een th e s p e c ie s .
B lu e b u n c h w h e a tg r a s s p l a n t s w e re e n t i r e l y
d o rm a n t ov_er___t_he
w in te r. They had th e h ig h e s t f i b e r and lo w e s t p r o te in c o n c e n tra tio n s .
The l o s s o f o v e rw in te rin g le a v e s im p lie d no f u r t h e r consequences to
th e p la n t.
I t was th e l e a s t p a la ta b le o f th e th re e s p e c ie s s tu d ie d .
Rough fe s c u e was th e in te r m e d ia te s p e c ie s .
Although i t was no t
m e a s u ra b ly a s p a l a t a b l e a s Id a h o f e s c u e , i t w as c e r t a i n l y th e m o st
p r e f e r r e d g r a s s on t h e S o u th Crow w i n t e r ra n g e .
I t d id h a v e th e
lo w e s t l i g n i n v a lu e s o f th e th re e s p e c ie s . Rough f e s c u e 's lo w e r l i g n i n
le v e ls ,
i n t e r m e d i a t e l e v e l s o f p r o t e i n and f i b e r c o n s t i t u e n t s and
g re e n is h
w in te r le a v e s
a ll
in d ic a te d
so m e l e v e l
o f w in te r
p h o t o s y n t h e t i c a c t i v i t y . F a l l f l o r a l p r i m o r d i a i n i t i a t i o n and slo w
w in te r developm ent have been r e p o r t e d i n ro u g h f e s c u e ( J o h n s to n and
McDonald 1967) f u r th e r in g th e demand f o r a w in te r r e s p i r a t o r y energy
supply,
58
Id a h o f e s c u e w as t h e m o st p h y s i o l o g i c a l l y a c ti v e p la n t through
th e w i n t e r .
P l a n t s h ad many g r e e n i s h w i n t e r l e a v e s .
c o n s titu e n t v a lu e s ,
to midsummer v a lu e s .
The w i n t e r
o th e r th an l i g n i n and h e m ic e llu lo s e , w ere s im ila r
L oss o f m ost w in te r le a v e s d e p re sse d l e a f grow th
i n t h e f o l l o w i n g s p r i n g . More i n d i v i d u a l Id a h o f e s c u e p l a n t s w e re
g ra z e d th a n e i t h e r o f th e o th e r two s p e c ie s i n s p i t e o f p e rio d s o f i c e
c r u s tin g .
F a ll i n i t i a t i o n o f f l o r a l p rim o rd ia h as a ls o been re p o rte d
i n Idaho fe s c u e (Hodgson i n J o h n sto n and McDonald 1967).
A ll o f th e se
c o n s id e r a tio n s su g g e ste d r e l a t i v e l y high w in te r m e ta b o lic demands, to o
high to be met by s to re d n u t r i e n t s alo n e.
WINTER NUTRIENT ALLOCATION PATTERNS
The w i n t e r c h e m ic a l c o n s t i t u e n t an d l e a f l o s s d a ta p lu s t h e
l i t e r a t u r e re v ie w ed su g g e ste d f a l l - w i n t e r n u t r i e n t s t r a t e g i e s f o r each
s p e c ie s .
Bluebunch w h e a tg ra s s i s one o f th e most e x te n s iv e ly s tu d ie d
ra n g e g r a s s e s . The f o l l o w i n g i s g e n e r a l l y s u p p o r te d by t h a t body o f
lite ra tu re
(p re v io u s ly
c ite d ).
A f te r
seedhead
fillin g
or
le a f
e x p an sio n i n n o n rep ro d u c tiv e p la n ts ,
new p h o to s y n th a te i s d ir e c te d to
r o o t crown n u t r i e n t s to ra g e p o o ls.
As summer w a te r s t r e s s deepens,
more a v a ila b le n u t r i e n t s ,
c e l l s o lu b le s ,
p r o te in s and some m in e ra ls ,
a r e t r a n s l o c a t e d fro m a e r i a l t i s s u e t o t h e r o o t c ro w n s.
When f a l l
m o is tu re o c c u rs a d d itio n a l p h o to s y n th a te i s produced and tr a n s lo c a tio n
may be r e v e r s e d . Upon c e s s a t i o n o f f a l l g ro w th and o f t e n c o i n c i d e n t
w ith th e f i r s t f r o s t , e a s i l y m e ta b o liz a b le com ponents from th e le a v e s
a re
ra p id ly
n u trie n t
tra n s lo c a te d
in to
r o o t c ro w n s t o r a g e
p o o ls ' s u p p o r t a v e r y low
p o o ls .
m a in te n a n c e l e v e l
T hese
o f w in te r
r e s p i r a t i o n and f u e l th e p ro d u c tio n o f th e i n i t i a l new s p rin g le a v e s .
So by th e th re e l e a f s ta g e c a rb o h y d ra te r e s e r v e s a re . n e a rly d e p le te d .
From midsummer through March,
Idaho fe sc u e ch em ical c o n s titu e n t
l e v e l s w ere c o n s ta n t. L ig n in and h e m i c e l l u l o s e w e re th e e x c e p t i o n s ,
both b ein g h ig h e r i n w in te r p la n ts ( ta b le 15). There ap p eared to be no
s i g n i f i c a n t sum m er o r f a l l
t r a n s l o c a t i o n fro m th e l e a v e s a n d /o r
c o n tin u e d p h o to s y n th e tic n u t r i e n t f i x a t i o n .
E v e rg re e n s h r u b s have b een show n t o h a v e lo w e r r a t e s o f c a rb o n
fix a tio n
d u rin g
th e
g ro w in g
season,
le s s
tra n s lo c a tio n
of
p h o to s y n th a te o u t o f t h e l e a v e s and low r a t e s o f p h o t o s y n t h e t i c
a c t i v i t y on f a v o ra b le w in te r days.
s i m i l a r s tr a te g y .
Idaho fe s c u e a p p e a rs to employ a
i
E vergreen p la n ts a ls o have th ic k l e a f c u t i c l e s t h a t
red u ce l e a f m o is tu re lo s s . The an alogous a d a p ta tio n i n Idaho fe sc u e i s
i t s v e ry d e n se ly tu f t e d grow th form .
Chapin e t a l . (1979) have sAown
t h a t th e c a e s p i t o s e g ro w th fo rm c r e a t e s a n i n t e r i o r th e r m a l re g im e
s e v e r a l d e g r e e s w a rm e r th a n a m b ie n t t e m p e r a t u r e s . A d e n s e ly t u f t e d
bunch a l s o r e d u c e s w in d and s o l a r e v a p o r a t i o n and i n c r e a s e s t h e
in te rn a l
re la tiv e
h u m id ity .
Id a h o f e s c u e ,
fo llo w in g
ra p id
and
m e t a b o l i c a l l y d e m a n d in g g ro w th s t a g e s , s e t t l e s i n t o a c o n d i t i o n o f
s t a s i s d u rin g w hich th e p la n t m a in ta in s i t s e l f through submaxima l and
o p p o r tu n i s tic p h o to s y n th e s is from summer through f a l l and w in te r.
Rough fe s c u e had an in te r m e d ia te s tr a te g y s h a rin g c h a r a c t e r i s t i c s
w ith
b o th o t h e r s p e c i e s .
le a v e s ,
Rough f e s c u e had many g r e e n i s h w i n t e r
but th e r e was a change i n ch em ical c o n s ti tu e n t v a lu e s betw een
sum m er and w i n t e r ( t a b l e 1 5 ). L ig n in w as t h e o n ly e x c e p tio n .
Rough
f e s c u e a p p e a r e d t o re s p o n d t o w i n t e r i n a m anner s i m i l a r t o t h a t
60
re p o rte d
fo r
E rio p h o ru m
v a g in a tu m .
W ith th e c o m p le tio n o f l e a f
e lo n g a tio n and re p ro d u c tio n , p h o to s y n th a te and some t i s s u e components
a r e tr a n s lo c a te d to r o o t crown s to ra g e p o o ls.
F o llo w in g f a l l reg ro w th
th e r a t e o f tr a n s lo c a tio n in c r e a s e s (Chapin e t a l . 1979,
I 980 and R o b erts and Woolhouse 1984a,b).
Chapin e t aL
Some p h o to s y n th e tic p ig m en ts
do re m a in i n t a c t a l l w in te r a s in d ic a te d by th e g re e n is h c o lo r.
That
l i g n i n d id n o t i n c r e a s e b e tw e e n sum m er and w i n t e r w h i l e a l l o t h e r
f i b e r c o n s ti tu e n ts d id , in d ic a te d some l e v e l o f w in te r p ro d u c tio n but
not enough to p re v e n t th e c a ta b o lis m o f some s t r u c t u r a l c a rb o h y d ra te s
f o r r e s p i r a t o r y energy.
Wew s p rin g grow th i s fu e le d
and c u r r e n t p h o to s y n th a te from o v e rw in te rin g le a v e s .
61
SUMMARY AND CONCLUSIONS
METHODS
1) The tr a n s e c t approach used i n t h i s p r o je c t was an am alg am atio n
of l i t t l e
u se d m e th o d s.
The t r a d i t i o n a l a p p ro a c h t o v e g e t a t i o n i n
ran g e management h as been on a p er a c re , p er p a s tu r e or p er v e g e ta tio n
ty p e
b a s i s . A sa r e s u l t
sam p lin g m ethods.
th e re
s u ita b le
no s ta n d a r d
in d iv id u a l
p la n t
F o r tu n a te ly th e s i t u a t i o n i s changing to w ard s more
em phasis on in d iv id u a l p la n ts .
be
a re
fo r
The method used i n t h i s p r o je c t would
m e a s u r in g o b s e r v a b l e p l a n t c h a r a c t e r i s t i c s
and
i d e n t i f i c a t i o n o f in d iv id u a l p la n ts . S p e c ie s w ith o b v io u s in d iv id u a ls
a re
e a s ie r
to
in d iv id u a ls .
deal
w ith
th a n
s m a lle r
p la n ts w ith i n d i s t i n c t
T his method would be d i f f i c u l t to a d a p t to rh izo m ato u s
s p e c ie s .
2) NIR p r e d i c t i o n
m o d e ra te ly s u c c e s s f u l,
th e procedure.
of
c h e m ic a l
c o n s titu e n t
v a l u e s w as o n ly
w hich i n p a r t d e m o n strated th e r e s i l i e n c e o f
Many re c o g n iz e d c a u tio n s and s a fe g u a rd s w ere v io la te d
and s t i l l th e r e s u l t s f o r th e tw o f e s c u e s w e re a c c e p t a b l e b a se d on
v a lu e s from th e w et c h e m istry and th e l i t e r a t u r e .
Most o f th e v a lu e s
f o r b lu e b u n c h w h e a tg r a s s w e re a l s o a c c e p t a b l e .
The m ain p ro b le m s
e n co u n tered i n t h i s p r o je c t w ere sam ple s iz e f o r c a l i b r a t i o n and th e
s m a ll volume o f in d iv id u a l sam ples f o r m u ltip le re a d in g s on th e NIR
sp e c tro p h o to m e te r.
The r e s u l t s o f t h i s p r o je c t should n o t be ta k en a s
a n a rg u m e n t a g a i n s t u s in g NIR, b u t r a t h e r a f i r s t e x p e r i e n c e w h ic h
e lu c id a te d
som e s p e c i a l
p ro b le m s a s s o c i a t e d w ith
te c h n iq u e f o r use i n ra n g e re s e a rc h .
a d a p tin g t h i s
^
62
SUMMER GRAZING EFFECTS
3) Bluebunch w h e a tg ra s s w in te r ch em ical c o n s ti tu e n ts form ed th r e e
d i s t i n c t c l u s t e r s r e p r e s e n tin g c e l l u l o s e , h e m ic e l l u l o s e and p r o t e i n .
The p r o t e i n
c lu s te rs .
c l u s t e r w as i n v e r s e l y
c o rre la te d w ith
th e f i r s t
tw o
Rough fe s c u e w in te r ch em ical c o n s ti tu e n ts form ed a lo o s e ly
c o r r e la te d f i b e r c l u s t e r and a p r o te in c l u s t e r .
Idaho fe s c u e w in te r
chem ical c o n s ti tu e n ts form ed f i b e r and d i g e s t i b i l i t y c l u s t e r s .
4) D is c rim in a n t a n a ly s is o f th e w in te r ch em ical c o n s titu e n t d a ta
show ed t h a t g r a z i n g Id a h o f e s c u e a t some p h e n o lo g ic a l s t a g e s d id
a f f e c t su bsequent w in te r n u t r i t i o n a l v a lu e s . But th e r e d id n o t ap p ear
to be a management im p lic a tio n a s s o c ia te d w ith th e se r e s u l t s .
WINTER GRAZING EFFECTS
5) W in te r g r a z i n g had no a p p a r e n t e f f e c t on ro u g h f e s c u e and
b lu e b u n c h w h e a tg r a s s p l a n t h e i g h t s i n th e n e x t s p r i n g ’ s new g ro w th
p e rio d .
In some c a se s ung razed Idaho fe s c u e p la n ts w ere t a l l e r th an
grazed p l a n t s .
6) The p resen ce o f s ta n d in g dead r e s u l t e d i n t a l l e r Idaho fe scu e
p l a n t s on m o st s i t e s
and i n
ta lle r
ro u g h f e s c u e
and
b lu e b u n c h
w h e a tg ra s s p la n ts on th e South Crow I I s i t e .
7) I t w as c o n c lu d e d t h a t g r a z i n g and s ta n d i n g d e ad w e re i n f a c t
tw o m e a s u re s o f l i t t e r a c c r u a l to t h e s i t e , and t h a t th e im p a c t o f
s t a n d i n g d e ad w as due to
re la tio n s h ip s
s h a d in g w h ic h im p ro v e d p l a n t m o i s t u r e
63
PLANT WINTER PHYSIOLOGICAL ACTIVITY
8) Bluebunch w h e a tg ra s s was dorm ant d u rin g th e w in te r.
l e v e l s w e re lo w e r
P r o te in
and f i b e r l e v e l s w e re h ig h e r th a n i n th e tw o
f e s c u e s . Rem oval o f s t a n d i n g d e a d i n t h e w i n t e r d id n o t im p a c t t h e
n ext s p r in g 's l e a f grow th.
9) Id a h o f e s c u e had t h e h i g h e s t l e v e l o f w i n t e r p h y s i o l o g i c a l
a c t i v i t y o f th e t h r e e s p e c i e s .
W in te r c h e m ic a l c o n s t i t u e n t v a l u e s
w ere not d i f f e r e n t th a n summer v a lu e s , and w in te r p r o te in l e v e l s w ere
h ig h e r th a n f o r th e o th e r two s p e c ie s .
The l o s s o f o v e rw in te rin g l e a f
m a t e r i a l re d u c e d l e a f g ro w th i n t h e n e x t s p r i n g .
L e a v e s re m a in e d
l i g h t g reen d u rin g th e w in te r.
10)
R ough
fe sc u e
p h y s io lo g ic a l a c t i v i t y .
had
an in te r m e d ia te
of w in te r
W in ter ch em ical c o n s titu e n t v a lu e s tended to
l i e b e tw e e n t h e o t h e r tw o s p e c i e s .
su m m e r t o w i n t e r ,
le v e l
but lig n in
G e n e r a lly f i b e r i n c r e a s e d fro m
d id n o t in c r e a s e .
The l o s s o f
o v e r w i n t e r i n g l e a v e s d id n o t a f f e c t l e a f g ro w th i n t h e n e x t s p r in g .
Rough fe s c u e d id have l i g h t g re e n w in te r le a v e s .
MANAGEMENT IMPLICATIONS
11) The p r im a r y w i n t e r f o r a g e m anagem ent c o n s i d e r a t i o n on t h e
South Crow w in te r ran g e i s th e amount o f fo ra g e a v a il a b le ,
because no
summer g ra z in g tr e a tm e n t s u b s t a n t i a l l y im proved w in te r fo ra g e q u a lity .
12) Some g ra s s e s a re not dorm ant d u rin g th e w in te r.
13) A key t o g r a z i n g t o l e r a n c e
p h o to s y n t h e s i z e d u r in g th e w i n t e r .
f e s c u e may b e t t e r
to le ra te
may be a p l a n t 's a b i l i t y
to
P l a n t s su ch a s ro u g h and Id a h o
p e r e n n i a l g ro w in g s e a s o n u t i l i z a t i o n
64
b e c a u s e o f w i n t e r p h o t o s y n t h e t i c p ro d u c tio n , Idaho fe sc u e more th an
rough f e s c u e .
14) C o n c lu s io n s 12) and 13) (a b o v e ) im p ly t h a t i f su ch s p e c i e s
a re grazed a l l y e a r lo n g they w i l l be damaged.
15) P e r e n n i a l g r a s s e s o f g e n e r a w ith n o r t h e r n o r i g i n s , su ch a s
F e s tu c a and F o a . may be h i g h e r q u a l i t y w i n t e r f o r a g e s th a n o t h e r
g ra sse s.
65
LITERATURE CITED
66
LITERATURE CITED
A. A.C. C. 1962. C e r e a l l a b o r a t o r y , m e th o d s ( 7 th
A s s o c ia tio n o f C ereal C hem ists. S t. P au l, MN.
E d.)
A m erican
A. 0. A. C. 1 970. O f f i c i a l m e th o d s o f a n a l y s i s ( 1 1 th E d.) A s s o c ia ­
t i o n o f O f f i c ia l A n a ly tic a l C hem ists. W ashington, D. C.
A n d e rso n , W. E. and D. L. F ra n z e n .
R a n g e la n d s 5:118.
1983.
Rough f e s c u e i n O regon.
A n d e rso n , J . E. and S. J . M cN aughton.
1973. E f f e c t s o f low s o i l
te m p e ra tu re on tr a n s p i r a t i o n , p h o to s y n th e s is , l e a f r e l a t i v e w a te r
c o n t e n t , and g ro w th among e l e v a t i o n d iv e r s e p la n t p o p u la tio n s.
E col. 5 4 :1 2 2 0 -1 2 3 3 .
B a i l e y , A. W. and M. L. A nderso n .
1 978. P r e s c r i b e d b u r n in g o f a
F e s tu c a - S tip a g ra s s la n d .
J. Range Manage. 31:446-449.
Bezeau, L. M. and A. Jo h n sto n . 1962. In V itro d i g e s t i b i l i t y o f range
f o r a g e p l a n t s o f t h e F e s tu c a s c a b r e l l a a s s o c i a t i o n .
Can. J.
P l a n t S c i. 42:692-697.
B l a i s d e l l , J . P. 1958. S e a s o n a l d e v e lo p m e n t and y i e l d s o f n a t i v e
p l a n t s on th e u p p e r S nak e R iv e r p l a i n s and t h e i r r e l a t i o n t o
c e r t a i n c l i m a t i c f a c t o r s . U. S. D. A. Tech. B u ll. no. 11 90.
B l a i s d e l l , J . P. and J . F. P e c h a n e c . 1949. E f f e c t s o f h e rb a g e
re m o v a l a t v a r i o u s d a t e s on v i g o r o f b l u e bunch w h e a tg r a s s and
a rro w le a f b a lsa m ro o t. E col. 30:298-305.
B l a i s d e l l , J. P., A. C. W iese and C. W. H odgson. 1952. V a r i a t i o n s i n
chem ical c o m p o sitio n o f bluebunch w h e a tg ra s s, a r r o w le a f b alsam r o o t, and a s s o c ia te d ran g e p la n ts . J. Range Manage. 5:346-353.
B o lto n , J. K. and R, H. Brown. 1980. P h o to s y n th e sis o f g ra s s s p e c ie s
d i f f e r i n g i n c a rb o n d io x id e f i x a t i o n p a th w a y s . V. R esponse o f
P anicum m aximum. P anicu m m i l i o i d e s and t a l l f e s c u e ( F e s tu c a
a ru n d in a ce a ) to n itr o g e n .
P la n t P h y sio l.
66:97-100.
B ro o k s , J ., I I I . , M. A n d e rso n and P. J . U rn e s s .
re fle c ta n c e a n a ly s is o f fo rag e q u a lity fo r
Manage. 4 8 :2 5 4 -2 5 8 .
1984.
e lk .
In fra re d
J . W ild l.
C a ld w e ll, M. M., J . H. R ic h a r d s , D. A. J o h n s o n , R. S. Nowak and R. S.
D zurec. 1981. Coping w ith h e rb iv o ry : P h o to s y n th e tic c a p a c ity and
re s o u rc e a l l o c a t i o n i n tw o s e m i a r i d A g ro o v ro n b u n c h g r a s s e s .
O ecologia. 5 0 :1 4 -2 4 .
67
C a s a g ra n d a , L. and R. G. J a n s o n c 1957. W i l d l i f e i n v e s t i g a t i o n s
( D i s t . 4 ): B ig Game S u rv e y s and I n v e s t i g a t i o n s . M ontana D ept,
o f F is h and Game. Fed. Aid Com p i. Rep. P r o j. W -7 4 -R -2 , J - A l ,
P t . 3. 19P.
C h a p in , S. F ., I l l , D. A. Jo h n so n and J . D. MeKen d r ic k . 1980.
S easonal movement o f n u t r i e n t s i n p la n ts o f d i f f e r i n g grow th form
i n a n A la s k a n t u n d r a e c o s y s te m : I m p l i c a t i o n s f o r h e r b iv o r y . J.
E co l. 6 8 :1 8 9 -2 0 9 .
C h ap in , S. F., I I I . , K. V anC leve and M. C. C h ap in .
1 979.
S o il
te m p e ra tu re and n u t r i e n t c y c lin g i n th e tu s so c k .grow th form o f
Eriophorum vaginatum .
J. Ecol. 67:169-189.
C la r k e , S. E. an d E. W. T i s d a l e . 1945. The c h e m ic a l c o m p o s itio n o f
n a t i v e , f o r a g e p l a n t s o f s o u th e r n A lb e r ta and S a s k a tc h e w a n i n
r e l a t i o n to g r a z i n g p r a c t i c e s . Can. D ept. A gr. T ech. B u l l . 54.
C o n s ta n , K. J . 1967.
B ig h o rn s h e e p ra n g e u s e , fo o d h a b i t s and
r e l a t i o n s h i p s t o m u le d e e r and e l k i n G a l l a t i n Canyon. M. S.
T h e sis, Montana S ta te U niv., Bozeman. 43p.
Cook, C. W. 1966. C a r b o h y d ra te r e s e r v e s i n p l a n t s (a l i t e r a t u r e
r e v ie w ) . U tah S t a t e A g r ic. Exp. S ta . L ogan, UT. 49p.
D aer, t . an d E. E. W i l l a r d . 1981. T o t a l n o n s t r u c t u r a l c a r b o h y d r a t e
tr e n d s i n bluebunch w h e a tg ra ss r e l a t e d t o grow th and phenology.
J. Range Manage. 34:377-379.
D eSim one, R., M. Thom pson and K. G ro v e r. 1 984. E lk h o rn M o u n ta in s
w i l d l i f e m o n i t o r i n g p ro g ra m .
P r o g r e s s r e p o r t J a n u a r y , 1983
th ro u g h S e p te m b e r, 1 984. M ontana D ep t, o f F is h , W i l d l i f e and
P a rk s . 119p.
D e P u it, E. J. and M. M. C a ld w e ll. 1975. Gas e x c h a n g e o f t h r e e c o o l
sem i d e s e r t s p e c ie s i n r e l a t i o n to te m p e ra tu re and w a te r s t r e s s .
J . E c o l. 6 3 :8 3 5 -8 5 8 .
D ixon, W. J . ( e d .) . 1983. BMDP s t a t i s t i c a l s o f t w a r e .
C a lif o r n ia P ress. B erk eley , CA. pp.725.
U n iv e rs ity of
E u s ta c e , C. D.
1967.
Food h a b i t s , ra n g e u se and r e l a t i o n s h i p s
betw een e lk and l iv e s to c k i n th e G ra v e lly M ountains, Montana. M.
S. T h esis.
Montana S ta te U niv., Bozeman. 55p.
Gammon, D. M. and B. R. R o b erts. 1978. P a tte r n s o f d e f o l i a t i o n d u rin g
c o n tin u o u s and r o t a t i o n a l g r a z i n g o f th e M atopos S a n d v e ld o f
R h o d e s ia . Rhod. J . A g r ic. R es. 1 6 :1 1 7 -1 3 1 .
G o e rin g , H. K. and P. J . V a n S o e st.
1 970. F o ra g e f i b e r a n a l y s i s .
A gr. H andbook 379 ARS, U. S. D. A. 2 0 p.
68
G ordon, F. A. 1968. Range r e l a t i o n s h i p s o f e l k and c a t t l e on e l k
w i n t e r r a n g e , Crow C re e k , M ontana. M. S. T h e s is . M ontana S t a t e
U n iv ,, Bozeman. 52p.
H a rris , J. A. 1967» Some c o m p e titiv e r e l a t i o n s h i p s betw een Agroovron
spicatum and Bromus teoto ru m , EcoL Monogr. 37:89-111,
H a rris , L, E, 1970. N u tr it io n a l re s e a rc h te c h n iq u e s f o r d o m e stic and
' w ild r u m i n a n ts . V o l, I . L0 E0 H a r r i s Pub. L ogan, UT0
H siao, T. C. 1973. P la n t re s p o n se s t o w a te r s t r e s s .
Ply s i o l. 2 4 :5 1 9 -5 7 0 .
Ann. Rev, P la n t
Jo h n sto n , A. 1961. Comparison o f l i g h t l y g razed arid u n g razed ran g e i n
th e fe sc u e g ra s s la n d of s o u th w e ste rn A lb e rta . Can. J. P la n t ScL
41 :6 1 5 -6 2 2 .
J o h n s to n , A, and L0 M0 B ezeau. 1962. C h em ica l c o m p o s itio n o f ra n g e
fo ra g e p la n ts o f th e F e s tu c a s o a b r e lla a s s o c i a t i o n .. Can. J. P la n t
S c i. 42:105^-115.
J o h n s to n , A, and M. D. M acD onald. 1967. F l o r a l i n i t i a t i o n and s e e d
p ro d u c tio n i n F e s tu c a s o a b r e lla T o rre Can. J. P la n t S c i0 47:577583.
K asw orm , W. F0, L0 R0 I r b y and H. B0 I h s l e Pac.
1984. D ie ts o f
u n g u la te s u s in g w in te r ra n g e s i n n o r th c e n tr a l Montana. J 0 Range
M anage. 37:67-71.
K ir s c h , J . B, 1963. Range u s e , r e l a t i o n s h i p t o lo g g i n g , and fo o d
h a b i t s o f th e e l k i n t h e L i t t l e B e l t M o u n ta in s , M ontana. M. S0
T h e sis, Montana S ta te Univ0, Bozeman. 44p.
K n ig h t, H. G., F0 E. H ep n er and A, N e ls o n .
1908.
Wyoming f o r a g e
p l a n t s and t h e i r c h e m ic a l c o m p o s i t i o n - s t u d i e s No. 3. Wyo. Agr0
E x p t. S t a 0 B u l l . 7 6. 119p.
K n ig h t, R0 R.
66p.
I 970.
The Sun R iv e r e l k h e r d .
W ild l0 M onogr.
No. 2 3 .
K n ig h t, R. R. 1961. E lk p o p u l a t i o n t r e n d s , fo o d h a b i t s and ra n g e
r e l a t i o n s h i p s i n th e Sun R iver a re a .
B ig Game R esearch P ro je c t
W-9 8 -R-1, J-B -4 . Montana F ish and Game Dept, 25p»
Kufe ld , R. C0
M anage.
1973. Foods e a te n by th e Rocky M ountain e lk .
26:106-113.
J. Range
Lund, R0 E0 1 983.
MSUSTAT - an i n t e r a c t i v e s t a t i s t i c a l a n a l y s i s
package. Montana S ta te U n iv e rs ity , Bozeman, MT. 124p.
69
M cCall, R. 1939. S easonal v a r i a t i o n i n th e c o m p o sitio n o f blue bunch
f e s c u e . J . A g r ic. R es. 5 8 :6 0 3 -6 16.
M cCall, fl. 1940. The d i g e s t i b i l i t y o f m atu re ran g e g r a s s e s and range
m ix tu re s fe d a lo n e and w ith su p p lem en ts. J. A g ric. Res. 60:3950.
M cC arty, E. C. and R. P r i c e . I 942. G row th and c a r b o h y d r a t e c o n te n t
o f im p o rta n t m ountain fo ra g e p la n ts i n c e n tr a l Utah a s a f f e c te d
by c l i p p i n g and g r a z i n g . U. S. D. A. Tech. B u ll. 81 8 .
M cC reary , 0. C. 1931.
Wyoming f o r a g e p l a n t s and t h e i r c h e m ic a l
c o m p o s i t i o n - s t u d i e s Wo. 9 . Wyo. Agr. E x p t. S ta . B u ll. 184. 23p.
M cC reary , 0.
1927.
Wyoming f o r a g e p l a n t s and
c o m p o s i t i o n - s t u d i e s No. 8.
Wyo. Agr. E x p t.
p p .91-106.
t h e i r c h e m ic a l
S ta . B u ll. 157.
M cIlv ai ne, S. K. 1942. C arbohydrate and n itr o g e n tr e n d s i n blue bunch
w h e a tg ra s s , Aeronvron sn ic a tu m . w ith s p e c ia l r e f e r e n c e to g ra z in g
in flu e n c e s .
P la n t P h y sio l.
17:540-557.
M cLean, A. an d S. W ikeem . 1985a. I n f l u e n c e o f s e a s o n and i n t e n s i t y
o f d e f o l i a t i o n on b l u e bunch w h e a t g r a s s s u r v i v a l and v i g o r i n
s o u th e rn B r i t i s h Columbia. J. Range Manage. 38:21-26.
McLean, A. and S. Wikeem. 1985b. Rough fe sc u e re sp o n se to season and
i n t e n s i t y o f d e f o lia tio n .
J. Range Manage. 38:100-103.
McLean, A. and E. W. T i s d a l e . I 972. R eco v ery r a t e o f d e p l e t e d ra n g e
s i t e s u n d e r p r o t e c t i o n fro m g r a z i n g . J . Range M anage. 2 5 :1 7 8 184.
McLean, A. and E. W. T i s d a l e . I9 6 0 . C h em ica l c o m p o s itio n o f n a t i v e
f o r a g e p l a n t s i n B r i t i s h C o lu m b ia i n r e l a t i o n to g r a z i n g p r a c ­
t i c e s . Can. J . P l a n t S c i. 4 0 :4 0 5 -4 2 3 .
M o r r is , M. S. and J . E. S c h w a r tz .
1957.
M ule d e e r and e l k fo o d
h a b i t s on t h e N a tio n a l B iso n R ange. J. W ild l. M anage. 2 1 :1 8 9 193.
M ueggle r , W. F. 1 972. I n f l u e n c e o f c o m p e t i t i o n on t h e r e s p o n s e o f
b lue bunch w h e a tg ra s s to c lip p in g . J. Range Manage. 25:88-92.
M u eg g le r , W. F. 1 975. R a te and p a t t e r n o f v ig o r r e c o v e r y i n Id a h o
fe s c u e and b lue bunch w h e a tg ra ss. J. Range Manage. 28:198-204.
N o r r i s , K. H., R. F. B a rn e s , J . E. M oore and J . S. Shenk.
1976.
P r e d ic tin g fo ra g e q u a l i t y by i n f r a r e d r e f l e c t a n c e s p e c tr o s c o p y .
J . Anim. S c i. 4 3 :8 8 9 -8 9 7 .
70
N o rto n , B, E. and P. S. J o h n so n .
1981. P a t t e r n o f d e f o l i a t i o n by
c a t t l e g ra z in g c r e s te d w h e a tg ra ss p a s tu r e s . H t h I n t L G ra ssla n d s
Cong. P ro c . L e x in g to n , KY. pp. 4 6 2 -4 6 4 .
P ark, Y. W., . M. J . A n d e rso n , K. H. Asay and A. W. M ahoney.
1983.
P r e d i c t i n g s o l u b l e n i t r o g e n and f i b r o u s f r a c t i o n s i n c r e s t e d
w h e a tg ra s s w ith n e a r - in f r a r e d r e f le c ta n c e sp e c tro sc o p y . J. Range
M anage. 36:529-533.
Q u in to n , D. A,, A. McLean and D. G0 S to u t .
1982.
V e g e ta tiv e and
re p ro d u c tiv e grow th o f blue bunch w h e a tg ra s s i n i n t e r i o r B r i t i s h
Columbia. J. Range Manage. 35:46-51.
R ic k a rd , W. H., D. W. U re sk and J . F. C lin e . 1975. Im p a c t o f c a t t l e
g r a z i n g on t h r e e p e r e n n i a l g r a s s e s i n s o u th - c e n tr a l W ashington.
J. Range Manage. 28:108-112.
R obertson, K. P. and H. W. Wool house. 1984a. S tu d ie s o f th e sea so n al
c o u r s e o f c a rb o n u p ta k e o f E rio p h o ru m v a e in a tu m i n a m o o rla n d
h a b i t a t . I . L e a f p r o d u c tio n and s e n e s c e n c e . J E c o l. 7 2 :4 2 3 435.
R obertson, K. P. and H. W. Wool house. 1984b. S tu d ie s o f th e sea so n al
c o u r s e o f c a rb o n u p ta k e o f E rio p h o ru m v a e in a tu m i n a m o o rla n d
h a b i t a t . I I . The s e a s o n a l c o u r s e o f p h o t o s y n t h e s i s .
J. E c o l.
72:685-700.
R ouse, R. A.
1957.
E lk fo o d h a b i t s , ra n g e u se and m o v em en ts.
G r a v e l ly M o u n ta in s , M ontana. M. S. T h e s is , M ontana S t a t e U n iv .,
Bozeman. 29p.
S a u e r, R. H. 1978. E f f e c t o f re m o v a l o f s t a n d i n g d e a d m a t e r i a l on
grow th o f Agropvron sjpicatum. J. Range Manage. 31:121-122.
S c h a l l e n b e r g e r , A. I 966. B ig game f o r a g e c o m p e t i t i o n i n Sun R iv e r
Canyon. M. S. T h e sis. Montana S t a t e U niv., Bozeman, 43p.
S henk, J . S., I . L anda, M. R. H oover and M. 0. W e s te rh a u s .
1981.
D e s c rip tio n and e v a lu a tio n of. a n e a r in f r a r e d r e f le c ta n c e sp ec tr o c o m p u te r f o r f o r a g e and g r a i n a n a l y s i s . Crop S c i. 2 1 :3 5 5 358.
S m ith , W. K., A. K. Knapp, J . A. P e a r s o n , J . H. V arm an, J . B. Y a v i t t
an d D. R. Young. 1983. I n f l u e n c e o f m i c r o c l i m a t e and g ro w th
fo rm on p l a n t t e m p e r a t u r e s o f e a r l y s p r i n g s p e c i e s i n a .h i g h e le v a tio n p r a i r i e .
Am. MidL N at. 109:380-389.
S m o lia k , S. and A. J o h n s to n . 1968. G e r m in a tio n and e a r l y g ro w th o f
g r a s s e s a t f o u r ro o t-» zo n e t e m p e r a t u r e s . Can. J . P l a n t S c i.
4 8 :119-127.
71
S t e e l , R. G. D. and J . He T o r r i e . . I 960. P r i n c i p l e s and p r o c e d u r e s o f
s t a t i s t i c s . M cG raw -H ill, In c. New York. 48lp.
S te v e n s, D. R. 1965» Range r e l a t i o n s h i p s o f e lk and li v e s t o c k i n th e
Crow C reek d r a i n a g e , E lk h o rn M o u n ta in s , M ontana. M. S. T h e s is .
Montana S ta te U niv., Bozeman. 68p.
S t o d d a r t , L. A.
1946.
Some p h y s i c a l and c h e m ic a l r e s p o n s e s o f
Agroovron sn ic a tu m to herbage rem oval a t v a rio u s sea so n s.
Utah
S t a t e A g r ic. Exp. S t a. B u l l . 324. pp.24.
S t o u t , D. G., A. M cLean a n d D. A. Q u in to n .
1981.
G ro w th an d
p h o n o lo g ic a l d e v e lo p m e n t o f ro u g h f e s c u e i n i n t e r i o r B r i t i s h
Columbia. J. Range Manage. 34:455=459.
T r l i c a , M. J . 1977. D i s t r i b u t i o n and u t i l i z a t i o n o f c a r b o h y d r a t e
r e s e r v e s i n ra n g e p l a n t s P. 7 3 -9 6 .
I n : R. E. S o se b e e (e d .)
R a n g e la n d P l a n t P h y s io lo g y . R ange S c i. S e r i e s No. 4 , Soc. f o r
Range Manage., Denver, CO.
U. S. D. A.
1970.
N a tio n a l F o re s t.
W h ite , L. M. 1 973.
Range Manage.
G ra z in g a c t i v i t y ,
H elena, MT.
E lkh o r n s.
Mimeo.
C a r b o h y d ra te r e s e r v e s o f g r a s s e s :
26:13-18.
H elen a
a r e v ie w .
J.
W illm s , W., A. W. B a ile y and A. McLean. 1 980. E f f e c t o f b u r n in g o r
c lip p in g Agroovron so icatu m i n th e autum n on the s p r in g fo ra g in g
b eh av io r o f m ule d e e r and c a t t l e . J. A pplied Ec o l.
17:69- 84.
W ils o n , A. M., G. A. H a r r i s and D. H. G a te s .
1966.
C u m u la tiv e
e f f e c t s o f c l i p p i n g on y i e l d o f b lu e b u n c h w h e a tg r a s s . J . Range
M anage. 19:90-91.
\
APPENDICES
73
APPENDIX A
R eg re ssio n 'E quations f o r NIR P re d ic te d
Chemical C o n s titu e n t V alues
Table 2 1.
NDF
R eg ressio n e q u a tio n s 1 used f o r th e NIR p r e d ic tio n o f rough fe sc u e chem ical c o n s titu e n t
v a lu e s .
.
3622(1818)1 - 8281(1759) + 4953(1722) +363.8(1445) - 625(1680) + 60.3912
ADF
1101(2348) - 1227(2139) - 227.7(1982) +3251(1759) - 4758(1722) + 1786(1680) + 53.13
ADL
-301.9(2348) + 130.3(2310) +421.5(2270) + 286.4(2139) - 198.1(1778) - 523.1(2100) + 178.6(1445) - .3584
CELL
2724(2348) - 2582(2310) - 189.1(2230) - 165.8(1982) +69.94
HC
1996(2139) + 5895(1818) - 2387(2100) - 475.6(1940) + 1694(1445) - 6204(1680) - 59.86
PRO
89.07(2190) - 1945(1778) +77.64(1940) + 1054(1734) - 314.7(1445) + 1074(1680) + .2152
DM
179.3(2336) - 261.7(2180) +725(1778) - 592.2(1734) - 65.78(1445) + 95.13
ASH
1035(2208) - 1111(2180) + 2618(1734) - 2416(1722) - 185(1445) +48.01
IVOMD
2195(2230) - 2178(2180) + 4030(1818) - 353.3(1940) + 1427(1445) - 4887(1680) - 39.24
1I n each p a ir of
2The last number
numbers the fir s t
is the coefficient and the second is the f ilte r
in each equation is the y- intercept .
WAVELENGTH (NM).
Table 2 2 .
NDF
R eg ressio n e q u atio n s^ used f o r th e NIR p r e d ic tio n o f Idaho fe sc u e chem ical c o n s titu e n t
v a lu e s .
,
-1910(2270)1+ 2962(2230) - 3960(2208) + 2596(2139) + 3442(1778) - 3047(1722) + 13612
ADF
2341(2336) + 595.9(2348) - 2667(2310) + 389.1(2230) - 503.7(2100) - 321.7(1940) + 351.4(1445) - 6.59
ADL
487.3(2348) + 211.6(2230) - 789.7(2139) + 24.07(1982) - .3831
CELL
2555(2348) - 3478(2310) + 2505(1982) + 1082(2100) - 2107(1940) + 496.4(1445) - 829.5(1680) +35.08
HC
-4149(2336) +6162(2348) - 1991(2310) - 269.5(1445) + 64.76
PRO
160.6(2310) + 517(2190) - 517.2(1982) - 638.9(2100) + 440.7(1940) + 5.547
DM
742.6(2270) + 748.8(2190) - 2088(2139) + 282.6(1818) + 533.6(2100) + 52.54(1940) - 251.2(1734) + 79.16
ASH
'
347.8(1982) - 4999(1818) + 1370(1778) + 4013(1759) - 876.6(1734) - 8.11
IVOMD
0.0001(2208) - .0001(2180) +824.2(1940) + 8933(1722) - 8350(1680) +99.26
1IN EACH PAIR OF NUMBERS THE FIRST IS THE COEFFICIENT AND THE SECOND IS THE FILTER WAVELENGTH (NM).
2The
last number in each equation is the y - intercept .
76
T able 23.
R eg ressio n e q u atio n s^ used f o r th e NIR p r e d ic tio n o f
bluebunch w h e a tg ra ss chem ical c o n s titu e n t v a lu e s .
2096(2310)1- 1030(2270) -s- 1776(2230) - 3613(2208) + 201A I2
ADF
-1682(1734) + 1330(1680) + 145.4
ADL
-1 1 4 .6 (1 7 3 4 ) - 43.95(1680) + 39.87
CELL
-904(1734) + 4 2 4 .4(1445) + 137.5
HC
1628(2310) - 1668(2270) + 307.6(1445) - 788 .3 (1 6 8 0 ) +107.1
PRO
-9 4 .7 1 (1 9 8 2 ) + 535.3(1778) - 23.64(2100) - 176.1(1445) - 35.62
DM
889.3(1722) - 1062(1680) + 121.9
ASH
-1 5 7 .1 (2 2 3 0 ) + 580.1(1778) - 310.9(1680) - 3.886
IVOMD
-3090(2230) - .0001(1818) + .0001(1778) - 834.7(1445) + 4 7 2 .6
I In each p a i r of numbers th e f i r s t i s th e c o e f f ic ie n t and th e second
i s th e f i l t e r , w av elen g th (nm).
2 The l a s t number i n each e q u a tio n i s th e y - i n t e r c e p t .
77 .
Table 24.
NIR p r e d ic te d rough fe sc u e summer chem ical c o n s ti tu e n t
v a lu e s and tr e a tm e n ts .
CHEMICAL CONSTITUENTS1
TREATMENT2
10
10
10
10
10
10.
10
10
10
10
11
11
11
11
11
11
11
11
11
11
11
11
NDF
ADF
ADL
CELL
HC
PRO
DM
ASH
IVOMD
%
%
%
%
%
%
%
%
%
66.3
65.7
67.2
66.6
67.5
70.6
65.9
69.0
69.9
69.1
67.3
68.7
67.4
70.5
65.4
65.1
67.1
69.5
66.4
65.5
64.9
62.4
45.1
45.3
44.8
4 5 .0
44.8
4 7 .2
46.4
4 7 .0
45.4
47.1
44.2
46.9
45.4
47.0
4 5 .0
45.6
45.2
4 4 .4
44.6
4 3 .8
45.1
43.6
32.6
3 3 .9
34.8
3 1 .8
34.9
37.9
30.5
32.7
34.4
36.7
33 .3
3 6 .2
34.6
37.6
34.7
35.4
33.9
3 4 .9
33.3
31.6
34 .9
33.3
27.2
2 4 .7
22 .7
23.0
22 .2
21 .8
2 7.0
26 .9
2 4.9
2 2 .3
22.8
18.2
20.9
21.5
21.0
2 0.9
23 .4
2 3.0
24.8
2 8 .3
22.5
20.6
96.6
96.6
96.1
96.1
96.1
96.2
96.1
96.0
96.1
96.1
96.1
96.4
96.5
96.8
95.7
96.0
96.4
96,5
96.2
96.0
96.1
95.7
10.7
10.8
10.8
11.8
10.9
10.3
11.7
9 .9
10.0
11.7
10.3
13.1
11.4
11.9
12.3
12.7
11.8
10.3
10.5
10.1
12.2
12.6
3 .3
3 .3
3 .0
3 .2
2 .8
3 .6
3 .6
3 .9
3 .5
3.1
4 .3
4.1
4 .2
4 .3
3.1
2 .7
3 .3
3 .7
3 .0
2 .9
2.4
2 .3
5 .7
5 .3
6 .3
6 .9
6 .0
5 .7
7.1
6 .0
6 .3
6 .2
6.6
7.5
6 .3
6.1
9.1
7 .8
6 .8
6 .8
6 .0
6 .6
6 .7
8 .3
48.4
4 8 .7
42.1
4 1.8
42.1
4 4 .9
45.1
48.1
45.8
44.6
42.5
40.7
42 .2
45.7
31.9
35.6
42.643.5
44.9
44.2
37.6
32.4
1 The chem ical c o n s ti tu e n ts a re n e u tr a l d e te r g e n t f i b e r (NDF), a c id
d e te r g e n t f i b e r (ADF), a c id d e te r g e n t l i g n i n (ADL), c e ll u lo s e (CELL),
h e m ic e llu lo s e (HC), p r o te in (PRO), d ry m a tte r (DM), a sh (ASH), and
i n v i t r o o rg a n ic m a tte r d isa p p e a ra n c e (IVOMD).
I The tre a tm e n ts a re c o n tr o l (10) and burn (1 1 ).
^ V alues a r e % o f t o t a l sam ple w eig h t on a dry m a tte r b a s is .
78
Table 25.
NIR p re d ic te d Idaho fe s c u e summer chem ical c o n s ti tu e n t
v a lu e s and tre a tm e n ts .
CHEMICAL CONSTITUENTS1
TREATMENT2
10
10
10
10
10
10
10
10
10
10
11
11
11
11
11
11
11
NDF
<
ADF
ADL
CELL
HG
PRO
%
%
%
%
%
68.7
67.0
74.1
74.4
81.2
7 5 .8
73.4
71.9
73.1
69.6
61.7
66.6
72.1
70.1
69.4
6 8 .9
68.6
45.2
4 6 .2
47.0
47.6
52.6
4 8 .8
4 6 .7
45.2
48.4
48.1
46.3
42.1
45.8
37.8
4 1 .8
50.9
46.5
36.5
35.4
38.9
3 9 .8
46.3
41.2
39.1
37.0
41.3
36 .8
36.7
34.7
38.1
3 2 .2
34.6
42.6
39.8
22 .0
19.6
18.7
20.6
19.6
21.6
22.9
21.4
21.4
20.6
15.0
21.1
21.3
23 .9
23.1
21.6
22 .9
3.1
3 .3
1.8
1.9
1.1
2.1
2 .7
2 .7
2.1
2 .3
1.7
2 .2
2.1
2 .6
2 .9
1.9
2 .3
6 .0
5 .8
4.9
4 .9
3.4
4 .6
4 .9
5 .3
5 .2
6.1
5.7
6 .5
5.5
6 .5
6.6
5 .4
6 .3
DM ' ASH
%
%
95.9
96.7
95.6
94.8
94.1
94.5
95.2
95.4
95.6
95.6
97.5
96.5
95.6
95.8
96.6
95.8
96.0
10.8
9 .8
7 .4
9 .3
6.1
8 .5
9.6
10.4
7 .5
10.0
10.4
10.0
6 .6
9 .9
9 .2
9 .3
8.9
IVOMD
'
%
59.4
55 .8
47.7
4 8 .4
34.2
4 4 .4
50.5
55.3
43.0
4 9 .2
53.2
52.6
49.8
65.4
63.0
41.6
46.9
I The chem ical c o n s ti tu e n ts a r e n e u tr a l d e te r g e n t f i b e r (NDF), a c id
d e te r g e n t f i b e r (ADF)» a c id d e te r g e n t l i g n i n (ADL), c e ll u lo s e (CELL),
hem ic e llu lo s e (HG), p r o te in (PRO), dry m a tte r (DM), ash (ASH), and
i n v i t r o o rg a n ic m a tte r d isa p p e a ra n c e (IVOMD),
^ The tre a tm e n ts a r e c o n tr o l (10) and burn (1 1 ).
3 V alues a r e % o f t o t a l sample w eig h t on a dry m a tte r b a s is .
79
APPENDIX B
Frequency of P h e n o lo g ic a l S ta g e s
Dates
4/18
5/10
6/18
Treatment
6/24
100-,
G r a z in g
Legend
EZD GRAZED
BBB UNGRAZED
F
r
e
q
IDO-,
n
c
S ta n d in g
D ead
y
Legend
ZZi n o SD
BH SD
P h e n o lo g ic a l S tages
F ig u re 4 .
Frequency o f Agsp p henolo g lc a l s ta g e s a f t e r two tre a tm e n ts on th e JGI s i t e , s p rin g 1984
Dates
4/18
5/10
6/12
Treatment
6/18
100
6 /2 4
G ra zin g
75-
Legend
EZ3 GR A Z E D
B S UNGRAZED
25-
C XJ
%50“
F
r
e
0-
e
n
c
y
100 -I
S ta n d in g
D ead
75-1
% 50-
Legend
25-
Z 3 NO SD
GBB SD
0m
i
P h e n o lo g ic a l S ta g es
F ig u re 5.
Frequency of Agsp phenolo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e JG II s i t e , s p rin g 1984
Treatment
Date s
4/18
5/10
6/ 1 2
100-1
6/18
6/ 2 4
G r a z in g
Legend
%50-
f Z 3 GR AZ ED
EEB U N G R A Z E D
S ta n d in g
D ead
Legend
Z 3 NO so
BH
SD
P h e n o lo g ic a l S tages
F ig u re 6 .
Frequency of Agsp p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e ML s i t e , s p rin g 1984
Dates
4/18
5/10
6/12
Treatment
6/18
6/24
IOOn
G ra zin g
Legend
C U2
EZ3 GR AZ ED
H S UNGRAZED
100-1
S ta n d in g
Dead
Legend
Z 3 NO SD
BH SD
P h e n o lo g lc a l S tages
F ig u re 7 .
Frequency o f Agsp p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e SCI s i t e , s p rin g 1984.
Dates
4/18
5/10
Treatment
6/12
6/18
6/24
100G ra zin g
75
-I
Legend
% 50-j
F
r
e
25 n
c
0e
IOOn
n
c
75-
I T
I
I
I
I
I
I
I
I
E D GRAZED
BBS U N G R A Z E D
I
r
I
I
S ta n d in g
Dead
y
%5 0 25-
O-
I
Ii
/
Legend
Z Z NO SD
BBS SD
/
P h e n o lo g ic a l S tages
F ig u re 8 .
Frequency o f Agsp p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e SCII s i t e , s p rin g 1984.
Dates
4/18
5/10
6/12
100-1
Treatment
6/18
6/24
G ra zin g
Legend
C >n
EZ3 GR A Z E D
K S UNGRAZED
100
S ta n d in g
Dead
Legend
Z3
NO S D
BH
SD
P h en o lo g lca l S tages
F ig u re 9 .
Frequency o f Feid p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e JGI s i t e , s p rin g 1984
Dates
4/18
5/10
6/12
Treatment
6/18
6/24
100
G ra zin g
75-
Legend
%50F
r
e
25-
q
0
CZD GR AZ ED
EEa U N G R A Z E D
- -
U
e
n
c
y
100-1
S ta n d in g
D ead
75% 50-
Legend
25-
Z Z NO SD
HS SD
0—
P h e n o lo g ic a l S ta g es
F ig u re 10.
Frequency of Feid p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e JG II s i t e , s p rin g 1984
Dates
4/18
5/10
6/12
Treatment
6/18
6/24
100
G ra zin g
Legend
EZ3 GR AZ ED
BB U N G R A Z E D
O4 P
S ta n d in g
Dead
Legend
ZZl NO SD
HS SD
F ig u re 11.
Frequency o f Feid p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e ML s i t e , s p rin g 1984
Treatment
Dates
4/18
5/10
6/18
6/12
6 /2 4
G razin g
Legend
CZD GRAZED
HB U N G R A Z E D
t
I_ _ _ _ _ _ _ _ _ I_ _ _ _ _ _ _ _ _ I
% 50-
I
y
i_ _ _ _ _ _ _ _ _ I_ _ _ _ _ _ _ _ _ I
J _ _ _ _ _ _ _ _ _ I_ _ _ _ _ _ _ _ _ I
n
c
I- - - - - - - - - - - - - - 1- - - - - - - - - - - - - - 1
O4 P
100 -I
Ij
<
S ta n d in g
Dead
Legend
ii I:
Av >
C
6
^
Z 3 NO SD
BBS SD
u
P h e n o lo g ic a l S tages
F ig u re 12.
Frequency of Feid p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e SCI s i t e , s p rin g 1984
Dates
4/18
5/10
6/12
100-1
Treatment
6/18
6/24
G ra zin g
75-
Legend
%5 0 -
C *Q
F
r
e
f Z 3 GRAZED
H S UNGRAZED
25-|
Oj -
e
IOOn
n
c
75-
S ta n d in g
Dead
y
%5 0 -
Legend
25-
Z 3 NO SD
B H SD
0
P h e n o lo g ic a l S tages
F ig u re 13
Frequency of Feid
p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e SCII s i t e , s p rin g 1984
Treatment
Dates
4/18
5/10
6 /1 2
lOO-i
6/18
6/24
G ra zin g
Legend
GR A Z E D
B S UNGRAZED
[ZJ
cr D
mo-.
S ta n d in g
Dead
Legend
Z 3 NO SD
BH SD
P h e n o lo g ic a l S tages
F ig u re 14.
Frequency of Fesc p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e JGI s i t e , s p rin g 1984.
Date s
4/18
5/10
6/12
T reatm en t
6/18
6/24
100
75-
I
G ra zin g
Legend
%501
fZD GR A Z E D
HS U N G R A Z E D
2 5 -|
C U3
F
r
e
I I i i r
e
100-1
n
c
75 -
S ta n d in g
D ead
y
%50 -
Legend
25-
Z 3 NO SD
B H SD
0—
IiW
P h e n o lo g ic a l S ta g es
F ig u re 15.
Frequency o f Fesc p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e JG II s i t e , s p rin g 1984
Date s
4/18
5/10
6/12
Treatment
6/18
6/ 2 4
G r a z in g
Legend
EZ3 GR AZ ED
BBS U N G R A Z E D
CT 3
e
n
c
100-1
S ta n d in g
Dead
75-
y
Legend
5025-t
X
I
I
Z 3 NO SD
BH SD
P h e n o lo g ic a l S ta g es
F ig u re 16
Frequency o f Fesc p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e ML s i t e , s p rin g 1984
Treatment
Da te s
4/18
5/10
6/12
6/18
6/24
G ra zin g
Legend
( 2 3 GRAZED
@ 0 UNGRAZED
C &
F
r
e
e
S ta n d in g
Dead
y
Legend
Z 3 NO SD
HS SD
O ^4"
P h e n o lo g lc a l S tages
F ig u re 17
Frequency o f Fesc p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e SCI s i t e , s p rin g 1984
Dates
Treatment
G ra zin g
Legend
C
fZD GR AZ ED
BEB U N G R A Z E D
100-1
S ta n d in g
Dead
75%50-
Legend
25-
ZZl NO SD
BB SD
0—
P h e n o lo g ic a l S tages
F ig u re 18 .
Frequency o f Fesc p h en o lo g ic a l s ta g e s a f t e r two tre a tm e n ts on th e SCII s i t e , s p rin g 1984.
O
M?7P
D788
c .2