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

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A HABITAT-USE AND DIETARY A N A L Y S I S 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 S a s k a t c h e w a n , B.Sc,  1976  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 I N PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES Department o f Animal  Science  We a c c e p t t h i s t h e s i s a s c o n f o r m i n g to/the requi^fe^ standard  THE UNIVERSITY OF B R I T I S H COLUMBIA O c t o b e r , 1984 ® SUSAN KAREN PRESTON, 1984  In p r e s e n t i n g  t h i s t h e s i s i n p a r t i a l f u l f i l m e n t of  requirements f o r an advanced degree a t the  the  University  o f B r i t i s h Columbia, I agree t h a t the L i b r a r y s h a l l make it  f r e e l y a v a i l a b l e f o r reference  and  study.  I further  agree t h a t p e r m i s s i o n f o r e x t e n s i v e copying of t h i s t h e s i s f o r s c h o l a r l y purposes may  be granted by the head of  ;  department o r by h i s or her  representatives.  my  It is  understood t h a t copying or p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l gain  s h a l l not be allowed without my  permission.  Department of  o4fV\\  WV  I  -^S>  The U n i v e r s i t y of B r i t i s h Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date  DE-6  (3/81)  CIA  e/wg  written  ABSTRACT Interactions  between f e r a l  studied  on f o r e s t e d  British  Columbia  objectives  f r o m May, 1979 t o A u g u s t ,  included  collecting  and c a t t l e , and  data  only,  f o r moose.  i n t h e 200 km2 s t u d y a r e a , was e s t i m a t e d  t o be 89.7 p e r c e n t  cattle,  moose.  study  The e q u i n e  harem g r o u p s , each  7.4 p e r c e n t  population  h o r s e s a n d 2.9  was o r g a n i z e d  composed o f a s i n g l e s t a l l i o n  mares and immature animals 5 t o 14.  of both  Surplus  g r o u p s o f b e t w e e n one a n d f o u r  adult  sexes,  nor considered  of  the study  on  macrovegetation  closed  forest,  five adult  i n group bachelor  animals.  seven general  was n o t p r a c The b r e a k d o w n  h a b i t a t types  was b a s e d  a s s o c i a t i o n s w h i c h seemed t o r e f l e c t  broad  and i n c l u d e d open f o r e s t , s e m i - o p e n f o r e s t ,  meadow, s h r u b c a r r , i n t e r f a c e z o n e and ' o t h e r ' 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  c o l l e c t i o n , was done o v e r t h e two summers o f f i e l d  Positive but  with  m a l e s formed f o u r  e s s e n t i a l i n t h i s study.  into  patterns  (e.g. r o a d s ) . plant  area  percent  into  ranging  A d e t a i l e d a n a l y s i s of a l l h a b i t a t types  habitat-use  Based  equal, 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  ticable  Research  s e l e c t i o n f o r horses  and d i s t r i b u t i o n  from  1980.  on h a b i t a t - u s e ,  habitat-use  size  were  information  and d i e t  approximately  a n d range c a t t l e  range o f the C h i l c o t i n Plateau i n Central  distribution  on  horses  identification  was a c h i e v e d  f o r 148 p l a n t  work.  species,  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 t h e v a r i o u s  species q u a n t i t a t i v e l y . Fecal  epidermal  a n a l y s i s was c a r r i e d o u t o n f e c a l  samples  collected and  and c a t t l e  f o r horses i n winter.  plant  species  species 86%  f o r horses  were  were used found  of the horse  herbivore plant  winter,  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 by t h e two h e r b i v o r e s .  to consitute  Horses  and w h i l e  shrub-use  from only  utilized  7.4% o f t h e a v a i l a b l e  fewer p l a n t  t h e use o f g r a s s e s  was  species  i n the  r e d u c e d , sedge and  increased.  were  used  on a r e g u l a r ,  d i s t r i b u t i o n and h a b i t a t - u s e Depending  o f b e t w e e n 18 t o 25 km i n  consecutive  data f o r horses,  on t h e i n d i v i d u a l t r a n s e c t s ,  degree o f s p a t i a l o v e r l a p and  plant  8 0 % o f t h e c a t t l e d i e t s and  A s y s t e m o f random t r a n s e c t s length  Eleven  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  d i e t s were d e r i v e d  species.  f r o m J u n e t h r o u g h September  the concentration  basis  to collect  c a t t l e and moose.  a greater  or l e s s e r  was i n d i c a t e d f o r t h e t h r e e  herbivores  o f use i n any a r e a v a r i e d w i d e l y  between  species. The transect  designated and were  habitat assumed  for  the e n t i r e study  the  habitats  horses (though also  area.  i n proportion  and c a t t l e  were measured a l o n g  to reflect  habitat  None o f t h e t h r e e to habitat  each  availability  herbivores  availability.  used Both  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  not necessarily  indicated  types  i n t h e same l o c a t i o n s ) , a n d  a preference  used open f o r e s t , c l o s e d  f o r i n t e r f a c e zone, while  cattle moose  f o r e s t and s h r u b c a r r d i s p r o p o r t i o n a t e l y  more.  iii  TABLE OF CONTENTS  Page Abstract.  i i  List of tables.  vi  L i s t of figures  v i i  Acknowledgement  ix  C h a p t e r 1:  Introduction  1  C h a p t e r 2:  M a t e r i a l s and m e t h o d s  2.1  Study area  2.2  Equine study  2.3  System o f l i n e t r a n s e c t s  2.4  B o t a n i c a l p r o f i l e of study  2.5  Establishment  2.6  Animal d i s t r i b u t i o n s  20  2.7  Fecal analysis  22  C h a p t e r 3:  6 population  9 9 area.  of h a b i t a t types  . . . . .  12 12  Results  3.1  Observations  3.2  Animal 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  Diet Estimates  43  C h a p t e r 4: 4.1  on e q u i n e p o p u l a t i o n  Discussion H a b i t a t use r e l a t e d t o  animal's  perception of a v a i l a b i l i t y 4.2  4.3 C h a p t e r 5:  24  Distribution patterns  and  60 spatial  separation  68  D i e t c h o i c e b y h o r s e s and c a t t l e  73  Summary  85 iv  TABLE OF  CONTENTS  (cont)  Page Bibliography Appendix 1  96 L i s t of plants  identified  on  the  study s i t e 2  H o r s e and  107 cattle diets  determined  by f e c a l a n a l y s i s 3  117  Paired  * t ' test for  'long-term*  versus  'short-term' f e c a l counts  on  transect V 4  Presence  134  of plant types i n o r i g i n a l  habitat types 5  Nutrient  139  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  L I S T OF  TABLES  Table 1  Page C o m p o s i t i o n and  sightings of f e r a l  horse  groups 2  10  C h a n g e s n o t e d i n harem g r o u p  no.  2, f r o m  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 May 3  2 4 , 1979  t o August  a  from  1, 1980  27  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  4  The  33  percent d i s t r i b u t i o n of h a b i t a t  types  along the f i v e t r a n s e c t s 5  38  H a b i t a t use v e r s u s p e r c e n t  habitat  availability 6  k-Independent C h i Square T e s t f o r habitat  7  39  Horse  comparing  use between s p e c i e s  41  and c a t t l e d i e t s e s t i m a t e d f r o m f e c a l  samples c o l l e c t e d i n June, J u l y , August  and  September 8  Average  44  u s e p e r month o f t h e 11  forages  c o m p o s i n g 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 9  cows  49  G r a s s s p e c i e s e a t e n by h o r s e s - s u m m e r v e r s u s horses-winter  10  R u s h and  52  s e d g e 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 11  Z-Values  for differences  use o f b o t a n i c a l groups  54 i n proportional on a s e a s o n a l b a s i s vi  58  L I S T OF FIGURES  Figure 1  Page  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 . o f  the  five transects  7  2  Young h e t e r o g e n o u s  s e r a i h a b i t a t type  3  Young homogenous s e r a i h a b i t a t t y p e  15  4  Maturing s e r a i h a b i t a t type  16  5  Maturing climax habitat type. . . .  16  6  D r y meadow h a b i t a t t y p e  18  7  Wet ( s e d g e )  8  Shrub  9  Edaphic  meadow h a b i t a t t y p e  . .  18  (meadow) c a r r h a b i t a t t y p e climax  19  (altered) h a b i t a t types  21  Logged/burned  11  I n t e r f a c e zones  12  Map o f minimum home r a n g e s  21  f e r a l horse groups, based Transects I through V distributions  19  (spruce) h a b i t a t type  10  13  15  f  f o r a l l observed on s i g h t i n g s  showing  25  frequency  of f e c a l droppings  for horses,  c a t t l e a n d moose. 14  31  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 droppings  for horses, c a t t l e  and moose o n a l o n g - t e r m b a s i s sedge p l a n t d u r i n g  35  15  Whole u n g r a z e d  16  Inner p a r t o f dormant sedge p l a n t , showing shoots  winter  d o r m a n c y . . 55 green 55  vii  L I S T OF  FIGURES  Figure 17  Page  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 m a j o r  botanical  g r o u p s 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 b a s e d fecal analysis  on 56  viii  ACKNOWLEDGEMENT T h i s s t u d y was f u n d e d b y t h e B o r e a l I n s t i t u t e f o r  Northern  S t u d i e s i n A l b e r t a , t h e B.C. F o r e s t S e r v i c e , a n d a L i f e Grant  from the U n i v e r s i t y  indebted for  t o these  of B r i t i s h  I am d e e p l y  o r g a n i z a t i o n s a n d a number o f i n d i v i d u a l s  t h e unique opportunity for f i e l d  afforded.  Columbia.  Sciences  research which t h i s  Always l o g i s t i c a l l y d i f f i c u l t ,  study  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  worthwhile I thank  experience. my s u p e r v i s o r , D r . D. M. S h a c k l e t o n ,  for  h i s perseverance  and  final  p r e p a r a t i o n o f my d a t a  In a d d i t i o n , and  and e x c e l l e n t a d v i c e  D r . R. M.  Beames, Animal  provided study the  Science  committee,  form.  D e p a r t m e n t , U.B.C.,  reviewed  a thesis  draft  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 ,  valuable assistance during  site  identifying  Provincial  able  to identify  the property  thankful  days  and sedges.  spent  o n my  Leon Pavlick of  i n V i c t o r i a , was f o r t u n a t e l y I  collected.  was p r o v i d e d b y t h e c o r r a l and c a b i n I r e n t e d o f Wayne a n d T i n a  f o r other  help  P l u m m e r , t o whom I am a l s o  at critical  provided  summer's f i e l d  S . C. C h a t w i n  three  t h e 'unknown' p l a n t s p e c i e s  Knut A t k i n s o n  first  grasses  Museum H e r b a r i u m  E s s e n t i a l support  and  a presentable  p r o v i d e d u s e f u l comments. Anna R o b e r t s  on  on t h e a n a l y s i s  D r . F. L . B u n n e l l , F o r e s t r y D e p a r t m e n t , U . B . C ,  m e m b e r s o f my e x a m i n i n g and  into  particularly  work.  f o rh i shelp  important I would  times.  Janie  Heywood  assistance during the e s p e c i a l l y l i k e t o thank  i n t h e mud-frought  frustration  of  f e r r y i n g h o r s e s , gear  and  vehicles  at times, almost i n a c c e s s i b l e study  x  site.  from Vancouver t o t h e ,  C h a p t e r 1:  Introduction  Broadly  defined,  animals,  of  resources  the  are  not  seeking  the  that  the process"  same o r  supply  i n short  "competition  of  (Birch  r  when a number  different species, which  supply,  resource  occurs  i s short;  competition  nevertheless  1957).  The  or  i f the  occurs  selection  (Diamond, 1978).  resource,  i s an  extremely  (1978),  because  by  White  animals  appear  'Interaction grazing  h a r m one  out  by  or  Schoener  1964; and  and  1978;  Arnold  of  the  and  1964),  M a i l e r , 1977;  1979;  feeding  S m i t h fit a l . ,  strategy  non-random use a l s o be  of  (1982),  oscil-  biological natural  limitation  concept  as  of  pointed  mechanisms  be  feeding  1981;  which Such  Arnold  experience  Leuthold,  out  b e h a v i o r a l , as  (Arnold,  1977),  Leuthold,  a  in and  (Arnold,  food  1977;  habits  Carpenter  1979), ' s p e c i a l i s t ' vs. 'generalist*  (Ellis  and  habitat  T r a v i s , 1975;  Nudds, 1980),  ( S a n d e r s o n , 1966).  The  or  mechanisms  p h y s i c a l , i n c l u d i n g the 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  the  Arnold  1978;  and  or  in  remains  direct interaction.  diurnal cycles Arnold,  the other  a rare  range of  avoid  a d a p t a b i l i t y ( L a y c o c k , 1978;  £t al-r  may  'Shortage' nebulous  employ to  animals  a major d r i v i n g f o r c e of  A v o i d a n c e M e c h a n i s m s ' may  times  Dudzinski,  to  and  resources  when t h e  l a t i n g between the extreme views of i t s b e i n g ( W i e n s , 1977)  common  concept of c o m p e t i t i o n  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  phenomenon  utilize  of  Dudzinski,  s p e c i e s ' morphology  M o e h l m a n , 1 9 7 4 ) , s i z e and  maintenance parameters  ( H u n g a t e §£  1946;  e c o l o g i c a l aspects  J a r m a n , 1 9 7 4 ) , and  (Leuthold,  - 1  -  a j , . , 1959;  Elsden  1977;  physical et a l . ,  of a n i m a l metabolism  ( K l e i b e r , 1961; these  latter  G e i s t , 1974;  i n the  following  tolerate  d i e t s that depart  animals,  particularly  in  quality."  profound  starvation,  on  the  the  to the  interaction  and  begins  t o be  extremely It  (1971) suiranmarized  "Small animals  can t o l e r a t e  departures  shortage  exert  relative.food ecology  of  animals,  food  shortage  b e t w e e n an a n i m a l and  between the  animal  and  ceases  to  of be  i t s environment,  ambiguous. i s important  t h a t the  two  components of  the  theory  Park  (1954)  Exploitation  refers  recognized, distinguished  as  'exploitation'  and  which  point  a  i t s associates, i s  c o m p e t i t i o n be  the  can  to the  of  to  can  from t h e optimum i n q u a n t i t y ; l a r g e  p o i n t at which  due  Bell  statement:  non-ruminants,  Although  effect  B e l l , 1971).  'interference.'  r e l a t i v e s u c c e s s o f two have  free  access  o r more i n d i v i d u a l s , o r s p e c i e s  to a l i m i t e d  is  w h e n some f o r m  of  is  the  competition  object  of  space  by  resource.  i s substituted (Miller,  Interference,  f o r a resource  1969;  Ayala,  and  1970).  I n t e r f e r e n c e i s a more s t r a i g h t f o r w a r d concept t h a n 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 b e h a v i o r  o f two  s p e c i e s of chipmunks  in  Nevada  of  Competitive  Gause  (Brown, 1971).  (1934).  (Eutamias  Brown's s t u d y s u p p o r t s t h e  Exclusion formulated Exploitation,  to t e s t because of the becomes l i m i t i n g ' ,  d o r s a l i s and g.. u m b r i n u s )  by  Grinnell  Principle  (1904)  h o w e v e r , i s much more  e l u s i v e n e s s of  f o r as W h i t e  and  difficult  ' a t what p o i n t a r e s o u r c e  (1978) p o i n t s out  'there  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 . '  Slobodkin  (1962) was  inspired  to  t h a t s p e c i e s cannot  coexist  rephrase  the  Gause P r i n c i p l e  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 - 2 -  ecological  procedure This  rather  than  a verified  or v e r i f i a b l e  i s not to say the e x p l o i t i v e  proposition.  form of competition,  which  i n c l u d e s d i e t o v e r - l a p , does n o t e x i s t , b u t r a t h e r t h a t evidence indicative and on  of i t s occurrence  should  r e s u l t s n o t be a n t i c i p a t e d . the subject  Boer  (Birch,  be c a r e f u l l y  A review  1979) s u p p o r t s  of current  must be c o n s i d e r e d  field-work  t h e c o n t e n t i o n o f Den  (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 form,  exclusion'  evaluated  'competitive  a s o n l y an e x c e p t i o n a l outcome  of t h e 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 . Historically, behind  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  t h e e c o l o g i c a l studies of A f r i c a n versus  ungulates;  t h e two t o g e t h e r  on  herbivore  in  Africa  make u p t h e b u l k  interactions.  The n u t r i t i v e  and A s i a has f o c u s e d  separation  (Leuthold,  and T a l b o t ,  North  America  around  has c e n t e r e d  the spatial  relations  premise t h a t competition (McMahan,  Willms e t a l . ,  ungulates  1969).  The work i n  t h e degree o f d i e t  among h e r b i v o r e s ,  between s p e c i e s  with  overlap  the basic  i s inherent for limited  1 9 6 4 ; H a n s e n , 1 9 7 6 ; S t o r r a r §_£. aJL. , 1 9 7 7 ;  1979).  came a b o u t b e c a u s e  1880  on  1 9 7 7 ; G r i m s d e l l and F i e l d , 1 9 7 6 ; B e l l ,  Jarman, 1974; T a l b o t  forage,  work  literature  o n s e l e c t i v i t y and e c o l o g i c a l  1971;  range  of  American  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 c o m p l e x  communities  and  North  I t i s possible that this  the western  due t o o v e r s t o c k i n g  range  lands  of domestic  latter  emphasis  were depleted of  animals  as e a r l y as  ( P a r k e r , 1 9 5 4 ) , and s e v e r a l n a t i v e u n g u l a t e s were d r a s t i c a l l y  reduced their before  i n numbers  (e.g.Cervus elaphus nelsoni)  natural predators broad  (e.g. Canis  along  with  l u p u s , Ursus a r c t o s )  long  spectrum e c o l o g i c a l s t u d i e s began. -  3  -  Recently,  however,  the  p e n d u l u m has  begun to  m u l t i - s p e c i e s r a n g e management At  this  been proven  time, to  similarities Similarity  occur in  of  but  i n regard  and  other  no  use  of  feral  herbivores  and  equids  1958  and  1959;  literature  (Koehler,  1961;  H a n s e n e t a_. , 1977)  to  competition  competition, E.  , 1 9 7 7 ) , and  W i l l m s e t a i . , 1975; •similarity,'  asinus)  historically in  recent  Hansen,  1976;  'overlap'  ' j u x t a p o s i t i o n , w i t h the concept of  Management d e c i s i o n s , i n c l u d i n g complete removal of in localized  in  United  the  Canada  forms as  of  wildlife  i t may  the  that  and  exists  they  and  occupy  1958;  the  b e t w e e n f e r a l h o r s e s and and  with  livestock.  with  However, as  emphasized t h a t r e a l i s t i c  i s very  scarce,  that other  obvious  knowledge  i n the v a r i o u s  particularly  in  management  made w i t h a d e q u a t e knowledge, and  d i e t s of f e r a l equids  both  and  assumption burros  feral  past,  Thomas, 1979)  H u d s o n , 1 9 7 8 b ) , on  o n l y be  habits  h a v e b e e n made i n t h e  (McKnight,  s e e m i t must be  d e c i s i o n s can of  States  (Salter  competition  areas,  and  competition.  1  equids  only  have been i n d i c a t e d .  been a t r e n d , b o t h  equate  have  America,  ( E q u u s c a b a l l u s and  S t o r r a r e_  of  Hudson, 1980).  i n North  diet  i n favor  n e c e s s a r i l y imply  t h e r e has  (McKnight,  exploitive  among u n g u l a t e s  does not  back  ( R i c h m o n d , 1980;  habitat-use  to  sometimes o n l y  cases  swing  on  niches  forested  ranges. The on  the  b r o a d g o a l o f t h e p r e s e n t s t u d y was i n t e r a c t i o n s between horses  regarding alces) but  not  habitat-use,  were  included  i n the  diet  and  to obtain information cattle  (Bos  taurus)  distribution  and  diet.  Moose  (Alces  i n habitat-use  and  distribution  data,  analysis.  Specific  - 4 -  o b j e c t i v e s were  as  follows: 1.  Establish  g e n e r a l h a b i t a t t y p e s and d e t e r m i n e t h e p e r c e n t  availability 2.  of each  type  in  the entire  study  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 whether  area.  available,  h o r s e s , c o w s , 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  habitat types. 3.  Determine  whether  t h e r e were d i f f e r e n c e s  i n the choice  o f h a b i t a t t y p e s among h o r s e s , cows and moose. 4.  Investigate  whether  h o r s e s , c o w s a n d moose were  distri-  b u t i n g themselves d i f f e r e n t l y i n space. 5.  Establish  what  plants  ( 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 to the animals i n the study area. 6.  D e t e r m i n e , b y 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 c o m p r i s e d t h e d i e t s o f h o r s e s and c o w s , and t o what t h e d i e t s were  7.  Determine  degree  similar.  whether  t h e same b o t a n i c a l  groups  were being  u s e d i n t h e same p r o p o r t i o n s b y h o r s e s and b y c o w s . 8.  Compare t h e d i e t s fall  9.  of horses  f o r the spring,  summer and  with their winter diets.  Determine  the broad  social  o r g a n i z a t i o n of f e r a l  horses  f o u n d i n t h e 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 of the groups.  -  5  -  C h a p t e r 2: 2.1  M a t e r i a l s and  Study Area The  study  divided  into  1,  March  1979;  1980.  began three 1 to  study  Lake  (123°  B.C.,  at km  24* an  W,  periods  as  follows:  M a r c h 10,  lodgepole  poplar,  topographical  and  volcanic traverse  occurrence  of  boulder  features  The  area  subzone  extremely tivity, the  Zone  climax  a  October  June 1 t o August the  study  S.  W.  15,  site.  L a k e and  Gay  of A l e x i s Creek,  covered  approximately  flat  with  contorta). with  gently  r e s u l t of  combined  area i s rather  forest density of  localized  l a k e s , m a r s h e s and meadows.  the  glacial  a f f e c t the  and  and  the  glacial  difficult widespread  volcanic  distribution  and  origin.  movements  stock. i n the  ( A n n a s and i n Douglas  (Douglas  to  was  I n t e r i o r Douglas F i r  (IDF)  C o u p d , 1979), i n c l u d i n g a l l f i r on  mesic  sites.  f i r - p i n e g r a s s , northern  The  phase)  IDF  is  an  i m p o r t a n t f o r e s t e d subzone i n terms of range produc-  and  provides  c e n t r a l B.C.  a r e a was  the  is classified  Biogeoclimatic that  are  Much o f  domestic  km  (Pinus  fields  also  23  work  a f o r e s t o f D o u g l a s f i r (Pseudotsuoa  features  of  May  i s generally  interspersed with  because  o f w i l d l i f e and  areas  area  field  between Haines  m a s l , and  pine  activity.  and  N) , 24.3  The  the  s p e n t on  located 57'  1).  The  These  51°  and  1980;  m o n t h s was  e l e v a t i o n o f 1050  and  s p r u c e and  in  1979  t e r r a i n , c o v e r e d by  menziesii)  (b)  of  a r e a was  (Figure  2  rolling  to  i n May  A t o t a l o f 6 1/2 The  200  Methods  located  the m a j o r i t y region  i n the  o f summer g r a z i n g  (Annas and  C o u p e , 1979).  more w e s t e r l y -  6  -  portions  for The  of the  cattle study IDF(b)  Figure  1:  Map  of  study  area,  showing  arrangement  o f t h e f i v e t r a n s e c t s , r a n g i n g from 18 t o 25 km, from the  center  of  (discussed i n sec.  the  study s i t e ,  2.3).  - 7  -  "Wilson's  radiating Meadow"  subzone. The  study area included  a l l o t m e n t s where c a t t l e  p a r t s of t h r e e d i f f e r e n t  (Herefords predominating)  range  ranged  from  June through October with a peak g r a z i n g d e n s i t y i n July-August. A p o p u l a t i o n of f e r a l  h o r s e s a l s o used the study area d i v i d e d  f o r the most p a r t , i n t o d i s t i n c t s o c i a l 2.2  Equine Study P o p u l a t i o n A fluctuating  area  number o f domestic horses shared the study  w i t h 65 h o r s e s  identified  as f e r a l .  were o r g a n i z e d i n t o  at l e a s t  individual  w i t h o u t a harem.  stallion  composition of the groups 2.3  groups.  eight  The  feral  horses  d i s t i n c t groups, p l u s one Table 1 shows the b a s i c  identified.  System of L i n e T r a n s e c t s Although  aerial  photographs  were used  d u r i n g the study  to e s t a b l i s h t r a n s e c t s , and t o supply more s p e c i f i c i n f o r m a t i o n of l a n d f e a t u r e s than i s found on t o p o g r a p h i c a l maps, an o v e r a l l photo of The  interpretation  each  habitat  type  of the area to determine available  r e g i o n i s composed  proved  of h a b i t a t s  t o be  the p e r c e n t impractical.  which a r e i n t o o  a mosaic t o 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 b l a c k and white graphs.  Such h a b i t a t  t y p e s as  'Interface  Zones' and  tight photothose  based on the p r e c i s e s e p a r a t i o n of f o r e s t d e n s i t y , are d i f f i c u l t to  r e c o g n i z e , a l t h o u g h t h e s e a r e a s p r o b a b l y 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 p r o b a b l y be f e a s i b l e by a e r i a l photograph i f c o l o r e d photographs  interpretation  were a v a i l a b l e f o r the r e g i o n . -  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 H o r s e G r o u p s  Locations  Stallion Groups  Mares  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  Sightings  Bachelor  F o a l s , up t o two y e a r s  Total*  Groups  4  4  6  3  2  2  7  2  5  5  8  4  6  5  9  1  •including  stallion  * * g r o u p most c l o s e l y o b s e r v e d . 15 m o n t h s  A l l group changes noted  (Table 3 ) .  - 10 -  over  A method  of  random  transect  s t u d y w h i c h more a c c u r a t e l y a v a i l a b l e t o animals. of  a modified  all  The b a s i c method i n v o l v e d  transect  The  cabin  final  was  i n the  t h e development  system i n the form o f f i v e  located  layout  used  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  o r i g i n a t i n g and r e t u r n i n g  study  a n a l y s i s was  at  'circuits,  1  t o t h e c e n t r a l a r e a where t h e 'Wilson's  Meadow'  (Figure 1).  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 b y  practical  considerations,  i n c l u d i n g where s a d d l e h o r s e s  could  maneuver  and  w h i c h c o u l d be c o v e r e d o n a d a i l y and  seasonal  the distance  basis.  The  length  (Figure  on  final  be known  collected. along  ranged  1 ) , and were d i v i d e d  into  18 t o 25 km i n  0.5-km i n t e r v a l s  accumulated was  recorded,  was  transects.  enabling  production  and a s a c c u r a t e l y a s p o s s i b l e .  removed t o a v o i d This  s u c h as f e c a l  two summers' movements  encouraging  field  or hoofprint work, that  of animals  Each t r a n s e c t the order  survey  Minimal  animals  t o use  i s an o b v i o u s d a n g e r when u s i n g a r e p e a t i n g  was  the transects  i n the area.  ridden  of r i d i n g  There was no i n d i c a t i o n  concentrations)  o f a n y t r a n s e c t was a d o p t e d a s a r e g u l a r  (i.e.  the  type  c l e a r e d of overhanging branches t o f a c i l i t a t e  t r a n s e c t s y s t e m i n h e a v i l y wooded t e r r a i n . (e.g.  of each h a b i t a t  I n a d d i t i o n , e a c h t r a n s e c t was r i b b o n e d w i t h  them repeatedly  vegetation  distance  being  p i c t u r e o f t h e h a b i t a t t y p e s a v a i l a b l e on e a c h  and l i g h t l y  riding  the  from  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  transect  a composite  tape  with  The  each  transect.  the  transects  t h e g r o u n d t o e n s u r e t h a t t h e r i d e r ' s p o s i t i o n on t h e t r a n s e c t  would  of  five  during the  unduly  affected  C e r t a i n l y no segment travel  route.  on a r e g u l a r , c o n s e c u t i v e  the transects - 11 -  was  not  basis  changed),  during  which  a l l fresh  3 m of either side sheets. signs can  horse,  of the saddlehorse  a  i n the form  be  quickly  of feces,  and e f f i c i e n t l y  problem  i n studies  e l k a n d moose  removed  and  September, 1979; and t h r e e  1200 km.  a  to carry  clear  quently, plant  three times  distance  Each  times each each  such  ridden  were  transect  i n August  i n J u n e and J u l y ,  f o r a l l transects  idea  t o e s t a b l i s h some s y s t e m o f h a b i t a t  was  species  classification  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 of plant  species  present  t o have  i n the area.  Subse-  a b o t a n i c a l s u r v e y o f a l l p l a n t s o b s e r v e d , and c o n c o m i t a n t  collection,  work.  no  a s h a s b e e n shown t o be  B o t a n i c a l P r o f i l e of Study Area. In order  and  and  Saddlehorse defecations  o f 12 t i m e s ,  nearly  are large  from horseback i n  when t h e y o c c u r r e d .  ridden  2.4  feces  1968).  was  The a c c u m u l a t e d  because  L i t t l e d i f f i c u l t y was e n c o u n t e r e d  from the t r a n s e c t s  1980.  species  o f more c l o s e l y r e l a t e d s p e c i e s  (Neff,  a total  recorded  within  r e c o r d e d on d a t a  b e d s , and t r a c k s  i d e n t i f y i n g the respective  as  were  T h i s method w o r k e d w e l l f o r t h e s e t h r e e  the d i f f e r e n t h a b i t a t types. in  cow, a n d moose s i g n s  Positive (Appendix  w e r e made o v e r  identification 1).  t h e two summers o f  was a c h i e v e d  Due t o t i m e  a t t e m p t was made t o d e t e r m i n e  f o r 148  field plant  a n d manpower c o n s t r a i n t s ,  availability  of the various  plant species q u a n t i t a t i v e l y . 2.5  Establishment The  of Habitat  breakdown  Types  of the study - 12 -  area  into general  habitat  types  was  there to  based  i s no  animals,  was  types  used  macro-vegetation  doubt  a n a l y s i s of  I n some c a s e s ,  the  b a s i s of  habitat  d e s i g n a t i o n was  such  the  as  spacing  used  on  wooded  terms of  were  m o r e on  not  large variations in this  In other  variation instances,  spatial differences, of  lodgepole.  (R.A.B.) s u c c e s s i o n a l s t a g e  terminology  Habitat  relative  important  a l l h a b i t a t t y p e s was  separation.  based  Although  often  Consequently,  i n d e f i n i n g h a b i t a t types sites.  are  between the t r e e s i n stands  Resource A n a l y s i s Branch was  there  between h a b i t a t s .  as  associations.  that micro-habitats  a detailed  practicable. plant  on  ( W a l m s l e y et. a _ . ,  d e s i g n a t i o n was  d e n s i t y from v i s u a l  1980)  also described  e s t i m a t i o n s of  in  foliage  cover. Initially, transects;  10  habitat  types  a model h a b i t a t s i t e  were d e s c r i b e d  i n each case being  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 .  to  n o n - c l i m a x s p e c i e s w h i c h h a v e n o t y e t gone t h r o u g h  of  n a t u r a l t h i n n i n g s , and  old.  The  little 60  to  140  1.  years  "Maturing  o f age  and  has  of  even h e i g h t  serai"  Young  a series 60  years  there i s  gone t h r o u g h  the f i r s t  thinning  some r e g e n e r a t i o n .  were:  heterogenous  typical  and  refers  i s g e n e r a l l y between  a c q u i r i n g c o m m u n i t y s t r u c t u r e and  h a b i t a t types  than  the  described  "Young s e r a i "  generally less  is essentially  regeneration.  stages, The  canopy  are  along  young  serai  on  south-facing  of  some  Douglas  juxtaposition  serai:  Essentially  features  slopes f i r .  except  subscribing  t e n d i n g t o be  w h i c h seems t o f a v o r t h e These  to maturing - 13  areas  climax. -  a l s o tend Density  to  found  presence t o be  in  estimate  is  semi-open 2.  Young  (Figure 2).  homogenous s e r a i :  More s t r i c t l y  typical  young s e r a i  form  cover.  Density estimate  with  little  conforming  to  t o moderate ground  i s d e n s e t o v e r y dense  (Figure  3). 3.  Maturing  serai:  There  fallen  t r e e s tend  fairly  uniform  to  4.  Maturing  climax:  d o m i n a n t and 'veteran'  litter  serai  e s t i m a t e i s semi-open  a.  forest  floor.  may  g r a s s and  represented.  Regeneration  Wet  with Douglas f i r A few  o f b o t h f i r and  wet  fire-scarred lodgepole.  (Figure 5).  be  low  p o i n t s i n l a r g e open a r e a s ) .  Forb,  some s e d g e r e p r e s e n t a t i o n .  Dry,  open  hillsides  overgrazing  shrub  tending  cover.  t o be  slightly  ( s e d g e ) meadow:  grazing years.  dry  more areas,  Both types are susceptible  (Figure 6). Low  some t e n d i n g t o s t a y wet to  Density  sub-types.  w i t h more t r e e and  6.  Usually  ( i . e . lodgepole).  p r o d u c t i v e i n v a r i e t y of s p e c i e s than the f l a t ,  to  and  (Figure 4).  lodgepole well  Two  'veterans',  Open meadow - o f t e n i n j u x t a p o s i t i o n t o s e d g e meadows  (which  b.  no  Community s t r u c t u r e  firs.  D r y Meadow:  t o be  the  species  D e s i t y e s t i m a t e i s open 5.  tend  areas  year  pressure.  of water  round.  accumulation,  Reasonably  Sometimes i n a c c e s s i b l e  Relatively  low  - 14  -  resilient in  plant species variety  very and  F i g u r e 2:  Young heterogenous s e r a i h a b i t a t type  little  t r e e or shrub growth  (Figure 7 ) . t  7.  Shrub  (meadow) c a r r :  thaw u n t i l season and  June, w i t h moist s o i l  i n an a v e r a g e  sedges.  drier  Edaphic  Bog b i r c h  i s open, climax  (Heyes,  erratically  or  consistently  shrub  carr  cattle  with  (Picea glauca). shrub  Tending  cover  areas  with  moose.  f o r i t t o accumulate with  with  Density  (Figure 8 ) . either  (ravines);  w e t meadows  P r o v i d e favored bedding  areas c o i n c i d e n t a l  grasses  t o be f o u n d  low l y i n g  i n juxtaposition  and p a r t i c u l a r l y  tendency  dense  interspersed  areas.  1979) w i t h  (Betula spp.) is, dominant,  (spruce):  in  of  year  f o r m a j o r i t y o f summer-fall  'pockets' o f w h i t e spruce  estimate 8.  A r e a s f r e q u e n t l y f l o o d e d from s p r i n g -  and  sites for  F r e e wa'ter a b s e n t , b u t i n d e p r e s s i o n s and edges  shrub margins  - areas  hence  i  always  damp and c o o l  - reflected mosses).  ( p r o l i f i c mosquito-breeding  i n hydrophilic  plant  species  grounds)  ( e . g . Sphagnum  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  (Figure  9). 9.  Logged/burned at various (i.e.  10.  disjunct.  regenerative  l o g s ) c a u s i n g h e r b l a y e r t o be c o n s i s -  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  (Figure 10).  Interface of  T e n d i n g t o be  a g e s ; 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  l i m b s , stumps,  tently open  (altered) s i t e s :  Zones:  quite  ' 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  Spruce-Aspen-Lodgepole  of varying  ages. i  - 17 -  zones  Tending t o  F i g u r e 7:  Wet meadow h a b i t a t - 18 -  type  F i g u r e 9:  Edaphic  climax - 19  (spruce) h a b i t a t -  type  be  vegetatively  (and o f t e n  physically)  complex - h a v i n g  high v e g e t a t i v e d i v e r s i t y i n keeping w i t h Ecotone The In tions  d e n s i t y e s t i m a t e i s dense t o semi-open the  final  were  habitat  (Figure  r e d u c e d t o s e v e n c a t e g o r i e s w h i c h seemed t o  Open F o r e s t :  11).  a n a l y s i s , the ten o r i g i n a l designa-  broad h a b i t a t - u s e p a t t e r n s . 1.  biology.  reflect  They were:  c o m b i n i n g m a t u r i n g c l i m a x and  logged/burned  sites. 2.  Semi-open F o r e s t : genous young  combining  maturing  s e r a i and h e t e r o -  serai.  3.  Closed Forest:  4.  Meadow:  homogenous y o u n g  combining  wet  and  dry  serai. (spatially  difficult  to  separate). 5.  Shrub  Carr.  6.  Interface  Zone:  combining  edaphic  c l i m a x and  interface  zones. 7.  'Other :  2.6  Animal  1  To any  point  Distributions  determine  level  presence  roads, e t c .  whether  of a s s o c i a t i o n or absence,  along a l l five  or not  the  t h r e e s p e c i e s showed  among e a c h o t h e r , I c o n s i d e r e d t h e i r  relative  to each  transects. - 20  o t h e r , at each  S p e c i e s were t h u s -  .5  km  tabulated  Figure 10:  Logged/burned  (altered) habitat  type  as  either  alone,  none p r e s e n t . C,  was  Fecal  The  a contingency  and  of  of  f o u n d , was  for  the  m a t e r i a l , each  two  one  a l w a y s on  collecting  I t was  13  fecal  was  the  c o l l e c t i o n s of  possible  due  the to  vegetative  immersed  i n 90%  the  of  were e v e n l y  then  were c o l l e c t e d work.  consisted  t a k e n from e i g h t same s i t e .  The  experience  only  of  One eight  individual, opportunistic wherever encountering  that eight  individual  space o f 1 t o 2 days.  m a t e r i a l f o r both species  from  June  e f f e c t of  through  c o l l e c t i o n and  stored  The  of  in air-tight  t o t a l monthly  respect  to  to  plant  fecal material, jars,  a l l pertinent information  d i s t r i b u t e d with  were  September,  concentrations  'growth s p u r t s . '  a l c o h o l , was  Between  and  recorded.  collections,  time.  f e c a l s a m p l e s were s e n t t o t h e Washington S t a t e U n i v e r s i t y  Wildlife the  was  Test.  summers' f i e l d  Samples f o r a n a l y s i s , chosen from the  of  (Smith,  coefficient  always  the  fecal  ameliorate  The  or  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  e a c h month  date  -1.0  m a t e r i a l , w h e n e v e r and  d e t e r m i n e d by  made t h r o u g h o u t  types  to  adopted because o f the u n p r e d i c t a b i l i t y of  piles  and  each  d i e t estimates  found f o r both s p e c i e s w i t h i n the 7  f r o m +1.0  f o r each species  f r e s h dung p i l e s , n o t  fecal  vary  Chi Square  cows d u r i n g  sample  teaspoons  horses.  a l l together,  a c o e f f i c i e n t of a s s o c i a t i o n ,  s i g n i f i c a n c e of  samples  horses  approach  can  species,  Analysis  Fecal  month's  other  From t h e s e v a l u e s  702).  d e t e r m i n e d by  for  one  c a l c u l a t e d which  1980;  2.7  with  Habitat  composite  Lab  for analysis.  samples  of  eight  - 22  -  They c o n s i s t e d  of  four  i n d i v i d u a l s each, f o r both  h o r s e s and c o w s , f o r J u n e , J u l y , A u g u s t for  example,  II).  In  were  f o r cows i n June  addition,  spent  at  the  of  These for a  one  diet  The  single  the  referred  from  were  each  examined  consisting as  each  sample  t o as  96  an  tests  top of the the  study  i n d i v i d u a l dung p i l e s .  undetermined  two  samples  time  f o r horses w i l l  '^prsersLura^er'  period.  be  collec-  to distinguish  i t from  analysis.  material  points'),  Due  to  or  d i e t was of  800  economic  were examined,  that  could  slides  be i d e n t i f i e d ,  level  of  5%  a  presented i n Chapter  200  views  per  determined from four diet views  each,  f o r both  r e s t r a i n t s , o n l y two  diet.  samples,  horses winter  and diet  analysis  views.  probability  of hypotheses.  t o t a l of  susequently the w i n t e r d i e t  i s b a s e d on a t o t a l o f 400 A  only.  Thus t h e w i n t e r d i e t s r e p r e s e n t  over  for plant  month c o n s i s t s  samples  on  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  each monthly  cattle.  frozen  days  o f 25 f i e l d s o f v i e w p e r 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  'cover  Because  or  feces  w e r e a l l c o m p o s i t e d and  taken out.  'Horse-winter' d i e t For  are  fresh  spring/summer/fall diet analysis  tively  to  each  samples  analysis  larger  either  (Appendix  1979-80, t e n  random c o l l e c t i o n t h r o u g h o u t  teaspoon  individual  J-C2# e t c .  study s i t e c o l l e c t i n g horse  representing  site  J-C^;  d u r i n g the w i n t e r of  Samples were c o l l e c t e d snow,  as  and S e p t e m b e r - d e n o t e d ,  Specific  was  details  3.  - 23  chosen  -  a priori  fora l l  o f t h e a n a l y s e s used  Chapter 3.1  3:  RESULTS  O b s e r v a t i o n s on Equine P o p u l a t i o n For  and  a total  groups,  considered 4 percent towards and  o f 90 h o r s e  f o r a l l time feral,  sightings,  spent  28 p e r c e n t  unknown.  i n the f i e l d , domestic,  Not s u r p r i s i n g l y ,  meadows, w i t h  including  86 p e r c e n t  individuals  65 p e r c e n t  3 percent sightings  of horses  mixed and  were  found  biased  i n them,  8 p e r c e n t i n i n t e r f a c e zones, 3 p e r c e n t i n semi-open f o r e s t ,  2 percent  i n closed forest,  i n open f o r e s t and shrub The high  home  priority  range  1 percent  o b s e r v a t i o n s were  opportunistic,  on  number  o f minimum  of sightings  home  ranges  i s r a t h e r tenuous  H o w e v e r , 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 areas horse  of use, relative groups  to the entire  Harem  group  #2, b a s e d  the  coincidence of i tbeing In addition,  G r o u p #2 w e r e a l s o r e c o r d e d Animal In  at best.  approximate  located i n 'group  range  given i s  perhaps  centrally  due t o  i n the study  composition  o f Harem  (Table 2 ) .  Distributions  addition  percentages  o n 29 s i g h t i n g s ,  changes  based  study area, f o rthe e i g h t  ( F i g u r e 1 2 ) . The most a c c u r a t e  for  area.  as a  was n o t p l a c e d o n o b t a i n i n g b e h a v i o r a l o r g r o u p and t h e mapping  a limited  i n o t h e r , and 0 p e r c e n t  carr.  information,  3.2  were  t o p r o v i d i n g an e s t i m a t e  o f each  habitat  from t h e t r a n s e c t s f a l l  o f t h e t y p e s and  type a v a i l a b l e , the data  collected  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 1 2 : Map o f minimum home r a n g e s f o r a l l b a s e d on s i g h t i n g s .  - 25 -  observed f e r a l horse groups,  - 26 -  TABLE 2: C h a n g e s n o t e d i n Harem G r o u p No. 2, f r o m a t o t a l o f 29 s i g h t i n g s in  the period  f r o m May  2 4 , 1979 t o  a d d i t i o n s and l o s s e s t o t h e band.  - 27 -  August  1, 1 9 8 0 , s h o w i n g  Date of Siqhtinq May  24,  Oriqinal  1979  Composition  Chestnut S t a l l i o n , 13 y r s .  Additions  Losses  about  Heavy b l a c k m a r e , a b o u t 12 y r s . - w e a r i n g o l d leather 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 m a r e s , a b o u t 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 J u n e 14,  1979  Br. male f o a l b o r n t o one o f brown m a r e s .  J u n e 24,  1979  Br. female f o a l born to o t h e r b r . mare.  Aug.  10,  1979  M a r c h 7,  1980  B r . m a l e 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 hours. Later found r e u n i t e d . Red m a l e f o a l b o r n t o b l a c k mare a f t e r m i d A u g u s t , 1979.  1 b r o w n mare & male f o a l o f p r e v i o u s June continued...  Table 2 continued... Date o f Sighting  Original  Composition  Losses  New c h e s t n u t m a r e .  M a r c h 7, 1980 (cont.)  New b a y m a r e . F i n a l composition.  J u n e 3, 1980  Chestnut J u l y 3 1 , 1980  Additions  Stallion.  B l . mare & r e d f o a l o f Aug. 1 9 7 9 . 1 b r . mare w i t h new f o a l and y e a r l i n g f e m a l e o f J u n e , 1979. 1 (2-3 y r . o l d ) b r . mare w i t h new f o a l . 1 r o a n mare.  Br. f o a l born t o b r . y e a r l i n g o f 1979.  Parti-colored male.  New Roan mare.  2 New m a r e s i n March.  Br. f o a l born t o remaining o r i g i n a l b r . mare - w i t h y e a r l i n g female.  seen  1.  R e l a t i v e use and  2.  o f each h a b i t a t type  horses  moose ( s e c t i o n 3 . 3 ) , and  Distribution  by  each  animal  the e n t i r e study area The  by c a t t l e ,  latter  was  species  i n relation  (i.e.a l l habitats available).  achieved  by  graphing  the percent  fecal every  material  i n relation  to the distance along  the transects  0.5  km,  to give  idea  separation  the  species.  levels use  This  of the s p a t i a l  approach  of i n t e r a c t i o n .  meadows  they  an  to  i s important  Although  as a h a b i t a t type  cows  d i d n o t u s e t h e same i n d i v i d u a l  o f i n t e r a c t i o n w o u l d be m a r k e d l y  i n clarifying  and  i n excess  between  horses  the  both  may  of a v a i l a b i l i t y , i f  meadows, t h e n  the  levels  reduced.  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 of  of s p a t i a l  use  i n any  o v e r l a p was  area  The d i s t r i b u t i o n  indicated  v a r i e d widely  and t h e c o n c e n t r a t i o n  between t h e t h r e e  species.  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  frequencies  alone, with  one  were p r e s e n t , is  shown  other  with  which  the three  seen  s p e c i e s , o r a l l t o g e t h e r , and when none  a t e a c h 0.5 km i n t e r v a l  i n Table  s p e c i e s were  3.  along  the f i v e  transects  C a l c u l a t i o n s of the c o e f f i c i e n t s  of  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 , together 1)  with  horses  expected  and cows were by c h a n c e  moose w e r e chance  the C h i Square  found  alone  h c  together n m  showed  the  a s s o c i a t e d w i t h each other  (C =0.07;  (C =0.03;  values  following: more  X = 4 . 4 , d f = 1 ) ; 2) h o r s e s 2  as o f t e n  X =0.33, df 2  - 30 -  as  one  =1);  would and  expect  3) m o o s e  than and by and  FIGURE 1 3 : Transects  I through  V,  showing  frequency  distributions  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 a n d moose a t 1 Km along  each  transect  f o r the June,  period.  - 31 -  July,  August  of  intervals  and September  a a  ""  . s i  __ •____5___ ^SI5____  > i-  -a  -a 9  I-a  55555  f -  - 32 -  8  9  a  Table  3:  Coefficient moose  o f A s s o c i a t i o n f o r horses.  Frequencies  with  which  M = moose; H = h o r s e s ) all on  together, a l l five  obtained number  each  species  C a t t l e and  (C =  cows;  were s e e n a l o n e , two t o g e t h e r ,  or none,  a t each  0.5  t r a n s e c t s combined.  km  interval  Frequencies  were  on a p r e s e n c e o r a b s e n c e b a s i s o n l y ; a c t u a l of fecal  piles  p e r 0.5  km  interval  were  ignored.  Total # 0.5 km intervals 201  C  H  M  HC  HM  MC  HCM  None  55  4  14  64  8  17  33  6  - 33 -  cows a r e f o u n d 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 X =17.81,  I  estimate  used  the  to  the relative  t h e number  16  and  abundance o f t h e t h r e e  of i n d i v i d u a l  transects with  cattle,  moose  fecal  the available  data  24  hour  recorded  that indicate  period  a l . , 1 9 7 6 ; M i q u e l l e , 1983; cows: From  piles  species along  horses,  a l l have an a v e r a g e d e f e c a t i o n r a t e o f  e l i m i n a t i o n s per  obs.).  m c  df = 1).  2  To  (C =-0.51;  these  data  (moose:  Hafez,  the r e l a t i v e  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  15  F r a n z m a n §_£  1969; h o r s e s :  pers.  p r o p o r t i o n s were  and 2.9 p e r c e n t moose  89.7 (Table  8). In the  order  three  Transect  species with (V) was  indications old,  t o compare  were  Transect  as  a  'control.'  However,  investigated  use o f t h e a r e a f o r  collected  use, estimated  recorded. V was  'long-term*  the data  used  of animal  the  i n this Normally  t o be l e s s  before  data  in detail.  study, only  than  total  fresh  s i x days  collection The  began, 18.5  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 signs  recorded  material  could  When t h e s e the  within  graph  represent  'long-term* f o r 'fresh*  on T-V, t h e two g r a p h s similar Rohlf, was  ( F i g u r e s 13 1969:  r u n on  'fresh' graphs  4 m each  side of the center  at least  data  were  years  graphed  material collected  km  animal  line.  The  accumulation.  and compared over  two  with  summers  ( i g n o r i n g s c a l e ) a p p e a r e d t o be e x t r e m e l y and  14).  A  205), with Arcsin the three  f o r each  five  one  being  ' t 'test  Transformation  sets of data,  of the three  t o be d i f f e r e n t ,  Paired  just  - 34 -  (Appendix I I I )  i . e . comparing  species. into  ( S o k a l and  ' o l d ' and  R e s u l t s showed t h e  the s i g n i f i c a n t  range  FIGURE 14: Transect  V,  showing  frequency  distributions  f e c a l droppings  f o r horses, c a t t l e  data c o l l e c t i o n  f o r the study.  -  35  -  of  and moose p r i o r  accumulated to regular  -  Percent  9e  -  Frequency  of KJ  o  i  ////// yrvr// // // // // // o  3 —tfD —} in ro  NJ  UJ  // // // // JZ\ ////// SSSS3  A  Y // // // /T7m In  00  65 KJ.  O  V // // // //H  BHSlfe.7/  / / / / / / //]  Observation (jn L_  CO  O i  CO  cn i  for a l l three horses  vs.  sets of data:  horses  (t =  cows v s . cows  2.51)  and  ( t = 2.35,  36  moose v s . moose  d.f.),  (t =  2.68)  (Appendix I I I ) . 3.3  Habitat The  based  Use  percent  on  the  T a b l e 4.  total  (X  in  to  regard  contained  the  was  type  for  along  each  a l l transects,  = 24)  transect,  i s shown  e x i s t s among t h e  r e l a t i v e amount  of  each  transects  habitat  type  plant  species  i n dry  found i n each h a b i t a t The  meadow h a b i t a t ,  and  the  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 m e a s u r e d on e a c h d i r e c t measurement  transect, the  an  estimate  the of  entire  habitat  4  habitats  and in  horses and  zone,  while  ).  None  proportion and  more,  5  to  cattle also moose  between  6) , i n d i c a t i n g s p e c i e s  three  open  availability  These  species  - 37  -  used  the  (Table  5).  for  forest  differences  interface and in  were s i g n i f i c a n t  s p e c i f i c selection for  habitats.  transects  disproportionately  preference  forest, closed  more.  three  a  of  availability  herbivores  meadow h a b i t a t  indicated  used  the  the  habitat  c a t t l e used  disproportionately  selection  of  transect  length  e n t i r e s t u d y a r e a when c o m b i n e d f o r a l l f i v e  (Tables  Both  provides  along  species  minimum number  f o u n d i n y o u n g homogenous s e r a i h a b i t a t .  by  they  varied  maximum number o f  was  each  in  4).  between h a b i t a t s . ,  found  obtained  carr  distance  numbers o f  considerably  for  habitat  = 18,020.7, d . f .  2  (Table  The  (26)  each  A k - I n d e p e n d e n t C h i S q u a r e T e s t showed t h a t a s i g n i f i c a n t  difference  (58)  of  the  shrub habitat (Table  available  TABLE 4:  The five  percent*  transects.  meters each  distribution Numbers  of transect  of habitat  type  i n parentheses  covered  by each  along the are actual  habitat  type,  on  transect.  SemiOpen F  Closed F  Meadow  Interface zone  Transect  Open F**  I  2.3 (2280)  8.9 (8880)  3.1 (3060)  5.7 (5730)  0.0 (0.0)  0.2 (150)  0.2 (240)  II  0.7 (720)  8.0 (7980)  3.9 (3870)  3.9 (3930)  0.0 (0.0)  1.0 (1020)  0.0 (0.0)  III  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 transects  **F = Forest  - 38 -  Shrub carr  ( i . e . 99,690m)  Other  TABLE 5: Habitat  use  versus percent  1.  T o t a l f e c a l c o u n t s per for a l l transects.  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 availability f o r a l l t r a n s e c t s f o r t h e 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 based for a l l three species.  Habitat  habitat  Habitat A v a i l a b i l i t v type  f o r horses,  on e q u a l  average  cows and moose  defecation rates  SemiInterShrub Open Open C l o s e d face F F zone O t h e r Type F Meadow c a r r  % Available Total  14.5 28  Relative Population Estimates 6 d.f. X  33.4  13.7  28.1  2.6  5.7  1.9  100%  100  32  331  7  15  2  515  2  336.7*  Horses % Total  5.4 19.4 289  551  6.4  64.2  1.4  2.9  0.4  96  4739  103  392  46  Cows  6216 7402.8*  %  4.6  8.9  1.5  76.2  1.7  6.3  7.4%  89.7%  0.7 continued  Table 5  (cont.)  SemiOpen Open C l o s e d H a b i t a t Type F F F %  Available Total  Shrub Meadow c a r r  Interface zone Other  X2  14.5 33.4  13.7  28.1  2.6  5.7  1.9  100%  37  84  31  13  4  0.0  200  31  Moose  6 d.f.  161.9*  %  15.5 18.5  42.0  15.5  6.5  2  0.0  Total fecal  count:  6931  Relative Population Estimates  2.9%  TABLE 6  ^^\k=7  k - I n d e p e n d e n t C h i S q u a r e T e s t f o r c o m p a r i n g H a b i t a t Use B e t w e e n S p e c i e s , showing 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 three species.  H a b i t a t Open F. S-Open F. C l . F. Meadow S h r u b C. I . Z . O t h e r F e c a l C o u n t 14.5 33.4 % 13.7 28.1 1.9 2.6 5.7 Totals X Cow  Horse  Total: Cow  289  551  96  4739  103  392  46  4.6%  8.9%  1.5%  76.2%  1.7%  6.3%  .7%  28  100  32  331  7  15  2  5.4%  19.4%  6.4%  64.2%  1.4%  2.9%  .4%  317  651  128  5070  110  407  48  6731  289  551  96  4739  103  392  46  6216  6216  2  d.f.=6 129.0*  515  1327.71* Moose  Total:  31  37  84  31  13  4  15.5%  18.5%  42.0%  15.5%  6.5%  2%  320  588  180  4770  116  396  -  200  46  6416  continued...  Table 6 (cont) H a b i t a t Open F. S-Open F. C l . F. Meadow S h r u b C. I . Z . O t h e r F e c a l C o u n t % 14.5 33.4 13.7 28.1 2.6 5.7 1.9 Totals X 2 Horse  28  100  32  331  7  15  2  d . f . = 6  515 216.1*  Moose  31  37  84  31  13  4  -  200  Total:  59  137  116  362  20  19  2  715  3.4  Diet Analysis From  the four  composite  fecal  samples  collected  each  month f o r b o t h cows and h o r s e s - s u m m e r , and f r o m t h e two s a m p l e s for  horses-winter,  presence (Table that  average  7).  The  at least  results  54  (Table 7 ) .  o f 80 p e r c e n t  found  to constitute  horses  were  indicates diets thus plant 11  diets  were  species.  species  which  they  among m o n t h s  80%  (Table  a  diet,  on t h e s t u d y  o f 7.4  percent  relatively  I t should  chosen  minimum  and  were  differed  I ) , this  herbivores* species,  of s e l e c t i v i t y f o r w e r e t h e same  the proportions i n  b e t w e e n t h e two h e r b i v o r e s  showed  of  use) by  month  and  (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 8)  of  148 p l a n t  (Appendix  t h a t these  h e r b i v o r e s , although  were  86 p e r c e n t  o f t h e two  degree  be n o t e d  show  by t h e  of the a v a i l a b l e  high  month  used  of a t l e a s t  f o r both used  each  11 p l a n t s p e c i e s  area  percent  f o r the  analyses  were  an a r b i t r a r i l y  As a t o t a l  80  fecal  species  of the c a t t l e ' s  8).  at least  composed  indicating  Using  of the t o t a l  identified  that  plant  determined  i n the diet  of the monthly  different  value  species  were  of the various plant species  two h e r b i v o r e s  the  values  that both  t o month  rubescens.  Poa  neglecta,  Stipa  the average horses  consumption  and  cattle  f o r the f o l l o w i n g iuncifolia,  Stipa  comata, and  was f o u n d t h a t h o r s e s a l w a y s  ( i . e .concentration  varied  significantly  s i x forages: occidentalis,  Hordeum  (Table  iubatum.  u s e d C_. r u b e s c e n s  from  Calamaqrostis Calamaqrostis Further, i t  more t h a n  cattle,  b u t i t s a v e r a g e u s e by h o r s e s p e a k e d i n A u g u s t a n d t h e n d e c l i n e d , - 43 -  7:  TABLE  H o r s e and C a t t l e D i e t s e s t i m a t e d f r o m f e c a l s a m p l e s in  June,  July,  August  and  September.  b a s e d on p e r c e n t p r e s e n c e i n f e c e s :  Plant  -  species  use  averages from 4 composite  f e c a l samples p e r a n i m a l s p e c i e s p e r month.  - 44  collected  PLANT SPECIES  MONTHS (% d i e t )  H 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17.  •Calamagrostis rubescens *Poa j u n c i f o l i a * S t i p a comata *Hordeum j u b a t u m *Calamagrostis neglecta •Glyceria borealis Stipa richardsonii Agropyron sp. Muhlenburgia richardsonis Oryzopsis a s p e r i f o l i a *Stipa occidentalis Poa i n t e r i o r Alopecurus aegualis Agrostis sp. Spartina g r a c i l i s Beckmannia s p . Unidentified grasses  TOTAL GRASSES  July  June  Grasses  C  H  August C  18.55 11.25 17.75 10.6 7.73 4.48 7.65 5.6 4.78 8.0 2.75 4.48 1.4 .93 .18 1.28 1.03 .9 1.0 .48 .8 1.33 .3 .23 .13 .08 .68 .08  15.83 10.08 9.05 2.83 6.1 4.88 4.4 7.13 6.23 9.23 8.28 3.6 1.53 .53  63.59 50.96 64% 51%  56.7 57%  5.15 10.6 9.1 11.73 7.7 9.5 3.58 4.3 2.18 7.1  4.25 2.45 15.35 19.95 7.13 9.15 2.1 1.6 3.13 2.33  .83 1.6 6.1  1.3 .83 6.55  .95 .2 .3  .25  H  42.08 21.18 4.0 6.88 5.75 1.43 2.5 2.15 1.3 3.2 1.8 3.75 .38 .65 .13 .85 6.53  1.08 52.6 53%  C  September H  C  30.0 23.63 3.53 7.63 4.3 1.83 2.4 4.18 5.48 1.0 3.33 4.05 .6  1.3 1.5 4.83  1.3 .4 5.95  .6 2.08 4.88  .18  .13 .35 1.43  .28 .65 .5  62.08 51.37 51% 62%  Winter H 7.0 1.65 3.95 2.8 6.15  5.0  8.15  60.23 50.28 34.7 60% 50% 35%  Rushes /Sedcres 18. 19. 20. 21. 22.  •Juncus b a l t i c u s *Carex a q u a t i l i s *Carex r o s t a t a Carex a t h e r o i d e s *Carex c o n c i n o i d e s  4.88 6.2 9.63 11.28 9.23 6.18 2.18 2.7 4.45 5.23  5.23 17.15 4.25 8.78 21.25 12.35 5.38 16.15 10.38 .4 .35 5.0 1.15 3.83 continued.  PLANT S P E C I E S  MONTHS (% d i e t )  Grasses H 23. 24.  Eleocharis sp. U n i d e n t i f i e d rushes/sedges  TOTAL RUSHES/SEDGES  July  June  .25 1.3  C  5.73  35.43 42.79 35% 43%  H 7.48 .95  August C  6.03  40.39 41.5 40% 42%  H 1.88  C 1.78  34.87 30.75 35% 31%  September H 1.48 .23  Winter  C  H  .15  .95  30.52 36.19 47.75 31% 36% 48%  • D o m i n a n t 11 f o r a g e s :  Other 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40.  (Forbs: Shrubs: Trees)  Astragalus miser Pinus contorta Rosa a c i c u l a r i s Picea sitchensis •Fern' Artemesia f r i g i d a Arctostaphylos uva-ursi P o t e n t i l l a hippiana Shepherdia canadensis V i c i a sp. Achillea millefolium S a l i x spp. Sisyrinchium angustifolium Geum t r i f l o r u m Hippuris vulgaris Aster conspicuous  .08  .38  .15  .38  .08  .58  .88  .25 .2  1.48  .25  .1  .68  .35  .3 .13 .08 .08  .18 .33 .43 .2  .03  1.6  .35 .45 1.38  1.63 .55 .15  2.58 .73 .6  .3 .4 .98  .15  .18  .4 .98 1.65 .63 .93 .73 .98 .8  .58 .28  .35  .23  .13  .15 .68 .18 .2 .18 .25  .33 .5 .98 1.8 1.43 1.13 .3  4.4 3.2 1.95 1.25 1.0 1.0  .5  continued.  PLANT SPECIES  MONTHS (% d i e t )  Grasses  , -j ,  June  July  August  H  C  H  C  Taraxacum o f f i c i n a l e Fragaria virginiana Spiraea b e t u l i f o l i a Antennaria p a r v i f o l i a Antennaria sp. Eriogonum sp. Sedum l a n c e o l a t u m Juniperus sp. Betula glandulosa Arnica cordifolia Solidago spathulata V i o l a adunca Cerastium arvense Heuchera c y l i n d r i c a Penstemon p r o c e r u s 'mosses* Thalictrum sp. U n i d e n t i f i e d Forbs U n i d e n t i f i e d Shrubs  .05  .08 .45  .13 .38  .13 .25  TOTAL FORBS; SHRUBS; TREES  1.02 1%  41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59.  .1  .3  .08  .2  5.8 6%  .08 .08  .38  3.02 3%  6.08 6%  H  C  .43 .63 .68  .23  .33 .75 .23  .18  .13  .13 .48 .9  C  .08  .05 .25  .28 .38 .23 .15 .08 .08 .13  H  98  September  .23 .28 5.43  3.24 17.5 3% 18%  .58 .15 .15 .1 .08  .38  Winter H  .5  .8  .55 .38  2.18 1.68  .63  2.8  9.17 13.88 17.4 9% 14% 17%  whereas  cattle  u s e o f C. r u b e s c e n s  t h e summer a n d p e a k e d i n S e p t e m b e r . difference  i n pattern  increased  steadily  T h e r e was a l s o a s i g n i f i c a n t  o f use by horses  and c a t t l 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 Hordeum  -iubatum.  and G l v c e r i a  over  borealis.  f o r the balticus>  Finally,  i n only  t h r e e s p e c i e s , i _ a i u n c i f o l i a . J u n c u s b a l t i c u s and S t i p a c o m a t a . was  the difference  to  be s i g n i f i c a n t l y  8).  i n use between dependent  horses  and c a t t l e  on a p a r t i c u l a r  I t s h o u l d be n o t e d t h a t f o r b o t h  month  i n an i n t e r a c t i o n  because  of the former  and  always  context,  than  cattle,  less of the l a t t e r . Considering  forages of  the consistently  species actually  available  t o them.  the  area  study  Reduced  and June  horse  was a s f o l l o w s :  = 4.69%, A u g u s t  t o how many  O f 88 f o r b s  horses  (Table 7 ) .  diets  o f t h e 11  (Appendix  utilized  Winter  s p e c i e s were  I) i d e n t i f i e d i n  a p p r o x i m a t l y 16, and  dependence diets.  plant  on f o r b s  The p o r t i o n = 2.75%,  identified  (Appendix  cattle  was r e c o r d e d i n  of forbs  June  i n horse  = 1.02%,  = 3 . 0 1 % , a n d S e p t e m b e r = 7.53%.  of g r a s s e s and sedges  number  e a t e n b y t h e two h e r b i v o r e s p e c i e s  considered i n relation  winter  high proportion  i n t h e d i e t s o f b o t h h o r s e s and cows, t h e t o t a l  plant  was  20  a t e more  (Table  C. r u b e s c e n s o r , C_. a q u a -  t i l i s . m o n t h was n o t i m p o r t a n t horses  found  July  Of 43 s p e c i e s  I ) , horses  utilized  22 a n d c a t t l e o n l y 19 ( T a b l e 7 ) . A  factor  further  the use o f t h e sedges they =  formed  indicative  i s found i n  C a r e x a q u a t i l i s a n d Q_. r o s t a t a .  a significant  19%) and c a t t l e  of s e l e c t i v i t y  portion  of the diet  (X = 2 4 % ) , b u t w e r e - 48 -  found  Together  of horses  (X  o n l y i n sedge  TABLE 8 A v e r a g e u s e p e r month o f the 11 f o r a g e s composing a t l e a s t 8 0 % o f the d i e t s o f b o t h h o r s e s and cows, with an a n a l y s i s of v a r i a n c e (using transformed data) comparing the 11 plant species f o r u t i l i z a t i o n per month, between s p e c i e s and on an i n t e r a c t i o n b a s i s : Plant Species  June  July  Calamagfostis H rubescens C  15.8 18.6 5.5 5.7 + 11.3 10.2 ± 3.2 ± 1.4  42.1 5.9 21.2 ± 8.3  30.0 + 19.2 23.6 + 5.3  Carex aquatilis  9.1 15.4 2.3 + 8.0 11.7 20.0 ± 3.7 ± 2.8  9.6 7.1 11.3 ± 6.4  8.8 1.5 21.3 ± 11.4  Ppa juncifoiUa  Carex rostata  Juncus balticus  H C  H C  H C  H C  +  +  17.8 3.9 + 10.6 ± 2.5 ±  9.1 4.9 2.8 1.9  7.7 2.4 + 9.5 2.2 ±  7.1 2.3 9.2 5.3  10.6 4.7 ± 5.2 + 1.9 ±  4.3 2.9 2.5 1.4  +  + + +  xa  August September +  ±  + ± + ± ± ±  4.0 2.3 6.9 3.6 9.2 2.6 6.2 3.2 6.2 1.2 4.9 2.6  +  ± ±  ±  ±  ±  10.4 4.2  +  ±  3.3  ±  + ±  5.4  4.3 2.1 5.2 1.5  Month  26.6 + 15.9 6.44* 16.5 ± 7.70  7.6 1.1 3.5 2.2  +  F  +  ± ±  10.7 5.7  16.1  2.61  F  Species interaction 8.11*  1.12  5.82*  1.25  7.8  9.6 6.0 16.2* 5.9 4.0 8.6 3.0 7.6" 3.8 6.4 2.9 5.2 1.9  .108  1.88  12.1*  .80  7.88*  4.54*  2.23  3.63*  continued...  Table 8 Plant  (cont) Species  Stioa occidentalis  June H C  Calamaarostis H nealecta C  + + + + +  Stipa comata  H C  Carex concinoides  H C  Hordeum jubatum  H C  +  Glycepia borealis  H C  +  a  5T = O v e r a l l  + + +  +  +  July  0.8 1.1 + 1.3 1.5 +  6.1 4.6 6.6 2.2  4.8 4.8 + 8.0 2.5 +  6.2 3.3 9.2 3.3  7.7 2.3 + 4.5 0.5 +  6.1 1.7 4.9 2.1  2.2 1.6 + 7.1 4.2 +  3.1 2.6 2.3 1.4  5.6 2.9 + 7.7 1.7 +  4.4 2.2 7.1 2.  2.8 2.0 + 4.5 2.3 +  3.6 0.6 8.3 1.7  mean f o r 4 mo n t h s .  A u g u s t September + + + + + + + + + + + +  6.5 2.9 4.8 2.3 1.3 .6 3.2 1.8 1.4 1.7 5.8 3.0 5.2 5.4 4.5 2.6 2.5 1.6 2.2 1.1 1.8 0.2 3.8 3.3  + + + +  6.0 2.8 4.9 1.7 5.5 4.5 1.0 1.2  +  1.8 1.0 4.3 1.0  +  5.0 3.6 3.8 2.9  +  + + + +  2.4 1.3 4.2 1.1 3.3 1.4 4.1 1.4  F  + + + + + + + + + + + +  4.8 3.7 4.5 2.5 4.4 3.8 5.4 4.1 4.3 3.2 4.9 1.8 3.9 3.5 4.4 3.2 3.7 2.3 5.3 2.7 3.0 1.7 4.9 3.2  Month  F  Species  F  Interaction  6.15*  .1  5.61*  .72  2.78  5.04*  .83  6.99*  .69  .21  1.56  9.9*  5.8*  1.04  2.91  7.04*  1.17  .53  meadows and  which  were  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  i n scattered  locations.  formed  a dominant  portion  cattle  were  only  found  Other  plant  of the diets  i n d r y meadows  C_. a t h e r o i d e s . a  and G l y c e r i a  disproportionately  for  both  total  horses  plant  formed  borealis)  s p e c i e s which  of both  (Appendix  IV).  Although  was  indicated  ( T a b l e 5) , a s m a l l  species available  comata.  ( i . e . C. n e o l e c t a .  h i g h u s e o f meadow h a b i t a t  and c a t t l e  h o r s e s and  ( i . e .S t i p a  S_. o c c i d e n t a l i s ) . o r i n s e d g e meadows o n l y  area,  number o f t h e  i n wet a n d d r y meadows  actually  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 animal s p e c i e s  (Appendix The  IV, Table 7 ) . presence  identified outstanding  of Eleocharis  sp. , i n t h e d i e t s ,  b y t h e W.S.U. l a b o r a t o r y , example  indicative  positively  t  appears  t o be t h e most  of s e l e c t i v i t y .  Based  on t h e  p e r c e n t o c c u r r e n c e i n h o r s e 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 forage plant actually  (X^  s u m m e r  identified  Eleocharis  SJD  v  =  2.77%. T a b l e 7 ) .  on t h e s t u d y s i t e  makes up a r e l a t i v e l y  H o w e v e r , i t was  and i t i s assumed small  proportion  vegetation cover i n the study area, or i s extremely The  overall,  average  number  of plant  summer X = 2 9 ; H o r s e - w i n t e r X" = 2 1 ; T a b l e most  drastically  reduced  13 g r a s s  the that  winter;  species,  7).  (i.e. The  was t h e g r a s s e s  a  while this  reduction  was  of almost  the category of "unidentified  t o b e 'new' g r a s s e s , b u t r a t h e r  decreased  half.  of the  localized. by  Horse-  botanical  (Table 9 ) .  I n t h e summer h o r s e s u s e d a maximum o f 16 and a m o n t h l y of  that  s p e c i e s used  h o r s e s was f o u n d t o be l e s s i n w i n t e r t h a n summer  group  never  average  t o seven i n  I t s h o u l d be  noted  g r a s s e s " i s assumed n o t  'known' g r a s s e s i n a n u n i d e n t i -  - 51 -  TABLE  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 . showing o r d e r  b a s e d on  p e r c e n t use , and  of s p e c i e s i n the w i n t e r  Order  Summer grass species  (%)  the  reduction  diet.  Order  Winter grass species  Y 1. 2. 3. 4. 5. 6. 7.  X  Calamagrostis 26.6 rubescens Poa j u n c i f o l i a 9.6 Stipa 4.8 occidentalis Calamagrostis 4.4 neglecta S t i p a comata 4.3 Hordeum j u b a t u m 3.7 G l y c e r i a b o r e a l i s 3.0  1. 2. 3. 4. 5. 6. 7.  S u b - t o t a l : 56.4 8. 9. 10. 11. 12. 13. 14. 15. 16. 17.  (%)  Oryzopsis 1.0 asperifolia Stipa .8 richardsonii Muhlenburgia .8 richardsonii Unidentified grass .7 Agropyron sp. .3 Spartina g r a c i l i s .3 Beckmannia spp. .2 Poa i n t e r i o r .1 Alopecurus .1 A g r o s t i s sp. .1 TOTAL: 60.8  - 52  -  Unidentified grass Calamagrostis rubescens Glyceria borealis Stipa occidentalis S t i p a comata Calamagrostis neglecta Poa j u n c i f o l i a  8.2 7.0  Total:  34.9  6.2 5.0 4.0 2.8 1.7  fiable  condition  Habitat  Lab, 1981).  made and  (Bruce  up t h e b u l k  Davitt,  The same c o r e - g r o u p o f f i v e of that  b o t a n i c a l grouping  summer d i e t s o f h o r s e s  amount  P e r s . comm., W.S.U. W i l d l i f e  o f use o f these  (Table  five  f o r both  winter  1 0 ) . However, t h e i n c r e a s e d  species  n o t a b l e , i n c r e a s i n g f r o m 33.2 p e r c e n t in winter of the t o t a l diet  rushes-sedges  during  the winter  was  i n summer t o 48.0 p e r c e n 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 content of  i n t h e w i n t e r t h a n some g r a s s e s , b e c a u s e o f t h e p r e s e n c e  green  material  analysis winter  carried  i n the center  plants  (dry b a s i s ) , parts  A nutrient  o u t on sedge m a t e r i a l c o l l e c t e d during t h e  o f 1979-80,  sedge  o f sedge p l a n t s .  showed  had a crude while  that  a sample  protein  from whole  (CP) l e v e l  ungrazed  o f 3.78  percent  t h e s e l e c t e d green m a t e r i a l from t h e i n n e r  of the plants  h a d a CP v a l u e  o f 6.5 p e r c e n t  (Figures  o f s h r u b s was n o t e d  i n horse  15 a n d 1 6 ; A p p e n d i x V ) . An diets  increased  concentration  f r o m s p r i n g t o w i n t e r ; where J u n e = 0.0%,  August  = 0.23%, September  Conifer winter  needles (6.25%)  Based fecal  occurred  = 2.96%, only  winter  i n September  =  14.65%.  (0.55%)  and i n  counts  i n the  diets.  on t h e p r o p o r t i o n  analysis  and  J u l y = 0.58%,  (Section  of total  cover  2.6), the concentration  compared f o r t h e t h r e e b o t a n i c a l g r o u p s : and  'other'  ( i . e . forbs, trees  the  following Z-proportional  and s h r u b s )  formula  -  grasses,  53  -  rushes-sedges  (Figure  (Walpole,  o f u s e was  17), using  1982: 266-74):  TABLE 10:  Rush and sedge winter,  species  showing  n b y H o r s e s i n summer  incre  use  o f sedges  versus  i n the winter  diet.  Order  Summer sedge s p e c i e s  (%)  Order  Winter sedge s p e c i e s  X  X  1.  Carex  2.  aquatilis  (%)  10.7  1.  Juncus  Carex r o s t a t a  8.6  2.  Carex r o s t a t a  16.2  3.  Juncus  7.4  3.  Carex  aquatilis  12.4  4.  Carex  3.9  4.  Carex  concinoides  5.  Eleocharis spp.  2.6  5.  E l e o c h a r i s spp.  balticus concinoides  S u b - t o t a l : 33.2 6.  Carex  atheroides  7.  Unidentified Rush-sedges  balticus  17.2  1.2 1.0  TOTAL: 48.0  2.4 .6  TOTAL: 36.2  - 54 -  FIGURE The  percent  occurrence  i n t h e d i e t s o f h o r s e s and  of  the  17: t h r e e major  c a t t l e based  - 56  -  botanical  on f e c a l  groups  analysis.  where:  n-^ and <32  *  1  2  = 800*  ~ 2 D  n Horse-winter = The  horses in  =  n  July  and  groups  proportional  and  the  400  c a t t l e proved  i n s t a n c e s of August was  use  t o be  grasses  (Table 11).  significantly  and w i n t e r d i e t s  of  the  three  botanical  significantly in July,  The  and  proportional  - 58  -  by  d i f f e r e n t , except rushes-sedges use  d i f f e r e n t i n horses  (Table 11).  groups  in  of the three  between  summer  TABLE 1 1 :  Z-Values  f o r differences  g r o u p s on a s e a s o n a l  CATTLE vs. HORSES  HORSES vs. HORSES  i n proportional  use of b o t a n i c a l  basis:  (Grasses)  (R-Sedges)  (Other)  June  5.2*  3.2*  5.5*  July  1.6  0.8  3.0*  August  4.4*  1.7  10.0*  September  4.0*  2.1*  3.13*  8.8  4.3  6.5  summer  *Z g r e a t e r  winter  vs.  t h a n 1.96 = S i g n i f i c a n t  - 59 -  Chapter 4.1  4: D i s c u s s i o n  Habitat-use The  of  the  r e l a t e d to animal p e r c e p t i o n of  t r a n s e c t system  habitat availability  the  IDF  not  surprizing  was  forested, including  (b)  subzone  closed forest  from  habitat-use  a  t h a t o n l y one true  and  of  to  dry  cattle  entire as  these  habitat available,  interface  1942;  become  species  following:  essentially  cows  take  differ  non-forest  area.  Since  67.3  open f o r e s t ,  zone.  i t is percent  semi-open  However, c o n s i d e r e d  perspective, i t is interesting  Smith,  three  the  study  estimate  f o r e s t e d range,  h a b i t a t types  grasslands  have  adaptation and  the  and  human i n f l u e n c e s .  around an  of  reliable  to  note  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  Camp and  under  f o r the  a  f o r e s t - d w e l l e r (Franzman, 1978).  adapted 1974;  provided  is classified  that  forest,  a  used  availability  and  cattle  to  Walker, 1975).  cosmopolitan The  crux  in a How  of  the  ( i . e . how moose  primarily  parkland  (Groves,  However b o t h  in their  adaptive  habitat  are  exploitation  similar  horses  What  are  they)?  h a b i t a t w i t h i n the  environment?  cows  does  How  of  revolves  and  form  in utilization  horses  distribution  f o r e s t e d environment  f o r e i g n environment?  from  Horses are  to  their  do  horses  forest  versus  In general, i t  w o u l d 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 forest,  and  an  overall  distribution  t h e r a n g e o f n o r t h e r n t r e e s and (Coady, 1 9 8 2 ) , c o u l d  be  shrubs  expected  this  or  p e r s p e c t i v e , i t would - 60  t h a n t o any  to exploit  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 From  more c l o s e l y  -  related  other  to  factor  forested habitats  cattle. appear  from  the  study  r e s u l t s t h a t moose a r e 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 of  a l l fecal  appear  counts  somewhat  open a r e a s  were  shrub  65.3  percent  f o r horses  only  1.4  1.7  carr.  Horses  predictable i n being  (meadow and  and  in forest.  percent,  cows  fecal  76.9  percent  than  cattle.  respectively  Horse  fecal  f o r cows, were  although  found  in  counts  indicated  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 . there i s a hidden habitat of  availability,  the  forest  observations that horses with for  factor. diet  species  during use  B a s e d on  two  of  itself  summers on  distinct  difference  on  open  my  study  graze  the  overuse  study  by  and  site,  areas  counts.  horses general  i t appears  found  i n open a r e a s  i n these  two  forms  forests  there  o f o p e n a r e a s was  with  over-using  There  observed  is a  forested habitat (e.g.  f o r e s t e d areas,  i n getting the  fecal  or i n f o r e s t  t h a t where o p e n a r e a s  is difficulty  without  of  forest  their  ( i . e . I n t e r f a c e Zones).  clearcuts) alternate  site,  Cows u s e d  o f t h e t i m e , and  (1979) a l s o o b s e r v e d or  34.1  However,  use  rubescens):  the  fecal  o n l y 15 p e r c e n t  to  grasslands  a  t w i c e as much a s c o w s , a c c o u n t i n g  a l l horse  adjacent  McLean  ( i . e .higher  Calamaqrostis  c o n c e n t r a t i o n s were o t h e r w i s e  use.  forest  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  percent  h a b i t a t by  shrub  f e c a l concentrations versus  results  open h a b i t a t s ) a t l e a s t 31.2  to  concentrations being  H o w e v e r , h o r s e s a p p e a r t o be more i n c l i n e d t o use  habitat  also  s t r o n g l y attracted  carr),  and  and  percent  cattle  open a r e a s .  t o o c c u r on my  study  as to  Such  site.  Moose t e n d e d t o use t h o s e h a b i t a t t y p e s ( i . e . o p e n , s e m i - o p e n , closed  forest  cattle  used  and  the  shrub  least.  c a r r ) that horses In  addition, - 61  -  and,  although  particularly, the  pattern  of  habitat  u s e was  more  s i m i l a r between horses  and  t h a n b e t w e e n 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  cattle  herbivores  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 . This  variability  spatial  i n habitat  separation  of the three  this  form  of spatial  all,  that  the transects  the  relative  highest on  amount  fecal  f o r horses  amount  of both  herbivore  the highest  Meadow a n d  Interface  contiguous  forest  ( i . e .not including  it  was  were  f o r cows The  made  was  on  greatest  I n d i c a t i v e of the  appear  a habitat  to  types  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  horses  carr?  species  Zone, t h e h a b i t a t  which  on T - I .  of  and s e c o n d l y , t h e  count  t o a v a i l a b i l i t y , were f o u n d on T - I V .  found  i n regard  on T - I , and f o r moose o n T-V.  t h a t c a t t l e appeared  with  type  of  Supportive of  significantly  habitat  f o r each  a degree  were t h e f i n d i n g s , f i r s t  varied  of each  counts  herbivores.  separation  different transects:  T-IV,  s e l e c t i o n produced  T-V  had  type  t o move  by  through  f o r e s t , t h e most  Interface  f a r the largest  moose u s e d  a v a i l a b l e , and which  freedom  Zones),  amount  of  was  shrub  d i s p r o p o r t i o n a t e l y more  than  h a v e a t t r a c t e d them t o  this  may  transect. Subsequently, standing  o f what were f e l t  involved  i n habitat  move a b o u t (discussed of  horses  that  the data  more  t o be t h e t h r e e  use, i n that  i n S e c . 4.2);  cattle  appear  b o t h h o r s e s and c a t t l e d i f f e r  use o f h a b i t a t s w i t h i n t h e  under-  than  horses cattle patterns  d i f f e r e n t ; and  g r e a t l y f r o m moose i n t h e i r  environment. - 62 -  indicated  the adaptive  somewhat  an  considerations  environment  that  t o be  toward  major  evidence  readily i n a forested  further and  d i d contribute  There  i s a potential  the  habitat  types  and  b) a r e m o s t a c t i v e  and  Telfer  f o r misinterpretation i n determining  i n which  time  that  moose  (1980), noted  i n upland  time, Cairns  range, b i s o n (Bison  g r a s s l a n d s , but spent the  tremuloides).  most a c t i v e  t h e most  ( N e f f , 1968).  t h a t on A l b e r t a  i n aspen (Populus were  a) s p e n d  (e.g. t r a v e l l i n g )  b i s o n ) , were most a c t i v e most  animals  i n forest,  They a l s o  but spent  t i m e , p r o b a b l y f e e d i n g and b e d d i n g , i n s h r u b l a n d .  noted  t h e most  I t i s important  t o n o t e t h a t i t i s n o t 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 these  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  unless in  i t c a n be d e m o n s t r a t e d  association  immediately  t h a t d e f e c a t i o n s occur  w i t h some o t h e r a c t i v i t y ,  adjacent  t o bedding  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 for  particular  habitat  types  sites.  its  more  than  ways, but i ti s  General observations  i n open f o r e s t .  t o c l o s e d f o r e s t , which  would  availability,  spruce  have been expected  and tended  t h e y used  three  i n proportion to  t o bed i n r a v i n e s w i t h heavy  attempt  was  made  by  Stcarrar - e t :al>'f19?7 )^*to f establish:' -;a s.  r e l a t i o n s h i p b e t w e e n h a b i t a t s e l e c t i o n and f o r a g e feral  horses.  a positive (i.e.  Moose  cover.  An  in  moose and  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  a strong a f f i n i t y  times  as o c c u r r i n g  Horses,  i n different  t o bed i n meadow-margins, and h o r s e s had  primarily  i n t h i s study, a l l selected  n o t p o s s i b l e t o s a y f o r e x a c t l y what purpose. throughout  such  counts,  availability  Using m u l t i v a r i a t e a n a l y s i s of f e c a l  response  by h o r s e s  to increasing  f o r e s t h a b i t a t ) was i n d i c a t e d .  however, due t o l a c k  of diet data.  counts,  pinegrass  R e s u l t s were  cover  inconclusive,  S a l t e r and Hudson  (1978a;  1979)  felt  that  determinant Alberta, and  of  availability  for habitat  and  of  forage  was  the  primary  s e l e c t i o n of f e r a l horses i n southern  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  grasslands.  since  the  This  conclusion  i n t h e i r study the  h o r s e s was  a grass  i s somewhat c o n f u s i n g ,  however,  s i n g l e most s i g n i f i c a n t d i e t c o m p o n e n t species  most o f t e n  found  in  forested  most  strongly  habitats. In for  but  fecal  rubescens.  findings 1968;  of  value  use.  of  of  strictly  a  cattle total  showed  various  highest  not  fecal  f o r e s t , the  the  plant  Salix  spp.  counts  1979;  based  (42%)  supports  authors  the  (Neff, estimate forage  done on moose, i n t e r p r e food  habits  contention  forage  f o r moose were the  was  established that  fecal  selection. found  in  lowest plant species  The  closed  variety.  a v a i l a b i l i t y and l o c a t i o n ,  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 1978;  P e e k , 1 9 7 4 ) , and  i f a v a i l a b l e , i n the of  w h i c h are  depend  summer  heavily  (Peterson,  f o u n d t o any  extent  forest habitat.  Johnson regarding  on  related to  Coady, 1982), n e i t h e r  in closed  not  v a r i a b l e , d e p e n d i n g on  plants,  other  a l s o supported the  (Franzman,  aquatic  This  communities to animals for  data  directly  t h e y t e n d t o use  transect  most d o m i n a n t f o r a g e  forest species.  h a b i t a t type with  Moose d i e t s a r e  f e c a l counts from  t h a t p e l l e t g r o u p s do not r e l i a b l y  researchers, are  selected  e i a l . , (1981) and  habitat-use  other  counts  on  shown by  analysis  Miller  and  A l t h o u g h a d i e t a n a l y s i s was  tation  but  horses  L e c k e n b y , 1968)  the  by  study,  meadow h a b i t a t as  data, C.  my  (1980) warns a g a i n s t  making  absolute  statements  u s a g e v e r s u s a v a i l a b i l i t y d a t a , because o f t h e - 64  -  inherent  .arbitrariness habitat the a  use  can  the  case  delimited into  the  natural a  the be  values.  a  habitat types,  selection  in  of  result  or  the  simply  of  range c a t t l e ,  niche  being  Additional  equation  being  with  the  Only  t h a t of  two  question  environments has  been c o l l e c t e d  and  why,  horses  With such  as  been p l a c e d i n a  seasonal  quantity  and  but  uniform.  introduced their  showing  t o more h e a v i l y f o r e s t e d  by  f e r a l horses  i n forested  S t o r r a r . e t al(-1977) .Not• enough data',.;  a b l e t o understand  under what c o n d i t i o n s ,  habitats.  variability,  and  ' r e a l ' versus  quality  from  fluctuate  season  example, P r i c e  buselaphus  season  to only  digestible nutrient one  availability when w a t e r  forage  t h e a b i l i t y o f t h e h a b i t a t t o meet n u t r i t i o n a l H a b i t a t s may  For  least  'apparent*  to  wildly  season,  in  or  was  levels  ranged  percent  observed  relatively  the  effect  d u r i n g my  i n sedge a r e a s , - 65  -  i n Coke's h a r t e b e e s t  from  i n the  i n t a k e v a r i e d from  i n s t a n c e of  to  (1978) showed t h a t a c t u a l p r o t e i n  cokei), 17  nutritive  from year  maintenance needs o f the h e r b i v o r e remain  (Alcelaphus  At  be  g r a z e r , but  i n t a k e , r e l a t i v e t o t h e presumed r e q u i r e m e n t s  wet  may  or  certain  of f e r a l horses because of  select particular  requirements.  the  area,  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 c o n s i d e r e d  availability,  year,  defined  h a v e a l r e a d y made  in a particular  open p l a i n s  in  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  1978;  t o be  investigator  arbitrariness  h a b i t a t use  (Salter,  the  having  adaptation  other  of  how  found  i n the case  s u c c e s s f u l secondary  habitats.  of  example, a r b i t r a r i n e s s  f a c t that animals  by  area.  For  148  dry  percent  season;  in  total  196 p e r c e n t t o 56 p e r c e n t . of  c l i m a t e on  field  p o n d s and  study.  habitat In  1979,  c r e e k s were v e r y  low  o r d r y , a l a r g e n u m b e r o f w e t meadow a r e a s  grazed high  and t r a m p l e d , by September. summer r a i n f a l l ,  levels, to  I n 1 9 8 0 , due t o  and subsequent  many s e d g e a r e a s  were  unseasonably  abnormally  inaccessible,  heavily  high  forcing  water animals  u s e d r y meadow a r e a s a n d , p e r h a p s f o r e s t , more h e a v i l y . Van  the  Valen  paradox  green This  (1973 i n White,  where  i s the paradox  which  food  affect  its  limited  o f ' r e a l ' versus  t h e way i n w h i c h  a p o p u l a t i o n of domestic one y e a r ' s  1 percent and  diet  of the t o t a l  forage  I t i s not always  animals  will  i n Australia,  perceive  80 p e r c e n t  of plants comprising  forage a v a i l a b l e t o the animals  D u d z i n s k i , 1978).  avail-  t h e r e a r e s o many v a r i a b l e s .  sheep  was c o m p o s e d  i n a  features of a habitat  attractiveness t o animals.  to estimate  food  under-used.  'apparent*  to other  the usefulness o f t h e i r environment, For  by t h e i r  i s apparently greatly  a n d a l s o may a p p l y  possible  1 9 7 8 ) w o n d e r e d how t o r e s o l v e  of herbivores being  world,  ability,  of  were  Alternatively,  only  (Arnold  t h e favored food  might  be c o i n c i d e n t a l l y t h e most a b u n d a n t , o r a n i m a l s may have l e a r n e d to  favor i t .  vegetation were but  Animals  do n o t n e c e s s a r i l y  uniformly.  observed  In the present  utilize  study,  horse  t o u s e n o t o n l y t h e same meadows  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  habitat or groups  repeatedly,  meadows.  Some  open a r e a s  t h a t a p p e a r e d t o h a v e good f o r a g e w e r e n o t t o u c h e d  by  a l l summer, w h i l e s i m i l a r  cattle  grazed, type  a n o b s e r v a t i o n made a l s o b y Heyes  were  repeatedly  (1979) i n t h e same  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 .  e_ _ L . , in  areas  (1979) o b s e r v e d  B.C., t h a t a l t h o u g h  i n t h e w i n t e r d i e t s o f deer b i g sagebrush - 66 -  Willms  (O.hemionus)  c o n t r i b u t e d 26 p e r c e n t  of  the diet,  only 7 percent of available  plants  were  used,  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  complicate matters  may a p p e a r quality. range deer the  t o be adequate  'real'  i n quantity,  f o r a g e on o f f e r  but n o t adequate i n  W a l l m o §_t_ a l . , ( 1 9 7 7 ) d e t e r m i n e d  f o r a deer w i n t e r  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 a d e q u a t e b u t i t c o u l d n o t meet p r o t e i n  energy  being  requirement. caloric the  further,  need  l i t t l e  The l a t t e r  nutrient  being c r i t i c a l  o f a l l nutrients are linked  On a s e a s o n a l b a s i s ,  energy  requirements;  50 p e r c e n t o f t h e e s t i m a t e d  h a s been shown t o be t h e p r i m a r y  requirements  herbivores  over  and energy  f o r 14,000  shortage  (Owen ejt_ a l . , 1 9 7 8 ) .  since the  need t o which  (Blaxter,  1962).  i s a common p r o b l e m t o  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 b e i n s h o r t s u p p l y , c o m p o u n d i n g t h e p r o b l e m . McLean of  and T i s d a l e  i n the Southern  classes  markedly,  and t h a t  crude  fiber  o f common f o r a g e f r o m s p r i n g t o f a l l  forest  ranges  ranges.  It i s likely  that  increased, i n a l l on b o t h g r a s s l a n d  f o r a g e on t h e f o r e s t  p r o b a b l y d o e s n o t meet c a t t l e g r o w t h  requirements  a b o u t m i d - A u g u s t o n w a r d s , b u t may meet m a i n t e n a n c e through October The  (McLean a n d T i s d a l e ,  variabilities  availability  support  arbitrariness  in  investigator a patchy  involved Johnson's  from  requirements  1960). i n the concept  of habitat  (1980) c o n t e n t i o n o f t h e  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 the  Interior  B.C., t h a t t h e p e r c e n t a g e s o f c r u d e p r o t e i n and phosphorous  decreased  and  (1960) f o u n d ,  t o t h e a n i m a l , a n d seems t o d e p e n d enters the picture.  environment  i s a rather - 67 -  o n when  Optimal foraging idealistic  theory  approach  to  resource and  utilization  M a c A r t h u r and  will  forage  maximizing the of  in the  t i m e and this  with  an  Pianka  intake  4.2  'the  i s the  e n e r g y and  assumption  ( P y k e e_t_ a i . , 1 9 7 7 ) . horses,  choice  of  showed of  requirements.  minimizing  Inherent t o the that  animals  fashion,'  n u t r i e n t s , and  "fitness  usefulness associated  b e e n m a x i m i z e d by  natural  s e l e c t i o n , are  and  fecal  that  has,  by  spatial  natural  i t i s possible  s e l e c t i o n , and not  directly  range  comparable  separation  counts per  there  was  habitat  type  a significant  horses,  cattle  and  along  difference  the in  moose i n d i c a t i n g  t o some d e g r e e , 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  Evidence  expressed  by  several  to  cattle  in  their  §_£  a i . , 1977;  supported  authors  habits  that  than  S a l t e r , 1978).  concentrations  idea that  (1966)  patterns.  habitat  t h a t each s p e c i e s  Emlen  In t h i s context,  undergoing  D i s t r i b u t i o n patterns analysis  the  by  most e c o n o m i c a l  e n e r g y r e q u i r e d to. do i t .  t h e i r h a b i t a t use  transects the  in  a n i m a l ' s f o r a g i n g b e h a v i o r has  feral  An  developed  (1966) b a s e d on  of  c a t t l e , under a r t i f i c i a l in  animals,  habitats  theory  selection" that  by  the  horses are  they In  contention  are  to  previously  more s i m i l a r  moose  (Storrar  addition, although  i n d i c a t e d h o r s e s u s e d f o r e s t m o r e , and  fecal meadows  l e s s t h a n c a t t l e , b o t h s p e 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 in  the  animals  any  study by  other allowed  habitat  l a t t e r provided by  the  habitat  habitat  type.  The  transect  i n v e s t i g a t i o n i n t o the type  and  a l s o , i n regard  system used  distribution to  space.  of The  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 a n a l y s i s , i . e . are - 68  -  animals with  an  apparent  overlap (i.e.  the  i n habitat choice,  i n the  same p l a c e ) .  When t h e  l e v e l s of  of  separation)  seen  to  be  chance.  H o r s e s and  nor a v o i d i n g  t o be  avoiding  habitats.  and it  types  was  with  given  are  several  that  90  percent  cattle  almost an  are  If indicate  the  moose were  each other's  the  of  impossible  and  c o w s , and  five  transect  both  moose, the between  the  total  and  outnumbered  essentially  'lost'  might even choose  horses  use  shown  to  so  many c a t t l e  herbivore  population)  Therefore,  question  i s whether  h o r s e s i n any  given  ( i . e . that  together. - 69  -  the  i t would  horse  animals  association?  area,  i n c a t t l e numbers), or t h a t the be  of  a c e r t a i n amount o f a s s o c i a t i o n ,  indifference  to  associated  these r e s u l t s .  are  'everywhere.'  to avoid  even  species.  there  large  used  further,  concentration  cattle  fact that  unanswered  probable  seen  horses  graphs that  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 avoid  cattle  by  preferred  ( i . e . moose  i n t e r p r e t a t i o n s of  literally  important  seek, are  expected  However, l o o k i n g  a s s o c i a t i o n of  reflect  cattle  neither interacting  same r e s u l t  horses  area varied widely  could  (nearly  by  than with  example, the  occur,  but  s e e n on  other  There For  h o r s e s and  i n d i c a t e d , c a t t l e and h o r s e s were more o f t e n  each  i n any  the  meadows h e a v i l y ) .  clearly  or  conversely,  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  l e a s t used  cows u s e d  t h o u g h , as  be  not  (or  C a t t l e and  e i t h e r each other,  T h i s was  same h a b i t a t s  more t h a n w o u l d be  other.  essentially indicated  habitat  association  moose were f o u n d t o be each  the  were e v a l u a t e d ,  associated  with,  had  spatial  degree  were  necessarily using  i t would  numbers two  are  species  H o w e v e r , when h o r s e s  are  frequently in  certain places,  attempt in  found i n considerably  t o avoid  with  respect  use  only  horses by  choose t o d i s t r i b u t e themselves  A l l f i v e t r a n s e c t s show numerous i n s t a n c e s  are heavily  are frequently Considering  apparently  concentrated  strong  not found  bution  the very  small  affinity  of both  o f t h e two s p e c i e s  cows a r e e n t i r e l y meadow  habitat), meadow  (already  that at Kilometer  frequency  percent,  conversely,  used  extensively  horses  and cows f o r  s u c h an u n e q u a l  distri-  as i n d i c a t e d by t h e g r a p h s .  For  i f movements o f h o r s e s a n d motivated  i n terms  shown n o t t o be s o , f o r f o r e s t 14 o n T - l l l ,  ( i . e . one s u r r o u n d e d  percent  and  cattle  numbers o f h o r s e s , and  random and i d e n t i c a l l y  habitat  that  28.1 p e r c e n t o f t h e t o t a l  i t seems s u r p r i s i n g t o f i n d  e x a m p l e , i t d o e s n o t seem l i k e l y ,  of  i n areas  i n areas  meadow h a b i t a t , w h i c h c o m p r i s e s o n l y study area,  cattle  or a d e f i n i t e difference  moderately, s l i g h t l y , or not at a l l ;  cows.  the  species,  and c a t t l e  t o space.  where horses  frequency than  i t w o u l d 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  the other  t h e way h o r s e s  greater  w h i c h was a n i s o l a t e d  by e x t e n s i v e  of use i n d i c a t e d  a n d f o r cows, o n l y  f o r e s t areas) the  f o r h o r s e s w o u l d be 22.5  2.5 p e r c e n t .  As t h i s  example  i s n o t a n i s o l a t e d i n s t a n c e , i t would appear t h a t t h e d i s t r i b u t i o n of  horses  i s d i f f e r e n t from that  of cattle.  in  d i s t r i b u t i o n between t h e two s p e c i e s  This  underlines  difference the 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 a r e n o t n e c e s s a r i l y equal that  i n terms of animal  I t i s p o s s i b l e t h a t two a r e a s  s e e m 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  animal could  use.  perception. account  There  i n terms o f  a r e numerous v a r i a b i l i t i e s  f o rdifferences  that  i n u s e such as t r a d i t i o n a l use  patterns, distance  from water, d i s t a n c e  from s i m i l a r  sites,  human h a r r a s s m e n t , and many o t h e r s d e p e n d i n g on t h e s i t u a t i o n . In  the specific  T-III,  k e e p them from g a i n i n g access  forage area,  area.  Due t o a h i s t o r y  h o r s e s may  less  have  accessible  experience and  'learned'  areas  that  t h i s had a marked e f f e c t  range appears  t o share  lack  cattle  factors. from to  those  h a v e a much  either to  horses  moose  distribution  animals), 1974,  Forested  ranges' i n t h i s (e.g.  trails  v e g e t a t i o n use by c a t t l e . between  seem t o be a t t r i b u t a b l e  horses  to several  o f moose a r e  divergent  a n d moose h a v e b e e n  attachment  or cattle,  shown  t o f o r e s t e d h a b i t a t than  which would  preclude  interaction  S e c o n d l y , a s shown by t h e t r a n s e c t d i s t r i b u t i o n  tend  n o t t o be q u i t e a s w i d e s p r e a d  areas.  o f moose  and t h e i r  cited  ranges,  i n their  a s 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  use i n fewer  population  'hilly  i n d i c a t e h a b i t u a l routes  or c a t t l e ,  stronger  a great extent.  graphs,  of  with  of a l l , the habits  of horses  (1951)  ' g r a z i n g r o u t e s ' on h i l l y  o f a s s o c i a t i o n shown t o o c c u r  First  and  C e r t a i n l y h a b i t and  on t h e u s e o f v e g e t a t i o n .  a n d moose w o u l d  i n the  perception of a v a i l a b i l i t y ,  i n t e r - c o n n e c t i n g meadows) a f f e c t The  acceptable  W e a v e r a n d Tomanek  attributes  r e s p e c t , a s my o b s e r v a t i o n s  or  t o an o t h e r w i s e  of the region.  developed  t o use f o r e s t e d a r e a s  t o u s e t h e more i s o l a t e d  i t s distribution.  cattle  14 o n  o f human h a r r a s s m e n t  i n f l u e n c e an a n i m a l ' s  therefore  noted  and  o f t h e meadow a t K i l o m e t e r  t h e apparent reluctance of c a t t l e  could  and  instance  This (only  could  b e d u e t o t h e much  2.9% o f t h e t o t a l  relatively  i n Coady, 1982).  small  lower  number o f a l l  home r a n g e s  (LeResche,  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 a v o i d both horses and An  in  the  interest  i n s e e i n g whether t h e r e were c o n s i s t e n c i e s  distributions  period  of  time  short-term  of the  three h e r b i v o r e s over  l e d t o the comparison of the  accumulations  amount o f v a r i a t i o n  a  i n animal  distribution  and  from year t o year due  v a r i a t i o n , and  reduced  longer  long-term  of f e c a l m a t e r i a l on T-V.  p a t t e r n s c o u l d be expected as c l i m a t i c  cattle.  A certain  habitat  due  to  death  leadership,  factors  horses  cattle.  for  and  long-term  and  o f dominant important  or expanded animal  individuals  to s o c i a l  Statistically,  the  numbers.  Comparison  of  only just  the  two  travel  or changes i n  h e r b i v o r e s such two  graphs of  short-term d a t a , were shown to be  but t h i s d i f f e r e n c e was  use  to such factors  V a r i a t i o n c o u l d a l s o occur from changes i n ' t r a d i t i o n a l ' routes  and  as T-V,  different,  significant.  graphs,  h o w e v e r , y i e l d e d some  i n t e r e s t i n g i n f o r m a t i o n which tended  t o c o r r o b o r a t e the findings  f r o m t h e h a b i t a t and  distribution  a n a l y s i s made on the  •recent' transects.  T h e r e were t h r e e n o t a b l e  between the  two  transect  moose, t h e  of the  by  graphs:  'isolated*  and  14;  and  15,  which  the  The  r a t h e r pronounced  similarities use  of  use by h o r s e s , but l i g h t use by  meadows d i s t r i b u t e d  'absence' o f c a t t l e  consisted essentially  five  this  cattle  between K i l o m e t e r s  11  between K i l o m e t e r s 8 and  of s o l i d  forest  with  only  s m a l l , s c a t t e r e d meadows. The that  long-term  overall  versus the short-term data f o r T-V  distribution time,  indicated  p a t t e r n s o f the t h r e e s p e c i e s were  different  over  while  maintaining certain consistencies  involving  habitat preferences. -  72  It i s possible that variations -  in  habitat  The  use  area  use  are c y c l i c  of horse  may  or at l e a s t p a r t i a l l y  'dung p i l e s *  signal  changes  predictable.  i n v a r i o u s p a r t s o f the  in distribution  patterns,  study  as  some  go unmarked f o r a p e r i o d o f a y e a r o r more ( p e r s o n a l e s t i m a t i o n ) , while  at other s i t e s ,  begin  t o be  marked  a l o n g - n e g l e c t e d dung p i l e may  and  become ' a c t i v e *  o n c e more.  suddenly  This could  r e s u l t f r o m t h e a b s e n c e o f h o r s e s i n an a r e a , o r some u n i d e n t i f i e d reason  for lack  pile.  The  of  i n t e r e s t by s t a l l i o n s  significance  been f u l l y e s t a b l i s h e d 4.3  1964;  fact  Slade  similarity North  are:  has  1982).  The  indicate  limited and  a  a m o u n t o f work done  other species  horses  range  cattle  indicated  of s e l e c t i v i t y  and  e l k , and  (Antilocapra  total  plant  diet,  factors  80-86% of  the d i e t s  least  t h a t d u r i n g t h e months f r o m c a t t l e were exercising,,  i n t h e i r choice of forage  but  fairly  species. percent  r e p r e s e n t e d o n l y 7.4  percent  t o be a v a i l a b l e t o them.  ( i . e . the high degree o f cows and  a  June  80  species determined  T h e s e two  2)  americana).  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  148  g£  shown i n g e n e r a l t h a t d i e t s of f e r a l to  Hansen and  (Hansen  1977;  S e p t e m b e r , h o r s e s and  more o f t h e  potential  Clark,  analysis  degree  the  1982) , w o u l d  Hansen, 1977;  most s i m i l a r  For both  of  and  not y e t  horses are both g r a z e r s (Hafez,  t o moose, d e e r and p r o n g h o r n  through  and  and  diets.  has  dung  cows  with horses  Olsen  1977)  Diet  high  Godfrey,  America  1)  similar  and  cattle  in their  a l . . 1977; Salter,  that  to horses  ( S l a d e and G o d f r e y ,  D i e t c h o i c e by h o r s e s and The  in  o f dung p i l e s  i n a particular  of s e l e c t i v i t y  horses, respectively,  and being  composed in  o f t h e same 11  the choice of d i e t .  analysis  indicated  were  necessarily  not  forages) indicated  a large  H o w e v e r , h a b i t a t u s e and  diet  selections  made b y  distribution  horses  i n t h e same l o c a t i o n .  overlap  and  cows  Attempts  were  made t o a n s w e r t h 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 t h e s i g n i f i c a n t  amount  was  of  overlap,  which  whether  a difference  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 ? when and how  were p l a n t s p e c i e s u t i l i z e d ?  In o t h e r words,  Did either herbivore  h a v e 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 Upon apparent  analysis,  were  found  (e.g. g r a s s e s ) ?  to exist  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 .  in  t h e use  and  Carex  and  32.6  cattle,  differences  o f t h e two  major  For  i n the example,  f o r a g e s , Calamagrostis rubescens  a q u a t i l i s , w h i c h composed an a v e r a g e o f 37.3 percent, respectively,  not  existed  only d i d horses  of the d i e t s  always  percent  of horses  and  rubescens  (as  e a t more C.  much a s 2 1 % more i n A u g u s t ) , and cows c o n s i s t e n t l y more _. t i l i s ,  but  over  r u b e s c e n s was Horse  use  t h e summer p e r i o d ,  the p a t t e r n  significantly different  of t h i s  s p e c i e s peaked  f o r t h e two  aqua-  o f use  for  herbivores.  i n August, 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 varied  average  significantly  on a m o n t h l y could more  consumption  be  the  growth  o f s i x o f t h e 11 d o m i n a n t f o r a g e s  in their  use  by  both  h o r s e s and  b a s i s from June t h r o u g h September. result  o f a number o f  spurts occurring  factors,  in a plant  This variation such  indicates  for  cattle  both  and  this  as one  or  s p e c i e s making 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 . dentalis  cattle  Stipa  occi-  type of 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 ,  h o r s e s , use - 74  i s low i n June but i n c r e a s e s -  i n J u l y and r e m a i n s f a i r l y s t e a d y t h r o u g h September. O p p o r t u n i s t i c foraging  could  particularly  seek  happen t o f i n d significant six  explain  out a plant  i t .  differences  was u s e d  whereas horses diet  content  use  from  4.5 p e r c e n t  and  then  came b a c k  September.  their  used  than c a t t l e used  by h o r s e s .  grass species,  cattle  In addition,  increased their  i n J u n e t o a peak i n J u l y  (8.3 p e r c e n t ) ,  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 G l y c e r i a  (not, i n c l u d i n g  borealis)  showed  use between t h e two h e r b i v o r e s over  Horses  an average  3.0  flexibility i n  the four  o f 4.3 p e r c e n t m o r e P o a  p e r c e n t more t h a n  horses.  was o n e o f t h e t h r e e g r a s s e s , i n w h i c h  and  rubescens.  months.  juncifolia  i n t h r e e out o f f o u r months, b u t i n August,  nearly  used  t o the 2 to 3 percent level of  E i g h t of t h e forages  aquatilis  and c a t t l e  G l y c e r i a b o r e a l i s . f o r example,  more t h a n  held steadily  they  f o r C. r u b e s c e n s . t h e r e w e r e  i n t h e way h o r s e s  by c a t t l e  for this  i f a n i m a l s do n o t  s p e c i e s b u t e a t i t when  As i n d i c a t e d  o f t h e 11 d o m i n a n t f o r a g e s .  always  C.  some v a r i a b i l i t y  Poa  cattle  juncifolia  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 was f o u n d t o be s i g n i f i c a n t l y dependent on  a particular  variations  that  example, Carex September the d i e t s use  month. occur  It i s difficult  f o rthe  i n t h e u s e o f some p l a n t s p e c i e s .  rostata.  i n cattle  t o account  decreases  diets,  steadily  from June  through  while i t increases steadily i n  o f horses c o i n c i d i n g with a d e c l i n e i n o v e r a l l  by h o r s e s  from  July  For  through  September.  I think  sedge that,  outside of the top three or four forage choices f o r a particular geographical l o c a t i o n , i t i s probable that the order of specific plant  selection  by horses  and c a t t l e - 75 -  would  be s e e n  to vary  over I  a number o f y e a r s  base t h i s  and  lower  and  also  feral of  with  r e l a t i v e use because  the  Hansen, there  most o f  and  1978;  the  have shown a s i m i l a r  The  and  C l a r k , 1977;  H a n s e n and In the  pattern species,  decreasing) Reid,  present  1975; study,  a v e r a g e summer  C_. r u b e s c e n s  on  (26.6%), the  use fifth  neglecta  ( 5 . 4 % ) , and  the  rubescens eleventh  (4.4%).  of  horses  of b o t a n i c a l groups  grasses,  difference in  r u s h - s e d g e s and  u s e d an a v e r a g e o f 61 p e r c e n t  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  browse.  Horses c l e a r l y  are  more o r i e n t e d t o w a r d s  of  are  cattle  This  i s not  fiber for that  the  by  utilization  s p r i n g , summer a n d  rather dramatic surprising  horses  ruminants  switch  (Janis,  1 9 7 6 ) , and  i n the  Equid  -  species  particularly  t o sedges i n the  the  with  higher  in  winter.  tolerance  dental  than  to  adaptations  d e n t a l development exceeds  combination - 76  grass  fall,  c o n s i d e r i n g the  a highly fibrous diet. of  of  grasses,  grasses  of  average  'other.'  percent  i n the  an  the  51  light  used  diet,  (6.4%) and t h e e l e v e n t h c h o i c e , <_. n e g l e c t a  c h o i c e , C.  utilization  cattle  plant  species rapidly  cows showed a s i g n i f i c a n t  summer d i e t s  whereas  few  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  horses  monthly  a very  d i f f e r e n c e s among t h e  concinoides  erratic  s t u d i e s done t o d a t e  f o r c a t t l e , t h e f i r s t c h o i c e b e i n g , C.  (16.5%), the f i f t h  In  diet  other  choice of horses,  (3.0%); s i m i l a r l y  c h o i c e , C.  o f t h e somewhat  H a n s e n §_£ aj.. , 1 9 7 7 ) .  large  first  of  H a n s e n and  1976;  were  use  c h o i c e , Juncus b a l t i c u s  by  the  range c a t t l e  percentage  were done.  of the other p l a n t species i n the  ( i . e . h e a v y d e p e n d e n c e on  (Salter,  of  c o n t e n t i o n on t h e e v i d e n c e  horses  use  i f consecutive d i e t analyses  of premolar  complexity,  movement f o r w a r d o f t h e m a i n c e n t e r o f m a s t i c a t i o n and adaptations  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 t h e  dibulum, * a and  lower  structural  deep  invagination filled  incisors  found  1912;  Weller,  1968;  fecal  analysis  showed  only  with  c e m e n t on  i n modern e q u i d s  S i s s o n , 1953). that both  Although,  cattle  and  ' infunupper  (Lydekker, in general,  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 o f b o t a n i c a l groups  indicates  cattle  f o r lower f i b e r content  forb  their and in  species  utilization  holds the  decreased  eaten. of  and  At  shrub  i t s nutrient  fall  somewhat more  selective  material.  In w i n t e r , horses and  were b e i n g  winter  the  the  same t i m e ,  material,  content  number o f g r a s s , sedge  which  increased  i s more  b e t t e r than  (Nagy, 1969;  they  available  g r a s s e s or f o r b s  Cook, 1956).  This  sort  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  year-round  solely but  on  has  energy grass  not  yet  i n t a k e shown t o o c c u r i n h o r s e s k e p t  pasture  (Owen §i a ! . ,  1978).  It i s possible,  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 food i n t a k e during 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 such  as  deer  and  moose  rush-sedges  from  (Robbins,  1983). The  increased winter-use  summer t o 48% attributable exposed,  i n w i n t e r by h o r s e s , a l s o may to  unlike  Snow d o e s n o t  of  have been  availability.  Rush-sedges tend  forages  meadow a n d  appear  i n the  to c o l l e c t  i n the  33.2%  in  partially to  remain  forested areas.  sedge areas  despite  t h e i r open a s p e c t , o r p e r h a p s b e c a u s e o f i t ( i . e . wind e f f e c t s ) , combined  with their  inherent surface uneveness. - 77  -  The  sedge  a r e a s were o b s e r v e d plants  even  freezing offer  on w i n t e r  (pers.  obs.,  'pristine'  the w i n t e r , summer d u e  t o have  s h a l l o w , pooled water around  d a y s when t h e t e m p e r a t u r e was March,  grazing,  1980).  as  being inaccessable  f a r below  O f t e n the sedge  some c a n  only  be  the  areas  utilized  in  t o a n y t h i n g b u t moose i n t h e  t o w a t e r d e p t h and mud.  I t i s possible that  sedges  r e t a i n a higher n u t r i e n t content during the winter than grasses or  forbs.  for  The  mobile  the e x p l o i t a t i o n  sedge  plants.  rostata  lips  of horses would  of the green m a t e r i a l  This material,  ( B e r n a r d , 1974)  and C a r e x a q u a t i l i s  of  h o r s e s , had  Weathered  grasses i n winter  CP  values.  in  a  Comparative  'weathered'  October study  and  2.8%,  spp.  3.3%  a v e >  (Demarchi, collected 2.7  protein would  and  4.5  of  6.5  percent.  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  collected  s p . 1.89%, Poa and  Tisdale,  March,  during  t h e months o f  of s p e c i e s found i n the  Agropyron  1973), both Agropyron in late  level  spp.  spp. 3.3%  1960).  3.08%, Bromus . , and  Stipa  In a second  study  s p i c a t u m and  a v e  Stipa  such  1974) , and visible it  as  percent, respectively.  rhizomes  I t would  appear  by sedge p l a n t s i n t h e f a l l  the subsequent  portions  decline  of the p l a n t s  w o u l d be n u t r i t i o n a l l y  comata,  w e r e f o u n d t o h a v e a v e r a g e CP v a l u e s  even w i t h t h e t r a n s l o c a t i o n of m a t e r i a l s t o underground areas,  winter  e x p e c t e d t o have l o w e r  follows:  (McLean  Carex  (Gorham and Sommers,  percent of the t o t a l  November, r e p r e s e n t a t i v e  Koeleria  f o r both  be  condition,  a r e a , were as  spp.  of  a crude  allow  i n the center of  shown t o o c c u r  1 9 7 2 ) , w h i c h t o g e t h e r composed 28.5 diet  physically  that  storage  (Bernard,  i n n u t r i e n t content i n the  (McLean  and  Tisdale,  1960),  advantageous t o i n c r e a s e t h e u t i l i z a t i o n - 78  -  of  sedge  plants  over  grasses  so b e c a u s e p i n e g r a s s , by forage, basis  although  (McLean and  forage.  f a r the  nutritionally  M c L e a n and  Tisdale,  a decisive  i n c r e a s e i n the  is  i n the  related  for  bution for  and  use  C.  cattle  horses  and  cattle  diet  show  rush-sedges  by  both.  w e r e b o t h on t h e s t u d y s i t e  from  choice appeared  to a m e l i o r a t e the  two  species.  However, because cattle  t h e m i n t h e summer, t h e r e i s a p o s s i b i l i t y t h a t  cattle  from  was  adversely effect  June through part  single  actually use  w i t h the  some w e t tend  rush-sedges  potential  i n t h e w i n t e r , and  moderate  again,  results  September, the 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 -  r o s t a t a were the  combined  diet  1969;  selectivity  However, the  only  the  Freyman,  t o the a v a i l a b i l i t y of these p l a n t s , t o  significant  horses  (McLean and  content  whether i t  summer-use c o u l d and  (2.7%) and phosphorous  o f b r o w s e and  w i n t e r , i t i s not  d e p e n d h e a v i l y on  utilized  geographical  a p p e a r s t o be a p o o r w i n t e r  Although  i n t e r f e r e n c e between the  horses  a  horse  p o s s i b l e to say  June through  of  1960).  on  an u n u s u a l l y h i g h s i l i c a  higher n u t r i e n t content, or to While  a  variable  Freyman, 1969),  m a t u r e p i n e g r a s s has  Particularly  m o s t i m p o r t a n t summer  e x c e e d s 10 p e r c e n t by O c t o b e r  horses  winter.  In a d d i t i o n t o low crude p r o t e i n  levels, which  i n the  by  two  sedges  of  the  important  Juncus b a l t i c u s horses  and  Carex  u s e d most by  S e p t e m b e r , and  (28.5%) most  horses.  aquatilis horses  together they  winter winter  diet forage  (17.2%), which  cattle  of  and  formed horses.  plant for sustained  i n t h e summer.  This,  apparently exclusive winter-accessibility  meadow a r e a s  (even  i n very  dry  summers)  would,  to ameliorate the e f f e c t of p o s s i b l e i n t e r f e r e n c e . - 79  -  By  the  same t o k e n ,  range to c a t t l e . the  two  few  horses,  Salter  there  by  most v u l n e r a b l e cattle,  but  seem  diet  of s p r i n g  and  the  relatively  probability  5 percent  of  between  of  this.  non-forested  i n A l b e r t a showed e v i d e n c e o f  horses.  t o use  little  only  value  use-differences  shown t o o c c u r ,  that  site  on  the  spring  F o r b s w o u l d be e x p e c t e d t o be  by h o r s e s p r i o r  to the a r r i v a l of  analysis indicated only  a  the  range  1 percent  forb  by h o r s e s i n J u n e . The  estimated  d i e t s a l s o showed an  c o n i f e r needles  observed in  would  h i s study  (June) g r a z i n g  of  already  (1978) e s t i m a t e d on  undermine  However, based  herbivores  habitat  use  h o r s e s may  the  t o be  winter  expel  the  d i e t s of  horses,  I do  not  but  f e e l the  i s accidental considering  unwanted m a t e r i a l taken d e s i r e a b l e items  apparently w h i c h may content  winter  the  phenomenon  increased  use  of c o n i f e r  the  ability  of  1981).  oils  horses  A l s o , some w i l d  to  ruminants  of c o n i f e r needles during  ( p e r s . comm. w.  become a d a p t e d t o t h e essential  (1978)  considered  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  i n winter  H a b i t a t Lab,  Salter  use  i n t o t h e m o u t h , e v e n when m i x e d  (pers. obs.).  i n c r e a s e t h e i r use be  (6.25%).  i n the  i n c r e a s e i n the presence of c o n i f e r needles  accidental.  needles  with  a similar  during  increase  winter,  carbohydrate  B r u c e D a v i t t , W.S.U. W i l d l i f e  T h e r e i s e v i d e n c e t h a t rumen m i c r o o r g a n i s m s 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  found  in  conifer needles  (Wilson,  1969).  M o n o t e r p e n e a l c o h o l s a p p e a r t o be t h e w o r s t o f f e n d e r s i n d i g e s t i v e inhibition  ( A r n o l d , 1981), but  t h e i r e f f e c t , i f any,  no  r e f e r e n c e was  found  regarding  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  horses  of  a b r a s i v e roughages,  fecal  w i t h a d e n t i t i o n maximally developed  epidermal  drawback  of the  digestion general  of  f o r b s would  analysis  technique l i e s  browse,  forbs,  that  (Equus b u r c h e l l i i )  sedges  i t i s a good  largely  (Owaga, 1 9 7 7 ) ,  burros  (Woodward, 1976),  Reid,  1975).  Grass  Work to  and  f o r b s which  i s currently  determine  a  fragments  retain  i n the  and  grasses.  technique  horses  cattle  are  and  being  done a t W a s h i n g t o n  differential  digestion  cattle  diet  to  forbs significantly.  use  a total  level  utilized  e a c h month...  were t h e  o n l y ones which  out  by  Davitt  Geum).  remain  (pers.  comm., 1 9 8 1 )  quite  technique  and  resistant  than  University for  the  i s applied.  to date, horses Salter  have  (1978)  3 p e r c e n t f o r b s were  consistently."  that  possibly  t h e r e f o r e can  t h e r e may  of  5 t i m e s more t h a n  be  As p o i n t e d  t h e f o r b s t h a t do show  t h o s e t h a t have h a i r s  suggests  Fecal  less  appeared  further  epidermal  the  comparisons  of  I t i s suggested  u n d i g e s t e d and  4 to  (Hansen  " s p e c i e s i n the P o t e n t i l l a - G e u m groups  t e n d t o be b i a s e d t o w a r d s and  zebra  Hudson,  coefficient  versus  that  as the  State  In horse  states  animals  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 .  s p e c i e s to which  shown  is  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 ,  animal  been  for  elk  generally  major  There  ( S a l t e r and  various  not  The  by  differential  of g r a s s e s , such  1979)  unlike  identified  i n horses than i n c a t t l e .  i s made up  to digestion  be  analysis  agreement  whose d i e t  fewer  for utilization  be  the  factor  being  Potentilla  h a i r s tend to  identified.  a correction  are p r e s e n t l y  (like  up,  Davitt for forbs  identified  by  analysis. analysis  i n t h i s study d i d not n e c e s s a r i l y - 81  -  indicate  a weakness i n the was  relatively  technique  i n t h e i r u s e 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 o r  sedges.  did indicate  between c a t t l e  there horses  results  difference  f o r b s , as and  The  little  for identifying  consistently  higher  selection  for  f o r b s i n t h e d i e t by c a t t l e f o r J u n e , J u l y , August and September. Seegmiller  and  Ohmart  (1981) showed by  fecal  diet  analysis,  that  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 t h e p r e f e r r e d f o o d s  when  they  were  composing  as  s e e m s no  green  much a s  reason  forbs are  February  percent  total  of the  to  June,  diet.  There  diets  should  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  present.  be  justification  in  technique  and  56.5  s u c c u l e n t from  t o assume t h a t f o r b s i n h o r s e  be more d i f f i c u l t the  and  Conversely,  for attributing  rather than  t h e r e does not  'lack*  selectivity  on  of the  seem  forbs to p a r t of  horses  total  diets  of  horses  and  cattle  represented  a f r a c t i o n of the p l a n t s p e c i e s comprising the b o t a n i c a l example, both  rush-sedges  such  as  cattle.  Of  both  although  and  found  43  cows  grass 19.  h e r b i v o r e s was Carex a q u a t i l i s  For  important  were the  horses, forage  cattle  used  only  consistently and  t o be  by  Out horses  Overall, pinegrass was  the  most  o f 88 and  - 82  20  by  forage  (Calamagrostis rubescens) .  rubescens  -  forb  utilized  important  u t i l i z e d by c a t t l e o n l y  more t h a n  13  locations  s p e c i e s i d e n t i f i e d , horses  h o w e v e r , C.  plant;  utilized  groups.  relatively  i n various  C_. p a r r y a n a .  only  s i x of the  i n t h e s t u d y a r e a , i g n o r i n g some  s p e c i e s found  16  22,  less.  and  Carex pachystachya  species,  only  horses  found  widespread  for  error  cows. The  For  to  16  was  percent  slightly  by f a r t h e most utilized  than  the  next  favorite  (_.  aquatilis.  Salter  (1978)  determined  t h a t the 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 habitats the  on  h i s study  area  most dominant s i n g l e  d i e t component o f c a t t l e value  of  i n southwestern  forage  of  feral  F e s t u c a spp.,  most i m p o r t a n t component i n f e r a l h o r s e rubescens  on  B.C.  on  f o r e s t e d l a n d , and  acres)  ranges.  of t h i s  (Tisdale is  the  the  lies  largest  up  i s an  f a r the  bulk  i n the  s u b z o n e and  S t o u t and  but  varies  i n the  s p r i n g and,  the  fall.  Around  early  t o g r a z i n g ( S t o u t e_t a l . , studying  of a v a i l a b l e  and was in  my  July  forage,  The  partly  on  showed a p r e f e r e n c e  ( a l t h o u g h p i n e g r a s s was  zone  may  (Stout  m o i s t u r e , one  i t i s most  provided  50  readily  different  in  prior  sensitive  Brook, 1980).  (McLean, 1980).  McLean percent  acceptable mid-August,  Fecal analysis p a t t e r n , i n that i n June  t h e i r second c h o i c e ) , and -  on  growth s p u r t s ,  f o r Carex a q u a t i l i s  - 83  IDF(b)  i s s l o w i n g down  t h a t i t was  a slightly  (IDF)  exclusively  s u m m e r , b e c a m e u n p a l a t a b l e by  indicates  (6 m i l l i o n  Pinegrass  h a v e two  S t o u t and  a g a i n used i n September  is  forested habitat types.  growth  determined  species  i n B.C.  Brook, 1980).  t h a t i s the time 1980;  second  D o u g l a s F i r Zone  range where p i n e g r a s s  in early  study  cattle  July,  the  m i l l i o n ha  Coupe, 1979).  between  major  'clumped'  forage  i s l o c a t e d almost  depending  summer d o r m a n c y , a n d  to c a t t l e  was  rangelands  3.2  Interior  (Annas and  one  (1967)  which  t o 80 p e r c e n t o f t h e g r o u n d a r e a i n t h i s  a l . . 1980),  The  diets.  of  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  to  horses.  important  approximately  M c L e a n , 1957;  Fraser Plateau  occupy et  and  By  A l b e r t a , composed  i n S a l t e r ' s study (ignoring the  s e d g e s p e c i e s ) was  Calamaqrostis  i n forested  and  essentially  doubled  their u t i l i z a t i o n of pinegrass  My r e s u l t s of  Heyes  meadows  are also d i r e c t l y contradictory t o the observations  (1979) r e g a r d i n g from  late  and  palatability  use  by c a t t l e  the preference  o f t h e upland  of pinegrass,  in  August and September  my  own  observations  by  than  usual  which  1973;  sedge  forage d e c l i n e s .  t h e major  upland  The h e a v i e s t  f o r a g e , was i n  i n J u n e and J u l y .  In addition,  i n 1 9 7 8 , a d r y summer w h e n m o r e were a c c e s s i b l e t o c a t t l e ,  showed  sedge that  d r y a n d wet meadows u s e d b y c a t t l e  and h a d s u s t a i n e d h e a v y t r a m p l i n g damage,  meadows a r e e s p e c i a l l y  Heyes, 1979).  quality  u s e o f s e d g e s b y c a t t l e was l e s s  than  t h e end o f August, both  were s e v e r e l y grazed to  by c a t t l e o f w e t l a n d  summer o n , w h e n t h e n u t r i t i o n a l  September, and t h e o v e r a l l  meadows  f o r A u g u s t and September.  From June  through  vulnerable  (Millar,  September,  pinegrass  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 horse  the  even c l o s e were Poa j u n c i f o l i a i n  only  species  coming  June and Carex a q u a t i l i s pinegrass  roughly  i n July.  Horses c o n s i s t e n t l y u t i l i z e d  1.5 t o 2 t i m e s more t h a n  - 84 -  diets,  cattle.  Chapter  5:  Summary  Comparative herbivores  selectivity  (largely  t o some e x t e n t .  equids)  o f food m a t e r i a l s by monogastric and r u m i n a n t s  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 . idea  has been t h a t  forages such the of  o f poor  factors  1976;  nutritive  slow  quality  0rskov,  T r a d i t i o n a l l y , the p r e v a i l i n g  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  as t h e need  relatively poor  quality  (Swenson, 1970).  rate  of passage,  f o r a g e by r u m i n a n t s  hinder the utilization such  Other  high  quality  i n summer t o p r o d u c e  ruminants,  upon i n t h e w i n t e r  l a r g e amounts o f c e l l u l o s e capacity f o r increasing food q u a l i t y  of  food  (Janis,  they tend  such  as deer,  require  t h e f a t reserve they  ( S h o r t , 1981).  Deer cannot  digest  (Nagy, 1969) a n d h a v e a v e r y  limited  1973 c i t e d  i n W a l l m o fit al.,  1977).  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  choice  i n b o t h summer a n d w i n t e r ( M a c k i e _ t ai., 1 9 8 2 ; S h o r t ,  1981).  Until  required  quite  recently,  high quality  i t was a s s u m e d  t h a t the horse  forage, but recent evidence  indicates  the  ability  o f equids i n g e n e r a l t o e x p l o i t environments  are  either  o p t i m a l or sub-optimal f o r ruminants.  due  to their  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  i n the d i e t than ruminants (Olsen results  a  f o o d i n t a k e t o accommodate f o r d e c r e a s e d  (Amman e_t_ a l . ,  Subsequently  as c a t t l e  1 9 7 5 ; Hogan and W e s t o n , 1970; A n n i s o n , 1970;  1969).  depend  However,  f o r s u f f i c i e n t b a c t e r i a l energy and  Corbett, diet  h a s been examined  ( J a n i s , 1976).  and Hansen, 1977) and A f r i c a of f i e l d  which  This i s content  I n b o t h North America  (Bell,  1971; 1970) t h e  s t u d i e s h a v e shown r u m i n a n t  feeding strategy  - 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 . Data  collected  zebra  by  Bell  (1970) i n a s t u d y  (Equus burchej3,i b o h m i ) c o m p a r e d t o f o u r r u m i n a n t  showed  t h a t zebra:  content,  2)  utilized  the  A l t h o u g h h o r s e s and tative  o f two  h i g h e s t amount o f  completely different  niche  they  exploit,  they e x p l o i t the n i c h e . by  Evans  as  efficient  is  probably  levels  are  species protein  r o u g h a g e , and  3)  herbage.  mammalian f a m i l i e s  Subsequently, they  remain  though  are  with  convergent  divergent i n  To d a t e , t h e g e n e r a l c o n s e n s u s ,  (1977) i s t h a t h o r s e s i n the  by  cows a r e b o t h g r a z e r s , t h e y a r e r e p r e s e n -  different digestive strategies. the  use  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  c h o s e t h e commonest and most a c c e s s i b l e  in  of herbage  'how'  expressed  approximately two-thirds  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  this  t r u e w h e n 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 f e d ad_ l i b i t u m .  However, t h e r e are  a number  of  utilization  of  other factors to consider: 1)  The  horse,  with  more  efficient  direct  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 depends  less  on  products  b a c t e r i a l p r o t e i n ) t h a n do 2)  The  rate  of passage  of b a c t e r i a l  while  3)  the  digestion  of d i g e s t a i s f a s t e r  i n the  than  activity  of the  rumen  ( B a l c h and  the  (cited  S l a d e , 1974); A l e x a n d e r ,  The  horse  and  ruminants,  caecum  c o m p a r a b l e to t h a t o f i n Robinson  horse  i n t h e same t i m e p e r i o d  V a n d e r N o o t e ^ ajL., 1967)  cellulolytic  (and  cattle.  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 ( A l e x a n d e r , 1963;  sources,  remains  Johnson,  1950  1963).  seems t o be a b l e t o s w i t c h e a s i l y f r o m e x t e n s i v e - 86  -  d e p e n d e n c e on on  bacterial  such 4)  as  Under  raonogastric digestion  that  required  identical  digestion w i t h o u t an  'adjustment'  period  ruminants  ( S l a d e e£ a l . ,  1970).  by  experimental conditions,  t h e major p r o d u c t s o f b a c t e r i a l fatty  acids,  large  t o e q u a l dependence  has  intestine  absorption  f e r m e n t a t i o n , the  b e e n s h o w n t o be s i m i l a r and  the  ruminant  forestomach  1966; A r g e n z i o §_t_ a l . ,  Janis  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  for  the use  of herbage  the  greater  quantity  volatile  f o r the equine  and S t e t t l e r , (1976)  (Stevens  1974b). be  essential  above a g i v e n f i b e r l e v e l and, whereas  o f f o o d r e q u i r e d by h o r s e s u n d e r  adverse  c i r c u m s t a n c e s m i g h t be i n t e r p r e t e d a s i n e f f i c i e n t , J a n i s out that a  diet  this which  maintain foragers'  digestive  strategy  ruminants  of s i m i l a r  themselves. by  becoming  of  Horses  may  selective,  enables horses to exist body s i z e  well  prove  i.e. a  simply t o be  1977;  on  cannot  'optimal  'specialist,'  t h e r e i s f o o d i n a b u n d a n c e , and u n s e l e c t i v e , i . e . a when f o o d i s s c a r c e (Pyke e t a l . ,  points  when  'generalist,'  N u d d s , 1980; B e l o v s k y ,  1978). A  number  of  large  h e r b i v o r e s have been shown t o  h i g h l y v a r i a b l e d i e t s , depending and e n v i r o n m e n t . 1978;  Peek,  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  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  1 9 7 4 ) , mule deer  (Short,  1981)  and  1978).  D i e t s t u d i e s c o n d u c t e d on f e r a l h o r s e s and  either  separately,  adaptability W e l s h , 1975;  have  (Franzman,  elk  (Boyd,  range c a t t l e ,  t o g e t h e r o r w i t h o t h e r s p e c i e s , h a v e shown  t o various environments  ( C a s e b e e r and K o s s ,  F e i s t and M c C u l l o u g h , 1976; - 87  -  1970;  L e u t h o l d , 1977; B e r g e r ,  1975).  F o r e x a m p l e , i n t h e R e d D e s e r t o f Wyoming, y e a r - r o u n d  dominant spp.  foods  (36%);  was  of  range  cattle  fall  1978) .  s p p . ( 3 7 % ) and S t i p a  s p p . (54%) ( O l s e n and Hansen, 1977).  forage In dry  i n s o u t h e r n C o l o r a d o , t h e d o m i n a n t summer f o r a g e s  were D a n t h o n i a  Carex  were Agropyron  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  Agropyron  mountain  of horses  s p p . and Fescue  spp., while i n the  s p p . i n c r e a s e d t o 58% o f t h e d i e t  On t h e d r y , W h i t e  Sands M i s s i l e  (Hansen and R e i d ,  B a s e i n s o u t h e r n New  M e x i c o , t h e d o m i n a n t f o o d s o f h o r s e s i n t h e summer were dropseed ( S p o r o b o l u s spp.) and mesquite thistle  (Salsola  amount has  kali)  of comparative  shown a t e n d e n c y  forages  (Prosopis i u l i f l o r a )  i n w i n t e r (Hansen, work  conducted  on h o r s e s  Salter,  1977;  Olsen  1978; Hansen and C l a r k ,  and Hansen, 1977).  (Hubbard  has been  and  However, d i s t i n c t  little  attempt  cattle similar  and H a n s e n ,  1977; Hansen  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 . there  The l i m i t e d  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  w i t h i n the d i f f e r e n t environments  1976;  1976).  and r u s s i a n  a l . .  differences In general,  to coordinate diet  analysis  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 i n s t a n c e s s o many d a t a h a v e b e e n 'lumped' t h a t t h e r e s u l t s a r e o f l i t t l e value  i n interpreting  interactions. annual to  dietary  the signficance  I n an a r t i c l e  animal  b y H a n s e n e__ a i . , ( 1 9 7 7 ) , t h e  overlap f o r horses  b e 77 p e r c e n t , b u t f e c a l  of potential  a n d c o w s was  material  determined  was c o l l e c t e d  without  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 preference other in  than  diet  or apparent i tbeing  age ( i . e . d e t e r i o r a t i o n )  * u n d e c o m p o s e d . ' The 77 p e r c e n t  c h o i c e may b e v a l i d ,  of feces similarity  but i t i s only a small part of  - 88 -  the be  whole  picture,  other  date,  most d i e t  between h e r b i v o r e s  has  studies involving  s p e c i e s have u t i l i z e d  what percentage  of  been d i r e c t l y  Hansen,  1976;  O h m a r t , 1981;  The  similarity common  (Anthony study,  a pair  H a n s e n et_ a l . ,  and  Smith,  1977;  a similarity shared I feel  index  by  on by  placed  use  of  and  bias  The  most c o n f u s i n g a s p e c t  i t towards misuse of  the  same way,  and  or e s s e n t i a l l y  Hansen,  1977).  percentage  present  In the  of the dominant  n e a r l y 70  overlap, a  tend  concept  i n the  use  percentage  to obscure  concept  of  percent  restrictions  thinking  competition.  of d i e t  overlap i s  of  particular  the  same p l a c e and  i t altogether.  i n t h e way indicates  h o r s e s , cows and moose u t i l i z e t h e i r the  need  f o r a comprehensive  i n v e s t i g a t i o n of herbivore  interactions:  - 89  -  at  the  compo-  'sameness,'  R e s u l t s f r o m my  s u m m a r i z e d b e l o w , show t h a t d i f f e r e n c e s as w e l l a s exist  use  1956).  can g r e a t l y a l t e r the degree of  negate  and  Seegmiller  same t i m e , w h e r e a s i n r e a l i t y , t h e s e a r e a l l i n d e p e n d e n t nents, each of which  this  species  of d i e t  the  indicate  (Hubbard  rather stringent  of  and  o r more a n i m a l  c o w s was  p l a n t s p e c i e s may  s p e c i e s i n the  overlap  summing t h e  i t i m p l i e s 'sameness,' i . e . the  plant  identical,  O l s e n and  two  and  interpretation  identical  index' to  f o r t h e a v e r a g e use  unless  are  and  may  horses  C l a r k , 1977;  Oosting,  horses  that,  diet  1977;  i s based forages  on  as  was  H a n s e n and  shared  (69.7%).  diets  interpreted  index  feral  'similarity  of  of  forages  that  a  S a l t e r , 1978;  and  11  interactions  subtle. To  in  as  study  similarities environment,  approach  to  the  1)  I t was in  determined  Closed  Forested Closed  Zone  Forest  of the study  site  was  ( 1 4 . 5 % ) , Semi-open F o r e s t  (33.4%),  ( 1 3 . 7 % ) , Meadow ( 2 8 . 1 % ) , S h r u b c a r r  (2.6%),  (5.7%) and  and  'Other'  (1.9%).  I n t e r f a c e Zone, c o m p r i s e d  a total  of  Habitat,  as  of the area.  a p p e a r e d t o be  78 p e r c e n t the  s q km  types  h a b i t a t , i n c l u d i n g Open F o r e s t , S e m i - o p e n F o r e s t ,  67.3 p e r c e n t Moose  200  Open F o r e s t  Forest  Interface  3)  the d i s t r i b u t i o n of habitat  the approximately  as f o l l o w s :  2)  that  attracted to Forest  of a l l f e c a l counts  highest  ( 7 6 % ) , were  fecal  counts  were made i n f o r e s t , whereas  f o r horses  made i n o p e n h a b i t a t  (65.3%) and  cows  ( i . e . meadow and  shrub  carr). 4)  Cows u s e d Meadow a n d more and  5)  than  i t was  available,  moose f o r Open and C l o s e d  On  the basis  to  use c o n t i g u o u s  with to  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  of f e c a l  use I n t e r f a c e Zones  Semi-open and, 7)  found  were  found  f o r e s t ( i . e . f o r e s t not i n conjunction  open h a b i t a t s ) more t h a n  Moose were  f o r Meadow  F o r e s t , and S h r u b c a r r .  c o n c e n t r a t i o n s , horses  o p e n h a b i t a t s ) more t h a n 6)  as d i d h o r s e s  tending  ( i . e . f o r e s t i n conjunction  with  horses.  t o use those  and C l o s e d  cows, the l a t t e r  h a b i t a t types  f o r e s t and  p a r t i c u l a r l y , cows u s e d  A l l t h r e e h e r b i v o r e s showed  ( i . e . Open,  Shrub carr) that  horses  least. some s p e c i e s s p e c i f i c  - 90 -  selection  for the available 8)  The f i v e  habitats.  transects  a l l varied  significantly  t o t h e r e l a t i v e amounts o f e a c h h a b i t a t 9)  The h i g h e s t made  f e c a l counts  w h i c h had t h e h i g h e s t  type.  f o r each h e r b i v o r e  on d i f f e r e n t t r a n s e c t s .  i n regard  Cows u s e d  species  T - I V t h e most  p e r c e n t a g e o f b o t h 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 of  contiguous  had  were  amount  f o r e s t , a n d moose u s e d T-V t h e m o s t , w h i c h  the highest  moose u s e d more  amount than  o f Shrub c a r r , a h a b i t a t  type  i t s p e r c e n t a v a i l a b i l i t y would have  indicated. 10)  Although for  f e c a l counts  Meadow h a b i t a t  that  forage  i n d i c a t e d h o r s e s and cows  most s t r o n g l y ,  availability  plays  there  a major  selected  was no e v i d e n c e role  i n habitat  s e l e c t i o n , a s t h e d o m i n a n t f o r a g e i n b o t h h o r s e and c a t t l e diets  i s a forest  meadow h a b i t a t 11)  species  i n the study  B a s e d on f o o d h a b i t s forage in  d i d n o t seem t o p l a y a d o m i n a n t r o l e  s e l e c t i o n , as t h e i r highest  w e r e made i n c l o s e d 12)  area.  e s t a b l i s h e d f o r moose i n t h e l i t e r a t u r e ,  availability  t h e i r habitat  (C. r u b e s c e n s ) , n o t f o u n d i n  f o r e s t , with  a low food  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  moose w e r e s e e n  were n e i t h e r  availability.  were  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 , and  f e c a l counts  evaluated,  while  cattle  t o b e s e p a r a t e , a n d h o r s e s and moose  associated  nor separate. - 91 -  13)  H o w e v e r , t h e f i v e t r a n s e c t g r a p h s showed t h a t t h e c o n c e n tration  o f use i n any g i v e n  area  varied  w i d e l y between  species. 14)  Based  on a p p r o x i m a t e l y  cattle to  populations  be  equal  f o r the study  90% of the t o t a l  population;  average d e f e c a t i o n  and h o r s e s  horse,  cattle,  Despite  the large difference  lations, habitat  and  instances  A  trend  (i.e.  was  17)  A  to  numerous  t o use  'isolated'  areas  much m o r e t h a n  travel  routes  were d i s t r i b u t e d  meadows  areas)  cattle,  linking  and which  meadows. f o r horses  differently  over the  area. and ' s h o r t - t e r m ' c o m p a r i s o n  made o n T-V,  statistically  different.  comparison  patterns  of the three  time,  of fecal material  and two g r a p h s made, w h i c h  graphical  over  popu-  o f meadow  showed  despite the association indicated they  'long-term'  be  amount  by e x t e n s i v e f o r e s t  t o use h a b i t u a l  cattle,  was  limited  horse  and v i c e v e r s a .  f o r horses  forest  Subsequently,  study  seen  surrounded  contiguous  and  herbivore  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  those  tended  moose  respectively.  t h e t r a n s e c t graphs  c a t t l e used v e r y s l i g h t l y , 16)  estimated  i n cow v e r s u s  the relatively  (28.1%),  were  a n d moose w e r e 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% a n d 2.9%, 15)  area  rates,  were  However, a l t h o u g h  indicated that overall  - 92 -  the  distribution  s p e c i e s were somewhat  t h e r e were c e r t a i n  shown  different  consistencies i n habitat  preference,  i n c l u d i n g t h e p r o n o u n c e d u s e o f T-V b y moose;  t h e u s e b y h o r s e s , b u t o n l y s l i g h t l y by c a t t l e , o f ' i s o l a t e d ' meadows; and t h e 'absence' o f c a t t l e  i n contiguous  forest  areas. 18)  Diet  a n a l y s i s i n d i c a t e d that both  from  June  high  19)  degree  through  of s e l e c t i v i t y  forages  cattle,  of the t o t a l  7.4%  study  area.  comprised  o f t h e 148  of horses  i n their  choice  86% f o r h o r s e s ,  diet.  D i f f e r e n c e s were found patterns  and  horses,  September, were e x e r c i s i n g a  Eleven  only  cattle  plant  T h e s e 11 f o r a g e s species  to exist  of  fairly  forages.  and 80% f o r represented  identified  on t h e  i n the apparent f o r a g i n g  and cows on t h e b a s i s o f t h e 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 in and  t h e w a y s a f o r a g e p l a n t was u s e d b y h o r s e s  and c a t t l e ;  w h e t h e r 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  context. 20)  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 g r o u p s by  horses  in  their  and 21)  and  cattle  utilization  'other'  showed a s i g n i f i c a n t p e r month o f g r a s s e s ,  ( i . e . mainly  I n summer d i e t s , h o r s e s  difference rush-sedges  f o r b s and b r o w s e ) . utilized  an a v e r a g e o f 61% g r a s s e s ,  w h e r e a s cows u s e d a n a v e r a g e o f 5 1 % g r a s s e s , w i t h a h i g h e r proportion  of sedges, forbs  w e r e somewhat more s e l e c t i v e content. - 93 -  and browse, i n d i c a t i n g  cows  f o r forage w i t h lower  fiber  22)  In winter, horses and  forb species  decreased eaten,  t h e number o f g r a s s ,  and i n c r e a s e d t h e i r  sedge  utilization  o f s h r u b m a t e r i a l up t o 1 4 . 7 % . 23)  Also,  i n winter, horses  Rush-sedges, to 24)  from  increased  33.2% average  their  dependence on  i n t h e summer  months,  48% i n w i n t e r .  Because horses inner  were c o n s i d e r e d  m a t e r i a l found  able t o e x p l o i t the green,  i n sedge p l a n t s during the winter,  a n u t r i e n t a n a l y s i s was done on t h e s e l e c t e d g r e e n m a t e r i a l . This  was  found  consistently types  t o have a Crude P r o t e i n l e v e l  higher  than  f o u n d on t h e s t u d y  collected  i n late  CP v a l u e s  site,  fall  Although heavily horses  horses  f o r grass  i n a 'weathered' c o n d i t i o n ,  or winter.  dropped d r a m a t i c a l l y i n the w i n t e r 25)  reported  o f 6.5%,  Horse use o f g r a s s e s (from 60% t o 3 5 % ) .  u t i l i z e C a r e x a q u a t i l i s and C a r e x r o s t a t a  i n the winter  (28.5%) which  are both  grazed  and cows d u r i n g t h e summer, t h e s i n g l e most  winter forage p l a n t f o r horses 17.2%, and used  only  important  was Juncus b a l t i c u s ,  v e r y m o d e r a t e l y by e i t h e r  by  utilized  herbivore  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 u s e o f c o n i f e r needles  during the winter  (6.3%).  T h i s was n o t 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 o f horses. 27)  The a  total  fraction  diets  of horses  of the plant  and c a t t l e  represented  species comprising  - 94 -  only  the botanical  groups only  found  on t h e s t u d y s i t e .  6 o f t h e 13 a v a i l a b l e  forbs,  horses  used  s p e c i e s , h o r s e s used 28)  The  dominant  rubescens. used  only  utilized £. 29)  Horses  rush-sedge  16 a n d c a t t l e  and cows species;  used o f 88  2 0 , a n d o f 43 g r a s s  22 a n d c a t t l e 1 9 .  forage f o r both horses  or pinegrass, slightly  less  C_. r u b e s c e n s  and c a t t l e  although Carex by c a t t l e .  16% more t h a n  was  aquatilis  Horses, the next  £. was  however, favorite,  aquatilis.  Horses from  always  June  used  through  m o r e C. a q u a t i l i s by c a t t l e  more  rubescens  September), than horses.  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The e f f e c t s o f f a l l g r a z i n g o r b u r n i n g b l u e b u n c h w h e a t g r a s s r a n g e o n f o r a g e s e l e c t i o n b y d e e r and c a t t l e in spring. C a n . J . A n i m . S c i . 60: 113-122. , A. M c L e a n a n d C. K a l n i n . 1980c. 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 grazing. C a n . J . P l a n t S c i . 60: 131-137. W i l s o n , A. D. of grazing  1969. A review of browse i n the n u t r i t i o n animals. J . Range Manage., 2 2 : 23-27.  W o o d w a r d , S. L . a n d R. D. O h m a r t . 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 ) , C h e m e h u e v i 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 List  o f p l a n t s i d e n t i f i e d on  and  1980.  1:  the study  - 107  -  s i t e during  1979  List and to  of plants identified 1980:  A total  species  level.  on t h e s t u d y s i t e d u r i n g 1979  o f 148 s p e c i e s were i d e n t i f i e d  FAMILY  GENDS S P E C I E S  1. BETULACEAE (birch family) Betula glandulosa (2 v a r i e t i e s )  at least  COMMON NAME Bog  or Scrub  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 Forget-me-not L i t h o s p e r m u m r u d e r a l e Lemon-weed  3.  CAPRIFOLIACEAE (honeysuckle family)  4. CARYOPHYLLACEAE (pink f a m i l y )  Linnaea  borealis  Symphoricarpos Cerastium Arenaria  5.  arvense lateriflora  Stellaria Lychnis  albus  longipes  drummondii  COMPOSITAE (aster family)  Erigeron  speciosus  E. p h i l a d e l p h i c u s E.  loncophyllus  E.  compositus  E.  flagellaria  Aster  eatonii  birch  Twin-flower Waxbe r ry/Snowbe r r y Field  chickweed  Blunt-leaf  sandwort  Long-stalk  starwort  Drummond c a m p i o n Large  purple  fleabane  Philadelphia  fleabane  Spear-leaved  fleabane  Cut-leaved  fleabane  Eaton's a s t e r  A.  ciliolatus  Lindley's aster  A.  conspicuous  Large  A. p a n s u s  purple aster  Tufted white aster  prairie  continued... - 108 -  A.  compestris  Antennaria  Meadow a s t e r  parvifolia Nuttall's  pussytoes  A. m i c r o p h y l l a  Rosy  A. p u l c h e r r i m a  Showy  pussytoes  Antennaria  Field  pussytoes  A. u m b r i n e l l a  Umber  pussytoes  Artemesia  Pasture  neglecta  frigida  pussytoes  wormwood/ Sagewort  Groundsel/Ragwort  Senecio streptanthifolius S. c a n u s Crepis  tectorum  Arnica  cordifolia  A. f u l g e n s  Narrow-leaved hawksbeard Heart-leaved  arnica  Twin a r n i c a  A. c h a m i s s o n i s v a r . incana Achillea  millefolium  Yarrow  Solidago  canadensis  Canada/Meadow goldenrod  S. s p a t h u l a t a Agoseris  glauca  Cirsium sp. Taraxacum  officinale  Tragopogon p r a t e n s i s - 109 -  Spike-like Pale  goldenrod  agoseris  Thistle Common d a n d e l i o n Goatsbeard/Salsify continued...  Appendix 1 (cont.) Hieracium s c o u l e r i  6.  CRASSULACEAE (stonecrop family)  7. CRUCIFERAE (mustard family)  Grindelia sguarrosa  C u r l y - g u p gumweed  Sedum l a n c e o l a t u m  Lance-leaved stonecrop  Descurainia richardsonii Lepidium  virginicum  Arabis sp. 8.  COPRESSACEAE (juniper family)  9.  CYPERACEAE  Hounds-tongue  J u n i p e r u s communis  (sedge f a m i l y ) Carex  aquatilis  C.  rostrata  C.  sitchensis  C.  praegracilis  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.  lasiocarpa  C. a u r e a C. p a r r y a n a C.  atherodes  C.  pachystachya  Eleocharis sp.  Tansy mustard Tall  peppergrass  Rockcress Common j u n i p e r  Water  sedge  Beaked Sitka  sedge sedge  Clustered f i e l d Liddon's  sedge  N. w e s t e r n  sedge  Low N. s e d g e Slender  sedge  Golden sedge Parry  sedge  Awned s e d g e Thick-headed Spike  sedge  rush  continued... - 110 -  sedge  Appendix 1  (cont.)  10. ELAEAGNACEAE (oleaster family) 11.  Shepherdia canadensis  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.  12. GENTIANACEAE (gentian family) 13.  Buffalo berry  (monensis) uniflora  Greenish  wintergreen  Woodnymph  P. a s a r i f o l i a  Common p i n k  Arctostaphylos uva-ursi  Kinnikinnick/Bearberry  Gentiana  N. g e n t i a n  amarella  GRAMINAE (grass family) Oryzopsis  asperifolia  0. p u n g e n s Stipa  richardsonii  wintergreen  Ricegrass  Richardson  needlegrass  S. s p a r t e a  Porcupine  S. c o m a t a  Needle-and-thread grass  S. o c c i d e n t a l i s  W.  Beckmannia s p .  Sloughgrass  Hordeum j u b a t u m  Foxtail barley  H. b r a c h y a n t h e r u m  Meadow b a r l e y  Calamagrostis  Pinegrass  rubescens C. n e g l e c t a C.  grass  needlegrass  Reedgrass N. R e e d g r a s s  inexpansa  continued, - Ill  -  Puccinellia nuttalliana (airoides)  Nuttall  Poa  Alkali  bluegrass  Inland  bluegrass  P.  juncifolia interior  alkali  grass  P. p r a t e n s i s  Kentucky  Agropyron  Bluebunch wheatgrass  A.  spicatum  trachycaulum (3 v a r i e t i e s )  Spartina  gracilis  Koeleria  micrantha (cristata)  Muhlenbergia richardsonis Festuca saximontana (ovina) Danthonia  bluegrass  Beardless Alkali  June  Mat  wheatgrass  cordgrass  grass  muhly  Sheep  i n t e r m e d i a Timber  fescue  oatgrass  Bromus a n o m a l u s  N o d d i n g brome  Alopecurus  Short-awn  Glyceria  aequalis  borealis  N.  foxtail  mannagrass  Deschampsia caespitosa  Tufted hairgrass  Agrostis  Bentgrass  scabra  continued. - 112  -  Appendix 1  (cont.) Distichlis spicata var. s t r i c t a  14.  15.  GROSSULARIACEAE (currant family) Ribes sp.  Gooseberry  HIPPURIDACEAE (mare's t a i l family) Hippuris vulgaris  Common m a r e ' s t a i l  16. IRIDACEAE (Iris family)  Sisyrinchium angustifolium  Blue-eyed  Baltic  17.  JUNCACEAE (Rush f a m i l y )  Juncus b a l t i c u s  18.  JUNCAGINACEAE, (arrowgrass family)  Triglochin  LILIACEAE ( l i l y family)  Smilacina  19.  Desert s a l t g r a s s  rush  maritimum Seaside  stellata  grass  arrowgrass  Star-flowered Solomon's s e a l  Zigadenus venenosus Allium 20.  cernuum  LEGUMINOSAE (legume family) Astragalus A.  miser  Meadow d e a t h camus Nodding  onion  Timber m i l k  vetch  alpinus  Vicia  americana  American  Lathyrus  ochroleucus Yellow  Oxytropis  deflexa  vetch  pea  Pendent-pod  crazyweed  continued... - 113 -  Appendix 1 21.  22.  23.  24.  25.  26.  27.  (cont.)  LINACEAE (flax family)  ONAGRACEAE (eve. p r i m r o s e family) ORCHIDACEAE (orchid family)  POLEMONIACEAE (phlox f a m i l y ) POLYGONACEAE (buckwheat family)  RANUNCULACEAE (buttercup family)  ROSACEAE (rose family)  Linum perenne var. l e w i s i i  Epilobium angustifolium  Orchis  rotundifolia  Wild  blue  flax  Narrow-leaved fireweed  Round-leaved  orchid  Spiranthus romanzoffiana  Ladies-tresses  Polemonium h u m i l e  Jacob's  Eriogonum heracleoides  Parsnip-flowered eriogonum  Rumex s p .  Dock/Sorrel  Polygonum s p .  Smartweed/Knotweed  Ranunculus cymbalaria  Shore  Anemone  G l o b e anemone  multifida  ladder  buttercup  Aquilegia brevistyla  Blue  columbine  Thalictrum sp.  Meadow r u e  Geum t r i f l o r u m  O l d man's w h i s k e r s  Fragaria virginiana var. glauca  Wild  strawberry  continued... - 114 -  Appendix 1  (cont.) P o t e n t i l l a anserina  Silver  leaf  cinquefoil  P. g r a c i l i s  T a l l yellow  cinquefoil  P. h i p p i a n a  Wooly  P. p e n s y l v a n i c u s  Prairie  cinquefoil  Rosa a c i c u l a r i s  Prickly  rose  cinquefoil  Spiraea b e t u l i f o l i a Flat-topped  28.  29.  30.  Amelanchier sp.  Saskatoon  RUBIACEAE (madder f a m i l y )  Galium boreale  N.  SALICACEAE (willow family)  Salix  SANTALACEAE (sandalwood family)  spp.  Comandra  livida  C. u m b e l l a t a v a r . pallida 31.  SAXIFRAGACEAE (saxifrage family)  berry  bedstraw  Willow  Bastard  Pale  toadflax  comandra  Heuchera c y l i n d r i c a Round-leaf alum  P a r n a s s i a p a l u s t r i s N. g r a s s 32.  spirea  SCROPHULARIACEAE P e n s t e m o n (figwort family) fruticosus P. p r o c e r u s  of parnassus  Shrubby penstemon Slender  blue  C a s t i l l e j a miniata  Indian paint  Orthocarpus luteus  Yellow  penstemon brush  owl c l o v e r  continued... - 115 -  root  Appendix 1 33.  34.  (cont.)  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  VIOLACEAE (violet family)  Blue  V i o l a adunca  violet  TREES: 35.  PINACEAE (pine f a m i l y )  Picea P.  glauca  englemannii  Pinus  contorta var. latifolia  White  spruce  Englemann s p r u c e  Lodgepole  pine  Pseudotsuga m e n z i e s i i Douglas f i r 36.  SALICACEAE (willow family)  Populus tremuloides  Trembling  aspen  OTHER: LICHENS  P e l t i g e r a sp. Cladonia  sp.  - 116 -  Reindeer l i c h e n  APPENDIX H o r s e and  2:  C a t t l e D i e t s - d e t e r m i n e d by  -  117  -  fecal analysis  HORSE DIET - JUNE Diet H - I Plant  Species  Poa j u n c i f o l i a Juncus b a l t i c u s Carex a q u a t i l i s Calamagrostis rubescens S t i p a comata Carex r o s t a t a Carex a t h e r o i d e s Agropyron Hordeum j u b a t u m Carex c o n c i n o i d e s Glyceria borealis Stipa richardsonii Calamagrostis neglecta Muhlenburgia richardsonis Stipa occidentalis Oryzopsis a s p e r i f o l i a  Diet H - 2 % Diet 21.7 13.5 12.3 12.1 11.0 5.4 4.8 4.4 4.2 3.0 2.6 2.3 0.9 0.8 0.7 0.3  plant  Species  Poa j u n c i f o l i a Calamagrostis rubescens Juncus b a l t i c u s Carex r o s t r a t a Carex a q u a t i l i s S t i p a comata Carex a t h e r o i d e s Glyceria borealis Hordeum j u b a t u m Carex c o n c i n o i d e s Muhlenburgia richardsonis Stipa richarsonii Alopecurus aequalis Oryzopsis a s p e r i f o l i a Calamagrostis neglecta  - 118 -  % Diet 19.3 16.8 14.2 11.0 8.7 7.8 6.3 4.8 4.1 2.1 1.7 1.2 0.9 0.7 0.4  HORSE DIET - JUNE Diet Plant  Diet  H - 3  Species  Calamagrostis rubescens Poa j u n c i f o l i a Hordeum j u b a t u m Juncus b a l t i c u s Calamagrostis neglecta Carex a q u a t i l i s Carex r o s t a t a S t i p a comata Carex a t h e r o i d e s Rush-Sedges Stipa richardsonii Oryzopsis a s p e r i f o l i a Poa I n t e r i o r Eleocharis Agropyron Muhlenburgia richardsonis Agrostis P o t e n t i l l a hippiana  % Diet 19.5 17.4 9.9 10.6 8.4 8.1 7.3 5.9 2.7 2.0 1.5 1.2 1.2 1.0 0.7 0.6 0.5 0.5  Plant  H - 4  Species  Calamagrostis rubescens Poa j u n c i f o l i a Calamagrostis neglecta Carex a q u a t i l i s Carex r o s t a t a S t i p a comata Hordeum j u b a t u m Juncus b a l t i c u s Glyceria borealis Carex c o n c i n o i d e s Carex a t h e r o i d e s Rush-Sedges Stipa occidentalis Oryzopsis a s p e r i f o l i a Sisyrinchium Muhlenburgia richardsonis Stipa richardsonii Geum t r i f l o r u m Achillea millefolium Astragalus miser Hippuris vulgaris Aster conspicuous Spartina g r a c i l i s P o t e n t i l l a hippiana Beckmanaia s p . Taraxacum o f f i c i n a l e  - 119 -  % Diet 25.8 12.6 9.4 7.1 7.1 6.2 4.2 4.0 3.6 3.6 3.4 3.2 2.3 1.8 1.2 1.0 0.6 0.5 0.4 0.3 0.3 0.3 0.3 0.3 0.3 0.2  HORSE DIET - J U L Y Diet H - 1 Plant  Species  Diet H - 2 Plant  % Diet  Carex a q u a t i l i s 26.3 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 r u b e s c e n s 10.1 Calamagrostis neglecta 7.0 Carex r o s t a t a 5.9 Carex c o n c i n o i d e s 5.4 Juncus b a l t i c u s 4.8 S t i p a comata 4.5 Stipa occidentalis 4.1 Hordeum j u b a t u m 3.9 Glyceria borealis 3.7 Carex a t h e r o i d e s 2.8 Sedges-Rush 2.2 Juncus/Eleocharis 1.8 Rosa a c i c u l a r i s 1.2 Muhlenburgia richarsonis 1.2 Oryzopsis a s p e r f o l i a 1.1 Beckmania s p . 0.8 Unknown f o r b 0.6 Spartina g r a c i l i s 0.5 P o t e n t i l l a hippiana 0.1 Aster conspicuous 0.1  Species  Poa j u n c i f o l i a Calamagrostis rubescens Carex a q u a t i l i s Calamagrostis neglecta Juncus b a l t i c u s S t i p a comata Hordeum j u b a t u m Carex r o s t a t a Glyceria borealis Juncus/Eleocharis Stipa occidentalis Carex c o n c i n o i d e s Sedges-Rush Fragaria virginiana Carex a t h e r o i d e s Oryzopsis a s p e r i f o l i a Muhlenburgia r i c h a r s o n i s P o t e n t i l l a hippiana Astragalus miser Unknown g r a s s  - 120 -  % Diet 14. 3 12. 6 11. 2 10. 5 9. 2 8. 1 6. 8 6. 7 3. 7 3. 5 2. 4 1. 9 1. 6 1. 5 1. 2 1. 1 1. 0 0. 9 0. 6 1. 2  HORSE DIET - J U L Y Diet Plant  H - 3  Species  Calamagrostis rubescens Juncus/Eleocharis Stipa occidentalis Carex a q u a t i l i s Poa j u n c i f o l i a Carex r o s t a t a Hordeum j u b a t u m S t i p a comata Calamgrostis neglecta Spartina g r a c i l i s Glyceria borealis Stipa richardsonii Oryzopsis a s p e r i f o l i a Forb flower Taraxacum o f f i c i n a l e Muhlenburgia richardsonis  Diet % Diet 22.3 16.9 12.7 7.9 6.4 5.4 5.2 4.8 4.1 3.3 2.8 2.5 2.5 2.4 0.5 0.4  Plant  H - 4  Species  Calamagrostis rubescens Carex a q u a t i l i s Carex r o s t a t a Juncus/Eleocharis S t i p a comata Carex c o n c i n o i d e s Stipa occidentalis Carex a t h e r o i d e s Glyceria borealis Poa j u n c i f o l i a Stipa richardsonii Calamagrostis neglecta Juncus b a l t i c u s Hordeum j u b a t u m Oryzopsis a s p e r i f o l i a Achillea millefolium Rosa a c i c u l a r i s Artemesia f r i g i d a Muhlenburgia richardsonis Unknown f o r b  - 121 -  % Diet 18. 3 16. 0 10. 5 7. 7 6. 8 5. 2 5. 2 4. 4 4. 2 3. 6 3. 6 3. 3 3. 0 1. 7 1. 7 1. 4 1. 1 1. 0 0. 7 0. 6  HORSE DIET - AUGUST Diet H - 1 Plant Species Calamagrostis rubescens Carex c o n c i n o i d e s Stipa occidentalis Carex r o s t a t a 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 a q u a t i l i s Unknown G r a s s Astragalus miser Monocot F o r b Oryzopsis a s p e r i f o l i a Glyceria borealis Juncus s p . Hordeum j u b a t u m Calamagrostis neglecta Unknown F o r b Achillea millefolium Spiraea b e t u l i f o l i a  Diet H - 2 % Diet 27.6 11.9 9.7 8.9 6.9 5.8 5.6 4.6 4.3 4.1 3.0 1.9 1.3 1.0 0.9 0.8 0.7 0.7 0.3  Plant Species Calamagrostis rubescens Stipa occidentalis Carex a q u a t i l i s Carex r o s t a t a Juncus b a l t i c u s S t i p a comata Hordeum j u b a t u m Glyceria borealis Carex a t h e r o i d e s Poa j u n c i f o l i a Calamagrostis neglecta Juncus s p . Achillea millefolium Unknown F o r b  Diet H - 3 Plant Species Calamagrostis rubescens Carex r o s t a t a Juncus/Eleocharis Carex a q u a t i l i s Stipa occidentalis Poa j u n c i f o l i a S t i p a comata Hordeum j u b a t u m Juncus b a l t i c u s Juncus s p . Carex c o n c i n o i d e s Stipa richardsonis Oryzopsis a s p e r i f o l i a Astragalus miser Calamagrostis neglecta Glyceria borealis Muhlenburgia richardsonii Antennaria p a r v i f l o r a  % Diet 62.7 7.2 7.2 6.9 5.9 2.5 1.7 1.4 1.2 0.8 0.8 0.8 0.7 0.2  Diet H - 4 % Diet 46.2 8.2 7.5 6.5 6.4 5.6 3.2 2.7 2.3 1.8 1.7 1.5 1.5 1.5 1.4 1.2  Plant Species Calamagrostis rubescens Carex a q u a t i l i s Carex r o s t a t a Carex c o n c i n o i d e s Juncus b a l t i c u s Hordeum j u b a t u m Poa j u n c i f o l i a Glyceria borealis Stipa occidentalis Carex a t h e r o i d e s Calamagrostis neglecta Juncus sp. Astragalus miser Arctostaphylos uva-ursi  0.5 0.3  - 122 -  % Diet 31.8 20.2 12.9 7.3 5.9 4.6 4.0 3.3 2.8 2.7 2.0 1.2 0.7 0.6  HORSE DIET - SEPTEMBER Diet H - 1  Diet H - 2  Plant Species  % Diet  Calamagrostis rubescens Carex r o s t a t a Carex a q u a t i l i s Poa j u n c i f o l i a Carex c o n c i n o i d e s Stipa occidentalis Juncus b a l t i c u s Astragalus miser Hordeum j u b a t u m Carex a t h e r o i d e s Glyceria borealis Calamagrostis neglecta Spartina g r a c i l i s S t i p a comata Oryzopsis a s p e r i f o l i a Arnica cordifolia Rosa a c i c u l a r i s Forb Flower Monocot F o r b  47.3 13.5 9.4 7.0 6.9 4.5 2.9 1.7 1.6 1.2 0.9 0.7 0.5 0.5 0.4 0.3 0.3 0.3 0.1  Plant Species Calamagrostis rubescens Poa j u n c i f o l i a Carex a q u a t i l i s Carex r o s t a t a Stipa occidentalis Calamagrostis neglecta Juncus/Eleocharis Carex c o n c i n o i d e s Hordeum j u b a t u m Glyceria borealis Juncus sp. S t i p a comata Oryzopsis a s p e r i f o l i a Muhlenburgia richardsonis Unknown F o r b Astragalus miser Unknown G r a s s P o t e n t i l l a hippiana Achillea millefolium Sheperdia canadensis Sedum l a n c e o l a t u m Juniperus sp. Betula glandulosa Carex a t h e r o i d e s Rosa a c i c u l a r i s  - 123 -  % Dj,?t 45.3 8.1 7.2 7.2 5.4 3.7 2.9 2.4 2.3 2.3 2.1 1.7 1.2 1.1 1.0 0.8 0.8 0.8 0.7 0.7 0.6 0.6 0.4 0.4 0.3  HORSE DIET - SEPTEMBER Diet H - 3 Plant  Species  Carex r o s t a t a Calamagrostis neglecta Carex a q u a t i l i s Calamagrostis rubescens Carex c o n c i n o i d e s Poa j u n c i f o l i a Unknown S h r u b s Juncus sp. Glyceria borealis Stipa occidentalis Unknown F o r b S t i p a comata Monocot F o r b G r a s s Seed Muhlenburgia richardsonis Hordeum j u b a t u m Unknown G r a s s Carex s p . Astragalus miser V i c i a sp. Pinus contorta  Diet H - 4 Plant  % Diet 14.4 11.4 10.6 9.4 9.1 8.9 5.9 5.6 4.7 3.8 2.7 2.6 2.1 1.8 1.5 1.4 1.2 0.9 0.7 0.7 0.6  Species  Calamagrostis rubescens Stipa occidentalis Carex a q u a t i l i s Poa j u n c i f o l i a Carex r o s t a t a Juncus s p . Calamagrostis neglecta Glyceria borealis Hordeum j u b a t u m Astagalus miser Juncus/Eleocharis Muhlenburgia richardsonis S t i p a comata Eriogonum s p . Unknown G r a s s P o t e n t i l l a hippiana Carex c o n c i n o i d e s Pinus c o n t o r t a Unknown F o r b Beckmania s p . Fern Taraxacum o f f i c i n a l e Monocot f o r b V i c i a sp. Unknown s h r u b Arctostaphylos uva-ursi  - 124 -  % Diet 18.0 10.1 7.9 6.5 6.4 6.4 6.1 5.4 4.3 3.3 3.0 2.6 2.5 2.3 1.9 1.9 1.6 1.6 1.6 1.4 1.2 0.9 0.9 0.8 0.8 0.6  HORSE DIETS - WINTER Diet H - 1 Plant Species  Diet H - 2 % Diet  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 Glyceria borealis 8.3 Unknown G r a s s 6.9 Calamagrostis rubescens 6.5 Stipa occidentalis 6.0 Pinus contorta 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 Calamagrostis neglecta 2.5 Artemesia f r i g i d a 2.0 Juncus/Eleocharis 1.9 Betula glandulosa 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 Picea sitchensis 1.2 Salix sp. 1.1 Unknown s h r u b 0.5 Fern 0.3  Plant  Species  Juncus s p . Carex r o s t a t a Carex a q u a t i l i s Unknown G r a s s S t i p a comata Calamagrostis rubescens Unknown S h r u b Pinus contorta Glyceria borealis Stipa occidentalis Calamagrostis neglecta Picea sitchensis Rosa a c i c u l a r i s Fern Antennaria p a r v i f l o r a Carex c o n c i n o i d e s Poa j u n c i f o l i a Arctostaphylos uva-ursi  - 125 -  % Diet 18.8 14.4 10.6 9.4 7.9 7.5 5.3 4.3 4.0 4.0 3.1 2.7 2.6 2.2 1.0 0.8 0.7 0.7  CATTLE DIETS - JUNE Diet Plant  C - 1  Species  Diet % Diet  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 r u b e s c e n s 10.1 Rush-Sedges 9.8 Calamagrostis neglecta 9.7 Hordeum j u b a t u m 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 Glyceria borealis 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 Muhlenburgia richardsonis 2.0 Hippuris vulgaris 1.7 Stipa richardsonii 1.3 Solidago spathulata 1.0 V i o l a adunca 0.9 Oryzopsis a s p e r i f o l i a 0.9 Fragaria virginiana 0.7 P o t e n t i l l a hippiana 0.6 Stipa occidentalis 0.4 Astragalus miser 0.4 Antennaria p a r v i f o l i a 0.3 Aster conspicuous 0.3 Achillea millefolium 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  Plant  C - 2  Species  Poa j u n c i f o l i a Carex a q u a t i l i s Calamagrostis rubescens Carex c o n c i n o i d e s Rush-Sedges Carex r o s t a t a Hordeum j u b a t u m Calamagrostis neglecta S t i p a comata Glyceria borealis Juncus b a l t i c u s P o t e n t i l i a hippiana Spartina g r a c i l i s Carex a t h e r o i d e s Stipa occidentalis Oryzopsis a s p e r i f o l i a Stipa richardsonii Muhlenburgia richardsonis Fragaria virginiana Geum t r i f l o r u m Solidago spathulata Aster conspicuous Antennaria p a r v i f o l i a Astragalus miser  - 126 -  % Diet 13.9 13.2 12.9 12.6 9.0 6.3 5.3 4.3 4.0 4.0 3.9 2.8 1.4 1.3 1.1 1.0 0.9 0.7 0.6 0.2 0.2 0.1 0.1 0.1  CATTLE D I E T S - JUNE Diet C - 3 Plant Species Calamagrostis rubescens Carex a q u a t i l i s Carex r o s t a t a Poa j u n c i f o l i a Hordeum j u b a t u m Calamagrostis neglecta Glyceria borealis S t i p a comata Carex c o n c i n o i d e s Stipa occidentalis Juncus b a l t i c u s Achillea millefolium Potentilla hippiana Sedge-Rush Stipa richardsonii Carex a t h e r o i d e s Geum t r i f l o r u m Agropyron trachycaulum Cerastium arvense Astragalus miser Fragaria virginiana Unknown f o r b Arnica cordifolia Muhlenburgia richarsonis Spartina g r a c i l i s Taraxacum o f f i c i n a l e Solidago spathulata Heuchera c y l i n d r i c a Penstemon p r o c e r u s Sedum l a n c e o l a t u m Aster conspicuous  Diet C - 4 % Diet 14.6 12.7 9.8 9.7 7.5 8.5 5.5 5.0 3.9 3.8 3.3 2.4 2.4 1.7 1.5 1.3 0.8 0.7 0.6 0.6 0.5 0.5 0.4  Plant  Species  Carex a q u a t i l i s Carex r o s t a t a Carex a t h e r o i d e s Calamagrostis neglecta Hoedeum j u b a t u m Poa j u n c i f o l i a Carex c o n c i n o i d e s Calamagrostis rubescens Juncus b a l t i c u s S t i p a comata Glyceria borealis Stipa occidentalis Sedge-Rush Spartina g r a c i l i s Sisyrinchium angustifolium Arnica cordifolia Muhlenburgia richardsonis Astragalus miser Aster conspicuous P o t e n t i l l a hippiana  0.4 0.3 0.3 0.3 0.3 0.3 0.3 0.1  - 127 -  % Diet 14. 7 11. 2 9. 6 9. 5 8. 5 8. 1 8. 0 7. 4 6. 1 4. 8 3. 6 2. 4 2. 4 1. 0 0. 7 0. 7 0. 5 0. 4 0. 3 0. 1  CATTLE DIETS - J U L Y Diet Plant  Diet  q - l  Species  Carex a q u a t i l i s Sedge-Rush Calamagrostis neglecta Calamagrostis rubescens Glyceria borealis Hordeum j u b a t u m S t i p a comata Carex r o s t a t a Stipa occidentalis Carex c o n c i n o i d e s Poa j u n c i f o l i a Juncus b a l t i c u s Rosa a c i c u l a r i s Potentilla hippiana Astragalus miser Achillea millefolium Carex a t h e r i o d e s Arnica c o r d i f o l i a Salix sp. Unknown f o r b Geum t r i f l o r u m Fragaria virginiana Spartina g r a c i l i s  %  Diet 15.8 10.8 10.8 10.0 8.4 6.3 5.7 5.1 4.9 4.3 4.2 2.8 2.4 1.6 1.2 1.0 1.0 0.8 0.7 0.7 0.6 0.6 0.3  Plant  q - 2  Species  qarex a q u a t i l i s Glyceria borealis Calamagrostis rubescens Stipa occidentalis Sedge-Rush S t i p a comata Calamagrostis neglecta Hordeum j u b a t u m Juncus b a l t i c u s Carex r o s t a t a Poa j u n c i f o l i a Carex c o n c i n o i d e s Carex a q u a t i l i s Muhlenburgia richardsonis Oryzopsis a s p e r i f o l i a Mosses Potentilla hippiana Rosa a c i c u l a r i s Sedum l a n c e o l a t u m Stipa richardsonii Spartina g r a c i l i s Arctostaphylos uva-ursi Penstemon p r o c e r u s Antennaria p a r v i f o l i a Aster conspicuous Taraxacum o f f i c i n a l e Betula glandulosa Achillea millefolium Salix sp. Unknown f o r b Geum t r i f l o r u m  - 128 -  %  Diet 21.9 9.2 8.5 7.8 7.5 7.4 5.5 4.8 4.2 4.1 3.3 2.3 2.0 1.9 1.7 1.1 0.8 0.8 0.8 0.7 0.7 0.7 0.5 0.5 0.2 0.2 0.2 0.2 0.2 0.2 0.1  CATTLE DIETS - JULY Diet C - 3 Plant  Species  Diet C - 4 % Diet  20.9 Carex a q u a t i l i s Carex r o s t a t a 14.3 C a l a m a g r o s t i s rubescens 10.0 7.9 Hordeum jubatum 7.6 Calamagrostis neglecta Glyceria borealis 6.8 5.8 Rush-Sedge Stipa occidentalis 4.5 Poa j u n c i f o l i a 3.8 S t i p a comata 3.5 Muhlenburgia richardsonis 2.7 2.0 Juncus b a l t i c u s Carex c o n c i n o i d e s 1.6 Oryzopsis a s p e r i f o l i a 1.6 Stipa richardsonii 1.4 P o t e n t i l l a hippiana 1.0 0.9 Carex a t h e r o i d e s 0.8 Mosses 0.6 Unknown f o r b Antennaria p a r v i f o l i a 0.5 Achillea millefolium 0.3 Rosa a c i c u l a r i s 0.3 Fragaria virginiana 0.2 A s t e r conspicuous 0.2 Arnica c o r d i f o l i a 0.2 A r c t i s t a p h y l o s u v a - u r s i 0.6  Plant  Species  Carex a q u a t i l i s Carex r o s t a t a Calamagrostis n e g l e c t a Calamagrostis rubescens Hordeum jubatum Stipa occidentalis Glyceria borealis Carex a t h e r o i d e s S t i p a comata Carex c o n c i n o i d e s Juncus b a l t i c u s Achillea millefolium Muhlenburgia richardsonis P o t e n t i l l a hippiana A s t e r conspicuous Arctostaphylos uva-ursi Taraxacum o f f i c i n a l e A s t r a g a l u s miser S a l i x sp. Fragaria virginiana Antennaria p a r v i f o l i a  - 129 -  % Diet 21.2 13.1 13.0 11.8 9.5 9.0 8.7 4.5 2.9 1.1 0.8 0.8 0.6 0.5 0.5 0.3 0.3 0.3 0.2 0.2 0.2  CATTLE DIET - AUGUST Diet C - X Plant  Species  Carex a q u a t i l i s Glyceria borealis S t i p a comata Calamgrostis 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 Calamgrostis neglecta Carex c o n c i n o i d e s Juncus/Eleocharis Hordeum j u b a t u m Unknown f o r b Stipa occidentalis Stipa richardsonii P o t e n t i l l a hippiana Oryzopsis a s p e r i f o l i a Salix sp. Astragalus miser Shepherdia canadensis Geum t r i f l o r u m Rosa a c i c u l a r i s Aster conspicuous Achillea millefolium  Diet  Plant  % Diet 16.2 10.8 10.0 9.3 8.7 8.1 7.3 5.9 4.9 3.8 2.9 2.7 2.0 1.6 1.3 0.8 0.7 0.7 0.6 0.5 0.4 0.4 0.2 0.2  C -  2  Species  Calamgrostis rubscens Carex a q u a t i l i s Poa j u n c i f o l i a S t i p a comata Stipa occidentalis Carex r o s t a t a Carex c o n c i n o i d e s Juncus/Eleocharis Hordeum j u b a t u m Juncus b a l t i c u s Shepherdia canadensis Calamagrostis neglecta Oryzopsis a s p e r i f o l i a Rosa a c i c u l a r i s Sisyrinchium angustifolium Muhlenburgia richardsonis V i c i a americana Stipa richardsonii P o t e n t i l l a hippiana Dicot forb Geum t r i f l o r u m Salix sp. Fragaria virginiana Thalictrum sp. Monocot f o r b Antenaria sp. Spiraea b e t u l i f o l i a Arctostaphylos uva-ursi Glyceria borealis Betula Aster consipcuous Achillea millefolium Solidago spathulata P o t e n t i l l a hippiana Moss  - 130 -  % Diet 25.1 16.6 5.6 5.0 4.9 3.5 3.5 3.4 3.2 2.7 2.6 2.2 2.2 1.7 1.7 1.4 1.4 1.3 1.3 1.2 1.2 1.2 1.1 1.1 1.0 0.7 0.7 0.5 0.5 0.3 0.3 0.2 0.2 0.2 0.1  CATTLE DIET - AUGUST Diet Plant  )  C - 3  Specie?  Calamagrostis rubscens Poa juncifolia Carex r o s t a t a Carex c o n c i n o i d e s Stipa occidentalis S t i p a comata Rosa a c i c u l a r i s Carex a q u a t i l i s Juncus b a l t i c u s Glyceria borealis Shepherdia canadensis Muhlenburgia richardsonis Oryzopsis a s p e r i f o l i a P o t e n t i l l a hippiana Stipa richardsonii Calamagrostis neglecta Geum t r i f l o r u m Unknown f o r b Juncus/Eleocharis Beckmania Achillea millefolium Hordeum j u b a t u m Astragalus miser Fragaria virginiana Solidago spathulata  %  Diet Plant  Diet 28. 1 11. 6 10. 8 8. 2 6. 5 5. 2 3. 4 3. 3 2. 9 2. 7 2. 6 2. 4 1. 7 1. 6 1. 3 1. 2 1. 2 1. 2 0. 8 0. 7 0. 6 0. 6 0. 6 0. 4 0. 3  C - 4  Species  Calamagrostis rubscens Unknown f o r m Carex a q u a t i l i s Stipa occidentalis Juncus b a l t i c u s Carex r o s t a t a Calamagrostis neglecta Poa j u n c i f o l i a S t i p a comata Achillea millefolium Carex c o n c i n o i d e s Sisyrinchium angustifolium Hordeum j u b a t u m Spiraea b e t u l i f o l i a Pinus contorta v a r . latifolia Taraxacum o f f i c i n a l e Muhlenburgia richardsonis Oryzopsis a s p e r i f o l i a Forb flower P o t e n t i l l a hippiana Arctostaphylos uva-ursi V i c i a americana Salix sp. Fragaria virginiana Glyceria borealis Shepherdia canadensis Geum t r i f l o r u m Astragalus miser  •  131  -  %  Diet 22.2 15.0 9.0 6.3 5.8 4.5 4.5 3.0 2.8 2.7 2.3 2.2 2.1 2.0 1.8 1.7 1.4 1.4 1.3 1.3 1.1 1.1 1.0 1.0 1.0 0.9 0.4 0.2  CATTLE DIET - SEPTEMBER Diet Plant  C - 1  Species  Calamagrostis rubescens Carex r o s t a t a Astragalus miser Carex a q u a t i l i s Glyceria borealis Stipa occidentalis Oryzopsis a s p e r i f o l i a Juncus b a l t i c u s Achillea millefolium Hordeum j u b t a u m Carex c o n n c i n o i d e s S t i p a comata Vicia sp. Beckmannia s p . Muhlenburgia richardsonis Cerastium arvense Taraxacum o f f i c i n a l e Sisyrinchium angustifolium Fragaria virginiana V i o l a adunca Poa j u n c i f o l i a Geum t r i f l o r u m Salix sp.  Diet % Diet 16.5 9.6 8.4 8.0 7.7 7.2 6.4 5.5 4.9 4.9 4.3 3.3 2.2 1.7 1.7 1.5 1.3 1.2 1.0 1.0 1.0 0.5 0.2  Plant  C - 2  Species  Calamagrostis rubescens Carex a q u a t i l i s Juncus b a l t i c u s Poa j u n c i f o l i a S t i p a comata Stipa occidentalis Carex r o s t a t a Glyceria borealis Hordeum j u b a t u m V i c i a sp. Stipa richardsonii Fragaria virginiana P o t e n t i l l a hippiana Sheperdia canadensis V i o l a adunca Oryzopsis a s p e r i f o l i a Astragalus miser Spiraea b e t u l i f o l i a Beckmannia s p . Achillea millefolum Pinus contorta Carex c o n c i n o i d e s Muhlenburgia richardsonis Arnica c o r d i f o l i a Calamagrostis neglecta Geum t r i f l o r u m  - 132 -  % Diet 29.0 15.6 7.1 6.0 5.3 5.2 4.4 3.4 3.9 3.1 2.4 2.0 1.6 1.5 1.2 1.1 0.9 0.9 0.9 0.8 0.8 0.8 0.7 0.5 0.5  CATTLE DIET - SEPTEMBER Diet C - 4  Diet C - 3 Plant  Species  % Diet  29. 6 Carex a q u a t i l i s Calamagrostis rubescens 25. 5 Carex c o n c i n o i d e s 7. 6 Glyceria borealis 4. 1 Juncus b a l t i c u s 3. 7 S t i p a comata 3. 6 Stipa occidentalis 3. 5 2. 8 Hordeum j u b a t u m Calamagrostis neglecta 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 sp. 1. 9 Carex r o s t a t a 1. 7 S a l i x sp. 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 Spartina g r a c i l i s 1. 1 Arnica cordifolia 1. 0 Oryzopsis a s p e r i f o l i a 0. 8 Sheperdia canadensis 0. 6 Juncus/Eleocharis 0. 6 Potentilla hippiana 0. 4  Plant  Species  Carex a q u a t i l i s Calamagrostis rubescens Carex r o s t a t a Hordeum j u b a t u m S t i p a comata Juncus b a l t i c u s Poa j u n c i f o l i a Stipa occidentalis Salix sp. Carex c o n c i n o i d e s Unknown f o r b Pinus contorta Unknown G r a s s Sheperdia canadensis Glyceria borealis Astragalus miser Calamagrostis neglecta  - 133 -  % Diet 31.8 23.5 5.8 5.1 5.0 4.6 3.9 3.6 2.9 2.6 2.5 2.1 2.0 1.8 1.0 1.0 0.8  APPENDIX Paired  ' t ' test  c o u n t s on T r a n s e c t  3:  f o r 'Long-term' V  - 134  -  versus  'Short-term'  fecal  Cows v e r s u s Cows Data  f o rPaired  ' t ' test  (with  t i o n - ( S o k a l a n d R o h l f , 1969: (old)  versus  'Short-term'  T r a n s e c t V i n .5 km  No. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33.  F e c a l Counts Old 286 289 425 416 231 259 292 384 252 224 234 206 165 175 165 181 33 24 10 4 1 14 10 1 4 3 2 13 8 20 167 100 38  36 d . f . ) - A r c s i n  Transforma-  205) f o r comparison o f 'Long-term' (new) f e c a l  counts  recorded  on  units:  Prop, o f Tot Transect-Old  No.  5.4 5.4 7.9 7.8 4.3 4.9 5.5 7.2 4.7 4.2 4.4 3.9 3.1 3.3 3.1 3.4 .62 .5 .2 .1 .02 .3 .2 .02 .1 .1 .04 .2 .2 .4 3.1 1.9 .7  1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33.  F e c a l Counts New 116 116 177 164 110 129 102 33 68 61 76 9 5 20 61 9 1 0 0 0 0 0 0 0 0 0 1 4 1 0 8 16 0  Prop, ofT o t . Transect-New 7.7 7.7 11.8 10.9 7.3 8.6 6.8 .1 4.5 4.1 5.1 .6 .3 1.3 4.1 .6 .1 0 0 0 0 . 0 0 0 0 0 .1 .23 .1 0 .5 1.1 0  continued... - 135 -  A p p e n d i x 3 - Cows v e r s u s Cows 34. 35. 36. 37.  41 94 287 278 5336  .8 1.8 5.4 5.2  d = 55.38 d =  1.497  ( d ) d2  (cont.) 34. 35. 36. 37.  2  2 15 38 163 1505  = 3066.94 =  622.23  - 136 -  t » 2.35  .1 1.0 2.5 10.8  APPENDIX  No. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37.  Fecal  Counts Old  Horses versus  Horses  Prop, of Tot. F e c a l Counts Transect-Old No. New  23 19 33 23 15 15 9 29 61 49 16 164 129 134 245 102 40 30 22 3 0 4 220 32 96 14 5 54 27 12 37 28 20 20 25 56 46 1857  1.2 1.0 1.8 1.2 .8 .8 .49 1.6 3.3 2.6 .86 8.8 6.9 7.2 13.2 • 5.5 2.2 1.6 1.2 .2 0 .2 11.9 1.7 5.2 .8 .27 2.9 1.5 .65 2.0 1.5 1.1 1.1 1.4 3.0 2.5  d = 105 .8 d =  3:  2.859  1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37.  ( d ) 2 = 11,193.6 d  2  =  2036.3 t = 2.51  - 137 -  Prop, o f Transect  0 0 1 0 0 0 0 1 0 2 5 0 0 10 3 3 0 0 0 0 0 0 7 4 3 0 0 0 0 1 0 0 0 0 0 1 _1 44  0 0 2.3 0 0 0 0 2.3 0 4.6 11.4 0 0 22.7 6.8 6.8 0 0 0 0 0 0 15.9 9.1 6.8 0 0 0 0 2.3 0 0 0 0 0 2.3 6.8 d . f . = 36 ArcSin »  % = x  InvSin x = ArcSin  APPENDIX 3: Fecal No. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37.  Counts Old  Moose  Prop, of Tot• F e c a l Counts Transect-Old NO. New  0 2 0 0 0 1 3 6 7 7 6 8 9 15 95 52 30 5 15 6 14 10 16 34 13 25 26 32 15 33 8 8 3 14 30 68 _ l i 647  0 0 0 0  1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37.  .31  .2 .46 .93 1.1 1.1 .93 1.2 1.4 2.3 14.7 8.0 4.6 .8 2.3 .93 2.2 1.5 2.5 5.3 2.0 3.9 4.0 4.9 2.3 5.1 1.2 1.2 .46 2.2 4.6 10.5  d = 9 1 .4 d =  Moose v e r s u s  2.469  ( d) d2  2  = 8 3 5 3 . 96 = 1358.3  - 138 -  Prop, of Transect  0 0 0 0 0 0 0 0 0 0 1 0 0 0 14 4 4 2 5 2 1 0 0 2 5 6 0 6 0 1 0 0 2 0 0 4 62 t = 2.675  0 0 0 0 0 0 0 0 0 0 1.6 0 0 0 22.6 6.5 6.5 3.2 8.1 3.2 1.6 0 0 3.2 8.1 9.7 0 9.7 0 1.6 0 0 3.2 0 0 6.5 4.8  APPENDIX Presence of p l a n t species o r i g i n a l 10  4: identified  habitat types  - 139  designated  -  in  the  present i n one or more of the o r i g i n a l h a b i t a t types Plant species  2.5)  (Present = x)  plant  O r i g i n a l H a b i t a t Type  Species  Grasses-Rushes-Sedges Calamagrostis  (Section  rubescens  3  4  5  6  7  8  x  X  O r y z o p s i s pungens  x  X  Koeleria  micrantha  X  X  Carex  concinnoides  X  X  Stipa  richardsonii  X  X  X  X  F e s t u c a saximontana  X  X  X  X  Poa  X  X  X  X  X  X  X  X  interior  Agropyron  trachycaulum  Oryzopsis  asperifolia  Poa spp.  (juncifolia & pratensis)  X  X X  X X  Bromus anomalus  X  Hordeum jubatum  X  Stipa spartea  X  Muhlenbergia  X  richardsonis  continued,  Appendix 4 Juncus Carex  (cont.)  balticus  5  6  7  8  x  x  x  x  x  X  X  X  praegracilis  Spartina  gracilis  Puccinellia Danthonia  nuttalliana  intermedia  X  Agropyron spicatum  X  Stipa  comata  X  Stipa  occidentalis  X  Carex  petasata  X  Calamagrostis  neglecta  X  Carex  aquatilus  X  Carex  rostata  X  Beckmannia sp.  X  Carex  sitchensis  X  Carex  atheroides  X  Alopecurus aegualis  X  Hordeum b r a c h y a n t h e r u m  X  Carex pachystachya  X  continued.  Appendix 4 (cont.) Deschanpsia Glyceria  caespitosa  borealis  Carex  lasiocarpa  Carex  aurea  x X  Carex p a r r y a n a Calamagrostis  X  inexpansa  X  Carex c o n c i n n a Eleocharis  sp.  ?  FORBS Linnaea  borealis  x  X  X  X  X  Astragalus miser  X  X  X  X  X  Epilopium spp.  X  X  Anemone m u l t i f i d a  X  X  Achillea  millefolium  X  X  Fragaria  virginiana  X  X  Solidago  spathulata  X X  X  Violoa  adunca  X  X  X  X  X  X  X  X  continued,  Appendix 4 Pyrola  chlorantha  Castilleja Aster  (cont.)  8 x  miniata  x  ciliolatus  Agoseris  X  spp.  Senecio  X  spp.  Polemonium humile  X  Antennaria  mycrophylla  X  Antennaria  neglecta  Geum t r i f l o r u m Lithospermum ruderale Heiracium Sedum  cynoglossydes  lanceolatum  Gentiana  amarella  Erigeron  speciosus  Galium Aster  X  boreale  X  X  X  conspicuous  X  L y c h n i s drummondii Arnica  cordifolia  X  continued.  4  Appendix  6  (cont.)  x  T r i g l o c h i n maritumum Potentilla Solidago  anserina canadensis  Taroxacum  officinales  x  Artemesia  fricida  X  Potentilla  hippiana  X  Penstemonprocerus  X  Cerastium  X  arvense  Heuchera  cylindrica  X  Zigadenus  venenosus  X  Linum perenne Aster  lewisii  campestris  Lepidium  X  Vicia  spp.  Lathyrus Allium  lutens  ochroleucus  cernum  X  X  X  X  X X  Orthocarpus  X  X  X  Eriogonum h e a c l e o i d e s virginicum  X  X X  X  X  X  X X  continued.  Appendix 4 (cont.) Arabis  1  8  9  spp.  Ranunculus spp. Aster  eatonii  Polygonum spp. Hippuris  , i—• *. Ul 1  vulgaris  Potentilla  pensylvanicus  Potentilla  gracilis  Aster  pansus  Smilacina  stellata  x X  Sisyrinchium angustifolium Comandre  livida  Pyrolaceae monensis u n i f l o r a Caryophylaceae Arenaria 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  X X X  C i r s i u m spp.  X  Antennaria  pulchrima  X  Thalictrum  occidentale  Arnica  X  chamissonis  Antennaria  parviflora  continued.  Appendix 4  (cont.)  Erigeron loncophyllus Other P l a n t s Cladonia Peltigera General Other  X  Fungus  x  lichen  Mosses  x  (general)  X  X  X  X  X  X  X  X  X  X  Trees/Shrubs Symphoricarpus Pinus  albus  X  contorta  X  Pseudotsuga m e n z i e s i i Populus  tremuloides  Juniperus Shepherdia Rosa Salix  communis canadensis  acicularis spp.  Amelanchier  spp.  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  continued,  Appendix Spirea  4  (cont.)  6  betulifolia  Arctostaphylus Picea  glauca  Ribes  spp.  uva  ursi  X  x  X  X  X  8  X  9  10  X  X  X  X X  B e t u l a g l a n d u l o s a and  Approximate  x  7  pumila  TOTAL SPECIES  X  26  34  31  58  28  43  44  37  52  APPENDIX N u t r i e n t a n a l y s i s on  5:  c o l l e c t e d sedge  material  Results  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 M a r c h 1 9 8 0 . *  Sedge M a t e r i a l  Fibre Dry TDN: Protein; Calcium; Phosphorus; A.D.F. M a t t e r A s As As As (% D r y ) (%) Fed Dry Fed Dry Fed Dry Fed Dry  1.  Ungrazed, whole sedge plant ( i . e . o l d material with green material mixed).  45.6  96.8  50  52 3.7  3.78 0.34  0.35 0.07  2.  S e l e c t e d green m a t e r i a l 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 outer growth).  41.0  96.7  55  57 6.4  6.5  0.14  * S o i l , F e e d , and T i s s u e - t e s t i n g L a b o r a t o r y ,  0.14  0.072  0.15 0.15  1873 S p a l l R o a d , K e l o w n a , B.C. V1Y 4R2  

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