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The effects of thinning on forest bird communities in dry interior Douglas-fir forests Booth, Barry P. 1994

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THE EFFECTS OF THINNING ON FOREST BIRD COMMUNITIES IN DRY INTERIOR DOUGLAS-FIR FORESTS by BARRY P. BOOTH B.Sc,  University of B r i t i s h  Columbia,  1988  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS MASTER  FOR THE DEGREE OF  OF SCIENCE  THE FACULTY OF GRADUATE  STUDIES  (Department o f F o r e s t S c i e n c e ) We a c c e p t t h i s t h e s i s as c o n f o r m i n g to the r e q u i r e d s t a n d a r d  UNIVERSITY OF BRITISH October ©  COLUMBIA  1994  B a r r y P. B o o t h ,  1994  In  presenting this  degree at the  thesis in  University of  partial  fulfilment  of  of  department  this thesis for or  by  his  or  scholarly purposes may be her  representatives.  permission.  Department of The University of British Columbia Vancouver, Canada  DE-6 (2/88)  for  an advanced  Library shall make it  agree that permission for extensive  It  publication of this thesis for financial gain shall not  Date  requirements  British Columbia, I agree that the  freely available for reference and study. I further copying  the  is  granted  by the  understood  that  head of copying  my or  be allowed without my written  Abstract I studied F o r a g e and species  P r o j e c t on  abundance i n t h e  three  was  result,  there  (<10  cm  t r e a t m e n t and  were s i g n i f i c a n t l y dbh)  lower) i n t r e a t e d  significant  increase  This  the  higher  s l a s h on  s i n c e treatment,  o r an  Elevated  t u s s o c k moth, and  forest  s t r a t a may  abundance o f  size.  l e v e l s of  as  Individual  a  tree  d i f f e r e n t between  dbh  were  hectare  significantly  T h e r e was  no  i n treatment  d e a d woody d e b r i s  was  sites. e f f e c t on  may  to  have p r e v e n t e d  the  foliage-feeding species.  the  habitat  predicted  time  failure  increase  The  of  thinning,  s p r u c e budworm and  potential foraging  forest  amount o f  explain  species  the  e i t h e r because  insufficient  insufficient  bird  thinned  Douglas-fir  shrub cover  little  or a l l f o u r ,  ground-and s h r u b - f e e d i n g predicted.  fewer s m a l l  Poor u n d e r s t o r y r e s p o n s e ,  site,  three  in  bird  I  s i z e c l a s s e s , and  cm  in treated had  1991.  e a c h 2 5 ha  sites.  e f f e c t s of c a t t l e g r a z i n g ,  leaving  fir  <10  i n h e r b and  thinning t r i a l  community.  and  Canopy v o l u m e s p e r  P e r c e n t c o v e r o f down and  significantly  bird  trees  s t r u c t u r e and  a b u n d a n c e on  significantly  control sites.  3  (60%  1990  i n treatment p l o t s .  (m /ha) o f D o u g l a s - f i r  sites.  bird  at smaller  c a n o p y v o l u m e s were n o t  lower  summers o f  s t r u c t u r e and  targeted  T h o m p s o n / N i c o l a Mule Deer  vegetation  unthinned study s i t e s ,  Thinning  trees  e f f e c t s of the  Slashing  sampled stand and  the  of  as Douglasin  other  reduction  increased  in  amount  o f down and increase  dead wood l i k e l y  i n woodpecker use  C h i - s q u a r e and that  accounted  of thinned  discriminant  f o r the  modest  sites.  function  analysis  suggested  several within-site bird/habitat associations  Northern f l i c k e r s , were a s s o c i a t e d  V e s p e r s p a r r o w s and  w i t h open f o r e s t h a b i t a t s  Yellow-rumped warblers, associated  Chipping  and  exist.  sparrows  in control  sites.  Dusky f l y c a t c h e r s were  w i t h dense, u n t h i n n e d h a b i t a t  i n treatment  sites.  Ruby-crowned k i n g l e t s , Orange-crowned w a r b l e r s and  Dusky  f l y c a t c h e r s were a s s o c i a t e d  in  control  with r i p a r i a n  sites.  I recommend t h a t g r a z i n g the be  effect piled  of g r a z i n g  i t to  l i e where i t was  left  as p a r t  of the  habitat heterogeneity.  both vegetation particularly Slashing  vegetation  modified  response.  felled.  and  i f the  bird  to  assess  Slash  species  could instead  Even i f n o t  f o r snag management, u n t h i n n e d p a t c h e s  c o n t i n u e t o be spatial  on  r e g i m e s be  t o b e n e f i t ground f o r a g i n g / n e s t i n g  of allowing required  habitats  treatment to  should maintain  Long-term m o n i t o r i n g  communities  of  i s recommended  T h o m p s o n / N i c o l a M u l e Deer F o r a g e  and  P r o j e c t becomes a model f o r t h i n n i n g p r o j e c t s  these f o r e s t  types.  in  iv  Table of Contents Abstract  i i  Table of Contents  iv  List  of Tables  vi  List  of Figures  viii  Acknowledgments  ix  1.  Introduction..  1  2.  Study s i t e s  5  3.  E f f e c t s of thinning  on t h e p l a n t  community  Methods Vegetation p l o t s . Canopy volume..  10 10 -11  Data a n a l y s i s  14  Results Stocking  10  14 density:  Canopy volume:  Large t r e e s Small t r e e s  14 18  Large t r e e s Small t r e e s  20 20  C o v e r measurements  23  Discussion Ground c o v e r T r e e canopy v o l u m e s  25 26 29  Conclusions  32  4.  E f f e c t s of t h i n n i n g community  on t h e f o r e s t  34  Methods Censusing Composite v e g e t a t i o n Data a n a l y s i s :  bird  maps  Community s t r u c t u r e R e l a t i v e abundance Bird/habitat relationships  34 34 36 37 38 39  V  Results Community s t r u c t u r e Bird  abundance:  Bird/habitat relationships:  Number o f t e r r i t o r i e s Number o f d e t e c t i o n s  43 43 45 52  C o m p o s i t e maps 52 B i r d communities 55 W i t h i n p l o t s i m i l a r i t y a n a l y s i s . . . . 64 Chi-squared a n a l y s i s 67 Discriminant function analysis 70  Discussion Community s t r u c t u r e R e l a t i v e abundance Bird/habitat relationships  76 76 77 82  5.  Conclusions  88  6.  Literature cited  92  Appendix  1  100  Appendix  II  101  Appendix  II  102  vi  List  of Tables  T a b l e 1. Number o f t r e e s sampled f o r c a n o p y volume measurements i n c o n t r o l and t r e a t m e n t s i t e s by s p e c i e s and d i a m e t e r c l a s s  13  T a b l e 2. Mean d e n s i t i e s o f l i v e t r e e s (stems/ha) > 10 cm dbh i n t r e a t m e n t and c o n t r o l s i t e s by t r e e s p e c i e s and d i a m e t e r c l a s s  15  T a b l e 3. Mean d e n s i t i e s o f s n a g s (stems/ha) > 10 cm dbh i n t r e a t m e n t a n d c o n t r o l s i t e s by t r e e s p e c i e s and d i a m e t e r c l a s s  17  T a b l e 4. S t o c k i n g d e n s i t y ( t r e e s p e r ha) o f s m a l l t r e e s (<10 cm dbh) f r o m f i x e d a r e a p l o t s  .19  T a b l e 5. Mean canopy volume (m ) o f i n d i v i d u a l D o u g l a s - f i r and P o n d e r o s a p i n e t r e e s > 10 cm dbh by d i a m e t e r c l a s s i n c o n t r o l and t r e a t m e n t s i t e s . . .  21  T a b l e 6. T o t a l mean canopy volume (m /ha) f o r D o u g l a s - f i r and P o n d e r o s a p i n e t r e e s > 10 cm dbh i n c o n t r o l and t r e a t m e n t s i t e s  22  T a b l e 7. Mean i n d i v i d u a l t r e e c a n o p y volume (m ) o f t r e e s <10 cm dbh i n c o n t r o l and t r e a t m e n t s i t e s  23  T a b l e 8. T o t a l mean canopy o f t r e e s <10 cm dbh (m /ha) i n c o n t r o l and t r e a t m e n t s i t e s  23  T a b l e 9. Percent cover and c o n t r o l g r i d s  24  3  3  3  3  estimates  T a b l e 10. Number o f 50x50-m c e l l s h a b i t a t type  from  treatment  i n each 41  T a b l e 11. Rank o r d e r o f t h e number o f d e t e c t i o n s i n b o t h t r e a t m e n t s and c o n t r o l s  46  T a b l e 12. Mean number o f t e r r i t o r i e s f o r minimum and maximum d e n s i t y e s t i m a t e s i n c o n t r o l s and t r e a t m e n t s i n 1990 and 1991 by b i r d s p e c i e s  47  T a b l e 13. Mean number o f t e r r i t o r i e s f o r minimum and maximum d e n s i t y e s t i m a t e s .in c o n t r o l s and t r e a t m e n t s by b i r d s p e c i e s  49  T a b l e 14. D e n s i t y o f g r o u n d - f e e d i n g s p e c i e s f o r b o t h minimum and maximum d e n s i t y e s t i m a t e s f o r c o n t r o l and t r e a t m e n t s i t e s  50  vii  T a b l e 15. D e n s i t y o f f o l i a g e - f e e d i n g s p e c i e s f o r b o t h minimum and maximum d e n s i t y e s t i m a t e s f o r c o n t r o l and t r e a t m e n t s i t e s  51  T a b l e 16 Mean number o f d e t e c t i o n s f o r 23 s p e c i e s i n c o n t r o l and t r e a t m e n t s i t e s  53  T a b l e 17 Mean number o f d e t e c t i o n s o f f o u r s p e c i e s groups  54  T a b l e 18 A v e r a g e p e r c e n t c o v e r a g e o f e a c h h a b i t a t t y p e i n each 50x50-m c e l l f o r c o n t r o l s and t r e a t m e n t s  56  T a b l e 19. Rank o r d e r , number o f d e t e c t i o n s and p e r c e n t a g e o f t h e t o t a l number o f d e t e c t i o n s r e c o r d e d t h a t o c c u r r e d i n dense h a b i t a t i n both c o n t r o l and t r e a t m e n t s i t e s  58  T a b l e 20. Rank o r d e r , number o f d e t e c t i o n s and p e r c e n t a g e o f t h e t o t a l number o f d e t e c t i o n s recorded t h a t occurred i n f o r e s t h a b i t a t i n both c o n t r o l and t r e a t m e n t s i t e s  59  T a b l e 21. Rank o r d e r , number o f d e t e c t i o n s and p e r c e n t a g e o f t h e t o t a l number o f d e t e c t i o n s r e c o r d e d t h a t o c c u r r e d i n open h a b i t a t i n b o t h c o n t r o l and t r e a t m e n t s i t e s  60  T a b l e 22. Rank o r d e r , number o f d e t e c t i o n s and p e r c e n t a g e o f t h e t o t a l number o f d e t e c t i o n s r e c o r d e d t h a t occurred i n r i p a r i a n h a b i t a t i n both c o n t r o l and t r e a t m e n t s i t e s  61  T a b l e 23. S i m i l a r i t y i n d i c e s f o r c o m m u n i t i e s i n e a c h h a b i t a t t y p e based on p r e s e n c e / a b s e n c e d a t a u s i n g J a c c a r d ' s s i m i l a r i t y index  65  Table for  24  Summary  of s i g n i f i c a n t  each s p e c i e s i n treatment  Table  25.  Summary  Chi-squared  and c o n t r o l  tests  sites  68  o f d i s c r i m i n a n t f u n c t i o n a n a l y s i s . . . . 72  T a b l e 26. D i s c r i m i n a n t f u n c t i o n c o e f f i c i e n t s and p o o l e d w i t h i n group c o r r e l a t i o n s f o r d i r e c t discriminant function a n a l y s i s f o r treatments and c o n t r o l s T a b l e 27. Mean p e r c e n t c o v e r a g e o f a l l f o u r h a b i t a t t y p e s i n 50x50-m c e l l s (group means) where each s p e c i e s was e i t h e r a b s e n t o r p r e s e n t f o r each o f t h e s i g n i f i c a n t d i s c r i m i n a n t functions  73  74  viii  List Figure  1.  Location  of  of study  Figures  sites  7  F i g u r e 2. Comparison o f s t o c k i n g d e n s i t y and canopy volume / ha o f D o u g l a s - f i r s i n c o n t r o l s and treatments..30 F i g u r e 3. Dominance-diversity and c o n t r o l s  curves  for  treatments 44  F i g u r e 4. Dominance-diversity curves s i t e s i n each o f f o u r h a b i t a t t y p e s  for control  F i g u r e 5. Dominance-diversity curves s i t e s i n each o f f o u r h a b i t a t t y p e s  for  57 treatment  F i g u r e 6. The p o t e n t i a l p r o b l e m s w i t h u s i n g 50x50-m c e l l as t h e u n i t o f a n a l y s i s i n bird/habitat relationships  63 a 87  Acknowledgments  I owe Bunnell.  a g r e a t d e a l o f t h a n k s t o my  I t was  Dr.  Forestry/Wildlife Dr.  B u n n e l l has  B u n n e l l who  i s s u e s i n 1987.  scholastic  Dave Low.  who  me  helped  I would  M i k e Burwash, Sandra K u r t a , in particular,  was  Sarah Bagshaw.  invaluable.  T h a n k s t o a l l of my  my  friend  not  SPSS o v e r  and  Dr.  J . P.  family.  encouragement has  and  and  logistics  Savard  both  help  in  interesting special was  there  life. Her  thanks  to  for  reach  Finally, support  this  goal.  and  me  I would love,  and  t e c h n i c a l e x p e r t i s e i n t h e use  immeasurable.  me  Booth,  Jackie  for their  Lavallee.  Their continued  helped  people  throughout t h i s p r o j e c t .  Loraine  t o mention h e r  out  Wilson,  office  downs o f g r a d u a t e  partner,  t h e y e a r s was  t o t h a n k my  Barry  c o l l e a g u e , V a n e s s a C r a i g , who  t o thank my  support  b u d g e t s and  graduate c o l l e a g u e s  d u r i n g a l l o f t h e ups like  p r o j e c t with  Thanks a l s o t o  I would a l s o l i k e t o e x p r e s s  and  and  Environment  Vyse,  s t a y a t U.B.C. a p l e a s a n t  experience.  in now  inspiration  A r t B l u n d e l l , Darcy  Dr. V a l Lemay and  very h e l p f u l advice  making my  of  to thank a l l those  Frances  Jackie's help with  provided  like  to c o l l e c t data:  interest  and p r o f e s s i o n a l  f r i e n d s h i p of M i n i s t r y of  biologist,  my  Fred  From t h a t t i m e u n t i l  I c o u l d not have c o m p l e t e d t h i s  t h e h e l p and  Johnson.  sparked  been a t r e m e n d o u s s o u r c e  h e l p t h r o u g h o u t both my endeavors.  first  s u p e r v i s o r , Dr.  I would  of  like  1  1.  Introduction  Thinning of overstocked of B r i t i s h  Columbia.  of t h i n n i n g volume  ( S m i t h 1986).  nutrients  yield  Thinning  increase  which u l t i m a t e l y  (Mann and L o h r e y  temporary r e d u c t i o n  of gross t o t a l  reduction  increases  of stand density  i n diameter  it  structure  lead t o  1972, S m i t h  can a f f e c t  increases  1986).  The  p r o d u c t i v i t y due t o t h e i s t h e n o f f s e t by  important  consequences f o r structure  I t i s w i d e l y acknowledged t h a t the composition  ( M a c A r t h u r and M a c A r t h u r Balda  allows f o r  because o f t h e changes i n h a b i t a t  can induce.  i n the  growth.  T h i n n i n g has p o t e n t i a l l y wildlife  from  u p t a k e o f w a t e r and  The s u b s e q u e n t  i n d i a m e t e r growth  initial  timber  reduces competition  f o l i a g e volume, and c a r b o h y d r a t e p r o d u c t i o n extension  objective  o f merchantable  and a l l o w s g r e a t e r  from t h e r o o t s .  crown and r o o t  i n many a r e a s  The p r i m a r y s i l v i c u l t u r a l  i s to increase  neighbouring trees  s t a n d s i s common  of bird  that  habitat  communities  1961, A n d e r s o n and S h u g a r t 1974,  1975, Maurer e t a l . 1981, Morgan e t a l . 1989).  Alterations  i n habitat  structure  habitat  complexity u s u a l l y  species  diversity  result  that  leads  i n increases  ( M a c A r t h u r and M a c A r t h u r  Meslow 1978, M o r r i s o n and Meslow 1983). changes t h a t specific  dead t r e e s , potential  lead to the e l i m i n a t i o n  habitat  components,  t o increases i n  1961, K a r r 1961,  Conversely,  or reduction of  such as shrubs o r  can lead t o t h e r e d u c t i o n  e x t i r p a t i o n o f some s p e c i e s  in bird  standing  i n abundance a n d (Kilgore  1971, P r y a h  2  and J o r g e n s e n 1974," B u n n e l l and A l l e y e - C h a n 1984, R a p h a e l and White bird  1984, P e t i t  e t a l . 1985).  community t o h a b i t a t  species  alteration will  v a r y among  i n accordance w i t h t h e magnitude o f t h e s t r u c t u r a l  alteration Medin  The r e s p o n s e o f any  ( S z a r o and B a l d a 1979, Freedman e t a l . 1981,  1985). The  s t r u c t u r a l changes  that thinning  can produce  been u s e d a s t o o l s t o b o t h enhance a n d d i s c o u r a g e Initially, for  t h i n n i n g was u s e d a s a means t o p r o d u c e  ungulates  Bunnell  pests  t o r e d u c e damage t o c r o p t r e e s by f o r e s t and S u l l i v a n 1988;  G e o r g e and Young 1989; Murua and R o d r i g u e z t h e changes  can produce,  browse  1985; S e v e r s o n and U r e s k 1988)  ( S u l l i v a n and Moser 1986; S u l l i v a n  Recently,  wildlife.  ( B u n n e l l and Eastman 1976, R o c h e l l e and  1978, Whitmer e t al.  and a s a t o o l  have  i n vegetation  especially  1989).  structure that  thinning  t h o s e r e g a r d i n g t h e enhancement o f  a s u p p r e s s e d u n d e r s t o r y , have been s e e n a s a means t o manipulate, The  or affect  biological  e f f e c t s of thinning  l a r g e l y dependent prescription.  diversity  on f o r e s t  on t h e s p e c i f i c  Increased l i g h t  Smith and  1965, 1970,  Zeveloff  Uresk  from heavy  i n u n d e r s t o r y growth Agee and B i s w e l l  understory shrubs  et a l . , (Zeedyk  1989),  communities  reaching the forest  are  thinning can  1973, D i e n  1986, S e v e r s o n and  increased  and Evans  floor  ( M c D o n n e l l and  1970, H a l l s  1980, D o e r r and S a n d b u r g  1988, H i l t  bird  1990).  silvicultural  and d e c r e a s e d c o m p e t i t i o n f o r w a t e r l e a d t o an i n c r e a s e  (Hunter  1975),  fruit  yields  from  and i n c r e a s e d  3  seed production  from g r a s s e s and legumes  (Mann and L o h r e y  1972) . Increases i n understory vegetation result the  t h a t occur as a  o f t h i n n i n g have been a s s o c i a t e d  with increases i n  numbers o f ground and s h r u b n e s t i n g / f o r a g i n g  (Hagar 1990, S t r i b l i n g Heavy t h i n n i n g  e t a l . 1990, D e G r a f f e t a l . 1992).  a l s o can a l t e r  b e h a v i o u r o f some s p e c i e s vegetation (Rolstad other  bird  and Wegge 1989).  species  behaviour  foraging  smaller  (Rolstad  canopy g a p s t h a t  cavity-nesting birds decreases i n stand  limit  the a b i l i t y  foraging,  study,  Any  (snags) o r  stand  to support  Initial  and h e n c e c a n o p y volume, c a n  o f some b i r d  a greater  insectivorous  In t h i s  species.  trees  of that  species The  growth i n t h e w i d t h and l e n g t h  however, c a n c r e a t e  growth,  b o t h n e s t i n g and  bird  1969, F r a n z r e b and Ohmart 1 9 7 8 ) .  increased  can c l o s e  1989).  (Welsh e t a l . 1 9 9 2 ) .  the density  on t h e  (Welsh e t a l . 1992),  f o r some f o r e s t  density  cover  on u n d e r s t o r y  t h a t removes dead s t a n d i n g  t r e e s , may l i m i t  also affect  affect  can a l s o p o t e n t i a l l y  opportunities  prescription  Balda  understory  More m o d e r a t e t h i n n i n g  abundance a n d d i v e r s i t y  Thinning  dying  because o f i n c r e a s e d  h a v i n g any s i g n i f i c a n t  or b i r d  t h e d i s t r i b u t i o n p a t t e r n s and  (Zwank e t a l . 1988) o r d e c r e a s e d canopy  hand, c r e a t e s  before  birds  birds  feeding (Dien  (Morse 1967, eventual  of tree  area  f o r crown-  and Z e v e l o f f  I examined t h e e f f e c t s  crowns,  1980) .  of the  T h o m p s o n / N i c o l a Mule D e e r F o r a g e and S l a s h i n g  P r o j e c t on  4  the  forest  bird  (scientific I)  community  i n dry i n t e r i o r  names f o r a l l p l a n t  Douglas-fir  s p e c i e s appear i n Appendix  forests. This  t h i n n i n g p r o j e c t had t h r e e m a i n  objectives  (Kurta  1991) :  1) t o p r o v i d e h a b i t a t d i v e r s i t y  for wildlife,  2) t o improve t h e snow i n t e r c e p t i o n a b i l i t y o v e r s t o r y t r e e s as l a r g e r crowns 3) t o improve u n d e r s t o r y  of  develop,  and  growth.  To meet t h e s e o b j e c t i v e s , t h e M i n i s t r y o f E n v i r o n m e n t Lands and P a r k s (Kamloops) s p o n s o r e d a l a r g e - s c a l e project. thinning  The p r o j e c t e d  stocking  d e n s i t i e s f o r these  t r i a l s were 200-600 s t e m s / h a  lower than  Columbia M i n i s t r y of F o r e s t s s t a n d a r d s Given t h a t  Lloyd  biogeoclimatic  e t a l . 1990), and t h a t  Douglas-firs predicted in  sub-zones  strata  was  f o u r ways.  increase.  Second,  increase.  Third,  would d e c r e a s e .  First,  individual  removed,  h e r b and s h r u b c o v e r tree  canopy volume Fourth,  xh2;  of the  t h a t t h e s t r u c t u r e o f t h e s e s t a n d s would  at least  increase.  interior  (IDF x h l and IDF  a large portion  i n the lower f o r e s t  British  ( L l o y d e t a l . 1990).  t h e s e s i t e s were i n t h e h o t , d r y  Douglas-fir  thinning  i t was  change would  c a n o p y volume  would  on a p e r h e c t a r e  basis  down and dead woody m a t e r i a l  would  5  In  light  of the p o t e n t i a l consequences of t h i n n i n g  the  p l a n t community,  the  i m p a c t o f t h i s t r e a t m e n t on t h e f o r e s t b i r d  1) An i n c r e a s e  three  p r e d i c t i o n s were made  i n the percent  2)  Removal  potential  regarding community.  c o v e r o f s h r u b s and  h e r b s , would l e a d t o an i n c r e a s e ground/shrub-foraging  on  i n t h e abundance o f  birds.  of a p o r t i o n  of the overstory  foraging habitat  would  limit  and p r o d u c e a d e c r e a s e i n  abundance o f f o l i a g e - f o r a g i n g b i r d s . 3)  Because t h i n n e d  material  woodpecker use o f t h i n n e d  was  left  on  site,  areas f o r foraging  would  increase.  My  approach t o e v a l u a t i n g  forest three  how  the predicted  structure affected the forest bird steps.  thinned  First,  (ground/shrub-feeding species,  species, Third, spatial  and woodpeckers) were  because the treatment  different  associations  structure in  and t h r e e  the species  foliage-feeding  a f f e c t e d by  thinning.  a l t e r e d the quantity  d i s t r i b u t i o n o f some h a b i t a t  within-site  had  S e c o n d , I examined how  r e l a t i v e abundance o f i n d i v i d u a l s p e c i e s groups  community  I e x a m i n e d community  and u n t h i n n e d a r e a s .  changes i n  types,  of i n d i v i d u a l bird  s p a t i a l d i s t r i b u t i o n s of h a b i t a t  and  I examined t h e species types.  with  6  2. The  sites  T h o m p s o n / N i c o l a Mule Deer F o r a g e and S l a s h i n g  p r o j e c t was c o n d u c t e d forest  Study  types  i n IDF x h l and IDF xh2.  e x i s t as a band a l o n g a e l e v a t i o n a l  f r o m 400-1200 m, i n t h e Thompson, and C o l d w a t e r valleys  and i n t h e Okanagan v a l l e y  Region.  T h e s e two  Together  they  gradient, River  i n t h e Kamloops F o r e s t  encompass 604,741 ha o r 7.5% o f t h e  Kamloops F o r e s t R e g i o n o f B.C. ( L l o y d e t a l . 1990). I chose s i x study Cherry sites  s i t e s t h a t were s o u t h  C r e e k , B. C. ( F i g u r e 1 ) . i n thinned  control  areas.  areas  All  stands  as being  had been h a r v e s t e d o f methods  and/or h i g h diameter took  the past  t o t h e 1970's  (Kurta  than  Flagrant contraventions  species also exist  (stems/ha) t h a n  including  o f these  o f my f i e l d  i n these  i n the understory.  T h e r e were a v a r i e t y shrubs  1991).  seasons.  and P o n d e r o s a p i n e were t h e most  t r e e species i n the o v e r s t o r y  Ponderosa pine  of both  mill,  a s l o n g a s stems a r e l e s s  g u i d e l i n e s occurred d u r i n g both Douglas-fir,  s e v e r a l decades  The most r e c e n t h a r v e s t i n g  i s permitted  30 cm dbh.  t o 1990.  ( t i e , cordwood, p o r t a b l e  limit).  place p r i o r  over  Each o f  o f p o o r t o low f o r e s t  Firewood c u t t i n g  common  i n unthinned,  Thinning occurred three years p r i o r  using a variety  activity  sites  study  s i t e was 25 ha i n s i z e .  my s i x s i t e s was c l a s s i f i e d productivity.  T h e r e were t h r e e  and t h r e e study  Each s t u d y  and e a s t o f  stands.  Both  D o u g l a s - f i r was more i n each  c o n i f e r o u s and  Saskatoon b e r r y ,  abundant  spirea,  stratum. deciduous  common j u n i p e r ,  7  Figure  1.  Location  of study  sites  8  r o c k y mountain j u n i p e r , r o s e , k i n n i k i n i k , snowberry. herbs.  The f o r b l a y e r was made up o f b o t h g r a s s e s and  Grasses  included:  p i n e g r a s s , bluebunch  r o u g h f e s c u e , and K e n t u c k y b l u e g r a s s . lemonweed, pussy leafed  arnica,  trembling  toes, yarrow,  aspen, r e d a l d e r , Red-oiser  were:  balsamroot,  In r i p a r i a n  heart-  areas,  a n d p a p e r b i r c h were t h e most  dogwood, D o u g l a s maple,  were common u n d e r s t o r y  twinflower,  wheatgrass,  Common h e r b s  fireweed,  and s t r a w b e r r y .  common t r e e s . and  s h e p e r d i a , and  shrubs.  s t a r Solomon's s e a l ,  willow,  Common h e r b s  were  n o r t h e r n b e d s t r a w , and  horsetail. Continuous the  forest  return  fire  structure i n these  interval  B.C. M i n i s t r y  Kamloops Region,  maintained Without  these stands  parkland structure,  5 m tall  with  roads,  skid  from p a s t  trails  (Dave Low,  formerly structure.  this  D o u g l a s - f i r had r e - s e e d e d  (dbh).  fire  o f E n v i r o n m e n t , Lands and  t o maintain  i n very  and 2.5-15 cm d i a m e t e r  openings  years  i n a n open p a r k l a n d  and now o c c u r s  1.3 m above ground  seven  A natural  p e r s . com.) o n c e  f r e q u e n t ground f i r e s  understory  stands.  of approximately  senior biologist, Parks,  s u p p r e s s i o n had d r a m a t i c a l l y a l t e r e d  open i n the  dense patches  o f t r e e s 2-  o f o u t s i d e bark  measured a t  These patches  a r e juxtaposed  logging operations  (e.g., o l d  and l a n d i n g s ) , s m a l l meadows, and open  f o r e s t s t h a t d i d not r e - s e e d  t o dense D o u g l a s - f i r .  9  The  goal  Slashing of  450  t o a c h i e v e an  l i v e stems/ha.  healthy  contractor c a n o p i e s and  Ponderosa pines  possible varied  T h o m p s o n / N i c o l a Mule Deer F o r a g e  P r o j e c t was  thinning  m.  of the  This prescription  to save D o u g l a s - f i r remove c o m p e t i n g  1991).  The  f r o m 172  t o 1100  t r e e s per  density  North-facing  was  slopes >  15  cm  dbh  quality. provided height  were not  south-facing  f e l l e d unless t h a n 50  t h a t workers d i d not overstory  t h a t was  being  pockets of unthinned  cm  on  fall  trees plus thinned. areas.  each  site  in  better  t o between 400-600 or h o t t e r ,  were  dryer Any  stems  poor  retained,  trees within the  4  position.  t h e y were v e r y dbh  1 to  wherever  t o between 250-300 s t e m s / h a .  Snags g r e a t e r  of the  material in  R i d g e t o p s and  full,  Variability  representing  were t h i n n e d  the  stems w i t h i n  to topographic  or wetter slopes,  were t h i n n e d  ha.  density  trees with  degree of t h i n n i n g  related  p o t e n t i a l growing s i t e s , stems/ha.  required  o f a l l s i z e s were r e t a i n e d  (Kurta  stocking  average stocking  and  height  of  one  tree  the  Snags, t h e r e f o r e ,  occur  10  3.  E f f e c t s of t h i n n i n g  The  on t h e p l a n t  community  T h o m p s o n / N i c o l a Mule Deer F o r a g e and  P r o j e c t was d e s i g n e d t o i n c r e a s e  Thinning  h a b i t a t d i v e r s i t y by  p r o m o t i n g u n d e r s t o r y g r o w t h and t o i m p r o v e t h e snow interception ability  of overstory  assessed the e f f i c a c y desired  changes.  production  thinning  I t was p r e d i c t e d  would i n c r e a s e ,  would i n c r e a s e , decrease,  of t h i s  trees.  This  trial  study  t o produce  t h a t h e r b and s h r u b  i n d i v i d u a l t r e e canopy volume  canopy volume on a p e r h e c t a r e  and down and dead woody m a t e r i a l  basis  would  would  increase.  Methods Vegetation  plots  Vegetation  s t r u c t u r e was sampled a t 20 s y s t e m a t i c a l l y  l o c a t e d p l o t s i n each o f t h e s i x s t u d y A u g u s t o f 1991. according stratified  Despite  t o slope  the thinning  by s l o p e  p o s i t i o n f o r three small  t o sample them e f f e c t i v e l y .  difficult  t o d e t e r m i n e where r i d g e  slopes  ended.  and s o u t h s l o p e s  varying  s a m p l i n g was n o t reasons.  First,  i n s i z e making i t  difficult  or south slopes  i n J u l y and  treatment  position, vegetation  r i d g e t o p s were o f t e n v e r y  north  sites  Second,  i t was  often  t o p s began and where  Third,  ridge  were n o t p r e s e n t  tops,  north  i n a l l study  sites. Variable-radius snags g r e a t e r  than  p l o t s were u s e d t o sample t r e e s and  10 cm dbh.  I used a w i d e - f i e l d  11  relaskope with a basal-area f a c t o r p l o t s were c h o s e n b e c a u s e t h e y efficient sampling  than  fixed  densities.  e s p e c i a l l y when t h e s e  s p e c i e s and s t a t u s  o c c u r a t low  A l l trees greater  10 cm dbh were t a l l i e d  (live  than  according  o r dead) w i t h i n a 10-m  (0.03  fixed radius plot. Percent cover of t r e e ,  dead woody m a t e r i a l , b a r e needles), 10-m  i n the  S p e c i e s , dbh, and h e i g h t were r e c o r d e d f o r e a c h  2.5 cm dbh and l e s s t h a n  ha),  time  and a r e e f f e c t i v e  t r e e t h a t was deemed " i n " t h e p l o t .  to  Variable radius  a r e o f t e n more  area p l o t s ,  of large trees,  o f 1.  ground,  herb, litter  area p l o t  Canopy c o v e r a b o v e 4 m from  e s t i m a t e d u s i n g a 'moose h o r n ' In  each v e g e t a t i o n p l o t ,  at  2.5-m  (primarily  a s i n Walmsley e t a l . t h e ground  random number  was 1989). times  long, perpendicular  a t the centre of the p l o t .  17 canopy c o v e r measurements were t h e n  transect direction  i n t h e same  ( B u n n e l l and V a l e s  i n t e r v a l s a l o n g two 20-m  an o v e r a l l  pine  c a n o p y c o v e r was measured 17  transects that intersected  determine  g r a s s , down and  and moss p l u s l i c h e n were e s t i m a t e d  r a d i u s (0.03 ha) f i x e d  (1980).  shrub,  estimate  averaged  to  o f canopy c o v e r .  was a compass  The  The  bearing generated  initial from  a  table.  Canopy volume Canopy volume measurements were t a k e n and were r e c o r d e d  after  sampling  was c o m p l e t e d  (1968).  A l l volume measurements were r e c o r d e d  vegetation  as i n Sturnam on t r e e s  12  encountered trees  in earlier  vegetation  f o r each s p e c i e s  sampled p r o p o r t i o n a l  than  to occurrence. on t r e e s  I sampled up t o 60 t r e e s  trees  I previously  Canopy volume o f t r e e s tallied  i n fixed  both greater  10 cm dbh a t each s t u d y s i t e  > 10 cm dbh t h a t  The numbers o f  and i n e a c h d i a m e t e r c l a s s were  < 10 cm dbh were r e c o r d e d plots.  surveys.  (Table  sampled,  from each d i a m e t e r c l a s s .  Two  1) .  area  t h a n and l e s s Of t h e t r e e s  I randomly  Douglas-fir  selected t r e e s of  d i a m e t e r c l a s s 1 (10-20 cm dbh) a n d one f r o m d i a m e t e r  class  2  until  (21-30 cm dbh) were sampled a t e a c h v e g e t a t i o n  q u o t a s were f i l l e d .  Trees  i n the l a r g e r diameter  were sampled o p p o r t u n i s t i c a l l y a s t h e y o c c u r r e d vegetation For three  p l o t u n t i l q u o t a s were  Douglas-fir  trees  plot  i n each  filled.  l e s s than  t r e e s were s e l e c t e d w i t h i n  classes  10 cm i n d i a m e t e r ,  a 10-m  radius  from  each  plot  center.  plot  d e t e r m i n e d by a r a n d o m l y c h o s e n compass b e a r i n g  distance  I sampled t h e c l o s e s t t r e e t o a p o i n t  from t h e p l o t c e n t e r .  t r e e s were e n c o u n t e r e d , denser p l o t s t o f i l l  3 trees  Small Ponderosa pine  were sampled o p p o r t u n i s t i c a l l y a s t h e y o c c u r r e d vegetation  plot until  q u o t a s were  basis.  density  to y i e l d  trees  i n each  species  meters of f o l i a g e ,  and d i a m e t e r c l a s s .  T h e s e volumes were t h e n m u l t i p l i e d by t h e stocking  on  filled.  Canopy volumes, e x p r e s s e d a s c u b i c were c a l c u l a t e d f o r e a c h t r e e  and  I n p l o t s where l e s s t h a n 3  I s a m p l e d more t h a n  quotas.  i n the  appropriate  c a n o p y v o l u m e on a p e r  hectare  13  T a b l e 1. Number o f t r e e s s a m p l e d f o r canopy volume measurements i n each c o n t r o l and t r e a t m e n t p l o t by s p e c i e s and d i a m e t e r c l a s s . Diameter class  Tree species  Number sampled  Douglas-fir  0 1 2 3 4 5  (2.5-9.9 (10-20 (21-30 (31-40 (41-50 (>50  cm) cm) cm) cm) cm) cm)  60 35 10 3 3 3  Ponderosa pine  0 1 2 3 4 5  (2.5-9.9 (10-20 (21-30 (31-40 (41-50 (>50  cm) cm) cm) cm) cm) cm)  3 3 3 3 3 3  14  Data  analysis To  examine t h e  differences  between t r e a t m e n t s and trees  and  controls  s n a g s f o r e a c h dbh  (Douglas-fir of v a r i a n c e condition  and  analysis  differences of  (<10  cm  dbh)  c o n i f e r s , and S o k a l and  interpreted despite  controls total  on  Rolf  for  Ponderosa  The  on  the  same  for  stocking  dead D o u g l a s - f i r  arcsine  transformed  (1981), main e f f e c t s  and  individual  cover were  nested  of variance  between t r e a t m e n t s  canopy volume o f  and  in  d e n s i t i e s f o r a l l small  c a n o p y volume/ha f o r e a c h dbh  (Douglas-fir  analysis  used to t e s t  A one-way a n a l y s i s  for differences the  species  o c c a s i o n s where s i g n i f i c a n t  e f f e c t s were d e t e c t e d . used to t e s t  and  stocking the  was  controls  live  trees  Following  total  live  i n Appendix I I ) .  of v a r i a n c e  the  data.  tree  cm)  c o n t r o l ; ANOVA t a b l e  Ponderosa pine, small  (>10  p l o t s were n e s t e d  between t r e a t m e n t s and small  densities  I u s e d a one-way  i s presented  n e s t e d one-way a n a l y s i s  large  c l a s s and  Ponderosa p i n e ) ,  ( t r e a t m e n t and  and  of  i n which v e g e t a t i o n  vegetation  density  i n stocking  and  trees  c l a s s and  was  tree  and  the  species  pine).  Results Stocking  density:  Densities  of  live  d i a m e t e r c l a s s e s >10 densities not  differ  of  Large trees  cm  dbh  each s p e c i e s  significantly  trees (stems/ha) by  are  and  summarized  species  and  i n T a b l e 2.  i n each diameter c l a s s  between t r e a t m e n t  and  did  controls  The  15  T a b l e 2. Mean d e n s i t i e s o f l i v e t r e e s (stems/ha) > 10 dbh i n t r e a t m e n t and c o n t r o l s i t e s by t r e e s p e c i e s and diameter c l a s s . DBH class  Species  Control  Treatment  cm  p-value  >0 .25  Douglas  1  (10- 20  cm)  198 . 1  171. 8  -fir  2  (21- 30  cm)  51. 6  64 . 2  >0 . 25  3  (31- 40  cm)  17. 4  16. 6  >0 .25  4  (41- 50  cm)  5 2  4 .9  >0 .25  5  (>50  2 8  2 8  >0 .25  Ponderosa  1  (10- 20  cm)  7  6 6  >0. 25  pine  2  (21- 30  cm)  5 9  5 3  >0 .25  3  (31- 40  cm)  3 5  3 7  >o .25  4  (41- 50  cm)  1 8  0 .7  0 . 25<p<0  5  (>50  0 .6  1 .0  >0 .25  cm)  cm)  13  16  (a=0.05).  The h i s t o r y o f s e l e c t i v e l o g g i n g  i n a diameter d i s t r i b u t i o n t h a t trees.  For both D o u g l a s - f i r  favours  has r e s u l t e d  younger/smaller  and Ponderosa  pine,  y o u n g e r / s m a l l e r t r e e s were t h e most common a g e / s i z e  class.  Stocking  class  density  decreased with  i n c r e a s i n g diameter  i n b o t h t r e a t m e n t s and c o n t r o l s . common o v e r s t o r y sites,  tree species  and c o m p r i s e d  summarized  93% a n d 83% o f o v e r s t o r y  Densities  significantly  Smaller Douglas-fir  t h e most common s i z e c l a s s .  Stocking  density  and  Ponderosa p i n e  extremely reflects of  i n b o t h t r e a t m e n t s and c o n t r o l s .  the logging  h i s t o r y of these stands.  generally  targets  Douglas-fir  treatment  The  The r e m o v a l  of large  In addition,  l a r g e r diameter  live  firewood  snags,  skewing t h e diameter d i s t r i b u t i o n i n f a v o u r snags.  size  low d e n s i t i e s o f l a r g e r d i a m e t e r c l a s s e s o f s n a g s  i n t o t h e snag p o p u l a t i o n .  cutting  treatments  snags i n t h e s m a l l e s t  l a r g e r l i v e trees reduced recruitment  trees  between  o f snags  i n c r e a s i n g diameter c l a s s i n both  c l a s s e s were a b s e n t  and i n  s n a g s were  decreased with controls.  density  and d i a m e t e r c l a s s e s a r e o f each s p e c i e s  each diameter c l a s s d i d n o t d i f f e r t r e a t m e n t s and c o n t r o l s .  stem  respectively.  o f s n a g s by s p e c i e s  i n T a b l e 3.  was t h e most  i n t r e a t m e n t and c o n t r o l  i n t r e a t m e n t and c o n t r o l s i t e s , Densities  Douglas-fir  further  of smaller  was t h e most common s n a g s p e c i e s i n  and c o n t r o l s i t e s ,  b o t h t r e a t m e n t and c o n t r o l  and c o m p r i s e d  sites.  95% o f s n a g s i n  17  T a b l e 3. Mean d e n s i t i e s o f s n a g s (stems/ha) > 10 cm dbh i n t r e a t m e n t and c o n t r o l s i t e s by t r e e s p e c i e s and d i a m e t e r class. DBH class  Species  Control  Treatment  p-value  23 . 3  >0. 25  4. 1  3 .1  >0 . 25  cm)  1.5  1.2  >0. 25  cm)  0.7  1.2  >0. 25  1.0  1.2  >0 .25  cm)  0.0  0.0  (21-30  cm)  0.3  0.8  >0. 25  3  (31-40  cm)  0. 1  0.1  >0 .25  4  (41-50  cm)  0.7  0.3  5  (>50  0.5  0.4  Douglas  1  (10-20  cm)  22 . 4  -fir  2  (21-3 0  cm)  3  (31-40  4  (41-50  5  (>50  Ponderosa  1  (10-20  pine  2  cm)  cm)  n/a  0.25<p<0 >0. 25  18  Stocking  density:  Small  I n b o t h t r e a t m e n t s and common s m a l l Douglas-fir  tree  t r e a t m e n t and Douglas-fir of  controls Douglas-fir  dbh)  98%  i n the  and  99%  were dead  (21%  and  i n treatments  o f dead D o u g l a s - f i r  significantly  30%,  (rocky did  of  d i d not  controls.  small  large standard  (both  sites  were due  individual,  sites  f r o m 0 t o 6000  and aspen)  species  and  more c o n i f e r s  i n b o t h c o n t r o l and  (0.025  in control The the  treatment  (variability)  Estimates of  i n d i v i d u a l 0.03-ha v e g e t a t i o n  trees/ha.  The  'other'  with estimates of  plots.  the  dead) i n  (0.025 < p < 0.01).  extreme p a t c h i n e s s  b a s e d on  in  were  (0.025 < p < 0.01).  0.03-ha v e g e t a t i o n  of D o u g l a s - f i r ranged  to the  l i v e and  between c o n t r o l s  errors associated  d e n s i t i e s of D o u g l a s - f i r  The  higher  there  t r e e s of a l l s p e c i e s  than i n treatment  the  significantly  Together,  T h e r e were s i g n i f i c a n t l y  < p < 0.01), and sites  differ  birch, trembling  differ significantly  treatments.  both  respectively).  dead Ponderosa p i n e s  mountain j u n i p e r ,  not  In  4).  stems  (0.025 < p < 0.01), w h i l e  more D o u g l a s - f i r  l i v e and  (Table  understory  significantly  c o n t r o l p l o t s than i n treatments densities  of the  the  a s u b s t a n t i a l p o r t i o n of  l i v e D o u g l a s - f i r was  between t r e a t m e n t s and  was  understory  controls, respectively.  control sites  c o n t r o l s than density  cm  represented  i n t r e a t m e n t s and  density  (<10  trees  of  density plots  19  H  0)  XX xx  rH  > I  TS  e  ft  o  o • o in V CN ft V o in A CN o  in CN o o v ft v in o  o H V  in in in in CN o o o o o o in in in o • • CN CN CN o • • • V V V ft ft ft ft O o o V v V V V A A A in rH rH rH o o o o  in CN o o v a v in o  >1 o o «  01 <D  CD  u  -P C.  -p rH •  rd 01  e -p 03 C 0)  <M g  CO  in  (0 0)  vo VD cn o o m rH in  e -p  r~  rH rH  rH  rH  H  rH  CN  co  vo  VO  CM  in rH rH  n  co o in  o  CN  in  rH CN  o  rH  JH  x:  -P  >H  TS  a rd  SH  SH CO  o  rH  -P C O CJ  CN  o  m in  co oo  CN  cn  CN  rH  n  in  H  in  o  VO CM  CN  rH n n n  CN  CN  co rco rrH  o n  CN  rH  u  SH  C CO  ft 01 rd  0) o  IT  -P -P  —c  C  o >. o •p  TS rd CO  C  -H  TS  01 - H  c  0">r-H  T3  O  a  C  -H  X O O  rd 0)  rH  - P (0  SH  X! rd  -rH U rd  >  CO TS CO • X  ^J"<w-H cu  rH e XI o rd  rH  E H <+H  c -H  o o -p w  -rH  <H  TS  >  <D  -H rH  >H -rH 4H  C  U  -rH <4H  1 01 1 1 01  01 rd  rd  rd  rH  rH  rH  CP  CP  CP  0  o  O  -rH  rd 01 o  u CO TS  c o  a>  rH  >  X!  e a)  •H rH  SH  CO C  -rH  ft  ft  rd 01  rd 01  O !H CO  O  TS  -P  CO UT  rd d)  01 -P  QJ TS  ft rd ft  o rH  01 01 CO  xi U  h X!  EH  — a) rd  rd  •  -P rd  0  TS C  u  TS  rd 0)  TS  > rH  C  XI -P  0) Xi -P  O  O  a.  O  CO  rd  rH  C 0 Oi  rd  -rH rH  CO TS  CO  TS  CO  vw  SH CO  SH CO  rH rH  4H --H  XX -P o  rd  c o 0  e 01  TS  O SH  -H 0 <4H -H  1  -P 01 SH rrj 0 rH  Q  Q  CM  rH  rH  rH  rH  rH  rH  rH  rH  rH  rH  rH  fd -p 0  rd -P  rd -P  rd -P  rd -P  rd -P  rd -p  rd -P  rd -P  rd -P  O  O  rd -P 0  O  O  SH  EH  FH  EH  EH  &< Q  O  O EH  O  EH  O EH  O  -  ft O  CP  3 0  SH  3 Q)  rH  a  O  -H  c  c 3  •H  C  Q  (H  01 QJ • TS  ^H  a)  •t—1 c  - P ro O -P c II p 0 rd E  20  Canopy volume: T h e r e was  no  Large  significant  canopy volumes f o r e i t h e r between c o n t r o l s and (Table 5).  On  significantly  treatments (0.05  difference  f o r each diameter  a per h e c t a r e b a s i s , higher  of i n d i v i d u a l  D o u g l a s - f i r or Ponderosa  treatments  cm  dbh  (0.025 < p <  pine class  for  0.01).  canopy v o l u m e s / h a were s i g n i f i c a n t l y than  higher i n  i n c o n t r o l s f o r P o n d e r o s a p i n e s >50  < p < 0.025; T a b l e  Canopy volume:  tree  c a n o p y volumes were  i n c o n t r o l s than treatments  P o n d e r o s a p i n e s 31-40 Conversely,  trees  cm  dbh  6).  Small  trees  I n d i v i d u a l t r e e canopy volumes f o r b o t h D o u g l a s - f i r and  P o n d e r o s a p i n e were n o t s i g n i f i c a n t l y  treatment 7) .  sites  v o l u m e s on  8) .  i n c o n t r o l s as was  Because of the s i g n i f i c a n t l y  Douglas-fir  for  than  in control plots  differ  higher d e n s i t y of  live  canopy  a p e r h e c t a r e b a s i s were s i g n i f i c a n t l y  higher  total  plots  (0.01  < p < 0.005;  Table  canopy volume o f P o n d e r o s a p i n e d i d n o t  significantly  control  p r e d i c t e d (Table  (Table 4), the t o t a l  Douglas-fir in control The  higher i n  different  between t r e a t m e n t  and  plots.  C o v e r measurements The greater H e r b and  percent cover i n treatment low  shrubs  o f g r a s s was  sites (< 1 m)  than  not  significantly  in control  cover  also  sites  showed no  (Table 9). response  21  T a b l e 5. Mean canopy volume (m ) o f i n d i v i d u a l D o u g l a s - f i r and P o n d e r o s a p i n e t r e e s > 10 cm dbh by d i a m e t e r c l a s s i n c o n t r o l and t r e a t m e n t s i t e s . 3  dbh class  Species  Control  Treatment  p-value  0 . 25<p<0.1  Douglas  1  (10-20 cm)  8.1  5.7  -fir .  2  (21-30 cm)  26.9  15.1  0.25<p<0.1  .3  (31-40 cm)  61.5  43.1  >0.25  4  (41-50 cm)  95.5  80.3  >0. 25  120.6  108.1  >0. 25  5  (>50 cm)  Ponderosa  1  (10-20 cm)  9.6  7.8  >0.25  pine  2  (21-30 cm)  30.6  18.2  >0.25  3  (31-40 cm)  50.4  37.5  0. K p < .0.05  4  (41-50 cm)  61.5  100.1  >0.25  144.6  184.5  0.25<p<0.1  5  (>50 cm)  22  T a b l e 6. T o t a l mean canopy volume (m /ha) f o r D o u g l a s - f i r and P o n d e r o s a p i n e t r e e s > 10 cm dbh i n c o n t r o l and treatment s i t e s . 3  dbh class  Species  Control  Treatment  p-value  Douglas-  1  (10-20 cm)  1623 . 5  940. 2  0.25<p<0.1  fir  2  (21-30 cm)  1398 . 6  976. 5  0.25<p<0.1  3  (31-40 cm)  1079 . 4  749 . 2  0.25<p<0.1  4  (41-50 cm)  493 . 5  420. 5  >0.25  5  (>50  389. 0  399 . 8  >0.25  Ponderosa  1  (10-20 cm)  171. 3  54 . 1  pine  2  (21-30 cm)  190. 6  101. 3  >0. 25  3  (31-40 cm)  175. 5  146 . 4  0.025<p<0.01  4  (41-50 cm)  116. 7  58 . 1  5  (>50  93 . 1  259 . 1  cm)  cm)  0.25<p<0.1  >0. 25 0 . 05<p<0.025  23  Mean i n d i v i d u a l t r e e canopy volume <10 cm dbh i n c o n t r o l and t r e a t m e n t s i t e s . Table  7.  Species  Control  Treatment  (m ) o f t r e e s 3  p-value  Douglas-fir  1.6  2. 2  0.25<p<0.1  Ponderosa p i n e  2.3  3. 3  >0.25  T a b l e 8. T o t a l mean c a n o p y v o l u m e s o f t r e e s (m /ha) i n c o n t r o l and t r e a t m e n t s i t e s .  <10 cm dbh  3  Species  Douglas-fir Ponderosa  pine  p-value  Control  Treatment  1332 .4  227.4  0.01<p<0.005  66. 2  11.3  >0.25  24  T a b l e 9. sites. Cover  P e r c e n t c o v e r e s t i m a t e s on t r e a t m e n t  variable  Control  Down and dead  12.6  Litter  11. 3  Grass  Treatment  23 . 0  and c o n t r o l  p-value  0.01<p<0.005  9 .4  >0. 25  12 . 5  14 . 8  >0.25  12 .9  10. 5  0.25<p<0.1  Tree  5.5  3 .4  0.01<p<0.005  S h r u b <1 m  9.8  5. 3  0.25<p<0.1  S h r u b 1-4 m  2. 7  0. 10  0.05<p<0.025  Bare  ground  Herbs  12 . 0  10. 7  >0.25  Moss & l i c h e n  22.8  22 . 7  >0.25  Canopy  38 . 3  25. 9  >0.25  cover  25  to the  treatment.  however, was  Percent cover t a l l  significantly  greater  shrubs  (1-4  m),  in control sites  (0.05  <  p < 0.025) . Because t h i n n e d c o v e r o f down and treatment Due  sites  to the  sites,  the  closure  dead was  stocking  higher  recorded  on  significantly  density  (0.01  site,  the  higher  (0.0K  of t r e e s  percent  in  p <  0.005).  in control  The  where s t o c k i n g  comparatively  illustrates  density  dead  canopy c o v e r  (Table  was  < p < 0.005) i n c o n t r o l s t h a n  the  i n b o t h t r e a t m e n t s and  canopy was  left  p h y s i c a l space t h a t t r e e t r u n k s occupied  sites.  overstory  was  than i n c o n t r o l s i t e s  higher  significantly treatment  material  low  estimates of  canopy  patchiness  of  the  controls.  In  addition,  a large proportion  in  was  high,  of  the  4),  further c o n t r i b u t i n g to  low  thinning  alter  estimates.  Discussion The forest  main g o a l  of t h i s  s t r u c t u r e f o r the  designed s p e c i f i c a l l y believed  t o be  to  the  trees  overstory  sampling regarding  t o promote the  (Kurta two  to  Thinning  g r o w t h o f what  suppressed understory  improve t h e  indicated that the  b e n e f i t of w i l d l i f e .  a previously  community and  p r o j e c t was  was  plant  snow i n t e r c e p t i o n a b i l i t y  1991). of the  e f f e c t s of t h i n n i n g  Results four  from  initial  occurred  as  was  of  vegetation predictions  expected.  As  26  was  i n i t i a l l y predicted,  increase  i n the  a significant Douglas-fir g r o u n d and did  the  thinning  amount o f down and  decrease i n the  trees.  shrub cover d i d not  Several  significant  d e a d woody m a t e r i a l  c a n o p y volume p e r  In c o n t r a s t ,  canopy volume o f  stands.  resulted in a  contrary  increase  hectare  nor  in treated  have c o n t r i b u t e d  n e g l i g i b l e r e s p o n s e o f t h e s e two  of  predictions,  significantly,  individual trees  factors could  to  and  to  the  habitat variables after  thinning.  Ground  cover  Cattle grazing grazing  can  negatively (Stoddart  may  change t h e  s t r u c t u r e of g r a s s l a n d  a f f e c t i n g plant vigour, e t a l . 1975)  (Kosco and  o r by  B a r t o l o m e 1983)  composition  can  a l s o d i s t u r b the  grassland  communities.  This  cyanophytes s t a b i l i z e s  l a y e r of  the  soil  1982), e n h a n c e s s o i l  water r e t e n t i o n  R u s h w o r t h 1983), and  increases  fixation  by  Trampling lead to  cyanobacteria  can  reduce the  soil  layer  of  bryophytes,  ( A n d e r s o n and (Brotherson nitrogen  of t h i s  f u r t h e r a l t e r a t i o n of the  Cattle  lichens,  (Cameron and extent  density  species  (Ryder 1980).  cryptogamic  plant  by  productivity  vegetation  thereby a l t e r i n g  Cattle  vegetation  g r o w t h , and  reducing  of p l a n t communities  trampling  and  have a f f e c t e d g r o u n d c o v e r .  Fuller  Rushforth and  content  via  1960).  l a y e r which community.  can  27  C a t t l e have been g r a z e d  i n t h e IDF  x h l and  xh2  stands  i n t h e Kamloops a r e a s i n c e t h e l a t e  1800's  com.).  c a t t l e were b r o u g h t  D u r i n g my  two  field  t o my  study s i t e s e a r l y  until  l a t e June,  elevations.  i n May.  I n 1990,  they  They were t h e n r e t u r n e d t o my In 1991,  treatment  1 control)  and  c a t t l e remain  cattle  remained  throughout  was  s u b j e c t t o c a t t l e g r a z i n g f r o m May  have been o n g o i n g  to  On  the treatment  similar  On my  technician,  four other s i t e s  fashion to that  on t h e s e s i t e s decades.  influence  Although  response. physical  Leaving slash  on  in a  C a t t l e h a v e been g r a z e d last  several to,  and  the  predicted. have b e e n a f a c t o r  f o r the  other  lack of understory  s i t e may  have r e p r e s e n t e d a  b a r r i e r t h a t p r e v e n t e d ground  r e s p o n d i n g as i t might  fences  Kamloops F o r e s t  have p r e v e n t e d  c a t t l e g r a z i n g may  explanations could account  there  been moved  i n these stands p r i o r  from r e s p o n d i n g a s  that  c a t t l e were r o t a t e d  i n 1990.  s i n c e t h e t i m e o f t h i n n i n g may understory  site  from  t o September,  i n a s i m i l a r manner f o r t h e  Their  (1  stands  problems w i t h p o o r l y m a i n t a i n e d  range  in  September.  w h i c h a l l o w e d c a t t l e t o r e t u r n a f t e r t h e y had (Mike D e e d l e ,  sites  i n t h e s e lower e l e v a t i o n  t o September i s u n u s u a l .  Service).  on  remained  study s i t e s  on two  May  May  out  pers  a f t e r w h i c h t h e y were moved t o h i g h e r  l a t e August.  Having  seasons  (Dave Low,  cover  have i f s l a s h had  from  been removed.  28  Insufficient thinning  t i m e may  on u n d e r s t o r y  a p p r e c i a b l e way. thinning from  to manifest  themselves  Most s t u d i e s e x a m i n i n g  7 t o 22 y e a r s a f t e r  1972,  Hilt  f o r the e f f e c t s  and  S m i t h 1970,  Doerr  and  Sandburg  Sonderman 1989,  1986,  Alaback  Hagar 1 9 9 0 ) .  examination.  understory vegetation a f t e r  1965), 4 y e a r s (Crouch  of t o t a l years  (Barrett 1986).  Research potentially  1-2  the r e s u l t s types  shrub  In each of  significant  3 years 1970,  Crouch  these  of response  increases i n  (Zwank e t a l .  (McDonnell  A u s t i n and  and  1988,  Smith  U r n e s s 1982)  (1986) a l s o n o t e d  has  annual  shown t h a t t h r e e y e a r s  l o n g enough t o e x p e c t  a significant  and  5  gains  i n the  ( e . g . , A u s t i n and Stribling  U r n e s s 1982,  e t a l . 1990) .  after thinning in similar  was  response  p l a n t community.  r e p o r t e d above, however, were from  cover  1988,  after thinning.  elsewhere  Zwank e t a l . 1988, and  Herman  a t the time  years  the p r o d u c t i o n of the u n d e r s t o r y  forest  and  p l a n t p r o d u c t i o n on h e a v i l y t h i n n e d p l o t s  immediately  1970,  Sassaman e t  S t u d i e s t h a t measured v e g e t a t i o n  e t a l . 1990),  anywhere  i n c r e a s e s i n t h e amount o f  a f t e r thinning, recorded  Stribling  of  Biswell  G r e l e h e t a l . 1972,  u n d e r s t o r y v e g e t a t i o n were r e c o r d e d  years  i n an  the e f f e c t  t h i n n i n g (Agee and  other studies, s i g n i f i c a n t  shortly  of  on u n d e r s t o r y d e v e l o p m e n t were p e r f o r m e d  McDonnel and al.  have passed  Most  in of  moister  Crouch  1986,  Increased forest  types  herb  29  occurred 1965,  i n the absence of g r a z i n g (McDonnell  Barrett  1970).  o v e r t h e l a s t 50+ vegetation  The  been e x c l u d e d  from  have p r e v e n t e d  that  in similarly  these s i t e s  after  of c a t t l e .  thinning.  take a long time (McLean and  a significant  p e r c e n t cover of ground c o v e r . did significantly  diameter 60%  occur with  to occur Tisdale  t h e e x t e n t o f t h e t h i n n i n g may  s e v e r e enough t o g e n e r a t e  trial  not  the shown  reduce  even 1972).  n o t have  thinning  stocking density of smaller  c l a s s e s of D o u g l a s - f i r s ,  and  did result  (Figure 2).  i n up  Given  e x t e n t o f t h e r e d u c t i o n o f c a n o p y v o l u m e / h a on t h e s e i t i s not unreasonable  been  change i n t h e  However, t h i s  r e d u c t i o n o f canopy volume/ha  sites,  had  Significant  P r e v i o u s r e s e a r c h has  have been e x c l u d e d  Finally,  understory  d r y a r e a s a p p r e c i a b l e changes i n  u n d e r s t o r y v e g e t a t i o n may when c a t t l e  the  stands  as p r e d i c t e d even i f c a t t l e  i n c r e a s e s i n u n d e r s t o r y v e g e t a t i o n may continued presence  Smith  g r a z i n g r e c e i v e d by t h e s e  y e a r s may  from r e s p o n d i n g  and  to expect  an  to a  the xeric  increase i n the  p e r c e n t cover of ground v e g e t a t i o n .  T r e e canopy volumes There individual  a r e two  p o t e n t i a l r e a s o n s why  t r e e s d i d not respond  r e l e a s e of t r e e s a f t e r tree vigour  (Barrett  thinning  1969,  canopy v o l u m e s o f  as e x p e c t e d .  First,  i s p r i m a r i l y governed  Helms and  S t a n d i f o r d 1985).  by Any  30  1 0 0 0 -,  2 0 0 0 -i  F i g u r e 2. Comparison o f s t o c k i n g d e n s i t y and canopy volume / h a o f D o u g l a s - f i r s i n c o n t r o l a n d t r e a t m e n t s i t e s  31  factors that negatively potential  tree vigour  b e n e f i t s of t h i n n i n g .  Douglas-fir  epidemic l e v e l s  At  (Orgyia  t u s s o c k moth  S p r u c e budworm  Forest  affect  (Choristoneura  E n t o m o l o g i s t , Kamloops F o r e s t  a f f e c t s on  Douglas-fir  expected  increase  can  their  Previous  anywhere f r o m 2-5  and  Helms and three  Ethridge  1976, 1985).  i n such d r y increase  treatment period, s p r u c e budworm and vigour,  likely  canopy volume.  negative Schmidt prevented  individual  insufficient  time  i n the  1977,  i t can  This  forest  Douglas-fir  A short  negative  of thinning  only  increase  in  be  a post-  effects  t u s s o c k moth on  l a c k of  take  1973,  stands  types to produce  coupled w i t h the  may  amount o f t i m e may  i n c a n o p y volume.  led to the  Gordon  I examined my  trees.  predicted  a marked i n c r e a s e  Herring  years after thinning.  significant  comm.).  h e n c e t r e e canopy g r o w t h a f t e r  Standiford  insufficient  have  have shown t h a t  years to detect  d i a m e t e r g r o w t h and (Herring  studies  pers.  influence  have p a s s e d f o r t h e s e s t a n d s t o r e s p o n d direction.  near  e t a l . 1982,  i n c a n o p y volume o f  S e c o n d as w i t h g r o u n d c o v e r ,  Western  Maclauchlan,  Service,  (Carlson  e t a l . 1983), i t i s p o s s i b l e t h a t  and  were a t  insect pests  vigour  study,  pseudotsugata)  fumiferana)  forest  delay  time of  i n these stands(Lorraine  G i v e n t h a t t h e s e two  the  the  can  of  Douglas-fir Douglas-fir  32  Conclusions The respond this  v e g e t a t i o n on t h e t h r e e t r e a t m e n t as p r e d i c t e d .  t h i n n i n g have been c o n f o u n d e d  factors. and  It is likely  First,  the e f f e c t  p r e s e n t , may  itself  may  Fourth,  s p r u c e budworm and  g r a z i n g , both  past  felled  Second,  may  have Third,  D o u g l a s - f i r t u s s o c k moth o f c a n o p y volume t o  t i m e s i n c e t r e a t m e n t may  the  may  thinning.  h a v e been i n s u f f i c i e n t  o f e i t h e r u n d e r s t o r y v e g e t a t i o n and  R e s u l t s o f my  study  indicate that  to generate the d e s i r e d  quickly.  different  change i n u n d e r s t o r y v e g e t a t i o n .  have h i n d e r e d the response  permit response  of  have n o t b e e n s e v e r e enough t o  a significant  unable  a number o f  i n c r e a s i n g as e x p e c t e d .  produce  volume.  by  t h e growth o f u n d e r s t o r y v e g e t a t i o n .  thinning t r i a l  Finally,  t h a t the e f f e c t s  of c a t t l e  s l a s h on s i t e where i t was  inhibited  d i d not  have p r e v e n t e d t h e u n d e r s t o r y v e g e t a t i o n  o f t h e s e s t a n d s from leaving  sites  and  canopy  thinning  predicted  was  outcomes  Clearly,  t h e s e s t a n d s n e e d t o be  re-surveyed to  assess the a b i l i t y  of t h i n n i n g t o generate  desired  increases be  i n u n d e r s t o r y and  advisable to r e s t r i c t  assess  i t s potential  thinned wildlife The concern.  canopy volume.  cattle  impact.  stands without r e s t i n g  Without  regenerating,  of f i r e fire  I t would  Continued  g r a z i n g of  these  them may  not b e n e f i t  the  intended.  i n these stands  i s also  to prevent Douglas-fir  the t r e a t e d  also  g r a z i n g i n some a r e a s t o  f o r w h i c h t h e t r e a t m e n t was absence  to  stands w i l l  soon  of  from  revert to  their  33  previous  structure.  cycle of stagnation,  This w i l l  lead to a r e p e t i t i o n  insect infestation,  need f o r a d d i t i o n a l e x p e n s i v e t r e a t m e n t s . e n v i r o n m e n t a l and p o l i t i c a l fire  and t h e u l t i m a t e However,  concerns regarding  social,  t h e use o f  a s a management t o o l may make i n t e r v e n t i o n o f t h i s  kind d i f f i c u l t important, tool  of the  be  i f not impossible  (Bunnell  however, t h a t t h e u s e o f f i r e  explored.  1991).  Iti s  a s a management  34  4.  E f f e c t s o f t h i n n i n g on t h e f o r e s t b i r d  Habitat forest  structure  i s w i d e l y acknowledged t o e f f e c t  b i r d communities.  I t was p r e d i c t e d  that the  T h o m p s o n / N i c o l a Mule Deer F o r a g e and S l a s h i n g alter  four aspects of h a b i t a t s t r u c t u r e  stands:  1) h e r b and s h r u b c o v e r w o u l d  i n d i v i d u a l t r e e canopy volume w o u l d volume on a p e r h e c t a r e and  community  Project  i n these f o r e s t increase,  increase,  3) canopy  increase.  I examined p o t e n t i a l c h a n g e s i n t h e f o r e s t community due t o t h i s t h i n n i n g I examined b i r d  unthinned s i t e s . abundance  species, Third, spatial  i n three  community s t r u c t u r e  bird ways.  i n t h i n n e d and  Second, I e x a m i n e d how t h e r e l a t i v e  shrub-feeding  and woodpeckers)  and t h r e e  species,  species  groups  foliage-feeding  were a f f e c t e d by t h i n n i n g .  b e c a u s e t h e t r e a t m e n t a l t e r e d t h e q u a n t i t y and d i s t r i b u t i o n o f some h a b i t a t t y p e s w i t h i n - s i t e s , I  examined t h e a s s o c i a t i o n s spatial  project  of i n d i v i d u a l species  (ground- plus  2)  b a s i s w o u l d d e c r e a s e , and 4) down  d e a d woody m a t e r i a l w o u l d  First,  would  of i n d i v i d u a l species  d i s t r i b u t i o n s of four habitat  with the  types.  Methods Censusing Forest sites.  b i r d s were s a m p l e d i n 3 t h i n n e d  Because o f t i m e c o n s t r a i n t s , o n l y  t r e a t m e n t s and 2 i n c o n t r o l s )  were s a m p l e d  and 3 u n t h i n n e d  4 sites  (2 i n  i n 1990.  An  35  additional 1991  t r e a t m e n t and  season.  study s i t e , at  50-m  control  s i t e were added f o r t h e  A l l 6 s i t e s were 25 ha a g r i d was  intervals.  in size.  Within  s u r v e y e d w i t h s t a t i o n markers p l a c e d  B i r d s were s u r v e y e d u s i n g t h e s p o t -  mapping t e c h n i q u e as o u t l i n e d  by V e r n e r  v i s i t s were made t o e a c h  i n both  site  (1985).  1990  and  Eight 1991.  1990,  c e n s u s i n g began on J u n e 1 and  ended on J u l y  1991,  c e n s u s i n g began on May  ended on June 26.  12  each year 2 people performed participated hour  after  On  and  bird  during both y e a r s .  censusing.  e a c h day,  n o r when wind  1 t r e a t m e n t and  each census p e r i o d .  O b s e r v e r s walked  When p o s s i b l e ,  a.m.  end  of  lines  singing,  of simultaneous  Clusters  At  least  be c o n s i d e r e d a t e r r i t o r y .  territory  territory within  f o r each  3 detections  c e n s u s i n g p e r i o d s were r e q u i r e d  0.5  study  singing After  o f d e t e c t i o n s were t h e n u s e d  t h e number o f t e r r i t o r i e s  of each season.  nearest  20  d a t a were t r a n s p o s e d on t o summary s h e e t s  f o r each s p e c i e s . determine  No  i n each g r i d f o r  m a l e s and movements o f s i n g i n g m a l e s were r e c o r d e d . e a c h c e n s u s , raw  In  observer  b i r d s on maps o f each  locations  In  exceeded  along grid  d u r i n g c e n s u s i n g and marked t h e l o c a t i o n  site.  1.  1 c o n t r o l were s u r v e y e d .  location  and v i s u a l l y - d e t e c t e d  In  C e n s u s i n g began o n e - h a l f  in rain,  C e n s u s i n g began a t a d i f f e r e n t  calling,  One  s u n r i s e and d i d n o t e x t e n d p a s t 11:00  c e n s u s e s were c o n d u c t e d kph.  each  each  t o be c o n s i d e r e d a f u l l  species at the  on 3 d i f f e r e n t  for a cluster  Territories  to  of p o i n t s  to  were summed t o t h e  study s i t e . territory,  For a a l l of t h e  36  detections the  that  bounds o f the  considered  0.5  detections, be  species  territory,  For  or at l e a s t o n e - h a l f  a conservative  The  2 out  study  status  d i d not (e.g.,  forest bird  (those  s h r u b s >2-m red-oiser  h a b i t a t types w i l l  be  the  estimate  g r i d s were u s e d t o  tall  dogwood  referred to  as  m) 2  e a c h h a b i t a t t y p e was  then estimated  B e c a u s e o f the way the  map  habitat  and  require areas  with  willow,  (henceforth  these  d e n s e f o r e s t , open Percent cover  f o r e a c h 50x50-m  structure  r e s o l u t i o n of  and  f o r e s t , open  s u c h as  riparian, respectively).  of t h i n n i n g ,  included  i n the  a r e a s t h a t w o u l d not  D o u g l a s maple, and  result  were  I w a l k e d t h r o u g h each g r i d  open meadows ( o p e n i n g s >25  open and  detection  c l u s t e r s of  mapped a r e a s o f d e n s e - u n t h i n n e d c o n i f e r o u s  forest,  territories  maps  coarse habitat a t t r i b u t e s .  d e c i d u o u s t r e e s and  to  territories.  In each study s i t e ,  thinning)  had  f o r each  include  w h i c h were e x c l u d e d  Composite v e g e t a t i o n  forest  of 3  site.  maximum number o f t e r r i t o r i e s  o f minimum number of  within  be  simultaneous s i n g i n g detections  c l u s t e r s of d e t e c t i o n s  coniferous  to  detections  minimum number o f  estimate that  where no  to f a l l  number o f t e r r i t o r i e s  The  c l u s t e r s of questionable  had  a territory  of the  borders of the  were d e r i v e d .  recorded).  territory  a minimum o f  estimates of the  detections  the  study s i t e .  located within Two  was  identified  the  changed as  of  cell. a  h a b i t a t maps  37  differed sites, an  between t r e a t m e n t and c o n t r o l s .  open h a b i t a t  obvious j u x t a p o s i t i o n  juxtaposition forest is,  was c l e a r l y e v i d e n t of habitat  o f open h a b i t a t  habitats  types.  This  i n treatment s i t e s ;  the simplification of the habitat of small  as i t r e p r e s e n t e d  w i t h d e n s e f o r e s t , and open  was n o t a s . c l e a r  made t h e d e t e c t i o n  In untreated  i n treatment  sites  open meadows l e s s o b v i o u s .  a r e s u l t , composite v e g e t a t i o n  maps f o r t r e a t m e n t  d o m i n a t e d by dense f o r e s t h a b i t a t . was e a s i e r  comprised  to detect  In a d d i t i o n ,  i n controls  a larger proportion  and  As  sites  were d o m i n a t e d by open f o r e s t , w h e r e a s c o n t r o l p l o t s  habitat  that  were  open  therefore  o f t h e area than i n  treatments.  Data a n a l y s i s : To  Community  structure  identify potential differences  structure,  i n overall  I plotted dominance-diversity  c u r v e s and  examined t h e d i s t r i b u t i o n s o f b i r d s p e c i e s t r e a t m e n t s and c o n t r o l s .  I also  i n d i c e s t o examine t h e s i m i l a r i t y t r e a t m e n t and c o n t r o l p l o t s . coefficient  from s p e c i e s similarity  detections i n  calculated  lists,  using  two s i m i l a r i t y  o f b i r d communities i n  I used Jaccard's  t o t e s t the degree o f s i m i l a r i t y  t r e a t m e n t s and c o n t r o l s  similarity  between  simple presence-absence  and I u s e d M o r i s i t a ' s  detection  data  data  index o f  t o compare t h e d e g r e e o f s i m i l a r i t y  log-transformed species  community  b a s e d on  (Kr.ebs 1989) .  38  Data a n a l y s i s :  Relative  abundance  I u s e d two-way a n a l y s i s o f v a r i a n c e (treatment vs.  control)  and  year  (1990,  ' t r e a t m e n t ' v a r i a b l e s , t o examine t h e territories. density  Only 11  species  analysis. foraging  I placed strategies  foliage-feeders)  species  t o change as  s p a r r o w s , and for bird plus  result  shrub cover.  comprised breasted  The  species  Chipping-  II) . to  for  e a c h o f 22  Groundas  a n t i c i p a t e d increase  f o l i a g e - f e e d i n g group  Yellow-rumped warblers.  in  Red-  Foliage-  i n abundance thus,  habitat.  over each study  from the site  as  1991  abundance.  I used a l l types  detections  and  to avoid  visual)  r e s u l t s towards t h o s e d e t e c t i o n s  data  another  means t o a s s e s s r e l a t i v e (songs, c a l l s  a  was  Western tanagers,  number o f d e t e c t i o n s  species  names  increase  b e c a u s e o f the d e c r e a s e i n c a n o p y v o l u m e and  I summed the  The  Vesper  (all scientific  were e x p e c t e d t o d e c r e a s e  abundance o f f o r a g i n g  species  comprised of  were p r e d i c t e d  o f Mountain c h i c k a d e e s , n u t h a t c h e s , and  and  of t h i n n i n g .  i n Appendix  of t h i n n i n g because of the  h e r b and  feeding  species  statistical  American robins,  are p r e s e n t e d  shrub-feeding  sufficient  e f f e c t s on  a result  Townsend's s o l i t a r e s  species  bird  shrub-feeders  g r o u p was  s p a r r o w s , Dark-eyed j u n c o s ,  as  i n t o groups of s i m i l a r  for general  shrub-feeding  in  to warrant  (ground- p l u s  to t e s t  groupings predicted ground- plus  several  1991)  condition  number o f  occurred  (number o f t e r r i t o r i e s )  using  biasing  w h e r e b i r d s were most  of  39  conspicuous  (i.e.,  open a r e a s ) .  males s i n g i n g  I excluded  from t h i s  t h a t were t r a n s i e n t i n t h e i r Red  crossbills,  infrequently shinned One  (e.g.,  low  reason f o r t h i s  territories  t o examine t h e  I a l s o used t h e  later  that  siskins,  of each  g r o u p s n o t e d above p l u s , woodpeckers  a n a l y s i s to only  down m a t e r i a l .  variance  as  The Three-  I removed  woodpecker g r o u p t o that  restrict feed  I a l s o u s e d a one-way a n a l y s i s  t h a t used f o r d e t e c t i o n s  of  individual  on of  species.  Bird/habitat relationships  I sub-sampled t h e  50x50-m c e l l s  h a b i t a t maps i n t r e a t m e n t s and distribution  without  flickers,  Red-naped s a p s u c k e r s .  t h o s e woodpecker s p e c i e s  dead and  Data a n a l y s i s :  data.  woodpeckers, N o r t h e r n  Red-naped s a p s u c k e r s f r o m t h e  of  species  B l a c k - b a c k e d woodpeckers,  P i l e a t e d woodpeckers, and  or  sites.  (woodpeckers and  woodpeckers, H a i r y  size,  I u s e d one-way a n a l y s i s  species  woodpecker group i n c l u d e d  the  analysis  large t e r r i t o r y  number d e t e c t i o n s  two  t o examine  eluded  Red-naped s a p s u c k e r s ) t o a n a l y z e d e t e c t i o n  my  Pine  occurred  a n a l y s i s was  species  control  a d d i t i o n a l groups  toed  species  hawk, C o o p e r ' s hawk, S h a r p -  because of  between t r e a t m e n t and  two  or  (e.g.,  in  hawk).  numbers o f d e t e c t i o n s .  variance  behaviour  Red-tailed  posts  a n a l y s i s those  Evening grosbeaks),  frequency of occurrence of b a s e d on  from e l e v a t e d  of d e t e c t i o n s  of the  composite  c o n t r o l s t o examine  i n the  e x t r e m e s of e a c h  the habitat  40  type  (e.g. those c e l l s w i t h a h i g h p e r c e n t a g e  forest, that  open f o r e s t ,  examining  open, o r r i p a r i a n  c e l l s with high l e v e l s  h a b i t a t type best addressed h a b i t a t t y p e on b i r d  open and c e l l s  the potential  >9% f o r r i p a r i a n .  lists  The shapes  criteria  o r d e r e d t h e number a s dominance-  communities o c c u r  I used p r e s e n c e - a b s e n c e  of similarity  t h a t were p r e s e n t i n d i f f e r e n t  habitat  t r e a t m e n t s and c o n t r o l s  1989) .  (Krebs  a uni-variate  i n any o f t h e f o u r d a t a from s p e c i e s coefficient in between types  communities  within  and a m u l t i - v a r i a t e a n a l y s i s t o  examine t h e e f f e c t s o f w i t h i n - s i t e on b i r d d i s t r i b u t i o n s .  i n each  o f t h e s e c u r v e s were examined t o  to calculate Jaccard's s i m i l a r i t y  I used  i n each  Once c u t - o f f  The r e s u l t s were p l o t t e d  o r d e r t o t e s t t h e degree  In t h i s  number o f d e t e c t i o n s i n e a c h site.  retaining  I summed t h e number o f d e t e c t i o n s by  i fdistinct  habitat types.  cells  open f o r e s t and  c u r v e s t o examine community s t r u c t u r e  h a b i t a t type. determine  I chose  The numbers o f c e l l s  s p e c i e s f o r each h a b i t a t t y p e and r a n k  diversity  respective  effects of that  type while  h a b i t a t a r e summarized i n T a b l e 10.  of detections.  I assumed  T h i s c h o i c e generated t h e  habitat  foranalysis.  were e s t a b l i s h e d ,  o f each  f o r dense f o r e s t ,  g r e a t e s t extremes o f e a c h enough c e l l s  habitat).  species distribution.  t h a t h a d >59% c o v e r a g e  o f dense  vegetation variability  analysis,  50x50-m c e l l  I t h e n used t h e c o m p o s i t e  I summed t h e a t each  study  v e g e t a t i o n maps t o  41  Table  10.  Number o f c e l l s  Habitat type  Condition  i n each h a b i t a t type  % coverage i n each c e l l  # of cells  % of cells  Riparian  Control Treatment  >9 >9  15 7  5 2  Open  Control Treatment  >59 >59  18 6  6 2  Open f o r e s t  Control Treatment  >59 >59  50 271  16 90  Dense f o r e s t  Control Treatment  >59 >59  126 14  41 5  42  determine r e l a t i o n s h i p s of b i r d composition.  detections  to  cell  I used a C h i - s q u a r e a n a l y s i s t o t e s t i f b i r d  species  were a s s o c i a t e d  forest,  open f o r e s t , open, r i p a r i a n ) more t h a n would  e x p e c t e d by assigned  chance.  one  any  this  sum  of a l l h a b i t a t  Cover c l a s s e s w e r e :  occurred  to the p r o p o r t i o n  0=0%,  25%  dense c o v e r made up  the  detections  there  was  no  should  different  number o f  to the p r o p o r t i o n  for  results.  use  land  type  containing  1-  base, t h e n 50%  of  category class)..  o f any  base t h a t  the  be  percent  cover c l a s s e s species  f i t test  (dense  analysis  several  was  significant  however, were d e r i v e d  f i n d i n g suggests t h a t  w i t h c e l l s which had  This  type  separately.  Chi-square a n a l y s i s y i e l d e d Most of t h e s e ,  types  the  similar  each h a b i t a t  4 habitat  controls  If  h a b i t a t type,  t y p e would  open f o r e s t , open, r i p a r i a n ) .  intermediate This  land  i n that  i n each o f  done f o r t r e a t m e n t s and The  of  i n each h a b i t a t  of the  detections  I p e r f o r m e d a C h i - s q u a r e goodness o f  each s p e c i e s ,  forest,  occur  i f cells  the  cells  (spatial  (e.g.,  disproportionate  number o f d e t e c t i o n s  represented.  50%  for  1 = 1-25%, 2 =  i n which t h a t h a b i t a t  i n each s t u d y s i t e  type  c l a s s e s t o t a l e d 100%  h e t e r o g e n e i t y ) , I w e i g h t e d e x p e c t e d number o f according  be  was  f o r each h a b i t a t  Because the  e a c h h a b i t a t c a t e g o r y was  (dense  a n a l y s i s , each c e l l  26-50%, 3 = 51-75%, 4 = >75%. in  h a b i t a t type  of f i v e cover c l a s s e s  such t h a t the each c e l l .  For  with  (i.e., may  from  classes  have been  a p a r t i c u l a r combination of  1 and  2).  associated habitat  43  attributes.  Therefore,  discriminant  function analysis using  attempt to c l a s s i f y detected  The  evaluated b a s e d on  cells  single discriminant by  the  Wilks  lambda.  1  effectively.  The  (Nie  1983)  species  were  h a b i t a t types of  of the  function to  proportion  statistic techniques  separate  groups  i n d i c a t e d the  controls  Analysis  success  was  separately.  were c a l c u l a t e d t o t e s t f o r t h e  variance/covariance  each  of cases c o r r e c t l y c l a s s i f i e d  between-group c l a s s i f i c a t i o n . t r e a t m e n t s and  an  was  Chi-square  I used c l a s s i f i c a t i o n  in  detections  function derived  t o a c t u a l g r o u p membership  c o n d u c t e d on scores  the  s i g n i f i c a n c e of the  ability  of the  i n which b i r d  (absence) b a s e d on  t o assess the  according  SPSSx  ( p r e s e n c e ) compared t o t h o s e where no  were r e c o r d e d cell.  I used a one-group d i r e c t  Box  homogeneity of  M the  matrix.  Results Community  structure  D o m i n a n c e - d i v e r s i t y c u r v e s f o r c o n t r o l s and were v e r y  s i m i l a r (Figure  d o m i n a t e d by T h e r e was  Chipping  3).  abundant t o the  B o t h c o m m u n i t i e s were  sparrows, the  a p r e c i p i t o u s drop  this  next most a b u n d a n t s p e c i e s . less steeply  n i n t h most a b u n d a n t s p e c i e s .  group of e i g h t s p e c i e s  treatment  most abundant  sites,  species.  i n abundance from the  s t e e p drop both curves d e c l i n e d a f t e r the  treatments  i s the  although the  The  most  After and  this plateaued  composition  same i n c o n t r o l  rank order  and  i n abundance.,of  of  44 in CN  C Q) e rd  o o o  o o  o  o  o  o  o  o  o  o  o  o oo  o  o  o in  o  o n  o  o  o  (%)  sDuepunqe a A p e p j  p3ZTT.i2uiJON  45  these  s p e c i e s i s somewhat d i f f e r e n t  n i n t h most abundant s p e c i e s , b o t h  ( T a b l e 11).  After  communities are  the  comprised  o f many r a r e s p e c i e s . The  two  measurements o f community s i m i l a r i t y  i n d i c a t e t h a t the communities p r e s e n t c o n t r o l p l o t s were v e r y s i m i l a r . data,  Jaccard's  transformed was  0.96.  similarity)  Bird Of only had  detection data,  t o 1 (complete  similarity).  abundance:  Number o f 50  enough t o t e s t  exceptions.  similarity  r a n g e from 0  (Appendix  enough d e n s i t i e s  and  occurred  sparrows  The  t h e r e were  f o r these  start  of censusing  s p e c i e s was  T h e r e were no  (i.e.,  and  differences in years with  the  significantly  (minimum d e n s i t y e s t i m a t e )  in  III)  frequently  maximum d e n s i t y  d i f f e r e n c e s i n abundance  years  (no  (maximum d e n s i t y e s t i m a t e ) ,  (minimum and  V e s p e r sparrows  significant  between t h e two  I n 1991  more M o u n t a i n c h i c k a d e e s  territories  of  log-  territories  i n high  There were no  t h e number o f t e r r i t o r i e s  (Table 12).  Using  f o r d i f f e r e n c e s between t r e a t m e n t  control plots.  and  0.92.  species recorded  s m a l l enough t e r r i t o r i e s )  and  presence/absence  M o r i s i t a ' s index i n d i c e s can  11 s p e c i e s o c c u r r e d  Chipping  was  These s i m i l a r i t y  t h e more t h a n  following  i n treatment  Using  index o f s i m i l a r i t y  also  likely  estimate) than  in  1990  between  because of the  early  1991.  significant  d i f f e r e n c e s i n t h e number  f o r e i t h e r maximum o r minimum d e n s i t y  of  estimates  46  o c o 0)  iH O O O  -P  o a) -p  VO CM CM CM H  H  iH  0)  -a QJ  U  a)  rH  a) XI <u rH 03  QJ  u a)  rH  u  rH  a)  r-l  X!  -p 0)  u rH fd tPX! X o QJ O rH rH C Ul -p o o O (d 0) 3 u •H 3 ft a) • i H O 0) rH X! Ul 0) T 3 r * rH o C rH £ O ft o O ft o o ^ o C P 0) X! a) T 3 o fd C T 3 - P «3 a) - h fd - H a) 3 T S •H O -P ftxi a-r-« a) XI Ul VH id Ul ftrH T3 rH O c a) ui u e T3 <4H 0) - H O rrj O -P o C Ul ^ > s TS (0 3 T 3 Ul >ITJ o S-l Ul ft 0) - P s fj> C VH 0) fd c -a Ul rH Q) o c o 0 o c C - h i >i a) fd c MH ft 5 5H XI rH a) •p c MO I (d - P 1 u l-H (0 S QJ ^ U <D 0) 0 Ul rV -P (0 0) 0) r j > 0) >i c ft-P O I X! 1 C QJ - P C rH - P ft* i a) a c rH ^ I • H iH r l Ul IH Ul Ul T 3 XI fd •H ' 3 rH rH TS Q) 3 a) fd o rH O --H C QJ JH < E H > a) (0 a) K 5 5 a X! 0 o CQ W fc U S >H Q «  -P  fd  EH  a)  U  -P C 0)  QJ •H  o  m ui o c o U -H a) - P  in  r> CO o CM in o CM CM H H H CM  (N  co  QJ  o vo  CO  rH  ft Tf  0)  CM rH t> CO cn r> CO CO CO CM CM rH H H  rH  O  QJ  rH  QJ X! rH rH  •H  (0  >> CT> Ul  c c H o  >1  X)  H H  O CO rH  O H  Ul rH rH (0 -iH  rH  X!  rH 0)  U -P (0  5 O  rH  X! QJ rH QJ  XI -P 3  QJ QJ rX  rH  O  (0  X3  S  fd  U  rH  •H o ft w  rH  rH  QJ  (0 -P  >1-  >i  rH >1  rH T3 fd rH  & o  rH  <  Q  -P  ft  (d T5 rH rH  C H - O ui a) - P O ft c O C VH >i (d <4H C C QJ QJ id O QJ rH (d u > rH fd • H rH I O -P QJ C Q) aj - P ftX! ftUl I - P >ix: •H ft I rH H 3 U l Ti •H T3 X. C QJ a) Ul VH - P rH Xi QJ id (D O U QJ Ul > > « QJ (d o O U « Q >H S & ffi z e 3 o CP fd  C  O  rH QJ 0) r * rH rH O QJ  O  X! QJ  3 rH QJ QJ 0) •H XJ rH £ Ul QJ X X O O O ft o o> o o QJ •p Ul rH fd (d QJ -H rH Ul (d C ftrH o Ul rd (d T 3 tn ft-0  U ^ C C O u U 3 0) H id TS •r-iftx! ft QJ O Ul 4-> Ul QJ  01 0)  rH  QJ  ^ o  CP 0) X C ft O  rH  QJ • H  —i  TS  XI  QJ  rH  T3 QJ  QJ  rH  s  •H >  C  c •H H  S Q) id  o  O  O O  O oI XI rd ui -  rH  QJ  I  &X O C O Ul fd rH XI rH U rH -H 3 JC  Id  OH  XI  rH  id  EH  EH  HcMcotj-invDr^cocriOrH ( M n - j m i O M o m o H N H H H H H H H r l r M N N  BUI •P  0.25  0.25  A  A  A  0.25  0.25  V in o o  0.25  :p<0.  a  . i<  >0.25  A  A  A  o  c as H C CT Ti i H  CM CO CM co H cn CM  CM CO H CO CO in in rH CO CO rH CO  O CTi cn rH  o  CO CO  ax  CO CO >i  V in o o  o v a  TS  fi fi •H  -cH  *—*  0.25  >i  -P  in (—%  :p<0.  i a  in CM o  si  c •i-i co 0) -p  >0.25  •valu  con trols  CO  in CM o  S  fi  cn co cn  CO C O cn CO CO rH  00  CM CM  in VD VD rH  rH  O rH  >0.25  c CO TJ  >0.25  •P  •rH 10  fi  -H  fi CO a  (0 O <4H T )  fi fi •H  CO - H CO X I •rH U >i 0 Si •p -rH H JH cn  G •iH  SH  SH 0)  £  CTI H  T3  C  <W O (0  0) X!  5<p<0.  in CM o  >0.25  o  >0.25  >0.25  >0. 25  o  o o  VO CO  CO CO CO CM  CM CO H  CM CO  o cn cn  VD H cn VO VD CO in in o\ <* CM C M r~ CM CM CO i-H CM o rH H  iH  H  CO CM CM CM  H  rH t>  rH  cn  O  Cn cn  e H c •c H c (0 C O CO - P s c CO fi • -p CM rrj rH CO SH C O -P rH XI  •  in o  cn cn rH  rH  -P  SH  o o  a  rH  Kp<0.  0  in CM o  25<p<0  -H  •value  CO C -H  •H  density  mum s.  and  fi  5<p<0.  (0  >0.25  X  TS  rd C H rd  SH  X!  O -P  > 0 C •H XI  o  $H  CO CO  -H 0  CO a CO  c rd  0 -H  >H  0)  fi  <  SH  rd co tn  C -H  fi a  •H X!  U  CD rd CD X I -P U 0 0) rd P O X! X c 0 O C 0) o • H •P TS SH P •rH X ! •l—i rd as O O -p > TS >i co CO rH • CH rd > 1 4H >i cu SH CO rd SH rd X I P •P 1 >l -H l X X C rH SH TS CO rd p o CD o a a 2 CO TS  CO  cu rd  rH  rH  XI  >H  -P  •rH rH  o CO -CO TS  C CO CO c o  EH  SH  o u u  rd a CO  0) IT  rd C rd  •P  rd TS  0)  a fi  P SH  1 QJ CO rH o a - P rH CO CO CO CO CD u  >  C  SH  48  between t h i n n e d and u n t h i n n e d  sites  s p e c i e s examined  Furthermore,  difference control affected  i n t h e number o f t e r r i t o r i e s  sites  of the s p e c i e s t h a t  by t h i s  feeding-species, were a l m o s t 14).  (Table 13).  f o r any o f t h e 11  treatment.  i n treatment or  Densities  f o r t h e ground-  ground-feeding  i n t r e a t m e n t and c o n t r o l  S i m i l a r l y , t h e r e was no s i g n i f i c a n t  number o f t e r r i t o r i e s control  group,  sites  (Table  difference  i n the  of the foliage-feeding  and t r e a t m e n t s i t e s  little  were e x p e c t e d t o be most  and f o r t h e e n t i r e  identical  t h e r e was  group between  (Table 15).  When a l l s p e c i e s were c o n s i d e r e d t o g e t h e r t h e r e no significant control estimate  difference  sites  b e t w e e n t h e number o f t e r r i t o r i e s i n  than i n t r e a t e d  sites  for either  (-^minimum number o f t e r r i t o r i e s  control  •"-minimum number o f t e r r i t o r i e s  treatment  =  ^maximum number o f t e r r i t o r i e s  control  70.5,  number o f t e r r i t o r i e s was a l s o  and  differences  for either  number o f t e r r i t o r i e s treatment  control =  number o f t e r r i t o r i e s territories  52.3, p >0.2 5; ^maximum  i n the total  There  number o f  o f t h e 11 most a b u n d a n t s p e c i e s between c o n t r o l  treatment s i t e s  territories  58.5,  =  t r e a t m e n t ~ 68.4, p >0.25).  no s i g n i f i c a n t  territories  =  density  treatment  4 7  estimate  51.8,  ^minimum  - 2 , 0.25 < p <0.1;  control =  =  density  =  63.1,  x  (^minimum number o f  ^maximum  maximum number o f  57.6, 0.25 < p < 0 . 1 ) .  49  Ul rH  o  OJ 3  u  rH fd > i  -p  c o o  ft  -rH  in  •  o A  ft  in CM  in CM  in CM  in CM  O V  in CM  in CM  in CM  O V  in CM  in CM  O A  o A  O A  O A  V in CM  o A  o A  o A  V in CM  o A  o A  •  ft  •  ft  •  ft  •  ft  Ul  OJ 4-> «J g •H  c OJ g  -P  3  Ul  g •H  OJ  >i  X  •H  QJ g •P  id OJ rH  CM  in  CO  vo  CO  o>  CO  CO  CO  o  CO  o  CO  en  CTl  OJ  in CM • o A  in CM • o A  in CM • o A  C  3 rH  id  fd  g  > I  ft  3  g •H  O  r-  o  CO  CM  CO  CM  rH  co  in  en  VO  CTl  CTl  in  CTl  CO  in  rH  CM  CM  H  in CM • o A  in CM • o A  in CM • o A  CTl  o  in  ft  ft  ft  O V  rH  ft  V in CM  in CM • o A  ft  O  o V  in CM • o A  in CM • o A  ft  V in CM  rH  ft  ft  ft  o  O V  ft  V in CM  ft  ft  O  -P  c  •H Ul  •H g  C OJ  o  TS  <W  g  3  Ul QJ •H rH  g -H  -P C  OJ g •p fd OJ rH  in  CO  CM  in  CO  H  in  co  CM  VO  H  CTl  CM  CO  co  H  CM  CM  O  VO  CTl  VO  CM  H  H  rH  EH  c  o  -p • • H Ul rH QJ rH - H 0) O - P 0) ft 4H Ul O  T3  rH rH OJ - H  -H  a  o  o  rH •P  G  CM  ui C Q)  fd  «H  Q)  Q)  -P  rH rH Ti  C  fd  u u  u  Ul  Ul QJ -H  0  u  ft W  0  •H  c id u -H  QJ  CTl  CO  u  XI  c X! 0  g  co  H  0  fd - P  -P  H CO  u  >i  •  CO CO  H  0  XI  EH  O  EH  H  T(  fd  ft  o  u  fd g  X!  o  H  X  QJ  ft  •  c  •P C  g g •H  a  •  -P  U  -a  C  ft  ft  O  OJ  C  ft  •  a  Ul  3  ft  •  fd  •P  g  •  o  -P  ui  ft  QJ g  <  fd  ft  D> C •H  0 0  c 3  •r-i  n  QJ >i QJ  U QJ  QJ QJ TS fd  X XI 0 0 -P  fd O  >i  rH <W  -rH  XI 0 c -H rd -p  X! -P 3 C  QJ  -P  Ul rd QJ rH  a 1 >1 X! a X XUl c 1 3 T3  -H  XI u  u rd  Q  3 Q  o  a  QJ Oh  XJ  -P  0  U >  Ul  -H  rH id -P  •H rH O  in  u  -H rH  o Ul  QJ  T3  QJ rH  QJ rH rd  -p rd  -  T3  C QJ Ul C  > o  EH  0  rH rH rd  a Ul  U QJ  a Ul  QJ  >  rH fd  QJ  CP fd  c fd -p  c rH OJ •p Ul Q)  QJ  ft g  3  5H  1 o rH rH QJ  ><  rH rH  50  CO  fi  3  3  rH (0 > I  fi -H X  (0  a  e TJ  ID ID ID ID ID CN  CN  CM  CM  CN  ID  O  O  O  O  O  O  a  a  a  a  a  a  CM  ID  VO  O  r>  CN  OA  CO  CM  CO  CT\ ID  •tf  CM  CM  r>  A  A  A  A  A  CM A  •P  c  -H CO  rd  C CO  fi 3 fi  •0  fi 3 fi  •H C  •H fi  -H  •P  c QJ e -p (0  CO  QJ  H  U EH  X  rd  -P  a  o  XI  O  rH  u  -P C  o  co o co cr>  o u  <tH  CO SH CO  TJ CO CO  QJ  4H  3  rH (0 > I  c  CO 3 O >H - P C" CO  C  «H  fi  O  -P  (0 CO CO CO rH •H - P >H O Tf •P  •H  C  td  rH rH CO •P  <n  a  ID CM  ID CM  ID CM  ID CN  o A  O A  O A  O A  O A  a  a  a  a  a  c CO TJ  fi 3 fi  •H  c  CJ rH  rQ o u fi  •p c QJ fi -p rd  CO  CO  VO  rH  H  CM  ID  t>  CM  CM  VO  QJ  rH EH  a  rH  o  r~  o  •* vo cn  co  JH -P C  CM  r>  r>  vo  U td  -P  c •H  a-H  •p  -p CO - H  H  CO  X!  C CO TJ  rd EH  X!  •P  O  •H  SH  o o  u rd  U a CO  CO  O  (0 CO >,  H  CO  rd (0 CO fi  H  CM  o u  0)  •  V V \V in CM  ID  CO  3 C C  V a o  -P  -H CO  •  •H O JH -P C  o o  JH CO  s*  LO CM  co QJ  •H  o QJ a  co  c  rd O  •H SH QJ  fi  <  tT>  c  TS QJ >i  SH SH  rd Tl  a CO  CO  SH  a a x  I  0)  SH  CO  •H  X! CJ  rd Q  a QJ  >  <  i  51  QJ  3 rH rd  >i  > I  -P -H  Ul  -P  c  •H  OJ TS  C  Ul OJ  g  TS  3  g  g  -H  3  X  g  rd g  •rH  IT) (M o A  ft  o  V  O V  v in  v in  CM  CM  in CM  o A DH  -P C Q)  g  -P rd Q)  "t  o  o  n  ^  H  to  rH EH  X  rd  TS  c  a O  (Ti  -p c o u  • co  • in  rH  OJ  in  3 rH  CM  O V  o v  v in  v in  CM  CM  o  o  u  rd g g  •  o H  •  X3 •P O  43 iH  o  rd > I  4H  a)  •H  •  0 ui a) OJ ft+j ui - H ui cn  c -p  •H TS OJ OJ  C OJ  g -P  <M rd  1  OJ  rd  OJ  u  -H  rH  TS C  O  rd  <(H  <w O  a  -P •H  Ul  Ul  c TS  g  3 g •H  c  Ul C OJ  a m  g  in  CO  cr>  H  rd  co  n  o  t>  •p QJ !H EH  •H  a O SH  4->  rH  vo  CO  H  CO  C  O U  O  OJ  0) TS  *  rd  0  rH o  •H  MH Q)  C  Ul  Ul  QJ  rd g  X?  -P  OJ •H U OJ  •r<H d  -P 3 C  0)  •P  Ul rd 0) SH  X!  0  TS Q)  a  0J  r0) H Cn  TS  -p  c 3  rH  Xi 0 -p rd Xi  U  X! 0  OJ  a  OJ  -P  EH  o A  c  rH  rd Ul  CM  •P  rH •p >i c •P o  -H  o A  in  1  rd C rd  •P  rH X! rrdH TS  0J a g  3  SH C  r0)H  -P  Ul 0J 3=  1  3! O  rH rH 0J >H  52  B i r d abundance:  Number o f d e t e c t i o n s  A s i g n i f i c a n t d i f f e r e n c e i n t h e number o f d e t e c t i o n s was  found  f o r o n l y one s p e c i e s  chickadees  (Table 16).  Mountain  were d e t e c t e d more o f t e n i n t r e a t m e n t  i n c o n t r o l s (0.05 < p < 0.025).  sites  Mountain chickadees  than  were  t h e o n l y s p e c i e s o f t h e f o l i a g e - f e e d i n g g r o u p t o be d e t e c t e d more o f t e n i n t r e a t e d s i t e s .  The o t h e r  species of  t h i s g r o u p , and t h e g r o u p a s a whole were n o t d e t e c t e d s i g n i f i c a n t l y more o f t e n i n c o n t r o l s i t e s sites  than  i n treatment  ( T a b l e 16 and 1 7 ) . Individual ground-feeding  equally  i n treatment  s p e c i e s were  and c o n t r o l s i t e s  Furthermore, the ground-feeding equally  i n treatment  Finally, equally  detected  (Table 16).  s p e c i e s were  and c o n t r o l s i t e s  detected  (Table 17).  each o f t h e s p e c i e s o f w o o d p e c k e r s was d e t e c t e d i n treatments  a group these treatments  and c o n t r o l s .  When t a k e n  s i x s p e c i e s were a l s o d e t e c t e d  and c o n t r o l s ( T a b l e  when Red-naped s a p s u c k e r s  17).  together as  equally i n  However,  were removed f r o m t h i s g r o u p a n d  t h e a n a l y s i s was r e d o n e , t h e woodpecker g r o u p was d e t e c t e d s i g n i f i c a n t l y more o f t e n treatment  sites  ( b u t o n l y m a r g i n a l l y so)  in  (0.1 < p < 0.05; T a b l e 1 7 ) .  Bird/habitat  relationships:  C o m p o s i t e maps  In c o n t r o l s , t h e most common h a b i t a t was dense f o l l o w e d by open f o r e s t ,  open a n d r i p a r i a n .  open f o r e s t was t h e most common h a b i t a t t y p e ,  forest  In treatments, f o l l o w e d by  53  T a b l e 16. Mean number o f d e t e c t i o n s c o n t r o l and t r e a t m e n t s i t e s .  Species  Control  Black-backed Three-toed Hairy  woodpecker  woodpecker  woodpecker  f o r 22 s p e c i e s i n  Treatment  p-value  7. 3  13 .3  p >0.25  2. 7  7 .7  p >0.25  10. 7  15. 3  p >0.25  10. 3  14 . 0  p >0.25  Northern  flicker  Pileated  woodpecker  3 .3  10. 0  sapsucker  16. 0  22 . 3  p >0.25  59. 3  84 . 0  0.05<p<0.0  nuthatch  109 . 7  64 . 7  0. 25<p<0.  warbler  75 . 7  75. 3  11. 3  3 .3  225. 0  217 . 7  p >0.25  80. 7  74 . 7  p >0.25  V e s p e r sparrow  22 . 3  27 . 3  p >0.25  American  50. 0  49. 3  p >0.25  35. 3  42 . 7  p >0.25  35. 7  21. 3  0. 25<p<0.  Swainson's t h r u s h  3 .3  0. 0  p >0.25  Ruby-crowned  4 .7  1. 0  p >0.25  Red-naped Mountain  chickadee  Red-breasted Yellow-rumped Western  tanager  C h i p p i n g sparrow Dark-eyed  junco robin  Townsend's  Dusky  solitare  flycatcher kinglet  0.25<p<0.  p >0.25 0. 25<p<0.  Warbling  vireo  1. 0  1. 7  p >0.25  Solitary  vireo  9 .0  12 . 0  p >0.25  Orange-crowned w a r b l e r  7 .7  1. 3  p >0.25  Wilson's  5 .7  0. 0  warbler  0.25<p<0.  54  T a b l e 17. Number o f d e t e c t i o n s c o n t r o l and t r e a t m e n t s i t e s .  Species  Control  of four  species  groups i n  Treatment  p-value  Foliage-feeders  64.0  56.8  p >0.25  Ground-feeders  82.7  82.3  p >0.25  Woodpeckers  8.4  13.8  0.25<p<0.1  Woodpeckers minus sapsuckers  6.9  12.1  0.1<p<.0.05  55  dense f o r e s t , h a b i t a t was  open, and  patchy  d e t e c t d i s c r e t e openings  The  and  Open h a b i t a t was  s i t e s than expected  h a b i t a t was  (Table 18).  i n both treatment  more so i n c o n t r o l s . treatment  riparian  because  extremely r a r e  Smaller, i s o l a t e d  o t h e r c o n t r o l p l o t and  one  l e s s common i n inability  sites.  a r e a s were f o u n d plot.  especially  controls.  The  i n one  large the  when combined  with  another. Bird/habitat relationships: The  Bird  communities  d o m i n a n c e - d i v e r s i t y c u r v e s f o r each h a b i t a t  i n c o n t r o l s i t e s are p r e s e n t e d dense f o r e s t ,  open f o r e s t  and  i n F i g u r e 4. open h a b i t a t s  community  sparrow) .  i s dominated  There  also  by  one  show a  from  After  o f many r a r e  t o the  t h e n i n t h most abundant level  s p e c i e s ( T a b l e s 19-21) .  h a b i t a t d e v i a t e s somewhat f r o m decline  T h i s drop i s  t h e second  s p e c i e s the curves f o r a l l t h r e e communities are comprised  (Figure 3).  f r o m t h e most  abundant t o t h e n e x t most a b u n d a n t s p e c i e s .  n i n t h most abundant s p e c i e s .  similar  s p e c i e s (Chipping  i s a sharp drop  f o l l o w e d by a more g e n t l e d e c l i n e  type  Communities i n  pattern to overall dominance-diversity curves Each  to  Riparian  standard d e v i a t i o n of a l l e s t i m a t e s i l l u s t r a t e s  one  but  o c c u r r e d i n one  treatment  h e t e r o g e n e i t y of these s i t e s ,  forest  sites,  i n b o t h t r e a t m e n t s and  main p o r t i o n o f r i p a r i a n h a b i t a t  control plot.  control  o f my  i n treatment  Dense  this  pattern.  The  o f f and Open initial  from t h e most a b u n d a n t s p e c i e s i s s t e e p e r , and  the  56  T a b l e 18. A v e r a g e p e r c e n t c o v e r a g e o f e a c h h a b i t a t t y p e i n e a c h 0.25 ha c e l l f o r c o n t r o l s and t r e a t m e n t s . Control  Treatment  Habitat  Mean  (SE)  Dense  49.9  (14.5)  9.3  (11-4)  Open f o r e s t  27.5  (14.5)  86.9  (13.5)  Open  21.3  (10.7)  3.5  (7.1)  1.3  (3.1)  0.3  (1-4)  forest  Riparian  Mean  (SE)  (%)  SDuepunqE  BAiqeiBi  pazTXBULZON  58  to c o  rH (0 -P  -H -p  o  0) -p 0) TJ • CO <4-l CO O - P  •H 5-1 CO Q) XI-P  e  c  3 CO C fi -P  H  (0 rd CO  o  EH  oV CO CO O  <4H  o  CO fi -p fO CO  CO C  CO *Xi  c  SH  EH  D>o rd o  -p CXi CO-P 0 O  SH  CO SH CO -P SH SH CO XI CO CO - P CO COrH SH X! as u cu<a rd X l CO SH * * tr> rdTSxl CO U 5 CO ft SH S (0 SH O O C -P O - P TS SH SH CO 3TS CO co d C O - H O 3 O rd CO o 0 ) * T S SH O O r H 0 * X ! C H S C O * CO C u * SH C P O COX! O co x i CO O ftTS SHTS C O O - H O (0 CO C - P > SH SH S H - H rd O T ) rd CO 3 - H . p X t C O T l T S C ftS rH (0 ftftrnx! rd O CO CO-HrH CO O CO SH CO fi O O U t f l * - P >MH £ C rd rdTS O 10T3 UC O 3 TS > i - O C f l TS S ft-P O CJISH CO C r H C - O rd rd > i C COTS O to O 0 c o cn w C I >i-HiwrdCXICOSHSHftCOSH C> 1 O H - >C CO 10 I H C • H S CO rd OCOISnrdCOrd+JOSHSH ft O I 4-> > r H CO * XI - P X ! C rd I C0C0>,cnCC0rHO C V H * C * rH C O I - H - P I CO >ift4-> SH C - H COX! CO •HrHMPCOCO^fdTirHSHTSrHXitfltO-HrdrdM^rH X l C O r d O d f i O r H C O O O CO-H d 0) CO rd SH > X ! rd-H SH  o o  CO  •P  O O SH -P  H  SH  IT) •P  rH  OCnr^r^VOin'l'COCOCnCMCMrHrHrHrHOOOOOO  o\<>  -P  4H  H^COCNCTiCOCOr^Htn^COCOOHOOOOOOO  CO  X! fi  0-P  rH  inoHco^'tfrHLninrHcocoriocMOOooooo O rH  • P SH  TS CO C XJ rd  nvO'JOJ^COCOO^'MrMHOHvMOlO^OriOO ID CM CM ID VO -tf CM •tf CTi co "tf vo co CO H ^ CM VO CM CM CM rH rH H  s  CO CO •<H CJ CO Si CO  UXi CO  >s  OaC  TS  -H  C + J  (0 (0  (0 +J  •P  CO-H CXI O (0 •HXJ 4J 04-> CO CO • P CO CO >H TS O <w MH O CO  to  C CO CO X!TS fi 3 c C-H SH  -TS CO >-l TS S-i >-l >H CO  o o o * o c  o  EH  incricninLnr^HvocMcoocT\cM>tfin<^'^ocr>cooo H H H rH rH rH CM rH CM CM CM CM rH H H  CO O  CO  4H O  X! fi 55  SH  -P C O  CO CO  X! -p  CO  CO H X!  TS SH O U  tO CO EH S-I  inM^cocovornr^HcriMn^^rnNHHHHOO OVOtCOCMCMCMHrH  o  a  CO  * c rd  SH  cu X  X! 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SH C ft* C o > H - P I C OJ > r H - H 0JX5-P Ul ft* I fdrHXirHfdrHrHUlrH • H S n T i r H Q) d Ul U) ^ f d - H SHTS X ! fd Q) 0) 6 O fjj d O r H fd o OJ J - I - H d o S J X I fd Q J - H UQf^^i<S>QHfX!a:S(^OCMf^WWEHSS3i  P  u •p c o U  CM'd'rHrHOinincMCMrHrHrHtHOOOOOOOOO 'J' H H H H  -P  U  HcMco^invor>a)O^OrHCN]co^invor>a)CT>OrHCM rHHrHrHrHrHrHrHrHrHCMCMCM  61  o -<op EH  o< \>  0H 3 r rH rH CO 0) o <4H X! o  CM  P C C D g P ( 0 <D U  EH  n^^H^MOvDcovotNMO^Hnooonooo in vo cn vo CM in co CN •* co "a* co CM CM H VD  H  CM CM  CM H  H  inHioinh^on^ri'j^oooooooooo CM'tfrOrHCMCMOrHiH'tfCNCMOOOOOOOOOO rH H rH  vocTir>r^vovoconcNCMCMrHOOOOOOOOOO  g  u  z  CO rH * CO OrH4J  u  rH  O rHXt ftHrH COC * (0u CO C OrHC O rH O TS r0 & > CO* rH X ! (0 C OuCO n o co H O COrH o ( 0 co C^*CO S C*O SO T l -H* O SHd-HTS CO D \ Q rH O rH ( ffl+Jrfj rrj roj * . a 0C • H O O O C O C O X i O O 0 3 C O X ! 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CO 0 r H) +JCO O 0) CO rH *CO COO SH X P -H C O S n r H r f l X ! SH(0 CO* x iO C ftCO C O ( 0 0\C O TO S*O T S 5 C-P ( 0 O 0 3 ft C OO r"HrHO *O 03ftCOXl O C O OC OOST3 * ftS Xi rH OC*TS*C C n O O O O C H O CO T 5 X I 03ft03-H C O (0 U-P 0 5COM TS-O H UO SH ro 3X1-HC TQ)ST0 S)COftrO H C TS(-H rH rO Op ftr-iOXi > 0 3 T S > 0 3 ( 0 > * < W £ (0 03TS O "S C P g > O T I P C O O ft > • 0 3 3 C O >iC O CnrO CTS 03 r H CP a) C CO (0 03 O 4H C >i(0-H CO - C 5 ft>H O C-P CX! SH CO (0r003>Hl-rHl C OSH -H CO O ( U S c > i C-P CCH 0C O rH*C XO !X r! OCP 0 >-tft -H p0o~iCO O X! O S H I H p O P 0) ft * ft* G C >i O0 3H 03-H SnCOSrCXiTSrHr H Hr003rHrHfOSHrH03 dxirCgOSHpCOQ) rOTSr CO O £ COX! (OrH O-H CO SH  SH  in  SH  *  SH  O ^H -cp 03 CO O •H CJ O CO ft  CO  SH  S H - P SH  >  (0  -H  3  ^  <D-H  OJ 3  SH  3SHTS  O  CO CO  SH  SH  SH  SH  I  SH  TS  I  SH  I I  rH  QUQp<2O««>tStHffi«WC0SEHSmZP-.>  * c  (0  icMco^invor^cocriOrHCNco'tfinvDr>coCT\OrHcN HrHrHrHrHrHrHrHrHrHCMCMCN  62  plateau  i n abundance o c c u r s e a r l i e r .  s u p p o r t s fewer s p e c i e s Riparian other one  habitat  habitats.  species  Second, t h e flycatchers, species.  than the  this habitat  same e x t e n t  rank o r d e r not  three  habitat  habitats.  as  i s not  the  dominated  other  three  s u c h as  by  habitats.  sparrows are  the  most abundant  Orange-crowned w a r b l e r ,  Ruby-  crowned k i n g l e t , W i l s o n ' s w a r b l e r a r e more abundant i n habitat  (Table  22).  decline  i n species  Finally,  there  i s a more  community  h a b i t a t c l o s e l y mimics t h a t dominance-diversity  curve  i n the  f o r the  (Figure  5).  c o v e r a g e make up  90%  followed  of the  in controls, there  by  total  are  the  decline  d o m i n a t e d by forest species  number o f c e l l s  from the  A f t e r the  species  species  communities were abundance was  h a b i t a t t h a n any  to the (Tables  other  three  the  and  20) .  open h a b i t a t were  and  l e s s pronounced  of the  most  is  levels off  same e x t e n t 19  This  second t o  (Table  i n both dense f o r e s t and one  (Table  n i n t h most abundant  c u r v e f o r open f o r e s t h a b i t a t of many r a r e  not  % open f o r e s t  next most a b u n d a n t s p e c i e s .  a more g e n t l e  i s made up  Communities  >59  is  i s a s h a r p d r o p from t h e  n i n t h most abundant s p e c i e s . species  open f o r e s t  This  that those c e l l s with  a b u n d a n t t o the  gradual  over a l l treatment  s u r p r i s i n g given  As  this  abundance i n r i p a r i a n h a b i t a t .  In treatments, the  10).  the  o f abundance i s d i f f e r e n t ; Dusky  Chipping  Species  other  open  i n c o n t r o l s d i f f e r s m a r k e d l y from  First,  t o the  Further,  21) .  that The  open decline  i n dense f o r e s t  habitat  not  types.  in  63  in CN  64  Riparian habitat species  and open h a b i t a t  (Table 22).  specific  community  supported the least  There i s l e s s  evidence of a r i p a r i a n  i n treatments than i n controls,  due t o t h e s c a r c i t y o f t h i s h a b i t a t  likely  i n treatments (Table  18) .  Bird/habitat  relationships:  In  communities i n dense f o r e s t  controls,  habitat  were t h e most  (Table 23).  similar  Within plot analysis  and r i p a r i a n  o f a l l p a i r w i s e comparisons  These two c o m m u n i t i e s were t h e most  because c e l l s with r i p a r i a n h a b i t a t amount o f dense f o r e s t i n dense f o r e s t  habitat.  habitat  usually  similar  contain a  As a r e s u l t ,  large  communities  c o n t a i n many o f t h e s p e c i e s  a s s o c i a t e d with r i p a r i a n h a b i t a t . forest  similarity  and dense f o r e s t  habitat  Communities  were a l s o  i n open  quite  similar  b e c a u s e o f t h e s h a r e d p r e s e n c e o f some o f t h e s p e c i e s a s s o c i a t e d w i t h more open h a b i t a t  (e.g.Northern f l i c k e r ,  and V e s p e r sparrow) and t h e s h a r e d a b s e n c e o f some o f t h e riparian  species  dissimilar forest Dense  (e.g. Ruby-crowned  pairings  and open  and open h a b i t a t  and r i p a r i a n  were d i s s i m i l a r  b e c a u s e many o f t h e s p e c i e s n o t e d a s o c c u r r i n g h a b i t a t s were d e t e c t e d open h a b i t a t s  The two most  o f c o m m u n i t i e s were t h o s e between  and open h a b i t a t , forest  kinglet).  i n dense f o r e s t  (e.g., O r a n g e - c r o w n e d  warbler,  Swainson's t h r u s h ,  riparian  h a b i t a t s were t h e l e a s t  dense  habitat. primarily i n riparian  habitat,  but not i n  warbler, Wilson's  and W a r b l i n g v i r e o ) . similar  Open and  communities  65  T a b l e 23. S i m i l a r i t y i n d i c e s f o r communities i n each h a b i t a t t y p e based on p r e s e n c e / a b s e n c e d a t a u s i n g J a c c a r d ' s s i m i l a r i t y index. Controls Dense forest  Open forest  Open  Dense f o r e s t  1.00  Open f o r e s t  0.80  1. 00  Open  0.65  0.71  1. 00  Riparian  0.81  0.70  0.48  Riparian  1. 00  Treatments Dense forest  Open forest  Open  Dense f o r e s t  1.00  Open f o r e s t  0.8 3  1. 00  Open  0.7 5  0.71  1.00  Riparian  0.69  0.71  0.72  Riparian  1.00  66  because a l l of the species p r i m a r i l y riparian  habitat  were a b s e n t  i n open h a b i t a t ,  s p e c i e s a s s o c i a t e d w i t h open h a b i t a t riparian  habitat.  relatively  associated  Open f o r e s t  with  and t h o s e  were a b s e n t i n  and r i p a r i a n  s i m i l a r due t o t h e s p i l l i n g  habitat  were  over of r i p a r i a n  s p e c i e s s u c h as W i l s o n ' s and O r a n g e - c r o w n e d w a r b l e r s cells  w i t h h i g h amounts o f open f o r e s t  forest  reasons.  communities contained s p e c i e s t h a t riparian In  communities riparian,  would u s u a l l y  Dense f o r e s t  i n dense f o r e s t  open f o r e s t  and open  be f o u n d  lack  10) .  and open, open f o r e s t  open and r i p a r i a n The l a c k  Those  and open, d e n s e f o r e s t and  similar  o f v a r i a b i l i t y was l i k e l y  dense f o r e s t ,  forest  (Table 23).  open and r i p a r i a n were e q u a l l y  (Table  forest  t r e a t m e n t s i t e s t h e r e was much l e s s v a r i a b i l i t y i n  c o m m u n i t i e s were t h e most s i m i l a r  This  Open  as might  habitat.  s i m i l a r i t y measures.  and  Open  and open communities were n o t a s s i m i l a r  have been e x p e c t e d f o r s i m i l a r  in  habitat.  into  t o one a n o t h e r .  due t o t h e l a c k o f  habitat  of r i p a r i a n  and r i p a r i a n ,  i n treatment  habitat  l e d to the  absence o f three s p e c i e s a s s o c i a t e d w i t h r i p a r i a n S w a i n s o n ' s t h r u s h , W i l s o n ' s w a r b l e r and W a r b l i n g This resulted  i n a much l e s s  h a b i t a t s o f treatment  sites.  sites  d i v e r s e community  habitat: vireo).  i n riparian  67  Bird/habitat relationships: Of t h e 22  s p e c i e s and  Chi-square  f o u r s p e c i e s g r o u p s examined,  s p e c i e s and  two  s p e c i e s g r o u p s were f o u n d  significant  a s s o c i a t i o n s w i t h one,  h a b i t a t types. Table  24.  First,  Those s i g n i f i c a n t  From t h i s  analysis  a n a l y s i s two  only f i v e s i g n i f i c a n t  t o have  or a combination results  of,  are presented  Chi-square  values  represented  >50%  that habitat).  Chi-square  were d e r i v e d from  remaining  low  and  significant  intermediate cover  Second, many o f t h e s i g n i f i c a n t derived  from  a low p e r c e n t a g e  in  main p o i n t s emerge.  c e l l s w i t h extreme h a b i t a t c l a s s e s ( i . e . , The  12  Chi-square  coverage  of  values  classes.  v a l u e s were  o f t h e t o t a l number o f  detections. These s i g n i f i c a n t interpret  Chi-square  because i t i s h a r d  values are d i f f i c u l t  to determine  i f a species  a s s o c i a t e d with that p a r t i c u l a r h a b i t a t type, another  h a b i t a t type,  combination  of h a b i t a t t y p e s .  interpretation significant w h i l e >50% habitat Two  or o c c u r r i n g i n a c e l l  of the t o t a l  avoiding with  a  for a particular  t o have a  extreme h a b i t a t  number o f d e t e c t i o n s o c c u r  i n other  t y p e s , o r o t h e r c l a s s e s o f t h e same h a b i t a t t y p e .  s p e c i e s , Northern  flicker  d e t e c t e d c e l l s w i t h >75% control  was  F u r t h e r problems i n  a r i s e when a s p e c i e s i s f o u n d  association  to  sites.  and  coverage  Vesper  s p a r r o w were  o f open f o r e s t h a b i t a t i n  Dusky f l y c a t c h e r s were d e t e c t e d coverage  in  sites  of dense f o r e s t .  Red-  w i t h between 51%  and  75%  naped s a p s u c k e r s  and  w o o d p e c k e r s were d e t e c t e d  in  sites  68  ui  O 55  c o  •H -p u OJ -p  co  rH  QJ  Q  CTl  ft  X  ui X)  o  o  H  H  in  ro o  VO  rH rH  in  CTl  OJ H  CTi OJ CTl OJ OJ  O OJ rH  ro  H VO o  ro  ro  rH  rH  o in H  in  in CTl  CO CTl rH O H H CO o  ro  rH rH  VO VO  rro H rro H  ro  r- vo  OJ OJ  rH ro VO > -* M VO OJ  rH  co vo r>  vo  O in ro rH VO VO  CTiOJ OJ  ro  VO  ro o  OJ iH  OJ  rH OJ rH  o  o  in  ro r-  OJ rH  rH  -3"  VO vo VO  CTl  OJ  OJ  in  ^ ^ ^ H H  H  OJ  OJ rH rH  OJ rH  rH  O  <tf CTl  ro  SH  QJ X  SH  QJ rH X!  O  QJ ft TS O  c •H ui  OJ •H u 0) a  X! 0 JH  o  TS  c u o  SH SH TJ  QJ X  rd 0  •iH SH OJ  -P  c QJ g -p rrj 0) SH EH  rd X)  1 X 0  rd  rd  ft Ul  fji C  •iH ft ft -H  rH CQ  x:  -P  -P  C  OJ g -P  id  QJ  SH EH  >H QJ QJ X! X!  3 = O  0  C  g  c  o  JH  u  c QJ g  -P  id  Q J JH EH  SH  .  O  o  -P  rd  -p rd  0  0  rH rH HH< < H >1 >1  XX Ul 3  Ul 3  a a  rH rH O 0 JH  JH  -P  •P  C O  a  )H JH QJ XQ!J QJ SH  x: 0  0  -P  -P  x: O  -P  rd rd rd  0 0 0 >i >i >i  rH rH rH < + H4H 4H  >!>.>.  XX  Ul Ul Ul 3 3 3  Q  Q  Q  -P  -P  -P  C  C  C  QJ QJ QJ g g g -P  -P  -P  rd rd id  C O  Q J QJ QJ JH SH JH  U  EH  EH  EH  id JH  JH  QJ QJ XX O  0  -<H- H rH rH ( W <H  > TS  Q J C 0 )H  C  J-i  JH  QJ QJ X! X: - P -p rH u o o 55 55  0  1 Q J tr> C  rd JH O  -P  -P  -P  QJ OJ QJ rH r-\ r-\ tn C  C  t7> C  ***  •<H T(  -H  TS  -H  TS  QJ QJ QJ  C C C S J S O O 0 SH  JH  0  0  I  JH  U 1 1  >. >i >i XI XI XI 3 3 3 OH OS  OS  > id O r-l  H  rH  rH  rH OJ  ro  H rH  rH rH o o SH  rH 0  rH rH rH O O 0 JH JH -P -P  -P  SH -P  JH -P  JH -P  C O  C O  C O  C 0  C O  C O  U  U  U  U  U  U  -p id  -P  •H  Ul  X  •P  Ul  SH  Q J SH  Ul  O  O  O  <4H  <4H  <+H  QJ  X!  rd  -P  o  •  o  •  Q J SH  a  •  c rd •H SH  -P  C  id •H SH  rd rd  ft ft •H -rH OS OS  -p  Ul Ul JH  O  O  II Q  OJ  11 -P  o\° Ul  in  QJ OIJ SH O H  II  rH  4H  Q  rH  o  O <H OJ  Ul  O Q J SH II  O  O <4H  c  QJ ft  •• Ul O Ul  II  id  -P  SH  O  rH Ul U Q J SH -P  C  QJ QJ- id H  SH  J VO Q OJ U) C QJ  Q J Ul  u u •p  O 4H  I 1  -P  ui  QJ U)  in  o\°  id >H  O  -p Ul Q J SH  SH  MHVM rd ft o • r H Q O OH  -P  Ul  Ul  SH  SH  O  O  QJ QJ  <4H  •  <JH  Q o  «  c rd  C  C  C  id id id  SH  -rSH H JH -rHSH-rH  ft -OH rH  ft ft ft -H -H -rH OH OH OH  -H  rd  rd rd id  QJ <4H  >  o • u o  69  03  H  o  C  2;  •rH O  H  H  VO VO VO  r> o  vo  O VO VO VO rH  r> rH  vo vo CM CM  CM CM CN CM  CM CO VO  CN CN CN  CN  VD O  •tf  CM CN CN  -P  U CO  rH IT) IT)  -p  o< \>  CO Q  •  ro  •  03  •  •  CM  •  •  •  CO  •  •  CM ON CO  ON CM  vo in in  co n -cr  CO CO •tf CTi  •tf  VD  CO  CO  in  o -tf  H  CO CN  CTi  00  H  o in o  CN rH rH  rH in CM rH  rH  O in  co r~ r-  \T  H  n  CO H O H rH rH  in in  t> (Xl CN CO CO CN rH rH  vo  rH CN  r-rH  CO  in  CM  ON  CM  SH 0)  rl  rl  rl  CO CO CO  XXX 0 u 3 03  3 0)  o 3 0)  ft ft ft  (0 rO rO 03  •H  U CO  ft  CO  c  •H •H  TS 0)  ft ft ft  (0 (0 rO C C C  1 1 1 TJ TS Ti 0)  QJ  P4  &  •P  -P  0)  in  CTi CO  ON ON rH  c  X3  X!  rH  (0  SH  03 03  SH SH  SH SH  o  rH SH  rO (0 (0  -Cft ft ft TS QJ  03  SH  03  SH  03  SH  X! >H  (0 03  —  C 0  CO <D QJ C ft ft ft 5 03 03 03  rH  O  -H  03  QJ >  QJ >  QJ >  03  &  TS 0)  ft fi 3 SH  1  > O  rH rH rH rH O  0 o  rH rH VH  -P  -P  -P  £  QJ  ft ft fi fi 3 SH  3 SH  o  O  11  rH  -P  SH -P  QJ QJ 4H  1 QJ Cn (0  •rH rH  o u  o u  O U  C  -p  QJ SH  0) SH  SH  EH  EH  EH  rO CO  O  in • i  (0 QJ  03  n  H  H  CM rH  CO  rH  rH  O  rH  rH  C  03  01  01  SH 0)  SH 0)  SH QJ  XXX O  QJ  O QJ  O QJ  ft ft ft  TS TI T> O O  O O  O O  (0  -p -H  Xi CO  X  CN  II  in  -P 03 QJ  CM SH  1O  rH  II rH  •P  C  -P  -P  -P  -P  (0 ro ro  0) 0) 0) SH SH SH EH EH EH  H  ro  rH  O  II O  •• 0) 01 01  ro  0  Q U3  CO  C (0  SH  -rH  O  SH  <w C (0 CO ft •  ft-H  O O PS  -P -P 03 03  CO SH O  QJ SH O  -P 03 QJ SH O  tH VM <U  C rO  •rH  C QJ  •  •  •  ft  O Q O O  VH  rO  ft  •H  -P 03 QJ SH O  vw •  Q  -P -P 03 03  -P 03  QJ  QJ  -P 01 QJ  SH SH O O <+H <*H  SH O <*H  SH O <4H  QJ  •  .  Q Q  •  Q  C cO  •H  C  CO  ft  o  II -p 01  0) SH SH O QJ 4_|  > O  •  u O  ft  II II  ft-H  o O  QJ SH O <4H  SH  C ro QJ  •  Q -P 03  -P  c c 0) 0) QJ fi fi fi  rH  -P  -p  QJ  QJ  rH  u  VIH  II Q  01  CO 03 > fO  03 QJ SH O  VD CM 03  TS  c QJ fi -P  -P  -  •H  -P  c QJ C O fi fi  CO - p  Cn C  -P  -P  CN CN rH rH  ft 03  0 Cn  C  vo  QJ  0 ) QJ >H >H  -P  c  QJ  •tf in  o  0) >H  o  U  TS TS  rH rH rH  03 QJ -H  H rH rH rH  O  U  U  rH VO  rH  rH rH  o  c c c c  0 o o u u u  rH  X!  c0 c0  QJ  o o o  SH QJ  rH rH  -P  SH SH SH EH EH EH  o u rO  u  SH 0)  rH  rH  EH  rO rfl rO CO QJ QJ  TJ  u  T3 0)  e e g -P -P -P  -p  O  0)  (0 •P •H  03  C C C CD Q) QJ  o  rH  TS  03  QJ SH  rH  in  O  CO  O •  •  •  CM H  H H  X!  03  •  rH  H H  CN CO CM  ft x w  m n o\  M i l CO *tf rH H  H  CN  •  P4  70  w i t h between 51%  and  75%  c o v e r a g e o f open h a b i t a t .  c a s e , however, t h e s e s p e c i e s detected  and  species  i n each of t h e s e h a b i t a t  g r o u p s were  time.  In a d d i t i o n , e x c e p t f o r Orange-crowned  and  Ruby-crowned k i n g l e t i n c o n t r o l s i t e s ,  not  that  e x c e e d 50%  led to s i g n i f i c a n t of the  Those s p e c i e s values  and  had  >50%  detections  of did  Chi-square  i n that  closely associated  In c o n t r o l s i t e s ,  number  detections.  both s i g n i f i c a n t  of t h e i r  of  warbler,  Chi-square values  number o f  t h a t had  c l a s s a p p e a r e d t o be habitat.  total  the  each  only  t y p e s l e s s t h a n 25%  the  detections  In  habitat  with r i p a r i a n  Ruby-crowned k i n g l e t s  avoided  s i t e s w i t h o u t r i p a r i a n h a b i t a t more t h a n e x p e c t e d , and detected  i n c e l l s with both  low  r i p a r i a n more t h a n e x p e c t e d .  Also  crowned w a r b l e r s were d e t e c t e d r i p a r i a n h a b i t a t more t h a n  and  bird  species  forest,  open, and  with  Orange-  low  amounts  Discriminant analysis  four h a b i t a t  types  riparian habitat)  function  conducted with  22  (open f o r e s t , dense  as  predictors  of  p r e s e n c e o r absence o f b i r d  detections  i n 50x50-m  cells.  These analyses  significant  functions.  yielded five  A l l significant  discriminant  Box's M t e s t f o r h o m o g e n e i t y o f matrices,  suggesting  However, d i s c r i m i n a n t  of  expected.  f u n c t i o n a n a l y s i s was  using  amounts o f  in controls,  in sites  Bird/habitat relationships: Discriminant  high  were  caution  the  discriminant functions  variance/covariance  i n i n t e r p r e t i n g the  function  failed  analysis  i s robust  functions. to  71  violations  of the homogeneity of the v a r i a n c e / c o v a r i a n c e  matrix with equally sized observations was  met  square  or l a r g e samples  i n each g r o u p ; F r e e s e  by a l l 5 s i g n i f i c a n t values, p-values,  % correct  classification  T a b l e 25.  The  coefficients  (greater than  1964).  This  Comrey  of each f u n c t i o n are p r e s e n t e d  function are presented  and  each  i n T a b l e 26.  loadings of l e s s than  0.45  Following  were n o t  Group means f o r e a c h d i s c r i m i n a n t f u n c t i o n  are presented  i n Table  27.  In c o n t r o l s i t e s ,  three variables  d i s c r i m i n a t i o n o f c e l l s w i t h and The  loading matrix  predictor variables coverage  ( T a b l e 26)  o f dense f o r e s t was  d i s t i n g u i s h i n g occupied  and  sparrows occurred i n c e l l s forest  (*=presence  = 3 9  s p a r r o w s were a b s e n t  without  sparrow  of c o r r e l a t i o n s  between  lower those  8  (*absence  Vesper  cells  where  Also  detected  ••  percent cover in  of  cells  cover  were b i r d s were  Finally,  Vesper  contributing  s p a r r o w s were found  in cells  2 6  Vesper  amounts of d e n s e  g r o u p s was  ( p r e s e n c e 5 . 1 % ) than (^absence"  variable  cells.  51.7%).  =  w i t h a h i g h e r amount o f open f o r e s t = 3  t h a t percent  unoccupied with  the  Vesper  suggest  • % ) than  open f o r e s t h a b i t a t .  contributed to  the primary  t o t h e s e p a r a t i o n o f t h e s e two  x  in  l o a d i n g matrices of the pooled w i t h i n  interpreted.  detections.  and  standardized canonical discriminant function  and  (1973),  Chi-  i n each g r o u p ,  g r o u p c o r r e l a t i o n s between p r e d i c t o r v a r i a b l e s discriminant  criterion  discriminant functions.  number o f c a s e s  30  Vesper  not  s p a r r o w s were  X  o  PQ  TS  o  o  o  o  o  o V  o V  o V  o V  o V  00 • VO  •  •  VO  VO  CM  in CM in r-~  ^  OJ >i  • H rH M-l - P -H O  |oV> Ul J-I rd M rH  CO  cn •  •  co co in  O  o o  -p c 0)  Ul  r>  CT\  CM  e Z I - P  01  Ul XI fd  ft  CM  in o in vo  CO VO OA  CM  H  o o  o o  H o  H o  H o  o V  o V  o V  o V  o V  CM • CO CM CM  O  in •  in co m CM CM  CTl CO • • • CO CM rH rH rH  u rH  X! JH  JH QJ  x;  fd S  U TS -P Q) rd ft  0 e  >i H JH <«H I  >i O * rH Ul rH  3 0)  Q  >H  Z 0 JH JH  JH QJ  XI  0 o  rd - P ft rd ui  JH JH  rd ft  O  >i  ui  tn rH  ft ft*  -H  JH  QJ ft Ul •H Ul QJ x: d > >1  u Q  73 T a b l e 26. D i s c r i m i n a n t f u n c t i o n c o e f f i c i e n t s and p o o l e d w i t h i n group c o r r e l a t i o n s f o r d i r e c t d i s c r i m i n a n t function a n a l y s i s f o r t r e a t m e n t s and c o n t r o l s . Standardized Canonical Discriminant Function Coefficients  Pooled Within Group Correlations  Treatments Dusky flycatcher  Open f o r e s t Dense f o r e s t Open  4 . 52 2 . 97 2 .20  Open f o r e s t Dense f o r e s t Riparian Open  0. -0. -0. -0.  90 83 42 27  Yellow-rumped warbler  Open f o r e s t Dense f o r e s t Open  1. 09 1. 68 - 0 . 06  Dense f o r e s t Open Open f o r e s t Riparian  0. -0. -0. -0.  76 61 29 24  Chipping sparrow  Open f o r e s t Dense f o r e s t Open  4 . 35 4 . 01 2 .71  Riparian Open f o r e s t Open Dense f o r e s t  -0. 0. -0. -0.  87 50 34 06  Dusky flycatcher  Open f o r e s t Dense f o r e s t Open  4 . 75 4 . 52 3 .48  Riparian Open f o r e s t Dense f o r e s t Open  -0. o. -b. 0.  97 28 14 08  Vesper sparrow  Open f o r e s t Dense f o r e s t Open  2.42 1.54 1. 88  Dense f o r e s t Open f o r e s t Riparian Open  - 0 . 85 0 . 63 -0 . 45 0. 41  Controls  co CO 5-1  c (0  o  PH  •H  •  CM co VO rH  (N  co o  •  <H O  •  H  •  o  •  •  CO o  •  SH  fO  a  - H  CO XI  04  • o  CO CO  •  ft o QJ  rH  i n CO CO  u cm  C CO  in  CO VO • • • • H CO o rH  CM • • o O  •  •  •  •  o o in CM CM CM CO  VO rH i n  cn CM i n VO  •  X!  VO O  •  «  i n i n rH  •  •  •  •  •  •  CM CO i n  cn H o rH CM CM CM  CO CM  cn  -P tO -P  •H  XI  cn  (0  •  QJ  -p [TJ  CO  u Pu  5-1 O  c  in  CO CO ft 5-1 O O 4H  •  cn co 5-1  •  PH  cn  cn  ft  0)  QJ 2  to  54  S  •  •  CO  i n CO CO H  •  •  •  •  •  •  •  •  in  •  in H  •  •  15  5H QJ  •  VO rH CO VO CM CM CM CM  X!  U  cn  •  --rPH O  XI CO C fO Xi 3 EH  •  rH  CO C X! - H O -P fO u O  qo  H  •  CM o cn rH i n i n •* i n  CO  • o cn c CM Q) o QJ  CO VO •>*  5H  t> ro CO  •  rH  ft  X!  •  in  cn cn CO 0 0 CO CO  C 0 QJ  i n CO  in 00 CM CM CM CO  •  •H  rO  •  cn H ON i n CO  H CO CO 00  0) to 0 Ss QJ  •  •  •  cn o  X!  u  •  co CO CO  X!  -p  •  CM o CO rH CM rH  4H  -p  cn CO  - H  0  CO  ft CO  c  CO g -p  CO QJ SH  c  U (0  >  rH  o  SH  1  i u > CO W  - H  *  g  rH0 rH  3 CO  <a  C U X! S 5H O O X! rO - P 5H O ft ( 0 SH SH CO o tO  T3  CO ft g >i 3  fO m  O  >  - H  cn  rH 0  rH  -p  c o u  C (0  > i ft t n < H oi  C MH  O - H ft  - H  ft*ft SH  >I  0)  SH QJ - H  cn CO g X! 3 CO  < U Q  >  75  found  in cells  ( presence x  One  °.1%)  =  detections  w i t h lower amounts o f r i p a r i a n h a b i t a t than i n c e l l s  (* bsence  without Vesper  1.5%).  =  a  significant discriminant  f u n c t i o n was  Yellow-rumped warblers i n t r e a t m e n t matrix  i n T a b l e 26 i n d i c a t e s t h a t  forest  and  x  open h a b i t a t , d i s t i n g u i s h c e l l s  w a r b l e r s were not d e t e c t e d  w i t h and  (*absence  =  5.3%).  amount o f open h a b i t a t  without  Yellow-rumped w a r b l e r d e t e c t i o n s  One  (*absence  (Xpresence  In a d d i t i o n ,  responsible  habitat  in cells  w i t h low p e r c e n t c o v e r a g e = 0  -  8 %  .  (Table 26).  v s  -  x  The  these groups  riparian  (*presence  = 1  •'  c  t o have e  l l  generated f o r  primary was  variable  t h e amount o f  C h i p p i n g sparrows  were  absence= .4%).  In a d d i t i o n ,  3  the  also contributed  separation  detections.  sparrows  w i t h and  without  were d e t e c t e d i n c e l l s  o f open f o r e s t h a b i t a t  where t h e y were not d e t e c t e d  28.8%) t h a n  (*absence  =  21.7%).  to the  Chipping  with a higher percent  (^presence"  found  of r i p a r i a n h a b i t a t  p e r c e n t c o v e r o f open f o r e s t h a b i t a t of c e l l s  s  a h i g h e r amount  f u n c t i o n was  in control sites.  for separating  had  tended  5.3%).  =  significant discriminant  C h i p p i n g sparrows  without  where Yellow-rumped  w i t h Yellow-rumped w a r b l e r s d e t e c t i o n s  open h a b i t a t  loading  of dense f o r e s t  a lower  of  The  T h e s e b i r d s were f o u n d i n  13.5%) compared w i t h c e l l s  =  generated f o r  v a r i a b l e s , dense  with a higher percent coverage  ( presence  cells  sites.  two  Yellow-rumped warbler d e t e c t i o n s . cells  sparrow  cover  in cells  76  Dusky f l y c a t c h e r s were t h e o n l y s p e c i e s t o have significant controls.  discriminant  functions  i n b o t h t r e a t m e n t s and  In c o n t r o l s the l o a d i n g m a t r i x  ( T a b l e 26)  s u g g e s t s t h a t r i p a r i a n h a b i t a t was t h e o n l y v a r i a b l e contributed  t o the s e p a r a t i o n of c e l l s  Dusky f l y c a t c h e r d e t e c t i o n s . cells  ( presence x  ( absence x  role  =  3  •than  • %)•  = 0  6  I  i n predicting  in cells  lower x  Dusky f l y c a t c h e r  = 8 9  %)•  T  n  e  presence  (Xp  habitat  r e s e n c e  this  h a b i t a t than  s p e c i e s was  in cells  ( presence •°%/ = 1  x  r e s e n c e  found  = 0  than  in cells  = 2 0 . 2 % , ^absence in cells  where t h e y were  absence  =73%,  Dusky f l y c a t c h e r s were f o u n d i n  (Xp  where t h e y were a b s e n t  with  was t h e n e x t  c e l l s w i t h h i g h e r amounts o f d e n s e h a b i t a t  x  (Table 26).  responsible f o r the  amount o f d e n s e f o r e s t  most i n f l u e n t i a l v a r i a b l e .  Finally,  absent  Dusky f l y c a t c h e r s were d e t e c t e d i n c e l l s  amounts o f open f o r e s t  absence  habitat  three variables played a  Open f o r e s t was t h e v a r i a b l e p r i m a r i l y separation.  found i n  where t h e y were  treatments,  n  w i t h and w i t h o u t  T h i s s p e c i e s was  w i t h a h i g h e r amount o f r i p a r i a n  that  = 7  w i t h more  - ^)• 3  riparian  absent  • %)• 2  Discussion Community  structure  This thinning  project  the s t r u c t u r e of the f o r e s t diversity  appeared bird  curves of communities  s i t e s were v e r y s i m i l a r  (Figure  t o have l i t t l e  community.  affect  Dominance  i n t h i n n e d and u n t h i n n e d 3).  Each  community  on  77  c o n s i s t e d o f one 8 relatively species.  very  abundant s p e c i e s , and  I n b o t h t r e a t m e n t and  s p e c i e s were t h e  same a l t h o u g h  somewhat d i f f e r e n t and  common s p e c i e s  (Table  t h i n n i n g had  their  11).  The  further i l l u s t r a t e on o v e r a l l  junco,  (either  i n terms o f the  and  vegetation relative  but  logging  al.  1990,  and  shrub cover.  fire  D e G r a f f e t a l . 1991)  1  In my  increase  study  ground-feeding species of t h i s  (Hagar 1960,  the  to  n e g l i g i b l e response of the  only h a b i t a t v a r i a b l e that  for  Lynch  the  Stribling  increases  community vegetation  increased  1976,  1970, et  i n ground  l a c k of response of  bird  in  understory  Flack  (Hagar 1990,  due  species  feeding/nesting  (Bock and  thinning  of  Townsend s  trend  shrub  the  Dark-eyed  There i s a general  Omhart 1978), and  that  on  of the  in  d i d not.  R a p h a e l e t a l . 1987)  the  number  s u c h as  sparrow,  abundance o f g r o u n d and  and  Jaccard's  0.96,  anticipated increase  species to increase a f t e r Franzreb  of  impact  A m e r i c a n r o b i n were e x p e c t e d t o  abundance because o f an  were  affect  o f any  shrub foragers  V e s p e r sparrow, C h i p p i n g  solitare,  little  o r number o f d e t e c t i o n s ) Ground and  uncommon  community s t r u c t u r e .  a l s o seems t o have had  examined.  and  the minimal  Thinning  territories  values  (0.92  7-  9 most a b u n d a n t  rank orders high  abundance  abundance  sparrows),  relatively  c o n t r o l s the  Relative  relative  The  14-15  M o r i s i t a ' s i n d i c e s of s i m i l a r i t y  respectively)  with  (Chipping  the  is consistent community.  significantly  due  78  t o t h i n n i n g was significant and  shrubs  down and  increase (Table  i n the  9).  piles  logging  (Franzreb  afforded junco,  foraging  posts.  Slash  s l a s h was  arranged  These s l a s h particularly  opportunities, in this  i n other  (at l e a s t  i n the  species.  and  The  availability  (Sturnam 1968,  amount o f of  food  species  S z a r o and  Balda  1979).  foliage-feeding species.  Balda  such  as  with the  foliage-feeding species  fire  1986).  foliage-feeding  (Flack  1976,  Conversely, species  in  close  the in  after  R a p h a e l e t a l . 1987, i n the  canopy  abundance  have been n o t e d  decreases  per  species  relative  Increases  a  in  been r e l a t e d  In a d d i t i o n ,  where f o r e s t c a n o p i e s d e v e l o p and or  on  Red-breasted  for foliage-feeding  of  abundance o f  plane)  affect  c a n o p y volume has  a l s o been a s s o c i a t e d  and  and  where i t  i n c a n o p y volume on  volume has  harvesting  Dark-eyed  Western t a n a g e r s were e x p e c t e d t o d e c r e a s e  basis.  instances  piles  left  h a v e any  Foliage-feeding  abundance b e c a u s e o f a r e d u c t i o n  several  the  in discrete  horizontal  Yellow-rumped w a r b l e r , M o u n t a i n c h i c k a d e e ,  t o the  in  studies.  foliage-feeding  hectare  of  escape cover  s t u d y was  t h i n n i n g treatment d i d not  nuthatch,  grass  increase  increases  no  T h i s a p p r o a c h r e s u l t e d i n a more u n i f o r m  d i s t r i b u t i o n of s l a s h  The  where t h e  Omhart 1 9 7 8 ) .  T h e r e was  cover of herbs,  a t t r i b u t e d to  slash, the  and  increased  felled.  than  was  ground-feeding s p e c i e s ,  observation was  percent  In s t u d i e s  ground-feeding species amount o f  dead woody m a t e r i a l .  Szaro  abundance  have b e e n c o r r e l a t e d w i t h  of  decreases  79  i n canopy volume a f t e r Morgan e t a l . 1989). less dbh  harvesting I n my  canopy volume p e r  however, t h e r e  was  abundance o f any estimate.  no  than  decrease  species  i t was  believed  that  i n f l u e n c e o f one abundant i n the within  this  The  plot. Dam  and  of the  hence r e d u c t i o n  treated  (spruce  bird  This  due  to  was  more  the very  I located 7  nests  s i t e s was  based on  limiting  f o r t h i s group o f  volume.  Thinning  component o f t h e  in this forest  proportion  First,  the not  Second, an  Douglas-fir  l o s s of a v a i l a b l e f o r a g i n g decline  community may  factors.  limiting.  budworm and  predicted  significant  15).  greatest.  canopy volume, may  s e v e r e enough t o make f o o d  The  detected  plot.  l a c k of response of the  negated the  relative  Mountain  largely  Lake s t u d y s i t e .  following  food  7);  c h i c k a d e e s w o u l d be were  cm  density  M o u n t a i n c h i c k a d e e s were  a t t r i b u t a b l e t o the  of  (Table  (Tables  number o f  c h i c k a d e e s i n t r e a t m e n t s i t e s was  1978,  t r e e s <10  i n the  f o r each  opportunities  d i f f e r e n c e i n the  Omhart  significantly  M o u n t a i n c h i c k a d e e s were  a b u n d a n t where f o r a g i n g However, t h e  was  in control sites  s i g n i f i c a n t l y more i n t r e a t m e n t s i t e s u n e x p e c t e d as  and  of D o u g l a s - f i r  significant  individual  In f a c t ,  study there  hectare  i n treatment s i t e s  (Franzreb  be treatment  have been abundance  t u s s o c k moth) habitat.  in foliage-feeding species the  assumption t h a t  species  of t h i s  was  because of reduced  p r o j e c t was  stratum  food  in  d i r e c t e d at a  ( t r e e s <15 s t r a t u m was  cm  dbh).  dead o r  canopy  specific A dying,  in  80  b o t h c o n t r o l and t r e a t m e n t s i t e s and  a t t h e time of study  21% r e s p e c t i v e l y ; T a b l e 4) t h e r e b y r e p r e s e n t i n g  t h a t was  already  Had t h i n n i n g  unavailable  occurred  to foliage-feeding  p r i o r to the extensive  e f f e c t that t h i s thinning  habitat could are  trial  have been g r e a t e r .  amount o f f o r a g i n g  form of o v e r s t o r y (Table  6) .  limiting  Douglas-fir  Therefore,  In a d d i t i o n , these  stands  erroneous.  Because t h i n n i n g this  habitat  To make f o o d  still  remained i n t h e  and P o n d e r o s a p i n e  t r e a t m e n t may  limiting,  was  forest, a  the assumption t h a t  as a r e s u l t o f t h i s  t u s s o c k moth, foraging  a t t h e younger t r e e s w i t h i n  significant  species.  had on l i m i t i n g  managed on an un-even aged b a s i s .  directed  habitat  d e f o l i a t i o n of  t h e s e s t a n d s by s p r u c e budworm and D o u g l a s - f i r the  (3 0%  food  foliage was made  have been  and t h e r e f o r e  t o see t h e  d e c r e a s e i n t h e number o f f o l i a g e - f e e d i n g s p e c i e s ,  thinning  w o u l d have had t o have been d i r e c t e d a t r e m o v i n g a  greater  number o f h e a l t h i e r t r e e s ,  and/or t r e e s  i n larger size  classes. The p r e c e d i n g  argument assumes  that  a reduction i n  canopy volume would have l e d t o a c o r r e s p o n d i n g r e d u c t i o n the  availability  Douglas-fir  of f o o d .  However,  t u s s o c k moth were  t h e s e stands that the B r i t i s h it  ( B a c i l l u s  associated  t h u r i g e n s i s ) .  Columbia F o r e s t  Insect  with f o r e s t pests,  s p r u c e budworm and  both s u f f i c i e n t l y  n e c e s s a r y t o s p r a y some o f my  in  study  abundant i n  Service  s i t e s with  outbreaks,  felt  BTK  e s p e c i a l l y those  and s p e c i f i c a l l y  spruce  budworm, have been c o r r e l a t e d w i t h n u m e r i c a l r e s p o n s e s o f  81  some s p e c i e s Morris  of birds  ( M a c A r t h u r 1958, B l a i s  e t a l . 1958, M a t t s o n e t a l . 1968, C r a w f o r d and  J e n n i n g s 1989).  Therefore,  elevated  budworm may have l e d t o i n c r e a s e d species  Whether  negative  effects of the habitat  l e v e l s o f some  the presence of t h i s  abundant f o o d  source could  reduced a v a i l a b i l i t y  from t h i n n i n g .  i n any s p e c i e s o r source l i k e l y  i n t h i s manipulation.  superabundant food  of foraging  manipulation.  t u s s o c k moth  l e d to a numerical response  confounding a f f e c t  the  of spruce  population  s p r u c e budworm and D o u g l a s - f i r  populations  this  levels  i n b o t h c o n t r o l s and t r e a t m e n t s t h e r e b y damping t h e  potential  not,  and Banks 1964,  had a  The p r e s e n c e o f  easily habitat  When budworm p o p u l a t i o n s  have made up f o r that  crash,  i n t h e abundance o f f o l i a g e - f e e d i n g s p e c i e s  may  resulted  differences become  apparent. The modest detections result  significant  f a v o r i n g t r e a t m e n t s o v e r c o n t r o l s may have been a  of bias  in detectability  have been more e a s i l y controls. detect  detected  (Table  birds visually.  Woodpeckers  may  i n treatments than i n  However, i f v i s i b i l i t y  o r groups of s p e c i e s ,  more o f t e n  i n treated  was t h e  i n t h e number o f d e t e c t i o n s ,  have been more s y s t e m a t i c .  species,  16).  The openness o f t r e a t m e n t s d i d make i t e a s i e r t o  reason f o r differences should  d i f f e r e n c e i n woodpecker  sites.  In other  should This  words,  h a v e been  the bias more  detected  was n o t t h e c a s e .  Only  M o u n t a i n c h i c k a d e e s were d e t e c t e d  more o f t e n  sites.  f o r the difference i n the  A more l i k e l y  explanation  i n treated  82  number o f d e t e c t i o n s  i s t h a t w o o d p e c k e r s were u s i n g  treatment s i t e s p r e f e r e n t i a l l y opportunities. m a t e r i a l on  The  contribute to  Hodges 1985,  a r e major p r e d a t o r s 1984) , and  like passerines,  numerical  and  abundance  (Conner and  1969,  represented  treatment supports  sites. this  1974,  probable  elevated  source  Observations  idea.  Safranyik  increased Knight  During  of  this  food  1958,  during  the t h i n n i n g  Koplin  dead m a t e r i a l  beetle  in  populations,  t h a t drew woodpeckers i n t o f o r a g i n g woodpeckers study,  a l l 5 species  woodpeckers e n c o u n t e r e d were s e e n f o r a g i n g on t h e felled  1987,  Woodpeckers  (Moeck and  a b u n d a n t down and  a food  beetle  h a v e been known t o show b o t h  Crawford  The  t r e a t m e n t s i t e s and likely  (Moser e t a l .  functional responses to  O t v o s 1970).  i n bark  C o n n e r e t a l . 1991).  of bark b e e t l e s  foraging  leaving cut  increases  populations  f r o n t a l i s )  N e b e c k e r and  increased  p r a c t i c e o f t h i n n i n g and  s i t e can  (Dendroctonus  because of  of  material  treatment.  Bird/habitat relationships Dominance-diversity illustrates  two  main p o i n t s .  appears t o support four h a b i t a t types. not  analysis for control  t h e most d i s t i n c t  riparian  sparrows.  Wilson's warblers,  more p r o m i n e n t i n t h i s  and  community  the  only  of  Ruby-crowned  Swainson's thrushes  were  (Table  open  22)  the  community  I n a d d i t i o n , many  s u c h as O r a n g e - c r o w n e d w a r b l e r s ,  kinglets,  habitat  community o f any  T h i s community was  d o m i n a t e d by C h i p p i n g  species  First,  sites  Second,  83  habitat This  i s not  areas and  supported the  s u r p r i s i n g given the  (i.e.,  as  t h e y do  s u c h , do  Similarity  not  richness  found of  the  responsible  (Table  open h a b i t a t  also  No  The  grassland  the The  o f two  four  the  whereas i n t r e a t m e n t s i t e s , abundant  (Table of  areas of  18).  species  primarily  due  t o the  association  association  sites.  open h a b i t a t  l a n d s c a p e by habitat  changing  types.  open f o r e s t was  types.  In  the  may  type,  most  evident  s i t e s as t h e  expansive that  have changed  following  in control  of t h e s e a r e a s  should disappear,  isolated  I t i s then possible i n the  u s e s open c a n o p y f o r e s t ,  s h o u l d be  the  control  Thinning e f f e c t i v e l y created  associations  relative rarity  distinct  extreme  most a b u n d a n t h a b i t a t  open canopy f o r e s t .  I f a species  dense  i n treatment  d e n s e - u n t h i n n e d f o r e s t s u r r o u n d e d by  bird/habitat  treatment  likely  forest  of the  d e n s e f o r e s t was  the  c o m m u n i t i e s a p p e a r e d t o be  r i p a r i a n , d e n s e f o r e s t and  distribution  reduced  b e t w e e n i t and  o v e r a l l community s t r u c t u r e  of  habitat  community).  support  community was  similarity  T h i n n i n g a l t e r e d the  the  a true  these  open c o m m u n i t i e s .  i n open h a b i t a t s .  communities.  scarcity  ways.  23)  r i p a r i a n and  low  islands  types.  l i m i t e d e x t e n t of  constitute  l a c k o f d i s t i n c t n e s s was  sites,  a l l habitat  support a very d i v e r s e  f o r the  from the This  of  community i s s u b s t a n t i a l l y d i f f e r e n t f r o m  community  forest  not  indices  d i s t i n c t n e s s of the riparian  fewest s p e c i e s  or  this  s i t e s because  in controls.  This  become l e s s a p p a r e n t ,  availability  of  this  of  habitat  in  84  increases. may  Any  species  t h a t uses dense-unthinned  undergo s i m i l a r c h a n g e s .  most abundant h a b i t a t t y p e  Because dense-unthinned  in control sites,  a s s o c i a t i o n w i t h t h i s h a b i t a t t y p e may (i.e.,  species  d i s t r i b u t i o n w o u l d be  However, i n t r e a t m e n t s i t e s abundant. should  As  a result,  become a p p a r e n t  C h i - s q u a r e and  not  more  any  be  discriminant  uniform).  for this habitat  with high was  less  type  a n a l y s i s were p e r f o r m e d  of a s s o c i a t i o n e x i s t e d .  f o l l o w the  The  However, and  Dusky  expected pattern.  N o r t h e r n f l i c k e r s were d e t e c t e d  Both i n areas  amounts o f open f o r e s t i n c o n t r o l s i t e s more t h a n  expected.  These a s s o c i a t i o n s were n o t  apparent  in  t r e a t m e n t s i t e s where open f o r e s t was  very  24).  hand, were d e t e c t e d  Dusky f l y c a t c h e r s , on  areas with high (Table  to  bird/habitat  Vesper sparrows, N o r t h e r n f l i c k e r s  V e s p e r s p a r r o w s and  strong  apparent  r e l a t i o n s h i p s y i e l d e d m o s t l y ambiguous r e s u l t s .  f l y c a t c h e r s appeared t o  the  there.  u n i - v a r i a t e Chi-square a n a l y s i s of  species,  is  d e n s e - u n t h i n n e d h a b i t a t was  preference  d e t e r m i n e whether t h i s k i n d  three  habitat  24).  the  other  abundant  amounts o f d e n s e f o r e s t i n t r e a t m e n t  This  a s s o c i a t i o n was  not  present  (Table in  sites  in control  s i t e s where dense f o r e s t i s f a r more a b u n d a n t . Results  from d i s c r i m i n a n t  a n a l y s i s suggest t h a t  changes i n t h e  d i s t r i b u t i o n o f h a b i t a t s may  changes i n t h e  spatial  control sites,  Chipping  d i s t r i b u t i o n of three s p a r r o w s and  a p p e a r e d t o be most s t r o n g l y  Vesper  associated  the  have l e d t o species.  the In  sparrows  with areas that  had  85  more open f o r e s t c h a r a c t e r i s t i c s dense f o r e s t , 26).  and more abundant open  In treatment s i t e s ,  noted.  (i.e.,  however,  areas with  forest  this  habitat;  o f dense f o r e s t h a b i t a t not noted  Table  a s s o c i a t i o n was  C o n v e r s e l y , Y e l l o w - r u m p e d w a r b l e r s and  f l y c a t c h e r s were found most o f t e n  little  not  Dusky  i n a r e a s w i t h h i g h amounts  i n treatments.  This association  i n c o n t r o l s i t e s where d e n s e f o r e s t h a b i t a t  was  was  more a b u n d a n t . Results the Chi-square a n a l y s i s  further  suggest  r i p a r i a n h a b i t a t s u p p o r t e d a somewhat d i s t i n c t p l a y e d a major r o l e Five  i n the d i s t r i b u t i o n  flycatchers) detected  sapsuckers  and one s p e c i e s g r o u p  o f some s p e c i e s .  and  Orange-crowned  Ruby-crowned  k i n g l e t s and t o a l e s s e r  t o be a s s o c i a t e d w i t h r i p a r i a n  extent  Dusky  s p e c i e s t h a t a r e known  habitat  (Brown  s a p s u c k e r s and t h e woodpecker  d e t e c t e d more o f t e n t h a n e x p e c t e d  1985). g r o u p were  in riparian  s a p s u c k e r s were o f t e n  water b i r c h t r e e s t h a t a r e found  habitat  observed  H a i r y woodpeckers, and P i l e a t e d aspen t r e e s  woodpeckers  ( B l a c k - b a c k e d and  w o o d p e c k e r s were e x c e p t i o n s i n t h i s  also  (Table  f e e d i n g on  only i n wetter areas.  naped s a p s u c k e r s , and t h e g r o u p o f w o o d p e c k e r s ,  Trembling  by c h a n c e  w a r b l e r s , W i l s o n ' s w a r b l e r s , and  f l y c a t c h e r s are long d i s t a n c e m i g r a n t  Red-naped  kinglet,  Dusky  i n r i p a r i a n h a b i t a t more t h a n e x p e c t e d  Red-naped  and  (woodpeckers) were  (Table 24).  24).  community  s p e c i e s (Orange-crowned w a r b l e r , Ruby-crowned  W i l s o n ' s w a r b l e r , Red-naped  that  Red-  particularly  often nested i n  Three-toed  group) r e s u l t i n g i n  86  numerous d e t e c t i o n s  i n c e l l s with r i p a r i a n habitat.  r i p a r i a n habitat being controls,  extremely rare  i t appears t h a t  Despite  i n b o t h t r e a t m e n t s and  even a s m a l l  amount o f r i p a r i a n  h a b i t a t c a n have a p o s i t i v e a f f e c t on s p e c i e s d i v e r s i t y . T h i s phenomenon has a l s o been n o t e d by M o r r i s o n and Meslow (1983), and Hagar It  i s unclear  were d e t e c t e d  (1990). why Red-naped s a p s u c k e r s and woodpeckers  i n o p e n i n g s more t h a n was e x p e c t e d by c h a n c e .  O p e n i n g s u s u a l l y do n o t c o n t a i n material  much down and dead  t h a t may have drawn some o f t h e woodpecker  i n t o open a r e a s . were d e t e c t e d nest  very  What i s m o r e ' l i k e l y  i s t h a t these  i n o p e n i n g s when en r o u t e  t o o r from  associations analysis.  involved using  their  with the bird/habitat  50x50-m c e l l s  The use o f 50x50-m c e l l s  contributed  as the u n i t o f  as t h e u n i t of a n a l y s i s  t o b o t h t h e low number o f c e l l s  p e r c e n t a g e c o v e r a g e o f any h a b i t a t  type,  with a  Chi-square values i n  intermediate  The r e a s o n t h a t u s i n g  cover c l a s s e s .  as t h e u n i t o f a n a l y s i s was p r o b l e m a t i c  Figure  entire  6.  In t h i s  50x50-m c e l l ,  between f o u r c e l l s h a b i t a t A.  instance  habitat  high  and t h e  preponderance of s i g n i f i c a n t  in  species  sites. One o f t h e p r o b l e m s a s s o c i a t e d  cell  species  a 50x50-m  is illustrated  B would c o v e r an  but because o f i t ' s p o s i t i o n , i s s p l i t  e a c h c o n s i s t i n g o f 25% h a b i t a t  Any d e t e c t i o n s  t h a t m i g h t be  B and 75%  restricted  Figure a  6.  The p o t e n t i a l p r o b l e m w i t h  50x50 m c e l l  bird/habitat  as t h e u n i t o f a n a l y s i s i n  relations.  Habitat  an e n t i r e 50x50 m c e l l .  B would  cover  However, b e c a u s e o f  the placement of the g r i d contains  using  cell,  each  cell  75% o f H a b i t a t A a n d 25% o f h a b i t a t  B. 50 m  Habitat  A  Habitat  B  50  88  t o h a b i t a t B would t h e n up  o f low  be  grouped with  amounts o f h a b i t a t B and  cells  t h a t a r e made  h i g h amounts o f h a b i t a t  A.  Conclusions Many a u t h o r s  b e l i e v e t h a t t h i n n i n g c o u l d p r o v e t o be  u s e f u l t o o l to manipulate Zeveloff belief  1980,  DeGraff  i s based mainly  microhabitat diversity diversity.  bird  e t a l . 1992, on and  T h i n n i n g has  developing  the a b i l i t y  the  forest  the stands  aged b a s i s and, understory. b e n e f i t those  x h l and  as a r e s u l t ,  Increased  forests.  xh2  resulted  a r e managed on  At an  recruited  this  present uneven-  tolerant  sites,  likely  resource. u n l i k e some  e t a l . 1992)  o f new  and  already  production w i l l  study  (e.g. DeGraff  i n the  have a l s o been  e t a l . 1992).  shrub  i n the r e c r u i t m e n t  s p e c i e s t h a t c o u l d be  mid-story.  a l r e a d y h a v e a shade  i n my  Long-billed curlews  understory  o r more s p e c i e s t h a t  s p e c i e s t h a t p r e s e n t l y use  other t h i n n i n g t r i a l s , result  enhancing  to increase w i l d l i f e  o f one  h e r b and  T h i n n i n g as c o n d u c t e d  likely  by  c a n o p y has  i n f r e q u e n t l y (DeGraff  i n t h e IDF  structural  in diversity  a s s o c i a t e d w i t h the o c c u r r e n c e are detected  This  community t h a t d i d n o t  Increases  and  increase  a shade t o l e r a n t ,  d e v e l o p m e n t o f an u n d e r s t o r y (Hagar 1990).  to  within-stand  been an e f f e c t i v e t o o l  d i v e r s i t y when o p e n i n g  (Dien  H u n t e r 1990).  Those g o a l s a r e a c h i e v e d  v e g e t a t i o n and  exist  species diversity  a  will  species into  not  these  Mountain b l u e b i r d s are  into  these  stands.  Both  two  89  were s e e n on the  study.  of very  occasion  However, b o t h s p e c i e s  l a r g e openings.  openings of the inappropriate perspective species  i n these stands over the  From a s i l v i c u l t u r a l  f o r e s t type.  be  t o a l t e r the  that  affecting bird will  controlled present it  likely  species  grazing  a c c o m p a n i e d by  diversity.  p r a c t i c e d should  Douglas-fir  (those  woodpeckers f o r n e s t i n g  and  t r e a t m e n t s and  (Table  controls  management i s i m p e r a t i v e  and  Chipping  benefit  from t h i s  from the  two  some f o r m  of  s n a g management but  as  should  program r e g a r d i n g firewood  be the  cutting  snags i n used  were r a r e  by  i n both  E f f e c t i v e snag  species  species  form o f t h i n n i n g  happen,  or a l t e r a t i o n s of  going to  i n the  study suggest t h a t  sparrows a r e  rather  t h i s to  i f these stands are  to support c a v i t y - n e s t i n g Results  For  sizes generally  foraging)  stands  species  Ponderosa pine  3).  these  best  i n these  enforcement of  and  are  continued,  an e f f e c t i v e e x t e n s i o n  l a r g e r diameter c l a s s e s  able  be  be  b e n e f i t of  a c c o m p a n i e d by  In a d d i t i o n ,  o f s n a g s i n f o r e s t s and  restrictions.  play  some r e s t r i c t i o n s  patterns.  would  wildlife  abundance o f  have t o be  b u r n i n g and  i s presently  role  relative  creation  elevations.  main r o l e t h a t t h i n n i n g w i l l  will  thinning  lower  of  perspective  as t h e s e s p e c i e s  managed i n more open s t a n d s a t The  From a  creating l a r g e openings f o r the  i s also inappropriate  years  would r e q u i r e the  s i z e needed f o r t h e s e s p e c i e s  in this  two  be  future.  Vesper  sparrows  that will potentially  applied.  Both  species  90  a p p e a r t o be  associated  made more abundant as  w i t h open c a n o p y f o r e s t s t h a t  a r e s u l t of t h i s  P a t c h e s of u n t h i n n e d in  thinned  likely  treatment.  f o r e s t should  c o n t i n u e t o be  s i t e s even i f snag management does  necessitate  it.  i n t r i c a t e part  of  this  would not  of D o u g l a s - f i r .  completely  eradicate  open  In a d d i t i o n , r e s u l t s from t h i s  associated  they provide  valuable  an  s e c u r i t y cover.  was  left  around a  Slash felled. nesting  could  Piling and  Results  seeking  study  Douglas-fir.  obvious s t r u c t u r a l d i v e r s i t y , On  w i t n e s s e d a P i l e a t e d woodpecker e s c a p i n g a C o o p e r ' s hawk by  parkland  Yellow-rumped  w i t h dense p a t c h e s of  In a d d i t i o n to p r o v i d i n g  were  a l l dense p a t c h e s  s u g g e s t t h a t a t l e a s t Dusky f l y c a t c h e r s and w a r b l e r s are  not  f o r e s t system.  F r e q u e n t w i l d f i r e s would have m a i n t a i n e d an but  left  Patches of dense-unthinned D o u g l a s - f i r  always an  forest,  are  cover  i n an  one  occasion,  I  from the p u r s u i t  unthinned patch  that  snag. be  piled,  rather  of s l a s h c o u l d  feeding  species,  from t h i s  have a s i g n i f i c a n t  than  left  b e n e f i t some o f t h e  particularly  study suggest that  a f f e c t on  where i t  the  Dark-eyed  was  ground juncos.  thinning did  forest bird  not  community.  Poor u n d e r s t o r y response t o t h i s  thinning  trial,  because of the  grazing,  leaving slash  site,  a f f e c t s of  cattle  ineffective thinning,  insufficient  period  of time,  or the or  passing  a l l four,  of may  In a d d i t i o n , e l e v a t e d  l e v e l s of  either on  an explain  n e g l i g i b l e response of ground-plus shrub-feeding species.  of  the  bird  s p r u c e budworm  and  91  Douglas-fir  t u s s o c k moth may  h a v e masked any  differences  i n t h e abundance  of foliage-feeding  These r e s u l t s ,  however,  d a t a and i l l u s t r a t e both vegetation  should  be c o n s i d e r e d  significant  as  species. preliminary  t h e need t h e f o r l o n g - t e r m m o n i t o r i n g o f  and b i r d  necessary p a r t i c u l a r l y  species.  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Manage. 52:253-260.  100  Appendix I . L i s t used i n t e x t .  o f common and s c i e n t i f i c  names o f p l a n t s  Trees Douglas-fir Ponderosa pine T r e m b l i n g aspen Red a l d e r Paper b i r c h  Pseudotsuga  m e n z i e s i i  Pinus  ponderosa  Populus  t r e m u l o i d e s  Alnus  r u b r a  B e t u l a  p a p y r i f e r a  Cornus  s t o l o n i f e r a  Shrubs r e d - o i s e r dogwood common j u n i p e r r o c k y mountain j u n i p e r rose kinnikinik Saskatoon berry sheperdia snowberry D o u g l a s maple spirea willow  J u n i p e r u s  communis  J u n i p e r u s Rosa  scopulorum  s p p .  A r c t o s t a p h y l o s A m e l a n c h i e r  u v a - u r s i a l n i f o l i a  Sheperdia  canadensis  Symphoricarpos  a l b u s  Acer  glabrum  S p i r a e a S a l i x  b e t u l i f o l i a  s p p .  Herbs pine grass rough fescue Kentucky bluegrass bluebunch wheatgrass lemonweed pussy toes yarrow fireweed balsam r o o t heart-leafed arnica strawberry s t a r Solomon's s e a l twinflower n o r t h e r n bedstraw horsetail  C a l a m a g r o s t i s F e s t u c a Poa  rubescens  s c a b r e l l a  p r a t e n s i s  Agropyron  s p i c a t u m  Lithospermum  r u d e r a l e  A n t e n n a r i a A c h i l l e a  s p p . m i l l e f o l i u m  E p i l o b i u m  a n g u s t i f o l i u m  B a l s a m o r h i z a A r n i c a F r a g a r i a  v i r g i n i a n a  S m i l a c i n a L i n n a e a Galium Equisetum  s a g i t t a t a  c o r d i f o l i a s t e l l a t a b o r e a l i s b o r e a l i s s p p .  101  Appendix I I .  Source of v a r i a t i o n  Cond Plot(Cond)  Anova t a b l e  f o r vegetation  analysis  F-test  DF  2-1=1  o  2  2* (3-1)=4  a  2  Error  2*3(20-1)=114  Total  120-1=119  e  + nka  e  + nka  2 r  2 r  +  rikfyc  102 Appendix I I I . List observed i n c o n t r o l Red-tailed  hawk  Cooper s hawk 1  Buteo  a  a  S h a r p s h i n n e d hawk American Ruffed  kestrel  grouse*  a  a  Great-horned o w l  c  Common n i g h t h a w k  a  swift  Pileated Hairy  Dusky  Common  wren  3  a r c t i c u s  P i c o i d e s  v i l l o s u s  C o l a p t e s  a u r a t u s n u c h a l i s t r i d a c t y l u s  Epidonax  o b e r h o l s e r i  Contopus  s o r d i d u l u s b i c o l o r  chickadee  3  Parus  3  Parus  nuthatch  3  nuthatch  3  c o r a x brachyrhynchos a t r i c a p i l l u s gambeli  S i t t a  c a n a d e n s i s  S i t t a  c a r o l i n e n s i s  T r o g l o d y t e s  kinglet  3  3  3  i n both c o n t r o l s  found o n l y  canadensis  Coruvs  thrush  k= f o u n d o n l y =  p i l e a t u s  P i c o i d e s  Coruvs 3  robin  Swainson's  c  c  b  Ruby-crowned  found  v a u x i  T a c h y c i n e t a  White-breasted  =  minor  P i c o i d e s  3  crow  Red-breasted  3  3  3  Mountain chickadee  American  v i r g i n i a n u s  P e r i s o r e u s  Black-capped  Winter  gnoma  Sphyrapicus  3  raven  American  3  woodpecker  flycatcher  jay  u m b e l l u s  Dryocopus 3  3  W e s t e r n woodpeewee Gray  3  3  sapsucker  Tree swallow  s p a r v e r i u s  C h o t d e i l e s  woodpecker  flicker  Three-toed  s t r i a t u s  Chaetura  woodpecker  Red-naped  A c c i p t e r  Bubo  woodpecker  Northern  c o o p e r i i  G l a u c i d i u m  a  Black-backed  j a m a i c e n s i s  Bonasa 3  c a l e n d u l a  Turdus  m i g r a t o r i u s u s t u l a t u s  and t r e a t m e n t s  i n treatments  t r o g l o d y t e s  Regulus  Catharus  in controls  names o f b i r d s  A c c i p t e r  F a l c o  3  N o r t h e r n pygmy-owl*  Vaux's  o f common and s c i e n t i f i c and t r e a t m e n t s i t e s .  103 Appendix I I I continued. L i s t o f common and s c i e n t i f i c names o f b i r d s o b s e r v e d i n c o n t r o l and t r e a t m e n t s i t e s . Mountain  bluebird  Townsend s  solitare  1  C e d a r waxwing Solitary  vireo  Warbling  vireo  a  Orange-crowned  warbler  warbler  Yellow-rumped  Vesper  junco  sparrow  Brown-headed Western Red Pine  siskin  Cassin's Evening  a  =  3  3  B o m b y c i l i a  cedroum  V i r e o  s o l i t a r i u s  V i r e o  g i l v u s c e l a t a p u s i l l a  Vermivora  p e r e g r i n a  Dendroica  c o r o n a t a  Junco  p a s s e r i n a  h y e m a l i s  Pooecetes 3  gramineus  M o l o t h r u s  3  P i r a n g a  3  L o x i a  3  finch  l u d o v i c i a n a c u r v i r o s t r a  Grosbeak  Carpodacus 3  found i n both c o n t r o l s  p i n u s c a s s i n i i  C o c c o t h t a u s t e s  and t r e a t m e n t s  found only  in controls  =  found only  i n treatments  ,  a t e r  C a r d u e l i s 3  k= c  townsendi  S p i z e l l a  3  cowbird  crossbill  Myadestes  W i l s o n i a  3  tanager  c u r r o c d i d e s  Vermivora  3  warbler  sparrow  Dark-eyed  3  9  Tennessee w a r b l e r Chipping  S i a l i a a  a  a  Wilson's  9  V e s p e r t i n u s  

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