Open Collections

UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

Some problems in testing provenance with special reference to the co-operative Douglas-fir provenance… Reuter, Franz 1971

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Item Metadata

Download

Media
831-UBC_1971_A6 R48.pdf [ 5.58MB ]
Metadata
JSON: 831-1.0075382.json
JSON-LD: 831-1.0075382-ld.json
RDF/XML (Pretty): 831-1.0075382-rdf.xml
RDF/JSON: 831-1.0075382-rdf.json
Turtle: 831-1.0075382-turtle.txt
N-Triples: 831-1.0075382-rdf-ntriples.txt
Original Record: 831-1.0075382-source.json
Full Text
831-1.0075382-fulltext.txt
Citation
831-1.0075382.ris

Full Text

SOME PROBLEMS IN TESTING PROVENANCE WITH SPECIAL REFERENCE TO THE CO-OPERATIVE DOUGLAS-FIR PROVENANCE TEST AT THE UNIVERSITY OF BRITISH COLUMBIA RESEARCH FOREST  by  FRANZ REUTER o  Dlplomforstwirt, U n i v e r s i t y of F r e i b u r g , Germany, 196?  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF Master o f F o r e s t r y i n the F a c u l t y of Forestry  We a c c e p t t h i s t h e s i s as conforming t o the r e q u i r e d standard  THE UNIVERSITY OF BRITISH COLUMBIA February, 1971  In p r e s e n t i n g t h i s  thesis  in p a r t i a l  f u l f i l m e n t o f the requirements f o r  an advanced degree at the U n i v e r s i t y of B r i t i s h the I  Library shall  f u r t h e r agree tha  for  scholarly  by h i s of  make i t  this  written  permission  It  for financial  i s understood that copying o r gain shall  Forestry  The U n i v e r s i t y o f B r i t i s h Vancouver 8, Canada  Date  r e f e r e n c e and  f o r e x t e n s i v e copying o f t h i s  permission.  Department of  for  I agree  Columbia  February 3 r d , 1971  that  study. thesis  purposes may be granted by the Head of my Department  representatives. thesis  freely available  Columbia,  or  publication  not be allowed without my  ABSTRACT Importance of r e s e a r c h on the provenance problem, which i s b a s i c a l l y one of seed t r a n s f e r from c o l l e c t i o n  site  t o o u t p l a n t i n g a r e a , i s d i s c u s s e d w i t h s p e c i a l emphasis on c o a s t a l D o u g l a s - f i r (Pseudotsuga  m e n z i e s i i (Mirb.) Franco v a r .  menziesii). The begun i n 1957  "Co-operative D o u g l a s - f i r provenance t e s t , " and i n v o l v i n g s i x t e e n c o a s t a l seed sources  B r i t i s h Columbia,  from  Washington and Oregon, i s d e s c r i b e d i n d e t a i l .  Height measurements c o l l e c t e d 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 , when the t r e e s were e l e v e n years o l d ,  are analysed and  the r e s u l t s are d i s c u s s e d .  g e n e i t y and young age  Due  to s i t e hetero-  of the Co-operative t e s t , no  h e i g h t growth d i f f e r e n c e s between provenances  significant  can be shown,  although the l o c a l seed source, from 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 , seems t o be the f a s t e s t growing  the southernmost  and  o r i g i n , Butte F a l l s , the slowest of a l l prov-  enances i n v e s t i g a t e d .  The Co-operative t e s t i s c r i t i c a l l y  e v a l u a t e d and s p e c i f i c a t i o n s f o r f u r t h e r s t u d i e s are recommended .  ACKNOWLEDGMENTS Above a l l I wish t o thank Dr. P.G. Haddock who suggested  the s u b j e c t f o r t h i s t h e s i s and p a t i e n t l y p r o v i d e d  help and guidance. The measurements were obtained w i t h a s s i s t a n c e from Mr. J . Walters who c o n s t r u c t i v e l y c r i t i c i z e d t h e s i s  drafts.  I am indebted t o Dr. A. Kozak and Miss L . Cowdell f o r t h e i r a s s i s t a n c e i n a n a l y z i n g the d a t a . S p e c i a l thanks a r e due Dr. 0. S z l k l a i f o r c h a i r i n g the t h e s i s committee. S e v e r a l graduate t h e i r p r e c i o u s time.  f o r e s t r y students gave me much o f  The h e l p of Messrs. D. Ormerod,  L. L a c e l l e and G. P a i l l e w i l l be g r a t e f u l l y remembered. Dr. K.K. Chlng of the Oregon S t a t e U n i v e r s i t y o b l i g i n g l y answered many q u e s t i o n s . Mr.  C. Heaman l e n t me h i s e x c e l l e n t r e p o r t on the  "Co-operative D o u g l a s - f i r provenance t e s t " i n B r i t i s h Columbia. I am deeply o b l i g e d t o my w i f e R i t a f o r her c o n t i n u a l support.  i l l  ABSTRACT  i  ACKNOWLEDGMENTS TABLE OF CONTENTS  i i i i i  LIST OF TABLES . LIST OF FIGURES  *  v vi  CHAPTER I. II.  III.  INTRODUCTION AND OBJECTIVES  1  LITERATURE REVIEW ON PROVENANCE  1  D e f i n i t i o n o f Provenance The Provenance Problem O b j e c t i v e s of Provenance T e s t s Provenance T e s t s and F o r e s t Tree Improvement B r i e f H i s t o r y o f Provenance T e s t i n g Problems Commonly I n v e s t i g a t e d Drought r e s i s t a n c e . . . . . . S i t e ecotypes Aspect r a c e s Bud b u r s t i n g Lammas shoots • Frost resistance C l i n e or ecotype? Climate Seed movement .  2 2 6 7 7 8 9 9 10 11 l4 l6 19 20 22  E a r l y Tests  24  MAJOR PROVENANCE TESTS WITH DOUGLAS-FIR. . . .  2?  T e s t s Made Outside I t s N a t u r a l Range . . . . Major Provenance T e s t s With D o u g l a s - f i r  27  Within I t s Range IV.  32  THE CO-OPERATIVE DOUGLAS-FIR PROVENANCE TEST .  38  General. Hypotheses t o be T e s t e d . . . . . . . . . . . Design Co-operators and Seed Sources Phases .  38 39 40 41 46  Page S e l e c t i o n of o u t p l a n t i n g areas and seed collection . Nursery phase. P l a n t i n g of the n u r s e r y s t o c k . . . . . . F i e l d examinations  49  Results. V.  METHODS  55  The  55  Study A r e a General d e s c r i p t i o n of 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 . . . Climate. . . L o c a t i o n of the c o - o p e r a t i v e provenance t e s t on the Research F o r e s t . . . . . . H i s t o r y of the t e s t s i t e . . . . . . . .  S t a t i s t i c a l Analysis  6l  RESULTS AND  6l  DISCUSSION  6l 65 67 73 75 Earlier 76  Results? EVALUATION OF THE  CO-OPERATIVE PROVENANCE TEST  IX.  CONCLUSIONS AND  79  RECOMMENDATIONS  81  SUMMARY  83  LITERATURE CITED APPENDIX.  77 77 77  Good P o i n t s Weak P o i n t s VIII.  58 58 60  Height a t Age E l e v e n R e s u l t s by I n d i v i d u a l B l o c k Early Tests Brush Competition. . The L o c a l Provenance How do These R e s u l t s Compare w i t h  VII.  55 56  Measurements.  Height  VI.  46 47 48 48  S c i e n t i f i c and i n the t e x t  common names of s p e c i e s  cited 93  Page Table 1  Go-operators  and l o c a t i o n  Table 2  Distribution  of the seed  sources  44  Table 3  Height performance a t age e l e v e n  66  Table 4  Height performance.  Block I  69  Table 5  Height performance.  Block II  70  Table 6  Height performance.  Block I I I  71  Table 7  Height performance.  B l o c k IV  72  Table 8  Height performance between the years1964-1967  of the t e s t  sites  43  74  Page Figure 1  Diagram  of p l o t  designations  Figure 2  Blow-up of p l o t X i n p l a n t a t i o n Y  Figure 3  Geographic d i s t r i b u t i o n of seed sources i n the P a c i f i c Northwest D o u g l a s - f i r provenance t e s t  Figure 4  Ormerod's  Figure 5  L o c a t i o n map of the c o - o p e r a t i v e provenance t e s t 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  map  4l 42 L  5  63 68  I.  INTRODUCTION AND OBJECTIVES  In 1954 the Oregon S t a t e Board of F o r e s t r y met t o c o n s i d e r the p o s s i b i l i t y of s t u d y i n g the g e n e t i c v a r i a b i l i t y of the c o a s t a l form of D o u g l a s - f i r (Pseudotsuga m e n z i e s i i (Mirb.) Franco v a r . m e n z i e s i i ) from s i x t e e n sources i n Oregon, Washington  and B r i t i s h Columbia.  The f o l l o w i n g major reasons  made t h a t study a n e c e s s i t y 1 1. the importance of the provenance  problem,  2. the l a c k of knowledge on the performance of D o u g l a s - f i r provenances i n the P a c i f i c  Northwest.  The p r e s e n t t h e s i s was w r i t t e n w i t h two o b j e c t i v e s 1 t o review l i t e r a t u r e on the provenance  problem,  t o i l l u s t r a t e t h i s review by an a n a l y s i s of height measurements of the "Co-operative D o u g l a s - f i r provenance t e s t , " taken 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 .  II.  LITERATURE REVIEW ON PROVENANCE  No attempt i s made i n the f o l l o w i n g t o give an exhaustive  review on t h i s s u b j e c t , which i s f a r t o o v a s t t o be  condensed w i t h i n the scope o f t h i s t h e s i s .  An e x c e l l e n t survey  of p e r t i n e n t D o u g l a s - f i r provenance l i t e r a t u r e has r e c e n t l y been p u b l i s h e d by Haddock e t a l . (1967).  D e f i n i t i o n of Provenance A c c o r d i n g t o Wright (1962), provenance (or proveni e n c e ) means "the geographic pollen)."  source  of a l o t of seed  (or  T h i s d e f i n i t i o n p a r a l l e l s very c l o s e l y t h a t g i v e n by  Snyder (1959) and L i n e s (1967).  Lines statedi  synonymous w i t h p o p u l a t i o n .  term 'race* r e f e r s t o one  The  "Provenance i s or  more n a t u r a l p o p u l a t i o n s showing d e f i n e d c h a r a c t e r i s t i c s .  Prov-  enance does not c a r r y t h i s i m p l i c a t i o n , hence the broader provenance i s more g e n e r a l l y a p p l i c a b l e when p o p u l a t i o n s  term are  investigated." In the present t h e s i s "source" or " o r i g i n " w i l l used synonymously w i t h "provenance" t o a v o i d  The  be  repetition.  Provenance Problem Species e x h i b i t n a t u r a l v a r i a t i o n as a r e s u l t  of  e v o l u t i o n a r y a d a p t a t i o n t o d i f f e r e n t environments i n space and time.  A c c o r d i n g t o Haddock (1967)» the major f a c t o r s respon-  s i b l e f o r the development of l o c a l l y , g e n e t i c a l l y  different  p o p u l a t i o n s , a r e b e l i e v e d t o be  "the g r e a t topographic  a s s o c i a t e d c l i m a t i c and  v a r i a t i o n i n the environment."  edaphic  T h i s v a r i a t i o n , the nature and p l o r e d f o r any discontinuous  extent of which a r e h a r d l y  t r e e s p e c i e s , can be continuous (ecotypical).  and  ex-  ( c l i n a l ) or  When t r a n s p l a n t e d t o a new  en-  vironment w i t h i n or o u t s i d e the n a t u r a l range of the s p e c i e s , d i f f e r e n t geographic  sources w i l l r e a c t d i f f e r e n t l y .  f e r e n c e i n r e a c t i o n cannot be p r e d i c t e d i n d e t a i l .  This d i f Good per-  formance of a p a r t i c u l a r provenance i n one p l a c e i s no  assurance lieved  of I t s s u p e r i o r i t y elsewhere.  (e.g., S i l e n 1966)  I t i s g e n e r a l l y be-  t h a t l o c a l sources a r e b e s t  t o t h e i r p a r t i c u l a r environment and w i l l outperform  adapted  t r e e s of  n o n - l o c a l o r i g i n i n the l o n g run, e s p e c i a l l y i n areas w i t h o c c a s i o n a l c l i m a t i c extremes such as e x c e p t i o n a l f r o s t s . T r a n s f e r of seed should only be made between r e g i o n s showing as much c l i m a t i c s i m i l a r i t y as p o s s i b l e i n order t o a v o i d u n d e s i r able s i l v i c u l t u r a l r e s u l t s Larsen ( 1 9 5 6 ) ,  (Isaac 1 9 ^ 9 ) .  the f a t h e r of modern f o r e s t g e n e t i c s ,  does not agree w i t h the i d e a of s t r i c t l y a d h e r i n g t o c l i m a t i c a l l y s i m i l a r areas when t r a n s f e r r i n g seed. optimism  and  He preaches  bold  i l l u s t r a t e s h i s p o i n t w i t h the s u p e r i o r perform-  ance i n Denmark ( a t 5 6 °  l a t i t u d e North) of Norway spruce  a b l e s (L.) K a r s t ) from a Romanian source of d i s s i m i l a r c l i m a t e .  (PJLcea  (46° l a t i t u d e N o r t h ) ,  The f a c t t h a t n o n - l o c a l s u p e r i o r i t y  has  not y e t been proven f o r D o u g l a s - f i r does not n e c e s s a r i l y mean t h a t l o c a l provenances are always s u p e r i o r , i t merely  indicates  the l a c k of b o t h knowledge and e x p e r i m e n t a t i o n i n t h i s  field.  Some i n d i c a t i o n s of " n o n o p t i m a l i t y " of l o c a l sources f o r s e v e r a l s p e c i e s , i n c l u d i n g l o b l o l l y p i n e (Pinus taeda L . ) , were r e c e n t l y r e p o r t e d by Namkoong ( 1 9 6 9 ) *  A p p a r e n t l y evidence  Is  mounting t h a t an o p t i m a l growth zone of t h i s s p e c i e s e x i s t s a l o n g the s o u t h e a s t e r n border of i t s range.  Further experi-  ments w i l l prove whether seed from t h i s zone w i l l l o c a l stock  outproduce  elsewhere.  Wheat ( 1 9 6 6 ) l i s t e d s e v e r a l reasons  f o r there b e i n g  an important provenance problem w i t h D o u g l a s - f i r i n the  Pacific  Northwest.  They a r e b r i e f l y J  1. Lack of l o c a l seed i n s u f f i c i e n t q u a n t i t i e s due t o i r r e g u l a r seed c r o p s . 2. Urgency  of immediate  r e f o r e s t a t i o n due t o the  c a p a b i l i t y of h i g h s i t e l a n d s i n the P a c i f i c Northwest t o grow a dense cover of b r u s h as f a s t or f a s t e r than D o u g l a s - f i r would grow. 3. Lack o f s u f f i c i e n t knowledge on the a d a p t a b i l i t y o f other sources t o the l o c a l s i t u a t i o n , i . e . l a c k of f i e l d - t e s t e d seed t r a n s f e r  rules.  4. Lack of c o n t r o l s on seed movement. C o n s i d e r i n g the number of y e a r s i t takes t o grow a stand t o m a t u r i t y and the time t h a t might e l a p s e b e f o r e a provenance shows i t s i n f e r i o r i t y ,  i t i s e c o n o m i c a l l y and s i l v i -  c u l t u r a l l y unsound t o b l i n d l y buy a s e e d l o t of which the o r i g i n , and t h e r e f o r e performance, a r e not known.  I t i s b e t t e r i n such  a case n o t t o p l a n t b u t r a t h e r t o w a i t f o r good seed. Thus, i n p r a c t i c a l f o r e s t r y , the provenance  problem  i n essence i s t o f i n d p o p u l a t i o n s o f t r e e s , whether the s p e c i e s be indigenous or n o t , o f which the seed w i l l grow f o r e s t s a r e w e l l adapted t o t h e i r environment  that  and t h a t produce more  wood than t r e e s from any other p o p u l a t i o n of the same s p e c i e s d u r i n g the same p e r i o d of time.  The provenance  problem i s  P r o d u c t i v i t y may not simply be e q u a l t o r a p i d growth. In a p a r t i c u l a r provenance t e s t , the c r i t i c a l f a c t o r i n p r o d u c t i v i t y may be the a b i l i t y t o w i t h s t a n d f r o s t ( e . g . S i t k a spruce i n Western Norway) or the a b i l i t y t o produce f e r t i l e seed (e.g. Scots p i n e (Plnus s y l v e s t r l s L.) i n Northern F i n l a n d ( L i n e s 1967).  i n t i m a t e l y connected  w i t h the problem of f i n d i n g the b e s t  spe-  cies f o r a particular area. T e c h n i c a l problems such as s t a t i s t i c a l d e s i g n , l a y o u t and c o n t r o l of provenance experiments have been thoroughly cussed by Edwards ( 1 9 5 6 ) and L i n e s  dis-  (1967).  A c c o r d i n g t o Schmidt ( 1 9 6 2 a ) ,  provenance t r i a l s  can  be d i v i d e d i n t o the f o l l o w i n g c a t e g o r i e s ! 1.  S t u d i e s of broad  geographic  variation.  S t u d i e s of broad geographic v a r i a t i o n are not s p e c i f i c a l l y designed t o p r o v i d e d a t a f o r seed t r a n s f e r r u l e s ; they u s u a l l y i n v o l v e a s m a l l number of provenances which may not be a s u f f i c i e n t l y r e p r e s e n t a t i v e sample of a g i v e n s p e c i e s . T h i s type of study u s u a l l y i n d i c a t e s t h a t l o c a l seed i s b e s t , a l t h o u g h t h a t might not always be c o r r e c t . 2.  T e s t s of s t o c k from improved seed p r o d u c t i o n a r e a s . S t u d i e s i n v o l v i n g stock from seed p r o d u c t i o n areas can be c r i t i c i z e d on the b a s i s t h a t the seed p r o d u c t i o n areas may no l o n g e r be f u n c t i o n i n g by the time the r e s u l t s a r e obtained and t h a t they u s u a l l y i n v o l v e too few provenances.  3.  Comprehensive  trials.  Comprehensive t r i a l s r e q u i r e much ground work, such as c l i m a t i c s t r a t i f i c a t i o n of the range of the s p e c i e s i n q u e s t i o n . Such a s t r a t i f i c a t i o n i n i t s e l f i s an overwhelming t a s k i n some a r e a s . R e c i p r o c a l p l a n t a t i o n s have t o be e s t a b l i s h e d w i t h i n each zone. Seed samples may be too few t o adequately capture the v a r i a t i o n w i t h i n the species. 4. F a c t o r i a l  tests.  The i n f l u e n c e of the i n d i v i d u a l c l i m a t i c (and geographic) v a r i a b l e s c o n s i d e r e d t o be of major importance t o e c o t y p i c d i f f e r e n t i a t i o n are determined by f a c t o r i a l t e s t s which are of g r e a t v a l u e i n areas where c l i m a t e shows l i t t l e , or a t l e a s t , p r e d i c t a b l e v a r i a t i o n w i t h changes i n geographic f a c t o r s such as l a t i t u d e .  5. Random p o p u l a t i o n  trials.  Random p o p u l a t i o n t r i a l s i n v o l v e a l a r g e number of randomly p i c k e d provenances and a l a r g e numb e r of o u t p l a n t i n g s i t e s r e p r e s e n t a t i v e of the c l i m a t i c v a r i a t i o n w i t h i n the range of the s p e c i e s . Although some c l i m a t i c d a t a are r e q u i r e d b e f o r e e s t a b l i s h i n g the t e s t t o a s s i s t i n s e l e c t i n g the o u t p l a n t i n g s i t e s , the b u l k of the i n f o r m a t i o n can be accumulated d u r i n g the s t u d y . T h i s approach was adopted by Schmidt ( 1 9 6 7 ) t o study the v a r i a t i o n w i t h i n the B r i t i s h Columbia c o a s t a l form of D o u g l a s - f i r . Provenance t r i a l s should not be mistaken f o r progeny t e s t s which f o l l o w them c h r o n o l o g i c a l l y . determine the g e n e t i c worth of p o p u l a t i o n s  Provenance s t u d i e s of t r e e s ; progeny  t e s t s determine the g e n e t i c worth of an i n d i v i d u a l t r e e  (Lines  1967).  Objectives  of Provenance Out  of the p r e v i o u s  o b j e c t i v e s can be 1.  Tests paragraphs, the f o l l o w i n g major  definedt  To study extent and  nature of the v a r i a t i o n i n p a r t s  or i n the e n t i r e range of a s p e c i e s populations  2.  to l o c a t e  of t r e e s , the seed of which w i l l  w e l l adapted, p r o d u c t i v e ( L i n e s 1967  i n order  and  Place  f o r e s t s i n a given  produce region  1969).  To d e f i n e the g e n e t i c and  environmental components  of t h i s phenotypic ( i . e . m o r p h o l o g i c a l and/or p h y s i o l o g i c a l ) v a r i a b i l i t y between t r e e s from d i f f e r e n t geographic sources ( L i n e s 3.  1967)*  To e s t a b l i s h seed t r a n s f e r r u l e s (Schmidt 1 9 6 7 ) .  Provenance Tests and F o r e s t Tree Improvement A c c o r d i n g t o Nanson (1964a), t r e e improvement programs b a s i c a l l y aim a t three 1. q u a n t i t y  pointsi  ( o f wood and/or other organic  products),  2. q u a l i t y ( o f v i s i b l e , e.g. stem s t r a i g h t n e s s  and/or  i n v i s i b l e c h a r a c t e r i s t i c s such as s p e c i f i c g r a v i t y ) , 3. r e s i s t a n c e  ( t o b i o t i c environment, t o s o i l and t o  climate)• The  f o l l o w i n g t o o l s can be used t o r e a c h these  objectivesi 1. Provenance  studies  2. Seed p r o d u c t i o n  areas  3. Seed orchards 4. I n t e r - and i n t r a - s p e c i f i c h y b r i d i z a t i o n 5. Induced mutations and p o l y p l o i d y 6. V e g e t a t i v e  propagation.  A c c o r d i n g t o Nanson (1964a), provenance t e s t s should be designed s t a t i s t i c a l l y  i n such a way t h a t growth performance,  form and r e s i s t a n c e of the v a r i o u s can be a s s e s s e d . step i n genetic  o r i g i n s under  Investigation  Provenance t e s t s c o n s t i t u t e the f i r s t research.  logical  A l t h o u g h a r e l a t i v e l y easy endeavour,  provenance t e s t s a r e a b l e t o y i e l d p r a c t i c a l r e s u l t s q u i c k l y . They o f t e n show without a d d i t i o n a l c o s t , which stand or r e g i o n t o buy seed from.  B r i e f H i s t o r y of Provenance The  Testing  Frenchman de V i l m o r l n  i s traditionally  considered  t o be t h e f a t h e r o f provenance t e s t i n g .  I n the e a r l y 1820's  he e s t a b l i s h e d Scots pine p l a n t a t i o n s of known geographic o r i g i n a t his estate a t l e s Barres.  I n 1862 he wrote a r e p o r t  on the growth of these t r e e s and concluded t h a t there was a d e f i n i t e d i f f e r e n c e i n s e v e r a l important c h a r a c t e r i s t i c s between t r e e s from d i f f e r e n t r e g i o n s .  Unfortunately,  c o v e r i e s went unheeded (Larsen 1 9 5 6 , Revel C i e s l a r ( 1 9 0 7 ) and E n g l e r existence spruce. pioneer  these d i s -  i960).  ( 1 9 1 3 ) demonstrated the  o f g r e a t d i f f e r e n c e s between v a r i o u s o r i g i n s o f Norway The 1912 D o u g l a s - f i r h e r e d i t y study c o n s t i t u t e s the  e f f o r t i n provenance t e s t i n g i n North America.  I t was  i n i t i a t e d by the U n i t e d S t a t e s F o r e s t S e r v i c e which a l s o prov i d e d the seed f o r the D o u g l a s - f i r t e s t s s t a r t e d by Schwappach and  Muench around 1912 i n Germany.  The f i r s t  t e n t a t i v e seed  t r a n s f e r r u l e s were e s t a b l i s h e d i n Sweden i n the 1 9 3 0 s . ,  With the growing awareness o f the importance o f the problem and the p o t e n t i a l gains t o be d e r i v e d from the s e l e c t i o n o f adequate-seed sources, more popular  provenance t e s t s became more and  i n f o r e s t r e s e a r c h around the w o r l d .  They a r e t o o  numerous t o be r e p o r t e d here i n d e t a i l .  Problems Commonly The  Investigated  f o l l o w i n g i s an a r b i t r a r y d i s t i l l a t i o n of l i t e r -  a t u r e f o r some major a u t e c o l o g i c a l a s p e c t s  o f the provenance  problem, w i t h s p e c i a l r e f e r e n c e t o D o u g l a s - f i r .  Drought  resistance A c c o r d i n g t o F e r r e l l and Woodland (1966), drought r e -  s i s t a n c e may  i n v o l v e e i t h e r drought avoidance o r drought h a r d i -  ness, or b o t h .  Drought avoidance i s accomplished through r o o t  e x t e n s i o n and r e d u c t i o n i n l e a f a r e a or through e a r l y onset of dormancy.  Drought h a r d i n e s s i s the a b i l i t y  t o s u r v i v e i n a dry  e x t e r n a l environment c a u s i n g severe t i s s u e d e h y d r a t i o n .  The  authors s t u d i e d the e f f e c t of seed o r i g i n on drought r e s i s t a n c e of D o u g l a s - f i r and found t h a t I n t e r i o r mountain  provenances  showed s i g n i f i c a n t l y g r e a t e r drought r e s i s t a n c e than those from areas west of the Cascades. notable exception.  S e e d l i n g s from C o r v a l l i s were a  They proved no more drought r e s i s t a n t than  s e e d l i n g s from h i g h e r p r e c i p i t a t i o n , lower temperature areas elsewhere west of the Cascades. By two drought h a r d i n e s s t e s t s , time t o death and s o i l moisture content a t the death p o i n t , P h a r i s and  Ferrell  (1966) showed t h a t D o u g l a s - f i r s e e d l i n g s from t h r e e c o a s t a l sources were l e s s drought r e s i s t a n t than those from f i v e sources.  inland  They found needle moisture t o be a good index f o r de-  t e r m i n i n g p l a n t v i a b i l i t y under w e l l watered c o n d i t i o n s . l i n g s c o u l d be c l a s s i f i e d  Seed-  i n t o c o a s t a l and i n l a n d groups on the  b a s i s of t h e i r needle moisture content w i t h the e x c e p t i o n o f an A r i z o n a provenance behaving l i k e the c o a s t a l  group.  S i t e ecotypes A b a s i c t e n e t of g e n e t i c s holds t h a t the phenotype i s a f f e c t e d by the genotype and the environment.  Provenance  s t u d i e s a r e designed t o enable the r e s e a r c h e r g e n e t i c d i f f e r e n c e s between v a r i o u s  t o d e t e c t the  seed s o u r c e s .  T h i s can  b e s t be done by growing the t r e e s on a h o p e f u l l y uniform under the same environmental c o n d i t i o n s .  site  Since s i t e i s one of  the major components o f the environmental complex, i t . i s n o t s u r p r i s i n g t h a t some f o r e s t e r s have t r i e d t o f i n d out whether populations particular  had adapted themselves g e n e t i c a l l y t o growing on a site.  The  f i r s t authors t o r e p o r t on t h i s problem i n  D o u g l a s - f i r were Munger and Morris age  (1936) who found t h a t the  o f the parent t r e e , i t s growing space, i t s c o n d i t i o n s as t o  fungus i n f e c t i o n and i t s s i t e index, had no e f f e c t upon h e i g h t growth of the progeny.  Gathy (19^1 and 19&7) t e s t e d  various  provenances of c o a s t a l D o u g l a s - f i r and came t o s i m i l a r c o n c l u sions »  age and growing s i t e of the parent t r e e s seemed t o have  no b e a r i n g  on height  growth of t h e i r o f f s p r i n g .  found no 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 h i g h e l e v a t i o n D o u g l a s - f i r provenances. been c o l l e c t e d from a stand  Heaman (1968)  growth between two  One l o t o f seed had  showing r e l a t i v e l y good phenotypic  c h a r a c t e r i s t i c s on a good s i t e w h i l e the other l o t o r i g i n a t e d from t r e e s growing on a much poorer s i t e , b u t i n a s i m i l a r climatic  Aspect  zone.  races Although there has been no d e f i n i t e proof  so f a r of  s i t e ecotypes i n D o u g l a s - f i r , i t i s q u i t e i n t e r e s t i n g t o note t h a t ecotypes have evolved  t h a t a r e adapted t o the p a r t i c u l a r  c o n d i t i o n s p r e v a i l i n g on s o u t h e r l y a s p e c t s . land  F e r r e l l and Wood-  ( 1 9 6 6 ) found t h a t s e e d l i n g s produced from t r e e s growing on  a south s l o p e were more drought r e s i s t a n t than those from t r e e s growing on a n o r t h s l o p e , a s h o r t d i s t a n c e away. r e s u l t s were obtained  Interesting  by Hermann and Lavender ( 1 9 6 8 )  i n a growth  chamber experiment w i t h D o u g l a s - f i r from v a r i o u s a l t i t u d e s and a s p e c t s i n southern Oregon.  Progeny o f t r e e s from south f a c i n g  a s p e c t s e x h i b i t e d a s h o r t e r growing p e r i o d and l a r g e r r o o t s i n r e l a t i o n t o t h e i r tops than s e e d l i n g s facing slopes.  from parents on n o r t h  T h i s was i n t e r p r e t e d t o be a r e s u l t of n a t u r a l  s e l e c t i o n f o r e a r l y c e s s a t i o n o f growth i n h a b i t a t s t h a t a r e p a r t i c u l a r l y d r y i n summer.  These r e s u l t s a r e an I n t e r e s t i n g  p a r a l l e l t o e a r l i e r f i n d i n g s by S q u i l l a c e and Bingham suggesting pine  Bud  the e x i s t e n c e  (Pinus m o n t l c o l a  of aspect  (1958)  ecotypes i n western white  Dougl.).  bursting Bud  b u r s t i n g , as d i s c u s s e d  i n t h i s chapter,  refers to  the opening of v e g e t a t i v e buds only and i s synonymously used with " f l u s h i n g . " 1.  The i n f l u e n c e of g e n e t i c c o n t r o l on the t i m i n g o f f l u s h i n g . Observations by Morris  e t a l . (1957) suggest  strong  g e n e t i c c o n t r o l over time of bud b u r s t i n g i n Douglasfir.  The most c l a s s i c a l study on t h i s q u e s t i o n  made by S i l e n ( 1 9 6 2 ) ,  who t r i e d t o minimize  was  environ-  mental i n f l u e n c e s by g r a f t i n g s e v e r a l r e p l i c a t i o n s o f s c i o n s from t r e e s w i t h l a r g e , known d i f f e r e n c e s i n  f l u s h i n g dates onto limbs o f seven t r e e s o f a seventeen year o l d clone.  As a c o n t r o l , a limb o f the c l o n a l  t r e e was c u t and r e - g r a f t e d .  S i l e n estimated the gen-  e t i c component of the bud b u r s t i n g t r a i t a t n i n e t y - f o u r and  n i n e t y - s i x percent of the t o t a l v a r i a t i o n f o r the  two  y e a r s of o b s e r v a t i o n s .  A t the same time he p r e -  sented evidence t h a t l o c a l environmental d i f f e r e n c e s may c o n s i s t e n t l y d e l a y bud b u r s t i n g of g e n e t i c a l l y s i m i l a r m a t e r i a l f o r as much as two weeks. a t i o n s by the United Griffith  States  Similarly, public-  Forest Service  (1964) and  (1968) show evidence o f s t r o n g g e n e t i c  over f l u s h i n g i n D o u g l a s - f i r .  The t r e e s under obser-  v a t i o n f o r several years c o n s i s t e n t l y flushed same order, w i t h i n a day o r two. altltudlnal. pattern  control  i n the  The p a t t e r n was n o t  Walters and Ching (1969) s t u d i e d the  of bud b u r s t  testi? t o be r e p o r t e d  i n the D o u g l a s - f i r  provenance  l a t e r I n t h i s t h e s i s , and found  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 enance t o be a l a t e f l u s h e r , r e g a r d l e s s  prov-  of p l a n t a t i o n ;  the Salem source proved t o be the e a r l i e s t f l u s h e r o f t e r m i n a l and l a t e r a l buds, r e g a r d l e s s  of p l a n t i n g  Evidence was put forward f o r a s t r o n g  i n f l u e n c e of  l o c a l climatic conditions 2.  area.  on bud b u r s t i n g .  The e f f e c t s of t h e environment on f l u s h i n g . The  p u b l i c a t i o n s c i t e d I n the p r e v i o u s c h a p t e r , s t a t i n g  t h a t under u n i f o r m environmental c o n d i t i o n s bud b u r s t ing  i s under v e r y s t r o n g g e n e t i c  c o n t r o l , a l s o mention  the e f f e c t of the environment, p a r t i c u l a r l y the matic c o n d i t i o n s expression served  of any  p a r t i c u l a r year, on  of t h i s t r a i t .  Morris  occur as much  as a month l a t e r than i n other y e a r s .  weeks t o two Griffith  the  et a l . (1957) ob-  t h a t i n some y e a r s f l u s h i n g may  a t i o n p l a n t a t i o n s , buds may  In h i g h  (1968) t r i e d t o c o r r e l a t e date of f l u s h i n g found t h a t i n the  University bursting  i n f l u e n c e d by weather c o n d i t i o n s d u r i n g the  The  two  elevations.  of B r i t i s h Columbia Research F o r e s t bud  3.  elev-  open anywhere from  months l a t e r than a t low  w i t h c l i m a t i c d a t a and  day  cli-  p e r i o d p r i o r t o May  was  forty-two  the s i x t h .  disadvantages of e a r l y bud  bursting.  Time of f l u s h i n g i s important l n r e l a t i o n t o s p r i n g frosts.  Since bud b u r s t i n g i s a h i g h l y  hereditary  t r a i t , seed from e a r l y f l u s h e r s cannot be p l a n t e d advantage i n areas where l a t e f r o s t s are l i k e l y occur.  Irgens-M^ller  to  (19&7) n o t i c e d a d i f f e r e n c e i n  time of bud b u r s t i n g of f i f t e e n t o eighteen  days  tween p l a n t s o r i g i n a t i n g from areas separated twenty t o f o r t y m i l e s .  He  by  spp.)  spring f r o s t s .  were found t o t h r e a t e n  f i r trees  ( M i t c h e l l and  only  un-  Needle midges ( C o n t a r l n l a e a r l y f l u s h i n g Douglas-  Nagel, 1969). w h i l e l e a v i n g  l a t e f l u s h e r s more or l e s s unharmed. advised  be-  s t r e s s e d the importance of  choosing a seed source t h a t can be expected to be a f f e c t e d by  to  Schober (19&3)  a g a i n s t D o u g l a s - f i r provenances from B r i t i s h  Columbians I n t e r i o r Wet B e l t f o r use i n areas w i t h l a t e f r o s t s or w i d e l y f l u c t u a t i n g weather c o n d i t i o n s during  s p r i n g , such as occur i n Western Europe,  because they f l u s h e a r l i e r than c o a s t a l provenances. T h i s e a r l i e r f l u s h i n g makes them more s u s c e p t i b l e t o the D o u g l a s - f i r needle b l i g h t (Rhabdocllne pseudotsugae Syd.). ions.  Haddock e t a l . (19^7) came t o s i m i l a r c o n c l u s To summarize, e a r l y f l u s h i n g i s h a r d l y an a s s e t ,  s i n c e s p r i n g f r o s t s , i n s e c t s and f u n g i may damage the trees 4.  severely.  Implications  o f time o f f l u s h i n g on the provenance problem.  When choosing seed l o t s , geographic and c l i m a t i c d a t a should be supplemented w i t h c a r e f u l observations date o f bud b u r s t i n g c o v e r i n g  several years.  on  The suc-  cess o f whole p l a n t a t i o n s may hinge on time of f l u s h i n g i n connection  with s p r i n g f r o s t s , insects or f u n g i .  Late f l u s h i n g e l i m i n a t e s a l l these dangers and has no negative  e f f e c t on t o t a l h e i g h t growth.  I d e a l proven-  ances a r e those t h a t grow f a s t , f l u s h l a t e and s e t t h e i r buds e a r l y , thus a v o i d i n g b o t h s p r i n g f r o s t s and fall  frosts.  Lammas shoots According and  t o Walters and Soos ( 1 9 6 l b ) ,  " ... l a t e r a l  t e r m i n a l v e g e t a t i v e buds of young t r e e s f r e q u e n t l y break  dormancy In l a t e summer and produce e x t r a - s e a s o n a l lammas shoots a f t e r the o l d E n g l i s h harvest  shoots named  f e s t i v a l s o f Lammas  on August 1 . "  The authors s t u d i e d t h i s phenomenon on Douglas-  f i r s e e d l i n g s from v a r i o u s o r i g i n s grown a t s e v e r a l  elevations  on the U n i v e r s i t y o f B r i t i s h Columbia Research F o r e s t .  The  c o n c l u s i o n s were t h a t lammas growth 1 ) was i n f l u e n c e d more by environmental than g e n e t i c factors, 2) decreased w i t h i n c r e a s i n g age, 3) i n c r e a s e d as s i t e q u a l i t y  increased,  4) i n c r e a s e d as c u r r e n t h e i g h t growth i n c r e a s e d f o r s e e d l i n g s of a p a r t i c u l a r provenance. Lammas growth was sometimes r e f l e c t e d l n f a l s e annual rings.  Forked l e a d e r s caused by lammas shoots were overcome  quite rapidly i n Douglas-fir. Sweet ( 1 9 6 5 )  found t h a t D o u g l a s - f i r s e e d l i n g s h a v i n g  more than one annual growth f l u s h i n t h e i r second year do n o t make a g r e a t e r h e i g h t increment than provenances w i t h fewer such t r e e s .  Hoffman ( 1 9 6 5 )  experienced other r e s u l t s with  Norway spruce, where the i n c i d e n c e o f lammas shoots v a r i e s geog r a p h i c a l l y and decreases w i t h i n c r e a s i n g e l e v a t i o n o f seed source as an a d a p t a t i o n t o the l o c a l c l i m a t e .  For s i t e s with  no e a r l y f r o s t s , Hoffman recommended provenances w i t h a h i g h occurrence of lammas shoots t h a t would outgrow dangers i l e stages more q u i c k l y .  Schmidt-Vogt  of juven-  (1966) a l s o noticed  that  lammas growth decreases w i t h i n c r e a s i n g a l t i t u d e of seed source i n Norway spruce, and he found i t s occurrence on one and twoyear o l d s e e d l i n g s t o be a workable  e a r l y t e s t t o check on  e l e v a t i o n a l a u t h e n t i c i t y of seed. To conclude, lammas shoots c o n s t i t u t e a disadvantage  on s i t e s where e a r l y f r o s t s are common, s i n c e the s o f t shoots can be damaged e a s i l y by t r e e s of poor form may  frost.  I f t h i s happens  result.  repeatedly,  On s i t e s f r e e of e a r l y f r o s t  hazards, lammas shoots are an advantage In overcoming competi t i o n from l e s s e r v e g e t a t i o n ,  Frost  and  p o s s i b l y from deer browsing.  resistance Schoenbach ( 1 9 5 8 ) evaluated  s e l e c t i o n of D o u g l a s - f i r s e e d l i n g s s e l e c t i o n had i v e , the  g i v e n two  f r o s t r e s i s t a n c e has  for frost resistance.  populationsi  other f r o s t r e s i s t a n t .  the e f f e c t s of a mass  one  mainly f r o s t s e n s i t -  Schoenbach concluded  s i n g l e gene i n h e r i t a n c e and  homozygous c o n d i t i o n .  He  The  that  appears i n a  s t r e s s e d the need t o s t a r t provenance  s t u d i e s on a v e r y l a r g e b a s i s , t h a t i s , I n c l u d i n g as many seed sources as p o s s i b l e , i n order to f i n d p o p u l a t i o n s r e s i s t a n c e genes.  A c c o r d i n g t o Scheumann ( 1 9 6 5 ) » the  f i v e p o i n t s have t o be be  containing  considered  following  when f r o s t r e s i s t a n c e i s t o  studied! 1.  readiness  t o harden o f f (= e a r l y f r o s t r e s i s t a n c e ) .  2 . e x t e n t of hardening o f f (= w i n t e r f r o s t r e s i s t a n c e ) . 3.  s t a b i l i t y of dormancy (Important d u r i n g w i n t e r s w i t h h i g h l y v a r y i n g temperatures).  4.  time of f l u s h i n g and  5.  regeneration damage)•  flowering  (late frost resistance).  p o t e n t i a l ( a b i l i t y t o overcome f r o s t  These f i v e p o i n t s make i t q u i t e obvious t h a t t - f r o s t r e s i s t a n c e i s a dynamic p r o c e s s t h a t cannot be s t u d i e d as a s t a t i c c o n d i t i o n , as has been done traditionally.  - t h e r e cannot "be a s i n g l e t e s t w i t h which t o evaluate a l l a s p e c t s of f r o s t r e s i s t a n c e i n plants * Previous based on f i e l d  knowledge on t h i s s u b j e c t has always been  observations  Scheumann ( 1 9 6 5 ) d e v i s e d extent  and  a f t e r extremely c o l d weather.  a l a b o r a t o r y t e s t w i t h which  s t a b i l i t y of h a r d e n i n g - o f f  t r o l l e d temperature c o n d i t i o n s .  can be  Seedlings,  readiness,  s t u d i e d under conneedles or young  twigs can be used l n t h i s t e s t where temperature i s g r a d u a l l y lowered t o below f r e e z i n g p o i n t . and  extent  ature  of h a r d e n i n g - o f f  Is r e p e a t e d l y  p o i n t and mancy.  After early frost  have been e v a l u a t e d ,  the  r a i s e d f o r some time above the  then lowered a g a i n  i n order  resistance temper-  freezing  to t e s t s t a b i l i t y  T h i s i s a method t o r a p i d l y mass s e l e c t and  of dor-  rank v a r i o u s  provenances (or progenies f o r t h a t matter) f o r s e v e r a l a s p e c t s of f r o s t r e s i s t a n c e . D o u g l a s - f i r and  Scheumann t r i e d t h i s  " s i m p l i f i e d t e s t " on  on European l a r c h ( L a r i x decldua M i l l ) .  He  found s i g n i f i c a n t d i f f e r e n c e s i n f r o s t r e s i s t a n c e of two-year o l d D o u g l a s - f i r hybrids  (glauca x v i r l d i s ) .  a t i o n s between the c l i m a t e  He  found c o r r e l -  i n which the l a r c h t r e e s grew and  the b e h a v i o u r of the twigs i n the f r o s t r e s i s t a n c e t e s t . Scheumann concluded t h a t t h i s p o s s i b i l i t y of f i n d i n g g e n e t i c d i f f e r e n c e s i n f r o s t r e s i s t a n c e among p l u s - t r e e s and  t h e i r pro-  genies g i v e s new  resist-  hope f o r s u c c e s s f u l l y t e s t i n g f r o s t  ance i n f o r e s t t r e e s .  Schoenbach and  on the same glauca x v i r l d i s h y b r i d s .  Bellmann ( 1 9 6 7 ) The  hybrids were a l l  s i g n i f i c a n t l y more f r o s t r e s i s t a n t than the Since  reported  they were a l s o growing f a s t e r than the  "green"  parents.  "blue" v a r i e t y ,  the h y b r i d s were recommended by Schoenbach and the c o l d c l i m a t e s  Bellmann f o r  of western Europe's mountains.  In a study i n v o l v i n g t h i r t y - o n e seed sources of c o a s t a l D o u g l a s - f i r from Washington and  Oregon, Gathy  found three Washington provenances from medium and ( Y a c o l t , 200  ations 400  f e e t , C a s t l e Rock, 1300  low  f e e t , and  (1961) elevForks,  f e e t above sea l e v e l ) t o be most f r o s t r e s i s t a n t and  fore best  s u i t e d f o r the maritime c l i m a t e  of Belgium.  there-  Nanson  ( 1 9 6 4 b ) s t u d i e d the e f f e c t of the severe w i n t e r of 1 9 6 2 - 1 9 6 3 on c o a s t a l D o u g l a s - f i r provenances i n Belgium, but confirm  could  not  the s u p e r i o r i t y of Washington sources over Oregon  gins In terms of f r o s t r e s i s t a n c e .  According  ori-  t o Nanson, a l l  provenances of c o a s t a l D o u g l a s - f i r t e s t e d r e q u i r e adequate prot e c t i o n from severe f r o s t , e s p e c i a l l y when they are young. Lacaze ( 1 9 6 4 ) examined D o u g l a s - f i r s e e d l i n g s the 1 9 6 2 - 1 9 6 3 "deep f r e e z e " and  i n France a f t e r  found c o n s i d e r a b l e  variation in  f r o s t r e s i s t a n c e which g e n e r a l l y i n c r e a s e d w i t h i n c r e a s i n g  lat-  i t u d e and  points  49°  a l t i t u d e of the seed source.  l a t i t u d e North, 1800  f e e t e l e v a t i o n , roughly  r e s i s t a n t from the n o n - r e s i s t a n t Stem  (1966),  f i r seedlings  reviewing  A l i n e j o i n i n g the  provenances.  divides  the  In a note by  r e s u l t s of a r e c e n t experiment, Douglas-  from A r i z o n a ,  New  Mexico and  Mexico were found t o  be more f r o s t r e s i s t a n t than provenances from c o a s t a l B r i t i s h Columbia, Washington and and  Oregon, t r a d i t i o n a l l y t e s t e d i n Europe  recommended by Schober ( 1 9 6 3 ) .  the 1912  Silen (1966),  evaluating  D o u g l a s - f i r h e r e d i t y study (see below) a f t e r the  November f r o s t , formulated  1955  the hypothesis t h a t l o c a l provenances  have adapted  themselves  over the c e n t u r i e s t o long-term  weather  extremes and a r e t h e r e f o r e l e s s damaged by severe f r o s t s  than  are n o n - l o c a l s o u r c e s .  C l i n e or  ecotype? In 1936  L a n g l e t p u b l i s h e d an a r t i c l e on the p h y s i o -  l o g i c a l v a r i a b i l i t y and i n Sweden.  i t s r e l a t i o n t o c l i m a t e f o r Scots pine  The dry matter content of needles proved  t o be  h i g h l y c o r r e l a t e d w i t h the number of days d u r i n g the growing of s i x degrees  (43°  season w i t h an average  temperature  Celsius  F a h r e n h e i t ) , or more.  L a n g l e t showed a n o r t h - s o u t h v a r i a t i o n  i n Scots pine i n Sweden and claimed t h i s v a r i a t i o n t o be tinuous.  Wright and Baldwin  ( 1 9 5 7 ) based  con-  their criticism  of  L a n g l e t * s a r t i c l e on o b s e r v a t i o n s from a Scots p i n e provenance t e s t i n New  Hampshire.  They agreed  t h a t there i s a n o r t h - s o u t h  v a r i a t i o n i n Scots p i n e , but t h e i r s t a t i s t i c a l a n a l y s i s  indic-  a t e d t h a t most of the geographic v a r i a t i o n i s d i s c o n t i n u o u s , not c l i n a l . r u l e s based  T h i s would mean t h a t the Swedish seed  transfer  on the c l i n a l hypothesis and l i m i t i n g cone c o l l e c t -  i o n s t o areas w i t h i n 250 k i l o m e t e r s (approximately 150 n o r t h or south and 300 meters (approximately 1000  miles)  feet) d i f f e r -  ence l n e l e v a t i o n from the p l a n t i n g s i t e , should be r e v i s e d t o c o l l e c t i n g seed w i t h i n the boundaries ed f o r the p l a n t i n g a r e a .  T h i s may  of the ecotype b e s t  suit-  permit the s a f e t r a n s f e r of  seed f o r s e v e r a l hundred m i l e s or l i m i t i t t o a few m i l e s . Langlet  ( 1 9 5 9 ) used Wright and Baldwin's  d a t a t o show where  they had gone wrong and he s t a t e d t h a t v a r i a b i l i t y c o n t i n u e s i n  the same degree as the d e t e r m i n i n g environmental  f a c t o r s vary-  continuously. N e i t h e r the c l i n a l n o r the e c o t y p i c a l h y p o t h e s i s has been d l s p r o v e n .  I t i s p o s s i b l e t h a t In i s o l a t e d  Scots pine has developed  occurrences  ecotypes t h a t cannot be e x p l a i n e d by  d i f f e r e n c e s i n the environment, whereas i n l a r g e , areas o f i t s v a s t range the v a r i a t i o n i s c l i n a l , g r a d u a l changes i n the environment.  contiguous reflecting  In o t h e r words, ecotypes  a r e n o t n e c e s s a r i l y In c o n t r a s t w i t h c l i n a l v a r i a t i o n . e t a l . (1967) mentioned the v a r i a b i l i t y  Haddock  o f ponderosa pine  (Plnus  ponderosa Laws.) s t u d i e d by v a r i o u s a u t h o r s , some o f whom support the c l i n a l , some the e c o t y p i c a l h y p o t h e s i s .  Haddock  e t a l . concluded t h a t no matter which a l t e r n a t i v e one s u b s c r i b e s to, " ... one cannot d i s p e n s e w i t h a thorough knowledge of the geography of a r e g i o n and an u n d e r s t a n d i n g of the i n f l u e n c e of topography on l o c a l c l i m a t e , e s p e c i a l l y i n r e g i o n s as c o n t i n u o u s l y mountainous as southern B r i t i s h Columbia."  Climate S i n c e weather measurements can only i n d i c a t e p a r t o f the complex c a l l e d c l i m a t e , and s i n c e the e s t a b l i s h m e n t and maintenance of weather s t a t i o n s i n remote areas a r e c o s t l y , Schmidt (1962b) t r i e d t o I n t e r p r e t c l i m a t e from p h e n o l o g i c a l observations.  F o r t h i s purpose,  time of i n i t i a l p o l l e n r e l e a s e  i n D o u g l a s - f i r was chosen, because i t i s the f i r s t  external i n -  d i c a t i o n of p h y s i o l o g i c a l a c t i v i t y i n s p r i n g and i t i s easy t o  a s s e s s , compared f o r i n s t a n c e w i t h d e t e r m i n i n g the s t a r t of cambial a c t i v i t y .  The r e s u l t s were not encouraging, and they  r a i s e d f u r t h e r problems because male bud development  does not  seem t o be r e l a t e d t o a simple e x p r e s s i o n of temperature. Irgens-Mjrfller ( 1 9 6 5 ) q u e s t i o n e d the v a l u e of c l i m a t i c d a t a l n a s s i g n i n g o f f - s o u r c e seed t o p l a n t i n g s i t e .  Both p l a n t -  i n g l o c a t i o n and seed source may be f a r away from the n e a r e s t weather s t a t i o n .  In a d d i t i o n , some provenances  may  tolerate a  wide v a r i e t y of c o n d i t i o n s , others might be narrowly adapted t o specific conditions.  ( 1 9 6 6 ) d i v i d e d the  Haddock and S z i k l a i  range of D o u g l a s - f i r l n B r i t i s h Columbia and western A l b e r t a i n t o n i n e seed c o l l e c t i o n zones based p r i m a r i l y on the ence of c l i m a t e on geographic v a r i a b i l i t y the d i s t r i b u t i o n of a s s o c i a t e d s p e c i e s .  influ-  of p o p u l a t i o n s and Both authors were  aware o f the m e r i t s and l i m i t a t i o n s of such a z o n a t i o n , and admitted t h a t p o p u l a t i o n s w i t h i n the i n d i v i d u a l zones a r e f a r from b e i n g homogeneous, s i n c e remarkable c l i m a t i c  differences  due t o e l e v a t i o n , topography and p r e c i p i t a t i o n e x i s t w i t h i n these zones.  I t i s not s u f f i c i e n t f o r a f o r e s t e r t o i n d i c a t e  the g e n e r a l zone from which he wants seed.  He should p r o v i d e  d e t a i l e d c l i m a t i c d a t a f o r the p l a n t a t i o n s i t e ,  including at  l e a s t mean annual temperature, a b s o l u t e minimum temperature, mean annual p r e c i p i t a t i o n d u r i n g growing season, and l e n g t h of f r o s t f r e e p e r i o d or dates of e a r l i e s t and l a t e s t f r o s t s . way  the seed c o l l e c t o r may  have a b e t t e r chance  s u i t a b l e seed (Haddock and S z i k l a i ,  This  of p r o v i d i n g  1966).  Newnham ( 1 9 6 8 ) p u b l i s h e d an a r t i c l e on the  c l a s s i f i c a t i o n of c l i m a t e  and  i t s r e l a t i o n s h i p to tree  species,  i n c l u d i n g D o u g l a s - f i r , u s i n g d a t a from seventy weather s t a t i o n s from many p a r t s analysis  of B r i t i s h Columbia,  on the m a t r i x of c o r r e l a t i o n c o e f f i c i e n t s w i t h n i n e -  teen v a r i a b l e s r e c o r d e d by three new  With p r i n c i p a l component  the weather s t a t i o n s , he  v a r i a b l e s a c c o u n t i n g f o r ninety-two p e r c e n t of  t o t a l v a r i a t i o n between weather s t a t i o n s . was  computed  a g e n e r a l Index of w i n t e r and  of the growing season, the tween s p r i n g and  The  f a l l climate  first and  second r e p r e s e n t e d the  summer temperatures and  the  variable  of the  length  contrast  be-  p r e c i p i t a t i o n , the  t h i r d v a r i a b l e , of l e s s e r importance, r e p r e s e n t e d merely a measure of l a t i t u d e .  When the weather s t a t i o n s were grouped,  the  p a r a l l e l s t o Chapman's ( 1 9 5 2 ) c l i m a t i c r e g i o n s were s u r p r i s i n g . Newnham's approach, w i t h f u r t h e r r e f i n e m e n t s , may u s e f u l f o r z o n i n g the n a t u r a l range of t r e e s and f i n d i n g good matches f o r t r e e s p e c i e s Newnham b e l i e v e s  that  prove very assist in  Introduction  elsewhere.  " ... the components can be used  as  measures of c l i m a t i c s i m i l a r i t y between d i f f e r e n t provenances within a  species."  Seed movement Isaac (19^9) based h i s r u l e s f o r D o u g l a s - f i r  seed  c o l l e c t i o n s on L a n g l e t ' s (19^5) p u b l i c a t i o n d e a l i n g w i t h seed movement l i m i t a t i o n s i n Sweden. as  These r u l e s , s t i l l v a l i d , were  followsi 1.  C o l l e c t seed w i t h i n 100 m i l e s n o r t h or south of planting s i t e i f at similar elevations.  2.  A 500 f o o t r i s e or drop i n e l e v a t i o n from p l a n t i n g s i t e i s a l l o w a b l e i f seed source i s not more than t e n m i l e s t o the n o r t h or south.  3.  For each a d d i t i o n a l t e n m i l e s n o r t h of p l a n t i n g s i t e , the a l l o w a b l e e l e v a t i o n f o r seed c o l l e c t i o n i s reduced by f i f t y f e e t , up t o 100 m i l e s north.  4.  F o r each a d d i t i o n a l t e n m i l e s south, the e l e v a t i o n may be Increased by f i f t y f e e t , up t o 100 miles south.  5.  In a rough, broken c o u n t r y , c l i m a t e should guide more than d i s t a n c e or e l e v a t i o n . Average annual temperature of the seed source should be w i t h i n two degrees F a h r e n h e i t p l u s or minus and f r o s t f r e e p e r i o d should be s i m i l a r t o t h a t of the planting s i t e .  6.  The seed source stand should be t h r i f t y and making average or b e t t e r than average growth f o r the l o c a l i t y .  7. I n d i v i d u a l seed t r e e s should be of good form should not be e x c e s s i v e l y limby.  and  Isaac (19^9) s t a t e d e x p l i c i t l y t h a t these r u l e s are not t o be c o n s i d e r e d optimum c o n d i t i o n s but r a t h e r l i m i t a t i o n s for  seed c o l l e c t i o n s .  He recommended t o c o l l e c t seed d u r i n g  heavy c r o p years and s t o r e i t f o r poor y e a r s . l a c k of seed c e r t i f i c a t i o n laws.  Isaac noted  the  Despite the concern about  the provenance problem, no r e f i n e d seed t r a n s f e r r u l e s have been d e v i s e d y e t f o r D o u g l a s - f i r , and l o c a l seed Is s t i l l r e commended as s a f e s t and probably b e s t (Bingham 1 9 6 6 ) . (1966) f e l t  t h a t the Increase l n a r t i f i c i a l  D o u g l a s - f i r l n Washington and  Wheat  reforestation with  Oregon and the l a c k of seed  from  adequate sources made i t easy t o ignore r u l e s of s a f e p r a c t i c e . He a l s o d e p l o r e d the l a c k of c o n t r o l s on the movement of seed. Haddock ( 1 9 6 6 ) noted t h a t n e i t h e r d i s t a n c e of seed movement i n m i l e s nor the change i n e l e v a t i o n i t s e l f  are  really  the q u e s t i o n of importance, but r a t h e r the e f f e c t s of these changes  on the l o c a l t o t a l environment.  Haddock s t r e s s e d the  importance of a thorough knowledge o f the geography  of a r e g i o n  and the i n f l u e n c e o f topography on l o c a l c l i m a t e i n r e s p e c t t o seed movement problems.  Schmidt's ( 1 9 6 7 ) provenance t e s t  h o p e f u l l y y i e l d enough i n f o r m a t i o n f o r new  will  seed t r a n s f e r r u l e s  f o r c o a s t a l D o u g l a s - f i r i n B r i t i s h Columbia l e s s than f i f t e e n y e a r s from  now. P h y s i c a l p r o p e r t i e s of wood have been shown t o be an  Important a s p e c t of the provenance problem, but one which  needs  f u r t h e r i n v e s t i g a t i o n (Haigh, 1 9 6 l > S i l e n , 1964$ B r a m h a l l ,  19661  McKlmmy, 1 9 6 6 ) .  Early Tests The l o n g r o t a t i o n s r e p r e s e n t one of the major h a n d i caps of f o r e s t r y .  Whereas most a g r i c u l t u r a l crops r i p e n a f t e r  one growing season, i t u s u a l l y takes a f o r e s t stand l o n g e r than an average human l i f e t o grow t o m a t u r i t y .  In an e f f o r t t o  overcome t h i s shortcoming In provenance and progeny  testing,  some f o r e s t e r s have been t r y i n g t o d e v e l o p e a r l y t e s t s . work i n t h i s f i e l d was done by Schmidt (Sweden) around 1930  Pioneer  (Germany) and L a n g l e t  (Nanson 19^5)••  E a r l y t e s t s i n v e s t i g a t e provenance v a r i a t i o n i n order t o g i v e the e a r l i e s t i n d i c a t i o n of the l a t e r performance of the trees (Lines 1 9 6 7 ) .  If statistically significant  can r e p e a t e d l y be found between e a r l y and l a t e r  correlations  performance  ( f i v e , t e n or even f i f t e e n y e a r s a r e not good enough f o r  management) i n s e v e r a l t e s t s of the same s p e c i e s or v a r i e t y , c o v e r i n g a l a r g e range o f environmental c o n d i t i o n s , i t should be p o s s i b l e t o s e t up equations p r e d i c t i n g l a t e r performance the b a s i s o f measurements taken a t an e a r l y age. the r e l a t i v e i n f e r i o r i t y  In t h i s way  or s u p e r i o r i t y of a p a r t i c u l a r  ance ( o r parent t r e e i n the case of progeny  on  proven-  t e s t i n g ) c o u l d be  r e c o g n i z e d e a r l y and f o r e s t e r s would know which stand t o choose f o r seed c o l l e c t i o n f o r l a r g e s c a l e p l a n t i n g programs. E a r l y t e s t i n g begins w i t h the seed, a l t h o u g h c l i m a t i c and geographic d a t a on seed source c o n s t i t u t e v e r y important background  information.  Growth chambers f a c i l i t a t e the study  of the d i f f e r e n t i a l r e a c t i o n of provenances environment.  under a c o n t r o l l e d  Irgens-Mjftler*s (1957) i n v e s t i g a t i o n s on the  e f f e c t s of v a r i o u s temperatures and photoperiods on D o u g l a s - f i r s e e d l i n g s from d i f f e r e n t sources a r e a good example o f t h i s . A l t h o u g h l a b o r a t o r y t e s t s a r e very h e l p f u l i n provenance  trials,  c a u t i o n must be used i n p r o j e c t i n g the r e s u l t s t o > l a t e r growth i n the f i e l d .  Nursery performance  ( e . g . Ching and Bever,  i960)  may y i e l d some u s e f u l I n f o r m a t i o n , b u t a g a i n the t r u e v a l u e of those o b s e r v a t i o n s w i l l be confirmed only many y e a r s l a t e r . Orr-Ewing  (1967) s t r e s s e d t h i s need f o r c a u t i o n when b a s i n g  premature c o n c l u s i o n s on e a r l y r e s u l t s .  He stated»  "Progeny  t e s t s can y i e l d enough i n f o r m a t i o n a t twelve t o f i f t e e n y e a r s a f t e r p l a n t i n g f o r a t l e a s t the poorer provenances  t o be r e c o g -  nized."  The same t h i n g can probably be s a i d of provenance  trials.  McKlmmy  (1966), s t u d y i n g s p e c i f i c g r a v i t y of Douglas-  f i r from s e v e r a l seed s o u r c e s , concluded t h a t t r e e s should be  over t w e n t y - f i v e  years  o l d f o r stand p r e d i c t i o n s .  another p u b l i c a t i o n on the 1912  According  Douglas-fir heredity  (United S t a t e s F o r e s t S e r v i c e 1 9 6 4 ) ,  to  study  the r e s u l t s were d i s a p -  p o i n t i n g when s e e d l i n g h e i g h t s were compared t o mature h e i g h t s . In a provenance study w i t h European l a r c h , Leibundgut found t h a t height growth d u r i n g the f i r s t years was r e l a t e d w i t h l a t e r height  (1962)  poorly  cor-  growth.  Some experiments, however, showed high c o r r e l a t i o n s i Wright and Baldwin (1957) found a s t a t i s t i c a l l y correlation  ( r = 0 . 9 3 3 and  f o u r year's  h e i g h t and  provenance t e s t i n New  0.861  significant  r e s p e c t i v e l y ) between t h r e e  seventeen year's  height  i n a Scots  Nanson ( 1 9 6 5 ) *  Hampshire.  or  pine  r e p o r t i n g on  an i n t e r n a t i o n a l Norway spruce experiment i n Belgium d a t i n g back t o 1 9 3 8 ,  found the t w e n t y - f i v e  y e a r growth t o be  strongly  c o r r e l a t e d w i t h the f o l l o w i n g " e a r l y " c h a r a c t e r i s t i c s ! - l e n g t h of growing season a t seed source -  1000  seed weight  - f r e s h weight o f - dry weight of  seedlings  seedlings  - bud b u r s t i n g date - h e i g h t s a t v a r i o u s ages. Nanson f e l t c o u l d be  c o n s i d e r a b l y reduced i f the p l a n t i n g check c o u l d  eliminated. ficient  t h a t the l e n g t h of the t e s t i n g c y c l e  Haddock e t a l . ( 1 9 6 7 ) found a c o r r e l a t i o n c o e f -  of over 0 . 9 3 8 a t the 0 . 0 1  h e i g h t a t age e l e v e n years  be  two  and  p r o b a b i l i t y l e v e l between  h e i g h t a t f i v e , s i x , seven, e i g h t  i n a D o u g l a s - f i r study a t the U n i v e r s i t y of  and  B r i t i s h Columbia Research F o r e s t .  For the same s p e c i e s , Lacaze  ( 1 9 6 8 ) c o u l d show a h i g h l y s i g n i f i c a n t c o r r e l a t i o n between h e i g h t growth a t two, f i v e , e i g h t and t h i r t e e n y e a r s a f t e r p l a n t i n g ( r between two and t h i r t e e n y e a r s was  O.96).  E a r l y t e s t s can a l s o be used as a check on the authe n t i c i t y of seed l o t s c o l l e c t e d by commercial seed d e a l e r s . Sweet ( 1 9 6 5 ) found t h a t b o t h two-year h e i g h t growth and time l a g between l a t e r a l and t e r m i n a l bud b u r s t of D o u g l a s - f i r provenances t e s t e d i n New  Zealand were s u f f i c i e n t l y s t r o n g l y c o r -  r e l a t e d t o c l i m a t e a t seed source t o do a check on the d a t a of o r i g i n s u p p l i e d w i t h commercial seed l o t s .  Schmidt-Vogt  (1967)  showed the same t h i n g f o r Norway spruce and Scots p i n e .  III.  MAJOR PROVENANCE TESTS WITH DOUGLAS-FIR  T e s t s Made Outside I t s N a t u r a l Range D o u g l a s - f i r was d i s c o v e r e d by A r c h i b a l d Menzies i n 1797 a t Nootka Sound on Vancouver I s l a n d  (Krajina 1 9 5 6 ) .  David  Douglas sent cones and o t h e r m a t e r i a l , c o l l e c t e d on the banks of the Columbia R i v e r , t o Europe as e a r l y as 1 8 2 7 ,  The  first  provenance experiments w i t h D o u g l a s - f i r were s t a r t e d much l a t e r . In 1909  and 1910, the U n i t e d S t a t e s F o r e s t S e r v i c e sent Douglas-  f i r seed, c o l l e c t e d under the a e g i s of Zon a t d i f f e r e n t  points  of i t s range from the P a c i f i c Coast t o the Rocky Mountains, t o P r o f e s s o r Schwappach a t Eberswalde, Germany, and t o Count von Berg i n L i v o n i a , R u s s i a .  The Count noted t h a t t r e e s from Chelan  " ... i n the same range of the Cascades as the seed from  Snoqualmle, but on the e a s t e r n s l o p e , have grown v e r y n e a r l y as h i g h as the s e e d l i n g s from Snoqualmle, but stand u p r i g h t and have s c a r c e l y s u f f e r e d a t a l l from the f r o s t . " 1912,  (Zon 1913).  P r o f e s s o r Muench s t a r t e d another provenance  In  test i n  K a i s e r s l a u t e r n , Germany, w i t h seed from t e n provenances, sent by P r o f e s s o r Schwappach who  was  experimenting w i t h n i n e t e e n  provenances h i m s e l f , two from low e l e v a t i o n s i n Western Washi n g t o n , t h r e e from h i g h e l e v a t i o n s i n C a l i f o r n i a , west of the S i e r r a Nevada summit, the remainder from more c o n t i n e n t a l mates. Germanyi  In 1933.  t h r e e other provenance  cli-  t e s t s were s t a r t e d i n  s e r i e s one a t the f o o t of Mount F e l d b e r g i n the B l a c k  F o r e s t , s e r i e s two i n Freienwalde and Braunlage, s e r i e s t h r e e i n Gahrenberg.  These experiments have been r e p o r t e d by  Schober  (195*0, Schober and Meyer (195^, 1 9 5 5 ) . Jahn (1955) t Rohmeder ( 1 9 5 6 ) and o t h e r s .  In Germany I t soon appeared t h a t t h e r e were  g r e a t d i f f e r e n c e s between provenances as t o growth r a t e , and d i s e a s e r e s i s t a n c e , as w e l l as morphology. the  1933  experiments was  frost  The seed f o r  from i l l - d e f i n e d provenances,  there-  f o r e morphology  was  (Decker 1 9 6 7 K  A l t h o u g h t h i s d i d not prove v e r y easy due t o  the  sometimes r e s o r t e d t o f o r c l u e s on o r i g i n  great morphological v a r i a b i l i t y  of D o u g l a s - f i r , the proven-  ances were a s c r i b e d t o three v a r i e t i e s 1 vlrldls  green or c o a s t a l form, c a l l e d m e n z l e s l l i n North America  varietas  glauca  b l u e or Colorado or Rocky Mountain D o u g l a s - f i r by L i t t l e ( 1 9 5 3 )  oaesla  grey form from the i n t e r m o u n t a i n or more n o r t h e r n i n t e r i o r provenances from Schenck's ( 1 9 3 9 ) c l i m a t i c r e g i o n 110 B.  At p r e s e n t most North American a u t h o r i t i e s D o u g l a s - f i r do not u n i v e r s a l l y agree beyond the glauca  separation  ers s t i l l  f r e q u e n t l y w r i t e about the c a e s l a v a r i e t y .  speaking f o r e s t e r s and  s o l v e d by  simply  against  fir.  Although there are  c a l l i n g Pseudotsuga m e n z i e s i i  great v a r i a b i l i t y  L a r s e n ( 1 9 5 6 ) » who  valid  the hyphen, t h i s problem c o u l d e a s i l y  "Douglas" as some Europeans The  r a g i n g among E n g l i s h  b o t a n i s t s regards the hyphenation prob-  i n s p e l l i n g Douglas (-)  arguments f o r and be  menzlesll-  1 9 6 5 ) t a l t h o u g h many German f o r e s t -  (Fowells  Another minor c o n t r o v e r s y  lem  on  (Mirb.) Franco  do.  of D o u g l a s - f i r was  stressed  by  wrote as f o l l o w s i  " ... one has t o t r a v e l very w i d e l y throughout the n a t u r a l range of D o u g l a s - f i r i n order t o get an impression of d i f f e r e n c e s i n geographi c a l type, but s t a n d i n g i n one p l a c e one can, without moving a f o o t , see many i n d i v i d u a l s d i f f e r i n g widely i n t h e i r s t r u c t u r e ; i t i s o f t e n more d i f f i c u l t t o p i c k out those t h a t r e semble another ... I t does not matter i f one chooses i n C a l i f o r n i a a s i t e i n the Coast Range or l n the S i e r r a Nevada, passes through Oregon and Washington, or i n B r i t i s h Columbia s e l e c t s a p l a c e on Vancouver I s l a n d or i n the Rocky Mountains; everywhere one i s bound t o be imp r e s s e d by the g r e a t i n d i v i d u a l v a r i a t i o n of t h i s t r e e s p e c i e s ... " Orr-Ewing ( 1 9 6 8 ) concluded from h i s i n b r e e d i n g that Douglas-fir opportunities  studies  i s a most heterozygous s p e c i e s , l e a v i n g ample  for further selection.  In a comprehensive paper presented i n Stockholm, Schober ( 1 9 6 3 ) summed up Douglas-fir.  European provenance s t u d i e s w i t h  He concluded t h a t i n C e n t r a l Western Europe, prov-  enances from areas west of the  Cascades i n Washington, e s p e c i a l l y  from e l e v a t i o n s below 700 meters  (2000 f e e t ) were f a s t growing  and f a i r l y r e s i s t a n t t o Rhabdocllne pseudotsugae Syd. b u t a t the same time r a t h e r s e n s i t i v e t o w i n t e r f r o s t .  However, l a t e  f r o s t s d i d n o t a f f e c t them because of t h e i r l a t e bud b u r s t . In the c o o l e r c l i m a t e s of N o r t h e a s t e r n Europe, the C e n t r a l European Mountains and the t r a n s i t i o n zones t o c o n t i n e n t a l E a s t e r n Europe, c e r t a i n provenances from the i n t e r i o r of B r i t i s h Columbia grew as f a s t o r f a s t e r than t r e e s from c o a s t a l Washington.  However, they showed g r e a t e r s u s c e p t i b i l i t y t o b o t h  Rhabdocllne and l a t e f r o s t s , due t o t h e i r e a r l y f l u s h i n g , a l though they were r e s i s t a n t t o w i n t e r f r o s t s .  In c o a s t a l r e g i o n s  or i n maritime European c l i m a t e s , the provenances from B r i t i s h Columbia's I n t e r i o r showed only average t o poor performance. Schober expanded on the g r e a t v a r i a b i l i t y  of these sources and  t r i e d t o e x p l a i n i t w i t h i n f o r m a t i o n from Galoux's ( 1 9 5 2 ) publ i c a t i o n on the phytogeography and paleobotany of D o u g l a s - f i r . U n f o r t u n a t e l y , many European f o r e s t e r s l a c k Galoux's knowledge on the s u b j e c t and do not r e c o g n i z e the e x i s t e n c e of B r i t i s h Columbia's I n t e r i o r Wet B e l t and the extreme c l i m a t i c In B r i t i s h Columbia.  variation  T h i s i s p a r t i c u l a r l y s i g n i f i c a n t f o r prov-  enances of D o u g l a s - f i r around Shuswap Lake, f o r i n s t a n c e . Haddock and S z i k l a i ' s  ( 1 9 6 6 ) seed zone map can h e l p seed c o l -  l e c t o r s t o a v o i d gross e r r o r s In the f u t u r e .  H o p e f u l l y these  seed zones w i l l be i n t e r p r e t e d e c o l o g i c a l l y and not c o n s i d e r e d t o be u n i f o r m e n t i t i e s .  Schober ( 1 9 6 3 ) recommended the I n t e r i o r  Wet B e l t o r i g i n s f o r a p p r o p r i a t e p a r t s of Europe.  Those from  B r i t i s h Columbia's d r y I n t e r i o r , c o r r e s p o n d i n g t o Haddock and  Sziklai*s  (1966) seed zones f i v e and e i g h t or t o Rowe's (1959)  Montane F o r e s t Region, were found t o be slow growing and susc e p t i b l e t o Rhabdocllne i n Europe.  Seed from c o a s t a l  Oregon  and C a l i f o r n i a showed good r e s u l t s l n the warmer c l i m a t e s of I t a l y , b u t they proved u n s a t i s f a c t o r y i n C e n t r a l and Northern Europe where they grew s l o w l y and s u f f e r e d from w i n t e r f r o s t s . Provenances from e a s t o f the Cascades, p a r t i c u l a r l y those from the Rocky Mountains, always showed g r e a t s u s c e p t i b i l i t y t o Rhabdocllne and e x h i b i t e d poor growth  (Schober 1963)•  A c c o r d i n g t o Schober (1959)t the c a e s l a and g l a u c a provenances a r e s u s c e p t i b l e t o Rhabdocllne because they a r e not used t o the h i g h a i r humidity of the European c l i m a t e s which does not b o t h e r the c o a s t a l form. the 1963  In the t a b l e s a t t a c h e d t o  p u b l i c a t i o n , Schober c l e a r l y showed t h a t i n most Euro-  pean c o u n t r i e s , s u i t a b l e D o u g l a s - f i r provenances outproduce a l l indigenous s p e c i e s . Rohmeder 1956)  195^,  Many European f o r e s t e r s (e.g. Schober  f e l t t h a t i t was  time t o sample  i n more d e t a i l  the p o p u l a t i o n s from the g e n e r a l areas where the provenances t h a t have proven s u i t a b l e f o r Europe, o r i g i n a t e d . a c t l y what some Danish f o r e s t e r s  (Barner 1 9 6 6 )  T h i s i s ex-  had i n mind when  they sent a team o f cone p i c k e r s t o western N o r t h America f o r several consecutive years, s t a r t i n g i n 1966.  The seed i s d i s -  t r i b u t e d by I.U.F.R.O. t o i n t e r e s t e d r e s e a r c h groups.  Many  more experiments w i t h D o u g l a s - f i r a r e going on i n Europe.  Stern  ( 1 9 6 6 ) f o r i n s t a n c e r e p o r t e d on three t o f o u r y e a r s ' performance of so f a r "unexplored" o r i g i n s from h i g h e l e v a t i o n stands i n A r i z o n a , New  Mexico and Mexico, grown a t Schmalenbeck.  He  pointed  out t h e i r advantages of f a s t e r growth, l o n g e r  p e r i o d , g r e a t e r f r o s t and The  majority  bourg (Decker 1967) D o u g l a s - f i r i s one  and  drought r e s i s t a n c e .  of European c o u n t r i e s , i n c l u d i n g LuxemRussia  "where the c o a s t a l form of  of the most r a p i d l y growing, v a l u a b l e  promising f o r e s t r y species 1 9 6 7 ) t are  growing  i n the western U k r a i n e "  and  (Brodovich  i n t e r e s t e d i n growing s u i t a b l e provenances of t h i s  s p e c i e s , which some a u t h o r i t i e s h a i l as Europe's most important exotic. D o u g l a s - f i r i s a l s o grown In other p a r t s of the w o r l d , Zealand (Sweet 1965)  f o r instance  i n New  tremely w e l l  (Spurr 1 9 6 3 ) «  i n areas o u t s i d e  i t s n a t u r a l range, D o u g l a s - f i r has mainly been Byrnes e t a l . ( 1 9 5 8 )  s e v e r a l provenances In P e n n s y l v a n i a and best  ex-  In e a s t e r n North America, t h a t i s ,  grown f o r Christmas t r e e purposes.  v a r i e t y was  where i t i s d o i n g  found t h a t the  i n terms of s u r v i v a l , growth and  tested  glauca  hardiness.  V i r i d i s , though s l i g h t l y f a s t e r In growth, s u f f e r e d most from c o l d and glauca  drought.  Likewise,  Baldwin and  Rock ( 1 9 6 1 )  rated  s u p e r i o r t o v l r l d l s a f t e r nine growing seasons i n  New  Hampshire.  Major Provenance T e s t s With D o u g l a s - f i r Within The  most famous D o u g l a s - f i r provenance t e s t i n North  America i s the c l a s s i c a l 1912 States Forest Service.  h e r e d i t y study of the  United  I t c o n s i s t s of progenies from 120  corded mother t r e e s , r e p r e s e n t i n g ranging  I t s Range  e l e v a t i o n a l l y from 100  re-  t h i r t e e n c o a s t a l provenances  t o 3 . 8 5 0 f e e t and  planted  at  f i v e e l e v a t i o n s between 1,1.00 and 1,400 and Washington.  f e e t i n western  Oregon  I t i s both a provenance study and a progeny  t e s t from which a w e a l t h of u s e f u l i n f o r m a t i o n has been d e r i v e d . Munger and M o r r i s (1936) found f o r i n s t a n c e t h a t the age of the parent t r e e , the q u a l i t y of i t s growing s i t e , i t s growing space and i t s c o n d i t i o n as t o fungus i n f e c t i o n had no e f f e c t upon the height growth o f i t s progeny.  Wright (1962), u s i n g d a t a from  t h a t p u b l i c a t i o n , c r i t i c i z e d the s t a t i s t i c a l d e s i g n of the study f o r l a c k o f r e p l i c a t i o n and r a n d o m i z a t i o n and concluded premat u r e l y t h a t " ... i n a l l f i v e t e s t areas one or more n o n - l o c a l provenances grew f a s t e r than the l o c a l provenance."  The  1955  November f r o s t , k i l l i n g many non-dormant t r e e s , l o c a l and nonl o c a l , was Silen  going t o change t h i s p i c t u r e d r a s t i c a l l y ,  (1966) t o s t r e s s the importance of s u r v i v a l and  combined,  leading growth  i n s t e a d of growth performance a l o n e , when d i s c u s s i n g  provenance performance.  S i l e n hypothesized that inherent  growth  r a t e of a r a c e has developed toward the maximum t h a t can be s u s t a i n e d i n each l o c a l i t y a g a i n s t impacts of long-term weather extremes.  T h i s h y p o t h e s i s does not exclude the p o s s i b i l i t y of  n o n - l o c a l r a c e s o u t p r o d u c i n g l o c a l ones a t s h o r t r o t a t i o n s on protected s i t e s .  Among o t h e r important f i n d i n g s , the f o l l o w i n g  p o i n t s a r e worth remembering (United S t a t e s F o r e s t S e r v i c e 1962). 1. Seed o r i g i n was more c r i t i c a l i n h i g h - e l e v a t i o n (above 2,000 f e e t ) than i n l o w - e l e v a t l o n p l a n t ations. 2. S u p e r i o r performance of a g i v e n provenance a t one p l a n t i n g s i t e was no assurance of s u p e r i o r performance elsewhere.  3» Slenderness or s t o c k i n e s s o f progeny was cons i s t e n t l y r e l a t e d t o i t s female parent l n a l l plantations. M o r t a l i t y r e s u l t i n g from the severe f r e e z e o f November 1955 was a l s o r e l a t e d t o the i n d i v i d u a l p a r e n t . A c c o r d i n g t o the U n i t e d S t a t e s F o r e s t S e r v i c e (1964), no r e s u l t was more s t r i k i n g than the evidence o f a geneenvironment i n t e r a c t i o n .  Low e l e v a t i o n sources performed p o o r l y  a t h i g h e l e v a t i o n s and v i c e - v e r s a .  I t was a l s o found t h a t en-  vironment g e n e r a l l y has a l a r g e r e f f e c t on growth than source of seed.  The p a t t e r n o f bud b u r s t d i d not appear t o be a l t i -  tudinal.  Wide v a l l e y o r i g i n s b u r s t t h e i r buds f i r s t ,  by those on open s l o p e s .  Those from narrow v a l l e y s  l a s t , probably representing a s e l e c t i o n against l a t e  followed  flushed frosts.  Sweet (1965) came t o s i m i l a r c o n c l u s i o n s l n New Zealand. Other a u t h o r s have used d a t a from the 1912 study. For i n s t a n c e Isaac (1949) e s t a b l i s h e d seven seed t r a n s f e r r u l e s or l i m i t a t i o n s r e l a t i v e t o the c o l l e c t i o n o f D o u g l a s - f i r seed. These r u l e s , l i s t e d  on page 22, a r e based p r i m a r i l y on average  annual temperature and f r o s t f r e e p e r i o d and r o u g h l y p a r a l l e l L a n g l e t ' s (1945) system used l n Sweden f o r Scots p i n e .  Jahn  (1955) compared N o r t h American w i t h German e x p e r i e n c e on D o u g l a s - f i r provenance. In 1954, f o r e s t e r s i n Oregon, r e a l i s i n g the l a c k of knowledge on the D o u g l a s - f i r provenance problem, planned a regionwlde study which comprised s i x t e e n provenances from Oregon, Washington and B r i t i s h Columbia.  T h i s s o - c a l l e d "Co-  o p e r a t i v e Oregon D o u g l a s - f i r provenance study" w i l l be d i s c u s s e d in detail later i n this thesis.  Orr-Ewing (1966) r e p o r t e d on  i n t r a s p e c i f i c c r o s s e s w i t h D o u g l a s - f i r from v a r i o u s o r i g i n s i n the U n i t e d S t a t e s and Canada.  Some North American  research  i n s t i t u t e s a r e growing seed from the I.U.F.R.O. c o l l e c t i o n mentioned e a r l i e r .  Schmidt  (1967) r e p o r t e d on the c r i t i c a l  stages o f an i n t e n s i v e provenance study the major aim of which i s t o s e t up seed t r a n s f e r r u l e s f o r the c o a s t a l range o f D o u g l a s - f i r i n B r i t i s h Columbia.  The a r e a was s t r a t i f i e d ob-  s e r v i n g two c r i t e r i a : 1) r e c o g n i t i o n of known or suspected c l i m a t i c differences, 2) p o s s i b i l i t y t h a t g e o g r a p h i c a l l y s e p a r a t e d areas might have produced d i f f e r e n t ecotypes d e s p i t e only minor c l i m a t i c d i f f e r e n c e s . From a t o t a l of n i n e t y provenances t o be t e s t e d , s i x t y - n i n e a r e from the c o a s t ( f i f t y - s e v e n from B r i t i s h  Columbia,  twelve from Washington and Oregon), t h i r t e e n from c o n t i n e n t a l c l i m a t e s o f B r i t i s h C o l u m b i a ^ I n t e r i o r , and e i g h t from CoastI n t e r i o r t r a n s i t i o n zones.  Cones f o r t h a t study were c o l l e c t e d  In 1966, p a r t l y In c o n n e c t i o n w i t h I.U.F.R.O. t o a v o i d d u p l i c a t i o n o f time and e f f o r t .  F i e l d t e s t i n g w i l l be conducted a t  over f o r t y o u t p l a n t i n g a r e a s , d i s t r i b u t e d over a comprehensive range o f c l i m a t e s w i t h i n the c o a s t a l range of D o u g l a s - f i r . M e t e o r o l o g i c a l d a t a w i l l be obtained a t the f i e l d t e s t In 1967 Haddock e t a l . p u b l i s h e d a paper coastal with i n t e r i o r Douglas-fir origins. s t r e s s e d the Importance  sites.  comparing  The a u t h o r s , who  of a thorough knowledge o f the geography  of an a r e a and an u n d e r s t a n d i n g o f the I n f l u e n c e of topography on l o c a l c l i m a t e , confirmed European e x p e r i e n c e t h a t c o a s t a l provenances outgrow  I n t e r i o r sources i n m i l d c l i m a t e s , w h i l e  b e i n g a t the same time l e s s f r o s t r e s i s t a n t and l e s s s u s c e p t i b l e t o Rhabdocllne i n f e c t i o n s .  Once more, l o c a l or n e a r - l o c a l  o r i g i n s were found t o produce the b e s t growth. l i c a t i o n s have appeared  Many more pub-  on D o u g l a s - f i r provenance  i n North  America. M o r p h o l o g i c a l problems i n s t a n c e by A l l e n ( 1 9 6 0 a , 1 9 6 l )  have been i n v e s t i g a t e d f o r who  d e s c r i b e d an easy method of  s e p a r a t i n g c o a s t a l from B r i t i s h Columbia  i n t e r i o r sources.  Tusko ( 1 9 6 3 ) concluded a f t e r an e x t e n s i v e study on the v a r i a b i l i t y i n c e r t a i n Douglas-fir populations i n B r i t i s h  Columbia,  t h a t t h e r e a r e only two s u b s p e c i e s - one c o a s t a l , and one ior.  Sziklai  ology o f 1 , 3 3 5 kZ°0?  inter-  ( 1 9 6 9 ) s t u d i e d v a r i a t i o n i n cone and seed morpht r e e s from the I.U.P.R.O. c o l l e c t i o n between  and 5 3 ° 3 7 l a t i t u d e n o r t h and found " ... a c l e a r l y  ex-  pressed c l i n a l v a r i a t i o n i n cone and seed l e n g t h w i t h an i n c r e a s i n g t r e n d from n o r t h t o south.  The o t h e r c h a r a c t e r i s t i c s  such as wing l e n g t h , wing width and seed width, d i d not show a similar c l i n a l variation pattern." Growth chamber experiments under c o n t r o l l e d were c a r r i e d out by Irgens--Miller ( 1 9 5 7 ) who response t o temperature and p h o t o p e r i o d .  light,  found e c o t y p i c  High e l e v a t i o n p l a n t s  appeared t o have a d e f i n i t e p h o t o p e r i o d i c response w i t h r e g a r d t o date of bud b u r s t i n g .  The magnitude of t h i s response  i n c r e a s e d by low n i g h t temperatures.  was  Irgens-Mjrfller concluded  t h a t the n a t u r a l s e l e c t i o n t o which D o u g l a s - f i r a t h i g h e l e v a t i o n s has been exposed may  have r e s u l t e d i n p l a n t s f o r which  the o p t i m a l l e n g t h of the day f o r bud b u r s t i n g occurs a t a time  when the danger of n i g h t f r o s t i s u s u a l l y low. concluded  t h a t p h o t o p e r i o d i c ecotypes  V a a r t a j a (1959)  have e v o l v e d as an I n d i v -  i d u a l mechanism of t r e e s t o s e a s o n a l l y changing c l i m a t i c ors.  Allen  ( 1 9 6 0 b ) separated c o a s t a l from i n t e r i o r seed  factlots  on the b a s i s of t h e i r germination behaviour a t v a r i o u s tempera t u r e s , f o l l o w i n g a s h o r t p e r i o d of s t r a t i f i c a t i o n .  Revel  ( i 9 6 0 ) compared c o a s t a l w i t h i n t e r i o r provenances grown i n a greenhouse, and found t h a t germination was  f a s t e r and  height  growth ceased much sooner f o r i n t e r i o r provenances which a l s o r e q u i r e more c h i l l i n g t o break dormancy f o r v e g e t a t i v e growth. S i m i l a r l y , N i c h o l s o n ( 1 9 6 3 ) found h i g h e r germination f o r i n t e r i o r provenances.  He separated i n t e r i o r from c o a s t a l provenances  by u s i n g s h o r t day  treatments.  However no d e f i n i t e r e g i o n a l  grouping of the c o a s t a l provenances was Sorensen  ( 1 9 6 7 ) was  distinguishable.  a b l e t o separate two y e a r - o l d s e e d l i n g s  r e p r e s e n t i n g s e v e r a l provenances from a west-east Oregon i n t o three d i s t i n c t groups, growth and the date of bud  t r a n s e c t In  on the b a s i s of t h e i r h e i g h t  f o r m a t i o n In the f i r s t  year.  Haddock and Schmidt (1957) wrote t h a t i t i s b e s t t o assume t h a t there are a l a r g e number of unexplored s i n c e the e c o l o g i c a l behaviour greatly with s i t e , of i t s range."  ecotypes  of D o u g l a s - f i r o b v i o u s l y v a r i e s  " ... even w i t h i n the P a c i f i c Coast p o r t i o n  Bingham ( 1 9 6 6 ) f e l t  t h a t many cases of a b e r r a n t  performance In " l o c a l " sources i n v o l v e movement of seed between unrecognized, but n e v e r t h e l e s s d i s t i n c t l y d i f f e r e n t e n v i r o n ments.  He recommended adherence t o the maxims  s a f e s t and probably b e s t , and  l o c a l seed i s  he c a u t i o n e d a g a i n s t s t r e t c h i n g  the concept  of " l o c a l , " e s p e c i a l l y where steep c l i m a t i c  gradi-  ents or s o l i changes are known or suspected between seed and p l a n t i n g a r e a .  Haddock and S z l k l a l  source  ( 1 9 6 6 ) e s t a b l i s h e d seed  c o l l e c t i o n zones f o r D o u g l a s - f i r i n Canada, based  primarily  c l i m a t i c d a t a and the d i s t r i b u t i o n of a s s o c i a t e d s p e c i e s . authors recommended t h a t seed u s e r s who cific locality  cannot  on The  indicate a  spe-  or stand from which they want seed, should pro-  v i d e p e r t i n e n t c l i m a t i c d a t a f o r the proposed i n order t o a v o i d u n d e s i r a b l e r e s u l t s .  plantation  site  Haddock ( 1 9 6 7 ) s t r e s s e d  the need f o r more p r e c i s e provenance d e s i g n a t i o n s because of the great topographic and a s s o c i a t e d c l i m a t i c and edaphic  variation  b e l i e v e d t o have shaped the e v o l u t i o n and development of l o c a l l y , g e n e t i c a l l y d i f f e r e n t p o p u l a t i o n s over l o n g p e r i o d s of time.  The author a l s o warned a g a i n s t r e l y i n g too much on morph-  o l o g i c a l f e a t u r e s used by taxonomlsts  " ... we  should r e l y more  on l e s s e a s i l y measured c h a r a c t e r i s t i c s such as g e n e r a l p h y s i o l ogy, phenology, c o l d , heat and drought  resistance,  susceptibil-  i t y t o i n s e c t s and d i s e a s e , growth form and wood q u a l i t y . "  IV.  THE  CO-OPERATIVE DOUGLAS-FIR PROVENANCE TEST*  General The g r e a t Importance of D o u g l a s - f i r In the r e g i o n a l economy and the l a c k of knowledge on the v a r i a t i o n w i t h i n t h i s  In t h i s t h e s i s " c o - o p e r a t i v e t e s t " or " c o - o p e r a t i v e study" are used as a b b r e v i a t i o n s f o r "the c o - o p e r a t i v e D o u g l a s - f i r provenance t e s t . "  s p e c i e s l e a d i n 195^ F o r e s t and  t o a p u b l i c a t i o n by the P a c i f i c Northwest  Range Experiment S t a t i o n , e n t i t l e d  "A Program of  Tree Improvement f o r the P a c i f i c Northwest," s t a t i n g t h a t  the  tolerance  of Northwest s p e c i e s t o changes i n geography, e l e v -  a t i o n and  c l i m a t e , had not been f u l l y explored  and  hensive program of provenance t e s t i n g should be soon as p o s s i b l e .  a compre-  initiated  as  That same y e a r , s t a f f of the Research Div-  i s i o n of the Oregon S t a t e Board of F o r e s t r y  (now  i n the  Research L a b o r a t o r y , Oregon State U n i v e r s i t y ) began t o  Forest organize  a regionwide provenance study of c o a s t a l D o u g l a s - f i r , a f t e r c o n f e r r i n g w i t h many i n t e r e s t e d p a r t i e s (Anon. 1 9 5 5 & K  Hypotheses t o be  Tested  A f t e r reviewing  l i t e r a t u r e on the D o u g l a s - f i r prov-  enance problem, Ghing and  Bever (19->0) f e l t t h a t i n the  experi-  ments reviewed, the sampling of the v a r i a b l e s of a l t i t u d e , l a t i t u d e and  l o n g i t u d e was  n e i t h e r p r e c i s e nor  systematic  enough t o e l u c i d a t e the q u e s t i o n whether v a r i a t i o n i s c l i n a l or d i s c o n t i n u o u s .  To improve on t h i s , an e f f o r t was  made t o  get as many c o - o p e r a t o r s as p o s s i b l e i n order t o t e s t the f o l l o w i n g hypotheses (Anon. 1 9 5 5 a ) • 1.  D i s t i n c t r a c e s of D o u g l a s - f i r are a s s o c i a t e d w i t h temperature as measured by f r o s t f r e e days d u r i n g the growing season.  2. D i s t i n c t r a c e s altitude.  of D o u g l a s - f i r are a s s o c i a t e d  with  3.  of D o u g l a s - f i r are a s s o c i a t e d  with  D i s t i n c t races latitude.  4. D i s t i n c t r a c e s of D o u g l a s - f i r are a s s o c i a t e d w i t h temperature, a l t i t u d e and l a t i t u d e , as measured  by comparable f r o s t f r e e days and comparable photoperlods d u r i n g the growing season. 5.  D i s t i n c t r a c e s of D o u g l a s - f i r have r e s u l t e d from the development of g e n e t i c s t r a i n s i n l o c a l i z e d a r e a s , and n o t from any o f the v a r i a b l e s or comb i n a t i o n s of v a r i a b l e s s t a t e d i n the p r e c e d i n g f o u r hypotheses.  6. D i s t i n c t r a c e s of D o u g l a s - f i r do n o t e x i s t w i t h i n the a r e a covered by t h i s study.  Design The d e s i g n was t o f o l l o w the working p l a n f o r the southern pine provenance study by Wakeley ( 1 9 5 3 ) who had found a t l e a s t one o f the f o l l o w i n g s i x weaknesses i n o t h e r  proven-  ance s t u d i e s 1 1.  O c c a s i o n a l a i m l e s s and i l l o g i c a l s e l e c t i o n of seed s o u r c e s . T h i s mistake was t o be avoided through r e l a t i v e l y s y s t e m a t i c sampling w i t h i n the range of c o a s t a l D o u g l a s - f i r i n Oregon, Washington and B r i t i s h Columbia.  2. Mixing of s i n g l e t r e e and b u l k cone c o l l e c t i o n s . A r i g i d cone c o l l e c t i o n p r o s p e c t u s , a s k i n g f o r a sample o f a t l e a s t f i f t y t r e e s p e r seed source was aimed a t overcoming t h i s shortcoming. 3.  Inadequate t e s t p l a n t i n g s . T h i s e r r o r was t o be c o r r e c t e d by e s t a b l i s h i n g p l a n t a t i o n s of a l l sources near each c o l l e c t i o n s i t e t o compare the l o c a l provenance w i t h the other provenances.  4. Inadequate s t a t i s t i c a l d e s i g n . Four r e p l i c a t i o n s w i t h random provenance a l l o c a t i o n was t o s a t i s f y the p r e r e q u i s i t e s f o r s t a t i s t i c a l a n a l y s i s (see Fig. 1). 5.  I n s u f f i c i e n t number o f t r e e s p l a n t e d f o r adequate s u r v i v a l and measurement up t o r o t a t i o n age. An e i g h t by e i g h t f o o t s p a c i n g i n square p l o t s w i t h  Since Wakeley's p u b l i c a t i o n was n o t seen, the Information on " d e s i g n " i s mainly taken from Heaman's (19^3) t h e s i s .  e l e v e n times e l e v e n s e e d l i n g s was t o be adopted; the two o u t e r rows were t o a c t as a b u f f e r s t r i p , m i n i m i z i n g edge e f f e c t s (see F i g . 2 ) , 6.  Lack of maintenance a f t e r e s t a b l i s h m e n t of the t e s t , and subsequent I r r e g u l a r measurements. Good c o - o p e r a t i o n and the r a t i f i c a t i o n of a ten-year agreement by a l l c o - o p e r a t o r s was t o remedy t h i s . In  a d d i t i o n t o the above p o i n t s , confounding  nursery  e f f e c t s were t o be e l i m i n a t e d by growing a l l s t o c k i n one n u r s e r y , namely C o r v a l l l s .  Fig.  1  Diagram of p l o t d e s i g n a t i o n s .  B l o c k A=l  B l o c k B=2  B l o c k C=3  Block  13  8  14  7  4  10  1  13  14  6  1  10  2  6  1  4  12  9  15  5  2  8  2  16  4  16  15  5  11  16  7  8  14  5  13  9  12  9  3  10  2  12  3  3  15  7  ll 6  16  Plantation I  11  11  D=4 8  14  6  4  12  5  3  7  10  1  9  13  15  Plantation II  Each o u t p l a n t i n g a r e a has two  " p l a n t a t i o n s , " u s u a l l y from  one-  q u a r t e r m i l e t o one-half m i l e a p a r t , and c o n s i s t i n g each of two a d j a c e n t b l o c k s .  Co-operators and Seed  Sources  T a b l e s 1 and 2 show the names of the c o - o p e r a t o r s as w e l l as the d i s t r i b u t i o n of the seed s o u r c e s .  Figure 3  illus-  t r a t e s the geographic d i s t r i b u t i o n of the provenances i n c l u d e d i n the c o - o p e r a t i v e t e s t . letters  The provenances were f i r s t  assigned  (A t o P ) , then g i v e n numbers f o l l o w i n g the order i n  Fig.  2  Blow-up of p l o t X i n p l a n t a t i o n Y.  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X 7 X 8 X 21 X 22 X 35 X 36 X 49 X  X  X  X 6 X 9 X 20 X 23 X 34 X 37 X 48 X  X  X  X 5 X 10 X 19 X 24 X 33 X 38 X 47 X  X  X  X 4 X 11 X 18 X 25 X 32 X 39 X 46 X  X  X  X 3 X 12 X 17 X 26 X 31 X 40 X 45 X  X  X  X 2 X 13 X 16 X 2? X 30 X 4l X 44 X  X  X  X 1 X 14 X 15 X 28 X 29 X 42 X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  X  »3  Features of each s m a l l -  121  - 8 x 8 '  ploti  trees spacing  - a l l t r e e s of the same provenance - only 4 9 t r e e s i n the i n n e r groups w i l l be measured. They a r e marked w i t h numbered cedar s t a k e s .  Table  1  C o - o p e r a t o r s and L o c a t i o n of the T e s t  Seed Source British Co lurabia  Washington  Oregon  Provenance Number  Sites Elevation L a t i - ( F e e t ) range tude of c o l l e c t i o n  Co-operator  A  4  Canadian F o r e s t Products L t d .  50°30  B  2  Crown Z e l l e r b a c h (Canada) L t d .  C  1  D  3  E  15  University of B r i t i s h  F  16  Weyerhaeuser Company  47°30'  G  6  Simpson Olympic Tree Farm  47°15'  H  5  Dept. Nat. Resources, S t a t e of Wash.  I  7  J  Year o f Collection  400-600  1956  49°45'  1300-1700  1956  MacMillan Bloedel Limited  49°10'  2600-2900  56/57  B. C. F o r e s t  48°50'  570-750  1955  49°10'  500-700  1957  Service Columbia  !  39-4100  Location of Test Site Nimpkish R i v e r Valley Courtenay  Sugar Loaf Mountain, Chemainus R i v e r Robertson V a l l e y near Lake Cowichan University of B r i t i s h Columbia Research F o r e s t  1957  Snoqualmie  100-500  1956  Shelton  46°45'  1850-2000  1956  Elbe Area  45°30'  1600-2200  54/55  9  S t a t e Board of F o r e s t r y , Oregon Crown Z e l l e r b a c h C o r p o r a t i o n  45°10'  1600-2000  1956 "1  K  10  Crown Z e l l e r b a c h C o r p o r a t i o n  45° 10'  3200-3800  1956  L  8  Jack Stump and Kenneth McCrae  44°50'  200  1956  M  11  Oregon S t a t e C o l l e g e  44°30'  1800-2000  55/56  N  12  U.S. F o r e s t  Service  43°45'  1800-2000  1954  0  13  U.S. F o r e s t  Service  43°45«  2500-3000  1956  P  14  Medford  42°20'  2700-3300  1955  Corporation  ;  Area  Area  T i l l a m o o k Area  J  Clackamas  Tree Farm  Mola11a A r e a Willamette V a l l e y , Salem A r e a McDonald F o r e s t , C o r v a l l i s Area Oakridge Area High P r a i r i e , Oakridge Area Butte F a l l s  Table 2  D i s t r i b u t i o n o f the Seed Sources  a)  British  Columbia  Elevation  Location 0-1000 N. Vancouver I s l a n d  1000-2000  Robertson V a l l e y ( D )  Mainland  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 ( E )  Sugar l o a f Mtn.(C)  U.S.A.  Location  Coast Zone ~~0-1000  Oregon  Courtenay(B)  S., Vancouver Is land  Washington  2000 +  Nimpkish(A)  C. Vancouver Is land  b)  (feet)  Shelton(G)  1000-2000  V a l l e y Zone 2000 +  Cascade Zone 1000-2000  ~2000-3000  Elbe(H) Tillamook(I) Corvallis(M)  3000 + Snoqualmie(F)  Willamette(L) Molalla(J) Butte F a l l s ( P )  Oakridge(N)  Oakridge(O)  Molalla(K)  which the seed l o t s reached the n u r s e r y (Table l ) . provenances  E and F were the l a s t ones t o be sent t o G o r v a l l i s  and were thus a s s i g n e d the numbers 15  and  16.  The f o l l o w i n g persons a c t e d as r e g i o n a l Oregon Washington B r i t i s h Columbia The f i r s t  F o r example,  i « i  J . F . Gartz J.W. D u f f i e l d A.L. Orr-Ewing  o v e r a l l c o - o r d l n a t o r was  State Board  co-ordinatorst  D.N.  Bever of the Oregon  of F o r e s t r y ? P r o f e s s o r K.K.  Chlng of the Oregon  S t a t e U n i v e r s i t y r e p l a c e d him l a t e r  on.  Phases The study was  separated i n t o f o u r d i s t i n c t  S e l e c t i o n of o u t p l a n t i n g areas and seed  phasesi  collection  The s e l e c t i o n of the t e s t s i t e s was  left  t o the  indiv-  i d u a l c o - o p e r a t o r w i t h a p p r o v a l by the r e g i o n a l c o - o r d i n a t o r . The t e s t s i t e s were t o meet the f o l l o w i n g requirements! i ) be i n the D o u g l a s - f i r type i i ) n o r t h t o northwest a s p e c t i l l ) l a r g e enough t o handle two s i x - a c r e p l a n t a t i o n s iv)  be on l a n d expected t o remain i n permanent ownership  v) not t o be r e l o g g e d or have a heavy b r u s h or herbaceous c o v e r . In  areas where e x c e s s i v e deer browsing was  an e i g h t f o o t fence was I t was requirements.  expected,  recommended.  not always p o s s i b l e t o s t r i c t l y  meet a l l the  Most good growing s i t e s s u f f e r from a b r u s h  invasion.  A l s o , mountainous s o i l s are extremely v a r i a b l e  t h e r e f o r e there may  and  be u n d e s i r a b l e s i t e h e t e r o g e n e i t y w i t h i n  as s m a l l an a r e a as s i x a c r e s .  The cone c o l l e c t i o n areas were  d e f i n e d as having a r a d i u s of t w e n t y - f i v e m i l e s i n a d e s i g n a t e d a r e a , a t an e l e v a t i o n not d i f f e r i n g from t h a t of the o u t p l a n t ing  a r e a by more than two hundred f e e t  f o u r hundred f e e t ) . at  (or e x c e p t i o n a l l y up  to  A maximum of f i f t y t r e e s should be p i c k e d  random t o sample as much v a r i a t i o n as p o s s i b l e , and t o pro-  vide 3*2  pounds of seed from each a r e a (Anon. 1 9 5 5 t > ) »  Seed  crop p e r m i t t i n g , the c o l l e c t i o n s were s t a r t e d i n the f a l l 195^*  The l a s t two l o t s of seed  Columbia Research i.e. 1957.  (from the U n i v e r s i t y of B r i t i s h  F o r e s t , i . e . seed source 15.  seed source l 6 )  Seed was  e x t r a c t e d by v a r i o u s o r g a n i s a t i o n s . obtained, the  s t o r e d i n a c o l d storage a t z e r o degree  h e i t a t the Oregon S t a t e F o r e s t Nursery Bever,  Snoqualmle,  from f o u r t e e n t o  A f t e r an e i g h t y - f i v e percent c u t t i n g t e s t was c l e a n seed was  and  a r r i v e d a t the n u r s e r y i n C o r v a l l l s i n f a l l  The number of t r e e s sampled ranged  eighty-nine.  of  In C o r v a l l l s  Fahren-  (Ching and  i960).  Nursery  phase In  seed was  order t o a v o i d confounding n u r s e r y e f f e c t s , a l l  raised i n Corvallls.  Naked s t r a t i f i c a t i o n was  used  by s o a k i n g the i n d i v i d u a l seed l o t s i n p l a s t i c bags w i t h water for  f o r t y - e i g h t hours.  A f t e r the water was  bags were p l a c e d i n a c o o l room a t and  d r a i n e d o f f , the  - 37 degrees  Fahrenheit  95 percent r e l a t i v e humidity, f o r three weeks (Ching and  Bever,  i960).  The  seeding r a t e was  v i a b i l i t y of the seed l o t . s t a r t e d on May  15  Sowing of f o u r t e e n seed l o t s  and completed  on May  d i c a t i o n of f i e l d germination was l i n g s were l i f t e d  a d j u s t e d a c c o r d i n g t o the  17,  1957*  n o t i c e d on May  i n l a t e February 1959  was  The f i r s t i n 28.  ( i . e . as 2 - 0  The  seed-  s t o c k ) and  c u l l e d a c c o r d i n g t o c r i t e r i a advocated by Edwards ( 1 9 5 6 ) .  All  s e e d l i n g s w i t h l e s s than a f o u r Inch top, were d i s c a r d e d .  Due  t o a poor cone c r o p , seed l o t s E (from 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 ) and F (Snoqualmle) were shipped  t o the n u r s e r y one y e a r a f t e r the other provenances, i n the s p r i n g of 1958}  consequently  and sown  t r e e s from these two  origins  were t o remain one y e a r younger than the others throughout  the  d u r a t i o n of the c o - o p e r a t i v e study.  P l a n t i n g of the n u r s e r y s t o c k The  o r i g i n a l p l a n s p e c i f i e d one p l a n t i n g crew i n  each r e g i o n (Oregon, Washington, B r i t i s h Columbia). not prove f e a s i b l e , and p l a n t i n g was c o - o p e r a t o r s i n f a l l 1959 p r e v i o u s l y been adequately  Field  c a r r i e d out by  or s p r i n g i 9 6 0 .  The  This d i d individual  test sites  had  prepared.  examinations Co-operators were p r o v i d e d w i t h thermometers, r a i n  gauges and s t a n d a r d i z e d f i e l d t a l l y s h e e t s . h e i g h t growth was  Current and  t o be assessed i n m e t r i c u n i t s and  kept on f l u s h e s , f r o s t and  other damage.  total  records  Results Ching ( 1 9 5 8 ) r e p o r t e d on f i r s t y e a r performance o f f o u r t e e n provenances  (the U n i v e r s i t y of B r i t i s h Columbia Re-  s e a r c h F o r e s t s e e d l p t and the Snoqualmle  source were sown one  year l a t e r than the o t h e r s ) In the n u r s e r y a t C o r v a l l l s . Height growth, bud b u r s t and bud s e t , as w e l l as f r o s t damage were a s s e s s e d and g e n e t i c v a r i a t i o n s found.  S e e d l i n g s from the  Nlmpkish V a l l e y , Vancouver I s l a n d , and from B u t t e F a l l s , I.e.  Oregon,  from b o t h l a t i t u d i n a l extremes of the study, showed a d e f -  i n i t e t r e n d o f i n f e r i o r growth.  Some s i g n i f i c a n t  differences  In the s u s c e p t i b i l i t y t o f r o s t damage were found, b u t c a u t i o n was expressed t o a v o i d premature  conclusions.  Ching and Bever ( i 9 6 0 ) r e p o r t e d on two y e a r s ' n u r s e r y performance. the  They found no c o r r e l a t i o n between the h e i g h t of  s e e d l i n g s and the a l t i t u d e of t h e i r p l a c e o f o r i g i n , a l -  though they d i d f i n d s i g n i f i c a n t d i f f e r e n c e s i n needle l e n g t h between v a r i o u s provenances and some c o r r e l a t i o n between needle l e n g t h and t o t a l h e i g h t growth.  Except f o r the Nlmpkish  Valley  provenance, the Vancouver I s l a n d sources compared f a v o r a b l y w i t h the  l o c a l W i l l a m e t t e V a l l e y provenance, as f a r as h e i g h t  was concerned.  growth  Under the s h o r t e r p h o t o p e r i o d i n Oregon, the  Vancouver I s l a n d s o u r c e s , e x c e p t i n g the Nlmpkish one, formed t h e i r buds e a r l i e r and terminated t h e i r growth e a r l i e r than others.  The f a c t t h a t the southernmost provenance  (Butte  F a l l s ) d i d not show i n c r e a s e d h e i g h t growth under the s l i g h t l y l o n g e r p h o t o p e r i o d a t C o r v a l l l s , was I n t e r p r e t e d t o be a r e s u l t of n a t u r a l s e l e c t i o n f o r e a r l y c e s s a t i o n of growth In an  environment c h a r a c t e r i z e d by severe summer droughts.  High  e l e v a t i o n o r i g i n s showed a tendency toward l a t e bud b u r s t i n g , as an a d a p t a t i o n  to late spring f r o s t s .  No c o r r e l a t i o n was  found between time o f bud b u r s t and annual height Ching ( i 9 6 0 ) r e p o r t e d  growth.  on s u r v i v a l o f f o u r t e e n  prov-  enances on eleven t e s t s i t e s a f t e r the f i r s t growing season i n the f i e l d .  Height was measured i n f o u r p l a n t a t i o n s . *  Except  f o r the Nlmpkish V a l l e y provenance, the Vancouver I s l a n d sources compared very f a v o r a b l y w i t h a l l o t h e r s , the Sugar-Loaf Mountain o r i g i n  (C) b e i n g among the three b e s t i n a l l f o u r  plantations. Walters and Soos ( 1 9 6 l b ) based t h e i r s t u d i e s  regarding  lammas growth on the U n i v e r s i t y o f B r i t i s h Columbia Research F o r e s t on D o u g l a s - f i r s e e d l i n g s , I n c l u d i n g the c o - o p e r a t i v e test.  T h e i r f i n d i n g s , e.g. t h a t lammas growth I s g r e a t l y  enced by environment have been reviewed above. (196la)  influ-  The same authors  i n v e s t i g a t e d the e f f i c i e n c y of v a r i o u s chemicals t o  prevent hares from damaging the young D o u g l a s - f i r t r e e s .  The  r e p e l l e n t s used d i d not prove adequate; p r o t e c t i o n of the natu r a l predators  of the hares was suggested as a b e t t e r means of  keeping the v a r y i n g hare p o p u l a t i o n Heaman ( 1 9 6 3 ) r e p o r t e d Columbia f o r the years 1 9 5 ^ - 1 9 6 1 .  i n check.  on the p r o j e c t i n B r i t i s h M o r t a l i t y and height  growth  a f t e r the i 9 6 0 and the 1 9 6 l growing seasons were assessed f o r the f i v e o u t p l a n t i n g areas In the p r o v i n c e and weather data  Unless otherwise d e f i n e d , " p l a n t a t i o n " I s synonymous w i t h " t e s t s i t e " or " o u t p l a n t i n g a r e a . "  were d i s c u s s e d .  Heaman's c r i t i c i s m covered the f o l l o w i n g  points t 1.  Scope of the The  study.  scope of the c o - o p e r a t i v e study was  c o v e r i n g e i g h t degrees of l a t i t u d e .  too wide,  According to  Schmidt's (1962a) c l a s s i f i c a t i o n , the c o - o p e r a t i v e t e s t f a l l s Into the category of " s t u d i e s of broad geographic  variation."  be expected ations.  T h i s type of study cannot  to y i e l d more than broad g e n e r a l i z -  Information gained from such a p r o j e c t can  be used t o d e s i g n a more I n t e n s i v e one  i n the f u t u r e ,  but as the r e s u l t s cannot be r e l i a b l y i n t e r p r e t e d f o r a t l e a s t twenty y e a r s , t h i s Is a l e n g t h y approach. P r a c t i c a l a p p l i c a t i o n s , such as seed t r a n s f e r r u l e s , cannot be 2.  expected.  Cone c o l l e c t i o n . The cone c o l l e c t i o n was way  by a l l c o - o p e r a t o r s .  not c a r r i e d out i n a u n i f o r m Apart from t h i s , the twenty-  f i v e m i l e c o l l e c t i o n r a d i u s was  too l a r g e , d e s p i t e  the l i m i t a t i o n of c o l l e c t i n g w i t h i n f o u r hundred, pref e r a b l y w i t h i n two hundred f e e t of the e l e v a t i o n of the t e s t i n g s i t e .  Heaman supported  his c r i t i c i s m  w i t h r e c o r d s from f i v e weather s t a t i o n s l y i n g w i t h i n those l i m i t a t i o n s i n a t w e n t y - f i v e m i l e r a d i u s of the Robertson  Valley.  The average f r o s t f r e e p e r i o d , the  major c l i m a t i c c r i t e r i o n of the c o - o p e r a t i v e  study,  between two extreme s t a t i o n s , v a r i e d from f i f t y - o n e  t o two hundred and f o r t y - f o u r days! 3«  Weather r e c o r d s . Heaman d e p l o r e d the l a c k of c o n t i n u i t y and s t a n d ardization.  Some c o - o p e r a t o r s measured  temper-  a t u r e s a t two f e e t , o t h e r s a t f o u r , o t h e r s s t i l l a t f i v e f e e t above the ground. site in i960,  A t the Nlmpkish  test  a f r o s t f r e e p e r i o d of e l e v e n weeks  was r e c o r d e d a t two f e e t above the ground and  twenty-  two weeks a t f i v e f e e t above the ground. 4.  Site  selection. Heaman f e l t t h a t more care c o u l d have been e x e r c i s e d in site selection.  For i n s t a n c e , i t might have been  r e c o g n i z e d from the s t a r t t h a t the Robertson V a l l e y p l a n t a t i o n was  l o c a t e d i n a f r o s t pocket.  A frost  f r e e p e r i o d of only s i x weeks was measured a t two f e e t above the ground i n i 9 6 0 . Heaman found no c o r r e l a t i o n between h e i g h t growth and e l e v a t i o n or l a t i t u d e of seed s o u r c e .  There was no  correlation  between e i t h e r seed weight and h e i g h t growth or between germina t i o n percentage and seed weight d u r i n g the f i r s t y e a r . At Nlmpkish and Courtenay, the low e l e v a t i o n source G (Shelton) and the h i g h e l e v a t i o n source C (Sugar-Loaf Mounta i n ) showed b e s t performance, a l t h o u g h they d i d not prove  sig-  n i f i c a n t l y s u p e r i o r on a l l B r i t i s h Columbia t e s t s i t e s . *  The  Heaman d i d not t e s t the provenance x b l o c k i n t e r a c t i o n , a l t h o u g h he p o i n t e d t o the s i t e h e t e r o g e n e i t y 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 .  unusual  e a r l y growth of S h e l t o n  (G), I f endorsed by more r e l i -  a b l e d a t a i n the f u t u r e , was i n t e r p r e t e d as a p o s s i b l e conf i r m a t i o n of the h y p o t h e s i s  t h a t " ... d i s t i n c t r a c e s o f  D o u g l a s - f i r have r e s u l t e d from the development o f g e n e t i c s t r a i n s i n l o c a l i z e d areas n i f i c a n t deductions  ... "  Heaman concluded  that  sig-  c o u l d n o t be made because o f the s h o r t  d u r a t i o n of the experiment, the incompleteness  and l a c k o f  s t a n d a r d i z a t i o n o f weather d a t a , and because of the e x c e s s i v e damage a t some t e s t s i t e s by f r o s t The  and browsing.  f o l l o w i n g were Heaman's recommendationst  1. Provenance s t u d i e s should be of a s m a l l e r scope than the c o - o p e r a t i v e t e s t , and sample the range of a s p e c i e s much more i n t e n s i v e l y w i t h i n much narrower l a t i t u d i n a l l i m i t s . 2. S t a n d a r d i z a t i o n and c o n t r o l s i n a l l phases of a c o - o p e r a t i v e study by one c o - o r d i n a t o r i n constant p e r s o n a l c o n t a c t w i t h a l l I n volved, are e s s e n t i a l . 3.  Cone c o l l e c t i o n areas have t o be c l e a r l y d e f i n e d and based on a thorough l o c a l knowledge •  4. Growth s t u d i e s should be based on p h e n o l o g i c a l o b s e r v a t i o n s Instead of t e d i o u s weather measurements. 5. A comprehensive study of a l l o u t p l a n t i n g areas i s necessary r e g a r d i n g c l i m a t e , s o i l and microtopography. 6.  The p l a n t a t i o n s have t o be p r o t e c t e d e f f i c i e n t l y from animal damage, i f e a r l y a s s e s s ments a r e t o have any meaning, u n l e s s i t can be e s t a b l i s h e d t h a t the damage i s evenly d i s t r i b u t e d over a l l s o u r c e s . Ching  (1965) assessed  s u r v i v a l and growth a f t e r the  The Robertson V a l l e y p l a n t a t i o n was s e r i o u s l y damaged by frost.  first  three years i n the f i e l d  eight outplanting areas. Nlmpkish.  f o r f o u r t e e n provenances  E a r l y s u r v i v a l was  at  h i g h e s t near  A t most p l a n t a t i o n s , t r e e s from the l o c a l  grew as w e l l as those from the t h r e e b e s t s o u r c e s .  source Trees from  seed c o l l e c t e d a t the n o r t h e r n and southern extremes  (Nlmpkish  and B u t t e F a l l s , r e s p e c t i v e l y ) grew l e a s t i n the three years (as  they had i n the n u r s e r y ) .  Never damaged t r e e s were no more  than s l i g h t l y d i f f e r e n t i n h e i g h t growth from those t h a t had been damaged. Christmas t r e e growers s y s t e m a t i c a l l y e v a l u a t e d t r e e s In three l o c a t i o n s i n 1964 provenance  (L) was  (Douglass 1967)*  The Salem  found t o be the b e s t Oregon source and  S h e l t o n (G) the b e s t r e p r e s e n t a t i v e from Washington. ertson Valley investigated.  The Rob-  (D) o r i g i n ranked h i g h e s t i n a l l t h r e e l o c a t i o n s I t combined a number of d e s i r a b l e  qualities,  such as h i g h v i g o r , dark green c o l o u r , u p r i g h t growth h a b i t , n a t u r a l tendency  f o r b u s h l n e s s , and an a t t r a c t i v e needle  arrangement. Walters and Ching (1969) s t u d i e d the p a t t e r n of bud b u r s t of the s i x t e e n provenances* 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 and i n the Willamette V a l l e y .  s u l t s have been reviewed on page 12.  The r e -  In a r e p o r t on f i e l d  per-  formance a t age n i n e , Ching (1967) a s s e s s e d s u r v i v a l and h e i g h t growth i n t e n l o c a t i o n s .  Trees from Oakridge  (N) and from the  * Sources E ( 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 ) and F (Snoqualmle) were i n c l u d e d f o r the f i r s t time. Since t h e i r e s t a b l i s h m e n t i n 1959.  the f o l l o w i n g p l a n t a t i o n s  southernmost source P (Butte P a l l s ) were c o n s i s t e n t l y poor In h e i g h t growth.  Ching concluded t h a t h i g h e l e v a t i o n s t o c k  can  s a f e l y be p l a n t e d a t lower a l t i t u d e s , whereas i t would be  un-  wise t o do the o p p o s i t e . a t i o n and  P l o t t i n g h e i g h t growth a g a i n s t  elev-  l a t i t u d e d i d not r e v e a l a s i g n i f i c a n t c o r r e l a t i o n .  F i n a l l y , another I n v e s t i g a t i o n ( M i t c h e l l and  Nagel,  r e v e a l e d t h a t a t t a c k by D o u g l a s - f i r needle midges C o n t a r l n l a pseudotsugae Condrashoff) was  1969)  (mainly  c o r r e l a t e d t o date of  bud b u r s t , the damage b e i n g h i g h e s t on e a r l y f l u s h i n g t r e e s .  V.  The  METHODS  Study Area  General d e s c r i p t i o n of the U n i v e r s i t y of B r i t i s h Research F o r e s t The a t k9°l8*  Columbia  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 ,  l a t i t u d e n o r t h and  l22035f  l o n g i t u d e west, i s s i t u -  a t e d f o u r m i l e s n o r t h of Maple Ridge, f o r m e r l y  Haney, i n the  F r a s e r V a l l e y of B r i t i s h Columbia, on the s o u t h f r i n g e of  the  Coast Mountains, a t a d i s t a n c e of 36 m i l e s from the U n i v e r s i t y  have been l o s t i 1 . D - Robertson V a l l e y , b o t h p l a n t a t i o n s abandoned due t o f r o s t damage. 2 . G - Shelton. One p l a n t a t i o n l o s t t o grass f i r e . 3 . I - Tillamook. Both p l a n t a t i o n s h e a v i l y damaged by rabbits. 4. J - M o l a l l a . Both p l a n t a t i o n s s u f f e r e d from r a b b i t damage. 5 . N - Oakridge a r e a . Both p l a n t a t i o n s l o s t t o deer damage. 6 . P - Butte F a l l s . Both p l a n t a t i o n s l o s t t o drought. (Ching 1 9 6 9 . p e r s o n a l communication.)  of B r i t i s h Columbia Campus i n Vancouver mittee 1959)*  (U.B.C. F o r e s t Com-  I t comprises approximately twelve thousand  seven  hundred a c r e s o f f o r e s t e d l a n d , and i s bounded on the n o r t h and east by G a r i b a l d i P r o v i n c i a l Park and by P i t t Lake on the n o r t h E l e v a t i o n s range from sea l e v e l t o 2600 f e e t .  west.  The a r e a  l i e s w i t h i n the southern P a c i f i c c o a s t s e c t i o n ( C . 2 ) of the c o a s t a l f o r e s t a c c o r d i n g t o Rowe (1959) and i n the c o a s t a l western hemlock (Tsuga h e t e r o p h y l l a (Raf.) Sarg.) zone a c c o r d i n g to Krajlna (1959)•  The main a s s o c i a t i o n comprises the two  c o a s t a l dominants western r e d cedar (Thuja p l l c a t a Donn.) and western hemlock, w i t h c o a s t a l D o u g l a s - f i r and s c a t t e r e d western white p i n e , P a c i f i c s i l v e r f i r  (Abies a m a b l l l s (Dougl.) F o r b e s ) ,  S i t k a spruce ( P l c e a s l t o h e n s l s (Bong.) C a r r . ) and y e l l o w cedar (Chamaecyparls n o o t k a t e n s l s (D. Don) Spach) (Walters and Soos, I96lb).  Climate The c l i m a t e , d e s c r i b e d i n d e t a i l by G r i f f i t h  (1968)  who used d a t a from the f o u r weather s t a t i o n s on the Research F o r e s t , Is c o n s i d e r a b l y i n f l u e n c e d by the r e l a t i v e l y warm and moist a i r o f the P a c i f i c Ocean and by the Coast  Mountains.  The summers a r e warm and d r y , the w i n t e r s r e l a t i v e l y m i l d and wet.  Temperature  seldom reaches 90 degrees F a h r e n h e i t , the  a b s o l u t e minimum on r e c o r d b e i n g f i v e degrees F a h r e n h e i t below zero.  The annual p r e c i p i t a t i o n averages about 8 9 inches i n the  southern p a r t o f the F o r e s t , where the provenance t e s t s i t e i s l o c a t e d , b u t i n c r e a s e s i n the h i g h e r e l e v a t i o n areas t o the  north.  The h i g h amount of p r e c i p i t a t i o n has a pronounced  e f f e c t on the s o i l s which a r e s t r o n g l y leached podsols and a r e very a c i d i c . The weather s t a t i o n o u t s i d e the A d m i n i s t r a t i o n B u i l d i n g a t an e l e v a t i o n of 475 f e e t above sea l e v e l i s the one c l o s e s t t o the study a r e a , the c l i m a t e o f which can be assumed t o be p r a c t i c a l l y the same.  F o r the years 1 9 5 9 t o 1967 i n -  c l u s i v e , i . e . the y e a r s r e l e v a n t t o t h i s t h e s i s , the f o l l o w i n g d a t a were p u b l i s h e d by G r i f f i t h  (1968)t  Precipitation! Annual average October t o March A p r i l t o September J u l y ( d r i e s t month) December (wettest month) Winter s n o w f a l l Number o f days w i t h measurable p r e c i p i t a t i o n  8 9 . 0 9 " (Min. 75.88" - Max. 95.83") 63.57"or 71.35$ of the annual t o t a l 25.52"or  2.70"or l2.64"or 24.4"  28.65#  2.97$ 14.19$  *  192  Temperaturei Annual mean J u l y ( h o t t e s t month) January ( c o l d e s t month) Absolute minimum Absolute maximum Frost free period, average  4 9 . 1 ° F (Min. 4 7 . 6 degrees - Max. 63.2OF 5 1 . 0 degrees F a h r e n h e i t ) 35.9°F 2 OF  98°F  200 days (Min. 1 6 5 - Max. 243)  Average date o f l a s t s p r i n g f r o s t 1 Average date of f i r s t f a l l f r o s t i  April l4 November 1  May f r o s t s a r e f a i r l y frequent b u t September f r o s t s are r a r e .  F o r the f i v e - y e a r p e r i o d 1953-1957, average l e n g t h  of the growing season, as i n d i c a t e d by camblal a c t i v i t y  result-  i n g In diameter increment, was 146 days ( r a n g i n g from a minimum of 107 days t o a maximum of 163 days) a c c o r d i n g t o G r i f f i t h  (i960).  Since the c o - o p e r a t i v e provenance t e s t s i t e s , a t an  e l e v a t i o n of approximately  500 f e e t above sea l e v e l , are a t a  s l i g h t l y lower e l e v a t i o n than the weather s t a t i o n used by G r i f f i t h i n h i s i 9 6 0 study, the growing season there can assumed t o be  be  of s l i g h t l y l o n g e r d u r a t i o n .  L o c a t i o n of the c o - o p e r a t i v e provenance t e s t on the Research Forest P l a n t a t i o n I ( i . e . b l o c k s 1 and 2) tween Mainroad F and Spur A - 1 0 ; and 4 )  i s situated  be-  plantation II ( i . e . blocks 3  i s a d j a c e n t to Branch Road A on the e a s t e r n s i d e of the  North A l o u e t t e R i v e r .  Both p l a n t a t i o n s are w i t h i n l e s s  than  one-half mile of each other and l e s s than a mile from the A d m i n i s t r a t i o n B u i l d i n g i n the southeast c o r n e r of the Research (see F i g . 5 ) »  Forest  H i s t o r y of the t e s t  site  Prior to logging i n 1955-1956, the f o l l o w i n g volumes ( i n thousand board  the a r e a f e e t , B.C.  Live Dead Douglas Douglas L i v e Dead W.Hem-fir - f i r Cedar Cedar l o c k T o t a l Pltn  I*  Pltn II  15  9  95  4  132  255  260  -  435  69  208  972  The a r e a of p l a n t a t i o n I was which caused  *  relatively l i t t l e  1  supported rule)*  Vol/ acre  Cedar poles  18.4 43.2  logged by h i g h - l e a d  s o i l d i s t u r b a n c e , the a r e a of  Area was  p a r t of a 14 acre  stand.  Area was  p a r t of a 22 a c r e  stand.  69  p l a n t a t i o n I I was yarded by t r a c t o r , r e q u i r i n g an i n t e n s i v e network of skldroads slash, burning  r e s u l t i n g i n heavy s o i l d i s t u r b a n c e .  The  p o o r l y i n a f i r s t attempt, p a r t i c u l a r l y I n p l a n -  t a t i o n I , was p i l e d and r e - b u r n t  i n 1958 (Heaman 1 9 6 3 ) .  Since  the m a t e r i a l was c o l l e c t e d by hand and no heavy equipment was used f o r s i t e p r e p a r a t i o n , there was minimal mechanical disturbance.  Much d e b r i s remained.  Vine maple (Acer  soli  circln-  atum Pursh.) was s u c c e s s f u l l y sprayed i n s p r i n g 1959 w i t h 2,4-D, 2,4,5-T.  D r a i n i n g was c a r r i e d out i n p o r t i o n s o f p l a n t a t i o n I I  by b l a s t i n g a d i t c h . The The  nursery  s e e d l i n g s were p l a n t e d  p l a n t i n g stock  i n the s p r i n g of 1959«  ( 2 - 0 D o u g l a s - f i r ) was g e n e r a l l y of  poor q u a l i t y and d e s p i t e c l o s e s u p e r v i s i o n of the p l a n t i n g o p e r a t i o n , examination o f A r m l l l a r l a m e l l e a r o o t systems s e v e r a l years  ( F r ) Kumm. i n f e c t e d  l a t e r suggested t h a t the presence of  the d i s e a s e c o u l d p o s s i b l y be r e l a t e d t o a poor p l a n t i n g t e c h nique (Walters  1964).  The a r e a was covered by fireweed  (Eplloblum a u g u s t I f o l l u m  L.) s i x f e e t t a l l , b u t the r e a l problem  the f i r s t year was browsing by hares. line,  chemical  In a d d i t i o n t o a trap  r e p e l l e n t s were used In May 1959» b u t they d i d  not prove adequate I f used alone  (Walters  and Soos, 1 9 6 l a ) .  By 1 9 6 l I t appeared t h a t almost a l l o f the s e e d l i n g s had grown away from the browsing zone f o r hares, treatments were planned (Walters M o r t a l i t y was f a i r l y i n f a l l 1959. 380 d i e d i n i 9 6 0 , apply  t h e r e f o r e no a d d i t i o n a l  1961).  highi  3 6 5 t r e e s were found dead  and 140 i n 1 9 6 1 .  These f i g u r e s  only t o the "Inner b l o c k s " w i t h f o r t y - n i n e numbered  stakes  (see under " d e s i g n " ) . of I960 and 1961 of poor q u a l i t y .  A l l m o r t a l i t y was  r e p l a c e d i n the s p r i n g  by s e e d l i n g s t h a t were themselves In 1963  the a r e a was  frequently  weeded by machete and  b r u s h hooks, and then c o n c e n t r a t i o n s of v i n e maple and salmonb e r r y (Rubus s p e o t a b l l l s Pursh.) sprayed w i t h 2,4-D, and 2,4,5-T i n d i e s e l o i l (Walters 1964).  Vacolnlum spec,  was  never t r e a t e d but p o s s i b l y impeded t r e e growth wherever i t o c c u r r e d In t h i c k c o n c e n t r a t i o n s (see below). were a g a i n t r e a t e d c h e m i c a l l y i n August  1965*  The f o u r b l o c k s I n f e c t i o n s by  A r m l l l a r l a c o n t i n u e d d u r i n g t h a t year (Walters 1966). the s i x t e e n provenances time  (Walters 1967).  of I960,  1961.  1966  In  produced f l o w e r s and cones f o r the  first  Height growth had been a s s e s s e d a t the end  1964 and  1965.  Height Measurements For t h i s t h e s i s , the h e i g h t of a l l s u r v i v i n g numbered t r e e s was  measured a f t e r the 19.67 growing season when most t r e e s  were e l e v e n years o l d .  S i n c e t r e e s of the l o c a l source (Univ-  e r s i t y of B r i t i s h Columbia  Research F o r e s t ) and from Snoqualmle  (F) were one y e a r younger than the r e s t . I t seemed reasonable t o measure them one y e a r l a t e r , a l t h o u g h no two growing are q u i t e the same.  The primary o b j e c t i v e i n a n a l y z i n g the  h e i g h t measurements taken a t the U n i v e r s i t y of B r i t i s h Research F o r e s t , was  seasons  Columbia  t o determine whether s i g n i f i c a n t growth  d i f f e r e n c e s between provenances  were showing a t an e a r l y  age,  and I f so, t o t r y and c o r r e l a t e them w i t h geographic v a r i a b l e s . A secondary o b j e c t i v e was  t o e v a l u a t e the s i t e homogeneity a t  the Research F o r e s t .  S t a t i s t i c a l Analysis Height measurements f o r simple were a n a l y z e d  a n a l y s i s of  variance  on computers a t the U n i v e r s i t y of B r i t i s h Colum-  bia.  VI.  Height a t Age The  RESULTS AND  DISCUSSION  Eleven a n a l y s i s of v a r i a n c e  of h e i g h t measurements  showed the b l o c k x provenance I n t e r a c t i o n t o be  significant.  T h i s means t h a t t a) The i n d i v i d u a l provenances behave d i f f e r e n t l y the f o u r b l o c k s . b) The  p l a n t a t i o n s i t e Is not homogeneous.  c) The  f o u r b l o c k s have t o be e v a l u a t e d  In  separately.  d) No s t a t i s t i c a l c o n c l u s i o n as t o the o v e r a l l performance, such as s u p e r i o r growth, of any part i c u l a r provenance, can be drawn, i . e . the provenances cannot be ranked on a s t a t i s t i c a l l y sound b a s i s . e) U n l e s s the p r e r e q u i s i t e s f o r c o - v a r i a n c e a n a l y s i s can be met, which would a l l o w comparisons of o v e r a l l performance of a l l provenances, the r e s u l t s 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 do not j u s t i f y a t t h i s stage a c o n f i r m a t i o n or r e j e c t i o n of any hypothesis t h a t the c o - o p e r a t i v e study s e t out t o t e s t , nor do the r e s u l t s j u s t i f y a p o s i t i v e answer t o the q u e s t i o n whether s i g n i f i c a n t growth d i f f e r e n c e s between provenances are showing a t an e a r l y s t a g e . Simi l a r l y , seed t r a n s f e r r u l e s cannot be e s t a b l i s h e d w i t h these r e s u l t s . An attempt was  t h e r e f o r e made t o f i n d a q u a n t i f i a b l e  parameter r e l a t e d t o s i t e ,  t h a t would a l l o w an a c c u r a t e  explan-  a t i o n o f the d i f f e r e n t behaviour o f the I n d i v i d u a l provenances In the f o u r b l o c k s , a f t e r b r i n g i n g e v e r y t h i n g down t o the same basis.  Griffith  (i960) s t u d i e d the growth of D o u g l a s - f i r i n  r e l a t i o n t o c l i m a t e and s o i l a t the Research F o r e s t and found t h a t a v a i l a b l e s o i l moisture i n the B-horizon ing  d u r i n g the grow-  season was the most Important s i n g l e v a r i a b l e a f f e c t i n g t r e e  growth.  T h e o r e t i c a l l y i t i s easy t o get a v a l i d Idea of the  s o i l moisture regime of an a r e a by b r e a k i n g major i n d i c a t o r s i slope, aspect.  slope  i t down i n t o i t s  ( p e r c e n t ) , s o i l depth, p o s i t i o n on  I t would indeed be h e l p f u l t o know what p e r -  centage o f each p l o t  s u f f e r s from adverse c o n d i t i o n s , such as  e x c e s s i v e moisture or abnormal s o i l d i s t u r b a n c e ,  f o r instance.  By weighing the performance of each p l o t a c c o r d i n g l y , t h a t i s , by t r y i n g t o e l i m i n a t e v a r y i n g environmental I n f l u e n c e s , a b e t t e r p i c t u r e of the growth p o t e n t i a l o f t h a t p a r t i c u l a r provenance c o u l d be a r r i v e d a t . Mr.  The f o l l o w i n g map was drawn by  D. Ormerod, In an attempt t o i l l u s t r a t e  regime i n the f o u r b l o c k s Despite  moisture  (see F i g . 4 ) .  the help of the map, i t proved d i f f i c u l t i n  the f i e l d t o get an a c c u r a t e the extreme v a r i a b i l i t y various p l o t s .  the s o i l  estimate  of s o l i m o i s t u r e , due t o  I n microtopography even w i t h i n the  I t was f e l t t h a t such an e v a l u a t i o n would t o o  e a s i l y be s u b j e c t i v e and t h e r e f o r e i n c o r r e c t and inadequate. Mr.  *  L. L a c e l l e , a graduate student  P l o t means any provenance i n any b l o c k .  In s o i l  science  ?LANTA  T/DM  T  PlAA/r/ff/o// Jl  BLOCK  1  BLOCK "I  BLOCK JL  *  LPiCSLLE's  8*. ock  iy  SOIL PIT  t UILL/NQ rows  APPK.OX.  SCALZ--  /"* 3 CM/AJS  LECf€AJl)  V  V  v  see/>A<5_ StJftMP  V  2>^/e>e  Figure  i  -6  Ormerod's  <5*Avei  SLl<fH-T  RiHS  i 3*us/f  Hirer  4  •exposed  map  SOIL PIT  a t the U n i v e r s i t y of B r i t i s h Columbia, k i n d l y dug s e v e r a l  soil  p i t s i n b o t h p l a n t a t i o n s and came t o the c o n c l u s i o n t h a t the s o i l was roughly  the same i n a l l f o u r b l o c k s , except f o r s m a l l  areas t h a t showed the e f f e c t s o f mechanical d i s t u r b a n c e  due t o  l o g g i n g , and a l s o e x c e p t i n g areas w i t h a d i f f e r e n t water regime due  t o t h e i r microtopography.  Mr. L a c e l l e d e s c r i b e d  the s o i l  i n b l o c k B, p l a n t a t i o n I , as a m l n i - o r t h i c - h u m o f e r r i c moderately w e l l d r a i n e d t o w e l l d r a i n e d .  According  podzol.  to Willing-  ton ( 1 9 6 8 ) , the s o i l i n b l o c k C, p l a n t a t i o n I I , i s a moderately well drained  o r t h i c p o d z o l , developed i n outwash parent  i a l o v e r l y i n g Whatcom g l a c l o m a r i n e .  mater-  Walters ( 1 9 7 0 ) t h i n k s t h a t  the cemented l a y e r u n d e r l y i n g the s o i l a t v a r y i n g depths, has t o take p a r t o f the blame f o r the v a r i a b i l i t y  i n s i t e i n some  p l o t s by a f f e c t i n g the d i r e c t i o n o f the seepage A co-variance  flow.  a n a l y s i s t h e r e f o r e c o u l d n o t be c a r r i e d  out due t o the l a c k of a d e t a i l e d a n a l y s i s o f the p h y s i c a l , chemical  and h y d r o l o g l c a l s o i l p r o p e r t i e s o f the d i f f e r e n t  p l o t s which would each r e q u i r e a g r e a t number o f samples due t o the extreme v a r i a b i l i t y i n microtopography.  I t i s hoped t h a t  such a study w i l l be made and more c o n c l u s i v e i n f o r m a t i o n  will  be gained  from the provenance study a t the Research F o r e s t .  According  t o a personal  uate student  communication by Ching ( 1 9 7 0 ) ,  a grad-  of Oregon State U n i v e r s i t y , c o l l e c t e d many  samples from e l e v e n  outplanting areas.  soil  His r e s u l t s w i l l  prob-  a b l y throw some l i g h t on the e f f e c t of microedaphic d i f f e r e n c e s on t r e e growth.  Heaman ( 1 9 6 3 ) p o i n t e d  out the s i t e  variability  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  e x e m p l i f i e d by a g u l l e y i n p l a n t a t i o n I , c a t e r p i l l a r roads, a g r a v e l p i t , and swampy areas i n p l a n t a t i o n I I .  However i t  should be noted t h a t , g i v e n the l i m i t e d a r e a and the knowledge a t the time of the s o i l s , the s i t e chosen was by f a r the b e s t one a v a i l a b l e .  In another D o u g l a s - f i r provenance study a t the  Research F o r e s t , Haddock e t a l . ( 1 9 6 7 ) found a s i g n i f i c a n t provenance x b l o c k i n t e r a c t i o n , t o o , r e f l e c t i n g p o s s i b l y the heterogeneous  " ...  nature of the s i t e , t y p i c a l f o r c o a s t a l  mountain country i n B r i t i s h  Columbia."  R e s u l t s by I n d i v i d u a l B l o c k s Separate a n a l y s e s o f v a r i a n c e f o r h e i g h t  performance  were c a r r i e d o u t . The r e s u l t s a r e l i s t e d i n Table 3« The wide range, s t r i k i n g l y e x e m p l i f i e d by three provenances (Nlmpkish, T i l l a m o o k and h i g h e r e l e v a t i o n  Molalla),  showing a d i f f e r e n c e of t h i r t e e n ranks, out o f a p o s s i b l e maximum of f i f t e e n , between t h e i r s h o r t e s t and t h e i r t a l l e s t  per-  formance, can be I n t e r p r e t e d t o mean e i t h e r t h a t t 1. The g e n e t i c v a r i a b i l i t y  of D o u g l a s - f i r i s such t h a t  f o u r r e p l i c a t i o n s of 4 9 t r e e s each f o r every provenance a r e n o t enough t o cover i t . To support t h i s one c o u l d p o i n t t o the f a c t  that,  when w a l k i n g through the study a r e a a t the Research F o r e s t , one i s s t r u c k n o t only by the v a r i a b i l i t y i n growth between some provenances  (e.g. seed source 4 , Nlmpkish,  and 1 6 ,  Snoqualmle  i n b l o c k I I I ) b u t a l s o by the f a c t t h a t i n every p l o t , no matter how poor I t s average  performance,  there i s a t l e a s t one  t r e e d o i n g much b e t t e r than the r e s t .  The b e s t example of t h i s  can be found i n b l o c k IV, p l o t 4, Nlmpkish, where one p a r t i c u l a r Ranking o f Provenances Block Provenance # 1=G 2=B 3=D 4=A 5=H 6=G 7=1 8=L 9=J 10=K 11=M 12=N 13=0 14=p 15=E l6=F  Origin Sugar Loaf Mountain, Vancouver I s l a n d Courtenay a r e a , Vane. I s l a n d Robertson V a l l e y , " . "' Nlmpkish V a l l e y , " E l b e a r e a , Washington Shelton area, " Tillamook area, Oregon Salem a r e a , " M o l a l l a , lower " elevation Molalla, higher " elevation C o r v a l l l s area, " Oakridge, lower " elevation Oakridge, h i g h e r " elevation Butte F a l l s , " U.B.C. Research F o r e s t , B r i t i s h Columbia Snoqualmle a r e a , Washington  Table 3  Average Range  I  II  III  IV  6  3  1  8  ^.5  7  8 3 10 12 4 5 11 13  2 8 11 7 1 6 13 5  7 13 2 8 3 16 12 11  12 9 15 5 4 3 6 10  7.2 8.2 9.5 8.0 3.0 7.5 10.5 9.8  10 10 13 7 3 13 7 8  2  15  15  2  8.5  9 14  9 16  9 4  14  16  4  14  7  8.2  12  15 1  12 10  10 5  16 1  13.2 4.2  6 9  7  14  6  11  9.5  8  10.0 12.0  13 4 12  *  Height performance a t age e l e v e n . Ranking i s from t a l l e s t ( l ) t o s h o r t e s t ( l 6 ) .  t r e e i s growing v i g o r o u s l y i n a swampy p l a c e , w i t h most o f i t s neighbours  l o o k i n g c h l o r o t i c and s t u n t e d i n a p p a r e n t l y the same  micro-environment.  T h i s I s an I n t e r e s t i n g p a r a l l e l t o a f i n d i n g  In the 1912 D o u g l a s - f i r h e r e d i t y study by the U.S. F o r e s t Servi c e (1964)i  "An anomaly t o the i n t e r a c t i o n e x i s t s I n the  f o l l o w i n g sensei  Almost without  e x c e p t i o n the progenies o f  every parent i n c l u d e a t l e a s t one s u p e r i o r i n d i v i d u a l a t each The l o c a l provenance.  location.  This indicates s u r p r i s i n g genetic d i v e r s i t y within  a r e s t r i c t e d source of germ-plasm such as a s i n g l e seed l o t . " The wide range i n r a n k i n g c o u l d a l s o mean that« 2.  T h i s s i t e i s extremely  heterogeneous.  This a l t e r n -  a t i v e i s supported by the s i g n i f i c a n c e of the i n t e r a c t i o n term and  i l l u s t r a t e d f o r i n s t a n c e by  a b o v e - c i t e d source 4 tallest  (Nlmpkish).  I t i s second  i n b l o c k I I I where I t s o u t s t a n d i n g  ance can be d e t e c t e d from a 1 photograph.  the  perform-  i 6300 s c a l e a e r i a l  In b l o c k IV I t i s the t h i r d  poorest  i n terms of h e i g h t growth, probably because i t has t o grow i n a r a t h e r swampy p l a c e . A e r i a l photographs of the study a r e a show evidence In f a v o u r of the  " s i t e h e t e r o g e n e i t y " - r a t h e r than the  etic variability" alternative.  Patches  of poor growth (marked  w i t h "P" on F i g u r e 5) show up v e r y w e l l . boundaries  "gen-  They i g n o r e p l o t  and q u i t e l i k e l y r e f l e c t edaphic d i f f e r e n c e s . Tables 4 t o 7 show height performance w i t h i n the  i n d i v i d u a l blocks f o r 1967. comparison.  1964  and 1961  are i n c l u d e d f o r  The b r a c k e t s show the r e s u l t s of Duncan's  New  M u l t i p l e Range T e s t .  Early  Tests T r y i n g t o f i n d c o r r e l a t i o n s between the i 9 6 0 ,  and 1967  measurements ( a t age 4 ,  8 and 11  y e a r s ) i s almost  e x e r c i s e In f u t i l i t y f o r the f o l l o w i n g reasons 1 1.  Duncan's New  1964  M u l t i p l e Range T e s t shows t h a t  an  /'//  Figure 5  L o c a t i o n map of the c o - o p e r a t i v e provenance t e s t 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 .  Prov. Rank  10 2 1 15 3 6 11 14 16 7 5 9 4 12 8 13  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16  Ht(cm)  % of Mean  Prov.  49.27 39.71 38.92 38.02 37.65 37.57 36.47 35.67 34.59 34.31 33.43 33.31 32.84 32.14 31.49 20.65  139 112 110 108 106 106 103 101 98 97 95 94 93 91 89 58  15 10 3 6 _ 1 7 2 11 16 4 8 5 9 14 [12 13  Table 4 Any  two provenances  Ht(m)  % of Mean  2.5980 2.5927 2.3959 2.3857 2.3653 2.2687 2.2330 2.1787 2.1511 2.1142 2.0777 2.0719 1.9826 1.8511 1.7789 1.5447  125 119 110 109 109 104 103 100 99 97 96 95 91 85 82 71  Height performance.  Prov. 15 10 3 6 7 1 16 2 11 4 8 5 9 12 14 13  Ht(ra)  % of Mean  Rank  5.7333 5.2458 5.1694 5.1163 4.7766 4.7633 4.6405 4.6045 4.5410 4.5349 4.4957 4.4043 4.2523 3.8682 3.7109 3.6081  124 114 112 111 103 103 100 100 98 98 97 95 92 84 80 78  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16  Block I  covered by the same b r a c k e t a r e not s i g n i f i c a n t l y  different.  OS  Prov. Rank  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16  6 5 1 2 9 7 13 11 14 3 10 4 12 8 15 16  Ht(cm)  % of Mean  47.76 43.94 43.81 41.71 40.39 39.96 39.89 37.33 35.47 34.85 31.35 30.33 29.12 28.53 27.39 26.58  132 121 121 115 112 110 110 103 98 96 87 84 80 79 76 73 Table 5  Any  two provenances  Ht(ra)  of Mean  2.7354 2.4056 2.3833 2.3830 2.3605 2.3372 2.3262 2.1228 2.0292 2.0273 1.9939 1.9622 1.9221 1.9156 1.8323 1.6664  127 112 111 111 110 109 108 97 94 94 93 91 89 89 85 77  Ht(m)  % of Mean  Rank  5.2543 5.0133 4.9556 4.8581 4.8045 4.7905 4.6804 4.5974 4.4109 4.4043 4.3667 4.3200 4.3111 4.0958 4.0673 3.8930  116 110 109 107 106 105 103 101 97 97 96 95 95 90 89 86  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16  7o  Prov. 6 "2 9 1 7 5 13 14 11 3 10 8 4 15 16 12  Height performance.  Prov. 6 2 1 13 9 7 5 3 11 15 4 14 8 16 10 12  Block II  covered by the same b r a c k e t a r e not s i g n i f i c a n t l y  different.  1960  1964  Prov. 6  1  40.61  of Mean  Prov.  Ht(m)  133  1  2.4781  134  4  2.3553  128  2.3178  126  2  4  38.35  125  3  1  36.71  120  _ 6  of Mean 7o  7»  of Mean  7o  Ht(cm)  1967  Prov.  fl r-L  4  Ht(m)  Rank  5.1375  132  1  5.0000  128  2  6  4.3238  111  3  4.0804  105  4 5  4  2  34.47  115  11  1.9087  103  12  5  5  34.10  111  14  1.8967  103  15  4.0390  103  16  3.9786  102  6 7 8  6  11  —  31.81  2  104  1.8890  102  101  12  1.8670  101  2  3.8317  98  98  5  1.8062  98  5  3.8062  98  8  1.7512  95  11  3.7111  95  9  3.6795  94  10  3.6565  94  11  8  3.6325  93  12  89  3  3.5980  92  13  1.5844  86  13  3.2907  84  14  1.4844  80  10  3.2675  84  15  1.3400  73  7  3.2646  84  16  _  7  30.86  8  3  30.  9  14  10  12  29.96  98  15  1.7354  94  14  9  29.29  96  16  1.7198  93  9  26.17  85  9  1.6796  91  25.90  85  3  1.6459  24.42  80  7  7  11  8  12  13  13 14  _ 1 6  10  29.96  98  .  15  15  23.77  78  13  16  10  21.78  71  10  Table 6 Any  two provenances  Height performance.  Block I I I  covered by the same bracket a r e not s i g n i f i c a n t l y  different.  Prov. Rank  10 7 15 5 ' 1 3 6 2 9 13 "16 4 12 8 11 14  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16  Ht(cra)  % of Mean  43.88 41.00 40.67 38.22 33.30 32.86 32.50 31.94 31.39 30.06 27.32 26.66 26.47 25.79 24.67 22.69  137 128 127 120 104 103 102 100 98 94 85 83 83 81 77 71 Table 7  Any  two provenances  Prov.  .[Do  7 5 8 6 1 13 3 9 11 2 16 12 4 14  Ht(m)  X of Mean  2.7622 2.6435 2.4098 2.2940 2.1596 2.1000 2.0600 2.0284 1.8378 1.7937 1.7735 1.7232 1.7170 1.6625 1.5133 1.4685  137 131 120 114 107 104 102 101 91 89 88 86 85 83 75 73  Height performance.  7c o f Prov. 15  rio L 7 _  r  6 5 8 13 1 3 9 16 2 11 12 4 14  Ht(m)  Mean  Rank  5.9204 5.1404 4.9128 4.5889 4.5755 4.4553 4.3683 4.0911 4.0347 3.8612 3.8372 3.7275 3.6163 3.6159 3.2441 2.9545  140 122 117 109 109 106 104 97 96 92 91 88 86 86 77 70  1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16  B l o c k IV  covered by the same bracket a r e not s i g n i f i c a n t l y  different.  -N3  the provenances do not respond four blocks.  s i m i l a r l y i n the  2. Three o b s e r v a t i o n s of such v a r i a b i l i t y are not enough f o r s i g n i f i c a n t r e g r e s s i o n e q u a t i o n s . Assuming these o b s e r v a t i o n s t o be on a s t r a i g h t l i n e i s making a very rough approximation. I t was  t h e r e f o r e not s u r p r i s i n g t h a t m u l t i p l e r e -  g r e s s i o n a n a l y s i s d i d not l e a d to any Although  spectacular findings.  the t r e n d i n h e i g h t growth w i t h i n the same b l o c k seems  t o be comparable, the s l o p e s are s i m i l a r , the h e i g h t s thems e l v e s are not, except w i t h i n very s m a l l groups of two provenances, s i n c e the i n t e r c e p t s are s i g n i f i c a n t l y  or three  different.  Even i f the s i t e were homogeneous the t r e e s might s t i l l be young f o r meaningful  Brush  early  too  tests.  Competition Vine maple, salmonberry  and  other s p e c i e s were r e p e a t -  e d l y t r e a t e d c h e m i c a l l y and m e c h a n i c a l l y them from competing w i t h D o u g l a s - f i r . Vacclnlum  t a l l , account  few y e a r s .  There i s no doubt t h a t  spp. o c c u r r i n g i n s e v e r a l c o n c e n t r a t i o n s (marked  f o r brush on Ormerod's map,  geneity."  i n an e f f o r t to prevent  page 63)  and  growing up t o f o u r f e e t  f o r some of the v a r i a b i l i t y termed " s i t e  The experimental  "b"  hetero-  t r e e s outgrew t h i s b r u s h i n the  Table 8 shows the h e i g h t increment  of the l a s t  last three  growing seasons ( 1 9 6 ^ - 1 9 6 7 ) b e f o r e the measurements f o r t h i s t h e s i s were taken.  T h i s d i f f e r e n c e was  c a l c u l a t e d i n an attempt  t o e l i m i n a t e , i n theory, the e f f e c t of brush c o m p e t i t i o n h e i g h t growth.  on  I t would be i n c o r r e c t to assume t h a t once the  t r e e s are t a l l e r than the brush, they grow completely  unimpeded.  Vacclnlum  spp. i s d e f i n i t e l y s t i l l competing f o r s o i l  and n u t r i e n t s .  N e v e r t h e l e s s , the h e i g h t increment  three years shows a s l i g h t l y d i f f e r e n t  moisture  i n those  r a n k i n g f o r many p l o t s  where some t r e e s o b v i o u s l y s u f f e r e d from b r u s h c o m p e t i t i o n .  It  Ranking of Provenances Block Provenance # 1=C 2=B 3=D 4=A 5=H 6=G 7=1 8=L 9=J 10=K 11=M 12=N 13=0 l4=P 15=E l6=F  Origin Sugar L o a f Mountain, Vancouver I s l a n d Courtenay a r e a , Vane. I s l a n d Robertson V a l l e y , " Nlmpkish V a l l e y , " Elbe a r e a , Washington Shelton area, " Tillamook area, Oregon Salem a r e a , " M o l a l l a , lower " elevation M o l a l l a , higher elevation C o r v a l l l s area, " Oakridge, lower " elevation Oakridge, higher " elevation Butte F a l l s , U.B.C. Research F o r e s t , B r i t i s h Columbia Snoqualmle a r e a , Washington  Table 8  Average Range  I  II  III  IV  11  3  1  10  6.2  10  12 2 6 8 3 7 9 13  1 5 7 12 6 8 10 9  10 12 2 8 6 16 9 7  13 8 14 7 4 3 6 11  9.0 6.8 7.2 8.8 4.8 8.5 8.5 10.0  12 10 12 5 3 13 4 6  4  16  11  2  8.2  14  10 14  11 14  '2  15  4  16 1 5  15 12  12.8 11.0  5 10  14  5  9.5  11  15 2  13 3  16 1  15.0 1.8  3^ 2*  13  5  9  8.0  8  Height performance between the years 1964-1967. Ranking i s from t a l l e s t (1) t o s h o r t e s t ( l 6 ) .  i s t r u e , however, t h a t the provenance x b l o c k I n t e r a c t i o n i s still  significant  Vacclnlum  i n the a n a l y s i s o f v a r i a n c e , meaning t h a t  cannot take a l l the blame f o r the h e t e r o g e n e i t y i n  site.  The  l o c a l provenance.  The L o c a l Provenance In terms of h e i g h t increment i n c l u s i v e , the l o c a l source  f o r the years  1965-1967  ( U n i v e r s i t y of B r i t i s h Columbia Re-  s e a r c h F o r e s t ) a l l of a sudden ranks second t a l l e s t i n b l o c k I I where I t h e l d a meager t e n t h p o s i t i o n f o r t o t a l h e i g h t , when brush was n o t taken i n t o account. f i f t h to third position. the narrowest  In b l o c k I I I i t moves from  Of a l l provenances t e s t e d , i t shows  range (from t h i r d p o s i t i o n i n b l o c k I I I over  second i n b l o c k I I t o f i r s t  i n b l o c k s I and I V ) .  One i s tempt-  ed t o c a l l i t " the b e s t of a l l provenances as t o o v e r a l l  per-  formance I f i t were n o t f o r the harsh laws o f s t a t i s t i c s  for-  b i d d i n g t h i s c o n c l u s i o n a g a i n s t the background r e f r a i n of a s i g n i f i c a n t b l o c k x provenance i n t e r a c t i o n .  The performance of  the l o c a l source a t t h i s stage might i n d i c a t e t h a t s e v e r a l y e a r s from now the e f f e c t s of temporary " j u v e n i l e handicaps"  such as  brush c o m p e t i t i o n , f o r i n s t a n c e , w i l l d i s a p p e a r and i n h e r e n t d i f f e r e n c e s t h a t the c o - o p e r a t i v e study s e t out t o t e s t , appear.  I t may a l s o be s p e c u l a t e d t h a t the i l l  will  e f f e c t s of s o i l  compaction caused by l o g g i n g , as i l l u s t r a t e d by Pearse  (1958),  may g r a d u a l l y fade as the r o o t s l o o s e n the s o i l . In t a b l e s 4 t o 7 t h i s  "upwards" t r e n d o f the l o c a l  source can be f o l l o w e d f o r the years In b l o c k  1960-1964-1967.  I i t moves from 4 t h p o s i t i o n II " " " 15th " III " " " 15th " IV " " " 3rd "  In b l o c k s I and IV the l o c a l source  #  Brush c o m p e t i t i o n i s s t i l l a problem In t h i s  to 1st " 10th* " 5th " 1st  ( U n i v e r s i t y of  plot.  B r i t i s h Columbia  Research F o r e s t ) i s s i g n i f i c a n t l y  different  from ( i . e . s u p e r i o r t o ) a l l other sources a f t e r the 1967 i n g season. provenance  Table 6 a l s o shows t h a t every B r i t i s h ranks a t l e a s t once, f i r s t  any provenance  grow-  Columbia  or second t a l l e s t .  If  can be r a t e d as i n f e r i o r i t w i l l be B u t t e F a l l s ,  T h i s i s n o t s u r p r i s i n g c o n s i d e r i n g the d i f f e r e n c e In c l i m a t e between Butte F a l l s and the U n i v e r s i t y o f B r i t i s h Columbia s e a r c h F o r e s t where the temperature pronounced  than i n southern Oregon.  Re-  regime d i f f e r e n c e s a r e more The B u t t e F a l l s  provenance  Is d o i n g p o o r l y i n every b l o c k , even i n b l o c k C, where i t can be s a i d t o p r o f i t from a s l i g h t amount o f seepage water.  How do These R e s u l t s Compare w i t h E a r l i e r R e s u l t s ? Ching ( 1 9 6 5 ) i n h i s r e p o r t on e a r l y growth, d i d n o t f i n d any s i g n i f i c a n t d i f f e r e n c e s between provenances U n i v e r s i t y of B r i t i s h Columbia 1961.  a t the  Research F o r e s t f o r i 9 6 0 and  He l i s t e d the sources as t o t h e i r o v e r a l l  w i t h o u t mentioning the provenance  performance,  x b l o c k i n t e r a c t i o n which was  found t o be s i g n i f i c a n t i n the p r e s e n t study.  In h i s summary,  Ching s t a t e d t h a t " ... t r e e s from the l o c a l seed source grew as w e l l as d i d those from the b e s t three s o u r c e s . " a f t e r the 1967  The r e s u l t s  growing season a t the U n i v e r s i t y of B r i t i s h  umbia Research F o r e s t c e r t a i n l y do n o t d i s p r o v e t h a t  Col-  statement  and i t Is f e l t t h a t p o s s i b l y the l o c a l source w i l l m a i n t a i n i f not improve  i t s position i n future years.  The f i n d i n g s of Ching  t h a t " t r e e s from seed c o l l e c t e d a t n o r t h e r n and southern extremes of the study grew l e a s t In the t h r e e y e a r s " a r e supported  by the p r e s e n t t h e s i s f o r the southernmost provenance (Butte P a l l s ) , whereas the northernmost source v e r y w e l l a t l e a s t In one b l o c k . maintained  (Nlmpkish) I s doing  Seed source G (Shelton) has  I t s p o s i t i o n as one of the v e r y b e s t .  Ching and  Bever ( i 9 6 0 ) as w e l l as Heaman ( 1 9 6 3 ) d i d n o t f i n d any c o r r e l a t i o n between h e i g h t growth and geographic as l a t i t u d e and a l t i t u d e .  Nothing was found  study t o support such a c o r r e l a t i o n  v a r i a b l e s such i n the present  either.  VII. EVALUATION OF THE CO-OPERATIVE PROVENANCE TEST  Good P o i n t s The  i n i t i a t i v e t o convince a l a r g e p o r t i o n of p r i v a t e  and p u b l i c f o r e s t Industry i n the P a c i f i c Northwest, I n c l u d i n g B r i t i s h Columbia, of the importance of the provenance problem, to  spend money, time and e f f o r t on t h e c o - o p e r a t i v e  c e r t a i n l y deserves  praise.  study,  The attempt t o f i n d an answer t o  p r a c t i c a l and t h e o r e t i c a l a s p e c t s of the provenance problem was honest. ity  The mistakes  made (see below) I l l u s t r a t e the complex-  o f the problem and help t o make the t a s k e a s i e r f o r i n v e s -  t i g a t i o n of f u t u r e provenance t e s t s .  Weak P o i n t s The of  f o l l o w i n g e v a l u a t i o n i s n o t meant t o be c r i t i c a l  the o r i g i n a t o r s o f the study, b u t should be i n t e r p r e t e d as  a help f o r f u r t h e r provenance  trials.  Since  temperature and  f r o s t f r e e p e r i o d were to be  the major c r i t e r i a i n determining  the e x i s t e n c e  of d i s t i n c t  " r a c e s " of c o a s t a l D o u g l a s - f i r , more emphasis should have been put on the s t a n d a r d i z a t i o n of weather measurements (see Heaman's c r i t i c i s m , page 51)•  In such a mountainous r e g i o n as the  covered by the c o - o p e r a t i v e  provenance study, mere a l t i t u d e and  l a t i t u d e , or t h e i r combination (see page 39)  "hypotheses t o be  tested,"  cannot be expected t o d e s c r i b e or t o determine t r u e  climatic differences.  Aspect, s l o p e , l o c a l topography, proxim-  i t y of the ocean, e t c . are too i n f l u e n t i a l i n modifying e f f e c t s of a l t i t u d e and area.  area  The  the  latitude, especially within a limited  a r e a covered was  too l a r g e and  the number of proven-  ances i n c l u d e d too s m a l l f o r meaningful c l u e s on the nature the v a r i a t i o n w i t h i n c o a s t a l D o u g l a s - f i r . sampling was from 750  was  altitudlnal  not uniform, s i n c e no r e p r e s e n t a t i v e was  t o 1250  c a r r i e d out.  The  The  of  selected  f e e t e l e v a t i o n , where much l o g g i n g i s b e i n g cone c o l l e c t i o n r a d i u s of t w e n t y - f i v e  miles  too l a r g e , areas of great c l i m a t i c d i f f e r e n c e s can  be  covered by a f i f t y - m i l e diameter, as i l l u s t r a t e d by Heaman ( 1 9 6 3 ) f o r the a r e a around Lake Cowichan on Vancouver I s l a n d . Besides  t h a t , the cones were not c o l l e c t e d a c c o r d i n g t o  same standards by a l l c o - o p e r a t o r s .  One  c o u l d almost  the  say  t h a t a provenance t e s t i s as good as the t e s t s i t e s .  If  these are heterogeneous (as 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 t e s t s i t e i s , f o r i n s t a n c e ) i t i s very  diffi-  c u l t t o t e s t any  hypothesis  a l t h o u g h the d i f f i c u l t y  related to various  provenances,  i n f i n d i n g homogeneous s i t e s , l a r g e  enough t o accomodate two  three-acre  p l a n t a t i o n s i n the rugged  topography on the P a c i f i c c o a s t , i s f u l l y a p p r e c i a t e d . by hare and deer c o u l d have been p a r t l y avoided The  Damage  through  fences.  f r o s t problem In the Robertson V a l l e y c o u l d have been r e -  cognized  by more c a r e f u l study of e x i s t i n g weather d a t a .  VIII.  Since  CONCLUSIONS AND  the a n a l y s e s  RECOMMENDATIONS  of v a r i a n c e show a  significant  provenance x b l o c k i n t e r a c t i o n , no c o n c l u s i o n as t o o v e r a l l performance of any  p a r t i c u l a r provenance can be drawn from the  measurements taken a t the U n i v e r s i t y of B r i t i s h Columbia s e a r c h F o r e s t when the t r e e s were e l e v e n years  old.  of t h i s , none of the hypotheses t h a t the c o - o p e r a t i v e ance study s e t out t o t e s t can be accepted  Re-  Because proven-  or r e j e c t e d .  Sig-  n i f i c a n t growth d i f f e r e n c e s between provenances do show up w i t h i n I n d i v i d u a l b l o c k s , but  the r e s u l t s are i n c o n s i s t e n t and  t h e r e f o r e not c o n c l u s i v e , due  t o the extreme s i t e  eity.  heterogen-  Observed, but s t a t i s t i c a l l y not proven d i f f e r e n c e s tend  t o I n d i c a t e t h a t the l o c a l provenance i s by no means i n f e r i o r t o any  other seed source t e s t e d , and  prove t o be the b e s t i t c o u l d be  one  i n f u t u r e years  i t might  of a l l i n terms of height growth, as  shown t o move up the ranks over the l a s t  years.  Thus, the l o n g - h e l d view t h a t l o c a l seed i s s a f e s t , and ably best  (e.g. Bingham, 1 9 6 6 ) ,  Is s t i l l v a l i d .  The  prob-  southern-  most provenance i s d o i n g very p o o r l y 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 .  It i s possible that several  years from now  the e f f e c t s of J u v e n i l e handicaps,  such as  brush  c o m p e t i t i o n and maybe s o i l compaction, w i l l have faded away and g e n e t i c d i f f e r e n c e s , i f any, w i l l show up The r e s u l t s found provenance problem and geneous s i t e .  statistically.  I n d i c a t e the complexity  of the  the overwhelming i n f l u e n c e of a h e t e r o -  F u r t h e r provenance s t u d i e s are necessary  to i n -  v e s t i g a t e the nature and extent of v a r i a t i o n w i t h i n c o a s t a l D o u g l a s - f i r and  t o e s t a b l i s h d e t a i l e d seed t r a n s f e r r u l e s  seed c o l l e c t i o n zones.  and  Based on the f i n d i n g s 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 and  on the weak p o i n t s of  the c o - o p e r a t i v e t e s t , the f o l l o w i n g recommendations f o r provenance t e s t s seem l o g i c a l 1.  General  (see a l s o Heaman, 1 9 6 3 ) «  recommendations  a) S t a n d a r d i z a t i o n i n a l l phases of a c o - o p e r a t i v e study i s i m p e r a t i v e .  C o n t r o l s are necessary  and  should be e x e r c i s e d by one c o - o r d i n a t o r , i n constant personal contact with a l l co-operators. b) I n t e n s i v e sampling  of a s m a l l e r a r e a than t h a t  covered by the c o - o p e r a t i v e study, i s l i k e l y  to  y i e l d more i n f o r m a t i o n . c) Cone c o l l e c t i o n areas should be c l e a r l y d e f i n e d , based on a thorough l o c a l knowledge. d) O u t p l a n t i n g areas have t o be  thoroughly s t u d i e d .  Besides topography and v e g e t a t i o n , c l i m a t i c should be looked I n t o , the s o i l should be  data  analyzed  i n an attempt t o have a t l e a s t homogeneous b l o c k s . e) More s t u d i e s should be c a r r i e d out t o f i n d  one  criterion for soil fertility,  or f o r s i t e  differ-  ences, so t h a t b e t t e r s t a t i s t i c a l r e s u l t s may  be  obtained. f ) Weather measurements should be supplemented  with  phenological observations. g) Damage from animals and f u n g i has t o be kept to a minimum. 2.  Recommendation f o r management P l a n t l o c a l seed.  C o l l e c t i t from s e l e c t e d  d u r i n g good crop years and s t o r e i t .  T h i s way  trees  unpleasant  s u r p r i s e s due t o seed c r o p f a i l u r e s and purchase of seed of unknown o r i g i n , can be a v o i d e d .  IX.  Importance,  SUMMARY  scope and nature of the provenance  lem a r e d i s c u s s e d i n t h i s t h e s i s . provenance America  Experience gained from major  t r i a l s w i t h D o u g l a s - f i r i n both Europe and  i s reviewed.  of D o u g l a s - f i r are apparent.  "Co-operative D o u g l a s - f i r provenance  d i s c u s s e d In d e t a i l .  North  From t h i s , the complexity of the problem  and the extreme g e n e t i c v a r i a b i l i t y The  prob-  I t was  s t a r t e d i n 1957  test" i s  and i n c l u d e d  six-  teen c o a s t a l o r i g i n s from Oregon, Washington and B r i t i s h Columb i a , p l a n t e d c l o s e t o the seed c o l l e c t i o n a r e a s , one of which i s l o c a t e d on the U n i v e r s i t y of B r i t i s h Columbia F o r e s t , where h e i g h t growth was e l e v e n years o l d .  Research  measured when the t r e e s were  The o b j e c t i v e s i n a n a l y z i n g these data were  to f i n d whether  s i g n i f i c a n t d i f f e r e n c e s between provenances show  a t an e a r l y stage and whether 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 t e s t s i t e i s homogeneous. The s t a t i s t i c a l a n a l y s i s shows a s i g n i f i c a n t provenance x b l o c k i n t e r a c t i o n , i n d i c a t i v e of a heterogeneous  site.  The r e s u l t s a r e very c o n f u s i n g and no c o n c l u s i o n can be drawn y e t as t o o v e r a l l performance of any i n d i v i d u a l  provenance.  A f t e r removing the e f f e c t s of b r u s h , randomly s c a t t e r e d over the t e s t s i t e , the r e s u l t s tend t o become more m e a n i n g f u l . A l though the i n t e r a c t i o n term i s s t i l l c o n c l u s i o n s a r e drawn.  s i g n i f i c a n t , the f o l l o w i n g  The l o c a l provenance i s by no means i n -  f e r i o r , and i t might prove t o be the b e s t one of a l l  i n a few  years when b r u s h c o m p e t i t i o n w i l l have become l e s s important. The southernmost provenance from Butte F a i l s , Medford, is inferior to a l l  other o r i g i n s  Oregon,  tested.  I t i s hoped t h a t the measurements taken a f t e r the 1970  growing season by a l l c o - o r d i n a t o r s w i l l y i e l d more con-  c l u s i v e I n f o r m a t i o n than the measurements from 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 a l o n e .  The c o - o p e r a t i v e  t e s t i s c r i t i c a l l y e v a l u a t e d and recommendations provenance s t u d i e s a r e i n c l u d e d .  f o r future  I t i s f e l t t h a t many more  t e s t s w i t h D o u g l a s - f i r a r e n e c e s s a r y t o answer the q u e s t i o n s r a i s e d by t h i s and by other s t u d i e s .  LITERATURE CITED  A l l e n , G.S. 1960a. A method of d i s t i n g u i s h i n g c o a s t a l from i n t e r i o r Douglas f i r seed. B r i t i s h Columbia Lumberman 4-4-» 8 1 2 6 - 3 0 . . 1960b. F a c t o r s a f f e c t i n g the v i a b i l i t y and germi n a t i o n of c o n i f e r o u s seed. IV. S t r a t i f i c a t i o n p e r i o d and i n c u b a t i o n temperature. Pseudotsuga m e n z i e s l i (Mlrb.) Franco. The F o r e s t r y C h r o n i c l e 3 6 i l 8 - 2 9 . 196l. T e s t i n g Douglas f i r seed f o r provenance. Proc. I n t . Seed T e s t i n g A s s o c . 2 6 * 3 * 3 8 8 - 4 0 3 . Anon.  1955a. A c o o p e r a t i v e seed provenance study f o r P a c i f i c Northwest Douglas f i r . Mimeo. 8 pp. 1955b. Provenance s t u d y . p e c t u s . Mimeo. 4 pp.  Cone c o l l e c t i o n  pros-  Baldwin, H.I., and D.A. Rock. 1 9 6 1 . Douglas f i r as a New Hampshire Christmas t r e e . Fox F o r . note No. 80, 2 pp. Barner, H. 1966. C i r c u l a r l e t t e r of A p r i l 2 0 , 1 9 6 6 , of working group on procurement of seed f o r provenance research. I.U.F.R.O. S e c t i o n 2 2 , 10 pp. Bingham, R.T. 1966. Seed movement i n the Douglas f i r r e g i o n (Symposium summary and recommendations). Proceedings 1965 Western F o r e s t Genetics A s s o c i a t i o n . Pacific Northwest F o r e s t and Range Experiment S t a t i o n , pp. 30-34. B r a m h a l l , G. 1966. P e r m e a b i l i t y of D o u g l a s - f i r heartwood from v a r i o u s a r e a s of growth i n B.C. B r i t i s h Columbia Lumberman 5 0 « 4 1 9 8 , 1 0 0 , 1 0 2 . B r o d o v i c h , T.M. 1967. P r o p a g a t i o n of Pseudotsuga by g r a f t i n g i n the U k r a i n i a n S o v i e t S o c i a l i s t R e p u b l i c . Transl a t e d by Jan Freyman. F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8, B.C. T r a n s l a t i o n No. 5 5 . 7 PP. Byrnes, W.R.; H.D. Gerhold, and W.C. Bramble. 1958. Douglas f i r v a r i e t a l t e s t s f o r Christmas t r e e p l a n t a t i o n s i n P e n n s y l v a n i a . Prog. Rep. Pa. A g r i c . Exp. S t a . No. 1 9 8 , 6 pp. Chapman, J.D. 1952. The c l i m a t e of B r i t i s h Columbia. T r a n s . 5 t h B r i t i s h Columbia N a t u r a l Resource Conference, pp. 8 - 5 ^ .  Ching, Kim K. 1958. F i r s t y e a r n u r s e r y r e s u l t s of the coo p e r a t i v e seed provenance study f o r P a c i f i c Northwest Douglas f i r . Progress Report. Mimeo. 17 pp. . i960. Provenance study of Douglas f i r i n the P a c i f i c Northwest. Oregon F o r e s t Research Center, Corvallls. P r o g r e s s Report 2. Mimeo. 4 pp. . 1965» E a r l y growth of Douglas f i r i n a r e c i p r o c a l p l a n t i n g . F o r e s t Management Research. F o r e s t Research L a b o r a t o r y . School of F o r e s t r y , Oregon S t a t e U n i v e r s i t y , C o r v a l l l s , Oregon. Research paper 3 . 20 pp. . 1967» Provenance study of Douglas f i r i n the P a c i f i c Northwest Region I I . F i e l d performance a t age n i n e . Progress Report. F o r e s t Research Labora t o r y . School of F o r e s t r y , Oregon S t a t e U n i v e r s i t y , C o r v a l l l s , Oregon. 7 PP« .  1969*  Personal  communication.  .  1970.  Personal  communication.  , and Dale Bever. i 9 6 0 . Provenance study of Douglas f i r i n the P a c i f i c Northwest r e g i o n I . Nursery performance. S i l v a e G e n e t l c a 9«li11-17. C l e s l a r , A. 1 9 0 7 . Die Bedeutung k l i m a t i s c h e r V a r i e t a e t e n u n s e r e r H o l z a r t e n f u e r den Waldbau. Centralbl. gesamte Forstwesen 3 3 « 1 - 1 9 . 49-62. Decker, M a r c e l . 1 9 6 7 * Typewritten.  Die D o u g l a s i e i n Luxemburg.  130  pp.  Douglass, Bernard S. 1 9 6 7 . Christmas t r e e g e n e t i c improvement i n the P a c i f i c Northwest. American Christmas t r e e J o u r n a l , August, pp. 3-8. Edwards, M.V. 1956. The d e s i g n , l a y o u t and c o n t r o l of provenance experiments. Z e l t s c h r i f t f u e r F o r s t g e n e t i k und F o r s t p f l a n z e n z u e c h t u n g 5 » 5 - 6 i l 6 9 - l 8 0 . E n g l e r , A. 1913. E i n f l u s s der Provenlenz des Samens auf d i e E l g e n s c h a f t e n der f o r s t l i c h e n Holzgewaechse. Mitt. Schweiz. C e n t r a l a n s t . F o r s t l . Versuchswesen 1 0 . 1 - 3 8 6 . F e r r e l l , W i l l i a m K., and E. Steve Woodland. 1 9 6 6 . Effects of seed o r i g i n on drought r e s i s t a n c e of Douglas f i r (Pseudotsuga m e n z l e s l l (Mirb.) F r a n c o ) . Ecology 4 7 » 3i499-503. F o w e l l s , H.A. 1965. S i l v i c s of f o r e s t t r e e s of the U n i t e d S t a t e s . U n i t e d S t a t e s Department of A g r i c u l t u r e , F o r e s t S e r v i c e . A g r i c u l t u r a l Handbook 2 7 1 , 762 pp.  Galoux, A. 1 9 5 6 . Le s a p i n de Douglas e t l a phytogeographle. S t a t i o n de Recherches, Groenendaal. ( S e r i e B) No. 20. 131 PP. Gathy, P. 1 9 6 1 . Rapport p r e l l m i n a i r e s u r un t e s t d ' o r i g l n e s de Douglas v e r t . S t a t i o n de Recherches, Groenendaal. ( S e r i e B) No. 26. 44 pp. . 1967. Encore l e s o r i g i n e s du Douglas v e r t . B u l l e t i n Soc. F o r . B e l g . 74:7i353-358. G r i f f i t h , Braham G. i 9 6 0 . Growth of Douglas f i r a t the Unive r s i t y o f 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 s o i l . F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8, B.C. B u l l . No. 2. 58 pp. . 1968. Phenology, growth, and f l o w e r and cone p r o d u c t i o n of 154 Douglas f i r t r e e s on the U n i v e r s i t y Research F o r e s t as I n f l u e n c e d by c l i m a t e and f e r t i l izer, 1957-1967. F a c u l t y o f F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8, B.C. B u l l . No. 6. 70 pp. Haddock, P h i l i p p G. 1 9 6 6 . I n f o r m a t i o n a v a i l a b l e f o r other western s p e c i e s h a v i n g s u f f i c i e n t g e n e r a l i t y t o be a p p l i e d t o Douglas f i r seed movement problems. Western F o r e s t Genetics A s s o c i a t i o n , P r o c e e d i n g s . Olympia, Wash. Dec. 6-7, 1965. PP. 8-22. . 1967. The Importance of provenance i n f o r e s t r y . Proceedings I n t e r n a t i o n a l P l a n t Propagators S o c i e t y , Annual Meeting, Vancouver, B.C. pp. 91-98. , and Ralph Schmidt. 1957* V e g e t a t i o n and phytogeography of the Douglas f i r r e g i o n . Chapter V i n 1 An i n t r o d u c t i o n t o the f o r e s t s o i l s of the Douglas f i r r e g i o n of the P a c i f i c Northwest. F o r e s t S o i l s Comm i t t e e of the Douglas f i r r e g i o n . U n i v e r s i t y of Washi n g t o n , S e a t t l e . XIV Chapters. Chapter V, pp. 1-24. , and Oscar S z i k l a i . 1966. Seed c o l l e c t i o n zones f o r Douglas f i r i n Canada.. Proceedings of the S i x t h World F o r e s t r y Congress. Madrid. V o l . I I , pp. 1467-  1474.  ; John W a l t e r s , and A n t a l Kozak. 1967. Growth of C o a s t a l and I n t e r i o r Provenances of Douglas f i r (Pseudotsuga m e n z l e s l l (Mirb.) Franco) a t Vancouver and Haney i n B r i t i s h Columbia. F a c u l t y o f F o r e s t r y , Unive r s i t y of B r i t i s h Columbia, Vancouver 8, B.C. Research paper No. 79* 32 pp.  Haigh, R.W. 1 9 6 l . The e f f e c t o f provenance and growth r a t e on s p e c i f i c g r a v i t y and summerwood percentage o f young Douglas-fir. B.S.F. t h e s i s . F a c u l t y o f F o r e s t r y , U n i v e r s i t y o f B r i t i s h Columbia, Vancouver 8 , B.C. 57 PP. Heaman, J.C. 1 9 6 3 . A provenance study o f c o a s t a l D o u g l a s - f i r i n the P a c i f i c Northwest. A r e p o r t on t h e p r o j e c t i n B r i t i s h Columbia 1 9 5 ^ - 1 9 6 1 . B.C.R.F. t h e s i s # 4 l l . Confidential. P e r m i s s i o n was granted t o use m a t e r i a l f o r the present t h e s i s . 1968. E.P. 4 7 7 . 17-18.  A study o f phenotypes I n Douglas f i r . B.C. F o r e s t S e r v i c e Research Review, pp.  Hermann, R i c h a r d K., and Denis P. Lavender. 1 9 6 8 . E a r l y growth o f Douglas f i r from v a r i o u s a l t i t u d e s and a s p e c t s i n southern Oregon. S i l v a e Genetlea I 7 » 4 i 1^3-150. Hoffmann, K. 1 9 6 5 * Bedeutung des A u g u s t t r i e b e s f u e r den Fichtenanbau im P l e i s t o z a e n d e r DDR. S o z i a l . Forstw. I5i204-207. I r g e n s - M i l l e r , Helge. 1957. E c o t y p i c response t o temperature and p h o t o p e r i o d i n Douglas f i r . F o r e s t Science 3 * 1 « 79-83. . 1 9 6 5 . Of what v a l u e a r e c l i m a t i c d a t a i n a s s i g n i n g o f f - s o u r c e seed t o r e f o r e s t a t i o n s i t e ? P r o c . West. Forest Genetics A s s o c i a t i o n . Olympia, Wash., Dec. 6 - 7 . pp. 2 3 - 2 4 . 1967. P a t t e r n s o f h e i g h t growth i n i t i a t i o n and c e s s a t i o n i n Douglas f i r . S i l v a e Genetica l 6 i 2 t 5 6 - 5 8 . Isaac, Leo. 1949* B e t t e r Douglas f i r f o r e s t s from b e t t e r seed. U n i v . o f Wash. P r e s s , S e a t t l e . 64 pp. Jahn, G i s e l a . 1955* Gegenueberstellung d e u t s c h e r und amerik a n l s c h e r D o u g l a s i e n Provenienzen. A l l g . F o r s t und Jagdzeltung 1 2 6 I 4 I 6 8 - 7 6 . K r a j l n a , V l a d i m i r J . 1 9 5 6 . A summary o f the nomenclature o f D o u g l a s - f i r , Pseudotsuga m e n z l e s l l . Madrono 1 3 » 2 6 5 - 2 6 7 . . 1959* B i o c l l m a t i c zones i n B r i t i s h Columbia. B o t a n i c a l S e r i e s No. 1 . U n i v e r s i t y o f B r i t i s h Columbia, Vancouver 8 , B.C. 4 7 pp. Lacaze, J . F . 1 9 6 4 . Note s u r l a r e s i s t a n c e du Douglas s u i v a n t l ' o r l g i n e des g r a i n e s . Revue f o r e s t i e r e f r a n c a l s e l 6 t 3i225-7.  . 1968. Comparalson de quelques provenances de Douglas dans l'arboretum des B a r r e s . Revue f o r e s t l e r e franca!se 2 0 i l i 2 8 - 3 2 . L a n g l e t , 01of. 1936. S t u d l e r over t a l l e n s f y s i o l o g i s k a v a r l a b l l i t e t och dess samband med k l l m a t e t . Medd. f . Statens S k o g s f o r s o k s a n s t a l t 29.421-470. • 1945* Om m o j l i g h e t e r n a a t t skogsodla med gran och t a l l f r o av ortsframmande p r o v e n l e n s . Sartryck n r 1. Svenska Skogsvardsforeningens T i d s s k r l f t 431 68-78. . 1959* A c l l n e or not a c l l n e - a q u e s t i o n Scots p i n e . S i l v a e G e n e t i c a 811«13-22. Larsen,  C. Syraoh. 1956. Genetics l a t e d by Mark L. Anderson. burgh and London. 224 pp.  in silviculture. O l i v e r and Boyd.  of  TransEdin-  Leibundgut, Hans. 1962. Der Photoperiodismus a l s M i t t e l der Laerchenrassenforschung. Schwelz. Z e i t s c h r i f t f u e r Forstwesen 11316i332-333* L i n e s , Roger. 1967. (Compiler). S t a n d a r d i z a t i o n of methods f o r provenance r e s e a r c h and t e s t i n g . S e c t i o n 22. XIV. I.U.F.R.O. Congress, Munich, pp. 672-718. Little,  E l b e r t L., J r . 1953* Check l i s t of n a t i v e and n a t u r a l i z e d t r e e s of the U n i t e d S t a t e s ( i n c l u d i n g A l a s k a ) . A g r i c u l t u r a l Handbook No. 4 l . F o r e s t S e r v i c e , Washi n g t o n , D.C. 472 pp.  McKimmy, M.D. 1966. A v a r i a t i o n and h e r i t a b i l l t y study of wood s p e c i f i c g r a v i t y i n 46 y e a r o l d Douglas f i r from known seed s o u r c e s . Paper presented a t the t h i r d TAPPI F o r e s t B i o l o g y Conference, November 1-2, Madison, Wisconsin. 31 PP* (not a l l numbered). M i t c h e l l , R.G., and W.P. N a g e l . 1 9 6 9 . Tree s e l e c t i o n f o r c o n t r o l l i n g midges on D o u g l a s - f i r . American Christmas Tree J o u r n a l 1 3 I 4 I 1 1 - 1 3 . M o r r i s , W i l l i a m G.j R.R. S i l e n , and H. I r g e n s - M i l l e r . 1957. C o n s i s t e n c y of bud b u r s t i n g i n D o u g l a s - f i r . J o u r n a l of F o r e s t r y 55»3«208-210. Munger, Thornton R., and W i l l i a m G. M o r r i s . 1936. Growth of Douglas f i r t r e e s of known seed s o u r c e . Technical B u l l e t i n No. 537. U n i t e d S t a t e s Department of A g r i c u l t u r e , Washington, D.C. 40 pp.  Namkoong, Gene. 1 9 6 9 * N o n o p t i m a l i t y of l o c a l r a c e s . Proceedings of the Tenth Southern Conference on F o r e s t Tree Improvement. Houston, Texas, June 17-19. pp. 149-153. Nanson, Alphonse. 1 9 6 4 a . Les grandes l l g n e s de 1 • a m e l i o r a t i o n des essences f o r e s t l e r e s par v o l e g e n e t i q u e . B u l l . S o c i e t e Royale F o r e s t l e r e de B e l g i q u e , 7 l » 2 i 121-135. 1964b. Enquete s u r l a r e s i s t a n c e de d i v e r s e s provenances de Douglas v e r t a l ' h l v e r 1 9 6 2 - 6 3 en Belgique. B u l l . S o c i e t e Royale F o r e s t l e r e de Belgique 7 1 « 1 - 1 - 1 1 . • 1 9 6 5 . C o n t r i b u t i o n a l a v a l e u r des t e s t s p r e c o c e s . 1 . Experience I n t e r n a t i o n a l e s u r l ' o r i g i n e des g r a i n e s d ' e p i c e a ( 1 9 3 8 ) . Travaux. S t a t i o n de Recherches, Groenendaal. S e r i e E, No. 1 . 60 pp. Newnham, R.M. 1 9 6 8 . A c l a s s i f i c a t i o n o f c l i m a t e by p r i n c i p a l component a n a l y s i s and i t s r e l a t i o n s h i p t o t r e e species distribution. F o r e s t r y Science l 4 t 3 i 2 5 4 - 2 6 4 . N i c h o l s o n , J e f f e r s o n J.M. 1 9 6 3 * A provenance study of Douglas f i r grown under c o n t r o l l e d l i g h t and tempera t u r e c o n d i t i o n s . B.S.F. t h e s i s . F a c u l t y o f F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8 , B.C. 59 PP. Orr-Ewlng, A.L. 1 9 6 6 . I n t e r - and i n t r a s p e c i f i c c r o s s e s w i t h i n the genus Pseudotsuga. B.C. F o r e s t S e r v i c e Research Review, pp. 13-24. . 1967. A progeny t e s t of Douglas f i r t o demons t r a t e the importance of s e l e c t i o n i n f o r e s t p r a c t i c e . B.C. F o r e s t S e r v i c e Research Notes. No. 4 3 . Victoria. 23 PP. . 1 9 6 8 . Inbreeding s t u d i e s w i t h Douglas f i r , Cowichan Lake. E.P. 4 7 4 . B.C. F o r e s t S e r v i c e Research Review, p. 17. Pearse, P.H. 1 9 5 8 . A study on the e f f e c t s o f s o i l compaction on the e a r l y development o f s e e d l i n g s o f Douglas f i r and western hemlock. U n i v e r s i t y of B r i t i s h Columbia F o r e s t Club Research Notes. No. 1 6 . 7 pp. P h a r i s , R i c h a r d P., and W i l l i a m K. F e r r e l l . 1966. Differences i n drought r e s i s t a n c e between c o a s t a l and Inland sources of Douglas f i r . Canadian J o u r n a l of Botany 44il65l-l659.  Place, I . C M . 1969* Tree b r e e d i n g i n Canada. Chronicle 4 5 i 6 » 3 7 5 - 3 7 7 .  The Forestry-  R e v e l , W.J. i 9 6 0 . Provenance i n D o u g l a s - f i r . B.S.F. t h e s i s . F a c u l t y o f F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8 , B.C. 67 PP. Rohmeder, E . 1 9 5 6 . P r o f e s s o r Muench's Anbauversuch m i t Douglasien v e r s c h i e d e n e r Herkunft und anderen Nadelbaumarten im Forstamt K a i s e r s l a u t e r n - O s t 1912-195^. S i l v a e Genetica 5 » 5 / 6 « 1 4 2 - 1 5 6 . Rowe, J.S. 1959. F o r e s t Regions o f Canada. B u l l e t i n 1 2 3 . F o r e s t r y Branch. Canada Department o f Northern A f f a i r s and N a t u r a l Resources, Ottawa. 71 PP. Scheumann, Werner. 1 9 6 5 * M o e g l i c h k e i t e n und E r g e b n i s s e d e r F r o s t r e s i s t e n z p r u e f u n g i n d e r Douglasien- und L a e r chenzuechtung. Ini Moeglichkeiten der F o r s t p f l a n zenzuechtung. Tagungsberlchte Nr. 6 9 . pp. 1 8 9 - 1 9 9 . Schenck, C a r l A l w i n . 1939* Fremdlaendlsche Wald - und Parkbaeume. V o l . 2 . Die N a d e l h o e l z e r . B e r l i n , Paul Parey. Schmidt, Ralph L . 1 9 6 2 a . Provenance s t u d i e s . Unpubl. paper. Meeting of r e s e a r c h o f f i c e r s . V i c t o r i a , March 5 - 9 . B.C. F o r e s t S e r v i c e . Mimeo. 10 pp. . 1962b. The i n t e r p r e t a t i o n of c l i m a t e from pheno l o g l c a l assessments. Research D i v i s i o n B.C. F o r e s t Service. Mimeo. 8 pp. . 1 9 6 7 . B.C. F o r e s t S e r v i c e i n i t i a t e d Douglas f i r provenance study. B.C. Lumberman 5 1 » 3 » 7 2 - 7 4 . Schmidt-Vogt, H . 1 9 6 6 . Der J o h a n n i s t r i e b t e s t b e l F i c h t e , e i n F r u e h t e s t f u e r d i e Bestandesanerkenriung i n hoeheren Lagen. F o r s t p f l a n z e n - F o r s t s a m e n 4 J 2 1 - 2 2 . . 1 9 6 7 . Die Bedeutung der F r u e h t e s t e . M i t t e i l u n g e n des V e r e l n s f o r s t l . Standortskunde f u e r Waldbau und Forstpflanzenzuechtung, 1 7 » 8 6 - 9 1 . Schober, R. 195^. Douglasien Provenienzversuche I . melne F o r s t - und J a g d z e l t u n g I 2 5 i 5 i l 6 0 - 1 7 9 »  Allge-  . 1959. E r g e b n i s s e von Douglasien P r o v e n i e n z v e r suchen i n Deutschland, H o l l a n d und Daenemark. A l l g e melne F o r s t z e l t s c h r i f t 8 i l 4 5 - l 5 2 . . 1 9 6 3 . Experiences w i t h the Douglas f i r i n Europe. FAO World C o n s u l t a t i o n on F o r e s t G e n e t i c s . Stockholm. FA0/F0RGEN - 6 3 / - 4 / 5 pp. i i i - 1 8 .  , and H. Meyer. 195^. Douglasien Provenienzversuche I I . Allgemelne F o r s t - und J a g d z e l t u n g 1 2 5 I 5 I 160-179. 1955* D o u g l a s i e n Provenlenzversuche I I . A l l g e melne F o r s t - und J a g d z e l t u n g 1 2 6 t l l / l 2 i 2 2 1 - 2 4 3 . Schoenbach, Hans. 1958. Die Zuechtung d e r D o u g l a s l e . Goehrei Die Douglasle und i h r H o l z . Akademie Verlag, B e r l i n , pp. 309-367.  In  , and E r i c h Bellmann. 1967. F r o s t r e s i s t e n z der Nachkommenschaften von Kreuzungen "gruener" und " b l a u e r " Formen der Douglasle (Pseudotsuga m e n z l e s l l (Mirb.) F r a n c o ) . A r c h i v f u e r Forstwesen Band l6«6/9» 707-711. S i l e n , Roy R. 1962. A study of g e n e t i c c o n t r o l of bud b u r s t i n g i n Douglas f i r . J o u r n a l of F o r e s t r y 6o»7»472-475. . 1964. Regeneration a s p e c t s of the 50-year-old Douglas f i r h e r e d i t y s t u d y . Proceedings of the 1964 Annual Meeting of the Western R e f o r e s t a t i o n C o o r d i n a t i n g Committee of the Western F o r e s t r y and Conserv a t i o n A s s o c i a t i o n . P o r t l a n d , Oregon, pp. 35-39* 1966. A 50-year r a c i a l study of Douglas f i r i n western Oregon and Washington. (Abstract) Western F o r e s t G e n e t i c s A s s o c i a t i o n Proceedings 1965* Olympla, Washington, Dec. 6-7. pp. 6-7. Snyder, E.B. 1959. G l o s s a r y of f o r e s t t r e e improvement workers. American Soc. F o r e s t . 22 pp. Sorensen, Frank. 1967. Two-year r e s u l t s of a west-east t r a n s e c t provenance t e s t of Douglas f i r i n Oregon. U.S. F o r e s t S e r v i c e Research Note. P a c i f i c Northwest F o r e s t Range and Experiment S t a t i o n No. PNW72. 8 pp. Spurr, Stephen H. 1963. Growth of Douglas f i r i n New Zealand. T e c h n i c a l Paper No. 4 3 . F o r e s t Research I n s t i t u t e . New Zealand F o r e s t S e r v i c e . 54 pp. S q u i l l a c e , A.E., and R.T. Bingham. 1958. v a r i a t i o n i n western white p i n e . 20-34.  Localized ecotypic F o r e s t Science 4»lt  S t e m , K. 1966. Zur Herkunftsfrage der D o u g l a s l e . melne F o r s t z e i t s c h r l f t , 2 1 i 2 5 . 4 3 0 , 4 3 2 ,  Allge-  Sweet, G.B. 1965. Provenance d i f f e r e n c e s i n P a c i f i c Coast Douglas f i r seed and s e e d l i n g c h a r a c t e r i s t i c s . Silvae Genetlca l4i2«46-56.  Sziklal,  Oscar. 1 9 6 4 . P r e l i m i n a r y notes on v a r i a t i o n i n cone and seed morphology of D o u g l a s - f i r (Pseudotsuga menzle s l l (Mirb.) F r a n c o ) . Second World C o n s u l t a t i o n on F o r e s t Tree B r e e d i n g . Washington. 11 pp.  Tusko, F e r e n c . 1963* A study of v a r i a b i l i t y i n c e r t a i n D o u g l a s - f i r p o p u l a t i o n s i n B r i t i s h Columbia. Ph.D. thesis. Department of B i o l o g y and Botany, U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8, B.C. 173 PP. U.B.C. F o r e s t Committee. 1959* The f i r s t decade of management and r e s e a r c h . U.B.C. F o r e s t 1949-1958. Faculty of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8, B.C. 82 pp. U.S.  Department of A g r i c u l t u r e , United S t a t e s F o r e s t S e r v i c e . 1962. Annual Report f o r 1 9 6 l , P a c i f i c Northwest F o r e s t and Range Experiment S t a t i o n , U.S. Department of A g r i c u l t u r e , P o r t l a n d , Oregon. 105 pp. . 1964. The 1912 D o u g l a s - f i r h e r e d i t y study. Annual Report f o r 1 9 6 3 , P a c i f i c Northwest F o r e s t and Range Experiment S t a t i o n , U.S. Department of A g r i c u l t u r e , P o r t l a n d , Oregon, pp. 4 - 7 .  Vaartaja, 0 . 1959» Evidence of p h o t o p e r i o d i c t r e e s . E c o l . Monogr. 2 9 1 9 1 - 1 1 1 .  ecotypes i n  Wakeley, P.C. 1953. Working p l a n f o r the southwide pine seed source study. Mimeo. Walters, John. 1 9 6 1 . Annual Report of 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 , Haney, B.C. F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8 , B.C. pp. 9-11. . 1964. Annual Report of 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 , Haney, B.C. F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver B.C. p. 1 5 .  8,  . 1966. Annual Report of 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 , Haney, B.C. F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver B.C. p. 2 5 .  8,  . 1967. Annual Report of 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 , Haney, B.C. F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver B.C. p. 1 1 . . 1968. Annual Report of 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 , Haney, B.C. F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver B.C. p. 1 2 .  8,  8,  .  1970. P e r s o n a l  communication.  , and Kim K. Ching. 1969* P a t t e r n and bud b u r s t i n a D o u g l a s - f i r provenance study. Unpublished. , and J . Soos. 1 9 6 l a . The r e l a t i v e e f f i c i e n c y of three h a r e - r e p e l l e n t s i n p r o t e c t i n g D o u g l a s - f i r seedlings. F o r e s t r y C h r o n i c l e 37.22-28. . 196lb. Some o b s e r v a t i o n s on the r e l a t i o n s h i p of lammas shoots t o the form and growth of D o u g l a s - f i r seedlings. F a c u l t y o f F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8 , B.C. Research Paper 40. 8 pp. Wheat, Joseph G. 1 9 6 6 . The seed movement problem i n the Douglas f i r r e g i o n . Western F o r e s t Genetics A s s o c i ation. Proceedings, 1965* PP. 3-5» W i l l i n g t o n , Robert P. 1 9 6 8 . Some e f f e c t s o f s l a s h b u r n i n g , c l e a r c u t t i n g and skidroads on the p h y s i c a l - h y d r o l o g i c p r o p e r t i e s o f coarse g l a c i a l s o i l s In c o a s t a l B r i t i s h Columbia. M.F. t h e s i s . F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8 , B.C. 1^9 pp. Wright, Jonathan W. 1962. Genetics of f o r e s t t r e e improvement. F o r e s t r y and F o r e s t Products S t u d i e s No. 1 6 , FAO, Rome. 399 pp. , and H.I. Baldwin. 1957. The 1938 I n t e r n a t i o n a l Union Scots pine provenance t e s t i n New Hampshire. S i l v a e Genetica 6il/2«2-l4. Zon,  Raphael. 1913* Douglas f i r .  E f f e c t o f seed source upon the growth o f Forestry Quarterly 111^99-502.  S c i e n t i f i c and common names of s p e c i e s c i t e d In the t e x t  A b i e s a m a b i l l s (Dougl.)  Forbes  Pacific silver f i r  Acer c i r e i n a t u r n Pursh.  Vine maple  A r m l l l a r l a m e l l e a ( F r . ) Kumm.  A r m i l l a r i a root r o t fungus  Ghamaecyparls n o o t k a t e n s l s (D. • Spach C o n t a r l n l a pseudotsugae  Don)  Condrashoff  Yellow cedar D o u g l a s - f i r needle midge  E p l l o b l u m a n g u s t l f o l l u m L.  Fireweed  L a r l x decldua  European  Mill.  larch  P l c e a a b l e s (L.) K a r s t  Norway spruce  Plcea sltchensls  S i t k a spruce  (Bong.) C a r r .  Plnus montloola  Dougl.  Western white p i n e  Plnus ponderosa  Laws.  Ponderosa  Plnus s y l v e s t r l s  L.  Scots pine  Plnus taeda L. Pseudotsuga  L o b l o l l y pine  menzlesll  (Mlrb.) Franco  Rhabdocllne pseudotsugae Rubus s p e o t a b l l l s  pine  Syd.  Pursh.  Douglas-fir D o u g l a s - f i r needle b l i g h t fungus Salmonberry  Thuja p l l c a t a Donn.  Western r e d cedar  Tsuga h e t e r o p h y l l a (Raf.) Sarg.  Western hemlock  

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
http://iiif.library.ubc.ca/presentation/dsp.831.1-0075382/manifest

Comment

Related Items