" 37%;; A COMPARATIVE AUTECOLOGICAL STUDY OF BLACK SPRUCE. JACK PIN£ LODGEPOLE PINE. AND HYBRID PINE FOR DIRECT sesame IN NORTHERN ONTARIO Thesis for the Degree of M. S. MICHIGAN STATE UNIVERSITY DAVID ARTHUR WINSTON 1973 “3'5?ng APR 3. 0 23:1 ABSTRACT A COMPARATIVE AUTECOLOGICAL STUDY OF BLOCK SPRUCE, JACK PINE, LODGEPOLE PINE, AND HYBRID PINE FOR DIRECT SEEDING IN NORTHERN ONTARIO BY David Arthur Winston A series of laboratory and field experiments were performed to compare germination, survival and growth of jack pine (Pinub banhbiana Lamb.), black spruce (Picea maniana (Mill.) B.S.P.), lodgepole pine (Pinub contonta var. Latifiolia) and a hybrid pine (P. contnnzu var. zatiéoiia x P. bankbiana) under a variety of soil moisture, soil fertility and field site conditions. In a study to determine the cardinal temperatures for germina- tion, jack pine seed germinated rapidly over a wide temperature range, from 15.6 C (60 F) to 32.2 C (90 F). Lodgepole pine and hybrid pine were more sensitive to temperature. Optimum germination of lodgepole pine was restricted to the range of 21.1 C (70 F) to 26.7 C (80 F). Two greenhouse pot studies compared the growth of seedlings of the three species and hybrid pine in two soil types at controlled soil moisture and soil fertility levels. The two soils were obtained from an experimental field location near Manitouwadge, Ontario, which covered two sites, a moist upland black spruce site, and a dry upland jack pine site. Black spruce seedlings grew best at soil moisture levels near 1/3 atmosphere suction, but they did survive and grow at 10 atmospheres soil moisture suction. A11 pines produced greater top and root weight than black spruce at all moisture levels. Both jack pine and lodgepole pine appear better suited for growth on low moisture David Arthur Winston content soils than either black spruce or hybrid pine. Increased growth of black spruce was obtained on soil from the moist site at fertility levels increased to those prescribed by Wilde (1958) as Grade B. Optimum growth of pines was obtained at a lower fertility level than black spruce. Seed germination and seedling survival of the three species plus hybrid pine were evaluated in field seeding trials at Manitouwadge in 1971 and 1972. Black spruce seed germinated in both years and on both moist and dry site conditions. Stocking of black spruce seedlings was poor in the 1971 trial but attained a high (75 percent) level in the 1972 seeding after the first growing season. Soil moisture levels near field capacity, and careful seedbed preparation are the two major factors affecting black spruce seedling survival. Jack pine and lodgepole pine seed germinated well in both seeding trials, with stocking exceeding 80 percent. Neither soil moisture nor seedbed preparation were deemed as critical for these two species as for black spruce. Hybrid pine seed germination was low in both 1971 and 1972 seeding trials, but, germination of those seeded in 1971 increased in 1972. Jack pine seed germinated sooner than the other species in the 1972 trial, during a period of high soil surface temperature and this is partly due to the differences in species germination response to temperature. Screening of seedspots, to determine if rodents were present and destroying seed on the cutover, were inconclusive. A COMPARATIVE AUTECOLOGICAL STUDY OF BLACK SPRUCE, JACK PINE, LODGEPOLE PINE, AND HYBRID PINE FOR DIRECT SEEDING IN NORTHERN ONTARIO By David Arthur Winston A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE College of Agriculture and Natural Resources Department of Forestry 1973 VITA David Arthur Winston Candidate for the degree of Master of Science Biographical Note Born: Stockport, Cheshire, England, September 13, 1946. Married, wife, Helen; daughter, Elaine Education: University of Toronto, B.Sc.F., 1968 Michigan State University, M.S., 1973 Employment: Present: Research Officer, Canadian Forestry Service, Canada Department of Environment, Great Lakes Forest Research Centre, Sault Ste. Marie, Ontario. April 1968 - April 1969: Land Use Planner, Ontario Department of Lands and Forests, Recreation Inventory Unit, Richmond Hill, Ontario. Organizations: Canadian Institute of Forestry Canadian Pulp and Paper Association Xi Sigma Pi Botanical Society of Sault Ste. Marie, Ontario 1i ACKNOWLEDGEMENTS The author wishes to express his gratitude to Dr. G. Schneider for the many hours of discussion and work he has contributed. The author is indebted to Drs. D. P. White, J. W. Hanover, and R. J. Kunze for their advice and criticism while acting as members of the Guidance Committee. The support of the Canadian Forestry Service, Great Lakes Forest Research Centre, Sault Ste. Marie, Ontario, is gratefully acknowledged. The author wishes to especially thank Mr. J. W. Fraser, Research Scientist, for the advice, guidance and assistance that he has so generously given. The author wishes to dedicate this dissertation to his wife, Helen, who has struggled through with spirit and determination, while spending countless hours working on the research and thesis preparation. iii TABLE OF CONTENTS TABLE OF CONTENTS LIST OF TABLES LIST OF FIGURES CHAPTER I. II. III. INTRODUCTION REVIEW OF LITERATURE l. JACK PINE 2. BLACK SPRUCE 3. EXOTIC SPECIES 4. SEED DEPREDATION METHODS 1. ORIGIN AND COLLECTION OF SEED FOR USE IN EXPERIMENTS 2. CARDINAL TEMPERATURES FOR GERMINATION 3. EXPERIMENTAL FIELD LOCATIONS A. General B. Soil-Site Description and Analytical Methods 4. GREENHOUSE POT TRIALS A. General B. Soil Moisture Levels C. Soil Fertility Levels iv Page iv vii ll 13 15 15 16 16 16 17 18 18 19 20 CHAPTER III (continued) Page 5. EXPERIMENTAL FIELD SEEDING IN 1971 AND 1972 MANITOUWADGE, ONTARIO 21 A. 1971 Seeding 21 B. 1972 Seeding 24 IV. RESULTS AND DISCUSSION OF EXPERIMENTS 28 1. SEED ORIGIN AND COLLECTION 28 2. CARDINAL TEMPERATURES FOR GERMINATION 28 3. SOILS AND VEGETATION AT FIELD LOCATIONS 32 4. GREENHOUSE POT TRIALS 37 A. Soil Moisture Levels 37 1. Height growth: 37 2. Length of the longest root: 38 3. pr weight: 38 4. Total root weight: 38 5. Chemical nutrient analysis of tops and roots: 40 B. Discussion of Soil Moisture Experiment Results 40 C. Soil Fertility Levels 41 1. Height growth: 41 2. Mean length of longest root: 43 3. Top weight: 43 4. Total root weight: 43 5. Chemical nutrient analysis of tops and roots: 45 D. Discussion of Soil Fertility Experiment Results 45 CHAPTER IV (continued) Page E. Practical Importance of Soil MOisture and Soil Fertility Pot Trials 47 5. RESULTS AND DISCUSSION OF 1971 AND 1972 FIELD SEEDING EXPERIMENTS 49 A. Results and Discussions of 1971 Seeding 49 B. Results and Discussions of 1972 Seeding 54 V. SILVICULTURAL IMPLICATIONS OF RESEARCH RESULTS 70 LITERATURE CITED 72 APPENDIX A 77 APPENDIX B 78 APPENDIX C 82 APPENDIX D 88 APPENDIX E 94 vi Table 10 ll 12 13 LIST OF TABLES Mean percent germination of jack pine, lodgepole pine, hybrid pine and black spruce at different temperatures and days after seeding Detailed soil pedon description for Manitouwadge moist and dry sites Soil horizon chemical analysis 1971 seeding area Soil physical and chemical properties for moist and dry site bulk samples Soil moisture content (percent by weight) for moist and dry site bulk samples Soil nutrient analysis and nutrient amendments required to increase soil fertility levels to treatment standards Weekly rainfall and air temperatures measured at Manitouwadge weather office, and field soil surface temperatures for July - September, 1971 Soil and air temperatures, and weekly rainfall totals measured on the 1972 field sites The effect of screening on germination of the three spec- ies and hybrid pine seeded on both moist and dry sites in 1972 Test of significance based on Duncan's Multiple Range Test for germination by treatment and assessment date Thirty-year climatic normals for Canada Department of Transport weather stations near Manitouwadge Growth measurements for seedlings extracted in August, 1972 from both the 1971 and 1972 seeding trials Details of seed origin and initial germination tests vii Page 29 33 34 35 35 42 50 56 59 61 65 69 77 Table 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 3O Cardinal temperature study: 7-day germination percent by species and population Cardinal temperature study: 14—day germination percent by species and population Cardinal temperature study: 21-day germination percent by species and population Cardinal temperature study: 28-day germination percent by species and population Effects of soil moisture levels pole pine seedling growth Effects of soil moisture levels pine seedling growth Effects of soil moisture levels pine seedling growth Effects of soil moisture levels spruce seedling growth Effects of soil moisture levels foliage nutrient concentrations Effects of soil moisture levels nutrient concentrations and and and and and and site site site site site site on lodge- on hybrid on jack on black on seedling on seedling Effects of soil fertility levels and site on lodgepole pine seedling growth Effects of soil fertility levels hybrid pine seedling growth Effects of soil fertility levels pine seedling growth Effects of soil fertility levels black spruce seedling growth Effects of soil fertility levels foliage nutrient concentrations Effects of soil fertility levels root nutrient concentrations and and and and and site site site site site on on jack on on seedling on seedling Soil surface temperature and moisture levels at the 1972 seeding location viii Page 78 79 80 81 82 83 84 85 86 87 88 89 9O 91 92 93 94 Figure 10 11 12 LIST OF FIGURES Map of the Boreal Forest Region in Ontario General view of dry site cutover near Manitouwadge, Ontario Bracke scarifier-seeder pulled by Timberjack wheeled skidder, preparing seed spots on dry site skid road and cutover Schematic presentation of plot layout for 1971 seed- ing location Example of independent randomly paired plot layout, 1972 Mean seed germination for three species plus hybrid pine at several temperatures, 7 and 28 days after seeding Soil moisture curves for bulk soil samples obtained from Manitouwadge, Ontario Seedling height growth and by length of longest root at three moisture regimes for two sites Seedling height growth and length of longest root at three fertility levels for two sites Germination, stocking and mortality for three species and hybrid pine seeded in 25 scalps on the moist site in 1971 Germination, stocking and mortality for three species and hybrid pine seeded in 25 scalps on the dry site in 1971 Mean soil and air temperatures, and total rainfall, by weekly periods for 1972 field seeding location ix Page 23 27 3O 36 39 44 51 52 55 Figure 13 14 15 16 Stocking, germination and mortality of the three species and hybrid pine in 25 scalps in the 1972 moist site Stocking, germination and mortality of the three species and hybrid pine in 25 scalps in the 1972 dry site Seedlings from the 1971 and 1972 dry site field seeding trials Black spruce seedlings from the dry site field seeding trials Page 57 58 66 66 CHAPTER I INTRODUCTION The technique of direct seeding for forest regeneration is currently undergoing renewed interest and application throughout North America. Although direct seeding is not a new technique, and indeed is Nature's method of regeneration, planting of seedlings has been practised by forest managers, to the neglect of seeding. Recently, several reports have been published, indicating a return to direct seeding. Scott (1970), Smithers (1965), McQuilkin (1965), Derr and Mann (1971) have reported on seeding trials and programs established in their respective areas. Rising interest in seeding can be attributed to several economic factors. Most notable are the high cost of planting and the increased number of acres that are harvested annually with the advent of mechanical harvesting equipment. This requires a cheaper and faster method of regenerating the increased acreage. Scott (1970) reports that seeding is now the recommended method for regenerating jack pine (Pinus banksiana Lamb.) in Ontario. However, even for this species, direct seeding sometimes results in unexplained regeneration failures. Seeding trials with black spruce (Picea mariana (Mill.) B.S.P.) in Ontario have usually been failures. Successful seedings of black spruce have been reported elsewhere, by van Nostrand 1) (1971) in Newfoundland, Johnston (1971a) in Minnesota, and Marek in 1) Personal communication, Mr. G. Marek, Ontario Ministry of Natural Resources, Beardmore, Ontario. 2 Ontario following modified types of clear—cut and intensive site preparation. Presently in northern Ontario, the standard harvesting technique is to clear-cut large areas, necessitating the planting of seedlings in order to obtain satisfactory regeneration. Financial resources of the Provincial Government do not permit site treatment and the planting of desired species on every acre harvested. This results in an ever-increasing back-log of non-regenerated forest land. The ability to obtain successful regeneration of black spruce and jack pine by direct seeding would provide a partial solution to Ontario's forest regeneration problems. This study provides information on environmental and climatic factors which favor the germination and survival of black spruce and jack pine on clear-cut areas in the boreal forest of Ontario. It also evaluates the potential use of lodgepole pine (Pinus contorta var. Zatifblia Dougl.) and a hybrid pine, jack pine x lodgepole pine (P. banksiana x P. contorta var. Zatifblia) as possible substitutes for seeding on black spruce and jack pine sites in northern Ontario. CHAPTER II REVIEW OF LITERATURE The effects on local environments of large clear-cut areas are well referenced. Johnson et al. (1971) summarize some of the most important ones, which, depending on the particular circumstances may be either beneficial or detrimental to regeneration or alternative land USES: 10. 11. Abrupt increase in incident light which encourages proliferation of ground vegetation. Increase in sunlight reaching soil surface layers. Increase in soil surface temperatures depending upon the color and texture, often to levels lethal for germinating seedlings, and also contributing to drying effects. Increase in soil temperatures below the surface by as much as 5 C (9 F). Increase in wind speeds and turbulence. Reduction in atmospheric humidity. Reduction in minimum temperatures and frequently an increase in maxima. . Increased incidence of frost. Penetration of frost more deeply into soil layers, with an increase in frost heaving and soil detachment. MOre rapid snow melt. More violent impact of rainfall and, until 3 4 vegetation becomes established, more rapid surface run-off. 12. Frequent increases in soil moisture and occasional water-logging. l3. Accelerated oxidation and breakdown of organic layers. A large portion of Ontario is covered by the Boreal Forest Type (Figure 1). This consists of extensive stands of jack pine, black spruce, white spruce (Picea gZauca (Moench) Voss), balsam fir (Abies balsamea) (L.) Mill), white birch (Betula papyrifera Marsh.) and poplar (Populus spp.) as single species or mixed species stands (Rowe, 1959). Broad site classification systems have attempted to delineate sites most favorable to the growth of several species but little investigation has been done on a microsite basis (Hills, 1959; Chrosciewicz, 1963). In current direct-seeding practices in Ontario, jack pine is seeded on a variety of sites, from dry to moist, including many from which a black spruce crop has been harvested. Thus, future stands will be predominantly jack pine. The development of the pulp and paper industry in Ontario has been and still is based mainly on a supply of black spruce fibre. If the supply of black spruce is seriously reduced, it is conceivable that a multi~million dollar industry will also suffer, since fibre quality is the major factor in the competitive edge that the Ontario industry still retains. Thus, attempts to obtain successful seeding of black spruce as well as jack pine are very important to Ontario's economic future. l. JACK PINE In his summary of direct seeding projects in Ontario, Scott (1970) indicates that successful seeding of jack pine has been consistently obtained on a wide range of sandy sites when seedbeds were prepared. Unexplained failures, however, have occurred and the optimum site requirements for successful germination such as seedbed condition, and degree and extent of scarification, have not been determined. Other factors which have been studied to a limited extent for their effects on seeding success are: soil and air temperature, precipitation, season of seeding, frost, rodents, vegetation, soil moisture and nutrition. Jack pine grows primarily on a variety of soils of the Spodosol order (Podzols). It is capable of growing on very dry coarse, and medium sands and on gravelly soils, but it also occurs on fine sands, sandy loams, loams, clay-loans, and on relatively thin soils over rock outcrop. Only rarely does it occur on poorly drained soils. Generally, maximum growth occurs on moist upland till slopes that vary in texture from fine sand to clay (Cayford et al., 1967). Jack pine is a pioneer species with high light requirements (Logan, 1966). Fire promotes the establishment of jack pine by remov- ing vegetation, exposing mineral soil and causing its serotinous cones to open and release their seed. It frequently occurs as a mono-species and stands of extensive size grow on dry outwash sand plains. Burning, to shmulate wildfire effects on seedbeds, requires expert knowledge about the influence of weather on fire behavior in many types and conditions of slash, and experience in manipulating fire to achieve desired results. Prescribed burning has been used 7 successfully elsewhere (Chrosciewicz, 1970; Beaufait, 1962) as a method of seedbed preparation, but has not yet received widespread approval in Ontario. The common method of site preparation used in Ontario is mechanical scarification to expose mineral soil seedbeds or planting spots, but what constitutes the best seedbed remains controversial. Studies by Cayford (1958, 1963) and by Sims (1970) indicate that a mineral soil seedbed, prepared either mechanically or by burning, is generally very favorable for germination, whereas undisturbed litter and humus are very poor seedbeds. An unpublished study by Waldronz) showed germination and early survival to be similar, in years of above-normal precipitation, on three different seedbed types (mineral soil, mineral soil mixed with humus, and humus). Beaufait (1960) found poor growth of young jack pine seedlings on micro-sites where the upper soil horizons had been removed, possibly due to a reduction in soil moisture-holding and nutritive levels. He noted that bare sand was cooler than burned-over or non-disturbed soil surfaces and this may be advantageous in preventing seedling dessica- tion due to high temperatures. Studies by Fraser and Farrar (1953) indicated that conditions favoring a high~moisture content surrounding the seed, such as a fine textured seedbed, watering, shading, and sowing beneath the surface, resulted in good germination, whereas, conditions favoring drought resulted in lower germination. Partial shade, provided by slash or snags can help considerably in creating moisture conditions favorable for germination. 2) Noted in Cayford et al., 1967, p. 8. 8 Sims (1970) and Chrosciewicz (1960) also found the best germination on moist sites, with germination decreasing as the soil moisture regime decreased. \ Miller and Schneider (1971) have reported that partial shade and wind protection greatly increased germination of jack pine seed. Survival was improved significantly by shading but not by high soil moisture content nor by removal of vegetation competition. All of the latter treatments did, however, increase seedling growth. Fraser (1970a) and Ackerman and Farrar (1965) studied temperature requirements for germination of jack pine seed in the laboratory. Fraser obtained maximum germination in only seven days at temperatures from 21.1 C (70 F) to 32.2 C (90 F), and after 28 days, had similar germination from 15.6 C (60 F) to 35.0 C (95 F). Ackerman and Farrar found no difference in germination at 15.6 C (60 F), 21.1 C (70 F) and 26.7 C (80 F) under continuous light conditions. Fraser obtained little or no germination at temperatures below 10.0 C (50 F) or above 37.8 C (100 F) after 28 days. Generally, jack pine is a pioneer species capable of establish- ing itself on a wide variety of sites including those with low nutrient and moisture conditions. It requires adequate moisture and temperature for germination and survival. Failures of seeding have been attributed to poor seedbed preparation, climatic extremes, seed depredation by birds and rodents, frost, competition fromdvegetation, insects and disease. Additional research is needed to examine seeding, both natural and artificial, under combinations of seedbed preparations and various nutritional values, and to determine the range of site condi— tions on which jack pine can germinate and survive. 2. BLACK SPRUCE Traditionally black spruce has been regarded as a swamp and peatland forest species, and in Ontario most regeneration research has been carried out on such areas. However, it also grows in extensive pure and mixed stands, with a shorter rotation period, on well-drained upland sites. Delineation of sites suitable for black spruce regenera— tion has been inadequate as has the determination of the environmental requirements needed to achieve successful direct seeding. Linteau (1955), studying black spruce in Quebec, found that it grew best on a moist loam or sandy loam soil. It is commonly found growing on shallow or deep sands, or gravel tills, and occasionally on moss caps over bedrock (Lafond, 1958). Black spruce is sparse or absent on very dry sites, but frequently succeeds jack pine on these sites, particularly if a black spruce seed source is available. Vincent (1965) concluded that most existing stands are of fire origin. As previously mentioned, although successful seeding of black spruce has not been obtained in Ontario, it has in other areas of North America where harvesting techniques differ. Johnston (1971b) recommended burning of slash within a year after harvesting, followed by seeding of 250 M seeds per hectare (100 M seeds per acre) to obtain adequate stocking. On upland sites, he stressed that burning should be severe enough to expose mineral soil. Richardson (1970) obtained adequate stocking on burned, cut— over, moist to very moist upland sites. Stocking was reduced on wet sites. The organic mantle was relatively thin (average depth 3.7 cm, 1.5 inches) providing a moist but not excessively wet seedbed with temperatures favorable for germination. He felt that because of the 10 thin humus layer, seedling root systems were able to become established in or just above the mineral soil before the organic horizon dried out. He suggested that seeding must be done shortly after burning, as the humus material probably becomes dry and matted and unsuitable for 4“” retaining adequate moisture for germination. A cool and moist climate was considered more favorable for black spruce germination and survival than either a hot and dry, or a cold and moist one. Several researchers have examined the effects of seedbed condi— tion on germination and concluded that a seedbed's ability to retain moisture was most important (Place, 1955; Linteau, 1957; LeBarron, 1944). Vincent (1965) rated seedbed receptivity in order of priority as: 1) beds of slow-growing Sphagnum mosses; 2) moist rotten wood; <9” 3) mineral soil with a light cover of Polytrichum; 4) moist bare mineral soil; 5) litter; 6) beds of fast-growing Sphagnum; 7) feather mosses, living and dead. He cautioned, however, that each may have disadvantages. For example, Sphagnum moss growth may engulf seedlings; litter, and rotten wood often dry out leaving roots without adequate moisture supply. LeBarron (1944) found least seedling mortality due to heat on mineral and scarified soil, and most mortality on burned duff. However, growth was better on the duff, possibly due to a nitrogen deficiency where organic matter was absent. Place (1955) found that burned organic surfaces became very hot in sunlight, and this could cause a complete failure of germination. LeBarron (1944) found that shade cast by slash is beneficial to germination and survival in dry seasons. Logan (1969) found optimum seedling growth at light levels above 45 percent of available sunlight, and extremely reduced growth at 13 percent light, indicating that 11 daily exposure to full sunlight for even a short period is desirable for young germinates, since healthier, more developed seedlings are better able to withstand environmental stresses. Fraser (1970b), in studies of cardinal germination temperatures for black spruce, found that seed provenance had a significant effect on germination at different temperatures, and that optimum temperatures for maximum germination differed by provenance. Optimum temperatures for seed from Ontario site regions 5E, 4E and 3E (Ontario, 1963) were 10.0 C (50 F) to 26.7 C (80 F), 15.6 C (60 F) to 21.1 C (70 F) and 15.6 C (60 F) to 26.7 C (80 F), respectively. Germination did not occur below 7.2 C (45 F) or above 35.0 C (95 F). The best application of existing knowledge could improve the results of current black spruce seeding operations. However, additional research is necessary to determine Optimum site requirements for germination and survival. Accurate climatic prediction would also permit timing of seeding Operations to take advantage of favorable conditions. 3. EXOTIC SPECIES The introduction of exotic tree species to several areas of the world, most notably Australia, has afforded these areas a rapidly grow— ing timber crop. The slow growth, long rotation period, and difficulty in obtaining regeneration of native northern Ontario conifer species makes attractive the possibility of introducing exotic species to Ontario. Lodgepole pine and the hybrid pine are two conifers that may be satisfactory alternatives to jack pine and black spruce. Jack pine and lodgepole pine are both species of the Boreal 12 Forest whose respective ranges overlap in several locations in Alberta. Considerable controversy still continues about the origin of the two species since they are similar in some physical characteristics and hybridize naturally (MOss, 1949). The two species either developed independently or one species is a diverging glacial refugia of the other species (Yeatman, 1967). Mirov (1956) and Critchfield (1957) have found that the major positive means of distinction is a biochem— ical difference; jack pine consists ofIO<'- and B-pinene, while lodge— pole pine is mostly B-phellandrene. Lodgepole pine (84 MI- 352 M clean seed/kg) has a slightly larger seed than jack pine (156 M - 551 M clean seed/kg) (United States Department of Agriculture, 1948). By comparison, black spruce seed is minute with 738 M - 1123 M clean seed/kg. The available literature contains little information comparing lodgepole pine and jack pine growth, yield or site requirements. A personal assessment by the author indicates that although lodgepole pine is capable of growing on most sites occupied by jack pine and black spruce, it does grow better than jack pine on moist sites. Some winter injury of lodgepole pine was observed by the author in frost pockets at Longlac, Ontario, and by MacHattie (1956) in Alberta. Smithers (1961) summarized much of the available literature on lodgepole pine silvics, ecology and silviculture. Like other conifer species, lodgepole pine seed requires adequate moisture and moderate temperature for germination and survival. Ackerman (1955) found that mineral soil exposure by scarification was necessary to obtain regeneration of lodgepole pine. This is similar to the requirements of jack pine as previously described. 13 Moss (1949) described the locations and stands of naturally occurring hybrid pines in Alberta where jack pine and lodgepole pine inter-mix. The hybrids exhibit a wide range of characteristics in stem, foliage, cones, etc., that are commonly identified with one or the other parent. Although the possibility of superior hybrid progeny is always present, little investigation has been done to resolve it. Yeatman and Teich (1969) discuss some of the implications and import- ance of provenance selection for screening species and hybrids. They also mention the few breeding studies that have been initiated in Canada to investigate natural and artificial hybrids (jack pine x lodgepole pine). Yeatman and Holst (1972) have reported on the winter hardiness of natural and artificial hybrids, and jack pine provenances. The natural hybrid from Alberta crossed to Petawawa (Ontario) jack pine was found to have suffered little winter damage and compared favorably with local Ontario jack pine provenances. In a Lakes States study, Anderson and Anderson (1965) found lodgepole pine and hybrid pine to be more susceptible than jack pine to sweet fern blister rust. The evidence for and against seeding trials using lodgepole pine and hybrid pine is inconclusive, and I feel that further research is warranted to determine their potential as substitutes for jack pine and black spruce. 4. SEED DEPREDATION Rodent and bird repellents were developed for use in direct seeding and were employed extensively in the southern United States (Mann, 1968). The most successful of these was a combination of arasan, endrin and aluminium with a latex sticker. This combination was 14 adopted for operational use in Ontario, but it has not been clearly established that rodents or birds present a threat to direct seeding in this province. Recent investigations by Radvanyi (1970) and Crouch and Radwan (1971, 1972) indicate that repellent treatments can reduce germination and may be more harmful than the rodent and bird depredation. Future research on seeding in Ontario should include determination of the extent and degree of rodent damage. CHAPTER III METHODS 1. ORIGIN AND COLLECTION OF SEED FOR USE IN EXPERIMENTS Jack pine, lodgepole pine, hybrid pine and black spruce seeds were obtained in 1970-1971 from provenances within the 2000 to 25000 growing-degree—day range (Boughner and Kendall, 1959). Lodgepole pine and hybrid pine cones were collected in June, 1970 from individual trees at Marlborough and Fox Creek, Alberta.3) Jack pine cones were collected in August, 1970 from two stands in Nimitz Township, Chapleau Forest District, Ontario. Cones from the two species plus hybrid pine were shipped to Petawawa Forest Experiment Station for seed extraction, cleaning and basic germination tests.4) Black spruce seed, obtained in May, 1971 from the Ontario Department of Lands and Forests office at Manitouwadge, Ontario, had been extracted, cleaned and tested for germination at their Tree Seed Plant at Angus, Ontario, in 1970. Based on initial germination tests at Petawawa, seeds from lodgepole pine tree #1, and jack pine stand #1 were selected for all future field and greenhouse tests, except for the experiment to determine germination at several temperatures. See Appendix A for details of seed origin and initial germination trials. Seeds used in this study were not treated with repellents. 3) Courtesy, Mr. R. Ackerman, Northern Forest Research Centre, Edmonton, Alberta. 4) Courtesy, Mr. B.S.P. wang, P.F.E.S., Chalk River, Ontario. 15 l6 2. CARDINAL TEMPERATURES FOR GERMINATION Three lodgepole pine populations, one hybrid pine population and two jack pine populations were tested to determine their optimum temperatures for germination, and the germination response at several controlled temperatures. Seeds were randomly selected in lOO-seed batches from individual seedlots, treated by flotation to eliminate empty seed, pre-treated to break possible dormancy by cold soaking in dilute H2804 for 36 hours at 2 C (35 F), surface sterilized to inhibit mould, and seeded on double layers of saturated germination paper in sterile petri dishes. The procedures used followed the format described by Fraser (19703). Dishes for each species were allocated by random methods to constant temperature cabinets set at temperatures from 7.2 C (45 F) to 35.0 C (95 F) in 2.7 C (5 F) intervals and to shelves within these cabinets. Seeds received constant low level illumination. Germination was tallied daily for 28 days and seeds were tallied as germinates when radicles reached 2 mm (.08 inches) in length. Species and temperature differences were analyzed by analysis of variance and Duncan's New Multiple Range techniques. 3. EXPERIMENTAL FIELD LOCATIONS A. General Field germination and survival of the four SGeletS was assessed under two widely divergent conditions of soil moisture. TWO experi- mental areas were located near Manitouwadge, Ontario, (approximately 49 N Latitude, 86 W Longitude) in the Central Plateau Section, B8, of the Boreal Forest (Rowe, 1959). Two field experiments were initiated in 17 the areas; the first in 1971 and the second in 1972. The cover type prior to cutting on the 1971 area was jack pine (70 percent), black spruce (25 percent), white spruce, white birch and poplar. Sixty percent of the timber volume was black spruce on the 1972 area with lesser amounts of jack pine and poplar. On each location, two sites were selected arbitrarily and called an upland dry site and an upland moist site. The former had originally supported a stand of jack pine; the latter, a stand of black spruce. B. Soil-Site Description and Analytical Methods Soil pits were dug and profiles described in each of the two field sites in both 1971 and 1972. A representative sample was taken from each horizon of each pit and placed in an individual plastic bag for transfer to the laboratory. In addition, a large bulk sample was removed, consisting of the A1 (Ah), A2 (Ae) and t0p 10 to 15 cm (3.9 to 5.9 inches) of the B2(Bf) horizons. The horizon samples and a sub—sample of each of the two bulk samples were transferred to the Great Lakes Forest Research Centre in Sault Ste. Marie, Ontario for nutritional analysis. A double check on nutrition levels of the two soil samples was obtained by sending sub- samples to the M.S.U. Soil Testing Laboratory. Nitrogen was analyzed by the semi~micro Kjeldahl technique. Phosphorus was determined by using a Bray P solution compared to 1 standards in a colorimeter. Potassium was measured in a flame emission spectrophotometer and calcium and magnesium were analyzed in an atomic absorption spectrophotometer. The pH was measured using a pH meter with a 1:1 soil:water ratio. Organic matter was measured by loss on 18 ignition (gravimetrically). An additional sub-sample of each bulk sample was analyzed for texture using a standard S.S.S.A. hydrometer technique (Black, 1965). Vegetation assessments were made on both sites to determine major species returning after harvest. In addition, an assessment of dead mosses was done to aid in site classification. The remainder of the 1971 bulk soil samples were transferred in sealed cans to Michigan State University. A soil moisture retention analysis was made at l/lO, 1/3, 1, 2, 10 and 15 atmospheres tension levels with standard pressure plate equipment. 4. GREENHOUSE POT TRIALS A. General Two greenhouse experiments were established to determine and compare the growth of young jack pine, lodgepole pine, hybrid pine and black spruce seedlings in the two field soil mixtures at: a) different soil moisture levels, and b) different soil fertility levels. The bulk field samples were air dried and sieved through a 2-mm sieve. Styrofoam potss) 22.5 cm (8.6 inches) high x 13.0 cm (5.1 inches) top diameter were filled with soil to obtain 96 pots of each soil type. Forty-eight pots of each type were randomly selected for a moisture stress experiment and the remaining 48 of each type were assigned to a fertility study. The two experiments were established in January, 1972 under greenhouse conditions of 21.1 C (:2) (70 F) temperature, and 16 hour 5) Courtesy, Dart Container Co., Mason, Michigan. 19 light, with available sunlight supplemented by 2500 ft-c artificial light. B. Soil Moisture Levels The experiment was a completely randomized factorial design with four replications. Treatments were: four seedlots, two soil types (called moist and dry to indicate their site origin), and three moisture levels. Each pot of each soil was randomly assigned a moisture and species treatment and allocated a spot on the greenhouse bench. In mid-January, 15 seeds of the assigned species were sown in each pot. Air—dried pot weight was obtained at this time and then the pots were soaked. On the tenth day after sowing, as primary needle initiation began, the best five seedlings in each pot were selected for retention and all other seedlings were removed. At this time soil moisture treatments were initiated. Treatments were based on the soil moisture retention studies and selected to represent low, intermediate and high levels of soil moisture stress at moisture levels equivalent to 1/3, 2 and 10 atmospheres (one-third atmosphere is hereafter referred to as field capacity; however, this may not be applicable in the field sites). Pots, based on weight, were allowed to reach the weight equi- valent to a percent moisture content prescribed as a treatment level. Moisture recharge was done daily to return the pot to its predetermined weight. water was added throughout the pot soil by a syringe needle which was supplied by an automatic pipette. Pots were removed from the greenhouse on May 8, 1972 and the l6—week-old seedlings were extracted and measured as follows: 1) for each seedling; top height, and length of the longest root; 2) for all seedlings in a pot; top (including stem and needles) and total root 20 oven—dry-weight; and 3) nutrient analyses of needles and roots. Needles were analyzed for nitrogen with a semi-micro Kjeldahl procedure digesting in the presence of a mercury catalyst. Phosphorus was extracted using a nitric-hydrochloric acid procedure and was deter— mined colorimetrically upon development of a phospho—molybdenum blue. Potassium, calcium and magnesium were determined by atomic absorption spectroscopy. Total roots were analyzed using the above methods but only for phosphorus, potassium, calcium and magnesium. C. Soil Fertility Levels The experiment was a completely randomized factorial design with four replications. Treatments were: four seedlots, two soil types (called moist and dry to indicate their site origin), and three fertility levels. Pot preparation, seed sowing and seedling selection were performed in a manner similar to the Soil Mbisture Level experiment (43). Soil fertility levels selected, based on assessment of nutrient levels, were: 1) control, 2) Grade C, a low level of nursery soil fertility satisfactory only for pioneer pines, and 3) Grade B, a moderate level of nursery soil fertility satisfactory for the majority of conifers. The latter two treatments are levels recommended by Wilde (1958) as necessary in order to obtain healthy seedling growth. Nutrient solutions were mixed so that sufficient N, P and K could be added to each soil and fertility-level treatment combination to raise the level of NPK to the prescribed level. Solutions were added in three bidweekly applications to the soil surface commencing three 21 weeks after sowing. The soil in each pot was soaked throughout the experiment, but care was taken to avoid over-watering to a level that might cause leaching. Pots were removed from the greenhouse on May 10, 1972; seed- lings were removed and analyzed as described in 43. Results for 4B and 4C were analyzed by the variance method and Duncan's New Multiple Range test was used to test significance of treatment means . 5. EXPERIMENTAL FIELD SEEDING IN 1971 AND 1972, MANITOUWADGE, ONTARIO A. 1971 Seeding The experimental seeding in 1971 was completed on June 2, on the moist and dry sites (Figure 2) previously described. The seedbed was prepared by a Bracke scarifier—seeder (Bergman, 1971) pulled by a Timberjack wheeled skidder. The rotating teeth on the Bracke flipped over the upper 8 to 15 cm (3.2 to 5.9 inches) of the organic matter and soil horizons, exposing soil of the A (As) and B horizons. 2 Scalps were made at a spacing of 1.2 m (4 feet). At the time of seed- bed preparation, the air was extremely hot and dry, and the soil was dusty (Figure 3). The seeding experiment was designed as a completely random block with four replications. Each block.was to contain four rows of 25 seedbed scalps per row. Each row was to be randomly assigned a species for seeding. This design was established in the moist site but had to be modified in the dry site due to an error by the tractor operator. Instead, two blocks were established, each block having four rows, each row having 40 to 50 scalps per row (Figure 4). 22 Figure 2. A general view of dry site cutover near Manitouwadge, Ontario. Figure 3. Bracke scarifier-seeder pulled by Timberjack wheeled skidder, preparing seed spots on dry site skid road and cutover. 23 MOIST SITE 25 SEEDSPOTS PER ROW Figure 4. Schematic presentation of plot layout for 1971 seeding location. j? = jack pine, 1? = lodgepole pine, hP = hybrid pine, b8 8 black spruce. 24 Seeding was done by hand using a bottle shaker obtained from the Ontario Department of Lands and Forests. Seed was shaken onto the exposed seedbed, and stepped on lightly to ensure seed-soil contact. Attempts to calibrate the seed shaker to release five to eight pine seeds or 9 to 12 spruce seeds were not very successful and the actual number of seed per scalp is unknown. ‘Soil temperature was measured and recorded by means of record- ing thermographs with 12.7-cm (five—inch) chromium-plated sensors that were placed on the soil surface. One thermograph was placed on each site, and seedbed temperature was recorded for the duration of the 1971 growing season. Additional temperature and rainfall information for the area was obtained from the Manitouwadge Ontario Department of Lands and Forests office, approximately 24 km (15 miles) southwest of the experimental areas. Germination was assessed periodically throughout the 1971 and 1972 field seasons. Seedlings on randomly selected scalps were excavated for measurement in late August, 1972. B. 1972 Seeding The 1972 seeding was established on sites similar to those selected in 1971 with all the four tree seedlots. It differed markedly in field procedure, and had an additional treatment, rodent protection, to determine if rodents influenced germination and survival. The experiment was established as a random block factorial design with two replications. Treatments were: two sites, four seedlots, and two rodent prevention levels, (screened scalp, or non- screened scalp). 25 Each replication contained eight rows of 25 seedbed scalps per row. Scalps were located at 1.8 m (6 feet) intervals from a base line. All scalps were hand prepared by fire rake and mattock to ensure seed- bed uniformity. Organic matter was removed to expose the A1 (Ah) horizon and the latter material was mixed lightly with the underlying A2 (Ae) soil. Each pair of rows (one and two, three and four, five and six, seven and eight) within a block was randomly assigned a species for seeding. Then within a pair of rows, one of each pair of side-by—side scalps was randomly assigned a rodent protection treatment. This treatment consisted of a 45.7-cm (18-inch) conical-shaped mesh which completely covered the seed scalp. Thus, one of each pair of scalps was covered with a screen and the other was exposed (Figure 5). Seeding was done by hand on June 25, 1972. Five seeds, of v the species allocated to a row, were carefully placed in each scalp and stepped upon lightly. Screens were then placed over those scalps assigned to receive rodent protection. Germination was assessed weekly. Colored toothpicks were used to mark individual seedlings as they germinated. A different color was used for each weekly period. Detailed environmental observations were recorded from mid- June to early September, 1972 on both sites. Observation techniques and instruments used were as follows: 1. Temperature at the seedbed surface was measured at weekly intervals at nine randomly-assigned spots in each block, using a 24-gauge copper- constantan thermocouple (Fraser, 1968) connected 26 to a portable potentiometer. Temperature was measured at mid-afternoon. 2. Continuous recordings of the seedbed temperatures in each block were obtained by thermographs with 12.7-cm (5-inch) chromium-plated sensors placed on the soil surface. 3. Air temperature and relative humidity were continuously recorded using a Fuess hygrother- mograph located on each site. Each hygrother- mograph was placed in a standard Stevenson screen 114.3 cm (45 inches) above ground level. 4. Soil moisture content by weight was obtained twice weekly for the upper 5 cm (2 inches) of soil using gravimetric methods, for nine randomly selected seedbeds in each block. 5. Precipitation was measured by means of a standard Canadian 65.9-sq cm (lo-square inch) rain gauge, at three locations per site. Rainfall was measured daily until September, then periodically until October. 6. Solar radiation was continuously recorded on each site by actinographs. In late August, lOdweek-old seedlings were excavated from several scalps in each site, washed and measured for top and root lengths. 27 x o o x x o g o jP = jack pine o x o x o x x 0 IP = lodgepole pine x o o x o x x o hP = hybrid pine o x x o o x o x bS = black spruce o x o x x o o x spacing: 1.8 m (6 feet) X o x x o o x o x 1.8 m (6 feet) x-SuMnduMInuhd o-fiaunisummuluuanuhd Figure 5. Example of independent randomly paired plot layout, 1972. CHAPTER IV RESULTS AND DISCUSSION OF EXPERIMENTS l. SEED ORIGIN AND COLLECTION Seed from the jack pine and hybrid pine collections were of relatively poor quality. Germination for the two jack pine seed collections averaged 75 percent and 63 percent and germination for the hybrid pine seed was only 60 percent. Lodgepole pine germination varied among populations, but all populations had significantly higher germination than both jack pine and hybrid pine. Black spruce germina- tion was significantly better than all pine species. Details of initial germination tests for all species are shown in Appendix A. 2. CARDINAL TEMPERATURES FOR GERMINATION A detailed summary of germination by population and temperature after 7, 14, 21 and 28 day periods is provided in Appendix B. A summary of mean percent germination by species for each temperature is found in Table 1. Table 1 includes a summary of germination results for black spruce from Site Region 3E adapted from Fraser (1970b). Figure 6 presents germination by species and temperature, 7 and 28 days after seeding. Lodgepole pine and hybrid pine are both more temperature sensitive than either jack pine or black spruce. Both jack pine and black spruce seeds germinate quickly over a wide range of temperatures. The optimum range for jack pine germination was determined to be 15.6 C (60 F) to 32.2 C (90 F). Optimum black spruce germination was obtained by Fraser at 15.6 C (60 F) to 26.7 C (80 F), with acceptable 28 29 swamp muaumquEMu coaumnwaumw abEHumo n. ouosov meson musumuwnewu vocaauso .mmfiommm mo Ancemav Humane sown emuamu< « mm N o o o o o o o o o o o o o o As mas o N.“ mm mm OH O H o o o N o o 0 mm ON 0 0 Am onv o o.oa “WSI.MM._NQ 0 HH a o o m o s o No mm mm o Am mmv o m.~H .3 also 2 S S 3 o S 8 S o .wmlomummdr H C 08 o 0.2 "8 Na romlm maidenmfl m “mime: 3.4.2 .2 fl 2 rmm E a: o 3: "No om R we ”rmmINNImINMIImW "mm mm mm rwlm "on me we so Am 08 u Tam “om om mm mm mm mm mm on "an On me No .mo me me no Am mmv o m.mu .3 mm 3 No me me as aw lemhllmlnlmm "an 3 .K 3 Am 08 05.3.. Pm...man.mm.umw on OH m N am am on sH "me me nu ma As mas o s.m~ us an we mm m o e N as mm mm on .mo.lmm.lmmu.mw Am Gay 0 «.mm mm mm Hm NH w o a N sq on on ma mm mm an an Am nmv o o.mm mm am «a m mm an «a 5 mm am «a 5 mm am «a 5 Amy 0 muaumumaama «madman xumHm mafia maunzm mafia maomemvog mafia 30mm msfipmmm umuwm when was mmuaumumdewu udmummwfiv um «weaken xowan was mean manna: .mcfia maoamwpoa .mafid sown mo soaumcfiawow assumed new: .H manme § ‘1 9 §§§§$§ 3O PERCENT GERMINATION a 3 S 6 8 8 20* 28 DAYS Ho 4: so 53 do 65 7'0 73 0'0 3'5 9'0 9'5 160 CM 7.2 no.0 12.: I“ 13.3 21.1 23.9 26.7 29.4 32.2 35.0 37.: TEMPERATURE -- .............. ~Black Spruce ———lodgopolo Pine ....... Jack Pine —‘-—~—-- Hybrid Pine Figure 6. Mean seed germination for three species plus hybrid pine at several temperatures, 7 and 28 days after seeding. 31 germination still obtained at the extremes of 32.2 C (90 F) and 10.0 C (50 F). Lodgepole pine germinated rapidly at temperatures in the range 21.1 C (70 F) to 26.7 C (80 F), but was delayed at 18.3 C (65 F) and germination was reduced at all other temperature levels. Optimum temperature for germination was 21.1 C (70 F). Hybrid pine germinated rapidly at only one temperature, 21.1 C (70 F), and attained maximum germination at only 18.3 C (65 F) and 21.1 C (70 F). Germination levels for hybrid pine were 17 percent higher in the cardinal tempera- ture experiment than in the initial germination test. This is possibly due to after-ripening of immature seed during the time lapse between the initial tests and subsequent cardinal temperature studies. All populations of lodgepole pine show a significant reduction in germina- tion at 23.9 C (75 F). This can only be attributed to a species germination characteristic. Results obtained for jack pine agree with those reported by Fraser (1970a) and by Ackerman and Farrar (1965). Optimum germination from lodgepole pine seed (86 percent in 28 days, Table l) agrees with results obtained by Ackerman and Farrar (1965) for this species (88 percent in 21 days at 21.1 C (70 F)). However, they also obtained germination at "alternating 15.6 C (60 F) and 26.7 C (80 F), and 15.6 C (60 F) constant for four days followed by 32.2 C (90 F) constant for the remainder of the test", and suggested that a two-stage germina- tion process might be necessary. The first stage requires temperatures in the 15.6 C (60 F) to 21.1 C (70 F) range, while the second stage requires temperatures of 21.1 C (70 F) to 32.2 C (90 F). Alternating temperatures were not used in this study. While constant temperature studies provide valuable information on species seed characteristics, 32 further studies should be initiated to simulate field temperatures which fluctuate from 23.9 C (75 F) to 35.0+ C (95+ F) during the day and from 1.8 C (35 F) to 12.8 C (55 F) at night, during the growing season . 3. SOILS AND VEGETATION AT FIELD LOCATIONS The soils are Spodosol (Podzol) sand tills, several feet in depth with a thin 5 to 10 cm (2 to 4 inch) organic mantle. Individual horizon characteristics are detailed in Table 2 and chemical properties are presented in Table 3. All soils studied were acidic. The soil pH was higher in moist sites (5.9) than dry sites (3.6) and increased with soil depth (Table 3). The pH values obtained for the bulk samples are slightly higher than the horizon sample levels. This may be due to a several month delay in measurement of pH for the bulk samples. The nutrition levels determined for the bulk samples were confirmed by the M.S.U. Soil Testing Laboratory. Textural analysis of the bulk soil samples indicate sand con- tents greater than 75 percent in soils from both moist and dry sites. Silt content was slightly higher in moist site soils (Table 4). Results of the soil moisture retention studies are found in Table 5. The moist site soil has a higher moisture content at all tension levels (Figure 7). This can be attributed to its higher silt and organic matter content compared to the dry site soil. Ground vegetation at the time of seeding was minimal. Logging slash and needle litter formed a heavy layer over much of the cutover. The major moss species on the moist site was Pleurozium schreberi 33 Table 2. Detailed soil pedon description for Manitouwadge moist and dry sites Moist site soil Classification: Typic Cryorthod* (Orthic Humo Ferric Podzol)** Depth Horizon (cm) Description L—F 5.0- 2.5 Raw to fermented litter. F-H 2.5- 0.0 Raw to fermented litter. Al (Ah) 0.0- 3.0 Black organic matter, sand (7.5 YR 2/0). Abundant roots. A2 (Ae) 3.0-10.5 Grayish brown (10 YR 5/2) loamy sand. Abundant roots, friable when moist. B2 (Bf) 10.5-51.0 Dark brown (7.5 YR 4/4) sand. Occasional roots. Blocky structure, firm when moist. C 51.0+ Light brown (7.5 6/4) sand and gravel. Hard and cemented. No roots. Dry site soil Classification: Haplic Cryorthod (Mini Humo Ferric Podzol) Depth Horizon (cm) Description L-H 3.5- 0.0 Mixture of raw to decomposed litter. A1 (Ah) 0.0— 2.5 Black (2.5 Y 2/0) sand and organic material. Abundant roots. A2 (Ae) 2.5- 7.0 Light brownish gray (10 YR 6/2) loamy sand. Discontinuous at intervals. Abundant roots. Friable when moist. B2 (Bf) 7.0-80.0 Dark yellowish brown (10 YR 4/4) sand. Fine texture, with iron concentrations. Firm.when moist. Occasional roots. C 80.0+ Very pale brown (10 YR 7/3) sand and gravel. Firm when moist. No roots. * United States Department of Agriculture, 1960 ** Canada Department of Agriculture, 1970 34 00.0 00.0 00.0 0.0 0.00 0.00 0.00 0.0 000.0 0 00.0 00.0 00.0 0.0 0.00 0.00 0.00 0.0 000.0 0000 0 00.0 00.0 00.0 0.0 0.00 0.00 0.000 0.0 .000.0 0000 00 00.00 00.00 00.0 0.0 0.000 0.000 0.0000 0.0 000.0 0000 00 00.00 00.00 00.0 0.0 0.000 0.0000 0.000 0.00 000.0 0000 000 00.0 00.0 00.0 0.0 0.00 0.000 0.00 0.0 000.0 0 00.0 00.0 00.0 0.0 0.00 0.000 0.00 0.0 000.0 0000 00 00.0 00.0 00.0 0.0 0.000 0.0000 0.00 0.0 000.0 0000 0< 00.000 00.00 00.0 0.0 0.000 0.0000 0.00 0.0 000.0 00<0 00 00.00 00.00 00.0 0.0 0.000 0.0000 0.000 0.00 000.0 0000 0000: 00 000000a0 000 E00 000 z 000000: 0000 kufiumamo 000008 Hmuoa mwcmnuxw ownmwuo mm mz mu M m 000000 mwum mcfipmmm Hmma mwmhamam Hmowawzu 600000: 000m .m wanes 35 Table 4. Soil physical and chemical properties for moist and dry site bulk samples Texture (Z) kg/ha available Site Sand Silt Clay ZN P20 K o 5 2 Mg pH Z O.M. Moist 77 14 9 .166 13 65 4925 342 6.2 7.41 Dry 82 9 9 .041 25 53 29 5.1 2.74 Table 5. Soil moisture content (percent by weight) for moist and dry site bulk samples Soil moisture (Z by weight) Tension (atm) Moist site Dry site 0.10 17.5 9.3 0.33 12.3 6.4 1.00 11.9 5.0 2.00 9.3 5.0 3.00 9.0 4.7 10.00 6.6 3.5 15.00 6.3 3.2 184? § E § 84% ex» 41F Soil Moisture Content (PERCENT BY WEIGHT) 2J¥ 0.0-I Figure 7. 36 MOIST SITE DRY SITE I I I 1 1 V V 2.0 4.0 0.0 0.0 10.0 12.0 14.0 “To Soil Water Suction (cums) Soil moisture curves for bulk soil samples obtained from Manitouwadge, Ontario. 37 (BSG.) Mitt. with occasional patches of Dicranum Spp., and Hypnum crista-castrensis Hedw. Pleurozium schreberi was again most frequent on the dry site. Hypnum crista-castrensis and Dicranum spp. were more frequent than on the moist site. A vegetation survey in August, 1972 indicated considerable regrowth of vegetation on the moist site but very little on the dry site. Major plant species occurring in abundance on each site are as follows (not in order of frequency of occurrence); Moist site: Equisetum sylvaticum L., Fragaria virginiana Duchesne, Rbsa acicularis Lindl., Epilobium angustijblium L., Cornus canadensis L., and Oryaopsis spp. Dry site: Vaccinium spp., Rosa acicularis, Ledum groenlandfcum Oeder and Salim spp. It should be noted that Vaccinium spp. were not observed on the moist site while Fragaria virginiana and OryzoPsis spp. were not found on the dry site. 4. GREENHOUSE POT TRIALS A. Soil Moisture Levels A detailed tabling of the results for the growth parameters measured for all treatments is presented in Appendix C. A brief summary and discussion is reported in the text. 1. Height growth: all species produced significantly greater height EIOWth at 1/3 atmosphere; however. growth of lodgepole pine and black spruce in the moist site soil at 10 atmospheres tension was not significantly different than growth at field capacity. Site effect was only significant for black spruce grown in the moist site. Hybrid 38 pine and jack pine grown at field capacity in the moist site were significantly better than all other species, except for hybrid pine grown at field capacity in the dry site soil (Figure 8). 2. Length of the longest root: all species, except jack pine, produced significantly longer roots in the moist site soil than in the dry site soil. Lodgepole pine and hybrid pine produced significantly longer roots at field capacity than at other moisture levels, but there was no difference between the three pine species at field capacity. All pine species were significantly better than black spruce at field capacity on both the moist and dry sites (Figure 8). 3. Top weight: increased soil moisture content produced significantly greater oven-dry top weight for all species, except for lodgepole pine and black spruce at the two atmosphere level. The effect of site type varied from species to species. Black spruce did equally as well at 10 atmospheres suction as at two atmospheres, and in the dry site soil performed better at 10 atmospheres. It should be noted, however, that a few black spruce seedlings died at the 10 atmOSphere, dry site treatment. This can be attributed to surface drying and resultant root dessication when roots were still developing. Jack pine grown in the moist site at field capacity was significantly better than all other species at the equivalent treat— ment. Top weight production by lodgepole pine was considerably less than by the other pines except at the two atmosphere moisture level. 4. Total root weight: within treatment variation occurred and statistical analysis did not indicate any significant differences between treatments with one exception. Black spruce root weight was significantly lower than the pines at all moisture and site treatment 39 '/:1 2 IO N\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\. nun V3210 60‘AMM 45-1 #103: m0» Zwa kufiaauuom 000o monouosfi ou pmufisvou munmEmnmEm 0:00003: mam mwmmamam unowuunc H0om .o manna 43 growth than jack pine and lodgepole pine but not hybrid pine. Growth of black spruce in the moist site was significantly greater than in the dry site soil at all fertility levels, despite the addition of nutrient solutions to bring the two soils to comparable levels of N, P and K. Pines grown in the moist site at control produced significantly better growth than in the dry site soil, but there was no difference in pine growth between sites at the low and high fertility levels (Figure 9). 2. Mean length of longest root: root length varied inversely with increased fertility level for all pine species except hybrid pine grown in the dry site soil. There was no significant difference among pines at comparable levels, but all pines had significantly longer roots than black spruce. Black spruce and hybrid pine were the only species that had significantly longer roots in the moist site. Black spruce was not significantly affected by increased fertility levels (Figure 9). 3. T0p weight: the low fertility level (Wilde's Grade C) produced significantly increased top weight, compared to control, for all pines in both soils, except for lodgepole pine and hybrid pine in the moist site soil. Considerable variation occurred among black spruce treatments, resulting in no significant differences between sites or fertility levels despite the four-fold difference in mean weights between moist site and dry site soils at high fertility. 4. Total root weight: increased fertility had no significant effect on root weight of any species in the moist site, but was highly significant for all pines in the dry site. 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Seedlings from the 1971 and 1972 dry site field seeding trials (reduced x 2.5). Left to right: lB—month-old black spruce, hybrid pine, jack pine, lodgepole pine; right: lO-week-old lodgepole pine, black spruce, jack pine and hybrid pine. e x Figure 16. Black spruce seedlings, from the dry site.field seeding trials, (actual size). Left: lS-month-old; right: 10- week-old. 67 critical temperature requirements. Mortality of all species was minimal. About 50 percent of those killed had been snipped-off just below the cotyledons by insects. Mortality of the other 50 percent can be attributed to flooding, soil surface drying, or failure of the hypocotyl to develop. Measurements of seedlings removed from both the 1971 and 1972 seeding trials indicate slightly greater growth by jack pine than by the other pines of the same age, except on the 1971 dry site where the few surviving hybrid pine germinates produced greater top growth (Figure 15, Table 12). Growth of black spruce was considerably poorer and indicates that this species may be much more susceptible than the pines to extremes of climate. This poorer growth of black spruce, particularly root development (Figure 16), may account for the increased mortality experienced in the 1971 field seeding trial. Seedlings, such as pines, with larger root systems would be less susceptible to soil washing and root exposure than black spruce. At no time in the 1972 study did site have a significant effect on germination. As mentioned previously, jack pine germinated significantly faster than the other species. Lodgepole pine germina- tion rates became similar to those of jack pine, but not until the third assessment was made on August 1. Screening placed over the seedspots increased germination, but this was found to be significant only for lodgepole pine on the dry site. There were no other main effects or interactions. (Table 9)- Results of the screening study were quite variable and to date are inconclusive. Screens did not significantly alter tempera- tures. The screens may have shaded the seedspots and may have 68 intercepted precipitation. Based on the variable results, it appears that the screens slightly altered the microsite, but they certainly did not indicate a rodent population or problem. 69 Table 12. Growth measurements for seedlings extracted in August, 1972 from both the 1971 and 1972 field seeding trials Mean* length Date of Mean* top height of longest root seeding Site Species mm (inch) mm (inch) 1971 Dry Jack pine 30.2 (1.19) 154.6 (6.09) Hybrid pine 36.3 (1.43) 166.0 (6.54) Lodgepole pine 27.1 (1.07) 180.2 (7.09) Black spruce 16.4 (0.65) 58.5 (2.30) 1971 Moist Jack pine 45.3 (1.78) 181.4 (7.14) Hybrid pine 21.7 (0.85) 101.7 (4.00) Lodgepole pine 23.9 (0.94) 142.4 (5.61) Black spruce 22.1 (0.87 69.3 (2.73) 1972 Dry Jack pine 17.0 (0.67) 91.3 (3.59) Hybrid pine 14.3 (0.56) 48.1 (1.89) Lodgepole pine 13.8 (0.54) 64.5 (2.54) Black spruce 7.2 (0.28) 31.7 (1.25) 1972 Moist Jack pine 18.0 (0.71) 67.4 (2.65) Hybrid pine 12.6 (0.50) 41.2 (1.62) Lodgepole pine 15.2 (0.60) 61.3 (2.41) Black spruce 8.7 (0.34) 26.1 (1.03) * Mean of 10 seedlings. CHAPTER V SILVICULTURAL IMPLICATIONS OF RESEARCH RESULTS Contrary to past seeding results reported in Ontario, black spruce seeds were found to germinate and seedlings survived on care- fully prepared seedbeds, on a large cutover in a year of above normal precipitation. Many sources of variation occurred in the 1971 seeding which obscure comparisons between the 1971 and 1972 seeding trials. Nevertheless, black spruce seeds did germinate in both years and on both sites despite climatic and seedbed-preparation differences. The results indicate that black spruce requires a more carefully prepared seedbed than the pine species tested; they also verify reports that a humus and mineral soil mixture is desirable for germination and survival. Thus careful attention must be paid to seedbed preparation in order to obtain successful field germination of black spruce seed. Both jack pine and lodgepole pine germinated and survived well in the 1971 and 1972 seedings, and on both site types. The two species are well suited to harsh cutover conditions and do not require as carefully prepared seedbeds as does black spruce. Based on the experimental results in both field and pot trials, there is little difference in the growth of the two pines or in their ability to survive. Field trials must be continued over several years to deter- mine if any benefits can be obtained by substituting lodgepole pine for jack pine. At the seedling stage, jack and lodgepole pines would appear to be good substitutes for black spruce; however, hybrid pine appears less attractive because of its seed's inability to germinate satisfactorily in the year of seeding. 70 71 All field sites tested were deficient in phosphorus and potassium. Nitrogen was deficient in dry site soils, and in some pits sampled in the moist sites. The fertility pot trials indicated that significant increments in growth could be obtained by the addition of fertilizers to the field soils. Nutrient additions should be formu- lated to raise soil fertility levels to those prescribed by Wilde (1958) as Grade C, for pines, and Grade B for black spruce. Black spruce seedlings were found to survive and grow in soils at moisture levels as low as 10 atmospheres suction. This indicates that survival is affected by factors other than soil moisture alone. The three pines tested grew better at high soil moisture levels than at low levels. Jack pine and lodgepole pine appear to be the best species for seeding on dry sites. LITERATURE CITED LITERATURE CITED Ackerman, R.F. 1955. The regeneration of lodgepole pine after clear cutting following mechanical scarification and fire as methods of seedbed preparation. Canada, Dept. Northern Affairs and Natural Resources, For. Br., For. Res. Div., Unpubl. Ms. Ackerman, R.F. and J.L. Farrar. 1965. The effect of light and temperature on the germination of jack pine and lodgepole pine seeds. Univ. Toronto, Fac. For., Tech. Rep., No. 5, 41 p. Anderson, N.A. and R.L. Anderson. 1965. The susceptibility of jack pine and lodgepole pine and their hybrids to sweetfern rust (Cronartium camptoniae) and eastern gall rust (C. quercuum). United States, Dept. Agriculture, For. Serv., Lake States For. Exp. Sta., Res. Note No. LS-56, 4 p. Beaufait, W.R. 1960. Influences of shade level and site treatment, including fire, on germination and early survival of Pinus banksiana. Mich. Dept. Conservation, For. Div., Un-numbered Mimeo Rep., 79 p. 1962. Procedures in prescribed burning for jack pine regeneration. Michigan Coll. Mining and Technology, Ford For. Cent., Tech. Bull. 9, 39 p. Bergman, F. 1971. Direct seeding of Scots pine, Pinus silvestris. In: Techniques in silvicultural operations. Papers presented at the XV IUFRO congress, IUFRO Div. No. 3, Publ. No. 1,: 143-149. Black, C.A. Ed. 1965. Methods of soil analysis. Part 1. Agronomy. Amer. Soc. Agron. Inc., Madison, Wisconsin, No. 9, p. 562— 567. Boughner, C.C. and G.R. Kendall. 1959. Growing degree-days in Canada. Canada, Dept. Transport, Meteorological Br., Circ. No. 3203, 21 p. Canada, Department of Agriculture. 1970. The system of soil classifi- cation for Canada. The Queen's Printer of Canada, Ottawa, 249 p. Canada, Department of Transport. 1968a. Climatic normals. Tempera— ture. Meteorological Br., 1: 66. 72 73 Canada, Department of Transport. 1968b. Climatic normals. Precipita- tion. Meteorological Br., 2: 110. Cayford, J.H. 1958. Scarifying for jack pine regeneration in Manitoba. Canada, Dept. Northern Affairs and National Resources, For. Br., For. Res. Div., Tech. Note No. 66, 14 p. 1963. Some factors influencing jack pine regeneration after fire in southeastern Manitoba. Canada, Dept. Forestry, For. Res. Br., Publ. No. 1016, 16 p. Cayford, J.H., Z. Chrosciewicz, and H.P. Sims. 1967. A review of silvicultural research in jack pine. Canada, Dept. Forestry and Rural Development, For. Br., Publ. No. 1173, 255 p. Chrosciewicz, Z. 1960. A spot seeding trial with jack pine. Canada, Dept. Northern Affairs and Natural Resources, For. Br., For. Res. Div., Mimeo 60-1, 8 p. 1963. The effects of site on jack pine growth in northern Ontario. Canada, Dept. Forestry, For. Res. Br., Publ. No. 1015, 28 p. 1970. Regeneration of jack pine by burning and seeding treatments on clear—cut sites in central Ontario. Canada, Dept. Fisheries and Forestry, Canad. For. Serv., Ontario Region, Info. Rep. 0-X9128, 13 p. Critchfield, W.B. 1957. Geographic variation in Pinus contorta. Harvard university, Maria Meors Cabot Foundation, Publ. No. 3, 118 p. Crouch, G.L. and M.A. Radwan. 1971. Evaluation of R—55 and mestranol to protect Douglas-fir seed from deer mice. United States, Dept. Agriculture, For. Serv., Pacific Nthwest For. Range Exp. Sta., Portland, Oregon, Res. NOte, PNW-170, 6 p. 1972. Arasan in endrin treatments to protect Douglas—fir seed from deer mice. United States, Dept. Agriculture, For. Serv., Pacific Nthwest For. Range Exp. Sta., Portland, Oregon, Res. lap. PNW—136, 7 p. Derr, H.J. and W.F. Mann, Jr. 1971. Direct seeding pines in the south. United States, Dept. Agriculture, For. Serv., Agric. Handb. No. 391, 68 p. Fraser, J.W. 1968. A.method of constructing and installing thermo- couples for measuring soil temperatures. Canad. J. Soil Sci. 48: 366-368. 1970a. Cardinal temperatures for germination of white, red and jack pine seed. Canada, Dept. Fisheries and Forestry, Canad. For. Serv., Petawawa For. Exp. Sta., Chalk River, Ontario, Info. Rep. PS-X-lS, 29 p. 74 Fraser, J.W. 1970b. Cardinal temperatures for germination of six provenances of black spruce seed. Canada, Dept. Fisheries and Forestry, Canad. For. Serv., Petawawa For. Exp. Sta., Chalk River, Ontario, Info. Rep. PS-X-23, 12 p. Fraser, J.W. and J.L. Farrar. 1953. Germination of jack pine on humus. Canada, Dept. Resources and Development, For. Br., For. Res. Div., Silvi. Leafl. No. 91, 2 p. Hills, C.A. 1959. A ready reference to the description of the land of Ontario and its productivity. Ontario, Dept. Lands and Forests, Div. Res., Preliminary Rep., 142 p. Johnson, J.H., R.F. Cerezke, F. Endean, G.R. Hillman, A.D. Kill, J.C. Lees, A.A. Loman and JNM. Powell. 1971. Some implica- tions of large-scale clearcutting in Alberta. A literature review. Canada, Dept. Environment, Canad. For. Serv., Northern For. Res. Cen., Edmonton, Alberta, Info. Rep. NORrX—6, 114 p. Johnston, W.F. 1971a. Broadcast burning slash favors black spruce reproduction on organic soil in Minnesota. For. Chron. 47 (1): 33—35. 1971b. Management guide for the black spruce type in the lake states. United States, Dept. Agriculture, For. Serv., North Central For. Exp. Sta., Res. Pap. N.C.-64, 12 p. Lafond, A. 1958. Some soils, vegetation, and site relationships of the climatic and subclimatic black spruce forest in north- eastern America. In: First North American For. Soils Conf., Sept. 8—11, Michigan St. Univ., Agric. Exp. Sta., East Lansing. LeBarron, R.K. 1944. Influence on controllable environmental condi- tions on regeneration of jack pine and black spruce. J. Agric. Linteau, A. 1955. Forest site classification in the Northeastern Coniferous Section, Boreal Forest Region, Quebec. Canada, Dept. Northern Affairs and National Resources, For. Br., For. Res. Div., Bull. 118, IX + 85 p. 1957. Black spruce reproduction on disturbed soil condi- tions. Canada, Dept. Northern Affairs and National Resources, For. Br., For. Res. Div., Tech. Note No. 54, 14 p. Logan, K.T. 1966. Growth of tree seedlings as affected by light intensity. II. Red pine, white pine, jack pine and eastern larch. Canada Dept. Forestry, For. Res. Br., Publ. No. 1160, 19 p. ‘ 1969. Growth of tree seedlings as affected by light intensity. IV. Black spruce, white spruce, balsam fir and eastern white cedar. Canada, Dept. Fisheries and Forestry, For. Br., Publ. No. 1256, 11 p. 75 Mann, W.F. Jr. 1968. Ten years experience with direct seeding in the South. J. For. 66: 828-833. MacHattie, L.B. 1956. Winter injury of lodgepole pine foliage. Canada, Dept. Forestry, For. Res. Br., Cont. No. 536: 301—307. McQuilkin, W.B. 1965. Direct seeding in the northeast - Status and needs. In: Direct seeding in the north—east. A symposium. University Massachusetts, Amherst, p. 25-27. Miller, E.L. 1970. Studies of environmental factors affecting jack pine (Pinus banksiana Lamb.) regeneration. A Thesis for the degree of Doctor of Philosophy. Michigan State Univ., Dept. Forestry, 102 p. Miller, E.L. and G. Schneider. 1971. First year growth response of direct seeded jack pine. Michigan State Univ., Agric. Exp. Sta., Res. Rep. No. 130, 8 p. Mirov, N.T. 1956. Composition of turpentine of lodgepole x jack pine hybrids. Canad. J. Bot. 34: 443-457. Moss, E.H. 1949. Natural pine hybrids in Alberta. Canad. J. Res. 27 (5C): 218-229. Nostrand, R.S. van. 1971. Strip cutting black spruce in central Newfoundland to induce regeneration. Canada, Dept. Fisheries and Forestry, Canad, For. Serv., Publ. No. 1294, 21 p. Ontario, Department Lands and Forests. 1963. Ontario resources atlas. Oper. Br., Toronto, Ontario, 33 p. Place, I.C.M. 1955. The influence of seedbed conditions on the regeneration of spruce and balsam fir, Canada, Dept. Northern Affairs and National Resources, For. Br., Bull. 117, 87 p. Radvanyi, A. 1970. A new coating treatment for coniferous seeds. For. Chron. 46 (5): 406-408. Richardson, J. 1970. Broadcast seeding black spruce on a burned cutover. Canada, Dept. Fisheries and Forestry, Canad, For. Serv., Publ. No. 1272, 14 p. Rowe, J.S. 1959. Forest regions of Canada. Canada, Dept. Northern Affairs and National Resources, For. Br., Bull. 123, 71 p. Scott. J.D. 1970. Direct seeding in Ontario. For. Chron. 46 (6): 453-457. Sims, H.P. 1970. Germination and survival of jack pine on three prepared cutover sites. Canada, Dept. Fisheries and Forestry, Canad. For. Serv., Publ. No. 1283, 19 p. 76 Smithers, L.A. 1961. Lodgepole pine in Alberta. Canada, Dept. Forestry, Bull. 127, 153 p. 1965. Direct seeding in eastern Canada. In: Direct seeding in the north-east. A symposium. University Massachusetts, Amherst, p. 15-23. Swan, B.S.D. 1970. Relationships between nutrient supply, growth and nutrient concentrations in the foliage of black spruce and jack pine. Pulp and Pap. Res. Inst. Canada, Woodl. Pap., W.P./19, 46 p. 1972. Foliar nutrient concentrations in lodgepole pine as indicators of tree nutrient status and fertilizer require- ment. Pulp and Pap. Res. Inst. Canada, Woodl. Rep., W.R.42, 19 p. United States, Department Agriculture. 1948. Woody-plant seed manual. For. Serv., Misc. Publ. No. 654, 416 p. . 1960. Soil classification (7th Approximation), Soil Conservation Service, 265 p. Vincent, A.B. 1965. Black spruce a review of its silvics, ecology and silviculture. Canada, Dept. Forestry, Publ. No. 1100, 79 p. Wilde, S.A. 1958. Forest soils. Their prOperties and relation to silviculture. Ronald Press Co., New York, 563 p. Yeatman, C.W. 1967. Biogeography of jack pine. Canad. J. Bot. 45: 2201-2211. Yeatman, C.W. and A.H. Teich. 1969. Genetics and breeding of jack and lodgepole pines in Canada. For. Chron. 45 (6): 428-433. Yeatman, C.W. and M.J. Holst. 1972. Hardiness of jack pine hybrids and provenances in the boreal forest. For. Chron. 48 (1): 30-31. APPENDICES APPENDIX A Details of seed origin and initial germination tests 77 .Hhuooommn uuonm no oaoavou m :uw3_vouoc«auow meson moan hufiafinmswahou N I hufiawnma> N .Ammmn HNV uowuom umou ca mafiavomm Ou vomoao>on was woumcaakom mcmmm I muHHHnmcaauou N o.ma o.¢m coauomaaoo ovwslaoawou mm cofimwu cashew aoum mamawm mufim oaumuno xomam m.qo m.mo vcmum Gaoum sumo moauooa q.¢~|m.- w.HMIm.o~ N Houvoa ounumaaa .Smmsuao vcmm noon z.mq oNq .umA m.on m.mn vnmum meson me m.oa|n.ma o.mHIN.oH a 3.m~ omw .msoa Houvom ousumaafi .oaumuno mafia .Smmsuso vamm noon .smoammso xumh z.mH oem .uma o.«o m.oo cOHumHmwmp HOH o.mN «.H« H 3.«m oOHH .waoH wanouw uaofiusxsa .muuonH4 mafia mama muHNm och .xoouo xom vaunhm o.oN n.mN Ho>maw,ammuum «0H N.oN «.NN N um>o Houvom 3oHHm£m o.am n.mm Hm>amm,ammuum Hm «.ON N.NN N um>o Hoswom soHHmnm z.N¢ 0mm .umq m.om m.¢m masons Hmwomdw, no m.H~ 0.0m H 3.0a oNHH .wcoq uo>o moon so N.nq .muumnd< mafia owvfiu Houuoa Noamm .cmsouonaumz maomowcoq hufiafinmfi> aowum mxumawu Hmuoamu Amumohv AEV Aaov vnmum coaumuoa mofiomam N Icfisuoo ow< unwaon .m.m.a no omua N Heuoa mummu GOHumawapow Hmfluwafi mam awwfiuo poem mo mafimuma .mH mHan APPENDIX B Cardinal temperature study 7*, 14-, 21-, 28-day germination percent means by species and population 78 o o o o o o Anew N.N O O O O O O HOOO O.OH O O O O O O Hmmv O.NH O O O O H O NOOO O.mH O O OH NO ON OO NOOO N.OH NO ON ON ON mm HN NONO H.HN Om OO NO «O HO ON HONO O.MN ON NO ON NN ON ON NOOO N.ON N OH OH HH .. NN NOOV «.ON N O OH OH NO NO NOOO N.Nm N OH NH OH on O« HOOO O.mm mcHa ON maHa Ne maHa HN mcHa Ne HN HNO O vaunmm maomomcog oaomomuoa oaomomvoq mafia xomm mafia sown musumumaamy coaumasaoa was moaomam ha names ucmouma coeumCHEHmm hmwln "manum ounumuoaawu Hmcfiwumo .qa manna 79 o o o o o o Amqv «.5 O O O O O O NOOO O.OH O O « O O« «N NOOO O.NH H« ON NO NO OO OO NOOO O.OH NN NO HO OO OO ON NOOO N.OH ON OO HO OO OO NN AONV H.HN NN OO NO ON NO ON HONV O.NN H« NN NO ON ON ON NOOV N.ON O OH NN OH .. NN NOOO «.ON « OH ON «N OO NN NOOO N.NN « ON ON HN OO HO NOOO 0.0N maHO NO maHO NO maHO HO maHO NO HO NOV O vNunmm odommwcoa oaomowuoa oHoaowvoq mafia xomh mafia xomw ousumumaawa GONumaamoa was mmfiowmm mp mamas Hcmouma cowumcaeuow zmvlqa "mvsum unaumuoaewu HmCNnHmu .mH magma 80 o o o o o o Amev N.“ O H O H NN O NOOO 0.0H O N OH N OO HO NOOO O.NH NO NO OO OO NO HN NOOV 0.0H «N HO NO ON OO OO NOOO N.OH ON OO HO OO HO NN NONO H.HN ON NO OO NN NO ON NONV O.NN N« «N «O NN ON ON NOOO N.ON OH NN ON HN .. NN NOOO «.ON O ON N« ON OO NN NOOO N.NN O HO O« H« OO NO NOOO 0.0N OOHO NO 9:O NO OOHO HO OOHO NO HO ONO O vaunmm mHoaomvoa oaomwmvoq maoamwvoa ocwa xomh mafia xomh unnumumaswe coaumHsaom mum mmNommm an mcmoa unmouma CONuchahmw Outlaw ”knsum manumquEOO Hmawwumo .oa manme 81 o o o o o . o Amcv N.n H N O « O« ON NOOO 0.0H HH N OH O NO OO NOOO O.NH «O NO ON OO NO «N NOOO 0.0H ON NO NO HO NO HO NOOO N.OH NN OO HO OO HO NN NONO H.HN ON OO OO ON NO ON NONO O.NN N« «N OO «N ON ON NOOO N.ON HH ON NN ON .. NN NOOO «.ON O NN HO «O OO NN NOOO N.NN O ON ON HO OO NO NOOO 0.0N OOHO NO «ONO NO OOHO HO OOHO NO HO NOV O uNunhm maoammwoa waoamwwoq maomowvou mafia gosh maNm xomh unnumumaewh GONumasmoa was mowommm hp mamma unmouoa coNuchaumw mmwlwm "zvsum manumuomawu Hmcwvumu .NH maan APPENDIX C Effects of soil moisture levels and site on the growth and tissue nutrient levels of seedlings grown in a greenhouse experiment 82 Table 18. Effects of soil moisture levels and site on lodgepole pine seedling growth Mean top height (mm) Mean tOp weight (g) Site Moisture level Moist Dry Moist Dry 1’3 8‘“ 36.503’b 44.70a 0.089c 0.140a 2 Atm 35.15b 33.80b 0.102b 0.089C 10 Atm 35.80%"b 30.20b 0.081d 0.066c Mean length of longest root (mm) Mean root weight (g) Site Moisture level Moist Dry Moist Dry 1/3 Atm 183.05a 157.05b 0.1338 0.1848 2 Atm 164.253’b 147.351”c 0.116a 0.1133 10 Atm 174.455"b 128.55C 0.1058 0.1058 Treatments not having a common letter for each set of values are significantly different at the 5% level. 83 Table 19. Effects of soil moisture levels and site on hybrid pine seedling growth Mean top height (mm) Mean top weight (g) Site Mbisture level Moist Dry Moist Dry 1/3 Atm 57.253 51.953 0.155b 0.1608 2 Atm 35.80b 32.80b 0.086c 0.0748 10 Atm 36.70b 31.40b 0.080d 0.070e Mean length of longest root (mm) Mean root weight (g) Site Moisture level Mbist Dry Moist Dry 1/3 Arm 172.00a 180.208’b 0.1468 0.1038 2 Atm 156.55b 134.50c 0.1298 0.1078 10 Atm 167.05b 132.90c 0.1343 0.0818 Treatments not having a common letter for each set of values are significantly different at the 5% level. 84 Table 20. Effects of soil moisture levels and site on jack pine seedling growth Mean top height (mm) Mean top weight (g) Moisture level Moist Dry Moist Dry a a,b a b 1/3 Atm 50.40 45.80 0.172 0.130 2 Atm 39.401”c 36.20c 0.088d 0.0778 10 Atm 39.701“c 32.13C 0.107c 0.065f Mean length of longest root (mm) Mean root weight (g) Moisture level Moist Dry Moist Dry 1/3 Atm 163.808 162.058 0.1138 0.1308 2 Atm 164.208 149.158 0.074a 0.1553 10 Atm 158.058 142.138 0.0813 0.0848 Treatments not having a common letter for each set of values are significantly different at the 5% level. ll III,I III] II I [I I}! 85 Table 21. Effects of soil moisture levels and site on black spruce seedling growth Mean top height (mm) Mean top weight (3) Site Moisture level Moist Dry Moist Dry 1/3 Atm 42.60a 29.95b 0.040a 0.030b b c b c 2 Atm 28.85 18.72 0.029 0.016 10 Atm 33.93b 11.69c 0.033b 0.029b Mean length of longest root (mm) Mean root weight (g) Site Moisture level Moist Dry Moist Dry 1/3 Atm 126.70a 73.95b 0.0208 0.0163 2 Atm 112.83a 81.28b 0.0198 0.0143 10 Atm 126.87a 82.31b 0.033a 0.014a Treatments not having a common letter for each set of values are significantly different at the 5% level 86 .oanmaaoun: mum vocfiouno muasmom NO mmN.o mon.H mwN.o «mq.N mma.o mae.o wmc.o wmm.a ass 0H mHH.o mam.o Hem.o moN.N NNH.o Ham.o can.o mmm.a au< N monumm Hma.o mem.o mNm.o mum.N sma.o oa muaumfioz mofioomm muwm Nun muww umfioz AoHanNm>m NV GONumHuamocoo uGONuunz mcoaumuucmoaoo ucofiuunc wwmfiaom wcfiavmmm :0 mafia mam mam>oa muaumwoa HNom mo muommmm .NN maan 87 .oanmaaoua: one voawmuno muasmmm NO NNN.O N«O.N NON.O OHN.O «O0.0 No«.O aH< OH OO0.0 O«N.O NON.O «NN.O HHN.O NON.O aO< N weaken ONN.O OON.O ON«.O ONN.O OON.O OO«.O au< NOH OOOHO NO0.0 OOH.O «N0.0 ONN.O NN0.0 OON.O au< OH HO0.0 NHN.O «N«.O OON.O NO0.0 ON0.0 aOO N mafia NOH.O NON.O OO0.0 ONN.O OHN.O ONO.H OOO NOH NOON ON0.0 «H«.O OO0.0 NOH.O OON.O ON0.0 au< OH HO0.0 «NH.O OO0.0 NOH.O NON.O O«0.0 EON N mafia NNH.O HO«.O NOO.O NNN.O ON0.0 NON.O aO< NOH OHOONO HN0.0 NON.O OO0.0 O«N.O NNO.H OON.O EON OH ON0.0 HHN.O OO«.O OON.O OON.O «NN.O aO< N mafia OO0.0 OON.O OHN.O OON.O OO0.0 NON.O EON NNH OHOOOOOOH m2 mo M m w: mo M m Hm>mH unaumfioz mowowam Nm» NO OOOO Nun OOHO OOHoz AoHanNmpm NV cONumHucooaoo unmfiuusz mcoNumuucmocoo ucmfiuusa uoou mafiawmom so mufim mam mHm>wH onnumfloa HNom mo muommmm .mN mHan APPENDIX D Effects of soil fertility levels and site on the growth and tissue nutrient levels of seedlings grown in a greenhouse experiment llll.lI! i II ‘I Table 88 24. Effects of soil fertility levels and site on lodgepole pine seedling growth Mean top height (mm) Mean top weight (g) Site Fertility level Moist Dry Moist Dry. Control 40.75a 23.95b 0.145“:b 0.056c Low 38.65a 35.77a 0.125b 0.1488"b High 40.908 44.838 0.1611"b 0.191a Mean length of longest root (mm) Mean root weight (g) Site Fertility level Moist Dry Moist Dry Control 233.955"b 207.953:b 0.287“:b 0.136c Low 239.40a 227.71"“’b 0.182b’c 0.332a High 202.101“C 172.96c 0.2288:b 0.2818 b Treatments not having a common letter for each set of values are significantly different at the 5% level. 89 Table 25. Effects of soil fertility levels and site on hybrid pine seedling growth Mean top height (mm) Mean top weight (g) Site Fertility level Moist Dry Moist Dry Control 39.85a 24.75b 0.125b 0.043c Low 41.90a 46.25a 0.1345"b 0.1655"b High 50.02a 44.258 0.1783 0.1818 Mean length of longest root (mm) Mean root weight (g) Site Fertility level Moist Dry Mbist Dry Control 228.105“b 190.70“d 0.157b 0.110b Low 236.55a 210.298"c 0.149b 0.377a High 198.80b’c 162.49d 0.177b 0.361a Treatments not having a common letter for each set of values are significantly different at the 5% level. 11.1 117.! Ill-I'll 90 Table 26. Effects of soil fertility levels and site on jack pine seedling growth Mean top height (mm) Mean top weight (g) Site Fertility level Moist Dry Moist Dry Control 32.005"b 26.10b 0.091b 0.046b Low 38.853 39.708 0.1458 0.1658 High 36.508’b 52.053 0.1583 0.1883 Mean length of longest root (mm) Mean root weight (g) Site Fertility level Meist Dry Moist Dry Control 210.502:b 199.15":c 0.093b 0.101b Low 232.75a 218.858’b 0.181b 0.334a High 194.55b’C 168.25c 0.169b 0.403a Treatments not having a common significantly different at the 5% level. letter for each set of values are 91 Table 27. Effects of soil fertility levels and site on black spruce seedling growth Mean top height (mm) Mean top weight (g) Site Fertility level Moist Dry Moist Dry Control 37.301“C 22.85d 0.042a 0.016a Low 41.10b 27.79“d 0.0453 0.0308 High 55.25a 20.55d 0.0746 0.0178 Mean length of longest root (mm) Mean root weight (g) Site Fertility level Moist Dry Moist Dry Control 132.00a 94.25b 0.023a 0.012a Low 140.55a 96.10b 0.0168 0.0238 High 154.80a 87.55b 0.0333 0.0108 Treatments not having a common letter for each set of values are significantly different at the 5% level. 92 oHAoNHoua: mum voswmuno ouasmom NO HNN.O ON0.0 OON.H OHO.N NON.O «H0.0 OOO.H OOO.H OOHO OOH.O HN«.O HH0.0 OHO.H OOH.O NO0.0 ONN.H NON.H soH oosuan HOH.O OO0.0 NH0.0 NOO.H HOH.O NH0.0 NHN.H H«O.N HonoooO HOOHO ««N.O N«H.O O«N.H OON.H NNN.O OO0.0 NNO.H OON.H HOHO OOH.O «OH.O OO0.0 OON.H OOH.O HH«.O NHH.H O«O.H soH O NOH.O ONN.O HO0.0 OO«.H «OH.O OO«.O ON0.0 ONN.H HonuaoO “WHO N«N.O O«H.O OOO.H ONH.N OHN.O OO«.O NHO.N O«O.H OOHO «OH.O NON.O HO0.0 HNN.H HOH.O ONN.O NON.H OON.H soH o OOH.O NNN.O ON0.0 NON.H ««H.O OH«.O NO0.0 NNO.H HoNOooO OHMMMO NHN.O NOH.O «NO.H «NN.N HHN.O «H0.0 OHH.N NOO.H OOHO NOH.O OHN.O ON0.0 NNO.H «NH.O NO«.O OOO.H HON.H aoH O NNH.O OON.O NOO.H «NO.H ONH.O OO«.O N«0.0 ONO.H HonoooO OHoOOwwwH Oz no N NOO 2 Oz no N NON z HO>OH OOHOOOO NONHHONCM ouNm mun ouww umNoz AoapmHNm>m NV nONumuucooooo uaowuusz mGOHUMHuGOUCOU quHHuSC wamfifiow wCHHmem GO muwm USN mHm>mH %uHkuHmw HHOm MO muuwwwm .wN mHan 93 oHnmNHoua: mum vocamuno muHonoM NO OOH.O ONN.O ON0.0 NON.O HOO.H OON.H OOHO HNH.O ONN.O OO0.0 NOH.O «O0.0 OON.O soH mosses «NH.O ON0.0 NH0.0 ONH.O OOO.H HNN.O Houoooo HOOHO OOH.O NOH.O OO0.0 «ON.O O««.N ONO.H OOHO NOH.O «ON.O OO0.0 ONN.O ONO.H «O0.0 soH mafia OO0.0 HHN.O ON«.O ONN.O «OH.H ON0.0 HonoooO NOON ONO.O OOH.O NNO.O NON.O O«N.H HHO.H OOHO NHH.O ONH.O ON0.0 «ON.O «NN.O «NN.O soH mafia N«H.O OHN.O HO«.O «OH.O NO0.0 NNO.O HonOOoO OHHONO O«H.O ONN.O NNO.O OON.O NHO.N ONH.H OOHO. NOH.O O««.O NO0.0 N«N.O HOO.H NO0.0 soH mafia NNH.O NON.O HO«.O NHN.O O«0.0 ON0.0 HonOooO oHoOoOOoH w: mu M Amp wz mo M Amm Ho>oa mowoomm NOHHHOHOO OOHO ONO oOHn OOHoz AoHanNm>m NV cowumuucooaoo ucmwuunz meNumuusmuaoo uaofiuuss uoou chHvoom co mufim Cam mHo>mH Muwawuumm HHom mo muoowmm .mN manna APPENDIX E Soil surface temperature and moisture levels at the 1972 seeding location 94 Table 30. Soil surface temperature and moisture levels at the 1972 seeding location Moisture content Temperature C (F)* percent (by weight)** Date Moist site Dry site Moist site Dry site June 24 22.0 26 31.6 (89) 34.4 (94) 28 38.3 (101) 39.4 (103) 23.7 19.2 July 4 22.2 (72) 26.6 (80) 20.5 18.7 7 31.6 (89) 30.5 (87) 25.5 18.5 10 20.5 (69) 20.5 (69) 31.2 25.8 15 30.9 24.6 18 25.0 (77) 33.1 25.7 19 28.2 27.8 26 18.3 (65) 20.0 (68) 33.9 24.8 Aug. 6 28.3 (83) 26.6 (80) 29.5 16.1 10 35.8 20.9 18 21.6 (71) 21.0 (70) 36.7 30.4 21 44.3 29.7 25 30.7 26.3 29 23.3 (74) 31.6 (89) Sept. 4 27.5 20.3 * Soil surface temperature recorded by thermocouple. ** Soil moisture levels in upper 5 cm of soil determined gravimetrically. ill fill it! 1‘11 0'. HICHIGRN STQTE UNIV. LIBRQRIE 3102522012