ABSTRACT ANATOMICALvCHARACTERS AND CHEMICAL»COMPOSITION OF EUROPEAN BLACK PINE NEEDLES AS INFLUENCED BY GEOGRAPHIC ORIGINS AND NITROGEN FERTILIZATION by Chen Hui Lee European‘black pine seed, collected from 27 native stands throughout the species range, was sown in the Michigan State University Bogue Nursery in the spring of 1959. Each seedlot consisted of seed from.about 10 average trees per stand. In 1961 two-year old stock was used to establish the permanent test plantation at the Kellogg Forest, Augusta, Michigan following a randomized complete block design with 10 replications.r Each replicate has 27 four-tree plots. In the spring of 196h, two ounces of the RS-percent urea pellets were applied to the northernmost two trees in each plot and the southernmost two trees were left unfertilized. The plantation follows a split plot design with fertilizer treatment as the main factor, and prove- nance as the subfactor. Three growth, 19 anatomical and 18 chemical sets of data were studied. Western provenances were susceptible to winter burn in lower Michigan, Corsican trees being the most injured. Austrian trees were winter hardy. However, they were of considerably slow growth. No flowering was observed. Most characters were useful in the study of geographic variation pattern. Needle length, needle length/width ratio, number of resin canals, Chen Hui Lee - 2 and number of rows of stomata were of special importance and there was a steady east-west trend in those characters. Nitrogen fertilizer had little effect on growth and anatomical characters but significantly affected foliar chemical composition. Nitrogen fertilization significantly increased the uptake of N and Mn; and suppressed the absorption of K, P, Mg, B, Zn and Al. The result agreed best with that found in citrus trees. The level of foliar nutrient ions had nothing to do with growth rate. It failed to establish a relationship between foliar nutrient level and growth rate . Based on a combined judgment of growth, anatomical and chemical data it was possible to divide black pine into five distinct groups or races. The features which are most characteristic for each race and the correct varietal name were suggested and discussed. Var. poiretiana had most winter burn and characteristic curly leaves. Var. mnaica and var. cebennensis had moderate winter injury and long and soft needles. Var. musica differed from var. cebennensis in their yellow branchlets and longer needles. Var. austriaca was slowest growing among winter-hardy seedlots. Var. caramanica and var. 'Balkan' were winter hardy, however, var. caramnica was faster growing, had more rows of stomata and larger endoderm than var. 'Balkan' . ANATOMICAL CHARACTERS AND CHEMICAL COMPOSITION OF EUROPEAN BLACK PINE NEEDLES AS INPLUENCED BY GEOGRAPHIC 1‘ ORIGINS AND NITROGEN FERTILIZATION By Chen Hui Lee A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Forestry 1966 ACKNOWLEDGEMENTS I sincerely express my gratitude to Drs. J. W. Andresen, J. H, Beaman, C. L. Hamner, I. w. Knobloch, D. P. White and J. w. Wright, under whose guidance and supervision the present study was completed. Thanks are due to Prof. Luis Ceballos of Spain, Dr. Pierre Bouvarel of France, Prof.?Riccardo Morandini of Italy and Prof. Max Schreiber of Austria, for their valuable comments on the natural distri- bution map. I am also indebted to the late Dr. T. D. Stevens, former Chairman of the Forestry Department, for his approval of a doctoral program and for his generous assistance which enabled my wife and daughter to come to join me in early 1965. 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". ii TABLE OF CONTENTS INTRODUCTION ----------- ------- ----------------------.------------- DISTRIBUTION OF EUROPEAN BLACK PINE ------------------------------- Source of Information -------------------—----------------- General Range -------------------------------------.---.--- Climate ---------O----O----------.--------O- ..... ----- ..... ‘Detailed Range - ----- -- ------ - ----- --- -------------- - ------ Spain, France and Corsica, Italy, Austria, Yugoslavia, Albania, Bulgaria, Rumania, Greece, Russia, Turkey, Cyprus, Algeria. Planted Distribution ----------- ------ -------- -- -------- --- New Zealand, Great Britain, The United States of America, Argentina, South Africa. General Description of European Black Pine - ----- ---------- mmkm AND METHODS -Q---0---ouuoaococccncounnfO-cnouno. ..... on--- Seed Procurement and Nursery Practice, Description of Study Plantation, Field Measurements,.Anatomical Study, Chemical Study, Statistical Treatment. GrWth Traits -----------CCU-OCCOOOCOOODO ..... ago-coccnonon Growth Rate, Branch Angle, Winter Burn (1963-64), General Appearance Ana conical Characteria tics ccoccuunconnuuncuo-o-ouuuocaccq- Needle Length, Needle Curvature, Needle width, Needle Length/Width Ratio, Needle Color, Height/Needle Length Ratio, Number of Resin Canals, Position of Resin Canals, Serrations, Stomata(Ventral and Dorsal Rows), Dorsal] Ventral Stomatal Ratio, Hypodermal Layers, Sclerenchyma, iii Page 27 30 32 39 40 48 TABLE OF CONTENTS (CONT'D) Page Distance Between Fibrovascular Bundles, Endoderm (Length and Width), Endodermal Length/Width Ratio Folisr Chemical Composition ------------ ------ - -------- ---- 65 Determination of Error Variance, Nitrogen, Potassium, Phosphorus, Sodium, Calcium, Magnesium, Manganese, Iron, Copper, Boron, Zinc, Aluminun, NIP, N/K, K/P, Na/K, 08MB. Mil/PG EFFECT OF flTROGBN FERTILIZATION ------- - ------ --- ----------------- 77 Growth and Anatomical Characters --------------------- ----- 77 Growth Rate, Needle Length, Number of Resin Canals, Serrations , Number of Stomata, Hypodermal Layers P01“: Chuical magician can-cucocuonaocucooucccon-ac--- 81 Nitrogen, Potassium, Phosphorus, Magnesium, Manganese, Boron, Zinc, Aluminum, Overall Geographic Trend, Simple Correlation Between Mineral Composition and Growth Rate mmmum -------------------C-----------..-------------- ..... -O- 87 Taxonomic Interpretation of the Growth, Anatomical and mtri‘nt Data -C-----------.----------------O--.------.---- 87 Varietal Equivalents of the Groups ------------------------ 9o Pinus nigra Arnold, g. nigrs var. austriaca, g. nigra var. poiretiana, g. nigra var. cebennensis, g. nigra pygenaica, g. nigra var. caramanica, g. nigra cr. 'Balkan' SUMRY ------------------ -------------------- ----- ------- ---- ----- 100 LITERATURE CITED -------------------------------------------------- 102 11_T_A_ - ------------------- ------------------------ ------------------ 111 APPENDIX ----------------— ----------- ,-- ------------- ------ ----- ---- 113 iv Table 8a 8b 8c 8d 10 LIST OF TABLES Growth and needle characteristics of Pinus nigra from different origins ------------- ----- ---- ---------- --- ..... Needle characteristics of Pinus nigra from different origins .-CO------------------------------O---------------- Differences in surface needle characteristics associated With geographic origin ------------- -----------.---------- East-west variation patterns in number of layers of hypodermal and sclerenchymatous tissue in black pine needles ..... -----------------------------ODD-------------- Differences in internal anatomy of black pine needles ----- Statistical significance of the effect of fertilization and provenances on growth and needle characteristics - ----- Coefficients of variability for 12 mineral elements measured in European black pine (l plantation) and Scotch pine (3 plantations) ------------- ------------------------- Mineral element (N, R, P, Na) concentration in mature needles Of blaCk pme 0:181” ------------ ..... nun-uncncauu Mineral element (Ca, Mg, Mn, Fe, Cu) concentration in mature needles of black pine origins ---------------------- Mineral element (B, Zn, Al; NIP and N/K ratios) concentration in mature needles of black pine origins ----- Mineral element (K/P, Na/K, Ca/Mg, and Mn/Fe ratios) concentration of black pine origins ----------------------- Effect of application of nitrogen fertilizer on foliar mineral composition of European black pine measured 5 months later -------------------------------------------- Percent of characters by which various provenances differ firm Greek seedlots -- OOOOOO uces-u-n-c ----- u. ..... on- ..... -- Page 44 53 58 62 63 67 69 7O 71 72 83 89 LIST OF FIGURES Figure 1 Natural distribution of Burcpean black pine ------ - -------- 2 Natural distribution of European black pine in Spain ------ 3 Natural distribution of EurOpean black pine in Southwestern France and Corsica ---------------------------------------- 4 Natural distribution of European black pine in Italy ------ 5 Natural distribution of European black pine in Yugoslavia - 6 Natural distribution of EurOpean black pine in Greece ----- 7 Natural distribution of European black pine in Turkey ----- 8 Photomicrographs showing abnormal features of needle anatomy -cocnooouunuccuuunuucuoo ..... u ..... -Q-------------- vi Page 5 10 l3 17 21 23 25 SO LIST OF APPENDICES PRELIMINARY STUDY ------ ........ .-------- ...... -.--------------- IABLE A-l Results of statistical analysis of 5 needle A-2 to A-12 A-l3 to A-lB A-19 to A-28 A-29 A-30 to A-39 A-4O to A-42 Characteris tics .o-n-oocuoouco-c-u------------g Analysis of variance of 27 different black pine provenances for growth and needle characters from Kellogg planting (Augusta, Michigan) ----- Analysis of variance of 27 different black pine provenances based on one composite sample from Kellogg planting (Augusta, Michigan) ---------- Analysis of variance of 8 different black pine provenances based on one composite sample from Kellogg planting (Augusta, Michigan) ---------- Foliar composition of 27 black pine seedlots in percentage for N, K, P, Ca and Mg; as ppm. for other elements ---------------------------- Plot means (fertilized and unfertilized) and standard deviations for the measurement of 27 black pine seedlots ----------------- ------- Plot means (fertilized and unfertilized) and standard deviations for the measurement of 8 black pine seedlots ------ ----- ----- ..... ---- vii Page 114 115 116 127 133 143 146 156 CHAPTER 1 INTRODUCTION European black pine (giQEEDEiggg_Arnold) is a widely distributed species in southern Europe. It has been planted extensively in other parts of the world and in many areas is one of the most important timber trees. In the United States it has been used mostly as an ornamental but it also has a good potential for timber production. Provenance studies in other widespread species have shown a great amount of genetic variability associated with the geographic origin of the seed. European black pine appears to be no exception. During the past century taxonomists have described.many varieties and forms. A few European provenance studies have also shown striking differences among trees grown from seed collected in different parts of the range. However, taxonomic treatment by various authors differ considerably and the pro- venance studies were not replicated. Thus it seemed that new work*was needed to aid in understanding the genetics of the species. My study was undertaken as part of a long-range project for the improvement of black pine planted in north central United States. that project was started at Muchigan State University in 1958, when seeds were obtained from 27 different stands cmostly native) in Europe. The trees were grown in a research nursery for two years, then transferred to permanent field plantings at 14 locations in the north central states. My work is based in large part on one of those plantations located at the U. K. Kellogg Forest in southwesternNMichigan. 2 The Kellogg plantation is well replicated. Thus it is possible to separate enviromental from genetic effects and to make more definite conclusions on some taxonomic points than can be made with phenotypic data alone. Also, it is possible to incorporate various types of growth data with the morphological characteristics which are usually considered in taxonomic work. On the other hand, there are limitations. The trees are now 7 years old and represent only 27 different origins. Thus it was necessary to concentrate on certain types of questions which could be answered well with the material available. My study had three primary objectives. First was to determine the genetic variability pattern in many different morphological, growth, and physiological characteristics and to thus clarify the taxonomic status of the species. Second was to determine the underlying evolutionary factors responsible for the variation pattern and if possible to relate the morphology and growth characteristics to the internal physiology. Third was to obtain better trees for reforestation work. CHAPTER 2 DISTRIBUTION OF EUROPEAN BLACK rm Sources of Information My main source of information was a German researcher Roehrig's (1957) excellent work published in Silvae Genetics. Other sources such as Macdonald gt 5;. (1957) of Great Britain, Sexton (1947) of New Zealand, and Pritz Heller's (1951) World Porest Atlas published in Berlin were also very useful. Detailed climatic data were taken from Eiel's (1944) book. References pertaining to individual countries are cited as used. Another valuable source of information was obtained by corre- spondence with research foresters in Europe. I asked them to supply the latest and most accurate story of the distribution in their particular countries. They responded very cooperatively. Their maps were used to supplement recently published papers in the preparation of figures 1 through 7. General we European black pine m n_igg_a_ Arnold, syn. m laricio Poir.), has a large natural range in central and southern Europe and possibly northern Africa. The southern-oat range (35°l.) includes part of both Morocco and Algeria. Data on these north African stands are very limited, however. Eikli (1943) showed one occurrence in Morocco. Salvador (1927) and Sexton (1947) describedgthe occurrence in Algeria. This species is distributed from Spain in the west to Asia Minor and the Crimes in the east; from Algeria (Mount Edough) in the south to near Vienna in the north (Pig. 1). It has a 13-degree latitudinal range from 35° to 48°N. and a 48- degree longitudinal range from 6°w. to 42°E. That range is slightly larger than indicated by earlier writers such as Macdonald _e_t 31. (1957, 35° to 45°N.) or Sexton (1947, 35° to 48°N., now. to 42°E.). Black pine is predominantly a mountain tree, but can also be found at sea level along the shores of the Adriatic Sea. In Spain, Corsica, and Italy black pine is limited to highland areas between elevations of 2,600 to 5,000 feet above sea level. On the French mainland it is found between elevations of 800 and 2,600 feet. In Austria and Yugoslavia, it is distributed between 1,000 and 3,000 feet; in southern Greece and Turkey it occurs up to 4,300 feet; in the southern Taurus Mountains of Turkey it occurs higher than 6,000 feet. With respect to other cos-on Mediterranean species, black pine generally grows at higher elevations than maritime pine (Li_g_u_s_ pinaster Ait.) or most oaks (Quercus ssp.). It is generally at lower elevations than Scotch pine (Li_nus_ szlvestris 1"), although the tin species sometimes grow together. The black pine forests are always lower than those of beach (Legs; ssp.) or birch (Betula ssp.). Black pine grows on dry, usually calcareous soils. The soils are usually shallow and the slopes moderate to steep. It is sometimes able to grow in cracks in rocks. In Corsica, where it is found on deep soils, it grows most rapidly. The natural stands are very scattered. The length of the largest continuous distribution area is about 70 miles. In many cases the stands are less than a mile across. It forms pure stands under the most favorable growth condition and is mixed with other species at its lower and upper Pig. 1.-- Natural distribution of European black pine (prepared from figures 2 through 7). Can be reproduced by asking Michigan State University Photographic Laboratory Nega- tive File Number 65-1148. M 4—.— distribution limits. glimate The majority of the natural range is characterised by a Medi- terranean type of climate -- dry and hot summer with cool, moist winters. A few black pine stands are found at high altitudes where winter frost and snow frequently occur. Annual precipitation in the areas covered by black pine varies from 16 to 60 inches or more. However, there are considerable differ- ences among countries. In Spain the west and northwest coast stations recorded more than twice as much rainfall as those on the southeast coast. In Italy the western flanks of the Apennine Mountains facing the Ligurian and Tyrrhenian Seas receive much more rainfall than the eastern flanks facing the Adriatic Sea. A striking case of the lee-side effect is the aridity of the innermost valleys of the Alps. In the very heart of the Alps such as thaupper Rhone Valley the necessity of artificial irrigation has been known since Roman days. In the Balkans the entire coast of Yugoslavia, Albania, and Greece is extremely rich in winter precipitation. Because of orographic and frontal nature, the extreme of 180 inches was recorded around the Bay of Eotor. Enormous differences between.windward and lee slope precipitation were also recorded in the Caucasus Mountains. There is 100 inches at Datum, but only 10 inches at Baku. Most of the precipitation falls in the‘winter, when the mean monthly precipitation.ranges from 4 to 8 inches. In Spain, trance, Italy, Yugoslavia, Albania, Greece and Turkey, mean monthly precipitaion during the summer months seldom exceeds 2 inches. In Cyprus and.North,Africa, mean monthly precipitation during the summer falls as low as 0.4 inch. 8 There is a decreasing length of dry season from the south to the north. It has a period of five months in southern Spain; four months in.Asia Minor, Greece, and Sicily; three months in southern Italy; and two months in central Italy. Precipitation in a form of snow occurs throughout the natural range of black pine. Generally there are 10 to 20 days of snow per year in the Mediterranean area. (As many as 50 to 100 days of snow per year have been recorded in the interior mountains and plains of Yugoslavia. If the annual range of temperature is defined as the difference between mean temperature of the warmest and the coldest months, much‘of the Mediterranean coast has a typical marine climate with an annual range of temperature of less than 60°F. The coasts of France and Italy, the eastern coasts of the Adriatic and Ionian Seas and the shores of Asia Minor fall in a transitional zone from marine to continental climate with an annual range of between 60° and 68°F. A continental type of climate is found on the shores of Bulgaria, Romania and Russia where the annual range a 68° to 105%. The increase in continentality u from west to east. Summers are warm everywhere on the coast, winters are mild in the west and severe in the east. Winters are very mild in Sardinia and Corsica. W ‘ gpggg (Fig. 2).-- European black pine is commonest in the eastern part of Spain. only a few stands occur west of Madrid. It is absent as a native tree in Portugal. However, some plantations have been success- ful on the western side of the Peninsula (Taylor, 1949). According to Professor iuis Ceballos of Ciudad University, Madrid (personal corre- spondence. October 5, 1964) there are approximately 875,000 acres of natural black pine stands in Spain. There are three important natural 9 distribution areas.. One occurs in the highlands of Aragon in northern Spain (Province Teruel); a second is found in the mountains of Cuenca (Province Cuenca); third lies in the mountains of Segura and Cazorla in southeastern Spain (Province Jean). The climate is dry and extreme, modified in part by the relief of the mountain ranges. Annual precipitation is between 16 and 28 inches throughout the natural range in Spain. The highlands of Aragon receive 16 inches of annual precipitation; the mountains of Cuenca, about 24 inches; the mountains of Segura and Cazorla, between 20 and 28 inches. The precipitation is unevenly distributed with a regular period of summer drought and a further dry period in autumn. Most of the precipitation falls during winter. According to Biel (1944), the monthly mean precipi- tation during July and.August is less than 1.2 inches. During winter (October - December) it is between 4 and 5.6 inches. In highlands of Aragon where the climate is severe, the mean January temperature is 36°F., and the mean July temperature is 68°F. The climate of Cuenca is milder than that of Aragon. Segura and Cazorla have higher summer temperatures than Aragon or Cuenca. Segura and Casorla have mild winters with infrequent frost and snow. Black pine is most common between elevations of 2,600 and 5,000 feet. In Aragon, it occurs between 1,650 and 4,600 feet; in Cuenca, it is mostly found between 3,300 and 5,000 feet; in Segura and Casorla it is . usually seen between 3,300 and 6,000 feet. Black pine usually grows on calcareous soils on steep slopes. The soil is, according to Stevens (1934), derived froanurassic and cre- taceous formations. It is also able to grow on siliceous soils in Spain (personal correspondence with Professor Luis Ceballos of Ciudad University, 10 Fig. 2.-- Natural distribution of European black pins in Spain (personal correspondence from Professor Luis Ceballos, Ciudad University, Madrid. Scale, 1 : 6,250,000) «44 O (\l C O 0 CO C a“ (O CO o O 12 Madrid). Growth is vigorous even under the conditions found in Aragon. According to Eoehrig (1957), a 150-year old black pine reached 66 feet in height and 1.7 feet in diameter. In Cuenca tree growth is very satis- factory. Straight stsms and slender branches are characteristic for that area. Although tree size is smaller in Segura and Cazorla, yet the pro- duction of timber is noteworthy. Black pine is'a mid-mountain species mixed with maritime pine at its lower range, and mixed with Scotch pine at its higher range. France and Corsica.(Fig. 3).-- Black pine occurs on the French- side of the Pyrenees starting from Pradas (Dept. Pyrenees - Orientales) and ranging south to the region of Gorneilla-de-Conflent near Vernst-les- Bains and Escaro. Separated about 90 miles from the eastern Pyrenees lies the next black pine zone in the most southwestern outlet of the Cavennas and northern Montpelier in the neighbor of St. Guilhem-le-Desert (Dept. Herault). It also is found on the southeastern and southern borders of the Cevennes, especially in Col d'Uglas and Gagnieres. In Corsica, it is found on the south slope of high mountains in the northern half of the island and west of Corte city. There is also a considerable black pine forest in the southeastern part of the island. The total area of natural bldck pine stands is about 30,000 acres. Annual precipitation at Pic du‘Midi station on the French Pyrenees is about 63 inches. The lowest mean.month1y precipitation (3 inches) occurs in July. The highest is in January (7.2 inches). Annual precipi- tation at a coastal station such as Perpignan (on the Gulf of Lion) drops sharply probably due to lee-side effect. It has only 20 inches of annual in Fig. 3.-- Natural distribution of European black pins in Southwestern France and Corsica (Debazac, 1963 and 1964 and personal correspondence from Dr. Pierre Bouvarel, National Institute of Agronomy Research, Nancy. Scale, 1 : 2,000,000). UHmMOU Om Ohms O.” . \\..m aloe/coo.\lai Unmomm Z O H A h C m A D 0 \ 303$ / dd... H50 . “m mom acvmw MUZAmmm om: we 15 precipitation with 0.8 inch in July and 2.4 inches in October. Data on the Cevennes area are not available. On Corsica, the annual precipitation is abundant, ranging between 55 and 60 inches. Most occurs during the winter. At coastal stations, the highest monthly mean is 73°F. (July), the lowest monthly mean is 45°F. (January). Thmperature data for the Cevennes are not available (Biel, 1944). The mean annual temperature on Corsica is 54°F. 'Winter temper- ature is usually low. Autumn frost is frequently the case. Black pine finds its upper distribution limit around 3,300 feet on the French Pyrenees, but is more common between 1,300 and 2,000 feet. The optimum altitude in the Cevennes is approximately 2,300 feet. In Corsica, the best development occurs between 2,600 and 3,900 feet, where it forms pure stands. The soil is of limestone nature in the Pyrenees. It is of dolo- mite, quartz sandstone and conglomerate nature in the Cevennes. Dry, stony and shallow soils are characteristic of both areas. rh- soils are welldweathered and deep, and derived from granite or porphyry in Corsica. Trees seldom reach 60 feet height on the French mainland. The stems are generally crooked. Under the most favorable conditions in Corsica, trees reach 120 feet or more in height and over 3 feet in di- ameter macdonald, gt _a__l_., 1957). Growth is rapid there. On the French mainland, black pine is oftenlmixed with Scotch pine and other broadleaved species. In Corsica, it is mixed with chestnut at its lower limit, and with beach at its higher limit. 16 L§3;1(Fig. 4).-- Black pine is found in the eastern Italian Alps (Professor Riccardo'Morandini of Silvicultural Experiment Station, Firense. Personal correspondence. September 1, 1964). Small isolated occurrences are found in the Abruszi Mountains (Province Aquila) of Central Italy. Large stands occur in Aspromonte and in the Sila of southern Italy. In Sicily, it occurs on Mt. Etna. The area of black pine in southern Italy and Sicily is about 87,500 acres. In southern Italy, annual precipitation ranges from 50 to 80 inches, the majority of it falling in winter. Summer is dry. The weather record taken at Chieti in Central Italy shows 40 inches of annual precipitation ‘with November the highest month (5 inches) and July the lowest.month (1.7 inches). Corisia, the nearest station to natural black pine in the eastern Italian.Alps, has annual precipitation of 64 inches. The wettest month is October (6.8 inches) and the driest months are January and February (3 inches in each). About 5.1 inches is recorded for July. The mean July temperature in Catania (eastern Sicily) is 79%., the mean.January temperature is 51°F., the annual range being 28°F. The meathuly temperature in Chieti (Central Italy) is 73°F., and the mean January temperature is 39°F., the annual range being 34°F. The mean July temperature in Corizie is 73°F., the.mean.January temperature is 37°F., the annual range being 36°F. The annual range of temperature shows an increasing tendency from south to north.. Black pine is generally found between elevations of 3,300 and 6,000 feet in southern Italy. No data are available to indicate its elevational range in both Central Italy and the eastern Italian.Alps. In southern Italy, soils are derived from crystalline schist and granite, which support vigorous growth of black.pine forest. On the 17 Fig. 4.-- Natural distribution of European black pine in Italy (personal correspondence from Professor Riccardo Horandini, Silvicultural Experiment Station, Firense. Scale, 1 : 7,200,000) n O O 0 0 mo C6 A. 0 no no 4 4 11+ A.” no 79 kI 1 o . I s a l u6 06. 4‘ 1 2 u ‘ U 34 ,A\ , 4 I u e... weee\\ 1’ v 1 3.. ”a .. I Y o ... l A 2 an... I L 0 II. a A ‘ Oh u. 1 .. T o ... o f .. 4|. (c... .— n. 00 1 e. U as a; N“ A .\ O .“.Jf Q." .‘Q 8 O 0‘ 0‘. fe 0 o o a nmc 06 4. O 8 6 I: 4 I . A: 7t”. 71% 19 slope of Mount Etna, Sicily the soils are derived from volcanic material. According to Stevens (1934), there are great variations in cone types, needle characters, bark and branching habit. he considered the Italian black pine to have gradations from the typical Austrian pine to the typical Corsican pine. Austria (Fig. l).-- A large natural stand occurs south of Vienna. The second, less well known, occurrence is in the Central Alpine Zone of southwestern Austria. In,the Karawanken.uountains of southern Austria there are a few small natural stands (Professorluax Schreiber, nochschule fuer Bodenkultur, Institut fuer waldbau, Wien. Personal correspondence. September 2, 1964). The total area of black pine stands in.Austria is estimated to be 62,500 acres. Annual precipitation ranges between 22 and 28 inches. Host of the rain comes during the summer months. Snow occurs infrequently and lasts short period of time. Mean annual temperature in the region is 48°F.; mean January temperature is 28°F. and mean July temperature is 66°F. Soils are weathered products of Alpine limestone. Occasionally they are derived from.dolomite or gneiss. They are uniform in nature. Yugoslavia (Fig. 5).-- The natural range extends from northwest to southeast in Yugoslavia, mainly in the western part of the country facing the Adriatic Sea. Numerous stands are found in Dalmatia, Bosnia, and Herzegovina. Sarajevo (elevation 1,800 feet) in Central Yugoslavia is the nearest station to the natural stands from which seeds of HSFG 415 (mount Tara, Kremna, Serbia, Yugoslavia. Elevation 4,000 feet) were collected. Annual precipitation at Sarajevo is 39 inches. The precipitation is 20 equally distributed throughout the year. The highest monthly mean (4 inches) occurs in June, the lowest in February (2 inches). The temperature pattern at Sarajevo is as follows. The mean July temperature is 66°F., the meah‘January temperature is 30°F. Albania (Fig. l).-~ Black pine occurs in the northern part of the country.. Bulgaria (Fig. 1).-- In the Rhodope (southern Bulgaria) and Pirin (southwestern Bulgaria) mountains some black pine stands can be found. The greater part of the Bulgarian forest, largely conifers, is in those areas. Rumania (Fig. 1).-- Black pine stands occur north of the Danube River in the vicinity of Iron Gate. 95339; (Fig. 6).-- Black pine forest occurs in Macedonia, Thrace, Thessaly, Bpirus, and the Pleponnesian Peninsula. It is found between elevations of 2,800 and 5,300 feet. .At its upper limit there is a zone of overlap with Scotch pine. Natural black pine stands occur in various soil conditions. Soils of schist and granitic nature are con-non in addition to calcareous ones. gaggig,(Fig. 1).-- Black pine occurs in the southwest of the Crimean Peninsula and the northwest parts of the Caucasus Mountains. gggkgz_(Fig. 7).-- Black pine occurs in the Pontus Mountains along the Black Sea and also in the Taurus Hountains of souhern Turkey. In the Pontus, it is found at elevations between 1,700 and 5,000 feet; in the Taurus between 3,300 and 6,600 feet. 922535 (Fig. 1).-- An extensive forest occurs in the Troodoe Mountains. It is generally found between 4,300 and 5,300 feet above sea level. 21. Fig. 5.-- Natural distribution of European black pine in Yugoslavia (From world Forest Atlas by Fritz Heller Publishing Co., Berlin. 1951; Vidakovic, 1957; Eoehrig, 1957. Scale, 1 : 6,000,000) 2' \ 23 Fig. 6.-- Natural distribution of European black pine in Greece (From World Forest Atlas by Fritz Heller Publishing Co., Berlin. 1951. Scale, 1 : 3,200,000) r40 59 \t 22° 21° ,. o 45.1. 50 35/ 25 Fig. 7.-- Natural distribution of European black pine in Turkey (From‘World Forest Atlas by Fritz Heller Publishing Co., Berlin. 1951. The map agrees well with one prepared in 1962 by the Turkish Forest Service. A few small stands in the northeastern part of the country were not shown in the 1962 map, due either to destruction or overlapping with other species. Scale, I : 10,000,000) 27 Algeria (313. 1).-- Black pine is found on Mount Edough, Dept. Constantine, northeastern Algeria (Salvador, 1927; Sexton, 1947). The mountain rises to 3,307 feet, receives heavy precipitation up to 70 inches annually. It is covered with cork oak (Quercus suber L.) forest. Planted Distribution European black pine has been extensively planted outside its natural range. Promising results were reported in New Zealand (Hinds 35 21., 1957; Weston, 1957); in Great Britain (Stevens, 1934; Nacdonald 25 31,, 1957); and in the United States of America (Aughanbaugh, l96l; and E. J. Eliason, personal correspondence. September 28, 1964). New Zealand.--.According to Hinds gtflgl. (1957) Corsican pine is an important species below 38° south latitude. It was ranked third in the acreage among the planted exotic species. Actual acreages planted to this variety were as follows. North South Total Island Island -------- Act“ a--.---- State Forest, pure stands 41,000 17,700 58,700 State Forest, mixed stands 1,600 2,000 3,600 Private and Others -- -- l,000 It grows satisfactorily and forms productive stands with 40 to 60 inches of precipitation. It grows on a wide range of soils including those rich in bases. However, poor drainage and/or a shallow soil pan limits its growth. The altitudinal limit for successful planting is 3,000 feet in the central North Island and 2,000 feet in the South Island. 28 Corsican pine is sensitive to spring frost. Although it is rela- tively free from diseases and insect pests the Tortrix moth may attack buds in early years and may cause forking. Exceptionally straight trunks are characteristic of Corsican pine. It has earned a great reputation in New Zealand. Estimated rate of current planting is 1,500 to 2,000 acres per year (Weston, 1957). Austrian pine was probably introduced in 1866, approximately the same time that Corsican pine was introduced. It has been a coarse tree of little value. great Britain." Corsican pine is generally considered to have been introduced into Great Britain in 1759 (Stevens, 1934). Until 1840 it was planted primarily as an ornamental or in windbreaks. Extensive planting , started about 1870. h The 1947 statistics showed that total area planted to this species was 38,503 acres, of which 83 percent or 32,004 acres were operated by the Forestry Cousission (Nacdonald _e_t_ .a_1_., 1957). Corsican pine has been superior to other races of black pine in growth rate, small branch size habit and straightness of stem. It is therefore highly recouended for planting in the warmer parts of Great Britain. Adequate soil drainage is one of the most inortant site factors to consider in planting Corsican pine. In general frost and winter kill have not been a problem. The fungus Brunchorstia d__e_s_t_r_u_e_ns_ Erikss. has caused occasional dieback in areas of heavy precipitation. A pine sawfly Mbyrus app.) occasionally causes partial defoliation. Black pine varieties other than Corsican have also been planted to a large extent in Great Britain. Calabrian trees have replaced much 29 Scotch pine in the warmer parts of England, and locally in Scotland and wales. Austrian trees are planted pricipally in coastal areas and on calcareous soils. Spanish sources have been inferior to Calabrian and Corsican ones in survival and height growth based on two small provenance. studies started in 1951 (Nacdonald, $5 9_1_., 1957). l The United States of America.-- In the eastern United States only the Austrian variety has been planted to any extent. It has been used commonly as an ornamental, in windbreaks, and recently as a Christmas tree. There are a few forest plantings. The oldest one in the State of New York is about 30 years old. It grows throughout that State, but planting has been done mostly in the southern half. The State of New York has 75,000 acres ofiAustrisn pine plantations with a current planting rate of 500 acres per year (E. J. Eliason, personal correspondence. September 28, 1964). At Kellogg Forest, Augusta, Michigan there is a 30-yesr old plan- tation. It has grown at the same rate as red pine but slightly larger branches. It is now being thinned for pulpwood. Near the edge are several vigorous gig£g_x densiflora hybrids. These are seeded naturally from.the old stand. In Michigan black pine is a minor species (Stone and Chase, 1962). Its susceptibility to European pine shoot moth was moderate, based on examination. This investigation of six lower Michigan plantations ranging from 8 to 12 years of age (Miller and Heikkenen, 1959). In western Ohio, Austrian pine has been used as a windbreak on calcareous soils. Fast growth was reported fromla 3-acre experimental plantation established in 1938 in.wayne County, Ohio. After 23 growing seasons the trees were 35 feet tall (Aughanbaugh, 1961). 3O Trees of an unknown nonaAustrian origin grew better than Austrian ones in the Secrest Arboretum, Wooster, Ohio. The growth performance of both types is shown below Wat 11., 1958). NonrAustrian Austrian Age dbh height dbh height (inches) (feet) (inches) (feet) 10 3.6 19.8 3.8 17.4 25 7.0 44.6 7.1 38.8 30 7.5 62.0 7.0 47.7 40 9.0 65.6 8.8 56.0 Argentina.-- Eighteen-year-old Corsican trees reached 26 feet in height and 8 inches in diameter. That is considered slow growing in Argentina (Barrett, 1952). It is generally planted as an ornamental on calcareous soils in the milder parts of the Southern Andes. Damage caused .by aphid and breakage of branches due to snow have been observed. South Africa.-- Austrian pine is of minor importance in South Africa. The oldest trees are about 55 years old. In most cases they are crooked and unhealthy (Poynton, 1957). General Description of Eurgpggn Black Pine European black pine belongs to the group Lariciones, subsection Pinaster, section Diplggylon of the genus giggg, Large ray cell pits, cones dehiscent at maturity and thin seed wings are characteristic of Lariciones. Its articulated seed wings and persistent fascicle sheaths are characteristic of Pinaster. The decurrent fcscicle bracts and two fibrovascular bundles in the leaves are characteristic of‘Diplogzlon 31 (Shaw, 19110. Pilger (1926) subdivided the genus differently. According to his classification, EurOpean black pine is in the section Eupitys of the subgenus Diploglon . Dallimre and Jackson's (1961) description is included at this point to describe the range of variation in the species as a whole. Their description is as follows: "It is a large tree up to 120 - 150 feet high. Crown is pyramidal when young, flat-headed at maturity. Rough bark is grayish-brown or dark brown. It has dense branch system. Buds are ovoid- oblong or cylindrical in shape and 1/2 to 1 inch long; they are abruptly contracted to a sharp point with resinous and light brown scales. Leaves are in pairs, persisting about h years, very dense on the branchlets, stiff, stout, straight or curved, 4 to 6 inches long, margin minutely toothed, apex a thickened horny point, 12 to 1h lines of stomata on each surface, resin canals medial, basal sheath about 1/2 inch long at first, gradually wearing away as leaves age. The flowers fora about the end of m. The female catkins are in two's or three's or more, together, with rather long psduncles from the extremity of the young branches; they become 1/2 inch long and l/h inch wide and elliptical in July. The cones are tam- yellow, becoming brown as the seeds ripen, solitary or in clusters, sub-sessile, ovoid-conic, 2 to. 3 inches long and l to 1 1/4 inches wide before opening. They do not arrive at mturity until October in its second year. The scale is about 1 inch long transversely heeled near the apex which often ends in a more or less persistent prickle. Seeds are greyish or brownish, more or less mottled, about 1/8 inch long, with a wing several times its length . " CHAPTER 3 MATERIAL.AND METHODS Seed_procuremegt and nursery practice.-- In the summer of 1958 Dr. J. H. Wright requested black pine seed from several research foresters in southern Europe. Each was asked to send one ounce of seed from a native stand in his vicinity. They were very cooperative and included information on locality and altitude of origin, collection data and collector, soil material and age. Each collection was to include seed from 10 average trees situated within a radius of one mile. Upon re- ceipt, a ZO-seed sample of each seedlot was weighed. Then the seeds were stored dry at 35°F. until sowing. On May 13, 1959 the seeds were sown in a 4-replicated, randomized complete block design. Bach plot consisted of one b-foot row. The rows were 6 inches apart and were thinned to a density of about 40 seedlings per row (equivalent to 20 seedlings per square foot). These were to pro- vide material for the study of 2-year seedling characters reported elsewhere (Wright and Bull, 1962). A fifth set of plots was broadcast sown to provide stock for perma- nent outplanting. The study was conducted in Michigan State University's Bogus Nursery. The seedlings received conventional treatments in the nursery such as fumigation of seedheds with methyl bromide, application of water, fertilizer and mineral spirits weed killer as needed, occasional hand weeding, and a half-inch water mulch of sawdust. 32 33 Description of study plantation.-- Plantation No. HSPGP-S-él is one of 14 permanent black pine outplantings in the North Central Region of the United States. It is located in Compartment 26-D, Kellogg Forest, one mile north of Augusta, Kalamazoo County, Michigan. It was planted on an.8 x 8 foot spacing on March 23, 1961 by T. Godde, N. Barry, W. Lemmien, P.8tsncel, 0. 0. Wells and J. W. Wright. The field had not been used since 1948 and had a heavy sod of blue-grass. This was furrowed before planting. The soil is Oshtemo loamy sand to sandy loam. The site is rolling with eastern and southern aspects. The slopes are from 0 to 25 percent. woody vegetation was killed'with 2,4,5-T during 1958-59. On May 4, l9t4‘Hr. Lemmien applied a directed spray of simaaine and amino- triaaole to 2-foot strips containing the trees. The spray was at the rate cf uh pounds simszine plus two gallons of amino-triazole per acre treated. The plantation follows a randomized complete block design with 10 replicates (blocks). Each replicate consists of 27 four-tree plots. The northernmost two trees in each plot were fertilized with 45- 0-0 urea pellets at the rate of two ounces per tree. The fertilizer was applied twice; the first on April 10, and the second on.April 28, 1964. Each time one ounce of fertilizer was uniformly placed on the surface soil of a 16-inch diameter circle around the tree. The southernmost two trees- in the plot remained unfertilized. The plantation is now considered as following a split plot design with fertilizer treatment as the main factor, and provenance as the subfactor. Growth of the trees is satisfactory in.most cases. Those from Corsica and southern France suffered heavy winter hill but the remainder are healthy. Differences in height growth among the provenances are 34 noticeable to the naked eye and response to nitrogen fertilizer is recog- nizable in many provenances. The fertilized trees generally have much greener foliage than the untreated ones. Figlggmeasurements.-- Total height was measured in late 1963, 1964 and 1965 to the nearest half-inch. Branch angle was measured in 1964 to the nearest 5 degrees with the aid of specially made protractor. Mortali- ty was counted annually. The numbers of winter-damaged trees were counted in.April 1964. If 50 percent or more of the total foliage-was brown the tree was considered as damaged. Anatomical study.-- Sampling for the study of anatomical characters of the needles was conducted on September 15, 1964, five months after the first application of nitrogen fertilizer. In each plot one fascicle was collected from each living tree. The fascicles were taken from the south side of the leader and placed in FAA solution (a mixture of 1 part of 401 Formaline, 1 part of glacial Acetic acid and 18 parts of 502,Alcohol) on the same day. A preliminary study showed no significant difference be- tween the two needles of the same fascicle for several characters studied. Therefore the single needle per fascicle seemed to be sufficient to pro- vide the necessary data (see Appendix A). The following foliage characters were studied in the laboratory: needle length to the nearest millimeter, needle width to the nearest 40 microns, length/width ratio, number of serrations per 2 millimeters, number of resin canals, position of resin canals (external, medial, in- ternal, medial-external, or medial-internal), number of layers of hypo- dermal cells on the ventral and dorsal sides and on the corners, number of sclerenchyma layers over the phloem, distance between the two fibrovascu- lar bundles to the nearest 40 microns, length of endoderm to the nearest 35 40 microns, the width of endoderm to the nearest 40 microns, number of rows of stomata on the ventral and dorsal surfaces, ratio between endo- dermal length and endodermal width, and ratio between rows of stomata on the dorsal and ventral surfaces. All the anatomical characters were ob- tained in transverse section across the middle location of each leaf furnished by free hand sections and stained by Safranin 0 and Past Green PCP. Paraffin sections cut at 20 microns thick and stained by the Safranin O - Fast Green FCT staining schedule (Johansen, 1940), were used to obtain photomicrographs. Chemical study.-- I collected 70 foliage samples in mid-November 1964 following Dr. D. P. White's recommendation (1954). Each sample con- tained needles from trees of single seedlot and fertilizer treatment. In most cases a sample was from all 10 replicates; for 8 seedlots there were separate samples for the first 5 or second 5 replicates. The foliage samples were quickly brought back from the field and oven-dried immediately at 149°F. (65°C.) for 48 hours. When dried they were ground in a Wiley mill and made to pass a 20—mesh screen. The samples were analysed spectrographically for phosphorus, calcium, magnesium, iron, manganese, copper, sine, aluminum, boron and sodium. Nitrogen was determined by the micro-Kjeldhal method and potassi- um by the flame photometry. All analyses were undertaken at the Plant Nutrition Laboratory, Department of Horticulture. The instrumental errors were calculated by Dr. A. L. Kenworthy (unpublished data) from data on 18 samples analyzed 5 times. These are shown below. 36 Element Mean concsn- Coefficient of tration variability Phosphorus 0.328 I 2.60 \ Calcium 1.09 " 5.31 Magnesium - 0.339 " ' 4.87 Sodiua ' 349.0 ppm. 21.20 Manganese 156.0 " 5.30 Iron 93.0 " 8.43 Copper 16.3 " 14.29 Boron 29.2 " 7.37 Zinc 39.2 " 40.00 Aluminum 107.0 " 8.63 In addition ground tissue was scored for color. This was done independently by Dr. Wright and myself. In doing this bottles containing the samples were sorted into 17 color grades. The "eye" or instrumental error was 1 standard deviation 3 3.5 color grades. The Munsel equivalents of the extremes were as follows. Color grads Mnnsel equivalent 1 5.0Y6/6 l7 5.0Y6/8 Statistical treatment.-- The Kellogg Forest plantation contained a total of 540 subplots. Of these, 36 (s 6.7 percent) were missing in that both trees had died. Substitute values were calculated by computing the average for the surviving plots of the same provenance and treatment. The degrees of freedom for the error term were reduced accordingly. 37 After substituting the missing plot values each set of field or ' anatomical measurements was subjected to analysis of variance for which the degrees of freedom were as follows. Source Degrees of freedom Blocks 9 Fertilizer 1 Error A (Block x Fertilizer) 9 Provenance 26 Fertilizer x Provenance 26 Error B (Block x Provenance plus Fertilizer x Provenance x Block) 432 Total 503 For the chemical study the degrees of freedom were as follows. Source 1 d. f. Fertilizer l Provenance 26 Error (Fert. x Prov.) 26 Total 53 The above analysis could show the significance of the main effects (fertilizer and provenance). It could not show significance of the Fert. x Prov. interaction. To do that, data from the four bulked samples (fertilized or unfertilized from replicates l to 5 or 6 to 10 respectively) from each of the selected provenances were subjected to special analyses of variance. For these the degrees of freedom were as follows. 38 Source d. f. Block 1 Fertilizer 1 Error A (B x F) '1 Provenance 7 Fertilizer x Provenance 7 Error B (B x P plus F x B x P) 14 Total 31 On the assumption that these 8 provenances were a random sample of the entire 27, the error terms derived from the above analyses can be used to test the significance of the fertilizer x provenance interactions for the entire study. To do this the F x P mean squares based on 27 prove- nances were divided by the F x P x B mean squares based on 8 provenances. CHAPTER 4 GEOGRAPHIC VARIATION PATTERN Growth, anatomical and chemical data for the Kellogg plantation are presented in Tables 1 through 8. Included in those tables are means for each seedlot, arranged roughly in west to east order. These means apply to fertilized and unfertilized trees. The effects of fertilization are treated separately in a later chapter. An analysis of variance was performed for each trait. That showed whether the between-origin differences were significant, but not whether trees of one country were sufficiently different from those from another country to be classed as a separate race. Ordinarily, this would be learned from another analysis of variance, using seedlot means as items. However, that was unwise because some countries were represented by one or a few origins. There was an adequate number (13) of seedlots from Greece to sample the variability in that country. Therefore, the Creek popu- lation was used as-a standard against which all others could be compared. This was done character by character. If one or all the seedlots from another country fell outside the 95 percent confidence limits for the Greek population it could be considered that the Greek and other popu- lation differed enough to be treated separately. Another technique was used occasionally. The variability of the Greek population could be considered as the standard. Were the trees from two other countries more different from each other than two different Greek seedlots were from.each other? A standard deviation (applicable to a provenance mean) was calculated from the Greek population, and compared 39 40 with the difference between the non-Greek seedlots. Both procedures left something to be desired. The gross aspects -- but not the fine details -- of the variation pattern could be detected. More intensive range sampling would have been necessary to obtain defini- tive answer on genetic discontinuities and other important theoretical questions. The procedures are similar to Critchfield's graphical method (1957). He used the range of observations to find the likelihood of two samples being from the same population. If the range overlapped, the two samples were from the same population; if they did not overlap, there was little likelihood of the two samples being drawn from the same population. Most statistical analyses were carried out with an electronic computer, CDC 3600. Growth Traits The relationship between geographic origin and genotype has been demonstrated in many forest tree species. Each geographic origin has a particular habitat and has adopted itself to local growth conditions for many generations. Take Scotch pine for example. Trees from most cold Scandinavia are much slower growing and have much shallower roots than do trees from Germany with milder climate. The Austrian trees generally grow on calcareous soils. In their native habitats, they are characterized by a rather slow growth and less desirable tree form. On the other hand, Corsican trees dwell on soils of igneous origin (granite). They are rapidly growing and have beautiful tree form. From an unreplicated geographic origin test conducted in Belgium, it is inter- esting to learn that the Corsican trees are rather indifferent to the 41 mineral content of soil, but, are sensitive to soil drainage (Delevoy, 1949 a). Roehrig (1957) stated in his paper that a large provenance test was set up in Spring, 1957 in West Germany. Twenty-five seedlots were included. He gave no details of seed collection or experimental design. Growth rate::? The Kellogg plantation has the highest survival of any of the NC-Sl g, gigs; tests established in 1961. Also, its growth is among the best. The 6-year heights are low in comparison with 6-year heights of other species which get off to a more rapid start. However, they are above average in comparison with most other black pine plan- tation in the state. In 1961, the first year after plantihg, the trees grew little. Host trees were firmly established by 1962 and were capable of normal growth. By the end of 1964, most evidence of transplanting shock had been overcome. The 1964 total heights for each provenance are given in Table 1. First, let us turn our attention to the Greek population, which contained the tallest (HSFG 425 -- 80 cm.) and one of the shortest (MSFG 416 -- 49 cm.) seedlots. There was possibly a slight pattern within the country. The three seedlots from the Peloponnesus Peninsula (416, 421, 429) were all below average, and the three from the eastern part of the mainland (422, 424, 425) were average or above average. Those from the Greek islands (419, 426, 427) and the main portion of the country were above average. Three seedlots which were classified as Corsican were the slowest growing. One (412) was grown from seed collected in Corsica; another (411) was grown from seed collected from Corsican trees in the.Arboretum 42 des Barres, Nancy; the third (413) from a supposedly native stand of another type in the French Pyrenees. It was difficult to see any pattern in the rest of the range. The single Austrian seedlot (423) was considerably below average, and the single Crimean seedlot (408) was considerably above average. Growth rate of the Spanish, French, and Turkish trees fell within the range of the Greek material. The poor performance of the Austrian trees indicates one practical reason for a study such as this -- Austrian pine is now the standard variety being planted in this country and far from the best. There was a general correlation between the 6-year heights and the 2- and 3-year data reported by Wright and Bull (1962) for the same material in the nursery. They, too, found that the Corsican material was among the shortest and that Austrian pine grew more slowly than.most others. However, there were exceptions. They reported that seedlot 416 was one of the ‘ tallest. Perhaps this discrepancy is due to damage suffered in lifting. There is a record that, due to partially frozen soil, the lifter blade accidentally rose and cut the roots of that seedlot abnormally short. Maybe the trees have not yet fully recovered. A strong correlation between the 1964 heights (the 6-year heights) and the 1963- and l965-data was also established for the same material in the same study plantation. The trends found in the 1964 heights were generally applicable to the 1963 and 1965 heights. ' The juvenile mean height growth (at ages 10 and 15) observed on several sites in Belgium was in the following order: Corsican trees were the fastest growing among several origins tested; Calabrian trees were next fastest growing; the trees from Southwest France were of moderate growth rate. Crimean trees were unsatisfactory in their juvenile height 43 growth, however, their growth potential increased with age. At age 36 they were as tall as Corsican trees (Delevoy, 1949 b). The same author (Delevoy, 1950) later made a comparison of volume growth between Corsican and Austrian trees. he found at age 15 that Corsican trees had 50 percent greater volume than.Austrian ones. There seems to be no doubt that Corsican trees grow quite satis- faclra'ily under Belgian climatic conditions. Gathy (1957) also confirmed this point. When the growth of Corsican trees is compared with Scotch pine, the former exhibited faster growth and better stem quality according to unreplicated test plantations in North Belgium (Miegroet and Janssens, 1956). In Great Britain, Robinson (1945) reported that Corsican trees were fast timber producers. The yield was greater than for Scotch pine. For example, the mean annual increment of Corsican trees was 125 cubic feet per acre for the first 25 years whereas the corresponding figure for Scotch pine was 78 cubic feet per acre. What is the reason for the contrast between European and Michigan performance data? The primary reason is possibly that Corsican trees have been planted in warmer parts of European countries where oceanic climate is more or less in effect. In'Hichigan the climate is continental, a temperature of 20°F. below zero being common in winter. A sudden drop of temperature results in crystalization of cell sap which may be fatal to non-resistant Corsican trees. It is interesting to compare the pattern in this species with that found in Scotch pine by Wright and Bull (1963). The two species have. roughly parallel ranges in the Mediterranean area, the Scotch pine being found at higher elevations. The two patterns agree in that there are not 44 Table 1. Growth and needle characteristics of Pinus nigra from different.origips. 'COUNTR?% HEIGHT eRANCH nlNTtk Neauue NtEDLE ORIGIN 1964 ANGLE JURN LENGTH elDTH NUMBER 1963-64 (11 (21 (31 (4) (6) NM DEGREE PERCCNT Mm MM CORS1CA 411 471 13700 0705 105 10.58 412 443 4000 72.: 100 1.32 413 460 64.3 67.6 103 1.36 AVERAGE “DU “.340 7 Che“ 103 1 0.15 SPAIN 402 606 61.6 7.6 123 1.37 403 640 61.4 7.5 121 1.30 AVERAGE 62: 61.6 7.6 122 1.34 FRANCE 407 004 59.a a7.s 109 1.3: 410 629 60.6 10.0 100 1.37 414 993 51.9 45.0 117 1.41 AVERAGE 6339 57e6 6402. 109 1 031 AUSTRIA 423 969 04.: o be 1.39 YUGO. 416 066 63.9 0 d? 1.26 GREECE 416 469 63.2 o 77 1.30 417 617 60.6 2.5 68 1.26 418 606 63.1 o 68 1.26 419 636 69.3 2.5 92 1.29 420 679 66.3 2.6 74 1.26 421 507 64.7 0 75 1.25 424 644 61.6 0 92 1.34 425 601 no.7 0 66 1.33 426 666 66.2 2.6 90 1.31 427 06‘ 65.6 2.5 101 1.43 428 646 66.3 0 90 1.31 429 677 66.6 0 60 1.31 AVERAGE 61” (11.7 100 C13 1.31 TURKEY 401 606 69.4 0 76 1.32 404 662 64.4 0 95 1.37 405 012 6304 2.6 91 10‘0 AVERAGE 627 63.1 .6 67 1.36 cutmca 406 701 67.2 0 102 1.40 45 many differences between the Greek, French and Turkish populations. They disagree in other respects. In Scotch pine, seedlings from Yugoslavia and farther north clearly belong to a faster growing central European variety. Also, Spanish Scotch.pine is measurably slower growing than other south European types. Height growth begins with bud burst and terminates with bud set. Morris 55.51, (1957) observed strong genetic control over the time of bud burst in Douglas-fir based on a plantation established in 1915. In general, bud burst occured earlier for southern seed sources at a given testing sites. They continued to grow late in autumn and hence were sub- ject to early frosfi'injury. Approximately the same trend has been found for low and high elevational races of ponderosa pine Cuirov 55 21., 1952; Irgens-Noller, 1957; Callaham ggflgl., 1961). A few observation was made to determine whether the same factors were operative in European black pine. Seemingly, they were not. Time of bud burst varied by only one or two days among origins and dormant winter buds were visible on all origins by mierune. Admittedly, however, the data were not conclusive because weekly measurements were not made to de- termine the exact time of growth start and cessation. Branch angle.4- This characteristic is important from an esthetic point of view and also important to the forester. It has been studied little. Differences in this trait among the 27 seedlots were noticeable to the naked eye and were statistically significant (Table 7). The branch angle for each provenance is given in Table l. The pattern in the Greek population seems to be a random one. Of the 13 Greek seedlots, the following had acute branch angles: HSFG 421, one of three Fe10ponnesian seedlots; 419 and 426, two of the three seedlots from Greek 46 islands; 422 and 425, two of the three seedlots from the eastern mainland and 417, one of the four seedlots from the rest of the Greek mainland. The Corsican, Spanish and French provenances all from the western range were characterized by much more acute branches than the Greek trees, Corsican trees had the acutest branch angle of all. The Austrian, Yugo- slavian, Turkish and Crimean provenances were characteristically flat branched. ,. Winter burn.-- Most species with a wide natural range show great variation in winter hardiness. European black pine is no exception. In general, western provenances were susceptible to winter burn in Michigan (Table l). Corsican trees were most heavily damaged. Some of their terminal buds were killed. French and Spanish trees suffered less damage. Greek and Turkish trees were only slightly injured. Trees from Austria, Yugoslavia, and the Crimea were not injured. In November 1961, 6 months after planting, overall mortality was 22 percent. Mortality was highest (75 percent) in seedlot 416 from Greece. That seedlot suffered noticeable root damage when lifted. Three seedlots from Corsica (411, 412, 413) suffered next highest mortality (40 percent). Those three Corsican seedlots had brown tops in the nursery. This hidden injury combined with planting shock may explain the high mortality. Dead trees were replaced in the spring of 1962. Overall mortality was about 10 percent in the September 1962, 11 percent in November 1963. Cathy (1957) believed that Corsican trees derived from different altitudes differed in resistance to extreme climatic conditions. He ob- served at Louveigne, Belgium (about 1,000 feet altitude) that trees from high altitudes generally grew better and suffered no winter damage. In Great Britain, Robinson (1945) recommended Corsican pine of high alti- 47 tudinal origin for planting purposes. In my experiment, seedlot 412 was collected from a natural stand at 3,000 feet elevation in Corsica. It suffered severe winter injury. Evidently, if there are altitudinal races on the island, none is sufficiently hardy for Michigan purposes. There was a strong correlation between the 2- and 3-year data (Wright and Bull, 1962) and the 6-year data on winter injury. 11:... authors were able to recognize the Corsican and Non-Corsican ecotypes by this particular character. For the Greek population it is interesting to note that all three island-inhabiting seedlots (419, 426, 427) suffered slight winter injury. It reflected a long term influence of the mild oceanic climate. I The results suggested that the Corsican and southwestern French seed sources should not be used by Michigan tree growers. General appearance.-- Some seedlots show distinct characteristics to the naked eye. They are easily identifiable to any one who visits the study plantation for the first time. Curved leaves are characteristic of all three seedlots from Corsica. Seedlot 414 from France is characteristically bushy. Seedlot 423 from.Austria has moderate growth rate and white, resinous winter buds. The leaves grow at right angles to the leading shoot so that the terminal buds are fully exposed and visible from a distance. In contrast to the Austrian trees, Corsican trees and Crimean trees bear leaves forming a small angle with the leading shoot. The leaves hide the terminal buds, which are not easily seen from a distance. By the differences in leaf curvature, one is able to distinguish between I I 48 Corsican and Crimean trees. Young branchlets in the first whorl are slender and characteristi- cally orange-colored in Spanish trees, which also have long needles. Anatomical Characteristics Leaf anatomy has been studied by several students of geographic variation in pine species. The works of Coulter and Rose (1886), and of Doi and Morikawa (1929) on internal leaf structure are of special im- portance. Harlow (1931) was the first worker to present a series of photomicrographs dealing with the native and introduced pine species in the United States. Recently, Vidakovic (1957) fomd the following ana- tomical traits important in the identification of European black pine from different parts of Yugoslavia: number of stomata, number of resin canals, structure of epidermis, structure of hypodermis, breadth and heightof transverse section of the needle, nunber of layers of trans- fusion parenchyma, number of sclerenchyma in the vascular bundles, and endoderm structure. Leaf anatomy is an useful tool for verifying pine hybridity when the specimen is young and bears no cones. Mergen (1958) and Vidakovic (1958) so used needle characters. Keng and Little‘(l962) at the Institute of Forest Genetics, Placerville, California found that F1 interspecific hybrids had needle characters intermediate between those of the parent species. Forty-two pine hybrids were involved in their study. Concerning needle characteristics, Kriebel 35.21. (1965) warned that there was more variability in needle anatomy within taxa and within a single F1 hybrid family than was indicated by earlier investigators. 1 also found a considerable variation in needle anatomy within the Greek 0 I 0 population. 49 Some abnornl and rarely occurring features were found in leaf cross sections. Three fibrovascular bundles were noticed in single leaves from seedlots mm hos from Spain; MSFG 1.20, 11211 and 1129 from Greece (Fig. 8.1),. One needle from a tree of MSN 418 (Greece) was completely devoid of resin canals (mg. 8-2). Analyses of variance indicated that there were highly significant differences along the 27 seedlots for most anatomical traits. Only two characters (the ratio of number of rows of dorsal and ventral stomata, and the distance between two fibrovascular bundles) were constant. Needle leng;t_h_.- - Needle length is one of the most dependable in- tensively studied characters used to differentiate geographic varieties. There are several examples. In Scotch pine, Wright and Bull (1963) found that needles were longest in trees from central Europe and shortest in trees to the north (Finland) and south (France, Spain). Morgen (1963) found in eastern white pine that needles were shortest in trees from northern latitudes. A geographic pattern in needle length was also evi- dent in lodgepole pine. Critchfield (1957 ) stated that needles of the inland populations were longer than those of the coastal ones. Needle length showd a decreasing trend from west to east. Spanish trees had the longest needles of all. Turkish and Greek trees had the shortest needles. The one Crimean seedlot formed an exception to the general trend -- it had moderately long needles. There was a slight pattern within the Greek population. Trees from island-inhabiting seedlots (419, 426, 427) had above-average needle length; those from the Peloponnesus Peninsula (416, 421, 429) had below-average needle length. Of the remining seven seedlots from the Greek mainland, five had above-average needle length . Figs 8a-- (1) (2) (3) (4) 50 Photomicrographs showing abnormal features of needle anatomy Three fibrovascular bundles found in seedlot MSFG 424 from Greece; Absence of resin canals in one tree of seedlot 418 from Greece; High number of resin canals (3 on the ventral, S on the dorsal portion of the needle) in a leaf from a fertilized tree of Greek seedlot MSFG 422; Unfertilized tree of the same seedlot showing only 4 resin canals. 52 Needle curvature.-- Needles in the 6-year Corsican trees were curly. Wright and Bull (1962) also reported curved needles in the 2- and 3-year seedlings for all three seedlots from Corsica. Other seedlots generally had straight needles. Needle width.-- Data on needle width are presented in Table 1. Analyses of variance showed that differences among 27 seedlots are sig- nificant at the 1 percent level. Crimean trees had the widest needles, Austrian trees had the next widest. Seedlots from Spain, France, Corsica, Yugoslavia and Turkey fell within the range of the Creek population. The range in this character was relatively small. The extreme difference between seedlot means was 0.24 mm, approximately 18 percent of the mean. The range of seedlot means from Greece was 0.20 mm, about as great as for the species as a whole. Needle lengthlgidth ratio.-- This ratio is an indication of needle slenderness. The larger the ratio, the more slender and flexible the needles. The ratio for all 27 seedlots are presented in Table 2. Differ- ences in this character was highly significant among seedlots. The trend in this character is similar to that for needle length. The ratio showed a decreasing trend from west to east. Spanish trees had the largest ratio of all which confirmed my optical observation in the study plantation. Turkish trees had the smallest ratio. The single Crimean seedlot formed an exception to the general trend -- it had moder- ately large ratio but was smaller than any of the three western Popu- lations (Spain, France and Corsica). There was a slight pattern among the Greek seedlots. Trees from island-inhabiting seedlots (419, 426, 427) had above-average ratio; those Thhh32. origins. COUNTRY. UHIGIN NUMBCH CORSICA SPAIN FRANCE AUSTRIA YUGO. GREECE TURKEY CRIMEA 53 Needle characteristics of Pinus nigra from different 4l1 412 413 AVERAGE 402 403 AVdHAGE 407 410 414 AVERAGE 423 415 416 417 41a 419 420 421 422 424 425 426 427 426 429 AVLRAGE 401 .0. son AVERAGE 408 NttULE LENGTH! uIUTH RATIO (6) NUMUER 76.1 7b.8 77.6 76.6 59.8 93.1 91.9 50.7 73.0 96.7 83.5 63.3 65.0 39.2 69.8 09.8 71.3 57.8 00.0 59.5 Obo7 03.7 65.7 09.7 05.7 01.1 95.2 57.6 69.3 65.0 64.0 72.9 NtuuLt COLUH (71 GRAUdS 12.0 13.0 12.0 12.3 10.0 7.0 8.3 3.0 12.0 10.5 10ed HcIoHt/ NEfiULE LtNGTH RATIO (U) NUMUER 4.49 4.43 4.3u 4.43 4.95 5.29 9.1:: 6.09 6.29 5.10 3.83 6.70 7.05 5.35 7.01 6.91 5.93 7.55 6.76 8.59 7.00 9.36 0.31 0.33 7.15 7.21 7eZJ 6.00 0.95 5.73 7.23 6.87 HtSlN CANAL (9) NUMUER 4.. 4.3 4.7 4.4 54 from the Peloponnesus Peninsula (416, 421, 429) had below-average ratios. Needle color.-- Differences in needle color among various seed sources have been reported for tree species. Take Scotch pine for example. Autumnal and winter color differences have been noticed for years (Wright, 1963; Wright and Bull, 1963). Northern provenances usually turn yellow in winter, central European ones make less discoloration, but southern Euro- pean ones remain green round the year. The time of yellowing varies from year to year, however, this phenomenon tends to intensify from autumn to late winter. The recovery of green color of northern provenances parallels approximately with the beginning of the growing season. Photoperiod and temperature played important roles in the seasonal discoloration of sugar maple (Olmsted, 1951) and Scotch pine (Gerhold, 1959 b). The scoring of needle color in the present study was on the basis of ground samples. Dr. Wright and I did the scoring in late November 1964, shortly after the ground samples were prepared from fresh needles. Total number of color grades was 17, varying from yellowish brown (low color grades) to bluish green (high color grades). There were significant between-seedlot differences in the ground foliar color. Data are presentedin Talbes 2 and 6. The variation pattern seemed to be a random one. The foliar color of Corsican and Crimean seedlots was bluish green (high color grades), of Spanish and French seedlots was green (intermediate color grades) and of the remaining seedlots (Austria, Yugoslavia, Greece and Turkey) was yellowish brown (low color grades). It is interesting to compare the pattern found in European black pine with that found in Scotch pine re- ported by Wright and Bull (1963). They recognised four distinct color 55 classes for Scotch pine in the Mediterranean area. Italian Scotch pine. had low color grades (intermediate between yellow and green), the Austrian- Yugoslavian trees had medium color grades (green), the Creek-Turkish had higher color grades (between green and dark gree) and the Spanish Scotch pine had the highest color grades (blue-green). The most striking dis- crepancy in the color grades was that the Austrian-Yugoslavian seedlots differed from the Greek-Turkish ones in Scotch pine, whereas in European black pine those four seed origins had the same color grades (yellowish brown). It was difficult to see any pattern within the Creek seedlots. Of the 13 seedlots, one (416) had bluish green color, 6 seedlots (417, 419, 420, 421, 426, 429) were of green color, the remaining 6 seedlots (418, 422, 424, 425, 427, 428) had yellowish brown color. ggightjneedle length ratio.-- According to a regression analysis using seedlot means as items there was no relationship between height and needle length. Differences in height/needle length ratio were significant among the 27 seedlots, according to analysis of variance. Details are shown in Tables 2 and 6. There was a distinct pattern to the geographic variation. In general, seedlots from the western range (Spain, France, Corsica) were characterized by lower height/needle length ratios than the Greek material, the Corsican seedlots having the smallest ratio of all. On the other hand, seedlots from the eastern range (Austria, Yugoslavia, Greece, Turkey and Crimes) were much high ingthis ratio -- larger than six. The single Yugoslavian seedlot had the highest ratio. 56 Within the Greek population, the three island-inhabiting seedlots (419, 426, 427) and the three from the Peloponnesus Peninsula (416, 421, 429) had below-average ratios. Of the remaining seven seedlots four were below'average. Number of resin canals.-- Mean number of resin canals presented in Table 2 for the 27 seedlots is far less than previously reported. Vidakovic (1957) who reported a range from 7.4 to 11.1 in the mean resin canals of European black pine five races grown in Yugoslavia. In eastern whité‘pine, there were also geographic differences in this anatomical character. Merges (1963) observed more resin canals for the specimens from northern latitudes than the specimens from sorthern latitudes. Differences in number of resin canals among the 27 seedlots were significant at the 1 percent level (Tables 2 and 6). There was a geographic trend to the variation pattern in European black pine. Mean number of resin canals decreased from west to east, the Corsican trees having highest number of all (4.4 resin canals). However, both Spanish and French seedlots were exceptional to this general trend. ‘ Although they are in the west of Corsica, yet they had fewer number of resin canals than the Corsican seedlots. Schwarz (1936) found the same west - east trend. Some pattern was recognizable within the Greek population. The three seedlots (416, 421, 429) from the Peloponnesus Peninsula were low in number of resin canals. 0n the contrary, the three (422, 424, 425) from the eastern part of the Creek mainland were considerably high in number of resin canals. 57 One tree of Greek seedlot 418 was devoid of resin canals (Figure 8 - 2). Complete absence of resin canals has been reported in the coastal and Mendocino Plain populations of lodgepole pine (Engelmann, 1886; McMillan, 1956; Critchfield, 1957). Number of resin canals possibly increase with age. Mean resin canals in Austrian pine trees on the Michigan State University campus was 6.2 in five 20-yearrold trees; it was 10.3 in five 40-year-old trees. Position of resin canals.-- The position of resin canals is an important anatomical trait in pine identification. According to Doi and Morikawa (1929) and Harlow (1931), European black pine was described as medial (resin canals are completely surrounded by the green tissue, meso- phyll). The majority of European black pine seedlots had medial resin canals. However,!bther positions were found. For typical external, medial and internal positions scores of l, 2 and 3 were assigned respectively. There were several cases of resin canals that touched both hypoderm and endoderm. Data on positions of resin canals are presented in Table 3. There were significant between-seedlot differences in this anatomical character (Table 6). There was a west-east trend in resin canal positions. Resin canals were most nearly external in seedlots from Spain, France, and Corsica. They were more nearly medial in seedlots from Austria, Yugo- slavia, and Greece. Turkish and Crimean seedlots had more nearly external canals than did the Greek material. Serrations, number_per 2 mm.-- This character gives an indication of needle smoothness. It is considered important by many researchers. Eastern white pine originating in southern latitudes was less serrated 58 Table 3. Differences in surface needle characteristics associated with geographic origin. COUNTRYs RESIN StHRATIONS STOMATA STOMATA STOMATA 0R1 G 1 N CARI“. PEN 2N”. VtINTHAL DOHbAL DORfi/VEN. NUMBKR RATIO (10) (11) (12) (Id) (14) P051 T 1UN NUMUEH R0135 R0 H 6 NUMBER CORS1CA 411 1069 606 706 1106 1051 “12 1090 902 703 1009 1049 413 1093 301 705 1102 1049 QVERAGE 1091 507 705 1103 1050 SPA1N 402 1061 607 703 1105 1060 403 1006 700 703 1300 1064 AVERAGE 1064 609 703 1106 1062 FRANCfi 407 1094 601 706 1109 1057 410 1994 U02 700 1201 1056 414 1004 004 700 1100 1037 AVhRAGE 1091 706 705 1107 1036 AUSTRlA 423 1096 701 707 1200 1056 YWCe 415 1099 701 70‘ 1102 1051 GRbECE 416 2000 500 705 1102 1049 417 2004 703 704 1103 1053 416 2001 706 701 1009 1054 419 1095 607 006 1002 105“ ‘20 1096 701 70‘ 1100 1049 421 1099 706 609 1004 1051 422 1096 704 702 1105 1060 424 1099 705 706 1201 1055 425 1099 701 704 1109 1061 426 1095 709 709 1200 1052 427 1099 600 607 1302 1052 428 1096 706 704 1104 1054 429 1090 707 606 1008 1059 AVERAGE 1099 705 704 1104 10:4 TURKEY 401 1090 707 705 1106 1055 404 1096 706 706 1105 1066 405 1099 802 709 1109 1051 AVERAGE 1096 708 707 1105 1054 CR1MEA 408 1096 705 800 1204 1055 59 than the northern ones (Hergen, 1963), Analysis of variance indicated that there were significant differ- ences in needle serrations among the 27 seedlots (Tables 3 and 6). There was little pattern in needle serrations. A Stomata, number of rows.-- This morphological characteristic is relatively independent of the environment according to Mergen's (1958) finding with slash pine from 12 different geographic locations. He found that seedlings of eastern origins had more stomata per unit length than did seedlings of‘western origins. ‘Mergen also demonstrated a geographic trend with eastern white pine (1963). More stomata were found in trees from southern latitudes than in trees from northern ones. Number of rows of stomata has been used to verify interspecific hybridity. Mergen (1959) examined the number of stomata in 4 to 6 year old hybrid seedlings of several combinations. It was intermediate between the two parents . In general, the number of rows of stomats increased from west to east in European black pine. Spanish seedlots had the fewest rows of dorsal and ventral stomata (11.8 and 7.3 respectively). The number of rows of dorsal and ventral stomata increased to 12.4 and 8.0 respectively in the Crimean seedlot. A slight pattern was recognizable in the Greek material. The four seedlots (417, 418, 420, 428) from the Greek mainland were average or below average in number of rows of dorsal and ventral stomata. A correlation analysis was undertaken (using seedlot mean as item) to determine the relationship between needle width and number of rows of stomata. The relationship was strong and positive (r : 0.746 for dorsal and r : 0.750 for ventral rows of stomata). 60 Stomata, dorsal/ventral ratio.-- This ratio was constant among the 27 seedlots (Tables 3 and 6). Hzpodermal layers.-- European black pine leaves have uniform, thick-walled hypodermal cells (Doi and Morikawa, 1929). The number of layers of the hypodermal cells usually varies from 2 to 3. I measured number of hypodermal layers at the three locations of a needle cross section -- on ventral and dorsal surfaces and at the corner having the greatest number of layers. There were more layers of hypodermal cells at the corners than on either surface. More layers of hypodermal cells were observed on the dorsal surface than on the ventral surface in all seedlots (Tables 4 and 6). The between-seedlot differences in this anatomdcal character at the three locations were significant at the 1 percent level. There was possibly a slight pattern to the geographic variation. The three western populations (Spain, France, Corsica) had a below-average umber of hypodermal layers. The rest of the seedlots fell within the range of the Greek material. The pattern was con» for the hypodermal cells observed in all three parts of the needles. Sclerenchy_m__aI nunber of layers." Sclerenchymatous cells could be present or absent in European black pine leaves. When present, they formed one or more irregular layers on the phloem (external) side of the fibro- vascular bundles. - There were significant between-seedlot differences in nuaber of sclerenchymatous layers (Tables 4 and 6). masher of layers of sclerenchyma was below average for the Corsican, French and Austrian seedlots, Corsican trees have the fewest. The rest of the range fell within the 95-percent confidence 1131': of the Greek material. 61 Number of sclerenchymatous layers was similar for all Greek seedlots. Distance between fibrovascular bundles.-- This character was constant for the entire range. No significant between-seedlot differ- ences were detected (Tables 5 and 6). gnggggnn, length.-- Endoderm is the boundary tissue between stelar tissue surrounding the fibrovascular bundles and the mesophyll (the green tissue). It is elliptical in cross section in European black pine. According to Doi and Morikawa (1929), the endoderm is composed of equal sized cells with uniformly thickened walls in this pine species. Differences in the length of endoderm were significant among the 27 seedlots as shown in Tables 5 and 6. The Austrian and Crimean seedlots had long endoderm cross sections. Within the Greek material there was a slight pattern recognizable. The three seedlots from the Peloponnesus Peninsula (416, 421, 429) were below average in endodermal length. There was no discernible pattern for the remainder of the range. Endoderml width.-- The width of endoderm differed significantly among the 27 seedlots (Tables 5 and 6). The single Crimean seedlot had much wide endoderm than did other seedlots. No pattern.was recognizable for the rest of the range. W." There were significant between- seedlot differences in endodermal length/width ratio, although the range between the two extremes was rather small (Table 5). The single.Austrian seedlot had a high ratio. The three island- inhabiting seedlots from Greece had a low ratio. No pattern‘was visible for the rest of the range. 62 Table 4. East-west variation patterns in number of layers of hypodermal and sclerenchymatous tissue in black pine needles. COUNTRY 0 HYPODEHM HYPUDERM HYPODEHI“ SCLEREN" OH1G1N VENTHAL CORNER DORSAL CHYNA NUMBER LAYERS LAYERS LAYERS LAYERS (1D) (16) (17) (18) ---- NUMUER ---- CORS1CA 411 100 106 10d 04 4 1 2 1 e 0 1 0 9 1 o 1 o 2') 413 100 106 101 07 AVERAGE 100 107 101 05 SPA1N 402 100 109 102 09 403 101 200 102 09 AVERAGE 101 200 102 09 FRANCE 407 100 109 105 07 410 101 105 103 0? 41¢ 100 106 101 09 AVERAGE 100 108 103 06 AUSTR1A 423 1003 8.01 300 06 YUGOe 415 102 202 108 09 GREECE 416 101 204 202 100 417 101 £01 109 100 ‘13 103 202 200 09 ‘19 103 201 108 09 ‘20 103 2.2 106 09 “21 103 801 109 09 422 103 204 200 100 ‘2‘ 104 403 200 100 425 105 202 201 100 426 102 201 109 100 427 106 £05 202 100 428 103 204 200 100 429 104 205 202 100 AVERAGE 103 203 200 100 TURKEV ‘01 102 203 202 100 404 102 203 200 100 ‘05 106 206 200 100 AVERAGE 103 30‘ 201 100 CRIMEA 408 104 204 202 100 63 Table 5. Differences in internal anatomy of black pine needles . COUNTRY 0 D [STANCE ENDODERa'u ENUODERM ENDODEHH ORIGIN bETwhEN LENGTH w l DTH LENGTH/ NUMBER F I UROVAS . w I DTH EUNDLES RATIO (19) (20) (an (22) t". I CRONS .51 l CRONS m l CHONs NUMUER CORSICA 4!) 86.6 776.0 39 1.96 412 5200 72500 372 1.95 413 71.2 738.8 383 1.91 AVERAGE 79.6 746.7 383 1.94 SPAIN 402 77.2 738.0 375 1.97 403 80.0 799.2 393 2.03 AVERAGE 78.6 768 .6 384 2 .00 FRANCE 407 70.4 770.0 397 1.94 410 78.4 775.2 392 1.98 014 7.3.6 732.0 384 1.91 AVERAGE 74 . I 759 . I 391 1.9a. AUSTfllA 423 81.2 838.0 400 2.1.0' YWO0 ‘13 740‘ 74°00 367 2.03 GREECE 416 72.0 765.2 394 1.94 417 74.4 747 .2.’ 389 1.92 418 85.2 744.0 367 2.03 419 62.0 722.0 379 1.91 420 70.4 731 .2 368 1.99 421 76.0 763.4: 380 2.01 .22 700‘ 72000 358 1.97 424 76. 0 792.0 41.4 1.91 425 70.4 796.3: 402 1.98 426 5106 77000 398 1.95 427 87.6 857.4”: 4414 1.93 428 86.0 781.2 401 1.95 429 74. 0 766.0 396 1.93 AVERAGE 76. 8 766 . 2 39 2 1. 95 TURKEY 40 I 73.2 762.0 381 2.00 404 76.4 814.0 us 1.95 408 78.4 843.2 1413 2.01;- AVERAGE 76.0 806.4 40.. 2,00 cameA 408 75.0 046.0 .141 1.92 64 Table 6. Statistical significance of the effect of fertilization and provenances on growth and needle characteristics 1. Z. 3. 4. 6. 7. 5. 9o 10. ll. 13. 14. lb. 16. 17. 18. 190 CO. ‘11. CHARACTtRIsTIC F VALUE DUt T0 FERTILIZATION PHDVENANCE HEIGHT 0R0 TN 19o4 13.4a** uQANCH ANGLL .vl wxmran uuew lacs-ca ~- NLCOLL LLNCTH c.37 NtLULL wIUTH 1.75 NELDLL LENGTH/“IUTH RATIO .70 NtLDLt COLOR .23 HthHT GHOuTn/wccULu LcNGTH RATIO 4.10 RESIN CANAL NUmucH 17.76wfi RESIN CANAL PUSITION .57 SEHHATIONS NUmutH PER 4 mm. 9.57% STOMATA NUMBER OF VENTHAL Hons b.00* STOMATA.NUNULH uF UURSAL Rois 10.37% STUmATA DUKan/thTRAL «ATlu 1.50 HYPOJfiRmAL LAYCNS Ow VthRAL blot 3.44% HYPUDEHMAL LAYcHS UN CUNNER .Du HYPODERMAL LAYthb Uh OUHSAL quE 1.98 SCLLNLNCHYMA a'éU‘~.ut.H UF LAYtHb 3. l b UIbTANCL. LJL.T\J.—L.'V- FI'UHUVAS uU-JULtS 1.39 ENDUDERN LtNQTh 1.13 ENUODtRm ulDTH ' 1.6J ENUODEHm LcNGTn/nIuTn RATIO .01 o.35** 3000*”; 0106.5")? 1.50 15‘7”} 4 . 37*‘3" 19.69** 4 od7“'* 10.db** b 044*‘54‘ J.ll** 7 odb** 3.66** 4.C7** 1.10 6.07** 11.91** 27.34** 7.41** lon 4050** 4 .4145“? l.b7* 65 Foliar Chemical ngggsition Differences in inherent ability of seedlings to absorb mineral nutrient ions from the soil medium have been demonstrated in forest tree species. Youngberg (1950) observed great differences in foliar chemical composition of Norway spruce. Seeds were collected from northern, central and southern Wisconsin and also from Sweden and Austria. He recognized three major groups which differed in the capacity of nutrient uptake: high (central and southern Wisconsin), medium (Austria) and low (northern Wisconsin and Sweden). The nutrient uptake capacity of Norway spruce originating in central Wisconsin was not depressed by extremely low fer- tility of parent soils. The work on Scotch pine by Gerhold (1959 a) and Steinbeck (1965) was of particular interest because this species has an overlapping range with European black pine in southern Europe. Gerhold's study included six seedlots; Steinbeck' work included more extensive sampling to cover the entire natural range. He found that the mineral composition varied not only between seedlots but also between plantations. Determination of error variance.-- Determination of error variance served two objectives. The first objective was to measure the signifi- cance of genetic, fertilizer, and genotype 3 fertilization effects. The second objective was to compare variability in.gingg.gig£3 and other species. This section is mainly for discussion on the first objective. I collected 32 samples from 8 of the 27 seedlots. Samples for each seedlot were kept separate fur fertilised and unfertilized trees from replicates l to S and 6 to 10 respectively (8 x 2 x 2 : 32 samples). 66 The coefficients of variability (a C. V.) for the 12 mineral ele- ments are shown in Table 7. They were calculated by dividing the standard \deviation by the plot mean and then multiplying by 100. The smaller the C. V., the more dependable the data. There was a close relationship between the C. V.'s of European black pine and Scotch pine, although the coefficients were slightly higher for black pine (Table 7). To confirm this a correlation analysis was con- ducted to compare variability patterns in the two species, using nutrient- element means as items. The correlation coefficient of the C. V. between European black pine and Scotch pine at Russ plantation was 0.906; between European black pine and Scotch pine at Newaygo plantation was 0.823; between European black pine and Scotch pine at Higgins Lake plantation was 0.863. The three values were all statistically significant (1 percent level) with 10 degrees of freedom. In other words, elements which were most variable in Scotch pine were also most variable in black pine. The strong correlation suggests that the error term derived from these two species may be applicable to others. Nitrogen.-- There were significant between-seedlot differences in foliar nitrogen level (Tables 8. and 9). Foliar nitrogen was lowest in the slow growing Corsican seedlots. Within the Greek population a slight pattern was recognizable. The three island-inhabiting seedlots (419, 426, 427) and the three (422, 424, 425) from the eastern part of the Creek mainland had nitrogen content below those from the rest of Greek seedlots. It is interesting to compare foliar nitrogen in.8uropean black pine with that found in Scotch pine by Steinbeck (1965). The range between the most extreme seedlots for European black pine was 1.28 to 1.60 percent; 67 Table 7. Coefficients of variability fer 12 mineral elements measure? in European black pine (l plantation) and Scotch pine (3 plan- tations . Mineral Species and plantation elements P. nigra P. sylvestria Kellogg Russ Newago Higgins ........ ----‘2352223_.--------------- N 6.2 6.1 h.6 h.7 K 7.11 7.2 7.7 73 P 6.1+ 7.11 6.2 7.2 Na h1.7 36.7 38.8 26.0 Ca 12.8 15.h 18.6 8.7 Mg 19.5 18.9 32.2 8.6 Mn 18.9 21.h 17.6 15.2 Fe 16.9 16.5 15.9 18.3 Cu 28.2' 17.5 22.9 27.1 B 19.5 12.9" 11.2 13.5 Zn 26.9 18.5 25.8 15.7 .Al 22.h 15.1 10.6 13.5 68 that for Scotch pine from the Russ Forest was 1.68 to 2.22 percent. Potassium.-- Poliar potassium content ranged from 0.55 to 0.66 percent in European black pine. According to Steinbeck (1965) the corre- sponding range for Scotch pine (Russ Forest) was similar (0.44 to 0.62 percent). Between-seedlot differences were significant at the 1 percent level (Tables 8a and 9). There was a slight geographic pattern to the variation. Corsican seedlots had less foliar potassium than did most others. Also, the three seedlots from the eastern part of the Creek.main- land had low potassium contents. Phosphorus.-- Analysis of variance showed that there were signifi- cant differences in foliar phosphorus content among the 27 seedlots (Tables 8a and 9). The Corsican seedlots had considerably below-average phosphorus content. For the rest of the range there was no particular pattern. The between-seedlot range in European black pine was from 0.15 to 0.18 percent; that in Scotch pine (Russ Forest) was from 0.19 to 0.27 percent. §g§igg.-- The between-seedlot differences in foliar sodium content were significant at the 1 percent level. Means for each seedlot are shown in Table Be. There was same pattern in foliar sodium content. Austrian and Corsican trees were characteristically low in this mineral element. Also, the three seedlots each from the Peloponnesus Peninsula (416, 421, 429) and from the eastern part of the Greek mainland (422, 424, 425) had low sodium content. The sodium content of European black pine ranged from 36 to 127 ppm. which overlapped with that of Scotch pine (18 to 134 pne.). 69 Table 8 a. Mineral element .110, K, P, He) concentration in mature needles of black pine origins. COUNTRY. ORIGIN NUMuaH CORSICA 411 412 413 AVERAGE SPAIN 402 403 AVERAGE FRANCE .07 «no .14 AVERAGE AUSTRIA 423 YUGO. 415 GREECE 416 417 416 419 “20 421 422 424 425 426 427 428 429 AVERAGh TURKfiY 401 404 405 AVERAGE CRINEA 408 1231 1241 PckCENT PcRCENT 1.31 .590 1.26 .576 1.33 .556 1.31 .573 1.46 .636 1.40 .600 1.44 .616 1.60 .636 1.49 .630 1.39 .595 1.49 .620 1.52 .650 1.52 .610 1.46 .665 1.56 .616 1.46 .610 1.42 .610 1.60 .636 1.40 .600 1.36 .656 1.37 .566 1.39 .650 1.42 .626 1.42 .640 1.48 .630 1.30 .610 1.43 .609 1.37 .630 1.44 .623 1.39 .630 1.40 .626 1.48 .630 (29) PERCfiNT .163 .147 .153 .152 .169 .161 .166 .173 .169 .175 .172 .173 .173 .167 .177 .177 .161 .167 .161 .149 .157 .161 .166 .169 .157 .153 .163 .167 .161 .149 .156 .163 NITROGEN POTASSlUM PHUbPHUHUS §ODIUM ‘26) PPM 40.0 55.0 54.0 50.7 96.6 61.0 66.0 72.0 36.0 69.0 59.0 43.5 67.0 66.5 66.0 121.0 127.0 76.5 50.5 43.0 48.0 58.0 52.0 79.5 67.0 69.0 72.7 116.5 67.0 56.5 66.3 61.9 70 um. 8b. 81mm]. .1...nt (6.. la, an, P., Cu) commutation COUNTRY. 0R1G1N NUHBER GORS1GA 411 SPAIN FRANCE AUSTRIA 7000. GREECE TURKEY CRIMEA 412 413 AVERAGE 402 403 AVERAGE 407 410 414 AVERAGE 423 415 416 417 418 419 420 421 422 424 425 426 427 428 429 AVERAGE 401 404 405 AVERAGE 408 1. mar. men... of black pin. origin. . 1271 (201 PERCENT PERCENT .315 .075 .315 .065 .346 .000 .326 .000 .395 .130 .375 .110 .385 .130 .330 .100 .360 .105 .360 .126 .350 .110 .345 .145 .340 .120 .335 .090 .375 .130 .406 .100 .315 .110 .890 .105 .270 .100 .339 .130 .330 .130 .306 .130 .270 .135 .266 .105 .330 .090 .270 .110 .311 .112 .310 .115 .330 .125 .360 .096 .333 .112 .340 .135 CALCIUM MAGNCS1UM MANGANESE (291 PPM 39&.0 266.0 304.5 267.5 296.0 295.0 296.6 266.5 336.0 332.0 315.6 305.0 265.0 226.6 294.0 297.6 349.5 360.6 406.5 272.0 336.0 317.0 300.0 296.5 328.0 312.0 314.6 247.5 336.0 263.0 262.5 260.0 1RON COPPER (301 1311 PPM PPM 43.6 6.66 45.0 7.30 42.0 7.25 43.5 7.72 66.0 11.00 58.5 10.75 63.3 10.33 43.0 11.66 45.0 9.00 53.0 7.96 46.7 9.63 53.0 0.40 43.5 10.95 40.5 6.50 54.0 17.50 64.5 13.00 61.5 9.46 66.5 7.50 60.0 10.30 05.6 9.00 46.0 10.76 60.0 10.70 49.5 9.00 54.0 11.65 63.0 10.75 66.5 7.70 56.6 10.36 52.5 9.05 51.0 10.30 49.5 13.90 51.0 11.00 47.0 11.00 71 Table 8c. Mineral element (B-'-, Zn, A1; NIP and Ill! ratios) concentration in mature needles of black pine origins . COUNTRY! BUNCH ZINC ALUMINUM NIP N/K ORIGIN RATlO RATIO NUMBER (32’ (33) (34) (35’ ‘36, PPM PPM PPM NUMbER NUMBER CORS!CA “1‘ 03065 3000 35405 5055 2024 412 11070 2300 87905 0059 30¢J “13 I4070 2700 38105 8069 20Q0 AVERAGE 1303b 2703 33605 6061 2029 SPIIN 402 ’9093 3905 36105 6079 4036 ‘03 10030 Q90b 30105 0069 2033 AVERAGE 09023 44.5 33605 507‘ 2035 FRANCE ‘07 13015 2905 30900 9024 2052 410 l‘070 3705 29605 605‘ 2035 41‘ 16090 3605 35300 7097 2035 ‘VERAGE IQ09Z 3403 30606 5067 ZoQZ AUSTRIA ‘23 19050 3405 39000 5053 2035 YUGO0 ‘15 1507b 3600 3330b 6060 605‘ GREECE 4‘6 {6070 2‘05 ‘9‘05 5070 2.:3 “I7 17090 4305 22400 “077 2053 41b 200l0 360: 31900 5036 2043 Qi9 20070 3205 35905 8052 2033 ‘20 18000 2505 40305 8093 3037 Q2! I709: 2205 33‘00 6076 C033 ‘22 2°0IO 2305 3‘505 90‘: 20‘5 424 2003§ 3000 26300 6079 2035 ‘25 2Z095 2405 30505 6063 3092 ‘26 17090 2305 Z7400 D061 £087 427 19063 3103 37400 “0‘2 2023 428 2(030 3905 4‘200 9042 2037 429 I8043 2600 33900 5049 20.3 AVERAGE 19053 2907 3'602 6077 3035 TURKEY 40! 37090 4'05 25700 6073 20‘? ‘04 20065 3505 42305 8095 3030 405 150¢b 3905 2‘600 9033 2022 AVERAGE 17093 3902 32005 9001 2033 CRIMEA ‘03 19000 4305 32000 9005 2036 72 “his 8d. Nina-01 0100001: (It/P, Ila/K. Gill's. and finite ration ammunhmndon>N>>Hv>> XXfiNKfiXflMX '4 I H 9‘ z 0'! O 8 H U \D The F values for each character are shown in.Appendix Table A - 1. In no case were differences significant between two needles in the same fascicle. Therefore, one needle per fascicle was considered adequate for the anatomical study. 114 Table A-1. F VALUL DUE TO AGL CLASS YEAR CLASb TREt tFFtCT NEEULL A A X X Y T K T N LFFCCT 115 Results of statistical analysis of 5 needle characteristics. NLLULC LtNGTH 4Je70** 1.33 IQCCOOL’37'x’X' 0.48 430782125 eYooso%* 0007 547090** ao9.39** 0004 0.46 0.74 mecuLL bLNKAWIUNb JIUTH PER a 0.04 l77.vl*% 0.001 3.00 0.4.v 96..YT** 0.0v 1.00 0.10 1.00 0.41 uo.10** 0.04 0.73 0.11 4.34 0.001 0.27 0.03 Gob“ 0.07 0.10n 0.000 0.47 0.001 0.03 0.001 0.84 QTUI'tAT A VL‘JVTl‘iAL RUJb 4.5 o UJ'X' ‘3“ 15.71** 100 0 ‘J3';(' ‘7‘ 0°13 4.14 IJOUJ'X‘ 0044 LJo‘J‘J 0.31 1.17 1000 0040 0.31 STUHAT A UUNbAL HUHD 001.01** 0.0C 3e0.79** 0.38 bled7** bdso7** 0°30 bl.vv** 0.02 a.ao 10.77% JOQU lebO OeUl 116 TableA-z, Analysis of variance of 27 different black pine provenances for growth and needle characters from Kellogg planting (Augusta, Michigan) Height Growth, 1964 SOURCE or DEGS. 0F SUM VARIANCF FREEDOM ‘OF SQUAHFS MEAN SQUARE F STATISTIC BLOCKs 9 6656.60135195 628.51131689 7.35818 48(- MAIN (A) 1 1322.2685185?1322.26851852 15.48022 ERROR 1 9 768.75000002 85.41666667 *3!- SUB (H) 26 19919.4oonon43 766.13076925 6gfl£6 AB INTFRACTION 26 3026.08148151 116.38774929 .96141 ERROR 2 11,32 52310.14814026 ':120.715726h6 TOTAL(AFTER rFAN)5xB 83003.2500U164 Branch Angle SOURCE OF DEGS. OF $0M VARIANCE FR§E00fi__HQF SOUAHES MEAN SQUARE F STATISTIC BLOCKS 7 15144.201360192163.45734130 6.76049 MAIN (A) 1 162.55787n37 162.55787037 .50947 ERRQR 1 7 2233.49768523 319.07109790 §** sue (m 26 37099.074074651426.88746440 3,878]? AB INTERACTION 26 5720.37962978 220.014.0114 .5980? spam: 2 3365123986.1759277o""‘369.006u7§ TOTALtArtea HEAN1le_184345.88639974 117 Table A-3. Analysis of variance of 27 different black pine provenances for growth and needle characters from Kellogg planting (Augusta, Michigan) TOTAL (AFTER MEAN) 503 103“”.59259808 Winter Burn, 1963-Bu SOURCE OF DECS. OF SUM VARIANCE FREEDOM OF SQUARES MEAN SQUARE F STATISTIC BLOCKS 9 65.12 7.236 u7.29** A 26 2us.16 9.u29 61.63** ERROR 231 35.28 0.153 TOTAL 266 395.56 Needle Length TSOURCE:OF DECS. OF SUM :VARIANCE FREEDOM OE SQUARES MEAN SQUARE F STATISTIC BLOCK 9 719.33333333 79.92592593 9.9229u MAIN (A) 1 38.nooooooo 38.nooooooo 2.36520 ERROR 1 9 196.11851852' 16.23539095 SUB (B) 26 uoo2.0925926u 153.92663818 13.1823** AB INrERACTION 26 378.70000000 lu.56538u62 1.2u7u ERROR 2 932 5059.9u81us26 11.7128u293 118 Table/H}. Analysis of variance of 27 different black'pine provenances for growth and needle characters from Kellogg Planting (Augusta, Michigan) Needle Width SOURCE OF OEOS. VARIANCE FREEDOM BLOCKS 9 MAIN (A) 1 ERROR 1 9 SUB (B) N 26 A9 INTERACTION 26 ERROR 2 ' h32 TOTAL1AFTER MEAm>503 SUM OF SOUARES MEAN SQUARE F STATISTIC 2.62666667 .232?9630 1.20915519 6.60970370 1.81770370 24.10314615 36.6037U427 .29185185 .23229630 .13435391 .25421937 .06991168 ~05563hl9 Needle Length/Width Ratio SOURC? OF DEGS. VARIANCE FREED0M_ BLOCKS . 9 MAIN (A) 1 ERROR 1 9 SUB (8) 26 AB INTFPACTION 26 ERROR 2 -h32 TOTAL( AFTER MFAN)EQ_3. SUM 2.17226 1072899 b.5695 1.2566 0F SQUARFS MEAN SQUARE F STATISTIC 34.16378504 .90942194 10.77571556 625.37972083 42.43968363 529.46735783 1243.13670121 3.79597612 .90942194 1.1973017? 24.05306653 1.6322955? " 1.2218h77 3.17044 075956 ** 19.6858 1.3359 119 TableA-s, Analysis of variance of 27 different black pine provenances based on one compoSite sample from Kellogg planting (Augusta, Michigan) Needle Color ‘DOURCF 0F DEBS. OF SUM 1. - . VARIANCF FREEDOM 0? SQUARES MEAN SQUARE F STATISTIC bLOCKS 1 , 1.18518519 1.18518519 .2337? ** ‘ 26 553.14314817 ;?1.65954416 ”_ 4.27227 ERROR 26 131.814P1479 5.06980n57 TOTAL(AFTFR HFAM153 696.14814814 Tablefl-J‘Analysis of variance of 27 differnet black pine provenances for growth and needle characters from Kellogg planting (Augusta, Michigan) Height Growth/Needle Length Ratio SOURCE Or 0668. OF SUM VARIANCE FREEUQM _0F SQUAHFS MEAN SQUARE F STATISTlC BLOCKS 9 3.21357020 .35706336 .96664 MAIN (A) 1 1.51353352 1.51353352 4.09742 ERROR 1 9 3.32448355 .36938706 *3!- SUB (R) 26 131-55224079 5.05970157 lOJfiDl A8 INTFRACTIUN 26 15.79354960 .60744422 Ju30fl3 ERROR 2 11132 201.88809416 .h658956 TOIALtAFfER REAN1603 357.28547109w 120 Tablekb. Analysis of variance of 27 different 'black pine provenances for growth and needle characters from Kellogg planting (Augusta, Michigan) Resin Canal Number SOURCE OF DECS. VARIANCE FREEOOM BLOCKS 9 MAIN (A) 1 ERRnR 1 9 SUB (P) 26 AR INTFRACTION 26 ERROR 2 E32 IOTAL1AFTER MEAN)503 0F SUM OF SUUARES MEAN SQUARE F STATISTIC “3.00195165 74.816606o7 37.9“555556 706.0067037? 07.43336363 1398.19299261 7397.3557U353 Resin Canal Position SOURCE OF HESS. VARIANCF FREEDOM BLOCKS 9 MAIN (A) 1 ERROR 1 9 SUB (D) 26 Ag IMTERACTIQM 26 ERROR 2 h32 TOTALtAFTFR MEAN)503 SUM 9.22242798 74.61666667 4.21172840 27.15398960 3.74743590 3:31 2.18970 {Hi- 17.76389 8.hlS7 1.161h nF SQUARES MEAN SQUARE F STATISTIC 1.19490741 .03750000 .59750000 5.0759259} 1.85000000 32.59259250 V42,33842588. .13276749 .03750000 .06527778 .23368946 .07115385 .07521367h 2.03388 .5744? 3.1070’ .9h&) *3!- 121 TableAJZ.Analysis of variance of 27 differnet black pine provenances for growth and needle characters from Kellogg filnating (Augusta, Michigan) Number per 2 mm. 0F SQUARES MEAN SOUARE F STAT!STIC Serrations. SOURCE OF DEBS. SUM VARIANCF FREEDOM_ BLOCKS 9 96.82222222 MAIN (A) 1 17.06666667 ERROR 1 9 15.56296296 SUB (R) 26 827.83333354 AB INTERACTION 26 98.63333333 ERROR 2 h32 SOURCF OF OEGS. VARIANCE FREEDOM BLOCKS MAIN (A) ERROR 1 SUB (R) 26 AR INTERACTION 26 ERROR 2 U32 TOTAL(AFTFR MEAN)593 1755.01481481 2810.9333506? RUM OF SQUARES MEAN SQUAR 113.75000000 ‘8.93518519 43.82407407 359.80000001 “1.81481481‘ A675.i?592594 .2§?3.2SOOOSOO 10.75802469 17006666667 1.72921811 31.83925359 3.79358974 h.OsOO3A2 Stomata, Number of Ventral ROWS 12.63658669 38.93518519 4.96934156 14.22307692 3.14672365 3.8656752 6022132 1* 9.86959 7.8616 .9367 E r STAT!STIC 2.59561 * 7.99599 3.6793 .81M3 *4! 122 Tablefl-B. Analysis of variance of 27 differnet black pine provenances foo growth and needle characters from Kellogg Planting (Augusta, Michigan) Stomata. Number of Dorsal Rows SOURCE O1 OEOS, VARIANCF FREEDOM BLOCKS 9 MAIN (A) 1 ERROR 1 9 SUB (R) 26 AR INTFHACYION 26 ERROR 2 O32 TOTAL(AFTER MEAN)SO3 SUM OF sOUARES MEAN SQUARE F STATISTIC 196.69629660 38.4OODUOUO 33.340740/4 850.10370371 218.2000UOOO 3314.36200299 4641.10371409 20.74403292 38.40000000 3.70452675 32.69629630 8.39230769 7.6h85299, Stomata, Dorsal/Ventral Ratio SDURCF OF OEOS. VARIANCF FRFEOOM BLOCKS 9 MAIN (A) ‘ 1 ERROR 1 9 SUB (R) 26 AB IMTFRACTIWN 26 ERROR 2 U32 TOTAL(AFTER MEAN)503_ SUM 5.59964 : * 10.36570 ** h.27h8 1.0972 OF SQUARES MEAN SQUARE F STAT!STIC .43476827 .05033923 .24372839 .93992066 .74541241 14.25668599 16.72035537 .05350759 .05033923 .02708093 .03615080 .02866971‘ .03290 1.98692 1.85884 1.0988_ .87lh 123 Tableflqfl.Ana1ysis of variance of 27 different black pine provenances for growth and needle characters from Kellogg planting (Augusta, Michigan) Hypodermal Lavers. 0n Ventral Side SOURCE OF WEGS. OF SUM VARIANCE FREEDOM OF SOUARES MEAN SQUARE F STATISTIC BLOCKS 9 25.41.66.666? 2.82407407 8.615751% MAIN (A) 1 1.77962963 1.77962963 5.44305 ERROR 1 9 2.94259259 .32695473 ‘18? SUB (R) 26 65.39148148 2.51467236 OJMBQ AR INTERACTION 26 5.67037057 .21809117 .526h ERROR 2 1132 179.54074OM oblb32h8 IOTAL(AFTFR MEANISO3 290.7314B156 HYPodermal Layers, On Corner SOURCE OF OEGS. OF SUM VARIANCF FREEDOM OF SQUARES MEAN SQUARE F STATISTIC BLOCKS 9 12.0914814R 1.34238683 6.55020 RAIN (A) 1 .11851852 .11851852 .57631 ERROQ 1 o 1,54444444 ,20493827 SUB (R) 26 (63.10000000 5.66846154 11.2xn1_ Ag INTERACTION 26 8.18148148 .31467236 .63m1 ERROR 2 1.32 214.2740'7408 oh9hh786b,’ TOTALHFTER MCAV>SO3V 389.639non39 124 Table/HQ Analysis of variance of 27 different black pine provenances for growth and needle characters from Kellogg Planting,(Augusta, Michigan) Hypodermal Layers, On Dorsal Side SOURCF OF OEGS, VARIANCF FREEOON BLOCKS 9 MAIN (A) 1 ERROR 1 9 SUB (R) 26 AR INTFRACTION 26 ERROR 2 h32 IOTAL(AFIER NFAN)SO3 SUM OF sOUARES MEAN SQUARE F STATISTIC 14.66851852 ,97962903 4.59444444 311,58925926 16.27037067 262,53703704 570.90925963 1.62983539 .97962963 .51049383 12.75227920 .62578348 .h68085h7 Sclerenchvma, NUmber of Layers SOURCF OF OEGS. VARIANCE E'I""_-Ef.:.00’4 BLOCKS 9 MAIN! (A) l ERROR l 9 SUB (R) 26 AB INTERACTION 26 ERROR 2 h32 Iguu .. . 3.19266 1.91898 %*. 27.2U3S 1.3369 OF SQUARES MEAN SQUARE F STATISTIC 9.32592593 .47407407 1.34074074 49.93333333 4.37592593 112.33333333 177.73333335‘ 1.03621399 .47407407 .14897119 1.92051262 .16638177 .2592308 6.95580 3.18232 7.h085 061.118 125 TableA-fl, Analysis of variance of 27 different black pine provenances for growth and needle characters from Kellogg planting (Augusta, Michigan) Distance Between Fibrovascular Bundles SOURCF 0+ DECS. or sun VARIANCE FHFEDOM or SQUARES MEAN SQUARE F STATISTlc BLOCKS 9 38.84305556 4.31589506 4.27957 MAIN (A) 1 1.40046296 1.40046296 1.38868 ERRnR 1 9 9.07638850 1.00848765 SUB (R) 26 1 45.76759259 1.76029202 1.3176 AR INTFHACTI”N 26 37.01203704‘ 1.42353989‘ 1AM66 ERROR 2 L532 578.90555557 1.3359359 TOTAL(AFTFR HFAN)SO3 711.0n509264 Endoderm. Length SOURCE OF RERS. OF RUM , VARIANCF FREEDOM or SQUARES MEAN SQUARE F STATISTIC BLOCKS 9 7n9,16491483 78.79609053 2.71709 MAIN (A, 1 32.75740741 32.75740741 1.12956 ERROR 1 9 2‘1.on195155 29.00020576 sue (R) 26 9084.33333336 79.01282051 hxfifl9 AR INTFRACTIRN 26 424.7025926016.33817664 1xmus ERROR 2 .1132 7048.13636345 16.26h9231 TOTAL(AFTEH McAu>§gg ’1n510.19334957 126 TableA42.Analysis of variance of 27 different black pine provenances for growth and needle characters from Kellogg planting (Augusta, Michigan)_ Endoderm, Width SOURCF nF DECS. VARIANCF FRFEDOM BLOCKS 9 MAIN (A) 1 ERROR 1 9 SUB (R) 26' AR INTFRACTIDN 26 ERROR 2 hgg TOTALtAFTFR MFAM)§O3 RUM OF SQUARES MEAN SQUARE F STATISTIC 316.44629630 7.8?407407 38.56481451 .543.67037069 66.4?592593 9119.08898891 3112.0?037054 35.16069959 7.82407407 4.28497942 70.91039886 2.55484330 h.936359 Endoderm, Length/Width Ratio SnuRcc nF HESS. VARIANCE FREEDOM BLOCKR 0 MAIN (A) ' 1 ERRnR 1 9 SUB (R) 26 AR IMTFRACTION 26 ERROR 2 hBé TOT1L(AFTE& MFAU)SQ§ quu 8.20557 1.82593 b.2360 -5176 *4? GP SQUARES MEAN SQUARE F STATISTIC 1.6991211“ .00022743 .15984367 1,29952899 .616P6060 13.76479306 17.53947569 .18879123 .00022740 .01772710 .04998188 .02370234 .03176h9 10.64987 .01283 1.5735 .7h62 {- 127 TableAJAAnalysis of variance of 27 different black pine provenances based on one comoosite sample from Kellogg planting (Augusta, Michigan) Foliar Nitrogen Content SQURCF OF DECS. OF SUM VARIANCF FRFtDOM OF SOUARES MEAN SQUARE F STATISTIC W“ BLOCKS 1 .04800185 .04800185 29.68141 v A 26 .32815926 .01262151 _1.228°4§Z_ ERROR 26 .042n4814 .00161724 TOTAL(AFTER MFAN) 53 .41820925 Foliar Potassium Content SOURCE or ‘ DEGSJ 0F SUM V‘RIANCF FREEQOM or SQUARES MEAN SQUARE r STATxSTIc BLOCKS 1 : .05226667 .05226667 94.an9s1*% A . 26 ,04228148 .00162621 2.949R7** ERRnR 26 .01433333 .00055128 TOTAL(AFTER MFAN) 5§_ .10888148 111. _( _ Foliar Phosphorus Content SOURCE OF DECS. 0F RUM VARIANCE FREEDOM or SQUARES MEAN SQUARE F STAT!STIC BLOCKS 1 .on034757 .00034757 6.18575* A 26 .00400433 .00015401 2.7«1096wr ERROR 25 .00146093 .000U5619 TOTAL(AFTER MEAN) 53 .00581283 128 Tablé‘4éfinalysis of variance of 27 different black pine Provenance based on one composite sample from Kellogg planting (Augusta, Michigan) Foliar Sodium Content MEAN SQUARE F STATISTIC SOURCF OF UEGS. OF 8UM VAR!ANCF FREEDOM OF SQUARES BLOCKR 1 1688.96296298 1688.96296298 A 26 3ne74.33333421 1187.47435R99 ERROR 26 12766.03703918 491.00142458 TOT‘L(AF'TFR MEAN) 53 45379033336665 Foliar Calcium Content SOURCE OF HESS. or VARIANCF FREEDOM OF ULOCKS 1 A 26 ERROR 26 TOTAL(AFTER MFAN)53 SUM SOUARES .00009074 .07350000 .02315926 .09675000 MEAN SQUARE .00009074 .00282692_ .00089074 Fbliar Magnesium Content SOURCE OF OEOS. OF SUM VARIANCE FREEDOM BLOCKS 1 .00106667 A 26 .01773333‘ ERROR 26 TOTAL(AFTFR MRAU)53 00593333 02473333 .00106667 .00068205 .00022821 3.43983 2.41847 910187 3.17368 0F SQUARES MEAN SQUARE F STATISTIC 4.67416 2.98876 F STATISTIC * #8? 3“! 4H? 129 TableAdfiAnalysis of variance of 27 different black pine provenances based on one composite sample from Kellogg planting (Augusta, Michigan) Foliar Manganese Content SOURCE OF OEGS. 0F SUM VARIANCE FREEDOM OF SQUARES HFAN SQUARE r STATISTIC ULOCKR 1k 699n8.16666710 68908.16666710 57.83063** A 26. 73450.9259273? 2825.03561255 2.37oR9* ERROR 26A 3n9R0.33331335 1191.55128130 TOTAL(AFTFR REAM)53'17333R.42590650 Foliar Iron Content SOURCE OF DECS. 0F SUM VARIANCE FREEDOM OF SQUARES MEAN SQUARE F STATISTIC BLOCKR 1 .296?9630 .29629630 .00675 A 26 3166.81481467 121.41595442 2.76500** ERROR 26 1141.70370056 43.91168079 TOTALtAFTER MEAN)53 4298.814R1177 Foliar Cooper Content SOURCE OF OEOS. 0F SUM VARIANCE FREEDOM. OF SQUARES MEAN SQUARE F STATISTIC BLOCKS 1 12.61500000 12.61500000 2.72530 A 26 296.01814816 11.38531339 2.45964* ERROR 26 120.35000030 4.62884617 TOTAL(AFTER HEAN)53 439-983‘4846w— 130 TableAJQAnalysis of variance of 27 different black pine provenances based on one composite sample from Kellogg planting (Augusta, Michigan) Foliar Boron Content SOURCF OF OECS. OF 9U» VARIANCE FREEDOM OF SOUARES MFAN SQUARE F STATISTIC BLOCKS 1 1‘6.40018520 156.40018520 43.55130** A 26 4n9.51333332 15.75051232 _ 4.5R643** ERROR 26 R3.754915% 3.22133906 TOTALmanw¢ERb~oeso~qxru\4m<>hsmcno~&c> we» bu) 0 o Table A-29. (continued) M K P NA CA "G 419F110 1.48 0.60 0,169 77 0.36 0.11 4190110 1.36 0.69 0.153 177 0.?7 0.11 420F105 1.54 0.55 0.177 82 0.?7 0.10 4200105 1.44 0.65 0.161 82 0.30 0.10 420F610 1.54 0.65 0.161 77 0.30 0.11 QZOUGIO 1.48 0968 09169 6.3 ”027 0910 421F110 1.38' 0.55 0.145 38 0.27 0.09 4210110 1.42 0.69 0.177 63 0.27 0.11 422F110 1.40 0.53 0.153 54 0.30 0.1? 54220110 1.32 0,50 0,145 32 0.27 0.14 423F105 1.52 0.62 0.169 36 0.51 0.14 4230105 1.44 0.68 0.169 32 0.30 0.15 423F610 1.60 0.62 0.169 42 0.27 0.1? 4230610 1.54 0.60 0.105 63 0.30 0.16 424F110 1.40 0.55 0.145 54 0.33 0.1? 4240110 1.34 0.6? 0.169 42 0.33 0.1? 425F110 1.48 0.55 0.161 58 0.27 0.11 4250110 1.30 0.55 0.161 50 0.30 0.15 426F110 1.44 0.60 0.161 50 0.27 0.12 4260110 1.40 0.65 0.169 54 0.27 0.19 427F110 1.42 0.58 0.161 77 0027 001“ 4270110 1.42 0.70 0.177 a? 0.30 0.11 428F110 1.54 0.58 0.161 72 0.36 0.09 4280110 1.42 0.60 0.153 63 0.30 0.09 429F110 1.34 0.60 0.153 50 0.24 0.10 4290110 1.?6 0.62 0.153 8»0 0-30 0-1? 144 401 THROUGH 429--SEED ORIGIN NUMBER F--FERTILIZED SUBPLOT U--UNFERTILIZED SUBPLOT 105--COMPOSITE SAMPLE.REPLICATES 1 THROUGH 5 610--COMPOSITE SAMPLEcREPLICATES 6 THROUGH 10 ...__-_H .—— llO--COMPOSITE SAMPLEoREPLICATES 1 THROUGH 10 ~H—_q~ .2... MN 379 320 434 350 344 314 434 379_ 326 218 391 274 303 251 326 326 320 314 320 280 373 224 416 240 367 297.. FE 48 75 69 75 54 66 60 60 54 57 51 51 60 40 48 48 57 63 57 42 48 60 69 57 63 54 C) L: 1.; 14 P1P ~04H+a Hb‘Hwb , mcrhsorohsowacapcuxum~qaumc:\Hacaoc2arv\q» O O I O O O C O C O O O O O ocn01u1461m0004mwnxdo~qo~044~491uCDa33‘d\JN 145 Table A—29. (continued) 3 ZN AL 8 ZN AL 401F105 16.8 34 232 419F110 17.9 34 32? 401u105 17.9 48 329 419u110 23.5 31 397 401F610 16.8 42 307 420F105 19.0 26 444 401U6lo 20.1 42 279 '420U105 24.6 36 510 502F105. 19.0 39 444 420F610 14.7 24 207 402u105 22.3 42 444 420U610 16.8 28 452 402F610 13.7 34 213 9215110 14.7 21 245 402U610 24.6 42 344 1421u1lo 21.2 24 397 403F110 15.8 45 279 1422F110 20.1 19 300 403U110 21.2 54 344 .422U110 20.1 26 397 4040110 22.3 42 468 423v105 21.2 39 307 405F110 14.7 34 213 9231:8111 16.8 28 300 405U110 15.8 45 279 9923116117 2041 42 381 407F110 12.6 31 252 “24F110 16.8 21 226 407mm 13.7 28 366 “2“”“0 23.5 39 300 .AOBFIOSL 14.7 48 351 425F110 20.1 21 P66 408U105 16.8 57 374 “250110 25.8 28 351 408F610 12.6 34 219 426F110 17.9 21 272 408U61o 15.8 ,54 336 ‘26U**° 17.9 26 272 410F110 14.7 35 3‘5 427F110 15.8 2‘ 351 4100110 14.7 39 27? “27W” 23.5 39 397 4115110 12.6 26 351 ”28“” 22.3 34 42n 411U,10 14.7 34 358 428““0 22.3 45 404 412F110 9.7 24 245 429F110 17.9 24 272 412U110 13.7 26 314 “9W0 119.0 28 358 413F110 12.6 26 ‘412 413u110 16.8 28 351 314F105 13.7 31 232 414u105 20.1 39 404 414F610 14.7 31 23? 414U610 19.0 45 544 415F105 18.8 31 397 415u105 22.3 42 358 415F610 15.8 28 239 41SU6+9- 20.1 42 300 416F105 14.7 19 188 416U105 16.8 28 93 416F610 11.8 19 170 416U610 23.5 31 314 417F110 18.8 42 176 417u14o» 19.0 45 272 418F110 20.1 31 272 418U110 20.1 42 366 Table A-30. FEEL UNFER. MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG 401 1402 1403 404 4405 407' 408 410 411 412 4421 414 415 416 417 418 419" 420 421 422 423 424 hflfiaG—AZSL MSFG MSFG MSFG 426 427 428 146 Plat means (fertilized and unfertilized) and standard deviations for the measurement of 27 black pine seedlots Height Growth, 1964 MEAN 49.14 46.01 47.85 47.85 50.35 52.05 48.15 52.30 55.20 49.50 37.05 34.90 36.15 46.70 52.35 38.50 43060 47.85 50.20 45.60 39.85 50.70 46.40 50.65 63.05 44.65 52.05 50.85 Nfiflfl31429 45.40 Overall Mean & S. D. 47.58 5.0. 11.492 13.098 9.943 8.362 *8.393 9.087 8.387 14.907 11.482 13.975 11.019 9.391 11.793 12.139 6.945 7.030 19.896 10.168 12.878 12.604 11.412 7.462 8.665 10.121 16.040 14.426 Branch Anghe MEAN 121.72 120.49 118.87 123.12 122.87 128.75 130.75 118.43 134.37 121.25 114906 97.62 104.68 103.75 127.81 126.37 121.56 126.25 119.06 132.50 109.37 119.31 129.06 123.56 119.31 110.31 14.416 131.25 9.964 132.50 11.080 123.12 12,409 121.10 SOD. 20.087 21.287 18.205 20.645 14.916 13.964 11.952 22.782 15.692 14.776 24.373 19.649 22.543 27.958 14.602' 10.506 26.502 15.545 19.080 20.575 24.281 19.327 16.655 21.096 23.570 18.117 14.083 18.073 9.756 20.681 Needle Length MEAN 5.0. Needle Width MEAN SOD. 18.974.381 2.66 15.15 24.65 24.25 18.95 18.15 21.85 20.30 19.90 21.00 20.10 21.05 23.30 17.35 15.35 17.60 17.60 18.40 14.85 14.90 14.90 17.60 18.45 17.25 17.95 20.15 18.00 16.00 18.70 2.758 3.313 3.041 3.170 3.963 5.294 3.113 3.338 4.255 3.782 5.286 3.798 2.497 2.996 1.465 2.500 3.393 2.906 3.851 3.275 2.799 2.314 3.252 5.742 2.575 3.879 4.380 2.63 2.54 2.59 2.74 2.79 2.69 2.80 2.76 2.66 2.70 2.41 2.56 2.60 2.52 2.52 2.58 2.55 2.49 2.52 2.77 2.67 2.69 2.62 2.90 2.61 2.62 2.64 .250 .268 .227 .260 .204 .230 .284 .360 .218 .250 .260 .396 .330 .189 .163 .288 .143 .186 .176 .173 .288 .180 .197 .145 .184 .160 .339 .223 .214 .260 Table A-Bl. 147 Plot means (fertilized and unfertilized) and standard deviations for the measurement of 27 black pine seedlots Needle, Length/ Width Ratio MEAN SQDO FEEL 7,1 1,50 UNFER. 7,0 1.53 MSFG 401 5.7 .93 MSFG 402 9.7 1.24 MSFG 403 9.3 1408 MSFG 404 6.9 .89 MSFG 405 6.4 1.23 MSFG 407. 8.0 1.13 MSFG 408 7.2 1.07 MSFG 410 7.2 .94 MSFG 411 7,5 1.36 MSFG 412 7.7 1.26 MSFG 413 7.7 1.46 045502-414 9.6 1.42 MSFG 415 6.7 .86 MSFG 416 5.8 .70 MSFG 417 6.9 .54 MSFG 418 6.9 .89 MSFG 419 7.1 1.14 msasp420 5.8 .96 MSFG 421 5.9 1.12 MSFG 422 5,8 1.05 MSFG 423 6.3 2..83 MSFG 424» 6.8 .91 MSFG 425 6.4 .83 massanze. 6.8 1.06 MSFG 427 6.8 1.47 MSFG 428 6.8 .81 MSFG 429 EH1 1.54 Overall Mean 84 7.0 1.51 S. D. Height/ Needle Length Ratio MEAN 2.69 2.59 3.?6 1.95 2.11 2.79 2.75 2.38 2.71 2.52 1.76 1.74 1.74 2.02 3.06 2.60 2.77 2.72 2.74 3.05 2.86 3.52 2.66 2.75 3.70 2.50 2.75 2.87 2.97 2.64 5.0. .788 .837 .983 .297 .494 .563 .701 .331 .409 .761 .365 .334 ,443 .549 .478 .692 .582 .474 .662 .598 .200 .729 .471 .445 .025 .682 .952 .688 .896 .814 Resin Canal Number MEAN 7.58 6,83 6.60 8,80 8.35 6.30 5.70 7.20 6.10 7.55 8.65 8.45 9.40 8.35 6.95 5.55 7.20 6.70 5.45 6.35 5.55 8.60 8.60 7.?5 8.15 7.35 6.20 7.30 6.00 5.0. 2.097 2.057 1.846 2.462 2.158 1.174 1.341 2.546 1.119 2.114 2.580 1.904 2.161 2.368 1.394 1.190 2.015 1.688 .944 1.308 1.190 2.112 2.280 1.208 1.755 1.531 1.399 1.780 1.716 7.20 2.108 MEAN 3.89 3.91 3.00 3.62 3.72 3.95 3.97 3.87 3.92 30R7 3.77 3.90 3.85 3.67 3.07 4,00 4.07 4.02 3.05 3.95 3.97 3.95 3.95 3.97 3.97 3.95 3.97 3.00 3.95 Resin Canal Position Table A-32. FERT. UNFER. MSFG_ MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG- MSFG MSFG MSFG MSFG, MSFG MSEG_ MSFG MSFG MSFG MSFG MSFG 401 4o2 403 404 405 407 4oa 410 411 412 413 414 415 416 417 418 419 420 .421 422 423 424 425 426 BHNFG_AUIZ MSFG MSFG 1428 «429 Overall Mean & S. D. 148 Plot means (fertilized and unfertilized) and standard deviations for the measurement of 27 black pine seedlots Serrations, Number per 2 mm MEAN 15.35 15.00 15.35 13.35 13.90 15.15 16.30 16.10 15.05 16.35 17.60 18.35 16.25 12.75 14.10 15.90 14.65 15.15 13.35 14.15 15.25 14.70 14.15 14.90 14.25 15.75 16.05 15.65 15.30 15.17 SOD. 2.380 2.172 1.871 1.926 1.552 2.158 1.559 1.889 1.503 1.755 2.414 3.082 2.672 2.314 1.552 1.832 1.225 1.785 1.089 1.871 3.058 1.949 1.565 1.860 2.173 1.970 1.959 1.225 1.65T 2.283 Stomata, No. of Ventral was MEAN 15.18 14.64 15.10 14.30 14.65 15.25 15.80 15.20 16.05 15.55 15.65 14.65 .15.05 13.90 14.05 14.45 14.70 14.10 13.25 14.4.; 13.80 14.35 15.30 15.50 14.70 15.70 17.30 14.70 13.60 14.91 SOD. 2.046 2.074 2.337 1.625 1.268 1.831 2.067 2.647 2.139 2.305 2.230 2.134 2.350 2.174 1.495 2.187 1.174 2.023 1.551 1.704 2.284 1.755 1.750 1.147 1.838 1.688 2.735 1.454 1.569 2.076 Stomata. No. of Dorsal was MEAN 23.30 22.77 23.15 23.06 24.00 23.50 23.75 23.75 24.80 24.20 23.55 21.35 22.45 21.95 22.35 22.40 22.60 21.80 20.40 29.05 20.75 22.95 24.00 24.25 23.75 24.00 26.35 29.85 21.00 23.04 5.00 2.899 2.950 2.942 2.665 2.176 2.212 1.551 3.611 2.764 3.155 3.347 3.375 3.203 2.999 1.980 3.500 1.818 2.117 1.667 2.855 2.613 1.731 2.000 1.915 3.226 1.622 4.704 2.134 2.663 2.934 Stomata, Dorsal/ Ventral Ratio MEAN SOD. 1.54 1.56 1.54 1.61 1.65 1.55 1.52 1.58 1.56 1.56 1.51 1.49 1.50 1.60 1.51 1.50 1.54 1.56 1.55 1.49 1.52 1.61 1.58 1.56 1.62 1.54 1.53 1.56 1.59 1.55 .178 .173 .189 .150 .216 .176 .165 .208 .201 .138 .191 .114 .148 .230 .107 .177 .138 .194 .141 .166 .204 .182 .180 .101 .232 .173 .211 .145 .149 .176 149 (fertilized and unfertilized) for the measurement of 27 black Table A-33. Plot means and standard deviations pine seedlots Hypodermal Hypodermal Hypodermal Sclerenchyma. Layers, 0n Layers, 0n ILayers, On No. Of Layers Ventral Side Corner Dorsal Side MEAN $000 MEAN $.00 MEAN SOD. MEAN SOD. FERT. 2.49 .765 4.28 .572 3.551.014 1.78 .563 IDEER. 9,37 .671 4.75 .829 :3y471.044 1.72 .583 MSFG 401 5?Uhi.409 4.50 .760 4.30 .571 1.95 .223 MSFG_402 2.00 .000 3.80 .615 7.30 .571 1.75 .444 MSFG 403 2.10 .447 3.95 .604 7.45 .759 1.75 .550 MSFG 404 2.45 .759 4.65 .670 3.90 .552 2.00 .000 MSFG 405 2.90 .788 5.25 .716 4.00 .458 1.90 .307 MSFG 407 2.00 .000 3.85 .366 2.95 .825 1.45 .686 MSFG 408 2H75 .850 4.70 .854 4.45 .604 2.00 .000 MSFG 410 2.10 .447 3.55 .604 2.651.039 1.35 .875 MSFG 411 2.00 .000 3.15 .312 2.30 .571 .85 .933 MSFG 412 2.05 .223 3.75 .716 2.15 .366 1.00 .858 MSFG 413 2.00 .000 3.10 .342 2.15 .489 1.30 .732 MSFG 414 2.00 .000 3.65 .670 2.10‘ .447 1.85 .489 MSFG 415 2.40 .598 4.40 .598 3.65 .745 1.80 1.410 _MSFG_-416 2.15 .489 4.85 .670 4.40 .680 1.95 .686 M5522“; 2.1.5 .366 4.15 .409 3.00 .615 1.90 .307 MSF’G 418 2.65 .875 4.45 .759 3.90 .447 1.75 .444 MSFG 419 2.55 .686 4.20 .615 3.60 .753 1.70 .571 MSFG 420 2.50 .760 4.40 .753 3.65 .812 1.80 .410 MSFG 42. 2.50 .760 4.1.0 .552 3.70 .864 1.85 .366 MSFG 422 7.60 .753 4.75 .638 3.90 .552 2.00 .000 MSFG 423 2.55 .686 4.25 .444 3.95 .394 1.65 .587 MSFG 424 2.75 .850 4.6-5 .812 4.00 .725 1.90 .307 MSFG 425 2.95 .944 4.35 .670 4.10 .447 2.05 .223 MSFG 426 7.35 .670 4.10 .307 3.75 .850 2.00 0561 MSFG 427 3.20 .894 4.95 .759 4.35 .933 2.00 ‘.000 0051:5423 2.95 .944 4.75 .786 3.95 .944 1.90 .307 MSFG 429 ?0RO .894 4095 6825 4045 I759 Zeno .000 Overall Mean & 2.43 .721 4.26 .850 3.51 1.029 1.75 .574 S. D. Table A-34. and standard deviations for the measurement of 27 black 150 Plot means (fertilized and unfertilized) pine seedlots FERT. UNFER. MSFG MSFG MSFG MSFG- MSFG MSFG MSFG MSFG MSFG M556— MSFG MSFG MSFG MSFG MSFG 401 1402 403 1404 1405 407' «408 410 411 4121 413 414 415 416 417 JWSFG%4¥%8» MSFG MSFG MSFG MSFG MSFG 4419 42K) 421 4422 4423 NEfiflS—AéyL MSFG MSFG MSFG MSFG MSFG Overal Mean & S. D. 4925 4261 427' 4255 429? 1 Distance Between Fibrovas. Bundles MEAhl 3.87 3.77 3.65 3.05 4.00 3.02 3.92 3.52 3.90 3.92 4.27 4.10 3.55 3.67 3.72 3.60 3.62 4.25 3.10 3.52 3.00 3.52 4.05 3.00 3.52 4.07 4.37 4.30 3.70 3.82 SOD. 1.197 1.097 1.039 .762 .903 1.330 1.054 1.141 1.220 1.227 .910 1.198 .841 .748 .751 .994 .915 1.381 1.020 1.081 1.866 .834 .930 1.229 1.381 1.127 1.653 1.239 1.140 11.148 Endoderm Length MEAN 38.87 39.38 38.10 36.90 39.95 40.70 42.15 38.50 42.30 38.75 33.90 36.40 36.65 36.60 37.30 38.25 37.35 37.20 36.10 36.55 38.15 36.00 41.90 39.60 39.75 38.80 42.85 39.05 38.30 38.62 500. 4.462 4.371 4.024 4.339 3.804 3.435 4.955 5.296 4.341 4.063 4.948 4.672 4.132 3.315 1.976 4.115 2.796 2.839 3.782 3.203 5.460 4.611 4.140 2.741 3.552 2.839 5.650 3.872 4.219 4.420 Endoderm Width MEAN 19.76 19.52 19.05 18.65 19.65 20.90 20.65 19.05 22.05 19.60 19.75 18.60 19.15 19.20 18.35 19.70 19.45 10.35 10.95 10.40 19.00 18.30 20.00 20.70 20.10 19.90 22.20 20.05 10.00 19.64 . - 500. 2.428 2.163 2.109 2.641 2.125 2.758 2.796 2.187 2.414 2.244 1.788 3.030 1.989 1.565 2.473 ‘1.276 2.058 2.012 1.875 1.919 1.719 2.513 1.080 1.372 2.531 3.088 2.855 2.238 2.402 Endoderm. Length/ Width ' Ratio MEAN 1.97 1.97 2.01 1.98 2.05 1.95 2.05 1.94 1.92 1.98 1.97 1.95 1.94 1.91 2.04 1.94 1.92 2.04 1.90 1.99 2.00 1.97 2.10 1.91 1.97 1.96 1.94 1.96 100-3 j_.97 SOD. .168 .192 .215 .150 .213 .154 .152 .139 .151 .192 .178 .232 .127 .204 5134 .194 ”216 .137 .182 .193 .216 .183 .122 .117 .179 .202 .207 .132 .180 Table A-35. 151 Plot means (fertilized and unfertilized) and standard deviations for the measurement of 27 black pine seedlots Nitrogen g MEAN 5.0. PERT. {1.5 .000 UNFmi 1.39 .077 MSFG 401 1.37 .014 MSFG 402 1.48 .077 MSFG 403 1.40 .056 MSFG 404 1.44 .028 MSFG 405 1039 0014 MSFG 407 1.60 .056 MSFG 408 1,45 .070 MSFG 410 1,49 .070 MSFG 411.1,31 .042 MSFG 41211.28 .028 MSFG 413 1,33 .042 MSFG 414 1,39 .007 MSFG 415 1.52 .106 MSFG 416 1.46 .035 MSFG 417 1.55 .014 MSFG 418 1.48 .056 MSFG 419 1.42 ],oa4 MSFG 420 1.50 .056 MSFG 421 1.40 .073 MSFG 422 1.36 .056! MSFG 423 1.52 9049' MSFG 424 1.37 .042- MSFG 425 1.39 .127: MSFG 426 1,42 .028? MSFG 427 1,42 .000 MSFG 428 1.48 .084 MSFG 429 1.3 .056 Overall Meang!‘ 1,42 0088 8.1L Potassium McAN 5.0. .580 .029 .642 ‘.036 .630 .028: .035 .077' .600 .0282 .625 .035: 0630 9070' .635 .021' .630 .086 .630 .070 .590‘ .084 .575 .063 .555 .035 .595 .063 .610 .042 .655 .021 .615 .049 .610 .014 .610 .014 .635 .049 0600 .028 .555 .035 .650 .042 585 .049 .625 .035 .640 .084 .630 .070 .610 .014 .611 .045 Phosphorus MEAN 5.0. .160 .0098 .165 .0106 .157 .0056 .169 .0120 .161 .0000 .161 .0113 0349 .0056 .173 .0056 .163 .0028 .169 .0000 .153 .0000 0149 .0056 .175 .0028 .973 .0000 .167 .0084 .177 .0113 177 .0000 .161 .0113 .167 .0028 .149 .0056 9173‘ .0056 .1571 .0169 .161 .0000 9365 .0056 0169‘ 00113 .157 .0056 .153 .0000 .362‘ .0104 Smfinm pun MEAN 64.5 75.7 115.5 96.5 81.0 87.0 56.5 72.0 51.5 36.0 40.0 58.0 54.0 69.0 87.0 68.5 05.0 121.0 127.0 76.5 50.5 43.0 43.5 48.0 58.0 52.0 79.5 67.5 69.0 70.1 SoD. 21.96 34.58 45.96 19.09 25.45 33.94 9.19 7.07 2.12 2.82 2.8? .00 5.65 1.41 21.21 2.12 4.24 55.15 70.71 4.94 17.67 15.55 6.36 8.48 .00 2.82 3.53 6.36 26.87 29.24 Table A-36. 152 Plot means (fertilized and unfertilized) and standard deviations for the measurement of 27 black pine seedlots F ERT . UNFER. MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MbFG MSFG 401 402 403 404 405 407 408 410 411 412 413 414 415 416 417 418 419 420 421 422 423 424 425 426 427 428 429 Overall Phan1& 8.1L Calcium % MEAN SoD. '329 0046 .327 .039 .310 .014 .395 .071 .375 0021 .330 .000 9360 .000 0330 .000 .340 .028 .360 .042 ~315 .063 .315 .021 .345 .021 9360 .000 9340 .014 9335 .071 .375 .021 0405 .021 0315 .063 0790 .000 0270 .000 .285 00?1 .330 .000 .285 .021 9?70 .000 Q?85 .021' .330. .042 o?70 .042 0328 .042‘ Mmymmimm MEAN 0106 .115 .115 .130 .110 .125 .095 .100 .135 .105 .075 .085 .080 .125 .120 .090 0130 .100 .110 .105 .100 .130 .145 .120 .130 .135 .105 .090 .110 9111 .0216 lhngumse PM“ MEAN 5.0. 338.2 45.57 266.8 44.03 247.5 21.92 296.0 14.14 995.0 69.29 338.0 49.49 263.0 24.04 288.5 12.02 260.0 24.04 336.0 96.16 992.0 73.53 266.0 67.88 304.5 113.84 322.0 14.14 265.0 22.62 228.5 ‘70.00 294.0 28.28 297.5 65.76 349.5 41.71 360.5 40.30 406.5 38.89 972.0 76.36 305.0 59.39 326.0 .00 317.0 4024 300.0 26.28 298.5 105.35 328.0 124.45 312.0 77.78 302.5 57.18 Inn1 ppm MEAN 5.0. 53.? 7.74 53.0 10.26 52.5 2.12 68.0 .00 58.5 2.12 51.0 12.72 49.5 2.12 40.0 '8.48 47.0 .00 45.0 4.24 43.5 2.12 45.0 .00 42.0 4.24 53.0 4.24 43.5 2.12 40.5 9.19 54.0 .00 64.5 2.12 61.5 19.09 66.5 06.36 6090 000 55.5 2.12 53.0 4.24 48.0 .00 60.0 4.24 4915 10060 54.0 8.48 63.0 8.48 58.5 6.36 53.1 9.00 Table A-37. pine seedlots FERT. UNFER. MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG 401 qumr ppm 9.58 9.05 40211.00 40310.75 40410.30 40513.90 40711.65 40811.00 410 411 412 413 414 9.00 8.55 7.35 7.25 7.95 41510.95 416 6.50 41717.50 41813.80 419 9.45 420 7.50 42110.30 422 9.00 423 3.40 42410.75 42510.75 426 9000 42711.65 42810.75 429 7.70 1.27 (Nenfll an1& 5.1L MEAN SOD. 2.15 10.54 3.37 .63 2.12 .63 1.27 5.09 .63 .28 .56 2.47 1.76 .63 .35 2.19 .00 6.50 1.27 2.47 .28 .000 1.83 .28 4.31 .63 3.11 .63 .63 10.06 2.84 153 Rnbn PW“ MEAN 16.22 19.62 17.90 19.95 18.50 20.65 15.25 13.15 15.00 14.70 13.65 11.70 14.70 16.90 18.75 16.70 17.90 20.10 20.70 18.90 17.95 20.10 19.60 20.15 22.95 17.90 19.65 22.30 18.45 17.92 5.00 2.866 3.279 1.555 5.020 3.518 2.333 .777 .777 1.538 .000 1.484 2.828 2.969 3.818 3.464 4.949 1.555 '.000 3.959 2.687 4.596 ”.000 1.555 4.737 4.030 '.000 5.444 .777 3.501 Plot means (fertilized and unfertilized) and standard deviations for the measurement of 27 black fine PW“ MEAN 5.0. 29.4 7038 37.0 8.84 41.5 4.94 39.5 3.53 49.5 6036 36.5 7.77 39.5 7077 29.5 2.12 48.5 10.60 37.5 2.1? 30.0 5.65 25.0 1041 27.0 1041 36.5 7.77 36.0 8.48 24.5 7.77 43.5 2.12 36.5 7077 32.5 2.12 28.5 4.94 22.5 2.12 22.6 4.94 34.5 9.19 30.0 12.72 24.5 4.94 23.5 3.53 31.5 10.60 39.5 7.77 26.0 2.8? 33.2 8.93 Xhmdnmn PPm MEAN 8.0. 289.0 63.97 353.6 65.48 287.0 24.04 361.5 45.96 311.5 45.96 428.5 55.86 246.0 46.66 309.0 80.61 320.0 49.49 258.5 19.09 354.5 4.94 279.5 48.79 381.5 43.13 353.0 171.11 323.5 7.77 191.5 17.67 224.0 67.88 319.0 66.46 359.5 53.03 403.5 109.60 321.0 107.48 348.5 68.58 350.0 8.48 263.0 52.32 308.5 60010 272.0 .00 374.0 32.52 412.0 11.31 315.0 60.81 321.3 71.93 Table A-38. and standard deviations for the measurement of 27 black pine seedlots Nitrogen- PhOSphorus Ihtio M5401 5.8. FEET. 9.09 .366 IHWER. 8.46 .392 MSFG 4015-73 040‘ MSFG 4028.79 1.092 MSFG 4038169 1351 MSFG 4048.96 .453 MSFG 4059.33 .449 MSFG 4079.24 .024 MSFG 4089.08 .591 MSFG 4108.81 .418 MSFG 411 3.58 .277 MSFG 4128.59 .136 MSFG 4135.69 .27 MSFG 4147.97 .189 MSFG 415 8.81 .613 MSFG 4168.78 .658 MSFG 4178.77 .480 MSFG 4188.36 .319 MSFG 4198.82 .092 MSFG 4208.98 .186 MSFG 4218.76 1.056 MSFG 4229.12 .033 MSFG 4238.82 .574 MSFG 4248.79 1.220 MSFG 4258.63 .790 MSFG 4268.61 .466 MSFG 4278.42 .563 MSFG 4289.42 .200 MSFG 4298.49 .369 Overall Meanéc 8.78 .493 8.1L Nitrogen- Pawnmimn Patio MEAN 2.51 2.18 2.17 2.36 2.33 2.30 2.22 2.52 2.36 2.38 2.24 2.23 2.40 2.35 2.51 2.23 2.52 2.42 2.33 2.37 2.33 2.45 2.35 2.35 2.52 2.27 2.23 2.37 2.13 SoUo .136 .114 .120 .412 .204 .085 .271 .173 .324 .380 .395 .198 .229 .264 .348 .126 .130 .149 .193 .274 .062 .258 .229 .271 .231 .174 .296 .400 .142 .210 Plot means (fertilized and unfertilized) Pbtmxflnm- rhomflmuus lhfiio MEAN 3.62 3.89 4.01 3.73 3.72 3.88 4.?2 3.67 3.86 3.72 3.85 3.85 3.62 3.39 3.52 3.92 3.47 3.44 3.80 3.80 3.75 3.73 3.75 3.73 3.41 3.78 3.77 4.02 3.98 3.76 8.0. .224 .261 .035 .193 .175 .053 .314 .242 .280 .418 .554 .280 .231 .308 .245 .072 .057 .079 .354 .330 .351 .378 .122 .088 .000 .084 .249 .595 .092 .276 $xfinm- Rfiwsshmm fhtio WCAN 8.0. 110.9 35.50 118.1 55.33 185.1 81.26 151.? 11.53 134.1 36.10 137.8 46.50 89.4 4.55 11302 7935 82.? 10.75 57.? 1.93 68.8 14.69 101.4 11.23 9708 16042 1160S 10008 141.7 24.91 104.6 6.62 13803 4023 197.3 85.84 90609 111.12 121.1 17.23 83.5 25.52 76.5 33.03 66.7 5.43 82.9 21.5? 105.4 .00 83.2 .18 124.9 11.04 108.3 22.26 112.6 41.43 114.5 46.19 155 Table A-39. Plot means (fertilized and unfertilized) and standard deviations for the measurement of 27 black pine seedlots Calcium- Manganese- Magnesium Iron Needle Ratio Ratio Color MEAN SOD. NEAN SOD. MEAN 5.0. FEET. 3.14 .575 6.45 1.137 7.6 3.73 UNFER. 2.98 .846 5.12 .895 7.9 3.52 MSFG 4012.73 .380 4.70 .227 6.0 2.82 MSFG 4023.09 .509 4.35 .207 10.0 1.41 MSFG 4033.40 .192 5.06 1.368 7.0 2.32 MSFG 4042.67 .454 6.96 2.708 6.0 4.24 MSFG 4053.80 .282 5.30 .258 8.0 4,2. MSFG 4073.30 .000 6.08 .824 8.0 1.41 MSFG 4082.53 .188 5.53 .511 12.0 4.24 MSFG 4103042 0173 7039 1.439 1200 1.41 MSFG 4114.25 1.250 6.67 1.364 12.0 1.41 MSFG 4123.72 .559 5.91 1.508 13.0 .00 MSFG 4134.31 .265 7.14 1.988 12.0 1.41 MSFG 4143.00 .848 6.10 .755 10.5 2.1? MSFG 4152.83 .117 6011 .313 410 .00 MSFG 4163072 0235 5098 .‘59 1300 2.8? MSFG 417 2.97 .809 5.44 .523 8.0 1.41 MSFG 4184.10 .792 4.59 .868 2.0 1.41 MSFG 4192.86 .578 6.08 2.566 7.0 .00 MSFG 4202.76 .186 5.47 1.130 10.0 .00 MSFG 4212.72 .385 6.77 .648 9.0 .00 MSFG 4222.?1 9404 4993 1.564 300 2.82 MSFG 4237.43 .795 5072 .662 465 3.53 MSFG 4242.75 .000 6.79 .000 6.0 1.41 MSFG 4252.22 .321 5.29 .445 2.0 1,4. MSFG 4262.02 .318 6.14 .744 8.0 1.41 MSFG 4272.71 .019 5.75 2.854 5.0 2.32 MSFG 4283.66 .471 5.11 1.285 6.0 1.41 MSFG 4292.45 .070 5.29 .753 7.0 .00 Overall 8. D- Table A-uo. Pine seedlots NEE. UNNHL MSFG MSFG MSFG MSFG MSFGu415- MSFG MSFG MSFG 401 402 408 414 416 420 423 Ovemfll an1& 8.1L Efifl. UNFHL MSEG_404 MSFG MSFG MSFG MSFG- MSFG 402 408 414 415 416 MSEG_420 MSFG 423 Ownmll rbanét S. D. Nitrogen % 1.37 1.50 1.48 1.39 1.52 1.46 1.50 1.52 1.47 SoD. .102 .094 .095 .108 .084 .064 .139 .151 .048 .066 .104 thflnm MEAN .337 .335 .307 .390 .337 .360 .337 .330 .285 .345 .336 8.0. .064 .048 .015 .042 .066 .024 .015 .054 .017 .110 .055 156 Potassium % MEAN .594 .661 .630 .632 .627 .595 060% .650 .632! 800‘ .035 .046 .024 .101 .045 .071 .053 .024 .056 .650' 0034 .628 .053 lkgmxfium MEAN .11 .12 .11 913 .13 .12 .11 .09 .10 .14 .11 5.0. .027 .028 .032 .023 .020 .052 .005 .011 .005 .017 .027 Plot means (fertilized and unfertilized) and standard deviations for the measurement of 8 black Phosphorus MEAN .165 .171 .157 .169 .163 .175 .173 .167 .167 .1731 .3,68 S000 .011 .011 .008 .016 .007 .004 .013 .017 .008. .011 1%ngmxme PW“ NEAN 309.00 261.75 247.50 295.75 260.00 321.75 264.75 228.00 360.50‘ 304175 285.37 SOD. 58.23 50038 31.20 29.03 25.42 39.25 50015 70.80 51.46 61.31 58.69 fixfimm PW“ MEAN 5.0- 76.37 75.12 44.480 24.746 79.483 23.902 10.874 11.044 17.492 9.949 8.981 115.25 96.00 51.25 68.75 87.00 68.50 76.00 43.25 75.75 13.793 35.412 Innl MEAN 52.87 52.37 52.50 67.50 46.50 52.50 43.50 40.00 66.00 52.50 PP“ 8.0. 9.666 12.706 9.000 0.244 1.732 7.141 3.872 8.602 8.831 5.190 52.62 11.108 TabRBAflnlo Nfifl. U NFER. MSFG MSFG MSFG MSFG MSFG MSFG MSFG MSFG 401 1402 440E} 414 4415 4516 42H) 42K3 Ownnll Phan.& 5.1L UNNHL MSFG MSFG MSFG MSFG MSFG 415 MSFG 416 MSFGF420 MSFG 423 401 402 408 414 Ovemfll Phanék 8.11 157 Plot means (fertilized and unfertilized) and standard deviations for the measurement of 8 black pine seedlots Cepper PP“ MEAN 8.90 9.13 9.00 10.97 10.97 7.92 10.95 6.45 7.47 8.37 9.01 5.0. 2.051 2.066 I 1.110 1.920 1.920 .450 1.819 .404 .861 .450 2.028 Nitrogen- Phosphorus Ratio MEAN 9.17 8.37 8.72 8.95 9.09 7.97 8.81 8.80 8.99 8.82 8.77 50D. .597 .435 .469 1.056 .576 .297 .505 .931 .396 .511 .653 Ebrm1 ppm MEAN 15.71 20.12 17.90 19.90 1‘Q97 10.87 18.75 16.05 18.77 19.55 17.92 5.0. 2.372 2.830 1.555 4.729 1.800 3.147 2.996 5.041 4.261 1.905 3.407 Nitrogen- Potassium Ratio MEAN 5.0. 2.55 2.17 2.17 2.42 2.37 2.37 2.53 2.25 2.39 2.35 2.36 .247 0182 .206 .435 .262 .319 .389 .279 .285 .203 .288 Zfinc PW“ MEAN 31.00 41.06 41.50 39.25 48.25 36.50 35.75 24.25 28.50 34.25 36.!‘3 5.0. 7.702 8.028 3.774 10.210 6.806 7.320 6.184 5.259 7.320 9.275 Potassium- Phosphorus Ratio MEAN 3.62 3.87 4.01 3.72 3.85 3.39 3.52 3.92 3.80 3.75 3.75 5.0. .385 .259 .225 .246 .278 .378 .369 .385 .462 .176 .347 Almmbmm MEAN 286.68 ppm 5.0. 94.182 360.56 104.120 986.75 41.843 301.25 109.500 320.00 09.123 353.00 150.956 323.50 09.004 191.25 91.620 403.25 134.097 350.00 55.238 323.62 104.623 Sodium- Potassium Ratio MEAN SOD. 329.07 74.781 114.71 41.687 185.91 135.891 154.87 46.641 62.10 18.996 116041 229983 142.58 20.726 105.32 140419 121.58 23.270 766.37 19.444 121.89 60.000 Table A-42. Efiw. UNFER. MSFG MSFG MSFG MSFG MSFG— MSFG MSFG MSFG 401 1402 4Cfl3 414’ 4455 1416 420 1423 Ovemfll Mean & 5.1L 158 Plot means (fertilized and unfertilized) and standard deviations for the measurement of 8 black pine seedlots Calcium- Magnesium Ratio MEAN 3.05 2.84 2.87 3.05 2.56 3.22 2.94 3.71 2.78 2.44 2.94 5.0. .585 .734 .684 .534 .438 .898 .249 .748 .146 .815 .662 Manganese- Iron Ratio MEAN 5.0 5.91 1.003 5.08 .633 4078 .731 4.41 .670 5.58 .445 6.17 .844 6.07 .898 5.61 .593 5.52 .935 5.82 1.231 5.50 .928 Ne edle Qflor MEAN 7.75 0975 0.00 10.00 12.00 10.50 4.00 13.00 10.00 4.50 8.75 SOD. 4.374 3.492 2.581 2.000 3.404 1.914 1.154 3.205' 2.581‘ 3.4151 4.024