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The effect of some environmental factors on the morphological characteristics of western hemlock seedlings… Soos, Joseph 1961

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THE E F F E C T OF SOME ENVIRONMENTAL FACTORS ON THE MORPHOLOGICAL CHARACTERISTICS OF WESTERN HEMLOCK (Tsuga h e t e r o p h y l l a  (.Rafo)  SEEDLINGS  ISargo)  by Dipl«  JOSEPH SOOS F o r . Engo U n i v e r s i t y o f S o p r o n H u n g a r y 1953.  A THESIS SUBMITTED I N PARTIAL FULFILMENT OF THE REQUIREMENTS FOR DEGREE OF MASTER I N FORESTRY  in  the Faculty of Forestry  We a c c e p t t h i s t h e s i s a s c o n f o r m i n g t o t h e required standard  THE UNIVERSITY OF B R I T I S H COLUMBIA A p r i l , 1961  In the  presenting  requirements  of  British  it  freely  agree for  that  an  advanced  for  available  I  copying  gain  shall  by or  his  in  p a r t i a l  degree  of  fbres//*/  the  shall  reference  and  study.  I  extensive  may  be  allowed  The U n i v e r s i t y o f B r i t i s h Vancouver $, Canada.  of  copying  granted  this  without  by  of  the  It thesis  is  Columbia,  make  further this  Head  of  thesis my  understood  for  my w r i t t e n  of  University  Library  publication be  at  the  representatives.  not  fulfilment  that  for  purposes  or  agree  for  permission  that  Department  thesis  Columbia,  scholarly  Department  this  f i n a n c i a l  permission.  ABSTRACT, The objective of the study reported i n t h i s t h e s i s i s to assess the influence of l i g h t and s i t e quality on the morphology of western hemlock Tsuga heterophylla (Raf<>) Sarg, seedlings  0  The importance of western hemlock to the present future economy of B r i t i s h Columbia i s emphasized v i c a l l i t e r a t u r e on the species i s reviewed,  0  and  The  sil-  The inadequate  treatment given western hemlock i n the technical l i t e r a t u r e i s noted and the point i s made that, as the hemlock f o r e s t s are exploited and new  f o r e s t s regenerated, information on the  s i l v i c a l c h a r a c t e r i s t i c s of the species w i l l be needed,  At  the present time, knowledge of the s i l v i c s of western hemlock i s very l i m i t e d .  This thesis attempts to add to knowledge  i n that f i e l d with p a r t i c u l a r reference to some factors ( l i g h t microsite and s i t e ) which influence the juvenile growth of western hemlock on the University of B r i t i s h Columbia Research Forest, near Haney, B C 0  0  The e f f e c t of l i g h t i n t e n s i t y was  studied through measure  ments of natural regeneration growing on 10 p l o t s , under f u l l l i g h t and under d i f f e r e n t degrees of canopy closure, i n a 77year-old stand.  Measurements included height, diameter and  number of green and dead branches. microsites was  The influence of d i f f e r e n t  studied on three-year-old wild seedlings  on 1 -J- 1 planted seedlings, with p a r t i c u l a r reference  and  to  height, stem diameter at ground l e v e l , number of branches, length of needles, angle of branches, diameter of branches  p  weight  and d e p t h  five-year-old  o f r o o t s , and r o o t / s h o o t  w i l d l i n g s growing  i n d i c e s were s t u d i e d w i t h needle length results  with  v i v e with  increased  light  daysj«  significantly  on f i v e  different  morphological  particular greater  o f n e e d l e s and a n g l e  to levels  of branches of  growth„  associated  with  on d i f f e r e n t  site  maximum  I t was c o n c l u d e d  t o estimate  quality.. needle  that, within a i s of  f a c t o r s , i n super-  on t h e b a s i c f o r m o f w e s t e r n  The p o s s i b i l i t y  studied  two y e a r s , t h e  provenance, t h e i n f l u e n c e of environment  needle length  showed  on p l o t s o f known  involved branch angle,  imposing m o d i f i c a t i o n s  days.  external c h a r a c t e r i s t i c s i n height,  s i g n i f i c a n c e than h e r e d i t a r y  seedlingso  on b r i g h t  T h e s e c h a r a c t e r i s t i c s were t h e n  were m o d i f i e d  sur-  ( m e a s u r e d on  c h a r a c t e r i s t i c s o f hemlock p l a n t e d  and h e i g h t  cannot  i n t e n s i t y was f o u n d  The r e s u l t s show t h a t , a f t e r  These m o d i f i c a t i o n s length,  light  The  increased  of microsites  w i l d l i n g s of l o c a l provenance  indices.  sites,  types  shoots«,  seedlings  of f u l l  site  t o maximum  growth  f o r a 12-hour p e r i o d  studied  internode.  planted  site  year's  The a v e r a g e minimum l i g h t  d i a m e t e r , maximum l e n g t h last  and t h a t  l e s s than t e n p e r cent  Seedlings  on  of last  Four-and  0  known  p a r t i c u l a r reference  and b r a n c h a n g l e  t o b e 117 f o o t - c a n d l e s  the  on p l o t s w i t h  i n d i c a t e t h a t d i a m e t e r and h e i g h t  directly  cloudy  ratio  hemlock  o f u s i n g b r a n c h a n g l e and maximum  site  q u a l i t y i s noted.  X X I  C O N T E N T S Page I  0  IA o II. IIIo  ABSTRACT ACKNOWLEDGEMENTS INTRODUCTION  1  DESCRIPTION OF STUDY AREA  8  Ao  B IV.  University  Research  Forest  8  Climate  9  0  METHODS A  0  Bo  The I n f l u e n c e o f L i g h t I n t e n s i t y on Hemlock S e e d l i n g s T h e E f f e c t o f M i c r o s i t e on t h e E x t e r n a l C h a r a c t e r i s t i c s of Three-Year-Old Seedlings  11  14  a.  Microsite  1  14  b.  Microsite  2  15  c«  Microsite  3  15  d  Microsite  4  Io  Microsite  5  Io  0  e« Co  Vo  o f B r i t i s h Columbia  The I n f l u e n c e Seedlings  o f S i t e on Hemlock  The I n f l u e n c e  of L i g h t  17  RESULTS Ao  I n t e n s i t y on  Hemlock S e e d l i n g s  20  a.  T h e amount o f l i g h t  20  bo  The i n f l u e n c e o f l i g h t i n t e n s i t y on t o t a l h e i g h t o f s e e d l i n g s The i n f l u e n c e o f l i g h t i n t e n s i t y on t h e age t o r e a c h b r e a s t h e i g h t  21  c. d e  0  0  The e f f e c t o f l i g h t diameter growth  21  i n t e n s i t y on  The i n f l u e n c e o f l i g h t on d e n s i t y o f f o l i a g e  22 intensity 23  CONTENTS (Cont*d)  B„  Light i n t e n s i t y and form quotient  g»  Natural pruning and l i g h t i n t e n s i t y  23  The E f f e c t of Microsite on the External C h a r a c t e r i s t i c s of Three-Year-Old Seedlings  24  a  T o t a l height  24  b«  Base diameter  24  c  e  Weight of roots  25  d  0  Depth of roots  25  e  0  Weight of needles  e  VI,  26  f.  Root/shoot r a t i o  2b  g  9  Total number of branches  27  h  0  The average number of branches by internodes  27  i»  Average branch angle of internodes  27  3o  Average length of largest needles by internodes  28  k  a  The average length of branches by internodes  29  1  0  a  0  Page 23  f<»  m C  iv  The average base diameter of branches by internodes  29  Bark thickness  30  The Influence of S i t e on Hemlock Seedlings  30  a,  Height growth  30  b»  Branch angle  30  Co  Length of largest needles  31  DISCUSSION A The Influence of Light Intensity on Hemlock Seedlings 33 B o The E f f e c t of Microsite on the External Characteristics of Three-Year-Old Hemlock Seedlings 35 0  Co  The Influence of S i t e on Hemlock Seedlings  Vo  CONTENTS ( C o n t * d )  Page  VII.  CONCLUSIONS  42  VIII,  TABLES  44  IX,  GRAPHS  64  BIBLIOGRAPHY  85  X  0  XI,  APPENDIX  vi Number  10  T A B L E S Page Occurrence of ground v e g e t a t i o n on the t h r e e m i c r o s i t e s f o r study of the e f f e c t of m i c r o s i t e s on the e x t e r n a l c h a r a c t e r i s t i c s of hemlock s e e d l i n g s 0  44  2 e P h y s i c a l c h a r a c t e r i s t i c s of the f i v e p l o t s f o r study of the e f f e c t seedlingso  3« 4.  of s i t e on hemlock  Depths of s o i l h o r i z o n s and the t r e e - r o o t p e n e t r a t i o n of f i v e p l o t s f o r study of the e f f e c t s of s i t e on hemlock s e e d l i n g s .  46  Occurrence of ground v e g e t a t i o n on the f i v e p l o t s f o r study of the e f f e c t of s i t e on hemlock s e e d l i n g s  47  The i n f l u e n c e of l i g h t i n t e n s i t y on hemlock seedlingso L i g h t i n t e n s i t y data on a b r i g h t day f o r e l e v e n and s i x - y e a r - o l d seedlings,,  48  The i n f l u e n c e of l i g h t i n t e n s i t y on hemlock seedlingso L i g h t i n t e n s i t y data on a cloudy day (complete o v e r c a s t ) f o r eleven and s i x year- old s e e d l i n g s „  49  The i n f l u e n c e of l i g h t i n t e n s i t y on hemlock seedlings« Some e x t e r n a l c h a r a c t e r i s t i c s of e l e v e n - y e a r - o l d hemlock s e e d l i n g s on two p l o t s of v a r i o u s degrees of shade  50  The i n f l u e n c e of l i g h t i n t e n s i t y on hemlock seedlingsa Some e x t e r n a l c h a r a c t e r i s t i c s of e l e v e n - y e a r - o l d hemlock s e e d l i n g s on t h r e e p l o t s of v a r i o u s degrees of shade,  51  The i n f l u e n c e of l i g h t i n t e n s i t y on hemlock seedlingso Some e x t e r n a l c h a r a c t e r i s t i c s of s i x - y e a r - o l d hemlock s e e d l i n g s on two p l o t s of v a r i o u s degrees of shade  52  The i n f l u e n c e of l i g h t i n t e n s i t y on hemlock seedlingso Some e x t e r n a l c h a r a c t e r i s t i c s of s i x - y e a r - o l d hemlock s e e d l i n g s on t h r e e p l o t s of v a r i o u s degrees of shade  53  Some e x t e r n a l c h a r a c t e r i s t i c s of t h r e e - y e a r old hemlock s e e d l i n g s on M i c r o s i t e 1  54  Some e x t e r n a l c h a r a c t e r i s t i c s of t h r e e - y e a r old hemlock s e e d l i n g s on M i c r o s i t e 2.  55  0  5o  b  0  7©  0  bo  9o  0  10e  0  11  0  3  12o  45  vii TABLES (Canted) Page  Number 13 a  Some e x t e r n a l c h a r a c t e r i s t i c s or' t h r e e - y e a r o l d hemlock s e e d l i n g s on M i c r o s i t e 3o  56  14a  Some e x t e r n a l c h a r a c t e r i s t i c s of t h r e e - y e a r o l d hemlock s e e d l i n g s on M i c r o s i t e 4o  57  15o  Some e x t e r n a l c h a r a c t e r i s t i c s of t h r e e - y e a r o l d hemlock s e e d l i n g s on M i c r o s i t e 5,  58  C h a r a c t e r i s t i c s of branches of t h r e e - y e a r o l d hemlock s e e d l i n g s i n r e l a t i o n t o i n t e r node and m i c r o s i t e {.Averages of 10 s e e d l i n g s per m i c r o s i t e ) ,  59  Student*s t - t e s t v a l u e s f o r the comparison of the mean t o t a l height of hemlock s e e d l i n g s on f i v e m i c r o s i t e s ,  60  Student*s t - t e s t v a l u e s f o r the comparison o f the mean base diameter of hemlock s e e d l i n g s on f i v e m i c r o s i t e s ,  60  Student*s t - t e s t v a l u e s f o r the comparison of mean branch angle of t h i r d i n t e r n o d e o f hemlock s e e d l i n g s on f i v e m i c r o s i t e s .  61  Student*s t - t e s t v a l u e s f o r t h e comparison o f mean l e n g t h of l a r g e s t needles of t h i r d i n t e r node o f hemlock s e e d l i n g s on f i v e m i c r o s i t e s ,  61  Student's t - t e s t v a l u e s f o r t h e comparison of mean oven-dry weight of needles ( i n gramsj o f hemlock s e e d l i n g s on f i v e m i c r o s i t e s ,  62  The i n f l u e n c e of s i t e on hemlock s e e d l i n g s . The mean l e n g t h of l a r g e s t n e e d l e s , branch angle of l a s t yearns i n t e r n o d e , and height growth of hemlock s e e d l i n g s  63  lb.  17«  18o  19o  20  o  21,  22©  e  23o  The i n f l u e n c e of s i t e on hemlock s e e d l i n g s . Student's t - t e s t v a l u e s f o r comparison o f mean branch angle of l a s t y e a r s i n t e r n o d e of hemlock s e e d l i n g s on f i v e p l o t s , 8  24o  The i n f l u e n c e of s i t e on hemlock s e e d l i n g s , Student*s t ~ t e s t v a l u e s f o r t h e comparison of mean l e n g t h of l a r g e s t needles of l a s t yearns internode of hemlock s e e d l i n g s on five plotSo  64  65  viii TABLES l C o n t * d ) Number 25.  Page The i n f l u e n c e of s i t e on hemlock s e e d l i n g s F i d u c i a l l i m i t s at 0 01 l e v e l calculated f o r t h e mean branch angles (.in degrees) o f hemlock s e e d l i n g s on f i v e p l o t s <,  0  0  26o  66  The i n f l u e n c e of s i t e on hemlock s e e d l i n g s F i d u c i a l l i m i t s at 0 01 l e v e l c a l c u l a t e d f o r t h e mean of l a r g e s t needles I i n m i l l i meters) of hemlock s e e d l i n g s on f i v e p l o t s  6b  E s t i m a t i o n o f s i t e index of m i c r o s i t e s by u s i n g needle l e n g t h and branch angle of hemlock s e e d l i n g s  67  0  o  0  27o  0  ix  0  ILLUSTRATIONS Figure l  a  2o 3e 4o  Page Location or' p l o t s established f o r various studies« Appendix Cumulative l i g h t i n t e n s i t y of various degrees of shade f o r 12-hour period of bright day. 68 Cumulative l i g h t i n t e n s i t y of various degrees of shade f o r 12-hour period of bright day©  °9  Cumulative l i g h t i n t e n s i t y of various degrees of shade f o r 12-hour period of cloudy day  70  Cumulative l i g h t i n t e n s i t y of various degrees of shade f o r 12-hour period of cloudy day  71  0  5o  0  bo  Average cumulative length of internodes of 11-yearold seedlings on plots of various degrees of shade  72  Average cumulative length of internodes of s i x year- old seedlings on plots of various degrees, of shadeo  73  Average cumulative diameter of internodes of eleven-year-old seedlings on plots of various degrees of shade  74  Average cumulative diameter of internodes of s i x year- old seedlings on plots of various degrees of shadeo  75  Average cumulative number of green branches of eleven-year-old seedlings on p l o t s of various degrees of shade,  76  9  7o  b  0  0  9o  10o  lie  12© 13a 14• 15 a 16o  Average cumulative number of green branches of six-year-old seedlings on p l o t s of various degrees of shade a  77  Average t o t a l height of three-year-old seedlings on various micrositeso  78  Average base diameter of three-year-old seedlings on various micrositeso  78  Average oven-dry weight of roots of three-year-old seedlings on various micrositeso  79  Average depth of roots of three-year-old seedlings on various micrositeso  79  Average oven-dry weight of needles of three-yearold seedlings on various microsites,  80  ILLUSTRATIONS (Ccnt^d) Figure 17o  Page Average number of green branches of t h r e e - y e a r - o l d s e e d l i n g s on v a r i o u s m i c r o s i t e s  80  The average branch angle various micrositeso  81  0  18  s  of t h r e e i n t e r n o d e s of t h r e e  on  19o  Average l e n g t h of l a r g e s t needles nodes on v a r i o u s m i c r o s i t e s ,  inter-  20o  Average l e n g t h of branches of t h r e e i n t e r n o d e s various micrositeso  on  21B  The average base diameter of branches of t h r e e nodes on v a r i o u s m i c r o s i t e s ,  inter82  22o  R e g r e s s i o n l i n e based on branch angle and s i t e f o r study of the i n f l u e n c e of s i t e on hemlock seedlings<y3  23o  R e g r e s s i o n l i n e based on l e n g t h of needles and f o r study of the i n f l u e n c e of s i t e on hemlock seedlings,  °"1 ^2  site 84  XX o  ACKNOWLEDGE*® NTS  The author wishes t o express h i s thanks t o ;  Dr„  PoGo Haddock,associate p r o f e s s o r w h o d i r e c t e d and a d v i s e d s  the i n v e s t i g a t i o n , , Mr,  Jo W a l t e r s ,  r e s e a r c h f o r e s t e r , who a d v i s e d and encouraged the author throughout the study,  Dr, JoG.H. Smith, a s s o c i a t e p r o f e s s o r , f o r h i s and Mr  D  L  0  suggestions  criticisms  Orloczy f o r h i s assistance i n i d e n t i f y i n g the species of minor v e g e t a t i o n .  I N T R O D U C T I O N  72 p e r c e n t o f t h e l a n d  Almost is  forested.  heterophylla  I n terms  immature,  7o2  million  8,684,876  acres.  a n d 0,3 m i l l i o n  mature and overmature  o f hemlock i n t h e contemporary  the Province,  billion  cubic feet) i nB r i t i s h  double that the  Of  acres,  and f u t u r e  economy  155«6  Columbia, i s second only t o  billion  c u b i c f e e t ) , and i s a l m o s t  o f D o u g l a s f i r (30,6 b i l l i o n  cubic f e e t )  0  On  c o a s t t h e volume o f w e s t e r n hemlock i s t r e b l e t h a t o f  Douglas bia,  f i r (Continuous Forest  1957)o  western  These  industry  arity  the  Colum-  importance  hemlock.  of theprovince.  pronounced  Inventory of B r i t i s h  d a t a s e r v e t o emphasize  Hemlock i s t h e p r i n c i p a l  o f hemlock i s t h a t  s p e c i e s used i n the pulp  I t has a l i g h t e r  g r a i n than Douglas  uses than Douglas  fir.  c o l o u r and l e s s  The c a u s e  o f unpopul-  i t i s less desirable  for certain  f i r o r more d i f f i c u l t  t o p r o c e s s , The  m o i s t u r e c o n t e n t o f hemlock i s h i g h e r t h a n D o u g l a s it  species,  stands, respectively.  T h e volume o f w e s t e r n hemlock  o f t h e s p r u c e s (92,8  that  of  Tsuga  l a r g e a r e a s u p p o r t i n g mature s t a n d s , i n d i c a t e s t h e i m -  portance of  of stands covering  a r e a , 1,1 m i l l i o n ,  support The  o f a r e a , western hemlock,  Columbia  ( R a f ) S a r g , i s t h e f o u r t h most i m p o r t a n t  b e i n g a component this  area of B r i t i s h  takes longer t o dryj  f i r and  s a n d i n g i s l e s s s a t i s f a c t o r y and  l e s s face veneer i s produced.  The advantages  o f hemlock  - 2 i  v  t o t h e lumber  and  l i g h t e r than f i r , more a t t r a c t i v e  plywood  i n d u s t r i e s are that  thus i t costs  and  less to ship  i t s painting  0  the product i s The  characteristics  grain i s  are  better  t h a n f i r (Wellwood 1957)° Western  h e m l o c k grows a l o n g t h e P a c i f i c  Alaska t o Northern C a l i f o r n i a . narrow,  scattered  pattern  Coast,  from  I t a l s o ranges i n l a n d  in a  along the United States-Canadian  b o r d e r , and t h e n s p r e a d s out t h r o u g h N o r t h e a s t e r n W a s h i n g t o n , Northern Idaho,  N o r t h w e s t e r n Montana and  Southeastern B r i t i s h  Columbia.  Average  annual p r e c i p i t a t i o n  region v a r i e s from  38  throughout  i n the c o a s t a l  i n c h e s t o o v e r 100  hemlock  i n c h e s (.Berntsen  195«Jo  According t o Halliday  and Brown (1943.J > t h e b e s t s t a n d s o f  w e s t e r n hemlock a r e on t h e Queen C h a r l o t t e thern part interrupted  of Vancouver by  Island„  the I n t e r i o r Dry  The  distribution  British  b) area  nor-  Selkirk,  on  Purcell  Mountains„  Krajina in  the  eastward i s  B e l t , but i t reappears  t h e r a i n y westward s l o p e s of t h e Columbia, and R o c k y  I s l a n d s and  (1959) d i s t i n g u i s h e d two w e s t e r n h e m l o c k z o n e s  Columbia:  I n t e r i o r Western  a)  Hemlock Z o n e ,  o f t h e I n t e r i o r Wet  F l o w e r i n g and  fruiting  c r i b e d by t h e F o r e s t  C o a s t a l Western  Belt,  habits  Hemlock Zone  and  Rowe (1959) c a l l e d  the  t h e "Columbia  Forest  Region".  o f w e s t e r n h e m l o c k were d e s -  S e r v i c e U.S.  Department  of A g r i c u l t u r e  - 3 (1948), and by B e r n t s e n (1958), are monoecious The  pollen  and a r e on d i f f e r e n t  s h e d d i n g b e g i n s about  than i n Washington The dency  The m a l e and  ( D a l l i m o r e and J a c k s o n 1 9 4 8 )  average weight  o f one  1,23  grams p e r t h o u s a n d  Western  thousand  s e e d s was  1,7  c r o p a t an a v e r a g e i n t e r v a l white pine type  10 m i l l i o n  Garman  An  Douglas f i r , Haig  and  a bum-  86 pounds o f  (1951) f o u n d t h a t more t h a n near A l b e r n i ,  Van-  hemlock seed c r o p  at the University  An e s t i m a t e d w e i g h t  Re-  o f 113  p r o d u c e d , p e r a c r e , i n a young  hemlock,  seed  years i n the western  which produced  o b s e r v e d by t h e a u t h o r i n 1959  o f hemlock s e e d was  weight  Island,  p r o d u c i n g a heavy  e x c e p t i o n a l l y heavy  s e a r c h F o r e s t n e a r Haney,  Charlotte  (1951) t h e a v e r a g e  s e e d s p e r a c r e were r e l e a s e d  couver I s l a n d ,  ten-  (1952) r e -  Godman (1953) r e p o r t e d  p e r c r o p i n A l a s k a d u r i n g 1951, hemlock s e e d p e r a c r e ,  Wiksten  grams on V a n c o u v e r  o f 2,1  ( H a i g 1941)«  e  s e e d s i n t h e Queen  hemlock i s v e r y p r o l i f i c ,  tree.  i n Alaska  t h o u s a n d s e e d s shows a  I s l a n d s , w h i l e a c c o r d i n g t o Garman o f one  flowers  o f t h e same  one month l a t e r  t o decrease from south t o n o r t h ,  ported  was  branches  female  pounds  stand of  cedar,  (1941) f o u n d t h a t , i n I d a h o , w e s t e r n hemlock r e -  l e a s e s i t s seeds c o n t i n u o u s l y i n g summer,  Allen  f r o m midsummer t o t h e  (1958) f o u n d t h a t S e p t e m b e r 15  follow-  represents  approximately the date at the U n i v e r s i t y F o r e s t i n C o a s t a l B.C.  when a l m o s t a l l s e e d s h a v e m a t u r e d  g e r m i n a t e and t o s t o r e  well.  sufficiently  to  - 4 According t o Siggin*s data western  hemlock seed  slowest  of i t s associateso  Wiksten  was 2.6 f e e t / s e c o n d , w h i c h i s t h e  (1953) r e p o r t e d t h a t t h e number o f s e e d  per acre decreased sourceo  (1933) t h e r a t e o f f a l l o f  w i t h i n c r e a s i n g d i s t a n c e from  He a l s o f o u n d  that lightest  seed  caught  t h e seed  t r a v e l l e d the  s h o r t e s t d i s t a n c e s , and t h e h e a v i e s t , f l e w a b o u t 5 c h a i n s * Sutton  (1954) s t u d i e d t h e g e r m i n a t i o n  and s u r v i v a l  hemlock s e e d l i n g s on r o t t e n ' w o o d , l i t t e r , soil found  and v e r m i c u l i t e , w i t h no s i g n i f i c a n t  t o seedbed Allen  or pH  tification Bientjes of seed  s p e c i a l a t t e n t i o n t o pH.  t h a t seed  collected  reduces  of western  on S e p t e m b e r 15,  He c o n c l u d e d  rate of that  stra-  t h e v a r i a t i o n i n r a t e o f germination,,  (1954) i n d i c a t e d  highest germination  t h a t r a t e and amount o f g e r m i n a t i o n  hemlock a r e h i g h l y v a r i a b l e used,  percent„  (1958) a r e i n p a r t i a l  conflict  20 d e g r e e s The r e s u l t s with these  were c o n s i d e r e d n e c e s s a r y  f o r germination  C„ i n c u b a t i o n t e m p e r a t u r e  8  Of t h e t h r e e  C. p r o d u c e d  the  r e p o r t e d by  Ching  o f A l l e n and  B i e n t j e s i n d i c a t i n g t h a t n o p r e s o a k i n g and n o  ation  attributable  c a p a c i t y , but a slower  later collections.  i n c u b a t i o n temperatures  degrees  He  0  (1958) f o u n d  than  humus, m i n e r a l  d i f f e r e n c e s i n germination  showed a h i g h g e r m i n a t i v e germination  of western  stratification  t e s t s , b u t t h a t 20  a l s o gave t h e b e s t  germin-  results. One o f t h e most i m p o r t a n t  mination  factors f o r successful ger-  i n t h e f o r e s t , and i n t h e n u r s e r y , i s t h e d e p t h o f  - 5 cover*  I n one s t u d y  exceeded  i t was f o u n d  that, i f t h e depth  one q u a r t e r o f a n i n c h ,  the seeds germinated  t h e y d i d n o t a p p e a r above t h e g r o u n d  (Hoffmann  Olson,  Stearns  and N i e n s t a e d t  germination  process  o f e a s t e r n hemlock.  ponds t o t h a t o f w e s t e r n  of cover  1918)  but e  (1959) d e s c r i b e d t h e This process corres-  hemlock, as observed  b y t h e author,.  The  r o o t grows a t a r a t e o f two o r t h r e e m i l l i m e t e r s a d a y ,  for  the f i r s t  of  s e v e r a l days,  a f t e r which t h e stem p o r t i o n  the hypocotyl a l s o begins  t o grow.  Normally,  a pause i n development a f t e r t h e c o t y l e d o n s is  there i s  o p e n , and i t  a t t h i s p o i n t t h a t g e r m i n a t i o n might a r b i t r a r i l y  s i d e r e d t o end.  During the early  stages  be  con-  o f growth, the  l e a v e s a r e formed i n t h r e e s around t h e stem, a t about t h e same l e v e l ,  so that they  appear  E a r l y b u d s on v i g o r o u s in  one g r o w i n g s e a s o n  whorled.  shoots  occasionally  (lammas g r o w t h ) ,  1961),  Mortality  of seedlings begins  diately  a f t e r germination,  e a r l y m o r t a l i t y was c a u s e d  Haig  flush  twice  ( W a l t e r s and S o o s , i n the forest  imme-  (1941) p o i n t e d o u t t h a t  p r i n c i p a l l y by b i o t i c  agents  ( i n s e c t s , b i r d s and f u n g i ) and o c c u r r e d when s e e d l i n g s were still  s u c c u l e n t and t e n d e r .  Later losses, beginning i n late  J u n e and e a r l y J u l y , were p r i n c i p a l l y drought.  He f o u n d  that i n f u l l  due t o i n s o l a t i o n a n d  sunlight  the surface of d u f f  becomes much h o t t e r t h a n t h e s u r f a c e o f m i n e r a l s o i l . caused  lesions  producing  a t t h e ground l i n e  t h e death  o f many,  hot weather t h e temperature  This  on t h e s u c c u l e n t s e e d l i n g s ,  Isaac  (1938), f o u n d  that i n  o f unshaded f i r e - b l a c k e n e d s u r -  f a c e s was c o n s i d e r a b l y h i g h e r  (seven t o e i g h t e e n degrees  F)  than that  of u n d i s t u r b e d  occur  a t an  soils  He  a  temperature  a i r temperature  reported  o f 125  o f 85  that i n -  degrees  the deepest rooted rooted  species  d r o u g h t was  and  w e s t e r n hemlock as  He  0  heaviest  one  of the  of shallow-  found t h a t the m o r t a l i t y caused  under f u l l  Drought l o s s e s at the  F.  degrees F ) ,  (1941) c l a s s i f i e d w e s t e r n w h i t e p i n e a s one  Haig  est  -  mineral  j u r y began a t a s u r f a c e s o i l ( w h i c h may  6  full-sun  shade f o r b o t h and  species  by  0  p a r t - s h a d e s t a t i o n s would A  have b e e n r e l a t i v e l y killed  higher  i f i n s o l a t i o n had  l a r g e numbers o f r e d  Godman and  Gregory  c e d a r and  1951  1952,  same s p e c i e s , was  Observations (195b)o  He  133  of days,  Griffith s  re-  E  155  d a y s and  153  d a y s i n 1954  August  1955*  and  B,C©  o f l e a d e r g r o w t h were r e p o r t e d by  found that  terminated  Buckland  l e a d e r growth s t a r t e d e a r l y i n 25»  at Cowichan Lake,  May  Godman and  Gre-  (1955) r e p o r t e d t h a t l e a d e r g r o w t h i n A l a s k a b e g a n a t  gory the  d a y s and  respectively, i n Alaska,  r e s p e c t i v e l y , a t Haney,  and  127  duration  (1959) i n d i c a t e d t h a t t h e d u r a t i o n o f r a d i a l g r o w t h f o r  sults the  and  previously  hemlock,  (1955) f o u n d t h a t t h e  r a d i a l g r o w t h o f w e s t e r n h e m l o c k was in  not  end  Walters  of May  and  terminated  (I960) p o i n t e d  out  on September 17,  that the  point  in  1952,  of t e r m i n a t i o n  a n n u a l g r o w t h i n w e s t e r n h e m l o c k i s marked u s u a l l y by extra  l o n g b r a n c h immediately below the Wellner  (194b)  well i n height  growth a f t e r a c l e a n i n g i n a mixed hemlock s e e d l i n g s had  height  check p l o t s , but  as t h o s e  on  the  an  a n n u a l node,  observed t h a t western hemlock  Moderately cleaned  of  the  responded stand.  same a v e r a g e  heavy c l e a n i n g  re-  s u i t e d i n b5 p e r cent b e t t e r average height„ Griffith f o r a thinned  (1959) found t h a t the average height  growth  hemlock stand was s l i g h t l y l e s s than the average  height  growth of t r e e s on the c o n t r o l a r e a .  served  t h a t the thinned  However, he ob-  t r e e s grew i n b a s a l area more than  t w i c e as f a s t as the c o n t r o l t r e e s , W a l t e r s , Soos, and Haddock (i960) d e s c r i b e d hemlock p l u s t r e e s a t Haney  0  western  The angle of branches i s an  important f a c t o r f o r p l u s - t r e e s e l e c t i o n r e g a r d l e s s species  (Toda 1957, Z o b e l I960),  of t r e e  Trees with good b o l e  form  are f r e q u e n t l y r e j e c t e d i f the angle of branches i s much l e s s than 90 degrees.  On t h e b a s i s of the study  reported  i n t h i s t h e s i s , i t appears t h a t t h e s i t e or even the micros i t e , has a s i g n i f i c a n t e f f e c t on some e x t e r n a l istics  of hemlock  The  character-  seedlings.  s i t e s and m i c r o s i t e s s u p p o r t i n g  l i n g s studied, are described  t h e hemlock seed-  i n d e t a i l , and the e f f e c t of  l i g h t i n t e n s i t y and s i t e q u a l i t y on some e x t e r n a l characteristics study.  of s e e d l i n g s , i s d i s c u s s e d  w i t h i n t h e scope of the  The r e l a t i o n s h i p between s i t e q u a l i t y and angle of  b r a n c h i n g r e c e i v e s p a r t i c u l a r emphasis.  D E S C R I P T I O N  OF  S T U D Y  A R E A  UNIVERSITY OF BRITISH COLUMBIA RESEARCH FOREST The U n i v e r s i t y sists  of B r i t i s h Columbia  o f 9*774 a c r e s s i t u a t e d  m i l e s n o r t h o f Haney and B C o  i n the F r a s e r V a l l e y ,  t h i r t y - f i v e miles east  E l e v a t i o n s range from sea l e v e l  0  Research Forest  of  four Vancouver,  t o 2,600 f e e t  area i s s i t u a t e d  (U B C  F o r e s t Committee 1959)o  The  Southern C o a s t a l S e c t i o n  (C^) o f R o w e s (1959) F o r e s t  jina s  m a i n a s s o c i a t i o n c o m p r i s e s t h e two  Tsuga fir,  red cedar, Thuja p l i c a t a heterophylla Pseudotsuga  Chamaecyparis  Donn, and  western  ( M i r b ) F r a n c o and  (Douglo) Forbo  Spach,  w e s t e r n p o r t i o n o f t h e F o r e s t was  and  regenerated n a t u r a l l y  and now  hemlock, Douglas wes-  silver f i r ,  and S i t k a  spruce,  (Bongo) C a r r .  The  was  wes-  scattered  Pacific  t o a mixed  t e r n h e m l o c k and w e s t e r n r e d c e d a r s t a n d o tion  The  yellow (or Alaska) cedar,  n o o t k a t e n s i s (D« Don)  Picea sitchensis  1868  Region  c o a s t a l dominants,  t e r n white pine Pinus m o n t i c o l a Douglo, Abies amabilis  o  Hemlock Zone o f K r a -  (Raf.) Sargo, i n combination with  menziesii  0  the  (1959) B i o c l i m a t i c Z o n e s i n B r i t i s h Columbia©  8  tern  within  e  e  o f Canada and i n t h e C o a s t a l W e s t e r n  con-  l o g g e d and b u r n e d s u p p o r t s mixed  l o c k , and w e s t e r n grees of s t o c k i n g .  s w e p t by f i r e i n Douglas The  eastern por-  b e t w e e n t h e y e a r s 1920  stands of Douglas  f i r , wes-  and  1931  f i r , western  hem-  r e d c e d a r o f v a r i o u s c o m p o s i t i o n s and Scattered  old-growth f o r e s t  numerous i n t h e f o r e s t where t h e f i r e  de-  types are  d i d not touch or l i g h t l y  - 9 burned the o r i g i n a l s t a n d .  The t o p o g r a p h i c c o n d i t i o n s on the  whole area are g e n e r a l l y rugged, w i t h numerous rock  out-  er oppings. Most s o i l s o r i g i n a t e d from g l a c i a l t i l l . e x c e e d i n g l y rocky and  The s o i l i s  of a sandy loam t e x t u r e , w i t h a v a r y i n g  depth, from a few i n c h e s t o t h r e e or more f e e t  ( G r i f f i t h 1960JL  C L I M A T E A weather s t a t i o n has been maintained an e l e v a t i o n of 550 f e e t , s i n c e 1946.  The  a t the F o r e s t a t climate i s i n -  f l u e n c e d by the P a c i f i c Ocean and by the Coast Mountains. The main f e a t u r e s of the c l i m a t e are the m i l d , wet  winters  and the comparatively warm, dry summers. P r e c i p i t a t i o n on the F o r e s t f o l l o w s , g e n e r a l l y , the same p a t t e r n as the southern c o a s t a l r e g i o n of B r i t i s h umbia.  Col-  The six-month p e r i o d from October t o March, i n c l u s i v e ,  i s very moist, having an average i n c h e s per month.  p r e c i p i t a t i o n of  11*24  But the growing season i s r e l a t i v e l y  with an average monthly r a i n f a l l of 3.98 p r e c i p i t a t i o n occurs mostly as r a i n ; s n o w f a l l f o r the p e r i o d 194b  - 1957  inches.  dry  Winter  however, the i s 58 i n c h e s .  average Of the  t o t a l p r e c i p i t a t i o n , 40.76 per cent occurs d u r i n g the w i n t e r months, 20.72 per cent d u r i n g the s p r i n g , 11.64 the summer and drought  26.88 per cent i n the autumn.  s i n c e 1946  per cent i n  The l o n g e s t  o c c u r r e d i n the summer of I960, and  lasted  f o r f i v e weeks. Temperatures are r e l a t i v e l y m i l d i n w i n t e r and summer.  The average mean temperature  cool i n  f o r the w i n t e r months  - 10 is F  0  The  34.66 d e g r e e s F„ and f o r t h e summer months, 59<>66 d e g r e e s The a v e r a g e a n n u a l mean t e m p e r a t u r e i s 47 d e g r e e s F„ l o w e s t r e c o r d e d t e m p e r a t u r e was m i n u s 5 d e g r e e s F , w h i c h G  occurred  i n J a n u a r y 1950  o f 100 d e g r e e s F The shortest  c  and t h e l o n g e s t  The h i g h e s t  o c c u r r e d i n August  a v e r a g e number period  0  occurred  of f r o s t - f r e e i n 1956  recorded temperature  I96O0 d a y s i s 193 d a y s  w i t h 156 f r o s t - f r e e  i n 1947* w i t h 225 d a y s  0  a  The  days  - 11 THE  INFLUENCE OF LIGHT INTENSITY ON HEMLOCK SEEDLINGS„  A r e p r e s e n t a t i v e area i n compartment 3 B (.Figure 1 Append i x ) was chosen t o study the e f f e c t hemlock s e e d l i n g s reproduced western  of l i g h t i n t e n s i t y on  The e n t i r e area was burned i n Ibbb and 1  0  n a t u r a l l y t o Douglas f i r , western  red cedar  0  f ollowed by western  hemlock and  Douglas f i r was e s t a b l i s h e d f i r s t , hemlock and western  r e d cedar, as e v i -  denced by the younger ages of t h e l a t t e r two s p e c i e s 1960)  The average  e  age of t h e stand, f o r Douglas f i r , a t  the time t h e study was made was 77 y e a r s the vicinity  (Griffith  The area i s i n  s  of P l o t b ( G r i f f i t h i 9 6 0 ) and has an e l e v a t i o n  of 1,100 f e e t on a southwesterly a s p e c t , w i t h a s l o p e of n o t more than f i v e p e r cento g l a s f i r and western pectively  The estimated s i t e i n d i c e s of Dou-  hemlock were 140 f e e t and 115 f e e t , r e s -  0  The f o l l o w i n g c h a r a c t e r i s t i c s were d e s c r i b e d by G r i f f i t h (1960) W  C  M  o  The average  depths of s o i l h o r i z o n s A % m  ®B and n  were f o u r , e i g h t e e n and e i g h t e e n i n c h e s , r e s p e c t i v e l y  e  The depth of h o r i z o n s was found t o be f a i r l y uniform and t h e s o i l was c l a s s i f i e d as b e l o n g i n g t o the i m p e r f e c t l y d r a i n e d groups o f Brown P o d s o l i c s o i l s c o l o r with sandy loam t e x t u r e  The "A2 h o r i z o n was grey i n K  e  w 0  B  n  horizons varied  y e l l o w i s h brown t o dark brown i n c o l o r , with m o t t l i n g a t the lower l e v e l s  0  from  conspicuous  The t e x t u r e was c o a r s e , sandy  loam, o f t e n c o n t a i n i n g v a r i e d amounts of f i n e gravel« M  C  W  h o r i z o n s were of g l a c i a l t i l l  grey i n color«  The  f o r m a t i o n , l i g h t t o dark  T e x t u r a l l y they were c o a r s e , sandy loam, and  o f t e n contained v a r i e d amounts of f i n e gravelo  - 12 77=year o l d  The by  -  s t a n d n e a r L o o n L a k e Road was  p a r t i a l l o g g i n g o f D o u g l a s f i r and  vigorous natural  the  light  the  stand  of western  roadside extending i n t o  p e n e t r a t e d from the  canopy of  road c o n s t r u c t i o n  reproduction p r i n c i p a l l y  took p l a c e along the  r o a d and  also  the  stand  were  t h e r e , under d i f f e r e n t d e g r e e s of canopy c l o s u r e The  plot  s u p p o r t i n g no  a control  plot.  ( P l o t 1,  cated  where t h e  h e m l o c k s e e d l i n g s was  From t h i s p l o t , 2,  Plot  closure  of  2,  Plot  3,  canopy d e c r e a s e d  Plot  1 and  were  7084 m e t e r s , 7<>23 m e t e r s and  Plot  Plot  Hemlock s e e d l i n g s on  Plot  and  5 was  established  3 and  h a r v e s t e d i n 1953 year- old  sunlight.  The  years in  old  were c h o s e n f o r f u r t h e r vance r e g e n e r a t i o n ,  study.  and  grew  repro-  p r e v i o u s s t a n d had  T h e s e were, o f  been  Eleven-  d e c a y i n g wood i n t h i s  remaining a f t e r the  4*  Plot  compartment  because of m i s t l e t o e i n f e c t i o n .  s e e d l i n g s g r o w i n g on  road,  respectively.  n e a r L o o n L a k e Camp, where a v i g o r o u s n a t u r a l  d u c t i o n grew i n f u l l  as  d i s t a n c e s between  Plot  located  1)«  were l o -  toward the  b«,47 m e t e r s , were 11  broken  established  The  0  3*  Plot  these p l o t s  o n l y i n d e c a y i n g wood. 3 A,  2 and  4)  Plot  where  (Figure  four other plots  and  A  0  hemlock  from the  Nine f o u r - m i l a c r e p l o t s  0  disturbed  area,  course,  a r e a was  ad-  logged  in  1953o The was  the  location same as  Plot  4  Plot  I I and  (Figure  of P l o t  described 1),  Plot  I, Plot for Plot  II, Plot 1,  Plot  D i s t a n c e s between P l o t I I I , and  Plot  I I I and  Plot  I I I and  Plot  2,  3  Plot  I and IV,  Plot were  IV,  and II,  5*47  - 13 meters, 4o47 meters and 3o80 meters, r e s p e c t i v e l y ,  Seedlings  growing on P l o t I , P l o t I I , P l o t I I I and P l o t IV were s i x years o l d and the r o o t s grew mostly i n A M  ra Q  horizon.  Plot V  was l o c a t e d on the same s i d e o f Loon Lake Road, i n compartment 3 C, where P l a n t a t i o n 27 ( F i g u r e 1) was e s t a b l i s h e d , to  study the growth of hemlock s e e d l i n g s .  The hemlock seed-  l i n g s v a r i e d from s i x t o seven y e a r s of age, and grew i n f u l l sunlight.  The p l o t s having 11-year-old  s e e d l i n g s were r e -  corded by a r a b i c numbers, l a r g e r numbers r e p r e s e n t i n g i n creased l i g h t i n t e n s i t y .  The p l o t s having 6-year-old seed-  l i n g s were coded with Roman numbers, the l a r g e r numbers r e presenting increased l i g h t  intensity.  The l i g h t i n t e n s i t y on each p l o t was measured f o r a 12hour p e r i o d , on a cloudy and on a sunny day, w i t h the l i g h t meter 4o5 f e e t above ground l e v e l . the  The l i g h t m e t e r used was  Brosh t y p e , which gave the i n t e n s i t y i n f o o t - c a n d l e s .  S i n c e the o b s e r v a t i o n showed t h a t each p l o t had d i f f e r e n t l i g h t i n t e n s i t i e s , ten s e e d l i n g s were chosen from each p l o t for  further analysis.  The p o s i t i o n o f nodes was determined  on every s e e d l i n g w i t h each i n t e r n o d e r e p r e s e n t i n g one yearns growth.  The base and mid-diameter of each i n t e r n o d e were  measured w i t h a c a l i p e r , o b t a i n i n g a p r e c i s i o n o f 1/10  milli-  meters.  The l e n g t h of each i n t e r n o d e was measured i n m i l l i -  meters.  Numbers of green and dead branches were counted  and recorded f o r every i n t e r n o d e , was examined  A t o t a l o f 100  seedlings  and analysed on t e n f o u r - m i l a c r e p l o t s .  i  - 14 THE E F F E C T OF MICROSITE  ON THE EXTERNAL  OF THREE-YEAR-OLD HEMLOCK The  area t o r the study  three-year-old IC  SEEDLINGS  0  or' t h e i n f l u e n c e o f m i c r o s i t e on  s e e d l i n g s was l o c a t e d i n Compartments IB a n d  i n the southern  growing i n f u l l Seedlings  CHARACTERISTICS  part of the Research  sunlight  Forest,  Seedlings  on f i v e m i c r o s i t e s were s t u d i e d .  on m i c r o s i t e s 1 a n d 4 were p l a n t e d b u t s e e d l i n g s  on m i c r o s i t e s 2, 3, a n d 5 were The  natural.  d e t e r m i n a t i o n o f m i c r o s i t e q u a l i t y based on t h e t o t a l h e i g h t o f s e e d l i n g s , M i c r o s i t e 1,  Microsite from an  1 was l o c a t e d i n Compartment 1 0 , t e n m e t e r s  the road.  old-growth  Until stand  logged  i n 1956, t h e a r e a  c o n s i s t i n g mainly  i n g l o g g i n g t h e s l a s h was p i l e d site  index The  indicative The  of Douglas f i r .  and b u r n e d .  of the Polystichum  evident  forest  Follow-  The e s t i m a t e d  o f t h e p r e v i o u s s t a n d was 160 f e e t  ground v e g e t a t i o n s t i l l  supported  f o r Douglas  on t h e a r e a was  type  ( K r a j i n a 1^59)•  "A" h o r i z o n s were d i s t u r b e d c o n s i d e r a b l y d u r i n g t h e  logging,  "B" h o r i z o n s were 70 c e n t i m e t e r s d e e p , m o i s t , and  possessed  a high  organic m a t e r i a l content.  m a t e r i a l was g l a c i a l The  The p a r e n t  till.  a r e a was f l a t ,  but well-drained.  The h e m l o c k  l i n g s s t u d i e d were p l a n t e d on t h e a r e a i n November 1 + 1  stock of l o c a l  blished of  fir.  t o study  growth.  provenance.  the effect  seed-  1958 a s  The p l a n t a t i o n was e s t a -  o f s p a c i n g on v a r i o u s  aspects  - 15 •/  i  The North  M i c r o s i t e 2. a r e a was l o c a t e d i n Compartment  study  Fork  o f t h e A l o u e t t e R i v e r a n d was p a r t i a l l y  1955 and c l e a r - c u t  i n 1959.  w h i c h were o n l y m o d e r a t e l y timeters© and  M  B2  The t i l l .  d i s t u r b e d b y l o g g i n g , was 12 c e n -  The B " h o r i z o n s were 47 c e n t i m e t e r s t h i c k e  A study  b e f o r e i t was  n  0  h o r i z o n was d e r i v e d f r o m  t h e Blechnum f o r e s t  logged  type  but well-drained  on t h i s m i c r o s i t e o r i g i n a t e d  that the area (Krajina  0  Seedlings  growing  3.  3 was l o c a t e d a d j a c e n t t o M i c r o s i t e 2o  S e e d l i n g s g r o w i n g on t h i s m i c r o s i t e o r i g i n a t e d on wood i n a s t a g e  of well-advanced  7<>5 c e n t i m e t e r s  0  D o u g l a s - f i r l o g s were t a k e n  tent  T h e l o g s were  Two s a m p l e s o f wood f r o m t o the Forest Products  V a n c o u v e r , B.C., where t h e s p e c i f i c  The  decay©  naturally  on t h e s u r f a c e and i n t o t h e wood f o r a d e p t h o f  more t h a n  moisture  1959)  naturally.  Microsite  decayed  glacial  0  a r e a was f l a t  Microsite  water  of minor v e g e t a t i o n i n d i c a t e d  supported  The  "Bj™  n  m a t e r i a l o f t h e "C  probably  logged i n  The t h i c k n e s s o f t h e "A" h o r i z o n s ,  h o r i z o n s were s e p a r a t e d b y s e e p a g e  M  IB n e a r t h e  content  estimated  after  12 days'  specific gravity  1  gravity  decaying  Laboratory,  a n d maximum  s o a k i n g were  determined  a n d t h e maximum m o i s t u r e  0  con-  o f t h e two wood s a m p l e s were 0.146 and 0 2 1 b , a n d 826*4 o  and  b3b.b p e r c e n t , r e s p e c t i v e l y .  mum  moisture  content  Specific  gravity  were, a p p a r e n t l y , i n v e r s e l y  and maxi-  related.  - lb Microsite The Microsite  location,  characteristics,  and t o p o g r a p h y o f  4 were t h e same a s f o r M i c r o s i t e  Seedlings stock,  soil  4,  of l o c a l  and s o i l  1  0  p r o v e n a n c e were p l a n t e d  covered  by a slowly  s l a s h and b a r k o f Douglas  fir  decaying  as 1 + 1  mixture of  Q  M i c r o s i t e 5«> Microsite The  5 was l o c a t e d 12 m e t e r s n o r t h  "A" h o r i z o n s  were c o m p l e t e l y  loggingo  T h e "B" h o r i z o n s  contained  only s l i g h t  o f M i c r o s i t e 2o  removed i n t h e p r o c e s s o f  were 105  centimeters  seepage water,  Stoniness  t h i c k , and was e s t i -  mated a t 60 p e r c e n t . The  area  was f l a t  Seedlings  on t h e a r e a  l o g g i n g i n 1955o  first sition site  but well-drained  originated naturally after the  Examination  of the lesser vegetation  was r e p r e s e n t a t i v e  0  of the species  compo-  indicated that t h e o r i g i n a l  o f t h e Moss f o r e s t  type  (Krajina  1959)o The and  of shrubs,  l i c h e n s g r o w i n g on m i c r o s i t e s 1,  Table  f e r n s , herbs, 2 a n d 5,  mosses  arelisted i n  1, The  and  dominant s p e c i e s  s e e d l i n g s were c h o s e n r a n d o m l y f r o m e a c h m i c r o s i t e  t h e f o l l o w i n g c h a r a c t e r i s t i c s measured:  base diameter internode  of each internode,  angle  The l e n g t h a n d  of branches  on e a c h  and t h e l e n g t h a n d b a s e d i a m e t e r o f b r a n c h e s .  branch angle  was d e f i n e d  as t h e angle  between t h e  o f t h e s t e m above t h e b r a n c h a n d t h e m a j o r a x i s  major  The axis  of the branch  - 17 f o r a d i s t a n c e of 50 m i l l i m e t e r s *  The instrument used t o  measure t h e branch angle was a p r o t r a c t o r * One sample branch r e p r e s e n t i n g t h e average l e n g t h was taken from each i n t e r n o d e t o measure t h e l e n g t h of t h e seven largest needles* The bark t h i c k n e s s was measured f o r every i n t e r n o d e and seedling, with a c a l i p e r *  The depths of r o o t s were a l s o r e -  corded* The average v a l u e s of t h e f o l l o w i n g s e e d l i n g c h a r a c t e r i s t i c s were obtained f o r each m i c r o s i t e :  t o t a l height,  base diameter, branch angle of l a s t y e a r ' s s h o o t s , depth o f r o o t s , weight of l e a v e s , r o o t / s h o o t r a t i o , t o t a l number of branches, number and age of branches on each i n t e r n o d e , t o t a l l e n g t h o f branches on each i n t e r n o d e , t h e l e n g t h of l a r g e s t needles of each i n t e r n o d e p e r s e e d l i n g , and bark t h i c k n e s s of each i n t e r n o d e * THE INFLUENCE OF SITE ON HEMLOCK SEEDLINGS* The e f f e c t of s i t e on e x t e r n a l c h a r a c t e r i s t i c s of heml o c k s e e d l i n g s was s t u d i e d i n compartments 2A, 3A, 2B and 3B* E i g h t p l o t s were e s t a b l i s h e d p r e v i o u s l y t o study t h e growth of Douglas f i r on v a r i o u s s i t e s ( G r i f f i t h I960).  Five of  these p l o t s , P l o t 1, P l o t 5, P l o t 7, P l o t 8, and P l o t 4, were logged and t h e s l a s h and brush were p i l e d and burned.  The  s i t e i n d i c e s c a l c u l a t e d f o r Douglas f i r ranged from 85 f e e t , t o 180 f e e t .  S i t e index was determined f o r hemlock by u s i n g  the e q u a t i o n developed by J.H.G. Smith. + 0.663 S I of Douglas f i r . )  (SE  £  (Hemlock S I - 22.34  + 16.4').  The d e s c r i p t i o n  - 18 of these p l o t s was reported by G r i f f i t h c h a r a c t e r i s t i c s o f p l o t s a r e recorded The  (19b0).  The p h y s i c a l  i n Table 2.  average age of Douglas f i r on P l o t 1, P l o t 5, P l o t 7  and P l o t b, was 7b y e a r s , whereas on P l o t 4> t h e average age was bO y e a r s .  The s o i l was f a i r l y uniform  and was c l a s s i f i e d as b e l o n g i n g  on a l l p l o t s  t o the imperfectly drained  group of Brown P o d s o l i c s o i l s , with a maximum depth of 60 i n c h e s and a minimum depth of two i n c h e s . p l o t was e x c e e d i n g l y  The s o i l on every  stony, w i t h components v a r y i n g i n s i z e  from f i n e g r a v e l t o l a r g e b o u l d e r s . The  " A j " h o r i z o n s were u s u a l l y gray i n c o l o r and d i s -  continuous,  with v a r y i n g t h i c k n e s s .  The s o i l was non-  f r i a b l e , n o n - p l a s t i c , n o n - s t i c k y and had a pH range from 3*8 t o 4.9.  The "B" h o r i z o n s v a r i e d from yellowish-brown t o  d a r k brown, g e n e r a l l y with conspicuous m o t t l i n g a t t h e lower levels.  The t e x t u r e was coarse, sandy loam, o f t e n c o n t a i n i n g  f i n e g r a v e l and s m a l l c o n c r e t i o n s .  The s o i l was c l a s s i f i e d  as f r i a b l e t o f i r m , n o n - s t i c k y and with a n o n - p l a s t i c consistency.  The pH ranged from 4»8 t o 5»7»  were of g l a c i a l t i l l  surface, the C W  M  M  horizons  Where t h e bedrock was c l o s e t o  h o r i z o n was absent.  ness of the *C * h o r i z o n was 31 i n c h e s . I  The maximum t h i c k The t e x t u r e was  coarse sandy loam, o f t e n c o n t a i n i n g f i n e g r a v e l . the C " h o r i z o n ranged from 5.3 t o 5»9« n  W  o r i g i n , l i g h t t o dark gray i n c o l o r ,  often with reddish mottling. the  The C  The pH o f  The depths of s o i l  h o r i z o n s and t r e e root p e n e t r a t i o n f o r the p l o t s were summa r i z e d by G r i f f i t h  (I9b0) i n h i s Table 3.  - 19  -  S e v e r a l s p e c i e s - Pseudotsuga m e n z i e s i i (Mirb) Franco, Tsuga h e t e r o p h y l l a (Rat) Sarg, Abies a m a b i l i s  (Dougl.),  Thuja p l i c a t a Donn., Abies g r a n d i s L i n d . , P i c e a s i t c h e n s i s (Bong), L a r i x decidua M i l l , L a r i x l e p t o l e p i s ( S i e b , and L a r i x e u r o l e p i s (Henry) and  Populus spp* - were p l a n t e d on  logged-over p l o t s as a p a r t of P r o j e c t 57-10 Committee, 1959)*  The  Ten  two  the  (U.B.C* F o r e s t  s e e d l i n g s were p l a n t e d i n s u b - p l o t s  36 f e e t by 36 f e e t , i n a r e p l i c a t e d randomized, b l o c k Each p l o t contained  Zucc*L,  s u b - p l o t s f o r every  design*  species*  hemlock s e e d l i n g s , s e l e c t e d at random, were measured  on each s u b - p l o t , except P l o t 4, where only s i x s e e d l i n g s were a v a i l a b l e because of high m o r t a l i t y * measured f o r every hemlock s e e d l i n g . on the l a t e s t i n t e r n o d e was  The  measured and  Height angle  growth  was  of branches  recorded*  s e e d l i n g , the l e n g t h of the seven l a r g e s t needles  For each was  ob-  t a i n e d from the samples taken from the l a t e r a l branches of the l a t e s t  internode*  -  20 -  R E S U L T S THE INFLUENCE OF LIGHT INTENSITY ON HEMLOCK SEEDLINGS The The  total  amount o f l i g h t  d a y s and c l o u d y d a y s . the  control  Figures on  amount or* l i g h t  plot  and on P l o t s  except  Plots  the  of l i g h t  intensity  of cumulative  V and 5* i n c r e a s e d  ceased  light  are n e g l i g i b l e  on sunny d a y s , t h e l i g h t  sun's r a y s s t r u c k t h e p l o t s  graphs  on sunny on  I and 1 ( T a b l e s 5 and b ) .  1400 h o u r s , when t h e y were f r e e crease  on e a c h p l o t  These d i f f e r e n c e s  2 and 3 show t h a t ,  each p l o t ,  varied  o f shadow.  gradually  This  soon a f t e r  until  rapid i n -  1700 h o u r s , when  a t an a c u t e a n g l e .  intensity  intensity  The  on a c l o u d y d a y  (Figures  4 and 5) show a more u n i f o r m p a t t e r n , b e c a u s e t h e shadows o f the  tall  crease until  trees  were n o t o b v i o u s i n t h i s  of l i g h t  intensity  case,  A greater i n -  b e g a n a t 1200 h o u r s and  lbOO h o u r s , when t h e l i g h t  intensity  lasted  decreased  slowly  (Table b ) . The for  r e l a t i v e per cent  each p l o t ,  under p a r t i a l This  by d i v i d i n g  of l i g h t  e n c e s , because the t o t a l  (Tables light  on open p l o t s  5 and b ) .  readings  intensity  the t o t a l  shade by t h e t o t a l  r e l a t i v e per cent  was g r e a t e r  of l i g h t  value  value  on open  days  (Table  readings  the  different  days  of t h e c u m u l a t i v e  o) i s u s e d i n t h i s  c o m p a r i s o n s between v a r i o u s s e e d l i n g  plots.  differ-  on s u n n y d a y s , t h a n on c l o u d y  for  plotso  readings  shows g r e a t  intensity  The r e l a t i v e p e r c e n t  on c l o u d y  obtained  of l i g h t  received  intensity  of the l i g h t  was  characteristics  thesis on  - 21 -  f  T h i s measurement or' l i g h t i n t e n s i t y i s p r e f e r r e d because the shadows of t r e e s were minimized under these c o n d i t i o n s The  0  average r e l a t i v e percentages of l i g h t i n t e n s i t y f o r  P l o t s I , I I , I I I , IV, 1, 2, 3, and 4, were 1 3 ° 5 9 , 31c4b,  45o42, 56,33, 10,32, 22,72, 42,54, and 55*90 per cent r e s pectively  (Table  b).  The i n f l u e n c e o f l i g h t i n t e n s i t y on t o t a l height o f s e e d l i n g s . The data  (Tables 7 and 8) and graphs ( F i g u r e s t> and 7)  show t h a t t h e r e i s a great d i f f e r e n c e i n t o t a l height between those s e e d l i n g s which had grown i n the open and those which had grown under the shade of f o r e s t canopy.  The a v e r -  age height o f 11-year-old s e e d l i n g s i s 37&0 m i l l i m e t e r s on P l o t 5 while s e e d l i n g s growing under a l i g h t i n t e n s i t y , per cent of t h a t i n the open on P l o t 1 (Table b), average height of 307 m i l l i m e t e r s .  10,32  show an  The e f f e c t of l i g h t i n -  t e n s i t y on t o t a l height i s the same on s i x - y e a r - o l d seedl i n g s as on 11-year-old s e e d l i n g s .  The average t o t a l  height  of hemlock s e e d l i n g s grown i n the open on P l o t V i s 1617,0 m i l l i m e t e r s while s e e d l i n g s grown i n an average r e l a t i v e l i g h t i n t e n s i t y of 13»59 per cent on P l o t I have an average t o t a l height of 133ol m i l l i m e t e r s (Tables 9 and 10) o The  i n f l u e n c e of l i g h t i n t e n s i t y on the age t o reach b r e a s t height o I t i s obvious  from the data presented  i n T a b l e s 7,  8,  9, and 10, and from F i g u r e s b and 9 . t h a t s e e d l i n g s grown under d i f f e r e n t l i g h t i n t e n s i t i e s w i l l reach b r e a s t height a t d i f f erent ages w i t h i n s i m i l a r s i t e s . l i n g s reached  Open grown hemlock seed-  b r e a s t height a t s i x y e a r s , w h i l e  seedlings  -  22  -  growing on P l o t 4 w i t h a r e l a t i v e l i g h t i n t e n s i t y per  cent needed e i g h t y e a r s t o grow t o b r e a s t  of 55*90  height.  S e e d l i n g s growing on P l o t 3 grew i n a r e l a t i v e l i g h t sity  inten-  of 4 2 , 5 4 p e r cent and needed 9 ° 5 y e a r s t o r e a c h b r e a s t  height o The e f f e c t  of l i g h t i n t e n s i t y  on diameter growth.  Diameter growth showed a response t o i n c r e a s e d i n t e n s i t y s i m i l a r t o t h a t of h e i g h t growth.  The average  base diameter of 1 1 - y e a r - o l d s e e d l i n g s growing under on P l o t 5 i s 51o91 m i l l i m e t e r s .  light  8),  2,44 millimeters respectively  2,and 1 a r e 29.16, ( T a b l e s 7 and  The average base diameters of s i x - y e a r - o l d  growing on P l o t s 5* 4* 3 2 03,  S  2 and 1 a r e 2 5 . 7 0 ,  and 1 , 3 b m i l l i m e t e r s r e s p e c t i v e l y  o  full  The average base  d i a m e t e r s of s e e d l i n g s grown on P l o t s 4* 3s 15»95> 7 . 5 , a n d  light  seedlings  7.17, 2 , 8 0 ,  ( T a b l e s 9 and 1 0 ) ,  The graphs of cumulative mid-diameter of i n t e r n o d e s show the same t r e n d  ( F i g u r e s 8 and 9) as the graphs of cumulative  p e r i o d i c l i g h t - i n t e n s i t y measurements ( F i g u r e s 2, 3» 4> and 5)  The comparison o f the mid-diameters of the l a s t y e a r s 8  0  i n t e r n o d e s shows a tendency t o decrease from P l o t 5 t o P l o t The mid-diameters of s e e d l i n g s on P l o t 5 , 4* 3 5,1,  2 , 4 5 , 1,82, 1,11  y  2, and 1 were  and 0 7 5 m i l l i m e t e r s r e s p e c t i v e l y . o  The  mid-diameters of l a s t y e a r ' s shoots of s e e d l i n g s on P l o t V, IV, I I I , I I and I were 4 . 3 5 , 1,80, 0 . 7 0 , 0 . 5 9 and 0 , 5 0 m i l l meters  respectively.  1,  -  The  i n f l u e n c e of l i g h t  The old  average t o t a l  seedlings  77,4,  34*5  -  23  intensity  of  number o f g r e e n b r a n c h e s  on P l o t 5 *  4,  3s  2 and  8,6 r e s p e c t i v e l y *  and  on d e n s i t y  1  The  0  11-year-  of  133*5,  was  foliage  104,6,  average t o t a l  number  of green branches  of s i x - y e a r - o l d s e e d l i n g s  on P l o t s V,  III,  8 9 . 5 , 41*0,  3 d respectively  I I and  (Tables  7,  I was 8,  9 and  10),  16.6,  The  8 * 0 and  graphs  o f c u m u l a t i v e number  o f g r e e n b r a n c h e s f o r b o t h a g e s show a s i m i l a r graphs  of cumulative l i g h t  Light Tables first  internode  defined diameter the  8,  7,  9 , and  10  for a l lplots*  of the i n t e r n o d e ) *  of P l o t s i , 2 ,  respectively.,  3,  The  The  (Form q u o t i e n t  respectively.  The  base  year's internode  5 are 0 * 8 , 0 . 7 5 ,  0 76, o  when i s  and  f r o m P l o t s I , I I , I I I , IV and of 0*82,  0*76, 0 * 7 8 ,  average form quotient  t o decrease with increased Natural  was  average form q u o t i e n t s f o r  4,and  Seedlings  of the  difference i s evident  of the l a s t  have an a v e r a g e f o r m q u o t i e n t  dency  quotient*  a s t h e r a t i o b e t w e e n t h e m i d d l e d i a m e t e r and  compared between p l o t s .  0,68  form  11)o  show t h a t t h e f o r m q u o t i e n t  i s similar  average form q u o t i e n t  trees  and  trend t o the  ( F i g u r e s 1 0 and  intensity  intensity  IV,  p r u n i n g and  0*63  V  0.78 and s  showed a  ten-  light.  light  intensity.  Dead b r a n c h e s were n o t f o u n d on s i x - y e a r - o l d s e e d l i n g s in  Plots I to V  dead b r a n c h e s 6.4,  and  (Tables  9 and  on P l o t s 1 ,  10).  2, 3 ,  The  4,and  5 , 0 r e s p e c t i v e l y (Tables  a v e r a g e number o f  5 i s 0,2, 1 * 5 ,  7 and  8),  5,0,  Seedlings  growing  - 24 on t h e h e a v i l y s h a d e d P l o t 1 were a l m o s t f r e e h a v i n g b e e n p r u n e d n a t u r a l l y on t h e f i r s t  o f dead  five  branches  internodes  In  contrast,  2,  3* and 4 i n d i c a t e d t h a t n a t u r a l p r u n i n g on t h e f i r s t  internodes  t h e number o f d e a d b r a n c h e s on s e e d l i n g s  was o n l y b e g i n n i n g s  o f P l o t 5 were s t i l l  light  i n the very  p r u n i n g w i t h dead b r a n c h e s THE  Seedlings  0  on P l o t s five  growing i n the  full  e a r l y stages of n a t u r a l  on t h e f i r s t  four internodes  only  0  E F F E C T OF MICROSITE ON THE EXTERNAL CHARACTERISTICS OF THREE-YEAR OLD HEMLOCK S E E D L I N G S 0  heighto  Total The  average t o t a l heights  1, 2, 3* 4,  site  and  5  of seedlings  74b 2, 73b 2, 620 9, 522,7 a n d  are  s  s  4b9«4  millimeters  15)o  S t u d e n t s t - t e s t was a p p l i e d  respectively  No s i g n i f i c a n t 1 a n d 2,  between M i c r o s i t e s differences existed sites  1 a n d 3,  and 2 and 3  3 and 5 ( T a b l e  lings  17)o  o f each m i c r o s i t e  Microsite  1 t o 5»  3 a n d 4* a n d 4 a n d 5e  Highly  0  significant  The a v e r a g e t o t a l h e i g h t  2 a n d 5* of seed-  showed a t e n d e n c y t o d e c r e a s e f r o m 12)  (Figure  0  Diameter  0  i s different for  T h e a v e r a g e b a s e d i a m e t e r s on M i c r o s i t e s  2, 3, 4  and  meters,  respectivelyo  are  Micro-  differences  2 a n d 4*  The a v e r a g e b a s e d i a m e t e r o f s e e d l i n g s  5  Significant  l e v e l between t o t a l h e i g h t s o f  Base  each micrositeo  differences  l e v e l between  1 a n d 4* 1 a n d 5*  on M i c r o s i t e s  the  and  d i f f e r e n c e s were f o u n d  o  o  and  t o evaluate  at the0 05-per-cent  were o b s e r v e d a t t h e 0 01 seedlings  o  11, 12, 13* 14  (Tables  8  between m i c r o s i t e s o  g r o w i n g on M i c r o -  llo44*  b 09, o  7olb, b«53,  and  5o95  1,  milli-  S t u d e n t * s t - t e s t was u s e d t o e v a l u a t e  - 25 the  -  d i f f e r e n c e s i n base diameter f o r each m i c r o s i t e ,  significant  d i f f e r e n c e s were f o u n d b e t w e e n v a l u e s  sites  3,  2 and  ficant  4,  3 and  differences exist  s i t e s 1 and and  3 and  2,  5 (Table  grown on  1 and  18)  3;  The  0  5  at the 1 and  lings  on  (Figure  the  different  pattern  diametero  The  3,  2,  3<>75 and  and  as  4 were p l a n t e d  fact  weight for  root that  of  than  i s that  decreasing  2  seedlings  from  Microsite  seed-  d i d not  f o l l o w the  same  o r d e r ) was  seedlings  artificially  and  on M i c r o s i t e  on M i c r o s i t e s  evaluation  height  and  base  12,7,  4*32,  14) »  The  1, 4°21,  explan-  g r o w i n g on  Microsites  t h e y had  better  a  originating naturally. 2 had  3 and  of m i c r o s i t e  less  average  4 excluded the  need  d i f f e r e n c e s based  roots.  However, t h e  in  and  of r o o t s f o r M i c r o s i t e s  roots.  W e s t e r n h e m l o c k i s known a s  different  4,  of hemlock  observed f o r t o t a l  Depth of  in  2 and  Micro-  of r o o t s  microsites  seedlings  statistical  species.  5,  signi-  roots»  system than s e e d l i n g s  of r o o t s  on w e i g h t  1 and  0  order  Highly  l e v e l between  2 98 grams r e s p e c t i v e l y ( F i g u r e  1 and  The  of  5 ( i n decreasing  of t h i s  5°  f o r Micro-  13)•  average weight  ation  developed  4,  average oven-dry weight  decreasing  4,  0,01  4 and  decreases gradually  Weight The  and  average base diameter of  each m i c r o s i t e  1 to Microsite  5,  No  effect  a typically  shallow-rooted  of d i f f e r e n t m i c r o s i t e s  depths of r o o t i n g .  The  average depth of  order, i s 220,6, 191,1, 178,7, 133<»0 and  resulted roots, 120,4  - 26 millimeters f o r Microsites 1, 5* 2, 3 and 4 respectively,, (Figure 15)o  Although the contrast between Microsites 1 and  4 i s great, seedlings on both microsites had the same average depth of roots one year before the study was made since they were grown i n the nursery.  Root depth f o r seedlings on  Microsite 1 was closely followed by that of seedlings on Microsite 5 which had the lowest rate i n height and diameter growth. Weight of needles. The average oven-dry weight of needles of seedlings growing on Microsites 1, 2, 3* 4, and 5 was 1 5 . b 4 , 9o94, 8 . 0 , 4<>95 and 5.66 grams respectively (Figure lb)„  The average  weight of needles of seedlings follows the same pattern as the average height and average base diameter, with the exception of the values f o r seedlings of Microsite 5* which have a s l i g h t l y greater average weight than those on Micros i t e 4«  Student s t - t e s t was applied t o evaluate the d i f f e r B  ences between microsites.  Mo s i g n i f i c a n t differences were  found between Microsite 2 and 3, and 4 and 5©  Significant  differences existed at the 0 . 0 5 l e v e l between Microsite 1 and 2, 2 and 5* and 3 and 5» Highly s i g n i f i c a n t differences were' at the 0«01 l e v e l between Microsite 1 and 3y 1 and 4* 1 and 5 , 2 and 4, and 3 and 4 (Table 21)  0  Root/Shoot r a t i o . The average r a t i o f o r Microsites 1, 2, 3? 4, and 5 was O 3b9, 0.19b, 0.286, 0*432, and 0 3 0 7 respectively o  0  11, 12, 13» 14 and 15)o Microsites i n decreasing root/shoot  r a t i o are 4  S  (Tables order of  1* 5* 3* and 2. The r e s u l t s indicate  - 27 t h a t t h e r a t i o was h i g h e s t t o r p l a n t e d s e e d l i n g s and lowest t o r those which o r i g i n a t e d  naturally.  T o t a l number o f branches. 1,  The average t o t a l number o f branches f o r M i c r o s i t e s  2, 3, 4 and 5 was 38,4, 39o0, 34ol, 30,6 and 31eb r e s p e c t i v e l y 1,  ( F i g u r e 17)©  The d e c r e a s i n g order o f M i c r o s i t e s i s 2,  3,  The d i f f e r e n c e s between M i c r o s i t e s 1 and 2  5,  and 4°  and 4 and 5 are s o s m a l l , t h a t they a r e n e g l i g i b l e .  Gener-  a l l y t h e t o t a l number o f branches o f M i c r o s i t e s f o l l o w e d t h e same p a t t e r n as t o t a l height and base. The average number of branches by i n t e r n o d e s . The average number of branches on the f i r s t i n t e r n o d e f o r M i c r o s i t e s 1, 6,4  respectively.  2,  3,  4,  0  6,9,  and 13,3  of poor p l a n t i n g t e c h n i q u e and i s  2,  3, 4,  respectively.  and 5 was 18,8,  16Jo  14.1,  13 o 2, 8,8  15o5,  The average number of  branches on the t h i r d i n t e r n o d e f o r M i c r o s i t e s 1, and 5 was 17,0,  5e4, and  The average number of branches on t h e second  imternode f o r M i c r o s i t e s 1, 14o0, 16,4,  9o4,  The s m a l l number of branches on s e e d l i n g s  on M i c r o s i t e 1 i s a r e s u l t discussed l a t e r  and 5 was 2,6,  and 12,2  2,  respectively  3,  4,  (Table  The average number o f branches on the t h i r d i n t e r n o d e  shows a decrease from M i c r o s i t e 1 t o 5, w i t h t h e e x c e p t i o n of M i c r o s i t e 5» Average branch angle of i n t e r n o d e s . The average branch angle o f t h e f i r s t i n t e r n o d e on M i c r o -  s i t e 1, 2, 3, 4, and 5 was 35.00, 47o98 42,54, 43o70, and s  37«81 degrees r e s p e c t i v e l y .  The average branch angle of t h e  - 2b -  on M i c r o s i t e s 1,  second i n t e r n o d e  2, 3* 4* and 5 was 49ol5*  48.54* 43o57, 3b 03 and 43«79 degrees r e s p e c t i v e l y . The 0  average branch angle of the t h i r d i n t e r n o d e  on M i c r o s i t e s  1, 2, 3* 4 and 5 was 39.3* 3b 8 2, 34.4* 27 o 8 and 2b09 degrees respectively. a decreasing The  The branch angle o f t h e t h i r d i n t e r n o d e p a t t e r n from M i c r o s i t e s 1 t o 5 ( F i g u r e  d i f f e r e n c e s between m i c r o s i t e were evaluated  Student's t - t e s t .  shows  18)  c  by u s i n g  No s i g n i f i c a n t d i f f e r e n c e s were found  between M i c r o s i t e s 1 and 2, 2 and 3* and 4 and 5.  Highly  s i g n i f i c a n t d i f f e r e n c e s a t the 0.01 l e v e l were observed between M i c r o s i t e s 1 and 3, 1 and 4* 1 and 5, 2 and 4* 2 and  5* 3 and 4* and 3 and 5 (Table  19).  Average l e n g t h of l a r g e s t n e e d l e s by i n t e r n o d e s . The internode  average l e n g t h of t h e l a r g e s t needles on t h e f i r s t of s e e d l i n g s  on M i c r o s i t e s 1* 2, 3* 4* and 5 was  14.9b, 17.36, 16.36, 12.87* and 14.96 m i l l i m e t e r s r e s p e c t i v e l y . The  average l e n g t h o f t h e l a r g e s t needles of t h e second  inter-  node on M i c r o s i t e s 1 , 2, 3* 4* and 5 was 17.22, 17.59* 16 81, 0  15«55* and 15.31 m i l l i m e t e r s r e s p e c t i v e l y . length  The average  of t h e l a r g e s t n e e d l e s of t h e t h i r d i n t e r n o d e on  M i c r o s i t e s 1, 2, 3* 4, and 5 was 13»79, 1 2 . 2 6 , 11.44* 10.14* and  9.01 m i l l i m e t e r s r e s p e c t i v e l y ( F i g u r e 19)«>  length  o f t h e l a r g e s t needles on t h e t h i r d i n t e r n o d e  the same d e c r e a s i n g and  The average showed  p a t t e r n from M i c r o s i t e s 1 t o 5* a s height  average base diameter and branch a n g l e .  between m i c r o s i t e s based on l e n g t h by u s i n g Student's t ~ t e s t  0  The d i f f e r e n c e s  of needles were  evaluated  No s i g n i f i c a n t d i f f e r e n c e s were  -  2 and  found between M i c r o s i t e s a t 0*05 l e v e l  differences 1  2,  and  2 and  ences e x i s t e d 4*  and  1  The of  151*97, The  Microsites  1,  87*76, and  87*25  2,  3,  4,  4*  3 and  4,  5  1,  2,  3,  1*87  millimeters  of b r a n c h e s  internode  4,  180 73,  5 was  and  internode  (Figure  third  internode  1*14,  0*92,  21)o  The  The  The  1*32,  on M i c r o s i t e s  0*48,  and  length  average  The  22*67  and  internodes* of f i r s t  1*15,  average base  and  1*21  2,  3,  internode  4,  1,  1*14, diameter 2,  3,  millimeters  5 was  respectively showed s i m i l a r  t o 5 as h e i g h t ,  of l a r g e s t n e e d l e s *  and  internode  1*78,  on M i c r o s i t e s  a  1  1,  on M i c r o s i t e s  2*78, 1 * 7 6 ,  0 7 1 millimeters  data of the t h i r d  on  20)*  5 was  1,  0  117*83,  average base diameter of branches  from M i c r o s i t e  a n g l e and  and  0  respectively  4b*48, 12*87, 1 8 * 6 7 ,  respectively*  2*54, 1*56,  respectively*  pattern  4,  Q  171*08, 1 2 4 * 3 2 ,  of the second internode  5 was  and  3*  1  on t h e f i r s t  respectively*  of the t h i r d  2,  3,  and  internodeso  average base diameter of branches 1,  1  differ-  of the second i n t e r n o d e  5 was  millimeters  Microsites  20)  145*6 millimeters  and  52*24,  (Table  o f b r a n c h e s by  of branches  respectively  on M i c r o s i t e s  Significant  significant  a v e r a g e b a s e d i a m e t e r o f b r a n c h e s by  The  4,  3,  5 was  and  millimeters  and  2,  Highly  of branches  of branches  The  length  on M i c r o s i t e s  length  5«  and  l 6 l * 8 4 , 1 0 5 * 9 4 and  average  length  5,  average l e n g t h  seedlings  3 and  and  l e v e l between M i c r o s i t e s  2 and  average  3,  were o b s e r v e d between  4 and  and  a t 0*01 5,  and The  4,  -  29  of the 1*57, (Figure decreasing  base diameter, branch  - 30 Bark The  average bark thickness  1,  Microsites 0. 69  5  and  0.77,  0.58  1,  3* 4,  2,  millimeters  o  5,  7,  8,  millimeters Plots  and  1959  256.0, and  8,  64.0  differences  4 was  i n 1959  5, 7,  1,  4  and  in  8  i960  4  was  49o02,  grees r e s p e c t i v e l y . P l o t s was ficant  The  e v a l u a t e d by  the  difference  significant  was  o  was  The  o  Micro-  0 36  and  o  was  measured  h e i g h t g r o w t h on  108.0, 106.4 The  and  in Plots  44.2  h e i g h t growth  on  307.8, 3 1 6 . 9 , 283o4, (Table  I960 g r o w i n g  22)o  The  already  differences  season.  angle. o f I 9 6 0 i n t e r n o d e s on  difference  35»07  and  Student*s t - t e s t .  were o b t a i n e d  5 and  Plots  24.72  i n branch angle  f o u n d between P l o t s  differences  was  HEMLOCK SEEDLINGS.  41*01, 3 9 « 9 2 ,  using  on  0 37^  growing season, these  during  5  respectively.  internode  respectively  average branch angle and  4 and  g r o w t h b e t w e e n p l o t s were  1959  the  thickness  growth.  19b0  millimeters  and  16).  S I T E ON  and  3*  Q©47, 0 . 5 8 ,  o  Branch  5, 7,  third  respectively.  became e v e n g r e a t e r  The  2,  millimeters  128.6, 1 0 8 . 3 ,  i n height  obvious during  1,  on  0.75*  average bark  growth of hemlock s e e d l i n g s  growing seasons of 1,  0.99,  (Table  INFLUENCE OF  height  0,86,  0 6l,  Height The  1.42,  of the  was  respectively  THE  internode  The  0 54  and  5  and  first  on M i c r o s i t e s  average bark t h i c k n e s s  sites  of the  was  respectively.  second i n t e r n o d e  0.75,  1.18, The  3, 4*  2,  millimeters  of the  thickness.  No 7«  1,  de-  between signiHighly  comparing P l o t s  1  and  - 31 5,  1  and  and 7, 1 and 8 , 1 and 4, 5 and 8 , 5 and 4, 7 and 4, and 8 and 4 r e s p e c t i v e l y (Tables  23)©  7  The upper and  lower f i d u c i a l l i m i t s were c a l c u l a t e d f or each p l o t a t t h e 0 0 1 levels o  The v a l u e s are given i n Table 25o  there was no o v e r l a p p i n g  Generally  between p l o t s except i n t h e case  of P l o t 5 and F l o t 7 where the lower l i m i t of P l o t 5 overlapped the upper l i m i t o f P l o t 7o  T h i s f a c t i s understandable  because t h e s i t e - i n d e x d i f f e r e n c e between those p l o t s i s only 6 f e e t ,  (The e x c e p t i o n of t h i s l a t t e r case i n d i c a t e s  t h a t t h e means of P l o t s 1 , 5, 8 , and 4 a r e d e r i v e d heterogeneous p o p u l a t i o n )  0  from a  The upper and lower f i d u c i a l  l i m i t s were p l o t t e d over the s i t e i n d i c e s ,  The c o r r e l a t i o n  e x i s t i n g between p l o t s i t e index and average branch angle was found t o be l i n e a r , 0 984, o  The c o r r e l a t i o n c o e f f i c i e n t i s  which i n d i c a t e s a h i g h l y s i g n i f i c a n t r e l a t i o n between  s i t e index (X) and branch angle ( Y )  c  (Correlation coefficient  i s s i g n i f i c a n t f o r 3 Df a t the 0 , 0 1 l e v e l a t 0,959)o  The  c a l c u l a t e d equation o f the l i n e i s Y <=> 0 , 3 7 4 X =5*88 and i s i l l u s t r a t e d i n Figure 22, Length of the l a r g e s t n e e d l e s . The  average l e n g t h  of needles of t h e lybO i n t e r n o d e on  P l o t s 1, 5, 7, 8 and 4 was l b , 7 4 , 8,14  respectively.  13*76,  14,17, 12,68 and  The d i f f e r e n c e s between p l o t s were eva-  l u a t e d s t a t i s t i c a l l y by u s i n g Student's t ~ t e s t .  No s i g n i -  f i c a n t d i f f e r e n c e s were found between P l o t s 5 and 7«  Highly  s i g n i f i c a n t d i f f e r e n c e s e x i s t e d between P l o t s 1 and 5, 1 and  7,  1  and 8 , 1 and 4, 5 and 8 , 5 and 4, 7 and 8 , 7 and 4, and  8 and 4 (Table  24)«.  The upper and lower f i d u c i a l l i m i t s were  -  calculated except 7  i n the  0  case  with a value  of  o  the f a c t feet  level.  of  over the  site  was  calculated  The  0  u p p e r and  (X)  appears  equation  t o 5«  The  needle  l e n g t h and  t o be  of l i n e  a n g l e was  average The  direct  fiducial  linear.  l e n g t h of the  largest  to only  of each p l o t .  The  needles  The  0 , 1 3 2 X  (Y)  correlation of  0 , 9 8 8 ,  - 2 , 3 8 ,  l e n g t h i t was  poss-  indices f o r Microsites 1 angle  characteristics  In M i c r o s i t e 5 the  g r e a t e r than  of  were  with a value  on n e e d l e  of these 27o  limit  Plot  limits  r e l a t i o n s h i p between b r a n c h either  of  i n d i c e s was  index  is Y =  t h e unknown s i t e  i s shown i n T a b l e  branch  the lower  lower  site  highly significant  t o determine  index  upper l i m i t  l e n g t h of l a r g e s t  U s i n g the equations based ible  The  overlap  T h i s o v e r l a p p i n g i s due  1 3 o 3 1 «  corresponding  indices  coefficient The  7o  higher than  c o r r e l a t i o n between average and  These v a l u e s d i d not  5 and  was  14«67  -  t h a t t h e d i f f e r e n c e between s i t e  (Table 2 6 )  plotted  0 1  of P l o t  5 with a value  Plot  6  at the  32  in "icrosite needles  was  4,  and  and  site  average  while  the  significantly  less.  r e l a t i o n s h i p s d e m o n s t r a t e d f o r M i c r o s i t e s 1 t o 4 become  obscured sonable  f o r M i c r o s i t e 5.  It i s difficult  explanation f o r this  variability.  to offer  a  rea-  - 33  -  D I S C U S S I O N THE  INFLUENCE OF  Baker upon a s  LIGHT INTENSITY ON  (1950) had  t h i s t o say:  HEMLOCK SEEDLINGS.  "The  f o r e s t may  a c e l l u l o s e f a c t o r y i n w h i c h w a t e r and  be  carbon  a r e u s e d t o make wood t h r o u g h t h e  agency of s u n l i g h t ,  mineral  compounds a r e  n u t r i e n t s and  only parts  nitrogenous  of the machinery  of the  light  can  g o v e r n e d , t h e i r f o r m and  be  be  modified,  certain be  the  limits  strength  and  the  sun  nature©  i s the  1959,  t i m e and  i n t e n s i t y may  of the  trees  crown  changed  can  within can  degree." of l i g h t  for plants  which o r i g i n a t e d from  intensity  Laverick  1958,  i n f l u e n c e the  m y c o r r h i z a e , and  and  quality  t h a t Kramer and of l o b l o l l y  intensity  up  to f u l l  p e t i n g hardwoods was  Light  growth of p l a n t s , f o r m a t i o n  Decker  pine  other  (Gerhold  K o z l o w s k i 1957)o  d i s c o l o r a t i o n of f o l i a g e ,  synthesis  (1944) f o u n d t h a t  increased with  light,  Kozlowski  at  increase  one-third  (1957)  photoof  whereas p h o t o s y n t h e s i s  greatest  of  light  o f com-  or l e s s  of  full  intensity, Bourdeau  was  manipulation  amount o f r e p r o d u c t i o n  a l l - i m p o r t a n t source  the necessary  B o u r d e a u and  light  character  The  However s e v e r a l e x p e r i m e n t s have shown i t i s  with  reported  By  0  o f t h e wood c a n be  p o s s i b l e t o grow p l a n t s i n l i g h t sources  factory  dioxide  essentially  r a t e of growth of i n d i v i d u a l  governed t o a c o n s i d e r a b l e The  in  f a c t o r the  of the  looked  reduced  Gerhold resulted  (1958) s t a t e d t h a t t h e  c o n s i d e r a b l y by  (1959) r e p o r t e d i n decreased  that  growth of e a s t e r n  exposure t o low  light  reduction i n light  hemlock  intensity,  intensity  needle d i s c o l o r a t i o n of Scots  pine.  - 34 The of l i g h t results great  literature to forest of t h i s  varied  on  creased  of l i g h t  the  sunny d a y s  5 and  b)  0  f o u n d t o be  on a b r i g h t and Expressing  average l i g h t  d i v i d i n g the s h a d e by  the  to f u l l  light  total  light  total  i n the  ages under u n i f o r m  buted t o the  intensity,  open  height  As  0  seedlings  o f up  site  day  candles  i n per 10  of  open a r e a  0  related to light a consequence  i n f l u e n c e of v a r i o u s l i g h t  cent  hemlock calculated under growth  intensity  of the i n -  reached b r e a s t  growth.  cent  per  Height  height  at  Differences i n  t o f o u r o r more y e a r s  same manner a s h e i g h t  for  i n foot-candles)  conditions.  on  respectively  i n t e n s i t y was  were  attri-  intensities.  Diameter growth responded t o i n c r e a s e d the  change  intensxty for a  found that  light  directly  also  days, than  for survival  on t h e  amounts  noon,  r a t e of  l a t t e r value  (expressed  light  various  to breast  The  lly-foot  i t was  average r e l a t i v e  light  in  Seedlings  0  intensity  cloudy  cloudy  intensity  creased  age  on  and  this  intensity,  t h e minimum r e l a t i v e The  117  a  Light intensity i n -  a v e r a g e d a i l y minimum l i g h t  h o u r p e r i o d was  the  i n t e n s x t y has  light  afternoon.  more g r a d u a l  o f hemlock s e e d l i n g s was up  by  received various  The  e  of day.  g r a d u a l l y i n the  The  seedlings. by  and  t o the time  hemlock s e e d l i n g s  was  light  stand,  importance  i s supported  growth of hemlock s e e d l i n g s  i n t e n s i t y was  sunny days.  of the  of the  g r a d u a l l y i n the morning, r e a c h e d i t s peak a t  decreased  (Tables  T h i s evidence  canopy of t h e  cloudy  according  ample e v i d e n c e  w h i c h shows t h a t  i n f l u e n c e on t h e  of l i g h t  12  growth,,  study  growing under the  and  provides  light  intensity  D i a m e t e r measurements  - 35 from root c o l l a r t o t e r m i n a l shoot were s i m i l a r l y by v a r i a t i o n i n l i g h t The creased  influenced  intensity.  average number of green branches i n c r e a s e d with i n light intensity.  N a t u r a l pruning  was not e v i d e n t on  s i x - y e a r - o l d s e e d l i n g s and no dead branches were found on such s e e d l i n g s .  Eleven-year-old  s i g n s of n a t u r a l p r u n i n g .  s e e d l i n g s , however, showed  The r a t e of n a t u r a l pruning  inversely related to light intensity.  was  The t a p e r of s e e d l i n g s  growing s l o w l y under r e l a t i v e l y low l i g h t i n t e n s i t i e s , was apparently  l e s s than t h a t of s e e d l i n g s growing v i g o r o u s l y  under r e l a t i v e l y h i g h l i g h t  intensities.  THE EFFECT OF MICROSITE ON THE EXTERNAL CHARACTERISTICS OF THREE-YEAR-OLD  HEMLOCK SEEDLINGS.  T o t a l height was a t a maximum f o r 1 + 1 planted  on M i c r o s i t e 1.  nursery  Although s e e d l i n g s u s u a l l y  a set-back immediately f o l l o w i n g p l a n t i n g , p l a n t e d M i c r o s i t e 1 was l a r g e r than n a t u r a l r e g e n e r a t i o n 2.  T h i s d i f f e r e n c e i s probably  explained  stock experience  stock on  on M i c r o s i t e  i n term of the  d i f f e r e n c e s of root systems as w e l l as the p l a n t i n g s i t e s . Natural regeneration  growing i n decaying  s i t e 3 was l a r g e r and more v i g o r o u s growing i n the poor m i n e r a l  soil  wood i n M i c r o -  than n a t u r a l  of M i c r o s i t e 5.  regeneration Hemlock  seedlings, r e p r e s e n t i n g a homogeneous p o p u l a t i o n when p l a n t e d , developed, w i t h i n one y e a r , h i g h l y s i g n i f i c a n t d i f f e r e n c e s i n height growth on M i c r o s i t e 1 compared t o M i c r o s i t e 4. S i m i l a r d i f f e r e n c e s were observed f o r base diameter measurements except where h i g h l y s i g n i f i c a n t d i f f e r e n c e s were  -  demonstrated Microsite  statistically  1 and n a t u r a l  3b -  between a r t i f i c i a l  regeneration  on M i c r o s i t e  base diameter decreased from M i c r o s i t e The that  microsites,  seedlings  average weight nutrients  height  result  and  heavier  2 o  T h i s may mean t h a t  was t h a t  that and  this  species  relative.  vigor  inhabit  0  This  factors.  varied in  The a v e r a g e w e i g h t  with microsite.  The w e i g h t  p h o t o s y n t h e s i s , and t h e r e f o r e  to follow  t h e same t r e n d  fact  zone.: i n t h e and n u t r i -  seedlings  of needles i s of importance  might  as height  of  depth  demon-  capacity  of needles  height  systems.  root  t h e most f a v o r a b l e  growing media i n terms o f m o i s t u r e - h o l d i n g tive  indicates  microsites  of root  had a g r e a t e r  t h a n t h o s e w h i c h grew i n d e c a y i n g w o o d the roots  study  typically  as e x p r e s s e d by  with depth  growing i n m i n e r a l s o i l  strates that  t o be a  of t h i s  seedlings  of roots*  v a r i a b l e between  Seedling  was n o t f o u n d t o b e a s s o c i a t e d Seedlings  established  g r o w i n g on M i c r o s i t e  considered  The r e s u l t  0  shallowness i s quite  i s only  3 had a b e t t e r  w i t h t h e weight  Western hemlock i s u s u a l l y shallow-rooted  growing i n w e l l -  t h e t o p development o f young  associated  more  originating naturallyo  seedlings  system than s e e d l i n g s  not n e c e s s a r i l y  I ti s  0  b e c a u s e t h e y may have r e c e i v e d  a d v a n c e d d e c a y i n g wood on M i c r o s i t e  is  indicate  1 and 4 had t h e b e s t  i n the nursery than seedlings  root  of roots  and d i a m e t e r s  on M i c r o s i t e s  of roots  Another s t r i k i n g  Average  r e l a t i o n s h i p e x i s t s between t h e v a r i o u s  than f o r t o t a l  reasoned that  2 .  on  1 to Microsite 5 o  data f o r average oven-dry weight  a different  regeneration  Of c o u r s e b e e x p e c t e d  g r o w t h and d i a m e t e r  growth.  - 37  -  These trends appear to be useful i n evaluating s i t e influences© i f the l i g h t factor remains constant.  Root/shoot r a t i o  was  the highest f o r the planted seedlings on Microsites 1 and 4« Seedlings on Microsite 2 had the second best height and diameter growth, but the lowest root/shoot r a t i o , fact indicates that the root/shoot r a t i o i s of l e s s  \This importance  i n natural w i l d l i n g s , than i n nursery stock). Moreover, the r a t i o i s not constant but changes from microsite to micros i t e and i s not connected with height or diameter growth. Number of branches on seedlings increased with microsite quality.  The average number of branches appears to be a good  basis to evaluate the vigor and growth p o t e n t i a l of young seedlings.  The average number of branches per internode  followed the same trend.  Total number of branches per i n t e r -  node increased with s i t e q u a l i t y .  The t o t a l number of branches  on the f i r s t internode on Microsite 1 was less than on the other microsites. The reason f o r t h i s i s that the seedlings on Microsite 1 were planted too deep, k i l l i n g branches.  the lower  It i s recommended that i n further studies i n t h i s  f i e l d the average number of branches on the l a s t internode offers a convenient measure to estimate the r e l a t i v e vigor (based on t o t a l height) of seedlings growing on d i f f e r e n t micrositeso The average branch angle of internodes decreased with s i t e quality.  Branch angles on the f i r s t , second, and t h i r d i n t e r -  node showed generally s i m i l a r tendencies.  Branch-angle  ferences on the t h i r d internode ( t o t a l age 3)  dif-  were evaluated  -  statistically ing  on d i f f e r e n t  branch angles  19)  (Table  found t h a t  seedlings  m i c r o s i t e s have s i g n i f i c a n t l y  different  geneous p o p u l a t i o n  angles  a f t e r planting„  highly  significant  i t was  o r i g i n a t i n g from a reasonably  i n the  c h a r a c t e r i s t i c s before  nursery  and  p l a n t i n g developed d i s s i m i l a r Seedlings  This  The first  height  average l e n g t h  internode  length  on  was  characteristic  decreasing  dent  on  that  the  the  had  seedlings  of hemlock  seed-  q u a l i t y beside  of l a r g e s t n e e d l e s  unrelated  the  of b r a n c h e s on  to micrositeso the  internode.  The  second i n t e r n o d e  microsite quality.  third  branch  growth.  o f l a r g e s t n e e d l e s on  with  uniform  g r o w i n g on M i c r o s i t e 1  a p p e a r s t o o f f e r a measure o f s i t e  e v a l u a t i o n based  grow-  homo-  presumably with  d i f f e r e n c e s i n branch angle from  grown on M i c r o s i t e 4« lings  and  Seedlings  0  -  38  The  This trend statistical  was  the  average decreased most  evi-  a n a l y s i s proved  d i f f e r e n c e s i n n e e d l e l e n g t h were s i g n i f i c a n t  between  micrositeso The  average base diameter  s e c o n d , and with had  third  decreasing an  only  than those The  internode  o f b r a n c h e s on  showed a t e n d e n c y t o  microsite quality.  slightly  higher  on M i c r o s i t e  Seedlings  first, decrease  of  branches  4.  gradually  of s e e d l i n g s  g r o w i n g on  from M i c r o s i t e 1  various  to Microsite  5 i n d i c a t i n g a c l o s e r e l a t i o n s h i p between b a r k t h i c k n e s s microsite quality. p r e s u m a b l y had  5  on M i c r o s i t e  average base diameter  average bark t h i c k n e s s  m i c r o s i t e s decreased  the  Seedlings  g r o w i n g on M i c r o s i t e s 1  s i m i l a r bark thicknesses  before  and  and  planting  4  but  -  one  -  3 9  year a f t e r p l a n t i n g the average bark t h i c k n e s s  l i n g s g r o w i n g on M i c r o s i t e 4 d e c r e a s e d t o M i c r o s i t e 1. importance  THE  1959 for  o f hemlock  decrease i n d i c a t e s the apparent  sites  hemlock, i n c r e a s e d  ences i n height  seedlings„  ranging with  from s i t e  site  index.  Considerable  however, i n comparisons o f h e i g h t on P l o t s 1 and 4 r e p r e s e n t i n g  i n April  i n d i c e s 79 t o 142 Only s l i g h t  growth e x i s t e d between s e e d l i n g s  1 and 5 r e s p e c t i v e l y .  differ-  on P l o t s  d i f f e r e n c e s were  g r o w t h between  evident,  seedlings  t h e extremes of s i t e  index.  r e s u l t s o f p r e l i m i n a r y work on t h e d i f f e r e n c e s o f  external c h a r a c t e r i s t i c s revealed  isons  on t h e e x t e r n a l  g r o w t h i n 1959 o f w i l d l i n g s p l a n t e d  on d i f f e r e n t  with  compared  INFLUENCE OF S I T E ON HEMLOCK SEEDLINGS.  height  The  4 7 p e r cent  o f m i c r o s i t e a s an i n f l u e n c e  characteristics  The  This great  of seed-  site  quality.  that branch angle  decreased  T h e s e r e s u l t s were s u p p o r t e d b y compar-  o f measurements o f t h e a v e r a g e b r a n c h a n g l e  of t h e I960 g r o w i n g s e a s o n  on d i f f e r e n t  sites  of shoots  (Table 22).  T h e s e c o m p a r i s o n s r e v e a l marked c h a n g e s i n s e e d l i n g  morphology.  A l s o many c h a n g e s become r e a d i l y a p p a r e n t w i t h i n two after  planting.  These c o n c l u s i o n s  portant  and a r e s u p p o r t e d  tically  significant  ence exceeded in  years  a r e b e l i e v e d q u i t e im-  statistically.  They a r e s t a t i s -  i n a l l c a s e s where t h e s i t e - i n d e x  s i xf e e t .  Such r a p i d changes i n b r a n c h  p a r t i c u l a r , were u n e x p e c t e d b e c a u s e s e v e r a l a u t h o r s  1957,  Zobel  height,  19b0) r e p o r t e d  stem t a p e r ,  that  straightness  the h e r i t a b i l i t y of b o l e ,  crown  differangle, (Toda  of t r e e diameter,  bark t h i c k n e s s ,  branch angle,  n o t e d however t h a t fically  40  these  -  were a l l h i g h .  I t should  r e p o r t s were n e i t h e r d i r e c t e d  t o w e s t e r n hemlock nor  generally  by  weight  the  fact  of branches.  This p o s s i b i l i t y  that branch length  and  (I9b0j stated  i f small  speci-  to seedlings.  change i n b r a n c h a n g l e i s p o s s i b l y a r e f l e c t i o n erent  be  The  of the  diff-  i s strengthened  needle length  increased  with s i t e o Mergen vironment  are  phenotype the It  was  that  r e f l e c t e d by trait  pronounced m o d i f i c a t i o n  i s under l o o s e  f a r beyond the  aim  genetic  of t h i s  statements about g e n e t i c  f a c t o r s and  of w e s t e r n hemlock.  fact  The  that  branch a n g l e w i t h changes i n s i t e tion  that  the  e n v i r o n m e n t may  hemlock t h a n i n o t h e r external  The  species,  length  of the  sites  the  t o the  assump-  or the  heritability  of  some  stage  Using the  on  significant  the  last  year's  c o r r e l a t i o n with  xndex d i f f e r e n c e s exceeded s i x f e e t .  emphasized  b r a n c h a n g l e i t was  o f hemlock s e e d l i n g s  the  importance  equations based  on  of  growing  on  environmental  needle length  p o s s i b l e t o estimate the  i n d i c e s f o r M i c r o s i t e s 1 to 5 The  respect  changed  q u a l i t y leads  l a r g e s t needles  change o f n e e d l e l e n g t h  factorso  in  stages,  i n d e x , where s x t e  various  0  support  heritability  seedlings  the  have more i n f l u e n c e i n w e s t e r n  shoots a l s o i n d i c a t e d a h i g h l y sxte  control  thesis to  en-  in  c h a r a c t e r i s t i c s i s l e s s marked i n j u v e n i l e  than i n l a t e r The  changes i n the  unknown  and site  a  p o s s i b l e a p p l i c a t i o n of t h e s e r e s u l t s t o s i t e  index  - 41 determination require confined no  by  using  seedlings  further investigation. t o the  attempt  material  is justified  study t o t r e e s  older than f o u r years The  scope of t h i s  described.  o l d e r than f o u r  years.  study  I t i s emphasized  here t o p r o j e c t the  would  r e s u l t s of  is  that this  - 42 C O N C L U S I O N S THE INFLUENCE OF LIGHT INTENSITY ON HEMLOCK S E E D L I N G S The  height  lated  to light  height  within  influence  growth o f hemlock s e e d l i n g s intensityo  Some d i f f e r e n c e s  was d i r e c t l y r e i n age a t b r e a s t  p l o t s o n s i m i l a r s i t e s were a t t r i b u t e d t o t h e  of l i g h t  intensity.  Diameters a t i n t e r v a l s from t h e root m i n a l s h o o t were a l s o i n f l u e n c e d tensity.  e  Form q u o t i e n t  collar  t o the t e r -  by v a r i a t i o n o f l i g h t i n -  was i n v e r s e l y  related t o light i n -  tensity. The  a v e r a g e number o f g r e e n b r a n c h e s p e r s e e d l i n g i n -  creased with l i g h t dent on  intensityo  on s i x - y e a r - o l d  11-year-old  of n a t u r a l  Natural  seedlings.  seedlings  0  p r u n i n g was n o t e v -  Natural  p r u n i n g had s t a r t e d  On 1 1 - y e a r - o l d  seedlings  p r u n i n g was i n v e r s e l y r e l a t e d t o l i g h t  the rate  intensity.  THE E F F E C T OF MICROSITE ON THE EXTERNAL CHARACTERISTICS OF THREE-YEAR-OLD HEMLOCK SEEDLINGS., The  height  was i n f l u e n c e d ditions  growth o f t h r e e - y e a r - o l d by m i c r o s i t e s  resulted  year a f t e r planting.  had  better  Differences observed The ment  significant  Seedlings  growing  growth than s e e d l i n g s  con-  differences,  on d e c a y i n g  i n mineral  wood  sub-soil©  observed i n d i a m e t e r growth a r e s i m i l a r t o t h o s e  f o r height. average weight  o f young hemlock The  seedlings  D i f f e r e n t environmental  in statistically  one  height  0  hemlock  d i dnot r e f l e c t  the develop-  seedlings.  depth of roots  with height  of roots  was v a r i a b l e , b u t was n o t c o r r e l a t e d  growth.  Microsite  quality  (based  on t o t a l  h e i g h t ) and weight o f  - 43 needles  -  o f s e e d l i n g s were c l o s e l y  Root/shoot quality«  The  r a t i o was  not a s s o c i a t e d  average t o t a l  increase i n site  quality,  correlated.  number  and  ,  with  microsite  of b r a n c h e s  increased  t h e same t r e n d was  observed  b e t w e e n number o f b r a n c h e s p e r i n t e r n o d e on v a r i o u s The  Branch-angle d i f f e r e n c e s  i n t e r n o d e were s i g n i f i c a n t f o r only  teristic  microsites  average branch angle per i n t e r n o d e decreased with  m i c r o s i t e quality,.  ing  with  on t h e  third  e v e n when s e e d l i n g s had b e e n growmicrositeso  one y e a r on d i f f e r e n t  of hemlock s e e d l i n g s a p p e a r s t o o f f e r  This  charac-  a measure o f  qualityo  site  The third  length  of the l a r g e s t  i n t e r n o d e was  closely  n e e d l e s on t h e s e c o n d  correlated  and  with microsite  diff-  erences,, The  average base diameter  t o decrease with microsite The  THE  growth,  local  o f s e e d l i n g s was  INFLUENCE OF S I T E environment  had  had  These  cal  site,  correlated  HEMLOCK S E E D L I N G S  d i f f e r e n c e s were n o t  b r a n c h a n g l e , and  highly significant  based  ON  a significant  i n d e x d i f f e r e n c e d i d not exceed The  tendency  quality«  b r a n c h a n g l e , and n e e d l e l e n g t h  seedlings.  showed a  quality„  average bark t h i c k n e s s  with microsite  The  of branches  of western  estimate s i t e  length The  index,,  height  hemlock site-  0  of l a r g e s t calculated  on n e e d l e l e n g t h and b r a n c h a n g l e were n e a r l y  and b o t h c o u l d  on  o b s e r v e d when t h e  six feet  correlation,,  effect  e  needles equations identi-  0  - 44 Table 1  O c c u r r e n c e or' G r o u n d V e g e t a t i o n on t h e T h r e e Microsites used t o r S t u d y or' t h e e f f e c t o f M i c r o s i t e s on t h e E x t e r n a l C h a r a c t e r i s t i c s o f Hemlock S e e d l i n g s *  0  0  Microsite  Acer circinatum Agrostis  alba  Pursh  L  0  e  Anaphalis margaritacea Alnus rubra  X  X  X  X  X  X  f e m i n a (JL») R o t h  Blechnum s p i c a n t  commutata  ( L ) Scop 0  Cornus c a n a d e n s i s L shallon  IRegelj  Fern  0  Var.  0  e  Pursho  angustifoiium L  X  ffl  Linnaea b o r e a l i s L, P o l y t r i c h u m jjuniperum Hedw  0  P o l y s t i c h u m muni turn ( K a u l f o ) Populus t r i c h o c a r p a  Torr  0  Underw  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  0  & Gray  X  Douglo  X  Rubus s p e c t a b i l i s  Pursho  X  X  X  X  Cham  & Sch  0  0  S a l i x Hookeriana Barr, Saiix  sitchensis  Sambucus  Spiraea douglasii  Thuja p l x c a t a  X  Sanson  pubescens  Taxus b r e v i f o l i a  X  X  Rubus l e u c o d e r m u s Rubus v i t i f o l i u s  5  X  Betula papyriphera  Epilobium  e  2  Bong  Athyrium f i l i x  Gaultheria  L  1  Number  X  (Michx)  X  Hook »  X  Nutto  Donn  Tsuga h e t e r o p h y l l a  X X  X  X  X  X  X  X  X  a  I R a f o ) Sarg»  Vaccinium p a r v i f o l i u m  X  Smith  * P l a n t s were i d e n t i f i e d  by L  B  0rloczy  o  -  Table 2 .  45  -  P h y s i c a l C h a r a c t e r i s t i c s of the F i v e P l o t s used f o r Study of the E f f e c t of S i t e on Hemlock S e e d l i n g s ,  P L  Characteristics  0  N U M B E R  T  1  5  7  Site Index - Feet ( f o r hemlock) 2  142  128  122  E l e v a t i o n - Feet  690  1,000  1,080  Aspect  SW  Slope - Per cent  Source:  2.  13  SW 12  SW 10  8  115  4  79  1,100  1,370  SE  SE 8  12  G r i f f i t h B.G. 1900. Growth o f Douglas F i r a t the U n i v e r s i t y of B r i t i s h Columbia Research F o r e s t as R e l a t e d t o Climate and S o i l .  S i t e index was c a l c u l a t e d f o r hemlock by u s i n g the e q u a t i o n developed by J.H.G. Smith Hemlock SI » 22.34 + 0.663 S I of Douglas F i r (SE + l(>.4«) E ~  - 46  Table 3.  Depths o f S o i l H o r i z o n s and Tree Root P e n e t r a t i o n of F i v e P l o t s used f o r Study of the E f f e c t of S i t e on Hemlock Seedlings.*-  Plot  Average depth of h o r i z o n s (inches)  Depth t o Bedrock (inches)  No. "A"  1.  -  Max.  Min. Ave.  Root p e n t r a t i o n (inches) L a y e r of main conMax. centration  1  3  22  27  60  30  50  28  6-10  5  5  19  18  54  32  37  48  7-16  7  3  19  11  46  22  30  38  2-10  8  4  18  18  42  30  36  38  2-10  4  4  16  5  36  2  20  34  2-10  Source:  G r i f f i t h B.G. I960. Growth o f Douglas F i r at the U n i v e r s i t y of B r i t i s h Columbia Research F o r e s t as r e l a t e d t o C l i m a t e and Soil.  - 47 T a b l e 4»  O c c u r r e n c e o f G r o u n d V e g e t a t i o n on t h e F i v e P l o t s u s e d f o r S t u d y o f t h e E f f e c t o f S i t e on Hemlock Seedlings e  PLOT 1 Acer Circinatum Pursh X Achlys t r i p h y l l a D C X Blechnum s p i c a n t ( L . ) S c o p X ChimaphiJLa__ u m b e l l a t a (L „) N u t t „ Cornus c a n a d e n s i s L Galium b o r e a l e L X G a u l t h e r i a s h a l l o n Pursh« G o o d y e r a m e n z i e s i i Lindlc Gymnocarpium d r y o p t e r i s L« Linnaea b o r e a l i s L Mahonis a q u i f o l i u m f u r s h X Menziesia f e r r u g i n e a Smith Monotropa u n i f l o r a L O p l o p a n a x l v H o r r i d u s ( S m i t h ) B. & Ho  N  U M B E R  5  7  8  X  X  X X  X  X X X X  X  X  4  0  0  0  e  0  X  e  0  X  X X  6  0  P o l y s t i c h u m munitum ( K a u l f . ) Underw» Pteris aquilina L P t e r o s p o r a andromedea N u t t Rubus p e d a t u s S m i t h Rubus s p e c t a b i l i s Pursho Sambucus p u b e s c e n s M i c h x Streptopus amplexifolius D C Tiarella trifoliata L T r i e n t a l i s l a t i f o l i a Hooka T r i l l i u m ovaturn Pursh« V a c c i n i u m membranaceum D o u g l V a c c i n i u m o v a l i f o l i u m Smith Vaccinium p a r v i f o l i u m Smith Viola g l a b e l l a Nutt.  X  X X  X  X X  X X  X X X  X  X  X  X X X  X X X  0  X  e  D  8  0  c  1.  Source*  X X  X  X 0  X X X  X X X  X  X X  Griffith B G 1960 Growth o f D o u g l a s F i r a t the U n i v e r s i t y o f B r i t i s h Columbia R e s e a r c h F o r e s t a s r e l a t e d t o C l i m a t e and S o i l . 0  o  0  Table  5o  The i n f l u e n c e Light  i n t e n s i t y data year  P l o t V, and . Plot 5  Control  00 00 00 00 00 00 00 00 00 00 00 00  7500 9000 11250  16 24 60 96 120 106  Total  Time o f Observation 07  08 0y 10 11 12 13 14 15 16 17 18  11700 13500  12900 12900 2100  of l i g h t  Plot  70  I.  II.  32  90  34  47 125 70  125 100 180 100 125  96 100 190 110 IbQ  21150  50 240 110  3600  32  290  114450  1024  1408  7846  13950 3900  85*33  Average 9537.50 Average relative light i n tensity 100*0 percent 2  0.89  1, 2, +  on a b r i g h t d a y f o r 1 1 - a n d 6o l d seedlings.  0 XXX o  P L  250 750 250 1500 3900 450  100  i n t e n s i t y on hemlock s e e d l i n g s  180  110 125  T  N U M B E R IV.  125 160  370 230  400  260  1100 7800 1200 980 500  400 375 820 1500 9000 7500 500 550  13605  21590  190 250 750  Date of Observation J u l y 9, I960,  1 28 34  70 125 250  190 120 180 110 125  2  3  50 84  90 110 125  500 130  260  60 195 190 375  130 520 375  725  180  180 200 260 220  750 1500 500 375  1562  2373  5460  150  117o33 653*83 1113*75 1799*16 130*16 197*75 455*0  1*23  6.85  11,89  18*86  1.36  2*07  4*77  M e a s u r e m e n t s were t a k e n i n f o o t - c a n d l e s , C a l c u l a t i o n of t h e average r e l a t i v e l i g h t - i n t e n s i t y p e r c e n t d e s c r i b e d on Page 34 o F o o t c a n d l e v a l u e s were i d e n t i c a l f o r b o t h p l o t s .  4 125  130  165 340 250 500  750 1200 7500 7500  600 450 19520 1626*66  17*06  Table 6 .  The i n f l u e n c e Light  Time o f Observation  07 00 00  08  oy oo  10 00 11 00 12 00 13 o o 14 00 15 00 l b 00 17 o o  l b 00 Total Average  F l o t V, and ^ Plot 5  i n t e n s i t y data on a cloudy day (completg f o r 11-and 6-year-old hemlock s e e d l i n g s .  Control  700  30  48 64 90  125 90  94  it>50  1800 iy20 1800 850 185 13935  0  +  90 170  110  220 155 230 150  500 350 450  170  290  275  iyo  120  200  885 73.75  1895 157.91  4385 365.41  b.35  13o59  3lo4b  lb  1 2,  II,  195 155  94 bO  Average relative light i n tensityg 100 percent  I*  550 550 550 500  110 b4  llbl.25  P L0 T  plot  yoo  y«o 1050 1100 1000  of l i g h t i n t e n s i t y oh hemlock s e e d l i n g s .  50  bo  overcast) Date of Observation September 1, I960,  N U MB E R III. 170 275  490 550  575 650 725 800 870  725  390 110  6330  IV.  280 375  600 725 800  750 950 950 950 900 450 120  7850  527*50 654ol6  45*42  56.33  1  64  90 100 130  200  140  125 200 125  125 100  2  3  4  92  170  280 375  150  200  300 400  250  375 400 375 375 190  250 470 520  575 600 725 725 725 700 375  b0 94 3167 5929 119.91 263.91 494*08 40  1439  10„32  22.72  730  800 750  950 1000 870 900 425 110 7790  55o90  42.54 55.90  Measurements were taken i n f o o t candles© C a l c u l a t i o n o f the average r e l a t i v e l i g h t i n t e n s i t y percent d e s c r i b e d on Page 34. Foot candle v a l u e s were i d e n t i c a l f o r both p l o t s s  600  Table 7 °  The i n f l u e n c e of l i g h t i n t e n s i t y on hemlock  seedlings  0  Some e x t e r n a l c h a r a c t e r i s t i c s of 11-year-old hemlock s e e d l i n g s on two p l o t s o f v a r i o u s degrees of shade? Flot Characteristics No. of 10 s e e d l i n g s measured on each plot Average length(mm) Average middle diameter {mm) Average form quotient Average number of green branches Average number of dead branches  I N T E R N O D E  1  2  3  4  5  6  N U M B E R  7  8  10  9  Q2  9.6 11 5  2 34  2.28 2.17  2 . 0 3 1.91 1 . 8 1 1 . 7 0 1.51  Q 96  0.98 0.97  0.9b 0.9b 0.95 0.94  0  e  0  0  -  Total  13.6 16.3 2 0 . 1 26.8 26.7 31«7 5 7 . 5 8 4 . 8 307.8  - 0 . 1  11  0„6 1.1  -  -  0  .  0 92 0*92 o  0.8 l 1  -  lo29  o  0 1.2  -  -  lo05 0.75 0.88  0.81  2 . 8 1.1 -  -  8 b Q  0 2 o  Average length(mm) 2 5 . 2 3 0 . 9 33e0 41«4 5 3 . 1 47o8 80.2 8 8 . 7 9 7 . 7 183.8 169.2 851«0J diameter  (nun)  Average form quotient  Average number of green branches Average number of dead branches  7 . 0 5 6 . 8 9 6 . 4 1 5*90  5.67  0.94 0.98 0.97 0.96 0 96 o  -  0 . 4 1*3  l o 5 2.0  0.4 0.2 0.2 0.3 0.3  5 . 0 7 4o40 3<>69 2 . 9 8 2 . 0 9  1.11  0.95 0.94 0.91 0 89 0.88 0.75 o  2.5 -  3o4 4 o l 4o2 1 0 . 5 -  °«1  -  -  4o6 34o -  l.i  - 51 H O C8 e +3 •O 0 ON H H H  H  N  ©  •O Ni  o  CQ W  a  •H  H  •o  CD 0) CQ  J* 0 0 iH  CD 0 TJ 0 (3  H  O H  >o*  ON  00  o  o  NO CN  00 00  O.  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CD &t CS U U CS CD CD - r i CD 3 CD U CD CD CD CD •rl CD 3 CD t, CD CD CD CD - r i CD 3 CD U CD CD > > TJ > cr > > TJ > >cr > bD > TJ> > TJ >cr > bO > TJ  *> s 0 9 H »  CO  •rl  0»  JC  <<  o  CM 0  <  <  CQ CD u CD x ; CD X) O XJ a fi s 3 CO  <  g g  O H  o  CN  o  CO  t  o  ©  o  FH  <<  -3-  0  <  <  <  << to  <  <  <  Table 9o  The i n f l u e n c e of l i g h t i n t e n s i t y on hemlock s e e d l i n g s  0  Some e x t e r n a l c h a r a c t e r i s t i c s of s i x - y e a r - o l d hemlock s e e d l i n g s on two p l o t s of v a r i o u s degrees of shade.  Plot No, XQ  II.  C h a r a c t e r i s t i c s of 10 s e e d l i n g s measured on each p l o t  I N T E R N O D E  1  Average l e n g t h (mm) Average middle diameter (mm) Average form q u o t i e n t Average number of green branches Average number of dead branches  9.7  Average l e n g t h (mm) Average middle diameter (mm) Average form q u o t i e n t Average number of green branches Average number of dead branches  20,5  1.3  0.95  -  2  3  10.0  16.0  N  4 27.5  U M B E R  5 36.1  6  Total  33.8  133ol  1.20  1.04  0.91  0.85 0.85  0.66  0.50  0.1  0.3  3. o 1  1.4  0.2  3.1  47*6  41.5  207.5  0.98  20.5  -  30.9  -  46o5  0.82  -  0.82  -  1.79  0.87  1.54 0.93  1.36  0.92  1.15 0.89  0.81  0.86  0.59 0.76  Ool  0.8  1.6  2.6  2.6  0.3  8.0  Table 10.  The i n f l u e n c e of l i g h t i n t e n s i t y on hemlock s e e d l i n g s Some e x t e r n a l c h a r a c t e r i s t i c s o f s i x - y e a r - o l d hemlock s e e d l i n g s on t h r e e p l o t s of v a r i o u s degrees of shade.  Plot No  0  III.  IV.  V.  C h a r a c t e r i s t i c s of 10 s e e d l i n g s measured on each p l o t  INT  1  Average l e n g t h (mm) Average middle diameter (mm) Average form q u o t i e n t Average number of green branches Average number of dead branches  20.1  2 19.6  E R N0 D E  3 31e0  NUMB  4 110.7  E R  6  5 118,5  Total  80.0  2.83 0.94  2.62 0.96  2.46 0.96  2.01 0.86  1.25 0,79  0o7 0o78  -  0.3  1.7  6.4  7.5  0.7  0.1  0.2  -  -  -  Average l e n g t h (mm) Average middle diameter (mm) Average form q u o t i e n t Average number of green branches Average number of dead branches  28,8  24.1  Average l e n g t h (mm) Average middle diameter (mm) Average form q u o t i e n t Average number of green branches Average number of dead branches  80.5 148.0  31.8  173.0  6.61 0.92  6.02 0.95  5.72 0.97  5.0 0.91  0.7  0.8  1.9  9.6  „  =>  -  -  23ol 17.8 0.90 0.85 6.5  10.0  122.5  193.0  16.4 0.93  13.1  15.0  10.5  0.89  293.0 3.59 0.82 16.2 -  414.0 8.95 0.80 26.0 —  318.4  379«9  16.6  0.3 869.1  1.8 0,78 11.8  41.0  659.5  1617,5  -  4.35 0.68 21.5 —  89.5  Table 11.  Some e x t e r n a l c h a r a c t e r i s t i c s of 10 t h r e e - y e a r - o l d hemlock s e e d l i n g s on M i c r o s i t e 1©  Characteristics T o t a l height (mm) Base diameter(mm) Average l e n g t h of l a r g e s t needles on f i r s t internode Average l e n g t h of l a r g e s t needles on second i n t e r n o d e Average l e n g t h of l a r g e s t needles on t h i r d internode T o t a l number of branches Weight of r o o t s (grams) Weight of needles (grams) Root/shoot r a t i o Depth of roots(mm) Double bark t h i c k ness at the middle diameter of f i r s t internode (mm) Double bark t h i c k ness a t the middle diameter of second internode (mm) Double bark t h i c k ness a t the middle diameter of t h i r d internode (mm)  10,7  11.5  N U M B E R 8 7 642 725 834 14.0 10.0 10.9  19.9  16,9  15.3  11. 8  13.5  15.0  23.1  17.7  14.9  16.6  14.5  13.93 15.16  13.37  11.83  12.54  46  30  31  39  15.6  10.6  10.5 13.2  S E E 1  2  691 10o3  767 12.0  13.6  16.1  16.0  17©4  13.36 46 8.4  13.4  0„30 142  3 73b 11.9  20,6  16,6 0.41 0 , 3 0 146 141  D L I N G 4  681  0.38 182  5 756  6  9  10 841  Average  11.4  746.2 11.4  13.2  13.8  14.9  20.5  14.9  16.1  17.2  13.26  15.4  14.10  14.87  13.7  37  33  36  44  42  38.4  14,6  24.0  10.7  11.5  10.5  10.6  12.7  18,4  25.0 0.46 201  13.7  12.6 11.4 0.39 0o5l 192 595  13.5  15.8 0.38 220.6  0.38 165  0.37 202  787  11.7  O.36 240  1©9  XQ X  lo7  1.1  1.3  1.7  Xe2  1.4  1.4  1.4  1.42  lo3  0.9  X0 1  1.1  1.2  1.4  X <? 2  1.2  1.3  1.1  1.18  1*1  0,5  0.6  0.6  0.5  0.7  0.6  0.6  0.3  0.6  0.61  T a b l e 12o  Some e x t e r n a l c h a r a c t e r i s t i c s o f 10 t h r e e - y e a r - o l d hemlock s e e d l i n g s on M i c r o s i t e 2.  Characteristics Total height (mm) Base diameter(mm) Average l e n g t h of l a r g e s t n e e d l e s on f i r s t internode Average l e n g t h of l a r g e s t n e e d l e s on second i n t e r n o d e Average l e n g t h of l a r g e s t n e e d l e s on t h i r d internode T o t a l number o f branches Weight of r o o t s (grams) Weight of n e e d l e s (grams) Root/shoot r a t i o D e p t h o f roots(mm) Double bark t h i c k ness at the middle diameter of f i r s t internode (mm) Double bark t h i c k ness at the middle diameter of second internode (mm) Double bark t h i c k ness at the middle diameter of t h i r d intejraode (mm)  1 8 1 5  2 3  704  3  S  E E D L I  8 5 9  4  8 0 8  5  N V M B E R  N G  7  6  6 7 8  7 6 4  1 0 . 1  7.0  6 3 3  9  8  7 5 3  7.4  10  Average  6 3 4  6 8 4  7 3 6 . 2  6,4  7.1  8.0  9.9  8.5  7.6  19*6  1 7 o 8  17.0  1 3 . 2  15.7  2 0 . 8  17.0  1 5 . 5  1 9 . 8  1 6 . 8  1 7 . 3  1 8 . 2  15.5  1 8 . 1  14.5  17.4  1 9 . 3  1 6 . 8  1 9 . 0  19*2  17.4  1 7 . 5  12.7  1 2 . 4  9.8  1 3 . 8  10.3  12.3  13.3  1 3 . 1  1 2 . 2  4 9  2 6  4 5  39.0  1 1 c  4 7  40  8.7  5.1  2 3 . 6  1 3 . 9  0.21 1 3 5  0.18  107  3.X  0  3  5 . 6  4 5  3 6  40  31  3 1  4.0  2.8  4.7  2.7  1.5  2.3  2.1  5.4  10.7  7.4  4.7  6.2  4.3  0.17  0.17  0.23  1 6 5  2 4 2  2 1 2  222  13.0 0.17 1 4 6  0.20  144  0.19  204  0.21  3.6  1 0 . 3 0.20 2 1 0  3.7 9 . 9 4  0.19  1 7 8 . 7  0.6  0.8  0.9  0.7  1.4  0 . 8  0.6  0.8  1.0  1.0  0.86  0.5  0.8  0.5  0.5  1.2  0.5  0.9  0.8  0.9  0.9  0.75  0.4  0.4  0.6  0.3  0.5  0.3  0.5  0.4  0.8  0.5  0.47  Table 1 3 o  Some e x t e r n a l c h a r a c t e r i s t i c s o f 1 0 t h r e e - y e a r - o l d hemlock s e e d l i n g s on M i c r o s i t e 3 «  Characteristics Total height (mm) Base diameter(mm) Average l e n g t h of l a r g e s t n e e d l e s on f i r s t internode Average l e n g t h of l a r g e s t n e e d l e s on second i n t e r n o d e Average l e n g t h of l a r g e s t n e e d l e s on t h i r d internode T o t a l number o f branches Weight of r o o t s (grams) Weight o f n e e d l e s (grams) Root/shoot r a t i o D e p t h o f roots(mm) Double bark t h i c k ness a t the middle d i a m e t e r of f i r s t internode (mm) Double bark t h i c k ness a t the middle diameter of second internode (mm) Double bark t h i c k ness at the middle diameter of t h i r d internode (mm)  S E E 1  2  D L I N G 4  3  5  6  46b 6.3  457 5.5  514 5.9  617  783 6.8  611  6.0  20.4  11.5  14.9  16.4  21.5  14.3  17.6  11.3  14.3  17  23  2.8  2.5  3.6  2.4  7.3 0.22  4.4  5.0 0.40 115  0.20  95  0.31 91  0.9  0.9  0.5  0.6  N U M B E R 8 7  9  10  Average  840 9.8  620.9 7.18  527  5.4  757 9.4  17.2  16.5  16.2  14.7  17.1  16.4  16.1  17.4  13.4  14.6  16.8  15.4  18 o 3  18.3  16.8  10.7  10.6  10.6  11.6  12.1  10.3  12,1  10.4  11.4  30  35  42  49  40  27  35  43  34.1  4.3  3.7  4.3  3.5  7.1  7.9  8.6  0.26  6.7 0.30  4.7  6.3  105  51  1.1  0.8  0.6  0.9  0.4  0.6  8.0  637 8.7  4.2  5.5  5.0  0.32  245  0.40 227  0.26 85  0.9  0.8  1.3  0.8  0.9  1.5  0.99  0.6  0.7  0.7  1.0  0.7  1.2  0.8  0.77  0.4  0.7  0.5  0.6  0,5  0.6  0.9  0.58  6.5  151  0.17  165  8.0 0.28 133.0  Table  14.  Some e x t e r n a l  S  Characteristics Total height (mm) B a s e diameter(mm) Average l e n g t h of l a r g e s t n e e d l e s on first internode Average l e n g t h of l a r g e s t n e e d l e s on second i n t e r n o d e Average l e n g t h of l a r g e s t n e e d l e s on t h i r d internode T o t a l number o f branches Weight of r o o t s (grams) Weight of n e e d l e s (grams) Root/shoot r a t i o D e p t h o f roots(mm) Double t h i c k n e s s of m i d d l e d i a m e t e r of f i r s t internode(mm) )ouble bark t h i c k less at the middle i i a m e t e r of second Internode (mm) )ouble bark t h i c k l e s s at the middle i i a m e t e r of t h i r d Internode (mm)  characteristics  1  2  E E D  3 530 4.1  o f 10 t h r e e - y e a r - o l d on M i c r o s i t e 4o  L I N G 4 5  616  535 6.4  360  13*5  13.0  XX o 7  11.4  13.2  lbo4  13.2  13.1  15.1  17.7  12e4  12 © 1  10.8  10.4  25  28  30  35  5.4  6.0  569 7.5  9.7 40  hemlock  seedlings  N U 6 476 6.6  M BER 8 7 552 521  9  io  593 7.7  1  Average 522.7 6.5  6.8  8.2  475 6.6  13.2  12.0  14.7  12.4  13.3  12.8  14.2  16.3  18.1  14.6  16.5  15.5  7.9  7.8  26  29  8.2 25  X2© X 32  9.7 36  10.1 30.6  5.3  3.2  2.0  4.7  5.6  3.0  3.1  5.7  3.6  7.0  5.4 0.43  3.6  0.48  2.7  0.36  6.6 0.43  3.7 0.35  3.7 0.43 95  6.7 0.40  3.9 0.46 145  7.5 0.46  130  4.7 0.47 105  0.9  0.6  0.8  0.6  0.9  0.8  0.5  0.9  0.8  0.7  0.75  0.8  0.4  0.7  0.5  0.7  0.6  0.5  0.7  0.6  0.3  0.58  0.4  0.2  0.2  0.3  0.5  0.4  0.4  0.6  0.5  0.2  0.37  120  105  160  132  112  100  4.3 4.9 0.43  120.4  Table 15o  Some external c h a r a c t e r i s t i c s of 1 0 three=year-old hemlock seedling on Microsite 5o S E E D L I NG 2  1  T o t a l height(mm) 44b Base diameter(mm) 7„2 Average length of l a r g e s t needles on f i r s t internode 13.b Average length of largest needles  500  bob 12  0  N U M B E R  3  4  5  6  485  448 4.3  511 5.4  537 7.1  441 5.2  lb.1  lb.0  14.4  b.8  3 lbo4  14.5  7  8  9  39b  581 5.7  on second i n t e r n o d e l 5 » 9 13o3 13.8 15.8 lb . 2 14.b 1 5 . 7 Average length of largest needles on t h i r d internode 8 . 2 7 7.29 7*42 7 . 5 4 7.54 8.47 8.54 T o t a l number of branches 2b 38 29 24 37 37 31 Weight of roots (grams) 2 9 W01 4.9 2o7 4.5 1.4 1.8 Weight of needles 7.0 2.8 (grams) 4.1 9.5 4.6 b 0 b.4 Root/shoot r a t i o 0 . 3 b 0.24 0.38 0.39 0.24 0.28 0 . 2 0 Depth of roots(mm) 1 7 0 215 231 170 1 8 5 182 Ibb Double thickness at the middle diameter 0.6 of f i r s t internode(mm)007 0 o 7 O.b 0.9 0.5 0.7 Double thickness a t the middle diameter of second internode(mm)0 b 0 b 0.5 0.5 0.5 O.b 0.3 Double bark thickness at the middle diameter of t h i r d internode (mm) 0 , 3 0o5 0.2 0.4 0.4 0.5 0.3 e  o  o  o  5.1  10  Averag  547 5.9  489.4 5.9  15.2  15.6  14.9  14.9  15.1  15.3  17.0  15.3  7.80  8.31  8.94  27  31  34  8.01  31.6  2.5  3.0  4.0  3.7 0.42 145  0 28  5.4  7 01 0.34 222  0.6  0.7  0.9  Q.b9  O.b  0.7  0.5  0.54  0.4  0.2  0.4  0.3b  o  225  2.9 5.6 0.30 191.1  Table l b .  C h a r a c t e r i s t i c s or" branches o f t h r e e - y e a r - o l d hemlock s e e d l i n g s i n r e l a t i o n t o i n t e r n o d e and m i c r o s i t e . (Averages of 1 0 s e e d l i n g s p e r microsite). M I C R O S I T E S  Characteristics Average l e n g t h of branches (millimeters) 180.73 Average base diameter of c branches(millimeters) 2.78 u CD Average branch angle (degrees) 35.0 •P a Average number of t h r e e - y e a r 1.1 H o l d branches Average number of two-yearU o l d branches 1.5 •rl Average number of one-yearo l d branches Average l e n g t h of branches 171.08 (millimeters) Average base diameter of CD 2.54 branches ( m i l l i m e t e r s ) GO 49.2 O C Average branch angle (degrees) OU CD CD Average number of two-year15.b W-P o l d branches C H Average number of one-year3.2 o l d branches Average l e n g t h of branches 52.24 (millimeters) CD Average base diameter of 1.57 branches ( m i l l i m e t e r s ) UC 3 9 .3 Average branch angle (degrees) •HSH XlCD Average number of one-yearH-P 17.0 o l d branches a CD  -  *s H  151.97  lbl.84  1.7b  1.78  105.94  1.14  145.b  1.87  42.5  43.7  37.8  2.b  3.7  1.1  2.0  b.4  3.0  4.1  4.2  0.4  0.2  0.2  0.2  117.83  87.7b  87.25  1.32  1.21 43.8  47.9  124.32 1.5b 48.5  43.5  1.15 3b.0  9.0  11.1  13.b  b.9  b.5  2.9  2.8  b.4  4b.48  12.87  18.bb  22.b7  1.14  0.92  0.48  0.71  3b.2  34.4  27.8  28.9  14.1  13.2  8.8  12.2  - 60 T a b l e 17.  Student's t - t e s t v a l u e s t o r t h e comparison o f t h e mean t o t a l h e i g h t of hemlock s e e d l i n g s on f i v e microsites.  M i c r o s i t e number 1 2  1  2  X  0.29 NS X  3  3  5  4  2.bl  * 7.2b**  2.29  * 60b * * 7.66** 2.02NS  X  4  9 . 4 4 * *  3.07**  1.14NS  X  5  X  Significant  Table 18.  t v a l u e s f o r 18 DF  .05 .01  2.101* 2.878**  Student's t - t e s t v a l u e s f o r t h e comparison o f the mean base diameter of hemlock s e e d l i n g s on five microsites.  M i c r o s i t e Number  1  1  2 x  2  6.70** X  3  5  4  3 7.10**  9 . 4 4 * *  1.42NS  3.12**  4.55**  1.08NS  2.01NS  X  1.20NS  X  4  5 Significant  11.68**  X  t  v a l u e s f o r 18 DF  .05 .01  2.101 * 2.878**  - 61 -  Table 19*  Student's t - t e s t v a l u e s t o r t h e comparison o f mean branch angle of t h i r d i n t e r n o d e of hemlock s e e d l i n g s on f i v e m i c r o s i t e s .  Microsite Number 1  1  2  3  1 . 5 7 NS  X  2  X  3  5  4  2.88 **  6.76**  8.06**  o.78 NS  3.81  3.57**  3.46**  3.00**  X  4  X  0.46NS  5  X  Significant  T a b l e 20.  v a l u e s f o r 18 DF  .05 .01  2.101* 2.878**  Student's t - t e s t v a l u e s f o r t h e comparison o f mean l e n g t h of l a r g e s t needles of t h i r d i n t e r n o d e of hemlock s e e d l i n g s on f i v e microsites.  Microsite Number 1  t  1  2 2.73*  X  2  X  3  5  4  3 5.73**  4.10**  1.41 NS  2.30*  8.85**  1.46NS  3.17**  X  4  X  5  14.40**  2.56* X  Significant  t  v a l u e s f o r 18 DF  .05 .01  2.101* 2.878**  - 62 -  Table 21.  Student's t - t e s t v a l u e s f o r t h e comparison of mean oven-dry weight of needles ( i n grams) o f hemlock s e e d l i n g s on f i v e microsites.  Microsite number 1 2 3 4 5  1 X  2  3  4  5  2.56*  4.88**  7.77**  7.27**  X  0.42NS  2.93**  2.52*  2.89**  2.22*  X  X  0.88NS X  S i g n i f i c a n t v a l u e s f o r 18 DF .05 2.101* .01 2.878**  - 63 Table  22.  The i n f l u e n c e of s i t e  on hemlock  seedlings.  The mean l e n g t h of l a r g e s t n e e d l e s , branch angle of 1960 i n t e r n o d e , and h e i g h t growth of hemlock s e e d l i n g s .  Plot 1 Plot 5 SI 142« SI 128  1  Plot 7 Plot 8 Plot; S I 122« SI 115• SI79  r  Mean l e n g t h of l a r g e s t needles (millimeters)  16.74  13.76  14.17  12.68  Mean branch angle I960 i n t e r n o d e (degrees)  49.0  41.0  39.9  35.1  27.4  Mean height growth 1959 ( m i l l i m e t e r s )  128.6  108.3  108.0  106.4  44.2  Mean height growth I960 ( m i l l i m e t e r s )  307.8  316.9  284.4  256.0  64.0  8.11  Mean l e n g t h of l a r g e s t n e e d l e s based on 7 0 70 70 70 42  samples samples samples samples samples  on on on on on  Plot Plot Plot Plot Plot  1 5 7 8 4  Mean branch a n g l e based on  77 94 68 69 18  samples samples samples samples samples  on on on on on  Plot Plot Plot Plot Plot  1 5 7 8 4  Mean h e i g h t growth 1 9 5 9 based on  2 0 seedlings 2 0 seedlings 2 0 seedlings 2 0 seedlings 1 2 seedlings  on on on on on  Plotl Plot 5 Plot 7 Plot a Plot 4  Mean height growth  46  seedlings seedlings seedlings seedlings seedlings  on on on on on  Plotl Plot 5 Plot 7 Plot 8 Plot4  I960  based on  46 29 29 14  - 64 -  Table 23.  The i n f l u e n c e of s i t e on hemlock s e e d l i n g s . Student's t - t e s t v a l u e s f o r comparison of mean branch angle of l a s t year's i n t e r n o d e of hemlock s e e d l i n g s on f i v e p l o t s *  Plot Number 1  5  1 x  5  8  7  5.88** X  7  4  5.90**  8.94**  9.34**  0.083NS  5.30**  9.75**  3.84**  8.26**  X  8  5.28**  X  4  X  Significant v a l u e s between  DF  P l o t s 1 and  5  169  1.974  2.606  P l o t s 1 and  7  143  1.977  2.614  P l o t s 1 and 8  144  1.977  2.614  P l o t s 1 and 4  93  1.986  2.630  t  t .05*  .01**  P l o t s 5 and  7  160  1.975  1.608  P l o t s 5 and  8  161  1.975  1.608  P l o t s 5 and 4  110  1.982  2.622  P l o t s 7 and  8  135  1.978  2.613  P l o t s 7 and 4  84  1.989  2.637  P l o t s 8 and 4  85  1.989  2.635  - 65 -  Table 24*  The i n f l u e n c e of s i t e on hemlock s e e d l i n g s * Student's t - t e s t v a l u e s f o r t h e comparison of mean l e n g t h of l a r g e s t needles of l a s t y e a r ' s i n t e r n o d e of hemlock s e e d l i n g s on five plots*  Plot Number  1  5  8  7  11.92**  9.51**  19.33**  31.85**  1.56NS  4.94**  22.48**  6.47**  21.53**  1  X  5  X  X  7  X  X  X  8  X  X  X  X  4  X  X  X  X  Significant v a l u e s between  DF  4  t  .05*  23.89** X  t  .01**  P l o t s 1 and  5  138  1.977  2.613  P l o t s 1 and  7  138  1.977  2.613  P l o t s 1 and  8  138  1.977  2.613  P l o t s 1 and  4  110  1.982  2.622  P l o t s 5 and  7  138  1.977  2.613  P l o t s 5 and  8  138  1.977  2.613  P l o t s 5 and  4  110  1.982  2.622  P l o t s 7 and  8  138  1.977  2.613  P l o t s 7 and  4  110  1.982  2.622  P l o t s 8 and  4  110  1.982  2.622  - 66 Table  25»  The i n f l u e n c e of s i t e on hemlock s e e d l i n g s * F i d u c i a l l i m i t s a t 0.01 l e v e l c a l c u l a t e d f o r the mean branch angles (.in degrees) o f hemlock s e e d l i n g s on f i v e p l o t s .  P l o t number  Upper l i m i t  Lower  limit  1  52.02  46.02  5  42.82  39.20  7  41.88  37.96  8  37.37  32.77  4  28.63  20.81  Table  26.  The i n f l u e n c e of s i t e on hemlock s e e d l i n g s . F i d u c i a l l i m i t s a t 0.01 l e v e l c a l c u l a t e d f o r the mean of l a r g e s t n e e d l e s ( i n m i l l i m e t e r s ) of hemlock s e e d l i n g s on f i v e p l o t s .  Flot  number  Upper l i m i t  Lower  limit  1  17.22  16.26  5  14.21  13.31  7  14.67  13.67  8  12.97  12.39  4  8.54  7.74  - 67  -  Table 27. E s t i m a t i o n of hemlock s i t e index of m i c r o s i t e s by u s i n g needle l e n g t h and branch angle of hemlock s e e d l i n g s .  Estimated s i t e  index  1  M i c r o s i t e Number 2 3 4 5  Based  on needle l e n g t h  123  112  104  94  78  Based  on branch angle  121  113  107  91  93  68  Fig- 2 Cumulative light intensity of various degrees of shade for 12 hour period of bright day 30,000  Plot V  Plot IV 20,000k to O  o  c  Plot III  c  a> 10,000  3  o  Plot II  Plot I Control 0700  1000  1400 Time of Observation  Line excised because of high value  1800  -  Fig- 3  69  -  Cumulative light intensity of various degrees of shade for  12 h o u r  30,000r-  period o f bright d a y  / Plot 5  20,000^  Plot  4  Plot  3  1  o  I Ii. c CO c ®  «  > 10,000h o  E o  .Plot 2 Plot I Control 0700  1000 1400 Time of Observation  Line excised because of high value  1800  - 70 Fig- 4  C u m u l a t i v e l i g h t i n t e n s i t y o f v a r i o u s d e g r e e s o f s h a d e f o r 12 hour  period of cloudy d a y  Time of Observation  - 71 Fig- 5 Cumulative light intensity of various degrees of shade for 12 hour period of cloudy day 15000  Plot 5  CO  a> TJ  c o o lOOOOh o .o  CO  c  co  « o 3  E 3  o 5000  Control Plot 0700  8  I  10  I II  I 1 12 13 Time of Observation  14  15  16  17  1800  Fig- 6  Average cumulative length of internodes of 11-year old seedlings on plots of various degrees of shade  4000h  5 Base  6  Internodal Number  Tip  - 73 -  Fig- 7  Average cumulative length of internodes of 6-year old seedlings on plots of various degrees of shade  Base  Internode Number  Tip  Fig- 8  Average  cumulative diameter  v a r i o u s d e g r e e s of  shade  of internodes  of 11-year  old seedlings on  plots  of  Fig- 9  Average cumulative diameter of internodes of 6-year old seedlings on plots of various degrees of shade  Plot  /V  4 0 h  30 h  Base  Internode Number  Tip  F i g - 10  Average cumulative number of green branches of 11-year old s e e d l i n g s on plots of various d e g r e e s of  shade  Fig-  II  Average  cumulative number of green  on  of various degrees of shade  plots  branches of 6-year old seedlings  Plot  V  Plot IV  P l o t III P l o t II —  Base  Internode Number  Plot  Tip  Fig- 12 Average total height of 3-year old seedlings on various microsites  Microsite I  Microsite 2  Microsite 3  Microsite 4  Microsite 5  Microsite  Microsite 2  Microsite 3  Microsite 4  Microsite 5  Fig- 13 Average base diameter of 3-year old seedlings on various microsites  - 79 -  14 Average ovendry weight of roots of 3-year old seedlings on various microsites 01  e o  2 0  o o  cr  E? 5  T3 C  10  CD >  o  CD  o  a> > <  I  1 1 l_ Microsites  15  Average depth of roots of 3-year old seedlings on various microsites  = 80 -  16  Average ovendry weight of needles of 3-year old seedlings on various microsites  1 20 o <»>  '5>  * 10 >. c  Q> >  o  •  in 2f CT>  $5  17  •  Microsites  Average number of green branches of 3-year old seedlings on various microsites co CP JE  o  § 40 CD CO  JO  E  Z320 D  ,o CD  a> o CO  >  < Microsites  - 81 F i g - 18  T h eaverage  branch angle of three internodes o n  various microsites 50-  <  25  o  « o  I  Microsite I  19  Microsite 2  Microsite 3  Microsite 4  Microsite 5  T h e average length of largest needles o f three internodes onvarious microsites  CO  I  20-  9  z •s O  £ *C  -—10. k. »  • E  oo> — =  a> £ —•  <  Microsite I  I Microsite 2  I Microsite 3  I Microsite 4 I Microsite 5  Legend First internode (from base) f[[Jj Second internode  Q  Third internode  Q  82 -  Fig- 2 0  The average length of branches of three internodes on various microsites  200-  $ w  CQ »*-  o 100o» CO  _l % o E o> —  11  > <  EL  —  Microsite I |  Fig- 21  Microsite 2 ] Microsite 3  | Microsite 4 | Microsite 5  The average base diameter of branches of three internodes on various microsites  3 " •0  s  1 ' 2  is  3  Microsite I  Microsite 2  Microsite 3 I Microsite 4 Legend  First internode (from baa Second internode Third internode  ] ^  Microsite 5  |  Fig  2 2  R e g r e s s i o n line b a s e d o n b r a n c h a n g l e a n d s i t e hemlock  f o r study o f t h e influence of site on  seedlings  60h  50  +  aa>> k.  I  «  •a  » c40l <  5  -P  c  o o m  8 4-  OJ  2 30 a> > <  Equation of line  Y= 0-374  X -  4  5-88  +  20 *  Site index is calculated for hemlock  150  140  130  ±  120 Site Index  110  100  90  80  Site index is calculated for hemlock  J  150  I  140  I  130  I  Site  120  I  Index  110  I  100  I  90  L.  80  - 85 B I B L I O G R A P H Y Allen  G©S  0  1941 L i g h t a n d t e m p e r a t u r e a s f a c t o r s i n t h e germination of seed o f Douglas f i r ( P s e u d o t s u g a t a x i f o l i a ) Lambo B r i t t , F o r , C h r o n „ 17 (3) (99-109) 1958  Factors a f f e c t i n g the v i a b i l i t y mination behavior of coniferous For© C h r o n . 34 (3) (200-298)  OOOOOOOOOOO A l l e n G,S„,  and g e r seed 0  I , K B a r b e r a n d I a n Mahood 1955 - T h e 1951 a e r i a l b a i t i n g and s e e d l i n g p r o j e c t , Ash R i v e r T r a c t , M a c M i l l a n and B l o e d e l Limited F o r , C h r o n , 31 (1) (45-59) 0  0  Atkins  E,S,  1957 L i g h t measurement i n a s t u d y r e p r o d u c t i on, For, Research Div, T e c h n i c a l  of white N o t e No,  pine bO,  (5-18) B a k e r S, F r e d e r i c k 1929© E f f e c t o f e x c e s s i v e l y h i g h temp e r a t u r e s on c o n i f e r o u s reproduction. Reprinted from J o u r n a l o f F o r e s t r y , V o l , XXVII, No, 8. (949-974) o o o o o e o o e o o e 1948  A s h o r t method o f d e t e r m i n i n g l e a f a r e a and volume g r o w t h i n p i n e t r e e s , H i l g a r d i a , (Journal of A g r i c u l t u r a l Science, P u b l i s h e d by the C a l i f o r n i a A g r i c u l t u r a l E x p e r i m e n t S t a t i o n ) (335-355)  oo,oooo,oo9«l950 P r i n c i p l e s o f S i l v i c u l t u r e , Book Co,,  New Y o r k ,  McGraw  Hill  (125-150)  B e r n t s e n C M , 19b0, P l a n t i n g s i t k a s p r u c e and D o u g l a s f i r on d e c a y e d wood i n c o a s t a l O r e g o n R e s e a r c h Note No, 197 (1-5)• U.S,D,A, P a c i f i c N o r t h w e s t F o r e s t a n d Range E x periment S t a t i o n , P o r t l a n d , Oregon, 0  oooeoo,,oo  ,1958  S i l v i c a l c h a r a c t e r i s t i c s o f w e s t e r n hemlock, S i l v i c a l s e r i e s No, 3 ( l - l b ) U,S,D A, P a c i f i c N o r t h w e s t F o r e s t a n d Range E x p e r i m e n t S t a t i o n , Portland, Oregon, 0  0  B e v e r D a l e N,  1954©  Evaluation of factors affecting natural reproduction of forest trees i n Central Western Oregon, R e s e a r c h B u l l e t i n No, 3 (l-46)e Oregon S t a t e Board o f F o r e s t r y , Salem, Oregon,  - 86 B i e n t j e s W. 1954  The e f f e c t s o f t e m p e r a t u r e , s e e d m o i s t u r e , and s t r a t i f i c a t i o n on t h e g e r m i n a t i o n b e h a v i o r o f western hemlock s e e d R e s e a r c h N o t e No. 11 (1-7) U.B.C. F o r e s t Club R e s e a r c h Committee, Vancouver, 0  B o u r d e a u P,F, a n d M i r i a m L , L a v e r i c k 1958 a Tolerance of photosynthetic adaptability t o l i g h t i n t e n s i t y i n w h i t e p i n e , r e d p i n e , heml o c k and A i l a n t h u s s e e d l i n g s .  F o r , S c , 4 (3)  (196-207)O  B u c k l a n d D . C 1956,  T e r m i n a l shoot growth o f f o u r western c o n i f e r s f o r a s i n g l e growing season. F o r , C h r o n , 32 (4) (307-399).  C h i n g T e May 1958  Some e x p e r i m e n t s on t h e optimum g e r m i n a t i o n c o n d i t i o n s f o r western hemlock. J o u r n a l o f F o r e s t r y , 56 (4) (277-279).  Dept, o f Lands and F o r e s t s , Continuous f o r e s t inventory o f B r i t i s h C o l u m b i a 1957. Dept, o f L a n d s a n d F o r e s t s B , C , B,C, F o r e s t S e r v i c e , V i c t o r i a , (1-38), Dallimore Downs R.Jo  W, & J a c k s o n A , B r u c e 1948, A handbook o f c o n i f e r a e , E d . 3. L o n d o n (682 p p ) . a n d A,A, P i r i n g e r 1958. E f f e c t s of photoperiod and k i n d o f s u p p l e m e n t a l l i g h t on v e g e t a t i v e growth of p i n e s .  F o r . S c . 4 (3)  (185-195).  D u f f i e l d , J.W, 1955. S e l e c t i n g p l u s t r e e s f o r o u r seed orchards. Industrial Forestry Association, P o r t l a n d , Oregon. (15 p a g e s ) . Garman E,H, 1951  Gerhold  H.D. 1959  Seed p r o d u c t i o n by c o n i f e r s i n t h e c o a s t a l r e g i o n o f B r i t i s h Columbia r e l a t e d t o d i s s e m i n a t i o n and r e g e n e r a t i o n . T e c h n i c a l P u b l i c a t i o n T, 35 (1-45). Department o f Lands and F o r e s t s B r i t i s h Columbia F o r e s t S e r v i c e , V i c t o r i a . Seasonal d i s c o l o r a t i o n o f Scotch pine i n relation t o microclimatic factors.  F o r . S c . 5 (4) Godman R.M. 1953  (333-343).  Seed d i s p e r s a l i n S o u t h - e a s t A l a s k a . T e c h n i c a l N o t e No. 16 (2 p p ) o f t h e A l a s k a F o r e s t Research Center U»S.D.A. Juneau, Alaska.  - 87 Godman R M C  a n d R,A  0  G r e g o r y 1955o Seasonal d i s t r i b u t i o n of r a d i a l and l e a d e r growth i n the s i t k a s p r u c e w e s t e r n hemlock f o r e s t s o f s o u t h east Alaskao J o u r n a l o f F o r e s t r y 53 (11) (827-835)0  0  1957  G r e g o r y R©A©  A comparison between l e a d e r growth o f western c o n i f e r s i n A l a s k a and Vancouver Islando T e c h n i c a l N o t e s N o 36 2 p p Alaska Forest Research Center U S D A « Juneau, Alaska© e  c  0  G r i f f i t h B.G©  1959  The e f f e c t o f t h i n n i n g of w e s t e r n hemlocks  For. 0  9  0  0  0  0  0  0  0  0  I960  C h r o n , 35 ( 2 )  on a y o u n g  a  stand  (114-133)o  Growth o f D o u g l a s f i r a t t h e U n i v e r s i t y of B r i t i s h C o l u m b i a R e s e a r c h F o r e s t a s r e l a t e d t o c l i m a t e and s o i l . Faculty of F o r e s t r y o f U.B.C© Forestry Bulletin No. 2. ( 1 - 5 8 ) , V a n c o u v e r , B C 0  H a c s k a y l o E , & A,G©  0  0  Snow, 1959s Relation of soil nutrients and l i g h t t o p r e v a l e n c e o f m y k o r h i z a e on pine seedlings S t a t i o n P a p e r No. 125 (1-13) N o r t h e a s t e r n F o r e s t Experiment S t a t i o n U.S.D.A. U p p e r D a r b y . , P a . 0  H a i g I . T . 1936  Factors c o n t r o l l i n g i n i t i a l establishment of w e s t e r n w h i t e p i n e a n d a s s o c i a t e d species. Yale University School Fore s t r y Bui© 41o 149 PPo New H a v e n , Connecticut.  H a i g I . T . , K e n n e t h P D , a n d R.H. Weidmann 1941o Natural reproduction i nthe western white pine type. T e c h n i c a l B u l l e t i n No. 767 (1-98) UsS.D.A. N o r t h e r n R o c k y M o u n t a i n F o r e s t and Range E x p e r i m e n t a l S t a t i o n F o r e s t S e r v i c e , W a s h i n g t o n , D©C© 0  H a l l i d a y W E©D© 0  1937  0  A f o r e s t c l a s s i f i c a t i o n f o r Canada. F o r e s t R e s e a r c h D i v i s i o n B u l l e t i n No, 89 ( 5 - 2 7 ) j F o r e s t r y B r a n c h Canada D e p a r t ment o f R e s o u r c e s a n d D e v e l o p m e n t , Ottawa.  Brown A.W.A. 1943  T h e d i s t r i b u t i o n o f some i m p o r t a n t t r e e s i n Canada© Ecology© 24 (3) J u l y 1943 (3^4-366)  H a r l o w & H a r r a r 1941  T e x t b o o k o f D e n d r o l o g y . M c G r a w - H i l l Book Co© I n c . , New Y o r k a n d London© (503 p p )  H a t c h A.B.  forest  a n d Doak K D . 1933 M y k o r r h i z a l and o t h e r f e a t u r e s of t h e r o o t s y s t e m s o f Pinus.©., J o u r n a l o f t h e A r n o l d A r b o r e t u m © Vol© X l V . (85-99) 0  -  H e l l m e r s Henry  1959  88 -  P h o t o p e r i o d i c c o n t r o l o f bud d e v e l o p ment i n c o u l t e r p i n e a n d b i g c o n e Douglas f i r .  For. Sc. 5 ( 2 )  Hoffmann J . V . 1918  (135-141)  The i m p o r t a n c e o f s e e d c h a r a c t e r i s t i c s i n the natural reproduction of coniferous f o r e s t s . B u l l e t i n o f t h e U n i v e r s i t y o f Minnes o t a No, 2 ( 8 - 2 5 ) Minneapolis. The n a t u r a l r e g e n e r a t i o n o f D o u g l a s f i r i n t h e P a c i f i c Northwest. Department B u l l e t i n No. 1 2 0 0 ( 1 - 5 5 ) U.S.D.A, W a s h i n g t o n , D.C.  I960  Hough A . F .  S i l v i c a l c h a r a c t e r i s t i c s of eastern hemlock. S t a t i o n P a p e r No. 1 3 2 (1-16), U.S.D.A. N o r t h e a s t e r n F o r e s t Experiment S t a t i o n , Upper Darby, P a 0  Isaac  LoAo 1928  H o p k i n s H.G.  1937  Measured f l i g h t o f seed of Douglas f i r and i t s a s s o c i a t e s . P a c i f i c Northwest F o r e s t a n d Range E x p t . S t a t i o n U n p u b l i s h e d ( 7 Pp) T y p e w r i t t e n . The f o r e s t s o i l o f t h e D o u g l a s f i r r e g i o n a n d c h a n g e s wrought upon i t b y l o g g i n g and s l a s h b u r n i n g . R e p r i n t e d f r o m E c o l o g y 1 8 ( 2 ) (264-279)  actors eeooooo o o o o o « o X 9 3F ^  a f f e c t i n g establishment of Douglas f i r s e e d l i n g s . C i r c u l a r No. 4 8 6 ( 1 - 4 5 ) U.S.D.A., W a s h i n g t o n , D.C.  K i n g h o r n J.M. 1 9 5 4  Kittredge  I . 1945  K o z l o w s k i T . T . 1957  The i n f l u e n c e o f s t a n d c o m p o s i t i o n on the m o r t a l i t y of various c o n i f e r s , caused by d e f o l i a t i o n by western l o o p e r on V a n c o u v e r I s l a n d , B.C. F o r . Chron. 3 0 ( 4 ) ( 3 8 0 - 4 0 0 ) . E s t i m a t i o n o f t h e amount o f f o l i a g e o f t r e e s and stands. J o u r n a l o f F o r e s t r y 4 3 (1) ( 9 0 5 - 9 1 2 ) E f f e c t of continuous, high l i g h t i n t e n s i t y on p h o t o s y n t h e s i s of forest tree seedlings. For.  Krajina V.I,  Kramer P.J.  1959  1957  Sc. 3  (3)  (220-223).  B i o c l i m a t i c zones i n B r i t i s h Columbia. U.B.C. B o t a n i c a l S e r i e s . No. 1 ( 1 - 4 5 ) Some e f f e c t s o f v a r i o u s  combinations of  - 89 -  ooooeoeoooeoooo.o 1957  o f day and n i g h t temperatures and p h o t o p e r i o d on t h e h e i g h t g r o w t h o f l o b l o l l y pine seedlings.  For, Sc, 3 U ) Kunz R, 1953  (45-54).  Morphologische Untersuchungen i n naturlichen Fohrendickungen M i t t e i l u n g e n d e r Schweizenschen A n s t a l t f u r das F o r s t l i c h e Versych0  swen 29 12)  1335-402),  L e B a r r o n R u s s e l K, 1944  Influence of c o n t r o l l a b l e environm e n t a l c o n d i t i o n s on r e g e n e r a t i o n of j a c k p i n e and b l a c k s p r u c e . R e p r i n t e d from J o u r n a l o f A g r i c u l t u r a l R e s e a r c h , 68 (3) 197-119), W a s h i n g t o n , D,C,  Lindquist  G e n e t i c s i n Swedish F o r e s t r y P r a c t i c e , S t o c k h o l m , Sweden 5,173 p a g e s )  B . 1948  L o g a n K.T. 1959  Some e f f e c t s o f l i g h t on g r o w t h o f w h i t e p i n e seedlings© Technical Note No, 82 ( 1 - 1 8 J F o r e s t r y B r a n c h Canada D e p a r t m e n t o f N o r t h e r n A f f a i r s and N a t u r a l R e s o u r c e s ,  Ottawa,  Mergen F . 1958,  D i s t r i b u t i o n o f r e a c t i o n wood i n e a s t e r n hemlock a s a f u n c t i o n o f i t s t e r m i n a l growth.  F o r , S c . 4 (2)  oooooooooooooooo  N o r d i n V . J . 1950  Olson  I960  (98-109)  V a r i a t i o n and h e r i t a b i l i t y o f p h y s i o l o g i c a l and m o r p h o l o g i c a l t r a i t s i n Norway s p r u c e . Fifth World F o r e s t r y Congress SP/105-11U o S . A . (5 p a g e s ) , A g a l l d i s e a s e o f w e s t e r n hemlock i n B r i t i s h Columbia. The F o r e s t r y C h r o n i c l e . 24 (4) (2 p p )  J . S e , F o r e s t W. S t e a r n s a n d H. N i e n s t a d t 1959. E a s t e r n hemlock s e e d s a n d s e e d l i n g s r e s p o n s e t o p h o t o p e r i o d and t e m p e r a t u r e .  B u l l e t i n No. 620 ( 5 - 7 0 ) .  The C o n n e c t i c u t E x p e r i m e n t New Haven. Papp L . 1959  Station,  T h e q u a l i t y o f s e e d l i n g s and t h e success of p l a n t i n g . E r d o 8 (9) (335-343)o  - 90 Schubert  H G 0  C  1952  Germination of various coniferous seeds a f t e r c o l d s t o r a g e Forest R e s e a r c h N o t e , No, 38 (1-7)« U,S,D,A, C a l i f o r n i a F o r e s t a n d Range E x p e r i m e n t Station, Berkeley, C a l i f o r n i a , 0  S i g g i n s H W, 0  1933  D i s t r i b u t i o n and r a t e o f f a l l o f c o n i f e r seeds. J o u r n a l A g r , R e s e a r c h 47  (119-128),  S i n n o t t W,E,  a n d W i l s o n K,S, 1955« Botany, P r i n c i p l e s and P r o b l e m s (60-130), M c G r a w - H i l l Book Co, I n c , New Y o r k , T o r o n t o , L o n d o n 0  0  The i n f l u e n c e o f s e e d b e d c o n d i t i o n s on t h e r e g e n e r a t i o n o f e a s t e r n w h i t e  S m i t h D „ M , 1951  pine.  B u l l e t i n 545 (27-56),  Connecticut A g r i c u l t u r a l S t a t i o n , New Haven,  The  Experimental  S u t t o n R,C, 1954  Some i n f l u e n c e s o f s e e d b e d on g r o w t h and s u r v i v a l o f w e s t e r n h e m l o c k . R e s e a r c h Note No, 10 (1-4)» U,B,C, F o r e s t C l u b R e s e a r c h Committee, Vancouver,  T o d a , R, 1957  V a r i a t i o n a n d h e r i t a b i l i t y o f some quantitative characters i n Cryptomeria. Government F o r e s t E x p e r i m e n t a l S t a t i o n , N i y a z a k i , Japan ( 8 7 - 9 3 )  U,S,  Department  o f A g r i c u l t u r e 1948 Woody p l a n t s e e d manual, (416 pp)« U S o Government P r i n t i n g Office, W a s h i n g t o n , D,C, 0  0  Walters J , i 9 6 0  The b r a n c h a r r a n g e m e n t o f w e s t e r n heml o c k (Tsuga h e t e r o p h y l l a ) ( R a f ) S a r g , R e s e a r c h N o t e No, 29 (2 p p ) F a c u l t y of F o r e s t r y U « B , C , Vancouver, 0  W a l t e r s J « , J , S o o s a n d P,G Haddock I960, The s e l e c t i o n o f p l u s t r e e s on t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a R e s e a r c h F o r e s t , Haney, B,C, 0  R e s e a r c h P a p e r No, 33 (1-11), Faculty W a l t e r s J , and J  0  of F o r e s t r y  o f U,B,C,,  Vancouver,  S o o s 1961 Some o b s e r v a t i o n s on t h e r e l a t i o n s h i p o f lammas s h o o t s t o t h e f o r m and g r o w t h o f D o u g l a s - f i r s e e d l i n g s , (in press), 0  - 91 W e l l n e r C.A. 1946  Improving composition i n young western white pine stands. R e s e a r c h Note No. 43 (6 p p ) . U.S.D.A. N o r t h e r n R o c k y M o u n t a i n F o r e s t a n d Range E x p e r i m e n t S t a t i o n , M i s s o u l a , Montana.  W e l l w o o d R.W. 1956  Some e f f e c t s o f d w a r f m i s t l e t o e on west e r n hemlock. The F o r e s t r y C h r o n i c l e 32 (3) (282-296)  OOOOOGOOOOOOOO1957  U n p o p u l a r s p e c i e s and t h e i r i n c r e a s e d utilization. The F o r e s t r y C h r o n i c l e 33 (1) (7-9).  W i k s t e n A . 1953  R e p o r t on r e p r o d u c t i o n s t u d i e s Queen Charlotte Islands. Unpublished. R e s e a r c h P r o j e c t No. 8 (1-25) Mimeographed.  W r i g h t J . G . 1943  Measurement o f t h e d e g r e e o f s h a d i n g o r crown canopy d e n s i t y i n f o r e s t s i t e s . The F o r e s t r y C h r o n i c l e V o l . 19 (3)  (183-185).  Zobel  B . J . 1957  O . O O O O . . . O O . . 1 9 6 0  F o r e s t G e n e t i c s and P u l p I n d u s t r y . T a p p i V o l . 40 No. 3 ( 4 2 - 4 4 ) . Inheritance of c e r t a i n important charact e r i s t i c s f o rconifers. Southern Lumberman (153-158).  Fig-1  L o c a t i o n o f plots e s t a b l i s h e d f o r v a r i o u s  studies  Plot  2  A  2 B  Ic LEGEND Plots for light intensity study Plots for microsite study Plots for study of the effect of site Compartment boundary  Scale: | = IOOO  1000  500  0  JL  • 1000 Feet  2000  I  3000  Fig- 2  Cumulative for  12 h o u r  light intensity o f various d e g r e e s o f shade period o f bright d a y  / *  30,000k  /  Plot  V  Plot IV 20,000 CO 0>  TJ  c o  o  o o u.  CO  P l o t III  c  cn « 10,000 E 3  o  P l o t II  Plot I Control 1 1400 Time of Observation Line excised because of high value 0700  —1 1000  1 — — i  1  P  1800  Fig- 3  Cumulative light intensity of various degrees of shade for  12 h o u r  30,OOOh  period o f bright d a y  /  / Plot 5  CO  20,000  Plot  a>  4  o O o o  CO  c  0)  JC >  o 3  10,000  £ 3  CJ  Plot 3  Plot 2 Plot I Control 0700  1000  1400 Time of Observation  Line excised because of high value  1800  Fig- 4  C u m u l a t i v e l i g h t i n t e n s i t y o f v a r i o u s d e g r e e s o f s h a d e f o r 12 hour period o f cloudy d a y  Time of Observation  Fig- 5 Cumulative light intensity of various degrees of shade for 12 hour period of cloudy day 15000  Plot 5  Time of Observation  Fig 6  Average cumulative length of internodes of 11-year old seedlings on plots of various degrees of shade  40001£3500  •| 3000  £2500 <D  "12000 3  6 o 1500 o> o w  > < lOOOh  500  Base  Internodal Number  Tip  Fig- 7  Average  cumulative length o f internodes of 6-year o l d  seedlings o n plots of various degrees o f shade  Base  Internode Number  Tip  Fig- 8  Average  cumulative diameter  v a r i o u s d e g r e e s of  shade  of internodes  of 11-year old s e e d l i n g s on  plots  of  Fig- 9  Average  cumulative diameter of internodes of 6 - y e a r old seedlings  on plots of various degrees of shade 90r-  80h  60h  40h  30h 20r-  ioh  3 Internode Number  4  Fig-  10  Average cumulative number of green branches of 11-year old s e e d l i n g s on plots of various d e g r e e s of  shade  Fig-  II  Average on  c u m u l a t i v e n u m b e r of g r e e n  plots of various degrees  of  b r a n c h e s of 6 - y e a r old  seedlings  shade  90rCO CD  x: o c  Plot  V  Plot  IV  80  D  w  m 70h CD CD  o  H_  60  CD  E  3  50  Z  CD  £a 4 0 3  E  3 CD  30  cn o CD  >  20  P l o t III  10  P l o t II Plot I  Base  Internode Number  Tip  F i g - 12  A v e r a g e total height of various  Fig-  13  Average  3-year  old seedlings  on  microsites  b a s e d i a m e t e r of 3 - y e a r  various microsites  old seedlings  on  Fig- 14 Average ovendry weight of roots of 3-year old seedlings on various microsites CO  E o i_  cr>  o cr  o  '55  l>0  c a> > O CD  cn a CD  5 Microsites  Fig-15  Average depth of roots of 3-year old seedlings on various microsites  CO  0)  -t— CD  E E 200  o oor £" ioo  o  CD  cn D  w.  CD > <  Microsites  Fig-16  Average  ovendry  weight of needles of 3 - y e a r old seedlings  on various microsites { 20 o  £  10  •D C  <u > O co  o>  ^ in  I o fl) 2  Fig  17  I  2  3 Microsites  Average number of green branches on v a r i o u s  microsites  CO  a>  § 40 CO CD  E  f  a  20  .o CT  k.  o  CD  > <  Microsites  •  of 3 - y e a r  •  old seedlings  F i g - 18  T h eaverage  branch angle of three internodes o n  various microsites  co CD CD i>_ cn CD  50-  T3  .CD cn  JZ 2 5 o c D  V-  CD  CD cn D w CD  Microsite I  | Microsite 2  | Microsite 3 | Microsite 4 | Microsite 5  F i g - 19 T h e a v e r a g e l e n g t h o f l a r g e s t n e e d l e s o f t h r e e internodes o nvarious microsites CO  a>  | z  20-  to CD cn  10cn oj  c  L_  co a> - a> J> E 1  cn —  a —  S£ > >—  <  Microsite I  Microsite 2  I Microsite 3  I Microsite 4 I Microsite 5  Legend First internode (from base) Second internode Third internode  Q  Fig- 2 0  T h e average  length of branches o f three  internodes  on v a r i o u s m i c r o s i t e s 20010 CP  o c o  k.  m  100-  CJ) CO  —' "5 a> E u> —  11 >^ < Fig- 21  Microsite I | Microsite 2 | Microsite 3  £1  | Microsite 4 | Microsite 5  T h e average base diameter o f branches o f three internodes o n various microsites  3 ' CO  a) "5  1' 2  co  aj E o CL) CO a  CO  co  o  CP  >  <  Microsite I  I Microsite 2 | Microsite 3 | Microsite 4 | Microsite 5 Legend First internode (from bqse)f|T[| Second internode Third internode  Fig- 2 2  R e g r e s s i o n line b a s e d o n b r a n c h a n g l e a n d s i t e hemlock  f o r study o f the influence o f site on  seedlings  60-  50  4-  to  I  OJ  a > k_ o > a>  T3  «>  5  c?40 <  - P  o c o  8  L_  +  CD  a> o> S 30 a> > <  Equation o f line  Y *0-374  X -  4  5-88  +  20 *  Site index is calculated for hemlock  150  140  130  i 120 Site Index  110  100  90  80  Fig- 2 3  Regression  line b a s e d o n length o f needles a n d site f o r  study of the influence o f site on hemlock  + 5  seedlings  + 7 8  E q u a t i o n o f line  Y =0132  X - 2-38  Site index is calculated for hemlock  150  140  130  Site  120  Index  110  100  90  80  Fig-1  L o c a t i o n o f plots e s t a b l i s h e d f o r v a r i o u s  studies  LEGEND Plots for light intensity study  •  Plots for microsite study  •  Plots for study of the effect of site  |  |  Compartment boundary  Scale: l"= IOOO'  IOOO I  500 1  0 1  1000 1 Feet  2000 1  3000 1  

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