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Deer food production in certain seral stages of the coast forest Gates, Bryan Rodd 1968

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. L E E R FOOD PRODUCTION I N C E R T A I N S E R A L STAGES OF TEE COAST F O R E S T  by  B.Sc.  • BRYAN RODD GATES U n i v e r s i t y o f B r i t i s h C o l u m b i a , 19&2  A T H E S I S SUBMITTED I N P A R T I A L FULFILMENT THE. R E Q U I R E M E N T S FOR THE DEGREE OF ' MASTER OF S C I E N C E IN  THE DEPARTMENT  OF  ZOOLOGY  We a c c e p t t h i s t h e s i s ; the required  THE U N I V E R S I T Y  as c o n f o r m i n g t o standard  OF B R I T I S H . C O L U M B I A  AUGUST, 1 9 6 8  OF  In p r e s e n t i n g the  t h i s th.eais  r e q u i r e m e n t s f o r an  in partial  advanced degree a t the  of  British  it  f r e e l y a v a i l a b l e f o r r e f e r e n c e and  Columbia, I agree that  the  agree - t h a t p e r m i s s i o n f o r e x t e n s i v e for  s c h o l a r l y p u r p o s e s may  D e p a r t m e n t or by  his  be  c o p y i n g or p u b l i c a t i o n  gain  s h a l l not  of  study.  Columbia,  University s h a l l makeI  further'  c o p y i n g of, t h i s the  It'is  this thesis  a l l o w e d w i t h o u t my  D e p a r t m e n t of ZOOLOGY, The U n i v e r s i t y o f B r i t i s h V a n c o u v e r 8, C a n a d a .  Library  g r a n t e d by  representative..  that  be  f u l f i l m e n t . of  Head o f  thesis my  understood  for f i n a n c i a l  written  permission..  ABSTRACT  This stages at  study  was d e s i g n e d  of the regenerating  deer  foods, used  tailed  COdocoileus  deer  coastal levels  British soon  by deer.  reproductive  rumen  success  content  obtained  then  reflated  serai  An  Cover  and w i n t e r quality  ing.  type  summer..  seas-  a r e most  to reach  o f a deer  three  were  blackin  maximum  range i n  influencing  determined,  serai  through  composition  up t o f o u r  estimates  of quantity,  of the important stages..  t o which  food  These  deer  by t h e abundance  salal-catsear  develops  early  as  Richardson)  as a " f a c t o r  preferences  as i n d i c a t e d  association This  suggested  i n different  early  stages  columbianus  The e f f i c i e n c y  to the i n t e n s i t y  stage,  available  have been r e p o r t e d  analysis.  and n u t r i e n t  were  efficient  and t h u s , p o p u l a t i o n d e n s i t y .  forage  h e i g h t , a n d summer  variety  energy  serai  l o g g i n g and s l a s h - b u r n i n g , • and t o d e c l i n e  has been  Seasonal  a r e most  efficient  hemionus  Columbia  after  food  into  which  Populations of Columbian  as. s u c c e s s i o n a d v a n c e s . producing  forest  and i f t h e most  intensively  in  coast  c o n v e r t i n g r a d i a n t energy  onal  t o determine  feet  species  data  utilized of pellet  were each groups.  CGaultheria-Hypochaeris)  to five  years  was p r e f e r r e d by d e e r  after  during  Herbaceous p l a n t s formed  slash-burns p r i n g and  6Q p e r c e n t  of  the  spring-summer  covered  more  in  serai  this  A ation type  stage  winter  12  species and  area,  serai  stage  than, i n  tein,  mineral,  foods  were  these  formed  70  the  clusive,  particularly  in  declines  occurring  ever,  seasonal  and a  in  the  the  in  important  the  covered  forage the  and  autumn-  species,  addition,  in  crude  this pro-  evergreen  than  serai  at  other  stages  where  available.  of  key  was  foods  evidence  plants  and  of  Tests-  serai  fourteenth  years  i s  a  i n i t i a l  influencing  in  range  to  were of  years.  food  by  succession  likelihood four  s i g n i f i -  selection  as  there the  changed  available.  s u c c e s s i o n a l changes  factor  of  available  nutrient levels  since  Shrub  (autumn-winter)  there  fourth  icant  of  cent  more  occurred  most  nutritious  between  by  In  contents  winter.  per  other.  and  declines  be  80  more  nutrient content  advanced  to  to  produced  use  were  forage  .  slash-burning.  and  represented  ash.  available  This'  after  autumn  any  species,  years  during  season,  other.  more  more  associ-  deer,  most  produced  by  f i r CG a u 1 1 h e r i a -P s e u d o t s u g a )  Heaviest  the  represented  any  eaten  demonstrate  believed  in  when  with  of  and  higher  The  deer  15  and  and  evergreens  cantly  than  were  surface  were  area,  to  preferred  diet  times.  and  salal-Douglas  coniferous  more  surface  develops was  diet  inconsignifHow-  quality  selection.  are  It logged at  unit  which  munity  i s concluded of coast  food  t h a t t h e numbers  forest  i s produced.  are affected  Numbers  units  f o r each  suggested  which  states that  will  i n a logged  inevitably  cause  season.  coast  within- a  by t h e  efficiency  within  by t h e a v a i l a b i l i t y  producing  expected  are affected  of deer  the whole  of ideal  A further  1  food-  hypothesis i s  sustained populations forest  because  r e g r e s s i o n of range  com-  serai  cannot  be  succession  quality.  T A B L E OF  CONTENTS  ' •  Page  INTRODUCTION  i  Ob j e c t i v e s THE  STUDY  ,.  AREA  Location • Topography  Climate  5  and- S o i l s  .............. •  Vegetation Logging h i s t o r y  5 5  7  8 10  METHODS OF S A M P L I N G AND  ANALYSIS  14  P e r i o d o f Study Rumen A n a l y s i s Range S u r v e y Range P r o d u c t i v i t y C h e m i c a l C o m p o s i t i o n o f Food S p e c i e s Deer Use o f S e r a i S t a g e s E s t i m a t i o n o f Deer Numbers ..  14 14 15. 17 18 19 22  RESULTS Food  3  '24 Habits Annual d i e t Spring-Summer Autumn d i e t Winter diet  . ....  rC . 5  transitional diet •  24 26 30 32 33  Range C o m p o s i t i o n a n d C h a n g e s w i t h Serai Succession S e r a i c o m m u n i t i e s a t N o r t h w e s t Bay ..... 1) Senecio-Epilobium association 2) G a u 11 h e r i a -Hy p o c h a e r j. s a s s o c i a t i o n .. 3) G a u i t h e r . i a - P s e u d o t s u g a a s s o c i a t i o n .. 4) Pasudotsuga subclimax .•  33 34 39 40 43 44  Species  45  Range  Variety  i n Serai  Productivity  Communities.  48  vi  Page  C h e m i c a l C o m p o s i t i o n o f Deer Forage S p e o i e s The e f f e c t o f s e r a i s u c c e s s i o n o n forage nutrients The e f f e c t o f s e a s o n o n f o r a g e nutrients " Deer  Use o f S e r a i C o m m u n i t i e s Seasonal use i n r e l a t i o n t o s e r a i succession ... The e f f e c t o f s l a s h - b u r n i n g o n range use E s t i m a t e d number o f d e e r i n t h e study area  54 54 57 .. .  65 66 71 72  DISCUSSION  77  L i m i t a t i o n s of the data The r e l a t i o n s h i p b e t w e e n f o o d a n d t h e s e l e c t i o n o f s p r i n g a n d summer r a n g e The r e l a t i o n s h i p b e t w e e n f o o d a n d t h e selection  o f autumn  and w i n t e r  77 82  range  85  CONCLUSIONS LITERATURE APPENDIX  87 CITED  89 . '  9^  LIST  OF  TA3LES  Table  1.  2.  3.  4.  5.  6.  7.  8.  9.  10.  Page  The l o g g i n g h i s t o r y o f t h e 1 0 . 1 s q . m i l e (6,464 a c r e ) s t u d y a r e a a t N o r t h w e s t B a y The r e l a t i v e i m p o r t a n c e o f f o o d i t e m s t o b l a c k t a i l e d d e e r a t N o r t h w e s t Bay as i n d i c a t e d b y s t o m a c h c o n t e n t s - o f 72 a n i m a l s sampled i n three d i f f e r e n t seasons  11  ,.  The t e n d o m i n a n t i t e m s i n t h e a n n u a l d i e t s of C o l u m b i a n b l a c k t a i l e d deer occupying three separate ranges I  27  29  The p e r m a n e n t r a n g e s u r v e y a n d p e l l e t - g r o u p study p l o t s e s t a b l i s h e d a t Northwest Bay, 1959 t o 1 9 6 1  36  Annual production, o f deer foods i n v a r i o u s p o s t - f i r e s e r a i s t a g e s , e x p r e s s e d as pounds per acre wet-weight '.  49  Winter a v a i l a b i l i t y o f deer foods p o s t - f i r e s e r a i stages, expressed per acre wet-weight  49  i n various as pounds  Results of a series of four analysis of v a r i ance t e s t s d e s i g n e d t o determine, t h e e f f e c t o f s e r a i s u c c e s s i o n on t h e n u t r i e n t c o n t e n t of i m p o r t a n t deer f o r a g e s p e c i e s T h e a v e r a g e s e a s o n a l c o m p o s i t i o n o f some important deer forage s p e c i e s a t Northwest Bay, V a n c o u v e r I s l a n d Analysis of Variance f o r differences i n nutrient composition of s i x important deer f o r a g e s p e c i e s c o l l e c t e d I n J u l y , December and M a r c h . Results applied protein for the Table 9  • 55  58  .  62  o f D u n c a n ' s new m u l t i p l e r a n g e t e s t t o t h e s e a s o n a l means f o r c r u d e and n i t r o g e n f r e e e x t r a c t d e r i v e d a n a l y s i s o f v a r i a n c e shown i n 62  Mean i n d i c e s o f serai sites  deer-use in d i f f e r e n t - a g e d  Age c l a s s i f i c a t i o n o f p e l l e t g r o u p s e n c o u n t e r e d d u r i n g t h e r e g u l a r s y s t e m a t i c May c o u n t s o f 1961 and 1962 E s t i m a t e s o f t h e number o f d e e r s u p p o r t e d by e a c h a g e - c l a s s o f r e g e n e r a t i n g l a n d , and the estimated deer p o p u l a t i o n i n the study area ,...  TABLES  IN  APPENDIX  T e m p e r a t u r e and p r . e c i p i t a t i o n r e c o r d s t a k e n a t t h e Nanaimo A i r p o r t , and averages f o r t h e f o u r - y e a r s t u d y p e r i o d Seasonal d i e t s of b l a c k t a i l e d deer at N o r t h w e s t B a y , b a s e d on a n a l y s e s o f rumen samples from female deer . ....... Average cover-percentages f o r the major c a t e g o r i e s of v e g e t a t i o n , the rank of d o m i n a n t s p e c i e s , a n d t h e mean n u m b e r o f p a l a t a b l e species recorded i n burned s i t e s o f i n c r e a s i n g s u c c e s s i o n a l age Average cover-percentages f o r the major c a t e g o r i e s o f v e g e t a t i o n , and t h e rank o f dominant s p e c i e s , r e c o r d e d i n unburneds i t e s o f i n c r e a s i n g s u c c e s s i o n a l age ... N u t r i e n t values of four groups of forage s p e c i e s sampled i n d i f f e r e n t - a g e d s i t e s and a t d i f f e r e n t s e a s o n s .  ix  LIST  OF  FIGURES  Figure 1.  2.  3.  4.  '5.  6.  7. .  8. 9.  10.  11.  Page 'Southwestern B r i t i s h . Columbia showing the l o c a t i o n o f t h e Northwest Bay l o g g i n g c l a i m and t h e i n t e n s i v e s t u d y a r e a  .•  6  A e r i a l p h o t o g r a p h o f a p p r o x i m a t e l y 15 s q u a r e m i l e s o f t h e N o r t h w e s t Bay l o g g i n g c l a i m , i n c l u d i n g the study area  13  Generalized seasonal use-pattern of the m a j o r v e g e t a t i o n , t y p e s by N.orthwest Bay d e e r , as i n d i c a t e d by rumen c o n t e n t analysis  25  .Composition of the seasonal N o r t h w e s t Bay d e e r  diets  ...  of 31  The N o r t h w e s t Bay S t u d y A r e a s h o w i n g t h e major l o g g i n g roads and t h e l o c a t i o n and r e l a t i v e s i z e o f t h e 27 p e r m a n e n t s t u d y plots  35  S e q u e n t i a l development of four major s e r a i a s s o c i a t i o n s and r e l a t e d cover percentages by t h r e e v e g e t a t i o n t y p e s ; slash-burned sites only  38  Dense b r a c k e n f e r n g r o w i n g i n t h e m o i s t l o w e r e n d o f P l o t J , 14 y e a r s a f t e r slash-burning . ,  42  An e a r l y 15 y e a r s  Gaultheria-Pseudotsuga association a f t e r l o g g i n g a n d b u r n i n g ( P l o t M)  .  42  Trends i n the v a r i e t y of p a l a t a b l e deer f o o d s p r e s e n t a f t e r l o g g i n g and s l a s h burning ,  46  The w e i g h t and w i n t e r stages •  51  o f deer i n each  food produced o f the major  i n summer serai  The r e l a t i v e u t i l i z a t i o n by d e e r o f e a r l y s e r a i s t a g e s d u r i n g t h e summer a n d a u t u m n spring period  68  Figure  12.  13.  Page  A S e n e c i o — E p i l o b i ' u m • a s s o c i a t i o n : one y e a r • a f t e r l o g g i n g and b r o a d c a s t b u r n i n g ( P l o t A) .' ..,..  73  An u n b u r n e d  73  area  one y e a r  FIGURES .14a.  14b. 15a.  15b.  16a.  16b.  17a.  17b.  after  logging  IN APPENDIX  A S e n e c i o - E p i l o b i u T O a s s o c i a t i o n two y e a r s a f t e r l o g g i n g and s l a s h - b u r n i n g ( P l o t B) T h e same p l o t burning ..  s i xyears  after  ...  102  slash-  An e a r l y G a u l t h e r i a - H y p o c h a e r i s nine years a f t e r burning  10 2 association  T h e same p l o t a s a n a d v a n c e d . G a u l t h e r i a Hypochaeris a s s o c i a t i o n nine years a f t e r burning ..'  103  ...  10 3  An a d v a n c e d G a u l t h e r i a - H y p o c h a e r i s a s s o c i ation nine years.after logging and-spot burning. (Plot I ) . T h e same p l o t a.s a n e a r l y GaultheriaP s e u d o t s u g a a s s o c i a t i o n 13 y e a r s a f t e r deforestation An a d v a n c e d G a u l t h e r i a - H y p o c h a e r i s a s s o c i a t i o n 12 y e a r s a f t e r l o g g i n g a n d s p o t b u r n i n g ( P l o t K) .. P l o t K as a G a u l t h e r i a - P s e u d o t s u g a a s s o c i a t i o n 16 y e a r s a f t e r l o g g i n g a n d • burning .. ..,..  1°^  .  10 * 1  1  0  5  105  ACKNOWLEDGMENTS  This Fish  p r o j e c t was f i n a n c e d  and W i l d l i f e  Branch,  D r . James H a t t e r ,  am g r a t e f u l f o r t h e s u p p o r t members Dr.  o f the Branch.  tion  thought,  Drs.  Zoology during  D.H.  the preparation Department  statistical  age  and a s s i s t a n c e  of Graduate  Science,  D r . W.D. Science  Studies,  o f t h e Department o f  of' t h e m a n u s c r i p t .  during  assistance.  and c o n s t r u c t i v e  of Poultry  D i v i s i o n of Animal  ities  Cowan, Dean  advice  problems.  to  D i v i s i o n who, w i t h  much v a l u a b l e  C h i t t y a n d H.C. N o r d a n offered  o f f e r e d by  I am i n d e b t e d  Research  I  and d i r e c t e d t h e i n v e s t i g a -  offered  I a n McTaggart  fr^eely  Roberts,  the  designed  a n d who h a s s i n c e  Dr. and  In p a r t i c u l a r  Columbia  Director.  and encouragement  P . J . Bandy, Head, W i l d l i f e  meticulous  by. t h e B r i t i s h  Kitts  criticism  Dr. helped  C.W.  solve  many  and t h e s t a f f o f  provided  laboratory  the proximate  facil-  analysis of for-  samples.  Fellow on  the project  and  efficient  iated.  students  W.B.  Preston  a n d M.A.  as Research A s s i s t a n t s , methods of  Mr. D o n a l d  -  data  collection  Their  Bigg  worked  accurate  a r e much  C. T h o m a s c o l l e c t e d t h e r u m e n  apprecsamples,  which, were  subsequently  Mr.  Smith., p r o v i d e d ' s o u n d  I a n D.  computer  .supplied the  study  advice  was made p o s s i b l e  of the Northwest  Limited,  hy M i s s  Rosemary  Parker.  and a s s i s t a n c e w i t h  programming.  This tion  analysed  which  Bay D i v i s i o n  allowed  unlimited  through  the  of MacMillan  access  coopera-  Bloedel  to their  l a n d and  much o f t h e i n f o r m a t i o n on l o g g i n g h i s t o r y ;  British  Columbia  accommodation  at  Finally, many r e v i s i o n s  Forest  S e r v i c e , which  provided  and summer  Parksville. I wish  t o thank  of the manuscript.  my w i f e , S h a r o n ,  who  typed  INTRODUCTION  It fires few  has been p o s t u l a t e d t h a t  and p r i o r  blacktailed  Richardson) biome  tieth  deer  were  and Hines  stages  Northwest  conversion of climax  began.  these  Marked  workers  creation  industry,  C o l u m b i a n us  coniferous  forest  1945, Leopold  1961).  Early  forests  back  i n c r e a s e s i n deer  to coincide  1950,  i n t h e twenand a  to early  numbers- w e r e  serai noted,  w i t h t h e r e g e n e r a t i o n o f new  f o r favorable responses  of pioneer  and Derby  improved  nutritional  1932,  Buckland  Hines  1959),  preferred  floral  (1955),  and abundance  factors. that  and have  quality  1 9 4 1 , Cowan  as w e l l  food  have  been  been  plants  t o an  Increased  and shrubs  nutrient  regard, recent growth  1 9 5 0 , Dasmann and content  suggested  of the  larger  and  Hat-  as c a u s a t i v e  investigations  rates,  vari-  (Storer  ( E i n a r s e n 1946, Cowan, Hoar been  to the  d i s c u s s e d by  attributed  1945, Leopold  1956) have  more r a p i d  by u n g u l a t e s  of the range.  as improved  species  In this  conditions  of herbaceous  1 9 5 0 , a n d Swank  reveal  wild-  -  Reasons  ter  (Cowan  1959, and Brown  growth.  ety  by t h e m o i s t  of  century extensive logging operations developed,  wide-spread  DeWitt  o f the. l o g g i n g  (Odocoileu's hemionus  supported  of the P a c i f i c  Dasmann  .by  to the development  i n the absence  body  with  deer  size, i n -  creased from  productivity,  and  greater  population  a better nutritional  regimen  (Taber  Dasmann 1965,  1958,  Murphy  It erating  Severinghaus and  can  food.  at  land  which  Efficiency  deforestation, proached  to  number  of  ideal  numbers energy  Deer  Therefore, and  largest  Ullrey  of  deer  are  Klein  i f other  be  available  soon  after  be  or  ap-  expected  factors  constant  such  have  expected  u n i t s f o r each  the  into  c o n d i t i o n s are  n u m b e r s may  can  regen-  a f f e c t e d by  greatest  climax  populations  food-producing  1965,  a l . 1967).  i s converted  hunting.are  deer  Taberand  Bandy  et  result  therefore, that within  d e c l i n e as  1961).  weather,  effects,  and  1956,  1964,  i s b e l i e v e d t o be  and  (Brown,  the  solar  accordingly.  dation,  1966,  Tanck  be. h y p o t h e s i z e d ,  u n i t s of  efficiency  vary  Coates  and  densities  to  as  pre-  negligible when  season  the  i s at  a  maximum.  Within be  reflected  should  occur  regenerating  by  the  greatest  food  Decreasing  efficiency of  species  deer  tive  quality,  when  l a r g e amounts  or  and  occupies  should  use.  of  unavailable plants .  be  the  become  efficiency deer  of  occur of  use.  •  nutritious  greatest  reflected  coverage  energy  of  variety  I t should  surface  optimum  intensity  w h e n ' the' g r e a t e s t  productive  intensity  units,  by  surface a  trapped  decrease, in  It and area.  declining  when v a r i e t y ,  foods  should  nutriand  unpalatable  Studies ed  of  to b l a c k t a i l e d  and  Dasmann  ent  within  a  Brown  single  are  (1961),  serai  and  in isolated  t o work  Klein  thus  by  Taber These  ranges  of  demonstrating  herds.  to  relat-  (1965).  separate  deer  were- k e p t  t o measure  f o r a g e , as  limited  structure,  influences  attempted  q u a n t i t y of  c o n d i t i o n s on  differences  extraneous study  or  and  numbers,  compare  biotic  biological that  deer  (1958),  investigations different  quality  In  order  a minimum,, t h e  t h e s e . f a c t o r s as  they  pres-  occurred  range.  Obj e c t i v e s  Specifically of  changes  of  the of  test  the  detail: range  -  forest,  range  and  selection  (1)  the  by  seasonal  and  progresses;  (3)  ferent  communities;  serai  and  season  and  (5)  ities.  food  the  i n the  the  these  aspects  magnitude  of  (2)  (4)  use  the  the  changes  content by  deer  of  pat-  occupying  the  as  a  to  investigated  deer  for defining  to  areas  In order  were  h a b i t s of  nature  burned  deer.-  p r o d u c t i o n of  nutritive  and  changes  v e g e t a t i v e cover  annual  e x p l o r e the  i n logged  various plant species;  composition  to  blacktailed  information essential of  were  to relate  basic hypothesis, five  importance species  purposes  i n v e g e t a t i o n growing  coast  tern  the  in  the  relative changes  serai  deer  succession  foods  with  important  in  in  dif-  succession deer  of v a r i o u s s e r a i  foods; commun-  ' Theoretically, within  a  single  the  block-logged was  of  regenerating  extended 'period of ticable,  each  time.  4 these  unit  In  of  order  aspects  l a n d , but  only  be  of  studying  a number  units,  at  a different  stage  chosen.  c  studied  over  t o make .the. s t u d y  alternative each  could  an  prac-  of  adjacent,  of  regeneration,  5  THE  STUDY  AREA  Location  The  privately  owned  of  MacMillan  Bloedel  Division mately 18  135  miles  land  square  northwest  lies  cludes  on  the  Englishman west on  by  the  Okay  and  of  Nanaimo  the  Vancouver  (Figure  Englishman  Mt.  and  f t ) and  DeCosmos  middle  Mt.  Island,  1).  River  The  and  Arrowsmith  by  south  of  (5283  Moriarty  the  of  side  approxi-  I t i s bounded  and  area  f o r k s of  (4444- f t ) , a n d  on  at  A  s e l e c t e d f o r i n t e n s i v e study  i n the  middle  Topography  graphy  city  east  Bay  inthe  on  the  (5962 f t ) ,  the  east  by  (2826 f t ) .  centered  river  L i m i t e d encompasses  the  side  Northwest  M o r i a r t y Creek.  miles  prevail  south  the  River  The  south  the  of  of  Mt.  Mt.  the  of  on  land  drainages  south  square  miles  forest  northeast  f o r k s of  coordinates  and  Englishman  13'N  and  n o r t h e r l y aspect  throughout. and  49°  lying  between  River.  124°  ten  the  It  Is  18'W.  Soils  general  banks  the  corner,  included  A  few  to minor  i s intermediate  steep  areas  between  of  and  gently  slopes rock  that  are  rolling confined  outcrop.  typical  of  slopes to  the  Physiothe  Georgia  Fig.  .1.  Southwestern B r i t i s h Columbia showing the l o c a t i o n o f t h e N o r t h w e s t Bay l o g g i n g c l a i m and t h e i n t e n s i v e s t u d y a r e a ( c e n t e r ) .  7  Depression  (Nanaimo  typical  the  of  (Holland,  Outer  1964).  north-east  Lowland) of  corner  Mount a i n A r e a  ten  1960)  miles  are  Parent  with  south-west  1600  of  the  colluvial  soil  south  and  have  materials.  climates  with  dense  slightly  acid  and  from  i n the  groups  possess  Island the  corner.  Nanaimo  the  study  Lakes  area.  P l u t o n i c g r a n i t e and rock.  Glacial of  belong  to  cover.  the  They  a relatively  the  Upper drift  soil,  rock-outcrop-  characteristic  vegetative  that  Mueller-Dombois  c o n t r i b u t e d most  soils  and  f t -i n  c o n t r i b u t i o n s from  The  Gley  500  (Mc'Minn 19 57 ,  sedimentary  localized  Brown P o d z o l i c and  profiles  derived  deposits  and  f t i n the  r e p r e s e n t a t i v e of  sandstone  minor  Vancouver  to  m a t e r i a l s are  alluvial  of  from  considered  Cretaceous and  to  C o a s t a l Trough  . E l e v a t i o n s range  Descriptions area  the  thin  Brown of  or  warm,  are  well  organic  Redmoist drained>  layer.  Climate  Northwest couver  general  ably ed  at  1965).  during  tation  lies  I s l a n d Mountain  Guiguet, port  Bay  the  i n the  slightly Nanaimo.  Range  Weather four year  indication  of  study  This  12  field  than  i s based  shadow to  taken  study  the at  during  average on  the  of  the  (Appendix  that of  the  west  conditions.  itself the  rain  miles  records  climatic  area  higher  i n the  (Cowan  Nanaimo A)  period  a  precipiwas  inches  observations  and  Air-  provide  Annual  42.6  Van-  of  probrecordMueller-  Dombois a  ( 1 9 6 0 ) , who  significant  as  west-east  m u c h a s . a . 50  tance  of  14  reported  inch  that'in  precipitation  decrease  miles.  the Nanaimo gradient  in precipitation  Northwest  Bay  appears  Lakes  area  exists,  with  over  a  dis-  to experience  the  same g r a d i e n t .  Temperatures indicated and  by  a four  Airport. tions area  are  a four  500  y e a r mean J a n u a r y  Here  a g a i n , mean t e m p e r a t u r e s t o have  temperature  b e e n more  a t Nanaimo A i r p o r t ,  to  1600  from  the  the  stabilizing  f t higher i n  of  64.7 and  extreme  since  the  year  as  of  39.1  temperature  mean J u l y  further' inland 'is  throughout  year  assumed  than  are moderate  F at  Nanaimo  daily i n the  former  influence  F  fluctuastudy  i s two of the  miles sea  and  elevation.  Vegetation  Vegetation Zone The  described climatic  CPseudotsuga climax red  menziesii  ('Thuja  the  canopy.  imately range  of  50 , 000 from  (1959)  (Pinus  balsam  F i r Bioclimatic.  Mueller-Dombois mainly  (Tsuga  (1959).  of Douglas f i r the  edaphic  heterophy11a ) , western  f i r (Abies grandis)  and  c o n t o r t a ) c o m p r i s i n g the remainder  The  timber y i e l d  feet-board-measure 35 , 000  Douglas  and  consists  hemlock  plicata),  pine  of the  var m e n z i e s ii ) , with  western  lodgepole upper  Krajina  climax forest  species  cedar  by  is typical  f .b.m.  from  the  area  (f.b.m . ) per  t o 60.000  f.b.m. .  is  acre,  of  approxwith  Species  a  com-  position ern  averages  hemlock,  balsam eau,  fir,  and  Forest  Ltd.,  10  70  per  per cent  1 per  Engineer,  pers.  comm.  The  cent  Douglas  western  red  cent  western  N.W.  Bay  fir,  17  cedar,  white  Division  of  and  subclimax  understory  grape  (Mahonia  berry  (Arctostaphylos uva-ursi), pipsissewa  described be  by  nervosa),  form  or  these lichen  of  Krajina among  salal  (E.  west  TouzBloede  the  lesser  rose  (Linnaea  amounts  (Rosa  form  of  borealis).  either  following  bear  (Chimaphila  the  virgin  Mueller-Dombois  area,  by  Oregon  gymnocarpa),  a s s o c i a t i o n s of  study  i s dominated  (1959)  in their  and  can  natural  destruction.  (Pseudotsuga-Gaultheria)  forest  Two  the  of  salal-  (Pseudotsuga-Gaultheria-Peltigera) associations -  completely  ment  i n the  in residual  - the  dwarf  twinflower  of  with  K r a j i n a ( 1 9 5 2 ) and  recognized  cent  MacMillan  (Gault'heria shallon ),  seven  2 per  pine.  salal  All  cent  1963).  climax  u m b e l l a t a ) , and  per  willows  the  others.  Logging  and  slash-burning  most  the  edaphic  of  ( 1 9 5 9 ) and  these  (Rubus  dominate  are  radicata),  serai  Mueller-Dombois  red  vitifolius), (Salix  and  have  alder  (A1nus  encouraged . e s t a b l i s h species  (1960).  black.raspberry  (Rubus  everlasting  (Anaphalis  by  Noteworthy  rubra ), t r a i l i n g  species), hairy catsear  pearly  listed  blackber  leucodermis),  (Hypochaeris m a r g a r i t a c e a ),  vanilla  leaf  (Achlys triphy11a), fireweed  folium ) , bracken  fern  ( P t e r idiurn  a q u i l i n u m ) , and  ( P o l y s t i c f a u m mu n i t urn ) .'  .Logging  end  an  first  i n 1946,  work  of the mature  average  nual  began  annual  forest harvest  slightly  more  than  Logging stands  was  2300  by  was  removed  o f 3.5  acres  years  to ing  a maximum more  to ten adjacent  o f two  typical  the study.  subjected  intensity  and  thorough  vegetation  As  to at least  and  was  period  June,  remained.  30 t o  loading site.  adjacent  were  years  an-  1962  35  drag-hauling with a  settings  broadcast  practised  a result, some d e g r e e  of the f i r e s  was  per  spar  In the  cleared this  settings,  in a  was  reduced  thus  becom-  of patch-logging.  Slash-burning during  By  the  maximum  of selected  In the l a t t e r  or three  The  and  until  .Sixty-four  per cent.  f o l l o w e d by  early  operation.  year  of mature f o r e s t  or b u l l d o z e r to a roadside  continuous  i n 1945,  i n t h a t 18 y e a r  22 p e r c e n t .  cable  up  each  ( T a b l e -I)..  clear-cutting  (settings),  area  was' r e m o v e d  i n 1962  h a r v e s t , i n 1 9 4 8 , was  acre  - •.  i n the study  some t i m b e r  of the f i e l d  cent  •  swordfern  History  Logging except  (Epilobium angusti  varied  burns  regularly  almost  before  a l l settings  of burning. greatly,  i n which  destroyed., to l o c a l i z e d  from  almost spot  The  and were extent  effective  a l l slash burns  in  and which  11  TABLE  1  The  logging  acre)  Annual Acres  Year 1945 1946 1947 194 8 1949 1950 1951 19 5 2 1953 1954 1955 19 5 6 1957 1958 1959 1960 19 61 1962  and  99 . 9 nil 266 . 2 1436. 8 159 . 2 494 . 1 171. 9 390 . 1 205 . 5 73 . 8 26 . 4 179 . 1 92 . 5 184 . 4 103 . 1 121. 0 138 . 4 14 . 3  from  history  of the  10.1  study area at Northwest  Cut %  1 . 6 nil 4 . 2 22.2 2 . 5 7 . 6 2 . 7 6 . 0 3 . 2 1. 1 0 ,4 2 . 8 1.4 2 . 8 1.6 1 . 9 2 .10.3  sq. mile  (6464  Bay.  Accumulated Cut  Remaining Timber  1.6 1. 6 5.7 27.9 30 . 4 38.0 40.7 46.7 49.9 51. 0 51.4 54 . 2 5 5.6 58.5 6 0,. 1 62.0 64 . 1 6 4.3  98.4 98 . 4 94.3 72.1 69.6 62.0 59.3 53.3 51.1 49 . 0 48.6 45 . 8 43 . 4 41. 5 39 . 9 38.0 35 . 9 35.7  D a t a d e r i v e d f r o m l o g g i n g c o m p a n y o p e r a t i o n s maps p l a n i m e t r i c m e a s u r e m e n t s o'f a e r i a l p h o t o g r a p h s .  12  only  portions  of the l i g h t  s l a s h and v e g e t a t i o n  were  destroyed,  Conifer seed  drift  from  regeneration adjacent  some l a r g e , e x p o s e d Douglas ing  f i r seedlings  from.370  eer,  pers.  By created. ities, were  comm.  1962  serai  Only  reforested with  inches  high) CE.  upon  natural  portions  of  two-year o l d  at d e n s i t i e s  Touzeau,Forest  varyEngin-  1963).  a. m o s a i c o f d i s c r e t e l a n d different  (Figure  o f edge stages  were  primarily  trees.  trees per acre  Seventeen  present  floral  t o 450  mature  (10-14  interspersed with  abundance of  areas  depended  2).  was  was  composition,  u n i t s had  age-classes  irregular  stands  of serai of v i r g i n  communforest  A highly varied habitat with  created  f o r deer,  and  a v a i l a b l e f o r the study food  been  productions,  and  a wide of  choice  changing  deer-use.  an  Fig.  2  A e r i a l p h o t o g r a p h , o f a p p r o x i m a t e l y 15 s q u a r e m i l e s o f t h e N o r t h w e s t Bay l o g g i n g c l a i m , i n c l u d i n g t h e study area. M a t u r e and subraature f o r e s t stands o c c u p i e d a p p r o x i m a t e l y one t h i r d o f t h e a r e a w h e n the photograph was t a k e n i n J u n e 1 9 6 2 . (Photograph by A i r D i v i s i o n , P r o v i n c e o f B r i t i s h Columbia).  1  METHODS OF Period  of  field  1962, w i t h b r i e f  devoted  work  was  ANALYSIS  conducted  i n t h e s u m m e r s o f 19 5 9  p e r i o d s i n March and December  to the c o l l e c t i o n  analysis.  Ruman  AND  Study  Most to  SAMPLING  of plant  Rumen s a m p l e s  were  samples  1962  f o r chemical  c o l l e c t e d . i n 1963  and  1964.  Analyses  Rumen s a m p l e s  from  seventy—five, female  analysed  to determine  animals,  r e p r e s e n t i n g a l l a g e - c l a s s e s from  12-1/2  years  seasonal diets  (Thomas, 1963 ) , were  winter,  and i n the t r a n s i t i o n  feeding  seasons  One-pint preserved  d e s c r i b e d by  samples  i n the f i e l d  1958) and s u b s e q u e n t l y and  N'o . 5 s c r e e n s .  was  separated manually  secting  microscope  possible, ies  of  trees,  while other  at Northwest  collected  material  and  plants  U.S.  months  and  identified  identified  and  Standard  remaining  to and  summer  Dasmann, No.  on b o t h  3-1/2 screens  with the a i d of a  and f o r b s jwere  were  The  dig'e.sta 'were  (Taber  and r e f e r e n c e p l a n t  shrubs,  Bay.  i n t h e autumn  of thoroughly mixed  over  were  (1945).  i n 10°£ f o r m a l i n  The  five  between the s p r i n g Cowan  washed  deer  collection. identified  dis-  Where to  only to family.  specNo  15  attempt fine  was  made t o  r e s i d u e was  The ured  identify  listed  separated  volumetrically  as  grass  and  fragments  were  t o .the n e a r e s t 1945,  Brown  and  of  three  collection  centages  and  The three  average  most  periods  "consumption  For  this a  l e n g t h of This  combined  food  were  the  one  Island- b l a c k t a i l e d  and deer  determined  obtained  provided  species  consumed  during a total  spring  sampled,  and  and  late  on by  the  an  summer  therefore could  water  species  b a s i s of  Cowan  from the  cit.).  the  for  preferences  not  entered  to of  Vancouver consumption  f o r the  forage  was  May  mid-point  nine  of  period  early  The  rank  p e r i o d of  per-  (1945).  transition  with  importance  be  by  a modification  d e s c r i b e d by  ( C o w a n , op_.  meas-  calculated.  summer f e e d i n g s e a s o n s  indices  Early  plants,  closely  each  were  o n e - h a l f months,  corresponds  spring  For  and  p e r i o d s , frequency  spring-summer  and  millilitre  1961).  percentages  i n d e x " method  calculation  mid-June. the  important  sampled,  the  given  volume  sponge-dried  half  CCowan the  s p e c i e s , and a l l  "unidentified".  displacement f o r each  sedge  food  months.  into  were the  not compu-  tation.  Range  Survey  Permanent representative  range  stands  survey  of mature  plots  were  timber  and  established in a  in  chronological  16  s e q u e n c e 'of r e g e n e r a t i n g slope,  constant  community ous  were  and  selected  so  were  The a  aspect,  environmental  age-classes  surveyed  cover  (1954)  all  possible.100-foot were  through  the  of  along  .a t a u t  "hit"  between  (the  normal  The recorded even  as  though  significant after area  the had  vari-  of  minimal.  the  (see  ground  at  each  level  i n those  burn. been  and  ber  growing  few  of  sample  the four  their  successional  seasons  feet of  since  were "was  logging.  ground  for mark  or  item  level  recorded,  sites  since  was burning,  in early  spring  occurred  immediately  less  than  considered recorded  a  line  foot  plant  d e e r ) was  seasons  have  age  each  above  burned-over  i n which  fire  at  uppermost  p l o t s burned  to  base  size requirements  below);  from  (usually  transects  g r o w t h may  subjected  a base'line  of  growing  Those p l o t s  random  number  of  of  at  by  Dasmann  the  successional  amount  selected  from  the  height  number  follows  to  and  of  determined  angles  only  age  was  right  maximum b r o w s i n g  the  site  as  were  Methods  tape  Replicates  possible.  modified  plot;  by  100-foot  sites, of  run  the  governed counts  i n t e r a c t i o n s between  i n t e r v a l s along  r o a d ) and depth  plant  be  : transects  logging  group  that  homogeneous  wherever  method  Blaisdel  pellet  a relatively  composition  and  was  S i t e s e x h i b i t i n g minimum  f a c t o r s would  l i n e - p o i n t survey  selected  units.  half  an  in-  the  unburned as  the  num-  Cover-percentages determined describe  f o r each  Data  separately. surveyed  were  samples  and u n t u r n e d s i t e s  years after  their  composition ratings  used t o  were plots  were  treated were r e -  original  sur-  f o rpreviously  un-  Productivity  burned  plant  p r o d u c t i v i t y , was e s t i m a t e d  survey plots Four were  i n December.  sampled  25 f t l o n g ,  at  from  survey  selected  h i g h were  belts  sampled  i n the plot,  from  belt-  each  selected  100 f t r a n g e i n A u g u s t , two  2 f t wide  W i t h b o t h methods  a n d De-  and 4 f t  an average which  was  o f 0.10 available  clipped.  Clippings  weighed  i n July  was c l i p p e d  100 f t l o n g ,  c e n t o f a l l new g r o w t h  ported  sampled  a n d 4- f t h i g h ,  In those plots  sampled.  d e e r , -was  two i n August and  one-quarter of the permanent  transects.  randomly  2 f t wide  i n ten of  s i x d i f f e r e n t age-  i n July,  year's growth  transects random  representing  In those plots  cember, t h e c u r r e n t  to  a n d mean v a l u e s  were  age—classes.  classes.  per  and i t e m  I n t h e summer o f 1 9 6 2 , s e l e c t e d  Annual  four  f o r burned  thus providing  Range  species  f o r which r e p l i c a t e  one, two, or three  surveyed  the  study plot  those age-classes  obtained.  veys,  f o r each  were  placed  in air-tight  t o t h e l a b o r a t o r y , f o r -weighing. separately  and t o t a l  production  bags and  Each  trans-  species  was  was e x p r e s s e d i n  18  pounds in  net  weight  greater than  per  w e r e . r a t e d no  Cowan  (1945) were  sis,  of  taken  from  Four for  sampling,  nine,  and  infra-red  tight  to  remained  were  a. 40  bottle-,  each  collected  combining  serai  analythe  with annual plot.  A  small  growth  represen-  sub-samples  aluminum of  plants after was  ground  s t o r e d a t room  four,  March. 12  hr  and  than the  five,  collection,  Moisture hr  Deter-  in a  heated  five  were  Clippings  of  f o r 24  foil  less  chosen  Collections  (Ohaus  dried  were  undergone  and  determined  mesh s c r e e n , t h o r o u g h l y and  had  December  average  sample  production.  during  communities  which  Samples were  i n the  by  f o r proximate  a laboratory within  An  palatable"  of r e g e n e r a t i o n .  c o n s t r u c t e d of  lamps.  or  plants.  August,  contents  Subsequently, through  July,  material  same s u r v e y by  individual  Balance).  oven  moisture  obtained  fourteen years  moisture  porary  the  samples  food  supplemented  sites  transferred  mination  from  were  including  were  rumen  "moderately  foods  of  i n June,  i n the  appeared  Species  age-classes  made  and  study  was  many  Food  sufficient  randomly  sample  of  important  browse-production collected  than  species which  included in calculating  provide  specimens  tative  lower  Composition  To  Those  trace quantities  which  Chemical  acre.  tem-  by  per  cent  drying period.  i n a Wiley mixed,  temperature  Mill  placed until  to  i n an  pass air-  analysed.  3  19  Procedures, o u t l i n e d Agricultural analysis. the  Chemists Nitrogen  difference  centages Results the  were  drying  tested each  free  t h e . A s s o c i a t i o n , o f Of f i c i a l  were  followed  extract  1Q0  CN , F , E. ) was  per cent  f a t , crude  process  between  and  and  protein,  expressed  differences serai  crude  nutrient  into  and  arranging  a randomized  tested  an  the Computing Center  of variance  and  ash.  between were  values f o r  Percentages  transformation  of the U n i v e r s i t y  after  basis.  seasons  library  as  per-  remaining  percentage  design.  by means o f • t h e . a r c - s i n e analysis  of the  fibre  levels  between  proximate  calculated  on a d r y w e i g h t  transformed  at  t h e sum  in nutrient  stages,  f o r • s i g n i f i c a n c e by  with  f o r the  corrected, f o r r e s i d u a l moisture  Over-all species,  C1960)  between  of crude  by  were and  program  (MFAV)  of B r i t i s h  Colum-  bia.  Deer  Use  of Serai  Stages  Accumulated determine Circular tered  plots  line-point  the  0.8  p e l l e t group d e n s i t i e s  t h e r e l a t i v e u s e by  at every  erence  •  100  transects  per cent  (1958),.an  sq f t i n area  hundred-foot  stakes.  deer  and  (radius  marked  Sufficient plots sample  of various 5.64  i n t e r v a l along  were  were  used  serai  to stages.  f t ) were  the permanent .  f o r r e l o c a t i o n by  were  s i z e recommended  i n t e n s i t y designed' to provide  established by  cen-  t o meet  Robinette  at.least  ref-  70  e_t a 1. per  20  cent  c o n f i d e n c e with, t e n per  When e a c h September moved. deer  of each May  use;  use.  plot  counts  group  fication  of  Although  somewhat  counts  arbitrary,  "fresh"  have  been  and  dropped  subsequent were  assumed  were  last  groups plots were  within  to the  start  t o have  cleared  counted were  cleared  been  to accumulate  count  (the "time-lapse",  tive  (groups use  this  by  per  months  deer , and 1  was  index, deer latter  unit  per  the  growth; between  March.  groups  had  of  fresh",  season  of  Age  not  of  t o be square  age or  classi"aged".  assumed  the  "aged"  the  time  the  of  the  t h u s , a l l groups summer.  of the  Since this  in describing  group  converted to the  factor densrela-  time-  I t i s understood deer  time  subsequent  no  expresses r e l a t i v e  plots  classification since  -  groups  pellet  mile.  to  counting period  observed value  deer  deposition.  calculation  1958).  spring  summer  r e c o r d e d i n each  Hazzard,  re-  p r o v i d e d a.  of  necessary and  and  autumn t o  a s s i g n e d an  facilitated  a c r e ) were  counted  f r e s h " : g r o u p s were  constant for a l lplots,  constant that  the  of  d e p o s i t e d i n . the  of groups  taken  ities  were  categories  t h e p r e v i o u s May  known t o have  not  these  dropped  early  and. a g a i n i n M a y . a n d  "moderately  of plant  i n September  Removal  was  two  been  and  was  approximate  "moderately  error.  p r o v i d e d measures  i n May  "fresh",  means o f d e t e r m i n i n g t h e The  groups  p r o v i d e d measures  counted  either  sampling  established,  year, pellet  September  Each  was  cent  use  rather  than  the density  study  plot.  formula,  based  actually  calculation  m o d i f i e d from  where.  Factor  Its  of animals  was  Razzard  Cop,  the  number  of deer  a  the  number  of p e l l e t  b  the  time-elapse i n days  c  the  daily  per square  13 p e l l e t  r e p o r t e d by Rasmussen (1955) and E v e r h a r d t  Initial  i n each  ations  counts because  groups  and Van  t h e t i m e - l a p s e c o u l d n o t be  an a t t e m p t  to demonstrate  exploitation  attributable  burned  were  e q u a l age were  tested  Das-  (1956).  i n the  calcul-  regarding  i n the coast  forest,  determined.  differences  i n deer  to slash-burning, results f o r  analysed  Between-treatment  (1943),  available  pellets  deer-day  Etten  were  of deer  per  a n d Doman  data  thus  of  deer.  insufficient  and  sites  of  not i n c l u d e d  rate  (per day)  per . acre  were  decomposition  In  mile  plot  the  sites..  groups  defecation rate  mann a n d T a b e r  following  t  X  on work  given  c j t );  a x 640 a c r e s b x c  given the value  any  b y means o f t h e  X  c was  occupying  s e p a r a t e l y from differences  those  i n deer  for significance  of  unburned  use f o r s i t e s  w i t h a t_ T e s t f o r  paired observations.  Trends determined  by  i n use. w i t h a d v a n c i n g plotting  serai  s u c c e s s i o n were  mean u s e — i n d i c e s , a g a i n s t ' s u c c e s s i o n a l  age.... • R e g r e s s i o n c u r v e s  were  fitted'.through  these  p o i n t s by.  22  means o f a . p o l y n o m i a l ing  Center  ysis  of the University  was programmed  premise  that  deer  immediately Estimation  tial  of the land density  were  area  to support  was c a p a b l e  rived, for  that  square  square  required  miles  occupied  combined  with  logging  age  estimates  needed.  was  found- t h a t  within  any l o g g e d  age-class  history  According  of a  o f deer de-  equation.  o f t h e number  information  density area  site  The  of deer  by t h e number o f .Planimetric  taken•in.1962  As a n e x a m p l e  of- l a n d  deer  the study  the density  age-class.  photograph  the following  or burned  of land  poten-  use o f the p e l l e t  upon  meas-  (Fig. 2),  provided  the acre-  of the calculation,, i t  i n t h e summer o f 1 9 6 2 t h e r e  mile)  site  numbers o f  f o r obtaining  of land  by t h a t  an a e r i a l  regeneration.  sizes  was t h e p r o d u c t  from  square  on t h e  the ecological  by the " p o l y n o m i a l  f o r each  total  I t made  of supporting  urements  (0.29  deer.  sample  age-class  mile  use o f a  anal-  burning.  a n d was b a s e d  and ( 2 ) t h a t  given,age  per  The  the o r i g i n  f o r estimating  i n t h e 18 a g e - c l a s s e s  calculation  through  and f o r m e a s u r i n g  Cl.) t h a t  adequate,  Columbia.  Numbers  information  assumptions: indices  and  was d e r i v e d  i n the study  CPFIT) a t t h e Comput-  of B r i t i s h  f o r regression  logging  o f Deer  deer  program  do n o t make s i g n i f i c a n t  after  A method  group  library  were  w h i c h had undergone to the polynomial  184.4 four  results,  acres  years- o f four-  23  year to  old sitea  140  0.29 or old  deer square  about  41  sites.. :  classes  were' u s e d  per  square  The  deer  mile.  m i l e s x 140 deer  by  deer  p r o v i d e d the  of  intensity  equivalent  Therefore, per  were p o t e n t i a l l y sum  a t an  square  = 40.6  s u p p o r t e d by  t h e p r o d u c t s , so  estimate of  mile  total  derived herd  deer,  the f o u r - y e a r for. a l l  size.  age-  24  RESULTS Food  Habits  Interpretation sition, ity  of  serai  to  of  d i e t s of digesta  demonstrate  other  ranges  1958,  Brown  shrubs  and  preferred quantity  Fig.  that  1961,  and  animal 75  the  the  But  under  trees  and  winter.  volume,  c o n t r i b u t i o n to  the  shrubs  of  During  the  spring-summer  cent  of  annual  the  bulk.  peak, while  During  the  dominated  the  major  the  Grass shrub,  autumn,  1947,  Taber  The  anal-  knowledge Bay,  and  as  on  Dasmann  p r i m a r i l y upon  Herbaceous  and  and  trees other  plants  are  are  taken  plant  in  types  unavailable  make, t h e  greatest  year-  diet.  use  forbs  annual  Northwest  otherwise  Seasonal 3.  the  sufficient  coniferous  compo-  availabil-  at  food.  and  and  of  depend  These  necessary  by  deer  1961)  for  range  investigation.  Chatelain  McCullough  supply  a knowledge  blacktailed 1945,  with  production  rumens p r o v i d e d  succulent,  during  available  an  from  (Cowan  when  nutrients. round  the  deciduous  undoubtedly  per  succession,  factors dealing  n u t r i e n t s depends upon  seasonal ysis  of  plant  transition  rumen and  samples,  sedge  c o n i f e r , and  shrubs  classes  i s shown  period  newly  contributing  consumption l i c h e n use  replaced  in  forbs  as  59  reached were  the  low.  main  Fig.  3..  S e a s o n a l p a t t e r n o f use o f t h e m a j o r v e g e t a t i o n t y p e s by N o r t h w e s t Bay d e e r , as i n d i c a t e d by rumen c o n t e n t a n a l y s i s .  component, the palatable.  latter  but  73  of  cent  increased sedges  largely  M u s h r o o m s made up  autumn d i e t per  becoming  were the  decreased  important  samples.  slightly,  only  and  the  slightly,  u n a v a i l a b l e or  seven  i n that The  per  they  cent  between  the  of  the  were'present  consumption  consumption  un- .  of  in  conifers  of grasses transition  and and  autumn  periods..  In,winter five but  plant  annual forbs  of  less  bulk  markedly  samples  noted.  than  i n autumn.  Grasses,  were  s t i l l  use  of  remained  sedges,  and  forms the  the '  dominant  C o n i f e r s and  w i t h both  were more.abundant but  i n the  Shrubs  i n importance,  high , levels. available  autumn  greatest diversity  c a t e g o r i e s was  provided  increased  the  lichens  reaching  few  perennial  i n w i n t e r samples  relatively  unimportant  of  (1945)  than in  in  terms  bulk.  Annual  diet  index"  method  Appendix during are  B,  more  to  application the  shown  i n Table  cent than  important  of  top  period.  any  habits  importance The  results  2.  a l l food  than  relative  of  data  "consumption present  of  foods  of  the  in  consumed computations  -  eleven  a third  Cowan's  seasonal food  g i v e s the  a nine-month  The per  - The  sources  consumed the  other  total  account i n that diet  species.  for approximately period.  and  was  Red  cedar  Salal  three  formed  times  (.11.per  92  more  cent).-  27  Table  2  The r e l a t i v e i m p o r t a n c e o f f o o d i t e m s t o b l a c k t a i l e d ' d e e r a t N o r t h w e s t Bay as i n d i c a t e d by s t o m a c h c o n t e n t s o f 72 a n i m a l s s a m p l e d i n t h r e e , d i f f e r e n t , seasons.  Season  Spring-Summer Transition  Length of season i n months  Itern  Autumn  1-1/2  3  Seasonal Consumption  Salal 8 .'1 Red Cedar 2. 9 Arboreal lichens 11-. 0 Pearly everlasting 69 . 0 Grasses 2 2'. 6 T r a i l i n g b l a c k b e r r y • 6-. 9 Catsear 12 . 2 Mushrooms T Douglas f i r 3 .5 Bearberry 4 .1 • Red a l d e r T Other vegetation 9 .8  167 ,. 7 6 ,. 9 ' T 0 7 .,2 21. , 9 4 ,. 5 21, . 0 11. . 7 3 ., 3 ' 21., 6 34 .. 2  the  Total  4-1/2  9  Factors  Consumption Index A c t u a l 2 ch  136 ,. 4 94, . 0 59 ,. 4 . 0 36 .. 9 16 ,. 6 17 ,. 6 12, . 6 18 .. o . 23. . 0 3 ., 2 32 ,.4  312 .. 2 103 ., 8 70 , . 4 69 ., 0 66 ., 7 45 ., 4 34, , 3 33 ,, 6 33 ., 2 30 ., 4 24. , 8 76 ..4  Totals  T  Winter  34, . 7 11, •5 7, .8 7, ,7 7 .. 4 5 , .0 3, .8 3 ,8 3 ; .7 3 ,, 4 2 ,, 8 8 ,, 4  900 ,, 2 100 ,, 0  = . Trace ^ P r o d u c t of average volume % f o r each season number o f m o n t h s r e p r e s e n t e d by t h a t s e a s o n . 2  (1945).  Sum  of  Seasonal Consumption  Factors.  See  by  Cowan  28  ranked  second  pearly  everlasting  per  cent).  foods, ifers per  per  cent,  cent  Of  shrubs 15  was  f o l l o w e d by  ?  C8  the  and  Cowan  as  were  (.1945 ) a n d  red  these not  species Table  are  the  rate  2.  higher  For  this  annual  combined  data  of  study.  This  and  authors  are  Salal, important at  areas. the  and  Bay  heavy  Northwest  grasses  key con-  lichens 8 of  8  per  trailing periods  Bay,  Northwest'. Bay  was  foods  at  Cowan  (1945),  comparable  Brown  lists  i n Table.  rumen  i t is likely  list  spring,  in late  which  diet, than  they of  by  from  the  the  other  3,  areas.  Four  species  Bay. d i e t ' and  most  present  grasses  other  trailing  and  are  very  important  blackberry, arboreal  important  pearly everlasting  ten  in  the  a l d e r , and  also  samples  derived  presented  summer.  appear  the  ( 1 9 6 1 ) and  and  that.these,  f i r , red  cedar,  of  Douglas f i r ,  blackberry  during  studies  in early  a tentative  willow), are  Only  use. o f  reason  (.red  11  (7  cent,  cent,  in their  annual  Douglas  the  balance  i n the  given  sedges  4 6 per  per  The  (1961),  found  Northwest  i n a l l three  Northwest  lichens  at  and  cent),  vegetation.  deer,  two  11  cent.  Brown  alder,.and  obtained  important  two  mixed  per  c o n t r i b u t e d hy  plants  4 per  C8  grasses  t r e e s made up  b l a c k b e r r y , w i l l o w and  salal,  Since  volume  herbaceous  coastal'blacktailed  of  c e n t ) , and  total  mushrooms  grouped  trailing  per  deciduous  cent,  and  arboreal lichens.  i n one  cataear  of are  it" i s possible that  the  other  unique these  to  would  29  not  have ranked  available  i n the f i r s t  on l a t e  example,.ranked (Table  summer  food  habits.  i t i s possible  nual  diet  than  that they  t o deer  either, of these  Salal Red cedar Trail.blackberry Arboreal lichens G r a s s e s and sedges Douglas f i r Pearly everlasting Red alder Willow Catsear  by l a t e  sum-  forbs.  Southern Vancouver, Island '  Bay  index  a r e more i m p o r t a n t ' i n t h e a n -  The. t e n d o m i n a n t i t e m s , i n . t h e a n n u a l Columbian b l a c k t a i l e d deer.occupying separate ranges.  Northwest  were  Mushrooms, f o r  2 ) , . a n d s i n c e some a r e a v a i l a b l e  3  data  e i g h t h on t h e b a s i s o f t h e c o n s u m p t i o n  mer,,  Table  ten i fsufficient  Douglas f i r Salal Arboreal lichen Red a l d e r ; Willow. . Mushrooms. Bracken G r a s s e s and sedges Thimbleberry Equisetum ,  diets of three  Western ^ . Wash i n g t o n Trail.blackberry Salal Grasses Red alder Vine maple Western hemlock Douglas f i r Huckleberry Fireweed Red cedar  From p r e s e n t s t u d y w i t h i n t e r p o l a t i o n (1945) and Brown (1961).. See t e x t .  from  Cowan  2 From  Cowan  ( 1 9 4 5 ) . ..  From  Brown  (1961)  Western ington  deer,  Northwest than  hemlock,  was  Bay.  one p e r c e n t  never Dz.ubin  an. i m p o r t a n t  observed  food  to western  t o h a v e b e e n b r o w s e d at..  .(.19.51). r e p o r t e d t h a t  of the diet  Wash-  during winter  i t formed feeding  less  tests  30  with .blacktailed unpalatable m a p l e was was  t o deer  was  Equisetum, Bay  a n d Cowan  on s o u t h e r n  not present  of limited  latter  deer,  distribution.  rated important found  r a t e d i t as apparently-  Vancouver  In the study  area  Where  s e v e r e l y hedged.  b u t were  C194 5)  Island.  and r e d h u c k l e b e r r y  available,  Bracken,  however, the  t h i m b l e b e r r y , and  by Cowan, were  i n only  Vine *  s m a l l amounts  eaten  at  Northwest  i n the stomach  samples.analysed.  Spring-summer most  important  summer, leaves found  that this  c e n t , were  Hulcus  Catsear, diet  The  transition  berry,  i n May  samples  Douglas  early t h e new  Cowan  and sedges,  important  item.  formed half  g r a z i n g was  Usnea  by  deer  a t 15  recorded multiflora.  15 p e r c e n t  the  (1945 )  Among t h e  e x e r a t a , and L u z u l a  of the  samples.  barbata  but occurred  period progressed.  bearberry,  sampled.  upon w h i c h  than  arboreal lichens  were abundant the  most  r a t e d t h i r d , by v o l u m e , i n more  (Fig. 4).  Grasses  lanatus, Agrostis  and o c c u r r e d  and  apparently not eaten  Island.  i n the area  spring  was t h e  t h e p r e f e r r e d p o r t i o n s and were  o f the stomachs  the second  present  of the diet  s p e c i e s was  Vancouver  Pearly everlasting  during the-late  4.6 p e r c e n t  i n 79 per., c e n t  species were  item  d i e t -  and f l o w e r buds, were  Southern  per  food  forming  reported in  transitional  and A l e c t o r i a less  Salal,  f i r and r e d cedar  sp.  f r e q u e n t l y as.  trailing, blackeach  made up  A U T U M N  i  i g  b e ai s e r i a l  diets  of Northwest  Bav  deer  ' 3 2  less ant  than  because  cent was  each  taken  and v a n i l l a  the  from  latter  but  moisture  leave •in  "  ries  diet  and low f i b r e as o t h e r after  Thus,  a shift  i n pref-  as soon as  rumen  samples browsed. may  unidentifiable  (Bruggemann,  Giesecke,  it  1968).  an a v e r a g e  had dropped  autumn  season  year-round  and r i p e  food  i n Western  diet.  For the three-'  therefore, salal contributed  by volume  this  ber-  November, t h e l e a v e s  of the diet.  b u t made up o n l y  (1961) found  salal  o f t h e autumn  (Fig.4).  b l a c k b e r r y l e a v e s ;-were p r e s e n t  of t h e samples Brown  green  i n early  as a whole,  o f 56 p e r c e n t  Trailing  both  o f 79 p e r c e n t  month  and  quantities  o f t h e new g r o w t h  consumption  c o n t r i b u t e d 16 p e r c e n t  bulk.  i n increasing  succulent plants,  alone  cent  everlasting,  t o be e x t e n s i v e l y  content  the berries  average  species  while pearly  i n the spring  noted  After  an  of these  i n the spring.  -- When a v a i l a b l e ,  formed  each  import-  t o 71 p e r  to succulent- forbs i s indicated  s p r o u t s were  soon  50 p e r c e n t  period progressed.  Walser-Karst,  Autumn  taken  was n o t r e c o r d e d  i t , as w e l l  t h e rumen  and  were  become'available  the tender  High  quantities,  leaf  browse  Bracken  i n from  As w i t h l i c h e n s ,  i n decreasing  the transition  erence  o f t h e diet'. b u t a r e c o n s i d e r e d  was p r e s e n t  of the samples.  catsear as  s i xp e r cent  i n 91 p e r  7.3 p e r c e n t  t h e most  important  Washington,  of the autumn  Chatelain  (1947)  33  found  i t o f moderate  (1945) found on  •  southern  .  '  cent), were  i tof l i t t l e Vancouver  Red a l d e r Douglas  other  autumn  importance  importance  (.7.2- p e r c e n t ) ,  f i r (3.9 per cent) items  f i r occurring  achs.  of browse  the  March  diet  and g r a s s e s  ( 7 . 0 per  (2.4 p e r cent)  i n more  than  half  musht h e stom-  s p e c i e s a n d a f e w f o r b s made up  The c o n s u m p t i o n  was a b o u t  was s t i l l  food  half  eaten  (Fig. 4).  volume eal  mushrooms  balance.  Winter  ies  of the year  d u r i n g t h e autumn, w i t h  rooms and Douglas A variety  a t any time  a n d Cowan  Island.  leaves  important  i n Oregon,  logging  by almost  Red c e d a r ,  volume. Douglas  t o mature  Grasses  a m o u n t s by a t l e a s t  Range  were  eaten  Composition Cover  t h e key-  second  by  Arbor-  a n d b y snow a n d 13 p e r c e n t b y  c o n t r i b u t e d • 8 p e r Cent.  70 p e r c e n t  i n minor  but the spec-  a l l samples.  contributed  b l a c k b e r r y were  c a t s e a r and mushrooms  through  a n d was s t i l l  i n almost  trees,  amounts b u t by o n l y one t h i r d plants  a l l deer  by s t r o n g w i n d s  and sedges  f i r and t r a i l i n g  December  a t 21 p e r c e n t , r a t e d  made a v a i l a b l e  damage  from  o f t h e autumn p e r i o d  a n d i t t o o was p r e s e n t  lichens,  berry,  that  of s a l a l  taken  i n moderate  of the animals, while  were  taken  i n slightly  of the animals.  amounts "during t h i s  and Changes with. .Seral  a n d s p e c i e s compos i t i o n of  Many  bear-  greater other  season.  Succession various serai  com-  munities  were  derived  lished  and  surveyed  cation  and ' r e l a t i v e  t r a n s e c t .-lines ; The  logged  senting ing  during size  Table  patches  eleven  from  acres  forest. (Plot  each  lists  The  V)  to  sizes  69.0  plot,  acres  17  estab-  5 shows  the  lo  the ' d i r e c t i o n  of  burned  unburned  stands  of  and  plots  descriptive features  eight  two  study  Figure  included  age-classes, and  permanent  195.9-61.  of  studied  six age-classes,  climax  4  27  the  of  sites  sites  and  each  repre-  represent-  180-200 y e a r  p l o t s ranged  ( P l o t : 2)  of  old  from  sub  4.1  averaged  14.5  acres.  .Average and  the  rank  of  successional Appendix  D  individual Division  of  cover-composition dominant  age  are  p l o t s are the  ities  developing  Cowan  (1) (2) (3) (4)  those  (1945).  on  British  communities  to  given  at  described Four  in sites  file  with  Columbia  logging  the  of  (burned  areas)  f o r the  - The  major  or  burning  are  and  27  Research  Wildlife  Bay  Branch.  serai  commun  basically  Vancouver  a s s o c i a t i o n s were  Senecio-Epilobium association. Gaultheria-Kypochaeris association. Gaultheria-Pseudotsuga association. Pseudotsuga consocies. •  categories  increasing  Wildlife  F i s h and  f o r southern  distinct  C  plant  D e t a i l e d data  Northwest  after  major  i n Appendix  (unburned. a r e a s ) .  Serai  ilar  species  by  Island  sim by  recognized:  35  Fig.  5.  The N o r t h w e s t major l o g g i n g relative size p 1ots.  Bay S t u d y r o a d s and o f t h e 27  Area showing the the l o c a t i o n and permanent study  Table _  Plot A. •A B C R S E G H U P J N 0 L M W 1 B. D T Q I V K X 2 C. F 3  4  The p e r m a n e n t r a n g e s u r v e y a n d p e l l e t - g r o u p • s t u d y p l o t s e s t a b l i s h e d .at . N o r t h w e s t B a y , 1.9 5 9 t o 1 9 6 1 .  Age i n Y e a r s a t I n i t i a l j Resurvey Survey  Year Logged or Burned Burned-over 1958 1957 1957 ' 195 8 1958 1956 1954 1953 1956 1952 1949 1950 1950 1948 1948 1948 . 3.9 4 5 Unburned  sites  A* A A  1 2 2 3 3 4 5 6 6 8 10 10 10 12 12 13 15  A S A A S A A A A A A A A  9 .7 •13.1 13 . 6 •9 . 5 11.5 21. 7 8. 9 8. 9 11. 4 17 . 5 8.8 23 . 2 17 . 4 23 . 3 9. 8 11. 4 9.3  — —  •  —  4 —  -  -  9  —  —  '  —  14 —  -  sites  1957 1956 1954 1951 1953 19 4 7 1948 1947 Subclimax  Area (Acres  2 5 6 8 8 12 13 13  -  9.4 8. 8 16 . 2 . 8.9 4.1 9 .7 13 . 2 69.0  -.  8.3 15 . 0  —  -  forest sites  —  -  18 0-200 180-200  Average  A -Autumn  burn  S — Spring  burn  14.5  .  Cowan's " r o c k communities not  bluff",  were  considered  The they the  limited  of  the  sites, by  i n the  sample  size  and  to  site  tially  a result  of  differences  three  the  than  after  from  They by  per  fifth  and. t h e n of the  the  develop  the  the  are the  year Under  cent  of  and  third  year,  decline.  downward  shading  to  being  par-  is  The  apparent.  c o v e r - d e n s i t i e s about early  are  Gaultheria-  decline gradually  of minor  importance  Gaultheria-Pseudotsuga  the  forest,  year  mature  forbs cover  less  surface.  deciduous, t r e e s dominate through  cover  60  maintain the  consists  trend  attributable  (the early  young  By  in  intensity.  i n the  and  to  shown  Thereafter they  the  rapidly,  community  peak  are  latter  i n burning  burning, usually  association.  Shrubs tum  variability,  c o m p e t i t i o n and  one  graph  plants reach  association).  were  as  relationship  plant types  Herbaceous  fifteenth  that they  associations  the  s u c c e s s i o n , however,  Hypochaeris  by  and  p a t t e r n of  years  through  distribution  f o u r dominant  surface covered  Irregularities  general  "alder-willow"  detail.  i n burned-over  amount o f 6.  in  such  sequence  develop  Fig.  of  " s e d g e meadow", and  37  to to  at 65  least per  that level fifteenth  of  shrubs,  c o n t i n u e s beyond  cent  until  year and the  the  low  stra-  fifteenth  year.  of the  only  30  the  the  surface  tenth  year,  per  cent  i t is likely  that  fifteenth  year  as  S e q u e n t i a l d e v e l o p m e n t of. f o u r m a j o r s e r a i a s s o c i a t i o n s and r e l a t e d c o v e r p e r c e n t a g e s by t h r e e v e g e t a t i o n t y p e s ; slash-burned sites only. -  Senecio-  Gaultheria - Hypochaeris Association Gaultheria-Early-Intermediate— -AdvancedEpilobium Pseudotsuqa(Cirsium-Epilobium (Epiiobium-Rubu's (PseudotsutjC-Rubus Assoc. Subdom.) Subdom.) Subdom.) Assoc. 80-  Pseudotsuaa Subclimax {Gaultheria Understory)  shrubs the  a r e shaded  mature  ance  and r e p l a c e d , by c o n i f e r o u s  timber,  and r e a c h  shrubs  a peak  (.principally  cover  C o n i f e r s .comprise of  associations. ually,  remaining  13. y e a r s .  five  cover  per cent  Pseudotsuga  and i t i s a t t h i s  advances  into  dense  70 p e r cent'. proportions .  increases  that  This  years,  type,  second-growth  Douglas  v e g e t a t i o n , and e v e n t u a l l y i n t o  canopy  forest  a salal  The the  four  o f each  ing,  a sparse  the  first  sel  (Senecio  tuce  are presented  Senecio-Epilobium Immediately  (Lactuca  arvense)  after  CAppendix  a closed-  and t h e c h r o n o l o g y  follow.  of  Representative  i n the Appendix.  logging forb  and i n t e n s i v e s l a s h - b u r n a s s o c i a t i o n develops.  C) i t i s d o m i n a t e d  sylvaticus),  with the p r i n c i p a l  muralis), thistle  and w i l l o w h e r h  f i r with  association  and ephemeral  year  stages  a marked,  understory.  characteristics  fire-induced serai  photographs (1)  structural  12 t o  i n turn,  subordinate  or. m o s s  grad-  the G a u l t h e r i a -  little  with  very  f o r the i n i t i a l  time  a s s o c i a t i o n develops.  domin-  Gaultheria-Hypochaeris'  I n t h e f o u r t e e n t h and f i f t e e n t h  increase, occurs  quickly  below  ground  Under  regain  insignificant  and e a r l y  Coniferous  salal)  d e n s i t y of almost  almost  the Senecio—Epilobium  trees.  CCirslum  b y wood  . In  ground-  a s s o c i a t e s ,l e t  l a n c e o l a t u s - andC_.  CEp i l o b iurn m l n u t u m )'.  Shrubs  and  coniferous than  four  trees  p e r cent  In a  are rare  and t o t a l  t h e second  western  year,  hemlock  fireweed,  become  well  markedly  and  contributing equally.  relatively scorched  ( 2)  1959),  salal  C ) , a mixed  Rubus  species,  berry  and salmonberry a n d become  Associated and Most  moisture  the fourth  shrub  than  and f o r b  maintains  i t grows b e s t soil  a  on  conditions  more  to fourteenth community  rarely  ' Both  of these  blackberry,  and Oregon  herbaceous  with  grape  associates.  black  plants  and w i t h  advancing  are other  vary  rasp-  (Cirsium  rep-  years.  common  shrubs  greatly with  the intensity  by  species  spectabills ) are usually  abundant  are thistles  year  dominated  any o f t h e p r i n c i p a l  (Rubus  conditions  prominent  shrubs  association  including trailing  twinflower  cover  with  Wood g r o u n d s e l  i s recognizable.  f a r more a b u n d a n t  Willows,  Plant  20 p e r c e n t  but since  raspberry  season.  approximately  and c a t s e a r  resented,  black  i t and t h e a s s o c i a t i o n i t s e l f  Gaultherla-Hypochaeris  (Appendix  are  density,  i n the third  From  c a t s e a r and  and i s i n t o l e r a n t o f changing  (Mueller-Dombois thrive  thistles,  established.  to approximately  constant  soils  may Be l e s s  including salal,  increases forbs  cover  (Fig.6).  v a r i e t y o f woody s p e c i e s  and  plant  soil  of burn.  s p p . and Carduus.  spp.),  lettuce,  pearly everlasting.,  Wood g r o u n d s e l first ly  year  is.g r e a t l y  sun-exposed  Bracken  pockets  ment  of the t y p i c a l  sist  under  the  The  plants.  less  than  species,  forb stratum.  stages  and s h r u b s  with  The  than  association  from  present cent  (see F i g . 6). i s shaded  of the surface. serai  c a t s e a r and t r a i l i n g  i n moderate  more  advanced  stages  burns,  important  each  occupy  . Grasses  are  stage  but s t i l l  Salal  i s the dominant  b l a c k b e r r y the  occupy  important and  let-  are recognizable i n f i v e -  areas  more  i n which Shrubs  total  dominate  of the surface than  plant and  cover  may  herbaceous  coniferous species are generally  but f r e q u e n t l y occupy of the surface.  by  quantities.  40 t o 70 p e r c e n t .  The  per-  are repre-  are e q u a l l y abundant,  of the area.  s i x to eight times  plants.  of the surface  i n any o t h e r  thirteen-year-old  ranges  develop  h o w e v e r , may  and b l a c k r a s p b e r r y , w i l l o w s , t h i s t l e  present  local-  vigorously i n moist,  Salal,  of this  of. 15 p e r ' c e n t  two p e r c e n t  associates,  per  are only  bracken.  Forbs  abundant  cover  p e r c e n t * from, i t s  and sedges  develops  leaf.  7) and f r e q u e n t l y p r e v e n t s  one-third or less  more  to  fern  CFig.  earliest  an a v e r a g e  tuce  i n cover  by t h e t h r e e - and f o u r - y e a r - o l d s i t e s  Typically,  ing  reduced  abundance , and g r a s s e s  distributed.  sented  f i r e w e e d , and v a n i l l a  an a v e r a g e  Only  under  of less  ideal  soil  than  five  and  moist-  42  Fig.  7  Dense b r a c k e n f e r n g r o w i n g i n t h e m o i s t l o w e r end o f P l o t J , 14 y e a r s a f t e r s l a s h - b u r n i n g . In t h i s s i t e s a l a l p e r s i s t e d under the bracken.  Fig.  8  An e a r l y G a u l t h e r i a - P s e u d o t s u g a association f i f t e e n y e a r s a f t e r l o g g i n g and b u r n i n g ( P l o t M). D o u g l a s f i r r e g e n e r a t i o n was p a r t i c u l a r l y r a p i d i n t h i s s i t e , e v e n t h o u g h r e f o r e s t a t i o n was not attempted. When s u r v e y e d t h r e e y e a r s e a r l i e r t h i s p l o t was c l a s s i f i e d a s a n a d v a n c e d GaultheriaHypochaerls association.  ure  c o n d i t i o n s i s Douglas  usually the  most  abundant,  surface.  Catsear  face  but  ranks  intensive. Douglas tuce 1  (3)  occupying covers or  Trailing  are  the  salal  common i n t h e  hamper and  the  once  other  forbs  and  year-old this the  advanced  earlier  become  less  serai  C)  the  the  fire  along  f i r cover.  earliest start  of  surwas  with  burning,  let-  does  numbers  The  salal. not  Dense seriously  they  soon  f o u r t e e n - and the  earliest  i t is little  salal  Doug-  (Mueller-Dombois,  and  1959) surpass  fifteenstage  different  total  the shrub  of from  discover,  Trailing.blackberry,  c a t s e a r remain  with  of  approximately  after  stages  represent  reduced  w i l l o w and  the  cent  components.  conifers  Structurally  abundant  Since before  of  (Appendix  i n c r e a s e d Douglas  years  cent),  associate with  in sufficient  features being  grape,  of  where  Gaultheria-Hypochaeris association,  Oregon  per  is  association  i n abundance.  association.  tinctive and  sites  38  cent  averaging  fourteen years  establishment  shrubs  four per  principal  principal  established  of  i n a l l areas  (each  Gaultheria-Pseudotsuga  f i r becomes the  average  Salal  hemlock, willow,, black r a s p b e r r y ,  Approximately las  only  third  pearly everlasting  cent)  an  ( F i g . 8).  b l a c k b e r r y (2 p e r  f i r , - western  and  per  second  f i r dominant  as  subdominants  but  time.  logging occurred field  only  investigations,  fourteen more  advanced  stages  survey. that to  However, under  Douglas  2.0 y e a r s  (4)  Pseudotsuga.  tsuga  in  an  (Cowan  of Douglas  the understory  were  as t h e dominant  forest  1945),  f i r and w e s t e r n to only  community,  was r e p r e s e n t e d  timber.  i n both  restricts  t h e most  and s u p p o r t e d  tion few  were p r e s e n t  only  low-hanging  remainder plant  hemlock branches  a moss  i n this  shallow  was n o t e d  soils  Twin-  i s very  but, together  Limited  red.cedar  As n o t e d  association  (1959).  i n the s a l a l  of western  community  gymnocarpa) and  of the surface.  of the low s t r a t u m .  variety  (Rosa  on t h e s e  two p e r c e n t  of western  rose  growth  Rock-outcropping  Oregon  dwarf  consisting  studied, covering  floWer,  grape,  stands  shade-tolerant species.  of the surface.  sites,  by two  hemlock  t o t h a t d e s c r i b e d by M u e l l e r - D o m b o i s  covered  18  or "Pseudo-  canopy  similar  pipsissewa  species  A dense  s a l a l . d o m i n a t e d t h e two s i t e s  o f 65 p e r c e n t  present  c o n d i t i o n s i t i s presumed  salal  subclimax  180- t o 200-year—old  average  not a v a i l a b l e f o r  burning.  mature,  Tall-growing  were  subclimax  consocies"  mostly  average  f i r surpasses after  The  of  of t h i s . a s s o c i a t i o n  zones  regenera and a  made up t h e  b y Cowan limited.  (1945),  45  Speci.es  Variety  The  In  Serai  variety  Communities  of p a l a t a b l e species  of. r e g e n e r a t i n g r a n g e  i s probably  production  That  at  of  food.  a l l seasons  rumen  is clearly  analyses  reported  presented  by , o t h e r  1952 , B r o w n  Northwest  variety,  Bay,  as  i s shown  Only  species recorded  during  the  cantly  to  1945),  are  sis.  Since  the  (see  grasses are  not  variety  total  by  foods  results  food  habit  of  studies  C h a t e l a i n 194.7 ,  this  time  of  herbaceous  the  Thus,  Rieck  d e r i v e d are  given  i n g r e a t e r than  sedges  were  occurred s i x to of  i n the  C.  quantities signifi-  a b o v e , and  foods  Cowan  in this  identified  analy-  to  at  species  this  intermediate Gaultheria-  eight years  time  COdum, 1960 ) o c c u r r e d , the  data  i n Appendix  contribute  not  The  trace  habits results  p l a n t s important- to  important,  to  succession  included.  number  i t was  serai  i n F i g . 9.  " p a l a t a b l e " deer  and  to  graphically  food  association  At  indeed  and  the  i t is related  s u r v e y , 'and. k n o w n  diet  they  index"  were  i n c l u d e d as  Hypochaeris  ity  curves  range  Peak  equal.  by  CCowan 1 9 4 5 ,  the  species,  the  unit  c o n s u m e many- d i f f e r e n t  earlier,  from.which those  i m p o r t a n t , .as  demonstrated  workers  i n any  1961).  .Species at  deer  as  present  after  of  shrubs  deer  were  that and,  the  and  burning. the  number  approximately greatest  i f variety  greatest attractiveness  to  of deer  "diversfoo'ds  is  could '  Fig.  9.  Trends i n the v a r i e t y present after logging  SenecioEpiiobium Assoc.  o f p a l a t a b l e deer foods and s l a s h - b u r n i n g .  Gaultheria - Hvpochaeris -Early-  Association  -Intermediate—  Gaultheria-  -Advanced-  (Cirsium-Epiiobium  (E p i l o b i u m - R u b u s  (Pseudotsugq-Rubus  Subdom.)  Subdom.)  Subdom.)  Pseudotsuga  Pseu-dotsuqa Assoc.  Subclimax (Gaultheria Understory)  £.0  2 0 ail  species  'o 111  o  forbs  and  ferns-^  &  I  10  a.  conifers  8 years  1 iO sines  12 burning  14  16  180-200  47  be  expected.  Eleven conifers  were  herhaceous t h e ' maximum  plants., eleven  but each form  reached  time.  Forbs  quickly,  in the  the f i r s t  intermediate  years  after  occurred, forbs had  season.  decreased  Under  t o two.  browse  reaching  encroachment  The  three  r e d cedar,  present  i n regenerating  within  reach  until  (Fig. 9), of deer  only  variety  four  timber  development  palatable this  number  there  of s a l a l  rate  than  forbs,  of the late  Gaultheria-Pseudotsuga  variety  and balsam  diversity i n  a s s o c i a t i o n (6 t o 8  at a slower  was i n d i c a t i o n probably  owing  of a  sig-  to the  and Douglas f i r .  palatable coniferous  western  teenth, year  invaded  Subsequently,  different  •«  and e a r l y  d e c l i n e i n browse  continued  season  t h e mature  variety  Gaultheria-Hypochaeris  at a  decline i n forb  •  plants  maximum  associations.  variety  F o l l o w i n g maximum  a gradual  recorded.  three  w i t h , many b e c o m i n g e s t a b l i s h e d  and i n t h e f i f t e e n t h  were  nificant  maximum  Gaultheria-Hypochaeris  burning)  The not  growing  and  number's o f p a l a t a b l e s p e c i e s r e -  corded,  invaded  shrubs,  fir,  Douglas f i r ,  -were n o t c o n s i s t e n t l y  areas, u n t i l Red c e d a r  trees,  the fourteenth to  was t h e o n l y  under, t h e m a t u r e  conifer  fiffound  subclimax . forest .  • Range  Productivity  The varies fire  production  greatly  history  1962).  as  biotic the  (Halls  area,  and  plants  of  were  latter  plants  to  the  (Table  cannot  not  had  samples  gram  climax  amount of  follow  this  subclimax  of  the  conditions.  p a l a t a b l e new surface  trend  the  a relatively  (Table  and  browse  amount  of  p a l a t a b l e browse  expected the  for total  was  to  follow  available  annual  the  with level  In the  study  The  clipping  avail-  palatable former  investigation,  for  given  climax  low  by  5).  non-palatable  i n Appendix  in  same t r e n d  be-  foods,  and  production.  of- t h e  consumed  by  !  amount o f  deer  at  the  browse  time  Therefore,  the  crop  and  (near  the  end  During  the  clipping  relatively  and.in little  August  December. food  was  noted  to  estimates  the  collected.  season),  C.  December  evergreen  obtained  in July  and  Murphy  growth  covered  cover-percentages  a l r e a d y been  were  decreases.to  percent  measure  standing  growing  the  be  use,  increases for a period varying  include only  account  No  the  the  land  E h r e n r e i c h and  d e s t r u c t i o n of  then  total  of  they  1960,  communities  past  subtracting cover-percentages  Estimates 6)  structure,  Crawford  d e r i v e d from  by  from  of  foods, i n f o r e s t  serai  approaches  the  appear  figures  which  and  community  able,  do  and  production  estimates  cause  deer  with, s i t e ,  area,  the  of  Generally, after  s i t e s , , food the  48  have  represent of  already  the pro-  been  49  Table  5 A n n u a l p r o d u c t i o n of. d e e r f o o d i n v a r i o u s p o s t - f i r e s e r a i s t a g e s , e x p r e s s e d as pounds per a c r e wetw e i g h , t'. The .numbers i n p a r e . n t h . e s e s e x p r e s s p a l a t able food production as a percentage, of t o t a l newgrowth p r o d u c t i o n . Years since burning  F o r b s and Ferns Shrubs Conifers  grasses  Total  Figures  12  14  151 15 782 T  12 5 69 8 47 ; 249  13 140 7 91 28  97 16 6 7 44 105  61. 8  e x t r a p o l a t e d from  47 . 0  range  Mature Timber T 2 42 3 5  1114 (79.5)  430 (95.5)  43.4  6 7.1  survey  data.  6 Winter a v a i l a b i l i t y of deer foods i n v a r i o u s p o s t f i r e s e r a i s t a g e s , e x p r e s s e d a s p o u n d s per. a c r e wet-weight. The n u m b e r s i n p a r e n t h e s e s e x p r e s s f o o d a v a i l a b i l i t y d u r i n g w i n t e r as a p e r c e n t a g e of t o t a l a v a i l a b l e new-growth.  F o r b s and Ferns Shrubs Conifers  the  10  3-9 . 3  Years since, b u r n i n g  Total  4  94 8 1290 ' 972 (93.5) (93.0) (63.9)  Percent of ground c o v e r e d by p a l a t able species  Table  ,  grasses  4  53  14  T T 1.0 45  T 6 16 4 59  T 10 299 34  55 (82.4)  229 (87,7)  D e r i v e d f r o m summer e s t i m a t e s b y weights of evergreen p l a n t s .  343 (97.4)  including  Mature Timber T 2 421 5 428 (98 .  only  50  consumed, crop  so i t would  closely  approximate  Data total  food  followed  production  by a s l i g h t  ( F i g . 10A).  ually  through  forbs  and g r a s s e s  years  after  acre  through  were  forest. dance  from  forbs  and g r a s s e s ,  at which  years  time  trend,  annual  production after  browse  progressive  i n a l l age-classes,  available  d e c l i n e through  on e a c h  to decline  i n the  subclimax  when  approxi-  As w i t h t h e under  formed  the mature  the bulk  a n d was m o s t  Approximately  four  i n c r e a s i n g i n abun-  available.  growth  group,  151 pounds  tended  fern production  Shrub  burning.  were  were  fire,  grad-  quantities  quantities  an o p p o s i t e  a  four  As a  approximately  abundance  to trace  however,  was n e g l i g i b l e .  a  stage.  i n greatest  Their  per acre  timber  ferred  forest  after  continues  the f o u r t h to the fourteenth year,  166 p o u n d s  years  increase i n  the next  decline probably  were p r o d u c e d  showed  mately  ten  a rapid  decline f o r at least  available.  Ferns  crop.  indicate  to the subclimax  subsequent  of. t h e s t a n d i n g  f o r the f i r s t ' ten years  This  burning,  estimates  the t o t a l  f o r t h e summer  years  per  appear, t h a t  abundant  847 p o u n d s  acre  at this  to the subclimax  of the  of pre-  time, but  stage  was  indicated.  Estimates foods  of the annual  do n o t c o m p a r e  estimates  shown  production  favorably with  i n F i g , 6.  They  the  of coniferous  cover-percentage  do n o t f o l l o w t h e  expected  Fig.  10.  The w e i g h t o f d e e r f o o d a v a i l a b l e w i n t e r i n each of t h e major s e r a i  SenecioEpilobium Assoc. a.  GjluJlhexifl- Hvpochaeris -Early-  m summer a n d stages.  Association  -Intermediate—  -Advanced-  (Cirsium-Epilobium  (Epilobium-Rubus  (Pseudotsugc-Rubus  Subdom.)  Subdom.)  Subdom.)  Gaultheria-  Pseudotsuqa  PseudotsuaaAssoc  Subclimax (Gaultheria U n d e r story)  summer  200-  all  plants  900  600-  3 0 0 -  forbs-p  b.  conirers  P. -  winter  600r  4  8 years  10 since  12 burning  14  !6  180-200  • trend  of a very  followed is  by  likely  ation  in different I n two  annual  conifers  association  ter  foods  serai  after  burning,  nine-  and  i n F i g . 10B,  and most  was  and  deciduous  on e a c h  estimated  cession  sites  Under by  growth  were was  from  i s apparent an  Douglas  the weight  from  Table  was  accounted  f o r by t h e g r a d u a l  p a l a t a b l e evergreens  such  by  the  availabil-  herbaceous  were  growth available  salal.  6 that  deer  replacement as s a l a l  343  shrubs  of a l l annual  was  succes-  years  and  where of  win-  In the  with  428 p o u n d s  important  four  per acre  i n c r e a s i n g percentage  i n December  serai  f i r .  available,  of which  agree.  of p r e f e r r e d  available  pounds  were  do  coverage  s u b t r a c t i n g the weights  97 p e r c e n t  advanced,  regener-  subclimax.  the mature t i m b e r ,  available  by  229  This  Mueller-Dombois  constantly with  i n summer, a p p r o x i m a t e l y  acre,  It  of t h i s  respectively  dominant, s o u r c e .  estimated  increased  years  Gaultheria-Hypochaeris  availability  55 p o u n d s p e r a c r e  pounds p e r acre  by  surface  the Pseudotsuga  14-year-old  ity  as r e p o r t e d  i n the early  i n December  Only  types  14- y e a r s .  r a t e s of c o n i f e r  and. n e g l i g i b l e  and under  sion.  i n c r e a s e .at a b o u t  10 t o 12  stages, however , the r e s u l t s  recorded  shown  f o r the f i r s t  to variable  soil  production  were  As  increase  a more r a p i d  attributable  (1959 ). Low  gradual  52  as s e c o n d a r y o f new food.  growth This  of deciduous  and D o u g l a s  suc-  c a n be plants  f i r , and  53  points to  out the importance  wintering  theria—Hypochaeris of weight  cover.' of  from  subclimax  p a l a t a b l e foods  ly  the greatest producer with  rates  relatively  only  the s a l a l  early  moderate  in  winter  is  available  available  The ered  that  as w i n t e r  during  less  than  food.  i s potentialB u t com-  stages, i t  of annual  growth.  association  produces'  s u b t r a c t i o n of the  s i x per cent  Therefore,  t h e s p r i n g and  eaten  of the t o t a l  f o r a g i n g by d e e r i n t o f o r b s and  grasses  summer,  association  (.1945) t o b e t h e m o s t i m p o r t a n t when  cover  p l a n t s and by p l a n t s n o t  m u s t be c o n f i n e d m a i n l y  community, p a r t i c u l a r l y  winter..  high  foods.  serai  Moreover,  Gaultheria-Pseudotsuga  b y Cowan  winter  Gaultheria-Hypochaeris  by h e r b a c e o u s  i t capable  autumn and e a r l y  production  of food.  makes  a n d w i n t e r , arid  of evergreen  Gaul-.  of palatable  has a r e l a t i v e l y  summer  low i n t o t a l  indicates  association  i n terms  and c a t s e a r - d o m i n a t e d  amounts  amounts produced  this  i n both  to the  most p r o d u c t i v e i n  composition  spring to late  of  The  stages  the intermediate  is clearly  i t s varied  Pseu'dotsuga  pared  earlier,  association  Furthermore,  mature  5 and 6 a r e r e l a t e d  o f . f o o d , and r a t e s h i g h  supplying food  The  i n Tables  associations described  terms  serai  deer.  . When t h e d a t a serai  o f t h e more a d v a n c e d  supplemented  was  consid-  food-producing  by f r i n g e s  of sub-  climax  forest.  analyses tion and  and  spring Along  and  of  forage  Composition  purposes  (1958) suggests influencing  range  of  effect  indicate  s e l e c t e d groups  summarizes  the  ance  designed  tests  regeneration  Einarsen  by  from  on  the  However,  results, of  nutrient  test  the  hypothesis  forage  quality.  succession species  deer.  be  important  average food  fourth  the  changes  the  nutrient species  did  fourteenth and  differ-  significant.  forage  nutrients  four  analysis  i n f l u e n c e of For  et  D i e t z et a l .  may  to  that  (Cowan  Generally,  the  con-  differences in  important  a s e r i e s " of  show t h e  snow.  the  factors  generally  to  the  value  on  seasonal  s u c c e s s i o n on  during  are p r e f e r r e d .  ( 1 9 4 6 ) and  that  of  autumn  Species  to  these  associa-  limited  deep  s e a s o n a l , and  selection  below  serai  by  w i t h advancing  s p e c i e s were  of  Forage  s p e c i e s were  each'of  content  summer,  forbs  i t i s of  o b t a i n i n g data  significantly  between  stages  of  of- s u c c e s s i o n .  ences  The  that  rumen  than 'ideal  Deer  W o r k by  presented  composition  year  i s less  of  to measure  levels.  change  the  indicate, that this  is restricted  decreases  1950 ) , and  not  of  conditions for late  serai  availability  quality  results  early  v a r i o u s food  nutrient  in  optimum  f e e d i n g , but  other  The tent  results  summer w h e n ' v a r i o u s , s u c c u l e n t  with  Chemical  combined  r a n g e .survey  near  winter  when f o o d  al.  of. t h e  provides early  The  most  - Table of  7  vari-  advancing  s p e c i e s , means  55  Table  7  R e s u l t s o f a s e r i e s o f four, a n a l y s i s o f v a r i a n c e - t e s t s designed t o determine the e f f e c t of s e r a i s u c c e s s i o n on t h e n u t r i e n t c o n t e n t o f i m p o r t a n t deer forage s p e c i e s , The v a l u e s t e s t e d a r e g i v e n i n A p p e n d i x E , T a b l e s 1-4.  Test  1;  Source Between Between Error  . df Species Ages  Total 2:  Source  -Ash  6.18** 1.07  3.84* .26  7.. 5 8 * * .82  Crude .Fibre.  N.F.E.  73.66** .02  23.27** 1.73  F o u r - y e a r - o l d v s n i n e - y e a r - o l d communities; f o u r s p e c i e s sampled i n December. 1  df Species Ages  Total.  3 1 3  Values of F Crude Crude Protein Fat 14.72* .07  12.28* 2.72  Ash 10.74* .73  Crude Fibre  N.F.E,  75.60** .63  20.13* .02  7  Test  3:  Source Between Between Error  8 1 8  Values of F Crude Crude Protein Fat  17 Test  Between Between Error  F i v e - y e a r - o l d v s 1 4 - y e a r - o l d communities; nine s p e c i e s sampled i n March.  Nine-year-old ities; three December.  df species ages  Total  2 1 2 5  * p < — p <  0 ,05 , 0 ., 0 1  v s 1 4 — y e a r - o l d communspecies sampled i n  Values of F Crude Crude Protein Fat  Ash  Crude Fibre  2 3.13* .5.-2 5  2.05 .33  43.72* 6 6 3.67** .01 13 0 . 3 7 * *  16.40 .40  N.F.E.  56  Test  4:  F o u r — y e a r - o l d vs n i n e — y e a r - o l d -14~year-old communities; f.our species c o l l e c t e d i n June,  Source  df  Between Between Error  3 2 6  species ages  Total  derived  from  samples age  some  only  species  age-class  and  N . F .E  6 8.48"" .57  40.66-" .65  59 .8 5 .09  4.03 .07  volved  were a  i n the  i n each  Appendix  15.19".30  :  test  i n two  o r more  used  i n the  analysis.  single  sample  was  the  and  their  of  However,  obtained  interaction  a n a l y s i s of  sites  for a  equal for  given  f a c t o r could  variance.  The  nutrient values  not  species  are  given  inin  E.  In  only  attributable  to  (N.F.E.) l e v e l s  one  test  burning result  coniferous  were  age-of-site; i n Douglas  when t r e a t e d as  similar  collected  consequently  determined  after  Crude Fibre  < 0.05 < 0 . 01  successional  low,  Values of F Crude Crude Protein' ' Fat Ash.  11  p p  be  vs  than was  species  a  nine  significant  years by  in central  differences  December n i t r o g e n - f r e e  f i r , trailing  group,  reported  there  were after  higher  blackberry fourteen  extract and  years  b u r n i n g ..(.Test. 3 ) .  Cowan' :et a l . ( 1 9 5 Q ) f o r British  Columbia,  but  wil-  no  A three  57  physiological The in  levels  e x p l a n a t i o n c a n .he o f f e r e d f o r t h e i n c r e a s e .  of other  nutrients  December, March,  i n groups  a n d .June w e r e  of plants  not altered  collected  by s e r a i  suc-  cession.  In  g e n e r a l , these  C o w a n e t a l . Cop clines  cit.),  as f o r e s t s  be d i s c u s s e d  have  been r e c o r d e d been  years  analysed  effect  the  above  all  age-classes  for  each  Values in  later,  o f season results  however,  14  8).  early  winter  o f v a r i a n c e , and where  e x t r a c t e d from  The s i x s p e c i e s w e r e  tests  changes  data  than  - On - t h e  four  then  (December),  to  seasons.  a  f i r , western and w i l l o w .  '9 a n d 1 0 .  a  sampled winter  tested with  with  from  levels  provided  and l a t e  applicable,  i n Tables  obtained  species,  T a b l e ' 8, p o o l e d ,  Douglas  basis of  nutrient  browse  blackberry, red alder, a r e shown  might  i f samples  younger  seasonal  important  analysis  these  As  investigation  These  were  of  conditions.  for differences attributable  (July),  trailing  de-  significant  nutrients  (March),  salal,  climax  of  quality  i t was p o s s i b l e t o u s e v a l u e s  f o r s i x o f t h e most  test.  food  those  years.  on f o r a g e  (Table  of testing  that  r e g e n e r a t i n g areas  to d e r i v e average  species  summer  toward  i n the present from  do n o t s u p p o r t  concluded  o f age and o l d e r t h a n  The  means  who  regenerate  will  had  results  an  range red cedar, Results  T a b l e i3 Browse  The a v e r a g e s e a s o n a l compos i t i o n o f s p e c i e s a t K o r t h w e s t Bay , V a n c o u v e r  Species  some i m p o r t a n t Island •  Crude Protein  Douglas F i r J u l y Dec . Mar .  63.5 59.6 53.9  6.32 10 .17 7.63  16 . 43 12 .69 13 .76  2 .36 3 .97 2 . 60 .  20 . 20 20 . 31 18 . 96  Western Red C e d a r  July Dec . Mar .  61.0 58.5 51.6  4.68 7.38 4.75  18 . 04 18 .02 9 .55  •. 4.22 4 .48 3 . 40-  2 5 . 34 27 . 02 2 1 . 10  Salal  June July Aug . Dec . Mar .  78.4 71. 4 57.6 58.5 55.9  9.68 7.9 7 . 3.70 6 . 40 5.03  14 13 7 10 10  . 16 .63 . 20 .42 . 20  3 .83 . 4. 46 3 .85 5 .23 4 .66  18 . 2 6 2 2 . 93 26 . 05 24 . 48 19 . 60  July Aug. Dec . Mar.  66.3 61. 2 63.0 57.4  15 . 04 9.30 13.23 8.55  12 8 12 8  . 64 . 10 .70 .00  6 .22 5 . 60 6 . 57 5 . 40  July Aug . Dec . Mar .  56.5 59.4 46 . 5 53.5  17 . 47 13 .70 - 11.16 .7.90  13 10 14 17  .66 . 50 . 16 . 17  4 . 14 3 .30 . 3 . 00 2 .50  June July Aug . Dec . Mar .  69.1 62.9  "14.46 12 .35 5.40' 9 . 46 6 . 25  12' . 69 14 . 27 • 4 . 60 8 . 30 10 . 5 0  Trailing Blackberry  Red  Alder  Willow  —  5.C . 4 . 5 0.5  -  Percent of dry weight Ether Extract Ash Fibre  % Moisture  1  d e e r f-or a g e  6 . 30 6 . 46 5 .70 3 . 92 3 .40  -  13 . 13 15 . 40 13 . 92 1 1 . 40 .  N . F . E54.67 52.84 57.03 47 .71 43 . 09 '. 6 1 . 1 0  .  '  54.07 51.56 59.10 53.45 60.50 52.96 61.60 53.56 56.20  1 1 .4 1 16 . 50 23 . 47 2 1 . 17  53.32 55.90 47 . 47 51.23  15 . 78 16 . 26 22 . 90 29 . 54 24 . 0 0  50.77 50.63 61.40 48.7 6 56.10  Table  8  Browse  (cont)  Species  Moisture  Oregon grape  50 . 8 Aug. Dec . . 50 .2 M a r . • 50 . u  Black Raspberry  June July Aug . Dec . Mar .  75 . 9  -  53 . 0 61 . 5 41 . 6  Crude Protein 4 . 40 10 .12 7 . 53 18 15 8 6 6  .38 . 04 . 30 . 24 .90  Salmonberry  J u n e •' 6 9. 1 July 57 . 0 Aug . 51 . 1 Dec .  • 17 . 10 12 .35' 7 .00 7 . 37  Arbutus  Aug". M a r .••  56 . 0 55 . 1  4 .70 5 . 93  Rose  July Aug . Mar .  56 . 3 49 . 6 43 . 9  11 . 4 5 4 . 10 4 .13  Red Huckleberry  M a r . • 84 . 4  5 . 50  Thimbleberry  Aug .  65.6  1 1 .70  Percent of dry Ether Extract Ash  weight Fibre  N.F.E.  3.25 2.87 2.57  36.90 33.10 29.53  51.05 4 5.70 55.07  12 .17 12 .57 3.20 8.07 3.9 5  6.92 6.96 5.50 4.50 3.15-  13.79 .17.22 2 2.00 35.59 '3.1. 2 5  48 .74 48 . 21 51.00 45.59 54.75  12.50 13 .31 7 . 40 6 . 8,6  5.50 5 . 41 6.30 3.72  13.32 13.31 23.30 23.33  51.58 55.62 56.00 56.72  7.20 4.53  3 60 2 97  13.30 9.37  7 1 . 20 77.20  15 .15 6.20 4 . 37  6 32 6 20 6 10  18 .07 26.90 34.10  4 9.01 56.60 51.30  8.65  5.00  26.45  5 4 . 40  15 . 40  7.3 6  15.39  50.15  4.40 7 .61 5.30  Table  8  (Cont) Percent % Moisture  Herbaceous Catsear  of dry weight  Crude Protein  Ether Extract  Ash  Fibre  N . F'. E .  Species June July Aug .  84 . 4 8 0 ..5  15 . 50 13 .42 4 .10  16 .97 16 .83 5 . 30  8 . 67 8 .37 8 .20  14 .90 17 . 45 29 . 10  . 4 3 .95 4 4 . 93 53 . 30  Fireweed  June July  75 . 8 76 . 8  14 .85 13 . 17  13 . 42 14 . 18 '  6 . 89 5 .74  8 .77 8 . 81  56 . 12 58 . 10  Pearly Everlasting  July Aug .  76 . 2  10 . 86 5 . 00  14 .02 5 . 20  . 9 .79 5 . 80  18 .63 28 .90  46 . 60 5 1 . 10  June •July Aug .  81. 6 82 . 0  1 1 .28 12 .58 4 . 10  12 . 55 15 . 53 7 . 80  10 . 81 10 . 35 5 .40  16 . 73 26 . 84 35 .30-  48 . 63 34 . 69 47 . 4 0  July Aug .  77 . 0  12 •22 3 . 00  9 . 29 5 .00  6 .72 6 .40  19 .98 • 20  • 47 . 15 50 . 40  3 .40  5 .90  3 .70  36 .00  Thistle  Lettuce White Hawkwe e d Ferns  and  -  -  -  Aug .  '  3  5  5.1. 00  Grasses  Bracken  June July Aug .  85 . 6 60 . 6 59 . 3  33 . 24 • 1 1 .90 6 .65  15 . 25 14 .75 5 .00  8 .22 7 .15 .8 .75  9 .73 • 22 . 77 21 . 25  Swordfern  Dec .  61. 9  . 12 .48  10 . 84  4 .'5 6  28 .79  43 . 32  Grasses  July Aug .  53 . 0  9 . 84 11 . 80  • 5 . 30 6 .30  43 . 56 31 .40  33 . 20 54 . 40  8 . 10 3 . 10  35 . 55 . 43 .4 2 57 . 35  . higher  Crude (p  icantly  different  western  levels These July  species  i n December  t o December four  in'late.winter  of  than  0.01)..  shrubs.and  trees  winter  maintenance  alder, and  i n which  opposite  highest  (Table 8 ) .  to offset  (p  a  general of the  thus  increases  highest  that  The o n l y  was c u r r e n t  bordered  this  species i n was r e d  were r e c o r d e d  i n July  that  browsing  i n the seasonal  to the  trend  significant  only  on t h i s  i n late plant  ..  1  Differences  advances,  contributing significantly  N.F.E. l e v e l s  win-  N.F.E. 'content'  as phenology  of ungulates.  I t i s perhaps  were ^higher  0.05) o r e a r l y  was n o t c o n s i s t e n t ,, w i t h  noted.  general  f i r and  f o r the s i x species  (1965 ) r e p o r t e d  ration  and September  nutrients  having  enough  The two  i n p r o t e i n i n d i c a t e d by e a c h  generally  dormancy,  which  August.  August  signif-  i n protein  i n July  i n e i t h e r summer  Dietz  winter  group  both  levels  apparently  (Douglas  trends  species,  N.F.E. l e v e l s  toward  the  winter, hut not  i n the test  and lowest  decline  t o be • s i g n i f i c a n t l y ,  plants.  Average  (p  late  seasonal  four  were  found  summer a n d e a r l y w i n t e r .  included  of the other  two s p e c i e s  other  ter  between  r e d c e d a r ) .showed  those  were  .05) i n summer t h a n  coniferous  to  protein levels  levels  on s i g n i f i c a n c e ( a l l p  r e l a t i o n s h i p s and c o n c l u s i o n s  of the  remaining  0.14).  which can be.  .The-,  62  Table  9  Analysis of variance f o r differences i n nutrient 1 composition of s i x important deer forage species c o l l e c t e d i n J u l y , December and M a r c h . Values .of F a r e s h o w n . 2  Source  Crude Protein  df  Between Between Error  species seasons  Total  5 2 10  4.39' 4.65 =  Crude Fat-  Ash  1.78 2.31  5.942 . 67  Crude Fibre  N. F.E  4 . 21 3 . 11  1.85 9.9 7**  17  Douglas f i r ,western r e d cedar, s a l a l , trailing b l a c k b e r r y , r e d a l d e r and w i l l o w . 2 I n t e r a c t i o n s w e r e n o t s e p a r a b l e f r o m e r r o r mean s q u a r e s b e c a u s e r e p l i c a t e samples were n o t a v a i l a b l e f o r all species. " D i f f e r e n c e s s i g n i f i c a n t a t ; p < 0.05 * * D i f f e r e n c e s s i g n i f i c a n t a t p < 0.01  Table  10  R e s u l t s o f D u n c a n ' s new m u l t i p l e r a n g e t e s t a p p l i e d t o t h e s e a s o n a l means f o r c r u d e p r o t e i n and n i t r o g e n f r e e e x t r a c t d e r i v e d f o r t h e a n a l y s i s o f v a r i a n c e s h o w n i n T a b l e 9. Any two m e a n s n o t u n d e r s c o r e d b y t h e same l i n e a r e s i g n i f i c a n t l y d i f f e r e n t a t p <• 0 . 0 5 .  Mean p e r c e n t a g e Crude  for:  protein  Nitrogen-free  extract  Mar.  Dec.  July  6.6 3  9.6 3  10.46  58.69  49.91  51.81  63  suggested,, t h e r e f o r e , highest  during early  highest  d u r i n g summer  levels had  were  greens; both  crude  summer  growing  period  Dietz  crude  i n the deciduous  percentages  and l o w e s t  plants;  fiber  plants  containing  d u r i n g each  and  crude  levels  than  levels  fiber  were  were  high i n  seasonal  h i g h e s t d u r i n g t h e summer  during late  winter.  Colorado  t h e h i g h e s t amounts  season,  during  i n the ever-  no c o n s i s t e n t  et_ aJ_. ( 1 9 5 8 ) r e p o r t e d t h a t  browse  nutrients  by March;  and w i n t e r b u t showed  moisture  plants  were  winter f o r a l l s i x species but  f a t (ether extractive)  trend;  "choose  i n the deciduous  slightly lower  ash. [ m i n e r a l ) l e v e l s  winter i n the evergreen  highest i n early  declined  summer w e r e  are that:  especially  mule  deer  of important  i n t h e case  ofpro-  i'  tein".  A similar  relationship  Bay.  For example,  ically  the portion  i s indicated  at  Northwest  of bracken  fern  shoots,  June samples eaten  by d e e r ,  crude  p r o t e i n ' ( r a n g e 29.7 p e r c e n t  month  later,  frond  stage  creased per also  when a l m o s t and were  by a l m o s t  cent  two t h i r d s  t o 12.1 p e r c e n t ) .  shoots  than  fibre  33.2 p e r c e n t  t o 35.6 p e r c e n t ) . ,  a l l plants  rarely  h i g h e r , and crude  averaged  had r e a c h e d  browsed,  average  t o 11.9 p e r c e n t Moisture lower  typ-  One  t h e open  protein (range  had de11.6  and a s h c o n t e n t  i n the preferred  were  new  i n the mature f r o n d s .  Another  e x a m p l e was n o t e d  with red alder  and t h i m b l e -  64  berry.  Among  levels, leaves red at  August  samples,,  and t h e l o w e s t of these  alder, a time  fiber  two s p e c i e s  both  were  when o t h e r  the highest protein  .levels (Table  browsed plants  were r e c o r d e d 8).'  only i n late sampled  were  and f a t i n the  As n o t e d  above f o r  summer  and autumn,  of low n u t r i t i o n a l  quality.  Six red  cedar,  grape fats  salal,  species,  trailing  i n early  winter.  seasons  a group  about  western  and Oregon  of protein  compositions  60 p e r c e n t  fir,  fern,  some o f t h e s e  chemical  comprised  Douglas  high percentages  Although  when t h e i r  they  including  b l a c k b e r r y , sword  contained suprisingly  other as  evergreen  were had  and  eaten i n changed,  of the winter  diet.  Finally, pearly  the crude  everlasting  protein!,  f a t , and a s h c o n t e n t s o f  and c a t s e a r d e c l i n e d  and  A u g u s t , c o i n c i d e n t a lwith a decrease  the  rumen  :  black  A similar  rumen  two  plants a time  It  samples, were when  Although  signs of current browsing  observed' o n l y between their  nutrient  c a n be c o n c l u d e d ,  icant  seasonal variation  plant  groups  as w e l l  between  of these  relationship  r a s p b e r r y and s a l m o n b e r r y .  the  at  samples.  markedly  spring  was  June  foods i n  found'with  not abundent i n on t h e s e  latter  and mid-summer -  c o n t e n t s were h i g h e s t .  therefore,  i n the chemical  as o f i n d i v i d u a l  that  there  i s signif-  composition  species.  of  Since  major floral  65  composition changes stages  in. the must  variation the  differs  nutritional  also occur.  in nutrient  contention that  plants,  but  able  at  each  Deer  Use  of  the  equal  heavily  Serai  de e r , f o r treated the  verted  square was  are in  not  nutritious  used  by  serai  between  the  serai  a  the. a n i m a l s  as  the  seasonal  to  most  support  nutritious  association  stages  of  of  in a  as  used  i n preference  upon d a t a  mature  units  from  reflect  (Harper,  equal used  stages,  age.  avail-  of  in this  time.  here, to  85  study  use" The  simply  After  the  most  1967).  This  preference  (2)  are  index  per  actually  to  autumn  counts  age-classes  and  con-,  over  per. s q u a r e  decimal, numbers.  s p r i n g and  that  "deer-days  acre"  by  differently  occurring  "deer  ''deer-days use. o f  and  numbers  It i s understood  r e p r e s e n t i n g 17  timber.  community.  densities  "deer-days.of  short periods  serai  f u n c t i o n of. d e e r  basis for determining  a s s o c i a t e d w i t h the  adjacent  are  different-aged serai  m i l e " , or  early  these  habits tend  p r e f e r r e d communities  expressions  based  food  only.select  densities  density indices  chosen  fusion  group  or  serai  relatively  and  seasonal  Communities  spent  was (1)  deer  relationships  intervals,•highest  used  criterion  overpaid,  s t a t u s .of e a c h , o f  levels  most  stage,  season.  time  time  serai  The  deer  a l s o the  Pellet and  i n each  per mile",  avoid  con-  Results conducted the  66  A l t h o u g h , i t i s shown l a t e r nificantly means were  enhances  f o r both used  range  hurned  to describe  trends  a more r e a l i s t i c  ships  between, deer  since  burned,  i n site  approach  and f o r e s t  partially  logging  period  This  pro-  relation-  sites  area,  were  i n F i g . 11.  of developing, s e r a i  or burning  and r e a c h e d  per square  mile  peak  Gaultheria-Hypochaeris use  equivalent  was  recorded.  succession dense  may  the next  upward  reflect  I t may  to four years equivalent  s i x years  the  mile  beginning  a tendency  also reflect  A  late  (12 a c r e s  time-  per deer)  i n the 12th year  of deer  developing  after  t o 140  recorded.  into  after  i n the early .  p e r d e e r ) were  per square  trend  p r o t e c t i v e cover  stages..  soon  a s s o c i a t i o n f o l l o w e d , by w h i c h  t o 55 d e e r  A slight  three  t h e . summer,,  began  intensity  use-indices  (4.5 acres  d e c l i n e through  During  communities  time,  At t h a t  - Trends i n  and f o r the. a u t u m n - s p r i n g  disturbance.  serai  11  i n the study  succession  association,  of  i n Table  selection.  and unburned  Gaultheria-Hypochaeris  marked  given  regeneration  to serai  a r e shown g r a p h i c a l l y  exploitation  deer  the combined  to the e c o l o g i c a l  burned,  use i n r e l a t i o n  u s e f o r t h e summer  period  by d e e r ,  associated.  Seasonal deer  selection  slash-burning sig-.  and u n b u r n e d . a r e a s  vided  closely  that  to take  i n t h e more  advantage advanced  g r e a t e r .use o f t h e  G a u l t h e r ia-Hypochaer i s a s s o c i a t i o n and t h e e a r l y  of  late  Gaultheria-  Table  11  Mean i n d i c e s o f d e e r u s e i n d i f f e r e n t - a g e d s e r a i s i t e s . V a l u e s were d e t e r m i n e d by p e l l e t g r o u p c o u n t s a n d a r e e x p r e s s e d as d e e r p e r square mile.-'The n u m b e r o f r e p l i c a t e s a m p l e s o b t a i n e d f o r e a c h a g e - c l a s s i s shown i n p a r e n t h e s e s .  Years since l o g g i n g and burning 2 3 4 5. 6 7 8 9 10 11 12 13 14 15 16 17 t imber  deer  per  Summer U s e - p e r i o d Burned . Unburned Pooled Sites Sites data 106 . 6 118 . 8 181. 2 152 . 8 58 . 5 97 . 0 35 . 4 126 . 6 99 . 6 24 . 8 50 . 1 75 . 1 61.  -  44 . 2 4 7 .5  -  (1) (3) (5) (3) (2) (4) (2) (2) (1) • (3) (3) (3) (3) (1) (1)  99 . 3 113 . 4 166 . 2 85 . 5 37 . 8 Ill. 5 70 . 9 . 63 . 2 86 . 4  -  43 . 1 54 . 1 86 . 1 -  -  -  (1)  CD' (D. (D CD (D (2)  CD CD CD  (3)' (2)  106 . 6 113 . 9 169 . 9 156 . 1 52 . 2 85 . 2 51. 8 98 . 8 81. 4 40 . 2 50 . 1 67 . 4 57 . 8 86 . 1 44 . 2 47 . 5 55 . 0  See p a g e 22 f o r m e t h o d square m i l e .  of  (D C4) C6) (4) (3) (5) (3) (4) (2) (4) (3) • (4) (6) (2) (1) (1) (5)  Autumn Burned sites 110 . 4 I l l .7  to  Spring Use-period Unburned Pooled sites data  CD (4)  79. 3  (D  44. 2 60 . 5 69 . 4  (D  71. 8  (D  68 . 7  (D  29 . 4  (3)  57 . 4  (3)  •  -  -  -  -  -  -3 . 3  (2)  CD  (D  -  -  converting  pellet  -  25 . 0. (D 95 . 2 CD • 64 . 7 ( D  -  -  48 . U 45 . 9  -  • -  group  110 .4 ( D 103 . 2 C5)  -  58 60 47 95 66  • -  . 0 (2) . 5 (2) o (2) •. 2 (D .7 ( 2 )  29 . 4 ( 3 )  -  (2)  (D  5 3 . 7 (5) 45 . 9 ( D  -  3 . 3 (D  -  42 . 9 ( 2 )  densities  to  Fig.  11.  T h e r e l a t i v e u t i l i z a t i o n b y d e e r o-f e a r l y serai s t a g e s d u r i n g t h e summer a n d t h e a u t u m n - s p r i n g period.  Senecio-  GaultheriQ - Hypochaeris Association GaultheriaE a r l y -Intermediate— A d v a n c e d Epilobium (Cirsium-Epilobium (Epilobiurn-Rubus (Pseudotsuac-Rubus PseudotsuqaAssoc. Subdom.) Subdom.) Subdom.) Assoc.  S years  10 12 s i n c e burning  14  16  Pseudotsuqo Subclimax (Gauifheric Understory)  180-200  Pseudotsuga  association  mer a n d autumn b r o w s e The  shift  ceous  to these  growth  dries  From stages (Fig. in  parallel  Early  square  a b o v e , p r e f e r r e d sum abundant occurs  deer  was  sites  peak  densities  per deer) - about  arly,  the intermediate Gaultheria-Hypochaeris  Gaultheria-Pseudotsuga  autumn-spring Hypochaeris during  p e r i o d than  both  sample  later  indicating  study  area  being  the case,  ter,  deer  that  association, used  tend  Age  less  fewer  deer  i t ' w o u l d appear to concentrate  curve  i n the  will  be  intensive presented  w i t h i n the  p e r i o d than  i n summer.  that  autumn  during  and  This win-  late Gaultheriato other  of the p e l l e t  s p r i n g o f 1961 and 1962 ( t h o s e  spring  association,  The l a t e G a u l t h e r i a -  were p r e s e n t  i n preference  classification  deer  and t h e mature  Evidence  i n this  most  Simil-  however, r e c e i v e d e q u a l l y  periods.  association  were  summer.  intensively  i n summer.  i n the autumn-spring  Hypochaer is  the  were  association,  lower  26 p e r c e n t  than  use  recorded, during  serai  months  d e n s i t i e s . ' e q u i v a l e n t t o 100  ( s i x acres  subclimax  herba-  substantially  Gaultheria-Hypochaeris  mile  Pseudotsuga  after  o f t h e summer  lower  the  varied.  use o f the v a r i o u s  to that  age-classes  supporting  and  unpalatable.  to spring,  1 1 ) , but in.most  used,  a r e most  and becomes  was b a s i c a l l y  heavily per  species  as shown  associations likely  autumn  intensity.  where,  used  o f F i g . 11)' a r e s u m m a r i z e d  serai  groups  stages.  counted  in  t o d e r i v e t h e autumn In Table  12.  The  Table  12  Age c l a s s i f i c a t i o n o f p e l l e t g r o u p s encountered d u r i n g t h e r e g u l a r s y s t e m a t i c May c o u n t s o f 1961 and 1962. Age  Age o f . S i t e . (years ) 1 7  to to  6 Mature timber  "fresh" been  sified  s i x , 48.6 as  "fresh"  serai  stages  these  categories.  the  very  ful)  (23.0)  421  fresh"  24.3  curve  distinct  of  early  increases intensity  rapidly i n the  quickly  (where  were  can  of  soon  the  In  not  the  a a  -  clas-  placed the  were  more  Thus,  have  advanced  half  plentiful),  considered  only other  the  use  of  plenti-  about 75  per  autumn-  s y n t h e s i s of pattern  d e f o r e s t a t i o n , reaches  1  year;  and  and  an  in  advanced  spring  fourth year,  sixth;  were  were  i n s p r i n g , the  after  period  i n the  groups  to  age-classe.s  counted  deer-use:  or  In  s p r i n g foods  winter. be  third  after  1  spring.  and  patterns  the  (75.7  V  counting  fresh"; of  (51.4)  considered  approximately  occurred  F i g . 11  the  a l l groups  cent  stages  s p r i n g foods  seasonal  relatively  of  per  the- u s e  of  were  s p r i n g growth.  Therefore,  serai  of  groups  months  of  o c c u r r i n g i n autumn  spring  which  12 8  "moderately  i n the  (where  quarter  cent  peak  early  occurred  stages one  only  (%)  (.1.3 )  cent  or  Aged ' No . 169  onset per  Groups  (43.7)  w i t h i n two the  Pellet  14 4  "moderately  to  of  (4.9)  7  deposited  to  (%•)  16  and  subsequent one  Fresh No.  Categories Mod . Fresh No . (%)  declines autumn-  two  71  winter tion,  p a t t e r n which, r i s e s reaches  depending  on  relatively seasonal  peaks  effects  the  data  data  ences  was  were  cu  logging  that, even the  by  the  perhaps burned  than  range  than  use  sites  to  stage.  slash  were  remains  T- A  the  more  10  each  remove, as  per  intensive  freer  cent use  of by  that  They  much a s the deer  an  90  larger of  the  and  higher  levels, i n foods (.Ei.narsen,  and  the  average"  of  clear-cut also per  concluded cent  of  wood.  site.  sites It  foods,  growing  1966 ) .  which  differ-  burned  availability•of  of  for  0.01)  after  greater variety  i n unburned" s i t e s  The  (p <  acre  movement w i t h i n  nutrient  11  A t_ T e s t  available..  Douglas . f i r , f o r e s t s .  slash-fires  of  i n Table  age-classes  the.summer  on  two  comparison  areas.  (1937 ) r e p o r t e d  Hopkins  The  s l a s h - b u r n i n g w e r e -more  only  f o r both  and,  declines  presented  unburned  using  year,  . ..'  and  reflects  even  subclimax  0.05).  light  reflect  the  unburned  deer  s m a l l e r d e b r i s and  p'robably  to  p e r i o d .(p <  coastal  Therefore,  also  and  executed,  f t of  of  thirteenth  a p p a r e n t , i n F i g . 11.  significant  Isaac  deforesta-  d u r a t i o n of. s n o w f a l l ,  both, t r e a t m e n t s  autumn-spring  29,000  e l e v e n t h to  areas.subjected  used  from  gradually after  s l a s h - b u r n i n g on  that  intensively  and  through  are  of  i n the  depth  f o r burned  indicated  samples  the  little  The  these  a peak  very  in  may and  Figures two  logged  burned,  units,  one i n w h i c h  the other  groundsel, dant  12 a n d 13 c o m p a r e  i n which  lettuce,  i n the burned  early  serai  the slash  burning  was  b l a c k ' r a s p b e r r y and w i l l o w h e r b site;  residual  salal  site.  Estimated  Number  Area  study page  area, 22.  o f Deer  based  o f e s t i m a t i n g numbers on p e l l e t  The f i r s t  acy  o f t h e method  let  group  sampling  group  of deer  densities,  was f u l f i l l e d .  i n each  plot,  when  i s described  ( S t e e l e and T o r r i e ,  on  the accur-  Estimates  transformed  t r a n s f o r m a t i o n and a p p l i e d t o S t e i n ' s  procedure  plant  within the  o f t h e c o n d i t i o n s upon which  depends  densities  square-root  i n the Study  Wood  a r e abun-  i s the.only  i n q u a n t i t y i n the unburned  method  thoroughly  was n o t a . t t e m p t e d .  available  The  conditions i n  of  pel-  with the two-stage  1960: 8 6 ) , i n d i c a t e d  that  the desired sampling  level  o f 70 p e r c e n t  with  10 p e r c e n t  error  ( R o b i n e t t e e t a l . 1 9 5 8 ) was  sampling  obtained'.  The s e c o n d ' a s s u m p t i o n  fulfilled.  F o r some a g e - c l a s s e s  between r e p l i c a t e most, and  however,  t h e method  approximation  The Table  13.  estimates  may  c a n be e x p e c t e d  use were  to produce  noted.  For  an  acceptable  size.  of the calculations  In the winter  entirely  were r e a s o n a b l y • c o n s i s t e n t ,  to the true population  results  n o t have been  of land, large differences  of deer  the r e p l i c a t e s  confidence  are presented i n  o f 1961-62 t h e r e  were  estimated  to  4  Fig.  12 A S e n e c i o - E p i l o b i u m a s s o c i a t i o n o n e y e a r l o g g i n g and b r o a d c a s t b u r n i n g ( P l o t A ) . u s e i n t h i s p l o t was i n t e n s i v e .  after Deer  Fig.  13 An u n b u r n e d a r e a o n e y e a r a f t e r l o g g i n g . Prod u c t i v i t y was l i m i t e d t o s a l a l s u r v i v i n g from the mature timber stage. A c c e s s t o , and movem e n t t h r o u g h t h e a r e a b y d e e r was noticeably restricted.  Table  13  E s t i m a t e s o f t h e number o f d e e r e r a t i n g l a n d , and t h e e s t i m a t e d  s u p p o r t e d by e a c h a g e - c l a s s o f r e g e n deer p o p u l a t i o n i n t h e study a r e a .  A u t u m n - S p r i n g 1 9 6 1-62 .rs s i n c e .Total area Use i n d e x No. d e e r i n ;ging o r (sq mi) (de e r / s q age-class :rning m i ) (CD) CO (D) 0 1 2 3 4 5 6 n i  8 9 10 11 12 13 14 ' • 15 16 17 Timber Totals  . 22 . 19 .16 . 29 . 14 .28 . 04 . 12 . 32 -.61 .27 .78 .25 2 .25 .42 . 15 3 . 64 10 . 15  _  c  63 98 102 92 76 63 53 49 49 51 53 54 48 38 —  9 43  _ 12 . 0 15.7. 29 . 6 12.9 21.3 2. 5 6 .4 15.7 29 . 9 13 . 8 41.3 13.5 108.5 16.0 1.4 155.5 49 7  Total area ( s q m i ). (A) .02 .22 .19 .16 . 29 . 14 .28 . 04 . 12 .32 .61 . 27 .78 .25 .2.26 .42 ..16 3.62 10 . 15  Summer-1962 Use i n d e x No. d e e r i n (deer/sq age-class mi ) (AB) (B) _ 63 120 140 140 13 0114 . 8 6 79 65 56 53 ' 54 60 64 65 40 55  13.9 22.8 2 2.4 40.6 18 . 2 31 . 9 3.4 9.4 20 . 8 34.1 .14.3 42.1 15 .0 144,6 27.3  -  6 . 4 199.1 666  75  be  497 d e e r  for  every  i n the t e n square  13 a c r e s .  m i l e s - an a v e r a g e  I n t h e summer  creased  by a p p r o x i m a t e l y  animals  - slightly  less  o f 1 9 6 2 , numbers  34 per cent than  one d e e r  f o r every  i n c r e a s e i s b e l i e v e d t o be due p a r t l y  and  partly  Smith  (1968) concluded  Northwest after  fawning  average tween per  cent.  of  tagged  indicate  outside  as fawns  spend  the north  movement i n t o  because winter  climatic food  adjacent  is  perhaps  area  a t lower  considered  entirely  support  op.cit. )  summer  i n the  win-  i s probable  r i g o r o u s t h e r e and  numbers  reasonable.  to  elevations just  areas  second-growth  R i v e r appears  i n c r e a s e i n deer  attributable  A significant  peripheral  conditions are less  t o the Englishman  be  of the  (Smith,  which  boundaries.  the lower,  o f 20 t o 25  gathered  individuals  the winter•months  The  o b s e r v a t i o n s and r e t u r n s  i n the study  and e a s t  immediately  somewhere b e -  would  data-were  but limited  some  lies  above  i n the  by A u g u s t .  i n the order  p r o d u c t i o n i n the advanced  34 p e r c e n t  i n June  ranges.  o f 10 t o 14 p e r c e n t  No s p e c i f i c  that at least  study.area  a  two e x t r e m e s ,  assumption,  to birth  a s 35 p e r c e n t  probably  34 p e r c e n t ' i n c r e a s e n o t e d  deer  ter  may b e a s h i g h  I f so, a balance  latter  ten acres.  winter  and as l o w as 18.9 p e r c e n t  immigration.  this  adjacent  566  t h a t r e c r u i t m e n t through, b i r t h  r e c r u i t m e n t t o August  these  total to  Bay h e r d  from  deer  had i n -  t o an e s t i m a t e d  The  to immigration  o f one  stands  t o be h i g h .  from  winter  Thus,  t o summer  76  Also tion  considered reasonable  i n numbers between  t h e summer.of  1962-63.  This  population  t o t h e same l e v e l  1961-62.  Smith  years as  1960 t o 1 9 6 2 , a n n u a l  mortality,  additional  10 p e r c e n t .  that  e x c l u d i n g summer Therefore,  are consistent with  of the  i n each  hunting mortality  a n d summer p o p u l a t i o n s i z e s  densities  return  f o r the winter of  of the Northwest  cellaneous  herd .  estimated  (op.cit . ) concluded  reduc-  1962 and t h e w i n t e r o f  represents the t h e o r e t i c a l  m u c h a s 20 p e r c e n t  winter  i s a 26 p e r c e n t  accounted f o r  Bay h e r d , fawn  of the  and  deaths,  the differences  indicated  independent  mis-  f o r an between  by p e l l e t  group  studies of the  77  DISCUSSION  This tative  changes  mine  the  tion  by  of  the  role  content  The  and  implicated  as  The  the  broad-leaved  composition, nutrient  forage  s u c c e s s i o n , and range  range  be  selec-  that,  for four  use.  variety,  content  of  range  available  selection.  i n the  not  food  nutrient  found  to  occur  t h e r e f o r e c o u l d not  be  selection.  i n the  recognized  survey,  accuracy  the  growth  are  data,  due  cover forms,  not  percentages Only, t h e  recorded,  inherently  mainly  as' i n f l u e n c i n g  f o r e x a m p l e , may  t r a n s e c t p o i n t was shrubs  deter-  range  deer  food  seasonal  s p e c i e s were  to  relationship  consistent trends  limitations  different  each  a close  vege-  data  should  with, e q u a l  at  and  and  demonstrated  s u c c e s s i o n and  seasonal  influencing  of  used,  displaying plant  have  serai  describe  in influencing  associated with  serai  Certain  ated  i n the  important  Limitations  sults.  of  to  communities,  investigated,  significant  advancing  methods  results  stage  closely  of  designed  changes  occurring in floral  are  However,  these  factors  production,  with  of  between  Changes  was  o c c u r r i n g i n .serai  deer. five  exists  foods  investigation  '  more  and  likely  the  have of  to  the re-  estim-  plants  uppermost since  tall  t o be " h i t "  78  than  either  plants ses  such  and  as  been  may  case  here,  a measure  undertaken  measured tend  may  an  error  would  tend  most  advanced  serai  stages  forage as  availability.  food  production.  d e n s i t y , whether  separately in this the need  are normally  However, where  within  variability  occurs  stocking prevent  rate of s a l a l the development  associations artificial and  was  described  s t o c k i n g was  negligible  and  f i r ,  o f some o f t h e  limited  i n the s i t e s  As  error  only.  Cover  i f sizes  was esti-  i f  (diameters) uniform.  does  in a  stocking density the  developing.  or Douglas  above.  this  measurements  the chronology, stages  .was  (as i t u s u a l l y  i n modifying of s e r a i  investigation  are r e l a t i v e l y  be  t h e number  the  Thus,  f o r such  s h r u b - c o n i f e r community),  even  not  investigation.  species  are  n a t u r a l or a r t i f i c i a l ; ,  regenerating important  estimates  composition  distributed  each  shrubs  ' T h i s - was  clipping  of cover  t o be  where  serious i f cover  a separate  to estimates  individuals  as  the shrubs  t o overcome  species  gras-  Conversely,  to estimate  Stocking  of  Of  however,  restricted  or g r o u n d - l e v e l s p e c i e s ,  proportionately underrated  It is potentially  as  most  been  Such  i n t h e more  used  mates  forbs  of the community.  dominate.  not  have  overrated.  prevalent  is  leader  catsear, fireweed, pearly everlasting,  sedges  components have  single  sequence,  may., or  A high natural  f o r example,  may  catsear-dominated  pointed  to Douglas  out  earlier,  f i r seedlings,  s e l e c t e d f o r study.  Thus,  any  differences  between  equal-aged  The deer  production  given  season  ample,  figures  plants  i n early  substantially portionately stages. deer  mer m o n t h s  than  the standing  s o when  compared  i n these  exhibited  that  total  Therefore,  occur  i n estimates  food  standing  by d e e r . crop  with  that  produccrop  are  probably dispro-  f o r other  highest  the greatest  dominated  discrepancy  o f the key foods  pearly  and  F o r ex-  serai  spring-summer  ( F i g . 1 1 ) , and i n t h e s p r i n g plants  as a  of herbaceous  sites  those  by  which a t  p r o d u c t i v i t y , and  sites  herbaceous  4).  earlier  of total  used  factors  taken  i n s e r a i stages  Gaultheria-Hypochaeris  lower  already  between  heavily  estimating  crop  estimation  composition  to natural  was m e n t i o n e d  be g r e a t e s t  a r e most  I t was  u s e was  of the annual  the differences would  or i n cover  was d u e e n t i r e l y  accurate  Naturally  total  density  o f measurement  preventing  tion.  sites  proportion  at the time  factor  any  i n species  a n d sum-  the diet  would  (Fig.  probably  e v e r l a s t i n g and  catsear.  Taber which all of  produce  new-growth  and Dasmann errors  i n estimating  on p a l a t a b l e  certain behavioural  nate  habits' such  central  (1958) d i s c u s s e d  portions  plants  other  factors  a v a i l a b l e foods. is. p o t e n t i a l food  c h a r a c t e r i s t i c s s.hown b y d e e r .  as t h e t e n d e n c y of large  open  to avoid  areas  dense  (see also  Not because In-  t h i c k e t s and Harper, 1967 ) ,  80  and  the  trees  utilization  growing  on  considered.  of  only  hillsides  I f not,  Although  localized  unburned. s i t e s ,  methods,  Northwest food  Bay  few  and  tend  (see  the  upon  shrubs  food  and be  production  thickets  do  exist  topography,  in  patch-  slash-burning  program  the  effect  of  behaviour  the  availability  variety  of  may  a l s o depend  to  some  plants present.  dominated  by  a  appear  to  be  highly productive.  al  and  potential  of  the  lack of  the  dense  dominated the  of  gentle  minimize  of  a l s o C o w a n , 1945.) m u s t  impenetrable  extensive  to  sides  at on  availability.  Food  for  lovrer  over-estimates  result.  logging  a  the  single  variety. second  growth  a c t u a l consumption  preclude factor  the  the  after  r e g r e s s i o n of  by  Hopkins,  1937).  succession,  soil  as  be  would  low  because  estimates and  for  higher  ever  do  not  i s an  are  rich  salalthan  be..  completely important  concentrations  in  growth  initial  mayactu-  nutrient levels.  p o s s i b l y other the  may  being  analyses  forage  salal)  however,  f i r stands  succession  In  as  production  deer  slash-burning, soils  (Isaac  of  food  recognized  that  n i t r o g e n , and  years  this  Douglas  food  potassium, and  of  proximate,  possibility  causing  mediately  of  of  (such  In r e a l i t y ,  Therefore,  a s s o c i a t i o n s are  Results  Serai associations  palatable species  utilization  extent  Im-  calcium,  elements  two  decline  or  three  rapidly  81  through slowly  erosion, decaying  1958). the  For  present  effects  were  already  the  trends  1956). many  (Cowan But  with  ranging high  from  in early  of  place  and  plants,  land  for  were  trees  sampled  from  Major from  which in  and  t h i s . t y p e of  the The  samples  masking  Harris  analysed  in  testing.the  obtained.  important  sample  or  Dasmann,  when s a m p l e s  a _ l . 1950 , C o o k each  and  availability.  have been et  Taber  unsuitable  communities  recognized be  per  defecation  spring Taber  in this deer  that  slightly  rates  i n seasonal  a mid-winter  (Dasmann and  groups  deer  differences  calculations  may  taken  since  individual  been  shrubs  also  also  age-classes  nutrient  have  in perennial  was  error  1950 , pooled  is  consider-  minimal.  vary  13  on  storage (see  the  have  individual  Finally,  ed  may  c o l l e c t e d may  Swank  ed  reason,  (four-year-old)  nutrient  from  this  and  tissues  succession  c h a n g e s may  •age o f  plant  study  of  youngest  leaching,  of  17  low  of  food 10  groups  were  (Eberhardt  Van  use-indices • derived high.  Those  iod, however, are  probably  discussed  above,  almost  exhibited  during  the  for  the  reasonably h a l f the  use  autumn-spring  per  deer  on  deer  daily  Etten for  to  have been  a  autumn  record-  average  summer to  period  spring  because,  of  serai  period  appeared  of  i t is  accurate early  a  Since a l l  1956),  the  to  Daily"rates  ejt a l . 1958 ) .  based  and  habits.  groups  per  195 5 , R o g e r s study  have been r e p o r t e d  peras  stages to  have  82  occurred rates  i n early  would  tend  with, t h e a v e r a g e  The and  spring. to balance being  group  early  Gaultheria-Hypochaeris  after  three  during  tity use  There  of food of this  mental  available  support  four  habits,  this  argument.  was t h e  stage.developing In these  sites  t o 140 a n i m a l s p e r q u a l i t y and quan-  stimulating  as p r o t e c t i v e  i n conjunction Herbaceous  the spring-summer o f changes  indicated  abundant  after  diet  i n range  that  i n terms  and b u r n i n g ;  years  greatest  the serai  a n d summer  that  factors  spring  when  such  other  cover  heavy  environ-  are not r e a d i l y  here.  advanced,  logging  doubt  association, particularly  Measurements  most  that  of regeneration.  i s little  Food  dominate  spring  u s e was e q u i v a l e n t  such  rates,  selection of  association, a  are the primary  requirements  defecation  indicated  the.late  years  the•• i n t e n s i t y o f d e e r square, m i l e .  arid''the  densities  preferred  defecation:  t o 13 g r o u p s p e r d e e r .  food  stage  to five  low: w i n t e r  close  r e l a t i o n s h i p between summer r a n g e • ,  Pellet  I f ao, high, s p r i n g  these  logging  plants  were Bay  composition,  as  cover'three  succession  years  present  to s i x years  deer..  species  a v a i l a b l e on a w e i g h t  five  results,  shown t o  of Northwest  and b u r n i n g ;  v a r i e t y of species  the other  same h e r b a c e o u s  of ground most  with  after  basis  i n the after  were:  logging  and b u r n i n g .  nutrient summer other  foods  If  (1958),  this  period  superior  terms  into  plants  when J u n e  that  were  samples  maxwere  not a l l i n d i v i d u a l a t any g i v e n  by T a b e r foods  time.  taken  and Dasduring  adequate, but also  variety,  therefore, the early to provide  the growing  developing  an a v e r a g e  of active  dominating  of the  and t h e r e f o r e most nu-  as s u g g e s t e d  of a v a i l a b i l i t y ,  food  serai  and  nutritional  Gaultheria-Hypochaeris spring-summer  conversion  i s considered  of  condi-  solar  t o be  greatest  stage.  intensive exploitation  a s s o c i a t i o n s cannot  plant  this  ideal  season,  summer,  Gaultheria-Hypochaeris period  Although  nutrient levels  nutritionally  a v a i l a b l e , deer  early  compared t o  available.  appeared  During  In  the  that  the youngest  not only  when  at the beginning  i t c a n be c o n c l u d e d were  of spring-  of the year.  Nevertheless,  available,  of food,  association  this  1965).  to"others  In  in  time  not analysed  to select  plants  mann  energy  at this  trends i n  the quality  was g e n e r a l l y h i g h  i ti s likely  (Dietz,  tend  tritious  tions.  that  a r e a t t h e same p h e n o l o g i c a l s t a g e  deer  quality  seasonal  and had d e c l i n e d s l i g h t l y  collected plants  foods  were  season,  then  indicated  not eaten  samples  growing imal  composition  herbaceous  forage  Furthermore,  growth.  community  Most  continue  of  early  beyond  of the herbaceous  become u n p a l a t a b l e  as  they  84  approach forced  maturity.  to feed  stages,  Thus, w i t h  the advanced  Gaultheria-Pseudotsuga  noted  was r e f l e c t e d  salal  of various  berries,  ground  time,  been  This  latter  summer  higher  here  shift  i n range of use  (Fig. 11). foods,  and  Avail-  especially  an i m p o r t a n t ' f a c t o r s t i m u l a t i n g Abundance  i n terms  was h i g h e s t  at  this  of the preferred portions  under  could  of salal  production  quality  than  possibility  serai  intensity  and autumn  a n d new g r o w t h  and t h e n u t r i t i v e  have  age-classes  exploitation.  cover  This  by an i n c r e a s e d  i s likely  this, increased of  late  deer are  Gaultheria-Hypochaeris  associations.  i n the 12- t o 14-year  ability-  phenology,  more f r e q u e n t l y i n b r o w s e - d o m i n a t e d  particularly  preference  advancing  mature  timber  (Cowan  n o t be t e s t e d w i t h  may  1945).  the data  available.  Not trend be  associated with  logged  trend areas  after  of western  spring  serai  used, i n t h o s e mals ' h a b i t a t .  Brown  Washington.  bedding  similar  summer  seasons  summer  (1961) r e p o r t e d group  food,  and form  a corres-  densities i n  He a t t r i b u t e d  at Northwest  upward  ( F i g . 11) can  and p r o t e c t i v e c o v e r .  stages, p r o v i d e  and e a r l y  during  estimating pellet  i s undoubtedly  advanced  stages  feeding.  associated with  uation  c o n t r i b u t i n g t o the second  i n the use o f s e r a i  ponding  use  a l l activity  i t to The  Bay.  sit-  Although  very  little  i n t h e way o f .  they  are nevertheless  an i m p o r t a n t  part  heavily  of the ani-  The ber  intensity  i n summer  mile)  was s u r p r i s i n g l y  area.  Cowan  production,  only  at  Northwest  The and  late  accounts  be a t t r i b u t e d  spring  were  stages.  young  stages  winter. foods  forest  may  The h e a v y  support .  use of timber  of the• interspersion the patch-logged  f o r part  of the a c t i v i t y ,  b u t most  to bedding  food  and  occurred  and ofi t  protection.  and t h e s e l e c t i o n  accumulating  indicate  between  o f autumn  autumn  and  early  i n the two- t o f o u r - y e a r - o l d of three  t h a t most i n spring  other - aspects  deer  activity  r a t h e r than  of  this  i n these  i n autumn o r  of autumn-winter  shown  t o be l o w i n t h e e a r l y  serai  items  such  as . s a l a l ,  being, most  western  abundant  and under  autumn-winter  of  envi-  berries  numerous  association  important  of low food  p r o d u c t i o n and v a r i e t y  lichens  tsuga  the study  First.,  were  principal  mile.  square  as b e i n g  f e e d i n g on s a l a l  However, r e s u l t s  investigation  because  of coast  result  within  tim-  per  summer  groups  most  stands  that,  trees within  r e l a t i o n s h i p between w i n t e r ranges  Pellet  eal  per square  of mature  Some  mushrooms  timber  tracts  Bay i s a d i r e c t  stands  ronment.  of mature  undisturbed  few as one d e e r  55 d e e r  h i g h , a n d m u s t be i n t e r p r e t e d  (194-5) e s t i m a t e d  as  small  deer, . u t i l i z e , t h e m a t u r e  (equivalent to approximately  representative  must  to which  red cedar,  and a r b o r -  i n the Gaultheria-Pseudo-  the subclimax  foods  stages, the  were  forest.  not found  t o be  Second, nutri-  86  tionally If  they  b e t t e r i n the' y o u n g had  been,  the  young  stages  the  b e n e f i t s of  protection the  of  therefore, tions  early, of  of  use  i n the  stages,  and  groups So  derived  pellet  parent  from shift  advanced  and  salal-Douglas  available  at  growth to  on  the the  least  the  shrub-  Presumably,  intensities Data  accumula-  the  and  in  develop-  group  highest  conifer-dominof  use  in  concerning  i n s p r i n g counts  support  the  the  age  this  a n a l y s i s , of p o p u l a t i o n , estimates .  group  densities,  i n autumn  which  and  r e v e a l e d an  winter  into  the  apmore  associations.  basis 12-  f i r sites  winter food  the  in  override  i s .available  have r e v e a l e d  stages.  stages.  Third,.little  of p e l l e t  lowest  serai  animals  complex  protection,  autumn and  does  of  Thus, and  content.  advanced,  recorded  reasoning.  would  the  forb-dominated  pellet  annual  insufficient  until  specific.measurements  i n autumn a n d . w i n t e r  serai  of  w i n t e r .weather  forest  old serai  Gaultheria-Pseudotsuga.association.  intensities ated  he  nutrient  inclement coast  i n the  production  probably  a higher  from  the  scant  would  regenerating  ment  the  than  to  of  availability,  15-year-old  are.considered  ranges.  appears  food  t o be  Conversion most  comfort,  salal-catsear t o be of  the p r e f e r r e d  solar  efficient  and  energy  i n these  into sites.  87  CONCLUSIONS  This  study  indicates  •food- a r e - the- p r i m a r y blacktailed  In produce  factors  the greatest variety these  are sites  serai  quantity  stages  of shrubs  these  years  of regeneration.  ity  are sites  quantity  producer affects  The and  perhaps select  o f deer  range  serai  t h e most  statements  lished  that food  then The  deer  within  original  three  summer t o l a t e  which  winter,  twelve  forage.  deer,  and  and c o n i f e r s .  undergone  foods. to five  the greatest variety  the subclimax  Gener-  to  fifteen  food  availabil-  forest  becomes an  i tfollows  then  that  food  selection.  quality  of food  succession. foods  quality.also hypothesis  changes  with  Furthermore, available  i n the previous  a regenerating unit  -by t h e . e f f i c i e n c y  stages  undergone  salal)  have  nutritious  Considering  serai  When, s n o w f a l l r e d u c e s  nutritional with  have  produce  which  by.  secondary.  prefer  late  i n the regenerating areas,  important  to  which  (particularly  ally,  selection  and q u a n t i t y o f herbaceous  From  which  and q u a n t i t y o f  range  cover  and summer, d e e r  of regeneration.  prefer  governing  with protective  spring  Generally, years  deer,  that quality  i n each  paragraph,  affects  range  i s supported: of coast  a t which, r a d i a n t e n e r g y  deer  forest  season appear season.  i t i s estabselection.  t h e numbers o f are affected  i s converted  into .  88  suitahle  and ayai.lab.le  whole  logged  community  ideal  food—producing  food.  Largest  c a n be e x p e c t e d  populations  when t h e number o f  u n i t s f o r each season  i s at a  These . c o n c l u s i o n s l e a d t o a f u r t h e r that  sustained  realized and  only  deer  populations  i f the area  climax.conditions equals  regeneration forest  destruction.  maturity balance never  between  occurs.  Since  the rate than  to  Patch-logging than  clear-cut,  o r some  subclimax  other  into  form o f  of regeneration to  current  operations broadcast  harvest  rates,  a  deterioration  approach  the ideal  o p e r a t i o n s , b u t even  decline i s inevitable  i n the. food-producing  c a n be  stimulated  h a b i t a t c r e a t i o n and h a b i t a t  a population  regression  being  logging, burning  i s f r e q u e n t l y slower  more, c l o s e l y here,  through  forest  succeeding  the area  maximum.  hypothesis:  i n the coast  of land  w i t h i n the  because  capabilities  of  eventual  of the land.  89  LITERATURE  CITED  Association of O f f i c i a l A g r i c u l t u r a l Chemists. 1960. O f f i c i a l m e t h o d s o f a n a l y s i s o f t h e AOAC , 9 t h e d . A s s o c . • O f f i c i a l A g r i c . Chern. W a s h i n g t o n , D.C. Bandy, P . J . 1965. A study o f comparative growth i n four races of b l a c k - t a i l e d deer. Ph.D. t h e s i s , D e p a r t ment o f Z o o l o g y , U n i v e r s i t y o f B r i t i s h C o l u m b i a , Vancouver. 189p. B r o w n , E.R. 1961. T h e b l a c k - t a i l e d , d e e r of- w e s t e r n Washington. W a s h . S t a t e Game D e p t . B i o l . Bull. No. 1 3 . 124 p . B r u g g e m a n n , J . , D. G i e s e c k e a n d K. W a l s e r - K a r s t . 1968. Methods f o r s t u d y i n g m i c r o b i a l d i g e s t i o n i n rumina n t s p o s t mortem w i t h s p e c i a l r e f e r e n c e t o w i l d species. J . W i l d l . Mgmt.'32(l): 198-207. B u c k l a n d , D.C. 1941. Forest s u c c e s s i o n i n burns i n the coast forest. B.Sc. t h e s i s , Department o f Botany, U n i v e r s i t y o f B r i t i s h C o l u m b i a , Vancouver. 48p. C h a t e l a i n , E.F. 1947. Food p r e f e r e n c e s o f t h e C o l u m b i a n b l a c k - t a i l e d deer, ( O d o c o i l e u s hemionus columbianus R i c h a r d s o n ) on t h e T i l l a m o o k B u r n , O r e g o n . M.Sc. t h e s i s , Oregon S t a t e C o l l e g e . 69p. C o o k , C.W. a n d L . E . H a r r i s . . 1950. The n u t r i t i v e v a l u e o f r a n g e f o r a g e as a f f e c t e d by v e g e t a t i o n t y p e , s i t e . and s t a t e o f m a t u r i t y . Utah A g r i c . Expt. S t a . B u l l No. 3 4 4 , 4 5 p . , a n d T . V/. B o x . 1961. A comparison of the loop and p o i n t methods o f a n a l y s i n g v e g e t a t i o n . J . of R a n g e Mgmt. 1 4 ( 1 ) : 2 2 - 2 7 . C o w a n , I . McT. 1945. The e c o l o g i c a l r e l a t i o n s h i p s o f t h e food of the Columbian b l a c k - t a i l e d deer (Odocoileus hemionus columbianus R i c h a r d s o n ) i n t h e c o a s t f o r e s t r e g i o n o f s o u t h e r n Vancouver I s l a n d , B r i t i s h Columbia. E c o l . M o n o g . 15 ( 2 ) : 1 0 9 - 1 3 9 . , W.S. H o a r , a n d J . H a t t e r . 1950. The e f f e c t s of f o r e s t s u c c e s s i o n upon t h e quantity and upon t h e n u t r i t i v e v a l u e s o f woody p l a n t s u s e d by moose. Can. J . Z o o l . 20: 249-271.  90  C o w a n , I . McT., British  and C . J . G u i g u e t . 1965. The mammals o f Columbia. H a n d b o o k No. 1 1 , B.C. P r o v . Mu.s .  D a s m a n n , W.P. and J.A. B l a i s d e l l . 19.54. •. D e e r a n d f o r a g e r e l a t i o n s h i p i n the Lassen-Washoe I n t e r s t a t e w i n t e r deer range. C a l i f . F i s h a n d Game 4 0 ( 3 ) : 2 1 5 - 2 3 4 . •  , a n d R.D. deer census 225-228.  Taber. methods.  1955. A comparison of four C o l o . F i s h a n d Game 4 1 ( 3 ) :  , a n d V/. W. H i n e s . 19.59. Logging, plant success i o n and b l a c k - t a i l e d deer i n t h e redwood r e g i o n . Div. N a t . 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V a n E t t e n . 1956.' Evaluation p e l l e t - g r o u p c o u n t as a d e e r c e n s u s m e t h o d . W i l d l . Mgmt. 2 0 ( 1 ) : 7 0 - 7 4 .  of the J.  E h r e n r e i c h , J . H . a n d D.H. M u r p h y . 1962. A method o f evaluation habitat for forest wildlife'. N.A. Wildl. N a t . R e s . C o n f . 27: 3 7 6 - 3 8 3 . E i n a r s e n , . A.S. . 1946a. Crude p r o t e i n d e t e r m i n a t i o n of deer f o o d s a s 'an a p p l i e d m a n a g e m e n t t e c h n i q u e . Trans. N. Am. W i l d l . C o n f . 1 1 : 3 0 9 - 3 1 2 . 1946b. Management o f b l a c k t a i l e d W i l d l . Mgmt. 1 0 ( 1 ) : 5 4 - 5 9 .  deer.  Halls,  L . K . a n d H.S. C r a w f o r d J r . 1960. Deer-forest habitat r e l a t i o n s h i p s i n north Arkansas. J. Wildl Mgmt. 2 4 ( 4 ) : 3 8 7 - 3 9 5 .  Harper,' J.A. ' 1967. Deer-Roosevelt E l k Relationships. J o b C o m p l e t i o n R e p o r t , P r o j e c t W--59-R-4. Oregon S t a t e . Gam D e p t . 1 2 p . Hazzard,  L.K. 1958. A r e v i e w o f l i t e r a t u r e o n b i g game census methods. C o l o . F i s h a n d Game D e p t . , F e d . A i d P r o g . W-38-R-11.  Holland,  S.E. 1964. Landforms o f B r i t i s h Columbia - A physiographic outline.. B.C. D e p t . o f M i n e s £ P e t r o l e u m R e s o u r c e s , ' B u l l . '48.' 1 3 8 p .  Isaac,  L . A . a n d H.G. H o p k i n s . 1937. The f o r e s t s o i l o f the Douglas f i r r e g i o n , and changes wrought upon i t by l o g g i n g a n d s l a s h b u r n i n g . E c o l o g y 18: 264-279  Klein,  D.R. Ecol.  Krajina,  1965. Monog.  Ecology of deer 35: 259-284.  ranges  i n Alaska.  V.J. 1952. The e c o l o g i c a l c l a s s i f i c a t i o n o f f o r e s t s of t h e e a s t e r n p a r t o f Vancouver Island. M.Sc. t h e s i s , D e p a r t m e n t o f B o t a n y , U n i v e r s i t y o f B r i t i s h Columbia, Vancouver. 112p. . 1959. B i o c l i m a t i c zones i n B r i t i s h B o t a n i c a l S e r i e s #1, D e p a r t m e n t o f B o t a n y , s i t y - o f B r i t i s h Columbia. 12p.  Leopold,  A.S. sion.  1950. Trans.  Columbia. Univer-  Deer i n r e l a t i o n t o p l a n t s u c c e s N. Am. W i l d l . C o n f . 1 5 : 5 7 1 - 5 7 8 .  M c C u l l o u g h , D.R. 1961. An e c o l o g i c a l s t u d y o f t h e C o l u m b i a n b l a c k t a i l e d deer i n a logged environment. M.Sc. t h e s i s , O r e g o n S t a t e U n i v e r s i t y . 57p. M c M i n n , R.G. 1957. Water r e l a t i o n s i n t h e Douglas--fir r e g i o n on V a n c o u v e r I s l a n d . Ph.D. t h e s i s , D e p a r t ment o f B o t a n y , U n i v e r s i t y o f B r i t i s h Columbia, Vancouver, 103p. M u e l l e r - D o m b o i s . D. 1959. The D o u g l a s - f i r f o r e s t a s s o c i a t i o n s on V a n c o u v e r I s l a n d i n t h e i r i n i t i a l s t a g e s of secondary s u c c e s s i o n . Ph.D. t h e s i s , D e p a r t m e n t of Botany, U n i v e r s i t y o f B r i t i s h Columbia, Vanc o u v e r . 570p.  92  Murphy  Odum,  D.A. a n d J A. C o a t e s . 1966. p r o t e i n on d e e r . T r a n s . N.A 129-137. E.P. Co.  1960 546p  Fundamentals  Effects' of dietary Wildl. Conf, 21:  of ecology  W.B.  Saunders  R a s m u s s e n , D . I . a n d E.R. Doman. 1943. Census methods and t h e i r a p p l i c a t i o n s i n t h e management o f mule d e e r . T r a n s . N. A m e r . W i l d l . C o n f . 8: 3 6 9 - 3 7 9 . Rieck,  C.A. 1952. B l a c k - t a i l e d d e e r i n v e s t i g a t i o n s on a c l o s e d area i n western Oregon. M.S. t h e s i s , O r e g o n S t a t e C o l l e g e . 66p.  R o b i n e t t e , W.L., Problems counts. Rogers,  R.B. F e r g u s o n , a n d J.W. G a s h w i l e r . 1958 i n v o l v e d i n t h e use o f deer p e l l e t - g r o u p T r a n s . N.A. W i l d l . C o n f . 2 3 ( 4 ) : 4 1 1 - 4 2 5 .  G., 0. J u l a n d e r a n d W.L. R o b i n e t t e 1958 P e l l e t - g r o u p counts f o r deer census and range-use index.' J . W i l d l . Mgmt. 2 2 ( 2 ) : 193-199.  S e v e r i n g h a u s , C.W. a n d J . E . T a n c k . 1964. Productivity and g r o w t h o f w h i t e t a i l e d d e e r f r o m t h e A d i r o n d a c k R e g i o n o f New Y o r k . N.Y. F i s h a n d Game J . 1 1 ( 1 ) : 13-27. Smith,  I.D. 1968. The e f f e c t s o f h u n t i n g a n d s e r a i succ e s s i o n upon Vancouver Island b l a c k - t a i l e d deer. M.Sc. t h e s i s , Department o f Z o o l o g y , " U n i v e r s i t y of B r i t i s h Columbia, Vancouver. 140p.  Steel,  R. G.D. a n d J . H . S t o r r . i e . 1960. P r i n c i p l e s and p r o c e d u r e s of. s t a t i s t i c s . The M c G r a w - H i l l Book C o . I n c . , New Y o r k . 481p.  Storer  T.I. 1932. i f o r n i a , past  Factors influencing and p r e s e n t . Ecol  w i l d l i f e i n Cal13: 315-334.  S w a n k , W.G.. 1956. N u t r i e n t a n a l y s i s o f c h a p a r r a l browse species. A r i z . Game a n d F i s h D e p t . , W i l d l . R e s t o r . Div. P r o j . W-71-R-3. 15p. . 1956. P r o t e i n and phosphous c o n t e n t o f browse p l a n t s a s an i n f l u e n c e on s o u t h w e s t e r n d e e r h e r d levels. T r a n s . N. A m e r . W i l d l . C o n f . 2 1 : 1 4 1 - 1 5 8 .  93  Taber,  R.D. 1956. Deer n u t r i t i o n and p o p u l a t i o n i c s i n the n o r t h coast range o f C a l i f o r n i a . N. Am. W i l d l . C o n f . 2 1 : 1 5 9 - 1 7 2 .  dynamTrans.-  and R.F, Dasmann. 1958. The b l a c k - t a i l e d d e e r of t h e c h a p a r r a l . C a l i f . D e p t . o f F i s h and' Game, Game B u l l . N o . 8: 1 6 3 p . T h o m a s , D.C. 1963. Reproductive b i o l o g y o f female blackt a i l e d deer ( O d o c o i l e u s hemionus columbianus) o f Northwest Bay, Vancouver I s l a n d . Progress Report o f t h e Game M a n g . R e s . D i v . f o r 1 9 6 3 , B.C. F i s h ^ a n d W i l d l i f e B r a n c h : 18-20.' Ullrey',  D.E., W.G.Youatt, Bradley. 1967. t a i l e d deer fawns.  H.E. J o h n s o n , L . D. Fay-, a n d B . L . Protein requirement of whiteJ . W i l d l . Mgmt. 3 1 ( 4 ) : 6 7 9 - 6 8 5 .  Appendix  A  Temperature and p r e c i p i t a t i o n r e c o r d s t a k e n a t t h e Nanaimo and a v e r a g e s f o r t h e f o u r - y e a r s t u d y p e r i o d .  Airport,  Mean. D a i l y Temp . °F .  Jan .  Feb .  Mar .  Apr .  May  Jun .  Jul.  Aug.  Sep .  . Oct.  Nov .  Dec .  1959  36.6  38 . 2  41. 0  46 . 1  52.0  57.8  65.2  60.8  56.0  48.9  39.9  38.0  1960  35.6  38.4  40 . 5  47 .1  50.7  57.3  65.3  60.9  55.7  50. 6  40 . 9 •37.7  1961  38.8  41. 8  42 . 5  47 . 2  53.2  60.8  66.3  66.2  55.4  47 . 2  39.0  36.4  19 6 2  36.4  39 . 3  38.6  47 . 1  50.5  57.2  61.9 '60.4  57.0  50.2-  43 . 6  39.9  36 . 4  39.4  43 .1  46 . 8  51.6  58.3  64 . 7  56.0.  49 . 2  40 . 8  -38.0  Avera  Precipitat ion (.ins) J a n .  Feb .  Mar .  Apr .  May  Jun .  1959  5 . 84  2.96  4.23  1. 40  1.98  2.56  1960  7.62  5.10-  3.56  3.05  2 .25  .58  .07  1961  9.76  9.93  6.76  1 . 94  2.88  .39  19 6 2  3.32  2.43  3.78  3 . 57  1.99  1.35  4.58  2 . 49  2 . 29  1.22  A v g e . . 6 . en- 5 . 10  62.1  Sep .  Oct .  Nov .  Dec.  • Annual Pptn . (ins. )  2 . 85  2.02  4.53  3.40  •32.68  1.35  .56  3.05  8.67  4 .47 - 4 0 . 3 3  . 83  .95  1.95  4.09  5 . 14  7.76  52.3-8  ,72  2.62  2 .71  5.03  10 . 44  6 . 94  44.90  2.02  3.55  7 . 20  5 . 64  42.57  Jul. . 25  Aug . .66  . . 4 7 1.40  Appendix  B  Seasonal d i e t s of b l a c k t a i l e d deer at Northwest a n a l y s e s o f rumen samples f r o m f e m a l e d e e r .  Species Salal Red c e d a r Arboreal lichens Pearly everlasting Grasses and sedges Trailing blackberry Catsear Mushrooms Douglas f i r Bearberry Red A l d e r Vanilla leaf Hardhack Liverwort Willow Lichen Equisetum Rose Fireweed Moss . Sword f e r n Oregon grape Unidentified  Bay, based  S p r i n g - summer T r a n s - ^ 9 Autumn it ion V o l u m e %| F r e q u e n c y Volume % | F r e q u e n c y . % 5.4 1. 9 7. 3 46 . 0 15 . 1 4. 6 8.. 1 T 2 .3 2.7 T 4 . 4  ' T TV T-  71 57 29 79 57 54 42 8 54 50 12 17  55.9 2. 3 T  . T. T•  4 4  T .  17  Twenty-four  samples,  27 91 36 73 64 • 9 36  1.3  36  1.0 0.8 T T  . 9' 9 27 27  8.0  72.7  May  Winter Volume % l F r e q u e n c y  100 9 9  2.4 7. 3 1. 5 7.0 3.9 1.1 7. 2  8 12 4  %  5 to  30.3 20.9 13.2  -  June-15  samples,  °Thirty-seven  O c t . 2 t o N o v . 27  samples,  D e c . 10 t o M a r c h  25  94 94 49  8. 2 3. 7 3. 9 2. 8 4. 0 5.1 0 . 7  68 73 38 38 81 32 5  2. 5 1. 3  27 32  0. 9  41  T T T 1. 0  2 Eleven  on  • •  16 11 22 35  %  Appendix  Average cover-percentages f o r the major c a t e g o r i e s o f v e g e t a t i o n , t h e r a n k o f d o m i n a n t s p e c i e s , a n d t h e mean n u m b e r o f p a l a t a b l e species recorded i n burned s i t e s o f i n c r e a s i n g s u c c e s s i o n a l age.  C  Years Cover  • •  type  Herbaceous Shrubs and deciduous Coniferous All plants Bare s o i l Dominant  plants trees trees  species:  Total  ferns trees  slash-burning 5 4  2  2.8  7. 1  16 . 6  11.1  12 . 0 0. 7 24 . 8 44 . 2  14.7 0.4 31.6 18.2  27 . 0 0.4 . 33.5: 11 . 7  Salal Ep i l o b i u m spp Thistle Wood groundsel Catsear  Salal Catsear Thistle Epilobium spp Trailing blackberry  0.5 0. 1 3.5 60.9 Wood groundsel Lettuce. Ep i l o b i u m s p p "ThistleSalal  Mean number of P a l a t a b l e Species: Forbs and Shrub s Coniferous  since 3  1  S a l a l •. Catsear Trailing blackberry Pearly everlasting Willow  .6  11. 9  14 . 6  49.9 0. 9 62.7 5.7  38.6 1 . 6 54.8 4. 3  Salal Catsear Raspberry Trailing ' blackberry Pearly everlasting  Salal Catsear Raspberry Trailing blackberry Lettuce  7.0 4 . 0 2.0  7. 0 4.0 1.5  9.0 6.5 1. 5  9. 0 7. 5 1.5'  8. 0 9. 0 2. 0  13 . 0  12 . 5  17 . 0  18 . 0  19 . 0  •11.0 10.. 0 2.0 23.0  CO CD  Appendix  Cover  C  (cont)  8  type  Herbaceous Shrubs and deciduous Coniferous All plants Eare s o i l Dominant  plants trees trees  species:  Mean number of P a l a t a b l e Species: Forbs and Shrub s Coniferous Total  ferns trees ..  9  Years since slash-burning 12 13 , io  7. 2  14 .1  5. 5  10 . 5  53.5 4. 8 65.5 1. 6  23.4 4. 9 42.2 2. 0  59.6 2, 8 66.3 1.4  36.3 5.0 51.7 1.8  Salal Catsear Douglas f ir We stern hemlock Trailing blackberry  11. 0 10.0 2.0,' 23.0  Salal Trailing blackberry Catsear Vanilla leaf Black . raspberry 6. 0 9 . 0' 2.0 17 . 0  14  15  6. 3  7. 0  1. 3  43 . 4 1. 8 51.5 3. 1  30 .1 14 . 4 51.6 1.7  Salal Catsear Douglas f ir Western hemlock Willow  Salal Catsear Trailing black-. berry Willow Oregon grape  Salal Douglas fir Western hemlock Catsear Trailing blackberry  7. 5 9 . 5 2. 0  8.0 9. 0 2. 5  8. 0 11.0 2. 0  8. 0 9.0 3. 0  19 . 0  19 . 5  Salal Catsear Trailing blackberry Douglas fir Western hemlock  21.0  20.0  25 . 7 9. 2 36 . 2 4.8 Salal Douglas fir Oregon grape Red . A l d e r Catsear  4. 0 7.0 3. 0 14.0  Appendix  Cover  C  CCont)  Type  Herbaceous plants Shrubs and d e c i d u o u s Coniferous trees A l l plants Bare s o i l Dominant  species:  Mean number of P a l a t a b l e Species; F o r b s and f e r n s Shrubs Coniferous trees Total  Years since slash-burning Mature Timber  trees  0 . 6 67 . 1 1. 8 69.5 1. 3 Salal Western hemlock Dwarf rose Twin f l o w e r Western red cedar 2.0 8.0 1.0 11.0  CO  Appendix  D  Average cover-percentages f o r the major c a t e g o r i e s of v e g e t a t i o n , arid t h e r a n k ' o f d o m i n a n t s p e c i e ' s , r e c o r d e d I n u n b u r n e d s i t e s ' o f i n c r e a s i n g s u c c e s s i o n a l age.  Years Cover  since  logging  type  Herbaceous Shrubs and deciduous Coniferous A l l plants Bare s o i l Dominant  plants  12.6  trees trees  34 . 3 0 . 5 47 . 4 17.0'  species:  Salal Trailing blackberry Bedstraw Bracken Vanilla . leaf  2.8  4.6  66 .1 0 . 5 69.4 2 .2  68.5 2 . 0 7 5.2 .1.0  Salal Catsear G r a s s es Twinflower Douglas fir  Salal Trailing blackberry Catsear Lettuce Dwarf r o s e  8 . 0 4 4.7 3 . 3 56.1. 8.5'  :  12  13  2. 9  7 . 0  71.4 4,2 78.4 1.9  Salal Salal T r a i l i n g Western blackhemlock berry Douglas Catsear fir Catsear Douglas fir Trailing . blackWillow berry  41.3 4.8 . 53.4 2 . 5 Salal Catsear Douglas f ir Trailin, blackberry Willow  CO CO  100  Appendix  E  N u t r i e n t v a l u e s o f f o u r g r o u p s of. f o r a g e species s a m p l e d i n d i f f e r e n t - a g e d s i t e s and a t different seasons. The a n a l y s e s o f v a r i a n c e f o r . t h e s e data are given i n Table 7.  Age o f Site Cyrs . )  Species Test  1:  F i v e - y e a r - o l d vs  Willow Red  Alder  Red  cedar  Black  raspberry  Arbutus Rose Oregon  grape  Salal Douglas f i r  Test  2:  5 14 5 14 5 14 5 14 5 14 5 14 .5 14 5' 14 5 14  .  F o u r - y e a r - o l d vs December  Willow. Douglas f i r Black  Crude Protein  raspberry  Trailing blackberry  4 9 4 9 4 9 4 9  P e r c e n t Dry W e i g h t Ether Crude Extract Ash Fibre  14-year-old 5 6 7 8 4 .5 7 6 5 6 3 4 8 7 5 4 6 8  .- 8 0 .70 .60 .05 .40 .10 .60 .20 . 60 . 10 . 10 .65 .30 .15 . 30 .90 .30 . 30  7 13 17 17 9 9 4 3 3 5 6 ,3 3 6 11 9 9 15  sites . 50 . 50 .50 .00 .80 . 50 .60 . 30 .30 .15 .30 .40. . 40 . 25 . 30 .65 . 80 .75  n i n e -y e a r - o l d  8 .79 9 .80 12 .09 10 .24 6 .85 5 .64 12. .69 14 . 18  9 7 12 13 8 7 13 11  .32 .73 . 24 .18 . 69 .45 . 98 . 26  in  3 . 89 3. 67 2 . 87 4 . 44 4 . 21 4 . 79 7 . 65 7 . 14  N.F.E.  March  .3 . 00 ' 3 . 80 1. 50 3 . 00 4 .3 0 2. 5 0 . 3 . 50 2 .8 0 2 . 80 ; 3 . 05 8 . 00 • 5 . 15 . 2 . 60 2 .5 5 4'. 50 4 .7 5 2. 8 0 2 .5 0  sites  •  2 5 . 00 58 .70 22 . 50 53 .50 20 . 50 5 2 .90. 21. 5 0 50 .45 20 . 50 .61 .00 2 1 . 70 61 .20 53 . 7 0 •3 0. 60 3 1 . 90 55 .80 10 . 40 77 .90 8. 8 5 76 .85 32 . 90 49 .70 34 . 7 0 52 . 10 28 . 30 ,57 . 40 30 . 15 53 . 9 0 19 . 30 59 . 60 60 .95 19'. 75 61 .00 20 . 10 18 . 4 0 55 . 05  in 28 . 51 30 . 76. 2 1 . 19 19 . 40 35 . 54 35 . 64 1 1 . 87 14 . 56  49 . 49 48 . 05 51 . 61 52 .72 4 4 .71 46 . 48 53 .'8152 .86  101  Appendix  E  Age o f . P e r c e n t ' 'Dry We i g h t ' Site Crude Ether Crude Cyr.s.) ' ' P r o t e i n ' ' E x t r a c t ' A s h ' F i b r e N.F.E.  Species Test  3:  Douglas  Nine-year—old December fir  Trailing blackberry Willow  Test  4:  Willow  Catsear  Fireweed  Bracken  CCont)  vs 14-year-old  9 14 9 14 9 14  F o u r - y e a r - o l d vs s i t e s i n • June 4 9 14. ' 4 9 14 4 9 14 • 4 9 14  10 8 14 12 9 9  . 24 .13 .18 .82 .80 .48  sites  ,13 .18 ' 12 . 18 11 . 26 12 .88 7 .73 8 .42  nine-year- old 15 13 13 14 16 14 15 12 16 34 32 33  .72 . 52 . 51 .50 .86 .78 .39 . 11 .79 .13 .28 .79  11 12 14 17 16 15 12 15 13 14 12 13  .66 .90 .03 .94 . 76 .47 . 23 .46 . 16 .05 .25 .85  4 4 7 4 3 4  vs  in  . 44 . 14 .14 .93 .67 .47  19 ,.40 21.. 24 14 ,. 56 15 ,.34 30 ..76 28 ,.16  52 ,.72 .5 4,.31 52 ,.8 6 54 ,.03 48 ,.05 49 ,, 47  14-;y e a r -- o l d 6 . 2:8 5 . 86 6 .78 8 .62 8 . 86 8 .38 6 .89 6 .88 6 .74 8 .58 8 . 82 . 7 . 44  15 .. 5 0 50 ,.84 15 ,, 36 52 ,.36 16 .. 60 49 ,.08 15 .. 28 43 ,,67 15 ,, 24 42 ,,28 13 ,.50 47 .. 8 7 9 ,.07 56 ,.42. 9 .. 54 56.., 01 .54 .7 . 55 ,.78 34 .,55 8 ..69 10 ,. 12 . 36 ..53 9 ..86 35 .,06  102  Fig.  14a.  A S e n e c i o - E p i l o b i u m a s s o c i a t i o n two y e a r s a f t e r l o g g i n g and s l a s h - b u r n i n g ( P l o t B ) . Wood g r o u n d s e l and w i l l o w h e r b d o m i n a t e , w i t h c a t s e a r and t h i s t l e t h e p r i n c i p a l a s s o c i a t e s . Salal dominates i n unburned spots (upper left).  Fig.  14b.  The same p l o t s i x y e a r s a f t e r s l a s h - b u r n i n g . The a r e a h a s d e v e l o p e d i n t o an i n t e r m e d i a t e Gaultheria-Hypochaeris association, with s a l a l and c a t s e a r r e p l a c i n g t h e groundsel and w i l l o w h e r b .  103  Fig.  15a.  An e a r l y G a u l t h e r i a - H y p o c h a e r i s a s s o c i a t i o n five years after slash-burning ( P l o t G). T h i s t l e s , f i r e w e e d and t r a i l i n g blackberry a r e s u b d o m i n a n t s t o t h e s a l a l and catsear.  Fig.  15b.  The same p l o t a s a n a d v a n c e d GaultheriaHypochaeris a s s o c i a t i o n nine years a f t e r burning. T r a i l i n g b l a c k b e r r y and n a t u r a l Douglas f i r seedlings are the p r i n c i p a l associ ates.  104  Fig.  16a.  An a d v a n c e d G a u l t h e r i a - H y p o c h a e r i s a s s o c i a t i o n n i n e y e a r s a f t e r l o g g i n g and s p o t b u r n i n g (Plot I ) . Trailing blackberry i s subdominant and f o r b c o v e r i s l i g h t . Note the d e b r i s .  Fig.  16b.  The same p l o t a s a n e a r l y G a u l t h e r i a P s e u d o t s u g a a s s o c i a t i o n 13 y e a r s a f t e r d e forestation. Red a l d e r , t r a i l i n g blackb e r r y a n d w i l l o w a r e now t h e p r i n c i p a l associates.  Fig.  17a.  An a d v a n c e d G a u l t h e r i a - H y p o c h a e r i s a s s o c i a t i o n 12 y e a r s a f t e r l o g g i n g a n d s p o t b u r n i n g (Plot K) Fireweed i s abundant i n the foreground.  Fig.  17b.  P l o t K as a G a u l t h e r i a - P s e u d o t s u g a association 16 y e a r s a f t e r l o g g i n g a n d b u r n i n g . Douglas f i r has e n c r o a c h e d s u f f i c i e n t l y t o r e p l a c e most h e r b a c e o u s s p e c i e s .  

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