a V. I l rm”... ‘fkg/ x“ «a J? JUL 1 9’ 2383' ABSTRACT THE CORRESPONDENCE BETWEEN GENETIC, MORPHOLOGICAL AND CLIMATIC VARIATION PATTERNS IN SCOTCH PINE by John L. Ruby Six hundred and eighty-nine cone, seed and leaf specimens of Scotch pine were collected from 39 stands in 10 regions of Europe and Asia by cooperators in those countries. Nineteen characters or ratios were measured on these. The individual tree measurements were grouped by stand and region, and then were subjected to analyses of variance. Computations of the components of variance for the 8 most definitive characters showed that more than 85 per cent of the variance was attributable to between-region differences, and less than 3 per cent to stands-within-region. Comparisons were made of the regional grouping based on the parental measurements and a regional grouping based on an associated 122~origin provenance study in a uniform environment in Michigan. The groupings were nearly identical. In other words, it is possible to delimit a race or variety nearly as well by studying parental specimens collected in Europe as by growing their progenies in this country. However, it was not possible to forecast a race's perform- ance in Michigan from a study of the parental cones and leaves. John Lindley Ruby Character differences in juvenile performance were often re- lated to the climate of the parent races. The fibrous shallow roots of the northernmost provenances were associated with an area where precipitation was not critical, while the tap roots of the Spanish origins reflected the affects of summer drought and high temperatures. Similarly the blue-green leaf coloration of the Spanish seedlings was due to the development of protective waxes in a dry climate. The taxonomy of Scotch pine was reviewed for the last 60 years and summarized by geographic areas. The 10 geographic varieties defined in this study were compared and combined with other biometric and provenance results of Scotch pine studies and the varietal descriptions of previous authors and taxonomists. Based on this combined information 21 varieties of Scotch pine were named and their geographic location, taxonomic description and synonymy were detailed. As regards stands within a region, there was little corre- spondence between variation patterns exhibited by the parents and seedlings grown in this country. In other words, two stands which differed markedly in some cone or leaf trait might or might not differ appreciably in some seedling trait. This is thought to be so because the parental traits are subject to considerable environ- mental influences. Also it is likely that the genetic factors af- fecting mature cone or leaf traits and juvenile traits are different. It was not possible to forecast which of several German-Czecho- slovakian stands should have the tallest (or greenest or longest John Lindley Ruby needled) offspring in Michigan from a study of the microclimates of the stands in Europe. This general lack of climate-progeny correlation was also true for the western Germany-eastern France regions. Apparently micro-evolution lags so that every stand is not perfectly adapted to its particular local climate. In seven stands the parental data for growth rate, clear- stem length, crown size and length, and stem straightness, as well as for cone and leaf traits, were available for each parent tree. Two-year performance data for the offspring of those parents were also available. Coefficients of correlation were calculated between all parental characters and 20 juvenile characters in an effort to determine if any parental character could be used to predict juvenile performance in Michigan. Only 35 correlations were significant at the 5 per cent level out of a possible 1,260 combinations. There was little consistency from one stand to the next. The relative absence of correlations is thought to be due to genotype-environment interactions between Europe and America, and to the different ages at which parents and offspring were observed. THE CORRESPONDENCE BETWEEN GENETIC, MORPHOLOGICAL AND CLIMATIC VARIATION PATTERNS IN SCOTCH PINE B)’ John Lindley Ruby A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Forestry 1964 ACKNOWLEDGEMENTS The author wishes to acknowledge the assistance of the guidance committee: Drs. Jonathan W. Wright, John E. Grafius, Donald P. White, John H. Beaman, and Clinton E. Peterson. A special debt of gratitude is due to the many European and Asian Research Foresters who cooperated in the collection of paren- tal specimen materials. The study was financed in part by funds from the Cooperative State Research Service of the U. S. Department of Agriculture as part of regional project NC-Sl entitled ”Tree Improvement through Selection and Breeding.” This project involves active cooperation of numerous federal, state, and private agencies in the North Cen- tral United States. ii TABLE OF CONTENTS ACKNOWLEDGEMENTS . . . . . . . . . . . . . . . . . . . . . LIST OF LIST OF Chapter I. II. III. IV. VI. TABLES . . . . . . . . . . . . . . . . . . . . . . ILLUSTRATIONS O O O O O O O O O O O O O C O O O 0 INTRODUCTION . . . . . . . . . . . . . . . OBJECTIVES O O O O O O O O O O O O O 0 O O O O 0 MATERIALS AND METHODS . . . . . . . . . . . . . . . Specimen Procurement Data Concerning Area of Origin Specimen Handling Measurement Methods VARIATION IN PARENTAL CHARACTERS WITHIN AND BETWEEN REGIONS C O O O 0 O O O I O O O 0 O O O O O O 0 Methods Regional Variation Pattern The Naturalness of the Regional Grouping Sample Size RELATIONS INVOLVING SINGLE PARENTS AND THEIR PROGENY O C O O O O C O C O O O O O 0 0 Materials and Methods Parent-Parent Correlations Parent-Progeny Correlations Progeny-Progeny Correlations CORRELATIONS INVOLVING PARENTAL CHARACTERISTICS . Materials and Methods Correlations Within Stands Correlations Between Widely Distributed Stands and Between Regions Correlations Between Stands Within Regions Discussion iii Page ii viii l7 19 23 70 102 Chapter Page VII. CORRELATIONS INVOLVING PARENTAL AND JUVENILE CHARACTERISTICS OF ENTIRE STANDS . . . . . . 116 Materials and Methods Discussion VIII. CORRELATIONS INVOLVING CLIMATE . . . . . . . . 122 Materials and Methods Literature Review Weather Patterns Within Scotch Pine Distri- bution Area Correlations Involving Parental Character- istics and Climate Correlations Involving Juvenile Characteristics and Climate Ix. IAXONOMY o o o o o o o o o o o o o o o o o o 0 16]- Synonymy Validity of Named Varieties Discussion X. DISCUSSION 0 O O O O O O O O 0 O O O O O I O O 202 Problems in Methodology value of Parental and Climatic Data Trends in Evolution BIBLIOGRAPHY . . . . . . . . . . . . . . . . . . . . . 216 iv 10. 11. 12. 13. LIST OF TABLES Description of cone, seed, and leaf characteristics measured on parental specimens of Scotch pine . . . Significance of the variation between stands in the same region and between regions for various parental characters . . . . . . . . . . . . . . . . . . Components of the variance in parental characters supplied by trees-within-stands, stands-within-region, and regions of origin . . . . . . . . . . . . . . . Summary of significant parental character differences between regions . . . . . . . . . . . . . . . . . . . Description of parental stands . . . . . . . . . . Description of parental characters and grades . Description of juvenile characteristics and grades used in scoring Scotch pine progeny tests . . . . . . . . Correlations between parental characters in five East Gama 8 tands O O O O C C O C O O O I I O O O I O C O Parent-progeny and progeny-progeny correlations for nine Belgian trees from two stands . . . . . . . . . Parent-progeny and progeny-progeny correlations for eight Norwegian trees from a natural stand in southern Norway . . . . . . . . . . . . . . . . . . . Parent-progeny and progeny-progeny correlations for 20 trees (Nos. 321 to 340) from an even-aged stand near R5vershagen, East Germany . . . . . . . . . . . Parent-progeny and progeny-progeny correlations for 20 trees (Nos. 341 to 360) from an even-aged natural stand near Neustrelitz, East Germany . . . . . . . . Parent-progeny and progeny-progeny correlations for 20 trees (Nos. 361 to 380) from an even-aged stand near GUstrow, East Germany . . . . . . . . . . . V Page 27 29 3O 61 72 73 75 78 83 84 85 86 87 Table Page 14. Parent-progeny and progeny—progeny correlations for 20 trees (Nos. 381 to 400) from an even-aged stand near Nedlitz, East Germany . . . . . . . . . . 88 15. Parent-progeny and progeny-progeny correlations for 20 trees (Nos. 501 to 520) from an even-aged stand near Joachimsthal, East Germany . . . . . . . . . . 09 16. Parent-progeny and parent-ramet correlations re- ported by Nilsson (1958) for Scotch pine and Norway spruce in south-central Sweden . . . . . . . . . . . 94 1?. Correlations of parental characters within 39 stands expressed as the per cent of stands in which r exceeds the amount needed for significance at the 5 per cent level . . . . . . . . . . . . . . . . 111 18. Significant between stand correlations for 19 parent- al characters using stand means of 39 parent stands. 112 19. Significant between region correlations for 19 paren- tal characters using regional means for the 10 re- gions, A,C,G,H,I,J,K,L,M, and N . . . . . . . . . . 113 20. Significant between-stand correlations for 19 paren- tal characters, using stand means for regions A, C, and G only C O O C O O O O O O O O O O O O O O O 114 21. Significant between-stand correlations for 19 paren- tal characters using stand means for regions H, J, K, M, N, and Turkey . . . . . . . . . . . . . . . . 115 22. Significant correlations between parental and juve- nile characteristics within group 1, consisting of regions A, C, G, and E, group 2, consisting or re- gions I and L, and within group 3, consisting of regions J, K, M, and N, using stand means . . . . . 120 23. Description of juvenile characteristics used in corre- lations with meteorological and geographical data of origin and the grades used in scoring the Scotch pine progeny . . . . . . . . . . . . . . . . . . . . 125 24. Correlation of latitude, longitude, and elevation with precipitation data in regions A-F, G, H, and J, K, M, and N O O O I O O O O O O .0 O O O O O O O O O 130 25. Correlation of latitude, longitude, and elevations with temperature data in regions A-F, G, H, and J, K, M, and N O O O O O O O O O O O O O O O O O O 0 O 131 vi Table 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. Page Correlations between parental characteristics and meteorological data for north European Scotch pine (regions A, C, and G). . . . . . . . . . . . . . . . . 135 Correlations involving relationships between parental cone, seed, and leaf characteristics and meteorologi- cal and geographical data for regions H, I, J, K, L, M, and N . . . . . . . . . . . . . . . . . . . . . . . 136 Correlations involving relationships between juvenile characteristics and meteorological and geographical data of origin for regions A through F, northern Europe . . . . . . . . . . . . . . . . . . . . . . . . 142 Correlations involving relationships between juvenile characters and meteorological and geographical data of origin for region G, northeast Germany and Czecho- slovakia . . . . . . . . . . . . . . . . . . . . . . . 146 Correlations involving relationships between juvenile characteristics and meteorological and geographical data of origin for region H, western Germany, eastern France, and Belgium . . . . . . . . . . . . . . . . . 148 Correlations involving relationships between juvenile characteristics and meteorological and geographical data of origin for regions J, K, M, and N, the Medi- terranean area . . . . . . . . . . . . . . . . . . . . 151 Multiple correlations and regressions for the Medi- terranean stands for juvenile characters, winter fol- iage, December, 1961, and third-year height . . . . . 159 Principal comparative characteristics of four Scotch pine varieties as given by Carlisle (1958) . . . . . . 175 Summary of the named varieties of Scotch pine and the geographic regions in which they predominate . . . 181 Varieties of Scotch pine (Pinus sylvestris L.) and the regions in which they predominate. . . . . . . . . . . 186 vii Figure 1. LIST OF ILLUSTRATIONS Natural distribution of Scotch pins in Europe and provenances included in Wright and Bull (1963) test . Natural distribution of Scotch pine in Asia and pro- venances included in Wright and Bull (1963) test . . . Provenances included in this study of parental charac- ters O O O O O O O O O O O O O O O O O O O C O O O O O Seventeen year old Scotch pine 20-25 feet high and 13 year old red pine (Pinus resinnsa Ait.) 12-15 feet high in a wind erosion contrOL plantation in Newaygo Country, Michigan, June, 1960. . . . . . . . . . . . . . . . . . Natural reproduction of Scotch pine averaging 18 inches in annual growth from 22 year old parent trees 25- 35 feet high along U. S. Highway 2 near St. Ignace, Michi- gan . g o o o o o o o o o o e o e o o o o o o o o Thirty year old plantation of Scotch pine thinned in 1960 at Michigan State University's Kellogg Forest in Kalamzaoo County, Michigan . . . . . . . . . . . . . General view of seedling beds in Bogue Forest Research Nursery, East Lansing, Michigan, containing 4 repli- cates of the Scotch pine 122-origin provenance test . . View across seedling beds in Bogue Forest Research Nur- sery, East Lansing, Michigan, showing differences be- tween 1-0 Scotch pine seedlings from Spain and Norway in provenance test in September, 1959 . . . . . . . Scotch pine provenance test outplanting of 2-0 stock at Michigan State University's Russ Forest in Cass County, Michigan, one year after establishment in the spring of 1961. . . . . . . . . . . . . . . . . viii Page 10 12 12 14 14 16 16 Figure 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. Page Cone and cone scale measurements . . . . . . . . . . . 32 Regional variation pattern in length, width, and length/width ratio of Scotch pine seeds . . . . . . . 36 Regional variation pattern in cone length, width, and length/width ratio as determined from closed cones . . 38 Regional variation pattern in cone length, width, and length/width ratio as determined from open-cones . . . 40 Regional variation pattern in length and width of lar- gest apophysis and the length/width ratio of the lar- gest apophysis . . . . . . . . . . . . . . . . . . . . 44 Regional variation pattern in the cone length/length of largest apophysis ratio . . . . . . . . . . . . . . 47 Regional variation pattern in thickness of the largest apophysis, thickness of the opposite apophysis and width of the open cone . . . . . . . . . . . 50 Regional variation pattern in the basal angle of the cone and the length/thickness ratio of the largest apophysis . . . . . . . . . . . . . . . . . . . . . . 52 Regional variation pattern in leaf length and twist. . 55 Intra-regional variation pattern in region G . . . . . 69 ix CHAPTER I INTRODUCTION The variation in phenotypic expression within a given species resulting from the interaction of genetic diversity and environment is receiving increased attention in studies of the range of vari- ability within tree species. A thorough knowledge of this variability is a prerequisite to the intelligent use of tree species in silvi- cultural practice. The present common practice of utilizing only local seed sources or those from a narrowly restricted zone of similar environmental conditions seriously limits the development of our forest resources. Scotch pine (Pinus sylvestris L.) is a species widely dis- tributed throughout Europe and Asia. It has been greatly affected by the complex pattern of recent glaciation, particularly in Europe. The relatively recent changes in species distribution and in selec- tion pressures brought about by glaciation in the northern part of the northern hemisphere has been described by Hulten (1937, 1949) and others. Changing environments have resulted in genetic differen- tiation among continuous populations as well as in the isolated rem- nant populations of southern Europe and central Asia. Recently studies have been completed on phenotypic variation in the following pine species: Lodgepole pine (Pinus contorta Dougl.), Critchfield (1957). Black pine (Pinus nigra Arnold), Vidakovic (1960). Scotch pine (Pinus sylvestris L.), Staszkiewicz (1960, 1961, 1962). Jack pine (Pinus banksiana Lamb.), Schoenike et a1. (1959), Rudolph et a1. (1957). Loblolly pine (Pinus taeda L.), Thor (1961), Zobel and Thorbjornsen (1961). Virginia Pine (Pinus virginiana Mill.), Thor (1964). Monterey pine (Pinus radiate D. Don), Fielding (1953). Coulter pine (Pinus coulteri D. Don), Zobel (1952) (1953). Sand pine (Pinus clause (Chapm.) Vasey), Little and Dorman (1952). The few studies that have been made of the natural variation in the genus gingg have usually been limited in the coverage of the natural range of the species. Provenance tests to determine genetic differences in the same parental populations have been rare. Critch- field (1957) included both parental and seedling provenance studies in his study of geographic variation in lodgepole pine. The progeny provenance tests of jack pine made by Schantz-Hansen and Jensen (1954) were followed by a parental study of the same stands by Schoenike g£_gl. (1959), and Rudolph g£_al. (1959). Rudolph's §£_gl. (1957) study of jack pine parental variation in Minnesota was followed by vegetative propagation in a uniform environment. Thor (1964) has established a seedling provenance test in conjunction with his study of parental characteristics of Virginia pine. The genetic variability in pine species has been widely studied through seedling provenance tests. Recent Scotch pine studies include those conducted by Weidemann (1930), Langlet (1934), Veen (1952), Vincent and Polnars (1953), Holst (1953), Baldwin (1955, 1956), Wright and Baldwin (1957), Rudolf and Slabaugh (1958), Langlet (1959) comment on Wright and Baldwin, Bolsinger (1958), Echols (1958), Gerhold (1959), Wettstein (1958, 1959), Sanikov (1959), Bouvarel (1959) comment on Langlet (1959), Vojcal (1961), Lazarescu gtggl, (1961), Troeger (1962), Rubner (1962), Wright (1963), Wright and Bull (1963), Dengler (1938). Scotch pine is ideally suited to genetic and phenotypic study because of its great variability. It has been described as a complex by Semmler and Von Schiller (1927) and Mirov (1961). The utilization of this variability to improve the characteristics of stands planted under varying environments is a necessity. A knowledge of the genetic variability of the species and the parental phenotypic expression associated with the variability can materially improve and speed up the utilization of the best adapted races in a planting program. The natural distribution of Scotch pine covers a greater area than that of any other pine. According to Elwes and Henry (1908): . . . It is by preference a tree of siliceous soils, but occurs on almost all geological formations; and in Scotland, Norway, and Sweden grows on peat bogs too wet for the spruce to exist on. The area of distribution includes almost all Europe and the greater part of Northern Asia. The northerly limi}, com- mencing on the north-west coast of Norway at Alten 70 N. lat.), passes through Lapland, south of the Enara lake (68 50'), and touches Pasvig Fjord on the arctic sea at 69°30'. Extending through the Kola peninsula from Kola bay, it crosses the White sea a; 66 45' and in the Petchora territory goes as far north as 67 15' and crosses the Ural at about 64 . In Siberia it never reaches quite as far north as the arctic circle, though it nearly touches it on the Ob and Yenisei rivers; east of the Lens river it descends to aboug 64 . It reaches its extreme easterly point (about 150 E. long.) in the Wercho- jansk Mountains. The easu:n limit descends from there through the Stanovoi Mountains and the Seja territory to the upper Amur. According to Komarov (Flora Manshuriae, i. 175 (1901) it is a scarce tree on the banks of thefrivers in Manchuria. Its southerly limit in Siberia is not well known; but it is known to occur in the mountains of Dahuria in the territory around Lake Baikal, and in the Altai Mountains. Its southern limit in European Russia is a very irregular line, yhich be- gins in the Ural south of Orenburg at about latd 52 , is most to the north in the government of Tula jlat. 54 30'), and descends from thereoto Kharkof (lat. 49 ), passing into Ga- licia about lat. 50 . Far south of this line, and separated from it by the Russian Steppes, on which no pine trees grow, occurs an area of distribution, not yet well made out, which includes the Caucasus, the mountains of the Crimea, Asia Minor, and Northwest Persia. (g, sylvestris grows on the Armenian plateau, and has been described in Linnaea, xxii.296 (1849), as P. armgna, Koch; P. kochiana, Klotzsch; and P. pontica, Koch.‘ Cf. Moniteur Jardin Botanique Tiflis, 11.26 (1906). There is also an isolated area, in which the pine is found growing wild, in Macedonia, on Mount Nidjél From Galicia the southern limit in Europe (exclusive of the last mentioned area) passes southward to the Transylvanian Alps; thence it extends along the mountains to Serbia where the tree grows on the Kopavnik mountain (about lat. 43o ), continues through the mountains of Bosnia, Dalmatia, Illyria, Venetia, and through Lombardy to the Ligurian Appennines (about lat. 44 ). It passes into France, across the Maritime Alps, into the Cevennes, and reaches the Eastern Pyrenees; in Spain it descends through the mountains of Catalonia, Aragon, and Valencia to the Sierra Nevada in Andalusiad which is its extreme southerly point in Europe (lat. 37 ). The westerly limit beginning here, stretches northwest through the mountains of Avila to those of Leon in North Spain, and is continued through the mountains of Scotland to the north- west coast of Norway. In this vast area the pine is very irregularly distributed. The largest forests occur in the Baltic provinces of Russia, in Scandinavia, in Northern Germany, and in Poland. Towards the south it only occurs in mountains, and rarely forms pure forests of considerable extent. According to Huffel (Forets de la Roumanie, 6 (1890), it is rare in Roumanie, where he saw it at the confluence of the Lotru and Oltu rivers at 1700 feet altitude, and in the valley of Bistritza (only on calcareous soils in Muscel region). See figures 1 and 2. Scotch pine has been widely planted outside of its native range in Europe and Asia, particularly in the Northeast and North Central Regions of the United States, where it has propagated itself and exhibited rapid growth. Unfortunately many of the planted stands are of poor form, having originated from seed of doubtful origin or from areas known to produce trees of poor form. Scotch pine has been widely planted on relatively sterile soils in sand dune and soil stabilization projects where its pioneer qualities of survival and growth under poor conditions of moisture and soil have been outstanding. Stands of good form and quality in Michigan attest to the potential of Scotch pine as a timber source. Figure l.--Natura1 distribution of Scotch pine in Europe (shaded) and provenances included in Wright and Bull (1963) test (numbered dots). 15° 20° 25° 0° 5. l0° .00 } 2 ans nfll ’ fl lass W/ Figure 2.--Natura1 distribution of Scotch pins in Asia (shaded) and provenances included in Wright and Bull (1963) test (numbered dots). \ .* CA CO”. oON. so: .8. .00 .00 00h \\ M08 x a N. :\\\x\, \ I .u.\ _. \_ x _. r».\.\ w\ \ \ \\ \,. \ _ . . _ \\ «\w\\ x\\‘ 10 5'. I? 6' 0‘ 6' IZ' IS. 24' 30° 30° 42' >- 63' 0 22“. 4’5. 57' "oi. .. zm-efigs“ 2 .9. 7 5l' 2” 2. I') ‘7 4. ,‘I 45- '“ ‘z' . .“z' or” 3, ' - . . ' 2:2 1. a; ‘ moo, I""\.,. 7.: ‘ ”I“ 5. ..— 39° } 8' 0' 6' |2° [8° 24' 30' 36’ of parental Figure 3.--Provenances included in this study characters (numbered or lettered dots). 11 Figure 4.--Seventeen year old Scotch pine 20-25 feet high and 13 year old red pine (Pigus resinosa Ait.) 12-15 feet high in a wind erosion control plantation in Newaygo County, Michigan, June, 1960. Figure 5.--Natura1 reproduction of Scotch pine averaging 18 inches in annual growth from 22 year old parent trees 25-35 feet high along U.S. Highway 2 near St. Ignace, Michigan. 12 Figure 5. 13 Figure 6.--Thirty year old plantation of Scotch pine thinned in 1960 at Michigan State University's Kellogg Forest, in Kalamazoo County Michigan. Figure 7.--General view of seedling beds in Bogue Forest Re- search Nursery, East Lansing, Michigan, containing four replicates of the Scotch pins 122 origin provenance test. Figure 7 15 Figure 8.--View across seedling beds in Bogue Forest Research Nursery, East Lansing, Michigan, showing differences between 1-0 Scotch pine seedlings from Spain and Norway in provenance test in September, 1959. Each row represents a provenance. Figure 9.--Scotch pine provenance test outplanting of 2-0 stock at Michigan State University's Russ Forest in Cass County, Michigan, one year after establishment in the spring of 1961. This 4 tree plot 10 replicate planting clearly shows color differences, particularly the winter yellowing of foliage, and height differences. l6 V_.__!_.__i Figure 9. -.- "-\. - .— CHAPTER II OBJECTIVES This study is part of a long-range program directed toward the improvement of Scotch pine planted in the north central United States. The long-range program now includes a series of range- wide provenance tests to determine the best European origins. It also includes a series of l-parent progeny tests to determine the role and the best methods of applying selection for the betterment of the best natural populations. Ultimately it will include the establishment of seed orchards and breeding for specified goals. Broadly speaking, the objective of the present study was to determine the extent to which the performance of different Scotch pine genotypes when planted in Michigan could be forecast from the characteristics of these same genotypes in their native environments or the characteristics of those native environments. More specifically the objectives were (1) to determine the cor- respondence of the juvenile variation pattern found in the Michigan provenance tests with the temperature, precipitation, and daylength patterns of the localities of origin, (2) to determine‘the corres- pondence between the juvenile variation pattern found in Michigan and certain parental charcteristics such as the morphology of cone, seed, and leaf, (3) to determine the correspondence between juvenile performance in Michigan of the offspring of single trees and of the 17 18 parents growing in Europe, and (4) to determine the extent to which genetic variation patterns as studied in uniform-environ- ment growth tests are correlated with morphological variation patterns as studies in the field. CHAPTER III MATERIALS AND METHODS Specimen Procurement Cone and leaf specimens were collected in 1960, 1961, and 1962 by cooperators in Europe and Asia. Letters requesting the specimens were sent to forty-four cooperators who had pre- viously furnished seed for the 122-origin provenance test initi- ated by Doctor Jonathan W. Wright at Michigan State University in 1958 (Wright (1963), Wright and Bull (1963)). The requests were for one cone and one leaf from ten to twenty trees per stand with sufficient spacing between trees to avoid inbred parents. The single needle was to be the largest 1960 needle from the south side of the tree and was to be placed in an envelope with the largest 1960 cone from the same tree. The initial request was for collections from open-grown trees 3 to 6 meters high. There was an additional request for speci- mens from the same trees from which seed had been originally collected for the provenance test. Forty stand collections were received including eight individual tree collections from Norway, Belgium, and Germany matching those of the individual tree collection reported on by Wright (1963). The remaining collections were from the same stands as the original seed collections or from nearby stands. 19 20 One collection was too small to analyze so that the data for thirty-nine stands was used. The location of the stand col- lections is shown in Figure 3. The cones and needles were catalogued and stored by tree and stand as they arrived. They were assigned the same MSFG (Michigan State Forest Genetics) number given the original stands that were included in the seed sampling for the provenance test. Data Concerning_Area of Origin Meteorological data for the 122 stands represented in the provenance test of Wright (3219.) were collected from the cooperators, weather services of the local governments, the United States Department of Commerce Weather Bureau, and from the Great Britain Meteorological Office (1958). The remote location of many of the stands permitted the collection of monthly and annual data only for use with the collection as a whole. In some instances, corrections for altitude were applied where the stands were far removed from.the reporting meteorological stations. These corrections were made by the cooperators or with their advice. The original request for specimens also were for data on the geographic location, political subdivision in which stand occurred (province, county, or town name), elevation of stand above sea level, type of soil, age of tree, the size of the collecting area, the date of collection and shipment, and the name of the collector. 21 Spgcimen Handling All cone and needle specimens were packaged and identified by tree for each stand collection to preserve the identity of the trees. In eight stand collections these numbers also cor- responded with the individual tree provenance test data (Wright, 1963). The seeds were extracted and one typical seed with seed wing was attached to a card representing a stand collection. The remaining seeds were packaged by tree. The needles were placed in vials and the cones in com- partmented egg cartons. Measurement Methods Micrometer calipers were used for all length, width, and thickness measurements on cones, cone scales, and seeds. Needle length was measured on a flat scale. All measurements were in the metric scale to the nearest tenth of the unit of measure. All specimens were stored in a constant environment for one month before measurement to equalize moisture content. The open cones were measured for length and width of cone and length, width, and thickness of apophyses, then soaked in water for five minutes and placed under a polyethylene sheet for twenty-four hours before closed cone width was measured. During the period in which the cones were drying to a con- stant moisture content they were stored in open egg cartons and studied for measurable morphological differences. Form factors 22 of ratios of measurements were selected which would metrically describe the variability in shape. The manner of making specific measurements and of ana- lysing the data is discussed under the chapters concerned with the analysis of the data. CHAPTER IV VARIATION IN PARENTAL CHARACTERS WITHIN AND BETWEEN REGIONS The choice of parental characters used in this study was influenced by four factors: review of literature, estimated influence of environment, availability and transportability of materials, and ease and rapidity of measurements. Vidakovic (1958, 1960) studied the significance of seed, cone, and cone scale characters as taxonomic determinants in European black pine (Pinus nigra Arn.). The characters used were: 1. Seed color. 2. Seed mottling. 3. Color of scales. 4. Length, width, and thickness of seeds. 5. Length and width of cones. 6. Form of seed and cones. 7. Form of scales. 8. Weight of seed. He found that the italicized characters, 1, 2, and 6, were most useful in differentiating between populations. Staszkiewicz (1960, 1961, 1962) used the following cone characters in establishing morphological differences between Scotch pine populations: 23 24 1. Length of cones. 2. Width of cone. 3. Number of scales. 4. Length of apophysis. 5. Width of apophysis. 6. Thickness of apophysis. 7. Cone length/width ratio. 8. Relation of length of cone to number of scales. 9. Apophysis length/width ratio. 10. Apophysis length/thickness ratio. He based his study on cones from Poland, Czechoslovakia, Switzer- land, France, Scotland, Sweden and Finland. Staszkiewicz able to divide the material into 6 morphological types of cones, each type distinguished by some characteristic feature. Renvall (1914) studied the variation in leaf length within mature individuals of Scotch pine and found that the average needle length declines with increasing branch order and branch age. Gerhold (1959) studied the chloroplast pigments and nutrient elements in the needles of six geographic origins of Scotch pine growing in the New Hampshire IUFRO plantings. He found significant differences in needle color, total chlorophyll, magnesium, nitrogen, iron and calcium. Cvrkal (1958) used Sutherland's methods to determine dif- ferences in the essential oils of Scotch pine from several European countries and checked the accuracy of the results by infra-red spec- trums and chromatographic methods. 25 Fielding (1953) found distinct differences among three stands of Monterey pine (Pinus radiata D. Don) in cone and seed size, seed color, and needles per fascicle. A Critchfield (1957), in his study of geographic variation in lodgepole pine (P. contorta Dougl.), found that seven cone and leaf characters tended to follow a regional pattern while others conformed to elevation gradients. Resin canal number, leaf width, cone density, cone angle, cone symmetry, cone per- sistence, and apophysis form showed regional differences. He concluded that elevational variation was high in lodgepole pine, that each character studied had a unique variation pattern, and that there was no correlation with latitude in any characteristic. Schoenike‘g£_gl. (1959) reported in their study of cone variation in jack pine (P. banksiana Lamb) that cone closure and symmetry were useful in developing regional patterns. Thor (1961) studied 18 stand samples from 10 southern and southeastern states in his investigation of variation patterns in loblolly pine (P. taeda L.). Within-stand variation accounted for a large proportion of the total variation. The majority of the 13 morphological characteristics, including seed-wing length, seed length, needle length, cone length, and frequency of serrations on the needle margin, did not show regional trends. However, seed form, seed coat thickness, cone weight, cotyledon numbers and stomatal frequencies showed regional trends but no evidence of discontinuity. Thor (1964) also studied natural variation in the wood properties of Virginia pine (P. virginiana Mill.). Basing his W 26 results on 13 stands in Tennessee and Kentucky he found highly significant differences within stands for radial growth, specific gravity based on green volume, and specific gravity after removal of extractives. He found regional differences in length of summer- wood tracheids of the 10th and 25th year. There was a geographic pattern to the specific gravity variation if based on fresh wood but not if based on extracted wood. Wood characteristics, particularly tracheid length and specific gravity, have been studied by many researchers including Zobel (1961), Zobel and Rhodes (1955), Zobel and McElwee (1958), Zobel g£_gl. (1961), Kramer (1957) and Echols (1958). The earlier studies in wood properties showed conflicting results. No geo- graphic trend was found by Zobel and Rhodes in loblolly pine stands in east Texas, but a northerly and westerly trend was found by Zobel and McElwee in the southeastern states. In 1961 Zobel gtggl, reported their findings on the interrelationships of wood properties in loblolly pine. Methods The characters studied in the parental populations (Table 1) were principally those of the cone and seed because these have proven to be of value. Needle length and twist were also studied because they could be studied in the young progenies. Thirty-nine stands from 13 countries were sampled. In each stand one cone or one needle fascicle was collected from each of a number of young trees growing in full sunlight. The location of the 27 Table 1.--Description of cone, seed, and leaf characteristics measured on parental specimens of Scotch pine. Number Characteristics Unit of Measure P6 Cone length. cm. P7 Cone width, closed. cm. P8 Cone width, open. cm. P9 Ratio, cone length/cone width, closed. number P10 Ratio, cone length/cone width, open. number Pll Basal angle of the open cone. degrees (0 to 9 =o°to 45°) P12 Length of largest apophysis. mm. P13 Width of largest apophysis. mm. P14 Thickness of largest apophysis. mm. P15 Ratio, length/width of largest apophysis. number P16 Ratio, length/thickness of largest apophysis. number P17 Thickness of apophysis on opposite side of cone from largest apophysis. mm. P18 Asymmetry, ratio, thickness largest/ thickness of opposite apophysis. number P20 Ratio, cone length/length largest apophysis. number P28 Seed length. mm. P29 Seed width mm. P30 Ratio, seed length/seed width. number P35 Leaf length. cm. P38 Leaf twist. grade ( l = straight, 17 = 360 degree twist) 28 sample on each tree was standardized. The number of trees sampled per stand was usually 20 trees but in a few cases was as low as 10 or as high as 34. The period of collection for all countries except Spain was from August, 1960 to March, 1961. Spanish collections were made during the period December, 1961 to February, 1962. The 19 cone, seed, and needle measurements were sub- jected to analyses of variance using tree means, stand means and regional means to obtain a measure of variability within stand, between regions and within regions. The F-ratios are presented in Table 2. The components of variance are then determined from the same within-stand, within-region and between- region variances to determine the percentage of variability due to these variances. These percentages are shown in Table 3. The individual tree data for each stand were processed through "MISTIC," Michigan State University's electronic computer, to obtain means, standard deviations, and variances. The combined analyses of variance for each character was com- pleted by hand reprocessing the tree, stand and regional data output of MISTIC to obtain within-region and between-region variances for the species as a whole. In addition to Tables 2 and 3 the data are presented graphically in Figures 11 to 18 for a biometric comparison of the regions. 29 Table 2.--Significance of the variation between stands in the same region and between regions for various parental characters. Between Stands within Character regions regions P6 Cone length 27.80** 1.53* P7 Cone width, closed 15.20** 2.42** P8 Cone width, open 8.30** 3.70** P9 Cone length/width ratio, closed 3.49** 1.13 P10 Cone length/width ratio, open 3.87** 2.51** P11 Cone basal angle, open 3.09* 3.26** P12 Largest apophysis length 2.10 2.59** P13 Largest apophysis width 9.73** 2.75** P14 Largest apophysis thickness 9.76** 1.91** P15 Apophysis length/width ratio 6.76** l.85** P16 Apophysis length/thickness ratio 6.51** 2.51** P17 Apophysis thickness on concave side S.09** 3.96** P18 Index of cone asymmetrya 1.96 16.37** P20 Ratio, cone length to length of largest apophysis 21.86** 13.95** P28 Seed length 21.67** --- P29 Seed width 42.09** --- P30 Seed length/width ratio 5.76** --- P35 Leaf length 15.00** 2.59** P38 Leaf twist 1.41 --- w __.__ a See Figure 10 for explanation of measurement. * Significant at 5 per cent level. Greater than 2.27 or 1.50 for between-region and within-region comparisons respectively. ** Significant at 0.1 per cent level. Greater than 3.17 or 1.80 for between-region and within-region comparisons respectively. 30 Table 3.--Components of the variance in parental characters supplied by trees-within-stands, stands-within-region, and regions of origin. Per cent of total variance attributable to Trees Stands within within Between Character stands region regions P6 Cone length 7.5 .4 92.1 P7 Cone width, closed 8.8 1.2 90.0 P8 Cone width, open 10.7 2.9 86.4 P9 cone length/width ratio, closed 54.0 .7 45.3 P10 Cone length/width ratio, open 30.8 4.7 64.5 P11 Cone basal angle 30.6 6.9 62.5 P12 Largest apophysis length 22.2 3.5 74.3 P13 Largest apophysis width 12.0 2.1 85.9 P14 Largest apophysis thickness 16.5 1.4 82.1 P15 Apophysis length/width ratio 23.4 2.0 74.6 P16 Apophysis length/thickness ratio 18.9 2.9 78.2 P17 Apophysis thickness on concave side 16.3 4.8 78.9 P18 Index of cone assymmetrya 13.8 21.3 64.9 P20 Ratio, cone length to length of largest apophysis 1.1 1.5 97.4 P35 Leaf length 8.4 1.3 90.3 8See Figure 10 for method of measurement. 31 Regional Variation Pattern--Parenta1 Data Eight of the 19 characters (Table 1) studied proved to be the most definitive in separating the various stands into natural groupings. These are: P6. Cone length P7. Cone width, closed. P8. Cone width, open. P13. Largest apophysis width. P20. Ratio cone length to length of largest apophysis. P28. Seed length. P29. Seed width. P35. Leaf length. Only one of six ratios computed was of particular value in defining differences between stands or regions. In other words, shape of the organs remained relatively constant. The one ratio which proved of value was that of cone length to length of the largest apophysis. This ratio is essentially a measure of the number of scales per cone and reflects the increased number of scales per cone from north to south. The ratio of thickness of the dorsal apophysis to the ventral apophysis was used as a measure of cone symmetry (Figure 10). It proved to be too vari- able within stand and within region to be of value. The F-ratio for between-region variance in seed width was the largest of all the between-region ratios (F= 42.09, Table 2). Therefore seed width was one of the most definitive characters in developing regional patterns. 32 Figure 10.--Cone and cone scale measurements. Concave or ventral side Width ( ’ Length ._______1r_________. Thickness _AL__ o~ I‘- The method of measuring cones and cones scales on, a. open cone, b. closed cone, and c. cone scale. Basal angle is measured perpen— dicular to the cone axis. The largest apophysis on a reflexed cone is on the convex side of the cone and the opposite apophysis measured is on the concave side. The ratio of the two measurements is an index of asymmetry. 33 When seed width and length were plotted by regional group and latitude (Figurell) there was a clear north to south pattern. Northern populations had the smallest seeds and the populations in Spain and Turkey had the largest. The two stands sampled in Surrey, England (Nos. 269 and 270), did not conform to the general pattern. They had larger seeds than expected for their latitude. They are known to be of planted origin. Historical and progeny- test evidence indicates that the Surrey population is hybrid, probably between Scottish and German types (Wright and Bull, 1963) (Edlin, 1962). Possibly the larger seed is a manifestation of hybrid vigor. The Greek stand, an isolated population, had smaller seeds than expectedffor its latitude. The differences in seed width are sufficient to recognize a narrow-seeded Scandinavian population, a wide-seeded Spanish-Turkish- Yugoslavian population, and an intermediate French-German-Czechoslo- vakian population. Scotland is intermediate between the continental and Scandinavian groups. Spain and Turkey are areas of relatively light precipitation, generally averaging between 30 and 40 millimeters per month on an annual basis. The precipitation tends to be heavier in the growing season than in the winter, reaching a high of 40 to 60 millimeters in four of the five stands sampled in Spain and 74 millimeters in Turkey. Natural selection would favor the survival of the larger seeds which would have the germinative capacity to get roots down deep enough into the soil to survive through the first growing season and the subsequent dry winter. 34 Seed length does not present the same pattern as seed width. The Scotch, German (No. 253), French, Yugoslavian, and Creek are different from the Finnish populations, but the East German (Nos. 525 to 529) are not significantly different from the Swedish or Norwegian populations. The Spanish seeds are longer than all other origins and appear to belong to the group including England, France (No. 239), Yugoslavia, and Turkey. The patterns for the seed length/width ratio tends to parallel that for seed length, that is, the long-seeded Spanish and English samples also had the largest length/width ratios (Figure 11). Cone length and width, and length/width ratios are pre— sented graphically in Figures 12 and 13. Trees from north of the Arctic Circle had the smallest cones. Only one such stand was sampled. Hence it is not possible to say whether the nor- thernmost population differs significantly from the more southerly ones but that is a possibility. Also small-coned were the Greek and Yugoslavian parents, even though other trees from southern Europe (i.e., Turkey, southern France, and Spain) were large coned. The southern Scandinavian, German, Czechoslovakian, and Scottish parental populations were not separable on the basis of cone size. The two collections made in Surrey, England (Nos. 269 and 270), had the largest cones of all. Those were from planted stands. As previously mentioned under seed dimensions these 35 Figure ll.--Regiona1 variation pattern in length, width, and length-width ratio of Scotch pine seed. Scale length, cm. 5.5 4.0 4.5 5.0 Coun- Stand Lati-.Reg- width, cm. - ratio tgl‘ Ngabgrs tudg ign 1.5 2.0 2.5 5.0 o I I I I IIIII 226 67 A . . rnI 231-233 61-62 " SUE 545-545 59-60 c -. a, non 275-275 59-60 I \ sco 265-268 57 L 7- -’-.—- .' GER 525-529 50 G I 028 515-315 49 “T “‘5," \‘ -., ENG 269-270 51 I 4, :3, a, ‘. can 255: 50 H I I: ERA 2576.1: 49 T + I 334 239 45 I i a O, .....0 rue 242a,b 44 J .l "1.. \ GEE 2 \ 71 41 K . ° \. ,. \ SPA 218-219 40-42 N 245-247 ‘7“ ““3." ‘ ’0 O... TUB 2156,15 41 ,° L I - J I ‘ ‘ FINland, swsden, N0hway, SCOtland, GEhmany, CZEchoslovakia, ENGland, FRAnoe, YUGoslavia, GREsoe, SPAin, TUhkey. Seed length. oaooooooooooelo S..d 'idth. Seed length/width ratio. 37 Figure 12.--Regiona1 variation pattern in cone length, width, and length/width ratio as determined from closed cones. Scale Length, cm. 4.0 4.5 5.0 Coun- Stand Lati- Reg- Width, cm. - ratio tgya number; tude ion 2.0 245, 5.0 o l 7 1 PIN 226 67 A \ 2. rm 251-235 61-62 __- 5' sws 545-545 59-60 0 ". NOR 275-275 59-60 “T '-. sco 265-268 57 L 4“?“ ' I can 525-529 50 G .1... c2»: 515-515 49 “*5 \. 00...... ENG 269-270 51 I \, "I? .I GER 2 I rm 237l,b 49 —-l- E I FRA 239 45 m j I YUG 242a,b 44 J ._L 37'" I 0 Gas 271 41 K ( O. \. 0...... 524 218-219 40-42 N A“ -- .72: ‘ 245-247 I ‘0. I TUB 2156.6 41 ' 3 1‘ l J a FINland, SWEden, NOhway, SCOtland, GEnmany, CZthoslovakia, ENGland, FRAnco, YUGoslavia, GREece, SPAin, TURkey. Cone lengths 1..-nouns.» ClOBCd cone Width. ------- Cone length/closed cone width ratio. 39 Figure l3.-~Regiona1 variation pattern in cone length, width, and length/width ratio as determined from open cones. Sag}: length, width, on. 4.0 4.5 5.0 Coun- Stand Lati- Rog- ratio bsr t s on 1.0 1.5 2.0 o I T7 T FIN 226 67 A\ PIN 251-233 61-62 3'3 543-545 59-60 0 FOR 273-275 59-60 300 265-268 57 L - GER 525-529 50 023 313-315 49 ENG 269-270 51 I o" can 255 50 H FHA 23739b 49 FHA 239 45 H rue 2428,13 44 J GEE 271 41 x . soon-Loo...- SPA 210 219 40 42 11 ' “m.___.. ' 245-247 !. ‘ "' ' 0...... TUE 215‘9b 41 I o ‘f_ l L ._b._—o ‘ FIlland, SIEdon, NORway, SCOtland, GERmany, CZEehoslovakia, ElGland, FRAnoo, TUGoslavia, GREeoe, SPAin, TURkey. Cons length. IIIIIIIIIIIIIOI op.n con. '1dth. ........- Cone length/open cone width ratio. 41 trees may be hybrids and the unusual cone size may be a mani- festation of hybrid vigor. A hybrid origin is also a possible explanation for the unusually long cones from stand 233 in southern Finland. There is nothing in the origin data to suggest that this is other than a native stand. But pollination by a planted stand of continental provenance could have introduced other genes into the native forest. THE'peculiar nature of stand 233 was also noted in the progeny test. Large cones can either be an accommodation for more seeds or for larger seeds. The relatively large cones found in Turkey and Spain also contain the longest and heaviest seeds so that the adaptation is for larger seed in thkscase. Since the larger seeds have the best potential for survival, particularly in the semi-arid areas, this trend toward larger cones with larger seed in the Scotch pine endemic to these areas is possibly an answer to selection pres- sure. Cone width was measured in both the open- and closed-cone condition (compare Figures 12 and 13). Except for the cones from stand 239 in south central France, the variation pattern was the same for the two measurement methods. The error variance was least for the measurements made on closed cones. The ratio length/width of cone was of little value in the study of geographic variation. There was a tendency for the longer cones to be relatively narrower than the short cones. The apophysis is the raised portion of the cone scale which is visible when the cone is closed (Figure 10). Within a single 42 Scotch pine cone the size and shape of the apophysis varies within wide limits. It was thought that this measurement would be most meaningful if confined to the scale with the largest apophysis in the case of length and width or to the scale with the thickest apophysis in the case of thickness. Among stands within the same region there were differences in apophysis length. But this character was of little value in identifying the region of origin of a sample (Tables 2 and 3, Figure 14). The longer apophysis could be an indication of fewer seeds per cone if associated with greater apophysis width and short cones. However, apophysis length does not appear closely related to cone length or apophysis width throughout the range, suggesting a more ccmplex evolutionary pattern for the apcphyses from unknown factors, particularly in the highly variable southern limits of Scotch pine distribution. This may account for the lack of significance in the length of the apophysis when studied as a single factor. The ratio of cone length/length of largest apophysis was, however, of geographic significance (Table 2). Of all the characters studied, the percentage of the total variance attri- butable to between-region differences was highest for the ratio (Table 3). It is an indirect measure of the number of scales per cone,--the higher the ratio the higher the number of scales per cone. When the regional means are plotted by latitude, as in Figure 15, the ratio is seen to increase from north to south A 4 .3 Figure l4.--Regiona1 variation pattern in length and width of largest apophysis and the length/width ratio of the largest apophysis. Scale Coun- Stand Lati- hog- length, width, mm. - ratio IE!‘ numberg tuda ion 1.2 8,0 49,0 19.0 o I I 1 F1! 226 67 A FIN 251-255 61-62 SUE 545-545 59-60 C NOR 275-275 59-60 300 265-268 57 L ass 525-529 50 023 313-315 A 49 ENG 269-270 51 I GER 255 50 FHA 237a,b 49 F44 259 45 I rue 2428,b 44 J GEE 271 41 K 324 218-219 40-42 N 245-247 TUB 215a,b 41 l l L l ‘ FINland, swsden, NORway, SCOtland, GEhmany, CZEchoslovakia, ENGland, FRAnoo, YUGoslavia, GREeoo, SPxin, TURkey. — Length of largest apophysis. ..............- Width of largest apophysis. ------ Ratio length/width largest apophysis. 45 to a maximum in southern central France (Figure 15). Then it decreases and the southern populations (Spain, Turkey, Yugo- slavia, and Greece) have values similar to those from Germany and Czechoslovakia. The major exceptions to the general lati- tudinal trends were furnished by the English (larger than expected ratio), Yugoslav (smaller than expected ratio), and Spanish (larger than expected ratio) populations. Also, the previously mentioned Finnish sample from stand number 233 had an abnormally high ratio for its geographic location. The higher ratio of cone length to apophysis length would indicate that greater numbers of seed would be produced in Eng- land, West Germany, and France. This is an area of better soils and adequate moisture where survival of seedlings is not a problem but competition could be serious. Therefore greater seed quan- tities may be required. The decrease in the number of scales farther south would indicate more critical site conditions. In contrast to apophysis length, apophysis width had geo- graphic significance. So also did the length/width ratio for the apophyses. In neither trait, however, were there clear lati- tudinal or east-west trends. Perhaps the most noteworthy feature of the ratio was its small size for the south central French stand number 239, which had almost the widest apophysis of any sample. Thickness of the apophysis is of evolutionary significance in the genus,§igg§.according to Shaw (1914). He considered species with thick apophyses to be more advanced. Within the single species, Scotch pine, however, this characteristic had less geographic 46 Figure 15.--Regiona1 variation pattern in the cone length/ length of largest apophysis ratio. x; Coun- Stand Lati- neg- natio Jaw L0 5-5 6-0 o I I 1 III 226 67 A PI] 231-233 61-62 3'3 543-545 59-50 C '03 273-275 59-50 800 265-268 57 L GER 525—529 50 C23 513-315 49 ENG 269-270 51 I GER 253 50 FRI 257a,b 49 FHA 239 45 I rue 242a,b 44 J GEE 271 41 K SBA 218-219 40-42 N 245-247 TUB 215a,b 41 L l J FlNland, SWEden, HOhway, SCOtland, GEhmany, CZEchoslovakia, ENGland, Rhinos, YUGoslavia, GnEece, SFAin, TUhkey. Cone length/length of largest apophysis ratio. 48 significance than did cone or seed size (Tables 2 and 3). There was a noticeable tendency for this trait to parallel cone width; that is, for wide cones to have thick apophyses (Figure 16). Increased cone width and apophysis thickness could be a protective device that has developed in areas of more rigorous environments for Scotch pine. When the thickness of the largest apophysis was measured, an additional measurement was made of the thickness of the apophyses on the opposite side of the cone (character P17, thickness of op- posite apophyses, Figure 10). This gave a ratio which was a measure of asymmetry (character P18). Thickness of the opposite apophyses, however, closely follows the pattern of thickness of the largest apophyses (Figure 16) and the ratio of the two measures showed that there were no significant differences in asymmetry (character P18, Table 2). Another attempt to show cone scale variants was the use of the ratio of length of the largest apophyses over the thickness of the largest apophyses (Figure 17). Significant differences were obtained (Table 2). Shorter and thicker scales were found in the French, Turkish, and Spanish than in the other populations. The basal angle of the open cones (character P11, Figure 17) closely approximated the pattern of the ratio of length to thickness of the largest apophyses. There was a tendency for the trees with large basal angles to have short, thick cone scales. The Scandi- navian, Scotch, and Greek populations had the smallest basal angle and the Spanish and Turkish the largest. 49 Figure l6.--Regional variation pattern in thickness of the largest apophysis, thickness of the opposite apophysis (see Figure 10), and width of the open cone. Scale Coun- Stand Lati- heg- thickness, mm. - width, cm. trya numbers tune ion 2.0 2.5 5.0 1.5 o T T‘ 1‘ FT" FIN 226 67 FIN 251-255 61-b2 SW8 543-545 59-60 non 275-275 59-60 800 265-268 57 can 525-529 50 026 313-315 49 ENG 269-270 51 can 255 50 FRA 25789b 49 FHA 259 45 YUG 242a,b 44 HRS 271 41 SPA 218-219 40-42 245-247 TUE 215a,b 41 1L ll 1 .1 a FINland, SWEden, NOhway, SCOtland, Gasmany, CZEchoslovakia, ENGland, FRAnce, YUGoslavia, GhEece, SPain, TUnkey. .— Thickness of largest apophysis. mom-mm. Thickness of opposite apophysis. -----—- Closed cone width. 51 Figure l7.--Regional variation pattern in the basal angle of the cone, and the length/thickness ratio of the largest apophysis. Scalg degrees 10 20 5o 40 Coun- Stand Lati- Reg- - ratio trx‘ nggbgre tgdg ign 2.0 2.5 5.0 5.5 o I F f 1 PIN 226 67 A , FIN 251-255 61-62 ”ks 5'3 545-545 59-60 C *‘ “ -%3, -1; non 275-275 59-60 “a, 0..... 800 265-268 57 L ‘ r 1‘- ‘ far- can 525-529 50 023 313-315 49 ENG 269-270 51 I can 255 50 PEA 237:.b 49 FRA 239 45 I rUG 242a,b 44 J GEE 271 41 K 321 218-219 40-42 N 245-247 TUB 215a,b 41 l a PINland, SlEden, IOhway, SCUtland, GEflnany, CZEchoslovakia, ENGland, Rhinos, YUGoslavia, GhEece, SRAin, TUhkey. Length of largest apOphysis/width of largest apophysis. leeeeeeeeeeeee‘ 38.81 angle Of cone. 53 In extracting seed from the cones it was quite noticeable that the cones which opened the widest (small basal angle) also permitted the seed to be extracted moreeasily and quickly. Cones with large basal angles retained the seed in the lower half of the cone. The heavier scale and larger cone with the greater opening of the scales was more efficient in seed dispersal. This condition coincides with less favorable environmental conditions for seedling development. Shaw (1914) states that the heavier cone and scale is a higher evolutionary development. However, it was also noted in some of the northern stands, particularly in Norway, that there was an occasional serotinous or partially serotinous cone, also an indication of a higher development. Leaf length (character P35, Figure 18) increased from the Arctic Circle to England, West Germany, and eastern France. Then it decreased and was short in trees from Greece and south central France. Stover (1944) stated that leaf length in conifers was longest under mesic habitat and shortest under xeric conditions. Since growth would also be greatest under mesic conditions leaf length should be a measure of growth rate of the tree. The gen- eral pattern of leaf length does follow the growth rate of Scotch pine. The Scotch pine of the palatinate region of Ger- many has been noted for the quantity of wood which it produces. The English population is the result of natural reseeding by what appears to be hybrids showing hybrid vigor. If the above premise is correct, the English population should also have a 54 Figure 18.--Regiona1 variation pattern in leaf length and twist. \41 Scale . number 0 5 1O 15 Coun- Stand Lati- Eeg- on. tr!“ number; tgde ion 4.0 25.0 6.0 1.0 o T 1 I j III 226 67 A FIN 231-233 61-62 SUE 545-545 59-60 C 303 273-275 59-50 300 265-268 57 L GER 525-529 50 G 033 315-515 49 ENG 269-270 51 I f can 255 so 3 1a IRA 257.,» 49 ,° FHA 2 3 9 4 5 I ..eeeeee rUG 242a,b 44 .1 “"»§, 0.. GhE 271 41 K .«“ 3:1 218-219 40-42 x ' In :3. -- 245-247 ,2’ TUB 215a,b 41 if e—l—a I J J, I ‘ FINland, SIEden, NOBway, SCOtland, Gas-any, CZthoslovakia, ENGland, FRAnee, YUGoslavia, GREece, SRAin, TUhkey. — LOSf length. aneceeaeeeeee. LOaf t'iste 56 high growth rate in comparison to other geographical races of Scotch pine. The measurement of leaf twist (character P38, Figure 18) proved to be of little value except to indicate that leaves from English and Turkish populations have less twist per unit length than other.populations. The Naturalness of the Regional Grouping One of the initial objectives of the study of parental characteristics was to determine if population discontinuities existed, where they occurred, and which characters would prove most valuable in defining populations. The original parental data were grouped by natural region of origin in such a manner that the regions were as homogeneous as possible with regard to morphological characteristics. The final grouping is very similar to that used by Wright and Bull (1963) for progeny test data, the major exceptions being the separation of the Czechoslovakian from the German and the TUrkish from the Greek populations. For the analysis of variance the data for stand 226 in Finland (region A) and stand 271 in Greece (region K) were discarded because there was only one stand per region. Data from 10 trees in each stand were used. The degrees of freedom for the analyses were as follows: between regions-~8, between stands within regions--30, between trees within stands-- 351, total-~389. 57 When grouped in this manner, between-region variances accounted for 62 to 97 per cent of the total variances in the various characteristics. Stands-within-region variances gen- erally accounted for less than 5 per cent of the total vari- ances, the exception being the index of cone symmetry (charac- ter P18 in Tables 1, 2, and 3). The large between-region differences indicate that the regional grouping was natural. Furthermore, they indicate that it is possible to tell the region of origin from cone and leaf specimens. Perhaps the variation is too great to permit this on the basis of cones or needles from a single tree, such as might be found on an ordinary herbarium specimen. But, if mass collections are made from several trees in a stand, the stand as a whole can be characterized sufficiently to identify its place of origin. Turesson (1922) stated that an ecotype is one of a series of relatively discrete, genetically differentiated, natural enti- ties whose limits coincide with the distribution of some environ- mental effect. Huxley (1939) said that a cline is a geographical gradient in a phenotypic character with continuity and a genetic gradient implied. Langlet (1959) reviewed the concept of con- tinuous variability in his discussion of Wright and Baldwin's (1957) report on New Hampshire plantings of the 1938 IUFRO Scotch pine provenance test. He stated that "Discontinuity may thus very well occur-~where the conditional ecological factors vary discon- tinuously." 58 Stebbins (1950) in discussing geographical races stated "There is no doubt that in plants, as in animals, many species may be divided into races or groups of genetic types which are adapted to the different parts of their ranges, and that these sub-divisions are separated from each other by partial discon- tinuities in the variation pattern." Mayr (1942) was less flexible in his definition of a geographical race stating that such a race was "A complex of interbreeding and completely fertile individuals which are morphologically identical or vary only within the limits of individual ecological and seasonal variability. The typical characters of this group are genetically fixed and no other geographical race of the same species occurs within the range." The analyses of variance based on the sample of 39 stands indicates that regional discontinuities exist after removal of the within-stand or within-region variability. The use of the word ecotype has been avoided as being applicable to a certain situation and the term geographical variety has been used to describe a population which is different although there may be a continuity in the characteristic as it merges with another geographical area. Ecotypes, as defined by TUresson, do exist in Scotch pine where there are sharp breaks in the factors which have produced them. Glaciation disrupted the original continuity of Scotch pine during the complex patterns of glaci- ation in Europe and Asia according to Hulten (1937, 1949). The movement of Scotch pine back into areas of original distribution (.0 59 following the shrinking of the glaciers once more established continuity in distribution but not genetic continuity. Certain populations have remained isolated, particularly along the south and east of the area of distribution of Scotch pine. Langlet's insistence that the variability of Scotch pine can be called only clinal is well authenticated by other authors who have studied the species. The concept of continuity is not disputed in the case of continuous populations under continuous meteorological or edaphic conditions. The sampling upon which this study is based is too meager to prove discontinuities over such a wide range as that covered by Scotch pine. It does prove that if population samples are taken at discrete locations within the range that there are identifiable geographic races which pre- dominate at the location that also are genetically different from other samples taken at other locales. The range of Scotch pine covers a wide area of complex geographical and meteorological conditions that are often quite well defined. Even in the less well defined and less abrupt gradients differences have been found between populations which are of significant importance to a tree breeder, much as a superior tree within a given stand. However, there is an im- portant difference, a superior tree in a stand represents an improvement in some desirable character or combination of characters that are identified with an individual tree, an ideal situation only if vegetative propagation is planned. 60 Selection of a segment of a population permits the acquisition of sufficient breeding material without seriously restricting the gene pool required for a breeding program. Many authors who have reported their observations of Scotch pine have pointed out that although there are recognizably different varieties of Scotch pine these are contained in highly variable populations in which they predominate. In this meaning, a geographical variety is recognized here by the predominance of a characteristic which arose in answer to selection pressures based on local environmental conditions. The between-region differences have been summarized by character in Table 4. Finland north of the Arctic Circle (region A) is repre- sented by a single stand which is significantly different at the one per cent level from the southern Finland, Sweden, and Norway population (region C) in open-cone width, and seed and leaf length. The regions are significantly different at the five per cent level in cone length, and the cone length/length of largest apophysis ratio. Southern Finland, Sweden, and Norway are significantly different from the population of Scotland (region L) at the one per cent level in the open cone length/cone width ratio, width of the largest apophysis, and the length and width of seed. The regions are significantly different at the five per cent level in cone length and the cone length/length of largest apophysis ratio. l Table 4.-SuI-sry of significant parental character differences between regions. Stand. Country N. Re- 6,8,20,32,32 Finland, Sweden, Norway gion C L 8 i 3.3.1 23H3 ‘3Jal 210:] -N :flafi ens-N- no' ‘0 N 0 H . 8 >. O E 3 I U Czecho- slovakia I H 6]. 0,13,16,17 g; l. 18 3g 10,14,16,20 29,35 J 20,35 8.28.32.§ 6.9.335 NE.Greece 20,28,29 K 10,13,20,28, 29 18.39,. Central IgSee Table 1 for description of characters. N T Underlined characters significant at l per cent level, others at 5 per cent level of significance. 62 Scotland is significantly different from the East German-- Czechoslovakian population (region G) at the one per cent level in the open-cone length/width ratio, basal angle of the open cone, width of the largest apophysis, asymmetry, and the seed length/width ratio. These regions are significantly different at the five per cent level in cone length, thickness of opposite apophysis, and the cone length/length of largest apophysis ratio. The Scottish population is intermediate between regions C and G but not different in cone width, leaf length, and leaf twist. The cones, leaves, and seed of the Surrey, England, popu- lation (region I) exceeded the length of all others except for the Spanish seed. The English population was significantly different from that of the East Germany and Czechoslovakia in 13 characters and from the West German, eastern France in 10 characters. The population in eastern France and West Germany (region H) differed significantly from that in East Germany and Czecho- Slovakia in 10 characters and from the Central Massif of France in 4 characters, all at the one per cent level of significance. Region M, the Central Massif of France, represented by a single stand, differed from the Spanish (region N) population in the open cone length/width ratio, width of the largest apophysis, and the cone length/length of largest apophysis ratio at the one per cent level of significance, and in seed length and width at the five per cent level. 63 The Greek population, represented by a single stand in the mountains northeast of Drama near the Bulgarian border, was different from the Turkish population, represented by two stands from the mountains of north central Turkey near the Black Sea, in 12 characters, and from the Yugoslavian popu- lation represented by two stands, in 7 characters. The Greek population, however, differed in only two characters, cone length/length of largest apophysis ratio and leaf length, from the East German, Czechoslovakian. The Greek stand dif- fered in only four characters from region C and L, and in three characters from region A. The differences between regions outlined in Table 4, combined with the data presented in Figures 11 to 18, not only define the variability of the Scotch pine in Europe and Asia Minor, but permits comparative analysis of herbarium specimens and placement in the region of origin. The eleven regions defined are all separated by significant differences in their cones, seeds, and leaves except for regions C, L, and G, where there were no differences in leaf length. The parental data supports the regional patterns based on total similarities in juvenile characters reported by Wright and Bull (1963) except for the Turkish population. However, the data presented by Wright and Bull in their table of similarities and differences for region K does indicate a possible separation of the Turkish-Georgian populations from that of Greece. 64 Staszkiewicz (1960, 1961, 1962) studied the cones of Scotch pine populations from Finland, Sweden, Scotland, Czecho- slavakia, Hungary, Poland, Switzerland, and the Central Massif of France. He separated the populations on the basis of ten cone measurements into seven regional varieties having different types of cones as following: var. lapponica, northern Finland. var. suecica, Sweden. var. scotica, Scotland. var. polonica, Poland. var. subcarpatica, northeastern Czechoslovakia. var. meridionalis, Czechoslovakia, Switzerland, and Central Massif of France. Comparisons between his and my data show general agreement with regard to the first five varieties. They are separable in both cases. 0n the other hand, the population which he regards as var. meridionalis and as homogeneous, is very heterogeneous according to the present study. In a later chapter, these data on phenotypic varability patterns will be coordinated with past taxonomic treatments to give a recommended series of varietal names. Sample Size The specific question of how small a sample can be measured without appreciably changing the results is important. Staszkiewicz (1960) reported that his original sample size was 100 cones picked under 100 trees, or one cone per tree from 100 trees per stand. In 65 some instances he collected 2 cones per tree. Since the measure- ment of so many samples was laborious he halved some of his samples and found that the data obtained from sample sizes of 50 did not exceed the permissible error. Critchfield (1957) used a sample size of 5 cones and 5 needles per tree for 12 trees per stand. Thor (1961) used 20 sound and mature cones and 5 twigs from 15 trees in his study of variation patterns in natural stands of loblolly pine. Thor (1964) took two core samples from each of 15 trees per stand in his wood property study in Virginia pine. In the course of analysing the 19 characters utilized in this study correlations were run between the standard deviations of each sample and the size of that sample. The samples ranged in size from 10 to 34 cones (l cone per tree). There was no correlation between size of sample.and of the standard deviations. This shows that the disparity in sample sizes was unimportant and that one cone per tree from each of 10 trees would have been suf- ficient. If, in the present study, cone length and needle length are used to determine the actual size of an acceptable sample, the following procedure can be used: Stand 226 (Finland) was represented by a collection of 29 cones taken from 29 different trees. The analysis of variance for cone length produced the following information: 3(- 3 3.567 centimeters (cone length) 32.. .268 66 S = s51? 331 = .096 The 95% confidence interval for the mean== : (t 95) (S ) ' if = (2.048) (.096) = .1966 100 (t.95) (Si) _ 100(2.048) (.096) _ 5 57 Sampling error 3? “ 3.567 Assuming that the permissible error is 10 per cent of the mean, the confidence interval is: .10 (i) = .10 (3.567) = .3567 Since the confidence interval (.3567) is equal to (t 95)(Si) we can solve for S— =.;§§§Z = .158 X 2.26 The number of sample plots needed can be computed using the formula: 2 s. =.__.§_____. or n = S =,_;Z§§ 2 = .268 = llcones. x n— 5;?— (.158) 76273 Based on the cone-length data, sample size ranged from 3 to 14 cones for 9 different stands. The computed sample sizes for these 9 stands were 11, 7, 6, 4, 10, 3, 5, l4, and 11 cones. Therefore cones from 10 trees is not only a good average sample, but a sample sufficient for nearly any stand. At the 95 per cent confidence limit, the sampling error with regard to needle length was 10.4 per cent from a lS-tree sample. In other words, needles from 16 or 17 trees would have been required to give the desired 10 per cent sampling error. Using the formulas given previously on needle length data, the numbers of needles needed to give that sampling error for eight different stands were found to be 30, 9, 18, 19, 7, 12, 15, and 4. 67 In other words, needles from 15 or 20 trees would be needed to give the same accuracy as can be achieved with cones from 10 trees per stand. The specifications for needle and cone samples for the study called for one needle and one cone per tree from 10 to 20 trees per stand over an area which would minimize the chances of obtaining seed from two trees that had been pollenated by a single parent. Some cooperators sent in two cones and two needle fascicles per tree. A comparison of the results of measuring two cones and four needles per tree and one cone and one needle revealed no significant difference between the sampling. Samples based on single cones and leaves had samp- ling errors of the same magnitude as samples based on two cones and two needles. In other words, if cones or leaves are collected in a standard manner, within-tree variation can be disregarded. There were some intra-regional differences which should be noted, particularly in region G which included northeast Germany and Czechoslovakia. The north to south variation pattern here is apparently complicated by an east to west dif- ference due to geographical and meteorological differences. Seed length and width, and needle length follow the established trend of small to large from north to south but open cone length and width of the Czechoslovakian populations are smaller than that of the northeast German (Figure 19). Although the Czecho- slovakian material falls within the range of the German it does 68 not show the high variability of the German and is centered at the low end of the scale. Origin 233 in east Finland repre- sents another stand sample which is materially out of line with the remaining populations sampled in southern Finland. The cones of 233 are of significantly larger size but the seeds are normal to that region (Figure 18). 69 Figure l9.--Intra-regional variation pattern for region C, southern Finland, Sweden, and Norway, and region G, northeast Germany and Czechoslovakia. Scale Coun- Character try 200 300 4%9 500 1 ' T I P6 Cone FIN ‘h—d233 length SWE ~‘ NOR " CZE nan P8 Open- FIN '—=—4 cone SWE in width NOR ‘“' GER I—Iha CZE «It P28 Seed length FIN "4 SWE has NOR .u_n_s GER ‘-‘ CZE h“ P29 Seed width FIN In SWE "‘ NOR the GER use CZE add J . L I I 1FINland, SWEden, NORway, GERmany (northeast Germany), CZEchoslo- vakia. CHAPTER V RELATIONS INVOLVING SINGLE RARENTS AND THEIR PROGENY Materials and Methods A series of half-sib progeny tests was started in 1959 with Scotch pine seed received from eight stands in northern Europe (Table 5). Two of these stands are in Belgium and were sampled through the courtesy of Dr. Alain de Jamblinne (parents 285-304), Chief, Section of Genetics, Centrum voor Bosbiologisch Onderzoek, Bokrijk-Genk, Belgium. In addition to the stand data the collectors sent parental data on height, age, and diameter for nine trees which were progeny tested in Belgium. A stand in southern Norway was sampled by Dr. Tollef Ruden, Norwegian Forest Research Institute, Vollebekk, Norway, who also furnished information on the site characteristics and age and height of the parent trees. The sampling of the East German stands was done by Dr. Otto Schchk, Director of the Branch for Forest Research at Waldsievers- dorf of the German Academy of Agricultural Science of Berlin, Institute for Forest Science, Eberswalde. Dr. Schchk furnished data on many characteristics of the parental trees including age, height, diameter breast high, stem form, crown length, length of clear stem, crown diameter, flatness, and form, and foliage condition. 70 71 In each of the eight stands open-pollinated seeds were collected from either 10 or 20 randomly selected dominant trees. In general, the parents in any one stand were 100 feet or more distant from each other. Thus they probably did not have common female parents although they may have had common grandparents. After receipt in late 1958, the seeds were weighed and placed in cold storage. They were sown in the Bogue Forest Research Nursery on May 13 and 14, 1958. A variation of a randomized complete block design was used. The 10 or 20 seed- lots from each stand were randomized and grouped together. These progeny-groups were then randomly distributed (along with the stand-progenies from the geographic origin test) through each of the four replicates. This arrangement resulted in greater precision for the detection of within-stand than between-stand differences. The seedlings were grown for two years at an approximate spacing of 50 per linear foot (25 per square foot). Then they were thinned to an approximate spacing of 20 per linear foot (10 per square foot). They were partially lifted at the start of the third year and completely outplanted by the start of the fourth year. The European cooperators measured metric characters (height, etc.) to an accuracy of approximately one-twentieth of the range between extremes. They described other traits in words which were translated to numerical codes for purposes of analysis (Table 6). The linearity of these translated numerical grades is not known. 72 Asa ens ma w .muaov .umaa xooummsuumnsm own venom Hmunumz 111 ON oHH we .Hmsumawnomoh .wzem .wzazamo .m on NH an em -Hmm cowumucmaa vHo 1umom1oo~ hands: mama so om wuanawq son 6 Beam mwcfiHvoom Homeoe couH oH 1mm mo .Hmunoom .ZbHuAmm Hm m no Hm 1mmu camauo :Bocx HHOmnsm momma -e: .easum euussnm .wop-aamm On mm mm massage «mm Hmeua .zaHoamm on m an an -mmm ooH emu venom Honouwz 111 CH 1omH CNN vcmaaom .z .M<3moz an HH on an 1m~N Hanan: mummw muouma . 8 . o mxumamm mama Hwom eoamwmw JMMd .>0Hm muwawooH use huucsou «wwoq .usq .oz momma venom .umsm euuoz venom avenue unused mo cowuafiwomon11.m manna 73 suaouw ummm mmmwaom h>mo= =3ouo venom caouo umHm eaouu ovwz sumaoa waoa nuwcoa uuonm sous uanmuum nouoamgo owned :u3ouw numb nusouw 30am mmmHHom cHHH cacao vocam1mso cacao moueaom :3ouo souumz nowaoa uuonm nuwcoa umouo mo>uao ham: Houseman Hausa nuaouw Son munch aw owe assume a“ unwwon owumm mums» m on H avenue m on H seesaw m as H seesaw “~0qu Hmumz Munoz m on H «cease Houuawuaou Roam: was» euaono on sea 3 :oHuHeeou owsHHoa we wanna cacao he waHaouusHm nacho as umuoamwm c3ouo me Baum ummao mo summon «a :uwcoH caouo o>wq me Show Boom Na swan ummmua Houseman He semee Hence os on emewwmmu o=Hm> newsman ou moawwmmm 05Hm> umoaoq uceaouamsmz mo owe: Manama was nouosumno seesaw one muouowuwno Hmuamuma mo cowuawuomoO11.o manna 74 The seedlings were measured or scored periodically for all macroscopic traits in which preliminary examination showed the probable presence of between—progeny differences (Table 7). Measurements were made to an accuracy of approximately one- twentieth of the range between extremes. Non-metric characters were scored by numerical grades. In order to avoid the need for transformation the grades were so defined as to result in normal distributions and linearity. The details of progeny performance were described by Wright (1963) who performed an analysis of variance for each trait and each group of progenies from a single stand. In those analyses plot means were used as items. For the present paper all possible simple (product- moment) correlations were calculated among parental and progeny characters. values applicable to single parents or to progeny means were used as items. When analyzed in this manner, the word-interpretation of a significant relationship such as reported between parental character No. 44 and progeny character No. 22 is as follows: the offspring of trees with long clear boles had fewer lateral buds in May, 1960, than did the off— spring of trees with short clear boles. Because of the relatively small number of significant parent-progeny correlations it was deemed inadvisable to use multiple correlation or factor analysis. The correlations were calculated by MISTIC, the former Michigan State Integral Computer. 75 Table 7.--Description of juvenile characteristics and grades used in scoring Scotch pine progeny study seed wt. Character Unit of Lowest value Highest value and number Date measure Range assigned to assigned to 5 Seed weight 11/58 mg. 2.9-12.2 Light seed Heavy seed 6 Seed length 11/58 mm. 3.4-5.9 Short seed Long seed 7 Height, age 1 10/59 mm. 37-131 Short tree Tall tree 8 Height, age 2 7/60 mm. 57-387 Short tree Tall tree 9 Foliage color 6/59 Grade 4-8 Yellow-green Green 11 Foliage color 9/59 Grade 8-20 Red-maroon Green 12 Foliage color 10/59 Grade 8-20 Maroon Green 13 Foliage color 8/60 Grade 8-16 Green Dark-green 15 Foliage color 11/60 Grade 4-32 Yellow-green Dark-green l6 Foliage color 12/60 Grade 4-32 Yellow-green Dark-green 18 Bud color 5/60 Grade 4-24 Bright-green Brown 19 Time of bud 9/59 Day of 200-269 Early set Late set set year 2? Bud diameter 4/60 mm. 1.5-6.4 Small bud Large bud 22 Branched buds 5/60 Per cent 0-13 Small number Large number - of trees 23 Branched buds 10/60 Per cent 0-5 Small number Large number of trees 26 Presence of 5/60 Grade 0-12 None present Present on primary lvs. 3" of stem 27 Presence of 9/59 Grade 0-4 None present Present on 50% secondary lvs. of seedlings 28 Presence of 10/59 Grade 0-24 None present Present on 90% secondary lvs. of seedlings 29 Leaf length 8/60 cm. 25-43 Short needle Long needle 7/5 Ratio l-yg.ht. 5/60 Number 8-26 Low ratio High ratio 76 Parent-Parent Correlations The simple correlations among the parental characters in the five German stands are presented in Table 8. Because the stands were even-aged, total height and height growth rate are identical; relations involving one also involve the other. For the 9 traits and 5 German stands, a total of 180 simple correlations were calculated. Actual and expected numbers of cor- relations significant at various levels were as follows: Significance level 0.1 per cent 1 per cent 5 per cent r needed for significance .68 .56 .44 Number of r's exceeding amount needed for significance Actual 7 19 36 Expected .18 1.8 9.0 Approximately 25 per cent of the total significant correlations and 50 per cent of those for which .05 > P > .01 are presumed to be meaningless. Presumably the six correlations significant at only the 5 per cent level and for only one stand have little meaning. Thus clear-stem length, crown form (one-sided or round), and foliage condition were essentially uncorrelated with the other variables. The 36 correlations, which were above the level needed for significance at the 5 per cent level were distributed among the five stands as follows. 77 Table 8.--Correlations between parental characters in five East German stands. With 18 degrees of freedom for each stand the value of r needed for significance at the 5, l, and 0.1 levels was .444, .561, and .679 respectively. 78 Character Stand Character 11' 49. 41. Diameter Ravershagen '* {3 breast high Neustrelitz * 3:3 GUstrow NS 3 u Nedlitz NS 5 g Joachims thal NS 22 j; l. 42. Stem form RBvershagen NS .;* a Neustrelitz NS NS 3 GUstrow NS NS ‘“ Nedl 11:2 NS NS 5 J oachims thal NS NS 3'; 42. 43. Crown R3vershagen NS NS ** length Neustrelitz NS * NS Glis trow NS NS NS E '5 ,1: Nedl 1:2 *** * NS o g“ to Joachims thal *** NS NS 5 £3. 5 9.3.. "‘ 44. Clear-stem RUvershagen NS -* NS NS 5 length Neustrelitz NS NS NS NS : Gustrow NS NS NS NS H Nedlitz NS NS NS NS 8 Joachims thal NS NS NS NS '3 S U 4&5 o 45. Crown RBvershagen NS *** ** * -** .§ diameter Neustrelitz NS ** NS NS NS 'o GUstrow NS * NS NS NS g on Nedlitz NS ** NS NS NS 3 .5 Joachimsthal NS * NS NS -* cs 5 45. U 46. Crown RBvershagen NS ** NS ** -* ;;* [E flattening Neustrelitz NS NS NS NS NS *** ‘H GUstrow NS ** NS NS NS *** E Nedlitz NS ** NS NS NS *** a J oachims thal NS ** NS NS NS ** 0 5.19- a 47. Crown shape R8vershagen NS NS NS * -* NS * ‘g Neustrelitz NS NS NS * -* NS * m Gustrow NS NS NS NS NS NS NS g Nedlitz NS NS NS NS NS NS NS 8 Joachimsthal NS NS NS NS NS NS NS 9’ 47. 48. Foliage RBvershagen NS NS NS NS NS NS NS NS condition Neustrelitz NS NS NS NS NS NS NS NS GUS trow NS NS NS NS NS NS NS NS Nedlitz NS NS NS * NS NS NS NS Joachimsthal NS NS NS NS NS NS NS NS 79 Correlations greater §£§g§ than r = .444 vaershagen 15 Neustrelitz 7 Gflstrow 3 Nedlitz 6 Joachimsthal 5 Total 36 There was nothing in the origin data to account for the greater amount of intercorrelation in the R8vershagen stand. Nor was it accounted for by a greater range of variability in the measured traits; the other stands were just as variable. Bole diameter, crown diameter, and crown flattening were strongly inter-correlated in each of the five stands. Trees with large boles had large-diameter crowns which tended to be flat rather than pointed. These relations probably reflect cause-and- effect relationships. The large crowns are needed for rapid diameter growth. Also, large crowns are a sign of approaching maturity which is heralded by a flattening of the crown. Crown length was strongly correlated with total height in two stands. It was weakly correlated with bole and crown diameter, stem form, crown flattening, crown shape, and foliage condition in one or two stands. In the R8vershagen stand, length of clear stem was inversely correlated with bole and crown diameter, crown pointedness, and greater crown growth on one side than on the other. 80 Total height and diameter were weakly correlated in two stands, and were not correlated in the other three. This is contrary to what would normally be expected. It indicates that the seed collectors may have tried to select unusual trees which departed from.the normal behavior pattern. These results for the five German stands are in partial agreement with the results of previous investigators. Zieger (1928) (cited in Spurr (1948)), working with Scotch pine, found bole and crown diameter to be strongly correlated. Ilvessala (1950) (cited in Spurr (1960)), also working with Scotch pine, found that bole diameter could be estimated from crown length and total height. The standard error of estimate was about 10 per cent, less than if only one variable was used in the regression. Willingham (1951), working with longleaf pine (P. palustris Mill.) found a significant simple correlation (r ==.51) between total height and bole diameter. The correlation was much stronger (r = .82) if bole diameter was treated as a function of two independent variables--total height and crown diameter. Data from 9 parent trees in two Belgian stands and from 10 parent trees in the Norwegian stand were available for analysis. In each case, the trees were of differing ages so that the ratio total height (character 50) was considered as a measure of height age growth rate. Correlations were calculated between total height, height growth rate, bole diameter, and age for the Belgian material; between total height, length of clear stem, age, and height growth 81 rate for the Norwegian material. None were significant, probably because of the smallness of the samples. What do these correlations mean in terms of plus-tree selec- tion? Most of the answers must, of course, be applicable to the German stands for which the most data are available. First, there was a high loading of bole diameter, crown diameter, and crown flattening with a common diameter-growth or maturity factor. Selection for any one of the three characters would mean selection for that factor and the other two characters. In these five stands it would not be advisable to select for large diameters and pointed crowns. Second, there were several inter-correlations in the Raver- shagen stand so that selection for greater bole diameter would be accompanied by selection for greater total height and less clear bole length. Selection for greater crown diameter would also be accompanied by selection for greater bole diameter, fewer stem crooks, greater live-crown length, less clear stem length, and greater crown flattening. Regardless of whether or not these relationships had a common causal factor, they are present in that particular stand. Attempts to reverse the selection trend against one of the associated characters would be successful only if a very large number of trees were sampled. Third, in the other four stands there were relatively few inter-correlations. This means that in them selection for one trait will have relatively little effect on other characters. The presence of significant correlations is sometimes inter- preted as favoring indirect selection, for example selection for 82 large diameter to achieve a gain in height growth. When prac- ticing such selection, the rate of progress in the correlated character is r2 (r2== coefficient of determination) as great in the correlated character as in the character for which selec- tion is practiced. For the five German stands, the frequency of coefficients of determination was a follows: Number of correlations Range of r 145 below .2 ll .2 to .3 10 .3 to .4 6 .4 to .5 3 .5 to .6 3 .6 to .7 2 .7 to .8 O .8 to 1.0 Thus there were 7 cases in which 50 per cent as much progress could be expected by indirect as direct selection. Progress by indirect selection would be very slow (less than 20 per cent of maximum) in 145 cases. Parent-Prgggny Correlations The most impressive fact obtained from this portion of the individual tree study is the exceptionally low number of correlations between parental and progeny characters and the complete lack of a pattern in the correlations. In considering the 11 parental characters (only 4 for the Belgian and Norwegian stands) and the 20 progeny characters for the 7 stands under consideration there were 1,260 possible simple correlations (Tables 9-15). The actual and expected correlations significant at various levels were as shown on page 90. 83 Table 9.--Parent-progeny and progeny-progeny correlations for nine Belgian trees from two stands. Only characters involved in one or more significant correlations are included. PROGENY CHARACTERS a 8 f; to 0.88 6 S\3 8 In\\ 01o tnt: c: \v-IN mom 0 O HNOHH \‘U 00 U u r: -o as s 16 s m sass:ss.s 9.2.: M “Scum—114.40 é-u 0% up“: 010 ooo~mmm>~0\ oo 3 p114 u <3 0 o o o *1 a: in £01 :14 J35 U '00 0) U 'U‘UOOOO‘H'H‘H g figm-m. 33:36:13.3: as“: mmmmASAmHmmgm-HSH 5 6 7 8 9 15 16 18 19 21 23 26 27 28 PARENTAL CHARACTER 40. Total height NS NS NS NS * NS NS NS NS NS NS NS NS NS PROGENY CHARACTERS 6. Seed length ** -- -- -- -- -- -- -- -- -- -- -- -- -- 12. Leaf color 10/59 NS NS NS NS * -- -- -- -- -- -- -- -- -- 15. Leaf color 11/60 * NS NS NS NS -- -- -- -- -- -- -- -- -- 16. Leaf color 12/60 * NS NS NS NS * -- -- -- -- -- -- -- -- 18. Bud color 9/59 NS NS * NS NS NS -* -- -- -- -- -- -- -- 19. Time of bud set NS NS NS NS NS NS NS ** -- -- -- -- -- -- 21. Bud diameter 5/60 NS NS * NS NS NS NS NS NS 22. Br. buds 5/60 NS NS NS NS NS NS NS NS NS NS -- -- -- - 23. Br. buds 10/60 NS * NS * NS NS NS NS NS NS -- -- -- -- 27. Sec. lvs. 9/59 NS NS NS NS NS NS NS NS NS * NS NS -- -- 28. Sec. lvs. 10/59 NS NS NS NS NS NS NS NS NS NS NS -* ** -- 29. Leaf length * NS NS NS NS NS NS NS NS NS * NS NS NS 7/5. Height age 1 -* NS * NS NS NS -* * -* NS NS NS NS * Seed weight *‘r = greater than 0.67 needed for significance at 5 per cent level. ** r = greater than 0.80 needed for significance at 1 per cent level.‘ NS Correlation not significant (a)Parents and progenies analyzed were numbers 286, 288, 292, 294, 297, 298, 301, 303, and 304. 84 Table 10.--Parent-progeny and progeny-progeny correlations for eight Norwegian trees from a natural stand in southern Norway. Only characters involved in one or more significant correlations are included. a PROGENY CHARACTERS y Y o g :1 as 3‘. Q 8 S 1:: '2 1d as F1 a) _, 01 HS; m u 00‘ u ,5 1n 0 “U was mus .fl U 0 H H H H \ ”I 3 031!) W\ 00 no 00 O O O O a 001.0 \ o -H a m F1 F1 —1 14 Bus max o~1 m o O O O O U m H U o 3 1-1 1.! o o o 0 cu ~41: 16 {1:01:01 .G m '610 u 810 010 'U 'U a: $1 $1 41 41 LI 0:) m:> m:> a) a) -.-1 m as a: m 'U '35 a: fig 38 1% 1% :g :3 13 13 13 lg #13 figuxlh-d and 5 6 7 11 12 13 15 19 21 22 27 28 RARENTAL CHARACTERS 44. Clear length NS NS NS NS NS NS NS NS NS -** NS NS 50. szannual growth NS NS NS NS NS NS NS NS NS -* NS NS PROGENY CHARACTERS 6. Seed length ** -- -- -- -- -- -- -- -- -- -- -- 7. Height age 1 * NS 8. Height age 2 NS NS ** -- -- -- -- -- -- -- -- -- 11. Leaf color 9/59 NS NS * -- -- -- -- -- -- -- -- -- 12. Leaf color 10/59 NS NS * *** -- -- -- -- -- -- -- -- 13. Leaf color 8/60 NS NS NS * NS -- -- -- -- -- -- -- 16. Leaf color 12/60 NS NS NS *** *** * NS -- -- -- -- -- 27. Sec. lvs. 9/59 -* -* NS NS NS NS NS NS NS NS -- ~- 28. Sec. 1V8. 10/59 NS -* NS NS NS NS NS NS -* NS * -- 29. Leaf length 8/60 NS NS NS NS NS NS NS * NS NS NS NS 7/5. Height age 1 -* -** NS NS NS NS -* NS NS NS * * Seed weight * r = greater than 0.707 needed for significance at 5 per cent level. ** r = greater than 0.834 needed for significance at 1 per cent level. *** r = greater than 0.925 needed for significance at 0.1 per cent level. NS correlation not significant. (a)Parents and progenies analyzed were numbers 275,276 and 278 through 283. 85 Table 11.--Parent-progeny and progeny-progeny correlations for 20 trees (Nos. 321 to 340) from an even-aged stand near Ravershagen, East Ger- many. Only characters involved in one or more significant correlations are included. Li -*—‘ PROGENY CHARACTERS o 8 s. a\ C) xo \e u us a: ‘\ tn ‘g \ \ I-l . .4 cu so «n we 0‘ ufig m u .g tn m 'U HEB .c u o o u u u ~\ u u a a. cm a: no cm 0 c: o a\ o 3 3‘33” 3°335~33 co co :2 z: .3 .4 .3 a: :n 3 :5 :h.4 5 6 7 8 9 13 15 19 g_ 22 26 PARENTAL CHARACTERS 40. Tbtal height NS NS * NS NS NS NS NS NS NS NS 44. Clear length NS NS NS NS NS NS NS -** NS ** -* 47. Crown form NS -* NS NS NS NS NS NS NS NS NS PROGENY CHARACTERS 6. Seed length ** -- -- -- -- -- -- -- -- -- -- 7. Height age 1 *** NS -- -- -- -- -- -- -- -- -- 8. Height age 2 ** NS * -- -- -- -- -- -- -- -- 13. Leaf color 8/60 NS NS NS * NS -- -- -- -- -- -- 15. Leaf color 11/60 NS NS NS NS NS ** -- -- -- -- -- 16. Leaf color 12/60 NS NS NS NS NS * *** -- -- -- -- 21. Bud diam. 5/60 NS NS * NS NS NS NS NS -- -- ~- 22. Br. buds 5/60 NS NS * NS NS NS NS -** * -- -- 23. Br. buds 10/60 NS NS NS NS ** NS NS NS NS NS -- 26. Primary lvs. 5/60 NS NS NS NS NS NS NS ** NS -* -- 28. Sec. lvs. 10/59 NS NS NS NS NS NS NS -* NS NS -** 7/5. Height age 1 -*** -* NS NS NS NS NS NS NS NS NS Seed weight * r = greater than 0.44 needed for significance at 5 per cent level. ** r== greater than 0.56 needed for significance at 1 per cent level. *** r== greater than 0.68 needed for significance at 0.1 per cent level. NS Correlation not significant. .l!‘ in! lli.l .‘lld‘ ’ "I’!.---l'. ‘1 until.\ . 86 Table 12.--Parent-progeny and progeny-progeny correlations for 20 trees (Nos. 341 to 360) from an even-aged natural stand near Neustre- litz, East Germany. cant correlations are included. Only characters involved in one or more signifi- 7' TRUCENY'CHARKCTERS‘T <3 >» >. <3 so u u mo ~O\>~. g a: mono \OH '0 mm\\ mHg c: t: \\Ov-i O O HNQO‘H—Omhomm'fl 00m “A: WUQ'U'UHS MO‘mLfi .GUQQJHHHH\U':3:ICL munm\ no on no u) o o o 0 Ch moxJ: :3 \~ \~ c: 'thmmv-d—cv-cv-a Em mmmmma o o o o o o .u a .4 13 u o o 3 .4 u .u o u o o m le m .2 ::m czm :co .2 .c m tsm u o «:6 4’0 «:6 v'uoooomu-au-atu umgm> m>m> 8333888“335‘3w”838°’8 m'mmmgAASmmsqcfiafiH can-«61H 5 6 7 8 9 ll 12 15 19 21 22 23 26 27 28 PARENTAL CHARACTERS 40. Total height NS * NS NS NS NS NS NS NS NS NS NS NS -*** NS 41. Diameter NS * NS NS NS NS NS NS NS NS NS * NS * NS 42. Stem form NS NS NS NS NS NS NS NS NS NS NS NS NS NS -* 43. Crown length NS * NS NS NS NS NS NS NS NS NS NS NS NS NS 44. Clear length NS NS NS NS NS NS * NS NS NS NS * NS NS NS 47. Crown form NS NS NS NS NS NS NS NS NS * NS NS NS NS NS 48. Foliage condition NS NS NS NS NS NS NS -* NS NS NS NS NS NS NS PROGENY CHARACTERS 7. Height age 1 * NS -— -- -- -- -- -- -- -- -- -- -- -- -- 8. Height age 2 NS NS ** -- -- -- -- -- -- -- -- -- -- -- -- 16. Leaf color 12/60 NS NS NS NS NS NS NS***-- -- -- -- -- -- -- 18. Bud color 5/60 NS NS NS NS -* NS NS NS -- -- -- -- -- -- -- l9. Bud set 9/59 NS NS * NS NS NS NS NS -- -- -- -- -- -- ~- 21. Bud diam. 5/60 NS NS NS * NS NS NS NS NS -- -- -- -- -- -- 22. Br. buds 5/60 ** NS NS NS NS NS NS NS NS NS -- -- -- -- -- 26. Prim. lvs. 5/60 -* NS NS NS NS NS NS -* * -* -* NS -- -- ~- 27. Sec. lvs. 9/59 NS NS NS NS NS * NS NS NS NS NS NS NS -- ~- 28. Sec. lvs. 10/59 NS NS NS NS NS NS NS NS NS NS ** NS -* NS -- 7/5 Height age 1 -*** NS NS NS NS NS NS NS NS NS ** NS NS NS NS Seed weight *, **, *** = r greater than .44, .56, and .68 needed for significance at the 5, l, and 0.1 levels respectively. 87 Table 13.--Parent-progeny and progeny-progeny correlations for 20 trees (Nos. 361-380) from an even-aged stand near GUstrow, East Germany. Only characters involved in one or more significant cor- relations are included. T PROGENY CHARACTERS >5 .. o o >. g o‘ m \o \D H n \ \ \ t: \ o H N a o H N 0‘ o-i H H ox H Um u .c .n uc> mus .1: U m a) H H u H \ {3.0 m\ on an 60 on c: o o o o~ \~ C) H c: m m H H H H mm a)... m o o o o o u o o 3 .4 ‘2 '2 c) u u o g 5:» Gun 'o 'o co no 44 Lu .4 u. m"; 33 3 a: :3 '3 a :3 s a 3 as as co ca :2 :2 .4 e: e: e: a: and find 5 6 7 8 11 12 15 l6 19 26 28 PARENTAL CHARACTERS 44. Clear length NS NS NS NS NS NS NS ** NS NS NS 45. Crown diameter NS NS NS NS NS NS NS NS * ** gs PROGENY CHARACTERS 6. Seed length *** -- -- -- -- -- -- -- -- -- -- 7. Height age 1 *** ** -- -- -- -- -- -- -- -- -- 8. Height age 2 k * ** -- -- -- -- -- -_ -- _- 11. Leaf color 9/59 NS -** -** NS -- -- -- -- -- -- -- 12. Leaf color 10/59 -* -* NS NS ** -- -- -- -- -- -- 16. Leaf color 12/60 NS NS NS -** NS NS NS -- -- -- -- 19. Bud set 9/59 -*-***-*** NS *** * NS NS -- -- -- 22. Br. buds 5/60 NS NS NS NS NS NS NS NS -* -- -- 26. Prim. lvs. 5/60 NS NS NS NS ** NS NS NS *** -- -- 27. Sec. lvs. 9/59 NS NS NS * NS NS NS NS NS NS -- 28. Sec. lvs. 10/59 NS NS NS NS NS NS NS NS NS -* -- 29. Leaf length 8/60 NS NS NS NS NS ** NS NS NS NS NS 7/5. Height age 1 -*** -** NS NS NS NS NS NS NS NS * Seed weight * r = greater than 0.44 needed for significance at 5 per cent level. ** r:= greater than 0.56 needed for significance at l per cent level. *** r== greater than 0.68 needed for significance at 0.1 per cent level. NS correlation not significant. 88 Table l4.--Parent-progeny and progeny-progeny correlations for 20 trees (Nos. 381 to 400) from an evenvaged stand near Nedlitz, East Germany. Only characters involved in one or more significant cor- relaticns are included. PROGENY CHARACTERS >. >. u u 0‘ EB 53 -8 -8 In \ \ C G \ H N O O H O\ H H as 0 mm H u vs 0:» om~ u a 2’. s *a s 3 “a ”3 ”a «4 to H H H was 0:4 tun-l u o o o u u o o 3 :4 o u u a) a m cm» H 3 .c: m 00 mm .1: 'U on ma 44 ea m > m>' 'U 0 vi a m m 1: a m ‘03 ‘3 3 :73 1% S 3 .3 :3 $1.94 (EH :30: 5 7 ll 15 16 19 27 28 7/5 PARENTAL CHARACTERS 41. Diameter NS NS NS NS NS NS NS * NS 43. Crown length NS NS -* NS NS NS * NS NS 45. Crown diameter NS * NS NS NS NS NS NS NS 46. Crown flattening NS NS NS NS NS NS NS NS -* PROGENY CHARACTERS 6. Seed length * -- -- -- -- -- -- -- -- 8. Height age 2 NS NS *** -- -- -- -- -- -- 16. Leaf color 12/60 NS NS NS ** -- -- -- -- -- 19. Bud set 9/59 NS NS NS * NS -- -- -- -- 22. Br. buds 5/60 NS * NS NS -* NS -- -- -- 29. Leaf length 8/60 NS * NS NS NS NS NS NS -- 7/5. Height age 1 Seed weight -*** ** NS NS NS NS NS NS -- * r = greater than 0.44 needed for significance at 5 per cent level. ** r = greater than 0.56 needed for significance at l per cent level. *** r = greater than 0.68 needed for significance at 0.1 per cent level. NS Correlations not significant. 89 Table lS.--Parent-progeny and progeny-progeny correlations for 20 trees (Nos. 501 to 520) from an even-aged stand near Joachimsthal, East Germany. Only characters involved in one or more significant correlations are included. “W PROGENY CHARACTERS >. >. 8“ ts in \ \ \ Inc: C \ O H N O O F‘O‘ H H F4 (7‘ mo UO‘H u .I: In "Ucuosmtn U .5 U 0.) H H H H \ :lmtnmm: co m 000 O O O O‘ .0 \ O 00 'H G (69-! 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ADJ .m a.“ a.“ n. m. m m. a u u 8 S S 1 «Ha my. a.” a nn 3 a. .w a u. a S O .4 u a m-HMQEDZ USN .Hmn—gz mumuomuwno amucmumm umuumumno .moxuns new .2 .z .M .H .m macawmu now mamas w:Mum mafia: muouomumsu Hmuamumm ma uom acowumamuuoo vamum-cmmsuon”unmowMficwwm-.HN magma CHAPTER VII CORRELATIONS INVOLVING PARENTAL AND JUVENILE CHARACTERISTICS OF ENTIRE STANDS Materials and Methods The 19 parental characters measured on the cone, seed, and leaf specimens of 33 of the 39 parental stands of Scotch pine sampled were analyzed for simple correlations with 14 of the juvenile characters measured by Wright and Bull (1963) on the 122—origin replicated Scotch pine provenance test conducted at East Lansing, Michigan. Six of the parental stands sampled were not represented in the provenance test. All data were entered on punch cards and processed through the Control Data Corporation 3600 computer at Michigan State University on a correlation-regression program. The parental and juvenile data were grouped by areas of clima- tic similarity prior to analysis as follows: Group 1 Regions A, C, G, and H (19 stands). Group 2 Regions I and L (6 stands). Group 3 Regions J, K, M, and N (8 stands). Grouping in this manner reduced the degrees of freedom considerably, as compared with an analysis based on stands from the entire range. However, it had the advantage of eliminating the many meaningless correlations which almost always result when very diverse types of data are considered together. 116 117 The resulting correlations were assembled in Table 22, organised for ready comparison between the three area groupings. The most important fact presented was the failure of the corre- lations obtained to present a consistent pattern. The actual and expected numbers of significant correlations were as follows: Significance Level E 0.1 l 5 fi Per cent Per cent Per cent 3 t Value of I needed, ‘ group 1, l7 d.f. .693 .575 .456 ’ group 2, 4 d.f. .974 .917 .811 l; group 3, 6 d.f. .925 .834 .707 5' Number of r's exceeding amount needed Actual 25 102 190 Expected (912 .9 9.1 45.5 combinations) It can be seen that 24 per cent of the correlations significant at the five per cent level of significance can be expected to be meaning- less. Discussion Presumably the most meaningful correlations are those which were significant in all three areas (there are two examples of this, one involving a change in sign) or in two of the three areas. Thus there is strong evidence that stands having large seeds (characters P28, P29) produce progeny which are taller, greener in the winter, have longer needles, and produce buds later in the autumn. Also 118 that stands with long needles (character P35) produce progeny which are taller at age 3 and which have longer needles. Or that stands in which the ratio cone length/apophysis length is high produce progeny which are greener in the autumn, have later bud formation, or produce longer needles. Note that it is the same progeny traits in each case, and that they are associated with rather diverse parental characters. This fact hints that there is not necessarily a direct cause-and-effect relationship between a given parental trait and its associated progeny trait. Rather, it is more likely that the evolutionary factors which resulted in smaller seeds or more slender cones in the far northern stands also resulted in earlier bud set, deeper yellow foliage, and shorter needles in the seedlings. There are some instances in which the direction of a cor- relation is reversed between areas 1 (Scandinavia through Germany), 2 (Great Britain), and 3 (southern Europe). This certainly means that one cannot forecast for the entire species from a part, but it does not necessarily mean that the relationships are not valid for the respective areas. As already emphasized by Wright and Bull (1963), many progeny traits reach their highest development in the central portion of the species' range. For example, height growth and needle length of the progeny decrease as one proceeds outward from a center in northern France-Belgium-western Germany in almost any direction. That is not true for the parental traits, which vary in a more irregular manner. Hence the change in direc- tion of relationship as one proceeds from northern to southern Europe. 119 Table 22.--Significant correlations between parental and juvenile characteristics within group 1, consisting of regions A, C, G, and H, group 2, consisting of regions I and L, and within group 3, consisting of regions J) K, M, and N, using stand means. 1220 JUVENILE cgéggprsn n H H H k H H H U; .. .. 2. .9. 2 a 3 “a . ° 3 3 .. no .cao O O O O O 3 G U U Q Q U U U U U U H .99 0 >6 M H H D O O H H0 C H .: 9.: S 2. 8 S 8 : s “as 3.. a: .9. 3 a .8 a. as. s as as :3 ° .3 s; 8:: .. :1 z: 3:: '32 '33 :3 s: "a: “s 7’ “a as 83 s —' CHARACTER REGIONS J34 J7[5 J9 J12 J13 J16 J31 J20 J18 J19 J17 J27 J29 334 P6 Cone length A,c,c,N Ns Ns Ns NS Ns .53 Ns Ns Ns .47 NS NS .47 NS I & L NS NS NS 84 NS NS NS NS NS .96 NS NS .96 NS J,K,H,N NS NS NS NS .72 NS NS NS NS NS .81 .79 NS NS P7 Closed-cone A,C,G,H NS NS NS NS NS .54 NS NS NS NS NS NS NS NS width I,L .87 NS NS .87 NS NS NS NS NS .94 NS NS .94 NS ’ J,N,N,N Ns NS Ns NS Ns NS Ns Ns NS Ns NS .71 -.73 Ns P8 Open-cone A,C,G,H NS NS NS NS NS NS NS NS NS NS NS NS NS NS width I,L NS NS NS NS NS NS NS NS NS .84 NS NS .84 NS J,K,M,N Ns NS Ns NS NS NS Ns NS Ns Ns .81 Ns Ns NS 3 P12 Length larg- A, ,G,H NS NS NS NS NS NS NS NS NS NS NS NS NS NS est apophysis I, J- C L NS NS NS NS NS NS NS NS NS NS NS NS NS NS K,M,N NS NS NS NS NS NS NS NS NS NS NS NS NS .94 914 Thickness A,C,G,H NS NS NS NS NS Ns Ns Ns NS NS NS Ns NS Ns 1ergest I,L NS NS NS NS NS NS NS NS Ns .91 NS Ns NS NS apophysis J,K,M,N -.77 Ns .91 NS .83 .87 NS Ns NS NS Ns .73 -.84 Ns P17 Thickness of A,C,G,H NS NS NS NS NS NS NS NS NS NS NS NS NS NS opposite I,L NS NS NS NS NS NS NS NS NS NS NS NS NS NS J,K apophysis ,M,N NS NS .90 NS .89 .90 .72 NS NS NS .74 NS NS NS P19 Basel angle A,C,G,N NS NS NS NS NS NS NS NS NS NS NS NS NS NS of cone I,L -.81 NS NS NS NS NS NS NS NS NS NS NS NS NS J,K,M,N .71 NS NS NS NS NS NS NS NS NS NS NS .77 NS P28 Seed length 0,8,3 .70 NS -.69 .72 .71 .56 .61 .69 .62 .75 .68 .73 .74 Ns L .91 Ns NS .88 NS Ns NS NS Ns .86 NS NS .92 Ns K,M,N NS -.72 .75 NS NS .76 Ns NS Ns .79 .77 Ns Ns .77 P29 Seed width .c,N .86 .50 -.74 .85 .89 .63 .75 .63 .61 .84 .86 .90 .84 -.58 .92 NS Ns .92 Ns NS NS NS Ns .94 Ns Ns .91 NS C L K,M,N NS NS NS NS NS NS NS NS NS NS .77 NS NS NS C G N L K 059 NS NS NS .51 .63 060 I51 NS .62 .48 .52 .69 '.50 .81 NS NS .86 NS NS -.82 NS NS .99 NS NS .94 NS A I J P35 Leaf length A I J ,N,N NS NS NS NS NS NS NS -.86 NS NS NS NS NS NS P38 Leaf twist A,C,G,H NS NS NS NS NS Ns NS NS Ns NS Ns NS NS Ns I,L NS Ns NS NS NS NS NS Ns NS NS NS Ns NS NS J,K,M,N Ns NS Ns Ns NS Ns NS Ns Ns Ns NS Ns NS NS P9 Cone length A,C,G,H Ns Ns NS Ns NS Ns Ns Ns NS NS Ns NS Ns NS Closed-cone I,L NS NS .86 NS NS NS NS NS NS NS Ns .83 NS Ns width J,K,M,N NS NS NS NS NS NS NS NS NS NS NS Ns NS NS P10 Cone en th A,C,G,N .64 NS NS .74 .60 .62 .54 .59 NS .64 .63 .63 .54 us Open-cone I,L NS Ns NS Ns Ns NS Ns Ns NS NS Ns Ns Ns NS width J,K,M,N NS Ns NS Ns Ns Ns Ns Ns Ns NS NS NS Ns -.80 P15 Apophxniu 1n. A,C,G,H -.57 NS Ns -.60 -.58 -.55 -.53 NS Ns -.56 -.62 -.57 -.51 NS Apo. width I,L Ns Ns NS Ns NS NS Ns Ns Ns Ns NS NS NS NS 3.x ,N,N Ns NS NS NS Ns Ns NS Ns Ns Ns Ns —.71 Ns NS P16 Apophxaia 1n. A,C G,N Ns Ns NS Ns NS NS Ns Ns Ns Ns NS Ns NS NS Apo. thickness I,L Ns Ns NS NS NS Ns Ns Ns NS Ns NS NS NS Ns J,K, ,N .94 NS -.86 NS -.89 NS Ns NS Ns NS -.74 -.76 .75 NS M ,6,“ NS NS NS NS NS NS NS NS NS NS NS NS NS NS M P18 Ape. thickness A,C Apo. opposite 1,1 Ns NS NS Ns Ns Ns Ns Ns Ns NS Ns Ns Ns Ns thicknesu J,K, ,N NS Ns NS NS NS NS Ns -.71 -.85 Ns NS NS NS NS P20 Cone length A,C,G,H .68 NS -.46 .76 .61 .74 .65 .66 NS .76 .62 .65 .69 NS Apophysis I,L .91 NS NS NS NS NS NS NS NS .89 NS NS .93 NS 1ength J,K,H,N -.84 NS NS NS .83 NS NS NS NS NS .73 .89 NS NS P30 Seed length A,C,G,H NS NS NS NS ‘NS NS NS NS NS NS NS NS NS NS Seed width I,L NS NS NS NS NS NS NS NS NS NS NS NS NS NS J,K,M,N NS NS NS NS NS NS NS .74 NS NS NS NS NS .71 .—I N pd Generally speaking, the parental characters of seed size, leaf length, and the ratio cone length/apophysis length were of the most value in forecasting progeny performance, but even with these care must be taken to apply the relationships only within an area within which they are presumed to be applicable. The other measured parental characteristics are of value in delimiting populations which are gene- tically different but not in telling how the seedlings grow. CHAPTER VIII CORRELATIONS INVOLVING CLIMATE Materials and Methods Meteorological data was obtained for the 121 origins represented in Wright and Bull's (1963) provenance test and for the parental specimen stands where they differed from or added to the original provenance col« lection. The data was assembled from information received from the various cooperators who originally supplied seed or parental specimens, the weather services of the country of origin, the Great Britain Meteoro» logical Office (1958), and from the Foreign Area Section of the United States Weather Bureau, United States Department of Commerce. The remoteness of the stands and the wide area of distribution of Scotch pine made it impractical to assemble daily weather data. All information was assembled on a monthly and annual basis. The basis of the precipitation and temperature data was usually a 20-year record, although in some instances this period was as long as 50 years. While it would have been desirable to use comparable years of record, wars, catastrophes, and lack of long established weather sta- tions in some areas resulted in the use of the best data available. The precipitation data was recorded in millimeters as average monthly precipitation (weather characters 20—31), average monthly 122 123 precipitation on a yearly basis (weather character 32), and total annual precipitation (weather character 33). Temperature data was recorded in degrees centigrade as the number of plus six degree months in a year (character 34}, average monthly temperature (weather characters 35-46) and average monthly temperature on an annual basis (weather character 47). In addition to the climatic characterisitics listed above, 18 combinations of monthly precipitation and temperature figures were used to further measure plant-climate relationships as follows: Identifying Number Description of Meteorological Data 48 Average temperature for period April-June 49 Average temperature for period May-July 50 Average temperature for period july-Augnst 51 Average temperature for period July~September 52 Total precipitation, SeptemberwJune 53 September-June precipitation as per cent of total annual 54 Total precipitation, OctoberoApril 55 October-April precipitation as per cent of total annual 56 Total precipitation, April-June 57 April-June precipitation as per cent of total annual 58 Total precipitatzon, May June 59 May-June precipitation as per cent total annual 60 Total precipitation, July-August 124 61 July-August precipitation as per cent of total annual 62 Total precipitation, May—August 63 May-August precipitation as per cent of total annual 64 Ratio of average April-June temperature to pre- cipitation for April-June 65 Ratio of average July-August temperature to pre- cipitation for July-August The sampling of Scotch pine provenances cut across so many areas of diverse weather patterns that correlating the data as a whole resulted only in an abundance of meaningless but statistically significant corre- lations interspersed among those correlations of real biological signi- ficance. As an example, when the 16 juvenile characters used in this study (Table 23) were used to get simple correlations with 14 tempera- ture characteristics there were 172 correlations out of a possible 224 combinations significant at the one-tenth of one per cent level of sig- nificance. Many of these correlations were artificial and biologically meaningless as determined by scatter diagrams and the only real test was by the power of the correlation (r) and by stratifying the obser- vations. Another basis for stratification other than diverse weather pat- terns was through the knowledge that many of the parental and juvenile characteristics reached an optimum at midmrange, or at about 50 degrees north latitude. While not universally true, particularly considering some of the east-west gradients, it was considered correct enough to use in stratifying the data. 125 Table 23.--Description of juvenile characteristics used in corre- lations with meteorological and geographical data of origin and the grades used in scoring the Scotch pine progeny. Character Unit of Lowest value Highest value and Number Date Measure Range assigned to assigned to J5 Seed weight 11/56 mg. 2.9- Light seed Heavy seed 12.2 ‘ J6 Seed length 11/58 mm. 3.4- Short seed Long seed 5.9 J7/5 Ratio l-zr.ht. 5/60 Number 8'25 Low ratio High ratio seed wt. J9 Foliage color 6/59 Grade 4-8 Yellow-green Green J12 Foliage color 10/59 Grade 8—20 bmroon Green J13 Foliage color 8/60 Grade 8-16 Green Dark green J16 Foliage color 12/60 Grade 4-32 Yellow-green Dark green J17 Initiation of 1960 Day of 123- Early ini- Late initiation growth year 116 tiation J18 Bud color 5/60 Grade 4-24 Bright green Brown J19 Time of bud 9/59 Day of 200- Early set Late set set year 269 . J20 Earliness of Number 1-5 Early color Late color leaf color J24 Pullability 7/59 Number 8-18 Easy to lift Hard to lift from soil from soil J27 Presence of 9/59 Grade 0—4 None present Present on 50% secondary lvs. of seedlings J29 Leaf length 8/60 cm. 25—43 Short needle Long needle J31 Foliage color 12/61 Grade 4-40 Yellow Blue-green J32 Height, 3-yr. 1961 mm. 118- Short seed- Tall seedling 708 ling 126 The juvenile data was studied in relation to meteorological con- ditions by separating the regions into A through F, G, H, and J thrzagh N (less Region L, Scotland). The parental data, which contained fewer observations, was studied by grouping regions A through G, and H through N (including Turkey, which in the parental study was separated from Wright's re- gion K). All data were entered on punch cards and processed through Michigan State University's Control Data Corporation 36C0 data pro« cessing system on a correlation-regression program. Literature Review Lehotsky (1961) reported that altitude had an effect on seed and cone quality in Scotch pine. He collected seed from 4w5 trees at 600, 700, 870, and 1100 meters altitude. These seed were sown in nurseries at 640, 750, 870 and 1050 meters altitude. Measure- ment of seedling height at two years showed that at each nursery, stem height decreased with increasing altitude of provenance. Callaham (1959) reported a highly significant influence of elevation on the height and diameter growth of 20-year-old ponderosa pine and Jeffrey's pine (Pinus jeffreyi Grev. and Balf.) in a pro- venance test of seven altitudinal collection of Jeffrey pine (3,500-8,500 feet). McLemore g£_gl. (1961) found that the dry-matter content of the previous year's needles collected from two plantations of lob- lolly pine in Mississippi and Louisiana showed no correlation with 127 mean annual rainfall, January or July temperature, length of growing season, or the ratio of July temperature to mean annual rainfall. The provenances tested represented 25 counties from the geographical and climatic extremes of the range of loblolly pine. The authors state the different results of Langlet (1959) could be attributed to the greater importance of low temperature for Scotch pine. Langlet (1961), in reporting on Central European spruce pro- venances tested in Sweden, stated that growth was affected by lati- tude as much as altitude, i.e., that mean temperature and length of growing season were the important factors in spruce. However, as compared with Scotch pine, both racial character and climatic re- sponse depended far more on local climate than on latitude as such, and no effect of day length could be observed. Critchfield (1957) found that each of the principal morphological characteristics of lodgepole pine (leaves, cones, anatomy of leaves) was unique in its variation pattern. Leaf width was tied more closely to elevation than geographic location. He found that latitude, which has an important influence on local climate in California, showed no clear-cut association with any single morphological characteristic throughout the whole of the species range (California to Alaska and eastward to Colorado). Goddard and Strickland (1962) reported that longitude, latitude, and early and late rainfall, as independent variables, accounted for 88 per cent of regional variation observed in highly significantly different groups of wood specific gravity in slash pine (Pinus elliottii Engelm.). 128 Squillace and Silen (1962) reported on a 30-year-old (10- origin, 5 plantation) and a 45-year-old (20 origins in one planta- tion) provenance test of ponderosa pine. They found that growth rate was related to average annual temperature and the April-May temperature of seed source localities. Growth was not correlated with the number of frost-free days, however. The two strongest climatic variables found in simple correlations were September- throush-June precipitation as a per cent of annual and April-May temperature. Growth of the trees of different sources was un- related to springtime precipitation (April through June). Kraus (1962) reported that red pine shoot growth was found significantly correlated to the moisture deficiencies of the pre- vious year's food storage period (July lS-August 30) and the cur- rent yeor's shoot elongation period (May lS-July 15) for three sites in central lower Michigan. The study included 18 red pine trees in three stands and included meterological data of 10-20 years duration. Langlet (1959), in reviewing his studies of Scotch pine variability, stated that the dry matter content of needles shows a remarkably close relationship with the conditions in the native habitats of different provenances in respect tc latitude and dura- tion of the period of vegetation (expressed as the number of days with an average temperature of +6OC. or more). He found an even closer relationship between the dry matter content and the length of the first day of the year with a normal average temperature of +6°c. 129 Meyer and Barman (1963) found both a geographic and altitudinal cline in the ratio of cone/bract length in a phenotypic variation study of balsam fir (Abies balsamea (L.) Mill.). They also reported that leaf length was correlated with altitude. Schoenike (1963) in reporting natural variation in jack pine (ginus banksiana Lamb.) stated that, of six environmental factors, precipitation and latitude had the most effect on morphology and tree growth. Weather Patterns Within Scotch Pine Distribution Area In regions A through F, Scandinavia, Baltic area, Poland, and Northwest U.S.S.R., the per cent of the total annual rainfall which Occurs during the period April to August decreases with increasing latitude, increases with increasing east longitude, and increases with increases in elevation (Table 24). Temperatures during the period Maythrough September decrease with increasing latitude and temperatures during the period September through April decrease with increasing east longitude and with elevation increase (Table 25). The higher latitudes have an increased amount of precipitation during the April through June period in relation to temperature while increasing east longitude shows a reduction in precipitation during the same period in relation to temperature. In region G, Northeast Germany and Czechoslovakia, the rain- fall during the period April through August shows an even sharper decrease withincreasing latitude, particularly during the month of June, and there is an increasing amount of the total annual precipi- tation during the period May to August with increasing east longitude 130 Table 24.--Corre1ation of latitude, longitude, and elevation with precipitation data in regions A-F, G, H, and J, K, M, and N. NS ri’ Regions " A—F g. H J.K.M.& N Meteoro- . . . . . 163ml .3 f: 5 .5 :2" 5 .3 2° 5 .3 ‘2? E5 Data :3 .9. a :3 3 £3 a 3 a :3 .3 2:5 fivr 1 2 3 l 2 3 1 2 3 1 2 3 Actual precipitation in January NS -.57 NS NS NS NS NS NS NS NS NS NS February NS .,52 .35 NS NS NS NS NS NS NS NS NS March NS ~.59 .35 NS NS NS NS NS NS NS NS NS April .32 -.71 -.35 -.63 NS .51 -.53 NS NS NS NS NS May NS -.63 -.43 -.66 NS .56 -.66 NS .66 NS NS ;3 June NS NS NS -.84 NS .62 -.65 NS .73 NS NS NS July NS -.48 NS -.62 NS .49 NS NS NS NS NS NS August NS -.49 NS -.58 NS .45 NS NS NS .50 NS NS September NS 9.45 NS NS NS NS -.57 NS .57 NS NS NS OCtober .34 -.56 NS NS NS NS NS NS NS NS NS NS November NS -.58 v.34 NS NS NS NS NS NS NS NS NS December NS -.57 -.33 NS NS NS .48 .49 -.37 NS NS NS Monthly av. NS -.58 NS —.42 NS NS NS NS NS NS NS NS Sept-June NS -.58 -.33 NS NS NS NS NS NS NS NS NS Oct.-April NS -.59 ~.34 NS NS NS NS NS NS NS NS NS April—June NS -.62 -.34 -.78 NS .60 -.64 NS .69 NS NS NS May-June" NS -.51 NS -.79 NS .62 -.68 NS .70 NS NS NS July-Aug. NS -.51 NS -.62 NS .49 NS NS NS .42 NS NS May-Aug. NS -.52 NS «.76 NS .59 -.50 NS .58 NS NS NS Precipitation as per cent of annual total ’ Sept.-June .49 -.78 «.60 NS NS NS NS NS NS ~.80 NS .48 0ct.-April .53 -.86 -.49 .45 -.38 NS NS NS NS -.60 NS .50 April-June -.51 .58 NS -.59 NS .46 -.88 .65 .84 NS .51 .50 May-June -.54 .73 NS -.54 NS .43 -.82 .66 .70 NS .48 NS July-Aug. -.49 .76 .60 NS NS NS NS NS NS .80 NS ~.48 May-Aug. -.54 .80 NS -.49 .43 .37 -.49 .68 NS .55 NS -.46 ‘.05 .316 .368 .456 .368 ’.01 .407 .472 .575 .472 r.001 .505 .580 .693 .579 NS 8 Non-significant correlations 131 Table 25.--Corre1ation of latitude, longitude, and elevation with temperature data for regions A-F, G, H, and J, K, M, and N. f v ‘—V W *** Regions A-F G H J,K,M & N Meteoro- . logical u 2:0 S u 2° 5' 4.» 2:0 5 u 3:0 Data 3 .3 E3 .3 3 B .3 .3 E 3 3 w_ l 2 3 1 2 3 l 2 3 1 2 Average temperature in January NS -.84 -.60 NS -.72 -.55 .76 -.82 -.69 -.38 NS February NS -.78 NS NS -.74 -.53 .63 -.76 -.71 NS NS March NS -.86 -.60 NS -.45 NS NS NS -.54 -.41 NS April NS -.69 -.58 NS NS NS NS NS NS -.45 .51 May -.46 NS -.49 NS NS NS NS NS -.47 NS .63 June -.64 NS NS NS NS -.39 -.47 .54 NS -.42 .59 July -.49 NS NS NS NS NS -.56 .68 NS -.59 .60 August -.50 NS -.33 NS NS -.39 -.59 .67 NS -.67 .61 September -.35 -.49 -.55 NS NS NS -.46 NS NS -.56 .48 October NS ~.80 -.62 NS NS -.55 NS NS NS -.56 .48 November NS -.85 -.67 NS ~.50 -.70 NS NS NS -.57 .39 December NS -.86 —.60 NS -.48 -.41 .66 -.68 -.49 -.61 NS Monthly av. NS -.62 -.60 NS NS -.45 NS NS NS -.53 NS April-June -.41 -.48 -.50 NS NS NS NS NS NS -.41 .59 May-July -.57 NS NS NS NS NS -.38 .59 NS -.46 .62 July-Aug. -.51 NS NS NS NS -.37 -.59 .68 NS -.63 .61 July-Sept. -.46 NS -.38 NS NS NS -.64 .63 NS -.61 .57 6 months -.51 -.33 -.43 .42 NS NS NS NS NS NS NS Ratios Apr.-June prec. Apr.:3une temp. .64 -.38 NS -.56 NS .64 —.51 NS .67 NS NS July-Aug. prec. July-Aug. temp. NS -.52 NS -.48 NS .55 NS NS NS .47 NS V—V '1 There were no correlations in regions J,K,M, and N with elevation. NS a Non-significant correlation 132 and with elevation. Winter temperatures decrease with increasing east longitude and with elevation. Temperature during the period April through August increases more than precipitation as latitude increases, but decreases in relation to precipitation as elevation increases. In region H, western Germany, eastern France, and Belgium, precipitation decreases with increased latitude during the period April through August, and increases during the same period with increased east longitude and with elevation. Temperatures increase with latitude during the period December through February and de- crease during the period June to September, however, winter tempera- tures decrease and summer temperatures increase as east longitude increases and winter temperatures decrease as elevation increases. The amount of precipitation during the period April to June in- creases with relation to temperature as latitude increases but is reversed with elevation increases. In regions J, K, M, and N, Yugoslavia, Greece, south central France, and Spain, the per cent of total annual precipitation de- creases during the period September through April with latitude increases but increases with elevation. The per cent of total an- nual rainfall occurring during the period April to June increases with increasing east longitude. In summary, regions A-F have less rainfall in the east but a higher percentage of the annual amount occurs during the summer with lower summer temperatures in the north and lower winter temperatures to the east and at the higher elevations. Region G has decreased 133 amounts of summer rainfall in the north and increasing rainfall at higher elevations with lower winter temperatures as elevation in- creases and to the east. Region H has the same precipitation pat- tern, growing season temperatures are less to the north and higher to the east. Regions J, K, M, and N have a greater percentage of the total annual rainfall occurring during the summer to the north and east at the lower elevations. E These changing patterns of summer and winter precipitation E and temperature with latitude, longitude, and elevation changes make it impractical to study parental and juvenile correlations with h meteorological data for the distribution area as a whole. Correlations Involving Parental Characteristics and Climate Nineteen cone, seed, and leaf characteristics of parental speci- mens were analyzed for possible simple correlations with meteorological data. Separate analyses were made for northern Europe (regions A, C, G, northern and central Scandinavia, northeast Germany, Czechoslovakia, Table 26), and central and southern Europe (regions H, J, K, M, N, western Germany, Belgium, France, Spain, Turkey, Table 27). The northern area, comprising regions A through G had the fol- lowing actual and expected numbers of correlations significant at the various levels: 134 Significance Level 0.1 l 5 +7 __ Per cent Per cent Per cent _ value of r needed (17 d.f.) .693 .575 .456 Number of r's exceeding amount needed Actual 23 66 156 Expected (874 combinations) .87 8.7 43.5 Apophysis thickness increased as the percentage of total annual precipitation occurring during the period July-August increased. Apophysis thickness decreased with lower temperatures and the thick- ness in relation to length decreased with higher temperatures during the period September-December. Thinner apophyses are associated then with cold, dry winters and cold summers. Seed length and width increased as a higher percentage of the annual precipitation occurred during the April-June period and as temperature increased throughout the year. An increased amount of leaf twist is associated with increased temperatures during May and June. The various measurements of cone length and width, of apophysis width, and of leaf twist are omitted from Table 26. They were found not to be correlated with any climatic characteristics. On the other hand, three of the seven ratios involving some of those same traits were correlated, especially with temperature data. 135 Table 26.-~Correlations between parental characteristics and meteoro— logical data for north European Scotch pine (regions A, C, and G). fl W ._.7 Parental Characteristics .3 Ratios (D >. .e .o .c.c Meteoro" m m m '5. m ‘Jq'fi go a: 450:0 logical 7.: 2:: 2.8 53 88 88 n... as as <5 :5 .c a u N as M 8'2? 8'8. 5.533%" 8% tateaa'ae 85 as: as sass a"; 3.233 as as 852‘ +_ _"+ P12 P14 P17 P28 P22_,P35 P38 P15 P16 P20 Actual precipitation in January NS NS NS NS NS NS NS NS .62 NS February NS -.58 NS NS NS NS NS NS .62 NS April NS NS NS NS .56 NS NS NS NS *3 May NS NS NS .61 .68 NS NS NS NS NS June NS NS NS .50 .61 NS NS NS NS NS August .52 .63 .53 NS NS NS NS NS NS NS October .63 .63 NS NS -.62 NS NS .60 NS .47 April-June NS NS NS .55 .67 NS NS NS NS NS May-June NS NS NS .59 .68 NS NS NS NS NS July-Aug. NS -.48 .46 NS NS NS NS NS NS NS Entire year NS NS NS NS NS NS NS NS NS NS Precipitation as per cent of annual total Sept.-June NS -.61 -.60 NS ’NS NS NS NS .69 NS ApriléJune -.50 NS NS .52 .68 NS NS -;60 NS NS May-June' -.50 'NS NS .51 .62 NS NS -.59 NS .55 July-Aug. NS .61 .61 NS NS NS NS NS -.69 NS Average Temperature in January NS -.53 NS .68 .82 .56 NS NS NS NS February NS -.48 NS .67 .81 .65 NS -.57 NS NS March NS -.54 NS .86 .90 .48 NS -.49 NS .53 April -.48 -.54 NS .87 .89 .51 NS -.54 NS .61 May -.51 -.56 NS. .88 .85 .50 .49 -.48 NS .64 June -.51 -.52 NS .79 .70 .52 .4) —.46 NS NS July NS NS -.47 NS NS NS NS NS NS NS August NS -.61 -.53 .50 .55 NS NS NS .54 NS September NS -.63 -.46 .66 .77 .55 NS NS .54 NS October NS -.58 NS .70 .81 .52 NS NS .51 NS November NS -.60 NS .58 .74 .55 NS NS .55 NS December NS -.63 .50 .58 .73 .50 NS NS .60 NS AprilsJune -,51 -.56 'NS .87 .85 .52 NS -.51 NS .62 MayéJuly NS -.55 .47 .78 .70 .51 NS NS NS .53 July-August NS —.60 -.51 NS .59 'NS NS NS NS 'NS Entire year NS -.59 NS .77 .85 .55 NS NS .49 .49 No. 6° months NS -.66 NS .77 .88 NS NS ‘ NS .56 .55 r I :456,w357§: .693 for significance at the 5, l, and 0.1 per cent “— levels respectively. NS 3 Non-significant correlation 136 Table 27.-~Correlations involving relationships between parental cone, seed, and leaf characteristics and meteorological and geographical data of origin for regions H, I, J, K, L, M, and N. *v—v—r W v—v—v fi' Parental Characteristics Ratios Meteoro- -3 '3 3 U) (D logical 8 a .2 as as :5; Data 3 2 ass-ass .a.sss u o m m o.o e o 0.: m a» m-H seseesswsc’ s 33......” at» m u e44 o.o - o m c 'u u>'o u -- -. m --o'o 88 .23 as 2.: as H38 83 °S§88$ on: oeoamugumomam3§1~ and ch 3 m u .3 c I: .3 .3 Geograph- - m 3 .3 4.3 43 .3 3 ical £2 gov-l 5H 8 cu sag-818 '5 '3 5... 8.32.2? 21:2 21.2 . BE :8 8:: as s 353% .23 £33 5 84?. sass 3.9. a? w_" fi J5 J34 J31 J20 J18 J19 J17 J27 J29 24 Actual precipitation in January NS NS .34 NS NS NS NS NS -.47 NS February NS NS .33 NS NS NS NS NS -.44 NS March NS NS .41 NS NS NS NS NS NS NS April NS NS .48 NS NS NS NS NS -.52 NS May NS NS .35 NS .53 NS NS .45 NS NS June NS NS .34 NS .34 NS NS NS NS NS August NS NS NS NS .38 NS NS NS -.35 NS September NS NS NS NS .34 NS NS NS -.38 NS October NS NS NS NS NS NS NS NS -.51 NS November NS NS NS NS NS NS NS NS -.49 NS December NS NS .36 NS NS NS NS NS -.46 NS Monthly av. NS NS .34 NS .34 NS NS NS -.47 NS Sept.-June NS NS .34 NS .32 NS NS NS -.48 NS OcL.-April NS NS .36 NS NS NS NS NS -.48 NS April~June NS NS .37 NS .40 NS NS .36 -.37 NS May-June NS NS NS NS .46 NS NS .38 NS NS July-Aug. NS NS NS NS .35 NS NS NS -.39 NS May-Aug. NS NS NS NS .40 NS NS NS -.34 NS Per cent of total annual precipitation Sept.-June NS NS .38 .41 NS NS NS NS -.49 NS 0ct.-April NS NS .49 .37 NS NS NS NS -.61 NS April-June NS .41 NS NS NS NS .51 NS .63 NS May-June .36 .39 -.43 NS NS .41 NS NS .68 NS July-Aug. NS NS -.38 -.42 NS NS NS NS .49 NS May Aug. NS NS -.42 -.36 NS NS NS NS .59 NS r a .316 significant r u .407 significant r a .505 significant NS n Non-significant at the 5 per cent level of significance. at the l per cent level of significance. at the 0.1 per cent level of significance. correlation. Table 28.--Continued 143 J Juvenile Characteristics Meteoro- g g u, .U ,H logical f‘ '3 0 'g g g f," and m U U) H .Dfl H ‘H m o O ‘H m H Geograptv- .. J 8’. 2 '6‘ “3'3 0.: at .c E lcal asses :5 ° .§:588.ss Data cu-H 0H .—a\ H—l 'U 44 :60 on). «3:: F4 3’58 :3 :22 638 Ea’ 84?. 618383. 3.9. E. g J5 J34 J31 J20 J18 J19 J17 J27 J29 J24 Average temperature in January NS NS .51 .58 .32 .46 NS .46 -.40 -.53 February NS NS .49 .36 NS NS NS NS -.43 -.33 March NS NS .59 .59 .37 .55 NS .58 NS -.53 April NS .47 .52 .56 .44 .66 .36 .72 NS -.54 May NS .60 .43 .51 .55 .66 .48 .88 NS -.45 June .53 .54 NS NS .39 .43 NS .60 NS NS July .43 .36 NS NS NS NS NS .48 NS NS August .44 .51 .34 .42 .36 .50 .37 .67 NS NS September NS .55 .54 .61 .43 .68 .42 .79 NS -.45 October NS NS .61 .61 .36 .61 NS .65 NS -.53 November NS NS .54 .59 NS .50 NS .55 -.37 -.51 December NS NS .50 .56 NS .45 NS .48 -.38 -.51 Monthly av. NS .41 .44 .53 .44 .62 NS .63 NS -.54 April-June NS .57 .46 .52 .51 .67 .43 .79 NS -.47 May-June .48 .56 NS NS .46 .53 .41 .72 NS NS July-Aug. .44 .45 NS NS NS .39 NS .59 NS NS July-Sept. NS .52 NS .47 .39 .54 .38 .71 NS NS 6 months .41 .63 .51 .35 .47 .69 .52 .77 NS NS Ratios Apr.~June prec. Apr.-June temp. -.39 -.49 NS NS NS -.39 -.33 -.32 .58 NS July-Augg,prec. July-Aug. temp. NS NS NS NS NS NS NS NS -.47 NS Latitude -.58 -.74 NS NS -.39 -.63 -.57 -.54 -.72 NS Longitude .38 NS -.56 -.43 NS NS NS NS .59 .40 Elevation NS -.43 NS -.49 -.32 -.43 -.43 -.58 NS .45 144 Foliage color (character J31) was recorded with the low number representing yellow and the high numbers representing blue-green. The yellow predominated where the temperatures during the period August through May were higher and the percentage of total annual rainfall occurring during the winter period, October through April, was highest. Extreme yellowing of foliage then occurred in areas with milder winters and dryer summers. Earliness of yellow coloring followed the same pattern. Bud color, where green was scored low and tan high, showed that tan was associated with higher temperatures throughout the year but particularly in April to May and with increased amounts of rainfall during the period April to June. The formation of buds was scored as the date of the year that they formed. Late formation was correlated with warmer temperatures throughout the year, particularly during the April-June period, more 6° C. months, and a greater amount of the annual precipitation oc- curring during the May-June period. The early presence of secondary leaves was strongly correlated (r" .77) with an increased number of 6‘30. months, warmer tempera- tures throughout the year and a greater amount of precipitation during the April-June period. Leaf length was correlated with a greaterpercentage of annual precipitation occurring during the growing season, April-August, with colder winters, and with higher amounts of precipitation during the period April-June in relation to temperature for the same period. 145 In other words, growth characteristics were all positively cor- related with more favorable environmental factors of warmer tempera- tures and greater amounts of precipitation during the growing season. Or, they were negatively correlated with latitude, and elevation. Only leaf length was positively correlated with increasing east longi- tude. Region G, Northeast Germany and Czechoslovakia, had the lowest number of correlations. The actual and expected numbers of correla- tions are as follows: Siggificance Level 0.1 l 5 *** Per cent Per cent Per cent Value of r needed (27 d.f.) .368 .472 .580 Number of r's exceeding amount needed Actual 2 15 45 Expected (784 combinations) .78 7.8 39 Approximately 87 per cent of the correlations at the five per cent level of significance can be expected to be biologically meaningless. The correlations for this area are summarized in Table 29. Yellowing of the foliage was associated with higher temperatures, particularly during the winter and was negatively correlated with in- creased east longitude and increased elevation. This would be expected since the winters are colder to the east and at the higher elevations in this area. Table 29.--Correlations involving relationships between juvenile 146 characteristics and meteorological and geographical data of origin for region G, eastern Germany and Czechoslovakia. v T Juvenile Characteristics Meteorological H u u and .3 .3 .3 ‘3 >. Geographical 8 8 8 Is .3 m 3 Data .. 2:. a 2’. '6' a 2 33' as as as as ° 88 s G) -H H In H ‘\ H '\ "O U Q) H as as as as s as a V* J5 J9 J16 J31 J18 J27 J24 Actual precipitation in April -.53 NS NS NS NS NS NS May -.45 NS NS NS NS NS NS June NS NS NS NS -.37 -.47 .45 August NS NS NS NS NS NS .40 April-June -.48 NS NS NS NS -.39 .41 May-June -.44 NS NS NS -.38 -.43 NS July—Aug. NS NS NS NS NS NS .39 May-Aug. -.45 NS NS NS NS -.43 .45 Precipitation as per cent of total annual April-June NS .38 NS NS -.42 NS NS Latitude NS NS NS NS NS NS NS Longitude NS NS -.50 .59 NS NS NS Elevation NS NS -.54 NS NS NS NS Average temperature in January NS NS .57 NS NS NS NS February NS NS .47 .55 NS NS NS March NS .43 NS .55 NS NS NS April NS .42 NS .45 NS NS NS July NS NS .38 NS NS NS NS August NS NS .39 NS NS NS NS September NS .41 NS NS NS NS NS October NS NS .57 .41 NS NS NS December NS NS .49 NS NS NS NS July-Sept. NS .38 .38 NS NS NS NS Average annual NS .39 .48 .46 NS NS NS 6°months NS NS NS NS NS NS .89 Ratios Apr.-June prec. -.52 NS NS NS NS NS NS Apr.-June temp. - - JulyrAug;_preC. -.43 NS .38 NS NS NS NS July-Aug. temp. r I @368, .472, .580 for significance at the 5, l, and 0.1 per cent levels of significance. NS:§ Non-significant correlation. 147 Bud color is greener as the per cent of annual precipitation occurring during the growing season increases. Increasing amounts of rainfall in June appears to favor reten- tion of primary leaves and delay formation of secondary leaves. The development of a strong root system is highly correlated (r's .89) with an increase in the number of 6° C. months in the year. In region H, western Germany, eastern France and Belgium, the actual and expected number of correlations are as follows: Significance Level 0.1 l 5 Per cent Per cent Per cent Value of r needed (17 d.f.) .456 .575 .693 Number of r's exceeding amount needed Actual 9 26 57 Expected (784 combinations) .78 7.8 39 Approximately 69 per cent of the correlations significant at the five per cent level of significance and 30 per cent at the one per cent level are biologically meaningless. The correlations between juvenile characters and meteorological and geographical data of origin for region H are shown in Table 30. Third year seedling height is negatively correlated with increased amounts of the total annual rainfall occurring during the growing sea- son, April through June, and, in this region, is positively correlated with increasing latitude where the growing season temperatures are lower. 148 Table 30.--Correlations involving relationships between juvenile characteristics and meteorological and geographical data of origin for region H, western Germany, eastern France, and Belgium. Juvenile Characteristics Meteorological g 3 g 3 w, a, and >a H H H 0 11 cu u o o o s 0 Geographical ‘“ ° ° ° 3 3 '9 8 '4 Q Data .5 J“ 3) S» 3: g '3 ‘3': '6 g .5 U .13 Q! (00‘ a! HH 0 ECU 'UOO 00 Ha Hm How-IO 0) Gm “400 ”a “333333313032 35 <%.3 :% £213 :u.4 e.a> :n o a: Cluq cn n. .4.4 J6 J34 J9 J12 J13 J20 J18 J19 J27 J29 Actual precipitation in January NS NS NS NS NS .47 -.87 NS NS NS February NS NS NS NS NS NS -.91 NS NS NS April NS NS NS NS NS NS NS NS NS -.73 May NS NS NS NS NS NS NS .46 NS -.66 June NS NS .64 NS NS NS NS NS NS -.64 July NS NS NS NS NS NS NS NS NS -.57 August NS NS .63 NS NS NS NS NS NS -.55 September NS NS NS NS NS NS NS NS NS -.74 October NS NS NS NS NS NS NS NS NS -.57 November NS NS NS NS NS NS NS NS NS -.52 December NS .50 NS NS NS NS NS NS -.48 NS Total annual NS NS NS NS NS NS NS NS NS -.56 Sept.-June NS NS NS NS NS NS NS NS NS -.55 Oct.-Apr. NS NS NS NS NS NS -.75 NS NS NS ApriléJune NS NS .48 NS NS NS NS NS NS -.71 May4June' NS NS NS NS NS NS NS NS NS -.68 July-Aug. NS NS .54 NS NS NS NS NS NS -.57 May-Aug. NS NS .53 NS NS NS NS NS NS -.67 Per cent of total annual precipitation Sept.-June NS NS NS .48 -.51 NS NS NS NS NS Oct.-April NS NS NS NS NS NS -.75 NS NS NS April-June NS -.61 .49 NS NS NS NS .66 .64 -.47 May-June NS -.60 NS NS NS NS NS .60 .52 NS July-Aug. NS NS NS -.48 .51 NS NS NS NS NS r I .456 significant at the 5 per cent level of significance. r i .575 significant at the l per cent level of significance. - r u .693 significant at the 0.1 per cent level of significance. NS: Non-significant correlation. 149 Table 30.--Continued Juvenile Characteristics Meteorological . u r. u and a 2 .9. .2 “a a ".3 Geographical .3 8 8 8 .. .. 3 a 53. Data ~ m a) m g .3 'H'g 7. 8 .a u cw u‘ u: c o o u 'o c .c was smaaaoas°.g sacs ac: v! HWv-I\ «no HH '6 u 00 m: 833 g 8.35.2 £3.38 .3 88 33%; 331’. Vt _WW J6 J34 J9 J12 J13 J20 J18 J19 J27 J29 Average temperature in January -.68 NS NS NS NS NS NS NS NS NS February -.65 NS -.54 NS NS NS NS NS NS NS March NS NS -.67 NS NS NS NS NS ' NS NS April NS NS -.49 NS NS NS NS NS NS NS May NS NS -.68 NS NS NS NS NS NS NS August NS -.50 NS NS NS NS NS NS NS NS September NS NS .50 NS .62 NS NS NS NS NS October NS NS -.48 NS NS NS NS NS NS NS December -.47 NS NS NS NS NS NS NS NS NS NS NS -.60 NS NS NS NS NS NS NS April-June NS NS -.52 NS NS NS NS NS NS NS July-AugUst NS -.47 NS NS NS NS NS NS NS NS July-September NS -.53 NS NS NS NS NS NS NS NS Ratios Apr.-Junegprec. NS NS .58 NS NS NS NS NS NS -.66 Apr.-June temp. July-Augggprec. NS NS .52 NS NS NS NS NS NS NS July-Aug. temp. Latitude NS .59 NS NS NS NS NS-258 NS NS Longitude .61 NS NS NS NS NS NS NS NS NS Elevation NS NS .68 NS NS NS NS NS .67 NS 150 Late summer foliage yellowing is positively correlated with late winter lower temperatures and increased growing season precipitation. Green bud color is highly correlated (r’= .87, r t .91) with increased precipitation during January and February and to a lesser extent (r a .75) with increased precipitation during the October- April period. Leaf length is positively correlated with decreased amounts of precipitation during the period April-November and with decreased amounts of precipitation during the period April-June in relation to temperature. Since growing season precipitation decreases with lati- tude in this region the longer needles would occur at the higher lati- tudes, where the summers are also cooler. In regions J, K, M, and N, Yugoslavia, northeast Greece, southern France, Spain, and Turkey, there is essentially a Mediterranean climate. The actual and expected correlations in this region are as follows: Significance Level 0.1 1 5 Per cent Per cent Per cent Value of r needed (27 d.f.) .368 .472 .579 Number of r's exceeding amount needed Actual 8 61 136 Expected (784 combinations) .78 7.8 39 The correlations between juvenile characters and meteorological and geographic data of origin for regions J, K, M, and N are shown in Table 31. 151 Table 31.-~Corre1ations involving relationships between juvenile characteristics and meteorological and geographical data of origin for regions J, K, M, and N, the Mediterranean area. m Juvenile Characteristics Meteoro- g s s s s .. a t “81°31 2‘. s '3 '3 ’6 ° '3 .3 8 and Hen o o o o 3 .08 m .40 Geograph— u 4: 3&7 3’0 3 3’0 & 8 “313' on 153. 1°31 Jo's) 'obb'u'gogg 3% 3o .38 :13 mg 13‘; 53 Data as ssssszsasasass“ose 3:: 013 (0'4qu Flu-a Lin-c Ina) [141-4 NU 8'44 man mo. # J5 J6 J7/5 J31 J12 J13 J16 J20 J19 J17 J27 Actual precipitation in January NS. NS NS NS NS NS NS NS NS -.42 -.43 February NS NS NS NS NS NS NS NS NS -.43 -.47 March NS NS NS NS NS NS NS NS NS NS -.49 April NS NS NS NS NS NS NS NS NS -.40 -.45 May NS NS NS NS NS NS NS NS NS -.51 NS June NS -.52 .37 NS NS NS NS NS NS -.55 NS July NS -.56 .43 NS NS NS -.43 NS -.37 -.56 NS August NS -.65 .51 NS .37 NS -.44 NS -.43 -.52 NS September NS -.44 NS NS NS NS NS NS NS -.47 NS October NS NS NS NS NS NS NS NS NS -.43 —.46 November NS NS NS NS NS NS NS NS NS -.47 -.46 December NS NS NS NS NS NS NS NS NS -.43 -.53 Total annual NS NS NS NS NS NS NS NS NS -.49 -.42 Sept.~June NS NS NS NS NS NS NS NS NS -.47 -.44 Oct.-April NS NS NS NS NS NS NS NS NS -.43 -.46 April-June NS NS NS NS NS NS NS NS NS —.50 -.39 May-June NS -.44 NS NS NS NS NS NS NS -.54 NS July-Aug. NS -.60 .47 NS NS NS -.44 NS -.40 -.54 NS May-Aug. NS -.53 NS NS NS NS -.39 NS NS -.55 NS Per cent of total annual precipitation Sept.-June .37 .74 -.56 NS NS NS NS .43 .45 NS NS Oct.-Apri1 NS .48 NS NS NS NS NS NS NS NS -.41 April-June NS NS NS NS NS NS NS .38 NS NS .37 May-June NS NS NS NS NS NS NS NS NS NS .41 July-Aug. -.37 -.74 .56 NS NS NS NS -.43 -.45 NS NS May-Aug. NS -.49 NS NS NS NS NS NS NS NS NS f r I .368 significant at the 5 per cent level of significance. r ' .472 significant at the l per cent level of significance. r I .579 significant at the 0.1 per cent level of significance. NS . Non-significant correla tion 152 Table 31.--Continued Juvenile Characteristics Meteoro- 'g s s s s a 1. .. lexical a. a "a '3 *3 ° '3 .2 s and «1 o (J U U a .0 g ‘H .4 m Geograph- u .c u. ‘30 3’0 3’0 3’0 2 “3 I: o .c: '6 3 1°31 'U'ED ego-0'5) 33333. $3.338 :S ”E 3‘; 33 Data as 2:5 ssasssasaassosese co 3 card :2 fiaFlChvd k.a>Ih-a :3 o c3u4 an!» 0):; J5 J6VJ7[5 J31 J12 J13 J16 J20 J19 J17 J27 Average temperature in January NS NS NS .43 .40 NS .54 NS .39 NS NS February NS NS NS .39 .43 NS .54 .45 .51 NS NS March NS .39 -.41 NS NS NS NS NS NS NS NS April NS .43 -.52 NS NS -.39 NS NS NS NS NS May NS NS -.48 NS NS -.51 NS NS NS NS NS June NS .41 -.48 NS NS -.44 NS NS NS NS NS July NS .50 -.58 NS .39 -.47 NS NS NS NS NS August .39 .58 -.63 NS .44 -.49 NS .40 NS NS NS September NS .51 -.58 NS .48 NS NS NS NS NS NS October NS .51 -.57 NS .45 NS NS NS NS NS NS November NS .45 -.57 NS .42 NS NS .38 NS NS NS December NS .46 -.44 NS .37 NS NS .58 NS NS NS Monthly av. NS .48 -.55 NS .42 NS NS .38 NS NS NS April-June NS .40 -.50 NS NS -.46 NS NS NS NS NS May-July NS .43 -.52 NS NS -.48 NS NS NS NS NS July-Aug.' NS .54 -.61 NS .42 -.48 NS .37 NS NS NS July-Sept. NS .53 -.60 NS .44 -.44 NS .38 NS NS NS 6°months NS NS NS NS NS NS NS NS NS NS NS Ratios Apr.-June prec. Apr.aJune temp. NS NS NS NS NS NS NS NS NS -.47 NS July-Aug;_prec. July-Aug. temp. NS -.62 .51 NS NS NS -.45 NS NS -.52 NS LatitUde -052 -o67 .67 NS—OSB NS NS '0 62 “.46 NS NS Longitude NS NS NS -.50 NS -.86 -.37 -.50 NS NS NS Elevation NS NS NS NS NS NS NS NS NS NS NS 153 In this area, the ratio of first year height to seed weight, a measure of vigor, was positively correlated with increased growing season precipitation during June-August and with lower temperatures during the period March through December, but particularly durhng the months of July and August. Foliage color does not present any clear correlations. This was expected since foliage color was generally green to blue-green throughout the area. Earliness of winter color changes was correlated with a smaller per cent of the total annual rainfall occurring during the period July-August and with warmer winters. Early formation of buds was associated with increased precipi- tation during July and August. Early growth start was correlated with increased precipitation throughout the year, particularly during the months of June to August, and with increasing precipitation during the period April-June with respect to temperature increases. The presence of secondary leaves was negatively correlated with increased precipitation during the period October-April. Discussion The origins sampled within the Scotch pine distribution area covered such a wide variation in weather patterns that it was essential to treat the material as groups from more or less similar meteoro- logical conditions for study. The breakdown was not entirely satis- factory but further stratification would have resulted in groups with too few degrees of freedom for statistical study. The high percentage 154 of meaningless correlations resulted in reliance on the size of the correlation coefficient rather than on mere statistical significance. A study of the various weather patterns showed that precipi- tation in region A—F decreased from west to east with increasing summer rainfall to the east and at higher elevations. In regions G and H precipitation decreased from south to north in the growing season but increased during the same period with increases in ele- vation. In regions J, K, M, and N precipitation was greater in the summer than in the winter at higher latitudes but more in the summer than in the winter at higher elevations. Temperatures in region A-F decreased in the summer with latitude, and in the winter with increasing east longitude and elevation. In region G there was no pattern of correlation with latitude but winter temperatures decreased from west to east and with increasing elevation. In region B summer temperatures decreased with increasing latitude and winter temperatures increased, the reversal was true from west to east and with increasing elevation. Correlations of climatic data with the parental characteristics of cone, seed, and leaf measurements showed an overall pattern of greater temperature effects in the north area and greater precipitation effects in the southern area. Apophyses were shorter and thicker as temperature increased in the southern area, but were shorter and thicker as temperatures decreased in the north area. They were also shorter and thicker as winter precipitation decreasedin the northern area. .Because this is a protective mechanism for the seed it would be expected to 155 increase with less favorable environmental conditions within the area of growth. Seed length and width increased as average annual temperatures increased in the north. In the south they increased as growing- season temperature increased. However, larger seed were associated with more growing season rainfall in the north area and with less precipitation in the south area. The causal mechanisms were appar- ently different in the two areas since selection pressures would not be the same. In the north better environmental conditions during the growing season produced larger seeds, but in the south larger seeds were needed for survival during the dryer and hotter spring and early summer. The ratio of July-August precipitation/temperature was unim- portant in the northern area since precipitation was not a critical factor. In the southern area, however, this ratio was important. Decreasing precipitation/temperature ratio was positively correlated with increases in cone length, cone width, apophysis thickness, seed length and width, and cone scales. The hotter and dryer the July- August period the larger and thicker were the cones and seeds. Juvenile characteristics of seedlings grown in a provenance test in Michigan showed a few interesting correlations with meteoro- logical and geographical data of origin. Third.year height was positively correlated with increased pre- cipitation and temperature in the growing season in the north area, regions A-F. There were no correlations in region G. However, in region H third year height was negatively correlated with increased 156 precipitation (higher percentage of total annual precipitation) during the period April-June and with increased temperatures during the period July-September. There were no correlations for this characteristic in regions J’ K, M, and N. This reversal indicates that other factors may be responsible for the height growth, either at the point of origin (soils, planted stands, differing critical thresholds) or at the nursery site in the form of environmental interactions. Yellowing of foliage following the growth period was positively correlated with higher winter temperatures in the north region, higher summer and winter temperatures in region G, lower temperatures during the period February-May in region H and colder summer temperatures and higher winter temperatures in regions J,K,M, and N. This yellowing was positively correlated with more winter precipitation in the north region and less summer rainfall in the southern regions. Late bud formation was correlated with higher temperatures throughout the year in the north, not correlated with temperature in region G or H, and was correlated with the lower latitudes in the southern region. Late bud formation was correlated positively with increased precipitation during the period May-June in the north re- gion, with increased precipitation during the period April-June in region H. But it was negatively correlated with July-August precipi- tation in the south regions. In the north higher temperatures would indicate a longer growing season and later bud set. In the south, where the length of the growing season is not limiting, the late sum- mer dry period is the determining factor. 157 The early presence of secondary leaves is associated with higher yearly temperatures in the north, and greater percentages of the total annual precipitation occurring during the period April-June in region H and in regions J, K, M, and N. The more favorable environment in the temperature-critical north and precipitation-critical south ap- parently results in early presence of secondary leaves. Multiple Correlation Analyses An attempt was made to determine the effects of multiple cli- matic and geographic variables on the juvenile traits in order to better explain the variation in these characters. The initial multi- ple regression equations were set up on the best logical estimates of the variables responsible for correlations. Subsequent intelli- gence shows that new variables could have been used to better ad- vantage. Multiple correlations and regressions were run on third-year seedling height and on winter foliage color in December, 1961, as dependent variables with the following independent variables: Total annual precipitation North latitude East longitude Average monthly temperature Number of 6 degree months in the year The data for the four groups of stands were run separately on a correlation-regression program. The resulting output data for the Mediterranean group isshown in Table 32. The output data include 158 several factors which were useful in describing the effect of any one of the independent variables on the dependent variable. The beta weights, which are independent of units, give an accurate estimation of the value of the independent variable in determining the regres- sion coefficient. The F-value is used to test that any regression coefficient or beta weight is significantly different from zero. The R2 deletes show the coefficient of determination which would have been obtained had the independent variable not been included in the equation. Beta weights for the foliage color in December, 1961, indicates that the effect of each independent variable on the regression coef- ficient was in the following descending order of importance: Increasing east longitude Increasing north latitude Total annual precipitation Average monthly temperature Number of months per year averaging more than 6 degrees centigrade 2, coefficient of multiple determination, is equal to .378 2 The R for winter foliage color. Referring to the R deletes it can be seen that the deletion of the variable "number of months per year averaging 6 degrees centigrade" has an insignificant effect (R2 delete - .371). Increasing east longitude, on the other hand, reduces R2 to .146, indi- cating that this variable is the principle factor. In confirmation, a glance at Table 31 shows that east longitude is negatively correlated at the 5 per cent level of significance with winter foliage color, character P31, and that there are no other correlations. The use of 159 Table 32.--Multiple correlations and regressions for the Mediterranean stands for the juvenile characters, winter foliage color, December, 1961, and third-year height. Regression Beta R1 'W coefficient Independent Variable weight delete Winter Foliage Color, December, 1961, as Dependent Variable (R2 e .378) 69.499 -.002 grade per mm total annual precipitation -.286 .335 -.636 grade per degree of north latitude -.34U .306 -.217 grade per degree of east longitude -.591 .146 -.285 grade per degree of average monthly temperature -.132 .371 -.087 grade per month of av. 6 degree C. temperature -.036 .377 Third-year Height as a Dependent Variable (R2 I .290) 154.778 +. .034 mm. per mm total annual precipitation .529 .145 +-4.970 mm. per degree of north latitude .352 .213 '+ .579 mm. per degree of east longitude .209 .261 +-2.925 mm. per degree of average monthly temperature .179 .277 +-4.769 mm. per month of av. 6 degree C. temperature .260 .240 —Y 160 multiple correlations in this case would have assisted in developing a true picture of the multiple effects of the various variables mea- sured, but would not have changed the results of the simple correla- tions analysis. The results of the multiple regressions for winter foliage color showed that east longitude was the most important factor in the northern regions, that in Northeastern Germany, Czechoslovakian area east longitude had the greatest effect. However, even that effect was small. In western Germany, eastern France and Belgium north lati- tude was the most effective variable. In the Mediterranean area east longitude had the greatest effect on the multiple regression. Multiple regressions for third-year height showed that in the north there was little choice between the independent variables but that north latitude was the most important. In northeastern Germany and Czechoslovakia the number of 60 months and total annual precipi- tation were almost equally effective. In western Germany, eastern France and Belgium north latitude was most effective in the regres- sion, and in the Mediterranean area total annual precipitation ac- counted for the majority of the regression coefficient. In many instances the multiple correlations and regressions were not significant and the mass of data involved in this study discouraged their use since the simple correlations for the many variables involved were sufficient for the purpose. 3.3T.r .7T -.1' IF iJ-I CHAPTER IX TAXONOMY Shaw (1914) placed Scotch pine (Pinus sylvestris L.) in series Lariciones of the section Pinaster under the subgenus Diploxylon on the basis of large ray cell pits, cones dehiscent at maturity, and seed wing blades thin at base (Lariciones), an effective articulated seed wing and persistent fascicle sheath (Pinaster), and bases of the fascicle bracts decurrent and two vascular bundles in the leaves (Biploxylon). The cones, seeds, and leaves of Scotch pine, the subjects of this study, have been described by Shaw (1914) as: Leaves binate, from 3 to 7 cm. long; hypoderm inconspicuous; resin-ducts external; cones from 3 to 6 cm. long, reflexed, symmetrical or sometimes oblique, ovate-conic, deciduous; apophyses dull pale tawny-yellow of a gray or greenish shade, flat, elevated, or protuberant, and often much more prominent on the posterior of the cone, the umbo with a minute prickle or its remnant. Rehder (1940) states that the leaves are stiff, usually twisted, and bluish-green; that the cone-scale apophyses are flat, sometimes pyramidal, with a small umbo. The wide distribution of Scotch pine and the many isolated popu- lations along the southern and eastern limits of its range have re— sulted in considerable genetic diversity. Numerous Latin names have been proposed for various populations. Several authors have described infraspecific taxa without attempting to relate them to the remainder 161 162 of the species. Others have attempted to catalogue and describe the variation of the entire species. The more prominent of these monographic treatments are summarized below. The word "variety" is used here to describe a geographically limited population which is described on the basis of its phenotypic characteristics in the wild. The words ”race” or "ecotype" are re- stricted to populations whose genetic distinctness was studied by means of appropriate uniform-environment progeny tests. The descriptions are those of the authors whose publication is cited. Schott (1907) recognized the following geographic varieties. Although he did not cite authors, they have been added where it is certain that he was using a name in the same sense as an earlier author. var. lapponica (Fries) Hartm. Lapland, middle and northern Scandinavia, northern Finland. Crown spruce-like, slender, pointed, trunk predominately straight, branches inclined, ascending, or drooping; foliage dense, needles green, short, persisting for 4-7 years; cones more yellowish; apophyses convex and hooked; seed small, brown, seed wings reddish-brown. var. scotica Beissn. Scotland. Needles shorter and more blue- green than in the case of the principal variety (22523: gigg), 3-6 cm. long; cones shorter, about 3.5 cm. long, symmetrical, apophyses on the bottom of the cones smooth (flat), on the upper part more or less pyramidal. var . var 0 var. var. var. 163 borussica. Northeast German lowlands. Form between that of Eiggnsis (septentrionalis); cones violet-green to golden brown; seed not markedly brown but on the con- trary black, brown-mottled, seed wing gray-violet; needles larger than lapponica; the best of the pine in Germany, a very long period of life and greater height with less volume than in batava or superrhenana. batava. lower Rhine. Form between borussica and superr- henana; early flowering, longer needles, bears cones early, shorter life but fast growing. septentrionalis. South and west Scandinavia, northwest Russia. (syn. g. g. rigensis Desf.). Young trees very fast growing; bark very red; stem tall and straight; branches small, numerous twigs very dense; needles very glossy, long and flat. supperrhenana. Upper Rhine. (syn. g. g. rubra Endl., P. s. haguenensis Loud.) Form between bataga, Egggg- giga, and vindelica, about similar in height to bgggg- 3323 but with greater wood volume, poor trunk form and greater amount of branching, branches large and scat- tered in contrast with lapponica; foliage bluish-green, particularly in early stage, seeds large. vindelica. Northern pre-Alps, North Switzerland, Austria. Frequently straight growing and straighter than superrhenana, 164 although not as straight growing as the northern pines, needles short. var. pannonica. Western Hungary. Similar to superrhanana in habit, cone scales flat, crowned and reflexed, seed predominately black, smaller with superior ger- minating power over superrhanana; fast growing, pro- fuse flowering and regular ripening of cones. var. aguitana. Massif Centrale, France, and central Asia. Form resembling vindelica, early flowering time (April to early May), seed black, small, wings frequently pale violet, foliage deep-green, short and bright. Elwes and Henry (1908) stated that the following varieties, occurring in the wild state, have been distinguished. var. genuina Heer. This is the common pine, growing on good soil in Germany, southern Scandinavia, Poland, and northwestern Russia. Two subvarieties have been distinguished on the continent and in cultivation (rigensis and haguenensis). subvar. rigensis (syn. Pinus rigensis Desf.). Baltic Prov- inces of Russia, and, according to Willkomm occurs in north Germany, Poland, and Russia. Stem very straight and cylindrical, rising to a great height with few lateral branches; bark very red, stripping off above in very thin papery scales. 165 subvar. haguenensis Loud. Forest of Haguenau in Alsace. Vigorous in growth but defective on account of its tendency to form numerous irregular branches, so that the stem is not so clean and does not reach the same height as rigensis; bark is not red, and is not so fine scaled as in rigensis. var. scotica. Highlands of Scotland. Differs in redder bark of the stem; in the shorter, more glaucous leaves (1.5 inches long), often persistent for four years; and in the shorter cones (1.5 inches long), which are symmetrical, with apophyses usually flat near the base, tending to be pyramidal in the upper part of the cone. var. engadinensis Heer. Engadine Alps. Bark reddish; needles short, 1 to 1.5 inches long, thick and stiff, persistent to five years; buds resinous; cones ovoid-conic, 2 inches long, oblique at the base; apophyses convex on the outer side of the cone, umbo large and blunt. A small tree, rarely reaching 30 feet high. var. uralensis Fischer. Ural and Altai Mountains. Dis- tinguished by having short and stiff needles. 166 var. lapponica. (syn. Pinus lapponica Mayr). Northern Norway, Sweden, and Finland. This variety is con- sidered by Willkomm and Christ (Flora de la Suisse, 197, and Suppl. 31 (1907)) to be identical with vari- ety engadinensis, with which it agrees in the short, straight, stiff leaves, persistent for five years, in the resinous buds, and in the small cones with hook-like apophyses. Mayr, however, considers it to be a distinct species, and gives the characters which distinguish it from the common form of g. sylvestris without pointing out in what respect it differs clearly from variety engadinensis. var. nevadensis Christ. Sierra Nevada Mountains in south Spain. Needles broad, short, and stiff, very white on their flat surface; female flowers erect, purple- red; cone short stalked, nearly sessile, reddish-gray, lustreless, oblique; cone scales with very high curved pyramidal umbos on outer side of cone. var. reflexa Heer. High moors in Bern Canton, Switzerland, poor sandy soils of Prussia. Needles soft, about 6 cm. long; cones long and slender with long hooks (6 mm.) on the apophyses; cone scales dark red-brown and glossy; commonly a small tree with an irregular crown reaching 19 m. 167 Beissner (1909) listed Schott's varieties and also the following. var. genuina Heer. Haguenau area of France. This is the common form of the pine. Form usually pointed with tall stem; cones symmetrical and well developed; cone scales either flat or convex; bark ash-gray to gray-brown; buds gray or reddish-gray, female flowers pale or greenish-red. var. reflexa Heer. (Range and description as in Elwes and Henry). var. nevadensis Christ. (Range and Description as in Elwes and Henry). var. engadensis Heer. Engadine Alps, Tyrol. A branchy tree, slender pyramidal, prostrate form reaching a height of 10 m. with an umbrella shaped crown when with other trees; bark thin, reddish; needle short, thick and stiff, 3 cm. gray-green, sharp pointed and persistent 7 or 8 years, cones ovoid-conic, small, oblique at base, light yellow, symmetrical; cone scales glossy. yellow; apophyses convex on the outer of the cone, umbo large and blunt. 168 Beissner-Fitschen (1930) recognized Schott's varieties. They added some comments to some of the varieties and in addition listed varieties armena, turfosa, and katakeimenos. var. rigensis hort. (syn. Pinus rigensis Desf. Cat. Hort. Par. Arbr. II (1829) 61 and g. g, septentrionalis Schott, Forstwiss. Cbl. (1907) 278) South and west Scandinavia and Lithuania. Tree very fast growing when young. Bark very red; stem very straight and tall; branches small and numerous, twigs very thick; needles very long glossy and wide. var. scotica Schott Forstwiss Cbl. (syn. P. scotica Willd. Herb.) (1907) 278. Scotland. Needles shorter and more bluish-green than 3. _ylvestris, 36 mm. long; cones shorter, 3.5 cm. long, symmetrical; cone scales with the exposed portion flat near the base, tending to be pyramidal in the upper part of the cone. var. lapponica Fries K. Svenska V. et. Acad. (1888) (syn. Pinus friesiana Wichura Flora XLII (1859) 409; g. lapponica Mayr Fremdl. Wald- u. Parkb. (1906) 348. Lapland, middle and north Scandinavia, north Finland. (Description as in Schott.) var. engadinensis Heer Verh. Schweiz. Nat. Ges. Luzern (1862). In Engadine Alps, upper Inn valley by Martinsbruck and Finstermflnz. (Description as in Beissner (1909)). 169 var. armena. (syn. Pinus armena K. Koch Linn. XXII p. 297). Crimea, Caucasus, Asia Minor and Persia. Leaves are glossy green, acute, and densely cover the branches; staminate catkins nearly round or broad oval; conelets wide oval often with small points and the edge lightly serrate; young cones on thick very short penduncles, when matured they stand erect or horizontal; slightly glossy and yellowish-brown; upper cone scales usually with raised apophyses on upper half of cone slightly raised or flat on under side. var. turfosa Woerlein Ber. Bayr. Bot. Ges. III (1893). (syn. Pinus turfosa Willkomm). Bogs of northwest Germany, Baltic Sea shores on heather moors usually among Sphag- num, also on Scandinavian peninsula, Denmark, and northern Russia. Plant only .5 to 2 m. high, usually high shrub with a straight, erect central shoot, form- ing a small, flat crown later in life. The short needles (25 mm.) are curved and often fall in the second year. It produces numerous small cones with well deve10ped seed. var. katakeimenos Graebner Naturw. Wochenschr. XIV (1899) 546. Sand dunes on the Baltic coast, in Sweden, and northern Russia. A dwarf form with prostrate stems and branches up to 50 cm. long, but not raised more than 2.1 m. above the ground. The young shoots are 170 very long and slender, and bear normal leaves and cones o The varieties listed in Beissner (1909) as published by Schott (1907) and not listed above (septentrionalis, borussica, batava, superrhenana, vindelica, pannonica, and aguitana) were as described under Schott (1909). Rehder (1949) apparently used a very conservative approach and listed only four varieties. Varietal descriptions are from Rehder (1940). var. rigensis Loud. Bark very red, stem straight and tall. Silviculturally the most important variety. (Range and further description as in Beissner-Fitschen (1930)). var. scotica Beissner. Bark redder; leaves shorter, about 3.5 cm. long; cones shorter, about 3.5 cm. long. (Range and further description as in Beissner-Fitschen (1930)). var. lapponica (Fries) Hartman. 0f narrower pyramidal habit; leaves broader and shorter, remaining alive 4-7 years; cones more yellowish. (Range and further description as in Schott (1907)). var. engadinensis Hear. A slow growing pyramidal form with grayish-green, thick and rigid leaves 2.5 to 3.5 cm. long persisting for 7 or 8 years; cones oblique with 171 partly convex apophyses. (Range and further descrip— tion as in Beissner (1909)). Svoboda (1953) divided the varieties of Scotch pine into three principal groups on the basis of climatic type and topography. His first group, from central and northern Europe, was listed as, "Borovice severska (g. g. septentrionalis), klimatypy (areal vyznacen srafovane)," and included 16 varieties. The second group, from Asia Minor, southern and southwestern Europe, was listed as "Borovice hors- ka (2. g, montana), klimatypy (areal nesrafovan)," and included 15 varieties. The third group from south central Russia, Siberia, and north central Asia, was listed as "Borovice stepni (g. g, stepposa)," and included 4 varieties. The following are the 35 varieties recog- nized by Svoboda. var. lapponica Fries. (Range as in Schott.) (syn. Pinus lapponica Mayr.) var. scotica Schott. (Range as in Schott). var. norvegica. Subarctic and maritime Norway. var. suecica. Southern Norway, central and south Sweden (syn. var septentrionalis Schott). var. fennica. Finland. var. borealis. Northern Russia (extreme north is var. lapponica). var. uralensis (Fischer). (Range as in Elwes and Henry). 172 var. borussica Schott. (Range as in Schott). var. polonica. Mountainous area of south Poland. var. baltica. (syn. §.§. rigensis Desf.). (Range as in Elwes and Henry.) var rossica. Central Russia. var. ucrainica. Northwestern Ukraine. var. sarmatica (Zapal.). Southeast central Russia. var. baschirica. Northeast central Russia. var. obensis. Urals. var. jakutensis Sukac. Eastern Siberia. var. nevadensis (Christ). Sierra Nevada Mountains of south Spain. var. iberica. North central Spain. var. pyreneica. Central and western Pyrenees. var. aguitana Schott. (syn. P. g. avernensis Bayer). Massif Centrale of France. var. alpina. Upper Inn valley, southern Switzerland and northwest Italy. (syn. P. g. rhaetica BrUgger, P. g. engadinensis Heer.) var o V8! . 173 vindelica Schott. (Range as in Schott.) superrhenana Schott. (syn. P. haguenensis Loud.). (Range as in Schott.) var. pannonica Schott. (Range as in Schott.) var. hercynica Mflnch. (syn. P. g. bohemica (Sim.-Kav.)). var. var o carpatica Klika. Eastern Czechoslovakia. romanica. Mountains of northwestern Romania. var. rhodopaea. Mountains of northeastern Greece and southern Bulgaria. var. illxrica. Central Yugoslavia. var. pontica C. Koch. (syn. P. g. armena Koch, P. s. pontica var. Bayer.) Western Turkey. caucasica (Busch-Fisch.) (syn. P. hamata D. Sosn., P. 3. var. hamata Stev., P. kochiana Klotzch.). Northern Turkey, Armenian S.S.R., Georgian S.S.R. var. scythica. South central Russia in Steppe region. var. var. var . kasachstanica. East central Russia and Kazakh S.S.R. altaica Ledeb. Altai Mountains of Siberia. mongolica Litv. (Fomin). Mongolia. 174 Carlisle (1958) published a guide to the taxonomy of Scotch pine in which he names 13 geographic variants and an additional 13 possible variants. He considered varieties as collections of many forms (habit, cone, leaf, and flowering parts) where certain forms predominated to distinguish that variety when studied as a whole. His descriptions of the varieties he knew best were therefore com- parative rather than definitive. Table 33 summarizes Carlisle's views on the 4 varieties he knew best. The remaining 22 named varieties are listed but described only where additional character- istics have been added since the description was made by the origi- nal author. Carlisle's descriptions closely follow those found in Schott, Beissner, and Beissner and Fitschen. These 22 varieties are (with authorities as cited by Carlisle): var. borussica Schott. (Range and description as in Schott.) var. batava Schott. (Range and description as in Schott.) var. superrhenana Schott. (Range and description as in Schott.) var. vindelica Schott. (Range and description as in Schott.) var. hercygica Mflnch. Southern and mid-German mountains; to 1,0001m in the Schwarzwald and to more than 630-720 m. in mid-Germany. The stem is very straight and the crown narrow and pointed, with slender and flexible branches. In lower situations the stem remains straight, but the narrow, pointed crown is less marked. It forms valuable 175 ago» o>ooo .3 N cu H Ou maoo soHHoh nmwovmn guaca> omoaaocoua Mama .ouu suds vowcwu msouame ou :mHBoHHuh moan nu moaaumEOm .hmuu an uoz 3ouumz mamcomwu .EE maunm coma maoozmam .ao o.¢ somuwuoaouoummuo munch cum uoz omoum magmaowuucoumom noucwa a“ uoaoa> no now some soHHohnan:Souo msooaoaw naoHHoh coumo mo>mou .30 on .SOHHuhuamuum mums» use smog souuoz moficommma coma naounuaoouw .aa as .oaououoou «man .80 no shun» .asounnhouu mums» mum .msoosmau odoum woauoom mxumaom numcoq uoHou maoo mucoumwmuom ouwm navaa muowuw> wood can wood nacho moaumauouuoumno .Awnaav masseuse ho au>ww mm mowuowum> mafia nououm know no muwumwuouuoumno o>wumumaaou Hmmwoaaumnu.mn «snow 176 stands. A somewhat similar type occurs in northeast Germany, but var. hercynica is quite different from var. superrhanana (Mflnch, 1924). var. haguenensis Loudon. (Range and description as in Elwes and Henry.) var. engadinensis Heer. (Range and description as in Beissner (1909)). var aguitana Schott. (Range and description as in Schott.) var. subillyricum Corona. Trentino Province of Italy. var. pannonica Schott. (Range and description as in Schott.) var. nevadensis Christ. (Range and description as in Beissner (1909)). var. armena Beissner. (Range and description as in Beissner and Fitschen (1930)). var. altaica Ledebour. Altai Mountains of Russia. A tree 15 m. high with a compact pyramidal crown and short leaves. Carlisle stated that the following varieties had been mentioned in literature but that little is known of them except that they occur in certain geographic areas: A 177 var. caucasica Busch. (syn. P. caucasica Busch). Caucasus Mountains. var. altissima London. (Probably synonymous with var. caucasica). Caucasus Mountains. var. uralensis Fischer. Ural Mountains of Russia. Has short stiff needles. var. sibirica Ledebour. Altai Mountains of Siberia up to 900 m. and in Sungaria and Kirghiz in Russia. It has short leaves and cones with numerous scales, some being flattened and others bearing a prominent, pyramidal apophysis. (Carlisle considered it syn. with altaica.) var. jacutensis Sukacz. var. annulata Jurinson. var. funebris Kom. Mentioned by Kruberg (1937) as growing in eastern Siberia. var. mongolica. Mentioned by Kruberg (1937) as growing in Siberia. var. cretacea Kalenicz. Occurs in Russia on calcareous sites. Gaussen (1960) tended to discount the value of chemistry, wood anatomy and hybridization and to emphasize needle anatomy and the size 178 of pollen grains as a means of arriving at his species of the genus Pings. However, his determination of valid varieties was somewhat arbitrary. His monograph of the genus Pings lists the following 25 varieties of Scotch as authentic under 7 classifications or groupings (author citations are Gaussen's): I. Nordic variety. var. lapponica Fries. II. Baltic varieties. var. septentrionalis Schott var. rigensis Desf. III. Scotland variety. var. scotica Schott. IV. Varieties of the Western Mountains of Europe. var. hercynica Mflnch. var. aguitana Schott var. vindelica Schott var. pyreneica Svob. var. iberica Svob. var. catalaunica Gaus. var. brigantiaca Gaus. V. Varieties of the lower altitudes of the middle and southern areas. var. haguenensis Loud. var. batava Schott var. borussica Schott (considered var. sylvestris by Gaussen) var. engadinensis Heer var. nevadensis Christ var. Rannonica Schott var. carpatica Klika var. vocontiana Guinier and Gaus. VI. Non—geographic trees deformed by hostile environment. var. turfosa (Willk.) Woer. var. katakeimenos Graeb. 179 VII. Eastern forms. var. armena K. Koch (syn. g. kochiana Klotzsch) var. uralensis Fischer var. siberica Ledeb. var. yamazutai Uyek. Gaussen dismisses as not sufficiently known or characterized the following varieties although he states that they are probably varieties with special characters: var. sarmatica Zapal (along with the climatic types rossica Svob., and Baschirica Svob.) var. rhodopaea Svob. var. illyrica Svob. Table 34 summarizes the varieties of Scotch and the geographic regions where they predominate. Weidemann (1930) summarized the 1907 IUFRO Scotch pine European provenance tests. The results, based on 16 outplantings in Sweden, Germany, Belgium, Hungary, and the Netherlands, of 12 origins from the Ural Mountains of U.S.S.R., Northern Sweden, Scotland, Latvia, Prussia, northeast and western Germany, Belgium, Central Mountains of France, eastern and southern Czechoslovakia, and central Bulgaria, showed that geographical differences existed between the origins. The separate published reports summarized by Weidemann showed that the various provenances could be characterized as follows: North Sweden. Very slow growth and straight stems. Latvia. Moderately fast growth, straight stems, pointed crowns. Prussia. Similar to Latvian but with slightly more crooks. German. Fast growth, crooked stems, pointed crowns, variable form. 180 Table 34.-~Summary of the named varieties of Scotch pine and the geographic regions in which they predominate. 181 Auihgr and Qgte Elwes ghiissner and and Geographic Schott Henry Beissner Fitschen Rehder Svoboda Carlisle Stasskiewics Gaussen Area 1301 1908 1309 1930 19a9 1911* 1311 412§9L1fl3_§1___3292_ 1. N. Scandinavia, I. Russia LAPP LAPP LAPP LAPP LAPP LAPP LAPP LAPP LAPP 2. N. Russia SEPT RISE SEPT -— -- BORE -- -- SEPT 3. Cent. Norway, Sweden, Pin- land -- —- —- RICE -- -- SEPT -- SEPT b. 3. Norway, Cent. Sweden SEPT -- SEPT RIGE -- SUEC SEPT SUEC SEPT S. 3. Finland SEPT -- SEPT RISE -- PENN SEPT -- SEPT 6. 3. Sweden SEPT -- SEPT RICE -- SUEC SEPT SUEC SEPT 7. Norway SEPT -- SEPT -- -- HORV SEPT -- SEPT 8. Scandinavia, 1. Russia, H. Germany -- -- -- SEPT -- -- -- -- SEPT 9. S. and d. Scan- dinavia, NU. Russia SEPT RICE SEPT -- -- -- SEPT -- SEPT 10. La'via, Esth- onia SEPT RICE SEPT RICE RICE BALT RICE -- RICE 11. Poland -- -- -- -- -- -- -- POLO POLO 12. Bogs of NH. Germany and Poland -- -- -- TURF -- -- -- -- TURF I3. Dunes of 3. Haiti Coast -- -- -- KATA -- -- -- -- KATA 1;. Scotland SCOT SCOT SCOT SCOT SCOT SCOT SCOT -- SCOT 15. NE. Germany lowlands BORU -- BORU BORU -- BORU BORU -- -- lbs Se Polmd. ans. -- -- BCRU BCRU -- POLO -- SUBC POLO 17a Ea Cam. Germany -- -- -- -- -- HERC -- -- R033 18. Lower Rhine, NE France, Belgium BATA -- BATA BATA -- SUPE BATA -- BATA 19. Vosres Mtna., France BATA -- BATA BATA -- VINO BATA -- HERC 20. Alsace and Raa‘ad! areas of France and ' Germany BATA HAGU BATA BATA -- SUPE HAGU -- HAGU 21. Uryer Rhine SUPE -- SUPE SUPE -- SUPE SUPE -- VIND 22. N. Alps and Too'hills VINO -- VINO VIND -- VIND VIND -- VINO 23s 5e ‘rdi :on'e Jermany M'ns -- -- -- -- -- HERC HERC -- HERC 2L. En adine Alps, us'ria VINO ENGA ENGA ENGA ENSA ALPI ENGA -- BNGA 25. Swi'xerland, Bern Canion VINO REFL REFL VINO -- VINO -- -- ~- 26. SE. France, U. Switzerland, Jura Mina. -— -- —- -- -- VIND -- MERI VOCO 27. SW. Alps -- -- -- -- ¢- -- -- -- BRIG 28. 3. Swiss Alps and N. I'aly -- -- -- ~- -- ALPI SUBI -- -- 29. Cen'ral Massif, Francs AJUI -- AJUI AJUI -- AJUI AQUI HER! AQUI 30. S. Csn'ral Mas- sif, France AJUI -- AQU AJUI -- AJUI AQUI CAUS AJUI 31. Cent. and d. Pyrenees -- -- -- -- -- PYRE -— -- FY33 32. NE. 3;ain -- -- -- -- -- -- -- -- CATA 33. N. Cent. Spain -- -- -- -- -- IPER -— -- 195 34. 3. Con'. Jyain -- NEVA NEVA -- -- NEVA NEVA -- NEVA 35. E. Czechoslova- kia -- -- -- -- -- CARP -- MEET CARP 36. l. Hunrary FANN -— FAEN PANN -- FAN" PANN MEET PANN 37. Cent. Tuto- slavia -- -— -- -- -- ILL! -- -- -- 38. 3. Bulgaria, NE. Greece -- -- -- -— -- RHCD -- -- -- 39. Rulania -- -- -- -- -- scum -- -- -- LO. i. Turkey -— -- -- ARMS -- PORT ARM; -- ARMS L1. E. Turkey, Armen- ian and Georgian SJR., Iran, Crimea —- -- -— ARME -- CAUC ARMS -- ARME £2. Caucusus M'ns. of Seorgéan 33R and rkey -- -- -- ARME -- CAU CAUC -- ARC: L3. N. Cent.Russia -- -- -- -- —- n03; -_ -- -- 4. SE. Cent.Russia -- -- -- -- -- SAEZ -- -- -- L5. Ukrainian JJH -- -- -- -- -- UCRA -- -- -- L6. SE. Russia and Kazakh 35R -— -- -- -- -- KAJA -- -- -- ‘7e "8e cen's RUSU1. i. of Urals —- —- -- -- -— BAJC -- -- -- “as Siberia (“to 60° - 690), Ob to Yenesei R. -- -— -- -- -- 055" -- -- -- 59. Ural M’ns. -- URAL -- -- -- ossu URAL -- URAL 50. Sterro area of Russia -- -- -- -- -- SCYT -— -- -- 51. Altai Mina. Si- beria, S. *o Kirrhis and Sungaria -- URAL -- -- -- ALTA ALTA -- SIBI 52a I. COE.e 31b.r1a. Yenesei to Clenek and Lena rivers -- -- -- -- JAKU JAKU -- -- 53. E. Siberia to Hanchuria v— -- —- -- -- -- PURE -- YAMA 5L. Iongolia -- -- -- -- -- MONO MONO -- -- Key to varietal names: ALPXna, ALTAica, AQUI'ana, ARHEna, BALTica, BASChirica, BATAva, BORdalis, BORUsaica, BRIGantiaca, CARPatica, CAUCasica, CAUJsicola, ENGAdinensis, Fififlica, FUNEbris, HAGHenensis, HERCynica, THiRica, ILLYrica, JAFV‘ensis KAJAchstanica, KATAksmeinos, LATPonica, MERIdionalis, FKflfblisa ILVAdersis, NORVefiica, OEENsis, PAHHonica, FCLOnica, TFKTica, FTREneica, RHODoyaea, Ellirsis, Rfrjnica, HCJJica, SARMaYica, SCOTica, SCYThica, SIBIrica, SEFTontrionalia, SUBCarpaiica JUPIllyricun, SUsCica, SUPErrhenana, TURFoaa. UCRAinica. URALensis. VINDelica. VOCOntiana. YAHAsutai. 182 Scotland. Slow growth, variable form, many forked and a few crooked stems. - U.S.S.R. (Ural Mountains). Very slow growth, variable in form. France. (Central Mountains). Slow growth, variable in form. Belgium. Crooked stems, large branches, very fast growth. Mortality due to war, fire, and other causes reduced the amount of information that could be gained, particularly for the origins not included in the above summary. There is essential agreement in height growth between the relative rates of growth reported in provenance experiments by Weidemann (1930), Langlet (1936, 1937), Vincent and Polnar (1953), Wright and Baldwin (1957). The greatest growth rate was shown by the Belgian origin, followed by the origins from Germany, Nether~ lands, Latvia, Rumania, southern Scandinavia, Scotland, central Scandinavia, northern Scandinavia, in that order. In 1957 Wright and Baldwin reported on the New Hampshire planting of the 1938 IUFRO provenance test. They measured mor- tality, average height, diameter, branch diameter, basal sweep, lean, crooks, large crooks, porcupine damage, and fruiting. Sig- nificant differences between origins were found principally in height, and the per cent of trees with basal sweep, lean and crooks. The regions defined by these differences were: Northern Norway, Sweden, and Finland Central Norway and Sweden, and central and southern Finland South central Norway and Sweden Scotland 183 Latvia, Esthonia Poland, Germany, Czechoslovakia, and Hungary Belgium Romania Italy Netherlands Gerhold (1959) reported on analyses of color, pigments, and nutrient elements in needles of six of the geographic races repre- sented in the 1938 IUFRO plantings in New Hampshire. The six races tested were from the Netherlands, north central Germany, Poland, 0 Czechoslovakia, Sweden (640 latitude), and Norway (62 latitude). He found statistically significant differences in the concentra- tions of elements in needles between races. Trees from northern origins contained more chlorophyll in the summer and less in the winter than other origins. Gerhold stated that latitudinal effects (i.e. day length) etc. only could not explain the variation pattern. Staszkiewicz (1960, 1961, 1962) studied the populations of Scotch pine in Poland, Czechoslovakia, Switzerland. France, Scot- land, Sweden, and Finland and, on the basis of ten measurements taken from the cones, separated the populations as follows: Northern Sweden and Finland var. lapponica Central and southern Sweden and Prussia var. suecica Poland var. polonica Northeast Czechoslovakia and southern mountainous Poland var. subcarpatica Czechoslovakia, Switzerland, Italy, Hungary, central mount- ains of France var. meridionalis 184 Southern portion of the central mountains of France var. caussicola Wright and Bull (1963) reported the following geographic ecotypes of Scotch pine based on one year data for 64 provenances and three year data for 122 provenances grown in Michigan: Northern Norway, Sweden, and Finland North central Sweden Central and southern Norway, central Sweden, and southern Finland Southern Sweden and Latvia Siberia and Ural Mountains Poland South and East Germany, Czechoslovakia, and Austria Western Germany, eastern France, Belgium, Italy, and Hungary England East central France, central Austria, and Yugoslavia Greece, Georgian S.S.R., and Turkey Scotland Central Mountains of France Spain own» -4 .. ____.—.v,__‘l m”. CZO’IIMU . , . . ‘l'ii's. . '3 ZZF‘WHH Significant differences were found between provenances in seed measurements, height, foliage color, time of color change, bud color, form, and start and cessation of growda, lammas shoots, secondary leaf formation, leaf length, and ease of pulling (mea- sure of type of root formation and of root development.) The 1938 IUFRO test results and Michigan provenance test results are based on genetic differences in origins grown under a common environment. This study, based on the biometric differences in cone, seed, and leaf characteristics of specimens from living wild populations from 33 of the 122 origins included in the three year provenance 185 test reported on by Wright and Bull (1963), has confirmed the geographic ecotypes of the 1963 report except for areas B and D, which were not sampled. In addition, parental morphological differences indicate a separation of the TUrkish and Greek popu- lations and show that the Czechoslovakian population is different from the East German in cone characters and in leaf twist. It is also reasonable to assume that further differentiations could be made in Wright and Bull's regions E, G, H, and J, since they consist of areas which have been described by previous authors as separate entities, i.e., the populations in the Ural Mountains and in the Altai Mountains of the U.S.S.R., the populations in eastern France and Hungary, and the western Alps and Yugoslavia. Except in the instances of the isolated populations along the southern and eastern limits of the distribution of Scotch pine one would expect a gradual transition from one variety to another based upon the gradualness of the environmental differ- ences and the size and density of the populations. Selection is a slow continuing process and there have been relatively recent geological and meteorological changes wrought by the complex glaciation pattern of a large part of the area occupied by Scotch pine. While the genetic and morphological changes in Scotch pine may not be abrupt it is a fact that there are proven differences in populations. These populations can be defined biometrically and genetically. Table 35 is a list of geographical varieties based on genetic and biometric studies and descriptions based on field study and research by Ledebour (1833), Steven (1838), Schott 186 Table 35.--Varieties of Scotch pine (Pinus sylvestris L.) and the regions in which theywpredominate. Varietye; Geographical Area lapponica Northern part of Norway, Sweden, and Finland, and ’ northwest Russia. septentrionalis Central and southern Norway, central Sweden, southern Finland. rigensis Southern Sweden and Baltic Provinces of Russia. scotica Highlands of Scotland. polonica Poland. borussica Northeast German lowlands. hercynica South and east Germany and central Austria. haguenensis Western Germany, eastern France, and Belgium. vindelica Southeast central France, Switzerland, and western Austria. engadinehsis Engadine Alps, upper Inn Valley, and eastern Switzerland. aquitana Central mountains of France (Massif Centrale). iberica North central Spain. nevadensis Sierra Nevada Mountains of southern Spain. carpatica Northeastern and eastern Czechoslovakia. pannonica Western highlands of Hungary. illyrica Central mountains of Yugoslavia. rhodopaea Southern Bulgaria, northeast mountains of Greece. armena Northeast Turkey, Armenia, Georgian SSR, and Iran. uralensis Ural Mountains in USSR and area immediately to west. altaica Altai Mountains in USSR south to Kirghiz and Sungaria. mongolica Eastern Siberia and Manchuria. 'v 187 (1905), Mflnch (1924), Komarov (1934), Kruberg (1937), Sukatchev (1938), and others previously mentioned or cited as authors of the varietal names. The highly variable nature of Scotch pine makes it difficult to define varieties except on a population study basis. Most authors have pointed out that stands are mainly identifiable on the basis of the predominance of certain types over others rather than as distinctly different trees morphologically, particularly when speaking of contiguous areas. Some areas have easily identi- fiable forms such as those occupied by varieties lapponica, rigensis, Eggadinensis, nevadensis, or the noble race of the Vosges Mountains of France. Other varieties, however, are not so easily identified. The area of collection for the specimens of Linnaeus is in doubt. Annotations on sheet 1135.2 in the Linnaean Herbarium in apparently Linnaeus' hand, include the name "Kalm" with no other information as to the region of collection. Kalm, a student of Linnaeus, collected in Europe and in the United States. On another sheet (1135.1) is the annotation "ex Leche" indicating a Scandi- navian specimen. Elwes and Henry (1908) described Heer's variety genuine as containing two races, rigensis and haguenensis, and give the range of genuine as growing on good soil in Germany, south- ern Scandinavia, Poland, and northwest Russia. Many authors define varieties as being morphologically different from a standard or common Scotch pine without reference as to the locale of the typical variety sylvestris. Until a lectotype of Pinus sylvestris is desig- nated, it will not be possible to refer to a "typical" variety of the 188 species. It may also be necessary to typify a number of the other varieties as well as variety sylvestris. The synonomy which follows is based on the studies of geo- graphical varieties by previously cited authors, published prove- nance tests, and biometric studies of parental populations. The varieties accepted here are not considered complete since future investigations into serology, anatomy, etc., and the more mature characteristics of origins now being observed in provenance tests should provide additional information on genetic and phenotypic differences. ,3. sylvestris var. lapponica (Fries) Hartman, Handb. Skand. ed. ed. 5, 214 (1849) P. sylvestris var. lapponica Fries, Summa Veget. Scand. 1:58 (1846), nom. ,g. frieseana Wichura in Flora, 42:409 (1859) - Ascherson and Graebner, Syn. Mitteleur. F1. ed. 2, 1:345 (1912) pg. lapponica (Hartm.) Mayr, Fremdl. Wald- u. ParkaUme 348 (1906) pg. sylvestris var. borealis Svodoba, Lesni Dreviny A Jejich Porosty, Cast I, Praha. 203. (1953) .3: sylvestris var. frieseana Caspary (Wichura), Die Nuphar der Vogesen und des Schwarzwaldes. Halle, (1870). 3. frieseana Wichura. Flora XLII, 409 (1859). 189 Higher proportion of narrow crowns than var. scotica and var. septentrionalis with branches persisting low down on the stem; leaves not glaucous, often yellowish or violet in winter, short, averaging 41 mm., persisting 4-7 years; Cones short, mean length 3.6 cm; seed mean length 3.4 mm., mean width 2.1 mm., mean weight 4.4 mg. Mean pollen air-sac length 37 microns. Distribution: Northern Norway, Sweden, and Finland, north- western Russia to about 70 degrees North Latitude. Merging south- wards into var. septentrionalis. P. sylvestris var. rigensis Loudon, Arb. Brit. 4:2157 (1838). P. rigensis Loddiges. Cat. ex. Loudon, Hort. Beit. 4:387 (1830) nom - Desfontaines ex Moroques in Mem Soc. Agri. Orleans, Ser. 2, 25:43 pl. 5, fig. 10 (1885) pro. syn. E. rigensis Desfontaines, Cat. Hort. Par. Arbr. II (1829) 61. P. sylvestris var. baltica Svoboda, Lesni Dreviny a Jejich Porosty Praha (1953). P. sylvestris var. rigensis Hort. ex Beissner-Fitschen Nadelholz-Kundr (1930) 416. P. sylvestris var. rigensis (Desfontaine) Ascherson and Graebner Synopsis dev Mitteleur. Fl. 1. 13d Leipzig (1912). §._§ylvestris var. rigensis Loudon (Fischer)[ ? ] P. sylvestris var. septentrionalis Schott, Forstwiss. Cbl. 278 (1907) pro parta. 19C Crowns pyramidal, lance shaped, stems straight with ascending branches; bark pronounced reddish-yellow down to about 1-2 meters above soil; leaf mean length is 58 mm. without a pronounced glaucous sheen; seed mean length is 4.0 cm., buds are green-tan. Distribution - Southern Sweden and Baltic Provinces of U.S.S.R. P. sylvestris var. scotica Beissner, in Jager and Beissner, Ziergeh. ed. 2, 488 (1884). P. sylvestris var. scotica (Wi ldenow) Schott, in Klika, Siman, Novak, & Kavka, Jehlicnate, Nakladatelstvi Ceskoslovenske Akademie ved. Praha (1953) P. sylvestris var scotica Schott, Beissner, Handbuch der Nadelholzkunde. (1909) 431. - Schott, Forst- wiss. Cbl. (1907) 278 - Carlisle, A Guide to the Named Variants of Scots Pine (Pinus sylvestris L.). Forestry, XXXI: 204, (1958) - Svoboda, Lesni Dreviny a Jejich Porosty. Cast 1, Praha 201 (1953). P. sylvestris var. montana Sang, in Loudon, Encyclopaedia Plants. London (1855)- 2. scotica Hilldenow ex Endlicher Syn. Conf. 172 & Conf. (4 (1840)). P. sylvestris horizontalis Antoine (sec. Schott (1909))[?] P. horizontalis Don (of Forfar sec Schott (1909)) [?] 191 Older trees are flat crowned with narrow crowns rare. Leaves glaucous, 4.5 cm. long, twisted, green throughout the year; Persist 2-5 years; cones short, mean length is 4.5 cm. mean width closed 2.2 cm.; seed mean weight 7.2 mg., mean length is 4.5 mm., mean width 2.6 mm; buds green to green-tan; pollen air-sac length 38 microns. Distribution - Highlands of Scotland. P. sylvestris var. polonica Svoboda, Lesni Dreviny a Jejich Porosty. Cast 1, Praha. 206 (1953). Cone mean length 3.6 cm., mean width 1.9 cm. (Staszkiewicz (1960)). Distribution - Poland north of mountains along southern border. .P.‘sy1vestris var. borussica Schott. Forstwiss. Cbl. 278 (1907). ‘2. _ylvestris var. baltica Svoboda Lesni Typy. Praha. (1950). Crown variable, slender crowns in north and east with moderate branch development to heavier branches and wider crowns southward, vigorous growth; long life; leaf length cm.; seed mean length 4.0 mm., mean weight 6.2 mg. C Distribution - Northeast German lowlands. .2. sylvestris var. hercynica Mfinch, Allg. Forst- u. Jagdztg. c. 540, (1924). 192 Stem is straight and the crown narrow and pointed with slender, flexible branches except at lower elevations where the crown may be broader; leaf mean length 5.4 cm.; cone mean length 4.1 cm., mean width closed 2.1 cm.; seed mean length 4.3 mm. mean width 2.7 mm. Distribution - Southern and Mid-German mountains to 1,000 m. and in Black Forest to 630-720 m. merging into var. borussica in north- east. 'g. sylvestris var. haguenensis Loudon, Arb. Brit. (1838). ‘g. scariosa Loddiges ex Loudon, Encyc. Trees, 953. g. scariosa Loddiges Cat. of Plants in the Collection of Conrad Loddiges and Sons, London, ED XVI (1836). ,2. sylvestrig var. superrhanana Schott. Forstwiss. Cbl. 278 (1907). g. haggenensis Loudon ex Beissner. Handb. Nadglholzk. 228 (1891). .g. sylvestris var. Egggg Endlicher Synopsis Coniferarum (1847) Irv sylvestris batava Schott, Forstwiss. Cbl. 228 (1907). 'g. sylvestris var. genuina subvar. haguenensis Elwes and Henry, the Trees of Great Britain and Ireland. Edin- burgh (1908). pg. haggenensis Hort ex Lavallee. Arb. Segrez. 245 (1877). Stem form inferior to var. borussica but producing large volume of wood, tendency toward large horizontal branches; leaf mean length 6.9 cm.; cone mean length 4.9 cm., mean width closed 193 2.4 mm.; seed mean length 4.5 mm., mean width 2.7 mm., mean weight 6.7 mg. Distribution - Rhine River Valley, eastern France, and Belgium. ,g. syLvestris var. vindelica Schott, Forstwiss. Cbl. (1907) 278. lg. sylvestris var. vulgaris Bauhin [? ] Pu . genevensis Hort ex Carriere. Conif. ed. I. 372. Pu . genevensis Hort. ex Endlicher Synopsis Coniferarum. (1847) (4) g. sylvestris var. vulgaris ggnevensis Bauhin [?] Distribution - Southeast central France, Switzerland, and western Austria. g. _y1vestris var. septentrionalis Schott. Forstwiss. Cbl. 278 (1907) g. _y1vestris var. suecica Svoboda, Lesni dreviny a jejich porosty. I. Praha. 202 (1953). ,g. sylvestris var. fennica Svoboda, ibid. g. gylvestris var. horvegica Svoboda, ibid. Crowns variable but usually broad with branches persisting a short way down the stem, and a few narrow crowns with low per- sisting branches in eastern part of area. Leaves not glaucous, mean length 5 cm., persistent 3-6 years; cone short, mean length 4 cm.; seed mean length 3.9 mm., mean width 2.4 mm; mean pollen air-sac length is 37 microns. 194 Distribution - middle and western Scandinavia and northwestern Russia, merging into var. rigensis in south Sweden and Baltic Provinces of Russia. ‘g. szlvestris var. engadinensis (Heer) Hegi, Ill. F1. Mitteleur. 1:100 (1907). g. sylvestris subsp, engadinensis Ascherson and Graebner Syn. Mitteleur. Fl. ed. 2, l (1912). Small tree, reaching height of 15 m. with branches persisting low down on stem. Leaves short, less than 3.5 mm. thick, stiff, persistent 4-5 years; cones small with convex apophyses bearing large blunt umbos. Distribution - Engadine Alps, Tyrol, Upper Inn Valley, Switzerland. ‘g. szlvestris var. aguitana Schott. Forstwiss Cbl. (1907) 278. pg. sylvestris var. avernensis Bayer [?] g. szlvestris var. caussicola Staszkiewicz, Floristica and Geobotanica Ann. IX, Pars 2: 174-187, (1963). g. sylvestris var. pyreneica Svoboda, Lesni Dreviny a Jejich Porosty. Cast 1 213 (1953). Leaves 4.6 cm. long twisted; cones moderately long, mean length 5.0 cm., mean width closed 2.5 cm.; seed mean length 4.7 mm., mean width 2.7 mm. Distribution - Central Mountains of France (Massif Centrale). 195 g. gylvestrgg var. iberica Svoboda, Lesni Dreviny A Jejich Porosty Praha Cast 1. 212 (1953). ,g. gylvestris var. catalaunica Gaussen, Les Gymnospermes, Actuelles and Fossiles. Chap. XI, Travaux du Laboratoire Forestier de Toulouse. Tome II, sect. 1, vol. 1, part 2 (1960) 161. Leaves blue-green, mean length 4.8 cm. average twist about , 210 degrees; cone mean length 5.0 cm., mean width closed 2.5 cm.; seed mean length 5.4 mm., mean width 3.0 mm. Wflfiv.-. Distribution ~ central and northern Spain. g. sylvestris var. nevadensis Christ, Verh. d- Naturf. Ges. Basel, III Teil (1963) Heft. 4. ‘g. sylvestris var. hispanica Svoboda, Lesni Typy, Praha (1950). Poor timber form with short, broad, stiff needles. Distribution - Sierra Nevada Mountains of southern Spain up to 2,400 m. Southernmost natural population of Scotch pine (37 degrees north latitude). g. sylvestris var. carpatica Klika, Poznamky k rasam (ekotypum) Borovice v Ceskoslovensku. Vest Ceskoslov. Akad. Zamedelske 10: 2-3. 'g. giggestris var. carpatica 06min. (1937). g. _ylvestris var. subcarpgtica Staszkiewicz, Floristica et Geobotanica Ann. VII, pars 1, (1961) 158. 196 Leaf mean length 5.0 cm., twist approximately 280 degrees, cones, mean length 3.9 cm., mean width closed 2.0 cm., seed mean length 4.5 mm., mean width 2.7 mm. Distribution ~ Carpathian Mountains of Czechoslovakia and Poland and eastern Czechoslovakia. ‘g. sylvestris var. pannonica Sohott. Forstwiss. Cbl. (1907) 278. .E° _ylyestris var. meridionalis Staszkiewicz, Acta Botanica, Acad. Scientiarum Hungaricae, Tbmus VII, Fasciculi 3-4 (1961) 451-456. Pro parta. ‘g. sylvestris var. bohemica Siman (1923) Cone mean length 3.9 cm., mean width 1.9 cm. (Staszkiewicz, (1961)). Distribution - Western hill country of Hungary. Remarks - Staszkiewicz (1961) considers this population as part of his variety meridionalis on the basis of cone samples but states that it is possible that it may be a separate variety when other characters are analyzed. g. gylvestris var. illzgica Svoboda, Lesni Dreviny a Jejich Porosty, Cast 1, Praha. 212 (1953). g. sylvestris var. balcanica Svoboda, Lesni Typy. Praha, (1950) Pro parts. 197 Leaf mean length 5.8 cm., twist 280 degrees; cone mean length 4.7 cm., mean width closed 2.4 cm.; seed mean length 4.7 mm.; mean width 3.0 mm. Distribution - Central Mountains of Yugoslavia. ‘P. sylvestris var. rhodopaea Svoboda, Lesni Dreviny a Jejich Porosty. Cast 1. 219 (1953). ‘3. _ylvestris var. balcanica Svoboda, Lesni Typy. Praha, (1950), pro parta. Leaf mean length 4.1 cm., twist 180 degrees; cone mean length 3.8 cm., mean width closed 2.0 cm.; seed mean length 4.5 mm., mean width 2.6 mm. Distribution - Mountains of northeast Greece and southern Bulgaria. P. sylvestris var. armena K. Koch in Tchihatcheff Asie Min. 2 p. 496-499. Irv sylvestris var. Eontica K. Koch Q“ armena K. Koch, Linnaea 22: 296 (1849). sylvestris var. pontica Bayer [?] Izv Rd sylvestris var. caucasica (Busch-Fisch.) Izv hamata (Steven) Sosn. Fl. Tifl. 11 (1925). irv sylvestris var. hamata (Steven) Komarov. Fl. U.S.S.R. 1, 170 (1934) '51 kochiana Klotzsch ex C. Koch, Linnaea 22: (1849) 296. is sylvestris var. caucasica Fischer Irv sylvestris var. altissima Loudon 198 ‘2. altissima Ledebour ex Gordon. Pinatum 186. .20 hamata (Steven) Fomin. Mem. Acad. Sc. Ukraine, Cl. Sc. Phys. and Math. XI. 23 (1928). ‘2. sosnowskyi Nakai. Tyosen. Kaiho, No. 167 (Indig. Spec. Conif. and Taxads, Korba and Manchuria IV) 33 (1934) Leaf mean length 5.4 cm., twist 180 degrees; cone mean length 5.3 cm., mean width closed 2.7 cm.; seed mean length 4.9 mm., mean width 3.2 mm. Distribution - Crimea, Turkey, Caucasus, Armenian SSR, Georgian SSR and Iran. P. sylvestris var. uralensis Fischer [?] P. sylvestris var. obensis Svoboda, Lesni Dreviny a Jejich Porosty. Cast 1. 209 (1953). he . sylvestris var. sibirica Ledebour (1883) pro parts. he . padufia Ledebour ex Gordon. Pinatum 186. ‘P. sylvestris var. altaica Ledebour [? ] Distribution - Ural Mountains of U.S.S.R. P, sylvestris var. altaica Ledebour, F1. Altaica IV (1833). g. _ylvestris var. sibirica Ledebour, pro parta. ‘P. sylvestris var. altaica Komarov, P1. U.S.S.R., Leningrad (1934) ‘g. padufia Ledebour ex Gordon. Pinatum 186. Pro parta. 199 Distribution - Altai Mountains of U.S.S.R. south to Kirghiz and Sungaria. '2. sylvestris var. mongolica Litvinof, Hossiji Trudy Botan. muz. Akad. nauk l (1902). P. funebris Komarov in Acta Hort. Petrop XX (1901), Mitt. d. d. dendr. Gesllsch. (1903) 61. .2. yamazutai Uyeki Jour. Chosan Nat. Hist. Soc. No. 9, 20 (1929). he . takahasii Nakai (1939) [ ?] ‘P. sylvestris subsp. mongolica Fomin (1936) [ 2] ‘2. sylvestris var. lakutensis Sukac [? ] Distribution - Northeastern Mongolia, southeast central Siberia, Manchuria. Validity of the Named Varieties The International Code of Botanical Nomenclature (1961) re- quires that on or after January 1, 1953, the name of a taxon, to be validly published, must have been published through distribution of printed material containing a description or reference to a description previously published and a Latin diagnosis in a form which complies with that required by the International Code. In addition, valid publication on or after January 1, 1958, requires the designation of the nomenclatural type. The above requirements have not been met by Svoboda (1953) or by Gaussen (1960) and the names included in this discussion 200 cited with the above authors cannot be considered to have been validly published. Discussion The synonymy of Scotch pine is probably one of the most com- plicated of all the pine species. Linnaeus included Scotch pine in his Species Plantarum (1753) with the notation that the species had two needles per fascicle and glabrous primary leaves. Prior to and since Linnaeus's description there have been numerous tax- onomists, biologically oriented travelers, and horticulturists who have described a multitude of varieties and forms. Carlisle (1958) listed 144 varieties and forms in his resume of the Scotch pine. Recent provenance tests and biometric studies of Scotch pine indi- cate that there are differences between populations, particularly along the eastern and southern edge of the distribution area among the isolated stands, and within the more or less continuous popu- lation extending from northern Europe to southern Europe. This does not imply discontinuity within these areas but rather a dif- ference between the characters when measured at certain points. These samples are biometrically different and have been referred to as geographic varieties. Their existence has been proven gene- tically. Provenance tests conducted under uniform conditions of environment have proven the existence of races of Scotch pine. The results of this study and the biometric and provenance studies reviewed provide sufficient data for confirming some of 201 the varieties previously named or suggested, but are not considered sufficient evidence to preclude additional varieties or races, or further separations within the named varieties. Additional con- firming taxonomic information is needed in the fields of physiology, serology, phylogeny, genetics, morphology, and anatomy to properly separate definitively the varieties of Scotch pine. In addition, varieties of Scotch pine are continually being raised to the status of species by some as noted in Price (1963). Now that more is known concerning the natural populations careful studies should be made which include the examination or designation of type specimens so that the nomenclature may be ac- curately applied. CHAPTER X DISCUSSION Problems in Methodology The attainment of the objectives of this study required the acquisition of parental data and meteorological and geographical data of origin for as many of the original stands included in the 122-origin provenance test of Wright and Bull (1963) as possible. In addition intelligent and economical use had to be made of the acquired information in order to avoid an overpowering mass of data. The characters measured on the parental cone, seed, and leaf specimens were arrived at after a thorough study of the literature and of the specimens themselves. Scotch pine is pro- bably the most variable of the pine species and previous work on other pines was not always applicable to this species. Char- acter measurement was a tedious and time consuming operation. Only eight of the nineteen measurements proved necessary to es- tablish differences. However, important correlations were de- veloped because of the presence of the additional measurements. The dependence or independence of traits that are economically important, such as growth, needle length, and seasonal change of foliage color would not have been ascertained without the multi- plicity of characters measured. 202 203 The use of six foliage color measurements taken at dif- ferent periods of juvenile growth appears indefensible until one realizes that analysis of the data showed that change of color at various ages and times are controlled apparently by different mechanisms. The use of nineteen parental characters or combinations of characters also provided an insight into evolutionary trends in different areas that would not have been evident had fewer charac- ters been used. In the same way form ratios were not very useful in differentiating populations, but they were very useful in under- standing the variability relationships. The variability study of Scotch pine would have been expedited if climatic data had been available at the start. Unfortunately the climatic information was more difficult to obtain than the specimens so that several misadventures were taken before cause and effect relationships could be understood. The extremely wide distribution of Scotch pine over Europe and Asia, with the resultant highly vari- able environmental factors and occasional isolated populations, made the climatic study a necessity. The segmentation and movement of the original Scotch pine distribution by the glacial periods of the recent past were important factors to consider. The climatic data were organized into months, an arbitrary measure of time, and into natural periods of the four seasons. Ese sentially the same information and more accurate results were ob- tained by the use of natural seasons of winter, shoot development and elongation period, growth period, and food storage period as 204 climatic factors responsible for parent and juvenile characteristics based on physiological and biological phenomena. There was serious concern over the lack of evaporation-transpiration information. However, the use of the ratio of precipitation over temperature during periods of growth apparently sufficed. Here again, as in the use of multiple measurements and charac- teristics of the parent and juvenile specimens, the presence of apparently non—essential data provided an insight into the effect of the changes and of the timing of precipitation and temperature on parent and juvenile characters. In essence a form of multiple regression or correlation was achieved without formal programming. The statistical approach used would not have been possible had it not been for the availability of electronic computers. The cmmplicated analysis of variance of 689 individual specimens and their 19 characters by tree, stand, region and area of distribution and the calculations of their components of variance would have been impractical without the assistance of the computer. The use of punch cards for all data permitted rapid rearrangement of the information to explore suggested new avenues of approach. A study of sample size revealed that essentially the same information could have been obtained by the use of 10 cones and 20 leaves from each population. Mass collections of this magni- tude in populations with the high variability of Scotch pine would result in a maximum error of the mean of 10 per cent. 205 Value of Parental and Climatic Data Recognition of the genetically different populatigns.--The variation patterns in the cone, seed, and leaves of Scotch pine parent populations were most definitive in the following charac- ters. They are listed in descending order of their differentia- ting value: 1. Seed width. 2. Cone length. Cone len th 3' Ratio, Length ofilargest apophysis 4. Seed length. 5. Closed-cone width. 6. Leaf length. 7. Width of the largest apophysis. 8. Open-cone width. Analysis of the above characters showed that more than 85 per cent of their variability was attributable to between-region differences. The variation pattern based on the definitive dif- ferences of these eight characters indicated that the Scotch pine populations sampled could be separated into 10 regions. These grouped populations were identifiable entities based upon the significant differences existing between the regional means. The regions identified were: Region A. Northern Scandinavia in the vicinity of the Arctic Circle. Region C. Central and southern Scandinavia. Region G. Northeastern Germany and Czechoslovakia. 206 Region H. Western Germany, eastern France and Belgium. Region 1. England. Region J. Yugoslavia. Region K. Northeastern Greece (Macedonia), Region L. Scotland. Region M. South central mountains of France. Region N. Northern and central Spain. Region T. North central Turkey. The regions listed were found to coincide basically with those defined by Wright and Bull (1963) on the basis of three years of juvenile performance of Scotch pine in a 122-origin replicated provenance test at East Lansing, Michigan, with the exception of the Turkish population and the possible exception of the Czechos- lovakian. The juvenile characters measured in the provenance test pro- vided sufficient information for the establishment of races of Scotch pine based on discernible genetic differences under the con- ditions of a uniform environment test. The analyses of the parental data confirmed and strengthened this separation into geographic varieties. The analyses of the parental populations revealed that there was a remarkable similarity between the results of the two studies and that the regional pattern obtained by juvenile perfor- mance, when applied to the parent data, provided very high between region components of variance. Forecasting individualctree performance.--Eight stands in Belgium, Norway, and East Germany were analysed on a single-parent- 207 progeny individual tree basis to determine the relationships existing between single-parent-progeny performance in Michigan and character- istics of the parent tree in Europe. An exceptionally low number of parent-progeny correlations were found to exist, perhaps partially due to lack of comparable characters for comparison, genotype-environ- ment interactions, and silvicultural treatments of the parent stands. There were no correlations between growth rates, where they could most reasonably be expected to exist. These results indicate that the use of parental characteristics for prediction of juvenile per- formance in Michigan would be of little value. This would not be necessarily true of more mature information collected on the pro- venance test, particularly after outplanting in a sufficient num— ber of different environments to permit studies of genotype-environ- ment interactions. Analysis of silviculturally important traits such as clear stem length and diameter, crown shape and size, and foliage con- dition within parent stands supports previously reported relation- ships of crown diameter and stem diameter, and crown flattening with stem diameter and crown diameter. These relationships are those expected of trees approaching and reaching maturity. One stand, at RBvershagen, had twice the number of correlations present in each of the other four German stands. Selection in this stand for greater crown diameter would be accompanied by selection for greater stem diameter, fewer crooks, greater live-crown length, and less clear stem.length. In the other four German stands selec- tion for one trait would have relatively little effect on other 208 traits. Selection based on single traits would be expected to produce favorable results, but indirect selection would be slow as shown by the fact that the coefficients of determination based on the correlations of traits indicate that progress would be greater than 50 per cent in only 8 out of 145 correlations. Correlations between parent and progeny on an individual tree basis were exceedingly rare, 35 correlations significant at the 5 per cent level of significance among 1,260 combinations. Therefore, under the conditions of this test, two year juvenile performance characteristics under uniform environment conditions in Michigan as compared to parent performance in Europe, little progress would be achieved by attempting to base juvenile per- formance on the parental traits measured. As previously men- tioned though, later measurements on the provenance test may provide a better base for correlation and further research into genotype-environment interactions may provide a better understand- ing of the reasons for the lack of correlation. At the present little can be gained by parental selection. Progeny testing then becomes a necessity in a selection program for tree improvement. The relative lack of progeny-progeny correlations between characters infers that such traits as color, height, and time of bud set seem to have basically different physiological mechanisms. This result strengthens the previously noted situation in the parent-parent correlations in that one should treat each trait independently, and not expect selection in one trait to give a response in another. 209 Forecasting_stand:progeny and ecogype performance.--The within-stand correlation part of the study was accomplished by comparing the number of stands exhibiting the same correlations. In general cone characters were correlated among themselves, but there was little interrelationship involved in the majority of the stands. Changing climatic patterns tended to nullify expected correlations. The importance of selection for single traits was emphasized by the low coefficient of determination of one trait by another. When parental traits were analysed on a stand and regional basis many discrepancies were noted between the correlations ob- tained between regions and between stands. The increased number of correlations obtained on a stand basis pointed out that many of these apparent correlations were meaningless and due more to between region differences than to any gradual trends in cause and effect relationships. The deficiencies experienced in cor- relation by all stands was reduced by grouping the regions into areas of similarity of climate, a northern group, a Great Britain group, and a southern group. Comparison of the northern and southern group correlations showed that while there were the same number of significant correlations in each area the traits involved were different and at times the direction of the cor- relation was reversed. Cone characters in general were correlated in both areas but the basal angle of the cone was positively but not signifi- cantly correlated (r u .10) with apophysis thickness in the north 210 and significantly correlated negatively (r = -.77) in the south. Differences in the degree of correlation indicated basic form differences, probably resulting from differing environments and selection pressures.. The correlations resulting from unstratified data clearly indicates that there may be many apparent associations that may be meaningless, and that true correlations are increased over the meaningless by proper stratification of data. The experience gained in analysing the parental data was employed in analysing the stand parental-juvenile data. The data were analysed by grouping the stand data into three regions of apnrnximate similarity. Group 1, northern area, consisted of regions A, C, G, and H, group 2, Great Britain, of regions I and L, and group 3, Mediterranean area, of regions J, K, M, and N. While this grouping reduced the number of degrees of freedom it increased the validity of the correlations obtained. There were very few correlations, just as in the single-parent-progeny analysis, and only two instances of correlations between the same two characters in all three regions. In one of these instances seed length was significantly positively correlated with lateness of bud formation. In the other, height at age three was positively correlated with the ratio of cone length/apophysis length in the north and in Great Britain, but negatively correlated in the southern group. Significant correlations in two of the three groups give general trends and associations. 211 The correlations of precipitation and temperature data with geographic locations proved that the weather patterns of summer and winter varied between areas. In the northern area, for in- stance, the per cent of annual precipitation occurring during the growing season decreased with increasing latitude, but in- creased with increases in east longitude and elevation. In the southern or Mediterranean area the per cent of the total annual precipitation occurring during the period May to August increased with increasing latitude and decreased with elevation. These differences between area weather patterns were important in understanding correlations of climate and geography with paren- tal and juvenile characters. The parental characteristics showed an overall pattern of greater temperature effects in the north and greater precipitation effects in the southern area. Apophyses were shorter and thicker as temperature decreased in the north and shorter and thicker as temperature increased in the south. This seed protective mechanism was responding to a different set of critical factors. Large seed were associated with more growing season precipi- tation in the northern area and with less precipitation in the southern or Mediterranean area. In the north better environmental conditions during the growing season produced larger seed, but in the south larger seeds were needed for survival during hot and dry spring and early summer seasons. The juvenile characteristics of seedlings grown in Michigan had a few interesting correlations with meteorological and geo- graphical data of origin. 212 Third year height was positively correlated with increased precipitation and temperature during the growing season in the north, not correlated in eastern Germany and Czechoslovakia, negatively correlated in western Germany, eastern France and Belgium, and not correlated in the Mediterranean area. Yellowing or decrease of greenness of foliage in the winter was positively correlated with higher winter temperatures and more winter pre- cipitation in the north, higher summer and winter temperatures in eastern Germany and Czechoslovakia, lower temperatures during the period February to May in western Germany, eastern France and Belgium, and with cold summers and high winter temperatures and less summer rainfall in the Mediterranean area. The foregoing data reveal the dangers of applying infor- mation based on a limited area to a species as a whole or to the area of distribution. The diverse climatic situations in- volved within such a wide distribution pattern requires restraint in the use of local or regional relationships. Climatic data may provide a basis for inference in pre- dicting ecotypic performance but not assurance that the expected will be realized. Only population studies based on climatic infor- mation can be used to accurately determine the reaction of the plant to its environment. This is not to say that broad implica- tions can not be made on the basis of geography or climate as the limits of critical factors are approached, such as smaller trees as moisture or elevation becomes a critical factor to the develop- ment of a tree. 213 In much the same manner knowledge of parental performance can be used to develop only general relationships that can be expected to apply to ecotypes. The paucity of correlations ob- tained in the highly variable Scotch pine parent populations shows that the application of parental characteristics of a stand to other stands within an ecotype must be done with caution. The use of climatic or parental data to forecast progeny performance is also limited as evidenced by the lack of correla- tions, the lack of pattern in the correlations obtained, and the reversal of the directions of correlations in some instances. The general implications of the lack of pattern in corre- lations in a distribution wide or even a regional analysis of a character or between different characters is that progeny testing is the only assured method of acquiring performance information, and that parental or climatic data of origin is useful in develop- ing expectations, but not actual performance. The progeny test of Wright and Bull (1963) would seem to indicate at this stage that parent populations exhibiting good qualities in desired characters should be tested regardless of their origin. Trends in Evolution The general evolutionary trends in the Genus Piggg were listed by Shaw (1914) as moving from symmetric, dehiscent, and deciduous cones to oblique, serotinous and persistent cones, from uninodal to multinodal shoots, and from soft and thin to liquified and thick cones with larger seed with articulated wings or no wings. 214 Evolution within the Scotch pine, however, has been in re- sponse to local conditions and not always in the same direction, or toward the same objectives, unless these objectives be basically stated as survival. This study of Scotch pine emphasizes the point that evolution within a species as a whole is governed by its local environments, particularly in a widely distributed species covering a wide array of diversified climatic zones with their subsequent changes in selection pressures. Seed in the northern part of the distribution are smaller due to shorter growing seasons and lower temperatures, but more numerous to provide adequate propagation in a growing season where precipitation is not a critical item, but litter penetration and seed destruction by rodents may be. Seed in the Mediterranean area are large and well-stocked with the nutrients necessary to allow them to reach moisture in a less favorable growing season environment. The larger cones have developed under the more favorable development conditions in the optimum development area of Scotch pine and also as protective devices for seed in areas where greater protection was needed. The general evolutionary trends in the genus £2323 discussed by Shaw (1914) probably hold true when considering the overall long term development of the genus. However, evolution within a species is more directly associated with the environmental conditions, the plasticity of the species, and the response to selection pressures as discussed by Wright and Bull (1963). The distribution of Scotch 215 pine was radically changed during the recent complex glaciation of Europe and Asia and subsequent changes in environment. Some popu- lations were concentrated into islands of refuge. In some instances these populations have again expanded while others have remained as isolated populations, usually at the higher elevations. Selec- tion pressures due to changes in environment have slowly modified these populations. Great Britain was at one time completely within the distribution area of Scotch pine, but at present there are only remnant populations in northern Scotland. This situation and the climatic differentials involved in the various areas of population have resulted in a complex pattern of selection pressures. The area of distribution is not stable and the results of glaciation have not been fully realized. Changes due to varying selective pressures are slow. This study provides sufficient information to indicate the direction of development in response to local pressures on the cone, seed and leaves of Scotch pine. The direction of these evolutionary trends is toward sur- vival within an environment and the fact that Scotch pine is so widely distributed attests to its ability to adapt to changes in environment. The pioneer qualities of this species is indicated by its ability to survive, even in such hostile environments as the sand dunes and the bogs along the North and Baltic Seas, and propagate. 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The indirect determination of forest stand variables from vertical aerial photographs. Photo- grammetric Engr., 23(5):892-893. 225 Wright, J. and Baldwin, H. I. 1957. The 1938 International Union Scotch pine provenance test in New Hampshire. Silvae Genetica 6:2-14. Wright, J. W. 1963. Genetic variation among 140 half-sib Scotch pine families derived from 9 stands. Silvae Genetica. 12:83-89. Wright, J. W. and Bull, W. I. 1963. Geographic variation in Scotch pine. Silvae Genetica, 12:1-25. Zobel, B. J. 1953. Geographic range and intraspecific variation of Coulter pine. Madrone 12(1):1-7. Zobel, B. J. 1952. Geographic range and intraspecific variation of Coulter pine. Madrono 11:285-316. Zobel, B. J. 1961. Inheritance of wood properties in conifers. Silvae Genetica. 10:65-70. Zobel, 3., Cole, D., and Stonecypher, R. 1962. Wood properties of clones of slash pine. Presented at 1962 Annual Meeting Soc. Amer. For., Augusta, Ga. Mimeo. Zobel, B. J. and McElwee, R. L. 1958. Natural variation in wood specific gravity of loblolly pine and an analysis of con- tributing factors. TAPPI. 42:158-161. Zobel, B. J., McElwee, R. L. and Brown, C. 1961. Inter- relationships of wood properties of loblolly pine. Pro- ceedings of the Sixth Southern Conf. on Forest Tree Improvement, School of Forestry, University of Florida, Gainesville. 142-163. Zobel, B. J. and Rhodes, R. R. 1955. Relationship of wood specific gravity in loblolly pine (Pinus taeda L.) to growth and environmental factors. Texas For. Service Tech. Rpt. 11.32 pp. Zobel, B. J. and Thorbjornsen, E. 1961. Geographic, site and individual tree variation in wood properties of loblolly pine. Silvae Genetica. 9:149-158. John Lindley Ruby Candidate for the degree of Doctor of Philosophy Place of birth: Indianapolis, Indiana Date of birth: March 1, 1912 Marital status: Married Anne J. Bender, May 27, 1939 Children: Jo Anne L. Ruby, born May 10, 1944, married April 11, 1964 Education: Purdue University, BSF, 1934 U. S. Army Command and General Staff College, 1953 Michigan State University, MS, 1959 Michigan State University, Ph. D., 1964 Experience: Junior Forester, U.S. Forest Service, Region Nine. Acquisition, Recreation Survey and Plans, June, 1934 to July, 1936 U.S. Army, Field Artillery, July, 1936 to March, 1958 Retired on March 31, 1958 as Lt. Colonel. Teaching and research experience: Instructor, ROTC, Purdue University, Sept., 1940 to Oct., 1942 Assistant Professor, ROTC, Princeton University, April, 1952 to June, 1955. Research assistant and assistant instructor, Michigan State University, September, 1959 to August, 1964 226 227 Honorary societies: Xi Sigma Pi Alpha Zeta Scabbard and Blade Sigma Xi Foreign countries visited: Austria, Azores, Belgium, Canada, Denmark, England, France, Germany, Holland, Italy, Japan, Korea, Luxembourg, Norway, Portugal, Spain, Switzerland. t' «1 W .e:\, IV. nICHIan STATE UN [WWIWII‘[IWWIUIWHN [I 31293010 RQRIES SEES LIB "HIM” £22 5T7