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
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Morphological and anatomical variation of picea in southwestern British Columbia Scagel, Robert Kevin 1984
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Title | Morphological and anatomical variation of picea in southwestern British Columbia |
Creator |
Scagel, Robert Kevin |
Publisher | University of British Columbia |
Date Issued | 1984 |
Description | Relations between individual trees hypothesized to be representative of Picea enqelmannii and P. sitchensis and putative hybrids in southwestern British Columbia were examined using several morphological and anatomical variables measured on cones, needles, and twigs. 36 variables were selected that had an inter-individual variation that exceeded intra-individual variation. Such relative variation suggested that the variables selected were more genetically than developmentally variable. These variables were measured on 640 trees. Patterns of inter- and intra-individual variation of these variables were examined with multivariate analyses. Similar patterns of intra-individual variation with respect to position in the canopy were detected regardless of the age of the tree or the species, suggesting that intra-individual variation in Picea is not necessarily a function of the environment extrinsic to the individual. Further, the two hypothesized taxa were not especially different with respect to the pattern of intra-individual variation. In an inter-individual context, the intra-taxonomic variation was shown to be larger than inter-taxonomic variation. Inter-individual variation attributed to the hypothesized taxa was smaller than intra-population variation. Inter-taxonomic variation was only 25 percent larger than intra-individual variation. Further, the hypothesized taxonomic polarity of the data was not co-incident with the actual polarity of the data. What taxonomic structure emerged was evident only as the extremes of a continuum of variation over a large geographic area reflecting large environmental variation, suggesting that recognition of two taxa of Picea in southwestern British Columbia may not be appropriate or practically feasible. From the evidence available here, it is not possible to attribute the pattern of variation to hybridization followed by introgression or to the differentiation of a single, large polymorphic taxon. The pattern of inter-individual variation was only partially correlated with regional and local scales of geographic and environmental variation. Local patterns of variation were not co-incident in 16 separate geographic areas. The relation of the pattern of variation in nursery grown trees with respect to regional scales of geographic variation was not co-incident with the patterns of variation in naturally grown trees from the same area of origin. This variability of relations suggests that inter-individual variation is probably attributable to processes operating at an extremely local scale. Implications from these conclusions are presented and discussed with respect to the systematics of Picea and the utilization of Picea in forestry and tree improvement programmes. |
Genre |
Thesis/Dissertation |
Type |
Text |
Language | eng |
Date Available | 2010-05-16 |
Provider | Vancouver : University of British Columbia Library |
Rights | For non-commercial purposes only, such as research, private study and education. Additional conditions apply, see Terms of Use https://open.library.ubc.ca/terms_of_use. |
DOI | 10.14288/1.0096121 |
URI | http://hdl.handle.net/2429/24766 |
Degree |
Master of Science - MSc |
Program |
Botany |
Affiliation |
Science, Faculty of Botany, Department of |
Degree Grantor | University of British Columbia |
Campus |
UBCV |
Scholarly Level | Graduate |
Aggregated Source Repository | DSpace |
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