Habitat use and distribution of bison in Theodore Roosevelt National Park
by Jack Eugene Norland
A thesis submitted in partial fulfillment of the requirements for the degree Master of Science in Range
Science
Montana State University
© Copyright by Jack Eugene Norland (1984)
Abstract:
This study described the habitat use and distribution of bison in Theodore Roosevelt National Park.
Data was collected by direct observation of bison herds and by use of fecal transects. Additional data
was collected through the use of 4 marked bison cows. Both percent use from direct observation and
the fecal transect index gave comparable estimates of habitat use. The close agreement meant that
habitat use observed in 1982-83 was not different from the previous years, and absence of mature bulls
in the direct observations did not effect use patterns. Flat grasslands were preferred with use higher
than availability would indicate. Areas of rugged terrain or dominated by trees and shrubs were
avoided with use lower than availability. Habitat use changed over time with levels of use varying
widely in some habitats.
Home ranges of individuals and the overall distribution were very similar. Virtually the whole park was
used by bison, with few areas avoided. Restricted distribution was found during certain parts of the
year. Herds were uniformly distributed during a 15 day period in the immediate area of use. Grazing
activity was significantly different over the habitat types in the park, but not over a more generalized
land classification scheme. Resting and other activities were significantly different over the habitats.
Grazing and other activities had signifcantly different levels over time. Daily activities had a
crepuscular pattern with grazing being the deterministic factor. Bison -moved 1.65 kilometers per day
which did not significantly differ over time. Bison daily changed sites of use and habitats. The second
and third weeks after initial location the distances moved by bison individuals from the initial location
were in a random pattern.
The combined results characterize the bison as a nomadic animal which had preferences for habitats
and localities. Bison are very similar to other large, herding bovids which inhabit grasslands. HABITAT USE AND DISTRIBUTION OF BISON.
IN THEODORE ROOSEVELT NATIONAL PARK
by
Jack Eugene Norland
A t h e s i s su bm itted in p a r t ia l f u lf illm e n t
o f th e req u irem en ts fo r the degree
■ Of
'
Master o f S c ie n c e
in
Range S c ie n c e
MONTANA STATE UNIVERSITY
Bozeman, Montana
December 1984
APPROVAL
o f a t h e s i s s u b m i t t e d by
J a c k Eugene N o r la n d
T h i s t h e s i s h a s b e e n r e a d by e a c h member o f t h e t h e s i s c o m m it t e e
a n d h a s b e e n f o u n d t o be s a t i s f a c t o r y r e g a r d i n g c o n t e n t , E n g l i s h
u s a g e , f o r m a t , c i t a t i o n s , b i b l i o g r a p h i c s t y l e , and c o n s i s t e n c y , and i s
r e a d y f o r s u b m i s s i o n t o t h e C o l l e g e o f G r a d u a te S t u d i e s .
/& Aec-
/% /
D a te
/9 M
D a te
C o-C hairm an, G r a d u a te c o m m ittee
A pproved fo i
1^,
- A ^
v
D a te
Head, M ajor D e p a r tm e n t
Approved f o r t h e
D ate
G r a d u a te Dean
iii
STATEMENT OF PERMISSION TO USE
In p r e s e n t i n g
th is
th e s is
in
p a r tia l
fu lfillm e n t
of
th e
r e q u ir 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 ib 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 un der
r u le s o f th e L ib rary.
B r ie f q u o ta tio n s from t h i s t h e s i s are a llo w a b le
w ith o u t s p e c ia l p e r m issio n , p rovided th a t a ccu ra te acknowledgem ent o f
sou rce i s made.
P er m issio n fo r e x t e n s iv e q u o ta tio n from or rep ro d u ctio n o f t h i s
t h e s i s may be gran ted by my major p r o f e s s o r ( s ) ,
by th e D ir e c t o r o f L i b r a r i e s w hen,
or i n t h e ir ab sen ce,
i n t h e o p in io n o f e i t h e r ,
proposed use o f th e m a te r ia l i s f o r s c h o la r ly purp oses.
th e
Any cop yin g
or use o f th e m a te r ia l in t h i s t h e s i s fo r f in a n c ia l g a in s h a ll not be
a llo w e d w ith o u t w r it t e n p erm issio n .
D ate
fa - 3
iv
ACKNOWLEDGEMENT
I
w a n t t o th an k my c o m m it t e e , D rs. C la y to n M arlow , Lynn I r b y ,
John T a y lo r ,
and K r is H a v sta d ,
fo r a l l
t h e i r a d v ic e and h e lp .
I
e s p e c i a l l y want to thank both C layton Marlow and Lynn Irby who were
in stru m en ta l in my ed u c a tio n a l ex p erien ce h ere.
I ' g r e a t ly a p p r e c ia te d a l l th e h elp g iv en by the s t a f f o f Theodore
R o o s e v e lt N a t io n a l Park.
Thanks go t o H arvey W ickw are,, Bob P o w e ll,
Jim C u t le r , J im Z ahradka, G a r r e tt M oynihan, John R e i s e r , S k ip Snow,
and many o f th e o th er permanent and s e a so n a l em ployees.
I e s p e c ia l ly
want to thank J e f f Bradybaugh fo r a l l h is a s s is t a n c e .
Thanks' a ls o go to Kathy H irsch and M ich ele Girard who both helped
me w ith the v e g e t a t io n and h a b it a t s found in the Park.
Thanks w i l l a lw ays go to th o se unsung h ero es,
th e s e c r e t a r ie s in
th e Animal and Range S c ie n c e Departm ent, who made l i f e go much e a s ie r .
And thanks go to a l l th e f e llo w graduate stu d e n ts who encouraged me on
my way.
T his stud y was fin a n ced by Theodore R o o se v elt N a tio n a l Park, th e
N a t io n a l
Park S e r v i c e ,
Experiment S ta tio n
and in
p art
by th e
M ontana A g r ic u l t u r a l
V
' TABLE OF CONTENTS
PAGE
ACKNOWLEDGEMENT.
................................................
iv
TABLE OF CONTENTS .
LIST OF TABLES .
LIST OF FIGURES.
.
.
v
.
vi i
.....................................
.
.
ix
ABSTRACT.............................................................. ...... .........................................................
INTRODUCTION................................................................................................
LITERATURE REVIEW. .
............................................ ......
.
.............................. 2
STUDY S I T E ......................................................................................•.
METHODS
I
/
.
.....................................................................
.
-. -D ir e c t O b s e r v a t i o n s ................................................
F eca l T ra n sects.
. . . ..
.
. . . . . .
H a b ita t U s e ..................................
. I n d iv id u a l and Herd A s s o c ia tio n s .
Home Range and. D is t r ib u t io n .
.
S ea so n a l D is t r ib u t io n . . . .
.
. .
. .
A c t iv it y P a tte r n s . . . . . .
■ D a ily A c t i v i t i e s . .
M ovem ent..................................
CONCLUSIONS AND SUMMARY .
APPENDICES
. 24
.
.
.
.
.
.
.
24
.24
24
30
30
.3 4
35
39
40
..........................................
LITERATURE C I T E D ................................. ......
...................................
.
.
.
.
7
19
RESULTS ANDDISCUSSION -...................................................................
‘
.
.
' . .
.
.
.
45
.
.
..
-^
.
.
.
.
.
.4 9
.
Appendix.A — Area covered by th e h a b ita t d i v i s i o n s .
53
54
Appendix B - P ercent use by b iso n of. th e c l a s s e s
and h a b ita t d iv is i o n s . . . . .
.
. 60
Appendix C - D a ily movement o f herds in each tim e
p eriod .
75
Appendix D - A c t iv it y a n a ly s is o f v a ria n ce t a b le s .
87
vi
TABLE OF CONTENTS - Continued
PAGE
Appendix E - Average p ercen t o f th e h erd s engaged
in th e d if f e r e n t a c t i v i t i e s among
the time p e r io d s .
. . . . . . .
Appendix F - P ercent o f th e herds engaged in th e
■ d if f e r e n t a c t i v i t i e s during th e
d a y lig h t hours fo r each tim e
p e r i o d ............................ ...... .................................. 98
91
v ii
LIST OF TABLES
TABLE
PAGE
1
2
■
3
4
5
Comparision o f h a b ita t, ty p e s and mapping u n it
names used in t h i s study w ith o th er system s
developed in North Dakota
. ..
. . ..
.
.
14
L is t .of each v e g e ta t io n a l complex and th e propor­
t io n o f each h a b ita t type and. mapping u n it th a t
com prised th e complex ...............................................................
18
Time p e r io d s w ith s t a r t in g and ending d a te s along
w ith the number o f d ir e c t o b s e r v a tio n .d a y s in th e
p e r i o d .........................................................................................
20
Percent use o f c l a s s e s from d ir e c t o b se r v a tio n s and
f e c a l t r a n s e c ts i n th e South U nit . . . . . .
7
25
26
Average num ber.of b iso n f e c a l ch ip s per tr a n s e c t
and a v a i l a b i l i t y o f c l a s s e s .........................................................26
8
9
10
?
12
13
Number o f tr a n s e c t s and average number o f ch ip s
■ per f e c a l tr a n s e c t and standard erro r i n the
c l a s s e s fo r both u n it s .
. . .
6
11
.
Number o f h e c ta r e s c l a s s i f i e d i n th e d if f e r e n t
p h y s io g r a p h ic /v e g e ta tio n a l c l a s s e s fo r both
u n it s .................................................................................................
Percent use from the d ir e c t o b s e r v a tio n s and
p ercent coverage -o f h a b ita t ty p e s , mapping u n it s ,
,and com plexes in th e South U n it.
. . ..
. ..
.
28
F -v a lu e s o f th e main e f f e c t s , th e c l a s s e s and tim e
p erio d s or h a b ita t d iv is i o n s and time p e r io d s, fo r
th e th r e e d if f e r e n t a c t i v i t i e s : g ra z in g , r e s t in g ,
and o t h e r ..............................................
35
M ea n s.o f.th e-a v era g e p ercen t o f a herd engaged in
g r a z in g , r e s t in g , and o th er a c t i v i t y fo r each
c l a s s .........................................................................................
36
Means o f the average p ercen t o f a herd engaged in
g r a z in g , r e s t in g , and o th er a c t i v i t y fo r each
h a b ita t d i v i s i o n ................................
36
Means o f th e ,a v e r a g e p ercen t o f a herd engaged in
. g r a z in g a c t i v i t y in the c l a s s e s over th e 11 tim e
p erio d s
.
37
v iii
LIST OF TABLES - Continued
TABLE
PAGE
13
14
15 .
16
17
18
19
20
21
22
23 .
. .
Means o f the d a ily d is t a n c e s in k ilo m e te r s moved
by b iso n herds in th e tim e p e r io d s .
. . . . .
41
ANOVA ta b le o f th e lo g a r it h m ic a lly transform ed
v a lu e s o f th e d a ily d is ta n c e moved by b ison
h erd s Over th e 11 time p e r io d s ..........................................
41
G-t e s t s o f th e d is t a n c e s between two random p o in ts
a g a in s t r e lo c a t io n a number o f days l a t e r
43
H ecterage o f h a b ita t ty p e s , mapping u n it s , and
com plexes a s they occu r in each p h y sio g r a p h ic /
v e g e t a t io n a l c l a s s in th e North, U nit .............................
55
H ecterage o f h a b ita t ty p e s , mapping u n it s , and
com plexes a s th ey occur in each p h y sio g ra p h ic/
v e g e t a t io n a l c l a s s in th e South U nit . . . . .
57
ANOVA t a b le s o f g ra zin g a c t i v i t y among th e
c l a s s e s and tim e p e r io d s . . ..........................................
88
ANOVA t a b le s o f r e s t in g a c t i v i t y among th e
c l a s s e s and tim e p erio d s ........................................................
88
ANOVA t a b le s o f oth er a c t i v i t y among th e
" c la s s e s and tim e p e r i o d s .......................................................
89
ANOVA t a b le s o f g ra zin g a c t i v i t y among th e
h a b ita t d iv is i o n s and tim e p e r io d s .
. . .
.
.
89
. ANOVA .ta b le s o f r e s t in g a c t i v i t y among th e
h a b ita t d iv is i o n s and time p e r io d s ...................................
90
ANOVA t a b le s o f o th er a c t i v i t y among th e
h a b ita t d iv is i o n s and tim e p e r io d s ...................................
90
ix
LIST OF FIGURES
FIGURE
•
1
Map o f North U n it, Theodore R o o se v elt
N a tio n a l Park. . . . . . . . . . .
2
Map o f South U n it, Theodore R o o se v elt
N a tio n a l Park.
.
............................
.
3
4
■5
6
9 -3 6 .
37-t47
48-53
54-86
.
PAGE
.
.
.
8
.
.
.
9
A view o f th e p h y s io g r a p h ic /v e g e ta tio n a l c l a s s e s '
w ith in an id e a liz e d laridform o f th e North U n it.
.
10
A view o f th e p h y s io g r a p h ic /v e g e ta tio n a l c l a s s e s
w ith in an id e a liz e d landform o f th e South U n it.
.
II
Home range o f marked anim al 0001 du rin g th e
s t u d y ................................................................................... .
31
.
.
, Home range o f marked anim al 0002 du rin g th e
stu d y . ..............................................................
■■7. ' 8
.
,
32
■L o ca tio n s o f a l l herds ob served during th e
s t u d y ................................................................................................. 33
■ Average d is ta n c e and
i n i t i a l lo c a t i o n and
a f t e r v a r io u s number
average d is t a n c e and
random p o in t s .
. .
stand ard erro r between th e
r e lo c a t io n o f marked an im als
o f d ays, a lo n g w ith th e
standard erro r between two
. . . . . . .
42
.Percent use o f each c l a s s and h a b ita t d iv is i o n
determ ined from d ir e c t o b se r v a tio n o f b iso n h erd s.
'
D a ily m ovem ents,of a l l h erd s ob served fo r each
. . . tim e p eriod .. . . . . .
. . . . . . .
Average p ercen t o f ob served h erd s engaged in
th e d if f e r e n t a c t i v i t i e s among th e c l a s s e s and
h a b ita t d iv is i o n s in each tim e p eriod .
.
. The p ercen t o f observed herds engaged in th e
d if f e r e n t a c t i v i t i e s d u r in g .th e d a y lig h t hours
fo r e a c h ■tim e p eriod
61
.
76
■.
92
99
X
ABSTRACT
T his study d esc r ib e d th e h a b ita t u se and d is t r ib u t io n o f b iso n in
T h eod ore R o o s e v e lt N a t io n a l Park.
D ata w as c o l l e c t e d by d i r e c t
o b se r v a tio n o f b iso n h erd s and by use o f f e c a l t r a n s e c t s .
A d d itio n a l
d a ta w as c o l l e c t e d th r o u g h th e u se o f 4 marked b is o n co w s.
Both
p ercen t u se from d ir e c t o b s e r v a tio n and th e f e c a l t r a n s e c t in d ex gave
com parable e s t im a t e s o f h a b ita t u se.
The c lo s e agreem ent meant th a t
h a b i t a t u se o b s e r v e d i n 1 9 8 2 - 8 3 w as n o t d i f f e r e n t from th e p r e v io u s
y e a r s , and ab sen ce o f mature b u lls i n th e d ir e c t o b s e r v a tio n s did not
e f f e c t use p a tte r n s .
F la t g r a s s la n d s were p refer red w ith u se h igh er
th a n a v a i l a b i l i t y w o u ld i n d i c a t e .
A rea s o f ru g g ed t e r r a i n or
d o m in a te d by t r e e s and s h r u b s w e r e a v o id e d w it h u s e lo w e r th a n
a v a ila b ility .
H a b ita t u se ch a n g ed o v e r t im e w it h l e v e l s o f u se
v a r y in g w id e ly in some h a b ita ts .
Home ran ges o f in d iv id u a ls and th e o v e r a ll d is t r ib u t io n were very
s i m i l a r . V i r t u a l l y th e w h o le park w as u sed by b is o n , w i t h few a r e a s
a v o id e d .
R e s t r i c t e d d i s t r i b u t i o n w a s found during c e r t a in p a rts o f
th e year. Herds were u n ifo rm ly d is t r ib u t e d during a 15 day p eriod in
th e im m e d ia te a r e a o f u s e .
G r a zin g a c t i v i t y w as s i g n i f i c a n t l y
d i f f e r e n t o v e r th e h a b i t a t t y p e s i n th e p ark , but n o t o v e r a m ore
g e n e r a liz e d la n d c l a s s i f i c a t i o n schem e.
R estin g and o th er a c t i v i t i e 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 o v e r t h e h a b i t a t s . G r a z in g and o th e r
a c t i v i t i e s had s i g n i f c a n t l y d i f f e r e n t l e v e l s o v e r t im e .
D a ily
a c t i v i t i e s h ad a c r e p u s c u l a r p a t t e r n w i t h g r a z i n g b e i n g t h e
d e t e r m in is t ic f a c t o r .
B ison -moved 1.65 k ilo m e te r s per day which d id
not s i g n i f i c a n t l y d i f f e r over tim e. B ison d a ily changed s i t e s o f use
and h a b it a t s .
The second and th ir d weeks a f t e r i n i t i a l lo c a t io n th e
d is t a n c e s moved by b iso n in d iv id u a ls from the i n i t i a l lo c a t io n were in
a random p a ttern .
The combined r e s u l t s c h a r a c te r iz e th e b iso n a s a nomadic anim al
w h ic h had p r e f e r e n c e s f o r h a b i t a t s and l o c a l i t i e s . B is o n a r e v e r y
s im ila r to o th er la r g e , h erding b o v id s w hich in h a b it g r a ssla n d s.
I
INTRODUCTION
. K now led ge o f t h e h a b i t a t u se and d i s t r i b u t i o n o f an a n im a l i s
e s s e n t i a l t o u n d e r s ta n d in g i t s e c o l o g y and management.
Because th e
American b iso n has d e c lin e d from m il lio n s o f a n im a ls f r e e ly roam ing
th e g r a ssla n d s o f North America to sm a ll p o p u la tio n s r e s t r i c t e d to a
f r a c t i o n o f t h e i r form er, r a n g e , much o f th e h i s t o r i c a l in f o r m a t io n
d o e s n o t a p p ly t o t h e b is o n ! s c u r r e n t
1982).
s itu a tio n
(R e y n o ld s e t a l .
New s t u d ie s are needed to r e d e f in e how b iso n use t h e ir p resen t
h a b it a t s .
T h is s tu d y
s p e c ific a lly
lo o k e d
a t a . p o p u l a t i o n o f b is o n i n
Theodore R o o se v e lt N a tio n a l Park (TRNP), North Dakota.
The o b j e c t iv e s
w e r e t o d e s c r ib e t h e h a b i t a t u s e and d i s t r i b u t i o n o f b is o n i n t h e
p a rk , i n o r d e r t o d e f i n e
a b le r e s o u r c e s in th e park.
th e s p a t i a l and te m p o r a l u s e o f th e a v a i l ­
T his stu d y was done in co n ju n ctio n w ith a
l a r g e r stu d y m eant to d e te r m in e c a r r y in g c a p a c i t i e s o f b is o n i n th e
park (Marlow e t a l.
1984).
2
LITERATURE REVIEW
The A m erican b is o n i B i s o n b i s o n ) h a s been p r e s e n t on th e N o rth
A m erica n c o n t in e n t s i n c e i t s e v o l u t i o n 4 ,0 0 0 - 5 ,0 0 0 b e f o r e p r e s e n t
(M cDonald 1981).,
The b is o n i s
b e l i e v e d t o h ave o r i g i n a t e d on th e
N orthern Great P la in s and from th e r e r a d ia te d to o th e r p ro v in ces.
At
th e tim e o f c o n ta c t w ith w estern man, b iso n were d is t r ib u t e d from th e
f o o t h i l l s o f th e A ppalachian M ountains to th e Great B a sin and from th e
b o r e a l f o r e s t s o f n o r t h w e s t Canada s o u th to n o r t h e r n M exico.
The
o r i g i n a l .en v iro n m en t o f th e b is o n w as c h a r a c t e r iz e d by w id e open
sp a c e s covered by la r g e g r a ssla n d s.
E v en tu a lly ,
th e b iso n did adapt
t o h a b it a t s dom inated by f o r e s t s and shrublands.
B e f o r e th e c h a n g e s b ro u g h t by w e s t e r n c i v i l i z a t i o n b is o n w e r e
c h a r a c t e r i z e d a s h i g h l y m o b ile a n im a ls u t i l i z i n g l a r g e la n d a r e a s
d u r in g th e y e a r (G a r r e ts o n 19 3 8 , L a r so n 19 4 0 , Roe 19 7 0 , and Dary
1974). B ison, p refer red g r a s s la n d s, and t h e i r d i e t s w ere made up m o stly
o f gram inoid s p e c ie s .
b een d e b a te d f o r y e a r s .
How the b iso n in flu e n c e d i t s
environm ent has
W eaver and C le m e n ts (1 9 3 8 ) s t a t e d t h a t t h e
p la n t c o m m u n itie s i n th e g r a s s l a n d s w e r e i n f l u e n c e d p r im a r ily by
c lim a te and s o i l s , and b iso n a f f e c t e d only, sh o rt term changes.
Larson
( 1940) and M alin ( 1956) f e l t th a t b iso n were a m ajor in f lu e n c e k eep in g
th e g ra ssla n d in a d is c lim a x .
The b is o n 's e n v ir o n m e n t h a s u n d erg o n e many c h a n g e s s in c e th e
co m in g o f w e s t e r n c i v i l i z a t i o n .
The v a s t o r i g i n a l g r a s s la n d s h a v e
been fragm en ted le a v in g o n ly sm a ll t r a c t s o f g r a ssla n d in- which th e
b is o n ca n l i v e .
S in c e th e g r e a t s la u g h t e r o f th e h e r d s a lm o s t a l l
3
b is o n h a v e b een m anaged a s c a p t i v e a n im a ls w it h o n ly Y e llo w s t o n e
N a tio n a l Park and Wood B u ffa lo N a t io n a l Park h a v in g h e r d s d e s c e n d e d
from and
re se m b lin g w ild a n im a ls
(Reynolds e t a l.
S e v e r a l r e c e n t s t u d ie s have in v e s t ig a t e d
p r e se n t s it u a t io n .
Popp (1981),
1982).
th e b is o n in
th e ir
w orking in Wind Cave N a tio n a l Park,
r e p o r t e d t h a t b is o n c h o s e g r a s s s l a n d s f o r th e m a j o r it y o f u s e , and
f o r e s t e d a rea s w ere o n ly used a s t r a v e l co r r id o r s.
S i t e s dom inated by
c o o l s e a s o n g r a s s e s te n d e d t o be p r e f e r r e d i n th e s p r in g and e a r l y
summer by th e cow herds w h ile warm sea so n g r a ss s i t e s w ere p refer red
in
th e mid and l a t e
p refer red i n th e f a l l .
summer.
The c o o l
season
s ite s
w ere a g a in
B u ll groups had d if f e r e n t usage p a tte r n s w ith
th e warm s e a s o n s i t e s b e in g p r e f e r r e d th r o u g h o u t t h e y e a r .
f o r e s t op en in gs w ere used more o f t e n by th e b u ll groups.
A ls o ,
P r a ir ie dog
to w n s r e c e i v e d h ea v y u se d u r in g t h e r u t (m idsum m er t o e a r l y f a l l ) .
The l a r g e s t h e r d s fo u n d i n t h e park w e r e on t h e p r a i r i e dog to w n s
du rin g th e ru t.
Coppock e t a l . ( 19 8 3 ), w o r k in g i n Wind C ave, r e p o r t e d s p e c i f i ­
c a l l y on b iso n use o f p r a ir ie dog tow ns.
They found a d e f in it e s e l e c ­
t i o n f o r p r a ir ie dog tow ns during midsummer. Herds would use th e m ost
r e c e n t ly c o lo n iz e d a rea s fo r g r a z in g w h ile the o ld e r p o r tio n s o f th e
tow ns w ere used fo r o th er a c t i v i t i e s .
T his usage p a tte r n was r e la t e d
t o d i f f e r e n c e s i n v e g e t a t i o n s t r u c t u r e and f o r a g e q u a l i t y b ro u g h t
about by th e h erb iv o ry o f th e p r a ir ie dogs.
In U tah Van Vuren (1979 and 1 9 8 3 ),
w o r k in g w it h an i s o l a t e d
p o p u la tio n o f b iso n in th e Henry M ountains,
fe r e d from c a t t l e in s e v e r a l r e s p e c t s .
rep o rted th a t b iso n d i f ­
B ison tended to use h a b it a t s
4
fa r th e r away from w a ter so u rces,
tio n s .
s te e p e r lan d form s,
and h ig h er e le v a ­
B ison used h a b it a t s w hich were dom inated by t h e ir p referred
f o r a g e , g r a s s and g r a s s l i k e p l a n t s .
C a t t l e a v o id e d t h e sam e s i t e s
because o f th e d is ta n c e from w a ter, s te e p n e s s , and e le v a t io n .
Van Vuren (1979 and 1983) a d d it io n a lly rep o rted th a t b iso n w ere a
h i g h l y m o b ile a n im a l s t a y i n g an a v e r a g e o f 1.9 d a y s a t a s i t e b u t
r a r e ly lo n g er than 3 days.
The changes in b iso n d is t r ib u t io n during
th e s tu d y w e r e a t t r i b u t e d t o s e a s o n a l and y e a r l y d i f f e r e n c e s
in
f o r a g e and w e a th e r p a t t e r n s . The home r a n g e s o f s e v e r a l b is o n co w s
o v e r la p p e d w it h th e a v e r a g e s i z e b e in g 52 sq u a r e k i l o m e t e r s (km2 ).
Herds tended to in c lu d e 15 in d iv id u a ls or few er, but th e s i z e o f herds
w as p o s i t i v e l y c o r r e l a t e d w it h th e s i z e o f f o r e s t o p e n in g s . S o c i a l
s t r u c t u r e o f th e h e r d s and t h e a s s o c i a t i o n o f i n d i v i d u a l s w as d e ­
s c r ib e d a s f lu id .
L o tt and M inta (1983) rep o rted on group s t a b i l i t y and in d iv id u a l
a s s o c ia t io n s o f b iso n cows on Santa C a ta lin a I s la n d , C a lifo r n ia .
They
rep o rte d th a t m ature cows a s s o c ia t e d in a random p a tte r n , and th e o n ly
c lo s e a s s o c ia t io n rep o rted was betw een cows and c a lv e s .
The c o w -c a lf
a s s o c ia t io n e v e n t u a lly became random a f t e r c a lv e s reached a g es o f 8-19
months.
Herd s i z e d id not fo llo w a. random p a ttern but was in flu e n c e d
by. en viron m en tal f a c t o r s .
Some o f th e en vironm en tal f a c t o r s m entioned
were- w a te r in g lo c a t i o n s , topography, fo r a g e q u a lit y and phenology, and
s o c i a l i n t e r a c t i o n s su ch a s t h e r u t .
Home r a n g e s f o r co w s a v e r a g e d
56.1 km2 w it h p o r t i o n s o f a l l home r a n g e s o v e r la p p in g .
Movement o f
i n d i v i d u a l s i n a s t r a i g h t l i n e a v e r a g e d 2.8 k i l o m e t e r (km) p er day.
T h is d id n o t ch a n g e s i g n i f i c a n t l y o v e r th e s e a s o n s but. d id in c r e a s e
5
'som ew h at d u r in g t h e r u t .
The a u t h o r s c h a r a c t e r i z e d t h e b is o n a s a
h ig h ly m o b ile a n im a l, w it h s i m i l a r home r a n g e s , and b e in g d r iv e n
in d iv id u a lly by en viron m en tal ra th er then s o c ia l fa c t o r s .
S e v e r a l a u t h o r s have d e s c r ib e d h a b i t a t u se. by b is o n i n Wood
B u f f a l o (S o p er 1 9 4 1 , R ey n o ld s e t a l . I 9 82) and Y e llo w s t o n e N a t io n a l
P ark s ( McHugh 1958,
M eagher 19 7 3 ).
B oth p a rk s a re d o m in a ted by
f o r e s t s a lt h o u g h b is o n a lm o s t e x c l u s i v e l y choose a r e a s dom inated by
gram in oid s.
The d is t r ib u t io n o f the g r a s s la n d s li m it e d th e range o f
b iso n in th e parks.
Although o th er h a b it a t s and lo c a t io n s w ere chosen
d u r in g th e y e a r , t h i s w as th o u g h t to be r e l a t e d t o snow d e p th s and
lo c a l c l im a t ic c o n d itio n s .
McHugh
(1 9 5 8 )
in v e s t ig a t e d
th e
b e h a v io r
of
b is o n
Y ello w sto n e N a tio n a l Park and s e v e r a l o t h e r l o c a t i o n s .
herds
in
He r e p o r t e d
th a t th e home range fo r Y ello w sto n e N a tio n a l Park b iso n was ap p roxi­
m a t e ly 31 km
i n th e summer and 93 km
i n th e w in t e r .
Movement o f
b iso n in th e W hichita Mountains N a tio n a l W ild lif e Refuge averaged 3.4
km p er day and t h i s d a i l y m ovem ent w as common t o a l l th e h e r d s .
He
ob served b iso n g o in g to' w ater o n ly once a day.
P eden e t a l . ( 1 9 7 4 ) , w o r k in g w it h c a t t l e and b is o n i n n o r t h e a s t
Colorado,
rep o rted th a t b iso n used uplands more than c a t t l e .
w ere found mohe o f t e n in the drainagew ays.
w orking in th e same area,
C a ttle
Schw artz and E llis ( 1 9 8 3 ) ,
rep orted th a t b iso n w ere s e l e c t i n g fo r th e
m ost abundant fo r a g e the dominant g r a s s e s .
They th e o r iz e d th a t b iso n ,
because o f t h e i r la r g e s i z e and energy needs, would have to s e l e c t fo r
th e m ost a v a ila b le fo r a g e in an area.
6
From th e l i t e r a t u r e rev iew i t appears th a t th e b iso n o f today: I)
p r e fe r g r a s s la n d s ,
3)
2) w i l l , .change h a b ita t, p refer en ce during th e y ea r,
have d if f e r e n c e s i n h a b ita t p r e fe r e n c e s betw een th e s e x e s , 4) have
la r g e home ranges,. 5) w i l l u t i l i z e m ost a l l of. th e a v a ila b le h a b ita t,
6)
move c o n s t a n t l y from s i t e
t o site" , and 7 ) have a f l u i d s o c i a l
s t r u c t u r e w it h n o lo n g te r m . a s s o c i a t i o n s .
D i f f e r e n c e s i n th e home
range s i z e s w ere r e la t e d to th e d if f e r e n t s i z e s o f a v a ila b le p referred
habi t a t .
7
STUDY SITE
The e n t i r e s tu d y w as c o n d u c te d i n T h eod ore R o o s e v e lt N a t io n a l
Park.
The Park was cre a ted in 1978, but
had p r e v io u s ly been Theodore
R oo.sevelt Memorial Park e s t a b lis h e d in 1947.
The Park i s composed o f
tw o u n i t s , th e N orth U n it (NU) and S o u th U n it (SU) ( F ig u r e s I and
' ■'
.
.
2 ).
The SU i s 18 ,7 5 6 h e c t a r e s i n s i z e and th e NU 9 ,7 4 1 h e c t a r e s in
s iz e .
Both u n i t s a r e l o c a t e d i n w e s t e r n N orth D a k o ta , t h e SU n ea r
M edora, N orth D ak ota and th e NU 80 k i l o m e t e r s (km) d i r e c t l y n o r th .
B ison w ere rein tr o d u ce d in t o th e Park in 1956.
The dom inant f e a t u r e s in both u n it s are the L i t t l e M isso u ri R iver
and t h e v a l l e y
and b a d la n d s t h a t i t h a s fo rm ed .
D u rin g t h e m ost
r e c e n t g l a c ia t io n th e mouth o f th e L i t t l e M issou ri R iver was low ered
w hich a c c e le r a te d th e e r o s io n a l fo r c e s in th e s o f t sed im en tary rock
p r o d u c in g th e b a d la n d f o r m a t io n s fo u n d i n th e Park (H ansen et- a l .
1980).
The p a st and ongoing e r o s io n has produced a v a r ie d topography
w hich r e s u l t s i n a v a r ie t y o f v e g e t a t io n a l com m unities i n the Park.
Two d if f e r e n t schem es w ere used t o d e sc r ib e th e lan d in th e Park.
One schem e in v o lv e d d iv id in g the u n it s in t o p h y s io g r a p h ic /v e g e ta tio n a l
c l a s s e s (from now on r e f e r r e d t o a s c l a s s e s ) b a se d on a p p e a r a n c e ,
landform o r ig in , and the g r o ss s tr u c tu r e o f the a s s o c ia t e d v e g e t a t io n
(tr e e s,
g r a s s la n d s ,
sh ru b s,
e t c .) .
A to ta l
of
p h y s io g r a p h ic /v e g e t a t io n a l c l a s s e s w e r e i d e n t i f i e d .
w ere unique to one or the oth er u n it s ,
in both u n its (Table I).
each c l a s s i s found.
F ig u res 3
14
d iffe r e n t
S ev en c l a s s e s
w h ile the o th e r s w ere p resen t
and 4 in d ic a t e w here i n the u n it s
A g en era l d e s c r ip t io n o f each c l a s s f o llo w s :
BM
2636
North Un i t
KILOMETERS
F ig u re I .
Map o f N o r t h U n i t , T h o e d o re R o o s e v e l t N a t i o n a l P a r k .
PETRIFIED
FOREST -
KILOMETERS
f P t a ci t ul V i l t y
PoncP
Plowing W l l
F ig u re 2 .
Map o f S o u th U n i t , T h o e d o re R o o s e v e l t N a t i o n a l P a r k .
ACHENBACH
UPLAND
HILLS
GRASSLANDS
BREAKS
TOE SLOPE
WOODED DRAW
ROLLING /GRASSLANDS
TTOM
GRASSLANDS
SAGEBRUSH
F ig u re 3.
COTTONWOODS
BOTTOMS
A v ie w o f t h e p h y s i o g r a p h i c / v e g e t a t i o n a l c l a s s e s w i t h i n an i d e a l i z e d la n d f o r m
o f th e N o rth U n i t .
/ 0 LD
RIVER
TERRACE
SCORIA
HILLS
BREAKS
WOODED DRAWS
RI DGE
RAVINE
U P L / N D1 G R A S S L A N D
GRASSLAND
SAGEBRUSH
F ig u re 4.
FLATS
BOTTOMS
A v i e w o f t h e p h y s i o g r a p h i c / v e g e t a t i o n a l c l a s s e s w i t h i n an i d e a l i z e d la n d f o r m
o f t h e S o u th U n i t .
12
1. B r e a k s : c o n s i s t o f a r e a s n o t i c e a b l y d e v o id
v e g e t a t io n or i f v e g e t a t io n d oes e x i s t , i t
s it u a t e d on s te e p s lo p e s .
of
is
2 . Cottonwood F o r e s ts : g a ll e r y f o r e s t s a lo n g p eren n ia l
w a te r c o u r s e s d o m in a te d by p l a i n s c o t t o n w o o d
( P o p u lu s d e l t o i d e s ). L arge e x p a n s e s In t h e NU a r e
d evoid o f a dense shrub u n d erstory and have in s te a d
a dense cover o f g r a s s and f orbs.
3 . Wooded D raw s: w oody s t r i n g e r s d o m in a ted by e i t h e r x
g ree n ash CFraxinus p en n sv lv a n ica ) or aspen CPoduI us
t r e m u l o i d e s ).
The w ooded d ra w s a r e u n if o r m ly
s c a tte r e d over both u n it s o f th e Park.
4 . Upland G rasslan d s: l e v e l to r o l l i n g g r a s s la n d s found
on t h e p l a i n s a b o v e t h e r i v e r v a l l e y . T h ese la n d s
are t y p ic a l o f the north ern Great P la in s.
5 . Old R iver T erraces: l e v e l g r a ssla n d s 200 to 500 f e e t
a b o v e th e r i v e r w h ich a r e s i t u a t e d on t e r r a c e s
form ed b e fo r e th e ra p id d o w n -cu ttin g o f th e r iv e r .
6 . G r a s s la n d F l a t s : l a r g e f l a t g r a s s e d a l l u v i a l
d e p o s i t s fo u n d 100 to 200 f e e t a b o v e t h e r i v e r
v a ll e y .
7 . B ottom G r a s s la n d s : l a r g e f l a t g r a s s e d a l l u v i a l
d e p o s it s found on h ig h er f lo o d p la in s o f th e L i t t l e
M isso u ri and i t s la r g e r t r ib u t a r ie s .
8 . Toe S l o p e s : g r a d u a lly s l o p i n g la n d s fo rm ed
s lu m p in g and a l l u v i a l d e p o s i t i o n .
The la n d
covered w ith g r a s s , shrubs, and tr e e s .
by
is
9 . R o l l i n g G r a s s la n d s : l e v e l t o r o l l i n g g r a s s la n d s
fo u n d on th e g l a c i a t e d p l a i n s a b o v e t h e r i v e r
v a ll e y .
10.
A chenbach H i l l s : h i l l s fo u n d 650 f e e t a b o v e th e
r i v e r w h ic h a r e c o v e r e d by g r a s s and s h r u b s . The
la r g e b ou ld er f i e l d s on th e s lo p e s o f th e h i l l s are
the d e b r is o f a bedrock cap.
11. R id g e and R a v in e : l a n d s h i g h l y d i s s e c t e d by
w a te r c o u r se s and covered by v a r io u s g r a s s e s , shrubs,
and t r e e s .
12. S c o r ia H i l l s : la n d s in flu e n c e d by s c o r ia (a c lin k e r
form ed from the baking o f c la y s a d ja cen t to burning
co a l v e in s ) which produces a d i f f e r e n t i a l w ea th erin g
13
o f th e lan d . T his w ea th erin g produces a very rugged
and v a r i e d to p o g r a p h y w h ic h i s c o v e r e d by v a r i o u s
• g r a s s e s and shrubs.
13. Sagebrush Bottom s: f lo o d p la in s dom inated by s i l v e r
sageb ru sh (A r te m isia cana) a lo n g w ith a s u b s t a n t ia l
g r a s s cover.
14.. P r a i r i e Dog Towns: la n d s w h ic h h a v e b een or a r e
b e in g in f l u e n c e d by p r a i r i e d o g s. At th e e d g e s o f
tow ns p la n ts a re s t i l l c h a r a c t e r is t ic o f th e former:
p la n t community.
Nearer the c e n te r , v e g e t a t io n i s
a b sen t or dom inated by u n p a la ta b le p e r e n n ia l p la n t
s p e c ie s .
T able I g iv e s th e area o f each c l a s s found in th e d if f e r e n t u n its .
Table I .
Number
of
h e c ta r e s
c la s s ifie d
in
th e
d if f e r e n t
p h y s io g r a p h ic /v e g e ta tio n a l c l a s s e s fo r both u n it s ( r iv e r
ch a n n els in c lu d e d ) .
H ecterage
P h y sio g r a p h ic /V e g e ta tio n a l C la sse s
Breaks
Sagebrush Bottoms
Wooded Draws
Upland G rasslan d s
P r a ir ie Dog Town
Cottonwood F o r e sts
R iver Bottom
G rassland F la t s
Bottom G rassland
Ridge & Ravine
Toe S lo p es
Old R iver T erraces
Achenbach H i l l s
SU
357 8 .3
1174.8
424.1
TOI 0. 5
168.3
88.6
8 7 .4
1 9 4 9 .0
6879.5
1012.5
-
NU
2 9 04.1
911.4
37 3 .5
4 1 9 .5
48 .1
51 9 .6
171.4
8 9 5 .8
1956.3
7 7 .0
The second schem e was based on h a b ita t ty p es (HT) w hich have been
p r e v io u s ly d escrib ed by o th e r s w orking in th e im m ed iate area (Whitman
19 7 8 , H ansen e t a l . 19 8 0 , K. H ir s c h ( p e r s o n a l c o m m u n ic a tio n , N o rth
D ak ota S t a t e U n i v e r s i t y ) , M. G ira rd ( p e r s o n a l c o m m u n ic a tio n , N orth
14
D ak ota S t a t e U n i v e r s i t y ) , and S o i l C o n s e r v a t io n S e r v i c e (ra n g e s i t e
g u id e lin e s fo r th e Badlands o f North Dakota).
The h a b ita t ty p es were
d e fin e d a s th e v e g e t a t io n a ssem b la g es t h a t e v e n tu a lly would d ev elo p on
a s i t e g iv e n a s p e c i f i c c lim a te and s u b s tr a te .
Table 2 shows each HT
rec o g n ized and th e corresp on d in g c l a s s i f i c a t i o n by th e auth ors.
Table 2 .
Comparison ' o f h a b ita t typ e (HT) and mapping u n it (Mt)) names
used in t h i s stud y w ith o th er system s developed in North
Dakota (Whitman 1978, Hansen e t a l . 1980, K. H irsch (p er­
so n a l com m unication, North Dakota S ta te U n iv e r s it y ) , M.
Girard
(p erso n a l
com m unication,
North
Dakota
S ta te
U n iv e r s it y ) , and S o il C on servation S e r v ic e ( range s i t e
g u id e lin e s f o r th e Badlands o f North D akota).
Agropyron s m it h ii - S tip a v ir id u l a HT
H ir s c h :
Whitman
SCS:
Agropyron s m it h ii
H ir s c h :
Whitman:
Hansen e t a l . :
SCS:
Agropyron s m it h ii - S tip a v ir id u la *
Agropyron - S tip a v ir id u l a - B ou telou a and
Agropyron - S tip a v ir id u la - Mixed
Clayey range s i t e
S tip a comata HT
Agropyron s m it h ii
Agropyron - S tip a
Agropyron s m it h ii
S i l t y range s i t e
S tip a comata*
B outeloua
Carex f i l i f o l i a
S tip a comata - B ou telou a g r a c i l i s HT
H ir s c h :
Whitman:
Hansen e t a l . :
SCS:
S tip a
S tip a
S tip a
Sandy
comata - B outeloua g r a c i l i s *
- C alam ovilfa - Carex
comata - Carex f i l i f o l i a
range s i t e
A rtem isia eana HT
H ir s c h :
Hansen e t a l . :
A rtem isia cana*
A rtem isia cana - Agropyron s m it h ii
Andropogon sc o p a r iu s - Juniperus h o r iz o n t a l is HT
H ir s c h :
Andropogon s c o p a r iu s - Ju n ip eru s h o r iz o n t a lis *
Hansen e t a l . : J u n ip eru s h o r iz o n t a l is - Andropogon sco p a riu s
SCS:
Very sh allow range s i t e
15
Table 2 . Continued
A rtem isia t r id e n t a t a - A tr ip le x c o n f e r t i f o l i a HT
H ir sc h :
SCS:
A rtem isia t r id e n t a t a - A tr ip le x c o n f e r t i f o l i a 9
Shallow c la y range s i t e
A rtem isia t r id e n t a t a - B outeloua g r a c i l i s HT
H ir s c h :
Hansen e t a l . :
SCS:
A rtem isia t r id e n t a t a - B outeloua g r a c i l i s 9 x
A rtem isia t r id e n t a t a — Agropyron s m it h ii
Shallow c la y range s i t e
Andropogon s c o p a r iu s HT
H ir sc h :
Whitman:
Hansen e t a l . :
SCS:
Andropogon s c o p a r iu s 9
Andropogon - S tip a - B o u te lo u a •
Andropogon sc o p a r iu s - Carex f i l i f o l i a
Shallow range s i t e
Andropogon g e r a r d ii HT
H ir s c h :
Whitman:
Andropogon g e r a r d ii9
Andropogon - S tip a - Sporobolus
Agropyron s m it h ii - B outeloua g r a c i l i s - D i s t i c h l i s s p ic a t a
H ir sc h :
SCS:
Agropyron s m it h ii - B outeloua g r a c i l i s D i s t i c h l i s s p ic a t a v a r. s t r i c t a *
Thin c la y pon range s i t e
Grassed Sand F lo o d p la in s HT
SCS:
Sandy range s i t e 9
Hardwood Draws HT
Hansen e t a l . :
G irard :
F raxinus pennsyI v a n ic a - Prunus v ir g in ia n a and
Fraxinus p en n sy lv a n ica - Symphoricarpos
o c c id e n ta lis 9
(Has a s im ila r HT p rop osed )
Ju n ip eru s scopulorum - O ryzopsis m ircantha HT
Hansen e t a l . :
G ira rd :
Ju n ip eru s scopulorum - O ryzopsis m ircanth a9
(Has a s im ila r HT proposed)
Populus d e lt o i d e s - Ju n ip eru s scopulorum HT
G ir a r d :
P o p u lu s d e l t o i d e s -
J u n i p e r u s s c o p u lo r u m
16
Table 2 Continued
Pop u lu s trem u lo id es — B etu la o c c i d e n t a l is HT
Hansen e t a l . :
G irard:
Populus trem u lo id es - B etu la o c c id e n ta lis m
(Has a s im il a r 'HT proposed)
Marsh MU
SCS:
Wetland range s i t e 3
Brush MU
Hansen e t a l . :
Sym phoricarpos o c c i d e n t a l i s 3
( t h i s in c lu d e s o n ly a few
s i t e s , th e r e s t may have
been dom inated by Prunus
v i r g i n i a n a .. or Prunus
am ericana)
In trodu ced Grass MU
C la s s i f ie d by author
P r a ir ie Dog Towns MU
C la s s i f ie d by author
W illow s MU
C la s s i f ie d by author
Man-Managed MU
C la s s i f ie d by author
3D enotes the d e s c r ip t io n w hich was used in t h is study f o r th e h a b ita t
type or mapping u n it.
B e s id e s t h e 15 HT, s e v e n m apping u n i t s w e r e a l s o r e c o g n iz e d .
Mapping u n it s (MU) were d efin ed a s v e g e t a t io n a ssem b la g es which have
been d istu r b e d by n a tu ra l or an th rop ogen ic f o r c e s or w ere in th e e a r ly
s ta g e s o f s u c c e s s io n .
S e v e r a l MU had no co m p a ra b le d e s i g n a t i o n s by
th e p r e v io u s ly m entioned au th ors.
The c h a r a t e r is t ic s w hich were used
17
to d i f f e r e n t i a t e th e se MU w ere:
1 . INTRODUCED GRASSES MU - d istu r b e d a rea s r e p la n te d to
in tro d u ced g r a s s e s to r e d u c e e r o s io n h a z a r d s .
The
g r a s s e s m o st o f t e n u sed w e r e c r e s t e d w h e a t g r a s s
(.A gropyron c r i s t a t u m ) or sm ooth brome ( Bromus
jLnerm is),
2 . PRAIRIE DOG TOWNS MU - v e g e t a t i o n
a lte r e d by p r a ir ie dogs.
d r a s tic a lly
3 . RIVER BOTTOMS MU f lo o d in g .
to fr e q u e n t
areas
su b je c t
4 . WILLOWS MU - a r e a s d o m in a te d by d e n s e ly g r o w in g
w illo w ( S a lix spp.) w ith l i t t l e oth er p la n t growth.
5 . MAN-MANAGED MU - t h o s e la n d s whose v e g e t a t io n was
a l t e r e d by o n g o in g human a c t i v i t i e s ( r o a d s i d e s ,
campgrounds, e t c ) .
At s e v e r a l l o c a t i o n s i n b oth u n i t s ,
h a b i t a t t y p e s or m apping
u n i t s w ere so s m a ll or so in t e r m ix e d a s t o make i t
d i f f e r e n t i a t e s p e c i f i c a ssem b la g es.
c o m p le x e s .
c o m p le x w e r e
p la c e d
in to
to
These m osaics w ere . c l a s s i f i e d a s
T a b le 3 s h o w s t h e f o u r
c o n tr ib u tio n o f each HT and MU.
d iffic u lt
c o m p le x e s an d t h e p e r c e n t
Areas n ot c l a s s i f i e d a s a HT, MU, or
a se p a r a te
c a te g o r y :
u n v eg eta ted .
C o lle c t iv e l y th e HT, MU, and com p lexes w i l l be r e fe r r e d to a s h a b ita t
d iv is io n s .
A p p en d ix A sh o w s t h e a r e a o f ea ch HT, MU, and co m p le x a s i t
o c c u r s i n ea ch o f th e c l a s s e s .
M apping o f a l l c l a s s e s , .HT, MU, and
com p lexes was done w ith th e use o f c o l o r -in fr a r e d p h otos.
in te r p r e te d
Photos w ere
th ro u g h th e u se o f a p h o to e n h a n c in g co m p u ter and by
e x t e n s i v e ground t r u t h in g .
A ll c l a s s e s , HT, MU, and c o m p le x e s w e r e
p laced on a b ase map o f 1:12000 s c a le and d ig it iz e d .
18
Table 3•
L is t o f each v e g e t a t io n a l complex and th e p ro p o rtio n o f each
HT or MU th a t com prised th e com plex.
Achenbach H i l l s Complex .4
.3
.1 5 .
.05
. O5
.0 5
Agropyron s m it h ii - S tip a comata
S tip a comata - B ou telou a g r a c i l i s
Andropogon sc o p a r iu s
Andropogon s c o p a r iu s - J u n ip eru s h o r iz o n t a lis
Brush
Andropogon g e r a r d ii
P e t r if ie d F o re st Complex .4
.3
.1
.05
.1 5
Agropyron s m it h ii - S tip a comata
Agropyron s m it h ii - S tip a v ir id u la
Andropogon sco p a riu s
Andropogon g e r a r d ii
S tip a comata - B outeloua g r a c i l i s
R o llin g S c o r ia Complex .2 5
.2
.2
.15
.1
.1
Andropogon sco p a riu s
Agropyron s m it h ii - S tip a v ir id u la
S tip a comata - B outeloua g r a c i l i s
Agropyron s m it h ii - S tip a comata
Andropogon sc o p a r iu s - Ju n ip eru s h o r iz o n t a lis
Agropyron s m ith ii-B o u te lo u a g r a c i l i s - D i s t i c h l i s s t r i c t a
S teep S c o r ia Complex .3
.2 5
.15
.15
.1
.0 5
Andropogon sc o p a r iu s
Andropogan sco p a riu s - Ju n iperus h o r iz o n t a lis
Agropyron s m it h ii - S tip a v ir id u la
S tip a comata - B outeloua g r a c i l i s
' A rtem isia t r id e n t a t a - A tr ip le x c o n f e r t i f o l i a
Agropyron s m ith ii-B o u te lo u a g r a c i l i s - D i s t i c h l i s s t r i c t a
19
METHODS
Data w ere g a th ered i n two b a s ic c a t e g o r ie s : h a b it a t use by b iso n
and d is t r ib u t io n o f b iso n .
Data on h a b ita t use were c o lle c t e d by the
d ir e c t o b s e r v a tio n o f b iso n herds and
by th e use o f f e c a l tr a n s e c ts .
D ir e c t o b s e r v a tio n s were s ta r te d in J u ly 1982 and ended i n J u ly 1983.
No o b s e r v a tio n s w ere made from January to e a r ly March 1983. Observa­
t io n s averaged 8 hours per day and w ere taken betw een dusk and dawn.
During th e d ir e c t o b s e r v a tio n s th e f o llo w in g in fo r m a tio n was c o lle c t e d
a t each 15-m inute o b se r v a tio n p o in t:
1.
The l o c a t i o n o f th e h erd (r e c o r d e d on a I - 2 5 ,0 0 0
to p o g ra p h ic map).
2.
The number o f b is o n u s in g t h e c l a s s e s and h a b i t a t
d iv is i o n s .
3.
The number o f b is o n en g a g ed i n g r a z in g , r e s t i n g ,
and o th er a c t i v i t i e s w it h in th e c l a s s e s and h a b ita t
d i v i s i o n s . G ra zin g w as d e f in e d any a c t i v i t y t h a t
would end in a p la n t b ein g eate.ru R estin g was used
o n ly f o r b is o n r e c l i n i n g on t h e grou n d .
O ther
a c t i v i t y w as a n y t h in g t h a t w as n o t g r a z in g or
' r e s t i n g ; t h i s c o u ld in c l u d e s o c i a l i n t e r a c t i o n s ,
w a t e r in g , w a lk in g , r u n n in g , e t c . Each b is o n w as
c l a s s i f i e d a s d o in g o n ly one o f th e a c t i v i t i e s a t
any one o b se r v a tio n p o in t.
A t o t a l o f 5,281 o b se r v a tio n s w ere taken a t th e 15-m in u te ob ser­
v a tio n p o in ts which t r a n s la te d in t o 298,016 b ison m in u tes.
The hours
o f o b se r v a tio n w ere v a r ie d to prevent b ia s r e s u lt in g from o b se r v a tio n s
ta k e n d u r in g o n ly one p a r t o f th e day.
No n ig h t o b s e r v a t i o n s w ere
taken because o f th e d i f f i c u l t y and danger in v o lv e d , even though b iso n
w ere a c t iv e a t n ig h t.
Large h erd s were s e le c t e d fo r d ir e c t ob serva­
t i o n s w h e n e v e r p o s s i b l e b e c a u s e s m a ll h erd s c o n t a in e d o n ly a s m a ll
p o r t io n o f th e p o p u la t io n and c o u ld p o s s i b l y b i a s th e o b s e r v a t io n s
20
toward h a b it a t s used e x c e s s iv e ly by s m a ll herds.
A ll d ir e c t ob serva­
t i o n s w ere done i n t h e SU b e c a u s e o f t im e and t r a v e l c o n s t r a i n t s .
E leven d if f e r e n t tim e p erio d s w ere d e lin e a te d during th e study.
Table
4 g i v e s th e d a t e s o f ea ch p e r io d and how many o b s e r v a t io n d a y s w e r e
com p leted in each p eriod .
Table 4 .
Time p erio d s w ith s t a r t in g and ending d a te s a lo n g w ith
number o f d ir e c t o b se r v a tio n days in the p erio d .
Period
I.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
A p r il
May
June
August I
August 2
September I
September 2
October I
O ctober 2
November
December
End
S ta r t
3 /2 8 /8 3
5 / 1/83
6 /2 7 /8 3
8/ 3/82
8 /1 6 /8 2
9/ 1/82
9 /1 6 /8 2
10/ 2/82
10/ 16/82
1 1 /1 0 /8 2
12/ 4/82
to
to
to
to
to
to
to
to
to
to
to
4 /1 4 /8 3
5 /1 9 /8 3
7 / 7/8 3
8 /1 5 /8 2
8/ 31/82
9 /1 5 /8 2
9 /3 0 /8 2
1 0 /1 5 /8 2
10/ 28/82
11/ 26/82
12/ 15/82
th e
Number o f
O bservation Days
14
14
14
10
13
11
11
10
11
14
10
F eca l t r a n s e c t s c o n s is te d o f w a lk in g a t r a n s e c t 500 m eters lo n g
and I m eter w id e, w ith a l l r e c o g n iz a b le b iso n Chips counted w ith in th e
tr a n s e c t.
By running s e v e r a l t r a n s e c t s w ith in the same area t r a n s e c t s
o f t h i s le n g t h and w id t h a d e q u a t e ly sa m p led t h e d e n s i t y o f c h ip s
w ith in a 100 h e c ta r e area o f a c la s s .
T ra n sects were d is t r ib u t e d i n a
s t r a t i f i e d random m anner in b oth u n i t s .
The SU w as d iv id e d i n t o 9
b lo c k s and t h e NU i n t o 5 ■b lo c k s . W ith in ea ch b lo c k t r a n s e c t s w e r e
p la c e d ra n d o m ly i n e a c h o f th e c l a s s e s i n t h e b lo c k .
The number o f
t r a n s e c t s p la c e d i n t o ea ch c l a s s w as p r o p o r t io n a l t o th e a r e a i t
c o v e r e d i n t h e b lo c k .
A ll c l a s s e s had a minimum o f 3 t r a n s e c t s p er
21
u n it.
F eca l t r a n s e c t s w ere read d u rin g th e summer o f 1983.
A to ta l
o f 52 t r a n s e c t s w ere read in th e NU and 100 in th e SU (T able 5).
Home ra n g e and h erd a s s o c i a t i o n s w ere e s t a b lis h e d through the
use o f 4 c h a r a c t e r i s t i c a l l y marked a n im a ls.
v id u a lly r e c o g n iz a b le horn c o n fig u r a tio n s .
These a n im a ls had in d i­
L o ca tio n s o f th e se a n im a ls
w ere record ed whenever they were en cou n tered du rin g th e study.
S e v e r a l m eth od s o f s t a t i s t i c a l a n a l y s i s w e r e em p lo y ed i n t h e
study.
The C hi-square t e s t f o r independence was used to t e s t d i f f e r ­
e n c e s i n a v a i l a b i l i t y ( a r e a c o v e r e d by th e c l a s s ) and b is o n u se o f
d if f e r e n t c l a s s e s (Zarr 1984).
The C h i-s q u a r e t e s t f o r in d e p e n d e n c e
compared th e v a r io u s d is t r ib u t io n s o f b iso n to a random d is t r ib u t io n
( P o i s s o n ) , and com pared ea c h o f th e v a r i o u s d i s t r i b u t i o n s am on gst
t h e m s e lv e s . C o e f f i c i e n t s o f d i s p e r s i o n w ere c a lc u la te d fo r th e home
r a n g e s and v a r i o u s d i s t r i b u t i o n s o f b is o n b a sed on th e number o f
lo c a t i o n s per km2 ( Sokal and H ohlf 1981).
A c o e f f i c i e n t o f d is p e r s io n
o f I in d ic a t e s random d is t r ib u t io n , l e s s than I means th e d is t r ib u t io n
i s uniform , and g r e a te r than I means th e d is t r ib u t io n i s clumped.
A n a l y s is o f th e a c t i v i t i e s o f b is o n w as b a sed on th e p e r c e n t o f
th e herd engaged i n each a c t i v i t y a t th e 15-m in u te o b se r v a tio n p o in t.
A c t i v i t i e s w ere th e n a v e r a g e d o v e r a l l th e 15 - m in u t e o b s e r v a t io n
p o in ts w it h in each tim e p eriod.
Those a v era g es based on few er than 20
o b s e r v a t i o n s w ere c o n s id e r e d an in a d e q u a t e sa m p le and dropped fro m
fu r th e r a n a ly s is .
The r e g r e s s i o n m od el o f a n a l y s i s o f v a r ia n c e
(ANOVA) w ith unequal r e p l ic a t io n fo r a tw o -fa c to r ex p erim en t was used
t o t e s t f o r s i g n i f i c a n c e among t h e m ain e f f e c t s : e i t h e r th e c l a s s e s
and tim e p e r io d s , or th e h a b ita t d i v i s i o n s and tim e p erio d s (N eter and
22
W asserm an
1 9 7 4 ).
F -te sts
fo r
th e
m ain e f f e c t s
fo llo w e d
th is
eq u a tio n :
w h ere:
SSE(R)= E rro r sum o f s q u a r e s
the o th e r main e f f e c t .
f o r th e r e d u c e d m odel f o r
SSE(F)= Error sum o f sq u ares fo r th e f u l l model.
DF(R) = E rro r d e g r e e s o f fr e e d o m f o r th e r e d u c e d
fo r th e o th er main e f f e c t .
m od el
DF(F) = Error d eg rees o f freedom fo r th e f u l l model.
B e c a u s e o f th e u n eq u a l r e p l i c a t i o n no i n t e r a c t i o n e f f e c t s c o u ld be
a n a ly z e d , and i t w i l l be a ssu m ed t h e r e w ere none.
e f f e c t was s i g n i f i c a n t ,
I f o n ly one m ain
then th a t s i g n i f i c a n t e f f e c t was su b jected to
s e p a r a t i o n o f m eans a s a on e f a c t o r a n a l y s i s u s in g T ukey1s m ethod
(S o k a l and R o h lf 1 9 8 1 ).
I f b oth m ain e f f e c t s w e r e s i g n i f i c a n t , an
a d ju s tm e n t o f th e p e r c e n t s i n th e c l a s s e s and h a b i t a t d i v i s i o n s w as
done in th e form of:
Xi j -Xi =Z ij
w h ere:
x i j = The o r i g i n a l p e r c e n t o b s e r v a t io n i n e a c h c l a s s and
h a b ita t d iv is i o n .
X1 = The mean o f th e p e r c e n ts found in th a t tim e p erio d .
Zjj= The transform ed o b se r v a tio n .
The a d ju sted v a lu e s w ere then used in th e sep a ra tio n o f means by onef a c t o r a n a ly s is u sin g Tukey1S method.
The purpose o f th e ad ju stm en t
w as t o r e d u c e th e e f f e c t o f th e t im e p e r io d s on th e . v a l u e s i n th e
c l a s s e s and h a b i t a t d i v i s i o n s .
No s u i t a b l e form o f a d ju s tm e n t w as
23
found to reduce th e e f f e c t o f th e c l a s s e s and h a b it a t d iv is io n s 'o n th e
tim e p e r io d s , but they w ere g r a p h ic a lly r ep re se n ted so tren d s could be
ob served.
An a r c s in e tr a n sfo r m a tio n was done on a l l o f th e a c t i v i t y
p e r c e n ta g e s.
The d a ily d is ta n c e moved among th e tim e p erio d s was an alyzed by
a n a ly s is o f v a r ia n c e (Sokal and R olh f 1981).
A lo g a r it h im ic tr a n s fo r ­
m ation was done on th e d is ta n c e s because o f h e t e r o s c e d a s t ic it y and th e
f a c t th a t the standard e r r o r s were p r o p o rtio n a l to th e means.
The G-
t e s t (S o k a l and R o lh f 1 9 8 1 ) was u se d t o t e s t f o r in d e p e n d e n c e among
th e d is ta n c e s betw een two random p o in ts and the d is ta n c e s b iso n moved
from an o r i g i n a l l o c a t i o n .
The G - t e s t w as a l s o e m p lo y ed t o t e s t
in d e p e n d e n c e b e tw e e n th e d i s t a n c e s t h a t a random t r a n s e c t w as from
w a te r
com pared t o t h e d i s t a n c e from w a te r d u r in g a h e r d ’ s d a i l y
movement.
24
RESULTS AND DISCUSSION
D ir e c t O b servation s
The average herd s i z e in th e d ir e c t o b s e r v a tio n s was 71 an im als
(s ta n d a r d
error
(S E )= 25,
ran ge 2 5 -1 2 6 ).
The h e r d s
in
th e
Park
c o n s i s t e d o f a l l a g e s o f f e m a le s and m a le s up t o 4 y e a r s o f ag e.
M ature b u l l s w e r e fo u n d a s s o l i t a r y i n d i v i d u a l s or i n s m a ll g ro u p s.
The o n ly tim e when la r g e numbers o f m ature b u lls w ere found in herds
w as d u r in g t h e r u t , and t h i s w as th e o n ly tim e when d i r e c t o b s e r v a ­
t io n s in c lu d e d s i g n i f i c a n t numbers o f mature b u lls . Herds under ob ser­
v a tio n v a r ie d in s i z e betw een and w it h in days.
F eca l T ra n sects
The a v e r a g e number o f c h ip s p e r f e c a l t r a n s e c t and a s s o c i a t e d
standard erro r fo r each c l a s s appears in Table 5.
w ith in some o f the c l a s s e s e v id e n t ly r e f l e c t e d
The la r g e v a r ia tio n
th e d is p r o p o r t io n a t e
l e v e l s o f use among d if f e r e n t a rea s o f th e Park.
H ab itat Use
The p ercen t use o f each c l a s s in th e SU from both f e c a l t r a n s e c t s
and the d ir e c t o b s e r v a tio n s i s g iv e n in Table 6.
Both f e c a l tr a n s e c ts
and d ir e c t o b s e r v a tio n s gave com parable (r=.95) e s t im a t e s o f use fo r
the c l a s s e s ,
but th e methods did d i f f e r in what they w ere e s tim a tin g .
These d if f e r e n c e s w ere:
I.
The f e c a l t r a n s e c t s e s tim a te d th e accum ulated use
over two y e a r s or more, w h ile d ir e c t o b s e r v a tio n s
. e s tim a te d u se on ly during th e study p eriod .
25
2.
3.
The f e c a l t r a n s e c t s e s t i m a t e d u s e by th e w h o le
p o p u la t io n w h i le d i r e c t o b s e r v a t i o n s e s t i m a t e d
herd u se, ig n o r in g m ature b u ll use fo r most o f th e
year.
•
F e c a l t r a n s e c t s w e r e o n ly an in d e x o f u se w h i le
d ir e c t o b s e r v a tio n s w ere an a c tu a l measure o f use
(a t l e a s t du rin g th e d a y lig h t hours).
The h igh c o r r e la t io n betw een th e e s t im a t e s o f both methods su g g este d
th a t: I. use o f c l a s s e s in 1982-83 was s im il a r to th a t in th e p rev io u s
y e a r o r y e a r s , 2 . u se o f c l a s s e s by th e e n t i r e p o p u la t io n w as n o t
r a d i c a l l y d i f f e r e n t from th e u se e x h i b i t e d by th e h e r d s , and 3 . th e
f e c a l tr a n s e c t in d ex a r r iv e d a t the same r e l a t i v e use a s did th e r e a l
e s t im a t e s from the d ir e c t o b se r v a tio n s.
Table
5.
Number o f t r a n s e c t s and average number o f c h ip s per f e c a l
tr a n s e c t and standard e r r o r in the c l a s s e s f o r both u n it s .
C la ss
Breaks
Cottonwood f o r e s t s
Wooded draws
.Upland g r a s s la n d s
Old r iv e r te r r a c e s
G rassland bottom s
G rassland f l a t s
Toe s lo p e s
R o llin g g ra ssla n d s
Ridge & Ravine
S c o r ia h i l l s
Achenbach h i l l s
Sagebrush bottom s
P r a ir ie dog towns
tr a n se c ts
21
3
9
9
8
South Unit
c h ip s
SE
1.5
0.3
2.2
17.8
4 1 .9
—
11
8 .7
—
11
9 .6
15
8 .7
—
—
6 ;4
9
4 2 3 .5
2.0
0.6
1 .9
13.5
3 1 .2
tr a n se sts
North Unit
ch ip s
SE
9
6
5
4
1.0
2 .3
4 .0
3 3 .5
1 .2
1. 9
2.0
14.9
5
14.8
8 .9
5
5
3 1 .6
15.4
2 8 .3
9 .3
3
6
3
1.1.6
9 .5
5 0 .7
2.1
5 .6
51.1
7 .7
—
7 .9
5 .8
9 .2
1 2 .4
Table 7 compares the p ercen t a v a i l a b i l t y o f each c l a s s ,
i.e .
area o f co v era g e, and th e b iso n use e s tim a te d b y .f e c a l tr a n s e c ts .
th e
26
Table 6 .
P ercen t use o f c l a s s e s from d ir e c t o b s e r v a tio n s and f e c a l
t r a n s e c t s in th e South U n it.
C la ss
O b servation s
Breaks
Cottonwood f o r e s t s
Wooded draws
Upland g r a ssla n d s
Old r i v e r t e r r r a c e s
G rassland f l a t s
Ridge & Ravine
S c o r ia h i l l s
Sagebrush bottom s
P r a ir ie dog towns
1 .3
2 .9
1 .3
7 .2
• 3 5 .9
8.0
6.0
10.6
8.8
17.9 ■
Table
7.
1.2
0 .3
1.8
14 .7
3 4 .7
7 .2
7 .9
7 .2
5 .3
19.5
Average number o f b iso n f e c a l ch ip s per
a v a ila b ility
(p erce n t co v era g e) o f c l a s s e s .
found by u sin g C hi-square a n a ly s is .
South Unit
ch ip s
c la s s
% coverage
Breaks
Cottonwood f o r e s t s
Wooded draws
Upland g ra ssla n d
Old r iv e r te r r a c e
G rassland f l a t s
G rassland bottom s
Toe s lo p e s
R o llin g g r a ssla n d s
Achenbach h i l l s
Ridge and Ravine
S c o r ia h i l l s
Sagebrush bottom s
P r a ir ie dog town
A,B,C
T ra n sects
1 7 .3 A
0 .7 a
3 . Oa
5 .5 a
5 .8 a
1 1 .s c
1 9 .2 a
2 9 . 7A
5 .5 C
1 .5 a
1 .5 B
0 .3 b
2 . 2b
17. 8 b
41 .9 b
8 .7 C
9 .6 b
8 .7 B
6 .4 C
2 3 .5s
tr a n s e c t ’and
S ig n ific a n c e
North Unit
ch ip s
% coverage
3 7 .2 a
4 .6 C
3 .9 C
6 v3 a
I .Ob
2 .3 C
4 . oC
3 3 .5B
4 .5 a
19 . 1A
1 1 . 6°
1.-9A
1 4 .8 b
31 . 6 b
1 5 .4 °
1 1 .6 b
10. 3 A
0 .5 a
9 .5 b
5 0 .7 b
P a ir s w ith d if f e r e n t s u p e r s c r ip t s are s i g n i f i c a n t l y d if f e r e n t a t
p<0 . 0 1 .
Those c l a s s e s w ith use s i g n i f i c a n t l y d if f e r e n t from t h e ir a v a i l a b i l i t y
w ere id e n t i f i e d , u sin g th e C hi-square t e s t o f independence.
c a n t l y h ig h e r u se w as i n t e r p r e t e d a s
S ig n ifi­
p r e fe r a n c e fo r a c la s s ,
and
27
lo w e r u s e w as i n t e r p r e t e d a s a v o id a n c e o f a c l a s s .
w ere n e ith e r s tr o n g ly avoided nor p refer red .
S everal c la s s e s
The c l a s s e s w hich, were
p r e fe r r e d w ere f l a t and dom inated by g r a ssla n d s.
The c l a s s e s which
w ere avoided had e it h e r rugged topography or were dom inated by t r e e s
or shrubs.
Those c l a s s e s w ith no d e f in a t e p referen ce or aviodance had
c h a r a c t e r i s t i c s t h a t w e r e fo u n d i n b o th th e a v o id e d and p r e f e r r e d
c la s s e s .
The observed u se o f th e h a b ita t d iv is i o n s a lo n g w ith t h e ir a v a il­
a b i l i t y i s g iv e n i n T a b le 8.
o b s e r v a tio n s ,
no s t a t i s t i c a l
B e c a u se o f th e n o n -in d e p e n d e n c e o f t h e
t e s t s w e r e done.
T hose HT, MU, and
c o m p le x e s w it h u s e much h ig h e r th a n a v a i l a b i l t y w ere d o m in a ted by
g r a s s e s and o c c u r r e d on l e v e l or g e n t l y r o l l i n g s i t e s .
Those
w it h
lo w er u se were dom inated by woody v e g e t a t io n or w ere found on rugged
s ite s .
T hose i n w h ic h u se a p p r o x im a te d a v a i l a b i l i t y had ch a ra cte r­
i s t i c s o f both h ig h .u s e and low u se h a b ita t d iv is io n s .
The h ig h
p refer en ce
f o r g r a s s la n d s th r o u g h o u t t h e y e a r w as
p r o b a b ly t h e r e s u l t o f b is o n d i e t s b e in g com posed o f 87% g r a m in o id s
(Marlow e t a l. 1984), and b iso n b ein g la r g e a n im a ls w ith high a b so lu te
energy req u irem en ts
e t a l., 1 9 8 1 ).
and a to le r a n c e f o r low q u a lit y fo r a g e (Schwartz
In a d d i t i o n , b is o n had no n eed f o r p r o t e c t i v e c o v e r ,
e it h e r from en viron m en tal c o n d itio n s or from p red a to rs,
and fo r a g in g
i n th e w in te r i s e a s i e s t on g ra ssla n d s because o f th e lo w e r snow cover
(R e y n o ld s e.t a l . 1 9 8 2 ).
28
Table 8 .
P ercent use from th e d ir e c t o b s e r v a tio n s
and p ercen t
coverage o f h a b it a t ty p e s , ■ mapping u n it s , and com plexes in
th e South U n it.
HT,MU,& Complexes
%Use
^Coverage
Agropyron s m it h ii- S t ip a comata
Agropyron s m it h ii- S t ip a v ir id u la
S tip a com ata-B outeloua g r a c i l i s
A rtem isia cana
Andropogon sc o p a r iu s-J u n ip e r u s h o r iz o n t a lis
A rtem isia t r id e n t a t a - A tr ip le x c o n f e r t i f o l i a
A rtem isia tr id e n ta ta -B o u te lo u a g r a c i l l i s
Andropogon sc o p a r iu s
Hardwood draws
Ju n ip eru s scopulorum -O ryzopsis m ircantha
Populus d e lto id e s -J u n ip e r u s scopulorum
In trod u ced g r a s s e s
P r a ir ie dog town
.River bottom
Man-managed
S teep s c o r ia complex
R o llin g s c o r ia complex
P e t r if ie d f o r e s t complex
W atering area
U nvegetated a rea s & r iv e r
Brush
Populus trem u lo id es
2 2 .5
22.0
3 .9
6 .5
0 .9
2.2
0.2
4 .5
0 .4
4 .3
15.5
2.0
5 .5
3 .6
2 .3
0.6
10.1
2 .3
1.9
0 .5
1.0
1.0
0.5
0.2
1 3 .0
7 .8
0.1
—
0 .7
3 .9
17.8
2.8
2 .4
1.7
3 .1
0.1
0 .4
0.1
—
-
—
27 .8
0.1
0.1
S ea so n a l U se •o f H a b ita ts
The d i r e c t o b s e r v a t io n s a llo w e d s e a s o n a l u se t o be ex a m in ed i n
ea ch c l a s s and h a b i t a t d i v i s i o n .
Use o v e r t h e 11 d i f f e r e n t t im e
p erio d s fo r each c l a s s and h a b ita t d iv is i o n can be found in Appendix
B.
H ea v ily used c l a s s e s had w id e ly f lu c t u a t in g v a lu e s from p eriod to
p eriod w h ile th o se w ith l i t t l e use did n ot f lu c t u a t e a s w id e ly .
same w as tru e fo r th e h a b ita t d iv is io n s . Some o f the c l a s s e s ,
th e p r a ir ie dog tow ns and o ld r iv e r t e r r a c e s ,
p a tte r n to t h e ir u se.
The
such a s
had a d e f i n i t e sea so n a l
The p r a i r i e dog to w n s had h ig h u se d u r in g th e
summer and low use a t a l l oth er tim e s w h ile th e o ld r iv e r t e r r a c e s had
29
low u s e d u r in g t h e summer and h ig h u se a t o t h e r t im e s .
T h ere w e r e
o th er c l a s s e s ,
but showed
no s e t p a ttern .
such a s the s c o r ia h i l l s w hich flu c t u a t e d ,
Examples can a ls o be found i n th e h a b it a t d iv is io n s
o f s e a so n a l and ir r e g u la r use p a tte r n s.
P r a i r i e dog to w n s make up o n ly a s m a ll p a r t o f th e Park ( 1 .5 $ ) ,
b u t r e c e i v e d h ig h u se by b is o n .
T h is h ig h p r e f e r e n c e h a s b een n o te d
and i n v e s t i g a t e d by s e v e r a l o t h e r a u t h o r s (Popp I 981 , Coppock e t a l .
1 9 8 3 ).
Coppock e t a l . (1 9 8 3 ) r e p o r t e d t h a t p r e f e r e n c e i s t h e r e s u l t
o f p l a n t s on t h e e d g e s o f th e to w n s b e in g ch an ged p h y s i c a l l y and
g e n e t ic a lly by th e h erb iv o ry o f the p r a ir ie dogs.
Such changes cause
p l a n t s t o be m ore p r e f e r r e d and p a l a t a b l e t o b is o n .
c o u ld
be t h a t
a c tiv itie s
b is o n a r e drawn to p r a i r i e
(Popp 1 9 8 1 , S h u lt 1 9 7 2 ).
A n oth er r e a s o n
dog to w n s f o r r u t t i n g
Use o f p r a i r i e dog to w n s i n ­
c r e a s e d i n th e s tu d y from a lm o s t z e r o a t th e s t a r t o f th e r u t t o a
h igh l e v e l a t th e peak o f th e ru t and th en dropped back t o zer o a t th e
end o f th e r u t ( F ig u r e I 3 ).
Two o th er p refer red MU w ere th e r iv e r bottom s and man-managed.
D u r in g th e summer th e r i v e r b o tto m s becam e f a v o r i t e l o a f i n g s i t e s .
T h is w as b e c a u s e th e y p r o v id e e a s y a c c e s s t o w a te r f o r d r in k in g and
c o o l i n g - o f f , and b e in g n e a r ly d e v o id o f h e r b a c e o u s v e g e t a t i o n th e
r iv e r b ottom s became q u it e hot.
to f l i e s
T his produced c o n d itio n s u n favorab le
w h ic h w ere a n u is a n c e t o b is o n d u r in g th e summer.
managed, a r e a s may have b een p r e f e r r e d b e c a u se
m a n ip u la te d by m ow ing, w a t e r in g ,
Man-
t h e v e g e t a t i o n w as
and s e e d in g . . T h is m a n ip u la t io n
produces a p refer red and p a la ta b le fo ra g e.
30
I n d iv id u a l and Herd A s s o c ia tio n s
No lo n g -term a s s o c ia t io n s betw een th e four, i d e n t i f i a b l e cows w ere
ob served nor w ere in d iv id u a ls a s s o c ia t e d w ith s p e c i f i c h erd s.
In stea d
th e r e was o f t e n a d a y -to -d a y in terc h a n g e o f in d iv id u a ls w it h in herds.
At some p o in t during th e stud y every marked anim al was a s s o c ia t e d w ith
a l l o f th e o th e r marked an im a ls.
Home Range and D is t r ib u t io n
Home ran ges o f b iso n were d eterm ined fo r two o f th e fo u r charac­
t e r i s t i c a l l y marked a n im a ls (F ig u res 5 and 6).
s u r v iv in g b iso n appeared
The home range o f each
to encom pass th e w hole o f th e Park.
C ertain
a r e a s i n t h e Park had l o c a t i o n s clum p ed t o g e t h e r , i n d i c a t i n g b is o n
p r e fe r e n c e fo r c e r t a in c l a s s e s and h a b ita t d iv is io n s a t
th e se s i t e s .
A rea s d e v o id o f l o c a t i o n s w ere d o m in a ted , by a v o id e d c l a s s e s and
h a b ita t d iv is i o n s , la ck ed permanent w a ter, were not e a s i l y a c c e s s ib le
due to
rough te r r a in ,
and/or la ck ed s u it a b le fo ra g e.
C o e f f ic ie n t s o f d is p e r s io n e s t a b lis h e d f o r a n im a ls 0001 and 0002,
w ere 2.1 and 1.4, r e s p e c t iv e ly .
Both c o e f f i c i e n t s su g g e st th a t th ese
a n im a ls e x h ib ite d a clumped ra th er than a random or a uniform d i s t r i ­
b u tion .
T his clum ping was presum ably r e la t e d to th e p r e fe r e n c e s fo r
c l a s s e s and h a b ita t d iv is i o n s w hich w ere th em selv es not d is t r ib u t e d
e v e n ly over th e Park.
The o v e r a ll d is t r ib u t io n o f th e herds and b iso n in th e Park i s in
F igu re 7.
Both th e in d iv id u a l home ran ges and the o v e r a ll d is t r ib u ­
t i o n w e r e s i m i l a r i n t h a t l o c a t i o n s w e r e fo u n d from one end o f th e
SOUTH
F ig u r e 5 .
Home r a n g e o f m arked a n im a l 0001 d u r in g th e s t u d y .
UN
SOUTH
F ig u r e 6 .
Home r a n g e o f marked a n im a l 0002 d u r in g th e s t u d y .
UN
SOUTH
F ig u re 7
L o c a tio n s of a l l h e rd s observed d u rin g th e stu d y
UN
34
park t o t h e o t h e r .
Y et, t h e c o e f f i c i e n t o f d i s p e r s i o n 0.9 4 w as l e s s
than both th e in d iv id u a l home ra n g es, and anim al 0001 had a s i g n i f i ­
c a n t l y d i f f e r e n t (p < 0 .0 0 1 ) d i s t r i b u t i o n .
A p o s s ib le reason fo r th e
d i f f e r e n c e c o u ld be t h a t i n d i v i d u a l l y b is o n w ere s e l e c t i n g c e r t a i n
lo c a t i o n s fo r u se,
but c o l l e c t i v e l y th e r e was enough v a r ia t io n among
i n d i v i d u a l s t o p ro d u ce a more u n ifo r m d i s t r i b u t i o n .
A ll o f \ t h e s e
d is t r ib u t io n s s i g n i f i c a n t l y d if f e r e d from a random d is t r ib u t io n .
D ir e c t o b s e r v a tio n s recorded o n ly the d is t r ib u t io n o f herds.
The
f e c a l t r a n s e c t s on th e o th er hand d id reco rd m ature b u ll and herd u se.
In a r e a s w ith few i f any herd lo c a t i o n s ,
but w ith p refer red h a b it a t ,
u se w as r e c o r d e d i n t h e f e c a l t r a n s e c t s .
E v id e n t ly th e a r e a s t h a t
herds d id n ot freq u en t had some use by mature b u lls or by h erd s during
p rev io u s y e a r s.
Home ran ges o f mature b u lls w ere probably s im ila r to
or la r g e r than th o se o f cows s in c e m ost b iso n e sc a p in g from the Park
w ere m ature b u lls .
S ea so n a l D is t r ib u t io n
Each t im e p e r io d had a l l l o c a t i o n s p l o t t e d and a c o l l a p s i n g
p olygon p la ced around th e p erim eter p o in ts (Appendix C ).
Using o n ly
th e a r e a i n s i d e th e p o ly g o n a s th e e f f e c t i v e r a n g e f o r b is o n h e r d s
du rin g th a t tim e p erio d ,
a c o e f f i c i e n t o f d is p e r s a l fo r a l l lo c a t io n s
over th e tim e p e r io d s was, 0*84.
d i f f e r e n t from random.
A lo w c o e f f i c i e n t o f d i s p e r s i o n s u g g e s t s a
uniform d is t r ib u t io n o f lo c a t io n s .
an area to u se,
T his d is t r ib u t io n was s i g n i f i c a n t l y
A pparently,
once the b iso n chose
th e use occurred e v e n ly over i t .
D i s t r i b u t i o n o f th e b is o n h e r d s i n th e Park d id ch an ge o v e r th e
35
11 tim e p erio d s (Appendix C).
The most n o t ic a b le r e s t r i c t i o n in d is ­
t r ib u t io n o f b iso n h erd s was during th e e a r ly s p r in g when h erd s w ere
found e x c l u s i v e ly in th e n orth w est s e c t io n o f th e Park.
t io n li m it e d u se to b a s ic a ll y o n ly two c l a s s e s ,
and th e r id g e and r a v in e .
T his r e s t r i c ­
the o ld r iv e r te r r a c e
As la r g e r p a r ts o f th e Park w ere used b iso n
u se was spread over more c la s s e s .
A c t iv it y P a tte r n s
The a c t i v i t i e s
g r a z in g ,
r e s tin g ,
o f b is o n w ere d iv id e d i n t o
and o t h e r .
th r e e
c a te g o r ie s :
The s i g n i f i c a n c e o f th e m ain e f f e c t s
(e it h e r the c l a s s e s and th e tim e p e r io d s or th e h a b ita t d iv is i o n s and
th e tim e p e r io d s) on th e s e a c t i v i t i e s appears in T able 9 (Appendix D
c o n ta in s a l l ANOVA t a b le s used fo r th e F -v a lu e s).
T able 9 .
A c t iv it y
F -v a lu e s o f th e main e f f e c t s ,
th e c l a s s e s and tim e p erio d s
or h a b ita t d iv is i o n s and tim e p e r io d s , f o r the th ree
d if f e r e n t a c t i v i t i e s : g r a z in g , r e s t in g , and o th e r .
Main e f f e c t
C la ss X Time Period
c la ss e s
tim e p eriod
c la s s e s
time p eriod
c la ss e s
tim e p eriod
H a b ita t d iv is i o n X Time Period
g ra zin g
h a b ita t d iv is i o n
g ra zin g
tim e p eriod
r e s t in g .
h a b ita t d iv is i o n
r e s t in g
tim e p eriod
o th er
. h a b ita t d iv is io n
o th er
tim e p eriod
g ra zin g
g ra zin g
r e s t in g
r e s t in g
o th er
o th er
F -v a lu e
.DF
P -value
1.6
3 .4
3.1
1.1
3 .3
6.8
6,31
10,31
6,31
10,31
6 ,3 2
10,31
0.17
0.0 4
0. 01
0 .3 9
0. 01
0 .0 0
3.1
4 .9
2 .5
2.1
5 .2
8.0
9 ,3 6
10,36
9 ,3 6
10,36
9,3 6
10,36
0.01
0 .0 0
0.02
0.05
0.00
0 .0 0
36
T a b le 10 c o n t a i n s t h e s e p a r a t io n o f m eans f o r a l l t h r e e a c t i v i t i e s
among the c l a s s e s .
Table 11 c o n ta in s th e same f o r th e h a b ita t d i v i ­
s io n s .
Table 10.
Means1 o f th e average p ercen t o f a herd engaged in g ra zin g ,
r e s t in g , and o th er a c t i v i t y fo r each c l a s s .
grazing^
C lass
Sagebrush bottoms
Ridge & Ravine
Grassed f l a t s
Upland g ra ssla n d
Old r iv e r te r r a c e
S c o r ia h i l l s
P r a ir ie dog town
38. 4a
41 . 8 a
3 5 .3 a
30. 7 a
3 7 .5 a
34. Oa
3 4 . 8a
o th e r 2
r e s t in g
1 4 .7 a
17. 8A
22. 4a
2 7 .4 A
2 7 .5 a
2 8 .2 a
2 9 .4 A
4 6 .9A
4 0 .3 A
4 2 .3 ^
4 1 .8 ^
3 4 .4 ^
3 8 .8 AB
36 .2 B
1Means fo llo w e d by the same l e t t e r are n ot s i g n i f i c a n t l y d if f e r e n t a t
p>0 . 05.
2Means have been a d ju ste d to reduce th e a f f e c t o f th e tim e p erio d s.
^Grazing was n o t found to be s i g n i f i c a n t l y d if f e r e n t over the c l a s s e s .
Table 1 1 .
Means1 o f th e average p ercen t o f a herd engaged in g ra zin g ,
r e s t in g , and o th e r a c t i v i t i e s f o r each h a b ita t d iv is io n .
H a b ita t D iv is io n
S tip a com ata-B outeloua g r a c i l i s
In trodu ced g r a s s
P r a ir ie dog town
A rtem isia cana
Agropyron s m it h ii- S t ip a comata
Agropyron s m it h ii- S t ip a v ir id u la
R o llin g s c o r ia complex
Andropogon sc o p a r iu s
A rtem isia t r id e n t a t a A tr ip le x c o n f e r t i f o l i a
R iver bottom
g ra zin g 2
r e s t in g
o th e r 2
36. Oa
3 0 .9A
3 4 .7 ab
4 3 .2 B
3 3 .8A
3 7 .Sab
3 4 .7 a
4 2 .9 AB
3 4 .4 a
3 7 .3A
29.4a
2 6 . 3A
29 .6 A
2 4 . Oa
3 0 . 7A
21. 2a
2 9 .8A
3 1 . 8A
3 5 .8 A
3 0 .4 a
3 6 .6 A
3 8 . Oa
34 . 4 A
3 5 .9 a
3 8 . Gab
2 5 .6A
2 1 .9 a
1 7 .2 a
3 9 .5 AB
5 9 .5 B
1Means fo llo w e d by th e same l e t t e r are n ot s i g n i f i c a n t l y d if f e r e n t
p > 0 .0 5 .
2Means have been a d ju ste d to reduce th e e f f e c t o f tim e p e r io d s.
at
37
Trends i n a c t i v i t i e s l e v e l s over tim e can be examined i n Table
12 and Appendix E.
Table 12.
Means o f th e average p ercen t o f a herd engaged i n
a c t i v i t y i n the c l a s s e s over th e 11 time p e r io d s .
Time p eriod
means^
August I
August 2
September 2
September I
O ctober 2
November
June
A pril
O ctober I
December
May
2 7 .2 A
30.3AB
32.2AB
33.7AB
37.4AB
37.4AB
3 7 .5 AB
40.9AB
4 I . OAB
4 1 . SB
4 6 .9 B
g ra zin g
.
Means fo llo w e d by the same l e t t e r are n ot s i g n i f i c a n t l y d if f e r e n t a t
th e p>0 , 0 5 .
Grazing a c t i v i t y v a ried s i g n i f i c a n t l y among th e tim e p erio d s
both the c l a s s e s and th e h a b ita t d iv is i o n s .
in
Grazing was s i g n i f i c a n t l y
lo w e r d u rin g th e summer and h ig h er d u rin g th e w in te r and s p r in g (Table
12, and F ig u r e s 48 and 5 1 ).
The lo w summer g r a z in g a c t i v i t y c o u ld
have r e s u lt e d from lo n g , days which a llo w g ra zin g to be s tr e tc h e d out
over many d a y lig h t hours,
and
h igh d aytim e tem p era tu res which cou ld
have in h ib it e d d iu rn a l a c t i v i t y . Higher g ra zin g a c t i v i t y in th e w in te r
cou ld have been th e r e s u l t o f sh o r te r d a y lig h t hours g iv in g b iso n l e s s
t im e t o g r a z e i n and t h e l e a s t s t r e s s f u l e n v ir o n m e n ta l c o n d i t i o n s
o c c u r r in g i n th e d a y l i g h t h o u r s.
H ig h e r a c t i v i t y i n th e s p r in g may
h a v e b een a r e s u l t o f t h e b e s t e n v ir o n m e n t a l c o n d i t i o n s f o r g r a z in g
o c c u r r in g m o s t ly d u r in g th e d a y l i g h t h o u r s , and b is o n may have b een
38
r a i s i n g t h e i r in ta k e to m eet h ig h er p h y s io lo g ic a l needs fo r g e s t a t io n
and c a lv in g (R eynolds e t a l 1982).
R e s t in g a c t i v i t y
d id n o t v a r y s i g n i f i c a n t l y
p e r io d s ( F ig u r e s 49 and 5 2 ).
v a ria n c e o f r e s t in g a c t i v i t y
among t h e t im e
T h is can be a t t r i b u t e d
to th e w id e
among th e c l a s s e s and h a b ita t d iv is io n s
w ith in a tim e p eriod .
O ther a c t i v i t y v a r ie d s i g n i f i c a n t l y among t h e t im e p e r io d s i n
b o th th e c l a s s e s and h a b i t a t d i v i s i o n s .
O ther a c t i v i t y w a s h i g h e s t
d u rin g th e l a t e summer and lo w er in th e sp rin g , l a t e f a l l ,
(F ig u res 50 and 53).
and w in te r
Summer was h ig h because t h i s was th e tim e when
g r a z in g was lo w e s t and r u t t in g a c t i v i t i e s w ere occu rrin g.
G r a z in g a c t i v i t y d id n o t s i g n i f i c a n t l y v a r y among th e c l a s s e s ,
but d id v a r y s i g n i f i c a n t l y
among t h e h a b i t a t d i v i s i o n s .
H a b ita t
d iv is i o n s occurred i n much s m a lle r p a tch es than did c l a s s e s and w ere
e v e n ly d is t r ib u t e d throughout th e Park.
As b iso n t r a v e lle d du rin g a
day th e y d i f f e r e n t i a t e t h e i r a c t i v i t y among th e h a b i t a t d i v i s i o n s .
Because th e c l a s s e s o ccu p ied la r g e co n tig u o u s a rea s, t r a v e l to another
c l a s s was som etim es im p o s s ib le ,
so the b iso n ended up g ra zin g w here
th ey were.
R estin g and o th e r a c t i v i t y d id vary s i g n i f i c a n t l y among both th e
c l a s s e s and h a b i t a t d i v i s i o n s .
w id e ly ,
B e c a u s e r e s t i n g a c t i v i t y v a r ie d so
s i g n i f i c a n t d if f e r e n c e s w ere u n d e te c ta b le betw een th e l e v e l s
o f r e s t i n g a c t i v i t y among the c l a s s e s and h a b ita t d iv is io n s .
reason s fo r
th e d i f f e r e n t l e v e l s
. h a b ita t d iv is i o n s f o llo w :
o f a c tiv ity
Probable
i n t h e c l a s s e s and
39
1 . R iver b ottom s MU had h ig h er o th er a c t i v i t y because
b i s o n m a i n l y l o a f e d on t h e s i t e , a n d r u t t i n g
a c t i v i t y d id o c c u r on t h e s i t e .
Low g r a z i n g
a c t i v i t y w as due to th e lo w am ount o f a v a i l a b l e
fo r a g e . .
2 . A r te m isia cana HT had h ig h g ra z in g a c t i v i t y because
b is o n w o u ld f o r a g e i n th e HT b u t e v i d e n t l y a v o id e d
r e s t in g or d oin g o th er a c t i v i t y .
The avoidance was
p r o b a b ly r e l a t e d to th e h ig h d e n s i t y o f A r t e m is ia
cana which co u ld preven t v is u a l co n ta ct i n herds i n
t h e HT and c r e a t e m i c r o c l i m a t e c o n d i t i o n s
uncondusive to r e s t in g .
x
3 . In trodu ced g r a s s MU had low g ra zin g a c t i v i t y because
th e d o m in a n t .s p e c ie s A groovron c r i s t a t u m warn o n ly
p r e fe r r e d or p a la ta b le du rin g a sh o r t p eriod o f tim e
i n th e e a r ly sp rin g . With such low u se th e MU had a
la r g e buildup o f l i t t e r and sta n d in g dead m a te r ia l
w hich i s d e tr im e n ta l to g r a z in g a c t i v i t y (Norton e t
a l. 1982).
R estin g and o th e r a c t i v i t i e s w ere h ig h er
b e c a u s e o f t h e lo w g r a z i n g l e v e l a n d l a c k o f
u n fa v o ra b le c o n d itio n s fo r th e s e a c t i v i t i e s .
4 . S a g e b r u sh b o tto m s c l a s s had h ig h o t h e r a c t i v i t y
because i t in clu d ed th e r iv e r bottom s MU w hich had a
h ig h l e v e l o f o th er a c t i v i t y .
5 . P r a i r i e d og tow n c l a s s had lo w l e v e l s o f o th e r
a c t i v i t y becau se the c l a s s was fa v o red f o r r e s t in g
and g ra zin g .
D a ily A c t i v i t i e s
D a ily a c t i v i t y o f h erd s w ith in each tim e p erio d was an alyzed by
ta k in g th e p ercen t o f th e herd doing
p o in t and p lo t t in g th e se by
graphs
r e p r e se n tin g
each
each a c t i v i t y fo r an o b se r v a tio n
tim e o f day.
of
th e
Each p eriod then had th ree
th ree
a c tiv itie s
(A pp en d ix F ).
G razing a c t i v i t y i n the sp r in g peaked in th e morning and even in g,
s u b s t a n t ia l amounts o f g ra zin g a c t i v i t y
but
occurred throughout th e day.
As th e d a y s becam e lo n g e r th e p ea k s i n t h e m o rn in g and e v e n in g r e ­
m a in ed , but g r a z i n g d ro p p ed d u r in g t h e r e s t o f. th e day e x c e p t f o r a
40
peak a t midday.
The August 2 tim e p erio d b e st e x e m p lif ie s t h is .
The
f a l l and w in te r had the same p a tte r n a s th e sp rin g .
R estin g a c t i v i t y in th e sp r in g was co n cen tra ted i n th e m id d le o f
the day.
As days became lo n g er th e r e s t i n g a c t i v i t y spread ou t w ith
p ea k s o c c u r r in g d u r in g m id -m o r n in g and m id -a fte r n o o n .
The midday
drop i n r e s t in g a c t i v i t y was th e r e s u l t o f th e midday g ra zin g .
days grew s h o r te r th e r e s t in g a c t i v i t y
,As th e
again, was co n cen tra ted du rin g
th e m iddle o f the day.
O th er a c t i v i t y d id n o t e x h i b i t w e l l d e f in e d p e a k s . T here w as a
s l i g h t tren d tow ards h ig h e s t l e v e l s o c c u r r in g a f t e r th e m ajor g ra zin g
p e a k s.
The l e v e l o f o t h e r a c t i v i t y d id n o t n o t i c e a b l y ch an ge a s day
le n g th c h a n g es.
Movement
The movement p a tte r n s o f h erd s and in d iv id u a l b iso n were d e te r ­
m in ed from
a n im a ls .
d i r e c t - o b s e r v a t io n s o f h e r d s and l o c a t i o n s o f m arked
The a v e r a g e d i s t a n c e moved p er day f o r a h erd
(SE=I.5 8 ).
w a s 1.6 5 km
Table 13 show s th e average d is ta n c e moved by th e herds in
ea ch tim e p e r io d .
The d i s t a n c e a b is o n h erd moved d u r in g a day w as
t h e sam e f o r a l l p e r io d s (p>0.1 0) (T a b le 14) e v e n th o u g h th e p o r t io n
o f the Park used d id vary.
The number o f d a y s b e tw e e n l o c a t i o n s o f m arked b is o n w as n o te d
and the average d is ta n c e betw een th e i n i t i a l lo c a t i o n and r e lo c a t io n
fo u n d .
F ig u r e 8 c o n t a in s t h e a v e r a g e d i s t a n c e b e tw e e n t h e i n i t i a l
lo c a t i o n and su bseq u en t r e lo c a t io n f o r v a ry in g numbers o f days a f t e r
i n i t i a l lo c a t io n .
The average d is ta n c e moved by b iso n by th e second
41
day a f t e r a n : i n i t i a l lo c a t io n was 2 . 8 ' km..
Table 13.
Means o f the d a ily d is t a n c e s in k ilo m e t e r s ’ (km) . moved
b iso n herds in th e 11 d if f e r e n t tim e p e r io d s.
Time p eriod
•
A pril
May
June
August I
August 2
September I
September 2
October I
O ctober 2
November
December
Table 14.
By th e th ir d day a f t e r an
by
Mean d is ta n c e (km)
2.1
2.0
I .2
2 .5 .
2.2
1 .8
2.0
1 .4
1.1
1.1
0 .7
/
ANOVA-' ta b le o f th e lo g a r it h m ic a lly transform ed v a lu e s o f
' t h e • 'd a ily d is ta n c e moved by b iso n herds over ,the 11 tim e
p e r io d s .
Source
Time p erio d s
R esid u al
DF .
1.0
1.19
S .S .
MvS..
F - v a lu e ■
P -valu e
3.91
2 9 .6 3
.391
.249
1.57
.12
i n i t i a l l o c a t i o n ,' th e d i s t a n c e w as 4 km.
The c o n s t a n t m ovem ent by
b is o n i n th e f i r s t few d a y s a f t e r th e i n i t i a l l o c a t i o n r e s u l t e d i n
b is o n r a r e l y s t a y i n g i n t h e sam e l o c a l i t y
f o r m o r e . th a n 48 h o u rs.
B iso n , sh ow ed ho p r o p e n s it y to u se th e sam e s i t e ,
c la s s ,
or h a b i t a t
d iv is i o n fo r any extended p erio d o f tim e.
T h e .d ista n c e from th e i n i t i a l l o c a t i o n in c r e a se d a s the'num ber o f
d a y s b e tw e e n t h e i n i t i a l l o c a t i o n and r e l o c a t i o n in c r e a s e d .
w as r e a c h e d by th e I Oth day a t a d i s t a n c e o f 7.6 km.
A peak
The d i s t a n c e s
42
AVE.
DISTANCE(KM)
AND
SE
13"
14
NUMBER
OF
INITIAL
F i g u r e 8.
DAYS
20
30
RANDOM
FROM
LOCATION
A v e ra g e d i s t a n c e and s t a n d a r d e r r o r b e tw e e n t h e i n i t i a l
l o c a t i o n and r e l o c a t i o n o f m ark e d a n i m a l s a f t e r v a r i o u s
number o f d a y s , a l o n g w i t h t h e a v e r a g e d i s t a n c e and s t a n ­
d a r d e r r o r b e tw e e n two random p o i n t s .
th e n d e c l i n e d a f t e r t h e I Oth d ay, but n o t to th e l e v e l o f th e f i r s t 2
or 3 days a f t e r th e i n i t i a l lo c a t io n .
Table 15 shows th e r e s u l t s from
t e s t s o f independence betw een th e d is t a n c e s found f o r v a r io u s numbers
o f days a f t e r i n i t i a l lo c a t i o n to th a t o f the d is t a n c e s found betw een
two random ly g en era ted p o in ts .
No s i g n i f i c a n t d if f e r e n c e s w ere found
b e tw e e n th e d i s t a n c e s a t th e 7 t h ,
10th ,
1 4 th , and 2 0 th d a y s , a f t e r
i n i t i a l l o c a t i o n and th e random d is ta n c e s .
A pparently th e d is ta n c e s
moved by b iso n a f t e r th e second and th ir d w eeks were n o t in flu e n c e d by
w h e re t h e b is o n w e r e a w eek or tw o b e f o r e . T h is t r e n d may n o t h o ld
tr u e fo r d a te s beyond th e 30th day a f t e r i n i t i a l lo c a t io n because a t
th e 3 0 th day t h e d i s t a n c e s w e r e no lo n g e r random and b is o n w e r e
s t a r t in g to clump t h e ir lo c a t io n s around th e i n i t i a l lo c a tio n .
T able 15 .
G-t e s t s o f th e d i s t a n c e s •between two random p o in ts
r e lo c a t io n a number o f days l a t e r .
Days a f t e r
i n t i a l lo c a t i o n
5
7
10
14
20
30
G-s t a t i s t i c
P -value
22.1
11.9
5 .9
3 .5
4 .8
22.1
0. 00
0.1 0
0.66
0.8 9
0.84
0.0 0
a g a in s t
DF
7
8
7
8
9
7
The s h o r t e s t d is ta n c e to permanent w a ter was record ed fo r each
day a b iso n herd was observed.
Permanent w ater so u rces w ere d efin ed
a s a l l d evelop ed s p r in g s and th e L i t t l e
M issou ri R iver.
The average
d is ta n c e to permanent w a ter was I km (SE= 1.1 km) fo r th e study period.
The average d is ta n c e to' permanent w a ter f o r a randomly p laced tr a n s e c t
of
1.6 km w as 1.8 km (SE= 1.3 km ).
In a c o m p a r is o n
b e tw e e n t h e
44
observed d is t a n c e s to w a ter and th e random d is ta n c e s th e observed was
s i g n i f i c a n t l y d if f e r e n t (p<.05).
The o b s e r v e d d i s t a n c e s w ere c l o s e r
or more clumped around th e permanent w a ter so u rces,
b iso n were dependent on permanent w a ter so u rces.
s u g g e stin g th a t
B ecause most perma­
n e n t w a t e r s o u r c e s ' w e r e s i t u a t e d i n or around p r e f e r r e d h a b i t a t s ,
d eterm in in g w hether b iso n were s e l e c t i n g f o r w ater or fo r some o th er
a t t r ib u t e o f th e h a b ita t was not p o s s ib le .
Other o b s e r v a tio n s which
may supp ort th e argument th a t b iso n were not s e l e c t i n g fo r permanent
w a ter so u rces a re:
1. B ison w ere seen e a t in g snow when a v a ila b le and not
seen d rin k in g w a ter a t th o se tim e s.
2 . B ison , because o f t h e ir h igh m o b ilit y , were a b le to
u t i l i z e d if f e r e n t w ater so u r c e s each day.
• 3 . B ison used tem porary w ater so u rces when a v a ila b le .
4 . B iso n w ere observed g o in g w ith o u t w a ter fo r a t l e a s t
one day.
5 . B ison would go to w a ter once a day,and th e le n g th o f
s t a y a t w a t e r in g a r e a s w a s o f s h o r t d u r a t io n — one
hour or l e s s fo r even th e la r g e s t herd.
45
■CONCLUSIONS AND SUMMARY
Many a s p e c ts o f b iso n h a b ita t use and d is t r ib u t io n found in t h is
stu d y have been rep o rted in o th er s t u d ie s .
These being:
1. The p r e fe r e n c e fo r g r a ssla n d s.
2 . The home ran ges b ein g la r g e and s i g n i f i c a n t l y over­
la p p in g w ith th e r e s t o f the p o p u la tio n .
3« The s o c ia l s tr u c tu r e b ein g f l u i d w ith no lo n g term
in d iv id u a l or herd a s s o c ia t io n .
. ■•
4 . The m a tu re. b u lls u sin g a r e a s not n orm ally freq u en ted
by h e r d s .
5 . The co n sta n t movement o f b iso n in t o new areas.
6 . The s e a s o n a l c h a n g e s i n d i s t r i b u t i o n and h a b i t a t
u se .
The a v e r a g e d a i l y d i s t a n c e moved i n t h i s , s t u d y w as lo w e r th a n t h e
d i s t a n c e s r e p o r t e d i n th e o th e r s t u d i e s .
T h is may have been due t o
th e m eth o d s o f m e a s u r in g t h e d i s t a n c e s moved a n d /o r d i f f e r e n c e s
b e tw e e n stu d y a r e a s .
H a b ita t u se by b u l l s w as o n ly r e c o r d e d i n th e
th e w h ole p o p u la tio n h a b ita t use. w hich a ls o in clu d ed th e cow herd u se.
Because th e w hole p o p u la tio n h a b ita t u se was s im ila r to the cow herd
h a b ita t u se,
th ere w as probably l i t t l e
when compared to cow herd use.
in b u ll and cow herd u se.
d if f e r e n c e in o v e r a ll b u ll use
The o th er
s t u d ie s found d if f e r e n c e s
I t i s p o s s ib le th a t i f mature b u ll use had
been measured more in t e n s iv e l y more d if f e r e n c e s m ight have been found.
The d a ily a c t i v i t y p a tte r n s o f b iso n i n .t h e stu d y were s im ila r to
t h o s e r e c o r d e d fo r. many o t h e r u n g u la t e s (L e u th o ld I 9 7 7 , M lo s z e w s k i
I 9 8 3 ).
The b is o n had a d e f i n i t e c r e p u s c u la r b e h a v io r p a t t e r n w it h
g ra zin g b ein g the d e t e r m in is t ic fa c to r .
46
g r a z in g b ein g the d e t e r m in is t ic f a c t o r .
The b eh avior o f b iso n in th e Park was a com b in ation o f nomadism
and th e p r e f e r e n c e f o r c e r t a i n co m p o n en ts o f th e e n v ir o n m e n t.
The
nomadic b eh avior o f th e b iso n was based on:
1. B ison b ein g h ig h ly m o b ile , moving to new l o c a l i t i e s
and h a b it a t s a lm o st d a ily .
2 . The d i s t a n c e s moved by b is o n a p p a r e n t ly n o t b e in g
in f l u e n c e d by w here th e y w ere l o c a t e d t h e w eek
b e fo r e .
. 3* The d is t r ib u t io n o f b iso n herds resem b lin g a random
p a tte r n .
4 . The uniform d is t r ib u t io n o f lo c a t io n s over th e area
used du rin g a s in g le tim e period.
5 . Home r a n g e s o f i n d i v i d u a l b is o n b e in g a s l a r g e a s
th a t fo r th e w hole p o p u la tio n .
6 . The c o n s t a n t l y c h a n g in g a s s o c i a t i o n s b e tw e e n
i n d i v i d u a l s and th e la c k o f f i d e l i t y t o any one
herd.
7 . The g ra zin g a c t i v i t y not b ein g s i g n i f i c a n t l y d i f f e r ­
en t over the c l a s s e s .
8 . The b is o n 's a b i l i t y
so u r c e s each day.
to
u se
d iffe r e n t
w a t e r in g
The p r e f e r e n t ia l b ehavior o f th e b iso n was based on:
1. The b iso n 's p r e fe r e n c e f o r c e r t a i n h a b i t a t s .
2 . The ch an ge i n u se o f h a b i t a t s and a r e a s o f t h e park
as changes in the environm ent occurred.
3 . The clumped lo c a t io n s f o r in d iv id u a l b ison .
4 . The s i g n i f i c a n t d i f f e r e n c e s i n a c t i v i t y p a t t e r n s
among h a b it a t components.
5 . The clumped lo c a t io n s o f h erd s around w ater so u rces.
Both nomadism and th e a c t io n o f p refer en ce had a d a p tiv e s i g n i f i ­
c a n c e f o r th e b is o n .
Nomadism i s a d a p t iv e i n t h a t i t
a llo w e d th e
47
b is o n t o cop e w it h th e f l u c t u a t i n g f o r a g e p r o d u c t io n fou n d on t h e
g r a s s la n d s .
B i so n r e q u ir e l a r g e am o u n ts o f f o r a g e t o s u s t a i n t h e i r
la r g e s i z e and energy needs.
Because g ra ssla n d s vary w id e ly in fo r a g e
p r o d u c t io n b e tw e e n l o c a t i o n s and y e a r s , a n om ad ic b is o n w as m ore
c a p a b le o f f i n d i n g t h o s e a r e a s o f fa v o r a b le fo ra g e p rod u ction w hich
would s a t i s f y i t s fo r a g e needs.
The consequence o f nomadism,
e s p e c ia l ly w ith co n sta n t movement
and random d i s p e r s a l o v e r t im e , w a s t h a t p l a n t s w ere p o t e n t i a l l y
g r a z e d o n ly o n c e , i f a t a l l , i n a 3 - 4 w eek p e r io d .
th e r e l a t i o n s h i p
New t h e o r i e s on
b e tw e e n h e r b iv o r y and p la n t p h y s i o lo g y ,
( Grunqw 1 980) s u g g e s t
th a t
p la n ts a re
b e tte r
a b le
to
e c o lo g y
cope w it h
h erb iv o ry when i t occurred o n ly on ce, ra th er than o ccu rr in g a s s e v e r a l
r e g r a z in g s d u r in g t h e g r o w in g s e a s o n .
G r a ssla n d p l a n t s th e n a r e
b e t t e r a b le to cope w ith h erb iv o ry by a nomadic anim al ra th er than one
w hich i s sed en ta ry and r e g r a z e s th e same p la n t r e g u la r ily .
P re feren ce fo r c e r t a in en viron m en tal f a c t o r s i s a d a p tiv e b ecau se,
e v e n th o u g h b is o n w e r e n o m a d ic, no a n im a l can a f f o r d t o l e a v e a l l
a s p e c ts o f i t s b eh avior to chance and ex p ect to s u r v iv e (A lcock 1984).
B iso n w ere w e ll adapted to a gram i n i od d ie t ,
and so a s th ey t r a v e lle d
th e y c h o s e f o r t h o s e a r e a s d o m in a ted by g r a m in io d s , th e g r a s s la n d s .
P r e f e r e n c e f o r o t h e r h a b i t a t s w a s b a se d on t h e h a b i t a t s b e in g w e l l
s u it e d to th e o th er n o n -g ra zin g a c t i v i t i e s o f b ison.
Together th e se b eh a v io rs r e s u l t i n th e b iso n b ein g w e ll s u ite d t o
the g r a ssla n d environm ent.
o th er la r g e ,
These b eh a v io ra l t r a i t s w ere a ls o found in
h erd in g b o v id s,
e s p e c ia l ly th e A frica n b u ffa lo (Svncerus
c a f f e r ) (L euthold 1977, Owen-Smith 1982, M loszew ski 1983). • I t appears
48
th a t la r g e b ovid s in an environm ent dom inated by la r g e g r a ssla n d s w i l l
d ev elo p th e se b eh a v io rs,
even when th e g r a ssla n d s and a n im a ls' have a
d if f e r e n t e v o lu tio n a r y h is to r y .
LITERATURE CITED
50
LITERATURE CITED
A lcock, John. 1984. Animal b ehavior: an e v o lu tio n a r y approach. Sinauer
A s s o c ia te s I n c . , Sunderland, Mass.
C oppock, D.L., J.E. E l l i s , J.K. D e t l i n g , and M.I. D yer. 1 9 8 3 . P l a n t h e r ib v o r e i n t e r a c t i o n s i n a N o rth A m erica n m ix - g r a s s p r a i r i e : I I .
R e s p o n s e s o f b is o n t o m o d i f i c a t i o n o f v e g e t a t i o n by p r a i r i e d o g s.
O eco lo g ia 5 6 :1 0 -1 5 .
D ary, D.A. 1 9 7 4 . The b u f f a l o book: t h e sa g a o f an A m erica n sy m b o l.
Avon Books/Sw allow P r e ss, Chicago.
G arretson , M.S. 1938. The American b iso n . New York Z o o lo g ic a l S o c ie ty .
Grunow, J.O. 1980. P r i n c i p l e s and p r a t i c e s o f c o n t r o l l e d s e l e c t i v e
g r a z in g . Multicamp symposium. Maze ls p o o r t , South A fr ic a .
H ansen, P. L., R. B. H o p k in s, and G. R. H offm an. 198 0. An e c o l o g i c a l
s tu d y o f th e h a b i t a t t y p e s and t h e i r a n im a l co m p o n en ts a t T h eod ore
R o o se v e lt N a tio n a l Park, North Dakota. U n iv e r s ity o f South Dakota.
H ollan d er, M, and D.A. W olfe. 1973. N onparam etric s t a t i s t i c a l methods.
John W iley & Sons, New York.
L a rso n , F lo y d . 1 940. The r o l e o f th e b is o n i n m a in t a in in g t h e s h o r t
g r a s s p la in 's. E cology 2 1 :1 1 3 -1 2 1 .
L e u th o ld , W a lte r . 1 9 7 7 . A f r ic a n u n g u la t e s : a c o m p a r a tiv e r e v ie w o f
t h e i r e t h o lo g y and b e h a v io r a l e c o l o g y . S p r in g e r - V e r la g B e r l i n ,
H e id elb erg , Germany.
L o t t , D. F. and S. C. M int a. I 983. Random i n d i v i d u a l a s s o c i a t i o n and
s o c i a l . grou p i n s t a b i l i t y i n A m erica n B is o n ( B is o n B is o n ). Z.
T ie r p s y c h o l 61:153-172.
M arlow , 0. B. j L. R. I r b y i and J. E. N o rla n d . 1 984. Optimum c a r r y in g
c a p a c ity f o r b iso n i n Theodore R o o se v e lt N a tio n a l Park. Montana S ta te
U n iv e r s ity .
I
M cD onald, J e r r y N .'1 9 8 1 . N orth A m erican b is o n . U niv. o f C a l i f o r n i a
P r e ss, B erk eley and L .A ., C a lifo r n ia .
Malin , James C. 1956. The g ra ssla n d o f North America. Edwards B roth ers
I n c . , Ann Arbor, M ichigan.
McHugh, Tom. 1958. S o c ia l behavior
4 3 :1 -5 4 .
o f th e American B u ffa lo . Z o o lo g ica
51
M eagher, Mary M. I 973. The b is o n o f Y e llo w s t o n e N a t io n a l Park. Mat.
Park S e r v . S c i. Monogr. S e r ie s I.
M lo sz e w s k i , Mark J. I 9 8 3 . The b e h a v io r and e c o l o g y o f th e A f r ic a n
b u ffa lo . Cambridge U n iv e r s ity P r e ss, New York.
M eter,J . a n d .W. W asserinan.1 974. A p p lie d l i n e a r s t a t i s t i c a l
Richard D. Irw in In c ., Homewood, I l l i n i o s .
m o d e ls.
N o rto n , B.E., P.S. J o h n so n , and M.K. Owens. 1 982. I n c r e a s in g g r a z in g
e f f i c i e n c y . U t. Agr. Exper. S t a t io n 4 3 :1 1 0 -1 1 3 .
Owe n - S m ith , N. I 98 2 . F a c t o r s i n f l u e n c i n g t h e c o n s u m p tio n o f p la n t
p r o d u c ts by l a r g e h e r b iv o r s In : B.J. H u n tle y and. B.H. W alker (e d ),
E c o lo g y o f t r o p i c a l sa v a n n a s. S p r in g - V e r la g , B e r l i n , H e id e lb e r g ,
Germany.
P eden, D.G,., G.M. Van D yne, R.W. R ic e , and R.M. H ansen. 1 974. The
t r o p h ic e c o l o g y o f B is o n b is o n L. on s h o r t g r a s s p l a i n s . J. A ppl.
E e o l. 1 1 :4 8 9 -4 9 8 .
Popp, J e w e l Kay. 1 9 8 1 . Range e c o l o g y o f b is o n on m ix ed g r a s s p r a i r i e
a t Wind Cave N a tio n a l Park. M.S. t h e s i s , Iowa S t a t e U n iv e r s ity , Ames,
Iowa.
R eynolds, H.W., R.D. G lah olt* and A.W.L. Hawley. 1982. Chap. 49 B ison ,
i n J.A. Chapman and G.A. F eld h a m er ( e d ) , W ild mammals o f N o rth
A m erica : b i o l o g y , m anagem ent, and e c o n o m ic s . The John H op k in s
U n iv e r s ity P r e ss, B a ltim o re, Maryland.
Roe, F.G. 1970; The N orth A m erican b u f f a l o : a c r i t i c a l stu d y o f t h e
s p e c i e s i n i t s w i l d s t a t e . U n i v e r s i t y o f T o ro n to P r e s s , T o ro n to ,
Canada.
S c h w a r tz , C. C. and J. E. E l l i s . 1 9 8 1 . F e e d in g e c o l o g y and n ic h e
se p a r a tio n in some n a tiv e and d o m estic u n g u la tes on the s h o r tg r a s s
p r a ir ie . J. o f A pplied E cology 18:343-353.
S h u lt , M.J. 1972.. A m erican b is o n b e h a v io r p a t t e r n s a t Wind Cave
N a tio n a l Park. Ph.D. t h e s i s , Iowa S ta te U n iv e r s ity , Ames, Iowa.
S o k a l, R. R. and F. J. R o h lf . 1981. B io m e tr y . W.H. F reem an and
San F ra n cisco .
C o.,
Soper, J.D. 1941. H istory,-, range, and home l i f e o f th e northern b ison .
E co l. Mongr. 11:3 4 9 -4 1 2 .
Weaver, J.W. and F.E. C lem ents.
C o ., New York.
1938. P lan t eco lo g y .
McGraw-Hill Book
Whitman, Warren Ci 1978. A n a ly sis o f g ra ssla n d v e g e t a t io n on s e le c t e d
k ey a r e a s i n s o u t h w e s t e r n N orth D akota.: N orth D ak ota R e g io n a l
.Environm ental Program.
Van V uren, D irk ; 1 979. E c o lo g y and b e h a v io r o f b is o n i n t h e Henry
M ountains,' Utah. M.S. t h e s i s , Oregon S ta te -U n iv , C o r v a llis , Oregon.
Van Vuren, Dirk. 1983. Group dynam ics and summer home range o f b iso n
in sou th ern Utah. J. Mammalogy 6 4 :3 2 9 -3 3 2 .
Zarr,. J ero ld H. 1984. B i o s t a t i s t i c a l a n a ly s is . ’ P r e n tic e -H a ll Inc. New
Jersey.
APPENDICES
54
APPENDIX A
. Area Covered by th e H a b ita t D iv is io n s
55
Table 16.
H ecterage o f h a b ita t ty p e s , mapping u n it s , and com plexes as
they o ccu r i n each p h y s io g r a p h ic /v e g e ta tio n a l c l a s s i n the
North U n it.
Breaks
1.
2.
3.
4.
5.
6.
7.
8.
S tip a comata - B outeloua g r a c i l i s
Hardwood Draws
Populus trem u lo id es - B etu la o c c i d e n t a l i s .
Andropogon sc o p a r iu s - J u n ip eru s h o r iz o n t a lis
Juniperus scopulorum - O ryzopsis m ircantha
A rtem isia t r id e n t a t a - A tr ip le x c o n f e r t i f o l i a
Andropogon sc o p a r iu s
U nvegetated
2 .4
3. 5
10.1
7. 6
24 2 .4
123.6
Q.3
25 1 9 .7
Sagebrush Bottom
1.
2.
3.
4.
5.
6.
7.
8.
9 •.
Hardwood Draws
A rtem isia cana
R iver Bottom
W illow s
Marsh
Man-Managed
Brush
Grassed Sand F lo o d p la in s
U nvegetated
. 6 .3
7 4 8 .7
'16.9
3 0 .8
18.8
7. 0
1. 3
7 2 .8
15 . 1
Wooded Draws
In clu d ed in th e o th er c l a s s e s a s Hardwood Draw and
Populus trem u lo id es - B etu la o c c i d e n t a l is
Upland G rasslands
I.
2.
3.
4.
5.
6.
7.
8.
9.
Agropyron s m it h ii - S tip a comata
Agropyron s m it h ii - S tip a v ir id u l a
S tip a comata - B outeloua g r a c i l i s
Hardwood Draw
Andropogon sc o p a r iu s - Ju n iperus h o r iz o n t a lis
A rtem isia t r id e n t a t a - A tr ip le x c o n fe r t i f o l i a
Andropogon sc o p a r iu s
Brush
U nvegetated
172.1
115.1
9 7 .6
0.6
7 .8
1.9
15.7
0. 2
9.1
P r a ir ie Dog Town
I.
P r a ir ie Dog Town
48.1
56
T a b le 16.
C o n tin u e d
Toe S lo p es
T.
2.
3.
4.
5.
6.
7•
8.
910.
11.
12.
Agropyron s m ith iiS tip a comata
Agropyron s m it h ii - S tip a v ir id u l a
.
.
S tip a comata - B outeloua g r a c i l i s
Hardwood Draw
Populus -trem uloides
B etu la o c c id e n t a lis
A rtem isia cana
Andropogon sc o p a r iu s - Juniperus h o r iz o n t a lis
J u n ip eru s scopulorum - O ryzop sis m ircantha
A rtem isia- t r id e n t a t a — A tr ip le x c o n f e r t i f o l i a
Andropogon sc o p a r iu s
Brush
U nvegetated
122.7
609. 6
13.7
311 .2
13.7
6 .3
1 3 5 .0
116.0
3 1 0 .0
266.1
3 .7
3 8 0 .4
R o llin g G rassland
Agropyron s m it h ii - S tip a comata
Agropyron s m it h ii - S tip a v ir id u l a
' ■
• 3 . S tip a comata-r- B outeloua g r a c i l i s
.. 4 . Hardwood Draw
■ 5 . Po p u lu s trem u lh id es - B etu la o c c id e n t a lis
• 6 .: .Andropogon sc o p a r iu s - Ju n ip eru s h o r iz o n t a lis
- ■> 7 . A rtem isia t r id e n t a t a - A tr ip le x c o n f e r t i f o l i a
8 . Andropogon sc o p a r iu s
9 . In trodu ced G rasses
Brush
-" 10 .
1 1 . U nvegetated
1.
2.
2 2 .7
7 3 7 .9
7 .5
51.9
0.8
26.6
5 .3
4 8 .6
14.1
9 .0
76.8
, Achenbach H i l l s
■ I.
2.
Achenbach H i l l s Complex
; Popul u s . trem ul o id e s ^ .B etu la o c c i d e n t a l is
77.6
1 .4
Bottom G rasslands
1 . Agropyron s m it h ii - S tip a v ir id u l a
,.
2 . Hardwood Draw
3 . : Juniperus s c o p u l o r u m O ryzopsis m ircantha ..
• ■ 4 . Brush
■:
■ 5 . Grassed Sand. F lo o d p la in s
6 . U nvegetated
210.0
0. 1
0.2
1 .5
60.7
5 .0
Cottonwood F o r e sts
I.
. Populus d e lt o id e s - Juniperus scopulorum
88.6
57
Table 17.
H ecterage o f h a b ita t ty p e s , mapping u n it s , and com plexes a s
they occu r i n each p h y sio g r a p h ic /v e g e ta tio n a l c l a s s i n the
South U n it.
Breaks
1.
2.
3.
4.
5.
6.
7.
8.
9•
10.
11.
12.
13•
14.
15.
Agropyron sm ith i i - S tip a comata
Agropyron s m it h ii - S tip a v ir id u l a
S tip a comata - Bouteloua g r a c i l i s
Hardwood Draw
Populub trem u lo id es - B etu la o c c id e n t a lis
Andropdgon sc o p a r iu s - J u n ip eru s h o r iz o n t a lis
Juniperus scopulorum - O ryzopsis m ircantha
A rtem isia t r id e n t a t a - A tr ip le x c o n f e r t i f o l i a
Andropogon sc o p a r iu s
Man-Managed
A rtem isia t r id e n t a t a - B outeloua g r a c i l i s
S teep S c o r ia
R o llin g S c o r ia
Brush .
U nvegetated
8 .4
3 0 .4
12.3
9 3 .4
1 .0
7 2 .8
23 4 .6
'3 0 2 .7
134.2
6.8
63.5
3 0 .4
3. 5
1. 9
2701.2
Sagebrush Bottom s
1.
2.
3.
4.
5.
6.
7.
Agropyrori s m it h ii - S tip a v ir id u l a
Hardwood Draws
A rtem isia cana
Andropogon sc o p a r iu s
R iver Bottom
Man-Managed
U nvegetated
4 6 .9
14.5
978.9
1.6
93. 2
18.6
35. 6
Wooded Draws
In clu d ed in th e o th er c l a s s e s a s Hardwood Draws.
Upland G rassland
1.
2.
3.
4.
5.
6.
7.
8.
, 9.
Agropyron s m it h ii - S tip a comata
Agropyron s m it h ii- S tip a v ir id u l a
S tip a comata - Bouteloua g r a c i l i s
Hardwood Draw
A rtem isia t r id e n t a t a - A tr ip le x c o n fe r t i f o l i a
Andropogon sc o p a r iu s
In trodu ced G rasses
Brush
U nvegetated
P r a ir ie Dog Towns
I.
P r a ir ie Dog Town
7 7 .2
513.5
192.3
0 .3
0 .7 ■
25. 7
155.7
2 .6
4 2 .6
168.4
58
T a b le 17.
C o n tin u e d
Old R iver T errace
/-■ I . •
2. .
. . 3 .'
4^
5.
6;
7.
8.
9«
Agropyron s m it h ii - S tip a comata
Agropyron s m it h ii - S tip a v ir id u l a
S tip a comata - Bouteldua g r a c i l i s
Hardwood Draw
Andropogon sc o p a r iu s
Introduced G rasses
Man-Managed
P e t r if ie d F o re st Complex
U nvegetated
60 8 .6
1 8 4 .0
159.4
2 .9
7. 7
2 5 .9
5 .2
2 2 .0
G rassland F la t s
I.
Agropyron s m it h ii - S tip a comata
■2.
Agropyron s m it h ii - S tip a v ir id u l a
3 . Hardwood Draw
4 . A rtem isia cana
■5,
Andropogon sc o p a r iu s - Ju n ip eru s h o r iz o n t a lis
6 . J u n ip eru s scopulorum - O ryzop sis m ircantha
■7.
A rtem isia t r i d e n t a t a .- A tr ip le x c o n f e r t i f o l i a
■8 ; • . Andropogon s c o p a r iu s
9 . In trodu ced G rasses
10.
. Man-Managed
11.. A rtem isia t r id e n t a t a - B outeloua g r a c i l i s
12. R o llin g S c o r ia Complex
13. Brush
14.. U nvegetated
2 .3
1347. 3
13.8
I3. 6
7.1
.5 .4
9 ,4
47. 4
2 .9
2 .6
3 2 .7
12.8
2. 9
462.6
Ridge & Ravine
1.
2.
3 ..
■ 4.
‘5 .
6.
7.
8.
9.
10.
11.
.1 2 .
13.
14.
15.
Agropyron s m it h ii -■ S tip a comata
Agropyron s m it h ii - S t i p a v ir id u l a
S tip a comata 4 B outeloua g r a c i l i s
■Hardwood Draw
..
Populus trem u lo id es - B e tu la. o c c id e n t a lis
A rtem isia cana
Andropogon sc o p a r iu s - Ju n iperus h o r iz o n t a lis
J u n ip eru s scopulorum - O ryzop sis m ircantha
A rtem isia, t r id e n t a t a - A tr ip le x c o n f e r t i f o l i a
Andropogon sc o p a r iu s
Man-Managed
S teep S co ria Complex
R o llin g S c o r ia Complex
Brush
U nvegetated
103.6
37 2 .3
14.5
276. O
2 .4
21.0
58 9 .8
77 .5
9 4 .3
1652.8
I.3
12.4
18.2
4.4
1489.2
59
T a b le 17.
C o n tin u e d
S c o r ia H i l l s
I.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
Agropyron sm ith i i - S tip a v ir id u l a
Hardwood Draws
A rtem isia cana
Andropogon scd p a riu s - J u n ip eru s h o r iz o n t a lis
Juniperus scopulorum - O ryzop sis m ircantha
A rtem isia t r id e n t a t a - A tr ip le x c o n f e r t i f o l i a
Andropogon scd p a riu s
A rtem isia t r id e n t a t a - B ou telou a g r a c i l i s
S teep S c o r ia Complex
R o llin g S c o r ia Complex
U nvegetated
8 1 .7
23.1
8 .3
4 .6
6 3 .0
3 5 .7
10.6
14 .2
2360.9
140 6 .0
524.4
Cottonwood F o r e sts
I.
Populus d e lt o id e s - Juniperus scopulorum
88.6
APPENDIX B
. P ercen t Use by B iso n o f t h e .C la s s e s and H ab itat D iv is io n s
.Figures show ing th e . p ercen t use .determ ined from d ir e c t o b s e r v a tio n s o f
b is o n i n ea ch c l a s s and h a b it a t d i v i s i o n by tim e p e r io d (T a b le I ).
.N o n -p a r a m e tr ic c o n f id e n c e i n t e r v a l s are: shown (H ollander and W olfe
1973).
d en o tes l i m i t s o f non-param etric'..confidence in t e r v a l
61
TI ME
Cottonwood f o r e s t s c l a s s
USE
F igure 9.
PERI OD
T I ME PERI OD
F igu re 10.
R id ge & R a v in e c l a s s
62
TIME
Breaks c l a s s .
USE
Figure 11.
PERIOD
TIME
F igu re 12.
S co ria h i l l s
cla ss
PERIOD
63
TIME
P r a ir ie dog town c l a s s .
USF
Figure 13.
PERIOD
T I ME P E R I O D
F i g u r e 14 .
Wooded draw c l a s s .
64
Figure 15.
G rassland f l a t s c l a s s .
TIME
F igu re 16.
PERIOD
S a g e b r u s h b o tto m s c l a s s
65
TIME
Old r iv e r te r r a c e c l a s s .
USE
F igure 17.
PERIOD
TIME
F igure 18.
PERIOD
Upland g r a ssla n d c l a s s .
F igure 19.
Figure 2 0 .
TIME
PERIOD
TIME
PERIOD
Man-managed MU.
A rtem isia t r id e n t a t a - A tr ip le x c o n f e r t i f o l i a .
7
USE
67
Sn
21.
PERIOD
S tip a comata - B outeloua g r a c i l i s HT.
USF
rn
TIME
HME
F ig u r e 22.
S t e e p s c o r i a c o m p le x .
PERI OD
% USE
68
23.
PERIOD
U nvegetated a rea .
USF
*
TIME
TIME
F ig u re 2 4 .
A rtem isia t r id e n t a t a -
PERIOD
B o u t e l o u a g r a c i l i s HT.
69
TIME
Figure 25.
PERIOD
Introduced g r a s s e s MU.
70
TIME
R o llin g s c o r ia com plex.
USE
F igure 27.
PERIOD
TIME
F ig u re 2 8 .
Hardwood draw HT.
PERIOD
% USE
71
29.
PERIOD
Andropogon sc o p a r iu s - Ju n ip eru s h o r iz o n t a l is HT
USE
m
TIME
T I ME P E R I O D
F ig u re 3 0 .
R i v e r b o tto m s MU.
72
TIME
Figure 31.
PERIOD
P r a ir ie dog town MU.
TI ME P E R I O D
F ig u re 3 2 .
A r t e m i s i a can a HT.
USE
73
TIME
Figure 33.
Agropyron s m it h ii - S tip a v ir id u la HT.
TIME
F igu re 3 4 .
PERIOD
W a te r in g a r e a MU.
PERIOD
74
S-
LU
CO
=)
-I
4-
-f-
TIME
Populus d e lt o i d e s - Ju n ip eru s scopulorum HT.
TIME
Figure 36 .
4=
PERIOD
USE
F igure 35.
4-
PERIOD
Andropogon sco p a riu s HT.
75
■APPENDIX C
D a ily Movement o f Herds in Each Time Period
F ig u r e s o f th e d a i l y m ovem ent o f a l l h e r d s o b s e r v e d f o r ea ch tim e
p erio d .
- d en o tes s t a r t in g lo c a t io n . L e tte r s d en ote en d in g lo c a ­
tio n .
Those d a i l y m ovem en ts e n d in g i n t h e sam e l e t t e r w ere h e r d s
fo llo w e d in c o n s e c u tiv e days.
C
SOUTH
UNI T
N
F ig u re 37.
A p ril.
A
SOUTH
UNI T
N
D
X. .
SOUTH
UNI T
-A
/
B
F ig u re 39.
June.
SOUTH
UNI T
B ''
Va
F ig u re 41.
A u g u st 2 .
B1
SOUTH
F ig u re 42.
S ep tem b er I .
UNIT
SOUTH
UNI T
O
I
OD
r\;
F ig u re 4 3 .
S ep tem b e r 2 .
C
SOUTH
UNI T
CD
LU
F ig u re 4 4 .
O c to b e r I .
SOUTH
UNI T
N-
F ig u re 45.
O c to b e r 2 .
SOUTH
UNI T
N-
A
F ig u re 46.
November.
SOUTH
UNIT
N
-
CO
O'
)
F
F ig u re 47.
D ecem ber.
87
APPENDIX D
A c t iv it y A n a ly sis o f V ariance T ables
ANOVA t a b le s from th e a n a ly s is o f g ra zin g , r e s t in g , and o th e r a c t i v i t y
among th e c l a s s e s and tim e p e r io d s or th e h a b ita t d iv is i o n s and tim e
p e r io d s .
88
T a b le 1 8 .
ANOVA t a b l e s o f g r a z i n g a c t i v i t y among t h e c l a s s e s and t i m e
p erio d s.
F u ll model
S ource'
r e g r e s s io n
r e s id u a l
to ta l .
DF
16
31
47 ,
Reduced model o f th e c l a s s e s
Source
M
r e g r e s s io n
10
r e s id u a l
37
to ta l
47
SS
0 .1 8 0
0 .1 2 0
0.301
MS
0.0113
0. 0038
SS
0.1 4 2
0.158
0.301
ES
0.0142
0.0042
Reduced model o f th e tim e o e r io d s
Source
DF
SS
r e g r e s s io n
6
0. 047
r e s id u a l
41
0.253
to ta l
0.301
47. :
Table 19.
MS
0. 0079
0. 0061
ANOVA t a b le s o f r e s t in g a c t i v i t y among th e c l a s s e s and tim e
p erio d s
F u ll model
Source
r e g r e s s io n
r e s id u a l
to ta l
DF
16
31
47
Reduced model o f th e c l a s s e s
Source
DF
r e g r e s s io n
10
r e s id u a l
37
to ta l •
47
SS
0.436
0. 516 .
0 .9 5 2
MS
0.0212
0.0166
SS
0.131
0.821
0.952
MS
0. 0131
0.0221
Reduced model o f th e .time p erio d s
Source
DF ■
SS
r e g r e s s io n
6
0.252 .
r e s id u a l
41
0 .7 0 0
to ta l
0.952
47
MS
0. 0420
0. 0170
89.
T a b le 20.
ANOVA • t a b l e s o f
p erio d s.
F u ll model
Source
r e g r e s s io n
r e s id u a l
to ta l
DF
16
31
47
Reduced model o f th e c l a s s e s
Source
M
■
r e g r e s s io n
10
r e s id u a l
37
to ta l
47
o th er
a c tiv ity
among t h e c l a s s e s and t i m e
SS
0.341
0.117
0.4 5 8
MS
0.0213
0.0037
SS
0.265
0.193
0 .4 5 8
ES
0.0265
0.0052
Reduced model o f th e tim e p erio d s
Source
SS
M
r e g r e s s io n
6
0.084
r e s id u a l
41
0.374
to ta l
0.458
47
MS
0 .0 1 4 0
0. 0091
T a b le ■21. ■' -ANOVA t a b l e s o f g r a z in g a c t i v i t y
d iv is i o n s and tim e p erio d s.
F u ll model
Source
r e g r e s s io n
r e s id u a l ■
to ta l
DF
19
36
55
SS
0.253
0.109
0.362
MS
0.0133
0. 0030
Reduced model o f th e h a b ita t d iv is i o n s
Source
DF
SS
r e g r e s s io n
10
0.1 7 0
r e s id u a l
45
0.192
to ta l
55
0.362
MS
0. 0170
0. 0042
Reduced model o f th e tim e p erio d s
Source
DF
SS
r e g r e s s io n
9
0.103
r e s id u a l
46
0.2 5 9
t o t a l.
. .
0.362
55
MS
0. 0115
0.0056
among th e
h a b ita t
90
T a b le
22.
ANOVA'- t a b l e s o f r e s t i n g a c t i v i t y
d i v i s i o n s and tim e p e r i o d s .
F u ll model
Source
r e g r e s s io n
r e s id u a l
to ta l
DF
19
36
55
SS
0 .3 6 0
0.296
0.656
th e
h a b ita t
MS
0.0189
0.0082
Reduced model o f th e h a b ita t d iv is i o n s
Source
DF
SS
r e g r e s s io n
10
0.172
r e s id u a l
45
0.484
to ta l
55
0.656
IlS
0.0172
0.0107
.Reduced model o f th e tim e n erio d s
S ource.
DF
SS
r e g r e s s io n
9
0.186
r e s id u a l
46 .
0 .4 7 0
to ta l
55 . 0.656
MS
0.0207
0. 01 02
Table 23.
■ ■ -
am ong
ANOVA t a b le s o f o th er a c t i v i t y among th e h a b ita t d iv is i o n s
and :time p e r io d s.
F u ll model
Source
r e g r e s s io n
r e s id u a l
to ta l
DF
19
36
55
SS
0 .4 0 0
0.086
0.487
MS
0. 0211
0.0241
Reduced model o f th e h a b ita t d iv is io n
Source
DF
SS
r e g r e s s io n
10
0.287
r e s id u a l
45
0.2 0 0
to ta l■
55
0.487
MS
0.0287
0.0044
Reduced model o f th e tim e o erio d s
Source
DF
SS
■ regressio n
9.
0.207.
r e s id u a l
46 . .
0.2 8 0
to ta l ■
55
0.487
MS
0 .0230
0.0060
appendix
E
A v erage P e r c e n t o f t h e H erd s E ngaged i n th e
D i f f e r e n t A c t i v i t i e s Among t h e Time P e r i o d s
ACTI VI TY
AVE• % OF HERD ENGAGED I N
VO
rv
5 TlME6 PERIOD
F i g u r e 48.
A v e r a g e p e r c e n t o f o b s e r v e d h e r d s en g a g e d i n g r a z i n g a c t i v i t y among
t h e c l a s s e s i n e a c h tim e p e r i o d .
IN
% OF HERD ENGAGED
VO
AVE.
LaJ
5
6
7
8
TI ME PERIOD
F ig u re 49.
A v e r a g e p e r c e n t o f o b s e r v e d h e r d s e n g a g ed i n r e s t i n g a c t i v i t y among
t h e c l a s s e s i n e a c h tim e p e r i o d .
IN A C T I V I T Y
A
A
A
A
f
A
2▲
A
A
A
A
▲
A
A
▲
A
A
2
t
*
A
A
A
2-
A
A
S-
A
A
i
A
A
A
A
A
A
AVE.
% OF HERD ENGAGED
S-
5
F ig u re 50.
6
7
TI ME PERIOD
I
I
10
A v e r a g e p e r c e n t o f o b s e r v e d h e r d s en g a g e d i n o t h e r a c t i v i t y among
t h e c l a s s e s i n e a c h tim e p e r i o d .
% OF HERD ENGAGED I N A C T I V I T Y
AVE.
TI ME PERIOD
F ig u re 51.
A v e r a g e p e r c e n t o f o b s e r v e d h e r d s en g a g e d i n g r a z i n g a c t i v i t y among
t h e h a b i t a t d i v i s i o n s i n e a c h tim e p e r i o d .
AVE• % OF HERD ENGAGED IN A CT I V I T Y
A
4
I
A
A
A
A
A
A
A
I
A
A
I
A
A
▲
A
A
A
A
A
A
A
A
*
A
4
A
A
▲
A
A
A
A
A
A
A
A
A
A
A
T I ME PER7I OD
F i g u r e 52.
A v e r a g e p e r c e n t o f o b s e r v e d . h e r d s e n g a g ed i n r e s t i n g a c t i v i t y among
t h e h a b i t a t d i v i s i o n s i n e a c h tim e p e r i o d .
OV
I N ACT I V I T Y
AVE - % OF HERD ENGAGED
5 Tl ME PERI OD
F ig u re 53.
A v e r a g e p e r c e n t o f o b s e r v e d h e r d s en g a g e d i n o t h e r a c t i v i t y among
t h e h a b i t a t d i v i s i o n s i n e a c h tim e p e r i o d .
APPENDIX F
Percent', o f th e Herds Engaged i n the D if f e r e n t A c t i v i t i e s
During th e D a y lig h t Hours fo r Each Time Period
F ig u r e s sh o w in g th e p e r c e n t o f o b s e r v e d h e r d s en g a g ed in g r a z in g ;
r e s t in g , or o th er a c t iv it y , during d a y lig h t hours fo r each tim e period.
- 100
OF
ACTIVITY
I OOH
IN
HERD
ENGAGED
ENGAGED
OF
ACTIVITY
HERD
IN
4 : 00
6:00
8:00
10:00
I 2:00
14: 00
HOURS
F ig u re 54.
G r a z in g a c t i v i t y f o r A p r i l
I 6:00
I 8:00
20:00
2 2:00
IOCH
OF
IN A C T I V I T Y
-100
OF
IN A C T I V I T Y
HERD
ENGAGED
ENGAGED
HERD
4 : 00
6:00
8:00
10:00
12:00
14:00
HOURS
F ig u re 55.
R e stin g a c t i v i t y f o r A p r il.
I 6:00
18: 00
20: 00
22:00
OF
IN A C T I V I T Y
-100
OF
IN A C T I V I T Y
HERD
ENGAGED
ENGAGED
HERD
4: 00
8:00
10:00
12:00
14: 00
HOURS
F ig u re 56.
O th er a c t i v i t y f o r A p r i l
I 6:0 0
I 8:00
20:00
22:00
IOOH
OF
IN A C T I V I T Y
-100
HERD
IN
HERD
ENGAGED
ENGAGED
OF
ACTIVITY
4:00
6:00
8:00
10:00
I 2:00
14:00
HOURS
F i g u r e 57
G r a z in g a c t i v i t y f o r May
I 6:00
18: 00
20:00
22:00
IOOi
- 100
OF
HERD
ENGAGED
IN
ACTIVITY
4 : 00
6:00
8:00
10:00
1 2 :0 0
14:00
HOURS
F i g u r e 5 8.
R e s t i n g a c t i v i t y f o r May.
16:00
18:00
20:00
22:00
100
-100
90-90
>
>
80-
H
O
<
-80
•
•
"
70-
Q
LU
CD
<
CD
Z
LU
60•
•
••
•
•• •
30-
•:‘.v
20-
• :•
• • •• a
•
• .%
. • •• • N
•:
•%
••
y • • •
• • . .
v
••
••
• #
.-V
\ ••
• •• .
. %
••
6:00
8:00
I 0:00
I
I
12:00
•
. -
••
••
••v
%
••
-40
14:00
16:00
-30
>
O
H
-20
<
H
•
-<
/
-10
18: 00
CD
>
CD
m
O
Z
I---------- 1
-------
HOURS
O t h e r a c t i v i t y f o r May
^
• •
•
••
. .
•
10-
o -L__
A"
•
: .
•
-50
s-
•••
I
m
33
O
m
Z
•
40-
F i g u r e 5 9.
-60
*
•
4:00
•
O
n
20: 00
22:00
104
X
u.
O
a?
•
•
e *
50-
Q
CE
LU
-70
•
Z
as
ACTIVITY
OF
IN
HERD
ENGAGED
ENGAGED
OF
ACTIVITY
HERD
IN
4:00
6:00
8:00
10:00
I 2:00
14:00
HOURS
F ig u re 60.
G r a z in g a c t i v i t y f o r June
I 6:00
18:00
20:00
22:00
100-1
- 100
OF
HERD
ENGAGED
IN A C T I V I T Y
4:00
6:00
8:00
10:00
I 2:00
14:00
HOURS
F ig u re 61.
R e stin g a c t i v i t y fo r June.
16:00
I 8:00
20: 00
22: 00
ACTIVITY
OF
IN
HERD
ENGAGED
ENGAGED
IN
HERD
OF
ACTIVITY
4:00
6:00
8:00
10:00
12:00
14:00
HOURS
F ig u re 62.
O th er a c t i v i t y f o r Ju n e
I 6 :0 0
I 8:00
20:00
22:00
-100
OF
HERD
ENGAGED
IN A C T I V I T Y
8:00
12:00
14:00
HOURS
F ig u r e 63.
G r a z in g a c t i v i t y f o r A u g u st I .
16:00
I 8:00
20:00
22:00
-100
OF
<
50-
ENGAGED
70-
HERD
o
IN
ACTIVITY
30-
6:00
8:00
10:00
12:00
14:00
HOURS
F ig u re 64.
R e s t i n g a c t i v i t y f o r A u g u st I
16:00
I 8:00
20:00
22:00
ACTIVITY
OF
IN
HERD
ENGAGED
ENGAGED
OF
ACTIVITY
HERD
IN
4:00
6:00
10:00
12:00
I 4:00
HOURS
F ig u r e 65.
O th er a c t i v i t y f o r A u g u st I
20:00
22:00
IOCH
-100
OF
HERD
ENGAGED
IN
ACTIVITY
6:00
8:00
10:00
12:00
14:00
HOURS
F ig u re 66.
G r a z in g a c t i v i t y f o r A u g u st 2 .
I 6:00
18:00
20:00
22:00
- 100
OF
HERD
ENGAGED
IN A C T I V I T Y
4:00
6:00
8:00
10:00
I 2:00
14:00
HOURS
F ig u re 67.
R e s t i n g a c t i v i t y f o r A u g u st 2 .
I 6:00
18:00
2 0 : 00
2 2: 00
-100
OF
ACTIVITY
1 OOH
IN
HERD
ENGAGED
ENGAGED
OF
HERD
IN A C T I V I T Y
4:00
6:00
8:00
1 0: 0 0
12:00
14:00
HOURS
F ig u re 68.
O th er a c t i v i t y f o r A u g u st 2 .
16:00
18:00
20:00
22:00
ACTIVITY
OF
IN
HERD
ENGAGED
ENGAGED
OF
ACTIVITY
HERD
IN
4:00
8:00
10:00
1 2 :00
14:00
HOURS
F ig u re 69.
G r a z in g a c t i v i t y f o r S e p tem b e r I
16:00
18:00
20:00
22:00
-100
OF
ACTIVITY
100-1
IN
HERD
ENGAGED
ENGAGED
OF
HERD
IN A C T I V I T Y
4:00
6:00
8:00
10:00
12:00
14:00
HOURS
F ig u r e 70.
R e s t i n g a c t i v i t y f o r S ep tem b er I .
16:00
18 :00
20:00
22:00
ACTIVITY
OF
OF
IN A C T I V I T Y
HERD
ENGAGED
ENGAGED
IN
HERD
4:00
6:00
8:00
10:00
12:00
14:00
HOURS
F ig u re 71.
O th er a c t i v i t y f o r S ep tem b er I .
I 6:00
18: 00
2 0 : 00
22: 00
ACTIVITY
OF
IN
HERD
ENGAGED
ENGAGED
OF
ACTIVITY
HERD
IN
4:00
6:00
8 :00
10:00
I 2:00
14:00
HOURS
F ig u re 72.
G r a z in g a c t i v i t y f o r S ep tem b er 2,
I 6:00
1 8:00
20:00
22:00
I OCH
OF
ACTIVITY
-100
IN
HERD
ENGAGED
ENGAGED
OF
HERD
IN A C T I V I T Y
4:00
6:00
8:00
10:00
1 2: 0 0
14:00
HOURS
F ig u re 73.
R e s t i n g a c t i v i t y f o r S ep tem b er 2
I 6:00
18:00
20:00
22: 00
ENGAGED
IN
ACTIVITY
ENGAGED
HERD
IN
ACTIVITY
HERD
OF
OF
OF
ACTIVITY
-100
IN
HERD
ENGAGED
ENGAGED
OF
HERD
IN A C T I V I T Y
4:00
6:00
8:00
10:00
12:00
14:00
HOURS
F ig u re 75.
G r a z in g a c t i v i t y
f o r O c to b e r I .
1 6:00
1 8:00
20:00
22:00
- 10 0
OF
ACTIVITY
IOCH
IN
HERD
ENGAGED
ENGAGED
OF
ACTIVITY
HERD
IN
4:00
6:00
8:00
10:00
1 2: 0 0
I 4:00
HOURS
F ig u re 76.
R e s t i n g a c t i v i t y f o r O c to b e r I .
I 6:00
I 8:00
20:00
22:00
IN A C T I V I T Y
OF
IN
HERD
ENGAGED
ENGAGED
HERD
OF
ACTIVITY
4:00
6:00
8:00
10:00
I 2:00
14:00
HOURS
F ig u re 77.
O th er a c t i v i t y f o r O c to b e r I
I 6:00
I 8:00
20:00
22:00
I OCH
OF
IN A C T I V I T Y
-100
OF
IN A C T I V I T Y
HERD
ENGAGED
ENGAGED
HERD
4:00
6:00
8 :0 0
10:00
I 2:00
14:00
HOURS
F ig u re 78.
G r a z in g a c t i v i t y f o r O c to b e r 2 .
I 6:0 0
I 8:00
20:00
22:00
OF
-100
9 0-
-90
8 0-
-80
O
n
x
-70 m
x
□
-60 m
70
60
z
Q
50
-50
40
••
-40
• .
: •
100-
I
4:00
T
__ J-O
6:00
8:00
10:00
12:00
14
HOURS
F ig u re 79.
o
z
>
-30 O
—I
<
-20
H
-<
-10
30'
20-
>
O
m
R e s t i n g a c t i v i t y f o r O c to b e r 2 .
16:00
18:00
20:00
2 2:0 0
124
HERD
ENGAGED
IN
ACTIVITY
I OO-I
I OOH
OF
ACTIVITY
-100
IN
HERD
ENGAGED
ENGAGED
OF
HERD
IN A C T I V I T Y
4:00
6 :0 0
8:00
10:00
I 2:0 0
14:00
HOURS
F ig u re 80.
O th er a c t i v i t y f o r O c to b e r 2
I 6:00
18:00
20:00
22:00
-100
IOOi
70-
IN
..
•
30
20
r
6 :0 0
8:00
• / ••••
• •
• •
•
10:00
81.
* •
• e•
-60
-50
I'
.x
'.s*.
V
X
*
v-
-30
*
»
.-v
• .
• •
••• . • %
• •.
*
*.
-20
*•
-10
___> 0
I
I 2:00
i 14:00
HOURS
Figure
0
G r a z in g a c t i v i t y f o r November.
' 16:00
22:00
IN A C T I V I T Y
40-
i
S
XX.S
•
• r:. X -:
". •
:
v
•
• •• •
•
•\•
•
.
•
• .y
I
I
50-
4:00
•
••
ENGAGED
ENGAGED
HERD
HERD
h70
80-
60-
OF
80
OF
ACTIVITY
90
- 10 0
6?
80-
80
IN
f-70
HERD
50-
-50
40-
-40
I\
30
.
-30
.. .
V
20-
1004:00
-20
. •.
••
••
\
|
\
.
-10
*
■
_____ ____________ _____ _____ _____ _________________ _____ _____ ____________ J - O
6:00
8:00
10:00
12:00
14:00
HOURS
F ig u re 82.
R e s t i n g a c t i v i t y f o r November.
16:00
18:00
20:00
'
22:00
IN A C T I V I T Y
OF
60
ENGAGED
ENGAGED
60
HERD
70-
OF
ACTIVITY
I OOH
I OO-I
- 10 0
OF
HERD
ENGAGED
IN A C T I V I T Y
4:00
6:00
8:00
10:00
12:00
14:00
HOURS
F ig u r e 83.
O th er a c t i v i t y f o r November.
16:00
I 8: 00
20:00
22:00
IOCH
-1 0 0
OF
HERD
X
30-
IN A C T I V I T Y
50-
ENGAGED
<
4:00
6:00
8 :00
10:00
1 2:0 0
14:00
HOURS
F ig u re 84.
G r a z in g a c t i v i t y f o r Decem ber.
I 6:00
I 8:00
20:00
22:00
I 00-1
OF
ACTIVITY
-100
IN
HERD
ENGAGED
ENGAGED
OF
ACTIVITY
HERD
IN
4:00
6:00
8:00
10:00
I 2 :0 0
14:00
HOURS
F ig u r e 85.
R e s t i n g a c t i v i t y f o r D ecem ber.
16:00
I 8:00
20:00
22:00
1 OCH
OF
IN A C T I V I T Y
-100
OF
IN A C T I V I T Y
HERD
ENGAGED
ENGAGED
HERD
6 :0 0
F ig u r e 86.
8:00
10:00
12:00
14:00
HOURS
O th er a c t i v i t y f o r D ecem b er.
16:00
18:00
20:00
22:00
MONTANA STATE UNIVERSITY LIBRARIES
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