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UBC Theses and Dissertations

Morphological and anatomical variation of picea in southwestern British Columbia Scagel, Robert Kevin 1984

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MORPHOLOGICAL AND ANATOMICAL VARIATION  OF PICEA IN SOUTHWESTERN  BRITISH COLUMBIA. By ROBERT KEVIN B.Sc,  The U n i v e r s i t y  SCAGEL  of B r i t i s h  Columbia  1977  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE  in THE FACULTY OF GRADUATE (Department  We a c c e p t t h i s to  STUDIES  of B o t a n y )  thesis  the required  as c o n f o r m i n g standard  THE UNIVERSITY OF BRITISH COLUMBIA October  1984  (c) Robert K e v i n S c a g e l , 1984  >E-6  In p r e s e n t i n g  this thesis  r e q u i r e m e n t s f o r an B r i t i s h Columbia,  it  freely available  for  understood for  that  Library  s h a l l make  for reference  and  study.  I  f o r extensive copying of  h i s or  be  her  g r a n t e d by  s h a l l not  the  be  of  further this  this  Botany  The U n i v e r s i t y o f B r i t i s h 1956 Main Mall V a n c o u v e r , Canada V6T 1Y3 Date  (3/81)  thesis  a l l o w e d w i t h o u t my  Robert K. S c a g e l .  of  Columbia  my  It is  permission.  Department  thesis  head o f  representatives.  copying or p u b l i c a t i o n  f i n a n c i a l gain  University  the  s c h o l a r l y p u r p o s e s may by  the  the  I agree that  permission  department or  f u l f i l m e n t of  advanced degree a t  of  agree that  in partial  written  i i  ABSTRACT.  Relations  between  representative putative using  an  hybrids  several  cones,  of  southwestern  and  twigs.  variables patterns the  the  of  c a n o p y were d e t e c t e d  the  suggesting  necessarily  individual.  especially  a  different  intra-individual  the  with  intra-taxonomic  variation  inter-taxonomic  variation.  attributed  to  the  intra-population 25  percent  hypothesized with  the  emerged was  In  than  evident  to  respect  the  age  taxa  be  was  the  data.  as  the  e x t r e m e s of  position  the  tree in  were  or  Picea to  not  of context,  the  than  than  variation  variation.  the  to  variation  smaller  of  these  extrinsic  taxa  larger  data  of  Similar  of  inter-individual  Inter-taxonomic  of  variables  variation  pattern  Inter-individual  polarity  only  of  the  intra-individual  polarity  with  hypothesized  an  had  variable.  analyses.  environment  shown t o  hypothesized  taxonomic  actual  two  was  variation.  larger  the  respect  variation.  the  variation  intra-individual of  that  on  trees.  regardless  function  Further,  that  than d e v e l o p m e n t a l l y  variation  that  measured  intra-individual  intra-individual  intra-individual  were examined  were s e l e c t e d  suggested  640  be and  variables  exceeded  variation  i n t e r - and  Columbia  variables that  to  P_j_ s i t c h e n s i s  British  were m e a s u r e d on  of  and  were examined w i t h m u l t i v a r i a t e  species, not  36  variation  hypothesized  anatomical  were more g e n e t i c a l l y  Patterns  trees  enqelmannii  Such r e l a t i v e  These v a r i a b l e s  is  in  inter-individual  selected  in  Picea  m o r p h o l o g i c a l and  needles,  variation.  individual  was  not  was  only  Further,  the  co-incident  What t a x o n o m i c a continuum  structure of  iii variation  over a l a r g e  environmental Picea  i n southwestern  or  practically  is  not p o s s i b l e  hybridization  The with  followed  trees  of a s i n g l e , l a r g e  to regional  suggests  attributable  with  utilization programmes.  and l o c a l  of P i c e a  appropriate  taxon.  scales  Local  of g e o g r a p h i c  only  s c a l e s of patterns  of  geographic  of v a r i a t i o n i n n u r s e r y  areas.  grown  trees  v a r i a t i o n was n o t  of v a r i a t i o n i n n a t u r a l l y grown  of o r i g i n .  This  variability  of  i n t e r - i n d i v i d u a l v a r i a t i o n i s probably a t an e x t r e m e l y  from t h e s e c o n c l u s i o n s  respect  taxa  of v a r i a t i o n t o  i n 16 s e p a r a t e  to processes operating  Implications discussed  that  of two  n o t be  polymorphic  variation.  the patterns  from t h e same a r e a  relations  regional  of t h e p a t t e r n  with  may  o f i n t e r - i n d i v i d u a l v a r i a t i o n was  were n o t c o - i n c i d e n t  co-incident  Columbia  recognition  by i n t r o g r e s s i o n o r t o t h e  and e n v i r o n m e n t a l  respect  that  large  From t h e e v i d e n c e a v a i l a b l e h e r e , i t  c o r r e l a t e d with  relation  reflecting  to a t t r i b u t e the p a t t e r n  pattern  geographic variation  British  feasible.  differentiation  partially  area  v a r i a t i o n , suggesting  of  The  geographic  to the systematics  i n f o r e s t r y and t r e e  local  a r e p r e s e n t e d and of P i c e a  and t h e  improvement  scale.  iv  CONTENTS. ABSTRACT  i i  CONTENTS  iv  L I S T OF TABLES  v i i  L I S T OF FIGURES.  x i i  ACKNOWLEDGEMENTS  xv  PREFACE  1  I.  3  INTRODUCTION  II.  MATERIALS  AND METHODS  15  1. I n t r o d u c t i o n  15  2. C h a r a c t e r s 2.1 C h a r a c t e r s and v a r i a b l e s s e l e c t e d 2.2 V a r i a b l e measurement r e p e a t a b i l i t y 2.3 V a r i a b l e v a r i a t i o n between i n d i v i d u a l s  17 18 22 26  •  3. Samples a n d s a m p l i n g 3. 1 Study area 3.1.1 C l i m a t e 3.1.2 E d a p h i c e n v i r o n m e n t 3.1.3 A s s o c i a t e d v e g e t a t i o n 3.1.4 G e o l o g i c a l , v e g e t a t i o n , a n d c l i m a t i c frrstory. .. 3.1.5 H i s t o r y o f P i c e a i n w e s t e r n N o r t h A m e r i c a 3.2 T r a n s - i n d i v i d u a l c i r c u m s c r i p t i o n of samples 3.2.1 P o p u l a t i o n c i r c u m s c r i p t i o n o f s a m p l e s 3.2.2 T a x o n o m i c c i r c u m s c r i p t i o n o f s a m p l e s . 3.2.3 L o c a l g e o g r a p h i c c i r c u m s c r i p t i o n o f s a m p l e s . ... 3.3 S e l e c t i o n o f samples  31 31 35 37 39 40 42 44 44 47 48 49  4. A n a l y s e s 4.1 M a t h e m a t i c a l n o t a t i o n and f o r m u l a t i o n 4.2 S t a t i s t i c a l techniques 4.2.1 C o n f o r m i t y t o d i s t r i b u t i o n a l a s s u m p t i o n s 4.2". 2 Sample s i z e 4.2.3 D i s c r e t e v a r i a b l e s 4.2.4 U n i v a r i a t e a n a l y s e s 4.2.5 M u l t i v a r i a t e a n a l y s e s 4.3 C o m p u t a t i o n and t e s t s o f s i g n i f i c a n c e  53 54 54 56 58 61 64 65 73  III.  INTRA-INDIVIDUAL VARIATION  74  1. I n t r o d u c t i o n  74  2. M a t e r i a l s a n d methods 2.1 Materials 2.2 Analyses  83 83 85  V  IV.  3. R e s u l t s 3.1 Adventitious versus primary whorl branches 3.2 Primary versus secondary whorl branches 3.3 Year t o y e a r v a r i a t i o n 3.4 Inter-positional variation 3.5 I n t r a - p o s i t i o n a l v a r i a t i o n i n an i n t e r - i n d i v i d u a l context  89 89 90 92 94 105  4. D i s c u s s i o n 4.1 I m p l i c a t i o n s f o r sampling 4.2 Crown form m o r p h o g e n e s i s 4.3 Further research  107 110 113 116  and i n t e r p r e t a t i o n  INTER-INDIVIDUAL VARIATION: TAXONOMIC CIRCUMSCRIPTION.  120  1. I n t r o d u c t i o n  120  2. M a t e r i a l s and methods 2.1 Materials 2.2 Analyses  123 123 124  3. R e s u l t s 3.1 P o p u l a t i o n s of s t a n d a r d s 3.1.1 P_;_ e n q e l m a n n i i and P^ s i t c h e n s i s 3.1.2 P_j_ e n q e l m a n n i i , P. s i t c h e n s i s, and P_^_ g l a u c a . .. 3.2 Individual s t a n d a r d s and p u t a t i v e t a x o n o m i c representatives 3.2.1 P^ e n q e l m a n n i i and P_j_ s i t c h e n s i s 3.2.2 P\_ e n q e l m a n n i i , P. s i t c h e n s i s, a n d P_^_ g l a u c a . .. 3.3 Artifical h y b r i d s i n the c o n t e x t of p o p u l a t i o n s of s t a n d a r d s . . •. 3.4 A r t i f i c i a l and p u t a t i v e h y b r i d s 3.5 I n d i v i d u a l s t a n d a r d s , p u t a t i v e s , and " h y b r i d s " . ..  127 127 127 130  4. D i s c u s s i o n 4.1 Intra-population variation 4.2 Natural hybrids 4.3 I n t e r - i n d i v i d u a l v a r i a t i o n i n an context  144 145 147  133 133 134 136 139 140  inter-individual 150  V. INTER-INDIVIDUAL VARIATION: RELATIONSHIPS OF PATTERNS OF VARIATION  1 52  1. I n t r o d u c t i o n  152  2. M a t e r i a l s and methods 2.1 Materials 2.2 Analyses 3. R e s u l t s 3.1 Geographic v a r i a t i o n • 3.1.1 N a t u r a l v a r i a t i o n 3.1.2 N u r s e r y v e r s u s n a t u r a l v a r i a t i o n 3.2 Local geographic v a r i a t i o n  156 156 157 162 162 162 171 173  4. D i s c u s s i o n  .  180  vi  4.1 4.2 4.3 VI.  I n f e r e n c e s from e n v i r o n m e n t a l and g e o g r a p h i c correlation I n f e r e n c e s from v a r i a b l e s u i t e i n t e r c o r r e l a t i o n . . H y b r i d i z a t i o n or d i f f e r e n t i a t i o n i n a polymorphic taxon?  UNIFYING DISCUSSION  190 196  1. The n a t u r e o f P i c e a i n s o u t h w e s t e r n B r i t i s h Columbia 2. R e f l e c t i o n s on t h e taxonomy o f P i c e a 3. S o u r c e s o f v a r i a t i o n i n p o p u l a t i o n s t u d i e s o f conifers. 4. Ontogeny and p h y l o g e n y - t h e i n t e r f a c e VII.  181 186  EPILOGUE: IMPLICATIONS AND  APPLICATIONS FOR FORESTRY.  1. I m p l i c a t i o n s f o r t r e e improvement and s i l v i c u l t u r e . . 2. I m p l i c a t i o n s f o r P i c e a f o r e s t r y 3. S u g g e s t i o n s f o r i n d e p e n d e n t s t u d i e s and c o l l e c t i o n s .  197 200 205 208 213 214 219 223  BIBLIOGRAPHY  230  APPENDICES  274  Appendix suites  I.  Description  Appendix I I . L o c a t i o n t r e e s and p o p u l a t i o n s  o f v a r i a b l e s and v a r i a b l e 274  and d i s p o s i t i o n o f i n d i v i d u a l of t r e e s  Appendix I I I . Complete t a b l e s a b b r e v i a t e d i n body o f t e x t  281  o f ANOVAs and PCAs 293  vii  L I S T OF TABLES Table  1.  Characters  used  in previous  studies  21  T a b l e 2. A v e r a g e v a r i a b l e and v a r i a b l e s u i t e measurement r e p e a t a b i l i t y , i n t e r - i n d i v i d u a l v a r i a b i l i t y , and inter-population variability  24  T a b l e 3. Inter-individual v a r i a t i o n , repeatability estimates, reported f o r morphological characters in other c o n i f e r tree species 30 T a b l e 4. Summary c f some a n n u a l a v e r a g e c l i m a t i c r e p o r t e d f o r t h e s t u d y a r e a and a d j a c e n t a r e a s Table  5 Distribution  T a b l e 6. zones  of samples w i t h  Distribution  variables 36  respect to moisture  of samples w i t h  respect to  regime. 39  biogeoclimatic 41  T a b l e 7. D i s t r i b u t i o n occurrences.  o f s a m p l e s a s p o p u l a t i o n s and s i n g l e  Table  of samples  8.  Distribution  46  T a b l e 9. Summary o f s a m p l e s w i t h various variable suites  into  geographic  areas.  50  c o m p l e t e measurements f o r 53  T a b l e 10. Sample s i z e e s t i m a t e s f o r i n t r a - i n d i v i d u a l (n) and i n t e r - i n d i v i d u a l v a r i a t i o n ( t ) T a b l e 11. Summary o f H o t e l l i n g ' s on v a r i a b l e s t a t e f o r d i s c r e t e p u l v i n u s pubescence  .  variation 60  T f o r twig morphology based (presence/ absence) v a r i a b l e : 63 2  T a b l e 12. ANOVA m o d e l s u s e d i n a s s e s s i n g v a r i o u s a s p e c t s o f intra-individual variation  87  T a b l e 13. ANOVA m o d e l s u s e d i n a s s e s s i n g t h e c o n t r i b u t i o n o f i n t r a - i n d i v i d u a l v a r i a t i o n i n the context of i n t e r - i n d i v i d u a l variation 88  vi i i  T a b l e 14. ANOVAs o f i n d i v i d u a l v a r i a b l e s c o m p a r i n g and w h o r l p r i m a r y b r a n c h e s o f engelmanni i T a b l e 15. M u l t i v a r i a t e a p p o r t i o n m e n t v a r i a b l e s u i t e s due t o d i f f e r e n c e s branches T a b l e 16. ANOVAs c o m p a r i n g y e a r v a r i a b l e s i n P\_ e n g e l m a n n i i  adventitious 90  of v a r i a t i o n f o r s e p a r a t e between o r d e r s o f 91  t o year  v a r i a t i o n of of 93  T a b l e 17. M u l t i v a r i a t e a p p o r t i o n m e n t v a r i a b l e s u i t e s due t o d i f f e r e n c e s branches  of v a r i a t i o n f o r s e p a r a t e between p o s i t i o n s o f 94  T a b l e 18. M a h a l a n o b i s ' D d i s t a n c e s between a p i c a l - a n d basal-most branches and cones of e n g e l m a n n i i and P. s i t c h e n s i s 2  T a b l e 19. A v e r a g e M a h a l a n o b i s ' s t r e e s i n T a b l e 18  D  2  d i s t a n c e s between  95  individual 98  T a b l e 20. M u l t i v a r i a t e a p p o r t i o n m e n t of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s o f s e p a r a t e t r e e s due t o w h o r l p o s i t i o n s . 100 T a b l e 21. C o r r e l a t i o n s amongst component components from PCAs i n T a b l e 20  correlations  for first 101  T a b l e 22. M u l t i v a r i a t e a p p o r t i o n m e n t of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between i n d i v i d u a l t r e e s or t a x a e m p h a s i z i n g i n t r a - i n d i v i d u a l v a r i a t i o n i n an i n t e r - i n d i v i d u a l context 106 T a b l e 23. M u l t i v a r i a t e a p p o r t i o n m e n t of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between t a x a , p o p u l a t i o n s , and i n d i v i d u a l s o f s t a n d a r d s o f P^ s i t c h e n s i s a n d P. e n g e l m a n n i i 128  T a b l e 24. M u l t i v a r i a t e a p p o r t i o n m e n t of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between t a x a , p o p u l a t i o n s , and i n d i v i d u a l s o f s t a n d a r d s o f P_;_ e n g e l m a n n i i , P. g l a u c a , and P. s i t c h e n s i s 131  IX  T a b l e 25. M u l t i v a r i a t e apportionment of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between t a x a and i n d i v i d u a l s t a n d a r d s and p u t a t i v e s o f e n g e l m a n n i i and P. s i t c h e n s i s 133  T a b l e 26. M u l t i v a r i a t e apportionment of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between t a x a and i n d i v i d u a l s t a n d a r d s and p u t a t i v e s of engelmanni i , P. g l a u c a , and P_;_ s i t c h e n s i s 136  T a b l e 27. PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r a r t i f i c i a l h y b r i d s and p o p u l a t i o n s of s t a n d a r d s of P^ e n g e l m a n n i i and P. s i t c h e n s i s T a b l e 28. M u l t i v a r i a t e apportionment of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between a r t i f i c i a l and p u t a t i v e h y b r i d s of P^ e n g e l m a n n i i and P_j_ s i t c h e n s i s . . 139 T a b l e 29. M u l t i v a r i a t e apportionment of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o e i t h e r d i f f e r e n c e s between t a x a or i n d i v i d u a l s o f P_^ e n g e l m a n n i i , P. s i t c h e n s i s , and p u t a t i v e hybrids 142 T a b l e 30. r values linear regression 2  for individual variables g i v e n i n MODEL 4  from  multiple 164  T a b l e 31. M u l t i v a r i a t e apportionment of v a r i a t i o n f o r separate v a r i a b l e s u i t e s b a s e d on p r e d i c t e d and r e s i d u a l v a l u e s from m u l t i p l e l i n e a r r e g r e s s i o n (MODEL 4) 165 T a b l e 32. Correlations residual values  among PCAs o f o r i g i n a l ,  predicted,  T a b l e 33. I n t e r c o r r e l a t i o n s of v a r i a b l e s u i t e a n a l y s e s PCAs of r e s u l t s of r e g r e s s i o n a n a l y s i s  and 168  from 169  T a b l e 34. M u l t i v a r i a t e r e l a t i o n s h i p of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s f o r n u r s e r y and n a t u r a l l y grown m a t e r i a l s t o geographic v a r i a t i o n 171 T a b l e 35. M u l t i v a r i a t e r e l a t i o n s h i p of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s f o r separate geographic areas r e l a t e d to the environment 173  T a b l e 36. R a n k i n g o f v a r i a b l e s redundancy a n a l y s i s  on a d i s p e r s i o n  T a b l e 37. PCAs  C o e f f i c i e n t s and v a r i a b l e  Table  Correlation  38.  c r i t e r i o n and 224  v a l u e s used  in various 226  amongst PCAs g i v e n  i n Table  XXXVIII. 227  T a b l e 39. primary  ANOVAs and PCAs o f s e p a r a t e v a r i a b l e s u i t e s and s e c o n d a r y b r a n c h e s o f P^ s i t c h e n s i s  f o r whorl 293  T a b l e 40. ANOVAs a n d PCAs o f s e p a r a t e v a r i a b l e s u i t e s o f w h o r l b r a n c h p o s i t i o n s a n d m a t u r e a n d immature P^ e n g e l m a n n i i and P. s i t c h e n s i s 294 T a b l e 41. PCAs of s e p a r a t e v a r i a b l e s u i t e s o f w h o r l b r a n c h p o s i t i o n s o f i n d i v i d u a l t r e e s o f P_j_ e n g e l m a n n i i and P. s i t c h e n s i s a n d %SS(mva) f o r e a c h i n d i v i d u a l t r e e . .. T a b l e 42. PCAs o f s e p a r a t e v a r i a t i o n i n the context %SS(mva)  295  v a r i a b l e s u i t e s of i n t r a - i n d i v i d u a l o f i n t e r - i n d i v i d u a l v a r i a t i o n and 296  T a b l e 43. ANOVAs and PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r sampled p o p u l a t i o n s o f s t a n d a r d s o f P^ e n g e l m a n n i i and P. s i t c h e n s i s  297  T a b l e 44. ANOVAs a n d PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r s a m p l e d p o p u l a t i o n s o f s t a d a r d s o f P_;_ e n g e l m a n n i i , P. g l a u c a , and P. s i t c h e n s i s 299 T a b l e 45. ANOVAs and PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r s a m p l e d i n d i v i d u a l s t a n d a r d s a n d p u t a t i v e s o f Pj_ e n g e l m a n n i i and P. s i t c h e n s i s 301 T a b l e 46. ANOVAs a n d PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r sampled i n d i v i d u a l s t a n d a r d s a n d p u t a t i v e s o f P_^_ e n g e l m a n n i i , P. g l a u c a , and P_;_ s i t c h e n s i s 303 T a b l e 47. PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r a r t i f i c i a l h y b r i d s and p o p u l a t i o n s o f s t a n d a r d s o f P^ e n g e l m a n n i i and P. s i t c h e n s i s 305  xi  T a b l e 48. ANOVAs and PCAs o f s e p a r a t e v a r i a b l e s s u i t e s f o r a l l i n d i v i d u a l s o f a r t i f i c i a l h y b r i d s and p u t a t i v e h y b r i d s of P. e n g e l m a n n i i and P^ s i t c h e n s i s 306  T a b l e 49. ANOVAs and PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r a l l i n d i v i d u a l s o f Pj_ e n g e l m a n n i i , P. s i t c h e n s i s , a n d p u t a t i v e hybrids. 307 T a b l e 50. ANOVAs a n d PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r s t a n d a r d s , p u t a t i v e s , and " h y b r i d s " o f P^ e n g e l m a n n i i and P. s i t c h e n s i s b a s e d on c a l c u l a t e d p r e d i c t e d and r e s i d u a l v a r i a t i o n from m u l t i p l e l i n e a r r e g r e s s i o n w i t h 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 309 T a b l e 51. PCAs o f s e p a r a t e v a r i a b l e P. s i t c h e n s i s c o m p a r i n g n a t u r a l l y grown i n t h e n u r s e r y T a b l e 52. PCAs o f s e p a r a t e variable suites  s u i t e s of standards of o c c u r r i n g t r e e s and t r e e s 311  geographic  areas  f o r separate 312  xii  L I S T OF  FIGURES  Figure  1.  Illustrations  Figure  2.  Summary of v a r i a b l e v a r i a t i o n  individuals  without  of v a r i a b l e s u s e d .  referance  25  based  to taxonomic  on a l l circumscription. 27  F i g u r e 3. Illustration l e a f anatomy Figure  4.  of range  of v a r i a t i o n  Figure  6.  28  Maps o f l o c a t i o n of samples  F i g u r e 5. Elevational, latitudinal, d i s t r i b u t i o n o f samples Location  encountered for  and  study a r e a .  ...  32  and l o n g i t u d i n a l 37  of g e o g r a p h i c a r e a s c i r c u m s c r i b i n g  samples. 49  F i g u r e 7. S c h e m a t i c r e p r e s e n t a t i o n o f s o u r c e s of i n t r a - i n d i v i d u a l v a r i a t i o n a s s o c i a t e d with branch architecture in e n q e l m a n n i i and sitchensis.  52  F i g u r e 8. S c h e m a t i c r e p r e s e n t a t i o n o f i n d i v i d u a l t r e e s of P. enqelmann i i and P^ s i t c h e n s i s from w h i c h i n t r a - i n d i v i d u a l sampling conducted 84 F i g u r e 9. O r d i n a t i o n s of f i r s t two components of PCAs of s e p a r a t e v a r i a b l e s u i t e s of w h o r l p r i m a r y and s e c o n d a r y b r a n c h e s of P_;_ s i t c h e n s i s  91  F i g u r e 10. O r d i n a t i o n s o f f i r s t two s e p a r a t e v a r i a b l e s u i t e s of w h o r l P. e n g e l m a n n i i and P_;_ s i t c h e n s i s  97  F i g u r e 11. anatomy  components o f PCAs o f b r a n c h p o s i t i o n s from  E x a m p l e s of i n t r a - i n d i v i d u a l v a r i a t i o n f o r some t r e e s shown i n F i g u r e 8  of  leaf 102  F i g u r e 12. S c a t t e r s of means and s t a n d a r d d e v i a t i o n s o f s c o r e s of f i r s t components of PCAs f o r i n d i v i d u a l b r a n c h e s o f i n d i v i d u a l t r e e s a g a i n s t whorl branch p o s i t i o n s . ...... 103  F i g u r e 13. O r d i n a t i o n s o f f i r s t two components of PCAs of i n d i v i d u a l t r e e s c o m p a r i n g p o s i t i o n of c o n e s  105  F i g u r e 14. O r d i n a t i o n s of f i r s t two components o f PCAs of separate v a r i a b l e s u i t e s f o r i n t r a - i n d i v i d u a l variatio-n i n t h e c o n t e x t of i n t e r - i n d i v i d u a l v a r i a t i o n 108 F i g u r e 15. Summary p a r t i o n i n g o f . s o u r c e s variation.  of  intra-individual 109  F i g u r e 16. O r d i n a t i o n s of f i r s t two components o f PCAs of s e p a r a t e v a r i a b l e s u i t e s f o r p o p u l a t i o n s of s t a n d a r d s of P. e n g e l m a n n i i and P_^_ s i t c h e n s i s  129  F i g u r e 17. O r d i n a t i o n s of f i r s t two components of PCAs of s e p a r a t e v a r i a b l e s u i t e s f o r p o p u l a t i o n s of s t a n d a r d s o f P. e n g e l m a n n i i , P. g l a u c a , and P_^ s i t c h e n s i s  132  F i g u r e 18. O r d i n a t i o n s o f f i r s t two components of PCAs of s e p a r a t e v a r i a b l e s u i t e s f o r a l l s t a n d a r d s and p u t a t i v e s of P. e n g e l m a n n i i and P^ s i t c h e n s i s 135 F i g u r e 19. O r d i n a t i o n s o f f i r s t two components o f PCAs of s e p a r a t e v a r i a b l e s u i t e s f o r a l l s t a n d a r d s and p u t a t i v e s of P. e n g e l m a n n i i , P. g l a u c a , and P_^ s i t c h e n s i s 137 F i g u r e 20. O r d i n a t i o n s o f f i r s t two components o f PCAs of s e p a r a t e v a r i a b l e s u i t e s f o r sampled p o p u l a t i o n s of standards and a r t i f i c i a l h y b r i d s 138 F i g u r e 21. O r d i n a t i o n s o f f i r s t two components o f PCAs of separate v a r i a b l e s u i t e s f o r i n d i v i d u a l a r t i f i c i a l hybrids and p u t a t i v e h y b r i d s 141 F i g u r e 22. O r d i n a t i o n s of f i r s t two components o f PCAs of s e p a r a t e v a r i a b l e s u i t e s f o r a l l i n d i v i d u a l s of P. e n g e l m a n n i i , P. s i t c h e n s i s , and " h y b r i d s "  143  F i g u r e 23. O r d i n a t i o n s o f means of f i r s t components from PCAs of s e p a r a t e v a r i a b l e s u i t e s f o r s t a n d a r d s , p u t a t i v e s , and " h y b r i d s " a g a i n s t 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 . . 163  xiv  F i g u r e 24. O r d i n a t i o n s of f i r s t two components from PCA o f s e p a r a t e v a r i a b l e s u i t e s f o r s t a n d a r d s , p u t a t i v e s , and " h y b r i d s " u s i n g c a l c u l a t e d p r e d i c t e d and r e s i d u a l v a l u e s m u l t i p l e l i n e a r r e g r e s s i o n i n MODEL 4  from 167  F i g u r e 25. Average v e c t o r s of s e p a r a t e v a r i a b l e s u i t e s p l o t t e d i n t o c o - o r d i n a t e s o f f i r s t two a x e s from PCAs o f a l l i n d i v i d u a l t r e e s b a s e d on o r i g i n a l and p r e d i c t e d and r e s i d u a l v a l u e s from m u l t i p l e r e g r e s s i o n 170  F i g u r e 26. O r d i n a t i o n s of means o f component s c o r e s f o r i n d i v i d u a l t r e e s from PCAs of s e p a r a t e v a r i a b l e s u i t e s of n a t u r a l l y and n u r s e r y grown t r e e s p l o t t e d and r e g r e s s e d s e p a r a t e l y a g a i n s t l o n g i t u d e and l a t i t u d e  172  F i g u r e 27. O r d i n a t i o n s of f i r s t two components f r o m PCAS of s e p a r a t e g e o g r a p h i c a r e a s and s e p a r a t e v a r i a b l e s u i t e s . 176 F i g u r e 28. P a r t i t i o n i n g of s o u r c e s o f v a r i a t i o n f o r n a t u r a l l y o c c u r r i n g P i c e a i n study area 180 F i g u r e 29. O r d i n a t i o n s of means, s t a n d a r d d e v i a t i o n s , and r a n g e s o f f i r s t component s c o r e s f o r a p r i o r i i d e n t i f i c a t i o n s and b i o g e o c l i m a t i c z o n e s 229  XV  ACKNOWLEDGEMENTS. Thanks and a p p r e c i a t i o n a r e e x t e n d e d t o t h e f o l l o w i n g p e o p l e f o r t h e i r t i m e and e f f o r t s p e n t on my b e h a l f d u r i n g t h e c o u r s e of t h e p r e s e n t s t u d y and t h e r e s e a r c h l e a d i n g up t o t h e f o r m u l a t i o n of t h e s t u d y . Bob G i t t i n s , J a c k Maze, Gary B r a d f i e l d , Ned G l i c k , and Y o u s r e y E l - K a s s a b y provided e n c o u r a g e m e n t and c l a r i f i c a t i o n on many a s p e c t s of t h e p h i l o s o p h y , a s s u m p t i o n s , and d e t a i l s of v a r i o u s s t a t i s t i c a l analyses. John Emanuel, M a l c o l m G r e i g , and Nancy R e i d p r o v i d e d valuable advice a s s o c i a t e d with computation. Fred Nuszdorfer and K a r e l K l i n k a p r o v i d e d h e l i c o p t e r t i m e and a c c e s s t h r o u g h p r e v i o i u s employment t o many a r e a s . W i t h o u t t h e i r h e l p much of t h e s a m p l i n g c o u l d not even have been c o n s i d e r e d . I owe an e s p e c i a l debt t o K a r e l K l i n k a who f i r s t i n t r o d u c e d me t o t h i s p r i c k l y p r o b l e m and p r o v i d e d t h e o r i g i n a l i m p e t u s f o r w r i t i n g the f o r e s t r y e p i l o g u e . The WFGA p r o v i d e d t h e r e s o l v e t o w r i t e the e p i l o g u e . U l f B i t t e r l i c k , R e i d C a r t e r , Don F o w l e r , D a n i e l Gagnon,and Helmar Hahn a l l p r o v i d e d s a m p l e s and expressed c o n t i n u e d i n t e r e s t i n t h e s t u d y even when r e s u l t s were not immediately forthcoming. J i m P o j a r p r o v i d e d comments w i t h r e s p e c t t o many a r e a s o u t s i d e t h a t s t u d i e d h e r e . F i e l d work w o u l d have been f a r l e s s e x c i t i n g and m o t i v a t i n g w i t h o u t the h e l p of P a u l C o u r t i n , Z o n g - h i Chen, and F r e d L a i n g R i d g e . S p e c i a l t h a n k s a r e e x t e n d e d t o N e l s a and "Buck" Buckingham and Ed T e i c h f o r d i r e c t i n g me t o P a t ' s P r a i r i e p l u s p r o v i d i n g a h o s p i t a b l e a t m o s p h e r e w h i l s t c o l l e c t i n g on t h e O l y m p i c Peninsula. G r a n t D i c k i n s o n and G e o r g e S h i s h k o v measured many of t h e o r i g i n a l s a m p l e s c o l l e c t e d i n 1979. Anna S c a g e l w i l l i n g l y p r o v i d e d t i m e t h a t was h e r s and h e r s a l o n e by i n d u l g i n g me i n c o l l e c t i n g , b o i l i n g c o n e s , e n t e r i n g d a t a , and back p a t t i n g . She a l s o p r o v i d e d a c l e a r view of t h e r e a l i t y a s s o c i a t e d w i t h Botanical Trivia Pursuit.  1  PREFACE. There beyond these has  are  science. reasons  conducting occurred  of  Regardless  me  this  r e s e a r c h and  on  analytic  reduced.  As  the  a summarization  Furthermore,  certain  investigate,  deficiency  the be  resulting  addressed  'collecting fortuitous  from  variation  three  events  a n a l y s e s , the i n the  study  results  I would have  be  of  to  number  were  i s not  as  liked.  addressed  the a n a l y t i c  ignoring to t h i s  effectively.  because  indicate  was  finished.  collections  The  wanted  of where  such  type  of  study,  long a f t e r field  trip  p r o v i d e d some d a t a into  biological variation  came t o o  I had and  that  thought some  proved  f a i l u r e to c a p i t i l i z e  on  intra-individual  i n f o r m a t i o n r e p r e s e n t s a source to i n d i c a t e  a  to  the  data  critical  intra-individual  I have e n d e a v o u r e d  in  extremely  of  of  late  occurred during  the nature  as a source  dissatisfaction.  A quick  and  intra-individual  This realization  p r o c e s s of m e a s u r i n g ,  in p r e s e n t i n g i n s i g h t variation.  used  I have t r i e d  realization  t r a d i t i o n a l approach  reflective  In  i s s u e s w h i c h , a t t h e o u t s e t , I had  size.  what  have o c c u r r e d .  Second, the deficiency  say  or c o l o u r e d the c o n c l u s i o n s  c o u l d be  as  go  s t u d y i n g t h e c o n s e q u e n c e s of  s i m p l y c o u l d not  i n sample  limitations  of  I cannot  the t h e s i s  that  study  spruces.  t h e p r e s e n t a t i o n of  parsimonious  to  especially  study  of any  study.  of m u l t i v a r i a t e  that  Consequently,  the  limited  results  approaches  on  writing  a result  a scientific  the o b j e c t i v i t y  trees,  which e f f e c t i v e l y  size  of  impact  to study  were drawn.  sample  for conducting  invariably  attracted  that  reasons  better  of  2  design  in sampling  incorporate The  such  third  and  a n a l y i s w h i c h would  intra-individual  event,  the  one  changing perception  genetics  f o r e s t r y and  of  to  w r i t i n g the  genetics.  the  principles  or  legislative  and  biology  the  where t h e r e  foresters  work and  operational  to t h i s  perceived  epilogue.  For  research  t o be  of  communicated  otherwise,  is a total  Forestry  R e s e a r c h Development  that  value,  1984).  that  scientific  of  There  the  under  i s as  which  an  I offer  the  (Young  the  studied.  the  results  to p o t e n t i a l  loss  for  than  conditions It  weeks  programmes  recognition  disinterest  r e c o m m e n d a t i o n s must be research  a concern  rather  f o r e s t r y i s conducted.  antithesis  any  any  few  forest  organisms being  was  forest  last  on  studies.  about,  of  the  administrative  the  s i t u a t i o n s , and  the  by  commitments  instances  In  a conference  industrial  problems,  on  relation  and  of  in future  most b i t t e r  forest biology.  t h e s i s I attended  of  political  were few  of  I am  T h e s e m e e t i n g s were p e r v a d e d  continuation met  information  that  c o n c e r n s my  effectively  and  users;  1972;  Secretariat  3  I.  Picea  A.  Diet,  i s a genus of w i d e l y  hemisphere c o n i f e r o u s species  that  occur  taxa  1971,  the  from  scale  - stiff  the  size,  or  form, and  1968;  (Wellendorf review tend  Sudo  and  i n La  to support  of  these  and  1968)  and the  edaphic  anatomical as  Dugle  chemical  Wellendorf  1968)  and  1968,  1972,  (Pravdin,  v a r i a b l e s 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 i n the 1974;  Garman  Simonsen  1979;  and  see  these the  content  e_t a_l.  1976;  1965c; P r i c e , e t a l . 1 9 7 3 ) .  further substantiated  clarified  1955;  DNA  Moir  have  (Wright  and  various  other  Evaluation  b a s i s of  between  species  only  cone  the  little  of  are  on  vary  Burley  to define  variables  and  species  cone  All  have been u t i l i z e d ,  r e c o g n i t i o n of  the  the  or f l a t t e n e d ;  habit.  variables  w e l l as  1977;  of  to d i s t i n g u i s h taxa or  1963).  traditionally  used  be  ancient  (Florin  traits  Chromosomal v a r i a t i o n  Evaluation  species  these  nature  variables.  1977;  of  shape - r h o m b o i d  can  informal 1966;  in Asia  traits:  40  Gaussen  twigs.  used  and  I t i s an  traditional  Fox  35  p u b e s c e n c e on  geographic  Kaufmann  Roi  Most  elevations.  needle  features  of more m o r p h o l o g i c a l Colleau  1955;  1977).  morphological  or a b s e n c e of  Other  crown  Wright  northern  into several  Cretaceous times  flexuous;  combinations  taxa.  i n c l u d e s between  genus have been d i s t i n g u i s h e d  three  presence  possible nine  of  subalpine  late  i n the  basis  1936;  Schmidt-Vogt  a t montane and  Species  and  (Flous  1973;  genus d a t i n g  on  trees that  distributed  have been v a r i o u s l y g r o u p e d  supraspecific Bobrov  INTRODUCTION.  1957;  g r o u p s of  d i s t i n c t n e s s of genus  (Duman  Marco  1931,  taxa  1957; 1939;  species  and  have  Daubenmire von  Rudloff  4  1975;  T a y l o r and  Sih-kin  1966).  b a s e d on between Gordon  Patterson As  these  1980;  La  Roi  w e l l as a t t e m p t s  variables,  Mikkola  1978,  1969).  earlier  1980,  1982;  B o n g a r t e n and  isolation  work on  and  (Johnson  Fowler,  Hanover  Two  groups  systematic Europe  1983)  t r e e s used  i n the  1965), and  of N o r t h  conducted  1977; the  P^  America.  b e c a u s e of  the  I n v e s t i g a t i o n of  focused  the  diversity  A m e r i c a : P_j_ s i t c h e n s i s P.  engelmanni i P a r r y  (Parry  ex  Martinez  separated  the  Engelm.  v a r i o u s taxa  from each o t h e r traditionally  genus.  flattened  only  species  l e a v e s , P^  inadequate the  subjected  - P_j_ o b o v a t a complex Lindquist  ("Glaucoides",  of  some of  P^  glauca  the  mexicana the  these  on  the  i s the  only  engelmannii  i n t h e complex w i t h  and  pubescent  to  has North  engelmanni i  (Martinez)  P^  mexicana  Taylor  complex  three  species  Fowler  (Moench.) V o s s ;  ssp.  the  1948;  species  i n western  P_;_ g l a u c a  in  been  complex  species  (= P_;_ g l a u c a  primarily  small  to i n t e n s i v e  pungens Engelm.; and,  of  in  crosses.  used to d i s t i n g u i s h  P_;_ s i t c h e n s i s  with  w e l l as  (Bong.) C a r r . ; P\_ g l a u c a  (= P_;_ e n g e l m a n n i i s s p . The  as  complex  the  shown by  Engelm.) T a y l o r ) P^  Patterson).  criteria  ex  1982;  i n t e n s i v e r e s e a r c h has  the  1955;  Santamour  e t a l . 1976;  importance  forestry. on  glauca  Such  Wright  t o some i n a d e q u a c i e s  P_j_ a b i e s  Pravdin,  and  crosses  e t a l . 1980,  of p a r e n t s  the  1939;  c r o s s e s owing t o t h e  i n these  Lee  relationships  (1982) and  genus have been  investigation:  (Schmidt-Vogt  Andersson  controlled  identification  number of p a r e n t  1968;  a l a r g e number of c o n t r o l l e d  (1967) however, have c a l l e d . a t t e n t i o n the  Dugle  to understand  s p e c i e s have been p e r f o r m e d 1976b,  and  and  are  morphological  other i n the  s p e c i e s of complex  P_J_ m e x i c a n a a r e branches,  and  the  5  P.  glauca  i s the only  species  i n t h e complex  with s t i f f  most a n c i e n t  fossils  from w e s t e r n N o r t h  cone  scales.  The referable  to Picea  a r e of M i o c e n e  t h e e x t a n t E\ c h i h u a h u a n a Madre O c c i d e n t a l e Klamath recent  region  macrofossils  scattered findings  P.  and  P^  and C a l i f o r n i a .  and m i c r o f o s s i l s  suggest the e x i s t e n c e dating  widely d i s t r i b u t e d  of P i c e a  Tertiary  lineage,  t h e P^ m a r i a n a  arcto-boreal  in eastern  P.  g l a u c a complex,  separately complex early  remain  Tertiary  (Critchfield  The P.  The  and  recent  Picea  - P^  relationships  the  more  of P i c e a  in  t o t h e present.: t h e  t h e remnants  rubens  o f a more  forest  in  lineage  of  and,  the  latutudes.  These  t o have m i g r a t e d  Patterson  migrant  o b s c u r e as d o e s  1980),  i n the l a t e  t h e P^  Cretaceous or  among t h e s e t h r e e  the r e l a t i o n  groups  to f o s s i l s  from t h e s o u t h e a s t e r n U n i t e d  glauca  of  from a  States  1984).  similarity  a b i e s complex  America  (Taylor  b e i n g t h e most  Tertiary.  spruces late  from A s i a  Sierra  These  groups  North America  considered  and  1983) .  widespread at a r c t o - b o r e a l  t h r e e groups are g e n e r a l l y  resemble  are widely  and montane e l e v a t i o n  western North America; latitudes  and  of t h e  Tertiary  (Crabtree  from t h e l a t e  temperate  1968)  of t h r e e major  b r e w e r i a n a - P^ c h i h u a h u a n a  1963)  b r e w e r i a n a S. Wats..of  i n western North America  North America  (Florin  M a r t i n e z (Gordon  of M e x i c o  o f Oregon  age  America  (Hills  o f e x t a n t forms  to the Miocene  and O g i l v i e  1970)  of t h e P^ g l a u c a  P_^ b a n k s i i  of a r c t i c  l e n d s weight  and North  to H u s t i c h ' s  (1953) h y p o t h e s i s c o n c e r n i n g a common d e r i v a t i o n  o f t h e P^  abies  6  and  glauca  complexes.  extant  species  by  the  Pleistocene  of  this  lineage  post-glacial  the  the  La  Oligocene.  differentiated and  by  has  by  the  of the  Wright's  the  P^  (1939) c o l l e c t i o n s ,  1980).  these  e v o l u t i o n of  liberal  the  glauca  complex  D u g l e , and  O g i l v i e and  work o f  emphasized latent P.  research the  suggests  that  much e a r l i e r  during  been  geographic  that  isolation  b a r r i e r s or  La  Roi  and  suggest  taxonomy  s i t c h e n s i s as  of  through  Dugle  (1968)  instead that  rather  than  taxa  i n the  s i t c h e n s i s as  the  a hiatus during  work of  research  Picea  to  was  an  this  glauca  work i s t h e  a distinct  species  continues  a distinct  in  1969  ( 1 9 5 7 ) , and  and  This  with Roi  by  recognition  i n the  i n the  Horton  recent  the  and  the (1959).  work  has A  of  Pj_ g l a u c a of  area  Roche's  P_j_ e n g e l m a n n i i .  a tradition  species  1968  e l a b o r a t i o n on  been c o n d u c t e d . P^  western  D a u b e n m i r e , La  followed  ( 1 9 5 9 ) , Garman  has  assumption  of  Rudloff  r e l a t i o n s of  assumption  Indeed, t h i s P.  Taylor  the  von  T h i s a b u n d a n c e of  More r e c e n t  by  breeding  reached  p u b l i c a t i o n of  earlier  a  Crabtree  (1955) s u g g e s t s  was  systematics  coincident  work.  1984).  t o have  differentiation.  to  been  f o l l o w i n g mountain b u i l d i n g  Wright  species  characteristic  1973), has  also Critchfield  mid-Tertiary  the  evolution.  R e s e a r c h on of  1972,  (1955) comments, and  been s u b j e c t e d  conservative  h y b r i d i z a t i o n , so  Bobrov  (see  that  were a l l d i f f e r e n t i a t e d  have been d i f f e r e n t i a t e d  strong morphological counter  the  1955;  Motte's  Patterson  than  that  complex  P_;_ e n g e l m a n n i i a p p e a r s  differentiation rather  and  (Wright  s i t c h e n s i s may  (Taylor  glauca  phenomenon  (1984) c i t i n g P.  of  O g i l v i e (1972) c o n t e n d s  complex.  recognizing  genus b a s e d on i t s  7  flattened hybrids  leaves  and  have been o b s e r v e d w i t h  Beckwith  1977;  Hanover  Daubenmire  1968)  have been  regarded  the  validity  the  rest  spite  patterns  source  complex  within  complex.  recognition  (Taylor  concerning  P^  s i t c h e n s i s that  they  and W i l k i n s o n  e t a l . 1976;  felt  t o be  1968;  1953;  presumed  l a c k o f d i s c r e t e n e s s between  species in  Roche  1975).  subsumed (Taylor 1968; 1969).  The  continued  (Daubenmire  1968); subsumed a s a s u b s p e c i e s O g i l v i e and von R u d l o f f  generally  von R u d l o f f  r e m a i n t o be r e s o l v e d :  status  Columbia  rest  merely as a  1970); La R o i and D u g l e  varietal  P. g l a u c a  clinal  o f t h e P_;_ g l a u c a  are i d e n t i f i e d  O g i l v i e and von R u d l o f f  (Hustich  have  f o r other  species  o f P_;_ e n g e l m a n n i i a s a s p e c i e s  1959;  the  d e m o n s t r a t e d f o r P^ g l a u c a  conventions  and D u g l e  s i t c h e n s i s from  o f P^ s i t c h e n s i s f r o m t h e  Where o t h e r  i n the v a r i a t i o n  nomenclatural  influence  This p o s i t i o n i s held in  P_;_ e n g e l m a n n i i i s g e n e r a l l y  1974;  they  has r e s u l t e d i n i n v e s t i q a t i o n s o f  of c o m p a r i s o n ( P o l l a r d ,  Daubenmire  Roi  complex.  i n d e p e n d e n t of any c o n s i d e r a t i o n  1959; Hanover  1969;  does not  d i s c r e t e n e s s of  have been c o n s i d e r e d ,  entirely  Roche  phenomenon t h a t  discreteness  of v a r i a t i o n  By c o n t r a s t ,  (Copes and  P^ s i t c h e n s i s and P_;_ e n q e l m a n n i i .  presumed  P^ g l a u c a  complex  1970;  (1969) o b s e r v a t i o n s  between  proceeded the  as a l o c a l  o f t h e assumed  t h e P\_ g l a u c a  e i t h e r P_;_ g l a u c a  Where  or P_^ e n q e l m a n n i i ( K l i n k a , e_t a_l. 1982)  of R o c h e ' s  This  a t low e l e v a t i o n s .  and W i l k i n s o n  of t h e E\ g l a u c a  variation  of  i t s occurrence  o f E\  1974;  La  glauca  1968); o r r e d u c e d t o  L o v e and Love  1966).  P^ e n g e l m a n n i i  The and  h a s been a d o p t e d i n f o r e s t r y a p p l i c a t i o n s i n B r i t i s h where t h e complex  i s r e f e r r e d t o as simply  "Interior  8  spruce"  in  (Kiss  1976).  Contrary  to the  the  that  glauca  suggest  related the  complex a r e  t h a t E\  t o P^  b a s i s of  assumption  that  the  remarks c i t e d  e n g e l m a n n i i be  sitchensis  than  suggest  of P\_ g l a u c a  the  a similarity presence  areas  reported of  the  suggested P.  e n g e l m a n n i i may  attention  generally  prevented Nash  1978b).  sitchensis  seem t o be variation  and  An  area,  investigation  Fox  on  (1977)  the  basis  work a s s o c i a t e d other  the  1969; of  with  of  complex.  the a r e a  has  1968;  i n Ching  particular  study  special  and  the v a r i a t i o n  The  as  for  (Daubenmire  others  in  Columbia  a phenomenon  isolation  with  observations  British  to b e t t e r c i r c u m s c r i b e the  Pj_ g l a u c a  and  P_^ s i t c h e n s i s  extensive c o l l e c t i o n Roche  impression  (1957) c a l l e d  P_^ e n g e l m a n n i i i n t h i s  necessary i n the  local  Garman  m a t e r i a l s of  1978;  Moir  P_j_ s i t c h e n s i s  have been as  from t h i s  2  of  this  on  1  southwestern  Although  to c o l l e c t i n g  F a l k e n h a g e n and  P.  not  suggested.  engelmanni i  Sziklai  of  closely  complement .  i n K l i n k a , e t a l . (1982) and coast mainland  (1957)  F r a n k l i n (1961),  t o P_;_ s i t c h e n s i s  made i n c o n j u n c t i o n w i t h  that h y b r i d i z a t i o n  previously  P.  Alternatively,  of a/3-chromosome  Observations that  more  s e e d l i n g morphology, c o r r o b o r a t e s 1974).  is discrete  by Garman  considered  t o P_j_ g l a u c a •  ( s e e however J e f f e r s  of  sitchensis  and  of area  range  would  of  presented  here  G u y l a K i s s , B r i t i s h C o l u m b i a M i n i s t r y of F o r e s t s , V e r n o n c o n f i r m s t h e o b s e r v a t i o n s of M o i r and Fox (1977) and o f f e r s t h e o b s e r v a t i o n t h a t P_;_ e n g e l m a n n i i l a c k s a (b-chromosome complement. 1  2  be  F o r t h e sake of b r e v i t y , P^ s i t c h e n s i s and P^ e n g e l m a n n i i will r e f e r r e d t o c o l l e c t i v e l y as P i c e a u n l e s s o t h e r w i s e n o t e d .  9  addresses  this  issue  Conceptually, explaining taxa.  of v a r i a t i o n  such a study  the v a r i a t i o n  Analytically,  and  correlating  are  two s c a l e s  variation.  research  intra-  description  must be b a s e d on an e s t i m a t e Explanations  not  Analytic  been c o n s i d e r e d  consideration- has -  against  which  offers  inter-individual numerical  in conifers.  under two g e n e r a l  variation  concerning in addition  Where  analytic scale  c a n be compared.  variation  A  potentially  the nature of t o the r e s u l t s of  hybridization  explanations  variation  could  f o r emergent  be t e n d e r e d ,  processes: hybridization  hypothesized process  study w i l l  tendered  v a r i a t i o n has  i t provides a convenient  of s p e c i f i c  inter-individual  Neither  in conifers  comparisons.  A plethora of  generally  variation  variation.  although  to intra-individual  explanations  variation  Where t h e y have been  view o f i n t r a - i n d i v i d u a l  further  There  have been made f o r c e r t a i n  inter-individual  developmental  variation  attention  been made  scales.  of i n t e r - i n d i v i d u a l  t o the environment,  developmental c o n s i d e r a t i o n s conifers.  at various  describing  and i n t e r - i n d i v i d u a l  for intra-individual  have been r e l a t e d  o u t by  of i n t r a - i n d i v i d u a l  have n o t commonly been p r o p o s e d . they  o f t h e two h y p o t h e s i z e d  i s carried  of v a r i a t i o n  of v a r i a t i o n :  Effective  i n v o l v e s e x a m i n i n g and  i n the trees  such  patterns  of P i c e a .  constitutes  explanation,  a d d t o our knowledge c o n c e r n i n g  evolutionary  consequences  i n Picea  (Wright  but they  fall  and d i f f e r e n t i a t i o n .  need be m u t u a l l y  a valid  patterns  exclusive. then  If  the present  t h i s p r o c e s s and i t s 1955; B o b r o v 1972,  10  1973). the  If d i f f e r e n t i a t i o n  results  of  evolutionary and  such a study  or more, s p e c i e s  Owing t o t h e c o n d u c t e d and, individual  large  s c a l e upon w h i c h t h i s  be  considered.  i n examing  patterns  p r e v i o u s l y i n both  variation  b a s e d on  1968;  Pollard, (Pollard,  1967;  1965a,  Lines  1977,  1978; and  variation  1976;  to  sampling that  general  taxa  species  every  present  specific  immediate and  larger  and  controlled  are variation  over  such a  and  Burley  have been o b s e r v e d  of  variables.  (Moir  Daubenmire  Over  several patterns (Falkenhagen I975b,c,  Fox  have  1974;  the been Daubenmire  1980b; M i k s h e  1977); c l i n a l  1971;  variation  1966b,c; O ' D r i s c o l l 1976a;  I l l i n g w o r t h 1976;  McLean  and  f o r s e v e r a l s c a l e s of  variation  Moir  e t a l . 1976;  El-Kassaby  cannot  is  Such p a r e n t a g e - s p e c i f i c  1966c; F o r r e s t  e_t a_l. 1976;  from c l i n a l and  the  discontinuous  Falkenhagen  the  s e v e r a l types  r a n g e of  Burley  study  organisms  respect  of v a r i a t i o n  described  observed:  versus  area.  Independent  geographic  one  i n t e n s i v e procedures  geographic  contrast  O n l y more d i s t a n t a n c e s t r y  require detailed  counter-productive  essential  in sexually reproducing  can  - labour  The  and  species d i f f e r e n t i a t i o n  some d e g r e e u n i q u e ,  would  ecological  then  merging.  selection.  selection  crosses  single  for r e l a t i o n s h i p s with  p a r e n t a g e and  explanations  address  in Picea.  geographic  that  i s to  explanations  scale  may  considerations  g e n e r a l i z a t i o n s are:  two,  seems a p l a u s i b l e h y p o t h e s i s ,  1983; and 1968;  Mergen and  Daubenmire  Fox  1977;  Burley  Thielges  1968); d e v i a t i o n s  Forrest 1965b,  1980b; L e w i s  1966b;  Cannell  11  and  Willett  1975; F a l k e n h a g e n  Hanover and W i l k i n s o n 1980;  Falkenhagen  1966a,c; F o r r e s t reported and  these  1970); u n i f o r m i t y  (Yen a n d  1980b).  Other  observation  (1966a), H a r r i s  invoke  several causes:  and  and, e v o l u t i o n a r y b o t t l e n e c k s .  in  under  the p r i n c i p a l  expectations  The m a j o r  taken  t h e two s p e c i e s ,  al  1982; G r i f f i n  occurrence  source of  latitude,  process  that  patterns natural  whereas  (Fowells  1965; H o s i e  and C r i t c h f i e l d  of P i c e a  i n the study  e t a_l. 1982).  selection  area  and  occurrence  1975; K r a j i n a , e t  1976; L i t t l e  1971).  The  i s f a r more s p o r a d i c  than  by t h e maps i n t h e s e p u b l i c a t i o n s Assumptions  b a s e d on t h e p r e s u m p t i o n  have  and the t h e o r e t i c a l  f r o m t h e presumed d i s t r i b u t i o n  c o n t i n u i t y suggested  (Klinka,  f o r the observed  t h e assumed v a l i d i t y  operative  of  are  hybridization; isolation  i s c o r r e l a t e d with  of the e x p l a n a t i o n s  been t e n d e r e d  the  refugium;  P^ e n g e l m a n n i i i t i s c o r r e l a t e d w i t h e l e v a t i o n .  Most  is  differentiation;  i n Ching  Explanations for  glacial  selection;  i n Pj_ s i t c h e n s i s  1970; B u r l e y  ( 1 9 7 8 ) ; and o t h e r s  migration;  variation  El-Kassaby  on v a r i a t i o n a r e  (1978b) and O ' D r i s c o l l (1976b)..  patterns  drift;  e t a l . 1975;  1976; Hanover a n d W i l k i n s o n  i n Burley  Sziklai  1978; P o l l a r d ,  that  large  need t o be r e c o n s i d e r e d inter-breeding  when  populations  continuuous.  On a l o c a l concerning largest  the nature  source  populations. seedling  s c a l e where o b s e r v a t i o n s of p o p u l a t i o n  of v a r i a t i o n These  populations  have been  variation  i s invariably  s t u d i e s of p o p u l a t i o n s r a t h e r than  mature  reported  i n these  contained  taxa, the  within the  have been b a s e d on  individuals,  a s have  12  most o t h e r s observed  studies  high  observations  on  intra-population reported  species.  Explanations  Picea,  well  as  Furthermore,  conifers  as  f o r the  other  specific  the  Picea  - however  m a g n i t u d e of  1974).  majority  of  other  intra-population  explanations  i t may  be  possible  intra-population  the  v a r i a t i o n in been  cannot  for circumscribing  with  coniferous  c o n i f e r g e n e r a have not  local  The  variation is consistant  for high  h e r e owing t o a c o n c e r n of  (Falkenhagen  be  tendered.  presented  general  to provide  an  variation  estimate  v a r i a t i o n compared  of  to  inter-population variation. Studies systematic various  of  v a r i a t i o n serve  purpose.  parts  species.  of  Both  their  successional  species  r a n g e and  Furthermore,  reforestation  in  the  they are  programmes as species.  both s p e c i e s  reforestation  and  comparable  P.  s i t c h e n s i s to  widely  study  present  in that  planted  are  species  It  of  Britain.  f o r e s t r y has  commercial  been  species  1982).  consideration  be  Dietrichson given  occur  as  of  tree  in  important  i n European  British  that  the  Columbia  are  that  observations  t o have r e l e v a n c e The  in  Columbia.  i s worth n o t i n g  and  a  primary  British  exotics  Britain,  prove  as  commercial  is particulary  interior  H y b r i d s between F\_ s i t c h e n s i s and (Faulkner  important  u s e d as  southern  well  landscape dominants  in southwestern  s t u d y may  part  economic as  p o t e n t i a l l y valuable  aforestation.  t o t h o s e of  made i n t h e forestry  this  are  both  central  Additionally,  of  are  P_;_ e n g e l m a n n i i  r e f o r e s t a t i o n i n the  latitudes  an  to  s i g n i f i c a n c e of  r e a l i z e d since in B r i t a i n P_j_ g l a u c a  (Pearce are  (1971) s u g g e s t s  t o P_;_ e n g e l m a n n i i  i t i s the  also  that  i n Norway.  most  1976). of  interest  more Various  13  p r o v e n a n c e s of  e n g e l m a n n i i a l s o may  p r o v e u s e f u l as an  in  some p l a c e s  i n western North A m e r i c a  et  a l . 1981).  Alternative explanations  differentiation forestry  or c o a l e s c e n c e  relating  "inter-specific"  to the  can  be  Roche  1977;  Phelps et  Roche and  the  species  hypothesis P.  being  areas  hypothesis not  of  of  Krajina,  specific and  present  not  Hunt  conclusions i n the  taxa and  1965;  from  this  succeeding  observations  are  also  given  further discussion i s provided framework.  study  i s concerned with individuals  the  the  range  been d e s c r i b e d p r e v i o u s l y . there The  then  an  are  two  Such a s t u d y  of  The  taxa:  inference derived examination  continue  nature  of  i n a p o r t i o n of  engelmani i .  taxa.  1972;  the  Fowler  Dobbs  i s appropriate  two  ecology  1970;  P_^ s i t c h e n s i s  P^  and  of  to support  Forests  is: if  trees  from  the  seeks to uncover  Helmar Hahn, B r i t i s h C o l u m b i a M i n i s t r y of F o r e s t R e g i o n , p e r s . comm. 1  of  species, further  and  sampled p r e v i o u s l y w i l l  recognition  use  Christensen  tested i s that  s i t c h e n s i s and  this  and  1975;  analytic  t h a t has  these  occurrence  in naturally occurring  e n g e l m a n n i i and  these  of  have been d e t a i l e d  and  summary, t h e  of v a r i a t i o n P.  Ruth  literature  conceptual  In  importance  Folwer  Additionally,  from more r e c e n t of  implications for  of p r o d u c t i o n  Where r e l e v a n t , s p e c i f i c  literature  chapters.  species  b i b l i o g r a p h i e s ( K r a j i n a 1969;  H a r r i s and  a l . 1982).  c o r p u s of  economic  in various  1973;  for  have  feasability  p e r t a i n i n g to the  found  (Shepperd,  1  crosses.  Owing t o t h e information  may  exotic  and  ,Vancouver  1 4  describe  patterns  geographic  To  variation.  The  offer  inter-individual  must be  primary  and  c o n s e q u e n c e of The  explanations  the  given  question  being  the  differentiation  secondary concern  addressed  the  of a  two  of  conceptual  and  intra-individual i s whether and  taxa,  single  i s to present  patterns  of  various  patterns  variation,  hybridization of  at  f o r the  to aspects  differentiation  f o r these  variation  explanations  i s a c o n s e q u e n c e of  introgression  taxon.  assess  attention  variation  inter-individual  s c a l e s and  described. analytic  of  subsequent or  simply  polymorphic  possible  differentiation.  the  15  II. 1.  AND METHODS.  Introduction. The  phenomena  phenotypic The  requiring explanation  variation  identification  towards p r o p o s i n g Quantification the  of P i c e a  of a g i v e n  and s u b s e q u e n t  focuses  explanations.  about  surrouding  sampling,  and how  As  completely. characters there  in this  case,  information  the c h a r a c t e r i s t i c s of  i s an e s t i m a t e  must be t e n d e r e d  that  of the  respect  the b i a s  T h i s amounts t o c a r e f u l l y  of t h e s e  c h a r a c t e r s , the n a t u r e of  t h e measurement e r r o r a s s o c i a t e d  so t o o i s i t i m p o s s i b l e  with  a n a l y s i s so t h a t  Indeed,  there  by i t s  t o measure a c h a r a c t e r  are probably  an i n f i n i t e  by w h i c h an o r g a n i s m c a n be d e s c r i b e d  number o f  - so t o o a r e  number o f ways t o measure a c h a r a c t e r .  Measurements o r d e s c r i p t i o n s a r e s i m p l y  that  a c t of q u a n t i f i c a t i o n  a r e n o t emburdened by an o b s e r v a t i o n a l b i a s .  an i n f i n i t e  number  are required prior to  one c a n n e v e r d e s c r i b e an o r g a n i s m c o m p l e t e l y  characters,  step  d e s c r i p t i o n o f t h a t phenomenon and  c h a r a c t e r s c a n be m i n i m i z e d d u r i n g  explanations  a major  phenomenon.  i t i s manifest  estimates.  the nature  quantifying  f o r that  Since, q u a n t i f i c a t i o n  i n these  considering  represents  Columbia.  t h e s p e c i f i c , phenomenon and t h e c o n d i t i o n s  phenomenon, e x p l a n a t i o n s inherent  British  a p o r t i o n of the p o t e n t i a l  i t soccurrence;  organism.  phenomenon  However, t h e v e r y  a t t e n t i o n on o n l y  available  are v a r i o u s aspects of  i n southwestern  an e x p l a n a t i o n  c o n d i t i o n s under w h i c h  tendering  an  MATERIALS  t o an o b j e c t  (O'Grady  1982).  r u l e s used  to assign a  I t i s important  c h a r a c t e r s a r e the m a n i f e s t a t i o n s  of i n d i v i d u a l  to realize organisms  1 6  by  w h i c h we  individual  d e s c r i b e , communicate, organisms.  independent 1973; and  1982).  characters,  multivariate distinction  here,  as  of an  Although  we  i t is specific  between c h a r a c t e r  and  p u r p o s e as sample,  elsewhere,  draw  see  actually  and  fundamental concept, little  to a l l e v i a t e  Measuring sources  analytic  but  intertwined with  i n any  theoretical  Several  a n a l y s i s and systematic  technique  well  g r o u n d s may  be  the  variation  variables.  and  s u i t e d to inadequate  c o v a r i a t i o n , sampling  analytic  of  does  such  collecting  are  unique  inappropriate  one  as  system. on  with  collection  limitations,  to  i s that  of a b i o l o g i c a l  nature  are  appropriate  r e s o l v i n g a problem  the  usage.  determining  framework used  or  us  a  from  sampling  s e t of d a t a  protocol allows  around  1981),  of an  for a given  Understanding  It i s crucial  and  and  selection  investigation  measurement e r r o r f o r a g i v e n for a given  arise  of  same  a subject  samples,  conceptual  the  usage of  measuring  aspects  The  world  (Wiley  t h a t may  parts,  methodologically  the on  intuitive  to r e s o l v i n g that problem  samples and  of  a character  ambiguities  their  limited  the  to bear  variables, collecting  the  analytic  respect  s u c h as  synthesis.  appplied An  the  of e r r o r a s s o c i a t e d w i t h  inextricably  trees,  perceptions  an Heywood  estimator.  how  about  d i f f e r e n c e between  to acknowledge c l e a r l y  acknowledging  and  measure.  r e c o g n i z i n g the  and  Merely  as  v a r i a b l e serves  conceptually constrained preceptions  study.  (Davis  variables in a  or e s t i m a n d and  b r i n g these  exist  organisms,  and  we  inferences  does not  organism  p e r s p e c t i v e t h a t we  epistemological population  A character  manifestation  O'Grady  and  of  of v a r i a b l e and  w e l l as  to s e l e c t  an  associated the  caveats  1v  appropriate effort  a n a l y t i c methodology.  was  spent  determining  In  equally  inferences central of  the  that  t o be  that  an  considerable and  appropriate  to  recognizing  to bear  state clearly from  t o be  on  a  the  the  analytic  drawn h e r e  nature  genes  (Solbrig  1968).  look  of of  study, i t the  Specifically,  i s that,  i n d i v i d u a l s that  perceptive  subject  such a s t u d y .  number o f  besides  the  a l i k e are  i n common and  are  the effect  assumed  to  thus capable  of  Characters. Phenotypic  genetic the of  so  brought  drawn  environment,  interbreeding  2.  are  important  large  error  to e x p l i c i t l y  inference  share a  end  employed.  addition  constraints is  be  this  in d e f i n i n g v a r i a b l e s , sampling,  associated  method c o u l d  To  v a r i a b l e s are  information  and  the  i n t e r n a l environment the  o r g a n i s m , and  that  are  than  inter-individually  less  buffered  the  more v a r i a b l e between  are  genetic,  d e g r e e of  that  immediate  local  of  information  external  environment.  with  environment Variables  individual (intra-individual)  environment  individuals.  r e g a r d e d as  than are  B o t h t y p e s of  d i f f e r e n t e x t r e m e s of  being  those  that  variables  a c o n t i n u u m of  buffering.  Assessing relativistic variation  at  an  expression  ( i n t e r - i n d i v i d u a l ) are  are  albeit  the  surrounding  more v a r i a b l e w i t h i n  against  an  i n t e r a c t i o n of  and  the  r e g a r d e d as  differences  exercise.  within  a taxon  between t a x a  T h e r e must be  is, analytically,  information  (intra-specific)  as  well  a  about as  between  the  18  taxa  (inter-specific).  variables  t h a t have an  intra-taxonomic contend and  with  inter-taxonomic  variation  are  individuals,  intra-individual hypothesized their  taxa,  variation,  concerning Davis  variation and  taxonomic  provide  g i v e s an an  are  example w i t h  central  quantification  Similarly,  can  an  i n f e r e n c e be Heywood  to p o p u l a t i o n  of  study  in  made  1973).  respect  Newhan and  respect  aspect  this  of p h e n o t y p i c  variation  Columbia.  Specifically,  between  individual  trees.  relationships the  of  As  to  Jancey  this (1983)  differentiation.  individual  which  certain The  phenomenon, out  Picea  of  these  c h a r a c t e r s may  characters  a character  communicate infinite  had  be  of  selected for this  form of p a r t s of t o meet two  the  in  southwestern  i s in distinguishing the  product  information  about  characters  individual,  only  value. study  organism.  criteria.  i s the  number of  i t i s p o s s i b l e to d e s c r i b e a given  general  selected  of  a  o b j e c t i v e i s to understand  t r e e s and  of an  i s to p r o v i d e  interest  the  n e c e s s i t y t o d e s c r i b e and  particular  the  then,  inter-correlated  information with  British  by  than  Variables selected. The  of  If  more v a r i a b l e between  only  example.  must  inter-individually.  v a r i a b l e s are  further  than  variable variation  r e l a t i o n s h i p s ( D a v i s and  (1983) p r o v i d e s  view and  and  and  larger  However, one  inter-individual  these  then,  variation  analytically,  intra-  t h a t have a h i g h e r  in d e f i n i n g taxa,  preferred.  and,  c o - v a r i a t i o n expressed  variables  2.1  Traditionally,  First,  had  to  The  variables  the  circumscribe  accuracy  and  a  19  precision was  (Cochran  assessed  measurement S e c o n d , as those  and  only  interest  was  of  a high  variables  degree  of  "measurement e r r o r " ) were s e l e c t e d .  were  variation,  inter-individual  only  than  used.  in this  leaf  these  study  characters length).  are  morphological  refer  to the  Anatomical  form of a s t r u c t u r e ( e . g .  and  external  characters  number o f  form  define  endodermal  in c r o s s - s e c t i o n ) . The  characters  genome of selected  the  s e l e c t e d can  plant.  of  fundamentally other  the  genome.  those  and  theoretical  architecturally represent organism.  the  of  complex  to the  canopy.  employed d i d not  allow  the  the  better  criteria  the  adopted  w o r k e r s when d e a l i n g Some w o r k e r s  i n t e r p r e t a t i o n s to certain upon t h e  objectives  of  and  researcher.  d e s c r i p t i o n of a l a r g e ,  and  size  characters could  the  sample of  characters  variables.  depending the  a  o r g a n i s m , c h a r a c t e r s were s e l e c t e d t h a t  reproductive  Owing  reproductive portion  genetic  others,  a general  other  chemical  p r e d i s p o s i t i o n of  provide  selection  by  as  t h e more  individual,  The  adopted  morphological  v a r i a b l e s than  regarded  Developmentally,  would a t t a c h more s p e c i f i c  To  be  from more p o r t i o n s of an  representation  these  v a r i a b l e s with  (=  Morphological  internal  with  measurement  in i n t e r - i n d i v i d u a l  used  a s t r u c t u r e (e.g.  are  those  variation  characters  anatomical.  cells  the  variables displaying greater  The  the  of  repeatability  intra-individual  of  1977)  v e g e t a t i v e p o r t i o n s of  of  the  not  be  S i m i l a r y the  the  organism, v e g e t a t i v e examined  from the  preparatory  and  same  techniques  f o r a one-to-one correspondence  to  be  20  made between c e r t a i n v e g e t a t i v e  C h a r a c t e r s a r e grouped there the  variables.  together  i n "character  s u i t e s " as  e x i s t s a one-to-one developmental correspondence  characters  f o r any g i v e n  sample.  Developmentally, a  one-to-one c o r r e s p o n d e n c e does not e x i s t samples  from d i f f e r e n t c h a r a c t e r  characters  o f c o n e s , cone  between  suites.  scales  between  individual  F o r example,  and b r a c t s ,  twig  although  morphology,  t w i g anatomy, n e e d l e m o r p h o l o g y , and n e e d l e anatomy a l l c o u l d examined  f o r each  variables analyze  on s e p a r a t e p a r t s  for variable  variables  except  aspects  tree  The  i n t e r - c o r r e l a t i o n s between  f o r separate  trees.  Correlation  t h e canopy o f a  but a r e not meaningful i n  s e l e c t i o n of a c h a r a c t e r  and d e s c r i p t i o n of  was b a s e d upon p r e v i o u s work on P i c e a  1956; J a n s s o n a n d Bornmann  1953;  Dalgas  1973; J e f f r e e ,  1983; M a r t i n e z  Colleau  1968; S t o v e r  Carlson  and B l a k e  1981; Duman  1961 i n T a y l o r  1944) a n d o t h e r  conifers  1968)  was n o t u s e d b e c a u s e o f t h e h i g h e r r o r (see a l s o  Parker,  (Fulling  1934;  t o Holmgren a n d Keuken  o f many s a m p l e s .  this variable  1976a;  i n the f i e l d and  acronyms a c c o r d i n g Cone s c a l e  1;  and P a t t e r s o n 1980;  1969) a n d e x a m i n a t i o n  (UBC, V, WS;  (see Table  1957; de L a u b e n f e l s  e t a l . 1971; Gordon  1974)  measuring  between, s a y ,  sense.  Facey  herbarium  s u i t e s of  t o mean v a l u e s c a l c u l a t e d f o r  are a n a l y t i c a l l y possible,  initial  variables  al.  Thus one c a n n o t  of n e e d l e morphology and cones w i t h i n  a developmental  et  t h e y a r e measured a s  o f t h e same t r e e .  as i t a p p l i e s  these c h a r a c t e r s  single  individual tree,  be  phyllotaxis  Cannell  (Daubenmire,  associated  and Bowler  with  1978).  T a b l e  1.  C h a r a c t e r s  u s e d  e x c l u d e d .  S t u d i e s :  D a u b e n m i r e  1 9 7 4 ;  1 9 5 7 : 7  G a r m a n 1982:  1 0  O g i l v i e R e e d  -  -  -  i -  n p r e v i o u s  1 9 5 9 : 8  L a R o i a n d D u g l e  a n d v o n R u d l o f f  P a t t e r s o n  -  -  V A R I A B L E S C O N E  -  K h a l i  1968: R o c h e  T a y l o r .  1 9 6 8 : 2  R a t i o -  a n d N a s h  1 9 6 8 ;1 3  1 9 4 9 ; 1 5  1 9 8 0 : 1 8  s t u d i e s .  D a u b e n m i r e  F a l k e n h a g e n  H o r t o n  a n d F r e y t a g  a n d  1 4  1 1 -  1  -  1 9 7 4 : 9  M i t t o n  P a r k e r .  -  -  1 9 7 2 : 3  F u n s c h  K l i n k a ,  a n d A n d a l o r a S t r o n g  1 9 7 5 : 19  -  -  1 9 7 5 :6 1 9 8 1 ; 1 2  1 9 7 8 : 1 4  1 9 7 8 : 1 7  -  c o n e  T a y l o r 1 9 5 9 .  l e n g t h  c o n e  w i d t h  m e a n  c o n e  m e a n  c o n e  c o n e  1 , 2 . 3 . 1 7 . 1 8 1.2.3,  a p e x  c o n e  4 . 5 . 8 . 9 . 1 3 . 1 9  w i d t h  8 . 9 . 1 3 . 1 9 S . 10  s h a p e  6.  w e i g t n  S C A L E  10  8  M E A S U R E M E N T S .  s c a l e  p h y 1 1 o t a * y  s c a l e  s h a p e  s c a l e  m a r g i n  s c a l e  1 7 . 18  l e n g t h  t e x t u r e  c o n e  C O N E  1 . 2 . 3 . 6 . 1 0 . 1 7 . 1 8  l e n g t h  c o n e  m e d i a n  1.2.3 10  t h i c k n e s s  13  t e x t u r e  s c a l e  a p e x  s c a l e  m a r g i n  ' 0  s h a p e  7 . 1 0 . 1 3 . 1 9  f o r m  7.  1 0 .1 2 . 1 3 .  1 9  s e a l e  l e n g t h  1 . 2 . 3 . 5 . G . 8 . 9 . 1 0 . 1 2 . I S . 1 6 . 1 7 . 1 8  s c a l e  w i d t h  1 . 2 . 3 . 5 , 6 . 8 , 9 . 1 0 . 1 2 , 1 5 , 1 6 . 1 7 , 1 8  s c a l e  t a p e r  s c a l e  2 . 7 . 9 . 1 2 . 1 3 . 1 6  w i d t h  f r e e  2 m m  b e l o w  15 2 . 3 . 9 . 1 3 . 1 6 . 1 7 . 1 8  s e e d  i m p r e s s i o n  l e n g t h  s e e d  i m p r e s s i o n  w i d t h  B R A C T  a p e x  s c a l e  1 5 . 1 6 9  M E A S U R E M E N T S .  b r a c t  a p e x  b r a c t  m a r g i n  b r a c t  s h a p e  6 . 7 . 1 0 . 1 3  f o r m  10  l e n g t h  6 . 9 , 1 3 . 1 5  b r a c t  w i d t h  9  b r a c t  t a p e r  9  S E E D  M E A S U R E M E N T S ,  s e e d  w i n g  l e n g t h  s e e d  w i n g  w i d t h  s e e d  l e n g t h  s e e d  w i d t h  T W I G  4 4 4 4  M E A S U R E M E N T S ,  b r a n c h  d i a m e t e r  s t e r t g m a t a  5  p r o j e c t i o n  s t e r i g m a t a  1 . 2 . 3 . 6 . 1 0  a n g l e  1 . 2 . 3 . 6 .  p u l v i n u s  l e n g t h  5  p u l v i n u s  s h a p e  13  1 0  p u b e s c e n c e  1 . 2 . 3 . 5 . 6 .  t w i g  13  c o l o u r  M E A S U R E M E N T S , b u d  s c a l e  b u d  s c a l e  b u d  s c a l e  a p e x  b u d  s c a l e  m a r g i n  b u d  s c a l e  f o r m  L E A F  l e n g t h  13  w i d t h  13 s h a p e  13  f o r m  13 1 1  M E A S U R E M E N T S , l e a f  s h a r p n e s s  l o n g e s t a v e r a g e s t o m a t a  1  l e n g t h  1 . 2 . 3 .  l e a f  l e n g t h  5 . 1 3 . 1 4 . 1 9  1  < " d o r s r e s  i n  s a c  l e a f  w e i g h t  l e a f l e a f  l n e s  I. i o n  l e n g t h  c o l o u r o r i e n t a t i o n o d o u r  I I .  1 4  11.14 13 5 5 5  1 7  2 . 3 . 6  tv e n t r a 1 l t y " 1  s a cp o s 1 t  r e s i n l e a f  5 . 1 1 . 1 3  l e a f  1 0 .1 1 . 1 2 . 13.  -  -  T a y l o r  S T U D I E S  m e d i a n  -  a±.  e t  S I Z E . l a r g e s t  B U D  -  a n d M c l a c h l a n  1 9 6 9 ; 1 6 e _ t a±  v a r i a b l e s  D a u b e n m i r e 1 9 7 8 : 5  1 7 . 1 8 . 19  22  Ratio of  variables  t h e poor  review  calculated  statistical  in Phillips  p r o p e r t i e s a s s o c i a t e d with  of v a r i a b l e s  a number of d i f f e r e n t  manners.  continuous,  categorical,  an  flexibility  generally with  allow.  from  r a t i o s (see  that other  nature  Continuous types  variables  of v a r i a t i o n  genetic control  c h a r a c t e r s may be done i n  C h a r a c t e r s c a n be e x p r e s s e d  or counts.  Continuous  the continuous  hypothesized  because  1983).  The t r a n s l a t i o n  analytic  by o t h e r s were n o t c a l c u l a t e d  over  their  variables  of v a r i a b l e s  as  offer  do n o t  a r e a l s o more i n k e e p i n g of c h a r a c t e r s and the expression  (Falconer  1981)  2.2 V a r i a b l e measurement Samples u s e d selected  randomly  i n determining  measurement  from~the t o t a l  number o f samples  (Appendix  II).  precision  of measuring.  repeatability  repeatability.  Measurement  of these  repeatability  The i n i t i a l variables  measurement  repeatability  assessment  observers.  was made w i t h  were  available  i s an e s t i m a t e  was c a r r i e d  measurements made by two d i f f e r e n t  repeatability  of the  o f measurement  out using  replicate  Assessment of  an a n a l y s i s  of v a r i a n c e  model o f t h e f o r m : (MODEL 1.) y = A + e.  where y i s a measurement observer, residual  f o r a given  A i s the hypothesized variation.  variable  effect  by a g i v e n  of an o b s e r v e r ,  and e i s  23  Variables displaying attributed  on  another  the d e s c r i p t i o n  sample.  of a v a r i a b l e  v a r i a b l e s which appeared sample by  Following greater  than  two  and  repeatability  was  estimates over  suite.  ambiguous.  invariate  in fact  on  over  spent  Figure  four  assumed  showed t h e  1 illustrates  includes  t h e method o f s a m p l i n g ,  of  during  in Table f o r each  form  variable  number of  proved  eventually  specimen  the  2.  t o be  amount o f measurement  variable  to  repeatability  with a larger  descriptions  f o r each  Variables  measurement  Average  the v a r i a b l e s  full  possessing  samples  separate times  symmetry  I c o n t a i n s the  technique  on  all.  repeatability  largest  still  sample.  estimates are given  Appendix  measurement  at  i n measuring.  g i v e s the average  that  Quantitative  several  re-examination,  measured a t f o u r  r e f e r e n c e p o i n t s and  t o make  were r e - m e a s u r e d  yet another  they v a r i e d  second  these  sought  and  between o b s e r v e r s were a g a i n  In g e n e r a l , t h o s e v a r i a b l e s  variables  variation  observers at a higher m a g n i f i c a t i o n .  were r e - e x a m i n e d  this  c o u r s e of t h e y e a r  2 also  t o be  re-examined  whether  Following  Table  less  re-measurement, t h o s e v a r i a b l e s  invariate  determine  Re-specification  10 p e r c e n t v a r i a t i o n  re-specified proving  10 p e r c e n t  t o o b s e r v e r s were s u b j e c t e d t o r e - s p e c i f i c a t i o n  re-measured  another  g r e a t e r than  error.  selected.  f o r the v a r i a b l e s p r e p a r a t i o n , and  suite.  the  and  T a b l e  2.  A v e r a g e  v a r i a b l e  i n t e r - i n d i v i d u a l M e a s u r e m e n t f o u r  r e p l i c a t e s ( M O D E L  p e r c e n t a g e c  ir c u m s c r  a n d  1 ) .  o f  t h e  a n d  d u r i n g  p o p u l a t i o n  s u i t e A N O v A  t h e  v a r i a b i l i t y a n d  t o t a l  s q u a r e s  o f  i n  c o a r s e  R e p e a t a b i l i t y s u m s  m e a s u r e m e n t  i n t e r - p o p u l a t i o n  o n  M O D E L o f  ( V - S S ^ ) .  a v e r a g e d  M O R P H O L O G Y  o n  s a m e  e x p r e s s e d  M E A S U R E M E N T  I N T E R -  I N T E R -  R E P E A T A B I L I T Y  I N D I V I D U A L  P O P U L A T I O N  7 8  4 7  A B X S T O M  3 . 2  8 2  3 3  4 1 4 8  A D X S T O M  0  0  € 5 . 0 6  3 8 . 0 3  4 . 3  0  0  R E S C V N O R E S C Y L O C  M O R P H O L O G Y  1 . 0 5  7 5 . 2 9  4 6 . 6 8  6 4  .8 7  4 0 . 2 1  7 6  6 8  1.7  7 2  .94  0  x  3  8  1.8  R E S C _ Y L E N  T W I G  3 8 . 2 3 3 9  . 2 8  S U I T E  P U L V L E N  7 . 0  8 9 . 4 7  5 4 . 9 0  T I P W I D  3 . 2  7 6 . 3 3  3 6 . 5 1  T I P D E P  6 . 3  6 6 . 3 6  P U L V P U B  0 . 0  9 7 . 1 7  x  4 . 1  8 2 . 3 3  L E A F  A N A T O M Y  1 4 . 7 9 7 4 . 1 9 4 5  . 1 0  S U I T E  N E E D W I D  1.1  9 1  . 2 8  5 6 . 5 1  N E E D E P  7 . 6  9 1 . 6 3  5 8 . 0 6  A B X A N G  2 . 0  8 7 . 2 6  A D X A N G  3 . 0  7 1 . 2 0  4 4  . 9 2  C E N C ' W I D  7 , 5  9 1 . 0 9  4 6  . 4 0  C E N C Y L A T  2 . 0  9 1 . 3 3  5 9  . 4 6  C E N C Y A B X  1.2  9 2 . 9 3  7 3 . 6 9  7 3 . 9 1  1.6  81.77  E N D O N U M  3 . 0  8 4 . 6 8  3 9 . 2 0  P H L E N D  9 . 8  8 8 . 5 3  5 3 . 2 2  X / L E N D  4 . 6  8 5 . 6 8  4 . 0  8 7 . 7 2  C O N L E N  0 . 0  6 6 . 7 5  5 4 . 1 2  C O N W I D  0 . 0  7 2 . 8 2  3 6 . 9 2  S C A L E N  5 . 2  6 2 . 1 8  40.18  S C A L W I D  4 . 3  7 6 . 7 6  4 8 . 3 0  C E N C Y AD X  x C O N E  M O R P H O L O G Y  3 8  . 3 6  4 7 . 0 8 5 3  .7 1  S U I T E  S C A L T A P  9 . 6  5 8 . 7 6  3 5 . 3 8  W I N G W I D  6 . 7  6 9 . 6 1  3 2 . 0 5  W I N G T A P  6 . 2  5 2 . 2 3  1 9 . 9 2  F R E E S C A L  4 . 8  6 7 . 3 1  3 5 . 8 7  B R A C T L E N  9 . 2  7 9 . 2 2  6 7 . 2 1  B R A C T W 1 0  6 . 2  6 7 . 0 2  4 1 . 8 2  B R A C T A P  7 . 7  8 2 . 3 8  6 7 . 9 9  x  5 . 5  6 8 . 6 4  7 4 . 1 9  T W I G  A N A T O M Y  S U I T E  P I T H D I A  8 . 3  3 2 . 1 4  7 0 . 8 4  C O R T H I K  4 . 2  1 3 . 3 5  6 8 . 7 7  P E R I T H I K  2 . 3  3 6 . 4 9  7 9 . 3 3  V B T H I K  6 . 2  3 5 . 7 6  7 7 . 8 3  5 . 3  2 9 . 4 4  7 4 . 1 9  S H C O L E N  -  -  S O . 9 7  S H C O W I D  -  -  4 4 . 2 3  L O C O L E N  -  -  5 5 . 9 6  L O C O W I D  -  -  4 8 . 9 1  -  -  5 0 . 0 2  x C O N E  C O L L E C T I O N  S U I T E  x T O T A L  ^ w i t h o u t  7  f o r m a s  N o . t a x o n o m i c  S U I T E  N E E D L E N  t w i g  a n a t o m y ) 4 . 1  6 2  . 14  o v e r  m e a s u r e m e n t .  b a s e d  v a r i a b i l i t y  r e p e a t a b i l i t y  v a r i a b i l i t y . 1.  i p ti o n .  V A R I A B L E  L E A F  v a r i a b l e  b a s e d  p e r f o r m e d  I n t e r - i n d i v i d u a l A N O V A  a n d  v a r i a b i l i t y ,  r e p e a t a b i l i t y  4 6  . 3 4  o f a  F i g u r e  1.  I l l u s t r a t i o n s  o  A p p e n d i x  I .  =  P j . s i t c h e n s i s .  '  r  c  ,  rnZTcZ  « .  C O N L E N ,  Znllir,  C O N W I D .  c r o s s - s e c t i o n  f v a r i a b l e s  T r i a n g l e s . ' ' '  N u m b e r s U  S  " "  L E A F L E N . f o l l o w s  u s e d .  P . e n g e l m a n n i i :  the  a  '  e  a  :  f i g u r e  S H C O L E N .  S c h e m a t i c c o n v e n t i o n  a b b r e v i a t i o n s s q u a r e s .  b e s i d e  P .  S H C O W I D .  r e p r e s e n t a t i o n o  f C o l l e a u  g i v e n  i  n  g l a u c a -  are~"sampTe~numbers L O C O L o f  E N .  L O C O W I D .  leaf  1 9 6 8 .  PUIV L E N  CENCYLAT  •  «o^V^  C E N C Y W | D  RESCYLEN RESCYLOC PITHDIA  4 ADXSTOM  ABXSTOM  ADX  ABX  SCALTAP SCALWID SCALEN  BR AC  BRACT  TAP  WID  BRACT  LEN  FREESCAL .WINGWID W ING  B 7O10  TAP  26  2.3  Variable The  variation  variables  between  selected  s a m p l e s o f t h e two t a x a .  individuals.  were measured on r e p r e s e n t a t i v e  The n a t u r e  of v a r i a b l e  e x a m i n e d w i t h an ANOVA o f t h e same form a s g i v e n 1; a  however, h e r e y i s a measurement given  of  sample  from an i n d i v i d u a l ,  the i n d i v i d u a l  residual  variation  Those v a r i a b l e s the  variables  further  2 summarizes  inter-population variation (1968,  of t h i s  variation  in leaf  variation  of t h i s  estimates  et a l .  f o r t h e P_;_ a b i e s  Estimates  Table  exceeded  anatomy  2 gives  in this  t h e amount  selected.  some of t h e  study.  Daubenmire  1957; H o r t o n 1959)  encountered  i n cone  (1978) i l l u s t r a t e  morphology.  t h e range of  complex.  variation  form o f ANOVA a r e a l s o 1981).  variation.  f o r further  3 illustrates  (Garman  of i n t e r - i n d i v i d u a l  (Falconer  effect  v a r i a t i o n and  anatomy e n c o u n t e r e d  Pravdin,  made on  and e i s  that  removed t w i g  f o r the v a r i a b l e s  some o f t h e v a r i a t i o n  an a s i d e ,  rule  Figure  1  variation  were r e t a i n e d  inter-individual  variation .  variable  variation),  was  i n MODEL  intra-individual  consideration.  1972, 1974) and o t h e r s  illustrate  results  variation  Application  from  case,  w i t h an i n t e r - i n d i v i d u a l  inter-individual  Figure  As  or, in this  above  A i s the hypothesized  (inter-individual  the i n t r a - i n d i v i d u a l  measurement.  of  tree  f o r a given  variation  b a s e d on t h e  known a s  As a s t a t i s t i c  from  repeatability ANOVA,  Between p o p u l a t i o n v a r i a t i o n i s a s s e s s e d w i t h a model o f t h e same form a s i n MODEL 1 g i v e n above f o r i n t e r - i n d i v i d u a l v a r i a t i o n , t h e e x c e p t i o n h e r e b e i n g t h a t "A" i s t h e e f f e c t o f the p o p u l a t i o n ( i n t e r - p o p u l a t i o n variation). 1  27  F i g u r e 2. Summary o f v a r i a b l e v a r i a t i o n b a s e d on a l l individuals without r e f e r e n c e to taxonomic c i r c u m s c r i p t i o n . %S5'.ndividcui' amount o f v a r i a t i o n b e t w e e n i n d i v i d u a l t r e e s . %SS p o f u U + i e r x amount o f v a r i a t i o n b e t w e e n i n d i v i d u a l p o p u l a t i o n s . Variable suites: T - twig anatomy; C - cone c o l l e c t i o n ; P - twig morphology; S - cone s c a l e m o r p h o l o g y ; N - l e a f m o r p h o l o g y ; A - l e a f anatomy Values g i v e n a r e t h o s e i n T a b l e 2.  100i c o  AA  P  AP  c.  SS  5  o  SA  o.  N  A  C  A  N  0  repeatability the  i s i n t e r p r e t e d as a  heritability  heritability,  negate  (Falconer  nature  practical  means  biological  philosophical "components only  stressing  means  the s p e c i f i c  to  1961); Owing  (Sokal  1978) a s s u m p t i o n s  genetic  to the size  sense  of  and  traits.  and Rohlf  1969),  1983) a n d  engendered  repeatability  that  does  i s t h e most  heritabiity.  aspects  of broad  however, t h i s  1963; J a c q u a r d  u s e o f ANOVA,  of  cautious  the h e r i t a b i l i t y  of a s s e s s i n g  estimate  approximation  repeatability  1981; R o b i n s o n  of variance"  appropriate  utility.  the s t a t i s t i c a l  (Kempthorne  100  sample-specific  i s subject  of estimating  (Falconer  I'  As an  of c o n i f e r s ,  considering  I  1981; Z o b e l  i t scomparative  perennial  Indeed,  of a t r a i t .  repeatability  interpretation not  I  VeSS individual  some  by t h e  may  Rather  be t h e than  researchers  28 F i g u r e 3. I l l u s t r a t i o n of range of v a r i a t i o n encountered f o r leaf anatomy. Numbers i d e n t i f y I n d i v i d u a l t r e e s l i s t e d i n Appendix I I .  R enqelmonnii  |05mm |  a s s o c i a t e with h e r i t a b i l i t y , r e p e a t a b i l i t y  will  be used  here  simply as a r e l a t i v e e x p r e s s i o n of the amount of inter-individual v a r i a b i l i t y . The  inter-individual  variation  values given  comparable to those given f o r r e p e a t a b i l i t y other morphological  i n Table 2 are  estimates  shown by  c h a r a c t e r s i n other c o n i f e r o u s s p e c i e s  29  (Table  3).  These r e p e a t a b i l i t y  than p u b l i s h e d species  heritabilities  (Hattemer  of h e r i t a b i l i t y heritabilities physiological differences  effects  considered are  the  as  i n the  are  variables in tree  reflect  broad  size  simply  reflect  of  the  the  and  basic  heritability  result fact  of  that  (1965) r e s u l t s ,  presented  l a r g e , but  i t should  be  large geographic Engelm. may  in sampling  (Sorensen  example of  population-specific  be  nature  and  genetic  estimate  could  area.  small  as  in Table that  3,  the  sample  are  these  samples  Sorensen's data a result  T h e s e may  genetic  for  of s e l e c t i o n serve  as  (1961) r e m a r k s c o n c e r n i n g  of  be  3.  noted  1964).  Zobel's  as  hypothesized t r a n s - i n d i v i d u a l  Andersson's  convenient  However,  a more h e t e r o g e n o u s  in Table  imposed  over-estimation  physiological variables.  been c o n s i d e r e d  elliottii  the  sense.  have not  were from a v e r y  generally larger  most commonly c a l c u l a t e d f o r g r o w t h  s t u d i e s - the  particularly  Pinus  probably  between m o r p h o l o g i c a l  interpreted other  and  for other  variables i t could  Additionally,  than  1963)  estimates  a the  i n t e r p r e t a t i o n of  such  estimates.  Fig.  As  a group,  2)  are  vegetative  the  cone c h a r a c t e r s  more v a r i a b l e w i t h i n variables.  The  British  Columbia,  selected  subject  to l e s s  vegetative  characters.  individual  i n f e r e n c e drawn  southwestern are  an  measured here  This  the  than  is that,  reproductive  genetic  (Table are  in Picea  of  characters  c o n t r o l than are  conclusion  2,  the  i s c o n t r a d i c t o r y to  general  expectation  that  reproductive  characters  subject  to environmental  perturbation  than are  are  less  vegetative  the  30  T a b l e 3. Inter-individual variation, repeatability estimates, reported f o r m o r p h o l o g i c a l c h a r a c t e r s in other c o n i f e r o u s t r e e s p e c i e s . V a l u e s r e p o r t e d i n t h e t a b l e a r e p e r c e n t a g e o f t o t a l v a r i a t i o n due t o d i f f e r e n c e s between i n d i v i d u a l t r e e s . V a l u e s a r e c a l c u l a t e d from c i t e d a u t h o r i t i e s and r e f e r t o the t o t a l i n t e r - i n d i v i d u a 1 v a r i a t i o n w i t h o u t r e s p e c t to h y p o t h e s i z e d t r a n s - i n d i v i d u a 1 s o u r c e s or var i at i on.  Picea glauca (Khalil 1974) cone weight 86 . 16 cone l e n g t h 95 .07 cone width 7 1 . 33 scale length 78 . 74 s c a l e width 62 . 1 1 78.68  A b i e s b a l s a m e a ( L e s t e r 1968) cone l e n g t h 70.00 scale length 85.00 seed length 77 .00 bract length 85.00 stalk length 76 .00 77 . 80  P i c e a mar i a n a ( P a r k e r e t aj_ . cone l e n g t h cone diameter scale length scale width scale concavity dark band w i d t h 1 i g h t band w i d t h seed length seed+wing l e n g t h leaf width leaf thickness twig pubescence l e a f apex shape r e s i n canal separat ion vascular bundle diameter l o n g e s t bud scale length l a t e r a l bud 1ength x 62  P i c e a ab i e s  1965)  P i n u s e l 1 i o t t i i ( S o r e n s e n 1964) leaf length 48 .67 f a s i c l e volume 54 .95 41 . 83 leaf divergence sheath length 78 . 74 77 83 bud s c a l e l e n g t h ~ 60 . 40  70 0 0  P i nus  36 . 80 75 . 80 71 . 30 .59  P s e u d o t s u g a menz i e s i i (Chen, et a [ . , u n p u b l . ) x n e e d l e anatomy  (Andersson  1983) 67 . 30 68 . 70 70 . 90 71 .90 61 . 40 70 .00 67 . 60 73 .80 70 .60 48 . 40 49 . 20 6 0 .'60 30 . 30  86.32  cone weight 90.01 cone 1ength 91.67 .•'seeds/ c o n e 81.22 seed weight/ cone 85.29 x 86 . 17  kes i ya ( B u r l e y a n d B a r r o w 1972) leaf length 76.84 % 3-leaf f a s i c l e 67.96 leaf/ fasicle 67.88 "x 70.89  P i c e a mar i ana ( K h a l i l 1975) cone l e n g t h 65.93 cone w i d t h 64.91 "x 65.42  31  (Stebbins the  1950;  Heywood  o b s e r v a t i o n s of P a r k e r  Taylor  (1959).  variables than  3.  D a v i s and  The  i s that  1973).  This also  e t a l . (1983)  (see T a b l e  general trend in Table  t h e y have a l a r g e r  contradicts 3)  and  3 of cone m o r p h o l o g y  inter-individual  variation  vegetative variables.  Samples and  sampling.  Once v a r i a b l e s attention relate  were s e l e c t e d ,  to the o r i g i n  of  the  that  necessary  samples and  to the v a r i o u s a p r i o r i  organization  i t was  how  these  trans-individual  have been h y p o t h e s i z e d ,  to  direct  samples  levels  of  i . e . populations  and  taxa.  3.1  Study  area.  The  main a r e a  of  Columbia  including  both  (Fig.  4A).  address study  The  specific  area.  P.  the Coast  situations  better  of  areas  the  sought  southwestern  effect  study  was  to  i n t h e main p o r t i o n  of  of b o t h  i n the  permits study  of s a m p l e s the  third a  to species,  better  area.  from  relation  the  to  s p e c i e s and  of a p o s s i b l e  to: identify  Ranges  expanded  sampling  inclusion  British  Cascade Mountain  a r e a s were s e l e c t e d  the v a r i a t i o n  the v a r i a t i o n  Specifically, additional  arising  Such s u p p l e m e n t a l  understanding  and  i n c l u d e d i n the  t h e e x i s t e n c e and  glauca.  i s mainland  These supplemental  circumscribe consider  area  study  these between  32  F i g u r e 4. Maps of l o c a t i o n s of samples arid study area. A - study area; B - common garden samples of s i tchens i s ; C - n a t u r a l l y growing c o l l e c t i o n s from southern p o r t i o n of range of P_^ s i tchens i s ; D samples of engelmanni i o u t s i d e of study area. Tree and p o p u l a t i o n numbers correspond to those g i v e n i n Appendix I I .  A  B  C  D  p. enqelmanni i and Pj_ g l a u c a ; i d e n t i f y the r e l a t i o n between P. enqelmanni i  i n the study area t o that reported f o r a d i s j u n c t  l o c a t i o n of the P^ enqelmanni i on the Olympic 1970;  Peninisula  (Sharpe  H i t c h c o c k , et a l . 1969) ( F i g . 4D); i d e n t i f y the r e l a t i o n  between E\ enqelmanni i  i n the study area to that  and d r i e s t p o r t i o n s of the i n t e r i o r of B r i t i s h ( F i g . 4D); and, i d e n t i f y the r e l a t i o n between the study area and that  i n the wettest  Columbia s i t c h e n s i s in  i n more s o u t h e r l y l o c a t i o n s  ( F i g . 4B, C)  33  The  possibility  P. g l a u c a was  Knight  necessary  P. g l a u c a , P. g l a u c a  Inlet  could  t o examine  albeit  the r e l a t i o n  The s u s p e c t e d  the K l i n k a k l i n i  west  o f T a t l a Lake  reconnaissance  Such an o c c u r r e n c e  s i t u a t i o n s have been (Daubenmire  reported  1968; Garman  a l . 1982; Hanover and W i l k i n s o n  1977;  F a l k e n h a g e n a n d Nash  Coola  River  Several  fully  specific  o f P_^ g l a u c a  Inlet  along the since  f o r more n o r t h e r l y 1957; Roche  inlets:  1969; Coupe,  1970; Copes a n d B e c k w i t h  (Garman  2  (Roche  previous  i s not unexpected  1978); Dean R i v e r  ( P o j a r ) ; Nass R i v e r  River  during  i n d i c a t e d the presence  low e l e v a t i o n p a s s t o K n i g h t  River.  Skeena R i v e r  nature  of  o f Remote C r e e k a l o n g  the r e l a t i v e l y  Bulkley  occurrence  f o r trees east  1  et  engelmanni i and  form and cone  research  similar  between  As s u c h i t  i s b a s e d on t h e crown  A d d i t i o n a l l y , personal  Klinaklina  a priori.  Inlet  River.  in  and/ or i n t r o g r e s s i o n of  n o t be d e n i e d  superficially.  i n Knight  morphology  of t h e o c c u r r e n c e  (Pojar ); 2  1957; Roche  Bella  1969); a n d  1969).  collections  of i n t r a - i n d i v i d u a l  were made t o d e s c r i b e t h e  variation.  These a r e d e s c r i b e d  more  i n Chapter I I I . Samples were c o l l e c t e d  under n a t u r a l  from a v a r i e t y o f common g a r d e n  situations.  r e p r e s e n t a t i v e s o f P^ s i t c h e n s i s from range of t h e s p e c i e s  s i t u a t i o n s as w e l l as  were sampled  First,  throughout  the n a t u r a l  i n a common g a r d e n  situation  34  in  the Chilliwack  from  seed.  Valley  Comparison  grown t r e e s may p r o v i d e population useful  under are  given  Fredericton,  this  to nursery  surrounding  comparison  from o t h e r  may p r o v e  common  garden  h e r e b a s e d on p l a n t s c o l l e c t e d l o c a t i o n s and  common g a r d e n s were sampled; George, B r i t i s h  New B r u n s w i c k .  information  collections  provide  Secondly  regarding  the p r a c t i c a l i t y  hybrids.  More s p e c i f i c  i s provided  In a d d i t i o n data  1  and t h e o t h e r a t nurseries,  p a r e n t a g e were s a m p l e d .  information  hybrids.  Columbia  one a t Red Rock,  In b o t h o f t h e s e  2  o f known h y b r i d  al.  Further,  More e x a c t  individuals  study,  trees  grown  i n Appendix I I .  of P r i n c e  trees  had been  i n t o the f a c t o r s  the r e s u l t s reported  conditions.  Two o t h e r south  insight  to the r e s u l t s presented  natural  The t r e e s  of n a t u r a l l y o c c u r r i n g  differentiation.  in relating  research  ( F i g . 4B).  3  concerning  the d e s c r i p t i o n of  these c o l l e c t i o n s provide of r e c o g n i z i n g  information  about  Such  information  naturally  occurring  the parentage of these  i n Appendix I I .  t o the t r e e s  from specimens  (1982) were a l s o  specifically  sampled  sampled p r e v i o u s l y  for this  in Klinka, et  used . 3  B r i t i s h Columbia M i n i s t r y of F o r e s t s , Vancouver F o r e s t Region, R e s e a r c h B r a n c h - V a n c o u v e r 1981. B r i t i s h Columbia M i n i s t r y of F o r e s t s , P r i n c e Rupert F o r e s t Region, Smithers. U l f B i t t e r l i c k ; B r i t i s h Columbia M i n i s t r y of F o r e s t s 1  2  3  G y u l a K i s h ; B r i t i s h Columbia M i n i s t r y of F o r e s t s . Dan F o w l e r ; C a n a d i a n F o r e s t r y S e r v i c e . p e r m i s s i o n of K a r e l K l i n k a ; B r i t i s h Columbia M i n i s t r y of Forests. 1  2  3  35  3.1.1  Climate. In t h e  level  and  reported to the  study  2100  area  s a m p l e s came from e l e v a t i o n s between  m ASL.  w i t h i n the  Some of  study  extreme t o p o g r a p h i c  alteration  a s s o c i a t e d with  values  w e l l be  can  reported. climatic of  the  area  The  variation  are  indicated  such  to d i f f e r  r a t h e r than given  v a r i a b l e s should  be  responsible for l i m i t i n g  a given  site.  roughly  with  overview  (1978a,b) and of c l i m a t i c  Figure with  that P. any  to corroborate  P_j_ s i t c h e n s i s  i s a low  engelmani i i s a h i g h  apparent. areas. from t h e  the  It should highest  be  study  the  and  noted  the of  by  a  (1976).  general  area.  of  samples  These  from t h e  literature whereas  s p e c i e s ; however,  Daubenmire  "hybrids" occur  i n the  Fletcher  elevation, coastal species  1955;  P i c e a at  corresponds  elevation.  impression  only  Climate  r a n g e of d i s t r i b u t i o n  (Wright  P r e d i c t a b l y , the  observation  f o r the  these  the c l i m a t e  elevation, interior  discontinuity  survival  briefly  in  t h a t the c o l l e c t i o n s  1968)  i s not  intermediate o f F\  sitchensis  e l e v a t i o n s came from more n o r t h e r l y a r e a s  t h a t c o r r o b o r a t e s Daubenmire's  of  description  being  longitude.  those  impression  (1976b) p r o v i d e  to longitude, l a t i t u d e ,  serve  edaphic  O'Driscoll  variation  of  and  i s summarized  5 illustrates  respect  figures  variation  elevation, latitude,  r a n g e of P.? s i t c h e n s i s Schaeffer  the  from  However  i n t e r p r e t e d as g r o w t h and  Owing  climatic  accurate  location.  4.  climatic  4 g i v e an  p r o v i d i n g an  sample  not  local  substantially  in Table  factors  Further,  and  conditions  in Table  topography, a c t u a l  presented  c l i m a t e a t any  climatic  r a n g e of c l i m a t i c  variation  expected  values  the  sea  (1968) r e p o r t  -  that  an  T a b l e 4. Summary o f some a n n u a l a v e r a g e c l i m a t i c v a r i a b l e s r e p o r t e d f o r t h e Study a r e a and a d j a c e n t a r e a s . SSS - s t a n d a r d P_^ s i t c h e n s i s; SSP - p u t a t i v e P. s i t c h e n s t s ; SXE - " h y b r i d " ; ESP - p u t a t i v e P_^ enge1mann i i ; ESS s t a n d a r d P_^ e n g e 1 mann i i ; WSS - s t a n d a r d g 1 auca . - no d a t a a v a i l a b l e . S o u r c e s : U.S. Dept. o f Commerce ( 1 9 7 5 ) ; E n v i r o n m e n t C a n a d a ( 1 9 7 3 ; 1975a, 1975b).  STATION  ID  STUDY AREA B R I T T A N I A BEACH PORT A L I C E PORT HARDY RIVER JORDAN SOUAMISH TOFINO VANCOUVER - UBC  ELEV (m)  RAIN (mm t  SNOW  T  X  (C  )  min T (C )  maxT (C )  FROST ( day )  SSS  50 20 25 5 2 30 95  1985 3152 1660 1962 1916 3020 1258  782 587 706 2 18 1455 4 17 490  10 .0 9 .4 7. 9 9 .0 8 .9. 9. 2 9. 8  6 4 5 .7 4 .7 5. 2 4 7 5 .7 6 6  13 5 13 . 1 1 1. 2 12 .8 13 . 3 12 . 7 13 . 1  43 33 67 44 84 49 33  SSP  10 190 50  1636  f 101  1029 1 107 162 1  10 . 2 8. 9 9. 7  5 .7 4 .7 5 .1  14 .8 13 . 2 14 . 3  57 80 77  SXE  730  837  5936  5. 7  0. 4  10 .9  176  PEMBERTON MEADOWS SKAGIT RIVER  ESP  240 560  742 802  2825 3228  7. 2 7 .8  2 .1 1 .9  12 . 3 13 8  128 165  ALLISON  ESS  1470  486  9652  1 .8  -3 . 7  7. 5  255  ESP  570 470 980 6 30 500  218 • 189 204 7 17 703  752 602 1669 7364 4 115  7 .9 9. 7 1. 8 4. 2 7. 2  1 .7 4 .2 -5 . 8 -0. 9 1 .7  ESS  1305 1930  302 2 14  3106 3297  3. 2 2. 2  -3 . 3 -3 . 4  9 6 7, 7  SSS  60 120  9. 8  7 3  17 . 1  _  14 13  30 30 30  2056 15 19 2097  -  1 1 . 8 10 . 8 1 1 . 2  8.3 7 .4 6.3  15 14 16  -  -  663  193  5 8  0. 8  10. 8  SAMPLES SSS 10 15  1440 2350  -  4 8 3 4  - 1 2. -5 . 0  12 . 7 1 19 .  JUNEAU  10  1389  -  6 .0  -2 . 6  14 . 1  TERRACE. B.C. SANDSPIT B E L L A COOLA CHILLIWACK  70 10 20  917 1 182 1358  1816 785 1750  CHILLIWACK HANEY RESEARCH HOPE ALTA  FOR.  LAKE  PASS  OUTSIDE STUDY AREA HEDLEY KEREMEOS KLEENA KLEENE MICA DAM REVELSTOKE CHUTE LAKE HEDLEY MINE SEOUIM, WASH. ELWHA S T A T I O N .  WASH.  BROOKINGS. ORE. BANDON. ORE. CANARY. ORE.  SSS  OTTAWA,  WSS  ONTARIO  CHILLIWACK RIVER NURSERY KODIAK IS. ALASKA CORDOVA BAY  FORKS,  WASH.  CLOVERDALE . ORE . CANARY BROOKINGS  1448  -  -  -  6. 7 2 .7 7. 9 5 .1 7. 4 2 .8 < SEE ABOVE >  14 15 9 9 12  . . . . .  1 1 5 3 6  3 ,3 .3  10. 8 10. 6 12 .0  158 1 14 258 184 150 228 231  -  162  -  137 63 1 24  120  2956  -  9 5  3 .6  15 .6  -  5  2 147  -  10 9 < SEE < SEE  5 .9 ABOVE ABOVE  15 . 7  -  37 F i g u r e 5. E l e v a t i o n a l , l a t i t u d i n a l , and l o n g i t u d i n a l d i s t r i b u t i o n of samples. SSS - s t a n d a r d s i t c h e n s i s ; SSP - p u t a t i v e P_^ s i t c h e n s i s ; SXE - " h y b r i d " ; ESP - p u t a t i v e P_^ e n g e I mann i i ; ESS - s t a n d a r d P . e n g e 1 mann i i ; IESP - p u t a t i v e P_^ e n g e 1 mann i i i n t e r i o r ; IESS standard e n g e 1 mann 1 i i n t e r i o r .  2100  ELEVATION  LATITUDE  LONGITUDE  52 2  I  JJlTflfJI tr)  <0  Of  it/  <(/  48  0  128 2  I  I  I  I  I  I  115 B  ty ty  t h i s s p e c i e s o c c u p i e s a wider e l e v a t i o n range a t h i g h e r latitudes.  3.1.2 Edaphic  environment.  Parent m a t e r i a l s i n t h e study a r e a a r e p r i m a r i l y coarse t e x t u r e d q u a r t z - and g r a n o - d i o r i t e s ( H o l l a n d 1976).  Notable  e x c e p t i o n s a r e t h e l i m e s t o n e and sedimentary parent m a t e r i a l s o f the C h i l l i w a c k V a l l e y and some of i t s a s s o c i a t e d d r a i n a g e s . In a l l a r e a s these parent m a t e r i a l s a r e o v e r l a i n , t o v a r i o u s degrees, by an a c c u m u l a t i o n of g l a c i a l m a t e r i a l s (Ryder  t i l l s and c o l l u v i a l  1978).  S o i l s where P i c e a occur a r e p r i m a r i l y those a s s o c i a t e d w i t h f l u v i a l and a l l u v i a l l a n d f o r m s .  W e l l a e r a t e d and p e r v i o u s  c u m u l i c r e g o s o l s and l e s s a e r a t e d and s a t u r a t e d g l e y s o l s a r e the most f r e q u e n t .  On t h e o u t e r c o a s t , P i c e a may occur i n b r a c k i s h  c o n d i t i o n s a s s o c i a t e d w i t h sandy s o i l s of advancing beach f r o n t s (Cordes  1972).  38  Picea sites. Picea  o c c u r s on a v a r i e t y  In t h e n o r t h e r l y  of  coastal  o c c u r s on v a r i o u s d e g r e e s  "folisols" area,  (Pojar  P i c e a may  luvisols.  be  With  interior,  1982).  on  increasing  P i c e a i s found  t y p e s on more  portion  of  the  interior  portion  v a r i o u s degrees elevation  on  on  of  both  p o o r l y developed  upland  study  of h u m i f i e d p o d s o l s  In t h e  found  soil  area,  and  of t h e  study  illuviated  t h e c o a s t and podsols  the  and  brunisols. Like  climate,  geographic  location.  environment given  An an  - they i n the  collection  of the  classification  properties  regime  of t h e e d a p h i c relative  of  specific  w i t h E\  moisture  index  The  the  Table  to  textural  entirely  example, what c o n s t i t u t e s  of  in Table  independent  noted  illustrate "hybrids"  that  of l o c a l  a subxeric site  on  The  moisture  5 serves to  I t s h o u l d be  samples  5.  respect to  o f P^_ e n q e l m a n n i i and  sitchensis.  Briefly,  landscape  disposition  i s given  samples w i t h II.  paid  is a  by  f o r each  i s p r o v i d e d by  attention  regime  amplitude  i s not  i s provided  of m o i s t u r e  availability  soil.  i n Appendix  compared  environment  availabity  of m o i s t u r e  individual  edaphic  edaphic  possible  i n Walmsley, et a l . (1980).  with secondary  i s given  the wider  For  s e r v e t o d e s c r i b e the  respect to moisture  disposition  of the  with  area.  outlined  of the  roughly  t o be a c c u r a t e f o r any  Such q u a n t i f i c a t i o n  classification  classification  with  simply study  site.  corresponds  F u r t h e r , the d e s c r i p t i o n s  approximation  indication  this  variation  g i v e n h e r e a r e not meant  site  variation  edaphic  this  climate.  the outer c o a s t  39  T a b l e 5. regime  D i s t r i b u t i o n of samples w i t h r e s p e c t to m o i s t u r e s c h e m e f o l l o w s W a l m s l e y , e t al_. ( 1 9 8 0 ) .  XERIC  SUBXERIC  IDENTIFICATION P.  SUBMESIC  MESIC  sitchensis STANDARDS PUTATIVES  "HYBRIDS'  P.  enge1mann i i STANDARDS PUTATIVES  16  S E L K I R K MTNS. STANDARDS PUTATIVES  10 10  regime.  SUBHYGRIC  Moisture  HYGRIC  TOTAL  4 13  36 38  21 8  62 65  10  25  51  44  133  10 27  51 18  10 36  27  12  71 105  10 69  15  515  might  well  be s u b h y g r i c i n t h e i n t e r i o r .  A description not  necessarily  critical sampled of  of the e d a p h i c  reflect  study.  edaphic v a r i a b l e s  subject  the edaphic  description to a large  Additionally,  i s such  (Courtin,  besides that  p r e s e n t today  environment  t o t h e e s t a b l i s h m e n t and e a r l y for this  detailed  environment  that  does  was  growth of the t r e e s the l a t e r a l  variability  e_t a_l. 1983) t h a t  g i v e n here  is liable  more  t o be  measurement e r r o r .  3.1.3 A s s o c i a t e d v e g e t a t i o n . In a d d i t i o n  t o c l i m a t i c and e d a p h i c  many documented d e s c r i p t i o n s Picea. sample et al.  References sites  specific  c a n be f o u n d  a l . (1982);  Krajina  (1982); K l i n k a ,  v a r i a t i o n , there are  of v e g e t a t i o n found  growing  with  t o t h e s t u d y a r e a and a u x i l i a r y  i n :Krajina,  (1969); K r a j i n a  e t a _ l . ( 1978);  Krajina,  (1965); K l i n k a , e t  e t a l . (1980); K l i n k a ,  e t a l . (1979);  Jones  40  and  Annas As  ( 1 9 7 8 ) ; F r a n k l i n and D y r n e s s  with climate  vegetation  and e l e v a t i o n ,  approximation trees,  i s provided  biogeoclimatic  of  in this  zone"  longitude,  e_t a l .  study  indicated  is  given  i n Table  the edaphic  vegetation survival  may  from  establishment  to these a e r i a l l y  The b i o g e o c l i m a t i c 6.  smaller  d i s p o s i t i o n o f samples  The d i s p o s i t i o n o f i n d i v i d u a l samples  environment, contemporary that  associated  contributing  herb  species  and d e v e l o p m e n t  vegetation,  Evidence of recent study area.  to the  glaciation  of  1978).  seedlings.  of upland  soils  i s dominated  Holland  With  the exception  throughout the  have d e v e l o p e d  (1976) and Ryder  of the t r e e s  from  by t h e e f f e c t s o f  a r e v i e w a n d summary of t h e g e o m o r p h i c  area.  upon t h e  and c l i m a t i c h i s t o r y .  and t h e l a n d s c a p e  (Ryder  The  t r e e may be v a s t l y  g l a c i a t i o n i s pervasive  The m a j o r i t y  tills  on a l a r g e  t h e e f f e c t o f t h e same h e r b s p e c i e s  3.1.4 G e o l o g i c a l ,  study  e t a l . 1981;  and c h a r a c t e r i s t i c s o f t h e i n d i v i d u a l t o d a y .  different  provide  precise  e t a l . 1983; C o u r t i n ,  n o t be n e c e s s a r i l y  of a given  glacial  sample t o a  i n Appendix I I .  Like  effect  each  forest  1979), however t h e wide g e o g r a p h i c d i s t r i b u t i o n  impractical.  is  primarily  by a s s i g n i n g More  An  i s possible (Mitchell,  t h e samples makes a s s i g n m e n t  syntaxa  and l a t i t u d e .  1969).  circumscription  (1977).  i n t e r d e p e n d e n c e of  forest vegetation,  (Krajina  a l . 1981a, b; U t z i g ,  Klinka,  i s a strong  of a s s o c i a t e d  "biogeoclimatic  et  there  ( 1 9 7 3 ) ; Rowe  (1978)  h i s t o r y of the  sampled  i n Oregon  41  T a b l e G. D i s t r i b u t i o n of samples w i t h r e s p e c t to b i o g e o c l i m a t i c zones. CDF - c o a s t a l d o u g l a s - f i r z o n e ; CWH - c o a s t a l w e s t e r n h e m l o c k z o n e : MH m o u n t a i n h e m l o c k z o n e ; ESSF - e n g e l m a n n s p r u c e s u b a l p i n e f i r z o n e ; IDF i n t e r i o r d o u g l a s - f i r z o n e ; IWH - i n t e r i o r w e s t e r n h e m l o c k z o n e . A s s i g n m e n t t o b i o g e o c l i m a t i c z o n e b a s e d o n C o u r t e n , e_t aj_. ( 1 9 8 1 ) ; K l i n k a , e t a±. ( 1979, 1 9 8 0 ) ; M i t c h e l l , e t a]_. ( 1 9 8 1 a , b ) ; a n d , U t z i g , e t al_. ( 1 9 8 3 ) .  IDENTIFICATION  s i tchensi s STANDARDS PUTATIVES  P.  BIOGEOCLIMATIC ZONES CWH MH ESSF  CDF  13 1  "HYBRIDS"  116  15  engelmanni i STANDARDS PUTATIVES  7  7  C a l i f o r n i a , a l l trees  ice during  summary  sampled  came  71 56  35  10 50  5  (1978) and W o l f e  of the pre-Quaternary  14  from an a r e a  the l a s t c o n t i n e n t a l g l a c i a t i o n  Daubenmire  IWH  49 64  S E L K I R K MTNS. STANDARDS PUTATIVES  and  IDF  (Ryder  (1969) p r o v i d e  vegetation  covered  1978).  r e v i e w s and  of t h e a r e a .  Hebda  (1983) p r o v i d e s  a summary  for  Mack, e t a l . ( 1 9 7 6 ) , Hansen ( 1 9 5 5 ) , and Hebda  the c o a s t .  (1982) p r o v i d e Following to  summaries o f v e g e t a t i o n t h e most r e c e n t  or hypsithermal  was most p r o n o u n c e d  the c l i m a t e  than a t p r e s e n t .  period  i n the i n t e r i o r  occurred  areas  of the c o a s t  (Alley  (Barnosky  change  i s assumed  This  around  6000 yBp and  1976; Mack, e t  1976; Hansen 1955; D a u b e n m i r e 1975; Hebda  rainshadow  vegetation  changes f o r the i n t e r i o r .  glaciation  have become warmer and d r i e r  xerothermic  al.  of t h e p o s t - g l a c i a l  by  1981).  1982) and i n t h e The  hypothesis  42  of  the existence  of the x e r o t h e r m i c  more c o a s t a l a r e a s  The has  vegetation  been d i s c u s s e d  constitutes history. trends  3.1.5  the  today  i n respect  i n western North  i t i s necessary  regarding  to place  important  the occurrence  taxa i n of P i c e a  especiallyi f  forthcoming.  the Quaternary  h i s t o r y of  i s the p o s t - g l a c i a l p e r i o d . b a s e d on p a l y n o l o g i c  These  evidence.  As  o f P\_ s i t c h e n s i s a n d P_;_ e n q e l m a n n i i on p o l l e n s i z e i s  (Mathewes  retrace  the h i s t o r y of both taxa  g e n e r a l l y agreed south during  1973; W i l s o n  that  glacial  these  1963),  like  period.  It  many o t h e r s ,  For the c o a s t a l area  to is  retreated  species where  have been made p r i m a r i l y a t low e l e v a t i o n s , o f P^ s i t c h e n s i s .  Picea  In t h e  p o l l e n has been assumed t o be e i t h e r  P. e n g e l m a n n i i o r P^ g l a u c a , contemporary  this  e p i s o d e s a n d t h a t more montane  has been assumed t o be t h a t Picea  i t i s impossible  during  species,  t o lower e l e v a t i o n s .  investigations  interior,  climatic  America.  perspective,  in considering  tenuous  descended  history  t o a c t u a l dates but t h e  r e l a t i o n s h i p s a r e t o be  i n the study area,  separation  concerning  t h e v a r i a t i o n o f t h e two h y p o t h e s i z e d  interpretations are generally  pollen  inferences  i n t o some s o r t o f h i s t o r i c a l  Particularly  The v e g e t a t i o n  area  are similar.  examining  inferences  Picea  for derived  of Picea  study area  h i s t o r y of the study  by numerous w o r k e r s .  The a c c o u n t s d i f f e r  History  remains enigmatic f o r  1973).  and r e - v e g e t a t i v e  the b a s i s  reported  In  (Mathewes  period  species.  d e p e n d i n g upon t h e p r o x i m i t y o f  The s a n c t i t y o f t h e s e  i n t e r p r e t a t i o n s has  43  been c h a l l e n g e d by t h e r e c e n t Puget  lowlands  Daubenmire Picea  identified  important  i s s u e i n the Quaternary  North  America  refugia  of the s p e c i e s .  P_^ s i t c h e n s i s  (reviewed  this  frequently  invoked  variation tendered related  concerns  the h y p o t h e s i s  i n Daubenmire  1984).  hypothesis  In b o t h  by P o r s i l d  A similar  ( i n Garman  o f P_;_ s i t c h e n s i s  differentiation  (Illingworth  1976).  For present areas,  then  1968) and P^ g l a u c a cases  conclusions The h y p o t h e s i s i s  hypothesis  intra-glacial  i n the  has been  Post-glacial  A  genetic  (Yeh a n d E l - K a s s a b y  during p o s t - g l a c i a l  t h e c o a s t and i n t e r i o r ,  Picea pollen and r a i n  1980) and  migration  d i f f e r e n t i a t i o n of been h y p o t h e s i z e d  Picea pollen  i n the o l d e s t  by  i s generally  sequences.  For the c o a s t a l  d e c l i n e s g r a d u a l l y t o the p r e s e n t .  sha.dow a r e a s ,  increases after  hypothesized  tendered  (1974).  and abundant  interior  has been  discontinuities  P. e n q e l m a n n i i d u r i n g m i g r a t i o n has a l s o Daubenmire  glacial  1957) f o r P^ e n q e l m a n n i i .  i s s u e i s the h y p o t h e s i z e d  subsequent  of P i c e a i n  of a  remains e n i g m a t i c .  to e x p l a i n apparent  o f P_;_ s i t c h e n s i s .  depauperization  of the nature of  history  Such an h y p o t h e s i s  (reviewed  in Critchfield  concerning  review  1981).  lowlands.  western  for  of m a c r o f o s s i l s i n the  a s P_;_ e n q e l m a n n i i ( B a r n o s k y  (1968) p r o v i d e s a f u r t h e r  i n t h e Puget An  findings  Picea  5000 yBP, t e n d i n g  occurrence  initially  d e c l i n e s and  to support the  of a h y p s i t h e r m a l  In t h e  interval.  44  3.2  Trans-individual R e g a r d l e s s of  variation,  often  convention  rather  o c c u r r e n c e of that  the  hypothesized  s t i p u l a t e d by  balanced  v a r i a t i o n was  3.2.1  circumscription  Where t h e r e  not  location a population  within  of  years),  and  were g r o w i n g  conditions. population pollen et  This  (Wright  and  Colwell  p o l l i n a t i o n and  T r e e s were c o r e d individuals  had  where i n c r e m e n t height  was  potential and  not  as  coring  u s e d as  an  between  population  to  those  defined  this  of  the  1951;  Ibe  same age  (Mair  spatial  1973;  a  (+/-  and  definition  1983;  at  i f i n d i v i d u a l s were  of  available literature  Stern  10  edaphic a  pertaining  Silen  restriction  that  sampled.  age  not  1962; and  to  Wang,  Roche  minimizes  long  d i f f e r e n t aged  Under c o l l e c t i o n  estimator.  i n the  obviously  possible,  data  environmental  trees  i s such  samples.  was  a check  was  heterogeneity  the  dispersal.  been  meso-topographic  distance  This  or  possible.  individual trees  of  seed d i s p e r s a l  in forest trees.  distance  part  of  Columbia  in s i m i l a r physiographic  upon t h e  1953;  nature  for a l l sources  approximately  spatial  i s based  a l . 1969)  1974)  each other,  the  British  of  were a number of  particular 30m  reality,  sampling  trans-individual  Population  samples  s o u r c e s of t r a n s - i n d i v i d u a l  in southwestern  and  of  theoretical considerations  than b i o l o g i c a l  Picea  systematic  circumscription  diameter  at  Such s a m p l i n g  situations breast reduces  the  r e l a t e d t o d i f f e r e n t ages conditions.  The  restricted  also confines  the  definition  of  i n d i v i d u a l s that  are  p o t e n t i a l l y able  to  45  cross-pollinate 1974)  and  probably  parental  expected  t o be  lower  populations  individuals.  numbers of A  sampled  of  observed  be  represented  there be  unbalanced  of  sampling  a result, thereby  from  of  but  by  only  different  sites.  adjacent In  is broader  a  population, or  at  the  Such  samples  trees have  decline extent  of  level  was  1976).  found  7.  were been a  large  in  the  i n Appendix  Similarly  study  of  II.  lacked  a  amabilis  the  were w e l l c i r c u m s c r i b e d p h y s i o g r a p h i c a l l y or  the  Specific  in Abies  some s i t u a t i o n s i t was  large  distribution  were e v e n - a g e d and  s u c h as  large  population-intensive  The  i s given  a  e x p e n s e of  geographic  analysis.  more  were s u c h  to ten  at a population  Etheridge  environment  narrow  to a s i n g l e  i s i n d i c a t e d in Table  age-structure  were l a c k i n g on  the  populations  ( H e r r i n g and  physiographic  over  individual  such a  two  trees could  in sampling.  complicating  In most c a s e s stand  of  clearly  five  individual  sampling  into populations  distribution  restricted  were, p o t e n t i a l l y ,  assigned  samples  spent  Ledig  population.  s u c h c o n d i t i o n s where t h e r e  time  -  for studies with  would have r e s u l t e d i n a c o n c o m i t a n t  As  stands  from a  stringent definitions  some a r e a s  individuals,  distribution  complex  such  a  sampled c o u l d  Under  progeny  size"  intra-population variation  i n such p o p u l a t i o n s ,  sampling  samples  of  l a r g e r number of  investment  area.  the  that  trees that could  population.  made.  In  of  ("neighbourhood  a c o n s e q u e n c e of a d o p t i n g  than  definitions  a result  number of  As  of a p o p u l a t i o n ,  operational  the  each other  represent  generation.  definition  As  with  edaphic  and  - individuals  edaphically  p o s s i b l e to  define  46  T a b l e 7. D i s t r i b u t i o n of s a m p l e s as p o p u l a t i o n s and s i n g l e o c c u r r e n c e s . P o p u l a t i o n s - t o t a l number of s a m p l e s : t o t a l H p o p u l a t i o n s {0 p o p u l a t i o n s * H s a m p l e s p e r population).  IDENTIFICATION P.  POPULATIONS  s i tchens i s STANDARDS PUTATIVES  44: 21:  6(16;2*10:2*3:2} 7{6:4;3;4*2>  18 44  82:  1 3{ 16: 1 5: 1 3: 1 0 ; 7 ; 5 ; 2 * 3 ; 5 * 2 >  51  enge1mann i i STANDARDS PUTATIVES  66: 84:  7 { 2 1 ; 1 1 ; 1 0 ; 2 * 7;2 *5} 1 3 { 2 2 : 15; 1 3 : 2 * 6 ; 6 * 3 ; 2 * 2)  5 21  S E L K I R K MTNS. STANDARDS PUTATIVES  10: 66:  2*5 1 9 { 1 0 : 2 * 5 : 4 : 12*3 ; 3*2)  "HYBRIDS"  P.  TOTAL  and  where t h e r e  were o b v i o u s  differences. saplings)  S t a n d age  another.  were g r o w i n g general  the  from  to  coupled  Approximatly lone  40  trees.  hypothesized  by  coast,  spatially  environmental  obvious and  Dobbs  as  edaphic  of  Indeed,  t o be  soil,  age  the this  i n the  asked as  collections  of  of  collected  agreeing  with  sense d e f i n e d l o g g i n g or  restricted  inability  and  the  others.  a c o n s e q u e n c e of  populational level  causes questions  sites  perturbation  s a p l i n g samples  i t s early successional  percent  adjacent  d i f f e r e n c e s or  (1972) and  (i.e. relatively  with  142  d i f f e r e n c e s ( i . e . c o m p a r i n g mature  populations,  find  conditions  Picea  from  i n r e c e n t l y exposed m i n e r a l  were d i f f i c u l t  of  samples  Seedling  observations  Along  edaphic  collect  were a s s o c i a t e d w i t h  form o r  3  3 7 3 : 6 7 { 2 2 ; 2 1 ; 2 * 1 6 ; 2 * 1 5 ; 2 * 13; 11;5*10;3*7;3*6;7*5; 2*4;23*3;15*2)  populations  one  SINGLES  simply  edaphic  amplitude  occurrence).  made were  to s a t i s f y  trans-individual  whether or  above,  not  a  sampled the variation  "population"  is  47  indeed  a viable,  coastal  naturally  Picea c o l l e c t e d  3.2.2 Taxonomic  o c c u r r i n g s u b j e c t worthy o f study i n  in this  study.  c i r c u m s c r i p t i o n of samples.  A pre-requisite for investigating that For  reference  samples, d e s i g n a t e d  the purposes of t h i s  choosen: standards,  Trees that  fluvial  the  three  100m ASL and were g r o w i n g  as s t a n d a r d s  swamps, o r b e a c h  and w i t h i n  C a s c a d e Range and t h e more e a s t e r l y of edaphic  P. engelmann i i .  Owing t o t h e t a x o n o m i c  1974; Roche  representative  were  coast-mainland  in alluvial  in British  1969; T a y l o r  o f P_^ g l a u c a  Individual  500m o f t r e e l i n e i n ranges  Columbia  were o b t a i n e d  were,  as standards of  confusion  1959)  sites,  f r o n t s were  mountain  habitat, considered  P. e n g e l m a n n i i and P^ g l a u c a Daubenmire  and  r e p r e s e n t i n g P^ s i t c h e n s i s .  growing a t the t r e e - l i n e  regardless  are required.  groups of s t a n d a r d s  t h e immediate c o a s t  terraces, Lysichiton  declared trees  below  as "standards",  of taxa i s  p u t a t i v e s , and h y b r i d s .  growing along  occurred  study  the s i m i l a r i t y  between (Garman 1957;  standards f r o m t h e Ottawa  Valley . 1  Trees displayed the taxa  w h i c h were g r o w i n g a t i n t e r m e d i a t e "characteristic"  two t a x a being  crown and m o r p h o l o g i c a l  c h a r a c t e r s of  were d e c l a r e d t o be p u t a t i v e r e p r e s e n t a t i v e s o f t h e  investigated.  Samples c o l l e c t e d Quebec a M o n t r e a l . 1  e l e v a t i o n s and  The d e s c r i p t i o n  courtesy  o f " h y b r i d s " was  o f D a n i e l Gagnon, U n i v e r s i t e du  \  48  applied  to those  morphological individuals  t r e e s t h a t had  characters.  separated,  intermediate  an  Such an  Mountains along  alluvial  along  steep c o l l u v i a l  slopes with  expected  to favour  disposition given of  of  in Table  5.  The  i n Appendix  3.2.3  Local geographic  the  the  variation  areas.  are given  geographic  individual  local  taxa of  and  and  This that  of h y b r i d s .  should  be  individual  The  hybrids i s  and  elevation noted  in  samples i s  The  areas  general  be  divided  transition,  and  interior.  the physiography are  of t h e  variation  location  of  these  distribution  in Table  8.  The  it  to  and was  16  geographic  of  samples  into  distrubition  of  i n Appendix I I .  areas  can  between m o r p h o l o g i c a l  geographic  The  in given  trees i s given  samples.  somewhat a r b i t r a r i l y ,  i n F i g u r e 6.  areas  at  the Cascades  longitudinal,  two  relationship  samples and  geographic  growing  p u t a t i v e s , and  c i r c u m s c r i p t i o n of  with  These g e o g r a p h i c  to  of h y b r i d  terrances,  survival  disposition  to a s s i g n samples,  geographic areas  of t h e  and  II.  In e x a m i n i n g  necessary  and  standards,  standards  given  seepage.  latitudinal,  More s p e c i f i c  anatomical  fans, f l u v i a l  hybridization  5.  s i d e of  form  "hybrid" h a b i t a t c o i n c i d e s with  s a m p l e s as  s e p a r a t i o n of  Figure  the  t r e e s found  the c o a s t a l  Coast  of  identification  as a g r o u p ,  e l e v a t i o n s on  characterization  " i n t e r m e d i a t e " crown  c i r c u m s c r i b e the into  three  These broad study  area  broad areas  occurrence areas:  coast,  roughly  correspond  ( H o l l a n d 1976).  not meant t o r e f l e c t  climatic,  of  The  16  ecological,  49 F i g u r e 6. L o c a t i o n of g e o g r a p h i c areas c i r c u m s c r i b i n g samples. General geographic areas: C - coast; T - t r a n s i t i o n ; I - i n t e r i o r . Specific geographic locations: 1 - s o u t h e r n V a n c o u v e r Is. and O l y m p i c P e n n i n s u l a ( S V I O L Y ) ; 2 - Howe S d . a n d W h i s t l e r (HOWHIS); 3 - l o w e r F r a s e r V a l l e y ( L O F R A V ) ; 4 - T o b a I n l e t (TOBA ) ; 5 - B u t e I n l e t (BUTE); 6 - K n i g h t I n l e t (KNIGHT); 7 - n o r t h e r n V a n c o u v e r Is. (NVANCI); 8 C h i l l i w a c k V a l l e y ( C H I L L I ) ; 9 - F r a s e r C a n y o n ( H O P L Y T ) ; 10 - u p p e r L i l l o o e t t R i v e r ( P E M B R A ) ; 11 - Sumalo a n d S k a g i t R i v e r s (HOPMAN) ; 12 - S i m i l k a m e e n R i v e r (MANPRI ) : 13 - O k a n a g a n (OKAN); 14 - Mt. R e v e l s t o k e (MTREV); 15 - R o g e r s ' P a s s (ROGPAS 1 ; 16 - M i c a C r e e k (MICA ) .  edaphic, samples  or taxonomic  i n a given geographic  circumscribed local  i n s u c h an a r e a .  Selection The  and  simply group  of  that  the ground.  8  to i l l u s t r a t e the  variation  that  might  be  i l l u s t r a t e s the v a r i e t y  i n a narrowly  of  defined area.  samples.  s y s t e m a t i c sampling  pole pruner.  Table  c a n be f o u n d  arborescent habit  branches  together  a r e a and a r e g e n e r a l l y  by a major d r a i n a g e and a t t e m p t  identifications  in  They  m o r p h o l o g i c a l and e n v i r o n m e n t a l  encountered  3.3  groups.  o f P i c e a made f o r s p e c i f i c  of f o l i a g e ,  were o f t e n The l o w e s t  not e a s i l y whorl  branches,  problems  and c o n e s .  Foliage  o b t a i n e d even w i t h a 3m l o n g  branches  were o f t e n  C o n e s i n t h e two s p e c i e s a r e u s u a l l y  over  6m above  restricted  to  50  T a b l e 8. D i s t r i b u t i o n of samples i n t o geographic a r e a s . Numbers a n d a b b r e v i a t i o n s f o r g e o g r a p h i c a r e a s g i v e n i n c a p t i o n t o F i g u r e 6. SSS s i t c h e n s i s s t a n d a r d : SSP - P^ s i t c h e n s i s p u t a t i v e ; SXE " h y b r i d " : ESP - P^ e n g e l m a n n i i p u t a t i v e ; ESS - P^. e n g e l m a n n i i s t a n d a r d ; IESP - S e l k i r k M t n s . P_^ e n g e l m a n n i i p u t a t i v e ; IESS S e l k i r k Mtns. P_^_ e n g e l m a n n i i s t a n d a r d . Note, not a l l t r e e s c o l l e c t e d were a s s i g n e d t o a g e o g r a p h i c a r e a .  GEOGRAPHIC AREA SVIOLY HOWHIS LOFRAV TOBA BUTE KNIGHT NVANCI CHILLI HOPLYT PEMBRA HOPMAN MANPRI OKAN MTREV ROGPAS MICA  SSS  U) 1 2 3 4 5 6 7 8 • 9 10 1 1 12 13 14 15 16  SSP  9 38 5 1  2 6 14 3 18 1 1 3 2  7  IDENTIFICATION ESS IESP ESP  SXE 15 47  2  1 5 30  2  TOTAL  IESS  24 89 1 1 16 8 50 18 24 10 66 18 72 28 29 24 21  21 8 66 4 28 1  14  44 27 5  24 24 16  5  **  the t o p t h i r d Harrison  o f t h e canopy  not a l w a y s a v a i l a b l e damage,  as a r e s u l t  five  with  juvenille 1974).  foliage  Similar  reported  As w e l l ,  that  Cones were  or sporadic  of the f o l i a g e  cone-set,  of such  s u c h young  i sdistinct  and problems  small trees  between age a n d s i z e  f o r t r e e s younger  o f immature t r e e s  that  structures appear  from m a t u r e t r e e s  ( 1 9 7 5 ) , a l t h o u g h he c l a i m s  i s most p r o n o u n c e d  aspect  size  relations  by F u n s c h  had t o  predation.  the manipulation  d u r i n g measurement.  of cones u s u a l l y  t o t h e ground.  of poor  1976b;  y e a r s o l d o r y o u n g e r were n o t sampled  to the diminutive  encountered  age  had f a l l e n  or s q u i r r e l  Those t r e e s owing  (Owens and M o l d e r  and Owens 1983) and c o l l e c t i o n s  be made from c o n e s t h a t  insect  of the t r e e  508 * *  than  was o b s e r v e d  to display (Jeffers  have been that  t h e e f f e c t of  15 y e a r s .  regards  Another  pubescence.  51  It  appears  that  young t r e e s  may  lack  p u b e s c e n c e whereas  mature t r e e s  i n the  same s t a n d  have p u b e s c e n c e .  observations  at  Chilliwack  River  described  pubescent, are  as  (1948) r e p o r t s reasons  twig  the  Wherever p o s s i b l e ,  around  the  b a s e of  contamination It a  was  the  sampling Following the  amount  belonging  manner of  the  trees  collection,  to  of the  canopy. trees  For  these  immature  trees.  materials  Cones were c o l l e c t e d from  trees  collecting  was  minimized.  cones  represented  climbing, from  only  those cones  directly  or  fallen  c o n e s were e x a m i n e d and,  recent  possible  Cone c o l l e c t i o n s made  f r o m a h e l i c o p t e r or  the  are  Lindquist  reproductive  were made by  decomposition, most  for  from n e i g h b o u r i n g  u p p e r c a n o p y of standing  and  individual.  this  from  recorded  that  i n d i v i d u a l i n s u c h a manner t h a t  cones  assumed t h a t  random sample  from  of  by  the  not  on  species  f o r P_;_ a b i e s .  vegetative  were c o l l e c t e d from e a c h  Based  g l a b r o u s when young.  similar observations  m o r p h o l o g y was  Nursery,  the  on  trees.  the  basis  obviously  cone c r o p were r e t a i n e d  for  measurement.  B r a n c h and whorl primary  f o l i a g e c o l l e c t i o n s were made from t h e  branches  ( F i g . 7).  from t h e  m i d d l e of  were not  a l w a y s a v a i l a b l e and  different of  often  old  t o measure c h a r a c t e r s  orders  t o be to  For  g r o w t h was  n e e d l e s were  increment.  other  difficult  branches c o l l e c t e d .  where a n n u a l e x t e n s i o n possible  year  c a n o p y p o s i t i o n s had  b r a n c h i n g was  damage t o  a two  Twig and  of  identify  very  branches The  growing at  with  i t was twig  from  true  owing t o  small,  associated  sampled  Such c o l l e c t i o n s  collected.  trees  lowermost  order  previous  tree  line,  not morphology.  52  F i g u r e 7. S c h e m a t i c r e p r e s e n t a t i o n of s o u r c e s of intra-individua1 v a r i a t i o n a s s o c i a t e d w i t h b r a n c h a r c h i t e c t u r e in enge1mann i i a n d P. s i t c h e n s 1 s . A) F o u r - y e a r o l d s t e m (O) w i t h w h o r l (W) a n d i n t e r w h o r l (I) p r i m a r y (1) b r a n c h e s . B) T h r e e - y e a r o l d w h o r l p r i m a r y b r a n c h w i t h whorl and i n t e r w h o r l s e c o n d a r y (2) b r a n c h e s . Circles r e p r e s e n t whorl nodes. O r d e r i n g scheme i s a c r o p e t a l .  Generally,  vegetative  m a t e r i a l s were more d i f f i c u l t  t h a n were r e p r o d u c t i v e collections smaller  could  patchiness  unbalanced trees could  suites  of the s a m p l i n g  availability  evaluation  complete were  from  individuals created trees  ( i . e . not a l l  of v a r i a b l e s u i t e s measured). of samples c o m p l i c a t e s  of the d a t a .  A general  of i n d i v i d u a l s w i t h  i n Appendix I I .  More  where t h e t r e e s  v a r i a b l e s u i t e s i s given  disposition  i s given  9).  forms..  have t h e same number  of complete  Specific  crown  v a r i a b l e suites for separate  Such an u n b a l a n c e d statistical  (Table  be made i n t h e i n t e r i o r  and had n a r r o w e r  The  number  materials  to obtain  any p u r e  summary  i n Table  respect  of the  9.  to variable  53  T a b l e 9. Summary o f s a m p l e s w i t h c o m p l e t e m e a s u r e m e n t s f o r v a r i o u s v a r i a b l e suites. p - tt v a r i a b l e s p e r v a r i a b l e s u i t e . Note, cone morphology includes cone c o l l e c t i o n v a r i a b l e s .  IDENTIFICATION  P.  VARIABLE SUITES MORPHOLOGY LEAF CONE TWIG (P = 4) (p = 6) (p=15)  sitchensis STANDARDS PUTATIVES  "HYBRIDS"  (p=36)  46 35  44 19  24 16  62 65  102  99  100  77  50  133  45 63  53 60  53 52  65 71  37 38  7 1 105  10 55  10 45  10 45  10 4 1  10 29  10 69  Analyses. The  rather  are  data analyses  statistical  frequently  satisfying  general  indication  statistical associated  inferences,  attendant  Further,  this  part  (1941,  rather  than  confirmatory  that  in biology  1956).  ideal,  focuses  more  f o r p u r p o s e s of precise  approaches are less  with purely  1977).  t o the n e c e s s i t y of  The a n a l y t i c a n d b i o l o g i c a l  assumptions  stringent  statistical  exploratory  data  data-analytic  and s y s t e m a t i c s  of the i n f o r m a l  1949,  and Tukey  an e x p e r i m e n t a l  techniques  of a c o n f i r m a t o r y  approach  exploratory  S u c h an a p p r o a c h  and d e t e r m i n a t i o n  such h e u r i s t i c  merely a r e p h r a s i n g Anderson  t o make owing  I t i s worth n o t i n g  an i n t e g r a l  although  of s t a t i s t i c a l  those a s s o c i a t e d  approaches.  (Mosteller  assumptions.  inferences. with  here are p r i m a r i l y  in nature  difficult  the u t i l i z a t i o n  than  conducted  than c o n f i r m a t o r y  Precise  is  TOTAL AVAI LABI  46 35  S E L K I R K MTNS. STANDARDS PUTATIVES  on  TOTAL  42 59  P . enge1mann i i STANDARDS PUTATIVES  4.  ANATOMY LEAF (p=11)  data-analytic  analysis  protocol.  i s n o t new, methodology of  54  4.1  Mathematical The  been  mathematical  under  of  the  investigation  investigation.  analytic  intuition,  complexity  and  formulation  notation  and  notation  r e s u l t s are  the  text,  4.2  Statistical  the  i n t o the  actual  encumber  the  analysis.  owing t o  the  length  detail  presented  of  for  i n the  However, as  tables  are  to  a variety  regression,  and  of  estimate various  whereas m u l t i v a r i a t e  data.  MVAs p r o d u c e  are  l i n e a r c o m p o s i t e s of the  would not  variable  with  analytic the  in  results,  of  in these  of  the  text  discussion  is  many  tables  in Appendix  univariate  (MVA)  minimize  of in  III.  (analysis  the  Univariate  original variables  typically  variables and  are  r e l a t i o n s among samples where t h e o r d i n a r i l y permit  inter-correlations.  from  such e x p l o r a t i o n  the  used  dimensionality  and  of  multivariate  parameters are  summary s t a t i s t i c s the  and  (PCA)).  population  techniques  p a r a m e t e r s and  the  of  reader  correlation)  (UVA)  variables  an  communication  body of  presented  techniques  further  the  the  ( p r i n c i p a l components a n a l y s i s  explore  for  has  techniques.  (ANOVA),  maximize v a r i o u s  the  various  techniques  data  to a l l o w  b a s e d upon d e t a i l s e x c l u d e d  complete  here  a n a l y t i c model employed  and  does not  given  biological situation  allows  D a t a were s u b j e c t e d variance  the  Such b r e v i t y  minimized.  the  minimized  of  and  intentionally  formulation  t r a n s l a t i o n of  detail  Similarly, the  and  i n t e n t i o n a l l y s i m p l i f i e d and  appreciation  the  notation  to  of  which used  to  original because  55  MVA  methods have been u s e d most commonly a s  tendering  evolutionary  inferences, Barnes 1977;  aside  inferences.  T a i and  Goodman  1967;  M a k i n g more d i r e c t  (Lande  Tarn Neff  1979;  1980; and  T a i and  Smith  Leamy  De  recent  and  Morishima  1968;  S m i t h , e t a l . 1962;  and  Morishima  1969;  Atchley,  1977;  Becker  regarding  of  purposes,  character  1984).  To  example,  variation  ignore  the  they  generally and  potential effect  in accounting  of of  genetic  for observed  Oka  the  genetic  specific  multivariate 1981;  variation  effects,  patterns  however  for  intra-chromosomal  multi-genic  1968;  Although  appropriate  ignore  rare  Hashiguichi  (see  c o v a r i a t i o n (Arnold  such a s p e c t s  inter-chromosomal e f f e c t s , effects  often  Oka  1954;  UVA  "selection indices"). are  Williamson  C h e v r u d , e_t a l . 1983).  i n f e r e n c e s have been b a s e d on  from UVA  and  comparatively  Wright  e t a l . 1982;  for  genetic  1980;  M o r i s h i m a and  Most g e n e t i c  breeding  Jong  1979), a r e  and  inference derived  basis  from c o n t r o l l e d h y b r i d i z a t i o n ( S h a r i k  1971;  1967,  the  of  nature  Sterns ignores,  for  linkage,  and  cytoplasmic  phenotypic  variation.  It  i s the  t o s t u d i e s of be  v i e w e d as  different al.  inter-relation biological  time  scales.  Nothofagus a n t a r c t i c a .  of A b i e s . these  systems.  Development  Maze, e t a_l. ( 1984) such changes Mitton,  and  Most  evolution  with  and  for ovule  i s fundamental  respect  Scagel,  can to  et  development  of  e t a l . (1980) d e m o n s t r a t e  c h a n g e s between p o p u l a t i o n s  (1983) d e m o n s t r a t e s  on  v a r i a b l e s that  changing v a r i a b l e i n t e r - r e l a t i o n s  (1984) d e m o n s t r a t e  similar  of  of  Pinus  ponderosa.  s u c h c h a n g e s between p o p u l a t i o n  structured m u l t i v a r i a t e systematic  v a r i a b l e i n t e r - c o r r e l a t i o n s (Adams  1982;  and  Maze species  studies Campbell  rely and  56  Dearn  1980; MVA  for  S c a g e l and  Maze  methods a r e u s e d  simultaneous  1984).  throughout  quantification  time  for q u a n t i f i c a t i o n  and  n e g l e c t the  Quantification  and  inter-dependance systems.  evolutionary  4.2.1  statistical  distributed.  UVA  o f UVA and  test  falling  for  D'Agostino's  to  Lewis  and  MVA  often  assumption  nature  of  the  specified  D were f u r t h e r  Research  (g^).  and  satisfaction  assumptions  about  t h e d a t a be  plots  Lab.  1974)  was  used.  confidence l i m i t s characterized  (CFDs, U n i v .  1976),  of  measures of  Such c h a r a c t e r i z a t i o n  allows  and  aids  i n the d e t e c t i o n  (Tukey  1977),  data.  normally  Variables calculated  using  cumulative  Michigan, skewness  ( g t ) and  i n f e r e n c e s to (Bock  of o u t l i e r s  Transforming data to approximate  the data  the  of  distributions  t r a n s f o r m a t i o n s a p p r o p r i a t e to minimize  symmetrize  biological  developmental  require  i s that  of n o r m a l i t y ( Z a r  within  1978).  at a  assumptions.  distributional  distribution  normality,  one  variable  drawn c o n c e r n i n g t h e n a t u r e o f t h e d i s t r i b u t i o n suggests  methods  variables.  In a s s e s s i n g n o r m a l i t y of v a r i a b l e  not  kurtosis  linear  variables  o f whole o r g a n i s m s ,  A common d i s t r i b u t i o n a l  Statistical  integrated  to d i s t r i b u t i o n a l  sampling  frequency  attention  provide  interdependance.  Utilization  D'Agostino's  of  they  and  contrast,  of o n l y s i n g l e  r e s p e c t s the  Conformity  specific  By  inter-dependance  In t h e c a s e  s t u d y as  of v a r i a t i o n  i n t e r - i n d e p e n d a n c e of v a r i a b l e s . provide  the  be  1975),  departures  from  ( B a r n e t t and  n o r m a l i t y or  although useful  for  drawing  57  statistical For  c o n c l u s i o n s , may make i n t e r p r e t a t i o n  the v a r i o u s data  h a v e t o be t r a n s f o r m e d . skewed t o t h e r i g h t data.  This  jj_979). to  s e t s examined t h e v a r i a b l e s d i d n o t Most v a r i a b l e s a p p e a r e d  from t h a t e x p e c t e d  i s t h e same s i t u a t i o n  applications  o f MVA.  i s most e a s i l y distributions 1975;  normality  Pimentel  i s assumed  The a s s e s s m e n t  approached  Generally,  distributions,  approximating  1979).  sufficient  (Mardia  1968; Day 1969; M a r d i a  Machado  1978;  Andrews, e_t §JL. 1972) ,however  conformity  technique  1979; Reyment  small  s e t s of data.  proposed  by C a m p b e l l  (1980).  Purely are available and Zemrock  elegance i s  c o s t and r e s t r i c t i o n t o  An h e u r i s t i c  to multivariate normality  assessment of  i s provided Gnanadesikan  by t h e CFD (1977)  p r o v i d e s many e x a m p l e s o f t h e u s e o f CFD t e c h n i q u e s f o r characterizing  m u l t i v a r i a t e data  prior  i s not  1971; Cox and S m a l l  statistical  a t t h e expense of c o m p u t a t i o n a l  dimensionally  distributions  by i t s e l f  assessments of m u l t i v a i a t e n o r m a l i t y  1983; Malmgren  (Bock  multivariate normality normality  1975;  the  distributed  multivariate normality.  1970, 1974; Wagle  achieved  of m u l t i v a r i a t e n o r m a l i t y  t h e c o n d i t i o n a l and component  However, m a r g i n a l  to indicate  statistical  for statistical  i f the marginal  so w i l l  (Pimentel  v a r i a b i l i t y at  by a s s u r i n g t h a t t h e m a r g i n a l  approximate normality, thereby  either  error.  of the v a r i a b l e s are normally 1979).  distributed  may be a t t r i b u t e d  o r t o i n c r e a s e d measurement  Multivariate  slightly  r e p o r t e d by B a r k w o r t h , e_t a l .  Such skewed d i s t r i b u t i o n s  sizes  t o be  f o r normally  the consequence of i n c r e a s e d developmental  larger  difficult.  t o MVA.  58  For  most h e u r i s t i c  normality contain  does not  The  or g r o u p s of  group c o v a r i a n c e  data  multivariate  normally  were a p p a r a n t l y departure  be  matrices  s e t s examined h e r e  from t h e  curve  and  sensitive  (Gower  not  such  as  to  especially  with  repect  1972). as  being  S i n g l e or g r o u p s of t h e r e was  expected  data  Some MVA,  were a l l i n d i c a t e d  distributed.  non-existent,  t h a t the  outliers.  from m u l t i v a r i a t e n o r m a l i t y ,  individual  multivariate  assumed, m e r e l y  v a r i a t e s a n a l y s i s (CVA), may  departures to  have t o be  wild outliers  canonical  a p p l i c a t i o n s of MVA,  little  outliers  i n the  way  of  f o r a m u l t i v a r i a t e normal  distribution.  4.2.2  Sample  size.  Fundamental of  an  adequate  concerns  refer  t o any  study  of v a r i a t i o n  sample  size  (n)  t o the  sample a c c u r a c y .  for a s p e c i f i c  in' P i c e a have g e n e r a l l y u s e d an more t o t h e  e c o n o m i c s of  variability  of  of  these  the  1 for c i t a t i o n  Concern constitute  f o r the  a valid  However, w i t h o u t in  to the  No  of  criteria  information of  specific  the  placed  i s provided sizes  value  of  the  the in  any  used  r e s e a r c h may  f o r d e c i d i n g upon a g i v e n  some i n f o r m a t i o n c o n c e r n i n g  i n the  relating  studies).  e s t i m a t i n g a d e s i r e d p o p u l a t i o n parameter  confidence  Such  respecting  sample  e c o n o m i c s of c o n d u c t i n g  criterion  purpose.  r a t h e r than  adequacy  establishment  Previous i n v e s t i g a t i o n s  n b a s e d on  study  organism.  s t u d i e s as  (see T a b l e  the  i s the  sample  the  well  n.  adequacy of  t h e r e can statistic.  be  n  little Sample  59  size  directly  influences  t h e more r e l i a b l e Any  estimate  the  i s the  sample  statistic  statistic that  of n i s , by d e f i n i t i o n ,  required  to estimate  reliably  sample  e s t i m a t i o n technique  upon  the methodology o u t l i n e d  approach  seeks  to s e l e c t  correlation  matrix  n  A similar  f o r MVA.  for  individual  standard  error  measuring  was  an  made.  initial  size  The  al.  more  be  parameter.  employed h e r e  i s based  thereby  This of  providing a  e s t i m a t i o n methodology  i s less  sampling  technique  e s t i m a t e of  selects than  an  n a t which  the a c c u r a c y  of  technique.  were u s e d  were g e n e r a l l y t o o  as  the stable i s used the  the  that  small  (1972) i n  by Wood  these  of v a r i a t i o n  (Table  10).  sizes  suggested  of m u l t i v a r i a t e  by  sample the  less  of S c a g e l , e t  sample  size  In a r r i v i n g  sample-size  be  sequential,  However, u t i l i z i n g  initial  (1972)  (1972) r e q u i r e s  is a  Sample  univariate  the b a s i s f o r s e l e c t i n g  suites.  e x t e n s i o n of t h e  underestimation  by G r e e n  s a m p l e - s p e c i f i c methodology  (1984) i n d i c a t e d  multivariate  illustrated  (1972) t e c h n i q u e  a  by G r e e n  the p o p u l a t i o n c o e f f i c i e n t  for separate v a r i a b l e and  study  illustrated  illustrated  graphical  techniques  for this  technique  In c o n t r a s t , Wood's  parameter-  size  estimation technique  sample-specific these  n will  i n S c a g e l , e_t a_l. ( 1 9 8 4 ) .  where one  t h e mean  an  n at which the d e t e r m i n a n t  sample  with a g r a p h i c a l  used.  that  of  a given  stabilizes,  variables  the  size  concert  (|R|)  larger  The  device.  During sample  an  the  n is  parameter.  sample s p e c i f i c .  a collection  size  larger  e s t i m a t e s the  variable  The  of s a m p l e s ,  - the  at  estimates the  estimation a  sample s i z e  was  similar  observed  f o r the  60  T a b l e  1 0 .  S a m p l e  E s t i m a t e s d e c l i n e s s t a b i l i z e ( 1 9 7 2 ) o f  s i z e  b a s e d  o n  b e l o w  e s t i m a t e s s t a n d a r d s  m e a s u r e m e n t  i n t e r - i n d i v i d u a l  a n d W o o d  v a r i a b l e s  ( 1 9 7 2 )  n  N E E D L E N  ( n / t  p e r v a r i a b l e  ' eftt '  V A R I A B L E  ( 8  »  4 B X S T 0 M  ( 9  1  4 D X S T 0 M  ( 7 )  R E S C Y N O  ( 3 5 )  f  o  o f  r m t r a - i n d i v i d u a 1  b o t h  t a x a .  a c c u r a c y . v a r i a t i o n U S E D  v a r i a t i o n  U n i v a r i a t e  M u l t i v a r i a t e  t  b a s e d -  n  o n b a s e d  a v e r a g i n g o n  l a r g e r  n  b a s e d o f  i n t e r - i n d i v i d u a l  b a s e d  b a s e d  u p o n  u p o n o n  G r e e n  n  t r i e  o r  t  s a m p l e - s i z e o r  W o o d ' s  n  v a r i a t i o n  o r  t  a t  w h i c h  |R(  o r  e s t i m a t i o n  o f  a t  w h i c h  t e c h n i q u e .  p  -  s i t c h e n s  i s  n / t U S E D P .  e n g e 1 m a n n 5  i i 5  s ' t c h e n s  i s  P  2 0  15  5  5  2 5  2 0  10  5  10  10  1  10 1  1  1  1 1 9 )  10  2 0  10  2 0  ( 4 7 )  10  2 0  10  15  P U L V L E N  1 8 )  2 0  3 0  4 0  4 0  T I P W I O  ( 2 )  7  15  2 0  T I P O E P  ( 2 )  5  10  15  P U L V P U B  ( 1 )  1  N E E D W I D  ( 2 )  2 5  3 0  N E E D E P  ( 1 I  2 0  4 B X 4 N G  5  1  10 2 0  1 5  1  5 0  4 5 4 0  3 0  4 5  < 1 )  5  10  10  4 D X 4 N G  ( 3 )  8  2 0  2 0  15  C E N C Y W I D  ( 1 )  2 0  3 0  4 5  4 5  C E N C Y L 4 T  ( 2 1  2 5  3 5  7 0  5 0  C E N C V 4 B X  ( 2  )  2 0  3 5  3 0  4 0  C E N C Y 4 D X  1 8 )  2 5  3 5  5 0  4 5  E N D O N U M  ( 2 1  END  C O N L  E N  5  5  10  2 0  2 5  ( 4 )  2 0  4 0  5 0  4 5  ( 1 1 )  2 5  3 0  4 5  3 0  5  2 5  2 5  5  . 1 0  ( 3 )  3 0  C Q N W I D  ( 2 )  2 0  S C 4 L E N  ( 7 1  2 0 1 1  5  5  10  "  10 10  S C 4 L W I D  ( 6 )  2 0  2 5  2 0  3 0  S C 4 L T 4 P  I 1 2 )  2 5  3 0  2 5  3 5  W I N G W I D  ( 8 )  3 0  3 0  2 0  3 0  W I N G T A P  ( 1 0 )  2 0  3 0  2 0  2 0  F R E E S C 4 L  ( 2 8 )  15  3 5  2 5  3 5  B R A C T L E N  ( 2 0 )  2 5  2 5  3 0  3 5  B R A C T W 1 D  ( 1 2 )  10  5  10  15  B R 4 C T T 4 P  ( 3 0 )  2 5  10  2 5  2 0  -  S H C O L E N  ( 3 )  S H C O W I D  ( 2 )  L O C O L E N  ( 3 )  L O C O W I D  I 2 )  V A R I 4 B L E L E A F  4 N 4 T 0 M Y :  3 0  3 5  D'  2 5  3 0  C O L L E C T I O N :  M O R P H O L O G Y : < p = 6 1  T W I G  M O R P H O L O G Y : ( p = 4  C O N E  2 5  -  | R |  ( p » 1 1 ) L E A F  >5  2 5  15  10  2 5  2 0  5  10  S U I T E  ( p = 1 1 > C O N E  -  )  M O R P H O L O G Y : ( p = 4  )  1*1 D'  l«l  -  _  -  -  3 0  2 5  D'  2 0  10  |R| 0'  10  2 5  2 0  15  3 5  3 0  15  2 5  lei D  1  5  1  5  4 0  3 0  3 0  2 0  3 0  3 5  2 5  2 0  3 0  4 0  9  15  5  2 5 12  10  10  3 5  3 5  2 0  2 0  2 0  .  n u m b e r  e n g e 1 m a n n ;  R E S C Y L E N  X V L E N D  e  G r e e n  t P .  R E S C Y L O C  P H L  ( t ) . s D •  s u i t e .  n —  ( n ) a n d  e s t i m a t e s  e s t i m a t e s  i  61  technique proposed In  addition  correlation also  application variance  D  The  2  performing  of  analysis  here  f o r both  in a l l tables  were s m a l l e r  t e s t s were p e r f o r m e d  4.2.3  upon  (n o r t ) and d e g r e e s  employed  Discrete  Assessment  suites  i s an MVA.  analysis  of  and of  the to  s i z e estimates are given  in Table  and  10.  The  sample  inter-individual levels  freedom  <df)  figures.  The  those  after  sample  s t a n d a r d s of each  and  indicated  s a m p l i n g was  species.  Sample  have been g i v e n actual  sample  by T a b l e  sizes  10 as  the  completed.  variables.  rather  unlike  the o t h e r v a r i a b l e s  than q u a n t i t a t i v e  inclusion  of  variables  in a multivariate  inter-sample  was  2  statistical  is a pre-requisite  intra-  of  than  P u l v i n u s pubescence, qualitative  the  MVA.  variable  v a r i a t i o n are based  explicitly  2  and m u l t i v a r i a t e and  i n the  e x t e n s i o n t o CVA  (DFA). D  D  a p p l i c a t i o n of  s u c h as m u l t i v a r i a t e  i t s further  a structured  a l l variables  sizes  of M a h a l a n o b i s '  of  of t h e c o r r e l a t i o n m a t r i x  MVA  of M a h a l a n o b i s '  reported  of the d e t e r m i n a n t  i s an e s s e n t i a l s t a t i s t i c  function  Univariate  (1981).  f o r the m u l t i v a r i a t e  (MANOVA) and  stabilization  sizes  determinant  of a s t r u c t u r e d  discriminant  for  the s t a b i l i z a t i o n  statistic  Mahalanobis'  Jancey  t o the s t a b i l i z a t i o n  matric,  examined.  essential  by Newhan and  variable  is a  (Appendix  such a v a r i a b l e a l o n g w i t h q u a n t i t a t i v e  differences  when s t a n d a r d i z e d  analysis  than  (Barkworth,  may  contribute  the q u a n t i t a t i v e e t a_l. 1979;  I).  The  continuous more t o  variables,  S n e a t h and  even  Sokal  62  1973).  Realizing  that  variable  may b i a s  critical  t o determine  t h e i n c l u s i o n o f such a q u a l i t a t i v e  the r e s u l t s of a m u l t i v a r i a t e the degree  with other  analysis.  I f t h e v a r i a t i o n of s u c h a v a r i a b l e  the  of the other  variables  two t y p e s o f v a r i a b l e s A measure o f p a r t i a l  of  the c o n t r i b u t i o n  situation; of  however,  prior  such a v a r i a b l e  assoicated  that  variables  that  then  t o being  i n t h e same  i t  is  may be  submitted  t o an  i s similar to  i t i s permissable  to include  analysis.  c o r r e l a t i o n could  of the c a t e g o r i c a l  address  variable  i t does not a d d r e s s  t h e s i t u a t i o n n o r does  analysis,  in a bivariate  the m u l t i v a r i a t e  i t address the question  o f whether t h e  variable-state  variables.  R e p h r a s i n g , a b i v a r i a t e a p p r o a c h does n o t a l l o w one i f , assuming a q u a n t i t a t i v e  taxonomically  with  nature  qualitative  to determine  i sassociated  the problem  the q u a n t i t a t i v e  variable i s  s i g n i f i c a n t , the q u a l i t a t i v e v a r i a b l e i s  similarily significant. B a r k w o r t h , e t a_l. (1979) a d d r e s s e d MANOVA.  Briefly,  were d e t e r m i n e d quantitative test  by t h e s t a t e  characters  the multivariate  enabled  taxonomically quantitative  used  submitted  t o MANOVA  differences  and o n l y  Employing Wilk's A  between t h e g r o u p  to  centroids  whether a s p e c i f i c q u a l i t a t i v e  t o c a l i b r a t e t h e g r o u p s was i n d e e d  s i g n i f i c a n t ( i . e . shared v a r i a t i o n variables).  contribution  relationships  they  of a q u a l i t a t i v e c h a r a c t e r  were e m p l o y e d .  them t o d e t e r m i n e  variable-state  the  t h e g r o u p s o f samples  t h i s p r o b l e m by way o f  Such an a p p r o a c h t o t h e e v a l u a t i o n o f  of the q u a l i t a t i v e v a r i a b l e  amongst  with  t h e two s p e c i e s  to the  was t a k e n h e r e .  I t should  63  be  pointed  here  2  as o n l y  (pubescence: present  employeed T  out that  test  A  two v a r i a b l e  or a b s e n t ) ,  by B a r k w o r t h , e t aJL.  of the d i f f e r e n c e  summary  states  were  t h e MANOVA  involved  approach  (1979) s i m p l i f i e s t o H o t e l l i n g ' s  between two c e n t r o i d s  of the T s performed 2  i s given  (Morrison  in Tables  1976).  11.  In  T a b l e 11. Summary o f H o t e l l i n g ' s T ' f o r t w i g m o r p h o l o g y b a s e d on v a r i a b l e s t a t e f o r d i s c r e t e (presence/ absence) v a r i a b l e : p u l v i n u s pubescence. *, T s i g n i f i c a n t 9 p < 0.01; D v a l u e s g i v e n i n body of table. P . s - P_^ s i t c h e n s i s ; P . g g l a u c a ; P . e - P_^ e n g e l m a n n i i . !  7  COMPARISON  df 13,x)  INTRA-INDIVIDUAL ADVENTITIOUS v s . WHORL PRIMARY v s . SECONDARY 1979 V S . 1981 INTERWHORL P^ s i t c h e n s i s INTERWHORL P^ e n g e l m a n n i i  only  the  1 . 97 0 . 37 1 1 . 50  INTER-SPECIFIC P.s x P.e x P.s x P.e  984 919  1 . 14* 1 . 59*  a s showing  compared.  was e i t h e r  source  P g  were t h e c o n t i n u o u s v a r i a b l e s  indicated  significant, that  530 385 61  groups being  variable  i nvar i ate i nvar i ate i nvar i ate 2 . 97 i nvar i ate  INTER-INDIVIDUAL P. s i t c h e n s i s P. enge1mann i i P. g l a u c a  two i n s t a n c e s  morphology  8 98 9 52 58  significant  differences  For those comparisons  i n v a r i a t e or the a t t a i n e d  the v a r i a b l e of v a r i a t i o n .  was e x c l u d e d  from  of twig  T  2  between  where t h e was n o t  further  analysis for  64  4.2.4 U n i v a r i a t e  analyses.  ANOVA was u s e d  b o t h a s a means o f t e s t i n g  between means a s w e l l ANOVA was u s e d basis  as examining  of the t o t a l  various hypothesized sources ANOVA with and  f o r both h e u r i s t i c  Wilk  test  the homogeneity  of group  i s consistant  (Tukey  1962; Tukey  1978; O'Grady  in testing  a s "nude" m o d e l s  less  on t h e b a s i s  dependent  variances.  criterion  of t h i s  methods o f p a r t i t i o n i n g gave a p p r o x i m a t e l y  variables  of  rather  o f sums o f  Additionally, t h a n a mean of  equal  i t was o b s e r v e d  v a r i a n c e from  analysis"  w i t h t h e MVA.  the purpose  of  partitioning square individual  that  an h y p o t h e s i z e d  At the  t h e two source  results.  most o f t h e MVAs p e r f o r m e d , involved  1965).  upon t h e a s s u m p t i o n s  the v a r i a t i o n  study  used  (Nooney  f o r a s p e c i f i c h y p o t h e s i z e d source of v a r i a t i o n .  For  1982).  the e q u a l i t y  w i t h t h e " v a r i a n c e component  a l l o w s one t o compare  of c o m p l e t i o n  along  into  The use o f  purposes  o f ANOVA  of v a r i a t i o n  1969),  by a sums o f s q u a r e  time  (uva)).  on t h e  forms o f t h e ANOVA m o d e l s  i n the text  i s , i n comparison  ( S o k a l and R o h l f  groups  the v a r i a t i o n  1978; W i l k i n s o n  The s p e c i f i c  partitioning  criterion  of the d a t a .  v a r i a n c e s when t h e ANOVA model e m p l o y e d was b a l a n c e d  given e x p l i c i t l y  squares  ,e.  ( Z a r 1974) was u s e d  near-balanced.  The  for  50wRC  differences  sums o f s q u a r e s  and s t a t i s t i c a l  1966; Kempthorne  the g r o u p  are  (%SS  the d a t a - e x p l o r a t i v e purpose  Bartlett's  or  the s t r u c t u r e  a s a means o f p a r t i t i o n i n g  of the p e r c e n t a g e  the  were a l s o Although  UVAs o f t h e i n d i v i d u a l  performed  and t h e r e s u l t s  t h e s e UVAs a r e o f t e n  of the p o i n t s r a i s e d  here,  presented  unneccessary  they a r e a v a l u a b l e  65  adjunct  f o r two  may  be  not  allow  an  to  familiar  First,  with  for readers  independent  variables  ( i . e . Newman and  was  variables  of  between the  UVA  may  such  UVA  will  described. the  be  results  given  useful in  considering, s p e c i f i c 1983).  the  MVA  b a s i s f o r most of  sample v a r i a n c e  (as  technique  i n the data  samples without  and  a priori  in canonical c o r r e l a t i o n  upon a c o r r e l a t i o n  correlation  matrix  was  matrix,  r a t h e r than  absolute  examining character v a r i a t i o n  nature  N o y - M e i r , e_t a l . 1975 of  the  e l e m e n t s of R, meaningful  i n the  context  detail  and  comparison  some p  as  PCA  (as  was  variables. the d i s p e r s i o n  interest As  of  samples  was  in  many v a r i a b l e s were means of  c o v a r i a t i o n (see a l s o Noy-Meir  f o r s u c c i n c t d i s c u s s i o n on t o be  submitted  of m o r p h o l o g i c a l  an  intuitively  variation  as v a r i a n c e and  d i s c u s s i o n of PCA  to  the  Individual  c o e f f i c e n t s , rjk , a r e  m e a s u r e s of d i s p e r s i o n , s u c h Considerable  of  extraction technique).  correlation  a  maximizes  stratification  a scale-independent  d i s p e r s i o n matrix  eigenvalue-eigenvector  allows  In a l l c a s e s  variation.  and  summary  PCA  p r e f e r r e d r a t h e r than  commensurable, R p r o v i d e s  1973;  The  a n a l y s i s ) or  R,  or v a r i a n c e - c o v a r i a n c e m a t r i x ,  relative  used. t h e UVA.  c a n o n i c a l v a r i a t e s a n a l y s i s (CCA)).  matrix,  not  and  Jancey  principal  individual  performed The  the  provide  variables in  being  in extending  such  technique  used,  reader  Multivariate analyses. PCA  the  situation  research,  a sampling  s i t u a t i o n s where a  techniques  interested  designing  4.2.5  under  t h e MVA  a p p r e c i a t i o n f o r the  Secondly, here  reasons.  than  more  other  covariance.  i s provided  here  as i t  66  is  t h e most g e n e r a l i z e d of  techniques extended  u s e d and  to the  PCA,  like  techniques  other  other  results  vector-variables and  are  the  the  interpretative  MVA  MVA  are  and  linear  values  solution  the  original  variables,  respect et  referred  X-,  to the  i s the  t o sample  forming  size;  t o as  1969)  Karr  R  and  to each  other.  components  o r , more i s an  v a r i a n c e of  n,  original variables  respect  colloquially,  eigenvalue samples  relating  vector-variable.  samples,  p,  These the  of c o e f f i c i e n t s  new  number of  of  e a c h component  y_j i s a v e c t o r  variable  with  Dempster  A s s o c i a t e d with  component.  composites  or o r t h o g o n a l  (v-).  readily  eigenvector-eigenvalue extraction  variables",  eigenvector  are  used.  These v e c t o r - v a r i a b l e s a r e  "axes".  techniques  i n i_ v e c t o r - v a r i a b l e s .  independent  (="principal  eigenvector-eigenvalue extraction  For  must e x c e e d  a  (X, )  for that the  original  "stable"  the  number  ( s e e a l s o r e m a r k s above Martin  1981;  PCA  of with  Scagel,  a l . 1984).  PCA  maximizes the v a r i a n c e  variables  have been m e a s u r e d .  component  is referred  The that 0.0.  linear the  i n d e p e n d e n c e of  correlation  Cooley  calculation  The initial  t o as  and  of  Lohnes  data.  As  t o e a c h component,  the  the  scores e a c h of  component individual  scores  scores  serve the  n samples on  value  from  (1971) p r o v i d e  of component  component  The  the  the  of  sample  score  for that  components  separate  p  f o r each sample.  i s shown i n  components i s  a readable  a summarization  original  t h e component  of e a c h  for a particular  as  which the  account  of  the  PCA.  of  the  p variables contributes  s c o r e s have t h e d e s i r a b l e  67  property  of  providing  original  data.  more n o r m a l l y same sample 1980;  the  For  Statistical  single  of  PCA  significance. variation be  satisfied  (Ruymgaart  component  discussion  of  Prior  test  of  equality,  the  to  to  was  necessary.  of  and  It  even the  PCA  assumes  that  for  the  performing  do  UVA  and  the  Collins  from a  (1980)  statistical  in  robustness  under  normality  and  non-normal  i s seen  not  appreciation  contain  radical  application  MVA  of  to  cannot  PCA  analyses  of  frequently  Pimentel  (1975) f o r a more with  the  outliers  of  statistical  i s most  as  their  d i s t r i b u t i o n a l caveats  or  method  Collins  heuristic  heuristic  Crow  (1979),  Gittins  detailed  the  method  correlation  matrix  and  situations.  PCA,  to  the  d e t e r m i n e whether the  equicorrelation Chatfield  on  multivariate  techniques  caveats associated  E s t i m a t e s of  also  and  is derived  and  the  is referred  biological  R  marginal  for  data  and  reader  are  i f the  i s the  evaluated  scores are  Chatfield  multivariate  used  I s e b r a n d s and  application  by  data  clustering,  The  on  evidence  be  scores) that  ( 1 9 6 9 ) , and  factored  can  1981).  encountered.  based  interpretative  provided  (ordination,  1979;  However, C h a t f i e l d  PCA  the  original p variables  Pimentel  where r e s u l t s  i n the  component  of  theorem").  Satisfaction  important  the  o r i g i n a l data  associated  situations. being  1976;  (1976) p r o v i d e  and  than  inference  the  in dimensionality  same r e a s o n  limit  population.  Morrison  the  (Morrison  of  reduction  distributed  "central  normality  a  overall  structure  ( B a r t l e t t ' s or  Collins  1980)  in  and  fact of  to PCA  R are  Lawley's t e s t independence  be was provided of  68  (Anderson's rjk  test  of R.  I f the  component  no  can  r ^  to perform  considered  dimensionality  statistically  i s not  based  equal  component  otherwise  eigenvalue  meaningful  the  noted  1976;  of PCA  Cooley  has  already  noted, a l l r j k were  been made w i t h  interpretations the  total  the d a t a  and  from  of  attributed  to  the  (Anderson's  Lohnes  1971).  PCAs r e p o r t e d variation  and  been p r e s e n t e d  the p e r c e n t  t h a t t h e PCA  a correlation  i n the  variance  here have  i s d e r i v e d from  matrix:  t a b l e s of  i s just a  R  variation  i n d e p e n d e n c e of  residual  there  an  1.0.  have not  PCAs as  of  are  zero.  structure in R  a l l components from  variables  R where t h e  to  of t h e  residual  different  provided  For  a test  Morrison  g r e a t e r than  Eigenvalues of  from  1979)  unique  to zero)  otherwise  upon an  results  i s no  of  r e d u c t i o n of  Component-specific  ( % v a r . ) , and  significantly  matrix.  the  and  Unless  same nor  redundancy, G i t t i n s  of e q u a l i t y ,  results  the o r i g i n a l  1980)  variables  equal  upon c o n s i d e r i n g t h e p o r p o r t i o n of  extracted  are  statistically as  Collins  original  independent  of  and  non-zero t h e r e  I f the  have been b a s e d  n e i t h e r the  the component  Unless  but  necessary.  techniques.  (=intra-set  test  a PCA  interpretation  a number of are  Chatfield  equal  linearly  PCAs r e p o r t e d h e r e  • The  are  ( i . e . a l l r j ^ are  reason  be  sphericity,  p o s s i b l e to e x t r a c t .  independent is  of  the  as correlation  69  The  eigenvector  original the  variables  contribution  values,  provide  of  component.  1979), way  to  An  interpret  , (=component  estimate  the  alternate,  the  and  calculate  product-moment c o r r e l a t i o n s  variables  and  Pimentel  1979;  s q u a r e of equals  the  the  correlations  of  by  eigenvector  differences  eigenvector  v a l u e s are  of  signs,  differences.  The  then  a  of  the  is  to  original  correlations",  and  Lohnes  the  component  component  variance  (Gittins  original  correlation  such  average  component  extracted  where t h e  1971).  i s the  for a given  by  that  is  o r i g i n a l data product-moment  eigenvector  derived concerns  of  that  from the  (Pimental  approximately  for  to  values  in  that  tested.  a vector  variation  opposite  total  component  v a l u e s are  form of  primarily,  be  values  a given  approximating  that  Additionally,  inference  eigenvector  described  of  advantage over  can  additional  relationship  under c o n d i t i o n s  have an  the  Cooley  upon component  transformed.  significance  An  (i.e.  reliance  of  ("component  correlations the  of  between  structure", the  significance  s e p a r a t e components  scores  eigenvalue  necessary  have been  signs  the  component  The  especially  their  matrix  p e r c e n t a g e of  component.  the  to  the  component  "component  a correlation  correlations  may  s c o r e s of  the  the  more s t a b l e  contribution  to  the  of  l o a d i n g s ) of  original variables  variables  For  the  an  e a c h of  particular  a;|  the  isometry, component  the  relative sizes  form of  variation  1979).  If  is said  i n s i z e among s a m p l e s . markedly d i f f e r e n t the  component  d i s t i n c t i o n between  the  same s i z e and Pimentel  If  sign  1979),  to  the  r e f l e c t s s i z e and two  then  reflect,  i n m a g n i t u d e and  these  and  forms  of  are  shape  70  variation  i s important  the  differences  and  inter-correlation.  between  Correlation (Atchley, Blackith  et  where t h e  i t serves to  s i m p l e and  Reyment  Pimentel 1971)  the  c o r r e l a t i o n of  comparisons  of  |R|  may  comparing R  (Newman and  component  illustrate  otherwise  scores  the  length  of  the  submitted  the  to  by  valid  only  necessarily  the  or  a l t e r n a t i v e means of  Scagel  amount of  used  of  analysis  and  Maze  of  the  the  to provide  1984;  the  to  sequentially  addition  emphasize smaller  drawn  axes a r e  maximum d i s p e r s i o n  for  the  first  the  the  ordination  s p a c e of  f o r by  In  the  several emphasizes  original  a l l ordinations  a given  decreasing  component variance  is  component  scores  are  samples the given  associated  eigenvectors.  to o r d i n a t i o n ,  This  variance  Where means of of  of  component.  decreasing  in  equal.  the  scale  the  here are  co-ordinate  means were c a l c u l a t e d .  parenthetically  Unless  to  means i n t h e  v a r i a t i o n accounted  ordinations  samples.  presented  components.  illustrated,  from w h i c h t h e  In  an  v a r i a t i o n of  emphasizes  succeeding  dispersion  be  of  scales  the  extracted  with  1981;  axes c o r r e s p o n d  convention  the  as  a l l ordinations  graphical  samples are  can  pattern  indicated  s u c h a manner t h a t  sample  not  the  e t a l . 1984) .  The  The  of  c o r r e l a t i o n matrices  suffice  Jancey  but  PCAs  vectors";  Such c o m p a r i s o n s a r e  The  that  variation  separate  assessment  same v a r i a b l e s  also  respectively,  "theoretical  p r o v i d e s an  same s a m p l e s .  Scagel,  v a l u e s of  1979,  between t h e s e PCAs. PCAs s h a r e  reflect,  complex v a r i a b l e  among e i g e n v e c t o r  a l . 1982;  and  similarity  as  also  71  utilized  i n subsequent  techniques a  further  allow  not  component  scores  1979).  "%SS  S o u R c £  ( b u t s e e Moore a n d Maze  Total  multivariate variation  (mva)".  Owing t o t h e a d d i t i v e  from  assessment  of the t o t a l  components  ( S c a g e l and Maze  given,  interpretation  tests  PCA, t h e u t i l i z a t i o n  no s i m i l a r  tests  and r e g r e s s i o n s c o r e s has  1965; P i m e n t e l  1984; W h e e l e r and i s referred nature  t o here  of the  o f ANOVA may p r o v i d e an  multivariate variation 1984; W h e e l e r  of s i g n i f i c a n c e  and p r o v i d e  o f a component.  ANOVA o f component  1983; S c a g e l  components  Although  Univariate  in correlation  i s a common p r a c t i s e .  Maze a n d P a r k e r  Guires as  a component-specific  r e c e i v e d t h e same a t t e n t i o n  1979;  analyses.  method o f a s s e s s i n g t h e s i g n i f i c a n c e  Utilizing analyses  univariate  and G u i r e s  a s s o c i a t e d with  of s i g n i f i c a n c e  over  some p 1979).  %SS ( u v a )  are provided  are  f o r %SS  (mva).  During the v a l u e and  Maze  variation For  the completion  obtained  of t h i s  fortotal  1984) o f a g i v e n  example,  hypothesized been m e a s u r e d  given a simple  i t was o b s e r v e d  multivariate variation  source  of each v a r i a b l e  study  i s equal  t o be due t o a s i n g l e f o r three v a r i a b l e s ,  ( v , Scagel c  to the average  f o r t h e same h y p o t h e s i z e d situation  that  source.  where v a r i a t i o n i s  source,  A, and s a m p l e s  x-. : x, , x , x . 2  3  have  72  The  c a l c u l a t i o n of t o t a l  multivariate  would be ( s e n s u S c a g e l and Maze  v  can a l s o  c  This calculate  v a r i a t i o n due t o s o u r c e A  1984):  be c a l c u l a t e d by:  r e l a t i o n s i m p l i f i e s the a n a l y t i c steps necessary to %SS  (mva).  I n d e e d , t h i s r e l a t i o n p r o v i d e s a means by  which m u l t i v a r i a t e  v a r i a t i o n c a n be c a l c u l a t e d when a PCA c a n n o t  be  to either  performed  owing  dimensionality  the s t r u c t u r e  of the data.  previous univariate  studies  It also  p r o v i d e s a way  c a n be compared  studies.  A d d i t i o n a l l y , t h e same e m p i r i c a l  r  from  2  values  from  regression  multivariate the  regression  and u n i v a r i a t e  increases  scores.  relation exists for and r  of these  values  2  r e l a t i o n between  v a r i a t i o n serves to further  r e s u l t s with  the u t i l i t y  This  that  to multivariate  of i n d i v i d u a l v a r i a b l e s  o f component  i n c l u s i o n o f UVA  of R or  justify  t h e r e s u l t s o f MVA and  r e s u l t s for future  independent  analyses.  Although feature  reduction  o f PCA, O r l o c i  independent  specific  (1973,  means o f a s s e s s i n g  technique of ranking and  of d i m e n s i o n a l i t y  variance  variables  i s an e s s e n t i a l  1975, 1978) p r o v i d e s an dimensionality.  Orloci's  on a sums o f s q u a r e s  criterion  ( = r e d u n d a n c y ) was employed h e r e a s a means  73  of  providing  reduced  a summarization  character  undertaken  (i.e.  o f t h e d a t a u s e d and t o s u g g e s t a  set for u t i l i z a t i o n  an i n d e p e n d e n t c o r r o b o r a t o r y  (1975) p r o v i d e s an i l l u s t r a t i o n with  respect  significance  study).  of t h e u s e o f t h i s  and t e s t s  Beshir  technique  of s i g n i f i c a n c e .  h a s been made o f v a r i o u s employed  in different  stated  i n the text,  judged  t o be so a t a p r o b a b i l i t y  significance  sampling i s  to Pinus banksiana.  4.3 C o m p u t a t i o n Mention  i f further  a l l tests  are e x p l i c i t l y  tests  of s t a t i s t i c a l  analyses.  indicated  Unless otherwise  as being s i g n i f i c a n t a r e  o f p < 0.01.  referenced  L e v e l s of  i n a l l t a b l e s and  f igures. All  a n a l y s e s were p e r f o r m e d u s i n g  available  at the U n i v e r s i t y  programmes u s e d were: ANOVAR (Greig NTSYS size  and B j e r r i n g (Rohlf,  dispersion  (Greig  1980); MIDAS  e_t a l . 1980).  estimation  of B r i t i s h  and O r l o c i ' s  criterion  ranking  Emanuel of t h e F a c u l t y  British  Columbia.  plotting  routine  points.  As w e l l ,  1978);  of c h a r a c t e r s using  GENLIN  Columbia  sample  by a  a programme d e v e l o p e d a t the U n i v e r s i t y of  of t h e Botany  w h i c h accommodated several  Statistical  and u n i v a r i a t e  of F o r e s t r y  Bradfield  of B r i t i s h  and O s t e r l i n  Multivariate  by John  the U n i v e r s i t y  Columbia.  facilities  (Fox a n d G u i r e 1976); a n d ,  were p e r f o r m e d  Gary  the computing  provided  Department a t  a two-dimensional  identification  of p l o t t e d  programmes d e v e l o p e d a s s y s t e m  sub-routines a t the U n i v e r s i t y  of B r i t i s h  Columbia  were  used.  74  III.  1.  INTRA-INDIVIDUAL  Introduct ion.  In  morphologically  perception  and  small  assessment  inter-individual  of  variation  readily  apparent.  In  complex  organsims,  s u c h as  variation  i s not  inter-individual variation  of  statistical  As  as  as  1981), not  genetic that  can  and  be  a  the  of  assumptions  of  differences  of  the  and  is  assess.  Intra-individual  which  between  variation  or  genetic  such  an  variation-'srerves genetic  i s the are  another  level  buffered  are  the  However,  development.  genetic  variation  in  an  or such  As is  variation,  one.  intra-individual  assumption  "non-genetic" causes  of  of  of  genetic,  individual  interaction.  basis  localized  recognition  individuals  other  to  definition but  suspected  variation  of  respect  classical  i n the  s o u r c e s of  with  1984).  genetic  Implicit  often  anatomically  (Chapt. I I ) , developmental v a r i a t i o n  simply  n a t u r e of  purposes,  intra-individual  i s , by  against  the  and  result,  Talbert  amongst p a r t s  environmentally  The  a  to  genotype-environment  ignore  as  as  easy  result  compared.  mentioned e a r l i e r considered  trees,  nor,  as  somatic* m u t a t i o n s ,  result  less  spruce  large  developmental v a r i a t i o n ,  whereas d i f f e r e n c e s result  systematic  variation  developmental  heritable  i s , for  ( Z o b e l and  a comparative yardstick  variation  organisms,  i s most commonly c o n s i d e r e d  accuracy  e n v i r o n m e n t a l and  simple  intra-individual  apparent  variation  trees  and  morphologically  intra-individual  (Falconer  a  VARIATION.  better albeit  75  inter-individual  context  reasons o u t l i n e d  here.  being  used  in t h i s  i s important Having  s t u d y have a  intra-individual  variation  intra-individual  variation,  Tukey  (1977),  been  "sweep i t under  2)  rug".  intra-individual variation  by  developmental  assumes random  and  interpretation  variation.  I_n P.  o t h e r s have been e m p h a s i z e d Reference necessitated trees.  As  has  by  those p o s i t i o n s necessitated literature b a s e d on  the of  very  Hunt and  intra-individual  of  Garman  the  at  random  be  influenced  these p o i n t s  and  1980a).  sampling  problems  individual  complexity easily  the  in. even  sampled, Evidence  in  the  impressions  between  Addressing selected  ignore  be  strategies.  differences  the  orders, issue  variables  Parker,_et_al.  of  avoids  (1981)  by  (1983).  m o r p h o l o g i c a l and variation  i n P_;_  could  the  be  and  not  o c c u r r e n c e of  could  canopy. of  may  substantiates  those d i r e c t e d  F r a n c e and  wood anatomy  bole.  that  Picea  i n the  Rudloff  A number o f  (1979) and  on  variation  s u c h as von  tree  to  than  intra-individual  (1975b,  structural  s p e c i f i c sampling  ages  intra-individual critsicms  the  o b s e r v a t i o n s of  and  Forrest  form, and  attendant  available  field  positions,  size,  variables  Mosteller  sitchensis,  been made e a r l i e r  the  well,  by  the  However, t o  variation.  sampling  the  ignore  words of  variation  suggests that  that  could  intra-individual That  for  inter-individual  one  i n the the  consider  satisfied  larger  (Table or  to  of  Mexal  Picea  studies  to  of  have been c o n d u c t e d .  (1980) have r e l a t e d  sitchensis  (1957) r e l a t e d  anatomical  position  variation  of  the  i n the  scale  Denne  variability canopy  morphology  and to  76  position  i n the cone.  longitudinal  This variation  cone s e c t i o n s .  needle morphology w i t h i n  Taylor  i s seen r e a d i l y  (1959) r e p o r t e d  the canopy.  (1965) have  reported  in  ( L . ) K r a s t . and P^ p u n g e n s .  P_j_ a b i e s  positional also  variability  examined  direction reported  t h e most  intra-individual Similarly,  structures;  Funsch related  competition associated  Stover  variability Structural  (1963)  of branches  d e v e l o p more  not d e t e c t  sitchensis and l i g h t ,  not  quickly detect  intra-individual  of the canopy.  Variation  has been a t t r i b u t e d  growth  to  regulation  d o m i n a n c e , and t h e e x t e r n a l e n v i r o n m e n t  1944; W a r d l e  interest  1968;  has f o c u s e d  of p h y s i o l o g i c a l variation  L e v e r e n z and  h a s been  on t h e  Jarvis  intra-individual  and g r o w t h v a r i a b l e s  sometimes has been r e p o r t e d  Lewandowska and J a r v i s  Jarvis  orders  (1959) c o u l d  direction  in  these p h y s i o l o g i c a l  variation  compass  G r a c e , e t a l . 1975).  More r e c e n t l y  with  to  (1975)  i n Rocky M o u n t a i n P_^ e n g l m a n n i i .  t o compass  with a p i c a l  1927;  1980a,b;  they a l s o  (1975) c o u l d  for nutrients  to  of E"\ g l a u c a .  and l o w e s t  Taylor  variation  has been o b s e r v e d  (Larsen  apical  anatomy  Funsch  Wilson  and Owens 1983).  variation  variables,  i n t h e c a n o p y o f P_;_ e n q e l m a n n i i .  have t h e l a r g e s t (Harrison  In a d d i t i o n  related  of p o l l e n  Sifton  of needle  variability  on t h e v a r i a b i l i t y  on cone and  (1982) and  variation  of m o r p h o l o g i c a l  intra-individual  Generally,  that  on a g e - r e l a t e d  Ewers  in  studies 1978).'  related  1974), p o s i t i o n  in conjunction  ( L e v e r e n z and J a r v i s  Intra-individual  to order  (Baxter  in Picea.  of b r a n c h i n g  and C a n n e l l  1980a,b;  physiological (Norman  and  1978; F r a s e r , e t  77  al.  1964;  Jarvis  L e v e r e n z and  1978), and  Soikkeli  age  1978).  L e v e r e n z and reported  Jarvis  (Freeland  Schulze,  Jarvis  for  light  and  the  Of  intra-individual positions, reported  and  Fuchs,  With  the  by et  has  also  Phillips  P_^  be  1979;  been d i r e c t e d  abies  and  have  structural  by  competition  the  environment  relation  sunlight  ("sun  Zimmerman and  of  and  Brown  ( I 9 8 0 a , b ) i n P_;_  scale  1977;  growth.  and  been t h e  et  and  the  structure,  Schulze,  commercial  of  regulators,  Jarvis  P^  sitchensis  effected  incident  Kozloski  on  variation, like  to  the  to  a l . (1977) and of  and  increment  i n t e r e s t has  L e v e r e n z and  advent  attention  growth  variation  Kramer and  as  single  surrounding special  Fry  on  physiological  nutrients,  tree.  a  been h y p o t h e s i z e d  micro-environment of  1952;  (1980a) w o r k i n g  Intra-individual has  I980a,b; Lewandowska  e_t a_l. (1977) w o r k i n g  v a r i a t i o n within  variation,  1979,  to c y c l o - ,  1971)  sitchensis,  a l . (1977) i n E\  grafting  shade"  abies.  programmes peri-,  and  topophysis.  Some of variation  the  most e l e g a n t  in c o n i f e r s  contributions  of  and  Rudloff  ( 1 9 6 8 ) , and  Kaufmann, e t  emphasized  b o t h the  (1967,  at  aerial  portions  system  (Ford  and  a_l. ( 1 9 7 4 ) .  the  Deans  i n P^  the  a l . (1974), F o r r e s t  1975), O g i l v i e and  It should  least of  et  of i n t r a - i n d i v i d u a l  i n p a r t i c u l a r are  s p a t i a l and  c h e m i c a l compounds. variability,  Picea  Hrutfiord,  1980a), von  studies  These s t u d i e s  temporal be  noted  sitchensis,  tree,  but  also  1977;  E i s and  von  (1975a,b; Rudloff have  v a r i a t i o n of that  intra-individual  i s not  occurs Long  volatile  r e s t r i c t e d to  i n the  1972).  root  the  78  In  the p r e s e n t  intra-individual  study  there are several  variation  to  t h e age o f t h e i n c r e m e n t  of  branches.  coniferous 1983:  studies  b,  of growth,  model").  Indeed,  sitchensis  (Denne  1979; F r a n c e  have  formed  Since  encountered assessing  from  a variety  variation  branch  branches  the b a s i s  branches  from  the lowest from  statistical  result  primary  different  associated  a r e commonly  variation  nature  appreciation  of environmental  the lowest  primary  of p r i m a r y  branches;  whorl  whorl  and, w h o r l  i n the canopy.  respect to positions a d d r e s s i n g these  to inter-individual  of t h i s  i t will  The  i n the  s o u r c e s of  may n o t a l l o w t h e a s s i g n m e n t  of sampling,  branching  with:  and s e c o n d a r y  positions  Although  confidence limits  Adventitious  that  1975a,  o f t h e same age from t h e  of c o l l e c t i o n  whorl  examined.  the unbalanced  heuristic  branches  of cone morphology w i t h  intra-individual  1967; F o r r e s t  e m p h a s i s has been p l a c e d on  ( F i g . 7); primary  ( F i g . 7 ) ; the year  c a n o p y was a l s o  for entire  1980; S c h u l z e , e t  of p o s i t i o n s  primary  branches  variation  (Tomlinson,  intra-individual  i s i n the v a r i a t i o n  o f t h e same age from  branches  i n other  in vegetative materials associated  and w h o r l  lowest primary  primary  here  during c o l l e c t i n g ,  adventitious  primary  interest  of  relate  and o r d e r  architecture  1977; F r a s e r , e t a l . 1964; von R u d l o f f 1980a).  These  a r e common  studies  and M e x a l  s o u r c e s of  the p o s i t i o n ,  s o u r c e s of v a r i a t i o n  of  with material  to  c a n be q u a n t i f i e d .  s p e c i e s owing t o t h e s i m i l a r  "Massart's  variation  al.  These  that  obvious  of  variation  due  p e r m i t an  source of v a r i a t i o n .  in  inducement  sitchensis by e i t h e r  i s thought physical  t o be a  damage t o  79  the  tree  (Herman  1964) o r s t a n d damage  o b s e r v a t i o n s made i n t h e f i e l d , case  for  citation  effect  literature  indicating  anatomical  differences  positions  of branches  environments,  than  whorl  The  a r e o f t e n i n markedly  explicit  i s also  no  different  to expect  differences.  As t h e s e two  the e x i s t e n c e of  Adventitious  not u n i f o r m l y p r e s e n t , a r e e a s i e r  of branches  a number o f w o r k e r s . nature  of branching  determinant suggested  t o be t h e  t o sample  branches.  Orders by  There  t h e two p o s i t i o n s .  i t would be r e a s o n a b l e  although  Based on  t h e r e a r e m o r p h o l o g i c a l or  between  m o r p h o l o g i c a l and a n a t o m i c a l branches,  t h e r e h a s n o t been  i n the l i t e r a t u r e .  whether  1940).  t h e same w o u l d a p p e a r  engelmanni i a l t h o u g h to t h i s  (Issac  o f t h e form  significant  Where t h e s e invariably  have been more i n t e n s i v e l y I t i s an o b v i o u s of the t r e e  differences  as a  Some r e s e a r c h has  between o r d e r s o f b r a n c h i n g .  are observed  t o growth r e g u l a t i o n  of v a r i a t i o n .  i s interpreted  of the t r e e .  differences  source  investigated  they  are a t t r i b u t e d  a s s o c i a t e d with  apical  dominance. With  r e s p e c t t o temporal  literature  i s very  variation  extensive with  explanations  f o r such  variation  Explanations  for this  source  to e x t r i n s i c  factors  physical 1980). appear  that  this  (climatic  was c a r r i e d  source  o b s e r v a t i o n s and t e n d e r e d  (Stover  1944; A n d e r s s o n  of v a r i a t i o n  damage) and i n t r i n s i c As s a m p l i n g  both  i n c o n i f e r s , the  have been  changes, pathogen controls out over  of v a r i a t i o n  (mast  attributed  infestation,  seeding,  several  could easily  1965).  years  Silverton i t would  influence the  80  results. By  temporal  variation  reference  between two y e a r  o l d twigs  and n e e d l e s  during  separate  collection 1977  years.  year  F o r example, a two y e a r  during  1978.  extrinsic  initiation  Owens, e t a l . . 1977; S i n g h of v e g e t a t i v e  noted  that  there  i s a temporal  and  order  variation because  1976a,  persist  in  the canopy  that  organisms  events).  habit  1976).  that  years  source  growth  source  Indeed,  "vegetative  Quantifying  and f e r t i l e "  intra-individual  the p o s i t i o n  i t i s height  Certainly this i s  of i n t r a - i n d i v i d u a l  f o r within-canopy  morphology) and d e v e l o p m e n t a l  foliage,  i n Picea  of v a r i a t i o n i n  variation,  "heterophylly", are a t t r i b u t e d to environmental leaf  that  whorl, p o s i t i o n ,  a s s o c i a t e d with  from w h i c h a sample comes.  Explanations  be  of the organism  most c h a r a c t e r i z e s t r e e a r c h i t e c t u r e .  variation.  I t should  long. •  i s that  the most e x t e n s i v e l y documented  sequence and  A d d i t i o n a l l y the temporal  for eight  P e r h a p s t h e s i n g l e most o b v i o u s large arborescent  1977, 1979,1980;  a s s o c i a t e d with  (Fraser  be a s s e s s e d  the leaves  surrounding  a n d Owens 1981, r e g a r d i n g  component  from  1980 owing t o t h e  1976b,  because of t h e p e r e n n i a l  could  a t t h e end o f  1978 a n d 1979 ( s e e H a r r i s o n and  and r e p r o d u c t i v e  of branching  o l d vegetative  initiated  conditions  during  Owens 1983; Owens and M o l d e r  timing  during  and i n t r i n s i c  and e l o n g a t i o n  and e l o n g a t i n g  Such m a t e r i a l may d i f f e r  o l dmaterial collected  specific  initiated  made i n 1979 would have been  and e l o n g a t e d  two  i s made t o v a r i a t i o n  ("sun a n d s h a d e "  ( " j u v e n i l e and mature" p o s i t i o n s ) causes.  variation  i n P^ e n g e l m a n n i i  81  and  P_^ s i t c h e n s i s p r o v i d e s  more t a x a . immature  Addressing  aspects  of t h i s  i n d i v i d u a l s may a l l o w  variation  with  concerning such  a d e s c r i p t i o n of t h i s  respect  environmental  information  variation  f o r explanations  to various  competing  or developmental  may be o f v a l u e  1963; R o u l a n d  P. e n q e l m a n n i i (Bower useful  (Whitham  Attention  control.  anatomically  form. it  Further  species  of P i c e a  organisms.  variation form  (Grant  1965; L i n d q u i s t  there  i s a diversity  form  variation  For t h i s  i s poorly  various  more  than  s t u d y and  understanding  b a s e d on t h e d e m o n s t r a t i o n  under g e n e t i c  c o n t r o l i n many  1976, 1977; A l e x a n d r o v 1971;  1948; J a n k i e w i c z of o p i n i o n  r e l a t e d t o crown  variatiation.  and M i t t o n  (Schmidt-Vogt  reason  u n d e r s t o o d and q u a n t i f i e d ,  and S t e c k i  1976) a l t h o u g h  as t o the degree of g e n e t i c  Some have a t t r i b u t e d t a x o n o m i c  i n Picea  serves  i n t r e e s h a s been  i s reportedly  Roche  control.  of t r e e s with  and f u n c t i o n i n g o f m o r p h o l o g i c a l l y and  of i n t r a - i n d i v i d u a l  form  i n v e s t i g a t i o n s may be  I t i s e s s e n t i a l f o r an  has been assumed t o be i m p o r t a n t  Crown  such  o b j e c t i v e s demanded by a s y s t e m a t i c  A l t h o u g h crown  of p a t t e r n s  c a r r i e d out  1981).  complex  intra-individual  programmes  graft incompatibility  the i n t e r a c t i o n  practises.  the development  In a d d i t i o n ,  f o r P_^ s i t c h e n s i s o r  to intra-individual  the pragmatic  silvicultural of  1982).  i n understanding  pathogens  just  reported  of t h i s  in tree-breeding  1973), a l t h o u g h  p r o b l e m s have n o t been  i n mature and  hypotheses  where g r a f t i n g and r o o t i n g p r o j e c t s a r e b e i n g (Klaehn  variation for  s i g n i f i c a n c e t o crown  1977; J o n e s and B e r n a r d  1977) and  82  have  speculated  t o heavy  snow p a c k s  researchers P.  on a d a p t a t i o n  have  of v a r i o u s  (Alexandrov  emphasized  1971;  and Deans  1978;  1980), c o m p e t i t i o n (Ford  Lindquist  the a d a p t a t i o n  C o c h r a n e and F o r d  1948).  intrinsic Pollard  with  support  (Cannell  and Logan  frequently  1979)  of m e c h a n i c a l  and e x t r i n s i c c o n t r o l s have  speculation  of p h o t o s y n t h e t i c  1980a,b; Norman and J a r v i s 1979a,b; F i s h e r  and H i b b s  efficiency  1974;  Stecki  and  1976).  hierarchy which  reproduction  Crown  of u n i t s slight  and  Diggle  Horn  1976).  affect  1962;  proposed.  the stand  explanations  Fisher  stand  adaptation, structure,  (Brunig  1976;  trees  most  for  crown  Jankiewicz  Cochrane  changes  1971;  Tomlinson  form r e s p e c t  and F o r d  during  1982), w h i c h ,  trees  and  of a  the development  i n t e r a c t i o n amongst  form i s  composition,  c h a r a c t e r i s t i c s mentioned above.  and p h y s i c a l  The  and Honda  to consist  et a l . 1964;  i n t e r n a l and e x t e r n a l  of crown  1976;  ( L e v e r e n z and J a r v i s  1971;  form i s r e c o g n i z e d  (Fraser,  Honda  Both  e_t §_1.  r e s u l t i n s u b s t a n t i a l a l t e r a t i o n s t o t h e crown  and Moore  trees  1982).  agreed to influence  regeneration,  light  properties  (Cannell, been  and  concerns adapatation  R e g a r d l e s s of the h y p o t h e s i z e d generally  Ford  of  with other  (McMahon and Konauer  1974)  tendered  optimization  1976;  abies  Other  of t h e crown  for nutrients, competition  1976), and o p t i m i z a t i o n  associated  may  forms of P^  s i t c h e n s i s t o t h e i n t e r c e p t i o n of p r e c i p a t a t i o n  (Ford  in  crown  i n the  1978)  development  form  (Pearce  i n turn  may  These of i n d i v i d u a l stand.  83  2.  Materials  2.1  Materials.  All  methods.  i n v e s t i g a t i o n s were r e s t r i c t e d  considered Such a  and  t o be  limitation  inter-individual situation  standards  f o r the  simplifies  the  two  species  n a t u r e and  variation, permitting  under w h i c h  to i n d i v i d u a l s Chapt. I I ) .  expected degree  a more  intra-individual  (see  of  constrained  v a r i a t i o n can  be  examined.  All  the  considered one To  site do  could  not  be  encountered  intra-individual  variation  assessed  s i n g l e t r e e or a t  during  the  f o r any c o u r s e of  so w o u l d have n e c e s s i t a t e d  Adventitious individual and  s o u r c e s of  and  of  whorl primary  P_j_ e n g e l m a n n i i  secondary whorl  representative  of  whole t r e e  (Tree  218,  s i t c h e n s i s (Tree  P_^_ engelmann i i  (Tree  Inter-positional conditions. taxa crown  ( F i g . 8).  v a r i a t i o n was  sampling  been e i t h e r blown o v e r d u r i n g struck year P.  down by  a  old saplings  truck  (Tree  damage had  not  Primary  from an i n d i v i d u a l Appendix  compared  was  557,  at  obscured  a  under  representative  II).  Year  (Tree  Appendix  since 221,  f o r b o t h of  each whorl  II).  Six,  in t h e i r  ordering  of  the  the  the  trees  Appendix  single half-sib  the  three  i n t e r v a l s throughout  possible  a storm  representing  s i t c h e n s i s were sampled  weevil  II).  an  individual representative  r e g u l a r l y spaced  This  from  60705, A p p e n d i x I I ) .  S i n g l e , mature t r e e s  were s a m p l e d a t  sampling.  Appendix  527,  any  collections.  b r a n c h e s were compared  t o y e a r c o m p a r i s o n s were made f o r an of  the.field  b r a n c h e s were compared  P^  being  had  II)  or  eleven  family  1  of  canopies  branches.  where  84  F i g u r e 8. S c h e m a t i c r e p r e s e n t a t i n of I n d i v i d u a l t r e e s o f P_^ enge1manni i and s i tchens i s from which i n t r a - i n d i v i d u a l sampling conducted T1 - T 6 , 1 1 - y e a r o l d i m m a t u r e P_^ s i t c h e n s i s f r o m n u r s e r y : lateral b r a n c h I n c r e m e n t s shown a r e a v e r a g e d p e r w h o r l , c i r c l e s r e p r e s e n t whorl nodes. T 7 , m a t u r e P_^ s i t c h e n s i s (SXP 2 2 1 ) . T8. mature P• e n g e l m a n n i 1 (SXP 5 2 7 ) . T r i a n g l e s mark p o s i t i o n a t w h i c h s a m p l e s taken from mature t r e e s . Not a l l w h o r l p r i m a r y b r a n c h e s m a r k e d f o r i m m a t u r e t r e e s due t o m e a s u r e m e n t p r o b l e m s a s s o c i a t e d w i t h w e e v i l damage. * , p o s i t i o n s cones c o l l e c t e d from.  Including  immature t r e e s  inter-positional developmental.  allows  v a r i a t i o n may  a t e s t o f whether  be e n v i r o n m e n t a l o r  I f v a r i a t i o n i s a r e s u l t of e n v i r o n m e n t a l  causes,  t h e n mature and immature t r e e s  display  similar patterns  Including  would n o t be e x p e c t e d t o  of i n t r a - i n d i v i d u a l v a r i a t i o n .  immature i n d i v i d u a l s a l s o a l l o w s  m o r p h o l o g y and anatomy w i t h whether, w i t h i n substantial  that  o f mature t r e e s  the c i r c u m s c r i p t i o n  differences  that  a comparison  could  of a given r e s u l t from  p e r s . comm., U l f B i t t e r l i c k ; B r i t s h C o l u m b i a F o r e s t s , C h i l l i w a c k River Nursery 1  of  their  and t e s t  taxon,  there are  t h e age o f t h e  M i n i s t y of  85  trees.  The and  mature t r e e s  edaphic  disposition  populations. had  the  The t r e e s  been s e v e r e  saplings  loss  sampled  British  collected  appeared  to neighbouring were sampled  of the c u r r e n t  similar trees  i n l a t e autumn  the height,  individuals  For the  2.2  examining  context  included  i s given  of F o r e s t s  N u r s e r y of  I I : trees  Relevant  information  positions  of the  8.  t h e n a t u r e of i n t r a - i n d i v i d u a l v a r i a t i o n i n  of i n t e r - i n d i v i d u a l v a r i a t i o n ,  that  there  The  River  (Appendix  a g e , form, and s a m p l i n g  in Figure  before  y e a r ' s cone c r o p .  71601, 71602, 71603, 71604, 71605, 7 1 6 0 6 ) . concerning  i n the  were g r o w i n g a t t h e C h i l l i w a c k  Columbia M i n i s t r y  i n a g e , form,  were s t a n d a r d s o f  individual  trees  were  e n g e l m a n n i i a n d P_;_ s i t c h e n s i s .  Analyses. PCAs b a s e d upon c o r r e l a t i o n  variables  were p e r f o r m e d .  samples a v a i l a b l e , averaging  could  dimensionality were Where  first  m a t r i c e s of s e p a r a t e  Owing t o t h e r e s t r i c t e d number of  combining  separate  n o t be u s e d w i t h o u t constraints  tested  o f PCA.  suites  A l l data  submitted  f o r i n d e p e n d e n c e of c o r r e l a t i o n  analyses  Evaluating  of v a r i a b l e s  of t h e i n d i v i d u a l  differences  therefore  variables  o r component  t o PCA  coefficients. not necessary,  was  performed.  between v a r i o u s p o s i t i o n s  upon ANOVAs o f o r i g i n a l v a r i a b l e s  by  i n v a l i d a t i n g the  i n d e p e n d e n c e was a c c e p t e d and PCA  univariate  s u i t e s of  scores.  was  based  The  86  ANOVA m o d e l s u s e d a r e g i v e n  12.  that  owing  that  r e f e r s t o d i f f e r e n c e s among t a x a .  prevented from  to the p a u c i t y  in Table  a s t r u c t u r e d MVA,  being  Differences  these  of  o f samples  s u c h a s t r u c t u r e d MVA  Mahalanobis' D  2  distances  trees.  per p o s i t i o n a r e suboptimal  utilization  size  of D  trend  inspections  would  2  estimation may  rather  of D  suffice than  2  Although (Table  t h e sample  such a  i n g i v i n g an i m p r e s s i o n  specific  detail.  i n the data.  was p e r f o r m e d p r o v i d e d line  could  of d e t e r m i n i n g  of the  Preliminary scores  against  It an  trend  regression  d i s p e r s i o n o f samples a r o u n d t h e  distances  overall  normally  distributed  were a l s o c a l c u l a t e d a s a means  similarity  per i n d i v i d u a l  allowing  were a p p a r e n t ,  whorl  1977).  for inter-positional  available for  2  that  be c o n s i d e r e d  ( C h a t t e r j e e and P r i c e  Mahalanobis' D  If a trend  sizes  10), the behavior  was s u c h t h a t  of s c a t t e r p l o t s of component  were p r e s e n t  sampled  have  between a p i c a l - and  or p o s i t i o n were a l s o made t o d e t e r m i n e whether a g e n e r a l  regression  CVA,  p o s i t i o n s were c a l c u l a t e d a s a means o f c o m p a r i n g  t h e sample  general  The l a c k  i s no t e r m  appropriate.  p o s i t i o n s i n the various  available  there  between p o s i t i o n s i n t h e canopy were examined  two manners.  basal-most  that  be n o t e d  s u c h a s MANOVA and s u b s e q u e n t  performed; although  been a n a l y t i c a l l y  in  of m a t e r i a l s ,  I t should  of the i n d i v i d u a l  variation.  tree appeared  The sample (Table  trees sizes  10) t o be a d e q u a t e  such use D . 2  was i m p o s s i b l e  inter-individual  to quantify  intra-individual  or inter-taxonmic  context  using  variation in a single  T a b l e 12. ANOVA m o d e l s u s e d i n a s s e s s i n g v a r i o u s a s p e c t s o f i n t r a - i n d i v i d u a l v a r i a t i o n . v a r i a b l e v a l u e s o r component s c o r e s f o r a g i v e n s a m p l e : e - r e s i d u a l v a r i a t i o n .  SOURCE  MODEL  ADVENTITIOUS/  PRIMARY/  YEAR/  WHORL  SECONDARY  YEAR  INTER-POSITIONAL  y = A + e.  y = A + B(A)  branch p o s i t i o n : a d v e n t i t i ous whor 1  + e.  A - branch order: p r i mary secondary B(A) - b r a n c h w i t h i n o r d e r .  y = A + e.  c o 1 1 e c t i on y e a r  y = A + B ( A ) + e .  A + B(A)  VARIABLE  TERMS  + C(AB)  + e.  y  -  SUITE  TWIG MORPHOLOGY LEAF MORPHOLOGY LEAF ANATOMY  TWIG MORPHOLOGY LEAF MORPHOLOGY LEAF ANATOMY  TWIG MORPHOLOGY LEAF MORPHOLOGY LEAF ANATOMY  A - tree B(A) - p o s i t i o n  in  A - tree B(A) - p o s i t i o n C(AB) - cone i n  in t r e e position  tree  TWIG MORPHOLOGY LEAF MORPHOLOGY LEAF ANATOMY  CONE MORPHOLOGY  88  ANOVA a s s y s t e m a t i c undertaken. based  sampling  A comparison  from PCAs i n c l u d i n g  source of i n t r a - i n d i v i d u a l  the  is Table of  13.  the i n d i v i d u a l  that  variation.  respect  trees,  not be  of i n t e r e s t .  on t h e b a s i s  nature  Differences  of t h e change i n  of t h e ANOVAs a r e g i v e n  of v a r i a b l e s  circumscription  whose  d i d not m a r k e d l y d e c r e a s e  variation  were t o be p r e f e r r e d  were d r a s t i c a l l y a l t e r e d I t would be p r e d i c t e d  ANOVAs  and e x c l u d i n g t h e  t o the taxonomic  suites  variation  inter-individual  suites  The e x a c t  Without  intra-individual of  variation  t h e s e ANOVAs were a s s e s s e d  sums o f s q u a r e s .  could  was made between two d i f f e r e n t  on component s c o r e s  between  of a l l i n d i v i d u a l s  t h e amount  to those  by t h e i n t e r - i n d i v i d u a l  that  adding  s o u r c e s of  T a b l e 13. ANOVA m o d e l s u s e d i n a s s e s s i n g t h e c o n t r i b u t i o n o f i n t r a - i n d i v i d u a l v a r i a t i o n i n the c o n t e x t of i n t e r - i n d i v i d u a 1 variation. y - v a r i a b l e v a l u e f o r component s c o r e f o r a g i v e n sample: e - r e s i d u a l v a r i a t i o n .  MODEL y  = A +  VARIABLE  TERMS A -  e.  TWIG LEAF CONE LEAF  individual  or A -  intra-individual  variation  should decrease  due t o h y p o t h e s i z e d  possible,  specifying  of  taxa,  a priori  prevented taxonomic  MORPHOLOGY MORPHOLOGY MORPHOLOGY ANATOMY  taxa: P. sitchensis P. enqe1mann i i  variation  than  SUITES  t h e ANOVA  t h e amount o f  inter-individual  sources.  Where  i n t e r m s of i n d i v i d u a l s ,  rather  the unwarranted hypotheses.  imposition  and  reification  89  To  serve  variation, an  a s an h e u r i s t i c  o r d i n a t i o n s o f component  heuristic  scores  separate  context.  PCAs o f s e p a r a t e  dispersion  of samples over  ordination  facilitates  analyses  that  among  by t h e l a r g e s t  analyses.  Such  individual  analyses.  Results.  The  sample  a reliable  thus  Further,  10) were n o t s u f f i c i e n t  tests  cysts  perform  indicated that  there  was no  significant  be c o n s i d e r e d  independent,  unnecessary. 14, i n d i c a t e d t h a t o n l y  (RESCYLEN) c o u l d  between  the average  be c o n s i d e r e d  l e n g t h of  significantly  w h o r l and a d v e n t i t i o u s b r a n c h e s - t h o s e  whorl branches being  longer  (2.6 mm)  (1.2 mm).  than  those  adventitious  branches  Of  d i d not i n d i c a t e h e t e r o s c e d a c i t y  variances  to  of t h e independence of  that a l l v a r i a b l e s could  ANOVAs, T a b l e  different  branches.  among t h e g r o u p s of v a r i a b l e s i n t h e v a r i o u s Rs,  making PCA  resin  whorl primary  (Table  coefficients  co-variation indicating  sizes  PCA.  correlation  the  variation in  t r e e s , the s c a l e s of the  a l l separate  comparison  3.1 A d v e n t i t i o u s v e r s u s  the  were made t o p r o v i d e  In t h o s e  o r d i n a t e and t h e a b s c i s s a were d e t e r m i n e d  3.  intra-individual  a p p r e c i a t i o n f o r the i n t r a - i n d i v i d u a l  s u c h an i n t e r - i n d i v i d u a l compared  a p p r e c i a t i o n of  Bartlett's  test  of the  of the of the e q u a l i t y  of v a r i a n c e s  among  two p o s i t i o n s .  With the exception  of the s i n g l e  significant difference,  90  T a b l e 14. ANOVAs o f I n d i v i d u a l v a r i a b l e s c o m p a r i n g a d v e n t i t i o u s a n d whorl p r i m a r y b r a n c h e s of engelmanni1. *. s i g n i f i c a n t F-values @ p < 0.01. O t h e r s y m b o l o g y g i v e n i n T a b l e 12.  VARIABLES  LEAF MORPHOLOGY NEEDLEN ABXSTOM ADXSTOM RESCYNO RESCYLOC RESCYLEN x  (n-10)  TWIG MORPHOLOGY PULVLEN TIPWID TIPDEP PULVPUB  ( n = 10)  17.58  1 .45 0.00 2 . 44 i nvar i ate 1 . 30 x  SUITE  LEAF ANATOMY NEEDWID NEEDEP ABXANG ADXANG CENCYWID CENCYLAT CENCYABX CENCYADX ENDONUM PHLENO XYLEND ~  38.38 0.50 0.67 9.09 0.58 53.68*  TOTAL  the  VARIABLE  %SS-  SUITE  %SSc  (n=10) 7.93 749 20.41 2.11 2.14 22.17 2.72 20.29 26.60 18.55 9.35 12.71  10.. 53  a v e r a g e amount o f v a r i a t i o n r e s u l t i n g from  between t h e two o r d e r s accounted  f o r only  intra-individual repeatability provided  about  10 p e r c e n t  for a l l suites  the best  2).  As a g r o u p ,  separation  appeared  a s t h e measurement  l e a f morphology  o f t h e two p o s i t i o n s  to vary  of v a r i a b l e s  of the t o t a l  v a r i a t i o n - twice as l a r g e  (Table  Twig m o r p h o l o g y  of branches  differences  little  with  variables  of branches.  respect  to branch  posi t ion.  3.2 P r i m a r y v e r s u s ANOVAs, of  orders  branches.  that  differences  between  orders  s i g n i f i c a n t , c o n s i s t e n t l y accounted f o r  v a r i a t i o n i n the data  branches.  whorl  15, i n d i c a t e d  branches, although  less  of  Table  secondary  than d i f f e r e n c e s  between  The v a r i a t i o n a t t r i b u t e d t o d i f f e r e n c e s  o f b r a n c h e s was e x t r a c t e d  primarly  t h e PCAs, an i n t e r p r e t a t i o n s u p p o r t e d  individual between  by t h e f i r s t  component  by o r d i n a t i o n s i n  T a b l e 15. M u l t i v a r i a t e apportionment of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between o r d e r s of b r a n c h e s . Abbreviation o f PCAs a n d ANOVAs i n T a b l e 3 9 , A p p e n d i x I I I . Symbols g i v e n i n T a b l e 12. O r d i n a t i o n o f r e s u l t a n t component s c o r e s g i v e n i n F i g u r e 9.  %SS VARIABLE  Figure  larger  E  35.32 36.97 44.95  46.31 55.36 21.65  x VEGETATIVE  19.82  39.08  41.11  Component  correlations  measured on p r i m a r y  and mean v a l u e s s u g g e s t  whorl  those of the secondary  anatomy a p p e a r s  for  (mva) B(A)  18.37 7.67 33.42  than  evidenced  A  LEAF ANATOMY LEAF MORPHOLOGY TWIG MORPHOLOGY  9.  variables  SUITE  t o vary  by t h e v e r y  angles of the leaf  i n more  branches branches.  than  just  were  between t h e two o r d e r s o f b r a n c h e s  consistently  However,  leaf  s i z e of v a r i a b l e s as  low and n e g a t i v e component s u r f a c e (ABXANG,  that  correlations  ADXANG).  Differences  was most e m p h a s i z e d  by t w i g  morphology.  F i g u r e 9. O r d i n a t i o n s o f f i r s t two c o m p o n e n t s o f PCAs o f s e p a r a t e v a r i a b l e s u i t e s of whorl p r i m a r y and s e c o n d a r y b r a n c h e s of P. S i t c h e n s i s . S c o r e s f r o m PCA g i v e n i n T a b l e 3 9 . A p p e n d i x III. Open c i r c l e s - w h o r l p r i m a r y b r a n c h ; f i l l e d c i r c l e s - w h o r l s e c o n d a r y branch. Glyphs represent i n d i v i d u a l samples.  TWIG MORPHOLOGY  LEAF MORPHOLOGY  LEAF ANATOMY  K>  O CD  cP •  c0* o  8°  O CD * «D •O  The  total  variation  •  for a l l variables  •O ° O  over a l l s u i t e s of  92  variables about  of d i f f e r e n c e s  20 p e r c e n t and t h a t  branches The  as a r e s u l t  within  difference  difference  With  attributed  an o r d e r was n e a r l y  between b r a n c h  o r d e r s was  to differences  between  twice  ( i . e . 40%) a s l a r g e .  between o r d e r s was t w i c e a s l a r g e  between a d v e n t i t i o u s  and w h o r l  a s t h e mean  branches  (Table 14).  t h e e x c e p t i o n of the twig morphology v a r i a b l e  intra-branch proportion  (residual) variation  of the t o t a l  variation  accounted  f o r a much  suite, larger  t h a n any o t h e r h y p o t h e s i z e d  source of v a r i a t i o n . Ordinations consistent  of i n d i v i d u a l  relation  b r a n c h means  between t h e s e c o n d a r y  given  primary  ( i . e . p r i m a r y and s e c o n d a r y  whorl  primary  branch d i d not appear  secondaries  from  the  o b t a i n e d from ANOVAs  that  results  t h e r e was l i t t l e  branches,  except  emphasizes  that  branches samples  3.3 Year The 10).  other branches).  based  the v a r i a t i o n  branches  more s i m i l a r  from  than  The o r d i n a t i o n s o f component  t h e same  re-inforce  s c o r e s ( T a b l e 15)  between t h e two o r d e r s o f  among samples  from  Figure 9 also secondary than  branches.  variation. s i z e s were i n s u f f i c i e n t  Additionally,  tests  of R i n d i c a t e d  overall  t o a l l o w a PCA ( T a b l e  of the independence that  variation  of c o r r e l a t i o n  the pulvinus,  morphology, and cone morphology v a r i a b l e significant  subtending a  on t w i g m o r p h o l o g y were more v a r i a b l e  to year  coefficients  branch  i n the case of twig morphology.  from p r i m a r y  sample  separation  ( F i g . 9) showed no  suites  leaf had no  i n the data, suggesting that  93  variables  i n t h e s e g r o u p s were  One-way variables results the  ANOVA  distinguished  indicate  years  (Table  that  for only  16) t e s t e d  some  of the  between t h e two d i f f e r e n t  years.  7 variables:  SUITE  LEAF MORPHOLOGY NEEDLEN ABXSTOM ADXSTOM RESCYNO RESCYLOC RESCYLEN TWIG MORPHOLOGY PULVLEN TIPWID TIPDEP PULVPUB X  .90 . 1 1 OO .02* . 33 .82  (n=10) 81 .09* 55 . 17* 7 .44 1nvar i a t e 47 . 90  X  between y e a r s a c c o u n t e d  intra-individual  variation.  i s , i n comparison  branch p o s i t i o n  This  The between  TIPWID,  The a v e r a g e amount of  v a r i a t i o n of  VARIABLE  (n=10)  TOTAL  variation  to year  20.03  X  variation  SCALEN.  %SSfl  12 10 4 51 38 3  differences  RESCYNO, PULVLEN,  16. ANOVAs c o m p a r i n g y e a r enge1mann i i .  VARIABLES  whether  t h e r e were s i g n i f i c a n t  NEEDWID, CENCYLAT, CONWID, AND  Table P.  independent.  variables  SUITE  LEAF ANATOMY NEEDWID NEEDEP ABXANG ADXANG CENCYWID CENCYLAT CENCYABX CENCYADX ENDONUM PHLEND XYLEND ~  in  %SS  A  (n= 10) 59 . 37* 13 . 45 3 , 80 7 . 40 0.. 38 73 . 3 5 * 0.. 26 26 .44 0 OO 1 1. 36 . 5 . 77 18 . 33  CONE MORPHOLOGY <n = 34 ) 1 .50 CONLEN 46 .04* CONWID 27 . 5 4 * SCALEN 2 . 35 SCALWID 2. 1 1 SCALTAP 2 . 82 WINGWID 1 . 34 WINGTAP 3 . 27 FREESCAL BRACTLEN 0 . 84 3 . 10 BRACTWID BRACTAP 0 . 17 31.01 29.34  f o r 30 p e r c e n t amount o f  w i t h measurement  ( 1 1 % ) , and b r a n c h o r d e r  of the t o t a l  intra-individual repeatability,  (20%) a l a r g e  s o u r c e of  94  variation.  However,  intra-individual 15).  year  suggested  3.4  even  that  t h e two  variable  there  were  variables  source  increment  values  no  variable  were  of  variation  (Table  f o r t h e two y e a r s  consistent  larger  which  a  trends  (i.e.  values  than  significantly  of one the  different  years).  Inter-positional  amongst  large  as w i t h i n  consistently  those  Overall  variation.  eigen-structure  a l l variables  component  scores  variation  was  17).  o f mean  d i d not have  other, over  variation  Examination  growth  i t i s not as  i n each  indicated  attributed  However,  indicated variable  that  significant suite.  the largest  to inter-individual  considering  a l l  variation  ANOVA  single  of  source  differences  intra-individual  sources  of (Table  of  T a b l e 17. M u l t i v a r i a t e a p p o r t i o n m e n t of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between p o s i t i o n s of b r a n c h e s . A b b r e v i a t i o n o f PCAs a n d ANOVAs i n T a b l e 4 0 , A p p e n d i x I I I . Symbols g i v e n i n T a b l e 12. O r d i n a t i o n s of r e s u l t a n t component s c o r e s g i v e n i n F i g u r e 10..  VARIABLE LEAF LEAF TWIG CONE  ANATOMY MORPHOLOGY MORPHOLOGY MORPHOLOGY  x VEGETATIVE x TOTAL  variation, that  only  exceeded  hypothesized  A  %SS B(A)  . 30 . 96 . 23 .64  22 .90 23 . 90 46 .59 11.91  48.91 43 . 59  31.13 26 . 33  SUITE  leaf  55 44 46 27  anatomy  had an  intra-individual. t o be  the r e s u l t  of  The  (mva) CUB)  E 2 1 . 57 31.14 7.18 17 .49  42 .97  19 . 96 30.09  inter-individual largest  variation  inter-positional  variation was  effects  95  manifest  by t w i g  morphology.  cone morphology  (Table  The l a r g e s t s o u r c e  17) was a t t r i b u t e d t o d i f f e r e n c e s between  cones a t a s i n g l e p o s i t i o n i n the canopy. cones of  from d i f f e r e n t  whorls provided  D i f f e r e n c e s between  the second  largest  source  variation.  Mahalanobis' D branches in  o f v a r i a t i o n of  leaf  (Table  distances  2  between a p i c a l -  18) i n d i c a t e t h a t  there  anatomy w i t h i n mature t r e e s  and b a s a l - m o s t  were g r e a t e r  than  differences  immature t r e e s .  The  T a b l e 18. Mahalanobis' D d i s t a n c e s between a p i c a l - and b a s a l - m o s t b r a n c h e s a n d c o n e s o f P_^ e n g e 1 mann i i a n d P_^ s i t c h e n s i s . Diagonal e l e m e n t s (+), i n t r a - i n d i v i d u a l d i s t a n c e s between a p i c a l - and basal-most branches. Lower o f f - d i a g o n a l e l e m e n t s , d i s t a n c e s b e t w e e n basal branches. Upper o f f - d i a g o n a l e l e m e n t s , d i s t a n c e s between apical branches. Is, immature s i t c h e n s i s (T1 - T G ) ; Ms, m a t u r e p. s i t c h e n s i s ( T 7 ) : Me, m a t u r e P_^ e n g e 1 m a n n i i ( T 8 ) . P.enge1mann i i a n d P_^ s i t c h e n s i s. Note: D v a l u e s c o u l d n o t be c a l c u l a t e d f o r t w i g m o r p h o l o g y due t o m a t r i x singularity. 1  !  LEAF  ANATOMY  Is  LEAF Ms  Me  Is  Ms  Me  10.81  18.82  18 .04  20. 53  19 . 28  Is  3 . 10  34 . 50  77 . 82  Ms  5 . 23  41 . 98  44 . 23  2 .89  7 3.90  20. 12  Me  205.92  199 . 4 1 .  CONE  the  1 1 .83  2 . 39  MORPHOLOGY Ms  individual  MORPHOLOGY  Ms  43.76  Me  43.47  Me +  65.30 8.80  o f E\ e n q e l m a n n i i was d e c i d e d l y  different  mature and immature I \ s i t c h e n s i s , b u t o n l y  with  from  both  respect  to  96  leaf  anatomy. These d i s t a n c e s  differences  also  indicate  between t h e b a s a l  branches.  Leaves  were l e s s  similar  that  t h e r e were  branches than  from t h e b a s a l  most  t o those of  sitchensis  branches.Differences  were most e m p h a s i z e d  by c o m p a r i n g  Inter-individual  basis in  of l e a f  leaf  positions  and cone m o r p h o l o g y .  P_z_ s i t c h e n s i s  the a p i c a l  than  engelmannii  were  between  of both  leaves  t h e two t a x a  anatomy o f  differences  c o n s p i c u o u s when a p i c a l - m o s t  between  branches of  from t h e a p i c a l most  branches.  greater  basal  t a x a were most  were compared  on t h e  Inter-individual  a r e most o b v i o u s when c o m p a r i n g  differences  apical  branches.  Figure difference Further, greater  10, i n d i c a t e s  Figure  and  immature  2  needles.  that  there  This  pattern  i n Table  branches.  needles  i s evident  a  t h a n among  i n both  mature  19.  of d i f f e r e n c e s  These r e s u l t s  b a s e d on M a h a l a n o b i s '  indicate  exist  that the  between  the s i n g l e  o f P^ e n g e l m a n n i i and a l l i n d i v i d u a l s o f .  P^ s i t c h e n s i s  considering  discrete  trees.  P. s i t c h e n s i s  difference  a  i s consistently  the apical-most  inter-individual differences  individual  of  amongst  o v e r a l l assessment  i s given  largest  10 i n d i c a t e s  dispersion  basal-most  D  there e x i s t s  between t h e a p i c a l - and b a s a l - m o s t  the  The  that  By c o n t r a s t , are t r i v i a l  resulting  among  individuals  and t h e r e a p p e a r s t o be no  from t h e age e x c e p t ,  l e a f morphology.  P. s i t c h e n s i s  the d i f f e r e n c e s  That  i s no l e s s d i f f e r e n t  perhaps, i n  i s , the mature from  i n d i v i d u a l of  immature t r e e s  than  97  F i g u r e 10. O r d i n a t i o n s o f f i r s t two c o m p o n e n t s o f PCAs o f s e p a r a t e v a r i a b l e s u i t e s o f w h o r l b r a n c h p o s i t i o n s f r o m P_^ e n g e l m a n n i i and P_^ s i t c h e n s i s S c o r e s f r o m PCAs g i v e n i n T a b l e 4 0 , A p p e n d i x III. Open g l y p h s , a p i c a l - m o s t b r a n c h e s ; f i l l e d g l y p h s , basal-most positions. C i r c l e s - P^ s i t c h e n s i s; t r i a n g l e s - P^ e n g e l m a n n i i . I n t e r v e n i n g sampled p o s i t i o n s o m i t t e d from o r d i n a t i o n to emphasize polarity. Glyphs represent i n d i v i d u a l samples.  LEAF  differences With  between respect  apical-most p.  P.  had p u b e s c e n t  (Tree  70301, A p p e n d i x fallen  that  immature  pulvinii.  this  532, A p p e n d i x  trees. i t was o b s e r v e d t h a t  mature  f o r the apical-most branches  sitchensis  TWIG MORPHOLOGY  to branch pubescence,  b r a n c h e s were a l s o  on  individual  ^  b r a n c h e s o f t h e sampled  sitchensis  existed  ANATOMY  This  II).  situation  was more  situation  Similarly,  o b s e r v e d on an i n d i v i d u a l  from o t h e r t r e e s  same  of  from a p u t a t i v e  I I ) from t h e Ottawa V a l l e y .  branches  individual  the  pubescent  o f P_;_ g l a u c a  (Tree  O b s e r v a t i o n s made  i n the study area suggested  frequent  t h a n t h e samples  available  98  T a b l e 19. Average Mahalanobis' D d i s t a n c e s between i n d i v i d u a l t r e e s in T a b l e D v a l u e s below d i a g o n a l ( .--) - d i s t a n c e s between i n d i v i d u a l t r e e s . S y m b o l s g i v e n i n F i g u r e 8 a n d T a b l e 18. !  !  LEAF  ANATOMY  T1 T2 T3 T4 T5 T6  3 .06 5 .61 . 7 .66 . 6 . 88 10..01  T7  G . 99  Me T8  56 . 38  Is  Ms  LEAF Is  Ms  . 22 .59 .85 . 33  7 .61 9 . 42 1 1 . 69  2 . 39 1 1. 94  -7 . 18  5 . 74  13 . 23  1 1 .61  5 . 81  8 . 55  47 .06  G6 . 23  56 ..28  5 9 . 18  . -2.69 6.62  .-7.68 15.14  51 .49  5 . 88 0 . 95 2 . 18 3 . 77 1 1 .66.  T7  2 0 .. 77  19 .84  22 . 40  22 . 24  38 . 05  25 .65  39.73  39.39  TWIG  6 6 3 6  .08 .02 . 37 .88  0 . 87 4.20 9.4 1  31.54  MORPHOLOGY  T1 T2 T3 T4 T5 T6  3 .67 3 . 16 0 . 32 1 . 24 8 . 54  0 . 56 3 . 76 2 . 83 1 . 97  3 . 72 1 . 59 1 . 87  1 . 38 9 . 87  6 . 85  __  T7  4 .81  2.14  2. 1 1  4 . 02  2 . 26  6.12  Me T8  88 .95  88 .02  83 . 54  95.99  Ms  53 . 77  MORPHOLOGY  T1 T2 T3 T4 T5 T6  Me T8  Is  4 3 2 6  T 1  90.4 1 T2  87 .61 T3  T4  T5  T6  7 1 . 46 T7  .T8  18.  99  would  suggest.  In t h e s e  were p u b e s c e n t ,  As  the basal  differences performed  The  variation,  between  from t h e s e  first  largest  all  morphology trees).  percent  variation for  any o t h e r The  21)  variable  suggested  that  inter-branch variation  variation  emphasizes initial  the  were l e s s  between  samples. generally  cone  than 17  morphology  inter-positional  trend  ANOVAs  variation  exceeded  components.  within  (Table  scores  intra-branch  i n Figure  11 w h i c h  a n d between t r e e s .  (Fig.  position 12).  f o r each p o s i t i o n  b a s e d on B a r t l e t t ' s  that  Examples of the type  scores against  f o r each t r e e  o f component  correlations  suggested  anatomy a r e g i v e n  o f component  equal,  over  components were r e f l e c t i n g s i z e and  the s t r i k i n g v a r i a t i o n  variances  considered  the f i r s t  shown by l e a f  s u g g e s t e d a common  anatomy, and  f o r cone m o r p h o l o g y , a v e r a g i n g  (83%) t h a n  f o r the f i r s t  scatter  leaf  suite.  differences  only  that the  to differences  r e l a t i v e m a g n i t u d e o f t h e component  shape d i f f e r e n c e s  of  differences  Intra-position  was much l a r g e r  (Table 20).  74%, 53%, a n d 51% r e s p e c t i v e l y  differences  over a l l t r e e s .  The component  PCAs i n d i c a t e s  was a t t r i b u t e d  Inter-positional  intra-positional  suite.  f o r twig morphology,  (averaging  PCAs were  e x a m i n e d by ANOVA  from e a c h o f t h e s e  of v a r i a t i o n  p a t t e r n s of  suggested s i g n i f i c a n t  separate  f o r each v a r i a b l e  between w h o r l p o s i t i o n s leaf  trees,  PCAs were t h e n  component  source  a n d t h e ANOVAs  branches  glabrous.  may have c o n v o l u t e d  individual  f o r each t r e e  where t h e a p i c a l  branches appeared  t h e PCA o f a l l t r e e s  intra-individual  scores  situations  test,  An  or whorl  However a s could  regression  and  n o t be  T a b l e 20. M u l t i v a r i a t e a p p o r t i o n m e n t o f v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s o f s e p a r a t e t r e e s due t o whorl p o s i t i o n s . A b b r e v i a t i o n o f T a b l e 41 i n A p p e n d i x I I I . S y m b o l s g i v e n i n T a b l e 12 a n d F i g u r e 8. O r d i n a t i o n s o f r e s u l t a n t component s c o r e s g i v e n i n F i g u r e s 12 a n d 13.  %SS Is T 1 VARIABLE  ANATOMY  LEAF  MORPHOLOGY A 43 . 01 e 56 . 69  TWIG  MORPHOLOGY A 94 . 25 e 5 . 75  CONE  MORPHOLOGY A B( A) e  VEGETATIVE  x TOTA 1  T3  T4  T5  T6  Ms T7  Me T8  X  SUITE  LEAF  x  T2  (mva)  A 67 . 89 e 32 . 1 1  A 68 . 38 e 31 . 62 A e  53 . 39 46.61  6 0 . 72 39 . 28  67 . 82 32 . 20  53.21 46 . 79  35 . 16 64 . 84  59 . 05 4 0 . 95  29 ,90 70 . 10  53 . 39 46 .61  41 .89 58 . 1 1  42 .93 57 .07  42 . 92 57 . 08  7 2 . 89 27.11  52 47 47 .53  82 . 79 17 . 21  27 . 30 72 . 70  50 78 49 . 23  74 . 84 2 5 . 16  30. 9 0 6 9 . 10  74 . 14 25 . 87  24 . 85 62 . 72 12 . 42  8. 92 63 . 87 27 . 21  16 .89 63 . 30 19 . 82  72 . 22 27 . 87  29 . 07 7 0 . 93  59 . 44 40 . 56  6 0 . 38 39 . 63  24 . 03 75 . 97  48 .80 51 . 20  96. 55 3 . 45  69 . 10 3 0 . 90  101  T a b l e 21. C o r r e l a t i o n s amongst component c o r r e l a t i o n s f o r f i r s t components from PCAs i n T a b l e 2 0 . *. c o r r e l a t i o n s s i g n i f i c a n t p < 0.01. O t h e r s y m b o l s as i n F i g u r e 8.  LEAF IS  ANATOMY  T1 T2 T3 T4 T5 T6  . 599 . 726* .749* ' . 389 . 509  .912* .905* .849* . 863*  . 967* . 754* . 722  . 704 . 78 1 *  .7 19  Ms  T7  .435  .808*  . 827  . 702  .847*  .516  Me  T8  . 709  .689  .829  . 847*  . 550  . 377  . 490 .878 . 323 . 594  . 562 . 884 . 751  . 563 . 877  .831  LEAF Is  .611  MORPHOLOGY  T1 T2 T3 T4 T5 T6  - .671 . 688 . 924* . 790 . 975*  Ms  T7  . 955*  - . 713  . 763  .909  .837  .955*  Me  T8  .938*  - . 774  .693  .941*  . 758  .928*  .991*  T3  T4  T5  T6  T7  T1  -  T2  TWIG MORPHOLOGY Is  T1 T4  . 574  Ms  T7  .519  .519  Me  T8  .438  .438  .438  T1  T4  T7  T8  T8  102  F i g u r e 11. E x a m p l e s of i n t r a - i n d i v i d u a l some t r e e s shown i n F i g u r e 8.  statistical  comparison  These diagrams nature the  of v a r i a t i o n  similarity  (Fig.  and  individuals  indicating  12) f u r t h e r  for  appropriate.  the continuous  the exception  ( F i g . 8: T5, T 6 ) .  emphasize  component  scores  indicated  variation  was t h e l a r g e s t  from  presents  separate  PCAs  of l e a f pattern.  s c a t t e r s of  variation inter-positional  source of v a r i a t i o n Correlations  21) from PCAs  of the v a r i a b l e s  21  of i n t r a - i n d i v i d u a l  and s u b s e q u e n t  that  f o r P^ e n g e l m a n n i i . (Table  Table  of the n e c r o t i c  i n the c o n t r i b u t i o n  to the p a t t e r n  PCAs of i n d i v i d u a l t r e e s  similarity  anatomy  The d i a g r a m s a l s o  o f age w i t h  a striking similarity  correlations  emphasize  among t h e e i g e n v e c t o r - v a l u e s  anatomy v a r i a b l e s  except  leaf  of i n t r a - i n d i v i d u a l v a r i a t i o n of a l l the  regardless  correlations  of  was n o t c o n s i d e r e d  i n the t r e e s .  individuals stunted  of s l o p e s  variation  tree,  amongst component  i n Table  contributing  i n each  20, s u g g e s t  to the pattern  a general of  103  F i g u r e  12.  S c a t t e r s  c o m p o n e n t s a g a i n s t Tab  1e  4 1,  c o m p o n e n t  of  w h o r l  A p p e n d s  of  P C A s  means  f o r  b r a n c h  d r a w n  ix to  a n d  s t a n d a r d  i n d i v i d u a l  p o s i t i o n s . S y m b o I s  III. same  L E A F  s e a  d e v i a t i o n s  b r a n c h e s S c o r e s a s  i n  of  b a s e d F i g u r e  of  s c o r e s  i n d i v i d u a l o n  P C A s  8.  A l l  f i r s t  Ie.  ANATOMY  LEAF  TWIG  of  t r e e s  g i v e n  MORPHOLOGY  MORPHOLOGY  TB  i n  f i r s t  104  intra-individual  variation  correlations  indicated  variation  leaf  of  regardless wider  central  maturity,  Variation  bundle i n P_^_  Unlike  result  of  than  the  the  T h e r e was except  disposed  that  that  for  having  larger  central  with  the  leaf  was  the  of  location  P^  of  the  leaves. engelmanni i  central  cylinder  contributing  and  to  and  the  twig morphology,  between p o s i t i o n s ,  highly  correlated  no  consistent  trend  leaf  anatomy,  that  in  were a l s o  PCA  component to  be  related  Figure  13  p r e s e n t s an  did  not  show  with p o s i t i o n  in  inter-whorl  suggested Of  that  the  twig morphology  performed i n the  scores against  appear  although  the  more  variable showed  the  variation.  significant variation  linearly  as  a p i c a l - and  within  (Table  data,  position  ordination  s e p a r a t e PCAs, and  amongst c o n e s of  the  more b a s a l  individual  variables  was  from t h e s e  was,  (NEEDEP) and  cone m o r p h o l o g y v a r i a t i o n  trees  sitchensis  being  l e a v e s were more v a r i a b l e .  intra-position  individual  F\  leaves  were e x a m i n e d w i t h PCA,  PCAs of  inter-positional  apical  the  anatomy,  trend  variation,  component  the  significant differences  apically  of  The  sitchensis.  leaf  canopy.  of  i n d i v i d a l s of  upon a s p e c t s of  a consistent  least  a  deeper  among l e a v e s  variation  suites  of  pattern  CENCY.^.) t h a n  more d e p e n d e n t  showing  the  (or measurements a s s o c i a t e d  cylinder;  vascular  that  anatomy  (NEEDWID) and  cylinders  was  of  amongst a l l t r e e s .  the  20).  c a s e of  of  first  the  indicates basal-most  leaf  two  a virtual  there  scatters  canopy d i d  in the  of  Although  preliminary i n the  canopy  of  not  anatomy.  components overlap  cone b e a r i n g  branches.  1 05 F i g u r e 13. O r d i n a t i o n s o f f i r s t two c o m p o n e n t s o f PCAs o f individual t r e e s comparing p o s i t i o n s of cones. S c o r e s f r o m PCAs g i v e n i n T a b l e -4 1. A p p e n d i x I I I . G l y p h s as i n F i g u r e 10. Intervening sample p o s i t i o n s o m i t t e d from o r d i n a t i o n to emphasize p o l a r i t y All c o m p o n e n t s d r a w n t o t h e same s c a l e . Glyphs represent individual sea 1es.  (51.69 •..)  Like  t w i g and l e a f  two t r e e s  were n o t s i g n i f i c a n t l y c o r r e l a t e d  inter-individual  Without variation,  variation  including  the  (r=0.345).  of c o r r e l a t i o n  intra-individual  matrices  i n the data  the l a r g e s t  indicated  (Table  22).  source of v a r i a t i o n  between i n d i v i d u a l s .  of v a r i a t i o n  significant  of ANOVA, s u g g e s t i n g  sources of v a r i a t i o n  corroborating  the c o n c l u s i o n  only  that  were not  i n an i n t e r - i n d i v i d u a l c o n t e x t ,  ANOVAs was  that  correlated.  intra-individual variation  the r e s u l t s  intra-individual  were s i g n i f i c a n t l y  that  Eigen-vector  f o r t h e components o f t h e s e p a r a t e PCAs s u g g e s t e d  Interjecting altered  that  to differences  patterns  of t h e  i n the context of  s o u r c e s of s p e c i f i c  a l l evaluation  22) s u g g e s t e d  attributed  *'•>  variation.  t h e r e was s i g n i f i c a n t v a r i a t i o n  values  4  m o r p h o l o g y , cone m o r p h o l o g y v a r i a t i o n  3.5 I n t r a - i n d i v i d u a l  (Table  <&'"  minimally  these highly  thereby  from e a r l i e r a n a l y s e s .  Adding  106  T a b l e 22. M u l t i v a r i a t e apportionment of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due to d i f f e r e n c e s between i n d i v i d u a l t r e e s or t a x a , e m p h a s i z i n g d i f f e r e n t s o u r c e s of i n t r a - 1 n d 1 v 1 d u a 1 v a r i a t i o n i n an i n t e r - i n d i v i d u a 1 c o n t e x t . A b b r e v i a t i o n o f t h e PCAs a n d ANOVAs i n T a b l e 4 2 , A p p e n d i x I I I . WO - w i t h o u t any o t h e r i n t r a - i n d i v i d u a l v a r i a t i o n b e s i d e s i n t r a - i n c r e m e n t ; 1 ° / 2 ° - w i t h p r i m a r y a n d s e c o n d a r y o r d e r s o f b r a n c h i n g a n d b r a n c h e s w i t h i n o r d e r ; POS w i t h i n t e r - p o s i t i o n a l v a r i a t i o n ; %W0 - c h a n g e f r o m %SS WO. Other symbols g i v e n m T a b l e 13. O r d i n a t i o n of r e s u l t a n t component s c o r e s g i v e n i n F i g u r e 14.  %SS VARIABLE LEAF  SUITE  ANATOMY  A  WO 1 °/2° POS  LEAF  TAXA E  (mva)  %WO  A  INDIVIDUAL E  %W0  31 .87 36 .49 30 . 27  68 . 13 63 .51 69 . 27  0 .0 - 4 .62 1 . 14  87 . 72 86 .96 79 . 89  12 . 82 13 .04 20 . 1 1  0 .0 0 .02 7 . 29  MORPHOLOGY  WO  26 .58 1 °/2° 21 .80 23 .00 POS  73 . 42 78 .20 77 .00  0 .0 4 . 78 3 .58  79 .60 75 . 52 77 .09  20 .40 24 .48 22 .91  0 .0 4 .08 2 .51 ,  TWIG MORPHOLOGY  WO  35 . 42 1 °/2° 35 .05 POS 31 .27  64 , 58 64 . 95 68 .73  0..0 0.. 37 4 . 15  93 .68 90. .63 87 .41  6 , 32 9 , 37 12 .59  0 .0 3 .05 6 . 27  CONE  WO POS  19. ,02 17 ..87  80. .98 82 .. 13  0. 0 1 .. 15  67. . 73 63. .50  32 . 27 35 ..50  0. 0 3. 23  1 °/2° POS  31 . 29 31 .. 1 1 28 . 18  68. 71 68 ..89 7 1 82 .  0. 0 0 . 18 -3 . 1 1  87. 0 0 84 .. 37 81 ..46  13. 0 0 15 . 63 18 . 54  0. 0 2 . 63 5 . 54  WO POS  28 . 22 25 . 6 0  7 1 78 . 74 . 40  0. 0 2 . 62  82 . 18 76. 97  17 . 82 23 . 03  0. 0 5 . 18  MORPHOLOGY  x VEGETATIVE  x TOTAL  intra-individual  WO  sources  amount o f v a r i a t i o n expected.  of v a r i a t i o n  between i n d i v i d u a l s .  However, f o r l e a f  hypothesized  taxa  intra-individual  g e n e r a l l y decreased  sources  Such a d e c r e a s e i s  anatomy t h e v a r i a t i o n  increased suggesting of v a r i a t i o n  the  between  that the i n c l u s i o n could alter  of  a taxonomic  hypothesis. T h e s e ANOVAs s u g g e s t less  than  differences  inter-individual for  between i n d i v i d u a l s .  and i n t e r - t a x o n o m i c  cone m o r p h o l o g y Ordinations,  that d i f f e r e n c e s  than  Figure  between t a x a a r e much F u r t h e r , the  differentiation  f o r the o t h e r  variable  was  lower  suites.  14, c o r r o b o r a t e t h e i m p r e s s i o n  of s m a l l  1 07  inter-taxonomic open g l y p h s 18%  of  the  represent total  ordination d e g r e e of  differences. means of  varition  during  Indeed, b e a r i n g  the  individual  i n the  data  calculations  i n mind t h a t  trees  has  been  ( i . e . on removed  of means) t h e r e  o v e r l a p between p o i n t - s w a r m s a s s i g n a b l e  the  average  from  the  is a  large  t o the  two  taxa. The  results  and  summarized  the  three  Compared  to the  being  of  variation.  larger  than  sources  investigate  and,  Consideration  sources  of  the  17 have been  variation  are  s o u r c e s , of  within a  both  to  summarization  hypothesized  of  of  single  specified  these  should  the  and  accounting  development  of  the  affect of  patterns the  averaged  assumes  that  independent.  intra-individual  increment  is  indicated  inter-increment  results  variation  have been  p a t t e r n of  sources  leads the  p o s s i b l e to  these  few  to c o n s i d e r a t i o n s  understanding  inter-individual of  are  investigated.  just  patterns  of  sources concerning  and variation.  intra-individual  assumed e f f i c a c y  for intra-crown  o f crown  these  positions,  variation  of p a t t e r n s  into question  As  size  p a t t e r n s may  consideration calls  intra-individual  perhaps,  intra-individual  interpretation  in  This  14  Discussion. Other  how  other  in Tables  15.  the v a r i a t i o n  as  of  in Figure  hypothesized  variation,  4.  presented  Such  variation  extrinsic  of v a r i a t i o n  processes  and  the  form.  are  b a s e d on  t h e c o n c l u s i o n s must be  only  a  few  regarded  as  t r e e s and being  a  more  few  F i g u r e  14.  O r d i n a t i o n s  v a r i a b l e  s u i t e s  i n t e r - i n d i v i d u a 1 A p p e n d i x g l y p h s  i l l . a r e  v a r i a t i o n  o f  f o r  f i r s t  Open  g l y p h s  i n d i v i d u a l i n d i c a t e d .  two  c o m p o n e n t s  i n t r a - i n d i v i d u a l  v a r i a t i o n .  S c o r e s r e p r e s e n t  s a m p l e s O t h e r  f r o m  g l y p h s  o f  v a r i a t i o n b a s e d  a a s  o n  means s i n g l e i n  PCAs i n  PCAs  o f  o f  f o r  s e p a r a t e  c o n t e x t  i n d i v i d u a l  t r e e  F i g u r e  t h e  g i v e n t h e  i n  o f  T a b l e  t r e e s ; s o u r c e  4 2 . s o l i d o f  10.  LEAF MORPHOLOGY H VI2'  MLB  o  £  O CP 0  OA& AA  tt int«r-positional  ,o  ,  oS?»*  O  OO  •O O  A A  A A  TWIG MORPHOLOGY VIV  inter-post tional  O  LEAF  ANATOMY  fii.3f,.l  CONE  MORPHOLOGY  I49.76-'.)  It3.se-.)  109  F i g u r e 15. Summary o f p a r t i o n i n g o f s o u r c e s o f i n t r a - i n d i v i d u a l variation. Summary b a s e d on t o t a l '/SS (mva) T a b l e s 14 t o 17 S t i p l e d border of p i e i n d i c a t e s r e g i o n that i s r e f e r r e d t o i n a l l subsequent a n a l y s e s as " i n t r a - i n d i v i d u a l v a r i a t i o n " .  tentative suggest  than  conclusive  t h e need  research  forestry  to understand  ecological  interests  physiological phenology  f o r new r e s e a r c h  in seeking  developmental,  i n nature.  and y i e l d  (Dietrichson  and  Campbell Cannell,  directions  and e v o l u t i o n a r y  variables 1964),  concerned  investigations  As  with  1975), p a r t i c u l a r l y of the  these v a r i a b l e s  and L e s t e r  e t a l . 1976) s h o u l d  results  and a p p r o a c h e s t o  processes.  (Campbell  underlying  1974; R e h f e l d t  these  the i n t e g r a t i o n of  become i n c r e a s i n g l y  developmental assumptions 1965;  Regardless,  (Burley  1969; R e h f e l d t  be u n d e r t a k e n .  1983;  110  4.1 S a m p l i n g  These  implications.  results  suggest  that,  i n Picea,  provided that the  sampling  of v e g e t a t i v e m a t e r i a l s i s r e s t r i c t e d  branches  of i n d i v i d u a l s ,  difference sampled.  t o t h e lower  t h e r e s h o u l d n o t be a g r e a t d e a l o f  a s t o whether a w h o r l  or a d v e n t i t i o u s  branch i s  F u r t h e r , t h e o r d e r o f b r a n c h i n g and y e a r  may be u n i m p o r t a n t  to understanding  of c o l l e c t i o n  inter-individual  However, a s t h e s e e s t i m a t e s a r e sample and v a r i a b l e seems a p p r o p r i a t e t o s u g g e s t increasing  that  t h e p r e s e n t sample  observations previously  extending  u n r e p o r t e d and s i g n i f i c a n t  intra-individual developmental  variation.  differences  take  into  results  suggest  variation  inter-individual  disposition branches in  variation  when e x a m i n i n g  and d e r i v i n g  correlated  when making  with  variation resin  cyst  o f samples w i t h  s h o u l d be t a k e n  inter-individual  these  based  lengths  variation  First,  where p a t t e r n s o f  morphology a r e s t r o n g l y  (RESCYLEN), t h e n t h e or a d v e n t i t i o u s  consideration. based  explainable  by t h e o r d e r o f b r a n c h  collections  have been made o v e r  into  inter-individual  r e s p e c t to whorl  into  i n which  s h o u l d be t a k e n  on l e a f  appear  f e a t u r e s and t o  situations  p a t t e r n s of  inferences.  i n t h e canopy  interpretations.  three s p e c i f i c  s o u r c e s of i n t r a - i n d i v i d u a l consideration  e n v i r o n m e n t a l and  between p o s i t i o n s  i t would seem a d v i s a b l e t o n o t e  The  uncover  s o u r c e s a n d p a t t e r n s of  Where o b v i o u s  consideration  critical  form may w e l l  to e x i s t , them  specific, i t  s a m p l i n g and  s i z e s and making  on o t h e r a s p e c t s o f crown  variation.  Second,  anomalies  on t w i g m o r p h o l o g y might sampled.  several  Third,  years,  be  where  inter-individual  111  variation year  of  of  created  by  The  for of  b a s e d on  cone w i d t h  mature  the  Similarly,  cone m o r p h o l o g y  year  l a c k of  of  ages and  be  and  The  species  observations  t o p u b e s c e n c e and indicate  do  differ  of  Picea  here,  variables  examined here,  i n an  not  variation 1948;  Funsch respect  presented  especially  variation  here  large.  - mature  With respect  a trivial  with  samples.  II w i t h  inter-individual  a p p e a r s t o be  a l l ages  of  immature  analyses  in i n t r a - i n d i v i d u a l immature.  that,  remains unclear  i n Chapter the  and  suggests  (Lindquist  d i f f e r e n c e s are  more v a r i a b l e t h a n  variation  juvenile  of a g e - r e l a t e d  reported  t r e e s are  age-related  artifacts  sampled  t o a l a c k of  needle morphology,  that age-related  However t h e y  for  21)  situation  observations  in other  the  examined  conditions  t o P_;_ e n g e l m a n n i i owing  demonstrated  strongly correlated  u s e d when e x a m i n i n g p a t t e r n s  variation.  the  the  inter-individual  are  s i t c h e n s i s (Table  environmental  withstanding  1975)  be  of  d i s c r e t e d i f f e r e n c e s between  individuals could  respect  that  d i f f e r e n c e s in  collections.  i n d i v i d u a l s of  the  reflect  patterns  (CONWID) s h o u l d  inter-individual  Not  morphology c o u l d  collection.  variation with  twig  to  the  context,  source  of  var i a t i o n .  The  demonstrated  intra-individual  l a c k of  variation  of  assumption  (Chapt. II) that  collection  from the  suggest be  of  that  crown  disposition  s e r i o u s consequence  identifiable  cone morphology  cone c o l l e c t i o n s  i s not of  pattern  cones  necessary. i n the  in understanding  of  suggests that  the  represent  a  The  presented  data  tree w i l l patterns  random  probably of  not  11 2  inter-individual  variation.  On an i n t e r - i n d i v i d u a l used  in this  probably  requires  data  exceed  That  intra-individual  inter-individual  supposedly  represent  question  are separated  Further,  inter-specific  variation  raised  respect  here with  intra-individual consideration variation. the  spread  (Table  should  This  opinion  of samples  that  22) s u g g e s t i n g  that  the p o i n t s  and d e g r e e o f  be t a k e n  individual  these  than  of  into  careful  inter-individual  i s f u r t h e r r e - i n f o r c e d upon  from  (Table  The i n d i v i d u a l s i n  i s much l e s s  when e x p l o r i n g p a t t e r n s  of v a r i a t i o n  a n d 1400m e l e v a t i o n .  t o the p a t t e r n  variation  taxa  of v a r i a t i o n  two t a x a .  variation  inter-individual  o f t h e two  considering  by 6° l o n g i t u d e  the t r e e s  and a n a t o m i c a l l y  sources  sources  result  that  o f t h e two t a x a , a n d  t h e most m o r p h o l o g i c a l l y  i s a somewhat u n e x p e c t e d  individuals  standards  a v a i l a b l e , the d i s c r e t e n e s s  comment.  generally 17)  a n a l y s i s represent  represent  polarized  note, c o n s i d e r i n g  trees  i n these  considering ordinations  ( F i g . 14).  These r e s u l t s the  (Barnosky  P. e n g e l m a n n i i  1981).  could  re-examined  reported  be o f n e e d l e s  than  needles  of P i c e a m a c r o - f o s s i l s  also citations  these  of IP^ e n g e l m a n n i i  This  simply  Pj_ s i t c h e n s i s r a t h e r  findings see  the p a l e o b o t a n i c a l  f i n d i n g s of m a c r o f o s s i l s  lowlands  of  question  in light  of these  same r e m a r k s w i t h  f r o m t h e upper  o f P_^ e n g e l m a n n i i .  canopy Other  and O g i l v i e 1970;  1984) s h o u l d  findings.  respect  i n t h e Puget  f i n d i n g of  (e.g. H i l l s  in Critchfield  s i g n i f i c a n c e of  be c a r e f u l l y  Wilson  to m i c r o f o s s i l s .  (1963) e c h o e s  11 3  These of  other  Klinka, that  results  on  researchers et  al.  (Garman  1982).  i n t e r - s p e c i f i c and  to  the  incorporation  variation  into  variation. taxa  based  variables  on  1982;  1959)  should  be  considerations  positions  of  slight  as  of  living  could  are  to  understanding  Roche  to  of  1969;  of  the  relative  lends  sources  to  support  between  vegetative  conclusion  of  previous  cone m o r p h o l o g y  (Coupe,  1984;  These c o n c l u s i o n s into  size  of  differences  Critchfield  integrated  suggest  intra-individual  compared the  22  1968;  as  variation  smaller  et  Horton  and  subsequent  inter-individual  the  the  support  variation.  t h o s e of  significance canopy.  strongly  related  shown h e r e ,  If to  highly  an  the  physiological  patterns  even  tree  s i g n i f i c a n t and  and  physiological  then,  individual  previous  between o r d e r s  h e r e , when e x t r a p o l a t e d  canopy of be  here  i n the  variation  reported  prove  of  a s p e c t s of  presented  branches  morphological  large  about  those  Daubenmire  in Table  trans-individual  patterns  respect  of  as  contradict  morphogenesis.  characteristics  the  1978;  results with  on  have been  Crown f o r m  reports  on  re-considered.  interpretations  The  information  suggests that  primarily  1969;  observation  cone m o r p h o l o g y as  Strong  be  intra-individual of  22)  Roche  may  This  demonstration  (Table  work r e l y i n g  variation  research  The  1957;  values presented  variation.  of  4.2  The  intra-individual  inter-specific  al.  inter-specific variation  for  i f they are the  these  entire  as volume  differences  especially  m a i n t e n a n c e of  of  relevant  individual  in  trees.  11 4  The be  morphological important  fungal  and  in understanding  variable  apical  selective  insect  f o r the  and  to growth r e g u l a t i o n a s s o c i a t e d  The  d e g r e e of v a r i a t i o n  between b r a n c h e s w i t h i n  environment  and  manifestation  owing  the  factors.  t o the that  shade.  The  by  sampling  of p a t t e r n s  12 and of  environments. controls  are  intra-crown  result  The  observation  and  basal-most  of c l i n a l  the  pattern  this  regard  appears today  i n the  described and  intra-canopy  may  the sun  regarded  as  opposite  poles  a  Table  i n d i c a t e a high  20  individaul  Such an regarded  variation.  of  not  leaves will,  be  caused  in Picea,  as  being Further  the  and  artifact  d e g r e e of  over a  to tree  uniformity  similarity  small  but  different  i s unexpected  important  be  continuum.  i n markedly  observation  by  out  eventually,  a sampling  variation  trees  of branches  d i s c r e t e d i f f e r e n c e s between a p i c a l be  from  i t i s worth p o i n t i n g of  with  surrounding  However, b o t h m a t u r e  that  intra-individual  g r o u p of  could  environmental v a r i a t i o n  perennial duration  b r a n c h e s must  Figure  diverse  In  canopy  a t t r i b u t e d to  shading,  apical-  same p a t t e r n  suggesting  leaf  basal-most caused  show the  thus  environmental  the  of  shade" morphology.  trees  variation,  order  (i.e. direction,  d i f f e r e n c e s between t h e  immature  an  apical-most  i n s e c t or p h y s i c a l damage).  suggests the "sun  larger  ascribed high  b r a n c h e s and  of  be  dominance.  discrete  observed consistency  could  vegetation,  in  of  primary  local  that  nature  also  on  differences  that,  the  shown h e r e may  values  positions  as  variation  infestation.  Explanations  the  anatomical  of  i f extrinsic form the  and pattern  of  11 5  intra-individual interesting pattern tree.  t o note  appears These  saplings  physiological  and  suqgest  as o p p o s e d  that  may be t h e  to environmental  g r o w t h and d e v e l o p m e n t  of developmental  control  of  as i n  of i n t r a - c r o w n  directly  processes associated  to considering  with  growth  f o r t h e c o n t i n u e d c o - o r d i n a t e d and  p h y s i o l o g y , and development  complex  organisms  p r o c e s s e s may  o f young t r e e s ,  development  uniformity in  s i t c h e n s i s and  Such s u g g e s t i o n s l e a d  growth,  architecturally  development  in  a l t h o u g h n o t t o t h e same d e g r e e  can account  predictable  that,  of d e v e l o p m e n t a l  known p h y s i o l o g i c a l  regulation  of t h i s  It i s  by t h e v i g o u r o f t h e i n d i v i d u a l  t h e same s o r t  differentiation. how  controls.  form and i n t r a - c r o w n v a r i a t i o n  results  i s under  similar  the maintenance  suggest  crown  These  mature t r e e s  suggests  t o be a f f e c t e d  primarily,  processes.  that  results  P• e n q e l m a n n i i , result,  variation  in trees  as P i c e a .  be a d e q u a t e however  i n such  The known  i n addressing the  the c o - o r d i n a t i o n  r e a c h i n g over  large,  30m  i n height  of growth remains  enigmatic.  In  s u p p o r t of the a f o r e m e n t i o n e d  integration results  dealing  similarity branches Ogilvie  of developmental  based  and e v o l u t i o n a r y  with the assessment upon whether  a r e examined.  necessity  of  samples  Similar  f o r an  studies,  are the  inter-individual from a p i c a l  or b a s a l  o b s e r v a t i o n s have been made by  and von R u d l o f f (1968) c o n c e r n i n g t h e s i m i l a r i t y o f  intra-crown  variation  variation.  Stover  variation variation.  o f P_;_ e n g e l m a n n i i  (1944) c i t e s  o f E"\ e n q e l a m n n i i The same a p p l i e s  that  to elevational  a similarity corresponds  of i n t r a - c r o w n to edaphic  f o r t h e o b s e r v a t i o n made h e r e a n d  11 6  reported  elsewhere  by  Daubenmire  t h e a p p e a r a n c e of p u b e s c e n t individuals pubescent P.  w e l l removed  rubens).  expressed  These  then  of  the  Further  research.  Having  presented  intra-individual  some d i r e c t i o n  through  inter-taxonomic  of and  form  Jarvis  Riding  context.  during  that  reported for  i t seems a p p r o p r i a t e  to  inter-relations  i n an  of  research directed of  these  g r o w t h of  these  sources  and  and  such  T h i s same s u g g e s t i o n  has  been made by  example w i t h p r a c t i c a l  of  of  (1982) s u g g e s t s  respect to p h y s i o l o g i c a l  at  individual  inter-individual  Tomlinson  sources  form  programmes.  growth  the  the  (1974) w i t h  being  an  for understanding  (1976) p r o v i d e s an  grafting  of  stages  with  Specifically:  sampling  variation  in trees.  or  s k e t c h of p a t t e r n s of  t h e d e v e l o p m e n t and  is essential  mariana,  for future research d e a l i n g with  variation  intra-individual  P.  I t i s worth n o t i n g  in Picea  variation.  systematic  other  derived  development  r e p o r t e d here  same  with  t h a t , i n s t e a d of  at p a r t i c u l a r  a thumbnail  t h e d y n a m i c s and  and,  and  the  to  (1948).  exploring  approach  taxa  respect  b r a n c h e s on  shared  g r o w t h and  variation  intra-individual  trees;  at  between  situation  Lindquist  with  p o s s i b l e sympatry  hint  are only manifest  a b i e s by  suggest  the  glabrous  only at given p o s i t i o n s .  similarity  4.3  differences  1974)  ( e . g . P_;_ e n g e l m a n n i i ,  results  throughout  individual,  P.  from  s p e c i e s of P i c e a  developmental  and  and  (1968,  an  variability Norman  variation. application  in  1 17  Dealing variation, recently (Maze, been  one  e t a_l.  portions  1984).  here,  directed  with  individual  respect trees.  (1982) and o t h e r s , al.  1972;  static McGuire of  might  be  process. sampling  important  of crown  of F r a s e r 1976).  variation regard  w i t h t h e age o f would  be  research  form v a r i a t i o n  1976;  1981;  Nicholls  (Fraser,  of the canopy  et_ a l .  the  and would by  provide  Maillette  Namkoong, e t 1967)  and t h e  1964;  Fraser  changing proportion  of  and  sources  o v e r t h e c o u r s e of d e v e l o p m e n t  to understanding  S u c h a s t u d y would  age,  would a d d r e s s d i r e c t l y  and Wykoff  The  variation  with c a r e f u l  portions  ( F i g . 10) where  of t h e d y n a m i c s d e s c r i b e d  Fraser  intra-individual  than i n o l d e r  t o g r o w t h and d e v e l o p m e n t  Namkoong and C o n k l e  1969;  f o r more  t h e i n t e r - d e p e n d e n c e of p o s i t i o n ,  (Rehfeldt  descriptions  variability  anatomy  in this  Such a s t u d y  extension  intra-individual  h y p o t h e s i s a p p e a r s t o have  for leaf  important  g r o w t h and d e v e l o p m e n t a valuable  this  intra-individual  at u n r a v e l l i n g  and o r d e r  increased  In f a c t ,  Especially  of  o f t h e canopy  at least  is increasing  the t r e e .  of  with the dynamics  would p r e d i c t  derived  upheld  there  first  the c o n t r o l s  require  whole  tree,  of  this  systematic  mapping o f s a m p l i n g l o c a t i o n s ,  ages,  and  orders.  Extending different derive  a study,  populations  inferences  evolutionary s t u d y would addition  outlined  o f , say, s e v e r a l  concerning  processes. be  such as t h a t  the i n t e g r a t i o n  Additionally,  in intra-individual  t o t h o s e measured  taxa,  as  above, could  h e r e m i g h t a l s o be  several  be u s e d t o  of d e v e l o p m e n t i n  interest  variation,  to  i n such a  variables in considered.  1 18  Another discussed issue  above,  of r e s e a r c h , i s the matter  of a g e - r e l a t e d  concerning  variation  the p h y s i o l o g y  resultant in  area  selection.  several species  differentiation  that  and  s e l e c t i o n may  i n these  1983).  anomalous r e s u l t s  age-related  variation  age c o u l d  between g e n e r a t i o n s .  rather  than  of p a t t e r n s regarding  may p r o v e u s e f u l  of v a r i a t i o n ,  the morphogenetic  Such a s t u d y  degree t h a t cumulative and  effects  examine t h e c o n f o r m i t y  of these  such a  i n c o n i f e r s and form, a  the perceptions  i t would p r o v i d e f o r these  information patterns.  i n understanding  are responsible  study  of development  In P i c e a  i n understanding  important  as i n o t h e r  and Brown  a n a l y s i s o f crown  explanation  may be e s p e c i a l l y  known i n  sequential  The d e v e l o p m e n t  as w e l l  The  morphogenetic  the course  of the organism.  have a l l been b a s e d on s t a t i c study  i s well  would have t o i n v o l v e  Picea  variation  Sarg.) as w e l l  1979) and a t r u l y  would r e q u i r e many y e a r s .  pointed  differentiation  1981; Zimmerman  study  be  separated.  variation  scopulorum  maturity  (Maze, e t  a consequence of  o f t h e same i n d i v i d u a l s o v e r  through t o sexual  dynamic  of the  s t u d i e s of i n c r e a s i n g  genetic  ( W a r e i n g and P h i l l i p s  variation  addressed  of a g e - r e l a t e d  of these  Age-related  (e.g. Juniperus  of a g e - r e l a t e d  h a s been  However, i t s h o u l d  be s i m p l y  Kramer and K o z l o w s k i  sampling  i n the context  n o t have been a d e q u a t e l y  with  The  i s c e n t r a l t o t h e i n f e r e n c e s made  studies the e f f e c t  variability  1971;  variation.  o f j u v e n i l e from mature p o p u l a t i o n s  out  woody p l a n t s  of a g e - r e l a t e d  to that  o f e s t a b l i s h i n g s e e d l i n g s and any  of A b i e s  1981; Maze and P a r k e r  Juniperus  allied  Age r e l a t e d v a r i a t i o n  al.  apparently  partially  f o r crown  d e s c r i p t i o n s with  the form  respect to  1 19  hypothesized Fisher  models  and Honda  variations  (Honda  1971;  McMahon and  1979a; B a k e r , e t al.  (Fisher  and H i b b s  1982;  1973)  Hibbs  Kroneuer and  1981).  1976;  reported  1 20  IV.  1.  INTER-INDIVIDUAL VARIATION: TAXONOMIC  Introduct ion.  As exhibit  P_;_ e n q e l m a n n i i natural  layering  a n d P^ s i t c h e n s i s  cloning  (however  i n P^ s i t c h e n s i s  individuals  identical  correspond  we a r e i n t e r e s t e d  hypotheses:  a n d P_^ s i t c h e n s i s.  sources of v a r i a t i o n  of these hypotheses  the a n a l y t i c  there are  levels  of v a r i a t i o n  populational,  or  that  taxonomic  rather  than  on t h e d a t a .  and Dearn  emergent,  f o r and d e t e c t i n g  emergence o f a t r a n s - i n d i v i d u a l  a r e p r o v i d e d by C a m p b e l l  a  priori  Good examples o f source  of v a r i a t i o n  (1980) and W e l l s ,  a l . (1977). If  can  unique.  By a n a l y t i c a l l y  i s h e r e made t o e x a m i n i n g  hypothesized imposition  genecological,  as w e l l as  g e n e r a t i o n s , e c o t y p e s , p o p u l a t i o n s , and s p e c i e s -  P. e n g e l m a n n i i reference  i n whether  between  Except f o r  are genetically  emergent, t r a n s - i n d i v i d u a l  to a priori  differences  differences.  twins, a l l i n d i v i d u a l s  analytically  (1931) r e g a r d i n g  of g e n e t i c d i f f e r e n c e s  and e n v i r o n m e n t a l  Specifically,  do n o t g e n e r a l l y  s e e Cooper  ), p h e n o t y p i c  are a function  developmental  et  CIRCUMSCRIPTION.  t h e emergence o f t r a n s - i n d i v i d u a l  be d e m o n s t r a t e d ,  subsequent  explanation.  hypothesized  they  r e q u i r e q u a n t i f i c a t i o n and  Quantification  s o u r c e s of v a r i a t i o n  of  these hypotheses  of  variation.  variation  then  sources of v a r i a t i o n  and e x p l a n a t i o n f o r  without  examining  the v a l i d i t y  may b i a s c o n c l u s i o n s r e g a r d i n g t h e s e  Indeed,  levels  t e n d e r i n g e x p l a n a t i o n s f o r sources of  more c o n c e p t u a l t h a n  real  may l e a d  to  unwarranted  121  reification  and  (1965a) and  Langlett  (i.e.  misrepresentation  a taxonomic  importance  on  hypothesis)  confers  a named t a x o n o m i c imparls  heterogeneity  between g r o u p s ,  variation of  of  the  trans-individual  chapter two  the  an  as  the  For  acceptance  of  regarding  between g r o u p s of  that  such as  explanations  b a s e d on  or  of  the  The  and  variability  developmental, and  scales  i n t e n t of  taxonomic  this  hypothesis  s u c h a taxonomy w i l l  determine  of  and  explanation  l e a d to f o r the  for  population  s t r u c t u r e of  erroneous  observed  variation  Further,  in a  are  not  P_;_ s i t c h e n s i s ,  the  tendering  part  of  quantification  and  of be  explanations  i n a p o o r l y d e f i n e d or polymorphic  subjected  lead  to erroneous c o n c l u s i o n s variation is also  to a r b i t r a r y about  nomenclatural that  in geographically considered.  taxon  species  identifiable  p o p u l a t i o n a l a s s u m p t i o n s would  has  "taxa"  and  which p a t t e r n s  individuals.  that  of p o p u l a t i o n  of  i s more  individuals  Similarly,  variation  been  there  lacks genecological  sympatric  populations,  population  sense  subspecific levels  explanation  where a l a r g e number of  inappropriate.  t h a t a name  (hypothesis)  compared.  s e l e c t e d g e n o t y p e ( s ) may  conclusions  false  Burley  variation.  i n a taxon  prevalent,  sympatric  are  example, q u a n t i f i c a t i o n  variation  that  validity  the  out  Intra-individual,  variation  c i r c u m s c r i p t i o n of  inter-individual  or  a  reality.  of h o m o g e n e i t y w i t h i n  scale against  i s t o examine t h e taxa,  group  impression  w i t h i n groups.  provides  biological  (1959,1962,1963) p o i n t  automatically  between than  of  for  taxon  splitting  unless  juxtaposed  the and  may  nature  1 22  Such e x a m p l e s a r e taxa  being  i n v e s t i g a t e d here.  latitudinal  trend  Lines  may  1976)  perceived along  serve  a pronounced latitudinal  latitudinal variation  "populations"  the  are  1953  1977), may suggest  1  ; Daubenmire  serve  discontinuities  elevation  (Habeck and  1959;  If  of  two  results  the  two  Hulten between 1  Garman  hypothesized  1957;  taxa  sources  i n Chapter  of h y b r i d s may are  smaller  be  than  and  P^  difficult  as  glauca Beckwith  the  related von  or  to  Rudloff  1968;  1968).  n e e d s t o be 22)  perhaps  in variation  corresponding  e m e r g e n t , then  (Table  -  of  Additionally,  comparison  Dugle  of  1976b).  Copes and  of v a r i a t i o n  indeed  III  By  be  Such  a number  with  trend  Ogilvie  and  should  the a c t u a l n a t u r e  g e n e r a l l y been  1969;  La R o i  taxa are  by  trend.  1969;  in v a r i a t i o n .  Weaver  and  The  from e x p e c t a t i o n s  expected  natural hybridization  presented  recognition  the  (Lewis  1976).  chapters,  Roche  P_^_ e n g e l m a n n i i has  trans-individual  potential  (Wright  P_j_ s i t c h e n s i s  1968;  to obscure  of  Horton  of  pronounced  that there  1978b; O ' D r i s c o l l  in previous  two  r a n g e of P_j_ s i t c h e n s i s  l a c k of a p r o n o u n c e d  variation  the  Sziklai  l a c k of a  in point.  suggests  somewhat d i f f e r e n t  f o r the  case  geographic range  f o r the  sitchensis  been d e m o n s t r a t e d  reported hybridization  (Little  of  variation  has  and  However, as m e n t i o n e d  accounting  and  to envisage  reported  as a c o n v e n i e n t  latitudinal  (see Ching  easy  The  in populations  narrow e d a p h i c  a lengthy  workers  especially  the  question  addressed.  indicated  to  that  The the  d i f f e r e n c e s between  d i f f e r e n c e s between t r e e s .  (1968) c o n s i d e r s t h i s r e p o r t t o be b a s e d on a h y b r i d s i t c h e n s i s and P_^_ m a r i a n a r a t h e r than P^ glauca.  1 23  Similarly, of  variation  perceived  of  the  are  not  Further,  variation  as  arbitrarily  and  vital and  the  being area  variation  nature  of  of  such  prescriptions  and  are  levels  sources the  into i n which  owing t o the  acceptance two  taxa.  such t r a n s - i n d i v i d u a l  populations,  d i s c r e t e taxa,  in themselves,  levels and  offer  i n c o n s i s t a n c y between e x p e c t e d and  provenances are  individuals  without  to the  or  the  reference actual  sources  of  from  edaphic  phylogeny. variation  further detailed  may  a  viewpoint,  suggest  objectives in s i l v i c u l t u r a l  provides  quantification  From a more p r a c t i c a l of v a r i a t i o n  and  simply  r e p r e s e n t a t i v e of  these  explanation.  documentation  called  i n d i c a t e areas  populations  c i r c u m s c r i p t i o n f o r the  subsequent  be  of d i s c r e t e n e s s of  there  reported  geographic  Establishing  may  have been d e f i c i e n t  Often  microgeographic  taxa  t a x a may,  e r e c t e d as  trans-individual  emergent manner, t h e n  f i n d i n g s may  generations,  f o r the  a particular  these  whether  variation.  i n an  perception  between t h e s e  explanations actual  such  questionable  hybrids  manifest  c o n c l u s i o n s may  Establishing of  hypothesized  d i s c r e t e n e s s of  question. previous  i f these  and  alternate  tree-breeding  programmes.  2.  M a t e r i a l s and  2.1  Materials. All  of  the  methods.  t r e e s sampled were u s e d  trans-individual  sources  of  from t h e  sitchensis  of v a r i a t i o n  in examining  (this  i n c l u d e d the  Chilliwack River Nursery).  samples  Both  1 24  m a t u r e and  immature  (Table  19)  indicated  ages.  In p r o v i d i n g f u r t h e r  individuals assess  t r e e s were u s e d little  difference  parent  2.2  primary  analytic  form  Barrow  and  1972)  analytic  were i n c l u d e d i n some a n a l y s e s .  of  technqiue  the h y p o t h e s i s  the a v a i l a b l e  a p p r o a c h a l l o w s any  variation  t o be  Including  intra-individual  emergent  relative (i.e.  to v a r i a t i o n  developmental  would be  sources. taxon  emergent  samples of t h e i r  that  sources  The  i s dependent  inclusion  of  upon  PCA  maternal  size  (Table  i n these  source  t o be  able  are  intra-individual  indeed  variation  available  s i z e s m i t i g a t e a g a i n s t such  10).  t o be in  assessed nature  emergent,  to a  variation.  would be  (Table  allows  individuals  individual  intra-individual  of  Pragmatically, i f  to a s s i g n  of a g i v e n  and This  analyses  i s more i n t r a - i n d i v i d u a l genetic).  the  priori.  of v a r i a t i o n  accommodated and sample  10).  sources  imposed a  variation  respects  tested (Burley  trans-individual  of v a r i a t i o n  assignment  PCA.  being  sample  r a t h e r than  worthwhile  was  r a t h e r than  trans-individual  trans-individual  the  of  compared.  unstructured  it  To  Analyses. The  any  comparison,  t o r e c o g n i z e n a t u r a l h y b r i d s , samples  a r t i f i c i a l h y b r i d s and  were a l s o  different  i n c l u d e d i n some a n a l y s e s .  known a r t i f i c a l h y b r i d i z a t i o n Additionally,  analyses  between t h e  inter-taxonomic  of P_^ g l a u c a were  the a b i l i t y  as p r e v i o u s  could  to  these  hypothesized Although  be  a p p r o p r i a t e f o r a s t r u c t u r e d MVA, an  then  approach  the  1 25  In q u a n t i f y i n g t r a n s - i n d i v i d u a l populations  and t a x a ,  suites  of p o p u l a t i o n s  nested  design:  levels  of v a r i a t i o n of  ANOVA o f PCAs b a s e d on s e p a r a t e of standards  were p e r f o r m e d  variable  t h a t used the  (MODEL  2.)  y = A + B(A) + C(AB) + e.  where A i s an e f f e c t  b a s e d on d i f f e r e n t population  taxa,  attributed  to particular  individual  i n B, a n d e i s i n t r a - i n d i v i d u a l  B i s the e f f e c t  i n A, C i s t h e e f f e c t variation.  This  ANOVA was u s e d  i n e v a l u a t i n g PCAs o f E\ e n g e l m a n n i i a n d  P. s i t c h e n s i s ,  a s w e l l a s i n e v a l u a t i n g PCAs t h a t  populations for  hypothesized  than be  of standards  a test  populations  intra-increment comparatively  populations  serves  on t h e d i f f e r e n t i a t i o n  remembered t h a t  Using  o f P^ g l a u c a .  intra-individual  variation,  and t h a t  included  The i n c l u s i o n  a comparative among  o f a term  purpose  populations.  variation  refers to  some o f t h e p o p u l a t i o n s a r e  small.  just  those  standard  may u n d u l y  samples  that  p o l a r i z e the data  occurred i n  l e a d i n g t o an  assessment  o f r e l a t i o n s h i p s o f t h e two t a x a .  Additionally,  i t ignores  a major  a l a r g e number  populations  (Table  representative  assessment  source  of v a r i a t i o n  of t r e e s a r e not p r e s e n t 7).  Increasing  o f t h e two t a x a  circumscription environmental  rather  I t should  inappropriate  as  o f an  by a p o p u l a t i o n  and geographic  i n the data  in recognizeable  t h e number o f s a m p l e s  without  b e n e f i t of the  or r e s t r i c t i o n  t o extreme  l o c a t i o n s may a l l o w  a less  o f t h e d i f f e r e n c e s between t h e two t a x a .  biased  1 26  A nested  ANOVA: (MODEL 3.) Y = A + B(A) + e.  applied  t o PCAs o f s u c h d a t a  on  populations  just  bias  of standards  c a u s e d by d e a l i n g w i t h  environmental attributed  extremes.  to d i f f e r e n t  variation.  As w i t h  hybrids,  ANOVAs  i n d i c a t e the degree of and  ANOVA, A i s t h e e f f e c t  B the e f f e c t  of d i f f e r e n t  and e i s t h e i n t r a - i n d i v i d u a l  populations  f o r P_j_ e n q e l m a n n i i ,  to addressing  to the previous  recognizable morphological  taxa,  i n the taxa,  Prior  will  In t h i s  individuals  performed  and compared  of s t a n d a r d s ,  P. s i t c h e n s i s ,  the question  this  a n a l y s i s was  and P^ g l a u c a .  of n a t u r a l l y o c c u r r i n g  an e v a l u a t i o n o f t h e m o r p h o l o g y o f known h y b r i d s  context  of standards  whether  t h e d e t e c t i o n o f n a t u r a l h y b r i d i z a t i o n was p r a c t i c a l .  This  of both  was p e r f o r m e d  t o determine  e v a l u a t i o n was made u s i n g an o r d i n a t i o n from PCAs i n c l u d i n g  populations along  with  comparison  of standards  o f P^ e n g e l m a n n i i a n d P_^_ s i t c h e n s i s  samples o f known a r t i f i c i a l of a r t i f i c i a l  performed  to determine  reference  to parent  for  taxa  i n the  hybrids  A further  a n d t h e p u t a t i v e h y b r i d s was  the r e l a t i o n  taxa.  hybrids.  among h y b r i d s  These comparisons c o u l d  without be made o n l y  the v e g e t a t i v e v a r i a b l e s u i t e s as the a r t i f i c i a l  were s t i l l  immature and l a c k e d c o n e s .  between a r t i f i c i a l given  and p u t a t i v e h y b r i d s ,  i n MODEL 3, e x c e p t  different  In t h i s  comparison  t h e ANOVA u s e d was t h a t  t h a t A was t h e e f f e c t  groups of h y b r i d s : a r t i f i c i a l  hybrids  attributed  or p u t a t i v e .  t o the  1 27  PCAs o f s e p a r a t e  variable suites  f o r a l l t r e e s were a l s o  performed.  As w e l l a PCA b a s e d on a l l 36 v a r i a b l e s was a l s o  performed.  F o r t h e a n a l y s i s t h a t used  all  the a r t i f i c i a l  21 v e g e t a t i v e v a r i a b l e s were u s e d .  variables  or a l l v e g e t a t i v e v a r i a b l e s ,  each t r e e , averaging  thus  these  was n e c e s s a r y  The sample  size  i n Chapter  hypothesized  II.  PCAs o f a l l  were a v e r a g e d f o r  variation.  owing t o t h e l a c k o f an  Such  intra-individual  between v a r i a b l e s o f s e p a r a t e v a r i a b l e was c o n s i d e r e d  PCAs on t h e b a s i s o f t h e sample  outlined  values  ignoring intra-individual  one-to-one correspondance suites.  For those  hybrids,  adequate  size  estimation  ANOVAs b a s e d on t h e s e  term a t t r i b u t e d  3 subsumed by t h e r e s i d u a l  t o perform  to individuals  term as a r e s u l t  technique  analyses  had the  i n MODEL 2 a n d MODEL of a v e r a g i n g per  tree.  3.  Results.  3.1  Populations  3.1.1  P_^ e n g e l m a n n i i and P^ s i t c h e n s i s . Polarizing  P.  sitchensis  that two  of s t a n d a r d s .  the r e l a t i o n  by examing o n l y p o p u l a t i o n s  t h e r e were s i g n i f i c a n t , taxa  slight suites,  between P_^ e n g e l m a n n i i and  (Table 23).  Figure  d i f f e r e n c e between  morphology,  variation.  16 c o r r o b o r a t e s  t h e two t a x a .  than  It  be n o t e d  should  inter-individual (Table  this  23),  variation  exceeds the o f cone  accounts  variation that  impression of  For a l l v a r i a b l e  With the exception  inter-population variation  variation  indicated  b u t s m a l l d i f f e r e n c e s between t h e  t h e amount o f i n t r a - t a x o n o m i c  inter-taxonomic  of standards  within  the largest  f o r less populations. amount o f  1 28  Table 23. M u l t i v a r i a t e apportionment of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between t a x a , p o p u l a t i o n s , and i n d i v i d u a l s t a n d a r d s o f P_^ e n q e l m a n n i i a n d P_^ s i t c h e n s i s . A b b r e v i a t i o n o f PCAs a n d ANOVAs i n T a b l e 4 3 , A p p e n d i x I I I . Symbols g i v e n i n MODEL 2. O r d i n a t i o n of r e s u l t a n t component s c o r e s g i v e n i n F i g u r e 16.  VARIABLE LEAF LEAF TWIG CONE TOTAL x  ANATOMY MORPHOLOGY MORPHOLOGY MORPHOLOGY  VEGETATIVE  intra-individual leaf  morphology  taxa.  A  %SS B (A )  (mva) C( AB)  29.21 29 .09 41.81 20. 79 31 .54  24.49 16 . 16 16 . 74 27 .70 27 . 14  33 .61 33.50 36.84 20.35  33 . 37  19.13  34 .65  SUITE  variation  is attributed  variation  between i n d i v i d u a l  six  variables  variables  in Table  intra-taxonomic  PULVPUB,  BRACTLEN,  environmental, considered  Ignoring emphasized  As  that  Figure  variation  of separate  i s one o f r o t a t i o n  that  of c l i m a t i c ,  of t h e s e  samples  f o r a l l 36  t h a t as a r e s u l t  variable  ANOVAs of  CENCYABX, ADXSTOM,  between t h e two t a x a  variation  the primary  than  should  results.  variation  of the t o t a l  16 i l l u s t r a t e s ,  taxa.  variation  The d i v e r s i t y  intra-individual  be n o t e d  i s more  t h a t t h e r e were o n l y  (ABXANG,  i n a p p r e c i a t i n g these  17 p e r c e n t  ordinations variables  BRACTAP).  and g e o g r a p h i c  However, i t s h o u l d tree  23 i n d i c a t e d  variation  the d i s t i n c t i o n  variation is  trees within a population  t h a t had an i n t e r - t a x o n o m i c  exceeded  be  v a r i a t i o n between  Generally, there  between p o p u l a t i o n s o r between h y p o t h e s i z e d separate  12 . 86  t o cone m o r p h o l o g y and  and e x c e e d s t h e c o r r e s p o n d i n g  t o twig morphology.  12 .69 21 . 26 4 .62 31.14 41 .32  -  The s m a l l e s t amount o f i n t r a - i n d i v i d u a l  attributed  E  variables  ( F i g . 16).  of the a v e r a g i n g  by  has been removed.  d i f f e r e n c e between  s u i t e s and t h a t b a s e d on a l l  of c o - o r d i n a t e axes  r a t h e r than t h e  t F i g u r e 16. O r d i n a t i o n s o f f i r s t two c o m p o n e n t s o f PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r p o p u l a t i o n s o f s t a n d a r d s o f P_^ e n g e 1 m a n n i 1 a n d P. s 1 t c h e n s i s. S c o r e s b a s e d o n PCAs g i v e n i n T a b l e 4 3 . A p p e n d i x III G l y p h s r e p r e s e n t means f o r i n d i v i d u a l t r e e s . G l y p h s a s i n F i g u r e 10. Not a l l i n d i v i d u a l t r e e s c o u l d be p l o t t e d .  LEAF  ANATOMY  L E A F MORPHOLOGY  O O oo' ft O O  A A ^  iO O  _  TWIG  TOTAL  MORPHOLOGY  .v.\  o  A 'A A  A  CONE MORPHOLOGY  > AS'  ^  A A  A  A  o  o  1 30  emergence o f any f u n d a m e n t a l Based  upon  which account hypothesized SCALWID,  t h e component  f o r the l a r g e s t taxa  Further,  suggest  that  that  f o r the largest  not c o - i n c i d e n t  o f t h e components  between t h e  ABXANG, ADXANG,  best  CENCYABX,  RESCYLOC, PULVPUB, and describe  the p o l a r i t y  of these  i n the  component  p o l a r i t y i s not s t r i c t l y  a  reflection  T h e s e a r e n o t a l l t h e same v a r i a b l e s  These r e s u l t s suggest is  difference  t h e s i z e and s i g n  size differences.  account  correlations  BRACTAP, ADXSTOM, RESCYNO,  correlations of  pattern.  i t appears that  TIPWID a r e t h e v a r i a b l e s data.  new  differences  that  with  between h y p o t h e s i z e d  the hypothesized  that  taxa.  taxonomic p o l a r i t y  the p o l a r i t y of the d a t a .  3.1.2 P_;_ e n g e l m a n n i i , P. s i t c h e n s i s , and Pj_ g l a u c a .  Adding further  the populations  polarizes  intra-taxonomic (Table  24).  indicated  variation  In a d d i t i o n  that  Component  component, respect  variation  there  of other  is still  to the s i x i n d i v i d u a l  variables  really  components  variation  that only  for.  that were  the a d d i t i o n effected  also  o f two  the f i r s t  remained a p p r o x i m a t e l y  The a d d i t i o n  exceeded  between t h e t a x a .  and m a g n i t u d e o f component  accounted  larger  variation  suggest  glauca  a  Quebec  inter-taxonomic  to distinguishing  correlations  to sign  than  from w e s t e r n  v a r i a t i o n , NEEDEP and FREESCAL  important  populations  however  had an i n t e r - t a x o n o m i c  intra-taxonomic considered  the data,  o f P_^ g l a u c a  t h e same  correlations  with  and t h e  o f t h e two p o p u l a t i o n s o f  131  Table 24. M u l t i v a r i a t e apportionment of v a r i a t i o n f o r separate v a r i a b l e s u i t e s due to d i f f e r e n c e s between taxa, p o p u l a t i o n s , and i n d i v i d u a l s of standard engelmanni i , P. glauca, and F\ s i t c h e n s i s . A b b r e v i a t i o n of PCAs and ANOVAs g i v e n in Table 44, Appendix I I I . Symbols g i v e n in MODEL 2. O r d i n a t i o n s of r e s u l t a n t component scores g i v e n i n F i g u r e 17.  VARIABLE SUITE  p.  glauca  from  35.68 29 . 98 4 1 . 70 35.39 38 .90  27 13 16 19 24  39 90 48 39 15  20. 75 32 .68 35.92 18 .64  x VEGETATIVE  35 . 79  .19 . 26  29 . 78  given  morphology. variation based  on  the  Like  the  variability 23,  results  the  among t h e  three  other  taxa  two  glauca  taxa. with  appears  m o r p h o l o g y and variables,  like  distinction  Again,  to  variation  in Figure  of  data  the  anatomy and  BRACTLEN  t o the  16,  shown that  and  other  f o r BRACTAP a  lack  similarity  two  to  the  cone  morphology.  taxa  for  leaf  o r d i n a t i o n of a l l 36  f u r t h e r emphasized  the  the  P _ ^ s i t c h e n s i s. with  samples  for standards the  that  polarity.  anatomy and  The  twig  than  a p p e a r s t o be  striking  a polar position  variation  remembered  populations  inter-individual  there  and  leaf  morphology.  of  of  polarizing variables  between E \ e n g e l m a n n i i and  s e c o n d component  be  23,  13.17  P ^ engelmann i i i s i n t e r m e d i a t e  however c o n s t i t u t e s o n l y  range of  a result  substitution  intermediate  that  16.18 2 1 . 78 5.91 26.02 36 . 95  -  for leaf  The  emphasize  respect  twig  E  v a r i a b l e s were g r e a t e r  taxonomic  ( F i g . 17)  as  in Table  a d d i t i o n of CENCYLAT. between  . . . . .  only  variables.  above, w i t h  Ordinations  should  but  vegetative  reproductive  of c o - i n c i d e n c e  p.  the  in Table  b a s e d on  r e m a i n e d as and  (mva) C( AB)  LEAF ANATOMY LEAF MORPHOLOGY TWIG MORPHOLOGY CONE MORPHOLOGY TOTAL  increased  that  %SS B (A)  A  standards  of of  P .  glauca  to  the  within  the  respect fall  F\_ e n q e l m a n n i i . E*\ g l a u c a  and  It  F i g u r e 17. O r d i n a t i o n s o f f i r s t two c o m p o n e n t s o f PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r p o p u l a t i o n s o f s t a n d a r d s o f P_^ e n g e 1 m a n n i i , P. g l a u c a . a n d P_^ s i t c h e n s i s S c o r e s b a s e d o n PCAs g i v e n i n T a b l e 44, Appendix I I I . G l y p h s r e p r e s e n t means o f i n d i v i d u a l t r e e s a n d a r e a s i n F i g u r e 16 e x c e p t s q u a r e s r e p r e s e n t i n d v i d u a l s o f P. g 1 a u c a . Not a l l i n d i v i d u a l t r e e s c o u l d b e p l o t t e d d u e t o o v e r l a p .  LEAF  ANATOMY  ft  A '.^VA  L E A F MORPHOLOGY  <9o  OO  CONE  MORPHOLOGY  TWIG  TOTAL  MORPHOLOGY  O  oo oO A  A'/TS.  A  1#°  A  o°  A  I. A  A\  oo o o ax o oo o° ooo 9>  o  CO  ro  133  P. e n q e l m a n n i i came  from p o p u l a t i o n s on d i f f e r e n t  s i d e s of the  cont i nent.  3.2  Individual  3.2.1  decreased  reflected axes  sitchensis.  morphological  c o n s i d e r i n g more t h a n  few  and p u t a t i v e t a x o n o m i c r e p r e s e n t a t i v e s .  E"\ e n q e l m a n n i i and P^  The by  standards  just  and a n a t o m i c a l  p o p u l a t i o n s of s t a n d a r d s  i n t h e amount o f v a r i a t i o n ( T a b l e 25) compared  polarity  accounted  to those  created  was  f o r by t h e  in Table  23.  first  In s p i t e of  T a b l e 25. M u l t i v a r i a t e a p p o r t i o n m e n t of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between t a x a and i n d i v i d u a l s of s t a n d a r d a n d p u t a t i v e P^ e n g e l m a n n i i a n d P_^ s i t c h e n s i s. A b b r e v i a t i o n o f PCAs a n d ANOVAs g i v e n i n T a b l e 4 5 , A p p e n d i x I I I . Symbols g i v e n i n MODEL 3. O r d i n a t i o n s of r e s u l t a n t component s c o r e s g i v e n i n F i g u r e 18.  %SS VARIABLE LEAF LEAF TWIG CONE TOTAL x  this  The  ANATOMY MORPHOLOGY MORPHOLOGY MORPHOLOGY  VEGETATIVE  decreased  significantly that  polarity, correlated  fundamentally  new  amount o f v a r i a t i o n  percent  less  Figure  A  SUITE  than  that  18 f u r t h e r  (mva) B( A)  31 .84 20 . 93 24 .61 36.44 28 .01  51 56 67 23  25 . 79  58 . 45  E 16 .69 22 .43 8 . 24 29 .03 7 1.97  .47 . 73 . 16 . 28  -  15 . 76  t h e component c o r r e l a t i o n s with  those  sources  i n ' T a b l e 23, i m p l y i n g  of v a r i a t i o n  attributed given  given  t o taxa  in Table  were  have n o t been i n Table  25 i s o n l y 3  23.  corroborates that  new  added.  s o u r c e s of  1 34  variation Figure  have not  16,  the d i f f e r e n c e s  been o b s c u r e d . trees  been a d d e d .  i t was  On  t h a t the  t h e C a s c a d e M o u n t a i n s and  Oregon and The  were t h e the  study  variables  separate  variable  had  suites  with  decreased,  taxa  Similarly southern  was  than  ( T a b l e 23),  only  five  the a n a l y s i s  component  the d i f f e r e n c e s  of - p o p u l a t i o n s  25,  and  variation,  was  inter-individual  less  than  of more by  f o r the of  of  the  unchanged,  individuals compared  the h y p o t h e s i z e d  was  data  intra-taxonomic  8 percent  i t was  variation),  that  22.  polarity  inter-individual as  All  impression  remained v i r t u a l l y  r e m o v a l of  such  18  an  P_j_ g l a u c a .  the  between t a x a the  Figure  in Table  25,  addition  i n c r e a s e d the  Inter-population of t h e  The  variability  s u b s t a n t i a t e s the  in Table  variation  decreased  Like Table  variability.  correlations  (Table 26).  intra-taxonomic.  intra-taxonomic  sitchensis,  variation  25%  from  that there  variation  r e p o r t e d above  inter-taxonomic  24.  an  further  E\_ e n g e l m a n n i i , P.  Table  from  indicated  that exceeded  inter-taxonomic  separated  the  co-incident.  sitchensis  Further  variability  the  and  different  (NEEDEP, ABXANG, CENCYABX, ADXSTOM, PULVPUB) had  exceeded  As  are  of v a r i a t i o n  reproductive variables  3.2.2  for  have  area.  variation.  inter-taxonomic  poorly  Selkirks  more i n t r a - t a x o n o m i c  inter-taxonomic  scores  taxa  s a m p l e s of P_;_ e n g e l m a n n i i  samples of  apportionment  substantially  with  between t h e h y p o t h e s i z e d  i n s p e c t i n g t h e component  observed  co-incident  However, compared  to  effect  variability.  ( i . e . approximately obscured  by  F i g u r e 18. O r d i n a t i o n s o f f i r s t two c o m p o n e n t s of PCAs of s e p a r a t e v a r i a b l e s u i t e s f o r a l l s t a n d a r d s a n d p u t a t i v e s o f P_^ e n g e I m a n n i i P. s i t c h e n s i s . S c o r e s b a s e d on PCAs g i v e n i n T a b l e 4 5 . A p p e n d i x G l y p h s a s i n F i g u r e 16.  L E A F ANATOMY  L E A F MORPHOLOGY  CONE  MORPHOLOGY  TWIG MORPHOLOGY  and III.  TOTAL  OO  A  w  A AV'A  o  o  (40 5O •/.)  oi  1 36  T a b l e 26. M u l t i v a r i a t e a p p o r t i o n m e n t of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s between t a x a and i n d i v i d u a l s of s t a n d a r d a n d p u t a t i v e P_;_ e n g e 1 mann i i . P . g l a u c a . a n d P_^ s i t c h e n s i s . A b b r e v i a t i o n o f PCAs a n d ANOVAs g i v e n i n T a b l e 4 6 , A p p e n d i x III. S y m b o l s g i v e n i n MODEL 3. O r d i n a t i o n s of r e s u l t a n t component s c o r e s g i v e n i n F i g u r e 19.  %SS VARIABLE LEAF LEAF TWIG CONE TOTAL x  ANATOMY MORPHOLOGY MORPHOLOGY MORPHOLOGY  VEGETATIVE  inter-individual impression all  A  SUITE  51 54 66 65  .03 . 32 . 48 .72 -  16 . 90 22 . 53 8 . 52 20. 55 69 . 12  26 . 20  57 . 28  16 . 52  Figure  separated  the v a r i a t i o n  3.3 A r t i f i c i a l  19 f u r t h e r c o r r o b o r a t e s  taxa.  36 v a r i a b l e s f u r t h e r e m p h a s i z e d  within  E  32 . 10 2 1 50 . 25 . 0 0 13 . 73 30. 88  variation.  of p o o r l y  (mva) B (A )  this  The o r d i n a t i o n b a s e d on the p o l a r i t y  of  glauca  of P^ e n g e l m a n n i i .  hybrids  i n the context  of p o p u l a t i o n s of  standards.  The (Table  amount o f v a r i a t i o n  40, A p p e n d i x  in Table  23.  was a t e n d e n c y  maternal closer  component  f o r these  27) a r e s i m i l a r  to those  known h y b r i d s , even  f o r t h e two t a x a  Particularly  "P^ e n g e l m a n n i i "  to  separate  III = Table  f o r by t h e s e p a r a t e  The o r d i n a t i o n s ( F i g . 20) s u g g e s t e d  way o f i n t e r m e d i a c y  ordination.  accounted  sitchensis  scores  important  opposite  Brunswick  given  little  i n the  though  there  p o l e s of the  was t h e o b s e r v a t i o n  f o r t h e New  than  variable suites.  t o occupy  PCAs  hybrids  that the was  were i t s p r o g e n y b a s e d on t h e  Examination  for individual  of the v a r i a n c e of  t r e e s d i d not i n d i c a t e  a g r o u p , t h e h y b r i d s were any more v a r i a b l e t h a n  the  t h a t , as  standards.  F i g u r e 19. O r d i n a t i o n s o f f i r s t two c o m p o n e n t s o f PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r a l l i n i d i v i d u a l s t a n d a r d s and p u t a t i v e s o f P . e n g e 1 mann i i . P . g l a u c a . a n d P_^ s i t c h e n s i s . S c o r e s b a s e d on PCAs g i v e n in T a b l e 46, A p p e n d i x III. G l y p h s as i n F i g u r e 17.  LEAF ANATOMY  LEAF MORPHOLOGY  CONE MORPHOLOGY  TWIG MORPHOLOGY  F i g u r e 20. O r d i n a t i o n s o f f i r s t two c o m p o n e n t s o f PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r sampled p o p u l a t i o n s of s t a n d a r d s and a r t i f i c a l hybrids. S c o r e s b a s e d o n PCAs g i v e n i n T a b l e 4 7 , A p p e n d i x III G l y p h s a s i n F i g u r e 16. F i l l e d t r i a n g l e - m a t e r n a l P_^ enge1mann i i f o r New B r u n s w i c k h y b r i d s . H a l f - f i l l e d glyphs - a r t i f i c a l hybrids u p p e r h a l f f i l l e d - New B r u n s w i c k h y b r i d s ; l o w e r h a l f f i l l e d - Red Rock h y b r i d s .  L E A F ANATOMY  LEAF MORPHOLOGY  TWIG MORPHOLOGY  VEGETATIVE  00  co  139  The  o r d i n a t i o n o f t h e PCA b a s e d  (Fig.  20) c o r r o b o r a t e s  hybrids  a r e more l i k e  taxon.  The p o l a r i t y  suggests in  a similarity  Figures  the impression the maternal  that  taxon  of the h y b r i d s  the  than  artificial  the p a t e r n a l  on t h e s e c o n d component  t o the d i s p o s i t i o n  of Pj_ g l a u c a  standards  16 and 18.  3.4 A r t i f i c i a l  and p u t a t i v e  ANOVAs of PCAs o f j u s t (Table  on a l l 21 v e g e t a t i v e v a r i a b l e s  28) s u g g e s t e d  differences  that  hybrids. the a r t i f i c i a l  there  and p u t a t i v e  were s m a l l  between t h e two g r o u p s .  Most  hybrids  but s i g n i f i c a n t of the v a r i a t i o n i s  T a b l e 28. M u l t i v a r i a t e a p p o r t i o n m e n t of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o d i f f e r e n c e s b e t w e e n a r t i f i c a l a n d p u t a t i v e h y b r i d s o f P. enge1mann i i a n d P_^ s i t c h e n s i s. A b b r e v i a t i o n o f PCAs a n d ANOVAs g i v e n i n T a b l e 48, A p p e n d i x III. S y m b o l s g i v e n i n MODEL 3. O r d i n a t i o n s of r e s u l t a n t component s c o r e s g i v e n i n F i g u r e 21.  %SS VARIABLE  SUITE  LEAF ANATOMY LEAF MORPHOLOGY TWIG MORPHOLOGY VEGETATIVE  between  individual  variation  remains  Figure the  21  A 3.67 14.06 1.15 8.82  E  76.28 70.17 90.09 -  20.05 15.77 8.76 91.18  t r e e s and t h e amount o f similar  to that  indicates a virtual  two g r o u p s , an i m p r e s s i o n  inter-individual  (mva) B(A)  variation  presented  in Tables  23 and 24.  o v e r l a p of the v a r i a t i o n  corroborated  (Table  intra-individual  28).  of  by t h e h i g h  The p u t a t i v e  hybrids  140  constitute  a significantly  individuals As first are  the a r t i f i c i a l  evidenced  few  not  III).  than  more h e t e r o g e n o u s g r o u p o f  by  hybrids.  t h e amount of v a r i a t i o n  components of  these  as p r o n o u n c e d as  PCAs, t h e  those  variation  were b e i n g d e s c r i b e d .  variables  had  exceeded source  a variation  of v a r i a t i o n  Individual  the p o l a r i t y alter  was  of  the  two  between  two  p u t a t i v e s , and  t h e d a t a compared  responsible  individual  To  variable  BRACTAP.  appears to e x i s t when t h e  variation  " h y b r i d s " were a l s o  v e c t o r s of v a r i a b l e s taxonomic p o l a r i t y  23 and  of  largest  standards  only 24,  the  slightly  and  did  variables  t h e p a t t e r n of  suites  a r e : NEEDEP, PHLEND,  SCALEN. the  two  indicates taxa,  If only variables  examined.  polarity  t h a t what s e p a r a t i o n  is virtually The  t h a t best d e s c r i b e the  or a c t u a l  that  a r e : NEEDEP, ABXANG, CENCYABX,  F i g u r e 22  between t h e  The  s a m p l e s of  iterate,  i s considered,  for polarization  BRACTLEN, and  of h y b r i d s  decreased  to Table  XYLEND, ADXSTOM, RESCYNO, TIPWID, and inter-individual  p a t t e r n s of  "hybrids".  ( T a b l e 29)  component c o r r e l a t i o n s .  of t h e  Appendix  individual  types  h y b r i d s t o the  taxa  data  trees.  r e s p o n s i b l e f o r the p o l a r i z a t i o n  variation  the  40,  the  radically  w i t h i n a g r o u p of h y b r i d s .  standards,  of  None of  between t h e  the h y p o t h e s i z e d  and'putatives  most  (= T a b l e were  for  i n these  that p r e v i o u s l y undescribed  the v a r i a t i o n  Adding  not  27  However, t h e c o m p o n e n t - c o r r e l a t i o n s suggesting  3.5  polarity  in Table  different  accounted  of  obscured  c o - i n c i d e n c e of hypothesized  the data  are at  an  the  F i g u r e 21. O r d i n a t i o n of f i r s t two c o m p o n e n t s of PCAs o f s e p a r a t e v a r i a b l e s u i t e s f o r i n d i v i d u a l a r t i f i c i a l h y b r i d s and p u t a t i v e hybrids. S c o r e s b a s e d o n PCAs g i v e n i n T a b l e 48, A p p e n d i x III. H a l f - f i l l e d glyphs - a r t i f i c i a l hybrids. Open c i r c l e s - p u t a t i v e hybrids. G l y p h s r e p r e s e n t means of I n d i v i d u a l trees.  L E A F ANATOMY  L E A F MORPHOLOGY  TWIG MORPHOLOGY  VEGETATIVE  1 42  T a b l e 29. M u l t i v a r i a t e apportionment of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s due t o e i t h e r d i f f e r e n c e s between t a x a or i n d i v i d u a l s of P. e n g e l m a n n i i , P. s i t c h e n s i s, o r t h e i r p u t a t i v e h y b r i d . A b b r e v i a t i o n o f PCAs a n d ANOVAs i n T a b l e 4 9 , A p p e n d i x I I I . %SS (mva) for t a x a b a s e d o n o n l y s t a n d a r d s a n d p u t a t i v e s o f t h e two t a x a . O r d i n a t i o n s o f r e s u l t a n t c o m p o n e n t s c o r e s g i v e n i n F i g u r e s 22 a n d 2 3 .  %SS  (mva)  TAXA VARIABLE LEAF LEAF TWIG CONE TOTAL  x  angle  co-incidence  values, than  and  between i n d i v i d u a l s  between  19 . 52  85 . 27  14 .93  suggest  .00 .08 . 10 . 89  partial  to a l t e r  be i d e n t i f i e d taxa  no  Individual putative  were no more v a r i a b l e t h a n  populations  of standards,  although  Further, rather  as i n d i c a t e d  populations  examples of both  populations  of a  Owing t o t h e p r o x i m i t y  swarms o f s t a n d a r d s ,  and h y b r i d s .  and u s i n g  intermediacy  sitchensis  hybrids.  that contained  these  for less  variation  variation.  suggest  engelmanni i or  20 f o r t h e a r t i f i c i a l  on  accounts  the impression  and a n a t o m i c a l  of the p u t a t i v e h y b r i d s with  Based  individuals.  ignoring intra-individual  o v e r l a p of the point  parental  80.48  was e m p h a s i z e d .  d i f f e r e n c e s between t a x a  of the v a r i a t i o n  confusion  could  17 21 6 33  c o n s i d e r i n g taxonomic c i r c u m s c r i p t i o n , the  relation  in Figure  T h i s would  83.00 78.92 93 .90 66 . 1 1  and d a t a .  36. v a r i a b l e s does l i t t l e  than  . 30 . 63 .80 .05 .99  72 . 24  27 . 76  continuum of m o r p h o l o g i c a l the  64 75 77 82 71  70 37 20 95 07  22° t o e a c h o t h e r .  Analytically, all  . . . . .  E  A  E  "x  the r e l a t i v e  half  35 25 22 17 28  of h y p o t h e s i s  Without differences  ANATOMY MORPHOLOGY MORPHOLOGY MORPHOLOGY  VEGETATIVE  of about  INDIVIDUAL  A  SUITE  hypothesized  hybrid  some o f t h e i n d i v i d u a l  as a group they  were  F i g u r e 22. O r d i n a t i o n s o f f i r s t two c o m p o n e n t s o f PCAs o f s e p a r a t e variable suites for all i n d i v i d u a l s of P_^ enge1mann i i , P. s i t c h e n s i s. and " h y b r i d s " . S c o r e s b a s e d on P C A ' s g i v e n i n T a b l e 49, A p p e n d i x III. G l y p h s r e p r e s e n t means o f i n d i v i d u a l t r e e s a s i n F i g u r e 16. Ha 1 f - f 1 11ed g l y p h s r e p r e s e n t " h y b r i d s " . Not a l l i n d i v i d u a l t r e e s c o u l d be p l o t t e d .  1 44  significantly  4.  Discussion. The  in  larger.  morphological  southwestern  and  British  c o n t i n u u m of v a r i a t i o n trans-individual hypothesized P.  sitchensis  others  polarity The of  they  of t h e  inclusion  of  is  number  instructive  results  relation  taxa  by  the  two  the c o n c l u s i o n that more  than  p o p u l a t i o n a l or  features.  The  variables  between P_^ e n g e l m a n n i i  the  that  and  same as have been used  is attributable  m e a s u r e s , no  ratio  sources  s o u r c e s of v a r i a t i o n samples  from  areas  t o compare t h e s e the  of P a r k e r  of P^  a  p a t t e r n of  suggested  the o n l y v a r i a b l e s  intra-individual  (1976) d e a l i n g w i t h the  there exists  that  i n c o n c l u s i o n s r e g a r d i n g the  ipsative  of  sampled  determine  by  the  data.  inter-individual large  of P i c e a  t h e e x c e p t i o n of t h e p r e v i o u s l y  a r e not  difference  (no  that  further  anatomy v a r i a b l e s ,  the h y p o t h e s i z e d  analysis  that  a discrete  of p r o m i n e n t  polarity  are, with  leaf  - yet  results  morphological  the taxonomic  unreported  such,as  exaggerations  intra-populational  P.  suggest  r a t h e r than  The  variation  P_;_ e n g e l m a n n i i r e p r e s e n t l i t t l e  nomenclatural  support  Columbia  variation  taxa.  g l a u c a and  anatomical  glauca  and  McLachlan  t o P^ m a r i a n a .  variables),  c o u l d be  the  a g a i n s t which  compared,  and  sampled p r e v i o u s l y .  results -  t o t h e method of  of v a r i a t i o n  not  "P^ m a r i a n a  appropriateness  with  those  rubens"  of  Gordon  complex  (1978) d e a l i n g w i t h Clearly  the  It  and the  relation  shown  1 45  h e r e between discrete  as that  P. g l a u c a , as  s i t c h e n s i s a n d P_;_  nomenclatural  the  shown h e r e  (Mitton  i s i t as  a n d P^ e n g e l m a n n i i  1981).  R e g a r d l e s s of t h e  the r e s u l t s presented here  n o t be e x c l u d e d  from  suggest  i n v e s t i g a t i o n s of  complex.  variation.  l a r g e s t s o u r c e o f i n t e r - i n d i v i d u a l v a r i a t i o n was  attributed  t o v a r i a t i o n between  population.  This  researchers  corroborates  Similar  anatomical  v a r i a b l e s , as w e l l  individual trees  as well  results are reported  isoenzymes and other  have p r o p o s e d  that  this  high  variation  i s r e l a t e d to the s u c c e s s i o n a l  (Rehfeldt  and L e s t e r  1969), however  consistently  shown f o r many s p e c i e s  successional  status  hypothesized  relation  There  (Guires  variables,  and p h y s i o l o g i c a l v a r i a b l e s . materials  grown p r o v e n a n c e s , p r o g e n y , and f a m i l y  researchers  of other  f o r m o r p h o l o g i c a l and  as f o r growth and y i e l d  chemicals,  a  other  as the m a j o r i t y  T h e s e r e s u l t s a r e b a s e d on n a t u r a l l y o c c u r r i n g nursery  within  t h e f i n d i n g s made by  working with P i c e a ,  conifers.  as  convoluted  1968), n o r P^ e n g e l m a n n i i and  considerations,  4.1 I n t r a - p o p u l a t i o n The  and E\_ r u b e n s , n o r  between P^ g l a u c a  and A n d a l o r a  P_;_ s i t c h e n s i s s h o u l d P^ g l a u c a  mariana  However, n e i t h e r  a l s o La R o i and D u g l e  P. pungens  that  for  a n d P^ m a r i a n a .  the r e l a t i o n  (see  reported  engelmanni i i s not as  as w e l l  trials.  Some  within-population status  the l a r g e  of the s p e c i e s  variability  i r r e s p e c t i v e of t h e i r  1984) s u g g e s t s  that  this  may be e r r o n e o u s .  i s no i n d e p e n d e n t  information  consistantly avaialble  1 46  in  this  study  concerning Research not  t h a t would permit  the  into  source this  been a d d r e s s e d  undertaken most  of  of  (Adams  1981).  et  a l . 1981a,b; E h r l i c h  well  environment  tendered  the  and  Mitton  1976,  nature  longevity such a  of  long  suggested  life  vagrancies  variation suggests  others  1969;  Hamrick  that  explanations  1977;  shown i n  b a s e d on  associated high  selection  and  the  1978,  presumed  of  Allard  and  may  be  King  (1972)  environmental  Coles and  Dancik  variation  the  stands and  Fowler  yearly  1984)  sources  as  well  of (Maze  1984)  principal  in naturally occurring  (1961) comments and 1979;  over  mutatiqn~were I variation  extrinsic  ordering process  Rowe's  1977;  Stern  1981)  for within-population alone  sizes,  tendered  1983;  s t r u c t u r e ( K i n g and  inefficiency  herbs.  fecundity  family structure in forest  Shaw and  the  frequent  within population  of  Additionally,  concerning  et a l .  variation  plasticity.  reproduction  (Falkenhagen  selection  i n agreement w i t h  phenotypi'c  high  of b r e e d i n g  to account  Mitton,  1976)  edaphic  have  Linhart,  (1979a) has  e_t a l . 1977;  variation-generating stands.  Raven  explanations  to n a t u r a l  Rehfeldt  l a c k of  demonstrated  proposed  1981;  e t a l . 1981a,b; R e h f e l d t  Mitton,  the  span.  f o r the  the  Allard  the  has  Where s u c h s t u d i e s have been  respect  other  variation.  of c o n i f e r s g e n e r a l l y  with  and  t r e e s , and  the  exploration  conifer lineage, large population  demonstration  (Linhart, 1976;  the  b a s e d on  that  explanations The  of  the  explanations  1981)  and 1977;  (1983) t e n d e r e d  ancient  as  (Shaw and  known m i c r o g e o g r a p h i c  Mitton  variation  or h y p o t h e s e s a d v a n c e d  by  systematic  intra-individual  aspect  f r e q u e n t l y been  Grant  this  the  Burgar  1964;  preconditioning  those  of  Bjornstad  (including  1 47  maternal  effects) also  should  be  considered  in  such  explanat ions. These o b s e r v a t i o n s speculations concerning that  the  a t an  suggest  that  factors  by  emergent  extremely  reproduction  from an  mechanism  large and  local and  conditional  Natural  and  operative  upon t h e  specific  scale.  results  standards  may  these  the  suggest be  results  primary  n o v e l t i e s u l t i m a t e l y become If n a t u r a l s e l e c t i o n mediated accepted  results  and  hypothesis  s e l e c t i o n may  as  the  those  causal  of many  (see a l s o well  be  s c a l e would a p p e a r lineage evolving  from  suggest  the  that  the  comparison hybrids  both  be  the  of p r o n o u n c e d m a t e r n a l  at  variance  hybrids  result  with  other  where s u c h  pronounced.  The  in respect  others  Mitton  operative,  t o be .  (Rehfeldt  1984b).  -like  I n d e e d , b a s e d on  could  information  of  the  to  These  have not  This  may  results  are  been  artificial  t h a n E\  provided  as  of c o n t r o l l e d  a l s o suggest  rather  and  recognize  variability.  effects.  effects  disposition  maternal parent the  difficult  form and  large maternal  polar  c o n t r o l l e d hybrids  m u l t i v a r i a t e analyses  a v a i l a b l e maternal parent glauca  to  of  are  intermediates  P.  component  Further, be  and  Natural h y b r i d i z a t i o n . The  the  this  d i s p e r s a l may  to c o n t r a d i c t t h i s  for c o n i f e r s .  of  variation  e c o l o g i c a l processes  i s t o be  i n e v o l u t i o n , these  i t s effects  size  a n c e s t r a l taxon.  differentiation  would a p p e a r  4.2  the  which e v o l u t i o n a r y  population  but  inter-individual  major e v o l u t i o n a r y  occurring  1983)  on  as  hybrids  and  a more engelmannii.  f o r the  origin  of  at  1 48  least  the m a t e r n a l  (Tree  70418, A p p e n d i x  situation  i s not  the maternal elevation  the  hybrids  hypothesized the  easily without  results shares the  Roche's  Other are  p o s s i b l e , but  effects  the  the  i t would be  r a n g e of v a r i a b i l i t y new  patterns  here  f o r the to  hybrids  e n g e l m a n n i i x P^  the  remains enigmatic higher  be  to  or  taxa  identify  hybrids  two  increased  that P^  then glauca  sitchensis  as  This contradicts  t h a t P_^ e n g e l m a n n i i a p p e a r s and  o n l y cone  P_;_ g l a u c a  with  heterogeneity  respect of  the  ; however,  morphology.  putative hybrids  to the  t o the putative  data  artificial presented  so  hybrids  T h i s e x p l a n a t i o n i s not a d e q u a t e f o r t h e Red Rock m a t e r i a l as t h e m a t e r n a l p a r e n t s were p o l l i n a t e d i n m i d - w i n t e r i n V e r n o n , where t h e r e were no o t h e r t r e e s p r e s e n t t h a t were c o n t r i b u t i n g pollen, pers. comm., G u y l a K i s s , B r i t i s h C o l u m b i a M i n i s t r y of F o r e s t s , Vernon. 1  a  falls  species  r u l e d out,  suggest  sitchensis.  were b a s e d on  of  the  of  exprimental  artificial  e n g e l m a n n i i and  Roche's d a t a  The  Regardless  difficult  of  the  experimental  hypothesized  of v a r i a t i o n  between P_j_ s i t c h e n s i s  far.  the  or  two  high  a p p e a r a n c e of  primarily  parent  between t h e  a p p e a r a n c e of  This  of c o m p a r a t i v e l y  f o r the  I f expermental c a u s e s can  relation  explanation.  Rock h y b r i d i z a t i o n as  are  of m a t e r n a l  intermediate  hybrids  Red  explanations  (1969) s u g g e s t i o n s  The  likely  were from a v a r i e t y  same r e l a t i o n  h y b r i d P^  is a  f o r the  t h a t , at best,  presented the  Brunswick h y b r i d i z a t i o n  1  suggesting  variability.  New  i s o l a t i o n ..during p o l l i n a t i o n ) .  Further, within  the  II), this  poor d i s t i n c t i o n  would s u g g e s t hybrid.  of  obvious  parents  inadequate  error,  as  origins.  artificial (e.g.  parent  1 49  (Fig.  20,  21)  number of  could  parent  reflect:  trees  than  environmental  variability  common g a r d e n  conditions  c o n s e q u e n c e of assignment  of  the  that are  historical selection  here are  the  two  explanations  explorations conducted,  polymorphic outcome of  result  between  introgressive  of  such  of, at  of  requires  hybrids  further  least,  b a s e d on  co-incident  with  a the  of  enigmatic.  based  elaboration.  out  on  Until  either  within  Regardless to note  glauca with  rule  v a r i a t i o n encountered  differentiation  that  British  variation.  explanations  to  i t is  i n t r o g r e s s i o n or  observed  pattern  hybrids  contemporaneous,  d o e s not  i t i s important  from  the  inappropriate  in southwestern  This  f o r the  engelmannii  different  than the  an  hybrids  artificial  subsequent  remain  research  greater  hybrids;  partially  hybrids  h y b r i d i z a t i o n or  fundamentally  putative  putative  the  taxa.  the  greater  here.  tendering  t a x o n must  relationship  alone  putative  a  hybrids;  artificial  were o n l y  h y b r i d i z a t i o n or as  the  a p p e a r a n c e of  the  not  hybridization  further  of  i n d i v i d u a l t r e e s as  Owing t o  Columbia  artificial  among t h e  of v a r i a t i o n shown  suggested  the  c o n t r i b u t i o n of  i n t r o g r e s s i v e h y b r i d i z a t i o n ; or,  g r o u p of v a r i a b l e s t h a t patterns  the  a of  that  single the  the  appears  P_^_ s i t c h e n s i s -  this  1 50  4.3  Intra-individual  The  results  systematic  presented  one  inherent  values  actualities. every of  of  i n the i d e n t i f i c a t i o n taxa  from  dealing with  In t h i s  context,  of l o c a t i o n  from  causes  of i n d i v i d u a l  tendencies  deviation).  i n nature,  independent  trees to  c h a r a c t e r s and  ( i . e . a mean) t h e r e  that the v a r i a b i l i t y  experimental  that the  rather  than  i t i s worth p o i n t i n g out t h a t f o r  o b s e r v a t i o n s c a n be i g n o r e d .  part,  context.  in  indicate  on t h e b a s i s o f s i n g l e  ( i . e . a standard  demonstrated entirely  the importance  These r e s u l t s  results  statistic  spread  emphasize  variation.  of the a p r i o r i  single  i n an i n t e r - i n d i v i d u a l  s t u d i e s o f c o n s i d e r i n g and q u a n t i f y i n g  intra-individual difficulty  variation  is a  I f i t c a n be  among v a r i a b l e then  statistic  the spread  values i s or v a r i a b i l i t y  If the v a r i a b i l i t y  of e x p e r i m e n t a l  results, in  error,  then  this  spread  c a n be i g n o r e d o n l y a t t h e e x p e n s e of m i s r e p r e s e n t i n g t h e  nature  of the s i t u a t i o n  location spread  described.  i s o n l y as m e a n i n g f u l  and t h e sample  demonstrated anatomical  The  size  i n Chapter  variation,  random and t h u s  British  being  variation  differentiation,  o c c u r s and i s not  n o t be i g n o r e d .  further  Thus,  here  suggest  that  in  southwestern  r e s e a r c h aimed a t e x p l a n a t i o n s of  investigations  c o u l d be a p p l i e d o v e r  illustrated  As was  m o r p h o l o g i c a l and  need n o t c o n s i d e r p r e v i o u s l y h y p o t h e s i z e d  circumscription.  variation  i t s cause,  s t a t i s t i c of  are based.  II, intra-individual  c o n c l u s i o n s reached Columbia  as the accompanying  upon w h i c h b o t h  whatever  should  A s t a t i s t i c of  here  without  taxonomic  of say, p o p u l a t i o n the complete  having  range of  to consider the  151  "taxon" for  t o w h i c h t h e p o p u l a t i o n may  population variation  context  of g e o g r a p h i c  with d i f f e r e n t i a t i o n (Tigerstedt of  how  suggest  1973;  the problem  than  t h u s c o u l d be  variation  assigned.  Explanations  addressed,  say  investigated  between m a r g i n a l and  Soule  1973;  and  Layton  i s approached,  the  results  situation  by  concerned  populations  1979).  Regardless  presented  concerning Picea in  previously considered.  i n the  others  central  Yeh  a more complex  North America  be  here  western  152  V.  INTER-INDIVIDUAL VARIATION: RELATIONSHIPS OF PATTERNS  OF  VARIATION.  1.  Introduction. Having  demonstrated  southwestern  British  morphological and  taxa.  Indeed,  proposing  Two of  general  variation.  correlating  to  clinal  are required to interpret of h y p o t h e s i z e d  this  circumscribing  suggests  that the  for variation.  h y p o t h e s e s c a n be o f f e r e d t o e x p l a i n  These c a n be t e n d e r e d  morphological  as the r e s u l t  and a n a t o m i c a l variation  of v a r i a t i o n .  strongly  suggest  of p o s s i b l e causes than  Larger  sources  causes  than  of v a r i a t i o n  do s m a l l e r  tendered  the e x t r i n s i c  dispositions  variation  Strong  patterns  of e i t h e r with  or a p p o r t i o n i n g  sources  only  p a t t e r n of  p a t t e r n of v a r i a t i o n  and/ or e x t r i n s i c  often  exists in  between P^ e n g e l m a n n i i  these  are  there  of taxonomic c i r c u m s c r i p t i o n i s unwarranted i n explanations  intrinsic  variation  irrespective  the c l i n a l  IV t h a t  a complex  explanations  of v a r i a t i o n  imposition  Columbia  and a n a t o m i c a l  P_;_ s i t c h e n s i s ,  pattern  i n Chapter  variation  c o r r e l a t i o n s more poor c o r r e l a t e s .  more s t r o n g l y i n d i c a t e p o s s i b l e  sources  as i f they  of v a r i a t i o n . were m u t u a l l y  explanations  toward e x t r i n s i c  These  e x c l u s i v e or that  are acceptable.  or i n t r i n s i c  hypotheses  A  priori  explanations i s  inappropr i a t e .  Extrinsic intrinsic. of  explanations  a r e o f f e r e d more  Such e x p l a n a t i o n s  phenotypic  variation  with  result  f r e q u e n t l y than a r e  from c o r r e l a t i n g  the environment  patterns  from which the  1 53  samples a r e t a k e n . are  expected  variation.  Discontinuities  t o be a c c o m p a n i e d In s p i t e  extrinsic correlates  1976)  there  broad  with  variation.  longitude,  formulated  latitude, variation;  environment  variation  i s referred  speci.es t h a t  show c o r r e l a t i o n  that  Intrinsic  and, w i t h i n  individual  selection. with the  are widely  distributed  are expected to Large  t o show c o r r e l a t i o n i n the plant  In  with  between respect to  environment.  relations  on c o r r e l a t i o n s  respect  amongst  relations;  with  i s a continuum of  into  to the l i f e  individuals  variation  of the  o f v a r i o u s ages o r o n t o g e n y .  there  c a n be d i v i d e d  parent-offspring  between t h e  plasticity".  with e x t r i n s i c c o r r e l a t e s ,  with  local  "phenotypic  are expected  ancestor-descendant  relations  climate,  and e l e v a t i o n ;  o f m o r p h o l o g i c a l and a n a t o m i c a l  duration  such as  scales:  t o under  e x p l a n a t i o n s a r e based  that  phenotypic  several  between p h e n o t y p e and g e o g r a p h y .  plants  surrounding  variation  into  Thorpe  intrinsic  the observed  i s correlated  p h e n o t y p e o f o r g a n s and p o s i t i o n  pattern  pressures,  and i n t e r - i n d i v i d u a l v a r i a t i o n a r e  under arguments o f n a t u r a l  Intra-individual  the  explanations,  E x p l a n a t i o n s b a s e d on t h e c o r r e l a t i o n s  environment  individual  to affect  environmental v a r i a t i o n ,  and t o p o - e d a p h i c  general,  selection  T h i s c o n t i n u u m c a n be d i v i d e d  associated  external  in pheotypic  reflect  i s hypothesized  geographic  climatic  by d i s c o n t i n u i t i e s  i s a c o n t i n u u m o f b o t h e x t r i n s i c and  that  variation.  distribution  o f t h e e m p h a s i s on e x t r i n s i c  (i.e.  factors  i n geographic  intervals  of  variable  of the organism:  reflecting  intrinsic  distant  and, i n t r a - i n d i v i d u a l  long-term  ancestors; variation  As  1 54  between s e p a r a t e  o r g a n s or  e x p l a n a t i o n s are  formulated  whereas with  intra-individual  respect  to  extrinsic  E x p l a n a t i o n s of There  or  variation  intrinsic  scales  of  variation  geographic  variation.  the  are  Similarly,  individuals  are  studies,  s u c h as  those  possible  variation to  variation.  into  apportion For  versus  a means of  S u c h an approximation  addressing  apportionment for  further  c o n d i t i o n s with c o n t r o l l e d specific  intrinsic  with  and  of  of  more  more d i s t a n t l y each o t h e r .  or  into  In  b a s e d on  a  occurring  intrinsic.  these the  s o u r c e s of of  apportionment  s e r v e s as  research conducted m a t i n g aimed a t factors  scales  variation  scales  or  However, i t i s  these v a r i o u s of  widely  related  effectively discriminate  variation  extrinsic  the  variation  in that  are  nature.  broad  environmental  intra-individual  subsequent c o r r e l a t i o n s  in  under n a t u r a l l y  extrinsic  variation  scales  that  of  ontogenetic.  historical  to  example, a p p o r t i o n m e n t  inter-individual  s e r v e s as  to  in  between  be  here,  sampling  i t i s impossible  are  relations  closer  reported  intra-individual  apportion  are  regardless  evolutionary  reflected  assumed t o  i n d i i v i d u a l s that  situations,  formulated  relation  local  complexed w i t h p a r e n t ^ o f f s p r i n g  limited  are  variation  and  relations  than are  and  nature,  variation,  approximate one-to-one  Ancestral  separated  genetics,  e x p l a n a t i o n s are  inter-individual  variation.  is  under p h y l o g e n e t i c s and  intra-individual  i s an  geographic  Inter-individual  development.  E x p l a n a t i o n s of their  characters.  of  into variation  variation of  a  variation.  first  under common  isolating  o p e r a t i v e at  garden  the a  scale  155  that  accounts  I n d e e d , an occurring  for  variation sound  S u c h programmes organism  being  generalized  silvicultural  or  tree  by  the  and  aspect  just  between d i v e r s e Falkenhagen  than  by  scale  physiology  Such a t t e n t i o n  programme.  or  of  the  economic  for  a  suggest  i f adequate are  variables that  an  relations  the  variables  and  the  status  morphological  well,  t h e y p r o v i d e an  variation:  result  of  variation  morphological  a_l. 1982).  and  Oka  1968). and  structures,  (Mitton,  inferences  the  view  the  being of  necessarily  dependency.  of  n a t u r e of  taxon  alternate  under  e_t a_l. 1980).  between g r o u p s  developmental  i s not  also  Variable  genetic  of  Such  inter-dependence  1983)^  inter-relationships  various  of  o r g a n i s m grows  important  in understanding  As  of  intercorrelations  disequilibrium"  evolutionary  the  important  between o r g a n s ,  (Maze  carry  are  et  (Morishima  implications  the  variables  developmental  Such  also  to  demonstration  (Small,  maintained.  "linkage to  The  purposes  out  respect  inter-relationship  aid  the  a  variation  variation  with  variables  simply  of  improvement natural  data.  naturally  theoretical  between g r o u p s of  g r o u p s of  inter-correlations of  i s the  analytic  evolutionary  examined.  of  initiation  of  consider  variables.  (1974) p o i n t s  only  rubric  to  variation  inter-relationships  the  guided  rather  g r o u p s of  more t h a n  carry  be  i n the  organism.  inter-relationships  and  apportionment the  to a p a r t i c u l a r  important  survives  the  variation  to  should  inter-individual  for  of  s o u r c e of  is essential  studied  predisposition  variables  largest  investigation  biologically  An  the  adaptive,  1 56  In a d d i t i o n of  to providing  v a r i a t i o n , there  variation studies  to that  reported  i n the l i t e r a t u r e .  b a s e d on n a t u r a l l y o c c u r r i n g  situations variation  b a s e d on t r e e s  occurring  pattern  would  trees  patterns  Comparison  on common  with  garden  of p a t t e r n s of  from a common g a r d e n faciliate  Comparison  i s straightforward;  the r e s u l t s based  i s not as d i r e c t .  Materials  f o r observed  i s a l s o a need t o r e l a t e t h e o b s e r v e d  however, c o m p a r i s o n s w i t h  2.  explanations  to the n a t u r a l l y  such a comparison.  and methods.  2.1Materials. Except P. g l a u c a ,  f o r the a r t i f i c i a l a l l the trees  c h a p t e r s were u s e d variation. grown  and the s t a n d a r d s of  sampled and u s e d  i n c o r r e l a t i n g with  i n the previous  the various  comparison  grown t r e e s c o u l d  P. s i t c h e n s i s .  between n a t u r a l l y o c c u r r i n g a n d  be made o n l y  f o r standards of  S i m i l a r l y , as the nursery  grown t r e e s  immature, c o m p a r i s o n s w i t h n a t u r a l l y grown t r e e s using  only  The  vegetative  lack  pertaining  v a r i a t i o n ( i . e . exact  r e q u i s i t e r e p l i c a t i o n within  a v e r a g e s of v a r i a b l e s p e r t r e e about  20 p e r c e n t  simplifies  were  could  still  be made  variables.  of i n f o r m a t i o n  intra-individual and  s c a l e s of  Owing t o l i m i t a t i o n s o f t h e number o f p r o v e n a n c e s  i n the nursery,  nursery  hybrids  of t h e t o t a l  to correlates f o r position within  positions)  be u s e d . variation.  analysis considerably  the tree  necessitated  Such a v e r a g i n g This  that removes  averaging  b u t any c o r r e l a t e s p e r t a i n  only  157  to the  inter-individual  data.  T h i s assumes t h a t  inter-individual  p o r t i o n of there  variation  Although populations of  o r g a n i z a t i o n cannot  of  trees, especially  Where t r e e s do available  occur  be  on  i s no  and  of  the  total  variation  interaction  intra-individual  Picea  do  occur  the  coast,  d i d not  in populations,  the  variation.  as  extremely  v a r i a b l e making a n a l y s i s  T h i s d o e s not  mean t h a t  the  variation the  to the  pattern  i s r e j e c t e d , merely  s a m p l e s a v a i l a b l e does not  least  an  a n a l y s i s which w i l l  permit  yield  example, t h e  regular  interior  w e l l c o n s t i t u t e an  may  e x t r e m e s of study. total is  Table  morphological  23  of  in t h i s  nature s u c h an  defined  important  continuum  suggests that  to  number  sizes  difficult.  of  anatomical  the  study  and  a n a l y s i s , or results.  populations  at For  in  the  d i f f e r e n c e between identified  apportioning  variation  level  populations.  and  reproducible  25  in  the  results  to  the  this  percent  inter-population  w o r t h remembering when c o n s i d e r i n g  of  the  variation be  chapter.  Analyses. In  broad 29  occurrence  inter-individual  presented  2.2  the  the  this  population  of m o r p h o l o g i c a l that  in  population  are  differentiation  7)  a large  occur  c o n t r i b u t i o n of  the  between  (Table  addressed d i r e c t l y  of  and  examining  geographic Figure  longitude,  and  hypothesized  22  the  r e l a t i o n s h i p s of p a t t e r n s  variation,  the  results  from t h e  were p l o t t e d s e p a r a t e l y  e l e v a t i o n to assess relation.  the  of  against  general  variation  PCAs i n  Table  latitude,  form of  to  the  1 58  Using multiple  means o f e a c h o f t h e o r i g i n a l  linear  v a r i a b l e s per t r e e , a  r e g r e s s i o n of t h e form: (MODEL 4.)  y = elevation + longitude  was p e r f o r m e d a n d r e s i d u a l  + latitude  and p r e d i c t e d v a l u e s  + e.  were c a l c u l a t e d  for  each t r e e f o r each v a r i a b l e .  all  t r e e s e q u a l l y , r e g a r d l e s s of p o p u l a t i o n a l d i s p o s i t i o n .  residual for  and p r e d i c t e d v a l u e s  each v a r i a b l e s u i t e  examined by t h e ANOVA  Such a r e g r e s s i o n  were s u b j e c t e d  considers  to separate  a n d t h e components of t h e s e  The  PCAs  PCAs were  model: (MODEL 5.) y = A + e.  were A i s t h e e f f e c t  of the hypothesized  PCAs upon w h i c h t h i s  ANOVA was b a s e d u s e d a l l t h e a v a i l a b l e  trees, taxa  only  were u s e d  examining et  the scores  al.  f o r the standards  i n t h e ANOVA.  taxa.  and p u t a t i v e s o f t h e  Such an a n a l y s i s h a s been u s e d i n  subspecific variation  i n Pseudotsuga m e n z i e s i i  the hypothesized  anatomically taxa and  (Chen,  1984).  B a s e d on s u c h an a n a l y s i s o f r e s i d u a l if  Although the  variation  e l e v a t i o n should from  standardizes  a r e indeed  d i s c r e t e then  and c l i m a t i c  resulting  taxa  removing  and p r e d i c t e d  m o r p h o l o g i c a l l y and the e f f e c t s  a s s o c i a t e d with  not a f f e c t  the geographical  of a l l o p a t r y of  longitude,  the apportionment  r e c o g n i t i o n of taxa.  values,  T h i s approach  and h i s t o r i c a l  latitude,  of v a r i a t i o n effectively  d i f f e r e n t i a t i o n of  1 59  populations.  Substantial  variation  resulting  variation  as a r e s u l t  suggest  that  separation  recognition  of removing  discrete.  a l o n e might  similar  the r e l a t i o n s h i p  variation  suite  f o r each  correlated  Dugle  tree  f o r each  Elevation  (1969) a n d o t h e r s  of l o c a l  may a f f e c t  t h e time  multiple  linear  regression  m o i s t u r e and e l e v a t i o n that given  these geographic geographic  morphological encountered the broad  on e a c h  s c o r e s were  scores against  Although  emphasized  t o g e t h e r samples  to illustrate  variation  that  treated  variation,  that  As w e l l , a  I t s h o u l d be  simply group  p a t t e r n s of geographic  Singh and  of morphogenesis and  i n P_^ e n g e l m a n n i i .  and e n v i r o n m e n t a l  previously  1968) a s b e i n g  variation.  of i n i t i a t i o n  a r e a and a t t e m p t  each  and r e l a t i v e  (1983) have c o n c l u d e d  was p e r f o r m e d .  i n s u c h an a r e a .  e r e c t e d geographic  a n d von R u d l o f f  of mean component  areas  separate  1974, 1978; L a R o i a n d  geographic  other a s p e c t s of morphogenesis  geographic  h a s been s u g g e s t e d  (1981) a n d H a r r i s o n and Owens  elevation  local  PCA and t h e s c o r e s from  (Falkenhagen  1959; O g i l v i e  t h e major d e t e r m i n a n t Owens  A v e r a g e component  separately against elevation  1968; H o r t o n  geographic  o f t h e 16 a r b i t r a r i l y  6).  moisture a v a i l a b i l i t y . by Roche  with  PCAs were p e r f o r m e d  f o r each  ( T a b l e 7, F i g .  than  that  between p a t t e r n s o f  environmental  PCA  would  suggest.  and a n a t o m i c a l v a r i a t i o n  calculated  of geography  Such a c o n c l u s i o n would s u g g e s t  morphological  areas  of  of taxa and the p a t t e r n of  the e f f e c t s  a r e more g e n e t i c a l l y  In e x a m i n i n g  variable  of t h e a p p o r t i o n m e n t  t h e t a x a a r e more g e o g r a p h i c a l l y t h a n  morphologically individuals  from  re-arrangement  in a  the l o c a l might  be  independently of  this  i s an a n a l y t i c  160  convenience  r a t h e r than  interaction  o f t h e v a r i o u s s c a l e s o f p a t t e r n would  extensive  localized  Comparing  suite,  averaging  regressing The  throughout  were a v a i l a b l e  and n u r s e r y  separate  component  s c o r e s f o r each  ( F i g . 4B).  tree,  As w i t h  standards  o f P_;_ s i t c h e n s i s  environment  environment  variation  should  the nursery  geography,  then  be s t r o n g e r  grown t r e e s .  only  necessary  to r e s t r i c t  i s an i m p o r t a n t  i . e . the n a t u r a l l y  with the grown t r e e s  ( T a b l e 23) s u g g e s t s  that the converse  grown t r e e s would be  strongly correlated  with  geography than the  nursery  grown t r e e s a s a c o n s e q u e n c e o f t h e g r e a t e r  edaphic  variation  To  scores  and, p o t e n t i a l l y ,  to the t r e e s c o l l e c t e d  larger  of s e p a r a t e  suites  local  of p a r e n t s  site.  among t h e p a t t e r n s o f  of v a r i a b l e s ,  f o r each t r e e i n the separate  number  a t any one  examine t h e i n t e r - r e l a t i o n s h i p s  variation  a s p e c t of  The o b s e r v a t i o n o f a l a r g e  may  contributing  naturally  the s c a l e of  f o r the n a t u r a l l y  variation  t o be l e s s  geographic  linear  Using  the c o r r e l a t i o n  intra-population be e x p e c t e d ;  from  of e l e v a t i o n .  associated with  morphological  variable  and l o n g i t u d e .  to multiple  was  was  then  the separate  the e x t e r n a l environment, or a t l e a s t  expected  and  P^ s i t c h e n s i s  a g a i n s t l o n g i t u d e and l a t i t u d e .  the p o s s i b l e e f f e c t s  If  here.  PCAs on e a c h g r o u p f o r e a c h  t h e mean s c o r e s were s u b m i t t e d  occurring  r e q u i r e more  grown P_;_ s i t c h e n s i s  grown m a t e r i a l s r e p r e s e n t e d i t s range  regression  than  than  T e s t i n g f o r the  the scores s e p a r a t e l y against l a t i t u d e  nursery  areas,  sampling  naturally  made by p e r f o r m i n g  a t e s t e d assumption.  average  component  PCAs were c a l c u l a t e d .  This  161  a v e r a g i n g assumes t h a t vegetative  v a r i a b l e s are independent.  suggested  in Figures  vegetative  12 and 13.  and r e p r o d u c t i v e  variation in  i n t r a - t r e e v a r i a t i o n of r e p r o d u c t i v e  suggests that  selection  (Wiley  there  are several  being  1982; Maze  derivation predicted  lineage  traits,  more h i g h l y  constraints  given  These v a r i a b l e  t o be i n d e p e n d e n t .  Based  adjacent  w o u l d be e x p e c t e d  ( M o r i s h i m a and Oka  that  the twig  or derived  v a r i a b l e s measured here  i t would  morphology  then  anatomy.  sequence  i s predicted  and  (1968) d e a l i n g  that  with  (1983) d e a l i n g  and Maze  s u i t e should  be t h e l e a s t  by t h e l e a f m o r p h o l o g y and l e a f on t h e b a s i s  the development  of Owens and M o l d e r  from d i f f e r e n t For the  followed  Owens  s u i t e s of  1983).  inter-related, This  would  or d e v e l o p m e n t a l l y  1968; S c a g e l  s u i t e s of v e g e t a t i v e  predicted  i s , they  t o be  t o be more c l o s e l y i n t e r - c o r r e l a t e d  t h a n t h o s e on more removed p a r t s  be  suites  on t h e common  That  variables  different  process  some s u i t e s o f v a r i a b l e s would be e x p e c t e d  are s p a t i a l l y  the  i t w o u l d be  As a c o n s e q u e n c e ,  apices  or simply  i n Chapter I I ,  show a s i m i l a r d e v e l o p m e n t .  sequential  parallel  1983).  i n t e r - c o r r e l a t e d than o t h e r s .  that  The  a v a r i a t i o n generating  o f some t h e s u i t e s o f v a r i a b l e s , that  has r e s u l t e d  suggests e i t h e r  s u i t e s of v a r i a b l e s .  not a l l expected  independence of  interdependence.  and v e g e t a t i v e  Owing t o t h e s a m p l i n g  are  in this  interdependence  for reproductive  and B r o o k s  from  v a r i a b l e s b a s e d on i n t e r - i n d i v i d u a l  evolution  consequences of e v o l u t i o n  independence i s  Departures  t h e emergence o f d e v e l o p m e n t a l  appearance of such  Such  and  of D o u g l a s - f i r  (1976a) and H a r r i s o n  w i t h development  o f t h e work o f  of the shoots of  needles  a n d Owens sitchensis  1 62  and  Pj_ e n g e l m a n n i i . In a l l a n a l y s e s ,  retained  in order  presented  b a s e d more on than  supporting  chapters.  convention  the  3.1  Geographic  3.1.1  Natural Table  the  these  small. the  largest  23  22  of  the  results  r e c o g n i z i n g the  two  taxa  r e c o g n i t i o n of  illustrate  d e p i c t s the  latitude,  trend  emphasizing and an  low  of  separate  i n the  appearing the  T h e r e was  geographic  variation.  The  with  rather  taxa  in  and  the  are  It should  scores  The  generally  variation. little  variation  i n the of  and  noted  e l e v a t i o n a l d i s c o n t i n u i t y of a b o u t  way  of  a  morphological were elevation -  interior that  for  Latitudinal  t o l o n g i t u d e and  be  from  e l e v a t i o n account  putative hybrids  respect  between a l l  elevation.  figure  c o n t i n u i t y between c o a s t  elevation.  relation  s c a t t e r s of  inter-individual  i n the  the  l o n g i t u d e , and  included  much s m a l l e r .  anatomical  t o be  Figure  source  intermediate  these  Columbia.  values  2  was  non-linear  high  of  results  r e t e n t i o n of  Over a l l v a r i a b l e s , l o n g i t u d e  variation  and  and  PCAs a g a i n s t r  the  variation.  Figure  significant  with  was  variation.  29  trees.  The  of a l l i n d i v i d u a l s  interpretation  continued  southwestern B r i t i s h  Results.  comparison  a l s o a i d s the  the  3.  identification  to- p e r m i t  in earlier  identifications  the  there  as  well  as  appeared  200m between  the  F i g u r e  2 3 .  O r d i n a t i o n s  s e p a r a t e a g a i n s t g i v e n  v a r i a b l e e l e v a t i o n ,  i n  T a b l e  c o m p o n e n t  t h a t  <  49. f r o m  o f  s u i t e s  m e a n s f o r  l a t i t u d e , A p p e n d i x P C A s  i n  o f  f i r s t  s t a n d a r d s , a n d  c o m p o n e n t s p u t a t i v e s .  l o n g i t u d e .  1 1 1 .  G l y p h s  F i g u r e  2 2 .  f r o m  a s ".  i n  P C A  a n d  S c o r e s  F i g u r e  r' v a l u e s  o f  " h y b r i d s "  b a s e d 2 2 .  o n  P C A s F i r s t  s i g n i f i c a n t  0  p  0 . 0 1 .  CONE  MORPHOLOGY  LEAF ANATOMY  TWIG  r  MORPHOLOGY  =0 093*  r' .-OO0B  r' ;007<  TOTAL  9  O  L7ZS.'.  o  f-? c  r  =0577'  r  = 0158*  r ' = 0 573*  1 64  r e p r e s e n t a t i v e s o f P_;_ s i t c h e n s i s P• engelmann i i from co-incidence well  apart  noting  that  Figure  23 e m p h a s i z e s t h e  Mountain  F*\_ e n g e l m a n n i i as  Columbia  P^ s i t c h e n s i s .  t h e P^ e n g e l m a n n i i p o p u l a t i o n s  3° l o n g i t u d e  and 2° l a t i t u d e .  t h e P^ s i t c h e n s i s a r e even  larger:  It  is  a r e 400 km  The d i s t a n c e s 500 t o 1000 km  10 t o 15° l a t i t u d e .  Table for  o f C a s c a d e and S e l k i r k  spanning  separating over  300 t o 500m ASL.  a s t h e Oregon and B r i t i s h  worthwhile  and t h e p u t a t i v e h y b r i d s and  30 g i v e s  the r  the m u l t i p l e l i n e a r  values  2  a s s o c i a t e d with  regresion given  each v a r i a b l e  i n MODEL 4.  As a g r o u p ,  T a b l e 30. r v a l u e s f o r i n d i v i d u a l v a r i a b l e s from m u l t i p l e linearr e g r e s s i o n g i v e n i n MODEL 4. r' v a l u e s e x p r e s s e d a s a t o t a l o f inter-individua1 variation. P r e d i c t e d and r e s i d u a l v a l u e s c a l c u l a t e d on t h e b a s i s of t h e r e g r e s s i o n s . * . r ' v a l u e s s i g n i f i c a n t <a p < 0.01 . !  r. t  VARIABLES NEEDWID NEEDEP ABXANG ADXANG CENCYWID CENCYLAT CENCYABX CENCYADX ENDONUM PHLEND XYLEND X  PULVLEN TIPWID TIPDEP PULVPUB  X  C0NLEN C0NWID SCALEN SCALWID SCALTAP WINGWID WINGTAP FREE SCAL BRACTLEN BRACTWID BRACTAP x  26.60  2 1 25* . 16 . 6 6 * 1 .65 52 . 4 5 * 23.00  x  On t h e a v e r a g e ,  22 .02 21 .04 * 20.90* 13.71* 34.22* 9.44* 16.74* 6.66* 15.60* 54.46* 18.92* 53.27* 22.60  23.56  anatomy v a r i a b l e s a r e t h e most  geography.  1  4 .64* 44 . 4 1 * 33.46* 23.92* 23.73* 1 . 98  NEEDLEN ADXSTOM ABXSTOM RESCYN0 RESCYLOC RESCYLEN  5 . 29* 47 . 6 8 * 35 . 4 0 * 20. 8 9 * 9 . 12* 10. 32* 6 0 . 01 * 36 . 9 9 * 16 . 56* 28 . 73* 21 .6 6 *  TOTAL  leaf  r  VARIABLES  strongly associated  with  vegetative v a r i a b l e s are only  1 65  slightly  more c o r r e l a t e d w i t h  reproductive. correlated  (i.e. r  linear  2  > 50%) w i t h  the geographic  PULVPUB, BRACTLEN, AND  were r e l a t e d s i g n i f i c a n t l y  variables.  than a r e the  The v a r i a b l e s w h i c h a r e t h e most  NEEDEP, CENCYABX, variables  the geography  i s 24 p e r c e n t  variables are:  BRACTAP.  Nearly a l l  t o the geographic  Over a l l v a r i a b l e s , t h e a v e r a g e  regression  strongly  r  of the t o t a l  for this  2  multiple  inter-individual  variation.  Table residual  31 g i v e s  t h e r e s u l t s o f t h e PCAs b a s e d  and p r e d i c t e d  regressions  given  values  i n MODEL 4.  from t h e m u l t i p l e  on t h e  linear  The amount of v a r i a t i o n a c c o u n t e d  T a b l e 3 1 . M u l t i v a r i a t e a p p o r t i o n m e n t of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s b a s e d on p r e d i c t e d and r e s i d u a l v a l u e s from m u l t i p l e l i n e a r r e g r e s s i o n (MODEL 4 ) . O r i g i n a l v a l u e s b a s e d on PCAs a n d ANOVAs g i v e n i n T a b l e 2 9 . A b b r e v i a t i o n o f PCAs a n d ANOVAs g i v e n i n T a b l e 4 9 a n d 5 0 , Appendix I I I . O r d i n a t i o n s of r e s u l t a n t component s c o r e s g i v e n i n F i g u r e s 2 2 . 2 3 , and 2 4 .  VARIABLE  for  SUITE  ORIGINAL  %SSA (mva) PREDICTED  LEAF ANATOMY LEAF MORPHOLOGY TWIG MORPHOLOGY CONE MORPHOLOGY TOTAL  35.70 25.37 22.20 24.98 27.06  7 1.59 59.26 63.40 53.22 6 1.87  5.22 2.69 1.45 1.44 2.70  x VEGETATIVE  27.76  64 75  3.12  by t h e PCAs and t h e component  compared w i t h predicted relation values, should  RESIDUAL  those given  values among  these  in Table  c o r r e l a t i o n s should 29.  were t h e most p o l a r .  the t r e e s should  be e m p h a s i z e d  be  The PCAs b a s e d on t h e Figure  24 i l l u s t r a t e s t h e  b a s e d on t h e r e s i d u a l and p r e d i c t e d  be compared w i t h t h e PCA b a s e d  those  in Figure  on p r e d i c t e d  scores  22.  It  from t h e  1 66  multiple total  linear  r e g r e s s i o n s only accounts  variation  i n the  effect  l a r g e p o r t i o n of based  on  residuals  polarity.  As  variation from the  ( F i g . 24)  by  the a n a l y s i s  as  the a n a l y s i s  predicted  by  31)  of  of  the  variation;  emphasizes  this  the d e m o n s t r a t i o n  residual  values,  caused  by  taxon  the  that  removes a  the o r d i n a t i o n  l a c k of of  taxonomic  the d e c l i n e i n taxa  results  i s the d e m o n s t r a t i o n  of  recognition resulting  of p r e d i c t e d v a l u e s .  v a l u e s a l s o emphasize  indicate  regression also  r e c o g n i t i o n of h y p o t h e s i z e d  increased variation  from  ( T a b l e s 29 and  inter-taxonomic  important  caused  scores  of g e o g r a p h y  the  percent  data.  ANOVAs of component removing the  f o r 24  The  o r d i n a t i o n s of  i n t e r m e d i a t e appearance  of  the p u t a t i v e h y b r i d s .  B a s e d on variables  the  predicted values  from  LOCOWID) d i s p l a y e d an  that  inter-taxonomic  exceeded largest  (uva)  r  values  2  attributed  predicted  The  %SS^(uva) a t t r i b u t e d polarity  of  the  in Table  based  based  A  29  on  are 31  those  with  for residual  predicted values  of  taxon  the  emphasizes  on  values  Table  32  of  the d a t a  based  e m p h a s i z e s TIPDEP, TIPWID, CENCYWID, and  presents  the c o r r e l a t i o n  between  the  largest  values.  CENCYADX.  c o n t r a s t the p o l a r i t y  %SS^  on  and  By  with  largest  % S S ( u v a ) as a r e s u l t  in Table  to taxa  the data  g e n e r a l l y had  BRACTLEN, BRACTAP, ABXANG, ADXSTOM, CENCYABX, NEEDEP,  residual  8  variation  Those v a r i a b l e s  r e c o g n i t i o n ( T a b l e 31)  largest  for variables  inter-taxonomic  variation.  ( T a b l e 30)  t o taxon  values.  recognition  The  r e g r e s s i o n , only  (NEEDWID, NEEDLEN, TIPDEP, CONWID, SCALEN, SCALTAP,  FREESCAL, and  the  the  SCALEN.  PCAs based  on  1 67  F i g u r e  2 4 .  v a r 1 a b 1 e  O r d i n a t i o n s s u 1 t e s  f  o  p r e d i c t e d  r e g r e s s i o n  i n M O D E L  g i v e n  i  LEAF  n T a b l e  5 0 ,  o  f f i r s t  t w o c o m p o n e n t s  rs t a n d a r d s .  c a l c u l a t e d  p u t a t  a n d r e s i d u a l 4 .  G l y p h s  A p p e n d i x  a  s i  I I I .  ANATOMY  f  8  LEAF MORPHOLOGY  CONE COLLECTION predicted  residual  TWIG MORPHOLOGY  <2>  TOTAL  O O  iv e s .  v a l u e s  f r o m  P C A s  a n d " h y b r f r o m  nF i g u r e  id s "  m u l t i p l e  2 2 .  S c o r e s  o  f  s e p a r a t e  u s in g l i n e a r b a s e d  o  n P C A S  1 68  original,  p r e d i c t e d , and r e s i d u a l  each v a r i a b l e s u i t e .  values  In g e n e r a l ,  (Table  the o r i g i n a l  29 and 31) f o r data  a r e most  T a b l e 32. C o r r e l a t i o n s among PCAs o f o r i g i n a l , p r e d i c t e d , a n d r e s i d u a l values. *, c o r r e l a t i o n s s i g n i f i c a n t ® p < 0.01. PCAs o f p r e d i c t e d and r e s i d u a l v a r i a t i o n from T a b l e 30. PCAs of f o r o r i g i n a l , p r e d i c t e d , a n d r e s i d u a l v a l u e s g i v e n i n T a b l e 49 a n d 5 0 , A p p e n d i x III. - c o r r e l a t i o n m i s s i n g as e i g e n v a l u e l e s s t h a n 1.0 f o r the component i n q u e s t i o n .  PREDICTED II  RESIDUAL 11  III  111  ORIGINAL CONE I 11 III LEAF  ANATOMY I II  TWIG MORPHOLOGY I II LEAF  MORPHOLOGY I II III  .111 .669* . 107  .095  . 367" .882"  .512* . 328  -.121 -.081  .912* .361*  . 194 .653*  .045 -.003 .229*  . 738" . 547* . 236*  .177* . 267* .026  . 034 .316* .915*  .039 .531* .036  . 990* .018 . 043  .064 .010 .969'  .112 . 292* .018  .808''  .317* -.659*  .627* .454* . 158*  .005 . 159" . 296*  TOTAL I II I 11  . 843* . 137 - . 335*  strongly  c o r r e l a t e d with  residual  data.  are  .028 . 142 .154  .058 .069 .02 1  the r e s u l t s  By d e f i n i t i o n  of the a n a l y s i s of the  t h e p r e d i c t e d and r e s i d u a l  scores  uncorrelated.  Table of  .018 .009 .939  .821* . 400* .002  .554* .449* .049  33 p r e s e n t s  the i n t e r - c o r r e l a t i o n s  the v a r i o u s v a r i a b l e s u i t e s .  Figure  among t h e a n a l y s e s  25 d e p i c t s t h e a n g u l a r  co-incidence  of the v e c t o r s of the v a r i o u s v a r i a b l e s u i t e s  plotted  the f i r s t  into  two components f r o m t h e PCA g i v e n i n  169  T a b l e 33. I n t e r c o r r e 1 a t i o n s o f v a r i a b l e s u i t e a n a l y s e s b a s e d on PCAs o f r e s u l t s of r e g r e s s i o n a n a l y s i s . PCAs g i v e n i n T a b l e 49 a n d T a b l e 5 0 . A p p e n d i x III. Inter-correlations b a s e d on s c o r e s f r o m f i r s t component. * , c o r r e l a t i o n s i g n f i c a n t ®> p < 0 . 0 1 .  ORIGINAL TOTAL REPRODUCTIVE LEAF ANATOMY TWIG MORPHOLOGY LEAF MORPHOLOGY  .764* -.921* .238* -.745*  . 56 1 * .293* . 485*  .018 .636*  .25 V  PREDICTED TOTAL REPRODUCTIVE LEAF ANATOMY TWIG MORPHOLOGY LEAF MORPHOLOGY  .970* -.999* .979* -.964*  .959* .902* .875*  . 987* .973*  .997"  RESIDUAL TOTAL REPRODUCTIVE LEAF ANATOMY TWIG MORPHOLOGY LEAF MORPHOLOGY  Tables  variable  on l e a f  pattern  values  the p a t t e r n  anatomy most c l o s e l y  variable  anatomy  inter-correlation i s about  on p r e d i c t e d  suite  and l e a f  inter-correlated.  variables  .685* - . 126  on o r i g i n a l o r  of v a r i a t i o n  morphology  suites  on o r i g i n a l v a l u e s ,  samples  the t o t a l i s the l e a s t  with the t o t a l p a t t e r n variable  of  of  variation.  a r e t h e most  the average  between v e g e t a t i v e and r e p r o d u c t i v e 21 p e r c e n t .  variation  A contrast  Based  approximates  Twig  morphology  Based  - . 149  22 and 24.  from a l l 36 v a r i a b l e s .  correlated Leaf  . 065* . 199' . 163  29 and 31 and F i g u r e  residual based  .679* .729* .730* -.213*  Predictably,  are highly  a l l patterns  correlated.  t o t h e s e p a t t e r n s of v a r i a b l e  suites  suite  of  based  170  F i g u r e 25. Average v e c t o r s of s e p a r a t e v a r i a b l e s u i t e s p l o t t e d Into c o - o r d i n a t e s o f f i r s t two a x e s f r o m PCAs o f a l l i n d i v i d u a l trees b a s e d on o r i g i n a l and p r e d i c t e d and r e s i d u a l v a l u e s from m u l t i p l e regression. PCAs g i v e n i n T a b l e s 49 a n d 5 0 . A p p e n d i x III. C o m p o n e n t s i l l u s t r a t e d p r e v i o u s l y 1n F i g u r e s 22 a n d 24. C - cone m o r p h o l o g y : T - t w i g morphology: A - l e a f anatomy; M - l e a f morphology.  ORIGINAL  PREDICTED  intercorrelations  i s seen  intercorrelations  based  the  are  intercorrelations  those based  suites  suggesting  Figure  values  the other  Residual  values.  are c o r r e l a t e d  of v a r i a b l e  patterns  suite  of v a r i a t i o n  patterns  are generally less  of  from  variable  inter-correlated  Vegetative  and  by an a v e r a g e  r  2  than  reproductive of about  2  independence.  25 c o r r o b o r a t e s  reproductive  original and  on r e s i d u a l  on o r i g i n a l  of v a r i a b l e s  percent,  and  i n the p a t t e r n s  r e g r e s s i o n on g e o g r a p h y .  suite  RESIDUAL  variable emphasize  the d i f f e r e n c e s  suites.  vegetative  PCAs o f p r e d i c t e d and  the d i s p a r i t y  vegetative variable  between  between t w i g  suites.  morphology  171  3.1.2  Nursery  Table naturally  and n a t u r a l  variation.  34 g i v e s  the results  of the separate  occurring  and nursery  grown  Correlations  of eigenvector-values  of  from  variation  relationships different.  these  between  Over  PCAs  of  s itchens i s.  indicated that  were  samples  trees  PCAs o f  highly  correlated yet  t o the geographic  a l lv a r i a b l e s , natural  the patterns the  scale are  and nursery  PCAs  have  T a b l e 34. M u l t i v a r i a t e r e l a t i o n s h i p o f v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s f o r n u r s e r y a n d n a t u r a l l y grown m a t e r i a l s t o g e o g r a p h i c variation. A b b r e v i a t i o n o f PCAs a n d r e g r e s s i o n s i n T a b l e 51, Appendix I I I . r values g i v e n f o r m u l t i p l e l i n e a r r e g r e s s i o n against l o n g i t u d e and l a t i t u d e . r! values expressed as a percentage of the total inter-individua1 variation. O r d i n a t i o n s o f r e s u l t a n t component s c o r e s g i v e n i n F i g u r e 26. !  (mva)  VARIABLE  SUITE  NURSERY  LEAF ANATOMY LEAF MORPHOLOGY TWIG MORPHOLOGY VEGETATIVE  similar  polarities. of these  morphology  a n d anatomy  strongly  PCAs  occurring  were  collected  more  relationship collections  with  against  trees.  longitude  Although  f o r even  trees  location  was  were t h e from t h e  the naturally  t h e same those  The  more  than  the collections  i s apparently  i n morphological  of the f i r s t  and l a t i t u d e .  grown  extensive,  d i dnot display  There  . 56 . 52 . 52 20  ordinations  the geographic  geography  overlap.  discontinuity  26 g i v e s  latitudinally  materials  25 18 25 23  of the nursery  c o r r e l a t e d with  naturally nursery  4 . 58 8 . 10 10. 94 7 . 87  Figure  components  NATURAL  pronounced  areas no  or anatomical  where  t h e two  latitudinal variation  in either  F i g u r e  2 6 .  O r d i n a t i o n s  t r e e s  f r o m  g r o w n  t r e e s  1 a t i t u d e .  P C A s r  1  LEAF  ir s t  f  o  c o m p o n e n t  f m e a n s  s e p a r a t e a  v a l u e s  i n t e r - i n d i v i d u a 1 F  o  p l o t t e d  n  o  e x p r e s s e d  v a r i a t i o n . f r o m  f c o m p o n e n t s  v a r i a b l e  dr e g r e s s e d  P C A s  i  a *,  s a  r*  n T a b l e  v a l u e s  oco  f o r  i n d i v i d u a l  f n a t u r a l l y o  f  t  a  n  d  n u r s e r y  l o n g 1 t u d e  a g a i n s t  p e r c e n t a g e  LEAF  NUfiSERY  o  h  s i g n i f i c a n t  a  n  e t o t a l •  p  <  0 - 0 1 .  3 - 4 .  ANATOMT  NATURAL  s c o r e s  s u i t e s  s e p a r a t e l y  ANATOMY  .  NATURAL  O  OCCP  o o CD CO CD g> O o o o o CO  (DO  Sb  0° o  I E A F MOPFXOLOGT  o  LEAF  o  oo9  CJDO  MOBPMXOGY 0 CO  OCO  3)  3b 39ccD  o o ao CD  at  o  oar3> o  CCC  TWIG TwG '  MOCPHOLOGT  MGBP-XXCGY  ooo  a>  coco as o D go o o op>  0  CCD OO,^  O  o  o  3ro  o co  <9<bo oco o  CP  tr' =.0J! "  o CCD  O  CO O  OO  OO  O CD  OO  oc  cm ffboaP  tfcaiiasf o 10  3D O  OO  d  1  natural  or nursery m a t e r i a l s .  3.2 L o c a l g e o g r a p h i c  Table suites  73  variation.  35 g i v e s t h e r e s u l t s  f o r t h e 16 s e p a r a t e  o f t h e PCAs o f s e p a r a t e  geographic  areas.  variable  Also given in  T a b l e 35. M u l t i v a r i a t e r e l a t i o n s h i p of v a r i a t i o n f o r s e p a r a t e v a r i a b l e s u i t e s f o r s e p a r a t e g e o g r a p h i c a r e a s r e l a t e d to environment. A b b r e v i a t i o n o f PCAs and r e g r e s s i o n s i n T a b l e 52, A p p e n d i x III. r' v a l u e s g i v e n f o r m u l t i p l e l i n e a r r e g r e s s i o n a g a i n s t e l e v a t i o n and m o i s t u r e . Those g e o g r a p h i c areas without r v a l u e s c o u l d not be a n a l y z e d due to an i n a d e q u a t e sample s i z e . r' v a l u e s e x p r e s s e d as a p e r c e n t a g e of the t o t a l i n t e r - i n d i v i d u a 1 variation. O r d i n a t i o n s of r e s u l t a n t component s c o r e s g i v e n i n F i g u r e 27. !  r' GEOGRAPHIC AREA 1 2 3 4 5 6 7 8 9 10 1 1 12 13 14 15 16  Table  LEAF ANATOMY  LEAF MORPHOLOGY  TWIG MORPHOLOGY  1 1.27 32 . 17  44 . 12 18 . 27  29 . 93 10 . 37  34 .97  29 . 69  20 . 69  32 . 25  35 ..95  32 .00  14 20 16 32 14 19 43  14 24 5 14 19 36 54  25 24 9 17 14 36 25  . 64 . 47 .02 . 12 . 59 . 16 .01  35 a r e t h e r  each geographic moisture. moisture  2  area  As w e l l ,  CONE MORPHOLOGY 19 13 18 33 24 51 7 36 33 37 13 14 17 14 37 42  .93 .1 1 .09 .01 . 44 . 44 . 20  with  separate  x VEGETATIVE  x TOTAL  28 . 44 20 . 77  26 . 24 18 . 46  28 .45  34 . 25  33 ..40  34 ..09  18 . 48 23 . 10 10 .07 21 23 16 .02 30 .06 40. 87  23 . 18 20 . 7 1 1 1. 13 20 .31 15 .69 32 25 4 1 16 .  24 ..64  24 . 83  .63 .02 .41 .02 .84 . 58 . 38 . 16 .88 . 29 .54 .31 .54 .69 . 18 .03  25 . 4 1  23 . 75  v a l u e s o f t h e means  o f component s c o r e s f o r  r e g r e s s i o n s on e l e v a t i o n and  m u l t i p l e r e g r e s s i o n s u s i n g e l e v a t i o n and  are expressed  components a r e g i v e n elevation  86 . 73 . 10 . 55 .04 . . 20 . 40  26 . 26  23 .88  X  (mva)  or r e l a t i v e  as an r  as t h e s e  2  (mva).  Only  the  first  had t h e h i g h e s t c o r r e l a t i o n s  moisture.  with  174  The  PCAs of  relationship anatomical these 6),  r  between  variation  values are  2  they  are  16)  and  separate  geographic  t h e p a t t e r n of and  local  significant  correlated  correlated  morphology.  Over a l l v a r i a b l e s  r  value  2  moisture  the  of t h e  size  i s about  average  geographic  variation  geographic  areas  regression  of e l e v a t i o n  total is  variables  than  physiographic correlation  PCAs and degrees this  are  based  elevation,  on  2  value  moisture  local  areas  the  and  25  to these  leaf  that for half  scale and linear  percent  In g e n e r a l , cone  of  the  morphology  environmental  The  approximatly  correlations,  some v a r i a b l e s consistently  T h e s e were t h e from  l o n g i t u d e , and  on  three the  broad  same  environment.  i n e a c h of  residuals  based  f o r the m u l t i p l e i s about  ( F i g . 6) a l l had  (Table  Elevation is  Over a l l v a r i a b l e s  variation.  component  variability  f o r the broader  amount of v a r i a t i o n  of v a r i a t i o n  each area.  data  the  the  TIPDEP) a c c o u n t in  and  4).  13 p e r c e n t  the v e g e t a t i v e v a r i a b l e s .  areas  their  r  35:  These v a l u e s are  more s t r o n g l y r e l a t e d  with  B a s e d on  value  2  the average  slightly  percent.  ( F i g . 23).  inter-individual  only  r  11  Table  f o r MICA  a l l geographic  i s about  Where  i s only  p a t t e r n s of v a r i a t i o n  for elevation  relative  Inlet,  ( T a b l e 35:  and  and  variation.  twig morphology  c o n e m o r p h o l o g y a t TOBA  little  morphological  (e.g. Knight  particularly  average  local  indicates  R e l a t i v e moisture  with  with  areas  environmental  generally small.  significantly 35:  the  the  e x t r a c t e d by  the  these  t h e r e were v a r y i n g  separate  areas.  In s p i t e  of  ( i . e . SCALEN, CENCYWID, TIPWID,  f o r t h e major p a t t e r n o f same v a r i a b l e s multiple linear  latitude  (MODEL 4;  variation  that polarized  the  r e g r e s s i o n on Table  35).  There  175  was  no  single  pattern  of  Similarly  local  leaf  MICA  that accounted  geographic  t h e r e was  consistently (i.e.  variable  no  single  more v a r i a b l e  (Table  35:  morphology).  16)  these  27  with  of  The  single  to  be  geographic  area  local  major  morphology.  areas  t h e most v a r i a b l e  the  the  uniform  was  location  for  consistently  environment  respect to p o s i t i o n  over  a l l geographic  e n g e l m a n n i i and  ordinations, discrete  and  areas, P^  however  the  than  hybrids  ( T a b l e 35:  little  polarity  the presence  of  Averaging presented scales  two  and  in Tables  of v a r i a t i o n  of v a r i a t i o n assumption  of  any  areas  t o be  compared  There  is a  tendancy,  r e p r e s e n t a t i v e s of  geographic  taxa. both  area  In t h o s e  p o l e s of there  t a x a and. t h e i r  the  i s no  geographic  areas  putative  ( 8 ) , HOWHIS ( 2 ) , KNIGHT  (6)), there  that