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Geographic variation in seed weight, some cone scale measurements and seed germination of Douglas-fir… Yao, Chang 1971

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GEOGRAPHIC VARIATION IN SEED WEIGHT, SOME CONE SCALE MEASUREMENTS AND SEED GERMINATION OF DOUGLAS-FIR PSEUDOTSUGA MENZIESII (MIRB.) FRANCO by CHANG YAO B.Sc., Taiwan P r o v i n c i a l Chung-Hslng U n i v e r s i t y Taiwan, China, i 9 6 0 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF FORESTRY i n the Faculty of FORESTRY  We accept t h i s t h e s i s as conforming t o t h e r e q u i r e d standard  THE UNIVERSITY OF BRITISH COLUMBIA J u l y , 1971  In p r e s e n t i n g  t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f the  ments f o r an advanced degree a t the U n i v e r s i t y of  require-  British  Columbia, I agree t h a t the L i b r a r y s h a l l make I t f r e e l y a v a i l a b l e f o r reference  and  study.  I f u r t h e r agree t h a t p e r m i s s i o n f o r  e x t e n s i v e copying o f t h i s t h e s i s f o r s c h o l a r l y purposes may granted by the Head o f my I t i s understood t h a t  representarives.  copying o r p u b l i c a t i o n o f t h i s t h e s i s f o r  f i n a n c i a l g a i n s h a l l not  Department o f  Department o r by h i s  be  be a l l o w e d without my  pXi>Ht{ y[  The U n i v e r s i t y o f B r i t i s h Columbia Vancouver 8, Canada  written  permission.  ABSTRACT  The p r i n c i p a l o b j e c t i v e s o f t h e study were t o i n v e s t i g a t e geographic v a r i a t i o n o f , and r e l a t i o n s h i p between, 1000-seed weight and c o n e - s c a l e morphology and v a r i a t i o n o f g e r m i n a t i o n percent o f D o u g l a s - f i r ( Pseudotsuga m e n z l e s l l  (Mirb.) Franco )  from w i t h i n i t s n a t u r a l range i n Northwest America.  One hundred  twenty f o u r seed sources r e p r e s e n t i n g e i g h t c l i m a t i c r e g i o n s from B r i t i s h Columbia t o C a l i f o r n i a 117000' to 1 2 7 ° 2 7 ' )  (lat.  38°50'  to 5 3 ° 3 7 ' ,  long.  were c o l l e c t e d i n 1966 and 1968 by t h e I n t e r -  n a t i o n a l Union o f F o r e s t r y Research O r g a n i z a t i o n s , S e c t i o n 2 2 . From t h e seed samples, f i l l e d  seed (which c o n s t i t u t e d  seed weight) were s e l e c t e d u s i n g s o f t X-ray f l u o r o s c o p y .  1000Five  cone-scale measurements were taken; c o n e - s c a l e width, b r a c t width, cone-scale l e n g t h , 1 s t  prong l e n g t h and 2 n d prong l e n g t h .  The  p o s i t i o n o f t h e b r a c t i n r e l a t i o n t o t h e s c a l e was r a t e d . For  g e r m i n a t i o n t e s t i n g , $6 f i l l e d  seeds were s e l e c t e d t o  r e p r e s e n t each o f 12 t r e e s i n each o f l l 4 provenances. of  The t o t a l  7 6 , 6 0 8 seeds from 1 , 3 6 8 I n d i v i d u a l t r e e s were sown u n t r e a t e d  i n two r e p l i c a t i o n s on t e n r e l a t i v e l y u n i f o r m n u r s e r y beds d u r i n g May,  1969. Seed weights v a r i e d g r e a t l y .  One thousand-seed weight i n -  creased c l i n a l l y from low t o h i g h e l e v a t i o n and from n o r t h t o south.  L a t i t u d e appeared t o a f f e c t seed weights more than e l e v a -  tion. Cone-scale c h a r a c t e r i s t i c s d i f f e r e d s i g n i f i c a n t l y from t r e e to  t r e e , provenance t o provenance, as w e l l as s u b - r e g i o n t o sub-  region.  Cone-scale widths and l e n g t h s were o n l y s i g n i f i c a n t l y  I  d i f f e r e n t between r e g i o n s .  These c h a r a c t e r i s t i c s  a g a i n showed a  c l i n a l v a r i a t i o n which i n c r e a s e d from low t o h i g h e l e v a t i o n s and from n o r t h t o south i n some r e g i o n s , and r e v e a l e d t h a t affected  c o n e - s c a l e morphology more t h a n  latitude  elevation.  Thousand-seed weights were g e n e r a l l y p o s i t i v e l y  correlated  with c o n e - s c a l e s i z e . Germination percent was  significantly affected  by  latitude  around 36 days a f t e r sowing, but t h i s e f f e c t d i s a p p e a r e d by days.  50  E l e v a t i o n and l o n g i t u d e appeared not t o a f f e c t g e r m i n a t i o n  percent d u r i n g t h e observed p e r i o d  (0 - 92 days a f t e r  sowing).  The r e s u l t s o f t h i s study i l l u s t r a t e the importance o f geog r a p h i c o r i g i n as a source o f phenotypic v a r i a b i l i t y i n Douglasfir.  The  w r i t e r wishes t o express h i s thanks t o Dr. 0. S z i k l a i ,  P r o f e s s o r o f F o r e s t r y and Chairman o f t h e T h e s i s Committee, f o r s u p e r v i s i o n o f t h e experimental work and v a l u a b l e a s s i s t a n c e i n the p r e p a r a t i o n  o f t h e t h e s i s , a s w e l l as continuous encourage-  ment. Acknowledgement i s a l s o extended t o t h e other members o f my t h e s i s committeet Dr.  B. Van Der Kamp, F a c u l t y o f F o r e s t r y .  Dr.  G. W. Eaton, D i v i s i o n o f P l a n t  Science,  f o r c o n t r i b u t i n g t h e i r v a l u a b l e a d v i c e and f o r r e a d i n g t h e manuscript.  PAGE INTRODUCTION  1 ..3  LITERATURE REVIEW  .3  A.  Variation  B.  Variation  i n cone and seed  C.  Variation  i n seed g e r m i n a t i o n c h a r a c t e r i s t i c s  MATERIALS AND METHODS  characteristics..........3 .8 12  A.  Thousand-seed weight  12  B.  Cone-scale morphology  13  C.  R e l a t i o n s h i p between 1000-seed weight and cone-scale characteristics ... l 4  D.  Seed g e r m i n a t i o n t e s t  RESULTS AND DISCUSSION  l4 23 ...23  A.  Thousand-seed weight...  B.  Cone-scale morphology  C.  R e l a t i o n s h i p between 1000-seed weight and cone-scale characteristics 62  D.  Seed g e r m i n a t i o n t e s t  36  71  SUMMARY AND CONCLUSIONS  82  LITERATURE CITED.  85  LIST OF TABLES TABLE 1.  PAGE  L o c a t i o n o f 124  D o u g l a s - f i r provenances (No.1-91 were  c o l l e c t e d i n 1966, 2.  No.92-124 were c o l l e c t e d i n 1968)....19  Number o f provenances and t r e e s sampled In 1966  and  1968 3.  22  Maximum, minum and mean v a l u e s  o f 1000-seed weight f o r  each provenance In grams 4.  Average v a l u e s and  30  «...  standard  d e v i a t i o n s o f 1000-seed 33  weight by c l i m a t i c r e g i o n s i n grams 5.  R e l a t i o n s h i p between 1000-seed weight and  l o c a t i o n of 3^  seed source. 6.  R e l a t i o n s h i p between 1000-seed weight and l a t i t u d e 35  w i t h i n v a r y i n g range o f e l e v a t i o n 7.  Mean v a l u e s and  standard  d e v i a t i o n o f cone-scale  charac-  t e r i s t i c s by provenances i n m i l l i m e t e r s . . . . . . . . . . . . . . . . . 50 8.  Mean v a l u e s and  standard  d e v i a t i o n of cone-scale  charac-  t e r i s t i c s by c l i m a t i c r e g i o n s i n m i l l i m e t e r s 9/a.  R e l a t i o n s h i p between cone-scale  width and l o c a t i o n of 55  seed source 9/b. 10/a.  5^  A n a l y s i s o f v a r i a n c e of cone-scale  55  widths  R e l a t i o n s h i p between b r a c t width and l o c a t i o n of seed source  56  10/b.  A n a l y s i s o f v a r i a n c e o f b r a c t widths...  56  11/a.  R e l a t i o n s h i p between cone-scale  l e n g t h and  57  seed source 11/b.  A n a l y s i s of v a r i a n c e o f cone-scale  l o c a t i o n of  lengths  57  12/a.  R e l a t i o n s h i p between 1 s t prong l e n g t h and l o c a t i o n 58  o f seed source. 12/b.  A n a l y s i s o f v a r i a n c e o f 1 s t prong l e n g t h s  .58  13/a.  R e l a t i o n s h i p between 2 n d prong l e n g t h and l o c a t i o n .......59  o f seed source 13/b.  A n a l y s i s o f v a r i a n c e o f 2 n d prong l e n g t h s  lVa.  R e l a t i o n s h i p between r a t i n g o f b r a c t and l o c a t i o n  iVb. 15.  .....59  of seed source  60  Analysis of variance of r a t i n g of bracts.  60  Summary o f average cone-scale  characteristics f o r 6l  " c o a s t a l " and " i n t e r i o r " r e g i o n s 16.  C o r r e l a t i o n between 1000-seed weight and cone-scale .70  characteristics 17.  Germination percent  18.  Minimum, maximum and mean germination 50,  o f 114 D o u g l a s - f i r provenances.76 36,  72 and 92 days a f t e r sowing w i t h i n a c l i m a t i c 79  region 19.  percent  V a r i a t i o n o f germination  percent  from f o u r d i f f e r e n t  provenances throughout f o u r observed stages........81  LIST OP FIGURES FIGURE 1.  PAGE  Geographic d i s t r i b u t i o n o f seed sources i n t h e P a c i f i c Coast o f North America D o u g l a s - f i r  2.  View o f a b a x i a l and a d a x i a l provenances No.  3a.  3b. 4-6.  surface  study....16  o f c o n e - s c a l e from  1 , 1 1 , 1 2 and 124.  Diagrammatic r e p r e s e n t a t i o n and b r a c t  provenance  17  o f D o u g l a s - f i r cone-scale  showing f i v e b a s i c measurements  P o s i t i o n o f the bract  18  i n r e l a t i o n to the s c a l e . .  The r e l a t i o n s h i p between 1000-seed  ..18  weight and e l e v a t i o n  i n Regions 1 , 2 and 3 7-11.  The r e l a t i o n s h i p between 1000-seed  27 weight and l a t i t u d e  i n Regions 1 , 2 , 3 , 5 and 7 . . . . . 12a-12f.  The r e l a t i o n s h i p between 1000-seed  28 weight and l a t i t u d e  from south t o n o r t h f o r each 500 f e e t o f e l e v a t i o n  from  sea l e v e l t o 2 , 5 0 0 f e e t , and 2 , 5 0 1 t o 5 , 5 0 0 f e e t 13-14.  29  The r e l a t i o n s h i p between t h e width o f c o n e - s c a l e and e l e v a t i o n i n Regions 1 and 3  15.  4l  The r e l a t i o n s h i p between t h e width o f b r a c t and e l e v a t i o n i n Region 7  16-17.  4-1  The r e l a t i o n s h i p between t h e l e n g t h  o f c o n e - s c a l e and  e l e v a t i o n i n Regions 1 and 3 18.  i n Region 1 .  .....43 o f 2 n d prong and  The r e l a t i o n s h i p between the l e n g t h elevation  21-22.  o f 1 s t prong and  The r e l a t i o n s h i p between t h e l e n g t h elevation  19-20.  42  i n Regions 1 and 7  43  The r e l a t i o n s h i p between the r a t i n g o f b r a c t and e l e v a t i o n i n Regions 1 and 3 .  •  44  FIGURE 23-25.  PAGE  The r e l a t i o n s h i p between t h e width o f c o n e - s c a l e and l a t i t u d e i n Regions 1,3 and 5  26-28.  45  The r e l a t i o n s h i p between t h e l e n g t h o f c o n e - s c a l e and l a t i t u d e i n Regions 1,3 and 5 . .  29-32.  46  The r e l a t i o n s h i p between t h e l e n g t h o f 1st prong and l a t i t u d e i n Regions 1,2,3 and 5  33-36.  4?  The r e l a t i o n s h i p between t h e l e n g t h o f 2nd prong and l a t i t u d e i n Regions 1,2,3 and 5  37-38.  48  The r e l a t i o n s h i p between t h e r a t i n g o f b r a c t and l a t i t u d e i n Regions 1 and 3  39-42.  ^9  The r e l a t i o n s h i p between 1000-seed weight and width of cone-seale i n Regions 1,2,3 and 5 . . . . . . .  43-45.  65  The r e l a t i o n s h i p between 1000-seed weight and l e n g t h of c o n e - s c a l e i n Regions 1,3 and 5  46-49.  ....66  The r e l a t i o n s h i p between 1000-seed weight and l e n g t h of 1st prong i n Regions 1,2,3 and 5  50-53.  ....67  The r e l a t i o n s h i p between 1000-seed weight and l e n g t h of 2nd prong i n Regions 1,2,3 and 5  54-55.  68  The r e l a t i o n s h i p between 1000-seed weight and r a t i n g of b r a c t i n Regions 1 and 3  56.  69  The c o r r e l a t i o n between average g e r m i n a t i o n percent ( a f t e r 36 days) and l a t i t u d e i n D o u g l a s - f i r  57-60.  7^  Cumulative g e r m i n a t i o n percent o f 12 t r e e s w i t h i n provenance  (Provenance N o . l , Stoner, B.C., Provenance  No.122, Covelo, C a l i f o r n i a , Provenance No.12,  Jeune  Landing, B.C., and Provenance No.11, Golden, B.C.)...75  D o u g l a s - f i r , Pseudotsuga m e n z l e s l l  (Mirb.) Franco, not  only  i s a w i d e l y - d i s t r i b u t e d but a l s o an e c o n o m i c a l l y important f o r e s t tree species  i n western North America.  The  n a t u r a l range of  D o u g l a s - f i r extends more than 2 , 0 0 0 m i l e s from n o r t h e r n c e n t r a l B r i t i s h Columbia i n t o Mexico and Ocean t o the e a s t e r n  almost 1 , 0 0 0  m i l e s from P a c i f i c  s l o p e s o f Rocky Mountains.  under more d i v e r s e c l i m a t i c c o n d i t i o n s than any North American commercial t r e e s p e c i e s . graphic  s p e c i e s grows  o t h e r Important  However, a l t h o u g h geo-  r a c e s e x i s t over t h i s l a r g e a r e a , many o f them a r e  s u i t a b l e f o r commercial use The  The  un-  (Allen, 1961).  phenotypic v a r i a b i l i t y w i t h i n the s p e c i e s through i t s  geographic range has  been i n v e s t i g a t e d by many workers and  l a r g e p o r t i o n o f t h i s v a r i a b i l i t y was environment  (Irgens-Moller,  1958  and  a t t r i b u t e d t o the  local  1 9 6 2 i Bramhall, 1 9 6 6 ) .  i s s t i l l p o s s i b l e t o study the v a r i a b i l i t y wherever the grows, and  a  It  species  t h e r e a r e advantages i n doing so i n n a t u r a l f o r e s t s .  Cones as t h e g e n e r a t i v e  organ o f gymnosperms a r e l e a s t a f f e c t e d  by e x t e r n a l c o n d i t i o n s and  have c h a r a c t e r s  which w i l l permit  r e c o g n i t i o n o f lower taxonomlc u n i t s w i t h i n s p e c i e s .  the  Cone and  seed c h a r a c t e r i s t i c s a r e l e s s i n f l u e n c e d by environment and most revealing i n v a r i a t i o n studies t i o n i n cone morphology and  (Sziklai, 1964).  Although v a r i a -  seed weight as w e l l as the r e l a t i o n -  s h i p s between these c h a r a c t e r i s t i c s i n c o n i f e r s have been s t u d i e d by a number o f i n v e s t i g a t o r s ( S q u i l l a e e , 19571 1967» how  Sweet, 1965  and  Sziklai, I969),  Simak, i 9 6 0  and  these s t u d i e s d i d not show  cone-scale morphology i n f l u e n c e d seed weight i n D o u g l a s - f i r .  The  primary alms of t h i s study a r e d i v i d e d i n t o f o u r p a r t s ; P a r t  A - t o i n v e s t i g a t e the geographic v a r i a t i o n o f 1000-seed weight; P a r t B - cone-scale  morphology (cone-scale width,  l e n g t h , b r a c t width, 1st prong l e n g t h , 2nd  cone-scale  prong l e n g t h  r a t i n g of b r a c t ) ; P a r t C - r e l a t i o n s h i p s between these  and character-  i s t i c s f o r a wide range of D o u g l a s - f i r provenances; and P a r t D t o determine the range i n seed germination g r a p h i c f a c t o r s i n f l u e n c i n g germination  percent  and  the geo-  i n Douglas-fir.  Because of the fewer number of samples from Regions 4, 6 8  (4, 2 and  1 provenance r e s p e c t i v e l y ) , these r e g i o n s were not  i n c l u d e d i n subsequent r e g r e s s i o n The was  germination  investigated.  germination  and  analyses.  i n a r e l a t i v e l y uniform  nursery  environment  In p a r t i c u l a r , the r e l a t i o n s h i p s between seed  percentage and l a t i t u d e , l o n g i t u d e , and  the seed source were s t u d i e d .  e l e v a t i o n of  LITERATURE REVIEW  A..  Variation V a r i a t i o n w i t h i n D o u g l a s - f i r was  noted by L a r s e n i n  1937  as f o l l o w s J "One has t o t r a v e l v e r y w i d e l y throughout the n a t u r a l range o f D o u g l a s - f i r i n o r d e r t o get an i m p r e s s i o n o f d i f f e r e n c e s i n g e o g r a p h i c a l type, but s t a n d i n g on 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 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 o r i n the S i e r r a Nevada, passes through Oregon and Washington, o r 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 o r i n the Rocky Mountains} everywhere one i s bound t o be Impressed 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 tree-species." Stebbins  ( 1 9 5 7 ) s t a t e d as  followsi  "One advantage i s t h a t i t makes p o s s i b l e the a n a l y s i s of the i n d i v i d u a l c h a r a c t e r s o f t h e s e combinations and i s the f i r s t step toward the c a u s a l a n a l y s i s o f t h e s e d i f f e r e n c e s In terms o f s e l e c t i o n o r any o t h e r factors. I t a l s o f o c u s e s a t t e n t i o n on the continuous v a r i a t i o n i n q u a n t i t a t i v e c h a r a c t e r i s t i c s which i s present i n many wide-ranging s p e c i e s and i s o f g r e a t importance i n t h e i r a d a p t a t i o n t o t h e environment..." Sziklai  (1967)  emphasized the Importance of v a r i a t i o n i n the  f o l l o w i n g way* "The v a r i a t i o n p a t t e r n from t r e e t o t r e e and from stand t o stand, as w e l l as the p o p u l a t i o n composit i o n throughout the range o f the s p e c i e s , should be known b e f o r e any I n t e n s i v e f o r e s t r y work can be planned on a l o g i c a l b a s i s . "  B.  V a r i a t i o n In cone and W i l l i s and  seed c h a r a c t e r i s t i c s  Hoffmann ( 1 9 1 5 ) observed t h a t i n D o u g l a s - f i r  the s i z e of cone was  d i r e c t l y dependent upon the v i g o r of  the  cone-bearing shoot. Coover ( 1 9 3 3 ) r e p o r t e d t h a t seeds from the upper  P e r r y and  crowns o f s h o r t l e a f pine  (Plnus eohlnata  M i l l . ) and  p i t c h pine  (Plnus r l g l d a M i l l . ) were more v i a b l e than those from middle and lower crowns.  They a l s o found t h a t l a r g e r cones g e n e r a l l y  l a r g e r seeds i n p i t c h p i n e , but t h a t many s m a l l and cones contained  yield  medium-size  more and b e t t e r seeds than the l a r g e r cones i n  shortleaf pine.  They noted t h a t p i t c h p i n e cones v a r i e d i n s i z e  from t r e e t o t r e e with l i t t l e v a r i a t i o n w i t h i n t r e e s and  found  no a s s o c i a t i o n between cone s i z e and v e r t i c a l p o s i t i o n o f  the  cone i n the t r e e .  pine  Wright ( 1 9 4 5 ) found t h a t the f r e s h weight of E a s t e r n  white  (Plnus  small  strobus L.)  t o l a r g e cones and  seed i n c r e a s e d  from the apex t o the base o f the  cone.  G u s t a f s s o n (195*0 noted t h a t i n S c o t s pine  Simak and  s l l v e s t r l s L . ) , cone s i z e and seed p r o d u c t i o n ment and,  s i g n i f i c a n t l y from  cone weight d i d not  only  (Plnus  influence  and average seed weight but a l s o embryo develop-  through t h i s , the subsequent g e r m i n a t i o n  Seed weight per cone i n c r e a s e d  capacity  with r i s i n g cone weight and  de-  c r e a s i n g seed number i n the mother t r e e s but these c o r r e l a t i o n s c o u l d not be e s t a b l i s h e d i n the g r a f t s .  In m o r p h o l o g i c a l r e -  spects t h e r e a r e d i s t i n c t d i f f e r e n c e s between seed obtained n a t u r a l t r e e s and  from g r a f t e d i n d i v i d u a l s .  Slmak ( i 9 6 0 ) r e p o r t e d from two  on cone samples t h a t were c o l l e c t e d  t r e e s o f S c o t s p i n e c l o s e t o each other i n a stand  Bogesund, Sweden, i n d i f f e r e n t years - 1 9 5 2 / 5 3 , and  1955/56.  from  Through h i s I n v e s t i g a t i o n , i t was  1953/5*+,  at  195V55  found t h a t number  o f seeds p e r cone, and average s i z e o f seed i n c r e a s e d with i n c r e a s i n g cone weight.  The frequency o f empty seeds decreased  when weight i n c r e a s e d .  He concluded t h a t t h e r e l a t i o n s h i p s o f  these p r o p e r t i e s appeared t o be determined by t h e genotype o f the t r e e and t h a t they were s t r o n g l y m o d i f i e d by t h e y e a r l y climatic variation. Allen  (1961) found t h a t v a r i a t i o n w i t h i n a D o u g l a s - f i r cone  was random and o f about t h e same o r d e r as t h e v a r i a t i o n among the cones o f t h e same p a r e n t .  The g e n e t i c i m p l i c a t i o n s were  evident i n t h e case o f t h e s i n g l e cone, t h e s i n g l e t r e e and t h e v a r i a t i o n among t r e e s , and t h e s e were a t t r i b u t e d t o environment, both i n t h e cone and t h e t r e e as w e l l as f o r a l o c a l  population  of trees. Peace  (1948) s t u d i e d the n o r t h e r n p a r t o f t h e D o u g l a s - f i r  range and noted t h e l a r g e v a r i a t i o n i n cone within the species.  characteristics  He found t h e range i n cone l e n g t h was from  3.4 t o 8.4 c e n t i m e t e r s .  Cones from t h e coast tended t o occupy  the upper end o f t h e range.  The r e f l e x e d b r a c t  characteristics  were observed on t h e coast w h i l e , a l t h o u g h t h e r e v e r s e s i t u a t i o n had i n t h e past been c o n s i d e r e d t o p r e v a i l , cones "without r e f l e x e d b r a c t s " were found i n t h e Rocky Mountain a r e a . Willett  (1963) measured t h e D o u g l a s - f i r cone l e n g t h and  width from 22 coast and 8 I n t e r i o r provenances from Nlmpkish Lake (Vancouver I s l a n d ) t o Kananaskis ( A l b e r t a ) on 348 t r e e s . The average cone l e n g t h f o r provenances was 6.0  cm w i t h a range  from 5»1 t o 7«7 cm and cone width average 2.1 w i t h a minimum o f 1.8  cm and a maximum o f 2.4 cm f o r t h e d i f f e r e n t  provenances.  L o n g i t u d e , l a t i t u d e o f t h e c o l l e c t i o n a r e a , t h e h e i g h t , diameter a t b r e a s t h e i g h t , crown width and age o f t h e t r e e e x p l a i n e d o n l y 9 . 3 p e r cent f o r cone l e n g t h and 1 3 . 2 p e r cent from cone width of the v a r i a t i o n .  T h i s suggests that o t h e r environmental and  probably g e n e t i c a l v a r i a b l e s a r e a l s o important. Tusko  (1963)  i n v e s t i g a t e d D o u g l a s - f i r cone samples  from  4 3 provenances a c r o s s B r i t i s h Columbia from east t o west and found t h e average cone l e n g t h t o be 5 . 4 cm r a n g i n g from 3 . 2 t o 9.3  cm.  The average cone width was 2 . 1  1 . 6 and 2 . 7  cm.  cm w i t h a range between  A c e r t a i n o v e r l a p was observed i n t h e s e c h a r a c -  t e r i s t i c s so f a r as coast and i n t e r i o r o r i g i n s were concerned but g e n e r a l l y t h e coast provenances were l a r g e r . Robinson ( 1 9 6 3 ) i n v e s t i g a t e d t h e v a r i a t i o n i n s i z e o f seed o f D o u g l a s - f i r based on 348 t r e e s from 30 d i f f e r e n t  provenances  and 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 both the l e n g t h and width o f seed and wing, among t h e d i f f e r e n t provenances.  He a l s o  mentioned t h a t o n l y 2 1 . 6 percent o f t h e v a r i a t i o n i n seed weight can be e x p l a i n e d by t h e age o f mother t r e e , l a t i t u d e and l e n g t h o f seed. Sziklai  ( 1 9 6 7 ) I n v e s t i g a t e d D o u g l a s - f i r cone l e n g t h , l e n g t h  o f seed, width o f seed, l e n g t h o f wing and width o f wing from 91 provenances  ( l a t i t u d e range from 5 3 ° 3 7 '  range from 1 2 1 ° 2 7 '  to 1 7 7 ° 0 0 ' ) .  t o 44°24*j longitude  The average cone l e n g t h o f  c l i m a t i c s u b - r e g i o n was a range from 4 9 . 6 0 t o 6 6 . 8 8 mm.  The  cone l e n g t h f o r t h e " c o a s t a l " r e g i o n s appeared t o be l o n g e r (62.64 mm) than f o r " i n t e r i o r " r e g i o n s ( 5 5 . 4 3 mm).  The l e n g t h  o f seed was 6 . 6 9 mm f o r " c o a s t a l " and 6 . 4 9 mm f o r " i n t e r i o r " ;  width o f seed 3 . 9 1  the 7.98  and 3.80 mms  the width o f wing 5 ' 8 6  mm:  the l e n g t h o f wing 9 . 1 5  and 6 . 2 2  mm.  and  He p o i n t e d out  t h a t a c l e a r l y - e x p r e s s e d c l i n a l v a r i a t i o n was  observed i n cone  and seed l e n g t h with 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 i n v e s t i g a t e d such as l e n g t h o f wing and seed d i d not show a s i m i l a r c l i n a l v a r i a t i o n p a t t e r n . Roche ( 1 9 6 6 ) s t u d i e d geographic v a r i a t i o n o f cone morphology which he found t o be s t r o n g l y c l i n a l i n white spruce  .  (Plcea  g l a u c a (Moench) Voss) and Engelmann spruce ( P l c e a engelmannll Parry).  The p o p u l a t i o n s were continuous i n t h e i r  from Montane t o S u b - a l p i n e f o r e s t r e g i o n s .  distribution  Hybrid populations  between both t h e s e s p e c i e s e x i s t and have been r e c o g n i z e d and d e l i m i t e d on the b a s i s o f cone s c a l e morphology. Van Deusen and Beagle ( 1 9 7 0 ) r e p o r t e d t h a t ponderosa p i n e (Plnus ponderosa Laws.) cone samples 1968  from 75  c o l l e c t e d d u r i n g 1967  i n d i v i d u a l t r e e s i n the Bear Lodge Mountain  Wyoming, were q u i t e uniform i n l e n g t h , a v e r a g i n g 2 . 6 the  area.  from 8 , 2 ^ 7 cone was of  and of  Inches over  Number o f seeds per pound averaged 1 2 . 6 7 3 but ranged to 22,997 for individual trees.  Number o f seeds per  p o s i t i v e l y r e l a t e d t o cone l e n g t h .  There was an average  4 l 5 green cones p e r b u s h e l . Sweet ( I 9 6 5 )  i n New  Zealand examined 30 D o u g l a s - f i r prove-  nances from the west s i d e o f the Cascades and Nevada.  Samples  were c o l l e c t e d from l a t i t u d e s r a n g i n g from 3 8 ° 1 0 ' N t o 48 0 15*N i n 1956.  Another two were c o l l e c t e d from p l a n t a t i o n s i n New  Zealand i n 1955  and 1 9 5 6 .  He found h i g h l y s i g n i f i c a n t  ences between seed weight and:  (1)  altitude,  (2)  differ-  length of  f r o s t - f r e e growing  season and  month o f seed s o u r c e s .  (3) mean temperature  He noted the c o r r e l a t i o n betwesen seed  weight and a l t i t u d e o f seed source was  c o n s i d e r a b l y h i g h e r than  between seed weight and f e a t u r e s r e p r e s e n t i v e o f regime a t seed s o u r c e .  of coldest  Provenances  temperature  from h i g h e r a l t i t u d e s had  h e a v i e r seeds than those from lower a l t i t u d e s . Allen  ( i 9 6 0 and 1961)  had developed a method f o r d i s t i n -  g u i s h i n g between coast and i n t e r i o r o r i g i n s of D o u g l a s - f i r based on seed morphology.  G.  V a r i a t i o n i n seed g e r m i n a t i o n c h a r a c t e r i s t i c s The g e r m i n a t i o n of seed Is i n f l u e n c e d by many complex  factors.  F a c t o r s r e l a t i n g t o provenance,  including  photo-  p e r i o d i c requirements and f l o w e r i n g h a b i t s , the n a t u r e of p o l l i n a t i o n and f e r t i l i z a t i o n , t h e s i z e , weight and l o n g e v i t y o f the seed, t h e degree o f m a t u r i t y , the c h a r a c t e r i s t i c s of dormancy, the p o s i t i o n of the seeds i n the cone and the p o s i t i o n of t h e cone on the t r e e s , and t h e c h a r a c t e r i s t i c s t o secondary dormancy under u n f a v o u r a b l e e x t e r n a l c o n d i t i o n s a r e i n h e r e n t i n the seed itself  (Baldwin,. 1 9 ^ 2 ) .  Mirov  ( 1 9 3 6 ) p o i n t e d out t h a t the g e r m i n a t i o n t e s t was  used  t o determine the v a r i a b i l i t y o f seed, t o estimate the amount of seed t o be used i n the f i e l d , o r t o determine t h e requirements f o r optimum g e r m i n a t i o n under v a r i o u s environments. The time o f cones h a r v e s t i n g , and the techniques o f seed e x t r a c t i o n and s t o r a g e a f f e c t g e r m i n a t i o n .  Rohmeder ( 1 9 ^ 2 )  found t h a t the g e r m i n a t i o n c a p a c i t y of f u l l y matured Ulmus  montana (With.) seed was h i g h e s t a t t h e time o f h a r v e s t and g r a d u a l l y decreased t h e r e a f t e r .  Seed which was not f u l l y matured  was c h a r a c t e r i z e d by a g e r m i n a t i o n c a p a c i t y h i g h a t f i r s t ,  low  d u r i n g one o r s e v e r a l months o f a f t e r - r i p e n i n g , and a t i t s h i g h est  s h o r t l y a f t e r t h i s was completed.  He a l s o mentioned  that  prematurely h a r v e s t e d seed s t o r e d b e t t e r than f u l l y matured seed and i t was recommended t h a t Ulmus montana (With.) seed should be h a r v e s t e d l a t e i n t h e season, p r e f e r a b l y by c o l l e c t i o n from the ground.  Sowing i n f o r e s t o r n u r s e r y should take p l a c e as soon  as p o s s i b l e a f t e r seed h a r v e s t . Hebb (195*0 r e p o r t e d t h a t t h e most e f f e c t i v e way t o open pond p i n e (Plnus s e r o t l n a Mlchx.) cones was t o d i p t h e cones i n b o i l i n g water f o r a moment.  He found»  (1)  full  seed from q u i c k -  s c a l d e d cones germinated 9 6 . 9 p e r c e n t , the h i g h e s t g e r m i n a t i o n of any cone treatment; and  (3)  (2)  seed from baked cones 94.4  percent,  seed from a i r - d r i e d cones opened with a k n i f e 9 2 . ? p e r -  cent. Allen fir  (1957) a l s o found t h a t seed i n green uncured  Douglas-  cones showed v e r y heavy l o s s e s when s u b j e c t e d t o 104°F, where-  as seed In p r e - c u r e d s i m i l a r cones showed no i l l e f f e c t a t a 122°F k i l n temperature.  He a l s o s t a t e d t h a t D o u g l a s - f i r can be  s a f e l y d r i e d a t a k i l n temperature  o f up t o 122°F, but about  20  percent o r more l o s s i n D o u g l a s - f i r seed v i a b i l i t y when t h e k i l n temperature  was r a i s e d t o l4o°F.  The seed damaged by dewinglng  or having a d u l l , d u s t y - l o o k i n g seed coat produced s e e d l i n g s o f low v i g o u r a p p a r e n t l y s u s c e p t i b l e d e s t r u c t i v e contamination. T o o l et a_l, (1956) i n d i c a t e d t h a t t h e g r e a t v a r i a b i l i t y  of  temperature age,  requirements  between and w i t h i n s p e c i e s depended on  storage c o n d i t i o n s and other f a c t o r s . Stone (1957  ) r e p o r t e d t h a t s u g a r p i n e (Plnus l a m b e r t l a n a  Douglas) seeds s t o r e d a t 77°F and 0°F, were d r i e d a t room temperature 10 p e r c e n t .  i n d e s i c c a t o r s a f t e r seeds  t o an average moisture content of  A f t e r two-year storage a t the U n i v e r s i t y o f C a l i -  f o r n i a A g r i c u l t u r e Experiment S t a t i o n , seeds were g i v e n a  germi-  n a t i o n t e s t i n P e t r i d i s h e s f i l l e d with v e r m l c u l i t e , and i t was found t h a t the dry storage a t 0°F was more e f f e c t i v e i n maint a i n i n g the seeds " f r e s h " c o n d i t i o n than storage a t 36°F o r 77°F. Mirov  ( 1 9 4 6 ) r e p o r t e d t h a t germination o f seed from  21  s p e c i e s o f p i n e kept a t C a l i f o r n i a F o r e s t and Range Experiment S t a t i o n i n a i r t i g h t 5 - g a l l o n t i n cans a t 40°F f o r p e r i o d s ranging from 5 t o 15 y e a r s , showed t h a t seed o f some p i n e s w i l l keep f o r a l o n g time without l o s i n g t h e i r v i a b i l i t y . With r e g a r d t o the e f f e c t of seed s i z e on Wright  germination,  ( 1 9 ^ 5 ) r e p o r t e d t h a t medium-size seeds showed h i g h e r  germination percentage  than e i t h e r the l a r g e o r s m a l l seeds o f  e a s t e r n white p i n e i n both s t r a t i f i e d Baldwin  or u n s t r a t l f i e d .  (19*1-2) reviewed the work o f some i n v e s t i g a t o r s and  found t h a t the s i z e and weight of seed had a d e f i n i t e e f f e c t  on  g e r m i n a t i o n because the l a r g e s t and h e a v i e s t seeds were the b e s t , had the more food r e s e r v e s , germinated t h e most v i g o r o u s s e e d l i n g s .  more promptly,  On the c o n t r a r y , I l j i n  and produced (1952) r e -  ported on t e s t s o f S c o t s p i n e i n a Liebenberg germinator but ed t o e s t a b l i s h a r e l a t i o n s h i p between seed weight and c a p a c i t y and  energy.  fail-  germination  Tourney and K o r s t l a n ( 1 9 ^ 8 ) s t a t e d t h a t when t h e s i z e of seed was  not dependent upon the range o f geographic  distribution  but r a t h e r upon l o c a l c o n d i t i o n s , l a r g e r seed possesses a g r e a t e r g e r m i n a t i n g power and produces  more v i g o r o u s s e e d l i n g s .  Seed g e r m i n a t i o n i s not o n l y e f f e c t e d by seed s i z e , but a l s o by cone s i z e . s i z e groupsJ 1 . 5 - 2 . 5 cms  Kocharj (1)  and  t e s t made on 100  >4.5 (3)  ( 1 9 5 0 ) d i v i d e d S c o t s p i n e cones i n t o t h r e e cm l o n g x  < 3 x  < 1.5  >1.5 cm.  cm diameters  (2)  3-^*5  x  The l a b o r a t o r y g e r m i n a t i o n  seeds of each group showed t h a t group 2 had  h i g h e s t g e r m i n a t i o n c a p a c i t y and energy.  the  Group 1 showed p r a c t i -  c a l l y the same g e r m i n a t i o n c a p a c i t y but a c o n s i d e r a b l y lower energy  of g e r m i n a t i o n and group 3 was  v e r y much i n f e r i o r .  S t r a t i f i c a t i o n can a l s o have a marked e f f e c t on g e r m i n a t i o n . Allen  ( 1 9 5 8 ) r e p o r t e d t h a t a l t h o u g h many i n t e r i o r l o t s of  f i r seed germinated  r a p i d l y without pre-treatment  Douglas-  or s p e c i a l  con-  d i t i o n s , most c o a s t a l l o t s were s l u g g i s h u n l e s s p r e - t r e a t e d o r subjected to s p e c i a l conditions during incubation.  C o a s t a l seed  sown l a t e i n the s p r i n g i n the n u r s e r y o r f i e l d may  not  germinat-  ed u n t l l l the f o l l o w i n g y e a r u n l e s s p r e v i o u s l y s t r a t i f i e d . g e n e r a l , c o a s t a l , seed appeared ed but was  In  t o be more "dormant" when u n t r e a t -  a f f e c t e d by seed parent and s i t e , n u t r i t i o n p r o v i d e d  by the p a r e n t , cone and seed m a t u r i t y , cone s t o r a g e c o n d i t i o n s , p r o c e s s i n g , and seed s t o r a g e .  MATERIALS AND METHODS A.  Thousand-Seed  Weight  A t o t a l Of 124 provenances  (Table 1 and 2 , F i g u r e 1)  were  used I n t h i s study, o f which 91 provenances c o l l e c t e d from B r i t i s h Columbia, Washington and Oregon i n 1 9 6 6 , w h i l e t h e c o l l e c t i o n was made f o r t h e remaining 3 3 provenances from Oregon and C a l i f o r n i a i n 1968. The e x p e d i t i o n s were o r g a n i z e d by I n t e r n a t i o n a l Union of F o r e s t r y Research O r g a n i z a t i o n , S e c t i o n 2 2 .  Come samples were  c o l l e c t e d from t h e south a s p e c t o f t h e middle p a r t o f t h e crown o f the t r e e ; t e n t o twenty dominant t r e e s 160 t o 320 f e e t a p a r t i n each s t a n d were sampled  (Barner, 1966 and L i n e s , 1 9 6 7 ) .  Twenty  cones were shipped t o t h e 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 a f t e r c o l l e c t i o n each y e a r . E x t r a c t i o n , dewinging and c l e a n i n g were made c a r e f u l l y by hand a t room temperature, then t h e seeds were s t o r e d i n a c o l d s t o r a g e room  (0°-2°).  A f t e r seed e x t r a c t i o n , f i l l e d seeds were separated from t h e empty ones u s i n g X-ray f l u o r o s c o p y . used ( S z i k l a i ,  The f o l l o w i n g c l a s s e s were  1964)1  Endosperm. 1.  Seed completely empty o f endosperm.  2.  Shrunken endosperm i n h o r i z o n t a l and v e r t i c a l  position:  l e n g t h l e s s than 1 / 3 o f t h e t o t a l seed l e n g t h ; rounded i n shape, o r occupying t h e middle p a r t o f t h e seed cavity. 3*  Insect l a r v a  inside.  4.  Endosperm f i l l s hut  out most o f t h e seed c a v i t y .  A narrow  conspicuous empty space e x i s t s between t h e endosperm  and t h e seed coat. 5.  Endosperm f u l l y o c c u p i e s t h e seed c a v i t y .  Embryo« 1.  Embryo absent.  2.  P a r t o f embryo I s v i s i b l e , t h e t o t a l l e n g t h o f v i s i b l e p a r t i s l e s s than 50% o f t h e l e n g t h o f t h e seed.  3.  Same as 2 , but t h e v i s i b l e p a r t i s between 5®% and 75%*  4.  Same as 2 , but v i s i b l e p a r t i s more than 75%*  The s e l e c t e d seeds had a w e l l - d e v e l o p e d endosperm and embryo 7 5 - 1 0 0 $ o f t h e embryo c a v i t y .  that f i l l s  From each t r e e 500 f i l l e d  seeds, o r as c l o s e t o t h a t number  as p o s s i b l e , were weighed and t h e f i n d i n g s used t o c a l c u l a t e t h e 1000-seed  weight and t h e average weight o f seed o f a p a r t i c u l a r  provenance. The f i l l e d  seed o f a l l t h e samples were u n i f o r m l y separated  i n t o envelopes, a l l o f which were p l a c e d i n a d e s i c c a t o r f o r 48 hours b e f o r e weighing. b a l a n c e r e a d i n g t o 10 B.  Cone-Scale  A l l seeds were weighed with an a n a l y t i c a l  4 grams.  Morphology  From t h e same provenances used i n P a r t A, two cones were randomly s e l e c t e d from each t r e e and s i x s c a l e s w i t h b r a c t s were t a k e n from t h e middle o f each cone a f t e r seed e x t r a c t i o n .  These  s i x s c a l e s were mounted, t h r e e on t h e a b a x i a l and t h r e e on t h e a d a x i a l s i d e on sheets o f paper ( F i g u r e 2 ) .  On t h e average,  15 t r e e s from each provenance were r e p r e s e n t e d by 90  cone-scales.  F i v e measurements were made ( F i g u r e 3/a) on each s c a l e , width o f s c a l e  (Wi), width o f b r a c t  l e n g t h o f 1st prong (L2)» of the b r a c t  (W2),  length of scale ( L i ) ,  l e n g t h o f 2nd prong (L3)  and p o s i t i o n  (R) i n r e l a t i o n t o the s c a l e was r a t e d ( F i g u r e  A l l o f t h e s e f i v e measurements were measured t o the n e a r e s t m i l l i m e t e r u s i n g the Swedish Tree Ring measuring  3/b)« 0.1  equipment.  The c o n e - s c a l e width was measured a t t h e widest p a r t o f the s c a l e w h i l e the width o f b r a c t was measured a t t h e base o f the prongs. I f t h e two s i d e prongs were not t h e same l e n g t h , then the 1st prong l e n g t h was measured from t h e base o f t h e deeper i n d e n t a t i o n and the 2nd prong l e n g t h was measured from the base o f the same i d e n t a t i o n .  C.  R e l a t i o n s h i p Between Thousand-Seed acteristics  Weight and Cone-Scale Char-  Data from P a r t s A and B were used t o i n v e s t i g a t e these r e lationships.  D.  Seed Germination T e s t Out o f 124 provenances c o l l e c t e d 114 were used i n t h i s  experiment  (Table 1 ) .  F i f t y - s i x f i l l e d seeds were s e l e c t e d t o  r e p r e s e n t two r e p l i c a t i o n s t r e e i n a provenance.  (28 seeds per r e p l i c a t i o n ) o f each  A t o t a l of 1 , 3 6 8  i n d i v i d u a l t r e e s were  r e p r e s e n t e d and 7 6 , 6 0 8 seeds were sown. Seeds were sown without p r e s o a k i n g o r s t r a t i f i c a t i o n i n the U n i v e r s i t y o f B r i t i s h Columbia Southern Campus F o r e s t  Re-  s e a r c h Nursery between May  3-9,  1969*  Ten n u r s e r y beds were e s t a b l i s h e d d u r i n g the s p r i n g of 1969  The 30-cm n u r s e r y beds  w i t h t h r e e n u r s e r y beds i n a row.  were f i l l e d w i t h C a l i f o r n i a mix o f 25 cm.  Seeds were sown 0 . 6  sand w i t h 2 . 5  cm x 10  to a  depth  cm deep and covered w i t h f i n e  cm s p a c i n g .  An i r r i g a t i o n system was Germination  (C) (Baker, 1957)  a l s o provided.  counts were made a t 3 6 ,  50,  72  and 92 days  a f t e r sowing. The data were s t r a t i f i e d a c c o r d i n g t o seed c o l l e c t i o n zone maps g i v e n by Haddock and S z i k l a i  ( 1 9 6 6 ) f o r Canada, by Western  F o r e s t Tree Seed C o u n c i l (1966) f o r Washington and Oregon, and by Buck et a_l. ( 1 9 7 0 ) f o r C a l i f o r n i a .  Each c l i m a t i c r e g i o n  d i v i d e d i n t o s e v e r a l sub-regions a c c o r d i n g t o geographic and s o i l and c l i m a t i c c o n d i t i o n s ( F i g u r e  1).  was  factors  Figure 1  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 Coast of North America D o u g l a s - f i r provenance study.  Pseudotsuqa m e n z i e s i i (Mirb.) Franco  »• •  Provenance No.; i -1966. I. U. F. R. O.  Pseudotsuga menziesii (Mirb.) Franco Provenance No.; I -1966. I.U.F.R.O.  D X E H  r o  35  15 Faculty of (Wastry  Q7B.C V a n c o u v e r .  U.B.C. V a n c o u v o r .  Pseudotsuqa m e n z i e s i i (Mirb.) Franco  Pseudotsuqa menziesii (Mirb.) Franco  Proven a nee No.;11 -1966. I. U. F. R. O.  Provenance No.; 11 -1966. I. U. F. R. O.  •  V .  SB T O  Ma  E  lotus  GO* a t  IS Faculty  of  Forestry  U.B.C. v m i c o u v w .  Pseudotsuga meniiesii(MirbJFrancp P r o v e n a n c e N o ; 1 2 - 1 9 6 6 . I. U.F. R.  Faculty  of  Provenance No.; :t-1968. I. U. F. R. O.  U.B.C. Vancouver.  tfei*»try  Pseudotsuga m ^ n j ^ « i i (MjrbAFranco Provenance No.;H-1966. I. U. F. R. O.  acultPseudotsuga I Of  Pseudotsuga m e n z i e s i i (Mirb.) Franco  M  Forestry  U . B . C . Vancouwer.  menziesi i (Mirb.) Franco  Provenance P r o v e n a n c e No.;>lV-1fl[68. m  I.U.F.R.O.  U.B.C. Vancouver.  F i g u r e 2.  View o f a b a x i a l and a d a x i a l s u r f a c e o f cones c a l e s from Prov. No. 1,11,12 and 124.  - 18 -  r«  W,  •  F i g u r e 3 / a •D i a g r a m m a t i c r e p r e s cone-scale and bract showing five basic measurements-  .F i g u r e 3 / b • P o s i t i o n o f t h e b r a f c t i s c a l e •  T a b l e 1.  L o c a t i o n o f 124 D o u g l a s - f i r provenances (No. 1 t o 91 were c o l l e c t e d I n 1966, No. 92 t o 124 were c o l l e c t e d  i n 1968).  British  Columbia  Provenance No. 1. 2. 3. 4. 5. 6. 7. 8.  9.  10. 11. 12.  14. 15. 16. 17.  18. 19.  20. 21. 22.  23.  24.  25.  26. 27.  28.  29. 30. 31. 32. 33.  Stoner Dean Stuie* Alexandria W i l l i a m s Lake Kllnaklini* Tatla Barriere Clearwater Revelstoke Golden Jeune Landing Nlmpkish Owl Creek Merritt Chase Monte Creek Salmon Arm Tahsls I n l e t * Forbidden Plateau Courtenay Albernl Cassidy Sechelt Squamish C h i l l i w a c k Low C h i l l l w a c k High Nelson Caycuse Jordan R i v e r * San Juan R i v e r Duncan Sook  Average elevation feet 1900 20 750  2100 2000 10  2900 1400 1500 2000 2700 550 300 700  .2700 1650 2100 1550 50 2000 220 450 650 600 50 55© 3000  Latitude  53 37 52 48 52 22 52 41 52 06 51 07 51 ^4 51 12 51 39 51 00 5123 50 27 50 19 50 20 50 04 50 33 50 37  Jo 44  49 47 49 4o 49 4i 49 19 49 03  Longitude  122 40 126 57 126 00 126 26 122 00 125 36 124 44 120 09 120 00 118 12 117 00 127 27 126 53  122 120 119 119  119 13 126 38  125 09 125 03  124 51 123 57 123 53 123 09  121 48 121 42 117 16 124 26 124 14 124 05  15©  49 31 49 47 49 04 49 06 49 03 48 55 48 28 48 35 48 45 48 20  1000 200 200  48 04 48 02 48 32  124 00 124 00 122 19  2700 700 800 700  200  123 123  JK  No. o f trees  ll 6  16 16 10 16 15 16 15 15 15 13 1| 16 16  li 11 15 15 15 13 15 15 15  ll ll 16 13  15  Washington 34. 35. 36.  Lake Crescent Sequim Bay Sedro Woolley  14 14 16  Table 1.  (Continued)  Provenance No. 37. 38. 39.  40. 41.  42. 44*.'  46. 47. 48. 49. 50. 51* 52.  Jfc 55. 56. 57. 58. !6 90'. 61. 62.  ll: 65. 66.  U:  69. 70. 71. 72.  Arlington Granite F a l l s Concrete Darrlngton Bacon P o i n t P e r r y Creek Marblemount S l o a n Creek D i a b l o Dam Twisp Republic Newport Forks Hoh R i v e r Humptullps Matlock Matlock Shelton Gard S t a t i o n Enumclaw North Bend Chest Morse Lake Parkway Denny Creek Gold Bar Skykomish Keechelus Lake C l e ELum Chiwachum Spokane Naselle Skamokawa Cathlamet C a s t l e Rock Yelm Yale Cougar A l d e r Lake Handle Packwood GlenwoodRimrock Prindle Willard  U: 75*  76. 77. 78. 79. 80.  No. o f trees  Average elevation feet 300 300 1550 500 1650 2000  400  2150 1450 2600 2400  2400 300  800  450 1650  400  300 1500  800  500 2000  2400 1800 400 1000 2600 2100  1800  2000 150 700 650 500 200  400  1650  1400  1100 2150 1600 2500 1500 1650  48 13 48 05 48 39 48 16 48 36 48 0 3 48 35 48 05 48 43 48 2 3 48 36 48 12  122 122 121 121 121 121 121 121 121 120  47 48 47 19 47 18 47 15  47  59  47  15  118 117  04 02 43 38 23  28 24 18 07  24 44 03  124 24  45  37  123 58 123 5 4 123 26 123 25 123 12 123 0 5 121 56 121 45 121 40 121 34 121 32 121 39 121 20 121 22 121 07 120 4 4 117 12 123 4 4 123 30 123 16 122 52 122 44 122 22 122 18 122 17 122 0 3 121 40 121 00 121 02 122 08 121 4 l  45  13  123  48 00 47 16 47 28 47 22 47 02 47 24 47 47  47 2 3 4? 13 47 41 47  47  46 22 46 21 46 18 46 19 47 01 46 00 46 0 5 46 48 46 33 46 34 46 00 46 40 45 48  If 16 16 15  li li 15 15 if 16  14 14 14 14 14 16 15 15  16 16  li  15 15 15 If 16 15  li ll li 15 15  16  14  16 15 15 15  Oregon 81.  Hebo  500  51  15  T a b l e 1.  (Continued)  Provenance No.  82. 83. 84. 85. 86. 87. 88. 89. 90.  91. 92. 93.  94.  95. 96.  97. 98.  99.  100. 101. 102. 103. 104.  Grand Bond Vernonla Sandy Cherryvilie P i n e Grove Waldport Upper Soda Coquille Olalla* Brookings Burnt Woods Mary's Park Eugene Corvallls M i l l City Detroit Marlon Forkes* Roseburg Steamboat Oakridge Cave J u n c t i o n Wolf Creek Ashland  Average elevation feet 600 700 900 2200  2400  200 3250 200 1100 1000 1100 3250 700 250 550 1600 3500 900 5250  2900 1400 1400  4900  No. o f trees  Latitude 06  46  23  19  4 06 44* 24 5  23 12 05 07 36 30 01  42 48  44 30 19 22 54 42 11  41 05  123 123 122 122 121 123 122  36 13  18  08 23 52 12 124 10 123 34 124 12 123 42 123 34 123 23 123 13 122 24 122 10 122 00 123 30 122 31 122 22 123 40 123 23 122 39  16 15 15 15 15 15  15 14  8 16 16 15 15 16 15 16 6 16  14  16 16 15 15  California 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115.  Gasquet Happy Camp Sawyers Sawyers Bar S c o t t Bar Seiad V a l l e y . Hawkinsville* Dunsmuir Burney Areata Areata  400 4100 4750 3800 3300 2600 3500 3300 3350 1600 2900  117. 118. 119. 120. 121. 122.  B i g Bar Wildwood Weaversville F o r t Bragg* Covelo Covelo Alder Springs* Lower L a k e *  £300 3900 3750 200 3000 5100  116.  123. 124.  B i g Bar  3250  4500  3100  41 51 41 39 4 i 16 4 l 17 4 l 44 4 l 48 4 l 47 4 i 12 41 05 40 5 40 * 4o 43 4o 47 40 2 40 5 39 30 39 55  39 48 39 39 38 50  123 123 123 123 123 123 122 122 121 123 123 123 123 123 122 123 123 122 122 122  59 31 09  08  06 00 40  18  39  50  46  18  12 00 44 43  li li  16 12 15  li  16 16 15 15 16 16  18  ii  42  15 15 15  56 45  * provenance not i n c l u d e d i n t h e g e r m i n a t i o n t e s t s .  T a b l e 2.  Number o f provenances and t r e e s sampled i n 1966 and 1 9 6 8 .  Year o f c o l l e c t i o n  1966 Province or State  1968 Total  Number of  Provenances  Trees  —  33  474  —  47  710  191  24  342  20  292  20  292  33  483  124  1,818  Provenances  Trees  B r i t i s h Columbia  33  474  —  Washington  47  710  —  Oregon  11  151  13  California  ——-——.  —  Total  91  1.335  Provenances  Trees  RESULTS AND  DISCUSSION  Thousand-Seed Weight The mean v a l u e s of 1000-seed weight of t h e 124 c o l l e c t e d f o r the present study ranged from 6.9  provenances  t o 18.0 grams  (Table 3 ) : these v a l u e s d i f f e r from those o f both Ching and Bever  ( i 9 6 0 ) and Sweet (1964).  From a t o t a l of 1,818  t r e e s r e p r e s e n t e d i n t h i s study, the 1000-seed weight over these t r e e s ranged from 5*3 t o 24.8  from a l l  grams (Table 3), and  t h i s range covers the f i n d i n g s of Ching and Bever Sweet (1964).  individual  (I960)  and  The d i f f e r e n c e s were p o s s i b l y due t o t h e fewer  number of sample t r e e s i n the l a t t e r s t u d i e s when compared t o an average of 15 t r e e s r e p r e s e n t i n g each seed source i n t h i s study. Another p o s s i b l e f a c t o r i n f l u e n c i n g the seed weight ent y e a r of sample c o l l e c t i o n .  While Ching and Bever  Sweet (1964) c o l l e c t e d i n 195^ and 1956 respectively. i n g 1966 and  i s the d i f f e r -  and I n 1955  (i960)  and  and  1956  The seed samples i n t h i s study were c o l l e c t e d dur1968.  R e g r e s s i o n a n a l y s e s were c a r r i e d out between 1000-seed weight and l a t i t u d e and e l e v a t i o n of seed source (Part A ) , between cone-scale c h a r a c t e r i s t i c s and l a t i t u d e and e l e v a t i o n of seed source (Part B) and between 1000-seed weight and scale characteristics  cone-  (Part C) a l l based f i r s t l y on s u b - r e g i o n s .  No s i g n i f i c a n t r e l a t i o n s h i p s c o u l d be e s t a b l i s h e d , except some i n sub-regions l c , 3*>, and 3c, so the r e g r e s s i o n a n a l y s e s were then based on r e g i o n r a t h e r than s u b - r e g i o n .  S i g n i f i c a n t seed-weight v a r i a n c e s among d i f f e r e n t  geo-  g r a p h i c a l l o c a l i t i e s , t r e e s and stands have been r e p o r t e d bys e v e r a l a u t h o r s ( S q u i l l a c e , 1965? Sweet, 1 9 6 4 ; and Simak, 1 9 6 ? ) .  The d a t a (Tables 3 and 4)  Anderson, showed t h a t  19&5 1000-  seed weight v a r i e d g r e a t l y among provenances, sub-regions and regions.  These r e s u l t s were supported by Simak (19&7)  s t u d i e d seed weight o f European l a r c h from d i f f e r e n t provenances.  ( L a r l x decldua ( M i l l . ) )  He found s i g n i f i c a n t  i n seed weight among d i f f e r e n t  who  differences  regions.  There a r e s e v e r a l e x t e r n a l f a c t o r s which may modify t h e 1000-seed  weight f o r a t r e e .  P e r r y and Coover ( 1 9 3 3 )  reported  t h a t l a r g e r cones g e n e r a l l y y i e l d l a r g e r seeds i n p i t c h p i n e . Simak ( 1 9 5 ^ ) a l s o r e p o r t e d t h a t cone s i z e and cone weight were the f a c t o r s I n f l u e n c i n g t h e average seed weight i n S c o t s p i n e . He mentioned t h a t seed weights per cone i n c r e a s e d w i t h i n c r e a s i n g cone weight and d e c r e a s i n g seed number i n the mother tree. Other e x t e r n a l f a c t o r s , such as the p o s i t i o n o f a t r e e i n a stand, t h e p o s i t i o n o f t h e seed i n a cone, c l i m a t e and edaphic f a c t o r s , t h e age o f t h e t r e e and t h e number of cones produced by a t r e e can a l s o I n f l u e n c e t h e seed weight. However, Sweet ( 1 9 6 4 ) d i d not recommend seed weight as a u s e f u l measure f o r i n d i c a t i n g provenance d i f f e r e n c e s owing t o the extent t o which i t may  be a f f e c t e d by degree o f c l e a n i n g ,  year o f c o l l e c t i o n and age o f parent t r e e .  A l l the seed sam-  p l e s i n t h i s study were c l e a n e d c a r e f u l l y by hand.  Dewinging  was carried out i n such a way that nothing except that portion of the wing d i r e c t l y attached to the seed remained with i t , and so the data of seed weight i n t h i s study appears more r e l i a b l e * Simak ( 1 9 6 7 ) pointed out that as thousand-seed weight values of European l a r c h seed from d i f f e r e n t geographical regions were constant and s p e c i f i c they could be used as a c r i t e r i o n f o r the i d e n t i f i c a t i o n of the o r i g i n of l a r c h .  However, the thousand-  seed weight v a r i a t i o n among the regions had a c l i n a l character, which made i t d i f f i c u l t or impossible to determine the o r i g i n of provenance material l y i n g on the boundary of two neighouring regions. Figures 4, 5 and 6 show that 1000-seed weight was strongly correlated with elevation i n Regions 1, 2 and 3 .  In other words,  provenances from higher elevations developed heavier seed than those from lower elevations. the  While the r e s u l t s substantiate  findings of Sweet's (1964) work on Douglas-fir and of Simak's  (196?)  on European l a r c h , they are .contrary to the statements of  Mirov e l at,  (1952) on ponderosa pine.  No relationships were observed between 1000-seed weight and elevations of seed source i n i n t e r i o r Regions 5 and 7 (Table 5)» perhaps because elevations of the provenances collected from these two areas ranged only from 1,400 to 2 , 9 0 0 feet, and  1,500  to 2 , 7 0 0 feet respectively, or because of the difference between coast and i n t e r i o r . Correlations between 1000-seed weight and l a t i t u d e of seed source i n coastal Regions 1, 2 and 3 (1$ l e v e l ) , and i n i n t e r i o r Regions 5 and 7 (5# l e v e l ) were negative (Figures 7 to 11). c o e f f i c i e n t of determination R  2  The  of these relationships (Table 5)  was  c o n s i d e r a b l y h i g h e r than t h a t f o r the 1000-seed weight v e r s u s  e l e v a t i o n o f seed source i n both c o a s t a l and I n t e r i o r r e g i o n s . L a t i t u d e was  c l e a r l y more important than e l e v a t i o n i n a f f e c t i n g  seed weight, and t h e r e s u l t s I n d i c a t e t h a t seed weight was  sub-  j e c t t o c l i n a l v a r i a t i o n , and i n c r e a s e d from n o r t h t o south. The 124 provenances were then d i v i d e d i n t o f i v e d i f f e r e n t 500-feet e l e v a t i o n c l a s s e s from sea l e v e l t o 2 , 5 0 0 - f e e t .  Because  t h e r e were o n l y 30 provenances between 2,500 t o 5»5©0 f e e t o f e l e v a t i o n , t h e s e were combined  Into a simple s i x t h group t o make  t h e i r c o n t r i b u t i o n more even from n o r t h t o south. C o r r e l a t i o n a n a l y s i s was  c a r r i e d out between  weight and l a t i t u d e f o r each group.  1000-seed  F i g u r e s 12a t o 12f ( T a b l e 6)  i n d i c a t e s i g n i f i c a n t n e g a t i v e r e l a t i o n s h i p s between weight and l a t i t u d e f o r each group.  1000-seed  Seed weight appears t o i n -  crease from n o r t h t o south even w i t h i n a c e r t a i n range of e l e v a t i o n c o n f i r m i n g p r e v i o u s f i n d i n g s t h a t l a t i t u d e was more imp o r t a n t than e l e v a t i o n as a f a c t o r a f f e c t i n g seed weight.  This  agrees w i t h c l i n a l t r e n d s i n seed and cone l e n g t h o f D o u g l a s - f i r which i n c r e a s e d from n o r t h t o south ( S z i k l a i , 1969), and the n u c l e a r volume and DNA Sziklai,  1971).  content o f D o u g l a s - f i r (EL-Lakany and  -  LZ  -  The relationship between 1000-seed weight and latitude18  Y=3I 0 3 - 0 45 X  18  Sub-region Sub-region Sub-region Sub-region Sub-region  16 16  E  V)  E  2a 2b 2c 2d 2e  S 14  2 14  £ 121  12  10  10  Y=50 7 - 0 85X Sub-region Sub-region Sub-region Sub-region  Sr-  _L 40  c 38  la laa lb Ic  _L _L J. 42 44 46 Latitude (Degrees North)  _L 48  _L '38  50  Fig ure 7- in region Iv  40  _L _L 42 44 46 48 Latitude (Degrees North)  50  52  50  52  Figure 8- in region 2-  I8H  I8r-  16  16  E  14  |  I4|  2z  o>  S  12  * 12  10  8  10  O • A X A  '38  Sub-region Sub-region Sub-region Sub-region Sub-region  40  3a 3b 3c 3d 3e  o  42 44 46 48 Latitude (Degrees North)  %1  50  Figure 9- in region 3  38  40  42 44 46 48 Latitude (Degrees North)  Figure 10- in region 5-  „ 14  r 12 Y-5673-0-93X O •  10  Sub-region 7a Sub-region 7b  O  o o 6>38  40  42  44 46 48 Latitude (Degrees North)  Figure II- in region 7-  50  52  54  Figure I2a-I2f- The relationship between 1000-seed weight and elevation from south to north for each 500 feet of elevation from sea level to 2,500 feet,and from 2,501 to 5,500 feet-  Latitude (Degree* North)  Figure 12 b- from 501 to 1,000 feet-  Lotitude (Degrees North)  Figure I2e- from 2,001 to 2,500 feet-  T a b l e 3.  Provenance No. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 9 0 41 42 43 44 45 46 47 48  Maximum, mlnlmui 1 and mean v a l u e s o f 1000-seed weight f o r each provenance i n grams. Maximum  Minimum  10.4157 11.6410 13.1456 12.5810 12.9982 12.4570 13.4160 14.0160 13.1684 10.4410 IO.3789 12.5692 8.6244 11.7630 14.4784 12.7020 15.2941 11.0748 9.7637 10.0879 11.1611 11.2878 13.8250 9.9979 11.4429 10.9955 9.5923 12.1609 12.3010 8.1280 11.0671 13.3383 9.7581 11.7197 11.0112 12.1210 II.7I88 11.4486 13.2034 12.3391 12.5974 13.4100 13.8350 14.5243 14.1661 I8.6756 17.2660 13.4961  7.3442 6.5926 7.8697 7.1285 7.3761 8.7799 8.7580 7.7428 7.4llO 6.7078 7.4270 8.7835 5.7190 6.7380 8.3531 7.4768 8.5368 6.3776 5.4673 5.9247 8.9212 8.9323 7.1690 6.4736 7.5006 6.4779 5.8902 6.9940 8.1506 5.6000 6.4799 7.4621 5.2818 7.2520 6.7916 7.8681 6.7659  6.1281 8.8963 7.8600 6.7740 8.4724 8.4024 8.2532 9.1316 IO.6655 8.1290 8.8019  Mean 8.6076 9.1026 IO.3307 9.3839 10.4723 10.3733 10.7942 10.1530 9.6942 8.8204 9.0777 10.0874 7.1422 9.7725 11.3622 10.3224 10.6012 8.7206 8.1081 8.0032 9.8105 10.1293 9.1725 8.3230 9.0917 8.4357 7.3223 9.2489 10.1977 6.9189 9.0643 IO.2133 7.7601 9.0075 8.3056 10.0224 9.4333 7.5752 10.8358 10.2248 9.9574 10.9612 10.5608 11.8493 11.0600 13.2947 12.3406 11.4489  Table 3 .  (Continued)  Provenance No. 49 50 51 52  J*  II 57 58 59 60 61 62  ll 65 66  f7 68 69 70 71 72 7 7 7 £ 76  77 78 79 80 81 82  12  U  87 88 89 90 91 92 9 9 95 96 97 98 99  Maximum 1.5065 4.6888 0.0324 3.3207 3.2467 3.6776 1.5000  ^*5576 3.7705 3.1992 2.9394 2.4396 1.424? 3.4964 2.2926  1.4*41)4 6.6393 3.484? 2.2843 3.1826 I.6980 3.9639  4.3908  3.7189 2.1506 4.9367 0.1613 5.6414 6.4904 3.8138 5.9030 2.8976 3.5474 4.2596 5.3184 4.0440 7.4502 3.6275 5.3074 3.2626 7.3469 6.0316 2.5676 .4916 .9245 4.1150 4.2228 3.3039 2.2684 3.8706  I  Minimum 6.9489 8.2028 7.2561 8.0791 6.3729 8.7912 6.8756 8.0860 5.6535 6.8344 8.9466 9.2474 6.8340 7.9H7 7.7674 8.4684 8.4940 IO.6508 7.8562 8.3971 9.3753 8.5478 9.8874 10.4883 9.0957 8.3853 9.0173 6.5407 10.4312 12.0572 9.2682 8.2644 8.2129 8.3798 8.4120 6.9978 9.4015 9.2063 7.0437 9.73©4 8.O658 12.0451 7.6o47 8.1660 8.5444 9.2127 9.1581 8.2909 8.I863 10.3650 7.8593  Mean 8.8598 10.8120 8.8665 10.0426 9.9037 11.2815 9.0224 11.2797 9.1742 9.9714 10.?4l9 10.9442 9.69?6 10.704i 10.5624 10.3615 13.7989 13.5056 10.5983 10.5509 10.5858 9.9515 12.0033 12.2660 10.9332 9.93% 11.7957 8.4889 13.5443 13.7810 11.7812 12.2576 10.8803 11.0515 11.7173 11.1045 11.7882 14.2043 9.5850 12.3854 10.4077 14.213© 12.0091 9.8074 II.6376 12.1002 11.5786 11.2457 11.1186 11.1930 11.4927  Table 3 .  (Continued)  Provenance No. IOOJ 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124  32 -  Maximum 15.6991 18.8851 18.1974 14.392£ 19.4334 15.1628 18.7659 20.3956 18.3912 22.4733 20.5615 22.2198 20.6708 20.8763 14.7973 18.7719 20.9691 20.0613 21.0619 19.5550 12.5857 18.8295 22.7006 20.0666 24.7686  Minimum 9.0286 9.4846 11.6566 10.3434 13.8145 10.2818 11.9469 11.9888 11.7760 12.8328 14.8266 12.8112 10.9028 15.7712 9.7002 11.0146 15.0369 13.2789 13.0980. 11.5797 10.8928 11.8890 12.1657 14.0297 11.9000  Mean 12.5702 12.4379 14.6608 12.5340 15.8762 13.1355 15.2213 15.4023 15.4171 17.2340 16.9969 16.1968 15.1707 18.0451 11.9590 13.9353 17.8080 16.7025 16.1422 15.4638 11.6085 15.0325 17.4883 17.1401 16.0461  Table 4 .  Region  la laa lb lc  Average v a l u e s and standard d e v i a t i o n s o f 1000-seed weight by c l i m a t i c r e g i o n s i n grams. Total No. of Prov.  1000-seed weight X  ±SD  1  9.51^9 9.6752 II.6758 16.1499  1.5403 1.8886 I.8765 2.3421  5 15  2a 2b 2c 2d 2e  6 7 6  I  8.2456 9.8539 10.9166 10.9298 12.8700  1.5382 I.7017 1.4496 1.9904 1.9803  3a 3b 3c 3d 3e  3 11 13 9 2  9.4598 9.8484 10.3819 12.2069 16.8537  I.4708 1.9645 I.7187 l.?473 2.4813  9.7720  1.4528  4 5a 5b  5 8  10.4696 12.5120  1.6847 2.2363  6  2  9.0082  1.2938  3  9.3664 11.4500  7a 7b  2.28.12  Table 5.  Relationship between 1000-seed weight and l o c a t i o n of seed source.  Character  Elevation  Latitude  No .of Prov.  R2  r  R2  r  Region 1 1000-seed weight  0.78  0.88**  0.87  0.93**  29  Region 2 1000-seed weight  0.19  0.44*  0.59  0.77**  30  Region 3 1000-seed weight  0.23  0.48**  0.64  0.80**  38  Region 5 1000-seed weight  0.01  0.06NS  0.40  0.63*  13  Region 7 1000-seed weight  0.03  0.17NS  O.63  0.79*  7  R and r values i n the above Table represent the relationships between 1000-seed weight and elevation (Pigs. 4 - 6 ) , between 1000-8eed weight and l a t i t u d e (Figs. 7-11). Only the s i g n i f i cant relationships were graphed. NS • not s i g n i f i c a n t . * = s i g n i f i c a n t at 5% l e v e l . ** = s i g n i f i c a n t at 1% l e v e l .  T a b l e 6.  R e l a t i o n s h i p between 1000-seed weight and l a t i t u d e w i t h i n v a r y i n g ranges o f e l e v a t i o n .  Latitude  Character R 1000-seed weight e l e v a t i o n range 1-500 f t .  2  r  No. o f Prov.  0.33  -0.57**  31  1000-seed weight e l e v a t i o n range 501-1000 f t .  0.^3  -0.64**  22  1000-seed weight e l e v a t i o n range 1001-1500 f t .  0.50  -0.71*  12  1000-seed weight e l e v a t i o n range 1501-2000 f t .  0.33  -O.58**  18  1000-seed weight e l e v a t i o n range 2001-2500 f t .  0.62  -0.78**  11  1000-seed weight e l e v a t i o n range 2501-5500 f t .  0.74  -0.86**  30  R* and r v a l u e s i n t h e above T a b l e r e p r e s e n t the r e l a t i o n s h i p between 1000-seed weight and l a t i t u d e from f i g u r e s 12a t o 12f. * = s i g n i f i c a n t a t $% l e v e l . ** «= s i g n i f i c a n t a t 1% l e v e l .  Cone-Scale  Morphology  The average  width o f cone-scales  was 22.81 mm w i t h a range (Table 7) B a r r i e r e , B.C.) t o 2 6 . 1 7 mm  (W^) f o r a l l 124 provenance  from 1 9 . 5 7 (Prov. No. 8,  (Prov. No. 1 1 7 ,  B i g Bar, C a l i f o r n i a ) .  The average v a l u e o f cone-scale width o f d i f f e r e n t c l i m a t i c subr e g i o n s was w i t h a range (Table 8) from 20.1 (Region 6 - no subr e g i o n ) t o 2 4 . 9 mm  (sub-region 3 e ) .  A n a l y s i s o f v a r i a n c e (Table 9/b) from t h e data o f cone-scale width shows s i g n i f i c a n t d i f f e r e n c e s among t r e e s w i t h i n provenance, among provenances w i t h i n sub-regions, among sub-regions w i t h i n r e g i o n s and among r e g i o n s . The width o f cone-scale was c o r r e l a t e d with e l e v a t i o n i n 1 ( F i g u r e 13)  Region  {1% l e v e l ) and Region  3 (Figure l 4 ) ( 5 ^ l e v e l ) .  T h i s i n d i c a t e s t h a t provenances from h i g h e r e l e v a t i o n i n Regions 1 and 3 had wider c o n e - s c a l e s .  No r e l a t i o n s h i p c o u l d be e s t a b -  l i s h e d between width o f cone-scale and e l e v a t i o n s i n Regions 2, 5 and 7  (Table 9 / a ) .  Width o f b r a c t (W2)  tended t o be c o r r e l a t e d w i t h  and was found t o be s i g n i f i c a n t o n l y I n Region Table  ( F i g u r e 1 5 , and  10/a). The average  l e n g t h o f cone-scale from 124 provenances com-  b i n e d was I 8 . 6 9 mm with a range (Table 7) 17,  7  elevation  Monte Creek, B.C.) t o 2 2 . 0 5 mm  fornia). sub-region  The average  from 15-82 (Prov. No.  (Prov. N 0 . 1 2 3 ,  Covelo,  l e n g t h o f cone-scale o f d i f f e r e n t  (Table 8) from 1 6 . 4  (sub-region 7 a )  Cali-  climatic  t o 2 1 . 0 mm (sub-  region 3e) The l e n g t h o f cone-scales were s t r o n g l y c o r r e l a t e d  ( 1%  l e v e l ) with e l e v a t i o n i n Region 1  ( F i g u r e 16) w h i l e F i g u r e  indicates a significant correlation  17  ( 5 $ l e v e l ) with t h e e l e v a t i o n  i n Region 3 . N e i t h e r width n o r l e n g t h o f cone-scale were c o r r e l a t e d with e l e v a t i o n i n coast Region 2 and I n t e r i o r Regions 5 and 7 9/a  and 11/a).  (Tables  I n o t h e r words these c h a r a c t e r i s t i c s appeared t o  be independent and not I n f l u e n c e d by e l e v a t i o n i n these The average l e n g t h o f 1 s t prong ( L 2 )  regions.  f o r a l l 124 provenance  was 8 . 1 3 mm with a range from 5 . 8 5 (Prov. No.4o, D a r r l n g t o n , Washington) t o 1 0 . 7 5 mm  (Prov. N 0 . I O 3 . Wolf Creek, Oregon).  A n a l y s i s o f v a r i a n c e (Table 12/b) from t h e data shows t h a t  1st  prong l e n g t h s a r e s i g n i f i c a n t l y d i f f e r e n t among t r e e s w i t h i n provenances, among provenances w i t h i n sub-regions r e g i o n s w i t h i n r e g i o n s , but not among r e g i o n s .  and among sub-  From r e g r e s s i o n  a n a l y s i s c a r r i e d out between 1 s t prong l e n g t h and e l e v a t i o n , a significant relationship  (5% l e v e l ) ( F i g u r e 18) was found i n Re-  gion 1 only. In a d d i t i o n a r e g r e s s i o n a n a l y s i s was c a r r i e d out between 2nd  prong l e n g t h ( L 3 ) and e l e v a t i o n .  The l e n g t h o f 2 n d prong i s  s i g n i f i c a n t l y c o r r e l a t e d (5% l e v e l ) with e l e v a t i o n i n Regions 1 and  7  ( F i g u r e s 19 t o 20 r e s p e c t i v e l y ) .  2nd  prong l e n g t h from combined provenances was 3 « 7 0 mm with a  range 2.46 109,  The o v e r a l l average o f  (Prov. No.8, B a r r i e r e , B.C.) t o 4.48 mm  S c o t t Bar, C a l i f o r n i a ) .  (Prov. No.  A n a l y s i s o f v a r i a n c e o f 2 n d prong  l e n g t h (Table 13/b) shows s i g n i f i c a n t d i f f e r e n c e among t r e e s w i t h i n provenances, among provenances w i t h i n sub-regions and among sub-regions  within regions.  Analyses of v a r i a n c e  f o r the above-mentioned  c h a r a c t e r i s t i c s showed c l e a r l y the great  cone-scale  intraspeclfic variation  o f D o u g l a s - f i r w i t h i n i t s n a t u r a l range i n Northwest America. Larsen (1937) pointed  out t h a t great i n d i v i d u a l v a r i a t i o n of  D o u g l a s - f i r e x i s t s even from the same g e o g r a p h i c a l  area.  Haddock  (1962) s t a t e d "Considerable v a r i a t i o n i n cone and l e a f morphology, c o l o r of f o l i a g e , e t c . has species."  Sziklai  been noted throughout the range of  the  (1967) mentioned "The v a r i a t i o n i t s e l f i s a  product of d i f f e r e n c e s among I n d i v i d u a l s which a r e the e f f e c t o f environmental m o d i f i c a t i o n s , g e n e t i c recombinations and F i g u r e s 23 and  25  s h i p between cone-scale  show a h i g h l y s i g n i f i c a n t l i n e a r  increased  i n Region 3»  ( F i g u r e s 26  Regions 1 and  3  t o 28)  (1% l e v e l ) , and  The  i n Region 5  (mm2)  s i z e and  cone s i z e Sziklai  (1964)  strongly  cones u s u a l l y h a v i n g l a r g e r s c a l e  (Plnus montloola Dougl).  g i v e f u r t h e r d e t a i l s on whether cone l e n g t h was t o cone-scale  was  cone s c a l e s i n D o u g l a s - f i r .  (1957) r e p o r t e d t h a t average s c a l e s i z e was  i n western white pine  and  from n o r t h t o south.  longer  r e l a t e d t o cone l e n g t h , l o n g e r  length  These  5 both width  (1964) and S q u i l l a c e (1957).  found t h a t wider cones had Squillace  {5% l e v e l ) .  3 and  r e l a t i o n s h i p between cone-scale  s t u d i e d by S z i k l a i  and  were a l s o r e l a t e d t o l a t i t u d e i n  r e s u l t s tend t o show t h a t i n Regions 1, increased  5»  Width of  s i g n i f i c a n t l y from n o r t h t o south, and  o f cone-scale  l e n g t h o f cone-scale  relation-  l a t i t u d e i n Regions 1 and  width and  a c u r v e l l n e a r r e l a t i o n s h i p ( F i g u r e 24) cone-scale  mutation."  l e n g t h or cone-scale  width.  t h a t the average width of cone-scale with average l e n g t h of cone-scale  He d i d not  directly related  S q u i l l a c e a l s o showed  had a p o s i t i v e c o r r e l a t i o n  i n western white p i n e .  The f i n d i n g i n the present study i n d i c a t e d t h a t width  and  l e n g t h o f cone-scale show a c l i n a l v a r i a t i o n from n o r t h t o south i n Regions 1,  3 and 5.  T h i s agrees w i t h the r e g i o n a l c l i n a l v a r (19^9) i n D o u g l a s - f i r cone and  i a t i o n p a t t e r n found by S z i k l a i  seed l e n g t h s , which a l s o Increased from n o r t h t o south. The width and l e n g t h of c o n e - s c a l e were not a f f e c t e d  by  e l e v a t i o n and l a t i t u d e , however, i n Regions 2 and 7. u n l i k e Regions 1,  3 and 5 (Tables 9 /  a  11/a).  and  due t o Region 2 being on the P a c i f i c Coast  T h i s may  possibly  ( F i g u r e 1)  and  be  having  h i g h moisture and r e l a t i v e humidity c o n d i t i o n s ; a l t e r n a t i v e l y c o n e - s c a l e s i z e i n Region 2 c o u l d p o s s i b l y be under s t r o n g g e n e t i c control.  The l a c k o f any r e l a t i o n s h i p s i n Region 7 might be  be-  cause o f the s m a l l e r number of samples (7 provenances) c o l l e c t e d i n t h i s area. F i g u r e s 30 t o 32 show t h a t the l e n g t h s of 1st  prong have a  h i g h l y s i g n i f i c a n t n e g a t i v e c u r v e l l n e a r r e l a t i o n s h i p with  lat-  i t u d e i n Regions 2 and 3» and h i g h l y s i g n i f i c a n t n e g a t i v e l i n e a r r e l a t i o n s h i p i n Region 5»  F i g u r e 29 shows t h a t the 1st  prong  l e n g t h i n c r e a s e d from sub-region l a t o l b and then decreased sub-region l c . A s i m i l a r t r e n d was  found f o r t h e r e l a t i o n s h i p  between 2nd prong l e n g t h and l a t i t u d e i n Region ( F i g u r e 33) and In Regions 2, 36).  3 and 5 (1% l e v e l )  A c o n e - s c a l e with a l o n g e r 1st  1 (5%  level)  ( F i g u r e s 34 t o  prong p r o b a b l y a l s o has a  l o n g e r 2nd prong, g e n e r a l l y w i t h a r e g i o n a l c l i n a l  variation,  both prongs i n c r e a s i n g from n o r t h t o south. T a b l e s 9/a, 10/a,  11/a,  12/a  to  and 13/a  e l e v a t i o n had a weaker e f f e c t on c o n e - s c a l e  c l e a r l y show t h a t characteristics  than l a t i t u d e .  That l a t i t u d e a f f e c t e d the cone-scale  c l e a r l y shown i n the present Tusko ( 1 9 6 3 ) who  t h i s i s supported  was  by  d e s c r i b e d a s t r o n g c o r r e l a t i o n between the  width of the cones and Sziklai  study and  size  ( 1 9 6 4 ) who  c e r t a i n environmental f a c t o r s , and  by  demonstrated t h a t the width of D o u g l a s - f i r  cones appeared t o be more p l a s t i c o r g e n e t i c a l l y " l o o s e l y " cont r o l l e d than the length; of cone. and l e n g t h of 1 s t  and  2nd  Width and  l e n g t h of  cone-scale,  prongs i n c r e a s e d from n o r t h t o south  much more n o t i c e a b l y than t h e i r i n c r e a s e from low t o h i g h  eleva-  tion. The  cone-scale  measurements f o r the " c o a s t a l " r e g i o n s  appeared t o be g r e a t e r than f o r " i n t e r i o r " r e g i o n s These f i n d i n g s agree with those by Peace ( 1 9 ^ 8 ) , and  and  2.25  15).  by Tusko (19&3)  (I969).  by S z i k l a i The  (Table  average r a t i n g of b r a c t  (Table 15)  was  2.19  for "coastal"  f o r " i n t e r i o r " regions, i n d i c a t i n g that "coastal" r e -  gions have l o n g e r b r a c t s than " i n t e r i o r " ones. The  r a t i n g o f b r a c t had a c u r v i l i n e a r r e l a t i o n s h i p with  l a t i t u d e and a p o s i t i v e l i n e a r r e l a t i o n s h i p with e l e v a t i o n i n Regions 1 and  3  (Figures 2 1 ,  22,  37  and  38,  Table  l4/a).  This  i n d i c a t e s t h a t the r e l a t i v e l e n g t h of the b r a c t i n c r e a s e d  from  low t o high e l e v a t i o n and  Eleva-  from n o r t h t o south l a t i t u d e s .  t i o n appeared t o have more i n f l u e n c e than l a t i t u d e . A n a l y s i s of v a r i a n c e  (Table l4/b)  Indicates highly  signif-  i c a n t d i f f e r e n c e s among t r e e s w i t h i n provenances, among provenances w i t h i n sub-regions and among sub-regions w i t h i n No  s i g n i f i c a n t d i f f e r e n c e s were found among r e g i o n s .  regions.  The relationship between the width of cone-scale and elevation27  27  26 25 ? _E  24  a> 23 U to  3 i  « g u  *  22 Y=22 41+0 0003 X O • A X  20  Sub-region la Sub-region too Sub-region lb Sub-region Ic  O • A X A  9 20  19 -  Sub-region Sub-region Sub-region Sub-region Sub-region  3a 3b 3c 3d 3e  i  19 _L 500  _L 1000  1500  2000 2500 30O0 Elevation (feet)  3500  4000  _L 4500  Figure 13- in region I-  X  5000  L 500  lOOO  _L 1500  2000  2500 3000 Elevation (feet)  Figure 14- in region 3The relationship between the width of bract and elevation7 —  E E  6 —  Y= 4166+000057 X O •  Sub-region 7a Sub-region 7 b  4 — 500  1000  1500  2000 2500 Elevation (feet)  3000  3500  4000  4500  3500  4000  4500  -L 5000  5500  The relationship between the length of cone-scale and elevation22j21  x x  E 20| E  ft  19 18 €  Y= 18 32+0 0005 X  -a*  Sub-region la Sub-region laa Sub-region Ic Sub-region Id  17 16 15  X.  500  1000  1500  2000  Figure 16- in region I-  2500 3000 Elevation (feet)  3500  4000  4500  5000  5500  22121 E E 20  x  19  A  a * u  Y= 18 29+0 0003 X  18  Sub-region Sub-region Sub-region Sub-region Sub-region  ? 17  3a 3b 3c 3d 3e  16  151-  _L  500  _L  1000  ±  1500  2000  2500 3000 Elevation (feet)  3500  4000  4500  5000  5500  The relationship between the length of 1st prong and elevationII  -  JO E  e  O & M  - 8 o £  Y*8 0 1 + 0 0 0 0 2 X  C  O • A X  «! 7  500  1000  1500  2000  Figure 18- in region I-  2500  3000  3500  Sub-region la Sub-region laa Sub-region lb Sub-region Ic  4000  4500  5000  Elevation (feet)  The relationship between the length of 2nd prong and elevation-  2000 o  500  IOOO  eoo  Figure 19- in region 1  2500  3000  Elevation (feet)  3500  4O00  4500  5000  500  1000  1500  _L  2000  _L 2500  Elevation (feet)  3000  3500  4000  4500  The relationship between the rating of bract and elevationA-  x  a o A  Y<=2 14+0 00014 X O • A X  500  1000  1500  2000 2500 3000 Elevation (feet)  3500  Sub-region la Sub-region laa Sub-region lb Sub-region Ic 4000  4500  5000  5500  5000  5500  Figure 21- in region I 4-  E E3 o m '2-  Y«l 84+0 0002 X  r*x  Sub-region Sub-region Sub-region Sub-region Sub-region  I 0  500  Figure 22-  1000  1500  2000 2500 3000 Elevation (feet)  in region 3-  3500  4000  3a 3b 3c 3d 3e  4500  Width of Cone-scale N  ro  cw  1  —1—r  ro  (mm)  M  '  —f 3* -CD -1 CD O  TT  -4-  5'  3 to 3"  CD O  k  •5" <  31  <a'  oT  CD  (mm) ro ^  ro w  CD  03  00  ro ot  xc«0  -<  CO CO CO CO  lO  5*5  •region 1 -region 1 region 1 region 1  3  r  2  C  n  cr 0 a  u  Q.  $1  O O 3 CD I CO O Q_ CD  0)  ????  o  -.  3  Width of Cone-scale ro ro ro — ro CM  CD CD 3  O ro CD  0  2h  K  a 3  O A)  2*  la  cQ C  y  10  8  Width of Cone-scole (mm) ro ro ro  T  ro  0.  1—1—r  c  Q. CD  CD  OJ j  CD  ro 10 d> CM 1  o 6 o cu o  to •fc X  -  <H7  -  - 9*7 -  -tot-  n  Length of 2nd Prong (mm)  •? v  v  w  w  u  »  »  »  *  Length of 2nd Prong (mm)  rnTTtTTTTtTT  c  , oo  Ol Ol  O  ro  •fr  01 oo  V 6  H  c^TTTTTTTTTT  c OJ OJ  w  Ul 6 .CD  CD  <D  «s.  c-i:  at • *  o r»  f  i "83  ro  A  3 Q.  A O CO <D i5 4-  O O  O lO (Jj  I'l'l'l'oSo. no* o o °  Length of 2nd Prong (mm) cO  c  CD  A I  0)  I  do 6 I  I  ro I  1  -oi -  i  - -•ro  i r  Q.  Length of 2nd Prong (mm) A di  ro CD  OJ  0)  Ol  CD  CD  ^  oi a> I  I  01 ' CD  ro I  TT  Q  Q  c  Q. CD  5' 3  O 3 iQ Q  X  3 o  rolS a  01  -< II c/> (/>(/)(/)(/>  O  c ccrcr c ccrco* O1J cr CD £  5 CD CD  •Q <Q IU  «0  iO  p  o oo o" 5' X  ro roro ro r o  a no  IJO  The relationship between the rating of bract and latitudeY=22 843-0 02776 X' +000039 X O Sub-region 3o • Sub-region 3 b A Sub-region 3c X Sub-region 3d A Sub-region 3e 3  Y= 4 01-00007 X O Sub-region Io # Sub-region loa A Sub-region lb X Sub-region Ic A Sub-region Id  2  i -  i 38  40  42  44  46  48  Latitude (Degrees North)  Figure 37- in region I-  50  52  38  40  42  44  46  48  Latitude (Degrees North)  Figure 38- in region 3-  50  52  54  T a b l e 7.  Prov, No. 1 2  ?  4 5 6 7 8 9 10 11 12 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32  Mean v a l u e s and standard d e v i a t i o n s o f cone-scale c h a r a c t e r i s t i c s by provenances in millimeters. Wi Mean 20.43 21 01 23.98 19.71 20.80 23.33 22.46 19.57 21.27 20.16 20.31 22.87 20.09 23.02 20.91 21.30 20.85 20.20 22.28 22.13 21.83 21 78 23.01 21.15 22.35 22.51 21.88 19.66 22.77 21.81 23.33 22.85  L  Li  W  2  L3  2  R  ±SD  Mean  ±SD  Mean  ±SD  Mean  ±SD  Mean  +SD  Mean  +SD  1.55 2 38 1.57 1.66 1.96 1.22 2.07 1.42 2.55 1.84 1.89 1.58 2.34 1.84 1.38 1.61 1.74 2.14 2.15 1.92 0.87 1.60 1.89 1.95 1.49 I.67 1.79 1.62 1.27 2.34 2.31 1.93  4.59 4.71 5.32 4.62 4.65 4.99 5.01 4.62 5.05 5.45 4.99 4.53 4.39 4.85 4.?4 5.40 5.24 5.00 5.50 5.01 5.19 5.15 5.22 5.20 5.13 5.40 4.85 5.53 5.37 5.29 5.27 5.24  .54 .51 .77 .51 .57  17.22 16.79 21.08 17.33 16.16 19.09 18.79 16.70 16.49 16.51 16.74 17.98 16.88 19.74 17.19 16.72 15.82 16.67 18.63 19.46 19.41 18.77 18.91 17.31 19.13 18.66 17.18 16.48 19.65 16.54 19.35 19.46  1.34 1.75 2.35 1.47 1.98 1.33 2.10 1.55 2.09 1.55 1.3? 1.3? 1.84 2.34 1.03 1.73 1.27 5.56 1.73 2.07 1.55 1.88 1.39 1.51 1.37 1.97 1.12 1.23 2.26 1.77 2.04 1.49  7.01 7.29 8.82 6.61 6.15 7.47 6.55 6.60 7.52 7.91 6.97 7.53  1.31 0.81 0.59 0.79 0.75 0.89 1.43 1.15 1.07 0.87 1.44 0.84 1.51 1.44 1.15 1.4i 1.12 1.46 1.34 O.98 0.72 1.13 0.98 0. 82 1. ?4 1.43 1.06 0.89 1.4o 1.55 1.68 1.26  2.72 2.76 3.17 2.74 2.71 3.43 2.77 2.46 2.81 3.19 2.53 3.33 3.02 3.37 2.94 2.87 2.95 2.86 3.35 2.93 3.51 3.49 3.71 3.55 3.87 3.87 2.78 3.34 3.54 3.15 4.08 3.66  .43 .41 .80 .40 .44 .41  1.90 2.44 3.00 1.77 1.87 2.4l 2.63 1.73 2.21 2.16 2.33 1.74 2.26 2.47 2.48 2.30 2.4© 2.21 1.78 2.57 2.19 3.61 2.32 2.49 2.15 2.13 2.33 2.16 2.21 2.52 2.4l 2.29  .42 .35 .00 .65 .63 .77  •2 .67 1  .71 .71 .63 .68  •?? .46 .45 .60 .72 .65 .54 .67 .65  •?  9  .43 .63 .55 .65  •?  5  .49 .60 .59 .41  7.4^  7.58 8.11  7.75  8.42  7.94 6.60 7.29 7.63 7.90 6.83 8.65 8.97 6.79  8.44 7.76 6.46 8.02 8.31  :8? :  .34 .59 .95 .58 .50 .54 .50 .67 .69 .43 .59 .39 .66 .63 • 55 .55 .58 .38 .47 .69 .65 .56  :l  .63 .50 .51 .61 .57 .67 .50 .72 .58 .78 .64  :U .44 .49 .50 .62 .52 .56 .49 .58 .51 .66 .66  Table 7. Prov. No.  35  36  37 38 0 41 42 ft  45 46 % 48 4 9  50 51 52  53 54 55 56 57 58 59 60 61 62 64* 65  (Continued) Mean  Wi  21.94 22.11 22.11 22.93 21.88 20.98 23.30 21.87 21.17 22.22 22.05 23.67 22.24 22.43 20.23 20.47 21.86 22.43 22.75 23.30 23.75 23.98 21.89 22.51 21.17  23.18 22.95  24.59  22.87 22.72  22.41  22.06 23.63  +SD 2.3? 1.94 2.52 2.29 2.01 1.39 2.30 2.26 1.98 2.13 1.56 1.34 1.78 I.65 1.37 1.77  2.08 2.62 1.44 2.89  1.42  2.63 1.61 1.69 1.55 2.10 1.92 1.83 2.25 2.39 1.83 1.92 2.51  5.35 5.33 5.H  5.23 5.06  '.ll :U  Mean  I8.58  .68  19.02  17.04 17.96  18.04  .58  16.89  .78 .74 .76  17.51 17.59 18.97  5.18 4.87 5.99  :G .38  18.99  5.77 5.46 5.24 5.l4  :ll  5.04 5.20 5.18 4.78  .72  4.79 4.74  4.93  4.74 4.69 5.23  5.24  5.16  4.96  4.87 4.4o 4.90  5.39 5.13 4.58 4.79 5.14  .63  .98  .59 .57  19.67  I8.65  18.69  2.04 2.18 1.75  1.91  1.13 2.06 I.76 1.47 1.49 1.50  18.29  19.13 18.86  .74  17.61  .73 .45 .97 .82 .79 .64 .83  1.70  19.71  .73 .60 .61  1.95 1.80 2.40 1.42 2.24 2.22  2.13  20.06 19.61  .72  2.65 1.74  18.17 15.58 17.36  17.54 16.81 19.26 18.66 19.07 18.75 17.62 18.19 17.78 18.71  L3 Mean  +SD  1.42 2.22  8.31 7.04 6.96  7.42 7.58  7.28 7.14 5.85 7.29 6.96 8.45 7.60 7.35 8.57 7.03  8.32  1.33 2.4l  6.49 7.38 7.12 8.65 7.11 8.33 6.58 7.55 7.57 7.85 7.49 7.4l 8.27 8.00  2.07 1.85  8.29 7.77  1.88 1.46  1.60 2.16 2.01  2.04 2.00  6.82  1.55 1.13 I.89 1.10 1.4l 1.50 0.97 1.10 1.00 0.98 0.96 1.19 1.06 1.60  0.81 1.4l 0.99 1.21 1.11 1.92  0.90 1.48 1.11 1.12  1.24 1.04  0.49 0.66 0.80 0.94 0.62 I.67 0.92  4.00 3.37 3.43 3.58 3.73 3.26 3.12 2.87 3.37 3.09 4.04 3.30 3.45 3.45 2.86 3.17 3.12 3.51 3.29 3.50 3.01  3.81  3.19 3.25 3.39 3.42 3.23 3.12 3.95 3.55 2.94 3.53 3.4l  +SD .66  .53 .73 ,4o .59 .37 .29  .38 .27  .41 .54 .44 .38 .28 .41 .38 .53  :8  :il .50  .39  %  .35 ,53 .53 .27 .50 .69  R Mean 2.10 2.29 1.97 1.56 1.85 1.98 2.02 1.74  1.81  1.97 1.94 2.02 1.03 2.59 2.37 2.29  1.64 1.77 1.62 1.72 2.32 2.09  1.64  1.55 1.71 1.99 1.92 1.94 1.83 2.02 2.17 2.35 2.75  +SD .85 .69 .50 .58 .77 .74 .52 • 51 .60 .63 .63 .58  :ll .58 .66 .50 .60 .57 .59 .59 .62 .59 .67 .66 .57 .71 .47 .61 .56 .58 .52 .30  T a b l e 7. Prov. No. 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99  (Continued) Mean 20.42 24.19 23.61 23.67 22.19 22.82 23.18 24.20 24.00 23.86 22.98 22.81 23.83 24.59 25.00 23.54 23.71 23.60 2^9^ 24.87 24.17 25.39 23.74 23.96 22.35 23.91 23.33 22.74 22.12 22.87 22.39 21.50 22.64  Wl  +SD 1.84 2.23 2.27 1.12 2.32 1.48 2.00 1.49 1.69 1.99 O.96 2.36 1.82 1.84 1.82 1.53 I.67 2.48 1.62 1.02 1.22 1.31 1.62 2.07 1.91 2.60 1.12 1.19 1.32 1.04 1.73 2.06 1.83 1.70  I  23 , -97 4.62 5.13 5.21 5.30 5.18 5.15 5.26 5.41 5.14 5.24 5.65 5.23 5.59 5.21 5.07 5.29 5.21 5.49 5-. 3 7 .o4  I:  5. 5.54 5.01 81 84 69 53 88 31 5.15  5.64  .84 .77 .78 .66 .78 .77  .49  .73 .82 .59 .68 .69 .65 .76 .83  :K .72 .68  .64  •?  5  .56 .77 .71 .54 .80 .47 .5© .90 .51 .59 .39 .65  16.29 I8.96 18.19 18.19 19.09 19.71 I8.76 19.17 18.42 18.88 19.79 19.11 19.49 19.85 19.20 18.33 18.73 18.89 18.97 19.63 20.30 19.43 19.34 17.94 19.86 17.85 19.97 18.75 19.75 18.08 I8.4l 18.54 17.65 19.08  1.43 1.91 1.99 1.55 2.4l 2.65 1.58 1.49 1.42 2.40 1.36 2.06 1.85 1.31 2.10 1.58 2.23 2.04 1.64 1.16 1.25 1.4l 1.43 2.94 1.33 2.02 1.09 2.49 1.47 1.68 1.53 1.72 2.35 1.17  7.80 7.91 7.78 8.11 8.54 8.61 9.16 7.74 8.67 9.35 7.51 8.56 8.91 8.40 8.4l 8.07 8.57 9.09 8.35 8.31 8.93 8.40 8.90 8.48 8.94 7.92 10.04 9.51 9.93 10.16 10.18 10.56 8.42 9.44  0.-78 O.85 0.94 1.10 1.4l 1.45 1.59 1.94 1.35 I.25 I.07 1.34 1.37 0.79 1.10 1.47 1.31 1.84 1*31 0.73 0.82 0.93 0,90 0.99 1.56 1.24 1.81 0.84 0.90 1.46 1.29 1.55 0.47 1.67  Mean  +SD  Mean  2.86 3.52 3.31 3.25 4.oi 4.26 4.07 3.43 4.03 4.02 3.66 3.87 3.79 3.76 4.34 3.55 .96 .08 3.73 3.77 4.31 3.93  .45  2.11 I.65 1.82 1.71 1.95 2.33 1.93 1.96 1.91 2.12 2.30 2.51 2.13 2.01 2.47 1.07 2.00 2.09 1.95 2.37 2.58 2.07 2.06 1.84 2.79 2.22 1.71 2.03 2.34 2.52 2.53 2.32 2.78 2.35  l:U  4.94 3.84 4.46 4.32 4.84 4.81 4.80 4.59 3.28 4.44  .4\ .45 .55 .83 .25 .66 .67  .64  .77 .£9 .47 .55 .90 .66 .79  il .60 .54 .67 .69 .57  •il .51 .71 .80 .78 .89 .17 .76  ±SD  • ?  % 5  .75 .63 .58 .69  •. 6I9  7  .75 .50 .52  % .63 .63  :8  .35 .62 .62 .59 .32 .77 .87  .34 .35 .39 .38 .28 .60  Prov. No.  Mean  102 103 io4 105 106 107 108 109 110 111 112 113 Ii4 115 116 117 118 119 120 121 122 123 124  21.94 24.25 21.94 22.09 24.31 23.27 23.79 25.01 24.61 23.64 23.72 23.58 24.57 25.12 22,44 23.06 26.06 26.17 24.94 25.07 22.48 23.55 24.96 24.99 24.13  m  W2 Li L2 L3 B  = = » = = =  Wi  +SD 2.00 1.8? 1.94 1.28 1.90 2.28 2.04 1.62 2.07 1.32 2.12 1.72 1.73 2.28 2.19 1.45 1.46 2.10 2.58 1.72 1.06 1.97 2.32 1.70 2.08  W2 Mean ±SD 5.95 5.57 5.21 5.36 5.65 5.02 5.28 5.08 5.67 29 98 28 5. 5. 18 5. 19 5. 10 5. 24 5. 21 5. 33 5. 06 09 90 5. 13 5. 35 57 93  l\ 2:  I;  cone-scale width. b r a c t width. cone-scale length, 1 s t prong l e n g t h . 2 n d prong l e n g t h . rating of bract.  .69 .73 ? .56 .73 , 5  •I .60 7  .49 .80 .49  :U .61 .64 .7^ ti  .38 .38 .60 .68 .54 .62  Ll Mean 19.32 20.74 19.32 20.74 18.51 19.15 20.74 19.05 19.59 20.53 19.72 20.49 19.97 19.99 20.56 21.34 20.19 20.30 20.78 20.28 18.46 18.94 21.10 22.05 20.05  +SD 1.4l 2.16 1.4l 2.16 1.36 1.99 1.91 1.59 1.23 0.81 2.39  1.47 2.l4 1.65 2.11 3.00 1.49 2.30 2.08 1.62 2.44 1.37 1.81 2.47 2.27  L2 Mean +SD 8.92 10.59 9.71 10.75 9.25 9.95 8.89 8.77 8.62 8.89 7.95 8.43 8.98 8.85 8.65 9.21 9.48 9.09 8.61 8.21 8.82 9.04 8.70 8.21 7.83  0.90 1.82 1.02 0.84 1.45 1.36 1.29 1.48 1.14 1.27 1.20 1.80 1.64 1.54 1.10 1.57 1.35 i.o4 1.36 0.97 1.16 0.88 0.98 1.71 1.18  L  Mean 4.54 4.62 4.84 4.82 4.79 4.49 4.60 4.2 4.8 4.3;32 4 . 4+9 4.51 4.54 4.17 4.42 4.61 4.54 4.61 4.39 4.01 4.78 4.20 4.38 4.25 3.60  3  H  +SD  Mean  +SD  .39 .70 .66 .66 .73 .59 .90  2.89 2.5? 2.64 2.67 2.31 2.02 2.79 2.27 2.75 2.90 2.87 2.77 2.77 2.02 l.?92 2.96 2.51 2.79 2.56 2.62 1.44 2.29 2.7O 2.47 2.45  .21 .44 .34 .47 .60 .68 .36 .10 .34 .21 .25 .40 .41 .65 .62 .10 .32 .30  .73 .75 .78 .83 .57 .72 .61 .54 .52 .56 .50 .45 .77 .60 .65  •?? .44 .50 .70 .40 .60 .50  Table 8.  Region  Mean v a l u e s and standard d e v i a t i o n o f cone-soale c h a r a c t e r i s t i c s by c l i m a t i c regions i n m i l l i m e t e r s .  Cone-scale width  Bract width  Cone-scale length  1st prong length Mean  ±SD  +SD  3.6 4*. 4  .70 .82 .87 .82  3.5 3.5  lot  I'.k  .78 .91 .76  I.63 1.75 1.43 1.76 1.64  ll l:i 4.5  .69 .70 .73 .91 .80  7.6  1.76  3.2  .72  1.98 2.25  7.0 8.0  1.93 1.54  2.8 3.5  .60 .78  5 8  17.3  1.91  6.8  1.82  2.7  .68  2  16.4  1.87 1.99  7.9 8.2  1.77 1.38  2.9 3.1  .71 .58  3  +SD  Mean  +SD  2.06 2.06 1.75 2.19  5.2 5.2 5.5 5.3  .61  :?2 .65  18.6 18.2 18.7 20.3  1.91 2.4l 1.77 1.96  VA 9.9 8.6  1.42 1.83 1.68 1.50  22.2 22.9 23.8 23.2 22.9  2.34 2.43 2.00 2.11 2.02  5.0 5.2  .76 .93 .80 .76 .58  18.0 19.1 18.7 18.9 18.8  2.20 1.75 1.97 2.12 1.84  7.5 7.5 8.2 9.1 9.2  1.92 1.66 1.39 1.61 1.54  3a 3b 3e 3d 3e  22.2 22.2 23.1 23.5 24.9  1.77 2.04 2.16 2.34 2.10  5.2 5.0 5.4 5.2  .60 .71 .78 .78 .64  19.3 18.4 18.6 19.4 21.0  2.23 2.09 2.05 2.12 2.65  7.3 7.5 8.1 9.2 8.9  4  22.6  2.21  4.9  .98  18.9  2.75  5a 5b  21.0 22.8  2.00 2.39  4.9 5.3  .77 .80  17.1 18.3  6  20.1  1.76  4.6  .64  7a 7b  20.6 20.2  2.20 2.20  5.2 5.5  .72 .82  8  20.3  +SD  la laa lb lc  22.3 22.2 23.0 24.6  2a 2b 2c 2d 2e  5.4 5.1  5.0  Total No. o f prov.  Mean  Mean  Mean  2nd prong length  16.7 16.8  7.0  4.4  2.5  i  5 15  6 7 6  I 3 11 13 9 2  1  T a b l e 9/a.  R e l a t i o n s h i p between c o n e - s c a l e width and l o c a t i o n o f seed source* Elevation  Character  R2  Latitude  r  R  2  No. o f Prov. r  Region 1 Cone-scale width  0.66  0.81**  0.59  0.77**  29  Region 2 Cone-scale width  0.02  0.13NS  0.07  0.27NS  30  Region 3 Cone-scale width  0.11  0.34*  0.27  0.52**  38  Region 5 Cone-scale width  0.01  0.02NS  0.49  0.70**  13  Region 7 Cone-scale width  0.36  0.60NS  0.12  0.35NS  7  R^ and r v a l u e s i n t h e above T a b l e r e p r e s e n t t h e r e l a t i o n s h i p s between c o n e - s c a l e width and e l e v a t i o n ( P i g s . 13 and 14), and between c o n e - s c a l e width and l a t i t u d e ( P i g s . 2 3 - 2 5 ) . Only t h e s i g n i f i c a n t r e l a t i o n s h i p s were graphed. R r NS * ** 2  _ c o e f f i c i e n t o f determination, ss c o r r e l a t i o n c o e f f i c i e n t . » not s i g n i f i c a n t . = s i g n i f i c a n t a t 5# l e v e l . = s i g n i f i c a n t a t 1% l e v e l .  Table 9/b. Source  A n a l y s i s o f v a r i a n c e o f c o n e - s c a l e widths. DP  SS  MS  7  6872.0  681.7  4.06*  13  3145.0  241.9  6.28**  103  3967.0  38.5  3.73**  T/P/S/R  1701  1751.0  10.3  Error  3649  2304.0  0.6  Total  5473  33859.0  R S/R P/S/R  R = region S m sub-region P = provenance  P  16.31**  T • tree * = s i g n i f i c a n t a t 5% l e v e l . **= s i g n i f i c a n t a t 1% l e v e l .  Elevation  Character  R  Latitude R  r  2  No. o f Prov.  r  2  Region 1 B r a c t width  0.01  0.11NS  0.01  0.10NS  29  Region 2 Bract width  0.07  0.27NS  0.03  0.17NS  30  Region 3 Bract width  0.03  0.17NS  0.01  0.28NS  38  Region 5 Bract width  0.01  0.09NS  0.19  0.44NS  13  Region 7 Bract width  0.69  0.83*  0.33  0.57NS  7  R and r v a l u e s i n t h e above T a b l e r e p r e s e n t t h e r e l a t i o n s h i p s between b r a c t width and e l e v a t i o n ( F i g . 15)* and between b r a c t width and l a t i t u d e . Only t h e s i g n i f i c a n t r e l a t i o n s h i p was graphed. 2  R r NS * **  = c o e f f i c i e n t of determination, =s c o r r e l a t i o n c o e f f i c i e n t . = not s i g n i f i c a n t . = s i g n i f i c a n t a t 5% l e v e l . m s i g n i f i c a n t a t 1% l e v e l .  T a b l e 10/b.  A n a l y s i s of v a r i a n c e of b r a c t widths.  Source  DF  ;  ss  MS  F  7  121.8  16.1  1.48NS  13  141.6  10.9  3.22**  103  348.6  3.4  2.66**  T/P/S/R  1701  2161.2  1.3  8.64**  Error  36^9  536.3  0.2  Total  5473  3300.1  R S/R P/S/R  R « region S * sub-region P = provenance  _ tree * s i g n i f i c a n t a t 5% l e v e l . » « . s i g n i f i c a n t a t 1% l e v e l . =  Elevation  Character  2  r  fi  No. of Prov.  Latitude R2  r  Region 1 Cone-scale length  0.64  0.80**  0.52  0.72**  29  Region 2 Cone-scale length  0.01  0.10NS  0.03  0.18NS  30  Region 3 Cone-scale length  0.16  0.40*  0.25  0.50**  38  Region 5 Cone-scale length  0.01  0.10NS  0.41  0.64*  13  Region 7 Cone-scale length  0.02  0.15NS  0.10  0.32NS  7  R and r values In the above Table represent the relationships between cone-scale length and elevation (Figs. 16-17), and between cone-scale length and l a t i t u d e (Figs. 26-28). Only the s i g n i f i c a n t r e l a t i o n s h i p s were graphed. z  Br r NS * **  «s c o e f f i c i e n t of determination, = correlation coefficient. = not s i g n i f i c a n t . = s i g n i f i c a n t at 5% l e v e l . « s i g n i f i c a n t at 1% l e v e l .  Table 11/b.  Analysis of variance of cone-scale lengths.  Source  DF  MS  SS  F  7  4956.0  708.0  4.21*  13  2186.0  168.2  5.24**  103  3306.0  32.1  3.30**  T/P/S/R  1701  I6656.O  9.8  16.80**  Error  3649  2126.0  0.6  Total  5473  29203.0  R S/R P/S/R  R = region S a: sub-region P =s provenance  T = tree * ss s i g n i f i c a n t a t 5% l e v e l . **« s i g n i f i c a n t at 1% l e v e l .  Elevation  Character  R  2  Latitude  r  R  2  No. o f Prov.  r  Region 1 1st prong l e n g t h  0.14  0.37*  0.56  0.71**  29  Region 2 1st prong l e n g t h  0.10  0.32NS  0.41  0.64**  30  Region 3 1st prong l e n g t h  0.02  0.13NS  0.36  0.60**  38  Region 5 1st prong l e n g t h  0.02  0.12NS  0.54  0.73**  13  Region 7 1st prong l e n g t h  0.22  0.47NS  0.12  0.35NS  7  R* and r v a l u e s i n t h e above T a b l e r e p r e s e n t t h e r e l a t i o n s h i p s between 1st prong l e n g t h and e l e v a t i o n ( F i g . 18), between 1st prong l e n g t h and l a t i t u d e ( F i g s . 2 9 - 3 2 ) . Only t h e s i g n i f i c a n t r e l a t i o n s h i p s were graphed. R r NS * **  = c o e f f i c i e n t of determination, a* c o r r e l a t i o n c o e f f i c i e n t . ss n o t s i g n i f i c a n t . as s i g n i f i c a n t a t $% l e v e l . ss s i g n i f i c a n t a t \% l e v e l .  2  T a b l e 12/b.  A n a l y s i s of v a r i a n c e o f 1st prong l e n g t h s .  Source  DF  SS  MS  F  7  636.O  90.7  0.45NS  13  2600.6  200.0  7.87**  103  2621.3  25.4  4.66**  T/P/S/R  1701  9287.4  5.5  2.69**  Error  3649  7659.2  2.1  Total  5473  22804.6  R S/R P/S/R  R as r e g i o n S as sub-region P ss provenance  T as t r e e * s= s i g n i f i c a n t a t 5% l e v e l . **s= s i g n i f i c a n t a t 1% l e v e l .  R  r  2  No. o f Prov.  Latitude  Elevation Character  R  2  Region 1 2nd prong l e n g t h  0.29  0.54*  0.66  0.82**  29  Region 2 2nd prong l e n g t h  0.05  0.23NS  0.52  0.72**  30  Region 3 2nd prong l e n g t h  0.06  0.25NS  0.48  0.70**  38  Region 5 2nd prong l e n g t h  0.04  0.20NS  0.75  0.87**  13  Region 7 2nd prong l e n g t h  0.75  0.87*  0.08  0.28NS  7  R and r v a l u e s i n t h e above T a b l e r e p r e s e n t t h e r e l a t i o n s h i p s between 2nd prong l e n g t h and e l e v a t i o n ( P i g s . 19-20), and between 2nd prong l e n g t h and l a t i t u d e ( F i g s . 3 3 - 3 6 ) . Only t h e s i g n i f i c a n t r e l a t i o n s h i p s were graphed. 2  Br r NS * **  = c o e f f i c i e n t of determination, = correlation coefficient. ss not s i g n i f i c a n t . = s i g n i f i c a n t a t 5% l e v e l . ss s i g n i f i c a n t a t 1% l e v e l .  T a b l e 13/b.  A n a l y s i s o f v a r i a n c e o f 2nd prong l e n g t h s .  Source  DP  ss  MS  F  7  815.2  116.5  1.97NS  13  767.1  59.0  10.02**  103  606.2  5.9  5.76**  T/P/S/R  1701  1739.3  1.0  4.05**  Error  3649  921.0  0.3  Total  5473  4848.8  R S/R P/S/R  R ss r e g i o n S ss subr<reglon P ss provenance  1  T ss t r e e * ss s i g n i f i c a n t a t 5% l e v e l . **== s i g n i f i c a n t a t \ % l e v e l .  Elevation  Character  R  Latitude  r  2  R  2  No. o f Prov.  r  Region 1 Rating of bract  0.31  0.56**  0.29  O.54**  29  Region 2 Rating of bract  0.00  0.00NS  0.02  0.15NS  30  Region 3 Rating o f bract  0.41  0.64**  0.33  0.73**  38  Region 5 Rating o f bract  0.09  0.30NS  0.14  0.38NS  13  Region 7 Rating of bract  0.14  O.37NS  0.01  0.11NS  7.  Br and r v a l u e s I n t h e above T a b l e represent t h e r e l a t i o n s h i p s between r a t i n g o f b r a c t and e l e v a t i o n ( F i g s . 21-22), and between r a t i n g o f b r a c t and l a t i t u d e ( F i g s . 37-38). Only t h e s i g n i f i c a n t r e l a t i o n s h i p s were graphed. R* r NS * **  ss c o e f f i c i e n t o f d e t e r m i n a t i o n , ss c o r r e l a t i o n c o e f f i c i e n t . ss not s i g n i f i c a n t . = s i g n i f i c a n t a t 5% l e v e l . ss s i g n i f i c a n t a t 1% l e v e l .  Table l4/b.  A n a l y s i s of variance of r a t i n g o f bracts.  Source  DF  SS  MS  7  154.5  22.1  1.28NS  13  223.7  17.1  6.24**  103  283.9  2.8  2.95**  T/P/S/R  1701  1592.1  0.9  4.24**  Error  3649  805.0  0.2  Total  5473  3059.3  R S/R P/S/R  R =s r e g i o n S ss sub-region P ss provenance  F  T ss t r e e * ss s i g n i f i c a n t a t 5% l e v e l **=; s i g n i f i c a n t a t 1% l e v e l  T a b l e 15.  Summary o f average cone-scale c h a r a c t e r i s t i c s f o r " c o a s t a l " and " i n t e r i o r " r e g i o n s .  Regions  Character  Coastal  Cone-scale width B r a c t width  (W^)  (W ) 2  Cone-scale l e n g t h (5.^) 1st  prong l e n g t h ( L i )  2nd prong l e n g t h Rating of Bract  (L ) 2  (R)  Interior  23.14  mm  21.38  mm  5.22  mm  5.13  mm  19.00  mm  17.38  mm  8.24  mm  7.68  mm  3.84  mm  3.12  mm  2.19  rating*  2.25  rating*  * f o r r a t i n g o f b r a c t see F i g u r e 3/b page 18.  R e l a t i o n s h i p Between Thousand-Seed Weight and Cone-Scale Characteristics Thousand-seed  weight and c o n e - s c a l e c h a r a c t e r i s t i c s were  found t o v a r y g r e a t l y from provenance t o provenance and were a f f e c t e d more s t r o n g l y by l a t i t u d e than e l e v a t i o n . I t i s l o g i c a l t o examine the r e l a t o n s h i p s between 1000seed weigth and c o n e - s c a l e c h a r a c t e r i s t i c s i n each r e g i o n t o determine whether o r not they agree with t h e f i n d i n g s shown p r e viously. Thousand-seed Regions 1,  weight was a s s o c i a t e d w i t h e l e v a t i o n i n  2 and 3 and with l a t i t u d e i n Regions 1,  Width o f c o n e - s c a l e was  7.  c o r r e l a t e d w i t h e l e v a t i o n i n Regions  3 and 7 and w i t h l a t i t u d e i n Regions 1, earling.  3» 5 and  1,  3 and 5» as mentioned  F i g u r e 39 t o 42 show t h a t 1000-seed weigth was  sig-  n i f i c a n t l y c o r r e l a t e d w i t h width o f c o n e - s c a l e i n Regions 1 and 3 (1% l e v e l ) , and i n Regions 2 and 5 (5$ l e v e l ) . seed weight was  significantly correlated  Although 1000-  (5$ l e v e l ) w i t h the width  of c o n e - s c a l e i n Region 2 ( F i g u r e 40), the r e l a t i o n s h i p was weak» the c o e f f i c i e n t o f d e t e r m i n a t i o n (Table 16), as compared w i t h 0.66 I n Region 1,  R, 2  was only 0.18,  which I m p l i e d t h a t i n Region 2  the i n c r e a s i n g 1000-seed weight was due more t o environmental o r o t h e r u n c o n t r o l l e d f a c t o r s than t o the width o f c o n e - s c a l e . The 1000-seed weight was r e l a t e d t o t h e l e n g t h o f cone-scale ( F i g u r e s 43 t o 45) m Regions 1,  3 (1# l e v e l ) and i n Region 5  (5% l e v e l ) , which agree w i t h the p r e v i o u s f i n d i n g s .  Ther was  a  c l e a r t r e n d t h a t seed weight was d i r e c t l y c o r r e l a t e d w i t h the s c a l e s i z e , the l a r g e r c o n e - s c a l e u s u a l l y h a v i n g h e a v i e r seed. S q u i l l a c e (1957) r e p o r t e d t h a t average weight o f seed per cone was  directly  p o r r e l a t e d with cone l e n g t h and average cone-scale western white p i n e . s i z e was  size i n  E L i a s o n and H e i t (1940) found t h a t cone  r e l a t e d t o seed s i z e i n Scots p i n e .  P e r r y and  Coover  (1933) r e p o r t e d t h a t l a r g e r cones g e n e r a l l y y i e l d e d l a r g e r seeds i n p i t c h p i n e , and t h a t many small and medium-size cones cont a i n e d not o n l y more seed, but b e t t e r f i l l e d seed than the l a r g e r cones I n s h o r t l e a f p i n e .  They a l s o i n d i c a t e d t h a t t h e r e was  no  r e l a t i o n s h i p between cone s i z e and v e r t i c a l p o s i t i o n of cone i n p i t c h pine. Seed weight i s g e n e r a l l y r e l a t e d t o cone-scale s i z e which i n yurn depends on cone s i z e . seed weight and  Although  the r e l a t i o n s h i p between  cone weight i s not i n c l u d e d i n t h i s study, i t  c o u l d be assumed t h a t l a r g e r and h e a v i e r cones have h e a v i e r than s m a l l e r ones.  In t h i s connection, Simak and  seed  Gustaffsson  (1954) r e p o r t e d t h a t i n Scots p i n e , i n c r e a s i n g cone s i z e and  cone  weigth Increased not only seed p r o d u c t i o n , but a l s o the average seed weight per cone.  Simak (i960) f u r t h e r noted t h a t , i n Scots  p i n e , the number and the average s i z e of seed per cone i n c r e a s e d with l a r g e r cone weight.  He  concluded  appeared t o be determined mainly  t h a t these r e l a t i o n s h i p s  by the t r e e ' s genotype, but  were a l s o s t r o n g l y m o d i f i e d by environmental  factors.  F i g u r e s 46 t o 53 show t h a t 1000-seed weight became h e a v i e r as 1st  and 2nd prongs became l o n g e r .  Furthermore, l a r g e r cone-  s c a l e s were a p p a r e n t l y accompalned by l o n g e r prongs, but r e l a t i o n s h i p was Cone-scales  not observed  i n Region  this  7.  with r e l a t i v e l y shorter bracts g e n e r a l l y y i e l d e d  h e a v i e r seed In Regions 1 and  3 ( F i g u r e s 5^ and  55).  The r e l a t i o n s h i p s between 1000-seed weight and c o n e - s c a l e c h a r a c t e r i s t i c s e x i s t e d i n a l l r e g i o n s except Region 7 16).  (Table  The l a c k o f r e l a t i o n s h i p i n Region 7 c o u l d perhaps be ex-  p l a i n e d by Inadequate sampling.  1000-seed weight (grams)  c  I  CD  I  (0  5  I  I  I  =  I  ro 5  5  1000-seed weight (grams) 5  —  I I I  —  ro 5  5  5  5S  CD cO  Ss. 2 fol  ™rol ?  u.r rol  OJ OJ OJ OJ OJ CD O . <"> O " O  1000-seed weight (grams) c B - ^ o » t o o  =  i i j -i T i  ro  i ii  c o i o O CD  o  rol Of  CD if>  rol  CD  1000-seed weight (grams)  r o 5 S 5 i 5 >  ro ro  Ss ro  Ss Ol?rof  3 ro  = rol CD O J |  rol  • 0 co co c c cr a  coco co CO CO c c c c c era cr cr cr -  (ft 6)  cr o  Cft CD  •  rororororo » a. o o"  o  =  F 5 o i ^ u i o >  -  L9  -  18-  18-  16-  O • A X  16-  Sub-region la Sub-rtgion laa Sub-region lb Sub-region Ic  Sub-region Sub-region Sub-region Sub-region Sub-region  E 2-4  12  2a 2b 2c 2d 2e  12  10-  O  o o  10  8-  6 \  I  I  l  I  I  l  l  l  I  I  I  I  X^l  I I I I I I I I I I I I I I I 2 0 2-2 2-4 2-6 2-8 3-0 3-2 3-4 3-6 3-8 4 0 4-2 4-4 4-6 4-8 50 5-2 5 4 56  *20 2-2 2-4 2-6 2 « S-O 3-2 3-4 3-6 3« 4-0 4-2 4-4 4-6 4-8 50 5-2 5-4 Length of 2nd Prong (mm)  Length of 2nd Prong ( m m )  Figure 50- in region I16 —  O • A X  14  Figure 51- in region 2-  Sub-region 3a Sub-region 3b Sub-region 3c Sub-region 3d  O 9  Sub-region 5a Sub-region 5b  14  - 12  12  10 o o o  O  o O  6 —  T4 I  I I I I I 20 2-2 2-4 2-6 2fl 3-0 3-2 3-4 3-6 3 8 40 4-2 4 4 Length of 2nd Prong (mm)  Figure 52- in region 3-  1 1481 4-6  1 5-0 5-2 5-4  8  6-  _I__L  J  I  1  I 5 0 5-2 I 54 2-0 2-2 24 2-6 2-8 3-0 3-2 3-4 3-6 3-8 4 0 4-2 4-4 4-6 4-8  K  Length of 2nd Prong  Figure 53- in region 5-  (mm)  The relationship between 1000-seed weight and rating of bract-  Table 1 6 .  C o r r e l a t i o n between 1 0 0 0 - s e e d weight and cone-scale c h a r a c t e r i s t i c s . 1000-seed weight  Character R Wl  60 «  W2  i  0.28 0.49 O.36  0.53**  Wi W2  0.18  L2  0.46  0.42* 0.09NS O.34NS  R  0.02  L  I»2 3 R h  N  §  3 cn fl  wi w2  S?  Li L  3  0.81** 0.08NS 0.84**  0.66  0.01  fl  3  r  2  2  R^ >A  W<  fl  w2  0.70  0.01 0.12 0.39  0.68** 0.63**  0.38  0.62** 0.29NS  0.08  0.4l 0.19 0.33  0.64** 0.43** 0.49**  0.57*  0.32  0.01  R^  :8 0.24  0.63* 0.66*  Wi  0.04  fl 2 0 Ll  0.01  60  « i  L L  R  2 3  38  0.58**  0.24  L  W  30  0.13NS  0.10NS O.56*  o-  29  0.71** 0.60**  0.31  S? 2 « Lo  No. Of Prov.  0  13  0.48NS 0.20NS  0.17NS 0.06NS 0.26NS 0.21NS 0.19NS  0.03 0.07 0.05  o.o4  7  R* and r v a l u e s i n t h e above T a b l e r e p r e s e n t the r e l a t i o n s h i p 1 0 0 0 - s e e d weight and c o n e - s c a l e c h a r a c t e r i s t i c s from P i g s . 3 9 t o 55» Only t h e s i g n i f i c a n t r e l a t i o n s h i p s were graphed. Wi W2 Li Lg L3 R  = = = = = =  c o n e - s c a l e width width o f b r a c t cone-scale length 1st prong l e n g t h 2nd prong l e n g t h r a t i n g o f bract  R* r  NS *  40*  c o e f f i c i e n t of determination, correlation of coefficient, not s i g n i f i c a n t , significant at % l e v e l , s i g n i f i c a n t a t 1% l e v e l .  Seed g e r m i n a t i o n  test  R e g r e s s i o n a n a l y s i s was  c a r r i e d out between germination  percent and l a t i t u d e , l o n g i t u d e and e l e v a t i o n of seed f i r s t l y on sub-regions and r e g i o n s .  No s i g n i f i c a n t  source  relation-  s h i p s c o u l d be e s t a b l i s h e d , so the r e g r e s s i o n a n a l y s i s then based on 114 out o f 124 provenances i n t h i s  was  experiment.  An a n a l y s i s o f the d a t a 36 days a f t e r sowing i n d i c a t e d t h a t g e r m i n a t i o n percent was (Figure 5 6 ) .  However, i t was  termination, R ,  i s 0.14,  2  s i g n i f i c a n t l y a f f e c t e d by l a t i t u d e shown t h a t the c o e f f i c i e n t o f de-  which i m p l i e s t h a t o n l y 14 per cent  o f the v a r i a t i o n i n germination percent can be e x p l a i n e d by latitude.  The  e f f e c t of t h i s f a c t o r disappeared 50 days a f t e r  sowing the seeds.  There was no s i g n i f i c a n t r e l a t i o n s h i p between  germination percent and e i t h e r l a t i t u d e o r l o n g i t u d e . The lowest germination percent a f t e r sowing) was  (Table 17)  of 11.6  (36  o b t a i n e d from seeds c o l l e c t e d i n C a s t l e Rock,  Washington (Prov. No.70, l a t . 46°19*, l o n g . 1 2 2 ° 5 2 ' ) and highest, 61.9  percent was  l a t . 5 1 ° 4 4 ' , long.  c o l l e c t e d from T a t l a , B.C.  the  (Prov.  No.7,  124°44*).  The lowest germination percent a f t e r sowing) was  days  (Table 17)  found i n Sook, B.C.  l o n g . 1 2 3 G 4 4 ) and the h i g h e s t , 8 5 . 6  of 4 1 . 8  (92  days  (Prov. No.33, l a t . 48°24', p e r c e n t , seeds were c o l -  l e c t e d from P i n e Grove, Oregon, (Prov. No.86, l a t . 4 5 ° 0 6 ' ,  long.  121°23'). The range i n the germination percent w i t h i n each sub-region i s presented i n T a b l e 18.  As can be observed t h e r e was  a  con-  s i d e r a b l e d i f f e r e n c e among provenances w i t h i n a s u b - r e g i o n j  f o r example, a t end o f the germination p e r i o d , sub-region 3d had t h e h i g h e s t range between minimum and maximum germination p e r c e n t , i . e . , from 44.5 t o 84.3#, while sub-region 7b had a r e l a t i v e l y s m a l l e r range from 73.7  t o 78.6#.  These two  sub-  r e g i o n s a r e l o c a t e d i n Washington S t a t e . In some sub-regions, f o r example, samples from 2b, 2d, 4, 6 and 7a  (Table 18)  w i t h the h i g h e s t and  g e r m i n a t i o n percent remain the h i g h e s t and lowest the e n t i r e duration of the  sub-regions lowest  throughout  experiment.  The d a t a showed t h a t t h e provenances with the lowest h i g h e s t germination percent i n sub-regions l a a , 2b, 5b,  6 and 7a appeared  and  3«» 4,  5a,  t o have the most c o n s i s t e n t germination 18.  p a t t e r n s when compared with the o t h e r sub-regions i n T a b l e The d i f f e r e n c e s i n the germination percent o f seeds t h e f o l l o w i n g f o u r provenances,  from  n o r t h e r n (Prov. N o . l , Stoner,  B.C.,  l a t . 53°37't l o n g . 1 2 2 ° 4 o ' ) , easthern (Prov. No.11,  B.C.,  l a t . 5 1 ° 2 3 ' , l o n g . 177 ©0*), westhern (Prov. No.12, Jeune  Landing, B.C.,  Golden,  G  l a t . 5 0 ° 2 7 ' , l o n g . 12702?') and southern  (Prov.  No.122, C o v e l , C a l i f o r n i a , l a t . 3 9 ° 4 8 ' , l o n g . 1 2 2 ° 5 6 ' ) a r e shown i n T a b l e 19.  The a n a l y s i s o f v a r i a n c e showed s t a t i s t i c a l l y  s i g n i f i c a n t d i f f e r e n c e s a t 1% l e v e l among these f o u r provenances 36 days a f t e r sowing.  The g r e a t e r v a r i a t i o n of g e r m i n a t i o n p e r -  cent a t 36 days a f t e r sowing found between n o r t h e r n and  southern  provenances when compared w i t h westhern and easthern provenances i s c l e a r l y shown i n F i g u r e s 57 t o 60.  These r e s u l t s can be  con-  f i r m e d i n F i g u r e 56 t h a t l a t i t u d i n a l f a c t o r a f f e c t s the germinat i o n percent more than l o n g i t u d i n a l f a c t o r .  No  significant  d i f f e r e n c e s among these f o u r provenances were observed  after  50 days from sowing. The l n t r a - p r o v e n a n c e v a r i a t i o n i n g e r m i n a t i o n percent can a l s o be observed from these f o u r provenances. t r e e s w i t h i n provenances  The range among  ( F i g u r e s 57 t o 6 0 , T a b l e 19) was g r e a t e r  i n westhern and n o r t h e r n provenances than i n e a s t h e r n and s o u t h ern provenances.  T h i s may i n d i c a t e t h a t D o u g l a s - f i r moved g r a d -  u a l l y northwards a f t e r t h e i c e age.  Stronger s e l e c t i o n pressure  i n the i n t e r i o r e l i m i n a t e d t h e i n d i v i d u a l s not adapted t o t h e new c o n d i t i o n s and reduced t h e v a r i a b i l i t y w i t h i n p o p u l a t i o n (Figure 6 0 ) . variation  C o a s t a l provenances e x h i b i t a much wider range o f  (Figure 59).  Figure 56- The correlation between average germination percent (36 days after sowing) and latitude in Douglas-fir-  Cumulative germination percent of 12 trees within provenance  36  50  72  92  36  50  Age in Doys  72  Figure 57- Provenance No-1, Stoner, B-C-  Figure 58- Provenance No- 122 , Covelo, California'9 7 4  80  2,11 12 6 8,10  c §60  3  aco  5  o |£4 0  10,12,  •  o  36  72  50  92  Age in Days  92  Age in Days  Figure 59- Provenance Nol2,Jeun« Landing, B-C-  -L 36  ±  ±  50  72  Age in Days  Figure 60- Provenance No- II, Golden, B-C-  T a b l e 17.  Germination percent of 114 D o u g l a s - f i r  Germination percent days a f t e r Prov. No.  2i  SI  1. 2. 4. 5. 7. 8. 9. 10. 11. 12.  33.8 52.8  41.7 68.8  33.2  45.8 67.4 50.0 50.2 61.8 53.3 49.8 50.0 54.5 29.8 32.4 47.8 48.1  13. 14. 15. 16. 17.  18.  20. 21. 22. 23. 24. 25. 26. 27.  28.  29. 31. 32. )3. 34.  35. 36. 37. 38. 9. 0. 4l. 42. 43. 44. 45. 46.  47. 48. 49. 50.  19.4  61.9  30.8  28.0 48.4 44.2 44.8 47.6 40.0 21.8  18.5  31.6  31.0 4l.l  25.0 38.3 38.I 27.3 60.0 30.4 51.8 48.9 39.9 44.0 33.6 22.6 33.7  31.3 34.6 20.1 17.9  26.9 12.2 26.5  45.0 32.22 35.3 24.7 48.0 21.9 51.0  42.0 49.7  32.3  46.9  42.3 47.6 43.6  34.7 68.5 36.8 55.4 62.2 49.4 49.6 45.5 27.0 39.0 39.3  2 i  22  53.6  57.8  81.8 48.7 53.9 70.5 65.7 65.7 72.8  64.3  68.3  64.2  74.3 44.7 57.8 61.6 68.3  60.9 64.6 62.7 65.8 63.4  82.8  63.O 72.0 78.6 74.4 71.8 70.6 39.2 66.2  63.9  45.4  72.6  24.3  56.9  19.0  59.9  30.8  37.8  38.1 55.5  48.1 48.1 48.9 48.9  38.7 61.5 38.4 63.O 47.8 54.0  sowing  61.5  68.8  69.5 -.4 77.4 li 76.7  64__ 64.6 68.2 79.2 57.2 72.2 67.7 73.3  81.6  48.7 54.1 75.0 66.0 66.1 72.8 66.0 70.0  64.2  74.3 44.7  59.4  61.6 69.8 62.2 65.5  62.7 67.7 65.' 84.0 66.9 72.6 78.6 75.9 73.4 71.3 4l 66.2 63.9  72.6 65.I  56.9  68.8 61.3  70.2 77.4  77.2 65.6 71.6 79.8 60.4 73.7 70.1 74.8  provenances,  Prov. N O .  51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 9 2. 91. 9j>. 94. 95. 95* 96.  2i  50  22  22  36.6 39.0 35.6 17.0 56.3 29.5 22.2 31.4 17.1  41.7  68.5 63.3 59.7 58.7 72.3 68.6 67.6  41.7  43.1  70.2 63.3 62.9 58.7 73.3 68.6 68.5 64.1 44.5  27.5  j*4.9 4 4.1 41.5 36.2 4o.6 38.9 43.9 27.6 32.3 11.6 19.8 31.3 34.5 33.2 35.1 32.9 32.2 53.4 30.7 37.1 27.5 16.5 33.4 l4.8 22.9 25.6 20.1 35.3 25.4 31.5 32.7 31.9 29.8 26.8 26.8 34.3  43.7 38.6 27.1 59.7 43.2 36.6  26.2 43.5 46.8 51.1 55.2 50.0 52.2 62.7 54.8 33.1 42.7 19.3 32.8 41.7  4l.3 39.5 4Q.0 44.4 53.6 71.6 46.3 62.4 42.0 30.7 45.7 33.7 30.1 57.9 31.7 52.4 4l.4 45.3 42.1 46. >.4 50.0  43.5 47.8  63.O 63.O  73.0 62.8 72.6 64.2 73.5 78.0 73.7 47.6 69.2 53.8 59.7  67.O  62.2 69.2 70.7 70.1 78.9 83.2 68.5 81.3 60.8 58.8 76.5 64.9 47.6 85.6 56.1 70.9 68.6 62.7 70.7 63.7 71.9 75.2 73.0  63.O  73.8 63.9 73.5 66.8 75.0 79.5 73.7 50.6 69.2 54.8 60.0 67.7 63.7 72.3 70.7 84.3 78.9 83.2 70.5 81.3 61.8 60.1 78.4 66.8 48.1 85.6 56.3 72.2 69.5 62.7 72.8 64.5 72.0 75.2 73.0  Prov. No.  &  97. 99.  50.5 36.8 33.5  100. 101. 102. 103. 104. 105. 106. 107.  108. 109. 110. 112.  113*.  114. 115. 116. 117.  118. 119. 121. 122.  36.9 27.9  18.0 41.1 45.0 20.7 26.7 22.8  29. 66.4 58.8 42.0 58.2 53.9 50.3 56.3  57.0 66.0 42.9  2 i  92.  84.1  84.5  55.1 81.3 75.8 75.0  55.1 81.3  81.9  76.4  78.0 66.7 68.8 61.2  81.9  76.2  75.0  76.4 78.9 67.4  25.6  48.0  74.3  57.3  68.8 61.9 58.6  16.2 36.2 59.8 32.4 27.7 25.9 19.2 27.4 25.5  4?.8 64.5 52.7 46.0 4l.3 36.2 43.2 44.1  30.4  58.5 66.6 79.4 73.2 64.9 70.9 60.7 70.2 62.4  59.1 66.6 79.7 74.4 66.7 70.9 60.7 71.0 62.7  16.7  35.0 32.8  76.2  18.  Region la laa lb lc 2a 2b 2c 2d 2e 3a  3b  3o  3d  3e  Minimum And Maximum Germination Percent 36, 50, 72 And 92 Days A f t e r Sowing w i t h i n A C l i m a t i c Region  Total No. o f Prov.  5 4  4 12  4 7 6  Prov. No. 21 71  82 109  33  22.6  r 68  27.5  3  87 114  3  22  *36  70 84  7  11  36 da 7a a f t e r sowing Min. Prov. Max. Mean No. 38.1 25s0 23 31.7 56.8 31.8 19.8 55 I6.5 26.1 96 34.3 16.7 25.9 104 41.1  40  2  113  4  2  14  5a  5  47.6  20.1 36.2  13 50 67 91 115  38.3  20  11.6 14.8 16.2  62 101 112  44.7 36.9 21.6  40.0  2  17.0  12.2  49.7  43.9 32.7  39.8 36.2 32.5  Prov. No.  24 37  82  109  33  54 68  50 days a f t e r Min. Prov. No. 32 34.7 30.8 55 39.7 99 32.8 106 27.0 21.1  33.1  sowina Max.  Mean  45.5  42.3  59.7 58.8 66.0  4o.8  45.1 45.4  13 50 67 102 115  30.0 54.0 54.8 53.9  50.0 4i.7 44.1 44.4  29 25  49.4  47.9  28.4  87 114  31.7 49.8  39.5  46.9  19.8  20 40 70 85 113  30.4  101 112  44.5  45.7 37.5  52.8  46.4  14  54.5  2  68.6  61.6  59.8  4i.l 60.0  47.0  30.0 29.7  28.2  19.0  19.4  30.1  62  64.5  68.5  51.5  58,2  57.1  42.8  40.4  8  16  18.5  47  78  7  61.9 53.4  33.2 33.8  15 47  26.8  45.1  38.4  7 78  67.4  21.9  6  2  4  19.4  1  33.8  26.6  4  32.3  1  4i.7  37.0  7a 7b  4  18 66  31.0  10 43  48.4 51.0  46.3  34.9  17 28  47.8 62.2  48  10  61.8 68.0  51.9  5b  3  38.9  71.6  55.6  62.6  Region  72 days a f t e r sowing Prov. No.  24  Min.  92 days a f t e r sowing  Prov. No.  Max.  Mean  Prov. NO.  Min.  Prov. No.  Max.  Mean  36  67.4 64.3 72.2 67.2  21 71  65.5 60.0 60.1 58.6  36 99 104  72.6 73.3  76.4  68.6 65.5 72.5 67.9  63.4 59.7 58.8 57.3  99 104  72.6 72.3 81.9 76.4  33  54 68 87 ll4  39.2 58.7 47.6 56.1 66.6  12 50 83 102 115  68.3 73.3 76.5 75.8 79.4  60.9 65.3 60.1 66.9 74.6  33  54 68 87 Ii4  4i.8 58.7 50.6 56.3 66.6  31 50 83 102 115  73.4 74.8 78.4 76.2 79.7  62.5 66.6 67.4 67.7 75.0  60.9 56.9 43.1 47.6 58.5  29  60.0 69.1 64.2 67.6 64.2  20 38 59 85 113  29  25 61 86 112  74.4 82.8 73.0 85.6 69.8  62.2 56.9  44.5 48.1 59.1  76 86 112  75.9 84.0 84.3  2  20 38 39 85 113  70.4  66.9 70.2 66.2 68.2 64.6  4  2  14  74.3  2  81.2  77.9  14  74.3  2  81.6  78.6  5a 5b  5  8  47  44.7 57.2  7 78  70.5 83.2  58.5 73.8  47  44.7 60.4  7 78  75.0 83.2  59.8 74.9  6  2  4  48.7  1  53.6  51.8  4  48.7  1  57.8  53.3  7a 7b  4  17 48  61.6 72.1  28  10  72.8 78.6  67.2 76.2  48  17  61.6 73.7  28  10  72.8 78.6  67.6 77.3  la laa lb lG  5 4 12  2a 2b 2c 2d 2e  4 7  3a  3  3b 3c 3d 3e  i  Total No. o f Prov.  6 7 3 11 13  6  3  71  82  109  55  82  109  55  81.9  85.7  T a b l e 19.  V a r i a t i o n o f germination percent from f o u r d i f f e r e n t provenances throughout f o u r observed s t a g e s . (The v a l u e presented i n t h e T a b l e I s percent %)  days after sowing  1  Northern Prov. N o . l Stoner.B.C. Lat. 53°37' Long.l22°4o»  36 50 72 92  l4.3 23.2 39.3 41.1  23.2 28.6 4i.l 50.0  4l.l 44.7 51.8 60.8  Eastern Prov. No.11 Golden,B.C. L a t . 51°23» Long.ll7°00'  36 50 72 92  42.9 46.4 58.9 60.7  44.7 50.0 64.3 64.3  Western Prov. No.12 Jeune Landing.B.C Lat. 50°27' Long.l27°27'  36 50 72 92  67.9 69.7 91.1 94.8  Southern Prov. No.122 Covelo,Calif. Lat. 39°48' Long.l22°56'  36 50 72 92  12.5 28.6 42.9 42.9  Location  T r e e No. 2  3  4  5  6  7  10  11  12  A.V.  39.3 67.9 81.9 85.7  28.6 42.9 53-6 55.4  75.0 75.0 78.6 78.6  7.2 9.0 32.2 33.4  48.2 53.6 64.3 64.3  28.6 32.2 42.9 42.9  28.6 42.9 53.6 54.1  28.6 30.4 44.7 46.5  44.7 50.0 58.9 62.4  33.8 4l.7 53-6 57.8  39.3 48.3 62.5 69.7  39.3 39.3 53.6 53.6  39.3 51.8 69.7 69.7  42.9 48.2 58.9 58.9  48.3 50.0 60.0 60.0  53.6 54.7 57.2 57.2  53-6 70.8 71.5 75.0  62.5 69.7 73.3 76.8  29.0 4l.4 69.7 69.7  35.0 64.3 76.8 76.8  44.2 53.3 64.7 66.0  48.2 49.6 73*3 75.0  12.5 25.0 46.5 46.5  55-4 69.7 83.9 84.2  30.4 30.4 37-5 37.5  44.7 44.7 48.3 64.3  75-0 82.2 87.5 87.5  14.3 16.1 60.7 60.7  71.4 75.0 91.1 92.9  26.8 28.6 60.7 60.7  39.3 48.2 71.1 75.0  53-4 59.1 67.9 69.7  44.8 49.8 68.3 70.7  30.4 56.4 75.0 75.0  30.4 53.6 67.9 67.9  1.8 25.0 66.1 66.1  50.0 57.2 69.? 69.7  17.9 25.0 46.4 48.2  14.3 28.6 53.6 53.6  28.6 58.7 75.0 75.0  39.3 60.7 61.4 61.4  4l.4 67.9 80.4 80.4  25.0 4l.l 60.4 64.3  l4.3 26.8 50.0 50.0  25.5 44.1 62.4 62.7  8  9  SUMMARY AND CONCLUSIONS Geographic v a r i a t i o n and r e l a t i o n s h i p between  1000-seed  weights and c o n e - s c a l e morphology and t h e r e l a t i o n s h i p between these f a c t o r s and g e r m i n a t i o n percent have been r e p o r t e d f o r D o u g l a s - f i r from w i t h i n i t s n a t u r a l range.  One hundred and  twenty f o u r provenances r e p r e s e n t i n g e i g h t r e g i o n s from B r i t i s h Columbia t o C a l i f o r n i a  ( l a t . 38°50' t o 5 3 ° 3 7 » ,  long. l ^ O O * t o  1 2 7 ° 2 7 ) were c o l l e c t e d i n 1966 and 1968 by IUFRO, S e c t i o n 22. f  Sach r e g i o n was d i v i d e d i n t o s e v e r a l sub-regions a c c o r d i n g t o geographic f a c t o r s and s o i l and c l i m a t i c  conditions.  The s t u d i e s o f 1000-seed weights and c o n e - s c a l e morphology were based on a l l provenances, w h i l e g e r m i n a t i o n t e s t s were based on o n l y 114. Seeds e x t r a c t e d from cone samples were then i d e n t i f i e d as " f i l l e d " o r "empty" u s i n g X-ray f l u o r o s c o p y and s e p a r a t e d by hand.  Seed l o t s were weighed w i t h a balance r e a d i n g t o 10  grams, and average 1000-seed w e i g h t y was  computed f o r each  provenance. The c o n e - s c a l e c h a r a c t e r i s t i c s o f c o n e - s c a l e widths and l e n g t h s , b r a c t widths and 1st and 2nd prong l e n g t h s were measured in millimeters. R e g r e s s i o n a n a l y s e s between 1000-seed weights and cones c a l e c h a r a c t e r i s t i c s , and t h e l a t i t u d e and e l e v a t i o n o f seed sources were c a r r i e d out f o r each r e g i o n . Seed g e r m i n a t i o n f o r each provenance was t e s t e d under nursery conditions.  R e g r e s s i o n a n a l y s e s showed no r e l a t i o n s h i p  between g e r m i n a t i o n percent and e l e v a t i o n , l a t i t u d e and l o n g i t u d e  w i t h i n each r e g i o n , and  f u r t h e r regression analyses  were t h e r e -  f o r e based on combined provenances. Some r e s u l t s a r e summarized as f o l l o w s t 1.  Thousand-Seed Weight a.  Thousand-seed weight v a r i e d among t r e e s , provenances, sub-regions and  b.  regions.  Thousand-seed weight had a c l i n a l v a r i a t i o n i n c r e a s i n g from low t o h i g h e l e v a t i o n , observed mainly i n c o a s t a l r e g i o n s , and a c l i n a l Increase both c o a s t a l and  from n o r t h t o south i n  i n t e r i o r regions.  L a t i t u d e appeared  t o a f f e c t seed-weight more than e l e v a t i o n . 2.  Cone-Scale C h a r a c t e r i s t i c s a.  Cone-scale c h a r a c t e r i s t i c s d i f f e r e d s i g n i f i c a n t l y among t r e e s w i t h i n provenances, among provenances w i t h i n r e g i o n s and among sub-regions. widths and l e n g t h s had  However, o n l y  sub-  cone-scale  s i g n i f i c a n t v a r i a n c e s among r e -  gions. b.  Cone-scale widths and l e n g t h s were s i g n i f i c a n t l y r e l a t e d t o e l e v a t i o n i n o n l y two three  c.  d.  e.  s i g n i f i c a n t l y r e l a t e d to elevation  r e g i o n s , and t o l a t i t u d e i n f o u r  regions.  A s i g n i f i c a n t r e l a t i o n s h i p between b r a c t width  and  e l e v a t i o n was  width  was f.  significantly related to elevation  r e g i o n , and t o l a t i t u d e i n f o u r r e g i o n s .  2nd prong l e n g t h was i n two  to latitudes i n  regions.  1st prong l e n g t h was i n one  r e g i o n s and  found i n o n l y one  region.  not r e l a t e d t o l a t i t u d e i n any  A s i g n i f i c a n t number of cone-scale  Bract  region. and b r a c t measurements  had  a d e f i n i t e c l i n a l variation increasing  high elevation gj.  from low t o  and from n o r t h t o south i n some r e g i o n s ,  The average v a l u e s o f cone-scale c h a r a c t e r i s t i c s i n "coastal  M  r e g i o n s were g r e a t e r than i n " i n t e r i o r " r e -  gions. 3>_  R e l a t i o n s h i p Between Thousand-Seed Weight and Cone-Scale Morphology Thousand-seed weight was s t r o n g l y scale but  correlated  with cone-  s i z e , l a r g e r cone-scales producing h e a v i e r seeds i n a l l  one r e g i o n .  T h i s r e g i o n may have had an i n s u f f i c i e n t  number o f provenances sampled. 4.  Germination Percent - Seed Source R e l a t i o n s h i p Germination percent was s i g n i f i c a n t l y a f f e c t e d by l a t i t u d e some 36 days a f t e r sowing, but t h i s e f f e c t seemed t o d i s a p p e a r a t about 50 days a f t e r sowing.  E l e v a t i o n and  l o n g i t u d e appeared not t o a f f e c t g e r m i n a t i o n p e r c e n t . In Region 2 , g e n e r a l l y , t h e cone-scale s i z e s were not r e l a t e d t o t h e i r seed source. investigation  These r e s u l t s i n d i c a t e t h a t  i s needed t o prove t h a t  further  cone-scale s i z e i s i n -  f l u e n c e d by e i t h e r environmental e f f e c t s such as moisture and r e l a t i v e humidity, o r g e n e t i c e f f e c t s , o r a combination o f t h e two. C l i n a l v a r i a t i o n o f cone-scale c h a r a c t e r i s t i c s d i d not e x i s t universally,  but o n l y i n c e r t a i n r e g i o n s .  a l l tree-to-tree  T h i s and l a r g e r  v a r i a t i o n w i t h i n provenances i n d i c a t e  over-  that,  i n f u t u r e seed c o l l e c t i o n s f o r provenance t e s t s and a r t i f i c i a l regeneration, careful attention variation.  should be g i v e n t o intra-provenance  1957. B e t t e r h a n d l i n g o f a s c a r c e commodity. B.C. Lumberman 41 ( 7 ) 1 32-36  A l l e n , G. S.  1958. F a c t o r s a f f e c t i n g t h e v i a b i l i t y and g e r m i n a t i o n b e h a v i o r o f c o n i f e r o u s seed. I I I . Commerical p r o c e s s i n g and treatments s i m i l a r t o p r o c e s s i n g , Pseudotsuga m e n z i e s i i (Mirb.) Franco, and o t h e r s p e c i e s . F o r . Chron. 3^ (3)* 266-298. 1960. F a c t o r s a f f e c t i n g t h e v i a b i l i t y and g e r m i n a t i o n b e h a v i o r o f 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 l e a i l (Mirb.) Franco. F o r . Chron. 36 ( l ) i 18-29. 1961. T e s t i n g D o u g l a s - f i r seed f o r provenance. P r o c . I n t . Seed T e s t . A s s . 26 (3)1 388-403 Andersson, E.  1965« Cone and seed s t u d i e s i n Norway spruce ( P i c e a a b l e s (L.) K a r s t ) Stud. f o r . suec. Skogshogsk, Stockn, No. 23, 286 pp.  Baker, K. F.  1957. The U.C. System f o r P r o c u c i n g Healthy Container-Growth P l a n t s . U n i v e r s i t y o f C a l i f . , D i v i s i o n of A f r l c u l t u r e S c i . , A g r i c u l t u r a l Experiment S t a t i o n - E x t e n s i o n S e r v i c e . 331 pp.  Baldwin, H. I .  19^2. F o r e s t t r e e seed o f t h e n o r t h temperate r e g i o n with s p e c i a l r e f e r e n c e t o North America. Chronlce B o t a n i c a Co. 240 pp.  Barner, H.  1966. C i r c u l a r l e t t e r o f A p r i l 20, 1966 o f Worki n g Group on Procurement o f Seed f o r Provenance Research. I.U.F.R.O. S e c t i o n 22. 10 p.  Buck, J . M., R. S. Adams, J . Cone, M. T. Comkle, W. J . L l b b y , C. J . Eden and M. J . K n i g h t . 1970. California Tree Seed Zones. C a l i f . Region F o r e s t S e r v i c e , U..S. Dept. o f A g r l . San F r a n c i s c o , C a l i f . . 5 pp. Ching, K. K. and D. Bever. i960. Provenance Study o f Douglasf i r i n the P a c i f i c Northwest Region. 1. Nursery Performance. S i l v a e G e n e t i c a 9 . 11-17. E l i a s o n , D. J . and C. E. H e l t . 1940. The s i z e o f S c o t s p i n e cones as r e l a t e d t o seed s i z e and y i e l d . J . F o r . 381 65-66 EL-lakany, M. H. and 0. S z i k l a i . 1971. Intraspecific Variation In N u c l e a r C h a r a c t e r i s t i c s o f D o u g l a s - f i r . Paper presented a t t h e Working Group Meeting a t Gainesv i l l e , F l o r i d a . Mar. 14-20, 1971. 12 pp.  Hegg, E.. A.  1954. 770.  Haddock, P. G.  1962. D o u g l a s - f i r i n Canada 1 Seed c o l l e c t i o n zones based on geographic v a r i a t i o n of p o p u l a t i o n as i n f l u e n c e d by c l i m a t e . U.B.C. F a c u l t y o f F o r e s t r y . Mimeographed. 17 p.  How  t o open pond pine cones. J . F o r . 52»  and 0, S z i k l a i . 1966. Seed c o l l e c t i o n zone f o r D o u g l a s - f i r In Canada. S i x t h World F o r e s t r y Congress. Madrid. 6 CFM/E/C. T. 1?15. 16 pp. 1955. I n f l u e n c e o f the s i z e of Scots pine seed on t h e i r q u a l i t y . For. A b s t r . . l6» 255*  I l j i n , A. I .  I r g e n s - M o l l e r , H. 1958. Genotypic v a r i a t i o n i n the time of c e s s a t i o n o f h e i g h t growth i n D o u g l a s - f i r . For. S c i . . 4» 325-330. _,  1962. Genotypic v a r i a t i o n i n p h o t o p e r i o d r e sponse of Douglas f i r s e e d l i n g s . For. S c i . 81 360-362.  K o c h a r j , N. T.  1952. C o r r e l a t i o n of f i e l d g e r m i n a t i o n of Plnus s y l v e s t r l s seed with s i z e o f cone. For. Abstr. 13• 384.  L a r s e n , G. S.  1937. Genetics i n s i l v i c u l t u r e . T r a n s l a t e d by H. L. Anderson. 1956. O l i v e r and Boyd Edinburgh. 224 pp.  1967. S t a n d a r d i z a t i o n o f methods f o r provenance r e s e a r c h and t e s t i n g . I.U.F.R.O. XIV Kongress. Munchen 1967. S e c t i o n 22. I I I . 672-718.  L i n e s , R.  Mirov, N. T. ,  1936. A note on g e r m i n a t i o n methods f o r c o n i f e r o u s s p e c i e s . J . For. 34, 719-723. 1946. V i a b i l i t y o f p i n e seed a f t e r prolonged c o l d s t o r a g e . J . F o r . 44« 193-195.  , J * W. D u f f i e l d and A. R. L i d d i c o e t . 1952. Altitudinal r a c e s of Plnus ponderosa — a 12 year progress r e p o r t . J . For. 50. 825-831. Peace, T. R.  1948. The v a r i a t i o n of Douglas f i r i n i t s n a t i v e h a b i t a t (Pseudotsuga t a x i f o l i a B r i t . syn. Pseudotsuga d o u g l a s l i C a r r . JT F o r e s t r y 221 45-61.  P e r r y , G. S., and C. A. Coover. 1933. J . For. 31« 19-25.  Seed source and  quality.  Robinson, B. A. 1963* V a r i a t i o n i n seed c h a r a c t e r i s t i c s o f D o u g l a s - f i r i n B j . l t i s h Columbia. B.S.F. Theses, F a c u l t y o f F o r e s t r y , U.B.C. Vancouver, 68 pp. Roche, L .  1966. Spruce provenance r e s e a r c h i n B r i t i s h Columbia. Tenth Meeting o f the Committee on F o r e s t T r e e Breeding i n Canada, Univ. o f B.C., Vancouver B.C. Sept. 7-10. P r o g r e s s Report.  Rogmeder, E.  19*>2. Germination w i t h Ulmus montana (With.) F o r . A b s t r . 164.  Simak, M.  i960. I n f l u e n c e o f cone s i z e on the seed produced (Plnus s l l v e s t r l s L . ) . Medd S k o g s f o r s k n , . 4 9 1 16 pp. ,  1967. Seed weight o f l a r c h from d i f f e r e n t provenances ( L a r l x deoldua M i l l . ) . F o r e s t a l i a S u e c l a , Nr. 57» S t u d l a F o r . Suec. Skogshogskolan, Stockn, 25 pp. and A. G u s t a f s s o n . 1954. F r o b e s k a f f e n h e t e n hos modertrad och ymper av t a l l . (Seed p r o p e r t i e s i n mother t r e e s and g r a f t s o f S c o t s p i n e ) . Medd Skogsforskn., Stockholm 44t 73 pp.  •  S q u i l l a c e , A. A. 1957. V a r i a t i o n i n cone p r o p e r t i e s , seed weight i n western white p i n e when p o l l i n a t i o n i s c o n t r o l l e d . School o f F o r e s t r y . Montana S t a t e U n i v e r s i t y . M i s s o u l a , Montana. B u l l e t i n No. 5» 16 PP* S t e b b l n s , G. J . 1957* V a r i a t i o n and e v o l u t i o n i n p l a n t s . U n i v e r s i t y P r e s s , N.Y. 643 pp.  Columbia  Stone, E. C.  I957;, The e f f e c t o f seed s t o r a g e on s e e d l i n g s u r v i v a l o f sugar p i n e . J . F o r . 55* 816-820.  Sweet, G.  1965. Provenance D i f f e r e n c e s i n P a c i f i c Coast D o u g l a s - F i r . S i l v a e G e n e t i c a 141 46-56.  Sziklai,  .  B. 0.  1964. V a r i a t i o n and i n h e r i t a n c e o f some p h y s i o l o g i c a l and m o r p h o l o g i c a l t r a i t s i n Pseudotsuga m e n z i e s i i (Mirb.) Franco v a r . m e n z i e s i i . Ph.D. T h e s i s . F a c u l t y o f F o r e s t r y , U.B.C. 136 pp.  • .  I967. Variation of Douglas-fir i n i t s native h a b i t a t . Paper presented a t I n t e r n a t i o n a l S c i e n t i f i c Conference, Z v o l e n , C z e c h o s l o v a k i a , 11 pp.  ,  T o o l e , E. H.,  1969. P r e l i m i n a r y notes on the v a r i a t i o n i n cone and seed morphology o f D o u g l a s - f i r (Pseudotsuga m e n z i e s i i (Mirb.) Franco. Second World C o n s u l a t l o n on F o r e s t T r e e B r e e d i n g . F O - F T B - 6 9 - 6 / 9 , 11 pp. et a l . 1956. P h y s i o l o g y o f seed g e r m i n a t i o n . Annual Review o f P l a n t P h y s i o l . 7» 299-324.  Tourney, J , W., and G. P. K o r s t i a n . 1948. Seeding and P l a n t i n g i n t h e P r a c t i c s o f F o r e s t r y . John Wiley and Sons Inc., N.Y. Tusko, F. F.  1963. A study o f v a r i a b i l i t y i n c e r t a i n Douglasf 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. T h e s i s , Dept. o f B i o l , and Bot., U.B.C. 173 PP»  Van Deusen, J . L., and L . D. Beagle. 1970. Some Cone and Seed C h a r a c t e r i s t i c s o f B l a c k H i l l s Ponderosa P i n e . Rocky Mountain F o r e s t and Range Exp. S t a . U.S.D.A. F o r . Serv. Note RM - 179. 7 pp. Western F o r e s t Seed C o u n c i l . 1966. Tree seed zone map f o r Washington and Oregon. Meeting o f W.F.T.S.C. A p r i l , 1966. P o r t l a n d , Oregon. Willis,  C. P., and J . V. Hoffmann, 1915. A study o f D o u g l a s - f i r seed. Soc. Amer. F o r e s t e r s P r o c . 10t l 4 l - l 6 4 .  W i l l e t t , S. V. H. I 9 6 3 . V a r i a t i o n I n cone c h a r a c t e r i s t i c s o f D o u g l a s - f i r i n B r i t i s h Columbia. 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.B.C. 65 pp.  

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