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Functional relationships between salal understory and forest overstory Vales, David Joseph 1985

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FUNCTIONAL  RELATIONSHIPS  BETWEEN  S A L A L UNDERSTORY AND  FOREST  OVERSTORY  By DAVID B.Sc,  Iowa  JOSEPH  State  A T H E S I S SUBMITTED THE  VALES  University,  1981  IN PARTIAL FULFILLMENT  REQUIREMENTS MASTER  FOR  OF  THE  DEGREE  OF  OF  SCIENCE  in THE  FACULTY  OF  (Faculty  We  accept to  THE  this  GRADUATE of  Forestry)  t h e s i s as  the required  UNIVERSITY  OF  October  © David  Joseph  STUDIES  conforming  standard  BRITISH  COLUMBIA  1986  Vales,  1986  9  In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department  or  by  his or  her representatives.  It  is understood  that copying  or  publication of this thesis for financial gain shall not be allowed without my written permission.  Department of The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date  DE-6(3/81)  H.  /O  ABSTRACT  Abundance relationship forest  to forest  stands  strongly  cover  to single  Most  d i d not d i f f e r  calculated  from  variation  equations  overstory  Mean  cover  photosynthetically  The  forest  stand  components. diffuse  radiation  data shoot  of g l o b a l , active on  transmitted  stand by open  below below  Plots  sampled  over  salal  density  was  characteristics  differed  stands (r  2  between  salal  z  plant  Equations  accounted  several  O  a  biomass or  f o r more o  = 0.73-0.97)  height  direct,  12 p l o t s  The r e l a t i o n s h i p  among  topographical  associations.  from  solar  characteristics  scattered  extended  single  of radiation  radiation  differed was  from  was s t u d i e d  proportions  plant  i n immature  some  predicting  abundance  salal  stand  i t s  stands  than ( r  was g r e a t e s t  2  =  under  o f 65 t o 8 0 % .  Transmission  canopies  from  in salal  developed  0.39-0.92).  between  data  that  equations  Pursh) and  studied  ranges.  forest  equations  on  having  indicated  but predictive  associations.  were  Island  winter  levels  related  0.92-0.94)  the  of deer  of stocking  shall  overstory  on V a n c o u v e r  characteristics range  (Gaultheria  of s a l a l  the height the canopy  radiation during  sunny were  and d i f f e r e d between  of short  of radiation was h i g h e r  forest  days a  among  i n summer  function  of  radiation  the proportion of  and stand When  and d i f f u s e  through  transmitted  structures. crowns  diffuse,  characteristics direct  trees  beam  and crowns  sensors, than  radiation  diffuse  outside.  The  extent  to  which  characteristics  predict  were  Sum  explored.  density and  index  direct  predicts when  predict  obtained  =  of  i s extensive above  by  Differences of  of  of  radiation.  Photographic  transmitted  differed  and  (i  best.  Overstory  stand  methods  =  (i  and  0.80-0.95)  2  =  bases  cover 0.74) of  tree  characteristics gave  different  transmitted.  between  diffuse  estimates  from  2  law  stand  global  hemispherical  radiation transmitted  diffuse  Beer's  Reineke's  variation  components  found  follow  r a d i a t i o n best  Forest  sampling  were  of  inter-plot  the  diffuse  transmission  diffuse  stand  and  0.70-0.98)  sensors.  different  predictabilities  estimates  2  and  diameters  transmission  transmission  are  forest  transmission tree  (i  radiation  there  crowns  of  different  measured  photosynthetically of  measured  and  photograph  direct  active  radiation  transmission  of  characteristics  to  direct  radiation. The  r e l a t i o n s h i p s of  transmission  of  solar  Salal  density,  cover  a l l increased  transmission. transmission global,  basal  The of  diffuse,  r a d i a t i o n components area,  growth  direct or  fastest  transmission  low  to  shoot  moderate  rate  diffuse at  height  with  and  proportions  was 2  {i  salal  direct basal of  examined.  =  and  increasing more  closely  0.65-0.99)  photosynthetically  which of  were  p r o d u c t i v i t y , biomass,  of-salal  radiation  The  Salal  foliar  asymptotically  radiation. for  salal  abundance  related  than  to  active plateaued  was  radiation. diameter  were  largest  radiation transmitted.  for The  to  lowest  and g r e a t e s t  adversely  affect  productivity transmission. regulated a  site  by  proportions  salal  increased Salal  height  by a n  with  abundance and  of d i r e c t  solar  interaction  regime.  iv  seemed  to  and diameter.  asymptotically  maximum  t h e amount  and p r o b a b l y  shoot  transmitted  increased shoot  radiation  with  Shoot  a  size  are  received  site's  at  moisture  TABLE OF  CONTENTS  ABSTRACT  i i  TABLE OF CONTENTS  v  L I S T OF TABLES  viii X  L I S T OF FIGURES L I S T OF APPENDICES  xiii  ACKNOWLEDGMENTS  xvi  CHAPTER  1.  GENERAL  INTRODUCTION  1  REFERENCES CHAPTER 2.  3 RELATIONSHIPS BETWEEN  SALAL UNDERSTORY  AND  FOREST OVERSTORY  4  INTRODUCTION  4  STUDY AREAS  6  METHODS  10  Understory  10  Overstory  12  Definition  of v a r i a b l e s  13  Analyses  13  RESULTS  16  Regressions  predicting  salal  abundances  16  Form o f r e l a t i o n s h i p s  19  Differences  20  Relative Salal  among r e l a t i o n s h i p s  predictability  shoot  among  height  independent v a r i a b l e s  ....23 29  DISCUSSION  32  CONCLUSION  40  v  REFERENCES CHAPTER  3.  41 R E L A T I O N S H I P S EETWEEN  RADIATION  AND  FOREST  TRANSMISSION  OF  SOLAR  STAND C H A R A C T E R I S T I C S  45  INTRODUCTION  45  STUDY A R E A S  47  METHODS  49  Overstory Solar  sampling  radiation  Definition  49  sampling  50  of v a r i a b l e s  52  Analyses RESULTS  53  AND  DISCUSSION  Proportions  transmitted  Radiation-forest Equation Site  55  stand  55 relationships  59  differences  67  factors  Generality  67  of equations  71  CONCLUSION  76  REFERENCES  78  CHAPTER  4.  R E L A T I O N S H I P S OF  TO T R A N S M I S S I O N  OF  SOLAR  SALAL  (GAULTHERIA  R A D I A T I O N THROUGH  SHALLON) FOREST  CANOPIES  83  INTRODUCTION  83  STUDY A R E A S  86  METHODS  89  RESULTS  92  Form Salal  of response shoot  94  size  99  vi  Subzone Relative Effects  differences  102  predictability of r a d i a t i o n  among  radiation  components  on s a l a l  components  ....104 105  DISCUSSION  106  CONCLUSION  112  REFERENCES  113  CHAPTER  5.  MANAGEMENT  IMPLICATIONS  INTRODUCTION FOREST  MANAGEMENT  117 117  IMPLICATIONS  W I L D L I F E MANAGEMENT  IMPLICATIONS  REFERENCES  118 123 129  APPENDICES  133  vii  LIST  2.1  Descriptions  stand  2.2 a  used  TABLES  t o sample  salal  and  forest  characteristics  Simple  linear  function  CWHa=lO, 0.05.  2.3  of plots  OF  regression  o f MCC, p l o t ,  CWHb^S,  Regression  Multiple  function  9  equations  and prism  variables.  CWHb = 7.  Equations  equation:  Y = b  3  of salal  equations  of salal  o f MCC, p l o t ,  and prism  variables.  a t p r< 0 . 0 5 .  Sample  as  sizes:  at p <  fc,X  regression  significant  Sample  significant  +  0  variables  sizes:  17  variables  as a  Equations  CWHa=lO,  CWHb,=5,  CWHb =7  18  3  3.1  Descriptions  forest  stand  3.2  Daily  sums  3.3  Regression  radiation  of p l o t s  of solar  components  0.05.  Regression  predicting  solar  radiation  and 48  radiation  coefficients  sizes:  Indices  t o sample  characteristics...  Sample  3.4  used  from  of equations forest  CWHa=6, CWHb=6. equation:  of determination radiation  characteristics  using  components  stand  by p l o t  predicting  significant  Y = ae  2  components  from from  the regression  viii  solar  characteristics.  Equations  (i )  56  at p £ 60  regressions forest form  stand  Y = e  63  3.5  Comparison  from solar  4.1  of diffuse  hemispherical radiation  and d i r e c t  photographs  components  Descriptions of plots  site  factors  t o measured  obtained  proportions of  transmitted  used  t o sample  68  salal  and solar  radiation  4.2  Plot  88  means  a n d 95% c o n f i d e n c e  intervals  for salal  characteristics  4.3  Below-canopy  4.4  Regression  variables CWHa=6,  Tree  density  solar  from  solar  densities  statistics  of equations  radiation  Equations  equation:  index  radiation  coefficients  CWHb=6.  Regression  5.1  93  Y = (a(X-c))/(1  needed  as average  to maintain diameter  ix  predicting  components.  significant  94  Sample  salal sizes:  at p < 0 . 0 6 .  + a(X-c)/b)  a constant  changes  .....95  stand 120  LIST  2.1  Plot  U.B.C.  2.2  locations Research  FIGURES  on V a n c o u v e r  Island  and a t  the  Forest  The r e l a t i o n s h i p  intervals  OF  t o mean  7  of s a l a l  crown  density-irr  completeness.  CWHa a n d CWHb, c o m b i n e d  data  2  and 95% c o n f i d e n c e  Lower  (•), upper  equation  equation  CWHb  i s 3  (ffl)  2.3  24  The r e l a t i o n s h i p  and  95% c o n f i d e n c e  CWHa  ( • ) , CWHb,  of f o l i a r intervals  ( A ) , CWHb  3  2  productivity t o mean  (ffl).  (g-nr ;  crown  Equation  CAGBIOM)  completeness. i s f o r combined  data  2.4  25  The r e l a t i o n s h i p  of s a l a l  FOLBIOM) a n d 9 5 % c o n f i d e n c e completeness. represents  2.5  CWHa  ( • ) , CWHb,  combined  intervals  combined  ( A ) , CWHb  t o mean  ( A ) , CWHb  3  crown  (ffl).  Equation  3  total  biomass  t o mean  (ffl).  crown  Equation  2  (g-nr ;  TOTBIOM)  completeness. represents  data  27  The r e l a t i o n s h i p  confidence CWHb,  (g-nr ;  26  95% c o n f i d e n c e ( • ) , CWHb,  2  biomass  data  of salal  CWHa  foliar  intervals  The r e l a t i o n s h i p  and  2.6  for  of percent  intervals  ( A ) , CWHb  3  (ffl).  t o mean  cover  crown  Equation  data  of s a l a l  and 95%  completeness.  represents  CWHa ( • ) ,  combined 28  x  2.7  The r e l a t i o n s h i p  confidence (b)  3.1  Open and  3.2  intervals  stretch.  Solar  diffuse  crown  ( • ) , CWHb,  completeness  (•••••••),  between  line  the proportions  a n d mean  crown  i n a i s from  of salal  intervals  lines  basal  area  PPFD s o l a r  (o)  radiation  completeness.  Dotted  for individual (1959).  CWHa  (cm «m~ )  a n d 95%  of global,  direct,  2  radiation  2  components.  i n individual  subzones.  direct,  components.  of salal  diffuse,  Dotted CWHa  CWHa ....96  (ffl)  foliar  C A G B I O M ) a n d 95% c o n f i d e n c e  subzones.  global  of solar  Miller  to transmission  are relationships  Relationships  global,  canopy  62  and d i f f u s e  CWHb  for plot 27.  (ffl)  confidence  Dotted  30  59  Dashed  Relationships  diffuse,  (a) repose,  (ffi)  3  of time  i n a, b, a n d d a r e r e l a t i o n s h i p s  (•) , -CWHb  a n d 95%  of s a l a l  ( A ) , CWHb  as a function  transmitted  subzones.  4.2  height  (•)  Relationships  (•),  t o mean  (•) a n d d i f f u s e  components  4.1  CWHa  irradiance  global  lines  of average  lines  ( • ) , CWHb  intervals  and d i f f u s e  xi  (g»m~ ;  to transmission of PPFD s o l a r  are relationships (ffl)  2  productivity  radiation  in individual 97  4.3  Relationships  and  95% confidence  direct,  diffuse,  components. subzones.  4.4  of s a l a l  Relationships  diameter  intervals  and d i f f u s e  Dotted CWHa  lines  ( • ) , CWHb  of global  biomass  2  (g«nr ;  to transmission PPFD s o l a r  TOTBIOM)  of global,  radiation  are relationships  in individual  (ffl)  of average  (cm) o f s a l a l  transmission  total  with  shoot  98  height  (cm) a n d b a s a l  95% c o n f i d e n c e  and d i r e c t  intervals to  radiation.  CWHa  ( • ) , CWHb 100  (ffl)  4.5  Relationships  transmission (•),  CWHb  of average  of global  productivity  and d i r e c t  (ffi)  solar  of s a l a l radiation.  shoots to CWHa 102  xii  LIST  1  Forest  stand  2  Means a n d 95% c o n f i d e n c e  characteristics  characteristics  3  Correlation CWHa  4  6  3  coefficients  variables f o r  among  overstory  variables for  among  overstory  variables f o r 141  coefficients  variants  Allometric  Average  overstory  variables  for a l l 142  f o rstanding  crop  foliar  and t o t a l  of s a l a l  biomass  areas  CWHb,=5,  among  (n=22)  equations  143  ( i n grams)  intervals  Regressions  of salal and stand  CWHb =7.  Regression  3  95% c o n f i d e n c e  of salal  (backtransformed  variables  Y = b  of average intervals  from  Sample  significant +  0  leaves  against  densities.  Equations  equation:  The r e l a t i o n s h i p and  overstory  140  Correlation  basal  10  among  (n=7)  confidence  9  136  («=5)  biomass  8  f o r understory  139  coefficients  Correlation  three  7  intervals  133  of plots  coefficients  Correlation  CWHb  of plots.  (H=10)  CWHb,  5  OF A P P E N D I C E S  by p l o t square  plot  root)..146  and prism  sizes: at p  CWHa=lO,  < 0.05.  fc,X  basal  diameter  t o mean  xi ii  crown  and 95%  of salal  147  shoots  completeness..148  11  The r e l a t i o n s h i p 95%  confidence  Equation:  s  of twig  intervals  #CAGtwigs»m-  2  productivity t o mean  (# CAG t w i g s ) a n d  crown  = 193.81  completeness. 2  - 219.93-MCC  r =0.9l,  =10.7  149  yx 12  13  The r e l a t i o n s h i p  of salal  basal  confidence  intervals  Basal  = 2 0 . 5 7 - 22.80-MCC  area  The r e l a t i o n s h i p  t o mean  of s a l a l  area  crown  2  completeness.  2  r = 0.76,  shoot  2  (cm /m ) and 95%  posture  s  Equation:  =2.0  t o mean  150  crown  completeness  14  151  Distribution Distribution  of repose  heights  i s percent  of s a l a l  of shoot  density  shoots i n each  by  plot.  height  class  15  154  Percent  frequency  of occurrence  of species  found i n  quadrats  16  Average other  155  quadrat  than  List  18  Results  19  Regression  sizes:  of cover  of understory  species 156  classes  of site  f o rspecies  present  in plots.  ....157  diagnosis  coefficients  components  equation:  158  of equations  from  CWHa=6, CWHb=6.  Regression  density  salal  17  radiation  2  (0.25 m )  stand  characteristics.  Equations  Y = ae  predicting  significant  solar Sample  a t p ^ 0.10. 159  xiv  20  Regression radiation sizes:  components  equation:  Regression site  Y  =  derived  +  diffuse  relationships  solar  Sample  a t p £ 0.05.  significant  160  predicting  hemispherical  Regression  salal  photographs.  from Sample  equation:  a(X-c)/b)  161  density  to transmission  PPFD  solar  characteristics.  of equations  from  22 R e l a t i o n s h i p s o f s a l a l intervals  predicting  Y = e  CWHa=5, CWHb=6.  (C(X-C))/(1  stand  Equations  coefficients  factors  sizes:  of equations  from  CWHa=6, CWHb=6.  Regression  21  coefficients  2  ( # » n r ) and 95% c o n f i d e n c e  of g l o b a l ,  radiation  .direct,  components.  in individual  subzones.  d i f f u s e , and  Dotted  CWHa  lines are  ( • ) , CWHb  (ffl)  23  1 62  Relationships of percent intervals diffuse  to transmission  PPFD  relationships  solar  cover  of salal  of global,  radiation  i n individual  a n d 95% c o n f i d e n c e  direct,  components. subzones.  d i f f u s e , and  Dotted  CWHa  lines are  ( • ) , CWHb  (ffl)  163  24 R e l a t i o n s h i p s o f s a l a l and  PCTCOVER  D E N S I T Y , CAGBIOM,  t o Reineke's  SDI d e r i v e d  measurements  (SDI) and prism  are  relationships  separate  samples  i n CWHa  xv  from  FOLBIOM, plot  (BAFSDI).  a n d CWHb  3  TOTBIOM,  tree Dotted  lines  variants....164  ACKNOWLEDGMENTS  I  find  i t difficult  persons  have  of  project.  this  grandfather F.L.  made  who  Bunnell,  To  thanks  begin,  f o r development  learned  to  member  penetrating equipment B.C. and  loans  helped  and  of  always  thanks  technicians)  R.  go  to  field  McCann,  A.  field  under  trying  to  answer  my  some  my  thanks  go  questions  him  I  the have  researcher. thinking  from  with  for  the  essential  for  this  m e m b e r s M.D.  assistants  many  to  and  study  Pitt  and  provided  and  of  long,  good Black  also  solar  S.  (and  Jay,  xvi  D.  of  Hart  and  Your  frustrating  appreciated. finding  suggestions the was  for  probing  were  for always  radiation  study.  and  nature  for his  Because  Dexion  boring,  conditions.  T.A.  measurements.  Korol, monitoring  and  and  arranged  questions  r e s o u r c e f u l n e s s , and  meteorological R.  were  McLeod,  with  Special  also  many  insight.  p u t t i n g up  questions,  of  independent  Committee  graciously days  parents  funding,  developing  which  welcomed  and  supervisor,  Nyberg  photograph  costs.  My  Because  when  development  t o my  s t i m u l a t e d my  J.B.  Forests  comments  Many  Nyberg  the  go  data,  ideas.  confident,  J.B.  minimize  Smith  valuable  of  questions.  Ministry  J.H.G.  more  support.  background  impetus  Committee  acknowledgments  contributions to  provided  a  order  important  provided  be  to  on  suggestions included  time  in  of this  A  U.B.C.  Foresty the  Teaching  three  Council  1984  Career  graciously through  Access  Thanks involved foresters  who  were:  Products;  assistant  support  me  J . Kapitany  K. A l l e n ,  Harwijne  a n d R.  the friendly  with  Products  by  Housing  i n Woss  I am  maps  a n d M.  grateful  and t h e i r  to a l l of the  time.  Canadian  a n d A. W a l k e r ,  and others  Forest  British  B. G i l l m o r e , B.  VanBruen, Crown  Some o f  MacMillan  Kurtz, Bloedel;  F o r e s t ; R. M e h ,  at the Mesachie  Lake  Columbia  R.  Research  Miller,  Station,  of Forests.  Final  thanks  a r e extended  discussions provided study  had been  was  of Forests a t the  C. R a y , R. E l l i s  P. A f f l e c k ,  Thurlborn,  staff  Science  funded  '85 g r a n t s .  Forest  during  through  was p a r t l y  and E. Mulock,  B. M u r p h y ,  Ltd.;  financially  Station.  provided  J.  this  by t h e B.C.  t h e IWIFR p r o g r a m .  L o f t u s , B. S h u c k l e ,  whose  supported  a n d B.C. M i n i s t r y  Research  Products  Ministry  me  by C a n a d i a n  J.  and  supported  Service grants  go t o L . P e t e r s o n ,  with  F e l l o w s h i p and F a c u l t y of  a n d 1985 C h a l l e n g e  provided  Lake  was  Forest  Field  J . Kapitany  Mesachie  Forest  Research  and Canadian  Bunnell.  Graduate  Assistantship  years.  F.L.  those  University  to fellow  a much  coursework  richer  and f i e l d  xvi i  graduate education research  students, than i f alone.  1  CHAPTER  Manipulating technique domestic stands  t o encourage stock  overstory  canopy the  (e.g.,  affect  1986).  1976),  also  Maintaining  researchers  considered (e.g.,  studies  of this  a r e : 1) t o e x a m i n e Pursh) density,  height  differences  have  to forest  to manipulation been  and B e t t e r s with  trees  stands  of the forest drawn  between  1983).  nutrients  studied  f o r moisture  (e.g.,  Weetman e t  t o shade  outthe  (Black  potential  Anderson  have  e t a l . 1980;  factors regulating  e t a l . 1969; Z a v i t k o v s k i  had mixed  results.  and understory  of transmission  study  Causal  have  of solar  are treated  often  been  radiation  i n separate  foliar  stand  p r o d u c t i v i t y , biomass,  characteristics  of transmission  chapters (Gaultheria  the relationships of salal  i n t h e r e l a t i o n s h i p s among  relationships  examined t h e  1982).  Objectives  on  Studies  forest  overstory  as functions  Alaback  how c h a n g e s i n  competition  (e.g.,  between  forest  1981).  have  production  and these  T o manage  t o know  competes  dense  can reduce t h i s  relationships  and  e t a l . 1967).  see B a r t l e t t  as a  f o r w i l d l i f e and  and r e l a t i o n s h i p s have  and S p i t t l e h o u s e  understory  suggested  production  T a n e t a l . 1977) a n d l i k e l y  Few  shall  Young  vegetation  understory  and  forage  INTRODUCTION  h a s been  (understory)  two ( f o r r e v i e w  Black  stands  the understory.  (overstory)  (e.g.,  GENERAL  i t i s important  of forage  Understory  al.  forest  f o r forage  response  1.  cover,  and t o test f o r  sites;  2) t o s t u d y t h e  of global, d i r e c t ,  d i f f u s e , and  2  diffuse  photosynthetically  stand  characteristics;  salal  abundance,  3)  growth  transmission  of  and  management  wildlife  Emphasis  in this  immature  stands  Vancouver limited  work  and  shoot  previously  i s on  done  on  at  related  of  these  forest which  to the  forest  findings.  practices  in  Hemlock  Zone  on  will  supplement  this  study  untended,  understory-overstory  mature,  Group  to  to discuss  Western from  are  to  4)  forestry  results  in clearcut,  the F o r e s t r y - W i l d i f e  and  radiation  the extent  size  implications  i n the Coastal The  solar  to evaluate  radiation;  study  Island.  relationships by  solar  active  and  U.B.C.  old-growth  forests  done  3  REFERENCES  A l a b a c k , P.B. 1982. D y n a m i c s o f u n d e r s t o r y b i o m a s s i n S i t k a spruce-western hemlock f o r e s t s of s o u t h e a s t Alaska. Ecology. 63:1932-1948. Anderson, R.C, O.L. L o u c k s , a n d A.M. S w a i n . 1969. Herbaceous response t o canopy c o v e r , l i g h t i n t e n s i t y , and t h r o u g h f a l l p r e c i p i t a t i o n in coniferous forests. Ecology. 50:255-263. B a r t l e t t , E . T . a n d D.R. Betters (eds.). 1983. Overstory-understory r e l a t i o n s h i p s in western forests. W e s t e r n R e g i o n a l R e s e a r c h P u b l . No. 1. C o l o r a d o State U n i v . Exp. S t a . , F o r t C o l l i n s , CO. B l a c k , T.A. a n d D.L. S p i t t l e h o u s e . 1981. M o d e l i n g the water b a l a n c e f o r w a t e r s h e d management, pp. 117-129 in: D.M. B a u m g a r t n e r ( e d . ) . P r o c . I n t e r i o r West W a t e r s h e d Mgt. A p r i l 8-10, 1980. Spokane, WA. B l a c k , T.A., CS. T a n , a n d J . U . Nnyamah. 1980. T r a n s p i r a t i o n i n t h i n n e d and u n t h i n n e d s t a n d s . Can. J . S o i l S c i . 60:625-631. Tan,  C.S., T.A. B l a c k , a n d J . U . Nnyamah. 1977. Characteristics of s t o m a t a l d i f f u s i o n r e s i s t a n c e i n a D o u g l a s - f i r f o r e s t e x p o s e d t o s o i l water d e f i c i t s . Can. J . F o r . Res. 7:595-604.  W e e t m a n , G.F., A. G e r m a i n , a n d R. F o u r n i e r . 1 9 8 6 . Fertilizer s c r e e n i n g t r i a l s o f s t a g n a t e d S i t k a s p r u c e p l a n t a t i o n s on n o r t h e r n V a n c o u v e r I s l a n d , B.C. S o i l S c i . S o c . Am. J. (submitted). Young, J.A., D.W. H e d r i c k , and c o v e r and l o g g i n g - h e r b a g e mixed c o n i f e r o u s f o r e s t of 65:807-813.  R.F. K e n i s t o n . 1967. Forest and browse p r o d u c t i o n i n t h e northeastern Oregon. J . For.  Z a v i t k o v s k i , J . 1976. G r o u n d v e g e t a t i o n b i o m a s s , p r o d u c t i o n , a n d e f f i c i e n c y o f e n e r g y u t i l i z a t i o n i n some n o r t h e r n Wisconsin forest ecosystems. Ecology. 57:694-706.  4  CHAPTER  2.  RELATIONSHIPS  BETWEEN  SALAL  UNDERSTORY AND  FOREST  OVERSTORY  INTRODUCTION  In  the coastal  {Gaul theria  salal  component tree and is  shall  winter  (Odocoileus 1961;  nutrients  Crouch  foresters  abundance  Spittlehouse  1981),  regimes  might  that  1986).  abundance  In each to forest  than  t h e work  Koch  (1983),  are  not well  plant  anus  Richardson;  dense  commercial e t a l . 1986)  Second,  black-tailed  1975; R o c h e l l e  to maintain  Cowan  1980).  forest  salal  deer 1945; Brown  While  stands  t o reduce  ( B l a c k e t a l . 1980; B l a c k a n d  wildlife encourage case  managers salal  documented  forage  characteristics  and Turner  and Bunnell  are considering  (1975),  and Vales  (Nyberg e t  of salal  growth and  are c r i t i c a l . Stanek  (1986),  nor a r e they  thinning  abundance  the relationships  stand  o f Long  understory  with  e t a l . 1986).  f o rColumbian  columbi  of s a l a l  I t competes  Northwest  ( T a n e t a l . 1977; P r i c e  (Weetman  1968; J o n e s  may w a n t  of the P a c i f i c  i s an i m p o r t a n t  reasons.  forage  hemionus  o f much  Pursh)  s p e c i e s f o rmoisture  a major  al.  on  f o r two m a j o r  possibly  the  forests  tested  et a l .  these  Other (1979),  relationships  for different  associations. This  forest develop  study  stand  examines  the relationships  characteristics.  regressions predicting  Specific salal  between  objectives  density,  s a l a l and were  to:  productivity,  1)  5  biomass,  and  cover  from  identify  the  forms  of  variables  and  differences (Klinka  et  forest  i n the  1965)  identify  the  predicting  and  best  salal  relationship  of  the  stand  stand  and  plant forest  between  characteristics;  between  among  stand  height  and to  3)  examine  forest  stand  1965);  t o use  the  variants  subzones  1964,  characteristics 5)  for  biogeoclimatic  (Orloci  2)  salal test  biogeoclimatic  associations  abundance; salal  characteristics;  relationships  relationships  a l . 1984)  (Krajina  forest  4)  for  qualitative  characteristics.  6  STUDY  Twenty-two the  University  (Fig.  2.1).  Hemlock 1964, al.  were  of B r i t i s h  Ten p l o t s  - Douglas-fir  plant  1965).  Four  association  Coastal  Western  (G-WH-DF) p l a n t Island  Western  plots  also  Hemlock  by l e s s  Vaccinium  -  than  Gaul  theria  CWHb,.  was c o m b i n e d  plots  for analyses  moisture-nutrient Seven  additional  association Western  because regime  plots  with  sampled  Wet S u b z o n e  3  were  separated  by l e s s  than  All  (except  t h e UBCRF  i n immature  menziesii heterophylla  (Mirb.)  -  Douglas-fir the  Island  CWHb,  soil Klinka  1976).  plant  of the Coastal  seven  i n t h e same  CWHb, p l o t  containing  Sarg.).  a n d were  was l o c a t e d i n  variant These  plots  CWHb  3  plots  stand, and  8 0 0 m.  Other  i n o l d growth)  Douglas-fir  Franco) and western  (Raf.)  four  1964, 1965) i n  o n t h e same h i l l s i d e ,  stands  oft h e  i n t h e V-G-WH-DF  (CWHb ).  a l l located  were  plot  i t had a s i m i l a r  were  plots  stand  t h e Vancouver  i n t h e l e e w a r d submontane  Hemlock  These  (hygrotope-trophotope;  were  (CWHa;  variant  Hemlock  (Orloci  Klinka et  i n t h e G-WH-DF  (CWHb,).  - Western  association  plot  sampled  O n e UBCRF  (Orloci  D r y Subzone  i n t h e same  8 0 0 m.  UBCRF  (CWHa,;  submontane  Wet S u b z o n e  (V-G-WH-DF) p l a n t This  variant  were  and a t  - Western  association  Hemlock  i n t h e windward  Island  Research Forest,  i n t h e Gaultheria  a l l o n t h e same h i l l s i d e  separated  on Vancouver  Columbia  were  1984) o f t h e C o a s t a l  were  sampled  1965) i n t h e V a n c o u v e r  Krajina  the  plots  AREAS  hemlock  tree  species  {Pseudot  suga  (Tsuga encountered  Figure  2.1.  Plot  locations  on  Vancouver  Island  and  at  t h e U.B.C. R e s e a r c h  Forest.  were  western  (Abies  redcedar  amabilis  nootkatensis  Dougl.).  with  Oregon  (Klinka  examined, plots. annual  were  L F H a n d Ae l a y e r  CWHb,, a n d CWHb  3  moderately  medium. poor  t o medium.  developed  present. or  colluvial  d i f f e r e d most on most  among  plots. mm  Average in  d r y and nutrient dry to fresh  of plots  the  e t a l . 1984).  f o r t h e CWHa a n d CWHb  slightly  Characteristics  Smith) and  characteristics  t h e two ( K l i n k a  dry to slightly were  was p r e d o m i n a n t l y  i n t h e CWHa, 2 6 8 2  combinations  s  (Pinus  pine  on m o r a i n a l  Of t h e s o i l  i s 2 1 2 3 mm  white  commonly  development  i s between  CWHb, p l o t s  (Chamaecypari  parvifolium  Pursh) Podzols  was p o o r l y  Hygrotope-trophotope were  nervosa  e t a l . 1984).  precipitation  vegetation  (Vaccinium  Humo-Ferric  M o r humus  and western  Understory  (Mahonia  a Donn), a m a b i l i s f i r  Forbes), yellow-cedar  redhuckleberry  grape  Soils veneer  (Dougl.)  piicat  (D. Don) S p a c h ) ,  monticola salal  (Thuja  3  plots  poor t o and nutrient  are listed  i n Table  TABLE 2.1  Plot  D e s c r i p t i o n s of p l o t s used to sample s a l a l and f o r e s t stand c h a r a c t e r i s t i c s  Location name  Lat1tude-- Long 1tude Subzone var1 ant  Plant assoc.  Aspect  Elev. (m)  Slope  C )  Age  MCC  a  (%)  NTREEs" BA Avg. DBH (#/ha)(m'/ha) (mm) b  b  Avg. Ht (m)  C  S1te 1 ndex  1  Woss  50' 12' -- 126"28'  CWHa  G-WH-DF  230  430  30  28  0 91  2533  37 .7  131  12 . 4  36.4  2  Brewster 1  50' 07 ' -- 125*39'  CWHa  G-WH-DF  215  270  15  37  0. 78  1600  46 . 7  173  13 6  38.4  3  Brewster 2  50'07' •• 125'39'  CWHa  G-WH-DF  220  260  55  36  0. 55  533  21 .2  213  13 6  32.2  4  Mcrelght  50' 15 ' -- 125'37'  CWHa  G-WH-DF  218  150  48  53  0. 79  1778  53 .4  178  16 1  29.5  5  Youbou 1  48'54' •• 124'18'  CWHa  G-WH-OF  206  310  52  60  0 64  756  35 .0  235  25. 2  42.9  6  Youbou 2  48'54' •- 124'18'  CWHa  G-WH-DF  215  310  45  59  0 78  1733  34 .4  145  15. 6  35.3  7  Youbou 3  48'54' •- 124'18'  CWHa  G-WH-DF  173  200  9  34  0 35  267  14 . 8  219  15. 7  36.5  8  UBCRF M-G  49' 17 ' •• 122'28'  CWHa  G-WH-DF  115  375  10  44  0. 65  667  30 .7  222  18. 1  40. 4  9  Gordon R.  48'49' -- 124"19'  CWHa  G-WH-DF  261  630  43  42  0..90  2711  54 .9  149  15 .0  36.0  Raymond Cr.  48'52' -• 124'28'  CWHa  G-WH-DF  210  390  60  39  0, 75  1200  32 .5  165  15..5  39.7  49' 17 ' •- 122'28'  CWHb.  V-G-WH-DF  237  590  25  150+  0. 90  1244  61 .0  210  15. 6  30.0  12  H a r r i s Cr 1 48'37' -• 124' 15'  CWHb,  G-WH-DF  215  210  50  49  0. 64  844  25 .5  179  12.8  26.5  13  H a r r i s Cr 2  48'37' -• 124' 15'  CWHb.  G-WH-DF  221  200  40  50  0. 80  2000  62 .2  185  17. 6  36.5  14  H a r r i s Cr 3  48'37' -- 124' 15'  CWHbi  G-WH-DF  232  200  50  48  0. 84  2444  67 .6  171  17 4  35.0  15  H a r r i s Cr 4 48'37' •• 124" 15'  CWHb,  G-WH-DF  204  210  48  49  0. 79  2089  62 .0  182  13 9  27.9  889  10 . 1  110  9.7  37.8  10  1 1 UBCRF V-G  CWHbi  V-G-WH-DF  240  570  30  30  0. 58  48'36' -• 123'52'  CWHbi  V-G-WH-DF  240  570  30  30  0. 42  756  a .5  118  10. 3  37 .0  Weeks L. 3  48'36' -  123'52'  CWHb i  V-G-WH-DF  220  570  20  30  0. 81  2222  41 .6  144  13. 4  37.8  24  Weeks L. 4  48" 36' -• 123'52'  CWHbi  V-G-WH-DF  220  570  20  30  0.64  1111  26 . 1  159  14 .3  41 .0  25  Weeks L. 5  48'36' -• 123'52'  CWHb.  V-G-WH-DF  220  570  23  30  0. 74  1200  20 .0  138  13 .6  41.4  26  Weeks L. 6  48" 36' -• 123'52'  CWHb.  V-G-WH-DF  219  550  19  30  0.90  2622  38 .9  128  12 .7  37.0  17. 5  1 19  1 1 5.  37.7  21  Weeks L.  22  Weeks L. 2  23  27  1 48'38' -• 123'52'  Weeks L. 7  48'36' -• 123'52'  CWHbi  V-G-WH-DF  214  Mean crown completeness estimate of f o r e s t overstory cover. 'Derived from measurement of a l l trees > 8.0 cm within the p l o t . 100 year D o u g l a s - f i r reference based on t o t a l age. 'for western  hemlock at 100 years total age.  600  22  30  0. 76  1378  d  10  METHODS  Plots cover  selected  and t o have  winter  plots  plots  ranges  i s less  (Kojima  managed  replacements  (Nyberg stand  time  consuming  35-95%, slopes  winter  Under  Criteria  used  and homogeneity s t a n d ages  no t h i n n i n g ,  200-245°,  deer  winter  forests,  winter  young  ranges  ranges  may  serve  i n the future  to select  plots  canopy  cover  no d e c i d u o u s  and elevation  were:  structure  years, overstory  no f e r t i l i z a t i o n ,  aspect  locating  t r a n s i t i o n s , and  of forest  30-60  t o deer  Selectively  randomly  i n old-growth  f o r old-growth  overstory  similar  Although  deer  cover,  10-60%,  1975).  located  e t a l . 1986).  salal  of  1984). than  sites,  as black-tailed  immaturity  a range  and Bunnell  and K r a j i n a  are typically  forests  trees,  200-700  m.  story Plots  edge 0.5  Jones  and avoids heterogeneous  ecotones  and  t o encompass  topographic characteristics  (sensu  ranges  locating  as  were  9 x 25 m  parallel x 0.5 m  each  (0.0225  t o slope 2  (0.25 m )  contours. quadrats  of four  equi-spaced  Because  of s a l a l ' s  1983),  individual  therefore  a salal  unit.  A shoot  litter  layer  plants shoot  out with  (n =  clonal  difficult  the long  was s a m p l e d  systematically  complex, were  laid  Salal  transects  i n 13  located  a s any stem  had r o o t s .  along  52/plot). growth  form  (Koch  t o d i s t i n g u i s h and  was c o n s i d e r e d a s t h e b a s i c  was d e f i n e d  that  ha) were  emerging  In the f i e l d  sample  from t h e  i t was  often  11  difficult stems when  to distinguish  often  emerged  one stem  sometimes humus  found  layer.  connected Shoots  each  diameter  (point  height  intersection portion  previous  were  allometric  and  were  growing  counted.  clipped, equations  a  line  was  point  drawn  e s t i m a t e s were  (CAG) l e a v e s w e r e  crop  from  growth  randomly  natural to  uppermost  "stretch"  height  was  part  of  pulled  l e a v e s on a l l s h o o t s i n  of shoot  foliar  among  f o r basal  t o uppermost  samples  Standing  annual  a n d be  of rooting  the shoot  season's  Random  derived  shoot.  visually  from  growth),  when  predicting  a n d were  errors  surface),  of r o o t i n g  range  or  foliage,  measured  of a t l e a s t sizes  o v e n - d r i e d , and weighed  et a l . in prep.).  current  of s a l a l  were  from  twig  growth  roots  (to avoid  and l i t t e r  point  encompassed  excluded  growth  twig  cover  shoots  distance  from  were  of the plant.  growth  a horizontal  and c u r r e n t  that  biomass  with  season's  quadrat  (Vales  roots  (vertical  distance  upright),  plots  salal  stems  the l i t t e r  and n u t r i e n t s  of sampling),  of previous season's  (vertical  shoots  above  below  c o n s i d e r e d a s one  season's  A l lrooted  height, other  were  water  two  o t h e r , and  d i d not have  percent  date  t o each  to i t just  of the rest  with  next  stems;  that  obtain  quadrat,  associated  repose  rhizome  the current  estimated.  the  S e v e r a l stems  could  non-rooted  t o determine  t o be c o n n e c t e d  independent  excluding  from  the ground  pulled  roots  Within  plots  from  t o a main  with  somewhat  was  shoots  i n each  foliar  shoot  and  the allometric  sampled  salal  of ten  to develop  and t o t a l  biomass.  30  total equations  Current  from  each  biomass  annual plot  12  (except the at  for plot  fall  after  11 i n w h i c h  completion  65 ° C f o r 24 h o u r s  determine derived  average  Overst  ory Forest  ways are  used  use  "mean  al.  (1985:181).  crown  nearby  (less  corrections  from  the four  crown  height (HBLC;  attached inside  Watts  dominant  counted. factor species  height  (BAF„)  bored  using  s p e c i e s were  were  samples  prism  Site  where  Trees  trees  (1983:402).  topheight,  t o the bole)  Point  a g e was o b t a i n e d  edge)  measured  with a of each  by b o r i n g a t  and adding  t h e age  index  calculated  was  the equations  recorded  live  taken  t o base  branches  for a l l trees  a t the four  a n d DBH o f " i n " t r e e s  with  plot  were  a four  of  live  were £ 8.0 cm DBH  basal  corners and plot  recorded.  from  and diameter a t  < 8.0 cm DBH a n d > 20 cm t a l l were  I  i n the plot or  and height  the lowest  terms  by c a n o p y .  1967) a t t h e c e n t e r  a plot  of  Bunnell et  was m e a s u r e d  dominant  trees  Tree  (DBH),  theplot.  four  15 m f r o m  (1983:424).  breast  Stand  ( 1 . 3 m) than  plot  o f CAG  i n a variety  1 9 8 5 ) a n d many  (MCC) sensu  1947; Bonnor  (« = 5 2 / p l o t ) . height  Watts  MCC o f e a c h  1 mg t o  b y number  measured  and Bunnell  completeness"  (Robinson  breast  for  biomass  h a s been  oven-dried  b i o m a s s was  t o define the proportion of sky covered  moosehorn quadrat  see Vales  t o the nearest  leaf  was u s e d ) i n  growth,  CAG f o l i a r  average  overstory cover  ( f o r review  biomass  o f t h e season's  biomass.  by m u l t i p l y i n g per quadrat.  8 leaf  and weighed  leaf  leaves  plot  were area center;  13  Definition All cover  of  variables  u n d e r s t o r y measurements  are reported  quadrats. averages  as plot  Measurements of a l l shoots  of density,  averages  of s a l a l  derived  biomass, and from  52 0.25-m  height are reported  2  as plot  sampled.  DENSITY - a v e r a g e d e n s i t y o f s a l a l s h o o t s - n r CAGBIOM - a v e r a g e CAG f o l i a r b i o m a s s o f s a l a l ( g m" ) F O L B I O M - a v e r a g e f o l i a r b i o m a s s o f s a l a l (g-m - 2 excludes CAGBIOM) TOTBIOM - a v e r a g e f o l i a r + s t e m b i o m a s s o f s a l a l ( g - n r ; e x c l u d e s CAGBIOM) PCTCOVER- a v e r a g e q u a d r a t p e r c e n t c o v e r o f s a l a l REPHT a v e r a g e r e p o s e h e i g h t o f s a l a l s h o o t s (cm) a v e r a g e s t r e t c h h e i g h t o f s a l a l s h o o t s (cm) STRHT mean c r o w n c o m p l e t e n e s s o f p l o t ( a v e r a g e o f 52 MCC moosehorn samples and r e p o r t e d a s a f r a c t i o n ) number o f p l o t t r e e s - h a " ^ 8.0 cm NTREES b a s a l a r e a ( m ' h a ) o f p l o t t r e e s > 8.0 cm BA a v e r a g e DBH (mm) o f p l o t t r e e s £ 8.0 cm AVGDIA SUMDIA - sum o f DBH (mm/225 m ) o f p l o t t r e e s > 8.0 cm HT a v e r a g e t r e e h e i g h t (m) o f p l o t t r e e s ^ 8. 0 cm CRNDEP plot trees > a v e r a g e c r o w n d e p t h (HT - H B L C ) o f 8.0cm ) o f t r e e s > 8.0 BAFBA - a v e r a g e s t a n d b a s a l a r e a (m 'ha cm d e t e r m i n e d f r o m 5 B A F „ p r i s m s a m p l e s BAFDIA - a v e r a g e DBH (mm) o f t r e e s > 8.0 cm s a m p l e d b y 5 BAFn p r i s m samples B A F T R E E S - a v e r a g e number o f t r e e s - h a > 8.0 cm d e t e r m i n e d f r o m 5 BAF p r i s m samples B A F B A D I A - DBH (mm) o f t r e e o f a v e r a g e b a s a l a r e a d e t e r m i n e d from p r i s m samples BAFSDI - R e i n e k e ' s (1933) " s t a n d d e n s i t y i n d e x " d e t e r m i n e d from p r i s m sampling computed from BAFTREES-(BAFBADIA/25) • (Long 1985) 2  2  1  2  _ 1  2  2  -  1  1  fl  1  Analys  es Salal  for  6  quadrat  samples  normality within  significant,  plots  and shoot  measurements  (coefficient  Sokal and Rohlf  were  o f skewness  1981:174) a n d homogeneity  tested not of  1 4  variance were  among  plots  necessary  to  the  normalize  best  transformation  number  of  of  22  quadrat  plots  quadrat  are  reported  by  4  to  regressed  biomass  forward  Guire  1976),  Dixon  1983).  Best of  and  and  a  height;  the  (p  0.05;  >  I used  cube  minimum root  for  root  for  confidence means  of  each  salal  are  intervals  multiplied  subsets means  regression  were  used  means  were  conservative estimate those  having  the  of  lowest  ) and  no  trend  i n the  developed  on  untransformed,  was  variables  (MIDAS,  Fox  and  (BMDP  rather  regression estimates  backtransformed  (s  variable  selection  9R,  than  obtained usually  salal  the  from  lower  abundance.  standard  error  residuals.  yx Regressions reciprocally  were  transformed  transformation  usually  transformation  f o r most  For  c o n s i s t e n c y here,  independent  variables  independent  gave  a  better  equations  curvilinear reciprocally  and  log,  variables. equation  or A  than  independent  transformed.  reciprocal a  log  variables.  r e g r e s s i o n s have  a  largest  square  and  and  quadrat  backward  because  more  estimate  gave  were  not  plots.  m u l t i p l e independent  Backtransformed  r e g r e s s i o n s were  the  mean  a l l possible  on  which  was  2  single  means  plot  for other  Means  and  suitable  f o r one  used  shoot  Transformations  m.  selection,  based  data  A  significant  estimates.  1  against  provided  not  backtransformed  using  equations  the  and  backtransformed  transformed  variables.  Transformations  represent  The  and  g,  density, cover,  all  1976:68).  for a l l plots  having  plots).  Guire  transformation  single  14  and  for a l l salal  transformation necessarily  (Fox  the  There  seems  15  to  be  no  consistent curvilinear  published 1983).  equations  Equations  form  (see reviews  based  on  a  in Bartlett  reciprocal  independent  variable  give  The  reported  are the best  results  applicable equations  only  are developed  illustrated show  within  with  variability  regressions population Tests computed program  of p l o t  the range on  plot  salal  estimates  near  data  of the data. means,  Betters  transformation  for this  of the  zero.  s e t and are Because  regressions are  intervals  samples.  m e a n s may  and  around  Reported  underestimate  s^ the  plot  > j c  's  means t o  based  true  variation. of equal  with  regression slopes  analyses  (Le 1971).  statistical  unrealistic  95% c o n f i d e n c e of  of r e g r e s s i o n f o r  tests  of c o v a r i a n c e  Except were  at a  where =  and using  otherwise  0.05.  intercepts t h e UBC  were  SLTEST  s t a t e d ,a l l  on  16  RESULTS  Regressions  predicting  Simple abundances for  linear  a r e given  measurements  examination operating salal  from  based  on p l o t  prediction  (  s  variable  and l o c a t i o n s  2  live  index  a n d were  multiple  size  Variables  poor  ratio  regressions.  the  stands of  to variability  i n t h e CWHb,  only are  ranges of  i n t h e CWHa w e r e 2 . 2 ; CWHa  r e g r e s s i o n s used  always  more  density t h e same  representing the midstory,  (CRNDEP/HT),  independent  not c o n s i s t e n t l y  also  separately i n  2  (Table  factors  Regressions  ( r ) across  and narrow  3  permits  on a c c u r a c y o f  (CWHa) c o m p a r e d  Multiple  crown  were  design  plot  affecting  samples  are presented  i n t h e CWHb  2.3).  possibly  and prism  Equations  = 0.93).  format  samples).  Regressions  3  (Table  height, site  (CWHb ).  those  r  (prism  salal  are presented  on o v e r s t o r y c o v e r ,  and factors  variability  variables.  than  based  of  d e r i v e d from  This  o f sample  due t o low sample  MCC  approach  singly  t o show  3  Equations  samples.  Equations  x  estimates  variables  measurements  . )>  ages  independent  versus  y  a stand  excluded  the plot,  the influence  CWHa a n d C W H b  within  and prism  tree  2.2.  independent  outside the plot  demonstrate  and  i n Table  of relationships  within  different  abundances  regressions predicting  MCC a n d t h e b e s t  tree  tree  salal  species  variables  selected  when  composition used  forinclusion in  Simple l i n e a r r e g r e s s i o n equations of s a l a l v a r i a b l e s a s a f u n c t i o n o f MCC, p l o t , a n d p r i s m v a r i a b l e s . S a m p l e s i z e s : CWHa=lO, C W H b , = 5 , C W H b = 7 . Equations s i g n i f i c a n t at p S 0.05. Regression equation:  TABLE 2.2  3  Y = fc  + 0  X  Variant  DENSITY:  CWHa+b,  CAGBIOM:  -36.894  37.675  0.92  5.23  N.S. -26.456  82571 35631  0.86 0.73  7.30 9.50  1/MCC 1/SUMDIA 1/BAFSDI  -119.04 -45.059 -43.481  113.02 344340 45998  0.94 0.92 0.97  14.26 15.56 9.87  1/MCC 1/SUMDIA 1/BAFSDI  -33.804 N.S. -35.439  32.681 6851 1 38345  0.78 0.72 0.73  9.95 1 1 .25 10.89  1/MCC 1/SUMDIA 1/BAFSDI  -48.568  -18.752 -18.386  44.568 133040 17887  0.95 0.91 0.96  4.74 6.67  1/MCC 1/SUMDIA 1/BAFSDI  -38.575 -8.5865 -11.475  36.867 96966 17861  0.82 0.75 0.65  7.71 9.09 10.90  1/MCC 1/SUMDIA 1/BAFSDI  -84.614 N.S. -93.610  84.140 182100 102310  0.66 0.61 0.67  33.89 35.56 33.50  230.66 152.36 179.74  -227.48 -0.01817 -0.18985  0.73 0.89 0.88  24.34 15.84  1/MCC 1/SUMDIA 1/BAFSDI  -79.705 N.S. N.S.  86.729 206830 34535  0.62 0.66 0.58  30.49 29.04 32.90  CWHa  1/MCC 1/SUMDIA 1/BAFSDI  -295.55 N.S. -337.95  305.25 690550 378460  0.61 0.57 0.64  137.54 139.57 131 . 7 8  CWHb j  MCC SUMDIA BAFSDI  5 5 0 . 13 358.32 426.24  -547.89 -0.04302 -0.45503  0.76 0.88 0.89  55.28 38.19 36.20  All  1/MCC 1/SUMDIA BAFSDI  -258.12 N.S. 409.15  279.06 653580 -0.32395  0.61 0.61 0.43  100.33 100.23 121.79  CWHa  MCC SUMDIA 1/BAFSDI  76.344 51.230 N.S.  -73.637 -0.00531 18850  0.42 0.44 0.39  15.72 15.47 15.69  CWHb 3  MCC SUMDIA BAFSDI  100.65 61.623 74.591  -104.47 -0.00764 -0.08241  0.66 0.87 0.92  7.52 7.20 5.73  All  MCC SUMDIA BAFSDI  90.468 51.695 56.308  -91.103 -0.00509 -0.03968  0.58 0.48 0.48  12.18 13.53 13.58  CWHb  All  All  TOTBIOM:  PCTCOVER:  Prefix  3  3  CWHa  CWHb  fl  3  CWHa  FOLBIOM:  yx  1/SUMDIA 1/BAFSDI CWHb  1/MCC  by  bo  *Coefficient  MCC SUMDIA BAFSDI  3  3  3  3  1/ r e f e r s not  0  to  the  4  reciprocal  significant  (p >  of  the  0.10).  independent  4.12  16.51  variable.  18  TABLE  M u l t i p l e r e g r e s s i o n equations of s a l a l v a r i a b l e s and p r i s m v a r i a b l e s . Equations s i g n i f i c a n t at p CWHaMO, CWHb. =5, C W H b J =7  2.3  Var1 ant  DENSITY:  CWHa+b. CWHb 3  CWHa  CWHb 3  N.S. -B2.347 + 63.436-1/MCC + 668.57-1/BA 1465.4-1/BA 4201.2-1/AVGDIA -179.46  +  123480-1/BAFTREES  -79.352 -54.619  + +  41.170-1/MCC + 117970-1/SUMDIA  - 6 6 . 3 1 0  +  44369-1/BAFSDI  +  232 13•  0 . 98 0.. 9 4  1/BAFDIA  0 .99  294.32-1/CRNDEP + 353.17•1/CRNDEP +  0,. 9 3 0,.93 0 .82  205.64•1/CRNDEP  131.93 118.73-MCC 2.9369•CRNDEP 572.35-1/BA 1849.6-1/AVGDIA +  50383-1/BAFTREES  +  0..99 0.,93  8212.3-1/BAFDIA  0,.99  8 .71 13 . 8 8 6 .06  5 . 75 5 . 76 9 .63 2 .67 5 .96 2 . 10  9,, 5 0  CWHa  -246.51 -179.14 -216.45  0. 92 0. 93 0. 85  17 , 3 9 1 6 ,. 0 9 24 . 58  CWHb 3  N.S. 244.52 321.68  0. 93 0. 96  14 . 0 1 1 1 . 14  0.78 0.81  24.43 22.63  91 93 85  72.48 62.39 90. 23  93 97  33 .63 22 . 1 1  3  CWHb j  Al 1  3  + + +  -  1 1 4 . 3 1 - 1 / M C C + 1 0 4 6 . 1• 1 / C R N D E P 329570-1/SUMDIA + 1216.5•1/CRNDEP 126273 • 1/BAFSDI + 8 1 8 . 2 6 - 1/CRNDEP  0.05682 0.08421  NTREES BAFTREES  8.550•CRNDEP 12.828•CRNDEP  - 2 2 5 . 16+101.OO- 1/MCC+10960-1/AVGDIA+509.98• 1/CRNDEP -147.03+291140-1/SUMDIA+7868.3•1/AVGDIA+666.39•1/CRNDEP N.S. -947.04  +  426.67-1/MCC  -701.56 -848.24  + +  123930-1/SUMDIA 478028•1/BAFSDI  +  4209.7•1/CRNDEP + +  4877.7•1/CRNDEP 3399.3•1/CRNDEP  5 . 12 6 .94  N.S. 582.34 768.28  -  0.13432-NTREES O.20079 BAFTREES  -629.72  +  335.85-1/MCC  -433.45 N.S.  +  976490-1/SUMDIA  +  20.838•CRNDEP 31.062•CRNDEP  2638.4•1/CRNDEP +  3130.3-1/CRNDEP  0.74 O. 7 9  83.58 76.06  CWHa  159.61 120.64MCC 5.1860•CRNDEP 129.50 - 0.009344•SUMDIA 5.9870•CRNDEP 1287.6-1/BAFBA 12450-1/BAFDIA + 336.98•1/CRNDEP  0.69 0 . 78 0.82  12.17 10. 34 9 . 44  CWHb 3  N.S. N.S. 120.73  0.93  5.85  0.71 0.66 0.67  10.31 1 1 .49 1 1 . 38  A l l  3  143.24  0.03623-BAFTREES -  114.22-MCC  120. 8.1-0.02483 116.16-0.02646  a  x  0, 93 O, 8 6 0.,73  CWHa  PCTCOVER  y  -84.391+42.452-1/MCC-M951.1•1/AVGDIA+232.24•1/CRNDEP -43.995 + 115790-1/SUMDIA + 281.23 -1/CRNDEP 22224•1/BAFSDI 4020.7•1/BAFDIA  A l l  TOTBIOM:  3  MCC, p l o t , s i z e s :  Equat ion  -68.507 A l l  as a f u n c t i o n of < 0.05. Sample  P r e f i x  1/  r e f e r s  Slopes  of  a d d i t i o n a l  to  the  4.1137•CRNDEP  3.7744•CRNDEP  N T R E E S - O . 1 4 8 19 • A V G D I A - 3 . 6 3 17 • C R N D E P BAFTREES-O.11013-BAFDIA-3.0284•CRNDEP  r e c i p r o c a l  v a r i a b l e s  -  -  not  of  the  Independent  s i g n i f i c a n t l y  v a r i a b l e .  d i f f e r e n t  from  zero  (p  >  0.10).  19  Form  of  relationships  The  relationships  variables or  were  curvilinear  logarithmic  where  small, were  (ln(X))  l n ( X ) gave  differences  b u t where  characteristics CAGBIOM  characteristics  transformation. hyperbolically  to predict  narrower of  than  contributed  cover  (Table  related  of independent  2.2).  t o stand variables  i n t h e CWHa a n d may h a v e As w e l l ,  considerably  higher  to the linear  relationships for  linearly  the regression.  estimates  TOTBIOM  predicting  predicted  plots than  form. PCTCOVER  by BAFSDI  density  t o stand  an l n ( X )  characteristics  used  were  transformation  b i o m a s s was  equations  i  data.  related  t o stand  was  Ranges  of these  than  )  differences  salal  related  combined data  x  between  t h e 1/X  equations  .  I used a  crop  f o r most  3  better  y  better,  hyperbolically  2.2), with  s  equations  standing  also  CWHb .  f o r t h e range  (1/X)  For equations  (smallest  Salal  and  were  be u s e d .  predictability,  the relationship  (Table  consistently  however,  the best  was a l s o  hyperbolic  and logarithmic  B e c a u s e t h e 1/X was o f t e n  and  giving  could  and forest  that  predictability  1/X g a v e  to describe  Salal  density  and suggested  reciprocal  hyperbola stand  salal  equations  the best  between  large.  between  consistently i n t h e CWHa  except  when  BAFSDI  FOLBIOM a n d TOTBIOM,  characteristics i n t h e CWHb affected  3  in  the  were  the best  form  21 a n d 27 h a d b i o m a s s predicted  The forms were  o ft h e  mostly  i n t h e CWHa  and l i k e l y  linear,  (Table 2.2).  except  20  Differences  among  Subzone against  relationships  effects  G-WH-DF  within  CWHb,) a n d p l a n t  (G-WH-DF CWHb, a g a i n s t CWHa a g a i n s t of  slope  reciprocally and  prism  three  equations  ranges  however,  CAGBIOM  a n d 1/BAFTREES.  however,  with  variables. FOLBIOM  1/NTREES,  Because  analysis  valid.  regressions reciprocally  among  from  variants  a n d CWHb  the slope equal),  o f BA, B A F S D I ,  3  test to  indicating  Equations  variants  predicting a n d BAFBA,  differed  this, both  independent  found f o r 1/SDI,  significantly,  a n d 1/BAFBA  f o requation  using  were  1/MCC, 1/SUMDIA,  t h e CWHa a n d CWHb  Acknowledging  transformed  from  of relationships  to test  plant  CWHbi  nearly  among  1/BAFSDI,  forms  p l o t SDI,  variants.  Equations  d i f f e r e d between  covariance strictly  forms  significant differences predicting  NTREES,  effect.  associations.  among  variable.  i n t h e same  variables  reciprocal  Results  d i f f e r e n t among t h e  22 was d r o p p e d  plant  d i dnot d i f f e r  equations 1/BA,  from  (G-WH-DF  b u t d i dn o t d i f f e r  3  between  within  tested.  MCC, SUMDIA,  (CWHa, CWHb,, C W H b ) ,  (plot  were  CWHa  as a function of  significantly  of independent  density  No  density  i n d i c a t i n g no subzone  difference  salal  salal  were  differed  effects  the independent  t h e CWHa a n d CWHb, v a r i a n t s  association,  a  subzones  transformed  BAFTREES  variants  between  have  d i f f e r e d with  (l/X)  association  (G-WH-DF  3  3  predicting  association  V-G-WH-DF C W H b ) a n d a c r o s s  V-G-WH-DF C W H b )  tests  Equations  a plant  as  for 3  independent  TOTBIOM a n d  (Table  2.2),  differences  I still  i s not  tested the  untransformed and variables  to  explore  21  potential  differences.  1/SUMDIA,  1/SDI,  1/BAFSDI  and  and  1/BAFBA  among  variants  Tests  of equations  variants  when  equations  forms,  residual  with  did.differ.  d i d not  with  TOTBIOM.  PCTCOVER  were  tests  a l l equations would  variants  have  a t a = 0.05.  except  no  i s some  The  regression  considerable  resulted  i n high  not s i g n i f i c a n t .  Had  1/BAFSDI a n d  indicated  f o r equations  among  d i d not d i f f e r  All  slope  a difference  there  was  differ  among  equations  the F-test  with  variables.  t h e two d i f f e r e n t  o f FOLBIOM a n d TOTBIOM  predicting  1/BA  1/MCC,  a difference  not d i f f e r e n t .  variable.  TOTBIOM  and  Because  using  and thus  independent  indicated  FOLBIOM,  any independent  0.01,  Equations  1/NTREES  are likely  variance  Equations  equations  predicting  equations  using  1/BAFTREES; h y p e r b o l i c  of the r e s u l t s  variability  d i d not d i f f e r  f o r a l l untransformed  predicting  consistency  Equations  significantly  a been  1/BAFBA  set to  predicting  significant difference  predicting  CAGBIOM,  among  FOLBIOM, a n d  TOTBIOM. Reciprocally results  opposite  transformed those  BAFTREES  for salal  1/BAFSDI  i s a poorer  1/MCC, in are  1/SUMDIA,  part), likely  density  Results  of t r a n s f o r m e d  density  1/SDI,  and  Results  of slope  tests  variables  using  plot  SUMDIA,  than  plot  the plant  consistent  and  Because  samples of (Table  2.2  associations  among a l l  f o r FOLBIOM a n d  1/SDI  test  SDI, and  i n t h e CWHa+b,  between were  slope  relationships.  of density  1/NTREES  relationships  independent  MCC,  and biomass  predictor  different.  untransformed  BAFBA a n d BAFSDI g a v e  TOTBIOM.  1/BAFSDI t o  22  predict  salal  density  and biomass d i f f e r e d ,  between  SDI a n d BAFSDI  within  differences.  Equations  variants  always s l i g h t l y  were  differences  between  Differences prism  located stand 7,  plots  pattern  from  of dense  i n dense  t h e way t h a t  of v a r i a t i o n  surrounding  measurements  the  CWHa, b u t p r i s m  than  within  using  BAFSDI  plot  samples  (e.g.,  CWHb ;  2  being  located i n  dissimilar  plots  SDI  (e.g.,  slope  tests  samples accounted  samples  for  3,  may  the  from t h e  closer  3  more  measurements  maximum more  BAFSDI  range  Maxima  prism  samples  3  trees  more  BAFSDI  t h e same  S D I 1 3 3 5 i n CWHa,  of plot  plot  diverged  1461 i n CWHa, 8 1 8 i n C W H b ) .  f o r measurements  3  plot  minimum  followed  with  similar in  i n t h e CWHa,  a n d BAFBA 3  plot  1 and 2 4 ) .  than  i n t h e CWHb  2 3 3 i n CWHb ;  different  (maximum  were  t o t h e minima  s a m p l e s o f BAFSDI  SDI 2 9 5 i n CWHa,  between  (see Appendices  The minima  3  maxima  differed  a n d BA a n d B A F B A  a n d 238 i n C W H b ) .  with  overlapped  (r  SDI  sometimes  (e.g.,  had a higher  variables  i n t h e CWHb .  minimum  plot  were  Plots  o f BAFSDI  samples had a narrower  f o r prism  i n CWHa  pattern  of plots  o f SDI a n d BA w e r e  minima  than  no  measurements and  stands.  that  patches  o f SDI a n d B A F S D I ,  measurements  3  biomass  because p l o t s  o r open  and prism  range  with  between  prism  of independent  The  428  found  stands.  Ranges  but  variants  1, 2, 4, 1 3 , 15, 2 3 , 2 6 ) . T h e e q u i v o c a l  result  tree  plots  h a d a n SDI l e s s  10),and p l o t s  salal  better  not consistent  i n patches  gaps  predicting  f-tests  0.01 a n d 0 . 1 0 ) .  among  s a m p l e s were  and a c r o s s  but paired  than  1028  in  Ranges  (max-min SDI  23  733,  BA 2 6 . 8 ) t h a n  Discrepancies pronounced  Relative  from  Equations  salal  basal  was  a poorer  among  to  the  predictor  data.  and spacing.  number  differences  are illustrated in variable  of trees,  SDI d e r i v e d than  salal.  o r SDI from  prism  predicted  prism  samples  although  salal  variables  I n t h e CWHb  3  b y MCC i n 2.2).  than  MCC  other  MCC was g o o d f o r  predictor  inter-plot  's o f e a c h  separate  (Table  a n d TOTBIOM  was a p o o r e r  Other  in  y•x  predicted  by BAFSDI  o f FOLBIOM  considerable  in the s  variables  a n d TOTBIOM  3  i n t h e CWHa.  independent  for predicting  variable  i n t h e CWHb ,  BAFSDI  trees  SUMDIA.  independent  and density  was a g o o d  of trees.  f o r FOLBIOM  CWHa+b, w i t h  BAFSDI  plot  o n l y modest •*  measurements  combined  derived  variables  independent  area,  independent  o r number  except  CWHb  size  was t h e b e s t  on p l o t  were  the  forest  of values  contributed  t o MCC a s a p e r c e n t  SUMDIA  variable  3  less  abundances (Table 2.2).  measurements o f p l o t  area  variants  by  of salal  based  There  the  salal  was a b e t t e r  basal  were  12.8).  transformations  independent  (MCC) g e n e r a l l y  n o t a s good a s u s i n g  samples of  cover  2.2-2.6.  derived  ranges  and l i k e l y  3 9 0 , BAFBA  methods  Reciprocal  between  among  predicting  Relationships  were  methods  predictability  variable  (BAFSDI  results.  Overstory  Figs.  of trees.  the differences  t h e two s a m p l i n g  equivocal  samples  between t h e two s a m p l i n g  f o r number  accentuated by  for prism  of density i n  v a r i a t i o n of tree were  where  predicted  trees  were o f  well  24  175.0  -i  30  40  50  60  70  80  90  100  M E A N C R O W N COMPLETENESS (%)  2  F i g u r e 2.2. T h e r e l a t i o n s h i p o f s a l a l d e n s i t y - n r and 95% c o n f i d e n c e i n t e r v a l s t o mean c r o w n c o m p l e t e n e s s . L o w e r e q u a t i o n i s f o r CWHa a n d CWHb, c o m b i n e d d a t a ( • ) , u p p e r e q u a t i o n CWHb (ffl). 3  25  100.0  -i  2  F i g u r e 2.3. The r e l a t i o n s h i p o f f o l i a r p r o d u c t i v i t y (g-nr ; CAGBIOM) a n d 9 5 % c o n f i d e n c e i n t e r v a l s t o mean c r o w n c o m p l e t e n e s s . CWHa ( • ) , CWHb, ( A ) , CWHb (ffl). E q u a t i o n i s f o r combined data. 3  26  250.0 - i  200.0<N i  MEAN CROWN COMPLETENESS (%)  2  F i g u r e 2.4. The r e l a t i o n s h i p o f s a l a l f o l i a r b i o m a s s ( g - n r ; FOLBIOM) a n d 95% c o n f i d e n c e i n t e r v a l s t o mean crown c o m p l e t e n e s s . CWHa ( • ) , CWHb, ( A ) , CWHb (ffl). E q u a t i o n r e p r e s e n t s combined d a t a . 3  27  1000.0 - i  800.0-  3> CO  30  40  50  60  70  80  90  100  MEAN CROWN COMPLETENESS (%)  - 2  F i g u r e 2.5. The r e l a t i o n s h i p o f s a l a l t o t a l biomass ( g - m ; T O T B I O M ) a n d 9 5 % c o n f i d e n c e i n t e r v a l s t o mean c r o w n c o m p l e t e n e s s . CWHa ( • ) , CWHb, ( A ) , CWHb (ffl). E q u a t i o n r e p r e s e n t s combined data. 3  28  70.0  - i  30  40  50  60  70  80  9 0 100  MEAN CROWN COMPLETENESS (%)  F i g u r e 2.6. The r e l a t i o n s h i p o f p e r c e n t c o v e r o f s a l a l a n d 9 5 % c o n f i d e n c e i n t e r v a l s t o mean c r o w n c o m p l e t e n e s s . CWHa ( • ) , CWHb, ( A ) , CWHb (ffl). E q u a t i o n r e p r e s e n t s c o m b i n e d data. 3  29  relatively well.  The poorest  equations among  t h e same  variants.  individual different  Both  simple  either (Table  significant  equations,  added  variable,  regression related  (Table  Equations than  equations estimate  found  in salal for  to differ  f o r stand variation in often  equations  variants.  accounted f o r  multiple  little  additional  yielded  among  already  density,  very  data  o r no  additional  variables  were  (CRNDEP) was t h e m o s t  included Salal  i n a l l b u t one  important  multiple  was c o n s i s t e n t l y  inversely  depth.  predicting for  were  FOLBIOM  CAGBIOM. crop  more  a n d TOTBIOM  The d i f f e r e n c e s  biomass  variable  were  more may b e d u e  i n d i r e c t l y from  than  t h e more  allometric  direct  o f CAGBIOM.  s hoot Mean  hei ght  repose  variability generally, (Fig.  combined  were  equal  regressions  depth  standing  that  for  variables  t h e p r e d i c t a b i l i t y of biomass or  2.3).  those  estimating  a l l salal  t h e two methods  Where  crown being  t o crown  variable  2.3).  methods  of equal  accounted  and improved  cover  Salal  linear  variable  nearly  although  90% o f t h e v a r i a t i o n  variation  for  f o rt e s t s  predicted  equations  sampling  accounted  results  regressions  to  because  variants,  Because  BAFSDI  independent  was BAFSDI  characteristics  over  size,  2.7).  when shoot  and s t r e t c h related height  height  showed  t o MCC among peaked  CWHb, a n d CWHb  3  between  plots  considerable  t h e CWHa p l o t s , b u t 65 a n d 8 0 % MCC  sampled  within  stands  30  100.0-1  80.0-  60.0-  40.0-  20.0-  0.0 30  —r— 40  60  l  60  70  l  80  —1 90  1 100  MEAN CROWN COMPLETENESS (%)  F i g u r e 2.7. The r e l a t i o n s h i p of average height of s a l a l and 95% confidence i n t e r v a l s t o mean crown completeness (a) repose, (b) s t r e t c h . CWHa (•), CWHb, (A), CWHb ( a ) . 3  indicated  a similar  CWHa p l o t s MCC,  were  likely  relationship indicated  pattern  larger  than  due, i n part, of salal  similar  of height. i n CWHb  3  Shoots  plots  a t nearly  t o age d i f f e r e n c e s .  height  patterns,  t o other though  stand  more  i n some  ofthe  equal  The characteristics  variable  than  t o MCC.  DISCUSSION  Salal stand  abundance  Density  and f o r combined  was c l o s e l y  two p l a n t  variation CWHb  well  c h a r a c t e r i s t i c s i n the separate  associations,  the  was g e n e r a l l y  in salal  biomass  (Tables  due t o age a n d s i t e  plant  2.2 a n d 2 . 3 ) .  stand  Regressions and cover  to forest  variants,  to single  associations.  variants  3  related  data  related  characteristicsin  accounted  i n t h e CWHa  differences  f o rless  than  among  in  the  t h e CWHa  plots. Differences subzones, testing from  in relationships  o r between  f o r plant  most  plant  effects  T h e CWHb  2.2) o f g e n e r a l l y  smaller  other  plots.  among  predicting tests  current  results  and because  a r e somewhat  (possible  Type  II e r r o r ) .  relationships  variability  within  Most  variants  5 a n d 6 was g r e a t e r  level  o f an independent  than  variable  24) a n d c o n t r i b u t e d  CWHa e q u a t i o n s .  Slopes  Results  variants  not s t r i c t l y  a n d may b e  there  Figs.  to the greater  of equations  among  were  valid, the  tests  revealed  i s considerable  CWHa p l o t s  (e.g.,  of  misleading  2.4 a n d 2 . 5 ) .  other  density  2.7) t h a n t h e  among  though  (Figs.  predicted  biomass.  statistical  do n o t d i f f e r ,  plots  Appendix  equations  inconclusive,  when  d i d not d i f f e r f o r  foliar  were  found  (Fig.  variants  the tests  between  had a greater  shoots  growth  o f FOLBIOM a n d TOTBIOM  equivocal,  that  annual  were  on d e n s i t y  plots  3  (Fig.  Equations  variants,  associations  association  variables.  among  Biomass i n  at a  given  2.3-2.5;  variability variants  of  the  predicting  33  FOLBIOM a n d TOTBIOM differ,  however,  differ. than 6,  Plot  plot  (Chapters  found  BAFSDI  found  with  tests with  equations  salal  was  equations Soil  of  forest. plot  were  salal  from  SDI a  there  biomass.  were  salal Plot  5  in, I  feel  plant  differences from  no  with  NTREES  effect  on  untransformed of p l o t s plot  tree  and  plot  tree  sampled  site  a  of  data. of  among  function  of p l o t  trees  nearly  among  as  equations  measurements  were  t h e same a r e a  that  of equal  and  ranges  either  salal  differences  and  were  measurements.  measurements a r e t h e most  associations  and  and p a t t e r n  are partly  predicted  status  BA  equations  a measurement  and  plot  reciprocal  that' t e s t s  and m o i s t u r e  were  in relationships  Because  sampled  i n both  than  sample,  samples  nutrient  than  was  prism  plot  a  associations  than  using  with  cases  pronounced  different  No  i n biomass  for patchiness  Plot  and  equations  density  several  the d i f f e r e n c e s  derived  precise  similar  plots  method.  as BAFSDI,  predicting more  biomass  than  results.  predicting  variables  or between p l a n t  than  of  h a d a more  accounted  that  sampling  well  6 d i d not  equations d i d  and  radiation  regression  and  results  that  among  conclude  rather  5 and  i n a denser  Differences  BAFBA,  independent  samples  variation  of  6 was  the majority  BAFSDI,  of  variants  f o r FOLBIOM  solar  of equal  A l l equivocal  Prism  more  plots  productivity  plot  a n d BAFBA.  transformation  I  though  among  prism  overlap  excluding  3 and 4 ) .  inconsistent  BA.  6 had g r e a t e r  received  Several  were  MCC  the intercepts  5 even  however,  from  tree  regression appropriate. species  effects  were  alone  may  not  explain  short mean  the differences  and numerous density  Bunnell  by c o m p e t i t i o n  shoots  t o grow  competition affect stand be  other  Long  Long  (1977)  annual a  plots stands than  and Turner  stand  types  Betters  Pyke  that  a plant  found  a n d Zamora  (1982)  shrub  production  found  a hyperbolic  measured MCC.  with  Bunnell  because  changes  small  crop  density  be e x p e c t e d i n MCC  may No  density,  (1975), and  biomass,  from  and Morgan  current greater  weak  to  strong  i n a v a r i e t y of 1981;  Bartlett diverse.  r e l a t i o n s h i p of  Bunnell  linear  (1986)  may  stands.  a hyperbolic  and a  light  younger  of r e l a t i o n s h i p s a r e a l s o  and Vales  would  in a  and Long  r e l a t i o n s h i p of s a l a l  relationships  that  in salal  and o v e r s t o r y  i n Idaho.  a moosehorn  and c l o s e s ,  were  3  anything  see Specht  found  t o MCC  individual  r e s p e c t i v e l y , were  have  Forms  thinning  association effect.  standing  researchers  1983).  2  the greater  Turner  i n older  ( f o rreviews  (plot  i s increased  trends  than  are very  shoots«m" ,  and n u t r i e n t s  sampled,  understory  areas  self  allowing  i n t h e CWHb  and cover,  shoots  189  develops  stand  between  as  time,  there  (1975),  salal  biomass,  22-year-old  relationships  and  stand  and a l s o  rather  found  growth  Other  and  density  f o r moisture  Because  most  With  a r e a v a i l a b l e on s u c c e s s i o n a l  although  in  trees  an age e f f e c t  data  As a  i s reduced with  data).  reduces  larger.  salal. than  c l e a r c u t as high  unpubl.  caused  transmission  Salal  i n c l e a r c u t s a n d on e x p o s e d  i n a young  and V a l e s  in density.  and V a l e s  density  t o MCC  one b e t w e e n  stated  that  (1986)  salal  cover  hyperbolic  for productivity variables  a t low l e v e l s  o f MCC  have  a  35  greater  effect  transmission also MCC  that  be  necessarily Results and  of  and  truncating  were  found  narrow the  of  two  biomass  of  salal  were  Young  et  basal  as  cover  MCC  among SUMDIA  or  range  of  variants, for  combined  data  and  to  not  salal  the  CWHb  density  may  3  be  a  variable  Vales  1986)  or  the  (1975) p r e s e n t e d  salal  biomass  Pyke  were  Where was  predictor  to  equations  of  as  's)  an  foliage  of  of  good  to  they found  as,  or  believed  plot  were not  variants,  BAFSDI  because  equations  among  were  biomass  poorer  results  of  predicted  than  the  two  using  to  MCC  sampling  that  than basal  not  better  Within  salal  separate  (1962),  understory  salal  were  predicting  for  Ripley (1982)  acceptable,  almost  but  and  Zamora  equations  data.  between  and  > x  variables,  Shaw  relationships  trees  MCC  data.  s^  (lowest  dependent  better  estimates  difference  of  cover  linear  in  and  They  variable.  for  The  Turner  variables  among  variable,  for  (Bunnell  MCC.  may  productivity  independent  relationship  a  combined  differed  cover  hyperbolic  and  of  understory  regressions  Long  combined  was  of  of  cover.  the  light  overstory.  SUMDIA.  independent  The  some  and  levels  understory  for  for  Although  number  higher  were  linear  (1967),  area. or  for  at  single  here  consistent  a l .  a  independent  and  overstory  area  forest  best  variants,  to  plots.  curvilinear  throughfall,  relationships  distribution  inverse  The  than  because  regressions  a  influence  linear  related  relationships of  3)  expected be  biomass  result  competition,  (Chapter  stated would  on  as  good  differ than  was  a  good  variants from or  BAFSDI, SUMDIA.  methods  was  36  unexpected.  Forest  time-consuming. are  easier  inventories  Point  and quicker  samples  i n t e n s i v e l y with  with  probability proportional  sampled (Watts by a  reducing  In  sample  those  comm.,  variables  prism  where  using  were  variants  (Table  well  trees  of radiation  salal  samples  plot  samples.  point  trees are classes  the understory  o r by c o m p e t i t i o n . sampling  With  will  Smith,  related  to prism  Prism  the plot  samples which  f o rs o i l  moisture  transmission independent radiation  using  greater,  independent (prism)  S a l a l biomass  be  solar  influence  of the stand  radiation.  that  diameter  (J.H.G.  density  of solar  The  whereas  diameter  i n t e r - p l o t v a r i a t i o n was  transmission  influence  a r e sampled  frequency,  results of point  sampling  sampling  a r e sampled  trees  and larger  probably  from  2.2).  and outside  competing  trees  because  to tree  to size  of plot  derived  however,  inside  from  sampling  size,  predicting  than  obtained  Smaller  sampling a r e  U.B.C).  t h e CWHa  regressions  poorer  Smaller  enough  plot  i n t e n s i t y as smaller  transmission  approximate pers.  t h e same  1983).  large  plot  i sproportional with  from  t o sample.  more  sampling  derived  samples  were  and cover,  i n separate  included  trees  would  have  Trees  inside  both  an i n f l u e n c e  and nutrients,  the plot  on  would  and also  of radiation.  variables  selected  i s important  biomass.  Anderson  et a l .  postitive  r e l a t i o n s h i p s between  radiation  under  forest  (1969)  canopies.  i n regressions  f o rsalal  d e n s i t y and  and Zavitkovski understory Alaback  suggest  (1976)  abundance (1982)  found  and solar  believed  that  37  transmission growth,  of solar  but that  difficult  MCC,  1959), area  of solar  SUMDIA, with  competition SUMDIA.  (Horn  Because  variables,  structure  factors  affecting  understory.  area  h a s been (e.g.,  being  density  found  Wellner  better  foliage  1971) i s a l s o  density  with  intra-specific  competition  and r a d i a t i o n ,  predicting affects  salal  per unit biomass  transmission with  and  of overstory  amount  salal  abundance  examined  stand  Salal  height  CRNDEP,  through  be tree  radiation are  4.  has n o t been  30 a n d 9 1 % MCC,  and basal  area  based  on f i v e  plots.  and V a l e s  (1986)  a  moosehorn  on  relationship  Koch  between  relationship  linear  studied.  t o MCC  parabolic  inverse  extensively  height 1  spacing  between  found  2  by  which  (1983)  m «ha"  an  single  regressions  The r e l a t i o n s h i p s of s o l a r  to  a s by  and would  age and c o m p e t i t i o n  and t r a n s m i s s i o n  i n Chapter  MCC a n d  consequently  included  radiation  foliage.  basal  influenced  as well  Multiple  and cover  of solar  correlated  trees  area.  than  related  strongly  competition  biomass  related  1948; M i l l e r  with  strongly  i s probably  were  and consequently  correlated was  t o be  predictors  inter-specific  affecting  f o r understory  of f o r e s t  Overstory  salal  important  measures  a n d SUMDIA  3).  was  radiation  and b a s a l  MCC  (Chapter  plot  indirect  t o use as causal  Transmission to  radiation  Bunnell  of average  19 p l o t s .  study  of s a l a l  height  three  variants  indicates  height  t o MCC  The c o n s i s t e n t  under  MCC  that  between  of average  peak  shoot  0 t o 66 found  measured  found  with  in this  65 t o 8 0 % a m o n g t h e  a parabolic  relationship  may  a  38  apply may  for different  vary  with  age,  differences. in  more  1972)  no  incoming  areas  consequently dense  because  (Bunnell  et  for  such  allow  of  of  the  Tall  same a g e  peak,  Very  however,  be  stress  dense  to  be  canopies  intercept  and  tall  shoots  indicate  that  salal  under more  reduce  and  moderately  available  snow are  shorter (Swank  photosynthesis,  shoots  forage  may  moisture  limiting  canopies  a l . 1985)  the  light.  growth.  would  of  because  potentially  height  canopies  winter  either  location  ( c o m p e t i t i o n ) , or m i c r o s i t e  in stands  competition  light,  The  density  Shoots  open  or  sites.  during  effectively  less  likely  to  be  buried. The stand  characteristics.  whether with  results  salal  forage  predictions  abundance possible sampled  for here  specific (e.g.,  sites  plot  use.  I  of  equations  variables Because  with  point  among high  and  variants  compared  to  80%  tradeoff  between  MCC  f c r combined  snow  or  of  study.  to  are  (Tables  data  interception,  salal  about  and 2.2  are  aspects, sampling  which  equations  because these and  tests two  2.3).  foliar  double  equations.  salal  Salal  those  forest  SUMDIA  unequivocal  curvilinear,  biomass  agreement  inventory data  determine  MCC  predictabilities are  in  similar  method  to  however,  south-southwest  would  were  say,  in this  timber  The  either  is related  thinning  equations  untreated,  using  foliar  to  characteristics  samples)  relationships  productivity  to  p r e d i c t e d from  recommend  had  the  associations).  or  to  respond  from  (immature,  plant  is difficult  would  made  estimates  It  abundance  at  65%  The  MCC  optimum  productivity,  and  shoot  height Salal  may  occur  control  possible.  Salal  11,  having  and  (based  26 on  plot  through  abundances Reineke's  tree  1005,  respectively.  would  probably  effects  (e.g.,  considerable salal  than  SDI's  of  These  plots, were  open  stand  than  plot  Stands  have  an  expected  SDI  were  at  however,  were  i n dense the  plot  f o r a l l of  minimum  of  SDI's  or  on  steep  support  1397,  4,  these  salal,  higher  by  was  plots).  1,  9,  1243,  and  than  1000  although  site  slopes allowing  patches  (BAFSDI  also  indices  greater  1028,  influenced stand  1335,  13-15,  and  is  in plots  density  974,  effect  in plots  1544,  a  stand  having  may  MCC.  d e n s i t y management  (1933)  regime  (as  1388,  outside  stand  adverse  moisture  located  65-70%  measurements)  sidelighting)  1201,  they  around  abundances  and  23  with  of  plot  respectively). sidelighting  with  a  because  relatively  considerably  lower  CONCLUSION  The  functional  overstory  characteristics  associations effects.  studied,  Simple  characteristics more  of  the  The cover  between  forest plant  stand  regressions  biomass  and  improved developed  variable  stands  than  were The  predict  salal  tree  height  Although  mean  shape  the  of  variants  the  was  salal  hyperbolic, stand are  of  ages  density  greatest  height  was was  but  crown  for  or  plots. and  simple  including  and  90%  using  variation  in  salal  crown  depth  Relationships locations  within and  some  were  stands. forest  equations  characteristics  mean  and  equations.  different  age  different  samples,  understory  relationship  sampled.  58-82%  to  biomass  data  between  stand  shoot  plot  regressions of  due  sampled  not  developed  forest  or  are  relationships  abundance  diameters, Salal  of  generally  best  for  plant  accounted  combined  from  forest  overstory  productivity,  For  to  two be  among  characteristics  obtained  the may  with  density  salal  Multiple  Relationships  linear.  of  density  in  variables  salal  predictability  among  overstory  regressions  in  salal  difference  accounted  cover.  the  the  associations.  variables  of  different  independent  variation  independent  more  linear as  is  but  relationships  to  linear  relationship  to  use  completeness,  were  to sum  of  index.  between  65  less  the  the  in  same  and  80%  CWHb  among  3  the  MCC. variant, three  the  41  REFERENCES  Alaback, P.B. 1 9 8 2 . 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R E L A T I O N S H I P S BETWEEN RADIATION  AND  FOREST  TRANSMISSION  STAND  OF  SOLAR  CHARACTERISTICS  INTRODUCTION  Transmission  of shortwave  stands  h a s been  Miller  1959; V e z i n a  theoretically Norman al.  (e.g.,  1975; Norman  1985).  related  Solar  efficiency radiation  and Pech Anderson  1 9 6 6 ; Cowan  and J a r v i s  Miller  cover (e.g.,  (Bunnell  Zavitkovski canopies  characteristics  (e.g.,  Miller  has the advantage  a f f e c t e d by e x p o s u r e  transmission because 1948),  and stand  of stand o r methods  and  has been  stand  e t a l . 1969) o r 1959;  of being  widely  a p p l i c a b l e , but  Relationships among  basal  stand  density  (sum o f d i a m e t e r s )  may  composition  area,  have  between  sites  predictor variables  Crown c l o s u r e ,  number  been  with  Anderson  Hemispherical  1986).  and  solar  1969).  Vales  growth  production  using  a n d Coombe  s t r u c t u r e or species  et  4 ) , deer  Predicting  characteristics  1948; modeled  understory  1959; P e r r y  conditions.  of sampling  canopies  c a n be done  (Evans  Madgwick and B r u m f i e l d  photography is  photography  or  1976; C h a p t e r  1976).  forest  hemispherical  Wellner  1981; B a l d o c c h i  1959),  e t a l . 1986),  under  1964a;  forest  forest  1968; L e m e u r 1973;  1975; R o s s  under  (e.g.,  into  1964) a n d d i s c u s s e d  e t a l . 1969; Z a v i t k o v s k i  thermal  radiation  e m p i r i c a l l y (e.g.,  radiation  t o snow m e l t  (Anderson elk  evaluated  solar  differ  (Wellner (Bunnell  and  of t r e e s , and  t h e most  common  46  predictor  variables.  Some  researchers  radiation  components  Reifsnyder et  et  a l . 1984a,  have into  studied forest  a l . 1971/72; 1984b).  stands  Hutchison  Anderson  of  appropriate  separation  of  r a d i a t i o n components  al.  (1971/72)  importance of  of  and  direct,  by  et  the  a l . (1971) the  1977;  each  spatial  Baldocchi  design,  variation  and  are  Reifsnyder  demonstrated  1971;  the  sample  because  canopy.  Muller  stressed  instrumentation,  considering  study  solar  forest  examined  diffuse,  (measured  as  and  photon  radiation  stands  objectives  on  were  transmission  of  different assess  Anderson  predicts  1)  solar  degree  radiation.  of  as  immature  British  et the  of  penetration  i f the  the  diffuse,  active jim)  components  coniferous Specific  predicting from  forest  stand  r e l a t i o n s h i p s were s t r u c t u r e s ; and  hemispherical by  global,  Columbia.  regressions  different  adapted of  the  0.4-0.7  r a d i a t i o n components  which  transmission  PPFD,  into  Island,  develop  stands  (1964a)  p.m)  determine  to  of  photosynthetically  density,  (0.3-3.0  2)  penetration  diffuse flux  to:  between  the  the  Vancouver  characteristics;  of  Gay  Matt  separate  radiation. This  of  differently  of  (e.g.,  and  (1964b)  importance  intercepted  transmission  Vales  and  diffuse  3)  photograph Bunnell  PPFD,  and  method  (1986) direct  47  STUDY AREAS  Twelve Vancouver  plots  Island,  the  Coastal  and  6 were  separated  Columbia,  Hemlock  i n t h e CWH  at several  d r y subzone  wet s u b z o n e  CWHb p l o t s  than  800 m between  The  CWHb  stand  the  CWHa  (Table  represented  were  was y o u n g e r , 3.1).  by stands  references  Sixplots  (CWHb).  T h e CWHa  of d i f f e r e n t  the farthest  separated  smaller  were  stand plots.  than  t h e two s u b z o n e s  broad  t o subzones  trees  plots  ages and  i n t h e same  with  were i n  (CWHa; K r a j i n a 1 9 6 5 )  a l l located  Because with  l o c a t i o n s on  Canada.  g e o g r a p h i c a l l y and i n stands  less  future  sampled  British  Western  structures. with  were  stands i n  were  structural differences,  will  be synonymous  with  structural differences. All  plots  (Pseudotsuga het er ophylI pii  were  menziesii a  (Raf.)  cat a D o n n ) .  were  amabilis  yellow-cedar western  white  Plots cover.  treatment  (Mirb.) Sarg.),  stands  f i r (Abies (Chamaecypari  and western  pine  were  (Pinus  south-southwest  found  used  nookat  ensi s  monticola  stand  (Thuja  redcedar  i n t h e CWHb  stand  Forbes),  (D. Don) S p a c h ) ,  a range  and  were:  of overstory  homogeneous  a g e 30-60  no t h i n n i n g o r f e r t i l i z a t i o n ) , aspects,  (Tsuga  hemlock  Dougl.).  to locate plots trees,  western  (Dougl.)  s e l e c t e d t o encompass  no d e c i d u o u s (e.g.,  amabilis s  of Douglas-fir  Franco),  Additional species  Criteria  structure,  i n immature  a n d e l e v a t i o n 200-600  years, slopes m.  canopy  no  stand  10-60%,  TABLE  Plot  3.1  Descriptions  of plots  Lat i tude- Longi tude  used  Aspect  C) 1  d  2  d  e  d  7  d  9  d  Elev. (m)  Slope  solar  Age  radiation  MCC  a  (%)  and f o r e s t  stand  characteristics  C  Spp. Comp^ N T R E E S BA Avg. DBH DF-WH-MiSC U/ha) (m'/na) (mm) C  C  Avg. H t (m)  C  CRNDEP (m)  5 0 '  1 2 '  -  1 2 6 * 2 8 '  2 3 0  4 3 0  3 0  2 8  0 . 9 1  2 8 - 7 2 -  0  2 5 3 3  3 7 . 7  131  1 2 . 4  5 0 '  0 7 '  -  1 2 5 *  2 1 5  2 7 0  15  3 7  0 . 7 8  6 1 - 2 3 -  0  1 6 0 0  4 6 . 7  1 7 3  1 3 . 6  4 8 ' 5 4 '  -  1 2 4 ' 1 8 '  2 0 6  3 1 0  5 2  6 0  0 . 6 4  1 0 0 -  o -  0  7 5 6  3 5 . 0  2 3 5  2 5 . 2  7  4 8 ' 5 4 '  -  1 2 4 " 1 8 '  2 1 5  3 1 0  4 5  5 9  0 .  1 0 0 -  0 -  0  1 7 3 3  3 4 . 4  1 4 5  1 5 . 6  5 . 8  4 8 ' 5 4 '  -  1 2 4 ' 1 8 '  1 7 3  2 0 0  9  3 4  0 . 3 5  8 - 9 2 -  0  2 6 7  1 4 . 8  2 1 9  1 5 . 7  4 8 ' 4 9 '  -  1 2 4 ' 1 9 '  2 6 1  6 3 0  4 3  4 2  0 . 9 0  3 7 - 6 2 -  1  5 4 . 9  1 4 9  1 5 . 0  7 . 9  1 1 0  9 . 7  8 . 9  3 9 '  7 8  2 7 1  1  1 0 .  1  . 3  1 3 .  1  s  4 8 ' 3 6 '  -  1 2 3 ' 5 2 '  2 4 0  5 7 0  3 0  3 0  0 . 5 8  9 7 -  3 -  0  8 8 9  1 0 .  2 2  e  4 8 ' 3 6 '  -  1 2 3 ' 5 2 '  2 4 0  5 7 0  3 0  3 0  0 . 4 2  1 0 0 -  0 -  0  7 5 6  8  2 3  e  4 8 ' 3 6 '  -  1 2 3 ' 5 2 '  2 2 0  5 7 0  2 0  3 0  0 . 8 1  7 4 - 2 0 -  6  2 2 2 2  4 1 . 6  1 4 4  1 3 . 4  1 1 . 3  2 4  6  4 8 ' 3 6 '  -  1 2 3 ' 5 2 '  2 2 0  5 7 0  2 0  3 0  0 . 6 4  6 4 - 1 1 - 2 5  1 1 1 1  2 6  1 5 9  1 4 . 3  1 2 . 8  2 6  S  4 8 ' 3 6 '  -  1 2 3 ' 5 2 '  2 1 9  5 5 0  19  3 0  0 . 9 0  2 8 - 6 2 - 1 0  2 6 2 2  3 8 . 9  1 2 8  1 2 . 7  9 . 3  4 8 ' 3 6 '  -  1 2 3 ' 5 2 '  2 1 4  6 0 0  2 2  3 0  0 . 7 6  5 2 - 3 0 - 1 8  1 3 7 8  1 7 . 5  1  1 1 . 5  8 . 7  6  Mean  crown completeness  ^Percent  of species  °Determined  from  e s t i m a t e of f o r e s t  by b a s a l  trees  Plots  i n CWHa  subzone.  Plots  i n CWHb  subzone.  > 8 . 0  overstory  area contribution.  1  . 5  . 1  1  1 8  1 9  1 0 .  3  C  9 . 4  2 1  2 7  e  t o sample  9 . 8  cover.  D F = D o u g l a s - f i r , WH=western  hemlock,  Misc=other  coniferous  species.  cm DBH.  oo  49  METHODS  Overstory  sampling  Forest  overstory  (MCC) sensu  completeness" defined  cover  i s termed  Bunnell  t o include holes  within  as well  a s between  Plots  contours  a n d MCC was s y s t e m a t i c a l l y s a m p l e d  each  Bonnor  1967).  the  canopy Two  plot  using  plot.  designs  were  used  t o ph e i g h t ,  height  where  recorded  were  four  "in" trees  live  t o base  branches  were  plot  taken  only  corners  outside  recorded  samples  processes  theplot.  summarized Five  a 4 basal  and a t plot  i n theplot  Prism  integrate  with  for  area  complete  reflect  included trees acting  Plot  overstory  Point  (BAF„)  samples.  within as well  (HBLC;  the plot.  Species  acting  outside  height  t o the bole)  counted.  factor  of  stand  crown  attached  processes  slope  coverage  at breast  center.  theprism  tree  ( R o b i n s o n 1947;  of live  were  i s  a t 52 p o i n t s  additional  species, diameter  and  out along  f o r a l l t r e e s > 8.0 cm DBH i n s i d e  t r e e s were  plot.  plot  t o sample  and height  lowest  provided  < 8.0 cm DBH a n d > 20 cm t a l l  samples the  design  each  Tree  laid  10° a r c moosehorn  above  (DBH),  Trees  0.0225 h a ) were  The sample  characteristics.  were  a  crown  e t a l . (1985:181)  crowns.  within  ( 9 x 25 m,  "mean  prism at  a n d DBH o f  Samples o f within the  the plot and  a s from  angles  characteristics are  i n T a b l e 3.1. hemispherical  and analyzed  photographs  for diffuse  were  and d i r e c t  taken site  within factors  each (sensu  50  Anderson  1964a)  Bunnell  Solar  radiation  paired  Dome  solar  from  radiation  global  p a i r e d on e a c h  correction  factor  should  canopies, Because have  applied  t o both  relative diffuse  open  amounts  correction  term  below-canopy reflections  a t 1-minute diffuse  1.5 m a b o v e  Drummond's  term,  ground  (1956) outside a  developed  sensors  f o r under  would  (1977)  of brightness  results  through  under  from  made t o t h a t no  sensors  because  multiple  leaves,  a canopy  term  not affect  argued  be a p p l i e d t o d i f f u s e  radiation  correction  no c o r r e c t i o n s were  and Matt  and transmission  distribution  taken  a n d t h e same  and canopy  Hutchison  diffuse  o n CR-21  c o r r e c t i o n s f o r below-canopy  explored,  should  mounted  with  be a p p l i e d t o t h e b e l o w - c a n o p y  of radiation,  sensors.  recorded  be a p p l i e d t o s e n s o r s  appropriate  n o t been  Ltd.)  Ideally,  correction  should  um) was m o n i t o r e d  1969) s e p a r a t e d  sensors  stand.  forest  and that the  i sdifferent  from  above. One  area  by V a l e s a n d  o f samples  (Horowitz  with  and a d i f f e r e n t  sensors  averages  bands  forest  sensors.  (Lintronic  a s 3-minute  loggers  (0.3-3.0  radiation  solarimeters  Shadow  that  described  sampling  intervals.  and  t h e method  (1986).  Shortwave  data  using  pair  near  the plot  potentially located  of sensors  above  11 m a p a r t  recorded  t o determine t h e canopy. within  solar  radiation  t h e amount Two p a i r s  plots,  along  of  i n an open  radiation  of sensors  were  the centerline,  and 7 m  51  in  from  shaded  each and  density  &  Dome  Zonen  the  entire  and  Matt  150  W-nr  range  would was  have  been been  results  shortwave energy  Ross  2  of  The  open  was  quantum  (using  to  4.24  convert  to  waveband), active  but  are  Below-canopy  lower  diffuse  J  of  in  there  These than  from  the  sky  transmitted  reflected  diffuse,  and  scattered  (Reifsnyder radiation  et  al.  1971/72).  transmitted  that  was  was  The  diffuse  full-range radiation solarimeter  obtain  65%  an  of  The  diffuse  waveband 1972  2.756  photons  those  radiation  than  on  an  and  photosynthetically  values  radiation  over  less  canopy.  McCree  the  radiation  to  nm  reference  because  diffuse  that  0.4-0.7  ME/J  or  radiation.  as  Hutchison  data  diffuse  the  a  in  on  the  sensor  indicated the  cited  Had  PPFD a b o v e  in  center  curvilinear  sensors  amount  calibration  solar  (1975),  a l l diffuse  diffuse  was  flux  against  based  the  photosynthetically shortwave  calibration  overestimated.  radiation  1974  applied,  basis  1978  Hutchison  the  plot  were  W«irr .  irradiance  flux  McCartney active  the  calibrated  150  against  photon  the  exceeded  predicting of  in  sensor  samples.  were  and  to  nm)  Calibrations  linear  applied  calibrated  equation  A  quantum  photosynthetic  1-minute  (Matt  rarely  calibration  of  LI-190SB  0.4-0.7  solarimeter.  was  radiation  diffuse  solarimeters  1977). 2  LI-COR  (PPFD;  averages  The  One  monitored  radiation  3-minute  Kipp  edge.  of  is  J  of  in  Szeicz  composed the  beam  proportion  measured  of  per  are  through  direct  juE  diffuse  agreement (1974). of  diffuse  canopy, radiation  of  included  diffuse a l l  three  with  components diffuse  and i s not the true  sky radiation  Direct radiation each of  that  radiation  component  component  from  radiation.  Differences  canopy  consistent  variation between  of diffuse  global  demonstrated days  that  (ratio  monitored  clear  were  full  day, half-day  for  plots  Definition GLOBAL DIRECT DIFFUSE  PAR MCC SUMDIA BA DBH  g  global  over  sum  under t h e  little  spatial  canopies.  Differences  among  and  were  < 0.20). day.  plots  Only  used  f o r analyses  Each  plot  Because  permit  monitoring  (sunrise  to solar  was  variable  a plot noon)  Transmission  weather  during  one  monitoring was  averaged  two d a y s .  variables proportion of global solar r a d i a t i o n transmitted proportion of d i r e c t solar r a d i a t i o n transmitted proportion of d i f f u s e solar radiation transmitted ( i n c l u d e d d o w n w a r d s c a t t e r i n g o f d i r e c t beam i n tree canopies) photosynthetically active radiation as a p r o p o r t i o n o f d i f f u s e PPFD t r a n s m i t t e d mean c r o w n c o m p l e t e n e s s ( a v e r a g e o f 52 m o o s e h o r n samples and r e p o r t e d as a f r a c t i o n ) sum o f DBH (mm/225 m ) o f p l o t t r e e s £ 8.0 cm b a s a l a r e a ( m « h a " ) o f p l o t t r e e s ^ 8.0 cm diameter of tree of average basal area (quadratic mean d i a m e t e r ) number o f p l o t t r e e s « h a ~ ^ 8.0 cm 2  2  1  1  NTREES  sensors  due t o s u n f l e c k s .  sunny  by 2 f o r two c a s e s .  sampled  of  variation  one f u l l  d i d not always  multiplied  under  daily  sum o f o p e n  indicating  variable  or mostly  diffuse/open  was  the average  diffuse  plots  the diffuse  Transmission of  between  conditions clear  by d i v i d i n g  radiation  f o rat least  radiation.  by t h e d a i l y  among  sensors  by s u b t r a c t i n g  sensors  the spatial  were  open  global  was c o m p u t e d  of above-canopy  transmitted.  was o b t a i n e d  t h e two b e l o w - c a n o p y  were  was  fraction  53  SDI  - R e i n e k e ' s " s t a n d d e n s i t y i n d e x " ( R e i n e k e 1933) c o m p u t e d f r o m : SDI = N T R E E S • ( D B H / 2 5 ) • (Long 1985) BAFBA - a v e r a g e s t a n d b a s a l a r e a ( m ' h a " ) o f t r e e s £ 8.0 cm d e t e r m i n e d f r o m 5 B A F prism samples B A F T R E E S - a v e r a g e number o f t r e e s - h a " " £ 8.0 cm d e t e r m i n e d from 5 BAF prism samples BAFSDI - R e i n e k e ' s SDI d e t e r m i n e d f r o m p r i s m s a m p l i n g computed from BAFTREES*(BAFDBH /25) • 1  6  q  2  1  4  1  a  1  6  g  Anal  ys e s The  was  proportion  transmitted  regressed against  of each  radiation  stand c h a r a c t e r i s t i c s  component  using  simple — hV  linear,  second-degree  regressions. natural  error  of  were  regression  the  identified  regressions,  residuals.  best  form  components direct  For a f o r one  also  were  as having  s  ( . )r y  with  x  Ezekiel  2  r  was  2  i n the trend  the best not  in  form  necessarily  variable.  forms  regressions  standard  a n d no  variable,  independent  )  best  ( i  1970),  variable  i n the best  radiation  The  , ae  untransformed,  the lowest  and Fox  independent  (e  either  a high  g i v e n dependent  differed  nonlinear  transformed.  for a different  and g l o b a l  diffuse  variables  reciprocally  of the estimate  nonlinear the  Independent  l o g , or  regressions  p o l y n o m i a l , and  — hV  Radiation  of r e g r e s s i o n s  being different  with  from  regressions. — hV  Generally form  of  regression  independent does Saeki  the nonlinear  (1953),  form  for a l l radiation  variables  not depart  ae  i n the separate  was  ranked  the best  components a c r o s s a l l subzones.  This  form  g r e a t l y f r o m B e e r ' s law p r e s e n t e d by M o n s i - kz e , with stand c h a r a c t e r i s t i c s being  and  54  correlates  o f z, c u m u l a t i v e  leaf  area  index  (LAI).  I  also  — hV used  the e  form  intercepted give  by s t a n d  an u n r e a l i s t i c  coefficient Tests subzones SLTEST using  other  were  5  's t h a n  available  presented  within  a  factors  form,  were  This among  and the s t r e n g t h Paired used  t-tests  to test  a n d some A FORTRAN  y x  '  forms  variables  had only  modestly  tests  were n o t  subzones a r e data  of presentation (CWHa)  compared  to  of the  relationship  within  subzones and and d i r e c t  photographs  were  done  program  with  a  nonlinear  site  against  analyses  parameter-estimating  s u b r o u t i n e ( M o o r e 1984) c o m p u t e d s 's, and / ' s . 2  t h e UBC  on l i n e a r  the d i f f u s e  and DIRECT.  1976).  between  and f o r combined  stands  PAR,  manipulation  would  using  independent  manner  DIFFUSE,  derivative-free,  made  components  hemispherical  and G u i r e  analysis  in individual  from  Data  be  an a  and because  estimated  measured  with  may  form.  a n d PAR.  (CWHb)  a l l plots  were  transformed  t h e ae  a l l a r e combined.  across  Tests  the r e l a t i o n s h i p  stand  form  and equations  covariance  for a l l radiation  illustrates  areas  of regressions usually  developed  GLOBAL, DIRECT,  (Fox  with  f o r the nonlinear  Regressions  i n open  components  T h e ae  regression slopes  (Le 1971).  forms  radiation  1.0.  l o g and r e c i p r o c a l l y  larger  when  estimate  computed  these  how  characteristics.  than  of equal  program  because  of  to explore  using  MIDAS  regression  nonlinear regressions,  55  RESULTS  Proportions  transmitted  Daily are  of  tall  Below-canopy  Diffuse  21 y e a r - o l d  above  diffuse  t o 46% below  of direct  The  of diffuse  roughly  (Table  3.2).  global  global  shortwave  proportion  of diffuse  t h e canopy Variability  shortwave  energy  flux  f r o m 15%  and Matt  active  radiation  radiant  shortwave  solar  energy  t o t h e open  irradiance r a t i o s of  active  radiant  energy forest.  by t h e d e c r e a s e d  shortwave  radiation  (Table 3.2).  of d i f f u s e  was a s s o c i a t e d  o r was  found  was e v i d e n t  indiffuse  1976).  was e s t i m a t e d  22 a n d 2 9 % i n a h a r d w o o d  i n the ratio  (1975)  i n a 10-m  4.24 M E / J f o r c o n v e r s i o n ,  by t h e canopy  compared  Hardy  The i n c r e a s e i n  (Hutchison  e t a l . (1984a)  PPFD  f r o m 18  i s due t o r e f l e c t i o n a n d  radiation  shortwave  ratio  L. canopy.  diffuse  between  o f PPFD  openness of  Gay e t a l . (1971)  photosynthetically  Absorption  below  taeda  to diffuse  Baldocchi  was 7 t o 15%  ranged  of global.  photsynthetically  diffuse  M E / J PPFD  below-canopy to  beam  upon  t h e canopy  stand.  a t 2-24% by a s s u m i n g  0.1-1.02  depended  radiation  t o be 36% o f t h e t o t a l  a Pinus  scattering  below-canopy  diffuse  of the diffuse/global  of below-canopy  t h e canopy  shortwave  a n d open  below  Douglas-fir  proportion  in  1  and below  global  fluxes  radiation  an i n c r e a s e  amount  Open  7 a n d 23, r e s p e c t i v e l y )  below-canopy  reported  3.2).  22 t o 29 M J ^ m " ^ "  91% (plots  found  i n t h e open  2  from  plot.  loads  (Table  the total.  the to  radiation  summarized  ranged  AND DISCUSSION  with  PPFD  to diffuse  stand  structure.  TABLE 3.2  D a i l y sums of s o l a r r a d i a t i o n components by p l o t  Open r a d i a t i o n  Canopy r a d i a t i o n  Noon s o l a r Plot  1  e  Date  altitude C )  Global D i f f u s e Direct (Md • m-»-day " ' )  3  PPFD (Enr *d-')  Dif/G  b  0  0  Global Diffuse Direct (Md • nr-*-day- ) 1  PPFD (Em  3  PPFD/Dif )  22-23/7/84  60  28 .97  3 .08  25 .89  8 .50  0 . 106  0 . 763  0 . 546  0 .217  0 .084  0 . 153  1/8/84  58  25 .97  1 .79  24 . 19  4 .92  0 .069  1 .931  0,.644  1 .288 .  0,. 244  0,.379  19-20/8/84  54  22 .44  3 .27  19 . 18  9 .00  0 . 146  1 .642  0.. 750  0,.892  0,.632  0,.810  22/8/84  53  23 .00  3 .31  19 .69  9 . 12  0.. 144  2..082  0..946  1 .. 136  0,. 799  0..845  1/9/84  50  21 .99  2 .31  19 .68  6 . 37  0.. 105  10 . 230  1 .. 793  8 .437 .  1 .823 .  1.017  3-4/7/85  64  27..48  2 .51  24 .97  6 .92  0..091  1 .112 .  0.. 776  0.. 336  0. 079  0. 102  21  13/7/85  63  27..82  3.. 19  24..63  8 .80  0..115  7 . 701  4 .562 .  3.. 139  2.,295  0. 503  22  17/7/85  63  26..97  2..94  24..03  8 .09  0.. 109  15 . 279  5 . 283  9.,995  1 .750  0. 331  62  27.,50  2 .57 .  24 ,93 ,  7 .08  0. 094  1 .598 .  1 .462  0. 136  0. 257  0. 176  2 5  f  6  9  23  e  e  19-20/7/85  24  26/7/85  61  26 .91 .  2 .42 .  24 .49 .  6 .66  0. 090  3., 644  2 .785  0. 859  0. 747  0. 268  26  15/8/85  56  26.. 13  2..34  23..79  6 .44  0. 089  0. 908  0. 791  0. 117  0. 106  0. 134  27  17/8/85  55  25. 48  2 .09 .  23. 39  5 .76  0. 082  3 .614  2 .704  0. 911  0. 462  0. 171  a  Diffuse  p h o t o s y n t h e t i c photon f l u x  R a t i o open d i f f u s e / g l o b a l  density.  as an index of c l e a r sky, < 0.20 considered as sunny.  Average of two sensors. d  Below-canopy  r a t i o of p h o t o s y n t h e t i c photon f l u x d e n s i t y  to below-canopy  Average of monitoring two days. ^Computed from monitoring one f u l l  day and one half-day x 2.  ^Computed from monitoring one half-day x 2.  diffuse solar  irradiance  (j,E-d~').  d  There  were  correlations and  stand  between  e t a l . (1986)  of global  energy/global within  Ranges  canopy  were  0.005  between  t h e two s t a n d  a n d HBLC located  extended beam  reported  radiation  altitude  the sensors crowns  t o t h e ground) occurring higher  gambel also  i i  within  differed  dramatically of diffuse  w a s 0.18 t o 0 . 7 8 , b u t  higher  than  i n t h e CWHa.  and recorded  i n t h e CWHb Sensors (crowns  reflected  crowns.  Miller  of d i f f u s e  and global  portions  o f Populous  tremuloides  reported  canopies several  than cases  global  irradiance  exceeded  (1975)  found  73% and 31% o f d i f f u s e  the irradiance  t o t h e 5 m and ground  levels  above  direct (1967,  estimates  Nutt.  the  was 0.34 t o 1 . 8 0 . T h e  (1971)  that  1 a n d 22,  The p r o p o r t i o n o f  i n t h e CWHb p l o t s  tree  by t h e  26 a n d 7 ) , a n d  The p r o p o r t i o n  b y CWHb p l o t s  tree  (plots  (plots  Muller  transmitted  i n solar  transmitted  transmitted  i n t h e CWHa w e r e  i n t h e upper  Quercus  1 and 7).  b y CWHa p l o t s  was a b o v e  within  radiation  1969)  and  crowns  nearly  radiant  changes  of radiation  structures.  transmitted  stand were  radiation  transmitted  of tree  active  t o 0.43 f o r D I R E C T  shortwave  bases  i n t h e below-canopy  0.03 t o 0.57 f o r G L O B A L  diffuse  proportion  NTREES).  stand.  t o 0 . 2 9 f o r PAR ( p l o t s  radiation  of PPFD/diffuse  BA,  variation  energy with  of proportions  respectively), 0.01  found  shortwave  ratio  (MCC, SUMDIA,  photosynthetically  a single  forest  t h e below-canopy  characteristics  Baldocchi ratio  (p < 0 . 0 5 , n = 12) n e g a t i v e  significant  solar  t h e canopy.  where  below-canopy  i n t h e open. solar ina  Michx.  radiation 10m  tall  Hardy was  58  unthinned  Douglas-fir  Figure global  the  illustrates  and d i f f u s e  diffuse less  3.1  open  rises  above  t h e open  Number  from  found  prism  of trees  derived  prism  from  regressions  prism  samples  of saplings  larger  trees  were  variables  saplings),  tree  crown  depth  Other  studies  independent and  number  foliage  (trees worse  have  derived  of trees  as peaks i n diffuse  curve  on number  (Table 3.3).  from  plot  than  Basal  using  of trees  tree area  plot  BA a n d  d e t e r m i n e d by  Regressions  that  included  < 8.0 cm DBH a n d > 20 cm t a l l )  than  those  using  larger  the midstory  species  tree  be a u s e f u l  alone.  height, and  transmission.  r e l a t i o n s h i p s with  SDI i n t e g r a t e s  and should  of  trees  with  (number o f  composition,  single predictors  density.  stand  variables  predictors.  was p o o r e r  not reported  variable.  a r e much  a n d S D I , a n d BAFSDI  radiation  not s i g n i f i c a n t .  diameter,  and stand  are evident  SUMDIA  acceptable  samples  poor  and g l o b a l  t o be t h e b e s t  area  representing  were  diffuse  o f r a d i a t i o n on  of solar  developed  number  Stand  sampling  were  were  27 ( p r o p o r t i o n o f  curve.  MCC, p l o t  and basal  also  and canopy  relationships  transmission  measurements  the  diffuse  o f open  The below-canopy  of the regressions  characteristics  predicting  Canopy  curve.  stand  Results  plot  and sunflecks  global  Radiation-forest  obtained  1.29).  global  below-canopy  a time  r a d i a t i o n f o rp l o t  transmitted  than  stand.  size,  SDI a s a n  basal  area,  integrator of tree  1000-1  0400  0800  1200  1600  2000  TRUE SOLAR TIME  F i g u r e 3.1. Solar irradiance as a f u n c t i o n of 27. Open g l o b a l (•) and diffuse (•••••••), (o) and diffuse (•).  time for plot canopy global  TABLE  3.3  R e g r e s s i o n c o e f f i c i e n t s of e q u a t i o n s p r e d i c t i n g s o l a r r a d i a t i o n components from f o r e s t s t a n d c h a r a c t e r i s t i c s . Sample s i z e s : CWHa=6, CWHb=6. E q u a t i o n s s i g n i f i c a n t at  Y  p £ 0.05.  Subzone  R e g r e s s i o n e q u a t i o n : Y «= ae  a  X  b  /  2  5 yx  GLOBAL:  DIFFUSE:  0.0282  0.96 0.97 0.96  0.0363 0.0324 0.0364  5.1702 7.2250E-4 4.8465E-3 4.5990E-3  0.97 0.85 0.86 0.96  0.0366 0.0867 0.0830 0.0449  2.3547 0.9792 1.1656 1.3514  4.1668 4.7260E-4 3.7187E-3 3.7443E-3  0.86 0.85 0.88 0.84  0.0701 0,0725 0.0654 0.0761  MCC SUMDIA SDI  1 .5923 0.9120 1.1061  2.1929 1.8875E-4 1.4430E-3  0.85 0.72 0.70  0.0917 0.1255 0.1292  BAFSDI  N.S.  0.46  0.1732  2.4109  SUMDIA SDI BAFSDI  0.8657 1.2643 6.8557  4.8273E-4 3.4154E-3 6.2906E-3  CWHb  MCC SUMDIA SDI BAFSDI  4.8879 1.9707 1.4843 1.5890  Both  MCC SUMDIA SDI BAFSDI  CWHa  4.7957  6  4.8886 2.6581 2.4254 2.8328  2.2770 2.2341E-4 1.4755E-3 1.8436E-3  0.78 0.94 0.92 0.84  0.2848  CWHa  MCC SUMDIA SDI BAFSDI  3.3782 0.9725 1.5134 12.0179  6.0002 6.2694E-4 4.2891E-3 7.7863E-3  0. 98 0. 98 0. 98 0. 98  0. 0226 0. 0283 0. 0245 0. 0256  CWHb  MCC SUMDIA SDI BAFSDI  14.2767 1 .0 1.0 1.0  8.4899 6.6399E-4 5.3104E-3 4.8240E-3  0. 99 0.70 0. 77 0.86  0.0202 0. 0970 0. 0851 0. 0659  Both  MCC SUMDIA SDI BAFSDI  3.6343 1.2592 1 .5885  1.1168  5.8766 7.7261E-4 6.0770E-3 4.3206E-3  0. 93 0. 85 0. 74 0.63  0.0432 0. 0623 0. 0812 0. 0972  CWHa  MCC SUMDIA SDI BAFSDI  1 .1995 0. 4763 0. 6697 2. 621 1  4. 1655 3. 9642E- 4 2. 8915E- 3 5. 1752E- 3  0. 95 0. 95 0. 96 0.95  0. 0253 0. 0251 0. 0237 0. 0253  CWHb  MCC SUMDIA SDI BAFSDI  0. 9428 0.6171 0. 4694 0. 5200  3. 1579 4.6447E- 4 2. 8252E- 3 2. 9551E- 3  0. 67 0. 92 0.88 0. 84  0. 0635 0.0302 0. 0384 0. 0447  Both  MCC SUMDIA SDI BAFSDI  0. 9927 0. 5100 0.4930 0. 5551  3. 4685 4. 0491E- 4 2. 6627E- 3 2. 8154E- 3  0. 79 0. 93 0.87 0. 74  0. 0473 0. 0264 0. 0367 0.0522  "Exponent f o r power of .10. ^Equation  0  MCC SUMDIA SDI BAFSDI  CWHb  PAR:  0.98  MCC  CWHa  not s i g n i f i c a n t  (p > 0.05).  0.1519 0.1763 0.2435  61  Some the  of  the  CWHb) g a v e  equations  peculiar  (e.g.,  results  DIRECT  when  versus  SUMDIA,  SDI  extrapolated outside  in the  — AY  range  of  the  data  using  the  ae  form.  These  equations  were  — hV recalculated  as  intercept  1.0.  within  of  the  The  e  and The  range  of  negative  Table  3.3)  are  Solid  lines  illustrated  f o r DIFFUSE  DIRECT  PAR  subzones  The  line  i n the  is a  general  of  and  several studies  methods (1964)  were  used  presented  and  a  stands  used  to  was  i n the  sample  composition  used  estimate  graph  spp.  with  curves  a  overstory  or  cover  GLOBAL, in data.  from  from  the  Miller results  various  Vezina  and  transmission  i n Pinus  that  of  to  1964b),  1948),  MCC.  in  developed  closure.  related  (Anderson  for  i n which  moosehorn  be  developed  derived  than  from  f u n c t i o n of  i s reproduced  predicting  higher may  (equations  a  an  them.  i s f o r combined  stands  crown  (Wellner  forest  line  with  a p p l i c a b l e only  i n F i g . 3.2  relationship  also  radiation  species to  solid  relationship  Lamb,  as  3.3  develop  relationships  estimate  closure estimated that  the  to  equations  curves  the  i n Pinus to  used  are  relationships  global  crown  Differences  data  represent  represent  in Table  presented  i n F i g . 3.2  Dotted  individual  (1959)  stand  exponential  subzones.  dashed  presented  equations  the  individual and  are  banksi  Pech from ana  Miller  (1959).  different  methods  stand  s t r u c t u r e or  differences  in  ( B u n n e l l and  methods Vales  1986). Indices computed  for  components  of  determination  seven  using  independent  the  (/',  analagous  to  2  r)  were  variables predicting radiation — hV r e g r e s s i o n form e (Table 3.4). This  62  0.6-1  0.6  < oo 2 2  o  UJ  gc Q Z  0.4  o g  0 2  2  o 0.0  — i — i — i — i — i — i — i  0.3 0.4  0.6 0.6 0.7 0.8 0.9  C  1.0  1  -i 0.3 0.4  1  1  r — r 1  0.6 0.6 0.7 0.8 0.9  1.0  2.0-1  0.0-1  1  0.3 0.4  1  1  1  1  1 1  0.6 0.6 0.7 0.8 0.9  t.O  MEAN CROWN COMPLETENESS  o-i  1  0.3 0.4  1  1  1  1  1 1  0.6 0.6 0.7 0.8 0.9  1.0  MEAN CROWN COMPLETENESS  F i g u r e 3.2. R e l a t i o n s h i p s b e t w e e n t h e p r o p o r t i o n s o f s o l a r radiation c o m p o n e n t s t r a n s m i t t e d a n d mean c r o w n completeness. Dotted l i n e s i n a, b, a n d d a r e relationships forindividual subzones. Dashed l i n e i n a i s f r o m M i l l e r ( 1 9 5 9 ) . CWHa ( • ) , CWHb (ffl).  63 TABLE  3.4  I n d i c e s o f d e t e r m i n a t i o n (/*) f r o m r e g r e s s i o n s p r e d i c t i n g r a d i a t i o n components from f o r e s t s t a n d characteristics  X  Global  using  Direct  the regression  form  Di f f u s e  PAR  Y = e Avg.  rank*  CWHa: MCC  0.854  0.836  0.736  0.945  3.0  SUMDIA  0.951  0.975  0.707  0.791  2.8  NTREES  0.892  0.946  0.414  0.779  5.0  BA  0.913  0.906  0.647  0.941  3.3  SDI  0.955  0.954  0.694  0.910  2.3  BAFBA  0.706  0.698  0.427  0.821  5.5  BAFSDI  0.696  0.685  0.393  0.863  6.3  MCC  0.689  0.663  0.010  0.666  6.0*  SUMDIA  0.802  0.702  0.021  0.882  3.7  NTREES  0.751  0.659  0.007  0.876  5.0  BA  0.834  0.779  0.034  0.678  3.7  SDI  0.842  0.771  0.030  0.738  3.3  BAFBA  0.909  0.865  0.010  0.599  3.3  BAFSDI  0.910  0.863  0.010  0.653  3.0  MCC  0.729  0.751  0. 142  0.787  4.0*  SUMDIA  0.850  0.841  0.300  0.766  2.3  NTREES  0.534  0.646  0.221  0.086  6.0  BA  0.789  0.569  0.369  0.504  4.0  SDI  0.874  0.733  0.345  0.742  2.3  BAFBA  0.770  0.547  0.312  0.416  5.3  BAFSDI  0.809  0.631  0.280  0.600  4.3  components:  1=lowest,  6=highest.  CWHb:  Combined:  fl  Average  ^Excludes  rank  of s  di ffuse  across  yx radiat ion  rank.  3  64  form 3.3  has a regression  intercept  a n d 3.4 i n d i c a t e s  intercepts  i n Table  applicable  outside  the importance  3.3.  of the proportion  canopy.  Stand  integrated  HBLC's  there  (Tables  of diffuse  variables  Determining  MCC  often  made  being  and  therefore  MCC  measurements  the  direct  MCC  high  tree  than  also  were  within  of  plot  and t a l l  or prism  a n d PAR.  rather  of trees  variables  MCC was p o o r e r  with  overstory  b y MCC b e c a u s e  LAI.  of the v a r i a t i o n i n  sensors.  crowns  no  closely  measurements  than  cover  made a t t h e h e i g h t  of radiation  i s  variation  DIFFUSE  crowns  realistic  law r e l a t i o n s h i p and  more  at predicting  for  2  /  a  there  I n t h e CWHb, h o w e v e r ,  within  accounted  below  them,  a s i n t h e CWHa. of,  Extensive  or s l i g h t l y scatter of  i n t h e CWHb p l o t s  stand  structure  was  affected  measurements. There  were  slight  variables.  DIFFUSE  and  t o prism  BA t h a n  Much would in  when  inter-plot  radiation  be more  and give  as correlates  for  not representing  beam  would  i n t h e CWHb was d i f f i c u l t  the height  poorly  served  was e x t e n s i v e  3.3 a n d 3 . 4 ) .  plot  above  Beer's  ( C W H a ) , MCC a c c o u n t e d  transmission  than  transmitted  the theoretical that  form  with  of Tables  of the equation  of the data  characteristics  factors  Where  The e  t h e range  estimate  approximated  of 1 and comparison  of the diffuse be r e f l e c t e d  theplot  would  differences  was b e t t e r samples  between  related  S D I a n d BA  radiation  plot  and prism  to the plot (Tables  was s c a t t e r e d  downward  by t r e e  crowns.  therefore  be more  related  s a m p l e s SDI  3.3 a n d  direct  beam  Diffuse  3.4). which  radiation  t o overhead  canopy  65  on  sunny  days  GLOBAL  and  trees  Where  were  prism  there  spacing,  the  stand  Miller closure area  (1959)  c a n be p r e d i c t e d  and  tree  al.  (1969),  density. a n d my  a s SUMDIA Miller  radiation  did well is  plot  samples  as other  related  were  that  from  SUMDIA  more  Because s i z e and  accurately. i n tree  size  representative  was s u p e r i o r  of transmission. tree  1981),  diameters  SUMDIA  Jackson  and Harper  results  (Table  also  of  would  t o crown  Because  (e.g.,  (1955),  leaf  Gholz e t  integrate  t r e e LAI  Perry et  3.4) i n d i c a t e d  of transmission,  believed  predictor  was a l s o  not predict  the stand  GLOBAL  that  although  basal notas  SDI.  (1959)  an adequate  t o prism  t h e same  inter-plot variation  predictor  or  samples.  of r e l a t i v e l y  characterized  believed  1976; S p i t t l e h o u s e  of  were  a n d BA a s a m e a s u r e  good  than  sensors.  al.  not  related  stand  the  i sa useful  data  tree  t o prism  however,  area  combined  to plot  better  sampling  near  related  the plot.  I n t h e CWHb, h o w e v e r ,  was e x t e n s i v e  and  better  outside  3.3 a n d 3 . 4 ) .  i n t h e CWHb  spacing,  to trees  i n t h e CWHa a n d f o r  (Tables  DIRECT  than  a n d DIRECT were  measurements samples  i n summer  to tree  crown  of transmission  a function  transmission  stand  that  o f crown  that  integrate  tree  also  integrate  crown  size  and density. and density  depth.  because depth.  of d i r e c t and global  c h a r a c t e r i s t i c s (Table size  closure  3.4).  alone  was  attenuation MCC  alone  r a d i a t i o n as Crown  depth  Stand c h a r a c t e r i s t i c s such  a s SUMDIA  o r SDI  66  Studies radiation 1974; of a  within  Hardy  crown  coniferous  variable  or  because  MCC. were  estimate  overstory  moderately  wide  and  1982;  Zamora  1986)  and  not  moosehorn,  was  an  moosehorn  also  made  but  predict  crown  and  Bunnell  1986;  nature  of  the  understory  abundance  be  influenced  by  multiple  of  integrator  sampled  predicting  little  in  zero.  of  (Pyke  Vales  surface.  the  a  the The  overstory  transmission.  10° and  The  and  i s an  accurate  (Bunnell  and  Vales  1986)  (Chapter  with  and  bias  radiation  Any  crowns  Bunnell  intercepting  adequately  variable  of  better  As  stand  instrument  some d e p t h  a  influence  of  from  include  as  Jarvis  transmission  depth  an  of  attention.  effects  different  using  and the  little  confounding  serve  shows  Norman  will  however,  acceptable  not  for  distribution  1982)  received  did  the  cover  Vales  three-dimensional  (e.g.,  significantly  angle  possibly  arc  has  depth of  vertical  Knoerr  Coefficients  regressions of  and  stands  crown  the  stands  Sinclair  among  probably  density  demonstrated  1975;  depth  single  well,  have  predictor  which  would  4).  — h V  Coefficient pattern the  among  different  (rate  of  b  in  the  e  overstory  variables  radiation  components.  extinction  fastest)  followed  by  global,  Canopies  absorbed  direct,  direct  radiation  global  also  the  slowest.  equations  diffuse  Direct  and  PPFD  transmitted  contains  for  diffuse  was  for  rate  of  Slopes  then  diffuse  (Table  3.2).  greater  was  also  than  which  a  consistent  extinction  were  transmission  radiation  radiation  showed  of  of  steepest PPFD,  radiation. The  drop  global was  converted  in because  attenuated to  diffuse  67  radiation direct  which  caused  radiation  the steep  and shallow  slope  slope  for  attenuation of  f o re n r i c h e d  diffuse  radiat ion.  Equation  differences  Linear and  regression  reciprocally  differ  All  with  regressions  between  independent were  not  of subzone  tests  Si t e fact  between  factor and than  Anderson  transmission a l l plots  t h e measured  between  predicting  of stand  found  factor  and measured  higher  Vales  differences  PAR a n d  differing The t e s t s o f  structure  transmitted  and  transmission  between  on a l l b u t 2 p l o t s .  site  between  factor  was  lower  than  PAR  the direct  radiation site  excluding  site  radiation  and higher  the direct  Tests  1986) w i t h  the diffuse  of diffuse  of direct  11),a l t h o u g h  (estimated  subzones and  of diffuse  was f o u n d  factor  and Bunnell  11). The d i f f u s e  No d i f f e r e n c e  (p > 0 . 0 5 , n =  site  3.5) w i t h i n  transmission  3.5).  much  1964a;  (Table  (p < 0 . 0 5 , n =  (Table  plots  differed  of regression.  classes  p >  structures;  sometimes  of the d i f f u s e  and the proportions  PAR  DIFFUSE  d i d not  effects.  comparisons  photographically;  including  broad  from l o g  ors  Paired  measured  (stand  equivocal,  o r form  DIRECT  variables  of regressions  were  variable  equations  subzones  predicting  Tests  subzones  predicting  independent  between  (p < 0 . 0 5 ) .  subzones GLOBAL  transformed  significantly  0.05).  equations  site  over  a l l  factor  was  plots  7 and  TABLE  3.5  Comparison of d i f f u s e and d i r e c t s i t e factors o b t a i n e d from h e m i s p h e r i c a l photographs t o measured p r o p o r t i o n s of s o l a r r a d i a t i o n components t r a n s m i t t e d  Diffuse*  Diffuse* PAR  Direct site factor  Direct*  Plot  Diffuse site factor  1  0.092  0.177  0.010  0.139  0.008  2  0.119  0.360  0.050  0.180  0.053  5  0. 1 26  0.260  0.069  0.1 53  0.046  6  0.111  0.286  0.088  0.193  0.058  7  0.318  0.775  0.286  0.258  0.429  21  0.239  1 .429  0.261  0.170  0.127  22  0.248  1 .799  0.216  0. 185  0.416  23  0.112  0.569  0.036  0.089  0.006  24  0.150  1 . 1 53  0.112  0.114  0.035  26  0.072  0.338  0.016  0.151  0.005  27  0. 108  1 .293  0.080  0. 140  0.039  0  ^Estimated Measured  from  hemispherical  proportion  of solar  0  photographs. radiation  transmitted.  69  22  found  and  22  significant  were  the  heterogeneous Lower  most  than  of  sensor  points  and  direct  site  the  just  for  also  done  using  path  only  for  the  the  was  higher  Hourly  comparisons  were  direct  site  factor  noon.  Comparisons  were of  also  of  being  21  direct  site  open)  of  photos  assumed But  (0.5°),  reducing  the  assessment  of  consideration  factor was  than  the  An  obtained sampled.  for  for the  eight  calculated  transmitted  to  beam  in  the  these  consequently  to  site the  <  the  full  factor  solar  same  period  proportion because  i t to  more  may  be  of  diffuse  seen  on of  than  direct  A  solar  8-hour  low  smaller  on  measured  intensity  relationship  The  and  The  were  the  days  0.05).  been  that  diffusing  site  was  proportions  0.05).  were  This  centered  converting  canopy  gaps  <  the  factor  the  have  transmitted.  direct given  may  and  transmit  of  (p  from  for  site  for  analysis  factor  hours  direct  rather  summer.  (p  a  The  entire  measured  was  be  the  hourly  proportion  is  radiation.  and  gaps  7  photograph  September  sampled  many  diameter  23  site  direct  Small  radiation.  to  may  radiation.  direct  between  radiation. were  direct  significantly different  the  from  the  transmitted  Plots  spatially  between  radiation  scattering  of  for  radiation  used  9).  radiation  different  significantly different  direct  =  more  direct  March  sampled  (more  transmitted  were  variability  when  was  and  n  0.05,  plots.  for  days  factor  proportions  was  days  radiation  plots  locations  factor  <  (p  transmission  spatial  than  when  open  denser  measured  result  differences  the direct  the  beam,  sun's and  accurate made  among  if tree  height,  70  sun  diameter,  Muller  and distance  1971:3)  Anderson agreement Bunnell  from  and Vales  (1986) shall  factors  was p o o r e r  methods  used  (Chapter more  closely  radiation  related  during  sampled  year  from  the of  direct  beam  diffuse  photographic error  diffuse  sky r a d i a t i o n .  factor  estimation  f o r one p a r t i c u l a r  The  t h e time  i s an a v e r a g e a  by d i r e c t  originated  than  diffuse  here  time  on sunny  days  Because from  the diffuse  site  found  beam  proportion  (1964a)  over  standard  a short  radiation  Lower  than  Differences  of the d i f f u s e day.  solar  assuming  f o ronly  Anderson  was  used.  t h e measured  was h i g h e r  data  factors.  other  distribution.  radiation  radiation,  transmitted  I  enrichment  of sampling  other  abundance  of  site  site  by t r a n s m i s s i o n o f  distribution. sampled  and  from  salal  or direct  site  sky brightness  abundance  transmission  factors  between  Additional  that  sky radiation  being  or because  below-canopy  cover.  of the year  diffuse  of diffuse  a different  radiation for  plots  because  scattering, with  (1964a)  sky b r i g h t n e s s  arise  period,  time  and i s f o r d i f f u s e  overcast may  a short  transmission.  developed  may b e r e g u l a t e d  by t h e s i t e  Anderson's a  diffuse  found  the relationship  21) i n d i c a t e d  t o measured  (e.g.,  (1981)  canopy  understory  overstory  to either  however,  and Perttu  that  Pursh)  gap  location.  relationships  t o estimate  than  understory,  year  than  of a canopy  and measured  found  on  4 and Appendix  radiation  of  and L i n d r o t h  estimated  (Gaultheria  salal  the photograph  (1964a)  between  and angle  the scatter of  site  diffuse  site  stated factor  much o f  factor  that c a n be i n  factors  may  71  also  be a t t r i b u t e d  to photographs  underexposed  to obtain  overestimate  of the d i r e c t  Generality A this  of  Diffuse each  and d i r e c t  affected  discuss  stand,  by  by  a  among  that  forest  developed that  are affected  a n d among  are transmitted  transmitted.  by c l o u d s  the discussion  stands.  to a f f e c t the  I will  direct,  and s o l a r  by  differently, are  and i n t e r a c t  of d i f f u s e ,  of the  and  briefly  global  altitude  to differences  within  a  i n stand  stands.  of d i f f u s e  global  direct  by w e a t h e r .  beam  scattering.  available  i s reduced  Proportions  of d i f f u s e  radiation  direct-beam  scattering  should  The p r o p o r t i o n s  of d i f f u s e  radiation With in  i s less  i n canopies.  transmitted  decrease  transmitted  radiation  and there  for scattering  (1971/72)  Relationships  of d i f f u s e  t h e amount o f d i r e c t  radiation  et a l .  sky r a d i a t i o n  on t r a n s m i s s i o n  below-canopy  cloudiness,  and R e i f s n y d e r  affected  a r e based  radiation  cloudiness.  the  presented i n  proportions  transmitted  radiation  i s n o t much  above-canopy  not explain  of equations  o f how  within  (1970) c o n c l u d e d  here  increasing  does  factor.  structure,  the fraction  included  direct  stand  are affected  Anderson assumed  radiation  transmission  and a p p l y  structure  altitude  of global  how  radiation  components  and s o l a r  proportion  site  understanding  radiation  cloudiness  but t h i s  on t h e g e n e r a l i t y  requires  different  deliberately  equations  discussion  study  contrast,  being  with  that  include  increasing  radiation  transmitted  72  by  canopies  plots  that  different  were  of  clear  scattered Muller  open the  (1971)  source  a Pinus  to  from  much  scattering  i s dependent et a l .  affect  direct  canopy  openings above  beam  disk  1980).  scattering  i n the solar  the horizon  below  (Hutchison  crowns.  Scattered  et al.'s factor  sky radiation  accounted  canopies  f o r 50  in this  penetration  days,  was  radiation  site  beam  study. comes  transmission of  near  the path  of the  solar  altitude  will  (Hutchison path  canopies  of diffuse  Changing  radiation  conditions,  diffuse  openings  1976).  was g r e a t e s t i n  i n Reifsnyder  on c l e a r  on c a n o p y  diffuse  diffuse  of  of d i r e c t  clear-sky  sky r a d i a t i o n  by t h e  compared t o  and Matt  by t r e e  below  2  the influence  scattering  of d i r e c t  radiation  of the diffuse  (Hutchison  angle  canopy  the proportion  10° o f t h e s o l a r  diffuse sun  then  days  scatter  of the t o t a l  Ait.  92% o f t h e d i f f u s e  Because  (Hutchison  radiation  with  i n the proportion  demonstrating  under  days  clouds.  on o v e r c a s t  I f the estimated  reflects  transmitted,  a decrease  radiation  one-third  study.  accurately  of diffuse  resinosa  increasing  and that  on 8 o f 8  transmitted  of the total  that  cover  consecutive  downward  one-half  of direct  was a b o u t  (1971/72)  that  He c o n c l u d e d  scattering  below  stand,  spp. canopies,  primary  direct  found  radiation  reported  cloud  Proportions  transmitted  beam  was a b o u t  Pinus  2 full  1.0 w i t h  (1977)  i n a hardwood  stands.  from  over  than  radiation  direct  radiation below  less  and Matt  days  increased  conditions.  were  diffuse  with  sampled  cloud  CWHb p l o t s Hutchison  decreased  and Matt  generally et a l .  1976)  increase  and  with  1980; L i n d r o t h a n d  73  Perttu  1981).  canopies will on  not accurately  cloudy  days  information  with  increasing  increased  lack  three most  direct  was  open  radiation  on  i n which  was  factor  decreasing  solar  of eight  a n d may  site  would  transmitted  (Anderson  frequency  decreased  from  to horizon.  frequency  i n the solar  similar may of  at a l l solar  affect  a l l plots  the predictive  intercept altitude Jarvis under  would (/3)  effects  angles,  similarily  equations  change;  1 9 7 6 ) may  over  range  however, of d i r e c t of solar  were t h e  For  clear  the proportion  sampled  gap gap  was  in solar  altitude  skies.  The  slope  t h e same, b u t t h e  to account (e.g.,  the accuracy  at another  clouds,  a  term  The  If relative  clear  remain  errors  1970) p r o v i d e d  a change  equations  improve  of transmission  characteristics  a  sensor. decreased  a l l plots  under  would  Adding  sky c o n d i t i o n s  of increasing  relationship  then  to the regression  et a l . clear  among  The  be due t o  decrease  radiation  path  found  increased  factors.  direct  zenith  I  clouds  (global-diffuse)  of d i r e c t  direct  of  plots.  f o r the shaded  altitude  radiation  transmitted  calculated  transmission  and had the h i g h e s t  of d i f f u s e  the e f f e c t s  radiation five  tree  presented  transmitted.  unexpected  radiation  of c o r r e c t i o n  plots  days, of  transmission  t h e way  and  cloudiness  in  of the year.  i s available.on  of d i r e c t  beam  and the equations  times  of d i r e c t  the proportion  of d i r e c t  transmission  and a t other  the proportion  scattering  increases,  predict  that  in  i s less  as cloudiness  Little on  There  time  for solar i  a  of  e  X  ^  c  s  c  '  3  complicate  radiation altitudes.  ^  predictions  of the year.  would  1  to  The the  stand  74  Miller of  global  (1959)  Zavitkovski  opposite of  increasing  cloud  days.  global  angles, may  then  On  radiation, the  solar  a  on  but  other  path,  decrease  cloudiness  in  transmitted  to  extensive  days  would  be  global  may  of  cloudy  the  days  increase  direct sun  then,  because  to  of  the  path sunny  gaps  radiation  transmits  in areas  of  beam tree  the  proportion  plot of  on  cloudy of  ratios  cloudy  direct  other  and  During  than  plots  the  increasing  direct In  the  most clear  proportion  winter  because  increased global  in  showing  r a d i a t i o n on  r a d i a t i o n and  of  direct  radiation  proportion  days  and  overhead  little  transmitted..  and  days,  sun  transmitted  Those  direct  the  at a l l  dense  global  of  r a d i a t i o n with  minimal  crowns  a  canopy  days.  of  diffuse  are  of  global  proportion  diffuse  the  proportion  decrease.  scattering  the  through  global  with on  if a  on  penetration  of  of  in  the  than  r a d i a t i o n on  s c a t t e r i n g of  elevation,  angle  gaps  (1977)  plots  lower  hand,  the  reduced  transmitted  affect  Matt  increased  with  proportion  plots  8  plot  highest  open  of  in a  increase  the  are  direct  transmission  had  5  days  gaps  then  may  in  proportion  increasing and  i r r a d i a n c e was  would  there  proportion  has  that  the  with  Hutchison  is transmitted  cloudy  the  transmitted  If  light  the  decrease  canopy.  though  appreciable  days  and  found  structure  transmitted.  cloudy  where  radiation transmitted  cover,  Stand  transmitting on  global  I  studies increased  (1974)  results.  proportion  sunny  several  radiation transmitted  cloudiness. found  cited  with  low  solar  downward of  the  acute  albedo.  transmitted  radiation is entering  of  through  On may many  75  canopy  gaps.  direct  radiation during  transmitted Perttu  with  stand.  change  during  with  that  decreasing  Slaughter  i n a Picea  transmitting  the proportion  solar  with  mariana  L i n d r o t h and  of global  radiation  i n a Pinus  altitude  though,  appreciable  of global  increasing clouds.  transmission  (1983),  i n transmission  altitude  found  syl  vest ri s  no c o n s i s t e n t  changes  i n cloud  (Mill.)  B.S.P.  cover  stand  or solar sampled  May-September.  In diffuse  summary,  different  affected  would times  proportion  change  p r e d i c t i n g the proportion of  with  that  by w e a t h e r  Equations  conditions  Adding  a term  path  t o account  Equations  predicting transmission  structure.  cloud  cover,  would  and a r e h i g h l y  i n the solar  by t h e p r o p o r t i o n s  beam  among  stands  be  dependent  upon  angle  i frelative  gap  i s assumed  may  t h e same.  radiation are  and d i r e c t  and i n t e r a c t i o n  likely  forsolar  of g l o b a l  of diffuse  and a t  predicting the  the generality of the equations,  transmitted,  direct  radiation transmitted  frequency  affected  include  increasing cloudiness  of the year.  of direct  altitude.  broaden  the equations  radiation transmitted  scattering  solar  showed  canopy  summer,  may d e c r e a s e  (1981)  decreased L.  B u t f o r t h e same  with  radiation  stand  CONCLUSION  Although components different  equations  may  not  sky  demonstrate  sum  are as  of  has  been  used  variation  once  a  10°  i s an HBLC's  tree  not  angle  are  active  Equations  predicting  structure  resulting  in  equivocal  for  the  were  flux  affected  by  has  measured  merit  index  that  transmission.  and  is  of  inter-plot  useful  for  of  of  cover  global  diffuse  as  and  measured direct  predicted  using  sum  of  radiation  of  index.  classes  differences. of  use  index  overstory  transmission  transmission photon  broad  the  density  Extinction  transmission  two  suggesting  sky  diffuse  i s best  density  affected  equation  photosynthetic components  stand  law  stand  radiation  sensors, and  and  specific  integrator  Extinction  Beer's  or  good  using  do  solar  predict  radiation.  predicted  following  to  under  of  easily  diffuse  moosehorn.  between  i s an  above  of  studies  stand  apparently  diameters  differ  Stand  best  and  used  they  under  transmission  (1933)  radiation  transmission  period  other  of  when  altitudes,  between  predict  transmission  is  radiation  to  estimates  time  of  transmission  solar  previously  photosynthetically  by  given  variable,  cover  radiation  or  Reineke's  Overstory  predicting  a  results  diameters  predictor not  for  The  confirmed. a  accurate  relationships  characteristics  of  give  conditions,  the  conditions.  predicting  global  density,  of  direct of  diffuse  Equation and in  transmission  stand  did  structure. radiation  differnces  were  diffuse  part, of  because diffuse  these  two  radiation.  Hemispherical diffuse, direct  diffuse  radiation  proportions  photographic  photosynthetic were  estimates photon  significantly  transmitted.  of  flux  transmission density,  different  from  of  and  measured  REFERENCES  A n d e r s o n , M.C. 1964a. 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T r a n s m i s s i o n o f i n s o l a t i o n t h r o u g h f o r e s t c a n o p y , a s i t a f f e c t s t h e m e l t i n g o f snow. Schweiz. A n s t . f o r s t l . Versuchswesen. 35:57-79.  pine Mitt.  M i l l e r , P.C. 1967. L e a f t e m p e r a t u r e s , l e a f o r i e n t a t i o n a n d e n e r g y e x c h a n g e i n q u a k i n g a s p e n (Populous tremuloides) and g a m b e l l ' s o a k (Quercus gambellii) in central Colorado. Oecol. Plant. 2:241-270. M i l l e r , P.C. 1969. S o l a r r a d i a t i o n p r o f i l e s i n openings i n c a n o p i e s o f aspen a n d o a k . S c i e n c e . 164:308-309.  81  M o n s i , M. a n d T . S a e k i . 1 9 5 3 . U b e r d e n L i c h t f a k t o r i n d e n P f l a n z e n g e s e l l s c h a f t e n und s e i n e Bedeutung f u r d i e S t o f f p r o d u k t i o n . J a p . J . B o t . 14:22-52. M o o r e , C . 1 9 8 4 . UBC C U R V E . C o m p u t i n g C e n t r e . B r i t i s h Columbia, V a n c o u v e r , B.C.  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M e t e o r o l . 9:21-37.  R e i n e k e , L.H. 1933. P e r f e c t i n g a s t a n d d e n s i t y index f o r even-aged f o r e s t s . J . A g r i c . R e s . 46:627-638. R o b i n s o n , M.W. 1 9 4 7 . An i n s t r u m e n t t o m e a s u r e c o v e r . F o r . Chron. 23:222-225.  forest  crown  Ross, J . 1975. R a d i a t i v e t r a n s f e r i n p l a n t c o m m u n i t i e s , p p . 13-55 in: J.L. Monteith (ed.). Vegetation and the A t m o s p h e r e . V o l . 1. P r i n c i p l e s . A c a d e m i c P r e s s . New Y o r k , NY. Ross, J . 1981. The r a d i a t i o n regime a n d a r c h i t e c t u r e s t a n d s . D r . W. J u n k , P u b l . T h e H a g u e .  of plant  Sinclair, T.R. a n d K.R. K n o e r r . 1982. D i s t r i b u t i o n of p h o t o s y n t h e t i c a l l y a c t i v e r a d i a t i o n i n the canopy of l o b l o l l y p i n e p l a n t a t i o n . J . A p p l . E c o l . 19:183-191. S l a u g h t e r , C.W. 1983. Summer s h o r t w a v e s o l a r s u b a r c t i c f o r e s t s i t e . Can. J . F o r . Res.  r a d i a t i o n at 13:740-746.  S p i t t l e h o u s e , D.L. 1981. M e a s u r i n g and modelling evapotranspiration f r o m D o u g l a s - f i r s t a n d s . Ph.D. U n i v e r s i t y of B r i t i s h Columbia, V a n c o u v e r , B.C. S z e i c z , G. 1974. Solar Ecol. 11:617-636.  radiation  for  plant  growth.  V a l e s , D.J.. a n d F . L . Bunnell. 1986. Comparison of estimating f o r e s t overstory cover I. Observer p r e c i s i o n . Can. J . F o r . Res. (submitted).  J.  a  a  thesis.  Appl.  methods effects  for and  V e z i n a , P.E. a n d GY P e c h . 1964. Solar r a d i a t i o n beneath c o n i f e r c a n o p i e s i n r e l a t i o n to crown c l o s u r e . F o r . S c i . 10:443-451. Wellner, CA. in mature 46:16-19.  1948. Light i n t e n s i t y r e l a t e d to s t a n d s of the w e s t e r n w h i t e p i n e  stand type.  density J. For.  Z a v i t k o v s k i , J . 1974. S o l a r r a d i a t i o n measurements i n the E n t e r p r i s e R a d i a t i o n F o r e s t , p p . 3 3 - 4 5 in: T.D. Rudolph ( e d ) . The E n t e r p r i s e , W i s c o n s i n , R a d i a t i o n Forest. U.S. A.E.C. TID-26113. Z a v i t k o v s k i , J . 1976. Ground v e g e t a t i o n , biomass, production, and e f f i c i e n c y of e n e r g y u t i l i z a t i o n i n some n o r t h e r n W i s c o n s i n f o r e s t e c o s y s t e m s . E c o l o g y . 57:694-706.  CHAPTER  4.  RELATIONSHIPS  TRANSMISSION  OF  SOLAR  OF  SALAL  (GAULTHERIA  RADIATION  SHALLON)  THROUGH F O R E S T  TO  CANOPIES  INTRODUCTION  Salal species  {Gaultheria  widely  America.  little  (1961)  found  forest  canopy  on  distributed  Despite  forests,  shall  Pursh)  along  the  work  that  has  been  done  photosynthesis  was  16%  of  those  of  plants  was  an  concluded  that  salal  sun  p l a n t s were  shade  cover  was  plants.  photosynthetically canopies. competes et  F o r e s t e r s are with  a l . 1980) Salal  in  open  (Swank  trees and  has  areas 1972),  are  leaves.  Because  related  with to  for s o i l  numerous  in  leaves  shade  are  salal  of  short  taller,  responds  forest  to  Swank  a  (1972)  moisture  reported  coniferous  salal  (Tan  that  et  a l . 1977; et  growth  1985).  ( V a l e s and  a l .  and  Shoots thick  under  darker  openings  (Long  1986).  with  Shoots  Bunnell  Black  forms. 2),  larger,  small  forest  because i t  (Chapter  with  stocking  Sabhasr  and  greater  (Weetman  (Kelliher  increased vigor levels  with  different  and  open  species.  below  nutrients  coastal  diffuse  moisture  distinctly  small  and  concerned  North  ecology.  a l . (1979)  to  radiation  possibly  two  canopies  canopy  active  related  in  of  p l a n t s under  under  et  understory  Coast  i n the  intolerant  Stanek  positively  salal  salal  salal  salal  Pacific  on  that  than  forest  i t s ubiquitous distribution  concluded  stress  is a  green  in a  1986) Turner  forest  and  is  1975;  Chapter  2),  light  seems  to  be  an  important  factor  in  salal  growth. Studies  of  relationship Anderson was  et  with al.  positively  understory than  to  are  Direct sunfleeks Shirley  result  1945;  Sunflecks below  a  Miller  Gross  This shoot  radiation to:  1)  basal  transmission  and  1979;  little  solar  (Evans  1971) and  are  Tanner the  form  global,  relationship  radiation;  the  of  2)  status  and  to  and  et 1959).  radiation  of  regime  Slomka  1959;  diffuse  cover,  diffuse,  (measured  of  Specific  relationship  evaluate  salal  components  biomass,  direct,  active  influence  (Logan  additive  of  (e.g.,  Ustin  Czarnowski  canopies. of  the  and  1966).  transmission forest  plants to  1956;  and  examined  the  on  significantly  canopy  to  effect  Smith  form  1982),  (Young  photosynthetically PPFD)  Gross  the  1970).  photosynthesis  1979;  that  light  the  plants  productivity, of  in  moisture  through  area,  increased  but  Waring  canopies  1976).  abundance  between  influence  (Federer  explore  forest  positive  precipitation  1967),  Muller  size  throughfall  and  a  Zavitkovski  transmission  (Atzet  Chabot  shown  understory  Interactions  under  and  contribute  study  light  important  in  (e.g.,  that  to  have  (Hodges  have  forest  radiation  and  of  or  1959;  with  transmission.  may  1984),  found  responsive  radiation  growth  transmission  (1969) a l s o  more  photosynthesis  al.  light  probably  distribution  understory  correlated  was  light  moisture  forest  as if  solar  objectives of  and and  photon the  abundance  salal  were  density,  shoot  size  to  diffuse flux  density,  relationships  differed and most  3)  between  evaluate  important  stand  structures  which solar  and  radiation  for regulating  salal  biogeoclimatic component  growth.  may  subzones; be  the  86  STUDY  Twelve Vancouver  plots  Island,  the  Coastal  and  6 were  Western  Gaul t heri  (Orloci and  (Orloci  i n stands  were 800 most  of d i f f e r e n t  T h e CWHb  stands  Podzols  stand  on m o r a i n a l  (Klinka All  et a l .  plots  were  (Pseudotsuga  menziesii  heterophylla  (Raf.)  pii  cat a D o n n ) .  stand  were  western  white  species  were  parvi  folium  pine  Plots  plant  were  Vaccinium  in the plant  CWHb  veneer.  i n t h e CWHa  plots  by l e s s  smaller were  association  geographically  and separated with  1965)  i n t h e CWHa  and s t r u c t u r e s .  stand  were i n  - Douglas-fir  separated  or c o l l u v i a l  trees  than  than  Humo-Ferric  Average  and about  annual  2 3 5 0 mm  in  the  1984). i n immature (Mirb.) Sarg.),  (Pinus  stands Franco),  and western tree  f i r (Abies  primarily Smith),  were  Soils  (Chamaecyparis  yellow-cedar  hemlock  i n t h e CWHa.  Additional  amabilis  (CWHb).  was y o u n g e r ,  i s 2 1 2 3 mm  Sixplots  - Douglas-fir  ages  i n t h e same  sampled  precipitation CWHb  hemlock  on  (CWHa; K r a j i n a  1 9 6 5 ) a n d CWHb p l o t s  T h e CWHa p l o t s  a l l located m.  wet subzone  locations  Canada.  d r y subzone  - western  a - western  1965).  at several  Columbia,  Hemlock  i n t h e CWH  association  sampled  British  i n t h e Gaultheria  were  -  were  AREAS  species amabilis  nookatensis monticola  salal,  and Oregon  of  Douglas-fir  western redcedar found  (Tsuga  (Thuja  i n t h e CWHb  (Dougl.)  Forbes),  (D. Don) S p a c h ) , a n d Dougl.).  redhuckleberry grape  hemlock  (Mahoni a  Understory (Vaccinium nervosa  Pursh).  87  Plots  were  cover  and  were:  homogeneous  stand  age  or  30-60  to  abundances. forest  years,  fertilization),  elevation 4.1.  salal  selected  200-600  no  slopes m.  encompass Criteria  canopy stand  a  range  of  overstory  used  to  locate  structure,  no  deciduous  treatment  10-60%,  Descriptions  (e.g.,  no  south-southwest of  plots  are  plots trees,  thinning  aspects,  given  in  and  Table  TABLE  Plot  fl  4.1  Descriptions of plots solar radiation  Subzone  used  t o sample  s a l a l and  0  Aspect (°)  Elev. (m)  Slope (%)  Age  Cover  BA (m /ha) 2  1  CWHa  230  430  30  28  0.912  37.7  2  CWHa  215  270  15  37  0.782  46.7  5  CWHa  206  310  52  60  0.636  35.0  6  CWHa  215  310  45  59  0.783  34.4  7  CWHa  1 73  200  9  34  0.345  14.8  9  CWHa  261  630  43  42  0.899  54.9  21  CWHb  240  570  30  30  0.576  10.1  22  CWHb  240  570  30  30  0.416  8.5  23  CWHb  220  570  20  30  0.809  41 .6  24  CWHb  220  570  20  30  0.636  26. 1  26  CWHb  219  550  19  30  0.902  38.9  27  CWHb  214  600  22  30  0.757  17.5  Forest 1947)  overstory cover estimated a s a p r o p o r t i o n o f 1.0.  by moosehorn  (Robinson  89  METHODS  Detailed are 52  given 0.25-m  (0.0225  methods  f o r sampling  i n Chapters 2  quadrats  ha) p l o t s  contours.  Salal  2 a n d 3.  salal  shoots  for  basal  diameter  was v i s u a l l y  rooted  to top of previous year's  pulled  upright), Total  allometric  and current biomass  equations  current  biomass  was e s t i m a t e d  by  average  leaves  leaf  after  measurements  annual  shoot  measurements  =  52/plot)  basal  sampled  in a  when  the shoot  and biomass  are plot  averages  where  leaves  CAG  t h e number a random  Quadrat  was  were from  and d i d not foliar  o f CAG l e a v e s sample of  samples  normal  and shoot  distributions. samples  are reported backtransformed.  area,  Estimates of  are given  as p l o t  Estimates  of shoot  of measurements  and  averages  (n  h e i g h t and  of a l l shoots  plot. solar  solarimeters  loggers  ground  of transformed  2  Shortwave Dome  intervals  measured  was e s t i m a t e d  to obtain  and r e p o r t e d p e r m .  diameter  from  season.  transformed  and confidence  basal  obtained  quadrat.  from  (CAG) b i o m a s s .  by m u l t i p l y i n g  the growing were  shoot  slope  i n each  season's  in  9 x 25 m  counted,  e t a l . i n prep.)  growth  biomass  Means  density,  of each  (Vales  include  growing  was s a m p l e d  out along  were  growth  radiation  within  estimated  (distance  rooted  counted.  laid  i n the quadrat  and height  salal  located  t h e l o n g edge  cover  and s o l a r  Briefly,  systematically  with  All  salal  radiation  (Lintronic  as 3-minute  averages  (0.3-3.0  Ltd.)  um) was m o n i t o r e d  r e c o r d e d o n CR-21  of samples  taken  at  data  1-minute  with  90  intervals.  Shadow  from  radiation  global  mounted solar  m  above  radiation  amount of  1.5  of  centerline,  photosynthetic Mm)  i n the  samples.  in  subtracting  the  radiation.  Transmission  by  the  were one  daily  used full  always  2  for  over  diffuse  sum  of  of  daily open  day.  Because  two two  cases.  plot  variable  (sunrise  two  was  to  solar was  LI-190SB  (PPFD;  0.4-0.7  1-minute  obtained from was  from  days  global  that  by  full  n o o n ) was  averaged  sensors  were  for at  conditions did one  a  by  computed  monitored  during  the  below-canopy  Only  weather  plot  Transmission  equations  DENSITY;  CAGBIOM;  total  function  of  regression  the  along  calculated  component  of  pairs  diffuse  of  was  the  clear  clear least  not day,  multiplied  for plots  by  sampled  days.  Regression (density,  Each  a  monitor  component  Two  LI-COR  empirically  each sum  plots,  One  radiation  determine  canopy.  averages  radiation.  monitoring  monitoring  to  stand recorded  to  radiation  radiation  for analyses.  permit  half-day  Direct  the  diffuse  each  plot  within  used  P P F D was  on  sensors  the  edge.  3-minute  open  average  each  as  equation.  the  apart  and  separated  of  above  density  calibration  dividing  near  flux  center  Diffuse  m  from  shaded  photon  plot  pair  area  11  1969) sensors  One  potentially  7 m was  paired  open  located  and  sensor  with  i n an  were  (Horowitz  ground.  radiation  sensors  quantum  bands  basal  biomass,  transmission equation:  were area,  developed  for  SALALBA;  foliar  TOTBIOM; of  each  and  cover,  radiation  salal  abundances  productivity,  PCTCOVER) a s component  a  using  the  where  Y  i s the salal  variable,  radiation  component  asymptote  i s reached,  is a to  the level  transmitted, b  Y  type  intensity  (Thornley  regression,  x  were  )  i s 0.  i s the rate a t which of the asymptote,  This  form  used  an and c  of the equation  and g i v e s  of photosynthesis  i s often  of a  a  with  form  i s  similar  increasing  i n modeling  photosynthesis  1976).  Parameter  subroutine  that  a  relationship  the s a t u r a t i o n curve  light  i s the proportion  i s the l e v e l  o f X where  Michaelis-Menten  X  estimates  were  derivative-free (Moore  and index  computed  1984).  obtained  a  nonlinear  parameter-estimating  Standard  of determination  f o r each  using  error (/*:  regression with  FORTRAN  of t h e e s t i m a t e  Ezekiel  a n d F o x 1970)  a FORTRAN  program. •  92  RESULTS  Salal  estimates  2  density,  0.01-14.0  biomass,  0.006-64.7  for  average  basal  shoot  diameter  cm «m" g»m"  (Table  components  0.18-1.80  for  diffuse  radiation was  included  transmitted. proportions than  densities (Table  Form  of  Chapter  cm f o r  3).  average  0.026-0.57  for  direct,  The p r o p o r t i o n  for  of d i f f u s e  a n d 0.01-0.29 of d i f f u s e  maxima  r a d i a t i o n below  beam a n d  sky r a d i a t i o n  i n t h e CWHb c a u s e d  Daily  shoot  global,  scattering of direct  radiation transmitted  some  t o be  greater  and average  the canopies  flux  a r e summarized  response abundances  were  r a d i a t i o n components  subzones  (Table  transmission nonlinear, response  4.4).  upper  closely  related t o proportions of  transmitted  The response  of global, direct,  to transmission  nonlinearly with asymptote  (Figs.  within  and across  of a l l salal  and d i f f u s e  increasing asymptotically  of salal  increased high  4.3).  g«m" f o r  of the proportions of  were  0.005-0.43  2  0.06-634  4.3).  Salal the  Ranges  shoots«m" f o r  p r o d u c t i v i t y , 2 3 - 7 5 cm  a n d 0.26-0.54  structure  of diffuse  area,  f o rf o l i a r  proportion  of solar  basal  within-canopy  Stand  1.0 ( s e e  for  2  0.12-144.3  transmitted  (Table  not the true  2  from  4.2).  diffuse,  PPFD  2  height,  radiation  for  ranged  PPFD  (Figs.  4.1-4.3;  Table  were  4.1-4.3).  of diffuse  e i t h e r no o r an  abundances t o  The  radiation  unrealistically 4.4).  Diffuse  TABLE  P l o t  4.2  P l o t  DENSITY x  a  means  and  a  U-m--')  Conf.  1  0 . 7  2  10 .8  6 .9-  5  28 . 2  6  0 . 1-  int.  confidence  CAGBIOM x  i n t e r v a l s  ;  (g-m- )  Conf.  int.  a  for  TOTBIOM x  salal  (gnr' )  c h a r a c t e r i s t i c s  a  PCTCOVER  (%)  i n t .  b  HEIGHT  (cm)  Basal  c  d i a .  (cm)  Conf.  i n t .  x  0 .0-  1.0  0 .4  0 . 1-  1 .0  25 .3  18 . 9 - 3 2 . 5  44  10 . 0 -  35 .2  9 .6  6 .2 - 1 3 .6  23 .8  21 . 1- 2 6 .6  209  0 . 26 0 . 24- 0 , .28  Conf.  x  Conf.  i n t .  n  x  Conf.  i n t .  0 .02  0 .0-  0 . 1  15 .7  2 . 13  1 .0-  3 .9  20 .07  23 . 6 -  33 .2  2 8 . 13  20 . 0 - 38 .2  271 .07  189 . 4 - 3 7 3 . 3  51 . 1  43 .8 - 5 9 .0  47 . 2  43 .4 -51 . 1  354  0 . 39 0 . 37- 0 . .41  1 6 .. 5  12 . 8 -  2 0 .6  25 .56  15.. 1 - 3 9 . 9  4 0 0 . 28  242 . 9 - 614 .0  38 .5  32 .8 - 4 4 .6  74 .8  67 .5 - 8 2 .6  237  0,. 5 4  0 .49 - 0 ..59  7  6 8 .. 0  57 . 2 -  79 .8  6 4 .. 6 5  5 0 .. 4 - 81 .4  6 3 3 .. 57 466 . 9 - 835 .8  50 .3  43..5 - 5 7 .6  43.. 1  40 .9 - 4 5 .4  881  0,.41  0 ,. 39- 0 , .42  9  0.. 1  0 .4  0..01  0 ,. 2  3 4 ,.8  24 .0 - 4 7 .5  12  21  1 0 2 .. 1  8 5 .. 5 - 120 .6  22  1 4 4 .. 5  1 2 4 .. 9 - 1 6 5 . . 5  23  1 2 .. 7  8 ., 6 -  1 7 .. 7  24  50. 4  3 7 .. 9 -  6 4 ..8  19. 38  26  5 . 7  2.. 9 -  9..3  0. 58  27  40. 0  29. 2 -  5 2 ..5  13. 99  0 .0-  1 .7  95%  Average  of  52  0.25m  !  Average  of  52  0.25m  !  Average  of  a l l  shoots  0 . 23  0 .28 - 0 . 24  0 .3  0,, 1  2 7 8 .. 0 7  2 1 1 .. 9 - 356..7  52,,9  45., 7- 6 0 ., 5  2 7 ,. 2  26 . 1- 2 8 ,.3  1294  0..30  0.,29 - 0 . 3 0  2 8 9 . 17  2 1 6 .. 7 - 3 7 6 .. 1  4 8 ..7  4 0 ..4 - 5 7 . .7  22. 6  2 1 .. 8 - 2 3 . , 4  1750  0 .,31  0 .,31 - 0 . 3 2  5..5  9 2 . 61  4 6 . . 3 - 1 6 2 ., 4  12..2  7..7 - 1 7 . ,8  4 3 ..3  37..9 - 4 9 . . 1  209  0. 37  0.,35 - 0 . 40  1 3 . 6 - 2 6 .. 5  1 6 5 . 15  1 0 5 . , 6 - 2 4 3 ,, 6  33 . 5  25. 8 -42. 3  32 . 3  30..5 - 3 4 ., 1  680  0. 30  0 . 2 9 - 0 . 31  4. 5  24. 3  2 0 ..8 - 2 8 . 0  126  0. 27  0. 25 - 0 . 30  20. 6 -33. 4  43. 9  4 1 .. 6 - 4 6 . , 2  608  0. 34  0. 33 - 0 . 35  0 .0  0..06  3 6 .. 4 8  30.. 9 - 42 .7  55..59  4 5 . 8 - 66..6  0.. 0 -  3.. 33  quadrats  0 .21  1.8-  0. 2 -  x  1. 2  7 . 22  8. 9 - 20. 8  0 .0-  2 . 6-  15..3  2 .6  226. 96  139. 3 - 3 4 5 ..2  26. 6  sample  s i z e s .  4.  quadrats. measured.  See  n  for  0., 0 -  1. 2 -  0 ,. 34 0,. 26- 0 . 44  c  TABLE  4.3  Below-canopy  Plot  solar  GLOBAL  radiation  statistics  DIRECT  DIFFUSE  PPFD  3  b Max (Wm  c  1 2 5  d  6  Max (W-tn-  -«)  X  P.T.  Max (Wm-  n  X  P.T.  ')  Max X („E m -' s-  )  15  0 .026  75  4  0 .008  26  10  0 . 177  5  1 .6  0 .010  810  35  0 .074  779  23  0 .053  33  12  0 .360  9  4.5  0 .050  720  31  0 .073  676  16  0 .046  64  16  0 .260  35  11.9  0 .069  750  41  0 .091  710  22  0 .058  43  19  0 .286  40  15.9  0 .088  e  940  200  0 .465  860  165  0 .429  96  35  0,. 775  9  C  600  20  0 .040  579  6  0 .013  34  14  0. 310  9  21  890  129  0 .277  690  54  0 . 127  215  75  1 ,429 .  22  990  250  0 . 566  846  164  0.,416  165  86  120  27  0..058  78  96  0..006  65  24  350  62  0.. 135  272  15  0.,035  26  100  17  0.,035  84  2  27  560  65  0. 142  438  17  photosynthetic  photon  C  P.T. 1  90  Diffuse  ^Proportion c  P.T.  7  23  a  X  Average  of above-canopy  of monitoring  flux  43 . 1 0 .286 1 .4  0,.01 1  98  38 . 3  0 . 261  1 .799  66  28.7  0,.216  24  0. 569  9  4.3  0..036  140  47  1 .153  26  12.7  0.,112  0. 005  30  15  0. 338  4  2.0  0. 016  0. 039  120  48  1. 293  21  8.3  0. 080  density.  radiation  f o r two  104  transmitted.  days.  Computed  from m o n i t o r i n g  one  full  day  Computed  from m o n i t o r i n g  one  half-day.  and  one  half-day.  TABLE 4 . *  Regression c o e f f i c i e n t s of equations p r e d i c t i n g s a l a l v a r i a b l e s from s o l a r r a d i a t i o n components. Sample s i z e s : C W H a « 6 , CWHb-6. Equations s i g n i f i c a n t at  Y  DENSITY:  SALALBA:  TOTBIOM:  p < 0.06.  Subzone  R e g r e s s i o n e q u a t i o n : Y = -j—+ " a | x - c )/b  X  a  c  b  /  2  s  X  y  108. 1 0 . 1 0 0 .6 0. -3. 4 - 1 273. 7 0.  0290 0083 .7280 0081  0. 0. 0. 0.  93 93 83 95  7. 5 7.4 7 11 . 6. 1  559.4 1686.3 35.5 776.6  288. 183. -87. 369.  0315 0002 2575 0164  0. 0. 0. 0.  98 99 93 87  7. 5. 16. 22.  5 8 4 2  GLOBAL DIRECT DIFFUSE PPFD  520.9 1054.4 72.9 682.0  211. 146. 221167. 246.  0 . 0317 0. 0033 0 . 1811 0. 0163  0. 0. 0. 0.  86 7 1 80 76  17. 25. 21 . 23.  9 3 2 2  CWHa  GLOBAL DIRECT DIFFUSE PPFD  122.8 140.7 0.2 103.3  19. 3 0. 0324 18. 4 0. 0104 - 0 . 6 - 2 . ,5214 28. .6 0.,0139  0. 0. 0. 0.  88 85 69 94  2. 2 2. 4 3. 5 1 .5  CWHb  GLOBAL DIRECT DIFFUSE PPFD  53.4 3.1 80.7  24. ,2 0.,0255 16. ,2 - 0 . ,0027 -8. , 1 0..1737 24. ,8 o..0127  0. ,99 0.,99 0.,96 0.,85  0. ,3 0.,5 1 ,. 1 2., 1  71.0 134.4 5.7 87.2  21 ..3 0..0257 17. .9 0..0020 8804. ,0 - 0 . .1711 27. .2 0..0127  0.,89 0.,89 0..40 0..88  1 1 4 1  CWHa  GLOBAL DIRECT DIFFUSE PPFD  418.8 495.5 1 .3 324.0  CWHb  GLOBAL DIRECT DIFFUSE PPFD  Both  159.5  Both  GLOBAL DIRECT DIFFUSE PPFD  CWHa  GLOBAL DIRECT  6631.0 7730.0  DIFFUSE PPFD  N.S." 5665.0  9 0. 5 -0. 1 0. 9 0. 1 0 7 6  ,7 . .7 . ..0 . .8  829, .5 790, .7  0,.0325 0,.0106  0,.82 0,.79  125. .7 1 35. .2  1119,.3  0,.0141  0,.90  91 .9 .  0,.0330 0,.0016 0,.1992 0,.0151  0,.97 0,.92 0,.94 0..92  22. ,7 34. .5 29. .2 35. .3  497, .8 531 .7 ,  0,.0307 0,.0027  0,.60 0,.63  128. .4 122. .4  . 655,.3  0,.0123  0,,6B  114. .8  337, .3 309, . 1 -547155. 343, .5  CWHb  GLOBAL 4758.0 DIRECT 16027.0 DIFFUSE 196.6 PPFD 6241.0  Both  GLOBAL DIRECT DIFFUSE PPFD  5277.0 7405.0 N.S. 4679.0  CWHa  GLOBAL DIRECT DIFFUSE PPFD  476.8 537.2 0.6 406. 1  0,.0320 94, .6 90, .6 0,.0098 - 2 , .3 - 2 , .6543 1 57, .8 0,. 0 1 3 9  0,.88 0,.86 0..73 0,.94  9. .9 10. .6 14. .6 7., 1  CWHb  GLOBAL DIRECT DIFFUSE PPFD  204.6 603.2 14.4 317.6  1 16,.5 71 .3 , - 3 3 , .2 120, .7  0,.0374 0,.0016 0,.3522 0,. 0 2 1 9  0..99 0..99 0..94 0..84  2.,6 2..6 5.,9 9. .3  Both  GLOBAL DIRECT DIFFUSE PPFD  245.3 523.7 23.3 317.5  .1 13, 0,.0254 82. .9 0,.0044 899135, .0 - 0 , . 1637 162, .3 0,.0148  0..86 0,.90 0..33 0..86  8..7 7..4 19. ,0 8. .7  PCTCOVER: CWHa  GLOBAL DIRECT DIFFUSE PPFD  1245.0 1879.0 N.S. 1202.0  58, .6 53, .9  0,.0294 0,.0098  0..64 0..65  16. .3 16. .0  63, .6  0,.0111  0..79  12. ,5  CWHb  620. 1 GLOBAL DIRECT 2096.0 DIFFUSE 44.5 PPFD 557.8  62. .9 55.8 210, .6 84, .9  0,.0318 0.0013 0,.2818 0..0113  0..93 0,.93 0..87 0..99  5..7 5.,9 B..0 0.,2  Both  GLOBAL DIRECT DIFFUSE PPFD  984.5 1651.0 N.S. 905.3  56, .9 56, .6  0,.0303 0,.0040  0..73 0..72  1 1 ,6 . 1 1,8.  66 .7  0,.0121  0,.85  8.,7  CAGBIOM:  °Equation  not s i g n i f i c a n t  1  (p i 0 . 0 6 ) .  o.e 01 05 oa 0.4 o.a o.a PROPORTION GLOBAL. RADIATION TRANSMITTED  (To  0.1  OJ  OS  04  ^>.B  PROPORTION DIRECT RADIATION T R A N S M I T T E D  2  2  F i g u r e 4.1. R e l a t i o n s h i p s o f s a l a l b a s a l a r e a (cm 'm~ ) a n d 95% c o n f i d e n c e intervals t o transmission of global, d i r e c t , d i f f u s e , a n d d i f f u s e PPFD s o l a r radiation components. Dotted l i n e s a r e r e l a t i o n s h i p s in individual s u b z o n e s . CWHa ( • ) , CWHb (ffl).  97  oo o.a o.« o.e o.a to o 1 « to i t PROPORTION DIFFUSE RADIATION TRANSMITTED  oo oi O J o.a PROPORTION DIFFUSE PPFD TRANSMITTED  F i g u r e 4.2. R e l a t i o n s h i p s o f s a l a l f o l i a r productivity ( g » m ~ ; CAGBIOM) a n d 9 5 % c o n f i d e n c e i n t e r v a l s t o t r a n s m i s s i o n o f g l o b a l , d i r e c t , d i f f u s e , a n d d i f f u s e PPFD s o l a r r a d i a t i o n components. D o t t e d l i n e s a r e r e l a t i o n s h i p s i n i n d i v i d u a l s u b z o n e s . CWHa ( • ) , CWHb (ffi). 2  lOOO.O-i  _  000.0-  1000.0-i 000.0-  •oo.o-  PROPORTION GLOBAL RADIATION TRANSMITTED  PROPORTION DIRECT RADIATION TRANSMITTED  2  i g u r e 4.3. R e l a t i o n s h i p s o f s a l a l t o t a l b i o m a s s ( g « m ~ ; TOTBIOM) a n d 9 5 % c o n f i d e n c e i n t e r v a l s t o t r a n s m i s s i o n o f g l o b a l , d i r e c t , d i f f u s e , a n d d i f f u s e PPFD s o l a r radiation components. Dotted l i n e s a r e r e l a t i o n s h i p s in individual s u b z o n e s . CWHa ( • ) , CWHb (ffl).  99  irradiances daily  canopies  were  plots  about  equal  was a b o u t  Although  salal  an  with  asymptote  transmitted radiation  radiation,  Salal  (Fig.  however,  increased  to transmission  due t o s i t e  PPFD  Below-canopy  diffuse  of absorption  i n below-canopy  range  of shoot  shoot  and diameter  increased,  relationships  were  Salal  transmission  size  were  and direct  height more  shoot  of solar  stand  sizes.  small. were  by  diffuse  i n t h e CWHa, A  slight  o f t h e same a g e , At the lowest  l a r g e r , but as  pronounced  diameter f o rheight  was a d v e r s e l y  radiation.  Shoots  and basal  radiation  As t r a n s m i s s i o n  and basal  size  (height  variability  i n t h e CWHb  shoots  height  shoot  and age d i f f e r e n c e s .  was f o u n d  a narrow  diameter.  approached  increasing proportion of  of global  considerable  transmission  further  of d i f f u s e  with  size  relationship despite  with  increase  (Chapter 3 ) .  4.4) s h o w e d  probably  plot  of diffuse  apparently  3) b e c a u s e  o f PPFD  r e l a t i o n s h i p of s a l a l  diameter)  1 and d i f f u s e  unlimited  4.1-4.3).  (Chapter  T h e amount  s hoot The  4.4; F i g s .  average  of global.  an a l m o s t  canopy.  transmitted  plot  increasing proportion  (Table  When  r a d i a t i o n below t h e  radiation, salal  was l o w i n P P F D  forest  global  4.3).  4.3) t h e p r o p o r t i o n  showed  of diffuse  PPFD  (e.g.,  10x t h a t  transmission  low ( T a b l e  and d i f f u s e  23 o r 26 i n T a b l e  transmitted  the  were  irradiance of global  canopy 5,  under  increased,  transmission  decreased. than  affected were  The  f o r basal by  high  productive,  100  100.0-  100.0-  80.0-  00.0-  eo.o-  eo.o-  < 40.0-  40.0-E  zoo- .  20.0--  0.0-  0.0 0.1 O.J 0.3 0.4 o.e o.e PROPORTION G L O B A L RADIATION TRANSMITTED  0.0 0.1 0.2 0.3 0.4 O.S PROPORTION DIRECT RADIATION T R A N S M I T T E D  E  i  0.6  0.6  0.4  I 0.3-  0.2-  i  i  m  0.1  0.0 0.1 0.2 0.3 0.4 0.6 PROPORTION G L O B A L RADIATION TRANSMITTED  0.3  i B m  Q  $ o.i-  0.0-  0.0 0.1 0.2 0.8 0.4 0.6 PROPORTION DIRECT RADIATION TRANSMITTED  Figure 4.4. Relationships o f a v e r a g e s h o o t h e i g h t (cm) a n d b a s a l d i a m e t e r (cm) o f s a l a l w i t h 9 5 % c o n f i d e n c e intervals t o t r a n s m i s s i o n o f g l o b a l a n d d i r e c t r a d i a t i o n . CWHa ( • ) , CWHb (ffl).  101  however,  under  open  canopies  with  high  transmittance  (Fig. 4.5).  Subzone  differences  A qualitative differences were  assessment  of equations  available  differed  predicted  by d i f f u s e  4.4).  different Chapter  3 F i g . 3.2).  t h e two s t a n d  The  manner  maximum  abundance  transmitted rate  upper  would  a t which  asymptote  interaction (stand  differences subzone  radiation  regression of s a l a l likely  that  due t o  would  and the  have  t h e subzones  of d i f f u s e  of stand  i s reached  and s i t e  that  Table  was t r a n s m i t t e d ( s e e  were  radiation  b)  subzones might  by d i f f e r e n t  age and s i t e  coefficients.  o f t h e way i n w h i c h  effects  were  i s transmitted  be r e g u l a t e d  (coefficient  between  s a l a l was  4.1-4.3;  between  f o r a given  a maximum  structures)  tests  used.  the subzones  effects  and the e f f e c t s  to affect  where  radiation  in salal  no  structures.  i n which  structures  interact  The  between  Any s i t e  to differences  because equation  the equations  diffuse  potential  Figs.  by t h e d i f f e r e n t i a l t r a n s m i s s i o n  between  stand  (e.g.,  structure  way i n w h i c h  contributed masked  i n stand  subzones  i n a l l cases  radiation  between  of  of regression  strongly  Differences  differences  between  f o rt h e form  Equations  was made  Among  proportion by s i t e  stands, of  radiation  radiation  and age e f f e c t s .  (coefficient  are regulated  quality  a) a n d  by an  i s transmitted  and age e f f e c t s .  Although  are discussed,  the potential  cause  equations  t o be d i f f e r e n t a r e  102  O O JZ  1.75-1  w  3  1.60 H  >  1.25-  0 D  1.00-  Q  O tr Q.  0.75-  fe  O  X CO  UJ o  < cc  5 1X1  o o  0.50-  -ffl 0.25-j ^ • ' ^ 0.00 0.0  0.1  0.2  0.4  0.3  0.5  0.6  PROPORTION GLOBAL RADIATION TRANSMr TED  1.75-1  JZ  w  3  1.50 H  >  125-  3  1.00-  F u Q  O  CC CL  0.75-  fe  O  X  to UJ  <  CC UJ  5  0.50-83-  025 0.00  I  0.0  0.1  I  02  I  0.3  I  0.4  1  0.5  PROPORTION DIRECT RADIATION TRANSMITTED  Figure 4.5. R e l a t i o n s h i p s of average p r o d u c t i v i t y of s a l a l shoots t o transmission of g l o b a l and d i r e c t s o l a r r a d i a t i o n . CWHa ( • ) , CWHb (ffl).  103  confounded  by b r o a d  transmission  unit  between  predicting  area  transmission  from  4.4).  differed  Maximum  t h e CWHb  Slopes  4.4).  Site  density  size,  a small  shoots  plctt  7 density  4.2),  average  that  of shoots  global,  pronounced. SALALBA  Equations  between  than  the subzones  the  (Table (Table  Because t o shoot  a  biomass.  that  than  in  density  related  Although  of plot  22's (Table  7 was m o r e t h a n  having  among  from  quadruple  (Chapter  area,  less  the largest  coefficient  magnitude  of difference  CAGBIOM,  a n d PCTCOVER  radiation  3).  bf o r of  components.  transmitted  of d i r e c t  basal  transmission of were  the radiation  radiation  area  cover  salal  PPFD r a d i a t i o n  SALALBA,  and basal  greater  strongly  give  predicting  and greatest  Transmission  productivity  one-half i n plot  4.3; T a b l e  i n a lower  would  PPFD  22.  and d i f f u s e  of direct  was m u c h  i n shoot  and percent  predicting  proportion  size  biomass p e r  i n t h e CWHa  4 . 4 ) i n t h e CWHa.  i n equations  a differed  lower  differed  resulted  change  The subzone  coefficient  differ.  i n plot  a n d CAGBIOM  and  and d i f f u s e  Fig.  i sallometrically  biomass  productivity, direct,  also  i n shoot  was l e s s shoot  direct  (e.g.,  biomass  a)  (Fig.  greater  Differences foliar  subzones  total  biomass  change  proportionately  structure  and t o t a l  of global,  and age e f f e c t s  shoot  density  was c o n s i d e r a b l y  (coefficient  4.4) o f l a r g e r individual  salal  between  a n d maximum  CWHa.  i n stand  t h e two s u b z o n e s .  Equations  radiation  in  differences  generally  from t h e d i d not  d i dnot d i f f e r  Although  i n t h e CWHa p l o t  individual  shoot  7 was a b o u t  twice  1 04  that and in  o f CWHb p l o t therefore  SALALBA  t h e two s u b z o n e s  transmission and  salal  differ  very  Figs.  The  form  diffuse  radiation did values  probably  because  Relative  predictability  Although  proportions  abundance diffuse  of site  high  radiation  Results a l l salal  transmitted  shoot  size  subzones.  transmitted  differed  components r a d i a t i o n would  estimates  beyond  what  radiation did 2  was  ( F i g . 4.5) t o t r a n s m i s s i o n  of diffuse  regression  (i  components  structures.  proportion  between  adequately  poorer  better  predict  abundance here,  predict  Relationships  t h e subzones  abundances  of salal  was f o u n d  = 0.87-0.96).  r a d i a t i o n i n t h e CWHa w e r e  predicting  due, i n p a r t , t o  and age e f f e c t s .  transmitted  i n t h e CWHb  components.  likely  among radiation  of diffuse  from t h e  r a d i a t i o n appeared t o  s u b s t a n t i a l l y between  a given  transmission  unrealistically  transmission  for  structures  a n d PCTCOVER  r a d i a t i o n which  productivity  not d i f f e r  Because  d i dn o t d i f f e r .  of the r e l a t i o n s h i p s of s a l a l  4.4) a n d s h o o t  similar  A d d i t i o n a l l y , global and  by t h e two s t a n d  Absolute  at  PPFD  were  the stand  equations  CAGBIOM,  3).  area  one-half  4.1 a n d 4 . 2 ) .  i n t h e way t h e s e  (Chapter  PPFD c o n t a i n  differently  (Fig.  similar,  4.1 a n d 4 . 2 ) ,  differences  transmitted  diffuse  4.2; F i g s .  and d i f f u s e  was a b o u t  per unit  r a d i a t i o n between  were  of global  (e.g.,  potential were  (Table  p r e d i c t i n g SALALBA,  proportion  density  a n d CAGBIOM  of direct  abundance  Equations  of  22 ( F i g . 4 . 5 ) ,  than  using  differed,  than  other  salal  using  other with  PPFD  components  105  in  t h e CWHa, b u t P P F D g e n e r a l l y  than  other  components  more  closely  CWHa  than  were  better  related  well  biomass,  subzones.  Effects  radiation  Salal faster  radiation direct  generally 4.4).  a,  (coefficient  less  lower b,  transmitted  than  results  suggest  components.  (Chapter  3  i n t h e CWHb  equally  SALALBA,  well  about  by  Density, equally  radiation  reached radiation 4.4; F i g s .  upper  in  4.4),  asymptotes  than  f o r other  4.1-4.3).  f o rtransmission  Salal  of direct  and the proportion of  o r PPFD  that  better  salal  at levels  f o rg l o b a l  by t r a n s m i s s i o n  by o t h e r  Table  were  predicted  on  Table  though  equations.  and d i r e c t  of d i r e c t  consistently  These  regulated than  (coefficient  radiation  were  consistently  abundances  predictabilities. about  the  and d i r e c t  variation  combined  components  f o rtransmission  were  for  also  even  i n t h e CWHa  predicted  of global  abundances  components maxima  were  radiation  individual  of global  of s i t e  in  Salal  radiation  t o the better  by t r a n s m i s s i o n  of  and d i r e c t  and cover  3.3).  i n t h e CWHa  The lack  a n d PCTCOVER  and d i r e c t  3 Table  characteristics  contributed  CAGBIOM,  total  by s t a n d  3.3 a n d 3 . 4 ) .  have  global  of global  o f P P F D was  characteristics  by t r a n s m i s s i o n than  predictability  Transmission  stand  (Chapter  i n t h e CWHb  predicted  may  to forest  predicted  transmission  Tables  i n t h e CWHb.  i n t h e CWHb  radiation  had a lower  salal  of the direct  of s a l a l  near  (coefficient i s more radiation  zero  was  c, T a b l e  closely component  106  DISCUSSION  Salal shoot with  and  density, per  unit  increasing  photosynthetic 4.5).  The  the  area,  photon of  the  Because  relationships  diffuse  radiation  curve.  The  of  difference  photon  growth  diffuse  He  relations  concluded  stress  for  f . c ) , under on  that  used  Swank the  sunny  in  time in  (1972)  mid-light days  in  abundances the  a  intensity. is  lowered  net  compared  to  "shade"  plants.  shoots  Light  photosynthetic  that  The  of  of  the  below and  a  by  three  (400,  a  in  high  this  light  be  2600  greatest  under  2000  for  water  1600,  to  on  (1972).  greater  greater rate  plant  canopy.  Swank  under  than  of  influence  intensity  results  received  diffuse  its  Of  low,  saturation  understory  photosynthesis  and  in  transmission  experiment  intensity  resulted  that  light  light  plants.  less  of  to  was  were  usually  combination  indicated  shade  salal  response  demonstrated  plants  greenhouse  the  relating  radiation  was  to  importance  energy  4.1-4.3,  similar  PPFD and  the  diffuse  canopies  phase  to  solar  "sun"  found  to  and  (Figs.  below  diffuse  solar  than  summer  high  lower  opposed  salal  salal  longer  intensities  irradiance  demonstrate  as  of  radiation were  per  asymptotically  direct,  photosynthetic  shortwave  interaction  moisture  of  between  measuring  flux  global,  relationships  in  relationships  The  of  productivity  increased  density  salal  were  biomass,  cover  diffuse  shortwave  to  and  flux  relationship  intensity.  area,  proportions  forms  asymptotic  basal  canopies  f.c.  The  water  stress  "sun"  plants  study  proportion  of  107  direct  radiation  photosynthetic  were  rate  received  less  however,  increased  intensity, lowest. lowest less  Swank  productive because  growing light  in  by  ensuring  time  as  to  compensation (1961)  and  shoots  have  canopies. in  in  also  discussed  by  the  more  and  open  with  of  areas  for the  leaf  of  light.  adequate ideas  open  available  is  on  found  than  salal  that  under  population  age  Ellis,  B.C.).  "plants"  in  canopies,  unpubl.  Bunnell  open even  areas  data,  under B.C.  unit for  salal  forest  reproduction  salal  than  and  Sabhasri  determining  (R.M.  increasing  i s paid  Salal  areas  by  recruitment  structure  was  old-growth Ministry  of  and  Koch-Bakker  (1986)  had  more  shoots  than  canopy  plants  were  though  low  f i x e d per  canopies.  clearcut  for  tissue,  under  young  growth.  plants  structures.  areas  and  of  by  expense  (1986)  the  radiation,  material,  An  was  compensate  carbon  light  smaller  photosynthetic  Koch-Bakker  not  more  Plants  reproductive  in  were  and  open,  under  more  flowers are  area  to  low  received  "compensation"  same a m o u n t  reduced  Bunnell  by  the  At  that  light  environments.  that  photosynthesis  received  consistent  lowered  productivity,  transmitted  little  of  shoots  ( F i g . 4.5).  shoots  that  too  receiving  Victoria,  salal  "plants"  was  than  Shoot  that  that  shoots  He  that  canopies  Forests,  found  increase  Data  measured  found  than  light  plants  greater  reported  allocating assimilates  thus  rhizomes  ( F i g . 4.4).  radiation  there  low  size  forest  of  (1973).  leaf  as  here  because  stress)  asymptotically  r e s u l t s are  Whitehead  (possibly  moisture  (1972)  Results  My  or  radiation  proportion  likely  smaller  older.  108  Boardman between  sun  generally  (1977)  and  have  lower  dark  lower  stomatal  thicker  fixation. were  shade  plant  C0  with  and  Sabhasri a  higher  (1973)  transmitted that  had  between  leaves  in  plants  reviewed  by  a  in  basal  Chapter  with  area  sun than  leaf  (reduced  size  masses  proportions  plot  and  pronounced  form  of  height  and  darker  a  green  study,  of  than  open  areas.  i t is likely  that  salal  consistent with  the  my  radiation  transmitted.  in  physiology  to  Salal Although  sun  and  species  (1977).  proportions' transmitted infra-red  Sabhasri  with  relationship similar  proportions  to  proportion  r e l a t i o n s h i p between  this  red  sun  salal  salal  2  The  moderate  under  C0  observations.  to  greatest  from  surface  area  additional explanation  elongation.  higher  greater  that  with  consistent  Boardman  near  have  leaves  found  possible  of  plants  consistent  A  effects  Sun  3),  are  have  (1985)  by  shoot  shade  examined  shade  leaf  salal  unit  levels,  i n c h l o r o p h y l l , have  reported  per  plants  compensation  stomata  plants,  (1961)  stand  shade  consequently  that  mass  and  leaves.  of  and  Kelliher  increasing  average  thinner  shade  that  richer  density  radiation transmitted,  salal  are  found  than  stated  saturation  rates,  (1972)  had  He  diffusion  2  leaves.  Whitehead's  low  to  (1977).  increase  light  higher  thicker  leaves  not  and  physiological differences  plants.  density,  Swank  plant  of  lower  leaves  Boardman  shade  respiration  conductances  plants  reviewed  wavelengths  (1961)  light.  found  The  for  could  (NIR)  that  taller  on  salal  be  shoots due  to  under the  stem shoot  s p e c t r a l composition  length below  was a  109  coniferous 1966)  forest  a n d may  mentioned shoots  stimulate  that  a t low l i g h t  enrichment  different  smaller  4.4), f o l i a r  (Fig.  4.5).  either  losing  Leaves  may  (Swank  1972).  (Sabhasri  reproduction Ellis,  Sabhasri highest  with  size  by  early  light  intensity in salal  increased  proportions  greater  density  smaller  shoots  biomass.  Shoot  i s most  radiation  salal  density  small  transmitted. much  twigs  biomass,  dying  moisture  back.  stress  of flowers  i n high was  and  1986) o r  shoot  1986; density.  greatest  at the  (400 f . c ) . per unit  open  an o v e r a l l i n open  a  i s productive,  transmitted. under  transmitted  and Koch-Bakker  length  abundance  of shoots to give  root  were  rapid with  of high  resulting  he u s e d  Increases  size  increased  and Koch-Bakker  (Bunnell  that  o r NIR  i n shoot  r a p i d l y or having  1961; B u n n e l l  found  in older  rate,  do n o t a c c u m u l a t e  because  data)  also  whether  of r a d i a t i o n  of d i r e c t  open-grown  rhizomes  (1973)  per shoot  be a l l o c a t e d t o p r o d u c t i o n  unpub.l.  (1961)  and biomass  of i n c r e a s e  leaves  Tanner  transmitted.  proportions  Because  may  I t i s unclear  the differences  i n t h e open  their  and  elongation  p r o d u c t i v i t y per shoot  d i e back  assimilates  R.M.  shoot  The r a t e  shoots  stem  Evans  stress, photosynthetic  i n the proportion  Apparently  fruit  caused  proportions  (Fig.  (Federer  elongation.  can explain  salal  i n NIR  intensities.  at the highest  increase  stem  moisture  alone  Although  i s rich  etiolation  "compensation",  among  canopy  Within  canopies  increase  areas  area  may  were a  found  subzone the  compensated f o r  i n basal be h i g h  area  and  because of  1 10  increased  reproduction  (Whitehead  Bunnell  and Koch-Bakker  reduced  competition  found  under  competition  f o rlight  causes  thinning,  or because  (Chapter with  was  subzones  by d i f f e r e n t  differed  variables.  Within  an i n t e r a c t i o n  density CWHb,  stand  density, Keyes  Running  than  1977; G h o l z  coast  Among  or  1982).  strongly  with  biomass  (Table  are affected  4.4) a n d s t a n d  a s between biomass  as dependent  t h e CWHa a n d  and density  may  of radiation transmitted or of age d i f f e r e n c e s , or site  Wetter  Island  on t h e d r i e r  3) a n d  and biomass  stands,  1981),  (Chapter  variable, anda l l  radiation  because  of Vancouver  a r e found  Island.  differed  density  proportion  and G r i e r  1980)  developed i n  structures  and t o t a l  t h e CWHa, maximum  either  et a l .  equations  stand  density  of solar 2).  stand  inter-specific  Equations  a stand,  f o r a given  (e.g.,  north  (Chapter  and within  differ  with  higher  1986).  r a d i a t i o n a s an independent  equations  proportion of  b e a t t r i b u t e d t o t h e way r a d i a t i o n  and age d i f f e r e n c e s .  diffuse  by  could  shoots a r e  intra-specific  lower  (Black  (Weetman e t a l .  d i f f e r e n c e s between  transmitted  site  f o r moisture  data), or  Fewer  because  with  3 ) , and p o t e n t i a l  nutrients  Apparent  i sassociated  trees  (unpubl.  shoots.  possibly  densities  individual  among  canopies,  transmitted  possibly  and E l l i s  denser  radiation  competition  (1986)  1 9 7 3 ) a s was f o u n d b y  water  salal  support  interior  balance  sites much  nutrition ( G r i e r and  on t h e west a n d  larger  and eastern  shoots  parts  oft h e  111  Subzone area,  differences  productivity,  global  or  diffuse  or  difference  predicting  salal  All  salal  components the  most  was  (Table  in  photon  contribute  on  (1982)  level  is  to  stated a  plant's  sunflecks  become.  equations  predicting  (1982)  density,  of  moisture  stage)  and  steep  salal  direct  status,  water  My  shoot  3),  total  radiation, and  balance.  effects  are and of  the  suggest  would  site  age  had  radiation likely  a  canopy.  low  light  more  important of  the  support  that  salal  regulated  probably  radiation  of  diffuse  radiation  likely  no  cover.  rate  ( c o e f f i c i e n t a)  direct  of  radiation  average  point,  or  most  PPFD b e l o w  the  results  by  and  sunfleeks  slopes  size  showed  direct  lowest  closer  from  well  of  basal  relationships  below-canopy  compensation  The  and  the  the  statement.  biomass  interaction and  (Chapter  that  for  salal  transmission  radiation  predicted  the  to  from  for  productivity,  (maxima  Because  evident  subzones foliar  were  salal  flux  direct  Transmission  significantly  Gross  Gross'  area,  4.4).  fastest).  low  Only  variables  slightly predicted  between  basal  influence  increase  cover  PPFD.  pronounced  were  with  by  plant  an age  (successional  1 12  CONCLUSION  Salal size  were  through  density, well  related  forest  abundance  and  were  a  function  radiation biomass, direct  active  and  of  site  and cover  shortwave  of g l o b a l , radiation  equations stand  of  Relationships  transmission  between  biomass,  to transmission  canopies.  photosynthetically Differences  productivity,  were  Salal  were  responsive  solar  radiation.  between  and  shoot  radiation salal  and  diffuse  asymptotic. in different  effects  effects. most  solar  direct,  developed  structure  cover,  density,  on  subzones  transmission  productivity,  to transmission  of  of  113  REFERENCES  A n d e r s o n , R.C., O . 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C o u r t i n , a n d F . C Nuszdorfer. 1984. S i t e d i a g n o s i s , t r e e s p e c i e s s e l e c t i o n a n d slashburning g u i d e l i n e s f o r the Vancouver Forest Region, B r i t i s h C o l u m b i a . L a n d M a n a g e . R e p t . No. 2 5 . B.C. Ministry of F o r e s t s . V i c t o r i a , B.C. K r a j i n a , V . J . 1965. B i o g e o c l i m a t i c z o n e s B r i t i s h Columbia. E c o l o g y of Western 1:1-17.  and b i o g e o c o e n o s e s of North America.  L o g a n , K.T. 1 9 5 9 . Some e f f e c t s o f l i g h t o n g r o w t h o f w h i t e p i n e s e e d l i n g s . Can. Dept. N o r t h e r n A f f a i r s and N a t i o n a l R e s o u r . , F o r . B r . , F o r . R e s . D i v . T e c h . N o t e No. 82. L o n g , J.N. a n d J . T u r n e r . 1975. A b o v e g r o u n d b i o m a s s o f u n d e r s t o r e y a n d o v e r s t o r e y i n an age s e q u e n c e o f f o u r Douglas-fir stands. J . Appl. Ecol. 12:179-188. M i l l e r , D.H. 1959. T r a n s m i s s i o n o f i n s o l a t i o n t h r o u g h f o r e s t c a n o p y , a s i t a f f e c t s t h e m e l t i n g o f snow. Schweiz. A n s t . f o r s t l . Versuchswesen. 35:57-79. M o o r e , C. 1 9 8 4 . UBC C U R V E . C o m p u t i n g C e n t r e . B r i t i s h Columbia, Vancouver, B.C.  pine Mitt.  University  of  M u l l e r , R.A. 1971. T r a n s m i s s i o n c o m p o n e n t s of s o l a r radiation i n p i n e s t a n d s i n r e l a t i o n t o c l i m a t i c and s t a n d v a r i a b l e s . U.S.D.A. F o r . S e r v . , P a c . S o u t h w e s t F o r . a n d R a n g e E x p . S t a . R e s . P a p e r PSW-71. O r l o c i , L . 1965. The C o a s t a l W e s t e r n H e m l o c k Zone on t h e south-western B r i t i s h Columbia mainland. E c o l o g y of W e s t e r n N o r t h A m e r i c a . 1:18-34.  11 5  R o b i n s o n , M.W. 1 9 4 7 . An i n s t r u m e n t t o m e a s u r e c o v e r . F o r . Chron. 23:222-225.  forest  crown  S a b h a s r i , S. 1 9 6 1 . An e c o l o g i c a l s t u d y o f s a l a l (Gaultheria shall on P u r s h ) . P h . D . t h e s i s . U n i v . o f W a s h i n g t o n , S e a t t l e , WA. S h i r l e y , H.L. 1945. Light measurement. I I . Bot.  a s an e c o l o g i c a l f a c t o r Rev. 11:497-532.  and i t s  S t a n e k , W. , D. B e d d o w s , a n d D. S t a t e . 1 9 7 9 . F e r t i l i z a t i o n and t h i n n i n g e f f e c t s on a D o u g l a s - f i r e c o s y s t e m a t Shawnigan Lake on V a n c o u v e r I s l a n d . Can. F o r . S e r v . R e p t . No. BC-R-1. V i c t o r i a , B.C. S w a n k , W.T. 1 9 7 2 . Water b a l a n c e , interception, and t r a n s p i r a t i o n s t u d i e s on a w a t e r s h e d i n t h e Puget l o w l a n d r e g i o n o f w e s t e r n W a s h i n g t o n . Ph.D. t h e s i s . U n i v . o f W a s h i n g t o n , S e a t t l e , WA. Tan,  C.S., T.A. B l a c k , a n d J.U. Nnyamah. 1977. C h a r a c t e r i s t i c s of s t o m a t a l d i f f u s i o n r e s i s t a n c e i n a D o u g l a s - f i r f o r e s t e x p o s e d t o s o i l w a t e r d e f i c i t s . Can. J . F o r . R e s . 7:595-604.  Thornley, J.H.M. 1 9 7 6 . Mathematical models i n p l a n t p h y s i o l o g y . A c a d e m i c P r e s s . New Y o r k , NY. U s t i n , S.L., R.A. W o o d w a r d , M.G. B a r b o u r , a n d J . L . H a t f i e l d . 1984. R e l a t i o n s h i p s between s u n f l e c k dynamics a n d red f i r seedling d i s t r i b u t i o n . Ecology. 65:1420-1428. V a l e s , D.J. a n d F.L. B u n n e l l . 1986. under f o r e s t canopy I I . P a t t e r n J . E c o l . (in pr e p. ) .  S a l a l (Gaultheria shallon) analysis of distribution.  V a l e s , D.J., F . L . B u n n e l l , A . M c L e o d , a n d R.M. E l l i s , prep.). Biomass a l l o m e t r i c r e l a t i o n s h i p s o f s a l a l  (Gaultheria  (in  shallon).  W e e t m a n , G . F . , A . G e r m a i n , a n d R. F o u r n i e r . 1 9 8 6 . Fertilizer • screening t r i a l s of stagnated S i t k a s p r u c e p l a n t a t i o n s on n o r t h e r n V a n c o u v e r I s l a n d , B . C . S o i l S c i . S o c . Am. J . (submitted). W h i t e h e a d , F . H . 1973. The r e l a t i o n s h i p between l i g h t i n t e n s i t y a n d r e p r o d u c t i v e c a p a c i t y , p p . 7 3 - 7 5 in: Plant Response t o C l i m a t i c F a c t o r s . P r o c . U p p s a l a Symp., 1 9 7 0 . Unesco, Paris. Y o u n g , D.R. a n d W.K. S m i t h . 1 9 7 9 . I n f l u e n c e o f s u n f l e c k s on the t e m p e r a t u r e a n d water r e l a t i o n s o f two s u b a l p i n e understory congeners. Oecologia. 43:192-205.  Z a v i t k o v s k i , J . 1976. G r o u n d v e g e t a t i o n , b i o m a s s , production, and e f f i c i e n c y of energy u t i l i z a t i o n i n some n o r t h e r n Wisconsin f o r e s t ecosystems. Ecology. 57:694-706.  11 7  CHAPTER  5.  MANAGEMENT  IMPLICATIONS  INTRODUCTION  The for  results  managers.  salal  of  Forest  for moisture  or  stocking  that  managers  interested  suggest height from  to  those  variation must  be  chapters from  of  but of  used  to  will  be  timber  in this  may  of  regime,  caution.  The  be  the  with  illustrate  applied.  be  for  on  can salal  (Bunnell  the  seasonal  in Chapter  previous  additional  deer  et  and  equations  sites  predicting  in modeling  how  or  predicted  data)  the  MCC  forage  Equations  of  of  by  Wildlife  heterogeneity  results  integrated along to  can  canopies  spatial  level  optimum  cruise  aid  a  winter  are  study.  forest  radiation  can  as  purposes  competition  salal.  abundance  radiation  literature  chapters  salal  useful  with  suppress  Salal  two  identify  stocking that  under  because  serve  concerned  in producing  solar  with  published  previous  enough  sampled  the  can  n u t r i e n t s can  forest  environment  1986),  managers  i n v e n t o r i e s (e.g.,  transmission thermal  of  thesis  productivity.  forest  similar  al.  i s high  levels and  this  three  material  results  from  3  1 18  FOREST MANAGEMENT IMPLICATIONS  Tan and  et a l .  Price  et a l .  competitor sites, al.  with  (1986)  have  competitor  plantations  have  et a l . shown  trees  potential  suggested  for nitrogen  i nSitka  Black  Douglas-fir  and can reduce  (1986)  check"  (1977),  that  on n o r t h e r n  that  tree  is a  on d r y  Weetman e t  may b e a  serious  contributing  sitchensis  (Bong.)  Island  (1985),  serious  moisture  growth.  salal  Vancouver  Kelliher  salal  for soil  and phosphorus (Picea  spruce  (1980),  t o "growth  Carr.)  oligotrophic  peat  sites. Attaining densities salal.  Black  guide  than  salal.  Their  stand.  needed  stand  stocking  levels  for salal  diagrams  (Drew  thinning  densities  effectively  and  study  quality  salal  are located  (1979:523),  on a 2 5 - y e a r o l d  Stand  (SDI) c a n be u s e d t o SDI i s n o t  density  1979) c a n g u i d e a rotation.  density except  management  planting and  An SDI t h a t  c a n be m a i n t a i n e d .  a l l plots  t h e number o f  of the trees. index  on t h es t a n d  on average  closure and  suppression.  o r age.  throughout  controls  Flewelling  density  andFlewelling  crown  a r e dynamic,  on a g e a n d s i z e  stand  depending  was b a s e d  stands  (1933)  on s i t e  high  stand  "control"  recommended 1  Reineke's  dependent  this  (1981)  estimate  high  toeffectively  1500 t r e e s « h a " ,  Because  depends  andmaintaining  techniques  potential, t o maintain  Douglas-fir trees  closure  andSpittlehouse  greater  water  suppress  crown  are suggested  densities soil  rapid  Ifplots  diagram  from  o f Drew  3, 7, 2 1 , a n d 22  119  fall  above  above  the  their  crown  "lower  closure  limit  competition-mortality". line  have  salal  management might the at  to  zone  of  9,  near  or  above  less  than  15  shoots«m~ .  control  salal  of  imminent  competition-mortality.  lower  and zone  Maintaining residual  given  to  point  time  in  tree  The  5.1.  As  densities  management The occurs  with  salal  to  dense  are  not  the  to  SDI  1  =  5.1,  at  for  limit  of  densities  planting  and  used trees  calculated 1  /25) •  are  growth.  1200.  of  maintain  the  practical  6  at  (Long  SDI's  of  presented  extraordinarily operational  any  600, in  high  forest  Douglas-fir. which  little  i s not  water  balance  salal.  keep  to  sites  tree  of  higher  1200/(DBH  trees  1  SDI  be  optimum  inter-tree  number  could  An  lower Stand  d i f f e r e n t diameters  using  closure  the  normally  The  needed  in Table  ME«nr 'S" .  enough  trees  seen  suppress  2  1200  an  those  1985).  at  forest  stomatal  250-350  of  density  (1981) m o d e l e d needed  the  to  competition  potential  require  than  1200  regimes  salal  would  ttrees-ha"  number and  1200  i n Nyberg  by:  affect  corresponds  densities  maintain  1000,  Table  of  thinning  required  800,  SDI  above  experiencing  negatively  roughly  an  (densities  1985).  thus  be  are  Douglas-fir  zone  would  or  dry  stands  competition  near  on  the 2  maintain  however,  13-15  imminent  to  zone,  and  Plots  densities  regime  the  Plots  be  this  The  of  line.  Tan  began  Osberg  at  or  known. to  et a  (1986)  below-canopy  no  Black  suggest  a l .  competition  stand  (1977)  photon  flux  suggested energy  and  flux  Spittlehouse densities  reported density that  that of  stands  densities  be under  TABLE  5.1  Tree d e n s i t i e s needed to m a i n t a i n a constant stand d e n s i t y index as average diameter changes  DBH° q  fl  #  (cm)  600  2 4 6 8 10 1 2 14 1 6 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 50  34135 1 1 261 5886 371 5 2599 1 942 1517 1 225 1015 857 736 640 564 500 448 404 367 335 307 283 262 243 226 21 1 198  Diameter Reineke's  of  Trees/ha for 800  tree  stand  4551 4 1 501 4 7848 4953 3466 2589 2023 1 634 1353 1 1 43 982 854 751 667 598 539 489 446 409 377 349 324 302 282 264 of average density  SDI • •  b  1000  1200  56892 18768 9810 6191 4332 3236 2529 2042 1691 1 429 1 227 1067 939 834 747 674 61 1 558 512 471 436 405 377 352 330  68271 22521 1 1772 7429 5199 3883 3034 2451 2030 1715 1 472 1 281 1 1 27 1001 896 808 734 670 614 566 523 486 452 423 396  basal  index.  area.  121  20  2  W«m~ ,  thus  effectively  evapotranspiration solar  irradiance  Using  Osberg's  W«m~  2  above-canopy would  equations  (Table  global  salal,  than  abundances Crown the  time  growing  independent  of age.  of  an u n d e r s t o r y  foliage  7 0 % mean concept  processes  with  900  global 4  Working global  an e s t i m a t e d  Stands  having  an adverse  to touch  may  back  radiation SDI o f 1065  a n SDI  effect  support  on  higher  o f crown  species.  cover  as measured  crown  completeness.  below  because  t h e canopy  however,  Crown may  may  that i s by t h e  o r an  to a cover  t r e e crowns  estimate estimate  are overlapping,  correspond  Crown  as  the entire  of a stand  completeness,  c l o s u r e , but t h i s  precipitation).  a moosehorn,  When  by f o r e s t e r s  and cover  measure  Overstory  in research  acting  throughfall  gives  d e n s i t y and i s analogous  i s 100% c r o w n  useful  begin  i s a measure  tree  only  3 predicting  have  of  c i n Chapter  be z e r o .  or s i d e l i g h t i n g  It i s a useful  of  there  transmission  i s conventionally defined  crowns  area.  a n d a maximum  a t 3% t r a n s m i s s i o n o f  d e n s i t y would  likely  2  W«m" .  expected.  closure  when  moosehorn  effects  than  2  that  days the  800-1000  20 W - n r ,  Coefficient  combined  summer  can range  a n d 984 a t 3%.  1000 w o u l d  but s i t e  2%.  i n Chapter  for a l l plots  sunny  irradiance,  suggested  salal  2% t r a n s m i s s i o n  greater  to  global  the equation  SDI  a canopy  recommended  4.4)  understory  On c l e a r ,  be a b o u t  radiation  through  at  above  (1986)  radiation  from  rate.  reducing  roughly  completeness  i s a  i t i s a surrogate for (light  transmission or  completeness  n o t be a s  as measured  operationally  practical  as using  closure.  Bonnor  moosehorn  t o cover  found higher  that than  the  (1968)  photographs  related  to estimate  measurements  e s t i m a t e s o b t a i n e d from  e s t i m a t e s made  from  h i s e s t i m a t e s made  Relationships given  aerial  between  MCC  the ground from  made  aerial were  aerial  crown with the photos,  about  photographs.  and stand c h a r a c t e r i s t i c s  level  of stocking  c a n be d e v e l o p e d  desired  independent  variable  10%  and used  t o use t o p r e d i c t  fora  i f MCC i s salal.  123  WILDLIFE MANAGEMENT IMPLICATIONS  Because  salal  black-tailed managers winter al.  deer  1986).  t o avoid  on  forest  growth  during  spatially area  predicted natural  production stands  on southern  1980),  on deer  (e.g.,  Nyberg e t  aspects  were  prescriptions for salal conflicts  plot  from  areas  selection,  with  and forage  Management  production forest  change  of salal a mixed  increase  the d i g e s t i b i l i t y  during  which  Arboreal  feeding  deer  were  a large  results Exploiting  e t a l . 1986) i n  responses  i n MCC w i l l  result  i n forage  as a single  1980).  reported  are usually  here.  (Nyberg  was n o t  i n small  relationships are generally  Rochelle  (1961)  presented  patches  l a r g e r change  because  patches  p r e s c r i p t i o n s over  predictable  biomass  a small  of s a l a l  o f MCC may n o t y i e l d  the regressions  Because  cover  and salal  single  a level  may p r o v i d e  proportionately  unwise  homogeneous  heterogeneous,  hyperbolic,  8.0,  forest  andpotential  extensive  openings  patches.  be  sites  1961; R o c h e l l e  forage  stands  developing  t omaintain  small  site  food f o r  objectives.  Because found  t o encourage  Medium  winter  1945; Brown  by m a n i p u l a t i n g  studied good  (Cowan  may w a n t  ranges  i s an important  diet  production  winter  seems  of salal  s p e c i e s may  sarmentosa)  (Rochelle  trials  lasting  fed only  salal  1 7 . 5 , 9.5, a n d 2 2 . 2 % o f b o d y  forage  o r biomass.  toaid digestion  {Alectoria  lichen  i n a  1980).  (e.g., may Brown  3 0 , 4 0 , 4 1 , a n d 71 d a y s resulting  i nlosses of  mass, r e s p e c t i v e l y .  Black-tailed mass of  during  only  o r too  productivity,  although  Estimates  (Dietz  less  1965).  %nitrogen«6.25 as  4.4-6.3%,  5-8%, (1969) of  5-10.8  the  protein  crude  5-7.5%, (1970)  was  4.8 k c a l * g ~  could  1.6  kcal«g~  high  o f the low  gross with  in  an average Island  (converted  mass,  Brown  by R o c h e l l e  in  36 h o u r s . of current  vitro  (1979)  vitro  (1961) Webber  5%, a r a n g e  showed  that  and/or digestibility  volatile  andrate  (1959)  (1980), and  by t h i n n i n g  found  from  by D e a l y  (1976)  eta l .  are  maintenance  Klinka  a t 26-33%,  energy  winter  but  4.0%,  Stanek  over  varied  3.7-9.7%,  (1980)  winter  digestible  a Vancouver  for  protein  be a f f e c t e d  a n d 100%  1  not  (1968)  (1968)  1  in October,  is  Gates  o f 2.0 k c a l * g ~ ,  that  critical  may h a v e  was r e p o r t e d  o f 6.5%  (1985).  Rochelle  24 h o u r s ,  are  ( 1 9 6 1 ) a s 5% d r y  Dice  in  Gates  may b e l o w b e c a u s e  an average  foliage value  movement.  associations  necessary)  over  for  salal  where  fertilization.  Using  the  foliar  i ns a l a l  1  that  Salal  nitrogen  caloric  provided  7% r e q u i r e d  by McCann  salal  short,  than  4.4-8.6%  of  for  crude  (1961)  with  An u n d e r s t o r y  deer  of salal  2.3-3.8%,  1980).  body  species.  Sabhasri  Heilman  of their  sustain  salal  utilization  u p t o 25%  Rochelle  torestrict  of forage  generally  1965;  snowfall,  dense  that  diversity  (Bandy  may l o s e  may t h e r e f o r e  o f heavy  suggested  however,  winter  salal  periods buried  deer,  fatty  acid  of digestion  was 8 8 %  Seip  reported  (1979)  annual  digestibility  growth  o f 32.6%  foliage yielding  energy.  live  biomass  (W) o f 46 k g ( H o v e y  b l a c k - t a i l e d deer  and  dry  mass  1984) daily  125  forage  intake  could  eat  an  1  about  1346  kcal«g  _ 1  of  1  kcal«day"  i n 24  Mautz  requirements On  a  deer  loss  would  diets  volume  are An  of  Gates  higher  optimum  production  require  deer  energy  6  but  61  diet  Jones  During  buried.  Because  burial  rates  are  function  of  depth,  shoots  buried  in  the  of  tall  under 13)  (e.g., Harestad  6 5 - 8 0 % MCC shoots  higher  and  may  range  are  also  canopies, be  more  1979).  in this though,  readily  not  range may  buried  •  •  2.0  0.88 of  around  t o about  a  188  may  energy also  f o r the  may  are  and  than  depth  snow  be  salal  greatest  weaker  forage  forage  to  (Appendix  Mixed  1980).  less  the  only  deer  believed  the  be  1980).  buries  Tallest  decline.  average  however,  snow  have by  g  interception  and  is  acquire  deficit  (Rochelle  shrubs  taller  ( F i g . 2.7)  may  black-tailed  winter, erect  1984)  2473  (900  Rochelle  snow  Tall,  be  days  energy  a  f a t of m e t a b o l i z a b l e  rate  of  1975;  between  1  Salal,  availability.  can  salal  occurred,  intake  reducing  snow  or  translates  loss  loss.  in digestible  i s desired.  7 5  deer  If  (Hanley  energy  kcal«g~  take  winter  tradeoff  2  coefficient  i n an  biomass  biomass  1968;  of  only  deficit  i t would  the  average  in winter.  2»70W°-  eating  (assuming As  an  m e t a b o l i z a b l e energy  decline,  25%  reach a  1961;  day  increment  resulting  1978).  to  (Brown  of  The  diet  by  1  hours)  salal-only  20-50%  A  1984),  m e t a b o l i z a b l e energy  biomass  energy;  (Hanley  forage per  would  energy.  kcal-day" .  g«day~  of  deer  1  1127  g  7 5  (high) a c t i v i t y  • 0.85  digested  g«W -  900  average  kcal'day"  0  50  about  conservative used,  of  14). stems  readily a  linear  of  snow  shoots  are  percentage Shoots (Appendix  passing through  the  1 26  canopy. Salal in  may  be managed  the previous  required  by u s i n g  section.  for salal  Lower SDI's  control  would  biomass  equations  developed i n individual  1985)  used  f o r snow  and forage  that  the regressions  kg-ha"  1  under  8 0 % MCC  under  6 5 % MCC.  day  and  variants,  will  be  Because  MCC  among  and has been  (Bunnell  181  1  a  e t a l . 1985; McNay  be u s e d  in the  examples  important  1  o f FOLBIOM  CAGBIOM  a l l salal,  component  salal  CAGBIOM  i n a n 8 0 % MCC  a n d 537  was  could  of winter  be  kg-ha"  1  salal  2.23  a n d 5.39  Assuming in  immature  and deer  deer«ha"  deer«ha  FOLBIOM  could  intake  100% a v a i l a b l e ,  found  lichens i s  diets.  forage  support  stand  1984),  75  could  of arboreal  and Jones  70% o f t h e d r y m a t t e r 1  2, a n e s t i m a t e d  t h e abundance  (Bunnell  (560 g » d a y ~ ) ,  period  kg-ha"  kg-ha"  Because  was  i n Chapter  a n d 287  stands  a more  salal  stands find  o f CAGBIOM  i n young  become that  MCC  t h a n what w o u l d  no d i f f e r e n c e  interception  presented  follow. From  low  indicated  abundance,  concept  be n e c e s s a r y .  predicted  concept  well,  t h e SDI  _ 1  1  could  over a  i n a 65%  60 MCC  stand. Under likely  forage  however,  open  6 5 % MCC  buried  reduced  buried  a  might  more  provided  by  because  s h o o t s may  o f r e d u c e d snow  availability. have  readily  canopies.  canopy  weaker than  Harestad  shrubs would  Shoots  stems  be  shorter  interception under  denser  a n d more and thus canopies,  which  would  be d i s p l a c e d  shoots having  stout  stems  (1979)  believed  be b u r i e d  by  that  snow  under  75% o f  depths  and more  forage  between  50-110 stems If  cm. and  snow  still be  was  by  65%  by  MCC,  also  deer«ha  - 1  severe  winter  (Jones  1975).  assumptions,  pressure quality  of  the  information  that  functions. burial among  shoot  Shrub  burial  work  burial  levels  may  stands  1  could  of  80%  and  0.21  respectively.  The  >  many  80%  0.11  75  cm  snowpack  i s based deer  on  could  canopy.  reduce  affected  shoot  shoots of  by  work  by  many  be  Continued  abundance  on  should of  MCC. but stand should  example  posture.  and  as  a  the  level  of  growth  done  to  link,  of  I  burial the  forms  peaks MCC  Jay  shrub  include tests  s t r u c t u r e (e.g., be  and  critical  height  take  Because  developing  different Salal  should  is lacking  snow  done  work  be  Additional  an  deer  deer«ha~ in  example  functions  be  occurs age.  0.05  with  as  was  c o n d i t i o n s , FOLBIOM  preceding  burial  closed canopies  or  twice  the and  moderately may  days  than  canopies  supporting  preceding  that  like  f u n c t i o n s among different  ^60  availability  period  severe  of  shoots.  b u r i e d ) , and  and  day  spp.  higher  canopies,  resource  shrub  further  0.02  60  MCC  the  height  shrub  by  1%  readily  Under  65%  for displacement  years.  efforts  on  a  canopy  forage  identified  suggest  65%  subsequent  account  (1985)  and  Although  Modeling into  only  1971-72 h a d  a  only  potentially  i t indicates  on  more  over  consumed,  under  in  being  a l l salal,  80%  to  interception  CAGBIOM  under  supported  snow  account  e r e c t Vaccinium  tall,  respectively. be  not  abundance  stems  find  did  upon  salal  weaker  could  would  based  increased  supported  and  estimate  reduced  (assume offset  His  and  under  where  the  peak  sidelighting)  examine  the  128  consistency to  the  of  location  o v e r s t o r y sample  sensitive  t o MCC  coverage,  technique  weaker  increasing  for  at  support  growth),  or  of  the  too  used). MCC  rapid  work shoot  peak.  design  measurements  further  relationship  of  Attention  because (e.g.,  Because  (Appendix  peak  size salal  13;  of  plot,  shoots  lack  posture  be to  done MCC.  to  be  locations  of  e l o n g a t i o n f o r amount  should  must  verify  other  the  may  be  complete  may  of  given  become shoots  radial  stem  129  REFERENCES  B a n d y , P . J . 1965. A s t u d y o f c o m p a r a t i v e g r o w t h i n f o u r r a c e s o f b l a c k - t a i l e d d e e r . Ph.D. thesis, University of British C o l u m b i a , V a n c o u v e r , B.C. B l a c k , T.A., C S . T a n , a n d J . U . Nnyamah. 1980. Transpiration i n t h i n n e d a n d u n t h i n n e d s t a n d s . Can. J . S o i l S c i . 60:625-631. B l a c k , T.A. a n d D.L. S p i t t l e h o u s e . 1981. M o d e l i n g t h e w a t e r b a l a n c e f o r w a t e r s h e d management, p p . 117-129 in: D.M. Baumgartner ( e d . ) . P r o c . 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(submitted).  133  APPENDIX  Definition  1 Forest  of forest  stand  characteristics  of plots  variables  MCC  - mean c r o w n c o m p l e t e n e s s o f p l o t ( a v e r a g e o f 52 moosehorn samples and r e p o r t e d a s a f r a c t i o n ) NTREES - number o f p l o t t r e e s - h a " £ 8.0 cm NSAPL - # plot saplings«ha" < 8.0 cm a n d > 20 cm t a l l BA - b a s a l a r e a ( m « h a ~ ) o f p l o t t r e e s ^ 8.0 cm SUMDIA - sum o f DBH (mm/225 m ) o f p l o t t r e e s > 8.0 cm AVGDIA - a v e r a g e DBH (mm) o f p l o t t r e e s t 8.0 cm HT - a v e r a g e t r e e h e i g h t (m) o f p l o t t r e e s ^ 8.0 cm HBLC - a v e r a g e h e i g h t t o b a s e o f l i v e c r o w n (m) o f p l o t t r e e s £ 8.0 cm CRNDEP - a v e r a g e c r o w n d e p t h (HT - HBLC) o f p l o t t r e e s > 8. 0cm DF%# - p e r c e n t o f NTREES t h a t a r e D o u g l a s - f i r WH%# - p e r c e n t o f NTREES t h a t a r e w e s t e r n hemlock MISC%# - p e r c e n t o f NTREES o t h e r t h a n D o u g l a s - f i r a n d western hemlock DF%BA - p e r c e n t o f BA o c c u p i e d b y D o u g l a s - f i r WH%BA - p e r c e n t o f BA o c c u p i e d b y w e s t e r n hemlock M I S C % B A - p e r c e n t o f BA o c c u p i e d b y s p e c i e s o t h e r than D o u g l a s - f i r and western hemlock A V G B A D I A - DBH (mm) o f t r e e o f a v e r a g e b a s a l a r e a f o r t r e e s > 8.0 cm SDI - Reineke's " s t a n d d e n s i t y i n d e x " ( R e i n e k e 1933; L o n g 1985) c o m p u t e d f r o m N T R E E S • ( A V G B A D I A / 2 5 ) • 1  1  2  1  2  1  2  BAFBA  6  _ 1  - a v e r a g e s t a n d b a s a l a r e a ( m ' h a ) o f t r e e s ^ 8.0 cm d e t e r m i n e d from 5 BAF„ p r i s m samples BAFDIA - a v e r a g e DBH ( i n mm) o f t r e e s S 8.0 cm s a m p l e d b y 5 BAF prism samples BAFTREES- a v e r a g e number o f t r e e s « h a ~ ^ 8.0 cm d e t e r m i n e d from 5 BAF„ p r i s m samples BAFSAPL - a v e r a g e number o f s a p l i n g s « h a " < 8.0 cm determined from 5 BAF« p r i s m s a m p l e s B A F B A D I A - DBH (mm) o f t r e e o f a v e r a g e b a s a l a r e a determined from p r i s m sampling BAFSDI - Reineke's "stand d e n s i t y index" determined from p r i s m sampling computed from 4  1  1  1  BAFTREES-(BAFBADIA/25) • DOWN UP RIGHT LEFT CENTER  - average basal samples - average b a s a l samples - average basal samples - average b a s a l samples - basal area of  6  area  o f two d o w n s l o p e  area  o f two u p s l o p e  area  o f two r i g h t  area  o f two l e f t  prism  sample  corner  corner  corner corner  i n plot  prism  prism  prism prism  center  APPENDIX  Plot  1  Forest  MCC  stand  NTREES U/ha)  characteristics  NSAPL  BA  ( # / h a ) ( m /ha) J  of plots  SUMDIA AVGDIA (mm/225m ) (mm) I  HT (m)  HBLC (m)  CRNDEP (m)  DF%#  WH°/o#  MISC%#  DF%BA  WH%BA  MISC%BA  1  0 .912  2533  4178  37 . 7  7469  131  12 .4  3 .0  9 .4  25  75  0  28  72  0  2  0.. 782  1600  1378  46 .7  6212  173  13 .6  3. 5  10 . 1  28  36  36  61  23  16  3  0 .554  533  400  21 .2  2559  213  13 .6  2 .0  1 1 .6  50  17  33  68  14  18  4  O,. 793  1778  1911  53 .4  7112  178  16 . 1  8 .6  7. 5  82  18  0  86  14  0  5  0..636  756  44  35 .0  3991  235  25 . 2  17 .9  7. 3  100  0  0  100  0  0  6  0.. 783  1733  1022  34 .4  5660  145  15 .6  9 .8  5 .8  100  0  0  100  0  0  7  0.. 345  267  844  14 .8  1316  219  15 . 7  2 .6  13 . 1  33  67  0  8  92  0  8  0..651  667  800  30 .7  3333  222  18 . 1  5 .0  13 . 1  13  60  27  10  47  43  9  0..899  2711  3467  54. .9  9096  149  15 .0  7.. 1  7 .9  16  82  2  37  62  1  10  0.. 746  1200  711  32. .5  4461  165  15 .5  5.. 1  10 .4  96  4  0  99  1  0  1 1  0. 895  1244  622  61 . .0  5877  210  15..6  6, 3  9.. 3  0  79  21  0  62  38  12  0.,641  844  1822  25 . .5  3401  179  12 .8 .  4 ,0 ,  8 .8 .  68  16  16  91  4  5  13  0. 799  2000  1067  62. 2  8347  185  17 .6 .  9.. 3  8.. 3  62  25  13  78  12  10  14  0. 843  2444  3111  67. 6  9427  171  17.,4  9..9  7 .4 .  67  22  1 1  84  1 1  5  15  0. 792  2089  3289  62 ..0  8534  182  13..9  5..9  7 .9 .  72  7  21  90  2  8  21  0. 576  889  2133  10. 1  2194  1 10  9. 7  0. 8  8. 9  95  5  0  97  3  0.  22  0. 416  756  400  8 .5  2001  1 18  10. 3  0. 5  9. 8  100  0  0  100  0  0.  23  0. 809  2222  5556  41 .6  7206  144  13 .4  2 .1  1 1 .3  48  38  14  74  20  24  0. 636  1111  3200  26. 1  3972  159  14. 3  1 .5  12 .8  56  20  24  64  1 1  25  25  0. 736  1200  431 1  20. 0  3719  138  13. 6  2. 1  1 1 .5  41  33  26  57  22  21  2G  0. 902  2622  7111  38 .9  7576  128  12. 7  3. 4  9. 3  8  85  7  28  62  10  27  o. 757  1378  8000  17 .5  3676  1 19  1 1 .5  2 .8  8. 7  61  32  7  52  30  18  6  APPENDIX  Plot  1  continued  AVGBADIA (mm)  SDI  BAFBA (m'/ha)  BAFDIA (mm)  BAFTREES (#/ha)  BAFSAPL (#/ha)  BAFBADIA (mm)  BAFSDI  UP RIGHT LEFT CENTER DOWN (m*/ha)(m / h a ) ( m ' / h a ) ( m '/ha) (m'/ha) !  1  138  974  30 .4  168  2018  365  138  784  30  26  28  28  40  2  193  1055  40 .0  249  1312  1065  197  896  44  36  36  44  40  3  225  450  30 .4  256  927  0  204  671  34  34  42  26  16  4  196  1201  48 .0  230  1637  1121  193  1084  52  44  50  46  48  5  243  722  36 .8  385  462  0  318  681  36  28  28  36  56  6  159  839  31 .2  229  1 155  1087  185  716  36  32  36  32  20  7  266  295  18 .4  255  730  0  179  428  22  20  20  22  8  8  242  634  32 .8  288  638  0  256  662  32  36  30  38  28  9  161  1335  58 .4  182  3325  2661  150  1461  54  72  54  70  40  10  186  746  38 .4  268  1239  404  199  858  44  36  38  42  32  1 1  250  1243  59 .2  440  1062  731  266  1 176  48  72  66  54  56  12  196  572  28 , .8  234  909  208  201  640  30  30  36  24  24  13  199  1388  53 . .6  241  1711  0  200  1 195  52  48  50  50  68  14  188  1544  59 . 2  222  2546  1033  172  1400  48  60  54  54  80  15  194  1397  41 , .6  253  1361  648  197  932  36  32  44  34  52  21  120  275  12 .0 .  141  1095  5091  118  330  14  14  16  12  4  22  120  233  8 .8 .  144  741  564  123  238  12  8  4  16  4  23  154  1028  31 .2  190  1625  5954  156  767  32  28  36  26  36  24  173  616  25 .6  214  1027  393  178  597  24  26  28  22  28  25  146  505  24 .0  203  1390  5012  148  603  24  22  18  28  40  26  137  1005  31 .2  173  2250  5170  133  818  30  24  28  26  48  27  127  467  21 .6  181  1481  2064  136  561  26  18  22  22  24  136  APPENDIX  2 Means  and 95% c o n f i d e n c e characteristics  Definition  of  understory  intervals  f o runderstory  of plots  variables  2  DENSITY CAGBIOM  - backtransformed average density of shoots«m~ - backtransformed average current annual growth f o l i a r biomass (g«m~ ) p e r p l o t FOLBIOM - b a c k t r a n s f o r m e d a v e r a g e e s t i m a t e d s t a n d i n g c r o p f o l i a r biomass (g«nr ) p e r p l o t (excludes CAGBIOM) TOTBIOM - b a c k t r a n s f o r m e d a v e r a g e e s t i m a t e d s t a n d i n g c r o p f o l i a r a n d woody s t e m b i o m a s s ( g « m ~ ) p e r p l o t ( e x c l u d e s CAGBIOM) PCTCOVER- b a c k t r a n s f o r m e d a v e r a g e q u a d r a t ( 0 . 2 5 m ) p e r c e n t cover of s a l a l per plot REPHT - b a c k t r a n s f o r m e d a v e r a g e r e p o s e h e i g h t (cm) p e r shoot per plot STRHT - b a c k t r a n s f o r m e d a v e r a g e s t r e t c h h e i g h t (cm) p e r shoot per p l o t BASDIA - backtransformed average basal diameter (cm) p e r shoot per plot 2  2  2  2  APPENDIX 2 P l o t  Means DENSITY  and  Q m - ' ) 0 .95  X  95%  confidence  i n t e r v a l s  CAGBIOM  a  C . I  1  0 .65  0 .11-  1 .65  2  10 .83  6 .86-  15 . 7 0  3  40 .58  31 . 0 6 -  51 . 3 7  4  14 . 4 1  10 . 3 3 -  5  2 8 . 16  ( g m ' ) 0 .95  X  for  a  C . I .  understory  c h a r a c t e r i s t i c s  FOLBIOM  ( g m - ' )  X  O. 9 5  3  C. I .  0 .02  0 .00-  0 .08  0 .08  0 .01-  0 . 32  2 . 13  1 .00-  3 .88  5 . 57  2 .96-  of  p l o t s TOTBIOM X  ( g m - ' ) 0 .95  3  C.I  0 .23  0 .02-  0.95  9 .40  20 .07  10 . 0 5 -  35 . 2 0  1 1 .31  7 .91-15 .57  22 .54  15 . 4 7 -  31 . 4 8  106 . 2 7  19 . 16  6 .65  4 .11-10 .06  20 .04  11.48-  32 .05  66 .27  23 . 5 5 -  3 3 . 18  2 8 . 13  2 0 . 0 2 - 3 8 . 17  72 .50  52 . 7 4 -  96 .66  271 .07  189 . 4 2 - 3 7 3 . 34  16 . 4 7  12 . 8 1 -  20 .60  25 .56  15 . 1 5 - 3 9 . 8 8  93 .86  58 . 7 5 - 140 .76  4 0 0 . 28  242 .96-614 .03  7  67 .99  57 . 16-  79 . 75  64 .65  50 .40-81 .37  172 . 6 2  130 . 8 6 - 222 .43  633 .57  466 .92-835 .80  8  13 . 8 4  8.. 7 9 -  6 . 14  2 ,. 9 5 - 1 1 . 0 5  14 . 4 9  7 .27-  25 . 39  0 .00  0,. 0 0 -  0 .01  0 .01  0 .00-  8. 7 3 - 1 9 .74  29 .44  6  9  '  0 . 12  20 .04 0 . 38  0,, 0 1 -  10  15 . 6 5  1 2 ,, 3 7 -  19 . 3 1  13 . 5 0  11  1 .03  0., 3 4 -  2 .09  0 .01  12  20 .89  16. 7 6 -  25 .48  26 .09  13  14 . 5 0  1 1 .13-  1 8 ,. 3 1  5 . 58  3. 4 2 -  14  1 .57  0. 6 3 -  2 ,. 9 3  0 . 18  0. 05-  15  1 2 .. 12  9. 14-  15.. 5 2  21  1 0 2 .. 0 7  8 5 . 5 4 - 120..06  22  1 4 4 .. 4 7  124. 9 2 - 165.,45  23  1 2 .. 6 8  8. 5 9 -  1 7 ., 5 6  24  50. 44  37 . 8 9 -  64 . 77  25  2 4 ..51  16 . 8 3 -  26  5. 66  27  39. 96  Backtransformed  0. 0 0 -  0.03  17 . 6 8 - 3 6 . 8 1  0 . 20  70 . 5 0 - 1 5 2 .45 35 .56-1  46 .81  2 2 .. 3 3 -  0 .07  0 .06  0,. 0 0 -  17 . 4 9 -  45 .87  133,.79  0 .04-  0 .57  0..63  85 . 54  240,.84  10 . 9 6  84 . 72 0 ,. 3 1  80., 8 8 - 2 0 5 , .83 0. 11-  1 ., 8 6  53 .62  30 . 8 3 -  8 .48  1 6 ,. 0 7  7 .99-  28 .31  0 .44  0..29  0 .05-  0 .89  1 .. 3 9  14 . 6 1  8 . 70-22 .73  30,.64  15.. 3 5 -  53 .71  135.,47  5 8 . 6 2 - 2 6 0 . 58  36,.48  30. 90-42 .70  119..57  90,. 8 2 - 153 .83  2 7 8 .. 0 7  211. 9 7 - 3 5 6 . 66  55,. 59  45 . 8 7 - 6 6 . 58  1 1 9 .. 7 9  8 8 .. 6 4 - 1 5 7 . 5 1  2 8 9 . , 17  216 . 7 4 - 3 7 6 .  73.. 33  121 . 1 5 - 4 2 1 , , 0 7 37 . 5 9 - 1 2 6 . ,61 0. 24-  4 ., 2 1  14  5 .49  4 0 ..01  19.. 7 8 -  70 .74  9 2 . 61  46 . 3 6 - 1 6 2 . 43  1 9 ., 3 8  13. 6 8 - 2 6 , .49  72..05  4 6 ., 2 2 - 106 .06  1 6 5 . 15  105. 6 4 - 2 4 3 . 63  33 . 63  7 .. 9 8  4 . 5 3 - 1 2 ..86  6 2 ., 6 7  3 5 . , 4 5 - 1 0 1 ,. 1 2  143. 59  81 . 2 4 - 2 3 1 . 6 8  2. 9 3 -  9. 29  0. 58  29. 17-  52 . 45  13. 99  average  of  3.. 33  52  0.25m'  1 .82-  '  0. 22-  1 .. 2 2  8. 8 9 - 2 0 ..76 quadrats  x  4.  3. 05 97 . 98  6 .. 5 4  7 . 22  6 0 . 2 9 - 1 4 8 .. 8 1  226 . 96  1.09-  2 .63-  15. 34  1 3 9 . 3 6 - 3 4 5 . 19  APPENDIX P l o t  2  continued PCTCOVER  (%)  REPHT  (cm•shoot-')  n  X  0 .95  1  52  0 .38  0 . 0 5 - • 1 .01  44  15 . 4 0  12 . 3 6 - •18 . 7 7  2  52  9 .57  6 . 2 3 - •13 . 6 4  208  15 . 5 0  13 . 6 8 - •17 . 4 3  3  52  17 . 7 3  1 3 . 1 2 -• 2 3 . 0 3  567  17 . 9 3  4  52  17 . 8 8  1 3 . 3 6 -• 2 3 . 0 6  225  5  52  51 . 1 3  4 3 . 8 4 - •58 . 9 8  6  52  38 . 4 7  7  52  8  C. I .  n  X  0 .95  C . I .  STRHT X  n  0 .95  1  )  C. I .  BASDIA n  25 .26  18 . 9 3 - 3 2 . 4 9  44  209  23 . 79  21 . 14- 2 6 . 6 0  16 . 6 7 - •19 . 2 5  569  24 . 2 0  20 .69  18 . 6 0 - •22 . 8 8  227  350  30 .21  27 . 8 9 - •32 . 6 3  3 2 . 7 8 - •44 . 6 1  237  50 .09  50 .33  43 . 5 4 - 57 .61  878  52  1 4 .. 8 3  9 . 3 9 - 21 . 5 0  9  52  0.. 0 6  10  52  40.. 13  1 1  52  12  X  ( c m - s h o o t 0 .95  1  )  C . I .  0 .21  0 . 18- 0 . 2 4  209  0 . 26  0 . 2 4- 0 . 2 8  2 2 . 3 9- 2 6 . 0 8  570  0 .30  O . 2 9- o . 3 2  32 . 22  28 .81 - 3 5 .81  227  0 .29  0 . 2 7- o . 3 2  354  47 . 22  4 3 . 4 6 - 5 1 . 13  354  0 .39  0 . 3 7- 0 . 4 1  4 5 . 5 0 - •54 . 9 1  237  74 .85  6 7 . 5 2- 8 2 . 5 6  237  0 . 54  0,.49 - 0 . 5 9  35 .04  33 . 2 5 - 36 .88  881  .3 . 0 8  40..87 - 4 5 .35  881  0..41  0 . . 3 9- 0 . . 4 2  269  19 . 0 6  17 . 0 8 - 21 . 1 5  269  27 . 3 0  24.. 2 3 - 3 0 . 5 6  269  0..28  0 . . 2 6- 0 . . 3 0  0 .24  12  17 . 2 1  12 . 0 6 - 2 3 . 2 8  12  34 . 76  2 4 .. 0 4 - 4 7 . 4 6  12  0,. 3 4  0. 26 - 0 ..44  32 . 4 5 - 48 .63  221  31 . 2 3  28 . 5 7 - 34 .01  221  59 .06  5 3 .. 7 6- 6 4 . 6 0  221  0..47  0..44 - 0 . . 5 0  0..95  0.. 3 6 - • 1 .81  43  14 . 8 5  12 . 5 9 - •17 . 3 0  44  24 . 39  20..38 - 2 8 .76  43  0..24  0 . 21 - 0 . . 2 7  52  4 4 .. 0 5  35.. 9 0 - 53 .04  279  46 .43  41 . 6 5 - 51 . 4 8  280  67 .64  60..27 - 7 5 .43  280  0. 45  0 . 41'- 0 . ,49  13  52  26..38  20.. 2 1 - 33 . 3 6  217  30 .07  2 6 .. 7 2 - 3 3 . 6 1  217  39 .98  3 5 .. 2 3 - 4 5 . 0 4  217  0. 33  0 . 31 - 0 . , 3 6  14  52  0. 85  1 .54  53  1 9 ..31  1 4 .. 8 8 - 2 4 . 3 2  53  26 .88  20. 22 - 3 4 .49  53  0. 25  0. 20 -0. 30  15  52  28. 98  23., 6 8 - 34 .82  185  5 2 .,61  4 6 ., 5 3 - 5 9 . 0 6  186  76 . 7 5  67. 25 - 8 6 .87  186  0. 49  0 . 44 - 0 . 54  21  52  52 . 87  45..72- 60 . 54  1295  1 9 .. 7 0  18.. 9 5 - 2 0 . 4 7  1296  27 . 23  2 6 . 14 - 2 8 . 3 3  1294  0. 30  0. 29 - 0 . 30  22  52  48 . 66  40.. 4 2 - 57 .65  1751  1 5 ,. 9 6  15 . 4 3 - 16 . 5 0  1751  22 .60  21 . 79-- 2 3 . 4 3  1750  0 . 31  0 . 31 - 0 . 3 2  23  52  12. 22  7 .. 7 1 - 1 7 . 7 7  209  2 7 .. 7 4  2 4 . 4 0 - 3 1 ..31  209  43 .35  37 . 96 - 4 9 .09  209  0 . 37  0 . 35-- 0 . 4 0  24  52  33. 55  25 . 8 1 - 42 . 3 0  682  22. 42  21 . 2 9 - 2 3 . 5 9  682  32 . 3 0  3 0 . 53-- 3 4 . 1 1  680  0. 30  0 . 2 9 - 0 . 31  25  52  23. 05  17. 0 9 - 29 .90  399  28 . 02  25 . 9 9 - 3 0 . 12  399  39 .06  3 6 . 1 1-- 4 2 . 1 2  399  0 . 35  0 . 34 -O. 37  26  52  2 .63  4 .50.  126  16. 98  14 . 6 5 - 19 . 4 9  126  2 4 .. 2 8  2 0 . 83--28 .00  126  0. 27  0 . 25-- 0 . 3 0  27  52  26. 62  20. 6 3 - 33 . 37  608  2 8 . 25  2 6 . 7 7 - 29..77  608  4 3 .. 8 7  41 . 6 2 - - 4 6 . . 18  608  0 . 34  0 . 33-- 0 . 35  0 .00-  0.. 3 6 -  1 .2 6 -  44  (cm•shoot -  4  APPENDIX MCC NTREES NSAPL  C o r r e l a t i o n  3  1 .00 .92  c o e f f i c i e n t s  1 .00 .87 . 79 .97  1 .00 . 55 . 79  -.88  - . 75  AVGDIA  .69 . 83 .93 - .81  HT HBLC  - . 26 . 13  -  .40 .01  -.52 - . 27  CRNDEP DF%#  -  - . 6 0 - . 16  - . 24 - .48 .68 - . 28  BA SUMDIA  WH%# MISC%# DF%BA WH%BA MISC%BA AVGBADIA  -  .01 .08 - . 16 . 24 - . 18 - . 18  BAFTREES BAFSAPL BAFBADIA BAFSDI DOWN UP RIGHT LEFT CENTER  .90 .89 .69  .51 .71 .65 - . 36 . 74  - . 72 .88 . 78 - .62 . 77  .65 . 55  .58 . 59 . 53  -  -  . 55 .61 .61 MCC  WH%BA MISC%BA AVGBADIA SDI BAFBA BAFDIA BAFTREES BAFSAPL BAFBADIA BAFSDI  . 32 -.31 .04 . 1 1 - . 38 - . 92 . 89 .64  SDI BAFBA BAFDIA  .62  - .92 - . 42 - . 22 . 19 . 26 .34 - . 15 .07 .31  .60 .46 NTREES 1 .00 .04 .08 - . 10 - . 23 - .47 .31 .06 - .49 -  .05  DOWN  . 13 .43  -  .39  UP RIGHT  .01 . 33  .01 - .32  LEFT CENTER  .08 . 36 DF%BA  UP RIGHT LEFT  1 .00 .86 .94  CENTER  .32 UP  - .06 - .31 WH%BA 1 .00 . 76 . 30 RIGHT  - . 38 .53 - . 28 - . 74 .67 .40 - . 8 3 .85 . 57 - . 73 .59 . 28 .43 .30 . 39 . 28 NSAPL 1 .00 . 38 - . 26 - . 14 .21 - . 34 - . 3 0 . 34 -.22 - . 19 - .02 -.11 - .07 - . 2 0 MISC%BA  1 .00 . 52 LEFT  among  overstory  1 .00 .91 -  .53 .06  .28 -.62 - .02 .09 - . 12 .23 - . 17 - . 19 - . 6 0 .98 -  .90 .34 .70 .79  -.20 .88 .87 .72 .71 .83 . 75 BA  1 .00 - . 72 -.45 .72 - . 72 - .61 .64  1 .00 - .79 - . 28 . 13 - .65 -.11 . 24 - . 25 . 13 - .01 - . 33 - .84 .97 .78 - . 6 0 .86 -  .82 .47 .86 .73 .68 .64  .03 - . 18 . 32 - . 12 - .05 .43 .97 -.66 - . 36 .81 - . 73 - .62 .77 - . 54 - . 4 0 - .35 - . 39 -.34 - . 13 AVGDIA  1 .00 .87 -.47 .78 .83 - . 34  1 .00 - . 22 . 72 .82 -.11  .90 .83  -.40 - .46  . 72  1 .00 CENTER  1 .00 .63 .21 .49  .72 .61 SUMDIA  -.59 - .46  - .40 - .28 AVGBADIA  v a r i a b l e s  .70 .80 .69 SDI  for  CWHa  1 .00 .86 - . 2 2 .45 - .39 - . 24 .31 - . 34 .01 .52 - . 17 -  .03 .87 -.47 - . 28 .89  (n=10)  1 .00 - .69 .63 - . 50 - .44 .58 - .51 -.31 .08 .21 . 29 .60 - . 16 .08 . 65  1 .00 - . 58 .41 .50 - .68 . 49 .60 .59 - .64 - . 51 .09 . 37 . 54 .02 . 48 .53 . 32 .44  -  . 2 0 .06 . 14 . 25  . 1 1 . 23 .07 .04  -  -  .01 .43 HT  .21 .61  - .38 - . 5 5  .96  1 .00 - . 76 - .56 .96 -.47  .95 .91  - . 22 -.38  .86 .95 .65 BAFBA  - . 39 - . 24 .20 BAFDIA  HBLC  1 .00 .87 - .70 .88 .63 .80 .67 . 74 . 27 BAFTREES  CRNDEP  1 .00 - .50 .90 . 78 .87 .76 .86 .28 BAFSAPL  1 .00 -  .90 .45 .91  -.79 - .50 - .03 - .06 - .01 .45  1 .00 .03 .92  1 .00 - . 21  . 93 . 18  -.11 . 79  - . 10 . 15 .05  . 26 - . 18 - .09  -  - . 5 5  . 1 1  - . 3 5 - . 16 . 38 - . 13 . 16 - . 2 3 .06 - . 15  .51 . 26 - . 51 . 22 - . 15 .27 - .09 .21  - . 26 - . 19 . 17 - . 14 - . 0 6 .02 .04 - . 0 8  .20 DF%#  . 13 WH%#  - . 20 MISC%  1 .00 -.37  1 .00  - . 14 -.28 - .30 - . 15 .31 BAFBADIA  .91 .93 .87 .94 . 52 BAFSDI  1 .00 .83 .90 .88 .58 DOWN  APPENDIX 4 MCC NTREES NSAPL BA SUMDIA AVGDIA HT HBLC CRNDEP DF%# WH%# MISC%# DF%BA WH%BA MISC%BA AVGBADIA SDI BAFBA BAFDIA BAFTREES BAFSAPL BAFBADIA BAFSDI DOWN UP RIGHT LEFT CENTER WH%BA MISC%BA AVGBADIA SDI BAFBA BAFDIA BAFTREES BAFSAPL BAFBADIA BAFSDI DOWN UP RIGHT LEFT CENTER UP RIGHT LEFT CENTER  Correlation 1 .OO . 49 - . 14 .89 .60 .49 .66 .58 - .07 - .60 .62 . 13 - .65 .66 .63 . 52 .81 .93 . 58 .40 .58 . 39 .85 . 79 .85 .93 .90 .77 MCC - .99 - .99 -.98 - . 10 -.55 - .98 .32 - . 24 -.92 - . 30 - .43 - . 79 - .85 - . 54 - . 10 DF%BA 1 .00 . .96 .90 .61 UP  c o e f f i c i e n t s among o v e r s t o r y  1 .00 .58 .82 .99 - .46 .66 .82 - .88 .40 - . 36 - .42 . 34 -.33 - . 36 -.48 .90 . 54 - . 41 .87 .41 - . 58 .72 .51 . 15 . 18 .51 .86 NTREES 1 .00 .98 .97 . 10 . 58 .97 - .27 . 27 .90 . 34 .46 .83 .87 .58 . 13 WH°/„BA 1 .00 .89 .60 RIGHT  1 .00 . 16 .48 - . 75 - . 16 . 10 - .84 .71 - . 73 - . 15 .68 - .68 - .68 - . 73 .28 - . 21 - .63 .48 . 50 - . 75 .01 - . 37 - .43 - .43 - .27 . 16 NSAPL 1 .00 .99 .08 .49 .99 - .40 .20 .96 . 23 . 38 . 72 .81 .48 .04 MISC%BA  1 .00 .87 LEFT  1 .00 .89 . 13 . 75 .77 - .45 - . 19 .21 - .07 - .25 .26 . 24 .1 1 .99 .86 . 18 .64 .50 -.01 .88 . 77 .58 .68 .81 .91 BA  1 .00 - .04 . 39 .99 - .49 . 12 .98 . 12 .29 .66 .74 .39 -.08 AVGBADIA  1 .00 CENTER  variables  1 .00 -.31 .70 .82 - .79 .27 - . 24 - . 32 .21 - .20 - .22 -.34 .95 .63 - . 28 .81 .40 - .45 . 76 .60 . 24 .30 .59 .89 SUMDIA  1 .00 -.02 - .25 .82 - .94 .89 .67 - .94 .91 .97 .98 - .03 . 33 .96 - .57 - .02 .99 .06 .28 . 55 .68 .33 -.11 AVGDIA  1 .00 .80 .03 •72 ^48 - . 17 .86 .73 .48 -.56 .76 .93 SDI  1 .00 .41 .59 .47 .22 .96 .92 .90 .90 .99 .89 BAFBA  f o r CWHbi ( n = 5 )  1 .00 .97 -.37 - . 12 . 22 -.60 - . 16 .20 .09 .01 . 75 .87 - .01 . 76 . 15 - . 15 .93 .95 .65 .60 .90 .92 HT  1 .00 - .45 . 23 .96 . 15 . 28 .67' .76 .40 - .04 BAFDIA  1 .00 - .60 .1 1 - .01 - .68 .06 - .02 - . 13 - . 23 .81 .79 - . 22 .89 . 23 - . 38 .91 .85 . 50 .44 .80 .95 HBLC  1 .00 .46 - .65 .79 .56 . 32 .21 .58 .86 BAFTREES  1 .00 - . 73 .69 .55 - .69 .67 .72 .80 - .58 - . 15 . 74 - .84 - . 37 .88 - .40 -.11 . 20 . 23 - . 12 - . 58 CRNDEP  1 .00 .02 .50 . 14 . 49 . 48 .40 .45 BAFSAPL  1 .00 -.99 -.49 .99 -.99 -.99 -.99 -.03 - .50 - .98 . 36 - . 22 -.94 - . 25 - . 39 - . 77 -.82 - .50 - .04 DF%#  1 .00 - .05 . 16 . 50 .61 . 22 -.23 BAFBADIA  1 .00 .36 - .99 .99 .96 .96 .06 .57 .94 - . 26 . 22 .89 . 34 .47 .83 .85 .58 . 12 WH%#  1 .00 .91 . 78 . 75 .96 .97 BAFSDI  1 .00 - .48 .42 . 58 .60 - . 16 - . 21 .65 -.74 .09 .69 -.41 - . 36 - .06 . 15 - .27 -.46 MISC%,  1 .00 . 78 . 77 .96 .86 DOWN  o  APPENDIX 5  MCC NTREES NSAPL BA SUMDIA AVGDIA HT HBLC CRNDEP DF%# WH%#  msc%# DF%BA WH%BA MISC%BA AVGBADIA SDI BAFBA BAFDIA BAFTREES BAFSAPL BAFBADIA BAFSDI DOWN UP RIGHT LEFT CENTER WH°/oBA MISC%BA AVGBADIA SDI BAFBA BAFDIA BAFTREES BAFSAPL BAFBADIA BAFSDI DOWN UP RIGHT LEFT CENTER UP RIGHT LEFT CENTER  Correlation  1 .00 .88 .90 .82 .86 . 29 .56 .94 .01 - .91 .90 . 34 - .86 .87 . 37 . 34 .84 .90 .49 .94 .61 . 28 .94 .93 . 78 .80 . 78 .89 MCC - .92 - .61 - . 34 - .65 - . 79 - .56 - .83 - . 22 - .31 - .82 - .78 - .63 - . 56 - . 77 - .88 DF%BA 1 .00 .94 . 77 .83 UP  c o e f f i c i e n t s among o v e r s t o r y  1 .00 . 73 .92 . 98 . 24 .44 .81 -.05 -.84 .91 .08 -.70 .85 .02 . 26 .94 .84 . 25 .93 . 58 . 16 .89 .85 .67 . 75 .66 . 79 NTREES 1 .00 . 26 .09 .70 . 72 .25 .95 . 40 .01 .79 .73 .49 .50 .65 .80 WH%BA 1 .00 .62 .71 RIGHT  1 .00 .60 .67 . 10 . 38 .95 -.21 - . 74 . 77 . 17 - .84 .82 .41 . 15 .63 .74 .41 .83 .36 . 17 . 78 .84 . 58 .66 .65 .71 NSAPL  1 .00 .98 .58 .70 .68 .34 -.82 . 79 . 37 - .63 .66 .21 .61 .99 .94 . 52 . 77 .47 .51 .94 .91 .87 .89 .74 .83 BA  1 .00 .67 1 .00 . 18 .55 .69 . 50 .89 .89 . 13 . 12 -.06 -.26 . 74 .98 .45 .60 . 46 .58 . 57 .84 . 37 .69 .58 .60 .55 .57 MISC%BA AVGBADIA  1 .00 .94 LEFT  1 .00 CENTER  variables  1 .00 .42 .59 .76 . 15 - .85 .87 . 24 - .68 .77 . 12 .44 .99 .91 . 39 .87 .54 .34 .93 .90 .79 .83 . 72 .83 SUMDIA  1 .00 .93 . 18 .94 - .46 .20 .86 - .31 .06 .66 .99 .53 .66 .89 .08 -.11 .97 .56 .55 .79 .62 .62 .56 AVGDIA  1 .00 .93 .48 .81 .50 .45 .94 .91 .85 .88 .73 .83 SDI  1 .00 .73 .79 .43 .62 .99 .97 .95 .91 .87 .93 BAFBA  f o r CWHba (n=7)  1 .00 .48 .81 - . 72 .47 .91 - .59 . 36 . 76 .92 .66 .83 .95 . 36 .08 .90 .76 .74 . 88 .70 .84 .81 HT  1 .00 . 23 - .04 .92 .65 .68 .81 .63 . 78 . 70 BAFDIA  1 .00 - . 13 - .89 .92 .25 - .95 .94 .44 .21 .71 .80 .44 .92 .38 . 18 .85 .85 .61 .63 . 75 .85 HBLC  1 .00 .64 .02 .86 .80 .60 .65 .65 .82 BAFTREES  1 .00 - . 21 - .08 .86 -.03 - . 23 . 56 .90 . 27 . 40 . 78 - . 21 - . 16 .90 .29 . 27 . 59 . 37 .45 . 34 CRNDEP  1 .00 - . 19 .48 .43 . 39 .43 . 33 .45 BAFSAPL  1 .00 - .94 - .49 .93 - .89 - .49 - .48 - .83 - .91 - .60 - .88 - .43 - .40 - .94 - .87 -.78 - .69 - .87 - .98 DF%y  1 .00 . 18 - .90 .98 . 24 . 22 .82 .81 .32 .97 .48 . 12 .87 . 79 .61 .61 .71 .87 WH%#  1 .00 .52 1 .00 .55 .97 .78 .91 .64 .89 .57 .86 .95 .50 BAFBADIA BAFSDI  1 .00 - .40 .08 .83 .85 .32 .58 .94 .08 .02 .86 .50 .48 .71 .46 .71 .61 MISC%#  1 .00 .89 .90 .86 .89 DOWN  APPENDIX MCC NTREES NSAPL BA SUMDIA AVGDIA HT  C o r r e l a t i o n  6  1 .00 . 85 . 46 .74 .85 - . 21 .07  HBLC CRNDEP DF%# WH%# MISC%# DF%BA WH%BA MISC%BA AVGBADIA SDI BAFBA BAFDIA BAFTREES BAFSAPL BAFBADIA BAFSDI  -  .08  - .48 - . 2 5 .34 - . 16 - .04 . 16 - . 24  . 10 - .22 .80 .68 .06 .70 . 27 -  .02 . 75 .64  .42 .79 .54  -.32 .88 . 35 - . 35 .68  .58 .62 . 59  .44 .47  .70 MCC  .60 NTREES 1 .00 .21 .21 .06  - . 9 3 - . 56 - . 31  BAFDIA  - . 22 - . 15  - .05 - . 13  - .01 - .06 - . 13 -  -  .50 .42  WH%BA MISC%BA AVGBADIA SDI BAFBA  UP RIGHT LEFT CENTER  -  -.47 - . 35  CENTER  CENTER  .01 . 33 -.61  . 17  DOWN UP RIGHT LEFT  BAFTREES BAFSAPL BAFBADIA BAFSDI DOWN UP RIGHT LEFT  1 .00 .56 .69 .93 - .41  . 28  . 37 .05 - . 15 . 13  c o e f f i c i e n t s  . 0 4 . 26 . 12 . 18  - .03 DF%BA 1 .00 .93 .93 .64 UP  .09 .06 .29 .05  overstory  v a r i a b l e s  for  a l l  three  v a r i a n t s  (n=22)  1 .00  .42  - . 3 2  1 .00 .90 .27 .40 .54  1 .00 - . 10 . 17  1 .00 .74  .38  .49  .27 - .03  - . 18 .08 -.07 - .05  -  -  -.46 - .25  - . 52 -.24  . 35 - .05  .22 . 13 - .06 .05 .06  - . 2 0 . 24 .01 -.61 . 12 -.11 - .52 . 53 .64 -.59 -  .04 . 16 . 18 . 15 . 18  . 13 NSAPL 1 .00 . 33 -  .01 . 13  .42 - . 2 5 - .08  .20 .07 . 16  . 10  .01 WH%BA  . 10 MISC%BA  1 .00 .83 .67 RIGHT  1 .00 .69 LEFT  .26 .99 .93 . 35 .55 - . 13 .31 .92 .86 .80 .86 .82 .87 BA  1 .00 . 13 . 39 .83 - .40 - . 6 0 .83 . 22 . 38 .40 .43 .35 .22 AVGBADIA  1 .00 CENTER  1 .00 .88  . 19 - . 16 .04 .32  - . 0 3 - . 29  . 36 .04 .03 . 22 . 17  - . 10 . 14 .04  among  .01 .08 - . 14 - . 12 .96 . 78 - .04 .79 . 13 -.05 .86 .72 .63 .68 .68 .79 SUMDIA  1 .00 .90 .22 .65 -.03 . 18 .92  .22 . 1 1 . 32 .98 . 14  . 39 .82 -.41 - .61 .88  .70 - . 14 - .41  .56 .07 - . 34  - .01 - . 36 .02  .42 .34  . 36 :49  . 26 AVGDIA  . 54  .82 .79  .95 .94 .94 .94  .86 SDI  .83 BAFBA  .83 .76  .02 - .05 .06 .68 .31 .49  . 28 .46 -.48 . 34 .49  .82 . 36 .49 . 39  . 54 - . 21 .42 .97  1 .00 - .43  - . 19 - . 26 . 25 -.21 - . 2 0 .42 .49 . 59  .21 . 38 . 37  1 .00 .47  1 .00 - . 6 3 . 27  HT  1 .00 - .41 - .47 .93 .27 .43 .47 .50 .40 .37 BAFDIA  .67 .50 .60 .45 .46 .56 .62 HBLC  1 .00 .36 - .46 .71 .49 .52 .43 .56 .48 BAFTREES  -  . 23 .49  -  .41  - . 0 3 - . 4 3 -  .42  -  . 3 0 . 36 . 36 .41  CRNDEP  1 .00 - . 55 -  .09 . 25 .20 .22 .21  - .06 BAFSAPL  1 .OO - .93 -  1 .00 .07 .92  1 .00 - .23  .82 . 58 . 2 2 . 2 5 .28 . 15  .94 . 33 . 12 .24 .26 .05  -  - .33 - . 15  .45 . 19  - . 19 - .07 . 26  -  .44 .92  -  - .01 - .31 - . 19 -.38 -.27  -  -  - .20 DF%#  .30 . 17 WH%#  .06 . 13 MISC%#  1 .00 .21 .41 . 37  1 .00 .93 .92  1 .00 .88  .41 .37  .90 .93  .90 .93  .30  .35 BAFBADIA  .09 .32 . 17 .36 .22  .07 . 74 . 29 .08 . 13 .26  .81 BAFSDI  .07 .08 . 15 .21  . 73 DOWN  to  143  7  APPENDIX  Allometric  equations  total  Allometric sample shoot  of  sizes.  removed from  and  stem  selected  Current  the  nearest  0.01  for  quicker  drying  biomass  diameter,  these  two  used  Slope  plots  2  differences  <  (p  sampled  i n the  transformed  variables,  function.  Fox  and  1970)  equations.  A  regression  was  s  x  's  combination used  to  range  and  into  were  separated weighed  small  hours and  of  were  to  pieces  and  stem+foliar  developed  with  various combinations  among  were  a l l equations on  ln-ln  made w i t h  of  among  in plots  developed  transformed  ln(BASDIA)  variables.  found  Significant  equations 21,  and  22,  in a l l cases  and  24  which  plot  using  stand. were  developed  untransformed  I n d i c e s of were  the  variables.  developed same  cut  biomass  performed  were  random  twigs  hours  f o r 48  biomass)  independent  equations  nonlinear  °C  foliar  tests  tests  0.05)  was  65  and  a  l e a v e s were  f o r 24  h e i g h t , and  were  as  for equations  Allometric ln  relating  Separate  °C  at  l e a v e s and older  material  dried  from  r e p r e s e n t a t i v e of  65  independent  ln(BASDIA •STRHT)  were  and  equation  individual  except  Stem  stretch as  and  equations.  at  foliar  salal  determined  growth  standing crop  basal  for  g.  were be  standing crop  of  A l l living  dried  Equations (total  to  annual  discarded.  material,  weighed.  biomass  relationships  shoots  for  for ln  forward  identify  variables,  determination  computed of  f o r each  the  and best  2  (i ;  and  for  Ezekiel  transformed backward simple  or  stepwise multiple  a and  144  linear  regression  nonlinear computed  regressions with  From equation  the sets  transformed equations,  set.  the lowest  but they  were  s,  2  w  never  untransformed  (Moore  developed a  s  y x  d i d meet  Derivative-free  a•(BASDIA •STRHT)*  subroutine  of equations  equations  while  data  of t h e form  a FORTRAN  having  variance  f o r each  1984). f o r each  selected  better  plot  the  t o be u s e d .  than  the assumption a n d power  were  Log  untransformed of homogeneity of  equations  were  strongly  heteroscedastic. Time shoots  and l o g i s t i c  i n many  were  based  with  similar  different  the  best  applied  on p l o t s  Equations  either  growth  among  equations, similar  plots.  constraints  form  the plots  plots  equations  of s a l a l . used  for estimates  and p l o t s  of f o l i a r  Salal  t o unsampled  growth  form  salal  plots  locations  growth  following  t o which  and t o t a l  of  was  or very  allometric  had shoot  The  clipping  geographic  to develop  plots  sampled.  found  applied  in similar  but a l l unsampled  to those  prevented  form  table  presents  the equations  standing  were  crop  biomass.  E z e k i e l M. a n d K.A. F o x . 1970. M e t h o d s o f c o r r e l a t i o n a n d regression analysis. 3rd ed. John Wiley and Sons, I n c . New Y o r k , NY. M o o r e , C. 1 9 8 4 . UBC British"Columbia,  CURVE. Computing Vancouver, B.C.  Centre.  University  of  1 45 APPENDIX  7  S a l a l s t a n d i n g c r o p f o l i a r and t o t a l biomass allometric equations. Y1=F0LBI0M (g«shoot" ), Y2=TOTBIOM ( g . s h o o t " ) , X1=BASDIA, X2=STRHT, X 3 = X 1 , X 4 = X 2 , X 5 = X 1 « X 2 , X6=X3-X2, X7=X1-X4, X8=X3-X4 1  1  2  Plots  2  sampled  n  Equation  2  s  ^  Y1  =  0.02539-X2  42  0. 40  0. 66  1  Y2  =  0.3957-X5-0 .00224-X7  42  0. 76  1 .36  b  Y1  =  0.2921-X5-0 .000759-X40.4598-X6+0 .00297-X8  1 17  0. 52  0. 60  0. 87  1 .57  a  2  2  3  Y2  C  3  4 5  e  5  =  7.512-X3-0. 00489-X4+0. 7451-X51.352-X6+0. 0 1 2 8 1 « X 7 11 7  Y1  =  0.31695-X6  0  '60968  97  0. 53  0. 59  Y2  =  1.10036-X6  0  • 7 6 6 3 7  97  0. 88  1 .35  Y1  =  0.39828-X6  0  • 7 8 17 0  76  0. 77  1 .53  Y2  =  0.89957-X6  0  • 9 18 9 7  76  0. 93  3. 15  Y1  =  0.40376-X6  0  • 7 2 2 0 9  93  0. 81  3. 05  0.99106-X6  0  • 8 3 6 8 H  93  0. 90  9. 31  32  0. 95  2. 1 1  Y2 Y1  =  0.07619-X6  1  • 0 3 16 7  Y2  =  0.42300-X6  0  '98053  32  0. 98  5. 02  Y1  =  0.02246-X6  1  • 1 5 9 8 4  49  0. 85  10. 10  15  Y2  =  0.01716-X6  1  • 4 9 1 3 2  49  0. 85  81 . 31  ,22,24^  Y1 =  151  0. 85  1 .16  0 . 6 2 8 7 - 0 . 1 0 2 4 - X 2 + 0 . 3 9 7 1 •X50.3617^X6+0.00510«X7 151  0. 93  1 .86  10 1 5  s  Y2 =  ,22,24  -0.000784-X4+0.1208-X50.1942-X6+0.00352-X7  ^Equation  used  for plots  ^Equation  used  for plot  2.  c  Equation  used  for plot  3.  ^Equation  used  for plots  4 and  8.  ^Equation  1,  9,  and  used  for plots  5,  and  Equation  used  for plots  10,  ^Equation  used  for plots  12 a n d  Equation  used  for plots  21  6,  11.  7.  f J  R  13, a n d  -  15.  27.  14.  APPENDIX  8  Average biomass ( i n grams) o f s a l a l l e a v e s by p l o t a n d 9 5 % c o n f i d e n c e i n t e r v a l s ( b a c k t r a n s f o r m e d from square r o o t )  n  Plot  0.95  X  Confidence i n t .  1  330  0.0987  0.0914  -  0.1062  2  315  0. 1239  0.1148  -  0.1333  3  847  0.1133  0.1082  -  0. 1184  4  506  0.1432  0.1352  -  0.1513  5  646  0.2306  0.2188  -  0.2426  6  620  0.2429  0.2324  -  0.2536  7  672  0.1941  0.1839  -  0.2047  8  653  0.1294  0.1215  -  0.1375  9  1 62  0.0728  0.0644  -  0.0817  10  792  0.1831  0.1745  -  0.1919  12  • 865  0.2049  0.1954  -  0.2147  13  809  0.1774  0.1700  -  0.1850  1 4  583  0.1359  0. 1286 -  0.1434  1 5  922  0.1839  0.1762  -  0.1918  21  1 1 87  0.1338  0.1281  -  0.1395  22  1 450  0.1204  0.1154  -  0.1256  23  1 245  0.1118  0.1075  -  0.1161  24  1115  0.1303  0.1244  -  0.1363  25  1 003  0.1394  0.1337  -  0.1453  26  627  0.0935  0.0887  -  0.0985  27  1 094  0.1524  0.1462  -  0.1587  11°  a  Leaf  biomass  from  UBCRF  plot  8 was  used.  147  APPENDIX  9  Regressions of s a l a l v a r i a b l e s against plot and p r i s m b a s a l a r e a s a n d s t a n d d e n s i t i e s . Sample s i z e s : CWHa=10, CWHb,=5, CWHb = 7. Equations s i g n i f i c a n t a t p £ 0.05. R e g r e s s i o n e q u a t i o n : Y = b b ,X 3  t  Y  Variant  X  b^  bo  r  2  s  yx  DENSITY:  CWHa+b,  a  1/NTREES 1/BA 1/SDI 1/BAFTREES 1/BAFBA  N.S.* -14.094 -11 .505 N.S. -23.725  1 7731 1111.1 22351.0 19106 1499.0  0.86 0.81 0.86 0.31 0.68  6.73 8.02 6.97 1 4.76 10.47  CWHb  1/NTREES 1/BA 1/SDI 1/BAFTREES 1/BAFBA  -61.237 -25.221 -31 .878 -74.65 -34.597  140890 1350.8 38548.0 158560 1602.6  0.88 0.92 0.93 0.85 0.95  19.57 15.71 14.96 21 .84 12.01  CAGBIOM:  A l l3  1/NTREES• 1/BA 1/SDI 1/BAFTREES 1/BAFBA  N.S. N.S. N.S. N.S. N.S.  1 6807 442.33 11170 18784 471.21  0.54 0.59 0.70 0.34 0.56  12.09 1 1 .60 10.67 14.72 1 2.27  FOLBIOM:  A l l3  1/NTREES 1/BA 1/SDI BAFTREES BAFBA  N.S. N.S. N.S. 100.91 135.61  46956 1293.1 32432 -0.03828 -2.5425  0.42 0.59 0.66 0.29 0.57  37.24 31.17 29. 1 1 41 .86 32.52  TOTBIOM:  A l l3  1/NTREES 1/BA 1/SDI BAFTREES BAFBA  N.S. N.S. N.S. 331.49 385.04  156470 3785.9 97566 -0.12924 -6.7463  0.52 0.33 0.45 0.31 0.38  108.42 128.89 1 17.05 133.61 126.56  PCTCOVER:  A l l3  NTREES BA SDI BAFTREES BAFBA  49.774 47.809 50.226 50.373 51 .843  -0.0170 -0.63628 -0.3043 -0.01850 -0.78680  0.47 0.39 0.44 0.47 0.38  13.66 14.66 14.01 13.69 14.75  3  !  Prefix  1/ r e f e r s  'Coefficient  to the reciprocal  not s i g n i f i c a n t  (p >  of the independent  0.10).  variable,  148  0.6  0.5 H  E 3  CC UJ  0.4 H  i  hUJ  <  0.3  4  it  i  Q _J  < CO  <  0.2  CO O.H  0.0  30  40  50  60  70  80  90  100  MEAN CROWN COMPLETENESS (%)  Appendix 10. The r e l a t i o n s h i p o f a v e r a g e b a s a l d i a m e t e r of s a l a l s h o o t s a n d 9 5 % c o n f i d e n c e i n t e r v a l s t o mean c r o w n completeness.  149  175.0 - i  150.0-  I  100  MEAN CROWN COMPLETENESS (%)  A p p e n d i x 11. The r e l a t i o n s h i p o f twig p r o d u c t i v i t y (# CAG twigs) and 95% c o n f i d e n c e i n t e r v a l s t o mean c r o w n c o m p l e t e n e s s . E q u a t i o n : # C A G t w i g s •nr = 193.81 219.93-MCC r =0.9l, s =10.7. 2  2  #  150  20.0 n  2  2  Appendix 12. The r e l a t i o n s h i p o f s a l a l b a s a l a r e a (cm /m ) a n d 9 5 % c o n f i d e n c e i n t e r v a l s t o mean c r o w n c o m p l e t e n e s s . E q u a t i o n : B a s a l a r e a = 20.57 - 2 2 . 8 0 » M C C r =0.76, 2  151  APPENDIX  13 T h e r e l a t i o n s h i p  of s a l a l  crown  The  relationship  overstory  cover  posture  termed  heights  by: FI  FI  indicates  from  positive  A  relationship f o r combined  calculated  posture and  index  termed  strongly  increasing  potential  plot  index of  ( F I ) was d e r i v e d  of each  shoot  repose  repose  shoot  from  shoot  Although  variable,  of p o t e n t i a l 2  "flimsiness  maximum  MCC,  with a  repose  t o MCC i s  size  p  was  <  0.01).  also noted.  2" was d e v e l o p e d  from:  diameter)//!  ( F i g . 13b), i n d i c a t i n g  were  height  The  significant,  to overstory cover index  increasingly  per unit  a l l  95% c o n f i d e n c e  5^^=5.6%,  f o r shoot  t o MCC  shoots  (FI = 0 i f  "flimsiness"  (r =0.29,  accounting  related  i s reduced  height=stretch height).  i s illustrated  data  that  height  2 = 2( (STRETCH-REPOSE)/STRETCH/Basal  2 was  under  percent  and t h e r e l a t i o n s h i p  index, FI  have  o f F I t o MCC  second  shoot  An a v e r a g e  index"  maximum  ( F i g . 13a).  indicated  t o mean  (Z((STRETCH-REPOSE)/STRETCH)/n)•100  in a plot  intervals  salal  investigated.  the average  relationship  FI  =  posture  completeness  between  "flimsiness  the potential  shoots  This  was  shoot  basal  reduced  that from  diameter.  A p p e n d i x 13a. The r e l a t i o n s h i p of s a l a l shoot p o s t u r e ( f l i m s i n e s s i n d e x ) a n d 9 5 % c o n f i d e n c e i n t e r v a l s t o mean crown completeness.  1.5-i  1.3-  1.1-  ffl  0.9  0.7 A A  0.5  0.3 0.3  I  I  I  I  I  I  0.4  0.5  0.6  0.7  0.8  0.9  I  1.0  MEAN CROWN COMPLETENESS (%)  A p p e n d i x 13b. The r e l a t i o n s h i p o f s a l a l s h o o t posture ( f l i m s i n e s s i n d e x 2) t o mean c r o w n c o m p l e t e n e s s .  APPENDIX  14  D i s t r i b u t i o n of repose by p l o t . Distribution d e n s i t y i n each height  Repose  'lot  heights of s a l a l shoots i s percent of shoot class  height  class  (cm)  0-19  20-39  1  67.3  26. 1  6.5  0.  0.  0.  0.  2  51 .4  21.9  9.3  2.4  1 .0  0.  0.  3  58.4  26.1  12.1  3.5  0.  0.  0.  4  53.9  27.5  13.5  3.3  1 .6  0.  0.  5  38.2  32.7  15.6  8.3  3.8  1 .3  0.9  6  20.8  19.2  19.6  16.0  10.0  8.4  6.4  7  32.7  27.6  19.5  10.1  6.1  2.3  1 .5  8  55.7  26.2  11.4  4.4  1 .8  0.4  0.  9  61 .6  38.5  0.  0.  0.  0.  0.  10  30.3  42.2  16.3  6.8  4.0  0.4  0.4  1 1  68.2  31.9  0.  0.  0.  0.  0.  12  34.0  16.6  14.2  9.3  10.4  5.4  9.8  13  40.0  29.7  14.6  7.9  2.2  1 .2  3.9  14  60.8  25. 1  9.0  3.6  0.  0.  1 .8  15  24.0  15.1  15.2  15.7  13.1  21  55.6  33.4  8.8  1 .8  0.3  0.  0.1  22  66.3  29. 1  4.3  0.4  0.  0.  0.  23  43.8  27.8  13.9  8.8  3.4  0.9  1 .2  24  50.5  36.2  10.0  2.9  0.2  0.1  0.1  25  40.0  34.3  15.5  6.2  2.2  1. 1  0.4  26  67.2  24.0  7.6  1 .4  0.  0.  0.  27  35.0  42.2  15.4  4.1  2.4  0.6  0.6  40-59  60-79  80-99  100-119  5.2  120 +  11.2  155  APPENDIX 15  Percent  frequency  of o c c u r r e n c e  Speciei  1 Gauiint'ia  shot  Votcimum Mohoni  pan  a ntr  Ion i  folium  Thuja  96.2  8 6 ..5  51 ,. 9  3.8  19.2  21.2  1 5 ..4  36.5  46.2  2 3 . 1 59.6 1 3 . . 5  5.8  7.7  36.5  plicata  obi onti/ol1  Li imeta  boftal  Cr ami noea Hi t I oc I un  t  i chum  Ac hi,1  9.6  21.2  13.5  46.2  17.3  28.8  1.9 2 6 . 9  1.9 1 1 . 5 3.8  7.7  1 .9  19.2  11.5  1.9  7.7 3.8  .9 9.6  5.8  s mar gari  7.7  3.6  1 .9  5.8  3.8  55.6  66.r  19.2 6 3 . 5 1.9  7. 7  1.9 1 3 . 5  7. 7  13.5  9. 6  1.9  5.8  1.9  5.8  1.9 1 1 . 5  1.9  5. 8  3.8  9.6  1.9  1 .9  1.9  34. 6 9. 6  3.8 1 .9  1.9 15.4  15.4  1.9  9. 6  1.9  15.4  3. 8  1.9  1.9  1.9  21 . 2 3 2 . 7  1.9  1.9 7.7  1.9  3 . S i . 9  1.9  5.0  7.7  3 2 . 7 15.4  19.2 6 5 . 4  1.9 11.5  3.8 5.8  11.5  J.9  7.7  HIII  9.6  1  s pp.  t,9  1,9  1.9 \ .9  t ( f oi i um  abi (11  9.6  5.B  _g  3.8  1.9  5 g  3.8 1.9  1.9  col or  7.7  y g \  1.9  Hoi odi  g  j j  3.8  brtrifolia  3  t9.2  aimifolia  angut  1.9 g_g  1.9  racta  13.5  1  5 . 8 5 . 8  40.4  ttrtpiopotdei  di  5.8  27  1.9  19  Taxut  cut  94. 2 82.7  26  3. 8  ' •9  hier  25  1.9 19. 2 2 1 . 2  i cot a  Strtptopus  24  7.7  1.9  1.9  36.5  1 pp.  tptct  23. 1  3.8  7.7  Hoi a gymnocarpa  a  3.8  1.9 21 . 2  5.8  3.8  1 .9 ,  1.9  noot kai*  omebi 1I i  Eptlabium  23  9 8 . 1 84.6  5.8  mom  Ammtlone  99. 9  22  61 .5 1 7 . 3  5.6  11.5  21  53. 6 86. 5 59.6 3 2 . 7  1.9  5.8 42.3  tcouienana  Pyr ola  92.3  3.6  3.8  5.8  Abiet  Mubut  1.9  aquitinum  Pi nus  1  92.3  11.5  3.. 8  t.9  at a  Chamateyperit  Mom  96.2  11.5 4 0 . 4  9..6  1.9  iriphylla  Anaphait Sat i x  96.2 3 2 . 7  15.4  15  1. 9  latifolia  Pitridtum  12.3  13.5  13.5  1,9  mum t um  orbicut  Trtemalii  14  1.9 at at um  citiota  Viola  1 3  7 . 7  b'qut s el um s pp. yil  12  7.7  atbijlorum  jpp.  Lontcira  Plot n u m b e r 10 11  80.8  1.9  1.9  Mai ant hemum diI  fot  9  40.4  3.8  1 .,9  8  99.9  1.9  1.9  tpp.  Attn tri  7  in quadrats  7.7  a  iI  spp.  Ft agaria  9 8 . 1 94.2  found  3.6  3.8  spp.  Goadytra  6  76.9  1.9  Cblmopkilo  5  84. 6 30.8 19.2  1«  4  32.7  9010  ht 11r opkpt  Ttuga  3  21.2  urjfnuj  ftubui  2  o l species  3.9 1,9  APPENDIX  16  Average quadrat  1  (0.25 m )  d e n s i t y of u n d e r s t o r y  species other  Spftcies  Plot 1  Vaccinium  par rl  kanonta  nerrota  Xv.iif  ur 11  Tsuga  heteropnyl1  Pteudol  1 .60 1 .B7  Li i t a t a  boreal  .04 0.89 0.04  0.19  0.71  0.98  1.29 0.23 0.04  0.02 0.  menitetti  0.02  0.02 0.04  0.10  0.04  0.08 0.02  0.06  0. 19  0.04  0.02  ipp.  0.19  Agoterit  tpp.  0.04  Vi ol a  Achl ys  munition  0.02  0.38  0.12  .35  0.12 0.06  1 5  0.33 0,.58  21  1 .94 0.  54  0.04  lattfolla  0.04  Abiet  amabi l i s mom i col a  Ammei anchi  0.,02  0 .04 0..06  0..52  0.,27  0.,15  0.,02  0.  0 .08  0,.13  0 .15  0..25  0.02  0.04  0.,10  0.,02  0..06  0 .02  0,.06  0 .02  0 .12  0.,02  0 .06  0,.04 0 .10  0..02  0,.02  0 .02  0 .25  0 .02  0 .46  0 .06  0. 1 7  0 .02  0 .06  0.02 0.02  .13  0..04  1,.60  0,.08  0,.15  0,.02  0..58  1,. 17  0.02  0.02  0 .04 0 .02  0.02  0 .08  0 .08  1 .69  0.48  0  2.23 0.58  .54  0 .02 0 .42  0 .10 0.06  0 .12  0 .02  0 .04  0.04  0., 10 0 .04 0 .56 0 .08  0 .08  0  .10  0..08  0 .13  0.,09  0.04 0.02 0.04  ai ni fol  i a  0.04  0.01  0.13 0.02 0.02  discolor  0.02  0.02 0.08  angutIifofiurn  Jptetabi11  dtts  s  0.02  0.17 0.19  11reptopoi  Mont i a perfoliate Epilobium  0.  0 .48  0. OS  0.25 0.21  0.06  breti/oiia  Hot odi i cut  0 . 35 0., 1 50.  0.,02  0.02  tr  ,71  0.,77  1 .40  0.14  num  Pi nu,  opus  .19  27  0.,12  root tat eHIi s  lymnocarpa  26  0. 58  1.42  ertana  Rosa  25  0.04  mar gart t ace a  a ipp.  24  0.02  0 .02  0.25  a  aaui1i  Pyrcl  23  0.06 0,.08  0.13  0.02  or bi cul at a  j caul  22  0. 54 0. 88  0.10 0..10  0.06 0.02  0.04  0. 08  ,15  0.10  1  0.02 0.06  0. 0.27  1.29 0.21  14  0.06  0.04  Chamateyparii  Sub MI  0.27  13  0.02  t rtphyl1  Sal 1 x  Taxus  0.,27  0.12  0.35  0. 10  d i l a t e  Anaphalii  15  0.65  12  0.23  ipp.  Pleridium  Strtpi  arum  di 1 at et um  Trienlelti  0.06 0.08  number  10 11  0.04  um el b i l l  e era  0..14  9  0.15 0.04  Progeria  Lam  8  salal  0.10  11  Potystictium  0. 40  0.08  Ht tract  Earn j el um  0.31  7  0.50  ipp.  Mai ant Annum  6  0.04  oblongtfolia  Gremi i i a r a  5  0.42  a t pp.  Gaodyere  1  4  a  pi I eat a  Cki mepni1  3  0.46 0.48  nuj  luge  Thuja  folium  2  than  0.04 0.02  157  APPEMDII  17  List  of  cover clesse*  for  species  present  in  plots"  2i  TWOS: AOI ri  omoti 11 $  Al nut  i aifo  Ckomotc ppmr 11 ft aut  moot tot tot 11  mootIeol  Pt todolimgo  o  miott tn  Totui  oriwIfolI  Tkujo  fli  I  0  coio  Ttugo  ktltropkfil*  SHRUBS BBd  m m  Mat / «ac Cktmopkl I a Cloiol  i  ol ol f ol t o  «r  too.  kammut pjroi  I fl  Gout i ktno  tkotto*  Holtdttcoi  4i t color  Li oooto  orut  i op.  Lovi I I I M  tor tot  Lornctro  it  cl11oio  Lycopodl  um  kokom o  iff.  otrtoio  fol fttitkom  moot i M  ft tri di *M aami 11 * u Asia  gymoocor po  Rubm  ipoet obi 11 t  Kubui  urn  Solii  nut  icoulerigmo  Vottintum  otol i fol  Vaccintum  porn  i urn  folium  HSRBS: Ackl  f  ,  tri pkyl I o  Agotom  iff.  Aoopkelti  mar for I I Otto  Epttootum  oagutt  Ftagona  ifp.  Goodytro  ool oogi f el I e  Grcmi not a Httroti  i fol i um  tpp.  um olbt  ftorum  Mot em ktmum 41  I I K I M  *ff Ori km fyrola  m  ttcuoda tpf.  St *tpi oput  n r t fiopoi  Trltotoiit  4*t  leufolie o i i n i  Tri Hi  um  fl olo  orbi cut oio  MOSSKS end LICK CMSi Clo4.io  tpp.  Dlc r ooum I pp. Html Iomti  C0*(f 11 MH  Hyiocomlum Kt odbt r ft a  tpltodtot ortgooo  PI egi ot keci mm uodut at mm fbtytriekum  /  aoiptrioum  *k,t i dl a4tt pkm  I or tut  Mkyt i4i ad*I pkui  tn  cUsses  omtt r\  Klinke  et  et.  (I9flt>:  •  -  <\\, I -  i-< |, 5  2 -  5-<2Sl,  3 -  25-«5M.  4 -  50-<75\,  5 -  i75».  25  26  27  158  APPENDIX  Plot  18  Results  Hygrotope  of  site  diagnosis  Trophotope  Grid  No.  Site  unit  1  3  B-C  7  4  2  2-3  B-C  7  2  3  2  B  7  2  4  2-3  B-C  7  2  5  3  B-C  7  4  6  3  B-C  7  2  7  2-3  B-C  7  2-4  8  2-3  B-C  7  2  9  2-3  B-C  7  2  10  2-3  B-C  7  2  1 1  2-3  B  9  2  12  2  B  9  2  13  2-3  B-C  9  2  14  2-3  B-C  9  2  15  2  B  9  2  21  2-3  B-C  11  2  22  2-3  B-C  11  2  23  2-3  B-C  11  2  24  2-3  B-C  11  2  25  2-3  B-C  11  2  26  3  B-C  11  2  27  2-3  B-C  1 1  2  APPENDIX  19  R e g r e s s i o n c o e f f i c i e n t s of equations p r e d i c t i n g s o l a r r a d i a t i o n components from stand c h a r a c t e r i s t i c s . S a m p l e s i z e s : CWHa=6, CWHb=6. Equations s i g n i f i c a n t at  Y  p  £  0.10.  X  Subzone  TOTAL:  DIFFUSE:  a  PAR:  Y = ae  b  NTREES BA BAFBA  1 .0716 1.6708 5.8482  3.1539E-3 8.6639E-2 0.1381  CWHb  NTREES BA BAFBA  8.0679 1.3098 1.3583  Both  NTREES BA BAFBA  CWHa  NTREES BA  /  2  0.89 0.97 0.92  0.0610 0.0324 0.0485  3.5850E-3 0.1193 0.1097  0.92 0.84 0.93  0.0624 0.0880 0.0579  0.6640 0.8840 1.0616  1 . 1225E-3 7.2005E-2 7.9569E-2  0.63 0.79 0.77  0.1143 0.0850 0.0895  0.7272 1.2563  5.0976E-4 3.7167E-2  0.58 0.66  0.1527 0.1339  0.48  0.1709  N.S.*  0  CWHb  NTREES BA BAFBA  3.0165 2.3034 2.6304  7.5458E-4 3.5136E-2 4.3310E-2  0.93 0.89 0.80  0 . 1598' 0.2013 0.2727  CWHa  NTREES BA BAFBA  1.3409 2. 1132 11.2441  4.2749E-3 0.1078 0.1777  0.95 0.98 0.96  0.0394 0.0247 0.0353  CWHb  NTREES BA BAFBA  230.5546 213.0794 9.3088  8.3632E-3 0.7341 0.3536  0.99 0.98 0.98  0.0204 0.0263 0.0247  Both  NTREES BA BAFBA  0.7036 0.7069 0.8231  1.7435E-3 8.5952E-2 9.0618E-2  0.69 0.59 0.55  0.0885 0.1031 0.1070  CWHa  NTREES BA BAFBA  0.4157 0.8569 2.2549  1.6161E-3 7.4090E-2 0.1131  0.86 0.95 0.88  0.0430 0.0264 0.0396  CWHb  NTREES BA BAFBA  0.8099 0.4242 0.4800  1.5746E-3 6.7017E-2 7.1875E-2  0.88 0.85 0.83  0.0380 0.0419 0.0449  Both  NTREES BA BAFBA  0.3845 0.4213 0.4691  1.0378E-3 5.6009E-2 6.1045E-2  0.75 0.78 0.68  0.0515 0.0481 0.0579  Exponent  for  power  ^Equation  not  significant  a  equation:  CWHa  BAFBA  DIRECT:  Regression  of  10. (p > 0 . 1 0 ) .  APPENDIX  20  R e g r e s s i o n c o e f f i c i e n t s of e q u a t i o n s p r e d i c t i n g s o l a r r a d i a t i o n components from s t a n d c h a r a c t e r i s t i c s . S a m p l e s i z e s : CWHa=6, CWHb=6. Equations significant at  Y  p £ 0.05.  Regression  X  Subzone  equation :  b  Y =  /  2  s  yx TOTAL:  DIFFUSE:  DIRECT:  PAR:  CWHa  MCC  2.9899  SUMDIA SDI BAFSDI  5.5965E-3 2.8797E-3 2.7659E-3  0.85  0.0713  0.95 0.95 0.70  0.0414 0.0396 0.1028  CWHb  MCC SUMDIA SDI BAFSDI  2.3816 4.4934E-4 3.5185E-3 3.2876E-3  0.69 0.80 0.84 0.91  0 . 1232 0.0983 0.0878 0.0663  Both  MCC SUMDIA SDI BAFSDI  2.6362 4.8113E-4 3.2665E-3 3.0606E-3  0.73 0.85 0.87 0.81  0.0974 0.0725 0.0665 0.0817  CWHa  MCC SUMDIA SDI  1.4802 2.0949E-4  0.74 0.71 0.69  0.1217 0.1280 0.1310  1.3024E-3  a  BAFSDI  N.S.  CWHb  MCC SUMDIA SDI BAFSDI  N.S. N.S. N.S. N.S.  CWHa  MCC SUMDIA SDI BAFSDI  3.2988 6.4419E-4 3.2266E-3 3.0370E-3  0.84 0.98 0.95 0.68  0.0731 0.0284 0.0389 0.1015  CWHb  MCC SUMDIA SDI BAFSDI  3.2984 6.6399E-4 5.3104E-3 4.8240E-3  0.66 0.70 0.77 0.86  0 . 1032 0.0970 0.0851 0.0659  Both  MCC SUMDIA SDI BAFSDI  3.2985 6.5725E-4 4.4815E-3 4.0347E-3  0.75 0.84 0.73 0.63  0.0800 0.0829 0.0975 0.0731  CWHa  MCC SUMDIA SDI BAFSDI  3.7696 8.9596E-4 3.9498E-3 3.3928E-3  0.95 0.79 0.86  0.0270 0.0526 0.0346 0.0426  CWHb  MCC SUMDIA SDI BAFSDI  3.2611 6.6389E-4 5.4516E-3 4.9271E-3  0.67 0.88 0.74 0.65  6.0636 0.0378 0.0564 0.0648  Both  MCC SUMDIA SDI BAFSDI  3.4824 7.3004E-4 4.9477E-3 4.2759E-3  0.79 0.77 0.74 0.60  0.0473 0.0496 0.0521 0.0649  °Exponent  f o r power  ^Equation  not  of  10.  significant  {p  >  0.05).  6  0.91  APPENDIX  21  Regression c o e f f i c i e n t s of equations predicting s a l a l from s i t e f a c t o r s d e r i v e d from h e m i s p h e r i c a l p h o t o g r a p h s . S a m p l e s i z e s : CWHa=5, CWHb=6. Regression  Y  DENSITY:  SALALBA:  TOTBIOM:  CAGBIOM:  PCTCOVER :  Subzone  equation:  X  Y = 1 +°fl(x-c)/fc  c  2  s  a  b  CWHa D I F F U S E S F DIRECT SF  844 . 3 3 .3  105.8 -2.5  0. 0920 - 0 . 4638  0 . 96 0 . 86  5 .7 11 . 4  <0 .01 <0 . 0 5  CWHb D I F F U S E S F DIRECT SF  707 . 9 1 127. 5  46781946 83915.0  0 . 071 1 0. 0889  0 . 92 0 . 55  17 . 0 40 . 5  <0 .01 0 .09  B o t h D I F F U S E SF DIRECT SF  992 . 5 587 . 6  208.8 225.2  0. 0826 0. 0653  0 . 70 0. 1 3  26 . 0 44 . 5  <0 .01 0 .27  CWHa D I F F U S E S F DIRECT SF  173 . 8 1 06 . 7  21 . 4 11327 .4  0. 0879 0 . 1 326  0 . 78 0 . 76  3.1 3 .2  <0 . 0 5 0 .06  CWHb D I F F U S E S F DIRECT SF  60 . 5 94 . 2  19180.0 12179.0  0 . 0623 0 . 0821  0 . 89 0 . 51  1 .7 3 .7  <0 .01 0 . 11  Both  D I F F U S E SF DIRECT SF  77 . 4 75 . 4  52.9 1492B.8  0 . 0674 0 . 0872  0 . 82 0 . 47  2 .2 3 .7  <0 .01 <0 . 0 5  CWHa D I F F U S E SF DIRECT SF  9560 . 0 2994 . 0  879.8 -1252.3  0 . 0881 0 . 1 166  0 . 68 0 . 70  174 . 8 1 68 . 3  0 .09 0 .08  CWHb D I F F U S E SF DIRECT SF  5572 . 0  383.5  0. 0689  0 . 79  56 . 3  <0 . 0 5 0 .90  Both  1854 . 3 3 2 1 1 5 2 . 0 3133 . 5 4 7 6 2 1 6 . 0  0. 0372 0. 0917  0 . 57 0 . 53  131 . 9 1 38 .4  <0 .01 <0 . 0 5  D I F F U S E SF DIRECT SF  y•X  p  CWHa D I F F U S E S F DIRECT SF  787 . 9 29 . 6  101.0 -9.4  0. 0906 - 0 . 0209  0 . 84 0 . 76  12 . 0 14 . 3  <0 . 0 5 <0 . 0 5  CWHb D I F F U S E S F DIRECT SF  273 . 6 431 . 9  50096.0 B475943  0. 0759 0. 0916  0 . 90 0 . 53  7 .3 16 . 1  <0 .01 0 .10  Both  D I F F U S E SF DIRECT SF  299 . 9 356 . 6  501 . 3 71767.0  0 . 0718 0. 0972  0 . 85 0 . 51  9.1 16 . 5  <0 . 0 1 <0 . 0 5  CWHa D I F F U S E S F DIRECT SF  3183 .0 279 . 9  54.7 2468.1  0 . 0922 0 . 0757  0 . 61 0 . 28  17 . 1 23 . 0  0 .12 0 .36  CWHb D I F F U S E S F DIRECT SF  581 .8 322 . 2  100.2 70208.0  0 . 0671 0 . 0501  0 . 93 0 . 33  6 .0 18 . 0  <0 .01 0 .23  941 . 1 223 . 7  71.2 20414.0  0 . 0730 0 . 0281  0 . 67 0 . 24  12 . 4 18 . 9  <0 .01 0 .13  Both  DIFFUSE SF DIRECT SF  162  2  Appendix 22. R e l a t i o n s h i p s o f s a l a l d e n s i t y ( # « m ~ ) a n d 95% confidence i n t e r v a l s t o transmission of global, direct, d i f f u s e , a n d d i f f u s e PPFD s o l a r r a d i a t i o n components. Dotted lines are relationships i n i n d i v i d u a l subzones. CWHa ( • ) , CWHb (BB) .  163  PROPORTION GLOBAL RADIATION TRANSMITTED  PROPORTION DIRECT RADIATION TRANSMrTTED  Appendix 23. R e l a t i o n s h i p s o f p e r c e n t c o v e r o f s a l a l and 95% confidence i n t e r v a l s t o transmission of global, direct, d i f f u s e , a n d d i f f u s e PPFD s o l a r r a d i a t i o n components. Dotted l i n e s are relationships i n i n d i v i d u a l subzones. CWHa ( • ) , CWHb (ffi).  164  A p p e n d i x 2 4 . R e l a t i o n s h i p s o f s a l a l D E N S I T Y , CAGBIOM, F O L B I O M , T O T B I O M , a n d P C T C O V E R t o R e i n e k e ' s SDI d e r i v e d from p l o t t r e e measurements (SDI) and p r i s m samples ( B A F S D I ) . D o t t e d l i n e s a r e s e p a r a t e r e l a t i o n s h i p s i n CWHa a n d CWHb variants. 3  

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