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A habitat-use and dietary analysis of a monogastric versus a ruminant herbivore, on forested range 1984

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A HABITAT-USE AND DIETARY ANALYSIS OF A MONOGASTRIC VERSUS A RUMINANT HERBIVORE, ON FORESTED RANGE BY SUSAN KAREN PRESTON B.S.A., The U n i v e r s i t y o f Saskatchewan, 1976 B . S c , The U n i v e r s i t y o f B r i t i s h C o l u m b i a , 1984 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES Department o f A n i m a l S c i e n c e We a c c e p t t h i s t h e s i s as c o n f o r m i n g t o / t h e r e q u i ^ f e ^ s t a n d a r d THE UNIVERSITY OF BRITISH COLUMBIA O c t o b e r , 1984 ® SUSAN KAREN PRESTON, 1984 In presenting t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t f r e e l y available for reference and study. I further agree that permission for extensive copying of t h i s thesis f o r ; scholarly purposes may be granted by the head of my department or by his or her representatives. I t i s understood that copying or publication of t h i s thesis for f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of o 4 f V \ \ W V I -̂S> CIA e/wg The University of B r i t i s h Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date DE-6 (3/81) ABSTRACT I n t e r a c t i o n s b e t w e e n f e r a l h o r s e s and range c a t t l e were s t u d i e d on f o r e s t e d r a n g e o f t h e C h i l c o t i n P l a t e a u i n C e n t r a l B r i t i s h C o l u m b i a f r o m May, 1979 t o A u g u s t , 1980. R e s e a r c h o b j e c t i v e s i n c l u d e d c o l l e c t i n g i n f o r m a t i o n on h a b i t a t - u s e , d i s t r i b u t i o n and d i e t s e l e c t i o n f o r h o r s e s and c a t t l e , and h a b i t a t - u s e a n d d i s t r i b u t i o n d a t a o n l y , f o r moose. B a s e d on a p p r o x i m a t e l y e q u a l , average d e f e c a t i o n r a t e s , the r e l a t i v e abundance o f each s p e c i e s i n the 200 km2 study area, was estimated t o be 89.7 p e r c e n t c a t t l e , 7.4 p e r c e n t h o r s e s and 2.9 p e r c e n t moose. The e q u i n e s t u d y p o p u l a t i o n was o r g a n i z e d i n t o f i v e harem g r o u p s , e a c h composed o f a s i n g l e s t a l l i o n w i t h a d u l t m a r e s and im m a t u r e a n i m a l s o f b o t h s e x e s , r a n g i n g i n g r o u p s i z e f r o m 5 t o 14. S u r p l u s a d u l t males formed f o u r b a c h e l o r groups o f between one and f o u r a n i m a l s . A d e t a i l e d a n a l y s i s o f a l l h a b i t a t t y p e s was n o t p r a c - t i c a b l e n o r c o n s i d e r e d e s s e n t i a l i n t h i s s t u d y . The breakdown o f t h e s t u d y a r e a i n t o s e v e n g e n e r a l h a b i t a t t y p e s was based on m a c r o v e g e t a t i o n a s s o c i a t i o n s which seemed t o r e f l e c t broad h a b i t a t - u s e p a t t e r n s and i n c l u d e d open f o r e s t , semi-open f o r e s t , c l o s e d f o r e s t , meadow, shrub c a r r , i n t e r f a c e zone and ' o t h e r ' (e.g. r o a d s ) . A b o t a n i c a l p r o f i l e o f t h e a r e a , and c o n c o m i t a n t p l a n t c o l l e c t i o n , was done over t h e two summers o f f i e l d work. P o s i t i v e i d e n t i f i c a t i o n was a c h i e v e d f o r 148 p l a n t s p e c i e s , but no attempt was made t o de t e r m i n e a v a i l a b i l i t y o f the var i o u s s p e c i e s q u a n t i t a t i v e l y . F e c a l e p i d e r m a l a n a l y s i s was c a r r i e d out on f e c a l samples c o l l e c t e d f o r h o r s e s and c a t t l e f r o m J u n e t h r o u g h September and f o r h o r s e s i n w i n t e r . R e s u l t s i n d i c a t e d a t l e a s t 54 d i f f e r e n t p l a n t s p e c i e s were u s e d by t h e two h e r b i v o r e s . E l e v e n p l a n t s p e c i e s were f o u n d t o c o n s i t u t e 80% o f t h e c a t t l e d i e t s and 86% o f t h e h o r s e d i e t s , i n d i c a t i n g a t l e a s t 80% o f t h e two h e r b i v o r e d i e t s were d e r i v e d f r o m o n l y 7.4% o f the a v a i l a b l e p l a n t s p e c i e s . H o r s e s u t i l i z e d f e w e r p l a n t s p e c i e s i n t h e w i n t e r , and w h i l e t h e use o f g r a s s e s was r e d u c e d , sedge and shrub-use i n c r e a s e d . A s y s t e m o f random t r a n s e c t s o f b e t w e e n 18 t o 25 km i n l e n g t h were u s e d on a r e g u l a r , c o n s e c u t i v e b a s i s t o c o l l e c t d i s t r i b u t i o n and h a b i t a t - u s e d a t a f o r h o r s e s , c a t t l e and moose. D e p e n d i n g on t h e i n d i v i d u a l t r a n s e c t s , a g r e a t e r o r l e s s e r degree o f s p a t i a l o v e r l a p was i n d i c a t e d f o r t h e t h r e e herbivores and t h e c o n c e n t r a t i o n o f use i n any a r e a v a r i e d w i d e l y between s p e c i e s . The d e s i g n a t e d h a b i t a t t y p e s were measured a l o n g e a c h t r a n s e c t a n d w e r e assumed t o r e f l e c t h a b i t a t a v a i l a b i l i t y f o r t h e e n t i r e s t u d y a r e a . None o f t h e t h r e e h e r b i v o r e s used t h e h a b i t a t s i n p r o p o r t i o n t o h a b i t a t a v a i l a b i l i t y . B o t h h o r s e s and c a t t l e u s e d meadow h a b i t a t d i s p r o p o r t i o n a t e l y more ( t h o u g h n o t n e c e s s a r i l y i n t h e same l o c a t i o n s ) , and c a t t l e a l s o i n d i c a t e d a p r e f e r e n c e f o r i n t e r f a c e z o n e , w h i l e moose used open f o r e s t , c l o s e d f o r e s t and shrub c a r r d i s p r o p o r t i o n a t e l y more. i i i TABLE OF CONTENTS Page A b s t r a c t . i i L i s t o f t a b l e s . v i L i s t o f f i g u r e s v i i Acknowledgement i x Chapter 1: I n t r o d u c t i o n 1 Chapter 2: M a t e r i a l s and methods 2.1 Study a r e a 6 2.2 Equine s t u d y p o p u l a t i o n 9 2.3 System o f l i n e t r a n s e c t s 9 2.4 B o t a n i c a l p r o f i l e o f s t u d y a r e a . . . . . . 12 2.5 E s t a b l i s h m e n t o f h a b i t a t t y p e s 12 2.6 A n i m a l d i s t r i b u t i o n s 20 2.7 F e c a l a n a l y s i s 22 Chapter 3: R e s u l t s 3.1 O b s e r v a t i o n s on equine p o p u l a t i o n 24 3.2 A n i m a l d i s t r i b u t i o n s 24 3.3 H a b i t a t use 37 3.4 D i e t E s t i m a t e s 43 Chapter 4: D i s c u s s i o n 4.1 H a b i t a t use r e l a t e d t o a n i m a l ' s p e r c e p t i o n o f a v a i l a b i l i t y 60 4.2 D i s t r i b u t i o n p a t t e r n s and s p a t i a l s e p a r a t i o n 68 4.3 D i e t c h o i c e by h o r s e s and c a t t l e 73 Chapter 5: Summary 85 i v TABLE OF CONTENTS (cont) Page B i b l i o g r a p h y 96 Appendix 1 L i s t o f p l a n t s i d e n t i f i e d on t h e s t u d y s i t e 107 2 Horse and c a t t l e d i e t s d e t e r m i n e d by f e c a l a n a l y s i s 117 3 P a i r e d * t ' t e s t f o r 'long-term* v e r s u s ' s h o r t - t e r m ' f e c a l c o u n t s on t r a n s e c t V 134 4 P r e s e n c e o f p l a n t t y p e s i n o r i g i n a l h a b i t a t t y p e s 139 5 N u t r i e n t a n a l y s i s r e s u l t s on c o l l e c t e d sedge m a t e r i a l 148 v LIST OF TABLES T a b l e Page 1 C o m p o s i t i o n and s i g h t i n g s o f f e r a l h o r s e groups 10 2 Changes noted i n harem group no. 2, from a t o t a l o f 29 s i g h t i n g s i n t h e p e r i o d from May 24, 1979 t o August 1, 1980 27 3 C o e f f i c i e n t o f a s s o c i a t i o n f o r h o r s e s , cows and moose 33 4 The p e r c e n t d i s t r i b u t i o n o f h a b i t a t t y p e s a l o n g the f i v e t r a n s e c t s 38 5 H a b i t a t use v e r s u s p e r c e n t h a b i t a t a v a i l a b i l i t y 39 6 k-Independent C h i Square T e s t f o r comparing h a b i t a t use between s p e c i e s 41 7 Horse and c a t t l e d i e t s e s t i m a t e d from f e c a l samples c o l l e c t e d i n J u n e , J u l y , August and September 44 8 Average use per month of t h e 11 f o r a g e s composing a t l e a s t 80 p e r c e n t o f t h e d i e t s o f b o t h h o r s e s and cows 49 9 G r a s s s p e c i e s e a t e n by horses-summer v e r s u s h o r s e s - w i n t e r 52 10 Rush and sedge s p e c i e s e a t e n by h o r s e s i n summer v e r s u s w i n t e r 54 11 Z-Values f o r d i f f e r e n c e s i n p r o p o r t i o n a l use o f b o t a n i c a l groups on a s e a s o n a l b a s i s 58 v i LIST OF FIGURES F i g u r e Page 1 Map o f s t u d y a r e a showing arrangement.of t h e f i v e t r a n s e c t s 7 2 Young heterogenous s e r a i h a b i t a t t y p e 15 3 Young homogenous s e r a i h a b i t a t t y p e 15 4 M a t u r i n g s e r a i h a b i t a t t y p e 16 5 M a t u r i n g c l i m a x h a b i t a t t y p e . . . . 16 6 Dry meadow h a b i t a t t y p e 18 7 Wet (sedge) meadow h a b i t a t t y p e . . 18 8 Shrub (meadow) c a r r h a b i t a t t y p e 19 9 Edap h i c c l i m a x (spruce) h a b i t a t t y p e 19 10 Logged/burned ( a l t e r e d ) h a b i t a t t y p e s 21 11 I n t e r f a c e zones 21 12 Map o f minimum home ranges f o r a l l o b s e r v e d f e r a l h o r s e g r o u p s , based on s i g h t i n g s 25 13 T r a n s e c t s I t h r o u g h V f showing f r e q u e n c y d i s t r i b u t i o n s o f f e c a l d r o p p i n g s f o r h o r s e s , c a t t l e and moose. 31 14 T r a n s e c t V, showing f r e q u e n c y d i s t r i b u t i o n s o f accumulated f e c a l d r o p p i n g s f o r h o r s e s , c a t t l e and moose on a l o n g - t e r m b a s i s 35 15 Whole ungrazed sedge p l a n t d u r i n g w i n t e r dormancy . . 55 16 In n e r p a r t o f dormant sedge p l a n t , showing green s h o o t s 55 v i i LIST OF FIGURES F i g u r e Page 17 The p e r c e n t o c c u r r e n c e o f t h e t h r e e major b o t a n i c a l groups i n the d i e t s o f h o r s e s and c a t t l e based on f e c a l a n a l y s i s 56 v i i i ACKNOWLEDGEMENT T h i s s t u d y was funded by the B o r e a l I n s t i t u t e f o r N o r t h e r n S t u d i e s i n A l b e r t a , the B.C. F o r e s t S e r v i c e , and a L i f e Sciences G r a n t f r o m t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a . I am d e e p l y i n d e b t e d t o t h e s e o r g a n i z a t i o n s and a number o f i n d i v i d u a l s f o r t h e u n i q u e o p p o r t u n i t y f o r f i e l d r e s e a r c h which t h i s s t u d y a f f o r d e d . A l ways l o g i s t i c a l l y d i f f i c u l t , and o f t e n p h y s i c a l l y g r u e l l i n g , t h i s s t u d y o v e r a l l , p r o v i d e d me w i t h a t r u l y memorable and w o r t h w h i l e e x p e r i e n c e . I t h a n k my s u p e r v i s o r , Dr. D. M. S h a c k l e t o n , p a r t i c u l a r l y f o r h i s p e r s e v e r a n c e and e x c e l l e n t a d v i c e on t h e a n a l y s i s a n d f i n a l p r e p a r a t i o n o f my d a t a i n t o a p r e s e n t a b l e f o r m . I n a d d i t i o n , D r . F. L. B u n n e l l , F o r e s t r y Department, U.B.C, and D r . R. M. Beames, A n i m a l S c i e n c e D e p a r t m e n t , U.B.C., members o f my e x a m i n i n g c o m m i t t e e , r e v i e w e d a t h e s i s d r a f t and p r o v i d e d u s e f u l comments. Anna R o b e r t s o f t h e B.C. F o r e s t S e r v i c e i n W i l l i a m s L a k e , p r o v i d e d v a l u a b l e a s s i s t a n c e d u r i n g t h r e e d a y s s p e n t on my s t u d y s i t e i d e n t i f y i n g g r a s s e s and s e d g e s . L e o n P a v l i c k o f t h e P r o v i n c i a l Museum H e r b a r i u m i n V i c t o r i a , was f o r t u n a t e l y a b l e t o i d e n t i f y t h e 'unknown' p l a n t s p e c i e s I c o l l e c t e d . E s s e n t i a l s u p p o r t was p r o v i d e d by the c o r r a l and c a b i n I r e n t e d on t h e p r o p e r t y o f Wayne and T i n a Plummer, t o whom I am a l s o t h a n k f u l f o r o t h e r h e l p a t c r i t i c a l t i m e s . J a n i e Heywood and K n u t A t k i n s o n p r o v i d e d i m p o r t a n t a s s i s t a n c e d u r i n g t h e f i r s t summer's f i e l d work. I w o u l d e s p e c i a l l y l i k e t o thank S. C. C h a t w i n f o r h i s h e l p i n t h e m u d - f r o u g h t f r u s t r a t i o n of f e r r y i n g horses, gear and v e h i c l e s from Vancouver to the, at times, almost inaccessible study s i t e . x Chapter 1: I n t r o d u c t i o n B r o a d l y d e f i n e d , " c o m p e t i t i o n o c c u r s when a number o f a n i m a l s , o f t h e same o r d i f f e r e n t s p e c i e s , u t i l i z e common r e s o u r c e s t h e s u p p l y o f w h i c h i s s h o r t ; o r i f t h e r e s o u r c e s a r e n o t i n s h o r t s u p p l y , c o m p e t i t i o n o c c u r s when the a n i m a l s s e e k i n g t h a t r e s o u r c e n e v e r t h e l e s s harm one o r t h e o t h e r i n t h e p r o c e s s " ( B i r c h r 1957). The concept of c o m p e t i t i o n remains a c o n t r o v e r s i a l t o p i c as p o i n t e d out by Schoener (1982), o s c i l - l a t i n g between t h e extreme views o f i t s b e i n g a r a r e b i o l o g i c a l phenomenon ( W i e n s , 1977) and a major d r i v i n g f o r c e o f n a t u r a l s e l e c t i o n ( D i a m o n d , 1 9 7 8 ) . ' S h o r t a g e ' o r l i m i t a t i o n o f a r e s o u r c e , i s a n e x t r e m e l y n e b u l o u s c o n c e p t as p o i n t e d o u t b y W h i t e ( 1 9 7 8 ) , b e c a u s e o f t h e r a n g e o f mechanisms w h i c h a n i m a l s a p p e a r t o e m p l o y t o a v o i d d i r e c t i n t e r a c t i o n . Such ' I n t e r a c t i o n A v o i d a n c e M e c h a n i s m s ' may be b e h a v i o r a l , as i n g r a z i n g t i m e s and d i u r n a l c y c l e s ( A r n o l d , 1 9 8 1 ; A r n o l d and D u d z i n s k i , 1978; A r n o l d , 1 9 6 4 ) , f e e d i n g e x p e r i e n c e ( A r n o l d , 1964; A r n o l d and M a i l e r , 1977; L e u t h o l d , 1 9 7 7 ) , f o o d h a b i t s a n d a d a p t a b i l i t y ( L a y c o c k , 1978; L e u t h o l d , 1977; C a r p e n t e r £t a l-r 1979; Smith fit a l . , 1979), ' s p e c i a l i s t ' v s . ' g e n e r a l i s t * f e e d i n g s t r a t e g y ( E l l i s and T r a v i s , 1975; Nudds, 1 9 8 0 ) , o r non-random use o f h a b i t a t ( S a n d e r s o n , 1966). The mechanisms may a l s o be p h y s i c a l , i n c l u d i n g t h e h e r b i v o r e - s p e c i f i c g r a z i n g m e c h a n i c s d i c t a t e d by t h e s p e c i e s ' morphology ( L e u t h o l d , 1977; A r n o l d and D u d z i n s k i , 1978; Moehlman, 1974), s i z e and p h y s i c a l m a i n t e n a n c e p a r a m e t e r s (Hungate §£ a j , . , 1959; E l s d e n e t a l . , 1946; Jarman, 1974), and e c o l o g i c a l a s p e c t s o f a n i m a l metabolism - 1 - ( K l e i b e r , 1961; G e i s t , 1974; B e l l , 1971). B e l l (1971) suiranmarized t h e s e l a t t e r i n t h e f o l l o w i n g s t a t e m e n t : " S m a l l a n i m a l s can t o l e r a t e d i e t s t h a t d e p a r t from t h e optimum i n q u a n t i t y ; l a r g e a n i m a l s , p a r t i c u l a r l y n o n - r u m i n a n t s , can t o l e r a t e d e p a r t u r e s i n q u a l i t y . " A l t h o u g h r e l a t i v e . f o o d s h o r t a g e c a n e x e r t a p r o f o u n d e f f e c t on t h e e c o l o g y o f a n i m a l s , t o t h e p o i n t o f s t a r v a t i o n , t h e p o i n t a t w h i c h f o o d s h o r t a g e c e a s e s t o be due t o t h e i n t e r a c t i o n between an a n i m a l and i t s e n v i r o n m e n t , a n d b e g i n s t o be b e t w e e n t h e a n i m a l and i t s a s s o c i a t e s , i s e x t r e m e l y ambiguous. I t i s i m p o r t a n t t h a t t h e two components o f t h e t h e o r y o f c o m p e t i t i o n be r e c o g n i z e d , d i s t i n g u i s h e d by P a r k (1954) as ' e x p l o i t a t i o n ' and ' i n t e r f e r e n c e . ' E x p l o i t a t i o n r e f e r s t o t h e r e l a t i v e s u c c e s s o f two or more i n d i v i d u a l s , o r s p e c i e s w h i c h have f r e e a c c e s s t o a l i m i t e d r e s o u r c e . I n t e r f e r e n c e , i s when some f o r m o f s p a c e i s s u b s t i t u t e d f o r a r e s o u r c e and i s t h e o b j e c t o f c o m p e t i t i o n ( M i l l e r , 1969; A y a l a , 1 9 7 0 ) . I n t e r f e r e n c e i s a more s t r a i g h t forward concept than e x p l o i t a t i o n , and i s more e a s i l y shown t o o c c u r , as i n t h e a g g r e s s i v e behavior o f two s p e c i e s o f chipmunks (Eutamias d o r s a l i s and g.. umbrinus) i n Nevada (Brown, 1971). Brown's s t u d y s u p p o r t s the P r i n c i p l e o f C o m p e t i t i v e E x c l u s i o n f o r m u l a t e d by G r i n n e l l (1904) and Gause ( 1 9 3 4 ) . E x p l o i t a t i o n , h o w e v e r , i s much more d i f f i c u l t t o t e s t because of the e l u s i v e n e s s o f 'at what p o i n t a resource becomes l i m i t i n g ' , f o r as W h i t e (1978) p o i n t s out ' t h e r e can be s h o r t a g e w i t h o u t c o m p e t i t i o n . ' S l o b o d k i n (1962) was i n s p i r e d t o r e p h r a s e t h e Gause P r i n c i p l e t h a t s p e c i e s c a n n o t c o e x i s t u s i n g t o o s i m i l a r l y t h e same r e s o u r c e s , as a r u l e o f e c o l o g i c a l - 2 - p r o c e d u r e r a t h e r t h a n a v e r i f i e d or v e r i f i a b l e p r o p o s i t i o n . T h i s i s n o t t o s a y t h e e x p l o i t i v e f o r m o f c o m p e t i t i o n , which i n c l u d e s d i e t o v e r - l a p , does not e x i s t , but r a t h e r t h a t evidence i n d i c a t i v e o f i t s o c c u r r e n c e s h o u l d be c a r e f u l l y e v a l u a t e d and r e s u l t s not be a n t i c i p a t e d . A r e v i e w o f c u r r e n t f i e l d - w o r k on t h e s u b j e c t ( B i r c h , 1979) s u p p o r t s t h e c o n t e n t i o n o f Den B o e r (1980) t h a t a t l e a s t i n t h e e x p l o i t i v e f orm, ' c o m p e t i t i v e e x c l u s i o n ' must be c o n s i d e r e d as o n l y an e x c e p t i o n a l outcome of the p o s s i b l e i n t e r a c t i o n s between s p e c i e s . H i s t o r i c a l l y , i t i s i n t e r e s t i n g t o compare t h e p h i l o s o p h y b e h i n d t h e e c o l o g i c a l s t u d i e s o f A f r i c a n v e r s u s N o r t h American u n g u l a t e s ; t h e two t o g e t h e r make up t h e b u l k o f l i t e r a t u r e on h e r b i v o r e i n t e r a c t i o n s . The n u t r i t i v e work on u n g u l a t e s i n A f r i c a and A s i a has f o c u s e d on s e l e c t i v i t y and e c o l o g i c a l s e p a r a t i o n t o t r y t o e x p l a i n t h e a p p a r e n t harmony o f complex c o m m u n i t i e s ( L e u t h o l d , 1977; G r i m s d e l l and F i e l d , 1976; B e l l , 1 9 7 1 ; J a r m a n , 1974; T a l b o t and T a l b o t , 1 9 6 9 ) . The work i n N o r t h A m e r i c a has c e n t e r e d a r o u n d t h e degree o f d i e t o v e r l a p a n d t h e s p a t i a l r e l a t i o n s among h e r b i v o r e s , w i t h t h e b a s i c p r e m i s e t h a t c o m p e t i t i o n between s p e c i e s i s i n h e r e n t f o r l i m i t e d r a n g e ( M c M a h a n , 1964; H a n s e n , 1976; S t o r r a r §_£. aJL. , 1977; W i l l m s e t a l . , 1979). I t i s p o s s i b l e t h a t t h i s l a t t e r emphasis came a b o u t b e c a u s e t h e w e s t e r n r a n g e l a n d s were d e p l e t e d o f f o r a g e , due t o o v e r s t o c k i n g o f d o m e s t i c a n i m a l s as e a r l y as 1880 ( P a r k e r , 1954), and s e v e r a l n a t i v e ungulates were d r a s t i c a l l y r e d u c e d i n numbers ( e . g . C e r v u s e l a p h u s n e l s o n i ) a l o n g w i t h t h e i r n a t u r a l p r e d a t o r s (e.g. C a n i s l u p u s , Ursus a r c t o s ) l o n g b e f o r e b r o a d s p e c t r u m e c o l o g i c a l s t u d i e s b e g a n . R e c e n t l y , - 3 - h o w e v e r , t h e p e n d u l u m has begun t o s w i n g b a c k i n f a v o r o f m u l t i - s p e c i e s range management (Richmond, 1980; Hudson, 1980). A t t h i s t i m e , no c a s e s o f e x p l o i t i v e c o m p e t i t i o n have been p r o v e n t o o c c u r among u n g u l a t e s i n N o r t h A m e r i c a , o n l y s i m i l a r i t i e s i n h a b i t a t - u s e and d i e t have been i n d i c a t e d . S i m i l a r i t y o f u s e does n o t n e c e s s a r i l y i m p l y c o m p e t i t i o n , b u t i n r e g a r d t o f e r a l e q u i d s (Equus c a b a l l u s and E. a s i n u s ) and o t h e r h e r b i v o r e s t h e r e has been a t r e n d , b o t h h i s t o r i c a l l y ( M c K n i g h t , 1958 and 1959; S t o r r a r e_ , 1977), and i n r e c e n t l i t e r a t u r e ( K o e h l e r , 1961; W i l l m s e t a i . , 1975; Hansen, 1976; H a n s e n e t a_. , 1977) t o e q u a t e • s i m i l a r i t y , ' ' o v e r l a p ' and sometimes o n l y ' j u x t a p o s i t i o n , 1 w i t h t h e concept of competition. Management d e c i s i o n s , i n c l u d i n g complete removal o f f e r a l e q u i d s i n l o c a l i z e d a r e a s , have been made i n t h e p a s t , b o t h i n t h e U n i t e d S t a t e s ( M c K n i g h t , 1958; Thomas, 1979) and i n C a n a d a ( S a l t e r and H udson, 1 9 7 8 b ) , on t h e a s s u m p t i o n t h a t c o m p e t i t i o n e x i s t s b e t ween f e r a l h o r s e s and b u r r o s w i t h o t h e r f o r m s o f w i l d l i f e and w i t h l i v e s t o c k . However, as o b v i o u s as i t may seem i t must be emphasized t h a t r e a l i s t i c management d e c i s i o n s can o n l y be made w i t h adequate knowledge, and knowledge o f t h e h a b i t s and d i e t s o f f e r a l e q u i d s i n the v a r i o u s n i c h e s t h a t t h e y o c c u p y i s v e r y s c a r c e , p a r t i c u l a r l y on f o r e s t e d r anges. The b road g o a l o f the present study was t o o b t a i n i n f o r m a t i o n on t h e i n t e r a c t i o n s b e t w e e n h o r s e s and c a t t l e (Bos t a u r u s ) r e g a r d i n g h a b i t a t - u s e , d i s t r i b u t i o n and d i e t . Moose ( A l c e s a l c e s ) were i n c l u d e d i n h a b i t a t - u s e and d i s t r i b u t i o n d a t a , b u t n o t i n t h e d i e t a n a l y s i s . S p e c i f i c o b j e c t i v e s were as - 4 - f o l l o w s : 1. E s t a b l i s h g e n e r a l h a b i t a t t y p e s and det e r m i n e t h e p e r c e n t a v a i l a b i l i t y o f e a c h t y p e i n t h e e n t i r e s t u d y a r e a . 2. D e t e r m i n e , i n r e l a t i o n t o t h e way h a b i t a t s were a v a i l a b l e , whether h o r s e s , cows, and moose were s e l e c t i n g f o r p a r t i c u l a r h a b i t a t t y p e s . 3. D e t e r m i n e w h e t h e r t h e r e were d i f f e r e n c e s i n t h e c h o i c e of h a b i t a t t y p e s among h o r s e s , cows and moose. 4. I n v e s t i g a t e w h e t h e r h o r s e s , cows and moose were d i s t r i - b u t i n g t h e m s e l v e s d i f f e r e n t l y i n space. 5 . E s t a b l i s h w h a t p l a n t s ( i . e . v a r i e t y o f s p e c i e s ) were a v a i l a b l e t o t h e a n i m a l s i n t h e s t u d y a r e a . 6. D e t e r m i n e , by f e c a l e p i d e r m a l a n a l y s i s , what p l a n t s p e c i e s comprised t h e d i e t s o f h o r s e s and cows, and t o what degree t h e d i e t s were s i m i l a r . 7. D e t e r m i n e w h e t h e r t h e same b o t a n i c a l g r o u p s were b e i n g used i n t h e same p r o p o r t i o n s by h o r s e s and by cows. 8. Compare t h e d i e t s o f h o r s e s f o r t h e s p r i n g , summer and f a l l w i t h t h e i r w i n t e r d i e t s . 9. D e t e r m i n e t h e b r o a d s o c i a l o r g a n i z a t i o n o f f e r a l h o r s e s found i n the s t u d y a r e a , and e s t a b l i s h r e l a t i v e home-ranges o f t h e groups. - 5 - Chapter 2: M a t e r i a l s and Methods 2.1 Study A r e a The s t u d y began i n May o f 1979 and t h e f i e l d work was d i v i d e d i n t o t h r e e p e r i o d s as f o l l o w s : May 23 t o O c t o b e r 1, 1979; M a r c h 1 t o M a r c h 10, 1980; and June 1 t o August 15, 1980. A t o t a l o f 6 1/2 months was spent on t h e s t u d y s i t e . The s t u d y a r e a was l o c a t e d b e t w e e n H a i n e s L a k e and Gay L a k e (123° 24* W, 51° 57' N) , 24.3 km S. W. o f A l e x i s Creek, B.C., a t an e l e v a t i o n o f 1050 m a s l , and c o v e r e d a p p r o x i m a t e l y 200 km 2 ( F i g u r e 1). The a r e a i s g e n e r a l l y f l a t w i t h g e n t l y r o l l i n g t e r r a i n , c o v e r e d by a f o r e s t o f Douglas f i r (Pseudotsuoa m e n z i e s i i ) and l o d g e p o l e p i n e ( P i n u s c o n t o r t a ) . w i t h l o c a l i z e d s p r u c e and p o p l a r , i n t e r s p e r s e d w i t h l a k e s , marshes and meadows. The t o p o g r a p h i c a l f e a t u r e s a r e a r e s u l t o f combined g l a c i a l and v o l c a n i c a c t i v i t y . Much o f t h e a r e a i s r a t h e r d i f f i c u l t t o t r a v e r s e b e c a u s e o f t h e f o r e s t d e n s i t y and t h e w i d e s p r e a d o c c u r r e n c e o f b o u l d e r f i e l d s o f g l a c i a l and v o l c a n i c o r i g i n . T h e s e f e a t u r e s a l s o a f f e c t t h e d i s t r i b u t i o n and movements o f w i l d l i f e and d o m e s t i c s t o c k . The a r e a i s c l a s s i f i e d i n t h e I n t e r i o r Douglas F i r (IDF) B i o g e o c l i m a t i c Zone (Annas and Coupd, 1979), i n c l u d i n g a l l a r e a s t h a t c l i m a x i n D o u g l a s f i r on m e s i c s i t e s . The IDF (b) s u b z o n e ( D o u g l a s f i r - p i n e g r a s s , n o r t h e r n p h a s e ) i s an e x t r e m e l y i m p o r t a n t f o r e s t e d subzone i n terms o f range p r o d u c - t i v i t y , and p r o v i d e s the m a j o r i t y o f summer g r a z i n g f o r c a t t l e i n t h e c e n t r a l B.C. r e g i o n (Annas and Coupe, 1979). The s t u d y a r e a was l o c a t e d i n t h e more w e s t e r l y p o r t i o n s o f t h e IDF(b) - 6 - F i g u r e 1: Map o f s t u d y a r e a , s h o w i n g a r r a n g e m e n t of t h e f i v e t r a n s e c t s , ranging from 18 t o 25 km, r a d i a t i n g f r o m t h e c e n t e r o f t h e s t u d y s i t e , " W i l s o n ' s Meadow" ( d i s c u s s e d i n s e c . 2.3). - 7 -  subzone. The study area includ e d parts of three d i f f e r e n t range allotments where c a t t l e (Herefords predominating) ranged from June through October with a peak grazing density i n July-August. A p o p u l a t i o n of f e r a l horses also used the study area divided for the most part, into d i s t i n c t s o c i a l groups. 2.2 Equine Study Population A f l u c t u a t i n g number of domestic horses shared the study a r e a with 65 horses i d e n t i f i e d as f e r a l . The f e r a l horses were organized i n t o at l e a s t e i g h t d i s t i n c t groups, plus one i n d i v i d u a l s t a l l i o n without a harem. Table 1 shows the basic composition of the groups i d e n t i f i e d . 2.3 System of Line Transects Although a e r i a l photographs were used during the study to e s t a b l i s h transects, and to supply more s p e c i f i c information of land features than i s found on topographical maps, an overall photo i n t e r p r e t a t i o n of the area to determine the percent of each h a b i t a t type a v a i l a b l e proved to be i m p r a c t i c a l . The r e g i o n i s composed of h a b i t a t s which are i n too t i g h t a mosaic to d i s t i n g u i s h r e a l i s t i c a l l y on black and white photo- graphs. Such h a b i t a t types as 'Interface Zones' and those based on the precise separation of forest density, are d i f f i c u l t to r ecognize, although these areas probably have b i o l o g i c a l s i g n i f i c a n c e regarding animal u t i l i z a t i o n . Habitat-type separation would probably be f e a s i b l e by a e r i a l photograph interpretation i f colored photographs were available for the region. - 9 - TABLE 1: C o m p o s i t i o n and S i g h t i n g s o f F e r a l Horse Groups S t a l l i o n F o a l s , up S i g h t i n g s L o c a t i o n s Groups Mares t o two y e a r s T o t a l * 4 3 1 8 5 14 29** 8 2 3-5 2-7 6-13 6 4 3 3 1 5 5 5 4 9 3 13 1 1 5 6 3 10 B a c h e l o r Groups 4 4 6 3 2 2 7 2 5 5 8 4 6 5 9 1 • i n c l u d i n g s t a l l i o n **group most c l o s e l y o b s e r v e d . A l l group changes noted over 15 months (Table 3 ) . - 10 - A m e t h o d o f r a n d o m t r a n s e c t a n a l y s i s was u s e d i n t h e s t u d y which more a c c u r a t e l y r e f l e c t e d t h e v e g e t a t i o n a l components a v a i l a b l e t o a n i m a l s . The b a s i c method i n v o l v e d the development o f a m o d i f i e d t r a n s e c t s y s t e m i n the form o f f i v e ' c i r c u i t s , 1 a l l o r i g i n a t i n g and r e t u r n i n g t o t h e c e n t r a l a r e a where the s t u d y c a b i n was l o c a t e d a t ' W i l s o n ' s Meadow' ( F i g u r e 1 ) . The f i n a l l a y o u t o f t h e t r a n s e c t s was d e t e r m i n e d by p r a c t i c a l c o n s i d e r a t i o n s , i n c l u d i n g where s a d d l e h o r s e s c o u l d maneuver and the d i s t a n c e which c o u l d be c o v e r e d on a d a i l y and s e a s o n a l b a s i s . The f i n a l f i v e t r a n s e c t s r a n g e d f r o m 18 t o 25 km i n l e n g t h ( F i g u r e 1 ) , and were d i v i d e d i n t o 0.5-km i n t e r v a l s on t h e ground t o ensure t h a t t h e r i d e r ' s p o s i t i o n on the t r a n s e c t w o u l d be known w i t h r e a s o n a b l e a c c u r a c y w h i l e d a t a were b e i n g c o l l e c t e d . The a c c u m u l a t e d d i s t a n c e o f e a c h h a b i t a t t y p e a l o n g e a c h t r a n s e c t was r e c o r d e d , e n a b l i n g t h e p r o d u c t i o n o f a c o m p o s i t e p i c t u r e o f the h a b i t a t t y p e s a v a i l a b l e on each t r a n s e c t . I n a d d i t i o n , each t r a n s e c t was r i b b o n e d w i t h s u r v e y t a p e and l i g h t l y c l e a r e d o f o v e r h a n g i n g branches t o f a c i l i t a t e r i d i n g them r e p e a t e d l y and as a c c u r a t e l y as p o s s i b l e . M i n i m a l v e g e t a t i o n was removed t o a v o i d e n c o u r a g i n g a n i m a l s t o use the t r a n s e c t s . T h i s i s an o b v i o u s danger when u s i n g a repeating t r a n s e c t system i n h e a v i l y wooded t e r r a i n . There was no i n d i c a t i o n ( e . g . s u c h as f e c a l o r h o o f p r i n t c o n c e n t r a t i o n s ) d u r i n g the two summers' f i e l d w o r k , t h a t t h e t r a n s e c t s u n d u l y a f f e c t e d t h e movements o f a n i m a l s i n t h e a r e a . C e r t a i n l y no segment of any t r a n s e c t was adopted as a r e g u l a r t r a v e l r o u t e . E a c h t r a n s e c t was r i d d e n on a r e g u l a r , c o n s e c u t i v e b a s i s ( i . e . t h e o r d e r o f r i d i n g t h e t r a n s e c t s was n o t c h a n g e d ) , - 11 - d u r i n g w h i c h a l l f r e s h h o r s e , cow, and moose s i g n s w i t h i n 3 m o f e i t h e r s i d e o f t h e s a d d l e h o r s e were r e c o r d e d on d a t a s h e e t s . T h i s method worked w e l l f o r t h e s e t h r e e s p e c i e s because s i g n s i n t h e f o r m o f f e c e s , b e d s , and t r a c k s a r e l a r g e and c a n be q u i c k l y and e f f i c i e n t l y r e c o r d e d f r o m h o r s e b a c k i n the d i f f e r e n t h a b i t a t t y p e s . L i t t l e d i f f i c u l t y was en c o u n t e r e d i n i d e n t i f y i n g t h e r e s p e c t i v e f e c e s as has been shown t o be a p r o b l e m i n s t u d i e s o f more c l o s e l y r e l a t e d s p e c i e s s u c h as e l k and moose ( N e f f , 1 9 6 8 ) . S a d d l e h o r s e d e f e c a t i o n s were removed f r o m t h e t r a n s e c t s when t h e y o c c u r r e d . Each t r a n s e c t was r i d d e n a t o t a l o f 12 t i m e s , t h r e e t i m e s e a c h i n A u g u s t and S e p t e m b e r , 1979; and t h r e e t i m e s e a c h i n June and J u l y , 1980. The a c c u m u l a t e d d i s t a n c e r i d d e n f o r a l l t r a n s e c t s was n e a r l y 1200 km. 2.4 B o t a n i c a l P r o f i l e o f Study A r e a . I n o r d e r t o e s t a b l i s h some system o f h a b i t a t c l a s s i f i c a t i o n a n d t o c a r r y o u t a d i e t a n a l y s i s , i t was n e c e s s a r y t o have a c l e a r i d e a o f p l a n t s p e c i e s p r e s e n t i n t h e a r e a . S u b s e - q u e n t l y , a b o t a n i c a l survey of a l l p l a n t s observed, and concomitant p l a n t c o l l e c t i o n , were made o v e r t h e two summers o f f i e l d w o r k . P o s i t i v e i d e n t i f i c a t i o n was a c h i e v e d f o r 148 p l a n t s p e c i e s ( A p p e n d i x 1 ) . Due t o t i m e and manpower c o n s t r a i n t s , no a t t e m p t was made t o d e t e r m i n e a v a i l a b i l i t y o f the v a r i o u s p l a n t s p e c i e s q u a n t i t a t i v e l y . 2.5 E s t a b l i s h m e n t o f H a b i t a t Types The b r e a k d o w n o f t h e s t u d y a r e a i n t o g e n e r a l h a b i t a t - 12 - t y p e s was b a s e d on m a c r o - v e g e t a t i o n a s s o c i a t i o n s . A l t h o u g h t h e r e i s no d o u b t t h a t m i c r o - h a b i t a t s a r e o f t e n i m p o r t a n t t o a n i m a l s , a d e t a i l e d a n a l y s i s o f a l l h a b i t a t t y p e s was not p r a c t i c a b l e . I n some c a s e s , t h e r e were l a r g e v a r i a t i o n s i n p l a n t t y p e s b e t w e e n h a b i t a t s . C o n s e q u e n t l y , t h i s v a r i a t i o n was u s e d a s t h e b a s i s o f s e p a r a t i o n . I n o t h e r i n s t a n c e s , h a b i t a t d e s i g n a t i o n was b a s e d more on s p a t i a l d i f f e r e n c e s , s u c h as t h e s p a c i n g between the t r e e s i n s t a n d s of l o d g e p o l e . Resource A n a l y s i s Branch (R.A.B.) s u c c e s s i o n a l s t a g e terminology was u s e d i n d e f i n i n g h a b i t a t t y p e s ( W a l m s l e y et. a_. , 1980) on wooded s i t e s . H a b i t a t d e s i g n a t i o n was a l s o d e s c r i b e d i n t e r m s o f r e l a t i v e d e n s i t y f r o m v i s u a l e s t i m a t i o n s o f f o l i a g e c o v e r . I n i t i a l l y , 10 h a b i t a t t y p e s were d e s c r i b e d a l o n g t h e t r a n s e c t s ; a model h a b i t a t s i t e i n each case b e i n g d e s c r i b e d on R.A.B. h a b i t a t d e s c r i p t i o n f o r m s . "Young s e r a i " r e f e r s t o n o n - c l i m a x s p e c i e s which have not y e t gone t h r o u g h a s e r i e s o f n a t u r a l t h i n n i n g s , and a r e g e n e r a l l y l e s s t h a n 60 y e a r s o l d . The c a n o p y i s e s s e n t i a l l y o f e v e n h e i g h t and t h e r e i s l i t t l e r e g e n e r a t i o n . " M a t u r i n g s e r a i " i s g e n e r a l l y between 60 t o 140 y e a r s o f age and has gone t h r o u g h the f i r s t t h i n n i n g s t a g e s , a c q u i r i n g community s t r u c t u r e and some r e g e n e r a t i o n . The h a b i t a t t y p e s were: 1. Y o u n g h e t e r o g e n o u s s e r a i : E s s e n t i a l l y s u b s c r i b i n g t o t y p i c a l y oung s e r a i f e a t u r e s e x c e p t t e n d i n g t o be found on s o u t h - f a c i n g s l o p e s w h i c h seems t o f a v o r t h e presence o f some D o u g l a s f i r . T h e s e a r e a s a l s o t e n d t o be i n j u x t a p o s i t i o n t o m a t u r i n g c l i m a x . D e n s i t y e s t i m a t e i s - 13 - semi-open ( F i g u r e 2 ) . 2. Y o u n g homogenous s e r a i : More s t r i c t l y c o n f o r m i n g t o t y p i c a l y oung s e r a i f o r m w i t h l i t t l e t o moderate ground c o v e r . D e n s i t y e s t i m a t e i s d e n s e t o v e r y dense ( F i g u r e 3 ) . 3. M a t u r i n g s e r a i : T h e r e t e n d t o be no ' v e t e r a n s ' , and f a l l e n t r e e s t e n d t o l i t t e r t h e f o r e s t f l o o r . U s u a l l y f a i r l y u n i f o r m s e r a i s p e c i e s ( i . e . l o d g e p o l e ) . D e n s i t y e s t i m a t e i s semi-open ( F i g u r e 4 ) . 4. M a t u r i n g c l i m a x : Community s t r u c t u r e w i t h D o u g l a s f i r dominant and l o d g e p o l e w e l l r e p r e s e n t e d . A few f i r e - s c a r r e d ' v e t e r a n ' f i r s . R e g e n e r a t i o n o f b o t h f i r and l o d g e p o l e . D e s i t y e s t i m a t e i s open ( F i g u r e 5 ) . 5. Dry Meadow: Two s u b - t y p e s . a. Open meadow - o f t e n i n j u x t a p o s i t i o n t o sedge meadows ( w h i c h may be l o w p o i n t s i n l a r g e open a r e a s ) . F o r b , g r a s s and some sedge r e p r e s e n t a t i o n . b. D r y , o p e n h i l l s i d e s t e n d i n g t o be s l i g h t l y more p r o d u c t i v e i n v a r i e t y o f s p e c i e s t h a n t h e f l a t , d r y a r e a s , w i t h more t r e e and shrub c o v e r . Both t y p e s a r e s u s c e p t i b l e t o o v e r g r a z i n g ( F i g u r e 6 ) . 6. Wet ( s e d g e ) meadow: Low a r e a s o f w a t e r a c c u m u l a t i o n , some t e n d i n g t o s t a y wet y e a r round. Reasonably r e s i l i e n t t o g r a z i n g p r e s s u r e . S ometimes i n a c c e s s i b l e i n v e r y wet y e a r s . R e l a t i v e l y l o w p l a n t s p e c i e s v a r i e t y and - 14 - Figure 2: Young heterogenous ser a i habitat type  l i t t l e t r e e or shrub growth ( F i g u r e 7 ) . t 7. Shrub (meadow) c a r r : Areas f r e q u e n t l y f l o o d e d from s p r i n g - thaw u n t i l June, w i t h m o i s t s o i l f o r m a j o r i t y of summer-fall s e a s o n i n an a v e r a g e y e a r ( H e y e s , 1979) w i t h g r a s s e s and s e d g e s . Bog b i r c h ( B e t u l a s p p.) i s , d o m i n a n t , w i t h d r i e r ' p o c k e t s ' o f w h i t e s p r u c e ( P i c e a g l a u c a ) . D e n s i t y e s t i m a t e i s o p e n , w i t h d e n s e s h r u b c o v e r ( F i g u r e 8 ) . 8. E d a p h i c c l i m a x ( s p r u c e ) : T e n d i n g t o be f o u n d e i t h e r i n e r r a t i c a l l y i n t e r s p e r s e d l o w l y i n g a r e a s ( r a v i n e s ) ; o r c o n s i s t e n t l y i n j u x t a p o s i t i o n w i t h wet meadows and s h r u b c a r r a r e a s . P r o v i d e f a v o r e d b e d d i n g s i t e s f o r c a t t l e and p a r t i c u l a r l y moose. F r e e wa'ter a b s e n t , but t e n d e n c y f o r i t t o a c c u m u l a t e i n d e p r e s s i o n s and edges o f a r e a s c o i n c i d e n t a l w i t h s h r u b m a r g i n s - a r e a s hence i a l w a y s damp and c o o l ( p r o l i f i c m o s q u i t o - b r e e d i n g grounds) - r e f l e c t e d i n h y d r o p h i l i c p l a n t s p e c i e s (e.g. Sphagnum m o s s e s ) . D e n s i t y e s t i m a t e i s u s u a l l y q u i t e dense ( F i g u r e 9 ) . 9. Logged/burned ( a l t e r e d ) s i t e s : Tending t o be r e g e n e r a t i v e a t v a r i o u s ages; s i t e s a r e v e r y ' c l u t t e r e ' d * due t o d e b r i s ( i . e . l i m b s , stumps, l o g s ) c a u s i n g herb l a y e r t o be c o n s i s - t e n t l y d i s j u n c t . The d e n s i t y e s t i m a t e i s g e n e r a l l y q u i t e open ( F i g u r e 1 0 ) . ' 10. I n t e r f a c e Zones: E s s e n t i a l l y wet t o d r y t r a n s i t i o n zones o f S p r u c e - A s p e n - L o d g e p o l e o f v a r y i n g a g e s . T e n d i n g t o i - 17 - F i g u r e 7: Wet meadow h a b i t a t t y p e - 18 - F i g u r e 9: E d a p h i c c l i m a x ( s p r u c e ) h a b i t a t t y p e - 19 - be v e g e t a t i v e l y (and o f t e n p h y s i c a l l y ) complex - h a v i n g h i g h v e g e t a t i v e d i v e r s i t y i n k e e p i n g w i t h Ecotone b i o l o g y . The d e n s i t y e s t i m a t e i s dense t o semi-open ( F i g u r e 1 1 ) . I n t h e f i n a l h a b i t a t a n a l y s i s , t h e t e n o r i g i n a l d e s i g n a - t i o n s were reduced t o seven c a t e g o r i e s which seemed t o r e f l e c t broad h a b i t a t - u s e p a t t e r n s . They were: 1. Open F o r e s t : combining m a t u r i n g c l i m a x and logged/burned s i t e s . 2. S e m i-open F o r e s t : c o m b i n i n g m a t u r i n g s e r a i and h e t e r o - genous young s e r a i . 3. C l o s e d F o r e s t : homogenous young s e r a i . 4. Meadow: c o m b i n i n g wet and d r y ( s p a t i a l l y d i f f i c u l t t o s e p a r a t e ) . 5. Shrub C a r r . 6. I n t e r f a c e Zone: c o m b i n i n g e d a p h i c c l i m a x and i n t e r f a c e zones. 7. ' O t h e r 1 : r o a d s , e t c . 2.6 A n i m a l D i s t r i b u t i o n s To d e t e r m i n e w h e t h e r o r n o t t h e t h r e e s p e c i e s showed any l e v e l o f a s s o c i a t i o n among each o t h e r , I c o n s i d e r e d t h e i r p r e s e n c e o r a b s e n c e , r e l a t i v e t o e a c h o t h e r , a t e a c h .5 km p o i n t a l o n g a l l f i v e t r a n s e c t s . S p e c i e s were thus t a b u l a t e d - 20 - F i g u r e 1 0 : Logged/burned ( a l t e r e d ) h a b i t a t t y p e a s e i t h e r a l o n e , w i t h one o t h e r s p e c i e s , a l l t o g e t h e r , o r none p r e s e n t . From t h e s e v a l u e s a c o e f f i c i e n t o f a s s o c i a t i o n , C, was c a l c u l a t e d w h i c h c a n v a r y f r o m +1.0 t o -1.0 ( S m i t h , 1 9 8 0 ; 7 0 2 ) . The s i g n i f i c a n c e o f e a c h c o e f f i c i e n t was t h e n d e t e r m i n e d by a c o n t i n g e n c y C h i Square T e s t . 2.7 F e c a l A n a l y s i s F e c a l s a m p l e s f o r d i e t e s t i m a t e s were c o l l e c t e d o n l y f o r h o r s e s and cows d u r i n g t h e two summers' f i e l d work. One m o n t h ' s s a m p l e f o r e a c h s p e c i e s a l w a y s c o n s i s t e d o f e i g h t t e a s p o o n s o f m a t e r i a l , e a c h one t a k e n from e i g h t i n d i v i d u a l , f r e s h dung p i l e s , not always on t h e same s i t e . The o p p o r t u n i s t i c a p p r o a c h o f c o l l e c t i n g f e c a l m a t e r i a l , whenever and wherever f o u n d , was adopted because o f the u n p r e d i c t a b i l i t y of encountering h o r s e s . I t was d e t e r m i n e d by e x p e r i e n c e t h a t e i g h t i n d i v i d u a l f e c a l p i l e s was t h e u p p e r l i m i t o f what c o u l d be c o n s i s t e n t l y found f o r b o t h s p e c i e s w i t h i n t h e space o f 1 t o 2 days. Between 7 and 13 c o l l e c t i o n s o f f e c a l m a t e r i a l f o r b o t h s p e c i e s were made t h r o u g h o u t e a c h month f r o m J u ne t h r o u g h S e p t e m b e r , t o a m e l i o r a t e t h e p o s s i b l e e f f e c t o f c o n c e n t r a t i o n s o f p l a n t t y p e s due t o v e g e t a t i v e 'growth s p u r t s . ' The f e c a l m a t e r i a l , i m m e r s e d i n 90% a l c o h o l , was s t o r e d i n a i r - t i g h t j a r s , and t h e d a t e o f c o l l e c t i o n and a l l p e r t i n e n t i n f o r m a t i o n r e c o r d e d . Samples f o r a n a l y s i s , chosen from th e t o t a l monthly c o l l e c t i o n s , were e v e n l y d i s t r i b u t e d w i t h r e s p e c t t o t i m e . The f e c a l samples were sent t o the Washington Stat e U n i v e r s i t y W i l d l i f e H a b i t a t Lab f o r a n a l y s i s . They c o n s i s t e d o f f o u r o f t h e c o m p o s i t e s a m p l e s o f e i g h t i n d i v i d u a l s e a c h , f o r b o t h - 22 - h o r s e s and cows, f o r June, J u l y , August and September - denoted, f o r e x a m p l e , f o r cows i n June as J-C^; J-C2# e t c . (Appendix I I ) . I n a d d i t i o n , d u r i n g t h e w i n t e r o f 1979-80, t e n d a y s w e r e s p e n t a t t h e s t u d y s i t e c o l l e c t i n g h o r s e f e c e s o n l y . S amples were c o l l e c t e d e i t h e r f r e s h o r f r o z e n on t o p o f the s n o w , r e p r e s e n t i n g random c o l l e c t i o n t h r o u g h o u t t h e s t u d y s i t e o f one t e a s p o o n e a c h f r o m 96 i n d i v i d u a l dung p i l e s . T h ese i n d i v i d u a l s a m p l e s were a l l c o m p o s i t e d and two samples f o r d i e t a n a l y s i s t a k e n o u t . Thus th e w i n t e r d i e t s r e p r e s e n t a l a r g e r s i n g l e s a m p l e o v e r an u n d e t e r m i n e d t i m e p e r i o d . The s p r i n g / s u m m e r / f a l l d i e t a n a l y s i s f o r h o r s e s w i l l be c o l l e c - t i v e l y r e f e r r e d t o as '^prsersLura^er' t o d i s t i n g u i s h i t f r o m t h e ' H o r s e - w i n t e r ' d i e t a n a l y s i s . F o r e a c h o f t h e 34 d i e t a n a l y s e s , e i g h t m i c r o s c o p e s l i d e s w e r e e x a m i n e d f o r p l a n t m a t e r i a l t h a t c o u l d be i d e n t i f i e d , c o n s i s t i n g o f 25 f i e l d s o f v i e w per s l i d e ( a l t e r n a t i v e l y r e f e r r e d t o a s ' c o v e r p o i n t s ' ) , o r a t o t a l o f 200 v i e w s p e r d i e t . Because each monthly d i e t was d e t e r m i n e d from f o u r d i e t samples, e a c h month c o n s i s t s o f 800 v i e w s e a c h , f o r b o t h h o r s e s and c a t t l e . Due t o e c o n o m i c r e s t r a i n t s , o n l y two w i n t e r d i e t s a m p l e s were e x a m i n e d , s u s e q u e n t l y t h e w i n t e r d i e t a n a l y s i s i s based on a t o t a l o f 400 v i e w s . A l e v e l o f 5% p r o b a b i l i t y was c h o s e n a p r i o r i f o r a l l t e s t s o f h y p o t h e s e s . S p e c i f i c d e t a i l s o f t h e a n a l y s e s used ar e p r e s e n t e d i n Chapter 3. - 23 - Chapter 3: RESULTS 3.1 O b s e r v a t i o n s on Equine P o p u l a t i o n F o r a t o t a l o f 90 horse s i g h t i n g s , i n c l u d i n g i n d i v i d u a l s and g r o u p s , f o r a l l time spent i n t h e f i e l d , 65 p e r c e n t were c o n s i d e r e d f e r a l , 28 p e r c e n t d o m e s t i c , 3 p e r c e n t mixed and 4 p e r c e n t unknown. Not s u r p r i s i n g l y , s i g h t i n g s were b i a s e d t o w a r d s meadows, w i t h 86 p e r c e n t o f h o r s e s f o u n d i n them, and 8 p e r c e n t i n i n t e r f a c e z o n es, 3 p e r c e n t i n semi-open f o r e s t , 2 p e r c e n t i n c l o s e d f o r e s t , 1 p e r c e n t i n o t h e r , and 0 p e r c e n t i n open f o r e s t and shrub c a r r . The home r a n g e o b s e r v a t i o n s were o p p o r t u n i s t i c , as a h i g h p r i o r i t y was not p l a c e d on o b t a i n i n g b e h a v i o r a l o r group i n f o r m a t i o n , and t h e mapping o f minimum home r a n g e s b a s e d on a l i m i t e d number o f s i g h t i n g s i s r a t h e r tenuous a t b e s t . However, i t i s f e l t t h a t t h e o b s e r v a t i o n s do r e f l e c t a pproximate a r e a s o f u s e , r e l a t i v e t o t h e e n t i r e s t u d y a r e a , f o r t h e e i g h t h o r s e g r o u p s ( F i g u r e 1 2 ) . The most a c c u r a t e range g i v e n i s f o r Harem g r o u p #2, b a s e d on 29 s i g h t i n g s , p e r h a p s due t o t h e c o i n c i d e n c e o f i t b e i n g l o c a t e d c e n t r a l l y i n the s t u d y a r e a . I n a d d i t i o n , c h a n g e s i n 'group c o m p o s i t i o n o f Harem Group #2 were a l s o r e c o r d e d (Table 2 ) . 3.2 A n i m a l D i s t r i b u t i o n s I n a d d i t i o n t o p r o v i d i n g an e s t i m a t e o f t h e t y p e s and p e r c e n t a g e s o f each h a b i t a t t y p e a v a i l a b l e , t h e d a t a c o l l e c t e d from t h e t r a n s e c t s f a l l i n t o two major i n f o r m a t i o n a l c a t e g o r i e s : - 24 - FIGURE 12: Map o f minimum home ranges f o r a l l o b served f e r a l h o rse g r o u p s , based on s i g h t i n g s . - 25 - - 26 - TABLE 2: Changes n o t e d i n Harem Group No. 2, from a t o t a l o f 29 s i g h t i n g s i n t h e p e r i o d from May 24, 1979 t o August 1, 1980, showing a d d i t i o n s and l o s s e s t o t h e band. - 27 - Date o f S i q h t i n q O r i q i n a l C o m p o s i t i o n A d d i t i o n s L o s s e s May 24, 1979 Chestnut S t a l l i o n , about 13 y r s . Heavy b l a c k mare, about 12 y r s . - wearing o l d l e a t h e r c o l l a r . P a r t i - c o l o r e d male y e a r l i n g b e l o n g i n g t o b l a c k mare. 2 ( w i l d ) i d e n t i c a l brown mares, about 3-5 y r s . 1 b r . female y e a r l i n g , b e l o n g i n g t o one o f b r . mares June 14, 1979 Br. male f o a l b o r n t o one o f brown mares. June 24, 1979 Br. female f o a l b o r n to o t h e r b r . mare. Aug. 10, 1979 B r . male 2 mo. o l d f o a l l o s t from group f o r a t l e a s t 8 h o u r s . L a t e r found r e u n i t e d . March 7, 1980 Red male f o a l b o r n t o b l a c k mare a f t e r m i d - August, 1979. 1 brown mare & male f o a l o f p r e v i o u s June c o n t i n u e d . . . T a b l e 2 c o n t i n u e d . . . Date o f S i g h t i n g O r i g i n a l C o m p o s i t i o n A d d i t i o n s L o s s e s March 7, 1980 (cont.) June 3, 1980 J u l y 31, 1980 F i n a l c o m p o s i t i o n . Chestnut S t a l l i o n . B l . mare & red f o a l o f Aug. 1979. 1 b r . mare w i t h new f o a l and y e a r l i n g female o f June, 1979. 1 (2-3 y r . o l d ) b r . mare w i t h new f o a l . 1 roan mare. New c h e s t n u t mare. New bay mare. B r . f o a l b o r n t o b r . y e a r l i n g o f 1979. New Roan mare. B r . f o a l b o r n t o r e m a i n i n g o r i g i n a l b r . mare - w i t h y e a r l i n g f e m a l e . P a r t i - c o l o r e d male. 2 New m a r e s s e e n i n March. 1. R e l a t i v e use o f each h a b i t a t t y p e by c a t t l e , h o r s e s and moose ( s e c t i o n 3 .3), and 2. D i s t r i b u t i o n by e a c h a n i m a l s p e c i e s i n r e l a t i o n t o the e n t i r e s t u d y a r e a ( i . e . a l l h a b i t a t s a v a i l a b l e ) . The l a t t e r was a c h i e v e d by g r a p h i n g t h e p e r c e n t f e c a l m a t e r i a l i n r e l a t i o n t o the d i s t a n c e a l o n g t h e t r a n s e c t s e v e r y 0.5 km, t o g i v e an i d e a o f t h e s p a t i a l s e p a r a t i o n between t h e s p e c i e s . T h i s a p p r o a c h i s i m p o r t a n t i n c l a r i f y i n g the l e v e l s o f i n t e r a c t i o n . A l t h o u g h cows and h o r s e s b o t h may use meadows as a h a b i t a t t y p e i n exc e s s o f a v a i l a b i l i t y , i f t h e y d i d not use t h e same i n d i v i d u a l meadows, the n the l e v e l s o f i n t e r a c t i o n would be markedly reduced. Depending on t h e i n d i v i d u a l t r a n s e c t s , a g r e a t e r o r l e s s e r degree o f s p a t i a l o v e r l a p was i n d i c a t e d and the c o n c e n t r a t i o n o f use i n any a r e a v a r i e d w i d e l y between the t h r e e s p e c i e s . The d i s t r i b u t i o n o f h o r s e s , cows and moose a r e shown i n F i g u r e 13. The f r e q u e n c i e s w i t h w h i c h t h e t h r e e s p e c i e s were seen a l o n e , w i t h one o t h e r s p e c i e s , or a l l t o g e t h e r , and when none were p r e s e n t , a t each 0.5 km i n t e r v a l a l o n g the f i v e t r a n s e c t s i s shown i n T a b l e 3. C a l c u l a t i o n s o f t h e c o e f f i c i e n t s o f a s s o c i a t i o n between each p a i r e d c o m b i n a t i o n o f t h e t h r e e s p e c i e s , t o g e t h e r w i t h t h e C h i S q u a r e v a l u e s showed t h e f o l l o w i n g : 1) h o r s e s and cows were a s s o c i a t e d w i t h each o t h e r more t h a n e x p e c t e d by chance (C h c=0.07; X 2=4.4, d f = 1 ) ; 2) h o r s e s and moose were f o u n d t o g e t h e r as o f t e n as one w o u l d e x p e c t by c h a n c e a l o n e ( C n m = 0 . 0 3 ; X 2 = 0 . 3 3 , d f =1); and 3) moose and - 30 - FIGURE 13: T r a n s e c t s I t h r o u g h V, s h o w i n g f r e q u e n c y d i s t r i b u t i o n s o f f e c a l d r o p p i n g s f o r h o r s e s , c a t t l e and moose a t 1 Km i n t e r v a l s a l o n g each t r a n s e c t f o r the June, J u l y , August and September p e r i o d . - 31 - "" a a ŜI5____ . s i > i - __ •____5___ -a -a 9 55555 f - 8 I-a 9 a - 32 - T a b l e 3: C o e f f i c i e n t o f A s s o c i a t i o n f o r h o r s e s . C a t t l e and moose F r e q u e n c i e s w i t h w h i c h e a c h s p e c i e s (C = cows; M = moose; H = h o r s e s ) were seen a l o n e , two t o g e t h e r , a l l t o g e t h e r , o r n o n e , a t e a c h 0.5 km i n t e r v a l on a l l f i v e t r a n s e c t s combined. F r e q u e n c i e s were o b t a i n e d on a p r e s e n c e or absence b a s i s o n l y ; a c t u a l number o f f e c a l p i l e s p e r 0.5 km i n t e r v a l were i g n o r e d . T o t a l # 0.5 km i n t e r v a l s C H M HC HM MC HCM None 201 55 4 14 64 8 17 33 6 - 33 - cows a r e found t o g e t h e r l e s s f r e q u e n t l y t h a n e x p e c t e d (C m c=-0.51; X 2=17.81, d f = 1 ) . To e s t i m a t e t h e r e l a t i v e abundance o f t h e t h r e e s p e c i e s I u s e d t h e number o f i n d i v i d u a l f e c a l p i l e s r e c o r d e d a l o n g t h e t r a n s e c t s w i t h t h e a v a i l a b l e d a t a t h a t i n d i c a t e h o r s e s , c a t t l e , and moose a l l have an average d e f e c a t i o n r a t e o f 15 t o 16 e l i m i n a t i o n s p e r 24 h o u r p e r i o d (moose: Franzman §_£ a l . , 1976; M i q u e l l e , 1983; cows: H a f e z , 1969; h o r s e s : p e r s . o b s . ) . From t h e s e d a t a t h e r e l a t i v e p r o p o r t i o n s were 89.7 p e r c e n t c a t t l e , 7.4 p e r c e n t h o r s e s and 2.9 p e r c e n t moose (Table 8 ) . I n o r d e r t o compare t h e 'long-term* use o f t h e a r e a f o r t h e t h r e e s p e c i e s w i t h t h e d a t a c o l l e c t e d i n t h i s s t u d y , one T r a n s e c t (V) was u s e d as a ' c o n t r o l . ' N o r m a l l y o n l y f r e s h i n d i c a t i o n s o f a n i m a l u s e , e s t i m a t e d t o be l e s s t h a n s i x days o l d , were r e c o r d e d . However, b e f o r e d a t a c o l l e c t i o n began, T r a n s e c t V was i n v e s t i g a t e d i n d e t a i l . The t o t a l 18.5 km was s c r u t i n i z e d on f o o t and a l l f e c a l m a t e r i a l and o t h e r a n i m a l s i g n s r e c o r d e d w i t h i n 4 m each s i d e o f t h e c e n t e r l i n e . The m a t e r i a l c o u l d r e p r e s e n t a t l e a s t f i v e y e a r s a c c u m u l a t i o n . When t h e s e ' l o n g - t e r m * d a t a were g r a p h e d and compared w i t h t h e g r a p h f o r ' f r e s h * m a t e r i a l c o l l e c t e d o v e r two summers on T-V, t h e two graphs ( i g n o r i n g s c a l e ) appeared t o be e x t r e m e l y s i m i l a r ( F i g u r e s 13 and 1 4 ) . A P a i r e d ' t ' t e s t ( S o k a l and R o h l f , 1969: 2 0 5 ) , w i t h A r c s i n T r a n s f o r m a t i o n (Appendix I I I ) was r u n on t h e t h r e e s e t s o f d a t a , i . e . comparing ' o l d ' and ' f r e s h ' f o r e a c h o f t h e t h r e e s p e c i e s . R e s u l t s showed t h e graphs t o be d i f f e r e n t , b e i n g j u s t i n t o t h e s i g n i f i c a n t range - 34 - FIGURE 14: T r a n s e c t V, s h o w i n g f r e q u e n c y d i s t r i b u t i o n s o f accumulated f e c a l d r o p p i n g s f o r h o r s e s , c a t t l e and moose p r i o r t o r e g u l a r d a t a c o l l e c t i o n f o r t h e s t u d y . - 35 - - 9e - Percent Frequency of Observation o 3 ro —t- fD —} in KJ o i (jn L_ C O O i C O cn i ////// yrvr// // // // // // SSSS3 Y // // // /T7m // // // // JZ\ ////// A N J U J In 65 V // // // //H 0 0 BHSlfe. 7/ / / / / / / //] K J . O f o r a l l t h r e e s e t s o f d a t a : cows v s . cows ( t = 2.35, 36 d . f . ) , h o r s e s v s . h o r s e s ( t = 2.51) and moose v s . moose ( t = 2.68) (Appendix I I I ) . 3.3 H a b i t a t Use The p e r c e n t o f e a c h h a b i t a t t y p e a l o n g e a c h t r a n s e c t , b a s e d on t h e t o t a l d i s t a n c e f o r a l l t r a n s e c t s , i s shown i n T a b l e 4. A k-Independent C h i Square T e s t showed t h a t a s i g n i f i c a n t d i f f e r e n c e ( X 2 = 18,020.7, d . f . = 24) e x i s t s among the t r a n s e c t s i n r e g a r d t o t h e r e l a t i v e amount o f each h a b i t a t t y p e t h e y c o n t a i n e d (Table 4 ) . The numbers of p l a n t s p e c i e s found i n each h a b i t a t v a r i e d c o n s i d e r a b l y between h a b i t a t s . , The maximum number o f s p e c i e s (58) was found i n d r y meadow h a b i t a t , and t h e minimum number (26) was found i n young homogenous s e r a i h a b i t a t . The p e r c e n t o f each h a b i t a t t y p e measured on each t r a n s e c t o b t a i n e d by d i r e c t measurement a l o n g t h e e n t i r e l e n g t h o f e a c h t r a n s e c t , p r o v i d e s an e s t i m a t e of h a b i t a t a v a i l a b i l i t y f o r t h e e n t i r e s t u d y a r e a when combined f o r a l l f i v e t r a n s e c t s ( T a b l e s 4 and 5 ) . None o f t h e t h r e e h e r b i v o r e s used th e h a b i t a t s i n p r o p o r t i o n t o h a b i t a t a v a i l a b i l i t y ( T a b l e 5 ) . B o t h h o r s e s and c a t t l e used meadow h a b i t a t d i s p r o p o r t i o n a t e l y more, and c a t t l e a l s o i n d i c a t e d a p r e f e r e n c e f o r i n t e r f a c e z o n e , w h i l e moose u s e d open f o r e s t , c l o s e d f o r e s t and shrub c a r r d i s p r o p o r t i o n a t e l y more. These d i f f e r e n c e s i n h a b i t a t s e l e c t i o n b e t w e e n t h e t h r e e s p e c i e s were s i g n i f i c a n t (Table 6) , i n d i c a t i n g s p e c i e s s p e c i f i c s e l e c t i o n f o r t h e a v a i l a b l e h a b i t a t s . - 37 - TABLE 4: The p e r c e n t * d i s t r i b u t i o n o f h a b i t a t t y p e a l o n g t h e f i v e t r a n s e c t s . Numbers i n p a r e n t h e s e s a r e a c t u a l m e t e r s o f t r a n s e c t c o v e r e d by each h a b i t a t t y p e , on each t r a n s e c t . T r a n s e c t Open F** Semi- Open F C l o s e d F Meadow Shrub c a r r I n t e r - f a c e zone Other I 2.3 (2280) 8.9 (8880) 3.1 (3060) 5.7 (5730) 0.0 (0.0) 0.2 (150) 0.2 (240) I I 0.7 (720) 8.0 (7980) 3.9 (3870) 3.9 (3930) 0.0 (0.0) 1.0 (1020) 0.0 (0.0) I I I 4.0 (4020) 6.9 (6870) 3.3 (3330) 7.3 (7260) 0.8 (750) 1.5 (1530) 1.1 (1050) IV 1.8 (1740) 6.0 (6000) 1.2 (1170) 6.7 (6660) 0.3 (300) 2.4 (2430) 0.4 (420) V 5.7 (5730) 3.6 (3600) 2.2 (2220) 4.4 (4410) 1.56 (1560) 0.6 (570) 0.2 (210) •Based on t o t a l d i s t a n c e f o r a l l t r a n s e c t s ( i . e . 99,690m) **F = Forest - 38 - TABLE 5: H a b i t a t use v e r s u s p e r c e n t H a b i t a t A v a i l a b i l i t v 1. T o t a l f e c a l c o u n t s f o r a l l t r a n s e c t s . per h a b i t a t type f o r h o r s e s , cows and moose 2. C h i s q u a r e r e s u l t s f o r h a b i t a t use v e r s u s h a b i t a t a v a i l a b i l i t y f o r a l l t r a n s e c t s f o r the i n d i v i d u a l s p e c i e s , h o r s e s , cows and moose. 3. P o p u l a t i o n e s t i m a t e s b a s e d on e q u a l a v e r a g e d e f e c a t i o n f o r a l l t h r e e s p e c i e s . r a t e s H a b i t a t Type Open F Semi- Open F C l o s e d F Meadow Shrub c a r r I n t e r - f a c e zone Other X 2 R e l a t i v e P o p u l a t i o n E s t i m a t e s % A v a i l a b l e 14.5 33.4 13.7 28.1 2.6 5.7 1.9 100% 6 d . f . T o t a l 28 100 32 331 7 15 2 515 Horses 336.7* 7.4% % 5.4 19.4 6.4 64.2 1.4 2.9 0.4 T o t a l 289 551 96 4739 103 392 46 6216 Cows 7402.8* 89.7% % 4.6 8.9 1.5 76.2 1.7 6.3 0.7 c o n t i n u e d T a b l e 5 (cont.) H a b i t a t Type Open F Semi- Open F C l o s e d F Meadow Shrub c a r r I n t e r - f a c e zone Other X2 R e l a t i v e P o p u l a t i o n E s t i m a t e s % A v a i l a b l e 14.5 33.4 13.7 28.1 2.6 5.7 1.9 100% 6 d . f . T o t a l 31 37 84 31 13 4 0.0 200 Moose 161.9* 2.9% % 15.5 18.5 42.0 15.5 6.5 2 0.0 T o t a l f e c a l c o u n t : 6931 ^ ^ \ k = 7 TABLE 6 k-Independent C h i Square T e s t f o r comparing H a b i t a t Use Between S p e c i e s , s h o w i n g s i g n i f i c a n t d i f f e r e n c e s i n use p a t t e r n s f o r a l l t h r e e s p e c i e s . H a b i t a t % Open F. 14.5 S-Open F. 33.4 C l . F. 13.7 Meadow 28.1 Shrub C. 2.6 I.Z. 5.7 Other 1.9 F e c a l Count T o t a l s X 2 d.f.=6 Cow 289 4.6% 551 8.9% 96 1.5% 4739 76.2% 103 1.7% 392 6.3% 46 .7% 6216 129.0* Horse 28 5.4% 100 19.4% 32 6.4% 331 64.2% 7 1.4% 15 2.9% 2 .4% 515 T o t a l : 317 651 128 5070 110 407 48 6731 Cow 289 551 96 4739 103 392 46 6216 1327.71* Moose 31 15.5% 37 18.5% 84 42.0% 31 15.5% 13 6.5% 4 2% - 200 T o t a l : 320 588 180 4770 116 396 46 6416 c o n t i n u e d . . . T a b l e 6 (cont) H a b i t a t % Open F. 14.5 S-Open F. 33.4 C l . F. 13.7 Meadow 28.1 Shrub C. 2.6 I.Z. 5.7 Other 1.9 F e c a l Count T o t a l s X 2 d . f . = 6 Horse 28 100 32 331 7 15 2 515 216.1* Moose 31 37 84 31 13 4 - 200 T o t a l : 59 137 116 362 20 19 2 715 3.4 D i e t A n a l y s i s From t h e f o u r c o m p o s i t e f e c a l s a m p l e s c o l l e c t e d e a c h month f o r b o t h cows and horses-summer, and from t h e two samples f o r h o r s e s - w i n t e r , a v e r a g e v a l u e s were d e t e r m i n e d f o r t h e p r e s e n c e o f t h e v a r i o u s p l a n t s p e c i e s i n t h e d i e t each month ( T a b l e 7 ) . The r e s u l t s o f t h e m o n t h l y f e c a l a n a l y s e s show t h a t a t l e a s t 54 d i f f e r e n t p l a n t s p e c i e s were u s e d by the two h e r b i v o r e s (Table 7 ) . U s i n g an a r b i t r a r i l y chosen minimum v a l u e o f 80 p e r c e n t o f t h e t o t a l d i e t , 11 p l a n t s p e c i e s were f o u n d t o c o n s t i t u t e 80% o f t h e c a t t l e ' s and 86 p e r c e n t o f t h e h o r s e s d i e t s (Table 8 ) . As a t o t a l o f a t l e a s t 148 p l a n t s p e c i e s were i d e n t i f i e d on t h e s t u d y a r e a (Appendix I ) , t h i s i n d i c a t e s t h a t a t l e a s t 80 p e r c e n t o f t h e two h e r b i v o r e s * d i e t s were composed o f 7.4 p e r c e n t o f t h e a v a i l a b l e s p e c i e s , t h u s i n d i c a t i n g a r e l a t i v e l y h i g h degree o f s e l e c t i v i t y f o r p l a n t s p e c i e s . I t s h o u l d be n o t e d t h a t t h e s e were t h e same 11 s p e c i e s f o r b o t h h e r b i v o r e s , a l t h o u g h t h e p r o p o r t i o n s i n w h i c h t h e y were used d i f f e r e d between t h e two h e r b i v o r e s and among months (Table 8 ) . An A n a l y s i s o f V a r i a n c e w i t h A r c s i n T r a n s f o r m a t i o n (Table 8) showed t h a t t h e a v e r a g e c o n s u m p t i o n ( i . e . c o n c e n t r a t i o n o f u se) by b o t h h o r s e s and c a t t l e v a r i e d s i g n i f i c a n t l y from month t o month f o r the f o l l o w i n g s i x f o r a g e s : C a l a m a q r o s t i s r u b e s c e n s . Poa i u n c i f o l i a , S t i p a o c c i d e n t a l i s , C a l a m a q r o s t i s n e g l e c t a , S t i p a c o m a t a , and Hordeum i u b a t u m . F u r t h e r , i t was found t h a t h o r s e s always used C_. rubescens more th a n c a t t l e , b u t i t s average use by h o r s e s peaked i n August and t h e n d e c l i n e d , - 43 - TABLE 7: Horse and C a t t l e D i e t s e s t i m a t e d from f e c a l samples c o l l e c t e d i n J u n e , J u l y , A u g u s t and S e p t e m b e r . P l a n t s p e c i e s use based on p e r c e n t p r e s e n c e i n f e c e s : averages from 4 composite f e c a l samples per a n i m a l s p e c i e s p e r month. - 44 - PLANT SPECIES Grasses June H C 1. • C a l a m a g r o s t i s rubescens 18.55 11.25 2. *Poa j u n c i f o l i a 17.75 10.6 3. * S t i p a comata 7.73 4.48 4. *Hordeum jubatum 5.6 7.65 5. * C a l a m a g r o s t i s n e g l e c t a 4.78 8.0 6. • G l y c e r i a b o r e a l i s 2.75 4.48 7. S t i p a r i c h a r d s o n i i 1.4 .93 8. Agropyron s p. 1.28 .18 9. Muhl e n b u r g i a r i c h a r d s o n i s 1.03 .9 10. O r y z o p s i s a s p e r i f o l i a 1.0 .48 11. * S t i p a o c c i d e n t a l i s .8 1.33 12. Poa i n t e r i o r .3 13. A l o p e c u r u s a e g u a l i s .23 14. A g r o s t i s s p. .13 15. S p a r t i n a g r a c i l i s .08 .68 16. Beckmannia sp. .08 17. U n i d e n t i f i e d g r a s s e s TOTAL GRASSES 63.59 50.96 64% 51% Rushes /Sedcres 18. •Juncus b a l t i c u s 10.6 5.15 19. *Carex a q u a t i l i s 9.1 11.73 20. *Carex r o s t a t a 7.7 9.5 21. Carex a t h e r o i d e s 4.3 3.58 22. *Carex c o n c i n o i d e s 2.18 7.1 MONTHS (% d i e t ) Ju ly August September W i n t e r H C H C H C H 15.83 10.08 42.08 21.18 30.0 23.63 7.0 9.05 2.83 4.0 6.88 7.63 3.53 1.65 6.1 4.88 1.43 5.75 1.83 4.3 3.95 4.4 7.13 2.5 2.15 2.4 4.18 6.23 9.23 1.3 3.2 5.48 1.0 2.8 3.6 8.28 1.8 3.75 3.33 4.05 6.15 1.53 .53 .38 .65 .6 .83 1.3 .13 1.3 1.3 .6 1.6 .83 .85 1.5 .4 2.08 6.1 6.55 6.53 4.83 5.95 4.88 5.0 .95 .25 .13 .28 .2 .18 .35 .65 .3 1.08 1.43 .5 8.15 56.7 52.6 62.08 51.37 60.23 50.28 34.7 57% 53% 62% 51% 60% 50% 35% 4.25 2.45 6.2 4.88 4.25 5.23 17.15 15.35 19.95 9.63 11.28 8.78 21.25 12.35 7.13 9.15 9.23 6.18 10.38 5.38 16.15 2.1 1.6 2.7 2.18 .4 .35 3.13 2.33 5.23 4.45 5.0 3.83 1.15 c o n t i n u e d . PLANT SPECIES Grasses June H C 23. E l e o c h a r i s s p. .25 24. U n i d e n t i f i e d rushes/sedges 1.3 5.73 TOTAL RUSHES/SEDGES 35.43 42.79 35% 43% •Dominant 11 f o r a g e s : Other ( F o r b s : Shrubs: Trees) 25. A s t r a g a l u s m i s e r .08 .38 26. P i n u s c o n t o r t a 27. Rosa a c i c u l a r i s .08 28. P i c e a s i t c h e n s i s 29. •Fern' 30. A r t e m e s i a f r i g i d a 31. A r c t o s t a p h y l o s u v a - u r s i 32. P o t e n t i l l a h i p p i a n a .2 1.48 33. S h e p h e r d i a c a n a d e n s i s 34. V i c i a s p . 35. A c h i l l e a m i l l e f o l i u m .1 .68 36. S a l i x spp. 37. S i s y r i n c h i u m a n g u s t i f o l i u m .3 .18 38. Geum t r i f l o r u m .13 .33 39. H i p p u r i s v u l g a r i s .08 .43 40. A s t e r c o n s p i c u o u s .08 .2 MONTHS (% d i e t ) J u l y August September W i n t e r H C H C H C H 7.48 .95 6.03 1.88 1.78 1.48 .23 .15 .95 40.39 41.5 40% 42% 34.87 30.75 35% 31% 30.52 36.19 47.75 31% 36% 48% .15 .38 1.6 .35 1.63 2.58 .45 .55 .73 4.4 .58 .88 1.38 .15 .6 3.2 1.95 .3 1.25 .25 1.0 .4 .15 .4 .15 .33 1.0 .25 .98 .98 .68 .5 1.65 .18 .98 .63 .2 1.8 .35 .58 .35 .93 .18 1.43 .28 .73 1.13 .5 .98 .3 .18 .8 .25 .03 .23 .13 c o n t i n u e d . PLANT SPECIES Grasses June H C 41. Taraxacum o f f i c i n a l e .05 .08 42. F r a g a r i a v i r g i n i a n a .45 43. S p i r a e a b e t u l i f o l i a 44. A n t e n n a r i a p a r v i f o l i a .1 45. A n t e n n a r i a s p . 46. Eriogonum sp. 47. Sedum l a n c e o l a t u m .08 48. J u n i p e r u s sp. 49. B e t u l a g l a n d u l o s a 50. A r n i c a c o r d i f o l i a .28 , 51. S o l i d a g o s p a t h u l a t a .38 52. V i o l a adunca .23 -j 53. C e r a s t i u m arvense .15 , 54. Heuchera c y l i n d r i c a .08 55. Penstemon p r o c e r u s .08 56. 'mosses* 57. T h a l i c t r u m s p . 58. U n i d e n t i f i e d Forbs .13 59. U n i d e n t i f i e d Shrubs TOTAL FORBS; SHRUBS; TREES 1.02 5.8 1% 6% MONTHS J u l y H C .13 .13 .38 .25 .3 .2 .05 .25 .13 .48 .9 .38 (% d i e t ) August September W i n t e r H C H C H .43 .23 .33 .63 .75 .08 .68 .23 .08 .5 .18 .58 .15 .15 .08 .1 .8 .08 .38 .13 .55 .38 .23 .28 98 5.43 2.18 1.68 .63 2.8 3.02 6.08 3% 6% 3.24 17.5 3% 18% 9.17 13.88 17.4 9% 14% 17% w h e r e a s c a t t l e u se o f C. r u b e s c e n s i n c r e a s e d s t e a d i l y o v er t h e summer and peaked i n September. There was a l s o a s i g n i f i c a n t d i f f e r e n c e i n p a t t e r n o f use by h o r s e s and c a t t l e f o r t h e p l a n t s p e c i e s Carex a q u a t i l i s , Poa i u n c i f o l i a , Juncus b a l t i c u s > Hordeum -iubatum. and G l v c e r i a b o r e a l i s . F i n a l l y , i n o n l y t h r e e s p e c i e s , i _ a i u n c i f o l i a . Juncus b a l t i c u s and S t i p a comata. was t h e d i f f e r e n c e i n use b e t w e e n h o r s e s and c a t t l e f o u n d t o be s i g n i f i c a n t l y d e p e n d e n t on a p a r t i c u l a r month (Table 8 ) . I t s h o u l d be noted t h a t f o r b o t h C. rubescens o r , C_. aqua- t i l i s . month was n o t i m p o r t a n t i n an i n t e r a c t i o n c o n t e x t , b e c a u s e h o r s e s a l w a y s a t e more o f t h e f o r m e r t h a n c a t t l e , and l e s s o f t h e l a t t e r . C o n s i d e r i n g t h e c o n s i s t e n t l y h i g h p r o p o r t i o n o f t h e 11 f o r a g e s i n the d i e t s o f b o t h h o r s e s and cows, the t o t a l number o f p l a n t s p e c i e s a c t u a l l y e a t e n by t h e two h e r b i v o r e s p e c i e s was c o n s i d e r e d i n r e l a t i o n t o how many p l a n t s p e c i e s were a v a i l a b l e t o them. Of 88 f o r b s ( A p p e n d i x I) i d e n t i f i e d i n t h e s t u d y a r e a h o r s e s u t i l i z e d a p p r o x i m a t l y 16, and c a t t l e 20 ( T a b l e 7 ) . Reduced d e p e n d e n c e on f o r b s was r e c o r d e d i n w i n t e r and J u n e h o r s e d i e t s . The p o r t i o n o f f o r b s i n ho r s e d i e t s was as f o l l o w s : W i n t e r = 2.75%, June = 1.02%, J u l y = 4.69%, August = 3.01%, and September = 7.53%. Of 43 s p e c i e s of g r a s s e s and sedges i d e n t i f i e d (Appendix I ) , h o r s e s u t i l i z e d 22 and c a t t l e o n l y 19 (Table 7 ) . A f a c t o r f u r t h e r i n d i c a t i v e o f s e l e c t i v i t y i s found i n the use o f t h e sedges Carex a q u a t i l i s and Q_. r o s t a t a . Together t h e y f o r m e d a s i g n i f i c a n t p o r t i o n o f t h e d i e t o f h o r s e s (X = 19%) and c a t t l e (X = 2 4 % ) , b u t were f o u n d o n l y i n sedge - 48 - TABLE 8 A v e r a g e use per month of the 11 forages composing at l e a s t 80% of the d i e t s of both horses and cows, with an analysis of variance (using transformed data) comparing the 11 plant species for u t i l i z a t i o n per month, between species and on an in t e r a c t i o n basis: P l a n t S p e c i e s June July August September xa FMonth F S p e c i e s i n t e r a c t i o n C a l a m a g f o s t i s H 18.6 + 5.7 15.8 + 5.5 42.1 + 5.9 30.0 + 19.2 26.6 + 15.9 6.44* 8.11* 1.12 rubescens C 11.3 ± 3.2 10.2 ± 1.4 21.2 ± 8.3 23.6 + 5.3 16.5 ± 7.70 Carex a q u a t i l i s H 9.1 + 2.3 15.4 + 8.0 9.6 ± 7.1 8.8 + 1.5 10.7 ± 5.7 2.61 5.82* 1.25 C 11.7 ± 3.7 20.0 ± 2.8 11.3 ± 6.4 21.3 ± 11.4 16.1 ± 7.8 Ppa H 17.8 + 3.9 9.1 + 4.9 4.0 + 2.3 7.6 + 1.1 9.6 + 6.0 16.2* 12.1* 4.54* juncifoiUa C 10.6 ± 2.5 2.8 ± 1.9 6.9 ± 3.6 3.5 ± 2.2 5.9 ± 4.0 Carex rostata H 7.7 + 2.4 7.1 + 2.3 9.2 + 2.6 10.4 ± 4.2 8.6 + 3.0 .108 .80 2.23 C 9.5 + 2.2 9.2 ± 5.3 6.2 ± 3.2 5.4 ± 3.3 7.6" ± 3.8 Juncus b a l t i c u s H 10.6 + 4.7 4.3 ± 2.9 6.2 ± 1.2 4.3 + 2.1 6.4 ± 2.9 1.88 7.88* 3.63* C 5.2 + 1.9 2.5 ± 1.4 4.9 ± 2.6 5.2 ± 1.5 5.2 ± 1.9 continued... T a b l e 8 (cont) P l a n t S p e c i e s June J u l y August September F M o n t h F S p e c i e s F I n t e r a c t i o n S t i o a H + 0.8 1.1 + 6.1 4.6 + 6.5 2.9 + 6.0 2.8 + 4.8 3.7 6.15* .1 .53 o c c i d e n t a l i s C + 1.3 1.5 + 6.6 2.2 + 4.8 2.3 + 4.9 1.7 + 4.5 2.5 C a l a m a a r o s t i s H + 4.8 4.8 + 6.2 3.3 + 1.3 .6 + 5.5 4.5 + 4.4 3.8 5.61* .72 2.78 n e a l e c t a C + 8.0 2.5 + 9.2 3.3 + 3.2 1.8 + 1.0 1.2 + 5.4 4.1 S t i p a H + 7.7 2.3 + 6.1 1.7 + 1.4 1.7 + 1.8 1.0 + 4.3 3.2 5.04* .83 6.99* comata C + 4.5 0.5 + 4.9 2.1 + 5.8 3.0 + 4.3 1.0 + 4.9 1.8 Carex H + 2.2 1.6 + 3.1 2.6 + 5.2 5.4 5.0 3.6 + 3.9 3.5 .69 .21 1.56 c o n c i n o i d e s C + 7.1 4.2 + 2.3 1.4 + 4.5 2.6 + 3.8 2.9 + 4.4 3.2 Hordeum H + 5.6 2.9 + 4.4 2.2 + 2.5 1.6 + 2.4 1.3 + 3.7 2.3 9.9* 5.8* 1.04 jubatum C + 7.7 1.7 + 7.1 2. + 2.2 1.1 + 4.2 1.1 + 5.3 2.7 G l y c e p i a H + 2.8 2.0 + 3.6 0.6 + 1.8 0.2 + 3.3 1.4 + 3.0 1.7 2.91 7.04* 1.17 b o r e a l i s C + 4.5 2.3 + 8.3 1.7 + 3.8 3.3 + 4.1 1.4 + 4.9 3.2 a5T = O v e r a l l mean f o r 4 months. meadows w h i c h r e p r e s e n t a p p r o x i m a t e l y 15% o f t h e t o t a l a r e a , and were i n s c a t t e r e d l o c a t i o n s . Other p l a n t s p e c i e s which f o r m e d a d o m i n a n t p o r t i o n o f t h e d i e t s o f b o t h h o r s e s and c a t t l e were f o u n d o n l y i n d r y meadows ( i . e . S t i p a c o m a t a . S_. o c c i d e n t a l i s ) . or i n sedge meadows o n l y ( i . e . C. n e o l e c t a . C_. a t h e r o i d e s . and G l y c e r i a b o r e a l i s ) (Appendix I V ) . A l t h o u g h a d i s p r o p o r t i o n a t e l y h i g h use o f meadow h a b i t a t was i n d i c a t e d f o r b o t h h o r s e s and c a t t l e (Table 5) , a s m a l l number o f the t o t a l p l a n t s p e c i e s a v a i l a b l e i n wet and d r y meadows a c t u a l l y formed a dominant p o r t i o n o f t h e d i e t s o f e i t h e r a n i m a l s p e c i e s (Appendix IV, T a b l e 7 ) . The p r e s e n c e o f E l e o c h a r i s sp.t, i n t h e d i e t s , p o s i t i v e l y i d e n t i f i e d by t h e W.S.U. l a b o r a t o r y , appears t o be the most o u t s t a n d i n g example i n d i c a t i v e o f s e l e c t i v i t y . Based on the p e r c e n t o c c u r r e n c e i n horse d i e t s , i t was a r e a s o n a b l y s i g n i f i c a n t f o r a g e p l a n t ( X ^ s u m m e r = 2.77%. T a b l e 7 ) . However, i t was never a c t u a l l y i d e n t i f i e d on the s t u d y s i t e and i t i s assumed t h a t E l e o c h a r i s S J D v makes up a r e l a t i v e l y s m a l l p r o p o r t i o n o f the v e g e t a t i o n c o v e r i n t h e s t u d y a r e a , o r i s e x t r e m e l y l o c a l i z e d . The o v e r a l l , a v e r a g e number o f p l a n t s p e c i e s used by h o r s e s was found t o be l e s s i n w i n t e r t h a n summer ( i . e . H orse- summer X = 29; H o r s e - w i n t e r X" = 21; T a b l e 7 ) . The b o t a n i c a l g r o u p most d r a s t i c a l l y r e d u c e d was t h e g r a s s e s ( T a b l e 9 ) . In t h e summer h o r s e s used a maximum o f 16 and a monthly average o f 13 g r a s s s p e c i e s , w h i l e t h i s was d e c r e a s e d t o s e v e n i n t h e w i n t e r ; a r e d u c t i o n o f almost h a l f . I t s h o u l d be noted t h a t t h e c a t e g o r y o f " u n i d e n t i f i e d g r a s s e s " i s assumed not t o be 'new' g r a s s e s , but r a t h e r 'known' g r a s s e s i n an u n i d e n t i - - 51 - TABLE 9: Grass s p e c i e s e a t e n by Horses-summer v e r s u s H o r s e s - w i n t e r . showing o r d e r based on p e r c e n t use , and t h e r e d u c t i o n o f s p e c i e s i n t h e w i n t e r d i e t . Summer Wi n t e r Order g r a s s s p e c i e s (%) Order g r a s s s p e c i e s (%) Y X 1. C a l a m a g r o s t i s 26.6 1. U n i d e n t i f i e d g r a s s 8.2 rubescens 2. C a l a m a g r o s t i s 7.0 2. Poa j u n c i f o l i a 9.6 rubescens 3. S t i p a 4.8 3. G l y c e r i a b o r e a l i s 6.2 o c c i d e n t a l i s 4. S t i p a o c c i d e n t a l i s 5.0 4. C a l a m a g r o s t i s 4.4 5. S t i p a comata 4.0 n e g l e c t a 6. C a l a m a g r o s t i s 2.8 5. S t i p a comata 4.3 n e g l e c t a 6. Hordeum jubatum 3.7 7. Poa j u n c i f o l i a 1.7 7. G l y c e r i a b o r e a l i s 3.0 T o t a l : 34.9 S u b - t o t a l : 56.4 8. O r y z o p s i s 1.0 a s p e r i f o l i a 9. S t i p a .8 r i c h a r d s o n i i 10. M u h l e n b u r g i a .8 r i c h a r d s o n i i 11. U n i d e n t i f i e d g r a s s .7 12. A g r o pyron sp. .3 13. S p a r t i n a g r a c i l i s .3 14. Beckmannia spp. .2 15. Poa i n t e r i o r .1 16. A l o p e c u r u s .1 17. A g r o s t i s s p . .1 TOTAL: 60.8 - 52 - f i a b l e c o n d i t i o n (Bruce D a v i t t , P e r s . comm., W.S.U. W i l d l i f e H a b i t a t Lab, 1981). The same c o r e - g r o u p o f f i v e rushes-sedges made up t h e b u l k o f t h a t b o t a n i c a l g r o u p i n g f o r b o t h w i n t e r and summer d i e t s o f h o r s e s (Table 1 0 ) . However, t h e i n c r e a s e d amount o f use o f t h e s e f i v e s p e c i e s d u r i n g t h e w i n t e r was n o t a b l e , i n c r e a s i n g from 33.2 p e r c e n t i n summer t o 48.0 p e r c e n t i n w i n t e r o f t h e t o t a l d i e t (Table 1 0 ) . There i s a p o s s i b i l i t y t h a t sedges r e t a i n a h i g h e r n u t r i e n t c o n t e n t i n t h e w i n t e r t h a n some g r a s s e s , because o f the p r e s e n c e o f g r e e n m a t e r i a l i n t h e c e n t e r o f sedge p l a n t s . A n u t r i e n t a n a l y s i s c a r r i e d o u t on se d g e m a t e r i a l c o l l e c t e d d u r i n g t h e w i n t e r o f 1979-80, showed t h a t a sample from whole ungrazed sedge p l a n t s had a c r u d e p r o t e i n (CP) l e v e l o f 3.78 p e r c e n t ( d r y b a s i s ) , w h i l e the s e l e c t e d green m a t e r i a l from the i n n e r p a r t s o f t h e p l a n t s had a CP v a l u e o f 6.5 p e r c e n t ( F i g u r e s 15 and 16; Appendix V ) . An i n c r e a s e d c o n c e n t r a t i o n o f shrubs was noted i n horse d i e t s from s p r i n g t o w i n t e r ; where June = 0.0%, J u l y = 0.58%, A u g u s t = 0.23%, Sept e m b e r = 2.96%, and w i n t e r = 14.65%. C o n i f e r n e e d l e s o c c u r r e d o n l y i n Sept e m b e r (0.55%) and i n w i n t e r (6.25%) d i e t s . B a s e d on t h e p r o p o r t i o n o f t o t a l c o v e r c o u n t s i n t h e f e c a l a n a l y s i s ( S e c t i o n 2 . 6 ) , t h e c o n c e n t r a t i o n o f use was compared f o r t h e t h r e e b o t a n i c a l groups: g r a s s e s , rushes-sedges and ' o t h e r ' ( i . e . f o r b s , t r e e s and shrubs) ( F i g u r e 1 7 ) , u s i n g t h e f o l l o w i n g Z - p r o p o r t i o n a l f o r m u l a ( W a l p o l e , 1982: 266-74): - 53 - TABLE 10: Rush and sedge s p e c i e s w i n t e r , s h o w i n g i n c r e d i e t . Summer Order sedge s p e c i e s (%) X 1. Carex a q u a t i l i s 10.7 2. Carex r o s t a t a 8.6 3. Juncus b a l t i c u s 7.4 4. Carex c o n c i n o i d e s 3.9 5. E l e o c h a r i s spp. 2.6 S u b - t o t a l : 33.2 6. Carex a t h e r o i d e s 2.4 7. U n i d e n t i f i e d Rush-sedges .6 TOTAL: 36.2 n by Horses i n summer v e r s u s use o f sedges i n the w i n t e r W i n t e r Order sedge s p e c i e s (%) X 1. Juncus b a l t i c u s 17.2 2. Carex r o s t a t a 16.2 3. Carex a q u a t i l i s 12.4 4. Carex c o n c i n o i d e s 1.2 5. E l e o c h a r i s spp. 1.0 TOTAL: 48.0 - 54 -  FIGURE 17: The p e r c e n t o c c u r r e n c e o f t h e t h r e e major b o t a n i c a l groups i n t h e d i e t s o f h o r s e s and c a t t l e based on f e c a l a n a l y s i s . - 56 -  where: n-̂  and n 2 = 800* <32 = 1 ~ D 2 * n H o r s e - w i n t e r = 400 The p r o p o r t i o n a l use o f t h e t h r e e b o t a n i c a l groups by h o r s e s and c a t t l e proved t o be s i g n i f i c a n t l y d i f f e r e n t , e x c e p t i n t h e i n s t a n c e s o f g r a s s e s i n J u l y , and r u s h e s - s e d g e s i n J u l y and August (Table 1 1 ) . The p r o p o r t i o n a l use o f t h e t h r e e g r o u p s was s i g n i f i c a n t l y d i f f e r e n t i n h o r s e s between summer and w i n t e r d i e t s (Table 1 1 ) . - 58 - TABLE 11: Z - V a l u e s f o r d i f f e r e n c e s i n p r o p o r t i o n a l u se o f b o t a n i c a l groups on a s e a s o n a l b a s i s : ( Grasses) (R-Sedges) (Other) CATTLE v s . HORSES June J u l y August September 5.2* 1.6 4.4* 4.0* 3.2* 0.8 1.7 2.1* 5.5* 3.0* 10.0* 3.13* HORSES summer v s . v s . 8.8 4.3 6.5 HORSES w i n t e r *Z g r e a t e r t h a n 1.96 = S i g n i f i c a n t - 59 - Chapter 4: D i s c u s s i o n 4.1 H a b i t a t - u s e r e l a t e d t o a n i m a l p e r c e p t i o n o f a v a i l a b i l i t y The t r a n s e c t s y s t e m u s e d p r o v i d e d a r e l i a b l e e s t i m a t e o f t h e h a b i t a t a v a i l a b i l i t y f o r t h e e n t i r e s t u d y a r e a . S i n c e t h e IDF (b) s u b z o n e i s c l a s s i f i e d as f o r e s t e d range, i t i s n o t s u r p r i z i n g t h a t o f t h e h a b i t a t a v a i l a b l e , 67.3 p e r c e n t was f o r e s t e d , i n c l u d i n g h a b i t a t t y p e s open f o r e s t , semi-open f o r e s t , c l o s e d f o r e s t and i n t e r f a c e zone. However, c o n s i d e r e d f r o m a h a b i t a t - u s e p e r s p e c t i v e , i t i s i n t e r e s t i n g t o n o t e t h a t o n l y one o f t h e t h r e e s p e c i e s s t u d i e d (moose) i s c o n s i d e r e d a t r u e f o r e s t - d w e l l e r (Franzman, 1978). Horses a r e p r i m a r i l y a d a p t e d t o d r y g r a s s l a n d s and c a t t l e t o p a r k l a n d ( G r o v e s , 1974; Camp and S m i t h , 1942; W a l k e r , 1975). However b o t h h o r s e s and c a t t l e have become c o s m o p o l i t a n i n t h e i r d i s t r i b u t i o n under human i n f l u e n c e s . The c r u x o f t h e h a b i t a t e x p l o i t a t i o n o f t h e s e t h r e e s p e c i e s i n a f o r e s t e d e n v i r o n m e n t r e v o l v e s a r o u n d t h e f o l l o w i n g : How a d a p t i v e are h o r s e s and cows t o an e s s e n t i a l l y f o r e i g n e n v i r o n m e n t ? What f o r m does t h e i r a d a p t a t i o n t a k e ( i . e . how s i m i l a r a r e t h e y ) ? How do h o r s e s and cows d i f f e r f r o m moose i n u t i l i z a t i o n o f f o r e s t v e r s u s n o n - f o r e s t h a b i t a t w i t h i n t h e e n v i r o n m e n t ? I n g e n e r a l , i t would seem t h a t moose, t h r o u g h l o n g a s s o c i a t i o n w i t h t h e n o r t h e r n f o r e s t , and an o v e r a l l d i s t r i b u t i o n more c l o s e l y r e l a t e d t o t h e range o f n o r t h e r n t r e e s and shrubs t h a n t o any o t h e r f a c t o r (Coady, 1982), c o u l d be e x p e c t e d t o e x p l o i t f o r e s t e d h a b i t a t s more e f f i c i e n t l y t h a n e i t h e r h o r s e s or c a t t l e . From t h i s p e r s p e c t i v e , i t w o u l d a p p e a r f r o m t h e s t u d y - 60 - r e s u l t s t h a t moose are r e a s o n a b l y p r e d i c t a b l e i n t h a t 78 p e r c e n t o f a l l f e c a l c o u n t s were i n f o r e s t . H o r s e s and cows a l s o a p p e a r somewhat p r e d i c t a b l e i n b e i n g s t r o n g l y a t t r a c t e d t o open a r e a s (meadow and shrub c a r r ) , f e c a l c o n c e n t r a t i o n s b e i n g 65.3 p e r c e n t f o r h o r s e s and 76.9 p e r c e n t f o r cows, a l t h o u g h o n l y 1.4 and 1.7 p e r c e n t , r e s p e c t i v e l y were found i n shrub c a r r . However, h o r s e s appear t o be more i n c l i n e d t o use f o r e s t h a b i t a t t h a n c a t t l e . H o r s e f e c a l c o u n t s i n d i c a t e d a 34.1 p e r c e n t f o r e s t u t i l i z a t i o n and 21.3 p e r c e n t f o r c a t t l e . However, t h e r e i s a h i d d e n f a c t o r . Based on f e c a l c o n c e n t r a t i o n s v e r s u s h a b i t a t a v a i l a b i l i t y , d i e t r e s u l t s ( i . e . h i g h e r use by h o r s e s o f t h e f o r e s t s p e c i e s C a l a m a q r o s t i s r u b e s c e n s ) : and g e n e r a l o b s e r v a t i o n s d u r i n g two summers on the s t u d y s i t e , i t appears t h a t h o r s e s use f o r e s t h a b i t a t s ( i . e . f o r e s t not i n c o n j u n c t i o n w i t h open h a b i t a t s ) a t l e a s t t w i c e as much as cows, a c c o u n t i n g f o r 31.2 p e r c e n t o f a l l h o r s e f e c a l c o u n t s . Cows used f o r e s t h a b i t a t by i t s e l f o n l y 15 p e r c e n t o f t h e t i m e , and t h e i r f e c a l c o n c e n t r a t i o n s were o t h e r w i s e found i n open a r e a s or i n f o r e s t a d j a c e n t t o open a r e a s ( i . e . I n t e r f a c e Z o n e s ) . There i s a d i s t i n c t d i f f e r e n c e i n t h e s e two f o r m s o f f o r e s t e d h a b i t a t use. McLean (1979) a l s o o b s e r v e d t h a t where open a r e a s (e.g. g r a s s l a n d s o r c l e a r c u t s ) a l t e r n a t e w i t h f o r e s t e d a r e a s , as on my s t u d y s i t e , t h e r e i s d i f f i c u l t y i n g e t t i n g c a t t l e t o g r a z e t h e f o r e s t s w i t h o u t o v e r - u s i n g t h e open a r e a s . Such overuse o f open a r e a s was o b s e r v e d t o o c c u r on my s t u d y s i t e . Moose tended t o use t h o s e h a b i t a t t y p e s ( i . e . open, semi-open, c l o s e d f o r e s t and shrub c a r r ) t h a t h o r s e s and, p a r t i c u l a r l y , c a t t l e u s e d t h e l e a s t . I n a d d i t i o n , a l t h o u g h t h e p a t t e r n - 61 - o f h a b i t a t use was more s i m i l a r b e t w e e n h o r s e s and c a t t l e t h a n between h o r s e s o r c a t t l e v e r s u s moose, a l l t h r e e h e r b i v o r e s showed some s p e c i e s - s p e c i f i c s e l e c t i o n f o r t h e a v a i l a b l e h a b i t a t s . T h i s v a r i a b i l i t y i n h a b i t a t s e l e c t i o n p r o d u c e d a degree o f s p a t i a l s e p a r a t i o n o f t h e t h r e e h e r b i v o r e s . S u p p o r t i v e o f t h i s f o r m o f s p a t i a l s e p a r a t i o n were t h e f i n d i n g s , f i r s t o f a l l , t h a t t h e t r a n s e c t s v a r i e d s i g n i f i c a n t l y i n r e g a r d t o t h e r e l a t i v e amount o f e a c h h a b i t a t t y p e and s e c o n d l y , t h e h i g h e s t f e c a l c o u n t s f o r e a c h h e r b i v o r e s p e c i e s were made on d i f f e r e n t t r a n s e c t s : t h e h i g h e s t c o u n t f o r cows was on T-IV, f o r h o r s e s on T - I , and f o r moose on T-V. The g r e a t e s t amount o f b o t h Meadow and I n t e r f a c e Zone, t h e h a b i t a t t y p e s t h a t c a t t l e appeared t o f a v o r , on t h e b a s i s o f use i n p r o p o r t i o n t o a v a i l a b i l i t y , were found on T-IV. I n d i c a t i v e o f t h e freedom w i t h w h i c h h o r s e s a p p e a r t o move t h r o u g h f o r e s t , t h e most c o n t i g u o u s f o r e s t ( i . e . n o t i n c l u d i n g I n t e r f a c e Z o n e s ) , was f o u n d on T - I . T-V had by f a r t h e l a r g e s t amount o f shrub c a r r ? a h a b i t a t t y p e moose used d i s p r o p o r t i o n a t e l y more t h a n i t was a v a i l a b l e , and wh i c h may have a t t r a c t e d them t o t h i s t r a n s e c t . S u b s e q u e n t l y , t h e d a t a d i d c o n t r i b u t e toward an under- s t a n d i n g o f what were f e l t t o be t h e t h r e e major c o n s i d e r a t i o n s i n v o l v e d i n h a b i t a t u s e , i n t h a t e v i d e n c e i n d i c a t e d h o r s e s move about more r e a d i l y i n a f o r e s t e d environment t h a n c a t t l e ( d i s c u s s e d f u r t h e r i n S e c . 4.2); t h a t t h e a d a p t i v e p a t t e r n s o f h o r s e s and c a t t l e a p p e a r t o be somewhat d i f f e r e n t ; and t h a t b o t h h o r s e s and c a t t l e d i f f e r g r e a t l y from moose i n t h e i r use o f h a b i t a t s w i t h i n the envi r o n m e n t . - 62 - T h e r e i s a p o t e n t i a l f o r m i s i n t e r p r e t a t i o n i n d e t e r m i n i n g t h e h a b i t a t t y p e s i n w h i c h a n i m a l s a) s p e n d t h e most t i m e , and b) a r e most a c t i v e ( e.g. t r a v e l l i n g ) ( N e f f , 1968). C a i r n s and T e l f e r ( 1 9 8 0 ) , n o t e d t h a t on A l b e r t a range, b i s o n ( B i s o n b i s o n ) , were most a c t i v e i n u p l a n d g r a s s l a n d s , but s p e n t t h e m o s t t i m e i n a s p e n ( P o p u l u s t r e m u l o i d e s ) . They a l s o n o t e d t h a t moose w e r e most a c t i v e i n f o r e s t , b u t s p e n t t h e most t i m e , p r o b a b l y f e e d i n g and bedding, i n shrubland. I t i s important t o note t h a t i t i s not p o s s i b l e t o c l e a r l y d i f f e r e n t i a t e between t h e s e two u s e - c a t e g o r i e s o n l y on t h e b a s i s o f f e c a l c o u n t s , u n l e s s i t c a n be demonstrated t h a t d e f e c a t i o n s o c c u r p r i m a r i l y i n a s s o c i a t i o n w i t h some o t h e r a c t i v i t y , s u c h as o c c u r r i n g i m m e d i a t e l y a d j a c e n t t o b e d d i n g s i t e s . H o r s e s , moose and c a t t l e , on t h e b a s i s o f f e c a l c o u n t s i n t h i s s t u d y , a l l s e l e c t e d f o r p a r t i c u l a r h a b i t a t t y p e s i n d i f f e r e n t w ays, b u t i t i s not p o s s i b l e t o say f o r e x a c t l y what purpose. General observations t h r o u g h o u t t h e p e r i o d o f f i e l d work i n d i c a t e d c a t t l e p r e f e r r e d t o bed i n meadow-margins, and h o r s e s i n open f o r e s t . Moose had a s t r o n g a f f i n i t y t o c l o s e d f o r e s t , w h i c h t h e y used t h r e e t i m e s more t h a n w o u l d have been e x p e c t e d i n p r o p o r t i o n t o i t s a v a i l a b i l i t y , and t e n d e d t o bed i n r a v i n e s w i t h h e a v y s p r u c e c o v e r . An a t t e m p t was made by Stcarrar - et :al>'f 19?7 )^*to f establish:' -;a s. r e l a t i o n s h i p between h a b i t a t s e l e c t i o n and f o r a g e a v a i l a b i l i t y i n f e r a l h o r s e s . U s i n g m u l t i v a r i a t e a n a l y s i s o f f e c a l c o u n t s , a p o s i t i v e r e s p o n s e by h o r s e s t o i n c r e a s i n g p i n e g r a s s cover ( i . e . f o r e s t h a b i t a t ) was i n d i c a t e d . R e s u l t s were i n c o n c l u s i v e , h o w e v e r , due t o l a c k o f d i e t d a t a . S a l t e r and Hudson (1978a; 1979) f e l t t h a t t h e a v a i l a b i l i t y o f f o r a g e was t h e p r i m a r y d e t e r m i n a n t f o r h a b i t a t s e l e c t i o n o f f e r a l h o r s e s i n s o u t h e r n A l b e r t a , and t h a t t h e p r e f e r r e d h a b i t a t s o f h o r s e s were meadow and g r a s s l a n d s . T h i s c o n c l u s i o n i s somewhat c o n f u s i n g , however, s i n c e i n t h e i r s t u d y th e s i n g l e most s i g n i f i c a n t d i e t component o f h o r s e s was a g r a s s s p e c i e s most o f t e n f o u n d i n f o r e s t e d h a b i t a t s . I n my s t u d y , h o r s e s and c a t t l e s e l e c t e d most s t r o n g l y f o r meadow h a b i t a t as shown by t o t a l f e c a l c o u n t s from t r a n s e c t d a t a , b u t f e c a l a n a l y s i s showed t h e most dominant f o r a g e was C. r u b e s c e n s . s t r i c t l y a f o r e s t s p e c i e s . T h i s s u p p o r t s th e f i n d i n g s o f M i l l e r e i a l . , (1981) and o t h e r a u t h o r s ( N e f f , 1968; Leckenby, 1968) t h a t p e l l e t groups do not r e l i a b l y estimate t h e v a l u e o f v a r i o u s p l a n t c o m m u n i t i e s t o a n i m a l s f o r f o r a g e use. A l t h o u g h a d i e t a n a l y s i s was not done on moose, i n t e r p r e - t a t i o n o f h a b i t a t - u s e d a t a b a s e d on f o o d h a b i t s e s t a b l i s h e d by o t h e r r e s e a r c h e r s , a l s o s u p p o r t e d t h e c o n t e n t i o n t h a t f e c a l c o u n t s a r e n o t d i r e c t l y r e l a t e d t o f o r a g e s e l e c t i o n . The h i g h e s t f e c a l c o u n t s (42%) f o r moose were f o u n d i n c l o s e d f o r e s t , t h e h a b i t a t t y p e w i t h t h e l o w e s t p l a n t s p e c i e s v a r i e t y . Moose d i e t s a r e v a r i a b l e , depending on a v a i l a b i l i t y and l o c a t i o n , b u t t h e y t e n d t o use e a r l y s u c c e s s i o n a l woody browse, p a r t i c u l a r l y S a l i x s p p . ( F r a n z m a n , 1978; P e e k , 1 9 7 4 ) , and depend h e a v i l y on a q u a t i c p l a n t s , i f a v a i l a b l e , i n t h e summer ( P e t e r s o n , 1979; C o a d y , 1 9 8 2 ) , n e i t h e r o f w h i c h a r e found t o any e x t e n t i n c l o s e d f o r e s t h a b i t a t . J o h n s o n (1980) warns a g a i n s t m a k i n g a b s o l u t e s t a t e m e n t s r e g a r d i n g usage v e r s u s a v a i l a b i l i t y d a t a , because of the inherent - 64 - . a r b i t r a r i n e s s o f t h e v a l u e s . F o r e x a m p l e , a r b i t r a r i n e s s i n h a b i t a t use c a n be a r e s u l t o f how t h e i n v e s t i g a t o r d e f i n e d t h e h a b i t a t t y p e s , o r t h e f a c t t h a t a n i m a l s have a l r e a d y made a s e l e c t i o n by s i m p l y b e i n g f o u n d i n a p a r t i c u l a r a r e a , o r i n t h e c a s e o f r a n g e c a t t l e , h a v i n g been p l a c e d i n a c e r t a i n d e l i m i t e d a r e a . A d d i t i o n a l a r b i t r a r i n e s s may be i n t r o d u c e d i n t o t h e e q u a t i o n i n t h e case o f f e r a l h o r s e s because o f t h e i r n a t u r a l n i c h e b e i n g t h a t o f open p l a i n s g r a z e r , but showing a s u c c e s s f u l s e c o n d a r y a d a p t a t i o n t o more h e a v i l y f o r e s t e d h a b i t a t s . O n l y two o t h e r s t u d i e s i n N o r t h A m e r i c a have d e a l t w i t h t h e q u e s t i o n o f h a b i t a t use by f e r a l h o r s e s i n f o r e s t e d e n v i r o n m e n t s ( S a l t e r , 1978; S t o r r a r . ; e t al(-1977) .Not• enough data',.- has been c o l l e c t e d t o be a b l e t o understand under what c o n d i t i o n s , and why, h o r s e s s e l e c t p a r t i c u l a r h a b i t a t s . W i t h h a b i t a t a v a i l a b i l t y , s e v e r a l f a c t o r s must be considered s u c h as s e a s o n a l v a r i a b i l i t y , ' r e a l ' v e r s u s 'apparent* f o r a g e a v a i l a b i l i t y , and t h e a b i l i t y o f the h a b i t a t t o meet n u t r i t i o n a l r e q u i r e m e n t s . H a b i t a t s may f l u c t u a t e w i l d l y i n n u t r i t i v e q u a n t i t y and q u a l i t y f r o m s e a s o n t o s e a s o n , o r from y e a r t o y e a r , b u t maintenance needs o f t h e h e r b i v o r e remain r e l a t i v e l y u n i f o r m . F o r example, P r i c e (1978) showed t h a t a c t u a l p r o t e i n i n t a k e , r e l a t i v e t o t h e presumed requirements i n Coke's hartebeest ( A l c e l a p h u s b u s e l a p h u s c o k e i ) , r a n g e d f r o m 148 p e r c e n t i n t h e wet s e a s o n t o o n l y 17 p e r c e n t i n t h e d r y s e a s o n ; t o t a l d i g e s t i b l e n u t r i e n t i n t a k e v a r i e d from 196 percent t o 56 percent. A t l e a s t one i n s t a n c e o f t h e e f f e c t o f c l i m a t e on h a b i t a t a v a i l a b i l i t y was o b s e r v e d d u r i n g my f i e l d s t u d y . I n 1979, when w a t e r l e v e l s i n s e d g e a r e a s , ponds and c r e e k s were v e r y - 65 - l o w o r d r y , a l a r g e number o f wet meadow a r e a s were h e a v i l y g r a z e d and tra m p l e d , by September. In 1980, due t o unse a s o n a b l y h i g h summer r a i n f a l l , and s u b s e q u e n t a b n o r m a l l y h i g h w a t e r l e v e l s , many s e d g e a r e a s were i n a c c e s s i b l e , f o r c i n g a n i m a l s t o use d r y meadow a r e a s and, perhaps f o r e s t , more h e a v i l y . Van V a l e n (1973 i n W h i t e , 1978) wondered how t o r e s o l v e t h e p a r a d o x o f h e r b i v o r e s b e i n g l i m i t e d by t h e i r f o o d i n a g r e e n w o r l d , w h e r e f o o d i s a p p a r e n t l y g r e a t l y u n d e r - u s e d . T h i s i s t h e p a r a d o x o f ' r e a l ' v e r s u s 'apparent* f o r a g e a v a i l - a b i l i t y , and a l s o may a p p l y t o o t h e r f e a t u r e s o f a h a b i t a t w h i c h a f f e c t i t s a t t r a c t i v e n e s s t o a n i m a l s . I t i s not always p o s s i b l e t o e s t i m a t e t h e way i n w h i c h a n i m a l s w i l l p e r c e i v e t h e u s e f u l n e s s o f t h e i r e n v i r o n m e n t , t h e r e are so many v a r i a b l e s . F o r a p o p u l a t i o n o f d o m e s t i c s h e e p i n A u s t r a l i a , 80 p e r c e n t o f o n e y e a r ' s d i e t was composed o f p l a n t s c o m p r i s i n g o n l y 1 p e r c e n t o f t h e t o t a l f o r a g e a v a i l a b l e t o t h e a n i m a l s ( A r n o l d and D u d z i n s k i , 1 9 7 8 ) . A l t e r n a t i v e l y , t h e f a v o r e d f o o d might be c o i n c i d e n t a l l y t h e most abundant, or a n i m a l s may have learned t o f a v o r i t . A n i m a l s do n o t n e c e s s a r i l y u t i l i z e h a b i t a t o r v e g e t a t i o n u n i f o r m l y . I n t h e p r e s e n t s t u d y , h o r s e g r o u p s w e r e o b s e r v e d t o use n o t o n l y t h e same meadows r e p e a t e d l y , b u t a l s o t h e same p o r t i o n s o f t h e p a r t i c u l a r meadows. Some open a r e a s t h a t a p p e a r e d t o have good f o r a g e were not touched by c a t t l e a l l summer, w h i l e s i m i l a r a r e a s were r e p e a t e d l y g r a z e d , an o b s e r v a t i o n made a l s o by Heyes (1979) i n the same t y p e o f e n v i r o n m e n t t o t h e n o r t h e a s t o f my s t u d y a r e a . W i l l m s e_ _ L . , (1979) o b s e r v e d i n t h e w i n t e r d i e t s o f deer (O.hemionus) i n B.C., t h a t a l t h o u g h b i g s a g e b r u s h c o n t r i b u t e d 26 p e r c e n t - 66 - o f t h e d i e t , o n l y 7 p e r c e n t o f a v a i l a b l e p l a n t s were u s e d , w i t h n e i g h b o r i n g p l a n t s b e i n g unused. To c o m p l i c a t e m a t t e r s f u r t h e r , ' r e a l ' f o r a g e on o f f e r may a p p e a r t o be a d e q u a t e i n q u a n t i t y , b u t n o t a d e q u a t e i n q u a l i t y . W allmo §_t_ a l . , (1977) d e t e r m i n e d f o r a deer w i n t e r r a n g e i n C o l o r a d o t h a t f o r a g e q u a n t i t y was adequate f o r 14,000 d e e r b u t i t c o u l d n o t meet p r o t e i n and e n e r g y r e q u i r e m e n t s ; t h e e n e r g y b e i n g l i t t l e o v e r 50 p e r c e n t o f t h e e s t i m a t e d r e q u i r e m e n t . The l a t t e r n u t r i e n t b e i n g c r i t i c a l s i n c e t h e c a l o r i c need has been shown t o be t h e p r i m a r y need t o which t h e r e q u i r e m e n t s o f a l l n u t r i e n t s are l i n k e d ( B l a x t e r , 1962). On a s e a s o n a l b a s i s , e n e r g y s h o r t a g e i s a common problem t o h e r b i v o r e s (Owen ejt_ a l . , 1 9 7 8 ) . However, p r o t e i n and o t h e r n u t r i e n t s may a l s o be i n s h o r t s u p p l y , compounding t h e problem. M c L e a n a n d T i s d a l e (1960) f o u n d , i n t h e S o u t h e r n I n t e r i o r o f B.C., t h a t t h e p e r c e n t a g e s o f crude p r o t e i n and phosphorous d e c r e a s e d m a r k e d l y , and t h a t c r u d e f i b e r i n c r e a s e d , i n a l l c l a s s e s o f common f o r a g e from s p r i n g t o f a l l on b o t h g r a s s l a n d a n d f o r e s t r a n g e s . I t i s l i k e l y t h a t f o r a g e on t h e f o r e s t r a n g e s p r o b a b l y d o e s n o t meet c a t t l e growth r e q u i r e m e n t s from about mid-August onwards, but may meet maintenance r e q u i r e m e n t s t h r o u g h October (McLean and T i s d a l e , 1960). The v a r i a b i l i t i e s i n v o l v e d i n t h e c o n c e p t o f h a b i t a t a v a i l a b i l i t y s u p p o r t J o h n s o n ' s (1980) c o n t e n t i o n o f t h e a r b i t r a r i n e s s o f t h e a r r a y o f components t h e i n v e s t i g a t o r deems a v a i l a b l e t o t h e a n i m a l , and seems t o depend on when t h e i n v e s t i g a t o r e n t e r s t h e p i c t u r e . O p t i m a l f o r a g i n g t h e o r y i n a p a t c h y e n v i r o n m e n t i s a r a t h e r i d e a l i s t i c a p p r o a c h t o - 67 - r e s o u r c e u t i l i z a t i o n by a n i m a l s , d e v e l o p e d by E mlen (1966) and M a c A r t h u r and P i a n k a (1966) based on t h e i d e a t h a t a n i m a l s w i l l f o r a g e i n h a b i t a t s i n ' t h e most e c o n o m i c a l f a s h i o n , ' m a x i m i z i n g t h e i n t a k e o f energy and n u t r i e n t s , and m i n i m i z i n g t h e t i m e and energy r e q u i r e d to. do i t . Inherent t o the usefulness o f t h i s t h e o r y i s t h e a s s u m p t i o n t h a t " f i t n e s s a s s o c i a t e d w i t h an a n i m a l ' s f o r a g i n g b e h a v i o r has been maximized by n a t u r a l s e l e c t i o n " (Pyke e_t_ a i . , 1977). I n t h i s c o n t e x t , i t i s p o s s i b l e t h a t f e r a l h o r s e s , u n d e r g o i n g n a t u r a l s e l e c t i o n , and r a n g e c a t t l e , under a r t i f i c i a l s e l e c t i o n , a r e not d i r e c t l y comparable i n t h e i r h a b i t a t use p a t t e r n s . 4.2 D i s t r i b u t i o n p a t t e r n s and s p a t i a l s e p a r a t i o n An a n a l y s i s o f f e c a l c o u n t s p e r h a b i t a t t y p e a l o n g the t r a n s e c t s showed t h a t t h e r e was a s i g n i f i c a n t d i f f e r e n c e i n t h e c h o i c e o f h a b i t a t by h o r s e s , c a t t l e and moose i n d i c a t i n g t h a t each s p e c i e s h a s , t o some degree, d i f f e r e n t h a b i t a t s e l e c t i o n r e q u i r e m e n t s . E v i d e n c e s u p p o r t e d t h e c o n t e n t i o n p r e v i o u s l y e x p r e s s e d by s e v e r a l a u t h o r s t h a t h o r s e s a r e more s i m i l a r t o c a t t l e i n t h e i r h a b i t s t h a n t h e y a r e t o moose ( S t o r r a r §_£ a i . , 1 9 7 7 ; S a l t e r , 1 9 7 8 ) . I n a d d i t i o n , a l t h o u g h f e c a l c o n c e n t r a t i o n s i n d i c a t e d h o r s e s used f o r e s t more, and meadows l e s s t h a n c a t t l e , both spe c i e s appeared t o use meadows c o n s i d e r a b l y more t h a n any o t h e r h a b i t a t t y p e . The t r a n s e c t s ystem used i n t h e s t u d y a l l o w e d i n v e s t i g a t i o n i n t o t h e d i s t r i b u t i o n o f a n i m a l s by h a b i t a t t y p e and a l s o , i n r e g a r d t o s p a c e . The l a t t e r p r o v i d e d i n s i g h t i n t o t h e b a s i c c o n s i d e r a t i o n not answered by t h e h a b i t a t a n a l y s i s , i . e . a r e a n i m a l s w i t h an a p p a r e n t - 68 - o v e r l a p i n h a b i t a t c h o i c e , n e c e s s a r i l y u s i n g t h e same h a b i t a t s ( i . e . i n t h e same p l a c e ) . When t h e l e v e l s o f s p a t i a l a s s o c i a t i o n ( o r c o n v e r s e l y , t h e d e g r e e o f s e p a r a t i o n ) were e v a l u a t e d , h o r s e s and c a t t l e w e r e s e e n t o be a s s o c i a t e d more t h a n w o u l d be e x p e c t e d by c h a n c e . H o r s e s and moose were found t o be n e i t h e r i n t e r a c t i n g w i t h , nor a v o i d i n g e a c h o t h e r . C a t t l e and moose were s e e n t o be a v o i d i n g e i t h e r e a c h o t h e r , o r e a c h o t h e r ' s p r e f e r r e d h a b i t a t s . T h i s was not a s u r p r i s i n g r e s u l t , as h a b i t a t a n a l y s i s h a d e s s e n t i a l l y i n d i c a t e d t h e same r e s u l t ( i . e . moose u s e d h a b i t a t t y p e s l e a s t u s e d by h o r s e s and cows, and b o t h h o r s e s a n d cows u s e d meadows h e a v i l y ) . However, l o o k i n g f u r t h e r , i t was c l e a r l y s e e n on t h e f i v e t r a n s e c t g r a p h s t h a t e v e n though, as i n d i c a t e d , c a t t l e and horses were more o f t e n a s s o c i a t e d w i t h e a c h o t h e r t h a n w i t h moose, t h e c o n c e n t r a t i o n o f use i n any g i v e n a r e a v a r i e d w i d e l y between s p e c i e s . T h e r e a r e s e v e r a l i n t e r p r e t a t i o n s o f t h e s e r e s u l t s . F o r e x a m p l e , t h e a s s o c i a t i o n o f c a t t l e and h o r s e s shown t o o c c u r , c o u l d r e f l e c t t h e f a c t t h a t t h e r e a r e so many c a t t l e ( n e a r l y 90 p e r c e n t o f t h e t o t a l l a r g e h e r b i v o r e p o p u l a t i o n ) t h a t c a t t l e a r e l i t e r a l l y ' e v e r y w h e r e . ' T h e r e f o r e , i t would be a l m o s t i m p o s s i b l e t o a v o i d a c e r t a i n amount o f a s s o c i a t i o n , b u t an i m p o r t a n t u n a n s w e r e d q u e s t i o n i s w h e t h e r the a n i m a l s seek, a r e i n d i f f e r e n t t o , or a c t u a l l y t r y t o a v o i d a s s o c i a t i o n ? I f c a t t l e o u t n u m b e r e d h o r s e s i n any g i v e n a r e a , i t would i n d i c a t e p r o b a b l e i n d i f f e r e n c e ( i . e . t h a t h o r s e numbers are e s s e n t i a l l y ' l o s t ' i n c a t t l e numbers), or t h a t the two s p e c i e s m i g h t e v e n c h o o s e t o be t o g e t h e r . However, when h o r s e s are - 69 - f r e q u e n t l y f o u n d i n c o n s i d e r a b l y g r e a t e r f r e q u e n c y t h a n c a t t l e i n c e r t a i n p l a c e s , i t would t e n d t o i n d i c a t e e i t h e r an a c t i v e a t t e m p t t o a v o i d t h e o t h e r s p e c i e s , o r a d e f i n i t e d i f f e r e n c e i n t h e way h o r s e s and c a t t l e c h o o s e t o d i s t r i b u t e t h e m s e l v e s w i t h r e s p e c t t o sp a c e . A l l f i v e t r a n s e c t s show numerous instances w h e r e h o r s e s a r e h e a v i l y c o n c e n t r a t e d i n a r e a s t h a t c a t t l e use o n l y m o d e r a t e l y , s l i g h t l y , o r not a t a l l ; and c o n v e r s e l y , h o r s e s a r e f r e q u e n t l y n o t f o u n d i n a r e a s u s e d e x t e n s i v e l y by cows. C o n s i d e r i n g t h e v e r y s m a l l numbers o f h o r s e s , and t h e a p p a r e n t l y s t r o n g a f f i n i t y o f b o t h h o r s e s and cows f o r meadow h a b i t a t , which c o m p r i s e s o n l y 28.1 p e r c e n t o f t h e t o t a l s t u d y a r e a , i t seems s u r p r i s i n g t o f i n d such an unequal d i s t r i - b u t i o n o f t h e two s p e c i e s as i n d i c a t e d by t h e g r a p h s . F o r e x a m p l e , i t does n o t seem l i k e l y , i f movements o f h o r s e s and cows a r e e n t i r e l y random and i d e n t i c a l l y m o t i v a t e d i n terms o f meadow h a b i t a t ( a l r e a d y shown n o t t o be s o , f o r f o r e s t h a b i t a t ) , t h a t a t K i l o m e t e r 14 on T - l l l , w h ich was an i s o l a t e d meadow ( i . e . one s u r r o u n d e d by e x t e n s i v e f o r e s t a r e a s ) t h e p e r c e n t f r e q u e n c y o f use i n d i c a t e d f o r h o r s e s would be 22.5 p e r c e n t , a n d f o r cows, o n l y 2.5 p e r c e n t . As t h i s e x a m p l e i s not an i s o l a t e d i n s t a n c e , i t would appear t h a t the d i s t r i b u t i o n o f h o r s e s i s d i f f e r e n t f r o m t h a t o f c a t t l e . T h i s d i f f e r e n c e i n d i s t r i b u t i o n between t h e two s p e c i e s u n d e r l i n e s t h e concept t h a t t h e same h a b i t a t s i n d i f f e r e n t l o c a t i o n s are not n e c e s s a r i l y e q u a l i n t e r m s o f a n i m a l u s e . I t i s p o s s i b l e t h a t two areas t h a t seem s i m i l a r t o t h e human e y e , a r e u n i q u e i n t e r m s o f a n i m a l p e r c e p t i o n . T h e r e a r e numerous v a r i a b i l i t i e s t h a t c o u l d a c c o u n t f o r d i f f e r e n c e s i n use such as t r a d i t i o n a l use p a t t e r n s , d i s t a n c e f r o m w a t e r , d i s t a n c e f r o m s i m i l a r s i t e s , human h a r r a s s m e n t , and many o t h e r s depending on t h e s i t u a t i o n . I n t h e s p e c i f i c i n s t a n c e o f t h e meadow a t K i l o m e t e r 14 on T - I I I , t h e a p p a r e n t r e l u c t a n c e o f c a t t l e t o use f o r e s t e d a r e a s c o u l d k e e p them from g a i n i n g a c c e s s t o an o t h e r w i s e a c c e p t a b l e f o r a g e a r e a . Due t o a h i s t o r y o f human h a r r a s s m e n t i n t h e a r e a , h o r s e s may have ' l e a r n e d ' t o use t h e more i s o l a t e d and l e s s a c c e s s i b l e a r e a s o f t h e r e g i o n . C e r t a i n l y h a b i t and e x p e r i e n c e i n f l u e n c e an a n i m a l ' s p e r c e p t i o n o f a v a i l a b i l i t y , a n d t h e r e f o r e i t s d i s t r i b u t i o n . Weaver and Tomanek (1951) n o t e d t h a t c a t t l e d e v e l o p e d ' g r a z i n g r o u t e s ' on h i l l y r a n g e s , and t h i s had a marked e f f e c t on t h e use o f v e g e t a t i o n . Forested r a n g e a p p e a r s t o s h a r e a t t r i b u t e s w i t h ' h i l l y r a n g es' i n t h i s r e s p e c t , as my o b s e r v a t i o n s i n d i c a t e h a b i t u a l routes (e.g. t r a i l s and i n t e r - c o n n e c t i n g meadows) a f f e c t v e g e t a t i o n use by c a t t l e . The l a c k o f a s s o c i a t i o n shown t o o c c u r b e t w e e n h o r s e s o r c a t t l e and moose w o u l d seem t o be a t t r i b u t a b l e t o s e v e r a l f a c t o r s . F i r s t o f a l l , t h e h a b i t s o f moose a r e d i v e r g e n t f r o m t h o s e o f h o r s e s o r c a t t l e , and moose have b e e n shown t o have a much s t r o n g e r a t t a c h m e n t t o f o r e s t e d h a b i t a t t h a n e i t h e r h o r s e s o r c a t t l e , w h i c h w o u l d p r e c l u d e i n t e r a c t i o n t o a g r e a t e x t e n t . Secondly, as shown by the t r a n s e c t d i s t r i b u t i o n g r a p h s , moose t e n d n o t t o be q u i t e as w i d e s p r e a d i n t h e i r d i s t r i b u t i o n as h o r s e s and c a t t l e , and t h u s show c o n c e n t r a t i o n o f u se i n f e w e r a r e a s . T h i s c o u l d be due t o t h e much lower p o p u l a t i o n o f moose ( o n l y 2.9% o f t h e t o t a l number o f a l l a n i m a l s ) , and t h e i r r e l a t i v e l y s m a l l home r a n g e s (LeResche, 1974, c i t e d i n Coady, 1 9 8 2 ) . I t i s a l s o p o s s i b l e t h a t moose - 71 - may a c t i v e l y avoid both horses and c a t t l e . An i n t e r e s t i n seeing whether there were c o n s i s t e n c i e s i n the d i s t r i b u t i o n s of the three herbivores over a longer p e r i o d of time l e d to the comparison of the long-term and short-term accumulations of f e c a l material on T-V. A c e r t a i n amount of v a r i a t i o n i n animal d i s t r i b u t i o n and h a b i t a t use patterns could be expected from year to year due to such factors as c l i m a t i c v a r i a t i o n , and reduced or expanded animal numbers. Variation could also occur from changes i n ' t r a d i t i o n a l ' t r a v e l r o u t e s due t o death of dominant i n d i v i d u a l s or changes i n l e a d e r s h i p , f a c t o r s important to s o c i a l h e r bivores such as h o r s e s and c a t t l e . S t a t i s t i c a l l y , the two graphs of T-V, f o r long-term and short-term data, were shown to be d i f f e r e n t , but t h i s difference was only just s i g n i f i c a n t . Comparison of the two g r a p h s , however, y i e l d e d some interesting information which tended to corroborate the findings from the h a b i t a t and d i s t r i b u t i o n a n a l y s i s made on the f i v e • r e c e n t ' t r a n s e c t s . There were three notable s i m i l a r i t i e s between the two graphs: The rather pronounced use of t h i s t r a n s e c t by moose, the use by horses, but l i g h t use by c a t t l e of the ' i s o l a t e d * meadows d i s t r i b u t e d between Kilometers 11 and 14; and the 'absence' of c a t t l e between Kilometers 8 and 15, which c o n s i s t e d e s s e n t i a l l y of s o l i d f o r e s t with only small, scattered meadows. The long-term versus the short-term data for T-V indicated that o v e r a l l d i s t r i b u t i o n p a t t e r n s of the three species were d i f f e r e n t over time, while maintaining c e r t a i n consistencies i n v o l v i n g habitat preferences. It i s possible that variations - 72 - i n h a b i t a t u s e a r e c y c l i c o r a t l e a s t p a r t i a l l y p r e d i c t a b l e . The use o f h o r s e 'dung p i l e s * i n v a r i o u s p a r t s o f the s t u d y a r e a may s i g n a l c h a n g e s i n d i s t r i b u t i o n p a t t e r n s , as some go unmarked f o r a p e r i o d o f a y e a r or more (personal e s t i m a t i o n ) , w h i l e a t o t h e r s i t e s , a l o n g - n e g l e c t e d dung p i l e may s u d d e n l y b e g i n t o be marked and become ' a c t i v e * once more. T h i s c o u l d r e s u l t from t h e absence o f horses i n an area, or some u n i d e n t i f i e d r e a s o n f o r l a c k o f i n t e r e s t by s t a l l i o n s i n a p a r t i c u l a r dung p i l e . The s i g n i f i c a n c e o f dung p i l e s t o h o r s e s has not y e t been f u l l y e s t a b l i s h e d (Slade and G o d f r e y , 1982). 4.3 D i e t c h o i c e by h o r s e s and cows The f a c t t h a t c a t t l e and h o r s e s a r e b o t h g r a z e r s (Hafez, 1 9 6 4 ; S l a d e and G o d f r e y , 1982) , w o u l d i n d i c a t e a p o t e n t i a l s i m i l a r i t y i n t h e i r d i e t s . The l i m i t e d amount o f work done i n N o r t h A m e r i c a w i t h h o r s e s and o t h e r s p e c i e s (Hansen g£ a l . . 1977; O l s e n and H a n s e n , 1977; Hansen and C l a r k , 1977; S a l t e r , 1977) has shown i n g e n e r a l t h a t d i e t s o f f e r a l h o r s e s a r e : 1) most s i m i l a r t o r a n g e c a t t l e and e l k , and 2) l e a s t s i m i l a r t o moose, deer and pronghorn ( A n t i l o c a p r a a m e r i c a n a ) . D i e t a n a l y s i s i n d i c a t e d t h a t d u r i n g the months from June t h r o u g h September, h o r s e s and c a t t l e were exercising,, a f a i r l y h i g h d e g r e e o f s e l e c t i v i t y i n t h e i r c h o i c e o f f o r a g e s p e c i e s . F o r b o t h h e r b i v o r e s , t h e same 11 f o r a g e s c o m p r i s e d 80 p e r c e n t and more o f t h e t o t a l d i e t , b u t r e p r e s e n t e d o n l y 7.4 p e r c e n t o f t h e 148 p l a n t s p e c i e s d e t e r m i n e d t o be a v a i l a b l e t o them. T h e s e two f a c t o r s ( i . e . t h e h i g h d e g r e e o f s e l e c t i v i t y and 80-86% o f t h e d i e t s o f cows and h o r s e s , r e s p e c t i v e l y , b e i n g composed o f t h e same 11 f o r a g e s ) i n d i c a t e d a l a r g e o v e r l a p i n t h e c h o i c e o f d i e t . However, h a b i t a t use and d i s t r i b u t i o n a n a l y s i s i n d i c a t e d d i e t s e l e c t i o n s made by h o r s e s and cows w e r e n o t n e c e s s a r i l y i n t h e same l o c a t i o n . A t t e m p t s were made t o answer th e c r i t i c a l q u e s t i o n a r i s i n g from the s i g n i f i c a n t a m ount o f o v e r l a p , w h i c h was w h e t h e r a d i f f e r e n c e e x i s t e d i n t h e f o r a g i n g p a t t e r n s o f c a t t l e and h o r s e s ? In o t h e r words, when and how were p l a n t s p e c i e s u t i l i z e d ? D i d e i t h e r h e r b i v o r e have any p r e f e r e n c e f o r a s p e c i f i c b o t a n i c a l group (e.g. grasses)? Upon a n a l y s i s , d i f f e r e n c e s were f o u n d t o e x i s t i n t h e a p p a r e n t f o r a g i n g p a t t e r n s o f h o r s e s and c a t t l e . F o r example, i n t h e use o f t h e two m a j o r f o r a g e s , C a l a m a g r o s t i s rubescens and C a r e x a q u a t i l i s , w h i c h composed an average o f 37.3 p e r c e n t a n d 32.6 p e r c e n t , r e s p e c t i v e l y , o f t h e d i e t s o f h o r s e s and c a t t l e , n o t o n l y d i d h o r s e s a l w a y s e a t more C. rubescens (as much as 21% more i n A u g u s t ) , and cows c o n s i s t e n t l y more _. aqua- t i l i s , b u t o v e r t h e summer p e r i o d , t h e p a t t e r n o f use f o r rubescens was s i g n i f i c a n t l y d i f f e r e n t f o r t h e two h e r b i v o r e s . H o r s e u s e o f t h i s s p e c i e s p e a k e d i n A u g u s t , w h i l e c a t t l e use o f i t i n c r e a s e d from June t h r o u g h September. The a v e r a g e consumption o f s i x o f t h e 11 dominant f o r a g e s v a r i e d s i g n i f i c a n t l y i n t h e i r use by b o t h h o r s e s and c a t t l e on a monthly b a s i s from June t h r o u g h September. T h i s v a r i a t i o n c o u l d be t h e r e s u l t o f a number o f f a c t o r s , s u c h as one o r more g r o w t h s p u r t s o c c u r r i n g i n a p l a n t s p e c i e s m a k i n g i t more a t t r a c t i v e t o h e r b i v o r e s a t c e r t a i n s e a s o n s . S t i p a o c c i - d e n t a l i s i n d i c a t e s t h i s t y p e o f a v a i l a b i l i t y p a t t e r n i n t h a t , f o r b o t h c a t t l e and h o r s e s , use i s low i n June but i n c r e a s e s - 74 - i n J u l y and remains f a i r l y steady through September. O p p o r t u n i s t i c f o r a g i n g c o u l d e x p l a i n some v a r i a b i l i t y i f a n i m a l s do n o t p a r t i c u l a r l y s e e k o u t a p l a n t s p e c i e s b u t e a t i t when t h e y happen t o f i n d i t . As i n d i c a t e d f o r C. r u b e s c e n s . t h e r e were s i g n i f i c a n t d i f f e r e n c e s i n t h e way h o r s e s and c a t t l e u s e d s i x o f t h e 11 dominant f o r a g e s . G l y c e r i a b o r e a l i s . f o r example, a l w a y s was u s e d by c a t t l e more t h a n by h o r s e s . In a d d i t i o n , w h e r e a s h o r s e s h e l d s t e a d i l y t o t h e 2 t o 3 p e r c e n t l e v e l o f d i e t c o n t e n t f o r t h i s g r a s s s p e c i e s , c a t t l e i n c r e a s e d t h e i r use f r o m 4.5 p e r c e n t i n June t o a peak i n J u l y (8.3 p e r c e n t ) , and t h e n came back t o a p p r o x i m a t e l y 4 p e r c e n t f o r August and S e p t e m b e r . E i g h t o f t h e f o r a g e s (not, i n c l u d i n g r u b e s c e n s . C. a q u a t i l i s a n d G l y c e r i a b o r e a l i s ) showed f l e x i b i l i t y i n t h e i r u se b e t w e e n t h e two h e r b i v o r e s o v e r t h e f o u r months. H o r s e s u s e d an a v e r a g e o f 4.3 p e r c e n t more Poa j u n c i f o l i a t h a n c a t t l e i n t h r e e out o f f o u r months, but i n A u g u s t , c a t t l e u s e d n e a r l y 3.0 p e r c e n t more t h a n h o r s e s . Poa j u n c i f o l i a was one o f t h e t h r e e g r a s s e s , i n w h i c h t h e d i f f e r e n c e i n use between h o r s e s and c a t t l e was found t o be s i g n i f i c a n t l y dependent on a p a r t i c u l a r month. I t i s d i f f i c u l t t o a c c o u n t f o r t h e v a r i a t i o n s t h a t o c c u r i n t h e use o f some p l a n t s p e c i e s . For e x a m p l e , C a r e x r o s t a t a . d e c r e a s e s s t e a d i l y from June t h r o u g h S e p t e m b e r i n c a t t l e d i e t s , w h i l e i t i n c r e a s e s s t e a d i l y i n t h e d i e t s o f h o r s e s c o i n c i d i n g w i t h a d e c l i n e i n o v e r a l l sedge u s e by h o r s e s f r o m J u l y t h r o u g h S e p t e m b e r . I t h i n k t h a t , o u t s i d e o f t h e t o p t h r e e o r f o u r f o r a g e c h o i c e s f o r a p a r t i c u l a r g e o g r a p h i c a l l o c a t i o n , i t i s p r o b a b l e t h a t t h e o r d e r of s p e c i f i c p l a n t s e l e c t i o n by h o r s e s and c a t t l e w o u l d be s e e n t o v a r y - 75 - o v e r a number o f y e a r s i f c o n s e c u t i v e d i e t a n a l y s e s were done. I b a s e t h i s c o n t e n t i o n on t h e e v i d e n c e o f the somewhat e r r a t i c and l o w e r r e l a t i v e use o f t h e o t h e r p l a n t s p e c i e s i n t h e d i e t , a n d a l s o b e c a u s e most o f t h e d i e t s t u d i e s done t o d a t e on f e r a l h o r s e s and r a n g e c a t t l e have shown a s i m i l a r p a t t e r n o f use ( i . e . h e a v y d e p e n d e n c e on a v e r y few p l a n t s p e c i e s , w i t h t h e p e r c e n t a g e use o f o t h e r s p e c i e s r a p i d l y d e c r e a s i n g ) ( S a l t e r , 1978; Hansen and C l a r k , 1977; Hansen and R e i d , 1975; H a n s e n , 1976; Hansen §_£ aj.. , 1 9 7 7 ) . I n t h e p r e s e n t s t u d y , t h e r e w e r e l a r g e d i f f e r e n c e s among t h e a v e r a g e summer use o f t h e f i r s t c h o i c e o f h o r s e s , C_. rubescens ( 2 6 . 6 % ) , t h e f i f t h c h o i c e , Juncus b a l t i c u s (6.4%) and the eleventh c h o i c e , <_. n e g l e c t a ( 3 . 0 % ) ; s i m i l a r l y f o r c a t t l e , the f i r s t choice being, C. rubescens ( 1 6 . 5 % ) , t h e f i f t h c h o i c e , C. n e g l e c t a ( 5 . 4 % ) , and t h e e l e v e n t h c h o i c e , C. c o n c i n o i d e s ( 4 . 4 % ) . The a n a l y s i s o f t h e p r o p o r t i o n a t e use o f b o t a n i c a l groups by h o r s e s a n d cows showed a s i g n i f i c a n t d i f f e r e n c e i n t h e m o n t h l y u t i l i z a t i o n o f g r a s s e s , r u s h - s e d g e s and ' o t h e r . ' I n summer d i e t s h o r s e s used an average of 61 p e r c e n t g r a s s e s , w h e r e a s c a t t l e u s e d an a v e r a g e o f 51 p e r c e n t g r a s s e s w i t h a h i g h e r p r o p o r t i o n o f sedges, f o r b s and browse. Horses c l e a r l y a r e more o r i e n t e d t o w a r d s u t i l i z a t i o n o f g r a s s s p e c i e s t h a n a r e c a t t l e i n t h e s p r i n g , summer and f a l l , p a r t i c u l a r l y i n l i g h t o f t h e r a t h e r d r a m a t i c s w i t c h t o sedges i n the w i n t e r . T h i s i s n o t s u r p r i s i n g c o n s i d e r i n g t h e h i g h e r t o l e r a n c e t o f i b e r by h o r s e s ( J a n i s , 1 9 7 6 ) , and t h e d e n t a l a d a p t a t i o n s f o r a h i g h l y f i b r o u s d i e t . E q u i d d e n t a l development exceeds t h a t o f r u m i n a n t s i n t h e c o m b i n a t i o n o f p r e m o l a r c o m p l e x i t y , - 76 - movement f o r w a r d o f t h e main c e n t e r o f m a s t i c a t i o n and s t r u c t u r a l a d a p t a t i o n s t o w i t h s t a n d wear a b r a s i o n , i n c l u d i n g the ' i n f u n - d i b u l u m , * a d e e p i n v a g i n a t i o n f i l l e d w i t h cement on u p p e r a n d l o w e r i n c i s o r s f o u n d o n l y i n modern e q u i d s ( L y d e k k e r , 1 9 1 2 ; W e l l e r , 1 9 6 8 ; S i s s o n , 1 9 5 3 ) . A l t h o u g h , i n g e n e r a l , f e c a l a n a l y s i s showed t h a t b o t h c a t t l e and h o r s e s were b e i n g s e l e c t i v e as t o p l a n t s p e c i e s , t h e p r o p o r t i o n a t e use of b o t a n i c a l g r o u p s i n d i c a t e s c a t t l e were b e i n g somewhat more s e l e c t i v e f o r l o w e r f i b e r c o n t e n t m a t e r i a l . I n w i n t e r , h o r s e s d e c r e a s e d t h e number o f g r a s s , sedge and f o r b s p e c i e s e a t e n . A t t h e same t i m e , t h e y i n c r e a s e d t h e i r u t i l i z a t i o n o f s h r u b m a t e r i a l , which i s more a v a i l a b l e and h o l d s i t s n u t r i e n t c o n t e n t b e t t e r t h a n g r a s s e s or f o r b s i n t h e f a l l and w i n t e r (Nagy, 1969; Cook, 1 9 5 6 ) . T h i s s o r t o f f e e d i n g b e h a v i o r would t e n d t o o f f s e t the e f f e c t s of inadequate p r o t e i n and energy i n t a k e shown t o occur i n horses kept year-round s o l e l y on g r a s s p a s t u r e (Owen §i a!., 1978). I t i s p o s s i b l e , b u t has n o t y e t b e e n shown t o o c c u r , t h a t f e r a l h o r s e s reduce t h e i r v o l u n t a r y f o o d i n t a k e d u r i n g s t r e s s f u l periods ( i . e . winter) as do some o t h e r h e r b i v o r e s s u c h as deer and moose ( R o b b i n s , 1983). The i n c r e a s e d w i n t e r - u s e o f r u s h - s e d g e s f r o m 33.2% i n summer t o 48% i n w i n t e r by h o r s e s , a l s o may have been p a r t i a l l y a t t r i b u t a b l e t o a v a i l a b i l i t y . R u s h - s e d g e s t e n d t o r e m a i n e x p o s e d , u n l i k e f o r a g e s i n t h e meadow and f o r e s t e d a r e a s . Snow d o e s n o t a p p e a r t o c o l l e c t i n t h e s e d g e a r e a s d e s p i t e t h e i r open a s p e c t , or perhaps because o f i t ( i . e . wind e f f e c t s ) , c o m b i n e d w i t h t h e i r i n h e r e n t s u r f a c e u n e v e n e s s . The sedge - 77 - a r e a s were o b s e r v e d t o have s h a l l o w , p o o l e d water around the p l a n t s e v e n on w i n t e r d a y s when t h e t e m p e r a t u r e was f a r below f r e e z i n g ( p e r s . o b s . , M a r c h , 1 9 8 0 ) . O f t e n t h e sedge a r e a s o f f e r ' p r i s t i n e ' g r a z i n g , as some c a n o n l y be u t i l i z e d i n t h e w i n t e r , b e i n g i n a c c e s s a b l e t o a n y t h i n g b u t moose i n t h e summer due t o water d e p t h and mud. I t i s p o s s i b l e t h a t sedges r e t a i n a h i g h e r n u t r i e n t c o n t e n t d u r i n g the w i n t e r t h a n g r a s s e s o r f o r b s . The m o b i l e l i p s o f h o r s e s w o u l d p h y s i c a l l y a l l o w f o r t h e e x p l o i t a t i o n o f t h e g r e e n m a t e r i a l i n t h e c e n t e r o f s e d g e p l a n t s . T h i s m a t e r i a l , shown t o o c c u r f o r b o t h Carex r o s t a t a ( B e r n a r d , 1974) and Carex a q u a t i l i s (Gorham and Sommers, 1972), w h i c h t o g e t h e r composed 28.5 p e r c e n t o f t h e t o t a l w i n t e r d i e t o f h o r s e s , had a c r u d e p r o t e i n l e v e l o f 6.5 p e r c e n t . W e a t h e r e d g r a s s e s i n w i n t e r w o u l d be e x p e c t e d t o have lower CP v a l u e s . C o m p a r a t i v e v a l u e s o f CP l e v e l s f o r some g r a s s e s i n a ' w e a t h e r e d ' c o n d i t i o n , c o l l e c t e d d u r i n g t h e months o f O c t o b e r and November, r e p r e s e n t a t i v e o f s p e c i e s found i n the s t u d y a r e a , were as f o l l o w s : A g r o p y r o n s p p . 3.08%, Bromus s p p . 2.8%, K o e l e r i a s p . 1.89%, Poa s p p . 3 . 3 % a v e . , and S t i p a s p p . 3 . 3 % a v e > (McLean and T i s d a l e , 1 9 6 0). I n a second s t u d y ( D e m a r c h i , 1 9 7 3 ) , b o t h A g r o p y r o n s p i c a t u m and S t i p a comata, c o l l e c t e d i n l a t e M a r c h , were found t o have average CP v a l u e s o f 2.7 and 4.5 p e r c e n t , r e s p e c t i v e l y . I t w o u l d appear t h a t even w i t h t h e t r a n s l o c a t i o n o f m a t e r i a l s t o underground s t o r a g e a r e a s , s u c h as r h i z o m e s by sedge p l a n t s i n the f a l l ( B e r n a r d , 1974) , and t h e s u b s e q u e n t d e c l i n e i n n u t r i e n t c o n t e n t i n the v i s i b l e p o r t i o n s o f t h e p l a n t s (McLean and T i s d a l e , 1 9 6 0 ) , i t would be n u t r i t i o n a l l y advantageous t o in c r e a s e the u t i l i z a t i o n - 78 - o f s e d g e p l a n t s o v e r g r a s s e s i n t h e w i n t e r . P a r t i c u l a r l y s o b e c a u s e p i n e g r a s s , by f a r t h e most i m p o r t a n t summer horse f o r a g e , a l t h o u g h n u t r i t i o n a l l y v a r i a b l e on a g e o g r a p h i c a l b a s i s (McLean and F r e y m a n , 1969), appears t o be a poor w i n t e r f o r a g e . In a d d i t i o n t o low crude p r o t e i n (2.7%) and phosphorous l e v e l s , m a t u r e p i n e g r a s s has an u n u s u a l l y h i g h s i l i c a c o n t e n t w h i c h e x c e e d s 10 p e r c e n t by October (McLean and Freyman, 1969; M c L e a n and T i s d a l e , 1 9 6 0 ) . A l t h o u g h t h e d i e t r e s u l t s show a d e c i s i v e i n c r e a s e i n t h e use o f b r o w s e and r u s h - s e d g e s by h o r s e s i n t h e w i n t e r , i t i s n o t p o s s i b l e t o s a y w h e t h e r i t i s r e l a t e d t o the a v a i l a b i l i t y o f t h e s e p l a n t s , t o s e l e c t i v i t y f o r h i g h e r n u t r i e n t c o n t e n t , or t o b o t h . W h i l e h o r s e s and c a t t l e were b o t h on the s t u d y s i t e from June t h r o u g h September, th e d i f f e r e n c e s i n h a b i t a t - u s e , d i s t r i - b u t i o n and d i e t c h o i c e a p p e a r e d t o a m e l i o r a t e t h e p o t e n t i a l f o r i n t e r f e r e n c e b e t w e e n t h e two s p e c i e s . However, because h o r s e s depend h e a v i l y on rush-sedges i n t h e w i n t e r , and c a t t l e u t i l i z e d them i n the summer, t h e r e i s a p o s s i b i l i t y t h a t c a t t l e summer-use c o u l d a d v e r s e l y e f f e c t h o r s e s . C a r e x a q u a t i l i s a n d C. r o s t a t a were t h e two s e d g e s u s e d most by h o r s e s and c a t t l e f r o m J u n e t h r o u g h S e p t e m b e r , and t o g e t h e r t h e y formed a s i g n i f i c a n t p a r t ( 2 8 . 5 % ) o f t h e w i n t e r d i e t o f h o r s e s . H o w e v e r , t h e s i n g l e most i m p o r t a n t w i n t e r f o r a g e p l a n t f o r h o r s e s was a c t u a l l y J u n c u s b a l t i c u s ( 1 7 . 2 % ) , which s u s t a i n e d o n l y m o d e r a t e use by h o r s e s and c a t t l e i n the summer. T h i s , c o m b i n e d w i t h t h e a p p a r e n t l y e x c l u s i v e w i n t e r - a c c e s s i b i l i t y o f some wet meadow a r e a s ( e v e n i n v e r y d r y summers) w o u l d , a g a i n , t e n d t o a m e l i o r a t e t h e e f f e c t o f p o s s i b l e i n t e r f e r e n c e . - 79 - By t h e same t o k e n , h o r s e s may u n d e r m i n e t h e v a l u e o f s p r i n g r a n g e t o c a t t l e . However, b a s e d on u s e - d i f f e r e n c e s between t h e two h e r b i v o r e s a l r e a d y shown t o o c c u r , and t h e r e l a t i v e l y f e w h o r s e s , t h e r e w o u l d seem l i t t l e p r o b a b i l i t y o f t h i s . S a l t e r (1978) e s t i m a t e d t h a t o n l y 5 p e r c e n t o f n o n - f o r e s t e d h a b i t a t on h i s s t u d y s i t e i n A l b e r t a showed e v i d e n c e o f s p r i n g ( J u n e ) g r a z i n g by h o r s e s . F o r b s w o u l d be e x p e c t e d t o be the most v u l n e r a b l e t o use by h o r s e s p r i o r t o t h e a r r i v a l o f range c a t t l e , b u t d i e t a n a l y s i s i n d i c a t e d o n l y a 1 p e r c e n t f o r b use by h o r s e s i n June. The e s t i m a t e d d i e t s a l s o showed an i n c r e a s e i n t h e use o f c o n i f e r n e e d l e s d u r i n g t h e w i n t e r ( 6 . 2 5 % ) . S a l t e r (1978) o b s e r v e d a s i m i l a r i n c r e a s e i n t h e p r e s e n c e o f c o n i f e r n e e d l e s i n t h e w i n t e r d i e t s o f h o r s e s , b u t c o n s i d e r e d t h e phenomenon t o be a c c i d e n t a l . I do n o t f e e l t h e i n c r e a s e d use o f c o n i f e r n e e d l e s i s a c c i d e n t a l c o n s i d e r i n g t h e a b i l i t y o f h o r s e s t o e x p e l unwanted m a t e r i a l t a k e n i n t o t h e mouth, even when mixed w i t h d e s i r e a b l e i t e m s ( p e r s . o b s . ) . A l s o , some w i l d ruminants a p p a r e n t l y i n c r e a s e t h e i r use o f c o n i f e r n e e d l e s d u r i n g w i n t e r , w hich may be l i k e m i s t l e t o e , r e t a i n i n g a f a i r l y high carbohydrate c o n t e n t i n w i n t e r ( p e r s . comm. w. Bruce D a v i t t , W.S.U. W i l d l i f e H a b i t a t Lab, 1981). There i s e v i d e n c e t h a t rumen microorganisms become adapted t o t h e i n h i b i t o r y e f f e c t on d i g e s t i o n o f c e r t a i n e s s e n t i a l o i l s f o u n d i n c o n i f e r n e e d l e s ( W i l s o n , 1 9 6 9 ) . Monoterpene a l c o h o l s appear t o be the worst offenders i n d i g e s t i v e i n h i b i t i o n ( A r n o l d , 1981), but no r e f e r e n c e was found r e g a r d i n g t h e i r e f f e c t , i f any, on m o n o g a s t r i c d i g e s t i o n . I t i s e x p e c t e d t h a t as a r e s u l t o f more i n t e n s e m a s t i c a t i o n - 80 - by h o r s e s w i t h a d e n t i t i o n m a x i m a l l y d e v e l o p e d f o r u t i l i z a t i o n o f a b r a s i v e r o u g h a g e s , f e w e r f o r b s w o u l d be i d e n t i f i e d by f e c a l e p i d e r m a l a n a l y s i s i n h o r s e s t h a n i n c a t t l e . The major d r a w b a c k o f t h e a n a l y s i s t e c h n i q u e l i e s i n t h e d i f f e r e n t i a l d i g e s t i o n o f b r o w s e , f o r b s , s e d g e s and g r a s s e s . T h e r e i s g e n e r a l a g r e e m e n t t h a t i t i s a good t e c h n i q u e f o r a n i m a l s whose d i e t i s made up l a r g e l y o f g r a s s e s , s u c h as t h e z e b r a (Equus b u r c h e l l i i ) (Owaga, 1 9 7 7 ) , h o r s e s ( S a l t e r and Hudson, 1 9 7 9 ) b u r r o s (Woodward, 1 9 7 6 ) , c a t t l e and e l k (Hansen and R e i d , 1 9 7 5 ) . G r a s s f r a g m e n t s a r e g e n e r a l l y q u i t e r e s i s t a n t t o d i g e s t i o n and r e t a i n enough c h a r a c t e r s f o r i d e n t i f i c a t i o n , u n l i k e f o r b s w h i c h a r e much more e a s i l y and q u i c k l y d i g e s t e d . Work i s c u r r e n t l y b e i n g done a t W a s h i n g t o n S t a t e U n i v e r s i t y t o d e t e r m i n e a d i f f e r e n t i a l d i g e s t i o n c o e f f i c i e n t f o r t h e v a r i o u s a n i m a l s p e c i e s t o w h i c h t h e t e c h n i q u e i s a p p l i e d . I n h o r s e v e r s u s c a t t l e d i e t c o m p a r i s o n s t o d a t e , h o r s e s have n o t b e e n shown t o use f o r b s s i g n i f i c a n t l y . S a l t e r (1978) s t a t e s t h a t a t o t a l l e v e l o f l e s s t h a n 3 p e r c e n t f o r b s were u t i l i z e d e a c h month... " s p e c i e s i n t h e P o t e n t i l l a - G e u m groups were t h e o n l y ones w h i c h a p p e a r e d c o n s i s t e n t l y . " As p o i n t e d o u t by D a v i t t ( p e r s . comm., 1981) t h e f o r b s t h a t do show up, t e n d t o be b i a s e d towards t h o s e t h a t have h a i r s ( l i k e P o t e n t i l l a and Geum). I t i s s u g g e s t e d t h a t p o s s i b l y t h e h a i r s t e n d t o r e m a i n u n d i g e s t e d and t h e r e f o r e c a n be i d e n t i f i e d . D a v i t t f u r t h e r s u g g e s t s t h e r e may be a c o r r e c t i o n f a c t o r f o r f o r b s o f 4 t o 5 t i m e s more t h a n a r e p r e s e n t l y b e i n g i d e n t i f i e d by e p i d e r m a l a n a l y s i s . F e c a l a n a l y s i s i n t h i s s t u d y d i d not n e c e s s a r i l y i n d i c a t e - 81 - a w e akness i n t h e t e c h n i q u e f o r i d e n t i f y i n g f o r b s , as t h e r e was r e l a t i v e l y l i t t l e d i f f e r e n c e b e t w e e n c a t t l e and h o r s e s i n t h e i r use o f number o f s p e c i e s o f f o r b e s , g r a s s e s or sedges. The r e s u l t s d i d i n d i c a t e c o n s i s t e n t l y h i g h e r s e l e c t i o n f o r f o r b s i n t h e d i e t by c a t t l e f o r June, J u l y , August and September. S e e g m i l l e r and Ohmart (1981) showed by f e c a l d i e t a n a l y s i s , t h a t f o r w i l d b u r r o s i n A r i z o n a f o r b s were th e p r e f e r r e d foods when t h e y w e r e g r e e n and s u c c u l e n t f r o m F e b r u a r y t o J u n e , c o m p o s i n g as much as 56.5 p e r c e n t o f t h e t o t a l d i e t . There seems no r e a s o n t o assume t h a t f o r b s i n h o r s e d i e t s s h o u l d be more d i f f i c u l t t o i d e n t i f y than f o r b s i n burro d i e t s , p r o v i d i n g t h e f o r b s a r e p r e s e n t . C o n v e r s e l y , t h e r e d o e s n o t seem t o be j u s t i f i c a t i o n f o r a t t r i b u t i n g ' l a c k * o f f o r b s t o e r r o r i n t e c h n i q u e r a t h e r t h a n s e l e c t i v i t y on t h e p a r t o f h o r s e s and cows. The t o t a l d i e t s o f h o r s e s and c a t t l e r e p r e s e n t e d o n l y a f r a c t i o n o f t h e p l a n t s p e c i e s c o m p r i s i n g t h e b o t a n i c a l groups. F o r e x a m p l e , b o t h h o r s e s and c a t t l e u s e d o n l y s i x o f t h e 13 r u s h - s e d g e s f o u n d i n t h e s t u d y a r e a , i g n o r i n g some r e l a t i v e l y w i d e s p r e a d s p e c i e s f o u n d c o n s i s t e n t l y i n v a r i o u s l o c a t i o n s s u c h as C a r e x p a c h y s t a c h y a and C_. p a r r y a n a . Out o f 88 f o r b s p e c i e s , 16 w e r e f o u n d t o be u t i l i z e d by h o r s e s and 20 by c a t t l e . Of t h e 43 g r a s s s p e c i e s i d e n t i f i e d , h o r s e s u t i l i z e d o n l y 2 2 , a n d cows 19. O v e r a l l , t h e most i m p o r t a n t f o r a g e f o r b o t h h e r b i v o r e s was p i n e g r a s s ( C a l a m a g r o s t i s rubescens) . a l t h o u g h C a r e x a q u a t i l i s was u t i l i z e d by c a t t l e o n l y s l i g h t l y l e s s . F o r h o r s e s , h o w e v e r , C. r u b e s c e n s was by f a r the most i m p o r t a n t f o r a g e p l a n t ; more t h a n 16 p e r c e n t u t i l i z e d t h a n - 82 - t h e n e x t f a v o r i t e (_. a q u a t i l i s . S a l t e r (1978) d e t e r m i n e d t h a t t h e g r a s s Elymus i n n o v a t u s , found p r e d o m i n a n t l y i n f o r e s t e d h a b i t a t s on h i s s t u d y a r e a i n s o u t h w e s t e r n A l b e r t a , composed t h e most d o m i n a n t s i n g l e f o r a g e o f f e r a l h o r s e s . The major d i e t component o f c a t t l e i n S a l t e r ' s study ( i g n o r i n g the 'clumped' v a l u e o f sedge s p e c i e s ) was F e s t u c a spp., which was t h e second most i m p o r t a n t component i n f e r a l h o rse d i e t s . C a l a m a q r o s t i s r u b e s c e n s i s an i m p o r t a n t f o r a g e s p e c i e s on B.C. r a n g e s . By f a r t h e b u l k o f r a n g e l a n d s i n B.C. i s on f o r e s t e d l a n d , and a p p r o x i m a t e l y 3.2 m i l l i o n ha (6 m i l l i o n a c r e s ) o f t h i s l i e s i n t h e I n t e r i o r D o u g l a s F i r Zone (IDF) ( T i s d a l e and McLean, 1957; S t o u t and Brook, 1980). The IDF(b) i s t h e l a r g e s t s u b z o n e and i s l o c a t e d a l m o s t e x c l u s i v e l y on t h e F r a s e r P l a t e a u (Annas and Coupe, 1 9 7 9 ) . P i n e g r a s s may o c c u p y up t o 80 p e r c e n t o f the ground a r e a i n t h i s zone ( S t o u t e t a l . . 1 9 8 0 ) , b u t v a r i e s b e t w e e n f o r e s t e d h a b i t a t t y p e s . R e c e n t s t u d i e s show t h a t p i n e g r a s s may have two growth s p u r t s , one i n t h e s p r i n g a n d , d e p e n d i n g p a r t l y on m o i s t u r e , one i n t h e f a l l . A r o u n d e a r l y J u l y , g r o w t h i s s l o w i n g down p r i o r t o summer d o r m a n c y , and t h a t i s t h e time i t i s most s e n s i t i v e t o g r a z i n g ( S t o u t e_t a l . , 1980; S t o u t and Brook, 1980). McLean ( 1 9 6 7 ) s t u d y i n g r a n g e where p i n e g r a s s p r o v i d e d 50 p e r c e n t o f a v a i l a b l e f o r a g e , determined t h a t i t was r e a d i l y a c c e p t a b l e t o c a t t l e i n e a r l y summer, became u n p a l a t a b l e by mid-August, and was a g a i n used i n September (McLean, 1980). F e c a l a n a l y s i s i n my s t u d y i n d i c a t e s a s l i g h t l y d i f f e r e n t p a t t e r n , i n t h a t c a t t l e showed a p r e f e r e n c e f o r C a r e x a q u a t i l i s i n J u n e and J u l y ( a l t h o u g h p i n e g r a s s was t h e i r second c h o i c e ) , and e s s e n t i a l l y - 83 - d o u b l e d t h e i r u t i l i z a t i o n o f p i n e g r a s s f o r August and September. My r e s u l t s a r e a l s o d i r e c t l y c o n t r a d i c t o r y t o the o b s e r v a t i o n s o f Heyes (1979) r e g a r d i n g t h e p r e f e r e n c e by c a t t l e o f w e t l a n d meadows f r o m l a t e summer o n , when t h e n u t r i t i o n a l q u a l i t y and p a l a t a b i l i t y o f t h e upl a n d f o r a g e d e c l i n e s . The h e a v i e s t use by c a t t l e o f p i n e g r a s s , t h e m a j o r u p l a n d f o r a g e , was i n S e p t e m b e r , and t h e o v e r a l l use o f s e d g e s by c a t t l e was l e s s i n A u g u s t and Septe m b e r t h a n i n June and J u l y . I n a d d i t i o n , my own o b s e r v a t i o n s i n 1978, a d r y summer when more sedge meadows t h a n u s u a l were a c c e s s i b l e t o c a t t l e , showed t h a t by t h e end o f A u g u s t , b o t h d r y and wet meadows used by c a t t l e were s e v e r e l y g r a z e d and had s u s t a i n e d heavy t r a m p l i n g damage, t o w h i c h s e d g e meadows a r e e s p e c i a l l y v u l n e r a b l e ( M i l l a r , 1973; H e y e s , 1 9 7 9 ) . From June t h r o u g h S e p t e m b e r , p i n e g r a s s formed t h e h i g h e s t p e r c e n t o f any p l a n t s p e c i e s i n ho r s e d i e t s , t h e o n l y s p e c i e s c o m i n g e v e n c l o s e were Poa j u n c i f o l i a i n June and Carex a q u a t i l i s i n J u l y . Horses c o n s i s t e n t l y u t i l i z e d p i n e g r a s s r o u g h l y 1.5 t o 2 t i m e s more t h a n c a t t l e . - 84 - Chapter 5: Summary C o m p a r a t i v e s e l e c t i v i t y o f foo d m a t e r i a l s by m o n o g a s t r i c h e r b i v o r e s ( l a r g e l y e q u i d s ) and r u m i n a n t s has been examined t o some e x t e n t . R e c e n t l y p u b l i s h e d d a t a has l e d t o a c o n f u s i n g and c o n t r a d i c t o r y o v e r l a p o f i d e a s . T r a d i t i o n a l l y , the p r e v a i l i n g i d e a h a s been t h a t r u m i n a n t s were i d e a l l y s u i t e d t o d e a l w i t h f o r a g e s o f p o o r n u t r i t i v e q u a l i t y (Swenson, 1970). However, s u c h f a c t o r s as t h e need f o r s u f f i c i e n t b a c t e r i a l energy and t h e r e l a t i v e l y s l o w r a t e o f p a s s a g e , h i n d e r t h e u t i l i z a t i o n o f p o o r q u a l i t y f o r a g e by r u m i n a n t s s u c h as c a t t l e ( J a n i s , 1 9 7 6 ; 0 r s k o v , 1975; Hogan and W e s t o n , 1970; A n n i s o n , 1970; C o r b e t t , 1 9 6 9 ) . O t h e r r u m i n a n t s , s u c h a s d e e r , r e q u i r e a h i g h d i e t q u a l i t y i n summer t o p r o d u c e t h e f a t r e s e r v e t h e y depend upon i n t h e w i n t e r ( S h o r t , 1 9 8 1 ) . Deer cannot d i g e s t l a r g e amounts o f c e l l u l o s e (Nagy, 1969) and have a v e r y l i m i t e d c a p a c i t y f o r i n c r e a s i n g f o o d i n t a k e t o accommodate f o r decreased f o o d q u a l i t y (Amman e_t_ a l . , 1973 c i t e d i n Wallmo fit al., 1977). S u b s e q u e n t l y t h e y t e n d t o be h i g h l y s e l e c t i v e i n t h e i r c h o i c e o f f o o d i n b o t h summer and w i n t e r (Mackie _ t ai., 1982; S h o r t , 1 9 8 1 ) . U n t i l q u i t e r e c e n t l y , i t was assumed t h a t t h e ho r s e r e q u i r e d h i g h q u a l i t y f o r a g e , b u t r e c e n t e v i d e n c e i n d i c a t e s t h e a b i l i t y o f e q u i d s i n g e n e r a l t o e x p l o i t e nvironments which a r e e i t h e r o p t i m a l o r s u b - o p t i m a l f o r r u m i n a n t s . T h i s i s due t o t h e i r much g r e a t e r t o l e r a n c e o f h i g h f i b e r c o n t e n t i n t he d i e t t h a n ruminants ( J a n i s , 1976). I n b o t h North America ( O l s e n and H a n s e n , 1977) and A f r i c a ( B e l l , 1 9 71; 1970) t h e r e s u l t s o f f i e l d s t u d i e s have shown ruminant f e e d i n g s t r a t e g y - 85 - t o be more s e l e c t i v e f o r d i g e s t i b l e t i s s u e t h a n t h a t o f e q u i d s . D a t a c o l l e c t e d by B e l l (1970) i n a s t u d y o f h e r b a g e use by z e b r a (Equus burchej3,i bohmi) compared t o f o u r ruminant s p e c i e s showed t h a t z e b r a : 1) S e l e c t e d a d i e t w i t h t h e l o w e s t p r o t e i n c o n t e n t , 2) u t i l i z e d t h e h i g h e s t amount o f r o u g h a g e , and 3) chose t h e commonest and most a c c e s s i b l e herbage. A l t h o u g h h o r s e s and cows are both g r a z e r s , they are represen- t a t i v e o f two c o m p l e t e l y d i f f e r e n t mammalian f a m i l i e s w i t h d i f f e r e n t d i g e s t i v e s t r a t e g i e s . S u b s e q u e n t l y , though convergent i n t h e n i c h e t h e y e x p l o i t , t h e y r e m a i n d i v e r g e n t i n 'how' t h e y e x p l o i t t h e n i c h e . To d a t e , the general consensus, expressed by E v a n s (1977) i s t h a t h o r s e s a r e a p p r o x i m a t e l y t w o - t h i r d s as e f f i c i e n t i n t h e d i g e s t i o n o f f i b e r as r u m i n a n t s , and t h i s i s p r o b a b l y t r u e when e q u a l amounts o f f e e d s o f v a r y i n g f i b e r l e v e l s a r e f e d ad_ l i b i t u m . However, t h e r e a r e a number o f o t h e r f a c t o r s t o c o n s i d e r : 1) The h o r s e , w i t h more e f f i c i e n t d i r e c t u t i l i z a t i o n o f d i e t a r y s o l u b l e c a r b o h y d r a t e s and d i e t a r y p r o t e i n s o u r c e s , d e p e n d s l e s s on p r o d u c t s o f b a c t e r i a l d i g e s t i o n (and b a c t e r i a l p r o t e i n ) t h a n do c a t t l e . 2) The r a t e o f p a s s a g e o f d i g e s t a i s f a s t e r i n t h e h o r s e e n a b l i n g more f o o d t o be p r o c e s s e d i n t h e same time p e r i o d ( A l e x a n d e r , 1963; Vander Noot e^ ajL., 1967) than ruminants, w h i l e t h e c e l l u l o l y t i c a c t i v i t y o f t h e caecum r e m a i n s c o m p a r a b l e to t h a t o f t h e rumen ( B a l c h and Johnson, 1950 ( c i t e d i n Robinson and S l a d e , 1974); A l e x a n d e r , 1963). 3) The horse seems t o be a b l e t o s w i t c h e a s i l y from e x t e n s i v e - 86 - d e p e n d e n c e on raonogastric d i g e s t i o n t o e q u a l dependence on b a c t e r i a l d i g e s t i o n w i t h o u t an ' a d j u s t m e n t ' p e r i o d s u c h as t h a t r e q u i r e d by ruminants (Slade e£ a l . , 1970). 4) Under i d e n t i c a l e x p e r i m e n t a l c o n d i t i o n s , a b s o r p t i o n o f t h e major p r o d u c t s o f b a c t e r i a l f e r m e n t a t i o n , t h e v o l a t i l e f a t t y a c i d s , has been shown t o be s i m i l a r f o r t h e equine l a r g e i n t e s t i n e and t h e r u m i n a n t f o r e s t o m a c h ( S t e v e n s and S t e t t l e r , 1966; A r g e n z i o §_t_ a l . , 1974b). J a n i s (1976) s u g g e s t s t h e e q u i d s t r a t e g y may be e s s e n t i a l f o r t h e use o f h e r b a g e above a g i v e n f i b e r l e v e l and, whereas t h e g r e a t e r q u a n t i t y o f f o o d r e q u i r e d by h o r s e s under a d v e r s e c i r c u m s t a n c e s might be i n t e r p r e t e d as i n e f f i c i e n t , J a n i s p o i n t s o u t t h a t t h i s d i g e s t i v e s t r a t e g y e n a b l e s h o r s e s t o e x i s t on a d i e t w h i c h r u m i n a n t s o f s i m i l a r body s i z e s i m p l y c a n n o t m a i n t a i n t h e m s e l v e s . H o r s e s may w e l l p r o v e t o be ' o p t i m a l f o r a g e r s ' by b e c o m i n g s e l e c t i v e , i . e . a ' s p e c i a l i s t , ' when t h e r e i s f o o d i n abundance, and u n s e l e c t i v e , i . e . a ' g e n e r a l i s t , ' when f o o d i s s c a r c e (Pyke e t a l . , 1977; Nudds, 1980; B e l o v s k y , 1978). A number o f l a r g e h e r b i v o r e s have been shown t o have h i g h l y v a r i a b l e d i e t s , depending on t h e i r g e o g r a p h i c a l l o c a t i o n and e n v i r o n m e n t . T h i s a d a p t a b i l i t y i s t r u e f o r moose (Franzman, 1 9 7 8 ; P e e k , 1 9 7 4 ) , mule d e e r ( S h o r t , 1981) and e l k ( B o y d , 1978). D i e t s t u d i e s conducted on f e r a l h o r s e s and range c a t t l e , e i t h e r s e p a r a t e l y , t o g e t h e r or w i t h o t h e r s p e c i e s , have shown a d a p t a b i l i t y t o v a r i o u s environments (Casebeer and K o s s , 1970; Welsh, 1975; F e i s t and M c C u l l o u g h , 1976; L e u t h o l d , 1977; Berger, - 87 - 1 9 7 5 ) . F o r e x a m p l e , i n t h e Red D e s e r t o f Wyoming, y e a r - r o u n d d o m i n a n t f o o d s o f h o r s e s were A g r o p y r o n spp. (37%) and S t i p a s p p . ( 3 6 % ) ; and f o r c a t t l e , t h e o n l y s i n g l e s i g n i f i c a n t f o r a g e was A g r o p y r o n s p p . (54%) ( O l s e n and H a n s e n , 1 9 7 7 ) . I n d r y mountain range i n s o u t h e r n C o l o r a d o , t h e dominant summer forages o f c a t t l e were D a n t h o n i a s p p . and F e s c u e s p p . , w h i l e i n t h e f a l l C a r e x spp. i n c r e a s e d t o 58% o f t h e d i e t (Hansen and R e i d , 1978) . On t h e d r y , W h i t e Sands M i s s i l e Base i n s o u t h e r n New M e x i c o , t h e dominant foods o f h o r s e s i n the summer were dropseed ( S p o r o b o l u s spp.) and mesquite ( P r o s o p i s i u l i f l o r a ) and r u s s i a n t h i s t l e ( S a l s o l a k a l i ) i n w i n t e r (Hansen, 1976). The l i m i t e d a m ount o f c o m p a r a t i v e work c o n d u c t e d on h o r s e s and c a t t l e has shown a t e n d e n c y f o r t h e two h e r b i v o r e s t o s e l e c t s i m i l a r f o r a g e s w i t h i n t h e d i f f e r e n t e nvironments (Hubbard and Hansen, 1 9 7 6 ; S a l t e r , 1978; Hansen and C l a r k , 1977; Hansen a l . . 1977; O l s e n and H a n s e n , 1977). However, d i s t i n c t d i f f e r e n c e s have a l s o been seen i n a l l o f t h e s t u d i e s t o d a t e . I n g e n e r a l , t h e r e h a s b e e n l i t t l e a t t e m p t t o c o o r d i n a t e d i e t a n a l y s i s w i t h h a b i t a t - u s e and d i s t r i b u t i o n s t u d i e s ; and i n some instances so many d a t a have been 'lumped' t h a t t h e r e s u l t s a re o f l i t t l e v a l u e i n i n t e r p r e t i n g t h e s i g n f i c a n c e o f p o t e n t i a l a n i m a l i n t e r a c t i o n s . I n an a r t i c l e by Hansen e__ a i . , ( 1 9 7 7 ) , t h e a n n u a l d i e t a r y o v e r l a p f o r h o r s e s and cows was d e t e r m i n e d t o be 77 p e r c e n t , b u t f e c a l m a t e r i a l was c o l l e c t e d w i t h o u t r e g a r d t o s e a s o n , v e g e t a t i o n a n a l y s i s , a n i m a l movements, h a b i t a t p r e f e r e n c e o r a p p a r e n t age ( i . e . d e t e r i o r a t i o n ) o f f e c e s o t h e r t h a n i t b e i n g * undecomposed.' The 77 p e r c e n t s i m i l a r i t y i n d i e t c h o i c e may be v a l i d , b u t i t i s o n l y a s m a l l p a r t o f - 88 - t h e w h o l e p i c t u r e , as i n t e r a c t i o n s b e t w e e n h e r b i v o r e s may be s u b t l e . To d a t e , most d i e t s t u d i e s i n v o l v i n g f e r a l h o r s e s and o t h e r s p e c i e s have u t i l i z e d a ' s i m i l a r i t y i n d e x ' t o i n d i c a t e w h a t p e r c e n t a g e o f a p a i r o f d i e t s was i d e n t i c a l , and t h i s h a s b e e n d i r e c t l y i n t e r p r e t e d as d i e t o v e r l a p (Hubbard and H a n s e n , 1976; S a l t e r , 1978; Hansen and C l a r k , 1977; S e e g m i l l e r and Ohmart, 1981; Hansen et_ a l . , 1977; O l s e n and Hansen, 1977). The s i m i l a r i t y i n d e x i s b a s e d on summing t h e p e r c e n t a g e use i n common o f s h a r e d f o r a g e s by two o r more a n i m a l s p e c i e s ( A n t h o n y a n d S m i t h , 1977; O o s t i n g , 1 9 5 6 ) . I n t h e p r e s e n t s t u d y , a s i m i l a r i t y i n d e x f o r t h e average use o f t h e dominant 11 f o r a g e s s h a r e d by h o r s e s and cows was n e a r l y 70 p e r c e n t ( 6 9 . 7 % ) . I f e e l t h a t , u n l e s s r a t h e r s t r i n g e n t r e s t r i c t i o n s a r e p l a c e d on i n t e r p r e t a t i o n o f d i e t o v e r l a p , a p e r c e n t a g e use o f i d e n t i c a l p l a n t s p e c i e s may t e n d t o o b s c u r e t h i n k i n g a n d b i a s i t t o w a r d s m i s u s e o f t h e c o n c e p t o f c o m p e t i t i o n . The most c o n f u s i n g a s p e c t o f t h e c o n c e p t o f d i e t o v e r l a p i s t h a t i t i m p l i e s 'sameness,' i . e . t h e use o f t h e p a r t i c u l a r p l a n t s p e c i e s i n t h e same way, i n t h e same p l a c e and a t the same t i m e , whereas i n r e a l i t y , t h e s e are a l l independent compo- n e n t s , each o f which can g r e a t l y a l t e r t h e degree of 'sameness,' o r e s s e n t i a l l y n e g a t e i t a l t o g e t h e r . R e s u l t s f r o m my s t u d y summarized below, show t h a t d i f f e r e n c e s as w e l l as s i m i l a r i t i e s e x i s t i n t h e way h o r s e s , cows and moose u t i l i z e t h e i r environment, a n d i n d i c a t e s t h e need f o r a c o m p r e h e n s i v e a p p r o a c h t o t h e i n v e s t i g a t i o n o f h e r b i v o r e i n t e r a c t i o n s : - 89 - 1) I t was d e t e r m i n e d t h a t t h e d i s t r i b u t i o n o f h a b i t a t t y p e s i n t h e a p p r o x i m a t e l y 200 s q km o f t h e s t u d y s i t e was as f o l l o w s : Open F o r e s t ( 1 4 . 5 % ) , Semi-open F o r e s t (33.4%), C l o s e d F o r e s t ( 1 3 . 7 % ) , Meadow ( 2 8 . 1 % ) , Shrub c a r r ( 2 . 6 % ) , I n t e r f a c e Zone (5.7%) and 'Other' ( 1 . 9 % ) . 2) F o r e s t e d h a b i t a t , i n c l u d i n g Open F o r e s t , Semi-open F o r e s t , C l o s e d F o r e s t and I n t e r f a c e Zone, c o m p r i s e d a t o t a l o f 67.3 p e r c e n t o f t h e a r e a . 3) Moose a p p e a r e d t o be a t t r a c t e d t o F o r e s t H a b i t a t , as 78 p e r c e n t o f a l l f e c a l c o u n t s were made i n f o r e s t , whereas t h e h i g h e s t f e c a l c o u n t s f o r h o r s e s (65.3%) and cows ( 7 6 % ) , were made i n open h a b i t a t ( i . e . meadow and shrub c a r r ) . 4) Cows u s e d Meadow and I n t e r f a c e zone d i s p r o p o r t i o n a t e l y more t h a n i t was a v a i l a b l e , as d i d h o r s e s f o r Meadow and moose f o r Open and C l o s e d F o r e s t , and Shrub c a r r . 5) On t h e b a s i s o f f e c a l c o n c e n t r a t i o n s , h o r s e s were found t o use c o n t i g u o u s f o r e s t ( i . e . f o r e s t not i n c o n j u n c t i o n w i t h open h a b i t a t s ) more t h a n c ows, t h e l a t t e r t e n d i n g t o use I n t e r f a c e Zones ( i . e . f o r e s t i n c o n j u n c t i o n w i t h open h a b i t a t s ) more t h a n h o r s e s . 6) Moose were f o u n d t o use t h o s e h a b i t a t t y p e s ( i . e . Open, S e m i - o p e n and C l o s e d f o r e s t and S h r u b c a r r ) t h a t h o r s e s and, p a r t i c u l a r l y , cows used l e a s t . 7) A l l t h r e e h e r b i v o r e s showed some s p e c i e s s p e c i f i c s e l e c t i o n - 90 - f o r t h e a v a i l a b l e h a b i t a t s . 8) The f i v e t r a n s e c t s a l l v a r i e d s i g n i f i c a n t l y i n r e g a r d t o t h e r e l a t i v e amounts o f each h a b i t a t t y p e . 9) The h i g h e s t f e c a l c o u n t s f o r each h e r b i v o r e s p e c i e s were made on d i f f e r e n t t r a n s e c t s . Cows u s e d T-IV t h e most which had t h e h i g h e s t percentage of both meadow and i n t e r f a c e z o n e , h o r s e s used T-I t h e most, h a v i n g t h e l a r g e s t amount o f c o n t i g u o u s f o r e s t , and moose used T-V t h e most, wh i c h h a d t h e h i g h e s t a mount o f S h r u b c a r r , a h a b i t a t t y p e moose u s e d more t h a n i t s p e r c e n t a v a i l a b i l i t y would have i n d i c a t e d . 10) A l t h o u g h f e c a l c o u n t s i n d i c a t e d h o r s e s and cows s e l e c t e d f o r Meadow h a b i t a t most s t r o n g l y , t h e r e was no e v i d e n c e t h a t f o r a g e a v a i l a b i l i t y p l a y s a m a j o r r o l e i n h a b i t a t s e l e c t i o n , as t h e dominant f o r a g e i n b o t h horse and c a t t l e d i e t s i s a f o r e s t s p e c i e s (C. r u b e s c e n s ) , n o t f o u n d i n meadow h a b i t a t i n t h e s t u d y a r e a . 11) Based on food h a b i t s e s t a b l i s h e d f o r moose i n the l i t e r a t u r e , f o r a g e a v a i l a b i l i t y d i d not seem t o p l a y a dominant r o l e i n t h e i r h a b i t a t s e l e c t i o n , as t h e i r h i g h e s t f e c a l c o u n t s were made i n c l o s e d f o r e s t , w i t h a low f o o d a v a i l a b i l i t y . 12) When t h e l e v e l s o f s p a t i a l a s s o c i a t i o n were e v a l u a t e d , h o r s e s and c a t t l e were seen t o be a s s o c i a t e d , w h i l e c a t t l e and moose were s e e n t o be s e p a r a t e , and h o r s e s and moose were n e i t h e r a s s o c i a t e d nor s e p a r a t e . - 91 - 13) However, t h e f i v e t r a n s e c t graphs showed t h a t t h e concen- t r a t i o n o f use i n any g i v e n a r e a v a r i e d w i d e l y between s p e c i e s . 14) B a s e d on a p p r o x i m a t e l y e q u a l a v e r a g e d e f e c a t i o n r a t e s , c a t t l e p o p u l a t i o n s f o r t h e s t u d y a r e a were e s t i m a t e d t o be 9 0 % o f t h e t o t a l h o r s e , c a t t l e , moose h e r b i v o r e p o p u l a t i o n ; and h o r s e s and moose were e s t i m a t e d t o have p o p u l a t i o n l e v e l s o f 7.4% and 2.9%, r e s p e c t i v e l y . 15) D e s p i t e t h e l a r g e d i f f e r e n c e i n cow v e r s u s h o r s e popu- l a t i o n s , a n d t h e r e l a t i v e l y l i m i t e d amount o f meadow h a b i t a t ( 2 8 . 1 % ) , t h e t r a n s e c t g r a p h s showed numerous i n s t a n c e s where h o r s e s were h e a v i l y c o n c e n t r a t e d i n a r e a s c a t t l e used v e r y s l i g h t l y , and v i c e v e r s a . 16) A t r e n d was s e e n f o r h o r s e s t o use ' i s o l a t e d ' meadows ( i . e . t h o s e s u r r o u n d e d by e x t e n s i v e f o r e s t a r e a s ) and c o n t i g u o u s f o r e s t a r e a s much more t h a n c a t t l e , w h i c h t e n d e d t o use h a b i t u a l t r a v e l r o u t e s l i n k i n g meadows. S u b s e q u e n t l y , d e s p i t e t h e a s s o c i a t i o n i n d i c a t e d f o r h o r s e s and c a t t l e , t h e y were d i s t r i b u t e d d i f f e r e n t l y o v e r t h e s t u d y a r e a . 17) A ' l o n g - t e r m ' and ' s h o r t - t e r m ' comparison o f f e c a l m a t e r i a l was made on T-V, and two g r a p h s made, w h i c h were shown t o be s t a t i s t i c a l l y d i f f e r e n t . However, a l t h o u g h t h e g r a p h i c a l c o m p a r i s o n i n d i c a t e d t h a t o v e r a l l d i s t r i b u t i o n p a t t e r n s o f t h e t h r e e s p e c i e s were somewhat d i f f e r e n t o v e r t i m e , t h e r e were c e r t a i n c o n s i s t e n c i e s i n h a b i t a t - 92 - p r e f e r e n c e , i n c l u d i n g t h e pronounced use o f T-V by moose; t h e use by h o r s e s , but o n l y s l i g h t l y by c a t t l e , of ' i s o l a t e d ' meadows; and t h e 'absence' o f c a t t l e i n c o n t i g u o u s f o r e s t a r e a s . 18) D i e t a n a l y s i s i n d i c a t e d t h a t b o t h c a t t l e and h o r s e s , f r o m J u n e t h r o u g h S e p t e m b e r , were e x e r c i s i n g a f a i r l y h i g h d e g r e e o f s e l e c t i v i t y i n t h e i r c h o i c e o f f o r a g e s . E l e v e n f o r a g e s c o m p r i s e d 86% f o r h o r s e s , and 80% f o r c a t t l e , o f t h e t o t a l d i e t . These 11 f o r a g e s r e p r e s e n t e d o n l y 7.4% o f t h e 148 p l a n t s p e c i e s i d e n t i f i e d on t h e s t u d y a r e a . 19) D i f f e r e n c e s were f o u n d t o e x i s t i n t h e apparent f o r a g i n g p a t t e r n s o f h o r s e s and cows on t h e b a s i s o f the average amount consumed o f a p l a n t s p e c i e s p e r month; d i f f e r e n c e s i n t h e ways a f o r a g e p l a n t was used by h o r s e s and c a t t l e ; and whether a p a r t i c u l a r month was c r i t i c a l i n an i n t e r a c t i o n c o n t e x t . An a n a l y s i s o f t h e p r o p o r t i o n a t e use o f b o t a n i c a l groups by h o r s e s a n d c a t t l e showed a s i g n i f i c a n t d i f f e r e n c e i n t h e i r u t i l i z a t i o n p e r month o f g r a s s e s , r u s h - s e d g e s and ' o t h e r ' ( i . e . m a i n l y f o r b s and b r o w s e ) . In summer d i e t s , h o r s e s u t i l i z e d an average o f 61% grasses, whereas cows used an average o f 51% g r a s s e s , w i t h a h i g h e r p r o p o r t i o n o f s e d g e s , f o r b s and b r o w s e , i n d i c a t i n g cows were somewhat more s e l e c t i v e f o r f o r a g e w i t h lower f i b e r c o n t e n t . - 93 - 20) 21) 22) I n w i n t e r , h o r s e s d e c r e a s e d t h e number o f g r a s s , sedge and f o r b s p e c i e s e a t e n , and i n c r e a s e d t h e i r u t i l i z a t i o n o f s hrub m a t e r i a l up t o 14.7%. 23) A l s o , i n w i n t e r , h o r s e s i n c r e a s e d t h e i r d e pendence on R u s h - s e d g e s , f r o m 33.2% a v e r a g e i n t h e summer months, t o 48% i n w i n t e r . 24) B e c a u s e h o r s e s were c o n s i d e r e d a b l e t o e x p l o i t the g r e e n , i n n e r m a t e r i a l f o u n d i n s e d g e p l a n t s d u r i n g t h e w i n t e r , a n u t r i e n t a n a l y s i s was done on the s e l e c t e d green m a t e r i a l . T h i s was f o u n d t o have a C r u d e P r o t e i n l e v e l o f 6.5%, c o n s i s t e n t l y h i g h e r t h a n CP v a l u e s r e p o r t e d f o r g r a s s t y p e s found on t h e s t u d y s i t e , i n a 'weathered' c o n d i t i o n , c o l l e c t e d i n l a t e f a l l o r w i n t e r . Horse use o f g r a s s e s dropped d r a m a t i c a l l y i n t h e w i n t e r (from 60% t o 3 5 % ) . 25) A l t h o u g h h o r s e s u t i l i z e Carex a q u a t i l i s and Carex r o s t a t a h e a v i l y i n t h e w i n t e r (28.5%) w h i c h a r e b o t h g r a z e d by h o r s e s and cows d u r i n g t h e summer, the s i n g l e most important w i n t e r f o r a g e p l a n t f o r h o r s e s was Juncus b a l t i c u s , u t i l i z e d 1 7 . 2 % , and u s e d o n l y v e r y m o d e r a t e l y by e i t h e r h e r b i v o r e i n t h e summer. 26) The d i e t a n a l y s i s showed an i n c r e a s e i n t h e use o f c o n i f e r n e e d l e s d u r i n g the w i n t e r ( 6 . 3 % ) . T h i s was not c o n s i d e r e d a c c i d e n t a l because o f t h e d i s c r i m i n a t i n g a b i l i t y of horses. 27) The t o t a l d i e t s o f h o r s e s and c a t t l e r e p r e s e n t e d o n l y a f r a c t i o n o f t h e p l a n t s p e c i e s c o m p r i s i n g t h e b o t a n i c a l - 94 - g r o u p s f o u n d on t h e s t u d y s i t e . H o r s e s and cows u s e d o n l y 6 o f t h e 13 a v a i l a b l e r u s h - s e d g e s p e c i e s ; o f 88 f o r b s , h o r s e s u s e d 16 and c a t t l e 20, and o f 43 g r a s s s p e c i e s , h o r s e s used 22 and c a t t l e 19. 28) The d o m i n a n t f o r a g e f o r b o t h h o r s e s and c a t t l e was £. r u b e s c e n s . o r p i n e g r a s s , a l t h o u g h C a r e x a q u a t i l i s was u s e d o n l y s l i g h t l y l e s s by c a t t l e . 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N u t r i t i v e c h a r a c t e r i s t i c s o f g r a s s e s on s p r i n g r a n g e i n S o u t h C e n t r a l B.C. i n r e l a t i o n t o t i m e , h a b i t a t and f a l l g r a z i n g . Can. J . P l a n t S c i . 60: 131-137. W i l s o n , A. D. 1 9 6 9 . A r e v i e w o f b r o w s e i n t h e n u t r i t i o n o f g r a z i n g a n i m a l s . J . Range Manage., 22: 23-27. W o o d w a r d , S. L. a n d R. D. Ohmart. 1976. H a b i t a t use and f e c a l a n a l y s i s o f f e r a l b u r r o s (Equus a s i n u s ) , Chemehuevi M o u n t a i n s , C a l i f o r n i a . J . Range Manage., 29: 482-85. - 106 - APPENDIX 1: L i s t o f p l a n t s i d e n t i f i e d on t h e s t u d y s i t e d u r i n g 1979 and 1980. - 107 - L i s t o f p l a n t s i d e n t i f i e d on t h e s t u d y s i t e d u r i n g 1979 and 1980: A t o t a l o f 148 s p e c i e s were i d e n t i f i e d a t l e a s t t o s p e c i e s l e v e l . FAMILY GENDS SPECIES COMMON NAME 1. BETULACEAE ( b i r c h f a m i l y ) B e t u l a g l a n d u l o s a (2 v a r i e t i e s ) 2. BORAGINACEAE (borage f a m i l y ) M y o s o t i s s y l v a t i c a Lithospermum r u d e r a l e 3. CAPRIFOLIACEAE (h o n e y s u c k l e f a m i l y ) 4. CARYOPHYLLACEAE (pink f a m i l y ) L i n n a e a b o r e a l i s Symphoricarpos a l b u s C e r a s t i u m a r v e n s e A r e n a r i a l a t e r i f l o r a S t e l l a r i a l o n g i p e s L y c h n i s drummondii 5. COMPOSITAE ( a s t e r f a m i l y ) E r i g e r o n s p e c i o s u s E. p h i l a d e l p h i c u s E. l o n c o p h y l l u s E. compositus E. f l a g e l l a r i a Bog or Scrub b i r c h Forget-me-not Lemon-weed T w i n - f l o w e r Waxbe r ry/Snowbe r r y F i e l d chickweed B l u n t - l e a f sandwort L o n g - s t a l k s t a r w o r t Drummond campion Large p u r p l e f l e a b a n e P h i l a d e l p h i a f l e a b a n e S p e a r - l e a v e d f l e a b a n e C u t - l e a v e d f l e a b a n e A s t e r e a t o n i i A. c i l i o l a t u s A. c o n s p i c u o u s A. pansus Eaton's a s t e r L i n d l e y ' s a s t e r Large p u r p l e a s t e r T u f t e d w h i t e p r a i r i e a s t e r c o n t i n u e d . . . - 108 - A. c o m p e s t r i s A n t e n n a r i a p a r v i f o l i a A. m i c r o p h y l l a A. p u l c h e r r i m a A n t e n n a r i a n e g l e c t a A. u m b r i n e l l a A r t e m e s i a f r i g i d a S e n e c i o s t r e p t a n t h i f o l i u s S. canus C r e p i s t e c t o r u m A r n i c a c o r d i f o l i a A. f u l g e n s A. c h a m i s s o n i s v a r . i n c a n a A c h i l l e a m i l l e f o l i u m S o l i d a g o c a n a d e n s i s S. s p a t h u l a t a A g o s e r i s g l a u c a C i r s i u m s p. Taraxacum o f f i c i n a l e Tragopogon p r a t e n s i s - 109 - Meadow a s t e r N u t t a l l ' s p u s s y t o e s Rosy p u s s y t o e s Showy p u s s y t o e s F i e l d p u s s y t o e s Umber p u s s y t o e s P a s t u r e wormwood/ Sagewort Groundsel/Ragwort Narrow-leaved hawksbeard H e a r t - l e a v e d a r n i c a Twin a r n i c a Yarrow Canada/Meadow g o l d e n r o d S p i k e - l i k e g o l d e n r o d P a l e a g o s e r i s T h i s t l e Common d a n d e l i o n G o a t s b e a r d / S a l s i f y c o n t i n u e d . . . Appendix 1 ( c o n t . ) H i e r a c i u m s c o u l e r i Hounds-tongue G r i n d e l i a s g u a r r o s a C u r l y - g u p gumweed 6. CRASSULACEAE ( s t o n e c r o p f a m i l y ) 7. CRUCIFERAE (mustard f a m i l y ) 8. COPRESSACEAE ( j u n i p e r f a m i l y ) Sedum l a n c e o l a t u m D e s c u r a i n i a r i c h a r d s o n i i L e p i d i u m v i r g i n i c u m A r a b i s s p . J u n i p e r u s communis L a n c e - l e a v e d s t o n e c r o p Tansy mustard T a l l p e p p e r g r a s s R o c k c r e s s Common j u n i p e r 9. CYPERACEAE (sedge f a m i l y ) Carex a q u a t i l i s C. r o s t r a t a C. s i t c h e n s i s C. p r a e g r a c i l i s C. p e t a s a t a C. c o n c i n n o i d e s C. c o n c i n n a C. l a s i o c a r p a C. au r e a C. p a r r y a n a C. a t h e r o d e s C. p a c h y s t a c h y a E l e o c h a r i s s p . Water sedge Beaked sedge S i t k a sedge C l u s t e r e d f i e l d sedge L i d d o n ' s sedge N. w e s t e r n sedge Low N. sedge S l e n d e r sedge Golden sedge P a r r y sedge Awned sedge Thick-headed sedge S p i k e r u s h c o n t i n u e d . . . - 110 - Appendix 1 (con t . ) 10. ELAEAGNACEAE ( o l e a s t e r S h e p h e r d i a f a m i l y ) c a n a d e n s i s B u f f a l o b e r r y 11. ERICACEAE (heath f a m i l y ) P y r o l a c h l o r a n t h a P. (monensis) u n i f l o r a P. a s a r i f o l i a G r e e n i s h w i n t e r g r e e n Woodnymph Common p i n k w i n t e r g r e e n A r c t o s t a p h y l o s u v a - u r s i K i n n i k i n n i c k / B e a r b e r r y 12. GENTIANACEAE ( g e n t i a n G e n t i a n a a m a r e l l a N. g e n t i a n f a m i l y ) 13. GRAMINAE ( g r a s s f a m i l y ) O r y z o p s i s a s p e r i f o l i a 0. pungens R i c e g r a s s S t i p a r i c h a r d s o n i i S. s p a r t e a S. comata S. o c c i d e n t a l i s R i c h a r d s o n n e e d l e g r a s s P o r c u p i n e g r a s s N e e d l e - a n d - t h r e a d g r a s s W. n e e d l e g r a s s Beckmannia s p . S l o u g h g r a s s Hordeum jubatum H. brachyantherum C a l a m a g r o s t i s rubescens C. n e g l e c t a C. i n e x p a n s a F o x t a i l b a r l e y Meadow b a r l e y P i n e g r a s s Reedgrass N. Reedgrass c o n t i n u e d , - I l l - P u c c i n e l l i a n u t t a l l i a n a ( a i r o i d e s ) Poa j u n c i f o l i a P. i n t e r i o r P. p r a t e n s i s Agropyron s p i c a t u m A. t r a c h y c a u l u m (3 v a r i e t i e s ) S p a r t i n a g r a c i l i s K o e l e r i a m i c r a n t h a ( c r i s t a t a ) M u h l e n b e r g i a r i c h a r d s o n i s F e s t u c a saximontana ( o v i n a ) D a n t h o n i a i n t e r m e d i a Bromus anomalus A l o p e c u r u s a e q u a l i s G l y c e r i a b o r e a l i s Deschampsia c a e s p i t o s a A g r o s t i s s c a b r a N u t t a l l a l k a l i g r a s s A l k a l i b l u e g r a s s I n l a n d b l u e g r a s s Kentucky b l u e g r a s s Bluebunch wheatgrass B e a r d l e s s wheatgrass A l k a l i c o r d g r a s s June g r a s s Mat muhly Sheep f e s c u e Timber o a t g r a s s Nodding brome Short-awn f o x t a i l N. mannagrass T u f t e d h a i r g r a s s B e n t g r a s s - 112 - c o n t i n u e d . Appendix 1 (con t . ) 14. GROSSULARIACEAE ( c u r r a n t f a m i l y ) 15. HIPPURIDACEAE (mare's t a i l f a m i l y ) 16. IRIDACEAE ( I r i s f a m i l y ) 17. JUNCACEAE (Rush f a m i l y ) 18. JUNCAGINACEAE, (ar r o w g r a s s f a m i l y ) 19. LILIACEAE ( l i l y f a m i l y ) 20. LEGUMINOSAE (legume f a m i l y ) D i s t i c h l i s s p i c a t a v a r . s t r i c t a R i b e s sp. H i p p u r i s v u l g a r i s S i s y r i n c h i u m a n g u s t i f o l i u m Juncus b a l t i c u s T r i g l o c h i n maritimum S m i l a c i n a s t e l l a t a Zigadenus venenosus A l l i u m cernuum A s t r a g a l u s m i s e r A. a l p i n u s V i c i a americana L a t h y r u s o c h r o l e u c u s O x y t r o p i s d e f l e x a - 113 - D e s e r t s a l t g r a s s G ooseberry Common mare's t a i l B l u e - e y e d g r a s s B a l t i c r u s h S e a s i d e a r r o w g r a s s S t a r - f l o w e r e d Solomon's s e a l Meadow d e a t h camus Nodding o n i o n Timber m i l k v e t c h American v e t c h Y e l l o w pea Pendent-pod crazyweed c o n t i n u e d . . . Appendix 1 (c o n t . ) 21. LINACEAE ( f l a x f a m i l y ) Linum perenne v a r . l e w i s i i W i l d b l u e f l a x 22. ONAGRACEAE (eve. p r i m r o s e E p i l o b i u m f a m i l y ) a n g u s t i f o l i u m N a rrow-leaved f i r e w e e d 23. ORCHIDACEAE ( o r c h i d f a m i l y ) O r c h i s r o t u n d i f o l i a Round-leaved o r c h i d S p i r a n t h u s r o m a n z o f f i a n a L a d i e s - t r e s s e s 24. POLEMONIACEAE (phlox f a m i l y ) Polemonium h u m i l e J a c o b ' s l a d d e r 25. POLYGONACEAE (buckwheat f a m i l y ) Eriogonum h e r a c l e o i d e s Rumex sp. P a r s n i p - f l o w e r e d eriogonum D o c k / S o r r e l Polygonum s p . Smartweed/Knotweed 26. RANUNCULACEAE ( b u t t e r c u p f a m i l y ) Ranunculus c y m b a l a r i a Shore b u t t e r c u p Anemone m u l t i f i d a Globe anemone A q u i l e g i a b r e v i s t y l a B l u e columbine T h a l i c t r u m s p . Meadow rue 27. ROSACEAE (r o s e f a m i l y ) Geum t r i f l o r u m F r a g a r i a v i r g i n i a n a v a r . g l a u c a O l d man's w h i s k e r s W i l d s t r a w b e r r y c o n t i n u e d . . . - 114 - Appendix 1 ( c o n t . ) P o t e n t i l l a a n s e r i n a S i l v e r l e a f c i n q u e f o i l P. g r a c i l i s T a l l y e l l o w c i n q u e f o i l P. h i p p i a n a Wooly c i n q u e f o i l P. p e n s y l v a n i c u s P r a i r i e c i n q u e f o i l Rosa a c i c u l a r i s P r i c k l y r o s e S p i r a e a b e t u l i f o l i a F l a t - t o p p e d s p i r e a A m e l a n c h i e r sp. Saskatoon b e r r y 28. RUBIACEAE (madder f a m i l y ) G a l i u m b o r e a l e N. bedstraw 29. SALICACEAE ( w i l l o w f a m i l y ) S a l i x spp. W i l l o w 30. SANTALACEAE (sandalwood Comandra l i v i d a B a s t a r d t o a d f l a x f a m i l y ) C. u m b e l l a t a v a r . p a l l i d a P a l e comandra 31. SAXIFRAGACEAE ( s a x i f r a g e Heuchera c y l i n d r i c a R ound-leaf alum r o o t f a m i l y ) P a r n a s s i a p a l u s t r i s N. g r a s s o f p a r n a s s u s 32. SCROPHULARIACEAE Penstemon Shrubby penstemon ( f i g w o r t f a m i l y ) f r u t i c o s u s P. p r o c e r u s S l e n d e r b l u e penstemon C a s t i l l e j a m i n i a t a I n d i a n p a i n t b r u s h O r t h o c a r p u s l u t e u s Y e l l o w owl c l o v e r c o n t i n u e d . . . - 115 - Appendix 1 ( c o n t . ) 33. UMBELLIFERAE ( p a r s l e y f a m i l y ) Lomatium macrocarpum L a r g e - f r u i t lomatium 34. VIOLACEAE ( v i o l e t f a m i l y ) V i o l a adunca B l u e v i o l e t TREES: 35. PINACEAE (pi n e f a m i l y ) P i c e a g l a u c a P. e n g l e m a n n i i White s p r u c e Englemann s p r u c e P i n u s c o n t o r t a v a r . l a t i f o l i a L odgepole p i n e Pseudotsuga m e n z i e s i i Douglas f i r 36. SALICACEAE ( w i l l o w f a m i l y ) Populus t r e m u l o i d e s T r e m b l i n g aspen OTHER: LICHENS P e l t i g e r a s p . C l a d o n i a s p . R e i n d e e r l i c h e n - 116 - APPENDIX 2: Horse and C a t t l e D i e t s - d e t e r m i n e d by f e c a l a n a l y s i s - 117 - HORSE DIET - JUNE D i e t H - I D i e t H - 2 P l a n t S p e c i e s % D i e t p l a n t S p e c i e s % D i e t Poa j u n c i f o l i a 21.7 Poa j u n c i f o l i a 19.3 Juncus b a l t i c u s 13.5 C a l a m a g r o s t i s Carex a q u a t i l i s 12.3 rubescens 16.8 C a l a m a g r o s t i s Juncus b a l t i c u s 14.2 rubescens 12.1 Carex r o s t r a t a 11.0 S t i p a comata 11.0 Carex a q u a t i l i s 8.7 Carex r o s t a t a 5.4 S t i p a comata 7.8 Carex a t h e r o i d e s 4.8 Carex a t h e r o i d e s 6.3 Agropyron 4.4 G l y c e r i a b o r e a l i s 4.8 Hordeum jubatum 4.2 Hordeum jubatum 4.1 Carex c o n c i n o i d e s 3.0 Carex c o n c i n o i d e s 2.1 G l y c e r i a b o r e a l i s 2.6 M u h l e n b u r g i a S t i p a r i c h a r d s o n i i 2.3 r i c h a r d s o n i s 1.7 C a l a m a g r o s t i s S t i p a r i c h a r s o n i i 1.2 n e g l e c t a 0.9 A l o p e c u r u s a e q u a l i s 0.9 M u h l e n b u r g i a O r y z o p s i s a s p e r i f o l i a 0.7 r i c h a r d s o n i s 0.8 C a l a m a g r o s t i s n e g l e c t a 0.4 S t i p a o c c i d e n t a l i s 0.7 O r y z o p s i s a s p e r i f o l i a 0.3 - 118 - HORSE DIET - JUNE D i e t H - 3 P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s rubescens 19.5 Poa j u n c i f o l i a 17.4 Hordeum jubatum 9.9 Juncus b a l t i c u s 10.6 C a l a m a g r o s t i s n e g l e c t a 8.4 Carex a q u a t i l i s 8.1 Carex r o s t a t a 7.3 S t i p a comata 5.9 Carex a t h e r o i d e s 2.7 Rush-Sedges 2.0 S t i p a r i c h a r d s o n i i 1.5 O r y z o p s i s a s p e r i f o l i a 1.2 Poa I n t e r i o r 1.2 E l e o c h a r i s 1.0 Agropyron 0.7 Mu h l e n b u r g i a r i c h a r d s o n i s 0.6 A g r o s t i s 0.5 P o t e n t i l l a h i p p i a n a 0.5 D i e t H - 4 P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s rubescens 25.8 Poa j u n c i f o l i a 12.6 C a l a m a g r o s t i s n e g l e c t a 9.4 Carex a q u a t i l i s 7.1 Carex r o s t a t a 7.1 S t i p a comata 6.2 Hordeum jubatum 4.2 Juncus b a l t i c u s 4.0 G l y c e r i a b o r e a l i s 3.6 Carex c o n c i n o i d e s 3.6 Carex a t h e r o i d e s 3.4 Rush-Sedges 3.2 S t i p a o c c i d e n t a l i s 2.3 O r y z o p s i s a s p e r i f o l i a 1.8 S i s y r i n c h i u m 1.2 Mu h l e n b u r g i a r i c h a r d s o n i s 1.0 S t i p a r i c h a r d s o n i i 0.6 Geum t r i f l o r u m 0.5 A c h i l l e a m i l l e f o l i u m 0.4 A s t r a g a l u s m i s e r 0.3 H i p p u r i s v u l g a r i s 0.3 A s t e r c o n s p i c u o u s 0.3 S p a r t i n a g r a c i l i s 0.3 P o t e n t i l l a h i p p i a n a 0.3 Beckmanaia s p . 0.3 Taraxacum o f f i c i n a l e 0.2 - 119 - HORSE DIET - JULY D i e t H - 1 D i e t H - 2 P l a n t S p e c i e s % D i e t P l a n t S p e c i e s % D i e t Carex a q u a t i l i s 26.3 Poa j u n c i f o l i a 14. 3 Poa j u n c i f o l i a 11.9 C a l a m a g r o s t i s rubescens 12. 6 C a l a m a g r o s t i s rubescens 10.1 Carex a q u a t i l i s 11. 2 C a l a m a g r o s t i s n e g l e c t a 7.0 C a l a m a g r o s t i s n e g l e c t a 10. 5 Carex r o s t a t a 5.9 Juncus b a l t i c u s 9. 2 Carex c o n c i n o i d e s 5.4 S t i p a comata 8. 1 Juncus b a l t i c u s 4.8 Hordeum jubatum 6. 8 S t i p a comata 4.5 Carex r o s t a t a 6. 7 S t i p a o c c i d e n t a l i s 4.1 G l y c e r i a b o r e a l i s 3. 7 Hordeum jubatum 3.9 J u n c u s / E l e o c h a r i s 3. 5 G l y c e r i a b o r e a l i s 3.7 S t i p a o c c i d e n t a l i s 2. 4 Carex a t h e r o i d e s 2.8 Carex c o n c i n o i d e s 1. 9 Sedges-Rush 2.2 Sedges-Rush 1. 6 J u n c u s / E l e o c h a r i s 1.8 F r a g a r i a v i r g i n i a n a 1. 5 Rosa a c i c u l a r i s 1.2 Carex a t h e r o i d e s 1. 2 M u h l e n b u r g i a O r y z o p s i s a s p e r i f o l i a 1. 1 r i c h a r s o n i s 1.2 M u h l e n b u r g i a r i c h a r s o n i s 1. 0 O r y z o p s i s a s p e r f o l i a 1.1 P o t e n t i l l a h i p p i a n a 0. 9 Beckmania sp. 0.8 A s t r a g a l u s m i s e r 0. 6 Unknown f o r b 0.6 Unknown g r a s s 1. 2 S p a r t i n a g r a c i l i s 0.5 P o t e n t i l l a h i p p i a n a 0.1 A s t e r c o n s p i c u o u s 0.1 - 120 - HORSE DIET - JULY D i e t H - 3 D i e t H - 4 P l a n t S p e c i e s % D i e t P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s C a l a m a g r o s t i s rubescens 22.3 rubescens 18. 3 J u n c u s / E l e o c h a r i s 16.9 Carex a q u a t i l i s 16. 0 S t i p a o c c i d e n t a l i s 12.7 Carex r o s t a t a 10. 5 Carex a q u a t i l i s 7.9 J u n c u s / E l e o c h a r i s 7. 7 Poa j u n c i f o l i a 6.4 S t i p a comata 6. 8 Carex r o s t a t a 5.4 Carex c o n c i n o i d e s 5. 2 Hordeum jubatum 5.2 S t i p a o c c i d e n t a l i s 5. 2 S t i p a comata 4.8 Carex a t h e r o i d e s 4. 4 C a l a m g r o s t i s n e g l e c t a 4.1 G l y c e r i a b o r e a l i s 4. 2 S p a r t i n a g r a c i l i s 3.3 Poa j u n c i f o l i a 3. 6 G l y c e r i a b o r e a l i s 2.8 S t i p a r i c h a r d s o n i i 3. 6 S t i p a r i c h a r d s o n i i 2.5 C a l a m a g r o s t i s n e g l e c t a 3. 3 O r y z o p s i s a s p e r i f o l i a 2.5 Juncus b a l t i c u s 3. 0 F o r b f l o w e r 2.4 Hordeum jubatum 1. 7 Taraxacum o f f i c i n a l e 0.5 O r y z o p s i s a s p e r i f o l i a 1. 7 M u h l e n b u r g i a A c h i l l e a m i l l e f o l i u m 1. 4 r i c h a r d s o n i s 0.4 Rosa a c i c u l a r i s 1. 1 A r t e m e s i a f r i g i d a 1. 0 M u h l e n b u r g i a r i c h a r d s o n i s 0. 7 Unknown f o r b 0. 6 - 121 - HORSE DIET - AUGUST D i e t H - 1 P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s rubescens 27.6 Carex c o n c i n o i d e s 11.9 S t i p a o c c i d e n t a l i s 9.7 Carex r o s t a t a 8.9 Carex a t h e r o i d e s 6.9 Juncus b a l t i c u s 5.8 Poa j u n c i f o l i a 5.6 Carex a q u a t i l i s 4.6 Unknown Grass 4.3 A s t r a g a l u s m i s e r 4.1 Monocot Fo r b 3.0 O r y z o p s i s a s p e r i f o l i a 1.9 G l y c e r i a b o r e a l i s 1.3 Juncus s p. 1.0 Hordeum jubatum 0.9 C a l a m a g r o s t i s n e g l e c t a 0.8 Unknown Forb 0.7 A c h i l l e a m i l l e f o l i u m 0.7 S p i r a e a b e t u l i f o l i a 0.3 D i e t H - 3 P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s rubescens 46.2 Carex r o s t a t a 8.2 J u n c u s / E l e o c h a r i s 7.5 Carex a q u a t i l i s 6.5 S t i p a o c c i d e n t a l i s 6.4 Poa j u n c i f o l i a 5.6 S t i p a comata 3.2 Hordeum jubatum 2.7 Juncus b a l t i c u s 2.3 Juncus s p . 1.8 Carex c o n c i n o i d e s 1.7 S t i p a r i c h a r d s o n i s 1.5 O r y z o p s i s a s p e r i f o l i a 1.5 A s t r a g a l u s m i s e r 1.5 C a l a m a g r o s t i s n e g l e c t a 1.4 G l y c e r i a b o r e a l i s 1.2 M u h l e n b u r g i a r i c h a r d s o n i i 0.5 A n t e n n a r i a p a r v i f l o r a 0.3 D i e t H - 2 P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s rubescens 62.7 S t i p a o c c i d e n t a l i s 7.2 Carex a q u a t i l i s 7.2 Carex r o s t a t a 6.9 Juncus b a l t i c u s 5.9 S t i p a comata 2.5 Hordeum jubatum 1.7 G l y c e r i a b o r e a l i s 1.4 Carex a t h e r o i d e s 1.2 Poa j u n c i f o l i a 0.8 C a l a m a g r o s t i s n e g l e c t a 0.8 Juncus s p. 0.8 A c h i l l e a m i l l e f o l i u m 0.7 Unknown Forb 0.2 D i e t H - 4 P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s rubescens 31.8 Carex a q u a t i l i s 20.2 Carex r o s t a t a 12.9 Carex c o n c i n o i d e s 7.3 Juncus b a l t i c u s 5.9 Hordeum jubatum 4.6 Poa j u n c i f o l i a 4.0 G l y c e r i a b o r e a l i s 3.3 S t i p a o c c i d e n t a l i s 2.8 Carex a t h e r o i d e s 2.7 C a l a m a g r o s t i s n e g l e c t a 2.0 Juncus sp. 1.2 A s t r a g a l u s m i s e r 0.7 A r c t o s t a p h y l o s u v a - u r s i 0.6 - 122 - HORSE DIET - SEPTEMBER D i e t H - 1 P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s rubescens 47.3 Carex r o s t a t a 13.5 Carex a q u a t i l i s 9.4 Poa j u n c i f o l i a 7.0 Carex c o n c i n o i d e s 6.9 S t i p a o c c i d e n t a l i s 4.5 Juncus b a l t i c u s 2.9 A s t r a g a l u s m i s e r 1.7 Hordeum jubatum 1.6 Carex a t h e r o i d e s 1.2 G l y c e r i a b o r e a l i s 0.9 C a l a m a g r o s t i s n e g l e c t a 0.7 S p a r t i n a g r a c i l i s 0.5 S t i p a comata 0.5 O r y z o p s i s a s p e r i f o l i a 0.4 A r n i c a c o r d i f o l i a 0.3 Rosa a c i c u l a r i s 0.3 Forb F l o w e r 0.3 Monocot For b 0.1 D i e t H - 2 P l a n t S p e c i e s % Dj,?t C a l a m a g r o s t i s rubescens 45.3 Poa j u n c i f o l i a 8.1 Carex a q u a t i l i s 7.2 Carex r o s t a t a 7.2 S t i p a o c c i d e n t a l i s 5.4 C a l a m a g r o s t i s n e g l e c t a 3.7 J u n c u s / E l e o c h a r i s 2.9 Carex c o n c i n o i d e s 2.4 Hordeum jubatum 2.3 G l y c e r i a b o r e a l i s 2.3 Juncus sp. 2.1 S t i p a comata 1.7 O r y z o p s i s a s p e r i f o l i a 1.2 M u h l e n b u r g i a r i c h a r d s o n i s 1.1 Unknown For b 1.0 A s t r a g a l u s m i s e r 0.8 Unknown Grass 0.8 P o t e n t i l l a h i p p i a n a 0.8 A c h i l l e a m i l l e f o l i u m 0.7 S h e p e r d i a c a n a d e n s i s 0.7 Sedum l a n c e o l a t u m 0.6 J u n i p e r u s s p . 0.6 B e t u l a g l a n d u l o s a 0.4 Carex a t h e r o i d e s 0.4 Rosa a c i c u l a r i s 0.3 - 123 - HORSE DIET - SEPTEMBER D i e t H - 3 D i e t H - 4 P l a n t S p e c i e s % D i e t P l a n t S p e c i e s % D i e t Carex r o s t a t a 14.4 C a l a m a g r o s t i s C a l a m a g r o s t i s n e g l e c t a 11.4 rubescens 18.0 Carex a q u a t i l i s 10.6 S t i p a o c c i d e n t a l i s 10.1 C a l a m a g r o s t i s Carex a q u a t i l i s 7.9 rubescens 9.4 Poa j u n c i f o l i a 6.5 Carex c o n c i n o i d e s 9.1 Carex r o s t a t a 6.4 Poa j u n c i f o l i a 8.9 Juncus s p . 6.4 Unknown Shrubs 5.9 C a l a m a g r o s t i s n e g l e c t a 6.1 Juncus sp. 5.6 G l y c e r i a b o r e a l i s 5.4 G l y c e r i a b o r e a l i s 4.7 Hordeum jubatum 4.3 S t i p a o c c i d e n t a l i s 3.8 A s t a g a l u s m i s e r 3.3 Unknown Forb 2.7 J u n c u s / E l e o c h a r i s 3.0 S t i p a comata 2.6 M u h l e n b u r g i a Monocot Fo r b 2.1 r i c h a r d s o n i s 2.6 Grass Seed 1.8 S t i p a comata 2.5 M u h l e n b u r g i a Eriogonum s p . 2.3 r i c h a r d s o n i s 1.5 Unknown Grass 1.9 Hordeum jubatum 1.4 P o t e n t i l l a h i p p i a n a 1.9 Unknown Grass 1.2 Carex c o n c i n o i d e s 1.6 Carex s p . 0.9 P i n u s c o n t o r t a 1.6 A s t r a g a l u s m i s e r 0.7 Unknown Forb 1.6 V i c i a s p . 0.7 Beckmania s p . 1.4 P i n u s c o n t o r t a 0.6 F e r n 1.2 Taraxacum o f f i c i n a l e 0.9 Monocot f o r b 0.9 V i c i a s p. 0.8 Unknown shrub 0.8 A r c t o s t a p h y l o s u v a - u r s i 0.6 - 124 - HORSE DIETS - WINTER D i e t H - 1 P l a n t S p e c i e s % D i e t Carex r o s t a t a 17.9 Juncus sp. 15.5 Carex a q u a t i l i s 14.1 G l y c e r i a b o r e a l i s 8.3 Unknown Grass 6.9 C a l a m a g r o s t i s rubescens 6.5 S t i p a o c c i d e n t a l i s 6.0 P i n u s c o n t o r t a 4.5 Rosa a c i c u l a r i s 3.8 Poa j u n c i f o l i a 2.6 C a l a m a g r o s t i s n e g l e c t a 2.5 A r t e m e s i a f r i g i d a 2.0 J u n c u s / E l e o c h a r i s 1.9 B e t u l a g l a n d u l o s a 1.6 Carex c o n c i n o i d e s 1.5 A r c t o s t a p h y l o s u v a - u r s i 1.3 P i c e a s i t c h e n s i s 1.2 S a l i x s p . 1.1 Unknown shrub 0.5 F e r n 0.3 D i e t H - 2 P l a n t S p e c i e s % Diet Juncus s p . 18.8 Carex r o s t a t a 14.4 Carex a q u a t i l i s 10.6 Unknown Grass 9.4 S t i p a comata 7.9 C a l a m a g r o s t i s rubescens 7.5 Unknown Shrub 5.3 P i n u s c o n t o r t a 4.3 G l y c e r i a b o r e a l i s 4.0 S t i p a o c c i d e n t a l i s 4.0 C a l a m a g r o s t i s n e g l e c t a 3.1 P i c e a s i t c h e n s i s 2.7 Rosa a c i c u l a r i s 2.6 F e r n 2.2 A n t e n n a r i a p a r v i f l o r a 1.0 Carex c o n c i n o i d e s 0.8 Poa j u n c i f o l i a 0.7 A r c t o s t a p h y l o s u v a - u r s i 0.7 - 125 - CATTLE DIETS - JUNE D i e t C - 1 P l a n t S p e c i e s % D i e t Poa j u n c i f o l i a 10.7 Carex r o s t a t a 10.7 C a l a m a g r o s t i s rubescens 10.1 Rush-Sedges 9.8 C a l a m a g r o s t i s n e g l e c t a 9.7 Hordeum jubatum 9.3 Juncus b a l t i c u s 7.3 Carex a q u a t i l i s 6.3 G l y c e r i a b o r e a l i s 4.8 S t i p a comata 4.1 Carex c o n c i n o i d e s 3.7 Carex a t h e r o i d e s 2.1 Mu h l e n b u r g i a r i c h a r d s o n i s 2.0 H i p p u r i s v u l g a r i s 1.7 S t i p a r i c h a r d s o n i i 1.3 S o l i d a g o s p a t h u l a t a 1.0 V i o l a adunca 0.9 O r y z o p s i s a s p e r i f o l i a 0.9 F r a g a r i a v i r g i n i a n a 0.7 P o t e n t i l l a h i p p i a n a 0.6 S t i p a o c c i d e n t a l i s 0.4 A s t r a g a l u s m i s e r 0.4 A n t e n n a r i a p a r v i f o l i a 0.3 A s t e r c o n s p i c u o u s 0.3 A c h i l l e a m i l l e f o l i u m 0.3 Geum t r i f l o r u m 0.3 Rosa a c i c u l a r i s 0.3 D i e t C - 2 P l a n t S p e c i e s % D i e t Poa j u n c i f o l i a 13.9 Carex a q u a t i l i s 13.2 C a l a m a g r o s t i s rubescens 12.9 Carex c o n c i n o i d e s 12.6 Rush-Sedges 9.0 Carex r o s t a t a 6.3 Hordeum jubatum 5.3 C a l a m a g r o s t i s n e g l e c t a 4.3 S t i p a comata 4.0 G l y c e r i a b o r e a l i s 4.0 Juncus b a l t i c u s 3.9 P o t e n t i l i a h i p p i a n a 2.8 S p a r t i n a g r a c i l i s 1.4 Carex a t h e r o i d e s 1.3 S t i p a o c c i d e n t a l i s 1.1 O r y z o p s i s a s p e r i f o l i a 1.0 S t i p a r i c h a r d s o n i i 0.9 Mu h l e n b u r g i a r i c h a r d s o n i s 0.7 F r a g a r i a v i r g i n i a n a 0.6 Geum t r i f l o r u m 0.2 S o l i d a g o s p a t h u l a t a 0.2 A s t e r c o n s p i c u o u s 0.1 A n t e n n a r i a p a r v i f o l i a 0.1 A s t r a g a l u s m i s e r 0.1 - 126 - CATTLE DIETS - JUNE D i e t C - 3 P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s rubescens 14.6 Carex a q u a t i l i s 12.7 Carex r o s t a t a 9.8 Poa j u n c i f o l i a 9.7 Hordeum jubatum 7.5 C a l a m a g r o s t i s n e g l e c t a 8.5 G l y c e r i a b o r e a l i s 5.5 S t i p a comata 5.0 Carex c o n c i n o i d e s 3.9 S t i p a o c c i d e n t a l i s 3.8 Juncus b a l t i c u s 3.3 A c h i l l e a m i l l e f o l i u m 2.4 P o t e n t i l l a h i p p i a n a 2.4 Sedge-Rush 1.7 S t i p a r i c h a r d s o n i i 1.5 Carex a t h e r o i d e s 1.3 Geum t r i f l o r u m 0.8 Agropyron t r a c h y c a u l u m 0.7 C e r a s t i u m a r v e n s e 0.6 A s t r a g a l u s m i s e r 0.6 F r a g a r i a v i r g i n i a n a 0.5 Unknown f o r b 0.5 A r n i c a c o r d i f o l i a 0.4 Mu h l e n b u r g i a r i c h a r s o n i s 0.4 S p a r t i n a g r a c i l i s 0.3 Taraxacum o f f i c i n a l e 0.3 S o l i d a g o s p a t h u l a t a 0.3 Heuchera c y l i n d r i c a 0.3 Penstemon p r o c e r u s 0.3 Sedum l a n c e o l a t u m 0.3 A s t e r c o n s p i c u o u s 0.1 D i e t C - 4 P l a n t S p e c i e s % D i e t Carex a q u a t i l i s 14. 7 Carex r o s t a t a 11. 2 Carex a t h e r o i d e s 9. 6 C a l a m a g r o s t i s n e g l e c t a 9. 5 Hoedeum jubatum 8. 5 Poa j u n c i f o l i a 8. 1 Carex c o n c i n o i d e s 8. 0 C a l a m a g r o s t i s rubescens 7. 4 Juncus b a l t i c u s 6. 1 S t i p a comata 4. 8 G l y c e r i a b o r e a l i s 3. 6 S t i p a o c c i d e n t a l i s 2. 4 Sedge-Rush 2. 4 S p a r t i n a g r a c i l i s 1. 0 S i s y r i n c h i u m a n g u s t i f o l i u m 0. 7 A r n i c a c o r d i f o l i a 0. 7 M u h l e n b u r g i a r i c h a r d s o n i s 0. 5 A s t r a g a l u s m i s e r 0. 4 A s t e r c o n s p i c u o u s 0. 3 P o t e n t i l l a h i p p i a n a 0. 1 - 127 - CATTLE DIETS - JULY D i e t q - l D i e t q - 2 P l a n t S p e c i e s % D i e t P l a n t S p e c i e s % D i e t Carex a q u a t i l i s 15.8 qarex a q u a t i l i s 21.9 Sedge-Rush 10.8 G l y c e r i a b o r e a l i s 9.2 C a l a m a g r o s t i s n e g l e c t a 10.8 C a l a m a g r o s t i s rubescens 8.5 C a l a m a g r o s t i s rubescens 10.0 S t i p a o c c i d e n t a l i s 7.8 G l y c e r i a b o r e a l i s 8.4 Sedge-Rush 7.5 Hordeum jubatum 6.3 S t i p a comata 7.4 S t i p a comata 5.7 C a l a m a g r o s t i s n e g l e c t a 5.5 Carex r o s t a t a 5.1 Hordeum jubatum 4.8 S t i p a o c c i d e n t a l i s 4.9 Juncus b a l t i c u s 4.2 Carex c o n c i n o i d e s 4.3 Carex r o s t a t a 4.1 Poa j u n c i f o l i a 4.2 Poa j u n c i f o l i a 3.3 Juncus b a l t i c u s 2.8 Carex c o n c i n o i d e s 2.3 Rosa a c i c u l a r i s 2.4 Carex a q u a t i l i s 2.0 P o t e n t i l l a h i p p i a n a 1.6 M u h l e n b u r g i a A s t r a g a l u s m i s e r 1.2 r i c h a r d s o n i s 1.9 A c h i l l e a m i l l e f o l i u m 1.0 O r y z o p s i s a s p e r i f o l i a 1.7 Carex a t h e r i o d e s 1.0 Mosses 1.1 A r n i c a c o r d i f o l i a 0.8 P o t e n t i l l a h i p p i a n a 0.8 S a l i x s p. 0.7 Rosa a c i c u l a r i s 0.8 Unknown f o r b 0.7 Sedum l a n c e o l a t u m 0.8 Geum t r i f l o r u m 0.6 S t i p a r i c h a r d s o n i i 0.7 F r a g a r i a v i r g i n i a n a 0.6 S p a r t i n a g r a c i l i s 0.7 S p a r t i n a g r a c i l i s 0.3 A r c t o s t a p h y l o s u v a - u r s i 0.7 Penstemon p r o c e r u s 0.5 A n t e n n a r i a p a r v i f o l i a 0.5 A s t e r c o n s p i c u o u s 0.2 Taraxacum o f f i c i n a l e 0.2 B e t u l a g l a n d u l o s a 0.2 A c h i l l e a m i l l e f o l i u m 0.2 S a l i x s p . 0.2 Unknown f o r b 0.2 Geum t r i f l o r u m 0.1 - 128 - CATTLE DIETS - JULY Diet C - 3 Diet C - 4 Plant Species % Diet Plant Species % Diet Carex a q u a t i l i s 20.9 Carex a q u a t i l i s 21.2 Carex rostata 14.3 Carex rostata 13.1 Calamagrostis rubescens 10.0 Calamagrostis neglecta 13.0 Hordeum jubatum 7.9 Calamagrostis rubescens 11.8 Calamagrostis neglecta 7.6 Hordeum jubatum 9.5 Glyceria b o r e a l i s 6.8 Stipa occidentalis 9.0 Rush-Sedge 5.8 Glyceria borealis 8.7 Stipa o c c i d e n t a l i s 4.5 Carex atheroides 4.5 Poa j u n c i f o l i a 3.8 Stipa comata 2.9 Stipa comata 3.5 Carex concinoides 1.1 Muhlenburgia Juncus ba l t i c u s 0.8 richardsonis 2.7 A c h i l l e a millefolium 0.8 Juncus b a l t i c u s 2.0 Muhlenburgia Carex concinoides 1.6 richardsonis 0.6 Oryzopsis a s p e r i f o l i a 1.6 P o t e n t i l l a hippiana 0.5 Stipa r i c h a r d s o n i i 1.4 Aster conspicuous 0.5 P o t e n t i l l a hippiana 1.0 Arctostaphylos uva-ursi 0.3 Carex atheroides 0.9 Taraxacum o f f i c i n a l e 0.3 Mosses 0.8 Astragalus miser 0.3 Unknown forb 0.6 Salix sp. 0.2 Antennaria p a r v i f o l i a 0.5 Fragaria v i r g i n i a n a 0.2 A c h i l l e a millefolium 0.3 Antennaria p a r v i f o l i a 0.2 Rosa a c i c u l a r i s 0.3 Fragaria v i r g i n i a n a 0.2 Aster conspicuous 0.2 Arnica c o r d i f o l i a 0.2 Arctistaphylos uva-ursi 0.6 - 129 - CATTLE DIET - AUGUST D i e t C - X P l a n t S p e c i e s Carex a q u a t i l i s G l y c e r i a b o r e a l i s S t i p a comata C a l a m g r o s t i s rubscens Carex a t h e r o i d e s Juncus b a l t i c u s Poa j u n c i f o l i a Carex r o s t a t a C a l a m g r o s t i s n e g l e c t a Carex c o n c i n o i d e s J u n c u s / E l e o c h a r i s Hordeum jubatum Unknown f o r b S t i p a o c c i d e n t a l i s S t i p a r i c h a r d s o n i i P o t e n t i l l a h i p p i a n a O r y z o p s i s a s p e r i f o l i a S a l i x s p. A s t r a g a l u s m i s e r S h e p h e r d i a c a n a d e n s i s Geum t r i f l o r u m Rosa a c i c u l a r i s A s t e r c o n s p i c u o u s A c h i l l e a m i l l e f o l i u m D i e t C - 2 % D i e t P l a n t S p e c i e s % D i e t 16.2 C a l a m g r o s t i s rubscens 25.1 10.8 Carex a q u a t i l i s 16.6 10.0 Poa j u n c i f o l i a 5.6 9.3 S t i p a comata 5.0 8.7 S t i p a o c c i d e n t a l i s 4.9 8.1 Carex r o s t a t a 3.5 7.3 Carex c o n c i n o i d e s 3.5 5.9 J u n c u s / E l e o c h a r i s 3.4 4.9 Hordeum jubatum 3.2 3.8 Juncus b a l t i c u s 2.7 2.9 S h e p h e r d i a c a n a d e n s i s 2.6 2.7 C a l a m a g r o s t i s n e g l e c t a 2.2 2.0 O r y z o p s i s a s p e r i f o l i a 2.2 1.6 Rosa a c i c u l a r i s 1.7 1.3 S i s y r i n c h i u m 0.8 a n g u s t i f o l i u m 1.7 0.7 M u h l e n b u r g i a 0.7 r i c h a r d s o n i s 1.4 0.6 V i c i a a mericana 1.4 0.5 S t i p a r i c h a r d s o n i i 1.3 0.4 P o t e n t i l l a h i p p i a n a 1.3 0.4 D i c o t f o r b 1.2 0.2 Geum t r i f l o r u m 1.2 0.2 S a l i x s p. 1.2 F r a g a r i a v i r g i n i a n a 1.1 T h a l i c t r u m s p . 1.1 Monocot f o r b 1.0 A n t e n a r i a s p . 0.7 S p i r a e a b e t u l i f o l i a 0.7 A r c t o s t a p h y l o s u v a - u r s i 0.5 G l y c e r i a b o r e a l i s 0.5 B e t u l a 0.3 A s t e r c o n s i p c u o u s 0.3 A c h i l l e a m i l l e f o l i u m 0.2 S o l i d a g o s p a t h u l a t a 0.2 P o t e n t i l l a h i p p i a n a 0.2 Moss 0.1 - 130 - CATTLE DIET - AUGUST D i e t C - 3 ) D i e t C - 4 P l a n t S p e c i e ? % D i e t P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s C a l a m a g r o s t i s rubscens 22.2 rubscens 28. 1 Unknown form 15.0 Poa j u n c i f o l i a 11. 6 Carex a q u a t i l i s 9.0 Carex r o s t a t a 10. 8 S t i p a o c c i d e n t a l i s 6.3 Carex c o n c i n o i d e s 8. 2 Juncus b a l t i c u s 5.8 S t i p a o c c i d e n t a l i s 6. 5 Carex r o s t a t a 4.5 S t i p a comata 5. 2 C a l a m a g r o s t i s n e g l e c t a 4.5 Rosa a c i c u l a r i s 3. 4 Poa j u n c i f o l i a 3.0 Carex a q u a t i l i s 3. 3 S t i p a comata 2.8 Juncus b a l t i c u s 2. 9 A c h i l l e a m i l l e f o l i u m 2.7 G l y c e r i a b o r e a l i s 2. 7 Carex c o n c i n o i d e s 2.3 S h e p h e r d i a c a n a d e n s i s 2. 6 S i s y r i n c h i u m M u h l e n b u r g i a a n g u s t i f o l i u m 2.2 r i c h a r d s o n i s 2. 4 Hordeum jubatum 2.1 O r y z o p s i s a s p e r i f o l i a 1. 7 S p i r a e a b e t u l i f o l i a 2.0 P o t e n t i l l a h i p p i a n a 1. 6 P i n u s c o n t o r t a v a r . S t i p a r i c h a r d s o n i i 1. 3 l a t i f o l i a 1.8 C a l a m a g r o s t i s n e g l e c t a 1. 2 Taraxacum o f f i c i n a l e 1.7 Geum t r i f l o r u m 1. 2 Mu h l e n b u r g i a Unknown f o r b 1. 2 r i c h a r d s o n i s 1.4 J u n c u s / E l e o c h a r i s 0. 8 O r y z o p s i s a s p e r i f o l i a 1.4 Beckmania 0. 7 Forb f l o w e r 1.3 A c h i l l e a m i l l e f o l i u m 0. 6 P o t e n t i l l a h i p p i a n a 1.3 Hordeum jubatum 0. 6 A r c t o s t a p h y l o s u v a - u r s i 1.1 A s t r a g a l u s m i s e r 0. 6 V i c i a americana 1.1 F r a g a r i a v i r g i n i a n a 0. 4 S a l i x s p. 1.0 S o l i d a g o s p a t h u l a t a 0. 3 F r a g a r i a v i r g i n i a n a 1.0 G l y c e r i a b o r e a l i s 1.0 S h e p h e r d i a c a n a d e n s i s 0.9 Geum t r i f l o r u m 0.4 • A s t r a g a l u s m i s e r 0.2 131 - CATTLE DIET - SEPTEMBER D i e t C - 1 P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s rubescens 16.5 Carex r o s t a t a 9.6 A s t r a g a l u s m i s e r 8.4 Carex a q u a t i l i s 8.0 G l y c e r i a b o r e a l i s 7.7 S t i p a o c c i d e n t a l i s 7.2 O r y z o p s i s a s p e r i f o l i a 6.4 Juncus b a l t i c u s 5.5 A c h i l l e a m i l l e f o l i u m 4.9 Hordeum jubtaum 4.9 Carex c o n n c i n o i d e s 4.3 S t i p a comata 3.3 V i c i a s p . 2.2 Beckmannia s p . 1.7 Mu h l e n b u r g i a r i c h a r d s o n i s 1.7 C e r a s t i u m a r v e n s e 1.5 Taraxacum o f f i c i n a l e 1.3 S i s y r i n c h i u m a n g u s t i f o l i u m 1.2 F r a g a r i a v i r g i n i a n a 1.0 V i o l a adunca 1.0 Poa j u n c i f o l i a 1.0 Geum t r i f l o r u m 0.5 S a l i x s p. 0.2 D i e t C - 2 P l a n t S p e c i e s % D i e t C a l a m a g r o s t i s rubescens 29.0 Carex a q u a t i l i s 15.6 Juncus b a l t i c u s 7.1 Poa j u n c i f o l i a 6.0 S t i p a comata 5.3 S t i p a o c c i d e n t a l i s 5.2 Carex r o s t a t a 4.4 G l y c e r i a b o r e a l i s 3.4 Hordeum jubatum 3.9 V i c i a sp. 3.1 S t i p a r i c h a r d s o n i i 2.4 F r a g a r i a v i r g i n i a n a 2.0 P o t e n t i l l a h i p p i a n a 1.6 S h e p e r d i a c a n a d e n s i s 1.5 V i o l a adunca 1.2 O r y z o p s i s a s p e r i f o l i a 1.1 A s t r a g a l u s m i s e r 0.9 S p i r a e a b e t u l i f o l i a 0.9 Beckmannia sp. 0.9 A c h i l l e a m i l l e f o l u m 0.8 P i n u s c o n t o r t a 0.8 Carex c o n c i n o i d e s 0.8 Mu h l e n b u r g i a r i c h a r d s o n i s 0.7 A r n i c a c o r d i f o l i a 0.5 C a l a m a g r o s t i s n e g l e c t a 0.5 Geum t r i f l o r u m - 132 - CATTLE DIET - SEPTEMBER Diet C - 3 P l a n t S p e c i e s % D i e t Carex a q u a t i l i s 29. 6 C a l a m a g r o s t i s rubescens 25. 5 Carex c o n c i n o i d e s 7. 6 G l y c e r i a b o r e a l i s 4. 1 Juncus b a l t i c u s 3. 7 S t i p a comata 3. 6 S t i p a o c c i d e n t a l i s 3. 5 Hordeum jubatum 2. 8 C a l a m a g r o s t i s n e g l e c t a 2. 7 Rosa a c i c u l a r i s 2. 4 Poa j u n c i f o l i a 2. 3 V i c i a s p. 1. 9 Carex r o s t a t a 1. 7 S a l i x s p. 1. 4 Carex a t h e r o i d e s 1. 4 A r c t o s t a p h y l o s u v a - u r s i 1. 3 S p a r t i n a g r a c i l i s 1. 1 A r n i c a c o r d i f o l i a 1. 0 O r y z o p s i s a s p e r i f o l i a 0. 8 S h e p e r d i a c a n a d e n s i s 0. 6 J u n c u s / E l e o c h a r i s 0. 6 P o t e n t i l l a h i p p i a n a 0. 4 D i e t C - 4 P l a n t S p e c i e s % D i e t Carex a q u a t i l i s 31.8 C a l a m a g r o s t i s rubescens 23.5 Carex r o s t a t a 5.8 Hordeum jubatum 5.1 S t i p a comata 5.0 Juncus b a l t i c u s 4.6 Poa j u n c i f o l i a 3.9 S t i p a o c c i d e n t a l i s 3.6 S a l i x s p . 2.9 Carex c o n c i n o i d e s 2.6 Unknown f o r b 2.5 P i n u s c o n t o r t a 2.1 Unknown Grass 2.0 S h e p e r d i a c a n a d e n s i s 1.8 G l y c e r i a b o r e a l i s 1.0 A s t r a g a l u s m i s e r 1.0 C a l a m a g r o s t i s n e g l e c t a 0.8 - 133 - APPENDIX 3: P a i r e d ' t ' t e s t f o r ' L o n g - t e r m ' v e r s u s ' S h o r t - t e r m ' f e c a l c o u n t s on T r a n s e c t V - 134 - Cows v e r s u s Cows D a t a f o r P a i r e d ' t ' t e s t ( w i t h 36 d . f . ) - A r c s i n T r a n s f o r m a - t i o n - ( S o k a l and R o h l f , 1969: 205) f o r comparison of 'Long-term' ( o l d ) v e r s u s ' S h o r t - t e r m ' (new) f e c a l c o u n t s r e c o r d e d on T r a n s e c t V i n .5 km u n i t s : F e c a l Counts P r o p , o f Tot No. O l d T r a n s e c t - O l d 1. 286 5.4 2. 289 5.4 3. 425 7.9 4. 416 7.8 5. 231 4.3 6. 259 4.9 7. 292 5.5 8. 384 7.2 9. 252 4.7 10. 224 4.2 11. 234 4.4 12. 206 3.9 13. 165 3.1 14. 175 3.3 15. 165 3.1 16. 181 3.4 17. 33 .62 18. 24 .5 19. 10 .2 20. 4 .1 21. 1 .02 22. 14 .3 23. 10 .2 24. 1 .02 25. 4 .1 26. 3 .1 27. 2 .04 28. 13 .2 29. 8 .2 30. 20 .4 31. 167 3.1 32. 100 1.9 33. 38 .7 F e c a l Counts P r o p , o f T o t . No. New Transect-New 1. 116 7.7 2. 116 7.7 3. 177 11.8 4. 164 10.9 5. 110 7.3 6. 129 8.6 7. 102 6.8 8. 33 .1 9. 68 4.5 10. 61 4.1 11. 76 5.1 12. 9 .6 13. 5 .3 14. 20 1.3 15. 61 4.1 16. 9 .6 17. 1 .1 18. 0 0 19. 0 0 20. 0 0 21. 0 0 . 22. 0 0 23. 0 0 24. 0 0 25. 0 0 26. 0 0 27. 1 .1 28. 4 .23 29. 1 .1 30. 0 0 31. 8 .5 32. 16 1.1 33. 0 0 c o n t i n u e d . . . - 135 - Appendix 3 - Cows v e r s u s Cows (con t . ) 34. 41 .8 34. 2 .1 35. 94 1.8 35. 15 1.0 36. 287 5.4 36. 38 2.5 37. 278 5.2 37. 163 10.8 5336 1505 d = 55.38 ( d ) 2 = 3066.94 d = 1.497 d2 = 622.23 t » 2.35 - 136 - APPENDIX 3: Horses v e r s u s Horses F e c a l Counts P r o p , o f T o t . F e c a l Counts P r o p , o f No. O l d T r a n s e c t - O l d No. New T r a n s e c t 1. 23 1.2 1. 0 0 2. 19 1.0 2. 0 0 3. 33 1.8 3. 1 2.3 4. 23 1.2 4. 0 0 5. 15 .8 5. 0 0 6. 15 .8 6. 0 0 7. 9 .49 7. 0 0 8. 29 1.6 8. 1 2.3 9. 61 3.3 9. 0 0 10. 49 2.6 10. 2 4.6 11. 16 .86 11. 5 11.4 12. 164 8.8 12. 0 0 13. 129 6.9 13. 0 0 14. 134 7.2 14. 10 22.7 15. 245 13.2 15. 3 6.8 16. 102 • 5.5 16. 3 6.8 17. 40 2.2 17. 0 0 18. 30 1.6 18. 0 0 19. 22 1.2 19. 0 0 20. 3 .2 20. 0 0 21. 0 0 21. 0 0 22. 4 .2 22. 0 0 23. 220 11.9 23. 7 15.9 24. 32 1.7 24. 4 9.1 25. 96 5.2 25. 3 6.8 26. 14 .8 26. 0 0 27. 5 .27 27. 0 0 28. 54 2.9 28. 0 0 29. 27 1.5 29. 0 0 30. 12 .65 30. 1 2.3 31. 37 2.0 31. 0 0 32. 28 1.5 32. 0 0 33. 20 1.1 33. 0 0 34. 20 1.1 34. 0 0 35. 25 1.4 35. 0 0 36. 56 3.0 36. 1 2.3 37. 46 2.5 37. _1 6.8 1857 44 d = 105 .8 ( d)2 = 11,193.6 d . f . = 36 d = 2.859 d 2 = 2036.3 A r c S i n » % = x t = 2.51 I n v S i n x = A r c S i n - 137 - APPENDIX 3: Moose v e r s u s Moose F e c a l Counts P r o p , o f Tot • F e c a l Counts Prop, o f No. Old T r a n s e c t - O l d NO. New T r a n s e c t 1. 0 0 1. 0 0 2. 2 .31 2. 0 0 3. 0 0 3. 0 0 4. 0 0 4. 0 0 5. 0 0 5. 0 0 6. 1 .2 6. 0 0 7. 3 .46 7. 0 0 8. 6 .93 8. 0 0 9. 7 1.1 9. 0 0 10. 7 1.1 10. 0 0 11. 6 .93 11. 1 1.6 12. 8 1.2 12. 0 0 13. 9 1.4 13. 0 0 14. 15 2.3 14. 0 0 15. 95 14.7 15. 14 22.6 16. 52 8.0 16. 4 6.5 17. 30 4.6 17. 4 6.5 18. 5 .8 18. 2 3.2 19. 15 2.3 19. 5 8.1 20. 6 .93 20. 2 3.2 21. 14 2.2 21. 1 1.6 22. 10 1.5 22. 0 0 23. 16 2.5 23. 0 0 24. 34 5.3 24. 2 3.2 25. 13 2.0 25. 5 8.1 26. 25 3.9 26. 6 9.7 27. 26 4.0 27. 0 0 28. 32 4.9 28. 6 9.7 29. 15 2.3 29. 0 0 30. 33 5.1 30. 1 1.6 31. 8 1.2 31. 0 0 32. 8 1.2 32. 0 0 33. 3 .46 33. 2 3.2 34. 14 2.2 34. 0 0 35. 30 4.6 35. 0 0 36. 68 10.5 36. 4 6.5 37. _ l i 37. 4.8 647 62 d = 91. 4 ( d ) 2 = 8353. 96 t = 2.675 d = 2.469 d2 = 1358.3 - 138 - APPENDIX 4: P r e s e n c e o f p l a n t s p e c i e s i d e n t i f i e d i n the o r i g i n a l 10 h a b i t a t t y p e s d e s i g n a t e d - 139 - Plant species present i n one or more of the o r i g i n a l habitat types (Section 2.5) (Present = x) plant Species Grasses-Rushes-Sedges Calamagrostis rubescens Oryzopsis pungens Koeleria micrantha Carex concinnoides Stipa richardsonii Festuca saximontana Poa i n t e r i o r Agropyron trachycaulum Oryzopsis a s p e r i f o l i a Poa spp. ( j u n c i f o l i a & pratensis) Bromus anomalus Hordeum jubatum Stipa spartea Muhlenbergia richardsonis O r i g i n a l Habitat Type 3 4 5 6 7 8 x x X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X continued, Appendix 4 (cont.) Juncus b a l t i c u s Carex p r a e g r a c i l i s S p a r t i n a g r a c i l i s P u c c i n e l l i a n u t t a l l i a n a D a n t h o n i a i n t e r m e d i a Agropyron s p i c a t u m S t i p a comata S t i p a o c c i d e n t a l i s Carex p e t a s a t a C a l a m a g r o s t i s n e g l e c t a Carex a q u a t i l u s Carex r o s t a t a Beckmannia s p . Carex s i t c h e n s i s Carex a t h e r o i d e s A l o p e c u r u s a e g u a l i s Hordeum brachyantherum Carex p a c h y s t a c h y a 5 x 6 x 7 x 8 x x X X X X X X X X X X X X X X X X X c o n t i n u e d . Appendix 4 (cont.) D e s c h a n p s i a c a e s p i t o s a G l y c e r i a b o r e a l i s Carex l a s i o c a r p a Carex a u r e a Carex p a r r y a n a C a l a m a g r o s t i s i n e x p a n s a Carex c o n c i n n a E l e o c h a r i s s p . ? x X X X FORBS L i n n a e a b o r e a l i s A s t r a g a l u s m i s e r E p i l o p i u m spp. Anemone m u l t i f i d a A c h i l l e a m i l l e f o l i u m F r a g a r i a v i r g i n i a n a S o l i d a g o s p a t h u l a t a V i o l o a adunca x X X X X X X X X X X X X X X X X X X X X X X X X X X X X c o n t i n u e d , Appendix 4 (cont.) P y r o l a c h l o r a n t h a C a s t i l l e j a m i n i a t a A s t e r c i l i o l a t u s A g o s e r i s spp. S e n e c i o spp. Polemonium humile A n t e n n a r i a m y c r o p h y l l a A n t e n n a r i a n e g l e c t a Geum t r i f l o r u m Lithospermum r u d e r a l e H e i r a c i u m c y n o g l o s s y d e s Sedum l a n c e o l a t u m G e n t i a n a a m a r e l l a E r i g e r o n s p e c i o s u s G a l i u m b o r e a l e A s t e r c o n s p i c u o u s L y c h n i s drummondii A r n i c a c o r d i f o l i a 8 x x X X X X X X X X X X c o n t i n u e d . Appendix 4 (cont.) T r i g l o c h i n maritumum P o t e n t i l l a a n s e r i n a S o l i d a g o c a n a d e n s i s Taroxacum o f f i c i n a l e s A r t e m e s i a f r i c i d a P o t e n t i l l a h i p p i a n a Penstemonprocerus C e r a s t i u m arvense Heuchera c y l i n d r i c a Zigadenus venenosus Linum perenne l e w i s i i A s t e r c a m p e s t r i s Eriogonum h e a c l e o i d e s L e p i d i u m v i r g i n i c u m O r t h o c a r p u s l u t e n s V i c i a spp. L a t h y r u s o c h r o l e u c u s A l l i u m cernum 6 x x X X X X X X X X X X X X X X X X X X X X X X X X c o n t i n u e d . Appendix 4 (cont.) 1 A r a b i s spp. Ranunculus spp. A s t e r e a t o n i i Polygonum spp. H i p p u r i s v u l g a r i s P o t e n t i l l a p e n s y l v a n i c u s P o t e n t i l l a g r a c i l i s , A s t e r pansus i—• *. S m i l a c i n a s t e l l a t a Ul 1 S i s y r i n c h i u m a n g u s t i f o l i u m Comandre l i v i d a P y r o l a c e a e monensis u n i f l o r a C a r y o phylaceae A r e n a r i a l a t e r i f l o r a and C. S t e l l a r i a l o n g i p e s C i r s i u m spp. A n t e n n a r i a p u l c h r i m a T h a l i c t r u m o c c i d e n t a l e A r n i c a c h a m i s s o n i s A n t e n n a r i a p a r v i f l o r a 8 9 x X X X X X X X c o n t i n u e d . Appendix 4 (cont.) E r i g e r o n l o n c o p h y l l u s Other P l a n t s C l a d o n i a P e l t i g e r a G e n e r a l Fungus Other l i c h e n Mosses ( g e n e r a l ) Trees/Shrubs Symphoricarpus a l b u s P i n u s c o n t o r t a Pseudotsuga m e n z i e s i i Populus t r e m u l o i d e s J u n i p e r u s communis Sh e p h e r d i a c a n a d e n s i s Rosa a c i c u l a r i s S a l i x spp. A m e l a n c h i e r spp. x x X X X X X X X X X X X X X X X X X X X X X X X X X X X X c o n t i n u e d , Appendix 4 (cont.) S p i r e a b e t u l i f o l i a A r c t o s t a p h y l u s uva u r s i P i c e a g l a u c a R i b e s spp. B e t u l a g l a n d u l o s a and p u m i l a Approximate TOTAL SPECIES 6 7 8 9 10 x x X X X X X X X X X X X 26 34 31 58 28 43 44 37 52 APPENDIX 5 : N u t r i e n t a n a l y s i s on c o l l e c t e d sedge m a t e r i a l R e s u l t s o f N u t r i e n t A n a l y s i s on Sedge M a t e r i a l c o l l e c t e d March 1980.* F i b r e Dry TDN: P r o t e i n ; C a l c i u m ; Phosphorus; A.D.F. M a t t e r As As As As Sedge M a t e r i a l (% Dry) (%) Fed Dry Fed Dry Fed Dry Fed Dry 1. Ungrazed, whole sedge 45.6 96.8 50 52 3.7 3.78 0.34 0.35 0.07 0.072 p l a n t ( i . e . o l d m a t e r i a l w i t h g r e e n m a t e r i a l m i x e d ) . 2. S e l e c t e d green m a t e r i a l 41.0 96.7 55 57 6.4 6.5 0.14 0.14 0.15 0.15 from c e n t e r o f sedge p l a n t ( p r o t e c t e d by o u t e r g r o w t h ) . * S o i l , Feed, and T i s s u e - t e s t i n g L a b o r a t o r y , 1873 S p a l l Road, Kelowna, B.C. V1Y 4R2

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