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Effects of Trifolium-Rhizobium symbiosis on Pinus contorta regeneration, forest soil, and selected native… Trowbridge, R. L 1990

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EFFECTS OF TRIFOLIUM-RHIZOBIUM SYMBIOSIS ON PINUS CONTORTA REGENERATION, FOREST SOIL, AND SELECTED NATIVE PLANT SPECIES by R. L TROWBRIDGE Bachelor of Science, Colorado State University, 1970 A THESIS SUBMITTED IN PARTIAL FULFILMENT O F T H E REQUIREMENTS FOR T H E D E G R E E O F MASTER O F SC IENCE in T H E FACULTY O F G R A D U A T E STUDIES (Department of Forestry) We accept this thesis as conforming to the required standard T H E UNIVERSITY O F BRITISH COLUMBIA © R. L. TROWBRIDGE In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of F o r e s t r y  The University of British Columbia Vancouver, Canada Date March 26, 1990 DE-6 (2/88) ii ABSTRACT This study reports on the early effects of the Trifolium hybridum-Rhizobium symbiosis on Pinus contorta Doug, ex Loud (lodgepole pine), soil, and selected native plant species. Four rates of seeding (0, 10, 20, and 30 kg/ha) using inoculated Trifolium hybridum (alsike clover) seed were applied to three different site preparation treatments (broadcast burn, windrow burn, and mechanical scraping) using a split-plot design. Alsike clover and the Rhizobium inoculant were found to have excellent establishment and infectivity, and the symbiosis was assessed to be fixing nitrogen effectively. No effect of site preparation treatments was observed on establishment of the symbiosis, and clover-seeded plots averaged 76% cover by the end of the third growing season. The symbiosis had no significant (p < 0.05) effects on lodgepole pine total or incremental height or survival during the first three growing seasons, nor was there any observed effect on lodgepole pine foliar total nitrogen (N) concentration and 815N values at the end of the second growing season. Small, but significant (p < 0.05) decreases were observed for lodgepole pine total and incremental diameter in the second and third growing seasons, as well as needle mass in the second growing season. The growth decreases were probably attributable to the effect of shading by the clover cover. However, lodgepole pine seedlings overtopped the clover by the end of the third growing season and shade effects are likely to decrease as tree seedlings continue to grow. After one growing season, the symbiosis significantly (p < 0.05) increased mineralizable N in the forest floor and mineral (0-15 cm) soil layers. However, no significant changes in total N were detectable. The changes in mineralizable N were likely a measure of increased microbial biomass attributable to greater amounts of rhizosphere soil in clover-seeded plots compared to controls. Available phosphorus (P) in the forest floor significantly (p < 0.05) decreased as rate of seeding increased after one growing season. The decrease of forest floor available P may be attributed to greater assimilation of P in clover-seeded plots for plant and microbial growth, as well as the additional requirements for P in the supply of biological energy needed for active N2 fixation. All native plant species had low cover values which made interpretation of results difficult. However, percent cover of Calamagrostis canadensis, Rosa acicularis, and Spiraea betulifolia were iii significantly less in clover-seeded plots compared to controls at the end of the second growing season. Replacement of some herb and low-growing shrub species by legume-Rhizobium symbiosis may be desirable if the net result is an increase in site N without detrimental effects to tree crop species. It is recommended that the \egume-Rhizobium symbiosis be established in the early regeneration of lodgepole pine plantations on similar sites that are inherently N deficient and have experienced further site N depletion through forestry practices such as slashburning. iv T A B L E OF C O N T E N T S Page A B S T R A C T ii A C K N O W L E D G E M E N T S ix 1 INTRODUCTION 1 1.1 Background 2 1.2 Purpose and Objectives 2 1.3 Study Area 3 1.4 Experimental Design and Layout 5 2 E F F E C T S O F TRIFOLIUM-RHIZOBIUM SYMBIOSIS ON PINUS CONTORTA 8 2.1 Introduction 8 2.2 Methods 9 2.2.1 Study area and experimental design 9 2.2.2 Alsike clover and Rhizoblum 10 2.2.3 Lodgepole pine 11 2.3 Results and Discussion 12 2.3.1 Alsike clover-Rhizobium establishment 12 2.3.1.1 Infectivity and effectiveness of inoculant 12 2.3.1.2 Percent cover of alsike clover 13 2.3.1.3 Volume of alsike clover 16 2.3.2 Seedling growth and survival 17 2.3.2.1 Height and diameter increments 17 2.3.2.2 Survival 21 2.3.3 Seedling needle mass and nitrogen status 21 2.4 Conclusions 24 2.5 Summary 25 3 E F F E C T S O F TRIFOLIUM-RHIZOBIUM SYMBIOSIS ON SOIL . 26 3.1 Introduction 26 3.2 Methods 27 3.2.1 Study area and experimental design 27 3.2.2 Field sampling and preparation 27 3.2.3 Laboratory analysis 29 3.3 Results and Discussion 29 3.3.1 Nitrogen and carbon 33 3.3.2 Phosphorus 38 3.3.3 Cations, pH, and C E C 40 3.4 Conclusions 41 4 E F F E C T S O F TRIFOLIUM-RHIZOBIUM SYMBIOSIS ON NATIVE VEGETAT ION 42 4.1 Introduction 42 4.2 Methods 43 4.2.1 Study area and experimental design 43 4.2.2 Field sampling, measurement, and collection 43 4.3 Results and Discussion 44 4.3.1 Herbs 48 4.3.2 Shrubs 52 4.4 Conclusions 53 5 MANAGEMENT AND R E S E A R C H RECOMMENDATIONS 55 5.1 Management Considerations 55 5.2 Management Recommendations 55 5.3 Research Recommendations - 56 vi 6 L ITERATURE CITED 58 7 APPENDICES 64 Appendix 1 Summary of statistical analysis for Pinus contorta and Trifolium hybridum data 64 Appendix 2 Summary of statistical analysis for forest floor chemistry data 85 Appendix 3 Summary of statistical analysis for mineral soil chemistry data 116 Appendix 4 Summary of statistical analysis for native vegetation data 146 Appendix 5 List of plant species observed on the experimental site . . . 174 vii LIST OF T A B L E S Page 1. Analysis of variance table 5 2. Response of alsike clover and lodgepole pine seedlings to levels of N-treatments 14 3. Response of alsike clover and lodgepole pine seedlings to site preparation treatments 15 4. Soil nutrient content (kg/ha), cation exchange capacity (CEC), and pH pre-treatment, one year post-treatment, with pre- to one year post-treatment difference 30 5. Mineral soil bulk densities and forest floor mass estimates 32 6. Mineralizable N as a percent of total N for the different levels of N-treatment one year post-treatment 37 7. Percent cover of Trifolium hybridum and selected native species at the end of the first (1987) and second (1988) growing seasons 46 viii LIST OF F IGURES Page 1. Location of study area 4 2. Field layout of the experiment 6 3. Rhlzoblum infection causing typical abundant nodulation on roots of alsike clover 12 4. Clover percent cover by year . . . 16 5. Height increment in the 2nd and 3rd growing seasons 17 6. Diameter increment in the 2nd and 3rd growing seasons 18 7. Linear relationship of diameter increment as a function of clover percent cover on 2- and 3-year-old lodgepole pine seedlings 19 8. The interaction of N-treatment and site preparation on needle mass 22 9. The linear relationship of needle mass as a function of percent cover clover on and 2-year-old lodgepole pine needle mass 23 10. Mineralizable N in the forest floor and mineral soil layer one year post-treatment 33 11. Changes in forest floor P one year post-treatment 39 12. Percent cover of Calamagrostis canadensis at the end of the first two growing seasons 48 13. Site preparation x block interaction for Linnaea borealis in the first growing season 50 14. Site preparation x block interactions for Cornus canadensis in the first two growing seasons 51 15. Percent cover of Rosa acicularis at the end of the second growing season 52 16. Change in percent cover of Spiraea betulifolia from 1987 to 1988 53 17. Lodgepole pine seedlings and alsike clover in the third (1989) growing season 54 ix ACKNOWLEDGEMENTS I would like to thank my thesis supervisor, Dr. M.C. Feller, and committee members Drs. F.B. Holl, K. Klinka, and Mr. B. Downie for their individual and collective contributions towards the completion of this thesis. Dr. F.B. Holl deserves special recognition for his advice and assistance in all laboratory and field procedures relating to the legume-Rhizobium symbiosis. Dr. G. Eaton and Ms. W. Bergerud gave valuable assistance with experimental design and statistical analysis. The field assistance of B. Blackwell, M. Lavigne, A. Macadam, and S. Thomson were much appreciated. The patience and understanding of my sons, Aaron and Erik Trowbridge, were also appreciated. The Prince Rupert Forest Region, and in particular Mr. B. Downie, deserve recognition for their support throughout this study, and in granting me an educational leave of absence during 1986-87. Financial support was provided by the British Columbia Ministry of Forests, the Northern Interior Technical Advisory Committee of the Canada-British Columbia Forest Resource Development Agreement (1985-90), and a block grant administered by the Faculty of Forestry, University of British Columbia. I 1 INTRODUCTION Nitrogen is the principal nutrient limiting productivity of agricultural and forest crops in temperate North America. It is also readily lost from soils as a consequence of volatilization, leaching, denitrification, runoff and erosion, as well as through biomass removal during harvesting of the crop. In natural ecosystems, nitrogen inputs may be derived from small amounts deposited in rain water.and by living organisms through biological nitrogen fixation. On low nitrogen status soils (such as forest soils following severe fire, or severely disturbed and exposed mineral soils) recolonization by nitrogen-fixing organisms comprises the primary means of restoring nitrogen fertility over time (Burns and Hardy 1975). In forest and woodland ecosystems, it has been estimated that approximately 40 x 10s t of nitrogen are fixed annually by leguminous trees and woody actinorhizal plants throughout the world (Burns and Hardy 1975). In intensive agriculture and forestry, demands for increased productivity have been met by the addition of fertilizer nitrogen derived from fossil fuels. As interest in more intensive management of nitrogen fertility in forest ecosystems has increased, so have general concerns over the cost of forest fertilization and the consequences of extensive synthetic fertilizer use on soil and ground water quality (Pritchett 1979). Although immediate benefits from fertilization can sometimes be predicted from agricultural experience and short-term forestry experiments, it may be more appropriate for foresters to consider the long-term management questions: Can, or will, the productivity of our forest lands be maintained? How might long-term fertility needs best be met? The use of nitrogen-fixing organisms to enhance soil fertility during reforestation may be more appropriate than synthetic fertilizers to the natural time scale of forest regeneration. Legume species have been used in reforestation and related research projects in central Europe, Australia, New Zealand, and the USA (Jorgensen 1980). The primary function of the species has been to ameliorate soil nitrogen deficiencies via biological nitrogen fixation and, through eventual nutrient cycling, to supply the associated tree crop with a form of long-term, slow release available nitrogen. Other benefits of using herbaceous legumes in forestry may include: an economic return in forage where integrated multiple use is practised; reduction of vegetation that competes with crop trees; and the establishment of a more suitable micro-climate for tree seedling regeneration and growth. 2 The actual results and benefits from legume use, however, may vary widely. This variability is primarily a consequence of the plant-endophyte combinations used and their interaction with site, climate, and soil conditions. Satisfactory application of biological nitrogen fixation technology will require the appropriate use of selected legumes and compatible Rhizobium inoculant. 1.1 Background Prescribed fire is used to modify forest and range sites to meet specific management objectives. Prescribed fire is used for many purposes, but the impacts of slashbuming on site productivity are not clearly defined and were considered by local foresters to merit a high research priority. In response to this perceived need, a research program on the effects of slashbuming was initiated in 1980 in central British Columbia (Macadam and Trowbridge 1984). Since 1980 several experimental projects have been established and some preliminary measurements of site effects have been made. One of the consistent observations has been a net loss in total soil nitrogen (N) following fire treatments (Macadam 1987, Taylor 1987, Blackwell 1989). In conjunction with these results, fertilization screening trials indicate foliar N deficiencies on similar sites in all Age Class 3 stands of Pinus contorta Doug, ex Loud (lodgepole pine) diagnosed to date in the Prince Rupert Forest Region (Yole 1986, Carter 1989, Yole et al. 1989). Such results lead the British Columbia Ministry of Forests to anticipate N deficiencies in some new regenerating stands, particularly those that have undergone site preparation resulting in net N losses. 1.2 Purpose and Objectives Nitrogen loss following prescribed burning was anticipated to occur and has been confirmed by Blackwell (1989), in a recent study of the rehabilitation of repressed lodgepole pine stands. The present study was superimposed on selected areas of that large scale rehabilitation site, primarily to examine the amelioration of any negative consequences of burning and scraping on site nutrient-N status. 3 Specif ica l ly, the object ives of this study were : i. to se lect a n appropr iate nitrogen-fixing l egume spec ie s and inoculant strain combinat ion for site amel iorat ion t reatments in the study a rea ; ii. to a s s e s s the estab l i shment of the l egume spec ie s and to a s s e s s the ef fect iveness of the symbios i s ; iii. to determine the impact of different rates of l egume spec i e s seed ing ( 'nitrogen' treatments) on lodgepo le p ine seed l ing growth, survival, and foliar N status; iv. to determine the impact of these t reatments on se lec ted soi l c hem ica l propert ies in the upper soi l profile; and v. to determine the impact of these treatments o n se lec ted native plant spec ie s . 1.3 Study A r e a T h e study a r e a occu r s within the Lake s Forest District and is located north of Wistar ia, be tween F ranco i s and O o t s a L a k e s (Figure 1). T h e a rea c h o s e n for the exper iment occu r s within o n e dominant e c o s y s t e m - the mes i c bunchber ry -moss site ser ies of the Moist C o l d Sub -borea l S p ruce ( SBS ) b iogeoc l imat ic s ubzone (Pojar ef al. 1984). Th i s z o n e w a s cha rac te r i zed a s c o l d sub -borea l cont inental humid type, with severe, snowy winters and relatively wa rm, moist, a n d short s ummer s (Pojar et al. 1984). So i l s are dominant ly Bruniso l ic G ray Luv i so l s (Agriculture C a n a d a Expert Commi t tee on So i l Su rvey 1987) deve l oped o n mora ina l b lankets of d e e p (> 1 m) g lac ia l till. T h e soi ls are l oam to c lay loam, with 2 0 - 3 5 % coa r se fragments. T h e site is gently to moderate ly s lop ing with south to southeast aspect. A l though the site is genera l ly we l l d ra ined, there are imperfectly d ra ined depre s s i on s and rece iv ing a reas accumulat ing sur face runoff in the spr ing and after precipitat ion. C o m m o n plant s p e c i e s o n the site inc luded: Arnica cordifolia (heart- leaved arnica), Cornus canadensis (Canadian bunchberry), Epilobium angustifolium (fireweed), Calamagrostis canadensis (bluejoint), Rosa acicularis (prickly rose), and Spiraea betulifolia (b irch- leaved spirea). A list of plant s pec i e s ob se r ved o n the site is found in Append i x 5. FIGURE 1. Location of study area. 5 1.4 Experimental Design and Layout The experiment was superimposed on the lodgepole pine rehabilitation study of Blackwell et al. (1986), using a split-plot design (G. Eaton, Professor, UBC; and W. Bergerud, Biometrician, B.C. Ministry of Forests, pers. comm., 1987). Main plots were site preparation techniques, and the split-plots were the nitrogen-fixing treatments (Figure 2). There were three site preparation treatments and four nitrogen-fixing treatments. The nitrogen-fixing treatments were replicated in three rows in each of three blocks (blocks were considered to be a fixed effect). Therefore, the factorial experiment included: blocks (3) x rows in blocks (3) x site preparation techniques (3) x nitrogen-fixing treatments (4) = 108 plots. Table 1 shows the analysis of variance (ANOVA) sources and associated degrees of freedom. T A B L E 1. Analysis of variance table. Source df Block (B) 2 Site preparation (SP) 2 B x SP 4 Error A (B x Sp)Row-1 18 Nitrogen-fixing treatment (N) 3* N x B 6 N x SP 6 N X B x SP 12 Error B (residual) 54 Total 107 * Planned contrasts for Nitrogen-fixing treatment included: linear, quadratic, cubic, and control vs others -7 5 5 6 6 5 if IK w U " 02 • > as Us • J a - ••> • i a. a. a . 0. a . D' O s 1)2 O2 Os a. / 2 j s e s Q< as Di Os a « D« a« a i Q2 0'- as C l T l 5 3 B * D< • i D s a - O i • i • J O s C 2 T 2 L 2 / a 9 3 a e 0« D' 0< as 0 ' 0 ! o* as 0' 02 0; T 3 C 3 L 3 7 7 3 3 9 9 5 w 5 W -S" W 0- as Os Os as Os Os a« a* 0" 04 0' 0' 02 0 ' 02 Qi • « 0 - 0' 02 0' D2 Q2 LEGEND Example: / = Row No. {/ • 9 J W = Site Preporolion Treatment Q i = 0 kq/ha 0* = 10 kg/ha Os = 20 kg/ho 0 4 * 30 ka/ha IV = windrow burn S - mechanical scrape B * broadcast burn Note: T l , C ) , L I etc. designate plots of related experiment. Figure 2. Field layout of the experiment. 7 The site preparation techniques were broadcast burning, windrow burning, and mechanical scraping (areas between windrows). The rate of seeding levels in Nitrogen-fixing treatment (N-treatment) were 0, 10, 20, and 30 kg/ha of limed and inoculated Trifolium hybridum (alsike clover) seed. These latter treatments represented control (no seed), moderate, high, and very high rates of seeding in operational forestry practices for range forage seeding in clearcuts. The treatment effects were tested using SAS (SAS Institute Inc. 1985) 'General Linear Model' and 'Regression' procedures with statements to use correct error terms, Tukey's Studentized Range Test (HSD) and planned contrasts. Source effects were assumed to be significant at p < 0.05. Summaries of pertinent statistical analyses are found in Appendices 1 to 4. 2 E F F E C T S O F TRIFOLIUM HYBRIDUM-RHIZOBIUM SYMBIOSIS ON PINUS CONTORTA 8 2.1 Introduction Approximately 80% of the atmosphere surrounding the earth consists of nitrogen gas (N2). The capability to convert N2 gas into a usable form [e.g., ammonium (NH/) nitrogen] is universally restricted to prokaryotic organisms. In one large family of plants (Fabaceae, formerly Leguminoseae) a unique relationship has evolved with a class of microorganisms, which permits many of these plants to exploit the gaseous N2 resource. Management of biological nitrogen fixation in forestry has been considered for many years throughout the world (Assmann 1970, Beuter 1979, Haines and DeBell 1979, Rehfuess 1979, Jorgensen 1980, Granhall 1981, Fortin era/. 1984, Binkley 1986), and more recently in British Columbia (Hermansen 1976, Eichel 1979, Fahlman 1981, Kibbey et al. 1981, Carr and Ballard 1980, Beese and Kumi 1985, Croockewit and Trowbridge 1985, Trowbridge and Holl 1989). Representatives of the legume family can be infected through their roots by soil microorganisms of the genus Rhizobium. Within the root nodules that are formed as a result of this infection process, the plant and bacteria develop the complex biological machinery to convert N2 into NH/ for use by the plant. This complex symbiosis depends on the specific interaction of the legume host and Rhizobium species, and may be influenced by environmental conditions (Turvey and Smethurst 1983). The newly fixed nitrogen becomes available to other organisms once the plant tissues die and are incorporated into the soil. Small amounts of available nitrogen may be excreted from the root nodules directly into the soil (Russell 1973). Since most legumes are shade intolerant (Haines and DeBell 1979), the nitrogen fixation will mainly occur in the early stages of plantation development before canopy closure (15-30 years in interior British Columbia). During that time, 50-100 kg/ha/yr nitrogen should ideally be fixed (Jorgensen 1980). This newly fixed nitrogen may be used by any number of other organisms, released to the atmosphere through denitrification, or leached from the site. The primary interest of the legume-Rhizobium symbiosis in forestry has been directed to amelioration of soil nitrogen due to the inherent low nitrogen status of many forest ecosystems, or due to nitrogen losses caused by some forest management practices (Haines and DeBell 1979, Fort in et al. 1984). Before that amelioration can be achieved, the legume and tree seedling crops must be established in a regime that is compatible with forest management objectives. Haines and DeBell (1979) suggested several cropping systems for forest management. The current study is comparable to' their 'mixed species systems' - a nitrogen-fixing species beneath the overstory of a non-fixing commercial tree species for part of the rotation. There has not been an abundance of published literature regarding the effects of legumes on tree growth in North America. Fortin et al. (1984) found an absence of data reporting the effects of legumes on tree growth, most reports found were based on experiments not designed for that specific purpose. However, what literature is available overwhelmingly reports positive effects on tree growth. This section of the thesis describes the establishment of the alsike clover-Rhizobium symbiosis and it's effect on early growth, survival, and foliar nitrogen status of lodgepole pine. 2 . 2 Methods 2 .2 .1 s t u d y area a n d experimental d e s i g n The study area and experimental design have been described in Sections 1.3 and 1.4. Within the study site, the experiment was established incorporating three site preparation techniques. Each of these, broadcast burn, windrow burn, and scrape (areas between windrow), was split into four 20 m 2 plots ( 4 x 5 m), replicated in three rows in each of three blocks. These split-plots were randomly assigned one of four seeding rate treatments (N-treatments). The N-treatments consisted of 0 (control), 10, 20, and 30 kg/ha of inoculated alsike clover seed. There were 108 split-plots in total (see Figure 1, page 6). 10 2.2.2 Alsike clover and Rhlzoblum The selection of alsike clover was based on information available from local experience in the study area and recommendations based on the acid and drought tolerance of commercial cultivars. Since Rhizobium strains are known to have different infectivity and effectivity responses based on environmental conditions (Turvey and Smethurst 1983), nodules from alsike clover plants found in the study area were collected, and the endophyte isolated and tested for infectivity and effectivity in a pot experiment (Trowbridge 1987a) using the basic techniques described in Vincent (1970). Six individual colonies of effective Rhizobium leguminosarum biovar trifolii were ultimately selected and cultured for use in the experimental inoculant. Strains were cultured in yeast extract mannitol broth and combined prior to addition to packages of sterile ground peat to prepare the inoculant (Vincent 1970). Seed was weighed to correspond to the treatments, then each split-plot allocation of seed was coated with the peat inoculant and limed using procedures described in Trowbridge and Holl (1989). Seeding took place one day later, in early June 1987. The alsike clover and Rhizobium infection was assessed at the peak of the first three growing seasons (mid-August 1987, 1988, and 1989). Twelve 900 cm 2 (30 x 30 cm) subplots within a split-plot (centred at each seedling location) were assessed for percent cover and average height of the alsike clover (1296 samples in total) in 1987 and 1988. In 1989, one estimate of percent cover was obtained for each of the 108 split-plots. At the time of the preceding assessments, individual plants were excavated in each split-plot, the roots gently agitated in water, and then visually assessed for nodulation abundance and pigmentation. Nodule abundance was placed into one of four categories: none, few, many, and abundant. Nodules from the roots of these plants were cut open to assess if pink pigmentation was present. The pink pigmentation is due to the presence of leghemoglobin, a protein required during active nitrogen fixation in legumes, which indicates an effective symbiosis. Pigmentation was assessed as present or absent. 2.2.3 Lodgepole pine Twelve lodgepole pine seedlings (2-11 plugs) were planted in early June, 1987 in each split-plot at 1 x 1 m spacing prior to alsike clover seeding. Height and diameter of all seedlings were measured at the time of planting and later at the same time as the alsike clover assessments in each year. The difference between the 1987-88 and 1988-89 growing seasons was calculated as incremental growth, N-status was assessed in the second growing season (1988) by comparing treatment effects in needle tissue chemistry (total N and 1 5N). Natural abundance of 1 5N has been used to quantify N 2-fixation (Knowles 1980, Rennie and Rennie 1983, Hoefs 1987, Virginia et al. 1989) and is expressed as a 8 1 5N value calculated from the following equation: (1 5N/1 4N)sample - (1 5N/1 4N)atmosphere 8 1 5N = X 1000 (1 5N/1 4 N)atmosphere One lateral branch from the second whorl of each surviving tree was clipped and bulked with those from all other surviving trees per split-plot to yield a composite sample for each split-plot. The needles were oven dried at 70° C to determine needle mass (g dry weight/100 needles) by averaging the masses of three subsamples of 100 needles each for each split-plot. The remaining needles were analyzed for total N (%) (CHN-analyzer, Carlo Erba Model 1106) and 1 5 N (mass spectrometer, V G Isogas Ltd. Model PRISM) at the Department of Oceanography, University of British Columbia. 12 2.3 Resul ts and D i s c u s s i o n 2.3.1 A l s i k e clover-Rhizobium establ ishment 2.3.1.1 Infectivity a n d effect iveness of Inoculant Both infectivity a nd effectivity we re a s s e s s e d a s excel lent. In the samp l i ng s c h e m e , infection w a s to be a s s e s s e d by as s i gn ing one of four c l a s s e s of nodulat ion (ranging f rom none to very abundant), however nodulat ion w a s very abundant o n all s a m p l e s in all year s . (F igure 3). P ink p igmentat ion w a s ob se r ved in e a c h nodule cut o p e n in e a c h year. F igure 3. Rhizobium infection cau s i ng typical abundant nodulat ion on roots of a l s ike c lover. 13 2.3.1.2 Percent cover of a ls ike c lover The mean percent cover estimates for levels of N-treatment are shown in Table 2. In the first growing season there were many individual plants in each plot, although percent cover was very low (2-5%). There was a large increase in percent cover in the following years, ranging from 34 to 4 2 % in 1988 and 74 to 7 8 % in 1989 (Figure 4). There was no evidence of clover plants in the control plots in any year, although it was observed that re-seeding took place in the seeded plots in 1989. It was also observed that not all clover plants die back during the winter. To confirm this, during December 1987 snow was removed (and replaced) from selected plots. Clover cover of leaves and stems that remained green in colour at that time was estimated at 15%. The percent cover linear effect for N-treatment was highly significant (p < .01) in each year. However, the actual differences among the seeding rates were only a few percent in each year. It appeared that seeding rate made little real difference in percent cover - by the third growing season clover occupied a large amount of space in the plots regardless of increased seeding rate. This was further supported by the fact that means among the seeded plots could not be separated by Tukey's Studentized Range (HSD) test (Table 2). There was no significant effect of site preparation treatment in either year (Table 3). Likewise, there were no significant interactions in any year. Table 2. R e s p o n s e of a ls ike c lover and lodgepole p ine seedl ings to levels of N-treatment. 0 Seed rate (kg/ha): 10 20 30 Linear contrast: (D<0.051 Alsike clover: cover (%) 1987 1988 1989 Ob Oc Ob 2ab 34b 74a 3ab 38ab 76a 5a 42a 78a yes yes yes volume (m3/m2) 1987 1988 0.0 0.0b 0.001 0.084a 0.002 0.096a 0.005 0.108a yes yes Tree seedlings: Total height (cm) 1987 1988 1989 16.8 25.5 43.2 16.2 24.3 42.7 16.6 24.4 43.3 16.3 24.8 43.0 no no no Height increment (cm) 1987- 88 8.7 1988- 89 17.8 8.2 18.3 7.7 18.9 8.5 18.1 no no Total Diameter (mm) 1987 3.5 1988 6.7a 1989 11.3a 3.6 6.4ab 10.0b 3.6 6.3ab 10.1b 3.5 6.3b 9.8b no yes yes Diameter increment (mm) 1987- 88 3.1a 1988- 89 4.6a 2.8ab 3.6b 2.7b 3.8b 2.7b 3.6b yes yes Survival (%) 1988 1989 80 78 83 79 81 77 82 79 no no Needle mass (g/100) 1988 1.05a 0.93b 0.89b 0.90b yes Foliar N (%) 1988 1.46 1.47 1.43 1.49 no 8 1 5N 1988 9.13 9.10 8.97 8.81 no Note: means in the same line followed by the same letter are not significantly different (p < 0.05) using Tukey's Studentized Range (HSD) test. 15 Table 3. R e s p o n s e of a ls ike c lover and lodgepole pine seedl ings to site preparation treatments. Site preparation: Broadcast Scrape Windrow Alsike clover cover (%) 1987 1988 1989 1 25 56 3 31 54 3 29 61 volume (m3/m2) 1987 1988 0.000 0.063 0.002 0.075 0.005 0.079 Tree seedlinqs Total height (cm) 1987 1988 1989 17.2a 24.9 42.8 17.1a 25.7 42.1 15.1b 23.7 44.3 Height increment (cm) 1987- 88 1988- 89 7.7 17.9 8.6 16.4 8.5 20.6 Total Diameter (mm) 1987 1988 1989 3.5 6.4 10.5 3.6 6.4 10.1 3.6 6.5 10.4 Diameter increment (mm) 1987- 88 1988- 89 2.8 4.1 2.8 3.7 2.9 3.9 Survival (%) 1988 1989 79b 79ab 90a 86a 74b 69b Needle mass (g/100) 1988 0.96 0.89 0.95 Foliar N (%) 1988 1.36b 1.55a 1.47ab 8 1 sN(ppt) 1988 9.87a 8.07b 9.07ab Note: means in the same line followed by the same letter are not significantly different (p < 0.05) using Tukey's Studentized Range (HSD) test. 16 Clover % cover 100 r Year 1 Year 2 Year 3 Seed rate (kg/ha) H o E 2 i o W20 ^ 3 0 F igu re 4. C l o v e r percent c o v e r by year. 2.3.1.3 Volume of alsike clover V o l u m e es t imates (m 3/m 2) of a l s ike c l ove r b e t w e e n N-treatments are s h o w n in Tab l e 2. V o l u m e w a s ant ic ipated to be more sens i t ive to ac tua l d i f fe rences in b i o m a s s of c l o ve r p roduced than percent cove r . A l t hough percent cove r may b e the s a m e b e t w e e n plots, if the a ve rage heights were different then the ' total ' amount (volume) of c l o ve r w o u l d b e different. Howeve r , the da ta in Tab l e 2 s h o w that this a s sumpt i on is not neces sa r i l y the c a s e . Tukey ' s S t uden t i z ed R a n g e (HSD) test s h o w e d a s ignif icant d i f ference only b e t w e e n contro l and the rema in ing t reatments in the s e c o n d g row ing s e a s o n ( compare to Tab l e 2, percent c o v e r d i f ferences) , a l though the l inear effect for v o l ume of c l o ve r for the t reatments w a s highly s ignif icant (p < 0.001) in e a c h y ea r mea su red . T h e s e d a t a s u gge s t ed that there w a s no r ea son to c h o o s e v o l u m e ove r percent c o v e r es t imates to s epa ra te product ion be tween t reatments, so that vo l ume w a s not e s t imated in 1989. Fo r c l ove r vo l ume est imates, there w e r e no s ignif icant d i f fe rences b e t w e e n s ite preparat ion t reatments in e i ther y e a r (Table 3), nor w e r e there any signif icant interact ions. 17 2.3.2 Seedling growth and survival 2.3.2.1 Height and diameter There were no significant differences of N-treatment for total or incremental height in any year (Table 2). The overall total height of seedlings at the end of the 1989 growing season was 43 cm, while incremental growth was 8.3 cm for 1987-88 and 18.3 cm in 1988-89 (Figure 5). In most cases in 1989, current leader growth of seedlings was slightly above that of the maximum clover height. It is assumed that the seedlings have surpassed height growth of clover and will, in succeeding years, partially shade the clover. There were no significant post-treatment site preparation effects for height estimates, nor were there any interactions in any year. In addition, none of the planned contrasts were significant for total or incremental height (Table 3). Height i n c rement ( cm) 2 0 , - ^ '-2nd season 3rd season Seed rate ( k g / h a ) o E 3 i o W20 S 3 30 Figure 5. Height increment in the 2nd and 3rd growing seasons. The re were , however , s ignif icant effects of N-treatment o n total a nd incrementa l d i ameter a s s h o w n in T a b l e 2. E v e n though the effect on d iameter w a s s ignif icant, the actua l mea su remen t d i f fe rences w e r e very sma l l , thus render ing any b io log ica l o r pract ica l interpretation with caut ion. In relative va lue s , this effect r ep re sen ted a 1 3 % d e c r e a s e (0.4 mm) in d i amete r increment be tween no c love r and 3 4 - 4 2 % c l ove r c o v e r in the s e c o n d growing s e a s o n . In the third g rowing s e a s o n , at wh i ch t ime the c love r c o v e r r anged 7 4 - 7 8 % , the increment d i f fe rence rep re sented a 2 0 % d e c r e a s e (0.9 mm) be tween control a nd al l other c lover plots (F igure 6). Diameter i n c rement ( m m ) 4 -2 -2nd season 3rd season Seed rate ( k g / h a ) 0 ^ 1 0 WlO ^ 3 0 F igure 6. D iameter increment in the 2nd and 3rd g rowing s e a s o n s . 19 Clover percent cover had a significant linear effect for N-treatment, so that regression analysis was thought to be appropriate to determine if there was a significant relationship to diameter growth. The linear regression analysis for percent cover on diameter increment was highly significant (p < 0.01) in each year, but the r2 values were only 0.21 and 0.11 in 1987-88 and 1988-89 respectively. Despite the significant linear effect, only the control is strongly separated from the other N-treatments using Tukey's Studentized Range (HSD) test. The regression equations of diameter increment (Dl) as a function of clover percent cover (%CC) were: Dl = 3.38 - 0.02(%CC) for 1987-88, and Dl = 4.71 - 0.01 (%CC) for 1988-89. The regression lines are shown in Figure 7. D iameter i n c rement ( m m 8 6 4 2 --1 9 8 8 - 8 9 ^ * 1987 -88 —* i i i i i i i i i 0 10 20 30 40 50 60 70 80 90 100 Clover cover (%) Figure 7. Linear relationship of diameter increment as a function of clover percent cover on 2- and 3-year-old lodgepole pine seedlings. 20 No literature was found describing a comparable investigation between alsike clover and lodgepole pine. Hamilton (Plant ecologist, B.C. Ministry of Forests, pers. comm., 1989) recently found significant negative effects of vegetative shading on both seedling height and diameter increments in one-year old Picea glauca (Moench) Voss (white spruce) seedlings in the same biogeoclimatic zone, but in wetter and richer sites. Whereas white spruce is well known to begin growing slowly in interior British Columbia (Binder et al. 1989, Macadam 1989) , lodgepole pine generally grows comparatively faster in similar environments (Minore 1979). In this study, tree seedlings may have been responding to shading competition from clover by allocating its energy (physiological response) to increased height rather than diameter increment. In plots with no clover (virtually no competitive shading), the tree growth response may be balanced between increased height and diameter. The clover has reached its normal maximum height and will not likely increase shading given that lodgepole pine seedlings will continue to gain height increment and produce new whorls above the current shading level, thus any shading effects will likely reverse. Literature found on the use of legumes in forestry is overwhelmingly supportive of the observation of increased growth of associated trees. Finn (1953) found increased height and diameter growth of certain deciduous trees on all but one of his sites when the trees were associated with Robinia pseudoacacia L. (black locust). Haines et al. (1978) found increased height and volume (two- and threefold respectively) growth of Platanus occidentalis L. (sycamore) following establishment of clovers and vetch. Assmann (1970) summarized a study by Wiltch (1954) demonstrating increased height of Pinus sylvestris L. (Scots pine) associated with lupine (species not identified) resulting in 6 0 % volume increment over a 10-year period. Rehfuess (1979), in examining these results, reported overall growth reduced in lupine treatments for the first three or four growing seasons followed by improved growth after an initial lag period (3-7 years). The Scots pine experiments are confounded, in part, by simultaneous cultivation and other nutrient amendments. Kumi (1986) reported no overall significant effect of associated legumes on juvenile height growth of Pseudotsuga menziesii (Mirb.) Franco (Douglas-fir) except on poor sites, where there was a 1 3 % increase. 2.3.2.2 Survival N-treatment had no effect on survival (Table 2), nor were there any interactions with site preparation. Overall survival across N-treatment was 8 2 % in 1988 and 7 8 % in 1989. There were, however, significant effects of site preparation on survival - the scrape treatment resulted in higher survival than either broadcast or windrow burning in the first two post-treatment years (Table 3). Blackwell (1989) suggested that this effect is a response to increased soil temperature due to removal of forest floor in the scrape treatment, compared to broadcast burned areas. In addition, Blackwell (1989) suggested that some trees planted in windrowed areas were subjected to moisture stress and poor planting substrate. 2.3.3 Seedl ing needle m a s s and nitrogen status Needle mass showed a highly significant linear effect (p < 0.01) for N-treatment, however there was a weak site preparation by N-treatment interaction (p < 0.05). This interaction confounded the significant main effects since they were shown not to be totally independent. However, since N-treatment was so much more significant, some interpretation and discussion is warranted (Petersen 1985). The effect of windrow burning did not follow a similar response to the broadcast and scraping treatments (Figure 8). Windrow burning had a greater needle mass in the control, but lower or similar mass in the seeded plots. However, there was no main effect of site preparation on needle mass. No explanation was found to interpret the interaction result, although it may be attributable to experimental error. 22 Needle mass (g dry mass/100 needles) 0.5 1 ' 1 1 0 10 20 30 Nitrogen treatments (kg/ha seed) ~ — B r o a d c a s t b u r n S c r a p e - * - W i n d r o w F igure 8. T h e interact ion of N-treatment and site preparat ion o n need le mas s . T h e separat ion of need le m a s s m e a n s be tween leve l s of N-treatment cou ld on ly b e s h o w n be tween contro l a n d al l c l o ve r plots, not be tween the s eed i n g rates t hemse l ve s (Table 2). T h e l inear reg res s ion of need le m a s s a s a funct ion of % C C w a s highly s ignif icant but, a s with inc rementa l d iameter , the r 2 va lue w a s relatively low (0.34) (F igure 9). T h e regres s ion equat ion wa s : need le m a s s = 1.307 - 0.0071 ( % C C ) . 23 Needle mass (g mass/100 needles) 0 20 40 60 80 100 Percent cover clover 1988 • A c t u a l value P r e d i c t e d va lue F igure 9. T h e l inear re lat ionsh ip of need le m a s s a s a funct ion of percent c o v e r c lover o n 2-year-o ld lodgepo le p ine need le mas s . A s s h o w n in T a b l e 2, there w a s no N-treatment effect for e i ther total fo l iar N or 5 1 5 N, or interact ions b e t w e e n N-treatment a n d site preparat ion. Howeve r , e a c h var iab le s h o w e d signif icant s ite preparat ion effects. Tota l fol iar N w a s greatest in the s c r a p e a n d w ind row treatments, a n d least in b roadcas t burn t reatments (Table 3). Th i s result may be e xp l a i ned by so i l nutrient d i f ferences in the minera l so i l rooting z o n e . S c r ap i n g and w indrow burn ing resu l ted in g reater va lues of both minera l so i l N and minera l i zab le N (Chapter 3, T a b l e 4). Thus , it is l ikely that the higher so i l N status w o u l d result in h igher fol iar N Values. A c co rd i n g to J o r g e n s e n (1980), t rees initially r e spond to N f i xed by l e gumes by increas ing fol iar N. T h e N-treatments in the first post-treatment yea r of this s tudy h a d no s ignif icant effect o n lodgepo le pine fol iar N status b a s e d o n both m e a s u r e m e n t s - total fol iar N a n d 8 1 5 N . In this study, total N fol iar concent ra t ion w a s u s e d to separa te the N status a lone, w h e r e a s 8 1 5 N w a s u s e d to support e v i dence regard ing the transfer of newly f ixed-n itrogen f rom the s ymb io s i s to the seed l ing . T h e foliar N results are cons i stent with those reported by Kum i (1986) in wh i ch it w a s sugges ted that the lack of effect may be a c o n s e q u e n c e of the short durat ion of her exper iment (5 years ) . F inn (1953) f ound greater total fol iar nitrogen for t rees a s soc i a ted with b lack locust c ove r than control in a lmost every instance. Ha ine s et al. (1978) found higher fol iar N concentrat ions in S y c a m o r e a s soc i a ted with legumes, a s d id J o r g e n s e n (1980) in Liquidambar styraciflua L. ( sweetgum) and Pinus taeda L. (loblolly pine) fo l iage and G a d g i l (1979) in Pinus radiata D. Don (Monterey pine). H a d lodgepole p ine accumu la ted signif icantly more f ixed-nitrogen in the c lover plots, 8 1 SN va l ue s wou ld have b e e n pred icted to be signif icantly les s than those of control. Th i s is due to the fract ionation of N i sotopes dur ing the N cyc le . " N and 1 S N are two naturally occurr ing stable i sotopes found in a constant ratio in the a tmosphere, 1 5 N equal l ing 0.3663 a tom % or 3660 p p m (Renn ie and Renn ie 1983, Hoe f s 1987). Dur ing microbia l denitrif ication the i sotopes a re f ract ionated; 1 S N is d i scr iminated against and " N is r e l ea sed back to the a tmosphere (Renn ie et al. 1976, Hoe f s 1987). Th i s results in soi l N conta in ing a h igher proport ion of 1 5 N than a tmospher i c N. P lants that accumu la te N f rom the soi l thus conta in a h igher proport ion of 1 5 N than those wh i ch obta in N f rom nitrogen-fixing bacter ia. T h e s e latter plants theoretical ly have 81 5N ratios c l o se to 0, w h e r e a s the fo rmer will have va lues c l o se r to that of so i l . After s ome yea r s of c lover litter accumulat ion a n d decompos i t i on ( including be low ground s lough ing and root d ie back), one wou ld expect the soi l N poo l to have lower va lues of S 1 S N, and in turn lodgepo le p ine fol iage to a l so re spond with lower va lues . Th i s is not evident after only one full s e a s o n s ' nutrient cyc l ing (fall 1987 to m id - summer 1988). T h e s e ear ly results are not unexpec ted ; the ma in pu rpose of obta in ing the va lue s w a s to document initial levels of fol iar 81 5N status. In addit ion, B ink ley et al. (1985) found this method prob lemat ica l in trying to t race a lder-f ixed N, a l though they found significant d i f ferences in i sotopic compos i t i on of N poo l s within s ites, they d id not f ind cons i s tent patterns be tween sites. 2.4 Conc lus ions T h e exper imenta l endophyte ' s ability to infect a l s ike c lover and c a u s e effective nitrogen fixation, a s we l l a s the estab l i shment of a l s ike c lover o n the site, w a s a s s e s s e d a s excel lent. C l ove r percent cove r and vo lume inc reased with amount of seed ing , with s ome of the d i f ferences be tween levels of 25 N-treatment being statistically significant. Volume estimates of alsike clover were not more sensitive to production differences between rates of seeding than percent cover alone. Site preparation had no effect on the establishment of the symbiosis, nor were there any interactions between site preparation and N-treatment. There was no effect of the symbiosis on height or survival estimates of lodgepole pine. There were significant post-treatment differences for total and incremental diameter between the control and other levels of N-treatment. The linear regressions over two growing seasons of diameter increment as a function of clover percent cover were significant, but only accounted for 21 and 11% of the variability during the second and third growing seasons, respectively. Needle mass was significantly less where the seedlings were associated with clover compared to control, although the results are not totally independent of site preparation. There were no N-treatment effects for either total foliar N or 815N. 2.5 Summary In the first three years of the study, both alsike clover and lodgepole pine have become established and are initially growing acceptably well together. Alsike clover-Rhizobium symbiosis had no effect on lodgepole pine height increment, survival in the first three years, or foliar nitrogen status after one year. The symbiosis did have a negative effect on diameter and needle mass, but the absolute differences between no clover and increasing amounts of clover on these parameters are too small to make reasonable biological interpretations and predictions at this point. It will be important to continue these measurements in order to determine if these early results remain the same, disappear, or become increasingly significant as the crops mature while sharing and competing for site resources. 3 EFFECTS OF TRIFOLIUM-RHIZOBIUM SYMBIOSIS ON SOIL 26 3.1 Introduction Forest soils are often characterized by a low availability of nitrogen (Gordon and Dawson 1979) which is generally considered the principal nutrient limiting the productivity of forest crops. Natural ecosystem processes such as nitrification, denitrification, and leaching result in a loss of nitrogen from soils. But perhaps most important in intensive forest management, is the possibility of decreasing the levels of nitrogen and other nutrients as a result of biomass removal through forest harvesting (Fortin et al. 1984). Forest managers are recognizing the need to manage the soil as well as the crop in order to maintain or potentially improve site productivity. The conventional practice of maintaining site nutrients has been through the use of fertilizers, but concerns over the cost of fertilization and the consequences of extensive synthetic fertilizer use on soil and ground water quality have arisen (Pritchett 1979). Consideration of forest management over the long term may lead to increased recognition of the need for alternative methods of enhancing fertility. Legumes for forestry have been used in central Europe, Australia, New Zealand, and the USA (Jorgensen 1980). Fortin et al. (1984) summarized the rates of microbial symbiotic N-fixation to range from 85 to 300 kg/ha/yr. Gadgil (1979) estimated lupines to provide 60 kg/ha/yr under Pinus radiata (D.Don) until canopy closure. Whatever the actual fixation rate is on any given site under the prevailing environmental conditions, biological nitrogen fixation has the potential to ameliorate losses of N resulting from forestry practices. While the obvious benefit of using nitrogen-fixing plants is increased fixed nitrogen in the soil, other benefits included contributions to increased organic matter with its related improvements in soil structure, nutrient and moisture holding capacity, and environment for soil biota. In the Sub-Boreal Spruce (SBS) biogeoclimatic zone (Pojar et al. 1984), the use of prescribed fire has been shown to have an effect on soil nutrients, particularly loss of N (Macadam 1987, Taylor 1987, Ballard and Hawkes 1989, Blackwell 1989). Following prescribed fire in the SBS, Macadam (1987) reported average losses of 376 kg/ha from the forest floor and upper 30 cm of mineral soil, Taylor and Feller (1987) reported losses ranging from 470-650 kg/ha from the forest floor and slash, and Blackwell (1989) found 227 to 829 kg/ha lost in the forest floor alone over a range of fire severities. This study assesses the effects of the alsike clover-Rhizobium symbiosis on soil chemistry, and soil N status in particular, following broadcast and windrow burning in a SBS lodgepole pine ecosystem. 3.2 Methods 3.2.1 S tudy area a n d experimental d e s i g n The study area and experimental design have been described in Sections 1.3 and 1.4. Within the study site, the experiment was established incorporating three site preparation techniques. Each of these, broadcast bum, windrow burn, and scrape (areas between windrows), was split into four 20 m 2 plots (4 x 5 m), replicated in three rows in each of three blocks. These split-plots were randomly assigned one of four seeding rate levels (N-treatment). The N-treatment consisted of 0 (control), 10, 20, and 30 kg/ha of inoculated alsike clover seed. There were 108 split-plots in total (see Figure 1, page 6). 3.2.2 Fie ld s a m p l i n g and preparation Soil sampling took place following site preparation treatments (1985-86) in May 1987 before clover seeding (pre-treatment), and in early June 1988 (one year post-treatment). In all cases, the sample locations were randomly chosen from areas considered to represent acceptable planting spots, and avoiding standing water, excessive coarse fragments, and large accumulations of decaying wood. All samples were collected into pre-labelled plastic bags, temporarily stored in cool, shaded locations until returned to the laboratory within one to three days. The samples for chemical analysis were air dried at room temperature until mass became constant, then sieved to pass a 2 mm mesh. The samples taken for determination of mineral soil bulk density and forest floor mass were oven dried at 105°C until a constant mass was reached. Chemical analysis was conducted on one sample per experimental unit (108 units) on each of forest floor and mineral soil. Each sample consisted of a composite sample of four randomly located subsamples. The forest floor subsamples were each taken from an area 20 x 20 cm from the soil surface to the mineral soil interface in the broadcast burned areas. Forest floor was not present in the windrowed or scraped areas, except in distinctive microsites. The composite mineral soil sample was taken from the undisturbed mineral soil interface to a depth of 15 cm using an auger. In the windrowed areas, the undisturbed mineral soil layer was commonly several centimetres below a layer of ash. Ash was not sampled. For mineral soil bulk density estimates, one sample per row (nine per block, 27 in total) was collected. The mineral soil bulk density was determined by excavating and retaining a volume of no less than 1 L of soil including coarse fragments. The hole was then lined with a thin plastic bag and filled with silica gel to occupy the volume of material removed. The volume of the hole was then determined by filling a graduated cylinder with the silica gel. The coarse fragment content was determined by sieving out material finer than 2 mm, then weighing the remainder. An adjusted bulk density was then calculated using the method of Macadam (1987): Mtot " M cf = Df V t ot where M t o t is the oven-dry mass of the entire sample, M c f is the mass of the material larger than 2 mm in diameter, and V t o t is the total volume occupied by the sample (as determined in the field). Df, the adjusted bulk density expressed in g/cm3, is the proportion of the undisturbed soil volume occupied by materials finer than 2 mm in diameter. Five forest floor mass samples per block (15 in total) were obtained from four composite subsamples per sample, each subsample was taken from a randomly chosen 20 x 20 cm location in broadcast burned areas. The four subsamples were carefully excavated using a template and then composited into one sample. The composite samples were subsequently oven dried at 105°C, and the coarse fragment-free forest floor mass was then calculated and expressed in kg/m2. Mineral soil exposure was estimated by determining the percent cover of exposed mineral soil on each experimental unit in broadcast burned areas. This estimate was then applied as a correction factor to nutrient quantities to account for areas without forest floor. Finally, the mean estimate of mineral soil bulk density and forest floor mass for each block was applied to chemistry results in order to convert soil nutrient concentration to content (kg/ha). 29 3.2.3 Laboratory ana lys i s Methods of soil chemical analysis are those described in Page et al. (1982) except where indicated. Soil pH was determined in soil/0.01 M CaCI 2 suspensions using a pH meter. Total carbon (C) and nitrogen (N) were measured using Leco induction elemental analyzers. Available phosphorus (P) was extracted with acid-fluoride (Bray 1) and analyzed colorimetrically on a UV/visible spectrophotometer. Cation exchange capacity (CEC) and exchangeable calcium (Ca), magnesium (Mg), and potassium (K) were determined by the NH 4OAc/pH7 method. Released NH 4 for C E C estimation was determined by colorimetric analysis using a Technicon Autoanalyzer while the cations were determined by ICAP spectrophotometry. Mineralizable N was estimated through a two week anaerobic incubation at 30°C, followed by a 1N KCI extraction and colorimetric analysis for ammonium N. Analyses were done at the British Columbia Ministry of Forests Research Laboratory. 3.3 Resu l t s and D i s c u s s i o n Soil nutrient contents, C E C , and pH are presented in Table 4, while mineral soil bulk density and forest floor mass are presented in Table 5. Summaries of the pertinent statistical analyses are presented in Appendices 2 and 3. Table 4. S o i l nutrient content (kg/ha), ca t ion exchange capac i ty (CEC) , and p H pre-treatment, one year post-treatment, wi th pre- to o n e year post-treatment dif ference. S e e d rate (kg/ha): L i nea r contras t : S i t e p repa ra t i on 1 : 0 10 20 30 (p<0.05) B B S W B Nitrogen to ta l : forest f loor 2 pre 2 46 240 2 2 7 234 no 237 _ post 2 4 7 242 255 242 no 246 -d i f fe rence 2 2 28 8 no 9 _ mine r a l 3 pre 8 7 0 902 9 0 6 856 no 607b 8 1 4 b 1 2 3 0 a * post 9 4 0 832 850 800 no 5 0 6 c 8 3 2 b 1 2 3 0 a * d i f fe rence 70 -70 -55 -55 no -100 18 0 mineral izable :" forest f loor post 38 4 3 4 3 51 y e s 31 _ minera l post 2 6 26 28 30 y e s * 13b 2 5 a b 4 6 a C a r b o n :NHrogen forest f loor pre 60 62 63 59 no 61 _ post 4 4 47 43 47 no 45 d i f fe rence -16 -15 -21 -12 no 16 _ _ minera l pre 3 3 35 35 33 no 28b 3 3 b 4 1 a post 3 5 3 4 37 42 no 33 3 3 4 6 d i f fe rence 2 -1 3 8 no 5 0 5 P h o s p h o r u s forest f loor pre 11 11 11 12 no 11 _ post 13 13 11 11 y e s 12 - -d i f ference 2 2 -1 -1 y e s minera l pre 5 7 61 59 52 no 33b 5 0 b 8 8 a post 80 59 63 71 no 39b 5 0 b 115a d i f ference 2 3 -2 4 18 no 6 0 26 P o t a s s i u m forest f loor pre 2 6 28 28 30 y e s 28 _ _ post 20 21 21 22 no 21 _ _ d i f ference -5 -6 -7 -7 no -6 _ _ minera l pre 2 44 256 245 231 no 185b 2 0 1 b 3 4 6 a post 2 54 243 236 245 no 168b 198b 3 6 7 a * d i f ference -10 17 -10 25 no -16 -2 21 Table 4 continued. Calcium forest f loor pre 174 184 179 184 no 180 _ post 194 203 191 205 no 198 _ _ d i f fe rence 19 19 11 21 no 18 _ minera l p re 1028 991 1097 993 no 7 2 1 b 931 a b 1 4 3 0 a post 1365 1393 1345 1451 no 7 2 0 b 1 0 1 9 a b 2 4 2 6 a d i f fe rence 3 3 7 402 247 458 no -1 89 9 96 M a g n e s i u m forest f loor pre 24 25 24 27 no 2 5 - _ post 20 23 22 25 no 2 2 _ _ d i f fe rence -4 -1 -1 -1 no -2 _ _ minera l pre 224 236 238 216 no 159b 1 8 7 a b 3 3 9 a post 249 251 240 239 no 144b 1 9 1 a b 4 0 0 a d i f fe rence 26 16 2 23 no -15 3 61 CEC (meq/100g) forest f loor pre 88 90 90 88 no 8 9 - _ post 81 8 5 82 8 3 no 8 3 d i f fe rence -7 -5 -9 -5 no -4 _ _ minera l pre 14 14 15 14 no 14 13 16 post 14 14 14 14 no 12 12 18 d i f fe rence 0 0 0 0 no -1 -1 2 pH forest f loor pre 4.7 4.8 4.8 4.7 no 4.8 post 5.0 5.1 4.9 5.0 no 5.0 _ d i f ference 0.3 0.3 0.2 0.2 no 0.3 _ minera l pre 4.7 4.5 4.6 4.5 no 4.4b 4.3b 5.0a post 4.8 4.8 4.7 4.9 no 4.5b 4.6b 5.3a d i f fe rence 0.1 0.2 0.1 0.4 no 0.1 0.3 0.3 B B = b roadca s t bu rn ; S = s c rape ; W B = windrow burn. M e a n s in the s a m e l ine f o l l owed b y the s a m e letter a r e not s ignif icantly different (p < 0.05) u s i ng Tukey ' s S t uden t i z ed R a n g e (HSD) test. 2 forest f loor s a m p l e d on ly in b roadcas t burn treatment 3 m inera l = 0-15 c m mine ra l so i l 4 s a m p l e d only post-treatment * ind icates s ignif icant b lock interaction 32 Table 5. Mineral s o i l bulk densit ies a n d forest f loor m a s s est imates. Mineral s o i l : Bulk density (a/cm3) broadcast burn windrow burn scrape 0.7 (0.06) 0.9 (0.08) 0.9 (0.21) Forest f loor : Mass (kg/m2) Mineral soil exposure (%) block 1 block 2 block 3 3.26 (0.03) 3.23 (0.06) 4.13 (0.04) 18 15 25 1 standard error in parentheses Linear effects (contrast) (Hicks 1982, Ott 1984) are presented in Table 4 to show the significance of N-treatment effect. Remaining contrasts (quadratic, cubic, and control vs others) and Tukey's Studentized Range (HSD) test results for N-treatment are summarized in Appendices 2 and 3. Since post-treatment results only reflect effects due to the first growing season (1987), significant soil chemistry changes due to the N-treatments were not expected to be observed. However, after only one year, significant linear effects were observed for mineralizable N in both soil layers, and for available P in the forest floor. Where main effects for N-treatment were observed, no interactions with site preparation where found. Soil chemistry differences due to site preparation treatments largely agree with those of Blackwell (1989), and were of less specific interest to this study, except where an interaction might have occurred with N-treatment. Tukey's Studentized Range (HSD) test (p < 0.05) was chosen as the appropriate multiple comparison test (Zar 1984), and the results are presented in Table 3 to show site preparation effects. All comparisons were for the mineral soil layer only, since the forest floor layer was not present in the windrow burn and scrape areas. Pretreatment (before clover seeding but after site preparation) significant differences were found for total N, C:N ratio, K, Ca, Mg, and pH. Post-treatment (one year following seeding) significant differences continued to be observed for all the preceding variables, except for C:N ratio. In addition, post-treatment mineralizable N (sampled only post-treatment) was also found to have been significantly affected by site preparation treatment. 3.3.1 Nitrogen and Carbon For total N, no significant linear effect was observed among N-treatments, either in the forest floor or mineral soil layer. However, for mineralizable N there were significant linear effects observed for both the forest floor (p < 0.04) and mineral (p < 0.02) soil layer one year post-treatment as illustrated in Figure 10. Figure 10. Mineralizable N in the forest floor and mineral soil layer one year post-treatment. Mineralizable N is an index of N availability determined using an anaerobic incubation. Its strengths and weaknesses are summarized in Page et al. (1982). The result of the anaerobic incubation is a measurement of NH/-N, and has been operationally applied as a good predictor of soil N availability for agricultural and forestry crops. Most N in the soil is in organic form unavailable to higher plants. The major biological forms include proteins, microbial cell wall constituents, and nucleic acids (Paul and Clark 1989). While the amounts of inorganic N (1-5% is commonly cited) found in soil are small, it is of great importance as the primary source of N for most plants. The conversion of organic N to inorganic N is termed mineralization. Mineralization, or "release" of organic N in nature is the end product of enzymatic digestion by soil organisms which also results in inorganic N, usually N H / (Bartholomew 1977). Mineralization proceeds best in well-drained, aerated soils through the activity of many heterogenous organisms (Brady 1974), whose C source is derived from the hydrolysis of organic nitrogen compounds. The NH 4* is then immobilized by other soil organisms, fixed by clay minerals or organic matter, utilized by higher plants, or oxidized to N 0 3 by other soil micro-organisms (Brady 1974). Some of the NH 4* is part of the exchange complex, and small amounts may be volatilized as NH 3 (Paul and Clark 1989). The oxidation of NH 4 + to N0 3 , which is also readily available to higher plants, is referred to as nitrification. Autotrophic and heterotrophic organisms (all but one heterotroph are obligate aerobes) oxidize the mineralized ammonium. The predominant nitrifying organisms found in soil are in the bacterial genera Nitrosomonas and Nitrobacter (Alexander 1977). These bacteria are chemoautotrophic aerobes, their source of C is met by the assimilation of C 0 2 and carbonates (Paul and Clark 1989) and their activity in the soil is limited mainly by concentrations of NH 4* (energy substrate) and 0 2 (Schmidt and Belser 1982). The soil properties most likely influencing mineralization and nitrification of N are nutrients, aeration, temperature, moisture content, pH, C:N ratio, and plant-produced allelochemicals (Harmsen and Kolenbrander 1977, Robertson 1982). Although these properties may be important regulators, they may not be good predictors across a wide range of sites (Robertson 1982). This may be due to measurement techniques, interactions, seasonal variation, and other prevailing site conditions. The significant increase of mineralizable N (potentially available N) in this study could not be attributed to additions of newly cycled fixed-nitrogen (by the legume-Rhizobium symbiosis), since there were no significant overall increases observed for total N. This latter result in itself was not surprising since the initial production (see Table 2, page 14) and length of decay time for above-ground clover detritus were considered to be insufficient following the first growing season to have any detectable effect on soil N-status. The observed differences in mineralizable N must, therefore, be attributed to N-treatment effects on the relative distribution of N forms within the soil N-pool, excluding the contribution of any above ground clover detritus in the first year. The mineralizable N results indicated that the total N-pool in soils under clover was potentially more capable of being transformed from organic to inorganic N, suggesting that the clover root systems may at least in part, be responsible for the effect. Clover production below ground was assumed to increase with rate of seeding as was observed for the above ground production (see section 2.3.1). Therefore, some discussion of N mineralization and nitrification related to the rhizosphere is warranted in light of the significant effects observed. What impact might the clover root systems have on the soil environment that could lead to enhanced N mineralization? The rhizosphere has been defined as that part of the soil in which roots generally induce a proliferation of microorganisms (Balandreau and Knowles 1978). The rhizosphere can affect soil biota, chemistry, and physical properties through the penetration, and excretions and sloughing off of root tissues, all of which can in turn affect microorganism populations and mineralization rates. Microorganisms proliferate in the rhizosphere as a result of nutrients liberated by the plant (Giddens and Todd 1984). Rovira (1961) reported proliferation of microorganisms in the rhizosphere of most plants, and legumes in general support a higher rhizosphere population than do other plants (Rovira and Stern 1961, Curl and Truelove 1986). Alexander (1977) cited Lyon and Bizzell (1913) who reported soils taken under alfalfa nitrified more readily than those under timothy, and Stiven (1952) and Theron (1951) who reported that nitrification may be slow under grass in general, due to toxic substances in rhizosphere exudates. Davidson (1978) also reported that mineralization of N declines under perennial grassland, attributing the observation to secretion of antibiotic substances secreted by plants. 3 6 In the present study, plots containing clover contributed to an increased rhizosphere volume and content, which likely increased the activity and population of microorganisms in the soil. Myrold (1987) concluded that mineralizable N determined by anaerobic incubation is largely a measure of microbial biomass N. The mineralizable N results of this study could therefore be attributed to an increased microbial population stimulated by the presence of the clover roots. Other factors cited that are known to induce nitrification are increases in pH and P (Russell 1973), and decreases in C:N ratios might indicate enhanced changes (Bartholomew 1977, Harmsen and Kolenbrander 1977). However in this study there was no post-treatment evidence of clover significantly increasing pH and P, or decreasing C:N ratios. Nevertheless, there were generally post-treatment P increases in the mineral soil layer, C:N ratio decreases in the forest floor, and slight increases in pH in both soil layers; all of which may have contributed to increased N mineralization and nitrification independent of the N-treatment. Pre-treatment mineralizable N was not determined in this study and thus, it is not possible to confirm the contribution of these other factors to increased mineralization. Huss-Danell and Lundmark (1988) reported increases in the proportion of total N that could be mineralized after six years by annually adding fragmented leaves of alder [Alnus incana (L.) Moench] to field plots. They observed increases up to 2.5 and 8 % in the mineralizable N:total N ratio for N in the forest floor and mineral soil layer, respectively. In the present study, maximum increases in the mineralizable N:total N ratio were 7.7 and 1.0 % in the forest floor and mineral soil respectively, after one year of clover production in seeded plots compared to control plots with no clover (Table 6). Furthermore, these ratios in relative terms show that there were maximum increases of 57 and 3 6 % in mineralization rates comparing maximum ratio differences to that of control values for the forest floor and mineral soil respectively (Table 6). The forest floor might be expected to have higher initial mineralization rates due to the overall accelerated decay processes normally observed following disturbance by slashburning. In addition, for broadcast burn areas, the distribution of clover roots in the first growing season would have been established first in the forest floor, and whatever rhizosphere influences have on mineralization rates, as previously discussed, would therefore be first observed in the forest floor soil layer. 37 Table 6. Mineralizable N a s a percent of total N for the different levels of N-treatment o n e year post-treatment. Seed rate (kg/ha) 0 10 20 30 Relative maximum increase 1%) Forest floor 13.4 17.8 16.9 21.1 57 Mineral layer 2.8 3.1 3.3 3.8 36 The windrow burn mineral soil had significantly more total N than either scrape or broadcast burn mineral soils both pre- and post-treatment. This site preparation effect is attributable to additional concentrated slash loads, forest floor materials, and resulting ash in the windrowed areas. There was also significantly greater total N in the post-treatment mineral soil layer of the scrape treatment compared to that of broadcast burn. This can be attributed to the loss of total N in the broadcast burn mineral soil layer (-100 kg/ha) compared to essentially no N losses in either scrape or windrow treatments, although that loss is not easily explained and may be attributed to experimental error. When the entire soil N pool was compared between the broadcast burn (forest floor and mineral soil) to that of the scrape mineral soil (forest floor lacking), the comparison was no longer significantly different. Therefore, the two treatments produced similar soil N pools in regard to potential plant nutrition. Mineralizable N was significantly different between the windrow burn (greater values) and the broadcast burn areas (lower values), while in the scrape areas it was not found to be significantly different from the others. This effect is again attributable to the greater concentrations of slash, forest floor materials, and the resulting ash in the windrow areas. The mineralizable N:total N ratio was found to range from 3 to 4 % across site preparation treatments. Carbon:nitrogen (C:N) ratios were not significantly affected by N-treatment in either the forest floor or mineral soil layers. All levels of N-treatment showed post-treatment C:N ratio decreases, particularly in the forest floor layer where there was an overall 2 6 % decrease attributable to accelerated decomposition following burning. In the pre-treatment (prior to seeding) mineral soil layer, the windrow burn C:N ratio was significantly higher than that in the scrape or broadcast burn areas due to the increased amount of C derived from the concentration of organic materials by this type of site preparation. A similar difference was not detected in the post-treatment measurements, although the data reflected a similar trend, increased sampling variability likely contributed to the absence of any statistically significant differences. 3.3.2 Phosphorus Pre-treatment available soil P was similar between levels of N-treatment. A significant linear effect was observed one year post-treatment showing that forest floor P decreased as clover seeding rate increased. The forest floor P differences between N-treatments were small (ranging from 0 to 2 kg/ha), however Tukey's Studentized Range (HSD) test separated the means of control and 10 kg/ha seed rate from those of 20 and 30 kg/ha seed rates (Figure 11). The differences between pre- and post-treatment forest floor P values (Figure 11) also showed a significant linear effect, but Tukey's Studentized Range (HSD) test was unable to separate these treatment means. Similar effects were not observed in the mineral soil layer, where no significant differences were observed pre- or post-treatment. 1 4 1 2 1 0 8 F o r e s t f l o o r P ( k g / h a ) 39 2 0 - 2 1 y r p o s t - t r e a t m e n t D i f f e r e n c e p r e - p o s t 0 10 2 0 3 0 0 10 2 0 3 0 N-treatment (kg/ha clover seeding) F igure 11. C h a n g e s in forest f loor P o n e y ea r post-treatment. So i l P is f ound in inorganic and organ ic forms, most of wh i ch are poor ly so lub le and therefore unava i l ab le to plants. Avai labi l i ty of soi l P is med ia ted by microf lora, and the se m ic roo rgan i sms are concen t ra ted a round the roots, whe re it ha s b e e n es t imated that 9 0 % of rh i zo sphere microf lora are c a p a b l e of inc reas ing P avai labi l ity (Barber 1978). At the s a m e t ime, inorgan ic P may be immob i l i z ed by mic roorgan i sms. H a y m a n (1975), c i ted in G i d d e n s a n d T o d d (1984), states that m i c roo r gan i sms may remove as m u c h P f r om the so i l a s the c rop . L e g u m e s require ava i lab le so i l P a s do all p lants for growth, a n d wil l improve their P uptake by m e a n s of mycor rh iza l a s soc ia t i ons (Munns and M o s s e 1980, B o w e n a n d Sm i th 1981) a n d phosphate -d i s so l v i ng bac te r i a (Pau l and S u n d a r a R a o 1971). The i r pho spha te requi rement may be larger than ave rage b e c a u s e P is n e e d e d not only for plant growth but a l so for nodulat ion and N 2 f ixat ion. In addit ion, de f i c ienc ie s wi l l a l so inhibit N 2 - f ix ing s ymb io s i s ( J o r gen sen 1978, G ranha l l 1981). P h o s p h o r u s de f i c iency wil l o f ten affect n itrogen f ixation before it affects plant growth, attributed to its effect o n the net synthes i s of A T P (Granha l l 1981). N i t rogenase, the e n z y m e respons ib le for the reduct ion of N 2 to N H 3 , d e p e n d s o n a supply of b io log ica l energy v i a the A T P generat ing s y s t em (Quispel 1981). Increases in available soil P increase nodule number and weight, and nodule development and N-fixation will begin earlier with increasing P supply (Rovira 1978). Higher concentrations of P have been found in nodules of P-deficient host plants than host plants with adequate P (O'hara et al. 1988). The forest floor P depletion found in this study is attributable both to the general increase in vegetation (clover) and rhizosphere microorganisms requiring overall greater amounts of available P, and also to the requirement of the symbiosis in active N-fixation. However, it must be remembered that only the broadcast burn treatment had a forest floor layer. This effect was not observed in the mineral soil, which is the surface soil in the windrow and scrape treatments. Since the mineral soil layer in general contained five to six fold the amount of available P compared to the forest floor, small depletions might not be detectable or significant due to the greater relative amounts available. Mineral soil P was significantly different pre-treatment among all site preparation treatments; broadcast burn had the lowest values and windrow burn had the highest values. Post-treatment mineral soil P was no longer significantly different between the broadcast burn and scrape treatments. However, windrow areas continued to have significantly greater values than either of the other treatments. Changes in P availability are closely related to changes in pH (Tisdale and Nelson 1966). A significantly greater pH was observed in the windrowed areas (see section 3.3.3 following) and would partially account the greater availability of P in the windrow areas. Such greater availability can also be partially explained by P inputs from the large amounts of ash and other concentrated organic residues previously mentioned. 3.3.3 Cations, pH, and C E C N-treatment had no significant effect on any cations (K, Ca, and Mg), or on C E C . In addition, N-treatment had no significant effect on pH, although there were slight (0.2-0.3 units) increases post treatment across N-treatment. Windrow areas had significantly higher values for cations and pH, pre- and post-treatment. These results are again attributable to the concentration of large amounts of ash and other concentrated organic residues. The C E C is slightly (but not significantly) higher in the windrow areas, likely due to higher C concentration resulting in a greater capacity for exchange sites. 41 3.4 Conclusions Mineralizable N and available P were the two soil chemistry parameters which showed a significant response related to N-treatment. Mineralizable N increased in both the forest floor and mineral soil layers as rate of seeding increased, whereas forest floor P decreased as rate of seeding increased. While increases in mineralizable N could not be attributed to newly fixed-nitrogen, they probably indicate an increased amount of microbial biomass N. The clover rhizosphere may have provided an environment which stimulated increased activity and populations of soil micro-organisms. The requirement for P from increasing amounts of total vegetation and microorganisms, and that specifically associated with active nitrogen fixation, are likely the reasons for the small, but significant depletion in forest floor P. However, the same result was not observed for mineral soil P and is likely due to the large relative differences between forest floor and mineral soil P. Where greater amounts of nutrients or changes in pH and C E C were observed for site preparation treatments, they were associated with windrow bum areas and the results were attributable to the concentration into windrows of slash, organic debris, and ash. These results were observed in the spring (1988) following only one growing season, where the greatest ground cover of clover in the N-treatment was estimated to be five percent. In the most recent year of measurement (1989), the overall mean estimate in seeded plots was 7 6 % cover. The clover production, above and below ground, had dramatically increased and may have affected some very interesting changes in the soil. Resampling of the soils in 1990 is highly recommended in order to quantify the effects of the symbiosis after three growing seasons. 42 4 E F F E C T S O F TRIFOLIUM-RHIZOBIUM SYMBIOSIS ON NATIVE VEGETAT ION 4.1 Introduction The legume-Rhizobium symbiosis is largely utilized as a management practice to enhance soil nitrogen status, and thereby increase the productivity of the site with the purpose of increasing crop yield. Another consequence of introducing this management practice into forestry is the possible source of vegetation competition between the crop species (see Section 2) (Gordon 1983) and other non-crop species. This section of the thesis reports the early results of introducing inoculated Trifolium hybridum (alsike clover) at different seeding rates on some selected native species found on the experimental site. There was no published literature found reporting the effects of legume symbiosis on associated native vegetation in forestry in British Columbia. In British Columbia, some Forest Districts are using grass/legume mixtures to control native "brush" species and for range forage improvement, but quantitative assessments have yet to be reported. In general, the operational results in the interior have met with satisfaction from practising foresters and agrologists [D. Russell and G. Ellen (Range Officers), and O. Steen (Research Ecologist), pers. comm., BCFS, 1990]. Research trials established in coastal British Columbia have also shown promising results, particulary with grasses, but it was recognized that site properties and type of site preparation will greatly influence the results [R. Green (Research Pedologist), pers. comm., BCFS, 1990]. Kumi (1986) found Lotus corniculatus was able to provide some measure of weed control over several coastal sites, while other legumes were less effective and persistent. Klingler (1986) found significant reductions of Alnus rubra and Rubus parviflorus after several burned clearcuts were seeded with grass/legume mixtures in western Oregon, USA. Vegetation in the study area was not as abundant or diverse as it's productive capability suggested, given the site's ecological attributes (Pojar et al. 1984). This situation is mainly a consequence of the wildfire and stand history of the overdense, repressed lodgepole pine occupying the site (J. Parminter (Fire Ecologist), pers. comm., BCFS, 1986) and its resulting 'vegetatively poor" understory composition. The site is capable of moderate productivity for forestry and related 43 re sources (Pojar et al. 1984). T h e excel lent es tab l i shment of a ls ike c lover on the site is support ive ev i dence of its product ive capabi l i ty and is commensu ra te with that expec ted for a zona l e co s y s t em. At present, the vegetat ion commun i ty and soi l propert ies do not suggest the site to be a managemen t conce rn for un succes s fu l coni fer regenerat ion accord ing to the character i s t ics out l ined by Pojar et al. (1984). 4.2 Methods 4.2.1 S tudy area a n d experimental d e s i g n T h e study a r e a and exper imenta l des i gn have b e e n de sc r i bed in Sect ions 1.3 a n d 1.4. With in the study site, the exper iment w a s e s tab l i shed incorporat ing three site preparat ion techn iques . E a c h of these, broadcast b u m , windrow burn, and s c rape (areas between windrows), w a s split into four 20 m 2 plots ( 4 x 5 m), rep l icated in three rows in e a c h of three b locks. T h e s e split-plots we re randomly a s s i g ned one of four s eed i ng rate treatments (N-treatment). T h e levels of N-treatments con s i s ted of 0 (control), 10, 20; and 30 kg/ha of inoculated a ls ike c lover s eed , lightly raked into the s e e d b e d . The re were 108 split-plots in total ( see F igure 1, page 6). S i te preparat ion t reatments were comp le ted in fall 1985 for the s c r ape treatment, and in May - June 1986 for the burning treatments. T h e N-treatment w a s e s tab l i shed in J u n e 1987. 4.2.2 F ie ld s a m p l i n g , measurement, and co l lec t ion Vegetat ion samp l i ng w a s car r ied out in mid-August of 1987 (first growing season) , and 1988 ( second g rowing season) . Pe rcent cove r va l ue s for a l l s pec i e s we re es t imated to the nearest 1 % with the a id of pe rcentage charts (Walms ley et al. 1980), p lacement of wire f lags at 1 x 1 m spac ing in the 20 m 2 split plots, and sma l le r templates to ass i s t in accurate and cons i stent v i sua l est imates. S p e c i e s that were recorded as ' t race ' (present but not enough to a s s i gn 1%) in the f ield, we re g i ven a va lue of 0 . 1 % for da ta ana lys i s . A l l results are p resented to one dec ima l p l ace even though this may not confo rm to s tandard u se for signif icant f igures, in order to indicate c lear ly that a s pec i e s w a s present at cove r va lues be low 1% . V o u c h e r s pec imen s of all va scu la r plants were co l lec ted, p re s sed , mounted, and identification verif ied by a plant taxonomist (J . Pojar). Photographs 44 of e a c h split-plot we re taken f rom permanent locat ions in e a c h sampl ing year to ass ist in histor ical record keep ing . Entry into the plots w a s restricted to keep d i s turbance to a min imum. 4.3 Results and Discuss ion A list of plant s pec i e s ob se r ved o n the study site is found in Append ix 5. N ine native s pec i e s were se lec ted for ana l y se s to determine effects, if any, of exper imenta l treatments. S p e c i e s se lec ted were those that were ob se r ved to have a cove r of approx imate ly one percent or more by the end of the s e c o n d growing s ea son . S p e c i e s with le s s percent cove r were con s i de red not to be appropr iate for da ta ana ly ses . The se lec ted spec ie s inc luded: Arnica cordifolia, Calamagrostis canadensis, Cornus canadensis, Epilobium angustifolium, Petasites palmatus, Linnaea borealis, Rosa acicularis, Salix spp., and Spiraea betulifolia. Percent cove r of a l s ike c lover and native s pec i e s are p resented in Tab le 7. Resu l t s of l inear contrasts (Hicks 1982, Ott 1984) are inc luded in Tab l e 6 to s how the signif icant effects of N-treatment. The remain ing contrasts (quadratic, cub ic , and 'control v s others ' ) a nd Tukey ' s S tudent i zed R a n g e (HSD) test results are s ummar i z ed in Append i x 4. S i n ce post-treatment results only ref lected effects due to the first (1987) and s e c o n d (1988) g rowing s e a s o n s in wh i ch the c lover w a s becom ing es tab l i shed, few signif icant vegetat ion impacts due to the N-treatments were ant ic ipated. No signif icant l inear effects were ob se r ved at the end of the first g rowing s e a s o n and only one spec ie s , R. acicularis, s h o w e d a signif icant l inear effect in the s e c o n d growing s ea son . However , dec rea s i n g va lues of percent cove r for Calamagrostis canadensis a nd S. betulifolia s h o w e d a signif icant effect be tween control (no clover) and all other levels (clover s e e d e d plots) in the N-treatment ( 'control v s others ' ) . Vegetat ion d i f ferences due to site preparat ion t reatments we re of les s spec i f ic interest to this study, except where an interaction might have occu r red with N-treatment. Cornus canadensis a nd L. borealis we re signif icantly ef fected by site preparat ion treatment, but a l so s h o w e d significant block x site preparat ion interactions. The re were no interactions ob se r ved for these s pec i e s with N-treatment. In the s e e d e d plots, a ls ike c lover w a s the dominant spec ie s by the end of both growing s ea son s , and particularly so by the end of the s e c o n d growing s e a s o n with an overal l m e a n cove r of 3 8 % . D i s cu s s i on of a ls ike c lover establ i shment, and its effects on lodgepole pine, is found in sect ion 2.3, page 12. A l l native spec ie s had low percent cove r va lues, genera l ly less than one percent in 1987, and rang ing f rom approx imately o n e to s ix percent in 1988. At the end of the s e c o n d growing s e a s on , native s pec i e s genera l ly i nc reased percent cove r in the order: for herbs -P. palmatus < A. cordifolia < Cornus canadensis < Calamagrostis canadensis < L. borealis < E. angustifolum; and for shrubs - Salix spp. < R. acicularis < S. betulifolia. D i s cu s s i on of s u c h low cove r va lues, no matter how statistically significant, is highly speculat ive and shou ld be interpreted and app l ied with caut ion. Table 7. Percent c o v e r of Trifolium hybridum a n d selected native s p e c i e s at the e n d of the f irst (1987) a n d s e c o n d (1988) g r o w i n g s e a s o n s . S e e d rate (kg/ha) L i nea r contrast : S i t e p reparat ion 1 0 10 20 30 (p<0.05) B B S W B Trifolium hybridum 1987 0.0 1.7 3.2 4.9 y e s 1.0 2.9 3.4 1988 0.0 33.7 37.7 41.7 yes_ 24.9 31.5 28.5 H e r b s : Arnica cordifolla 1987 0.2 0.2 0.1 0.4 no 0.4 0.2 0.1 1988 0.8 0.9 0.8 0.9 no 0.9 0.9 0.8 d i f fe rence 0.7 0.7 0.7 0.5 no 0.6 0.7 0.7 Calamagrostls canadensis 1987 0.1 0.1 0.2 0.2 no 0.2 0.1 0.2 1988 2.0 1.1 1.9 1.1 no 1.8 1.4 1.4 d i f ference 1.8 1.0 1.7 1.0 no 1.6 1-3 1.2 Cornus canadensis 1987 0.4 0.1 0.2 0.3 no 0.6a 0.1b 0.1b * 1988 1.2 1.0 1.2 1.3 no 1.8a 1.0b 0.8b * d i f fe rence 0.8 0.8 1.0 1.0 no 1.2 0.9 0.7 Epilobium angustlfollum 1987 1.6 1.6 2.1 1.7 no 1.2 2.1 1.9 1988 5.2 5.2 6.4 4.8 no 4.6 5.5 6.1 d i f fe rence 3.6 3.6 4.2 3.1 no 3.4 3.3 4.2 Petasltes palmatus 1987 0.2 0.1 0.1 0.1 no 0.0 0.1 0.2 1988 0.9 0.6 0.7 0.7 no 0.5 0.8 1.0 d i f fe rence 0.8 0.5 0.6 0.6 no 0.5 0.6 0.8 Llnnaea borealls 1987 0.7 0.8 0.7 0.6 no 1.4a 0.5b 0.2b * 1988 2.0 2.4 2.1 2.3 no 3.3 1.8 1.5 d i f fe rence 1.3 1.7 1.4 1.8 no 1.9 1.3 1.4 CO Table 7 continued. S e e d rate (kg/ha): L i nea r contras t : S i t e p reparat ion : Shrubs: Rosa acicularis Sallx spp. Spiraea betullfolla 0 10 20 30 (p<0.05) B B S W B 1987 0.8 0.9 0.8 0.5 no 0.4 0.9 1.0 1988 2.1 1.9 1.6 1.2 y e s 0.9 1.9 2.3 d i f ference 1.2 1.0 0.8 0.7 no 0.5 1.0 1.3 1987 0.2 0.2 0.4 0.5 no 0.1 0.5 0.3 1988 0.5 0.6 0.7 1.0 no 0.5 1.0 0.7 d i f ference 0.3 0.5 0.3 0.6 no 0.4 0.5 0.4 1987 1.8 2.8 1.5 1.9 no 1.6 2.7 1.8 1988 3.4 3.5 2.5 2.8 no 3.3 3.2 2.5 di f ference 1.6 0.7 0.9 0.9 no 1.7 0.6 0.8 1 B B = b roadcas t burn; S = s c r ape ; W B = windrow burn. M e a n s in the s a m e line f o l l owed by the s a m e letter are not s ign i f icant ly different (p<0.05) us ing Tukey ' s S tudent i zed R a n g e (HSD) test. * ind icates signif icant b lock x site preparat ion interaction 4.3.1 H e r b s N-treatment s h o w e d no s ignif icant l inear effect for any herb a s s e s s e d dur ing either g rowing s e a s o n . T h e contrast ' contro l v s o ther s ' s h o w e d weak l y s ignif icant ef fects of N-treatment for Calamagrostis canadensis by the end of the s e c o n d g rowing s e a s o n (F igure 12). A l though percent c o v e r va l ue s for Calamagrostis canadensis we re sma l l , the relative effect (approx imately 3 0 % dec rea se ) h a s con s i de rab le operat iona l interest, a s the s pec i e s is c o n s i d e r e d a very aggres s i ve compet i to r for con i fers in northern interior Brit ish C o l u m b i a . W h e r e Calamagrostis canadensis g rows abundant l y and v igorous ly, it may c o m p e t e with con i fer roots, c a u s e s nowpre s s , or prevent natural regenerat ion (Haeu s s l e r a n d C o a t e s 1986). In addit ion, Calamagrostis canadensis is not con s i de red a des i rab le forage s p e c i e s in c o m p a r i s o n to a ls ike c lover (D. R u s s e l l ( R a n g e Off icer), pers. comm., B C F S , 1990). Calamagrostis canadensis c o ve r may b e benef ic ia l in r educ ing sur face eros ion, frost heav ing , a n d in add ing o rgan i c matter to soi l . A l s i ke c l ove r may b e the mo re des i rab le s pec i e s for forest m a n a g e m e n t d u e to its range fo rage va lue and ability to fix n itrogen, in addit ion to those benef i t s c i ted for Calamagrostis canadensis. 2.5 2.0 % cover Calamagrostis H Control -VZ1 Clover p < 0.07 p < 0.04 F igure 12. Pe rcen t c o v e r of Calamagrostis canadensis at the end of the first two g rowing s ea son s . Epilobium angustifolum may be a s ignif icant compet i tor with coni fers if abundant in the early s tages of reforestation, a lthough most d a m a g e to coni fers is primarily through snowpre s s (Haeus s le r and C o a t e s 1986). It to lerates a w ide range of so i l a nd site condit ions and is an aggres s i ve p ioneer spec ie s . It g rows erect and relatively tall (1-3 m), propagat ing shoots f rom extens ive lateral, rh i zome-like roots, or by seeding- in. A l though E. angustifolium had the greatest percent cove r for native herbs a s s e s s e d in the study a rea in both years , approx imate ly 2 % in 1987 and 5 % in 1988, it w a s not s ignif icantly af fected by N-treatment, probably due to its early p re sence a n d establ i shment pre-treatment. Th i s s pec i e s w a s in fact ob se r ved to beg in g rowing aboveg round within two w e e k s after e a c h p resc r ibed burn in the study. In addit ion, its v igorous and erect growth habit early in the s e a s o n may have p rec luded compet i t ion for light, and its extens ive lateral root s y s t em may have c o m p e t e d we l l with a l s ike c l ove r for moisture and nutrients. It has b e e n ob se r ved that grass/legume mixes u s e d in the C l ea rwate r Forest District c ompe te extremely we l l with E. angustifolium (G . E l l en (Range R e s ou r c e Officer), pers. comm., B C F S , 1990), and Haeu s s l e r and C o a t e s (1986) reported that a r ea s s e e d e d to g ra s s c a n have lower cove r and f requency of E. angustifolium than u n s e e d e d a reas . Therefore, a ls ike c l ove r may be ob se r ved to have an effect o n E. angustifolium in the future, particularly in relation to 1989 mea su rement s of a l s ike c lover cove r wh i ch a ve raged 7 6 % (see Sec t i on 2.3.1.2, page 13). Linnaea borealis s h o w e d a signif icant site preparat ion re sponse dur ing the first g rowing s ea son , but a l so s h o w e d a signif icant interaction with b lock (Figure 13). In the b roadcas t b u m treatment in b lock 1, L. borealis had a greater percent cove r va lue than for other b locks, but s imi lar va lues for a l l b locks we re ob se r ved for both s c rape and w indrow burn treatments. The site preparat ion effect for L. borealis w a s no longer apparent in the s e c o n d growing s ea son . Cornus canadensis b ehaved in a s imi lar w a y to L. borealis for both growing s e a s o n s (Figure 14a and b). B lackwe l l (1989) reported moderate severity of b roadcas t b u m s for b locks 1 and 2, and high severity for b lock 3. However , tho se different fire sever it ies do not account for the ob se r ved interactions s ince s imi lar percent cove r v a l ue s w e r e ob se r ved for b locks 2 and 3. In addit ion, B l ackwe l l (1989) reported no signif icant b i omas s d i f ferences for Linnaea or Cornus be tween the two fire sever it ies. There may have b e e n signif icantly h igher initial (pre-treatment) percent cove r va lues for these spec ie s in b lock 1 than in the other b locks in b roadcas t burn areas , a l though this content ion cannot b e conf i rmed s ince pre-50 treatment estimates were not obtained. Both scrape and windrow burn treatments appeared to have had similar lower percent cover values for L. borealis and Cornus canadensis. These results were likely associated with the complete removal of forest floor common to both the scrape and windrow treatments, which may have severely damaged or removed its propagules. Blackwell (1989) found no significant (p < 0.1) effects of site preparation treatments on the biomass of Epilobium, Linnaea, and Cornus spp. Arnica cordifolia and P. palmatus each had less than 1% cover after two growing seasons, and neither were significantly affected by N-treatment nor site preparation. All of the above results suggest that N-treatment had little interpretable effect on percent cover of native herb species in the study area, and effects due to site preparation are, in part, confounded by initial percent cover block differences before site preparation treatments. % cover 1987 Broadcast Scrape Windrow S i t e p r e p a r a t i o n t r e a t m e n t Figure 13. Site preparation x block interaction for Linnaea borealis in the first growing season. % cover 1987 Broadcast Block 1 Block 2 Block 3 Scrape Site preparat ion treatment Windrow % cover 1988 3.5 r Broadcast Block 1 Block 2 Block 3 Scrape Site preparat ion treatment Windrow F igure 14. S i te preparat ion x b lock interactions for Cornus canadensis in the first two growing s ea s on s . 52 4.3.2 Shrubs The only native shrub observed to have a significant linear effect was R. acicularis in the second growing season (Figure 15). Although the percent cover values were small, the overall relative decrease between the control and the highest rate of seeding was found to be 43%. Spiraea betulifolia showed a significant 'control vs others' contrast for the differences between 1987 and 1988 values (Figure 16). Neither of these shrubs are locally considered to compete with conifers. However, these species can be part of a larger shrub-complex plant community on some sites that may have an overall competitive effect [D. Coates (Research Siliviculurist), pers. comm., BCFS , 1990]. The ability of alsike clover to generally compete with shrubs is likely to be considered beneficial for forest regeneration, particularly if the net effect of clover is found to supply newly-fixed nitrogen to the site without providing more competition to tree seedlings than the native plant complex which would have grown up in the absence of clover. No significant N-treatment effect was observed for Salix spp., and no site preparation effects were observed for any of the shrubs assessed. % cover Rosa 2.5 r L inear contrast s ign i f icant at p ( 0.04 10 20 Rate of seeding (kg/ha) Figure 15. Percent cover of Rosa acicularis at the end of the second growing season. 53 D i f f e r e n c e i n % c o v e r Spiraea 1 9 8 7 - 8 8 Control Clover Figure 16. Change in percent cover of Spiraea betulifolia from 1987 to 1988. 4.4 Conc lus ions The establishment of the alsike clover-Rhizobium symbiosis had little apparent effect on native vegetation in this study. However, the initial low cover values for native vegetation may have accounted for the absence of significant effects on this site. Had the native vegetation on the experimental site been more abundant initially, similar relative reductions to those observed would have provided a clear basis for interpretation. Operational experience in interior British Columbia suggests that where grass/legume mixtures have been used in silvicultural prescriptions, they can compete effectively with native vegetation. Speculation on the results found in this study, together with operational experience suggests that introduced, nitrogen-fixing legumes can compete with, and reduce the percent cover of some herbs and low-growing shrubs. 54 Furthermore, replacement of native species by nitrogen-fixing species may be additionally desirable if the net effect on the growth of the crop species is not negative. Wollum and Davey (1975) agreed that elimination of competing vegetation is beneficial to young forest stands, but added that some vegetation (N-fixers) may co-inhabit a site and improve the N status sufficiently to more than compensate for any detrimental effects it may have otherwise. The results presented in Section 2 showed no negative effect on survival or height increment of three-year-old lodgepole pine seedlings. The small but significant decrease in total and incremental diameter may no longer be observed once the seedlings substantially overtop the clover (Figure 17) and the newly fixed nitrogen begins to contribute to the soil-plant ecosystem. Klingler (1986) found growth reductions of Pseudotsuga menziesii [(Mirb.) Franco] due to grass/legume mixtures to no longer be significant after five years. Thus, one might conclude that legumes (on similar sites) will not negatively affect the crop tree growth, and may contribute to crop tree enhancement by reducing some competing species while increasing site nitrogen status. Continued assessments of this experiment will provide the data necessary to formulate more precise conclusions. Figure 17. Lodgepole pine seedlings and alsike clover in the third (1989) growing season. 55 5 M A N A G E M E N T A N D R E S E A R C H R E C O M M E N D A T I O N S 5.1 Management Cons idera t ions The ability to incorporate research results into management practices is at times more of an art than science. Early results may be misleading for long-term planning, since experimental treatment effects may have not yet been detected, or may change over time. Managers also have economic, social, and political constraints to consider in making decisions, as well as individual styles to decision making. In addition, some scientists may be more willing to recommend changes in operational practices based on early results and underlying principles and processes, while other scientists may be reluctant to make operational recommendations at all. The management recommendations that follow are derived from combining the results of this study with knowledge from the literature and related ecological principles of forest and soil sciences. The individual manager will have to decide on his/her own willingness to incorporate the recommendations into operational practice. Nevertheless, I strongly encourage foresters at all levels to consider the recommendations as one of many steps that can be taken to improve overall forest and soil management in British Columbia. 5.2 Management Recommendat ions Soils require plant communities to replace organic matter losses derived from harvesting and site preparation techniques that are most commonly used in British Columbia, such as - clearcutting and slashburning. The organic matter is needed to maintain and improve soil conditions that promote site productivity. This study was concerned with selecting a plant-endophyte symbiosis that would become established given prevailing site conditions, add newly fixed nitrogen to sites that are inherently nitrogen deficient and have experienced further nitrogen depletion from forestry practices, and finally, not to negatively compete with lodgepole pine regeneration. Results from this study are encouraging in that trends observed are, for the most part, consistent with these goals. It is therefore recommended that legume symbioses be established with lodgepole pine plantations on similar sites, where similar site and soil management goals have been defined. Although inoculated alsike clover proved successful in this study, it is recommended that a 56 combination of legume species (and appropriate inoculants) be established on any given plantation. This experiment was part of a larger biological nitrogen fixation research program (Trowbridge 1987b, Thomson and Trowbridge 1988a, 1988b, and 1989, Douglas and Trowbridge 1990). Early experience from that research and other observations from the Ministry of Forests Research and Range programs, lead me to recommend a seed mix including equal proportions by weight of alsike clover, Lotus corniculatus [cv. Leo (birdsfoot trefoil)], and Trifolium repens [cv. Grasslands huia, (white clover)] at 10 kg/ha. The clover spp. should establish relatively quickly, but not so aggressively as to negatively compete with the tree crop. Birdsfoot trefoil does not establish as quickly as the clovers, although it can persist longer. The use of a mix not only allows for early establishment and persistence in the plantation, it also adds a degree of species/endophyte diversity that may ensure a successful prescription over a wide range of environmental and site conditions. It is stressed that this is a preliminary species mix recommendation which could, and should, be modified as operational experience is gained. Furthermore, the biological nitrogen fixation research program has identified that the inoculant and the inoculating process are likely to be determining factosr in the success of nodule infection and effectiveness (personal observation). Regionally adapted, host specific Rhizobium should be applied to the seed as recommended in Trowbridge and Holl (1989). 5.3 Research Recommendations The British Columbia Ministry of Forests should continue to support the concept and practice of biological nitrogen fixation in forestry. Research must include forming and testing of hypotheses, and transferring the knowledge gained to practising foresters. In order to effectively accomplish this, there must be an active and focused research program, with field experiments and trials, and demonstration areas throughout the province. In addition, there must be adequate laboratory facilities (and related technologies) to isolate, culture, and produce regionally adapted endophytes. Specifically the following recommendations are made: 1. establish a network of standardized field screening trials consisting of nitrogen-fixing species/endophyte combinations with crop tree species in silviculturally important biogeoclimatic zones in British Columbia; 2. establish an extension program surrounding these field trials to encourage operational practice and to transfer the technological knowledge needed to ensure effective practices; 3. activelyt seek a government/university/industry cooperative approach to ensure applied research and commercial production of effective inoculants; and 4. the British Columbia Ministry of Forests should coordinate the above efforts to ensure effective integration of the sum of knowledge into ecologically sound management practices This approach would ensure a systematic screening of legume and non-legume host species/endophyte combinations appropriate for forestry across British Columbia, and allow meaningful predictions of effects on crop trees and site properties. 6 L ITERATURE CITED 58 Agriculture Canada Expert Committee on Soil Survey. 1987. 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Min. For. Re P-880 03-PR. Vict oria B.C 63 Turvey, N.D. and P.J. Smethurst. 1983. Nitrogen fixing plants in forest plantation management, pp. 233-60 In: Biological nitrogen fixation in forest ecosystems: foundations and applications. J.C. Gordon and C.T Wheeler (eds.). Martinus Nijkoff/Dr W. Junk Publishers, The Hague. Vincent, J.M. 1970. A manual for the practical study of the root-nodule bacteria. Blackwell Scientific Publishers, Oxford, England. I.B.P. Handb. No. 15. Virginia, R.A., W.M. Jarrell, P.W. Rundel, G Shearer, and K.H. Kohl. 1989. The use of variation in the natural abundance of 1 5N to assess symbiotic nitrogen fixation by woody plants, pp. 375-394 In: Stable isotopes in ecological research. P.W. Rundel, J.R. Ehleringer, and K.A. Nagy (eds.). Springer-Verlag, New York. Wollum, A.G., and C.B. Davey. 1975. Nitrogen accumulation, transformation, and transport in forest soils, pp. 67-106 In: Forest soils and forest land management. B. Bernier and C.H. Winget (eds.). Proceedings of the Fourth North American forest Soils Conference. Laval University, Quebec. Walmsley, M., G. Utzig, T. Void, D. Moon, and J . van Barneveld. 1980. Describing ecosystems in the field. RAB Tech. Paper 2, Land. Manage. Rep. No. 7, B.C. Min. Envior. and B.C. Min. For., Victoria, B.C. Yole, D. 1986. Progress report on forest nutrition in the Prince Rupert Forest Region. Int. Rep. Forest Sciences, B.C. Min. For. Int. Rep., Smithers, B.C. Yole, D., R. Carter, and R Trowbridge. 1989. Evaluation of five single tree fertilizer trials in thinned lodgepole pine stands in the Prince Rupert Forest Region. B.C. Min. For. Draft Ms., Smithers, B.C. Zar J.H. 1984. Biostatistical analysis. 2nd Ed. Prentice-Hall, Inc., Englewood Cliffs, N.J. 64 APPENDIX 1. Summary o f s t a t i s t i c a l a n a l y s i s f o r P i n u s contorta a n d Trifolium hybridum d a t a 65 Dependent Va r i ab le : Height increment 1987-88 Contrast DF NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 Contrast SS Mean Square F Value 0.83622685 11.24557870 2.54204167 4.98529660 0.83622685 11.24557870 2.54204167 4.98529660 19 54 57 12 Pr > F 0.6658 0.1171 0.4522 0.2937 Source BLOCK PREP PREP*BLOCK DF 2 2 4 Type III SS Mean Square F Value 150.1111574 17.0235130 145.9794926 75.0555787 8.5117565 36.4948731 3.04 0.34 1.48 Pr > F 0.0728 0.7129 0.2501 NTREAT NTREAT * BLOCK PREP*NTREAT PREP *NTREAT* BLOCK 3 6 6 12 14.6238472 1.6231167 56.3445389 31.0404556 4.8746157 0.2705194 9.3907565 2.5867046 1.10 0.06 2.12 0.58 0.3574 0.9990 0.0660 0.8458 NUM MEAN STDERR CV BLOCK 1 36 6.54b 0.57 52.32 2 36 8.74b 0.41 28.49 3 36 9.27a 0.51 32.79 PREP1 B 36 7.64a 0.32 25.17 S 36 8.58a 0.51 35.77 W 36 8.33a 0.70 50.72 1 27 8.55a 0.50 30.28 2 27 8.11a 0.64 40.99 3 27 7.61a 0.70 47.89 4 27 8.46a 0.63 38.71 1 S i te preparat ion treatments: B = broadcast burn, S = between windrow and, W = windrow 2 Nitrogen treatments: 1, 2, 3, and 4 = 0, 10, 20, and 30 kg/ha inocu lated a l s i ke c lover seed re spec t i ve l y 66 Dependent Va r i ab le : Height increment 1988-89 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 1 1 1 1 3.61785185 10.70370370 2.21696296 8.86716049 3.61785185 10.70370370 2.21696296 8.86716049 0.35 1.02 0.21 0.85 5594 3169 6475 3619 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 489.9674074 2 330.2335185 4 721.6503704 244.9837037 2.69 0.0951 165.1167593 1.81 0.1919 180.4125926 1.98 0.1409 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 16.538519 13.152593 131.793148 127.040741 5.512840 2.192099 21.965525 10.586728 0.53 0.21 2.09 1.01 0.6665 0.9725 0.0689 0.4533 NUM MEAN STDERR CV BLOCK 1 36 15.31a 0.87 33.93 2 36 20.20a 0.74 21.98 3 36 19.33a 1.23 38.24 PREP B 36 17.87a 0.97 32.43 S 36 16.37a 0.72 26.36 W 36 20.60a 1.22 35.57 NTREAT 1 27 17.79a 1.22 35.74 2 27 18.32a 1.09 30.91 3 27 18.87a 1.27 34.87 4 27 18.15a 1.20 34.35 67 Dependent Va r i ab le : Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT P REP * NT REAT * B LOCK Tota l height 1987 DF Contrast SS 1 1 1 1 DF 2 2 4 3 6 6 12 1.17880167 0.50566759 5.02861500 3.35093364 43.19606852 98.94831852 14.73861481 6.7130843 11.2891685 9.8824519 11.5617037 Mean Square F Value 1.17880167 0.50566759 5.02861500 3.35093364 0.39 0.17 1.67 1.11 Type III SS Mean Square F Value 21.59803426 3.12 49.47415926 7.16 3.68465370 0.53 2.2376948 0.74 1.8815281 0.62 1.6470753 0.55 0.9634753 0.32 Pr > F 0.5346 0.6839 0.2023 0.2967 Pr > F 0.0684 0.0052 0.7131 0.5320 0.7107 0.7711 0.9828 NUM MEAN STDERR CV BLOCK 1 36 17.19a 0.32 11.12 2 36 15.65a 0.35 13.51 3 36 16.61a 0.35 12.72 PREP B 36 17.21a 0.32 11.10 S 36 17.11a 0.28 9.97 W 36 15.13b 0.35 13.91 NTREAT 1 27 16.79a 0.36 11.02 2 27 16.17a 0.40 13.00 3 27 16.66a 0.41 12.91 4 27 16.32a 0.47 14.87 68 Dependent Var i ab le : Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK Tota l height 1988 DF Contrast SS 1 1 1 1 DF 2 2 4 3 6 6 12 4.32553500 16.96940833 0.71941500 17.71006944 92.8933685 76.4400574 161.9511093 22.0143583 14.3010833 66.0472389 32.1655278 Mean Square F Value 4.32553500 0.63 16.96940833 2.47 0.71941500 0.10 17.71006944 2.58 Type III SS Mean Square F Value 46.4466843 38.2200287 40.4877773 7.3381194 2 .3835139 11.0078731 2.6804606 1 .26 1 .03 1 .09 1 .07 0 .35 1.60 0 .39 Pr > F .4310 .1219 ,7475 0.1142 Pr > F 0.3086 0.3760 0.3890 3703 9086 1644 9614 NUM MEAN STDERR CV BLOCK 1 36 23.79a 0.62 15.55 2 36 24.46a 0.52 12.86 3 36 26.01a 0.68 15.71 PREP B 36 24.90a 0.54 13.02 S 36 25.71a 0.56 12.98 W 36 23.66a 0.73 18.48 NTREAT 1 27 25.46a 0.63 12.88 2 27 24.34a 0.68 14.59 3 27 24.38a 0.86 18.37 4 27 24.85a 0.71 14.80 69 Dependent Var i ab le : Contrast .NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK Tota l height 1989 DF Contrast SS 1 1 1 1 DF 2 2 4 3 6 6 12 0.03488074 0.75668148 5.59777852 1.58200494 843.637391 88.585946 1520.544376 6.389341 44.472209 85.943387 219.202246 Mean Square F Value 0.03488074 0.75668148 5.59777852 1.58200494 421.818695 44.292973 380.136094 2.129780 7.412035 14.323898 18.266854 0.00 0.04 0.29 0.08 Type III SS Mean Square F Value 1.83 0.19 1.65 0.11 0.38 0.74 0.94 P r > F 0.9663 0.8440 0.5929 0.7760 P r > F 0.1892 0.8269 0.2057 9539 8867 6195 0.5116 NUM MEAN STDERR CV BLOCK 1 36 39.10a 1.33 20.38 2 36 44.67a 1.05 14.15 3 36 45.34a 1.68 22.18 PREP B 36 42.77a 1.39 19.45 S 36 42.08a 1.13 16.14 W 36 44.25a 1.75 23.79 NTREAT 1 27 43.24a 1.68 20.19 2 27 42.65a 1.53 18.67 3 27 43.25a 1.80 21.67 4 27 42.99a 1.72 20.82 70 Dependent Va r i ab le : Diameter increment 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 2.76490667 0.77690370 0.00008963 3.22003086 2.76490667 0.77690370 0.00008963 3.22003086 10.30 2 .90 0 .00 12.00 0022 0946 9855 0010 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 21.61575741 2 0.08217963 4 15.25859259 10.80787870 4 .56 0.0250 0.04108981 0.02 0.9828 3.81464815 1 .61 0.2151 NTREAT NTREAT*BLOCK PREP*NTREAT PREP *NTREAT * BLOCK 3 6 6 12 3.54190000 0.26996111 2.22500556 1.99780000 1.18063333 0.04499352 0.37083426 0.16648333 4 .40 0 .17 1 .38 0.62 0.0077 0.9843 0.2386 0.8155 NUM MEAN STDERR CV BLOCK 1 36 2.18b 0.15 41.85 2 36 3.13a 0.11 21.43 3 36 3.13a 0.17 32.32 PREP B 36 2.84a 0.13 27.71 S 36 2.77a 0.14 30.09 W 36 2.82a 0.21 44.86 NTREAT 1 27 3.11a 0.18 29.56 2 27 2.80b 0.18 34.08 3 27 2.65b 0.20 39.83 4 27 2.68b 0.18 35.81 71 Dependent Variable: Diameter increment 1988-89 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 1 10.61201852 1 3.44898148 1 2.94816667 1 16.31262346 10.61201852 3.44898148 2.94816667 16.31262346 12.22 3.97 3.39 18.78 0.0010 0.0514 0.0709 0.0001 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 12.54018519 2 2.76907407 4 42.23092593 6.27009259 1.38453704 10.55773148 1.41 0.31 2.38 2694 7360 0904 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 17.00916667 6 4.93388889 6 9.10277778 12 13.44166667 5.66972222 0.82231481 1.51712963 1.12013889 6.53 0.95 1.75 1.29 0008 4699 1280 2518 NUM MEAN STDERR CV BLOCK 1 36 3.59a 0.24 40.06 2 36 4.38a 0.20 27.25 3 36 3.75a 0.27 43.77 PREP B 36 4.08a 0.25 36.45 S 36 3.69a 0.18 29.45 W 36 3.94a 0.29 44.42 NTREAT 1 27 4.58a 0.30 33.53 2 27 3.64b 0.26 36.48 3 27 3.81b 0.29 39.15 4 27 3.59b 0.26 37.42 72 Dependent Va r i ab le : Tota l diameter 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.00486000 0.00486000 0 .11 0 .7373 NITROGEN-QUADRATIC 1 0.04645926 0.04645926 1 .09 0.3018 NITROGEN-CUBIC 1 0.03683630 0.03683630 0 .86 0.3574 CONTROL VS OTHERS 1 0.01661235 0.01661235 0 .39 0.5357 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 0.38555741 0.19277870 0.98 0.3952 PREP 2 0.13562407 0.06781204 0.34 0.7135 PREP*BLOCK 4 0.17767037 0.04441759 0 .23 0.9207 NTREAT 3 0.08815556 0.02938519 0 .69 0.5637 NTREAT*BLOCK 6 0.17989444 0.02998241 0.70 0.6497 PREP*NTREAT 6 0.13625000 0.02270833 0 .53 0.7821 PREP *NTREAT* BLOCK 12 0.39810000 0.03317500 0.78 0 .6719 NUM MEAN STDERR CV BLOCK 1 36 3 . 5 5 a 0.04 6. 29 2 36 3 .51a 0.04 6. 30 3 36 3 .65a 0.05 8. 68 PREP B 36 3 . 5 3 a 0.04 6 .71 S 36 3. 62a 0.04 6 .37 W 36 3 . 5 6 a 0.05 8 .75 NTREAT 1 27 3 .55a 0.06 8 .16 2 27 3 . 5 6 a 0.05 6 .83 3 27 3 . 6 2 a 0.05 7.52 4 27 3 . 5 5 a 0.05 7 .05 73 Dependent Va r i ab le : Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK Tota l diameter 1988 DF 1 1 1 1 DF 2 2 4 3 6 6 12 Contrast SS Mean Square F Value 2. 0. 0. 2. 64040296 40578148 06490074 86361605 Type III SS 24.15321667 0.12335000 16.37143333 3.11108519 1.04184259 3.39715370 2.45461852 2.64040296 0.40578148 0.06490074 2.86361605 08 09 17 68 Mean Square F Value 12.07660833 4 .68 0.06167500 0 .02 4.09285833 1 .59 1.03702840 2 .78 0.17364043 0 .47 0.56619228 1.52 0.20455154 0 .55 Pr > F 0.0102 0 .3015 0.6782 0.0076 Pr > F 0.0231 0.9764 0.2209 0.0497 0 .8306 0.1897 0.8724 NUM MEAN STDERR CV BLOCK 1 36 5.75b 0.13 13.74 2 36 6.67ab 0.13 11.53 3 36 6.81a 0.20 17.63 PREP B 36 6.38a 0.15 14.07 S 36 6.39a 0.15 14.34 W 36 6.46a 0.22 20.18 NTREAT 1 27 6.69a 0.20 15.75 2 27 6.39b 0.19 15.43 3 27 6.31b 0.21 17.32 4 27 6.25b 0.20 16.86 74 Dependent Va r i ab le : Tota l diameter 1989 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 24.04978074 6.33653333 4.02486000 33.29290000 56.1197241 2.8447241 108.1921037 34.4111741 7.2134315 13.4777204 19.5215407 Mean Square F Value 24.04978074 6.33653333 4.02486000 33.29290000 28.0598620 1.4223620 27.0480259 11.4703914 1.2022386 2.2462867 1.6267951 16 .31 4 .30 2 .73 22 .57 Type III SS Mean Square F Value 2.21 0.11 2.13 7.78 0.82 1.52 1.10 Pr > F ,0002 .0430 ,1044 ,0001 Pr > F 1381 8945 1184 0.0002 0.5630 0.1883 0.3770 NUM MEAN STDERR CV BLOCK 1 36 9.34a 0.35 22.37 2 36 11.05a . 0.30 16.40 3 36 10.56a 0.42 24.05 PREP B 36 10.46a 0.37 21.14 S 36 10.09a 0.30 17.55 W 36 10.40a 0.46 26.50 NTREAT 1 27 11.28a 0.44 20.30 2 27 10.03b 0.39 20.37 3 27 10.12b 0.45 23.35 4 27 9.84b 0.42 22.31 75 Dependent Va r i ab le : Percent su rv i va l 1988 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP *NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 15.57201852 15.94675926 92.83557407 61.62250000 1347.532407 4825.556296 469.232037 124.354352 661.240926 559.508148 3093.065741 Mean Square F Value 15.57201852 15.94675926 92.83557407 61.62250000 0 .07 0 .07 0.42 0.28 Type III SS Mean Square F Value 673.766204 2 .43 2412.778148 8.70 117.308009 0.42 41.451451 0 .19 110.206821 0.50 93.251358 0.42 257.755478 1 .16 Pr > F 0.7924 0.7900 0.5212 0.6009 Pr > F 0.1165 0.0023 0.7902 9053 8092 8633 0.3362 NUM MEAN STDERR CV BLOCK 1 36 85.16a 2.47 17.38 2 36 81.91a 2.69 19.68 3 36 76.59a 2.81 22.05 PREP B 36 79.14b 2.85 21.59 S 36 90.24a 1.64 10.93 W 36 74.27b 2.77 22.35 NTREAT 1 27 79.91a 3.43 22.28 2 27 82.67a 2.81 17.67 3 27 80.53a 3.18 20.51 4 27 81.76a 3.18 20.20 76 Dependent Var i ab le : Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK Percent su r v i v a l 1989 DF Contrast SS 1 0.89629630 1 6.25925926 1 56.06666667 1 1.49382716 DF Type III SS 2 226.462963 2 5469.462963 4 706.203704 Mean Square F Value Pr > F 0.89629630 0.00 0.9570 6.25925926 0.02 0.8867 56.06666667 0.18 0.6701 1.49382716 0.00 0.9445 Mean Square F Value Pr > F 113.231481 0.24 0. 2734.731481 5.87 0. 176.550926 0.38 0.8209 7869 0109 NTREAT NTREAT*BLOCK PREP*NTREAT P REP * NTREAT * BLOCK 3 6 6 12 63.222222 772.722222 393.055556 3759.500000 21.074074 128.787037 65.509259 313.291667 0.07 0.42 0.21 1.03 9762 8615 9707 4398 NUM MEAN STDERR CV BLOCK 1 36 77.81a 3.40 26.21 2 36 79.89a 2.53 18.97 3 36 76.36a 3.27 25.68 PREP B 36 78.72ab 3.22 24.54 S 36 86.36a 1.93 13.40 W 36 68.97b 3.22 28.01 NTREAT 1 27 77.81a 3.08 20.55 2 27 78.70a 3.24 21.42 3 27 76.85a 3.71 25.07 4 27 78.70a 4.23 27.89 77 Dependent Va r i ab le : Needle mass (g/100) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.34020540 NITROGEN-QUADRATIC 1 0.14622848 NITROGEN-CUBIC 1 0.00078723 CONTROL VS OTHERS 1 0.44266844 0.34020540 0.14622848 0.00078723 0.44266844 14.13 6.07 0.03 18.39 0.0004 0.0169 0.8572 0.0001 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP 0.24981902 0.11647369 1.72805276 0.12490951 0.05823684 0.43201319 0.64 0.30 17 .94 0.5382 0.7452 0.0001 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 0.48722111 0.04307794 0.32991350 0.12998961 0.16240704 0.00717966 0.05498558 0.01083247 6.75 0.30 2 .28 0 .45 0006 9351 0489 0.9345 NUM MEAN STDERR CV BLOCK 1 36 0.87a 0.05 37.22 2 36 0.97a 0.03 20.02 3 36 0.96a 0.05 28.65 PREP B 36 0.96a 0.05 28.48 S 36 0.89a 0.03 20.97 W. 36 0.95a 0.06 35.18 NTREAT 1 27 1.05a 0.05 26.00 2 27 0.93b 0.05 28.70 3 27 0.87b 0.05 31.38 4 27 0.90b 0.05 28.59 78 Dependent Va r i ab le : Percent cover Trifolium 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 22512.67490 22512.67490 183.82 0 .0001 NITROGEN-QUADRATIC 1 5960.54424 5960.54424 48 .67 0.0001 NITROGEN-CUBIC 1 1190.12860 1190.12860 9.72 0 .0029 CONTROL VS OTHERS 1 28799.34705 28799.34705 235 .15 0.0001 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 2 4 889.288709 775.584234 3735.927341 444.644354 387.792117 933.981835 1 .03 0.90 2 .16 3771 4248 1146 NTREAT NTREAT*BLOCK PREP*NTREAT PREP *NTREAT * BLOCK 3 29663.34774 6 931.98135 6 375.19972 12 2023.36587 9887.78258 155.33023 62.53329 168.61382 80 .73 1 .27 0 .51 1 .38 0.0001 0.2874 0.7977 0 .2057 NUM MEAN STDERR CV BLOCK 1 36 30.23a 3.86 76.56 2 36 24.23a 3.03 75.13 3 36 30.39a 4.14 81.64 PREP B 36 24.90a 3.86 92.93 S 36 31.45a 3.45 65.77 W 36 28.50a 3.81 80.20 NTREAT 1 27 0.00c 0.00 2 27 33.71ab 3.65 56.25 3 27 37.72ab 2.96 40.73 4 27 41.71a 3.30 41.07 79 Dependent Variable: Percent cover Trifolium 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 22512.67490 5960.54424 1190.12860 28799.34705 22512.67490 5960.54424 1190.12860 28799.34705 183.82 48.67 9.72 235.15 0001 0001 0029 0001 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 889.288709 775.584234 3735.927341 444.644354 387.792117 933.981835 1.03 0.90 2.16 3771 4248 1146 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 29663.34774 6 931.98135 6 375.19972 12 2023.36587 9887.78258 155.33023 62.53329 168.61382 80.73 1.27 0.51 1.38 0001 2874 7977 0.2057 NUM MEAN STDERR CV BLOCK 1 36 30.23a 3.86 76.56 2 36 24.23a 3.03 75.13 3 36 30.39a 4.14 81.64 PREP B 36 24.90a 3.86 92.93 S 36 31.45a 3.45 65.77 W 36 28.50a 3.81 80.20 NTREAT 1 27 0.00a 0.00 2 27 33.71a 3.65 56.25 3 27 37.72a 2.96 40.73 4 27 41.71a 3.30 41.07 80 Dependent Va r i ab le : Percent cover Trifolium 1989 Contrast DF NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 Contrast SS Mean Square F Value 76362.3375 34400.5208 6759.2782 117211.1304 76362.3375 34400.5208 6759.2782 117211.1304 1279.17 576.25 113.23 1963.44 Pr > F 0.0001 0.0001 0.0001 0.0001 Source BLOCK PREP PREP*BLOCK DF 2 2 4 Type III SS Mean Square F Value 306.004630 834.949074 1441.509259 153.002315 417.474537 360.377315 1.14 3.12 2.70 Pr > F 3405 0685 0639 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 117522.1366 6 289.6065 6 350.1065 12 940.7130 39174.0455 48.2677 58.3511 78.3927 656.22 0.81 0.98 1.31 0.0001 0.5678 0.4495 0.2385 NUM MEAN STDERR CV BLOCK 1 36 58.01a 5.90 61.02 2 36 58.47a 5.93 60.84 3 36 54.69a 5.55 60.92 PREP B 36 56.08a 5.80 62.04 S 36 54.25a 5.43 60.06 W 36 60.85a 6.10 60.19 NTREAT 1 27 0.00b 0.00 2 27 73.63a 2.33 16.42 3 27 76.19a 1.93 13.18 4 27 78.43a 2.19 14.52 81 Dependent Va r i ab le : Volume Trifolium (m3/m2) 1987 Contrast DF Contrast SS Mean Square F Value NITROGEN-LINEAR 1 0.00041178 0.00041178 5 .96 NITROGEN-QUADRATIC 1 0.00002374 0.00002374 0.34 NITROGEN-CUBIC 1 0.00000186 0.00000186 0 .03 CONTROL VS OTHERS 1 0 .00017575 0.00017575 2 .54 Pr > F 0.0180 0.5602 0.8704 0.1166 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 0.00040465 0.00033084 0.00084689 0.00020233 0.00016542 0.00021172 1.08 0 .88 1 .13 3619 4320 3754 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 0.00043738 0.00043899 0.00040277 0.00087436 0.00014579 0.00007317 0.00006713 0.00007286 2.11 1.06 0.97 1.05 1097 3987 4535 4157 NUM MEAN STDERR CV BLOCK 1 2 3 36 36 36 0 .00495a 0 .00088a 0 .00080a 0.00285 0.00029 0.00028 345.90 216.97 187.60 PREP B S W 36 36 36 0 .00041a 0 .00163a 0 .00458a 0.00015 0.00034 0.00287 211.40 125.82 375.20 NTREAT 1 2 3 4 27 27 27 27 0.00000a 0.00104a 0.00244a 0.00563a 0.00000 0.00037 0.00147 0.00354 185.90 313.39 343.43 82 Dependent Variable: Voume Trifolium (m3/m2) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.15114384 0.15114384 76.31 0.0001 NITROGEN-QUADRATIC 1 0.03579758 0.03579758 18.07 0.0001 NITROGEN-CUBIC 1 0.00708809 0.00708809 3.58 0.0639 CONTROL VS OTHERS 1 0.18672401 0.18672401 94.28 0.0001 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 2 4 0.02190621 0.00478617 0.07060037 0.01095311 0.00239308 0.01765009 1.27 0.28 2.05 0.3043 0.7606 0.1302 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 0.19402951 0.01713463 0.00632601 0.03421821 0.06467650 0.00285577 0.00105434 0.00285152 32.66 1.44 0.53 1.44 0001 2160 7813 1771 NUM MEAN STDERR CV BLOCK 1 36 0.08a 0.01 108.12 2 36 0.05a 0.01 84.84 3 36 0.08a 0.01 102.09 PREP B 36 0.06a 0.01 129.20 S 36 0.07a 0.01 69.88 W 36 0.08a 0.01 113.81 NTREAT 1 27 0.00b 0.00 2 27 0.08a 0.01 83.81 3 27 0.10a 0.01 75.34 4 27 0.11a 0.01 70.74 83 Dependent Var i ab le : Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP *NTREAT * BLOCK Atom-percent 15N isotope 1988 DF 1 1 1 1 DF 2 2 4 3 6 6 12 Contrast SS Mean Square F Value 1.62032667 0.10453333 0.00626963 0.60666790 31.00901296 58.55346852 26.57846481 1.73112963 10.58614259 9.73188704 16.82689074 1.62032667 0.10453333 0.00626963 0.60666790 .58 ,10 ,01 ,59 Type III SS Mean Square F Value 15.50450648 3.14 29.27673426 5.93 6.64461620 1.35 0.57704321 0.56 1.76435710 1.72 1.62198117 1.58 1.40224090 1.36 Pr > F 0.2147 0.7510 0.9380 0.4457 Pr > F 0.0675 0.0105 0.2913 0.6428 0.1348 0.1714 0.2119 NUM MEAN STDERR CV BLOCK 1 2 3 36 36 36 8.25b 9.49a 9.26a 0.29 0.27 0.23 21.31 17.28 14.84 PREP B BURN M SCRA WIND B 36 36 36 9.87a 8.07b 9.07ab 0.21 0.24 0.30 13.01 17.94 19.54 NTREAT 1 2 3 4 27 27 27 27 9.13a 9.10a 8.97a 8.81a 0.26 0.30 0.35 0.39 14.53 16.92 20.24 22.78 84 Dependent Var i ab le : Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK Tota l n i trogen (dry wt.%) 1988 DF 1 1 1 1 DF 2 2 4 3 6 6 12 Contrast SS 00384000 02370370 02507852 00000000 0.15222407 0.63653519 0.12198148 0.05262222 0.13062778 0.25420556 0.39994444 Mean Square F Value 0.00384000 0.02370370 0.02507852 0.00000000 0.07611204 0.31826759 0.03049537 0.01754074 0.02177130 0.04236759 0.03332870 19 18 25 00 Type III SS Mean Square F Value 1.38 5 .76 0 .55 0.88 1 .09 2 .11 1 .66 Pr > F 6633 2815 2682 0000 Pr > F 2778 0117 7005 0.4596 0.3821 0.0664 0.1017 NUM MEAN STDERR CV BLOCK 1 36 1.41a 0.03 13.23 2 36 1.50a 0.03 12.93 3 36 1.48a 0.03 12.17 PREP B BURN 36 1.36b 0.02 10.12 M SCRA 36 1.55a 0.03 11.33 WIND B 36 1.48ab 0.03 13.74 NTREAT 1 27 1.46a 0.04 14.98 2 27 1.47a 0.03 11.50 3 27 1.43a 0.04 14.26 4 27 1.49a 0.03 10.98 APPENDIX 2. Summary o f s t a t i s t i c a l a n a l y s i s f o r f o r e s t f l o o r c h e m i s t r y d a t a 86 Dependent Variable: Percent nitrogen 1987 Contrast DF Contrast SS NITROGEN-LINEAR 1 . 0 .00997556 NITROGEN-QUADRATIC 1 0.00490000 NITROGEN-CUBIC 1 0.00288000 CONTROL VS OTHERS 1 0.01080000 Source DF NTREAT 3 0 .01775556 BLOCK*NTREAT 6 0.01872778 Mean Square F Value 0.00997556 0.00490000 0.00288000 0.01080000 0.00591852 0.00312130 1 .26 0 .62 0 .36 1 .37 Type I II SS Mean Square F Value 0.75 0.40 Pr > F 0.2758 0.4411 0.5534 0.2574 Pr > F 0.5367 0.8724 NUM MEAN STDERR CV BLOCK 1 12 0 .73a 0.03 12 .57 2 12 0 .83a 0.02 8 .97 3 12 0 . 8 9 a 0.06 23 .84 NTREAT1 1 9 0.85a 0.06 20.35 2 9 0.83a 0.05 18.81 3 9 0.79a 0.05 19.41 4 9 0.81a 0.05 18.89 1 Nitrogen treatments: 1, 2, 3, and 4 = 0, 10, 20, and 30 kg/ha inoculated a l s i k e clover seed res p e c t i v e l y 87 Dependent Va r i ab le : Percent n i trogen 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.00016056 0.00016056 0 .01 0.9122 NITROGEN-QUADRATIC 1 0.00266944 0.00266944 0 .21 0 .6539 NITROGEN-CUBIC 1 0 .01073389 0.01073389 0.84 0 .3727 CONTROL VS OTHERS 1 0.00004537 0.00004537 0.00 0 .9533 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 6 0.01356389 0.06862778 0.00452130 0.01143796 35 89 0.7882 0.5222 NUM MEAN STDERR CV BLOCK 1 12 0 .82a 0.02 10 .43 2 12 0 .93a 0.03 12 .33 3 12 0 .87a 0.05 18 .05 NTREAT 1 9 0 .87a 0.03 10 .99 2 9 0 .86a 0 .05 17 .31 3 9 0 .90a 0.05 16 .94 4 9 0 .85a 0.04 14 .26 88 Dependent Va r i ab le : Percent n i trogen d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.00760500 0.00760500 0 .35 0.5603 NITROGEN-QUADRATIC 1 0.01480278 0.01480278 0 .69 0.4186 NITROGEN-CUBIC 1 0 .02473389 0.02473389 1 .15 0.2987 CONTROL VS OTHERS 1 0.00944537 0.00944537 0.44 0.5168 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 6 0.04714167 0.10086667 0.01571389 0.01681111 0 .73 0.78 0.5487 0.5974 NUM MEAN STDERR CV BLOCK 1 12 0 .09a 0.03 127 .55 2 12 0 .09a 0.04 161 .27 3 12 - 0 . 0 3 a 0.05 - 6 9 2 . 7 6 NTREAT 1 9 0 .02a 0.06 834.34 2 9 0 .03a 0.05 544 .67 3 9 0 .11a 0.05 139 .93 4 9 0 .04a 0.04 336 .61 89 Dependent Variable: Nitrogen (kg/ha) 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 1 1090.272222 1 354.694444 1 266.450000 1 1039.120370 1090.272222 1.51 0.2354 354.694444 0.49 0.4928 266.450000 0.37 0.5515 1039.120370 1.44 0.2463 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 6 1711.41667 1582.16667 570.47222 263.69444 0.79 0.36 0.5159 0.8919 NUM MEAN STDERR CV BLOCK 1 2 3 12 12 12 232.83a 199.75a 277.17a 8.46 5.19 19.09 12 .59 8 .99 23 .86 NTREAT 1 2 3 4 245.89a 239.56a 227.33a 233.56a 20.51 18.98 16.94 16.12 25.03 23.77 22.36 20.71 90 Dependent Va r i ab le : Nitrogen (kg/ha) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 1 5 .6888889 1 128.4444444 1 952.2000000 1 10.7037037 5.6888889 128.4444444 952.2000000 10.7037037 0.01 0.12 0.90 0 .01 0.9424 0.7320 0.3561 0.9211 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 6 1086.33333 5635.50000 362.11111 939.25000 0.34 0.88 0.7958 0.5258 NUM MEAN STDERR CV BLOCK 1 12 217 .00a 6.58 1 0 . 5 1 2 12 254 .08a 9.03 12 .32 3 12 268 .42a 13 .96 18 .02 NTREAT 1 9 247.44a 12.33 14.95 2 9 241.67a 13.77 17.09 3 9 255.11a 15.38 18.09 4 9 241.78a 15.09 18.72 91 Dependent Variable: Mineralizable N forest f l o o r (kg/ha corrected f o r mineral s o i l exposure) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 365.5125000 365.5125000 4 .65 0.0448 NITROGEN-QUADRATIC 1 7.0225000 7.0225000 0.09 0.7684 NITROGEN-CUBIC 1 53.9013889 53.9013889 0 .69 0.4184 CONTROL VS OTHERS 1 230.2712037 230.2712037 2 .93 0.1041 Source DF Type III SS Mean Square F Value Pr > F NTREAT 3 426.436389 142.145463 1 .81 0.1816 BLOCK*NTREAT 6 483.586111 80.597685 1 .03 0.4404 NUM MEAN STDERR CV BLOCK 1 12 25.01a 1.86 25.77 2 12 27.33a 1.92 24.39 3 12 40.40a 5.10 43.77 NTREAT 1 9 26.53b 2.93 33.14 2 9 30.69ab 3.12 30.51 3 9 30.26ab 3.12 30.96 4 9 36.18a 7.06 58.54 92 Dependent Variable: Percent carbon 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 31.41688889 14.95111111 1.92200000 3.06703704 31.41688889 14.95111111 1.92200000 3.06703704 0.97 0 .46 0 .06 0 .09 0.3374 0.5052 0.8102 0.7617 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 6 48.2900000 133.5350000 16.0966667 22.2558333 0.50 0 .69 0.6885 0.6620 NUM MEAN STDERR CV BLOCK 1 12 48.67a 1.46 10.41 2 12 49.11a 2.31 16.27 3 12 46.98a 1.39 10.22 NTREAT 1 9 48.76a 2.72 16.73 2 9 49.62a 1.79 10.83 3 9 48.17a 1.44 8.99 4 9 46.46a 2.06 13.33 93 Dependent Va r i ab le : Percent carbon 1988 Contrast DF Contrast SS NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 Source DF NTREAT 3 BLOCK*NTREAT 6 5.51250000 0.96694444 11.50138889 10.64083333 Type III SS 17.9808333 239.9816667 Mean Square F Value 5.51250000 0.22 0.96694444 0.04 11.50138889 0.47 10.64083333 0 .43 Mean Square F Value 5.9936111 39.9969444 0.24 1 .62 Pr > F 0.6421 0.8453 0.5035 0.5197 Pr > F 0.8652 0.1986 NUM MEAN STDERR CV BLOCK 1 2 3 12 12 12 3 9 . 3 6 a 4 1 . 2 3 a 3 6 . 0 0 a 1.44 1.57 1 .75 12 .64 13 .20 16 .80 NTREAT 1 9 3 7 . 9 2 a 1.70 13 .44 2 9 3 9 . 6 1 a 2 .40 18 .14 3 9 3 8 . 4 4 a 2.08 16 .24 4 9 3 9 . 4 8 a 1.74 13 .23 Dependent Va r i ab le : Percent carbon d i f f e rence 1987-88 Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source NTREAT BLOCK*NTREAT DF Contrast SS 1 63.24938889 1 8.12250000 1 4 .02005556 1 25.32675926 DF Type III SS 3 75.3919444 6 371.8105556 Mean Square F Value 63.24938889 1 .05 8.12250000 0 .13 4.02005556 0 .07 25.32675926 0.42 Mean Square F Value 25.1306481 0 .42 61.9684259 1 .03 NUM MEAN STDERR CV BLOCK 1 12 - 9 . 3 1 a 2 .26 - 8 4 . 0 0 2 12 - 7 . 9 2 a 2 .19 - 9 5 . 9 7 3 12 - 1 0 . 9 8 a 2 .04 - 6 4 . 4 9 NTREAT 1 9 - 1 0 . 8 6 a 2 .48 - 6 8 . 4 6 2 9 - 1 0 . 0 2 a 2 .37 - 7 0 . 8 6 3 9 - 9 . 7 3 a 2 .57 - 7 9 . 1 0 4 9 - 7 . 0 0 a 2 .69 - 1 1 5 . 2 7 95 ependent Variable: Carbon-nitrogen r a t i o 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 1.5867222 1.5867222 0.02 0.8763 NITROGEN-QUADRATIC 1 101.0025000 101.0025000 1.59 0.2237 NITROGEN-CUBIC 1 15.7827222 15.7827222 0.25 0.6244 CONTROL VS OTHERS 1 14.4467593 14.4467593 0.23 0.6394 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 6 118.371944 482.090556 39.457315 80.348426 0.62 1.26 0.6109 0.3223 NUM MEAN STDERR CV BLOCK 1 12 67.42a 2.14 10.99 2 12 59.92a 3.90 22.54 3 12 55.49a 4.21 26.29 NTREAT 1 9 59.84a 5.45 27.34 2 9 61.82a 4.17 20.23 3 9 63.41a 4.48 21.21 4 9 58.69a 3.47 17.72 96 Dependent Variable: Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source NTREAT BLOCK*NTREAT Carbon-nitrogen r a t i o 1988 DF 1 1 1 1 DF 3 6 Contrast SS Mean Square F Value 16.14005556 1.91361111 94.32272222 23.24083333 112.3763889 86.5994444 16.14005556 1.63 1.91361111 0.19 94.32272222 9.53 23.24083333 2.35 Type III SS Mean Square F Value 37.4587963 14.4332407 3.78 1.46 Pr > F 0.2179 0.6654 0.0064 0.1429 Pr > F 0.0289 0.2479 NUM MEAN STDERR CV BLOCK 1 12 48.30a 1.13 8.11 2 12 44.60a 1.02 7.94 3 12 41.88a 1.33 11.04 NTREAT 1 9 43.53ab 1.27 8.77 2 9 46.57a 1.99 12.81 3 9 42.82b 0.95 6.64 4 9 46.78a 1.72 11.03 97 Dependent Variable: Carbon-nitrogen r a t i o d i f f e r e n c e 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 28.4808889 28 .4808889 0.36 0.5563 NITROGEN-QUADRATIC 1 130.7211111 130.7211111 1.65 0.2153 NITROGEN-CUBIC 1 184 .4268889 184 .4268889 2.33 0.1445 CONTROL VS OTHERS 1 1 .0800000 1.0800000 0.01 0 .9084 Source DF Type III SS Mean Square F Value Pr > F NTREAT 3 343.628889 114.542963 1.45 0.2627 BLOCK*NTREAT 6 863.444444 143.907407 1.82 0.1525 NUM MEAN STDERR CV BLOCK 1 12 -19.10a 2.24 -40 .66 2 12 -15.32a 3.93 -88 .82 3 12 -13.62a 3.97 -101 .00 NTREAT 1 9 -16.31a 4.76 -87.46 2 9 -15.28a 3.69 -72.40 3 9 -20.56a 4.17 -60.81 4 9 -11.90a 3.27 -82.37 98 Dependent Variable: Phosphorus (kg/ha) 1987 Contrast DF Contrast SS NITROGEN-LINEAR 1 4.80200000 NITROGEN-QUADRATIC 1 0.58777778 NITROGEN-CUBIC 1 0.64800000 CONTROL VS OTHERS 1 5.51259259 Source DF NTREAT 3 6.0377778 BLOCK*NTREAT 6 14.9305556 Mean Square F Value 4.80200000 0.58777778 0.64800000 5.51259259 2.0125926 2.4884259 0.76 0.09 0.10 0.87 Type III SS Mean Square F Value 0.32 0.39 Pr > F 0.3944 0.7637 0.7523 0.3622 Pr > F 0.8114 0.8729 NUM MEAN STDERR CV BLOCK 1 12 15.63a 0.89 19.81 2 12 8.05b 0.50 21.57 3 12 9.86b 0.61 21.28 NTREAT 1 9 10.50a 0.98 28.09 2 9 11.32a 1.45 38.34 3 9 11.29a 1.40 37.33 4 9 11.60a 1.64 42.46 99 Dependent Va r i ab le : Phosphorus (kg/ha) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 24.93888889 0.21777778 12.80000000 7.15592593 24.93888889 0.21777778 12.80000000 7.15592593 8.47 0.07 4.35 2.43 0.0093 0 .7887 0.0516 0.1364 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 37.95666667 6 31.92500000 12.65222222 4 .30 0.0188 5.32083333 1 .81 0.1543 NUM MEAN STDERR CV BLOCK 1 12 11.18a 0.64 19.88 2 12 12.19a 0.67 19.09 3 12 11.81a 0.74 21.72 NTREAT 1 9 12.50a 0.69 16.47 2 9 12.98a 0.90 20.74 3 9 10.63b 0.86 24.36 4 9 10.80b 0.36 9.86 100 Dependent Variable: Phosphorus (kg/ha) differ e n c e 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 51.84200000 51.84200000 5.69 0.0282 NITROGEN-QUADRATIC 1 0.11111111 0.11111111 0.01 0.9133 NITROGEN-CUBIC 1 7.52355556 7.52355556 0.83 0.3754 CONTROL VS OTHERS 1 25.61814815 25.61814815 2.81 0.1108 Source DF Type III SS Mean Square F Value Pr > F NTREAT 3 59.4766667 19.8255556 2.18 0.1261 BLOCK*NTREAT 6 35.6950000 5.9491667 0.65 0.6874 NUM MEAN STDERR CV BLOCK 1 12 -4.43b 1.14 -89.44 2 12 4.16b 0.73 60.97 3 12 1.92b 1.06 190.32 NTREAT 1 9 2.01a 1.34 200.22 2 9 1.64a 1.56 284.66 3 9 -0.66a 2.03 -929.86 4 9 -0.80a 1.69 -635.04 101 Dependent Va r i ab le : Potassium (kg/ha) 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 69.44022222 69.44022222 5 .26 0.0341 NITROGEN-QUADRATIC 1 0.58777778 0.58777778 0.04 0 .8353 NITROGEN-CUBIC 1 3 .36200000 3.36200000 0 .25 0.6200 CONTROL VS OTHERS 1 54.32925926 54.32925926 4 .11 0.0576 Source DF Type III SS Mean Square F Value Pr > F NTREAT 3 73.390000 24 .463333 1 .85 0.1738 BLOCK*NTREAT 6 78.078333 13.013056 0 .99 0.4637 NUM MEAN STDERR CV BLOCK 1 12 43.56a 1.30 10.31 2 12 17.04b 1.57 31.97 3 12 22.72b 0.97 14.73 NTREAT 1 9 25.64a 4.31 50.41 2 9 27.69a 4.34 47.04 3 9 28.11a 4.54 48.43 4 9 29.64a 3.91 39.56 102 Dependent Variable: Potassium (kg/ha) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 15.60555556 15.60555556 2.33 0.1443 NITROGEN-QUADRATIC 1 0.01777778 0.01777778 0.00 0.9595 NITROGEN-CUBIC 1 0.93888889 0.93888889 0.14 0.7125 CONTROL VS OTHERS 1 11.47259259 11.47259259 1.71 0.2071 Source DF Type III SS Mean Square F Value Pr > F NTREAT 3 16.5622222 5.5207407 0.82 0.4975 BLOCK*NTREAT 6 67.8227778 11.3037963 1.69 0.1814 NUM MEAN STDERR CV BLOCK 1 12 19.27a 0.80 14. 46 2 12 19.62a 1.15 20. 38 3 12 25.07a 0.83 11. 47 NTREAT 1 9 20.34a 1.05 15. 46 2 9 21.27a 1.20 16. 88 3 9 21.42a 1.92 26. 88 4 9 22.26a 1.41 18. 97 103 Dependent Va r i ab le : Potassium (kg/ha) d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 19.47022222 19.47022222 0.74 0.4021 NITROGEN-QUADRATIC 1 0.40111111 0.40111111 0.02 0.9033 NITROGEN-CUBIC 1 0.72200000 0.72200000 0.03 0.8706 CONTROL VS OTHERS 1 16.02370370 16.02370370 0.61 0.4464 Source . DF Type III SS Mean Square F Value Pr > F NTREAT 3 20.593333 6.864444 0.26 0.8535 BLOCK*NTREAT 6 132 . 628333 22.104722 0.84 0.5580 NUM MEAN STDERR CV BLOCK 1 12 -24.29b 1.30 -18. 59 2 12 2.57a 1.66 223. 83 3 12 2.35a 1.18 174. 29 NTREAT 1 9 -5.30a 4.42 -250.23 2 9 -6.42a 4.09 -191.17 3 9 -6.70a 5.46 -244.58 4 9 -7.40a 4.89 -198.09 104 Dependent Variable: Calcium (kg/ha) 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 256.3280000 256.3280000 0.17 0.6864 NITROGEN-QUADRATIC 1 67.2400000 67.2400000 0.04 0.8359 NITROGEN-CUBIC 1 234.8408889 234.8408889 0.15 0.6991 CONTROL VS OTHERS 1 444.8948148 444.8948148 0.29 0.5954 Source DF Type III SS Mean Square F Value Pr > F NTREAT 3 558.4089 186.1363 0.12 0.9458 BLOCK*NTREAT 6 1228.8494 204.8082 0.13 0.9899 NUM MEAN STDERR CV BLOCK 1 12 250.86a 13.15 18. 16 2 12 115.21c 4.95 14. 87 3 12 175.00b 10.09 19. 97 NTREAT 1 9 174.27a 21.08 36.29 2 9 183.96a 21.69 35.37 3 9 179.49a 22.49 37.58 4 9 183.71a 25.71 41.99 105 Dependent Variable: Calcium (kg/ha) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 1 194.480056 1 42.466944 1 1044.976056 1 236.740833 194.480056 42.466944 1044.976056 236.740833 0.20 0.04 1.08 0.24 0.6600 0.8368 0.3135 0.6276 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 6 1281.92306 10082.46444 427.30769 1680.41074 0.44 1.73 0.7274 0.1715 NUM MEAN STDERR CV BLOCK 1 12 180.39b 9.77 18.77 2 12 183.91b 8.88 16.72 3 12 229.86a 10.16 15.31 NTREAT 1 9 193.61a 9.54 14.78 2 9 203.16a 13.16 19.43 3 9 190.78a 15.55 24.46 4 9 204.67a 15.46 22.66 106 Dependent Va r i ab le : Calcium (kg/ha) d i f f e rence 1987-88 Contrast DF NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC .1 CONTROL VS OTHERS 1 Contrast SS 4.2013889 218.5469444 290.0680556 32.7800926 Mean Square F Value Pr > F 4.2013889 0.00 0.9660 218.5469444 0 .10 0.7586 290.0680556 0 .13 0.7235 32.7800926 0.01 0.9052 Source NTREAT BLOCK*NTREAT DF 3 6 Type III SS 512.8164 15353.5244 Mean Square F Value 170.9388 2558.9207 0.08 1.14 Pr > F 0.9721 0.3797 NUM MEAN STDERR CV BLOCK 1 2 3 12 -70.46b 16.31 -80.20 12 68.73a 9.24 46.57 12 54.84a 14.76 93.25 NTREAT 1 2 3 4 19.36a 19.20a 11.28a 20.98a 23 .96 20 .35 27 .38 35 .47 371.38 317 .89 728 .23 507 .32 107 Dependent Va r i ab le : Magnesium (kg/ha) 1987 Contrast DF Contrast SS NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 Source DF NTREAT 3 BLOCK*NTREAT 6 13.01422222 15.47111111 14.33688889 2.25333333 Type III SS 42.822222 112.856111 Mean Square F Value 13.01422222 0.42 15.47111111 0.50 14.33688889 0.46 2.25333333 0.07 Mean Square F Value 14.274074 18.809352 0 .46 0.60 Pr > F 5265 4903 5064 7912 Pr > F 0.7153 0.7246 NUM MEAN STDERR CV BLOCK 1 12 33.66a 2.00 20.57 2 12 16.81b 0.99 20.42 3 12 23.83b 1.50 21.79 NTREAT 1 9 24.33a 3.41 42.10 2 9 24.69a 2.42 29.37 3 9 23.53a 2.44 31.16 4 9 26.51a 3.61 40.87 108 Dependent Variable: Magnesium (kg/ha) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 1 99.16088889 1 0.69444444 1 29.76800000 1 92.22259259 99.16088889 4.33 0.0521 0.69444444 0.03 0.8638 29.76800000 1.30 0.2694 92.22259259 4.02 0.0601 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 129.6233333 6 225.1950000 43.2077778 37.5325000 1.89 1.64 0.1683 0.1941 NUM MEAN STDERR CV BLOCK 1 12 18.76a 0.99 18.27 2 12 22.60a 1.35 20.73 3 12 26.56a 2.23 29.07 NTREAT 1 9 19.87b 1.25 18.82 2 9 23.26ab 1.72 22.24 3 9 22.30ab 2.20 29.64 4 . 9 25.13a 2.85 34.07 109 Dependent Va r i ab le : Magnesium (kg/ha) d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 40.61250000 23.20027778 2.81250000 66.42675926 40.61250000 23.20027778 2.81250000 66.42675926 0.77 0.44 0 .05 1 .26 0.3920 0.5158 0 .8201 0.2768 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 6 66.625278 405.143889 22.208426 67.523981 0.42 1 .28 0.7405 0.3157 NUM MEAN STDERR CV BLOCK 1 12 -14.88b 2.19 -50.97 2 12 5.78a 1.36 81.28 3 12 2.72a 2.76 351.46 NTREAT 1 9 -4.48a 4.01 -268.36 2 9 -1.42a 2.41 -508.39 3 9 -1.22a 3.42 -839.12 4 9 -1.38a 5.76 -1254.43 110 Dependent Va r i ab le : Cat ion exchange capac i ty (meq/lOOg) 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0 .00088889 0.00088889 0.00 0.9964 NITROGEN-QUADRATIC 1 36.40111111 36.40111111 0 .87 0.3630 NITROGEN-CUBIC 1 2 .40355556 2.40355556 0 .06 0.8132 CONTROL VS OTHERS 1 9.36333333 9.36333333 0.22 0.6416 Source DF Type III SS Mean Square F Value Pr > F NTREAT 3 38.8055556 12.9351852 0 .31 0.8182 BLOCK*NTREAT 6 91.7427778 15.2904630 0 .37 0.8910 NUM MEAN STDERR CV BLOCK 1 12 85 .24a 1 .73 7.02 2 12 90 .73a 1.89 7 .21 3 12 91 .35a 2 .05 7 .77 NTREAT 1 9 88. 2 9 89. 3 9 90. 4 9 87. 22a 2 .44 8 .30 77a 2 .53 8.44 46a 2 .39 7 .91 98a 2 .21 7.54 I l l Dependent Variable: Cation exchange capacity (meq/lOOg) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 2.04800000 2.04800000 0.03 0.8650 NITROGEN-QUADRATIC 1 7.11111111 7.11111111 0.10 0.7517 NITROGEN-CUBIC 1 59.39755556 59.39755556 0.86 0.3654 CONTROL VS OTHERS 1 21.51148148 21.51148148 0.31 0.5832 Source DF Type III SS Mean Square F Value Pr > F NTREAT 3 68.556667 22.852222 0.33 0.8025 BLOCK*NTREAT 6 626.785000 104.464167 1.52 0.2289 NUM MEAN STDERR CV BLOCK 1 12 7 4 . 5 4 a 2 .14 9 .92 2 12 89 .45a 2 .82 10 .93 3 12 8 4 . 1 9 a 4 .49 18 .46 NTREAT 1 9 81.39a 2.69 9.90 2 9 84.79a 4.97 17.60 3 9 81.56a 4.46 16.39 4 9 83.18a 5.11 18.44 112 Dependent Variable: Cation exchange capacity (meq/lOOg) di f f e r e n c e 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 2.13422222 2.13422222 0.01 0.9094 NITROGEN-QUADRATIC 1 11.33444444 11.33444444 0.07 0.7934 NITROGEN-CUBIC 1 85.69800000 85.69800000 0.53 0.4742 CONTROL VS OTHERS 1 2.49037037 2.49037037 0.02 0.9022 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 6 99.166667 899.825000 33.055556 149.970833 0.21 0.94 0.8908 0.4941 NUM MEAN STDERR CV BLOCK 1 12 -10.70a 2.85 -92.36 2 12 -1.28a 3.99 -1083.17 3 12 -7.16a 3.85 -186.16 NTREAT 1 9 -6.83a 2.80 -122.77 2 9 -4.98a 5.02 -302.48 3 9 -8.90a 3.96 -133.34 4 9 -4.80a 5.36 -334.88 113 Dependent Variable: pH 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0 .00280056 0 .00280056 0.02 0 .8965 NITROGEN-QUADRATIC 1 0.02300278 0 .02300278 0.14 0 .7097 NITROGEN-CUBIC 1 0 .00122722 0 .00122722 0.01 0 .9314 CONTROL VS OTHERS 1 0.01893426 0.01893426 0.12 0.7355 Source DF Type III SS Mean Square F Value Pr > F NTREAT 3 0.02703056 0.00901019 0.06 0.9820 BLOCK*NTREAT 6 0.46822778 0.07803796 0.49 0 .8107 NUM MEAN STDERR CV BLOCK 1 12 4.70a 0.11 8 .25 2 12 4.57a 0.10 7 .29 3 12 4.97a 0 .12 8 .49 NTREAT 1 9 4.71a 0.08 4.86 2 9 4.77a 0.17 10.57 3 9 4.77a 0.15 9.40 4 9 4.73a 0.16 9.83 114 Dependent Va r i ab le : pH 1988 Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS DF Contrast SS 1 0.04080056 1 0.00422500 1 0.06536056 1 0.00280093 Mean Square F Value 0.04080056 0.00422500 0.06536056 0.00280093 1 .07 0 .11 1 .72 0 .07 Pr > F 0.3138 0.7426 0.2062 0.7891 Source DF Type III SS Mean Square F Value Pr > F NTREAT BLOCK*NTREAT 3 6 0.11038611 0.70483889 0.03679537 0.11747315 0 .97 3 .09 0.4292 0.0292 NUM MEAN STDERR CV BLOCK 1 12 5 . 1 6 a 0.10 6 .85 2 12 4 . 8 9 a 0.07 5 .16 3 12 4 . 9 4 a 0.08 5 .80 NTREAT 1 9 5 . 0 1 a 0.11 6 .49 2 9 5 . 0 8 a 0.12 7 .26 3 9 4 . 9 3 a 0.08 4 .96 4 9 4 . 9 6 a 0.12 7.00 115 Dependent Variable: pH differe n c e 1987-88 Contrast DF Contrast SS NITROGEN-LINEAR 1 0.06498000 NITROGEN-QUADRATIC 1 0.00751111 NITROGEN-CUBIC 1 0.04867556 CONTROL VS OTHERS 1 0.03630000 Source DF NTREAT 3 0.12116667 BLOCK*NTREAT 6 1.20140000 Mean Square F Value 0.06498000 0.00751111 0.04867556 0.03630000 0.04038889 0.20023333 0.32 0.04 0.24 0 .18 Type III SS Mean Square F Value 0.20 0.99 Pr > F 0.5770 0.8490 0 .6289 0.6761 Pr > F 0.8946 0.4584 NUM MEAN STDERR CV BLOCK 1 12 0.46a 0.14 106.94 2 12 0.32a 0.12 135.55 3 12 -0.03a 0.11 -1243.98 NTREAT 1 9 0.30a 0.13 132.63 2 9 0.30a 0.18 180.67 3 9 0.17a 0.14 248.84 4 9 0.22a 0.20 264.03 APPENDIX 3 . Summary of s t a t i s t i c a l analysis f o r mineral s o i l chemistry data 117 Dependent Variable: Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK % Nitrogen 1987 DF Contrast SS 1 1 1 1 DF 2 2 4 3 6 6 12 0.00002241 0.00026759 0.00000907 0.00002500 Type III SS 0.00129630 0.02319074 0.03138148 0.00029907 0.00110370 0.00040926 0.00179630 Mean Square F Value 0.00002241 0.00026759 0.00000907 0.00002500 0.00064815 0.01159537 0.00784537 0.00009969 0.00018395 0.00006821 0.00014969 0.10 1.15 0.04 0.11 Mean Square F Value 0.25 4.54 3.07 0.43 0.79 0.29 0.64 Pr > F 0.7580 0.2893 0.8445 0.7448 Pr > F 0.7786 0.0253 0.0431 0.7347 0.5837 0.9382 0.7981 NUM MEAN STDERR CV BLOCK 1 36 0.07a 0.01 56.23 2 36 0.07a 0.00 29.58 3 36 0.07a 0.01 48.89 PREP B MIN 36 0.06b 0.01 59.68 M SCRAP 36 0.06b 0.00 32.78 WINDROW 36 0.09a 0.01 36.05 NTREAT 1 27 0.07a 0.01 46.87 2 27 0.07a 0.01 52.65 3 27 0.07a 0.01 41.77 4 27 0.07a 0.01 49.18 1 S i t e preparation treatments: B MIN = broadcast burn mineral s o i l , M SCRAP = between windrow mineral s o i l and, WINDROW = windrow mineral s o i l 2 Nitrogen treatments: 1, 2, 3, and 4 = 0, 10, 20, and 30 kg/ha inoculated a l s i k e clover seed re s p e c t i v e l y 118 Dependent Variable: % Nitrogen 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.00127574 0.00127574 2.11 0.1518 NITROGEN-QUADRATIC 1 0 .00011204 0.00011204 0.19 0 .6683 NITROGEN-CUBIC 1 0 .00031130 0 .00031130 0.52 0.4758 CONTROL VS OTHERS 1 0.00146944 0.00146944 2.43 0.1246 Source DF Type III ,SS Mean Square F Value Pr > F BLOCK 2 0 .00397963 0 .00198981 0.88 0 .4322 PREP 2 0 .03324630 0 .01662315 7.35 0.0047 PREP*BLOCK 4 0.04243704 0.01060926 4.69 0 .0091 NTREAT 3 0 .00169907 0 .00056636 0.94 0 .4287 NTREAT*BLOCK 6 0 .00566481 0.00094414 1.56 0 .1756 PREP*NTREAT 6 0.00166481 0.00027747 0.46 0 .8350 PREP*NTREAT*BLOCK 12 0 .00529630 0 .00044136 0 .73 0 .7150 NUM MEAN STDERR CV BLOCK 1 36 0.07a 0.01 73 . 08 2 36 0.07a 0.01 46. 44 3 36 0.06a 0.00 44. 59 PREP B MIN 36 0.05b 0.00 47. 60 M SCRAP 36 0.06b 0.00 43. 36 WINDROW 36 0.09a 0.01 55. 33 NTREAT 1 27 0 .07a 0 .01 54 .93 2 27 0 .07a 0.01 63.84 3 27 0 .07a 0.01 53 .90 4 27 0 . 0 6 a 0.01 63 .61 119 Dependent Variable: Nitrogen (kg/ha) 1987 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 2250.93750 44591.02083 940.10417 5967.56250 Mean Square F Value 2250.93750 0 .06 44591.02083 1.14 940.10417 0.02 5967.56250 . 0 .15 Type III SS Mean Square F Value 271158.875 7255881.375 5063175.250 47782.063 192797.125 71537.125 286730.750 135579.438 0 .41 3627940.687 11 .05 1265793.812 3 .86 15927.354 0 .41 32132.854 0.82 11922.854 0.30 23894.229 0 .61 Pr > F 0.8116 0.2912 0.8776 0.6981 Pr > F 0.6677 0.0007 0.0196 0.7494 0.5603 0.9323 0.8253 NUM MEAN STDERR CV BLOCK 1 36 948 .75a 100.20 63 .37 2 36 826 .92a 50 .02 36 .30 3 36 874 .92a 63.87 43 .80 PREP B MIN 36 606.58b 60.32 59.67 M SCRAP 36 814.00b 44.47 32.78 WINDROW 36 1230.00a 73.90 36.05 NTREAT 1 27 870.67a 81.20 48.46 2 27 901.94a 96.29 55.47 3 27 905.78a 81.20 46.58 4 27 855.78a 87.73 53.27 120 Dependent Variable: Nitrogen (kg/ha) 1988 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT P REP * NTREAT * BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 218406.6667 22794.0833 51333.7500 257725.4444 823546.292 9465698.667 7448788.083 292534.500 908356.375 306915.000 923818.250 Mean Square F Value 218406.6667 2.22 22794.0833 0.23 51333.7500 0.52 257725.4444 2.61 Type III SS Mean Square F Value 411773.146 4732849.333 1862197.021 97511.500 151392.729 51152.500 76984.854 1.09 12.57 4.95 0.99 1.54 0.52 0.78 Pr > F 0.1424 0.6325 0.4736 0.1117 Pr > F 0.3561 0.0004 0.0072 4048 1842 7914 0.6670 NUM MEAN STDERR CV BLOCK 1 36 943.46a 125.99 80.13 2 36 888.04a 77.23 52.18 3 36 736.82a 55.83 45.4 PREP B MIN 36 505.99b 40.14 47.59 M SCRAP 36 832.33b 60.15 43.36 WINDROW 36 1230.00a 113.43 55.33 NTREAT 1 27 940.72a 111.08 61.35 .2 27 832.44a 112.26 70.07 3 27 850.72a 93.81 57.30 4 27 800.56a 108.66 70.53 121 Dependent Va r i ab le : Nitrogen (kg/ha) d i f f e rence 1987-88 Contrast DF NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 Contrast SS Mean Square F Value 176312.6042 131147.5208 38380.1042 342127.5069 176312.6042 1 .13 131147.5208 0.84 38380.1042 0.24 342127.5069 2 .18 Pr > F 0.2935 0.3643 0.6226 0 .1453 Source BLOCK PREP PREP*BLOCK DF 2 2 4 Type III SS Mean Square F Value 741077.167 295351.042 1270637.833 370538.583 147675.521 317659.458 1 .26 0.50 1 .08 Pr > F 0.3074 0.6134 0.3954 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 345840.229 525894.833 603804.458 938493.167 115280.076 87649.139 100634.076 78207.764 0.74 0 .56 0.64 0.50 5352 7605 6958 9061 NUM MEAN STDERR CV BLOCK 1 36 - 5 . 2 4 a 65.51 - 7 5 0 6 . 7 9 2 36 61 .19a 71.58 701.82 3 36 - 1 3 8 . 0 8 a 68 .23 - 2 9 6 . 4 PREP B MIN 36 - 1 0 0 . 4 6 a 51 .08 - 3 0 5 . 0 6 M SCRAP 36 18 .33a 64 .87 2123.02 WINDROW 36 0 .00a 87 .23 NTREAT 1 27 7 0 . 0 6 a 90 .59 671.92 2 27 - 6 9 . 5 0 a 90.07 - 6 7 3 . 4 2 3 27 - 5 5 . 0 6 a 69.24 - 6 5 3 . 4 6 4 27 - 5 5 . 2 2 a 68.61 - 6 4 5 . 6 3 122 Dependent Variable: Mineralizable nitrogen (kg/ha) 1988 Contrast DF NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 Contrast SS Mean Square F Value 308.4177963 33.4445370 3.8001667 95.2792901 308.4177963 33.4445370 3.8001667 95.2792901 5.39 0.58 0.07 1.66 Pr > F 0.0241 0.4479 0.7976 0.2025 Source BLOCK PREP PREP*BLOCK DF 2 2 4 Type III SS Mean Square F Value 3862.15796 19810.48685 17874.73926 1931.07898 9905.24343 4468.68481 0.87 4.46 2.01 Pr > F 0.4357 0.0266 0.1355 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 345.66250 1498.20056 522.67167 1072.39778 115.22083 249.70009 87.11194 89.36648 2.01 4.36 1.52 1.56 1230 0012 1886 1315 NUM MEAN STDERR CV BLOCK 1 2 3 36 36 36 35.92a 24.75a 22.13a 7.63 1.85 2.28 127.40 44.75 61.82 PREP B MIN 36 12.77b 1.20 56.54 M SCRAP 36 24.53ab 1.65 40.40 WINDROW 36 45.51a 7.07 93.18 NTREAT 1 27 25.97a 4.55 91.09 2 27 26.04a 5.70 113.67 3 27 28.05a 5.41 100.25 4 27 30.34a 6.51 111.53 123 Dependent Variable: % Carbon 1987 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 0.02166000 1.53129259 0.01102519 0.73197531 5.24963519 91.66300185 41.75167037 1.56397778 3.28021667 2.22747222 5.42396667 Mean Square F Value 0.02166000 1.53129259 0.01102519 0.73197531 2.62481759 45.83150093 10.43791759 0.52132593 0.54670278 0.37124537 0.45199722 0.04 2.81 0.02 1.34 Type III SS Mean Square F Value 1.52 26.56 6.05 0.96 1.00 0.68 0.83 Pr > F 0.8428 0.0997 0.8875 0.2519 Pr > F 0.2452 0.0001 0.0029 0.4205 0.4338 0.6661 0.6213 NUM MEAN STDERR CV BLOCK 1 36 2.76a 0.32 69.29 2 36 2.29a 0.17 44.00 3 36 2.28a 0.19 48.64 PREP B MIN 36 1.57b 0.12 46.94 M SCRAP 36 2.05b 0.15 43.81 WINDROW 36 3.72a 0.24 38.83 NTREAT 1 27 2.30a 0.22 48.81 2 27 2.57a 0.33 66.25 3 27 2.56a 0.27 54.71 4 27 2.35a 0.27 59.69 124 Dependent Va r i ab le : Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK % Carbon 1988 DF 1 1 1 1 DF 2 2 4 3 6 6 12 Contrast SS Mean Square F Value 00000463 80655648 22611574 18446944 30.0561685 147.4743907 95.7929648 3.0326769 17.3920537 9.8314537 24.4787907 0.00000463 2.80655648 0.22611574 1.18446944 15.0280843 73.7371954 23.9482412 1.0108923 2.8986756 1.6385756 2.0398992 00 82 15 77 Type III SS Mean Square F Value 1.87 9.17 2.98 0.66 1.88 1.06 1.32 Pr > F 0.9986 0.1830 0.7033 0.3848 Pr > F 0.1830 0.0018 0.0474 5832 1013 3966 2336 NUM MEAN STDERR CV BLOCK 1 36 3.14a 0.54 103.53 2 36 2.04a 0.27 80.37 3 36 2.01a 0.22 65.76 PREP B MIN 36 1.40b 0.09 39.51 M SCRAP 36 1.76b 0.09 31.43 WINDROW 36 4.04a 0.55 82.39 NTREAT 1 27 2.58a 0.39 77.93 2 27 2.18a 0.47 113.12 3 27 2.30a 0.38 86.27 4 27 2.54a 0.52 105.98 125 Dependent Va r i ab le : Carbon-Nitrogen Ratio 1987 Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS DF 1 1 1 1 Contrast SS Mean Square F Value 4.89632667 76.54117037 0.07443630 46.72481975 4.89632667 76.54117037 0.07443630 46.72481975 0.19 2.91 00 78 Pr > F 0.6680 0.0939 0.9578 0.1883 Source BLOCK PREP PREP*BLOCK DF Type III SS 2 154.082457 2 3226.740457 4 80.190448 Mean Square F Value Pr > F 77.041229 0.92 0.4170 1613.370229 19.24 0.0001 20.047612 0.24 0.9125 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 81.511933 102.543861 86.578928 299.445611 27.170644 17.090644 14.429821 24.953801 1.03 0.65 0.55 0.95 0.3857 0.6904 0.7691 0.5080 NUM MEAN STDERR CV BLOCK 1 36 35.15a 1.50 25.60 2 36 33.87a 1.23 21.77 3 36 32.23a 1.28 23.87 PREP B MIN 36 27.74b 0.69 14.91 M SCRAP 36 32.55b 0.93 17.17 WINDROW 36 40.96a 1.27 18.66 NTREAT 1 27 32.61a 1.24 19.81 2 27 34.53a 1.67 25.15 3 27 34.65a 1.72 25.84 4 27 33.20a 1.58 24.71 126 Dependent Variable: Carbon-Nitrogen Ratio 1988 Contrast DF Contrast SS Mean Square F ' Value Pr • > F NITROGEN-LINEAR 1 771. 4442313 771.4442313 1.91 0. 1724 NITROGEN-QUADRATIC 1 181. 1446009 181.1446009 0.45 0. 5057 NITROGEN-CUBIC 1 15. 2644891 15.2644891 0.04 0. 8465 CONTROL VS OTHERS 1 162. 1802250 162.1802250 0.40 0. 5287 Source DF Type III SS Mean Square F Value Pr • > F BLOCK 2 2348 .437039 1174.218519 1.23 0. 3160 PREP 2 4152 .581450 2076.290725 2.17 0. 1427 PREP*BLOCK 4 5290 .336061 1322.584015 1.38 0. 2789 NTREAT 3 967.85332 322.61777 0.80 0. 4995 NTREAT*BLOCK 6 243.50922 40.58487 0.10 0. 9960 PREP*NTREAT 6 2709.11948 451.51991 1.12 0. 3636 PREP*NTREAT*BLOCK 12 3180.46746 265.03895 0.66 0. 7836 NUM MEAN STDERR CV BLOCK 1 36 39.62 a 3.21 48. 62 2 36 30.34 a 2.36 46. 58 3 36 40.76 a 5.31 78. 20 PREP B MIN 36 32.55a 2.70 49. 82 M SCRAP 36 32.49a 2.89 53. 35 WINDROW 36 45.67a 5.19 68. 24 NTREAT 1 27 34.78a 2.84 42. 49 2 27 33.91a 4.69 71. 82 3 27 37.31a 4.21 58. 58 4 27 41.62a 5.77 72. 07 127 Dependent Va r i ab le : Carbon-Nitrogen Ratio d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 652.1641849 652.1641849 1.60 0.2111 NITROGEN-QUADRATIC 1 491.5200000 491.5200000 1.21 0.2768 NITROGEN-CUBIC 1 17.4506694 17.4506694 0.04 0.8368 CONTROL VS OTHERS 1 34.8414738 34.8414738 0.09 0.7710 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 2 4 2704.723237 556.687491 4996.290840 1352.361618 278.343745 1249.072710 1.27 0.26 1.17 0.3060 0.7735 0.3573 NTREAT NTREAT*BLOCK PREP*NTREAT PREP *NTREAT * BLOCK 3 6 6 12 1161.13485 382.27615 2650.35632 3828.30751 387.04495 63.71269 441.72605 319.02563 0.95 0.16 1.08 0.78 4229 9869 3833 6648 NUM MEAN STDERR CV BLOCK 1 2 3 36 36 36 4.48a -3.51a 8.53a 3.39 2.44 5.10 454.76 -416.35 358.78 PREP B MIN 36 4.82a 2.55 317.05 M SCRAP 36 -0.04a 3.02 -41048.5 WINDROW 36 4.73a 5.44 690.81 NTREAT 1 27 2.18a 2.91 693.43 2 27 -0.61a 4.57 -3923.68 3 27 2.67a 4.44 863.37 4 27 8.42a 5.58 344.74 128 Dependent Va r i ab le : Phosphorous (kg/ha) 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 1 324.4770150 1 715.9530083 1 11.8429646 1 5.6829262 324.4770150 1.19 0.2796 715.9530083 2.63 0.1106 11.8429646 0.04 0.8355 5.6829262 0.02 0.8856 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 3885.62479 2 57182.17302 4 3334.50113 1942.81240 0.98 0.3950 28591.08651 14.40 0.0002 833.62528 0.42 0.7922 NTREAT NTREAT*BLOCK PREP*NTREAT PREP *NTREAT * BLOCK 3 6 6 12 1052.27299 1148.35045 1019.09195 2748.58391 350.75766 191.39174 169.84866 229.04866 1.29 0.70 0.62 0.84 0.2874 0.6480 0.7100 0.6081 NUM MEAN STDERR CV BLOCK 1 36 65.24a 7.11 65.37 2 36 55.34a 4.53 49.15 3 36 50.89a 4.66 54.95 PREP B MIN 36 33. M SCRAP 36 50. WINDROW 36 88. 00b 2.29 41.61 34b 2.89 34.47 12a 6.07 41.35 NTREAT 1 27 56.76a 6.28 57.47 2 27 60.95a 7.47 63.68 3 27 58.51a 6.58 58.45 4 27 52.40a 5.66 56.14 129 Dependent Va r i ab le : Phosphorous (kg/ha) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 1 1 1 1 738.434002 5392.290712 476.561378 4772.276669 738.434002 5392.290712 476.561378 4772.276669 0.87 6.34 0.56 5.61 0.3557 0.0148 0.4575 0.0215 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 18659.9470 2 118681.2624 4 9051.7896 9329.9735 59340.6312 2262.9474 3.34 21.22 0.81 ,0585 ,0001 ,5354 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 6607.2861 8349.2291 11309.0399 10016.7932 2202.4287 1391.5382 1884.8400 834.7328 2.59 1.64 2.22 0.98 0623 1553 0554 4783 NUM MEAN STDERR CV BLOCK 1 36 86.68a 9.89 68.48 2 36 58.11a 6.13 63.26 3 36 59.54a 8.44 85.03 PREP B MIN 36 39. M SCRAP 36 50. WINDROW 36 114. 36b 2.73 41.56 41b 4.05 48.21 55a 10.17 53.25 NTREAT 1 27 79.62a 12.24 79.89 2 27 59.39a 8.89 77.75 3 27 62.69a 7.08 58.67 4 27 70.73a 10.44 76.72 130 Dependent Variable: Phosphorous (kg/ha) differ e n c e 1987-88 Contrast DF NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 Contrast SS Mean Square F Value 84.41157 10035.04725 640.26667 5110.26408 84.41157 10035.04725 640.26667 5110.26408 0.06 7.59 0.48 3.86 Pr > F 0.8015 0.0080 0.4896 0.0545 Source BLOCK PREP PREP*BLOCK DF 2 2 4 Type III SS Mean Square F Value Pr > F 6554.76852 13651.44814 2458.90500 3277.38426 6825.72407 614.72625 1.65 3.44 0.31 2197 0544 8678 NTREAT NTREAT*BLOCK PREP*NTREAT P REP * NTREAT * BLOCK 3 10759.72549 6 9334.44297 6 17326.90675 12 12591.12707 3586.57516 1555.74050 2887.81779 1049.26059 2.71 1.18 2.18 0.79 0.0539 0.3327 0.0587 0.6551 NUM MEAN STDERR CV BLOCK 1 36 21.45a 5.97 166.99 2 36 2.78a 5.18 1117.09 3 36 8.66a 8.66 600.11 PREP B MIN 36 6.36a 2.82 265.77 M SCRAP 36 0.09a 3.21 21957.47 WINDROW 36 26.45a 10.66 241.87 NTREAT 1 27 22.88a 11.63 264.19 2 27 -1.55a 4.51 -1515.95 3 27 4.20a 5.45 675.26 4 27 18.33a 7.45 211.12 131 Dependent Variable: Potassium (kg/ha) 1987 i Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 3182.088375 4478.504823 542.703375 0.916274 11012.9547 571071.0193 249270.1915 8203.2966 25715.1920 24063.2325 42940.5456 Mean Square F Value 3182.088375 1.41 4478.504823 1.98 542.703375 0.24 0.916274 0.00 Type III SS Mean Square F Value 5506.4774 285535.5097 62317.5479 2734.4322 4285.8653 4010.5388 3578.3788 0.21 10.76 2.35 1.21 1.89 1.77 1.58 Pr > F 0.2411 0.1654 0.6265 0.9840 Pr > F 0.8146 0.0008 0.0934 0.3159 0.0989 0.1226 0.1255 NUM MEAN STDERR CV BLOCK 1 36 232.00a 24.49 63.35 2 36 243.08a 14.57 35.96 3 36 256.69a 19.88 46.47 PREP B MIN 36 184.58b 19.54 63.52 M SCRAPE 36 200.86b 8.34 24.90 WINDROW 36 346.33a 17.65 30.58 NTREAT 1 27 243.76a 22.02 46.93 2 27 255.80a 29.15 59.22 3 27 244.93a 22.39 47.50 4 27 231.20a 18.31 41.14 132 Dependent Va r i ab le : Potassium (kg/ha) 1988 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 1542.057409 2784.754890 157.021511 3323.458445 34463.8499 830951.5864 221070.5755 4483.8338 24949.9711 30368.4970 60160.1416 Mean Square F Value 1542.057409 2784.754890 157.021511 3323.458445 17231.9249 415475.7932 55267.6439 1494.6113 4158.3285 5061.4162 5013.3451 0.51 0.93 0.05 1.11 Type III SS Mean Square F Value 0.98 23.73 3.16 0.50 1.38 1.69 1.67 Pr > F 0.4767 0.3398 0.8200 0.2974 Pr > F 0.3929 0.0001 0.0394 0.6854 0.2375 0.1423 0.1001 NUM MEAN STDERR CV BLOCK 1 36 268.77a 28.42 63.45 2 36 239.05a 16.73 41.99 3 36 226.10a 14.88 39.48 PREP B MIN 36 168.06b 10.78 38.48 M SCRAPE 36 198.42b 6.88 20.81 WINDROW 36 367.44a 22.66 36.99 NTREAT 1 27 254.25a 25.32 51.76 2 27 242.87a 25.54 54.65 3 27 236.25a 22.18 48.78 4 27 245.19a 24.63 52.19 133 Dependent Va r i ab le : Potassium (kg/ha) d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 890.35128 890.35128 0.21 0.6508 NITROGEN-QUADRATIC 1 25598.19022 25598.19022 5.96 0.0180 NITROGEN-CUBIC 1 83.64630 83.64630 0.02 0.8895 CONTROL VS OTHERS 1 4475.38700 4475.38700 1.04 0.3120 Source DF Type III SS Mean Square F Value Pr > 1 BLOCK 2 40756.68210 20378.34105 1.76 0.1999 PREP 2 12883.11128 6441.55564 0.56 0.5823 PREP*BLOCK 4 36193.03930 9048.25983 0.78 0.5510 NTREAT 3 26572.1878 8857.3959 2.06 0.1161 NTREAT*BLOCK 6 25142.4766 4190.4128 0.98 0.4508 PREP*NTREAT 6 80036.8516 13339.4753 3.11 0.0110 PREP*NTREAT*BLOCK 12 50379.5380 4198.2948 0.98 0.4816 NUM MEAN STDERR CV BLOCK 1 36 32.93a 16.34 297.74 2 36 -4.93a 11.45 -1392.69 3 36 -10.97a 11.60 -634.72 PREP B MIN 36 -16.52a 1.35 -490.32 M SCRAPE 36 -2.44a 7.98 -1964.37 WINDROW 36 21.11a 22.12 628.62 NTREAT 1 27 -9.82a 17.37 536.61 2 27 16.82a 19.93 -1053.90 3 27 -9.62a 8.97 -484.42 4 27 25.32a 14.04 288.13 134 Dependent Variable: Calcium (kg/ha) 1987 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP *NTREAT * BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 0.7526 30048.0192 169831.4244 31.9602 2006791.669 9569311.165 1126326.143 199880.20 440746.69 310603.04 623173.32 Mean Square F Value 0.7526 0.00 30048.0192 0.23 169831.4244 1.30 31.9602 0.00 Type III SS Mean Square F Value 1003395.834 1.43 4784655.582 6.83 281581.536 0.40 66626.73 0.51 73457.78 0.56 51767.17 0.40 51931.11 0.40 Pr > F 0.9981 0.6335 0.2593 0.9876 Pr > F 0.2647 0.0062 0.8047 0.6772 0.7584 0.8783 0.9586 NUM MEAN STDERR CV BLOCK 1 36 870.98a 90.68 62.47 2 36 1007.81a 63.25 37.65 3 36 1203.17a 114.55 57.13 PREP B MIN 36 720.83b 76.56 63.73 M SCRAP 36 930.66ab 54.43 35.09 WINDROW 36 1430.47a 102.57 43.02 NTREAT 1 27 1028.26a 98.91 49.98 2 27 990.76a 122.28 64.13 3 27 1097.24a 114.13 54.05 4 27 993.02a 101.52 53.12 135 Dependent Va r i ab le : Calcium (kg/ha) 1988 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 60515.50417 41728.74454 72164.45602 20156.11188 13316555.76 59786089.52 49523810.70 174408.7 3241840.9 2945938.9 4801753.1 Mean Square F Value 60515.50417 0.34 41728.74454 0.23 72164.45602 0.40 20156.11188 0.11 Type III SS Mean Square F Value 6658277.88 1.03 29893044.76 4.63 12380952.68 1.92 58136.2 0.32 540306.8 3.01 490989.8 2.74 400146.1 2.23 Pr > F ,5636 ,6314 ,5284 ,7387 Pr > F 0.3765 0.0238 0.1510 0.8077 0.0129 0.0214 0.0226 NUM MEAN STDERR CV BLOCK 1 36 1880.02a 412.04 131.50 2 36 1081.93a 91.75 50.88 3 36 1203.23a 131.45 65.55 PREP B MIN 36 719.66b 58.12 48.45 M SCRAP 36 1019.22ab 68.30 40.21 WINDROW 36 2426.29a 388.00 95.95 NTREAT 1 27 1364.73a 292.98 111.55 2 27 1392.83a 349.26 130.30 3 27 1344.64a 258.89 100.04 4 27 1451.37a 307.78 110.19 136 Dependent Va r i ab le : Calcium (kg/ha) d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 59676.6241 59676.6241 0.16 0.6886 NITROGEN-QUADRATIC 1 142826.4468 142826.4468 0.39 0.5357 NITROGEN-CUBIC 1 463167.4619 463167.4619 1.26 0.2666 CONTROL VS OTHERS 1 21535.8886 21535.8886 0.06 0.8097 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 22765364.13 2 21897309.15 4 48635808.58 11382682.07 10948654.58 12158952.14 1.90 1.83 2.03 1786 1896 1334 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 665670.5 4287242.9 4104807.2 7216118.4 221890.2 714540.5 684134.5 601343.2 0.60 1.94 1.86 1.64 6155 0903 1046 1091 NUM MEAN STDERR CV BLOCK 1 36 1009.02a 391.28 232.67 2 36 74.30a 82.69 667.73 3 36 0.11a 101.74 99999.00 PREP B MIN 36 -1.03a 44.54 -25836.8 M SCRAP 36 88.63a 52.09 352.62 WINDROW 36 995.83a 407.78 245.69 NTREAT 1 27 336.68a 297.72 459.48 2 27 402.12a 362.22 468.05 3 27 247.43a 230.62 484.31 4 27 458.33a 253.19 287.05 137 Dependent Variable: Magnesium (kg/ha) 1987 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 551.480378 7890.453675 305.357120 816.975185 67266.0196 674743.6653 265555.2161 8747.291 20944.618 16165.685 47841.951 Mean Square F Value 551.480378 0.19 7890.453675 2.78 305.357120 0.11 816.975185 0.29 Type III SS Mean Square F Value 33633.0098 0.53 337371.8326 5.29 66388.8040 1.04 2915.764 1.03 3490.770 1.23 2694.281 0.95 3986.829 1.41 Pr > F ,6610 ,1011 ,7441 ,5937 Pr > F 5993 0156 4140 0.3876 0.3053 0.4678 0.1922 NUM MEAN STDERR CV BLOCK 1 36 217.53a 30.62 84.47 2 36 204.73a 10.83 31.74 3 36 262.90a 28.26 64.51 PREP B MIN 36 158.76b 23.47 88.70 M SCRAP 36 187.46ab 14.11 45.16 WINDROW 36 338.93a 24.94 44.15 NTREAT 1 27 223.62a 25.76 59.85 2 27 235.69a 32.12 70.81 3 27 238.18a 30.48 66.50 4 27 216.05a 28.06 67.47 138 Dependent Va r i ab le : Magnesium (kg/ha) 1988 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 2424.475045 73.227268 779.112735 675.624497 140872.161 1335963.691 1024358.538 3276.815 94878.531 102070.522 151985.350 Mean Square F Value 2424.475045 73.227268 779.112735 675.624497 70436.081 667981.845 256089.634 1092.272 15813.089 17011.754 12665.446 0.32 0.01 0.10 0.09 Type III SS Mean Square F Value 0.45 4.28 1.64 0.14 2.07 2.23 1.66 Pr > F 0.5755 0.9224 0.7507 0.7673 Pr > F 6438 ,0302 ,2075 .9337 ,0720 ,0542 .1031 NUM MEAN STDERR CV BLOCK 1 36 291.23a 61.94 127.62 2 36 203.13a 14.62 43.18 3 36 240.26a 25.67 64.11 PREP B MIN 36 144. M SCRAP 36 190. WINDROW 36 399. 03b 15.70 65.41 76ab 17.47 54.96 83a 56.65 85.01 NTREAT 1 27 249.20a 47.23 98.49 2 27 251.42a 56.62 117.02 3 27 239.97a 39.27 85.04 4 27 238.89a 40.86 88.87 139 Dependent Va r i ab le : Magnesium (kg/ha) d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 1 686.031958 1 6448.521490 1 2071.861936 1 3013.851402 686.031958 0.08 0.7850 6448.521490 0.71 0.4044 2071.861936 0.23 0.6357 3013.851402 0.33 0.5680 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 2 4 184661.3613 112447.6894 348291.6440 92330.6807 56223.8447 87072.9110 1.49 0.91 1.40 0.2523 0.4217 0.2728 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 9206.415 61626.773 122883.855 137750.379 3068.805 10271.129 20480.642 11479.198 0.34 1.13 2.24 1.26 0.7993 0.3603 0.0527 0.2706 NUM MEAN STDERR CV BLOCK 1 36 73.76a 37.46 304.70 2 36 -1.57a 12.07 -4624.91 3 36 -22.55a 18.91 -503.23 PREP B MIN 36 -14.68a 12.92 -528.13 M SCRAP 36 3.34a 10.19 1829.73 WINDROW 36 60.98a 41.14 404.85 NTREAT 1 27 25.70a 34.73 702.27 2 27 15.82a 40.73 1337.65 3 27 1.82a 17.92 5127.17 4 27 22.85a 22.03 500.92 140 Dependent Variable: Cation Exchange Capacity (meq/lOOg) 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.01557407 0.01557407 0.00 0.9598 NITROGEN-QUADRATIC 1 7.41564815 7.41564815 1.22 0.2737 NITROGEN-CUBIC 1 0.12757407 0.12757407 0.02 0.8852 CONTROL VS OTHERS 1 1.91361111 1.91361111 0.32 0.5766 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 87.446667 43.723333 0.42 0.6648 PREP 2 169.085000 84.542500 0.81 0.4614 PREP*BLOCK 4 1135.483333 283.870833 2.71 0.0628 NTREAT 3 7.558796 2.519599 0.42 0.7426 NTREAT*BLOCK 6 19.310370 3.218395 0.53 0.7825 PREP*NTREAT 6 37.036481 6.172747 1.02 0.4236 PREP*NTREAT*BLOCK 12 57.701852 4.808488 0.79 0.6555 NUM MEAN STDERR CV BLOCK 1 36 14.93a 1.02 40. 98 2 36 12.99a 0.54 24. 76 3 36 14.88a 1.25 50. 39 PREP B MIN 36 13.66a 1.26 55. 51 M SCRAP 36 13.12a 0.60 27. 42 WINDROW 36 16.01a 0.93 34. 85 NTREAT 1 27 14.03a 1.22 45. 10 2 27 14.49a 1.28 45. 75 3 27 14.57a 0.94 33. 62 4 27 13.97a 1.14 42. 37 141 Dependent Va r i ab le : Cat ion Exchange Capacity (meq/lOOg) 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 1.62251852 1.62251852 0.18 0.6737 NITROGEN-QUADRATIC 1 0.65333333 0.65333333 0.07 0.7892 NITROGEN-CUBIC 1 0.37340741 0.37340741 0.04 0.8398 CONTROL VS OTHERS 1 0.45938272 0.45938272 0.05 0.8226 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 332.172407 166.086204 0.80 0.4664 PREP 2 699.650185 349.825093 1.68 0.2150 PREP*BLOCK 4 1617.412037 404.353009 1.94 0.1478 NTREAT 3 2.649259 0.883086 0.10 0.9610 NTREAT*BLOCK 6 107.035741 17.839290 1.97 0.0859 PREP*NTREAT 6 78.160185 13.026698 1.44 0.2168 PREP*NTREAT*BLOCK 12 155.613148 12.967762 1.43 0.1800 NUM MEAN STDERR CV BLOCK 1 36 16.53a 2.13 77.50 2 36 12.44a 0.40 19.19 3 36 13.33a 0:88 39.41 PREP B MIN 36 12. M SCRAP 36 12. WINDROW 36 17. 25a 0.79 38.57 35a 0.49 23.74 70a 2.08 70.55 NTREAT 1 27 14.21a 1.45 52.97 2 27 14.15a 2.06 75.54 3 27 14.20a 1.32 48.23 4 27 13.83a 1.50 56.20 142 Dependent Va r i ab le : Cat ion Exchange Capacity (meq/lOOg) d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 1.32016667 1.32016667 0.08 0.7778 NITROGEN-QUADRATIC 1 3.66675926 3.66675926 0.22 0.6384 NITROGEN-CUBIC 1 0.06446296 0.06446296 0.00 0.9503 CONTROL VS OTHERS 1 4.24817901 4.24817901 0.26 0.6130 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 186.2635185 93.1317593 0.71 0.5068 PREP 2 193.6451852 96.8225926 0.73 0.4938 PREP*BLOCK 4 236.5437037 59.1359259 0.45 0.7723 NTREAT 3 5.051389 1.683796 0.10 0.9582 NTREAT*BLOCK 6 84.790556 14.131759 0.86 0.5295 PREP*NTREAT 6 90.186667 15.031111 0.92 0.4909 PREP*NTREAT*BLOCK 12 163.948889 13.662407 0.83 0.6173 NUM MEAN STDERR CV BLOCK 1 36 1.60a 1.47 550.20 2 36 -0.55a 0.61 -667.19 3 36 -1.55a 0.82 -318.77 PREP B MIN 36 -1.41a 0.76 -322.88 M SCRAP 36 -0.77a 0.57 -443.53 WINDROW 36 1.69a 1.51 536.67 NTREAT 1 27 0.18a 1.35 3959.33 2 27 -0.33a 1.59 -2483.64 3 27 -0.37a 0.84 -1189.58 4 27 -0.14a 0.96 -3556.63 143 Dependent Va r i ab le : pH 1987 Contrast DF NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 Contrast SS Mean Square F Value 22326000 00231481 17280667 25111235 0.31511852 10.70054074 0.46497593 0.39838148 0.41345185 0.44989630 1.34452037 0.22326000 0.00231481 0.17280667 0.25111235 0.15755926 5.35027037 0.11624398 0.13279383 0.06890864 0.07498272 0.11204336 2. 0. 2. 3. 81 03 18 16 Type III SS Mean Square F Value 0.65 22.09 0.48 1.67 0.87 0.94 1.41 Pr > F 0.0993 0.8650 0.1459 0.0809 Pr > F 0.5336 0.0001 0.7500 0.1836 0.5243 0.4712 0.1894 NUM MEAN STDERR CV BLOCK 1 36 4.50a 0.06 8.12 2 36 4.58a 0.09 11.47 3 36 4.63a 0.08 10.36 PREP B MIN 36 4.44b 0.03 3.57 M SCRAPE 36 4.27b 0.04 5.02 WINDROW 36 5.01a 0.09 10.43 NTREAT 1 27 4.66a 0.09 9.76 2 27 4.53a 0.10 10.89 3 27 4.60a 0.10 11.26 4 27 4.50a 0.07 8.33 144 Dependent Variable: pH 1988 Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS DF Contrast SS 1 0.03112963 1 0.15262593 1 0.01779630 1 0.00296420 Mean Square F Value 0.03112963 0.15262593 0.01779630 0.00296420 0.55 2.69 0.31 0.05 Pr > F 0.4624 0.1071 0.5781 0.8202 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 0.66851296 2 12.34264630 4 1.73904815 0.33425648 6.17132315 0.43476204 0.86 15.94 1.12 0.4386 0.0001 0.3768 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 0.20155185 0.31935370 0.68282037 0.78350741 0.06718395 0.05322562 0.11380340 0.06529228 1.18 0.94 2.00 1.15 0.3252 0.4767 0.0813 0.3426 NUM MEAN STDERR CV BLOCK 1 36 4.87a 0.11 13.27 2 36 4.68a 0.06 7.87 3 36 4.79a 0.07 9.14 PREP B MIN 36 4.51b 0.02 3.10 M SCRAPE 36 4.58b 0.02 3.22 WINDROW 36 5.26a 0.10 11.59 NTREAT 1 27 4.79a 0.10 10.72 2 27 4.75a 0.09 9.61 3 27 4.73a 0.09 9.39 4 27 4.85a 0.11 12.24 145 Dependent Variable: pH differ e n c e 1987-88 Contrast DF Contrast SS NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 0.42112296 0.11734815 0.30151407 0.19951111 1.51333889 1.02867222 3.65465556 0.83998519 0.95785370 2.03000926 2.11241852 Mean Square F Value 42112296 11734815 30151407 19951111 0.75666944 0.51433611 0.91366389 0.27999506 0.15964228 0.33833488 0.17603488 3.03 0.84 2.17 1.44 Type III SS Mean Square F Value 2.45 1.67 2.96 2.01 1.15 2.43 1.27 Pr > F 0.0875 0.3623 0.1466 0.2362 Pr > F 0.1144 0.2168 0.0484 1228 3475 0373 2653 NUM MEAN STDERR CV BLOCK 1 36 0.37a 0.07 119.34 2 36 0.10a 0.09 533.09 3 36 0.16a 0.08 293.97 PREP B MIN 36 0.07a 0.03 252.62 M SCRAPE 36 0.30a 0.03 67.54 WINDROW 36 0.25a 0.13 313.04 NTREAT 1 27 0.13a 0.11 427.21 2 27 0.22a 0.09 225.03 3 27 0.13a 0.08 329.19 4 27 0.35a 0.08 119.03 APPENDIX 4; Summary o f s t a t i s t i c a l a n a l y s i s f o r n a t i v e v e g e t a t i o n d a t a 147 Dependent Va r i ab le : Percent cover Arnica cordifolia 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.28474074 0.28474074 0.85 0.3600 NITROGEN-QUADRATIC 1 0.21333333 0.21333333 0.64 0.4277 NITROGEN-CUBIC 1 0.17785185 0.17785185 0.53 0.4688 CONTROL VS OTHERS 1 0.06530864 0.06530864 0.20 0.6602 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 2.48666667 1.24333333 1.58 0.2334 PREP 2 1.31166667 0.65583333 0.83 0.4509 PREP*BLOCK 4 6.22333333 1.55583333 1.98 0.1415 NTREAT 3 0.67592593 0.22530864 0.67 0.5715 NTREAT * BLOCK 6 0.87185185 0.14530864 0.43 0.8523 PREP*NTREAT 6 1.45574074 0.24262346 0.73 0.6304 PREP *NTREAT * BLOCK 12 3.53148148 0.29429012 0.88 0.5707 NUM MEAN STDERR CV BLOCK 1 36 0.08a 0.03 213.18 2 36 0.14a 0.06 252.86 3 36 0.43a 0.18 251.94 PREP1 B BURN 36 0.37a 0.18 291.31 M SCRA 36 0.15a 0.06 242.19 W BURN 36 0.13a 0.04 176.40 NTREAT2 1 27 0.17a 0.06 179.97 2 27 0.20a 0.08 215.91 3 27 0.14a 0.07 266.04 4 27 0.35a 0.23 342.03 1 S i te preparat ion treatments: B BURN = broadcast burn mineral s o i l , M SCRA = between windrow and, W BURN = windrow burn 2 Nitrogen treatments: 1, 2, 3,.and 4 = 0 , 10, 20, and 30 kg/ha inocu la ted a l s i k e c lover seed re spec t i ve l y 148 Dependent Va r i ab le : Percent cover Arnica cordifolia 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS 0.00185185 0.00592593 0.06666667 0.02086420 0.00185185 0.00592593 0.06666667 0.02086420 0.00 0.01 0.15 0.05 0.9492 0.9093 0.7026 0.8308 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 2 4 14.20796296 0.33462963 14.45648148 7.10398148 0.16731481 3.61412037 6.62 0.16 3.37 0.0070 0.8567 0.0317 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 0.07444444 0.89055556 3.23277778 6.82055556 0.02481481 0.14842593 0.53879630 0.56837963 0.05 0.33 1.19 1.26 0.9829 0.9194 0.3253 0.2715 NUM MEAN STDERR CV BLOCK 1 36 0.57a 0.09 93.74 2 36 0.63a 0.11 107.27 3 36 1.37b 0.19 81.42 PREP B BURN 36 0.93a 0.21 133.95 M SCRA 36 0.85a 0.10 73.87 W BURN 36 0.79a 0.11 84.03 NTREAT 1 27 0.83a 0.15 92.05 2 27 0.90a 0.15 88.42 3 27 0.83a 0.14 85.83 4 27 0.87a 0.23 140.88 149 Dependent Va r i ab le : Percent cover Arnica cordifolia d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.24066667 0.24066667 0.63 0.4301 NITROGEN-QUADRATIC 1 0.29037037 0.29037037 0.76 0.3864 NITROGEN-CUBIC 1 0.02674074 0.02674074 0.07 0.7920 CONTROL VS OTHERS 1 0.01234568 0.01234568 0.03 0.8578 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 4.89351852 2.44675926 5.80 0.0114 PREP 2 0.42740741 0.21370370 0.51 0.6109 PREP*BLOCK 4 1.92148148 0.48037037 1.14 0.3700 NTREAT 3 0.55777778 0.18592593 0.49 0.6919 NTREAT*BLOCK 6 0.34277778 0.05712963 0.15 0.9883 PREP*NTREAT 6 2.85555556 0.47592593 1.25 0.2962 PREP*NTREAT*BLOCK 12 2.88222222 0.24018519 0.63 0.8068 NUM MEAN STDERR CV BLOCK 1 36 0.50b 0.08 99.40 2 36 0.48b 0.09 114.13 3 36 0.94a 0.12 76.05 PREP B BURN 36 0.56a 0.12 130.76 M SCRA 36 0.71a 0.10 82.92 W BURN 36 0.66a 0.09 85.43 NTREAT 1 27 0.66a 0.12 92.58 2 27 0.69a 0.13 96.52 3 27 0.69a 0.13 95.10 4 27 0.52a 0.11 112.84 150 Dependent Va r i ab le : Percent cover Calamagrostis canadensis 1987 Contrast DF Contrast SS Mean Square F Value NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT P REP * NTREAT * BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 0.01557407 0.00231481 0.00535185 0.01114198 Type III SS 0.57388889 0.09388889 0.20222222 0.02324074 0.40537037 0.40537037 0.41851852 0.01557407 0.00231481 0.00535185 0.01114198 0.28694444 0.04694444 0.05055556 0.00774691 0.06756173 0.06756173 0.03487654 0.24 0.04 0.08 0.17 Mean Square F Value 1.96 0.32 0.35 0.12 1.05 1.05 0.54 Pr > F 0.6255 0.8506 0.7746 0.6796 Pr > F 0.1699 0.7299 0.8439 0.9480 0.4067 0.4067 0.8789 NUM MEAN STDERR CV BLOCK 1 36 0.10a 0.04 220.02 2 36 0.09a 0.03 183.93 3 36 0.25a 0.06 150.43 PREP B BURN 36 0.15a 0.04 179.70 M SCRA 36 0.11a 0.04 201.57 W BURN 36 0.18a 0.06 183.24 NTREAT 1 27 0.13a 0.05 197.04 2 27 0.14a 0.05 184.55 3 27 0.17a 0.06 182.29 4 27 0.16a 0.06 198.01 151 Dependent Va r i ab le : Percent cover Calamagrostis canadensis 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 3.86757407 3.86757407 1.91 0.1725 NITROGEN-QUADRATIC 1 0.17120370 0.17120370 0.08 0.7723 NITROGEN-CUBIC 1 12.42150000 12.42150000 6.14 0.0164 CONTROL VS OTHERS 1 7.14077160 7.14077160 3.53 0.0657 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 31.13574074 15.56787037 3.02 0.0741 PREP 2 3.60240741 1.80120370 0.35 0.7100 PREP*BLOCK 4 42.13203704 10.53300926 2.04 0.1315 NTREAT 3 16.46027778 5.48675926 2.71 0.0539 NTREAT*BLOCK 6 19.63611111 3.27268519 1.62 0.1602 PREP*NTREAT 6 15.90277778 2.65046296 1.31 0.2687 PREP*NTREAT*BLOCK 12 23.05833333 1.92152778 0.95 0.5066 NUM MEAN STDERR CV BLOCK 1 36 1.20ab 0.28 140.28 2 36 1.09b 0.18 99.09 3 36 2.28a 0.38 100.45 PREP B BURN 36 1.78a 0.32 108.50 M SCRA 36 1.42a 0.32 136.02 W BURN 36 1.37a 0.27 117.19 NTREAT 1 27 1.97a 0.41 107.67 2 27 1.11b 0.20 94.50 3 27 1.85ab 0.43 121.40 4 27 1.16ab 0.29 132.52 152 Dependent Variable: Percent cover Calamagrostis canadensis d i f f e r e n c e 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 4.37400000 4.37400000 2.41 0.1268 NITROGEN-QUADRATIC 1 0.21333333 0.21333333 0.12 0.7333 NITROGEN-CUBIC 1 11.91118519 11.91118519 6.55 0.0133 CONTROL VS OTHERS 1 7.71604938 7.71604938 4.24 0.0442 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 23.25574074 11.62787037 2.85 0.0841 PREP 2 3.83351852 1.91675926 0.47 0.6326 PREP*BLOCK 4 40.78148148 10.19537037 2.50 0.0791 NTREAT 3 16.49851852 5.49950617 3.02 0.0374 NTREAT*BLOCK 6 14.98648148 2.49774691 1.37 0.2420 PREP*NTREAT 6 12.08648148 2.01441358 1.11 0.3701 PREP*NTREAT*BLOCK 12 21.02518519 1.75209877 0.96 0.4940 NUM MEAN STDERR CV BLOCK 1 36 1.10a 0.26 143.57 2 36 1.00a 0.17 100.61 3 36 2.03a 0.35 105.03 PREP B BURN 36 1.63a 0.30 109.57 M SCRA 36 1.31a 0.30 136.60 W BURN 36 1.18a 0.25 125.16 NTREAT 1 27 1.84a 0.38 108.58 2 27 0.97a 0.18 93.90 3 27 1.69a 0.40 123.97 4 27 1.00a 0.26 137.20 153 Dependent Va r i ab le : Percent cover Cornus Canadensis 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.30578241 0.30578241 0.68 0.4141 NITROGEN-QUADRATIC 1 0.88020833 0.88020833 1.95 0.1683 NITROGEN-CUBIC 1 0.33500463 0.33500463 0.74 0.3928 CONTROL VS OTHERS 1 1.25315586 1.25315586 2.78 0.1015 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 5.38393519 2.69196759 6.20 0.0090 PREP 2 5.67143519 2.83571759 6.53 0.0074 PREP*BLOCK 4 12.34981481 3.08745370 7.11 0.0013 NTREAT 3 1.52099537 0.50699846 1.12 0.3479 NTREAT*BLOCK 6 2.11587963 0.35264660 0.78 0.5884 PREP*NTREAT 6 2.04171296 0.34028549 0.75 0.6092 PREP*NTREAT*BLOCK 12 4.56870370 0.38072531 0.84 0.6067 NUM MEAN STDERR CV BLOCK 1 36 0.57a 0.22 225.71 2 36 0.10b 0.03 165.83 3 36 0.10b 0.03 155.13 PREP B BURN 36 0.58a 0.22 221.87 M SCRA 36 0.10b 0.03 165.83 W BURN 36 0.09b 0.03 171.56 NTREAT 1 27 0.44a 0.26 308.46 2 27 0.12a 0.04 175.90 3 27 0.22a 0.08 198.68 4 27 0.25a 0.12 239.60 154 Dependent Va r i ab le : Percent cover Cornus canadensis 1988 Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS DF 1 1 1 1 Contrast SS Mean Square F Value 0.32266667 0.85333333 0.86400000 0.11111111 0.32266667 0.85333333 0.86400000 0.11111111 0.72 1.91 1.93 0.25 Pr > F 0.3996 0.1730 0.1704 0.6203 Source BLOCK PREP PREP*BLOCK DF Type III SS 2 11.49018519 2 22.08796296 4 21.57148148 Mean Square F Value Pr > F 5.74509259 3.87 0.0400 11.04398148 7.44 0.0044 5.39287037 3.63 0.0244 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 2.04000000 3.90611111 2.28388889 7.13666667 0.68000000 0.65101852 0.38064815 0.59472222 1.52 1.45 0.85 1.33 0.2199 0.2114 0.5370 0.2301 NUM MEAN STDERR CV BLOCK 1 36 1.53a 0.25 96.34 2 36 0.75b 0.08 63.94 3 36 1.26ab 0.14 67.59 PREP B BURN 36 1.81a 0.23 77.49 M SCRA 36 0.96b 0.12 75.68 W BURN 36 0.77b 0.10 75.32 NTREAT 1 27 1.24a 0.16 66.74 2 27 0.95a 0.15 79.54 3 27 1.24a 0.22 92.06 4 27 1.30a 0.28 109.65 155 Dependent Va r i ab le : Percent cover Cornus Canadensis d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 1.25667130 1.25667130 2.08 0.1550 NITROGEN-QUADRATIC 1 0.00020833 0.00020833 0.00 0.9853 NITROGEN-CUBIC 1 0.12300463 0.12300463 0.20 0.6536 CONTROL VS OTHERS 1 0.61797068 0.61797068 1.02 0.3164 Source DF Type III SS Mean Square F Value Pr > F BLOCK PREP PREP*BLOCK 2 2 4 4.77263889 5.70291667 2.48111111 2.38631944 2.85145833 0.62027778 1.60 1.92 0.42 2286 1760 7943 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 1.37988426 5.56643519 3.35726852 13.57037037 0.45996142 0.92773920 0.55954475 1.13086420 0.76 1.54 0.93 1.87 0.5207 0.1843 0.4837 0.0593 NUM MEAN STDERR CV BLOCK 1 36 0.96a 0.22 137.75 2 36 0.65a 0.08 69.34 3 36 1.16a 0.14 70.10 PREP B BURN 36 1.23a 0.23 109.99 M SCRA 36 0.87a 0.11 77.72 W BURN 36 0.67a 0.09 81.87 NTREAT 1 27 0.79a 0.22 142.74 2 27 0.83a 0.13 79.83 3 27 1.02a 0.19 95.54 4 27 1.05a 0.19 94.17 156 Dependent Var i ab le : Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source Percent cover Epilobium angustifolium 1987 DF 1 1 1 1 DF Contrast SS 1.11157407 1.22453704 2.60416667 1.07929012 Mean Square F Value 1.11157407 1.22453704 2.60416667 1.07929012 0.52 0.57 1.21 0.50 Type III SS Mean Square F Value Pr > F 0.4756 0.4541 0.2763 0.4820 Pr > F BLOCK PREP PREP*BLOCK 2 44.71166667 2 17.88666667 4 36.66833333 22.35583333 3.63 0.0474 8.94333333 1.45 0.2604 9.16708333 1.49 0.2475 NTREAT 3 4.9402778 1.6467593 0.76 0.5188 NTREAT * BLOCK 6 12.9305556 2.1550926 1.00 0.4345 PREP*NTREAT 6 14.2822222 2.3803704 1.11 0.3714 PREP*NTREAT*BLOCK 12 25.5561111 2.1296759 0.99 0.4713 NUM MEAN STDERR CV BLOCK 1 36 1.51ab 0.34 136.36 2 36 1.15b 0.19 98.56 3 36 2.66a 0.34 77.22 PREP B BURN 36 1.23a 0.23 112.11 M SCRA 36 2.21a 0.30 82.45 W BURN 36 1.86a 0.38 122.69 NTREAT 1 27 1.60a 0.36 117.40 2 27 1.62a 0.43 137.85 3 27 2.13a 0.36 88.61 4 27 1.73a 0.30 91.62 157 Dependent Va r i ab le : Percent cover Epilojbiu/n angustifolium 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 0.05601852 17.20009259 20.22268519 1.10250000 0.05601852 0.01 0.9329 17.20009259 2.20 0.1439 20.22268519 2.59 0.1137 1.10250000 0.14 0.7088 Source BLOCK PREP PREP*BLOCK DF 2 2 4 Type III SS Mean Square F Value 92.9590741 37.9924074 106.1292593 46.4795370 18.9962037 26.5323148 2.20 0.90 1.26 Pr > F 0.1394 0.4240 0.3228 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 37.4787963 78.0964815 32.7742593 127.1596296 12.4929321 13.0160802 5.4623765 10.5966358 1.60 1.66 0.70 1.35 0.2007 0.1477 0.6519 0.2165 NUM MEAN STDERR CV BLOCK 1 36 4.56a 0.50 65.23 2 36 4.94a 0.56 68.02 3 36 6.69a 0.64 57.34 NUM MEAN STDERR CV PREP B BURN 36 4.62a 0.57 73.71 M SCRA 36 5.53a 0.48 52.16 W BURN 36 6.06a 0.68 67.22 NUM MEAN STDERR CV NTREAT 1 27 5.23a 0.69 68.13 2 27 5.23a 0.71 70.42 3 27 6.37a 0.71 57.95 4 27 4.78a 0.59 63.68 158 Dependent Va r i ab le : Percent cover Epilobium angustifolium d i f f e r n c e 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 1.66666667 1.66666667 0.29 0.5943 NITROGEN-QUADRATIC 1 9.24592593 9.24592593 1.59 0.2123 NITROGEN-CUBIC 1 8.31296296 8.31296296 1.43 0.2366 CONTROL VS OTHERS 1 0.00012346 0.00012346 0.00 0.9963 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 18.79018519 9.39509259 0.60 0.5588 PREP 2 17.04685185 8.52342593 0.55 0.5889 PREP*BLOCK 4 88.64259259 22.16064815 1.42 0.2683 NTREAT 3 19.2255556 6.4085185 1.10 0.3555 NTREAT*BLOCK 6 66.8950000 11.1491667 1.92 0.0940 PREP *NTREAT 6 27.2738889 4.5456481 0.78 0.5869 PREP *NTREAT * BLOCK 12 78.2255556 6.5187963 1.12 0.3616 NUM MEAN STDERR CV BLOCK 1 36 3.06a 0.45 87.74 2 36 3.80a 0.50 78.63 3 36 4.04a 0.51 75.80 PREP B BURN 36 3.39a 0.41 71.99 M SCRA 36 3.31a 0.38 68.99 W BURN 36 4.19a 0.63 90.40 NTREAT 1 27 3.63a 0.61 87.96 2 27 3.61a 0.58 84.21 3 27 4.24a 0.50 61.71 4 27 3.05a 0.55 92.93 159 Dependent Variable: Percent cover Petasites palmatus 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0 .03112963 0.03112963 0.24 0.6263 NITROGEN-QUADRATIC 1 0 .01120370 0.01120370 0.09 0.7700 NITROGEN-CUBIC 1 0 .04090741 0.04090741 0.32 0.5768 CONTROL VS OTHERS 1 0 .06250000 0.06250000 0.48 ,0.4906 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 2 .26462963 1.13231481 3.36 0.0577 PREP 2 0 .65129630 0.32564815 0.97 0.3998 PREP*BLOCK 4 1 .49925926 0.37481481 1.11 0.3819 NTREAT 3 0 .08324074 0.02774691 0.21 0.8864 NTREAT*BLOCK 6 0 .17759259 0.02959877 0.23 0.9658 PREP*NTREAT 6 0 .60870370 0.10145062 0.78 0.5878 PREP*NTREAT*BLOCK 12 1 .13629630 0.09469136 0.73 0.7163 NUM MEAN STDERR CV BLOCK 1 36 0.33a 0.12 209. 69 2 36 0.02b 0.01 206. 43 3 36 0.03b 0.01 143. 43 PREP B BURN 36 0.03a 0.01 163. 53 M SCRA 36 0.14a 0.09 365. 57 W BURN 36 0.22a 0.09 239. 06 NTREAT 1 27 0.17a 0.11 350. 60 2 27 0.10a 0.07 382. 30 3 27 0.14a 0.08 305. 70 4 27 0.11a 0.05 243. 51 160 Dependent Va r i ab le : Percent cover Petasites palmatus 1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0. 24066667 0.24066667 0.26 0.6149 NITROGEN-QUADRATIC 1 1. 00148148 1.00148148 1.07 0.3065 NITROGEN-CUBIC 1 0. 80118519 0.80118519 0.85 0.3599 CONTROL VS OTHERS 1 1. 41345679 1.41345679 1.50 0.2254 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 25. 35185185 12.67592593 2.43 0.1163 PREP 2 4. 92074074 2.46037037 0.47 0.6315 PREP*BLOCK 4 33. 49925926 8.37481481 1.61 0.2161 NTREAT 3 2. 04333333 0.68111111 0.72 0.5416 NTREAT*BLOCK 6 9. 82000000 1.63666667 1.74 0.1291 PREP *NTREAT 6 5. 14444444 0.85740741 0.91 0.4932 PREP*NTREAT*BLOCK 12 12. 48888889 1.04074074 1.11 0.3735 NUM MEAN STDERR CV BLOCK 1 36 1.42a 0.37 154. 14 2 36 0.31a 0.11 214. 15 3 36 0.51a 0.15 179. 49 PREP B BURN 36 0.48a 0.16 196. 35 M SCRA 36 0.76a 0.27 210. 90 W BURN 36 1.00a 0.30 178. 89 NTREAT 1 27 0.94a 0.39 216. 76 2 27 0.56a 0.22 201. 76 3 27 0.74a 0.27 191. 87 4 27 0.74a 0.24 169. 91 161 Dependent Va r i ab le : Percent cover Petasites palmatus d i f f e rence 1987-1988 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0.09868519 0.09868519 0.16 0.6935 NITROGEN-QUADRATIC 1 0.80083333 0.80083333 1.27 0.2641 NITROGEN-CUBIC 1 0.48001852 0.48001852 0.76 0.3861 CONTROL VS OTHERS 1 0.88151235 0.88151235 1.40 0.2416 Source DF Type III SS Mean Square F Value Pr > F BLOCK 2 12.58129630 6.29064815 2.07 0.1553 PREP 2 2.00018519 1.00009259 0.33 0.7239 PREP*BLOCK 4 21.57259259 5.39314815 1.77 0.1781 NTREAT 3 1.37953704 0.45984568 0.73 0.5378 NTREAT*BLOCK 6 7.58907407 1.26484568 2.01 0.0800 PREP*NTREAT 6 5.01462963 0.83577160 1.33 0.2603 PREP*NTREAT*BLOCK 12 13.54592593 1.12882716 1.80 0.0724 NUM MEAN STDERR CV BLOCK 1 36 1.09a 0.28 154.79 2 36 0.29a 0.10 215.50 3 36 0.47a 0.15 184.31 PREP B BURN 36 0.45a 0.15 200.28 M SCRA 36 0.62a 0.20 193.21 W BURN 36 0.78a 0.24 182.83 NTREAT 1 27 0.77a 0.30 203.84 2 27 0.46a 0.16 179.92 3 27 0.61a 0.21 179.53 4 27 0.63a 0.23 190.53 162 Dependent Va r i ab le : Percent cover Linnaea borealis 1987 Contrast DF Contrast SS Mean Square NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT* BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 0.17785185 0.31148148 0.00362963 0.00012346 12.58574074 29.19240741 36.92537037 0.49296296 1.21203704 4.09870370 5.11462963 0.17785185 0.31148148 0.00362963 0.00012346 F Value 0.26 0.45 0.01 0.00 Type III SS Mean Square F Value 6.29287037 6.45 14.59620370 14.95 9.23134259 9.46 0.16432099 0.24 0.20200617 0.29 0.68311728 0.98 0.42621914 0.61 Pr > F 0.6147 0.5057 0.9426 0.9894 Pr > F 0.0077 0.0001 0.0003 8703 9386 4448 0.8206 NUM MEAN STDERR CV BLOCK 1 36 1.16a 0.28 147.28 2 36 0.49b 0.10 115.32 3 36 0.39b 0.12 189.93 PREP B BURN 36 1.39a 0.27 116.92 M SCRA 36 0.50b 0.13 154.47 W BURN 36 0.15b 0.04 175.66 NTREAT 1 27 0.68a 0.20 155.08 2 27 0.76a 0.20 135.66 3 27 0.71a 0.31 226.24 4 27 0.57a 0.17 157.49 163 Dependent Var i ab le : Contrast NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK Percent cover Linnaea borealis 1988 DF Contrast SS 1 0.79350000 1 0.20453704 1 2.38668519 1 1.82250000 DF Type III SS 2 53.62388889 2 63.45500000 4 63.96277778 3 3.3847222 6 2.3872222 6 4.6494444 12 14.7261111 Mean Square F Value 0.79350000 0.20453704 2.38668519 1.82250000 0.37 0.10 1.12 0.85 Mean Square F Value 26.81194444 2.78 31.72750000 3.29 15.99069444 1.66 1.1282407 0.53 0.3978704 0.19 0.7749074 0.36 1.2271759 0.58 Pr > F 5444 7580 2948 3593 Pr > F 0.0888 0.0607 0.2037 6642 9794 8989 0.8521 NUM MEAN STDERR CV BLOCK 1 36 3.20a 0.47 88.94 2 36 1.65a 0.18 67.20 3 36 1.76a 0.30 103.10 PREP B BURN 36 3.28a 0.42 76.00 M SCRA 36 1.79a 0.30 102.44 W BURN 36 1.54a 0.28 107.90 NTREAT 1 27 1.98a 0.37 96.87 2 27 2.41a 0.41 88.69 3 27 2.09a 0.50 125.70 4 27 2.34a 0.38 83.40 164 Dependent Va r i ab le : Percent cover Linnaea > borealis d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 1.72268519 1.72268519 1.49 0.2275 NITROGEN-QUADRATIC 1 0.01120370 0.01120370 0.01 0.9219 NITROGEN-CUBIC 1 2.20416667 2.20416667 1.91 0.1730 CONTROL VS OTHERS 1 1.79262346 1.79262346 1.55 0.2184 Source DF Type III SS Mean Square F Value Pr > 1 BLOCK 2 15.23018519 7.61509259 1.20 0.3237 PREP 2 7.55574074 3.77787037 0.60 0.5614 PREP*BLOCK 4 7.59703704 1.89925926 0.30 0.8743 NTREAT 3 3.9380556 1.3126852 1.14 0.3430 NTREAT*BLOCK 6 1.4505556 0.2417593 0.21 0.9724 PREP *NTREAT 6 1.6361111 0.2726852 0.24 0.9629 PREP*NTREAT*BLOCK 12 8.9377778 0.7448148 0.64 0.7949 CV 90.10 66.89 101.79 68.44 103.82 117.82 91.91 105.90 107.34 75.04 NUM MEAN STDERR BLOCK 1 36 2.04a 0.31 2 36 1.16a 0.13 3 36 1.37a 0.23 PREP B BURN 36 1.89a 0.22 M SCRA 36 1.28a 0.22 W BURN 36 1.39a 0.27 NTREAT 1 27 1.30a 0.23 2 27 1.65a 0.34 3 27 1.38a 0.28 4 27 1.77a 0.26 165 Dependent Va r i ab le : Percent cover Rosa acicularis 1987 Contrast DF NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 Contrast SS Mean Square F Value 1.25185185 0.68481481 0.04629630 0.19753086 1.25185185 0.68481481 0.04629630 0.19753086 1.47 0.80 0.05 0.23 Pr > F 0.2308 0.3740 0.8166 0.6321 Source BLOCK PREP PREP*BLOCK DF 2 2 4 Type III SS Mean Square F Value 9.30018519 6.49240741 0.88259259 4.65009259 3.24620370 0.22064815 1.19 0.83 0.06 Pr > F 0.3276 0.4523 0.9935 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 1.98296296 6 10.62648148 6 4.46537037 12 6.91518519 0.66098765 1.77108025 0.74422840 0.57626543 0.78 2.08 0.87 0.68 0.5127 0.0710 0.5206 0.7663 NUM MEAN STDERR CV BLOCK 1 36 0.47a 0.14 182.77 2 36 1.17a 0.27 140.71 3 36 0.67a 0.15 131.14 PREP B BURN 36 0.43a 0.13 185.71 M SCRA 36 0.88a 0.22 150.12 W BURN 36 1.00a 0.23 138.46 NTREAT 1 27 0.84a 0.27 165.57 2 27 0.87a 0.23 136.31 3 27 0.83a 0.22 140.76 4 27 0.54a 0.21 207.43 166 Dependent Va r i ab le : Percent cover Rosa acicularis 1988 Contrast DF Contrast SS Mean Square NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP *NTREAT* BLOCK 1 1 1 1 DF 2 2 4 3 6 6 12 11.67474074 0.08333333 0.01451852 6.30567901 10.08907407 38.12796296 5.63981481 11.7725926 21.0012963 3.6068519 23.4742593 F Value 11.67474074 0.08333333 0.01451852 6.30567901 57 03 01 47 Type III SS Mean Square F Value 5.04453704 0.55 19.06398148 2.06 1.40995370 0.15 3.9241975 1.54 3.5002160 1.37 0.6011420 0.24 1.9561883 0.77 Pr > F 0.0371 0.8574 0.9402 0.1220 Pr > F 0.5891 0.1565 0.9595 0.2157 0.2434 0.9631 0.6819 NUM MEAN STDERR CV BLOCK 1 36 1.34a 0.25 110.10 2 36 1.67a 0.35 125.52 3 36 2.08a 0.38 108.48 PREP B BURN 36 0.89a 0.21 141.08 M SCRA 36 1.89a 0.33 104.19 W BURN 36 2.31a 0.39 100.72 NTREAT 1 27 2.11a 0.39 96.49 2 27 1.86a 0.45 126.41 3 27 1.59a 0.33 107.72 4 27 1.22a 0.33 142.32 167 Dependent Va r i ab le : Percent cover Rosa acicularis d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 4.21350000 4.21350000 3.27 0.0762 NITROGEN-QUADRATIC 1 0.12675926 0.12675926 0.10 0.7550 NITROGEN-CUBIC 1 0.03112963 0.03112963 0.02 0.8771 CONTROL VS OTHERS 1 3.06250000 3.06250000 2.38 0.1291 Source DF Type III SS Mean Square F Value Pr > ] BLOCK 2 13.45388889 6.72694444 2.10 0.1518 PREP 2 11.90055556 5.95027778 1.85 0.1852 PREP*BLOCK 4 3.79555556 0.94888889 0.30 0.8769 NTREAT 3 4.37138889 1.45712963 1.13 0.3450 NTREAT*BLOCK 6 9.08388889 1.51398148 1.17 0.3336 PREP*NTREAT 6 5.97500000 0.99583333 0.77 0.5948 PREP*NTREAT*BLOCK 12 19.39555556 1.61629630 1.25 0.2727 NUM MEAN STDERR CV BLOCK 1 36 0.86a 0.20 137.02 2 36 0.50a 0.14 168.42 3 36 1.36a 0.29 129.06 PREP B BURN 36 0.46a 0.14 178.10 M SCRA 36 1.01a 0.22 129.49 W BURN 36 1.26a 0.28 134.73 NTREAT 1 27 1.20a 0.25 110.36 2 27 0.99a 0.34 177.44 3 27 0.76a 0.21 141.08 4 27 0.69a 0.23 171.19 168 Dependent Va r i ab le : Percent cover Salix spp. 1987 Contrast DF Contrast SS Mean Square F Value NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 1.26150000 1 0.08898148 1 0.19646296 1 0.34027778 DF Type III SS 2 2.72074074 2 3.74018519 4 2.21148148 3 1.54694444 6 7.74222222 6 4.68277778 12 7.04555556 1.26150000 0.08898148 0.19646296 0.34027778 1.36037037 1.87009259 0.55287037 0.51564815 1.29037037 0.78046296 0.58712963 1.79 0.13 0.28 0.48 Mean Square F Value 1.57 2.16 0.64 0.73 1.83 1.11 0.83 Pr > F 0.1866 0.7238 0.5998 0.4902 Pr > F 0.2354 0.1446 0.6423 0.5378 0.1105 0.3705 0.6170 NUM MEAN STDERR CV BLOCK 1 36 0.52a 0.19 221.56 2 36 0.15a 0.07 287.03 3 36 0.24a 0.15 379.11 PREP B BURN 36 0.08a 0.04 296.57 M SCRA 36 0.53a 0.21 234.57 W BURN 36 0.29a 0.14 277.86 NTREAT 1 27 0.20a 0.18 470.96 2 27 0.17a 0.07 213.76 3 27 0.38a 0.18 244.41 4 27 0.46a 0.21 245.15 169 Dependent Va r i ab le : Percent cover Salix spp. 1988 Contrast DF Contrast SS Mean Square F Value NITROGEN-LINEAR NITROGEN-QUADRATIC NITROGEN-CUBIC CONTROL VS OTHERS Source BLOCK PREP PREP*BLOCK NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 1 2.97779630 1 0.34453704 1 0.17424074 1 1.22225309 DF Type III SS 2 22.00907407 2 5.80907407 4 9.15148148 3 3.49657407 6 21.25981481 6 12.54870370 12 27.16407407 2.97779630 0.34453704 0.17424074 1.22225309 1.32 0.15 0.08 0.54 Mean Square F Value 11.00453704 6.77 2.90453704 1.79 2.28787037 1.41 1.16552469 0.52 3.54330247 1.58 2.09145062 0.93 2.26367284 1.01 Pr > F 2549 6970 7818 4642 Pr > F 0.0064 0.1957 0.2712 0.6715 0.1722 0.4812 0.4560 NUM MEAN STDERR CV BLOCK 1 36 1.34a 0.36 162.92 2 36 0.33b 0.14 248.42 3 36 0.44b 0.18 237.17 PREP B BURN 36 0.44a 0.14 189.74 M SCRA 36 1.01a 0.36 214.30 W BURN 36 0.67a 0.21 193.10 NTREAT 1 27 0.52a 0.23 233.46 2 27 0.63a 0.21 171.44 3 27 0.67a 0.27 210.71 4 27 1.00a 0.43 220.90 170 Dependent Variable : Percent cover Salix spp . difference 1987-88 Contrast D F Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 0. 36296296 0.36296296 0.43 0.5138 NITROGEN-QUADRATIC 1 0. 08333333 0.08333333 0.10 0.7540 NITROGEN-CUBIC 1 0. 74074074 0.74074074 0.88 0.3520 CONTROL VS OTHERS 1 0. 27271605 0.27271605 0.32 0.5713 Source D F Type III SS Mean Square F ' Value Pr > E BLOCK 2 9. 38388889 4.69194444 9.83 0.0013 PREP 2 0. 26166667 0.13083333 0.27 0.7633 PREP*BLOCK 4 4. 26277778 1.06569444 2.23 0.1060 NTREAT 3 1. 18703704 0.39567901 0.47 0.7038 NTREAT*BLOCK 6 5. 24574074 0.87429012 1.04 0.4097 PREP *NTREAT 6 4. 69685185 0.78280864 0.93 0.4801 PREP*NTREAT*BLOCK 12 10. 01203704 0.83433642 0.99 0.4679 NUM MEAN STDERR CV BLOCK 1 36 0.82a 0.23 165. 95 2 36 0.19b 0.08 243. 42 3 36 0.21b 0.08 219. 03 PREP B BURN 36 0.37a 0.12 190. 30 M SCRA 36 0.48a 0.18 232. 06 W BURN 36 0.38a 0.15 243. 52 NTREAT 1 27 0.32a 0.15 250. 26 2 27 0.46a 0.15 169. 78 3 27 0.29a 0.14 244. 61 4 27 0.55a 0.24 231. 67 171 Dependent Va r i ab le : Percent cover Spiraea betulifolia 1987 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 • 0.64757407 0.64757407 0.33 0.5663 NITROGEN-QUADRATIC 1 2.28231481 2.28231481 1.17 0.2835 NITROGEN-CUBIC 1 19.68446296 19.68446296 10.12 0.0024 CONTROL VS OTHERS 1 1.94447531 1.94447531 1.00 0.3218 Source DF Type III SS Mean Square F Value Pr > 1 BLOCK 2 18.6090741 9.3045370 0.78 0.4740 PREP 2 24.5946296 12.2973148 1.03 0.3775 PREP*BLOCK 4 109.3148148 27.3287037 2.29 0.0999 NTREAT 3 22.6143519 7.5381173 3.88 0.0139 NTREAT*BLOCK 6 7.9375926 1.3229321 0.68 0.6662 PREP*NTREAT 6 33.8853704 5.6475617 2.90 0.0158 PREP*NTREAT*BLOCK 12 33.0385185 2.7532099 1.42 0.1876 NUM MEAN STDERR CV BLOCK 1 36 1.41a 0.35 150.23 2 36 2.29a 0.36 95.56 3 36 2.29a 0.42 111.37 PREP B BURN 36 1.57a 0.35 132.88 M SCRA 36 2.66a 0.46 104.40 W BURN 36 1.76a 0.31 106.93 NTREAT 1 27 1.76ab 0.35 103.85 2 27 2.75a 0.62 116.41 3 27 1.53b 0.34 114.08 4 27 1.94ab 0.40 108.44 172 Dependent Va r i ab le : Percent cover Spiraea betulifolia 1988 Contrast DF NITROGEN-LINEAR 1 NITROGEN-QUADRATIC 1 NITROGEN-CUBIC 1 CONTROL VS OTHERS 1 Contrast SS Mean Square F Value Pr > F 9.57335185 0.54898148 7.99350000 4.38669753 9.57335185 0.54898148 7.99350000 4.38669753 2.12 0.12 1.77 0.97 ,1508 ,7285 .1886 ,3283 Source BLOCK PREP PREP*BLOCK DF Type III SS 2 0.0646296 2 13.1296296 4 290.5925926 Mean Square F Value 0.0323148 6.5648148 72.6481481 0.00 0.35 3.86 Pr > F 0.9983 0.7099 0.0195 NTREAT NTREAT*BLOCK PREP*NTREAT PREP*NTREAT*BLOCK 3 6 6 12 18.1158333 14.9094444 86.6422222 68.9066667 6.0386111 2.4849074 14.4403704 5.7422222 1.34 0.55 3.20 1.27 0.2712 0.7669 0.0092 0.2609 NUM MEAN STDERR CV BLOCK 1 36 3.06a 0.64 124.91 2 36 3.00a 0.47 93.61 3 36 3.01a 0.48 95.39 PREP B BURN 36 3.31a 0.62 113.21 M SCRA 36 3.22a 0.57 106.33 W BURN 36 2.53a 0.35 84.09 NTREAT 1 27 3.37a 0.61 94.60 2 27 3.45a 0.74 110.87 3 27 2.45a 0.46 97.21 4 27 2.81a 0.61 113.22 173 Dependent Va r i ab le : Percent cover Spiraea b e t u l i f o l i a d i f f e rence 1987-88 Contrast DF Contrast SS Mean Square F Value Pr > F NITROGEN-LINEAR 1 5.24118519 5.24118519 1.93 0.1702 NITROGEN-QUADRATIC 1 5.07000000 5.07000000 1.87 0.1772 NITROGEN-CUBIC 1 2.59029630 2.59029630 0.96 0.3328 CONTROL VS OTHERS 1 12.17234568 12.17234568 4.49 0.0387 Source DF Type III SS Mean Square F Value Pr > ] BLOCK 2 20.84574074 10.42287037 1.84 0.1877 PREP 2 28.44462963 14.22231481 2.51 0.1094 PREP*BLOCK 4 57.03703704 14.25925926 2.51 0.0778 NTREAT 3 12.9014815 4.3004938 1.59 0.2035 NTREAT*BLOCK 6 12.4557407 2.0759568 0.77 0.6003 PREP*NTREAT 6 17.9168519 2.9861420 1.10 0.3739 PREP *NTREAT * BLOCK 12 47.1925926 3.9327160 1.45 0.1727 NUM MEAN STDERR CV BLOCK 1 36 1.65a 0.40 145.98 2 36 0.71a 0.33 281.49 3 36 0.72a 0.25 212.54 PREP B BURN 36 1.74a 0.40 136.65 M SCRA 36 0.56a 0.34 366.20 W BURN 36 0.77a 0.23 182.31 NTREAT 1 27 1.61a 0.41 132.81 2 27 0.70a 0.30 219.17 3 27 0.92a 0.37 210.16 4 27 0.88a 0.47 278.90 APPENDIX 4. L i s t of p lant species observed i n the experimental s i t e . S c i e n t i f i c name Achillea millefolium Agoseris aurantiaca Agrostis scrabra Alnus viridis Alopecurus aequalis Arnica cordifolia Aster conspicuus Aster foliaceus Calamagrostis canadensis Carex adusta Carex athrostachya Carex macloviana spp. pachystachya Carex praticola Chamomilla suaveolens Cinna l a t i f o l i a Cladonia spp. Cornus canadensis Corydalis sempervirens Crepis capillaris Dactylis glomerata Dracocephalum parviflorum Dicranum spp. Epilojbiu/n angrustifoJium Epilobium ciliatum Equisetum arvense Equisetum sylvaticum Festuca occidentalis Fragaria virginiana Galium boreale Galium triflorum Geranium bicknellii Gnaphalium palustre Hieracium albiflorum Juncus bufonius Linnaea borealis Lonicera involucrata Lotus corniculatus Lupinus sericeus Lycopodium complanatum Marchantia polymorpha Petasites palmatus Phleum pratense Pinus contorta Pleurozium schreberi Polygonum aviculare Polytrichum juniperinum Populus tremuloides Potentilla norvegica Ribes glandulosum Ribes lacustre Rosa acicularis Rubus idaeus Rubus pedatus Common name yarrow orange f a l s e dandelion ha i r bent grass green a lder l i t t l e meadow f o x t a i l heart - leaved arn ica showy aster l ea f y as ter b l ue jo in t s lender-beaked sedge thick-headed sedge meadow sedge pineappleweed nodding wood-reed bunchberry pink coryda l i s smooth hawksbeard orchard grass American dragonhead f ireweed purp le - leaved wil lowherb common h o r s e t a i l wood h o r s e t a i l western fescue wi ld strawberry northern bedstraw sweet-scented bedstraw B i c k n e l l ' s geranium lowland cudweed white-f lowered hawkweed toad rush twinflower black twinberry b i r d ' s - f o o t t r e f o i l s i l k y lupine ground-cedar palmate c o l t ' s f o o t timothy lodgepole pine red-stemmed feathermoss common knotweed juniper haircap moss trembling aspen rough c i n q u e f o i l skunk currant black gooseberry p r i c k l y rose raspberry f i ve - l eaved bramble 175 Rubus pubescens Salix bebbiana Salix scouleriana Shepherdia canadensis Sibbaldia procumbens Spergularia rubra Spiraea betulifolia Stellaria calycantha Taraxacum officinale Tiarella trifoliata Trifolium hydridum Trisetum spicatum Vaccinium caespitosum Vaccinium membranaceum Vicia americana Viola orbiculata t r a i l i n g raspberry Bebb's willow Scou ler ' s wil low s o o p o l a l l i e creeping s i bba ld i a red sand spurrey b i r ch - l eaved sp i rea northern starwort common dandelion three- leaved foamflower a l s i k e c lover spike t r i setum dwarf b lueberry black huckleberry American vetch round-leaved v i o l e t R. Trowbridge Education: Colorado State University (1964-66, 1968-70), BSc. Agriculture (Agronomy, Foreign Service option) Western Washington State University, undeclared graduate studies in Education (1972-73) University of British Columbia (1986-87, 1988-90), MSc. Forest Science (Ecology) Special training at Universities: University of Pennsylvania and University of California (Riverside); language, cultural, and agricultural training for Peace Corps assignment in Gujarat, India (1966). College of the Virgin Islands and Arkansas State University; language, cultural, and agricultural training for Peace Corps assignment in East Cameroun, Cameroun (1971) Publications: Journal papers: Fox, C.A., R. Trowbridge, and C. Tarnocai. 1987. Classification, macromorphology and chemical characteristics of Folisols from British Columbia. Can. J . Soil Sci. 67:765-778. Trowbridge, and M.C. Feller. 1988. Relationships between the moisture content of fine woody fuels in lodgepole pine slash and the Fine Fuel Moisture Code of the Canadian Fire Weather Index System. Can. J . For. Res. 18-1:128-131. Contributions to books: Banner, A., J . Pojar, J . W. Schwab, and R. Trowbridge. 1989. Vegetation and soils of the Queen Charlotte Islands: recent impacts of development, pp. 261-279. In: G.G.E. Scudder and N. Gessler (eds.), The Outer Shores. Univ. B.C. Banner. A., R.J. Hebda, E.T. Oswald, J . Pojar, and R. Trowbridge. 1988. Wetlands of Pacific Canada, pp. 307-346, In: National Wetlands Working Group, Canada Committee on Ecological Land Classification, Wetlands of Canada. Polyscience Publ. Inc. Agriculture Canada Expert Committee on Soil Survey. 1987. The Canadian System of Soil Classification. Revision of Folisolic soils, pp. 82-92 in Chapter 9 "Organic Order". Specialized publications: Banner, A., J . Pojar, and R. Trowbridge. 1983. Ecosystem classification of the Coastal Western Hemlock Zone, Queen Charlotte Island Subzone (CWHg). B.C. Min. For., Smithers, B.C. Banner, A., J . Pojar, and R. Trowbridge. 1986. Representative wetland types of the northern part of the Pacific Oceanic Wetland Region. RR 85008-PR. B.C. Min. For., Smithers, B.C. Banner, A., J . Pojar, R. Trowbridge, and A. Hamilton. 1985. Grizzly bear habitat in the Kimsquit River valley, coastal British Columbia: classification, description, and mapping. Grizzly bear habitat symposium, Missoula, Mt., April 30-May 2, 1985. Douglas, M.J., A. Macadam, and R. Trowbridge. 1990. The conversion of multistoried bruch fields to coniferous plantations - a benchmark evalutation of alternative silicultural treatments: Treatment effects on soils. B.C. Min. For., Smithers, B.C. Klinka, K., R.N. Green, R.L. Trowbridge, and L.E. Lowe. 1981. Taxonomic classification of humus forms in ecosystems of British Columbia. Land Manage. Rep. No. 8., B.C. Min. For., Victoria, B.C. Lindeburgh, S., and R. Trowbridge. 1984. Production of forestry interpretive maps and management summaries from ecologically based polygons. RR 85001-PR, B.C. Min. For., Smithers, B.C. Macadam, A., and R. Trowbridge. 1984. Prescribed fire research in the Prince Rupert forest Region, British Columbia. RR 84010, B.C. Min. For., Smithers, B.C. Pojar, J., R. Trowbridge, and D. Coates. 1984. Ecosystem classification and interpretation of the Sub-Boreal Spruce zone, Prince Rupert Forest Region, British Columbia. Land Manage. Rep. No. 17 & Supplement. B.C. Min. For., Victoria, B.C. Pojar, J., R. Trowbridge, D. Coates, D. Holmes, and T. Lewis. 1986. A field guide to identification and interpretation of ecosystems of the Sub-Boreal Spruce Zone n the Prince Rupert Forest Region, British Columbia. Land Manage. Hand. No. 10. B.C. Min. For., Smithers, B.C, Pojar, J., R. Trowbridge, and T. Lewis. 1982. Biogeoclimatic zones of the Cassiar Timber Supply area, northwestern British Columbia. Int. Rep., B.C. Min. For., Smithers, B.C. Trowbridge, R. (ed.). 1980. First progress report of the working group on organic horizons, Folisols, and Humus Form classification. Agriculture Canada, Research Branch, Land Resource Research Institute. Ottawa, Ont. Trowbridge, R. (ed.). 1981. Second progress report of the working group on organic horizons, Folisols, and humus form classification. Agriculture Canada, Research Branch, Land Resource Research Institute. Ottawa, Ont. Trowbridge, R. 1984. Prediction of some soil thermal regimes in northwestern British Columbia with comparisons to the soil climate map of Canada and soil temperature classes used for Canada. Proceedings of the 5th annual meetings of the Expert Committee on Soil Survey. Agriculture Canada, Land Resource Research Institute, Ottawa, Ont. Trowbridge, R. 1986. Current fire research in British Columbia, pp. 38-47 In: Proceedings of the Fire Management Symposium, April 8 and 9, Prince George, B.C. Trowbridge. R. 1990. Effects of Trifolium-Rhizobium symbiosis on Pinus contorta regereration, forest soil, and selected native plant species. M.Sc. Thesis, Univ. B.C., Vancouver, B.C. Trowbridge, R., B. Hawkes, A. Macadam, and J . Parminter. 1986. field handbook for prescribed fire assessments in British Columbia: logging slash fuels. Land Manage. Hand. No. 11. B.C. Min. For., Victoria, B.C. Trowbridge, r., H. Luttmerding, and C. Tarnocai. 1984. Report on Folisolic Soil classification in Canada. Proceedings of the 6th annual meeting of the Expert Committee on Soil Survey. Agriculture Canada, Land Resource Research Institute, Ottawa, ONt. Trowbridge, R.L. and A. Macadam (compilers and eds.). 1983. Prescribed fire - forest soils symposium proceedings. Land Manage, rep. No. 16. B.C. Min. For., Victoria, B.C. Trowbridge, R., J . Pojar, and T. Lewis. 1983. Interim classification of the Boreal White and Black Spruce Biogeoclimatic Zone in the Prince Rupert Forest Region. B.C. Min. For., Smithers, B.C. 

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