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

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SOME PROBLEMS IN TESTING PROVENANCE WITH SPECIAL REFERENCE TO THE CO-OPERATIVE DOUGLAS-FIR PROVENANCE TEST AT THE UNIVERSITY OF BRITISH COLUMBIA RESEARCH FOREST by FRANZ REUTER o Dlplomforstwirt, University of Freiburg, Germany, 196? A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF Master of Forestry i n the Faculty of Forestry We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA February, 1971 In presenting th i s thes i s in p a r t i a l f u l f i lment of the requirements fo r an advanced degree at the Un ivers i ty of B r i t i s h Columbia, I agree that the L ibrary sha l l make i t f r ee l y ava i l ab le for reference and study. I fu r ther agree tha permission for extensive copying of th i s thes i s fo r scho lar ly purposes may be granted by the Head of my Department or by his representat ives. It is understood that copying or pub l i ca t ion of th i s thes i s f o r f i nanc ia l gain sha l l not be allowed without my wr i t ten permission. Department of Forestry  The Univers i ty of B r i t i s h Columbia Vancouver 8, Canada Date February 3rd, 1971 ABSTRACT Importance of research on the provenance problem, which i s b a s i c a l l y one of seed transfer from c o l l e c t i o n s i t e to outplanting area, i s discussed with sp e c i a l emphasis on coastal Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. menziesii). The "Co-operative Douglas-fir provenance t e s t , " begun i n 1957 and involving sixteen coastal seed sources from B r i t i s h Columbia, Washington and Oregon, i s described i n d e t a i l . Height measurements co l l e c t e d at the University of B r i t i s h Columbia Research Forest, when the trees were eleven years old, are analysed and the r e s u l t s are discussed. Due to s i t e hetero-geneity and young age of the Co-operative t e s t , no s i g n i f i c a n t height growth differences between provenances can be shown, although the l o c a l seed source, from the University of B r i t i s h Columbia Research Forest, seems to be the fastest growing and the southernmost o r i g i n , Butte F a l l s , the slowest of a l l prov-enances investigated. The Co-operative test i s c r i t i c a l l y evaluated and s p e c i f i c a t i o n s f o r further studies are recom-mended . ACKNOWLEDGMENTS Above a l l I wish to thank Dr. P.G. Haddock who suggested the subject f o r t h i s thesis and pat i e n t l y provided help and guidance. The measurements were obtained with assistance from Mr. J . Walters who constructively c r i t i c i z e d thesis d r a f t s . I am indebted to Dr. A. Kozak and Miss L. Cowdell f o r t h e i r assistance i n analyzing the data. Special thanks are due Dr. 0. S z l k l a i f o r cha i r i n g the thesis committee. Several graduate forestry students gave me much of th e i r precious time. The help of Messrs. D. Ormerod, L. L a c e l l e and G. P a i l l e w i l l be g r a t e f u l l y remembered. Dr. K.K. Chlng of the Oregon State University o b l i g i n g l y answered many questions. Mr. C. Heaman lent me his excellent report on the "Co-operative Douglas-fir provenance t e s t " i n B r i t i s h Columbia. I am deeply obliged to my wife Rita f o r her continual support. i l l ABSTRACT i ACKNOWLEDGMENTS i i TABLE OF CONTENTS i i i LIST OF TABLES . * v LIST OF FIGURES v i CHAPTER I. INTRODUCTION AND OBJECTIVES 1 I I . LITERATURE REVIEW ON PROVENANCE 1 D e f i n i t i o n of Provenance 2 The Provenance Problem 2 Objectives of Provenance Tests 6 Provenance Tests and Forest Tree Improvement 7 B r i e f History of Provenance Testing 7 Problems Commonly Investigated 8 Drought resistance . . . . . . 9 Site ecotypes 9 Aspect races 10 Bud bursting 11 Lammas shoots • l 4 Frost resistance l 6 Cline or ecotype? 19 Climate 20 Seed movement . 22 Early Tests 24 I I I . MAJOR PROVENANCE TESTS WITH DOUGLAS-FIR. . . . 2? Tests Made Outside Its Natural Range . . . . 27 Major Provenance Tests With Douglas-fir Within Its Range 32 IV. THE CO-OPERATIVE DOUGLAS-FIR PROVENANCE TEST . 38 General. 38 Hypotheses to be Tested. . . . . . . . . . . 39 Design 40 Co-operators and Seed Sources 41 Phases . 46 Page Selection of outplanting areas and seed c o l l e c t i o n . 46 Nursery phase. 4 7 Planting of the nursery stock. . . . . . 4 8 F i e l d examinations 4 8 Results. 4 9 V. METHODS 55 The Study Area 55 General description of the University of B r i t i s h Columbia Research Forest . . . 5 5 Climate. . . 56 Location of the co-operative provenance tes t on the Research Forest. . . . . . 58 History of the te s t s i t e . . . . . . . . 58 Height Measurements. 60 S t a t i s t i c a l Analysis 6 l VI. RESULTS AND DISCUSSION 6 l Height at Age Eleven 6 l Results by Individual Block 65 Early Tests 67 Brush Competition. . 73 The Local Provenance 75 How do These Results Compare with E a r l i e r Results? 76 VII. EVALUATION OF THE CO-OPERATIVE PROVENANCE TEST 77 Good Points 77 Weak Points 77 VIII. CONCLUSIONS AND RECOMMENDATIONS 79 IX. SUMMARY 81 LITERATURE CITED 83 APPENDIX. S c i e n t i f i c and common names of species c i t e d i n the text 93 Page Table 1 Go-operators and lo c a t i o n of the te s t s i t e s 4 3 Table 2 D i s t r i b u t i o n of the seed sources 4 4 Table 3 Height performance at age eleven 66 Table 4 Height performance. Block I 69 Table 5 Height performance. Block II 70 Table 6 Height performance. Block III 71 Table 7 Height performance. Block IV 72 Table 8 Height performance between the years-1 9 6 4 - 1 9 6 7 74 Figure 1 Diagram of plot designations Figure 2 Blow-up of plot X i n plantation Y Figure 3 Geographic d i s t r i b u t i o n of seed sources i n the P a c i f i c Northwest Douglas-fir proven-ance test Figure 4 Ormerod's map Figure 5 Location map of the co-operative provenance test at the University of B r i t i s h Columbia Research Forest Page 4 l 42 L5 63 68 I. INTRODUCTION AND OBJECTIVES In 1954 the Oregon State Board of Forestry met to consider the p o s s i b i l i t y of studying the genetic v a r i a b i l i t y of the coastal form of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. menziesii) from sixteen sources i n Oregon, Washington and B r i t i s h Columbia. The following major reasons made that study a necessity1 1. the importance of the provenance problem, 2. the lack of knowledge on the performance of Douglas-fir provenances i n the P a c i f i c Northwest. The present thesis was written with two objectives 1 to review l i t e r a t u r e on the provenance problem, to i l l u s t r a t e t h i s review by an analysis of height measurements of the "Co-operative Douglas-fir prov-enance t e s t , " taken at the University of B r i t i s h Columbia Research Forest. I I . LITERATURE REVIEW ON PROVENANCE No attempt i s made i n the following to give an ex-haustive review on t h i s subject, which i s f a r too vast to be condensed within the scope of t h i s t h e s i s . An excellent survey of pertinent Douglas-fir provenance l i t e r a t u r e has recently been published by Haddock et a l . (1967). D e f i n i t i o n of Provenance According to Wright (1962), provenance (or proven-ience) means "the geographic source of a l o t of seed (or p o l l e n ) . " This d e f i n i t i o n p a r a l l e l s very c l o s e l y that given by Snyder (1959) and Lines (1967). Lines statedi "Provenance i s synonymous with population. The term 'race* re f e r s to one or more natural populations showing defined c h a r a c t e r i s t i c s . Prov-enance does not carry t h i s implication, hence the broader term provenance i s more generally applicable when populations are investigated." In the present thesis "source" or " o r i g i n " w i l l be used synonymously with "provenance" to avoid r e p e t i t i o n . The Provenance Problem Species exhibit natural v a r i a t i o n as a r e s u l t of evolutionary adaptation to d i f f e r e n t environments i n space and time. According to Haddock (1967)» the major factors respon-s i b l e f o r the development of l o c a l l y , g e n e t i c a l l y d i f f e r e n t populations, are believed to be "the great topographic and associated c l i m a t i c and edaphic v a r i a t i o n i n the environment." This v a r i a t i o n , the nature and extent of which are hardly ex-plored f o r any tree species, can be continuous ( c l i n a l ) or discontinuous ( e c o t y p i c a l ) . When transplanted to a new en-vironment within or outside the natural range of the species, d i f f e r e n t geographic sources w i l l react d i f f e r e n t l y . This d i f -ference i n reaction cannot be predicted i n d e t a i l . Good per-formance of a p a r t i c u l a r provenance i n one place i s no assurance of Its s u p e r i o r i t y elsewhere. I t i s generally be-l i e v e d (e.g., S i l e n 1 9 6 6 ) that l o c a l sources are best adapted to t h e i r p a r t i c u l a r environment and w i l l outperform trees of non-local o r i g i n i n the long run, e s p e c i a l l y i n areas with occasional c l i m a t i c extremes such as exceptional f r o s t s . Transfer of seed should only be made between regions showing as much c l i m a t i c s i m i l a r i t y as possible i n order to avoid undesir-able s i l v i c u l t u r a l r e s u l t s (Isaac 1 9 ^ 9 ) . Larsen ( 1 9 5 6 ) , the father of modern forest genetics, does not agree with the idea of s t r i c t l y adhering to c l i m a t i c -a l l y s i m i l a r areas when t r a n s f e r r i n g seed. He preaches bold optimism and i l l u s t r a t e s his point with the superior perform-ance i n Denmark (at 5 6 ° l a t i t u d e North) of Norway spruce (PJLcea ables (L.) Karst) from a Romanian source (46° l a t i t u d e North), of d i s s i m i l a r climate. The f a c t that non-local su p e r i o r i t y has not yet been proven f o r Douglas-fir does not necessarily mean that l o c a l provenances are always superior, i t merely indicates the lack of both knowledge and experimentation i n t h i s f i e l d . Some indications of "nonoptimality" of l o c a l sources f o r several species, including l o b l o l l y pine (Pinus taeda L . ) , were recently reported by Namkoong ( 1 9 6 9 ) * Apparently evidence Is mounting that an optimal growth zone of t h i s species exists along the southeastern border of i t s range. Further experi-ments w i l l prove whether seed from t h i s zone w i l l outproduce l o c a l stock elsewhere. Wheat ( 1 9 6 6 ) l i s t e d several reasons f o r there being an important provenance problem with Douglas-fir i n the P a c i f i c Northwest. They are b r i e f l y J 1. Lack of l o c a l seed i n s u f f i c i e n t quantities due to i r r e g u l a r seed crops. 2. Urgency of immediate r e f o r e s t a t i o n due to the c a p a b i l i t y of high s i t e lands i n the P a c i f i c North-west to grow a dense cover of brush as f a s t or fas t e r than Douglas-fir would grow. 3. Lack of s u f f i c i e n t knowledge on the ad a p t a b i l i t y of other sources to the l o c a l s i t u a t i o n , i . e . lack of f i e l d - t e s t e d seed transfer r u l e s . 4. Lack of controls on seed movement. Considering the number of years i t takes to grow a stand to maturity and the time that might elapse before a prov-enance shows i t s i n f e r i o r i t y , i t i s economically and s i l v i -c u l t u r a l l y unsound to b l i n d l y buy a seedlot of which the o r i g i n , and therefore performance, are not known. It i s better i n such a case not to plant but rather to wait f o r good seed. Thus, i n p r a c t i c a l f o r e s t r y , the provenance problem i n essence i s to f i n d populations of trees, whether the species be indigenous or not, of which the seed w i l l grow forests that are well adapted to t h e i r environment and that produce more wood than trees from any other population of the same species during the same period of time. The provenance problem i s Productivity may not simply be equal to rapid growth. In a p a r t i c u l a r provenance t e s t , the c r i t i c a l f a c t o r i n productiv-i t y may be the a b i l i t y to withstand f r o s t (e.g. Sitka spruce i n Western Norway) or the a b i l i t y to produce f e r t i l e seed (e.g. Scots pine (Plnus s y l v e s t r l s L.) i n Northern Finland (Lines 1967). intimately connected with the problem of f i n d i n g the best spe-cies f o r a p a r t i c u l a r area. Technical problems such as s t a t i s t i c a l design, layout and control of provenance experiments have been thoroughly d i s -cussed by Edwards ( 1 9 5 6 ) and Lines ( 1 9 6 7 ) . According to Schmidt ( 1 9 6 2 a ) , provenance t r i a l s can be divided into the following categories! 1 . Studies of broad geographic v a r i a t i o n . Studies of broad geographic v a r i a t i o n are not s p e c i f i c a l l y designed to provide data f o r seed transfer rules; they usually involve a small number of provenances which may not be a suf-f i c i e n t l y representative sample of a given species. This type of study usually indicates that l o c a l seed i s best, although that might not always be correct. 2 . Tests of stock from improved seed production areas. Studies in v o l v i n g stock from seed production areas can be c r i t i c i z e d on the basis that the seed production areas may no longer be function-ing by the time the r e s u l t s are obtained and that they usually involve too few provenances. 3 . Comprehensive t r i a l s . Comprehensive t r i a l s require much ground work, such as c l i m a t i c s t r a t i f i c a t i o n of the range of the species i n question. Such a s t r a t i f i c a t i o n i n i t s e l f i s an overwhelming task i n some areas. Reciprocal plantations have to be established within each zone. Seed samples may be too few to adequately capture the v a r i a t i o n within the species. 4. F a c t o r i a l t e s t s . The influence of the i n d i v i d u a l c l i m a t i c (and geographic) variables considered to be of major importance to ecotypic d i f f e r e n t i a t i o n are de-termined by f a c t o r i a l tests which are of great value i n areas where climate shows l i t t l e , or at l e a s t , predictable v a r i a t i o n with changes i n geographic factors such as l a t i t u d e . 5. Random population t r i a l s . Random population t r i a l s involve a large number of randomly picked provenances and a large num-ber of outplanting s i t e s representative of the c l i m a t i c v a r i a t i o n within the range of the species. Although some c l i m a t i c data are re-quired before e s t a b l i s h i n g the test to a s s i s t i n s e l e c t i n g the outplanting s i t e s , the bulk of the information can be accumulated during the study. This approach was adopted by Schmidt ( 1 9 6 7 ) to study the v a r i a t i o n within the B r i t i s h Columbia coastal form of Douglas-fir. Provenance t r i a l s should not be mistaken f o r progeny tests which follow them chronologically. Provenance studies determine the genetic worth of populations of trees; progeny tests determine the genetic worth of an i n d i v i d u a l tree (Lines 1 9 6 7 ) . Objectives of Provenance Tests Out of the previous paragraphs, the following major objectives can be defined t 1 . To study extent and nature of the v a r i a t i o n i n parts or i n the entire range of a species i n order to locate populations of trees, the seed of which w i l l produce well adapted, productive forests i n a given region (Lines 1967 and Place 1 9 6 9 ) . 2 . To define the genetic and environmental components of t h i s phenotypic ( i . e . morphological and/or physiological) v a r i a b i l i t y between trees from d i f -ferent geographic sources (Lines 1 9 6 7 ) * 3 . To e s t a b l i s h seed transfer rules (Schmidt 1 9 6 7 ) . Provenance Tests and Forest Tree Improvement According to Nanson (1964a), tree improvement pro-grams b a s i c a l l y aim at three points i 1. quantity (of wood and/or other organic products), 2. q u a l i t y (of v i s i b l e , e.g. stem straightness and/or i n v i s i b l e c h a r a c t e r i s t i c s such as s p e c i f i c g r a v i t y ) , 3. resistance (to b i o t i c environment, to s o i l and to climate)• The following tools can be used to reach these objectives i 1. Provenance studies 2. Seed production areas 3. Seed orchards 4. Inter- and i n t r a - s p e c i f i c h y b r idization 5. Induced mutations and polyploidy 6. Vegetative propagation. According to Nanson (1964a), provenance tests should be designed s t a t i s t i c a l l y i n such a way that growth performance, form and resistance of the various origins under Investigation can be assessed. Provenance tests constitute the f i r s t l o g i c a l step i n genetic research. Although a r e l a t i v e l y easy endeavour, provenance tests are able to y i e l d p r a c t i c a l r e s u l t s quickly. They often show without a d d i t i o n a l cost, which stand or region to buy seed from. B r i e f History of Provenance Testing The Frenchman de Vilmorln i s t r a d i t i o n a l l y considered to be the father of provenance t e s t i n g . In the early 1820's he established Scots pine plantations of known geographic o r i g i n at his estate at l e s Barres. In 1862 he wrote a report on the growth of these trees and concluded that there was a d e f i n i t e difference i n several important c h a r a c t e r i s t i c s be-tween trees from d i f f e r e n t regions. Unfortunately, these d i s -coveries went unheeded (Larsen 1 9 5 6 , Revel i 9 6 0 ) . C i e s l a r ( 1 9 0 7 ) and Engler ( 1 9 1 3 ) demonstrated the existence of great differences between various origins of Norway spruce. The 1912 Douglas-fir heredity study constitutes the pioneer e f f o r t i n provenance t e s t i n g i n North America. I t was i n i t i a t e d by the United States Forest Service which also pro-vided the seed f o r the Douglas-fir tests started by Schwappach and Muench around 1912 i n Germany. The f i r s t tentative seed transfer rules were established i n Sweden i n the 1 9 3 0 ,s. With the growing awareness of the importance of the problem and the po t e n t i a l gains to be derived from the selec-t i o n of adequate-seed sources, provenance tests became more and more popular i n forest research around the world. They are too numerous to be reported here i n d e t a i l . Problems Commonly Investigated The following i s an a r b i t r a r y d i s t i l l a t i o n of l i t e r -ature f o r some major autecological aspects of the provenance problem, with s p e c i a l reference to Douglas-fir. Drought resistance According to F e r r e l l and Woodland (1966), drought re-sistance may involve ei t h e r drought avoidance or drought hardi-ness, or both. Drought avoidance i s accomplished through root extension and reduction i n l e a f area or through early onset of dormancy. Drought hardiness i s the a b i l i t y to survive i n a dry external environment causing severe tissue dehydration. The authors studied the e f f e c t of seed o r i g i n on drought resistance of Douglas-fir and found that I n t e r i o r mountain provenances showed s i g n i f i c a n t l y greater drought resistance than those from areas west of the Cascades. Seedlings from C o r v a l l i s were a notable exception. They proved no more drought r e s i s t a n t than seedlings from higher p r e c i p i t a t i o n , lower temperature areas elsewhere west of the Cascades. By two drought hardiness t e s t s , time to death and s o i l moisture content at the death point, Pharis and F e r r e l l (1966) showed that Douglas-fir seedlings from three coastal sources were les s drought r e s i s t a n t than those from f i v e inland sources. They found needle moisture to be a good index f o r de-termining plant v i a b i l i t y under well watered conditions. Seed-l i n g s could be c l a s s i f i e d into coastal and inland groups on the basis of t h e i r needle moisture content with the exception of an Arizona provenance behaving l i k e the coastal group. Site ecotypes A basic tenet of genetics holds that the phenotype i s affected by the genotype and the environment. Provenance studies are designed to enable the researcher to detect the genetic differences between various seed sources. This can best be done by growing the trees on a hopefully uniform s i t e under the same environmental conditions. Since s i t e i s one of the major components of the environmental complex, i t . i s not sur p r i s i n g that some foresters have t r i e d to f i n d out whether populations had adapted themselves gen e t i c a l l y to growing on a p a r t i c u l a r s i t e . The f i r s t authors to report on t h i s problem i n Douglas-fir were Munger and Morris (1936) who found that the age of the parent tree, i t s growing space, i t s conditions as to fungus i n f e c t i o n and i t s s i t e index, had no e f f e c t upon height growth of the progeny. Gathy (19^1 and 19&7) tested various provenances of coastal Douglas-fir and came to s i m i l a r conclu-sions » age and growing s i t e of the parent trees seemed to have no bearing on height growth of t h e i r o f f s p r i n g . Heaman (1968) found no s i g n i f i c a n t differences i n height growth between two high elevation Douglas-fir provenances. One l o t of seed had been c o l l e c t e d from a stand showing r e l a t i v e l y good phenotypic c h a r a c t e r i s t i c s on a good s i t e while the other l o t originated from trees growing on a much poorer s i t e , but i n a s i m i l a r c l i m a t i c zone. Aspect races Although there has been no d e f i n i t e proof so f a r of s i t e ecotypes i n Douglas-fir, i t i s quite i n t e r e s t i n g to note that ecotypes have evolved that are adapted to the p a r t i c u l a r conditions p r e v a i l i n g on southerly aspects. F e r r e l l and Wood-land ( 1 9 6 6 ) found that seedlings produced from trees growing on a south slope were more drought r e s i s t a n t than those from trees growing on a north slope, a short distance away. Interesting r e s u l t s were obtained by Hermann and Lavender ( 1 9 6 8 ) i n a growth chamber experiment with Douglas-fir from various a l t i t u d e s and aspects i n southern Oregon. Progeny of trees from south facing aspects exhibited a shorter growing period and lar g e r roots i n r e l a t i o n to t h e i r tops than seedlings from parents on north facing slopes. This was interpreted to be a r e s u l t of natural s e l e c t i o n f o r early cessation of growth i n habitats that are p a r t i c u l a r l y dry i n summer. These r e s u l t s are an Interesting p a r a l l e l to e a r l i e r findings by Squillace and Bingham ( 1 9 5 8 ) suggesting the existence of aspect ecotypes i n western white pine (Pinus montlcola Dougl.). Bud bursting Bud bursting, as discussed i n t h i s chapter, r e f e r s to the opening of vegetative buds only and i s synonymously used with "flushing." 1 . The influence of genetic control on the timing of f l u s h i n g . Observations by Morris et a l . (1957) suggest strong genetic control over time of bud bursting i n Douglas-f i r . The most c l a s s i c a l study on t h i s question was made by S i l e n ( 1 9 6 2 ) , who t r i e d to minimize environ-mental influences by g r a f t i n g several r e p l i c a t i o n s of scions from trees with large, known differences i n f l u s h i n g dates onto limbs of seven trees of a seventeen year old clone. As a con t r o l , a limb of the cl o n a l tree was cut and re-grafted. S i l e n estimated the gen-e t i c component of the bud bursting t r a i t at ninety-four and ninety-six percent of the t o t a l v a r i a t i o n f o r the two years of observations. At the same time he pre-sented evidence that l o c a l environmental differences may consistently delay bud bursting of genet i c a l l y s i m i l a r material f o r as much as two weeks. S i m i l a r l y , public-ations by the United States Forest Service (1964) and G r i f f i t h (1968) show evidence of strong genetic control over fl u s h i n g i n Douglas-fir. The trees under obser-vation f o r several years consistently flushed i n the same order, within a day or two. The pattern was not a l t l t u d l n a l . Walters and Ching (1969) studied the pattern of bud burst i n the Douglas-fir provenance testi? to be reported l a t e r In t h i s t h e s i s , and found the University of B r i t i s h Columbia Research Forest prov-enance to be a l a t e flusher, regardless of plantation; the Salem source proved to be the e a r l i e s t flusher of terminal and l a t e r a l buds, regardless of planting area. Evidence was put forward f o r a strong influence of l o c a l c l i m a t i c conditions on bud bursting. 2 . The e f f e c t s of the environment on f l u s h i n g . The publications c i t e d In the previous chapter, s t a t i n g that under uniform environmental conditions bud burst-ing i s under very strong genetic c o n t r o l , also mention the e f f e c t of the environment, p a r t i c u l a r l y the c l i -matic conditions of any p a r t i c u l a r year, on the expression of t h i s t r a i t . Morris et a l . (1957) ob-served that i n some years flushing may occur as much as a month l a t e r than i n other years. In high elev-atio n plantations, buds may open anywhere from two weeks to two months l a t e r than at low elevations. G r i f f i t h (1968) t r i e d to correlate date of f l u s h i n g with c l i m a t i c data and found that i n the University of B r i t i s h Columbia Research Forest bud bursting was influenced by weather conditions during the forty-two day period p r i o r to May the s i x t h . 3. The disadvantages of early bud bursting. Time of f l u s h i n g i s important l n r e l a t i o n to spring f r o s t s . Since bud bursting i s a highly hereditary t r a i t , seed from early flushers cannot be planted to advantage i n areas where l a t e f r o s t s are l i k e l y to occur. Irgens-M^ller (19&7) noticed a difference i n time of bud bursting of f i f t e e n to eighteen days be-tween plants o r i g i n a t i n g from areas separated by only twenty to f o r t y miles. He stressed the importance of choosing a seed source that can be expected to be un-affected by spring f r o s t s . Needle midges (Contarlnla spp.) were found to threaten early f l u s h i n g Douglas-f i r trees (Mitchell and Nagel, 1969). while leaving l a t e flushers more or l e s s unharmed. Schober (19&3) advised against Douglas-fir provenances from B r i t i s h Columbians I n t e r i o r Wet Belt f o r use i n areas with l a t e f r o s t s or widely f l u c t u a t i n g weather conditions during spring, such as occur i n Western Europe, because they f l u s h e a r l i e r than coastal provenances. This e a r l i e r f lushing makes them more susceptible to the Douglas-fir needle b l i g h t (Rhabdocllne pseudotsugae Syd.). Haddock et a l . (19^7) came to s i m i l a r conclus-ions. To summarize, early f l u s h i n g i s hardly an asset, since spring f r o s t s , insects and fungi may damage the trees severely. 4. Implications of time of fl u s h i n g on the provenance problem. When choosing seed l o t s , geographic and cl i m a t i c data should be supplemented with c a r e f u l observations on date of bud bursting covering several years. The suc-cess of whole plantations may hinge on time of flu s h i n g i n connection with spring f r o s t s , insects or fungi. Late f l u s h i n g eliminates a l l these dangers and has no negative e f f e c t on t o t a l height growth. Ideal proven-ances are those that grow f a s t , f l u s h l a t e and set t h e i r buds early, thus avoiding both spring f r o s t s and f a l l f r o s t s . Lammas shoots According to Walters and Soos (196lb), " ... l a t e r a l and terminal vegetative buds of young trees frequently break dormancy In l a t e summer and produce extra-seasonal shoots named lammas shoots a f t e r the old English harvest f e s t i v a l s o f Lammas on August 1 . " The authors studied t h i s phenomenon on Douglas-f i r seedlings from various origins grown at several elevations on the University of B r i t i s h Columbia Research Forest. The conclusions were that lammas growth 1) was influenced more by environmental than genetic f a c t o r s , 2) decreased with increasing age, 3) increased as s i t e q u a l i t y increased, 4) increased as current height growth increased f o r seedlings of a p a r t i c u l a r provenance. Lammas growth was sometimes r e f l e c t e d l n f a l s e annual rings. Forked leaders caused by lammas shoots were overcome quite r a p i d l y i n Douglas-fir. Sweet ( 1 9 6 5 ) found that Douglas-fir seedlings having more than one annual growth f l u s h i n t h e i r second year do not make a greater height increment than provenances with fewer such trees. Hoffman ( 1 9 6 5 ) experienced other r e s u l t s with Norway spruce, where the incidence of lammas shoots varies geo-graphically and decreases with increasing elevation of seed source as an adaptation to the l o c a l climate. For s i t e s with no early f r o s t s , Hoffman recommended provenances with a high occurrence of lammas shoots that would outgrow dangers of juven-i l e stages more quickly. Schmidt-Vogt ( 1 9 6 6 ) also noticed that lammas growth decreases with increasing a l t i t u d e of seed source i n Norway spruce, and he found i t s occurrence on one and two-year old seedlings to be a workable early test to check on elev a t i o n a l authenticity of seed. To conclude, lammas shoots constitute a disadvantage on s i t e s where early f r o s t s are common, since the s o f t shoots can be damaged e a s i l y by f r o s t . If t h i s happens repeatedly, trees of poor form may r e s u l t . On s i t e s free of early f r o s t hazards, lammas shoots are an advantage In overcoming compet-i t i o n from l e s s e r vegetation, and possibly from deer browsing. Frost resistance Schoenbach ( 1 9 5 8 ) evaluated the e f f e c t s of a mass se l e c t i o n of Douglas-fir seedlings f o r f r o s t resistance. The s e l e c t i o n had given two populations i one mainly f r o s t sensit-ive, the other f r o s t r e s i s t a n t . Schoenbach concluded that f r o s t resistance has single gene inheritance and appears i n a homozygous condition. He stressed the need to s t a r t provenance studies on a very large b a s i s , that i s , Including as many seed sources as possible, i n order to f i n d populations containing resistance genes. According to Scheumann ( 1 9 6 5 )» the following f i v e points have to be considered when f r o s t resistance i s to be studied! 1 . readiness to harden o f f (= early f r o s t r e s i s t a n c e ) . 2 . extent of hardening o f f (= winter f r o s t r e s i s t a n c e ) . 3. s t a b i l i t y of dormancy (Important during winters with highly varying temperatures). 4. time of f l u s h i n g and flowering (late f r o s t r e s i s t a n c e ) . 5 . regeneration p o t e n t i a l ( a b i l i t y to overcome f r o s t damage)• These f i v e points make i t quite obvious thatt - f r o s t resistance i s a dynamic process that cannot be studied as a s t a t i c condition, as has been done t r a d i t i o n a l l y . - there cannot "be a single t e s t with which to evaluate a l l aspects of f r o s t resistance i n plants * Previous knowledge on t h i s subject has always been based on f i e l d observations a f t e r extremely cold weather. Scheumann ( 1 9 6 5 ) devised a laboratory test with which readiness, extent and s t a b i l i t y of hardening-off can be studied under con-t r o l l e d temperature conditions. Seedlings, needles or young twigs can be used l n t h i s test where temperature i s gradually lowered to below freezing point. A f t e r early f r o s t resistance and extent of hardening-off have been evaluated, the temper-ature Is repeatedly raised f o r some time above the freezing point and then lowered again i n order to t e s t s t a b i l i t y of dor-mancy. This i s a method to rapidly mass select and rank various provenances (or progenies f o r that matter) f o r several aspects of f r o s t resistance. Scheumann t r i e d t h i s " s i m p l i f i e d t e s t " on Douglas-fir and on European l a r c h (Larix decldua M i l l ) . He found s i g n i f i c a n t differences i n f r o s t resistance of two-year old Douglas-fir hybrids (glauca x v i r l d i s ) . He found c o r r e l -ations between the climate i n which the l a r c h trees grew and the behaviour of the twigs i n the f r o s t resistance t e s t . Scheumann concluded that t h i s p o s s i b i l i t y of f i n d i n g genetic differences i n f r o s t resistance among plus-trees and t h e i r pro-genies gives new hope fo r successfully t e s t i n g f r o s t r e s i s t -ance i n forest trees. Schoenbach and Bellmann ( 1 9 6 7 ) reported on the same glauca x v i r l d i s hybrids. The hybrids were a l l s i g n i f i c a n t l y more f r o s t r e s i s t a n t than the "green" parents. Since they were a l s o growing f a s t e r than the "blue" v a r i e t y , the hybrids were recommended by Schoenbach and Bellmann fo r the cold climates of western Europe's mountains. In a study involving thirty-one seed sources of coastal Douglas-fir from Washington and Oregon, Gathy ( 1 9 6 1 ) found three Washington provenances from medium and low elev-ations (Yacolt, 200 feet, Castle Rock, 1300 feet, and Forks, 400 feet above sea l e v e l ) to be most f r o s t r e s i s t a n t and there-fore best suited f o r the maritime climate of Belgium. Nanson ( 1 9 6 4 b ) studied the e f f e c t of the severe winter of 1 9 6 2 - 1 9 6 3 on coastal Douglas-fir provenances i n Belgium, but could not confirm the s u p e r i o r i t y of Washington sources over Oregon o r i -gins In terms of f r o s t resistance. According to Nanson, a l l provenances of coastal Douglas-fir tested require adequate pro-t e c t i o n from severe f r o s t , e s p e c i a l l y when they are young. Lacaze ( 1 9 6 4 ) examined Douglas-fir seedlings i n France a f t e r the 1 9 6 2 - 1 9 6 3 "deep freeze" and found considerable v a r i a t i o n i n f r o s t resistance which generally increased with increasing l a t -itude and a l t i t u d e of the seed source. A l i n e j o i n i n g the points 4 9 ° l a t i t u d e North, 1800 feet elevation, roughly divides the r e s i s t a n t from the non-resistant provenances. In a note by Stem ( 1 9 6 6 ) , reviewing r e s u l t s of a recent experiment, Douglas-f i r seedlings from Arizona, New Mexico and Mexico were found to be more f r o s t r e s i s t a n t than provenances from coastal B r i t i s h Columbia, Washington and Oregon, t r a d i t i o n a l l y tested i n Europe and recommended by Schober ( 1 9 6 3 ) . S i l e n ( 1 9 6 6 ) , evaluating the 1912 Douglas-fir heredity study (see below) a f t e r the 1 9 5 5 November f r o s t , formulated the hypothesis that l o c a l provenances have adapted themselves over the centuries to long-term weather extremes and are therefore l e s s damaged by severe f r o s t s than are non-local sources. Cline or ecotype? In 1936 Langlet published an a r t i c l e on the physio-l o g i c a l v a r i a b i l i t y and i t s r e l a t i o n to climate f o r Scots pine i n Sweden. The dry matter content of needles proved to be highly correlated with the number of days during the growing season with an average temperature of s i x degrees Celsius ( 4 3 ° Fahrenheit), or more. Langlet showed a north-south v a r i a t i o n i n Scots pine i n Sweden and claimed t h i s v a r i a t i o n to be con-tinuous. Wright and Baldwin ( 1 9 5 7 ) based t h e i r c r i t i c i s m of Langlet*s a r t i c l e on observations from a Scots pine provenance test i n New Hampshire. They agreed that there i s a north-south v a r i a t i o n i n Scots pine, but t h e i r s t a t i s t i c a l analysis i n d i c -ated that most of the geographic v a r i a t i o n i s discontinuous, not c l i n a l . This would mean that the Swedish seed transfer rules based on the c l i n a l hypothesis and l i m i t i n g cone c o l l e c t -ions to areas within 250 kilometers (approximately 150 miles) north or south and 300 meters (approximately 1 0 0 0 feet) d i f f e r -ence l n elevation from the planting s i t e , should be revised to c o l l e c t i n g seed within the boundaries of the ecotype best s u i t -ed f o r the planting area. This may permit the safe transfer of seed f o r several hundred miles or l i m i t i t to a few miles. Langlet ( 1 9 5 9 ) used Wright and Baldwin's data to show where they had gone wrong and he stated that v a r i a b i l i t y continues i n the same degree as the determining environmental factors vary-continuously. Neither the c l i n a l nor the ecotypical hypothesis has been dlsproven. It i s possible that In is o l a t e d occurrences Scots pine has developed ecotypes that cannot be explained by differences i n the environment, whereas i n large, contiguous areas of i t s vast range the v a r i a t i o n i s c l i n a l , r e f l e c t i n g gradual changes i n the environment. In other words, ecotypes are not necessarily In contrast with c l i n a l v a r i a t i o n . Haddock et a l . (1967) mentioned the v a r i a b i l i t y of ponderosa pine (Plnus  ponderosa Laws.) studied by various authors, some of whom support the c l i n a l , some the ecotypical hypothesis. Haddock et a l . concluded that no matter which a l t e r n a t i v e one subscribes to, " ... one cannot dispense with a thorough knowledge of the geography of a region and an understanding of the influence of top-ography on l o c a l climate, e s p e c i a l l y i n regions as continuously mountainous as southern B r i t i s h Columbia." Climate Since weather measurements can only indicate part of the complex c a l l e d climate, and since the establishment and maintenance of weather stations i n remote areas are c o s t l y , Schmidt (1962b) t r i e d to Interpret climate from phenological observations. For t h i s purpose, time of i n i t i a l pollen release i n Douglas-fir was chosen, because i t i s the f i r s t external i n -di c a t i o n of physiological a c t i v i t y i n spring and i t i s easy to assess, compared f o r instance with determining the s t a r t of cambial a c t i v i t y . The r e s u l t s were not encouraging, and they raised further problems because male bud development does not seem to be related to a simple expression of temperature. Irgens-Mjrfller ( 1 9 6 5 ) questioned the value of c l i m a t i c data l n assigning off-source seed to planting s i t e . Both plant-ing l o c a t i o n and seed source may be f a r away from the nearest weather s t a t i o n . In addition, some provenances may tolerate a wide var i e t y of conditions, others might be narrowly adapted to s p e c i f i c conditions. Haddock and S z i k l a i ( 1 9 6 6 ) divided the range of Douglas-fir l n B r i t i s h Columbia and western Alberta into nine seed c o l l e c t i o n zones based primarily on the i n f l u -ence of climate on geographic v a r i a b i l i t y of populations and the d i s t r i b u t i o n of associated species. Both authors were aware of the merits and l i m i t a t i o n s of such a zonation, and admitted that populations within the i n d i v i d u a l zones are f a r from being homogeneous, since remarkable c l i m a t i c differences due to elevation, topography and p r e c i p i t a t i o n e x i s t within these zones. It i s not s u f f i c i e n t f o r a forester to indicate the general zone from which he wants seed. He should provide d e t a i l e d c l i m a t i c data f o r the plantation s i t e , including at l e a s t mean annual temperature, absolute minimum temperature, mean annual p r e c i p i t a t i o n during growing season, and length of f r o s t free period or dates of e a r l i e s t and l a t e s t f r o s t s . This way the seed c o l l e c t o r may have a better chance of providing suitable seed (Haddock and S z i k l a i , 1 9 6 6 ) . Newnham ( 1 9 6 8 ) published an a r t i c l e on the c l a s s i f i c a t i o n of climate and i t s r e l a t i o n s h i p to tree species, including Douglas-fir, using data from seventy weather stations from many parts of B r i t i s h Columbia, With p r i n c i p a l component analysis on the matrix of c o r r e l a t i o n c o e f f i c i e n t s with nine-teen variables recorded by the weather stations, he computed three new variables accounting f o r ninety-two percent of the t o t a l v a r i a t i o n between weather stat i o n s . The f i r s t v ariable was a general Index of winter and f a l l climate and of the length of the growing season, the second represented the contrast be-tween spring and summer temperatures and p r e c i p i t a t i o n , the t h i r d v a r i a b l e , of l e s s e r importance, represented merely a mea-sure of l a t i t u d e . When the weather stations were grouped, the p a r a l l e l s to Chapman's ( 1 9 5 2 ) c l i m a t i c regions were s u r p r i s i n g . Newnham's approach, with further refinements, may prove very useful f o r zoning the natural range of trees and a s s i s t i n f i n d i n g good matches fo r tree species Introduction elsewhere. Newnham believes that " ... the components can be used as measures of c l i m a t i c s i m i l a r i t y between d i f f e r e n t provenances within a species." Seed movement Isaac (19^9) based his rules f o r Douglas-fir seed c o l l e c t i o n s on Langlet's (19^ 5 ) publication dealing with seed movement l i m i t a t i o n s i n Sweden. These r u l e s , s t i l l v a l i d , were as follows i 1. C o l l e c t seed within 100 miles north or south of planting s i t e i f at s i m i l a r elevations. 2 . A 500 foot r i s e or drop i n elevation from plant-ing s i t e i s allowable i f seed source i s not more than ten miles to the north or south. 3 . For each ad d i t i o n a l ten miles north of planting s i t e , the allowable elevation f o r seed c o l l e c -t i o n i s reduced by f i f t y f e e t , up to 100 miles north. 4 . For each a d d i t i o n a l ten miles south, the elev-ation may be Increased by f i f t y f e e t , up to 100 miles south. 5 . In a rough, broken country, climate should guide more than distance or elevation. Average annual temperature of the seed source should be within two degrees Fahrenheit plus or minus and f r o s t -free period should be s i m i l a r to that of the planting s i t e . 6 . The seed source stand should be t h r i f t y and making average or better than average growth f o r the l o c a l i t y . 7. Individual seed trees should be of good form and should not be excessively limby. Isaac (19^9) stated e x p l i c i t l y that these rules are not to be considered optimum conditions but rather l i m i t a t i o n s f o r seed c o l l e c t i o n s . He recommended to c o l l e c t seed during heavy crop years and store i t f o r poor years. Isaac noted the lack of seed c e r t i f i c a t i o n laws. Despite the concern about the provenance problem, no refined seed transfer rules have been devised yet f o r Douglas-fir, and l o c a l seed Is s t i l l re-commended as safest and probably best (Bingham 1 9 6 6 ) . Wheat ( 1 9 6 6 ) f e l t that the Increase l n a r t i f i c i a l r e f o r e s t a t i o n with Douglas-fir l n Washington and Oregon and the lack of seed from adequate sources made i t easy to ignore rules of safe p r a c t i c e . He also deplored the lack of controls on the movement of seed. Haddock ( 1 9 6 6 ) noted that neither distance of seed movement i n miles nor the change i n elevation i t s e l f are r e a l l y the question of importance, but rather the e f f e c t s of these changes on the l o c a l t o t a l environment. Haddock stressed the importance of a thorough knowledge of the geography of a region and the influence of topography on l o c a l climate i n respect to seed movement problems. Schmidt's ( 1 9 6 7 ) provenance test w i l l hopefully y i e l d enough information f o r new seed transfer rules f o r coastal Douglas-fir i n B r i t i s h Columbia l e s s than f i f t e e n years from now. Physical properties of wood have been shown to be an Important aspect of the provenance problem, but one which needs further i n v e s t i g a t i o n (Haigh, 1 9 6 l > S i l e n , 1964$ Bramhall, 19661 McKlmmy, 1 9 6 6 ) . Early Tests The long rotations represent one of the major handi-caps of f o r e s t r y . Whereas most a g r i c u l t u r a l crops ripen a f t e r one growing season, i t usually takes a forest stand longer than an average human l i f e to grow to maturity. In an e f f o r t to overcome t h i s shortcoming In provenance and progeny t e s t i n g , some foresters have been t r y i n g to develop early t e s t s . Pioneer work i n t h i s f i e l d was done by Schmidt (Germany) and Langlet (Sweden) around 1930 (Nanson 19^5)•• Early tests investigate provenance v a r i a t i o n i n order to give the e a r l i e s t i n d i c a t i o n of the l a t e r performance of the trees (Lines 1 9 6 7 ) . I f s t a t i s t i c a l l y s i g n i f i c a n t correlations can repeatedly be found between early and l a t e r performance ( f i v e , ten or even f i f t e e n years are not good enough f o r management) i n several tests of the same species or va r i e t y , covering a large range of environmental conditions, i t should be possible to set up equations pr e d i c t i n g l a t e r performance on the basis of measurements taken at an early age. In t h i s way the r e l a t i v e i n f e r i o r i t y or supe r i o r i t y of a p a r t i c u l a r proven-ance (or parent tree i n the case of progeny testing) could be recognized early and foresters would know which stand to choose f o r seed c o l l e c t i o n f o r large scale planting programs. Early t e s t i n g begins with the seed, although c l i m a t i c and geographic data on seed source constitute very important background information. Growth chambers f a c i l i t a t e the study of the d i f f e r e n t i a l reaction of provenances under a controlled environment. Irgens-Mjftler*s (1957) investigations on the eff e c t s of various temperatures and photoperiods on Douglas-fir seedlings from d i f f e r e n t sources are a good example of t h i s . Although laboratory tests are very hel p f u l i n provenance t r i a l s , caution must be used i n projecting the re s u l t s to>later growth i n the f i e l d . Nursery performance (e.g. Ching and Bever, i960) may y i e l d some usefu l Information, but again the true value of those observations w i l l be confirmed only many years l a t e r . Orr-Ewing (1967) stressed t h i s need f o r caution when basing premature conclusions on early r e s u l t s . He stated» "Progeny tests can y i e l d enough information at twelve to f i f t e e n years a f t e r planting f o r at l e a s t the poorer provenances to be recog-nized." The same thing can probably be said of provenance t r i a l s . McKlmmy (1966), studying s p e c i f i c gravity of Douglas-f i r from several seed sources, concluded that trees should be over twenty-five years old f o r stand predictions. According to another publication on the 1912 Douglas-fir heredity study (United States Forest Service 1 9 6 4 ) , the r e s u l t s were disap-pointing when seedling heights were compared to mature heights. In a provenance study with European l a r c h , Leibundgut ( 1 9 6 2 ) found that height growth during the f i r s t years was poorly cor-related with l a t e r height growth. Some experiments, however, showed high c o r r e l a t i o n s i Wright and Baldwin (1957) found a s t a t i s t i c a l l y s i g n i f i c a n t c o r r e l a t i o n (r = 0 . 9 3 3 and 0.861 respectively) between three or four year's height and seventeen year's height i n a Scots pine provenance te s t i n New Hampshire. Nanson ( 1 9 6 5 ) * reporting on an i n t e r n a t i o n a l Norway spruce experiment i n Belgium dating back to 1 9 3 8 , found the twenty-five year growth to be strongly correlated with the following "early" c h a r a c t e r i s t i c s ! - length of growing season at seed source - 1 0 0 0 seed weight - fresh weight of seedlings - dry weight of seedlings - bud bursting date - heights at various ages. Nanson f e l t that the length of the t e s t i n g cycle could be considerably reduced i f the planting check could be eliminated. Haddock et a l . ( 1 9 6 7 ) found a c o r r e l a t i o n coef-f i c i e n t of over 0 . 9 3 8 at the 0 . 0 1 p r o b a b i l i t y l e v e l between height at age two and height at f i v e , s i x , seven, eight and eleven years i n a Douglas-fir study at the University of B r i t i s h Columbia Research Forest. For the same species, Lacaze ( 1 9 6 8 ) could show a highly s i g n i f i c a n t c o r r e l a t i o n between height growth at two, f i v e , eight and t h i r t e e n years a f t e r planting (r between two and th i r t e e n years was O . 9 6 ) . Early tests can also be used as a check on the auth-e n t i c i t y of seed l o t s c o l l e c t e d by commercial seed dealers. Sweet ( 1 9 6 5 ) found that both two-year height growth and time l a g between l a t e r a l and terminal bud burst of Douglas-fir prov-enances tested i n New Zealand were s u f f i c i e n t l y strongly cor-related to climate at seed source to do a check on the data of o r i g i n supplied with commercial seed l o t s . Schmidt-Vogt ( 1 9 6 7 ) showed the same thing f o r Norway spruce and Scots pine. I I I . MAJOR PROVENANCE TESTS WITH DOUGLAS-FIR Tests Made Outside Its Natural Range Douglas-fir was discovered by Archibald Menzies i n 1797 at Nootka Sound on Vancouver Island (Krajina 1 9 5 6 ) . David Douglas sent cones and other material, c o l l e c t e d on the banks of the Columbia River, to Europe as early as 1 8 2 7 , The f i r s t provenance experiments with Douglas-fir were started much l a t e r . In 1 9 0 9 and 1910, the United States Forest Service sent Douglas-f i r seed, c o l l e c t e d under the aegis of Zon at d i f f e r e n t points of i t s range from the P a c i f i c Coast to the Rocky Mountains, to Professor Schwappach at Eberswalde, Germany, and to Count von Berg i n Livonia, Russia. The Count noted that trees from Chelan " ... i n the same range of the Cascades as the seed from Snoqualmle, but on the eastern slope, have grown very nearly as high as the seedlings from Snoqualmle, but stand upright and have scarcely suffered at a l l from the f r o s t . " (Zon 1913). In 1912, Professor Muench started another provenance test i n Kaiserslautern, Germany, with seed from ten provenances, sent by Professor Schwappach who was experimenting with nineteen provenances himself, two from low elevations i n Western Wash-ington, three from high elevations i n C a l i f o r n i a , west of the S i e r r a Nevada summit, the remainder from more continental c l i -mates. In 1933. three other provenance tests were started i n Germanyi series one at the foot of Mount Feldberg i n the Black Forest, series two i n Freienwalde and Braunlage, series three i n Gahrenberg. These experiments have been reported by Schober (195*0, Schober and Meyer (195^, 1 9 5 5 ) . Jahn (1955) t Rohmeder ( 1 9 5 6 ) and others. In Germany It soon appeared that there were great differences between provenances as to growth rate, f r o s t and disease resistance, as well as morphology. The seed f o r the 1 9 3 3 experiments was from i l l - d e f i n e d provenances, there-fore morphology was sometimes resorted to f o r clues on o r i g i n (Decker 1 9 6 7 K Although t h i s did not prove very easy due to the great morphological v a r i a b i l i t y of Douglas-fir, the proven-ances were ascribed to three v a r i e t i e s 1 v l r l d l s green or coastal form, c a l l e d varietas menzlesll i n North America glauca blue or Colorado or Rocky Mountain Douglas-fir by L i t t l e ( 1 9 5 3 ) oaesla grey form from the intermountain or more northern i n t e r i o r provenances from Schenck's ( 1 9 3 9 ) c l i m a t i c region 110 B. At present most North American a u t h o r i t i e s on Douglas-fir do not u n i v e r s a l l y agree beyond the menzlesll-glauca separation (Fowells 1 9 6 5 ) t although many German f o r e s t -ers s t i l l frequently write about the caesla v a r i e t y . Another minor controversy raging among English speaking foresters and botanists regards the hyphenation prob-lem i n s p e l l i n g Douglas (-) f i r . Although there are v a l i d arguments fo r and against the hyphen, t h i s problem could e a s i l y be solved by c a l l i n g Pseudotsuga menziesii (Mirb.) Franco simply "Douglas" as some Europeans do. The great v a r i a b i l i t y of Douglas-fir was stressed by Larsen ( 1 9 5 6 )» who wrote as followsi " ... one has to t r a v e l very widely throughout the natural range of Douglas-fir i n order to get an impression of differences i n geograph-i c a l type, but standing i n one place one can, without moving a foot, see many individ u a l s d i f f e r i n g widely i n t h e i r structure; i t i s often more d i f f i c u l t to pick out those that re-semble another ... It does not matter i f one chooses i n C a l i f o r n i a a s i t e i n the Coast Range or l n the S i e r r a Nevada, passes through Oregon and Washington, or i n B r i t i s h Columbia selects a place on Vancouver Island or i n the Rocky Mountains; everywhere one i s bound to be im-pressed by the great i n d i v i d u a l v a r i a t i o n of t h i s tree species ... " Orr-Ewing ( 1 9 6 8 ) concluded from his inbreeding studies that Douglas-fir i s a most heterozygous species, leaving ample opportunities f o r further s e l e c t i o n . In a comprehensive paper presented i n Stockholm, Schober ( 1 9 6 3 ) summed up European provenance studies with Douglas-fir. He concluded that i n Central Western Europe, prov-enances from areas west of the Cascades i n Washington, e s p e c i a l l y from elevations below 700 meters ( 2 0 0 0 feet) were f a s t growing and f a i r l y r e s i s t a n t to Rhabdocllne pseudotsugae Syd. but at the same time rather sen s i t i v e to winter f r o s t . However, l a t e f r o s t s d i d not a f f e c t them because of t h e i r l a t e bud burst. In the cooler climates of Northeastern Europe, the Central European Mountains and the t r a n s i t i o n zones to continental Eastern Europe, c e r t a i n provenances from the i n t e r i o r of B r i t i s h Columbia grew as fast or f a s t e r than trees from coastal Washington. However, they showed greater s u s c e p t i b i l i t y to both Rhabdocllne and l a t e f r o s t s , due to t h e i r early f l u s h i n g , a l -though they were r e s i s t a n t to winter f r o s t s . In coastal regions or i n maritime European climates, the provenances from B r i t i s h Columbia's I n t e r i o r showed only average to poor performance. Schober expanded on the great v a r i a b i l i t y of these sources and t r i e d to explain i t with information from Galoux's ( 1 9 5 2 ) pub-l i c a t i o n on the phytogeography and paleobotany of Douglas-fir. Unfortunately, many European foresters lack Galoux's knowledge on the subject and do not recognize the existence of B r i t i s h Columbia's I n t e r i o r Wet Belt and the extreme c l i m a t i c v a r i a t i o n In B r i t i s h Columbia. This i s p a r t i c u l a r l y s i g n i f i c a n t f o r prov-enances of Douglas-fir around Shuswap Lake, f o r instance. Haddock and S z i k l a i ' s ( 1 9 6 6 ) seed zone map can help seed c o l -l e c t o r s to avoid gross errors In the future. Hopefully these seed zones w i l l be interpreted e c o l o g i c a l l y and not considered to be uniform e n t i t i e s . Schober ( 1 9 6 3 ) recommended the I n t e r i o r Wet B e l t origins f o r appropriate parts of Europe. Those from B r i t i s h Columbia's dry I n t e r i o r , corresponding to Haddock and S z i k l a i * s (1966) seed zones f i v e and eight or to Rowe's (1959) Montane Forest Region, were found to be slow growing and sus-ceptible to Rhabdocllne i n Europe. Seed from coastal Oregon and C a l i f o r n i a showed good r e s u l t s l n the warmer climates of I t a l y , but they proved unsatisfactory i n Central and Northern Europe where they grew slowly and suffered from winter f r o s t s . Provenances from east of the Cascades, p a r t i c u l a r l y those from the Rocky Mountains, always showed great s u s c e p t i b i l i t y to Rhabdocllne and exhibited poor growth (Schober 1963)• According to Schober (1959)t the caesla and glauca provenances are susceptible to Rhabdocllne because they are not used to the high a i r humidity of the European climates which does not bother the coastal form. In the tables attached to the 1963 p u b l i c a t i o n , Schober c l e a r l y showed that i n most Euro-pean countries, suitable Douglas-fir provenances outproduce a l l indigenous species. Many European foresters (e.g. Schober 195^, Rohmeder 1956) f e l t that i t was time to sample i n more d e t a i l the populations from the general areas where the provenances that have proven suitable f o r Europe, originated. This i s ex-a c t l y what some Danish foresters (Barner 1 9 6 6 ) had i n mind when they sent a team of cone pickers to western North America f o r several consecutive years, s t a r t i n g i n 1 9 6 6 . The seed i s d i s -tributed by I.U.F.R.O. to interested research groups. Many more experiments with Douglas-fir are going on i n Europe. Stern ( 1 9 6 6 ) f o r instance reported on three to four years' performance of so f a r "unexplored" origins from high elevation stands i n Arizona, New Mexico and Mexico, grown at Schmalenbeck. He pointed out t h e i r advantages of f a s t e r growth, longer growing period, greater f r o s t and drought resistance. The majority of European countries, including Luxem-bourg (Decker 1 9 6 7 ) and Russia "where the coastal form of Douglas-fir i s one of the most r a p i d l y growing, valuable and promising forestry species i n the western Ukraine" (Brodovich 1 9 6 7 ) t are interested i n growing suitable provenances of t h i s species, which some aut h o r i t i e s h a i l as Europe's most important exotic. Douglas-fir i s a l s o grown In other parts of the world, f o r instance i n New Zealand (Sweet 1 9 6 5 ) where i t i s doing ex-tremely well (Spurr 1 9 6 3 ) « In eastern North America, that i s , i n areas outside i t s natural range, Douglas-fir has mainly been grown f o r Christmas tree purposes. Byrnes et a l . ( 1 9 5 8 ) tested several provenances In Pennsylvania and found that the glauca v a r i e t y was best i n terms of s u r v i v a l , growth and hardiness. V i r i d i s , though s l i g h t l y f a s t e r In growth, suffered most from cold and drought. Likewise, Baldwin and Rock ( 1 9 6 1 ) rated glauca superior to v l r l d l s a f t e r nine growing seasons i n New Hampshire. Major Provenance Tests With Douglas-fir Within Its Range The most famous Douglas-fir provenance t e s t i n North America i s the c l a s s i c a l 1912 heredity study of the United States Forest Service. It consists of progenies from 120 re-corded mother trees, representing thirteen coastal provenances ranging e l e v a t i o n a l l y from 100 to 3 . 8 5 0 feet and planted at f i v e elevations between 1,1.00 and 1,400 feet i n western Oregon and Washington. I t i s both a provenance study and a progeny test from which a wealth of usef u l information has been derived. Munger and Morris (1936) found f o r instance that the age of the parent tree, the quality of i t s growing s i t e , i t s growing space and i t s condition as to fungus i n f e c t i o n had no e f f e c t upon the height growth of i t s progeny. Wright (1962), using data from that publication, c r i t i c i z e d the s t a t i s t i c a l design of the study f o r lack of r e p l i c a t i o n and randomization and concluded prema-ture l y that " ... i n a l l f i v e test areas one or more non-local provenances grew f a s t e r than the l o c a l provenance." The 1955 November f r o s t , k i l l i n g many non-dormant trees, l o c a l and non-l o c a l , was going to change t h i s picture d r a s t i c a l l y , leading S i l e n (1966) to stress the importance of s u r v i v a l and growth combined, instead of growth performance alone, when discussing provenance performance. S i l e n hypothesized that inherent growth rate of a race has developed toward the maximum that can be sustained i n each l o c a l i t y against impacts of long-term weather extremes. This hypothesis does not exclude the p o s s i b i l i t y of non-local races outproducing l o c a l ones at short rotations on protected s i t e s . Among other important findings, the following points are worth remembering (United States Forest Service 1962). 1. Seed o r i g i n was more c r i t i c a l i n high-elevation (above 2,000 feet) than i n low-elevatlon plant-ations. 2. Superior performance of a given provenance at one planting s i t e was no assurance of superior performance elsewhere. 3» Slenderness or stockiness of progeny was con-s i s t e n t l y related to i t s female parent l n a l l plantations. Mortality r e s u l t i n g from the severe freeze of November 1955 was also related to the i n d i v i d u a l parent. According to the United States Forest Service (1964), no r e s u l t was more s t r i k i n g than the evidence of a gene-environment i n t e r a c t i o n . Low elevation sources performed poorly at high elevations and vice-versa. It was also found that en-vironment generally has a larger e f f e c t on growth than source of seed. The pattern of bud burst d i d not appear to be a l t i -t u d i n a l . Wide v a l l e y origins burst t h e i r buds f i r s t , followed by those on open slopes. Those from narrow v a l l e y s flushed l a s t , probably representing a se l e c t i o n against l a t e f r o s t s . Sweet (1965) came to s i m i l a r conclusions l n New Zealand. Other authors have used data from the 1912 study. For instance Isaac (1949) established seven seed transfer rules or l i m i t a t i o n s r e l a t i v e to the c o l l e c t i o n of Douglas-fir seed. These r u l e s , l i s t e d on page 22, are based primarily on average annual temperature and f r o s t free period and roughly p a r a l l e l Langlet's (1945) system used l n Sweden f o r Scots pine. Jahn (1955) compared North American with German experience on Douglas-fir provenance. In 1954, foresters i n Oregon, r e a l i s i n g the lack of knowledge on the Douglas-fir provenance problem, planned a regionwlde study which comprised sixteen provenances from Oregon, Washington and B r i t i s h Columbia. This so-called "Co-operative Oregon Douglas-fir provenance study" w i l l be discussed i n d e t a i l l a t e r i n t h i s t h e s i s . Orr-Ewing (1966) reported on i n t r a s p e c i f i c crosses with Douglas-fir from various origins i n the United States and Canada. Some North American research i n s t i t u t e s are growing seed from the I.U.F.R.O. c o l l e c t i o n mentioned e a r l i e r . Schmidt (1967) reported on the c r i t i c a l stages of an intensive provenance study the major aim of which i s to set up seed transfer rules f o r the coastal range of Douglas-fir i n B r i t i s h Columbia. The area was s t r a t i f i e d ob-serving two c r i t e r i a : 1) recognition of known or suspected c l i m a t i c differences, 2) p o s s i b i l i t y that geographically separated areas might have produced d i f f e r e n t ecotypes despite only minor c l i m a t i c d i f f e r e n c e s . From a t o t a l of ninety provenances to be tested, sixty-nine are from the coast ( f i f t y - s e v e n from B r i t i s h Columbia, twelve from Washington and Oregon), th i r t e e n from continental climates of B r i t i s h Columbia^ I n t e r i o r , and eight from Coast-I n t e r i o r t r a n s i t i o n zones. Cones f o r that study were c o l l e c t e d In 1966, par t l y In connection with I.U.F.R.O. to avoid d u p l i c -ation of time and e f f o r t . F i e l d t e s t i n g w i l l be conducted at over f o r t y outplanting areas, d i s t r i b u t e d over a comprehensive range of climates within the coastal range of Douglas-fir. Meteorological data w i l l be obtained at the f i e l d test s i t e s . In 1967 Haddock et a l . published a paper comparing coastal with i n t e r i o r Douglas-fir o r i g i n s . The authors, who stressed the Importance of a thorough knowledge of the geography of an area and an understanding of the Influence of topography on l o c a l climate, confirmed European experience that coastal provenances outgrow I n t e r i o r sources i n mild climates, while being at the same time l e s s f r o s t r e s i s t a n t and less suscept-i b l e to Rhabdocllne i n f e c t i o n s . Once more, l o c a l or near-local or i g i n s were found to produce the best growth. Many more pub-l i c a t i o n s have appeared on Douglas-fir provenance i n North America. Morphological problems have been investigated f o r instance by A l l e n ( 1 9 6 0 a , 1 9 6 l ) who described an easy method of separating coastal from B r i t i s h Columbia i n t e r i o r sources. Tusko ( 1 9 6 3 ) concluded a f t e r an extensive study on the v a r i -a b i l i t y i n c e r t a i n Douglas-fir populations i n B r i t i s h Columbia, that there are only two subspecies - one coa s t a l , and one i n t e r -i o r . S z i k l a i ( 1 9 6 9 ) studied v a r i a t i o n i n cone and seed morph-ology of 1 , 3 3 5 trees from the I.U.P.R.O. c o l l e c t i o n between kZ°0? and 5 3 ° 3 7 l a t i t u d e north and found " ... a c l e a r l y ex-pressed c l i n a l v a r i a t i o n i n cone and seed length with an i n -creasing trend from north to south. The other c h a r a c t e r i s t i c s such as wing length, wing width and seed width, did not show a s i m i l a r c l i n a l v a r i a t i o n pattern." Growth chamber experiments under controlled l i g h t , were ca r r i e d out by Irgens--Miller ( 1 9 5 7 ) who found ecotypic response to temperature and photoperiod. High elevation plants appeared to have a d e f i n i t e photoperiodic response with regard to date of bud bursting. The magnitude of t h i s response was increased by low night temperatures. Irgens-Mjrfller concluded that the natural s e l e c t i o n to which Douglas-fir at high elev-ations has been exposed may have resulted i n plants f o r which the optimal length of the day f o r bud bursting occurs at a time when the danger of night f r o s t i s usually low. Vaartaja (1959) concluded that photoperiodic ecotypes have evolved as an Indiv-idual mechanism of trees to seasonally changing c l i m a t i c f a c t -ors. A l l e n ( 1 9 6 0 b ) separated coastal from i n t e r i o r seed l o t s on the basis of t h e i r germination behaviour at various temper-atures, following a short period of s t r a t i f i c a t i o n . Revel ( i 9 6 0 ) compared coastal with i n t e r i o r provenances grown i n a greenhouse, and found that germination was f a s t e r and height growth ceased much sooner f o r i n t e r i o r provenances which also require more c h i l l i n g to break dormancy fo r vegetative growth. S i m i l a r l y , Nicholson ( 1 9 6 3 ) found higher germination f o r i n t e r -i o r provenances. He separated i n t e r i o r from coastal provenances by using short day treatments. However no d e f i n i t e regional grouping of the coastal provenances was distinguishable. Sorensen ( 1 9 6 7 ) was able to separate two year-old seedlings representing several provenances from a west-east transect In Oregon into three d i s t i n c t groups, on the basis of t h e i r height growth and the date of bud formation In the f i r s t year. Haddock and Schmidt (1957) wrote that i t i s best to assume that there are a large number of unexplored ecotypes since the e c o l o g i c a l behaviour of Douglas-fir obviously varies greatly with s i t e , " ... even within the P a c i f i c Coast portion of i t s range." Bingham ( 1 9 6 6 ) f e l t that many cases of aberrant performance In " l o c a l " sources involve movement of seed between unrecognized, but nevertheless d i s t i n c t l y d i f f e r e n t environ-ments. He recommended adherence to the maxims l o c a l seed i s safest and probably best, and he cautioned against stretching the concept of " l o c a l , " e s p e c i a l l y where steep c l i m a t i c gradi-ents or s o l i changes are known or suspected between seed source and planting area. Haddock and S z l k l a l ( 1 9 6 6 ) established seed c o l l e c t i o n zones f o r Douglas-fir i n Canada, based primarily on c l i m a t i c data and the d i s t r i b u t i o n of associated species. The authors recommended that seed users who cannot indicate a spe-c i f i c l o c a l i t y or stand from which they want seed, should pro-vide pertinent c l i m a t i c data f o r the proposed plantation s i t e i n order to avoid undesirable r e s u l t s . Haddock ( 1 9 6 7 ) stressed the need f o r more precise provenance designations because of the great topographic and associated c l i m a t i c and edaphic v a r i a t i o n believed to have shaped the evolution and development of l o c -a l l y , g e n e t i c a l l y d i f f e r e n t populations over long periods of time. The author also warned against r e l y i n g too much on morph-o l o g i c a l features used by taxonomlsts " ... we should r e l y more on less e a s i l y measured c h a r a c t e r i s t i c s such as general physiol-ogy, phenology, cold, heat and drought resistance, s u s c e p t i b i l -i t y to insects and disease, growth form and wood q u a l i t y . " IV. THE CO-OPERATIVE DOUGLAS-FIR PROVENANCE TEST* General The great Importance of Douglas-fir In the regional economy and the lack of knowledge on the v a r i a t i o n within t h i s In t h i s thesis "co-operative t e s t " or "co-operative study" are used as abbreviations f o r "the co-operative Douglas-fir prov-enance t e s t . " species lead i n 1 9 5 ^ to a publication by the P a c i f i c Northwest Forest and Range Experiment Station, e n t i t l e d "A Program of Tree Improvement f o r the P a c i f i c Northwest," s t a t i n g that the tolerance of Northwest species to changes i n geography, elev-ation and climate, had not been f u l l y explored and a compre-hensive program of provenance t e s t i n g should be i n i t i a t e d as soon as possible. That same year, s t a f f of the Research Div-i s i o n of the Oregon State Board of Forestry (now i n the Forest Research Laboratory, Oregon State University) began to organize a regionwide provenance study of coastal Douglas-fir, a f t e r conferring with many interested parties (Anon. 1 9 5 5 & K Hypotheses to be Tested A f t e r reviewing l i t e r a t u r e on the Douglas-fir prov-enance problem, Ghing and Bever (19->0) f e l t that i n the experi-ments reviewed, the sampling of the variables of a l t i t u d e , l a t i t u d e and longitude was neither precise nor systematic enough to elucidate the question whether v a r i a t i o n i s c l i n a l or discontinuous. To improve on t h i s , an e f f o r t was made to get as many co-operators as possible i n order to test the following hypotheses (Anon. 1 9 5 5 a ) • 1 . D i s t i n c t races of Douglas-fir are associated with temperature as measured by f r o s t free days during the growing season. 2. D i s t i n c t races of Douglas-fir are associated with a l t i t u d e . 3 . D i s t i n c t races of Douglas-fir are associated with l a t i t u d e . 4. D i s t i n c t races of Douglas-fir are associated with temperature, a l t i t u d e and l a t i t u d e , as measured by comparable f r o s t free days and comparable photoperlods during the growing season. 5 . D i s t i n c t races of Douglas-fir have resulted from the development of genetic s t r a i n s i n l o c a l i z e d areas, and not from any of the variables or com-binations of variables stated i n the preceding four hypotheses. 6. D i s t i n c t races of Douglas-fir do not e x i s t within the area covered by t h i s study. Design The design was to follow the working plan f o r the southern pine provenance study by Wakeley ( 1 9 5 3 ) who had found at l e a s t one of the following six weaknesses i n other proven-ance studies 1 1 . Occasional aimless and i l l o g i c a l s e l e c t i o n of seed sources. This mistake was to be avoided through r e l a t i v e l y systematic sampling within the range of coastal Douglas-fir i n Oregon, Washington and B r i t i s h Columbia. 2. Mixing of single tree and bulk cone c o l l e c t i o n s . A r i g i d cone c o l l e c t i o n prospectus, asking f o r a sample of at le a s t f i f t y trees per seed source was aimed at overcoming t h i s shortcoming. 3 . Inadequate test plantings. This error was to be corrected by es t a b l i s h i n g plantations of a l l sources near each c o l l e c t i o n s i t e to compare the l o c a l provenance with the other provenances. 4. Inadequate s t a t i s t i c a l design. Four r e p l i c a t i o n s with random provenance a l l o c a t i o n was to s a t i s f y the prerequisites f o r s t a t i s t i c a l analysis (see F i g . 1 ) . 5 . I n s u f f i c i e n t number of trees planted f o r adequate s u r v i v a l and measurement up to ro t a t i o n age. An eight by eight foot spacing i n square plots with Since Wakeley's publication was not seen, the Information on "design" i s mainly taken from Heaman's (19^3) t h e s i s . eleven times eleven seedlings was to be adopted; the two outer rows were to act as a buffer s t r i p , minimizing edge e f f e c t s (see F i g . 2 ) , 6 . Lack of maintenance a f t e r establishment of the t e s t , and subsequent Irregular measurements. Good co-operation and the r a t i f i c a t i o n of a ten-year agreement by a l l co-operators was to remedy t h i s . In addition to the above points, confounding nursery e f f e c t s were to be eliminated by growing a l l stock i n one nursery, namely C o r v a l l l s . F i g . 1 Diagram of plo t designations. Block A=l Block B=2 Block C=3 Block D=4 13 8 14 7 4 10 1 13 14 6 1 10 2 11 14 8 6 1 4 12 9 15 5 2 8 2 16 4 16 6 4 12 15 5 11 16 7 8 l l 14 5 13 9 12 5 3 7 10 9 3 10 2 12 3 6 16 11 3 15 7 1 9 1 3 1 5 Plantation I Plantation II Each outplanting area has two "plantations," usually from one-quarter mile to one-half mile apart, and c o n s i s t i n g each of two adjacent blocks. Co-operators and Seed Sources Tables 1 and 2 show the names of the co-operators as well as the d i s t r i b u t i o n of the seed sources. Figure 3 i l l u s -trates the geographic d i s t r i b u t i o n of the provenances included i n the co-operative t e s t . The provenances were f i r s t assigned l e t t e r s (A to P), then given numbers following the order i n F i g . 2 Blow-up of plot X i n plantation Y. X X X X X X X X X X X X X X X X X X X X X X 1 2 3 4 5 6 7 X X X X X X X X X X X 14 13 12 11 10 9 8 X X X X X X X X X X X 1 5 16 17 18 19 20 21 X X X X X X X X X X X 28 2 ? 26 2 5 24 23 22 X X X X X X X X X X X 2 9 30 31 32 33 34 3 5 X X X X X X X X X X X 42 4 l 40 39 38 37 36 X X X X X X X X X X X »3 4 4 4 5 46 4 7 4 8 4 9 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Features of each small p l o t i - 121 trees - 8 x 8 ' spacing - a l l trees of the same provenance - only 4 9 trees i n the inner groups w i l l be measured. They are marked with numbered cedar stakes. Table 1 Co-operators and Location of the Test Sites Prove- Elevation Year of Seed nance L a t i - (Feet) range C o l l e c - Location of Source Number Co-operator tude of c o l l e c t i o n t i o n Test S i t e B r i t i s h A 4 Canadian Forest Products Ltd. 50°30 ! 400-600 1956 Nimpkish River Co lurabia V a l l e y B 2 Crown Zellerbach (Canada) Ltd. 49°45' 1300-1700 1956 Courtenay ; C 1 MacMillan Bloedel Limited 49°10' 2600-2900 56/57 Sugar Loaf Mountain, Chemainus River D 3 B. C. Forest Service 48°50' 570-750 1955 Robertson Valley near Lake Cowichan E 15 University of B r i t i s h Columbia 49°10' 500-700 1957 University of B r i -t i s h Columbia Re-search Forest Washington F 16 Weyerhaeuser Company 47°30' 39-4100 1957 Snoqualmie Area G 6 Simpson Olympic Tree Farm 47°15' 100-500 1956 Shelton Area H 5 Dept. Nat. Resources, State 46°45' 1850-2000 1956 Elbe Area of Wash. Oregon I 7 State Board of Forestry, 45°30' 1600-2200 54/55 Tillamook Area Oregon J 9 Crown Zellerbach Corporation 45°10' 1600-2000 1956 "1 Clackamas Tree Farm K 10 Crown Zellerbach Corporation 45° 10' 3200-3800 1956 J Mola11a Area L 8 Jack Stump and Kenneth McCrae 44°50' 200 1956 Willamette V a l l e y , Salem Area M 11 Oregon State College 44°30' 1800-2000 55/56 McDonald Forest, C o r v a l l i s Area N 12 U.S. Forest Service 43°45' 1800-2000 1954 Oakridge Area 0 13 U.S. Forest Service 43°45« 2500-3000 1956 High P r a i r i e , Oakridge Area P 14 Medford Corporation 42°20' 2700-3300 1955 Butte F a l l s Table 2 D i s t r i b u t i o n of the Seed Sources a) B r i t i s h Columbia Location Elevation (feet) 0-1000 1000-2000 2000 + N. Vancouver Island Nimpkish(A) C. Vancouver Is land Courtenay(B) S., Vancouver Is land Robertson Valley(D) Sugar loaf Mtn.(C) Mainland University of B r i t i s h Columbia Research Forest(E) b) U.S.A. Location Coast Zone Valley Zone Cascade Zone ~~0-1000 1000-2000 2000 + 1000-2000 ~2000-3000 3000 + Washington Shelton(G) Elbe(H) Snoqualmie(F) Oregon Tillamook(I) Willamette(L) M o l a l l a ( J ) Oakridge(O) Molalla(K) Corvallis(M) Butte F a l l s ( P ) Oakridge(N) which the seed l o t s reached the nursery (Table l ) . For example, provenances E and F were the l a s t ones to be sent to G o r v a l l i s and were thus assigned the numbers 15 and 1 6 . The following persons acted as regional co-ordinatorst Oregon i J.F. Gartz Washington « J.W. D u f f i e l d B r i t i s h Columbia i A.L. Orr-Ewing The f i r s t o v e r a l l co-ordlnator was D.N. Bever of the Oregon State Board of Forestry? Professor K.K. Chlng of the Oregon State University replaced him l a t e r on. Phases The study was separated into four d i s t i n c t phasesi Selection of outplanting areas and seed c o l l e c t i o n The s e l e c t i o n of the test s i t e s was l e f t to the i n d i v -idual co-operator with approval by the regional co-ordinator. The test s i t e s were to meet the following requirements! i ) be i n the Douglas-fir type i i ) north to northwest aspect i l l ) large enough to handle two six-acre plantations iv) be on land expected to remain i n permanent ownership v) not to be relogged or have a heavy brush or herbaceous cover. In areas where excessive deer browsing was expected, an eight foot fence was recommended. It was not always possible to s t r i c t l y meet a l l the requirements. Most good growing s i t e s suffer from a brush invasion. Also, mountainous s o i l s are extremely variable and therefore there may be undesirable s i t e heterogeneity within as small an area as s i x acres. The cone c o l l e c t i o n areas were defined as having a radius of twenty-five miles i n a designated area, at an elevation not d i f f e r i n g from that of the outplant-ing area by more than two hundred feet (or exceptionally up to four hundred f e e t ) . A maximum of f i f t y trees should be picked at random to sample as much v a r i a t i o n as possible, and to pro-vide 3 * 2 pounds of seed from each area (Anon. 1 9 5 5 t > ) » Seed crop permitting, the c o l l e c t i o n s were started i n the f a l l of 195^* The l a s t two l o t s of seed (from the University of B r i t i s h Columbia Research Forest, i . e . seed source 15. and Snoqualmle, i . e . seed source l 6 ) a r r i v e d at the nursery i n C o r v a l l l s i n f a l l 1957. The number of trees sampled ranged from fourteen to eighty-nine. Seed was extracted by various organisations. A f t e r an eighty-five percent c u t t i n g t e s t was obtained, the clean seed was stored i n a cold storage at zero degree Fahren-h e i t at the Oregon State Forest Nursery In C o r v a l l l s (Ching and Bever, i 9 6 0 ) . Nursery phase In order to avoid confounding nursery e f f e c t s , a l l seed was raised i n C o r v a l l l s . Naked s t r a t i f i c a t i o n was used by soaking the i n d i v i d u a l seed l o t s i n p l a s t i c bags with water for f orty-eight hours. A f t e r the water was drained o f f , the bags were placed i n a cool room at - 37 degrees Fahrenheit and 95 percent r e l a t i v e humidity, f o r three weeks (Ching and Bever, i 9 6 0 ) . The seeding rate was adjusted according to the v i a b i l i t y of the seed l o t . Sowing of fourteen seed l o t s was started on May 15 and completed on May 17, 1957* The f i r s t i n -d i c a t i o n of f i e l d germination was noticed on May 28. The seed-l i n g s were l i f t e d i n l a t e February 1959 ( i . e . as 2 - 0 stock) and c u l l e d according to c r i t e r i a advocated by Edwards ( 1 9 5 6 ) . A l l seedlings with l e s s than a four Inch top, were discarded. Due to a poor cone crop, seed l o t s E (from the University of B r i t -i s h Columbia Research Forest) and F (Snoqualmle) were shipped to the nursery one year a f t e r the other provenances, and sown i n the spring of 1958} consequently trees from these two origins were to remain one year younger than the others throughout the duration of the co-operative study. Planting of the nursery stock The o r i g i n a l plan s p e c i f i e d one planting crew i n each region (Oregon, Washington, B r i t i s h Columbia). This did not prove f e a s i b l e , and planting was c a r r i e d out by i n d i v i d u a l co-operators i n f a l l 1959 or spring i 9 6 0 . The test s i t e s had previously been adequately prepared. F i e l d examinations Co-operators were provided with thermometers, r a i n gauges and standardized f i e l d t a l l y sheets. Current and t o t a l height growth was to be assessed i n metric units and records kept on flushes, f r o s t and other damage. Results Ching ( 1 9 5 8 ) reported on f i r s t year performance of fourteen provenances (the University of B r i t i s h Columbia Re-search Forest seedlpt and the Snoqualmle source were sown one year l a t e r than the others) In the nursery at C o r v a l l l s . Height growth, bud burst and bud set, as well as f r o s t damage were assessed and genetic variations found. Seedlings from the Nlmpkish Valley, Vancouver Island, and from Butte F a l l s , Oregon, I.e. from both l a t i t u d i n a l extremes of the study, showed a def-i n i t e trend of i n f e r i o r growth. Some s i g n i f i c a n t differences In the s u s c e p t i b i l i t y to f r o s t damage were found, but caution was expressed to avoid premature conclusions. Ching and Bever ( i 9 6 0 ) reported on two years' nursery performance. They found no c o r r e l a t i o n between the height of the seedlings and the a l t i t u d e of t h e i r place of o r i g i n , a l -though they d i d f i n d s i g n i f i c a n t differences i n needle length between various provenances and some c o r r e l a t i o n between needle length and t o t a l height growth. Except f o r the Nlmpkish Valley provenance, the Vancouver Island sources compared favorably with the l o c a l Willamette Valley provenance, as f a r as height growth was concerned. Under the shorter photoperiod i n Oregon, the Vancouver Island sources, excepting the Nlmpkish one, formed t h e i r buds e a r l i e r and terminated t h e i r growth e a r l i e r than others. The f a c t that the southernmost provenance (Butte F a l l s ) did not show increased height growth under the s l i g h t l y longer photoperiod at C o r v a l l l s , was Interpreted to be a r e s u l t of natural s e l e c t i o n f o r early cessation of growth In an environment characterized by severe summer droughts. High elevation origins showed a tendency toward l a t e bud bursting, as an adaptation to l a t e spring f r o s t s . No c o r r e l a t i o n was found between time of bud burst and annual height growth. Ching ( i 9 6 0 ) reported on s u r v i v a l of fourteen prov-enances on eleven test s i t e s a f t e r the f i r s t growing season i n the f i e l d . Height was measured i n four plantations.* Except f o r the Nlmpkish Valley provenance, the Vancouver Island sources compared very favorably with a l l others, the Sugar-Loaf Mountain o r i g i n (C) being among the three best i n a l l four plantations. Walters and Soos ( 1 9 6 l b ) based t h e i r studies regarding lammas growth on the University of B r i t i s h Columbia Research Forest on Douglas-fir seedlings, Including the co-operative t e s t . Their findings, e.g. that lammas growth Is greatly i n f l u -enced by environment have been reviewed above. The same authors ( 1 9 6 l a ) investigated the e f f i c i e n c y of various chemicals to prevent hares from damaging the young Douglas-fir trees. The repellents used did not prove adequate; protection of the nat-u r a l predators of the hares was suggested as a better means of keeping the varying hare population i n check. Heaman ( 1 9 6 3 ) reported on the project i n B r i t i s h Columbia f o r the years 195^ - 1 9 6 1 . Mortality and height growth a f t e r the i 9 6 0 and the 1 9 6 l growing seasons were assessed f o r the f i v e outplanting areas In the province and weather data Unless otherwise defined, "plantation" Is synonymous with "test s i t e " or "outplanting area." were discussed. Heaman's c r i t i c i s m covered the following points t 1. Scope of the study. The scope of the co-operative study was too wide, covering eight degrees of l a t i t u d e . According to Schmidt's (1962a) c l a s s i f i c a t i o n , the co-operative t e s t f a l l s Into the category of "studies of broad geographic v a r i a t i o n . " This type of study cannot be expected to y i e l d more than broad generaliz-ations. Information gained from such a project can be used to design a more Intensive one i n the future, but as the r e s u l t s cannot be r e l i a b l y interpreted f o r at l e a s t twenty years, t h i s Is a lengthy approach. P r a c t i c a l a p p l i c a t i o n s , such as seed transfer r u l e s , cannot be expected. 2. Cone c o l l e c t i o n . The cone c o l l e c t i o n was not c a r r i e d out i n a uniform way by a l l co-operators. Apart from t h i s , the twenty-f i v e mile c o l l e c t i o n radius was too large, despite the l i m i t a t i o n of c o l l e c t i n g within four hundred, pre-ferably within two hundred feet of the elevation of the t e s t i n g s i t e . Heaman supported his c r i t i c i s m with records from f i v e weather stations l y i n g within those l i m i t a t i o n s i n a twenty-five mile radius of the Robertson Valley. The average f r o s t free period, the major c l i m a t i c c r i t e r i o n of the co-operative study, between two extreme stations, varied from f i f t y - o n e to two hundred and fo r t y - f o u r days! 3« Weather records. Heaman deplored the lack of continuity and stand-a r d i z a t i o n . Some co-operators measured temper-atures at two feet, others at four, others s t i l l at f i v e feet above the ground. At the Nlmpkish test s i t e i n i 9 6 0 , a f r o s t free period of eleven weeks was recorded at two feet above the ground and twenty-two weeks at f i v e feet above the ground. 4 . Site s e l e c t i o n . Heaman f e l t that more care could have been exercised i n s i t e s e l e c t i o n . For instance, i t might have been recognized from the s t a r t that the Robertson Valley plantation was located i n a f r o s t pocket. A f r o s t free period of only s i x weeks was measured at two feet above the ground i n i 9 6 0 . Heaman found no c o r r e l a t i o n between height growth and elevation or l a t i t u d e of seed source. There was no c o r r e l a t i o n between eithe r seed weight and height growth or between germin-ation percentage and seed weight during the f i r s t year. At Nlmpkish and Courtenay, the low elevation source G (Shelton) and the high elevation source C (Sugar-Loaf Mount-ain) showed best performance, although they did not prove s i g -n i f i c a n t l y superior on a l l B r i t i s h Columbia test s i t e s . * The Heaman did not test the provenance x block i n t e r a c t i o n , although he pointed to the s i t e heterogeneity at the University of B r i t i s h Columbia Research Forest. unusual early growth of Shelton (G), I f endorsed by more r e l i -able data i n the future, was interpreted as a possible con-firmation of the hypothesis that " ... d i s t i n c t races of Douglas-fir have resulted from the development of genetic str a i n s i n l o c a l i z e d areas ... " Heaman concluded that s i g -n i f i c a n t deductions could not be made because of the short duration of the experiment, the incompleteness and lack of standardization of weather data, and because of the excessive damage at some test s i t e s by f r o s t and browsing. The following were Heaman's recommendationst 1. Provenance studies should be of a smaller scope than the co-operative t e s t , and sample the range of a species much more intensively within much narrower l a t i t u d i n a l l i m i t s . 2. Standardization and controls i n a l l phases of a co-operative study by one co-ordinator i n constant personal contact with a l l In-volved, are e s s e n t i a l . 3 . Cone c o l l e c t i o n areas have to be c l e a r l y defined and based on a thorough l o c a l know-ledge • 4. Growth studies should be based on phenological observations Instead of tedious weather measurements. 5. A comprehensive study of a l l outplanting areas i s necessary regarding climate, s o i l and micro-topography. 6 . The plantations have to be protected e f f i c -i e n t l y from animal damage, i f early assess-ments are to have any meaning, unless i t can be established that the damage i s evenly d i s t r i b u t e d over a l l sources. Ching ( 1 9 6 5 ) assessed s u r v i v a l and growth a f t e r the The Robertson Valley plantation was seriously damaged by f r o s t . f i r s t three years i n the f i e l d f o r fourteen provenances at eight outplanting areas. Early s u r v i v a l was highest near Nlmpkish. At most plantations, trees from the l o c a l source grew as well as those from the three best sources. Trees from seed c o l l e c t e d at the northern and southern extremes (Nlmpkish and Butte F a l l s , respectively) grew l e a s t i n the three years (as they had i n the nursery). Never damaged trees were no more than s l i g h t l y d i f f e r e n t i n height growth from those that had been damaged. Christmas tree growers systematically evaluated trees In three locations i n 1964 (Douglass 1967)* The Salem provenance (L) was found to be the best Oregon source and Shelton (G) the best representative from Washington. The Rob-ertson Valley (D) o r i g i n ranked highest i n a l l three locations investigated. It combined a number of desirable q u a l i t i e s , such as high vigor, dark green colour, upright growth habit, natural tendency for bushlness, and an a t t r a c t i v e needle arrangement. Walters and Ching (1969) studied the pattern of bud burst of the sixteen provenances* at the University of B r i t i s h Columbia Research Forest and i n the Willamette Valley. The re-sults have been reviewed on page 12. In a report on f i e l d per-formance at age nine, Ching (1967) assessed s u r v i v a l and height growth i n ten loc a t i o n s . Trees from Oakridge (N) and from the * Sources E (University of B r i t i s h Columbia Research Forest) and F (Snoqualmle) were included f o r the f i r s t time. Since t h e i r establishment i n 1959. the following plantations southernmost source P (Butte P a l l s ) were consistently poor In height growth. Ching concluded that high elevation stock can safely be planted at lower a l t i t u d e s , whereas i t would be un-wise to do the opposite. P l o t t i n g height growth against elev-atio n and l a t i t u d e d i d not reveal a s i g n i f i c a n t c o r r e l a t i o n . F i n a l l y , another Investigation (Mitchell and Nagel, 1 9 6 9 ) revealed that attack by Douglas-fir needle midges (mainly Contarlnla pseudotsugae Condrashoff) was correlated to date of bud burst, the damage being highest on early f l u s h i n g trees. V. METHODS The Study Area General de s c r i p t i o n of the University of B r i t i s h Columbia Research Forest The University of B r i t i s h Columbia Research Forest, at k9°l8* l a t i t u d e north and l 2 2 0 3 5 f longitude west, i s s i t u -ated four miles north of Maple Ridge, formerly Haney, i n the Fraser Valley of B r i t i s h Columbia, on the south fringe of the Coast Mountains, at a distance of 36 miles from the University have been l o s t i 1 . D - Robertson Valley, both plantations abandoned due to f r o s t damage. 2 . G - Shelton. One plantation l o s t to grass f i r e . 3 . I - Tillamook. Both plantations heavily damaged by r a b b i t s . 4. J - Molalla. Both plantations suffered from rabbit damage. 5 . N - Oakridge area. Both plantations l o s t to deer damage. 6 . P - Butte F a l l s . Both plantations l o s t to drought. (Ching 1 9 6 9 . personal communication.) of B r i t i s h Columbia Campus i n Vancouver (U.B.C. Forest Com-mittee 1959)* It comprises approximately twelve thousand seven hundred acres of forested land, and i s bounded on the north and east by Garibaldi P r o v i n c i a l Park and by P i t t Lake on the north-west. Elevations range from sea l e v e l to 2600 f e e t . The area l i e s within the southern P a c i f i c coast section (C . 2 ) of the coastal forest according to Rowe (1959) and i n the coastal western hemlock (Tsuga heterophylla (Raf.) Sarg.) zone according to Krajlna ( 1 9 5 9 )• The main association comprises the two coastal dominants western red cedar (Thuja p l l c a t a Donn.) and western hemlock, with coastal Douglas-fir and scattered western white pine, P a c i f i c s i l v e r f i r (Abies amabllls (Dougl.) Forbes), Sitka spruce (Plcea sltohensls (Bong.) Carr.) and yellow cedar (Chamaecyparls nootkatensls (D. Don) Spach) (Walters and Soos, I 9 6 l b ) . Climate The climate, described i n d e t a i l by G r i f f i t h ( 1 9 6 8 ) who used data from the four weather stations on the Research Forest, Is considerably influenced by the r e l a t i v e l y warm and moist a i r of the P a c i f i c Ocean and by the Coast Mountains. The summers are warm and dry, the winters r e l a t i v e l y mild and wet. Temperature seldom reaches 90 degrees Fahrenheit, the absolute minimum on record being f i v e degrees Fahrenheit below zero. The annual p r e c i p i t a t i o n averages about 8 9 inches i n the southern part of the Forest, where the provenance te s t s i t e i s located, but increases i n the higher elevation areas to the north. The high amount of p r e c i p i t a t i o n has a pronounced e f f e c t on the s o i l s which are strongly leached podsols and are very a c i d i c . The weather st a t i o n outside the Administration B u i l d -ing at an elevation of 475 feet above sea l e v e l i s the one closest to the study area, the climate of which can be assumed to be p r a c t i c a l l y the same. For the years 1 9 5 9 to 1967 i n -cl u s i v e , i . e . the years relevant to th i s t h esis, the following data were published by G r i f f i t h (1968)t P r e c i p i t a t i o n ! Annual average October to March A p r i l to September July ( d r i e s t month) December (wettest month) Winter snowfall Number of days with measurable p r e c i p i t a t i o n Temperaturei Annual mean July (hottest month) January (coldest month) Absolute minimum Absolute maximum Frost free period, average 200 days (Min. 1 6 5 - Max. 243) Average date of l a s t spring f r o s t 1 A p r i l l 4 Average date of f i r s t f a l l f r o s t i November 1 May f r o s t s are f a i r l y frequent but September fro s t s are rare. For the five-year period 1953-1957, average length of the growing season, as indicated by camblal a c t i v i t y r e s u l t -ing In diameter increment, was 146 days (ranging from a minimum of 107 days to a maximum of 1 6 3 days) according to G r i f f i t h 89 . 0 9 " (Min. 75.88" - Max. 95.83") 63.57"or 71.35$ of the annual total 2 5 . 5 2"or 28.65# 2.70"or 2.97$ l2.64"or 14.19$ * 24.4" 192 4 9 . 1°F (Min. 4 7 . 6 degrees - Max. 6 3.2OF 5 1 . 0 degrees Fahrenheit) 3 5 . 9°F 2 OF 98°F ( i 9 6 0 ) . Since the co-operative provenance te s t s i t e s , at an elevation of approximately 500 feet above sea l e v e l , are at a s l i g h t l y lower elevation than the weather s t a t i o n used by G r i f f i t h i n his i 9 6 0 study, the growing season there can be assumed to be of s l i g h t l y longer duration. Location of the co-operative provenance test on the Research Forest Plantation I ( i . e . blocks 1 and 2) i s situated be-tween Mainroad F and Spur A - 1 0 ; plantation II ( i . e . blocks 3 and 4 ) i s adjacent to Branch Road A on the eastern side of the North Alouette River. Both plantations are within l e s s than one-half mile of each other and l e s s than a mile from the Administration Building i n the southeast corner of the Research Forest (see F i g . 5 ) » History of the test s i t e P r i o r to logging i n 1 9 5 5 - 1 9 5 6 , the area supported the following volumes ( i n thousand board feet, B.C. r u l e ) * Live Dead Douglas Douglas Live Dead W.Hem- Vol / Cedar - f i r - f i r Cedar Cedar lock Total acre poles P l t n I* 15 9 95 4 132 2 5 5 18 . 4 P l t n II 260 - 4 3 5 69 208 972 4 3 . 2 6 9 The area of plantation I was logged by high-lead which caused r e l a t i v e l y l i t t l e s o i l disturbance, the area of * 1 Area was part of a 14 acre stand. Area was part of a 22 acre stand. plantation I I was yarded by t r a c t o r , requiring an intensive network of skldroads r e s u l t i n g i n heavy s o i l disturbance. The slash, burning poorly i n a f i r s t attempt, p a r t i c u l a r l y In plan-t a t i o n I , was p i l e d and re-burnt i n 1958 (Heaman 1 9 6 3 ) . Since the material was c o l l e c t e d by hand and no heavy equipment was used f o r s i t e preparation, there was minimal mechanical s o l i disturbance. Much debris remained. Vine maple (Acer c i r c l n -atum Pursh.) was successfully sprayed i n spring 1959 with 2,4-D, 2 , 4 , 5-T. Draining was ca r r i e d out i n portions of plantation I I by b l a s t i n g a d i t c h . The seedlings were planted i n the spring of 1959« The nursery planting stock ( 2 - 0 Douglas-fir) was generally of poor q u a l i t y and despite close supervision of the planting operation, examination of A r m l l l a r l a mellea (Fr) Kumm. infected root systems several years l a t e r suggested that the presence of the disease could possibly be related to a poor planting tech-nique (Walters 1 9 6 4 ) . The area was covered by fireweed (Eplloblum augustIfollum L.) six feet t a l l , but the r e a l problem the f i r s t year was browsing by hares. In addition to a trap l i n e , chemical repellents were used In May 1959» but they d i d not prove adequate If used alone (Walters and Soos, 1 9 6 l a ) . By 1 9 6 l It appeared that almost a l l of the seedlings had grown away from the browsing zone f o r hares, therefore no add i t i o n a l treatments were planned (Walters 1 9 6 1 ) . Mortality was f a i r l y highi 3 6 5 trees were found dead i n f a l l 1959. 380 died i n i 9 6 0 , and 140 i n 1 9 6 1 . These figures apply only to the "Inner blocks" with forty-nine numbered stakes (see under "design"). A l l mortality was replaced i n the spring of I960 and 1961 by seedlings that were themselves frequently of poor q u a l i t y . In 1963 the area was weeded by machete and brush hooks, and then concentrations of vine maple and salmon-berry (Rubus spe o t a b l l l s Pursh.) sprayed with 2,4-D, and 2,4,5-T i n d i e s e l o i l (Walters 1964). Vacolnlum spec, was never treated but possibly impeded tree growth wherever i t occurred In thick concentrations (see below). The four blocks were again treated chemically i n August 1965* Infections by A r m l l l a r l a continued during that year (Walters 1966). In 1966 the sixteen provenances produced flowers and cones f o r the f i r s t time (Walters 1967). Height growth had been assessed at the end of I960, 1961. 1964 and 1965. Height Measurements For t h i s t h e s i s , the height of a l l surviving numbered trees was measured a f t e r the 19.67 growing season when most trees were eleven years old. Since trees of the l o c a l source (Univ-e r s i t y of B r i t i s h Columbia Research Forest) and from Snoqualmle (F) were one year younger than the r e s t . I t seemed reasonable to measure them one year l a t e r , although no two growing seasons are quite the same. The primary objective i n analyzing the height measurements taken at the University of B r i t i s h Columbia Research Forest, was to determine whether s i g n i f i c a n t growth differences between provenances were showing at an early age, and I f so, to t r y and correlate them with geographic v a r i a b l e s . A secondary objective was to evaluate the s i t e homogeneity at the Research Forest. S t a t i s t i c a l Analysis Height measurements f o r simple analysis of variance were analyzed on computers at the University of B r i t i s h Colum-b i a . VI. RESULTS AND DISCUSSION Height at Age Eleven The analysis of variance of height measurements showed the block x provenance Interaction to be s i g n i f i c a n t . This means thatt a) The i n d i v i d u a l provenances behave d i f f e r e n t l y In the four blocks. b) The plantation s i t e Is not homogeneous. c) The four blocks have to be evaluated separately. d) No s t a t i s t i c a l conclusion as to the o v e r a l l per-formance, such as superior growth, of any par-t i c u l a r provenance, can be drawn, i . e . the provenances cannot be ranked on a s t a t i s t i c a l l y sound ba s i s . e) Unless the prerequisites f o r co-variance analysis can be met, which would allow comparisons of o v e r a l l performance of a l l provenances, the re-s u l t s at the University of B r i t i s h Columbia Research Forest do not j u s t i f y at t h i s stage a confirmation or r e j e c t i o n of any hypothesis that the co-operative study set out to t e s t , nor do the r e s u l t s j u s t i f y a p o s i t i v e answer to the question whether s i g n i f i c a n t growth differences between provenances are showing at an early stage. Sim-i l a r l y , seed transfer rules cannot be established with these r e s u l t s . An attempt was therefore made to f i n d a quantifiable parameter related to s i t e , that would allow an accurate explan-atio n of the d i f f e r e n t behaviour of the Individual provenances In the four blocks, a f t e r bringing everything down to the same bas i s . G r i f f i t h (i960) studied the growth of Douglas-fir i n r e l a t i o n to climate and s o i l at the Research Forest and found that a v a i l a b l e s o i l moisture i n the B-horizon during the grow-ing season was the most Important single variable a f f e c t i n g tree growth. The o r e t i c a l l y i t i s easy to get a v a l i d Idea of the s o i l moisture regime of an area by breaking i t down into i t s major i n d i c a t o r s i slope (percent), s o i l depth, p o s i t i o n on slope, aspect. It would indeed be h e l p f u l to know what per-centage of each plo t suffers from adverse conditions, such as excessive moisture or abnormal s o i l disturbance, f o r instance. By weighing the performance of each plot accordingly, that i s , by t r y i n g to eliminate varying environmental Influences, a better picture of the growth pote n t i a l of that p a r t i c u l a r prov-enance could be arri v e d a t . The following map was drawn by Mr. D. Ormerod, In an attempt to i l l u s t r a t e the s o i l moisture regime i n the four blocks (see F i g . 4 ) . Despite the help of the map, i t proved d i f f i c u l t i n the f i e l d to get an accurate estimate of s o l i moisture, due to the extreme v a r i a b i l i t y In microtopography even within the various p l o t s . It was f e l t that such an evaluation would too e a s i l y be subjective and therefore incorrect and inadequate. Mr. L. L a c e l l e , a graduate student In s o i l science * Plot means any provenance i n any block. ?LANTA T/DM T BLOCK 1 BLOCK JL * LPiCSLLE's SOIL PIT PlAA/r/ff/o// Jl BLOCK "I 8*. ock iy t UILL/NQ rows SOIL PIT APPK.OX. SCALZ-- /"* 3 CM/AJS LECf€AJl) V see/>A<5_ StJftMP 2>^/e>e Hirer V V v •exposed <5*Avei SLl<fH-T RiHS i -6 i 3*us/f Figure 4 Ormerod's map at the University of B r i t i s h Columbia, kindly dug several s o i l p i t s i n both plantations and came to the conclusion that the s o i l was roughly the same i n a l l four blocks, except f o r small areas that showed the e f f e c t s of mechanical disturbance due to logging, and also excepting areas with a d i f f e r e n t water regime due to t h e i r microtopography. Mr. Lacelle described the s o i l i n block B, plantation I, as a mlni-orthic-humoferric podzol. moderately well drained to well drained. According to W i l l i n g -ton ( 1 9 6 8 ) , the s o i l i n block C, plantation I I , i s a moderately well drained orthic podzol, developed i n outwash parent mater-i a l overlying Whatcom glaclomarine. Walters ( 1 9 7 0 ) thinks that the cemented layer underlying the s o i l at varying depths, has to take part of the blame for the v a r i a b i l i t y i n s i t e i n some plots by a f f e c t i n g the d i r e c t i o n of the seepage flow. A co-variance analysis therefore could not be carr i e d out due to the lack of a det a i l e d analysis of the physical, chemical and hydrologlcal s o i l properties of the d i f f e r e n t plots which would each require a great number of samples due to the extreme v a r i a b i l i t y i n microtopography. It i s hoped that such a study w i l l be made and more conclusive information w i l l be gained from the provenance study at the Research Forest. According to a personal communication by Ching ( 1 9 7 0 ) , a grad-uate student of Oregon State University, c o l l e c t e d many s o i l samples from eleven outplanting areas. His r e s u l t s w i l l prob-ably throw some l i g h t on the e f f e c t of microedaphic differences on tree growth. Heaman ( 1 9 6 3 ) pointed out the s i t e v a r i a b i l i t y at the University of B r i t i s h Columbia Research Forest, as exemplified by a gulley i n plantation I, c a t e r p i l l a r roads, a gravel p i t , and swampy areas i n plantation I I . However i t should be noted that, given the li m i t e d area and the knowledge at the time of the s o i l s , the s i t e chosen was by f a r the best one a v a i l a b l e . In another Douglas-fir provenance study at the Research Forest, Haddock et a l . ( 1 9 6 7 ) found a s i g n i f i c a n t provenance x block i n t e r a c t i o n , too, r e f l e c t i n g possibly " ... the heterogeneous nature of the s i t e , t y p i c a l f o r coastal mountain country i n B r i t i s h Columbia." Results by Individual Blocks Separate analyses of variance f o r height performance were carr i e d out. The re s u l t s are l i s t e d i n Table 3« The wide range, s t r i k i n g l y exemplified by three prov-enances (Nlmpkish, Tillamook and higher elevation Molalla), showing a difference of thirteen ranks, out of a possible maxi-mum of f i f t e e n , between t h e i r shortest and t h e i r t a l l e s t per-formance, can be Interpreted to mean either thatt 1. The genetic v a r i a b i l i t y of Douglas-fir i s such that four r e p l i c a t i o n s of 4 9 trees each f o r every prov-enance are not enough to cover i t . To support t h i s one could point to the fac t that, when walking through the study area at the Research Forest, one i s struck not only by the v a r i a b i l i t y i n growth between some provenances (e.g. seed source 4 , Nlmpkish, and 1 6 , Snoqualmle i n block III) but also by the fac t that i n every p l o t , no matter how poor Its average performance, there i s at l e a s t one tree doing much better than the r e s t . The best example of t h i s can be found i n block IV, plot 4, Nlmpkish, where one p a r t i c u l a r Ranking of Provenances  Block Proven- Aver-ance # Origin I II III IV age Range 1=G Sugar Loaf Mountain, Vancouver Island 6 3 1 8 ^.5 7 2=B Courtenay area, Vane. Island Robertson Valley, " . "' 8 2 7 12 7.2 10 3=D 3 8 13 9 8.2 10 4=A Nlmpkish Valley, " 10 11 2 1 5 9.5 13 5=H Elbe area, Washington 12 7 8 5 8.0 7 6=G Shelton area, " 4 1 3 4 3.0 3 7=1 Tillamook area, Oregon Salem area, " 5 6 16 3 7.5 1 3 8=L 11 13 12 6 10.5 7 9=J Molalla, lower " elevation 13 5 11 10 9.8 8 10=K Molalla, higher " elevation 2 15 1 5 2 8.5 13 11=M C o r v a l l l s area, " 9 9 9 10.0 4 12=N Oakridge, lower " elevation 14 16 4 14 12.0 12 13=0 Oakridge, higher " elevation 16 4 14 7 8.2 12 14=p Butte F a l l s , " 15 12 10 16 13.2 6* 15=E U.B.C. Research Forest, B r i t i s h Columbia 1 10 5 1 4.2 9 l 6=F Snoqualmle area, Washington 7 14 6 11 9.5 8 Table 3 Height performance at age eleven. Ranking i s from t a l l e s t ( l ) to shortest ( l 6 ) . tree i s growing vigorously i n a swampy place, with most of i t s neighbours looking c h l o r o t i c and stunted i n apparently the same micro-environment. This Is an Interesting p a r a l l e l to a fi n d i n g In the 1912 Douglas-fir heredity study by the U.S. Forest Serv-ice (1964)i "An anomaly to the i n t e r a c t i o n exists In the following sense i Almost without exception the progenies of every parent include at l e a s t one superior i n d i v i d u a l at each The l o c a l provenance. l o c a t i o n . This indicates s u r p r i s i n g genetic d i v e r s i t y within a r e s t r i c t e d source of germ-plasm such as a single seed l o t . " The wide range i n ranking could also mean that« 2 . This s i t e i s extremely heterogeneous. This a l t e r n -ative i s supported by the si g n i f i c a n c e of the i n -t e r a c t i o n term and i l l u s t r a t e d f o r instance by the above-cited source 4 (Nlmpkish). It i s second t a l l e s t i n block III where Its outstanding perform-ance can be detected from a 1 i 6300 scale a e r i a l photograph. In block IV It i s the t h i r d poorest i n terms of height growth, probably because i t has to grow i n a rather swampy place. A e r i a l photographs of the study area show evidence In favour of the " s i t e heterogeneity" - rather than the "gen-e t i c v a r i a b i l i t y " a l t e r n a t i v e . Patches of poor growth (marked with "P" on Figure 5) show up very w e l l . They ignore plo t boundaries and quite l i k e l y r e f l e c t edaphic d i f f e r e n c e s . Tables 4 to 7 show height performance within the i n d i v i d u a l blocks f o r 1 9 6 7 . 1964 and 1961 are included f o r comparison. The brackets show the r e s u l t s of Duncan's New Multiple Range Test. Early Tests Trying to f i n d c o rrelations between the i 9 6 0 , 1964 and 1967 measurements (at age 4 , 8 and 11 years) i s almost an exercise In f u t i l i t y f o r the following reasons 1 1 . Duncan's New Multiple Range Test shows that / ' / / Figure 5 Location map of the co-operative provenance test at the University of B r i t i s h Columbia Research Forest. % of % of % of Prov. Ht(cm) Mean Prov. Ht(m) Mean Prov. Ht(ra) Mean Rank Rank 1 10 49.27 139 15 2.5980 125 15 5.7333 124 1 2 2 39.71 112 10 2.5927 119 10 5.2458 114 2 3 1 38.92 110 3 2.3959 110 3 5.1694 112 3 4 15 38.02 108 6 2.3857 109 6 5.1163 111 4 5 3 37.65 106 _ 1 2.3653 109 7 4.7766 103 5 6 6 37.57 106 7 2.2687 104 1 4.7633 103 6 7 11 36.47 103 2 2.2330 103 16 4.6405 100 7 8 14 35.67 101 11 2.1787 100 2 4.6045 100 8 9 16 34.59 98 16 2.1511 99 11 4.5410 98 9 10 7 34.31 97 4 2.1142 97 4 4.5349 98 10 11 5 33.43 95 8 2.0777 96 8 4.4957 97 11 12 9 33.31 94 5 2.0719 95 5 4.4043 95 12 13 4 32.84 93 9 1.9826 91 9 4.2523 92 13 14 12 32.14 91 14 1.8511 85 12 3.8682 84 14 15 8 31.49 89 [12 1.7789 82 14 3.7109 80 15 16 13 20.65 58 13 1.5447 71 13 3.6081 78 16 Table 4 Height performance. Block I Any two provenances covered by the same bracket are not s i g n i f i c a n t l y d i f f e r e n t . OS Rank 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Prov. Ht(cm) % of Mean 6 47.76 132 5 43.94 121 1 43.81 121 2 41.71 115 9 40.39 112 7 39.96 110 13 39.89 110 11 37.33 103 14 35.47 98 3 34.85 96 10 31.35 87 4 30.33 84 12 29.12 80 8 28.53 79 15 27.39 76 16 26.58 73 Prov. Ht(ra) 7o of Mean Prov. Ht(m) 6 2.7354 127 " 2 2.4056 112 9 2.3833 111 1 2.3830 111 7 2.3605 110 5 2.3372 109 13 2.3262 108 14 2.1228 97 11 2.0292 94 3 2.0273 94 10 1.9939 93 8 1.9622 91 4 1.9221 89 15 1.9156 89 16 1.8323 85 12 1.6664 77 % of Mean Rank 6 5.2543 116 1 2 5.0133 110 2 1 4.9556 109 3 13 4.8581 107 4 9 4.8045 106 5 7 4.7905 105 6 5 4.6804 103 7 3 4.5974 101 8 11 4.4109 97 9 15 4.4043 97 10 4 4.3667 96 11 14 4.3200 95 12 8 4.3111 95 13 16 4.0958 90 14 10 4.0673 89 15 12 3.8930 86 16 Table 5 Height performance. Block II Any two provenances covered by the same bracket are not s i g n i f i c a n t l y d i f f e r e n t . 1 9 6 0 1 9 6 4 1 9 6 7 Prov. Ht(cm) 7o of Mean Prov. Ht(m) 7» of Mean Prov. Ht(m) 7o of Mean Rank 1 6 4 0 . 6 1 1 3 3 1 2 . 4 7 8 1 1 3 4 f l 5 . 1 3 7 5 1 3 2 1 2 4 3 8 . 3 5 1 2 5 4 2 . 3 5 5 3 1 2 8 r-L4 5 . 0 0 0 0 1 2 8 2 3 1 3 6 . 7 1 1 2 0 _ 6 2 . 3 1 7 8 1 2 6 6 4 . 3 2 3 8 1 1 1 3 4 2 3 4 . 4 7 1 1 5 1 1 1 . 9 0 8 7 1 0 3 1 2 4 . 0 8 0 4 1 0 5 4 5 5 3 4 . 1 0 1 1 1 1 4 1 . 8 9 6 7 1 0 3 1 5 4 . 0 3 9 0 1 0 3 5 6 1 1 3 1 . 8 1 1 0 4 2 1 . 8 8 9 0 1 0 2 1 6 3 . 9 7 8 6 1 0 2 6 7 — 7 3 0 . 8 6 1 0 1 12 1 . 8 6 7 0 1 0 1 _ 2 3 . 8 3 1 7 9 8 7 8 3 3 0 . 1 0 9 8 5 1 . 8 0 6 2 9 8 5 3 . 8 0 6 2 9 8 8 9 1 4 2 9 . 9 6 9 8 . 8 1 . 7 5 1 2 9 5 1 1 3 . 7 1 1 1 9 5 9 1 0 12 2 9 . 9 6 9 8 1 5 1 . 7 3 5 4 9 4 1 4 3 . 6 7 9 5 9 4 1 0 1 1 9 2 9 . 2 9 9 6 1 6 1 . 7 1 9 8 9 3 9 3 . 6 5 6 5 9 4 1 1 1 2 8 2 6 . 1 7 8 5 9 1 . 6 7 9 6 9 1 8 3 . 6 3 2 5 9 3 1 2 1 3 1 3 2 5 . 9 0 8 5 3 1 . 6 4 5 9 8 9 3 3 . 5 9 8 0 9 2 1 3 1 4 _ 1 6 2 4 . 4 2 8 0 7 1 . 5 8 4 4 8 6 1 3 3 . 2 9 0 7 8 4 1 4 1 5 1 5 2 3 . 7 7 7 8 1 3 1 . 4 8 4 4 8 0 1 0 3 . 2 6 7 5 8 4 1 5 1 6 1 0 2 1 . 7 8 7 1 1 0 1 . 3 4 0 0 7 3 7 3 . 2 6 4 6 8 4 1 6 Table 6 Height performance. Block III Any two provenances covered by the same bracket are not s i g n i f i c a n t l y d i f f e r e n t . Prov. Ht(cra) % of Mean Prov. Rank 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 10 7 15 5 ' 1 3 6 2 9 13 "16 4 12 8 11 14 43.88 41.00 40.67 38.22 33.30 32.86 32.50 31.94 31.39 30.06 27.32 26.66 26.47 25.79 24.67 22.69 137 128 127 120 104 103 102 100 98 94 85 83 83 81 77 71 .[Do 7 5 8 6 1 13 3 9 11 2 16 12 4 14 X of 7c of Ht(m) Mean Prov. Ht(m) Mean Rank 2.7622 137 15 5.9204 140 1 2.6435 131 rio 5.1404 122 2 2.4098 120 L 7 4.9128 117 3 2.2940 114 6 4.5889 109 4 2.1596 107 5 4.5755 109 5 2.1000 104 _ 8 4.4553 106 6 2.0600 102 13 4.3683 104 7 2.0284 101 1 4.0911 97 8 1.8378 91 3 4.0347 96 9 1.7937 89 9 3.8612 92 10 1.7735 88 16 3.8372 91 11 1.7232 86 2 3.7275 88 12 1.7170 85 11 3.6163 86 13 1.6625 83 12 3.6159 86 14 1.5133 75 r 4 3.2441 77 15 1.4685 73 14 2.9545 70 16 Table 7 Height performance. Block IV Any two provenances covered by the same bracket are not s i g n i f i c a n t l y d i f f e r e n t . -N3 the provenances do not respond s i m i l a r l y i n the four blocks. 2. Three observations of such v a r i a b i l i t y are not enough f o r s i g n i f i c a n t regression equations. Assuming these observations to be on a straight l i n e i s making a very rough approximation. It was therefore not su r p r i s i n g that multiple re-gression analysis did not lead to any spectacular findings. Although the trend i n height growth within the same block seems to be comparable, the slopes are s i m i l a r , the heights them-selves are not, except within very small groups of two or three provenances, since the intercepts are s i g n i f i c a n t l y d i f f e r e n t . Even i f the s i t e were homogeneous the trees might s t i l l be too young f o r meaningful early t e s t s . Brush Competition Vine maple, salmonberry and other species were repeat-edly treated chemically and mechanically i n an e f f o r t to prevent them from competing with Douglas-fir. There i s no doubt that Vacclnlum spp. occurring i n several concentrations (marked "b" f o r brush on Ormerod's map, page 6 3 ) and growing up to four feet t a l l , account f o r some of the v a r i a b i l i t y termed " s i t e hetero-geneity." The experimental trees outgrew th i s brush i n the l a s t few years. Table 8 shows the height increment of the l a s t three growing seasons ( 1 9 6 ^ - 1 9 6 7 ) before the measurements f o r t h i s thesis were taken. This difference was calculated i n an attempt to eliminate, i n theory, the e f f e c t of brush competition on height growth. It would be incorrect to assume that once the trees are t a l l e r than the brush, they grow completely unimpeded. Vacclnlum spp. i s d e f i n i t e l y s t i l l competing f o r s o i l moisture and n u t r i e n t s . Nevertheless, the height increment i n those three years shows a s l i g h t l y d i f f e r e n t ranking f o r many plots where some trees obviously suffered from brush competition. It Ranking of Provenances  Block Proven- Aver-ance # Origin I II III IV age Range 1=C Sugar Loaf Mountain, Vancouver Island 11 3 1 10 6.2 10 2=B Courtenay area, Vane. Island 12 1 10 13 9.0 12 3=D Robertson Valley, " 2 5 12 8 6 . 8 10 4=A Nlmpkish Valley, " 6 7 2 14 7.2 12 5=H Elbe area, Washington 8 12 8 7 8 . 8 5 6=G Shelton area, " 3 6 6 4 4 . 8 3 7=1 Tillamook area, Oregon 7 8 16 3 8 . 5 13 8=L Salem area, " 9 10 9 6 8 . 5 4 9=J Molalla, lower " elevation 13 9 7 11 10.0 6 10=K Molalla, higher elevation 4 16 11 2 8.2 14 11=M C o r v a l l l s area, " 10 11 '2 1 5 12 . 8 5 12=N Oakridge, lower " elevation 14 14 12 11.0 10 13=0 Oakridge, higher " elevation 1 5 4 14 5 9 . 5 11 l4=P Butte F a l l s , 16 1 5 13 16 15.0 3 ^ 15=E U.B.C. Research Forest, B r i t i s h Columbia 1 2 3 1 1 . 8 2* l 6=F Snoqualmle area, Washington 5 13 5 9 8.0 8 Table 8 Height performance between the years 1964-1967. Ranking i s from t a l l e s t (1) to shortest ( l 6 ) . i s true, however, that the provenance x block Interaction i s s t i l l s i g n i f i c a n t i n the analysis of variance, meaning that Vacclnlum cannot take a l l the blame f o r the heterogeneity i n s i t e . The l o c a l provenance. The Local Provenance In terms of height increment f o r the years 1 9 6 5 - 1 9 6 7 inclusive, the l o c a l source (University of B r i t i s h Columbia Re-search Forest) a l l of a sudden ranks second t a l l e s t i n block II where I t held a meager tenth p o s i t i o n f o r t o t a l height, when brush was not taken into account. In block III i t moves from f i f t h to t h i r d p o s i t i o n . Of a l l provenances tested, i t shows the narrowest range (from t h i r d p o s i t i o n i n block III over second i n block II to f i r s t i n blocks I and IV). One i s tempt-ed to c a l l i t " the best of a l l provenances as to ov e r a l l per-formance I f i t were not for the harsh laws of s t a t i s t i c s f o r -bidding t h i s conclusion against the background r e f r a i n of a s i g n i f i c a n t block x provenance i n t e r a c t i o n . The performance of the l o c a l source at t h i s stage might indicate that several years from now the ef f e c t s of temporary "juvenile handicaps" such as brush competition, f o r instance,will disappear and inherent differences that the co-operative study set out to t e s t , w i l l appear. I t may a l s o be speculated that the i l l e f f e c t s of s o i l compaction caused by logging, as i l l u s t r a t e d by Pearse ( 1 9 5 8 ) , may gradually fade as the roots loosen the s o i l . In tables 4 to 7 t h i s "upwards" trend of the l o c a l source can be followed f o r the years 1 9 6 0 - 1 9 6 4 - 1 9 6 7 . In block I i t moves from 4 t h p o s i t i o n to 1 s t II " " " 1 5 t h " " 1 0 t h * III " " " 1 5 t h " " 5 t h IV " " " 3 r d " " 1 s t In blocks I and IV the l o c a l source (University of # Brush competition i s s t i l l a problem In t h i s p l o t . B r i t i s h Columbia Research Forest) i s s i g n i f i c a n t l y d i f f e r e n t from ( i . e . superior to) a l l other sources a f t e r the 1967 grow-ing season. Table 6 also shows that every B r i t i s h Columbia provenance ranks at l e a s t once, f i r s t or second t a l l e s t . If any provenance can be rated as i n f e r i o r i t w i l l be Butte F a l l s , This i s not s u r p r i s i n g considering the difference In climate between Butte F a l l s and the University of B r i t i s h Columbia Re-search Forest where the temperature regime differences are more pronounced than i n southern Oregon. The Butte F a l l s provenance Is doing poorly i n every block, even i n block C, where i t can be said to p r o f i t from a s l i g h t amount of seepage water. How do These Results Compare with E a r l i e r Results? Ching ( 1 9 6 5 ) i n his report on early growth, did not f i n d any s i g n i f i c a n t differences between provenances at the University of B r i t i s h Columbia Research Forest f o r i 9 6 0 and 1 9 6 1 . He l i s t e d the sources as to t h e i r o v e r a l l performance, without mentioning the provenance x block i n t e r a c t i o n which was found to be s i g n i f i c a n t i n the present study. In his summary, Ching stated that " ... trees from the l o c a l seed source grew as well as did those from the best three sources." The r e s u l t s a f t e r the 1967 growing season at the University of B r i t i s h Col-umbia Research Forest c e r t a i n l y do not disprove that statement and i t Is f e l t that possibly the l o c a l source w i l l maintain i f not improve i t s po s i t i o n i n future years. The findings of Ching that "trees from seed c o l l e c t e d at northern and southern ex-tremes of the study grew l e a s t In the three years" are supported by the present thesis f o r the southernmost provenance (Butte P a l l s ) , whereas the northernmost source (Nlmpkish) Is doing very well at le a s t In one block. Seed source G (Shelton) has maintained Its p o s i t i o n as one of the very best. Ching and Bever ( i 9 6 0 ) as well as Heaman ( 1 9 6 3 ) did not f i n d any cor-r e l a t i o n between height growth and geographic variables such as l a t i t u d e and a l t i t u d e . Nothing was found i n the present study to support such a c o r r e l a t i o n e i t h e r . VII. EVALUATION OF THE CO-OPERATIVE PROVENANCE TEST Good Points The i n i t i a t i v e to convince a large portion of private and public forest Industry i n the P a c i f i c Northwest, Including B r i t i s h Columbia, of the importance of the provenance problem, to spend money, time and e f f o r t on the co-operative study, c e r t a i n l y deserves praise. The attempt to f i n d an answer to p r a c t i c a l and t h e o r e t i c a l aspects of the provenance problem was honest. The mistakes made (see below) I l l u s t r a t e the complex-i t y of the problem and help to make the task easier f o r inves-t i g a t i o n of future provenance t e s t s . Weak Points The following evaluation i s not meant to be c r i t i c a l of the originators of the study, but should be interpreted as a help f o r further provenance t r i a l s . Since temperature and f r o s t free period were to be the major c r i t e r i a i n determining the existence of d i s t i n c t "races" of coastal Douglas-fir, more emphasis should have been put on the standardization of weather measurements (see Heaman's c r i t i c i s m , page 5 1)• In such a mountainous region as the area covered by the co-operative provenance study, mere a l t i t u d e and l a t i t u d e , or t h e i r combination (see "hypotheses to be tested," page 3 9 ) cannot be expected to describe or to determine true c l i m a t i c d i f f e r e n c e s . Aspect, slope, l o c a l topography, proxim-i t y of the ocean, etc. are too i n f l u e n t i a l i n modifying the e f f e c t s of a l t i t u d e and l a t i t u d e , e s p e c i a l l y within a l i m i t e d area. The area covered was too large and the number of proven-ances included too small f o r meaningful clues on the nature of the v a r i a t i o n within coastal Douglas-fir. The a l t i t u d l n a l sampling was not uniform, since no representative was selected from 750 to 1250 feet elevation, where much logging i s being car r i e d out. The cone c o l l e c t i o n radius of twenty-five miles was too large, areas of great c l i m a t i c differences can be covered by a f i f t y - m i l e diameter, as i l l u s t r a t e d by Heaman ( 1 9 6 3 ) f o r the area around Lake Cowichan on Vancouver Island. Besides that, the cones were not c o l l e c t e d according to the same standards by a l l co-operators. One could almost say that a provenance test i s as good as the test s i t e s . If these are heterogeneous (as the University of B r i t i s h Columbia Research Forest test s i t e i s , f o r instance) i t i s very d i f f i -c u l t to test any hypothesis related to various provenances, although the d i f f i c u l t y i n f i n d i n g homogeneous s i t e s , large enough to accomodate two three-acre plantations i n the rugged topography on the P a c i f i c coast, i s f u l l y appreciated. Damage by hare and deer could have been pa r t l y avoided through fences. The f r o s t problem In the Robertson Valley could have been re-cognized by more c a r e f u l study of e x i s t i n g weather data. VIII. CONCLUSIONS AND RECOMMENDATIONS Since the analyses of variance show a s i g n i f i c a n t provenance x block i n t e r a c t i o n , no conclusion as to o v e r a l l performance of any p a r t i c u l a r provenance can be drawn from the measurements taken at the University of B r i t i s h Columbia Re-search Forest when the trees were eleven years old. Because of t h i s , none of the hypotheses that the co-operative proven-ance study set out to test can be accepted or rejected. Sig-n i f i c a n t growth differences between provenances do show up within Individual blocks, but the r e s u l t s are inconsistent and therefore not conclusive, due to the extreme s i t e heterogen-e i t y . Observed, but s t a t i s t i c a l l y not proven differences tend to Indicate that the l o c a l provenance i s by no means i n f e r i o r to any other seed source tested, and i n future years i t might prove to be the best one of a l l i n terms of height growth, as i t could be shown to move up the ranks over the l a s t years. Thus, the long-held view that l o c a l seed i s safest, and prob-ably best (e.g. Bingham, 1 9 6 6 ) , Is s t i l l v a l i d . The southern-most provenance i s doing very poorly at the University of B r i t i s h Columbia Research Forest. It i s possible that several years from now the e f f e c t s of Juvenile handicaps, such as brush competition and maybe s o i l compaction, w i l l have faded away and genetic differences, i f any, w i l l show up s t a t i s t i c a l l y . The r e s u l t s found Indicate the complexity of the provenance problem and the overwhelming influence of a hetero-geneous s i t e . Further provenance studies are necessary to i n -vestigate the nature and extent of v a r i a t i o n within coastal Douglas-fir and to e s t a b l i s h d e t a i l e d seed transfer rules and seed c o l l e c t i o n zones. Based on the findings at the University of B r i t i s h Columbia Research Forest and on the weak points of the co-operative t e s t , the following recommendations for prov-enance tests seem l o g i c a l (see also Heaman, 1 9 6 3 ) « 1 . General recommendations a) Standardization i n a l l phases of a co-operative study i s imperative. Controls are necessary and should be exercised by one co-ordinator, i n con-stant personal contact with a l l co-operators. b) Intensive sampling of a smaller area than that covered by the co-operative study, i s l i k e l y to y i e l d more information. c) Cone c o l l e c t i o n areas should be c l e a r l y defined, based on a thorough l o c a l knowledge. d) Outplanting areas have to be thoroughly studied. Besides topography and vegetation, c l i m a t i c data should be looked Into, the s o i l should be analyzed i n an attempt to have at l e a s t homogeneous blocks. e) More studies should be carried out to f i n d one c r i t e r i o n f o r s o i l f e r t i l i t y , or f o r s i t e d i f f e r -ences, so that better s t a t i s t i c a l r e s u l t s may be obtained. f) Weather measurements should be supplemented with phenological observations. g) Damage from animals and fungi has to be kept to a minimum. 2. Recommendation for management Plant l o c a l seed. Collect i t from selected trees during good crop years and store i t . This way unpleasant surprises due to seed crop f a i l u r e s and purchase of seed of unknown o r i g i n , can be avoided. IX. SUMMARY Importance, scope and nature of the provenance prob-lem are discussed i n t h i s t h e s i s . Experience gained from major provenance t r i a l s with Douglas-fir i n both Europe and North America i s reviewed. From t h i s , the complexity of the problem and the extreme genetic v a r i a b i l i t y of Douglas-fir are apparent. The "Co-operative Douglas-fir provenance t e s t " i s discussed In d e t a i l . It was started i n 1957 and included s i x -teen coastal origins from Oregon, Washington and B r i t i s h Colum-b i a , planted close to the seed c o l l e c t i o n areas, one of which i s located on the University of B r i t i s h Columbia Research Forest, where height growth was measured when the trees were eleven years old. The objectives i n analyzing these data were to f i n d whether s i g n i f i c a n t differences between provenances show at an early stage and whether the University of B r i t i s h Columbia Research Forest test s i t e i s homogeneous. The s t a t i s t i c a l analysis shows a s i g n i f i c a n t proven-ance x block i n t e r a c t i o n , i n d i c a t i v e of a heterogeneous s i t e . The r e s u l t s are very confusing and no conclusion can be drawn yet as to o v e r a l l performance of any i n d i v i d u a l provenance. A f t e r removing the e f f e c t s of brush, randomly scattered over the test s i t e , the r e s u l t s tend to become more meaningful. A l -though the i n t e r a c t i o n term i s s t i l l s i g n i f i c a n t , the following conclusions are drawn. The l o c a l provenance i s by no means i n -f e r i o r , and i t might prove to be the best one of a l l i n a few years when brush competition w i l l have become les s important. The southernmost provenance from Butte F a i l s , Medford, Oregon, i s i n f e r i o r to a l l other origins tested. It i s hoped that the measurements taken a f t e r the 1970 growing season by a l l co-ordinators w i l l y i e l d more con-clu s i v e Information than the measurements from the University of B r i t i s h Columbia Research Forest alone. The co-operative test i s c r i t i c a l l y evaluated and recommendations f o r future provenance studies are included. It i s f e l t that many more tests with Douglas-fir are necessary to answer the questions raised by t h i s and by other studies. LITERATURE CITED A l l e n , G.S. 1 9 6 0 a . A method of d i s t i n g u i s h i n g coastal from i n t e r i o r Douglas f i r seed. B r i t i s h Columbia Lumber-man 4-4-» 8126 - 3 0 . . 1 9 6 0 b . Factors a f f e c t i n g the v i a b i l i t y and germ-ination of coniferous seed. IV. S t r a t i f i c a t i o n period and incubation temperature. Pseudotsuga menziesli (Mlrb.) Franco. The Forestry Chronicle 3 6 i l 8 - 2 9 . 1 9 6 l . Testing Douglas f i r seed f o r provenance. Proc. Int. Seed Testing Assoc. 2 6 * 3 * 3 8 8 - 4 0 3 . Anon. 1955a. A cooperative seed provenance study f o r P a c i f i c Northwest Douglas f i r . Mimeo. 8 pp. 1955b. Provenance study. Cone c o l l e c t i o n pros-pectus. Mimeo. 4 pp. Baldwin, H.I., and D.A. Rock. 1 9 6 1 . Douglas f i r as a New Hampshire Christmas tree. Fox For. note No. 80, 2 pp. Barner, H. 1 9 6 6 . C i r c u l a r l e t t e r of A p r i l 2 0 , 1 9 6 6 , of working group on procurement of seed f o r provenance research. I.U.F.R.O. Section 2 2 , 10 pp. Bingham, R.T. 1 9 6 6 . Seed movement i n the Douglas f i r region (Symposium summary and recommendations). Proceedings 1 9 6 5 Western Forest Genetics Association. P a c i f i c Northwest Forest and Range Experiment Station, pp. 3 0 - 3 4 . Bramhall, G. 1 9 6 6 . Permeability of Douglas-fir heartwood from various areas of growth i n B.C. 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FA0/F0RGEN - 63/ - 4 /5 pp. i i i - 1 8 . , and H. Meyer. 195^. Douglasien Provenienzver-suche I I . Allgemelne Forst- und Jagdzeltung 125 I5 I 160-179. 1955* Douglasien Provenlenzversuche I I . A l l g e -melne Forst- und Jagdzeltung 126tll/l2i2 2 1 - 2 4 3 . Schoenbach, Hans. 1958. Die Zuechtung der Douglasle. In Goehrei Die Douglasle und i h r Holz. Akademie Verlag, B e r l i n , pp. 309-367. , and E r i c h Bellmann. 1967. Frostresistenz der Nachkommenschaften von Kreuzungen "gruener" und "blauer" Formen der Douglasle (Pseudotsuga menzlesll (Mirb.) Franco). Archiv fuer Forstwesen Band l6«6/9» 707-711. S i l e n , Roy R. 1962. A study of genetic control of bud burst-ing i n Douglas f i r . Journal of Forestry 6o»7»472-475. . 1964. Regeneration aspects of the 50-year-old Douglas f i r heredity study. Proceedings of the 1964 Annual Meeting of the Western Reforestation Coordin-a t i n g Committee of the Western Forestry and Conser-vation Association. Portland, Oregon, pp. 35-39* 1966. 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Provenance differences i n P a c i f i c Coast Douglas f i r seed and seedling c h a r a c t e r i s t i c s . Silvae Genetlca l 4 i 2«46 - 5 6 . S z i k l a l , Oscar. 1 9 6 4 . Preliminary notes on v a r i a t i o n i n cone and seed morphology of Douglas-fir (Pseudotsuga menzl-e s l l (Mirb.) Franco). Second World Consultation on Forest Tree Breeding. Washington. 11 pp. Tusko, Ferenc. 1963* A study of v a r i a b i l i t y i n c e r t a i n Douglas-fir populations i n B r i t i s h Columbia. Ph.D. t h e s i s . Department of Biology and Botany, University of B r i t i s h Columbia, Vancouver 8, B.C. 173 PP. U.B.C. Forest Committee. 1959* The f i r s t decade of manage-ment and research. U.B.C. Forest 1949-1958. Faculty of Forestry, University of B r i t i s h Columbia, Vancouver 8, B.C. 82 pp. U.S. Department of Agriculture, United States Forest Service. 1 9 6 2 . 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Annual Report of University of B r i t i s h Columbia Research Forest, Haney, B.C. Faculty of Forestry, University of B r i t i s h Columbia, Vancouver 8 , B.C. p. 2 5 . . 1 9 6 7 . Annual Report of University of B r i t i s h Columbia Research Forest, Haney, B.C. Faculty of Forestry, University of B r i t i s h Columbia, Vancouver 8 , B.C. p. 1 1 . . 1 9 6 8 . Annual Report of University of B r i t i s h Columbia Research Forest, Haney, B.C. Faculty of Forestry, University of B r i t i s h Columbia, Vancouver 8, B.C. p. 1 2 . . 1970. Personal communication. , and Kim K. Ching. 1969* Pattern and bud burst i n a Douglas-fir provenance study. Unpublished. , and J . Soos. 196la. The r e l a t i v e e f f i c i e n c y of three hare-repellents i n protecting Douglas-fir seed-l i n g s . Forestry Chronicle 37.22-28. . 196lb. Some observations on the re l a t i o n s h i p of lammas shoots to the form and growth of Douglas-fir seedlings. Faculty of Forestry, University of B r i t i s h Columbia, Vancouver 8 , B.C. Research Paper 40. 8 pp. Wheat, Joseph G. 1 9 6 6 . The seed movement problem i n the Douglas f i r region. Western Forest Genetics Associ-a t i o n . Proceedings, 1965* PP. 3-5» Willington, Robert P. 1 9 6 8 . Some ef f e c t s of slashburning, c l e a r c u t t i n g and skidroads on the physical-hydrologic properties of coarse g l a c i a l s o i l s In coastal B r i t i s h Columbia. M.F. t h e s i s . Faculty of Forestry, Univer-s i t y of B r i t i s h Columbia, Vancouver 8 , B.C. 1^9 pp. Wright, Jonathan W. 1962. Genetics of forest tree improve-ment. Forestry and Forest Products Studies No. 1 6 , FAO, Rome. 399 pp. , and H.I. Baldwin. 1957. The 1938 International Union Scots pine provenance test i n New Hampshire. Silvae Genetica 6il/2«2-l4. Zon, Raphael. 1913* E f f e c t of seed source upon the growth of Douglas f i r . Forestry Quarterly 111 ^ 9 9 - 5 0 2 . S c i e n t i f i c and common names of species c i t e d In the text Abies amabills (Dougl.) Forbes Acer cireinaturn Pursh. A r m l l l a r l a mellea (Fr.) Kumm. Ghamaecyparls nootkatensls (D. Don) • Spach Contarlnla pseudotsugae Condrashoff Eplloblum angustlfollum L. Larlx decldua M i l l . Plcea ables (L.) Karst Plcea sltchensls (Bong.) Carr. Plnus montloola Dougl. Plnus ponderosa Laws. Plnus s y l v e s t r l s L. Plnus taeda L. Pseudotsuga menzlesll (Mlrb.) Franco Rhabdocllne pseudotsugae Syd. Rubus spe o t a b l l l s Pursh. Thuja p l l c a t a Donn. Tsuga heterophylla (Raf.) Sarg. P a c i f i c s i l v e r f i r Vine maple A r m i l l a r i a root rot fungus Yellow cedar Douglas-fir needle midge Fireweed European l a r c h Norway spruce Sitka spruce Western white pine Ponderosa pine Scots pine L o b l o l l y pine Douglas-fir Douglas-fir needle b l i g h t fungus Salmonberry Western red cedar Western hemlock 

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