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Geographic variation in picea glauca in British Columbia Roche, Laurence 1967

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The University of British Columbia FACULTY OF GRADUATE STUDIES PROGRAMME OF THE FINAL ORAL EXAMINATION FOR THE DEGREE OF DOCTOR OF PHILOSOPHY of LAURENCE ROCHE B .Agr . (Forest^.) J B.A., University of Dublin, Trinity College, 1960. M.F. University of British Columbia, 1962. MONDAY, DECEMBER 18, 1967 AT 2:00 P.M. ROOM 256, MacMILLAN BUILDING COMMITTEE IN CHARGE Chairman: I. McT. Cowan V.C. Brink G.G.E. Scudder P.G. Haddock 0. Sziklai D.P. Ormord T.M.C. Taylor External Examiner: Dr. H. Nienstaedt Institute of Forest Genetics Rhlnelandfec, jWi's'ooastnJUS S ...A, Research Supervisor: 0. Sziklai GEOGRAPHIC VARIATION IN PICEA GLAUCA IN BRITISH COLUMBIA ABSTRACT The study is divided into two parts, one of which is a genecological investigation of 150 spruce popula-tions grown from seed in a relatively uniform environ-ment during a period of two years. The second part is an investigation of geographic variation in mature popula-tions of spruce, and refers principally to a biometrical investigation in cone scale morphology which was carried out on a mass collection of spruce cones collected in 157 areas throughout the range of spruce in British Columbia. On the basis of the results obtained in both parts, zones of putative hybridization between white spruce and the other spruce species of British Columbia are demar-cated, and the following general conclusions are made. In regard to the white-Engelmann spruce complex in British Columbia, the environmental pressures which resul in microevolution, i.e. speciation. The faculty for normal development and survival of white spruce and its related forms in British Columbia, is conditioned by the cessation of; growth "and i n i t i a t i o n of dormancy. The genetic constitution of a natural population of white spruce, in any one region, is predominantly determined by the photothermal regime prevailing in that region. On the basis of these general conclusions, recom-mendations are made in regard to the silviculture of white spruce and its related forms in British Columbia, and also in regard to the f i e l d testing of the 150 spruce populations subjected to genecological investigation in the nursery. GRADUATE STUDIES Field of Study: Forest Genetics Advanced Plant Genetics: V.C. Brink Responses of Plants to Controlled D.P. Ormrod Environments ''. \ \v. Organic Evolution G.G„ Scudder Problems in Forest. Genetics 0 . Sziklai Forest Tree Seed 0 . Sziklai Taxonomy of Vascular Plants T.M.C. Taylor PUBLICATIONS Roche, L. The Lulu Island provenance of Pinus contorta. Irish Forestry, 18:50-56 (1962). Roche, L. The shore variety of Pinus contorta. Baileya 11: 11-14 (1963). Roche, L. Variation in lodgepole pine with reference to provenances planted in Great Britain and Ireland, Forestry 39: 30-39 (1966). Roche, L. Spruce provenance research in British Columbia Proceedings of the tenth meeting of the committee on Forest Tree Breeding in Canada. Part II, 107-121 (1967). Roche L. The value of short term studies in provenance research. Commonwealth Forestry Review, in press (1968). GEOGRAPHIC VARIATION IN PICEA GLAUCA IN BRITISH COLUMBIA by LAURENCE ROCHE B. Agr. (Forest), MA (Dublin) I960 M.F. University, of British Columbia, 1962 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF . DOCTOR OF PHILOSOPHY in the Department of FORESTRY We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA March, 1967 In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r an a d v a n c e d d e g r e e a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and S t u d y . | f u r t h e r a g r e e t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by t h e Head o f my Depar tment o r by h.i;s r e p r e s e n t a t i v e s . It i s u n d e r s t o o d t h a t c o p y i n g .or p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . D e p a r t m e n t o f p^it-tt<y  The U n i v e r s i t y o f B r i t i s h C o l u m b i a V a n c o u v e r 8, Canada Date / r ,/U7  ABSTRACT I The principal objective of the study is the determination of geographic variation in rtiite spruce in British Columbia. Since variation within this species in British Columbia is greatly influenced by hybridization with other spruce species, an attempt is made to demarcate zones of hybridization, and evaluate its effect on variation in white spruce. In a preliminary chapter the literature pertaining to principles and concepts of taxonomic and genecological investigation is critically examined in relation to infraspecific variation in tree species. The conclusions of this chapter constitute the assumptions of the investigation. A second chapter summaries the literature pertaining to the phylogeny and distribution of the spruce species of British Columbia, Photoperiodicity in forest trees is discussed in the third chapter. Following the chapters referred to above the study is divided into two parts, A and B. Part A is a study of the growth behaviour of 13>0 popula-tions of spruce grown in a relatively uniform environment during a period of two years. The seed, which was collected throughout the spruce complex of British Columbia,was sown at the British Columbia Forest Service research nursery on Vancouver Island in the spring of 1°65. Detailed measurements were made during the growing seasons of 1965 and 1966. In the laboratory seed samples of the same populations were X-rayed to determine embryo develop-ment and subsequently germinated at 25°C. Further seed samples were germinated at 15, 20, and 30°C. Part B is a study of geographic variation in mature populations of white spruce, and refers principally to a biometrical investigation of variation in cone scale morphology which was carried out on a mass collection I I of spruce cones c o l l e c t e d i n 157 areas throughout the range of spruce i n B r i t i s h Columbia during the summers of 1963 and 1964. On the basis of the r e s u l t s obtained i n parts A and B the following general conclusions are madet ( i ) In regard t o the white-Engalmann spruce complex i n B r i t i s h Columbia the environmental pressures which r e s u l t i n microevolution, i . e . i n f r a s p e o i f i o v a r i a t i o n , d i f f e r only i n degree rather than i n kind from the environmental pressures which r e s u l t i n macroevolution, i . e . s p e c i a t i o n . ( i i ) The f a c u l t y f o r normal development and s u r v i v a l of white spruce, and i t s r e l a t e d forms, i s conditioned by the cessation of growth and i n i -t i a t i o n of dormancy. ( i i i ) Time of i n i t i a t i o n of dormancy i n a population i n any one region where the species occurs n a t u r a l l y i s conditioned by i t s genetic c o n s t i t u t i o n . ( i v ) The genetic c o n s t i t u t i o n of a n a t u r a l population i s predominantly determined by the photothermal regime p r e v a i l i n g i n that region. (v) In so f a r as there i s a difference i n the photothermal regime between any two regions the genetic c o n s t i t u t i o n of the spruce populations occupying,; those regions w i l l d i f f e r . ( v i ) One of the most important external manifestations of t h i s d i f f e r e n c e i s the time of cessation of growth and i n i t i a t i o n of domancy. On the basis of these general conclusions, recommendations are made i n regard to the s i l v i c u l t u r e of white spruce and i t s r e l a t e d forms i n B r i t i s h Columbia, and a l s o i n regard t o the f i e l d t e s t i n g of the spruce populations r e f e r r e d t o i n part A of t h i s study. GEOGRAPHIC VARIATION IN PICEA GLAUCA IN: BRITISH COLUMBIA BT LAURENCE ROCHE I l l ACKNOWLEDGEMENT The writer is primarily and especially indebted to Dr. Oscar Sziklai of the Faculty of Forestry who is chief adviser and chairman of the examining committee, and who, since January 1961, has been his mentor in a l l matters pertaining to forest genetics. Acknowledgement is also due to Dr. P.G. Haddock of the Faculty of Forestry, Dr. V.C. Brink and Dr. D.P. Ormrod of the Division of Plant Science, Faculty of Agriculture, Dr. G.G.E. Scudder of the Department of Zoology, and Dr. T.M.C. Taylor of the Department of Botany, a l l of whom directly contributed to the development of the ideas presented in this study. Special acknowled-gement is given to Dr. A. Kozak of the Faculty of Forestry who gave major assis-tance with respect to statistical analysis, and also to Dr. J.H.G. Smith. In regard to the distribution of the spruce species in British Colum-bia, the writer, in numerous discussions, availed of the long experience of Dr. B.G. Griffith of the Faculty of Forestry, and Mr. E. Garman of the British Columbia Forest Service (now retired). To both these men the writer extends his thanks. The writer wishes to thank the University of British Columbia which, through the Faculty of Forestry, provided substantial financial assistance during the academic year 1965-66, Without this financial assistance i t would not have been possible to undertake post-graduate training. This study could not have been undertaken without close cooperation between the Faculty of Forestry of the University of British Columbia and the Research Division of the British Columbia Forest Service which, apart from the academic studies of the writer, financed the entire investigation. The writer, therefore, wishes to acknowledge his indebtdness to,the British Columbia Forest Service, and to its personnel in general, but particularly to those of his colleagues,, who by direct assistance in statistical matters, or by 17 constructive criticism contributed to the investigation. Therefore acknowledgement is particularly due to Mr. R.H. Spilsbury, head of the Research Division of the B.C. Forest Service, who fully supported the writer's relatively untested ideas in regard to provenance research, and provided the financial assistance for their implementation. Acknowledgement is also due to Mr. G»C. Warrack, who supervised the technical aspects of the work during the writer's absence, to Mr. A. Bamford who provided the ex-perimental material for Part A of the study, and to Mr. J. Revel who provided miscellaneous cone collections for part B, The writer is especially grateful to Mr.^ M. Kovats and Mr. A. Frazer for constant assistance in a l l aspects of statistical analysis. Special thanks is also extended to Mr. R. Bowden-Green who provided technical assistance throughout the course of this study, to Miss S. Lindley for her assistance in the laboratory, and to Miss E, Lemon for her unfailing help and good will over a period of k years in a l l matters pertaining to bibliography. Miscellaneous cone collectionsfor part B of the study were also pro-vided by Mr. H. Bunce of Columbia Cellulose Co. Ltd., who, together with Mr, W.H. VanHeek, assisted the writer in sampling in the Nass and Skeena river areas of the province. Finally, the writer wishes to acknowledge the indispensable help and encouragement provided by Dr. L. Daviault, Regional Director of the Federal Forestry Research laboratory at Quebec City during the winter of I966-I967 when'this thesis was being written. To a l l members of this laboratory who provided technical; assistance in the preparation of the manuscript, especially Miss E. Poisson, Miss- L. Lepage, Mr. R. Keable, Mr. R. Gagnon, and Mr. R. Charuest, the writer extends his thanks. V TABLE OF CONTENTS Page ABSTRACT 1 ACKNOWLEDGEMENT I l l TABLE OF CONTENTS V LIST OF TABLES VII LIST OF FIGURES IX LIST OF ILLUSTRATIONS... XII FRONTISPIECE XIII Caption for frontispiece. XIV INTRODUCTION 1 SYSTEMATICS IN RELATION TO GEOGRAPHIC VARIATION IN FOREST TREES: A STATEMENT OF ASSUMPTIONS. AND OBJECTIVES OF THE PRESENT STUDY...... 3 DISTRIBUTION AND PHYLOGENETIC RELATIONSHIPS OF THE SPRUCE SPECIES OF BRITISH COLUMBIA 17 PHOTOPERIODICITY IN FOREST TREES: A LITERATURE REVIEW" 23 MATERIALS AND METHODS 28 RESULTS PART A: GEOGRAPHIC VARIATION IN BMATURE POPULATIONS OF WHITE. SPRUCE ........ 51 DISCUSSION 79 The ecological significance of photoperiodic adaptation in white spruce...... •• 83 The relative importance of flushing and dormancy in the microevolution of white spruce 89 Germination behaviour in the laboratory. 95 RESULTS PART B: GEOGRAPHIC VARIATION IN MATURE POPULATIONS OF WHITE. SPRUCE 99 DISCUSSION • 1 0 6 The pattern of variation in the white-Engelmann spruce complexes The pattern of variation in the white-black spruce complex.. The pattern of variation in the white-Sitka spruce complex.. US' VI THE RELATIONSHIP BETWEEN PATTERNS OF VARIATION IN IMMATURE AND MATURE SPRUCE POPULATIONS 121 THE TAXONOMIC SIGNIFICANCE OF GEOGRAPHIC VARIATION IN WHITE SPRUCE IN BRITISH COLUMBIA 125 THE SILVTCULTURAL SIGNIFICANCE OF GEOGRAPHIC VARIATION IN WHITE SPRUCE. IN BRITISH COLUMBIA . . ... 13U GENERAL CONCLUSIONS 1U9 LITERATURE CITED 157 APPENDIX: FIGS. 9 to 1*6 170 LIST OF TABLES 1 Embryo and endosperm classes corresponding to those shown in Illus. 2 31 2 Measurements of germination and growth made on 150 spruce provenances 3 7 3 Geographic origin of 150 spruce provenances..... 38 U Geographic origin of 12 provenances sown in the Cowichan research nursery 3 9 5 Geographic origin of 157 spruce cone samples arranged in order of increasing elevation... hh 6 Cone scale morphology of black, white, Engelmann and Sitka spruce U7 7 Comparison of means of cone scale morphology of white and Sitka spruce U 8 8 Seed characteristics of 150 spruce provenances arranged in order of increasing embryo development....... i 62 9 Growth behaviour during the fi r s t year in the nursery 63 10 Growth behaviour during the second year in the nursery &h 11 Dates of flushing of 150 spruce provenances during the second year in the nursery 65 12 Dates of dormancy of 150 spruce provenances during the second year in the nursery... 66 13 Dates of flushing and dormancy of 12 spruce provenances on two soil types both inside and outside plastic greenhouse.. 67 Iii Relationship between germination behaviour and seed quality at 15°C 68 15 Relationship between germination behaviour and seed quality at 20°C 69 16 Relationship between germination behaviour and seed quality at 2 5°C. 70 1 7 Relationship between germination behaviour and seed quality at 30°c 71 18 Relationship between germination behaviour and factors of the environment • 72. VIII Table Page 19 Relationship between flushing and factors of the environment 73 20 Relationship between growth rate during second year and fac-tors of t he environment • lh 21 Relationship between- growth and dormancy during first and second year and factors of the environment............. 75 22 Correlations between variables included in the principal component analysis 76 23 Weighing for original variables in computed components...... 77 2li Percentage of variation accounted for by four principal com-ponents 78 .' 25 Percent of provenances in each of 5 elevational groups with a germination value of less than one at 15°C. 98 26 Average germination capacity of Engelmann spruce in comparison with other species... • 98 27 Means of foliage samples from 60 trees and geographic origin of each sample. 1°5 28 Percentage of 2-year-old seedlings of 13 spruce provenances dormant by August 22 at southern and northern nurseries in British Columbia. 137 LIST OF FIGURES IX Figure Page 1 Diagrammatic representation of spruce scale and bract showing five basic measurements.. • .. h$ 2 Characteristic curves obtained when cone scale morphology of populations from allopatric zones of white, Engelmann and Sitka spruce is compared by line of shape method. h9 3 Characteristic curves obtained when cone scale morphology of populations from allopatric zones of white and Engelmann spruce is compared by line of shape method...... 50 k Variation in embryo development in a. number of provenances from two spruce seed crops. 5? 5 Relationship between degree of dormancy at the Cowichan nursery on July 15 and altitude at place or origin of each provenance 58 6 Relationship between growth of 12 spruce provenances in a uniform environment and elevation at place of origin...... 59 7 Second years growth of 12 spruce provenances in k environ-ments 8 Relationship between altitude at place of origin and degree of dormancy of each provenance on July lU at the Cowichan nursery............... • 61 9 The relationship between continentality and temperature regime for representative climatic stations in British Columbia 170 10 The relationship between altitude and temperature regime for representative climatic stations in British Columbia... 171 11 The relationship between photoperiod and temperature at the Cowichan nursery and at Vaveriby in south central British Columbia 172 12 The relationship between temperature and photoperiod at the Cowichan nursery and at Babine Lake in central British Columbia. 13 The relationship between temperature and photoperiod at the Cowichan nursery and at Barkerville in east central British Columbia. • • Ik The relationship between temperature and photoperiod at the Cowichan nursery and at Terrace in northwestern British Columbia •• 173 17U 175 X Figure Page 15 The relationship between temperature and photoperiod at the Cowichan nursery and at Aleza Lake in east central British Columbia..... 176 16 The relationship between temperature and photoperiod at the Cowichan nursery and at Prince George in central British Columbia 3-77 17 The relationship between temperature and photoperiod- at the Cowichan nursery and at Allison Pass in southern British Columbia.... 178 18 The relationship between temperature and photoperiod at the Cowichan nursery and on Old Glory Mountain in southern British Columbia 179 19 Degree of correlation between factors of the environment and flushing and dormancy.... 130 20 Curves of flushing for Sitka, white and Engelmann spruce.... 181 21 Curves of dormancy for Sitka spruce and provenances from sympatric zones of white and Sitka spruce. • 182 22 Curves of dormancy for provenances from sympatric populations of white and Sitka spruce, and low elevation allopatric spruce provenances.. • 183 23 Curves of dormancy for spruce provenances from elevations between 2000 and 2300 ft , 18U 2k Curves of dormancy for spruce provenances from elevations between 2300 and 2700 f t . . . . . 1 85 25 Curves of dormancy for spruce provenances from elevations between 3000 and I|000 f t 186 26 Curves of dormancy for spruce provenances from elevations between U000 and 1£00 f t I 8 7 27 Curves of dormancy for spruce provenances from elevations between U600 and 5000 f t I 8 8 28 Relationship between elevation and spruce cone scale-seed . wing ratio. 1963 collection in the white-Engelmann complex. 189 29 Relationship between elevation and spruce cone scale-seed wing ratio. 1961; collection in the white-Engelmann complex. 190 30 Pattern of variation in a single measurement of cone scale morphology. 1963 collection 191 31 Pattern of variation in a single measurement of cone scale morphology. 196ij. collection..... ...» 192 XI Figure Page 32 Pattern of variation in a single measurement of cone scale morphology. Miscellaneous collections... 193 33 The pattern of variation in spruce cone scale morphology along a longitudinal transect from coastal Sitka spruce to montane white spruce forest......... • 19h 3k The pattern of variation in spruce cone scale morphology along an altitudinal transect at Stone Creek, South of Prince George, British Columbia......... 195 35 Variation in cone scale morphology in a sympatric population of white and black spruce. Sample 138 196 36 Variation in cone scale morphology in a sympatric population of white and black spruce. Sample 139. 197 37 Variation in cone scale morphology in a sympatric population of white and black spruce.. Sample l l ; 0 . . . . . . 198 38 Variation in cone scale morphology in a sympatric population of white and black spruce. Sample llj l 199 39 Variation in cone scale morphology in a sympatric population of white and black spruce. Sample l l i 2 . . . . . . . . . . . . . . . . . . . . . . 200 bP Variation in cone scale morphology in a sympatric population of white and black spruce. Sample 1U3 •...«. 201 I4I Variation in cone scale morphology in a sympatric population of white and black spruce.. Sample 114; • 202 kZ Variation in cone scale morphology in a sympatric population of white and black spruce. Sample lU5» •• 203 k3 Variation in cone scale morphology in a sympatric population of white and black spruce. Sample ll;6 201; I4J4. Variation in cone scale morphology in a sympatric population of white and black spruce. Sample lk7 •• 205 ll5 Sympatric and allopatric spruce populations as indicated by discriminant function analysis of cone scale data. 1963 collection 2 0 6 I4.6 Sympatric and allopatric spruce populations as indicated by discriminant function analysis of cone scale data. 1961; colls ction. •••• 207 XII LIST OF ILLUSTRATIONS Illustration Page 1 Layout of seedbeds at the Cowichan nursery 35 2 The X-ray assessment of embryo development in spruce seed..... 36 3 The cone scale morphology of Engelmann, Sitka, white and black spruce U6 k The differential growth behaviour of spruce populations from different elevations when grown in a plastic greenhouse on regular nursery s o i l . . . . . . . . 56 5 Cone scale morphology of white and Engelmann spruce, and in-termediate form.. 101 6 Cone scale morphology of sympatric populations of white and black spruce in the boreal forests of northern British Columbia 1 0 2 7 The variation pattern in cone scale morphology along a longi-tudinal transect from coastal Sitka spruce to montane white spruce forest..- 103 8 Cone scale morphology of Sitka and white spruce and inter-mediate form. 10 U 9 The subalpine Engelmann spruce forest.. 128 10 The white-Engelmann spruce complex in the Rocky Mountain Trench .. 129 11 The white-black spruce complex of the Alaska highway 130 12 The characteristic branching habit in high and low elevation spruce in British Columbia..... 131 13 Variation in branching habit in spruce in British Columbia.... 132 Ik Variation in spruce bark type in British Columbia 133 XIII 5 EXTREMES AND INTERMEDIATE FORMS OF THE SPECTRUM OF VARIATION IN CONE MORPHOLOGY IN THE WHITE-ENGEI21ANN SPRUCE COMPLEX IN BRITISH COLUMBIA. DRAWN TO SCALE FROM SAMPLES IDENTIFIED OVERLEAF PLACE OF ORIGIN OF THE CONE SAMPLES ILLUSTRATED IN FRONTISPIECE Sample No. Lat. Elev.(ft.) Locality 1 13 5n026' 2U00 3 miles ¥. of Fort St.James 2 k9 U9°07 f 2800 UO miles ¥. of Trail 3 50 U9°ll' 3300 31 miles S.W.of Princeton h 6U 52°12 * 2700 6 miles N. of Williams Lake 5 63 5o°oo' 3300 13 miles S. of Merritt 6 111 5oV 5200 52 miles N.E. of Kamioops XV The differences between trees of the same kind have already been considered. Now a l l grow "fairer and more vigorous in their proper positions; for wild, no less than cultivated trees, have each their own positions, some love wet and marshy ground, as black poplar, white willow and in general those that grow by rivers; some love exposed and sunny positions; some prefer a shady place. The f i r is fairest and tallest in a sunny position, and does not grow at a l l in a shady one. The silver-fir on the contrary is fairest in a shady place, and not so vigorous in a sunny one. Yew, pados and joint-fir rejoice exceedingly in shade. On mountain tops and in cold positions odorous cedar grows even to a height, while silver-f i r and Phoenician cedar grow, but not to a height, -for instance on the top of Mt. Cyllene; and holly also grows in high and very wintry positions. These trees then we may reckon as cold-loving; a l l others, one may say in general, prefer a sunny position* However, this too depends partly on the soil appro-priate to each tree; thus they say that in Crete on the mountains of Ida and on the white Mts. the cy-press is found on the peaks whence the snow never disappears; for this is the principal tree both in the island generally and in the mountains. Again, as has been said already, both of wild and of cultivated trees some belong more to the moun-tains, some to the plains. And on the mountains themselves in proportion to the height some grow fairer and more vigorous in the lower regions, some about the peaks. THEOPHRASTOS OF ERESOS (380-28? B.C.). !L¥TRODUCTION 1 Depending on the objectives of the investigation, a study of geographic variation in a widespread coniferous species may take several forms. Therefore, i t is particularly important for the investigator to make clear, at the outset, the objectives of the study he has undertaken. Ambiguity in this regard can result in considerable misunderstanding as to the silvicultural and/or genecological implications of the results of the investi-gation. For example, Critchfield's study of geographic variation in lodgepole pine (Pinus contorta Dougl.) (Critchfield 1957) and McDonald's criticism of this study (McDonald 1958) is a case in point. Wright and Baldwin's assessment of a large scale provenance t r i a l of Scots pine (P, sylvestris L.) in New Hampshire (Wright and Baldi*in 1957) and Langlet's detailed rebuttal (Langlet 1959) of both their premises and results even more clearly i l l u s -trates the validity of the above statement.. There i s , then, considerable diversity of opinion in regard to the purpose,, methods and scope of studies of geographic variation in tree species. For this reason, and rather than discuss the problem in a general introduction, the matter is treated separately, and in some detail, in the following section. The purely technical problems associated with an assessment of geographic variation in \h ite spruce in British Columbia are enormous in scope. There is for example the major problem of sampling - which has not been satisfactorily solved in the present study. The species has a vast range throughout the province. It is sympatric with other spruce species in nume-rous areas, and introgressive hybridization is known to occur extensively. The problem facing the forest geneticist in this instance, therefore, is analogous to that of the geographer who, going into a very large, rugged relatively unexplored territory must fi r s t plot the approximate location, 2 height and breadth of mountain peaks, the depth and direction of major valleys, and the location of major rivers and lakes (see p. 28 Stebbins 1 9 5 0 ) . Races, ecotypes, clines, and hybrid swarms are the "gross topo-graphy" of genetic variation within the white spruce complex of British Columbia, and i t is this variation with which the present study is principally concerned. This study, therefore, by no means exhausts the possibilities for further study of variation in white spruce in British Columbia, On the contrary i t is hoped that its results will lead directly to more detailed studies, particularly below the population level, in controlled and partially controlled environments. These studies can be conducted simultaneously with the long term assessments of the performance of experimental stock outplanted in diverse environments. In accordance with the tenets established by Anderson (19U-9), and in the belief that long tables of statistics do not satisfactorily illustrate patterns of variation in natural populations, a special effort is made to present the data in graphic form wherever possible. For con-venience of reference, and to avoid extensive divisions of the text, these graphs (Figs. 9 to 1;6) are placed in the appendix. 3 SYSTEMATICA IN RELATION TO GEOGRAPHIC VARIATION IN FOREST TREES: A STATEMENT OF THE ASSUMPTIONS AND OBJECTIVES OF THE PRESENT STUDY. INTRODUCTION.': Any student engaged in a study of geographic variation in a coni-ferous tree will soon realize, i f his interest extends beyond the establish-ment of reciprocal plantations, that i s , the provenance t r i a l , that informa-tion relevant to his study may be gleaned not only from the silvicultural y literature but also from the literature of taxonomy - classical, experimental and numerical - and genecology. This is so because the silvicultural pro* blem of infraspecific variation in a tree species, though not generally re-cognized as such in the silvicultural literature, is in fact a problem of microevolution, and therefore cuts across several branches of botanical science. It is not particularly surprising that Darwin's major thesis was completely anticipated in 1831 in a book entitled "On Naval Timber and arboriculture"• A high percentage of the studies of geographic variation in wide-spread coniferous species reported in the literature takes the form of field trials, generally referred to as provenance trials, of untested progenies, and without reference to the variation patterns in mature populations in the wild. The principal objective of provenance trials in this sense is "to compare the growth of crops of trees from different lots of seed in terms of their ability to produce forests and timber" (Edwards 1956). In general such trials are initiated by foresters. The characteris-tics assessed are silvicultural characteristics, and a provenance is fre-quently referred to as "good" or "bad" depending on whether i t exhibits rapid or slow growth during development. For example: "The fact that one provenance does badly in the nursery is a matter of importance, but i t ii vitiates the future of the experiment and its interpretation in relation to normal practice i f provenance experiments carry forward deficiencies which normally would be obliterated before the end of the nursery stage"(Edwards 1956). But the terms "good", "bad" or "deficient" when applied to total height growth in the nursery - or any other measurable characteristic - are meaningless in regard to vthe genecology of the species, and are not of much value as silvicultural terms either, since, depending on the environment in which they are outplanted,height growth may be reversed between any two pro-venances at a later date (Roche and Revel 1966). There are many exceptions to these generalizations concerning pro-venance tri a l s . Nevertheless, they hold good for a significant number of experiments which have as their objective the assessment, in silvicultural terms, of the effects of variation in tree species but ignore its causes. For example, nowhere in Edwards1 publication is there any indication that the results of assessment of within species variation in forest trees have any significance outside the field of practical silviculture. There i s , however, no real reason why the assessment of variation in tree species should be conducted as i f i t were only a silvicultural pro-blem and the broader implications of within species variation ignored. On the contrary, i t is the writer's belief that the failure to place the problem of infraspecific variation in forest trees in its botanical context is a major reason why the results of a large number of provenance trials are not commensurate with the time and expense incurred during their establishment, maintenance and assessment. what follows, therefore, is an attempt to place the problem of infraspecific variation in a tree species against the broad background of Post-Darwinian systematics. It is hoped that by so doing, unity and co-herance will be given to the subject as a wholej and the objectives of the present study brought into clearer perspective. The need for such an attempt i s , perhaps, indicated by the extent of the divergence of viewpoint ex-pressed in regard to various aspects of subject by Melville 19bP; Edwards 1956; Wright and Baldwin 1957,* MacDonald 1958; Wright, Bingham and Dorman 1958; Barber and Zobel 1959; Callaham, 1959; Nienstaedt I960; Langlet 1959, 1962, 1963a;; Stern 196U. To facilitate presentation, the subject matter is discussed under the following headings (1) Classical taxonomy (2) Experimental Taxonomy (3) Numerical Taxonomy (I4) Genecology. CLASSICAL TAXONOMY Classical taxonomy was a well established discipline before the de-velopment of evolutionary theory. Plants were classified on the basis of morphological characters, and, after Linnaeus, a binomial was given to a l l plants so classified. The publication of Darwin's "Origin of Species" destroyed the prevailing ideas concerning the fixity of species and argued that many species may have common ancestors. Since then attempts have been made to group plants on the basis of evolutionary relationships, or phylogeny Even, the very great advances in genetics, cytology and palebotany since the publication of "The Origin of Species", have not resulted in any radical change in the methods of plant classification followed since the time of Linnaeus. Classical Taxonomy, therefore, which is based to a considerable extent on the intuitive determination of affinities, is s t i l l the predominant method of classification in use at the present time. A very clear statement of the methods of classical taxonomy has been given by Sprague ( I 9 I 1 O ) , and i t is of interest to note that i t appeared in Huxley's "The New Systematics" (Huxley 19lj0), the book which to a considerable extent accelerated the development of biosystematics or experimental taxonomy. Good examples of the application of the methods of classical 6 taxonomy to a tree species, and the great importance of nomenclatural and b i b l i o g r a p h i c a l studies i n t h i s method of c l a s s i f i c a t i o n . , are given by Fernald (1945) i n h i s study of Betula i n Eastern North America, and Dugle (1966) i n her taxonomic study of Western Canadian species of the same genus. The p r a c t i c i n g f o r e s t e r i s seldom appreciative of the a c t i v i t i e s of the c l a s s i c a l taxonomist, and frequently f a i l s to understand how taxonomic i n v e s t i g a t i o n s , when app l i e d to a tree species, can have any bearing on s i l -v i c u l t u r a l problems (MacDonald 1958). Yet, as Wright et a l . (1958) have pointed out, "the taxonomic and h o r t i c u l t u r a l l i t e r a t u r e i s a l a r g e l y neg-l e c t e d source of information about genetic v a r i a t i o n i n f o r e s t trees even though i t contains scores of examples for every example contained i n the f o r e s t r y l i t e r a t u r e " . Furthermore, as M e l v i l l e (1940) has shown, the taxono-mists are not unaware of the s i l v i c u l t u r a l implications of t h e i r work when appl i e d t o a tr e e species. Wright and Baldwin (1957) used the data from al7-year-old, r e p l i c a t e d provenance t e s t of Scots pine i n New Hampshire as a standard against which to compare 19th century provenance s t u d i e s . "They found that herbarium studies made by leading 19th century taxonomists led to a better i n s i g h t i n t o the pattern of v a r i a t i o n than d i d many low-precision provenance t e s t s " (Wright 1962). Constance (1957) has compared the system of c l a s s i c a l taxonomy to a r e p o s i t o r y f o r the t o t a l i t y of a l l kinds of evidence. This i s a v a l i d compa-r i s o n , and the f o r e s t e r engaged i n provenance research would be unwise to neglect suoh a p o t e n t i a l source of information i n regard to the speoies he i s studying. For example no f o r e s t e r i n i t i a t i n g a genecological i n v e s t i g a t i o n of a western Canadian b i r c h species oould a f f o r t t o ignore the methods and r e s u l t s of Dugle's taxonomic inv e s t i g a t i o n s (Dugle 1966), however applied h i s aims, and though nomenclature per se i s not h i s i n t e r e s t . 7 EXPERIMENTAL TAXONOMY A general statement of the purpose, principles, and results of the experimental method in taxonomy has been given by Clausen, Keck and Hiesey (19i;0, 191&, 191*8) and Clausen and Hiesey (1958). These authors state that though morphological comparisons are of fi r s t importance in classification, cytological and genetical tests are the more conclusive indicators of re-lationships. Consequently the taxonomic units derived by the methods of experimental taxonomy are defined almost exclusively in genetic terms. White (1962), having described the biosystematic units recognized by Clausen and his co-workers stated, "Although no group can be considered to be fully worked taxonomically until its components can be arranged in biosystematic units similar to those of Clausen, Keck, and Hiesey, i t is doubtful whether such a classification will be entirely acceptable to practical taxonomists (those who write and use monographs) without modification.—- For most of the world's flora, and particular for slow growing tropical plants of l i t t l e economic importance, i t is unlikely that experimental data will ever be available in sufficient quantity to provide a basis for a biosystematic classification"• According to Good (1961;) there are at least 250,000 or more species of angiosperms alone in the world. Clausen and his co-workers have spent many years in assessing genetic relationships in a relatively minute number of species which were specially selected for their amenability to the methods employed. From a knowledge of both these facts, i t is not possible to com-pletely disagree with White's conclusion. However, there is no doubt that the methods of experimental taxonomy will continue to be applied on an ever-increasing scale, particularly by the genecologist, though his objective differs from that of the experimental taxonbmist. Callaham's advocacy 8 (Callaham 1959) of biosystematic methods in the study of variation in forest trees was, at the time of publication, symptomatic of this general trend. NUMERICAL TAXONOMY In a discussion concerning the classification of plant communities Webb, as early as 1951i, had this to say: For a scientific taxonomy i t is not enough to assure one's critics that with sufficient experience one can learn to recognize the units; there must be some means of defining them. Without this plant sociology can only be a craft to •which one is apprenticed, and not a science which one can learn. It is as i f organic chemistry were conducted without analysis, simply by relying on the sense of smell. Not only is such a procedure uncommunicablej i t is also unreliable (Webb 195U). Webb goes on to recommend a system of multifactorial recording, which, i f necessary, is capable of being carried on punched cards, as a basis for the classification of plant communities. Since the publication of this ar-ticle there has been a veritable flood of proposals in regard to numerical methods of classification of species and plant communities. Most of these publications are represented in the bibliographies of Sokal and Sneath(l963)* which is the first book devoted to numerical taxonomy, and in Greig-Smith (I96I4.) which deals with the numerical classification of plant communities. The development of numerical taxonomy cannot be ignored by anyone en-gaged in the study of natural variation, and since i t is the writer's opinion that Webb's remarks apply not only to classification of plant com-munities but also to problems of classification either above or below the species level, the question remains as to the bearing of numerical taxonomy 9 on investigations of infraspecific variation in long lived plant species such as the coniferous tree. At the outset i t is necessary to make the distinction between "numerical taxonomy", and the "numerical method in taxonomy". Because of this, and in view of what has been said above, the following must not be construed as adverse criticism of numerical methods in taxonomy. Nor is i t an analysis of the methods of numerical taxonomy, for the writer is qualified neither by training nor interest to make such an analysis. In any event i t has already been done (Heywood and McNeill 196k, Seal 196U, Williams and Dale 1965, McAllister 1966). What follows then, is an attempt to relate some of the basic assumptions of numerical taxonomy to problems encountered in genecological investigations.of forest trees. An excellent summary of some of the basic assumptions of numerical taxonomy is given by Ehrlich and Holm (1962), and is here quoted in f u l l : In summary, then, we would like to suggest that in broad investigations of the patterns of interaction and relationship among organisms the ar t i f i c i a l and stultifying fragmentation of population biology into divisions such as taxonomy, popula-tion genetics, and ecology should be ignored. Care should also be taken to scrutinize current concepts such as 'species', 'niche', and 'community'. If some emergent patterns seem to correspond to a degree with these concepts, then the concepts may be given operational definitions and the labels should be retained. If there is no such correspondence, then the concepts will have outlived their usefulness and should be discarded. The basic units of population biology are not communities, species, or even populations but individual organisms. In populations, variation, growth, genetic equilibria, selection, 10 behaviour and so on are not 'things' but relationships. There-fore, what is of interest in population biology is the pattern in which organisms are related in space and time. Since these assumptions of EhTlich and Holm have been critically examined in some detail by Stebbins (1963) no attempt will be made here to discuss, their validity at length. However, i t is important to note that according to these authors the various branches of botanical science which have a bearing on systematics should be ignored in investigations of relationships among organisms. In other words taxonomic relationships are evaluated purely on the basis of resemblances existing in the material at hand, and a total ignorance is presupposed concerning the ecology of the species or variety requiring classifications. It is also clear that Ehrlich and Holm (1962) think of evolution in terms of individuals rather than populations, but as Hardy (1965 p. 170) has pointed out, i t is the populations which are evolving not the individuals. Any forester who has closely studied within and between population variation in coniferous tree species will have no difficulty in agreeing with Hardy. The emphasis on descriptive morphology as the principal criterion of assessing relationships between organisms places the numerical taxono-mists in the tradition of classical taxonomy, and Stebbins' rhetorical ques* tion posed with regard to Ehrlich's paper may equally be asked of numerical taxonomy as a whole: "In essence, does Ehrlich's apparently new approach to classification and species relationships represent anything more than an elaboration of old-fashioned morphological taxonomy, dressed in a shiny new algebraic suit of cloths, and given additional allure by means of that largest and most costly of modern scientific status symbols, the digital computer?" (Stebbins 1 9 6 3 ) . 11 In reviewing the second edition of Huxley's "Evolution: the modern Synthesis" (Huxley 1963), De Beer (I963) stated:"—Although i t has been known since 1859 that evolution by natural selection was a unifying concept for a l l biology, i t is only now becoming clear how all-embracing i t i s , and how closely i t unites together the numerous and ever-increasing branches and experimental disciplines of biology". This is a statement of the synthetic theory of evolution, and i t is this theory (held not only by De Beer and Huxley, but also by Stebbins (1950) and Mayr (1963)) that is implicitly, i f not always explicitly, denied by the protagonists of numerical taxonomy. However, in general i t may be said that the literature of numeri-cal taxonomy (Sokal and Sneath 1963) is less a rebuttal of the synthetic theory of evolution than a demonstration of the belief that biology is a sub-division of physics. For example the work of Charles Darwin is referred to on five pages of Sokal and Sneath's book, whereas the work of T.T. Tanimoto (a mathematician in the research.division of International Business Machines) is referred to on no less than twenty-two pages. It is difficult to avoid the conclusion that numerical taxonomy has been designed to f i t the computor rather than to cope with the biological facts of variation and evolution. It is true that these facts are frequently inadequate. It i s , however, also true that in the past many of the most brilliant discoveries in biology have been made by imaginative men who exerci-sed their judgement on inadequate facts - a faculty which, perhaps fortunately, is not yet the prerogative of the computor. In conclusion i t is necessary to state that, in regard to the assess-ment of infraspecific variation in a tree species, numerical taxonomy is rightly considered a valuable contribution to the numerical method in tax-onomy - an important distinction. For there is a numerical method in tax-onomy which is evolving within the framework of. the .science. The objection to numerical taxonomy, therefore, does not l i e in the fact that i t uses numerical methods but in the fact that its basic assumptions preclude the application of other methods and the utilization of much knowledge of species -environment interaction which followed the development of post -Darwinian systematics. Therefore, no synthesis is possible. However " — f o r the under-standing of an important phase of raicroevolution the synthesis is indis-pensable, in the same way that a synthesis - on a broader canvas - was essential in the development of the argument cf the 'The Origin of Species' (Heslop - Harrison 196l±);n and, as already pointed out, the problem of in-fraspecific variation in forest trees is a problem of microevolution. GENECOLOGY As stated above, though the methods of the genecologist are similar to those of the experimental taxonomists, his objectives are different. The genecologist is primarily interested in determining habitat-correlated, gene-tically-based variation within the species (Turesson 1 9 2 3 ) , whereas the principal objective of the experimental taxonomist is classification. "Thus, ecological data and observations on genetic system, are a sine qua non of genecology, although by no means an essential part of taxonomy. Conversely, the nomenclatural, and biographical studies which are an obligatory part of any taxonomic study are not necessarily significant for a genecological inves-tigation of a species". (Heslop - Harrison I 9 6 U ) • The publication referred to above is a detailed and lucid review of kO years of genecology, and since i t contains an extensive and up-to-date bibliography no attempt will be made here to survey the literature of geneco-logy. The important point is that although classification is not the objective of genecological studies, such studies, particularly of plant species, continue to elucidate some important evolutionary principles, and consequently genecology has a definite, though indirect, bearing on phylogenetic classification. Genecology is essentially a synthetic science. It incorporates in modified form the approaches of the several disciplines outlined above. Its basic propositions have been stated succinctly by Heslop-Harrison (l°61i), and are quoted below: 1) Wide ranging plant species show spatial variation in mor-phological and physiological characteristics. 2) Much of this infraspecific variation can be correlated with habitat differences. 3) To the extent that ecologically-correlated variation is not simply due to plastic response to environment, i t i s attributable to the action of natural selection in moulding locally adapted populations from the pool of genetical variation available to the species as a whole. A provenance study is rightly considered an investigation of infra-specific variation in a plant species in relation to its environment. There-fore, in purpose, methods, and scope i t is a genecologic investigation, and its basic propositions are those stated above. Timofeef-Ressovsky (19UC-) has made the clear distinction between macro- and micro-evolution, and has pointed out that while the classical methods which gave a picture of macroevolution (the synthesis of palaeon-tological, morphological, embryological and biogeographical data) now seem more or less exhausted, l i t t l e has been done in the field of microevolution. He defined microevolution as the evolutionary process taking place within shorter limits of time, smaller groups of organisms, and lower systematic categories, and concluded that the main phenomenon of microevolution is geographical variability. 14 I f the main phenomenon of mioroevolution i s geographical v a r i a b i -l i t y then i t i s c l e a r that studies of geographical v a r i a b i l i t y i n f o r e s t t r e e s , i f r i g h t l y designed, could contribute s u b s t a n t i a l l y to the general understanding of microevolution. Stebbins (1950) has pointed out that the dominant evolutionary processes are d i f f e r e n t at each of the three l e v e l s of v a r i a t i o n i n n a t u r a l populations: " I n d i v i d u a l variation i s dominated by gene mutation and genetic recombination, microevolution by natural s e l e c t i o n , and maoroevolution by a combination of s e l e c t i o n and the development of i s o l a t i n g meohanisms...." Therefore, i t i s the population which i s characterized by i t s environment, f o r , as already pointed out, i t i s the population which i s evolving and not the i n d i v i d u a l - an important d i s t i n c t i o n often not observed i n studies of v a r i a t i o n i n f o r e s t t r e e s . Hence the unrewarding search f o r populations whioh are endowed with c h a r a c t e r i s t i c s randomly occurring i n i n d i v i d u a l t r e e s ; and the not infrequent attempts to r e l a t e v a r i a t i o n i n i n d i v i d u a l t r e e s , or very small numbers of trees, of d i f f e r e n t populations to f a c t o r s of the environment at t h e i r place of o r i g i n . In the h i g h l y heterozygous coniferous species many desirable s i l v i -c u l t u r a l c h a r a c t e r i s t i c s are most frequently the r e s u l t of genetic segrega-t i o n and recombination, and, therefore, are r i g h t l y studied at the l e v e l of i n d i v i d u a l v a r i a t i o n . These are not, generally, the r e s u l t o f s e l e c t i o n and adaptation. For example, as Langlet (1963) has pointed out, from an eoologi-o a l point of view, increment i n f o r e s t trees i s only of secondary importance since i t does not d i r e c t l y oondition s u r v i v a l or reproduction. The emphasis, therefore, on s i l v i c u l t u r a l c h a r a c t e r i s t i c s i n prove-nance t r i a l s , e x p e c i a l l y during the e a r l y phases of the t e s t , i s mis-placed. What is of principal interest is the growth rhythm of each provenance at the test site in relation both to the environment at the test site, and the envi-ronment at its place of origin. In this respect careful measurements of flushing and dormancy, for example, are of greater significance than increment measurements. It is only when some understanding of genetically-based, habitat-correlated variation has been obtained that the silvicultural poten-t i a l of a provenance is known. Furthermore, such an understanding can be obtained from carefully designed studies of the growth rhythm of diverse popu-lations at the juvenile stage in controlled and partially controlled environ-ments (Wareing 19ji>0, a, b, cj 1951, 1956) . Increment measurements have greater importance in fie l d tests, and are especially significant at the level of individual variation. The assessment of genecological differentiation within a tree species need not be confined to physiological studies on progenies grown in a uniform environment. As pointed out by Heslop-Harrison (I96J4. p. 2 1 7 ) there is no direct evidence that habitat-correlated variation in morphological features is non adaptive, and that i f populations in one type of habitat are regularly found to differ from those in another in any characteristic whatever, those differences have adaptive significance, and are due to the differential effect of selection in the two environments. Therefore, biometrical studies on mass collections of field specimens of cones and foliage can in many ins-tances also yield valuable information concerning the genecology of the species under study, as already indicated in the discussion of classical taxonomy. Such studies are likely to be of especial value in, for example, the forest regions of Canada where relatively l i t t l e is known of the dis-tribution, taxonomy and degree of introgressive hybridization of many coniferous species (Roche 196U)• CONCLUSIONS Infraspecific variation in a tree species is best considered as a problem in microevolution rather than a purely silvicultural problem, when so considered the problem is brought into clearer perspective, and objectives become more sharply defined. Genecology, a synthetic science which embraces the ideas and methods of the several botanical disciplines discussed above, is the study of infraspecific variation of plants in relation to their environment. Thus i t s basic propositions and methods apply directly to studies of geogra-phic variation in tree species. A purely numerical approach to the study of infraspecific variation in the manner of the numerical taxonomists is precluded because of the synthetic nature of genecology. In this context numerical taxonomy is considered simply as a statistical technique which may illustrate certain aspects of genecological differentiation, for example morphological variation in mass collections of field specimens. The synthetic theory of evolution as outlined by Hardy (1965) is the great unifying concept in biology, and i t is this theory which both illuminates and provides the framework for interpretation of a l l phenomena relating to variation in natural populations, including populations of forest trees. DISTRIBUTION AND PHYLOGENETIC RELATIONSHIPS OF THE SPRUCE SPECIES OF BRITISH COLUMBIA There are three species and one subspecies of spruce in B.C. which between them form a complex of sympatric and allopatric populations with a province wide distribution. These are Sitka spruce (Picea sitehensis (Bong.) Carr.), black spruce (P. mariana (Mill.) BSP), white spruce (P. glauca (Moench) Voss subsp. glauca) and Engelmann spruce (P. glauca (Moench) Voss subsp. engelmannii). The nomenclature follows that proposed by Taylor (1959). The taxonomic literature (Little 1953&) also recognizes the occurrence of two other varieties of white spruce in British Columbia. These are Porsild spruce (P. glauca var. porsildii Raup.) and western white spruce (P. glauca var. albertiana (S. Brown) Sarg). The distribution of the components of this great spruce complex is known only in very general terms indeed (see map 2, Stanek 1966) and literature references to distribution and taxonomic relationships are conflicting in the extreme. There are no detailed distributional,maps avail-able for British Columbia other than those published by Whitford and Craig in 1918. These distributional maps continue to be referred to in the l i -terature. For example, Hansen (1955) interpreted the results of bog pollen analysis in British Columbia, on the basis of the modern distribution of spruce species as given by ¥hitford and Craig. There are, however, a number of reports and studies dealing with the distribution and taxonomic relation-ships of white and Engelmann spruce in B.C. (McKinnon 1938, Griffith I9I4O, Garman 1957, Taylor 1959). Whitford and Craig (1918) placed the northern limits of Engelmann spruce almost at lat. 58°Q0,,and the southern limits of white and black spruce at approximately l a t . 53°30!. These distributional limits, therefore, 18 in the absence of complete altitudinal segregation of species, indicate a very large sympatric zone of white, black and Engelmann spruce between "latitudes 5 3 ° 0 0 ! and 5 8 ° 0 0 T . This zone, broadly speaking, corresponds to zone 9 (sub-boreal) of Krajina's classification of biogeoclimatic zones of British Columbia (Krajina 1 9 6 5 ) , though zone 8 (Englemann spruce-subalpine fi r ) of this classification gives the Yukon border (Lat. 6 0 ° 0 0 ' ) as the northern limit of Engelmann spruce in B.C. However, a northern distri-bution of this extent for Engelmann spruce is not corroborated by the fi n -dings of Griffith (19U0), Raup ( 1 9 U 5 ) or Garman ( 1 9 5 7 ) . In further contrast to the southern distributional limits of white, and the northern distributional limits of Engelmann spruce as given by Whitford and Craig, Garman ( 1 9 5 7 ) has reported the occurrence of white spruce at lat. U 9 ° 3 0 I in the East Kootenay valley at altitudes of 2 5 0 0 to 3 0 0 0 f t . and Recknagel ( 1 9 3 9 ) has stated that reconnaissance studies along the upper reaches of the Fraser river (along Lat. 5 U ° 0 0 ' between longitudes 1 2 0 ° 0 0 F and 1 2 3 ° 0 0 ' ) definitely determined that the prevailing species of spruce is white spruce and not Engelmann. Raup ( 1 9 U 5 ) has reported the extensive occurrence of sympatric populations of white and black spruce along the Alaska highway, particularly between Fort St-.John and Whitehorse. Garman ( 1 9 5 7 ) , who sampled spruce po-pulations along the Alaska highway north of Fort St. John, has also recorded the occurrence of a sympatric population of white and black spruce at Muncho Lake, lat. 5 9 ° 0 0 ' # However, he considered that the association of these two species in this area was not common. Wight ( 1 9 5 5 ) has. stated that although the range of black spruce approaches that of Engelmann and Sitka i t does not overlap them as does white spruce. Raup (19.1(5) has also reported the occurrence of Porsild spruce (P. glauca var. porsildii) in northern British Columbia. However, Garman (1957) found only white spruce in northeastern British Columbia, and detected no trees having the smooth-bark, and broad-crown characteristic of Porsild spruce. Most authors are in agreement that the variety P. glauca var. albertiana is the hybrid between white and Engelmann spruce (Little 1953, Wright 1955, Garman 1957, Taylor 1959); Garman (1957) is the principal authority in regard to its general distribution in the province of British Columbia. There has been no detailed study of introgressive hybridization between the spruce species of British Columbia, though Taylor's (1959) and Garman's (1957) studies, and Horton's (1959) study in Alberta have provided considerable evidence of its occurrence in populations of white and Engelmann spruce. The most important work in regard to crossibility in spruce is that of Wright (1955)* Wright crossed numerous spruce species, including those which are indigenous in British Columbia, and in addition surveyed the literature pertaining to crossability between the species.. The data presented overleaf, summarize that portion of Wright's Hbrk which has a bearing on the problem in British Columbia. Black spruce hybridizes freely with red spruce (Morgenstern and Farrar 1961;) and, since the publications of Wright's findings, Little and Pauley (1958) have located and described a natural hybrid between black and white spruce. There i s , then, evidence, both from studies of natural populations and a r t i f i c i a l hybridization that there are no major genetic barriers to hybridization between white and Engelmann, white and Sitka, and Engelmann and Sitka spruce. The evidence is less conclusive for black and white spruce but, nevertheless, i t appears that hybridization between both is also possible. 20 SUMMARY OF REPORTED CROSSES IN WHITE, BLACK, ENGELMANN MM) SITKA SPRUCE (FROM TABLE 2, WRIGHT 1955) Male parent Female Parent glauca  mariana  engelmannii sitchensis glauca F S S mariana U engelmannii sitchensis S S S - successful cross, F - unsuccessful cross, U - results of cross uncertain. 21 In regard to white, Engelmann, and Sitka spruce Wright (1955) concluded that morphological, distributional and genetic data indicate a common, relatively recent, origin of these species, and that they are related to the Old World spruce species through P. jezoensis (Sieb. and Zucc.) Carr. of Japan. However, Fowler (1966) has recently published in-teresting results of a r t i f i c i a l hybridization in a number of spruce species which led him to conclude that P. glauca and not P. jezoensis. is the connecting link between Old World and Western North American spruces. Wright (1955) also concluded that P. mariana is closely related to P. rubens Sarg.. and is a remnant of a more ancient migration than that which gave rise to the other three species. Studies of post-glacial forest succession based on bog pollen analysis, which are relevant to the present discussion, indicate that a l l four spruces which occur in B.C. survived the pleistocene in southern refugia, and re-invaded northern regions following the withdrawal of the ice (Hansen 19^7, 1955, Heusser I960, Watts and Wight I966) . However, Hansen (1955) has suggested that spruce may also have reinvaded central and south central British Columbia from refugia along the eastern flank of the Rocky Mountains in Alberta and British Columbia, and in west central Yukon during the late Wisconsin. It appears that the isolated stands of white spruce in Wyoming, Montana, and in the Black Hills of South Dakota are relics of the spruce populations of the southern refugia referred to above. The white spruce populations in the Black Hills are in association with ponderosa pine (Pinus ponderosa), dwarf birch (Betula glandulosa) and the shrub Shepherd- is canadensis (Watts and Wright I966). Watts and Wright carried out pollen and seed analysis in an alluvia-ted lowland 200 km. southeast of the white spruce stands in the Black Hills. Their results indicated that the Black Hills population is a relic of a semi-continuous distribution during late Wisconsin times, and that forests from Canada and the eastern Rockies were probably in contact in the Western Great Plains during Wisconsin. The late Wisconsin forest inferred from their results is described as xeric but cool spruce forest with dry openings, (see i l l u s . 10) . Watts and Wright found a relatively high incidence of the herb Artemisia in their pollen counts, and explained its presence with spruce fossils by suggesting that there may have been a mosaic-type vegetation marked by Artemisia-dominated cover in dunes, and virtually pure stands of white spruce in hollows, resulting in the forest type described above. It was concluded that this forest type may have no modern analogue. However, in this regard i t may be noted that Hansen (1955) has reported the occur-rence of Artemisia tridentata on open slopes adjacent to a spruce stand on a bog two miles south of Clinton in British Columbia, and that there are sympatric populations of ponderosa pine and spruce in the Rocky Mountain Trench in British Columbia (Eastham 19h9)• PHOTOPERIODICITY IN FOREST TREES: A LITERATURE REVIEW There is now considerable evidence that photoperiod or day length exercises a major influence on the growth rhythm of a number of tree species. Since the classic work of Garner and Allard (1920) i t has been a, known that the development of many herbaci'ous plant species is strongly influenced by photoperiod, and in recent years i t has become evident that not only development but growth also is so influenced. The increasingly sophisticated techniques of controlled environ-ment studies have been applied in recent years to assessing photoperiodic response in woody plant species, and there is now considerable experimental evidence that photoperiod exercises a major influence on the growth rhythm of many woody plants including a large number of deciduous and coniferous trees of the montane, subalpine and boreal forest regions. R. Van der Veen (1951) has studied the influence of day length on the dormancy of some species of the genus Populus by growing cuttings of several species in five different growth cabinets, and exposing them to different temperatures and photoperiods. His results showed that plants subjected to a short day treatment were the f i r s t to go dormant even though the temperature regime of the short day was higher than that of the long day treatment. Nevertheless, under the latter treatment a l l plants conti-nued growth. In a series of controlled experiments of considerable significance for foresters Wareing (1950a, 1950b, 1950c, 1951) showed that in a number of woody species dormancy is hastened by short day conditions, and that the cessation of cambial activity is also controlled by photoperiod, and takes place after the cessation of extention growth. Wareing's results also indicated that there was l i t t l e evidence that the breaking of dormancy in the spring under natural conditions is photoperiodically controlled, for 2k as he points out, i f plants have been previously exposed to low temperatures, dormancy i s readily broken at any time regardless of the photoperiod, simply by transferring the plants to warm conditions. Wareing also observed that in the case of Scots pine there is often a reduction in internode length tending towards a "rosette" growth-habit under short days (Wareing 1951)• Pauley and Perry (I95i|) demonstrated that high latitude clones of Populus tricocarpa Torr and Gray, when grown in the natural day regime at the latitude of Boston U.S.A. (ca. l42°N), cease height growth on or about the time of the summer solstice. However, i f photoperiod is lengthened by a r t i f i c i a l illumination they continue to grow. These authors concluded that photoperiod influences the onset of dormancy, but does not appear to be directly concerned with flushing. In a detailed study of geographic variation in Tsuga canadensis (L.) Carr. in controlled environments, Olson et a l . (1959) concluded that pro-venances from high latitudes and high elevations terminated growth before these from low latitudes and low elevations. These results suggested that there is a critical night-length for Tsuga canadensis of 8 to 9 hours at which there is a maximum change from active shoots to dormant buds. Olsen et a l . also concluded that the similarity in response .of hemlock seeds -and buds to photoperiod, temperature, and previous chilling requirements, suggest that the mechanism which regulates the germination of seeds is also related to such vegetative phenomena as the breaking of bud dormancy, growth and return to dormancy. Vaartaja (1959) conducted numerous studies of photoperiodic res-ponse in tree species. He concluded that for most of the species studied, including Picea glauca, decreasing day length after midsummer appears to be an important factor in initiating the rest period. Vaartaja also concluded that the critical day length for continental trees should be-longer than 25 that for maritime trees. To illustrate photoperiodic reactions of trees under outside conditions Vaartaja (1959) constructed a table from the data reported by Heikinheirao (195U). The table gives the fresh weight of one-year-old seedlings of Picea abies (L.) Karst (Norway spruce) from different sources when grown at five different latitudes li7°00 1 , 5l°00 1 , 55°00',, 60°00', and 66°00'N. Allowing for nursery effects other than climate i t is quite clear from the results presented that there is a strong interaction between growth and photoperiod at each nursery. Maximum differences between provenances occurred in the southern nursery at lat. 5l°00' where provenances from high latitudes showed least growth, and provenances from southern latitudes maximum growth. Besides the study of Olsen et a l , i t seems clear from the results of other investigations that provenances from high altitudes respond to photoperiod in a manner similar to that of provenances from high latitudes. In this regard Vaartaja's interpretation of Heikinheimo's data referred to above, and the results of investigations reported below are of considerable interest and relevance to the present study. Karschon (191+9) found that the lower the elevation of Scots pine provenances the greater the growth under a short day treatment. Vaartaja (I960) showed differences between high (63OO ft.) and low (500 ft.) eleva-tion provenances in response to photoperiod. Under short days the high elevation source grew less vigorously than the low elevation source. Special interest is attached to the studies of Irgens-Moller (1958) in that he investigated photoperiodic response in provenances of Pseudotsuga  menziesii (Mirb.) Franco (Douglas-fir) from British Columbia. Provenances from Salmon Arm in the interior of B.C. (Elev. ca 2000 ft.) when grown in a nursery at Corvallis, Oregon, entered dormancy before provenances from coastal British Columbia. When grown in controlled environments the same in-terior provenances grew more vigorously under a long photoperiod, and showed least growth under a short photoperiod. From similar investigations with the same species reported in 1962 Irgens-Moller concluded that differences in response to photoperiod do exist among seedlings of different origin in-dependently of possible differences in chilling requirements. In a recent paper Orr-Erwing (1966) showed the profound effect of the male parent on the time of cessation of growth and dormancy in the pro-genies of controlled crosses of coastal and continental Douglas f i r . He thus indicated the degree of genetic control on this phase of the growth rhythm of the species. One of the strongest correlations between factors of the environment at their place of origin and the growth behaviour of diverse provenances in a uniform environment is that demonstrated by Langlet for Scots pine (Langlet 1959). Stebbins (1950) has stated that this particular study is one of the best examples of a cline within a plant species. Langlet used as an index of the environment at the place of origin of each provenance the day length on that day of the year which first showed a temperature of plus 6°C. The correlation between this index of the environment and the dry matter content of 52 provenances of Scots pine was striking. The correlation coefficient was +• O.98. Thus approximately 97% of the original variance in regard to dry matter content in 2-k year old seedlings, was removed by eliminating the influence of day length during the first + 6°C. day. More importantly, Langlet (1959) has also shown that the growth behaviour of h£ different provenances of Scots pine at 17 years is similarly correlated. The investigations of photoperiodic response in tree species in controlled and partially controlled environments, reported above, provide experimental evidence of the profound effect of photoperiod, and its inter-action with temperature, in influencing the growth rhythm of tree species. The question remains as to what extent the growth behaviour of the diverse spruce provenances at Cowichan Lake has been similarly influenced. MATERIALS AND METHODS To facilitate presentation the study is divided into two parts, A and B. , Part A is a study of geographic variation in immature populations of white spruce, and refers principally to a study of physiological varia-tion of 150 different spruce provenances sown in a relatively uniform envi-ronment in the spring of 1965. Part A also incorporates references to a pilot investigation of variation in immature spruce populations. However, the data resulting from this investigation are ancilliary to the major study of physiological variation of l£0 spruce provenances referred to above. Part B is a study of geographic variation in mature populations,, of white spruce, and refers principally to a biometrical study of variation in cone scale morphology which was carried out on a mass collection of spruce cones collected in 157 areas throughout the range of spruce in British Columbia during the summers of I 9 6 3 and I96I4.. Part B also refers to other variable characteristics in mature spruce populations, for example foliage morphology, bark type, and branching habit. However, no detailed study was made of these characteristics, and as in part A, the data presented is purely ancilliary to the major study of variation in cone scale morphology. PART A Seeds from 150 spruce provenances were sown at the Cowichan Lake research station on Vancouver Island, British Columbia, in the spring of 1965 (illus. 1). Germination tests, which are referred to below, were conducted in the laboratory on a separate portion of each seed sample. Each provenance represents the bulked seed of a varying number of trees collected and registered under the supervision of professional personnel of the British Columbia Forest Service. Table 3 and f i g . 5 give details of the geographic origin of each provenance. Fig. 5 also shows-the location of the Cowichan research nursery. The provenances were randomized in each of six blocks and sown on a prepared soil mix (University of California 1957) in rows six inches apart and at right angles to the long axis of each bed. Germination was facilitated by covering the beds with plastic-sheeted, fiberglass flyscreens. When germination was complete the plastic sheeting was removed from the flyscreens, and the seedlings thinned to approximately 2 cm. apart by cutting superfluous seedlings at the root collar. When growth had proceeded beyond the cotyledon stage the fiber-glass screens were removed from the beds. During the winter the beds were again covered with plastic-sheeted, fiberglass screens. To facilitate measurement on the same seedlings during the growing season 10 seedlings per row were selected by placing a board with 10 evenly spaced teeth alongside each row and selecting that seedling nearest each tooth. A coloured chicken ring was placed over each selected seedling. At the end of the f i r s t growing season a sample of 10 seedlings was obtained for study from each row by placing a board with 10 evenly spaced teeth (so arranged as to li e between the permanently marked seedlings) and selecting that seedling nearest each tooth. The seedlings were cut at the root collar,, and the following measurements obtained in the laboratory; (1) epijjotyl length (2) root collar diameter (3) dry weight. Flushing and dormancy assessments were made at the beginning and at the end of the second growing season by scoring permanently marked seedlings in each row. A seedling was considered flushed when the bud scales were ruptured and needles were visible. A seedling was considered dormant when the terminal needles of the epicotyl were stiff and whorled, and a terminal bud visible. Shoot length was obtained every two weeks during the second growing season by measuring permanently marked seedlings. At the end of the second growing season the permanently marked seedlings were lifted by cutting at the root collar, and the following measurements obtained in the laboratory (l) shoot length (2) root collar diameter (3) dry weight. Using X-ray techniques, Scandinavian workers have demonstrated that embryo development in spruce and pine seed is highly correlated with germination behaviour. It is also possible that embryo development may influence early growth in the nursery. Consequently provenance differences in germination behaviour and even early growth (differences which, unlike embryo development, are genetically based) may be obscured i f no account is taken of this factor. In order, therefore, to obtain a measure of seed quality (expressed as embryo development) which would allow a more accurate interpretation of the ecological significance of the results of germination tests, k repli-cations of 100 seeds from each provenance were X-rayed. A X-ray unit pro-ducing soft rays was used for this purpose (Nippon super soft X-ray apparatus Type EM supplied by the Nippon Softex Co. Ltd. Tokyo), and the conditions of exposure were as follows (15 kV, 1.5 mA, f. 35 cm., 15 s e c ) . The quality of the radiographs thus obtained is shown in i l l u s . 2. Using a Bausch and Lomb, Stereoscopic binocular microscope, embryo development and f u l l seed percent were assessed from the resulting radiographs, and classi-fied in the manner indicated by Table 1 and Illus. 2. Germination tests were also conducted at 15, 20 and 30°C k replica-tions of 100 seeds each were germinated at each of these temperatures. In a l l germination tests conducted, the seed was unstratified and germinated on fi l t e r paper in petri dishes f i l l e d with vermiculite. Water was supplied only once, and at the beginning of each test. In a l l instances the indices of germination behaviour were calculated in the manner proposed by Czabator (1962), and the following table is from Muller-Olsen et al 1956. TABLE 1 EMBRYO AND ENDOSPERM CLASSES CORRESPONDING TO THOSE SHOWN IN ILLUS. 2 Embryo class 0 - Neither embryo nor endosperm (empty seed)• 1 - Endosperm but no embryo. II - Endosperm and one or several embryos, none of which is longer than half the embryo cavity. I l l - Endosperm and one or more embryos, the longest of which measures between half and three-quarters the embryo cavity. IV - Endosperm with one fully developed embryo completely, or almost completely, occupying the embryo cavity. Diminutive embryos rarely occur. Endosperm class A- - The endosperm almost f i l l s the seed coat to capacity and absorbs the x-radiation well. B - The endosperm only partially f i l l s the seed coat and is often shrunken or otherwise deformed. The X-ray absorption is inferior to that of class A. The pilot investigation of variation in immature spruce populations refers to 12 spruce provenances sown on two soil types in the spring of 196k at the Cowichan research nursery ('Table k)» In the f a l l of 196k the 12 provenances were sampled by placing a board with 10 teeth alongside each row of seedlings and taking that seedlings which was closest to each of the 10 teeth to give 10 seedlings per row. Since there were k replications, each provenance was represented by 1|.0 seedlings. Shoot length and root collar diameter were obtained for every seedling and the mean of a l l measurements calculated for the 12 provenances• During the early spring of the second year a plastic greenhouse was placed over two blocks, one of the a r t i f i c i a l soil mix, and one of local s o i l . A sampling and measuring procedure similar to that referred to above was followed in assessing the second year's growth. In addition, however, shoot extension was measured every two weeks during the growing season. Dates of flushing and dormancy were also obtained. A l l dataware subjected to statistical analysis in order to de-termine the relation between the variation observed and factors of the environment at the place of origin of provenances. This included principal component analysis and multiple regression analysis. Principal component analysis is a form of multivariate analysis which in recent years has found increasing application in genecological and taxonomic investigation. The mathematics of the technique have been discussed by Seal 19&k (pp. 101-122), and its application in regard to geographic variation and taxonomy of tree species has been demonstrated by Jeffers and Black 1963, and Gardiner'and Jeffers 1963. Jeffers 1965 has briefly summarized the objectives of principal component analysis, and his summary is given below in f u l l : 1. Examination of correlations between separate variables. 2. Reduction of the basic dimensions of the variability to the smallest number of meaningful dimensions. 3 . Elimination of variables which contribute relatively l i t t l e extra information to the study. U. Examination of the taxonomic groupings of the individuals 5. Determination of the objective weighting of the variables in the construction of taxonomic indices. 6. The: identification of individuals of doubtful or unknown origin. 7. The recognition of misidentified specimens. Since the objectives of the present study are genecological and not taxonomic, objectives 1 to 3 are of much greater importance than objectives 1; to 5 . The value of the principal component analysis in the present inst-ance, therefore, is that i t enables the investigator to identify, from the many variables measured, those variables which contribute most to the total pattern of variation. Consequently the identification of variables of greatest genecological significance is facilitated. Multiple regression analysis was used to determine the degree to which the variation observed is related to environmental factors at the place of origin of each provenance. In this way factors which exercise selection pressure on the species may be shown to have association with measurable geographic factors such as altitude, latitude or longitude. Because of the extreme paucity of climatic information for the montane, and particularly the subalpine forest regions of British Columbia, an attempt was made to calculate the number of days in the growing season at the place of origin of each of the 150 spruce provenances from the known data of altitude and latitude. 1 Data concerning forest regions in British Columbia, climatic data, and data concerning photoperiod have been obtained from the following sources: Rowe, J.S. 1959. Forest regions of Canada. Bulletin 123, Canada Dept. of Northern Affairs and National Resources Forestry Branch. 71 pp. 191,6. Tables of sunrise, sunset and twilight. Supplement to the American Ephemeris. Nautical Almanac office, United States Naval Observatory. Climates of the states for, Washington, Idaho and Montana. United States Department of Commerce, Weather Bureau. Climate of British Columbia annual reports. Department of Agriculture, British Columbia. Climatic summaries for selected meteorologicalstations in Canada: Frost data. Meteorological Division, Canada Department of Transport. 3k The altitude, latitude and number of days in the growing season were obtained for each of kSl weather stations representing a wide altitu-dinal and latitudinal range i n British Columbia and adjacent territories in the Yukon, Alberta, Idaho, Washington and Montana. These data were sub-jected to;regression analysis, the independent variables being altitude and latitude, and the dependent variable being the number of days in the growing season. Using the regression equation thus obtained, and by inserting altitude and latitude at the place of origin of each df the 150 provenances, i t was possible to obtain a measure of the number of days in the growing season at the place of origin of the same provenances. This measure is henceforth referred to as the index of the vegetative period. The index of the vegetative period calculated in the manner des-cribed above has a relative value only. Nevertheless i t is based on a body of data much more extensive than is obtainable in British Columbia alone, though at the same time obtained from territories on the periphery of this province. Furthermore the results obtained are in approximate agreement with results published in regard to European countries, in that they indicate that a displacement of one degree north shortens the growing season by 6 days, and that a displacement of 300 f t . upwards will have a similar effect (Wiersma 1963)• This index of the vegetative period i s , therefore, used in the present study, since even as a relative value i t has more biological sig- -nificance than altitude or latitude. Illus. 1. Upper photo gives general view of the layout of the nursery beds at Cowichan. Lower photo shows layout of seedlings in these beds. Tllus. 2. The X-ray assessment of embryo development in spruce seed. A. Diagrammatic representation of spruce embryo classes and insect infested seed: a - seed coat; b - cavity between seed coat and endosperm; c - endosperm; d - cotyledons; e - embryo cavity; f -radicle. B. Photographic representation (obtained from radiographs) of spruce embryo classes and insect infested seed corresponding to those illustrated in A. Note the polyembryonic condition of class II, O N TABLE 2' MEASUREMENTS OF SEED QUALITY, GERMINATION, AND GROWTH MADE ON 150 SPRUCE. PROVENANCES, AND MEASUREMENTS OF FACTORS OF THE ENVIRONMENT AT THEIR PLACE_OF ORIGIN. MEASUREMENT 1 2 3 h 5 6 7 8 9 10 11 12 13 lh A 5 _ 16 17" -18 19 20 21 22 CODE Altitude Latitude . . . . . . . . . . . Index of the vegetative period Day length of the first day of the year with average temperature of li3°C Percent of embryo development class III in each seed lot . . . . . Percent of embryo development class IV in each seed lot. . • • . . The rate of germination at 15, 20, 25, 30°C'» Mean daily germination at 15, 20, 25, 30°C . . . . . . . Actual germination percent . . . . . . . . . . . . . . . . . . . . Germination value (PVXMDG) at 15, 20, 25, 30°C . Rate of germination at 25°C. adjusted for embryo development . . . Germination value at 25°C. adjusted for embryo development • . . » Shoot length in fi r s t year . . . . . . . . . . . . . . . . . . . . Root collar diameter in fi r s t year . . . . . . . . . . . . . . . . Dry weight in fi r s t year . . . . . . . . . . A. L D DLl»3 ED3 EDl; PV15, PV20, PV25, PV30 MDQ15, MDG20, MDG25„ MDG30 AGP GV15, GV20, GV25, GV30 PVA25 GVA25 SL1 RCD1 DW1 Shoot' length in second year. .:. ....... .'. . Root collar diameter in second year. . . . . . . : . . . Dry weight in second year. . . . . . . . . . . . . . . . Percent flushed on April 6, l l i , 2 0 , 27 in 2nd year . • . Percent Dormant on June 3 0 , July 7, lit, 2 1 , 2 8 , August Shoot extension May 10, 2U, June 6, 20, July k> 18 during the second year. < » . . . . . . . . . . . . . < > ...< Shoot extension May 10, 2k, June 6, 20, July U, 18 during the second year expressed as a percent of total growth .....< SL2 ..RCD2 DW2 F L 6 , FLLU, F L 2 0 , FL27 DJ30, DJY7» DJYllt,, DJY21, DJY28, DAii. SMIO, SM2U, SJ6, SJ20, SJYU, SJY18. MLO, PM2ii, PJ6, PJ20, PJYU, PJY18. V J O - 0 TABLE 3 GEOGRAPHIC ORIGIN OF 150 SPRUCE PROVENANCES ARRANGED IN ORDER OF INCREASING ELEVATION 13 1U 15 16 17 18 1 9 20 2 1 22 23 21l 25 2 6 27 26 2 9 30 31 32 33 3U 3 9 3 6 3 7 36 3 ? Uo U l U2 UU US U6 147 U8 1.9 50 51 52 53 51. 55 5 6 5 7 58 59 60 6 1 62 6 3 6 1 6 5 6 6 6 7 68 6 9 73 75 * 96 103 131 132 101 93 101i 105 130 63 125 85 92 61. 116 117 1 2 6 121. 25 62 127 76 77 51 1 3 1 2 1 122 1 0 6 110 6 8 60 1.7 1 6 2 70 71 91 98 9 9 111 5 6 107 108 109 128 55 65 78 79 3 l l 5 1 1 5 7 5 8 5 9 123 1 2 9 1.8 1.9 118 32 75 112 1 1 9 5 2 9 692 71.8 B i o 1.61, 7U2 526 766 767 8 0 9 2 6 1 801 U26 525 2814 792 793 805 800 71.0 905 606 3 7 9 382 U 3 5 8 797 798 769 777 3 5 5 l i 9 779 3 6 7 l i 8 36ti 3 6 5 366 3 6 9 li51i 1.99 716 719 780 71.6 3 5 3 3 9 770 771 772 807 3 7 2 9 6 3 8 9 3 9 0 12 1.93 522 523 U31 50 l i * 370 U3 Ii6 li7 799 606 3 8 5 51 791i 3 8 7 373 781i 795 Let. 53 UO 52 22 50 03 55 05 5 l i 03 l i 9 08 55 05 Ii8 50 53 05 53 30 53 0 6 50 32 5U UO 5U I S 51 11 55 08 55 18 55 28 55 1 7 51 35 51 35 51i 5 1 55 3 6 55 36 53 25 55 26 51i l i 6 S l i U5 51i ID S l i ID S l i H i 55 3 0 51i 0 6 51i 56 55 10 5!t 56 51i 5 6 SU 56 5 l i 5 6 51i 56 S l i 05 51i 11 51i 11 5U 05 5U 0 6 S l i 5 6 5U 05 5 l i 15 SU 15 S l i 10 55 16 51i 05 5U 10 51i 00 5 l i o o S l i 30 S l i 30 52 20 52 25 51i 07 5 6 30 51i 20 55 20 5U 17 5k 05 SU 05 55 02 51. UO SU I S 5U 10 5U 38 5U 22 5U 05 5U 53 SU 07 Long. 132 20 1 2 6 U5 127 03 1 2 9 25 128 3 9 125 25 129 20 12U U5 132 OS 132 15 132 0 3 127 15 128 US 128 US 125 3U 127 2U 127 U2 127 U2 127 2U 119 51 1 1 9 51 127 15 127 5 0 127 50 122 UO 127 50 127 11 127 1 1 122 10 122 10 12 7 22 121 35 122 0 3 127 25 1 2 2 12 127 25 127 25 127 2 5 127 2 5 127 25 122 20 122 07 122 0 7 121 53 122 03 127 25 122 03 122 20 122 2 0 122 1 5 127 1 9 122 0 3 122 10 122 0 0 122 0 0 121, 0 6 122 UO 121 UO 121 25 121 50 121 0 6 122 UO 1 2 6 3 8 122 3 7 122 OS 122 05 1 2 6 30 127 15 122 15 123 10 127 12 122 30 127 15 126 UU 125 2S Elev. i n 1000 f t . Locatioi 0.1 Juskatla, Q. Charlotte I s . 0.1 B e l l a Coola 0.1 Kaouk R. II.W. of Zeballos 0.1 S.W. Aiyansh, Nass River 0.1 ? miles N. of Kitimat 0.1" Kennedy L S.E. of Alberni' 0.1 Nass R., S.W. of aiyansh 0.3 Klanawa R. Vancouver Island 0.3 Pallant Creek, Q.C.I. O.U Gold Creek, Graham I s . O.U Mosquitoe L., Moresby I s . 0.6 West of Port l i c K e i l l 0.7 Kitsura Kalum L., N.of Terrace 0.8 Kitsum Kalum L., N.of Terrace 0. ? K l i n a k l i n i R. Knight Inlet 1.0 Beaumont II. of Smithers 1.1 7 miles N. of Hazelton 1.2 tl. of Hazelton 1.2 Suskwa R., E.of Hazelton 1. U N.Thompson R., Birch I s . I . U N.Thompson r.., Birch I s . 1 .5 Evelyn, II. of Smithers 1.5 Kispiox R. 1.5 Kispiox R. 1.7 Hixon 1.7 20 miles N. of Hazelton 1.9 3.C.F.S.0fflce, Smithers 1.9 Bigelov Lake, Smithers 1.9 Fraser R.,1!.E. of Newlands 2.0 Fraser R,, N. of Giscome 2.0 McBride L., Korice R. 2.0 Peace R. Area 2.0 Aleza Lake Exp.Statlon 2.0 Doughty N. of Smithers 2.0 Pine River, Peace River 2.0 Doughty N. of Smithers 2.0 Doughty N. of Smithers 2.0 Doughty !.'. of Smithers 2.0 Doughty N. of Smithers 2.0 Doughty N. of Smithers 2.0 Fraser R., Giscome 2.0 li.rf. of Aleza Lake 2.0 N.W. of Aleza Lake 2.0 Fraser R., S.of Hansard 2.0 Aleza Lake Exp. Station 2.0 Doughty N. of Smithers 2.1 Aleza Lake Exp. Station 2.2 Fraser I!, of Giscome 2.2 Fraser R., N.of,Giscome 2.2 Fraser R., N.of Giscome 2.2 16 miles H.E. of Hazelton 2.2 Aleza Lake Zxpt* Station 2.5 H.E. of HewLand3 2.2 Aleza Lake 2.2 Aleza Lake 2.2 Fort S t . Janes 2.2 Davie L., N.of Prince George 2.2 Antoine L. E.of Soda Creek 2.2 Horsefly Lake E.of Soda Cr. 2.2 6 miles H.E* of Upper Frasei 2.3 N.W. Fort St. John 2.3 Summit L.,N.of Prince GeorgE 2.3 Fort Babine, Babine Lake 2.3 North of Prince George 2.3 Aleza Lake Exp. Station 2.3 Aleza Lake Exp. Station 2.3 "Smithers Landing, Babine L. 2.5 TelJara R., W.of Telkro 2 .5 N. of Giscome 2.5 Salmon R.,N.W.0f Pr.George 2.5 6 miles S.W, of Telxva 2.6 5 miles N.E.of Summit Lake 2.6 Korice R.,S.W.of Houston 2.7 Chapman L., N.E.of Smithers 2.7 S.E. of Burns Lake Prov. Reg. Kiev. No. No. Ho. Lat. Long. 1000 1 76 1U3 • 6 2 3 . 5 6 10 121 50 2 . 7 77 12 380 55 36 127 50 2.7 78 17 37U 5U 27 1 2 6 5U 2.7 79 6 9 3 5 9 5U OU 121 30 2.7 80 100 723 5U 15 12) 55 2.7 81 7 722 SU OU 121 27 2 . 6 82 113 7C9 SU 21 12S 2 6 2.9 63 115 791 55 05 127 20 2.9 eu 9 3 8 6 53 UO 122 25 3.0 85 18 3 5 6 5U 0 6 127 09 3.0 86 11U 790 5U OU 127 13 3.0 87 53 2U0 50 10 116 00 3.1 • 88 120 796 5U 51 127 00 3.2 ; 8 9 7U 371 SU 0 6 127 12 3 .3 50 U2 773 S3 20 122 10 3.U 91 20 U27 51 U2 120 10 3.5 92 90 U88 U9 OU 120 US 3.5 93 102 7U5 52 12 119 15 3.5 9U 21 U18 51 10 120 15 3.8 95 37 3UU U9 35 115 59 3.8 9 6 1U2 575 SO 00 118 00 3 . 6 97 28 33U U9 OO 116 U5 3.9 98 95 6 9 1 U9 0 7 120 53 u.o 99 11:6 6UU 50 38 116 00 li.O 100 eu U17 51 1U 120 15 U.o 101 31 3U6 U9 55 116 35 U.o 102 23 1U7 U9 10 120 35 U.o 103 22 150 U9 55 120 35 U.o 10U 30 393 U9 55 1 1 9 U6 U . l 105 13U 5 7 6 50 1 5 117 50 U.2 106 135 858 U9 00 116 UO U.2 107 67 U30 50 Uo 119 35 U.2 108 86 • U28 51 UU 120 05 U.2 109 52 725 50 39 117 32 U2 110 33 7U7 U9 20 117 15 U.2 111 26 738 50 U2 1 1 9 30 1.5 112 ' 2U 7U1 50 52 • 119 50 1.5 113 1 3 6 868 U9 03 116 5 8 U.3 11U UO 709 51 0 6 117 05 U.3 115 15 737 51 07 118 15 U.3 116 5U 715 U9 29 117 1U u.u 117 1U8 6 5 5 U9 05 115 U7 U.5 110 138 5 3 9 U9 55 11U U6 U.5 119 67 3U0 50 32 115 55 U.5 120 UU 732 U9 25 118 02 U.5 121 2 9 3li3 U9 U9 116 16 U.5 122 1U1 5U2 U9 55 11U U5 U.6 123 83 U16 50 51 119 52 U.6 12U 61 66 51 UO 120 10 U.6 125 39 3U5 U9 05 116 15 U.6 126 3 6 701 50 03 115 2 6 U.6 127 3U 710 U9 22 11U 35 U.6 1 2 8 1 9 U29 51 U6 120 10 U.6 1 2 9 88 UU7 51 o o 119 52 U.7 130 82 3 9 5 50 0 7 1 1 9 U2 U.7 131 80 392 50 OU 119 UI U.7 132 U3 898 51 0 7 117 15 U.7 133 35 699 U9 10 116 0 6 U.7 13U 1U9 6 5 7 U9 15 116 1U U.8 135 1U5 6U1 50 3U 1 1 9 2 9 U.9 1 3 6 1 3 9 5U0 U9 SS U U 55 U.9 137 US 3 3 8 U9 US 117 00 U.9 138 1UU 6 3 6 U9 U6 115 2U 5.0 1 3 9 137 888 50 05 117 U5 5.0 1U0 133 SU5 U9 03 11U UO 5.0 1U1 6 6 336 50 15 115 US 5.0 1U2 27 720 50 2 9 119 U5 5.1 1U3 97 703 U9 2 6 116 08 5.1 1UU 1U0 SU1 U9 55 11U UU 5.1 1U5 U6 335 U9 20 116 0 8 5.3 1U6 150 857 U9 15 117 US 5.5 1U7 38 3U1 U9 18 116 00 5.5 1U8 81 39U 50 01 119 U2 5.5 1U9 1U7 65U U9 0 6 11U UO 5.6 150 UI 337 U9 35 117 U6 5.7 Location  Nig Creek N. of Fort St.John Kispiox River IJ.W. of Barrett 12 miles N. of S i n c l a i r M i l l s N.E. of Aleua Lake N.E. of S i n c l a i r M i l l e Taltapin L. NJS. of Bums Lake Korice Town Stone Creek S.E. of Prince George S.W. of Houston KcEride L. S.W. of Houston Whatshan Creek, Upper Arrow Lake 12 mife a H.E. of Smithere Korice R. S.W, of Telkva tfaverahbu Rd. Prince George H. of Clearwater Manning Park Thunder R., N.E. of Blue R. S.W. of Barriere Perry Cr.,N.W. of Cranbrook Shelter Bay Upper Arrow L. Boundary Cr.,S.W. of Creston Gibsor. Pass Manning Park Upper Arrow Lake S.of Arrowhead Peterson Creek W.of Barriere Kettle R., E. of Kelowna Manning Park S.E. of K e r r i t t W. of Kelowna •Upper Arrow L.,S.W. of Hakusp Boundary Cr. S. of Creston Fl y H i l l s , Salmon Arm N. of Clearwater Trout Lake S. of Nelson F l y H i l l s , W. Salmon Arm 6 miles S.W. of HcGillvray L. Konk-nun Creek S.W.of Cre3ton Spillimacheen R.,S.E.of Golden Big Bend, Columbia R. Clear Creek, Nelson Spruce Tree Cr.,Tahk Valley Line Creek, N.of Natal Shushwap Creek, Radium Deer Park,Lower Arrow Lake St.Hary's R., W.of Kimberley Line Creek, N. of Natal 2 miles W. of McSillvray Lake Clearwater Boundary Creek,S,W. of Creston E. of Canal Flats S.E. of Fernie 'I. of Clearwater HcGillvray Lake H.W. of Kelowna H.W. of Kelowna S.W. of Golden S o f Cranbrook Kidd Creek S.W. of Cranbrook Spa Lake V/. of Salmon Arm North Fork Pas3, N. of Natal Cedar Creek S. of Kaslo Sumner L, S.E.of Skookemchuck McDonald Cr. South of Nakusp Gouldrey Creek W. of Flathead Upper Dry Creek,Columbia Lake S.E. of Westwold Perry Creek W. of Cranbrook North Fork Pass N. of Natal Koyie R. S.W. of Cranbrook S.W. of Castlegar Rabbit Ft.Cr.,S.W.of Cranbrook N.W. of Kelowna Cabin Creek W. of Flathead N. of Castlegar Lat. - latitudej Long. - longitude; Elev. - elevation TABLE k (GEOGRAPHIC ORIGIN OF TWELVE PROVENANCES SOWN IN THE .COWICHAN LAKE RESEARCH NURSERY E3BV. in No. RN Lat. Long. 1000 f t . Locality 12 779 5U°06' 122°03' 2.0 Aleza Lake 2 522 52°20' 121°U0! 2.2 Summit. Lake 8 UO 5U°30! 122°U0-? 2.3 Antoine. Lake, E. of Soda Creek k 387 5U°22I 122°30* 2.6 5 Miles N.E. of Summit Lake 6 388 53°U0I 122°25! 3.0 Stone Creek S.E. of Prince George 7 773 53°20-» 122°10J 3»k Naverabu Rd. Prince George 1 33U U9°00? 116°1|5! 1.0 Boundary Creek S.W. of Creston 3 3h3 h7°k9> 116016* 1*.5 West of Kimberly 9 702 UP^M 115°58? Iu7 Whitney Creek 10 338 U90U5' 117°00l U.8 Cedar Creek S. of Kaslo 11 335 U9°20? Il6°08i! 5.3 Moyie River S.¥. of Cranbrook 5 337 li9°35? 5.7 N. of Castlegar 1 Registration number PART B During the summer of I963 sampling was confined to the white-Engelmann spruce complex, and 57 samples were made. During the summer of 196U sampling was extended north to the Yukon border and west to the Coast; 100 areas were samples. The I96U sample, therefore, embraces elements of both black and Sitka spruce as well as white and Engelmann. Cones were collected from the ground at the base of each of 5 trees, which were at least 100 feet apart, in a l l sampled areas. To avoid contamina-tion from neighbouring trees, care was taken to collect cones as close as possible to the bole of the selected tree. At least 20 cones, which were kept separate by trees, were taken from the base of each tree at each sampled area. This gave a minimum of 100 cones per sampled area. General ecological observations were made at each area visited, and any notable variation in gross morphology of the trees sampled was noted and permanently recorded by means of photographs. In areas where field observa-tions indicated the presence of sympatric populations,sampling was more in-tensive and conducted along transects which passed from one broad ecological zone to another. Foliage samples were also taken in these areas. Table 5 gives the geographic origin of the 157 cone samples. In the laboratory a single cone scale and bract were taken from the midpoint of each of 100 cones per sampled area, and photographed to-gether with a ruler scaled in mm. (Illus. 3)« A l l subsequent measurements were made from these photographs utilizing the mm. scale cut from the photo-graph. A desk magnifier was used to facilitate measurement. Five basic measurements were made on each cone scale and bract and transferred directly to computor data sheets. Five further measurements were then derived from these basic measurements (Fig. l ) . A l l statistical analysis were carried out by electronic computor. ill Miscellaneous cone samples were supplied by the reforestation Division of the B.C. Forest service, by personnel of the Research Division and also by the Columbia Cellulose Company. These samples were measured in the manner described above, but the resulting data were not included in the main analysis. In order to clarify the complex pattern of variation in cone scale morphology, and to facilitate interpretation, recourse was made to extensive graphing of data. A modified form of the "line of shape" method developed by Jentys-Szaferowa (1959) was used, and is explained below. From the total number of trees sampled, 5 trees were selected from allopatric zones of white, Engelmann and Sitka spruce. Since no allopatric zone of black spruce was sampled i t was necessary to select individuals known to be pure black spruce trees from different sympatric zones of white and black spruce. These trees, representing the pure form of each of the four spruces occurring in British Columbia, are henceforth referred to as standard samples. The cone scale samples of these trees were measured in the manner already described, and mean measurements obtained for each of the four species in its pure form (Table 6 ) . Table 7 is derived from Table 6 and the figures in column 3 of Table 7 may be graphed in such a way as to allow a visual assessment of the line of shape of one species in relation to another (Fig. 2 ) . In Fig. 2:1 the straight line represents white spruce and is given the. numerical symbol 1. The angular line represents Sitka spruce, and the figures on the horizontal line represent the number of times each measurement of the Sitka spruce scale deviates positively or negatively from the corresponding measurement of the white spruce scale (See Table 7). The angular line, therefore, is the characteristic curve obtained when pure Sitka spruce is graphed on pure white spruce. The figures on the vertical line represent the 10 measurements of cone scale morphology illustrated in Fig. 1. The other species are similarly represented in Fig. 3. When a cone scale sample is from a sympatric zone, and is doubtful!' or unknown as to species - the rule rather than the exception in respect of such samples - i t can be graphed against the standard samples, using the means for the species represented in the sympatric zone. In this way i t is possible to determine the degree to X i r h i c h the unknown sample represents a known standard sample, and to assess the degree of putative hybridization occurring in the sympatric zone. A large number of samples have been graphed in this way. For example, the sampled area is represented by 5 trees, and i f the area is a sympatric zone of white and black spruce, each tree is graphed against standard samples of white and black, and on the same figure the characteristics curves of white on black,, and black on white (curves 11 and 12 on each figure) are also given (Figs. 35 to kk). In addition to the line of shape method of discriminating species and intermediate forms in sympatric zones, a l l cone scale data resulting from the cone collections made in 1963 and 196U were subjected to dis-criminant function analysis. Jeffers (1965) has discussed the uses of discriminant function analysis in the study of variation. In this same publication he lists the relevant literature. In the present instance the principal objective of the discriminant function analysis is to indicate sympatric and allopatric zones of white and Engelmann spruce, white and Sitka spruce, and white and black spruce, on the basis of variation in cone scale morphology. If for any sample of 5 trees the analysis indicates that two spruce species are present, then that sample is classed as sympatric for these species even though i t contains say four trees of one species and only one of another. The sample is classed as allopatric i f a l l five trees are of the same species according to the analysis. TABLE 5 GEOGRAPHIC ORIGIN OF 157 SPRUCE CONE SAMPLES ARRANGED IN ORDER OF INCREASING ELEVATION Sample E l e r . In 1 Sample Elevation in No. No. Lat. Long. 1000 f t . Location Ho. No. Lat. Long. 1000 f t . Location 1 129 SU 1U 128 32 0.3 9 Bil e s N. of Kltlmat 80 19 51 OU 119 56 3.0 UO miles N. of Kamloops 2 128 SU 15 129 27 0.3 55 m. E. of Prince Rupert 81 1U9 56 25 121 0 9 3.0 K i l e 6U.5 Alaska Highway 3 126 SU 15 129 U8 O.U 32 miles E. of Prince Rupert 82 78 5U UU 127 16 3.0 9.5 miles S.W. of Smithers U 127 5U 13 130 16 O.U 13 miles S.E. of Prince Rupert 83 136 SU 27 126 US 3.1 32 miles E. of Smithers 5 125 5U 15 130 8 0.5 1U miles E. of Prince Rupert 8U 1U8 56 UU 121 55 3.2 Mile 1 0 3 . 5 Alaska Highway 6 121 55 10 129 15 0.5 70 miles N.W. of Terrace 85 56 53 UO 122 25 3.2 30 miles S.E. of Prince George 7 119 55 2 129 33 0.5 89 miles N.W. of Terrace 86 1U1 5 9 16 129 37 3.2 152 miles N.E. of Telegraph Cr. 8 131 5U 35 128 23 0.5 1 2 . 5 miles E* of Terrace 87 79 5U U5 127 17 3.2 10 miles S.W. of Smithers 9 120 55 3 129 30 0.5 8U miles N.W. of Terrace 88 1UU 56 UU 12U 55 3.3 Mile U02 Alaska Highway 10 130 5U 3 7 128 U2 0.6 18 miles N. of Terrace 89 l l S 50 5U 119 U9 3.3 Ul miles N.E. of Kamloops 11 132 SU 5 7 128 23 0.7 U3 miles E. of Terrace 90 2U 50 55 120 25 3.3 30 miles N.W. of Kamloops 12 12U 55 16 129 02 0.7 70 miles N. of Terrace 91 SO U9 11 120 36 3.3 31 miles S.W. of Princeton 13 133 55 2 128 16 0.8 58 miles E. of Terrace 92 63 50 00 120 36 3.3 13 miles S. of Merrltt 111 13U 55 7 127 U6 1.1 80 miles E. of Terrace 93 85 SU 56 126 3U 3 . U 26 miles E. of Smithers 15 123 55 23 12 B 53 1.2 8U miles N. of Terrace 9U 109 51 30 117 18 3.U 30 miles w. of Golden 16 86 55 12 127 Ul 1.2 U8 miles S.W. of Smithers 95 8 S3 UO 122 25 3 . 5 30 miles S.E. of Prince George 17 135 55 23 127 36 1.2 16 miles N. of Hazelton 96 25 50 55 120 25 3.5 35 miles N.W. of Kanloops 18 31 51 28 120 10 1.3 97 80 SU U6 127 17 3.6 11 miles S.W. of Smithers 19 139 57 55 131 11 1.3 Telegraph Creek S.W. of Watson Lake 98 67 50 U2 119 23 3.6 9.6 miles S.W. of Salmon Arm 20 122 55 28 128 U7 I.U 10U miles 11. of Terrace 99 9 53 16 122 0 8 3.7 UO miles N.E. of Quesnel 21 117 51 35 119 53 I . U 91 mile3 N. of Kamloops 100 26 50 55 120 25 3.7 35 miles N.W. of Kanloops 22 1US 58 37 122 Ul 1.6 29 miles S. of Ft. Nelson 101 58 U9 OU 120 50 3.8 Manning Park 23 1U6 58 12 122 U U - 1.7 Mile 2UU Alaska Highway 102 20 51 03 119 53 3.8 UO miles N. of Kamloops 21i 1U3 5 9 3 9 126 53 1.7 Mile 532 Alaska Highway 103 108 51 21 116 3U 3.8 20 mile3 W. of Lake Louise 25 16 51 08 120 02 1.8 UO miles H. of Kanlobps 10U 1U7 S7 17 122 U5 3.9 K i l e 168.5 Alaska Highway 26 66 52 U6 122 26 1.8 58 miles N. of Williams lake 105 27 50 55 120 25 3.9 35 miles N.W. of Kamloops 27 7U SU SO 127 li> 1.9 6 miles N.W. of Smithers 106 57 53 UO 122 25 3.9 30 miles S.E. of prince George 28 LU SU US 127 08 2.0 10 miles S.E, of Smithers 107 99 U9 33 116 27 3.9 U8 miles N.W. of Cranbrook 29 67 52 55 122 27 2.0 69 miles Ti. of Williams Lake 108 11U 50 52 119 50 3.9 U3 miles N.E. of Kamloops 30 32 51 50 120 03 2.0 100 miles N. of Kamloops 109 60 U9 06 120 50 U.0 Harming Park 31 3 SU 06 122 OU 2.0 UO miles N.E. of Prince George 110 81 SU U7 127 17 U.0 12 miles S.W. of Smithers 32 U SU 02 121 UO 2.0 50 miles N.E. of Prince George 111 37 51 50 120 03 U.0 ICO miles N. cf Kamloops 33 69 5U 06 122 23 2.0 2U miles E. of Prince George 112 22 50 UU 119 11 U.0 10 miles N.E. of Salmon Arm 3U 1U2 59 58 128 09 2.1 30 miles S.E. of Watson Lake 113 59 U9 0 3 120 51 U .o Manning Park 35 51 53 UO 122 25 2 J 30 miles S.E. of Prince George 11U 88 50 U2 119 23 U.o 10.5 milea S.W. of Salmon Arm 3 6 151 55 37 122 0 0 2 ,1 .2.1 miles W. of Chetwynd 11? 105 SO U8 116 00 U . o 21 miles E. of Radium 3 7 17 51 10 120 0 5 2.1 50 miles N. of Kanloops 116 91 ur 6 116 5U U.l 5 9 miles S. of Nelson 38 65 52 21 122 20 2.2 2U miles N. of Williams Lake 117 UO 50 08 118 3U U.l UO miles E. of Vernon 39 12 53 00 122 30 2.2 2 miles N. of Quesnel 118 28 SO 55 120 25 U.l 35 miles N.W. of Kamloops 10 75 SU 50 127 15 2.2 6.5 miles N.W. of Smithers 119 62 5U U7 127 17 U.2 12.5 miles S.W. of Smithers Ul 15 5U 32 125 U5 2.2 15 milea N. of Burns Lake 120 1SS 53 OU 121 31 U.2 55 miles E. of Quesnel U2 153 SU 3 9 122 U3 2.2 150 miles S. of Chetwynd 121 106 51 0 2 115 SB U.3 39 miles E. of Radium U3 68 5U 6 122 U 2.2 Aleza Lake Nursery 122 29 50 55 120 25 U.3 35 miles N.W. of Kamloops UU 71 5U 1 12U 6 2.3 U miles W. of Vanderhoof 123 96 U9 6 116 56 U.3 55 miles S.E. of Nelson U5 156 5U 18 122 37 2.3 35 miles N. of Prince George 12U 113 50 50 119 50 U.3 U5 mile3 N.E. of Kamloope U6 7 SU 00 122 30 2.3 20 miles N.E. of Prince George 125 69 50 U2 119 2U u.u 11.6 miles S.W. of 5almon Arm U7 10 S3 U9 122 U7 2.3 10 miles S.W. of Prince George 126 ne 51 33 119 UU u.u 105 miles N. of Kamloops U8 52 S3 67 122 25 2.3 30 miles S.E. of Prince George 127 93 U9 UU 117 09 u.u 17.5 miles N.E. of Nelson U9 15U SU IB 122 36 2.3 3 5 . 5 miles N. of Prince George 128 102 U9 UU 115 31 U.5 25 miles N.E. of Cranbrook SO lllO 58 1U 130 38 2.3 3 9 miles N.E. of Telegraph Crk. 129 l d l i 50 3 9 116 00 U.S 5 milea E. of Radium 51 U7 U9 17 117 13 2 . U 20 miles 3 . of Nelson 130 36 51 50 120 0 3 U.S 100 miles N. of Kamloops 52 S 5U 20 122 37 2.U 30 miles N. of Prince George 131 U3 U9 03 117 00 U.6 UO miles E. of T r a i l S3 13 SU 26 12U 21 2.U 3 miles W. of Ft. S t . James 132 103 U9 U7 115 27 U.6 6U miles N.E. of Cranbrook SU au 55 3 126 30 2.U U7 miles E. of Smithers 133 21 51 00 119 50 U.6 UO miles N. of Kamloops 55 1S7 S3 US 122 52 2.U 1U miles W. of Prince George 13U 92 U9 UU 117 09 U.7 18 mlleo N.E. of Nelson 56 152 55 21 122 36 2.U 7U miles S. of Chetwynd 135 35 51 50 120 03 U.7 100 miles N. of Kamloops 57 73 SU 13 125 3 9 2.5 65 miles w. of Vanderhoof 136 23 50 U6 119 27 U.7 10 miles N.W. of Salmon Arm 58 76 5U UU 127 11 2.5 5 miles S.W. of Smithers 137 30 50 55 120 25 U.7 35 milea N.W. of Kamloops 5 9 53 53 UO 122 25 2.5 30 miles S.E. of Prince George 138 UB U9 12 118 00 U.7 20 miles N.W. of T r a i l 60 1 53 us 122 30 2.5 20 miles S.E. of Prince George 139 116 50 33 119 3D U.7 55 miles S.E. of Kamloops 61 11 53 Ul 122 55 2.5 20 miles S. of Prince George 1U0 110 50 5 9 119 33 U.7 5S miles H.E. of Kamloops 62 16 U9 02 117 10 2.5 UO miles W, of Creston 1U1 112 50 U6 119 52 U.8 U8 miles N.E. of Kamloops 63 61 U9 1U 121 0 6 2.5 Manning Park 1U2 62 U9 5 120 U6 U.8 Manning Park 6U 72 5U 0 3 12U U2 2.5 31 miles W. of Vanderhoof 1U3 90 50 U2 119 25 U.8 12.9 miles S.W. of Salmon Arm 65 33 51 50 120 03 2.6 100 miles H. of K&mloops 1UU 3U 51 50 120 03 U.9 100 miles N. of Kamloops 66 6U 52 12 122 06 2.7 6 miles N. of Williams Lake l i s 95 U9 6 116 58 U.9 53 miles S.E. of Nelson 67 70 53 5U 123 26 2.7 Ul miles W. of Prince George 1U6 101 U9 17 115 58 U.9 25 miles S.W. of Cranbrook 68 SU 53 UO 122 25 2.7 30 miles S.E. of Prince George 1U7 96 U9 2U 116 10 S . l 33 miles N.W. of Cranbrook 69 2 53 U5 122 15 2.7 30 miles S.E. of Prince George 1U8 U2 U9 0 3 117 00 5.1 UO miles E. of T r a i l 70 150 55 U5 120 38 2.8 19 miles w. of Dawson Creek 1U9 US U9 06 116 58 5.1 30 miles W. of Creston 71 18 51 OS 120 00 2.8 35 miles N. of Kamloops ISO 3 9 50 05 119 UO 5.1 UO miles S.W. of Vernon 72 U9 U9 0 7 118 35 2.8 UO miles W. of T r a i l 151 111 50 U7 119 52 5.2 52 miles N.E. of Kamloops 73 77 5U UU 127 12 2.8 5.8 miles S.W. of Smithers 152 UU U9 07 116 5U 5.U 25 miles W. of Creston 7U 137 SU 29 126 15 2.8 67 miles E. of Smithers 153 9U U9 6 117 1 5.U 51 miles S.E. of Nelson 75 138 5 9 U8 129 08 2.9 US miles S.W. of Watson Lake 15U 107 51 12 116 0 5 5.5 58 miles N.E. of Radium 76 83 SU US 127 02 2.9 10.5 miles £. of Smithers 155 100 U9 20 116 06 5.6 2ti miles S.W. of Cranbrook 77 38 51 SO 120 03 2 .9 100 miles N. of Kamloops 156 Ul U9 15 117 22 5.8 25 miles S. of Nelson 78 55 53 UO 122 25 2.9 30 miles S.E. of Prince George 157 97 U9 29 116 08 6.7 25 miles N.W. of Cranbrook — 7 5 — 6 -5U- 0 7 _ -121_UU_ 3.0 60 miles N.E. of Prince George Fig. 1 Diagrammatic representation of spruce scale and bract showing five basic measurements. The five derived measurements are L1/L2, L1/L3, Ll/Wl, LL/W2, and 11/12 X L3. The order of occurrence of these 10 measurements in.all subsequent figures and tables is as follows: 1 2 3 ii 5 6 7 8 9 10 Ll L2 :L3 WI W2 L1/L2 L1/L3 Ll/Wl Ll/W2: L1/L2 X L3 U6 ffffff 'ft ft ft ft 'ft 'I'* *f f ft ftft t't ft ft ftffft ft ft f tftfftff* f ft'#f fft *f ii'f'i'i'fH'ri »t't ' f t i f i f <«i# >f i f i f i f i f ' t ' t • f t t f i # >#•# i # i t 't 't •fi< l f t i f i f l » i V »t»f ' r t ft ' t i * i f i f ' f t f i f » • ft t ' # t ' t 'ft ft ft 'tf ft '•'•'ft 'ft 'ft ft •ft 'ft ft 'ft 'ft 'ft'ft 'ft 'ft 'ft ft ft ft 'ft ft ft 'tt ft ft ft ft ft ft ft ft ft' ft't'tftf t f f f t'tf t'tf t ft ft ft t't ft ft ft t 9 9 f ft 'ft 'ft ft tt ff ft ft f t ft ft ft ft ft ft ft 'ft ft ft ft ft ft ft ft tt tt ft ft ft ft ft 'ft ft ft ft Illus. 3« Gone scale morphology of Engelmann (1) Sitka (2) white (3) and black (U) spruce. The first column of scales in k are from a white spruce tree. The remainder are black spruce. Each column of 20 scales and bracts represents a single tree. TABLE 6 CONE SCALE MBRPHOLOGY OF BLACK, WHITE, ENGELMANN AND SITKA SPRUCE s P L 3 W^  w2> L1/L2 L l/L3 h/^ V ^ x l s 138-3 1.02 0.82 0.32 0.86 0.76 1.24 3.27 1.19 1.36 0.39 143-3 1.09 0.82 0.30 0.92 0.75 1.33 3,67 1.19 1.46 O.4O w I44-I O.84 0.68 0.22 0.68 0.60 ' I.24 3.81 1.25 1.41 0.28 o CP •11)5-5 1.13 0 .83 0.31 0.97 0.80 1.37 3.80 1.17 O.42-I46-4 1.12 0 .79 0.24 1.00 0.81 1.43 U.76 1.12 1.40 0.35 X 1.0U 0 . 7 9 0.28 0.89 0.74 1.32 3,86 1.18 1.42 0.37 64-I 1.16 0 . 9 9 0.1)5 0.91 0.82 1.18 2.63 1.29 1.42- 0.53 65-1 1.16 0.99 0.43 O.89 0.79 1.18 2.78 1.32 1.48 0.51 70-1 1.17 1.01 O.46 0.93 0.82 1.17 2.60 1.27 1 . 4 U 0.5U 1 : 71-1 1.11 O.98 0.45 O.84 0.77 1.14 2.52 1.33 1^ 5 0.51 72-1 1.20 1.00 0.42 1.05 0.92 1.20 2.96 1.14 1.32 0.50 X 1.16 0.99 0.44 0.92 0.82 1.17 2.7O 1.27 1.42 0.52 93-1 1.21 0 .86 0.59 0.92 O.67 1.41 2.09 1.32 1.92 0.82 95-1 1.32 1.00 0.54 0.88 0.62 1.32 2.49 1.49 2.16 0.72 1 99-1 1.51 1.15 O.64 0.97 0 .59 1.32 2.37 1.56 2.60 O.84 S2 116-1 1.25 0.92 o.5U 0.88 0.61 1.37 2.35 1.44 2.12 0.74 118-1 1.34 0.99 0.59 0.83 0.57 1.35 2.28 1.61 2.39 0.80 X 1 .33 1.00 0.58 0.90 0.61 1.35 2.32 1.1(6 2.24 0.78 119-1 1.22 1.00 0.82 0.62 0.52 1.23 1.51 1.97 2.35 1.00 120-1 1,1*8 1.06 0.87 O.76 0.51 I.4O 1.73 1.97 2.97 1,21 125-1 1.25 0.90 0.73 0.80 0.57 l . l j l 1.74 1.58 2.23 1.02 w M CO 128-1 1.62 1.11 O.98 O.89 0.52 I.46 1.66 1.83 3.15 1.43 129-1 1 .57 l . l l i 0.83 0.71 0.50 I.38 1.92 2.21 3.16 l . l U X 1.43 I.O4 0.85 O.76 0.52 1.38 1.71 1.91 2.77 1.16 S - species, P - provenance, Li - etc. - measurements as indicated in f i g . 1. TABLE 7 COMPARISON OF MEANS OF CONE SCALE MORPHOLOGY OF WHITE AMD SITKA SPRUCE 1 2 35 M WHITE SITKA SITKA/WHITE 1^ 1.16 1 1 . 2 3 : 1 % 0.99 1.0U 1.05 i 1 L3 O.UU 0.85 1.93 : 1 W^  0.92 0.76 0.83 :. 1 W2, 0.82 0.52 0.63 : 1 1.17 1.38 1.18 : 1 I^/L^ 2.70 1.71 0.63 : 1 1 LJ/W-L 1.27 1.91 1.50 L L / ^ 1.1(2 2.77 1.95 :• 1 LJL/LO; x L 3 0.52 1.16 2.2 :1 Column 3 represents the number of times each of the 10 measurements for Sitka deviates positively or negatively from the corresponding measurement for white spruce. M - measurements illustrated in f i g . 1. U9 Fig. 2 Characteristic curves obtained when cone scale morphology of popu-lations from allopatric zones of white, Engelmann and Sitka spruce is compared by line of shape method (see page Ul for further ex-planation) • 0 . 6 1.0 1.5 2 . 0 2.3 0 . 4 . 1.0 1.6 I 1 1 I I I I I 0 . 6 1.0 1.5 2.0 2.3 0 .4 1.0 1.6 The numbers 1 to 10 on the vertical bars represent the 10 cone scale measure-ments illustrated in f i g . 1. The numbers on the horizontal bars represent the number of times each measurement on one species deviates positively or negatively from the corresponding measurement on the species with which i t is being compared. So Fig. 3 Characteristic curves obtained when cone scale morphology of popu-lation from allopatric zones of white and Engelmann spruce is compared by line of shape method. The black spruce sample is not from an allopatric zone. (See page U l for further explanation). 0 . 6 1.0 1.6 0 . 6 1.0 I.S I 1 1 I I I I 1 1 I 1 1 0 6 1.0 1.6 0 . 6 1.0 1 5 The numbers 1 to 10 on the vertical bars represent the 10 cone scale measu-rements illustrated in f i g . 1. The numbers on the horizontal bars repre-sent the number of times each measurement on one species deviates positively or negatively from the corresponding measurement on the species with which i t is being compared. RESULTS PART A , GEOGRAPHIC VARIATION IN IMMATURE POPULATIONS OF WHITE SPRUCE GERMINATION AND GROWTH BEHAVIOUR OF 150 SPRUCE PROVENANCES SOWN IN THE SPRING OF 1 9 6 5 . There is no significant correlation between germination behaviour at 15°C. and the indices of seed quality ED4 and ED3 + E D 4 , but the index of seed quality ED3 and germination behaviour are significantly correlated at this temperature (see Table 2 for definition of ED3 and EDlt) • At 20°, 25° and 30°C. there is no significant correlation between the index of seed quality ED3 and germination behaviour, but germination be-haviour and the indices of seed quality ED4 and ED3 + EDU are highly corre-lated. The highest correlation between germination behaviour and seed qua-l i t y occurs at 25°C, and is between AGP (actual germination percent) and ED3 + E D 4 (Tables Ik to 17). Germination behaviour is correlated positively with altitude and negatively with the index of the vegetative period. The correlation is 1 strongest at extreme temperatures, that i s , at 15° and 30^3, and weakest at intermediate temperatures, that is 20° and 2 5 ° C The latter temperatures also result i n maximum germinative values (Table 18). Germination behaviour is also significantly correlated with l a t i -tude, but only at extreme temperatures. It is not significantly correlated with latitude at 20° and 25°C. However, when germination behaviour at 2 5 ° C is corrected for embryo development, the correlation between germination behaviour at this temperature and factors of the environment is increased, and rate of germination (PV) at this temperature is then significantly correlated with latitude (Table 1 8 ) • 52 Al l measures of total growth, except root collar diameter in the fir s t year, are highly correlated with altitude and the index of the vege-tative period both in the f i r s t and second year; negatively with altitude and positively with the index of the vegetative period. Of the fir s t year's measurements, root collar diameter (RCD) and the ratio of shoot length (SL) and root collar diameter (SL/RCD) are significantly correlated with latitude; RCD negatively, and SL/RCD positively (Table 21). The percent of seedlings flushed on April 6 in the second year, which is the fi r s t date on which flushing was assessed, is not correlated with any factor of the environment. However, percent flushed on April lu, and April 20 is correlated with a l l three factors of the environment, positively with latitude and the index of the vegetative period, and negatively with al-titude. Percent flushed on April 27 is not correlated with latitude, but is positively correlated with the index of the vegetative period and negatively with altitude. Therefore, low elevation provenances flushed before high elevation provenances (see Table 19)• The correlation between flushing and factors of the environment is much weaker than the correlation between dormancy and factors of the environment (Fig. 19)• Of the other second year's measurements SL/RCD, percent dormant on June 30, July 7, Iii, 21, are significantly correlated with latitude; SL/RCD positively, and percent dormant on a l l four dates negatively. Percent dormant on June 30, July 7, Iii, 21, 28, and August h is very highly corre-lated with altitude and the index of the vegetative period, positively with altitude and negatively with the index of the vegetative period. The highest correlation between any one measurement of growth behaviour and factors of the environment is that between percent dormant on July llj. and altitude. 53 The c o r r e l a t i o n c o e f f i c i e n t i n t h i s instance i s 0.862 (Table 21 and F i g . 8). There i s a high c o r r e l a t i o n between date of entering dormanoy and t o t a l growth of seedlings. Those seedlings whioh were the f i r s t t o enter dormanoy i n midsummer had the lowest dry weight. The c o r r e l a t i o n c o e f f i c i e n t f o r dry weight and percent dormant on J u l y 28 i s - 0.80 (Table 2 l ) . There i s a weak c o r r e l a t i o n between growth during the second growing season and l a t i t u d e . However, growth i s h i g h l y c o r r e l a t e d w i t h a l t i t u d e and the index of the vegetative period. Growth t o l a y 10, i s p o s i t i v e l y c o r r e l a t e d w i t h a l t i t u d e , and negatively with the index of the vegetative period (Table 20). Growth between May 10 and May 24 i s not c o r r e l a t e d with any f a c t o r of the emrironment, but growth from May 24 to June 6 i s again h i g h l y c o r r e l a t e d with factors of the environment as expressed by a l t i -tude, and the index of the vegetative period. However, at t h i s phase of the growing season the r e l a t i o n s h i p i s reversed, and growth from May 24 to J u l y 6 i s negatively correlated with a l t i t u d e , and p o s i t i v e l y with the index of the vegetative period (Table 20). The strongest c o r r e l a t i o n between growth and f a c t o r s of the environment i s that between the index of the vegetative period and growth on June 20 (Table 20). Percent flushed on the dates assessed i s c o r r e l a t e d with other measurements of growth, but these c o r r e l a t i o n s are considerably weaker than the c o r r e l a t i o n s between dormancy and the same measurements (Table 22). Seventeen of the variables assessed i n the 150 provenances at Cowichan Lake were subjected to p r i n c i p a l component analysis. These variables Sk are listed in Tables 22 and 23, and identified in Table 2. The results of the analysis are given in Tables 22 to 2k inclusive. Table 22 shows the degree to which these 1? variables are correlated. It i s obvious from this table that many of the 17 variables are highly correl-ated, and that a much smaller number of "fundamental dimensions0 (Gardiner and Jeffers 1963) could account for a high percentage of the total variation measured. Table 2k gives the percentage of variation accounted for by each of four new variates. These new variates or components, are based on the matrix of correlation coefficients given in Table 22. The fi r s t component alone accounts for 57.85 percent of the original variation, and a l l four components account for 87 percent of the variation. Table 23 gives the weighting for the original variables in the computed components. From this table i t is possible, to a considerable ex-tent, to assign biological significance to at least the f i r s t two components, which between them account for 72.77 percent of the total variation. Referring to Table 23 i t is clear that measurements of dormancy, and associated measurements of growth (variables 3, 6, l l i , 15, 16, 17 in Table 23) are strongly represented in the fi r s t component and that dormancy is one of the most significant measurements made in illucidating the variation pattern in immature spruce populations. A measurement of flushing is strongly-represented in the second component (variable k in Table 23). The third component is not easily identified, but measurements of germination rate appear to be incorporated in the fourth component. GROWTH BEHAVIOUR OF THE 12 PROVENANCES SOWN IN THE SPRING OF I964. Growth behaviour of the 12 provenances sown in the spring of 1964 in the Cowichan Lake nursery was significantly different in the f i r s t year at the 0.01 level of probability, and there was no interaction between the differential growth response of the 12 provenances and soil type. On the prepared soil the 12 provenances, segregated according to groxrth response, formed two intergrading groups. Group one comprised high-elevation prove-nances from southern latitudes, and group two low elevation provenances from northern latitudes (Fig. 6)• Maximum differences between provenances occurred in the greenhouse on the local soil during the second growing season. Growth rate during the second growing season is illustrated in Fig. 7 , and differences in total growth between high and low elevation provenances in the greenhouse is shown in Illus. 4. Table 13 gives the means of flushing and dormancy measurements made in the second year. 56 Illus, k» The differential growth behaviour of spruce populations from different elevations when grown in a plastic greenhouse on regular nursery so i l , 2 - 3 0 miles northeast of Prince George, elev, 2200 f t , 8 - u0 miles north of Prince George, elev, 2300 f t , 1 - 3 0 miles southeast of Nelson, elev. IjOOO f t . 5 - 2 5 miles northwest of Nelson, elev. 5700 f t . 5 7 Fig. k Variation in embryo development in a number of provenances from two spruce seed crops. Note the uniform quality of the l°6l crop in relation to the heterogeneity of the 1959 crop. A completely occluded circle indicates that the seed has an embryo development of 100$ class IV. (See LTJus- 2) F i g . 5 Relationship between degree of dormancy at the Cowichan Nursery on July 15 and altitude at place of origin of each provenance. Note that high elevation provenances go dormant f i r s t , V J \ then low elevation provenances followed by those from areas of coastal influence, e.g. the Mass and Sheena river basins. 59 30-ELEVATION IN THOUSANDS OF FEET Fig. 6 Relationship between growth of 12 spruce provenances in a uniform environment and elevation at place of origin. Each point on the curve represents the mean of 20 1-year-old seedlings. 1 and 5 ar t i f i c i a l soil mix inside the greenhouse; 2 and 6 regular nursery soil inside the greenhouse; 3 and 7 a r t i f i c i a l soil mix outside the greenhouse; l i and 8 regular nursery soil outside the greenhouse. RELATIONSHIP BETWEEN DORMANCY ON JULY 14 AND ELEVATION AT P L A C E OF ORIGIN OF EACH PROVENANCE o o @0(BaD@—°'0 o ° O (©O 8 0 0 0 0 0 0 0 o o o o o o o o 8 0 o o g o o o O OO o OO o o o 0 0 CTD o°o °© o o 00 o o 0 0 o bo 0 0 o o o o o 2.5 1 0 E L E - V A T l O N \t\ 3.5 10 0 0 Ft °o 0 (D5 Fig. 8 Relationship between altitude at place of origin and degree of dormancy of each provenance on July Ik at the Cowichan nursery. Each point on the curve is the mean of 60 seedlings. A seedling was scored as dormant when the terminal needles of the epycotil were s t i f f and whorled, and a terminal bud visible. j£ It is obvious that a more sophisticated method of assessing dormancy would reveal an even stronger relationship with altitude. Two provenances of uncertain origin have not been included in this graph. TABLE 8 SEED CHARACTERISTICS OF 150 SPRUCE PROVENANCES ARRANGED IN ORDER OF ... INCREASING EMBRYO, DEVELOPMENT 62 Prov. Reg. Elev. in FSP ED SW SA No. No. No. 1000 f t . C III C IV crna. year* 1 133 . 5k5 5 . 0 8 U . 5 U . 3 0 . 3 0 . 1 6 9 2 2 68 355 2 . 0 7 0 . 5 5.0 1U.0 O.120 7 3 75 373 2.6 23.0 3.3 U . 3 o . l 3 ; 7 U 11" 7 9 0 3 . 0 75-3 U.5 16.3 0 . 155 u 5 125 8 0 1 . 0.6 2 0 . 3 0 .3 2 0 . 0 O . I 3 6 U 6 129 8 0 8 2.11 2 8 . 3 1.5 25-3 0 . 1 5 0 " u 7 U 5 6UI k.9 6 9 . 0 3 2 . 8 27.8 0 . I U 9 2 8 137 888 5.0 5 9 . 5 19.8 29.5 0 . 155 ? S Ho 5 U I 5.1 8 6 . 3 3 6 . 8 3 0 . 0 0.196 2 10 72 366 2.0 ' 3 . 3 5 -0 33-0 0 . 1 3 9 7 11 81 •39U 5.5 8 6 . 5 3 3 . 5 3 3 . 0 0.I7U 7 U 127 806 1.5 U . o 3-5 33-8 0 . I 2 9 U 13 128 8 0 7 2.2 3 8 . 5 2.8 35-3 O.lUo U u 66 336 5.0 9 1 . 0 7.8 3 8 . 0 0.177 7 15 • 13U 576 ll.2 U8.0 5.8 3 8 . 0 O . I 9 8 2 16 73 369 2.0 U 9 . 0 2.3 Uo.o 0.117 7 17 85 1.26 0.7 U 7 . 8 0.5 u o . o 0.162 7 18 8 9 8 U.7 8 6 . 3 25-3 U l . o 0 . 1 7 U 2 19 1-2 773 3-k 55.5 3.8 H.3 0.186 U 20 23 U 7 u . o 8 I . 5 8.3 U1.5 0.211 9 21 135 8 5 8 U.2 UU.O 0.5 U1.8 0 . 1 9 U 2 22 12k 8 0 0 1.2 U 7 . 0 2 . 5 *2 .3 0 . U o u 23 u i 337 5-7 90,0 18 .0 U 3 . 0 0.18U 7 2k 18 356 3 . 0 5 6 . 8 9 . 5 U 3 . 0 0.160 7 25 U 1.93 2.2 5 3 . 0 3 . 0 UU.O 0 . 1 6 7 7 26 u 370 2-3 57.0 9.5 Uk.O 0 . 2 0 U 7 27 150 857 . 5.5 68.8 9 . 0 UU.3 0.187 2 28 7U 371 • 3.3 8 5 . 8 17.3 UU.3 0 . I 5 0 7 2 9 . 121 797 1 .9 5 0 . 5 1.0 "5.5 0.111 U 30 71 365 2.0 55-3 1.5 "5.5 0 . 1 7 6 7 31 u u 636 S.O 85.5 22.0 U 7 . 8 0 . 1 9 9 2 32 61 66. U.6 85.3 2 8 . 5 U8.0 0.216 9 33 82 395 U.7 59.5 7.5 U 8 . 3 0 . 1 6 5 7 y> 6U 2BU 0.9 53-3 2.5 U 9 . 0 0 . 1 3 5 8 35 139 5U0 U.9 8 7 . 0 25.3 50.5 0 . 1 9 0 2 36 17 374 2.7 57.5 1.5 50.5 0 . 1 3 U 7 37 U 7 6 5 " 5 . 6 9 k . 5 32.8 51.5 0.202 2 38 U 8 6 5 5 U.5 55.8 1.8 51.5 0 .175 2 39 '5 3 3 8 k.9 7 6 . 3 8.5 52.5 0 . 1 9 5 7 kO 87 U30 U.2 9U.8 2 8 . 0 52.8 0 .172 7 Ul 119 795 2.7 6 2 . 0 7-5 5 3 . 0 0.169 U lis I 3 8 539 k.5 8 8 . 0 30.3 5 U . 0 0.185 2 k3 I 2 0 796 3 - 2 99.0 17.8 5 5 . 0 0 .172 U UU 37 3UU 3 . 8 6 U . 0 2.0 56.3 0.169 7 k5 79 390 2.2 92.3 2.5 56.3 0 . 2 0 2 7 U6 U 9 657 U.8 9 2 . 3 2 8 . 5 57.5 0.218 2 "•7 88 lilt 7 U.7 8 9 . 8 2 3 . 0 57.5 0 . 1 7 2 7 U8 10k 766 0 .3 6 2 . 8 1.3 59.0 O . I 5 8 U l>9 22 150 u . o 6 9 . 8 k-5 59.0 0.197 9 50 70 36 •> 2.0 8 5 . 3 12.8 6 0 . 8 O . I 3 9 7 51 l t l 5k2 U.6 91.0 22.5 61.8 0.188 2 52 20 1.27 3-5 75.0 1.3 6 2 . 3 0.127 7 53 39 . 3k5 U.6 6o.8 9-3 63.O 0 . 2 0 6 7 5k 83 U16 U.6 9 U . 0 18.5 63.3 0.16U 7 55 8U kl7 U.o 6 6 . 5 . 2 . 0 63.3 0 . 1 3 1 7 56 38 3 k l 5.5 8 8 . 5 U.8 6 k . 0 0.186 7 57 122 798 1 .9 78.5 6 . 3 6 U . 5 0.168 <i 58 67 3M0 ".5 8 5 . 0 1 0 . 5 6 5 . 8 0.168 7 59 118 791. 2.5 81.3 3.3 6 6 . 5 0.200 U 6 0 136 868 k -3 75-0 5.5 6 7 . 0 0.186 2 61 111 7 8 0 2 .0 8 0 , 3 1.8 6 7 . 3 0.117 U 62 9 388 3 . 0 8 U . 3 9.8 6 7 . 3 0 . I 8 3 7 63 69 359 • 2.7 8 6 . 0 1 0 . 0 6 8 . 3 0 . U 6 7 6 k 32 387 2 .6 8 6 . 3 12.8 6 8 . 5 O . I 9 6 7 6 5 62 905 I.U 77.0 2.3 68.5 O . I 8 3 2 66 8 353 2 . 0 7 k . 0 2 . 0 68.8 0.166 7 67 108 771 2.2 8 2 . 8 u . o 69.O O . I56 U 68 28 33k 3 . 8 88.8 9 . 0 69.5 0 . I 8 0 7 69 8 0 392 U.7 96.5 16.0 7 0 . 0 0.16U 7 70 1U6 6 UU u . o 7 k . 5 1.5 7 0 . 8 O . I 9 6 2 71 115 791 2 .9 97.8 8 . 0 71.8 0 . 1 9 1 • 72 29 3"«3 U.5 8 3 . 8 " . 3 7 2 . 3 0.173 7 73 12 380 2.7 8 0 . 0 5 . 0 72.3 0.173 7 7" 1 0 ? 772 2.2 8 1 . 0 2.8 7 2 . 5 O . I 3 0 « 75 1.6 335 5 . 3 8 3 . 5 >.5 7 3 - 0 O . I 8 5 7 Prov. Reg. Elev. In FSP EO SW SA No. No. No. 1000 f t . C III C IV fcma. years 76 78 389 2.2 93.0 2.6 7 3 . 3 0 . I 6 6 7 77 126 8 0 5 1.2 81.5 5.5 7 3 . 8 0 . 2 0 2 u 78 19 U29 U.6 98.5 17.8 7 U . 0 0 . I 8 U 7 79 113 789 2.9 9 6 . 0 12.5 7 5 . 5 0.19U U 8 0 3 U2 2.2 7 8 . 0 1.8 75.8 0.209 10 81 77 382 1.5 85.3 5.5 76.3 0.171 7 82 I 3 0 8 0 9 O.U 81.8 2-3 7 6 . 5 0.211 U 83 57 <>i 2 . 3 8 6 . 3 3.8 77.3 0.212 10 8U UU 73? k.5 96.3 1-3 77.5 0.200 5 85 16 367 2.0 92.3 8.0 77.5 0 . U 2 7 86 59 >7 '•3 8 0 . 5 1.5 7 8 . 0 0 . 2 0 6 10 87 60 U9 2 . 0 85.3 2.8 7 8 . 8 0.190 10 88 11 523 2.2 8 2 . 8 2.0 79.0 0.1U2 7 89 110 777 1.9 9 U . 0 U.O 79.8 0.186 u 9 0 76 379 1.5 8 6 . 3 U.5 79.8 0.168 7 91 112 78U 2.7 97.8 11.0 8 0 . 5 0.177 u 92 90 U88 3.5 87.3 U.5 8 0 . 8 0.216 7 93 107 770 2.2 8 9 . 8 2.8 8 0 . 8 0.139 U 9U U 2 575 3.8 86,8 3 . 8 80.8 0.212 2 95 132 U6U 0 . 1 9 I . 8 5.3 61.3 0.19k ? 96 106 7 6 9 1.9 89.3 2.0 8?.o 0 . 1 6 2 u 97 U 3 <23 2.7 9 U . 8 6.5 8 2 . 3 0 . 2 0 0 2 98 U7 779 2 . 0 9 6 . 3 1.8 8 2 . 5 0.182 U 99 86 U28 U.2 96.3 7.8 83.5 0 . 1 7 0 7 100 58 U6 2.3 8 7 . 8 3.3 83.5 0 . 2 6 9 8 101 U8 385 2.5 8 9 . 0 2.0 8 U . 0 0.16U 7 102 9" 529 0.1 9 8 . 8 2-3 8 U . 0 0 . 1 9 5 7 103 15 737 k-3 8 7 . 8 1.0 8U.8 0 . 2 3 3 5 10U 96 6 9 2 o . l 93-0 3.3 8U.8 0.22U 5 105 10 522 2 . 2 91.8 2-3 8U.8 0 . I U 7 7 106 5 U9U 2.3 9 3 . 0 3-5 8 5 . 0 0.168 7 107 • 13 358 1.7 89.5 2.8 8 5 . 0 0.159 7 108 2U 7U1 U.2 ' 92.0 2-3 85.3 0 . 2 3 6 5 109 1 50 2 . 3 90.3 3 . 0 85.3 0.20U 10 110 I 2 3 799 2.3 9 6 . 8 6 . 3 8 5 . 8 0.182 u 111 30 393 U.l 93-3 3-5 8 6 . 5 0.181 7 112 31 3U6 U.O 98.5 U.S 8 6 . 5 0 . 1 9 3 7 113 5 0 U31 2.2 93-3 1.8 8 6 . 5 0 . 1 7 6 7 1 U 89 1.5k 2.0 97.0 6.3 87.O 0 . 1 9 U 7 115 51 Ul 1.7 9 0 . 8 1.5 8 7 . 0 0.22k 10 116 21 U18 3.8 97.3 6.0 87.5 O.Uo 7 117 6 T-6 2 . 0 9 0 . 5 1.8 87.5 0 .192 5 118 36 701 U.6 9 U . 8 1.5 8 7 . 8 0.197 5 119 55 37 2.2 91.3 2.3 8 8 . 3 0 . 2 0 6 10 120 116 792 1.0 9 8 . 5 U.5 8 8 . 5 0.189 u 121 5U 715 u . u 95.5 2.5 88.8 0.212 5 122 117 793 1.1 9 U . 8 3-8 89.3 0.201 U 123 92 525 0.8 93-3 1.8 89.5 O . I 8 5 7 12U 3* 710 U.6 95.3 3-3 9 0 . 0 0.20U 5 125 2 U8 2.0 93.8 2.5 9 0 . 0 O . 2 I 5 8 126 56 39 2.1 95-3 2.8 9 0 . 3 0 . 2 0 5 10 127 91 U99 2.0 97.8 3.8 90.3 0.173 7 128 26 7 3 8 U.2 98.3 1.5 91.3 O .232 5 129 25 7U0 I.U 9 6 . 0 1.6 91.3 0 . 2 3 3 5 130 105 767 0.3 99.3 3-3 91.8 0.2U2 U 131 UO 709 k-3 96.5 1.0 92.0 0.189 5 132 103 7«8 0.1 9U.3 0.8 92.5 0 . 2 0 3 5 133 27 720 5.1 9 8 . 0 2.0 9 2 . 3 0.193 5 13U 53 2U0 3.1 97-8 2.5 93-5 0 .252 8 135 97 703 5.1 9 8 . 8 3-3 93-5 0 . 2 2 0 5 136 35 699 U.7 99.3 1.8 9 3 . 8 0 . 2 1 8 5 ' 137 102 V>i 3.5 98.3 2.8 93.8 0 . 2 0 3 5 138 U 9 51 2.5 9 6 . 8 1.8 93.8 0.175 10 139 52 725 U.2 99.3 3-5 9 U . 0 0 . 2 3 5 5 lUo 33 7U7 U.2 9 8 . 8 1.8 9 U . 0 0 .217 5 U l 7 722 2.6 97.8 1.8 9 U . 0 0.193 5 U 2 63 281 0,U 100.0 3.0 9 U . 0 0 . 2 8 3 8 U 3 131 8U0 0,1 98.8 3.8 9 U . 0 0 .172 U UU 9 8 718 2 . 0 98.8 2.0 9 U . 0 0.187 5 U 5 101 7U2 0.2 97.8 1.8 9U.3 0 . 2 5 6 5 U 6 95 691 u.o 97.8 1.8 9 U . 5 0 .276 5 U 7 65 296 2 . 2 99.3 3 . 0 9 U . 8 0 .26U 9 ' U 8 99 719 2 . 0 97.3 1.3 ;> 95.0 0.171 5 U 9 100 723 2 . 7 9 9 . 3 2 . 0 9 5 . 8 0 . 2 0 U 5 150 93 526 0 .7 9 9 . 0 2 . 3 95.8 0.195 7 -FSP - f u l l seed percent; ED: - embryo development; C III - percent embryo class " " i l l (embryo measures between half and three quarters.embryo cavity); C IV -percent embryo class IV (embryo completely occupies embryo cavity); SA - seed age; SW - seed weight is the average of k replications of 100 seeds. TABLE 9 GROWTH BEHAVIOUR DURING THE FIRST YEAR IN THE NURSERY 63 S.L. R.C.D. Ho. No. L a t . LonK. E U v . D A Y S (c.) (C.I S . L . / f t . C . D . U » l 1 50 5 6 . 5 0 1 2 1 . 1 0 2 - 3 73 6 . 9 •133 5 2 . 9 0 1 .77 2 »8 5 5 - 6 7 1 2 2 . 2 0 2 . 0 85 7 . » . l k 2 5 2 . 2 1 2 . 0 6 3 »2 5 k . 5 0 l 2 k . 2 5 2 . 2 88 7.6 . l k l 5 k . 8 6 1 . 9 9 * * 9 3 5 k . 50 1 2 2 . 6 7 2 . 2 88 7 . 7 . 1 5 0 51.95 2.15 5 119k 5 k . 3 3 1 2 2 . 6 7 2 . 3 87 7 . 8 .15k 5 1 . 1 2 2 . 2 2 6 7k6 5 k . 1 0 1 2 2 . 0 5 2 . 0 95 9 . 0 . I 6 3 5 5 . 3 8 2 . 8 0 ; 722 5 k . 0 7 1 2 1 . k 5 2 . 8 77 8 . 9 . 1 6 0 5 5 . 9 9 2 . 5 9 8 353 5 k . 9 3 1 2 7 . k 2 2 . 0 90 8 . 1 . l k 2 58.17 2 . 0 1 9 388 53 - 6 7 1 2 2 . k 2 3 . 0 75 7 . 9 . 1 5 8 - k 9 . 8 l 2 . 2 9 10 522 52.33 1 2 1 . 6 7 2 . 2 102 7 . 9 .1*3 5 5 . 2 3 2 . 0 k 11 523 5 2 . k 2 1 2 1 . k 2 2 . 2 101 8 . 0 . l k 9 5 k . 1 2 2 . 0 3 12 38O 5 5 . 6 0 1 2 7 . 8 3 2 . 7 70 7.1 . l k 6 k 9 . 3 8 1 . 8 7 13 358 5 5 . * 7 1 2 7 . 8 3 1 .7 93 a.3 . 1 5 2 5 k . 6 9 2 . 1 k I t 370 5 5 . 3 3 1 2 6 . 6 3 2.3 8 0 7 . 8 .15* 5 1 . k 3 2.13 15 737 51 . 1 2 1 1 8 . 2 5 k . 3 62 8 . 5 .17* k 8 . 7 9 2 . 6 k 16 367 5 k . 9 3 1 2 7 . k 2 2 . 0 9 0 6 . 5 .123 5 0 . 7 5 1 . 3 9 17 37k 5 k . k 5 1 2 6 . 9 0 2 . 7 77 6 . 3 . 1 2 5 5 0 . 2 k l . k 2 18 . 356 5 k . 1 3 1 2 7 . 1 5 3 . 0 72 6 . 9 . l k o k 9.81 1 . 8 8 19 k 2 9 51 . 7 7 120.17 k . 6 51 7 .1 .159 kk.oo 2 . 0 2 20 k27 5 1 . 7 0 120.17 3 . 5 76 7 . 3 . l k 5 5 0 . 6 3 1 . 7 3 21 k l 8 51.17 1 2 0 . 2 5 3 . 8 73 7 . 2 •152 k 7.18 1 . 8 6 22 150 k 9 . 9 2 1 2 0 . 5 8 k.o 76 8 . 7 .168 5 2 . 0 3 2 . 6 6 23 l k 7 k9.17 1 2 0 . 5 8 k.o 81 7 . 8 .17k k k . 9 1 2 . k 3 2k 7 k l 5 0 . 8 7 1 1 9 . 8 3 k . 2 66 1 0 . 8 . 1 8 5 5 8 . 1 3 3 - 6 5 25 7ko 5 1 . 6 2 1 1 9 . 8 3 l . k 125 1 1 . 2 . 1 7 8 6 3 . 7 k 3 . 2 7 26 738 5 0 . 7 0 1 1 9 . 5 0 k . 2 67 1 0 . 8 •173 6 3 . 3 0 3 . 2 0 27 720 5 0 . k 8 119 . 7 5 5 . 1 k 8 7.k .159 k 6 . k 5 2 . 0 9 26 3 3 * k 9 . 0 0 1 1 6 . 7 5 3 - 8 87 6 . 8 . 157 k 2 . 9 9 1 .77 29 3*3 k 9 . S 2 116 . 2 7 k . 5 66 7 . 6 . 1 6 3 k 6 . 7 9 2 . 0 7 30 393 k 9 . 9 2 119 . 7 7 k . l 7k 9 . 9 . I 8 3 5 * . 5 k 3 . 0 1 31 3*6 k 9 . 9 2 U 8 . 5 8 k.o 76 8 . 6 .168 51.k 5 2.kk 32 387 5 » . 3 7 1 2 2 . 5 0 2 . 6 8 0 8 . 1 . 1 5 5 5 2 . 1 1 2.18 33 7k7 k 9 - 3 3 1 1 7 . 2 5 k . 2 76 8 . 2 .158 5 2 . 8 8 2.27 3 " 710 * 9 . 3 7 I l k . 5 8 k . 6 66 7 . 1 .157 k k . 7 k 2 . 0 0 35 6 9 9 k9.17 116.13 k . 7 65 8 . 6 • 173 ' 9 . 7 7 2 . 5 6 36 701 5 0 . 1 3 115. * 3 k . 6 61 9 . 0 . 1 7 6 5 1 . 5 5 2 . 8 0 37 3 k * k 9 . 5 8 1 1 5 . 9 8 3 . 8 83 8 . 8 .176 5 0 . 2 6 2 . 5 8 3« 3 k l * 9 . 3 0 1 1 6 . 0 0 5 . 5 k6 6 . 9 .156 k k . 9 9 1 . 8 k 39 3*5 k 9 . o 8 116 . 7 5 k . 6 6 8 7 . 8 .178 k 3.61 2 . k l ko 709 5 1 . 1 3 1 1 7 . 0 8 k . 3 . 62 8 . 0 .175 * 5 . * 5 2 . 5 k k l 337 • 9 . 5 8 1 1 7 . 8 0 5-7 ko 7 . 6 .170 k k . l l 2.15 k2 773 5 3 - 3 3 122.17 3-k 6 8 8 . 5 .168 5 0 . 3 9 2 . 5 9 »3 8 9 8 51 . 1 2 1 1 7 . 2 5 k . 7 53 7 . 9 . 1 7 0 k 6 . k 6 2 . 2 7 t> l> 732 k 9 . k 2 I I 8 . 0 3 k . 5 6 8 8 . 5 . I 6 3 5 1 . 2 9 2 . 7 8 " 5 338 k 9 - 7 5 1 1 7 . 0 0 k . 9 57 7 . 3 .172 k 2.ko 2.17 k6 335 k 9 - 3 3 I I 6 . I 3 5 . 3 51 6 . 9 .155 k k . 6 8 1 . 7 5 *7 779 5 k . 1 0 I 2 2 . 0 5 2 . 0 95 8 . 0 .156 5 1 . 7 k 2.13 * 8 385 5 k . 2 5 1 2 2 . 2 5 2 . 5 83 8.k . 1 5 9 5 2 . 9 3 2 . 3 8 *9 51 5 k . 3 3 1 2 3 . 0 8 2 . 5 82 8 . 8 .168 5 2 . 7 k 2 . 7 6 50 k 3 l 5 k . 1 2 1 2 2 . 8 3 2 . 2 9 0 8 . 1 . 1 5 5 5 2 . 7 2 2 . 2 k 51 k l 5 3 - * * 1 2 2 . 6 7 1 .7 106 8 . 3 .157 53.71 2.k6 52 725 5 0 . 6 5 117 . 5 3 k . 2 67 7 . 1 . I 6 3 k 3 . o 8 2 . 0 3 53 2k0 5 0 . 1 7 1 1 8 . 0 0 3 . 1 95 6 . 9 .156 k 3 . 5 0 2 . 1 5 5 * 715 * 9 . k 8 1 1 7 . 2 3 k.k 70 8 . 1 .169 > 7 . 8 5 2 . 3 8 55 37 5 k . 0 8 1 2 2 . 0 5 2 . 2 91 7 . 0 . 1 2 9 5 k . 36 1 . 7 9 56 39 5 k . 0 8 1 2 2 . 0 5 2 . 1 93 8 . 8 .162 5 k . 1 8 2 . 6 k 57 " 3 5 k . 2 8 1 2 2 . 6 2 2 . 3 87 8 . 6 . 1 6 2 5 3 . 2 9 2 . 6 2 58 k6 5 k . 0 8 1 2 2 . 0 8 2 . 3 88 8 . 0 .166 k 8 . 9 0 2 . 5 2 59 k7 5 k . 0 8 1 2 2 . 0 8 2 . 3 8 8 8 . 7 . 1 7 2 5 1 . k l 2.81 60 k 9 5 5 - 5 0 1 2 1 . 5 8 2 . 0 86 6 . 9 .136 5 0 . 5 9 1 .7k 61 66 5 1 . 6 7 120.17 k . 6 52 5 . 5 . 1 2 0 k k . 6 9 1 . 2 2 62 905 5 1 . 5 8 1 1 9 . 8 5 l . k 125 1 1 . 0 .165 6 6 . 7 8 3 . 0 1 «3 281 5 3 . 1 0 1 3 2 . 0 5 ,k I 3 8 1 2 . 2 .187 6 5 . 8 7 3 . 3 7 6k 28k 51.18 1 2 5 . 5 7 . 0 139 1 0 . 8 .178 6 0 . 9 5 2 . 8 8 « 5 296 5>.17 122.17 2 . 2 90 8 . 1 .151 5 3 . 7 9 2 . 3 8 6 6 336 5 0 . 2 5 U 5 . 7 5 5 . 0 52 7 . 7 .161 k7.91 2 . 0 7 67 3»o 5 0 . 5 3 1 1 5 . 9 2 k . 5 61 9 . 5 .177 5 k . 5 k 2 . 6 8 6 8 355 5 k . 2 3 1 2 7 . 3 7 2 . 0 9k 6 . 0 . 1 2 k k 7 . 6 6 1 . 2 k 6 9 359 5 * . 17 1 2 1 . 5 0 2 . 7 79 7 . 6 . l k 7 51.7k 2 . 0 0 70 36 k 5 * . 9 3 127.k 2 2 . 0 90 8 . 1 . l k l 5 8 . 0 1 1 . 8 k 71 365 5 k . 9 3 1 2 7 . k 2 2 . 0 90 7 . 9 . l k l 5 5 . 6 6 2 . 0 1 72 366 5 k . 9 3 1 2 7 . k 2 2 . 0 90 6 . 8 .129 5 2.51 1 . 5 0 73 3«9 5 k . 9 3 127.k 2 2 . 0 90 7 . 2 .129 5 6 . 2 9 1 . 7 1 7* 371 5k. 1 0 1 2 7 . 2 0 3 . 3 66 6 . 9 .136 5 0.91 1 . 7 7 75 373 5k.0 8 1 2 7 . 2 5 2 . 6 82 6.k . 1 3 0 k 9.81 I . 6 0 P r o v . 8 . 8 . S . L . R.C.D. S.L./R.C.D. Dry Wt. Ho. No. L a c . L O O R . E U v . Days ( c l C o l I B . I 76 379 5 5 . 6 0 1 2 7 . 8 3 1.5 97 8.8 •153 5 5 . 0 9 2 . k 2 77 382 5 5 . 6 0 1 2 7 . 8 3 1.5 97 8.7 . l k 9 5 9 . 0 8 2 . 3 5 78 389 5 k . 0 8 1 2 2 . 0 8 2 . 2 91 8.1 . 1 5 0 5*.23 2.19 79 390 5k. 08 1 2 2 . 0 8 2 . 2 91 8.1 .161 5 0 . 6 7 2.*3 80 392 5 0 . 0 7 1 1 9 . 6 8 k.7 6 0 7.6 .162 k 7 . 1 8 2 . 1 k 81 39k 5 0 . 0 2 1 1 9 . 7 0 5-5 k2 6.k .156 k o . 7 6 1 . 6 k 82 395 5 0 . 1 2 1 1 9 . 7 0 k.7 59 7.6 . I 6 3 * 7 . 0 9 2 . 0 5 83 k l 6 5 0 . 8 5 1 1 9 . 8 7 k.6 57 7.7 .165 k 6 . 3 5 2 . 2 0 8k k l 7 5 1 . 2 3 120.25 k.o 68 7 . 9 . 1 5 0 5 2 . 3 * 2 . 1 0 85 k26 5k.6 7 1 2 8 . 7 5 .7 121 1 0 . 0 .168 6 0 . 7 * 2 . 6 5 86 k 2 8 51.73 1 2 0 . 0 8 k . 2 6 0 6.8 .151 *5.16 1.81 87 ' 3 0 5 0 . 6 7 119 . 5 8 k . 2 67 7.1 .157 *5.*k I . 8 5 88 kk7 5 1 . 0 0 1 1 9 . 8 7 k . 7 5k 7 . 2 .159 kk , f l 2 2 . 0 1 89 k 5 k 5k.93 127.k2 2 . 0 9 0 8.6 .155 55 - 7 8 2 - k 5 90 k88 k 9 . 0 7 120.88 3-5 9 3 8.0 .167 k 7 . 8 2 2 . 2 k 91 k 9 9 5k.0 8 122-33 2 . 0 95 8 . 2 .162 5 0.61 2 - 5 3 92 525 5k.75 1 2 8 . 7 5 .8 118 8 . 3 . U 9 5 6 . 5 8 2 . 0 k 93 526 5 5 . 0 8 1 2 9 . 3 3 .7 118 9.6 ,16k 5 9 . 1 0 2 - k 5 9k 529 5 3 . 6 7 1 3 2 . 3 3 .1 l k l I 0 . 5 .172 6 0.*k 2 . 9 3 95 691 k 9 . l 2 120.88 k.o 82 8.k .190 k k . 5 1 2.88 96 692 52.37 1 2 6 . 7 5 .1 l k 9 l o . 1 . 1 5 9 6k. 1 3 2 . k 8 97 703 *9.»3 1 1 6 . 1 3 5.1 55 7.5 .177 "1.93 2.kk 98 718 5k.18 122.12 2 . 0 95 8.8 .161 5k.8 k 2 . 5 6 99 719 5k.18 122.12 2 . 0 95 8.2 .16k 5 0 . 6 8 2 . 5 8 100 723 5k.2 5 1 2 1 . 9 2 2.7 78 8.3 .162 5 1 . k l 2 . 6 3 101 7k2 k 9 . 1 3 1 2 5 .k2 . 2 168 13.9 . 2 0 2 7 0 . 5 8 3.77 102 7k5 5 2 . 2 0 1 1 9 . 2 5 3-5 73 8.0 .178 * * . * 3 2 . 5 9 103 7k8 5 0 . 0 5 1 2 7 . 0 5 .1 16k 11.9 .178 6 7.31 3 . 0 7 ?ok 766 k 8 . 8 3 l 2 k . 7 5 .3 I 6 7 12.1 .18k 66.k 5 3 . k l 105 767 5 3 . 0 8 1 3 2 . 0 8 .3 l k o 11.k . 1 7 8 6 3 . 8 8 3 . 0 9 106 769 5k.17 1 2 2 . 1 7 1.9 97 7.9 .1*9 5 2 . 8 0 2.12 107 770 5k. 25 1 2 2 . 3 3 2 . 2 90 7.8 . 1 5 2 5 1 . 6 9 2 . 2 1 1 0 8 771 5k.2 5 1 2 2 . 3 3 2 . 2 90 8.2 .157 5 2.7* 2 . 2 6 109 772 5k.17 1 2 2 . 2 5 2.2 9 0 7.k .1*7 51 . 0 7 2 . 0 6 110 777 5k.17 122.17 1.9 97 8 . 2 . 1 5 5 53-18 2 . 2 3 111 780 5k.0 8 121.88 2 . 0 95 7.0 . 1 3 k 5 2 . 2 1 1 . 7 2 112 78k 5*.88 126.73 2.7 7k 7.6 . l k 7 5 1 . 8 9 1 . 9 3 113 789 5k.35 1 2 5 .k3 2.9 73 7.6 . l k k 5 3 - 0 7 1 . 9 7 I l k 790 5k.0 7 127.22 3-o 73 7.6 .156 kS.15 2 . 2 k U 5 791 5 5 . 0 8 127.33 2.9 68 a.i .15k 5»-77 2 . 0 9 116 792 5 5 . 1 3 1 2 7 . k o 1 . 0 111 9.6 .153 6 2 . 9 0 2 . 7 5 117 793 5 5 . 3 0 1 2 7 . 7 0 1.1 108 8.5 . 1 2 7 67.33 1.81 118 79k 5k.6 3 1 2 7 . 2 0 2.5 80 8.k .160 5 2 . 9 0 2 - 5 3 119 795 5k.12 1 2 5 . k 2 2.7 79 8.7 . 1 5 2 5 7 . 2 6 2 . 3 2 120 796 5 5 . 0 0 127.00 3 . 2 62 7.6 .1*5 5 2 . 8 0 1 . 9 8 121 797 5k.8 0 1 2 9 . 0 3 1.9 93 8.8 .15» 5 7 . 1 0 2 . 2 6 122 798 5k.75 1 2 7 . 0 0 1.9 93 9.1 . 1 5 8 57.77 2 . 6 k 123 799 5 5 . 0 3 1 2 6 . 5 0 2.3 82 7.6 . l k k 5 3 - 2 0 1 . 8 5 12k 8 0 0 5 5 . 2 8 127.ko 1 . 2 106 9.3 .153 61.15 2 . 3 6 125 8 0 1 5 0.53 1 2 7 . 2 5 .6 150 12.5 .191 6 5 - 8 7 3.61 126 805 55.k7 1 2 7 . 7 0 1.2 105 10.1 . 1 5 1 6 7 . I 2 2 . 6 0 127 806 5k.9 0 127 . 2 5 1.5 101 7.9 . 1 3 8 5 7 . 6 k 1 . 9 2 128 807 5 5 . 3 0 1 2 7 . 1 5 2 . 2 83 9 . 0 . 1 5 1 5 9 . 8 9 2 . 2 5 129 8 0 8 5 k . 6 7 1 2 7 . 2 5 2.k 82 8.9 .155 5 7 . k 5 2.55 130 8 0 9 5 3 . 5 0 1 3 2 . 2 5 .k 135 1 0 . 0 .165 6 0 . 2 k 2 . 6 2 131 8ko 5 5 . 0 8 1 2 9 .k2 .1 132 10.0 . 1 6 3 6 1 . 6 2 2 . 5 0 132 k6k 5k.0 5 1 2 S . 6 5 .1 139 1 1 . 3 .167 6 7 . 9 0 2 . 7 9 133 5k5 k 9 . 0 5 I l k . 6 7 5.0 59 k.6 . 1 2 5 3 6 . 2 1 1.11 1 3 » 576 5 0.25 1 1 7 . 8 3 k . 2 70 8.6 .18k k 6 . 9 3 2.8* 135 8 5 8 k 9 . 0 0 1 1 6 . 6 7 k.2 78 9.1 .17k 5 1 . 9 8 2.5* 136 868 k 9 . 0 5 116.97 k.3 75 8.k . 1 7 k k 7 . 9 k 2 . 5 2 137 888 5 0 . 0 8 1 1 7 . 7 5 5 . 0 53 7.3 .168 * 3 - 3 l 2 . 1 k 136 539 k 9 . 9 2 U k . 7 7 k.5 65 8.k .160 52.17 2 . 2 2 139 5ko k9.92 I l k . 9 2 k.9 56 6.k .1*7 k 2 . 8 5 I . 6 3 l k o 5 k l k9.92 U k . 7 3 5.1 51 6.9 .157 * 3 . 7 0 .1 . 9 6 l k l 5k2 k 9 . 9 2 I l k . 7 5 k.6 6 3 7.7 . 1 3 9 5 5 - 2 9 1-75 l k 2 575 5 0 . 2 5 1 1 8 . 0 0 3-8 79 S.k .185 * 5 . 6 3 2 . 7 8 1*3 6 2 3 57 . 1 7 I 2 I . 8 3 2.7 60 6.5 .133 k 8 . 9 3 1.55 l k k 636 •9.77 1 1 5 .ko 5 . 0 55 7.5 .167 k 5 . 1 k 2 . 3 0 1*5 6 k l 50.57 119.18 k.9 52 7.6 .168 k5 - 3 2 2.18 l k 6 6kk 5 0 . 6 3 118.00 k.o 72 7.7 .172 k k . 7 6 2.kk l k 7 6 5 k *9.1o I l k . 6 7 5.6 k5 7 . 0 . 1 7 0 k l . 5 2 2 . 0 k I k S 6 5 5 » 9 . 0 8 1 1 5 . 7 8 k.5 70 8.6 .173 k9.61 2 . 5 6 l k 9 6 5 7 k 9 . 2 5 U 6 . 2 3 k.8 6 3 7.0 .159 kk.*6 1 . 9 7 I 5 0 857 k 9 . 2 5 117 . 7 5 5 . 5 k 7 8.1 .179 k5.Sk 2 . 5 3 Lat. - Latitude; Long. - Longitude; Elev. - Elevation; Days - days in growing season;'- S.L. - Shoot length; R.CD.^- Root collar diameter; Dry Wt. - Dry weight. 6U TABLE 10 GROWTH BEHAVIOUR DURING THE SECOND YEAR IN THE NURSERY S.L . R.C.D. Mo. No. L a t . Lona. E l e v . Dava leml ( c « l S . L . /R .C .D . ( n i l No. 1 50 56 50 l ? l . l o 2 . 3 73 15 . 0 . 2 3 0 6*.7? 9,07 76 ? k8 5 5 . 6 7 1 2 2 . 2 0 2 . 0 85 15 .5 . 2 k 9 6?.18 10 .77 77 3 k? 5* 50 l 2 k . ? 5 ?.? 8 8 l k . k . 2 3 4 61 . 7 9 10 .81 78 ii 1.93 5 * 50 1 ? ? . 6 7 2 . ? 88 1 5 . 0 . 2 b 9 5 9 . 99 I 0 . 0 5 79 . 5 lioli 5* 33 1 2 2 . 6 7 ?.3 87 15.» . 2 5 5 6 0 . 7 9 10 .29 80 6 7*6 5* 10 1 2 2 . 0 5 2 . 0 95 16.9 . 2 6 7 6 3 . 5 8 1 2 . 2 b 81 7 722 5 * . ° 7 1 2 1 . « 5 2 .3 77 17.? .271. 6 2 . 9 4 11 .8b 82 8 353 5» 93 127.w? 2 . 0 90 16 . 0 .?k9 6b . 5 2 1 0 . 0 7 83 9 388 53-57 1 2 2 .4? 3 . 0 75 15.8 . 2 6 5 6 0 .16 I 0 . I 3 8b 10 522 52 33 1 2 1 . 6 7 2.2 102 16.1 . ? 3 8 68.22 9 . 6 0 85 11 5 ? 3 52 «2 1 2 1 . 1 2 2.2 101 17 . 0 • 257 66.17 10 .57 86 12 380 55 60 1 2 7 . 6 3 2.7 70 15 .6 .?77 5 6 . 7 3 13 .19 87 13 358 55 •7 127 .83 1.7 93 16.7 .?61 6b.b 9 11.53 88 Ik 370 55 33 1 2 6 . 6 3 2 - 3 80 14.4 . ? 3 8 60.22 8 . 7 0 89 15 737 51 12 118 .25 « . 3 6? 15.8 . 2 7 0 5 8 . 8 9 11.53 90 16 367 5* 93 1 2 7 . « 2 2 . 0 90 1 3 . 5 . 2 3 1 5 8 . 5 9 7 . 3 6 91 17 37* 5» *5 126 . 9 0 ?.7 77 U . l . 2 3 5 59 . 9 8 8 . b 5 92 18 356 5 ' 13 127 .15 3 . 0 72 15.8 . 2 5 9 6 0 .97 11.15 93 19 1.29 51 77 120 .17 ».6 51 13.2 , 2 b o 5 b . 0 ? 8 . 0 9 9 b 20 k27 51 70 120 .17 3 . 5 76 I 6 . 5 . 2 5 8 6 3 .35 10 .35 95 21 kl8 51 17 1 ? 0 . ? 5 3 .8 73 1*.7 . 2 5 3 5 8 . 0 7 9 . 0 1 96 22 150 bo 92 I 2 0 . 5 8 k.o 76 17.1 . 2 7 8 61.71 1 2 . 7 2 97 23 1»T »9 17 1 2 0 . 5 8 4 . 0 81 17 . 0 . 2 9 8 5 7 . 0 8 13 ,95 98 2k 7*1 50 87 I I 9 . 8 3 4.2 66 2 0 . 0 . 2 8 7 7 0 . 1 0 1 5 . 2 0 99 ?5 7I10 51 62 I I 9 . 8 3 1.4 125 2 0 . 8 . 2 8 8 7 2 . 3 6 15.71 100 ?6 738 5 0 . 7 0 119 . 5 0 4.2 67 2 1 . 3 . 2 9 7 7 2 . 0 8 16.73 101 27 720 50.4H 119 .75 5-1 k8 16.1 . 2 7 0 59 . 0 0 11 . 1 2 10? 28 3 3 * »9 00 116 .75 3-8 87 1 3 . 8 . 2 b 9 5 5 - » 3 8 . 2 1 103 29 3«3 *9 82 116 .27 • • 5 66 1 5 . 0 . 2 5 8 59 . 5 6 9 . 3 0 lob 30 393 •9 92 119.77 4.1 7k 1 7 . 8 . 2 8 7 6 2 . 2 6 13.91 105 31 3*5 *9 92 U 8 . 5 8 it. 0 76 IS .3 . 2 6 3 6 I . 5 6 1 0 . 1 3 106 32 387 5> 37 1 2 2 . 5 0 2.6 80 15.8 . 2 5 0 6 3 . bb 9 . 5 5 107 33 ;»7 «9 33 117 .25 ».2 76 16.4 . 2 5 6 6».17 ' I 0 . 9 6 108 3 ' 710 *9 37 11» .5 8 4.5 66 I 3 . 8 . 2 5 1 5 * . 3 3 9 . 2 1 109 35 6 9 9 •9 17 U 6 . I 3 65 16.1 . 2 8 1 57.77 1 2 . 3 6 110 3* 701 50 13 115.<3 k . 6 61 17 .6 . 2 8 1 63-59 13 . 3 3 111 37 31, k *9 5S 115.98 3 . 8 83 17.9 . 2 8 0 - 6b. 1 5 12.71 11? 38 3*1 *9 30 116 . 0 0 5 . 5 I16 U . 5 . 2 b b 59 . 3 6 8 . 0 2 113 39 3*5 . 9 08 116.75 1.6 68 15.8 . 2 8 b 55-89 12 .19 l i b bo 709 51 13 U 7 . 0 8 * . 3 6? 16.1 . 2 7 5 5 8 .53 11.31 115 b l 337 *9 58 117 . 3 0 5 .7 ko 15.2 . 2 6 6 5 7 . « 9 . 7 8 116 b? 773 53 33 122 .17 3-» 68 15.8 . 2 6 3 60.47 1 0 . 6 0 117 " 3 8 9 8 51 12 117 .25 k.7 53 16 . 9 . 2 8 5 59 . 5 5 1 2 . 9 0 118 bk 732 •9 »2 I I 8 . 0 3 k . 5 68 I 6 . 5 . 2 7 5 59.8b 1 2 . ; 6 119 «5 338 "9 75 117 . 0 0 k.9 57 l k . 9 . 2 7 ? 5 4 . 6 0 1 0 . 3 8 120 • 6 335 »9 33 116.13 5 - 3 51 l b .2 . 2 5 2 5 6 . 2 3 7 . 8 3 121 k7 779 5* 10 1 2 2 . 0 5 2 . 0 95 15.8 . 2 5 6 6 3 . 6 9 10.81 122 • 8 385 5 ' 25 1 2 2 . 2 5 2 . 5 83 18 . 0 . 2 7 6 6 5 . 6 3 1 3 . 1 0 1?3 »9 51 5* 33 1 2 3 . 0 8 2 . 5 82 18.1 . 2 8 3 6 3 . 8 9 13 . 7 2 12b 50 *3 l 5* 12 1 2 2 . 8 3 ?.? 90 16.7 . 2 6 2 6 3 . 9 7 11 . 0 1 1?5 51 k l 53 »2 122 .67 1.7 106 16.8 . 2 6 0 6 b . 3 0 11 .07 126 52 725 50 65 " 7 . 5 3 4.? 67 l b . 5 . 2 5 1 57.1? 9 . 2 1 127 53 ?bo 50 17 118 . 0 0 3-1 95 l b .2 . 2 6 b 5 3 . k i 1 2 . 1 2 128 5» 715 . 9 • 8 117 . 2 3 b.l. 70 16 . 0 . 2 6 3 57 . 0 9 11.77 129 55 37 5 * 08 I 2 2 . 0 5 ?.? 91 l b . 8 . 2 b 6 59.9k U . 2 8 130 5« 39 5 * 0 8 1 2 2 . 0 5 2.1 93 15.9 . 2 6 0 61 . 3 8 11.19 13» 57 »3 5k 28 1 2 2 . 6 2 2 . 3 87 17.1 . 2 6 b 6 5 . bb I I . 6 5 13? 58 *6 5 ' 08 1 2 2 . 0 8 2 . 3 88 16.6 . 2 7 8 59.91 I 3 . 0 8 133 59 *7 5» 08 1 2 2 . 0 8 2 . 3 88 16.8 . 2 7 3 6 2 . 0 9 I 2 . 3 9 13b 6 0 »9 55 50 1 2 1 . 5 8 ?.o 86 l b . 8 . 2 b l 6 0 . 2 5 9.8b 135 61 66 51 67 120 .17 4.6 52 12.2 .22? 53 . 2 5 7 . 3 0 136 62 905 51 58 119 .85 1.* 125 2 0 . 6 . 2 8 b 7 3 . 0 6 15.37 137 «3 281 53 10 1 3 2 . 0 5 .ll 138 ?b.? • 3 5 9 68.35 2 3 . 0 8 138 6* 28b 51 18 1 2 5 . 5 7 .9 139 ?0.2 • 3?3 6 3 , 0 2 18.17 139 55 296 5 ' 17 1 2 2 . 1 7 2.2 90 16 . 0 . 2 6 ? 61 . 0 b 11.66 Ibo 66 335 5 ° 75 115.75 5 . 0 52 l b . o ,?bo 5 8 . 0 b 8.0 8 l b l 67 3*10 50 55 I I 5 . 9 ? k . 5 61 18.1 . 2 7 1 <7 . 3 9 1 2 . 4 3 ' l b ? 68 355 5* 23 127.37 2 . 0 9> 13-7 . ? 3 8 5 8 . b? 7.8b 1*3 69 359 5» 17 1?1 .50 2.7 79 l b .b . 2 b 6 5 8 . 8 6 9 . 4 3 lbb 70 36b 5» 93 l ? 7 . k 2 2 . 0 90 16 . 3 . ? b 6 6 6 . 5 7 9 . 8 2 l b 5 71 365 5 " 93 127.42 2 . 0 90 15 .5 . 2 b b 6 3 .76 11 . 6 7 1*6 72 366 5» 93 127.42 2 . 0 90 l b . o . ?3» • 59.35 8 . 5 0 147 73 369 5k 93 127.42 2 . 0 90 15.9 . 2 6 1 61 . 0 ? 9. ?b l b s 7* 371 5k 10 1 2 7 . 2 0 3 -3 66 1>.0 • 2?3 6?.61 6 . 8 5 l b 9 75 373 511 0 8 127 . 2 5 2.6 82 1 3 .9 • 2 3 3 59.51 8 , 5 0 150 Long. E l e v . Dava 379 38? 389 390 39? 39b 395 b l 6 b l 7 »26 b28 b30 bb7 454 b88 «99 525 526 529 691 6 9 ? 703 718 719 723 7»? 7»5 7»8 766 767 769 770 771 77? 777 780 78b 789 790 791 792 793 79b 795 796 797 796 799 8 0 0 8 0 1 805 606 807 8 0 8 809 8 b o b6b 5*5 575 8 5 8 8 6 8 8 8 8 539 5b o 5 b l 5 . 2 575 623 636 Sbl 6bb 65b 6 5 5 657 857 5 5 . 6 0 5 5 . 6 0 5 b . 0 8 5 b . 0 8 5 0 . 0 7 5 0 . 0 2 5 0 . 1 2 5 0 . 8 5 51 . 2 3 5 4 . 6 7 5 1 . 7 3 5 0 . 6 7 5 1 . 0 0 5 " . 93 4 9 . 0 7 5 b . 0 8 5 » .75 5 5 . 0 8 5 3 - 6 7 b 9 . l 2 5 2 - 37 b 9 . b 3 5 b .18 5 b .18 54. ?5 l>9.13 5?.?o 5 0 . 0 5 4 8 . 8 3 5 3 - 06 54.17 5 4 . ? 5 54. ?5 5 * . 1 7 5 * .17 5 b . 0 8 5 b . 8 8 5 » . 3 5 5 k . 0 7 5 5 . 0 8 5 5 . 1 3 5 5 - 3 0 5 > . 6 3 5 4 . 1 2 5 5 . 0 0 5 » . 8 o 5 ' . 7 5 5 5 . 0 3 5 5 . ?8 5 0 . 5 3 55 .47 5 b . 9 0 5 5 - 3 0 5 " . 6 7 5 3 - 5 0 5 5 . 0 8 5 4 . 0 5 4 9 . 0 5 5 0 . 2 5 b 9 . 0 0 4 9 . 0 5 5 0 . 0 8 4 9 . 9 2 b 9 . 9 2 » 9 . 9 2 4 9 . 9 2 5 0 . 2 5 5 7 .17 " 9 . 7 7 5 0 . 5 7 5 0 . 6 3 b 9 . 1 0 4 9 . 0 8 4 9 . 2 5 4 9 . 2 5 2 7 . 8 3 2 7 . 8 3 2 2 . 0 6 2 2 . 0 8 19.58 19 . 7 0 19 . 7 0 19 .87 20 . ? 5 ? 8 .75 2 0 . 0 8 19 . 5 8 19 . 8 7 27.k? ? 0 . 8 8 ??.33 ? 8 .75 2 9 . 3 3 3 2 . 3 3 20.83 2 6 .75 16.13 22 .12 22.12 21 . 9 2 2 5 . b? 19 . 2 5 2 7 . 0 5 24.75 3 2 . 0 8 2 2 .17 2 2 . 3 3 2 2 . 3 3 2 2 . ? 5 2 2 .17 21.98 2 6 .73 25-43 2 7 . 2 2 2 7 - 3 3 2 7 . bo 27 . 7 0 2 7 .20 2 5 . » 2 2 7 . 0 0 2 9 . 0 3 2 7 , 0 0 2 6 . 5 0 27.«0 27 . 2 5 2 7 . 7 0 2 7 . 2 5 2 7 . 1 5 2 7 . 2 5 3 2 . 2 5 29.42 2 8 . 6 5 l b . 6 7 1 7 . 8 3 1 6 . 6 7 16.97 17.75 1 » . 7 7 l b .92 1*.73 1».75 18 . 0 0 2 1 . 8 3 1 5 .bo 1 9 . 4 8 1 8 . 0 0 l b . 6 7 15 . 7 8 1 6 . 2 3 17.75 k.7 2 . 0 3-5 2 . 0 5-1 2 . 0 ? . 0 ?.7 1.9 ?.2 2.2 2.9 3.0 5-1 b.6 3.8 2.7 5.6 » .5 b.B 5.5 118 118 l b l 82 lb9 55 78 168 73 16b 167 lbo 95 74 73 73 6 8 111 103 80 79 6 ? • 93 93 8? 106 I 5 0 105 101 83 82 135 13? 139 59 70 78 75 53 65 56 51 63 S .L . I f ) 16.5 17-3 15.9 17.0 1 5 . 5 1 3 . * 15.? 15.8 15 . 1 19 . 1 l ? . b 15 . 3 15 .1 17.0 1 5 . 5 I 6 . 9 17 .* 17.3 21.8 1 5 . 2 19.b l b . 5 16.2 16.8 17 .5 ??.7 15. ? 21.» 21.7 20.8 15.4 15.8 16. ? 1 " ,7 15-9 1 5 - 3 1 5 . 0 l b . 6 1 5 . 5 17 .5 19.b 15.7 16.2 17.8 17 . 3 17.9 17. k 17.6 19.0 2 k . 1 18.9 15.1 17 .5 18.0 20.2~ 19.k 22 . 1 10.? 16.2 I 8 . 3 16.2 1 3 . 6 I 6 . 5 12.7 12 . 5 16.0 16.0 l b . 5 l b . b l b . 6 15.6 U . 2 17 .2 l k . 9 lb .b a . C D . Icml S . L . /R .C .D . • 293 . 2 6 3 . 2 b 9 . 2 8 3 . 2 6 5 . 2 3 8 . 2 6 3 . 2 6 5 . 2 b 6 . 3 1 1 .21b . 2 6 b . 2 5 6 .277 .271" . 2 6 9 .293 .27 1 * . 3 6 2 . 2 7 8 . 3 0 9 . 2 6 2 - 2 5 9 . 2 6 9 . 2 8 5 .3>6 .281 .319 . 3 3 6 -331 . 2 5 9 .261 . 2 5 7 .248 . 2 5 0 . 2 5 0 . 2 3 6 . 2 3 ! . . 2 5 7 . 2 7 6 . 2 8 b . 2 2 9 . 2 6 3 . 2 5 7 . 2 6 7 . 2 7 1 . 2 7 3 . 2 6 7 , ? 6 b . 3 6 7 . 2 7 2 . 2 3 9 . 2 6 2 .277 . 3 b o . 3 0 9 . 3 " ? .209 . 2 7 2 . 2 8 9 . 2 6 9 . 2 6 2 . 2 6 0 . 2 3 0 • 2 3 9 , ? 5 1 .271 . 2 2 6 . 2 6 8 .264 . 2 7 7 .261 . 2 7 8 .271 . 2 6 2 6 5 . 6 0 5 5 . 6 5 63 .91 6 0 . 8 0 58 .51 5 6 . 0 3 5 7 . 6 9 5 9 . 7 9 6 0 . 8 5 6 2 . 9 5 5 8 . 3 9 5 7 . 9 6 53 . 5 9 6 3 . 0 I 5 6 . 5 6 6 3 . 0 0 5 9 . k 6 6 3 . 6 I 6 0 . 10 54 . 2 b 6 2 . 9 k 5 5 - 3 3 6 3 . 6 0 6 2 . 0 7 6 I . 8 9 6 6 . 1 0 54.19 6 7 . 2 1 6 k . 3 9 6 ? . 7 ? 6 0 . 1 5 6 I . 2 5 6 3 . 2 2 5 9 . k 2 6 3 . 7 3 61 . 9 2 6 3 . bb 6 ? . 9 b 6 0 . 5 I 6 3 . k 9 6 8 . 6 0 6 8 . 6 k 61.5 6 6 8 . 9 7 6 6 . 0 2 6 6 . k 9 6 3 . 7 0 6 6 . 3 2 6 7 . 8 0 6 6 . I 3 6 9 . 7 8 6 k . k 2 6 6 . 7 5 6 k . 1 2 6 0 . 1 0 6 2 . 6 2 6 5 . 2 7 0 8 . 1 3 5 9 . k 6 6 3 . 8 6 6 0 . 1 1 5 2 - 96 6 3 . 19 5 * . 3 5 51 . 0 3 6 3 . 6 5 6 0 . 2 5 6 k . ? 8 5 3 - 28 5 k . 9 8 5 6 ,70 5 k . 3 9 6 2 . 0 9 5 5 . 2 1 5 k . 7 8 Dry u t . ' « • ) 1 3 . 2 3 I I . 6 3 9 . k 2 1 2 . 5 8 10.19 7-35 9 .51 9 . 8 6 9 .19 16 . 3 2 6 . 7 9 1 0 . 0 6 8 . 9 9 1 1 . 1 2 1 1 . 7 0 1 1 . b9 l k . 1 2 1 2 . 2 3 2 5 .19 11.92 17 . 1 2 10 .19 1 1 . 3 b 1 2 . 0 8 l b .15 2 3 . 0 5 1 2 . 5 2 19.16 19 -02 19 . 2 3 I 0 . 5 5 1 0 . 7 9 9 . 6 2 1 0 , 0 k 9 . k 6 9 . 5 k 8 . 2 k 7 . 8 9 10 .07 I 0 . 7 0 1 3 . ' 5 8 . 2 1 12 .19 1 1 . 2 8 1 2 . 2 1 1 2 . 1 2 U . 3 o 1 1 . 1 5 11 . 0 0 2 5 . k 9 1 1 .ko 9 . 3 k 1 1 . 1 0 1 2 . 8 7 2 1 . 0 0 15.91 19.48 5 .55 1 1 .ko 1 3 . 9 3 1 0 . 7 9 9 .27 9 . 6 2 7 . 7 6 6 . k 6 9 . 3 0 1 1 . 3 k 7 . 7 2 I 0 . I 3 9 . 7 3 11.1.6 9.59 1 2 . 3 8 9 . 9 6 1 0 . 1 1 Lat. - Latitude; Long. - Longitude; Elev. - Elevation; Days - days in growing season; S.L. - Shoot length; R.C.D. - Root collar diameter; Dry Wt. - Dry weight. TABLE 11 PERCENT FLUSHED ON k DIFFERENT DATES DURING THE SECOND YEAR IN THE NURSERY Prov . No. Reg. No. L a t . LonK. E l e v . Dave Apr .6 Apr . Ik A p r . 2 0 Apr.27 Prov. No. Reg. No. L a t . Long. E l e v . Days Apr.6 A p r . l b Apr .20 Apr . 2 7 1 50 56 50 121 10 2.3 73 10.0 6 3 . 3 95 . 0 l o o . o 76 379 5 5 . 6 0 127.33 1.5 97 0 . 0 0 33-3 35.0 93.3 2 b3 55 67 122 20 2 . 0 85 3-3 55-0 91.7 l oo .o 77 382 55 - 6 0 127.33 1.5 97 1.7 ' 5 . 0 .31.7 96.7 3 b2 5 k 50 12b 25 2.2 88 11.7 71.7 93-3 100.0 78 339 5 b . 0 8 122 . 0 8 2.2 91 1-3 b6 . 7 9 0 . 0 100 . 0 k h03 5» 50 122 67 2.2 88 3-3 k8.3 81.7 9 5 . 0 79 390 5 b . 0 8 122 . 0 8 2.2 91 6.7 35-0 8 5 , 0 100 .0 5 bob 5» 33 122 67 2.3 87 3-3 3 8 . 3 81.7 9 6 . 7 So 392 5 0 . 0 7 119.68 k.7 60 0 . 0 I 6 . 7 6 5 . 0 9 3 . 3 6 7b6 5b 10 122 05 2.0 95 1.7 3 6 .7 7 6 .7 95 . 0 81 39k 50^02 119 . 7 0 5-5 k2 0.0 10.0 6 0 . 0 9 3 . 3 7 722 5b 07 121 b5 2.8 77 0 . 0 31 .7 8 0 , 0 l o o . o 82 395 5 0 . 1 2 119 . 7 0 k.7 59 1.7 5 . 3 66.6 9 5 . 0 S 353 5b 93 127 b2 2.0 90 3-3 k l . 7 6 5 . 0 l oo .o 83 kl6 5 0 . 8 5 119.87 b.6 57 1.7 1 5 . 0 7 0 . 0 9 5 . 0 9 388 53 67 122 b2 3-0 75 0 . 0 I 8 . 3 6 3 . 3 95.0 8k kl7 51 . 2 3 120 . 2 5 b.o 63 0.0 16.7 5 6 . 7 91.7 10 522 52 33 121 67 2.2 102 3-3 51.7 9 6 . 7 9 8 . 3 85 k26 5k.6? 1 2 8 .75 .7 121 1.7 31-7 8 0 . 0 9 8 . 3 11 5 2 j 52 k2 121 b2 2 . 2 101 10.0 55.0 9 6 . 7 l oo .o 86 k 2 8 51 .73 120 . 0 3 k.2 60 0.0 10.0 6 6 . 7 9 6 . 7 1? 380 55 6 0 127 83 2.7 70 0 . 0 23-3 75-0 66.7 87 k 3 o 5 0 . 6 7 1 1 9 . 5 8 k.2 67 0.0 21 . 7 6 3 . 3 91.7 13 358 55 b? 127 83 1.7 93 0 . 0 35 . 0 8 I . 7 9 8 . 3 88 kk? 51.00 119.87 k.7 5k 0.0 20.0 36 . 7 100.0 lb 370 55 33 126 63 2-3 8 0 3.3 k8.3 9 6 . 7 9 8 . 3 89 k5k 5'-93 127.k2 2.0 90 1.7 3 6 .7 8 6 . 7 100.0 15 737 51 12 118 25 "•3 62 5.0 2 6 . 7 7 6 .7 9 6 . 7 90 k38 ko . 0 7 120.88 3-5 93 3-3 2 5 . 0 71.7 9 5 . 0 16 367 5* 93 127 »2 2 . 0 90 1.7 5 8 . 3 9 6 . 7 98.3 91 k99 5 b . 0 8 122-33 2 .0 95 0 . 0 35-0 3o.o 9 5 . 0 i ; 37b 5» b5 126 90 ,2.7 77 10.0 55-0 9 0 . 0 9 8 . 3 92 525 5k.75 1 2 8 .75 .8 U S 0 . 0 2 8 . 3 73.3 9 5 . 0 16 356 5» 13 127 15 3-0 72 0.0 33.3 8 6 . 7 100.0 93 526 5 5 . 0 8 129 .33 .7 118 0 . 0 bo.O 86.6 83-3 100 . 0 19 b29 51 77 120 17 b.6 51 3 . 3 13.3 7 6 .7 95.0 9k 529 53-67 132-33 .1 l k l 3-3 13.3 100.0 20 b?7 51 70 120 17 3 . 5 76 3 . 3 k5 . 0 93-3 95.0 95 691 k 9 .12 120.38 k.o 82 0.0 3.3 66 . 7 9 5 . 0 ? l • 18 51 17 120 25 3-8 73 1.7 33.3 6 8 . 3 95.0 96 692 52-37 1 2 6 .75 .1 lk9 0 . 0 2 6 . 7 7 5 . 0 100 . 0 22 150 bo 92 120 58 b.o 76 1.7 3 8 . 3 8 8 . 3 9 6 . 7 97 703 b o , b 3 U 6 . I 3 5 . 1 55 1.7 bo.O 73-3 0 5 . 0 23 l b , ba 17 120 53 b.o 31 0.0 20.0 73-3 9 6 .7 9 6 713 5b.18 122.12 2.0 95 3-3 53-3 9 0 . 0 9 3 . 3 2k 7bl 50 87 U 9 83 b.2 66 6.7 5 0 . 0 95 . 0 100 .0 99 719 5 k . 18 122.12 2 . 0 95 6 . 7 3 8 . 3 36 .7 9 3 . 3 ?5 7bo 51 62 119 83 I.b 125 3-3 k5.0 9 0 . 0 9 8 . 3 100 723 5 k . 2 5 121 . 9 2 2.7 78 3-3 k l . 7 8 6 . 7 9 6 . 7 26 738 50 70 119 50 b.2 67 13 .3 61.7 95.0 l o o . o 101 7*2 "•9.13 1 2 5 .k2 .2 168 1.7 5 0 .O 91.7 100.0 27 720 50 b8 119 75 5-1 bS 0 . 0 1 3 .3 6 3 . 3 95.0 102 7k 5 5 2 . 2 0 1 1 9 . 2 5 3-5 73 3-3 2 3 .3 70 . 0 95 - 0 28 33* bo 00 116 75 3.8 87 1.7 35 . 0 8 0 . 0 95.0 103 7k8 5 0 . 0 5 1 2 7 - 0 5 .1 16b 5 -0 k l . 7 91 .7 100 .0 29 3*3 b9 82 116 27 k.5 66 0.0 23-3 61.7 95.0 10k 766 b8 . 8 3 l2k .75 -3 167 0 . 0 23-3 81 .7 100 . 0 30 393 b9 92 119 77 b . l 7k 0 . 0 3 0 . 0 73-3 91.7 105 767 5 3 . 0 8 1 3 2 . 0 8 -3 lko 1.7 3 6 .7 36.7 100 . 0 31 3 " ko 02 118 58 b.o 76 0 . 0 I 8 . 3 81.6 9 6 . 7 . 106 769 5b.17 122.17 1.9 97 3.3 k l . 7 e 5 . o 95 - 0 32 387 5* 37 122 50 2.6 80 1.7 33.3 9 0 . 0 9 8 . 3 107 770 5k .2 5 1 2 2 . 3 3 2.2 90 1.7 5 8 .3 83.3 9 8 . 3 33 7*7 >9 33 117 ?5 b . 2 76 5 . 0 35.0 7 8 . 3 9 8 . 3 108 771 5b.2 5 122.33 2 .2 90 3.3 6 0 . 0 8 5 . 0 9 6 . 7 3k 710 bo 37 l i b 53 k.6 66 3.3 3 0 . 0 7 6 . 7 95.7 109 772 5 ' .17 122 . 2 5 2.2 90 S-7 5 0 . 0 38.3 9 6 . 7 35 6 9 9 b9 17 116 13 ».7 65 0 . 0 3 0 . 0 75-0 95.0 110 777 5k. 17 122 .17 1.9 97 3-3 kO.O ° 3-3 9 6 . 1 36 701 50 13 115 M 98 b.6 61 5.0 b5.o 8 5 . 0 100.0 111 780 5 b . 0 8 121.85 2.0 95 I 6 . 7 5 0 . 0 95.0 96.7 37 3bb »9 58 115 3.8 83 6.7 b 8 . 3 8 8 . 3 9 6 .6 112 78k 5 b . 83 126.73 2.7 7k 1.7 bo.o 8 6 . 7 100.0 38 3 b l 19 30 116 00 5-5 b6 0 . 0 1 3 .3 73.3 9 6 . 7 113 730 5 k-35 l 2 5 - * 3 2 . 0 73 1.7 55-7 8 S . 7 100.0 39 3*5 ko 03 116 75 b.6 63 5-0 21.7 6 8 . 3 91.7 l i b 790 5 k . 0 7 127-22 3- 1 73 0 . 0 2 5 . 0 ( 8 . 3 96.7 bo 709 51 13 117 0 3 >.3 62 3-3 31 .7 73.3 9 5 . 0 " 5 791 5 5 . 0 8 1 2 7 . 3 3 2.9 68 0 . 0 2 6 . 3 73-3 9 5 . ~> kl 337 bo 58 117 30 5.7 bo 3-3 I 6 . 7 76.7 9 6 . 7 116 792 55 -13 1 2 7 .bo 1.0 111 0.0 k6.7 9 3 - 3 9 8 . 3 k2 773 53 33 122 17 3." 68 1.6 11.? 71.7 93-3 117 793 55 - 3 0 1 2 7 - 7 0 1.1 108 0 . 0 18.3 61. 1 9 0 . 0 »3 898 51 12 117 25 k.7 53 6.7 k l . 7 75:o 9 6 . 7 118 79b 5k . 6 3 127 .20 2-5 80 5.0 66. 1 33.3 9 5 . 0 kb 732 »9 b2 118 0 3 k.5 68 1.7 21.7 8 3 . 3 l oo .o 119 795 5k.12 125.k2 2.7 79 5 -0 6 5 . 0 96.7 9 8 . 3 »5 338 >9 75 117 00 b.9 57 6.7 k l . 7 8 5 . 0 93-3 120 796 55-00 1 2 7 .00 3-2 62 5.0 5 1 .7 33.3 9 6 . 7 b6 335 bo 33 116 13 5.3 51 0 . 0 I 6 . 7 7 8 . 3 9 8 . 3 121 797 5k . 8 0 129 .03 1.9 93 16.7 55 - 0 9 6 .7 100.0 bT 779 *k 10 122 05 2 . 0 95 1 3 . 3 5 6 . 7 38.3 95.0 122 798 5'-75 127 .00 1.9 93 0.0 I 8 . 3 78.3 100.0 b8 385 5* 25 122 25 2.5 83 1.7 3 8 . 3 78.3 9 8 . 3 123 799 55 . 0 3 1 2 6 . 5 0 ?-3 82 1.6 k3.3 8 5 . 0 9 8 . 3 b. 51 5k 33 123 08 2 . 5 82 0 . 0 k6.7 8 8 . 3 9 6 . 7 12b 8 0 0 55 . 2 8 127.kO 1.2 106 0.0 2 8 . 3 75-0 9 6 . 7 50 »3 l 5' 12 122 83 2.2 90 3 . 3 5 6 .7 91.7 9 8 . 3 125 8 0 1 5 0 .53 1 2 7 . 2 5 .6 150 5.0 k l . 7 95-0 9 8 . 3 51 b l 53 b2 122 67 1.7 106 10 . 0 6 5 . 0 9 0 . 0 9 8 . 3 126 805 55. b? 1 2 7 . 7 0 1.2 105 5-0 3 6 .7 81 . 7 0 6 . 7 52 725 50 65 117 53 ' b.2 67 5 . 0 k6.7 8 8 . 3 95.0 127 806 5b.9 0 1 2 7 . 2 5 1 .5 101 3.3 33-3 6 8 . 3 93-3 53 2bo 50 17 118 0 0 3-1 95 1 5 . 0 k S . 3 8 5 . 0 9 6 . 7 128 307 55 . 3 0 127 .15 2.2 83 0.0 35-0 75-0 9 6 . 7 5* 715 bo b8 117 23 k.b 70 3 . 3 3 0 . 0 75-0 96.7 129 8 0 8 5k.6 7 1 2 7 . 2 5 2.k 82 5-0 33-3 8 3 . 3 100 .0 55 37 5b 08 122 05 2.2 91 6.7 5 6 . 7 91.7 9 8 . 3 130 809 53 - 5 0 1 3 2 . 2 5 .b 135 3-3 35 . 0 93-3 100.0 56 39 5k 08 122 05 2.1 93 3-3 5 1 . 7 8 8 . 3 93-3 131 8ko 5 5 . 0 8 129.b2 .1 132 3-3 38-3 8 0 . 0 9 8 . 3 57 »3 5b 28 122 62 2.3 87 1.7 5 0 . 0 8 6 . 7 100.0 132 k6k 5 b . 0 5 1 2 8 . 6 5 .1 139 3-3 »3-3 93-3 9 8 . 3 58 b6 5' 08 122 08 2-3 88 3-3 5 1 .7 93-3 9 8 . 3 133 5k5 k9 . 0 5 I l k . 6 7 5.0 59 3-3 21.7 73.3 9 8 . 3 59 >7 5» 08 122 08 2-3 83 I 8 . 3 b6.7 9 0 . 0 9 8 . 3 13k 576 5 0 . 2 5 1 1 7 . 3 3 k.2 70 3-3 55-0 95-0 100.0 6 0 bo 55 50 121 58 2.0 86 3-3 2 8 . 3 91.7 l o o . o 135 8 5 8 k9.00 1 1 6 . 6 7 k.2 78 0 . 0 I 6 . 7 73-3 9 8 . 3 61 66 51 67 120 17 k.6 52 3.3 2 5 .0 86.7 93-3 136 868 k9 . 0 5 116.97 *-3 75 0 . 0 3 0 . 0 86.7 9 8 . 3 62 905 51 58 119 85 l .k 125 3-3 51.7 86.7 9 8 . 3 137 838 5 0 . 0 3 117.75 5.0 53 0 . 0 2 5 . 0 7 0 . 0 9 1 . 7 63 281 53 10 132 05 .k 138 1 .7 b6.7 9 0 . 0 l o o . o 138 539 k9 . 9 2 I l k .77 k.5 65 1.7 3 5 . 0 91 . 7 100.0 6b 28b 51 18 125 57 .9 139 3.3 5 6 . 7 95 . 0 l oo .o 139 5ko k 9 . 9 2 I l k . 9 2 k.9 s6 3-3 3 5 . 0 75.0 9.8.3 65 296 5k 17 122 17 2 . 2 90 6 . 7 »3-3 8 3 . 3 l oo .o lko 5 U k9 . 9 2 I l k . 7 3 5-1 51 0.0 31-7 15-0 9 3 . 3 66 336 50 25 • 115 75 5-0 52 1 .7 2 6 . 7 73-3 9 6 . 7 l k l 5»2 k9 . 9 2 I l k .75 k.6 63 1.7 3 6 .7 8 3 . 3 100. 0 67 3 bo 50 53 115 92 k.5 61 3.3 6 0 . 0 9 0 . 0 9 8 . 3 lb2 575 5 0 . 2 5 118.00 3-8 79 1 .7 33-3 31.7 100 .0 68 355 5b 23 127 37 2 . 0 ob 1.7 *3-3 81.7 9 3 . 3 l k 3 6 2 3 57.1? 121 .83 2-7 60 5 . 0 6 0 . 0 93-3 1 0 0 . 0 69 359 5k 17 121 50 2.7 79 0 . 0 16.7 6 5 . 0 9 3 . 3 lbb 6 3 6 bo.77 1 1 5 .ko 5 -0 55 0 . 0 2 0 . 0 63.3 91 . 7 70 36b 5* 93 127 k2 2.0 90 13-3 75.0 9 8 . 3 l oo .o lb5 6b l 5 0 .57 119.k8 k.9 52 0.0 16.7 68 . 3 9 6 . 7 71 365 5» 93 127 b2 2.0 90 0 . 0 36-7 8 5 . 0 9 8 . 3 lkf, 6bb 5 0 . 6 3 11.3.00 k.o 72 11.7 55 - 0 36.7 100.0 72 366 5» 93 127 b2 2.0 90 3-3 b5.0 8 8 . 3 9 8 . 3 lb? 65b k9.10 I l k . 6 7 5.6 k = O.D 2 5 . 0 B 5 . 0 100.0 73 369 5k 93 127 b2 2.0 90 0 . 0 11.7 73.3 9 8 . 3 U S 6 5 5 k9 . 0 8 115-78 k-5 70 0.0 2 6 . 7 71 .7 9 8 . 3 7b 371 5k LO 127 20 3.3 66 0 . 0 2 3 . 3 8 0 . 0 9 6 . 6 lk9 657 b 9 . 2 5 116 .23 k.8 63 3.3 2 6 . 7 76.7 9 6 . 7 75 373 5» OS 127 25 2 . 6 82 3-3 b5.0 9 0 . 0 100.0 I 5 0 85 7 k9 . 2 5 117.75 5-5 k? 0 . 0 3 0 . 0 73-3 0 0 . 0 66 TABLE 12 PERCENT DORMANT ON 6 DIFFERENT DATES DURING THE SECOND YEAR IN THE NURSERY Prov. Reg. No. Ho. Lat. Long. Elev. Dava Jun.30 Jul.7 J u l . l b J u l . 2 1 J u l . 2 8 Aug.* Prov. Reg. 5 6 7 8 9 10 11 12 13 lk 15 50 . 8 »2 • 9 3 bol 7 k 6 722 353 388 5 2 2 523 380 358 370 737 3«7 3 7 " 355 " 2 9 . 2 7 .18 I 5 0 1 « 7 7 U 7bo 738 720 3 3 " 3 - 3 393 3*S 387 7«7 710 6 9 9 701 3bk 3>1 3»5 709 337 773 8 9 8 732 338 335 779 385 51 " 3 1 b l 725 2 bO 715 37 39 »3 be »7 • 9 66 905 281 28b 296 336 3bo 355 359 36b 365 360 3«9 371 373 5 6 . 5 0 5 5 . 6 7 5 k . 5 0 5 b . 50 5 k . 3 3 5 b . 10 5 » . ° 7 5 ' . 9 3 5 3 . 6 7 5 2 . 3 3 5 2 . b2 55.60 55.»7 5 5 . 3 3 51 . 12 5 * . 93 5 > . » 5 5 « . 1 3 51 .77 51 . 7 0 51.17 b 9 . 9 ? "9.17 5 0 . 8 7 51 . 6 2 5 0 . 7 0 5 0 . J 8 b9.oo b 9 . 8 ? » 9 . 9 2 b9.92 5 U . 3 7 » 9 . 3 3 k 9 . 3 7 •9.17 5 0 . I 3 • 9 . 5 8 • 9 . 3 " • 9 . 0 8 51 . 13 • 9 . 5 8 5 3 . 3 3 51 .12 • 9 . * 2 • 9 . 7 5 k 9 . 3 3 5 b . l o 5 b . 2 5 5 k . 3 3 5 » . l 2 5 3 . k 2 5 0 . 6 5 50.17 b 9 . b 8 5 * . 0 8 5 * . 0 8 5 " . 2 8 5k.03 5 b . 0 8 5 5 . 5 0 5 1 . 6 7 51 . 5 8 5 3 . 1 0 51.18 5k.17 5 0 . 2 5 5 0 . 5 3 5 k . 2 3 5k .17 5 * . 93 5 k . 9 3 5 * . 9 3 5 k . 9 3 5 k . 1 0 5 b . 0 8 1 2 1 . 1 0 1 2 2 . 2 0 1 2 b . 2 5 1 2 2 . 6 7 1 2 2 . 6 7 1 2 2 . 0 5 1 2 1 . k5 127."2 1 2 2 . b2 1 2 1 . 6 7 1 2 1 . k j I 2 7 . 8 3 127 .83 1 2 6 . 6 3 118 .25 1 2 7 . k 2 126 .90 127 .15 120.17 120.17 1 2 0 . 2 5 I 2 0 . 5 8 1 2 0 . 5 8 I I 9 . 8 3 119 .83 119 .50 119 .75 116.75 116.27 119.77 118.58 1 2 2 . 5 0 117 .25 l i b . 5 8 116.13, 115.k3 I I 5 . 9 8 116 .00 116 .75 117 .08 117 .80 122.17 117 .25 I I 8 . 0 3 117 .00 116.13 1 2 2 . 0 5 1 2 2 . 2 5 1 2 3 . 0 8 1 2 2.8 3 1 2 2 . 6 7 U 7 . 5 3 I I 8 . 0 0 1 1 7 . 2 3 1 2 2 . 0 5 1 2 2 . 0 5 1 2 2 . 6 2 1 2 2 . 0 8 1 2 2 . 0 8 1 2 1 . 5 8 120.17 119 .85 1 3 2 . 0 5 1 2 5 . 5 7 122.17 115 .75 U 5 . 9 2 127 .37 1 2 1 . 5 0 1 2 7 . k 2 127.k2 1 2 7 . k 2 127.k2 1 2 7 . 2 0 1 2 7 . 2 5 2 . 3 k . 3 2 . 0 2 . 7 3 - 0 k.6 3 . 5 3 - 8 k.o 75 102 101 72 51 7« 73 76 81 66 125 67 b8 87 66 7k 76 8 0 76 66 65 61 83 k6 6 8 53 68 57 51 95 83 82 90 106 67 91 93 87 88 88 86 52 125 138 139 9 0 52 61 9b 79 90 90 5.0 I 8 . 3 5.0 2 1 . 7 2 0 . 0 16 .7 3 3 . 3 1 1 . 7 15 .0 3 6 . 7 5 . 0 3 3 . 3 5 . 0 I 6 . 7 1 . 7 1 . 7 31.7 3 6 . 7 31.7 I 6 . 7 2 3 - 3 5 0 . 0 15 .0 k l . 7 2 1 . 7 3 6 . 7 3 - 3 0 . 0 3 . 3 bo.o 7 5 . 0 b 8 . 3 2 5 . 0 5 3 - 3 2 8 . 3 3 0 . 0 6 0 . 0 k 6 . 7 3 8 . 3 3 1 . 7 71 .7 5 6 . 7 5 0 . 0 5 0 . 0 * 3 . 3 bo.o k l . 7 5 0 . 0 7 0 . 0 2 3 . 3 15 .0 1 3 . 3 I 8 . 3 2 3 - 3 • 3 - 3 3 1 . 7 k 6 . 7 13 .3 2 8 . 3 2 0 . 0 1 0 . 0 . 3 6 . 7 2 0 . 0 ko.o 1 .7 0 . 0 0 . 0 2 5 . 0 bo.o U . 7 1 5 . 0 2 3 - 3 8 . 3 6 . 7 1 0 . 0 6 . 7 3 1 . 7 1 3 . 3 • 3 . 3 6 3 . 3 k l . 7 7 0 . 0 6 1 . 7 5 5 . 0 7 8 . 3 2 5 . 0 9 0 . 0 3 5 . 0 5 3 . 3 3 3 - 3 2 5 . 0 91 .7 7 5 . 0 7 6 . 7 7 0 . 0 7 3 . 3 8 3 - 3 5 6 . 7 8 0 . 0 61 .7 8 0 . 0 3 0 . 0 6 . 7 2 5 . 0 8 3 . 3 9 5 . 0 81 .7 So.o 9 1 . 7 7 8 . 3 8 3 . 3 9 8 . 3 7 1 . 7 7 8 . 3 7 8 . 3 9 3 - 3 3 3 - 3 k l . 7 3 3 - 3 5 8 . 3 * 3 . 3 3 5 . 0 6 0 . 0 1 0 . 0 7 1 . 7 13 .3 3 3 - 3 U . 7 8 . 3 6 6 . 7 5 8 . 3 51 .7 3 8 . 3 5 5 . 0 7 6 . 7 31.7 61 .7 b5,o 53 .3 I 5 . 0 5 . 0 1 0 . 0 6 I . 7 8 6 . 7 7 8 . 3 6 0 . 0 8 0 . 0 5 3 - 3 6 8 . 3 8 3 . 3 6 5 . 0 6 5 . 0 6 0 . 0 8 6 . 7 8 5 . 0 6 6 . 7 8 8 . 3 8 6 . 7 9 8 . 3 7 3 . 3 7 3 . 3 6 I . 7 8 I . 7 83 . 3 b5.o . 6 . 7 3 6 . 7 3 8 . 3 k5.o 5 3 . 3 5 5 . 0 7 3 . 3 25 . 0 k 3 - 3 5 0 . 0 2 5 . 0 5 8 . 3 k 3 - 3 6 6 . 7 0 . 0 0 . 0 3 6 . 7 61 .7 5 8 . 3 2 8 . 3 5 5 . 0 3 0 . 0 1 3 . 3 3 6 . 7 2 3 - 3 6 0 . 0 3 5 . 0 81 .7 8 6 . 7 8 0 . 0 91 .7 91 .7 7 0 . 0 7 0 . 0 6 8 . 3 5 5 . 0 6 8 . 3 71.7 7 5 - 0 8 5 . 0 U . 7 6 I . 7 6 0 . 0 6 8 . 3 7 6 . 7 6 3 . 3 7 3 . 3 1 0 . 0 0 . 0 0 . 0 6 0 . 0 7 8 . 3 8 0 . 0 ' 6 5 . 0 7 0 . 0 5 6 . 7 3 5 . 0 6 8 . 3 5 3 . 3 7 5 . 0 5 6 . 7 9 0 . 0 91 .7 8 0 . 0 8 8 . 3 8 5 . 0 8 6 . 7 9 3 . 3 7 3 . 3 9 6 . 7 7 6 . 7 91 .7 6 6 . 7 7 3 - 3 9 8 . 3 91 .7 9 5 . 0 9 5 . 0 91 .7 91 .7 9 8 . 3 9 5 . 0 9 0 . 0 91 .7 7 3 . 3 ko.o 61 .7 9 3 . 3 9 6 . 7 9 0 . 0 8 6 . 7 9 3 . 3 93 .3 9 8 . 3 1 0 0 . 0 9 3 . 3 9 5 . 0 9 6 . 7 9 5 - 0 9 8 . 3 91 .7 1 0 0 . 0 9 5 . 0 9 6 . 7 83 . 3 9 3 - 3 1 0 0 . 0 9 3 . 3 9 6 . 7 91 .7 9 0 . 0 9 8 . 3 8 5 . 0 9 0 . 0 9 5 . 0 8 3 . 3 8 8 . 3 9 0 . 0 8 8 . 3 9 3 - 3 9 3 - 3 8 5 . 0 5 0 . 0 5 . 0 1 .7 91 .7 9 3 - 3 9 6 . 7 9 3 - 3 8 5 . 0 9 5 . 0 7 3 . 3 9 0 . 0 7 8 . 3 9 0 . 0 9 3 . 3 93.3 93-3 8 5 . 0 93-3 8 5 . 0 90.0 95.0 8 I . 7 96.7 3 5 . 0 93-3 73-3 8 6 . 7 100.0 95.0 95.0 95.0 95.0 91.7 9 8 . 3 95 .0 91.7 93-3 91.7 90.0 8 5 . 0 96.7 96.7 91.7 93-3 93.3 98.3 98.3 100.0 9 3 . 3 95.0 9 8 . 3 95.0 9 8 . 3 91.7 100.0 9 6 . 7 100.0 8 8 . 3 9 8 . 3 100.0 95.0 loo.o 91.7 91.7 loo.o 8 8 . 3 9 3 . 3 9 5 . 0 8 8 . 3 93-3 95.0 93.3 96.7 95.0 95.0 73-3 6 3 . 3 36.7 96.7 93.3 96.7 9 6 . 7 8 6 . 7 95.0 8 5 . 0 91.7 8 0 . 0 93-3 91.7 No. No. Lat. Long. Elev. Daya Jun.30 76 379 55 . 6 0 1 2 7 . 8 3 1.5 97 1.7 77 382 55 . 6 0 1 2 7.8 3 1.5 97 0.0 78 389 5k.08 122 .08 2 . 2 91 16.7 79 390 5k.08 122 .08 2.2 91 I 8 . 3 8 0 392 5 0 . 0 7 119.68 k.7 6 0 kJ.J 81 39k 5 0 . 0 2 119 .70 5.5 k2 k3-3 82 395 50.12 119 .70 k.7 59 35.0 83 kl6 5 0 . 8 5 119.87 k.6 57 k8. 3 8k b l 7 5 1 . 2 3 120 .25 k.o 68 2 5 . 0 85 b26 5k.67 128.75 .7 121 0 . 0 86 k?8 51.73 120 .08 k . 2 6 0 kl.7 87 k 3 0 5 0 . 6 7 119 .58 k . 2 67 2 8 . 3 S3 bk 7 51 . 0 0 119.87 k.7 5k 36.7 8 9 k5k 5k.93 127.k2 2 . 0 90 5 . 0 9 0 k88 k9 . 0 7 120.88 3-5 93 26.7 91 b99 5k.08 122.33 2.0 95 U.7 92 5 2 5 5k.75 128.75 .8 118 0 . 0 93 526 55 . 0 8 129.33 .7 118 0.0 9k 529 5 3 - 6 7 132-33 .1 l k l 0 . 0 95 691 b 9 . l 2 120.88 k.o 82 3 5 . 0 96 692 52.37 126.75 .1 lk9 0 . 0 97 703 k9.k 3 116.13 5.1 55 5 1 . 7 9 8 718 5k.13 122.12 2 . 0 95 16.7 99 719 5 k .18 122.12 2 . 0 95 11.7 100 723 5k.25 121.92 2.7 78 15.0 101 7k2 k 9 . l 3 1 2 5 . k 2 . 2 168 0 . 0 102 7*5 5 2 . 2 0 1 1 9 . 2 5 3-5 73 33-3 103 7k8 5 0 . 0 5 127 .05 .1 16k 0 . 0 lob 766 b 8 . 8 3 12k.75 -3 167 0 . 0 105 767 5 3 . 0 8 1 3 2 . 0 8 • 3 lko 0 . 0 106 769 5k.17 1 2 2 . 1 7 1.9 97 21.7 107 770 5k.25 122.33 2 . 2 90 20.0 103 771 5k.25 1 2 2 . 3 3 2.2 9 0 18.3 I 0 9 772 5k.17 1 2 2 . 2 5 2 . 2 90 I 8 . 3 UO 777 5k.17 122.17 1.9 97 2 8 . 3 U l 780 5*.08 121.88 2 . 0 95 10 .0 112 78* 5k.88 126.73 2.7 7k 3 1 . 7 113 789 5k.35 125 .k3 2.9 73 2 5 . 0 Uk 790 5k.07 127 .22 3 . 0 73 U.7 U5 791 55 .08 127.33 2.9 68 6.7 116 792 55.13 127.ko 1.0 111 3-3 117 793 55 . 3 0 127 .70 1.1 108 1-7 118 79k 5k.63 127 .20 2 . 5 30 5 -0 119 795 5 b .1? 125.k2 2.7 79 8 . 3 1 2 0 796 5 5 . 0 0 1 2 7 . 0 0 3.2 6 2 2 1 . 7 121 797 5k.80 1 2 9 . 0 3 1.9 93 1.7 122 7 9 8 5k.75 1 2 7 . 0 0 1.9 93 1.7 123 799 55 .03 1 2 6 . 5 0 2.3 82 10.0 12k 8 0 0 55 .28 1 2 7 .ko 1 .2 106 3-3 125 8 0 1 50.53 1 2 7 . 2 5 .6 150 0.0 126 805 55-k7 1 2 7 . 7 0 1 .2 I 0 5 3-3 127 806 5k.90 1 2 7 . 2 5 1.5 101 1.7 1 2 8 807 55 -30 127.15 2.2 83 1.7 129 8 0 8 5k.67 1 2 7 . 2 5 2.k 82 5 . 0 130 8 0 9 5 3 . 5 0 1 3 2 . 2 5 .b 135 0 . 0 131 8ko 55 .08 129.k2 .1 132 0 . 0 132 k6b 5 k . 0 5 1 2 8 . 6 5 .1 139 0 . 0 133 5k5 k9 . 0 5 U k . 6 7 5 . 0 59 6 8 . 3 13k 576 5 0 . 2 5 1 1 7 . 8 3 k . 2 70 b8. 3 135 8 5 8 k9 . 0 0 1 1 6 . 6 7 k . 2 18 33-3 136 868 k9 . 0 5 116.97 k-3 75 k6.7 137 888 5 0 . 0 3 117.75 5 . 0 53 k8-3 138 539 kq . 9 2 Uk.77 k.5 65 31-7 139 5ko b9 .92 Ilk.9? k.9 56 k l . 7 lbo 5kl k9 . 9 2 Uk.73 5.1 51 55-0 l k l 5k2 ko.92 Uk.75 k.6 63 2 6 . 7 lk? 575 5 0 . 2 5 118 .00 3 . 8 79 2 0 . 0 l k 3 6 2 3 57.17 1 2 1 . 8 3 2.7 6 0 35-0 lhk 636 k9.77 1 1 5 .ko 5.0 55 kl.7 l b 5 6kl 50.57 119.k8 k . 9 52 k5.o lb6 6bb 5 0 . 6 3 118.00 k.o 72 k5.0 lk7 6 5 k k9.10 lib.67 5.6 k5 61.7 lk8 6 5 5 k9.o8 115-73 k.5 70 ko.o lk9 6 5 7 ko . 2 5 1 1 6 . 2 3 k.8 6 3 k6.7 150 857 k9 . 2 5 117.75 5.5 k7 k8.3 1.7 1.7 2 6 . 7 2 3 - 3 5 5 . 0 6 5 . 0 bo.o 5 6 . 7 3 3 - 3 0 . 0 b 8 . 3 >3-3 5 6 . 7 8 . 3 3 0 . 0 2 3 . 3 0 . 0 0 . 0 0 . 0 3 6 . 7 0 . 0 7 3 - 3 2 3 . 3 I 8 . 3 2 0 . 0 0 . 0 3 8 . 3 0 , 0 0 . 0 0 , 0 2 5 - 0 2 8 . 3 2 8 . 3 2 6 . 7 3 5 . 0 2 5 . 0 k l . 7 3 1 . 7 2 3 . 3 3 . 3 6 . 7 1 . 7 3 . 3 8 . 3 2 6 . 7 1 .7 3 - 3 2 0 . 0 3 . 3 0 . 0 3 - 3 3 . 3 1 .7 6 . 7 0 . 0 0 . 0 0 . 0 3 . 3 1 0 . 0 b8 .3 k 8 . 3 8 0 . 0 8 3 . 3 7 6 . 7 81 .7 51 .7 1 .7 6 8 . 3 8 0 . 0 7 8 . 3 2 5 . 0 5 6 . 7 b6.7 0 . 0 1 .7 1.7 6 0 . 0 0 . 0 8 6 . 7 b5.o 3 5 . 0 3 8 . 3 0 . 0 7 0 . 0 0 . 0 0 . 0 0 . 0 5 5 . 0 k3 .3 6 0 . 0 k 5 . 0 5 5 . 0 b 5.o 6 3 . 3 6 5 . 0 k l . 7 31.7 16 .7 U . 7 2 3 . 3 2 1 . 7 bo.o I 8 . 3 18.3 k8 .3 1 0 . 0 0 . 0 8 . 3 5 . 0 3-3 2 3 - 3 0 . 0 0 . 0 0 . 0 8 3 . 3 loo.o 56.7 71.7 k 3 - 3 5 5 . 0 65.O k 3 - 3 k5.o 6 3 . 3 3 8 . 3 3 3 . 3 3 8 . 3 51 .7 5 8 . 3 5 3 - 3 7 0 . 0 k 6 . 7 56.7 6 3 . 3 7 0 . 0 6 8 . 3 86 .7 71 .7 61 .7 6 5 . 0 8 0 . 0 8 5 . 0 7 3 . 3 8 5 . 0 8 0 . 0 7 6 . 7 8 3 . 3 13.3 I 5 . 0 6 0 . 0 6 8 . 3 90.0 95.0 8 5 . 0 9 0 . 0 70.0 1.7 8 8 . 3 8 6 . 7 83.3 51.7 73-3 76.7 0 . 0 1 .7 1.7 8 0 . 0 0.0 9 0 . 0 7 0 . 0 55,3 6 8 . 3 0 . 0 7 8 . 3 0 . 0 0.0 0 . 0 7 1 . 7 61.7 76.7 8 0 . 0 81.7 6 5 . 0 7 0 . 0 83.3 63.3 55.0 2 6 . 7 I 8 . 3 k3-3 k8.3 55.0 23.3 33-3 61.7 I 6 . 7 0 . 0 15.0 18.3 25.0 k6.7 0.0 0.0 0.0 loo.o 8 3 . 3 86.7 9 5 . 0 90.0 8 3 . 3 8 8 . 3 95-0 8 8 . 3 75.0 8 6 . 7 9 0 . 0 8 8 . 3 8 5 . 0 100.0 91.7 91.7 91.7 6 5 . 0 6 3 . 3 9 5 . 0 9 5 . 0 91.7 96.7 9 0 . 0 9 3 . 3 81.7 2 3 - 3 9 0 . 0 91.7 8 8 . 3 81.7 91.7 oo.O 26.7 20 . 0 16.7 9 3 . 3 3 . 3 9 3 . 3 9 0 . 0 8 5 . 0 8 6 . 7 0 . 0 9 3 . 3 1.7 1.7 5 . 0 91.7 8 6 . 7 0 8 . 3 91.7 86.7 8 5 . 0 91.7 96.7 91.7 8 6 . 7 71.7 71.7 8 0 . 0 91.7 86.7 8 0 . 0 5 8 . 3 9 3 . 3 5 3 - 3 1 0 . 0 6 3 - 3 6 8 . 3 5 5 . 0 8 6 . 7 3 . 3 16.7 2 0 . 0 loo.o 91.7 91.7 9 8 . 3 96.7 9 5 - 0 9 8 . 3 1 0 0 . 0 9 8 . 3 8 8 . 3 9 8 . 3 91.7 9 0 . 0 90 . 0 1 0 0 . 0 9 5 . 0 9 6 . 7 9 6 . 7 7 6 . 7 7 6 . 7 9 5 . 0 9 8 . 3 9 3 . 3 9 6 . 7 9 0 . 0 9 5 . 0 8 6 . 7 7 6 . 7 91 .7 9 8 . 3 90 .0 9 3 - 3 9 5 . 0 93-3 9 0 , 0 61 .7 5 5 . 0 9 6 . 7 5 0 . 0 9 6 . 7 91 .7 8 5 . 0 oo.O 21.7 9 3 . 3 k5.o 3 5 . 0 6 I . 7 91 .7 93.3 9 8 . 3 91 .7 91 .7 9 3 . 3 9 6 . 7 93.3 91 .7 8 8 . 3 36 .7 3 0 , 0 8 5 . 0 9 8 . 3 9 3 . 3 8 5 . 0 7 8 . 3 9 3 . 3 8 0 . 0 6 8 . 3 8 0 . 0 7 6 . 7 7 6 . 7 9 5 - 0 6 6 . 7 7 5 . 0 7 3 . 3 1 0 0 . 0 9 3 . 3 9 8 . 3 9 8 . 3 9 6 . 7 9 8 . 3 9 8 . 3 loo.o 9 8 . 3 90 . 0 1 0 0 . 0 9 3 - 3 9 3 . 3 9 5 . 0 loo.o 9 5 . 0 9 6 . 7 9 6 . 7 TABLE 13 DATES OF FLUSHING AND DORMANCY OF 12 SPRUCE PROVENS CES ON TWO SOIL TYPES BOTH INSIDE AND OUTSIDE PLASTIC GREENHOUSE CM inside greenhouse RVS inside greenhouse CM outside greenhouse RVS outside greenhouse Block 1 \ Block 2 Block 3 Block k  No. of ' No. of No. of No. of Average no. of days days days days " days between D & F between between between between in a l l four p E F D F & D F D F & D F D F & D F D F & D environments 12 2. 1 Mar. 16 Jun. 2 78 Mar. 16 Jun. 2 78 Apr. 12 Jun, 2k 73 Apr. 9 Jun. 17 69 7*1.50 2 2. 2 Mar. 16 Jun. 30 106 Mar. 16 Jul. 30 136 Apr. 12 Jun. 30 79 Apr. 6 Jun. 17 72 98.25 8 2. 3 Mar.. 16 May 25 70 Mar. 16 Aug. 13 150 Apr. 12 Jun*. 21; 73 Apr. 6 Jun. 2k 79 93.00 k 2. 6 Mar. 16 Jun. 2: 78 Mar. 16 Aug. 5 lii2 Apr. 12 Jun. 2k 73 Apr. 15 Jun. 17 63 89.00 6 3. 0 Jun. 16 Jun. 10 86 Mar. 16 Jun. 2 78 Apr. 15 Jun. 17 63 Apr. 9 Jun. 10 62' 72.25 7 3. 7 Mar. 16 Jun. 2 78 Mar. 16 May 25 70 Apr. 21 Jun. 30 70 Apr. 9 Jun. 10 62 70.00 1 3. 9 Mar. 16 May 25 70 Mar. 16 May 25 70 Apr. 12 Jun. 2k 73 Apr. 2 Jun. 10 69 70.50 3. a. 5 Mar. 16 May 25 70 Mar. 16 May 25 70 Apr. 12 Jun. 17 66 Apr. 9 Jun. 10 62: 67.OO 9 k. 7 Mar. 16 May 25 70 Mar. 16 May 25 70 Apr. 12 Jun. 10 59 Apr. 9 Jun. 17 69 67.OO 10 u. 9 Mar. 16 May 25 70 Mar. 16 May 25 70 Apr. 21 Jun. 17 57 Apr. 6 Jun. 10 65 65.50 11 5. 3 Mar. 16 May 25 70 Mar. 16 May 25 70 Apr. 12 Jun. 10 59 Apr. 12 Jun. 10 59 61+.50 5 5. :7 Mar. 19 May 25 67 Mar. 16 May 25 70 Apr. 15 Jun. 2k 70 Apr. 12 Jun. 10 59 66.50 RVS - Robertson Valley soil, CM - California mix, E„- Elevation, P - Provenance, F - Flushing, D - Dormancy j A provenance was considered flushed when more than 50$ of seedlings scored were flushed. Dormancy was similarly assessed. Since flushing and dormancy in each environment was assessed on the basis of only 10 seedlings, i t is likely that assessments based on a larger sample would exhibit a period of growth (last column in above table) more strongly correlated with altitude than that indicated in table. T A B L E l U R E L A T I O N S H I P B E T W E E N G E R M I N A T I O N B E H A V I O U R A N D S E E D Q U A L I T Y A T 1 5 ° C •H & a> o c o •H •s ED3 E D U RD3+EDl| M D G P V G V A G P ED3 1.00000 y ED/4 ED3 + EDU -0.1,1835 -0.05863 1.00000 0.93125 1.00000 Coefficients of correlation between pairs of variables. M D G 0.28526 O.OU9U8 0.16883 1.00000 P V 0.28527 O.O/48O8 0.16729 0.99992 1.00000 •H IS G V 0.25U23 0.03729 0.11+297 0.9)4381 0.9U1457 1.00000 Q> A G P 0.2852U 0.0U958 0.16893 0,99999 0.99991 0.9U379 1.00000 See Table 2 for explanation of symbols, r > .159 is significant at .05 level of probability, r > .208 i s significant at .05 level of probability, n = 150 ON C o TABLE 15 RELATIONSHIP BETWEEN GERMINATION BEHAVIOUR AND SEED QUALITY AT 20°C, & C2 O •H 1 0) o •H 0 ) ED3 EDl|. ED3 + EDU MDG PV GV AGP ED3 1.00000 -O.i.1.835 -0.05863 0.00028 -0.0011+8 -0.01925 0.00022 EDlj. ED3-*EDlr MDG PV GV AGP 1.00000 0.93125 0.62638 0.55U22 0.51*915 0.6261(3 1.00000 0.68856 0.60851+ 0.59584 0.68859 Coefficients of correlation between pairs of variables• 1.00000 0.96208 0.95685 1.00000 1.00000 0.97851 1.00000 0.96210 0.95692 1.00000 See Table 2 for explanation of symbols, r > .159 is significant at .05 level of probability, r > .208 is significant at .05 level of probability, n = 150 VO i TABLE 16 RELATIONSHIP BETWEEN GERMINATION BEHAVIOUR AND SEED QUALITY AT 25°C I a1 cn o •H •H C5 ED3 EDli ED3+ED1* MDG PV GV AGP !ED3 1.00000 EDl* ED3 + EDI* -0.1*1835 -0.05863 1.00000 0.93125 1.00000 Coefficients of correlation between pairs of variables. MDG 0.01*1*01 0.63975 0.72080 1.00000 o PV 0.06279 0.52722 0.601*65 0.95355 1.00000 havi GV 0.01395 0.53961 O.59868 0.91*61*2 0.9781I1 1.00000 <L> AGP 0.01*1*00 0.63977 0.72081 1.00000 0.95353 0.9h63h 1.00000 See Table 2 for explanation of symbols, r > .159 is significant at .05 level of probability, r > .208 is significant at .01 level of 'probability, n = 150 TABLE 17 RELATIONSHIP BETWEEN GERMINATION BEHAVIOUR AND SEED QUALITY AT 30°C. ED3 ED4 ED3+EDU MDG PV GV AGP •p • H ED3 1.00000 Seed qual ;ED4 ED3 + EDU -o.41835 -0.05863 1.00000 0.93125 1.00000 Coefficients of correlation between pairs of variables• MDG -0.03705 0.46839 0.49994 1.00000 c o PV -0.01525 0.44094 0.47851 0.99561 1.00000 mina iavio GV -0.06880 0.40867 0.42157 0.93589 0.94692 1.00000 i n . G O J 0) a ,n AGP -0.03714 0.46835 0,1.9986 1.00000 0.99560 0.93587 1.00000 See Table 2 for explanation of symbols, r > .159 is significant at .05 level of probability, r > .208 is significant at .01 level of probability, n « 150 TABLE 18 RELATIONSHIP BETWEEN GERMINATION BEHAVIOUR AND FACTORS OF THE ENVIRONMENT TO O -P o 3 Q) O 8 PQ S3 L A D PV15 PVIO PV25 PV30 GV15 GV20 GV25 GV30 PVJ GVJ 1 2. 3 k 5 6 7 8 9 10 11 12 13 L 1 1.000 A 2 -.680 1.000 Coefficients of correlation between pairs of variables D 3 .306 -.906 .999 See Table 2 for explanation of symbols pvi5 4 -.363 .540 -.490 1.000 r > .159 is significant at .05 level of probability r > .208 is significant at .01 level of probability PV20 5 -.121 .255 -.263 .591 1.000 n = 150 PV25 6 -.223 .346 -.322 .449 .852 1.000 PV30 7 -.336 .463 -.408 , .485 .711* .823 1.000 GV15 8 -.383 .452 -.365 .928 .1(68 .308 ,350 1.000 GV20 9 -.087 .210 -.221 .536 .91*9 .762 .621 .464 1.000 GV25 10 -.165 .288 -.280 .384 .878 .977 .791 .259 .776 1.000 GV30 11 -.298 .377 -.318 .377 .61*5 .750 .954 .277 .593 .756 1.000 PVA25 12 -.281 .456 -.431 .462 .727 .911 .761* .3U7 .702 .931 .712 1.000 GVA25 13 -.181 .337 -.334 .348 .708 .890 .760 .239 .705 .91*3 . 752 . 966 1.000 — J ro O Q CQ C U O O © f-t " P fe co C fe W If S 3 § 8 co H CD <D fe CO M L A D FL6 FL16 FL20 FL 27 TABLE 19 RELATIONSHIP BETWEEN FLUSHING AND FACTORS OF THE ENVIRONMENT 1 2 3 h 5 6 7 L 1 1.000 -.680 .306 ,1U8 .l+OO .329 .099 A 2 1.000 -.906 -.UP -.395 -.1|2U -.261 D 3 .999 .096 .281 .361 .280 FL6 k 1.000 •60U .496 .205 F L I 4 5 FL20 6 FL27 7 Coefficients of correlation between pairs of variables 1.000 .793 .331 1.000 .!i93 1.000 See Table 2 for explanation of symbols r > .159 is significant at .05 level of probability r > .208 is significant at .01 level of probability n = 150 U) TABLE 20 RELATIONSHIP. BETWEEN GROWILH DURING SECOND YEAR AND FACTORS OF THE ENVIRONMENT fa s o g § s EH H O > fa H W3 8 EH O s 1 L A D SM10 SM214 SJ6 SJ20 SJYi; SJY18 PM10 PM2li PJ6 PJ20 PJY4 PJY18 1 2 3 h 5 6 7 8 9 10 11 12 13 lit 15 L 1 1,000 A 2 -.680 1.000 D 3 .307 -.906 .999 Coefficients of correlation between pairs of variables. SM10 U »030 .067 -.105 1.000 * See Table 2 for explanation of symbols SM21* 5 .029 -'.083 .091 .865 1.000 r > .159 is significant at .05 level of probability r > .208 is significant at .01 level of probability SJ6 6 .135 -.336 .358 .686 .928 1.000 n = 150 SJ20 7 •Hl2 -.U50 .503 .559 .81* .975 1.000 SJYli 8 •na -.1*81 .51*3 .516 .812 .960 .995 1.000 SJY18 9 .130 -.liBU .553 .506 .803 .951; •99 k .996 1.000 PM10 10 -.121 .566 -.666 .1*6 -f.002 -.329 -.486 -.528 -.51*0 1.000 EM2ii 11 -.121 .01*5 .011 -.1*78 -.032 .065 .IO4 .118 -.113 -.617 ' .'999 PJ6 12 .331 -.700 .720 -.269 -.037 .1)02 .529 .561 .567 -.853 ,2b$ 1.000 PJ20 13 .115 -.653 .782 -.217 .076 .359 .55U .589 .601* -.816 .701 0U722 1.000 PJY4 lh .027 -.372 .1*68 -.330 *.173 .003 .105 .202 .176 -.U83 .16U 0.389 .Uio 1.000 PJY18 15 -.097 -.096 .181 -.036 .007 .068 .117 .089 .171 -.206 .048 0.1U7 .228 -.300 1.000 ~0 TABLE 21 RELATIONSHIP BETWEEN GROWTH AND DORMANCY DURING FIRST AND SECOND YEAR AND FACTORS OF THE ENVIRONMENT L A. D SL1 RC1 SL1/RC1 DW1 SL2: RC2 SL2/RC2 DW2: DJ30 DJY7 DJYll* DJY21 DJY28 DAl* 1 2 3 1* 5 6 7 8 9 10 11 12 13 11* 15 16 17 L 1 1.000 A D 2 3 - .682 .308 1.000 - .906 1.000 Coefficients of correlation between pairs of variables SL1 RC1 SL1/RC1 1* 5 6 -9020 -.555 .1*27 -.51+8 .128 -.811* .721* .151* .813 1.000 .61*5 .806 1.000 .078 1.000 See Table 2 for explanation of symbols r > .159 is significant at .05 level of probability r > .208 i s significant at .01 l e v e l of probability n = 150 DW1 7 -.190 - .286 .1*83 .892 .81*2 .521 1.000 SL2 8 .085 - . 5 9 1 * .721* .936 .530 .810 .805 .999 RC2 9 -.163 -.1*05 .622 .835 .706 .531* .798 .878 1.000 SL2/RC2 10 .1*1+6 - .596 .517 .625 .020 .821* .1*28 .696 .276 1.000 DW2 11 - .038 -.538 .722 .868 .598 .653 .793 .909 .951* .395 1.000 DJ30 12 -.661 .857 -.733 - .538 .11*0 - .821 - .308 - .623 - . 3 6 7 . -.717 -.510 1.000 DJY7 13 - .635 .856 -.71*5 - .566 .117 - .839 -.328 - .61*5 -.1+00 -.711 -.51*3 0.981 1.000 DJYlU 11+ -.562 .862 - .796 -.616 .065 - .857 -.361 - .678 -.1*61 -.671 - .603 0.928 .951 1.000 DJY21 15 -.lt30 .827 - .828 -.698 - .050 -.871 -.1(28 -.733 - .560 -.623 -.682 0.81*3 .869 .953 1.000 DJY28 16 -.131 .699 -.831* - .768 -.293 -.752 -.523 -.752 -.735 - .687 - . 8 0 0 0.607 .638 .760 .875 1.000 DAl* 17 - . 0 8 9 .656 - .803 - .727 - .271* -.713 -.1*97 -.672 -.61*1* -.361+ - .720 0.583 .610 .720 .813 .923 1.000 TABLE 22 CORRELATIONS BETWEEN MEASURED VARIABLES INCLUDED IN THE PRINCIPAL COMPONENT ANALYSIS PVA25 SU SL1/RC1 FLU* FL20 SL2 RC2 SL2/RC2 DW2 PM10 P J 6 PJ20 PJYl* DJ30 DJY7 DJYll* DJY21 1 2 3 1+ 5 6 7 8 9 10 11 12 13 11+ 15 16 17 PVA25 1 1.000 SL1 2 -.227 1.000 Note the high correlation between dormancy and measurements of growth and the relatively low correlation between flushing and the same measurements. SL1/RC1 3 - . 3 7 1 .807 1.000 See Table 2 for explanation of symbols FLU* k -.221* .151* .320 1.000 r > .159 is significant at .05 level of probability .365 r > .208 is significant at . 0 1 level of probability FL20 5 - . 3 6 7 .232 .793 1.000 n = 150 SL2 6 -.261* .936 .811 .201* .310 1.000 RC2 7 - . 2 1 6 .836 .538 - . 0 2 5 .111 .880 1.000 SL2/RC2 8 -.211 .627 .821* .1*63 .1*53 .697 .282 1.000 DW2 9 - . 2 5 7 .868 .651* .117 .239 .910 .955 .397 1.000 FMIO ID .333 - . 6 2 3 - . 6 2 1 .270 .123 -.61*9 - . 6 7 0 - . 2 9 1 - . 6 7 9 1 .000 P J 6 11 - . 2 7 1 .630 .722 -.022 .035 .679 .606 .1*57 .660 -.852 1.000 PJ20 12 -.391* .751 .711* .031 .178 .760 .750 .380 .796 - . 8 0 9 0.723 1.000 PJYU 13 -.251 .337 .328 - . 0 0 1 .11*2 .352 .1*11* .070 .1*1*9 - . 5 1 6 .386 .1*09 1.000 DJ30 111 .1*00 .51*2 -.822 -.1*03 -.1*02 - . 6 2 7 - . 3 7 7 -.71U - . 5 1 6 .501* - . 6 7 5 - . 5 7 0 -.211* 1.000 DJY7 15 .367 - . 5 7 0 -.81*1 - . 3 8 0 - . 3 8 7 -.61*9 -.1*08 - . 7 0 9 -.51*8 .51*3 - . 7 0 8 -.599 - . 2 3 7 . 980 1.000 DJYll* 16 .367 -.621 -.859 -.357 -.385 - . 6 8 2 -.1*72 - . 6 6 8 -.610 .613 -.760 - . 6 9 8 -.311* .927 . 950 1.000 DJY21 17 .369 - . 7 0 3 - . 8 7 2 -.253 - . 3 1 1 -.736 -.569 -.620 -.687 .718 -.803 - . 8 0 2 - . 3 9 2 .81*1 .867 .953 1.000 77 TABLE 23 WEIGHTING FOR ORIGINAL VARIABLES IN COMPUTED COMPONENTS Variable 'Components 1 2 3 h 1 PVA25 -.424 -.179 -.002 .676 2 SLI .85U -.175 .310 .250 3 SL1/RC1 .922 .160 -.094 .116 h FUU .275 .825 .321 -.056 5 FL20 .367 .1)15 .426 -.023 6 SL2 .903 -.120 .299 .223 7 RC2 .7U5 -.bkh .424 .080 8 SL2AC2 .701 .1*1 -.036 .348 9 DW2 .839 -.299 .377 .057 10 PKLO -.760 .521 .226 .160 11 PJ6 .827 -.256 -.300 -.001 12 PJ20 M -.286 .O46 -.11U 13 PJIU .1*1 -.287 .171 -.551 lh DJ30 -.839 -.31* .321 -.012 15 DJY7 -.861 -.303 •321 -.029 16 DJY1U -.902 ^.219 .279 .018 17 DJY21 -.930 -.051 .212 •OiO. See Table 2 for explanation of symbols, and pages 32, 33 and Sh for a dis-cussion of the significance of this table. TABLE 2k PERCENTAGE OF VARIATION ACCOUNTED FOR BY FOUR PRINCIPAL COMPONENTS. Component Percentage of Cumulative variation percentage accounted for 1 57.85 57.85 2 1U.92 72.77 3 9.67 8O.I4J4. k 6.56 87.OO 79 DISCUSSION From Tables ll* to 22, which show the correlation coefficients be-tween pairs of variables, i t is clear that environmental pressures asso-ciated with altitude have been the principal selective pressures resulting in the adaptation of spruce populations to a wide range of environments* Furthermore, i t appears that these factors of the environment associated with altitude vary progressively, resulting in a clinal pattern of variation (Fig. 8). Because of the inadequacies of available climatic data for British Columbia (Chapman 1952, Chapman and Brown 1966) i t is not possible to obtain climatic data for the place of origin of each of the 150 spruce provenances groxra at Cowichan Lake, or even for a small proportion of these provenances. Nevertheless, despite the inadequacies of the climatic data an attempt is made to explain the pattern of variation in the iimnature spruce populations grown at Cowichan Lake in terms of the environment as determined by altitude at their place of origin. In regard to provenances from the area of the Nass and Skeena rivers the variation pattern is explained in terras of the en-vironment as determined by continentality. There is sufficient evidence from Langlet's work (Langlet 1963a) to warrant the conclusion that, in the interpretation of the growth behaviour of diverse provenances grown in a uniform environment, the day length on the f i r s t day of the year which shows an average temperatures of plus 6°C.(li30F) is an extremely valuable index of the environment at the place of origin of these provenances. Consequently this index is used in the present study, and will henceforth be referred to as day length at 1 J 3 0 F or DLU3. Cowichan Lake is situated on Vancouver Island at ca.lat. k9°00x, and has an average frost free period of II4.7 days (Connor 19k9) • The annual c l i -matic pattern at this station is given in Figs. 9 and 10 and compared with the climatic pattern at meteorological stations at increasing altitudes (Fig. 10) and decreasing continentality (Fig. 9) . Figs. 11 to 18 show the relationship between temperature and photoperiod at different ecological zones in the interior of British Columbia, and at the Cowichan nursery, and compare each of these zones with the photoperiod and temperature regime at Cowichan Lake during growing season of 1966 "^. It will be seen from Figs. 11 to 18 that DLu3 at Cowichan Lake is generally shorter than that of high elevation provenances from the interior. The lower the elevation of the interior provenances the closer the line AB and CD, which respectively indicate the DLU3 at Cowichan Lake and at a repre-sentative weather station. For example at Vavenby, which is a DOT2 station of the same elevation, and is very close to the Birch Island provenances (25 and 62, elev. I4OO f t . , lat. 5l°35', long. n 9 0 5 0 ' ) the DLii3 is almost identical with that of Cowichan Lake (Fig. 11). The percent dormant on six different dates is given in Table 12 for a l l provenances, and in Figs. 11 to 13 for a number of representative high and low elevation provenances. From Figs.. 11 to 13 i t is clear that high elevation provenances entered dormancy during the 1966 growing season at Cowichan Lake when temperatures were increasing, but the photoperiod had begun to decrease. Low elevation provenances also entered dormancy when temperature was increasing, but at a much later date than the high elevation provenances. For example, the Clearwater provenance (61, lat. 5u°U0', In determining photoperiod for a given latitude i t was not considered necessary to adjust for altitude. ^ Department of Transport weather s t a t i o n . elev. 46OO ft.) showed 35 percent dormancy on June 30, when the Birch Island provenance from approximately the same latitude, but lower elevation, showed no sign of dormancy, and showed only 5 percent dormancy on July I4 when the Clearwater provenance was 65 percent dormant (Fig. 11). Similarly provenance I4.2 from a region south of Prince George, but at 3bP0 f t . showed 28 percent dormancy on June 30 when temperatures were increasing. Provenance 10 from Soda Creek, elev. 2200 f t . , showed only 3 percent dormancy on this date. As in the previous example, temperature was increasing and photoperiod decreasing at the outset of dormancy (Fig. 13). These examples are typical of the general pattern of the outset of dormancy in a l l populations from the white-Engelmann complex growing at Cowichan Lake. The pattern varies, however, in populations from the general region of the Nass and Skeena rivers. It seems clear that the intrusion of Sitka spruce genes, as a result of decreasing continentality, influences the pattern of variation in these populations. For example i t will be seen from Fig. 12 that provenance 128 from the region of Hazelton (lat. 55018', long. 121°19%t elev. 2200 ft.) and provenance I4 from Babine Lake (lat. 55°20!, long. 126°38', elev. 2300 ft.) differ significantly only in regard to longitude. Yet.there is a striking difference in time of entering dormancy. On June 30, the Hazelton provenance showed only 1 percent dormancy while the Babine Lake provenance showed 25 percent dormancy.,/ In regard to time of entering dormancy the Hazelton provenance is typical of a l l the provenances from the Nass and Skeena river drainages. A l l of these provenances entered dormancy much later than provenances from east of this general region. As in the previous two examples the Babine Lake and Hazelton provenances entered dormancy when temperature was in-creasing at Cowichan Lake, but photoperiod decreasing. Because of the clinal pattern of variation in the time of entering dormancy (Fig. 8) the relationship between temperature, photoperiod and degree of dormancy illustrated in Figs. 11 to 13 will vary similarly for a l l other provenances. It seems, therefore, that, as in the case of the many other tree species referred to in a previous section, the onset of dormancy in white spruce and its related forms is photoperiodically de-termined. r 83 THE ECOLOGICAL SIGNIFICANCE OF PHOTOPERIODIC ADAPTATION IN WHITE SPRUCE. In the last section evidence was provided in regard to the photo-periodic control of dormancy in the spruce populations grown at Cowichan Lake. In this section an attempt is made to clarify the ecological signi-ficance of, photoperiodic control of dormancy in these populations, and to . indicate the manner in which the photoperiodic stimulus may be received by the spruce seedling, and translated into metabolic activity leading to dormancy. It is well known that organs which are growing are much more sus-ceptable to injury by freezing temperatures than organs which are dormant. It i s also generally known that the decrease of growth activity in the buds of perennial plants "manifests itself in a lowered capacity to react imme-diately by continued growth to certain conditions which are growth-promoting during the active phase" (Vegis I 9 6 3 ) . The capacity to enter dormancy well in advance of the average occurrence of early f a l l frosts confers a survival advantage on that proportion of a population of trees which possesses i t . For such trees have the capacity to survive temperatures damaging to those which continue growth. The selective action of f a l l frosts operates both on seedlings and mature trees. But i t is clear that a temperature which is lethal for seedlings may only injure, or have very l i t t l e effect on mature trees (Parker 1°63). Most coniferous trees including white spruce are predominantly cross pollinators. Consequently, the individuals tend to be heterozygous and the population heterogenous. There i s , therefore, a wide range of genetic variability within any one population even though that population in nature occupies a relatively uniform environment. There is l i t t l e doubt that this intra-population variability is itself an adaptive characteristic for i t buffers the population against yearly fluctuations of the environment, as well as long term gradual changes in climate. It is this pool of genetic variability which is acted upon by the selective pressures of the environment. The date of-the occurrence of the fi r s t killing frost in any one environment will vary from year to year, and each year will exercise its maximum effect at the seedling and sapling stage. Now, that proportion of the population which enters dormancy at an early date, as a result of a capacity to receive and transmit a stimulus from a factor of the environment, other than temperature, will avoid the lethal effects of freezing temperatures which eliminate, i.e. select, those seedlings not possessing this capacity. These seedlings will survive, therefore, and pass on the capacity to their progenies. The factor of the environment must be continuous and increasing in intensity so that i f temperatures should rise again above the critical point growth will not consequently be induced. As indicated in Figs. 11 to 18, the one single factor of the en-vironment which shows a progressive, unfluctuating increase with the passing of time from summer to f a l l , and which does not fluctuate from year to year, is the length of the dark period, or nyctoperiod - and i t is the nyctoperiod and not the photoperiod which is the critical factor (Nitch 1°63 p. 176). If a portion of a population of spruce seedlings possessed a leaf pigment which, as a result of decreasing day length (increasing nyctoperiod), receives and transmits a stimulus resulting in dormancy, such seedlings would have a sur-vival advantage over those not possessing the pigment, for day length - and this is the important point - decreases before temperatures decrease (see Figs. 11 to 18). Hendricks and Borthwick (1°63) have shown that a bright blue protein termed phytochrome regulates many aspects of plant growth and de-velopment, including dormancy. Phytochrome has two interconvertable forms with absorption maxima at 660 top. and 730 mu. Hendricks and Borthwick re-presented the conversion reaction as follows: 660 mji darkness ^ ^ P 660 P 660 P 730 -^730 mu and gave four ways in which phytochrome may link the plant to its environment. These are given below in f u l l : (1) It changes with light quality independently of the intensity above low values. (2) It reverts in darkness from P 730 to P 660 and thereby de-termines photoperiodism. (3) The substrates upon which i t acts and the products that i t forms depend upon photosynthetic and reserve metabolic activity. (1+) The rates of the crucial reactions in which P 730 is involved, including its own dark transformation, are temperature dependent. The investigations of the manner and extent of the control of plant growth by phytochrome referred to above were not conducted on tree species. However, there is evidence that dormancy in trees is initiated as a result of the accumulation of growth inhibitors under short photoperiods (Giertych 196k page 298,Nitsch I963 page l8u, Steward 1963 page 206, Wareing 1956 page 205). Steward (1963) has reported that in conifer leaves and in buds, glutaraine and arginine levels f e l l with the onset of shorter days, lower temperature, and lower light intensities, and that as the glutaraine and arginine levels decrease, asparagine and proline increase. When plants enter into long days this trend is reversed. Metabolic changes such as these can have phenotypic expression. For example the onset of dormancy of the spruce populations at Cowichan Lakey particularly in the f i r s t year, was always accompanied by a vivid purple coloration of the leaves. From the work of Hendricks and Borthwick (1°63) i t appears likely that the metabolic processes leading to increased levels of inhibitors and eventual dormancy in white spruce seedlings are mediated by phytochrome, or a phytochrome-like substance, following reception of the stimulus of decreasing day length. It is possible, however, that the photoperiodic reaction is determined, not only by the dark conversion of P 730 to P 660, but is the result of an interaction between an endogenous circadian rhythm and phytochrome (Bunning l ° 6 l , Hamner 1963). For example, there is some indication that provenances from areas of short growing season, i.e., provenances from northern latitudes and/or high elevations, are more closely adapted to photoperiod than provenances from areas of long growing season, i.e., provenances from low elevations and/or low latitudes (Irgens-Moller 1957* Olsen et a l . 1959)• In the latter instance circadian rhythms may be of greater importance than photoperiod, though i t must be pointed out that the theories of Bunning in regard to circadian rhythms and photoperiod have not been universally accepted (Romberger 1963 p. 120). It is not, of course, possible to provide evidence concerning the effect of photoperiod on flowering in spruce from a study of geographic variation in immature spruce populations. There i s , however, evidence from other studies that photoperiod may have l i t t l e or no effect in regard to flowering in conifers. In a paper, which is of considerable significance in this connection, Mirov (1956) showed that there is no definite relationship between photoperiod and flowering in the genus Pinus. Most of the_pine..species of the world are represented at the Institute of Forest Genetics at Placerville, California, lat. 38°4U', and at the University of California at Berkeley, lat . 37 052 ,. In these two localities the longest day in summer is approximately 15 hours. Pine from north of these latitudes were not inhibited in the production of both male and female flowers. For example, Pinus sylvestris var. lapponica, which has a northern distribution between lat. 60°00' and 70°00', produced abundant male and female flowers at 29 years of age, despite the fact that the longest day in summer at latitudes spanning its natural range is approximately 19 hours. It is customary in Swedish forestry practice to take cuttings from trees of high elevations and high latitudes for grafting on stock near the coast where the climate is much less severe. The species concerned are Pinus sylvestris (Scots pine) and Picea excelsa (Norway spruce). Seed production is enhanced by this transfer, and i f flowering where under strong photogeriodic control i t is unlikely that such a result could be obtained. Kramer and Kozlowski (I960) Matthews (1963) and Anderson (1965) have reviewed the literature pertaining the flowering and seed setting in forest trees. From these reviews i t is clear that climate, other than photoperiod, edaphic conditions and the age of the tree are the principal factors affecting flowering and seed setting in conifers. Little evidence was provided indicating that photoperiod exercised as important an influence in flowering in conifers as i t obviously does in regard to dormancy. It is likely, therefore, that the ecological significance of photoperiodism in white spruce in British Columbia is confined to flushing and dormancy -particularly the latter - and is of l i t t l e significance in regard to flowering. Wareing (1956 p. 209) has pointed out that in regard to herbaceous species i t is generally assumed that the biological significance of photo-periodism lies i n the control of flowering, and that vegetative responses are secondary. He goes on to state: n±t does not follow that this is true also for woody species, however, in which the primary significance of photoperiodism may concern dormancy phenomena". This is an important hypothesis. It relates dormancy rather than flushing to photoperiod, and also gives dormancy greater significance than flowering as a photo-periodic response in woody species. The evidence presented above is in agreement with Wareing's re-marks concerning flowering and photoperiodism in woody species. Evidence will be presented in the following section which indicates that Wareing's special emphasis on dormancy in relation to photoperiodism in woody species is also essentially correct, at least in regard to the growth of immature spruce populations in British Columbia. 89 W RELATIVE IMPORTANCE OF FLUSHING AND DORMANCY AS ADAPTIVE FACTORS IN THE MICROEVOLUTION OF MITE SPRUCE In the last two s ections evidence was provided in regard to the photoperiodic control of dormancy in spruce, and the ecological significance of photoperiodic adaptation discussed. The manner in which the photoperiodic stimulus may he received by the spruce seedling and translated into metabolic activity leading to dormancy was also indicated. The question remains, however, as to the extent and importance of photoperiodic control of flushing as well as dormancy. A second and related question refers to t he strong correlation betx-reen factors of the environment and dormancy, and the relatively weak correlation between the same factors and flushing (Fig. 19). The writer has not found a satisfactory answer to these questions in the literature (reviewed by Vegis 1963), and because of this, and because of its obvious importance in relation to the latitudinal and altitudinal displacement of forest tree seed, the subject is treated here separately, and an explanation attempted. Surprisingly few studies of variation within tree species are designed to assess variation in times of flushing and dormancy. Indeed, much of our present knowledge concerning both of these vital phases in the growth cycle of forest trees comes, not from provenance tri a l s , but from physiological studies in controlled environments e.g. Wareing (1950 a, b, c; 1951, 1956), who has also stressed the silvicultural aspects of this work (1966). Wareing (1951 p. 51) has stated that there is no evidence that the breaking of dormancy in Scots pine in the spring under natural conditions is photoperiodically controlled. He pointed out that dormancy is readily bro-ken at any time from early January onwards simply by transferring the plants to warm conditions regardless of the length of the natural photoperiod. Pauley and Perry (195k) have shown that neither light nor its periodicity appears to be directly concerned in the breaking of dormancy in Populus. The onset of flushing occurred even in darkness. On the other hand, these authors conclude that the role of photoperiod in the annual growth cycle of Populus appears primarily to influence the timing of physio-logical processes concerned in the onset of dormancy. Morris et a l . (1957) have shown that the date of flushing in the same provenances of Pseudotsuga menziesii (Douglas fir) may vary as much as a month between years. They also reported that the time of flushing had no apparent relation to annual height growth. The results of Ching and Bever (I960) support the conclusion that flushing and height growth in Douglas f i r are not correlated. Burley (1966) has shown that flushing in Sitka spruce is controlled largely by temperature, and that bud formation in the same species is a response to photoperiod. In a study of the seasonal height growth of 5 conifers, including white spruce, Kozloxraki (1957) found that a l l trees started growth in the spring before danger of frost was over, and stopped growth in the summer usually long before f a l l frosts began. Hanover (1963), in a study of geo-graphic variation in Pinus ponderosa Laws, (ponderosa pine), found an identical situation in regard to flushing and dormancy, and has reported that a l l trees began growth in the spring before danger from frost has passed, and ceased growing before the beginning of f a l l frosts. Mitchell (1965) studied leader growth in U2 different conifers grox-ring at a single site in southern England. Two of his conclusions are 91 relevent to the present discussion. These are: (1) adverse climatic Condi-tions in the spring had no appreciable effect on the time of flushing (2) in the year of planting, trees usually began their height growth several weeks later than when they are established, but end at about the same time. These conclusions have particular significance when i t is considered that they are based on observations made during several growing seasons on 1|2 different^ species of conifers growing in a uniform garden. From the point of view of survival in white spruce - which was one of the species studied - during the f i r s t year of outplanting, and its obvious significance in regard to the re-lative importance of flushing and dormancy as adaptive factors in the same species, special interest is attached to the conclusion that the after effects of planting can delay flushing for several weeks whereas i t has no appreciable effect on the time of growth cessation. Fig. 1 ? shows the relationship between factors of the environment at the place of origin and flushing and dormancy in the white spruce po-pulations at Cowichan Lake. It will be seen that the correlation between dormancy and factors of the environment is high, and much higher than the correlation between flushing and factors of the environment. The correlation coefficient for dormancy on July l l * and altitude is . 8 6 2 . This is the highest correlation coefficient obtained between any one factor of the en-vironment, and a measure of growth behaviour. Furthermore, as will be seen from Table 21, the date of entering dormancy and dry weight are highly-correlated. The correlation coefficient between dormancy on July 28 of the 2nd year, and dry weight of two year old seedlings in - . 8 0 0 . The curves for dormancy and flushing of high and low elevation provenances, and for continental and maritime species are compared in Figs. 20 to 2 7 . It will be seen that there is very l i t t l e difference between high and low elevation provenances, and continental and maritime species in 92 the curves for flushing. On the other hand, there are striking differences between high and low elevation provenances, and between continental and maritime species in the curves for dormancy. Indeed the dormancy curve characterizes a provenance with considerable precision (see Fig. 22). From the data concerning the growth behaviour of 12 spruce pro-venances sown in the spring of 196k in the Cowichan Lake nursery (Table 1 3 ) i t will be seen that a l l 12 provenances inside the greenhouse flushed approximately 3 weeks before those outside the greenhouse, and a l l provenances from elevations above 3000 f t . inside the greenhouse entered dormancy by May 2 5 when temperatures were increasing, and moisture regime kept at optimum. Al l provenances from below 3 0 0 0 f t . inside the greenhouse showed considerable variation in time of entering dormancy. The average number of days between flushing and dormancy for provenances below- 3 0 0 0 f t . is approximately 9 0 days, and 7 0 days for provenances above 3 0 0 0 f t . The apparent discrepancy between Fig. 7 and Table 13 is attributable to the fact that a provenance was considered dormant when 5 0 per cent of the seedlings measured were dormant, even though one or two seedlings had not entered dormancy and continued shoot extension. There appears to be sufficient evidence both from controlled en-vironment studies, and from studies of periodicity both in the nursery and in natural stands of coniferous species, as well as from the results of the present study, to warrant the conclusion that flushing in vii it e spruce i s predominantly influenced by temperature, and that dormancy is predominantly under the control of photoperiod. This does not imply, of course, that temperature has no effect on dormancy, and that photoperiod has no effect on flushing. On the contrary i t is possible that both of these factors of the environment have some influence on each of these phases in the growth cycle of coniferous species. Similarly printer chilling exercises an import-ant influence on the plant's reaction to photoperiod (Nienstaedt 196?). Never-theless, the weight of evidence indicates that in regard to white spruce pho-toperiod exercises its predominant influence on growth cessation and dormancy, and temperature exercises its predominant influence on flushing. The following hypothesis is presented in regard to the apparent photo-periodic control of dormancy in white spruce and the lesser importance of photoperiod and greater importance of temperature in the control of flushing. It i s possible that this hypothesis can be extended to other tree species of the Pacific Northwest, e.g. the inland form of Douglas f i r . There are two major differences, which the writer has not seen pre-viously stated, between the environmental pressures operating on the plant in the spring, and those operating in the f a l l . In the spring the probabi-l i t y of occurrence of killing frost is decreasing with the passing of time. In the f a l l the probability of the occurrence of killing frost increases with the passing of time. Secondly in the spring temperatures and photoperiod are increasing simultaneously, whereas in the f a l l photoperiod decreases before temperatures decrease (see Figs. 11 to 18). Because the probability of the occurrence of lethal frost increases with the passing of time in the f a l l , i t is clear that in respect to dormancy a plant adapted to photoperiod has a greater survival advantage than one adapted to temperature alone. For the plant adapted to photoperiod will enter dormancy in advance of the plant adapted to temperature alone, and consequent-ly has a greater chance of avoiding the effects of freezing temperatures. In the spring a plant remains dormant until temperatures reach a certain thres-hold (approximately <43°F). Each day after the occurrence of temperatures say above h3°F the probability of the occurrence of lethal frosts decreases ra-pidly. This decrease is particularly rapid in areas of continentality. Therefore, no overwhelming survival advantage is confered on the plant in the spring by the photoperiodic control of flushing. These considerations also, to some extent, explain why dormancy is more closely adapted to the environment than is flushing. There is one other observation which may appropriately be made in this section. Fig. 8 shows the relationship between time of entering dormancy and elevation at place of origin of 150 spruce provenances. Though the pure forms of white, Engelmann and Sitka spruce and their intermediate forms are represented by these 150 ^provenances, there is no disjunct pattern of variation in time of entering dormancy, and a clinal pattern is obvious. It may be concluded, in regard to spruce species at least, that this is evidence justifying the assumption that environmental pressures which re-sult in microevolution, i.e. genecological differentiation, differ only in degree rather than in kind from the environmental pressures which result in macroevolution, i.e. speciation. Langlet (1963a)has concluded that the faculty for normal development and survival in Picea excels a (Norway spruce) is not conditioned by the time of flushing, but that on the contrary i t is the time of cessation of growth, and initiation of dormancy which determines its fitness for use in practical silviculture. From the evidence presented in the last three sections i t seems clear that this conclusion is equally valid in regard to the growth of white spruce in British Columbia. 95 'GERMINATION BEHAVIOUR IN THE LABORATORY Germination behaviour can be affected by a large number of factors other than those which are purely genetic, e.g. time of harvesting and methods of extraction and storage (Allen 1958, Roche 1965). For these rea-sons i t is obviously advantageous i f the experimental material is completely uniform, that i s , i f each provenance has been similarly treated, and i f the quality of the seed of each provenances is identical. This is rarely possi-ble, particularly when large numbers of provenances are involved; for almost certainly collections will have been made in different years, and the handling of cones and seed will also vary to some extent from year to year. There is one other major factor which will considerably increase the complexity of the problem. When forest tree seed is collected in natural stands, the processes of natural selection which would operate on that seed are bypassed. Consequently, seed thus collected contains a much greater range of genetic variability than is likely to be found in nature in the seedling progenies resulting from that seed had i t been allowed to propagate in its natural environment. These considerations also apply, of course, to the nursery propagation of collected seed. However, in this instance some selection will occur in the nursery bed, which in the optimum conditions of an incubator will not occur. The seed used in the present study is highly heterogenous in qua-l i t y . Only one non-genetic factor influencing germination behaviour i s , to some extent, accounted for, that i s , embryo development. The influence of other factors are unknown. Nevertheless, despite this heterogeneity in qua-l i t y some general trends are apparent in germination behaviour which can be attributed to selection and adaptation at the place of origin of the seed, rather than to the non-genetic factors discussed above. It i s clear from the results of this study that in assessing proven-ance difference in the germination behaviour of spruce, extreme temperatures are more effective than optimum temperatures (Table 18). Allen (1961), who obtained similar results with Douglas f i r , demonstrated that coastal and in-terior provenances of this species can be distinguished best on the basis of germination behaviour at 10 or 15°C. The effect of differences in embryo development on germination be-haviour is most pronounced at optimum temperatures, that is those tempera-tures at which seed is usually tested. At extreme temperatures, particularly at low temperatures, the effect of differences in embryo development appears to be overwhelmed by the effect of provenances differences (Tables Ik to 17). For example, certain provenances from the general region of the Nass and Skeena rivers behaved similarly to strictly coastal provenances, e.g. pro-venances 85, lat. 5U°U0', long. 128%5' (near Kitsum Kalum lake), provenance 116, lat. 55° 0 8', long. 127°24l (near Beaumont North of Smithers), in that they did not germinate at a l l , or had extremely low germination values at 15°C. The general relationship between germination values at 15°C and altitude is shown in Table 25• Germination behaviour is positively correlated with altitude at a l l four temperatures. This would indicate that in regard to germination be-haviour high elevation provenances are less narrowly adapted to temperature conditions than are low elevation provenances. In other words, high elevation provenances are capable of germinating over a wider range of temperatures than are low elevation provenances. This is also true in re-gard to species, for the Sitka spruce provenances did not germinate at a l l at 15°C. The relatively high germination vigor of the high elevation Engelmann spruce provenances is further illustrated by the data given in Table 26, which is compiled from the Woody-Plant Seed Manual (see Alexander 1958 p. 9). It is well known that interior provenances of Douglas f i r and lodge-pole pine germinate more rapidly than coastal provenances when incubated at the same temperatures (Allen 1958, Critchfield 1957, Roche 1962). This dif-ference in germination behaviour is the result of adaptation to continental and maritime climates. Similarly the differences in germination behaviour between high and low elevation provenances of white spruce can be attributed to the differential selection pressures prevailing in the subalpine and montane forest regions. The similarity in germination behaviour between the coastal pro-venances of Sitka spruce, and provenances from areas of coastal influence, but within the range of white spruce, suggest that there is an intrusion of Sitka spruce genes into the white spruce populations in this region. Since germination behaviour is habitat-correlated i t may be used in conjunction with measures of growth behaviour to discriminate clusters of similarity in the provenances studied. 98 TABLE 2 5 PERCENT OF PROVENANCES IN EACH OF FIVE ELEVATION GROUPS WITH A GERMINATION VALUE1 OF LESS THAN ONE AT 15°C Elev. (ft.) Percent 1 0 - 1000 100 2 1100 - 2000 U5 3 2100 - 3000 53 ll 3100 - 1+000 19 5 liioo - 5700 0 Germination value estimated i n the manner proposed by Czabator (1962) TABLE 26 AVERAGE GERMINATION CAPACITY OF ENGELMANN SPRUCE IN COMPARISON WITH OTHER SPECIES Species Germination Species Germination capacity capacity Engelmann spruce 69 Western white pine 54 Lodgepole pine 6k White spruce li9 Black spruce 61 Subalpine f i r 38 Sitka spruce 60 Grand f i r 28 Red spruce 60 Western larch 27 Western hemlock 56 Pacific silver f i r 2U 99 RESULTS PART B GEOGRAPHIC VARIATION IN MATURE POPULATIONS OF 1-JHITE SPRUCE Regression analysis of data resulting from sampling in the white-Engelmann spruce complex both in 19&3 and 196k showed that the index of cone scale morphology L1/L2 is highly correlated with altitude (Figs. 28 and 29). Fig. 3k shows the variability in cone scale morphology along an altitudinal transect at Stone Creek, 25 miles south of Prince George (lat. 53°UO', long. 122°2f>l)« This transect passes from montane white spruce forest to subalpine Engelmann spruce forest. Of the 10 measurements of cone scale morphology made the single measurement L1/L2 X L3 was most effective in distinguishing species and intermediate forms. Figs. 30 to 32 show the pattern of variation in this measurement for the 1963 and 1961; collections, and also for the miscellaneous collections. Sympatric populations of white and black spruce occur between latitudes 5U°00' and 60°00' (Figs. 35 to UU), and sympatric populations of white and Sitka spruce occur between latitudes 5U°30' and 55°30', and longitudes 127°30' and 128°U0'. Fig. 33 shows the pattern of variation in cone scale morphology in a longitudinal transect which passes from coastal Sitka spruce forest to montane white spruce forest. The actual samples used in constructing the line of shape for this transect are shown in Illus. 7. In a l l areas where white spruce is sympatric with any one of the other three spruce species which occur in British Columbia the line of shape of cone scales from these sympatric populations exhibited .characteristics of both species (Figs. 33, 3U, 35 - U l ) . The means and geographic origin of foliage samples are given in Table 27, and the results of the discriminant function analysis of a l l cone scale data for 1°63 and I96I4. collections are illustrated in Figs. hS and Ij.6. In the black-white spruce complex along the Alaska highway every gradation in bark type was observed from that of pure trhite spruce to that of pure black (Illus. lit). A spruce population exhibiting a bark type identical with that which characterizes Picea glauca var. Porsildii was observed at Telegraph Creek, lat. 57°52!, long. 131°12', elev. 1325 f t . (Illus. Ik). Individual trees and populations exhibiting atypical branching habit were located in numerous areas throughout the province (Illus. 13). z f t f t f t f t f t f t f t f t f t f t ' f t ' f t f t f t f t ' f t f t f t f t f t f t f t f t f t f t f t f t f t ' f t t f ' f t f t ' f t f t * f f t f t f t f t f t f t 1 1 f t f t t f t ' t f t t ' f t ' t t ' f t t ' t J t ' f t t ' t t f f f t ' f t ' f t ' f t - f t f t ' f t t f ( # f t ' f t ' f t f t ' f t ' f t f t f t f t ' f t f t ' f t f t ' f t f t ' f t t ' t f t t ' t ' f t f t f t t ' t ' t ' t ' f t ' f t ' f t t ' t f t ' f t ' f t f t ' f t f f f t ' f t f t f t ' f t t t f t ' f t t t ' t ' t ' t '# f t ' f f f t ' f t f t f t Mile 6k Alaska highway-No. Ik9 Lat. 5625 Elev. 3000 f t . ' f t f t ' f t ' f t t ' t ' f t ' f t t t t t f t t t t t t t t t t t f t 11 t f Birch Island 117 5135 moo f t . Manning Park 62 1905 U800 f t . Illus. 5 Cone scale morphology of pure white and Engelmann spruce samples (1U9 and 62) and intermediate form (117). Each column of scales and bracts represents a single tree. O f—' 1 'ft 'rt 't't t't 't't 't't rt t it •ft t't (rt r t * ¥ •t't •rt f t f t 't't f t 'ft f t f t f f f f f t f t f t f t f t •ft 'ft 't't •rt •rt t t tf i r t f t t f f t f t f t f t •ft t't f t f t tt r t t t r t f t f t f t f t f t f f f t •ft t't f t f t 't'9 f t f t f t •ft • t | | t r f f r t r t r*t f t t t f t f t f f t t f t f t 'ft 'tt f f_ f t f t t t f t Nr. Telegraph Creek No. UjO Lat. 5814 Elev. 2300 f t . Alaska highway l! l3 5939 1700 f t . Fort Nelson ihS 5837 1600 f t . Illus. 6 Cone scale morphology of sympatric populations of white and black spruce in the boreal forests of Northern British Columbia. o ro f t •ft f t 'ft f t f t ' ft 'ft •ft f t f f '«'# 'ft •ft f t f t f t •ft •t't f t f t f t 'ft •••• »t* t f t f t 'ft f t •t't f t f t 't't ' f t •t't ' ft f t f t •ft f t f •f f t 'ft •ft •ft •ft f f 'ft f i t 126 f t •ft f t •ft f t f t •ft f t f t f t t t 'ft '#'t 'ft •ft f t 'ft f# 'ft f t f t f t 'ft •t't •ft 'ft f t 'f# 'ft 'ft 'ft •ft 't't 'ft f t f t •ft f t 'ft 'ft 'ft •ft ' f f •ft •ft f t •ft 'ft 'ft 'ft 'ft f t f t f t • 'ft • I f f t 'H •rt •t't f t f • f t f t •t't f t f# f t •t't •ft f t f t f t 't*t 't't tff »ft f t f t f t f t f t f t •ft f t f t f t f t t f 't't 'ft f t f t •t't f t f t f t t t 'ft 131 f t 'ft f f f t - -I-s •ft ' f t f t f t f t f t 'ft f t f t f t f t f t f t f t 'ft ' f f t t f t 'V t • f t 't't 't't f t •ft 'ft f t f t f t f t f t f t f f f t f t f t f t f t f t f t f f f t f t 136 •ft 'ft •t't ' t f f t f f f t f t f t f t 't'f f t f t ' t f f f 'ft f t •t't f t f f 't't ' f t 't't ' t f 't't 'ft ' f t f f ' f t f f f t ' f t f f ' f t ' f t 'ft ' f t ' f t ' t f f t 'ft ' t •t't ' f t f t 'WW • f f 132 f t 'ft 'ft f t ' t f f t t t V K f f ' f t t ' t f t H 't't f t f t f t t f '9*9 ' f t f t f t 'ftp • f t ' f t f f ' f f f t f t ' f t t t ' f t f t f t f t ' f f t ' f f f f f f t f f f f f t f 0 f t f t f t f « f f f t f t t f * t t i f I i f f 'ft ' f f i f | f f f t f t •ft f t 'ft f f i f | f t 'ft ' f f 'f| f t 't't ' f f • f t f t f t f § i f I f f | f « r i ' f t ' f t f t ' f t I f I i f f f | y | • f | ' f t f t ' f t f t f t f t t ' t rt t t t t 133 I 34 83 'ft | t f t f t f t 'ft ' f f f t f t f t 'ft f f f t f t 't't 'ft f f f t 'ft 't't f t f t f t f t ' f t f t f f f t f f t ' t f t ' t f t't f t f f f t t f t t f t f f f t f t f t t f f f '.ft f t f t f t f t Illus, 7 The variation pattern in cone scale morphology along a longitudinal transect from coastal Sitka spruce to montane white spruce forest. See f i g . 3 3 for line of shape of samples 1 3 1 to 1 3 5 and 8 3 to 81* inclusive. TABLE 27 MEANS AND GEOGRAPHIC ORIGIN OF FOLIAGE SAMPLES FROM SIXTY TREES 105 Sample T r e e L a t . L o n g . E l e v . In L<mra) T(imn) L / T S i So S i / S o llO )to 1000 f t . ' l c 17 .0 2 U . 3 127 3 5U.13 130.16 0.1, 15.3 b 13 m i l e s SE o f - P r i n c e Rupert 13.7 5 12 .9 0.62 20.6 18. U 1 .0 18.1.0 0 1 0.72 19.s 13.7 1.0 13.70 G 1 0.57 , 26.U 12.7 1 .0 12.70 0 1 0.75 18.3 U . 3 1.0 111.30 0 0 0 . 6 9 18.7 1 2 . 0 1.7 7.05 a 0 i c 19.5 0.82 23 .8 12.0 1.3 9.23 0 0 2" 11.9 0.73 I 6 . 3 1U.S 2.5 5 .92 0 0 120 3° 55.3 129.30 0.5 19.8 0.91 21.8 18.2 U.8 3.79 G 0 W 61a m i l e a 1U o f T e r r a c e 111.7 0.79 18.6 u . 6 1.U 10 .U3 G 0 5° 10 . l i 0.77 13.Ii 16.1 3.1 5.19 0 0 1 21.3 1 .0U 20.5 5 . 0 5 . 0 1 . 0 0 G 1 2 17.7 1 .10 1 6 . 0 6 . U U.7 1 .36 G 0 3 57.55 131 .11 1 .3 17. l i 1.01 17.U 6 . 2 U . 8 1 . 2 9 G 1 U c 250 m i l e a SW o f Watson Lake 17 .8 1.31 1 3 . 6 7.3 5 . 8 1 .26 P 1 5 1 8 . 9 1.12 16 .9 6 . 6 5 . 1 1 .29 0 1 1= 111. 2 0.91 15.6 12.9 5.2 2.U8 a 0 2° 15 .U 0.99 15.5 10.U 3.7 2.81 G 0 122 3C 55.28 128.U7 I.U 1U.7 1.00 U . 7 13.6 5.7 2.38 c 0 U<= 10U m i l e s U of T e r r a c e 17.0 1.00 17.0 12.1 U.U 2.75 G 0 5 C 13.1 1.05 12.5 U . 9 6.0 2.U8 G 0 1 17.0 1.26 13.5 5.7 5.3 1.07 P 0 2 12.3 0.36 U . 2 5.7 3.3 1.70 0 0 15U 3 5U.13 122.36 2.3 16.1 1.09 16.6 5.6 U.O 1.U0 P 1 h 35 m i l e a t;. o f P r i n c e George 15.2 0.97 1S.5 • 6.7 ' U.7 1.U2 P 1 5 9.9 1.06 9.3 9.8 6.1 1.61 P 2 1<: 12.1 1.00 12.1 5.1 5.2 0.98 G 0 2<= 15.6 1.05 35.2 7.2 5.1 1.U1 G 0 157 3C 5U.U5 122.52 2.U 12.5 0.98 12.7 U.2 U.6 0.91 P 0 U<= U m i l e s W. o f P r i n c e George 16.8 0.92 16.3 U.U U.2 1.05 P 0 5= 16.9 1.02 16.7 5.3 U.O 1.32 P 0 l c 12.0 1.21 10.0 8.7 6.7 1.30 G 1 2 16.5 1.1U U . 5 6.1 6.U 0.95 P 0 138 3C 59.U8 129.08 2.9 7.2 0.97 7.U 8.8 6.5 1.35 P 2 U U5 m i l e s SW o f '.Vatson L a k e . 16.2 1.31 12.3 . 7.S 7.6 1.02. G 0 5= 17.6 1.09 I 6 . 3 6.9 7.0 0.9a a 0 l c 7.2 0.90 6.0 7.7 5.2 1.U8 p 2 2 8.8 1.11 8.0 6.9 6.U 1.08 G 0 U U 3 56.UU 12U.55 3.3 9.0 1.07 8.5 8.5 6.U 1.33 0 1 U K i l e s U02 A l a s k a Highway 8.7 0.98 8.9 7.0 5.1 1.37 G 0 5 10.8 0.99 11.0 7.7 5.1 1.51 G 1 l c 10.9 0.91 11.9 7.2 U.9 1.U7 P 2 2C 13.6 1.08 12.5 6.8 5.2 1.30 G 0 LU7 3C 57.17 122.U5 3.9 13.9 1.18 11.8 9.1 6.5 1.U0 P 2 U c m i l e s 166 A l a s k a Highway 11.U 1.20 9.5 10.0 5.5 1.81 P 2 5° 13.9 o.95 111 .7 7.0 U.9 1.75 G 0 1° 21.9 1.27 17.2 7.9 U.7 1.68 P 1 2C 15.8 1.1U 13.8 7.3 5.3 1.38 P 0 155 3° 53.0U 121.31 U.2 17.3 1.18 U . 8 7.U 5.2 1.U2 P 0 U c 55 m i l e s E . o f Quesne l 19.2 1.15 16.7 7.0 5.6 1.25 P 0 5° 13.2 0.9" 13.7 7.6 5.3 1.U7 • P 1 1 U . 2 1.09 13.0 7.6 6.5 0.69 P 0 2 17.9 1.16 15.2 7.5 6.U 1.17 P 0 106 3 51.02 115.58 U.3 1U.7 1.19 12.3 6.5 6.0 1.08 P 0 U 39 m i l e s £ . of Radium 13.3 1.2L 10.7 6.U 6.9 1.21 P 0 5 17.1 1.31 13.0 7.5 5.U 1.36 P 0 1 17.1 1.10 13.1 7.e 5-5 l . U l P 0 2 20.1 1.3U l S . c 6.7 6.2 1.U0 p 0 3 50.59 119.33 U .7 17 .U 1.22 U . 3 6.2 6.2 1.00 p 0 U 55 m i l e s M i o f hamloops 20 . f I . U 1S.3 7.9 5.0 1.S6 p 0 5 22.2 1.26 17.6 7.U 5.0 1.U6 p 1 L — needle length, T - needle thickness, S^  - stomatic lines on dorsal surface, $2 - stcmatic lines on ventral surface, R - no. of resin canals in needles; V - vestiture, P - pubescent, G - glabrous, c - cone samples taken from the same tree. 106 DISCUSSION It is clear from the results that geographic variation in cone scale morphology is habitat-correlated. The assumption is warranted, therefore, that this variation is genetically based (Heslop-Harrison I964 p. 217), and that variation in cone scale morphology is the phenotypic expression of a pattern of physiological variation resulting from (1) adaptation to the varying environments occupied by the species (2) hybridization and the pro-duction of hybrid swarms which are also subjected to the environmental pressures similar in kind to those acting on the parental forms. It is not possible to determine the extent to which variation in cone scale morphology per se confers a survival advantage on spruce popu-lations. The characteristic shape of the scale of high and low elevation populations in the white-Engelmann spruce complex, for example, may have a direct effect in facilitating pollination at varying elevations\ for the characteristic shape of the scale is already present in young female strobili at the time of pollination, both in high and low elevation populations. More: likely, however, this variation in cone scale morphology is a "neutral" character, and therefore a selective value must be ascribed not to the character as such, "but to the influence of the genes responsible for i t upon the relative viability of a specific gene combination in a specific local environment" (Timofeeff-Ressovsky 19u0 p. 123). The results show that white spruce is sympatric in British Columbia, not only with Engelmann spruce, but also with black and Sitka spruce. There-fore in the interest of coherence and clarity the pattern of variation in cone scale morphology will be discussed in three separate sections under the following headings: (1) The pattern of variation in the white-Engelmann spruce complex. (2) The pattern of variation in the white-black spruee complex (3) The pattern of variation in the white-Sitka spruce complex 108 THE PATTERN OF VARIATION IN THE MIITE-ENGEIMANN SPRUCE COMPLEX The results obtained in this study support Taylor's conclusion that the phylogenetic relationship between P. glauca and P.rengelmanii in British Columbia "is best indicated by regarding them as subspecies of a single : species..." (Taylor 1°59). It seems clear that both forms hydridize freely, and that the hybrid is occupying an ecological niche intermediate betxiieen that of the parental forms. It is unlikely that the hybrid can successfully compete in the eco-logical niche occupied by either parent, for the parental forms will almost certainly be better adapted to their respective environments than the hybrid form. Therefore, for a hybrid to be successful an ecological niche must be available for colonization. If the ecological niche is intermediate between that of the parental forms, and is in the vicinity of the zone of hybridiza-tion then the hybrid will colonize i t . In British Columbia such an ecological niche appears to be available to the hybrid between white and Engelmann spruce. For between the low eleva-tion, allopatric white spruce populations of the montane forest, and the high elevation, allopatric Engelmann spruce populations of the subalpine forest there lies a broad transition zone which is available to the hybrid. Such a hypothesis will explain the clinal pattern of variation in cone scale morphology obvious in the white-Engelmann spruce complex. The selection pressures associated with altitude will, as a rule, vary pro-gressively, and consequently i t is to be expected that the transition from pure white spruce, through the hybrid swarms to pure Engelmann will be 109 progressive. This s i t u a t i o n i s c l e a r l y shown by the pattern of v a r i a t i o n along a single a l t i t u d i n a l transect at Stone Creek, south of Prince George, i l l u s t r a t e d i n F i g . 34. In the s e c t i o n dealing with the d i s t r i b u t i o n and phylogenetic r e l a -t i o n s h i p of spruce i n B r i t i s h Columbia, evidence was presented which i n d i c a t e d that white spruoe during late-Wisconsin time was much more widely d i s t r i b u t e d than i n h i s t o r i c time. Brink and Farstad (1949) have observed that i n northern and c e n t r a l B r i t i s h Columbia i t i s apparent that aspen f o r e s t i s advanoing i n t o grassland, and have suggested that one of the conifers which often succeeds aspen i s P. glauoa. These authors a l s o point out that there i s evidenoe that Douglas f i r has been replaced by spruce i n north c e n t r a l B r i t i s h Columbia. / White spruoe i s normally associated with the northern montane and boreal f o r e s t region i n B r i t i s h Columbia. The present study, however, i n d i -cates that white spruoe and i t s r e l a t e d forms occur i n a wide v a r i e t y of environments outside the northern montane f o r e s t , not of course as large f o r e s t s , but as small, fragmented, though not i s o l a t e d , populations. From the evidence of p o s t - g l a c i a l f o r e s t succession presented i n a previous sec-t i o n , and from the observations of Brink and Farstad (1949), i t can be assumed that these populations are c o l o n i z i n g populations, and not remnants of past d i s t r i b u t i o n s . For these reasons s p e c i a l i n t e r e s t i s attached t o those small, f r a g -mented populations, such as the B i r c h Island population which i s discussed below. Sample 117, l a t . 51°35», e l e v . 1450 f t . was made at B i r c h Island, which i s approximately 90 miles north of Kamloops. The sample area i s s i t u a t e d i n the v a l l e y of the North Thompson River, and therefore i s i n the southern s e c t i o n of the Columbia f o r e s t region (Rowe 1959). This same 110 area is also represented in the immature spruce populations grown at Cowichan Lake, and the Birch Island provenance was the last provenance of the white-Engelmann complex to enter dormancy in the f a l l of 1966. Illus. 5:2 shows the pattern of variation in this population, which the discriminant function analysis showed to be a sympatric population of white and Engelmann spruce (Fig. 1*6), and i t will be seen that in cone scale morphology the population has elements of both species. It is clear, however, for the following reasons that the Birch Island population (and populations with similar cone scale morphology) is not simply a f i r s t generation hybrid between white and Engelmann spruce. Firstly there are no large allopatric populations of white and Engelmann spruce at this elevation (11*00 ft.) in the immediate area. Secondly, i f the Birch Island population were a fi r s t generation hybrid then i t is to be expected that the seed collected from this population would be extremely variable. On the contrary, the population at Cowichan Lake showed very l i t t l e variation. Furthermore, the delayed dormancy, and related growth be-haviour of this provenance at Cowichan Lake can be explained in terms of the environment at its place of origin rather than in terras of heterosis resulting from hybridization. The conclusion is that the low elevation, Birch Island population, and similar populations in the interior of British Columbia are the product of introgressive hybridization followed by selection and adaptation of fractions of the resulting hybrid swarms. Introgressive hybridization results in the production of great variability, and consequently the colonizing potential of white spruce is greatly increased. For example, sample 63 (elev:* 3300 f t . , lat. 50°00', long. 120°36l), which the discriminant function analysis classifies as a sympatric population of white and Engelmann-spruce,-was taken from trees - - I l l growing i n a s s o c i a t i o n with ponderosa pine 13 miles south of M e r r i t t and w e l l outside the normally accepted e c o l o g i c a l zone of white spruce i n B r i t i s h Columbia. For these reasons i t i s suggested that 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 i s one other f a c t o r whioh may account f o r the expansion of spruce i n north c e n t r a l B r i t i s h Columbia r e f e r r e d to by Brink and Farstad (1949). From the point of view of p r a c t i c a l s i l v i c u l t u r a l and tree improve-ment, these low e l e v a t i o n scattered populations of spruce i n southern l a t i t u d e s have considerable i n t e r e s t , and w i l l be ref e r r e d t o i n another s e c t i o n (page 134). The white spruce populations i n the Rocky Mountain Trench do not f i t i n t o the general pattern of v a r i a t i o n outlined above. In the f i r s t instance they occur at muoh higher elevations than white spruce i n a l l other areas of the provinoe east of long. 127°00'. West of long. 127° 00' white spruoe a l s o occurs at high e l e v a t i o n s . However, t h i s region i s more appropriately discussed i n r e l a t i o n t o the pattern of v a r i a t i o n i n the white-Sitka complex. The discriminant function analysis i n d i c a t e s that The Rocky Mountain samples 104 t o 106 i n c l u s i v e and sample 108 are from sympatric populations of white and Engelmann spruce. Samples 104 to 107 are a l l from above 4000 f t . Sample 108 i s from 3850 f t . The f l o r a i s generally sparse, and d i s t i n c t from that common to t y p i c a l Engelmann spruce stands at 4000-5000 f t . elsewhere i n the Province ( I l l u s . 10). One of the most common shrub i n many of these areas i s Shepherd!a canadensis. I t i s possible that these white spruce populations at r e l a t i v e l y high elevations i n the Rocky Mountain Trench, e.g. sample 108, are a modern analogue of the late-Wisconsin f o r e s t described by Watts and Wright (1966), and r e f e r r e d t o i n the s e c t i o n dealing with d i s t r i b u t i o n and phylogeny ( I l l u s . 10). 112 The results of this study indicate that in regard to cone scale morphology in the white-Engelmann spruce complex the change from pure white spruce to pure Engelmann spruce is a progressive one. Furthermore, i t seems clear that the hybrid swarms, representing every degree of backcrossing, and intercrossing between hybrid forms, are extending the range of spruce be-yond the boundaries of the ecological zone generally associated with white spruce. THE PATTERN OF VARIATION IN THE WHITE-BLACK SPRUCE COMPLEX Fifteen areas were sampled in the boreal forest lying approximately between latitudes 56°00' and 60°Oo'. The cone scale morphology of indivi-dual trees in 10 of these areas is illustrated in Figs. 35 to ljl*. It will be seen that black spruce is a component of each sample except sample 11*0 (forest section Stikine plateau, lat. 5 7 ° 5 5 ' , elev. 1300 ft.) and 139 (forest section upper Liard, la t . 59°U8', elev. 1300 f t . ) . Sample 11*0 is identified as pure white spruce. Two trees of sample 139 are identified as white spruce, but the cone scale morphology of the remaining three trees of the sample corresponds neither to white or black spruce, or the inter-mediate form of these species (Fig. 3 6 ) . In this regard special interest is attached to the fact that this population (139) at Telegraph Creek exhibited a bark type unlike the bark either of white or black spruce. (Illus. 11+). Another characteristic which distinguished the Telegraph Creek po-pulation was the fact that male and female stroboli were present at the base of the crown, whereas the common position for reproductive organs in white spruce is in the uppermost part of the crown. However, there is no striking difference in needle morphology of this population and other populations of white spruce (Table 27)» Sample I I 4 I , which is also from the Stikine plateau section of the boreal forest appears to have a black spruce component (Fig. 3 8 ) . Samples 138, 1U2, 1U3 and lij5 are from the upper Liard section of the boreal forest. The population represented by sample 138 was examined in some detail in the fie l d . Trees numbered 1 and 3 were felled, and foliage was collected from trees numbered 1, 3 and 5. It was not possible to distinguish clearly black and white spruce by field observations in these areas. Tree number 1, for example, had foliage which in general appearance looked like black spruce though its cones had the appearance of white spruce. The needles of this tree were 12 mm. long, which is closer to black than white spruce (Table 27). The cone scale morphology of tree number 3 is apparently intermediate between black and white spruce (Fig. 35)• Its needles are strikingly short, 7.2 mm in relation to the length of the needles of the other k trees sampled, and its twigs are pubescent. Both of these are black spruce characteristics, yet the line of shape clearly shows characteristics of both white and black spruce (Fig. 35* tree no. 3). Taylor (1959) has shown that foliage characteristics are not corre-lated with cone type in intermediate populations of white and Engelmann spruce, and suggests in explanation that both characteristics are probably due to the independent segregation of several genes. The lack of strong corre-lation between cone scale morphology and needle morphology illustrated by trees number 1 and 3 of sample 138 (Fig. 35) suggests that Taylor's ex-planation may also be extended to intermediate forms of black and white spruce in the boreal forests of northern British Columbia. Sample llj2 (Fig. 39) was made 30 miles south of Watson Lake near the Hyland River elev. 2075. Trees number h and 5 of this sample are inter-mediate between white and black spruce in cone scale morphology. Trees number 1, 2 and 3 are clearly white spruce. Sample 1U3 (Fig. i;0), elev. 1725 f t . was made 1U3 miles south of Watson Lake. Trees numbered 1 and 3 are classified as black spruce, and 2, k and 5 as white. Sample ll& was made at mile 281 on the Alaska highway, elev. 1650 115 f t . Tree number 1 is classified as white spruce, and trees 2, 3, U and 5 as black (Fig. I|2). Samples lUU, Uj.6, and 1U7 were made in the northern foothills sec-tion of the boreal forest (Illus. 11). Foliage samples were taken in sample area lUU, but except for tree number 1, the foliage was taken from trees other than those which provided the cones. It will be seen that, in general, needle morphology for the entire sample is more characteristic of black spruce than white. Tree number 1 is identified as black spruce, number 3 as intermediate, and numbers 2, U and $ as white (Fig. 1(1). See also Illus. lit which shows variation in bark type in this area. The line of shape for each of the five trees in sample 1U6, mile 2kh of the Alaska highway, elev. 1760 f t . , is relatively clear cut. Tree number 1 is classified as an intermediate, numbers 2, U and 5 as black, and number 3 as white (Fig. 4 3 ) . Foliage and cones were a l l taken from the same trees in sample I 4 7 , mile 168 of the Alaska highway, elev. 3900 f t . Again i t will be seen that there is no apparent correlation between foliage characteristics, and cone scale morphology (Table 27* Fig. 4 4 ) . Trees 1 and 3 are classified, accord-ing to line of shape, as black spruce, mean needle length is respectively 10.9 mm. and 13.9 mm. Trees number 2 and $ are classified according to line of shape as white, mean needle length is respectively 13.6 and 13.9 ram* Tree number 4 is classified, as intermediate, mean needle length is 11.U mm. Trees number 1, 3 and U are pubescent, whereas 2 and 5 are glabrous. (Table 27, Fig. UU). Sample 1U8 is not graphed for line of shape but from Fig. hr6 i t will be seen that i t is classified as an allopatric zone of white spruce. This sample was made at mile 103 of the Alaska highway, elev. 3200 f t . , and field observations indicated that the ecology of the population sampled 116 was quite different from a l l other areas to the north, and that the area is located in the mixedwood section of the boreal forest region. The deep moss covering (see Illus. 11) typical of a l l sampled areas to the north, except sample area 139, is absent. Sample area 11*9, elev. 3000 f t . , is also in the mixedwood section of the boreal forest, and the discriminant function analysis, as in the case of sample lJj.8, indicates that i t is from an allopatric population of white spruce. This area lies approximately 18 miles north of Port St. John. The ground flora in this area is similar to that of sampled area ll|8, and is quite distinct from a l l other northern samples, in that the deep covering of moss and Ledum is absent. The ecology of samples to the south of sampled area 11*9, except sample 15k, which will b e discussed separately, are in general typical of the spruce forests of the montane forest region at 2000 f t . in the general re-gion of Prince George, lat. ca. 5u°00'. It is clear from the results of this study that black spruce is a much more important component of the spruce forests along the Alaska high-way than has been hitherto acknowledged (Garman 1957)• Of the 10 areas sampled north of latitude 5700o', only one, sample area 139, at Telegraph Creek in the Stikine canyon, did not exhibit a black spruce influence. North of lat. 57°00' on the Alaska highway, black and white spruce appear to occupy the same ecological niche. South of this latitude (it is emphasized that lat. 57°00' is an arbitrary line for the change is transi-tional), there is a gradual change in the composition of the spruce popu-lations, and black spruce moves to bog areas, and is not generally found in association with white spruce (e.g. samples li+8 and lli9). Occasionally* however, white spruce will be found in association with black in poorly 117 drained areas south of lat. 57°00', e.g. sampled area 15>1|« It will be observed both from the results of the discriminant function analysis, and the results obtained by line of shape method, that Engelmann spruce is absent throughout the sampled areas north of latitude 56°00*. As will be seen from the results of the discriminant function analy-sis (Fig. 46) a l l three spruce species, white, black and Engelmann are re-presented in sample l$k* Sample 154 was made on the periphery of a bog near Summit Lake, 35 miles north of Prince George. Foliage samples were taken in this area, and i t will be observed that the foliage samples of k of the 5 trees sampled exhibited dense pubescence (Table 27)• .This bog, and its associated flora, represent a fairly typical eco-logical niche where black and white spruce may be found in association south of lat. 57°00'. However, in contrast to the situation north of latitude 57°00', such an association is atypical for white spruce in the montane forest region, where i t is usually found in pure stands, and where black spruce is relegated to areas of poor drainage. On the other hand, in the boreal forest region along the Alaska highway north of lat. 57°00', white spruce is typically associated with black spruce on muskeg over a very large area as indicated by samples 138 and 11(0 to llj.6. Consequently i t is not exceptional that intermediate forms of black and white spruce are represented in these samples. 1 118 THE PATTERN OF VARIATION IN THE WHITE-SITKA SPRUCE COMPLEX In the study of variation in immature spruce population at the Cowichan Lake nursery on Vancouver Island i t was observed that provenances from the general region of the Nass, Skeena and Bulkley river basins entered dormancy later than provenances from further east but at similar elevations. For example, a l l provenances from Doughty (lat. ca. 55 °00 ' , long. ca. 127°30', elev. 2000 ft.) north of Smithers, and from the region of Hazelton, entered dormancy much later than provenances from further east, e.g. Fort Babine (lat. ca. 55 ° 20 ' , long. ca. 126°28', elev. 2300 f t . ) . Furthermore, certain provenances from this same region, in regard to germination behaviour at lf> GC, behaved like Sitka spruce in that they did not germinate at a l l at this temperature, or showed very low germination values. The pattern of variation in mature populations in the same general region of the Nass, Skeena and Bulkley river basins, as determined by cone scale morphology, presents strong evidence as to why the progenies of spruce populations in this area behave as indicated above when germinated or grown in a uniform environment (Fig. 33 and Illus. 7). It seems clear that there is a broad sympatric zone of white and Sitka spruce in northwestern British Columbia, and that Sitka spruce genes appear to have penetrated along the valley bottoms as far east as Hazelton and Smithers, and south east into the valley of the Morice River. As will be seen from Fig. 33 and Illus. 7 there is every gradation in cone scale morphology from that of pure Sitka (sample 131) to that of pure white (sample 8U)5 though sample 81* s t i l l shows the influence of Sitka spruce. There i s , therefore, considerable evidence that introgressive hybridization 119 is occurring in this area. The variation pattern is quite striking, and obvious even in the f i e l d . For example, sample 132 was made in the Skeena valley 1*3 miles east of Terrace. The associated species in the sampled area are western red cedar and western hemlock. Field notes indicate that from west to east this was the f i r s t sample which showed a strong influence of white spruce. The cone scale morphology of tree number 5 of this sample is clearly intermediate between that of white and Sitka spruce (Illus. 7). It i s , perhaps, significant that i t was not possible to obtain more than 15 cones from this tree. Sample 133 was made 58 miles east of Terrace. The strong influence of Sitka spruce in this area is s t i l l quite obvious, and even by a visual examination of the photograph of the sample from this area i t is possible to identify intermediate forms of white and Sitka spruce. Fig. 33 and Illus. 7 show the pattern of variation in cone scale morphology in the Skeena and Bulkley river valleys only. It is important to note, however, that the pattern is similar in the Nass: river valley. A major complicating factor in the interpretation of the variation pattern in this general region is that intermediate forms of white and Sitka spruce are almost identical with pure Engelmann spruce in cone scale morpho-logy. This fact has emerged from the present study, and i s , perhaps, an ex-planation for the reported occurrence of Engelmann spruce at very low elevations, in areas of the Nass and Skeena rivers (Garman 1957)• Almost certainly i t is the reason why samples in this area have been classified as a sympatric population of Sitka, Engelmann and white spruce by the dis-criminant function analysis. The best way to discriminate between spruce species and their inter-mediate forms is by a combination of measurements. No single measurement by itself is sufficiently accurate. Nevertheless, i t is possible to distinguish between white, Engelmann and Sitka spruce, though not intermediate forms, by the measurement 11/12 x L3. The means of this measurement are graphed in Figs. 30 to 32 for both standard and miscellaneous samples. It will be seen that to the north and east of the general region of the Nass, Skeena and Bulkley valleys the cone scale morphology is that of pure white spruce. The samples to the immediate south are pure Sitka spruce. There are, then, in the vicinity of the sympatric zone extensive allopatric populations of white and Sitka spruce. From the results reported here, and from field notes, there is no indication that extensive allopatric populations of Engelmann spruce occur in the region under discussion. In fact at high elevations (above 1(000 ft.) to the north of the Skeena and Nass rivers the miscellaneous samples clearly indicate the presence of pure white spruce. To the writer's knowledge, therefore, the nearest extensive allopatric population of Engelmann spruce is several hundred miles to the east at 3500 f t . in the region south of the McGregor river and north of Sinclair Mills (lat. ca. 5u°00T, long. ca. ^ l 0 ! ^ ; 1 ) . The conclusion i s , therefore, that the variability in cone scale morphology in northwestern British Columbia is the result of introgressive hybridization between white and Sitka spruce in this region. Subsequent selection and adaptation gives a pattern of variation (Fig. 33, Illus. 7) which varies parallel with the progressive change in environment illustrated in Fig. 9. 121 THE RELATIONSHIP BETWEEN THE VARIATION PATTERN IN MATURE AND IMMATURE SPRUCE POPULATIONS IN BRITISH COLUMBIA As already stated the clinal pattern of variation in cone scale morphology in the white-Engelmann spruce complex is the phenotypic expres-sion of physiological adaptation to the varying environments occupied by these species and their intermediate forms. The variation pattern of cone scale morphology in the Nass and Skeena river basins indicates that there is a broad sympatric zone of white and Sitka spruce in this area, and that there is a clinal pattern of varia-tion in cone scale morphology along a longitudinal transect from coastal Sitka spruce forests to the interior montane white spruce forests. These results, therefore, and the results obtained in regard to the growth behaviour of the immature spruce populations at Cowichan Lake, are mutually corroborative. The results of both studies testify to the over-whelming influence of environmental pressures associated with altitude with respect to variation in the white-Engelmann complex. In regard to the pattern of variation in the white-Sitka complex both studies indicate the extent and effect of the penetration of Sitka spruce populations into popu-lations of white spruce in the montane forest region. Sampling in mature populations was more extensive than the sampling in immature populations; consequently, there is no corroborative evidence from the variation pattern in immature populations to support the results obtained in regard to the pattern of variation in cone scale morphology in the boreal forests of northern British Columbia. Obviously, i t would have been considerably better to have conducted the study of variation in cone scale morphology on cones which also supplied the seed for the study of variation in immature populations at Cowichan Lake. Had this been done i t is clear that a high correlation could be es-tablished between the cone scale morphology of the parents and the per-formance of the progeny in the nursery. However, even though cones and seed were obtained from different samples i t is obvious that seedlings from a seed sample obtained from cones with a morphology similar to that for high elevation spruce in Table 6 and Illus. 3 are likely to enter dormancy at a relatively early date in a coastal nursery. On the other hand seedlings from a sample with a cone scale morphology similar to that for low elevation spruce in Table 6 and Illus. 3 are likely to enter dormancy at a later date then the high elevation sample. These considerations do not, of course, apply to spruce populations from the boreal forest. One of the most recent, and most detailed taxonomic studies of natural populations of a coniferous tree species is Ruby's (1967) study of Scots pine. Six hundred and eighty-nine, cone seed and leaf specimens of Scots pine were collected from 39 stands in Europe and Asia. Nineteen variable characters of cone and seed morphology were measured. Compilation of the components of variance for eight of these characters showed that more than 95 "percent of the variance was attributable to between-region differences, and less than 3 percent to stands within regions. The particular significance of these results is that they could be compared with the results of an associated 122-origin provenance study of juvenile characters of Scots pine from the same regions but grown in a uniform environment in East Lansing, Michigan (Wright and Bull 1963). The regional groupings obtained by both studies were nearly identical, and i t was concluded that i t is possible to delimit a race or variety of Scots pine nearly as well by studying morphological variation 123 in parental specimens collected in natural stands in Europe as by growing their progenies in a uniform environment (Ruby 1967). Correlation between morphological and physiological characteristics are not unusual, and are the rule rather than the exception i f the morpholo-gical characteristics are associated with the reproductive organs. For example the seed of the coastal and interior forms of Douglas f i r (Fseudotsuga menziesii (Mirb) Franco) is distinct morphologically, and this difference is closely paralleled by a striking difference in germination be-haviour at certain temperatures (Allen 1961). As Diver (I9I4O1) has stated "morphological variation may only be a part of the whole field of variabili-ty, but there is l i t t l e variation in the rest of the field which is not correlated with some morphological change, however slight". The study of variation in spruce cone scale morphology referred to above is not a special case, and a number of similar investigations have yielded information which is of immediate value to the forester engaged in provenance research, e.g. Prichausser 1958; Parker 1963; Myers and Bormann 1963j Hall and Carr 1964. The principal value of studies of this nature is that they provide information concerning the broad pattern of variation in the wild, mature populations of tree species which have undergone selection and adaptation, and possibly (as in the case under discussion) hybridization in their natural habitat. A relatively large sample can be made and rapidly assessed, and in this way the scope of subsequent experimental work with immature popu-lations in controlled and partially controlled environments is narrowed. These tests in turn narrow the scope of the field tests which ultimately follow a l l preliminary assessments of variation. The correlation between physiological variation in immature popu-lations and morphological variation in mature populations obtained in this study supports the view that biometrial studies on field specimens of cones and foliage can provide valuable evidence concerning broad patterns of variation in the mature populations* 125 THE TAXONOMIC SIGNIFICANCE OF GEOGRAPHIC VARIATION IN TffilTE SPRUCE IN BRITISH COLUMBIA. The pure forms of white and Engelmann spruce in British Columbia are distinct taxonomically and occupy quite distinct ecological niches. As a result of introgressive hybridization, hoxrever, the intervening ecological zone is also occupied by hybrid swarms, with the result that white and Engel-mann spruce in British Columbia, as the present study indicates, are the extreme forms of a clinal pattern of variation which ranges from low elevation montane forest to high elevation subalpine forest. Therefore, though the tax-onomic relationship between white and Engelmann spruce in British Columbia is , somewhat analogous to that of Abies balsamea var balsamea and A. balsamea var phanerolepis described by Myers and Bormann (I963), there is l i t t l e doubt about the taxonomic validity of the subspecific status (Taylor 1959) of the extreme forms of the white-Engelmann cline. The question remains however, as to how the great arrey of variability along the cline can be classified taxonomically. Langlet (1962, I963) has discussed in some detail the whole question of nomenclature in regard to infraspecific variability in tree species. Langlet's main thesis is that a species distributed over a region with a continuously changing environment will exhibit a clinal pattern of variation, and that i t is futile and unnecessary "to attempt to construct special ter-minology in order to summarize, and at the same time discriminate between, the various patterns which may occur" (Langlet 1963). Langlet goes on to say that when dealing with facts and problems of ecological infraspecific adaptation and variability, as few terms as possible ought to be used. 126 The writer is in f u l l agreement with this view, and believes that the best way to identify a spruce population in British Columbia is to assign to the sample, whether i t is seed for future propagation, or foliage and cone specimens for an herbarium, the exact elevation, latitude and longitude at its place of origin. At a later date i f a name must be assigned to the sample then i t may be classified by referring to Table 6 and determining the line of shape of the cone scale morphology of the sample in the manner pre-sented here. If a small portion of cones of a l l seed lots, collected for reforest-ation purposes in sympatric zones of white and Engelmann spruce were retained i t would be possible to identify a l l such seed lots by the line of shape method, and classify them accordingly. In this way each provenance may be named simply as Picea glauca (Moench) Voss subsp. glauca; P. glauca (Moench) Voss subsp. engelmanii (Parry) Taylor; or P. glauca X P. engelmannii, de-pending on its classification (see Taylor 1 9 5 9 ) . The results of the present study strongly indicate that the simpli-fied nomenclature proposed by Taylor (1959) more accurately reflects the tax-onomic relationships of white and Engelmann spruce in British Columbia than that which existed prior to his recommendations. It is suggested, therefore, that this nomenclature be used in the registering of spruce seed in this province. Natural hybrids of white and Sitka spruce, and white and black spruce have already been identified and described. The white-Sitka hybrid was des-cribed from material collected on the Kenai Peninsula in Alaska. The white-black hybrid was described from material collected in Minnesota (Little 1953, Little and Pauley 1 9 5 8 ) . It appears, however, that the evidence presented in this study is the f i r s t indication of the possible occurrence of both hybrids in British Columbia, though, clearly, much more detailed taxonomic work in the sympatric zones demarcated for these species is required before i t will be possible to select such hybrids in British Columbia for tree improvement work. The varietal epithet albertiana should be discarded, for as Taylor (1959) has pointed out, and as indicated by the present study, i t refers to the white-Engelmann hybrid which is more accurately designated in the manner indicated above. There is no evidence from the present study which indicates that the Porsild variety of white spruce occurs extensively in northern British Columbia. It is possible that the small population at Telegraph Creek (sample area 139) is of this variety. If so, i t is obviously worthy of further investigation to determine its correct taxonomic status, and the value of the population in regard to tree improvement work. This observa-tion could be equally applicable to a l l the hybrid components of the spruce complex of British Columbia. Illus. 9 Subalpine Engelmann spruce forest. The upper photo gives a general view of sample area 103, 61* miles NE of Cranbrook, Lat. k9°kl\ long. U5°27'* elev. 1*625 f t . The lower photo illustrates the dense shrub understory common in subalpine Engelmann spruce forests. Sample area 112, 1*8 miles NE of Kamloops, Lat. 50°1*8 , long. U9°52 , elev. 1*800 f t . Illus. 10 The white-Engelmann complex in the Rocky Mountain Trench. The upper photo shows structures of stand at sample area 105. Lat. 50°U8', long. l l6 o 00' , elev. liOOO f t . and the lower photo sample area 108. Lat. £L°2l ' , long. U6 03lt', elev. 38£0 f t . Note the xeric appearance of the stands and the lack of dense ground vegetation. Contrast with i l l u s . 9« Illus. 11 The white-black spruce complex of the Alaska highway. Mile 1*02 Alaska highway, sample area liUi. Lat. 580UU'> Long. 12U°55', Elev. 3300 f t . Note the deep Moss layer. Illus. 12 Characteristic branching habit in high (1) and low (2) elevation spruce in B.C. 1 - 38 miles West of Needles, elevation U100 f t . 2 - 1*0 miles East of Prince George, elevation 2000 f t . H Variation in branching habit in spruce in B.C. 1-18 miles Northeast of Nelson; elevation 1*25 f t . 2 - U8 miles Northwest of Smithers; elevation 1200 f t . 3-30 miles North of Kamloops; elevation 3500 f t . h - 10U miles North of Terrace; elevation 1375 f t . Fig. Variation in spruce bark type in B.C. 2 3 Illus. lU Fig. l l t : l and li*:2 illustrate variation in bark type in a sympatric zone of white and black spruce on the Alaska highway in Northern B.C. in the general region of sampled area lUU, elev. 3300 f t . , lat. 58°aU'. Along the Alaska highway there is every gradation between these two extremes of bark type. Fig. 1H:3 illustrates the unusual bark type of the Telegraph Creek population (see f i g . 36), 14:1 black spruce, 14:2 white spruce. 13k THE SILVICULTURAL SIGNIFICANCE OF GEOGRAPHIC VARIATION IN WHITE. SPRUCE IN BRITISH COLUMBIA Turesson (1925), who proved the existence of maritime and inland, northern and southern, low and high altitude strains within species, was the fi r s t investigator to demonstrate that the greater the range of climates the species is able to occupy the greater the variability within the species. Furthermore, Turesson also demonstrated that climatic races of widely dif-ferent species showed parallel variation in regard to both morphological and physiological properties (Hiesey et a l . 191*2). The present study indicates that, in common with the numerous plant species investigated by Turesson, white spruce is adapted to a wide variety of environments, ranging from areas of coastal influence, e.g. the Nass, Skeena and Bulkley river basins, through the montane and Columbia forest to subalpine forests. The study also shows that the colonizing potential of white spruce has been increased by hybridization, and that this species and its related forms have penetrated a variety of ecological niches well outside the ecological zone normally associated with spruce. The general significance of photoperiodicity for the practice of forestry has already been discussed by Wiersma (1958) and will not be repeated here. It i s , perhaps, sufficient to note that a l l recommendations in regard to the silviculture and breeding of white spruce in British Columbia must take this factor into consideration. The ecotype concept of variation (Wright and Baldwin 1957) tends to oversimplify the complex pattern of variation normally found within a tree species. It is not intended to repeat here the lengthy debate as to what is an ecotype, and what is a cline (Langlet 1959). The writer be-lieves that the overwhelming evidence, including the evidence of the present study, indicates that the variation pattern in most widely distributed coniferous species is clinal and not ecotypic. Ecotypes in coniferous tree species are more frequently the artifacts of inadequate sampling or sta-t i s t i c a l procedures than a true expression of the variation pattern. However, i t is obvious that from the point of view of practical silviculture the ecotype concept is a useful one, and i t will be resorted to here in making recommendation, in regard to the silviculture of white spruce in British Columbia. It is emphasized, however, that when latitu-dinal and altitudinal boundaries are referred to, i t is not implied that these boundaries circumscribe an ecotype which, in its pattern of variation, is discrete from that of the populations immediately to the north or south in the case of latitude, or higher and lower in the case of elevation. White and Engelmann spruce in British Columbia are the extreme forms of a clinal pattern of variation associated with altitude. Therefore, there are no sharp zonal boundaries marking the region outside which a given population cannot be moved. Nevertheless, i f a population is dis-placed by 1000 f t . upwards i t is obvious that in general its growth rhythm will be out of phase with the more severe environment at its new position. It i t is displaced downwards, but retained at the same latitude its growth rhythm will also be out of phase with the environment at the lower elevation. But since in most instances this environment will be less harsh than at 1000 f t . higher no profound detrimental effect will follow as a result of a displacement downward. However, no silvicultural gain can be expected either• For example, the high elevation populations from approximately the same latitude as Cowichan Lake were unable to avail of the mild condition pre-136 vailing in the coastal environment, and very low elevation of the Cowichan Lake nursery. A l l these populations entered dormancy when temperatures were increasing. If, however, the displacement is downwards, and to the north, then i t is likely that a silvicultural gain will result from this transfer. There is no silvicultural gain obtained by propagating high elevation spruce pro-venances in a coastal nursery at southern latitudes in British Columbia as is the practice at present. On the contrary, such a practice will almost certainly increase the cost .• of planting stock because of the necessity of repeated transplanting to obtain seedlings of suitable size for outplanting. For these reasons i t is suggested that the propagation of high elevation spruce provenances in coastal nurseries at southern latitudes, either on Vancouver Island, or on the mainland of British Columbia should be discontinued. Such populations growing in coastal nurseries at southern latitudes enter dormancy early, and have a stunted "rosette" appearance resulting from decreased interhode length. On the other hand, these same populations are likely to produce seedlings with desirable silvicultural characteristics much more rapidly in a northern nursery, such, as Telkwa (lat. ca. 55°00') in west central British Columbia, which is approximately six degrees north of the coastal nurseries. Consequently, as will be seen from Fig. Iii, there is an appreciable gain in photoperiod during the growing season. Corroborative evidence for the conclusions stated in the above pa-ragraph is available from a study initiated by the writer in 1963 at the Telkwa nursery, and at the Duncan nursery on Vancouver Island, which is approximately the same latitude as the Cowichan Lake nursery, and six degrees south of the Telkwa nursery. Thirteen provenances of white and Engelmann 137 spruce were sown at both nurseries in the spring of 1963. Table 28 shows the percentage of seedlings at both nurseries which had entered dormancy by August 22, 1964 (Roche 1964a). It will be seen that a much greater percentage of seedlings of a l l provenances at the Duncan nursery had entered dormancy by the date given. TABLE 28 PERCENTAGE OF 2-YEAR-OLD SEEDLINGS OF 13 SPRUCE PROVENANCES DORMANT BY AUGUST 22 AT A SOUTHERN AND A NORTHERN NURSERY IN BRITISH COLUMBIA. Provenance Lat. Elev. (ft.) Percent Dormant Duncan Telkwa (ca.lat. l i9 o 00') (ca.lat. 55°00') 1 56°00' 2300 69 23 2 5 5 V 2000 52 15 3 54°20f 2000 54 16 4 54°io' 2000 37 17 5 54°07 ! 2300 42 18 6 5 3 V 1800 22 11 7 52°3o' 2200 34 12 8 S i V I4OO 1 0 9 5 l ° l 5 ' 3800 69 36 10 50°40' 1*000 55 18 11 3200 39 9 12 49°30' 4700 33 18 13 4 9 V 4700 63 23 138 High elevation southern populations cannot avail of a long growing season, but they can avail of increased day length. This is a major conclu-sion deduced from the performance of these populations of Cowichan Lake, and i t should be taken into consideration in a l l plans relating to the dis-placement and propagation of these populations, whether in nurseries or plantations. A l l spruce provenances south of latitude 5>3°00', and from below 2500 f t . may be expected to grow rapidly in coastal nurseries in southern latitudes. Indeed, some of these populations, i.e. the Birch Island popu-lation, will grow as rapidly as Sitka spruce in a coastal nursery. Most of these low elevations, southern latitude provenances are fragmented in their distribution, and l i e outside the main body of the species. Nevertheless, i t is clear that they have considerable silvicultural potential, and that they will represent an important component in any tree improvement program which may develop for white spruce in British Columbia* For these reasons, a special effort should be made to collect seed from these populations. This has already been done for the Birch Island popula-tion, and there is now ample seed of this population available for further experimentation. Spruce populations from below 2000 f t . between latitudes $ 3 0 0 0 ' and 55°00' produce seedlings with desirable silvicultural characteristics more rapidly in coastal southern nurseries than high elevation populations from southern latitudes. Nevertheless, even these populations will do less well, silviculturally speaking, in a coastal nursery at low latitudes than in a northern nursery. Indeed, the evidence of this study suggests that a nursery in the vicinity of Telkwa is likely to produce desirable spruce stock in a shorter period of time than coastal nurseries at southern latitudes. Assuming that the microenvironment at the nursery site is optimum, i t is possible that a nursery situated further east, but at the same latitude as Telkwa, is equally preferable for the propagation of high elevation provenances from southern latitudes. This remark, however, does not apply to the populations from low elevation and low latitudes already referred to. These conclusions appear to contradict the recommendations of Eis (1966) in regard to the propagation of white spruce seedlings in coastal nurseries. However, the contradiction is more apparent than real for the provenance effect was not investigated by Eis. However, i t is also clear from the results of this study that pro-venances from below 2f>00 f t . , and south of latitude $5°00% will respond extremely well to increased temperature irrespective of the day length (within limits). Therefore, these provenances can be grown on the coast with optimum silvicultural advantage i f temperatures are increased arti-f i c i a l l y , e.g. by the use of plastic greenhouse. Furthermore, these popu-lations do not require the stimulus of an a r t i f i c i a l s oil mix, and will respond vigorously to increased temperatures while growing on ordinary nursery soil (Illus. U ) . High elevation provenances on the other hand do not respond to in-creased temperatures to the same extent as low elevation provenances, and appear to require a long day length, such as that obtaining in a northern nursery. For this reason i f a nursery area is selected at a low elevation in southern latitudes in the interior of British Columbia i t is likely that high elevation spruce provenances will not thrive in such a nursery. Very few populations from north of latitude $$°QQ 1 and east of longitude 12U°00' are represented in the study of variation in immature spruce population at Cowichan Lake. Consequently i t is not possible to indicate, -with any accuracy, the effect expected as a result of a dis-placement to south of low elevation northern populations. Nevertheless, i t is to be expected that they will behave similarly to high- elevation po-pulations from southern latitudes. The results of the study of variation in mature populations show, however, that a black spruce component can be extected to occur in any extensive seed collections made to the north, and indicates the need for further genecological investigation in spruce popu-lations north of latitude 5 5 ° 0 0 ' . Frequently reforestation projects are'scheduled for areas for which no seed of local origin is available, and consequently seed from other regions is used instead. The results of this study indicate that a displa-cement of 1000 f t . upwards will almost certainly result in a pronounced de-trimental silvicultural effect in most instances. A displacement of 500 f t . is also likely to result in a measurable effect. In this regard, survival during the fi r s t 1 - 3 years is not a good measure of the ill-effects of the environment on a displaced population. A displaced population, which shows 1 0 0 $ survival during the fi r s t few years, could, nevertheless be quite i l l -adapted to its new environment. There are a number of reasons for this. In the f i r s t place the climatic pattern during a short number of years may be atypical, and not representative of the average environment. Secondly, the outplanting technique in relation to size of stock can have a drastic effect on survival. Survival is an important parameter three to four years after the establishment of the plantation. Though i n i t i a l survival may not be drastically affected by a dis-placement of seed upwards, growth will be affected. Consequently, i t is obviously unwise silviculturally to displace seed upwards more than 500 f t . anywhere in British Columbia. No serious detrimental effect may be expected U j l to result from a downwards displacement of seed, while a downwards and north-wards displacement will probably result in silvicultural gain. The question as to how far north high elevation southern populations and low elevation southern populations can be displaced without detrimental silvicultural effects can be determined in the long run only from the field test. In this regard i t is worth noting that certain populations of Norway spruce can be transferred 10 degrees northwards without any detrimental effect (Langlet 1963b). This is not meant to imply that spruce populations from southern latitudes in British Columbia can be moved similar distances. It is very likely however, that (other things being equal e.g. site index and moisture regime) a displacement of U-5 degrees north may be silvicultural-ly feasible for high elevation provenances from southern latitudes. On the other hand, i t is likely that a transfer north of 4 degrees will result in a detrimental silvicultural effect in southern populations from low elevations (Roche and Revel 1966). Not infrequently the question is asked by the practical forester concerning the extent to which the variation pattern in juvenile populations assessed in controlled environments (growth chambers) and semi-controlled environments (greenhouse and nurseries) adumbrates the variation pattern in the same populations at a later date. There is l i t t l e doubt that the optimum climatic requirements for growth and development in forest trees, as in other plant species (Cooper 1963 p. 392) vary at a l l stages of the l i f e cycle. Consequently the optimum climatic conditions for germination in a given species under natural conditions are not likely to be the same as those which are optimum for seedling growth. Similarly, the conditions best suited for seedling growth are not likely to be the same as those which result in maximum growth in a 50-year-old tree. However, natural selection in forest trees operates primarily at the seedling stage, and the growth rhythm of a population adapted to its particular climatic environment is determined at this stage. Therefore, i f 2-year-old seedlings of provenance A enter dormancy eight weeks before pro-venance B when both are grown in environment G, and i f they are both left in environment C, there is no reason to believe that this relationship between the two provenances, assuming that they both survive, will be reversed when the trees are 50 years old. Furthermore, i f this difference in time of entering dormancy shows a clear cut relationship with the environment at place of origin of each provenance, then, to a considerable, extent, i t is possible to predict the relative adaptability of both provenances A and B when outplanted in climatic zone D in the field test (Roche and Revel 1°66). It is important to note, however, that the characteristics measured in ju-venile populations must be those which are of adaptive significance, and are habitat-correlated. For example, the low elevation, Birch Island provenance, and similar provenances, did best, silviculturally speaking, at the Cowichan Lake nursery. This does not mean that, in another environment, the Birch Island provenance will always exhibit silvicultural characteristics superior to those of the other populations with which i t was grown at Cowichan Lake. On the contrary, i f the same provenances were transplanted to a nursery at 4OOO f t . in the interior of British Columbia, then i t is certain that the Birch Island pro-venance, and similar provenances, would do least well silviculturally, and that other provenances would do better. To say that this is a reversal of growth at a later date in the l i f e cycle of the Birch Island population is to misunderstand the principles of genecology. The vitally important point is that i f studies of variation in immature populations in controlled and partially- controlled; environments 1U3 are carried out on a sufficient large sample, and in sufficient detail, then the relative adaptability of any population in a different environment is to a considerable extent, predictable. In this regard i t is perhaps worth mentioning that the above sta-tement is not based on supposition, but has considerable experimental evidence to support i t (Went 1957, Hudson 1957, Evans 1963). To give but one example, Went (1957), by controlled environment studies, established the climatic requirements of the plant species (Veratum yiride. This plant had never been grown in cultivation, and field tests completely failed to give any indication of the factors controlling its growth. On the basis of growth chamber work i t was possible to suggest locations where i t might be grown, and when tried out in these places i t grew as predicted (see Fogg 1963, p. 21*3). Had 150 to 200 populations from a l l sections of the boreal forest of northern British Columbia been represented in the study of variation in immature populations carried out at Cowichan Lake, i t is certain that some criteria would be provided in regard to the silvicultural consequences of a displacement from south to north. For the variation pattern of these boreal populations in a southern nursery would reflect the environmental pressures operating in the diverse sections of the Northern Boreal Forest. Apart then from the study of variation in cone scale morphology reported here, there is no information concerning the genecology of the great spruce forests of northern British Columbia. The need for such in-formation may not be pressing at the present time, but with the increase in the pulpwood industry in British Columbia i t is certain that such information will be required in the near future. For these reasons, i t would be wise to anticipate increased logging and reforestation activities in these northern forests by initiatinggeneco-logical studies as soon as possible. As Haddock (l°6l) has pointed out in regard to Canadian spruce forests in general, and western Canada in parti-cular " i f planting is to be advocated as a widespread solution, even i f for only the best quality sites, much more must be learned about how to grow high quality (physiologically) and inexpensive nursery stock of the best possible genetic constitution". It is clear that the coastal influence penetrates well into the interior of British Columbia, and that in the Nass, Skeena and Bulkley river basins there are sympatric populations of white and Sitka spruce, and evidence of hybridization between these species in the sympatric zone. Assuming that there is hybridization between white and Sitka spruce in the areas stipulated, i t is not certain that the hybrid is of silvicul-, tural value. For example, Thaarup (1°1|5) reported that the; hybrid Sitka-white spruce has poor stem form, and warned against collecting seed in the zone of hybridization, and Mergen (1959) has pointed out that hybrid seed-lings do not possess any advantages over their parents in their native habitat. In any event, i t is likely that no silvicultural advantages can be gained by moving seed out of this zone into other parts of British Columbia, It is possible, however, that high elevation southern provenances, or low elevation northern provenances (to lat. 55°OOl) can be transferred into this area without serious detrimental effect. It is stressed that the general recommendations given above con-cerning the transfer of seed, and the propagation of seedlings in the nursery, are tentative. These are recommendations which will be modified as further knowledge accumulates concerning the genecology of white spruce in British Columbia. A major source of new information in this respect will be the fiel d trials following the study of variation in immature and mature spruce populations reported here. It is suggested that plantations of representative provenances should be established in six forest regions (Rowe 1959) according to the following general plan: Area Forest Region Approximate elev.(ft.) 1 Interior subalpine 1*500 2 Central Douglas f i r 3000 3 Southern Columbia 1500 k Interior subalpine (North) 2000 5 Montane transition 2000 6 Southern Pacific Coast Sea level to 500 All provenances below 1500 f t . , except numbers 25 and 62, should be planted only at areas 5 and 6 (see Table 3). Areas 1, 2 and 3 should be in the same general region between latitudes 5l°00' and 52O00l i f suitable sites can be found. In this general region these three plantations, es-pecially i f placed along a single mountain transect, will yield information concerning the effects of a gross altitudinal displacement which will allow the modification of the general recommendations already stated. Area k should be in the region of Germansen Landing at approximately latitude 56C)00,. This plantation will yield information concerning the effects of a gross latitudinal displacement. Area 5 should be situated in the region of Telkwa, latitude ca. 55c 001 longitude ca. 127°30'. Consequently, as indicated by the study in mature populations, this plantation will be in a transition zone between the northern pacific coast section and the montane transition section, and will yield information in regard to the effect of a gross displacement to the north 11*6 west, an area for which there is a chronic shortage of spruce seed. It has already been stated that a l l provenances below 1500 f t . except provenances 25 and 62, should be planted only at areas 5 and 6. These provenances represent Sitka spruce populations, both strictly coastal and inland forms, from a series of latitudes between latitude U9°00' and 56°00'. Therefore i f area 6 is selected on the south western tip of Vancouver Island much information will eventually be obtained concerning the effects of a latitudinal displacement, while the same populations at Telkwa will- yield information concerning the effects of c o n t i n e n t a l i t y on coastal populations. These same populations at Telkwa will therefore further clarify the pattern of variation in the sympatric zone of white and Sitka spruce. It will be seen from Table 3 that any one altitude (to the nearest 100 ft.) is represented by a number of .provenances. The results of the study reported here indicate that 5 provenances from each 100 f t . of altitude will adequately sample the'population at that altitude. I t is suggested, there-fore, that this criterion be applied in selecting provenances for outplanting. For some altitudes there are less than 5 provenances available, but for many altitudes there are more than 5. Where there is a choice i t is obvious that the five provenances selected at a given altitude should be selected to represent the widest area at that altitude. For example at 2000 f t . there are 16 provenances from which to select 5» Clearly i t is better to select only one from the region of Doughty near Smithers rather than select a l l 5 from this area which is well represented at this elevation (Table 3). If this procedure is followed, there will be a considerable re-duction in the number of provenances selected for outplanting with a con-sequent reduction in costs. The above remarks do not apply to provenances belox* 1500 f t . a l l of which should be outplanted as indicated above. 11*7 Each of the outplanting areas, except area 6, will be well re-presented by the provenances selected for outplanting. Consequently there will be a control available for comparison with alien populations. In regard to a tree improvement program, i t is suggested that the main task in the immediate future is to find and exploit by selection and breeding the great arrey of genetic variation in white spruce populations identified and demarcated by the present study. The X-ray method of assessing seed quality has considerable merit, and is obviously superior to methods in use at the present time. It is also clear that the environmental requirements for optimum germination vary for spruce seed from different elevations, and that an accurate assessment of germinability in the laboratory cannot be obtained by germinating high and low elevation provenances a l l at the same temperatures. The X-ray method of assessing germination capacity described here is accurate only when applied to seed undamaged physiologically as a result of handling prior to extraction, extraction methods, and conditions of storage. The low germination capacity of many provenances used in the pre-sent study cannot be fully explained in terms of embryo development or provenance effect, and i t seems clear that this seed has suffered damage during one or a l l of the processes referred to above. However, a comparison of figures for germination capacity obtained in this study with figures obtained as a result of routine germination tests following extraction, indicate that the condition of the seed prior to storage rather than storage conditions per se, is probably the principal cause of low germination ca-pacity of seed which has fully developed embryos, and shows no structural damage. There i s , nevertheless, a relationship between embryo development and germination behaviour even in seed which has a percentage of physiologi-148 o a l l y unsound seed. Consequently seed l o t s used i n research, p a r t i c u l a r l y research r e l a t e d t o d i r e c t seeding, should be assessed by radiography. Seed which has deteriorated p h y s i o l o g i c a l l y can a l s o be assessed by the X-ray method, but only i f the seed has been treated by s u i t a b l e contrast agents which vary wi£h species and whioh are absorbed d i f f e r e n t i a l l y by l i v e and dead t i s s u e , and consequently, present varying patterns of density i n the radiograph. The r e s u l t s of the study of germination behaviour reported here i n d i c a t e a considerable need for more d e t a i l e d studies of a l l f a c t o r s a f f e c t i n g germination behaviour i n white spruoe i n the manner of A l l e n ' s (1958) and S z i k l a i ' s (1965, 1966) comprehensive and d e t a i l e d work on Douglas f i r . GENERAL CONCLUSIONS 1. White and Engelmann spruce in British Columbia are the extreme forms of a clinal pattern of variation. 2. Throughout the Province south of lat. 55°00' and east of long. 123°00 intermediate forms of both species occupy the broad transitional zone be-tween the montane and subalpine forest regions. 3. The intermediate forms are found, not as large forests, but as small, fragmented, though not isolated populations. k» These populations are colonizing populations and not remnants of past distributions. They may be identified on the basis of cone scale morpho-logy. 5. The environmental factors, which exercise selection pressure on the white-Engelmann spruce complex, vary progressively from the montane to the subalpine forest region. 6. The clinal pattern of variation in the white-Engelmann complex re-ferred to in 1 above is the result of introgressive hybridization, followed by selection and adaptation of fractions of the hybrid swarm. 7. No populations of Engelmann spruce occur along the Hart and Alaska highways north of lat. 55o00'. 8. White spruce occurs throughout the Province from lat. U9°00! to the Yukon border, and west to long. 129°Oo' in the vicinity of the Nass and Skeena rivers, and long. 131°00' in the vicinity of the Stickine river. °. The pattern of variation in spruce populations in the transitional zone between coastal Sitka spruce forest and montane white spruce forest in the Skeena valley is clinal. 10. Intermediate forms of both species occur in the Skeena valley and may be recognized on the basis of cone scale morphology. 11. The intermediate forms of white and Sitka spruce are very similar to pure Engelmann spruce in regard to cone scale morphology. It is possible, therefore, that in the past these forms have been mistakenly identified as Engelmann spruce. 12. The environmental factors exercising selection pressure on the white-Sitka complex in northwest British Columbia vary progressively from coastal to montane forest. 13. The clinal pattern of variation referred to in 9?above is probably the result of introgressive hybridization between white and Sitka spruce followed by selection and adaptation of fractions of the hybrid swarm. Ik. It is probable that sympatric populations of Sitka and white spruce occur in other regions of the Province where coastal influences penetrate the valleys of the major rivers. 15. Sympatric populations of black and white spruce occur along the Alaska highway throughout northern British Columbia. 16. North of lat. j?7°00' on the Alaska highway, black and white spruce occupy the same ecological niche. South of this latitude there is a gra-dual change in the composition of the spruce populations, and black spruce moves to areas of poor drainage. 17. Individuals exhibiting characteristics of both black and white spruce occur in the sympatric zones referred to in 15 above. It is possible therefore that hybridization is occurring between these speoies. 18. These individuals ocour at random, and there is no evidence that intermediate forms:.of black and white spruce are successfully colonizing areas in the vacinity of the sympatric zones. 19. In regard to the white-Engelmann spruce complex in British Columbia there is sufficient evidence from this study to warrant the assumption that the environmental pressures which result in microevolution, i.e. geographic differentiation within the speoies, differ only in degree rather than in kind from the environmental pressures which result in macroevolution, i.e. speciation. 20. The faculty for normal development and survival of white spruce, and its related forms is conditional by the cessation of growth and initiation of dormancy. 21. Time of cessation of dormancy in a population in any one region where the speoies occurs naturally is conditioned by its genetic consti-tution. 22. The genetic constitution of a natural population is predominantly determined by the photothermal regime prevailing in that region. 23. In so far as there is a difference in the photothermal regime be-tween any two regions the genetic constitution of the populations occupying those regions wil l differ. 2u. One of the most important external manifestations of this differ-ence is the time of cessation of growth, and initiation of dormancy* 25. Time of cessation of growth and initiation of dormancy is a popu-lation parameter. Therefore, the dormancy curve will accurately character-ize a spruce provenance only i f assessed on the basis of a relatively large number of seedlings. 26. Between the time of the initiation of dormancy and it s completion in any one spruce provenance there is a period when the correlation between dormancy and factors of the environment at the place of origin is maximum for this provenance. Therefore, in order to determine this period a dor-mancy curve must be constructed. 27. On the basis of conclusions 19 to 23 inclusive recommendations may be made in regard to the silviculture of spruce populations from the interior of British Columbia. 28. High elevation spruce provenances should not be grown in coastal nurseries at southern latitudes on Vancouver Island or on the Mainland. Such provenances cannot avail of the long growing season at these nurseries, as they enter dormancy even when temperatures are increasing. The breaking of dormancy during the same growing season is the exception rather than the rule in these high elevation provenances. Consequently they have a stunted "rosette" appearance resulting from decreased internode length, and are inferior planting stock even after transplanting. 29. High elevation provenances cannot avail of a long growing season, but there is evidence that these provenances will respond to increased day length. Therefore, high elevation spruce provenances other things being equal e.g. conditions of so i l , general supervision, etc., are likely to produce seedlings xiiith desirable silvicultural characteristics much more rapidly in a northern nursery, such as Telkwa, which is approximately 6 degrees north of the coastal nurseries. 30. Assuming that the microenvironment at the nursery site is optimum, i t is possible that a nursery situated further east, but at the same l a t i -tude as Telkwa, is equally feasible for the propagation of high elevation provenances from southern latitudes. 31. A l l spruce provenances south of latitude $3°00x and from below 2^ 00 f t . may be expected to grow with relative rapidity in coastal nurseries in southern latitudes, and some will grow as rapidly as Sitka spruce in these nurseries. 32. These small, fragmented, populations, e.g. the Birch Island popu-lation, have considerable silvicultural value, and are of importance in tree improvement work. For these reasons, efforts should be made to collect seed from these populations. 33. Spruce provenances from below 2500 f t . , between latitudes 53°Oo' and 55° O o ' will produce seedlings with desirable silvicultural character-istics more rapidly in southern coastal nurseries than high elevation pro-venances from southern latitudes. These same low elevation provenances, however, are likely to do much better silviculturally speaking, in the Telkwa nursery also, where spring and summer temperatures are not much lower than those of the southern coastal nurseries, but where day length during the same season is considerably longer. 3k» Provenances from below 2500 f t . , and south of latitude 55°00' will respond extremely well to artificially increased temperature irrespect-ive of day length (within limits). Therefore, these provenances can be grown on the coast with optimum silvicultural advantage i f temperatures are increased artificially, e.g. by the use of plastic greenhouses. Further-more, these populations do not require the stimulus of a prepared s o i l , and will respond vigorously to increased temperatures while growing on regular nursery s o i l . High elevation provenances, on the other hand, do not so respond. 35• White and Engelmann spruce in British Columbia are the extreme forms of a clinal pattern of variation associated with altitude. Therefore, when latitudinal and altitudinal boundaries are referred to above, i t is not implied that these boundaries circumscribe an ecotype which, in its pattern of variation, is discrete from that of the populations immediately to the north or south in the case of latitude, or immediately above or below in the case of altitude. Therefore, there are no sharp zonal boundaries mark-ing the region outside which a given provenance cannot be moved. 36. Spruce provenances north of latitude 55°Oo' may contain a black spruce component. Such provenances from north of latitude 55°00!, whether from allopatric zones of white spruce or sympatric zones of white and black spruce, are likely to behave similarly to high elevation southern provenances in regard to time of entering dormancy in southern coastal nurseries. 37. Embryo development in spruce will vary with the seed crop year, date of harvesting, and place of origin within any one year. Germination behaviour is strongly influenced by embryo development. Consequently, care must be taken to ensure that seed lots used for experimental purposes, e.g. direct seeding tests, are standardized in regard to embryo development, 3 8 . An accurate, and quick method of assessing embryo development and seed quality in general is by radiography using the methods described in this study. 3 9 . In assessing population differences in germination behaviour, ex-treme temperatures are more effective than moderate temperatures. Maximum differences are observable at 15°C. 30°C. is also effective, but less so than 15°C. Least effective temperatures are 20 and 25°C. IjO, The temperature requirements for optimum germination are different for high and low elevation populations. I4I. Spruce provenances from sympatric zones of white and Sitka spruce in the Skeena, Bulkley and Nass river basins may be identified as to species i f germinated at 15°C At this temperature Sitka spruce will not germinate at a l l within lh days, or will show very low germination values. White spruce will germinate at this temperature. In making the test the germina-tion percent must be corrected on the basis of embryo development. Ij2, Reduced germination capacity of spruce seed during storage appears to result not from storage conditions per se but from the condition of the seed prior to storage. U3. In regard to a program of tree imporvement for white spruce i t is concluded that the main task in the immediate future is to find and exploit by selection and breeding the great array of genetic variation in white spruce identified and demarcated by the present study. 1*., Both the assessment of physiological differences in immature populations i n c o n t r o l l e d and p a r t i a l l y c o n t r o l l e d environments and the assessment of patterns of v a r i a t i o n i n mature populations i n the w i l d are e s s e n t i a l preliminary i n v e s t i g a t i o n s to the f i e l d t e s t , that i s , the pro-venance t r i a l . 45. When the problem of i n f r a s p e c i f i c v a r i a t i o n i n a tree species i s considered as a problem i n microevolution, rather than a purely s i l v i c u l -t u r a l problem, the emphasis i n research i s s h i f t e d t o the genecological i n v e s t i g a t i o n s r e f e r r e d t o i n 42 above. 46. Much greater emphasis, therefore, should be given both t o the assessment of p h y s i o l o g i c a l i n v e s t i g a t i o n s i n immature populations i n co n t r o l l e d and p a r t i a l l y c o n t r o l l e d environments, and the assessment of morphological v a r i a t i o n i n mature populations. 47. The f i e l d t e s t , that i s the provenance t r i a l , i s r i g h t l y considered the l a s t stage i n the assessment of v a r i a t i o n . Such t r i a l s then f a l l witlain the scope of the s i l v i c u l t u r i s t , and oan become part of an o v e r a l l p l a n t i n g program. The data concerning the performance of these plantations, which w i l l accumulate over the years, may be used t o modify the s i l v i c u l t u r a l re-commendations drawn up by the geneoologist on the basis of the in v e s t i g a t i o n s r e f e r r e d t o i n 44 above. 157 LITERATURE CITED Alexander, R.R., 1958. Silvical characteristics of Engelmann spruce. 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Fig. 10 The relationship between altitude and temperature regime for representative climatic stations in British Columbia. i i i 1 1 1 1 1 1 1 ' ' 1 Jo Jan Feb March April May June July Aug Sep Oct Nov Dec Note the general decrease in temperature with increasing altitude. 172 Fig. 11 The relationship between photoperiod and temperature at the Cowichan nursery and at Vaveriby in south central British Columbia. 20 9 0 - 7 0 - 6 0 o 10 Temp. Photoperiod Lai . Elev. (ft.) Period of Record (yrs.) co. 49 0 0 5 7 0 I ( 1966) 52 0 0 1465 3 0 90 8 0 7 0 6 0 -5 0 n -19 3 o 4 0 ^ H 3 0 - 10 20 I 31 I 31 30 30 2 9 2 9 2 8 2 7 2 7 2 6 2 6 Jan Feb March April May June July Aug Sep Oct Nov Dec 23 2 4 Note the striking similarity of the environments as represented by the Vaveriby station and the environment at Cowichan in respect to the day length on the f i r s t day of the year showing a temp, of U3°C. The Birch Island provenance is represented by the Vavenby station. Note, that i t is much slower to enter dormancy at Cowichan than a provenance from approximately the same lat. and long, but higher elevation, and that i t produced maximum shoot extension in the Cowichan nursery. Points A and C on the curves represent the daylength at each station when temperatures reach k3°C. 173 Fig. 12 The relationship between temperature and photoperiod at the Cowichan Nursery and at Babine Lake in Central British Columbia. 2 Or 100 Temp. Photoperiod Lot. Elev.(ft.) Period of Record (yrs.) co. 49 00 570 ' 55 15 '2360 ( 1966) Provenonce L o t Elev (fl.) | 128 HAZELTON 55 18 2.2 14 BABINE LAKE 55 20 2.3 90 80 70 60 -o CD o i o 4 0 a H 30 - 21 20 - 22 23 J 0 J 2 4 20 31 I 31 30 30 29 29 28 27 27 26 26 March April Moy June July Aug Sep Oct Nov Dec 1 Note the disparity between the two regions in this index of the environment and compare with the Terrace station. Note that the Hazelton provenance, which is from the general region of Terrace, was much slower to enter dormancy than the Babine Lake provenance which is further to the east, and in a region of greater continentality. (See f i g . lU for Terrace data)'* 171* Fig. 13 The relationship between temperature and photoperiod at the Cowichan nursery and at Barberville in east central British Columbia. 20 13-100 Photoperiod Lot. Elev. (ft.) CO. 49 0 0 5 70 5 3 0 0 41 80 Period of Record (yrs.) I (1966) 30 - 5 Provenance Lot Elev (ft.) 10 SODA CREEK 52 20 2 2 0 0 42 NAVER 53 20 3 4 0 0 31 I 31 30 30 29 29 2 8 2 7 2 7 2 6 2 6 Jan Feb March April Moy June July Aug Sep Oct Nov Dec 100 9 0 6 0 -3 o 4 0 a A 10 22 23 0 J 2 4 Note the disparity between the two environments and compare with the curves for Prince George and Aleza-Lake. Note also that the Soda Creek provenance, which is from a relatively low lat. and elev., was much slower to enter dormancy than the Naver provenance. In regard to dormancy behaviour the Soda Creek provenances is comparable to the Birch Island provenance and other provenances from low elevations and-low latitudes. 175 Fig. Hi The relationship between temperature and photoperiod at the Cowichan Nursery and at Terrace in Northwestern British Columbia. 20 S 10 2 -31 I 31 30 30 29 29 28 27 27 26 26 Jan Feb March April May June July Aug Sep Oct Nov Dec • | Note that on the basis of this index of the environment there is not a great disparity during the growing season between the environment at Cowichan and the environments represented by the Terrace station. The significant difference is not in temperatures during the growing season, but in winter temperatures• 176 Fig. 1$ The relationship between temperature and photoperiod at the Cowichan nursery and at Aleza Lake in east central British Columbia. Note the similarity of the Aleza Lake curves with those of Prince George, and contrast with those of Babine Lake and Barkerville. Fig. 16 The relationship between temperature and photoperiod at the Cowichan nursery and at Prince George in central British Columbia. Note the similarity of the curves with those of Aleza Lake, and contrast with those of Babine Lake, Barkerville, Allison Pass, and Vavenby. 178 Fig. 17 The relationship between temperature and photoperiod at the Cowichan nursery and at Allison Pass in southern British Columbia. 31 I 31 30 30 2 9 2 9 2 8 2 7 2 7 2 6 2 6 Jan Feb March April May June July Aug Sep Oct Nov Dec Note that on the basis of this index of the environment there is a considerable disparity between the environment at Cowichan and the environments represented by the weather station at Allison Pass. Compare with the curves for Old Glory mountain and contrast with Vavenby. 179 Fig, 18 The relationship between temperature and photoperiod at the Cowichan nursery and on Old Glory Mountain in southern British Columbia. 20 3 10 100 9 5 9 0 8 5 7 0 65 6 0 - S 5 0 a. e 4 5 - 4 0 3 0 2 5 4- 20 15 5 -Temp. Photoperiod Lot. Elev. (ft.) Period of Record (yrs) ca. COWICHAN 49 0 0 570 I (1966) OLD GLORY MT. 4 9 0 0 7 7 0 0 15 -6 I 31 I 31 3 0 30 2 9 2 9 2 8 2 7 2 7 2 6 2 6 Jan Feb March April May June July Aug Sep Oct Nov Dec 2 r-23 24 Note the great disparity between the environment at this elevation, which is at approximately the same latitude as the Cowichan nursery, and the environ-ment at Cowichan as measured, by day length on the fi r s t day of the year with a temperature of 1*3°C. Approximately 33 percent of the provenances grown at Cowichan are from above 1*000 f t . P E R C E N T F L U S H E D APRIL 6 LAT ALT DAYS = .148 -.140 .096 APRIL 16 LAT ALT DAYS = .400 -.395 .28 1 APRIL 20 LAT ALT DAYS = .329 -.424 .36 1 P E R C E N T D O R M A N T JULY 14 LAT ALT DAYS -.561 .862 -.796 JULY 21 LAT ALT DAYS -.430 .827 -.828 JULY 28 LAT ALT DAYS -.13 1 .699 -.834 APRIL 27 AUGUST 4 LAT ALT DAYS .099 -.26 1 .280 LAT -.089 ALT .656 DAYS -.803 Degree of correlation between factors of the environment and flushing and dormancy, r-correlation coefficient, LAT.-latitude, ALT.-altitude, DAYS-index of the vegetative period (See tables 2, 19, 21). Fig. 20 Curves of flushing for Sitka (1), white ( 2 ) , sympatric popula-tions of white-Sitka, and white-Engelmann (3), and Engelmann spruce ( 4 ) . DAYS FROM APRIL I Note the striking similarity of the curves for diverse provenances which vary in altitude from sea level to 47OO f t . P-provenance; E-elevation; L-latitude. 182 Fig. 21 Curves of dormancy for Sitka spruce ( 1 and 2) and provenances from sympatric zones of white and Sitka spruce (3 and !*)• DAYS FROM JUNE 25 Note that| a l l provenances in k except 62 are from the region of Hazelton in N.W. British Columbia. 62 is the Birch Island provenance and is. from a sympatricj zone of white and Engelmann spruce (see part B re sympatric spruce populations). Note also the similarity of the curves in U to the pure Sitka spruce populations, and compare with curves 5> and 6 in f i g . 22. I i I Fig. 22 Curves of dormancy for provenances from sympatric populations of white and Sitka spruce, and low elevation allopatric spruce pro-venances. Note the great sensitivity of the dormancy curve in distinguishing provenances. Z < £ OL O O L. 53 25 55 36 55 36 54 54 UJ O cn UJ Q. DAYS FROM JUNE 25 Provenance 51 in 5 is from Hixon south of Prince George in central British Columbia. 76, 77 and 127 are from N.W. British Columbia in the region of the Kispiox river. 25 is from the Birch Island region. In 6, 106 and 110 are from N.E. British Columbia: 13, 121 and 122 are from the Hazelton-Smithers region in N.W. British Columbia. Fig. 2 3 Curves of dormancy for spruce provenances from elevations between 2 0 0 0 and 2 3 0 0 f t . DAYS FROM JUNE 25 | ! Note that a l l these provenances except 8 in 9 and 1 2 8 in 1 1 have similar curves. Both of these provenences are from the region of Hazelton. Even at relatively high elevations, therefore, provenances from N.W. British Columbia have a characteristic dormancy curve which distinguishes them from provenances of the same elevation but further east. 18$ Fig. 2k Curves of dormancy for spruce provenances from elevations between 2300 and 2700 f t . DAYS FROM JUNE 2 5 -Note that the 5 provenances in 16 represent areas both in N.W. and N.S. British Columbia, hence the strikingly different curves for provenances from the same elevation. 186 "Note the gradual change in the shape of the dormancy curves, which at l|O00 f t . , show the rapidity with which provenances from high elevations enter dormancy in comparison with low elevation provenances from central British Columbia, and provenances from N.W. British Columbia. 167 Fig. 26 Curves of dormancy for spruce provenances from elevations between ipOO and 1(500 f t . DAYS FROM JUNE 2 5 Note that i n regard to elevation provenances 2k and 26 in 23 are of doubtful origin. Following observations in the nursery i t was confirmed by corres-pondence that both provenances were incorrectly registered. It is clear from the dormancy curves that these provenances are from low elevations, and are identical in growth rhythm to the Birch Island provenance. DAYS FROM JUNE 25 Note the change in form of the curves in comparison to those for pro-venances from elevations between 1500 and 2500 f t . 189 RELATIONSHIP BETWEEN ELEVATION AND SPRUCE CONE SCALE-SEED WING RATIO I 50T 14 5-ELEVATION IN THOUSANDS OF FEET "Fig. 28 Each point on the curve represents the mean of 100 cones. 1963 collection in the white-Engelmann spruce complex. 1.55^ 29 Relationship between elevation and spruce cone scale - seed wing ratio. Each point on the curve represents the mean of 100 cones. 196k collection in the white-Engelman spruce compl TJ./L2 - cone scale seed wing ratio. Fig» 30 Pattern of variation in a single measurement of cone scale morphology. Each bar represents the mean of 100 cones. 1963 collection. Fig 32 Pattern of variation in a single measurement of cone scale morphology. Each bar represents the mean of a varying number of cones. Miscellaneous collections. Fig. 33 The pattern of variation in spruce cone scale morphology along a longitudinal transect from coastal Sitka spruce forest to montane white spruce forest. Note the increase and decrease in CV (Coefficient of variation) as the transect crosses the sympatric zone. CV 10.0 11.3 12.0 12.6 22.2 15.6 15.1 2 5 0 7 10 1.9 20 0,7 1.0 1.5 ZD 0.9 10 1.5 0.5 _ J l 1 » I I 1 1 *• I 1 The numbers 1 to 10 on the vertical bars at the extreme left and right represent the 10 cone scale measurements illustrated in f i g . 1. In the top series of curves the mean for each longitude is plotted against pure white spruce in the bottom series against pure Sitka. The figures on the horizontal bars? represent the number of times each mean of the sample deviates positively or negatively from the corresponding measurement of pure:.white spruce (top series) and pure Sitka (bottom series). See page 41 for further explanation. CV refers to measurement 10, Ll/L2: x L3. Fig. 3k The pattern of variation in spruce cone scale morphology along an altitudinal transect at Stone Creek, South of Prince George (lat. ca. 5U°00'). C V 5 . 6 4.1 4 . 5 5.1 6 . 0 7.1 6 . 4 1 0 l . « 0.6 1.0 1.4 0.6 1.0 1.4 0.6 1.0 1.4 0.6 1,0 1.3 0 7 1.0 13 0 7 H • > 1 1 i 1 1 • 1 i i 1 p i i i i_ i f 1 i 1 1 i 1 i i r - • • -i i 1 1 i 1 1 i I i 0.6 1.0 1.4 0.6 1.0 1.4 0.6 1.0 1.4 0.6 1.0 1.4 0.6 1.0 1.3 0.7 1.0 1.5 0.7 1.0 1.8 In the top series of curves the mean for each altitude is plotted against pure white spruce, in the bottom series against pure Engelmann. The figures on the horizontal bars represent the number of times each mean of the sample deviates positively or negatively from the corresponding measurement for pure white spruce (top series) and pure Engelmann spruce (bottom series). Means are calculated from a 100-cone sample at each altitude ( see page41 for further explanation). CV refers to measurement 7, L1/L2. TN I 2 3 4 5 C W B I W B i 1 1 i 1 1 i 1 1 i 1 1 i 1 1 i 1 1 0.7 1.0 I.S 0.6 1.0 1.5 0.7 1.0 1.3 0.7 1.0 1.6 0.7 1.0 1.3 0.6 1.0 1.6 FigF-35 Variation in cone scale morphology in a sympatric population of white and black spruce. 11 is the characteristic curve of pure white spruce on pure black. 12 is the characteristic curve of pure black on pure white. TN - tree number, C - classification, B - black spruce, W - white spruce, I -intermediate form. ^ ON w w 1 i 1 : 1 i 1 1 I 1 1 1 i 1 1 i f-1.0 1.7 0.7 1.0 I.T 0.7 1.0 1.7 0.6 1.0 1.3 2.0 0.6 1.0 1.5 0.6 1.0 Variation in cone scale morphology in a sympatric population of white and black spruce. 11 is the characteristic curve of pure white spruce on pure black. 12 is the characteristic curve of pure black on pure white. TN - tree number, C - classification, B - black spruce, ¥ - white spruce, I - intermediate form, (it was not possible to classify trees 2-k)• T N I 2 3 4 5 C W W W W W i 1 1 i 1 1 i 1 1 1 i 1 1 i 1 i i 1 1 0.6 1.0 1.6 0.7 1.0 1.7 0 6 1.0 1.5 1.9 0.7 1.0 15 0.7 10 1.8 0.6 1.0 1.6 Fig. 37 Variation in cone scale morphology in a sympatric population of white and. black spruce. 11 is the characteristic curve of pure white spruce on pure black. 12 is the characteristic curve of pure black on pure white. TN - tree number, C - classification, B - black spruce, W - white spruce, I - intermediate form. co TN I 2 3 4 5 C W W I I W 0.7 1.0 I.B 0.7 1.0 1.7 O.B 1.0 1.5 0.8 1.0 1.6 0.7 1.0 1.6 0.6 1.0 1.6 I 1 1 » 1 1 t I I I I ' 1 I I ' I | I 1 1 I \ 1 I 1 1 I 1 1 I 1 1 I 1 1 0.7 1.0 1.8 0.7 1.0 1.7 0.8 1.0 1.5 0.8 1.0 1.6 0.7 1.0 1.6 0.6 1.0 1.6 Fig. 38 Variation in cone scale morphology in a sympatric population of white and black spruce. 11 is the characteristic curve of pure white spruce on pure black. 12 is the characteristic curve of pure black on pure white. TN - tree number, C - classification, B - black spruce, ¥ - white spruce, I -intermediate form. ^ NO TN I 2 3 4 5 C W W W I I 0.6 1.0 1.6 0.6 1.0 1.7 0.7 1.0 1.6 0.8 1.0 1.5 OB 1.0 1.6 0.6 1.0 1.6 • I I I I : I I I 1 I I I I I I I | I I I I I I 1 I I 1 I I 1 I ( — — — — , , 1 1 " 0.6 1.0 1.6 0.6 1.0 1.7 0.7 1.0 1.6 0.8 1.0 1.5 0.8 1.0 1.6 0.6 1.0 1.6 Fig. 39 Variation in cone scale morphology in a sympatric population of white and black spruce. 11. is the characteristic curve of pure white spruce on pure black. 12 is the characteristic curve of pure black on pure white. TN - tree number, C - classification, B - black spruce, ¥ - white spruce, I -intermediate form. § C B W B W W 0-6 1,0 1.8 0.6 1.0 1.7 0,6 1.0 1.4 0,7 1.0 1.5 0 7 1.0 16 06 10 16 ' 1 : 1 I | 1 I + 1 I 1 1 I 1 I i | i p 1 1 I 1 1 I 1 1 1 I 1 I 1 1 I I 1 0.6 1.0 1.8 0.6 1.0 1.7 0.6 1,0 1.4 0.7 1.0 1.5 0.7 1.0 1.6 0.6 1.0 1.6 Variation in cone scale morphology in a sympatric population of white and black spruce. 11 is the characteristic curve of pure white spruce on pure black. 12 is the characteristic curve of pure black on pure white. TK - tree number,, C - classification, B - black spruce, ¥ - white spruce, I -intermediate form. TN I 2 3 4 5 C B W I W W I I 1 I 1 I l 1 I I 1 1 I "r"' '—" 1 r ! 0.5 1.0 1.5 0.6 IJO 1.6 0.7 1.0 1.5 0.7 1.0 1.5 0.7 1.0 1.4 0.6 1.0 1.6 Fig, hi Variation~in cone scale morphology in a sympatric population of white and black spruce, 11 is the characteristic curve of pure white spruce on pure black, 12 is the characteristic curve of pure black on pure white. TN - tree number, C - classification, B - black spruce, W - white spruce, I -intermediate form, o ro T N I 2 3 4 5 C W B B B B 0.7 1.0 1.6 0.6 1.0 1.5 0.6 1.0 1.6 0.6 1,0 1.6 0.6 1.0 1.5 0.6 1.0 1.6 I I I I I 1 I I I I I 1 I I I I I | Tig. hZ Variation in cone scale morphology in a sympatric population of white and black spruce. 11 is the characteristic curve of pure white spruce on pure black. 12 is the characteristic curve of pure black on pure white, TN - tree number, C - classification, B - black spruce, W - white spruce, I -intermediate form. o I 1 i r- " I 1 I J i i i i i 1 1 • 1 ' O.B 1.0 1.4 0.6 1.0 1.5 07 1.0 1.6 1.4 1.0 1.8 0.5 1.0 1.7 0.6 1.0 1.6 Fig. U3 Variation in cone scale morphology in a sympatric population of white and black spruce. 11 is the characteristic curve of pure white spruce on pure black. 12 is the characteristic curve of pure black on pure white. TN - tree number, C - classification, B - black spruce, W - white spruce, I -intermediate form. o T N I 2 3 4 5 C B W B I W Fig, Uh Variation in cone scale morphology in a sympatric population of white and black spruce. 11 is the characteristic curve of pure white spruce on pure black. 12 is the characteristic curve of pure black on pure white. TN - tree number, C - classification, B - black spruce, ¥ - white spruce, I -intermediate form, o Fig. h$ Sympatric and allopatric spruce populations as indicated by discriminant function analysis of cone scale data. 1963 collection. Fig. U6 Sympatric and allopatric spruce populations as indicated by discriminant function analysis of cone scale data. l°61i collection. ^ o 

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